BIODIVERSITY AND
BIOGEOGRAPHY OF
CARIDEAN SHRIMPS OF
PAKISTAN
QUDDUSI B.KAZMI
AND
M. AFZAL KAZMI
2010
Marine Reference Collection and
Resource Center, University of
Karachi
Higher Education Commission
Islamabad, Pakistan
ii
Inside cover page: taken from hand painted greeting card sent to the
first author by Lilly Manning of USA and saved by the first author for
this publication.
iii
BIODIVERSITY AND
BIOGEOGRAPHY OF CARIDEAN
SHRIMPS OF PAKISTAN
QUDDUSI B.KAZMI
AND
M. AFZAL KAZMI
2010
Marine Reference Collection and
Commission
Resource Center, University of Karachi
Pakistan
Higher Education
Islamabad,
iv
Table of Contents
1.General Introduction to Caridean shrimps ........................................ 1
1.1 Diagnosis ............................................................................... 2
1.2 Classification ......................................................................... 2
1.3 History of Nomenclature of Infraorder Caridea .................... 3
1.4 Taxonomic Characters and biological success ...................... 6
1.5 Body Plan .............................................................................. 8
1.6 Biology ................................................................................ 15
2.Materials and Methods .................................................................... 22
3.Results ............................................................................................. 25
3.1Biodiversity ............................................................................... 25
3.1.1 Family Pasiphaeidae Dana, 1852 ...................................... 39
3.1.2 Family Oplophoridae Dana, 1852 ..................................... 60
3.1.3 Family Atyidae de Haan, 1849 ......................................... 66
3.1.4 Family Palaemonidae Rafinesque, 1815 ......................... 106
3.1.5 Family Processidae Ortmann, 1896 ................................. 215
3.1.6 Family Ogyrididae Holthuis, 1955 ................................. 223
3.1.7 Family Alpheidae Rafinesque, 1815 ............................... 232
3.1.8 Family Hippolytidae Bate, 1888 ..................................... 328
3.1.9 Family Pandalidae Haworth, 1825 .................................. 375
3.1.10 Family Crangonidae Haworth, 1825 ............................. 378
3.2. Biogeography ........................................................................ 389
3.2.1. Previous records ............................................................. 389
3.2.2. Discussion: ..................................................................... 392
3.2.2.1. Fresh and brackish water species: ............................... 393
3.2.2.2. Marine water forms: .................................................... 401
3.2.2.3. Discussion& Conclusion:............................................ 410
4.Species of interest to fisheries and aquaculture ............................ 414
v
4.1 Water Resources-Position, threats and Potentials .................. 414
4.1.1: Fresh water:.................................................................... 414
4.1.2. Marine and Brackish. ..................................................... 419
4.2. Shrimp Resources-position and potentials:........................... 420
4.2.1. Important Species: ......................................................... 422
4.2.1. Other uses: ..................................................................... 428
Addendum ........................................................................................ 430
Literature Cited ................................................................................ 439
Glossary ........................................................................................... 500
Taxonomic Index ............................................................................. 512
vi
This study is dedicated to the memory
of the renowned carcinologist (Late)
Professor Dr. L. B. Holthuis
vii
Acknowledgments
We bestow all due praise to Almighty God whose unlimited blessings
enabled us to pursue this study. A special note of appreciation and a sense
of reverence are extended to our long suffering mother for her patience and
constant prayers.
We are deeply indebted to Late Profs. Drs. M.N.Tirmizi ,University of
Karachi and L.B.Holthuis, National Museum
of Natural History
Naturalis,Leiden, from whom we gained much of our knowledge on the
subject. The publishers and authors whose illustrations/ views are adapted
are gratefully acknowledged, especially the M/S Science Publishers Inc.
USA & UK and E.J.Brill Leiden. Thanks are due to the scientists who have
helped with the identification (some of them are no more in this world) (Drs.
A. Banner, A.J. Bruce, F.A. Chace Jr., A. Anker,I.Marin & M. Apel) and to
Drs.P.K.Ng and Y.Cai during our visit to Singapore. We are grateful to
different organizations that have provided us number of specimens of fresh
water prawns. The first author wishes to acknowledge the directors of
Marine Reference Collection and Resource Center (MRC), University of
Karachi for providing lab facility.
Ms Razia Naushaba (MRC), our students, Mr. G. Asghar and
Mr.Imdadullah (PCSIR) and Mr. Abrar Ali (MRC) are thanked for
their help while preparing the MS.
The Higher Education Commission, Islamabad provided the funds for
publication.
The two anonymous referees are thanked for their helpful comments.
viii
List of abbreviations
ANT
ANTENNA
ANT.LE
ANTENNULE
ANT.FL
ANTENNAL FLAGELLUM
ANT.SP
ANTENNAL SPINE
APP.INT
APPENDIX INTERNA
APP.MAS
APPENDIX MASCULINA
BS.PO
BASIPODITE/CARPUS
CARP
CARAPACE
CAR.PO
CARPOPODITE/CARPUS
COX.BS
COXO-BASIS
COX.PO/COX
COXOPODITE/COXA
CF
CONFER
CL
CARAPACE LENGTH
DAC.PO/DAC
DACTYLOPODITE/DACTYLUS
DOR.SP
DORSAL SPINE
EN.PO
ENDOPODITE
EX.PO
EXOPODITE
FI.CH
FIRST CHELA
FIG
FIGURE
FL
FLAGELLUM
HOZ
HORIZONTAL
ix
IN.FL
INNER FLAGELLUM
IS.PO
ISCHIO PODITE/ISCHIUM
MM
MILLIMETER
MX.PED
MAXILLIPED
ME.PO
MEROPODITE/MERUS
MO.PRO
MOLAR PROCESS
OBL
OBLIQUE
OT.FL
OUTER FLAGELLUM
PEL.PO
PLEOPOD
PER.PO
PREOPOD
PR.COX
PRECOXA
PRO.PO
PROPODITE/ PROPODUS
RST
ROSTRUM
RST.SP
ROSTRAL SPINE
SE.CH
SECOND CHELA
ST
STATION
SP
SPECIES
SQ
SQUAMA
SU
SUTURE
TE
TELSON
TE.SP
TELSON SPINE
TER
TERGUM
TL
TOTAL LENGTH
x
URO.PD
UROPOD
List of Figures
1
A. Caridean abdomen; B. Penaeidean abdomen,
lateral view.
2-5
External anatomy of different Caridean families
6-7
Morphology and appendages
8
Caridean Life cycle
9
Bopyrid parasites of carideans
10
Map showing sampling stations
11-14
Eupasiphae gilesii
15-16
Leptochela irrobusta
17-18
Leptochela nasimae
19-20
Leptochela pugnax
21-22
Leptochela cf sydniensis
23
Pasiphaea sivado
24-26
Acanthephyra eximia
27
Caridina babaulti babaulti
28
Caridina nilotica
29
Caridina sumatrensis
30
Kalriana anissi
31-43
larval stages of Kalriana anissi
xi
44
Telsons of K. jhimphirensis, K. sunahrensis, K.
karachi
45-46
Exopalaemon styliferus
47
Leandrites celebensis
48
Leptocarpus potamiscus
49
Macrobrachium altifrons ranjhai
50
Macrobrachium dacqueti
51
Macrobrachium dayanum
52
Macrobrachium equidens
53
Macrobrachium idella idella
54
Macrobrachium lamarreii korangii
55
Macrobrachium lamarreii lamarreii
56-66
larval stages of Macrobrachium lamarreii lamarreii
67-68
Macrobrachium malcolmsoni kotreeanum
69-72
larval stage of Macrobrachium malcolmsonii
kotreeanum
73
Macrobrachium malcolmsonii malcolmsonii
74
Macrobrachium naso
75
Macrobrachium rude
76
Macrobrachium scabriculum
77
Macrobrachium shahpuri
78-79
Macrobrachium taunsii
80-81
Nematopalaemon tenuipes
82-83
Palaemon pacificus
xii
84-85
Palaemon semmelinki
86
Palaemon sewelli
87-88
Anchistus custos
89-93
Cuapetes elegans
94
Cuapetes longirostris
95-98
Cuapetes seychellensis
99-104
Processa compacta
105-106
Ogyrides orientalis
107
Ogyrides saldanhae
108-109
Alberta banneri
110
Alpheus albertai
111
Alpheus barbatus
112-113
Alpheus bisincsus
114A-B
Alpheus chiragricus
115-116
Alpheus edwardsii
117
Alpheus isodactylus
118-121
Alpheus lobidens
122
Larval stage of Alpheus lobidens
123
Alpheus manorensis
124
Alpheus pacificus
125
Alpheus pseudoedwardsii
126-129
Alpheus splendidus
130-131
Alpheus strenuus strenuus
xiii
132-133
Alpheus zulfaquiri
134A
Athanas arabicus
134B-137
Athanas dimorphus
138
Automate dolichognatha
139
Salmoneus brevirostris
140
Salmoneus cristatus
141
Synalpheus thai
142-146
Synalpheus tumidomanus
147-153
Post larvae of Synalpheus tumidomanus
154-157
Exhippolysmata ensirostris ensirostris
158
Heptacarpus pandaloides
159-160
161-163
164
Hippolyte ventricosa
Latreutes anoplonyx
Latreutes mucronatus
165-169
Lysmata vittata
170-174
Saron marmoratus
175
Philocheras parvirostris
176-180
Pontocaris pennata
181
Map showing Pakistan‟s water bodies
182
Macrobrachium sp
183
Alpheus sp
184
Undetermined alpheid
xiv
List of Plates (Colour Photographs):
1
A-Alpheus,B- Saron, C-Jelly fish showing Hippolytid
2
A- Acanthephyra eximia ,B-Caridina sumatrensis,CMacrobrachium macolmsoni kotreeanum,D- M. taunsai
3
A-M. lamarrei,B- M. lehiai,C- M. semmelinki,D- M.
equidens
4
A- A. splendidus,B- Alpheus lobidens,C- Synalpheus
tumidomanus,
5
A- Latreutes anoplonyx,B- Alpheus.rapax,C- Synalpheus
tumidomannus with bopyrus,D-S. neptunus,E- Alpheus.
barbatus
6
A-Alpheus sp.,B- A. zulfaquiri,C- Saron marmoratus, DLatreutes
anoplonyx (lot)
7
A- Lysmata vittata,B- Hippolyte ventricosa,in weed,CPontocaris pennata, D-Heterocarpus laevigatus
List of Tables:
1
Details of appendages of a shrimp
2
Composition of Pakistan known caridean fauna
3
Zoogeographical component of Pakistan marine
caridean shrimps
4
Landing analysis
5
Categories of size
xv
Preface
The caridean shrimps include a vast assemblage of species inhabiting
all types of aquatic habitats from high altitude streams to coastal marine
waters. They are wide distributed around the world and have considerable
aesthetic and food value to mankind. They are very good source of high
proteins and other by-products, also have shown good qualities for bioassay
studies.
As such they may occupy a significant position in the socio-economic
fabric of our country by providing nutritious food, income, leisure and
employment opportunities, if given their due status.
Although Pakistani research on noncaridean shrimps community has
been from the beginning organized to take in parallel environmental and
biological observations there is no comprehensive account for the much
neglected caridean shrimp resources of Pakistan. There are few scattered
reports on the group published by the Pakistani scientists focusing on
specific problems. This document seeks to provide a sketch of the existing
knowledge in Pakistan. An urgent need was felt in the interest of our
scientists to consolidate the scattered informations in the form of a
monograph. One of the major aims of the monograph is to communicate
among scientists and students the important up to date advancement on the
taxonomy of the group in world in a single volume for easy accessibility and
to solve some of the unsettled taxonomical problems.
The monograph contains 4 chapters. The first is the introduction to the
group. The second will be dealing with methodology and the third is the
result of with the taxonomy of these prawns, recent classification, keys to
identification of families, subfamilies, genera and species under Caridea,
detail descriptions find place in this chapter. The same chapter includes
analysis of the biogeography of the species under the Caridea where
distribution pattern of the Pakistani Caridea is given and zoogeographic
affinities are discussed for the Pakistani species based on published
xvi
distribution. The last chapter describes our water resources and the
caridean resources they embody.
This monograph is unique in providing information on the species
spread on a vast array of region spanning from Himalayas to Exclusive
Economic Zone of Pakistan in the Arabian Sea in a single volume. There
was room for something like a comprehensive treatment of the Pakistani
carideans. After 3-4 years of intermittent work the study could be finished. In
such a big endeavor in a fixed time frame work (there is never enough time)
shortcomings are unavoidable, and critical opinions not are surprising. Still
we hope that this monograph will be quite a value to all those interested in
carcinology throughout the world particularly for those in search of
information on biodiversity of our unique zoogeographical zones and will
stimulate interest in further research on these remarkable animals. This
information needs to be passed along to prevent its loss and before some of
the species at the verge of extinction disappear they need to be
documented, particularly in the present scenario when the Indus River could
see dramatically reduced flow, and we are among the countries hardest hit
by the climatic change, along with this the world oceans are becoming
acidic at a faster rate threatening disaster for marine life and food supplies
across the globe.
Actually, the present document was designed as a review of the state of
the knowledge, problem and challenges posed in quantifying the caridean
resources in Pakistan and we consider it as work in progress, it will be a key
reference on documentation of Pakistani caridean shrimps, particularly the
fresh water shrimps. When we are facing challenges to conserve our fresh
water resources with a large number of dams, and riverine development
planned in our species diverse country. It is also expected that a study of
this kind be bound to reveal provocative gaps in the knowledge of the
animal. Some of those appeared as this monograph project unfolded, can
be solved only by additional collecting; others by comparative population
analysis and still others by laboratory observations.
It is inevitable that there are errors and omissions in this monograph. Apologies are herewith
rendered for the editing mistakes. Blames therefore be charged to both of us having weak eye
sight that caused correcting more time consuming and to our selfishly strong desire for timely
rather than our post humous publication.
xvii
Quddusi Bashir Kazmi
Ex -Director
Marine Reference Collection
&Resource Center, University of
Karachi (qbkazmi@yahoo.com)
M. Afzal Kazmi
Ex- Chairman
Department of Zoology, University
of Karachi
(makazmi@yahoo.com)
CHAPTER-1
General Introduction to Caridean shrimps
The term shrimp is applied to a number of long tailed crustaceans not
very closely related to caridean shrimps, for example, fairy shrimps, tadpole
shrimps, opossum shrimps, mantis shrimps. However, the word shrimp is
characteristically used to refer to those crustaceans having complete set of
morphological features known as cardioid facies like long antennae, slender
legs, and a laterally compressed muscular abdomen that is highly adapted for
both forward swimming and backward escape movement. Prawn is often used
as synonym of shrimp for penaeoidean and caridean shrimps, especially those
of large size. Usage of shrimp and prawn varies regionally throughout the
English-speaking world. Shrimps groups differ in morphology and biology; the
following key may distinguish the Caridean from the Penaeidea as illustrated
(Fig.1)
Figure1: A- Caridean abdomen, lateral view; BPenaeidean abdomen, lateral view(Adapted from New
&Singholka,1982)
2
KEY
1.
Pleura of second abdominal somite overlapping those of first and third
somites. No chelae on third pereopods, Gills phyllobranchiate------------------------------------Caridea.
-
Pleura of second abdominal somite not overlapping that of first somite.
Third legs chelate never stronger than preceding legs, generally all
chelipeds of equal strength. Males with petasma. Gills
dendrobranchiate---------------------------------------Penaeidea.
1.1
Diagnosis
Decapod Crustaceans with 5 to 8 inches body plan; head consisting of
a preantennal region bearing the eyes, followed by five somites with two pairs of
antennae, three pairs of mouth parts, a thorax of eight somites and an abdomen
with six somites, most of which bear appendages. Female genital apertures or
gonopores positioned on the sixth thoracic somite and male apertures located
on eight thoracic somite.Abdomen terminating in telson, a non segmented
structure having anus. A cuticular fold –carapace extending backward from
head; extends over and fuses with all thoracic somites; the first three pairs of
thoracic legs or maxillipeds associated with mouth, leaving remaining five pairs
of legs, hence the name Decapoda, for a variety of functions, including
locomotion, food gathering, defense and offence and grooming. Gills enclosed
in a branchial chamber, its lateral wall formed by a gill cover or branchiostegite,
an extension of carapace. (From various sources).
1.2
Classification
Below is given the position of the Caridea within the classification of the
Arthropoda, subphylum Crustacea. Martin & Davis (2001) use the classification
adapted here.
Subphylum Crustacea
Class Cephalocarida (cephalocarids)
Class Branchiopoda (fairy or brine, tadpole, clam shrimps; water fleas)
Class Remipedia (remipedes)
Class Maxillipoda (barnacles, copepods)
3
Class Ostracoda (mussel or seed shrimps)
Class Malacostraca
Subclass Phylocarida (leptostracans)
Subclass Hoplocarida (stomatopods or mantis shrimps)
Super order Eumalacostraca
Super order Syncarida
Super order Peracarida (mysidaceans shrimps, isopods, cumaceans,
amphipods)
Super order Eucarida
Order Euphausiacea (krill)
Order Amphionidacea (amphionidacean shrimps)
Order Decapoda
Suborder Dendrobranchiata (penaeoidean and sergestoidean shrimps)
Suborder Pleocyemata
Infraorder Caridea (caridean shrimps)
Infraorder Stenopodidea (stenopodidean shrimps)
Infraorder Thalassinidea (mud or ghost shrimps)
Infraorder Astacidea (crayfishes, clawed lobsters)
Infraorder Palinura (spiny or nonclawed lobsters)
Infraorder Anomura (false crabs, hermit crabs, squat lobsters)
Infraorder Brachyura (true crabs)
1.3 History of Nomenclature of Infraorder
Caridea
It was Dana (1852) who was the first to recognize the group as distinct
and named it Caridea. Although rejected by Boas (1880) another term coined
Eukyphotes by him, the name Caridea is accepted by Holthuis (1955,1993) who
considered the reason for rejection not to valid and gave a treatment of the
history of nomenclature and classifications of caridean families, superfamilies
and other higher categories. According to Holthuis (1993), Dana (1852) divided
his sub tribe Caridea into four families. Bate (1888) divided his Phyllobranchiata
Normalia which coincided exactly with Dana‟s Caridea into four tribes. Ortmann
(1890) divided the Caridea, which he named Eucypihdea into 13 families.
4
Ortmann until 1898 kept on giving new versions. Balss (1927) gave a
classification, which was largely based on Borradaile's (1907) system. In 1955,
Holthuis himself divided Caridea into superfamilies: Pasiphaeoida,
Oplophoroida, Stylodactyloida, Pandaloida, Psalidopodoida, Palaemonoida, and
Crangonoida. Schram (1986) recognized two Infraorders, the Procarididea and
Caridea, placing them in the Suborder Eukyphida, of Boas for Carideans and
latter resurrected by Burkenroad (1981) and Schram (1984).
Thompson (1967) revised the arrangement proposed by Holthius
(1955) by removing the Nematocarcinidae from the Oplophororidea and by
placing them in the Bresilioidea together with the Bresiliidae and the
Disciadidae. The Eugonatonotidae, were placed by him in the Oplophoroidea.
He established a new super family Heterocarpodoidea.
Bruce‟s excellent work during 1967 to 2004 on palaemonids spans over
three decades. Banner & Banner in the eighties established and described
new genera and new species ,subspecies and did vast amount of revisionary
work (Titgen, 1991).
In 1971 a most aberrant caridean was discovered in an anchialine
habitat in the mid- Ascension Island (Atlantic Ocean)by Chace & Manning
(1972) as a new genus (Procaris) and species described it, which represents a
new family Procarididae and a new super family Procaridoidea.
Forest (1977) abandoned the status of the super family Bresilioidea. He
placed the Disciadidae, the Bresiliidae, and the Nematocarcinidae in the super
family Oplophoroidea, while the Eugonatonotidae and Rhynchocinetidae were
assigned to the Palaemonoidea.
Bowman & Abele (1982) divided the Infraorder Caridea into 11 super
families
(Procaridoidea,
Atyoidea,
Stylodactyloidea,
Pasiphaeoidea,
Rhynchocinetoidea,
Palaemonoidea,
Psalidopodoidea,
Alpheoidea,
Pandaloidea, Physetocaridoidea, and Crangonoidea); this was followed in most
instances.
Hart & Manning (1986) established the family Agostocarididae for their
new genus Agostocaris, which was later placed by Chace (1992) in the
Bresiliidae.
In various papers published between 1987 and 1990, Christoffersen studied
with cladistic methods various groupings within the Caridea with most
unexpected results. In 1987, Christoffersen examined the super families
Crangonoidea and Alpheoidea. In the former the families Barbouridae
5
(Barbouriidae), Lysmatidae, Processidae, and Crangonidae were recognized,
while to the Alpheoidea were assigned the families Nauticarididae, Alopidae,
Bythocarididae, Thoridae, Hippolytidae, and Alpheidae.In 1988 Christoffersen
divided the family Crangonidae into subfamilies. Christoffersen revised the
super family Pandaloidea in 1989a. In the same year, Christoffersen (1989b)
revised the super family Atyoidea and divided it into the following families:
Oplophoridae, Atyidae (with subfamilies Atyinae and Xiphocaridinae),
Pasiphaeidae, Alvinocarididae, Bresiliidae, Psalidopodidae, and Disciadidae.
Finally, in 1990, Christoffersen added the family Agostocarididae to the Atyoidea
completing the review of super families and families, the super family
Stylodactyloidea was contained by him as two families Stylodactylidae and
Campylonotidae. The Palaemonoidea consisted of the families Palaemonidae
and Rhynchocinetidae only. The Eugonatonotidae and Nematocarcinidae were
ranked super families and the Pandaloidea was similarly treated as in 1989, but
to the Crangonoidea the families Merguiidae and Glyphocrangonidae were
added. The arrangement of the Alpheoidea differed from that suggested by
Christoffersen (1987) by the addition of the families Merihippolytidae,
Pterocarididae, and Ogyrididae.
Recent works has forced caridean workers to look more closely at
some of the Christoffersens”suggestions (De Grave et al,2009).Now the
knowledge of DNA sequences has become indispensable for basic taxonomical
research, or systematics. The advent of DNA sequencing has significantly
accelerated caridean research and discovery,for example Baeze et al(2009)
and Bracken et al(2009).
. It was felt as a relief by the carcinologists when Chace (1992)
published a well-reasoned classification of the Caridean families. Chace
reduced the family number to 28. Vereshchaka (1997) provided a key for
caridean super families modifying slightly that of Chace (1992); Martin & Davis
(2001) subsequent to Bowman & Abele (1992) have changed some super
families. In this monograph, Martin & Davis (2001) are followed. Bauer (2004)
converted Chace‟s classification and expressed a traditional view of caridean
relationship. He is of the opinion that hypotheses proposed by Chace (1992)
and Christoffersen (1990) must be tested independently using DNA and RNA
sequence data and it is very likely that these classifications will be sooner or
later modified and phylogenetic relationships which remain contested will be
clear as researches are yielding new insight into the evolutionary history of the
Caridea.
6
1.4 Taxonomic Characters and biological
success
The caridoid facies has made caridean shrimps to successfully adopt
aquatic environment by making them highly stream lined. The carideans come
out for on top with number of species as compared to euphausiaceans and
penaeoideans. The high biological diversity of carideans reflects their much
wider range of habitat occupation in comparison with other caridoid groups. The
cardioids body plan, modified in a variety of ways, has been the basis from
which extensive invasion into and diversification in the benthic environment
have occurred. Schematic diagrams of important families are produced here to
show the various modifications (Figs. 2-5).
Figure 2: External anatomy of a typical Crangonid
(Adapted from Holthuis,1993)
Complete set of morphological characteristics that adapt a true shrimp
so as well to forward and backward swimming is known as the caridoid facies as
7
already mentioned. The term “carid” comes from the Latin word for shrimps,
the suffix “oid” means like and facies refers to general appearance or aspect.
Crustaceans that mostly exhibit the caridoid facies or shrimp like body plan are
species found in the decapod malacostracan suborder Eucarida. Taxa within the
Eucarida exhibiting a classic caridoid body form are the euphausiaceans (krill),
mysidaceans, (opossum shrimps) and sergestoideans, penaeoideans,
stenopodideans and carideans. Two other non-natatory features of caridoid
character complex are stalked eyes and the biramous antennules. The
compound eye is placed on a movable stalk; the eyes can detect movement and
form images over a wide field of vision. The outer branch of the antennules
carries the aesthetics, or olfactory hairs, which are sensitive to a variety of
chemicals dissolved in water. Both the inner and outer rami in pelagic shrimps
are quite long flagelliform, carrying a variety of tactile or touch receptors as well
as chemoreceptors sensitive to higher concentrations of chemical stimuli. The
long flagella of the two antennae, can sweep all around the animal in order to
detect and recognize nearby objects.
Figure 3: External anatomy of a typical Alpheid
(adapted from Kim & Abele,1988)
8
Figure 4: External anatomy of a typical Palaemonid
Figure 5: External anatomy of a typical Pandalid
1.5
Body Plan
The body of typical caridean shrimp (Fig. 6) can be divided into a head,
thorax and abdomen or pleon.
9
The head and first three thoracic somites are fused into a
cephalothorax, and a dorsal shield known as carapace covers the head and all
the thoracic somites dorsally, which extend downwards laterally to cover the
gills and often the bases of the thoracic limbs called pereopods. Anteriorly, the
carapace may project as a narrow, often armed rostrum, laterally flattened, as in
some hippolytids, or dorso-ventrally flattened, as in some crangonids. It may
possess teeth on its dorsal and ventral borders (as in Palaemon), may be
unarmed and acutely pointed (as in Athanas), or may be unarmed except for
small, rounded apical teeth (as in Processa).Occasionally some of the dor s sal
teeth may be movable. The orbits are situated at each side of the base of the
rostrum and these depressions partly surround the eyes. In some species of
Alpheus, the carapace is produced anteriorly to cover the eyes in dorsal view,
forming orbital hoods. The carapace may be furnished with a number of spines.
Above the orbit may be one or more supra-orbital spines; below the orbit the
antennal spine arises in the line with the antennular peduncle. The
branchiostegal spine projects over the proximal part of the antennal peduncle; at
the antero-ventral corner of the carapace, there may be a pterygostomian spine.
On the lateral surface of the carapace, at a considerable distance behind the
anterior margin, there may be a hepatic spine, which is sometimes joined to the
anterior margin by a hepatic groove in the carapace. The branchiostegal and
hepatic spines are never found together. A groove running parallel but just
behind the edge of the orbit is known as the post- orbital groove. The carapace,
pleon, and limbs may bear longitudinal ridges or carinae; in the Crangonidae, for
example, certain species posses carinae on the carapace, armed with spines or
protrusions. The compound eyes normally consist of a stalk of two segments
plus a cornea of varying shape on the end of the stalk. In some families
(Hippolytidae and Palaemonidae), an accessory eye is present, the function of
which is still not clear.
10
Figure 6: Body and appendages . A- Carapace and abdomen, left lateral
view; B- Left mandible, dorsal view; C- Left first maxillula, dorsal view; DLeft second maxilla, dorsal view; E- Left first maxilliped, dorsal view; FLeft second maxilliped, dorsal view; G- Left third maxilliped, ventral view .
11
Figure 7: Appendages. A- Left antennule, dorsal view; B- Left antenna,
ventral view; C- Left first walking leg, medial view; D- Left second walking
leg, lateral view; E- Left third walking leg, lateral view. F- Left fourth
walking leg, lateral view. G- Left fifth walking leg, lateral view; H- Left first
pleopod of a female in breeding dress, anterior view; I- Left first pleopod,
anterior view of non-breeding female; J-N from male; J- Left first pleopod,
anterior view; K- Left first pleopod, anterior view; L- Left fourth pleopod,
anterior view; M- Left fifth pleopod, anterior view; N- Left uropod, dorsal
view; O- Telson, dorsal view; P- Endopod of left male second pleopod,
anterior view.
Appendages: The nineteen pairs of appendages are grouped as
sensory, cephalic, mouthparts and abdominal. (Table 1, Figs.6-7)
Table 1: Details of appendages of a shrimp
BODY
BODY REGION
SESEGMENT #
REGIONAL
SEGMENT #
APPENDAGE
1
Cephalon
1
Antennules (first antennae)
2
"
2
Antennae (second antennae)
12
3
"
3
Mandibles
4
"
4
Maxillules (first maxillae)
5
"
5
Maxillae (second maxillae)
6
Thorax
1
First maxillipeds
7
"
2
Second maxillipeds
8
"
3
Third maxillipeds
9
"
4
First pereopods (usually chelipeds)
10
"
5
Second pereopods (usually chelipeds)
11
"
6
Third pereopods (1st walking legs)
12
"
7
Fourth pereopods (2nd walking legs)
13
"
8
Fifth pereopods (3rd walking legs)
14-18
Abdomen
1-5
First-fifth pleopods (swimmerets)
19
"
6
Uropods
Projecting from the anterior region of the carapace are two
sensory appendages, the antennules and the antennae. Antennules have a
stem, or peduncle of three segments, with two multi-articulate flagella arising
from the third, most distal segment. In the Palaemonidae and Hippolytidae,
certain genera have the outer flagellum bifid, the antennules thus becoming
triramous.The aesthetascs or chemo and mechano- receptors exhibit diversity in
number and shape depending on habitat and environment. In marine shrimp
both adult and larvae they are more numerous, longer, slender, and uniformly
thin walled. In adult freshwater shrimps, they are fewer, shorter, divided in thick
walled stalk and thin distal cob like portion structurally different and positioned
differently in freshwater families Atyidae (Caridina) and palaemonid genus
Macrobrachium(Shenoy et al,1993). The first segment of the antennular
peduncle may be laterally expanded to form a lobe or a spiniform process
13
known as stylocerite. This is of considerable importance in the taxonomy of
certain families (e.g. Crangonidae). The antennae comprise a long flagellum
arising from a peduncle and sometimes longer than the body of animal, plus an
exopod, which is laterally expanded and flattened into a scale-like process, the
scaphocerite. The latter varies in shape, is normally setose on its inner margin,
and in the most carideans has a spine on the external anterior corner.
The mouthparts, which deal with the handling and initial mechanical
breakdown of food, comprise a pair of mandibles, two pairs of maxillae from the
cephalic region and three pairs of maxillipeds. The maxillipeds are modified
from the three anterior pairs of thoracic limbs of the ancestral caridoid form and
are thus not appendages derived from somites of the head. The mandibles have
a grinding molar process, which is widely separated from a toothed incisor
process, used for cutting food fragments. Certain species also posses a
mandibular palp, which is often of taxonomic importance. In the Crangonidae,
both incisor process and palp are absent. The genus Hippolyte lacks the palp
only, whereas in some pasiphaeids the incisor process only is present. The first
maxilla is a flattened structure with a palp, and the second maxilla bears a
pointed lamellar expansion on its outer border forming the scaphognathite,
which by its rhythmic beating, creates water current over the gills. Following the
maxillae are the first three pairs of thoracic appendages-the maxillipeds as
mentioned earlier.
The first maxilliped has lost any vestige of its original pediform shape
and has become expanded and leaf-like. At the base of exopod is a process
known as the caridean lobe, this is the characteristics of the carideans as
mentioned earlier. The second maxilliped appears more limb-like, but the least
modification has occurred within the third maxilliped, which has retained its
pediform character, being frequently quite long and projecting beyond the
scaphocerite in some instances. The apical joint or dactylus may be absent or
fused to the next joint, and the third and fourth joints may also be fused. The
appendage may be quite heavily armed with spines or tubercles. The first two
pairs of maxillipeds are very much associated with food handlings. The exopods
have more specialized functions like blowing away the debris and preventing the
fouling.
The pereopods comprise the five pairs of thoracic appendages, usually
elongate limbs, used for feeding and locomotion. Each limb comprises seven
joints from the body outwards; these are termed the coxa, basis, ischium,
merus, carpus, propodus, and dactylus. In certain families (e.g. Hippolytidae),
the third joint or carpus of second pereopod may be subdivided, and this is
important in taxonomy of the group. In the Processidae, both the carpus and
merus of second pereopod are often subdivided into many joints. First and
14
second pereopods are usually furnished with terminal pincers, or chelae. In
Palaemonidae, the second pair is sexually dimorphic and in Alpheidae, they are
asymmetrical. In the Pandalidae, however, the chelae of the first pereopods are
microscopically small, and in the Processidae, a minute chela is present on one
of the first pereopods, whilst other ends in a simple pointed dactylus. In the
Crangonidae, the first pereopods are robust and sub-chelate, the dactylus and
propodus articulating at the right angles to the long axis of the limb. The long
and slender “fingers” of the chelae of first and second pereopods, the cutting
edges of these fingers being pectinate characterize the Pasiphaeidae, or comblike arising from the basis of some or all of the pereopods may be limb-like
processes, which are the exopods of the pereopods, relicts of the original
biramous nature of the limbs. Exopods are usually much reduced, although, in
the Pasiphaeidae, they may be quite long. Another spectacular variation of the
chela is found in the family Atyidae, where the fingers of the chelae terminate in
brushes of long complex setae. Certain genera (e.g. Pandalus )posses epipods,
which are small strap-like projections issuing posteriorly from the coxae of the
pereopods and often from the maxillipeds.
The abdomen or pleon is made up of six somites, the pleonites, the
sixth of which is modified to form, with the tail or telson, the “tail-fan” of the
animal. All pleonites are more or less extended laterally and downwards into
pleura. As already pointed out the pleonite second in carideans has a pleura
which overlaps those of pleonite first and third dorsally, pleonite third may be
extended posteriorly over part of the fourth pleonite (as in some hippolytids).
First to fifth pleonites each bears a pair of ventral pleopods, which are leaf-like,
biramous limbs used for swimming in some species. Each pleopod is made up
of a broad, flat endopod and an equally broad exopod. On the internal borders
of the endopods of pleopods 2-5 are found small, tubular appendages with
hooked ends, these hooks are termed cincinnuli. Each one of these
appendages is an appendix interna, and they serve to hook together laterally
adjacent endopods, thus improving the swimming beat of the pleopods and their
synchronization. It is significant that, in some wholly benthic species (e.g.
Crangonidae), the appendices internae are absent. The first pleopods differ
between the sexes in these decapods. In the male, the endopod is important in
copulation, usually having a modified terminal process, which is used when
pairing takes place. The second pleopod of the male also bears appendage,
which is used in copulation, and sperm transfer. This is the appendix masculina.
The endopod of the females‟ second pleopod is normally broader and more leaflike than that of the male and the pleopod does not bear the appendix
masculina. The appendages of the sixth pleonite are known as uropods and are
broad and foliaceous, projecting on each side of telson. The telson and uropods
make up the earlier mentioned “tail-fan” of the animal, which is used for rapid
15
escape movements by sudden flexor of pleonites, propelling the animal
backwards. There may be a varying numbers of spines on the dorsal surface,
on each lateral border of the telson, plus spines on the apex. This apex may be
flat, concave, deeply cleft, or convex. The exopod has a row of spines, generally
known as diaresis.
Gills: There are three types of gills found in these decapods; their
arrangement is often of taxonomic importance. Pleurobranchs are gills attached
to the sides of the body and occur dorsal to the pereopods. They are long gills
and are covered by the downward and lateral expansion of the carapace, which
thus forms a branchial chamber around them. Arthrobranchs are small gills,
which arise from the membrane joining the coxa of a pereopod to the animal.
Podobranchs are small gills, which differentiate from the epipods, and are thus
found on the coxae of limbs. Arthrobranchs and podobranchs are often
associated with the maxillipeds and pereopods, whereas pleurobranchs are
normally associated with pereopods only.
Genital openings: The genital openings differ between the sexes;
female ducts open into small apertures on the coxae of the third pereopods. The
male ducts open on the coxae or on medial surface of arthrodial membrane
between the coxae of fifth pereopod and the sternum, with a gonopore flap.
Sexual dimorphism is found in certain genera, sometimes only in the breeding
females (several species of Leptochela)
Protandry is scattered among caridean families particularly in
Pandalidae, Hippolytidae, Processidae, Crangonidae and Campylonotidae, that
for confirm sex change (Bauer, 2004)
The species evolve specialized phenotype to adapt local environmental
conditions.
1.6
Biology
Caridean Atyidae further study is necessary to shrimps occur in all the
world‟s oceans where they may be found intertidally, in the shallow sub littoral
zone and down to the abyssal depths. Some are pelagic, others are benthic,
and few have specific association with other invertebrates. The shallow sub
littoral species use the profuse algal growth as cover from predators. Some may
make shallow burrows covering themselves with sand to avoid predators
.Several species occur on offshore ground, which has good hydroid cover.
Offshore gravel may be rich in species, very fine sand, silt deposits, deep water
16
mud, hydrothermal vent and oozes are habitats for certain groups. Some are
found in the lower reaches and on the rocky ledges. Farther up estuary some
may be found being able to withstand wide range in salinity. On salt marshes
and in brackish pools with indirect contact with the sea, some paleamonids do
occur. In deep waters off shore, several species undergo extensive diurnal
vertical migrations. Rhythmic patterns of activity have been demonstrated in
palaemonids, becoming more active as the tide begins to ebb. Some carideans
are subterranean or troglobic. Taxonomy of the most of the troglobic species
points to a very long isolation. Some species also occurring in our area like
Caridina nilotica , Macrobrachium lamarrei have been found incidentally in
subterranean waters (Holthuis, 1964: 64). One family Alvinocarididae are
distinct component of chemosynthetic communities occurring at hydrothermal
vents or waters influenced by brine or hydrocarbon seeps (Komai & Segonzac,
2003).All oplophorid genera, few pandalids, one pasiphaeid and one
thalassocaridid produce luminous secretion. The secretion may be quite viscous
and quite retain its luminosity for many minutes .It is suggested that this
secretion is a hepatic product regurgitated from the mouth (Herring, 1985). The
secretion is probably a defensive response and can be sometime seen elicited
with a flashlight (Herring in Barnard, 1985). In addition to these cuticular
photophores are also present in few oplophorid genera. The luminescence of
these crustaceans has some commercial application to detect shoal size.
The aesthetascs or olfactory hair, setae and sensory (chemo
and mechano) receptors on antennule are frequently groomed with cleaning
brushes on the third maxillipeds. The long flagella of the antenna have vital
taste and touch receptors, frequently scrubbed and preened by the carpalpropodal cleaning brushes of the first legs. Chemoreceptors on the terminal
segment present in high concentration are scrubbed again by the third
maxillipeds. The aesthetascs of these shrimps play an important role as salinity
indicators (Shenoy etal, 1993). The other receptors comprise statocysts, the
stalked eye, and the dorsal organ.
Many pelagic carideans are active predators. Often these species take
part in extensive upward vertical migration at night, swimming down to the
bottom at dawn again feed at night on planktonic small crustaceans. Among
benthic species, types of food vary from group and season.
Shrimps posses rigid exoskeleton and can only grow by moulting, or
casting the outgrown skeleton and increase in size while the new skeleton is still
soft. The moult cycle is a complex one and dominates the life of the animal,
since the behavior, habits and physiology differs at different times of cycle. The
actual process is quite rapid, normally lasting from 9-25 seconds. Complete
hardening of the skeleton takes about two days. Due to fear of predators the
17
newly moulted animal becomes even more secretive than usual. Moulting is
normally of seasonal occurrence.Females prawns frequently grow at faster rate
than males and as such, the females of most species are larger than the males
at any given age. Moulting stops when a female carries eggs on its pleopods.
Certain features like small white spots are noticeable before moulting in
appearance.Their presence gives a useful indication of moult‟s cycle event,
especially in the field. No annual growth indicators are found in shrimps so age
of shrimp is difficult to estimate. Many carideans are colour changers, colour
modification mainly functions for concealment from visually hunting predator
such as fishes or birds. Some shrimps loose their colour at night. The colour
pattern is often a good species character; however, there may be striking
variation in colour pattern among individuals of the same species. Some species
produce sound.The rapid closure of chela results in a loud sound or snap.The
snap is a behavior observed most commonly during agonistic interactions.
Aggregations of animals sometime occur offshore, usually connected
with the onset of the breeding season. The shrimps are dioceous (gonochorus)
and exhibit sexually dimorphic characters. Intersexuality is common in certain
groups because of protandrous sex reversal. In some groups, for example
hippolytid and pandalid populations contain a varying percentage of primarily
females, which remain females throughout their life. This percentage may vary
with latitude. Copulation and fertilization is dependent upon the female moulting.
It occurs in such a position that the spermatophores are deposited on the
sternum of the female between fifth pair of legs. The appendix masculinae of the
males are involved in the placing of spermatophores. The latter are extruded
from the openings on the coxae of the fifth legs. Carideans have „unistellate‟ or
thumbtack shaped, completely immobile spermatozoa (Felgenhauer & Abele,
1983) with a main body and a spike. There are striking exceptions. The sperms
are packed in twin but simple spermatophores, one from each male gonopore.
Spawning of the eggs occurs soon after copulation, depending on the group at
the minimum 2-3 to 48h. Before spawning females usually engage in cleaning
activity, first two legs are being used to clean the pleopods, the pleon being
flexed to allow the action. For spawning females for different species take
different position - crangonids lie on inner side and bend the pleonites beneath
the thorax, palaemonids remain standing on spread out legs and process the
telson against the substratum. The pleon assumes a week S-shaped curve and
in this position, the eggs are extruded. The eggs released from the openings
pass over the spermatophores attached to the female sternum are fertilized
during this passage. They are passed backwards by the endopods of the first
pleopods and become attached to the endopods of pleopods 1-4, special
ovigerous setae which appear at the precopulatory moult of the female,
disappear with the next moult. The three anterior pleonites widen slightly and
18
the pleon deepens. Females fan the pleopods and groom the embryos with
cleaning chelipeds that is why the embryos stripped from female suffer heavy
mortality from microbial fouling in the absence of the female grooming. The
shrimps may locate odour of sex pheromones carried by water current and
move upstream by sing a positive rheotaxis.
The mating system of Macrobrachium rosenbergii, the giant freshwater
shrimp is a bit more complicated. There are three male morph types all capable
of mating, all without distinct genotypes: the large males having hypertrophied
second cheliped, small males with proportionately smaller chelipeds, and
intermediate size males with relative chelipeds size. All three types show
different mating system, when large males are finished due to one or other
reason the other size move the male social ladder, intermediate under go
changes and assume large males and small males to intermediate size. The
physiological changes responsible for this transformation are still unknown
(Bauer, 2004). Monogamy as extended guarding of female by the male has
been observed in Hymenocera picta(Seibt & Wickler, 1979).Protandry is
scattered
among
caidean
families
particularly
in
Pandalidae,Hippolytidae,Processidae,Crangonidae
and
Campylonotidae(Bauer,2004)The genera Exhipplysmata and Lysmata are
described as simultaneous protandric hermaphrodite (Kagwade,1982) or
euthermaphrodite(Fielder et al,2010) .
A
B
C
D
Figure 8: Life cycle of caridean. A- Egg; B- Larva; C- Post larva; DAdult(Adapted from New &Singholka,1982)
19
A typical caridean embryo hatches out of its surrounding membrane,
moults the embryonic cuticle, and enters the water as planktonic larva. The
larva is termed a zoea; it undergoes a metamorphosis in which larval
characteristics are lost. The young juvenile is recruited into the adult population
(Fig. 8). There is variation in life histories characteristics with latitude in near
shore, sub littoral marine carideans (Bauer, 2004 ,Table 6). The number of
larval instars varies from 5-9. Carideans with multiple generations usually have
one long term and overwintering generation and one short term summer
generation or mix (Omori et al, 1994). In many caridean species the number of
instars has been reduced, and is abbreviated. In the Palaemoninae, there is
direct relationship between abbreviated development and strictly freshwater
habitat. In some genera, for example Synalpheus all forms of development from
extended to abbreviate to direct occur.
The most obvious parasites of these shrimps are the bopyrid isopods
which often occur in branchial cavity of the shrimp, causing distortion in the form
of an obvious sewelling”Cheek” on one side of the carapace (Pl.1).
Bourdon (1968) has noted the affect that parasites have on their hosts,
usually the gonads are affected, and the morphological changes were noticed in
the secondary sexual characters (Kazmi, 1996) (Pl.1).
The bopyrids, which infest carideans all, belong to subfamily Bopyrinae
(Fig. 9). The Pakistani carideans: Saron, Alpheus, Synalpheus , Macrobrachium
, and Latreutes have been found bopyrized (see Parabopyriscus sp,
Probopyrus pica , Parabopyrella indica , P.niertsrazi and P.saronae in Kazmi et
al., 2002;Yaqoob,2006). Other less obvious parasites are hemiarthid isopods
(only once found during the present studies),ellobiopsid protozoans,
turbellarians, rhizocephalan cirripedes and nematodes (Smaldon et al, 1993).
20
Figure 9: Bopyrid parasites in pairs, females in dorsal and ventral views,
males in dorsal view A-C- From host Macrobrachium (After Chopra, 1923),
D-F- From host Alpheus (After Chopra, 1923), G- I- From host Synalpheus
(After Kazmi et al, 2002); J-L-From host Saron (After Bourdon & Bruce,
1979).
There are some amazing relationships between carideans and other
invertebrates and fishes. The invertebrate symbionts with the carideans are
corals, anemones, sponges, sea urchins, feather stars, clams, jellyfish, other
decapods, stomatopods, and echiuroid worms. Association of hippolytid
(Latreutes ) and medusa of Catostylus mosaicus was reported by Kazmi (1996)
(Pl.1). Shrimp symbioses are rarely reported in deepwater, polar, colder, and
freshwater environment. This may be explained by the absence or low
abundance and diversity in freshwater of the large sessile invertebrates that
serve as host in marine habitat.
21
22
CHAPTER -2
Materials and Methods
Materials: The results in the monograph are based on the new
samples collected by the authors, with the help of the Karachi university staff
and local fishermen by using one of these gears: Dragnets, Jali, Castnets, Dip
nets and Bottom nets or purchased from the fish markets, or samples sent for
study to the university from cruises ( Fridjtof Nansen Cruise, 1977) (Fig.10
Map)and Discovery cruise( 1993).The old unidentified collection housed in the
repositories of the Marine Reference Collection and Resource Centre, the
Zoology Departments of University of the Punjab , Lahore, Bahauddin Zakria
University, Multan , University of Karachi and the Zoological Survey
Department, Government of Pakistan were also studied. The Discovery cruise,
1993, samples although were from Oman, however helped in the biogeographic
studies.
23
Figure 10: Geographical distribution of the stations sampled by R/V Dr.
Fridtjof Nansen Cruise No. 1 January (1977- Joint NOARD/ Pakistan
project) showing Leptochela presence.
Methodology: The diagnoses of all higher taxa included in the text are
taken and adapted from already recognized characters used in literature and
thus a standard terminology is used. The information given in the monograph
are based on the material collected by the authors from coastal and inland
waters of Pakistan and based on a thorough review of the relevant literature,
which included scattered records of the presence of the various species shrimps
collected from various localities in Pakistan. The species are described with
synony
ms and detailed descriptions supplemented by information,
when available on colour, size, habitat, and geographical distribution. Each
species is fully illustrated. Some were photographed alive prior to preservation.
The genera and species itemized following the keys are those known for
Pakistan, whether or not seen and encountered by the authors. Almost all the
other authors‟ species from Pakistan are included without verification that was
not possible due to non-availability of their material. The status of many species
24
is still unclear which is why symbol “cf” if often is associated. It cannot be
said about these indeterminate records weather they refer to either to the new
species described herein, or other described or undescribed carideans of
indeterminate genus or to combination of both. They are included here for the
sake of completeness and to bring attention to their occurrence in the event that
more specimens eventually are found. The families are arranged in sequence
by order of appearance in the key to the super families and families. The family
account includes diagnosis and key to the Pakistani genera. Generic entries
comprise diagnosis, identification of the type species, the gender of the name
and key to the species. There has been an attempt to list important references
under the species. Literature cited is a full list of sources used. Extra information
is given under “Remarks” where necessary or appropriate. The biogeography is
traced for all species. A chapter on commercial species in our water bodies is
added to highlight the utilization of these shrimps.
The type specimens of the new species are housed in the
Marine Reference Collection and Resource Center, University of Karachi.
25
CHAPTER -3
Results
3.1Biodiversity
There are now at least 34 families, 351 genera and 3,268 described
species in the Caridea ( Zoo.Rec.,2001; ITIS;De Grave, 2008).However truly
exact counts are impossible because of occasional omission in the Zoological
Records and subsequent rejection or change of names by taxonomists. Out of
them 14 families, 35 genera, 84 species, and 8 subspecies are found in
Pakistani water bodies, both fresh and marine as reported here.
List of Pakistani species:
Super family Pasiphaeoidea Dana, 1852
Family Pasiphaeidae Dana,1852
Genus Eupasiphae Wood Mason & Alcock, 1893
Eupasiphae gilesi(Wood Mason,1892)
Genus Leptochela Stimpson, 1860
Leptochela irrobusta, Chace, 1976
Leptochela nasimae sp.nov
Leptochela pugnax de Man, 1916
Leptochela cf sydniensis, Dakin& Colefax, 1940
Genus Pasiphaea Savigny, 1816
Pasiphaea alcocki (Wood Mason & Alcock, 1891)
Pasiphaea sivado (Risso, 1816)
26
Super family Oplophoroidea Dana, 1852
Family Oplophoridae Dana, 1852
Genus Acanthephyra A. Milne Edwards, 1881
Acanthephyra eximia Smith, 1884
Super family Atyoidea de Haan, 1849
Family Atyidae de Haan, 1849
Genus Caridina H.Milne Edwards, 1837
Caridina babaulti babaulti Bouvier, 1918
Caridina nilotica (Roux, 1833)
Caridina sumatrensis de Man, 1892
Genus Caridella Calman, 1906
Caridella sp
Genus Kalriana Zuberi, 1990
Kalriana anissi Zuberi, 1990
Kalriana jhimpirensis , Zuberi, 1990
Kalriana karachi Zuberi, 1990
Kalriana sunahrensis Zuberi, 1990
Super family Nematocarcinoidea S.I. Smith, 1884
Family Rhynchocinetidae Ortmann, 1890
Genus Rhynchocinetes H. Milne Edwards, 1837
*Rhynchocinetes uritai (Kubo, 1940)
Super family Palaemonoidea Rafinesque, 1815
Family Palaemonidae Rafinesque, 1815
Subfamily Palaemoninae Rafinesque, 1815
Genus Exopalaemon Holthuis, 1950
Exopalaemon styliferus (H.Milne Edwards, 1840)
Genus Leandrites Holthuis, 1950
27
Leandrites celebensis (de Man, 1881)
Genus Leptocarpus Holthuis, 1950
Leptocarpus potamiscus (Kemp, 1917)
Genus Macrobrachium Bate, 1868
Macrobrachium altifrons ranjhai Tiwari, 1963
Macrobrachium dayanum (Henderson, 1893)
Macrobrachium dacqueti (Sunier, 1925)
Macrobrachium equidens (Dana, 1852)
Macrobrachium idella idella (Hilgendorf, 1898)
Macrobrachium lamarrei lamarrei (H.Milne Edwards, 1837)
Macrobrachium lamarrei korangii sub. sp nov.
Macrobrachium lehiai sp nov.
Macrobrachium mal colmsonii malcolmsonii (H.MilnEdwards,
1844)
Macrobrachium malcolmsonii kotreeanum Johnson, 1973
Macrobrachium naso (Kemp, 1918)
Macrobrachium naraensis Siddiqui, 1980 (Name only)
Macrobrachium rude (Heller, 1862)
Macrobrachium scabriculum (Heller, 1862)
Macrobrachium shahpuri n.sp
Macrobrachium sp
Macrobrachium taunsii n.sp
Macrobrachium tirmiziae Yaqoob & Kazmi (MS), 2000
Genus Nematopalaemon Holthuis, 1950
Nematopalaemon tenuipes (Henderson, 1893)
Genus Palaemon , Weber, 1795
Palaemon pacificus (Stimpson, 1860)
28
Palaemon semmelinkii (de Man, 1881)
Palaemon sewelli (Kemp, 1925)
Subfamily Pontoniinae Kingsley, 1878
Genus Anchistus Borradaile, 1898
Anchistus custos (Forskal, 1775)
Genus Cuapetes Clark,1919
Cuapetes elegans (Paulson, 1875)
Cuapetes longirostris (Borradaile, 1915)
Cuapetes seychellensis (Borradaile, 1915)
Family Hymenoceridae Ortmann, 1890
Genus Hymenocera Latreille, 1819
*Hymenocera picta Dana, 1852
Family Gnathophyllidae Dana, 1852
Genus Gnathophyllum Latreille, 1819
*Gnathophyllum americanum Guerin-Meneville, 1855
Super family Processoidea Ortmann, 1896
Family Processidae Ortmann, 1896
Genus Processa Leach, 1815
Processa compacta Kensley, 1971
Super family Alphoidea Rafinesque, 1815
Family Ogyrididae Stebbing, 1914
Genus Ogyrides Stebbing, 1914
Ogyrides orientalis Stimpson, 1860
Ogyrides saldanhae Barnard,1947
Family Alpheidae Rafinesque, 1815
Alpheid sp
Genus Alberta n.gen
29
Alberta banneri n.sp.
Genus Alpheus Fabricius, 1798
Alpheus alberti Kazmi, 1974
Alpheus cf barbatus Coutiere, 1897
Alpheus bisincisus de Haan, 1850
Alpheus chiragricus H.Milne Edwards, 1837
Alpheus edwardsii (Audouin, 1826)
Alpheus isodactylus Afzal ,Javed&Barkati, 1986
Alpheus lobidens de Haan, 1849
Alpheus manorensis Afzal ,Javed&Barkati, 1986
Alpheus pacificus Dana, 1852
Alpheus pseudoedwardsii Afzal, Javed&Barkati , 1986
Alpheus strenuus strenuus Dana, 1 852
Alpheus splendidus Coutiere, 1897
Alpheus zulfaquiri Kazmi, 1980
Genus Athanas Leach, 1814
Athanas arabicus Afzal Javed&Barkati , 1986
Athanas dimorphus Ortmann, 1894
Genus Automate de Man, 1888
Automate dolichognatha de Man, 1888
Genus Salmoneus Holthuis, 1955
Salmoneus brevirostris (Edmondson, 1930)
Salmoneus cristatus (Coutiere, 1896)
Genus Synalpheus Bate, 1888
Synalpheus neptunus Coutier, 1899
Synalpheus thai Banner & Banner, 1966
Synalpheus tumidomanus (Paulson, 1875)
30
Family Hippolytidae Bate, 1888
Genus Exhippolysmata Stebbing, 1915
Exhippolysmata ensirostris ensirostris (Kemp, 1914)
Genus Heptacarpus Holmes, 1900
Heptacarpus pandaloides (Stimpson, 1860)
Genus Hippolyte Leach, 1814
Hippolyte ventricosa H.Milne Edwards, 1837
Genus Latreutes Stimpson, 1860
Latreutes anoplonyx Kemp, 1914
Latreutes mucronatus (Stimpson, 1860)
Genus Lysmata Risso, 1816
Lysmata vittata(Stimpson,1860)
*Lysmata amboinensis (de Man,1888)
Genus Saron Thallwitz, 1891
Saron marmoratus (Olivier, 1811)
Super family Pandaloidea Haworth, 1825
Family Pandalidae Haworth, 1825
Genus Heterocarpus A.Milne Edwards, 1881
Heterocarpus laevigatus Bate, 1888
Super family Crangonoidea Haworth, 1825
Family Crangonidae Haworth, 1825
Genus Philocheras Stebbing, 1900
Philocheras parvirostris Kemp, 1916
Genus Pontocaris Bate, 1888
Pontocaris pennata Bate, 1888
*Records of all species marked by an asterisk are based upon a close-up under water photographs taken by Pakistani scuba
diver (Ali, 2006 unpublished list) therefore the information cannot be relied upon for identification; hence, the species are not
31
discussed elsewhere in the species account. However, they are included in the list for the sake of comprehensiveness, for
their verification further collection is desirable
The following key to the subfamilies, families, and super families is
largely based on published by Holthuis (1993), nearly quoted verbatim. When
one family is present in a super family, it is given immediately .
KEY TO THE SUPERFAMILIES, FAMILIES, AND SUBFAMILIES OF
CARIDEA (An asterisk marks Pakistani taxa)
1.
None of the pereopods chelate or subchelate. Third maxilliped
composed
of
seven
free
segments--------------Super
family----------PROCARIDOIDEA-----------Family Procarididae.
-Chelae present on first or second pereopods or on both. Third maxilliped with
fewer than seven free segments---------------------------------------------------------------------------------------------2.
2. First pair of pereopods chelate or simple------------------------------------------------------------ 3.
-First pair of pereopods subchelate or prehensile -----------Super family
CRANGONOIDEA--32.
3.First and second pereopods similar, with long, slender fingers that are
pectinate by the presence of long, narrow teeth on the cutting edge. Second
maxilliped without exopod -----------Super family PASIPHAEOIDEA--------------------------------------------------*Family Pasiphaeidae.
32
-Fingers of first and second pereopods not all pectinate with long slender
teeth; those legs often very dissimilar-----------------------------------------------------------------------------------------------4.
4.Carpus of second pair of pereopods entire, not subdivided. First pair of
pereopods always with well developed chelae----------------------------------------------------------------------------------5.
-Carpus of second pair of pereopods usually subdivided into two or more
segments; if not, first pair of pereopods not chelate --------------------------------------------------------------------------------26.
5. Last two segments of second maxilliped implanted side by side at end of
antepenultimate segment. Pereopods 1 and 2 similar; fingers extremely long
and slender, more than 10 times as long as high, and more than 5 times as long
as the short palm, without teeth but with long hairs--------- Super family
STYLODACTOIDEA---------------------------------Family Stylodactylidae.
Last segment of second maxilliped attached to penultimate, not
touching the antepenultimate. Fingers of first and second pereopods not
extremely long ---------------------- 6.
6.
First pair of pereopods with both fingers of chelae movable---Super
family---------- PSALIDOPODOIDEA-------------------------------------------------------------- Family Psalidopodidae.
-Chela of first pereopod with only one movable finger, the other finger is
immovably fused with the palm--------------------------------------------------------------------------------------------------------------7.
7. Epipods present on the pereopod, terminating in a naked appendix which
extends vertically far into the branchial chamber posterior to the corresponding
pleurobranch.First and second pereopods similar ----------Super family
OPLOPHOROIDEA----------------------------------------------------------------------------------------------------------------------------------*Family Oplophoridae.
Epipods of the pereopods, if present, not terminating in a long naked
appendix-----------8.
33
8.
First and second pereopods with the chelae similar, the fingers
usually with dense tuft of setae at the apex---------------- Super family
ATYOIDEA---------------------*Family Atyidae----9.
First and second pereopods without dense tufts of setae at the end of
the fingers--12.
9.
Supraorbital spines present on the carapace. Exopods on at least the
first two pereopods. Diaeresis with only one or two lateral spinules----------------Subfamily Paratyinae.
Supraorbital spines absent-------------------------------------------------------------------------10.
10.
Diaeresis with only one or two lateral spinules. Exopods usually present
on at least first and second pereopods ------------------------------------------------------Subfamily Typhlatyinae.
Diaeresis usually with numerous spinules along its entire length or the
distal part of it. No exopods on any of the pereopods----------------------------------------------------------------------11.
11.
Branchial formula incomplete, at most eight pairs of branchiae present.
No arthrobranch on the first pereopod-----------------------------------------------Subfamily Caridellinae.
-Branchial formula complete, 9 pairs of branchiae. An arthrobranch at the base
of the first pereopod----------------------------------------------------------------------------------- Subfamily Atyinae.
12.
First pair of pereopods stronger and heavier, though often shorter than
second ----13.
First pair of pereopods usually more slender than, rarely subequal to
second pair--17.
13.
Pereopods without strap-like epipods. Mandible with molar process
conical, laminar or vestigial----------------------- Super family BRESILIOIDEA---------------------- Family Bresiliidae.
34
-Strap-like epipods on at least the three anterior pairs of pereopods. Mandible
with molar process blunt, sub truncate with ridged grinding surface-----Super
family NEMATOCARCINOIDEA------------------------------------------------------------------------------------- 14.
14.
Rostrum finely dentate. Anterior two pairs of pereopods slender, fingers
not bearing conspicuous long spines---------------------------------------------------------------------------------------15.
Rostrum grossly dentate. Fingers of the chelipeds with lateral and
terminal spines distally, forming a basket-like cage when the fingers are closed -----------------------------------16.
15.
Marine, often deep-sea species. Third pereopod with dactylus simple,
unarmed. Last segment of second maxilliped applied as a narrow strip against
the distal margin of the much broader penultimate segment--------------------------------------------------Family Nematocarcinidae.
Fresh water species. Third pereopod with prominent curved spines on
the posterior margin of the dactylus. Last segment of second maxilliped longer
than broad and longer than the penultimate segment to which it is attached with
its narrow side---------------------------------------------------------------------------------------------------------------------------------Family Xiphocarididae.
16.
Rostrum movable or at least incompletely fused with the rest of the
carapace. Carapace without lateral ridges. Pereopods without exopod--------------------------------*Family Rhynchocinetidae.
Rostrum completely fused with the rest of the carapace, immovable.
Carapace with three strong longitudinal ridges on the lateral surface. All
pereopods with exopods----------------------------------------------------------------------------------------------------------Family Eugonatonotidae.
17.
Anterior four pairs of pereopods with an arthrobranch each. Dorsal
antennular flagellum simple, unbranched. Mandible not bifurcate----- Super
family CAMPYLONOTOIDEA-------------------------------------------------------------------------------------------------------------------------18.
35
No arthrobranchs on any of the pereopods. Dorsal antennular
flagellum provided with an accessory branch. Incisor and molar processes of
mandible distinctly separated (incisor process sometimes reduced) ---------------------- Super family PALAEMONOIDEA--------------19.
18.
Epipods present on the pereopods. Pereopods of the second pair sub
equal. Second maxilliped with the terminal segment applied as a narrow strip to
the mesial margin of the penultimate segment ------------------------------------------------------------- Family Campylonotidae.
Pereopods without epipods. Pereopods of the second pair very
unequal. Second maxilliped with the terminal segment forming an oblique
juncture with the penultimate segment-----------------------------------------------------------------------Family Bathypalaemonellidae.
19.
Third maxilliped slender, pereopod-like. First maxilliped with caridean
lobe of exopod distinctly overreaching the endite. Mandible usually with
prominent incisor process-----------20.
Third maxilliped with antepenultimate segment broadened, at least
proximally, sometimes operculate. First maxilliped with caridean lobe of exopod
not distinctly overreaching endite. Mandible with incisor process vestigial or
absent--------------------------- 25.
20.
First maxilliped with exopodal lash vestigial. Mandible with molar
process flared distally------------------------------------------------------------------------------------- Family Anchistioidae.
First maxilliped with exopodal lash fully developed. Mandible with molar
process conventional, not flared----------------------------------------------------------------------------------------21.
21.
Second maxilliped with terminal segment broadly ovate, penultimate
segment convexly produced mesiad, causing endopod to appear bilobate
distally. First maxilliped with palp broadly ovate.Supraorbital spines present----------------------------Family Desmocarididae.
36
Second maxilliped not markedly bilobate distally. First maxilliped with
palp not unusually broad------------------------------------------------------------------------------------------------- 22.
22.
First maxilliped with caridean lobe of exopod acutely produced
distally------- Family Typhlocarididae-------------------------------------------------------------------------------------------------23.
First maxilliped with caridean lobe of exopod not acutely produced
distally------------------Family Palaemonidae---------------------------------------------------------------------------------------24.
23.
Carapace with a longitudinal complete post-antennal suture on either
lateral surface. Third antennal flagellum partially fused with dorsal antennal
flagellum -----------------------------------------------------------------------------------------------------------------------Subfamily Typhlocaridinae.
Carapace without complete longitudinal suture. The three antennular
flagella entirely free, the third not fused with any of the others-------------------------------------------Subfamily Euryrhynchinae.
24.
Telson with two pairs of posterior spines and with one or more pairs of
hairs. A pleurobranch at the base of the third maxilliped-------------------------*
Subfamily Palaemoninae.
Telson usually with three pairs of posterior spines. Third maxilliped
without pleurobranch -------------------------------------------------------------------------*Subfamily Pontoninae.
25.
Third maxilliped with antepenultimate segment clearly articulated with
and much wider than next proximal segment--------------------------------------------*Family Hymenoceridae.
Third maxilliped with antepenultimate segment fused with and not much
wider than next proximal segment ------------------------------------------------------------------------------ *Family Gnathophyllidae.
37
26.
Carapace merging anteriorly into an inflated, indiscrete rostrum.
Second pereopod with fixed finger curving subrectangularly around the short
movable finger-------------Super family PHYSETOCARIDOIDEA------------------------------------------------Family Physetocarididae.
Rostrum, if present, discrete, not an inflated extension of the carapace
proper. Second pereopod, if present, with conventional chelae----------------------------------------------- 27.
27.
Right first pereopod chelate, left usually simple, terminating in a plain
claw-like dactylus; if both first legs chelate, the rostrum shows a distal setose
notch formed by a sub distal dorsal tooth; no other teeth on the rostrum. First
maxilliped with exopod abutting endite, displacing palp----------------Super family
PROCESSOIDEA------------------*Family Processidae.
Both first pereopods either simple or chelate. First maxilliped with
exopod far removed from endite--------------------------------------------------------------------------------------------28.
28.
First pair of pereopods
ALPHEOIDEA----------9.
distinctly
chelate-----------Super
family
First pair of pereopods with chela microscopically small or absent--Super family PANDALOIDEA--------------------------------------------------------------------------------------------------31.
29.
Eyes unusually elongate, reaching nearly to distal end of antennular
peduncle. First pair of pereopods about as robust as second pair ------------------------------*Family Ogyrididae.
Eyes normal in shape, short not reaching beyond the end of the first
segment of the antennular peduncle, sometimes covered by the carapace. First
pair of pereopods more robust than second pair--------------------------------------------------------------------------------------- 30.
30.
Carapace with cardiac notch in posterior margin. Eyes often partly or
entirely covered by the carapace. First pair of pereopods often unequal and
swollen----------------------------- *Family Alpheidae.
38
Carapace without cardiac notch (except in Saron and Yagerocaris).
Eyes free. First pair of pereopods usually equal, not swollen------------------------------------*Family Hippolytidae.
31.
Second pereopod with carpus undivided or with a single articulation.
First pereopod simple, without chela. First pleopod of male with endopod
enlarged, convoluted, and spinulose, petasma-like -------------------------------------------------------- Family Thalassocarididae.
Second pereopod with carpus usually composed of more than two
articles. First pereopod simple or microscopically chelate.First pleopod of male
with endopod laminar, not unusually large or convoluted----------------------------------------------------------*Family Pandalidae.
32.
Carpus of second pair of pereopods multi-articulate. First pereopod
prehensile, dactylus closing against the inner surface of propodus---------------Family Glyphocrangonidae.
Carpus of second pair of pereopods not subdivided. First pereopod subchelate,
dactylus closing against the sub truncate distal margin of the propodus which
often ends in a spine----------------------------------------------------------------------------------------------------*Family Crangonidae.
Species account
In the following account, the families are arranged as they
appear in the above key, while the genera and their species are in
alphabetic order. All the species are dealt with the current names and
supplemented by ecological informations including the larval
descriptions of species those were reared in either the author’s
laboratories or other Pakistani workers.
39
3.1.1 Family Pasiphaeidae Dana, 1852
Diagnosis: Rostrum either short or totally wanting, in some cases
represented by a spine arising behind frontal margin. Mandibular palp may be
present or absent, but molar process always present. Ultimate segment of
second maxilliped normally attached to penultimate segment (not as strip);
exopod rudimentary or absent, except in a few forms the chief part of
appendage. Exopods on third maxilliped and pereopods. First two pairs of
pereopods long and much stouter than others; their chelae elongate, fingers
slender; cutting edges of all the four movable fingers pectinate. Carpus of first
two pairs of pereopods short and unsegmented (From various sources).
Remarks: This is a family of pelagic shrimps, not all Pasiphaeidae
range so greatly throughout water column, and some heavier body genera
remain near the bottom at deep ocean depth. Pasiphaeids because of their
unusual mandibles and first and second maxillipeds disguise their affinities
about as well as any of the families (Chace, 1992).
The family Pasiphaeidae contains seven genera and 105 species
(Bauer, 2004) represented by the genera Eupasiphae Wood Mason, Pasiphaea
Savigny, and Leptochela Stimpson from the area under investigation. The
three Pakistani genera can be separated with the help of following key.
KEY TO THE PAKISTANI GENERA OF PASIPHAEIDAE
1. Mandible without a palp. Rostrum formed by an erect
postfrontal spine--------------------------------------------------------------------------------------------------Genus Pasiphaea.
- Mandible palp present. Rostrum a normal forward directed
prolongation of the carapace------------------------------------------------------------
40
------------------------------------2.
2. Fourth pereopod distinctly shorter than either third or fifth leg------------------------------------------------------------------------------------------------------------Genus Eupasiphae.
- Fourth pereopod longer than fifth leg, though sometimes
shorter than third ---------------------------------------------------------------------------------------------Genus Leptochela.
Genus Eupasiphae Wood Mason & Alcock, 1893
Diagnosis: Rostrum normal but short; post-orbital and post antennal
spines may be present. Mandible slender and provided with a 2-jointed palp.
Fourth pereopod shorter than the others.Telson apically narrow and truncated.
Two arthrobranchs on the third maxilliped. (After Holthuis, 1955).
Type species: Parapasiphae Gilesii Wood - Mason, 1892
Gender: Feminine
Remarks: Hanamura (1983) contradicts the possession of two
arthrobranchs on the third maxillipeds as the diagnostic character. He found
only one arthrobranch in his material of Eupasiphae.
There are four species included in the genus from the Indian
Ocean of which only E. gilesii has been recorded from Pakistan.
Eupasiphae gilesiim Wood-Mason, 1892)
(Figs.11-14)
Eupasiphae Gilesii Wood-Mason, 1892, Pl, 3, Fig, 8; Wood- Mason & Alcock,
1893:166
Parapasiphaea (Eupasiphaea) gilesii Alcock & Anderson, 1894:158; de Man,
1920; 3; Calman, 1939:187.
Parapasiphaea sp. B Chace, 1940: 140
41
Eupasiphae gilesi Fischer & Goldie, 1961:78;Tirmizi,1969:214; Foxton, 1970:
958; Crosnier & Forest, 1973: 150; Kensley et al, 1987:293; Kensley,
1981:23(distribution);
Hanamura, 1983:78; Burukovsky,1987:39; Crosnier,
1988: 786; Hanamura & Evans, 1994:52.
Eupasiphae gilesii Crosnier & Forest, 1973:150; Kensley, 1977:32
Description: The carapace is armed with two sharp spines; one
suborbital and one antennal spine. The antennal spine is slightly longer than the
suborbital one; both the spines are supported by well-developed carinae. The
carina of the antennal spine extends backward only for a short distance while
that of the sub-orbital spines runs backward, extending on the anterior one
fourth portion of the carapace. The position of the hepatic spine is marked by a
depression. The upper margin of the rostrum and the dorsum of the carapace
are armed with small teeth; each tooth is pointed and curved interiorly. The
rostrum is short, pointed, and slightly directed upwards.
The first and second abdominal somites are feebly carinated, while the
third is carinated only on its posterior two third portions; the fourth is sharply
carinated and it extends backward as a sharp spine which covers anterior one
fourth portion of the fifth somite; the fifth and sixth somites are also sharply
carinated, that of the sixth is produced into a small tooth. The telson tapers
towards the distal end; it is slightly longer than the sixth abdominal somite; on
the middle of its dorsal surface, a well-developed longitudinal groove is present.
The eye reaches the distal half of the basal segment of the antennular
peduncle. The cornea is globular. The main border of the ocular peduncle is
produced distally into a small protuberance.
42
Figure 11: Eupasiphae gilesii (Male). A- Lateral view of carapace; B-Lateral
view of third to fifth segments; C- Telson and left uropod; D- Right eye (
After Tirmizi,1969); E--Left scaphocerite in dorsal view;F Left, antennular
peduncle in dorsal view;
Figure 12: .Eupasiphae gilesii. A-Mandible of left side; B-Maxillule of right
side: C- Maxilla of left side. (After Tirmizi, 1969)
43
The antennular peduncle is slender. Its basal segment is about twice
the length of the second segment. The distal one is slightly longer than the
middle one; a well-developed, sharply pointed stylocerite is present which
extends nearly up to the distal margin of the basal segment. In the middle of the
stylocerite, a sharp longitudinal carina is present; a small spine is present on the
median side of the basal segment. The scaphocerite is well developed,
extending well beyond the antennular peduncle; the lateral spine of the
scaphocerite is very prominent and extends beyond the distal margin of the
squamose portion; the lateral margin of the scaphocerite is almost straight
except at the distal end where it curves inwards and forms the outer margin of
the spine. The mandible has a two-jointed palp; the basal segment is slightly
longer than the second segment; the incisor process bears nine prominent
teeth. The lower lobe of the inner lacinia of maxillule is subtriangular in shape;
the upper one slightly widens near the distal margin; the palp is unjointed; it
however, becomes rather narrow in its distal half. The maxilla has a rudimentary
inner lacinia; the palp is elongated and finger like. The first maxilliped‟s palp is
short. The second maxilliped is pediform. The third maxilliped is also pediform,
being long and narrow; its ultimate segment is shorter than the parapenultimate
segment; the exopod is well developed.
44
Figure 13: Eupasiphae gilesii. A- First right maxilliped; B-Second left
maxilliped; C-Third left maxilliped (After Tirmizi, 1969).
The first two pairs of pereopods are strong. The movable finger of the
first pereopod is much stronger than the immovable finger; the fingers are
shorter than palm; the movable finger is finely pectinate. The tips of the fingers
cross each other when closed; the carpus is short and more or less triangular;
its distolateral angle is produced into a spine; the median border of the merus is
armed with anteriorly curved spines, nineteen in all; it is three times as long as
the ischium which is unarmed. The exopod is well developed. The second pair
of pereopods is longer than the first; the fingers are finely pectinate; the carpus
is triangular and has a strong spine distally; a row of spines is present on the
merus and ischium. The exopod reaches almost to the middle of the merus. The
third pair of pereopods is very slender and flagelliform it reaches to the end of
the scaphocerite; the dactylus is long and narrow; the carpus is very short, while
the merus and dactylus are nearly of equal size. The fourth pereopod is shorter
than the third or fifth pereopod, its dactylus is flattened; the propodus is twice
the length of the dactylus; the segments bear setae on their posterior margins;
the carpus is rather small; the merus is the longest segment; the ischium has a
small spine. The fifth pereopod is slightly longer than the fourth; the ischium is
without any spine.
45
Figure14: Eupasiphae gilesii. A- Distal left part of first left pereopod; BDistal part of second right pereopod; C- Fourth left pereopod; D-Fifth left
pereopod.
The endopod of the first pleopod is short and broad, that of the second
pleopod bears both appendix interna and appendix masculina; the tip of the
appendix interna has few rows of coupling hooks, while the appendix masculina
bears long setae .On the third and fourth pleopods appendix internae are about
half of the length of the endopod.
Remarks: Considerable size related morphological variations both in
rostral shape and rostral dentition of the carapace has been observed in the
literature.
Size (male): 79mm in TL; 29mm in CL (including rostrum)
Locality: Northern Arabian Sea, 240 02N, 640 25E
Habitat: I0-1500m depth
Distribution: Indian, Atlantic and eastern Pacific Oceans; Andaman
Sea, off Cinque Island, Laxshadweep Sea off Goa coast, Central Arabian Sea,
Gulf of Oman, east coast of southern Africa, Bermuda, west of Cape Verde Isle,
near Madeira Canary Island.
Genus Leptochela Stimpson, 1860
Diagnosis: With conventional pasiphaeid rostrum. Carapace and
rostrum unarmed dorsally. Branchiostegal tooth and branchiostegal sinus
absent. Sixth abdominal somite with transverse carinate ridge near anterior end
of dorsal surface and long fixed posteriorly directed spine near posterior end of
ventrolateral margin. Telson with mesial pair of movable spines anteriorly, 1 or 2
pairs of dorsolateral movable spines, and 5 pairs of prominent posterior
movable spines, all but lateral pair (perhaps actually belonging to dorsolateral
series) of latter minutely on one or both lateral and mesial margins ( minute
additional pair of spines sometimes present between bases of median pair).
Mandibular palp broad, flattened and undivided. Third maxillipeds with two
arthrobranchs. Fourth pereopod shorter than third, longer than fifth. Exopods of
pleopods not unusually long. Both branches of uropod with series of movable
lateral spines. (Mainly from Chace, 1976)
Type species: Selected by Kemp, 1915:310. - Leptochela gracilis
Stimpson, 1860: 42
46
Gender: Feminine
Remarks: Chace (1976) considered to accord two subgenera
Leptochela and Probolaura in the genus, but not adapted by later workers and
hence not followed here. These little shrimps reaching in adult size up to 45 mm
total length are often abundant in certain parts of the world The chelate legs do
not appear in specimens measuring less than 2mm in CL. They are important
forage species of pelagic and benthic fishes. So far, 14 species of the genus are
recorded world wide grouped in two species group(Hayashi,1995). The material
at hand was collected from ten cruises of Dr.Fridtjof Nansen cruise in 1997.
From the leg of cruise closer to Pakistan four species are recognized, which can
be separated with the help of following key.
KEY TO THE SPECIES OF LEPTOCHELA IN THE PAKISTANI
WATERS OF THE ARABIAN SEA
1.
Telson armed with one pair of dorsomesial and two pairs of
dorsolateral spines in addition to posterior series; orbit armed with mesially
directed tooth on ventral margin or with tooth at suborbital angle-------------------------------------2.
Telson armed with one pair of dorsomesial and one pair of
dorsolateral spines in addition to posterior series. Ventral margin orbit and
suborbital angle unarmed (except in L.pugnax)-------------------------------------------------------------3.
2.
Pterygostomian
irrobusta.
spine
absent-----------------------------------------------L.
Pterygostomian
nasimae.
spine
present----------------------------------------------L.
3.
Suborbital
pugnax.
Suborbital
sydniensis.
angle
angle
dentate
----------------------------------------------------L.
unarmed--------------------------------------------L.
cf.
Leptochela irrobusta Chace, 1976
(Figs. 15-16)
Leptochela robusta Balss, 1915:17; Kemp, 1925: 252; Holthius, 1953:
52; Chace, 1955: 5
47
? Leptochela robusta Calman, 1939:188
Leptochela( Leptochela) irrobusta Chace, 1976: 19; Bruce, 1994:744;
Hayashi, 1995:93
Description: The rostrum has a straight dorsal margin and reaches
usually beyond the eyes, rarely extending slightly beyond level of distal margin
of basal segment of antennular peduncle. The carapace is simple, unarmed with
or without median or dorsolateral carinae or ridges in males and in non-breeding
females, but tricarinated in breeding females. The orbital margin is minutely
serrated; the suborbital angle has a mesially directed tooth or a tooth and two
spinules.
The abdomen is rounded dorsally on the four anterior somites, bluntly
carinated on posterior part of fourth and all of fifth; the sixth somite is provided
with two short slender spines on ventrolateral surface, and an acute tooth on
postero-dorsal lobe. The telson is armed with two pairs of dorsolateral spines,
posterior pair near mid length, in addition to an anterior dorsomesial pair; the
posterior margin bears pair of minute fused spines or knobs between bases of
mesial pair of 5 pairs of prominent spines.
Figure15: Leptochela irrobusta. A- Carapace; A’- Rostrum and orbit, lateral
view, magnified; A”- Rostrum and orbital margin, dorsal view; B-Posterior
end of sixth abdominal somite; C- Telson and uropods; C’- Posterior
margin of telson.
48
The stylocerite of antennular peduncle reaches about as far as the
distolateral margin of basal segment. The stylocerite has 2-4 simple setae on its
inner margin.
The antennal scale is half as long as carapace, its lateral margin is
slightly concave; the distal tooth is continuous with the mesial margin of scale;
the distal segment of antennal peduncle is nearly two third as wide as scale and
does not reach mid length of scale.
Figure 16: Leptochela irrobusta. A- First pereopod, magnified; A’- Same,
chela; B- Antennule, dorsal aspect; C- Antenna, ventral aspect.
The first pereopod overreaches the antennal scale by the length of
fingers; the latter are longer than the palm; the dactylus is armed with 23 spines
on opposable margin. The second pereopod overreaches the antennal scale by
less than length of its fingers; the fingers are longer than the palm. The third
pereopod overreaches anterior margin of carapace by combined length of the
dactylus and propodus; its exopod does not reach distal end of the ischium,
bearing row of three long spines near outer margin and a sub distal spine on
49
inner margin; its merus is armed with five or more similar spines near outer
margin; the dactylus is distinctly shorter than the propodus. The fifth pereopod is
similar to and little shorter than the fourth reaching to about the mid length of
ischium of third pereopod when both are extended anteriorly, the dactylus is
longer than the propodus.
The endopod of first pleopod is rather narrow, round distally; the
appendix masculina bears two long spines.
Size: 35 mm in CL
Habitat: Near shore station,Fridtjof Nansen Cruise
Distribution: Persian Gulf, Laxshadweep, Maldives, Andaman, Timor,
Nicobar, Myanmar, Vietnam, Philippine Islands, and Australia. This is first
record of the species from the northern Arabian Sea.
Leptochela nasimae sp. nov.
(Figs. 17-18)
Material Examined: Fridjof Nansen Cruise, St, 12 (Hoz), 2 specimens,
1 male CL 3.5mm (holotype) St.5 (Hoz), 2 specimens (1 immature male), St 26
(Hoz),7 specimens; St.27 (Hoz), 22 specimens; St 38 (Obl),3 specimens;
St.39(Hoz), 1 specimen; St 67 (Hoz), 1 specimen (carapace only)CL 2 to 4mm
(paratypes) (fig.10).
Description of Holotype: The dorsal margin of rostrum is straight
reaching up to eye. The median carina of carapace extends on anterior half of
its length. The orbital margin is serrate, with suborbital angle and
pterygostomian angle produced into a triangular small process.
The abdomen is regularly rounded dorsally but the ventral margins of
first two somites are serrated and those of last three are spined. The fifth somite
is carinated, without dorsal prominences but with sharp median tooth projecting
from the posterior margin. The sixth somite is slightly produced posteriorly;two
small spines on vetral surface; posterior lobe with pronounced acute tooth on
posteroventral margin. The telson is a long as the sixth somite, armed with two
pairs of dorsolateral spines in addition to anterior mesial pair; the posterior
margin has a fused pair of minute spines between bases of mesial pair of usual
five pairs of prominent spines.
The stylocerite nearly reaches a far as distolateral margin of the basal
segment; the second segment is shorter than the distal segment.
50
Figure 17: Leptochela nasimae Male Holotype. A- Carapace; A’- Orbital
region, lateral aspect; A’’-Rostrum and orbital margin, dorsal aspect; BPosterior margin of fifth abdominal somite and sixth abdominal somite; CTelson and uropods; C’ Posterior margin of telson.
The antennal scale has a convex lateral margin, which ends in an
apical concavity on the inner margin. The distal segment of antennal peduncle is
slightly more than half as wide, not reaching to middle of scale; the basal
segment has a distinct ventral spine. The first pereopod overreaches the
antennal scale; the fingers are longer than the palm; the dactylus is armed with
24 spines on the opposable margin. The second pereopod overreaches the
antennal scale by no more than length of fingers; the fingers are as long as
palm; the dactylus is armed with 21 spines on opposable margin.
51
Figure18:Leptochela nasimae. Male Holotype. A- Abdomen, dorsolateral
aspect; B-Antennule, dorsal aspect; C-Antenna, ventral aspect; D-First
pereopod; D’- Same, fingers; E-Second pereopod; E’-Same, fingers; FEndopod of first pleopod; G- Endopod of second pleopod showing
appendix masculina and appendix interna.
The endopod of first pleopod of male is rounded distally, the lateral
margin is slightly concave, and the appendix masculina bears only one spine.
The exopod of uropod is armed with nine movable spines.
Remarks: The specimens at hand have very peculiar combination of
characters. The spine on the posterodorsal end of the fifth abdominal somite is
not a common character in Leptochela, present only in L. gracilis Stimpson. The
other characteristic feature is the presence of a spine at pterygostomian region,
which is found in none of the species described by Chace (1976); the serrated
ventral margins of abdominal somites also do not match with any of the
described species. The serrated abdominal somites might have some relation
with the state of maturity of specimens. The characters numerated above may
justify the separation of the Fridtjof Nansen material under a new name.
52
Etymology: The species is named in the honour of late Dr. Nasima
M. Tirmizi, Ph.D. Supervisor of both the authors.
Leptochela pugnax De Man, 1916
(Figs. 19-20)
Leptochela pugnax de Man, 1916: 148; 1920:26 pl. 4: Fig. 8;Kemp,
1925: 255;Armstrong.1941:3; Kubo, 1955:101, fig. 4, 5; Holthuis & Gottlieb,
1958:22; Bruce, 1990: 612; Kazmi et al, 1990:235; Hayashi, 1995:95
Leptochela sp. de Man, 1920:30.
Leptochela robusta , Miyadi, 1940:141 [Not L. robusta Stimpson, 1960].
Leptochela (Leptochela) pugnax Chace, 1976:31; Fig. 25.
Description: The rostrum is distinctly upturned at the tip; the dorsal
margin is concave, provided with a dorsomedial carina. The orbital margin is
entire, not serrate, the ventral portion is without the mesially directed tooth; the
suborbital angle is armed with a prominent denticle.
The abdomen is regularly rounded dorsally on three anterior somites
and conspicuously carinated on fifth. The fifth somite is without the dorsal
prominences and without poster dorsal tooth; the spine on ventrolateral surface
of the sixth somite is distinct and long; the posterolateral lobe bears a distinct
acute tooth on the posterodorsal margin. The telson not including posterior
spines is about 1.3 times as long as sixth somite and is armed with 1 pair of
dorsolateral spines slightly anterior to midlenght, in addition to anterior mesial
pair; the posterior margin may or may not posses a pair of minute spines
between the bases of mesial pair of usual 5 pairs of prominent spines.
53
Figure 19: Leptochela pugnax. Entire animal, lateral view
The antennular peduncle is with stylocerite reaching nearly as far as
distolateral margin of basal segment, the second segment is distinctly shorter in
dorsal aspect.
The antennal scale has a concave lateral margin; the distal tooth is
continuous with mesial margin of blade. The distal segment of antennal
peduncle is narrower than the scale, reaching less than mid length of scale; the
basal segment is without the ventral tooth.
The first pereopod does not overreach the antennal scale; the fingers
are 1.5 to 2.0 times as long as palm; the dactylus is armed with seven spines on
opposable margin. The second pereopod is overreaching antennal scale by less
than the length of fingers; fingers are 1.9 to 2.6 times as long as the palm; the
dactylus is armed with 11 spines on opposable margin.
The lateral margin of exopod of uropods is armed with seven spines, in
addition to setae.
Remarks:The Arabian Sea specimens posses fewer spines on dactylii
of pereopods as compared to their Indo-Pacific counterparts(Chace,1976).
Dimorphic forms of females occur in this species (Kemp, 1925).
54
Figure 20: Leptochela pugnax A- Carapace; A’- Orbital region, lateral
aspect; B- Posterior end of sixth abdominal somite;C- Telson and
uropods; C’-Posterior margin of telson; D- Antennule, dorsal aspect; EAntenna, ventral aspect; F- First pereopod ; F’- Same, fingers; G- Second
pereopod ; G’- Same, fingers.
Size:
2.0 mm to 2.7 mm in CL
Habitat: In inshore water ranging in depth from seven to 140 meters on
both sandy and muddy sediment (Chace, 1976, Bruce, 1990). Locality:Near
shore stations of Fridtjof Nansen Criuse ,Port Qasim
Distribution:Red Sea, The Maldives Islands, eastward to eastern
Indonesia, Java to Moluccas, Hong Kong and northward to the south of Honshu,
Japan, penetrates in to the Mediterranean.
Leptochela cf sydniensis Dakin & Colefax, 1940
(Figs. 21-22)
55
Leptochela oculeocaudata Kemp, 1915:311(part); 1925:254; Menon,
1937:6; Kubo,1955:103; Pillai,1955:48;Fujino & Miyake 1970:239(not L.
oculeocaudata Paulson,1875)
Leptochela hainanensis Yu, 1936:87
Leptochela sydniensis Dakin & Colefax, 1940:153: Chace, 1976:40;
Bruce, 1986:612; 1988:277, 1994: Kazmi etal, 1990:235; Hayashi, 1995:98.
Description: The rostrum has a sinuous dorsal margin and reaches
up to the eye. The carapace has a median dorsal carina on anterior 1/3 or less
of length in males. The anterior lobe of carapace conceals the basal segment of
antennal peduncle from lateral view. The orbital margin is spinulose, without
distinct mesially directed tooth on ventral portion; the suborbital angle is
rounded and unarmed.
The abdomen is rounded dorsally on first 3 somites, carinated from
fourth to fifth, partially carinated on the fourth, distinctly carinated on the fifth; the
fifth somite is entire without any dorsal prominences or posterodorsal tooth, the
sixth somite is nearly one and a half as long as high, with a transverse
carinated swelling at the anterior end; the usual two smaller and along and
curved spine on the ventrolateral surface and distinct acute tooth is present on
the posterodorsal margin of posterolateral lobe.
Figure 21: Leptochela cf sydniensis. A- Carapace; B- Posterior margin of
sixth abdominal somite; C- Telson with uropods; D- Antennule; EAntenna; F- Second pereopod
56
The telson is nearly one ½ times, as long as sixth somite armed with
a pair of dorsolateral spines beyond mid length in addition to anterior mesial
pair; the posterior margin is without a pair of minute spines between the bases
of mesial pair of usual 5 pairs of prominent spines.
The stylocerite of antennular peduncle reaches beyond distolateral
margin of basal segment; the second segment is smaller than the distal
segment in both mesial and dorsal aspects. The antennal scale is narrow; the
lateral margin is distinctly concave near mid length; the junction of mesial
margin of blade with distal tooth is marked with a distinct shoulder. The distal
segment of antennal peduncle extends to about 1/3 the basal length of scale;
the basal segment is produced distoventrally as an acute tooth.
Figure 22: Leptochela cf sydniensis. A- First pereopod ; A’- Same, chela;
B- Second pereopod; B’- Same, chela
The first pereopod overreaches the antennal scale by as much as
length of the fingers; the dactylus is armed with 20-44 spines on opposable
margin. The second pereopod overreaches the antennal scale by more than
length of the fingers; the dactylus is armed with 17 spines on opposable margin.
The third pereopod overreaches extreme anterior margin of the carapace by
about the length of dactylus; the exopod is half of the ischium; the ischium is
57
armed with three spines; the merus has four spines; the carpus is provided
with three spines; the dactylus is distinctly shorter than the propodus. The fourth
pereopod reaches to half of the merus of third pereopod when both extend
anteriorly; the dactylus is distinctly longer than the propodus. The fifth pereopod
is similar but shorter in length of fourth; the dactytlus is about as long as
propodus.
Remarks: While using the key given by Chace, (1976) our specimens
are identified L. sydniensis Dakin & Colefax on the basis of (a) having an
unarmed ventral margin of orbit and suborbital angle (b) having telson armed
with one pair of dorsomesial and one pair of dorsolateral spines, (c) telson
lacking pair of minute spines between bases of median posterior pair and (d)
unarmed, fifth abdominal somite, But on detailed examination a relatively
shorter rostrum, an extended anterior lobe of carapace, longer tooth on poster
dorsal margin of posterodorsal lobe of telson, longer stylocerite, narrow
scaphocerite and smaller number of spines on second pereopod are the
characters enough to distinguish the specimens form those of L.sydniensis
given by Chace (1976:40) and L. chacei by Hayashi (1995). It seems
appropriate to keep it tentatively near to L.sydniensis Dakin & Colefax. This and
its allied species L. robusta, L .aculeocaudata, and L.pugnax make a complex
and are discussed at length by (Chace, 1976:44). He merged L. aculeocaudata
of the Indian authors like Kemp, 1925, Menon,1937 and Pillai,1955 with
L.sydniensis but was not sure about his decision, he opined that it was not “the
final taxonomic word on rather complex problem”, but „to extent towards such a
solution” The present material may prove to be part of this complex .
Size: 4 to 5mm in CL (females).
Habitat:
Planktonic
Distribution: Arabian Sea, Bay of Bengal, the South China, and Yellow
Seas, Japanese waters and Australia.
Genus Pasiphaea Savigny, 1816.
Diagnosis: Rostrum represented by a post-frontal gastric spine. Orbits
poorly defined. Fourth pereopod shorter than fifth. Mandible without a palp.
(After Chace, 1940)
Type species: Parapasipheae alcocki Wood-Mason & Alcock, 1891 by
58
monotypy.
Gender: Feminine
Remarks:This is a large genus which contains more than 60 species
arranged in 4 species groups(Hayashi,2004) At least 17 species of the genus
Pasiphaea are recorded from the Indian Ocean (Burukovsky, 1993). The
species are pelagic and vertical migrants.Hayashi (2006) has revised recently
the P.alcocki species group. One species P. alcockii Wood Mason, 1891, has
been recorded off Indus fan area and P. sivado from the section near coast. .
Pasiphaea alcocki( Wood Mason &Alcock, 1891)
Pasiphaea alcocki Wood
Mason&Alcock,1891:190;deMan,1920:6;Chace,1940:123;George&Rao,1966:23
8;Hayashi,2006:------Description:The carapace is not carinate dorsally.The abdomen is without
dorsal carination or armature
on any somite. The telson is sub-acute on the dorsal surface and not conspicuously
forked.The first
pereopod is armed with 4-6 spines on the merus and 1 spine on the carpus.The second
pereopod with
13-16 spines on the merus and 1 spine on the carpus. The fingers of the first chelae are
slightly more than half the length of the palm, while the fingers of the second chelae are
almost of the same length as the palm.
Size:63mm
Habitat:Demersal
Locality:Section near coast of Pakistan
Distribution: Bay of Bengal, Gulf of Manaar, and the Arabian Sea.
Pasiphaea sivado (Risso, 1816)
(Fig.23)
Alpheus sivado Risso,1816:93;Williams et al,1989: ;Hayashi,1999:523
Description:Five pleurobranchia are present,the last is small and rudimentary.
59
Figure 23: Pasiphaea sivado. Animal in lateral view (After Holthuis, 1993).
Remarks:P.sivado species group contains 9 species(Hayashi,1999)
Locality:Off the Sindh coast(947 fathoms)
Distribution: P. sivado is distributed in the Mediterranean, Atlantic, and
Indian Oceans
60
3.1.2 Family Oplophoridae Dana, 1852
Diagnosis: Rostrum present, immovable, highly variable in form;
mandible imperfectly cleft, palp segmented; third maxilliped and all pereopods
with exopods; first two pairs of pereopods chelate; carpii entire; pleopods with
appendices interna (Taken from Chace, 1986).
Remarks: The major characteristics of oplophorids are more a
combination of generalized, primitive characters than specialized features. The
unique thoracic epipods isolate the family from the other families. Adaptations to
pelagic life dominate the morphology and biology of the shrimps (Chace, 1940;
Omori, 1974).Species are capable of some form of bioluminescence (Herring,
1976). So little is known about the basic biology of these shrimps, which occupy
a world habitat of enormous extent. As such, this group is badly in need of
detailed study ,until now, 70 species have been recognized in the ten genera
(Bauer, 2004). From offshore waters of Pakistan, a single genus Acanthephyra
is recorded.
Genus Acanthephyra A. Milne-Edwards, 1881
Diagnosis: Rostrum with at least as many dorsal as ventral teeth;
carapace not denticulate dorsally usually without uninterrupted lateral carina
extending from near orbit to near posterior margin, without hepatic spine,
posterior slope of hepatic furrow usually not abruptly delimited by oblique carina;
abdomen dorsally carinate on at least third through sixth somite; telson
superficially blunt or sub truncate posteriorly not tapering regularly to sharply
acute posterior end, without spinose end piece; antennal scale without lateral
teeth proximal to distolateral spine; mandibles dissimilar; molar process with
transverse distal surface triangular on right member, compressed, sub-bilinear
on left, incisor process toothed along entire opposable margin; second maxilla
with proximal endite bearing papilla and sub marginal lamina; first maxilliped
with slender central lobe subdivided by 2 transverse sutures; second maxilliped
with distal segment sub triangular attached diagonally to preceding joint, third
maxilliped and first pereopod with exopod not unusually broad or rigid, legs with
61
ischium and merus not broadly compressed, fourth pair with epipod vestigial
or absent; appendix masculina present on second pleopods of male( from
Chace, 1986).
Type Species: Acanthephyra armata , A. Milne Edwards, 1881.
Gender: Feminine
Remarks: Chace (1986) considered genus Acanthephyra “a
polyphyletic one” encompassing 27 species, one species from southern Arabian
Sea (A. sanguinea) by George & Rao (1966) and one species Acanthephyra
eximia has been collected from the northern Arabian Sea(Pakistani waters).
Acanthephyra eximia Smith, 1884
(Figs. 24-26, Pl. 2B)
Acanthephyra Brachytelsonis Bate, 1888: 753
Acanthephyra eximia Smith, 1884: 377 (eximea: 376); Alcock, 1901:-;Calman, 1939:191 Siversten& Holthius, 1956:3; Holthuis & Gottlieb, 1958:112;
Kazmi, 1971a:169; ; Crosnier & Forest, 1973:34 (detailed synonymy);
Hanamura, 1983:73 Chace, 1986:186; Crosnier, 1987:699; Burukovsky,
1990:194;Pohla,
1992:1;
Hanamura
&
Evans,
1994:47;
Poupin,
1996:4;Cardoso&Young,2005:14
Description: The carapace is laterally compressed, leathery and
dorsally carinated and sinuous throughout its entire length, with a marked
depression in the region of the cervical groove. The antennal and
branchiostegal spines are well developed and sharply pointed. A suprabranchial
ridge is present. The rostrum is rather long and compressed not extending
beyond the scaphocerite; it is armed with eight teeth on the upper margin,
confined to the proximal region and four teeth on the lower margin, immediately
below to the unarmed dorsal portion.
62
Figure 24: Acanthephyra eximia. A-Female, in lateral view; B- Male, lateral
view (After Wood Mason, 1892).
The first abdominal somite is without a dorso-median carina, second is
feebly carinated, whereas, the rest of the somites are strongly carinated. The
carina of the third somite is strong and extends backwards as a sharp spine,
which reaches almost to the middle of the fourth somite. The dorso-median
carinae of fourth and fifth somites end in small spines that of the sixth are
produced in a large spine. Telson tapers gradually and ends in a strong spine,
there is a faint dorsal ridge, further, dorsal surface is armed with four pairs of
spines on its distal half, these spines gradually become larger towards the distal
end, and the tip of the telson is flanked by three spines: two on the right and one
on the left side.
The cornea is almost as wide as the stalk. The antennular peduncle is
short. Its basal segment is longer than the combined length of the distal two
segments; second and third segments are nearly of equal size; the anterior
margin of the basal segment is deeply notched on its upper surface, a well
developed stylocerite tapers gradually into a sharp and extend as far as the
distal end of the basal segment. The scaphocerite is an elongated leaf-like
structure; it is broad at the base, becoming narrower towards the distal end. The
outer margin is straight and ends in a small but distinct spine. The inner margin
is furnished with long plumose setae.
63
Figure 25:
Acanthephyra eximia. A- Lateral view of carapace; BAbdomen; C-Dorsal view of telson and left uropod; C’- Tip of telson; DRight eye; E- Antennule; F-Scaphocerite.
The mandible has a three-segmented palp; the basal segment is very
small, whereas, the median segment is very long; its inner margin becomes
convex near the distal end, giving its characteristic appearance; the incisor
process of the mandible bears nine sharp, unequal teeth. A distinct process is
also present. The maxillae are missing from the specimen. The first maxilliped
possesses a long slender three-segmented palp. The second maxilliped is
subpediform. The third maxilliped is elongated and pediform.
64
Figure 26: Acanthephyra eximia. A-Mandible of left side; B-First maxilliped
of left side; C-Second maxilliped of left side; D-Third maxilliped of left
side; E-Chela of first pereopod of right side; F- Appendix interna of
second pleopod of right side
The first pair of pereopods is small, the chelae are robust, the inner
margin of the dactylus is serrated, and the palm bears tufts of setae. The palm
is twice the length of the fingers. The chela is about twice as long as carpus.
The second pair of pereopods is longer than the first; its chelae are slender and
are about the same length as the carpus. The last two pairs of pereopods are
long and very slender. The fourth pereopod differs from the fifth in being slightly
stouter, shorter in length and bearing a row of spines on ischium and merus,
spines are absent on ischium of last pair of pereopods.
The endopod of each of the last four pairs of pleopods bears an
appendix interna, which is about one-third the length of the endopod. The tip of
the appendix interna is armed with a few rows of coupling hooks or cincinnuli.
Appendix interna and masculina of male are illustrated by Chace (1986, Fig.
9a.).
In the female, the thoracic sternites are armed. The first thoracic
sternum has a pair of small tooth-like projection; each is pointed upwards and
slightly curved forwards, so that in the posterior view it is somewhat convex. The
second, third and fourth sternites are more or less similar in structure, whereas
the fifth thoracic sternum has a single median plate which is notched anteriorly.
65
In between the two tooth-like projections, a strong longitudinal ridge runs
from the notch to the anterior border of the segment. The abdominal sternites
are unarmed except the last one where a small posteriorly directed spine is
present.
Colour: Crimson (When fresh) (Alcock, 1901).
Size:
95mm, in TL (Chace, 1986).
Habitat: It is a deep sea species can be found up to 1050m depth
(Poupin, 1996). The Arabian Sea material (one female) was collected at 1500m
depth (24002 N 64025E) by the PNS Ship “Zulfaquar.” So far, few males and
females are collected from the Northern Arabian Sea, it seems that the species
is not common in our seas.
Distribution: Western Atlantic off the coasts of North and South
America, Canadian Atlantic, Mediterranean, Red Sea and in the Indian and
Pacific Oceans from off Cape Natal, Africa, to the Peninsular Malaysia, French
Polynesia, Western Australia, Japan and the Hawaiian and Society Islands.
66
3.1.3 Family Atyidae de Haan, 1849
Diagnosis: Rostrum, if present, inflexibly attached to rest of carapace.
Carapace without longitudinal lateral ridges or suture and without cardiac notch
in posterior margin. Eyes neither usually long nor concealed beneath carapace.
Antennules with two flagella, neither with accessory branch. Mandible with palp,
with sub truncate molar process not distinctly separated from incisor process.
Second maxilla with endite well developed, scaphognathite with proximal lobe
tapering, bearing series of long setae, and extending far into branchial chamber.
First maxilliped with exopod terminating in lash, not in broad, partially detached
lobe. Caridean lobe not actually produced, not overreaching distally produced
endite. Second maxilliped with exopod, endopod composed of four segments,
not terminating in two segments attached side by side to preceding segment,
terminal segment attached to slender, sickle-shaped extension of preceding
segment. Third maxilliped composed of five segments, slender, pereopod-like.
Pereopods usually with strap-like epipods (mastigobranchs) on at least 3
anterior pairs, epipods without naked appendix extending vertically into
branchial chamber; 2 anterior pairs of pereopods similar, with fingers of chelae
usually terminating in tuft of setae; second pereopod with carpus undivided (
from Chace,1997).
Remarks: The Atyidae as whole is regarded a very primitive family of
the Caridea in spite of the fact that peculiarly modified chelae indicate a
considerable degree of specialization. Most of them live in fresh water lakes and
streams, some times in the backwaters (Thomas et al, 1973).
Rather surprisingly, only three of the 35 genera currently recognized
throughout the world are apparently known from the area. They are Caridina ,
Caridella, and Kalriana. The status of Caridella from Pakistan is uncertain since
it has not been encountered after its first preliminary record by Baqai et
al(1974).
Since (late) Prof. L. B. Holthuis (Q.B.K, pers. comm.) found no
indication of a type species for the new genus Kalriana Zuberi, he opined that
the name Kalriana Zuberi,1990 is unavailable nomenclaturally. His first
impression is that all species of Kalriana are Caridina , but that should be
67
verified. As we were unable to produce an effective response to his
arguments and did not want to delay this publication long enough to await a
verdict from the ICZN, it seems best for the purpose of this publication to include
it as it is.
KEY TO THE PAKISTANI GENERA OF ATYIDAE
1.
Carpus of second pereopod not deeply excavate----------------------------------------------------------------------------------------------Genus Caridina
_
Carpus and ischium of both first and second pereopod
excavate----------------------------------------------------------------------------------------------------------------------Genus Kalriana.
Carpus of first pereopod anteriorly excavate, that of second leg
not excavate--------------------------------------------------------------------------------------Genus Caridella.
68
Genus Caridina H.Milne Edwards, 1837
Diagnosis: Carapace without supraorbital spine, pterygostomian
margin usually rounded. Telson with posterolateral angles sometimes produced
posteriorly but never overreaching setiferous posterior margin. Eyes usually well
pigmented, not degenerate. Pereopods without exopods, second pair with
carpus not deeply excavate, distinctly longer than wide ( from Chace, 1997)
Type species: By monotype: Caridina typus H.Milne Edwards, 1837
Gender: Feminine
Remarks: The genus occurs in great number in every suitable locality
inhabiting both fresh and brackish water and ascending to at least 6000ft. Few
caridean, groups offer taxonomic problems of greater difficulty than do the
approximately 160 species and subspecies that are currently recognized in this
genus. Few populations can yet be named with satisfactory confidence that
range widely through the Indo-Pacific region and even Africa (Chace, 1997). As
already stated that the taxonomy of this genus is very difficult, and with the large
number of described species, elucidating the identities of the present Pakistani
specimens are not possible, as such the identity C. babaulti must be regarded
as tentative, to keep certain reservation on its denomination. The status of C.
babaulti is still a controversy in the world. The Pakistani material of Caridina
belongs to three species, separable from the following key.
KEY TO THE PAKISTANI SPECIES OF CARIDINA
1.
Rostrum unarmed anteriorly. First pereopod deeply excavated
for reception of proximal portion of chela----------C.weberi.
Rostrum with 1-3 sub apical teeth separated from rest of dorsal
series. First pereopod not deeply excavated--------C. nilotica
Rostrum unarmed anteriorly. First pereopod with a shallow
excavation for reception of chela-------------------C.cf babaulti babaulti.
Caridina cf babaulti babaulti (Bouvier, 1918)
(Fig. 27)
Caridina babaulti Bouvier, 1918:388
69
Caridina babaulti, Johnson, 1961: 136;, Jalihal et al, 1984:19 ;
Gorgin, 1996:666
Caridina babaulti babaulti Al-Adhub & Hamzah, 1987: 228
Description: The rostrum is short and slender, rather straight, with a
slight upward tilt at the distal end, it extends to the end of the second segment
of the antennular peduncle or slightly less far, rarely a little beyond the segment.
The tip of the rostrum terminates in a sharp point. Its dorsal armature consists of
16-27 closely packed teeth interspersed with prominent hairs (bristles). Six to
eight of the teeth are placed on the carapace behind the posterior limit of the
orbit. There are 4-11 (usually 6 or 7) ventral rostral teeth, situated in the distal
half of the rostrum. The terminal portion of the rostrum is unarmed, both dorsally
and ventrally. The carapace shows a strongly developed antennal spine,
situated on the lower angle of the orbit.
Figure 27: Caridina babaulti babaulti. A- Cephalic region in lateral view; BFirst leg; C- Second leg
The telson has a convex posterior edge with short, acutely pointed
median spine. The posterior margin bears three pairs of spines; the outer most
are the longest. There are also four pairs of thin intermediate hairs. The telson
has also 4-6(usually 5) pairs of dorsal spines. The exopod of the uropod bears a
series of 18 – 20 spinules along the diaeresis.
The antennule has a triangular, sharply pointed stylocerite extending to
about ¾ the length of the first peduncular segment. There is no distinct
antennular carina. The spine of the antennal scale extends well beyond the tip
of the antennular peduncle.
70
The chela of the first cheliped is more than twice as long as broad.
The carpus is deeply excavated anteriorly and attached to the chela ventrally;
the chela projects backward beyond the point of attachment. The carpus is
shorter than the chela and about one ½ times as long as broad. The chela of the
second cheliped is about twice as long as broad. The carpus is about 5-6 times
as long as broad.
The dactylus of the third pereopod is about ¼ the length of the
propodus and bears a ventral fringe of seven spines; the propodus is 8 times as
long as broad; the merus has four articulated spines whereas the ischium has a
single articulated spine or none. The dactylus of the fourth pereopod is about
1/5 the propodus length and bears a ventral fringe of 8-9 spines; the propodus
is about 10 times as long as broad; the merus has 2 articulated spines while the
ischium is lacking spines. The datylus of the fifth pereopod is about ¼ the
propodus length and bears a ventral fringe of about 37 spinules; the propodus is
about 10 times as long as broad; the merus has two articulated spines while the
ischium is without spines.
The endopod of the first pleopod of the male is less than half as long as
the exopod and narrows distally; proximally, it is leaf –like in shape with a
prominent inner sub terminal process which is about half as long as the
endopod.
Remarks: As already stated that the taxonomy of this genus is very
difficult, and with the large number of described species, elucidating the
identities of the present Pakistani specimens are not possible, as such the
identity C. babaulti must be regarded as tentative, to keep certain reservation on
its denomination. The status of C. babaulti is still a controversy in the world.
Size: Females are larger than males.
Habitat:Fresh water
Distribution: Central India, Iran and Iraq and for the first time from
Pakistan.
Caridina nilotica (P.Roux, 1833)
(Fig. 28)
71
Pelias nilotica P.Roux, 1833:73
Caridina nilotica - Barnard, 1950: 657; Holthuis, 1965:15; Mehr etal,
1988: 1291; Chace, 1997:17; Hussein & Obuid-Allah, 1992:121; Richard &
Clark, 2005:706
Caridina nilotica var brevidactyla J. Roux, 1919: 320
Description: The rostrum is very long, slender, and curved upward at
the tip. It usually reaches distinctly beyond the antennal scale and has the tip bi
or trifid. There is a distal unarmed portion on the dorsal margin behind the single
or double sub apical tooth. The proximal part of the upper border of the rostrum
bears 13-27 teeth 2 or 3 of which are placed behind the orbit. All teeth are
placed quite close together. The lower margin bears 6-22 teeth. The upper teeth
articulate with the rostrum, the lower do not. The lower orbital angle is very
distinct and separated from the antennal spine by a good distance. The
pterygostomian angle is rounded.
72
Figure 28: Caridina nilotica With individual variation. A,B- Anterior part
in lateral view; C- Tip of telson; D-E- First leg; F-G- Second leg; H-I- Third
leg; J-K- Fifth leg; L- Endopod of first pleopod of male; M- Outer part of
diaeresis (Modified from Holthius, 1965).
The dorsal surface of the telson bears 2-5, usually three or four pairs of
spinules. The posterior margin of the telson is rounded and shows no median
tooth; the outer pair of posterior spines is very short, being less than ¼ of the
length of the well-developed lateral pair. There are 4-8 intermediate spines,
which are undivided and distinctly shorter than the laterals.
The stylocerite is slender and pointed; it reaches beyond the middle of
the basal segment of the antennular peduncle but distinctly fails to reach the
end of the segment. The anterolateral tooth of this segment is sharply pointed; it
fails to reach the middle of the second segment. The spine near the base of the
scaphocerite is well developed and sharp.
The carpus of the first leg is longer than the merus; it is usually more
sometimes somewhat less, than twice as long as high. It is shorter than the
chela. The carpus of the second leg is longer than the chela; it is 3-6 times as
long as high. In the third leg, the dactylus bears 5-6 posterior teeth additional to
the distal claw; the propodus is 4-4.5 times as long as the dactylus. The
propodus of the fifth legs is 3.5 – 4.5 times as long as the dactylus.
The endopod of the first pleopod of the male is provided with a welldeveloped appendix interna. The exopod of the uropod bears 7-13 spinules on
the diaeresis. The preanal carina is blunt without a spine.
Remarks: C. nilotica in a protean species; Chace (1997), has merged
all varieties of C. nilotica with the main species. It is supposed to be restricted
to River Nile catchment area, material reported outside this area probably
belongs to other species(Richard & Clark,2005). This statement makes identity
of Pakistani material by Mehr et al.,1988 doubtful.
Colour: Semi-transparent with faint pinkish or orange brown speckling
(Barnard, 1950)
Size: 5mm in CL
Localities: Wah
Distribution: Over an extensive range from Eastern Africa to
Polynesia.
73
Caridina sumatrensis de Man, 1892
(Fig. 29, Pl. 2 B)
Caridina weberi var. sumatrensis de Man ,1892:375, pl.xxii, fig.23;
Bouvier, 1925:247, fig.571; Chopra & Tiwari ,1949:215; Holthuis & Rosa,
1965:9; Johnson, 1966:420; Jalihal et al, 1984:20; Hung et al, 1993:481:;
Richard & Chandran, 1994:257; Cai & Ng, 2000:942; Kazmi et al, 2002:43.
Caridina weberi
Chace, 1997:22; Cai & Ng, 2001:666
Caridina weberi sumatrensis Cai & Shokita, 2006:246
Description: The rostrum does not reach as far as distant end of the
antennular peduncle, the dorsal margin is horizontal and is turned upward at
the tip armed with up to 20 teeth does not reach extreme apex but extends on
dorsal margin of carapace, armed ventrally with 4-5 teeth. The suborbital angle
is indistinguishably fused with the antennal spines. The pterygostomian margin
is armed with a spinule; that of the right side is missing in one female. The eyes
are well developed.
The dorsal surface of telson is armed with three pairs of spines; the tip
of telson is convex, armed with 10 spines but without a median denticle.
The antennular peduncle is not slender, the stylocerite reaches up to
two third of the basal segment of antennular peduncle. The basal segment is
more than half-length of peduncle, anterolateral angle is distinct. The
scaphocerite is narrow, without distinct transverse suture. The mandible is
without palp, eleven small and large teeth are present at extremity of incisor
process.
74
Figure 29: Caridina sumatrensis. A. Front lateral view; B. Telson; B'.
Posterior margin of telson, magnified; C. Antennule; D. Antenna; E.
Mandible; E'. Same, cutting edge, enlarged; F. First pereiopod; F'-F"".
Same, setae on anterior margins of fingers; G. Second pereiopod; H.
Distal end of third pereiopod; H'.
Same, enlarged; I. Distal end of fifth pereiopod; I'. Same, enlarged; J. First
normal pleopod, J'. First abnormal pleopod; K. Second pleopod; K'. Same,
appendix interna enlarged.
75
The first pereopod is shorter and stouter than the second is;
the fingers are longer than the palm. Different kinds of setae are present apically
on first pereopod: the carpus variably excavates for the reception of chela, the
dactylii of third pereopods terminate in double spines, and there are about five
accessory spines on inner margin increasing in length distally. The dactylus of
last pereopod is as figured.
The first pleopod is as figured; in one female endopod equals to
exopod, the second pleopod is as illustrated.
Remarks: The specimens from Karachi were reported by Kazmi et al
(2002) as Caridina weberi var. sumatrensis, which is one of the most
misunderstood forms (Jalihal et al, 1984). Now we have changed our opinion,
have refrained here from giving a variety, and tentatively retained the nominal
name for our specimens.
Cai & Ng (2000) suggested that overall morphology of Kemp‟s (1918)
form Caridina weberi prox. var sumatrensis cannot be referred to any known
taxa, they are confident that it is new to science. However, they have not
included Chopra & Tiwari‟s (1949) report on the same form from Orissa in their
discussion. These authors suggested that the form represents distinct variety
but refrained from giving it a name. All the 6 subspecies are expected to be
elevated in future to the species while some to fall in to synonym of others (Cai
& Ng, 2001), for example C. weberi prox sumatrensis has fallen to C.
babaulti(Cai &Shokita,2006)
Colour: Transparent with red chromatophores arranged uniformly.
Size: 2.8 to 12mm in TL
Localities: Overhead water tank, Karachi University Campus ,Hub
Dam and Haleji Lake.
Distribution: India, Japan, Taiwan.
Genus Kalriana Zuberi, 1990
76
Diagnosis: Rostrum almost straight, longer than antennular
peduncle, 17-20 dorsal teeth on posterior and middle portion, 5 on preorbital
region, setae present in between teeth, anterior part of rostrum lacking teeth;
ventral teeth 4-7.Carapace smooth with strong antennal and branchiostegal
spines; former better developed and stronger; branchiostegal groove faint.
Antennal squama divided into two portions by a suture. Mandible without palp.
Arthrobranch on second maxilliped (Modified from Zuberi, 1990).
Remarks: Zuberi (1990) established four new species in the genus;
they are K. anissi, K. karachii, K. jhimphirensis, and K. sunahrensis and
provided a key for their separation.
KEY TO PAKISTANI SPECIES OF KALRIANA
(From Zuberi, 1990)
1.
Second maxilliped without suture between
dactylus--------------------------------------------------------------------2.
----------------3
2.
and
Second maxilliped with distinct suture on propodus-----
Telson with 6 pairs of dorsal spines ------------K. anissi .
-.
Telson with five pairs of dorsal spines----------------------
K. karachii .
3.
Ischia of fifth thoracic appendages with a strong spine.
Telson with five (or 6) pairs of dorsal spines-----------------------------------------------K. jhimphirensis .
Ischia of fifth thoracic appendages without a spine.
Telson with four pairs of dorsal spines------------------------------------------------------------- K. sunahrensis .
Kalriana anissi Zuberi, 1990.
(Fig. 30)
Kalriana anissi Zuberi 1990:75
Description: The ventral rostral teeth are always between five and
seven; their usual number is seven.
77
H
F
J
A
I
E
G
B
C
D
Figure 30: Kalriana anissi. A- Carapace, lateral view; B- Uropod ; CAntennule; D-Antenna; E-Second maxilliped; F- Third maxilliped; G- First
leg; H- Second leg; I- Third leg; J- Pleopod; -(Modified from Zuberi, 1990)
The telson possesses six pairs of dorsal spines arranged in two rows.
78
There is no suture between the dactylus and propodus of second
maxilliped.
The proportion between the length and breadth of dactylus of fifth
pereopod is four.
There is no spine on the anterior portion of the ischium of the fifth
walking leg.
Size:
15-20mm
Locality:
Keenjar Lake, Thatta, Sindh
Distribution:
Not reported out side Pakistan
Development: (Figs. 31-43) The just hatched larvae take 50 to 55 days
to become adult. There are usually 10 to 11 developmental stages in breeding
history of K. anisii, which were described by Zuberi, 1973 (unpublished) and
reproduced below:
Figure 31: Kalriana anissi Newly hatched larva (Modified from Zuberi,
1990)
Stage I larva:
Almost colourless, and possesses dark brown and orange coloured
patches on the uropod, cephalothorax and on the other parts of body. Rostrum
absent, carapace without spine, and eyes without stalks. Three pairs of
undeveloped biramous thoracic appendages and five abdominal segments.
Abdominal appendages absent, uropod with a deep groove and 16 setae.
Antennae possess 10 setae, but lack antennal feeler.
Stage II-5 Days Larva:
79
Almost colourless with dark brown colour patches on telson.
Rostrum developed, but seen with difficulty. Eyes stalked, mandible show no
cleft between incisor process and molar process. Antennular spine absent.
Antennae with 10 setae, the seta nearest to the developing flagellum the longest
and the remotest being the shortest. Maxillipeds still poorly developed. Thoracic
appendages developed and biramous. Abdominal appendages and their buds
absent. Telson as in stage 1, with rudiments of exopod and endopod.
Figure 32: Kalriana anissi Appendages of five days old larva. A- First
maxilla; B- Second maxilla; C- First maxilliped; D- Antenna; E- Thoracic
appendage; F- Fifth leg; G- Antennule; H- Mandible; I- Uropod (Modified
from Zuberi, 1973)
Stage III-10 Days Larva:
Almost colourless but patches of colour pigment may be seen on
carapace, appendages and uropod. Main pigment spot on the uropod
disappear. Rostrum as in stage-II, eyes stalked incisor teeth present in the
80
mandible. Exopod of third maxillipeds still not segmented. Outer and inner
flagella of the antennules present. Suture between coxa, basis, and precoxa
visible. Antennular teeth present. Suture on the squama of antenna absent but
the spine mildly developed. Thoracic appendages five pairs in number but not
developed. Abdominal appendages or their buds absent. Uropod divided into
the exopod and endopod at the posterior end of exopod, on endopod ten spines
present, suture of the exopod absent but possesses one movable spine, four
pairs of setae present at the tip of telson, outer most of them the longest while
the inner most the shortest. On the dorsal side of the telson, three pairs of
spines present.
81
Figure 33: Kalriana anissi
Appendages of ten days old larva. AAntennule; B- Antenna; C- First maxilla; D- Second maxilla; E- Mandible;
F- Thoracic appendage; G- Third leg; H – Third maxilliped; I- Telson; JUropod. (Modified from Zuberi, 1973)
82
Stage IV-15 Days Larva:
Larvae colourless but patches of colour pigment may be seen
throughout the body. Rostrum still not much developed. Mandible more
developed, molar process bears some teeth, between the two processes. A tuft
of hairs present, exopod of the second and third maxilliped still unsegmented.
Exopod of second maxilliped with two developing sutures, these sutures not
present on the endopod of third maxilliped but constrictions quite clear. Outer
and inner flagella of the antennules more visible. Suture between the
precoxa/coxa and coxa/basis visible but three teeth or spines on the suture not
present. Suture on the squama of the antenna not visible, the flagellum of the
antenna now three segmented but segmentation not very clear. The flagellum a
bit longer than the squama.
83
Figure 34: Kalriana anissi Appendages of fifteen days old larva. AAntennule; B- Antenna; C- First chela; D- Second maxilliped; E- Third
maxilliped; F- Second maxilla; G- Mandible; H- Second chela; I – Telson; JUropod; K- Pereopod. (Modified from Zuberi, 1973)
84
Thoracic appendages now uniramous gradually taking their usual
shape. In first thoracic appendage a bud like out growth visible, that also
undergoes some changes towards the formation of chelae but not very much
clear. Fourth thoracic leg has no suture, no teeth like structure on the dactylus.
Fifth leg with two visible spines present on the anterior end, two sutures also
present.
85
Figure 35: Kalriana anissi Appendages of twenty days old larva. AAntennule; B- Antenna; C- Mandible; D- Uropod; E- Telson (Modified from
Zuberi, 1973)
86
The buds of abdominal appendages visible but the distance between
them increased as compared to the previous stage. Exopod and endopod of the
uropod more clearly visible; spine on the outer margin of the exopod clearer but
suture still not present. Telson with three pairs of terminal and three pairs of
dorsal spines, but the distance between them increased as compared to the
previous stage.
87
Figure 36: Kalriana anissi Appendages of twenty five days old
larva. A- Antennule; B- Antenna; C- Third maxilliped; D- First
chela; E- Second chela; F- Fifth leg. (Modified from Zuberi, 1973
88
Stage V-20 Days Larva:
Larvae not colourless at this stage, the patches of pigment quite visible
from outside with naked eye.
Rostrum a bit more developed, mandible stouter and developed, both
processes quite clear and between them a row of small teeth present. Exopod
of first and second maxilliped still unsegmented, two sutures visible in the
endopod of second maxilliped as in the pervious stage, these sutures separate
merus ischium from the carpus and the carpus from dactylopropodus.
89
Figure 37: Kalriana anissi Appendages of twenty five days old larva. AFirst pleopod; B- Second pleopod; C- Third pleopod; D- Fourth pleopod;
E- Fifth pleopod; F- Telson; G- Uropod (Modified from Zuberi, 1973).
90
Outer and inner flagella of the antennules now become segmented.
Sutures between the segments clearly visible. Antennular spine at precoxa
more developed and stout. Suture on the squama of antenna not visible yet, the
flagellum four segmented and larger than the squama.
Thoracic appendages now much more clear. First thoracic appendage
with its chela a bit more developed. Second thoracic appendages also
developed the usual shape of second chela, the beginning of dactylopod visible.
Sutures between the segments quite visible in all the thoracic appendages.
Buds of abdominal appendages become biramous and more developed.
91
Figure 38: Kalriana anissi Appendages of thirty days old larva. AAntennule; B- Antenna; C- First chela; D- Second chela; E- Mandible; FThird leg; G- Fifth leg; H- First maxilliped; I- Uropod; J- Third maxilliped;
K- Second maxilliped (Modified from Zuberi, 1973).
92
Spine on the outer margin of the exopod of uropod remains unchanged,
its suture also clearly visible. Terminal spines of the telson become larger, outer
most of them the largest, while the inner most the smallest.
Stage VI-25 Days Larva:
Rostrum developed, pointed, and straight, not bent upwards or
downwards shorter than the scaphocerite of the antenna. Teeth present on the
ventral side.
93
Figure 39: Kalriana anissi Appendages of thirty five days old larva. AAntennule; B- Antenna; C- First maxilla; D- First maxilliped; E- Third
maxilliped (Modified from Zuberi, 1973).
94
Mandible and second maxilliped remain as in previous stage. Exopod
of the third maxilliped not segmented but in exopod, four sutures visible, one
between dactylo-propodus and carpus, other between the carpus and ischium,
third between the ischium and merus and the fourth between merus and coxa.
At the anterior end of the dactylus, three spines present in a row and one
terminal similar to adult conditions. A row of setae in the middle of the dactylopropodus and one spine present at the inner margin of it. A row of setae also
present at the suture between basis and carpus.
95
Figure 40 : Kalriana anissi Appendages of thirty five days old larva. AThird leg; B- Fifth leg; C- First pleopod; D- Second pleopod; E- Third
pleopod; F- Fourth pleopod; G- Fifth pleopod (Modified from Zuberi, 1973).
96
Thoracic appendages more developed. First and second thoracic
appendages acquired their normal structure. Suture between the dactylus and
propodus quite visible now. Tufts of hairs also appeared on the finger and the
palm of first and second chelae. The excavation at the joint of the carpus and
propodus quite clear now in the first chelae but not in the second chelae. Fourth
walking leg also acquired a bit more developed structure. Suture between the
dactylus propodus/ carpus and carpus ischio-merus quite visible, but suture
between ischium and merus yet undeveloped. Suture between the basis and
ischio-merus and one on dactylus present. Abdominal appendages quite
developed now. All of them acquired the normal structure, now possessing
coxa, exopod, endopod, and appendix interna.
97
Figure 41 : Kalriana anissi Appendages of forty days old larva. AAntennule; B- Antenna; C- Uropod (Modified from Zuberi, 1973).
Exopod of the uropod as in previous stage, the suture more visible with
two teeth. Terminal spines of the telson remain unchanged but the spines on the
dorsal side of telson now in four pairs.
98
Figure 42 : Kalriana anissi Appendages of forty five days old larva. AAntenna; B- Second maxilliped; C- Third maxilliped; D- Third leg; EFourth leg; F- Fifth leg (Modified from Zuberi, 1973).
99
Stage VII-30 Days Larva:
Rostrum well developed with twelve dorsal and two ventral teeth.
Antennal and hepatic spines absent. The structure and the sizes of rostrum and
scaphocerite remains unchanged. Rostrum straight and still shorter than
squama.
100
Figure 43: Kalriana anissi : Fifty days old larva: A- Antennule; B- First
maxilliped; C- First maxilla; D- First maxilliped; E- Mandible; F- Uropod
(Modified from Zuberi, 1973).
101
Mandible developed as typical of the adult. Teeth on the incisor
process and plates on the molar process may be observed. First maxilliped
bearing two palp like processes, a bulging portion almost in the middle, and four
bristles developed on the anterior terminal end. Exopodite of second and third
maxilliped unsegmented. Sutures between coxa-ischium-merus and meruscarpus quite clear in endopodite of second maxilliped. Sutures between carpuspropodus, and propodus-dactylus also visible but not very clear. Sutures clear
on the endopodite of third maxilliped. Ten teeth present at dactylo-propodus,
terminal hook is stronger.
Scutes quite clear in both flagella of antennules. Two small teeth visible
at the suture of coxa/basis, rest of the structure remains unchanged. Spines of
the squama and flagellum of antenna remain unchanged.
Thoracic appendages more developed. Sutures between propodite and
dactylopodite quite distinct now in the first thoracic appendage. Excavation
present at the suture of carpus and propodus and a bit deeper now. Second
thoracic appendage remains unchanged in the structure but sutures between all
the segments quite clear. Third and fifth walking legs more developed. Teeth
present, three at the dactylus, four on propodus, two on merus and one on
ischium of third walking leg. Similarly on the fifth thoracic appendage six teeth at
dactylus, five on propodus, one on carpus, two on merus and on ischium.
Abdominal appendages remained unchanged. Exopod of the uropod more
developed, its suture is complete, and six teeth appeared. Larger spines
developed on its endopod. Terminal and dorsal spines of the telson remain
unchanged.
Stage VIII-35 Days Larva:
Rostrum well developed, straight and pointed. Number of dorsal and
ventral teeth remains unchanged. Both spines, antennal and scaphocerite
remains the same. Rostrum still shorter than the squama.
The number of teeth increased on the inner margin of first maxilla. First
maxilliped acquired almost adult character. Second maxilliped remained
unchanged. On the endopod of third maxilliped all the sutures quite clear now.
Rows of teeth easily visible on dactylo-propodus. Two teeth present at the
suture of dactylo-propodus and propodus-carpus, five teeth may be seen at the
inner margin of carpus and four on the dorsal side almost in the middle and at
the anterior half. Three teeth at the suture of carpus and ischio-merus and one
tooth at the middle of the inner margin of ischio-merus. A spine also present on
the coxa.
Scutes of outer and inner flagella of the antennule quite clear and more
in number. Number of scutes greater in the inner flagellum than on the outer
102
flagellum. Two spines present at the inner margin of basis, three on coxa,
and three on precoxa. Antennular spines more developed and almost half of the
length of precoxa. Suture on the squama of the antenna almost half developed
and may be seen at high magnification, other structures of antenna remained
the same as in previous stage.
Thoracic appendages more developed. First and second thoracic
appendages remain unchanged. The number of teeth at dactylus remains
unchanged but at propodus only one spine on third walking leg, four teeth on
ischio-merus, one on anterior end of ischium and three teeth on the margin. The
number of teeth increases in fifth leg, there are ten teeth present at dactylus,
four on propodus, one on dorsal side of merus, two on ischium. Spicules also
visible on the outer margin of dactylus and propodus. Abdominal appendages
acquired the characters of the adult and bear exopod, endopod and appendix
interna. The structures of uropod and telson remain unchanged.
Stage IX-40 Days Larva:
Rostrum same as in previous stage. Numbers of dorsal teeth remain
unchanged but the number of ventral teeth increases from two to three and
located near the anterior end, but not very close to the tip. The structure of
mandible, first and second maxilla and first and second maxillipeds remain
unchanged. Number of teeth at the inner margin of dactylus increases from
three to four including hook at third maxilliped.
The number of scutes at outer and inner flagella of the antennule also
increases; other structures remained as in previous stage. Suture on the
squama of antenna became clearer.
Third, fourth and fifth walking legs also remain unchanged. Abdominal
appendages too remained as in previous stage. The number of spines
increases from six to eight at the suture of exopod of the uropod, two of them on
out side. Structure of telson remains unchanged.
Stage X-45 Days Larva:
Rostrum almost developed fully. Number of dorsal and ventral teeth
remains the same. Among the dorsal teeth, setae developed. The suture at the
squama of antenna about to complete. Other structures same as in previous
stage. In antennules, the number of scutes increases in outer and inner flagella.
At the suture of coxa-basis, three spines visible and at the suture of precoxacoxa a short but strong spine also present.
103
In the exopod of the third maxilliped no suture between the dactylopropodus and carpus, but present between carpus and ischio-merus. No suture
between the ischium and merus. In third maxilliped, the number of rows of teeth
on dactylus and propodus increases to eight. At the carpus five rows of spines
present almost acquiring the adult character.
Third, fourth and fifth walking legs now fully developed and acquired all
the adult characters. At the margin of dactylus of fifth walking leg, fifteen teeth
present, six on propodus, one on carpus, two on merus and one on ischium.
Remaining structures showed no change and at this stage and
remained as in previous stage.
Stage XI-50 Days Larva:
Rostrum, thoracic appendage, and abdominal appendages have
acquired all the adult‟s characters. First maxilla developed stout teeth on inner
margin and a strong palp. First maxilliped with poorly developed endopod and
highly developed exopod, acquiring all adult characters. In second maxilliped, a
suture developed to divide the ischio-merus into ischium and merus, no suture
between the dactylus and propodus.
Suture on the squama of antenna complete and very distinct now,
divides this suture squama into two parts, one from tip to suture and other from
suture to base. Scutes of inner and outer flagellum very distinct. Three spines
present at the suture of coxa/basis as in the previous stage. Five spines present
at the suture of coxa, precoxa, one marginal strong and stout spine present.
Eleven spines present at the suture of exopod of uropod. Out of them
three on the remote margin of the uropod, structure of uropod just as in adult.
Telson now developed six pairs of spines. Terminal spines remain unchanged
i.e. six in number.
Fifty days old, larvae have acquired all the characters of the adult.
Spines on the suture of the exopod of uropod vary in number from eleven to
sixteen. The number of movable spines may become four.
Kalriana jhimphirensis Zuberi, 1990
(Fig.44B)
Kalriana Jhimphirensis Zuberi, 1990: 76
Description: The ventral teeth of the rostrum between five and seven;
their usual number five.
104
The teslon possesses six dorsal spines on one side and five on the
other, in rows.
A very distinct suture present between dactylus and propodus of the
second maxilliped.
Figure 44: Telsons . A- Kalriana karachii; B- Kalriana jhimpirensis; CKalriana sunahrensis.(adapted from Zuberi,1973)
The proportion between the length and breadth of dactylus of fifth
pereopod is three.
A very distinct spine on the anterior portion of the ischium of fifth
pereopod.(from Zuberi,1990)
Localities: Keenjar Lake, Thatta, Sindh
Distribution: Not reported out side Pakistan
Kalriana karachii Zuberi, 1990
105
(Fig. 44A)
Kalriana Karachii Zuberi, 1990:75
Description: There are five to seven ventral teeth on the rostrum
The telson possesses five pairs of dorsal spines arranged in rows.
No suture between dactylus and propodus of the second maxilliped.
The proportion between the length and breadth of dactylus of fifth
pereopod is three.
No spine on the anterior portion of the ischium of the fifth walking
leg.(from Zuberi, 1990)
Locality: Keenjar Lake, Thatta, Sindh
Distribution: Not reported out side Pakistan
Kalriana sunahrensis Zuberi, 1990
(Fig. 44C)
Kalriana sunahrensis Zuberi, 1990: 76
Description: The ventral rostral teeth are never more than five, their
usual number is three.
The telson possesses four pairs of dorsal spines arranged in rows.
A very distinct suture is present between the dactylus and propodus of
second maxilliped.
The proportion between the length and breadth of dactylus of fifth
pereopod s three.
There is no spine on the anterior portion of the ischium of fifth pereopod
(from Zuberi, 1990)
Locality. Keenjar Lake, Thatta, Sindh
Distribution: Not reported out side Pakistan
106
3.1.4 Family Palaemonidae Rafinesque, 1815
Diagnosis: Rostrum compressed and usually dentate. Antennal spines
always present whereas the hepatic and branchiostegal spines may be present
or absent. Eyes well developd. Mandible with or without a palp, incisor, and
molar processes separated by a deep cleft. Ultimate segment of the second
maxilliped laterally attached with the penultimate segment, an exopod, an
arthrobranch and a pleurobranch may be present or absent. Chelae of the first
pair of pereopods small, while those of second pair large and robust. Carpus of
the second pereopod unsegmented. Epipods absent from all pereopods. Telson
narrow distally. Second pleopod with appendix masculina in male (Modified from
various sources)
Remarks: Serious attempts have been made to settle the classification
of this group of carideans (Bruce, 1986, Chace & Bruce, 1993). These authors
define their own work “far from definition”. Later authors refrained from a more
detailed classification of this group to avoid further confusion.
Morphologically abnormal specimens were encountered in
several species in otherwise normal natural stock as already mentioned by Dutt
& Ravindranath (1974).
The family Palaemonidae is further divided into two sub-families i.e.
Pontoninae and Palaemoninae and can be separated with the help of key given
in start.
Subfamily Palaemoninae Rafinesque, 1815
Diagnosis: Both rami of upper antennal flagellum fused in the basal
part. An appendix interna generally present on the second pleopod of the
female. Both, the appendix interna and appendix masculina present on the
second pleopod of male. Pleurobranch present on the third maxilliped. Posterior
margin of the telson provided with two pairs of spines and one or more pairs of
setae (From various sources).
Remarks: The subfamily includes 21 genera and 2 subgenera
(Jayachandran, 2001). The subfamily Palaemoninae in Pakistan pertains to
species belonging to genera Macrobrachium , Exopalaemon, Nematopalaemon
107
, Palaemon , Leandrites, Leptocarpus, and Palaemonetes. They can be
separated with the help of the following key.
The record of Palaemonetes by Qadri (1960) is not yet confirmed
therefore not treated further by us.
KEY TO THE PAKISTANI GENERA OF PALAEMONINAE
1.
-----2.
Branchiostegal spine present--------------------------------------------------------
-----6.
Branchiostegal spine absent --------------------------------------------------------
2.
----3.
Mandible without palp------------------------------------------------------------------
-----4.
Mandible with a palp-------------------------------------------------------------------
3.
First pleopod of male with well developed appendix interna on the
endopod. Branchiostegal groove absent. Propodus of fifth leg without transverse
rows of setae on the distal part of the posterior margin------------------------------------------------------ Genus Leandrites.
First pleopod of the male without appendix interna on the endopod.
Branchiostegal groove visible as sharp line. Propodus of fifth pereopod with
transverse rows of setae on the distal part of the posterior margin -------------------------------------------------Genus Palaemonetes.
4.
Rostrum with an elevated basal crest of teeth. Pleura of fifth abdominal
segment with apex broadly rounded. Mandibular palp three-segmented----------------5.
Rostrum without an elevated basal crest. Pleura of fifth abdominal
segment generally ending in a small sharp point. Branchiostegal groove
present------------------------------------------------------------------------------------------------------Genus Palaemon
5.
Dactylii of last three legs enormously lengthened, longer than carpi and
propodii together. No branchiostegal groove on the carapace. Stylocerite with a
large tooth on the upper surface------------------------------Genus Nematopalaemon
Dactylii of last three legs always shorter than propodii, never
excessively long. Branchiostegal grooves present on the carapace. Stylocerite
108
without a large dorsal tooth-----------------------------------------------------------------------------------------------------Genus Exopalaemon.
6.
Hepatic spine absent--------------------------------------------------------Genus
Leptocarpus .
-
Hepatic spine present------------Genus Macrobrachium
Genus Exopalaemon Holthuis, 1950
Diagnosis: Rostrum with highly raised dentate basal crest; carapace
with branchiostegal spine and branchiostegal suture. Hepatic spine lacking.
Fourth thoracic sternite without slender median process. Mandible with
prominent palp. Last three pereopods with simple and not biungiculate dactylii,
shorter than propodii. Endopod of male of first pleopod without appendix interna
(Modified from various sources)
Type Species: Palaemon styliferus H. Milne Edwards, 1841
Gender: Masculine.
Remarks: Six valid species are included within the genus, a thorough
examination of the genus is suggested by Jayachandran (2001), only the type
species has been recorded from here .
In an old sample without any clue of supporting data there are at least
30 specimens measuring 5mm in CL, close to E. styliferus to differing from it in
having almost no crest on rostrum and showing increased number of teeth,
since they are not in good preserved condition it is difficult to name them until
new fresh collection is available.
Exopalaemon styliferus (H. Milne Edwards, 1840)
(Figs. 45-46)
Palaemon longirostris H. Milne Edwards, 1837:394
Palaemon styliferus H. Milne Edwards, 1840:638; Rathbun, 1902:51;
Suvatti, 1937:50
Leander longirostris Henderson, 1893:439
109
Leander styliferus Kemp, 1915:273; 1917:214; 1925: 289; Balss,
1930:316; Panikkar, 1937:345; Chopra, 1939:223; 1943:5; Qadri, 1960:259;
Ahmed, 1967:17
Palaemon (Exopalaemon) styliferus Holthuis, 1950:47 Exopalaemon
styliferus Chace & Bruce, 1993:5; Dore & Frimbodlt 1987:72; Salman & Bishop,
1990:21; Jayachandran, 2001:25
Description: The rostrum is very long, extending far beyond antennal
scale by about 1/3 its length, basal crest is greatly raised; the upper margin of
the rostrum is provided with 6-8 teeth, of which one is post-orbital, 5-6 are
proximal and crowded at raised crest above the orbit, the tooth size increases
from proximal to distal teeth of this group. The rostrum beyond the basal crest is
very slender and prominently up curved. The distal half of the rostrum is
edentate. The distal end of the rostrum is armed with one or two teeth. The
ventral margin of the rostrum has 6-9 teeth. The carapace is smooth, the usual
antennal and branchiostegal spines are present, the branchiostegal groove lies
just above the branchiostegal spine.
The abdomen is glabrous; the pleurae of first to third somites are
broadly rounded at posteroventral angle, those of fourth and fifth are directed
backwards; that of sixth ends in a spine.
Figure 45: Exopalaemon styliferus, Animal in lateral view.
The telson is slender, elongate. The dorsal margin bears two pairs of
spines, both located in its distal half; the posterior end of telson is pointed.
110
The first pair of chelate legs is slender, almost reaching to the end
of the antennal scale; the ischium is shorter than the merus; the merus is
shorter than the carpus or may be equal, in some specimens, the fingers are
only slightly longer than the palm.
Figure 46: Exopalaemon styliferus. A- Anterior part of body in lateral view;
B-F- First to fifth legs (Modified from Jayachandran, 2001).
The second chelate pereopods are slender and longer than the first;
the ischium is longer than the merus; the merus is almost equal to or slightly
shorter than the length of the carpus, the carpus is shorter than the chela, the
palm is inflated; the fingers are slender; the tips of fingers are curved, and
distinctly longer than the palm.
The three pairs of non-chelate pereopods are slender, their dactylii are
simple.
The pleopod and uropods are typical in structure.
Colour: Body with fine dots fairly well developed on the rostrum, telson,
and uropods.
Size: 90mm in TL (male), 86mm in TL (female)
Localities: Shah Bunder and Keti Bunder, Thatta, in rice fields
Distribution: India, Bangladesh; Arabian Gulf and Iraq, Gulf of
Martaban; mouth of Ye River, Myanmar, Hingis Island, Green Island,
111
Tennasserim; Mergui Archipelago, Malay States; Pulu Burong, Sarawak;
Thailand.
Remarks: For larval development see Al-Abbad etal( 2008).
Genus Leandrites Holthuis, 1950
Diagnosis: Rostrum without elevated basal crest; carapace with sub
marginal branchiostegal spine, without hepatic spine or branchiostegal suture;
fourth thoracic sternite with slender median process; 3 posterior pairs of
pereopods with dactylii simple, shorter than propodus; endopod of male first
pleopod with appendix interna. ( from Chace & Bruce, 1993)
Type species: Leander celebensis de Man, 1881
Gender: Masculine
Remarks: Only four species have been included in the genus
Leandrites today. Out of these L. celebensis has been recorded from here.
Leandrites celebensis (de Man, 1881)
(Fig. 47)
Leander celebensis deMan, 1881:141
Palaemonetes hornelli Kemp, 1925: 318; Nataraj; 1942: 468
Leandrites celebensis , Holthuis, 1950: 36; Bruce, 1987:59; Chace
&Bruce, 1993: 7; Jayachandran, 2001:37; Kazmi et al, 2009:155
Description: The rostrum reaches to or a little beyond the end of the
antennal scale. It is straight rarely a trifle upturned at the tip and in lateral view is
deep. The upper border, very feebly convex, bears from 13 to 17 forwardly
directed teeth with setae in the interspaces; the first and second teeth are
placed on the carapace behind the orbit and the first is twice as far distant from
the second as the second is from the third. The remaining teeth on the upper
border are equidistant, the series extending to the apex; the middle teeth are the
largest, the foremost is very small and, though near the apex does not give it a
112
bifid appearance. The lower border of the rostrum is convex, but not deeply
excavated at the base. It is rather thickly fringed with setae throughout its length
and bears from three to five teeth, which are rather smaller than those of the
upper border are. The lower angle of the orbit is broadly rounded and the
antennal spine is strong. The branchiostegal spine is almost equally strong and
is set well back on the carapace, the tip reaching the margin.
Figure 47: Leandrites celebensis A-Rostrum and anterior margin
of carapace; B- Telson, ventral view of tip; C-D-First and second legs; EFifth leg
The telson is shorter than the uropods and bears, as usual two pairs of
dorsal spines; the firs pair is situated at, or a little behind the middle of the
telson-length and the second is nearer to the first than to the apex. The apex is
pointed in the middle with a single very stout seta and two spines on either side.
The outer spines are very short, only about one-fifth the length of the inner, the
latter much exceeding the setae in length and reaching beyond the ends of the
uropods. The external border of the outer uropod is setose.
The eye is large with a well-marked ocular spot in contact with the
cornea.
The antennular peduncle is not much shorter than the antennal scale.
The lateral process does not reach to the middle of the basal segment and the
outer margin of this segment ends in a spine, which does not usually reach as
far forwards as the strongly convex border, which intervenes between the
113
spines and between the spine and the articulation of the second segment.
The second and third segments are short, stout and sub equal in length. The
two rami composing the outer antennular flagellum are fused basally for a very
short distance. The antennal scale is more than 3 times as long as wide; its
outer margin is straight or very slightly convex and ends in a spine, which does
not reach as far forwards as the broadly rounded apex of the lamella.
The mandible is without a palp and the incisor- molar processes
terminate in three sharp teeth. The maxilla and first maxilliped are normal in
structure.
The first legs are slender and reach beyond the antennal scale; the
ischium is smaller than the chela; the merus is sub equal to carpus and the
fingers are longer than the palm. The second legs are very long, extending
beyond the antennal scale by the chelae and from one half to three- quarters of
the carpus. In females, the carpus is slightly longer than the merus and a little
shorter than the chela, the proportionate lengths of merus, carpus, and chela
being about as 12: 13: 15. The carpus is widened distally ; the palm is swollen
wider than the adjacent parts of the carpus . The finger meet throughout their
length when the claw is closed, each being provided at the base with a single
very small tooth. The fingers have in turned tips and are usually a little longer
than the palm. Except for a few very short setae on the fingers, the leg is naked.
In general, the legs of male closely resemble those of the female
except in second chelae where proportionate lengths of the segments are
different. The lengths of the merus, carpus, and chela are as 12: 17: 13.The
carpus thus being almost 1 times as long as the merus and nearly one third
longer than the chela. The fingers are a trifle longer than the palm.
The last three pairs of legs are very slender and increase in length from
before backwards; the third reaches beyond the antennal scale by about onethird of the propodus, the fifth by about half the propodus. In each pair, the
propodus is from 2 - 3 times the length of the dactylus. The latter segment is
slightly curved and extremely slender, from 12 to 14 times as long as its basal
breadth. The legs are almost naked except a few long setae on the anterior
margin of the dactylus and for a few spinules near the distal end of the propodus
of the fifth pair.
The sixth abdominal segment, measured dorsally, is rather more than
one and a half times the length of the fifth. In the first pleopod of the male, the
endopod is oval and bears on the inner side an appendix interna, which projects
far beyond its distal end. In the second pleopod of the male, the two appendices
are of equal length.
114
Remarks: Pillai (1974) has described the laboratory maintained
larvae. The adult specimens at hand show no difference from those described
by earlier workers except that the rotsral dentition has a lesser number of teeth
than found in the Australian specimens (Bruce, 1987).
Size: 4.5mm In CL (male) ,2.2 -3.5mm in CL (female)
Habitat: Often in brackish water
Locality: Ambro creek, Ghora Bari
Distribution: India, Indonesia, Celebes, Singapore, Northern territory
of Australia.
115
Genus Leptocarpus Holthuis, 1950
Diagnosis:
Medium- size slender bodied forms. Body subcylindrical. Rostrum well developed, long, with teeth on both margins. Basal
crest of rostrum generally slightly raised. Setae present between teeth of both
dorsal and ventral margins.
Upper margin with one row of setae, lower with two rows. Carapace
smooth, with only antennal spines located some distance below rounded orbital
angle. Distinct and sharp branchiostegal groove present.
Abdomen smooth. Pleurae of segments I-II broadly rounded along
posteroventral margin, those of IV and V directed backwards, that of VI spinous.
Segment VI distinctly longer than V.
Telson slender, elongate, triangular. Dorsal surface with two pairs of
spines. Posterior end sharp, flanked by two pairs of spines. Outer pair of these
spines much shorter than inner, inner pair overreaching tip of telson. Few
plumose setae present between longer spines. Eyes well developed, with
hemispherical cornea containing black pigment. Ocellus present. Basal segment
of antennular peduncle broad. Stylocerite sharp. Anterolateral spine slender,
reaching midlength of intermediate segment. Intermediate segment of peduncle
much shorter than distal one. Upper antennular flagellum distally bifid. Free part
of shorter ramus much longer than fused part. Antennal scale well developed.
Outer margin ends in sharp spine overreached by lamella. Mandible highly
chitinised. Molar process with flat chewing edges. Incisor process tridentate.
Palp 3- segmented. Maxillula usual in shape. Endite of maxilla deeply cleft.
Third maxillipeds slender and with exopod, endopod and epipod.
First pereopods very slender, fingers unarmed but with tufts of setae.
Carpus elongated. Second legs also slender. Carpus elongate. Last three pairs
of pereopods slender, non-chelate. Dacylii simple. Appendix interna present in
all pleopods, in second pleopod of male with appendix masculina also. Uropods
distinctly longer than telson. ( from Jayachandran, 2001)
Type species: Leander fluminicola Kemp, 1917
Gender: Masculine
Remarks:. Only three species have been reported under this genus to
date, all from fresh and brackish water, one species is present in our collection.
Rajyalakhsmy (1961) and Pillai (1973) have described the larval stages
116
Leptocarpus potamiscus (Kemp, 1917)
(Fig.48)
Leander potamiscus Kemp, 1917: 225, 1918: 270, 1925:288; Rai,
1933:886; Gordon, 1935: 629
Palaemon potamiscus Suvatii, 1937: 50
Leptocarpus potamiscus Holthuis, 1950:97; Liu et al, 1990: 252; Chace
& Bruce,1993:8; Jayachandran, 2001:43; Cai & Ng,2002: Li et al, 2004:520;
Kazmi & Kazmi, 2009:131.
Description: The rostrum is very long, extending beyond the antennal
scale by half of its length, its tip is strongly up curved; the basal crest is slightly
raised; the upper margin has 7 teeth, of which 1 tooth is post-orbital. The
rostrum beyond raised part is edentate, the lower margin is armed with nine
equidistant teeth.
Figure 48: Leptocarpus potamiscus. A- Rostrum and anterior part of
carapace; B- Telson; C- Antennule; D- Antennal scale; E- First leg; F-
117
Second leg; G- Distal part of third leg.( Modified from Jayachandran,
2001).
The carapace is provided with only antennal spine, the branchiostegal
groove is distinct.
The abdomen is smooth; pleurae of first to third segment are broadly
rounded at posteroventral margin, those of fourth and fifth are directed
backwards, that of sixth is spinous.
The telson is slender. The dorsal surface is provided with two pairs of
spines; of which first pair is slightly proximal to middle of the telson, second pair
is much in advance of midway point between first pair and apex. Distal end has
also two pairs of spines; a pair of feathered setae is present between inner pair
of spines.
Basal segment of the antennular peduncle is broad. The stylocerite is
sharp extending proximal 1/3 distance of it. The anterolateral spine is sharp,
extending slightly beyond convex anterolateral margin, long setae are present
on ventral part of anterolateral and anterior margin of the basal segment, the
middle segment is the smallest, the distal segment is elongate. The upper outer
antennular flagellum is bifid distally, fused basally for about nine joints. The
antennal scale is elongated and broadly ovate. Outer lateral spine is situated at
2/3 distance from the base. The mandibular palp is 3-segmented, basal two
segments are equal, distal segment is the longest.
.
Both the chelate legs are very slender. First pair reaches the apex of
antennal scale when extended. The ischium is equal to or sub equal to the
merus; the carpus is longer than the chela. The fingers are slightly shorter than
the palm and provided with short setae.
The second chelate leg reaches beyond the antennal scale by its chela
and portion of carpus, the ischium is longer than the chela but shorter than the
merus, the carpus is longer than the chela; the palm is longer than the fingers.
Fingers are not notably spoon-shaped.
The nonchelate pereopods are slender. The first pair extends beyond
the antennal scale by more than length of dactylus, the third pair by dactylus
and 1/2-2/3 length of propodus. The dactylii of all the 3 pairs are very short and
simple.
118
Remarks: The single specimen at hand is an ovigerous female, not
intact, and parts are missing therefore its identification, as L. potamiscus is only
tentative, the description of the missing parts is taken from the literature.
Colour: Brown coloration on dorsal part of the edentate rostrum.
Size:
48 mm (female)
Habitat: Inhabits slightly brackish waters.
Distribution: India, Andaman Archipelago, China, Penang, Thailand,
Malaya States, Sumatra, Java. The present record from Pakistan forms an
important enlargement of the range of distribution of the species.
Genus Macrobrachium Bate, 1868
Diagnosis: Body compressed and robust in general appearance.
Rostrum well developed, serrated and laterally compressed. Carapace armed
with antennal and hepatic spines. Branchiostegal groove present and visible as
a sharp line. Eyes generally well developed. Antennular peduncle consisting of
three segments, the basal one is the largest. Scaphocerite well developed.
Mandibles complete with an incisor, a molar process, and a three-segmented
palp. Exopods present on all the maxillipeds.First and second pairs of
pereopods generally symmetrical. First pair of pereopods slender. Fingers and
the palm nearly of equal size. Cutting edges of the fingers smooth. Second
pereopods robust, longer than the others. Endopod of the first pleopod in female
bearing only an appendix interna; in males both the appendix interna and
appendix masculina present (Modified from various sources).
Type species: Palaemon americanum Bate, 1868
Gender:
Neuter
Remarks: The genus is commonly seen in fresh, sometimes brackish
water; very few species are marine as juveniles. Some species are troglobite
and epigean (Li etal, 2006). Amongst the decapod crustaceans, the genus
Macrobrachium is one of the most successful groups invading the freshwater
(Jalihal et al, 1988).
The genus Macrobrachium is known to be taxonomically a very
complex group owing to its extreme range of variations and overlapping of
characters particularly in the juveniles. Males and females of many
Macrobrachium species reach sexual maturity long before attaining their final
size and shape (Holthuis, 1950).
Based on characters of size and second chelipeds, Tiwari (1952)
divided the Indo-Burmese species of Macrobrachium into two groups. Later
119
Jalihal et al (1988) divided further one of the Tiwari‟s groups into four
subgroups.
The two groups also differ in size with reference of sex, while males are
much bigger in large species; some times left and right legs are conspicuously
different in shape. It is the females, which are predominant and large in smaller
species.
The females also should be given due importance as the taxonomic
studies of both sexes is becoming imperative with their growing importance in
aquaculture The distinct patterns of larval development perhaps reflect that
Macrobrachium is not a homogenous genus and appears to be a polyphyletic
group. Jalihal et al (1993) divided the genus in three basic types, based on the
number of larval stages. Type 1: prolonged or normal type. Further divided into
groups. Group 1a: includes coastal, of medium sized to large species(50320mm), species which generally exhibit sexual dimorphism wherein the males
are larger and provided with larger and / or stouter second chelipeds e.g.: M.
dacqueti, M. malcolmsonii, M. idella, M. equidens. Group 1b: includes medium
sized (25-55mm) prawns that are generally found in slow moving or impounded
water bodies, require salt for metamorphosis, no representative is found in
Pakistan. Type 2: partial abbreviated type, inhabitants of upstream, hill stream
and impounded water bodies, medium size prawns (25-70mm), males do not
exhibit sexually dimorphic features but are generally smaller than the females
with some exceptions e.g.: M. lamarrei lamarrei, M. shahpuri. Type 3:
completely abbreviated type: hill stream species so far represented by in
Pakistan by M. dayanum, medium sized (30-65mm), males do not show sexual
dimorphism.
Jayachandran (2001) suggested a new subgenus namely Allobrachium
to accommodate the species with unequal second chelipeds.
According to Tiwari (1952) 10 new species and 4 new subspecies:
“emanate” from India, Pakistan, Myanmar, Andamans, and Sri Lanka, as types are
deposited in the zoological survey of India, Calcutta they were not seen. Later
workers (Cai & Ng, 2000) did not mention Pakistan in distribution of these.
Wowor et al(2009) did a molecular phylogenetic analysis of the diverse
Southeast and east Asian species of Macrobachiumand tracked the evolution of key
lifehitory traits.
Attempt to construct a key to the species so far recorded from Pakistan had
to be curtailed because the key has to be based on adult males only and a key
based on females is ineffectual.More than 180 valid species and subspecies are
120
now generally recognized from different parts of the world , 17 species have
been collected from Pakistan.
121
. Material of an indeterminate species probably of M. superbum
was sent to Dr. X.Li for determination.His response is awaited.
Synopsis to the Pakistani species of
Macrobrachium
numericals indicate rostral formula)
Telson extending beyond outer
Telson of same level as outer
lateral spine of uropodal exopod.
Chelipeds strongly unequal.
Generally scabrous carapace
12-15/13
M. scabriculum
5-14/2-8, chelipeds densely
spinulose
M. altifrons ranjhai
lateral spine of uropodal exopod.
Chelipeds equal or subequal.
Generally smooth carapace,
rarely with sparse prickles
Carpus of second leg
longer than chela
Carapace completely smooth
10-12/4-6.long appendix
masculina
M. lamarrei korangii
6-10/6-9
Chelipeds slender not
hairy
M. lamarrei lamarrei
,6-9/5-8, rostrum, slender,
Upper rostral teeth
with a subdistal gap
M. shahpuri
11/6, basal crest upward,
movable finger with
pubensence M.
Carpus of second leg shorter than
chela
Carapace almost smooth
5-11/4-7, carapace smooth,
fingers densely pubescent
on grooves M. dayanum
9-11/5-6
Fingers covered with velvety
hairs
M. equidens
8-11/5-8,Carapace smooth
M. naso
2-15/4-5, movable finger
completely pubescent
M. idella idella
11-14/11-13,Carpus shorter
than chela in smaller
specimens, equal in longer
specimens,
carapace
smooth
M. dacqueti
, rostrum sigmoid, tip trifid,
9/6 M. Sp
10-11/5, carapace scabrous with
minute spines, all segments of
second leg entirely velvety
pubescent
M. rude
rostrum lanceolate, 9/2,
carapace glabrous,
M. taunsi
m. malmalcolmsoni
, rostrum straight, 8/4,
carapace smooth
M. tirmizi
9-11,5-6, rostrum,long with
crest, M.m. kotreeanum
, 10-12/6, dorsal teeth
arranged in three groups,
carapace a smooth
M. lehiai
122
Macrobrachium altifrons ranjhai Tiwari, 1963
(Fig.49)
Macrobrachium altifrons ranjhai Tiwari, 1963:237; Chace & Bruce,
1993:9; Jayachandran, 2001: 74
Description: The rostrum is short, reaching as far as the third segment
of antennular peduncle but fails to reach the end of the squamose portion of
antennal scale. Proximal half of the rostrum is straight; the distal part is slightly
downwards. The upper margin is armed with 7-11 teeth of which 2-3 are post
orbital, small setae are present between teeth of both dorsal and ventral
margins. The ventral margin has 1-3 teeth. The carapace is scabrous; the
antennal spine is stronger than the hepatic spine; the hepatic spine is situated
below and behind the level of antennal spine. Abdominal somites are glabrous
except the last somite.
123
FG
F
GH
I
Figure 49: Macrobrachium altifrons ranjhai. A- Rostrum, dorsal view; BAbdominal somites fourth to sixth; C- Telson; D- Antennule; E- Mandible;
F-F— Second leg; G-G- - First leg; H- Fourth leg; I – Pleopod.
The telson is stout and scabrous posteriorly; the dorsal surface is
armed with two pair of spines, situated in the distal half, the distal end has two
pairs, the outer pair is smaller and immovable, the inner pair is movable and
surpasses the tip of the telson, long stiff setae are present between the inner
pair of spines.
The basal segment of antennular peduncle is the largest; the stylocerite
is one third of basal segment, the anterolateral spine fails to reach the end of
second segment. Spine of scaphocerite is shorter than squamose portion.
The oral appendages are normal.
The first pair of chelate legs is slender; it reaches beyond the apex of
antennal scale by entire chela when extended; the ischium is small, finger and
palm are more or less equal in size; the carpus is the longest segment, about
twice the length of chela.
124
The second pair of legs is slender, the ischium is the smallest
segment; the merus is longer than the carpus, the palm is compressed, the
fingers are almost equal in size and more than half of the palm; cutting edges of
both the fingers bear denticles, movable finger has 5 denticles in proximal half,
the ultimate and penultimale denticles are stronger. The last three legs are
slender, the dactylus of third leg are half the length of the propodus, posterior
margin is almost straight gradually narrowing and curved towards tip.
Remarks: The second legs are more slender than those of other
subspecies M. altifrons altifrons.
Size: 52- 70mm in CL
Habitat: Small streams, ponds, canals
Localities : Kabul River at Nowshera, Peshawar, Kala Pani, Rissalpur,
Salt Range in Jhelum, Ravi near Neaz Beg, and now from Mograh near Taunsa
Barrage, Muzaffargarh canal.
Distribution: This subspecies is confined to upper Indus basin.
Macrobrachium dacqueti (Sunier, 1925)
(Fig. 50)
Cancer (Astacus )carcinus-Herbst, 1792:58(not Cancer carcinus
Linnaeus, 1758).
Astacus carcinus-Fabricius,1798:479 (part) (not Cancer carcinus
Linnaeus, 1758).
Palaemon carcinus – Fabricius, 1798:402(part);Olivier, 1811: 659; de
Man, 1888:280; Henderson, 1893:340,441; Henderson &Matthai, 1910:281;
Kemp, 1918:255; Menon, 1938:292; Tiwari, 1955:232;Ahmed, 1958:23; Qadri,
1960: 256; Shakoor, 1968:5 (not Cancer carcinus Linnaeus, 1758)
Palemon carcinus –H. Milne Edwards, 1837:395(not Cancer carcinus
Linnaeus, 1758)
125
Palaemon (Eupalaemon) carcinus de Man, 1902:475, 763 (part);
Nobili, 1900:480;
Figure 50: Macrobrachium dacqueti. A- Anterior carapace, male; B- Same,
female; C- Telson and uropods; C’- Apex of telson; D- Right cheliped,
male; E- Same, female; F- Right cheliped of juvenile male
Plaemon d’ Acquiti Sunnier, 1925:117.
Macrobrachium
Linnaeus, 1758).
carcinus-Suvatti,
1937:49
(not
Cancer
carcinus
Macrobrachium
rosenbergii
-Holthuis,
1950:
111,
(part);Johnson,1961:56; Hedgecock et al., 1979:873 (part); Liu et al., 1990:104;
126
Ng, 1990: 197; 1994: 75; 1997:269; Naiyanetr, 1998:33; Jayachandran,
2001: 158; Mather & de Bruyn,2003:4 (part);de Bruyn et al.,2004:251 (part)
Macrobrachium rosenbergii schenkeli Johnson, 1973:277.
Macrobrachium rosenbergii dacqueti –Holthuis, 1995:148; 2000; 16 ;
Cai & Dai, 1999:233; Wowor & Choy, 2001:286;Cai & Ng, 2002:78 ; Cai et al,
2004:582.
Macrobrachium dacqueti Wowor & Ng, 2007:326.
Description: The rostrum is long, its tip distinctly extends beyond the
distal end of scaphocerite in male young specimens (24-29mm in CL), becomes
relatively shorter with age, in fully adult males (65 mm CL and larger ) the tip
slightly extends beyond or reaches the distal end of scaphocerite, the basal
crest is high to moderately high and sinuous not effected by sex and age armed
dorsally with 8-15 teeth including one apical tooth , the teeth above orbit are
closely spaced, and on to distal two third of rostrum are closely spaced, two or
three teeth are completely postorbital, ventral margin is armed with 6-16 teeth,
first tooth is located on about proximal one fifth or one fourth. The carapace is
spinulate.
The telson is moderate, stout, and glabrous.
The cornea is well developed.
The distal half of merus of second pereopods extends beyond end of
the scaphocerite. Elevated, widely spaced large spines are abundantly present
on all segments of the chelate legs of adult males except the dactylus, the latter
is thickly covered by pubescence, the chela is more than twice the carapace
length; the fingers are straight, the palm is sub-cylindrical, outer and upper
margins are more densely covered with smaller spines than inner and lower
margins; the fingers are less than half of the palm length, not gaping; the
dactylus is provided with two large teeth on proximal one third, the movable
finger has one large tooth at the distal end and two smaller teeth proximally on
the cutting edge; the carpus is slightly smaller than the palm, sub-cylindrical; the
merus is straight, longer than the ischium. The third pereopod extends beyond
the scaphocerite by distal half of propodus.
All the segments are armed with medium sized spines; the dactylus is
stout, curved.
127
Remarks: The species lies in M. rosenbergii species group and
weberi- rosenbergi complex in this group (Johnson, 1962) . It can be easily
separated from M. rosenbergii by having second pereopod covered with large
spines (except for dactylus) while third to fifth pereopods are covered with
medium sized spines only. The species has 15 associated synonyms. The case
was open wether it is just one species. It started with Johnson (1960), then
through morph metric and allozyme work and studies on mitochondrial DNA and
micro satellite by Chan et al( 2005), and Wowor&Ng, (2007) through multiple
discriminate analysis of the morph metric data set confirmed two distinct and
valid species within M. rosenbergi i.e. M. Rosenberg and M. dacqueti ;the
biology of their larvae and colouration is also found different.The last authors
commenting that the majority of research on giant freshwater prawn is in fact on
M. dacqueti and not on M. rosenbergi.
Colour: Carapace with several black bands; several dark blue areas
visible near margins of abdomen segments, telson and uropods, and reddish
brown (=dark orange) markings present on each pleural condoyle.
Size: Fully adult male 65mm in CL.
Habitat: In freshwater occurring in rivers and lakes, it is also commonly
collected in brackish/estuarine waters. ovigerous females are seldom collected
from fully freshwater habitats since breeding takes place in the estuarine
stretches of rivers and perhaps sometimes in the sea. After breading, the
females return to freshwaters and the whole process is repeated several times
(Johnson, 1973 as M. rosenbergi). The larvae also migrate upstream so that the
juveniles inhabit the freshwaters until reaching the adult stage after which the
breeding migration takes place (Wowor & Choy, 2001).
Localities: Sukkur to Keti bunder, Rahim Yar Khan
Distribution: Asian mainland to the western border of Huxley‟s Line
excluding Palawan, i.e. from eastern part of Pakistan, India, Sri Lanka and
Southern China up to Borneo and Java (Wowor & Ng, 2007).
128
Macrobrachium dayanum (Henderson, 1893)
(Fig. 51)
Palaemon Dayanus Henderson, 1893:443;
Palaemon dayanus; Nataraj, 1942:468; Chopra & Tiwari, 1949:215;
Ahmed, 1957:20; Shakoor, 1968:2;
Macrobrachium sintangense Kamita, 1974:7
Macrobrachium dayanum Holthuis, 1950:197; Siddiqui, 1976:70;
Jayachandran, 2001:89; Cai & Ng, 2002:61
Description: The rostrum is usually almost straight, and extends to the
end of the antennal scale, with the dorsal margin having 7-9 teeth and the lower
with 5-6 teeth on the upper margin; the six proximal teeth are equidistant, and
separated by a wider interval from two, or more rarely three, smaller sub apical
teeth, with are placed close together, while the second and occasionally the
third, proximal tooth is placed above the orbital margin; on the lower margin the
teeth are equidistant, and slightly decrease in size towards the apex. The
carapace is smooth, with the hepatic spine rather small, and a faint sulcus,
which commences below the level of the latter, extends back almost to the
middle of the side wall of the carapace.
129
Figure 51: Macrobrachium dayanum. A- Anterior carapace; B- Telson; CFirst leg; D- Second leg; E- Same, chela enlarged, hairs removed (
Modified from Jayachandran, 2001)
The abdomen is glabrous. The telson is shorter than the uropods, its
apex is rather broad, but with a short median spine; the inner spinules are
considerably longer than the median point, and more than twice the length of
the outer spinules.
The distal end of the antennal scale is rounded, and scarcely angulated
internally.
The first legs exceed the antennal scale by the length of their fingers.
The second legs are of equal size, and rather short, being shorter than the
body, but moderately stout; they are pubescent, and very slightly scabrous; the
merus and carpus are sub equal in length, the latter being very slightly the
longer; the carpus widens slightly towards its distal end, and is equal in length to
the palm or occasionally a little longer; the palm is cylindrical, and slightly wider
than the carpus; the fingers are two thirds or more the length of the palm, and
pubescent, on the proximal half of cutting edges; there are grooves on the
fingers of the chelae with sharp cutting edges in both sexes, and one or two
minute basal teeth; the three pairs of non-chelate legs are slender, their dactylii
are simple. The pleopods are typical in structure.
130
Remarks: M. dayanum belongs to the hendersoni group first used
by Tiwari (1952). Twelve other species are also in this group, presence of
longitudinal grooves in the fingers of second cheliped and presence of soft
velvety pubescence along the grooves easily separate these species from all
other congeners.
Koshy (1973) has studied the sexual dimorphism in the species and
Khanum et al (1985) its embryonic development.
Colour: Chelipeds are prominently banded with dark brown stripes and
carapace is flesh or ash coloured, densely mottled brown, the antennae are also
banded .
Size: 44mm in TL (female)
Localities: Lahore, Sheikhupura, Rawalpindi,Sanghar, Gharo Dhand,
Rohri and Hyderabad
Distribution: India, Bangladesh, Myanmar, Nepal
Macrobrachium equidens (Dana, 1852)
(Fig.52,Pl.3D)
Palaemon sundaicus auct. Kemp, 1918:261
Palaemon equidens Dana, 1852a:26.
Palaemon sulcatus Henderson & Matthai, 1910:289,pl.16, fig.4.
Macrobrachium equidens Holthuis, 1950:162, (part) fig.36; Holthuis &
Rosa, 1965:10; Johnson, ,1962: 56; 1973:283; Kensley, 1977: 42; Kazmi &
Kazmi ,1979:152; Hwang & Yu, 1982:161; Holthuis & Miquel, 1983:7; Kazmi et
al , 1991( larvae):63; Chace & Bruce, 1993:25; Ng, 1995:74; Yeo etal, 1999:
226;
Wowor & Choy, 2001:282; Jayachandran, 2001:96; Cai & Shokita,
2006:265;Li&Bruce,2006:
Description: The rostrum is long and reaches beyond the
scaphocerite. The upper margin of the rostrum bears 11 to 12 teeth; the lower
has 5 to 6 teeth. The carapace is armed with antennal and hepatic spines; the
hepatic spine lies close behind and below the antennal. The antennal spine
131
generally continues posteriorly in a strong carina, which almost reaches the
hepatic spine.
The telson is longer than the sixth abdominal somite; it tapers towards
the posterior end. The dorsal surface is armed with two pairs of posteriorly
directed spines. A shallow longitudinal groove runs dorsally on the proximal half
of the telson.
The antennular peduncle consists of three sub equal segments; the
basal segment is the largest. The stylocerite fails to reach the middle of the
basal segment. The second segment is longer than the third or ultimate
segment. The scaphocerite reaches well beyond the antennular peduncle. It is
3.5 to 4 times as long as broad. The lateral side is more or less straight and
ends in a well-defined spine, which is shorter than the squamose portion.
G
D
F
C
132
Figure 52: Macrobrachium equidens. A- Anterior carapace; B- Telson; CAntenna; D- Third maxilliped; E- First leg; F- Second leg; G- Same, chela
enlarged; H- Third leg ( Modified from Jayachandran, 2001).
The mandibular palp is three segmented. The incisor process bears
three teeth, of which the central one is the largest. The palp of the maxillula is
apically divided into two distinct unequal lobes. The maxillary palp is slender
and finger like.
The palp of the first maxilliped is slightly longer than the caridean lobe.
The lateral side of the caridean lobe is somewhat convex. The second
maxilliped is of typical caridean type. The third maxilliped fails to reach the end
of the antennular peduncle.
The first pair of pereopods is shorter than the second pair. The fingers
are nearly half of the chela. The carpus is longer than the merus. The members
of second pair are sub equal in length, similar in form, the palm is sub
cylindrical, covered with dense pubescence, dentition is present on opposable
margins in younger specimens, the fingers do not gape in full grown males; they
measure three fourth of the palm which is pubescent in younger specimens; the
chela is longer than the carpus, the palm is shorter than the carpus. Last three
pairs of pereopods extend beyond the scaphocerite by entire length of the
dactylii; the propodus of third leg is partially pubescent
Pleopods and uropods are normal in structure
Remarks: This species lies in a less closely-knit and less easily
defined group, the species have constantly being confused with each other in
the group . There seem to be two types of specimens in same population basing
to abundance of setae, already pointed out by Pillai(1990) that the two forms
may differ in colour and morphology and can be considered as two different
species.
133
Size: 65-75mm (male), 62-74mm (female)
Colour: Brown blotches on yellow background
Habitat: Widely distributed in estuarine water including mangroves
penetrating brackish water, rarely found far inland in complete freshwater.
Locality: Sandspit
Distribution:
Indo-West Pacific: From East to South Africa to South
China, Fiji, Taiwan, Peninsular Malaysia, Bay of Bengal, Sri Lanka, S. India,
Madagascar Brunei and North east Atlantic.
Development: see Ngoc-Ho,1976
Macrobrachium idella idella (Hilgendorf, 1898)
(Fig.53 )
Palaemon (Eupalaemon) idae var. idella Hilgendorf, 1898: 29
Palaemon (Eupalaemon) cf. idae var. idella Barnard, 1950:77
Palaemon (Eupalaemon) multidens Coutiere, 1900:1266
Palaemon multidens Coutiere, 1901: 327
Palaemon idae Henderson & Matthai, 1910:285; Panikkar, 1937: 346;
Nataraj, 1942: 468; Chopra, 1943:5; Yaqoob, 1986:375.
Macrobrachium idella idella, Holthuis, 1950 (with complete synonymy);
Jayachandran & Joseph, 1985:130 ;Jalihal et al, 1988:51; Jayachandran, 2001:
112.
Description: The rostrum is long extending up to distal end of antennal
scale, the basal crest is not much raised, the proximal dorsal margin is gently
sloped over eyes and the distal margin is slightly up curved; the dorsal margin is
armed with 12-15 teeth, of which 2 are post-orbital, the first dorsal tooth is
situated slightly in front of anterior ¼ length of the carapace; second is separate
from first by a distinct space; third is closed-set to second, second to tenth teeth
are equidistant; eleventh is slightly separated from tenth and twelfth from
eleventh by distinct gaps; the twelfth and thirteenth are smaller and sub distal.
The ventral margin is provided with 4-5 teeth situated beneath level of 7-11
134
dorsal teeth, small setae are present between teeth of both dorsal and
ventral margins. The carapace is almost smooth except for a few minute
spinules in dorsal part. Both the antennal and hepatic spines are present; the
latter is situated below and behind level of former.
Figure 53: Macrobrachium idella idella. A- Carapace; B- Telson; C-Second
leg, male; C’- Same, female (Modified from Yaqoob, 1986 as M. idae).
The abdomen is glabrous. The pleurae of somites first to third are
typical that of fourth and fifth is directed backwards and sixth ends in a spine.
The telson is basally broad narrowing distally, the tip of telson extends almost to
the level of distal end of the uropod; dorsal part of telson is armed with
tubercles. Two pairs of dorsal spines are present, the first pair is at about
midlength of the telson, and the second pair is slightly above midway between
first pair and tip of telson. The posterior end of the telson is pointed with two
pairs of spines; outer pair is smaller and immovable, the inner pair is longer and
movable reaching beyond tip of telson. A few plumose setae are present
between the inner pair of spines.
The antennae are typical. The mandible has a tridentate incisor
process. The palp is 3-segmented of which the first and second segments are
equal, the third is longer. The maxillae and maxillipeds are also typical. The
distal end of second segment of endopod of the third maxillipeds reaches to the
distal end of antennal peduncle.
The first chelate legs are slender, the ischium is inflated, about 1.5
times longer than the chela but shorter than the merus and carpus; the merus is
135
shorter than the carpus but longer than the ischium and chela; the carpus is
longer than all the other segments; the palm is equal to fingers.
The second chelate legs are equal. The ischium is flat and stout,
shortest of all the segments; merus is longer than the ischium and dactylus but
shorter than the carpus, propodus and the palm; the carpus is longer, than the
chela; the palm is about 2 times longer than the fingers, the fingers are equal,
slender; movable finger has denticles and immovable finger bears 1 prominent
denticle in the proximal part of the cutting edge. In this finger there may be 2-5
very minute denticles forming a longitudinal ridge between the prominent
denticle and the proximal part of that finger. This finger is covered with small
hairs only along the cutting edge whereas the movable finger is completely
pubescent. All segments are covered with prominent tubercles, last three pairs
of legs are simple, and non-chelate, the ischii are almost equal to carpi, or
slightly smaller than the propodii; dactylii are simple.
The pleopods and uropods are typical in structure
Remarks: An elaborate description of the species was given by
Henderson & Matthai (1910) under the name Palaemon idae. Holthuis (1950)
synonymies the various names and gave the valid scientific name for the
species as Macrobrachium idella. Jayachandran & Joseph (1985) described a
new subspecies, M. idella georgi; with the result this species is now known as
M. idella idella, which shows affinities with widely distributed M. idae (Cai & Ng,
2001).
A third subspecies is suspected to occur here as one specimen (CL 31)
recently collected from Thatta (Sindh) has characters differing from the previous
two subspecies. It needs more material to verify.
Size: 60-118mm
Locality: Sajawal Bridge, Thatta
Distribution: East Africa, Madagascar, and India
Macrobrachium lamarrei korangii sub-sp. nov.
(Fig.54)
Material Examined: Holotype, male; Paratypes, 8 females 11-16 mm
in CL, 6 males 10-16mm in CL.
Description of holotype: The rostrum is long, extending slightly
beyond the antennal scale; upper rostral teeth are present at equal interval; tip
of the rostrum is slightly concave and upturned; it is shorter than the carapace;
13-15 teeth are present on the upper rostral margin and 10-12 on lower margin
136
.The carapace is smooth with antennal and hepatic spine; antennal spine is
with a short carina.
The telson is slender, posterior end is sharply pointed and reaches as
far as the margin of endopod, dorsal surface has two pairs of spines situated in
posterior half of telson, distally ending in two pairs of spines having 1:3 ratio;
few plumose setae are present on either side of tip of telson.
Figure 54: Macrobrachium lamarrei korangii. A- Rostrum and anterior part
of carapace; B- First leg; C- Second leg.
The mandibular palp is three segmented, the distal most is the longest,
other oral appendages are typical.
The first pair of legs is symmetrical and reaching beyond the apex of
antennal scale by the length of middle of palm, ischium and merus are more or
less equal in size, inner margin is provided with short plumose setae, the carpus
is longer than the merus and its distal margin is thickened bearing few long
plumose setae; palm is slightly longer than the movable finger, the proximal part
of the palm bears a triangular short bunch of plumose setae. The second pair of
pereopods is strong and reaches beyond the apex of antennal scale by half of
carpus and entire chela. Small tubercles are present all over the leg, the
ischium, carpus and merus are more and less equal in size, and the chela is
longer than the carpus and merus. The fingers and palm are equal in size; the
cutting edges of fingers bear 1-2 denticles. Last three pairs of pereopods are
slender reaching beyond the antennal scale with dactylus, inner margin of
propodus has plumose setae, which become denser in fifth pereopod and fewer
in third; dactyli of last legs are simple.
137
Pleopods are normal, endopod of fifth pleopod is smaller, and the
endopod of second pleopod have appendix masculina extending beyond the tip
of endopod and,it is hairy. The uropod is without accessory spine on exopod.
Remarks: There are two subspecies in the nominal species as M.
lamarrei lamarrei and M. lamarrei lamarroides. This third sub-species exhibits
certain characteristic features such as 1) upper margin of rostrum with 10-12
teeth of which three are post orbital, ventral margin with 5-6 teeth, all the teeth
,are placed at distal half 2) the carpus of first leg is slightly longer than the
merus, 3) ishium and merus of second leg are equal, carpus is slightly shorter
than the chela (ratio 1:3) 4) long appendix masculina of second pleopod in
male.
Etymology: The subspecies is named after the type locality- Korangi
Creek, 15 km off Karachi
Macrobrachium lamarrei lamarrei (H. Milne -Edwards,
1837)
(Fig. 55,Pl.3A)
Palaemon Lamarrei H. Milne Edwards, 1837:397
Palaemon lamarrei Kemp, 1915:265; Nath, 1937:149; Chopra & Tiwari,
1949:214; Ahmad, 1957:19; Qadri, 1960:59; Jone, 1967:337; Shakoor, 1968:2
Macrobrachium
lamarrei Holthuis, 1950:119 (see for complete
references); Siddiqui, 1976: 70; Kurian & Sebastian, 1976: 93; Kazmi & Kazmi,
1979:151; Yaqoob, 1984:4
Macrobrachium lamarrei lamarrei Jalihal et al, 1988:2; Jayachandran,
2001:128; Cai & Ng, 2002:76; Sharma &Subba,2005:31(biology)
Description: The rostrum is very long and slender, reaching to or
beyond the antennal scale by 1/3 of its length; the basal crest is elevated and
the distal end upcurved. Upper margin is armed with 6-10 teeth of which two
teeth are situated on proximal elevated part, followed by wide edentate gap and
1-2 sub-distal teeth. The wide edentate gap is generally interrupted by one
tooth; the lower margin has 6-9 teeth generally arranged below the edentate
part, small setae are present between the teeth of both upper and lower
margins. The carapace is smooth and the antennal and hepatic spines are
present.
138
Figure 55: Macrobrachium lamarrei lamarrei. A- Rostrum and anterior part
of carapace; B- Telson; C- Second leg (from Jayachandran, 2001).
The abdomen is smooth. The pleurae of first to third segments are
broadly rounded posteroventrally, the first and fifth directed backwards and that
of sixth ends in a spine.
The telson is slender posteriorly, sharply pointed and reaching as far
as the tip of uropodal endopod; the dorsal surface is armed with 2 pairs of
spines situated in its posterior half of which distal pair is closer to proximal pair;
the terminal end has 2 pairs of spines, the outer pair is smaller and immovable,
the inner pair is longer and movable overreaching tip of the telson; a pair of
plumose setae is present between inner pair of movable spines.
139
Figure 56: Macrobrachium lamarrei lamarrei. First larval stage; ALateral view of entire larva; B- Dorsal view of head; C- Rostrum
The outer lateral spine of the antennal scale does not reach as far as
the distal end. The mandible is highly developed, the palp is three segmented,
and the distal segment is 1.5 times longer than the penultimate.
The first pair of chelate legs reaches up to outer lateral spine of
antennal scale; the ischium and merus are slender, latter slightly longer than
the former. The carpus is also slender, distal end somewhat thickened and
longer than the merus; the chela is half the length of carpus; the palm is slightly
shorter than the fingers.
The second chelate leg is larger than the first. The merus is longer than
the chela but 2/3 the length of carpus. The carpus is slender, thickens distally
and about twice longer than the chelae; the palm is only slightly longer than the
fingers; the latter are equal and slender.
140
A
C
B
E
D
Figure 57: Macrobrachium lamarrei lamarrei; First larval stage. AAntennule; B- Antenna; C- Mandible; D- Maxillula; E- Maxilla
The non-chelate legs are slender; their dactylii are simple. The
appendix masculina is distinctly longer than the endopod; it is almost non-hairy,
long, and slender. Exopods of the uropods are without accessory sub apical
spine.
Remarks: The rostrum in younger specimens is short barely reaching
the tip of antennal scale. The inner spine of telson tip distinctly overreach the
telson tip in adults but in juveniles the spinous telson ends relatively shorter
than the inner pair of spines as compared to the adults.
Colour: Body fleshy coloured with brown pigments sparsely distributed
on the rostrum, carapace, and body; inner flagellae of antennules reddish
brown in colour. The eggs are yellowish green.
Size: 44-58mm
Habitat: Normally found in fresh or brackish water
Localities: Hyderabad, Khairpur, Rawalpindi, Lahore, Multan, Rahim
Yar Khan, Chenab.
Distribution:
Nepal and Myanmar.
Confined to Indo Burmese coast, India,Pakistan,
141
Development: (Figs. 56-66) Development is partially abbreviated.
The three larval stages are described in the following account, a fourth larval
stage has been given by Sharma & Subba (2005).
B
A
C
D
F
G
H
E
Figure 58: Macrobrachium lamarreii lamarreii. First larval stage A- First
maxilliped; B-Second maxilliped; C- Third maxilliped ; D-H, walking leg.
First larval stage:
Age: 24 to 48 hours. Length 0.8 to 1.4mm., (from the tip of the rostrum
to the tip of the telson) body slightly bent between abdominal segments 2 and 3
more or less transparent except for a few localized chromatophores and
pigments in cephalic appendages; dorsal side of the cephalothorax and telson,
but it is never a permanent phenomenon, rostrum straight with slight curvature
at the tip reaching nearly ¾ of the basal segment of the antennular peduncle.
Carapace smooth, ventro-lateral margin produced into a sharp spine, large and
sessile compound eyes.
Antennules: Antennular peduncle long, narrow with two flagella at the
tip, inner flagellum long in the form of long plumose seta, outer flagellum in the
form of a rectangular with two aesthetascs and one long plumose seta at the
apex.
142
Antenna: Peduncle short, unsegmented with one long no plumose
seta, exopod with 11 plumose setae at the distal half; of which 9 long and 2
short, the last seta shortest .
Mandible: Mandible with three or four large sharp teeth on the incisor
part, one tooth like structure on the molar part, no palp present.
Maxillula: Small endopod with one spine on the inner upper part, upper
lacinia larger than lower lacinia one small tooth at the apex, no setae.
Maxilla: Exopod with two small plumose setae, no seta on the endopod
or palp. Outer margin of the scaphognathite wavy and upper half bears small
plumose setae.
Maxilliped I: Base almost rectangular with one small seta on the apex,
endopod small and nearly half of the exopod and with 4 plumose setae at the
apex, exopod long with 4 plumose at the apex, the outer margin of the caridean
lobe straight
C
B
D
E
A
F
Figure 59: Macrobrachium lamarrei
Telson;B-F- Five pleopods
lamarrei. First larval stage A-
143
Maxilliped II: Endopod elongated, five segmented, the last segment
with four short and one long plumose seta and one spine, exopod long with four
plumose setae at the apex.
Maxilliped III:Slightly longer than maxilliped II, endopod 5 segmented,
one pair of spines, each at the distal part of segment 3 and 4, 5 and 6 segment
with 2 short and one long seta, exopod long but shorter than endopod, 4
plumose setae at the apex.
Pereopods: All the five pereopods in advance stage of development,
formation of the chela initiated, pereopods 1, 2, and 3 with large exopods.
Telson: Triangular, posterior margin more than 3 times broader than
the anterior margin, seven pairs of plumose setae present.
Pleopods: Pleopods also in the advance stage of development with
endo and exopods, setation and appendices absent.
Second Larval Stage:
Age: 2 to 3 days, length 0.8 to 1.4mm. (From tip of the rostrum to the
end of the telson), body slightly bent between abdominal segments 2 to 4.
Transparent, few localized chromatophores on the cephalic appendages; dorsal
side of the cephalothorax and telson, rostrum slightly curved downwards at the
tip, nearly ¾ of the basal segment of antennular peduncle, with one dorsal
spine, carapace smooth, anteroventral margin of the carapace developed,
further forward to form pterygostomial spine.
Eye: Stalked and compound.
A
144
B
C
Figure 60: Macrobrachium lamarrei lamarrei Second larval stage. ALateral view ; B- Dorsal side of head; C- Rostrum.
Antennule: Three segmented peduncle bearing two flagella at the tip,
outer flagellum provided with 3 plumose setae, inner with 2 aesthetascs on the
outer side of first segment, 3 aesthetascs and one plumose seta on second, 2
plumose setae on third segment, 3 plumose setae on the outer lateral side and
on the tip also 3 plumose setae present, similarly on the inner lateral side of the
third segment, 3 plumose setae on the second segment and 4 plumose setae
on first segment.
Antenna:
Segmented antennal flagellum, antennal scale with one
sharp tooth like spine present at the distal end on the outer margin, squamose
portion with 18 to 22 plumose setae.
Mandible: Mandible with one long, and two short very sharp teeth on
the incisor part, molar part with only two short and blunt protuberances (not
shown in figure), no palp.
Maxillula: Proximal or inner lacinia with two long and one short strong
setae, distal or upper lacinia with only one seta and palp with two lobes.
Maxilla : Many plumose setae on the outer margin of the
scaphognathite, distal and proximal setae large, distal part of the protopod with
one long and one short plumose seta and proximal part of the protopod with two
long plumose setae, at the proximal part of the exopod before the long proximal
setae, one short and one long seta are present.
145
Maxilliped I: Base oblong rectangular with one seta on the apex,
endopod with two long non-plumose setae and one short seta in the middle of
the course of the length on the inner side, exopod long with 4 long plumose
setae at the tip. Caridean lobe somewhat concave.
Maxilliped II: Endopod 5 segmented, last segment with only one long
plumose seta and three short and three long non-plumose setae, exopod long
with 4 long plumose apical setae and one non-plumose short seta.
B
E
C
F
A
D
Figure 61: Macrobrachium lamarreii lamarrei. Second larval stage. ATelson; B- Antennule; C- Antenna; D- Mandible; E- Maxillula; F-Maxilla
Maxilliped III: Five segmented, first segment with one short seta on the
inner side, third segment with two long setae, and fifth or the last segment with
three long and two short setae, exopod with four long plumose apical setae and
one long simple seta.
First and Second Pereopods: Endopods chelate, longer than
exopods, both have two setae at second, and third segments, exopods have
three long apical plumose setae and one long plumose seta.
146
Third Pereopods: Endopod long five-segmented last segment
with apical claw and with 2 short setae, fourth segment with one long and one
short seta and one seta present on the upper part of the base in between the
endo and exopod. Exopod shorter with four apical long plumose setae.
Fourth and Fifth Pereopods: Exopod not present, only endopods
present, six segmented with apical claws. Fifth segment of fourth pereopod with
three setae on the third segment only one seta, on the first segment again 1
seta. On the fourth leg last segment with 1 long and 1 short seta, fifth segment
with 3 setae, first and second segments with 1 seta on each segment.
Pleopods 1 – 5: Developed with endo and exopod, setation and
appendices showing their appearance.
Telson : Triangular, slightly concave on the posterior side, 7 pairs of
stout, long plumose setae, one additional pair of short plumose setae present at
the centre of the posterior edge (setae omitted).
Third Larval Stage:
Age: 3 to 6 days, length 1.0 to 1.8 mm. One epigastric spine behind the
base of the rostrum, articulation between last abdominal segment and telson
present, chromatophores faded.
Figure 62: Macrobrachium lamarrei lamarrei Second larval stage. A-C- 1st3rd maxillipeds; D- H- 1st- 5th pereopods
147
Antennule: Peduncle 3 segmented bearing two flagella, outer one
is two segmented and provided with two long aesthetascs, the inner with the
appearance of segmentation and with one long and one short apical spine. First
segment with 3 aesthetascs on the outer side and 7 long plumose setae,
second with 4 short and 2 long plumose setae, the third and the last segment
provided with 4 long and 2 short plumose setae. Stylocerite starts developing
Antenna: Provided with long segmented flagellum, the proximal two
segments of the flagella have two plumose setae, one on each, a sharp spine
on the outer margin of the antennal scale at the distal part.
Figure 63: Macrobrachium lamarreii lamarrei Second larval stage. A-E- 1st5th pleopods
Mandible: Two stout and one short tooth on the incisor surface and
three large stout teeth present on the molar part, no palp.
Maxillula: Palp with two lobes, the upper lacinia bears few stiff-setae,
lower lacinia smaller than the upper lacinia and bears few setae.
148
E
D
F
C
A
B
Figure 64: Macrobrachium lamarrei lamarrei Third larval stage. AMandible; B-Maxillula; C- Maxilla; D-H- 1st- 3rd maxillipeds
Maxilla:
Palp well developed upper lacinia two lobed. The
scaphognathite well developed, margin of scaphognathite bears setae, setae on
the proximal half much longer.
Maxilliped I: Palp longer than caridean lobe, the tip bears one long
seta. Outer margin of caridean lobe convex. Tip of the exopod with four setae.
Maxilliped II: Seven segmented endopod, 1 long and one short apical
seta and one more short plumose seta on the fifth segment, exopod long with
four long plumose setae at the tip and one short plumose seta before the apical
setae.
Maxilliped III: Exopod shorter than endopod. The ultimate segment
nearly equal to parapenultimate segment, penultimate segment nearly half of
the parapenultimate segment, endopod 4 segmented with apical claw like stiff
setae and two long and 3 short plumose setae, 2 plumose short setae at the
third segment and 1 plumose seta on the second segment, exopod with 4 long
terminal plumose setae and one short plumose seta before the terminals.
First and Second Pereopods: Second pereopod the longest among
pereopods. Exopod reaches middle of the merus. Finger and palm of the first
149
pereopod equal, merus and carpus equal in size. Ischium smallest, movable
finger slightly longer than palm. Carpus and merus equal, ischium slightly
smaller. Exopod reaches the end of the merus.
D
C
F
E
F
H
G
A
B
Figure 65: Macrobrachium lamarrei lamarrei. Third larval stage. A- Telson;
B- Antennule; C- Antenna; D-H- 1st - 5th pleopods.
Third Pereopod: Endopod larger than exopod. Exopod reaches to the
end of merus. Propodus largest segment. Few setae present on each segment.
150
Figure 66: Macrobrachium lamarrei lamarrei. Third larval stage. A-E- 1st 5th pereopods
Fourth and Fifth walking legs: No exopod, bearing long apical claws
and numerous setae on each segment. Merus the longest segment.
Pleopods: First pleopod with long exopod and short endopod, setose,
second to fifth pleopods with long exopods and short endopods, setation and
appendices make their clear appearance.
Telson: More elongated than uropod, articulation between telson and
abdomen distinctly visible, concave at the posterior side, the uropod provided
with six pairs of long stout plumose setae, the additional central one not
plumose, and one pair of short lateral plumose setae. Exopod and endopod
formed exoped with 17 plumose setae but endopod without any setae. uropods
developed, outer twice as long as wide as inner; 17 plumose setae, central once
more longer than outers. Inner lobe setose.
Macrobrachium malcomsonii kotreeanum Johnson,
1973
151
(Figs. 67-68; Pl. 2C)
Macrobrachium malcomsonii kotreeanum Johnson,
1973:274,279; Chace & Bruce, 1993:15
Macrobrachium malcomsonii Qureshi, 1956:362; Siddiquij,
1976:72 ;Yaqoob,1987:90(larvae)
Description: The subspecies differs from the nominal
species in having longer rostrum exceeding the antennal scale, a
low rostral crest, rostral teeth being 9-11/5 or 6, palm of second
legs being longer than the merus in both adult male and female.
Colour: Dark grey on the dorsal side as well as on the
chelipeds; ventral side light grey; antennae and inner flagellae of
antennule grey, the outer flagellae brownish, the specimens
caught from dhands are darker in colour than those caught from
river.
Figure 67: Macrobrachium macolmsonii kotreeanum. Animal in lateral
view.
Size: 151mm
Remark: It can be considered here that workers like Qureshi (1956),
Siddiqui (1976) and Yaqoob, (1987) were dealing with Johnson‟s (1973)
152
subspecies instead of the nominal species.The duration of larval stages
may differ in different experiments as was noticed in the fourth stage achieved
by Yaqoob(1987) in 12-15 days that we obtained in 5 days.
Localities: Kotri, Thatta, Keti Bunder
Distribution: Confined to Indus River after Kotri Barrage
Figure 68: Macrobrachium macolmsonii kotreeanum.A-D, Rostral variant;
E- Carapace, anterior margin; F- Telson; G- Small cheliped, chela; HSame, merus; I- Large chela, tip of fingers; J- Cutting edges of fingers.
Description of Larvae: (Figs 69-73) .Larvae of first through eight
stages are described in the following lines taken from Yaqoob (1987).
First Larval Stage:
153
It extends from one to three days after hatching. Larvae ranging
from 2.0 to 2.2 mm were examined.
The rostrum was slender, slightly curved downward at the tip, reaching
to about two third length of the basal segment of antennular peduncle. The
anteroventral margins of carapace were produced into small spines. The eyes
were large and sessile.
The antennular peduncle was long, unsegmented with two flagella at
the tip; the inner flagellum was in the form of a long plumose seta; the outer
flagellum was somewhat rectangular with three aesthetasc, one long seta, and
one short stout plumose seta at the apex.
The antennal peduncle was short and unsegmented; the flagellum was
also unsegmented with 1 long plumose and 1 short non-plumose seta at the
apex; the exopod showed 5 annulations on the distal half, one small denticle
and 7 long plumose setae on its inner edge, 2 long plumose setae and one
short non-plumose seta at the tip, 1 short non-plumose and 1 long plumose seta
on its outer margin.
154
Figure 69: Macrobrachium malcolmsonii kotreeanum First larval stage. ALateral view ; B- Antennule; C- Antenna; D- Mandibles; F- Maxilla; G- First
maxillliped; H- Third maxilliped; I- Telson ( After Yaqoob, 1987 as M.
malcolmsonii).
The mandibles were very difficult to trace and only the right mandible
was studied with certainty. The incisor had three, one of which was blunt; while
the molar had two large sharp teeth and one bristle like tooth.
The maxillule was very small, the endopod was with 1 terminal and 1
sub terminal spine; the proximal lacinia was a little bigger than the endopod with
4 terminal setae, 1 outer seta and 1 seta on its inner edge; the distal lacinia was
about twice as large as proximal with 2 large and 3 small teeth at distal end.
155
The exopod of maxilla was with two terminal, 2 lateral and 1 large
proximal plumose seta; the endopod was with one long terminal setae, 2 setae
on inner edge of basal part; the protopod was divided into three lobes, the
proximal lobe was with four setae, others were with two setae each.
The base of first maxilliped was almost round with four setae on its
margin; the endopod was finger shaped with one long terminal and two sub
terminal setae; the exopod was long with four long plumose setae at the apex, 2
small non-plumose lateral setae.
The endopods of second and third maxillipeds were four segmented,
one finely serrated claw and two setae at the apex of each; the exopods were
long with four long plumose setae at the apex and two small non-plumose
lateral setae.
Figure 70: Macrobrachium malcolmsonii kotreeanum Second larval stage.
A- Dorsal view; B- Antennule; C- Antenna, tip of flagellum; D- Mandibles;
156
E- Maxillula; F- First maxilliped, base; G- Second thoracic leg, tip; HTelson (After Yaqoob, 1987 as M. malcolmsonii).
The first and second thoracic legs were biramous buds.
The telson was triangular; its posterior margin was broad, slightly
concave and notched in the middle with seven pairs of plumose setae and many
small stout spines, the first and second pairs were plumose on inner edges only.
Second Larval Stage:
It extends from three to nine days from hatching. The larvae were
measuring from 2.3 to 2.4 mm.
The antero-ventral margins of carapace were directed forwards to from
the pterygostomian spines. Two prominent supra-orbital spines were present
and the eyes were large and stalked.
The antennular peduncle was two-segmented. The antennal flagellum
carried one long plumose, two medium and one short seta at the apex.
H
A
B
G
C
E
D
F
157
Figure 71: Macrobrachium malcolmsonii kotreeanum. Third larval
stage. A- Antenna; B- Telson and uropod; C- Fourth larval stage, telson
and uropod; D- Fifth larval stage, telson; Sixth larval stage; E- Antennule;
F- Telson.(After Yaqoob, 1987 as M. malcolmsonii); G-H – Five days
larva,entire larva and eye.
The mandibles were asymmetrical, the right mandible was with four
stout teeth on the incisor, one of which was serrated on its inner margin, the
molar region was with five short teeth; the left mandible was with four teeth on
the incisor, the outer of which was very long and bifurcated at its tip, the molar
had six short teeth.
The endopod of maxillule was with one long and one short spine at the
apex; the proximal lacinia was with three setae at the apex, 1 seta each on
outer and inner margin; the distal lacinia was with four large and three small
teeth.
The exopod of maxilla was with two terminal, 4 lateral and 1 proximal
plumose seta.
The base of first maxilliped was swollen with eight setae; the endopod
was short with three terminal setae and one lateral seta on each side.
The first and second thoracic legs were biramous, each with fivesegmented endopod ending in a long curved smooth spine; the exopod was
with four long plumose setae at apex. Small biramous buds of third thoracic legs
were present.
The fifth abdominal segment had lateral pair of spines.
The telson had an additional (8th) pair of short non-plumose setae at
centre of posterior margin. The second pair of setae was plumose on both
edges. Small spines on posterior margin remain as in the first stage.
Third Larval Stage:
It lasts from 6 to 12 days post hatching reaching a size of 2.5 – 2.6 mm.
One (rarely two) epigastric spine was present behind the base of the
rostrum. The peduncle of antennule had three segments. The antennal
flagellum was also three segmented, the exopod was with 12 long plumose
setae, and the annulations were confined to the apex.
158
Figure 72: Macrobrachium malcolmsonii kotreeanum Seventh larval stage.
A- Antennule; B- First maxilliped; C- Second thoracic leg, chela; DTelson; Eighth larval stage; E- Rostrum; F- Antennule; G- Antenna; HTelson (After Yaqoob, 1987 as M. malcolmsonii)
The exopod of maxilla had eight plumose setae. The third thoracic leg
was similar to the first and the second legs. The fourth and fifth thoracic legs
159
were present as buds and the fifth and sixth abdominal segments had
lateral spines.
The articulation between sixth abdominal segment and telson had
formed. The telson had eight pairs of setae. First and eighth pairs of setae were
not plumose. Small spines on posterior margin remained as in the first stage.
Uropod was present; the exopod was more than twice as long as the endopod
with six long plumose setae on distal portion; the endopod was without any seta.
Fourth Larval Stage:
It takes 12-15 days post-hatching, ranging in length from 2.8 to 2.9 mm.
The carapace had two epigastric spines behind the base of the
rostrum; each spine was with few teeth on its front edge. The antennal exopod
was with seven plumose setae and one apical spine; no annulations seen.
The exopod of maxilla was with 9-10 plumose setae, the endopod and
lobes were with 1+2+3+3+2+3 setae, respectively.
The fourth thoracic leg was still a bud. The fifth thoracic leg was large
and segmented with very long curved terminal spine; the exopod was not
present.
The telson was much narrow; it bears three pairs of lateral spines and
five pairs of setae on posterior margin, three pairs of setae next to central pair
were plumose. Small spines on posterior margin were present. The exopod and
endopod of uropod were articulated with the protopod. The exopod was with 1011 plumose setae and one spine on its margin, 3-5 plumose setae were present
on its dorsal surface; the endopod was with 3+6 marginal plumose setae, while
two plumose setae were present on its dorsal surface.
Fifth Larval Stage:
It lasts from 15 to 33 days from hatching and ranges from 3.4 to 3.5
mm in length. The carapace bears 2 (or rarely 3) epigastric spines, each spine
is with few teeth on its front edge.
The proximal end of exopod of maxilla bears 1 long and 2 small
backwardly directed plumose setae, the anterior portion has eight plumose
setae. All thoracic legs are segmented; the fifth leg is the largest.
160
The telson is rectangular with 3 pairs of lateral spines and 5 pairs of
setae on posterior margin, 3 pairs of setae next to central pair are plumose,
small spines on posterior margin are present. The exopod of uropod bears one
plumose seta on its outer margin, 1 spine and 11-12 marginal plumose setae
and 4-5 plumose setae are present on its dorsal surface. The endopod is with
four plumose setae on its outer margin, 10-11 plumose marginal setae, and 2-3
plumose setae on its dorsal surface.
Sixth Larval Stage:
It extends from 19 to 38 days from hatching measuring 3.6 to 6.0 mm.
Three to four hairs are present on dorsal surface of carapace between
first and second epigastric spines. In one specimen, a small spine arises from
the base of the first epigastric spine; it is serrated on front edge. One to 3
plumose setae are present in front of second epigastric spine; each epigastric
spine is with six teeth on front edge. An antennal spine is present.
The inner flagellum of antennules is elongated, as long as the outer
flagellum and with three long setae at the apex; the outer flagellum is faintly
segmented and bears seven aesthetascs in two groups of three and four
proximal to distal.
The antennal flagellum is as long as the scale, the latter bears 18 (in
early larvae) to 28 (in late larvae) plumose setae on its edge.
The exopod of maxilla bears 13 to 18 plumose setae on its anterior
portion and 8 plumose setae on its proximal part.
The endopod of first maxilliped is indistinctly 2-segmented with
two long terminal and one sub-terminal seta on each side.
The first to fifth abdominal segments have pleopod buds. They are
prominent on second to fourth segments but become biramous as development
progresses.
The telson is much elongated with 3 pairs of lateral spines and 5 pairs
of setae on posterior margin; only 2 pairs of setae next to central pair are
plumose (in late larvae, 4 pairs of setae on posterior margin are present, the
inner two pairs are plumose). The exopod of uropod has two plumose setae on
its outer margin proximally, 4 to 5 setae on its outer margin distally, 1 spine and
22 to 24 plumose marginal setae and few setae are present dorsally; the
161
endopod has six plumose setae on its outer margin proximally, 19 to 20
plumose marginal setae and few setae are present dorsally.
Seventh Larval Stage:
It takes from 28 to 49 days from hatching ranging from 6.8 to 8.0mm in
length. Three to 4 plumose setae are present in front of second epigastric spine.
The inner flagellum of antennule is 3 to 4 segmented; the outer flagellum is split
longitudinally, the inner part is faintly segmented with 12 to 13 aesthetasc in four
groups as 3-4+two+3+4; the outer part is longer, 2-segmented and form
flagellum proper.
The exopod of maxilla bears about 49 plumose setae around its edge.
The endopod of first maxilliped is indistinctly 2-segmented with one short and
two long terminal setae.
The first and second thoracic legs are chelate. All five pairs of pleopods
are biramous, the exopods and endopods of both with plumose setae.
The telson is narrow and longer, three pairs of short lateral spines and
four posterior pairs of setae, the inner two pairs of setae are plumose. Some
times a central non-plumose setae is present.
Eighth (Last) Larval Stage:
It lasts 43-63 days from hatching and is 7.0 – 8.5 mm long.
The rostrum is with 4 to 9 teeth on upper margin, 4 to 5
plumose setae are present in front of second epigastric spine. The inner and
outer flagella of antennule are about equal in length; both are 5 to 6 segmented,
shorter outer flagellum is with about 17 aesthetes in seven groups as
2+2+2+2+2+3 and 4 terminal aesthtetasc.
The antennal flagellum is about 1.5 times as long as the scale and
consists of nine segments, the first of which is very long. The endopod of
maxillule is with two setae; the proximal lacinia is with 9 to 10 setae at apex and
one seta on its inner edge; the distal lacinia is with 7 to 8 long, 3 to 4 short teeth,
and its distal end.
The exopod of maxilla is with about 50 plumose setae around its edge;
the distal and median lobes of the protopod are well developed and with
plumose setae and spine; the proximal lobe is reduced. The basal endite of first
maxilliped is with 20 slender spines, the endopod is indistinctly 2-segmented;
162
the exopod is with 11 plumose setae on the outer edge of the enlarged
basal half. The fifth thoracic leg is the largest.
The telson is very much elongated; its posterior portion is narrow and
bears five pairs of setae, the inner two of which are plumose. The position of
first to lateral pairs of spines is not clear, while the third lateral pairs of spines is
merged with the posterior setae. The exopod of uropod is with three plumose
setae on its outer margin proximally.
Macrobrachium malcolmsonii malcolmsonii (H. Milne
Edwards, 1844)
(Fig .73)
Palaemon Malcolmsonii H.M.Edwards, 1844:8, pl.21
Palaemon choprai Tiwari, 1949:333
Marcobrachium malcolmsonii, Holthuis, 1950:121 (part); Patwardhan,
1937: 5,fig 1-65; Qureshi,1956:362;Ahmed,1957:25; Holthuis & Rosa, 1965;
Jones 1967:337; Rajyalakshmi & Randhir, 1969:903; Siddiqui, 1976: 72; Kurian
& Sebastian, 1976:223:11, fig. 21,pl.vii; Holthuis, 1980:99; Holthuis & Miquel,
1983 unpaginated; Jayachandran, 2001:141
Description: The rostrum is long, extending to or short of antennal
scale, it is only slightly broad, the basal crest is elevated, the distal width is
narrow and upcruved. The upper margin is provided with 11 teeth of which three
are postorbital. The proximal group
of irregularly placed ten teeth is
characteristic , the eleventh is slightly separated from the tenth, the twelfth is
widely separated leaving a distinct gap between eleventh and twelfth teeth, the
latter is at a level just in front of the antennal scale; the ventral margin is
provided with six teeth all equidistant; the first ventral tooth is located at a level
just in front of the ninth dorsal tooth, the third ventral tooth is located at level of
middle antennular peduncle segment; fourth to fifth ventral teeth are below the
distinct dorsal gap; sixth ventral tooth at the level of last dorsal tooth; small
setae are present between the teeth at both of dorsal and ventral margins.
The carapace is smooth, antennal and hepatic spines are present; the
latter is situated below and behind level of the former.
163
The abdomen is glabrous; the pleurae of first to third segment are
broadly rounded that of fourth and fifth are directed backwards that of the sixth
is ending in a spine.
The telson is basally broad, narrowing distally, ending in a spine; the
dorsal margin has two pairs of spines; the first pair is located at about middle of
the telson, the second pair is more close to the first pair than the distal end; the
posterior margin is also provided with two pairs of spines; the outer pair is
smaller and immovable while the inner pair is longer and movable, overreaching
the tip of telson extend beyond the inner spine; a few plumose setae are present
between the inner pair of movable spines.
Figure 73: Macrobrachium malcolmsonii malcolmsonii. A- Anterior
carapace; B- Telson ;C- First leg; D- Second leg; E- Part of second chela
enlarged (from Jayachandran, 2001)
The antennae are typical. The mandibular palp is three segmented; the
middle segment is the smallest. The maxillae and maxillipeds are also typical;
the second segment of endopod of third maxilliped reaches the level of antennal
peduncle;
The first pair of chelate legs is slender; the ischium is inflated, two third
lengths of the merus; the merus is shorter than the carpus; the carpus is long,
slender, and equal to the combined length of chela and ischium; the palm is
almost equal to the fingers. The second pair of pereopods is stronger sub equal;
164
the ischium is stouter, flattened, shorter than the merus, carpus and palm
but almost equal to the dactylus; the merus is shorter than the carpus and palm;
the carpus is shorter than the chela but longer than the palm; the palm is
cylindrical and slightly inflated; the fingers are slender, the movable finger is
armed with two prominent denticles and is covered with pubescent hairs, not so
in females, the immovable finger has one prominent and two smaller denticles;
all segments are tuberculate.
The pair of nonchelate legs is simple; ischium is equal to the carpus;
the dactylii are simple.
The pleopods and uropods are typical in structure.
Remarks: Dutt & Ravindranath (1976) and then Rao etal (1992)
observed differential growth rate in the species. The former author found a
distinct change in the rate of growth of the inner spine in relation to the spinous
end of the telson from juvenile through adult, in juveniles spines may be well
beyond the telson tip.
Localities: D.I. Khan, Nowshera
Distribution:
Indo-West Pacific; India, Myanmar, Bangladesh.
Macrobrachium naso (Kemp, 1918)
(Fig. 74)
Palaemon naso Kemp, 1918:19
Macrobrachium
naso Holthuis,
Jayachandran, 2001:148; Cai & Ng, 2002:63
1950:136;
Siddiqui,
1976:70;
Description: The rostrum is very long, slender, extends beyond the
apex of the antennal scale by about 1/3 of its length, in smaller specimens, the
rostrum is proportionately smaller, reaches beyond the scale by about ¼ its
length. The upper margin has 8-11 teeth of which generally three teeth are
situated on the carapace behind the orbit, in some specimens, the third tooth is
immediately above the orbital angle. The rostrum begins as a dorsal crest a little
in front of the midpoint of the carapace, the proximal half is straight and the
distal half is up curved, teeth on the proximal half are more close-set, those in
the distal half widely separated. One or two teeth are located sub distally in the
distal part and hence the distal end has a bifid or even trifid appearance; the
lower margin has 5-8 teeth, all of them equidistant. There is no well-marked
lateral keel. Small setae are present between teeth of both the upper and lower
margins.
165
Figure 74: Macrobrachium naso. A- Anterior carapace; B- Telson; C- First
leg; D- Second leg (Modified from Jayachandran, 2001)
The carapace is smooth and with usual antennal and hepatic spines,
the latter is situated below and behind the level of the former.
The abdomen is also smooth, pleurae of first to third somites are
broadly rounded along the posteroventral margin, that of fourth and fifth is
directed backwards and of sixth ends in a sharp point.
The telson is slender; the dorsal surface has two pairs of spines. The
anterior pair is situated slightly anterior to middle of the telson, posterior pair is
closer to anterior pair than to the distal end of the telson, and distal end
terminates in very sharp point flanked by two pairs of spines on either side.
Outer pair is smaller and immovable; the inner pair is very long and movable,
reaches much beyond apex of the telson, a few plumose setae are present
between the inner pair of the movable spines.
The eyes are short and broad.
The basal segment of the antennular peduncle is broad; its
anterolateral margin ends in a sharp spine, which extends to about middle of the
second segment of peduncle. The second and third segments are equal. The
upper antennular flagellum is distally bifid, its basal part is fused for about 8-9
joints. The outer lateral margin of antennal scale is almost straight.
166
The mandibular palp is three segmented. The end pod of the third
maxilliped reaches almost to the end of second segment of the antennular
peduncle.
The first pair of chelate legs are slender extend up to the end of
antennal scale; the ischium is smaller and inflated; The merus is slightly more
than ¾ length of carpus; The carpus is about 2.5 times longer than the chela;
the palm is equal to the fingers, latter are beset with tufts of setae.
The second pair of chelate legs are equal, smooth and slender and do
not differ in the two sexes. They are never more than half the total body length.
In full-grown animals, pereopods extend beyond the antennal scale by chela
and sometimes a small part of the carpus also; the ischium is the shortest
segment sub equal to the merus; the merus is shorter than the carpus; the
carpus is distinctly shorter than the chela; the palm is generally a little longer
than the fingers. Chela is slightly curved and fingers show no trace of denticles
on cutting edges.
The non-chelate pereopods are slender, their dactylii are simple.
The pleopods are normal in structure. Appendix masculina is fully
developed in small and large specimens.
Remarks: Cai & Ng (2002) commented on M. naso as endemic to its
types locality i.e. Myanmar. Most probably, they must have missed the record
from Pakistan by Siddiqui (1976). and ignored Indias|” record by Jayachandran.
The present record is being made following Siddiqui (1976). Here also is
another probability, Siddiqui, (1976) could have mistakenly taken M. dayanum
for M. naso, the two being very close. We are unable to make any further
comments due to the lack of specimens. The above description is based on
Jayachandran (2001).
Size: 72mm (male), 57mm (female)
Colour: General colour grey. Rostrum dark grey. Antennae and
antennules reddish. Fingers of chelae tinged red: a small grey spot at each side
near the upper limits of the lateral surface on a level with base of the rostrum.
An irregular grey bar sloping backwards and downwards from the lower orbital
margin; another still more irregular and broader, parallel to it a short distance
posteriorly; a backwardly directed lunate mark of the same shade about the
same distance behind the second bar, and finally a third irregular bar directed
straight downwards partly within the lunate mark, a short distance in front of the
posterior margin of the thorax. A grey spot on each side of the posterior margin
of each abdominal segment and above it a bar of same colour extending across
167
the dorsal surface. Telson irregularly cross-barred; uropods mottled or
clouded (Kemp, 1918).
Habitat: Occurs in riffle areas of streams and in flowing sections with
cobbles/ boulders substrate.
Distribution:
India, Myanmar.
Macrobrachium rude (Heller, 1862)
(Fig.75)
Palaemon rudis Heller, 1862: 527; 1865:114; Henderson & Mathai,
1910:291;Kemp, 1915:268; Menon, 1938:288; Chopra, 1939:223; 1943: 4.
Palaemon mossambicus Hilgendorf, 1898:29;
Palaemon ( Eupalaemon) sundaicus Hilgendorf, 1898:30;
Palaemon (Eupalaemon) Alcocki Nobil, 1903:9
Urocaridella borradaillei Stebbing, 1923:8
Palaemon (Eupalaemon) cf. sundaicus Barnard, 1950:775
Palaemon (Eupalaemon) cf. idea var. idella Barnard, 1950:777
Macrobrachium
Jayachandran, 2001:161
rude
Holthius, 1950:150; Johnson,
1973:285;
Macrobrachium equidens partim Holthius, 1950:162.
Description: The rostrum is short extending almost to the tip of
antennular peduncle; the dorsal margin is convex; the basal crest is not
elevated and apex is pointed forward; the dorsal margin has 10-11 of which 2
are post orbital; first tooth is at about anterior one fourth of carapace; first to
ninth teeth are almost equal and equidistant; two distal teeth are smaller; the
ventral margin has five teeth: the first ventral tooth is at level of fifth dorsal tooth;
the second is in front of sixth, third is in front of seventh, the fourth is at level of
eighth and the fifth is at level of ninth.
The carapace is scabrous due to presence of minute
inconspicuous prickles; its length is about twice that of rostrum. Both antennal
and hepatic spines are present; the latter is situated below and behind level of
former.
168
The abdomen is glabrous; the pleurae of first to third
segments are broadly rounded at posterolateral margin, those of fourth and fifth
are directed backwards and that of sixth is spinous.
The telson is sharply pointed, the distal end reaches beyond the level
of outer spine of uropodal exopod, the dorsal margin has two pairs of spines:
the anterior pair is at midlenght, the posterior pair is midway between anterior
pair and tip of telson; distal end is provided with two pairs of spines: outer pair is
smaller and immovable, inner pair is longer and movable overreaching the tip of
telson. A few plumose setae are present between the inner pair of movable
spines.
The mandibular palp is three segmented; the basal two
segments are equal. Other appendages are typical in structure.
Figure 75: Macrobrachium rude. A- Anterior carapace; B- Second leg; CThird leg (from Jayachandran, 2001)
The first chelate legs are slender; the ischium is slightly
inflated, longer than the propodus; the merus is slender, shorter than the carpus
reaching almost to the level of antennal flagellar peduncle; the carpus is the
longest segment; the palm is equal to the fingers. The second chelate legs are
longer and stronger than the first, more than 1.5 times longer than the total body
length; the ischium is the shortest segment; the merus is robust, longer than the
169
dactylus but shorter than the carpus and palm; the carpus is shorter than
the propodus but longer than the palm; the propodus is the longest segment, the
palm is cylindrical; the fingers are slender, equal, shorter than the palm; the
fixed finger is provided with one proximal denticle, the movable finger with two.
All the segments are entirely velvety pubescent.
Nonchelate legs are simple, almost equal; the ischii are equal to carpii,
the merii are almost equal to or slightly longer than the propodii, the dactylii are
simple.
The pleopods and uropods are typical in structure
Colour: Upper side of body greyish or of flesh colour; second legs grey
on the dorsal and lateral sides and bluish on the ventral sides; antennae grey;
under sides of the body flesh coloured.
Size: Males are larger than females measuring 130mm in TL
Locality: Sindh waters
Distribution: East Africa, Madagascar, India, Sri-Lanka
Macrobrachium scabriculum (Heller, 1862)
(Fig.76)
Palaemon scabriculus Heller, 1862:527; Henderson,
1893:442;Kemp,1915:272; Panikkar, 1937:346; Shakoor, 1968:3
Palaemon (s.s) Dolichodactylus Hilgendorf, 1879:840; Barnard,
1950:772
Palaemon (Paralaemon) Scabriculus Nobili, 1900; 1903
Palaemon dubius Henderson & Matthai, 1910:300
Macrobrachium
scabriculum (part) Holthuis, 1950:224;
Chace
&Bruce, 1993:37; Yeo et al., 1999:235; Jayachandran, 2001:165; Wower &
Choy, 2001:286; Cai & Ng, 2002: 75 (discussion);Cai & Shokita, 2006:266.
Description: The rostrum is very short extends only to the tip of
antennular peduncle. The upper margin in convex, distal end points downwards,
provided with 12-15 teeth, of which five are postorbital. The teeth are
170
equidistant, the ventral margin has one- three teeth, and small setae are
present between teeth of both dorsal and ventral margins.
Figure 76: Macrobrachium scabriculum. A- Anterior carapace; B- Large
second; C- Small second leg; D- Same, chela enlarged (from
Jayachandran, 2001)
The carapace is scabrous, minute spinules are present on the surface.
The telson is robust, extending almost to same level as outer lateral
spine of the uropodal exopod. The dorsal surface is provided with two pairs of
spines, of which proximal pair is situated about the midlenght of the telson, the
distal pair is situated midway between the proximal pair, and the distal end of
telson. The distal end also has two pairs of spines; the outer pair is smaller and
immovable, the inner pair is longer and movable, overreaching the tip of telson.
A few plumose setae are present between inner pair of spines.
The first chelate legs are typical in structure; the ischium is inflated,
slightly shorter than the merus, the merus is shorter than the carpus, which is
slightly more than twice the length of chela, the palm is slightly inflated and
equal to the fingers.
The second chelate legs are strongly unequal; either the right or the
left is longer, the ischium of the larger cheliped is shorter than the merus; the
171
merus is almost equal to the carpus; the carpus is almost equal to the palm
or slightly longer; the fingers are slender and distinctly longer than the palm;
entire cutting edge of finger has more than 20 denticles. Entire palm and basal
part of fingers are covered with thick pubescence. The ischium of smaller
cheliped is smaller than the merus; the merus is slightly smaller than the carpus;
the carpus is smaller than the chela but distinctly longer than the palm; the palm
is shorter than the fingers; the fingers are twice the length of the palm; the
cutting edges of the fingers have 5-6 small denticles, present in the proximal
part only. Stiff setae and spines are present throughout surface of second pair
of chelipeds, others segments are spinulose.
The three pairs of non-chelate legs progressively increase in length
from first to third, the ischii in these legs are almost equal to the carpii or slightly
shorter; the merii are longer than the propodii, the dactylii are simple.
The pleopods and uropods are typical in structure.
Remarks: M. scabriculum given by Jalihal et al (1988) has been
combined with M. lanatum, a new species established by Cai & Ng (2002).
Size: 48mm in TL
Colour: Flesh colour with grayish markings and cross bands on the
dorsal side of body.
Habitat: lives among weeds growing in back water and ascends to river
and streams or in caves.
Localities: Kotri, River Indus, Kalmat
Distribution: Juba, S. Italian Somaliland; “ Region des grands lacs”;
Zanzibar; Mozambique; Madagascar, India, Sri Lanka, Sumatra, Borneo,
Philippines, Peninsular Malayasia, Singapore, Brunei.
Macrobrachium shahpuri sp.nov
(Fig. 77)
Material Examined.: Holotype berried female C.L. 12mm; Paratypes
nine females 12-14mm C.L, date 12, June 2008
Type Localities: Canal from Shah pure Dam, Rainwater reservoir,
Wah stream and Fateh Ganj
172
Description of the Holotype: The rostrum is as long as the
carapace, extending much beyond the antennal scale. It is narrow with a slightly
upturned tip. The upper margin is armed with 6-9 teeth in which two are
postorbital. A large, smooth sub distal gap is present on upper margin between
apical (one or 2) and large and evenly spaced proximal teeth, slightly convex,
the lower margin is armed with 5-8 teeth. The carapace is smooth and armed
with antennal and hepatic spines.
Figure 77: Macrobrachium shahpuri. A- Animal in lateral view; A’- Rostrum; B- Eye; CD- Telson and uropod; E- Maxillula; F- Antennule; G- Antenna; H- First leg; I- Second
leg; J- Third leg; K- Fifth leg.
The eyes are well developed; cornea is broad and well pigmented.
The telson is slightly longer than the sixth abdominal somite and bears
two pairs of posteriorly directed spines in the posterior half, posterior margin
ends in a triangular median point, flanked by two pairs of spines of which inner
pair is stouter and at least 4.5 times longer than the outer, 2 pairs of short
plumose setae are ventrally, between the inner spine and one pair of small hairs
is present dorsally.
173
The stylocerite is reduced and anterolateral spine of basal segment
of antennular peduncle is slender and reaches up to the middle of second
segment. The scaphocerite is 2.25 times as long as broad and spine reaches up
to fourth tooth of lower margin of rostrum.
The oral appendages are normal. The mandibular palp is three
segmented; the third segment is twice the length of the second. The maxillulary
palp is as illustrated. The third maxilliped is reaching up to the middle of the
antennal scale.
The first pereopods are slender, equal and extend up to the tip of the
antennal scale. The fingers are as long as the palm. The carpus is more than
twice the length of chela. The second chelate legs are longer than the first. They
are slender, equal and similar in both the sexes and extending up to the tip are
even beyond the antennal scale with chela. The fingers are shorter than the
palm having a small proximal gap when closed and with delicate hairs on the
cutting edges, which are sharp but smooth; the chelae are always longer than
half times of the carpus, the carpus is longer than the merus. Last three pairs of
pereopods are similar in structure with simple dactylii.
The appendix masculina is shorter than the appendix interna in males
and provided with 3-4 stiffer setae terminally.
The uropods are longer than the telson and are characterized by the
absence of accessory sub apical spine on exopod. The exopod is longer than
the endopod.
The eggs are rather large, oval, or elliptical.
Remarks: Macrobrachium shahpuri sp.nov. lies in M. lamarrei group
and have a close resemblance with M. canarae, Tiwari, and M. sankollii Jalihal
& Shenoy. M. shahpuri and M. canarae can be easily separated from M.
sankollii by the presence of accessory apical spine on the exopod of the uropod.
In M. canarae, the rostrum is 1-1.75 as long as carapace where as in M.
shahpuri the rostrum is as long as carapace. The rostral formula of M. canarae
is 7-10/4-7 and can be distinguished from M. shahpuri where the rostral formula
is 6-9/5-8. The stylocerite of M. shahpuri is short, have 1:4 ratio with basal
segment whereas in M. canarae it is large, and have 1:3 ratios. The palp of
maxillula of M. shahpuri is quite different from that of M.canarae and M. sankolli.
In M. shahpuri the appendix masculina is shorter than the appendix interna and
provided with 3-4 stiffer setae terminally, whereas in M. canarae the appendix
masculina is 1.8 to 2 times as long as appendix interna.
174
Etymology: M. shahpuri is named after the location of sampling Shahpur Dam, Pakistan.
Macrobrachium taunsii sp.nov.
(Figs. 78-79,Pl.2D)
Macrobrachium sp Kazmi & Kazmi, 2006 (abstract only)
Material examined Holotype male 70mm in TL; Paratypes 3males 5076mm in TL, 1 female 65mm in TL. Muzaffar Garh, Zoological Survey of
Pakistan Reg. No. 1012; several specimens from stomach of Indus dolphin
collected at Taunsa, courtesy of WWF Pakistan, dt. 14-01-2005.
Type locality: Taunsa barrage, Muzaffar Garh canal, Punjab
Description of holotype: The rostrum is lanceolate in profile, being
highest just before the middle, with the tip is acutely pointed and feebly up
curved. The rostrum is almost horizontal, reaching up to or slightly beyond the
third antennular segment, but fails to reach the end of spine of scaphocerite.The
upper margin of the rostrum is armed with ten sub equal teeth of which two are
postorbital, the lower margin bears two teeth in distal half between eighth and
ninth tooth of upper margin. The lateral carina is distinct.
The carapace is entirely glabrous, and granular anterolateral region
bears large number of spinules behind the antennal spines. The hepatic spine is
much smaller than the antennal spine and is placed on a lower level.
The pleura of all the abdominal somites are smooth.
175
Figure 78: Macrobrachium taunsii. A- Anterior region, lateral view; BTelson; C- Second cheliped, chela and carpus, dorsal view; C’ Same,
ischium and merus; D- First leg; E- Third leg; F- Second pleopod; G- Outer
margin of exopod of uropod.
The telson is elongate, slender much longer than the sixth abdominal
somite. The dorsal surface bears two pairs of minute spines on the posterior half
of telson, the anterior pair is somewhat behind the middle, and the posterior pair
is at equal distance to the anterior pair and to the posterior end of the telson.
The posterior margin ends in a median process, which is flanked, by two pairs of
small spines; the inner pair is much stouter and longer than the outer, the latter
overreaches the tip of telson, few plumose setae are present in between the
inner pair of spines.
176
The eyes are typical. The peduncle is short, with globular and
reniform cornea. The ocellus is rather large.
The basal segment of the antennular peduncle is broad. The lateral
margin is slightly convex, its anterolateral angle being broadly expanded
forwards; it is rounded with a terminal tooth, which extends to the middle of the
second segments. A slender stylocerite is present failing to reach up to the
middle of first antennular peduncle. The distal two segments are sub equal in
length.
The antennal scale is broad, about two and a half times as long as the
maximum breadth. The lateral margin is straight tipping with a strong final tooth;
the anterior margin of the lamella is broadly expanded and more or less
produced anterointernally, far overreaching the lateral tooth. The basicerite of
the antennal peduncle is armed with a strong external tooth. The carpocerite is
broad and short.
Figure 79: Macrobrachium taunsii. A- Mandible; B- Maxillula; C- Maxilla; DF- First to third maxillipeds; G- Second right cheliped, chela and carpus,
ventral view; G’- Same, ischium and merus
177
The mouthparts are normal. The palp of the maxillula is distinctly
divided into two lobes; the outer lobe is narrower and setose, and the inner lobe
is with a seta distally. The endite of the maxilla is well developed and deeply
cleft. A finger like palp is present. The scaphognathite is rather elongate. The
mandible is three segmented; the distal segment is the longest, while the first
segment is the shortest. The first maxilliped has the endite separated into coxa
and basis by a distinct notch. The exopod is well developed with a narrow
caridean lobe; the epipod is broad and distinctly bilobed. The third maxilliped is
broad and pediform; the ultimate segment is a little shorter than the penultimate
with a curved claw terminally; the basal segment is twice the length of the
penultimate segment. The moderately developed exopod fails to reach the distal
end of the basal segment.
The first pereopods are equal, slender reaching beyond the antennular
peduncle by half of carpus, when extended; the carpus is longer than the
merus,both the fingers are equal, the palm is shorter than the ischium; movable
finger is curved distally to a pointed end,crossing over to immovable finger; the
cutting edges of the fingers are entire, without any tooth.
The second pereopods are equal, extending beyond the antennular
peduncle by chela when extended, and overreaching the scaphocerite by four
fifth of the length of the carpus; the immovable fingers bears 7-8 teeth whereas
the movable finger 6-7 teeth, the fingers are shorter than the palm, the tips of
the fingers are curved and crossed when closed; the carpus is slightly shorter
than the immovable finger; the palm is longer than the merus; ischium and
carpus are more or less equal. All segments are spinulose and pubescent
except distal part of the fingers.
The last three pereopods are similar in structure. The dactyli are
slender and curved. The propodus carries a row of both single and paired
spines at the posterior border. The ischium of the third leg is equal to carpus;
the merus is the longest segment, and longer than the carpus and ischium.
The second pleopod has a strong appendix masculina extending to the
middle of endopod, having stiff setae along the whole length; it is much longer
than the slender appendix interna.
The uropods are typical. The exopod has an extra movable spine.
Remarks: This palaemonid is of robust form showing a close
resemblance with Macrobrachium yui Holthuis, M. thai, M. roridens (Hilgendorf)
and M. mieni Dang, but it can be distinguished by the following characters 1)
size of the rostrum and rostral teeth of lower margin 2) finger of first legs being
shorter than the palm; 3)the palm of second leg larger than the fingers 4) palm
and fingers of first and second legs being more are less equal in size 5) more
178
tubercles present on third leg and its dactylus being large and curved in
wards and larger than the carpus.
Etymology: The species is named after the collecting site: Taunsa
Barrage.
Macrobrachium tirmiziae Yaqoob & Kazmi, 1987*
Material Examined: Holotype male 35mm in TL. Four paratypes,
Islamabad, date of collection 1986-87.
Type Locality: Islamabad, Rawal Dam.
Description: The rostrum is almost straight and slightly longer than the
antennal scale. The first (epigastric) and the second rostral teeth are placed
behind the orbital margin, five equidistant teeth on the rostrum and two teeth are
present near the tip of the rostrum. On the lower margin, four teeth are
equidistantly placed and slightly decrease in size towards the apex.
The carapace is smooth and with an antennal and hepatic spine.
The tip of the telson has two pairs of lateral spines in addition to a
central spine. Twelve long plumose setae (only eight illustrated) are present on
posterior margin of telson and four hairs are present on the dorso-posterior
region. Two pairs of spines are present dorsally on the telson.
The first pair of legs is as long as scaphocerite while the second pair of
legs is sub equal, moderately stout and exceed the rostrum. The ischium of the
second leg is shorter than the merus, which is shorter than the carpus; the
carpus widens slightly towards the distal end and is longer than the palm. The
palm is somewhat swollen. The fingers are shorter than the palm, with two
minute basal teeth on fixed and one on the movable finger. The ambulatory legs
are slender and setose.
Remarks: The rostrum in type series of M. trimiziae studied varies from
being in almost straight, as long as or slightly shorter or longer than the antennal
scales. Long plumose setae (8-14) are present around the posterior margin of
the telson. The new species resembles M. divakarani Jayachandran in the
number of rostral teeth and their distribution and the ratio of ischium, carpus
merus and palm but differs from it in the shorter rostral length and having an
unalleviated basal rostral crest.
179
Etymology: The name of this new species has been designated in the
honour of Dr. (Mrs.) N.M. Tirmizi, Professor of Zoology University of Karachi,
and the PhD supervisor of the authors..
* As cited the authors for this species are Yaqoob & Kazmi and the description of this
species is the responsibility of these two authors
180
Genus Nematopalaemon Holthuis, 1950
Diagnosis: Rostrum elongated, with elevated basal crest; carapace
with antennal and marginal branchiostegal spine, without branchiostegal suture;
mandible with palp; 3 posterior pairs of pereopods with dactylii simple, not
biunguiculate, longer thread-like, longer than propodus; first pleopod of male
without appendix interna on endopod (from Jayachandran, 2001)
Type species: Leander tenuipes Henderson, 1893
Gender: Masculine
Remarks: The genus consists of five species, of these only one i.e. N.
tenuipes is present here.
Nematopalaemon tenuipes (Henderson, 1893)
(Figs.80-81)
Leander tenuipes Henderson, 1893:440; Nobili, 1903:7; Kemp,
1917:207; 1925:289; Chopra, 1943:5
Palaemon luzonensis Blanco, 1939:201
Palaemon (Nematopalaemon ) tenuipes Holthuis, 1950:44
Nematopalaemon tenuipes Holthuis, 1980:108; Holthuis & Miquel,
1984: Palem Nemat 3; Chace & Bruce, 1993: 39; Tirmizi & Kazmi, 1995:75;
Jayachandran, 2001: 194
Description: The rostrum is elongated and slender, extends beyond
the apex of the antennal scale; the basal crest is highly elevated and armed with
5-7 teeth, of which 2-4 are postorbital; The teeth increase in size from behind
forwards, the hind most as a rule is quite rudimentary; the tip of rostrum is
strongly curved upwards and with single sub distal tooth; the ventral margin is
provided with 2-6 teeth
181
Figure 80: Nematopalaemon tenuipes. Animal in lateral view
The carapace has antennal
branchiostegal groove is lacking.
and
branchiostegal
spines
but
182
Figure 81: Nematopalaemon tenuipes. A- Anterior part; B- Telson; CMandible; D- Maxillula; E- Maxilla; F- First maxilliped; G- Second
maxilliped; H- Left fourth leg
183
The abdomen is glabrous, laterally compressed. The pleurae of first to
third abdominal somites are broadly rounded posterio-ventrally, that of fourth
and fifth are directed backwards, of sixth no more than 2/3 length of carapace
and ending in sharp point.
The telson is long, extends beyond the middle of uropodal exopod.
The first chelate leg reaches a little beyond the apex of antennal scale;
the carpus is a trifle shorter than the merus and about 1.5 times longer than the
chela; the fingers are 1.5 times longer than the palm.
The second pereopods in most cases reach beyond the antennal scale
by at least length of chela, sometimes by that of the chela, carpus, and a small
portion of merus. The carpus is the shortest segment; the ischium is longer than
the palm and carpus; the merus being the longest segment; the palm is strongly
inflated and longer than the carpus; the fingers are slender.
The last three pereopods are non-chelate, elongate, slender, their
dactylii are more than twice the length of propodus.
Size: 12 -16mm in CL
Habitat: Littoral to 17 meters, brackish and marine
Distribution: South Africa, Somalia? India, Myanmar, Thailand,
Philippines, Taiwan, New Zealand?.
Genus Palaemon Weber, 1795
Diagnosis: Rostrum well developed, toothed dorsally and ventrally,
without an elevated basal crest; upper margin bearing single row of setae
between dorsal teeth. Carapace smooth, bearing distinct branchiostegal spine
and groove; branchiostegal spine usually situated on carapace edge but may be
set back; groove usually running straight back from carapace edge in shallow
arc but may rise upwards and over spine before continuing posteriorly or into
and over but not past its posterior edge. Antennal spine generally strong, on
anterior margin of carapace some distance below rounded lower orbital angle.
Mandibular palp usually of two or three articles but may be absent. Eyes
distinctly pigmented, cornea well developed. Anterior margin of antennular
peduncle rounded, anterolateral spine small. Propodus of fifth pereopod with
184
several transverse rows of setae on distal end of the posterior margin. First
pleopod of male without or with rudimentary appendix interna on endopod.
Telson with two slender median setae on posterior margin. Pleura of fifth
abdominal somite usually ending in small sharp point ( from Walker & Poore,
2003)
Type species: Palaemon adspersus Rathke, 1837
Gender: Masculine
Remarks: Often reported from submarine caves(Denitto et al,2009).So
far, 35 species have been recognized under this genus (Jayachandran, 2001).
Of the total only three species are so far recorded from the Pakistani waters.
KEY TO THE PAKISTANI SPECIES OF PALAEMON
1
Only one tooth of dorsal rostra series situated on carapace posterior to
level of orbital margin --------------------------------------------------------------------------------P. semmelinkii.
-
Two or three teeth of dorsal rostral series situated on carapace
posterior to level of orbital margin--------------------------------------------------------------------------------------------2.
2
Rostrum ascending anteriorly with margins tapering slightly in anterior
half, basal antennular segment with distolateral spine distinctly
overreaching adjacent convex distal margin of segment--------------------------------------------------------------P. pacificus.
-
Rostrum ascending anteriorly with margin tapering abruptly in anterior
half, basal antennular segment with distolateral spine distinctly
reaching midway of adjacent convex distal margin of segment -------------------------------------------------------P. sewelli.
Palaemon pacificus (Stimpson, 1860)
(Figs. 82-83)
Leander pacificus Stimpson, 1860:40;Nobili,1906:73; Balss,1915:31;
Kemp, 1925:307, Barnard, 1950:784.
Palaemon pacificus, Rathbun,1906:924; Holthuis;1950:87; Tirmizi &
Kazmi, 1984:313; Liu et al, 1990: 236; Chace & Bruce,1993:41; Jayachandran,
2001:211; Li et al, 2004:522
185
Description: The rostrum is straight, slightly upturned at the tip
and reaches to or a little beyond the end of the antennal scale. On the upper
border, there are from eight to 11 teeth, two, or three of which are situated on
the carapace behind the orbit. The foremost dorsal tooth is placed close to the
tip and is frequently but not always separated from the next by a rather long
unarmed interval. The branchiostegal spine is nearly as strong as the antennal
and is situated on the margin of the carapace. Branchiostegal groove is visible
just above the spine, anteriorly straight and decurved posteriorly.
1.
Figure 82: Palaemon pacificus. A- Female in lateral view; B- H- Variant in rostrum; I-JVariation in mandibular palp (After Tirmizi & Kazmi, 1984)
The telson is normal, anterior pair of dorsal spines is situated behind
midlenght.
The outer margin of the basal segment of the antennular peduncle
ends in a strong spine, which extends beyond the convex margin intervening
between the base of the spine and the articulation of the first and second
segments. The free part of the shorter ramus of the outer flagellum consists of
21 to 26 segments, the fused part comprising seven or eight segments. The
outer margin of the free part is always conspicuously serrate and the total length
186
of the shorter ramous is equal to or slightly greater than that of the
peduncle. The spine at the end of the antennal scale does not reach as far
forwards as the broadly rounded apex of the lamellar portion. The mandibular
palp is three segmented.
Figure 83: Palaemon pacificus. A- Anterior region;B- Antennule; CAntennal scale; D-G- First to fifth legs.
The first legs reach slightly beyond the end of the antennal scale. The
carpus is about 7 times as long as it‟s distal breadth and is shorter than the
merus and longer than the chela; the palm is about equal in length with the
figures.
The second legs reach beyond the scale by part or all the length of the
chela. The merus is about one-fifth longer than the carpus in larger specimens
less in smaller individuals. The palm is about one-fifth longer than the fingers,
The last three non-chelate legs are comparatively short and shout; the
third reach about to the end of the scale, while the fifth falls slightly short of it.
The propodus in all the three pairs bears some fine spinules on its posterior
margin and is not dilated distally.
There is no appendix interna on the first pleopod of the male
Remarks: The species exhibits a considerable range of variation in the
rostrum and mandibular palp (Tirmizi & Kazmi, 1984), these authors illustrate
187
some of the profiles of the rostrum, and their Fig. 80F is closer to P.
ogasawaraensis Kato & Takeda in having 11 teeth on the upper margin.
Size:Maximum size 55 mm in male
Localities: Bakran creek, Manora, Bulleji
Distribution: Red Sea, India, South Africa, Cape of Good Hope, Hong
Kong, Japan,Taiwan, New Caledonia and Hawaii
Palaemon semmelinkii (de Man, 1881)
(Figs. 84-85 Pl.3C)
Leander semmelinkii de Man, 1881:137; Ortmann, 1890:517;
Leander modestus Henderson, 1893:441 (non-Heller, 1862).
Leander Semmelinkii Nobili, 1903a:8
Palaemon semmelinkii, Bruce, 1987:58; Chace & Bruce,
1993:41; Jayachandran, 2001:202, Kazmi & Kazmi, 2002:
12; Cai & Shokita, 2006:266
Palaemon (Palaeander) semmelinkii Holthuis, 1950:57;Jagadisha &
Sankolli, 1977:619 (larvae) .
Description: The rostrum is straight, with the tip curved upwards; it
reaches with 1/5 to 1/3 of its length (1/3 to almost ½ in the males) beyond the
scaphocerite. The upper margin is provided with 7 to 11 (mostly 8 or 9) teeth;
the first of these teeth is placed on the carapace behind the orbit, it is more
remote from the second tooth than the third is, the second tooth is situated
slightly behind or just over the posterior margin of the orbit. The first teeth are
more or less distinctly movable, the ultimate 2 to 3 teeth are immovable. The
teeth are placed in the proximal part of the rostrum only: slightly more than the
distal third to slightly less than the distal half of the rostrum is edentate; there is
no sub apical tooth near the apex of the rostrum; the lower margin of the
rostrum bears 2 to 5 (almost always 3) teeth, here too the distal part is entire.
The setae of the upper as well as of the lower margin of the rostrum are
arranged in a single row. In the males, the rostrum is more slender than in the
188
females, the teeth, especially the distals, are longer and more pressed
against the rostrum proper. The carapace has the branchiostegal spine slightly
smaller than the antennal; it is placed on the anterior margin of the carapace.
The abdominal pleurae are normal in shape. The sixth somite is about
1.5 times as long as the fifth. The telson is distinctly longer than the sixth
abdominal segment. Of the two dorsal pairs of spines the anterior is situated in
the middle of the telson, the place of the posterior pair is rather variable, mostly
is situated midway between the anterior pair and the posterior margin of the
telson, sometimes, however, it is placed closer to the anterior spines,
sometimes closer to the posterior margin. The posterior margin is ending in a
sharp median point, and is provided with two pairs of spines, the outer of which
is noticeably shorter than the inner; a pair of feathered setae is present between
the two inner spines.
.
189
Figure 84: Palaemon semmelinkii. A- Anterior part, male; B- Same, female;
C- Antennular peduncle;D-Antennal scale;E-Mandible;F-I-First to fifth
legs(excluding fourth)
The eyes have the
ocellus is present.
cornea distinctly shorter and broader than the eyestalk. An
C
A
B
D
E
F
G
H
Figure 85: Palaemon semmelinki. A- Endopod of 2nd Pleopod; (Male CL
4mm); B- 2nd Pleopod (Male CL5mm); C- 2nd Pleopod and Appendices
(Male CL5mm); D- 1st Pleopod (Male, CL6.5mm); E- 1st Pleopod ( Female,
CL9mm); F- Male genital sternum (CL 6.5mm); G- Female, sternites ( non
ovigerous CL 5mm); H- Ovigerous female sternum.
The stylocerite of the antennular peduncle is sharp and slender, but
just fails to reach the middle of the segment; a dorsal carina is visible on the
stylocerite. The outer margin of the basal segment is more or less straight and
ends in a strong anterolateral spine, which reaches beyond the middle of the
second segment of the antennular peduncle and distinctly overreaches the
convex anterior margin of the basal segment. The second segment of the
190
peduncle is distinctly shorter than the third; together these two segments
are slightly shorter than the first. The upper antennular flagellum has the two
rami fused for about 7 to 12 joints; the free part of the shorter ramous consists
of 11 to 17 joints.
The scaphocerite is rather broad; it is about thrice as long as broad and
reaches with 1/3 of its length beyond the antennular peduncle. The outer margin
is straight or slightly convex and ends in a strong final tooth, which is distinctly
overreached by the lamella.
The oral parts are quite typical for the present genus. The mandibular
palp is two jointed. The epipod of the first maxilliped is divided into two rounded
lobes. The third maxilliped reaches beyond the antennal peduncle; the ultimate
segment is somewhat shorter than the penultimate; the antepenultimate
segment is slightly less than twice as long as the ultimate.
The first pereopod reaches much beyond the end of the antennular
peduncle, but fails to reach the end of the scaphocerite. The fingers are a little
shorter than the palm; they bear some tufts of setae and have blunt tips. The
carpus is somewhat less than twice as long as the chela, it broadens distally.
The merus is distinctly shorter than the carpus. The ischium is about half as
long as the carpus.
The second pereopods are stronger than the first, they reach with the
fingers, sometimes even with half the palm, beyond the scaphocerite. The
fingers are slender; they measure about 2/3 of the length of the palm; the cutting
edge of the dactylus is provided in its proximal part with a small inconspicuous
tooth, the rest of the cutting edge of the dactylus as well as that of the fixed
finger is minutely spinulated; the palm is cylindrical and slightly swollen. The
carpus is as long as the chela, sometimes slightly shorter or somewhat longer; it
broadens distally. The merus measures about 4/5 of the carpus; the ischium is
not as long as the merus as given by earlier workers. All segments of the
second leg are setose.
The third pereopod reaches about to the end of the antennular
peduncle; the propodus measures somewhat less than thrice the length of the
barbed dactylus; the posterior margin of the propodus is provided with scattered
spinules; the carpus is somewhat more than half as long as the propodus; the
merus is much stronger than the propodus. The ischium is half as long as the
merus.
The fourth pereopod is slender; it reaches almost to the end of the
scaphocerite. The propodus is more than thrice as long as the dactylus. The
carpus is half as long as the propodus. The merus is about as long as the
propodus. The ischium is less than half as long as the merus.
191
The fifth pereopod resembles the fourth leg it is slightly more
slender. The propodus of fifth leg has transverse rows of setae in the distal part
of the posterior margin, these setae are closely packed together otherwise the
leg is almost nonsetose.
The pleopods are normal in shape. The endopod of the first pleopod of
the male is ovate and bears no appendix interna; its inner margin is concave.
The appendix masculina of the second pleopod of the male is strong and
reaches distinctly beyond the appendix internal, can be visible in males of
2.0mm in CL to sex the shrimp. The thoracic sternites differ not only in the two
sexes but in breeding and non breeding females
The uropods are elongate; they are of the normal shape.
Remarks: The species has a vulnerable status in India along its west
coast (Dholakia, 2004); it is one the commonest palaemonids in our waters.
Size: 2-7 mm in CL (male), 2-10mm in CL (female)
Habitat: Shallow coastal, sometimes brackish waters
Localities: Back water of Mauripur to Gharo creek, Bakran creek, Port
Bin Qasim and Ibrahim Hydri.
Distribution: Indo- Pacific
Palaemon sewelli (Kemp, 1925)
(Fig.86)
Leander sewelli Kemp, 1925:299
Palaemon sewelli Holthuis, 1950:8; Liu et al 1990:240; Ghani, 1999:37;
Jayachandran, 2001:203; Li et al, 2004:525
Description: The rostrum reaches about to the end of the antennal scale, but is
some times a little shorter and occasionally rather considerably longer. Its upper
border is straight proximally but it is usually a little upturned in its distal third.
There are altogether from 11 to 17 upper teeth of which from 3 to 5 are situated
on the carapace behind the posterior limit of the orbit; posterior tooth is
sometimes a little more distant from the second than the second is from the third
and the teeth are generally rather closely packed over the eyes and more widely
192
separated distally. The distal one or two teeth are sometimes, but not
always, separated by a distinct interval from those further back; the distal tooth
is not placed so close to the apex as to give it a bifid appearance. The rostrum
is deep and it bears on its lower border from three to five teeth. On both upper
and lower borders, there are setae in the interspaces between the teeth.
Figure 86: Palaemon sewelli. A- Entire specimen, lateral view, B- Anterior
part, lateral view; C- Telson ; D- Tip of telson; E- Setae on propodus of last
leg ( Modified from Ghani, 1999).
The branchiostegal spine is nearly as large as the antennal; it
is set well back from the margin of the carapace, but the tip reaches beyond the
margin.
The ocular spot is distinct and touches the cornea, the cornea itself
being rather narrow and not much wider than the stalk.
The sixth abdominal somite is less than twice as long as the fifth. The
telson bears the usual two pairs of dorsal spines; the foremost is placed a little
behind the middle of its length. The apex is normal, with two spines and one
feathered seta on either side of the median point.
The distal spine of the basal segment of the antennular peduncle
reaches to the middle of the second segment; its tip projects far beyond the
convex border, which intervenes between the base of the spine and the
193
articulation of the first and second segments. In the outer flagellum, the free
part of the shorter ramus consists of 10 to 13 elongate articles. The fused part
comprises 5 to 6 articles and the total length of the shorter ramus is less than
that of the peduncle.
The antennal scale is proportionately broader in females. The outer
margin is straight or slightly concave, terminating in a spine, which reaches
almost or quite as far forwards as the rather broadly rounded apex of the
lamella.
The mandibular palp is composed of three segments and the other oral
appendages are normal.
The first legs extend to the end of the antennular peduncle. The carpus
is slightly shorter than the merus and slightly longer than the chela. The fingers
bear a few setae and are a little longer than the palm.
The second legs of adults reach beyond the antennal scale by the
fingers and about half the length of the palm. The chela is longer than the
carpus, with the palm about 3 times as long as wide and equal in length with the
fingers.
The last three legs are slender. The third pair reaches almost to the
end of the antennal scale, while the fourth and fifth pair reaches a little beyond
it. The merii of these legs are long and the propodii are more than twice longer
than the dactylii, the dactylii are very slender, with a few setae on their anterior
borders.
There is no appendix interna on the first pleopod of the male.
Remarks: Sometimes mistakenly identified as P. serrifer.
Colour: Live specimens are deeply pigmented, dull reddish in colour,
pale dorsal blotches on abdominal somites are occasionally present .
Size: 32mm
Habitat: Generally lives in water of low salinity
Distribution: India, Bengal, Myanmar, and Northern South China
Sub-family Pontoninae Kingsley, 1878
194
Diagnosis: Upper antennular flagellum with both the rami fused in
the basal part. Both, the appendix interna and appendix masculina present on
the second pleopod of male and only an appendix interna present on the
second pleopod of female. Pleurobranch absent from the third maxilliped.
Posterior margin of the telson armed with three pairs of spines (except in
Anchistioides) (Modified from various sources).
Remarks: The most remarkable feature of this sub-family is the ability
its 450 members have shown in forming associations with other animals. Some
are found on sponges, others on Actinians, alcyonaria and madrepore corals, a
few are to be met with asteroids, and echinoids and many live on crinoids. A
considerable number of species occur in the mantle cavity of lamellibranch
molluscs and some are known from the branchial sac of ascidians. Many
species are also free living.
Pontoniinae is a large sub-family, containing 93 genera and 510
species (Marin, 2006), 44 species occur in North West Indian Ocean, two
genera Anchistus and Cuapetes occur in Pakistan. The third genus
Periclimenes as P. digitalis is given in an unpublished report from Pakistan
therefore not treated any further. Apparently, the presentation is certain to
increase when the rich coral reef and deep-sea fauna is investigated.
KEY TO PAKISTANI GENERA OF PONTONINAE
1.Carapace bearing hepatic spine------------------Genus Cuapetes.
-Carapace without hepatic spine -------------------Genus Anchistus.
Genus Anchistus Borradaile, 1898
Diagnosis: Rostrum short, cylindrical in basal part, laterally
compressed in distal half and frequently with small teeth at or near apex.
Carapace notably for disappearance of all spines (except for occasional
presence of antennal spine). Abdomen smooth. Pleurae of first five segments
broadly rounded at posterior margin, of VI bluntly spinous. Telson broad; dorsal
surface with two pairs of very small spines (rarely 3 pairs); posterior margin also
with three pairs of spines.
Eyes well developed. Cornea a little narrower than stalk and black
ocular spot distinct.
195
Antennule typical in structure. Antennal scale broad, oval, final
tooth failing to reach tip of lamella. Mandibles with incisor process tridentate;
molar process with blunt knobs and ridges and tuft-like spines. Mandibular palp
absent. Inner lacinia of maxillula broadened, palp generally not cleft at distal
end. Palp of maxilla broad. Scaphognathite well developed but not so broad. All
maxillipeds with exopods. Maxilliped normal in shape. Second maxilliped lacks
podobranch; arthrobranch present at base of third maxilliped.
First chelate legs slender, equal. Carpus entire. Second chelate legs
stouter, frequently unequal. Chela cylindrical, somewhat inflated. Fingers
elongate and with small denticles, which are never, hammer shaped. Dactyl of
non-chelate legs either simply hooked or scoop-shaped with distal part of
anterior border bent inwards and with accessory tooth.
Pleopods and uropods normal in structure. (After Jayachandran, 2001)
Type species: Harpilius Miersi de Man, 1888
Gender: Masculine
Remarks: Of the present genus seven valid species are known
(Jayachandran, 2001), represented in Pakistani waters by a single speciesAnchistus custos
Anchistus custos (Forskal, 1775)
(Figs.87-88)
Cancer custos, Forskal, 1775:94
Anchistus custos Holthuis , 1952:105 (for complete synonyms); Johnson &
Liang, 1966:433; Miyake & Fujino, 1968:415; Tirmizi & Kazmi, 1982:115; Titgen
(unpublished), 1982:86; Chace & Bruce, 1993:46; Bruce, 1996:205;2003:211;
Jayachandran, 2001:242; Li et al, 2004:528;Li&Bruce,2006:625
Anchistus inermis Barnard, 1950:792
Description: The rostrum is directed downwards, toothless and with
the apex broadly rounded in lateral view. The lower limit of the orbit is defined
by an acute projection from the frontal margin of the carapace; the antennal
spine is either altogether absent or is represented merely by a minute pointed
process.
196
Figure 87: Anchistus custos. A- Animal, dorsal view; B- Mandible;
C- Maxillula; D-Maxilla; E- First maxilliped; F- Second maxilliped; GThird maxilliped; H- First leg; H’- Same, chela enlarged.
The apex of the telson is generally armed with six spines. The two
forming the median pair are more slender than the intermediates; the outermost
are very short and inconspicuous and are occasionally missing. The dorsal
spines are very small and are sometimes absent. When present the anterior pair
is placed behind the middle of the telson, with the posterior pair midway
between the first pair and the apex.
The basal segment of the antennular peduncle is produced distally on
its outer side in the form of a convex lobe, the outer margin terminating in a
short spine. The fused portion of the outer antennular flagellum comprises five
segments. The antennal scale is broadly oval and little narrowed anteriorly; the
strongly convex outer border terminates in a rather small tooth, which does not
reach the distal end of the lamella.
The propodus of the third maxilliped is longer than the two distal
segments taken together and is very broad; its least breadth is more than three
times that of the dactylus. The lower margins of the basis and ischium of the first
197
pereopods are heavily fringed with setae. The carpus is little longer than the
merus and nearly twice as long as the chela; the fingers are much shorter than
the palm. The structure of the chela is very peculiar. The edge, both on its outer
and inner side, is produced to form a sort of flap, which is bent downwards and
is thickly fringed with long setae on its margin. The chela is thus deeply
hollowed in a longitudinal direction when viewed from below and in a transverse
section, the lower surface would be semicircular.
The second pereopods are unequal, either the right or left limb may be
enlarged. In the larger of the two, the merus is from 2.0 to 2.4 times as long as
broad; the carpus is very short, only one-sixth to one-eighth the length of the
chela and the fingers are a little more than half the length of the palm. The
dactylus is strongly convex externally. On the inner margin, it bears in its basal
half a very large triangular tooth and a rounded knob close to the articulation;
when the claw is closed both the tooth and the knob are received into large
socket in the fixed finger. The inner margin of the fixed finger is obtusely
produced in the middle and in the basal half, on a crest, which borders the
socket on its upper side, there are usually from 3-6 small denticles, the foremost
placed at the summit of the obtuse prominence referred to above. In all well
developed specimens, the distal half of each finger is internally concave. The
fingers of the smaller limb are similar, but the tooth on the dactylus is usually
less well developed. In the last three pereopods, the propodus is without
spinules on its posterior edge. The dactylus is strongly hooked, with the terminal
claw bent at right angle to the proximal portion. It is extremely slender, the
basal breadth being only about half that of the distal end of the propodus. The
endopod of the first pleopod of the male shows a laterally directed triangular
process in the distal part of the inner margin.
198
Figure 88: Anchistus custos. A- Rostrum with left antennae and eye, dorsal
view; B- Rostrum and eye, lateral view C- Telson and right uropod; D- Right
second leg outer view; E- Same, chela inner view;
Colour:Living specimens vary in colour from pale straw to bright
orange yellow. In female the entire body and legs are covered with minute white
dots and the eggs are pale straw yellow orange or brown. Males are
semitransparent and lack the white dots found in the female.
Size:
19.5mm inTL.
Habitat: The species generally known as watch man prawn have
adopted a secluded mode of life, living in the branchial cavity of bivalve
molluscs. The prawn probably enters its host when larvae and never leaves it
throughout the whole period of its life. However, Macnae & Kalk (1958) dissent
from this view and report that free-living individuals of A. custos are common in
their area. The spineless rostrum is an adaptation to more freedom of
movement on or with in its host (Bruce, 1976) whereas Johnson & Liang, (1966)
hold that canaliculation of the palm is a modification to collect mucus from the
host‟s gills and to consume it.
Locality:Bulleji
Distribution:
Indo-Pacific
199
Genus Cuapetes Clark,1919
Diagnosis: Small to medium sized pontoniine shrimps of sub
cylindrical body shape. Carapace smooth, glabrous, with rostrum well
developed, dorsally and ventrally dentate, with or without epigastric or
suborbital, spines, hepatic and antennal spine present, orbit obsolete, inferior
obital angle distinct, anterolateral angle rounded. Abdomen smooth, glabrous,
pleura rounded posterolateral angle bluntly rounded or dentate. Antennules with
short stylocerite; flagella well developed, upper ramus with shorter flagellum
multi-segmented. Antenna with basicerite with lateral tooth, scaphocerite well
developed. Eye well developed, elongate, and sub cylindrical, cornea globular,
ophthalmic somite without median process. Mandible without palp; molar and
incisor processes well developed. Maxillula with feebly bilobed palp. Maxilla
normal, with simple palp, basal endite simple, coxal endite obsolete. First
maxilliped with simple palp, basal and coxal endites feebly separate, broad,
exopod with reduced flagellum, caridean lobe small, broad; epipod large,
generally triangular. Second maxilliped with normal endopod, dactylar segment
narrow, exopod well developed, epipod small, subrectangular, with rudimentary
podobranch. Third maxilliped normal, ischiomerus not fused to basis, exopod
well developed, generally with numerous plumose setae distally, coxa with
rounded lateral plate, generally with small or rudimentary arthrobranch. Thoracic
sternites narrow, fourth with slender finger-like median process, posterior
sternites without acute processes. First pereopod slender, chela with fingers
simple. Second pereopod well developed, frequently slender, elongate,
generally equal or unequal, similar or dissimilar; major chela fingers without
molar process and fossa. Ambulatory legs slender, dactyls simple, without basal
process. Uropod with protopodite distolaterally dentate; exopod with small tooth
and mobile spine distolaterally. Telson with two pairs of dorsal spines, three
pairs of posterior spines. ( from Bruce, 2004 as Kemponia)
Type species: Anchistia grandis Stimpson, 1860
Gender: Feminine
Remarks: The genus Kemponia designated by Bruce (2004) has been
resurrected by Okunu(2009) and currently all its 24 species are placed in
Cuapetes Clark,1915 by Okunu(2009); of these only three are reported in
Pakistan .
200
KEY TO THE PAKISTANI SPECIES OF CUAPETES
1.Merus of second leg without a spine or tooth at distal end of
lower border------------------------------------------------------------------------------C. seychellensis
-Merus of second leg with a spine or tooth at distal end of
lower border------2.
2.Rostrum shallow. Last three legs long and slender, third pair
with merus at least 11 times as long as broad, fifth pair
reaching beyond antennal scale-------------------------------------------------------------------------------------- C. longirostris.
-Rostrum moderately deep. Last three legs stouter, third pair
with merus at most 10 times as long as broad, fifth pair not
reaching end of scale--------------------------------------------------------------------------------------------------- C. elegans
Cuapetes elegans (Paulson, 1875)
(Figs.89-93)
Anchistia elegans Paulson, 1875: 113, pl.17fig.1.
Periclimenes elegans Balss, 1915: 26; Bruce, 1971:7; Kazmi
&Qureshi,1974: 197; Kazmi &Kazmi, 1979:152; 2004:97;
Chace & Bruce, 1993:110; Jayachandran, 2001:296; Li,
2000:178
Periclimenes (Falciger) dubius Borradaile, 1915: 211
Periclimenes (Ancylocaris) elegans-Kemp, 1922: 215, text
figures 60-62; 1925:322.
Periclimenes (Harpilius) elegans- Holthuis, 1952:81,fig. 31;
201
, 1961:59; Miyake & Fujino, 1968: 406,fig.3.
Kemponia
elegans
Bruce,2006:641;Li,2008:214
Bruce,
2004:14;
2006:16;Li&
Cuapetes elegans Okunu,2009:67
Description: The rostrum is elongated and rather broad in lateral view.
It reaches beyond the scaphocerite. The basal half of the rostrum is more or
less horizontal, whereas, the distal half is directed upwards. The upper margin
of the rostrum is armed with eight prominent teeth. The epigastric tooth is
situated behind the level of supraorbital spine. Four teeth, which are separated
by a considerable gap, follow it; three distal teeth are situated near the apex.
The lower margin bears four teeth. Supraorbital and hepatic spines are placed
rather close behind the antennal spine, but on a lower level. The abdominal
somites are smooth; the poster lateral margins of all the somites are broadly
rounded, except the last two, each of which ends in a small acute process. The
poster lateral margin of the third abdominal somite covers one-third portion of
the fourth somite. The sixth abdominal somite is one and a half times as long as
the fifth.
Figure 89: Cuapetes elegans. A-Lateral view of carapace; B-Lateral view
of abdominal somites 1-6; C- Telson and right uropod
The supraorbital spine is small but distinct. The pterygostomian angle is
rounded.
202
The telson is about one third longer than the last abdominal somite.
The dorsal surface of the telson is armed with two pairs of spines; the first pair is
situated a little proximal to the middle of the telson, the second pair is situated
more or less half way between the first pair and the tip of the telson. These
spines divide the telson into three sub-equal parts; in this respect, the present
specimens differ from the Indian specimens, described by Kemp (1922) as
Periclimenes elegans. According to him, the dorsal spines of telson are so
arranged as to divide its length into three equal parts (1922:213). The telson
tapers gradually to a sharp distal end. On the dorsal surface of the telson, a
distinct longitudinal groove is present, which extends from the base to a little
distance beyond the second pair of spines.
The eye is short and broad. The cornea is hemispherical.
The basal segment of the antennular peduncle is three times longer
than its maximum width. It reaches up to the middle of the scaphocerite. The
lateral margin of the basal segment is straight, ending in a sharply pointed
spine, the median margin has a well-developed spine, and the stylocerite is
sharply pointed and just fails to reach the middle of the segment. Its lateral
margin is fringed with plumose setae. Second and third segment of the
antennular peduncle are nearly of equal size, the combined length of these two
segments is a little more than half the length of the basal segment. The outer
flagellum is cleft only for a short distance; the fused basal part is longer that the
peduncle.
The scaphocerite is slender, more or less arched and becoming narrow
towards the tip. The lateral side is concave and ends in a spine, which extends
a little beyond the squamose portion. Two distinct longitudinal carinae are
present in the middle of the scaphocerite.
The incisor process of the mandible has three teeth, the median one
being the smallest. The molar process is well developed and toothed, the
mandibular palp is absent. The maxillulary palp is bifid, its upper lobe is narrow
and sharply pointed, whereas the lower one is broad and truncated; the upper
lacinia is stronger than the lower lacinia. The palp of the maxilla is narrow and
fingerlike. The lacinia is divided into two sub equal lobes; the lower one being
just a trifle shorter and narrower than the upper one.
203
Figure 90: Cuapetes elegans. A- Right eye; B- Right antennular peduncle
in dorsal view; C- Right scaphocerite in dorsal view; D-Left mandible.
The first maxilliped has a very slender medially curved palp, which is
slightly longer than the caridean lobe. The lateral side of caridean lobe is
strongly convex and broadly rounded apically. In the second maxilliped, the
ultimate segment is attached laterally to the penultimate segment. The
anteromedian angle of the para penultimate segment is produced into an acute
spine. The third maxilliped is leg like; the ultimate segment is nearly 2.5 times
shorter than the parapenultimate segment. The outer margin of the
parapenultimate segment is armed with four spines. The number of spines
seems to be variable.
204
Figure 91: Cuapetes elegans. A-Right maxillula in dorsal view; A’-Same,
palp further enlarged; (in ventral view); B-Right maxilla in dorsal view; CE-Right first to third maxillipeds.
205
Figure 92: Cuapetes elegans. A- First pereopod; B-Second pereopod; B’Carpo-propodal joint of second pereopod, (further enlarged to show
spines); C-E- Third to fifth pereopods (All from right side);
All the pereopods are long and slender. The first pair of pereopods is
symmetrical. The fingers are slightly shorter than the palm. The carpus is the
longest segment, being 1.5 times longer than the chela, it suddenly broadens in
its distal half, it is 6.5-7.5 times as long as its maximum breadth. The second
pair of pereopods is also symmetrical, extends beyond the antennal scale, by
the length of carpus and chela, particularly so in the males. In the movable
finger, a blunt tooth is present near the proximal end. An excavation and an
acute tooth, which is situated in the middle of the cutting edge, follow it. The
immovable finger is armed with three sharp teeth; the dentition of the finger is
subjected to variation. Setae are present on the entire length of the outer and to
some extent the inner margin, they are not confined to tips only, the palm is 2.7
times as long as the movable finger. The chela is also much longer than the
carpus, being three times the length of the carpus. The carpus is also shorter
than the merus. The anterodistal margin of the carpus has two strong spines; a
longitudinal groove can be seen on the carpus. The inner distal margin of the
merus bears a strong spine. It may be pointed out here that the second leg
206
presents rugose appearance. The third, fourth and fifth pereopods are as
illustrated. The third pereopod is slightly longer than the last two pereopods. The
propodii of third, fourth and fifth pereopods are longer than the merii. The
propodii of last three pairs are armed with spines.
The first pleopod has a short endopod, which covers only half the
length of the exopod. The second pleopod bears an appendix interna and an
appendix masculina. The appendix masculina is slender and extends as far as
the tip of the enopod.The appendix interna is rather short, and curved, on its tip
a few rows of coupling hooks are present.
The exopod of uropod has a movable spine on its lateral side.
Figure 93.Cuapetes elegans. A-Left first pleopod of male; B- Left second
pleopod of male
Colour:Off white, anterior part pink (when fresh)
Size:35 mm inTL
Locality:
Manora Island
Habitat: Shallow waters,0-24m depth
Distribution: Throughout the Indo-West Pacific region from Red Sea,
Persian Gulf, Egypt, Saudi Arabia, Aden, Kuwait, Kenya, Bay of Bengal, the
Peninsular Malaysia to Hong Kong, Dampier Achipelago, Australia, Queensland
and Micronesia.
207
Cuaperes longirostris (Borradaile, 1915)
(Fig. 94)
Palaemonella longirostris Borradaile, 1915:210; 1917:359
Periclimenes (Falciger) affinis Borradaile, 1915:211;1917:372
Periclimenes (Ancylocaris) proximus Kemp, 1922:201
Periclimenes (Harpilius) longirostris Holthuis, 1958:3
Periclimenes longirostris Bruce, 1974:191; Chace & Bruce,
1993:117; Li, 2000:208
Periclimenes proximus Jayachandran, 2001:310
Kemponia longirostris Bruce, 2004:17; 2006:16
Cuapetes longirostris Okunu,2009:68
Description: The rostrum is slender and reaches almost to or a little
beyond the apex of the antennal scale. It is little up turned in its distal third and
bears on the slightly concave upper margin 6-7 large teeth, the posterior tooth is
placed on the carapace and is rather more distant from the second than the
second is from the third; the remainder are more or less evenly spaced and the
seventh tooth when present is much smaller than the rest and placed quite
close to the apex; on the lower border there are 2-3 teeth which are larger and
placed in advance of the middle of the rostrum. Supraorbital, antennal, and
hepatic spines are present. The eyes are large and depressed.
208
Figure 94:Cuapetes longirostris. A- Anterior part; B- Antennule; CAntennal scale; D- First leg; E- Second leg, male; F- Second leg, female;
G- Third leg ( Modified from Jayachandran, 2001 as Periclimenes
proximus).
The basal segment of the antennular peduncle has a short lateral
process, not reaching the middle of the segment; its terminal spine is long,
reaching the middle of the second segment, and the margin between the base
of this spine and the articulation is gently convex, the two distal segments are
slender. The antennal scale is 4-5 times as long as wide and is proportionately
longer in large males; the scale is very narrow at the apex, the outer margin is
strongly concave and terminates in a spine, which outreaches the end of the
lamella. The antepenultimate of the third maxilliped bears 1-2 spines on its outer
edge.
The merocarpal articulation of the first leg reaches the end of the
second segment of the antennular peduncle in adult males, not quite so in
female; the carpus in adults of both sexes is from 1.2-1.3 times the length of the
merus and about 1.4 times the length of the chela, the legs are stout, the fingers
are longer than the palm and are unarmed.
The second pereopods bear a conspicuous sub terminal spine on the
lower side of the merus, in large males they may be as much as six times the
length of the carapace, extending beyond the scale by more than length of the
carpus and chela. The legs of pair are equal or sub equal and similar in
209
structure. The second legs of males are closely covered with minute
spinules. In female the chela is always definitely longer than the carpus, the
chela is 1.4 times as long as the carpus, with palm 1.6 times as long as the
fingers.
The last three pairs of legs are slender, the dactylii are slender and curved with
a few setae in the middle of their anterior margin.
Colour: In specimens in alcohol, a bright spot is seen at the end of the
carpus of the second leg and a narrow red band across the fingers of the same
appendages.
Size: 17.5mm
Habitat: Recorded in association with Alcyonaria by Fransen (1994)
but most reports suggest a free-living species.
Distribution: Israel, Zanzibar, Seychelles Island, Chagos Island,
Andaman Island, Indonesia, Philippine Island, Queensland, Papua New Guinea,
and Marshall Island.
Cuapetes seychellensis (Borradaile, 1915)
(Figs.95-98)
Periclimenes ensifrons Nobili, 1899:234.
Periclimenes tenuipes (part) Nobili, 1899:235
Periclimenes (Falciger) seychellensis Borradaile, 1915:212;
1917,324, pl54, 55, fig. 14a-14j.
Periclimenes (Ancylocaris) seychellensis-Kemp, 1922:176
Text fig. 34-35pl 6, fig.7.
Periclimenes (Harpilius) seychellensis Holthuis,
1952:67,fig.25; Barnard, 1956:17; Johnson, 1961:58, table
1;Kazmi et al, 1975:105
Periclimenes seychellensis Bruce, 1971:8; 1974:192;
Chace & Bruce, 1993:121; Li,2000:235; Jayachandran,
2001:314; Kazmi, 2003:240
210
Kemponia seychellensis Bruce, 2004:19
Cuapetes seychellensis Okunu,2009:68
Description: The rostrum is elongated and rather broad in lateral view.
It is slightly shorter than the carapace. In small specimens (C.L. 9mm) it reaches
just up to the tip of the scaphocerite, while in the slightly larger specimens (C.L.
10mm) it extends beyond the scaphocerite and the antennular peduncle. The
upper margin bears seven teeth while the lower margin is armed with three to
four teeth .In shape, size and the position of the upper rostral teeth the Pakistani
specimens differ from Indian specimens described by Kemp. According to
Kemp, the rostrum has a “concave upper border. Dorsally it bears from seven to
nine teeth, usually 8. The two hind most teeth are situated on the carapace
behind the orbit” (1922:176), whereas, in our specimens the rostrum is more or
less straight, except at the tip where it is inclined downwards and instead of two,
there are three teeth behind the orbital angle (including the epigastric tooth).
The hepatic spine being placed rather close behind the antennal spine, but at a
lower level.
The abdominal somites are smooth. The posterolateral margins of the first three
somites are large and broadly rounded. The postero-lateral margin of the
second somite covers only one-third portion of the third, whereas, the third
somite covers three fourth portion of the next somite. Each of the last three
somites ends postero-laterally into a small acute process. The sixth somite is
one and half times longer than the fifth.
211
Figure 95.Cuapetes seychellensis. A- Animal in lateral view; B- Dorsal
view of telson with left uropod; C-Antennule; D- Antennal scale.
The telson is slightly longer than the sixth abdominal somite. It tapers posteriorly
and ends in to a sharp point. The dorsal surface of the telson is armed with two
pairs of spines, which divide the telson into three sub-equal parts. According to
Kemp, “The two pairs of dorsal spines on the telson are large and are placed so
as to divide its length into three equal parts” (1922:177). The tip of the telson
closely resembles with that of C. elegans.
An excavation and a papilla on the peduncle characterize the eye. The
excavation is always present though the papilla may be absent in some cases.
The basal segment of the antennular peduncle is twice as long as broad in the
middle. It reaches beyond the middle of the scaphocerite. The outer margin of
the basal segment is slightly convex and ends in a sharp spine. This spine is
well developed and may extend almost up to the middle of the second segment
of the antennular peduncle. The stylocerite is sharply pointed and reaches up to
the middle of the basal segment. The second and third segments are sub-equal
in length. The scaphocerite is three times as long as wide in the middle. The
lateral margin of the scaphocerite is slightly concave and ends into a spine. The
spine is short and fails to reach the end of the anterior margin of the squamose
portion.
The mandible is normal. The maxillulary palp is bilobed; upper lobe is sharply
pointed and spiniform, whereas, the inner lobe is peculiar in having two spines,
of which the lower is curved inwards, the lower lacinia is slender while the upper
lacinia is broad. The maxilla has a somewhat broadened palp, which ends into a
slightly curved distal end. The inner lacinia is divided into two sub-equal lobes.
The palp of the first maxilliped is slender and finger like in shape. It is a trifle
longer than the caridean lobe. The caridean lobe is broad and more or less oval.
212
Figure 96: Cuapetes seychellensis. A- Mandible of left side; B-Right
maxillula in dorsal view; B’- Palp of same (further enlarged); C- Maxilla; C’Palp of same (further enlarged)
The second and third maxillipeds are of typical caridean type as illustrated.
Figure 97: Cuapetes seychellensis. A-C-Right first to third maxillipeds.
213
The pereopods are slender and sparsely hairy. According to Kemp, “the first
peraeopods reach about to the end of the antennular peduncle” (p.177). We find
the length of the first pair of pereopod as variable. In one specimen (C.L. 9mm)
it extends beyond the antennular peduncle, while in the other it reaches as far
as the end of the antennular peduncle (C.L. 10mm) and in one specimen, which
is of the same size, it fails to reach the distal end of the antennular peduncle.
The length of the chela is not related to the size of the animal. It is in the smaller
specimen, that it is comparatively large. The cutting edges of fingers are
unarmed. The second pair of pereopods varies among the present specimens,
extending beyond the scaphocerite by the length of the chela to half length of
chela. The cutting edges are straight and with a few setae, the carpus is longer
than the palm. The carpus and merus are without any spine and are nearly
equal in size. The third pereopod is smaller than the others; the propodus bears
a row of setae on the posterior margin and is 3.5 to 4 times the length the
dactyls; the dactylic of last three pairs of legs are simple.
Figure 98: Cuapetes seychellensis. A - B, - Right first and second
pereopods. B’ - B”- Fingers of the chela; C- Right fourth pereopod
The appendix interna on each of the last pleopods is long and slender; the tip of
the appendix internae are furnished with rows of coupling hooks.
214
Colour: Live specimens are closely mottled pale, buff, lichen-green
and brown.
Size: 19mm
Habitat: Sea grass, including algae ,offshore floating Sargassum in
shallow water.
Localities: Korangi creek, Sandspit, Manora, Port Qasim, Younas
Abad, Goth Abdul Rahman and Sonari.
Distribution: Kenya, Zanzibar, Madagascar, Gulf of Suez, India,
Seychelles, Gulf of Manaar, Andaman Islands, Mozambique, Singapore,
Indonesia, China, Western Australia and New Caledonia.
215
3.1.5 Family Processidae Ortmann, 1896
Diagnosis: Rostrum discrete structure inflexibly attached to remainder
of carapace, unarmed except (usually) pair of teeth delimiting terminal setaefilled notch. Carapace without longitudinal lateral ridges, complete post antennal
suture, or cardiac notch. Telson bearing two pairs of posterior marginal spines
and one or more pairs of mesial setae. Eyestalks normal, neither abnormally
long nor concealed beneath carapace. Antennules with two completely separate
flagella, neither with accessory branch. Mandible without palp or incisor
process, latter obliquely truncate, sometimes slightly flared. Second maxilla with
endite reduced, scaphognathite with proximal lobe produced only moderately
into branchial cavity. First maxilliped with exopod abutting endite, without
partially detached lobe, lash well developed, and caridean lobe not much
produced distally, not distinctly overreaching endite. Second maxilliped with
exopod, endopod composed of four segments, not terminating in two segments
attached side by side to preceding segment, terminal segment narrow strip
attached obliquely to wide penultimate segment. Third maxilliped with exopod
composed of five segments, slender, pereopod-like, antepenultimate segment
fused with next proximal segment. Pereopods without epipods, anterior pair
more robust than second pair, often asymmetrical, second pair equal, with
undivided carpus, fixed finger not curving subrectangularly around short, broad
movable finger, fingers not concealed in dense setae. Third pereopod with
dactylus simple, unarmed on flexor margin. First pleopod of male with endopod
laminar, not unusually large, or elaborately convoluted. (d from Chace, 1997).
Remarks: Shrimps of the family Processidae are small, nocturnal
animals, which are abundant in shallow water, primarily on grass flats in tropical
and warm temperate latitudes. Little is known about the basic biology of
processids. Their robust first legs indicate predatory feeding (Bauer, 2004).
Of the five processid genera and recognized so far, only one genus
Processa is present in Pakistani waters.
Genus Processa Leach, 1815
Diagnosis: Rostrum usually slender in dorsal aspect. Telson with
distinct dorsolateral spines. Mandible with molar process more than 1/5 as wide
as minimal length. Third maxilliped usually with exopod. Anterior pereopods
without exopod, I member (usually right) chelate, other with simple dactyl.
Second pereopods with shorter member composed of more than six carpal
216
articles (After Chace, 1997).
Type species: Processa canaliculated Leach, 1815 by monotypy
Gender: Feminine
Remarks: More than 67 species of Processa have been recorded from
the world oceans. One species P. compacta belonging to P. edulis species
group is described from Pakistan .
Processa compacta Crosnier, 1971
(Figs. 99-104)
Processa barnardi Hayashi, 1975: 92; Jagadisha & Sankolli,
1977b: 601 (larvae)
Processa cf. edulis Barnard, 1947:386; 1950:178
Processa edulis crassipes Kazmi & Kazmi, 1973: 51
Processa edulis Kazmi & Kazmi, 1979:155
Processa compacta Crosnier, 1971: 577; Noel, 1986:273;
Hogarth, 1987:111; Chace, 1997: 37
Description: The carapace is globular, unarmed except the presence
of small antennal spine. The rostrum is short, slender, straight, and reaching
only up to the middle of the eyes. The tip of the rostrum is bifid, moreover, the
tip and the lower margin of the rostrum is hairy.
217
Figure 99: Processa compacta. A- Lateral view of carapace; A’- Tip of
rostrum; B- Lateral view of abdomen; C- Telson and uropods; (After Kazmi
& Kazmi, 1973 as P. edulis)
There is only one denticle on the postero-inferior angle of the fifth
abdominal somite; the pleuron of the sixth abdominal somite has a sub-median
process with a small sharp spine, the postero-inferior angle is produced into a
strong spine. The telson is two times as long as the sixth abdominal somite, it
tapers slightly towards the posterior end. The posterior margin is armed with a
small median spine, flanked by two to three pairs. The dorsal surface of the
telson is rather strongly sulcate and bears two pairs of spines.
218
Figure100: Processa compacta. A-Eye from right side; B-Right antennular
peduncle in dorsal view; C-Right scaphocerite in dorsal view; D-Mandible
of right side; D’-.Same, tip of molar process. (After Kazmi & Kazmi, 1973
as P. edulis)
The eyes are not very large; the cornea is hardly any wider than the
stalk, the stalk becomes suddenly narrow at the base; the eyes reach almost to
the distal portion of the basal segment of the antennular peduncle.
The basal segment of the antennular peduncle is slightly excavated
dorsally; it is longer than the combined length of the last two segments. The
stylocerite is short and truncated anteriorly; both the antero-median and anterolateral angles are produced into microscopic spines. The stylocerite reaches
nearly up to the one-third portion of the basal segment. The second segment is
longer than the third one. The scaphocerite extends a trifle beyond the
antennular peduncle. It is nearly five times as long as its maximum breadth. The
lateral margin of the scaphocerite is straight and ends into a small spine, which
is shorter than the squamose portion. The length of the antennal flagellum
exceeds the total length of the animal.
219
Figure 101: Processa compacta. A- Left maxillula in dorsal view; B- Left
maxilla in dorsal view; C-E- Right first to third maxillipeds. (After Kazmi &
Kazmi, 1973 as P. edulis)
The mandible is elongated, slightly curved and without an incisor
process or palp; the molar process is toothed. The maxillule is as illustrated, the
inner lacinia of maxillule is wanting; the palp is rather characteristic; it terminates
in a curved, claw like process. In maxilla, the inner lacinia is rudimentary; the
palp is well developed; the scaphognathite is large, and of moderate width, the
posterior margin of the scaphognathite bears a bunch of very long setae. The
first and second maxillipeds are as illustrated. The third maxilliped exceeds the
scaphocerite by a little less than the two distal segments. The penultimate
segment is nearly half of the parapenultimate segment.
220
Figure 102: Processa compacta. A-First pereopod of right side; B-First
pereopod of left side; C- Second pereopod of left side. (After Kazmi &
Kazmi, 1973 as P. edulis)
The first pair of pereopods is asymmetrical; the right pereopod is
chelate while the left has a simple claw like dactylus; when closed the fingers
are slightly less than half of the palm. The second pereopods are sub-equal; the
chela of left second pereopod is longer and larger than that of the right, the
fingers of the chela are nearly as long as the palm, the carpus, merus and
ischium are sub-divided; the number of the segment varies not only in different
individuals but also on the two sides of the same specimen; the carpus of the
right pereopod is sub-divided into 25-32 joints, the carpus of left pereopod has
17-18 joints, and merus has 5 joints, while the ischium is unsemgented.
221
Figure 103: Processa compacta. A-Third pereopod; B-Fourth pereopod;
C-Fifth pereopod. (After Kazmi & Kazmi, 1973 as P. edulis)
The third pair of pereopods is symmetrical; the ischium bears two large
spines and is a trifle shorter than the merus; the merus bears a row of welldeveloped spines, it is nearly as long as the carpus; the fourth pereopod is
longer than the third or fifth; the propodus of the fifth pereopod is slightly longer
than the carpus.
222
Figure 104: Processa compacta. A- Appendix interna and appendix
masculina of male; B- Right second pleopod of female. (After Kazmi &
Kazmi, 1973 as P. edulis )
The endopod of the first pleopod is small and notched at the apex. The
male second pleopod bears both appendix interna and appendix masculina.
Remarks: One specimen from Gwader measuring 6mm in total length
possesses only six joints in the carpus of second leg. This does not seem to be
related to size since even the post larva of the species has as many joints as
adult (Jagadeshi&Sankolli,1977).
Colour: Live specimens are off white and without any significant colour
marks
Size: 18-33mm in TL
Localities: Manora Island; Cape Monze, Gwadar.
Habitat:Shallow water among weeds.
Distribution: Indo-Pacific and Atlantic
223
3.1.6 Family Ogyrididae Holthuis, 1955
Diagnosis: First two pairs of legs chelate, nearly equal in size and
not much if any longer than other legs, first pair shorter and stouter than the
second does. Carpus of legs subdivided. Rostrum small or wanting.
Eyestalks long, slender, fully exposed but with reduced cornea. Telson
thick, obtusely pointed. Blades of uropods curved outwards. Thelycum
present in females (After Hay & Shore, 1918).
Remarks:
Williams (1984) summarized scant information on
ogyridid biology. These shrimps burrow into soft bottom of estuaries and
near shore; habitats emerging at nightfall and as such are nocturnally
planktonic. Their transparency and setose appendages are adaptation for
their nightly plankton. Bruce (1986) added that the ecological niche
occupied by ogyridid species has not been precisely identified. The slender
body form suggests that they may live in semi permanent tubes and burrow
rather than leading a mobile life. In the present study several specimens are
available for study from samples collected from shallow water through
sediment grab, no tube was seen in the samples but almost all specimens
carried the very slender fifth leg in elevated position on carapace, this may
be involved in maintaining an optimum position in a permanent burrow.
Much detailed work is needed on the biology of this little known but
potentially important inhabitant of seashore waters (Bauer, 2004)
There is only one genus Ogyrides in the family.
Genus Ogyrides Stebbing, 1914
Diagnosis: All characters of family. Telson with a stridulating
mechanism.
Type species: By monotypy Ogyris orientalis Stimpson, 1860
Gender: Masculine
Remarks: There are 12 species in the genus. The genus being extremely
abnormal in type exhibiting both primitive and specialized characters. Two
species O. orientalis Stimpson and O. saldanhae Barnard are present in
Pakistani waters. They are distinguishable with the help of following key.
224
KEY TO THE PAKISTANI SPECIES OF OGYRIDES
1Carpal articles of second leg with five joint lets. The inner spine of
the lateral pair of spines of telson surpasses the tip of telson-------------------------O.? saldanhae.
-Carpal articles of second leg with four joint lets. The inner spine of the
lateral pair of spines of telson not very longer than the outer spine and does
not surpasses tip of the telson------------------------------------------------------------------O. orientalis
Ogyrides orientalis (Stimpson, 1860)
(Figs. 105-106)
Ogyris orientalis Stimpson, 1860:105:171; Liu, 1955: 34
Ogyrides orientalis Fujino & Miyake, 1970:255; Banner & Banner,
1978:247; 1982:293 (discussion); Tirmizi, 1980a:107; Holthuis, 1980:123;
Bruce, 1986:586; Kazmi, 2003:241; Grabe & Lees, 1995: 958
Ogyrides sibogae- de Man, 1922:14
Ogyridas orientalis Kazmi & Kazmi, 1979:155
Description: The carapace is smooth, glabrous. The rostrum is
very short, acute, twice as long as the width across base, depressed, tip
reaching well beyond the level of the inferior orbital angles in dorsal view,
with tuft of sub terminal setae ventrally: four postrostral median spines are
present on the anterior third of carapace: the orbital notch is feebly
developed, the margin is setose; inferior orbital angle is bluntly rounded, the
antennal tooth is absent; the pterygostomial angle of branchiostegite is
bluntly obtuse; cardiac notch is absent, posterior marginal carina is well
developed.
The abdominal somites are smooth, glabrous, with the pleura of the
first five somites broadly rounded, non-setose; the sixth somite is longer
than the fifth, strongly compressed, with feeble semi acute posteroventral
angle and small blunt posterolateral angle.
225
Figure 105: Ogyrides orientalis. Animal in lateral view
The telson is longer than sixth abdominal somite, the former is
ventrally curved, the lateral margins are moderately convergent; glabrous,
but with angular expansions at about 0.35 of the length; two pairs of
posterolateral spines are present at 0.75 of the length, one small lateral
spine and much larger medial spine, the posterior border is strongly
produced, the sides are convex , with acute median process with about 8-9
long plumose marginal setae ventrally, except at the tip; the dorsal surface
has two pairs of small spines; the anterior pair is closer to the midline than
the posterior pair; the ventral surface is without anal tubercles, three distinct
semicircular ridges run proximolaterally, the central ridge is confluent with
the lateral margin, the fourth present proximally is feebly developed.
The antennular peduncle distinctly exceeds the carpocerite, does
not reach distal corneal margin; the proximal segment is proximally stout,
with well developed statocyst, two strong sub equal teeth are present
laterally, exceeding half of the segment length, the distal segment is
slender, sub cylindrical; sparsely settees; the intermediate segment is sub
cylindrical, with few long setae, about half of proximal segment length; distal
segment is also half of proximal segment length; the upper flagellum is
uniramous, made up of about 5 slender segments, and only single distal
group of aesthetascs is present, the lower flagellum is slender, slightly
longer than upper, comprised of 6 slender segments.
226
A‟
F
A
D
E
B
G
C
Figure 106: Ogyrides orientalis. A- Anterior part, dorsal view; A’Front, lateral view; B- Telson; C- Thoracic sternites; D- Maxilla; ESecond Leg; F- Endopod of first pleopod of male; G- Appendix
masculina and interna
The antennal basicerite is stout, with a small distoventral tooth; the
carpocerite is sub cylindrical; the flagella are generally broken, when
present they are long; the scaphocerite reaches to near middle of
intermediate segment of the antennular peduncle, far exceeded by the
carpocerite, distal lamella is moderately acute and produced well beyond
the strong distolateral tooth.
The eyes are extremely long, mainly glabrous, few short setae are
present proximally, The eye length exceeds length of antennular peduncle
by the diameter of cornea; the cornea is globular, well pigmented, with
227
lateral accessory pigment spot, the peduncle is broadly expanded
proximomedially, and slightly expanded distally,
The third maxillipeds are long, slender extending beyond cornea,
with numerous distal plumose setae; coxa without medial process. The
exopods are well developed.
The first pereopod is relatively robust, equal, similar, reaching to
about level of distal carpocerite: the palm is slightly compressed, the fingers
are slender, about 1.5 times the palm length, with distal cutting edges entire
and small hooked tips; the carpus is a little longer than the chela, slightly
tapered proximally, the dorsal and ventral margins are sparsely setose; the
merus is about carpal length, sparsely setose ventrally; the ischium is
smaller than the merus length, sparsely setose ventrally; the basis is short,
without any special feature; the coxa is robust,
The second pereopods are slender, sub equal, similar, exceed the
cornea by the length of chela; the palm is slightly compressed, the fingers
are slender, about 2 times the palm length, the distal cutting edges are
entire and with small hooked tips; the carpus is four segmented, more than
twice the chela length; proximal segments are more slender; the merus is
smaller than the carpus, sparsely setose; the ischium is about half the
carpus length, slightly broadened distally and sparsely setose; the basis is
short, much smaller than the ischium; the coxa is slender.
The third pereopod is robust, reaches to about tip of the
scaphocerite; the dactylus is slender, small, sub cylindrical, with three long
contiguous terminal setae; the propodus is 3 times the dactylus length,
compressed, 5-6 long coarsely plumose setae and 4-5 short serrulate
dorsolateral and ventrolateral spines are present; the carpus is about 1.5
times the propodus length having long coarsely serrulate setae along
dorsal and ventral borders, and 5 short spines along both dorsolateral and
ventrolateral margins; the merus is longer than the carpus length, slightly
widened distally with a strong mobile distolateral spine, dorsal and ventral
margins with several coarsely plumose setae; the ischium is longer than the
merus length, feebly tapered proximally, the distal half is with two strong
mobile lateral spines, otherwise sparsely setose; basis is without special
features, about sub equal to carpus length; coxa is robust and normal.
The fourth pereopod is longer and more slender than the third,
extends to about distal end of the scaphocerite; the dactylus is sub
cylindrical, slightly curved, with few long simple setae distally, the propodus
228
is armed, with long coarsely serrulate setae along dorsal, dorsolateral
and ventral margins; the carpus is about 1.25 times the propodus length,
with long feebly plumose setae along dorsal and ventral margins; the merus
is unarmed, provided with plumose setae distodorsally and ventrally; the
ischium is unarmed, longer than the merus , with numerous simple setae;
the basis and coxa are without any special feature.
The fifth pereopod is slender, extends anteriorly to about middle of
the scaphocerite; the dactylus is subcylindrical, tapering slightly distally, with
single long finely serrulate terminal seta, about 1.5 times the dactlylus
length; the propodus is setose, more robust, shorter than the dactylus
length; the carpus is shorter than the propodus , the segment tapers
proximally, with several short plumose setae; the merus is sub cylindrical, ,
unarmed and sparsely setose; the ischium is about 3.5 times the propodus
length, sub cylindrical, sparsely setose ventrally, unarmed; the basis is
short,simple; the coxa is slender and elongated.
The thoracic sternites are narrow and unarmed except the fourth
which has a median anteriorly directed process extending between coxae of
third pereopods, straight, with two acute processes distally, separated by a
U-shaped notch, with a median tubercle at about the midlenght, few setae
are also noticed on them.
The second pleopod of the male has sub equal endopod and
exopod; the appendix masculina is short, sub cylindrical, with three medial
and one distal spine; the appendix interna is long and slender, twice the
appendix masculina length.
The protopod of the uropod is short and broad, with dorsomedial
process, the posterolateral angle is acute, the lateral margin has plumose
setae; the endopod is sub equal to the telson length, but shorter than the
exopod, it is broadest proximally and tapering distally to an acute point,
without posterolateral spines, the lateral border is feebly concave, strongly
setose, with longer stiff setae .
Remarks: Banner & Banner (1972) although did not enter the
discussion of synonymy but gave some indication that it is possible that O.
delli Yaldwyn and O. mjobergi (Balss) are synonyms of O. orientalis if this is
resolved then the known distribution of O. orientalis will be greatly extended
.
Size:4 mm in CL
229
Localities:Jiwani, Gwadar, and Port Qasim
Habitat: At 9-535m depth on sandy bottom.
Distribution: Indo- west Pacific: India, Indonesia, Philippines,
Hong Kong, China, and Japan.
Ogyrides? saldanhae Barnard, 1947
(Fig. 107)
Ogyrides occidenalis (non Ortmann) Stebbing, 1914:32
Ogyrides saldanhae Barnard 1947; 387; 1950: 726; Kensley, 1981:26
Description: The carapace is setose, having plumose setae along
lower margin, 3 or 4 minute spines are present anteriorly on middorsal ridge; the
rostrum is short and triangular extends to the end of the antennular peduncle.
Antennal spine is absent.
The eyes are shorter than the antennular peduncle but longer than the
antennal peduncle.
The telson has slightly sinuous lateral margin, a pair of spines is
present in a notch in the distal half of lateral margin, the longer spine is very
long, surpasses the apex of the telson, the dorsal margin is armed with two
pairs of spines, some curved ridges are present on the ventral surface at the
base.
230
Figure 107:
Ogyrides? saldanhae. A- Carapace, lateral view; BTelson, dorsal view; C- Eye and antennule; D- Antennal scale; E-IFirst to fifth legs; J- Pleopod; K- Uropod.
The stylocerite ends in two slender spines, the inner being shorter. The
antennal scale is lanceolate, with straight outer margin; the inner margin is
obliquely beveled off from the apical point. The antennal peduncle fails to reach
the end of antennular peduncle.
The propodus of the first leg has a patch of strong spines on the lower
margin. It has fingers shorter than the palm; the carpus is slightly longer than
231
the chela. The carpus of second leg is divided into 4 segments. The third
leg is armed with a strong spine on the lower margin of merus, the dactylus of
third and fourth legs are thin plate like, elongate, narrower in the fourth than in
third leg, apically the third ends in a setule, fourth in to an unguis and two
setules. The dactylus of fifth leg is ensiform and provided with long spaced
plumose setae.
A bifurcated sternal plate arises between the basis of third legs, a pair
plumose setae arises from the anterior margin of the bifurcation.
Remarks: The present specimens are close to O. saldanhae with few
differences.
Size: 5-7mm in TL
Localities: Gwadar and Port Qasim
Distribution: South Africa, South East Atlantic and for the first time
from the Arabian Sea coast of Pakistan.
232
3.1.7 Family Alpheidae Rafinesque, 1815
Diagnosis: Rostrum short or absent, never spinose. Carapace smooth
and always provided with cardiac grooves. Antennal and pterygostomian spines
always absent, sometimes a supraorbital spine present. Eyes partially or wholly
covered by an anterior projection of the carapace. First pair of pereopods
usually strong, robust and chelate, often asymmetrical, especially in male.
Carpus of second pereopod divided in 3-5 segments. Last three pairs of
pereopods short, compressed, with propodi spinose, dactylii with one to three
ungui. Propodus of fifth pereopod bears a more or less well developed” brush”
of bristles arranged in transverse, oblique rows on its posterior margin. (From
various sources).
Remarks: The total biodiversity of this family is relatively unknown as
its taxonomy is difficult, with 46 genera and 600 valid species species, the
Alpheidae is a diverse family inhabiting shallow water coastal habitat except of
one genus, which lives in fresh to brackish water in mangrove swamps; there
are a few truly fresh water species.
1.
Some members of the Alpheid genera are among the smallest known
caridean shrimp with maximum TL of adults around 10mm or even less and can
be called microshrimps. Several crypto- species complexes in Alpheidae need
revision (Anker, 2001).
The Alpheidae genera Athanas, Automate, Salmoneus, Alpheus, Synalpheus,
and Alberta represent the Alpheidae in Pakistan. They can be separated with
the help of the following key. An undetermined genus close to Leptalpheus from
Gwadar is also included in the report.
233
KEY TO THE PAKISTANI GENERA OF ALPHEIDAE
1. Corneas of the eyes at most only partially concealed from anterior or
lateral view by carapace; dactylus of large chela without plunger fitting into
socket of fixed finger and often bearing teeth------------------------------------------------------------------------------------------2.
Corneas of the eyes entirely covered in dorsal, lateral, and anterior
view by orbital hoods; dactylus of large chela usually with a plunger that fits
into a socket of the fixed finger, never serrated---------------------------------------------------------------------------------------4.
2. Corneas of eyes reduced and entire eye stalk exposed ---------------Genus Automate .
Corneas of eyes usually well developed. Peduncles of eyes covered-------------------3.
3. Corneas of eyes exposed in dorsal and lateral view, infra and extra
corneal spines present. Posterior margin of telson without any notch------Genus Athanas
Corneas largely or entirely concealed in dorsal view, infra and extra
corneal spines absent; posterior margin of telson with “V” shaped notch----------Genus Salmoneus
4. Pterygostomian angle produced into a definite angle. Transverse and
longitudinal grooves absent from palm of large and small chela of first pair
of pereopods---------------------------------------------------------------------------------------- Genus Synalpheus -Pterygostomian angle rounded. ----------------------------------------------------------------------- 5.
5. Transverse and longitudinal grooves present on palm of large and
small chelae of first pair of pereopods------------------------------------------------------------------- Genus Alpheus.
No groove on the palm of chelipeds---------------------------------------------------Genus Alberta.
Genus Alberta new genus
234
Diagnosis: Carapace smooth, pterygostomial angle rounded.
Frontal margin without rostrum and orbital teeth. Eyes not visible in dorsal view.
Telson with two pairs of dorsal spines. Stylocerite not reduced. Antennal scale
with lateral tooth hardly overreaching margin. Mouth parts typical for family,
Mandible with two-jointed palp. Third maxilliped distally unarmed, with serrated
setae on medial margin of ultimate segment and acute lateral plate above
epipod. chelipeds similar, sub symmetrical folded back, cutting edges of fingers
with plunger and notch. Second leg short, carpus four segmented. Uropodal
diaraesis well marked.
Affinities: The shape of the chela and body shows that this specimen
belongs to those genera, which have the well-developed plunger at basal
portion of the finger and lack orbital hoods like Alpheus, Synalpheus ,
Metalpheus, Orygmalpheus, and Pomagnathus. It is difficult to place Alberta
unequivocally in proximity of any other alpheid genus.
Type species: By original designation Alberta banneri, new species, by
monotypy
Etymology: The new species is named after the late Dr. Albert H.
Banner of the Hawaii Institute of Marine Biology,Hawaii.
Alberta banneri n.sp.
(Figs. 108-109)
Material Examined: Holotype, ovigerous female 14mm in TL, 8mm in CL.
Type Locality: Bulleji, Karachi
Description of Holotype: The carapace, which completely conceals the
eyes, is smooth and unarmed, its frontal margin is produced in a broadly
triangular rostrum with a pointed and slightly deflected tip, reaching almost
to the middle of the second segment of the antennular peduncle;
pterygostomian angles are rounded.
235
C
A
E
D
B
Figure 108: Alberta banneri. A- Anterior part; B- Telson and left uropod ;
C- First leg, outer view; D- Same, inner view; E- Second leg
The antero-lateral margin of the first abdominal somite overlaps a
small portion of the carapace. An articulated plate is present on the posterolateral angle of the sixth abdominal somite. The telson is 2.7 times as long
as the posterior margin, the posterior margin is slightly convex, there exists
at ratio of 1:1:7, between anterior and posterior margin. In addition to the
two pairs of dorsal spines, each postero- lateral is armed with two sub-equal
spines. A row of small spines can be seen on the posterior margin.
The antennular peduncle is short and stout, with sub-equal
segments. The stylocerite is well developed, reaching as far as the distal
236
margin of the second segment of the antennular peduncle; the
scaphocerite extends up to the distal margin of the antennular peduncle; the
spine on the antero-lateral angle is small, not extending beyond the
squamose portion; the carpocerite is longer than the antennular peduncle; a
small but acute spine is present on the basicerite.
The incisor process of the mandible bears six teeth; the central
teeth are larger; the palp consists of two distinct segments; the maxillule has
lower lacinia slender; the upper is broad, the palp is deeply incised; the palp
of the maxilla is short and simple. The maxilla is typical.
Figure 109: Alberta banneri. A-Right antennular peduncle; B- Right
antennal scale; C- Mandible; D- Maxillula; E- Maxilla; F-H- First to third
maxillipeds
All the maxillipeds have well developed exopods; the palp of the
first maxilliped is longer than the caridean lobe; the ultimate segment of
second maxilliped is laterally attached with the penultimate segment; the
ultimate segment of third maxilliped is nearly half of the parapenultimate
segment; exopod reaches beyond the end of the parapenultimate segment;
each pereopod is 13mm long (coxa to dactylus) (chela = 5mm; merus =
3mm; ischium= 4mm). The pereopods of the first pair are symmetrical; the
237
chela is large, its palm is considerably swollen, a low oblique
tuberculated ridge extends dorsally on the proximal half of the palm, when
viewed from ventral view; a row of small tubercles can be seen on its inferior
surface and a few long setae on the distal half of the superior margin, as
illustrated; the fingers are 2.8 times as long as the maximum breadth; the
immovable finger is curved and with a strong tooth in the middle of the
cutting edge which fits in a notch on the fixed finger; the inferior margin of
the carpus is depressed and serrated; the combined length of the merus
and ischium is little more than the rest of the cheliped, an excavation runs
on the inferior surface of each and it is for the accommodation of the chela,
when flexed; the interio-external margin of the merus is serrated; in the
middle it enlarges and forms a prominent tooth; the ischium is slightly longer
than the merus, its superior margin bears 9-10 teeth, the interio-internal and
external margins are serrated; a small spine is present on the coxa; the
carpus of the second pereopod is four -segmented, the segments have the
following ratio 5:1:1:2.2
The third pereopod is relatively longer and stronger than the fifth,
the ischium is one third of the merus, bearing one spine on the internal
margin, the merus is 4.1 times as long as broad at its maximum breadth, the
carpus is slightly longer than the ischium bearing single feeble spine on the
interio-distal margin, the propodus bears three spine and it is shorter than
the merus; the dactylus is equally long to the width of the propodus breadth.
Unfortunately, the fourth pair of pereopods is missing. The carpus of the fifth
pereopod bears one spine at its inferior margin; the propodus bears two
spines: one on the proximal side and the other on the distal end. The
transverse suture of the exopod of the uropod bears a row of 20 spines
Remarks: Although only one specimen is at authors disposal, but
its peculiar combination of characters inspired us to describe it as new.
Nevertheless, there are chances that it is some variation of the already
recorded species.
Colour: Off white (when fresh).
Etymology: It is our pleasure to name this species for late Prof. Albert H.
Banner of Hawaii Institute of Marine Biology, Hawaii.
Genus Alpheus Fabricius, 1798
238
Diagnosis: Carapace anteriorly developed into orbital hoods,
which completely enclose eyes except on ventral side. Orbital hoods frequently
with spiniform apex and usually demarked from rostral base by more or less
pronounced depressions. Rostrum usually present and carapace often with at
least slight rostral carina. Pterygostomian margin of carapace rounded.
Antennule usually short, frequently with basal peduncular article and stylocerite
reduced. Scaphocerite frequently reduced; basicerite either armed with
inferolateral spine or rounded; carpocerite usually reaching to or beyond the end
of scaphocerite.Chelae of first legs of very asymmetrical development. Large
chela of variable form, from sub conical to compressed and twisted, surfaces
from smooth and entire to deeply sculptured with grooves and pronounced
“alpheopsidean” lobes.Dactylus usually with piston like process that fits into a
cavity on fixed finger and always with digital and palmer adhesive plaques.
Chela always carried extended from body. Large cheliped with hemispherical
carpus, with merus at least triangular in section. Small chela in simple form; at
times showing marked sexual dimorphism. Carpus of second leg of five articles
of variable proportions. Third and following legs robust, compressed; armature
and proportions of merus, carpus, and propodus variable, dactyls simple or
biunguiculate.Fifth legs with “brush” on propodus. Abdomen usually without
lateral compression, with pleura in females larger than those of males and
usually without acute projections on margins. Pleura of sixth abdominal segment
not jointed. Telson usually with posterior margin convex and lateral angles
distinct but not acute. Anal tubercles almost always well developed. (Adapted
from Banner & Banner, 1966).
Type species:
Alpheus avarus Fabricius, 1798
Gender:
Masculine
Remarks: The genus Alpheus or pistol shrimps are well known for their
production of sound underwater. The species are found in four major habitatson living and dead corals, in tide pools, in burrows in sand and muddy substrate
or under rocks, as symbionts with some invertebrates like anemones, echinoids,
annelids, and fish (Banner & Banner, 1966). Some particular features like the
subspatulate dactylus on third to fifth pereopods show the burrowing life style.
There are over 330 species and 10 subspecies in the genus. To facilitate this
cumbersome and large taxonomic unit the genus is subdivided earlier in five
groups by Coutiere (1894) and later the group crinitus into three subgroups.
Banner & Banner (1966) raised these subgroups to the group rank. Crosnier &
Forest (1966) made some changes. The grouping into seven groups that is
followed now is by Banner & Banner (1982): Brevirostris group, Crinitus group,
Diadema group, Edwardsii group, Macrocheles group, Obesomanus group and
239
Sulcatus group however, Chace, 1988 has not clearly accepted these
groups. Some of these groups are clearly not monophyletic.
This genus is represented in Pakistan by the following species, Alpheus
albertii, , Alpheus cf. barbatus,Alpheus bisincisus, Alpheus chiragricus ,Alpheus
edwardsii, , Alpheus isodactylus, Alpheus lobidens, Alpheus manorensis ,
Alpheus pacificus, Alpheus pseudoedwardsii ,Alpheus cf rapax ,Alpheus
splendidus , Alpheus strenuus strenuus, and Alpheus zulfaquiri. ,
Each of these species is associated with any of the above given
groups. Only the Pakistani species groups are characterized below (Characters
modified from Chace, 1988) and species found from Pakistan are listed under
the group to which the species belong. The critical information is lacking for A.
manorensis,and A. isodactylus and the type material is also of unknown
whereabouts but included here for the sake of completeness however, we were
able to assign them to one of the seven groups.
Brevirostris Group
Orbital teeth lacking, orbital hoods often prominent; major chela with
palm always compressed, more or less quadrangular in cross-section, often
with surfaces delimited by distinct angles, with or without “saddle” proximal
rd
to adhesive plaque; minor chela sometimes “ balaeniceps” in male; 3
pereopod with dactyl always simple, sometimes subspatulate, merus usually
unarmed on flexor margin. (Composition: A.cf barbatus, A. cf rapax, A.
zulfaquiri).
Crinitus Group
Many species of this group live in algal tubes or in groups.
Diadema Group
Intertidally to sub tidally, most of them confined to dead corals in
clean water and moderate wave action.
Edwardsii Group
Orbital teeth lacking except in 1 species; major chela with palm
compressed, with “saddle” proximal to adhesive plaque and usually with
shoulder on opposite margin proximal to fixed finger, “saddle: usually
extending on to both adjoining surfaces as triangular or quadrangular
rd
240
depressions; minor chelae often “balancipeses” in male; 3 pereopod
with dactyl usually simple, sometimes subspatulate, merus usually dentate
on flexor margin (Composition: A. lobidens A. chiragricus, A. bisincsus. A.
pacificus, A. strenuus strenuus, A. edwardsii, A. psudoedwardsii, A.
isodactylus, A. albertii, and A. manorensis)
Most species in this group are found in burrows in sandy or silty
bottom often constructed under rocks lying on the substrate, they frequently
penetrate in to brackish water.
Macrocheles Group
Many of the species of this group are only known from deeper
waters, the ones occurring in inter to sub tidal region seem to characteristic
of clean habitats.
Obesomanus Group
Some and perhaps most species live in protected galleries in the
crust of coralline algae.
Sulcatus Group
Rostrum sometimes with base flattened and delimited from
adrostral furrows; orbital teeth often present; major chela with palm never
markedly compressed, usually with longitudinal but without transverse
rd
grooves; minor chela never “balaeniceps” ; 3 pereopod with dactyl simple
or biunguiculate, merus with or without tooth on flexor margin (Composition:
A. splendidus)
Most species appear to occur in dead or living coral.
Alpheus albertai Kazmi, 1971
(Fig. 110)
241
Alpheus albertai Kazmi & Kazmi, 1979:154;1989:48(List only)
Description: The rostrum is very short triangular. It reaches up to the onefourth portion of the visible part of the basal segment. The frontal margins of
the orbital hoods are somewhat pointed. The carapace and abdomen are
finely pubescent.
C
A
E
B
D
Figure 110: Alpheus alberti. A- Anterior portion only left side shaded;
B- Telson and uropods; C- Left first leg; D- Right first leg; E-Left
second leg
The telson is slightly longer than the last abdominal somite. The
dorsal surface is armed with two pairs of posteriorly directed spines. The
242
posterior lateral angle of the telson is armed with one pair of spines; the
outer spine is twice as long as the inner spine.
The antennular peduncle consists of three sub-equal segments.
The visible part of the basal segment of antennular peduncle is equal to the
second segment; the stylocertie reaches up to the distal end of basal
segment; the second segment is twice as long as the third segment. The
scaphocerite has concave lateral margin, which ends into a spine. The
spine is curved inwards and it is slightly longer than the squamose portion.
The oral appendages are typical alpheid type.
The first pair of pereopods is asymmetrical. The larger chela is 2.5
times as long as broad; fingers occupy one fourth portion of the chela;
longitudinal and transverse grooves are present on palm as illustrated; the
small chela is 3.5 times as long as broad in the middle. The fingers are
slightly shorter than the palm. The postero-inferior angle of the merus bears
a well-developed spine.
The second pair of pereopods is symmetrical; the carpus is
five segmented, first and second segments are nearly of equal size; the
propodii or the last three pairs of pereopods bear spines. The dactylii are
spatulate.
Size:
28mm in TL
Locality:
Bulleji
Distribution:
Not reported outside Pakistan
Alpheus cf. barbatus Coutiere, 1897
(Fig. 111, Pl.5 E)
Alpheus barbatus Coutiere, 1897:235; 1899:230; de Man, 1911:387; Banner
& Banner, 1979a:25; 1981:8; 1982:163; Chace, 1988:16; Anker et al,
2007:21
Description: The body is compressed, sparsely setose; the rostrum
is short, sub rectangular, not reaching nearly as far as distal margin of first
antennular segment, it is slightly carinate in the midline, the carina does not
243
extend posteriorly beyond orbital hoods, its base does not abruptly
delimit from adrostral furrows. The carapace is without median tooth or
tubercle on the gastric region; the anterior margin is concave and unarmed
mesial to orbital hoods, the orbital hoods are unarmed, adrostral furrows are
shallow.
Figure 111: Alpheus cf. barbatus. A- Rostrum in lateral view; A’
Rostrum, dorsal view; B- Abdominal pleurites, lateral view; C- Telson
and right uropod; D- Antennular peduncle; E- Antennal scale, dorsal
view; E’- Same, lateral view; F- Third maxilliped; G- Large chela, inner
view; G’- Same, outer view; H- Small chela; I- Second leg J- Third
leg; K- Appendix masculina and appendix interna.
244
The telson is 2.7 times as long as posterior margin is broad. The
dorsal spinules are small and located in posterior third.
Second antennular segment is only slightly longer than wide,
stylocerite is truncated distally not reaching distal margin of first segment;
basal antennal segment (basicerite) is unarmed; The antennal scale with
lateral
margin
nearly straight, distolateral spine
unusually stout,
overreaching distal margin of blade.
Third maxilliped reaches to ultimate segment about two times as
long as penultimate, penultimate segment is broad distally.
First pereopod has an unarmed merus on flexor margin. The major
chela is noticeably compressed laterally, about twice as long as wide, the
dactylus is single ended, superior margin of palm with narrow and curving
transverse groove, palm has shallow longitudinal sulcus on lateral surface
extending from base of fixed finger to mid length of palm. The minor chela
is about 2 ½ times as long as wide, fingers are gaping, longer than palm,
lateral surface of dactylus and mesial surface of both fingers bear dense
fringes of fine setae filling gap, the proximal carpal article of second
pereopod is longer than second; fingers of chela are longer than palm.
Third pereopod dactylus is curved pointed, and simple, the propodus bears
five spines on the flexor margin; the carpus, merus and ischium are
unarmed on flexor margin. All the segments except propodus are sparsely
setose. Exopod lateral margin ends in an acute immovable tooth and spine,
the latter internally flanked by rounded lobes.
Remarks:
The present specimens belong to pan tropical A.
barbatus Coutiere, 1897 species complex (A. barbatus clade) of Anker et
al(2007), also is reported an association between A. barbatus and echuiran
Ochetostoma in the western Pacific by the same authours. For the
Pakistani specimens no such observation was made, although the same
echuirian genus also occurs here. A. barbatus species complex is
distributed in Indo-Pacific and Atlantic and is included in brevirostris group
of Alpheus which is large but relative homogenous species group, contains
over 30 described species (Bruce, 1994).
245
According to Anker et al (20 07), the A. barbatus
complex needs further phylogenetic, ecological, and taxonomic studies; they
expect further cryptic taxa in the Indo-Pacific. It is hoped that new collection
of the species will enable us to ascertain the status of the Pakistani
specimens. For the time being they are being described here as A.cf
barbatus.
Our A. barbatus resembles A. barbatus described by
Banner & Banner (1981, 1982) and Chace (1988) except that the ischium of
third leg being without a spine. The propodus is not only shorter than merus
when compared to that described by Banner & Banner (1982) and Chace
(1988), but shorter length of propodus allows to accommodate only three
spines on anterior margin, consequently the ratio between the propodus and
dactylus is not that given by Banner & Banner (1982), Kim & Abele (1988)
and Anker et al (2007). Pakistani specimens differ from A. aequus having
narrow large cheliped and its movable finger. We considered establishing a
new species (Kazmi & Kazmi, 2008) based on these differences but A.
Anker of BIMM (France) suggested (pers. comm.) to avoid to do so.
Colour: Mostly ivory whitish with orange tinge in some areas
Locality: Pacha
Distribution: Djibouti, Red sea, Maldives, Indonesia, Philippines,
Taiwan, Japan, East Africa, Australia, and Christmas Island and for the first
time from Pakistan.
Alpheus bisincisus de Haan, 1850
(Figs. 112-113)
Alpheus bisincisus de Haan, 1850: 179, pl. 45, fig. 3; Tiwari,
1963:304; Barnard, 1950:760; Banner & Banner, 1966: 125, fig. 46;1979;
26; 1982:263; Yang , Ko,&Kim, 2007:37.
Alpheus bisincisus variabilis de Man, 1911:405,pl. 22.fig. 95
Description: The tip of the rostrum reaches up to the middle of the
visible portion of the first antennular segment. It is flattened on top. The
base of lateral margins overhang the orbito rostral grooves and extend
posteriorly well past corneas. The frontal margins of orbital hoods are
246
flattened anteriorly and are separated from the rostrum by notch on
anterior margin.
Figure 112: Alpheus bisincisus. A- C- Left first to third maxillipeds; DRight first leg; E- Left first leg; F- Left second leg
The telson is larger than the sixth abdominal somite. It tapers
towards the posterior end. Each postero-lateral angle is armed with one pair
of spines; the outer spine is shorter.
The antennular peduncle consists of three sub-equal segments.
The second segment is twice as long as broad, in the middle. It is slightly
longer than the visible part of the basal segment and nearly two times as
long as third antennular segments. The spine of stylocerite reaches up to
the end of first antennular segment. The outer margin of scaphocerite is
slightly concave and ends into a spine, which is longer than the squamose
portion and reaches up to the end of the antennular peduncle. Carpocerite
is slightly longer than the antennular segment.
247
The oral appendages are as illustrated.
The large chela is 2.3 times as long as broad. A transverse deep
groove with proximal edge projects and overhangs, forming a sharp tooth.
The groove is continued on outer face as a shallow well-defined triangular
depressed area extending downwards beyond the middle of palm and is
continued on inner face as smaller depressed area. The dactylus is heavy,
narrow, and slightly twisted. The merus is 2.8 times as long as broad with
inferior internal margin armed by strong and sharp tooth.
Figure 113: Alpheus bisincisus. A- Telson and left uropod; B- Left
antennular peduncle; C- Left antennal scale; D-Left maxillula; E- Left
maxilla.
The small chela is 4.2 times as long as broad; fingers are slightly
shorter than the palm. The upper margin is provided with transverse groove
proximal to dactylus, which is continued into outer face as a small triangular
depressed area. The dactylar articulation is flanked on inner side by small
projection. The dactylus is broad and of balaeniceps shape.
The carpus of second pereopod is five segmented and has the
following ratio: 10:7:3:3:5. The ischium of third pereopod is unarmed. The
248
merus of third leg is 6.3 times as long as broad. The carpus is one
fourth of the merus both the margins are distally slightly projected, the
propodus is 0.7 times as long as merus and armed on its inferior margin
with 11 movable spines, the dactylus is simple and one third of propodus.
Colour: Reddish with white legs joints.
Size:
36mm in TL.
Habitat: Below stones or sponges, not burrowing at depth 1 to at
least 70m.
Locality: Manora Island
Distribution: Gulf of Suez, Sri Lanka, Indonesia and South Africa,
Singapore, Korea, Japan, Laxshadweep and for the first times from
Pakistan.
Alpheus chiragricus H.M. Edwards, 1837
(Figs.114A,B)
Alpheus chiragricus: H.M. Edwards, 1837:354, Coutiere, 1905: 912,
Pl. LXXXVI, fig. 51; Banner & Banner, 1982: 267; Chace, 1988:18,
Description: The rostrum reaches up to the middle of the first
segment of the antennular peduncle. The tip of rostrum is downward. The
carapace is smooth and without any carina and groove.
The telson is longer than the sixth abdominal somite. It is nearly 2.5
times as long as the posterior margin. The dorsal surface is armed with two
pairs of posteriorly directed spines; each postero-lateral angle bears one
pair of spines.
249
Figure 114 :A- Alpheus chiragricus. A- Anterior part; B- Telson and
uropods; C- Left antennular peduncle; D- Left antennal scale; E- Right
maxillula; F- Left maxilla
The antennular peduncle consists of three sub-equal segments; the
second segment is slightly longer than the visible portion of the first
segment. The second segment is 2.4 times as long as broad, the stylocerite
reaches up to the end of first antennular segment; the lateral margin of the
scaphocerite is slightly concave and ends into a spine, which is longer than
the squamose portion. The carpocerite is nearly equal to the spine of
scaphocerite.
The oral appendages are as illustrated.
The large chela is 2.5 times as long as broad in the middle. The
finger is one third of the chela; the shoulder of the palm is pointed; the
250
inferior angle of the merus bears a strong spine and it is 1.5 times as
long as broad at the distal side.
Figure 114B: Alpheus chiragricus. A-C First to third maxillipeds; DLarge cheliped; E- Small cheliped; F- Second leg.
The small chela is 4.1 times as long as broad in the proximal region;
the fingers are slightly longer than the palm. The merus is 2.8 times as long
as broad in the distal region. The fingers of the second pereopod are nearly
equal to the palm, the carpus of the second pereopod is five segmented and
has the following ratio of 7:4:2:2:3.
The ischium of third leg bears a movable spine. The propodus
bears 8-9 spines. The dactylus is nearly one four times shorter than the
propodus.
Locality:
Size:
Manora Island
38mm in TL
251
Distribution: Hong Kong, Mergui Archipelago, Eastern Africa,
Madagascar, Bay of Bengal, Indonesia and Australia and now from the
Arabian Sea.
Alpheus edwardsii (Audouin, 1826)
(Figs.115-116)
Athanas Edwardsii Audouin, 1826:91
Alpheus audouini Coutiere,1905:911,pl87:fig,52; Holthius & Gottlieb,
1958:41; Banner & Banner, 1966:135; Afzal et al, 1986:338
Alpheus edwardsi Banner & Banner,1973:1142, 1979:26; 1982:270, fig, 1;
Kazmi & Kazmi, 1979:154; Miya, 1984:93;Corfeild&Alexander,1995:675
Alpheus edwardsii, Barnard, 1950:759; Chace, 1988:25.
Description: The rostrum is long and acute, reaching up to the three fourth
of the visible part of the basal segment of the antennular peduncle; the
rostral carina reaches to the base of the orbital hoods; the orbitorostral
grooves are narrow and shallow; the lateral spine of the stylocerite reaches
definitely beyond the basal segment of the antennular peduncle; the outer
margin of the scaphocerite is slightly concave with the lateral spine being
slightly larger than the antennular peduncle and the squamose portion is as
long as the antennular peduncle; the carpocerite is as long as the
antennular peduncle; the lateral spine of the basicerite is well developed.
252
Figure 115: Alpheus edwardsii. A- Anterior region; B- Telson and
uropods; C- Large chela; D- Small chela; E- Second leg
The telson is two times as long as posterior margin is broad, spines
on the dorsal surface are small; the outer pair of terminal spines is as long
as the dorsal spines, the inner spines are a little longer.
The large chela is 2.7 times as long as broad with the fingers
occupying the distal 0.4 of the chela. The proximal shoulder of the superior
transverse grooves and the inferior shoulder are in the form of acute spines
overhanging the depressions. The superior groove is continued on the inner
face as a channel, which runs posteriorly near the superior margin
extending to the half of the palm, on the outer face the depression is
continued as a quadrangular area, the external depression of the inferior
shoulder is continued insignificantly on the inner and the outer face. The
dactylus is broadly rounded as the tip and is definitely shorter than the fixed
finger. The merus is 1.8 times as long as broad and carries strong acute
tooth on the distal margin of the inferior inner surface.
The small chela is sexually dimorphic in females; it is 4.4 times as
long as broad with fingers occupying the distal half. The palm is very
253
smooth. The dactylus is simple without fringe of setae. The meri of both
the sexes are 2.2 times as long as broad and bear an acute tooth distally on
inferointernal margin. The male small chela is 3.8 times as long as broad
with fingers 0.6 as long as palm, the superior margin of palm bears a small
groove proximal to the dactylus that is extended slightly.
Figure 116: Alpheus edwardsii. A- Left maxillula; B- Left maxilla; C- ELeft first to third maxillipeds.
254
Remarks:
The present material does not agree fully with
Henderson‟s description of A. edwardsii collected from Karachi coast in
1883. This is either due to the individual variation since the species is
variable; at least six characters show variation (Banner & Banner, 1972) or it
is a different species.
Colour: Ground colour is purplish brown, the dorsal halves of
abdominal somites bears 5 longitudinal rows of white or colourless short
stripes and some blue eyes spots- three on the first somite, two, four, four,
and two spots respectively on following somites
Size:11 mm in CL
Localities:Manora Island, Astola Island
Distribution: East Mediterranean evidently entered by way of
Suez Canal, Red Sea, and eastern and South Africa to Thailand,
Philippines, Indonesia, Australia, Marshall Island, and Caroline Island.
Alpheus isodactylus Afzal,Javed&Barkati, 1986
(Fig. 117)
Alpheus isodactylus Afzal et al, 1986: 339
Description: The rostrum is triangular reaching up to the middle or to the
end of the visible part of the basal segment of the antennular peduncle; a
notch is present between the rostral base and the anterior margin of the
orbital hoods; the rostral carina is acute and runs up to the base of the
orbital hoods; it is separated from the orbital hoods by broad shallow
grooves.
The telson is longer than the sixth abdominal somite. The two pairs
of dorsal spines are always present. The posterior margin bears two spines
on each posterolateral angle and a row of 12 to 20 spinules.
255
Figure 117: Alpheus isodactylus. A- Anterior region; B- Telson and left
uropod ;C- Large chela, male; D- Small chela, male; E- Second right
leg; F- Third right leg.
The lateral spine of the stylocerite is short and reaches to the end of
the basal segment of the antennular peduncle; the outer margin of the
scaphocerite is clearly concave, the lateral spine almost always reaches
beyond the antennular peduncle, with the squamose portion slightly longer
than the antennular peduncle, but always shorter than the lateral spine; the
carpocerite is almost equal in length to the antennular peduncle. The lateral
spine of the basicerite varies from reduced to well develop.
The large chela is 2.0 to 2.3 times as long as broad, with the fingers
occupying the distal one third; the dactlyus is broadly rounded at the tip in
both the sexes. The anterior margin of the tooth on the dactylus is abruptly
curved posteriorly in the male while in the female it gradually does so; the
groove of the superior margin of the palm is with proximal and distal
256
margins broadly rounded. The distal depression on the inferior shoulder
is well marked and continued superiorly on the inner surface to join the
extensive external midline; the merus is 1.5 to 2.0 times as long as broad,
and is unarmed.
The small chela shows the sexual dimorphism; that of the male is
balaeniceps-shaped, 2.6 times as long as broad. The transverse groove of
superior margin, depression on the palm and the inferior shoulder are
similar to the small chela of the male, far less pronounced but present in the
female specimens of all available sizes. The merus is almost 2.0 times as
long as broad, unarmed.
The second leg has the ratio of the carpal articles: 10:6:2:4. The
ischium of the third leg is with a movable spine. The merus is unarmed, 3.0
as long as broad, bearing 3 to 5 very much elongated stiff setae on the
superior margin of the anterior half besides many short scattered setae on
the inferior and the superior margins. The carpus is 0.6 as long as the
merus with either distal margin markedly projecting. The propodus is almost
as long as the carpus, bearing on its inferior margin 8 to 12 movable spines
of which the two are in the form of a pair at the base of the dactylus; the
dactylus is simple.
The endopod of the second pleopod in female bears only an
appendix interna. In males both the appendix interna and appendix
masculina are present and are almost of equal size. The tip of the appendix
interna bears a cap of tiny hooks.
Size:
10-48mm in TL (male) 8-50mm in TL (female)
Locality:Manora Island
Distribution: Not reported outside Pakistan
Alpheus lobidens de Haan, 1849
(Figs. 118-121, Pl.4B)
Alpheus lobidens de Haan, 1849:179; Kazmi & Kazmi, 1979:153; Banner &
Banner, 1981:29; Chace, 1988:34
257
Alpheus crassimanus Heller, 1862: 526; 1865:170;Bate, 1888:554; de
Man, 1902:880; Kemp, 1915:299; Barnard, 1950:756; Banner & Banner,
1959:147; 1966:138; Tiwari, 1963:307 (part); Tufail & Hashmi,1965:278
(larvae)
Alpheus inopinatus Holthuis & Gottlieb, 1958:42; Tirmizi& Kazmi, 1969:99;
Karim & Rehman, 1974:26; Afzal et al, 1986:338
Alpheus lobidens polynesica Banner & Banner, 1975:429
Alpheus lobidens lobidens Banner & Banner, 1982:252
Description: The carapace and abdomen are finely pitted. The rostrum is
short and sharply pointed. It fails to reach the distal end of the basal
segment of the antennular peduncle. Posteriorly it is continued as a welldefined carina as far as the base of the eyes. This carina is separated from
the ocular hoods by distinct grooves. The rostrum is smooth and may have
hairs. The posterior margin of the carapace has a distinct cardiac notch.
Postero-lateral angle of each of the first four abdominal somites is broadly
rounded that of the fifth is somewhat pointed. The antero-lateral angle is
broadly rounded.
258
Figure 118: Alpheus lobidens. Animal in dorsal view
(After Tirmizi & Kazmi,1969)
The telson is longer than the sixth abdominal somite. There are two
pairs of dorsal spines on the telson; the first pair is placed somewhat before
the middle of the telson, the second is situated slightly closer to the anterior
pair than to the posterior margin of the telson, some times one or (seldom)
two spines may be missing. The posterior margin is furnished with two
spinules on each postero-lateral angle and a fringe of setae.
The antennular peduncle consists of three sub-equal segments.
The stylocerite ends in a sharp point, it fails to reach the distal margin of the
basal segment. The second segment is longer than the last or ultimate
segment, the lateral margin of the scaphocerite is somewhat concave and
ends into a strong spine, which is longer than the squamose portion.
259
The mandibular palp is two segmented. Both the incisor and
molar processes are well developed; the incisor process bears 8-9 teeth.
The maxillulary palp is divided into a broad rounded outer and a slender
inner portion, the latter is provided with a strong bristle, the upper lacinia is
slender and distally curved. The maxillary palp suddenly becomes narrow
distally.
The palp of the first maxilliped is two segmented and reaches well
beyond the caridean lobe .The second maxilliped is as illustrated. The third
maxilliped reaches with the ultimate segment beyond the antennular
peduncle, its tip bears long setae, the ultimate segment is twice the length
of the penultimate segment, and the parapenultimate segment is three times
longer than the penultimate segment. The exopod reaches slightly beyond
the end of the parapenultimate segment.
260
Figure 119: Alpheus lobidens. A- Left antennular peduncle; B- Left
antennal scale; C- Left mandible; D- Left maxillula; E- Left maxilla.
The first pereopods reach with the chela beyond the antennal
peduncle, the larger leg has the upper margin of the dactylus evenly
rounded, forming almost semicircle, the ventral margin bears a very strong
blunt tooth, which fits in the cavity on the cutting edge of the fixed finger, the
palm is somewhat less than twice as long as the finger. Both the upper and
lower margins are provided with a distinct broad, blunt shoulder, the lower
incision is rather developed than upper, and continues on the outer surface
as an oblique groove, the palm shows a broadly triangular impression on
the inner surface just below the upper shoulder; the outer surface has a
similar depressed area, which however, is more quadrangular in outline. A
blunt tooth is present on the inner surface of the palm near the base of the
dactylus; this tooth is rather indistinct in the female, the carpus is short and
cup-shaped, the merus is about as long as the palm.
The smaller first leg in the male has the dactylus “balaenicep”
shaped, on both the inner and outer surface there is an oblique row of
closely set hairs, placed on a distinct ridge, which in its basal part
overhangs the cutting edges. The fingers are about as long as palm, their
edges are unarmed. The palm is more slender in the large leg. The dorsal
and ventral incisions are distinct. Here, too, is a triangular depressed area
on the inner surface of the palm below the dorsal incision. A distinct, rather
sharply pointed tooth is present on the inner surface near the base of the
dactylus. In the female the smaller leg differs conspicuously from that of the
male by the absence of the hairy ridges on the dactylus; the incisions on the
palm are less distinct here, but still noticeable. The merus is as long as the
palm.
261
Figure 120: Alpheus lobidens. A-C- Left first to third maxillipeds
The second pair of pereopods is symmetrical; the fingers are nearly
equal to the palm. The carpus is five segmented. Merus and ischium are
nearly of equal size. The third pereopod reaches with part of the propodus
beyond the antennular peduncle. The propodus is armed with spines on the
posterior margin and it is longer than the carpus. The fourth pereopod is
slender; the propodus is also armed with a row of spines on its posterior
margin. The fifth pereopod is more slender than the preceding two pairs, the
propodus bears 4-6 spines on the posterior margin, and oblique rows of
hairs are present on the distal two third portion of the segment.
262
Figure 121: Alpheus lobidens. A- Left cheliped; B-E- Left first to fourth
pereopods
In both the sexes, the endopod of the first pleopod is small,
whereas, the endopod of the second pleopod in female bears only an
appendix interna, in males both the appendix interna and appendix
masculina are present.
Remarks:
The Indian Ocean and western Pacific Ocean
specimens are greater in size than those of central Pacific are but no
dividing line was fixed (Banner & Banner, 1974). This species is perhaps the
most common alpheid found here and used as bait in fishing, used to be
collected at Karachi in the hundreds in one sample but now it is not so
abundant.
Anker (2001) indicated that his new collection may help to
determine the validity of A. inopinatus and A. audouini.
263
Colour: Greeny-brown, olive green, or smoky-grey, anterior
parts of abdomens often white, with, or without longitudinal stripes. Telson
and uropods apically blackish; chelae green- orange or green- brown, finger
and thumbs of large chelae orange, tips dull violet, palm with a more or less
brilliant cobalt – blue patch on inner surface; other legs dull pinkish (Kemp,
1915, Barnard, 1950) .In live specimens the color is variable, several
colour patterns have been observed of which one is photographed to be
included in this report.
Habitat:
At the low tide in large number from crevices, more
often from between the two layers of sandy stones.
Localities:
Manora Island, Hawks Bay, Somar Goth, Bulleji,
Astola Island, and Korangi Creek.
Distribution: Indo-Pacific – from Red Sea to Hawaii, as well as
eastern and central Mediterranean.
Development: (Fig.122).Only first larval stage has been
described (Tufail & Hashmi, 1965), later four larval stages were obtained
by Barkati (1980)but with no description was given.
First Larval Stage:
Length: 2.5mm.The general shape of the body is elongated with a
carapace, which lacks the grooves and details of the mature adult. Unlike
the ventral surface of the carapace, dorsal surface is smooth and curves
gradually down to the lateral edges. The ocular lobe does not project much
over the base of the eyestalks. The rostrum is very small, situated between
the bases of antennules.
264
H
Figure122: Alpheus lobidens. First larva. A- Entire latro-dorsal view;
B- Same, Ventral view; C- Antennule; D- Antenna; E- Maxilla; F-H-First
to third maxillipeds; I- Telson (Modified from Tufail & Hashmi, 1965 as
Alpheus crassimanus)
The body is clearly divided into two parts. The carapace covers the
anterior part, which is slightly less than one-third of the body length. The
posterior edge of the carapace is nearly straight and covers the basal
portion of the maxillipeds. The narrower posterior part is divided into
abdomen of six somites and unsegmented telson. The abdominal somites
are provided with lateral spines. Slight pointed abdominal spines mark the
junctions of the abdominal segments. The telson is much more elongated
but not forked, bears 16 spines, eight on either side of the midpoint. The
spines are barbed except their basal portion, which is smooth. The central
pair of spines is the smallest and is surrounded by a pair of slightly larger
spines. The latter in turn are enclosed by 4 larger spines on either side. The
spines on extreme ends of telson are small. Antennule is smaller than
antenna; divided at the distal end, one lobe bears three long plumose setae
while the other smaller one bears two plumose setae. Apical segment of
peduncle is produced into two flagella, outer and inner, the outer bears
three long plumose setae, and the inner flagellum two spines.
265
Antenna is biramous, the exopod, which is larger, bears eight
long, plumose setae while the endopod is smaller and possesses two
comparatively smaller plumose setae. Endopod is slender, and bears two
segments while the exopod is unsegmented. Scaphognathite bears 12
plumose small setae. First maxilliped has short endopod, which is
segmented and bears two small plumose setae at the last segment and one
at second segment; one at the first segment; three at the base. While the
exopod is long and unsegmented, and possesses four long apical plumose
setae at the terminal end. Second maxilliped endopod is segmented and is
ended in one long and two small spines, all are simple; exopod is also
segmented, bears a pair of long plumose setae at the base of last segment
and four long plumose setae at the tip of the last segment. Rudiments of
third maxilliped are seen as underdeveloped exopod and endopod is noted
to be without any setae.
Alpheus manorensis Afzal ,Javed&Barkati, 1986
(Fig.123)
Alpheus manorensis Afzal et al, 1986: 339
Description: The rostrum is short and narrow reaching to the
middle of the visible part of the basal segment of the antennular peduncle;
the orbital hoods are flattened dorsally. The rostral carina reaches to the
base of the orbital hoods. The orbitorostral grooves are broad and shallow.
The telson is 1.2 times as long as the posterior margin is broad.
266
Figure 123: Alpheus manorensis. A- Anterior region; B- Large chela,
male; C- Small chela, male; D- Second right leg; E- Third right leg.
The lateral spine of the stylocerite reaches to the end of the basal
segment of the antennular peduncle. The outer margin of the scaphocerite
is slightly concave, the lateral spine of the stylocerite reaches beyond the
antennular peduncle, and the squamose portion reaches to the middle of
the third segment of antennular peduncle. The carpocerite is definitely
longer than the antennular peduncle as well as the lateral spine of the
267
scaphocerite. The lateral spine of the basicerite is short and strong. The
second segment of the antennular peduncle is 1.4 times as long as the
visible part of the basal segment of the antennular peduncle. The third
segment is slightly shorter than the visible part of the basal segment.
The large chela is 2.3 times as long as broad with the fingers
occupying the distal 0.4 of the chela; the sculpturing on the palm is the
dactylus is pointed at the tip in both the sexes. In the male there is a
prominent ridge on the inner face of the dactylus, which bears a row of
sparse setae; the tooth on the dactylus is slanting posteriorly in male
whereas in the females it is a jutting out and knob like.
The small chela of the male and female are similar and not
balaeniceps in either sex; the chela is 3.9 times as long as broad with the
fingers occupying the distal half; the palm bears slight depressions on the
superior and the inferior margin but is smooth on the inner and outer faces;
the merus is 2 times as long as broad and unarmed. The dactylus is simple.
The ratio of the carpal joints of the second leg is 10:6:4:4:6; the
ischium of the third leg is armed with movable spine; the merus is 4 times as
long broad, unarmed. The carpus is half as long as the merus; the propodus
is 0.7 as long as the merus and bears eight movable spines on the inferior
margin; the dactylus is simple.
Size:
33mm in TL (male), 32- 37mmTL (female)
Locality:
Manora Island.
Distribution:
Not out side Pakistan
Alpheus pacificus Dana, 1852
(Fig. 124)
Alpheus pacificus Dana, 1852: 544; Coutiere, 1899:16, fig. 290; 1905:909;
Chace, 1962:610; 1988:45; Tiwari, 1963:315; Banner & Banner, 1966:143,
fig. 54; 1979:203; 1981:37; 1982a: 37; 1982b:217; Gillett & Yaldwyn,
1969:70, 110, fig. 41; Kazmi & Kazmi ,1979:154; Afzal et al, 1986:339;
Miya, 1984:96; Kim & Abele, 1988:100; Poupin, 1996:6; Yang & Ko,
2005:34
Alpheus gracilidigitus Miers, 1884:287
Crangon pacifica – Banner, 1953:138, fig. 50 [neotype established]
268
Crangon pacificus- Schmitt, 1939:12
Description: The rostrum is elongate, slightly overreaching middle
of visible part of the first antennular segment and bears few short setae on
lateral margins proximally. The rostral carina in rounded dorsally extending
backwards beyond posterior end of ocular hood; the ocular hood is slightly
inflated above the level of rostral carina and produced anteriorly to middle
of rostrum; the orbitorostral groove is moderately deep and reaching far
posterior to eye, anterior margin of ocular hood is rounded, very concave
near base of rostrum.
269
Figure 124: Alpheus pacificus. A- Anterior region; B- Telson and
uropods; C- Large chela, female; D- Small chela,male; E- Small chela,
female; F- Second left leg; G- Third left leg
The telson is about 1.6 times as long as broad at anterior end,
armed with two pairs of rather stout dorsal spines and with no longitudinal
median depression on dorsal surface. Posterior margin is rounded, bears
setae-like spines, and a pair of spines at each lateral end; the inner spine
is almost as long and wide as outer one.The pleurae of first four abdominal
somites are broadly rounded in both the sexes.
The first antennular segment bears very shallow, broadly V-shaped
carina which extends from ventral inner margin; the second segment is
about 2.3 times; as long as broad, 1.7 times as long as visible part of first
segment and 1.6 times as long as third segment; the stylocerite is
proximally broad, abruptly narrows to a sharp point, almost reaches to distal
margin of the first segment.
The scaphocerite is about 2.6 times as long as broad with lateral
margin slightly concave at the proximal 1/3, distal spine at least reaches to
distal end of antennular peduncle. Inner blade regularly becomes narrow
distally, and is far shorter than the adjacent distal spine, basicerite bears a
sharp triangular lateral spine; the spine is broad at base, almost reaches to
tip of the stylocerite.
The third maxilliped slightly overreaches the distal end of
carpocerite, ultimate segment is about 5.2 times as long as broad at
proximal end and 1.7 times as long as penultimate; the distal margin is
truncate, not much narrower than proximal end and bears long setae on its
superior and inferior margins as well as on distal margin; dense tufts of
setae are present on superior and inferior margins, especially distally; the
antepenultimate segment has sparse setae on inferior margin and few long
setae on superior distal margin; the exopod slightly overreaches distal end
of antepenultimate segment.
The major leg overreaches distal end of carpocerite by the length of
chela. It is about 2.3 times as long as broad; the fingers occupy distal 1/3.
Movable finger is regularly arched in profile and compressed laterally and is
acutely rounded; the immovable finger is directed upward, the superior
margin is very concave; the palm has a superior transverse and very deep
groove, the proximal shoulder overhangs the groove, the superior outer
270
palmer depression is well defined, quadrangular, extends to the oblique
suture, the superior inner palmer depression is elongated triangular in
shape and overreaches proximally, middle of groove is very deep slightly
slanting posteriorly; the proximal shoulder is heavy, rounded, slanted distally
and bears small granules; the inferior outer palmer depression is very
narrow, and obliquely, inverse V-shaped. The inner face of palm has a
shallow longitudinal depression near bottom of palm; the merus is about 1.8
times as long as broad, its inferior inner margin bears very short setae and
has no spine at distal end. The minor chela of first pereopods is about 3.5
times as long as broad, the fingers occupy slightly more than distal 0.6; the
inferior margin of palm has a broad shoulder below level of dactylar
articulation, the inner face of palm has blunt tooth flanking dactylar
articulation; both the fingers are slender and their tips are acute, movable
finger is not balaeniceps, series of dense setae are present on lateral
cutting edge that cross a similar series of setae on fixed fingers; tips of
fingers overlap but fingers gape between the tips and rest on movable finger
when closed. Minor chela of the female is smaller, four times as long as
broad with fingers only 2.5 times longer than the palm and bear only
scattered setae instead of rows of long hairs; the merus is about 2.5 times
as long as broad with very short setae on inferior inner margin and without
any spine at distal end.
The second pereopod overreaches distal end of carpocerite by ½ of
first carpal segment; the fingers of chela are almost as long as second
segment; the carpal articles of second leg bear ratio of 12:11:4:4:6.
The dactylus of third pereopod is simple, about 0.4 times as long as
propodus; the propodus is almost as long as carpus, with 7 movable spines
and a pair at distal end, the merus is about 4.5 times as long as broad, 1.5
times as long as carpus, bears few stiff setae on inferior margin, the ischium
has movable spines;
The appendix masculina is almost as long as appendix interna.
Colour: Tip of major chela brown, rest of fingers and palm banded
with irregular white, olive-green and blue-green. Fingers of small chela light
green, distal three-fourths of palm white and olive-green at base. Carpus
and dactylus blue; thoracic legs blue with white band at meral- carpal joint.
Antennae blue, antennules olive-green. Scaphocerite and carpocerite blue.
Carapace and abdomen reddish brown with faint brown line extending from
middle of carapace to sixth abdominal somite; on the lateral margins where
each abdominal somite meets the next is a diffuse white spot; telson of
271
same colour as abdomen, uropods light green. Colour pattern and
intensity varies from individual to individual and changes when kept in
aquarium (Banner & Banner, 1982:220).
Size: 34-35 mm in TL (male), 39-45mm in TL (females)
Habitat:
and rock spit.
Largely intertidal, living under rocks, rock shingle,
Localities: Manora Island,Nathia Gali,Bulleji,Light House and
Astola Island
Distribution: Australia; Red Sea and Madagascar; Mombasa
Kenya; Korea, eastern Pacific from Gulf of California, Costa Rica,
Clipperton Island and Galapagos Islands.
Alpheus pseudoedwardsii Afzal,Javed&Barkati,1986
(Fig.125)
Alpheus pseudoedwardsii Afzal et al, 1986: 330
Description: The rostrum is acute and triangular, twice as long as
broad at the base and reaches beyond the middle point of the basal
segment of the antennular peduncle; the lateral margin bears sparse and
short setae; the rostral carina extends as far as the base of the orbital hoods
from which it is separated by shallow and broad grooves.
The telson is 3.0 times as long as broad; its posterior margin is
broad. The anterior margin is 1.7 as long as the posterior margin. The
lateral margins are convex in the anterior third and narrowing slightly in the
posterior two third.
272
Figure 125: Alpheus pseudoedwardsii. A- Anterior region; B- Telson
and uropods; C- Large chela, female; D- Small chela, female; ESecond left leg; F- Third left leg
The second segment of the antennular peduncle is 3 times as long
as broad and 1.7 times as long as the visible part of the basal segment,
while the third segment is nearly as long as the basal segment; the acute
273
spine of the stylocerite reaches near the end of the basal segment. The
outer margin of the scaphocerite is slightly concave; the lateral spine
reaches beyond the antennular peduncle, the squamose portion reaches
the end of the antennular peduncle; the carpocerite is as long as the lateral
spine of scaphocerite; the lateral spine of the basicerite reaches almost to
the level of the tip of the stylocerite.
The large chela is 2.5 times as long as broad, slightly compressed;
the fingers are slender longer than the palm and occupy 0.6 of the total
length of the chela; the fingers are pointed at the tips; the dactylus is similar
to the fixed finger. It is sculptured with transverse grooves, which run
throughout the length of the superior margin on inner and outer face; the
grooves on the inferior margin run from the tip to the base of the tooth of the
dactylus. The tooth is an elongated inferior elevation of the dactylus, which
occupies most of the proximal half of the dactylus.The distal, and proximal
margins of the superior groove of the palm are broadly rounded. The groove
is continued on the outer face as roughly triangular areas, its tip reaches to
the middle of the palm. The notch on the inferior margin is much deeper
than the one on the upper, which has proximal shoulder projecting. The
merus is 2.5 times long as long broad, lacking a tooth on the inferior internal
margin but with a shallow notch in the proximal half.
The small chela is 4 times as long as broad with fingers longer than
the palm, occupying distal more than half of the chela; there is a tooth like
swelling on the inferior margin of the finger. No corresponding cavity is
present on the opposing face of the fixed finger. The merus is above 3 times
as long as broad and is unarmed. The ratio of the carpal joints of the
second leg is 10:7:2:3:5. The ischium of the third leg bears a strong
movable spine. The merus is 3.5 times as long as broad; the carpus is half
as long as the merus with both the distal end of the superior and inferior
margin projecting and acute. The propodus is 0.8 as long as the merus and
bears ten movable spines on its inferior margin. The dactylus is simple.
Size: 3mm in CL, 39mm in TL.
Habitat: Tide pools
Locality: Manora Island
Distribution: Not reported outside Pakistan
274
Alpheus cf. rapax Fabricius, 1798
(Pl. 5B)
Alpheus rapax Fabricius, 1798:405; de Man, 1909:147; Barnard,
1950: 752; Banner & Banner, 1966:121, 1982: 174; Thomas,
1970:1; Miya, 1984:97
Alpheus malabaricus Hilgendorf, 1878:832
Alpheus brevirostris de Man, 1888a:22:261
Description: The rostrum is acute, reaching variously from first
quarter of visible part of first antennular article to near the end of that article.
The rostral carina is sharp, reaching posteriorly to the base of orbital hoods.
The orbital hoods are inflated, forming deep orbitorostral grooves.
The telson is more than 2.5 times as long as its posterior margin;
the posterior margin is strongly arcuate.
The antennular peduncles are slender; the second article is longer
than the visible part of the first, and more than two times longer than the
third article. The stylocerite is flattened and leaf-like; the lateral spine is
small, reaching near end of first antennular article, the lateral margin of
scaphocerite is slightly curved, the lateral spine is a little longer than the
squamous portion; the carpocerite reaches to the end of antennular
peduncle.
The large chela is compressed almost 3 times as long as broad,
the fingers occupy distal one fourth, the superior margin of palm bears a
transverse groove proximal to dactylus, upper and lower inner margin of
palm bears rows of setae. The dactylus is heavy, its distal end is broadly
rounded, and the merus is nearly three times as long as broad, inferior
internal margin is armed with 4-7 movable spines and terminates in a strong
tooth.
The small chela is highly compressed, nearly five times as long as
wide, the fingers are 1.3 times as long as palm; their opposing faces are
flattened and fringed with a dense row of bristles, both fingers are strongly
hooked at tip. The merus is three times as long as wide, its armature similar
275
to that of the large chela. The large chela in males bears a short dense
row of bristles on inferior margin of dactylus.
The carpal articles of second leg bear a ratio of 10 :( 10-7):3:3:5.
The ischium of third leg bears a small spine; the carpus is almost half as
long as merus than the merus; the propodus is shorter than the merus and
bears on inferior margin nine movable spinules. The dactylus is 0.4 lengths
of propodus, subspatulate, laterally expanded, with inferior surface slightly
excavate.
Remarks: The unidetifiable specimens are only tentatively given
the name of A. rapax.
Colour: Black and white transverse bands across cephalothorax
and abdomen. Large chela with similar transverse markings clearly visible
on the inner side also. Tips of fingers white.
Habitat: Sandy muddy intertidal areas, burrowing in association
with gobies(Karplus,1987).
Distribution: Red Sea, S. Africa, Hawaii, Singapore, Japan,
Marshal and Caroline Islands, Indonesia, Australia, Mergui, India, Sri- Lanka
and now for the first times from Karachi.
Alpheus splendidus Coutiere, 1897
(Figs. 126-129,Pl.4A )
Alpheus splendidus
Coutiere, 1897:236; 1898: 197;1921: 426;
Johnson, 1962: 52; Crosnier & Forest, 1965:361, fig.4; Kazmi & Kazmi,
1979:153; Banner & Banner, 1982:56; Wicksten & Hendricks, 1985:571;
Afzal et al, 1986:338; Chace,1988:54
Alpheus pomatoceros Banner & Banner, 1966: 93, fig. 32.
Description: The body is a smooth and polished. The rostrum is
triangular and acute, in the males it extends beyond the basal segment of
antennular peduncle whereas in female it reaches as far as the distal
margin of the basal segment of antennular peduncle. A rounded carina is
276
present on the rostrum, which is continued backwards, extending over
the anterior one third of the carapace; short but deep orbitorostral grooves
separate the rostrum from the orbital hoods; the latter are anteriorly rounded
and from their upper surfaces, sharply pointed spines arise, directly above
the eyes. Each orbital spine has a short carina, which extends backwards
for a short distance; a shallow groove is present on the lateral side of each
orbital hood.
Figure 126: Alpheus splendidus. A- Carapace, lateral view; B- Telson
and uropods; C- Right antennular peduncle.
The telson is nearly as long as the last abdominal somite. It tapers
slightly toward the tip; the posterior margin is somewhat convex and armed
with two spinules on each postero-lateral angle, of which the inner one is
longer, besides these, two pairs of spines are present on the dorsal surface.
The antennular peduncle consists of three sub-equal segments; the
basal segment is the largest. It also bears an acute process near the distomedian angle and another, laterally at about the middle of the segment; the
stylocerite reaches up to the proximal half of the second segment of the
antennular peduncle. The second segment is longer than the third segment.
277
The spine of the scaphocerite is strong reaches beyond distal margin of
the blade.
Figure 127: Alpheus splendidus. A- Right antennal scale; B- Left
maxillula; C- Right maxilla; D-F- Left first to third maxillipeds.
278
Figure 128: Alpheus splendidus. A- First large cheliped; B- First
small cheliped; C- Second right leg; D- Third right leg
Seven incisor teeth are present in the mandible. The first maxilliped
has a slender palp.
The first pair of pereopods is asymmetrical; the large chela is
slender being three times as long as broad. The immovable finger is onethird the length of the palm; it is more or less club shaped, the apex being
broadly rounded; a spine is present on either side of the dactylus
articulation; the small chela is 4.3 times as long as broad; few spinules are
present as illustrated on the inferior margin of the merus.
Figure 129: Alpheus splendidus. A- Second pleopod, female; B- Same,
male
279
The endopod of the first pleopod in female is small, whereas,
the endopod of the second pleopod bears an appendix interna. In male both
the appendix interna and appendix masculina are present.
Colour: Body pale pink with two sub median longitudinal bands of
orange colour extending over the entire body length.
Size: 40-47 mm in TL (male), 29 mm in TL (female)
Locality: Manora Island
Distribution: Red Sea, eastern Africa, Seychelles, Malaysia,
Thailand, Hong Kong, Philippines, Indonesia, Australia, and eastern Pacific.
Alpheus strenuus strenuus Dana, 1852
(Figs. 130-131)
Alpheus strenuus Dana, 1852 a : 21: 1852b:543, pl. 3, fig. 4;
Barnard, 1950: 760; Kazmi & Kazmi, 1979:154
Alpheus strenuus var. angulatus Coutiere, 1905: 913, pl. 87 fig. 53.
Alpheus strenuus strenuus Banner & Banner, 1966: 140, fig. 53.
1982: 225; 1983:83; 1985:32; Chace, 1988:56
Description: The rostrum is slender reaches just beyond the end of
the first antennular segment; the rounded rostral carina extends up to the
base of the orbital hoods. The orbital hoods are only slightly inflated forming
moderate groove between orbital hoods and carina. The carapace is
smooth without any carina or groove.
280
Figure130: Alpheus strenuus strenuus. A- Left antennular peduncle; BLeft antennal scale; C- Right maxillula; D- Right maxilla; E- Telson and
left uropod
The telson is longer than the sixth abdominal somite, the dorsal
surface is armed with two pairs of posteriorly directed spines, and each
lateral side bears a pair of spines of which the outer one is smaller.
The antennular peduncle consists of three sub equal segments, the
second antennular segment is 2.3 times as long as broad in the middle and
it is 1.5 times as long as visible portion of the first; the stylocerite reaches
nearly up to or beyond the end of the first antennular segment. The lateral
side of the scaphocerite is concave and ends into a spine, which is longer
than the squamose portion; the carpocerite is nearly as long as the spine of
the scaphocerite.
281
B
F
A
D
E
C
Figure 131: Alpheus strenuus strenuus. A-C Left first to third
maxillipeds; D- Large chela; E- Small chela; F- Left second leg.
The oral appendages are typical.
The large chela is somewhat compressed, two and a half times as
long as broad; the dactylus is not noticeably curved, it is not double ended,
having a well developed plunger, the palm has a longitudinal groove on
inner surface near margin, at the base of fixed finger the saddle is situated
proximal to the adhesive plaque, the proximal shoulder is blunt, slightly
overhangs the saddle; the shoulder is strong behind the fixed finger, it is not
acute; the merus is 2.6 times as long as broad, the superior margin is
282
without any tooth whereas the inferior internal margin is armed with
several small spines and bears a strong acute tooth distally.
The small chela is 4 times as long as broad; the dactylus is
“balaeniceps” in both the sexes, shorter than palm. The merus is 2.3 times
as long as broad in the middle. The distal end of the superior margin is
rounded whereas the inferior internal margin is armed with several small
spines and bears a small acute tooth distally. The segments of the carpus of
the second pereopod have the following ratio-10:10:3:3:4. The third
pereopod has a simple pointed dactylus, the propodus has 10 spines on its
flexor margin, and the merus is unarmed and the ischium is armed with
movable spines.
Remarks: It appear to well known to importers as snowflake
shrimp of tropical animals for the marine aquarium in Europe under several
Latin names in the aquarium – oriented literature.
Size:
32-68 mm in TL.
Colour: Either greenish brown, often mottled or
striped more or less distinctly with white, when stripes are present they are
usually longitudinal.
Habitat:May share its burrow/tube
fireworm Eurythoe.
Distribution:
between large granite boulders with
Throughout Indo- Pacific and Red Sea.
Alpheus zulfaquiri Kazmi, 1982
(Figs. 132-133, Pl.6B)
Alpheus zulfaquiri Kazmi, 1982: 137
Description: The rostrum is short, triangular, reaching slightly
beyond middle of visible part of basal segment of antennular peduncle; the
rostral carina is somewhat angular between orbital hoods, but rounded
posteriorly; the orbital hoods are inflated, the orbito-rostral grooves are
moderately deep.
283
Figure 132: Alpheus zulfaquiri. A- Anterior portion in dorsal view; BTelson in dorsal view; C- Large cheliped; C’- Fingers of large chela;
C”- Merus and carpus of large chela; D- First small chela; E- Second
pereopod
The telson is nearly one and half times longer than sixth abdominal
somite, its posterior margin is convex, the postero-lateral angles bear two
pairs of spines, the outer spine is about half as long as the inner; the dorsal
surface of telson is armed with two pairs of spines; a shallow longitudinal
groove is present in posterior half of the telson.
The antennular peduncle reaches slightly beyond the scaphocerite;
the second segments is 2.4 times as long as the visible portion of first and
four times as long as broad, The tip of stylocerite is acute reaches almost to
the end of first antennular segment. Lateral margin of scaphocerite is
slightly concave, terminating in a tooth, reaching beyond squamose portion;
the carpocerite is as long as the antennular peduncle, spine of basicertie is
small and acute.
284
D
Figure 133: Alpheus zulfaquiri. A- Large chela in lateral view; B-DThird to fifth pereopods.
The large chela is three times as long as broad in the middle, the
fingers are slightly shorter than the palm, has low rounded ridges which start
from proximal region and end at distal region of palm; the superior margin of
palm has shallow rounded transverse groove which terminates abruptly on
either face, inferior shoulder is lacking, outer face of palm has low rounded
ridges which start from proximal region and end at distal region of palm, the
merus is twice as long as broad distally; the inferiointernal margin is armed
with six movable spines, terminating in a strong tooth. The small chela is 4.5
times as long as broad in the middle, the movable finger is nearly twice the
length of palm, the immovable finger is markedly shorter than the dactylus;
the inner-inferior internal margin of merus bears few movable spines, and
terminates in a strong tooth. The carpal articles of second leg have a ratio
of 3.1:3.1:1:1:1.3; the ischium of third leg is armed with a movable spine; the
merus is unarmed, it is five times as long as broad; carpus is 0.6 the lengths
of the merus with 5-6 spines; the dactylus is 0.3 as long as the merus, it is
spatulate in shape. The fourth and fifth pereopods are shorter than the third;
the carpus of fourth pereopod is slightly shorter than the propodus; the
propodus bears seven spines; dactylii of fourth and fifth pereopods are
spatulate.
Size: 45 mm in TL
285
Locality: Manora Island
Distribution: Not out side Pakistan.
Genus Athanas Leach, 1814 sensu stricto
Diagnosis: Body usually slender. Carapace usually smooth, sometimes
setose. Frontal margin with long, straight, slightly ascendant, or
descendant rostrum, with acute tip, without subdistal tooth on ventral
margin, rarely with teeth on dorsal margin. Extra-corneal teeth present,
acute, sometimes protruding beyond anterior margin of eye; ) supra- and
infra-corneal teeth absent or present. Orbital hoods absent; eyes exposed
in dorsal and lateral views. Pterygostomial margin usually rounded, rarely
sharply protruding or with small acute tooth. Cardiac notch well
developed. Sixth abdominal segment with articulated plate at
posteroventral angle; preanal plate rounded. Antennules with straight, not
serrated distal margin of 1st segment; ventromesial carina of 1st segment
with strong tooth; lateral flagellum with welldeveloped secondary ramus.
Antenna with normal, not particularly stout basicerite, distolateral tooth
present; scaphocerite variably oval to subrectangular, usually more or less
elongate. Mandible with palp, incisor process usually not expanded,
distally with 5-8 medium-sized teeth, rarely expande, with 12-15 min teeth.
Maxillule with bilobed palp, both ventral and dorsal lobes with setae.
Labrum not swollen or protruding. Third maxilliped without distal tooth on
dorsal margin of antepenultimate segment; penultimate segment
elongate, distinctly longer than wide; ultimate segment often with spines
on tip. First pereopods polymorphic, carried folded or extended with
dactylus in lateral position; coxa usually with(chelipeds) highly variable in
shape, size, and degree of asymmetry, often sexually dimorphic or
apressed subtriangular tooth mesially; basis with rudimentary exopod;
ischium unarmed mesially, often with spines on dorsal and ventral
margins;chelae without linea impressa on palm and without fossa-plunger
on fingers. Male chelipeds usually enlarged and/or elongate, equal,
subequal, or unequal, symmetrical, subsymmetrical, or asymmetrical
(with major cheliped situated either on left or right side); ischium
sometimes with lobes furnishedwith spines on dorsal margin; merus
usually stout, depressed or excavated ventrally; carpuselongate or cupshaped, sometimes flattened ventrally;chela highly variable, usually
conspicuously swollen or elongate, sometimes with tubercles
286
along ventral margin; fingers usually armed with teeth. Female
chelipeds variable from slender,slightly elongate, to greatly enlarged,
equal,
subequal,
or
unequal,
symmetrical,
subsymmetrical,
orasymmetrical (with major cheliped situated on left or right side); merus
variable, slender to stout, flattened or excavated ventrally; carpus
elongate or cup-shaped, sometimes flattened ventrally; chelaevariable
from slender to oval or elongate, sometimes with tubercles along ventral
nd
margin; fingers unarmed or armed with teeth. Carpus of 2 pereopod with
5 segments, exceptionally 4 or 6.
Third pereopod with ischium unarmed or bearing 1 spine on ventrolateral
margin; merus unarmed,
distoventral margin never acute or angular;
carpus unarmed; propodus usually with ventral spines,rarely with spinelike setae; dactylus more or less slender, simple or biunguiculate. Fifth
pereopod with well-developed propodal brush. Second pleopod with
appendix interna and appendix masculine in males, appendix interna in
females, exceptionally with appendix masculina in ovigerous
specimens.Telson with 2 pairs of dorsal spines and 2 pairs of
posterolateral spines; anal tubercles absent. Gill formula with somewhat
variable numberof epipods, as summarized in following table (untypical
condition indicated in parentheses).(from Anker&Jeng,2007)
Type species: Palaemon nitescens Leach, 1814 :401
Gender:
Masculine.
Remarks: The gulf and Indo- West-Pacific members of the genus
are 34,all small and inconspicuous .Some species are symbionts with other
animals. (Chace, 1988,Anker et al.,2010) of which only two A. dimorphus
Ortmann, 1894,belonging to a heterogenous species group and A. arabicus
Afzal, Javed&Barkati , 1986, have been recorded from Pakistan, they can
be separated with the help of following key.
KEY TO THE PAKISTANI SPECIES OF ATHANAS
1.
Infracorneal spine rounded. One movable spine on
propodus of third leg on the distal end. Telson armed with one spine on
each posterolateral angle ------------------------- A.dimorphus.
Infracorneal spine acute. A row of seven to eight spines on
propodus of third leg. Two spines on each posterolateral angle of telson------------------------------------------------------A. arabicus.
287
Athanas arabicus Afzal,Jaed&Barkati, 1986
(Fig. 134A)
Athanas arabicus Afzal et al ,1986:339
Description: The rostrum is long and triangular, its sides gradually
taper to pointed tip which reaches to the end of second segment of the
antennular peduncle; it bears a sharp carina running backwards up to the
level of the orbital margin. The supracorneal spine is absent. The extra
corneal spine is acute, reaching to the middle of the cornea, the infracorneal
spine is small, and acute .The cornea of the eye is partially concealed at the
base by the extra and infracorneal projections. The pterygostomian angle is
rounded.
G
A
E
B
D
F
C
Figure 134A: Athanas arabicus. A- Anterior part lateral view; B- Same
dorsal view; C- Telson and uropods; D- Third maxiliped; E- F- First
legs; G- Second leg
288
The telson is three times as long as broad at the posterior margin,
the anterior margin is 1.5 times as long as the posterior margin;it bears a
pair of spines on each posterolateral angle of which the inner is twice as
long as the outer one besides two pairs of spines are present on the dorsal
surface.
The second segment of the antennular peduncle is equal in length
to the visible part of the basal segment; the third segment is slightly longer
than the second; the stylocerite reaches to the middle of the third
segment of the antennular peduncle; the lateral spine of the scaphocerite
reaches as far as the distal margin of squamose portion; the carpocerite
reaches to the middle of the third segment of the antennular peduncle.
The first pair of pereopods exhibits sexual dimorphism, but in both
sexes, they are of symmetrical development; in the female the ischium of
the cheliped bears one or two spines on the superior margin; the merus is
flattened, unarmed, unexcavated on its inferior surface. In one female a
small exopod is present on the first pereopod; the carpus is as long as the
merus; the chela is equal to the merus, the finger is half as long as the
palm, the inner margins of the fingers do not gape when closed; in males
the first pair of pereopods is cylindrical and folded underneath the body; the
ischium has two to three spines on the superior margin; the merus is
spindle-shaped, unarmed and excavate on its inferior surface to
accommodate the propodus; the palm is sub cylindrical, broader at midsection, longer than merus; proximally a slight tooth-like projection is
present on the inferior surface; the movable finger is slightly longer than the
fixed finger; both the fingers have a small tooth on their inner margins, that
of the movable finger is small and insignificant;
The carpus of the second pereopod is five-segmented; the ratio of
carpal joints is 6:1:1:1:2; the chela is small but distinct; the fingers are
slightly shorter than the arm.
The ischium of the third pereopod is less than half the length of the
merus, it bears a fine spine on the distal superior margin; the merus is 4.5
times as long as broad in the middle; the carpus is nearly 0.5 as long as the
merus; the propodus is nearly as long as the merus, it bears a row of 7 to 8
spines on its inferior margin of which the distal most is longer and stronger.
The uropods are a little longer than the telson, and they bear a
strong spine on each posterolateral angle.,
289
Size: 6-17mm in TL (male), 6-12mm in TL (female)
Habitat: In small tide pools with dense vegetation
Locality: Manora Island
Distribution: Not reported outside of Pakistan.
Athanas dimorphus Ortmann, 1894
(Figs.134B-137)
Athanas dimorphus Ortmann, 1894: 12, pl. 1, fig. 1; Tattersall, 1921: 371,pl.
28,figs.23-24; Kazmi & Kazmi,1979:153; Bhuti et al, 1977: 591 (larvae) fig 24;
Banner
&
Banner,
1973:313;
1978:234;1981:41;1982:308;1983:76;Chace,1988:61;Kazmi,2003:240
Athanas leptocheles Coutiere, 1897a:381
Athanas dispar Coutiere, 1897b:233.
Athanas solenomerus Coutiere, 1897a: 381.
Athanas dimorphus seedaing Banner & Banner, 1966: 28
Description: The body is small and sparsely pubescent. The
rostrum is styliform. It reaches to the proximal portion of the second
segment to third segment of the antennular peduncle; a middorsal carina is
present which extends posteriorly to the base of the rostrum. The
supracorneal spine is wanting. The extracorneal spine is acute, reaching to
the middle of the cornea, infracorneal spine is small and rounded. Cornea of
the eye is concealed at the base of the extra and infracorneal projections.
The pterygostomian angle is somewhat rounded. The posterior margin of
the carapace has a well-developed cardiac groove and notch.
290
Figure 134B: Athanas dimorphus. A- Animal in lateral view; A’ -A’’’
variations in chela
The posterolateral margin of each of the first four abdominal
somites is generally rounded whereas, that of the fifth is pointed. Sixth
abdominal somite is longer than the fifth. The telson is nearly three times as
long as the posterior margin. It tapers slightly towards the posterior end.
Each of the postero-lateral angles of the telson is produced into a small
spine. The dorsal surface of the telson is armed with two pairs of spines.
The antennular peduncle consists of three sub-equal segments; the
basal segments being the largest, small spine is present on the median
side; the sylocerite is sharply pointed and extending almost up to the distal
margin of the second segment; the second and third segments are subequal; the outer flagellum is divided after the third joint, each of which
appears to be sub-divided, so that this part has the appearance of being six
jointed. The scaphocerite reaches slightly beyond the end of the antennular
291
peduncle. The lateral margin is more or less straight and ends in a
spine, reaching barely as far as the distal margin of the squamose portion;
the antennal peduncle reaches to about the middle of the distal antennular
segment.
The mandibular palp is two segmented; the basal segment is
slightly longer than the second segment. The incisor and molar processes
are well developed. The maxillulary palp is bilobed, the upper lacinia is
broad and truncated; the inner one is narrow and curved. The palp of the
maxilla is short and finger-like.
The palp of the first maxilliped is long, extending slightly beyond the
caridean lobe. The second maxilliped is as illustrated; the third maxilliped
barely overreaches the scaphocerite. The ultimate segment is nearly two
and half times longer than the penultimate segment. The penultimate
segment is small; the parapenultimate segment is five times as long as
broad in the middle; the exopod reaches to the end of the parapenultimate
segment.
Figure 135: Athanas dimorphus. A-Lateral view of carapace; B- Dorsal
view of telson and uropods; C-Left, antennular peduncle in dorsal
view; D- Left scaphocerite in dorsal view.
292
The chelipeds are sexually dimorphic, but of nearly symmetrical
development in both sexes. In female the ischium bears one or two spines
on the superior margin and is equal to the chela; the merus and carpus are
nearly of equal size; the chela is about more than half of the carpus. The
inner margins of the fingers approximate in such a way that they do not
gape. In males, the first pair of pereopods is folded beneath the body. In
some specimens, a small exopod is present on the first pair of pereopods
(not shown in the figure). The ischium has two to four spines on its lateral
side; the merus is flattened; the inferior surface is deeply excavated to
accommodate the propodus; the carpus is small and nearly half of the
ischium; the palm is sub-cylindrical, three times as long as broad at the
middle and it is slightly longer than the immovable finger; the movable finger
of the right pereopods bears a projection which is only slightly developed or
it is produced into a well developed tooth, the immovable finger bears two
teeth.
The immovable finger of left pereopods has a small tooth which fits
in a notch on the immovable finger.
The carpus of the second pereopod is five segmented; the first
segment is slightly longer than the other four segments together, second,
third and fourth are sub-equal and each of them is less than half as long as
the fifth. The chela is small but distinct and it is nearly equal to the last four
segments of the carpus; the fingers occupy more than one third portion of
the chela; the ischium of the third pereopod is slightly shorter than the
carpus; the merus is nearly five times as long as broad in the middle; the
propodus is nearly equal to the merus, it bears a row of spines on its inferior
margin; the distal spine is strongly developed; the ischium of the fourth
pereopod is longer than the carpus and bears a single spine distally. The
fifth pereopod of male is as illustrated.
293
.
Figure 136: Athanas dimorphus. A-Right maxillula in dorsal view; BRight maxilla in dorsal view; C-Left first maxilliped in dorsal view; DRight second maxilliped in dorsal view; E-Left third maxilliped in
dorsal view
The pleopods are rather large and well developed. In male the
appendix interna and appendix masculina are situated nearly at the middle
of the endopod. The appendix interna fails to reach the tip of the endopod,
both in male as well as in the female.
294
Figure 137: Athanas dimorphus. A-First pereopod of right side; B- First
pereopod of left side; C- Second pereopod of left side; D-Third
pereopod of left side; E-Fourth pereopod of left side; F- Fifth pereopod
of left side.
Size:
12-17mm in TL (males), 13-16mm in TL (females)
Colour:
Freshly caught specimens show two colour
patterns particularly on their abdomen, where the transverse bands are blue
in some and red and blue in others.
Habitat:
Collected at low tide from the rocky pools
Localities: Manora Island, Sonari,Pacha and Bulleji.
Distribution: Indo-Pacific: Red Sea at Suez, Perim, Djibouti,
Darussalam, Thailand, Philippines, Hong Kong, Japan, Australia, East
Africa and Northern Arabian Sea.
295
Genus Automate de Man, 1888
Diagnosis: Body not unusually compressed; rostrum, if present,
inconspicuous, sub triangular or lobate, unarmed extension of frontal margin
of carapace; carapace without high carina throughout length of dorsal
midline; abdomen without articulated triangular flap at posterolateral angle
of sixth somite; telson not terminating posteriorly in triangular tooth; both
eyes and eyestalks visible in dorsal aspect; mandible with palp and molar
process; third maxilliped not unusually broadened to form partial operculum
over other mouthparts; first pereopods dissimilar, carried extended with
movable finger dorsal or lateral, not ventral; major chela without molar-like
tooth on movable finger; second pereopod with fingers about as long as
palm, carpus with 5 articles; pereopods with strap-like epipods on 4 anterior
pairs; appendix masculina absent. ( from Chace, 1988).
Type species: by monotypy: Automate dolichognatha de Man,
1888:529.
Gender:
Feminine
Remarks:Eight species seem to have been valid in this genus
(Chace, 1988), only one member thus far known from Pakistani waters is A.
dolichognatha.
Automate dolichognatha de Man, 1888
(Fig. 138)
Automate dolichognatha de Man 1888: 529; Coutiere, 1897:234, Banner &
Banner, 1973:299, 1981:48; 1982:308;Bhutti et al,1977(larvae); Rios &
Carvacho,1982: 462; Chace, 1988:64
Automate Gardiner
Coutiere, 1902:337; Suvatti, 1937:47; Holthuis,
1958:17; Banner & Banner 1966: 37; Miyake&Miya,1966:137; Chace, 1972:
74; Kazmi et al, 1973: 287; Afzal et al, 1986:339; Kazmi, 2003:241; Kazmi &
Kazmi, 2004:97
Automate johnsoni Chace, 1955:13
Automate haightae Boone, 1931:184
296
Description: The rostrum in dorsal view is rounded or
triangular; its tip curves downward and fails to reach as far forward as the
anterior margin of the carapace; the pterygostomial margin is rounded.
The telson is 1.3 times as long as broad at the base; it bears two
pairs of dorsal spines, i.e. the anterior pair is placed on the middle of the
telson and the posterior lies on the distal fourth; the posterior margin is
broadly convex, bears two pairs of distal spines; the outer one is very small,
the inners are as long as the breadth of the posterior margin.
Figure 138: Automate dolichognatha. A- Anterior region, dorsal view;
B- Telson and uropods;C- Large cheliped; D- Small cheliped; ESecond leg
The visible portion of the first antennular segment is 1.5 times as
long as the eye peduncle, and is half to three-fourths of the second; the
297
stylocerite reaches the end of the first segment; the second segment is
twice to three times as long as the third, and varies from 2.4 to 4.3 times as
long as broad at the maximum portion; the thickened part of the outer
antennular flagellum consists of eight to fourteen joints. The carpocerite
slightly reaches two-third of the second antennular segment; in the smaller
specimens, it reaches the end of the second segment.
The third maxilliped exceeds the antennular peduncle by about the
length of the ultimate segment which is eight times as long as broad and
armed with numerous paired short spines on the superior margin; the
penultimate segment is 2.4 times as long as broad; the antepenultimate
segment is swollen, being 4.2 times as long as broad.
In the large cheliped the cutting edge of the immovable finger bears
a strong tooth near the apex; the tooth is separated by a distinct interval
from three small teeth in the medial portion, and two larger teeth on the
proximal third of the cutting edge; the movable finger has a single distal and
two similar proximal teeth on the cutting edge, being as long as the palm.
The ischium bears a small spine or a seta on the dorsal distal corner.
In the second leg, the first two joints of the carpus show a ratio of
1:1.25-2.0; the last three pereopods have several spines on the ventral
margin of the propodus and a spine on the ventral margin of the ischium.
Colour: Colourless,
chromatophores dorsally.
Size:
transparent
or
pale
orange,
with
red
14.6mm in TL.
Habitat:
Intertidal or shallow sub tidal
Locality:
Bulleji ,Manora
Distribution: Pan Tropical (except for eastern Atlantic): IndoPacific and eastern Pacific
Genus Salmoneus Holthuis, 1955
Diagnosis: Body not unusually compressed from side to side;
rostrum represented by triangular extension of carapace; abdome without
flap articulated at posterolateral angle of sixth somite. Telson not
298
terminating posteriorly in triangular tooth. Eyes at least partially
concealed from dorsal view, visible in anterior aspect; mandible with palp
and molar process; third maxilliped not unusually broadened to form partial
operculum over other mouthparts; first pereopods dissimilar and unequal,
major cheliped carried in flexed position, without molar-like tooth on
movable finger; second pereopod with fingers about as long as palm,
carpus with 5 articles; pereopods with strap-like epipod on 4 anterior pairs;
appendix masculina not overreaching exopod of second pleopod of male
(After Chace, 1988).
Type Species: By subsequent designation Jousseaumea
serratidigitus Coutiere, 1896:385
Gender: Feminine
Remarks: It is worth mentioning that the specimens of Salmoneus
cannot be sexed if they are not ovigerous as both sexes have a welldeveloped appendix masculina, often the specimens are collected in pairs
(Carvacho, 1989). Dworschak et al(2000) recognized two groups by the
development of first legs while Anker & Marin (2006) proposed to subdivide
Salmoneus into seven species groups. Out of 29 known species, two
species S. brevirostris and S. cristatus in S. cristatus group are found here.
They can be separated as follows.
KEY TO THE PAKISTANI SPECIES OF SALMONEUS
1.
Rostrum barely reaching to end of first antennular article;
orbital area, with slight crest running diagonally towards midline---------------------------------------------------------------------------S.brevirostris.
Rostrum reaching to or beyond end of second antennular
article; ridges or crest arising in orbital area and running parallel to median
crest----------------------------------------------------------S.cristatus.
Salmoneus brevirostris (Edmondson, 1930)
(Fig. 139)
299
Jousseaumea brevirostris Edmondson, 1930: 7
Salmoneus brevirostris Banner, 1953:12; Banner & Banner,1966:
39; 1981:54; Kazmi, 1974: 310; 2003:241; Kazmi & Kazmi, 1979:
153; 2002:102
Description: The rostrum is broad at its base, acute, reaches to
end of the second antennular segment; a slight rostral carina is present
reaching almost to middle of carapace. The orbital hoods continue as small
acute orbital teeth, smaller than the rostrum. The area between the base of
orbital teeth and dorsal carina is slightly concave.
Figure 139: Salmoneus brevirostris A- Anterior region, dorsal view ;
B- Telson with uropods; C- Large first leg; D- Small first leg; E- Second
leg (Modified from Kazmi, 1974)
The telson is 4 times as long as broad at posterior margin; the
anterior margin is 2 times as wide as the posterior margin. The terminal cleft
of telson is triangular, small; the distal end occupies about one third of
telson tip. Two strong spines are present on either side of the terminal cleft.
The antennular peduncle is short and heavy, the visible portion of
first and second segments is sub equal in length, almost as long as broad.
The stylocerite is strong, the tip reaches almost to end of second antennular
300
segment; the lateral spine of scaphocerite is equal to its squamose
portion and reaches to the end of antennular peduncle. The carpocerite
reaches to middle of third antennular segment; the basicerite has an acute
lateral tooth.
The first pair of pereopods is asymmetrical; the large chela is 3.3
times as long as broad, with fingers a little shorter than the palm. The
dactylus and fixed finger have eight obtuse teeth; the carpus is cup-shaped;
the merus is slightly longer than the palm, excavated on its inner margin to
accommodate palm when appendage is flexed. The small cheliped is 0.6 as
long as the palm of large chela; the ischium and merus are sub equal; the
carpus is broadened at its distal end and is equal in length to the merus.
The chela is 4.5 times as long as broad, palm and fingers are equal in
length.
The carpal articles of second leg have a ratio of 10:1:1:1:2. The
merus of third leg is 4.5 times as long as broad, inermous. The carpus is
almost as long as merus, armed distally with a blunt tooth on superior
margin and movable spinule on inferior margin and a pair of spinules
distally; the dactylus is 0.4 as long as carpus. The fourth and fifth pereopods
are more or less equal in size.
Size:
16-18mm in TL.
Colour: Bright orange yellow
Bulleji,Manora
Locality:
Habitat: intertidal zone under loose stones
Distribution:
Hawaiian Is., Phuket Island.
Salmoneus cristatus (Coutiere, 1897)
(Fig. 140)
Jousseaumea cristata Coutiere, 1897: 234
Salmoneus cristatus Holthuis, 1958:18; Banner & Banner, 1966:40;
1981:54
Description: The rostrum is triangular, having concave margins, the tip
reaches almost to end of antennular peduncle, the base is broad; the lateral
margins of rostrum are continued on carapace to posterior level of eyes as
301
strong lateral ridges or keels, giving rostral base a plateau-like
appearance; the rostral carina extends posteriorly to almost middle of the
carapace; short acute triangular orbital teeth lie lateral to keel and are
directed straight forward. The eyes are completely covered by rostral base
and orbital teeth in both dorsal and lateral views.
Figure 140: Salmoneus cristatus. A,B- Anterior region, dorsal and
lateral aspect; C- Large cheliped, inner face; D- Same, inferior face; ESame, superior face; F,G - Small chelipeds; H- Second leg; I – Third
leg; J- Telson (from Banner & Banner, 1966)
The telson is 3.6 times as long as posterior margin is broad; anterior
margin is 2 times as wide as posterior margin, the sides taper uniformly; the
terminal cleft is slight, triangular, the posterior margin is armed on one side
with three strong spines and two pairs of setiferous bristles in terminal cleft.
302
The articles of antennular peduncle are short and broad, sub
equal in length, the stylocerite is heavy, curved, reaching to tip of the
rostrum; the scaphocerite has a short lateral spine, equal in length to broad
squamose portion and reaches to end of antennular peduncle; carpocerite
reaches the distal end of second antennular article; the basicerite has a
short, sub-acute, lateral tooth.
The first pair of legs is asymmetrical; the fingers of large chela are a
little shorter than the palm. Their opposing edges are lined with eight
meshing obtuse saw-like teeth, plus three additional teeth on the proximal
end of fixed fingers, which meet against sharp ridge of dactylus; tips of both
the fingers are curved, acute, crossing each other. Medial side of basal
portion of palm has a strong excavation to accommodate carpus when the
carpal-propodal articulation is flexed; excavation is demarked by heavy
rounded projection of medial margin of palm; the merus is slender, concave
to accommodate palm when flexed; the internal margin is armed distally
with a rounded tooth. The small cheliped is slender, shorter than the
following thoracic legs, one-half as long as the large cheliped. The merus is
seven times as long as broad. The carpus is as long as the merus slightly
expanded distally. The chelae are three times as long as broad with fingers
equal to palm.
The carpal articles of second legs have the ratio of
10:1.7:1.7:2.7.The ischium of third leg is about as long as the merus; both
are unarmed, the carpus is slender, the superior margin terminates in an
obtuse projection; the inferior margin has a short spine. The propodus is as
long as the carpus carries four movable spinules on its inferior margin and
one pair distally; the dactylus is 0.5 as long as carpus, and slightly curved.
Colour: (See Holthuis, 1958:18)
Size: 10mm
Locality: Manora Island
Distribution: Red Sea,
Thailand. This is the
Israel,Kenya,Seychelles Is.Queensland and
first record of species from Pakistan.
303
Genus Synalpheus Bate, 1888
Diagnosis: Rostrum short and spiniform. Supra-orbital spines
smaller or equal to the rostrum. Eyes concealed under carapace. Upper
flagellum of antennule feebly biramous. Stylocerite well developed.
Squamose portion of scaphocerite shorter than lateral spine. First pair of
pereopods robust and asymmetrical, large chela carried extended, entire
and smooth, ovaloid, much longer than the small chela. Carpus of the
second pereopod bears five segments. Last three pairs of pereopods short
and compressed laterally. (After Banner & Banner, 1968)
Type species: Synalpheus falcatus Bate, 1888(=Alpheus
comatularum Haswell, 1882)
Gender: Masculine.
Remarks: Banner & Banner (1975) opined that established groups
within the genus by Coutiere (1909) may be coherent enough to continue to
be recognized, so are no more adopted. Of the approximately 115 currently
recognized species, this genus is represented here by three species:
Synalpheus tumidomanus, S. neptunus and
S. thai, which can be
separated as follows.
KEY TO THE PAKISTANI SPECIES OF SYNALPHEUS
Antennal scale with blade well developed, overreaching midlength of lateral
margin---------------------------------------------------------------------------------------------------------------------------2.
Antennal scale with blade vestigial or absent--------------------------------S. neptunus.
2.Telson with posterior angles projecting posteriorly as pointed teeth nearly
half as long as remainder of telson --------------------------------------------------------------------------- S. thai.
-Telson with posterior angles produced into shorter teeth, usually not
reaching level of midlength of adjacent spines-------------------------------------------------------- S. tumidomanus.
304
Synalpheus neptunus (Dana, 1852)
(Pl. 5)
Alpheus neptunus Dana, 1852a:22; Henderson, 1893:436.
Synalpheus neptunus.-Banner & Banner, 1972:24, fig. 3;
Miya, 1972:72
Synalpheus neptunus neptunus.- Banner & Banner, 1975:
317, fig. 11; 1977:282
Synalpheus neptunus Chace, 1988:82
Description: The rostrum does not overreach the first antennular
segment; its apex is not upturned and narrower at the base than the orbital
teeth.
The sixth abdominal somite does not project posteriorly at either
side of base of the telson, the posterior margin is unarmed mesially; the
telsonic dorsolateral spines are stout, prominent, the anterior pair is situated
on the anterior half of the telson, the posterior angles are sub rectangular.
The stylocerite falls short of or overreaches the distal margin of first
antennular segment; the basal antennal segment (basicerite) has a
ventrolateral tooth reaching about to the level of tip of stylocerite, its dorsal
angle is rounded, the blade of scale is sometimes moderately developed,
sometimes vestigial.
The movable finger of major chela does not appreciably overreach
the fixed finger, the palm terminates distally in a rounded prominence at
level of articulation with movable finger; the minor chela finger bears lateral
rows of regularly spaced setae but without patterned row on extensor
surface, both fingers are broad, excavate, and terminate in a single tooth;
the second pereopod has 5 carpal articles; the dactylus of third pereopod is
biunguiculate, the teeth are sub equal in length, the flexor tooth is slightly
stouter than the extensor tooth, the segment neither excavates nor is
swollen on the flexor margin; the merus is unarmed.
The uropod has transverse articulation on lateral branch in mature
specimens.
305
Remarks: This seems to be a very variable species (Chace,
1988). The older and the only record of S. neptunus from Karachi as
Alpheus neptunus by Henderson (1893) was not relied upon by Banner &
Banner (1972, 1975). Present specimens show geographical variations
when compared with Thai and Phillipino specimens.
Colour: Yellowish green
Size: 9 mm in CL, 1 ovigerous female with bopyrid parasite in left
branchial chamber.
Localities: Manora, Bulleji
Habitat: shallow sub tidal to 70 meters, association with corals is
given in literature.
Distribution: Arabian Sea, Red Sea to Japan, Philippines,
Indonesia, and Australia;the species is being reported from here for the first
time .
Development: The mother shrimp maintained for larval study
under laboratory conditions by Gory et al (2009) was initially identified as S.
coutierei. It is mostly probable that S. tumidomanus and S. coutierei are
synonyms. Larvae described in 2009 by Ghory et al differ from those
described for S. tumidomanus by Ghory & Siddiqui (2001); these differences
indicate the presence of another species in our waters later identified as
A.neptunus.
Synalpheus thai Banner & Banner, 1966
(Fig. 141)
Synalpheus thai Banner & Banner, 1966:61; 1974:427; 1979:246;
1985:56; Chace 1988:87; Hogarth, 1989:110; Kazmi et al,
1990:322; Kazmi, 2003:241
Description: The rostrum is awl-shaped, but the rounded tip
reaches slightly beyond middle of visible part of first antennular article.
Slight carina is present at the base of rostrum and extends posteriorly to
about middle of eyes. The orbital teeth have triangular base and rounded tip
reach almost to the end of rostrum. The tip of the orbital teeth and rostrum
306
bear short setae, the rostrum is strongly up-turned when seen in lateral
view; the orbital hoods are slightly inflated.
Figure 141: Synalpheus thai. Animal in dorsal view (After Kazmi &
Tirmizi, 1990)
The abdominal pleura of male are short and bear angular
projections, while those of female are expanded and rounded. The telson is
1.5 times, as long as posterior margin is wide, anterior margin is 1.5 times
wide than posterior margin, the lateral margins are anteriorly convex. The
lateral angles extend as acute teeth measuring, one third of the entire
telson, the margin between teeth is slightly arctuate, and the postereo
lateral spines are small.
The visible part of first antennular segment is more than 1.5 times
length of second segment; the second segment is 1.4 times as long as wide
and third segment is smaller in length of second segment; the stylocerite is
slender, reaches beyond distal end of the first antennular segment. The
scaphocerite has a concave margin. The lateral spine considerably
overreaches thus differs from that of type material (Banner & Banner, 1966),
where it reaches the distal end of the first antennular segment. The
squamose portion is very narrow, reaches to middle of the second
antennular segment; the inferior spine of basicerite is relatively broad at its
base, the tip reaches to the middle of second antennular segment; the
superior spine is acute, reaches just past middle of first antennular segment.
307
The chelipeds are not sexually dimorphic. The large chela is 1/3
times as long as broad; the fingers occupy its distal 0.3; superior margin
bears proximal to dactylus, an obtuse projection which is directed upward
and forward; below this projection on inner face, another projection is
present composed of two rounded cusps; the dactylus is strongly rounded, it
is longer than the fixed finger, bears an acute ridge along its superior
margin. The merus is armed with a small tooth distally on its superior
margin.
The small chela is 2.6 times as long as broad. Its fingers are 0.7 as
long as palm, bear setae along their opposing faces. The merus is slender,
the carpal articles of second pereopod show the following ratio: 10:2:2:2:5.
The dactylus of small chela bears a characteristics pattern of setae here
besides the setae on the opposite face, which cross those of the fixed
finger. The third leg is stout, the ischium is unarmed; the merus is three
times as long as broad; its carpus is shorter than the merus; the propodus
bears 9 spines on its inferior margin; the dactylus is biunguiculate; the
superior unguis is gradually curved inward; the inferior unguis is 0.4 as long
as the superior unguis;
Size:
11mm in TL.
Habitat: Shallow sub tidal
Locality:
PachaBulleji
Distribution: Thailand, southern Philippines, South China Sea,
Pearl Harbour, Indonesia, Hawaii, and Oman
Synalpheus tumidomanus (Paulson, 1875)
(Figs. 142-146, Pl.4,5)
Alpheus tumidomanus Paulson, 1875:101
Alpheus tumidomanus var gracilimanus Paulson, 1875:101.
Synalpheus tumidomanus exilimanus Coutiere, 1909:10.
Synalpheus neptunus Coutiere, 1899:436
Synalpheus hululensis Coutiere, 1921: 374; Chopra,
1923:173 (as host); Ramadan, 1936:19; Holthuis, 1958:31
308
Synalpheus Theophane de Man, 1911:261
Nec Synalpheus tumidomanus Kubo, 1940:90.
Synalpheus tumidomanus Holthuis, 1958:29; Banner &
Banner, 1966:56; 1968: 275; 1975: 377; 1981:83; 1983:107;
Miya, 1972:63;1984:90; Bhuti el al., 1977:595 (larvae);
Kazmi & Kazmi, 1979: 153; Afzal et al, 1986: 337; Chace
1988:90; Ghory & Siddiqui, 2001:113 (larvae); Kazmi,
2003:241
Description: The carapace and abdomen are smooth; the rostrum
is rather short, generally the tip of the rostrum fails to reach the distal end of
the basal segment of the antennular peduncle; the supra-orbital spines are
sharply pointed and nearly as long as the rostrum, or they may be slightly
shorter than the rostrum. A prolongation is present at the base of the
rostrum.
Figure 142: Synalpheus tumidomanus. Male in lateral view
The telson is nearly one and half times as long as the sixth
abdominal somite, its posterior margin is strongly convex; the dorsal surface
309
is armed with two pairs of spines. A shallow longitudinal groove runs on
the entire length of the telson reaches slightly beyond the one-third portion
of the adjacent movable spine.
Figure 143: Synalpheus tumidomanus. A- Anterior region, dorsal view;
B-Telson and uropods; C-E- Maxillipeds;F.G Right and left legs.
The antennular peduncle consists of three sub-equal segments; the
basal segment is longer than the combined length of the distal two
segments; its disto-median angle bears a small spine, the lateral margin is
produced into a blunt process; the stylocerite is large and reaches to about
the middle of the second segment of the antennular peduncle; the lateral
spine of the scaphocerite extends well beyond the distal margin of the
squamose portion; it reaches slightly beyond the antennular peduncle. The
lateral or larger spine of the basicerite is shorter than the stylocerite and the
inner spine reaches nearly up to one-third portion of the lateral spine of the
basicerite. In a large series of specimens, size of lateral tooth may be
variable.
310
The mandible is well developed having two-segmented palp; the
incisor and molar processes are also well developed;
Deep notch divides the tip of the maxillulary palp into two unequal
fingerlike projections; the upper lacinia is narrow at the base but broad and
truncated apically; the palp of the maxilla is short and fingerlike; the lacinia
is bilobed; the upper lobe is stronger than the lower one. The
scaphognathite is well developed. The palp of the first maxilliped is long,
extending beyond the caridean lobe; the second maxilliped is as illustrated.
The third maxilliped over-reaches the scaphocerite; its tip is armed with few
stout spines; the ultimate segment is nearly five times as long as the
penultimate segment; the parapenultimate segment is nearly two times
longer than the ultimate segment; the exopod fails to reach the distal margin
of the parapenultimate segment.
Figure 144: Synalpheus tumidomanus. A-Right antennular peduncle in
dorsal view; B-Right scaphocerite in dorsal view.
311
Figure 145: Synalpheus tumidomanus. A-Right, mandible in dorsal
view; B- Right, maxillula in dorsal view; C-Right maxilla in dorsal view.
The first pair of pereopods is well developed and asymmetrical; the
large chela is 2.5 to 3.1 times as long as broad in the middle, fingers are
sub-equal in length; the movable finger is 3.3 to 3.7 times to the total length
of the chela; the fingers are more or less equal in size; both the fingers bear
tufts of setae; the palm and merus are nearly equal in size.
312
Figure 146: Synalpheus tumidomanus. A-D- Right second to fifth
pereopod respectively; E- Second right pleopod; E’- Same, appendix
interna further enlarged
The second pair of pereopods is symmetrical; the fingers are
slightly shorter than the palm; the carpus is five segmented; the first (or
proximal) segment is longer being only a trifle shorter than the combined
length of the remaining four segments; the following three segments are
about the same size, the last or the fifth one is considerably longer than the
three proximal segments; the ischium is about the same length as merus;
the third pereopod is stronger than the last two pairs of pereopods; the
merus is 3.5 times as long as broad in the middle; the propodus is longer
than the merus; the posterior margin of the propodus bears 7 to 11 spines;
the dactylus is biunguiculate; the propodii of fourth and fifth pereopods are
armed with spines on their posterior margins. Moreover, oblique rows of
serrated spines are present on the distal two third portions of the propodus
of fifth pereopod; dactylii of last two pairs of pereopods are also
biunguiculate.
The endopod of the first pleopod is small; the appendix interna in
females is situated at the distal half of the endopod; the tip of the appendix
interna bears a few rows of coupling hooks.
313
Remarks: This a species complex and is divided into three
separate faunal realms. Many nominal species and subspecies have been
proposed on small number of specimens of this species. We have one
specimen as infested with the bopyrid parasite on the right chamber and
other on left chamber.
Colour: Pinkish orange
Size: 18 – 22mm in TL (males), 22 – 26mm in TL (females)
Localities:
Manora Island and Bulleji
Distribution: Indo-Pacific: Gulf of Thailand, Phuket, South Africa,
Red Sea, Sri-Lanka, Persian Gulf, Pacific Islands: Marshalls, Caroline Is.,
Phoenix, Samoa, Singapore, Indonesia, Philippines, Japan, Australia,
Mediterranean and Arabian Sea.
Development: (Figs. 147-153) Developmental stages of
Synalpheus tumidomanus were obtained by Ghory & Siddiqui (2001),
description and figures of these larvae are taken from them.
Post larva I
Size: TL = 3.75mm.
Duration: 6 days.
Carapace- Carapace smooth with few minute setae, rostrum basally
broad and distally pointed, reaching more than 3/4 length of distal end of
first antennular article, with a single epigastric tooth; eyes stalked and very
prominent free from carapace, their orbits produced into teeth.
314
C
A
B
D
I
G
E
H
F
J
A‟
Figure 147: Synalpheus tumidomanus. Post larva I; A- Lateral view;
A’- Frontal view of carapace; B- antennules; C- antenna; D- mandible;
E- maxillula, F- maxilla;
G-I- maxillipeds I-III J- pleopod II.
315
Antennules- Peduncle 3-segmented, basal segment largest,
each segment with a few fine setae, distal segment bears endopod and
exopod as articulated flagella; inner flagellum (endopod) with four long
plumose and few fine setae; as well as outer flagellum (exopod) with three
aesthetascs and a few fine setae.
Antenna - Peduncle 3-segmented, nearly equal in size, distal
segment bears a segmented flagellum (endopod) with few fine setae;
scaphocerite (exopod) with pointed distolateral spine, spine is shorter than
the squamose portion, which bears 16 marginal plumose setae, lateral
margin semi concave.
Mandible- Incisor and molar processes well developed.
Maxillule Coxal, basial endites and endopod rudimentry.
Maxilla - Coxal and basial endites without setae; endopod with 1
seta, scaphognathite with 17 marginal plumose setae.
Maxilliped I - Coxopod, basipod and endopod rudimentary; exopod
with three terminal plumose setae
316
.
D
B
A
C
E
F
I
G
H
J
Figure 148: Synalpheus tumidomanus. Post larva II: A- dorsal view; BAntennule; C- Antenna; D- Mandible; E, maxillula ;F- Maxilla; G-IMaxillipeds I-III; J- Pleopod II.
Maxilliped II - Coxopod and basipod without setae; endopod 3segmented, distal segment with 3 plumodenticulate setae; exopod 2segmented, proximal segment with 1 plumose seta, distal segment with 2
terminal and 2 sub terminal plumose setae.
Maxilliped III - Coxopod and basipod without setae; endopod 3segmented, distal segment with three terminal simple setae; exopod 2segmented proximal segment with two plumose setae, distal segment with 4
terminal plumose setae.
317
Pereopods I-V - Biramous pereopod I, II chelate endopod and
exopod partially segmented; exopod of pereopod I segmented and with 2, 4
plumose setae, exopod of pereopod II broken; exopod of pereopods III and
IV segmented and with 2 and 3 terminal natatory plumose setae
respectively.
Abdomen - Six somites with rounded posterolateral angles.
Pleopod Pleopods I-V biramous developed on each abdominal
somite; exopod and endopod with 2-3 setae; appendix interna present on
each pleopod.
Telson - Posterior margin with 2 simple and 10 plumose setae;
uropod (Fig. A) endopod with 3-4 small simple setae; exopod with 20-23
long plumose setae on their margins.
B
A
D
C
J
G
H
F
I
E
Figure 149: Synalpheus tumidomanus. Post larva III: A- Frontal margin
of carapace; B- Antennule; C- Antenna; D- Mandible; E- Maxillula, FMaxilla; G-I- Maxillipeds I-III; J-Pleopod II.
318
Post larva II
Size: TL = 3.84 mm. Duration: 5 days.
Carapace - Carapace smooth except few minute setae, rostrum
reaching near to distal end of first antennular segment; rostral tip bear 2
sub terminal setae, eyes sessile.
Antennules - Peduncle 3-segmented; lateral margin of basal
segment produced into a blunt process (stylocerite) reaching middle of
second segment of antennular peduncle, each segment of peduncle bear a
few fine setae, terminal segment with exopod and endopod; endopod 14
segmented with few setae, exopod 10-segmented with 3 aesthetascs and a
few fine setae.
Antenna - Endopod several segmented with small setae; distolateral
spine of scaphognathite (exopod) long and reaching middle of second
antennular peduncle, mesial margin with 17 plumose setae.
Mandible - Well-developed; endopod bud present with one plumose
seta.
Maxillule - Coxal endite with 11 setae; basial endite with 11
cuspidate and 9 plumodenticulate setae; endopod with one seta.
319
D
A
B
C
G
J
K
I
F
H
E
Figure 150: Synalpheus tumidomanus. Post larva IV: A- Lateral view;
B- Frontal margin of carapace; C- Antennule;
D- Antenna; E- Mandible; F- Maxillula, G- Maxilla;
H-J- Maxillipeds I-III; K- Pleopod II.
Maxilla - Coxal endite without setae; basial endite bilobed with 16 +
15 setae; endopod with 1 seta; scaphognathite with 33 marginal plumose
setae.
320
Maxilliped I - Coxopod without setae; basipod with 23
plumodenticulate setae; endopod with 4 plumodenticulate setae; exopod
with 4 terminal plumose and 1 lateral marginal plumodenticulate setae.
Maxilliped II - Coxopod without setae; basipod with 2
plumodenticulate setae; endopod 5-segmented, distal segment with 33
plumodenticulate setae; exopod 2-segmented with 4 terminal plumose
setae.
Maxilliped III - Coxopod without setae; basipod with 2 setae,
endopod 3-segmented each with several small plumodenticulate setae;
exopod with 4 terminal plumose setae.
321
Figure 151: Synalpheus tumidomanus. Post larva I: A- First large
cheliped; B-E- pereopods II-V; Post larva II: F- First (left) large
cheliped; G-First (right) small cheliped; H-K- Pereopods II-V.
Pereopods I - Unequal in size, large chela with broad movable
finger as compared to immovable finger, pereopods I (Fig. 135G right, small
cheliped) equal in size, both fingers similar to each other and provided with
322
fine tufts of setae as in pereopod I. Pereopods II-V well developed,
sparsely setose; dactylus of III, IV and V pereopods biunguiculate; superior
unguis longer than inferior.
Abdomen - Unchanged.
Pleopods - Endopod and exopod with 8-10 plumose setae;
appendix interna with 2 coupling hooks present on each endopod of
pleopods.
Telson - Dorsal surface with 2 pairs of spines, posterior margin with
2 pairs of spines and 10 long plumose setae, uropod (Fig. 133B) endopod
and exopod with 30-32 and 22 long plumose setae respectively.
323
Figure 152: Synalpheus tumidomanus. Post larva I: A- First (right)
large cheliped; B- First (left) small cheliped; C-F-Pereopods II-V. Post
larva IV: G- First (right) large cheliped; H- First (left) small cheliped; IL- Pereopods II-V
324
Post larva III
Size: TL = 3.88 mm. Duration: 7 days.
Carapace - Unchanged.
Antennules - No change in armature except addition of
aesthetascs.
4
Antenna - Unchanged.
Mandible - Unchanged.
Maxillule - Coxal endite with 11 plumodenticulate setae;
basial endite with 16 cuspidate and 7 plumodenticulate setae; endopod
bilobed with 1 plumodenticulate seta.
325
Figure 153: Synalpheus tumidomanus Post larva I: A- Telson with
uropods, dorsal view; Post larva II: B- Telson with uropods, dorsal
view; Post larva III: C- Telson with uropods, dorsal view; Post larva IV:
D- Telson with uropods, dorsal view.
326
Maxilla - Coxal endite with 2 plumodenticulate setae; basial
endite bilobed with 17 + 23 plumodenticulate setae; endopod with 1 terminal
simple seta, scaphognathite with 33 marginal plumose setae.
Maxilliped I - Coxopod without setae; basipod with several setae;
endopod with 4 plumodenticulate setae; exopod with 4 terminal plumose
and 6 lateral marginal plumodenticulate setae.
Maxilliped II - Coxopod broken; basipod with 2 simple setae;
endopod 5-segmented with several setae; exopod with 4 terminal and 2 sub
terminal plumose setae.
Maxilliped III - Coxopod without setae; basipod with 1 simple seta;
endopod and exopod unchanged.
Pereopods - Unchanged.
Peapod - Endopod and exopod with 6-10 plumose setae.
Telson - Posterior margin with 2 pairs of spines and 10 plumose
setae; roped unchanged.
Post larva IV
Size: TL = 4.45 mm.
Duration: Died with in 1 day.
Carapace - Unchanged except slight increase in size.
Antennules - Unchanged except 5 aesthetascs on inner flagellum.
Antenna - Unchanged.
Mandible - Endopodal bud 2-segmented, distal segment with 1
simple seta.
Maxillule - Coxal endite unchanged; basial endite with 9 cuspidate
and 15 plumodenticulate setae; endopod bilobed with 1 terminal seta.
Maxilla - Coxal endite broken; basial endite bilobed with 12 + 24
setae; endopod with 3 plumodenticulate setae; scaphognathite with 45
marginal plumose setae.
327
Maxilliped I - Coxopod with 5; basipod with several and
endopod with 5 plumodenticulate setae respectively; exopod with 4 terminal
plumose and 1 lateral marginal plumodenticulate seta.
Maxilliped II - Coxopod broken; basipod without setae; endopod 5segmented with several setae; exopod unsegmented with 4 terminal and 2
subterminal plumose setae.
Maxilliped III - Coxopod and basipod broken; endopod and exopod
unchanged.
Pereopods - Unchanged.
Abdomen - Unchanged
Pleopod - Endopod and exopod with 10-11 plumose setae.
Telson - Unchanged; uropod unchanged.
328
3.1.8 Family Hippolytidae Bate, 1888
Diagnosis: Rostrum may be short, absent, supra-orbital
spine present, or absent. Eyes well developed, but not covered by
carapace and never extremely elongated. Mandible with or
without an incisor process and palp. Third maxilliped with or
without an exopod; exopods also absent from all pereopods. First
pair of pereopods short and rather heavy but not swollen. Carpus
of the second pair of pereopods sub-divided into two or more
segments. Telson gradually tapering towards the posterior end.
Five to six pairs of gills present, with a varying number of epipods.
(Modified from various sources).
Remarks: Christoffersen during 1987 to 1999 erected several
families within the family Hippolytidae, Chace (1997) endorsing
the establishment of these families merged them with the original
family name. Majority of the members belonging to this family are
inhabitants of shallow waters. They are arboreal, anchialine and
marine, live to a depth of 3803 m. It is intresting fact that
Hippolytids are associated with medusae which are rejected by
pontonins as commensal partner. Sexual dimorphism is often
marked; Hippolytidae is a large reasonably homogenous family,
originally 40 genera were assigned to the family, but the numbers
of genera were eventually reduced. In the new literature, 39-42
genera and 308 species are given as described (Bauer, 2004). Of
these Saron, Lysmata, Exhippolysmata, Hippolyte , Heptacarpus,
and Latreutes are represented from the Pakistan coast. We
follow here Chace (1997) in keeping Lysmata within
Hippolysmata, which however is retained as valid by Holthius
329
(1993).Chace decision however, was considered as
misconception and resurrection is expected(Acoz,2004) .
KEY TO THE PAKISTANI GENERA OF HIPPOLYTIDAE
1.
Arthrobranchs present at base of first four pairs of
pereopods. Mandible with 3 segmented palp, many segments in
carpus of second pereopods---------------------------------------------------------------------Genus Saron.
No arthrobranch at base of pereopods -------------------------------------2.
2.
Mandible with two-segmented palp; carpus of second
pereopod composed of six to eight segments ---------------------------------------------------------------------------------------------------------------------------------- Genus Heptacapus.
Mandible without palp ------------------------------------------------------------3.
3.
Mandible with incisor process------------------------------Genus
Hippolyte .
Mandible without incisor process----------------------------------------------4.
4.
Carpus of second pereopod composed of three segments ----------------------------------------------------------------------------------------------Genus Latreutes
Carpus of second pereopod composed of many segments-------------5.
330
5.
Upper antennular flagellum unequally biramous--------------------------------------------------------------------------------------------------------------Genus Lysmata.
Upper antennular flagellum uniramous---------------------------------------------------------------------------------------------------------------Genus Exhippolysmata.
Genus Exhippolysmata Stebbing, 1915
Diagnosis: Integument not rigid. Rostrum overreaching
antennular peduncle, armed dorsally and ventrally, without ventral
tongue like lobe extending ventrally from lateral carina. Carpus
with dentate crest in midline at base of rostrum, with marginal,
unarticulated antennal and pterygostomian teeth but without
supraorbital tooth, depressed frontal or orbital regions, or
branchiostegal tooth; abdomen with 1st pleuron entire, not
bifurcate; 5th pleuron posteroventrally acute, not denticulate; 6th
somite without prominent spines, without articulated plate at
posteroventral angle and pleuron not curving around base of
uropod. Telson tapering to sharp posterior end, bearing 2 pairs of
dorsolateral spines. Eye with eyestalk movable, not concealed by
carapace, cornea not noticeably narrower than stalk, without
ocellus. Antennules with stylocerite not in vertical plane, not bifid;
2nd segment without sharp, curved lateral tooth; 3rd segment
without dorsodistal tooth or movable plate, dorsal flagellum
slender, not short or brush-like. Antennal peduncle not
overreaching antennular peduncle, without 3 strong ventral
spines; antennal scale overreaching antennular peduncle, lateral
tooth not near midlenght, lateral margin not spinose. Mandible
without palp or incisor process. First maxilliped with caridean lobe
clearly discrete from exopodal lash, epipod bilobate. Second
331
maxilliped with terminal segment narrow and applied
somewhat laterally to preceding segment, exopod not unusually
wide, with nonbilobate epipod and podobranch. Third maxilliped
with distal segment not flattened, with exopod, epipod,
arthrobranch, and reduced coxal exite. Pereopods without
exopods, with terminally hooked epipods on 4 anterior pairs,
without arthrobranchs. First pereopod with fingers shorter than
palm, not terminating in distal spines, chela 1 1/3 times as long as
carpus, latter not excavated to receive propodus, ischium not
produced into unusually long saber-shaped process. Second
pereopods symmetrical, fingers not longer than palm, carpus
subdivided into 12-22 articles. Third pereopod with dactyl tapering
gradually to acute apex, flexor margin
armed with about 4
spines, dactylus and propodus not prehensile in functional males,
propodus not subdivided, carpus not conspicuously spinose.
Uropod with lateral branch bearing distolateral movable spine
flanked laterally and mesially by sharp tooth (After Chace, 1997).
Type species: Hippolysmata ensirostris Kemp, 1914:118
Gender: Feminine
Remarks: There are five species and two subspecies of
which only Exhippolysmata ensirostris ensirostris is reported from
here.
Exhippolysmata ensirostris ensirostris (Kemp, 1914)
(Figs.154-157)
Hippolysmata ensirostris Kemp, 1914, 118, 7, fig.1-4; 1916a, 403,
fig.5; deMan, 1929:128, fig.7; Kazmi & Kazmi, 1979: 155
Exhippolysmata ensirostris Balss, 1933, 85; Holthuis 1980:125;
Holthuis & Miquel, 1983:3; Dore & Frimbodt; 1987: 70; Kazmi &
Kazmi, 2004, 97
332
Hippolysmata (Exhippolysmata) ensirostris, Holthuis, 1947, 74;
Johnson, 1962:48
Exhippolysmata ensirostris ensirostris Chace, 1997:65
Description: The rostrum is longer than the carapace, it
extends beyond the antennular peduncle by nearly two third of its
length. The upper margin is armed with 9-18 teeth (including the
epigastric tooth), of which the posterior 7-12 teeth form an
elevated basal crest. Anterior to the crest, the rest of the margin is
armed with 2-6 widely separated teeth. The lower margin has 716 teeth. The carapace is armed with antennal and
pterygostomian spines, which are more or less equal in size.
The telson is about twice the length of the sixth abdominal
somite. The dorsal surface is armed with two pairs of posteriorly
directed spines, but without the trace of lateral teeth.
Figure 154: Exhippolysmata ensirostris ensirostris. Animal in lateral
view.
The eyes are well developed; the basal segment being the
largest and nearly equal to the remaining two segments together;
333
the median side of the basal segment bears a spine, situated
nearly at the middle; the stylocerite is rather broad and pointed at
the anterior end; it fails to reach the distal margin of the basal
segment of the antennular peduncle, which is longer than the
third. The scaphocerite is three times longer than broad; the
lateral side is slightly concave and terminates in a strong spine,
which is shorter than the squamose portion.
Figure 155: Exhippolysmata ensirostris ensirostris. A- Telson and right
uropod; B- Right first leg; C- Right second leg; D- Third leg, last three
segments
334
The mandible has neither incisor process nor palp. The
maxillulary palp is distinctly notched, forming two lobes; the inner
lobe is slightly larger than the outer lobe. The upper lacinia is
distinctly large; the palp of the maxilla is also well developed; it is
slender and finger-like, becoming narrower in the distal half; the
lobes of the lacinia are un-equal; the upper lobe is much broader
than the lower lobe; the scaphognathite is of moderate width.
Figure 156: Exhippolysmata ensirostris ensirostris. A- Left mandible
B-Left maxillula C-Left maxilla.
The palp of the first maxilliped consists of three sub-equal
segments; the middle one being the largest; it extends slightly
beyond the caridean lobe. The second maxilliped is of a typical
caridean shape. The third maxilliped is slender, extending nearly
up to the middle of the rostrum; the tip of the maxilliped is thorny;
the parapenultimate segment is about two and half times the
335
length of the penultimate segment; the exopod fails to reach
the end of the parapenultimate segment.
The first pair of pereopods is rather stout, shorter than the
other pairs; the fingers are nearly one-third the length of the chela;
the cutting edges are in contact throughout their length; the chela
is slightly longer than the carpus; the merus is about five times as
long as broad in the middle; the second pair of pereopods is
longer and slender and reaches to about the end of the
scaphocerite. The chela is small; the palm is shorter than the last
segment of the carpus and only a trifle longer than the fingers; the
carpus is divided into 15 to 16 segments, the merus into 7 to 9
segments; the segments of carpus and merus may exhibit a great
variation. The third pair of pereopods extends well beyond the
scaphocerite; the dactylus measures about one fourth the length
of the propodus and its posterior margin is armed with 3-4
spinules; the propodus is twice as long as the carpus; the
posterior margin of the propodus is armed with a row of spines;
the fifth pair of pereopods is stouter and somewhat shorter than
the preceding two pairs.
336
Figure 157: Exhippolysmata ensirostris ensirostris. A-C Right first to
third maxillipeds.
An appendix interna is present on each of the second to
fourth pleopods of female. The appendix interna of second and
third pleopods is long and tubular in shape, measuring a little less
than half the length of the endopod ,that of the fourth is
considerably shorter. The tip of the appendix interna bears
several rows of small coupling hooks.
Size: 70 – 76mm in TL
Habitat:
Creeks, shallow seawater
Distribution: Indo-Pacific: India, Sri Lanka, Akyab,
Myanmar, Sumatra, Java and Indonesia
337
Genus Heptacarpus Holmes, 1900
Diagnosis: Rostrum long, Antennal spine present.
Supraorbital spine absent. Mandible with incisor process and
two-jointed palp. Third maxilliped without exopod. Carpus of
second leg with seven joints. Telson with four to eight pairs of
dorsal spines.(Adapted from Hayashi & Miyake, 1968)
Type species:
Hippolyte palpator Owen,1839:89
Gender: Masculine
Remarks: Heptacarpus is composed of over 29 species,
which are found in shallow water habitat. Two ecological and
morphological groups of species have been observed, one is of
short stout shrimps, which cling tightly to stones, algae, and
sessile invertebrates, and other group is of active omnivorous
shrimps in tide pools (Bauer, 1984)Komai & Ivano, 2008).
The sole Pakistani species Heptacarpus
pandaloides is included in the intermediate group as active
percher flitting from spot to spot in their respective rocky intertidal
and sea grass habitat.
Heptacarpus pandaloides (Stimpson, 1860)
(Fig. 158)
Hippolyte pandaloides Stimpson, 1860:103(34)
Spirontocaris propugnatrix de Man, 1906:404; Kemp,
1914:93
Spirontocaris pandaloides Kemp, 1916:386
Heptacarpus pandaloides Holthuis 1947:44;Miyake &
Hayashi, 1968: 374 fig. 1; Hayashi & Miyake, 1968:136;
Kikuchi, 1968:180; Komai & Ivanov, 2008:31 (key) .
338
Description: The rostrum is very long, about 1.3 to 2.0
times as long as the carapace; the upper border is provided with
eight to ten teeth usually nine teeth, the posterior two teeth are
always situated on the carapace behind the orbit; the lower border
bears 9 to 12 teeth. The carapace is smooth and provided with a
well-developed antennal spine; the pterygostomian angle is
rounded.
Figure 158: Heptacarpus pandaloides. Animal in lateral view;
(After Hyashi & Miyake, 1968)
The abdomen is smooth, not geniculate at the third somite.
The posterior angle of pleuron of the fifth somite is pointed. The
telson bears five pairs of dorsal spines and there are 3 pairs of
spines on the apex of telson.
The eye is long and sub cylindrical with a rounded
ocellus.The antennular peduncle reaches nearly to proximal third
of the rostrum The stylocerite reaches the tip of first segment of
339
the antennular peduncle; the second and third segment bears
a small marginal spine. The antennal scale is long and broad; the
outer distal spine falls short of the lamella.
The third maxilliped has an epipod but is without an
exopod. There are no epipods present on legs. The first pereopod
is rather stout, extending to the first segment of the antennular
peduncle. The second leg reaches as far forward as the middle of
the antennal scale; the carpus is subdivided into seven joints. The
merus of third pereopod bears seven to nine spines. There are
five to seven outer spines on the merus of the fourth leg, and two
to four similar spines on the fifth pereopod. The dactyli of the last
three legs bear six or seven spinules excluding the terminal claw.
Remarks: The present authors have not seen the material
of this species. According to earlier workers there is a variation in
the state of pterygostomian angle. Males are smaller than the
females.
Size:
33-53mm in TL
Locality:
Mouth of River Indus
Habitat:
Between 3.5 and 30m
Distribution: Japan and China
Genus Hippolyte L each, 1814
Diagnosis: Rostrum long. Supraorbital and antennal
spines present. Mandible with incisor process. Third maxilliped
with exopod but no epipod. Carpus of second leg divided into
three joints. Propodi of last three legs sub prehensile in mature
male, and normal in female. Telson with two pairs of dorsal
spines. (After Hayashi & Miyake, 1968a).
340
Type species: By monotypy Hippolyte varians Leach,
1814:431
Gender: Feminine.
Remarks:
Thus far, only one of the 28 recognized
species of Hippolyte : H. ventricosa has been recorded from
Pakistani region.
Hippolyte ventricosa H.Milne- Edwards, 1837
(Figs.159-160,Pl.7B)
Virbius australiensis Stimpson, 1860:104
Hippolyte orientalis Heller, 1862:227
Caridina cincinnuli: Haswell, 1882:183
Virbuis australiensis: Haswell, 1882: 186
Hippolyte ventricosus: H.Milne- Edwards, 1837:371;
Kemp, :96 pl.2 figs.; 1916a: 391
Hippolyte ventricosa Holthuis 1947: 16,55;Barnard
1950:704; Johnson, 1961:48; Hayashi & Miyake,
1968:140; Kensley, 1970:115; Hayashi, 1982:192; Tirmizi
& Kazmi 1984:313; Bruce, 1986:635; Chace, 1997:66; D‟
Acoz, 1999:72
Description: The rostrum is as long as carapace; it bears
only one tooth on the proximal fourth of the upper and two, rarely
three, teeth on the distal third or fourth of the lower border. There
is a considerable variation in rostral formula; in male both upper
and lower borders are sub parallel; in females the distal half of the
upper border is rather concave and the lower border is somewhat
expanded; the apex reaches the tip of the antennal scale in
females and it reaches only the tip of first or second segment of
the antennular peduncle in males. There are some tufts of
341
plumose hairs on the several portions of the carapace and
abdomen; the supraorbital spine is strong and acutely pointed; the
antennal spine is separated from the supraorbital angle, which is
blunt but projected nearly to the antennal spine, the
branchiostegal spine is set far back from the anterior margin of
the carapace.
Figure 159: Hippolyte ventricosa. A- Animal in lateral view;
A’-G- Rostral variation (after Tirmizi & Kazmi, 1984)
The third abdominal somite is gibbous and geniculated; the
pleura of the fourth and fifth somite are bluntly pointed. The telson
is slightly longer than the sixth somite and bears two pairs of
spines on the distal half near the lateral margin.
342
C
A
B
Figure 160: Hippolyte ventricosa.A-Second leg;B-Third leg of male;Csame of female
The eye is large and cylindrical, extending to the middle of
the rostrum; the eyestalk is about three times as long as the
cornea. The antennular peduncle falls short of the rostral apex;
the basal segment, which is armed with an outer distal spine, is
longer than the combined length of the second and third; the
stylocerite is slender, sharp and reaches beyond the middle of the
basal segment; the second is as long as third; the outer flagellum
is stouter and thicker in males than in females, and composed of
seven to eight joints in females and about 15 joints in males. The
antennal scale is large and broad; the inner angle of the lamella is
strongly produced far beyond the outer spine. The mandible bears
no palp.
The third maxilliped extends to the tip of the basal segment
of the antennular peduncle; the ultimate segment is flattened, with
minute teeth along the apex; there is a short exopod on the base.
343
The first pereopod is robust, not reaching to the tip of
the third maxilliped. The carpus of the second pereopod is
subdivided into three joints. The third leg reaches forward about
the distal tip of the antennal scale; merus has three or four spines
on the outer margin; propodus has seven or eight pairs of spines,
of which in males the fourth and fifth or fifth and sixth pairs are
long, with some minute hooks on each distal anterior border, while
in females these spines are normal; dactylus has 9 to11 spinules
excluding the terminal claw on ventral margin. The fourth
pereopod extends forward nearly to the end of the third
maxilliped. The merus is provided with two spines near the distal
articulation; the carpus, propodus and dactylus are same as in the
third leg. The fifth pereopod reaches forward about the tip of the
first leg; merus is armed with only one spine near the distal
articulation; the distal three segments are same as in the former
two legs.
Remarks: In different localities, it shows some
morphological differences with regard to the rostral formulae.
Colour: The species changes the body colour according to
its habitat; when obtained from green algal belt, they were green
and grey in colour, when they were captured from Sargassum
belt, the body was of a brownish black.
Size: Up to 24mm in TL
Habitat: In sea grass, meadows and Sargassum Sea
weed community in littoral to sub littoral zone.
Locality: Pacha
Distribution: Indo-West- Pacific region; D‟ Acoz, 1999
opines that the species is only known with certainty from India,
most other IndoPacific records are probably based on other
species.
.
344
Genus Latreutes Stimpson, 1860
Diagnosis: Integument not rigid. Rostrum overreaching
antennular peduncle, with ventral blade conspicuously developed
and projecting posteroventrally between bases of antennules,
without tongue-like lobe extending ventrally from lateral carina.
Carapace without dentate crest in midline at base of rostrum,
without longitudinal lateral carinae, without numerous appressed
teeth on lateral surface, without supraorbital tooth or abrupt
depressions on frontal or orbital regions, without hepatic tooth,
usually with branchiostegal tooth and denticles on branchiostegal
margin but without pterygostomian tooth.
Abdomen with somites not dorsally carinate or posteriorly
dentate, 1st pleuron not ventrally bifurcate, 5th pleuron rounded,
not denticulate, 6th somite not armed with 7 strong spines,
without articulated plate on pleuron curving around base of
uropod. Telson not tapering to sharp point, armed with 1-3 pairs
of dorsolateral spines, posterior margin not rounded,
posterolateral angles not sharply produced. Eyestalk movable, not
concealed by carapace, cornea without ocellus. Antennules with
stylocerite not lying in vertical plane, not bifid; second segment
without sharp curved lateral tooth; third segment without sharp
dorsodistal tooth or movable plate, dorsal flagellum often short,
stout and brush-like. Antennal peduncle usually not overreaching
antennular peduncle, not armed with 3 strong ventral spines;
antennal scale overreaching antennular peduncle, without lateral
tooth near midlenght or small movable lateral spines. Mandible
without palp or incisor process. First maxilliped not discrete from
exopodal lash, epipod usually bilobate. Second maxilliped with
terminal segment semicircular or sub triangular and applied
obliquely to preceding segment, exopod rather broad in proximal
½ with nonbilobate epipod but without podobranch. Third
maxilliped with exopod and epipod but without arthrobranch and
345
coxal exite. Pereopods with epipods with terminal hooks on at
least anterior 3 pairs, without exopods or arthrobranchs. First
pereopods with fingers shorter than palm, movable fingers
terminating in 4, fixed finger in 3, strong and interlocking spines,
chela 1-2 times as long as carpus, ischium not produced distally
into saber-shaped process. Second pereopods symmetrical,
fingers shorter than palm, carpus subdivided into 3 articles. Third
pereopods with dactyl and propodus not prehensile in functional
males, propodus not subdivided, carpus not conspicuously
spinose. (After Chace, 1997).
Type species:
Hippolyte planirostris de Haan, 1844
Gender: Neuter
Remarks: Nearly 20 species are known in this genus, of
these two are known from Pakistan, which can be separated from
the following key.
KEY TO THE PAKISTANI SPECIES OF LATREUTES
1.
Dactylii of last three pairs of pereopods
biunguiculate with conspicuous spines on margin---------------------------------L. mucronatus.
Dactylii of last three pairs of pereopods simple
claws, with microscopic spines on margin--------------------------------L.cf. anoplonyx.
Latreutes cf. anoplonyx Kemp, 1914
(Figs 161-163, Pl.5A&6D)
346
Latreutes anoplonyx Kemp, 1915: 104: 4, fig. 35;1916a, 339; de Man, 1929, 130, fig.8; Holthuis, 1947: 60; 1980:
128; Liu, 1955:42 pl.15.fig 15; Hayashi & Miyake, 1968
a:14;1968b:149 fig.2 & 4-b; Kazmi, 1971b: 417; 2003: 24; Kazmi
& Kazmi, 1979:155; 2004: 97; Chace, 1997:69; Yang, 2005:462
(larvae).
Description: The rostrum extends beyond the
scaphocerite and is triangular in lateral view. The epigastric tooth
is situated on the dorsum well behind the level of orbital angle. All
the teeth except the epigastric are situated on the distal half of the
rostrum. There are sexual differences in body length, rostrums
(the shape as well as spinulation) and the length of the upper
antennular flagellum; the rostrum is triangular in females and
much narrower except in one specimen where it is short and
broad, it is longer in males than in females; the upper border
has14-18 closed- set teeth and 9-15 on the lower border of the
rostrum in females and 7-10 teeth in three groups on the upper
border and 7-8 on the lower border in males; the tip in both sexes
in blunt and serrated not mentioned in original description but
given by Bruce(1986)for his Latreutes spA. The antennal spine is
strong and articulated on to the suborbital lobe; besides this the
pterygostomian angle is armed with a series of small spines
forming a comblike structure which may vary from 8 -13 teeth.
347
Figure 161 A: Latreutes anoplonyx. A- Ovigerous Female; B- Male,
(From Hayashi & Miyake, 1968)
The telson is slightly longer than the sixth abdominal
somite; it tapers posteriorly, ending in a median spine; the
posterior margin is flanked by two pairs of spines, of which the
inner one is smaller.
The eyes are short, reaching only up to the middle of the
first segment of the antennular peduncle; the antennular peduncle
is very short, consisting of three sub-equal segments; the basal
segment as usual is the largest and nearly equal to the remaining
two segments; the stylocerite is semi-circular in shape; the anterolateral side of the basal segment is produced into a spine; the
second segment is smallest, it is broader than long.
The lateral margin of the scaphocerite is more or less
straight may end in a sharp spine, which extends beyond the
squamose portion.
348
The mandibles bear neither incisor process nor palp;
the palp of the maxillula ends in a swollen tip; the upper lacinia is
broad and flat, whereas, the lower lacinia is much narrower and
strongly curved upwards; the maxillulary palp is small, slender and
fingerlike, becoming narrow and pointed at the tip; the inner
lacinia is divided into two lobes; the scaphognathite is large and
moderately broad.
349
Figure 161 B: Latreutes cf. anoplonyx. A,D- Carapaces of females,
lateral view; A’- Tipof rostrum; B- Right antennular peduncle; C- Right
antennal scale;E- Carapace of male; F- Telson tip; G- Distal two
segments of third leg
350
The palp of the first maxilliped far exceeds the caridean
lobe; the lateral side of the caridean lobe is convex; the ultimate
segment of the second maxilliped is laterally attached with the
penultimate segment.
Figure 162: Latreutes cf. anoplonyx.cf. A- Maxillula B- Maxilla;
C- First maxilliped; D- Second maxilIiped; D’ – Same, last three
segments; E- Third maxilliped
The third maxilliped extends well beyond the antennular
peduncle; the tip and distal part of the median margin of the
ultimate segment is spinose; the ultimate segment is nearly twice
the length of the penultimate segment; the parapenultimate
segment is one and half times longer than the ultimate segment;
the lateral margin of the parapenultimate segment is armed with a
row of spinules on the distal half; the exopod fails to reach the
middle of the parapenultimate segment.
351
Figure 163: Latreutes cf. anoplonyx. A- Appendix interna of second
pleopod; B- Same, fourth pleopod; C- Same, fifth pleopod
The first pair of pereopods is rather stout and shorter than
the others. It extends up to distal end of the parapenultimate
segment of the third maxilliped. Fingers are slightly shorter than
the palm, meeting only at their tips. The merus is nearly two times
longer than the finger. The entire chela is as long as the middle
segment of the carpus. The carpus is three segmented; the
middle one is the largest being nearly two times the length of the
first as well as the third segment. The last three pairs of
pereopods are similar in shape. The dactylus of each of the last
three pairs of pereopods is short being one-third the length of the
propodus, moreover, it is claw like and may be armed with 4-5
microscopic spinules on its posterior margin otherwise wholly
unarmed; the propodii each is armed with five spines on the
posterior margin.
352
An appendix interna is present on each of the second
to fifth pairs of pleopods, only their tips are capped with rows of
coupling hooks.
Remarks: The new collection of Latreutes having both
males and females helped us to reevaluate the earlier reports of
L. anoplonyx from India as new species by Kemp (1914) then
from Pakistan by Kamzi (1971) both basing on a single female.
The characters in sexual differences particularly in the shape of
rostrum shown by Hayashi & Miyake (1968) for Japanese L.
anoplonyx are not to the same extent in the present material, here
the female rostrum is similar in shape to that of Japanese males
(Fig.161), also the length and armature of rostrum in Pakistani
specimens is different. There are other differences therefore the
Pakistani specimens may turnout to be a new species. The
commensalism also seems to be rather loose than in L. anoplonyx
as free swimming individuals are also collected.
Size:
48mm in TL
Localities: Keti Bundar,Dubba,Makri creeks (one specimen
with isopod parasite).
Habitat: often associated with medusae.
Distribution: Indo-Pacific: India, Myanmar, Indonesia,
Philippines, Northern and Central China, and Japan .
Latreutes mucronatus (Stimpson, 1860)
(Fig. 164)
Rhynchocyclus mucronatus Stimpson, 1860:28
Latreutes mucronatus Kemp, 1914:101; 1916:396;
Barnard, 1950:706; Hayashi & Miyake, 1968:16; Titgen,
1982:82 (unpublished): 85; Ghani & Tirmizi: 1991:230;
Bruce, 1994:750; Chace, 1997: 69; Kazmi, 2003: 241
353
Description: The carapace is strongly arched dorsally.
It is not carinate in the median line but bears, as a rule, a single
stout fixed tooth behind the base of the rostrum: in rare instances,
three or four teeth are found in this position. There is a sharp
antennal tooth and a series of small spines, usually 11- 14, on the
antero-lateral margin.
Figure 164: Latreutes mucronatus. A- Female; B- Rostrum; C- Telson,
tip; D- First left leg; E- Second left leg; (Modified from Ghani & Tirmizi,
1991 )
In the female the rostrum reaches almost to or a little
beyond the end of the antennal scale. At its base it is anteriorly,
excavate for the accommodation of the eye. Anteriorly the rostrum
is sometimes almost circular in outline, but more often, it is
distinctly pointed and some times slightly upturned. The dorsal
and slightly upturned ventral teeth are borne only in its distal half.
In the male the whole form of the animal is far more
slender than the female, the rostrum is longer and much narrower
354
in lateral view; it extends well beyond the apex of the antennal
scale and with dorsal teeth 8-12 and 5-7 ventral teeth.
The last abdominal somite is about twice the length of the
fifth; the telson bears two pairs of dorsal spinules and terminates
in a narrow pointed process flanked by a pair of spines on either
side. The innermost of these is more than twice the length of the
outer and is often nearly twice as long as the median process
The eyestalk is a trifle wider than the cornea and bears a
conspicuous pointed process on its inner distal aspect. The
antennal scale is about three times as long as wide.
The third maxilliped reaches a little beyond the antennal
peduncle.
The second pereopods reach about to the apex of the
rostrum; the carpus is divided into three segments, of which the
first and third are approximately equal each being about half the
length of the middle segment; the palm is a little longer than the
last carpal segment and is decidedly longer than the dactylus; the
dactylii of the last three pairs of pereopods terminate in two stout
claws and bear three to five conspicuous spines on the posterior
margin. In the fifth pair the carpus is a little more than two-thirds
the length of the propodus; the dactylus is rather more than onethird the length of the propodus.
The exopod is shorter than the endopod and is about three
and a half times as long as broad.
Colour: Reddish brown
Size:
Female 4-5mm in CL, male 8.5mm in CL
Locality: Ibrahim Hydri
Habitat: Sea weeds (Kemp,1916); associated with
medusae ( Hayashi & Miyake, 1968).
355
Distribution: S. Africa, Persian Gulf, Red Sea, South
Coast of Arabia, India, China Japan, Korea, Thailand, Hong
Kong, Singapore, Indonesia and Australia.
Genus Lysmata Risso, 1816
Diagnosis: Integument not rigid. Rostrum armed dorsally
and usually ventrally, without ventral blade or tongue-like lobe
extending ventrally from lateral carina. Carapace not inflated, not
abruptly depressed
on frontal region, without numerous
appressed teeth on lateral surface, without supraorbital or sub
ocular tooth posterodorsal to orbital angle, latter not large or
obtuse, without hepatic tooth or branchiostegal tooth or denticles,
but with marginal, unarticulated antennal tooth and occasionally
pterygostomian tooth. Abdomen with 1st pleuron entire, not
bifurcate; sixth somite without prominent spines, without articulate
plate at posterovental angle and pleuron not curving around base
of uropod. Telson not tapering to sharp posterior end,
posterolateral angles not sharply produced, bearing 2 pairs of
dorsolateral spines. Eyestalk not concealed by carapace, cornea
not narrower than stalk. Antennules with stylocerite not in vertical
plane, not bifid; second segment without sharp, curved lateral
tooth; third segment without dorsodistal tooth on movable plate,
dorsal flagellum slender, not short or brush-like. Antennal
peduncle not overreaching antennular peduncle, without 3 strong
ventral spines; antennal scale not overreaching antennular
peduncle, lateral margin not spinose. Mandible without palp or
incisor process. First maxilliped with caridean lobe clearly discrete
from exopodal lash, epipod bilobate. Second maxilliped with
terminal segment narrow and applied somewhat laterally to
356
preceding segment, exopod not unusually wide, with
nonbilobate epipod and podobranch. Third maxilliped with distal
segment not flattened, with exopod epipod, arthrobranch, and
reduced coxal exite. Pereopods without exopods, with terminally
hooked epipods on 4 anterior pairs, without arthrobranchs. First
pereopod with fingers shorter than palm, not terminating in distal
spines, chela ¾ - 2 ¾ as long as carpus, latter not excavate to
receive propodus, ischium often produced into long saber-shaped
process. Second pereopods symmetrical, fingers not longer than
palm, carpus subdivided into 13-36 articles. Third pereopod with
dactyl and propodus not prehensile in functional males. (from
Chace, 1997)
Hippolysmata vittata Stimpson, 1860
Type species:
Gender: Feminine.
Remarks: A genus with diverse socio-ecological 36
species,only one recognized is known from here. The report of L.
amboiensis from Karachi by Ali (2006) can not be relied upon as
the specimens are not available for study.
ata vittata (Stimpson, 1860)
(Figs. 165-169, Pl. 7A)
Hippolysmata vittata, Stimpson, 1860: 26; Kemp, 1914:113, pl.6,
figs. 9-10; 1925:330; Gravely, 1927:137; Suvatti, 1937: 48;
Barnard, 1950:710
Nauticaris unirecedens Bate, 1888: 608, pl. 110, fig. 1
Lysmata (Hippolysmata) vittata Kubo, 1951:284
357
Hippolysmata (Hippolysmata) vittata, Holthuis, 1947, 20,67;
Johnson, 1961,48; Hayashi & Miyake, 1968,156,fig.17; Karim &
Rehman, 1974: 26, fig. 1-6; Bruce, 1986:601
Lysmata vittata Chace, 1997: 78; Kazmi, 2003: 241; Kazmi &
Kazmi, 2004:97
Description: The rostrum is considerably shorter than the
carapace it extends as far as the middle of the basal segment of
the antennular peduncle. It is directed slightly downwards except
at the tip, which is pointed upwards. The rostral teeth are variable
as already been pointed out by earlier workers. Six to fourteen
teeth are present on the upper margin and three to five on the
lower margin. The epigastric tooth is separated by a considerable
interval from the rest of the series, besides the epigastric tooth,
one or two teeth may be situated on the carapace. The carapace
is armed with strong antennal spine, which is separated from the
pointed suborbital angle, and a pterygostomian spine, of which
the antennal spine is stronger.
Figure 165: Lysmata vittata. Animal in lateral view
358
The posterolateral angle of each of the first four
abdominal pleura is broadly rounded, whereas that of the fifth is
acute and pointed.
The dorsal surface of the telson bears a shallow
longitudinal groove, from the base of which arises a bunch of
setae. Moreover, it has two pairs of posteriorly directed spines.
The eyes are well developed and reaching nearly up to half
the length of the rostrum.
The antennular peduncle consists of three sub-equal
segments. The second and third segments are more or less of
about the same size. The distal margin of each segment bears a
row of spinules.
The scaphocerite extends beyond the antennular
peduncle. The lateral side of the scaphocerite is more or less
straight and terminates in a strong spine, which extends only a
trifle beyond the squamose portion.
359
Figure 166: Lysmata vittata. A- Lateral view of carapace; B-Dorsal view
of telson and left uropod; C- Eye from right side of body.
The mandible of the right side differs from that of the left
side. The molar process of the right side bears a row of several
teeth, whereas, that of the left has only two, both of which are
large and sharply pointed teeth, the median surface (molar area)
is depressed and traversed by transverse ridges.
The palp of the maxillula is large and distinctly bilobed; the
palp of the maxilla is slender and finger-like, becoming narrower
toward the tip; the lobes of the inner lacinia are sub-equal; the
upper lobe is somewhat broader than the lower one. The
scaphognathite is large and moderately broad.
The palp of the first maxilliped is divided into three
segments; of which the middle one is the largest, moreover, the
palp extends a little beyond the caridean lobe; the outer margin of
360
the caridean lobe is somewhat convex; the second maxilliped
has the typical caridean structure; the third maxilliped is slender,
extending beyond the tip of the rostrum by the ultimate segment,
the parapenultimate segment is about three times the length of
the penultimate segment, the exopod fails to reach the distal
margin of the parapenultmate segment; the tip of the maxilliped is
thorny; the three segments have setae which are arranged in
small bunches (3-4 in each bunch). In addition to these, few long
setae are also present.
The first pair of pereopods is rather stout, shorter than the
others; but surpassing the end of antennular peduncle by chela;
the fingers of which are of equal size, meeting only at their tips.
The second pair of pereopods is long and slender. It reaches to
about the length of the scaphocerite. The chela is small. The
fingers are nearly equal to the palm. The segments of the carpus,
merus and ischium vary in number; the carpus may have 28-32
segments.
Figure167: Lysmata vittata. A- Right antennular peduncle; B- Left
antennal scale; C- Right mandible; C’- Left mandible; C’’- Same, molar
area
361
The third pereopod extends well beyond the tip of the
rostrum by the distal three segments; the merus is twice as long
as the carpus and bears 4 to 6 spines on its posterior side; the
merus of the fourth pereopod also has 4-6 spines, whereas, that
of the fifth pereopods has only two spines; the dactylus of each of
the last three pairs of pereopods is biunguiculate and with 4-5
spinules on its posterior margin.
Figure 168: Lysmata vittata. A- Right maxillula; B- Left maxilla;
C- Right first maxilliped; C’- Same, palp; D-E- Left second and third
maxillipeds.
The pleopods are biramous. In males, the endopod of the
second pleopod bears appendix masculina and appendix interna.
The appendix masculina is smaller than the appendix interna. On
the tip of the appendix interna, a few rows of coupling hooks are
present. Only an appendix interna is present on each of the last
three pairs of pleopods in males. In females, the second to fourth
endopods bear appendix interna only.
362
Figure 169: Lysmata vittata. A- E- Right, first to fifth pereopods.
Each of the first three abdominal sternites of the male has
a pair of upstanding spines, which are sharply pointed and curved
medially, the last three sternites are armed with a single spine,
and in females, and all the sternites are unarmed.
Size: 30-37mm in TL.
Colour: When fresh the body is translucent and marked
with numerous fine longitudinal striae of bright red colour.
Localities: Manora Island, Bulleji.
Distribution: Indo-Pacific: Red Sea, Persian Gulf,
Andaman Is., East Indies to Japan, Australia, and the Philippines .
Development:See Pillai(1974)and Bano(1999)
363
Genus Saron Thallwitz, 1891
Diagnosis: Integument not especially rigid. Rostrum
overreaching antennular peduncle, armed dorsally and ventrally,
with strong ventral blade, not projecting between bases of
antennules, or series of strong ventral teeth, without tongue-like
lobe extending ventrally from lateral carina. Carapace without
discrete dentate crest in midline at base of rostrum, without
longitudinal lateral carinae, without appressed teeth on lateral
surface, without supraorbital tooth, without abrupt depressions on
frontal or orbital regions, without sub ocular tooth posterodorsal to
orbital angle, latter not large, with antennal tooth, latter neither
sub marginal nor basally articulated, without hepatic tooth, with
branchiostegal margin not denticulate, with pterygostomian tooth.
Abdominal somites not dorsally carinate or posteromesially
dentate, 1st pleuron not bifurcate, 4th and 5th pleura pointed, not
denticulate, 6th somite not armed with 7 strong spines, with plate
articulated at posteroventral angle, pleuron not curving around
base of uropod. Telson not tapering to sharp point, posterior
margin sub truncate or slightly concave, and posterolateral angles
not sharply produced. Eyestalk movable, not concealed by
carapace, cornea with ocellus. Antennules with stylocerite not
lying in vertical plane, not bifid or semicircular, second peduncular
segment without sharp, curved lateral tooth; third segment without
sharp dorsodistal tooth or movable dorsodistal plate; dorsolateral
flagellum proximally stout but not unusually short or brush-like.
Antennal peduncle sometimes overreaching antennular peduncle,
not armed with 3 strong ventral spines; antennal scale
overreaching antennular peduncle, without lateral tooth near
midlenght or small movable lateral spine. Mandible with both palp
and incisor process. First maxilliped with caridean lobe quite
distinct from exopodal lash, epipod bilobate. Second maxilliped
364
with terminal segment elongate and applied somewhat
laterally to preceding segment, exopod not unusually wide, with
somewhat bilobate epipod and with exopod, epipod small
arthrobranch, and coxal exite, pereopods without exopods, with
epipods and arthrobranchs on 1st to 4th pairs, epipods with
terminal hook. First pereopod with fingers shorter than palm, not
terminating in interlocking spines, chela 1 ½ to 2 1/3 times as
long as carpus, carpus not deeply excavate for reception of chela.
Second pereopods symmetrical, fingers shorter than palm, carpus
subdivided into 10-17 articles. Third pereopod with dactyl not
gradually tapering to acute apex, armed with teeth on flexor
margin, dactyl and propodus not prehensile in functional males,
propodus not subdivided, carpus not conspicuously spinose.
Uropod with lateral margin of outer branch terminating in small
fixed tooth with larger movable spine mesial to it. (Modified from
Chace,1997).
Type species: Hippolyte gibberosus H. Milne Edwards,
1837
Gender:
Masculine.
Remarks:
Genus comprises of 4 known species, one
has been recorded from the Pakistani region.
Saron marmoratus (Olivier, 1811)
(Figs.170-174, Pl. 6C)
Palaeomn marmoratus Olivier, 1811:663;
Hippolite Krausii Bianconi, 1869: 2200
Hippolyte Hemprichii Heller, 1861:29,
Hippolyte Leachii Guerin Meneville, 1838: 37
Hippolyte marmoratus de Man, 1888: 533
365
Saron gibberosus, Thallwitz, 1881:100; de Man, 1902:
852; pl.26,fig.57; Parisi, 1919:74
Saron
marmoratus
Ortmann,
1894:15;
Borradaile,1898:1009; Coutiere, 1910: 1263; Kemp, 1914:
84; 1916:385; Kubo, 1940: 80; Holthuis, 1947: 25; Barnard,
1950: 688; Miyake & Hayashi, 1966: 144; Tirmizi & Kazmi,
1971: 283 (sexual dimorphism); Karim & Rehman, 1974:
25; fig. 1-6 Kazmi & Kazmi, 1979:155; Kensley, 1981:27;
Debelius, 1984:60; Kazmi,1996b:3; Chace 1997:89;
Description: It is difficult to give a description that would
be applicable to all the specimens under study.
Figure 170: Saron marmoratus. Animal in lateral view
The carapace is somewhat globular and smooth, but for a
few tufts of plumose setae, which are prominent on the mid-dorsal
line. A few setae may also be present on the lateral surface, near
366
the posterior margin. In preserved specimens, these tufts are
often wanting. The rostrum is longer than the carapace, it is
strongly curved upwards and armed on both of its margins. The
rostral formula being 1+6 / 6-7 beside the epigastric tooth, one or
in few cases, two teeth may be situated on the carapace. The
formula is more constant in females than in males. Beyond the
fifth tooth, the upper margin of the rostrum is unarmed for a
considerable distance. The last two teeth are minute and situated
near the tip. The lower rostral teeth are strong, gradually
diminishing in size towards the distal end. The lateral side of the
rostrum bears two prominent longitudinal carinae, of which the
upper one is very distinct and reaches well beyond the fifth upper
rostral tooth. The lower one is short and weak. The carapace is
armed with antennal, branchiostegal and pterygostomian spines.
The antennal spine is strong and well developed, while the
branchiostegal and pterygostomian spines are rather weak.
Figure 171: Saron marmoratus A- Carapace, lateral view; B- Right eye;
C- Right antennular peduncle; D- Same, ventral view; E- Right antennal
scale
367
The abdomen bears tufts of setae, which unlike the single
mid-dorsal row on the carapace are here arranged in two rows,
one on either side of the mid-dorsal line. The abdominal somites
of female bear more tufts of setae than those of male, in female
each sub-median row may have 1-2 tufts on the second, 3-4 on
third abdominal somite, whereas, in male a single tuft on the
second and one to three on the following somite. Further more,
the last two somites may be naked in the males, but in all the
females, available for study, there are two tufts of setae except in
one specimen (CL.15 mm), where they are wanting.
The telson is slightly longer than the sixth abdominal
somite. It tapers posteriorly, ending in a truncated tip, the
posterior margin is slightly sinuous and bears a small median
spine, flanked by two pairs of spines; inner one is smaller. The
sub-median spines are generally hidden by plumose setae. The
dorsal surface of the telson is armed with two pairs of spines. A
shallow longitudinal groove runs on the proximal half of the
telson. A bunch of long plumose setae arises from the base of the
groove and lies within it, some of the setae are nearly of the same
length as that of the groove.
368
Figure 172 Saron marmoratus. A- Left mandible; B- Left maxillula; CRight maxilla; D-F- First to third maxillipeds
The cornea of the eye is as wide as the maximum breadth
of the peduncle. The ocular spot or accessory eye is
semispherical, the basal segment of the antennular peduncle is
much longer than the combined length of the two following
segments and bears a small, sharp spine on its median margin.
The stylocerite is stout, sharply pointed and extending nearly up
to the distal margin of the ultimate segment, the second segment
of the antennular peduncle is slightly longer than the ultimate
segment when seen ventrally is produced forward into a sharply
pointed triangular process overlying the thick basal part of the
outer flagellum. The median margin of this process is slightly
sinuous. Moreover, a transverse row of small stiff setae is also
present in the middle of the segment.
369
The scaphocerite is slightly swollen at the base, narrow
in the middle, broad and truncated apically; the outer margin is
strongly concave and ends in sharp spine, which extends beyond
the squamose portion. The mandible consists of three-segmented
palp, an incisor process with four sharp, unequal teeth, and a
stout molar process. The molar process is partially sub-divided;
apically it bears two distinct tooth like projections and several rows
of short bristles. The maxillulary palp is rather small and broad; its
tip is truncated and setose apically; the upper lacinia is broad and
flat, its maximum breadth lies near the middle; the lower lacinia is
slender and curved. The scaphognathite of the maxilla is narrow,
particularly in its anterior half; the palp is slender and finger-like.
The palp of the first maxilliped is slightly longer than the
caridean lobe. The outer margin of the caridean lobe is strongly
convex. The second maxilliped is of typical caridean type. The
third maxilliped is one of those characters, which are subjected to
sexual dimorphism. However, in both the sexes the third
maxilliped reaches beyond the antennular peduncle; the
parapenultimate segment of the third maxilliped is nearly three
times as long as the penultimate segment; the anterior margin of
this segment is armed with spines - a strong dorsolateral and a
small dorso-median which is hidden by a bunch of long plumose
setae. Moreover, the lateral margin of parapenultimate segment
bears spines, which are arranged in groups of two to three and
situated more or less at equal distance. The antero-lateral margin
of the penultimate segment also bears a spine; the penultimate
segment is small being half as long as the ultimate segment; in
both the sexes the tip of the third maxilliped is thorny, spines
however are always more and rather stout in females than in the
males, the coxal median projection is subject to considerable
variation. It may be very small and only slightly projecting to
prominent elongated, broad and plate like. In females, the third
maxilliped is only a trifle longer than one third of the total body
length whereas, in males it is subjected to a greater range of
370
variation, being either more or even a little less than a third of
the body length.
Figure 173: Saron marmoratus. A-D- Endopods of second to fourth
pleopods of female A’-D’-Appendix internae of same; C- Tip of exopod
of second pleopods (After Tirmizi & Kazmi, 1971)
The males those have highly developed third maxilliped
also have the first pair of legs extremely long, sometimes attaining
monstrous size, approximately the same length as the total length
of the body. Here the ultimate segment is nearly four times the
length of the penultimate segment. In the males where the first
pereopod is highly developed, the fingers of the chelae are short,
stout, and strongly curved. Their tips are dark and notched so that
the two fingers can be interlocked. The gap between the fingers is
filled by long setae, arising from the cutting edges of the fingers.
371
Further more, the movable finger bears a large, rounded tooth
like projection, which is situated almost at the middle. It is easily
visible from the dorsal view, the palm is long, measuring
approximately three times the length of the fingers. In normal
males as well as in all the females the fixed finger differs being
less curved and hairy. A median coxal outgrowth similar to that of
the third maxilliped is present on the first pair of pereopods. In
females it is rather simple, short and pointed whereas, in males it
varies from a simple, spine-like projection to a large bilobed plate.
The first pereopods are underdeveloped in some males especially
which are bopyrized, (Kazmi,1996). The second pair of pereopods
is slender. The cutting edges of the fingers of the chelae are
unarmed; the fingers are slightly more than a third of the chela;
the carpus is sub-divided into 12-13 segments; the merus is
divided into two segments; the second pair of pereopods also has
a coxal outgrowth; it is rather simple in females. It may become
large and pointed whereas, in the males it is broad and plate like,
the outer margin is convex and serrated. The last three pairs of
pereopods are typical except in their armature on merii. In the
majority of the available specimens, the number of the spines is
reduced to one. The dactyli of the last three pairs of pereopods
are serrated.
372
Figure 174: Saron marmoratus. A- Thoracic sternites of male;
BSecond to fifth thoracic sternites of female. I- V- Coxae of thoracic
legs(After Tirmizi & Kazmi, 1971)
The endopod of the first pleopod is typical. In female, the
appendix interna is present on each of the second to fourth
pleopods, those on the second and third are small and knob like
and capped with rows of hooks.
In males, the appendix masculina on the endopod of the
second pleopod varies in size. In some males, it is well developed
and may or may not exceed the tip of the endopod very slightly;
the appendix interna of the third pleopod is wanting or hardly
noticeable in some males. The appendix interna of the fourth
pleopod, which though reduced, is always present. The earliest
stage is represented by a finger-like projection seen in a male
measuring in 5.5mm in CL.
In female, the second thoracic sternite has a pair of small
protuberances bearing a few plumose setae. Each half of the third
373
sternite has a transverse ridge, which becomes conspicuous
and heavily setose medially. The shape of the last thoracic
sternite of the female shows a considerable variation. It may be
slightly notched in the middle, the notch being flanked by
insignificant soft papilliform projections or short and hard spines or
the plate bears a single median spine; the abdominal sternites of
the females may be totally unarmed or may bear a single median
spine on the last somite.
In males, second to fifth thoracic sternites are armed. The
second thoracic sternite has a pair of small tooth-like projections,
pointed dorsally and slightly curved forward so that in posterior
view they are somewhat convex. The following sternite has similar
but larger projections. The fourth sternite has a pair of long,
narrow conical structures, which are strongly curved forwards.
Their posterior surface has a row of setae near the median
margin. The last thoracic sternite bears a pair of sharply pointed
spines, directed ventrally. Each of the first three abdominal
sternites has a pair of upstanding spines, which are sharply
pointed and curved medially ,each of the following three sternites
is armed with a single strong spine, directed posteriorly. The size
of these spines varies in different individuals, but in all cases, the
last spine is always smaller than two anterior ones.
Remarks: This dimorphism is one of gradual transition
rather than of true dimorphism, by which is implied either a
discontinuity in the development of the individual or a marked
dichotomy of evolution within the limits of a species (Kemp, 1914).
Colour: Fresh specimens are dark mottled, and marbled
with more or less circular-oval or crescent markings or rings of
varying shades of brown, not two specimens are exactly alike in
pattern.
Size: 38-83mm in TL (males), 65-81mm in TL (females)
374
Localities:
Mouth of Indus
Manora Island, Hawk‟s Bay, Bulleji,
Distribution: This species is spread over the whole Indowest Pacific region from Hawaii, Marquesas and Tahiti Is.,
Southward to Australia across the tropical Indian Ocean to Red
Sea and east coast of Africa.
Development: Complete larval series from first zoea to
seventh post larval stage has been obtained ( Ghory etal,2008,
abstract only).
375
3.1.9 Family Pandalidae Haworth, 1825
Diagnosis: Rostrum well developed, laterally compressed,
variable; antennular flagella simple, without accessory branches;
mandible with palp and with incisor and molar processes deeply
separate; first maxilliped with flagellum on exopod; second
maxilliped normal, 2 distal segments not arising side by side from
penultimate segment; first pereopod simple or microscopically
chelate; second pereopod chelate, carpus subdivided into 2 or
more articles; first male pleopod with endopod normal, not
unusually enlarged or elaborately convoluted (Modified from
Chace, 1985).
Remarks: This is a moderately diverse group including 24 genera
and 162 species of mainly epibenthic shrimps. Only Heterocarpus
has been recorded from Pakistan by Dore & Frimbotd (1987).
Therefore, this genus is further treated here.
Genus Heterocarpus A. Milne-Edwards, 1881
Diagnosis: Rostrum armed with teeth on both margins;
carapace without supraorbital spine, dorsally carinate nearly to
posterior margin, and with 1 or more longitudinal lateral carinae;
eye with cornea wider than eye stalk; second maxilliped with
terminal segment wider than long; pereopods with epipods on 4
anterior pairs, second pair distinctly unequal and dissimilar, left
376
member with 7 to 12 carpal articles, right member with 18 to
25 articles (After Chace, 1985).
Type Species: By original designation: Heterocarpus
ensifer A. Milne-Edwards, 1881:8
Gender: Masculine.
Remarks: No specimen from Pakistan is at our disposal. In
as much as information of occurrence in Pakistani waters of all but
one of the 20 species and subspecies assignable to this genus
has been available to us: H. tricarinatus Alcock & Anderson, H.
laevigatus Bate, H. woodmasoni Alcock and H. gibbosus Bate
have been reported in the Arabian Sea, H. laevigatus distribution
range includes Pakistan (see Dore & Frimbotd, 1986, map). Other
species are not dealt here with.
Heterocarpus laevigatus Bate, 1888
(Pl.7D)
Heterocarpus laevigatus Bate, 1888:636, pl. 112: fig.3; Alcock,
1899, pl.42: fig.1;1901:918; Rathbun, 1906:918; Barnard,
1950:684; Crosnier & Forest, 1973: 195,fig 61c; Kensley, 1972:
38; 1981, 28(list); Wilder, 1979:5, figs. 2,3; Moffitt, 1983: 435;
Chace, 1985: 33.fig; Hanamura & Takeda, 1987:107; Dore &
Frimbotd, 1987: 78; Poupin, 1996:8
Description: The rostrum varies from less than 1 ¼ to
more than 1 ½ times as long as the carapace in juveniles to
shorter than the carapace in adults. It is without any distinct lateral
carina and is armed dorsally with 6- 10, usually 6 or 7 teeth on the
carapace and basal part of rostrum, including 5, less commonly 4
377
or 6, on carapace posterior to the level of orbital margin and,
infrequently 1 situated directly above the posterior margin of orbit,
and ventrally with 6-13 teeth, most commonly 10 or 11; the
carapace is armed with blunt nearly complete intermediate carina,
an anteriorly sharp and posterior indistinct lateral carina, a short
and obscure anterior antennal carina, and a posterior sub
marginal carina.
The abdomen is unarmed dorsally, it is rounded on first
and second somite, and a blunt median ridge runs on third to fifth
somites, there is a slightly sulcate median area found on sixth
somite. The telson bears four pairs of dorsal teeth.
The antennal scale varies from little more than half as long
as carapace in large specimens to as much as three - fifth as long
in smaller ones; the blade and distolateral spine reaches
approximately to the same level.
The longer member of pair of second pereopods has 20
or 21 articles in carpus, shorter one has 6 or 7; the dactylus of
third pair is from nearly one- fourth as long as propodus, the carpi
of 3 posterior pairs are armed with 1 spine on third pair,
occasionally one on the fourth pair, and none on the fifth pair, the
meri are armed with 14-16 spines on the third pair, 6-19 on the
fourth, and 5-8 spines on fifth pair; the ischia are provided with
two spines on third and fourth pairs and are unarmed on fifth.
The first pleopods have bilobed endopods in males
(Modified from Chace, 1985).
Size: 50mm in Cl, 180mm in TL. This is probably one of
the largest members of the genus .
Habitat: The general bathymetric range is from 366-966m
Distribution: On Eastern Atlantic and in the Indo - West
Pacific.
378
3.1.10 Family Crangonidae Haworth, 1825
Diagnosis: Rostrum variable, usually short, depressed,
immovably attached to remainder of carapace; carapace often
sculptured with carinae, spines, or both; antennular flagella
simple, without accessory branches; mandible without incisor
process or palp; second maxilliped with terminal segment
semicircular, attached diagonally to penultimate segments; first
pair of pereopods subchelate, second pair, if present, with
undivided carpus. (After Chace, 1984).
Remarks: This is a moderately diverse family with 179
species and 20 genera currently recognized. Only two genera
Pontocaris and Philocheras are described from here. A third
genus Pontophilus includes a cosmopolitan P. gracilis has
Arabian Sea record also (see Dardeau & Heard, 1983, map).
Christophersen (1988) erected a subfamily Pontophilinae for
these genera. They can be separated with the help of the key
given below.
KEY TO THE PAKISTANI GENERA OF CRANGONIDAE
1Six or seven gills in each gill chamber, each with ventral
apex directed posteriorly.-------------------------------------------------------------Genus Philocheras.
- Eight gills in each gill chamber, each with ventral apex
directed anteriorly ----------------------------------------------------------------Genus Pontocaris.
Genus Philocheras Stebing, 1900
379
Diagnosis: Rostrum without lateral teeth in basal ½ of
length; carapace without postorbital longitudinal suture; eye with
well-developed cornea, pigmented; antennal scale with welldeveloped blade; 1st pereopod without trace on exopod; 2nd
pereopod chelate, usually overreaching merus of anteriorly
extended 1st pereopod.(After Chace,1984)
Type species: Selected by Holthuis, 1955:138: Crangon
nanus Kroyer, 1842 (Pontophilus) bispinosus Hailstone,
1835:271.
Gender: Masculine
Remarks:This is a large genus and our species
Pontophilus parvirostris and several other Arabian sea species of
Pontophilus have been placed in the genus Philocheras by Von
Vaupel Klein & Schram (2000). 38 to 45 are IndoPacific(Komai,2008). Two third species of genus Philocheras
apparently occur in depths of less than 50m.
Philocheras parvirostris (Kemp, 1916)
(Fig. 175)
Pontophilus parvirostris Kemp, 1916b: 372; 1925:341;
Kurian, 1954: 73; Kazmi & Kazmi, 1979: 156; 2004: 241; Tirmizi,
1980: 108
Philocheras parvirostris Jagadisha et al, 2000:582
(larvae);Han&Li,2008:47(larvae);Komai,2008:395(list)
Description: The rostrum is broad, triangular, and bluntly
rounded apically; it is deeply channeled longitudinally, the margin
380
forming a raised rim, which is contained laterally round the
orbits. In dorsal view, the carapace including the rostrum is a little
longer than broad the antennal spine is acute, the hepatic spine is
small. The branchiostegal spine is large and sharp; it is flanked by
a short carina and extends far beyond the level of the rostrum .
The telson is not sulcate above and is not much narrowed
distally; it has non-setose margin and without dorsal spinules; the
apex is formed by an acutely triangular tip, on either side at base
of the tip, a short spinule or one pair of large spines is present.
Figure175: Philocheras parvirostris. A- Carapace and eyes, dorsal
view; B- Telson; C- Antennular peduncle; D- Antennal scale;
EFirst leg; F- Second pleopod, (from Tirmizi, 1980)
The eyes are globular. The basal segment of the
antennular peduncle bears a spine at its outer distal end; its
lateral process is leaf like in outline and is pointed anteriorly;
second segment is equal to the third; the flagellae are sub
equal.The antennal scale is rectangular, outer margin is almost
straight and terminates in a large spine which does not reach as
381
far farwards as the rounded apex of the lamella. There are two
stout teeth on the basal segment of antennal peduncle.
The first legs do not posses exopod; the merus bears
externally a small procurved tooth a little behind its distal end; the
carpus is short and its inner angle is produced into several barbed
setae, the outer inferior surface has a spine. There are fine setae
on the cutting edge of the propodus and on margin of the merus.
The second pereopod has a remarkable chela.
The uropods are little longer than the telson; the exopod is
about three and half times as long as wide.
The thoracic sterna are broad posteriorly. The last four are
furnished with blunt carina in the middle, each of which terminates
anteriorly in a short spine. In front of them, a long and sharp spine
projects forwards between the bases of the first legs.
Size: 5mm in CL
Localities : Bulleji and Port Qasim
Distribution: Bay of Bengal, and S. India
Genus Pontocaris Bate, 1888
Diagnosis: Rostrum cleft apically, with one pair of lateral
teeth, at most; carapace with 7-12 teeth on first lateral carina, with
indistinct postorbital suture extending nearly to branchial region;
abdominal sterna armed with median spines; eye with cornea
well-developed, pigmented; antennal scale with well-developed
blade; first pereopod with exopod; second pereopod chelate,
overreaching merus of anteriorly extended first pereopod; 8 pairs
of gills, each curving anteriorly at ventral end (Modified from
Chace, 1984)
Type species: Pontocaris propensalata Bate, 1888:496
Gender: Feminine
382
Remarks: Of the ten species and one subspecies of
Pontocaris so far recognized(Chan,1996) two are present in the
Arabian Sea and a single species is present in our waters.
Pontocaris pennata Bate, 1888
(Figs. 176-180, Pl.7C)
Pontocaris pennata Bate, 1888:499, pl. 91; Kemp,
1916:12; Chace, 1984:43; Hayashi , 1986:145;
Kazmi,1972 (unpublished)137; 2003: 241; Sankolli &
Shenoy,1976:
(larvae);
Chan,1996:313;Han
et
al:2007:11;Han & Li, 2008:47 (larvae)
Pontocaris peunata Kazmi & Kazmi, 1979: 155 (Erroneous
spelling)
Aegeon pennata, de Man, 1920:294,pl.24: fig.70a-d.
Aegeon obsoletum Balss, 1914:76
Description: The rostrum is short, deeply notched at the
apex and extended up to the middle of the eyes; at the base of
the rostrum, each lateral margin is produced into an acute
process. The maximum breadth of the carapace lies at the front;
this is due to the enormous size of the anterolateral spines. The
carapace is armed with seven longitudinal carinae. The median
carina commences from a short distance behind the rostrum and
extends almost to the posterior margin. It is armed with a row of
eight prominent spines, which are directed anteriorly, the submedian carinae are convergent anteriorly. Each of the sub-median
carinea has nine spines. The lateral margin of the carapace is
formed by a row of spines, forming the intermediate carina, the
anterior most spine of which becomes greatly enlarged. The
383
lateral carina is not visible in the dorsal view. The antennal
spine is well developed.
Figure 176: Pontocaris pennata. Animal in lateral view
The anterior margin of the first abdominal somite bears
two mid-dorsal spines, each of which is produced backward into a
carina; laterally and in line with a sub-median carina of the
carapace, is a small acute process, behind which is another
similar process. The second abdominal somite bears a curved
spine situated at about a third from the anterior margin, it extends
backwards as a sharp mid-dorsal carina, which is continuous over
the following somites and finally ends as a small but sharp spine
on the posterior margin of the fourth somite. The fifth somite bears
two sub-median carinae, which diverge posteriorly and end in a
small spine. The carinae of the last somite run parallel to each
other, moreover, each carina has three posteriorly directed
spines, and another row of less prominent spines is situated,
lateral to each sub-median carina. The terminal spine of this
carina, however, ends in a well-developed spine. The pleuron of
the first abdominal somite is bluntly rounded; those of the
384
following two are narrow and acutely rounded. The posterior
margin of each of the last three somites is produced into spinose
processes. The remaining sculpture of the abdomen can be seen
in figure.
The telson is twice as long as as the sixth abdominal
somite. A shallow longitudinal groove runs on its dorsal surface.
The edges of the groove are raised into prominent carina. The tip
of the telson ends in a sharp point.
Figure 177: Pontocaris pennata. A- Carapace, dorsal view;
Abdomen, lateral view
B-
In natural position, the eyes are directed laterally and lie in
the orbit formed by the lateral margin of the rostrum and the
antennal spine. The cornea is reniform.
385
Figure 178: Pontocaris pennata. A- Telson and left uropod; B- Right
eye; C- Left antennular peduncle; D- Left antennal scale.
The antennal peduncle is rather short, the basal segment
is longer than the others are, its disto-lateral angle is produced to
form a tooth, and the stylocerite fails to reach the distal margin of
the basal segment. It is narrow and pointed apically, the second
segment is only a trifle longer than the third segment, which has
both its distal angles acutely produced, the third segment bears
two sub-equal flagella; the outer one is smaller and broader than
the inner one, the scaphocerite is short and broad, reaching
beyond the antennular peduncle. The lateral margin is straight
and ends in sharp spine, which is smaller than the squamose
portion.
386
Figure 179: Pontocaris pennata. A- Left mandible; B- Left maxillula; CLeft maxilla; D-F Right first to third maxillipeds.
The mandible is long and curved; it terminates in two sharp
spines. The maxillulary palp is slightly wider and notched distally;
the upper lacinia is broad, whereas, the inner one is narrow and
slender. The maxillary palp is characteristic, being swollen in the
middle, narrow and finger-like distally, the endopod of the second
maxilliped is four jointed. The distal two third of the exopod is
flagelliform. The third maxilliped is pediform, the distal part of the
exopod is also narrow and flagelliform like that of second
maxilliped fails to reach the distal end of the parapenultimate
segment and is nearly as long as the combined length of the last
two segments. Moreover, the inner margin is deeply concave near
the base. The ultimate segment is longer than the penultimate
segment; the tip of the maxilliped is armed with few microscopic
spinules.
387
The first pair of pereopods is symmetrical and subchelate. The anterior margin of the palm is oblique, deeply
notched to receive the tip of the dactylus; the carpus is short,
while the merus is slightly longer than the palm. The second pair
has two well-developed longitudinal carinae. The second pair of
pereopods is also symmetrical; the fingers are about half as long
as the palm. The carpus is entire and is longer than the merus.
The third pereopod is long and slender, reaching considerably
beyond the scaphocerite; the dactylus is styliform, long and
slender; the carpus is longer than the propodus; the merus and
ischium are sub-equal, the latter being the longer. The dactylus of
the fourth pereopod is nearly equal to the propodus, the propodus
bears longitudinal carinae; the carpus is shorter than the
propodus; the merus and ischium are nearly of equal size; the
dactylus of the fifth pereopod is shorter than the propodus, but
longer than the carpus; the propodus and merus are nearly of
equal size.
C
Figure 180: Pontocaris pennata. A-E- Left first to fifth legs;
Anterior portion of palm. n, notch
A’-
388
The endopod of the first pleopod is small. The endopod of
the second pleopod in females bears an appendix interna, capped
with rows of coupling hooks. In males, the endopod of the second
pleopod bears appendix interna and appendix masculina, both of
which are more or less of the same size.
The last thoracic sternite bears a ventrally directed spine.
Each of the first to fifth abdominal sternites in males is armed with
a single strong spine, directed posteriorly; in females, the sternites
are unarmed.
Size: 30- 35mm in TL (female), 33mm in TL (male).
Habitat: Highly benthic mainly living in soft bottom habitats
of sand, mud and gravel.
Locality: Fishing Area of Karachi
Distribution: Indo-Pacific, Red Sea, Persian Gulf.
389
3.2. Biogeography
Biogeography is studied based on range, population, and
species concepts, their environment and their interrelations.
3.2.1. Previous records: Turning to the zoogeographic
relationships of the shrimps under study, our entire conception of
biological diversity, biogeography is dependent on knowing how
many species are there and where they are found. For the review
of previous records of Pakistani carideans until to date, described
in detail in previous chapters, the period needs to be divided into
pre and post partition eras with misidentifications and mistakes in
literature remained uncorrected for long period. Some are still not
corrected.
In the following account, the species records are coming in
chronological order. The published caridean records from
Pakistan begin with a pasiphaeid Pasiphaea alcocki reported by
Wood-Mason&Alcock (1892) in prepartition era off the Sindh
coast. Several carideans were reported by Henderson (1893)they were four palaemonids: Exopalaemon styliferus
from
Karachi (Sindh Province), Macrobrachium scabriculum from the
River Indus, Macrobrachium, dayanus and Macrobrachium
altifrons from Lahore (Punjab Province) and two alpheidsAlpheus edwardsii and Synalpheus neptunus from Karachi
(Sindh Province). Another four species were recorded after a gap
by Kemp (1914), this time three hippolytids-Heptacarpus
pandaloides, Lysmata vittata and Saron marmoratus and one
palaemonid Exopalaemon styliferus was collected from Keti
Bunder (Sindh Province) in 1917 and Palaemon pacificus in 1925
by the same author from Karachi at the mouth of Indus.
Macrobrachium lamarrei was reported by Nath (1937) at Khalo
Garh (Multan), in the River Chenab (Punjab Province). An
390
offshore pasiphaeid Pasiphaea sivado was recorded by
Calman
(1939).
Exopalaemon
styliferus
from
Thatta,
Macrobrachium lamarrei and Palaemonetes sp. from Kotri near
G.M. Barrage, Palaemon sp. from Nara Canal and
Macrobrachium dacqueti from Sukkur to Keti Bunder, all of them
in Sindh province were first time reported by a Pakistani zoologist
(Qadri, 1960) in the post partition era. Macrobrachium altifrons
ranjhai was reported from Kabul River at Nowshera Kalapani
stream near Risalpur (NWFP), Salt Range, River Jhelum, and
River Ravi near Neaz Beg, Lahore (Punjab Province) by Tiwari
(1963). Ali (1968) reported some unspecified shrimp from Kohat
(NWFP) and Kurram Agency (FATA). In the same year Shakoor
reported
five palaemonids: Macrobrachium lamarrei from
Hyderabad, Khairpur (Sindh Province), Lahore, Sheikhupura and
Rawalpindi, (Punjab Province), M. dayanus from Sanghar,
Hyderabad (Sindh Province), Lahore Sheikhupura, Rawalpindi
(Punjab Province), M. scabriculus from Kotri (Sindh Province), M.
malcolmsonii from Sujawal, Thatta (Sindh Province), Kabul River
(NWFP) and M. dacqueti from Thatta (Sindh Province). A different
offshore pasiphaeid species Eupasiphae gilesi was recorded by
Tirmizi (1969) and in the same year Alpheus lobidens was
recorded upon by Tirmizi & Kazmi from Karachi. A new coastal
Alpheus species (A. albertai) and a deep-sea oplophorid
Acanthephyra exemia was reported by Kazmi, (1971a, b).
Johnson (1973) reported a subspecies of Macrobrachium
malcolmsonii- M. m. kotreeanum from Kotri (Sindh Province). A
processid Processa compacta (as Processa edulis crassipes)
was reported by Kazmi,1979(unpublished) and Kazmi & Kazmi
(1973) from Karachi, the same year Kazmi et al published record
of Automate dolichognatha as from Karachi. One alpheid Alpheus
lobidens, two hippolytids Saron marmoratus and Lysmata vittata
were reported upon by Karim & Rehman (1974) from the Karachi
coast. Salmoneus brevirostris was reported from Karachi by
Kazmi (1974) and Kemponia elegans was reported from Karachi
by Kazmi & Quershi (1974). Macrobrachium scabriculum ,M.rude,
391
M. lamarrei and M. malcomsonii were collected from the lower
Indus basin by Husain (1973), these species were also found in
Haleji Lake (Sindh Province) along with the atyids Caridella and
Cardinia (Baqai et al, 1974).
A pontonine Kemponia
seychellensis was reported by Kazmi et al (1975). Macrobrachium
naso, M. lamarrei, M. malcomsonii, M. naraensis (name only), M.
dayanus, M. rude, Exopalaemon styliferus were found at Kotri,
Rohri, Chilya Bund, Chahore Kalam, Kote Dhand (Sindh
Province) by Siddiqui (1976). In 1979, Banner & Banner reported
two alpheids from Astola Island (Balochistan Province). Kazmi &
Kazmi published a checklist in the same year reporting 12 new
records from different marine and brackish waters of Pakistan.
Four fresh water species were reported by Khatoon (1979) from
upcountry. Tirmizi (1980a) prepared a checklist of marine
decapods and stomatopods. Two marine species (Ogyrides
orientalis and Philocheras parvirostris) were added by Tirmizi in
the same year. A commensal pontonine Anchistus custos was
collected from a pinnid shell from Karachi by Tirmizi & Kazmi
(1982). Kazmi established a new Alpheus species in the same
year from Karachi. A new record of Hippolyte ventricosa was
made from Karachi by Tirmizi & Kazmi (1984). Yaqoob (1984,
1986) reported M. malcomsoni, M. idella, M.lamarrei, M.
scabriculum,
and
Exopalaemon
styliferus.
Synalpheus
tumidomanus, Alpheus splendidus, A. edwardsii and three new
alpheids were published by Afzal et al (1986) from Karachi. One
atyid Caridina nilotica was reported by Mehr et al (1988) from
Wah (Punjab Province). Three planktonic pasiphaeids, belonging
to the genus Leptochela from offshore waters of Pakistan were
reported by Kazmi et al (1990). The genus Kalriana of Atyidae
and its three new species from Kalri Lake were established by
Zuberi (1990). Latreutes mucronatus, a hippolytid was added to
the faunal list from Karachi by Ghani & Tirmizi (1991).
Macrobrachium equidens was collected from the Karachi Fish
Harbour by Kazmi et al (1991a) and a new record of Synalpheus
thai was published by Kazmi etal (1991b). Majid et al (1992)
392
reported Exoplaemon styliferus from brackish and estuarine
water of Sindh. Nematopalaemon tenuipes was reported by
Holthius & Miquel (1984) and then by Tirmizi & Kazmi (1995) from
Sindh brackish water whereas Palaemon sewelli was reported
from landings of Karachi Fish Harbour by Ghani (1999). Caridina
weberi was reported from the Karachi University campus by
Kazmi etal (2002). Kazmi & Kazmi (2002) while reviewing the
information upon shallow water palaemonids from Pakistan
brought Macrobrachium semmelinki to notice. An unpublished
report of the survey conducted by Ali (2005-2006) from Karachi
Scuba Diving Centre at Hawks Bay and Bulleji, Karachi (Sindh
Province) enlists four carideans: one hippolytid Lysmata
amboiensis, one hymenocerid Hymenocera picta, one
rhynchonetid Rhynchonetes uritai, and one gnathophyllid
Gnathopyllum americanum. Lately Leandrites celebensis and
Leptocarpus potamiscus were reported by Kazmi et al (2009) and
Kazmi & Kazmi (2009). The present study reveals several new
records and several new species await publication from Pakistan.
Laboratory rearing of some of the above mentioned
species has been done in Pakistan by Tufail & Hashmi (1956),
Barkati (1980), Yaqoob, (1980,1987, 1999) ,Tirmizi & Kazmi
(1986), Zuberi (1990) and Ghory etal (2001,2008,2009).
3.2.2. Discussion:
Biogeographically Pakistan is located at the cross roads of three of
the world‟s six zoogeographical regions, as such Pakistan does not
have a separate entity in biogeographically terms.. The provinces lie
in nine major ecological zones. As we see nothing has been done of
a zoogeographic nature of fresh water caridean shrimp, beyond
reporting where have been collected. Caridean biodiversity in fresh
water systems is distributed in fundamentally different patterns from
that in marine systems. They Bin the sea live in media that is more
or less continuous over extensive regions and species adjust their
393
ranges to some degree as climate or ecological conditions
change .But fresh water habitats are relatively discontinuous and
many freshwater species do not disperse easily across the land
barriers.
Table 2. Composition of the Pakistani known caridean
fauna.
Fauna
Families
Genera
Species
Marine Benthic
10
15
38
Pelagic coastal
1
1
4
Offshore
2
3
3
Sindh Province
13
9
75
Punjab Province
2
2
8
NWFP Province
1
1
2
Balochistan
Province
3
3
4
The species composition is discussed here with separately
as two ecological groups i.e. the fresh and brackish water, and
marine water forms.
3.2.2.1. Fresh and brackish water species:
The fresh water caridean shrimps are known almost exclusively
from the families Atyidae and Palaemonidae. There are however
a small number of freshwater shrimps from third family Alpheidae,
394
found in world deltaic region above the tidal limit in tidal areas
in a wide range of salinities from mesohaline to oligohaline to
almost pure freshwater, in freshwater caves, forest streams, in
acid water forest streams (Yeo & Ng, 1996), but one species
reported so far from Port Qasim may be considered in this kind.
Family Atyidae: In high mountains, which have almost
vertical waterfalls atyids prawns have not made their way there,
they have been denied access by the sheer impossibility of the
terrain. In all other kinds of waterfalls, even if the water is riparian
these prawns have found niches (in literature). We have no idea
of these prawns occurring at high altitudes in Pakistan. No such
study has been made on shrimps of this area due to
inaccessibility to these habitats.
The atyids are undoubtedly marine in origin. Their marine
ancestors must have invaded freshwater early in caridean history;
many atyids live in streams with high current flow. Adaptation to
freshwater has resulted in considerable variation in life history,
some have full, planktonic larvae, which are swept down streams
into the sea, juveniles, make the long migration in upstream,
some atyids have escaped their ancient marine bonds by passing
through abbreviated development or direct development. Man
made barriers do present a problem for juveniles (Bowles et al,
2000) thus effecting their abundance and distribution as is the
case of Sindh‟s most valued fish Tenualosa illisha which is
blocked to migrate due to ill designed barrages and dams as such
its catch has declined from 70% to 15% (WWF 2008). However,
this type of study is not made for any shrimp in Pakistan.
The family Atyidae occurs in lakes and streams in Sindh,
Balochistan and Punjab provinces in the genera Caridina ,
Caridella, and Kalriana. The genus Caridella is mentioned in
literature without any further details in a faunal list of Kalri Lake
(Sindh province) (Baqai et al, 1974). Since then it is encountered
nowhere. Caridina is represented by C. weberi, C. nilotica, and
395
C. babaulti. C. babaulti until now was known from central
India, in the east it extends to Malaya and in the west to Iraq. It is
said to be the only species found in permanently fast flowing
streams (Johnson, 1961). We have collected from the backwater.
C. weberi occurs in fast though seldom in torrential waters but not
confined to these part (in literature) tolerating oxygen from 3094.7%, pH 6.8 and alkalinity ø. 408 (Johnson, 1966). The species
whose eastern limit is Sumatra extends in the west up to Dubai
(Titgen, 1982).
No atyid was collected by the authors further west of Hub
Dam on Makran coast. Which otherwise makes a continuous belt
with Iran where the family is reported from S.W. Iran (Gorgin,
1996). Other atyid genera like -Paratya from Assam hilly areas
and Atyopsis (as Atya) from Orissa hilly areas are likely to occur in
northern areas in similar niches, as there are evidences of the
close affinity between Indian sub continental and Malayan faunas
(Tiwari, 1951).
Family Palamonidae: There is a consensus in the
literature that members of the family Palaemonidae are of very
recent origin, they disperse readily and as a result have little or no
value in the investigation of past geographical history (Strenth,
1976). However, Thompson (1967) gives evidence to support an
ancient origin of the super family Palaemonoidea. Tiwari (1955)
places the origin of the genus Palaemon and Macrobrachium
during the Pleistocene or post Miocene while the immigration of
palaemonids from marine into freshwater is of recent occurrence
(Sollaud, 1923). Fresh water species have acquired fresh water
habitat by migration from sea to the interior of land through rivers.
The process of adaptation is not yet complete because many
species are found in estuaries and still depend in brackish water
for breeding. Several have completely acclimatized to freshwater
and are found in inland rivers and hill streams. This idea is
reported as the Satpura hypothesis presented by Tiwari (1955),
supported by other authors.
396
The
paleomonids
belonging
to
the
genus
Macrobrachium have gradually evolved or are continuing to
evolve during their process of freshwaterisation (Jalihal et al
1993). These features have some significance from aquaculture
point of view. The family Palaemonidae has freshwater, estuarine,
and marine representatives in Pakistan. There are no truly marine
species within the genus Macrobrachium. Of the four great
palaemonine genera generally Palaemon is the most diverse in
marine waters, Macrobrachium is the most diverse in freshwater
and Palaemonetes is intermediate between the two in this
regards. Palaemon species are tolerant of low salinities commonly
ranging up into brackish water estuaries and bays and into fresh
water as well. The estuarine and marine species posses relatively
small and numerous eggs and go through a long and complex
larval development including as many as 13 stages before
reaching the juvenile stge. Strictly freshwater species posses
relatively few large eggs and go through abbreviated larval
development. As there is only one report of Palaemonetes from
Pakistan by Qadri (1960), according to him a sizeable population
it at Kotri was present at his time, there are probabilities that he
has mistaken Kotri‟s population of Macrobrachium for
Palaemonetes as juveniles of Macrobrachium can easily be
mistaken for Palaemonetes since they are of same size and
similarly found in shallow vegetated areas along the water edge,
not encountered in fresh collection.
Macrobrachium species: One cannot speak of the
palaemonids without marveling over the genus Macrobrachium.
They are apparently restricted to still waters at low levels with no
records above 500ft. Identification of taxanomic position and
species boundaries within the genus is important to obtain reliable
information for application in aquaculture and biodiversity
conservation programmes.A number of species of Macrobrachium
has a large range of distribution, some for instance range from E.
Africa or India to Peninsular Malaysia andTaiwan ( M.
397
scabriculum), and others are confined to smaller areas (M.
lamarrei, M. malcolmsoni, M. naso, M. dayanus) or known from
Indo-Burmese area. The genus Macrobrachium found in Pakistan
is associated with the Indus plain, which comprises about 40%
area of Pakistan drained by the river Indus and its tributaries. The
Indus water system is isolated from the Tibetan plateau and
separated from Brahmaputra watershed. However these barriers
have not effected on the dispersal of fish fauna of rivers of central
Asia states, Western China in Pakistan .Rafique (2001) have put
forward many explanations for this phenomena. No such study
has been made on shrimp of this area. As if atyids palaemonids
are also not successful in Balochistan i.e. the Palaearctic region
probably, they could not find their way to Balochistan Rivers due
to lack of interconnection of rivers of Balochistan with those of
other provinces. Besides there is an ecological limitation as the
temperature and salinity requirement of larvae that appears to be
one of the factors limiting the westward spread of Macrobrachium.
The presence of juveniles of Macrobrachium in the Hub Dam and
few sub adults from Lasbella District indicates survival of some
isolated populations. The Balochistan Rivers Hub, Porali, Hingol,
and Dasht share at high level the fish fauna with lower Indus
(Rafique etal, 2008), these authors have discussed the geological
history to explain this, and the same theories can be applied here
for shrimps.
Macrobrachium naraensis: It appears to be endemic to
Nara Canal Tharparkar Distt. (Sindh), initially Qadri (1960)
mentioned of presence of an undetermined species of Palaemon
near P. mirable Kemp from Nara Canal then Siddiqui (1976)
collected and reared a species which he named M. naraensis
naming after the locality, but did not describe it as the material of
any of the two authors is not traceable, it is assumed here that
they are the same taxa, the population seems to be dwindled as
no specimens were collected after these reports.
398
Macrobrachium dacqueti: This species has been
separated as valid species from the Asian stock of M. rosenbergii
species group. The recent distributional study of M. dacqueti by
de Bruyn etal (2004) suggest that its distribution follows Huxley‟s
Line, an extension of Wallace‟s Line which is a well known
biogeographical boundary that runs through Indonesia. However,
Wowor & Ng‟s study of 2007 shows that the distribution of M.
rosenbergii and dacqueti does not exactly conform to either
Huxley‟s or Wallace‟s Line. The easterly boundary line for M.
dacqueti begins after Palawan in the Philippines and Bali in
Indonesia. In the author‟s opinion the distribution of the two
species is more affected by the larval dispersal factors such as
high salinity tolerance combined with sea current patterns rather
than tectonic history. This species can be collected from Sukkur to
Keti Bunder from river Indus and its tributaries in different
salinities.
Macrobrachium scabriculum: The species occurring in
lower Indus is a typical tropical Indian form extending in the south
up to E. Africa. Shakoors report of 1983 from Kalmat need
verification.
Macrobrachium dayanum: This seems to be restricted to
Indo Burmese region and with other nine species in the area
forms one compact group from the phylogenetic point of view.
Macrobrachium
altifrons ranjhai: Tiwari (1963)
separated out the population of nominal species from Pakistan
and gave it subspecies name M. altifrons ranjhai confined to
Pakistan (Kabul River, all streams of Peshawar, Nowhere Road,
Risalpur, Indus River, and Kalabagh River, near Niaz Beg). New
localities have been found during the present survey. The
subspecies may be endemic to Pakistan.
Macrobrachium malcolmsonii: The populations at Kotri
were given a new status M. malcomsonii kotreeanum by Johnson
(1963); the species, a sister clade to M. rosenbergii ,is confined to
399
lower Indus basin down Kotri. The reports of M. malcolmsonii
from Kabul River and D.I. Khan are possibly based on the other
subspecies M. m. malcolmsonii.
Macrobrachium equidens: It is not a true freshwater
species but it can be called a brackish water species, it may be
collected from fishing area of northern Arabian Sea. It breeds in
mangroves bordered creeks. Smaller specimens are commonly
found in mangrove creeks. It has a wide range distribution in the
IndoPacific area reaching to Taiwan through mainland China.
Macrobrachium rude: It inhabits both fresh and brackish
water reported from Rohri-Kotri area. It has a somewhat limited
range worldwide starting from continent Africa not extending
beyond east coast of India.
Macrobrachium idella Idella: It was reported by Yaqoob
(1986) from Thatta as M. idae. It is characteristic of slow flowing
or still water, so likely to occur in other water body of similar
nature. Jalihal et al (1988) are of the opinion that true idae is
distributed only in East Africa and Peninsular Malaysia and not in
the subcontinent India where is appears to be replaced by idella,
the same is followed here.
Macrobrachium lamarrei lamarrei: It is one of the oldest
known species and widely distributed in the Indian subcontinent
described in two subspecies. This subspecies is found in the
deltaic area of River Indus and freshwater bodies of Pakistan. A
third subspecies was separated during the present study from
Karachi.
Macrobrachium siwalikense: It has been reported from
the hilly areas of Indian Punjab by the Indian authors, it is likely to
occur here also in the Dhok Pathan area of the middle Siwaliks of
Pakistan. No specimen is yet at our disposal as no survey has
been made in this region. This species with M. dayanum and
400
other seven species form one compact group of Indo Burmese
freshwater forms.
Macrobrachium shahpuri: This species is restricted to
the upper Indus basin streams and canals.
Macrobrachium naso: It was reported by Siddiqui (1976)
from Rohri, Kotri, Gharo Dhand, Indus River delta, and Kinjhar
Lake. The species is only known from Myanmar (Cai & Ng, 2002).
Its report in the Pakistani waters is although of some significance
but still Siddiqui‟s (1976) identification needs verification.
Nematopalaemon tenuipes: It is a littoral species, both in
brackish and marine waters of the Arabian Sea replaced by
another congener in the Bay of Bengal.
Palaemon semmelinki: It is distributed from Pakistan in
the west to Celebes in east. It occurs abundantly in shallow
marine sometimes-brackish waters of Pakistan.
Palaemon sewelli: It has a very discontinuous form of
distribution having one location in Europe (Portuguese), and then
in the Indo Burmese region (Pakistan, India and Myanmar). It
occurs in creeks.
Exopalaemon styliferus: It has a wide local range, found
in seepage water around Haleji Lake, in Kinjhar Lake in the deltaic
area spawning occurs in more saline areas and juveniles migrate
to the estuary, it has a limited Indopacific distribution not beyond
Indonesia in the east in west up to Pakistan.
Leptocarpus potamiscus: Since it is a brackish water
species extending in the east up to Sumatra is restricted to spread
further east into the Pacific area due to land barriers since they
could less easily circumvent it but a full satisfactory explanation of
other brackish water species that availed themselves of this route
and reached further east is not possible.
401
3.2.2.2. Marine water forms:
The western Indian Ocean fauna in general is the part of Indo
West Pacific region, which is a predominantly tropical and
subtropical marine region. The Arabian Sea being its
northwestern part in the Indian Ocean, biogeographically known
as Arabian Sea Region II. Its coast is considered the western
most extent of vast Indo- Polynesian province (Kelleher etal,
1995).
The northern and western boundaries are limited by
Pakistan, Iran, Arabia, and eastern Africa. Some zoogeographers
consider the Persian Gulf and Gulf of Aden as extension of the
western Indian Ocean. Some like Briggs (1974) do not. The
eastern boundary is loosely established by western India,
southward Islands Mauritius and Madagascar form the southern
boundary for benthic species, and benthic fauna does extend
further south along African coast due to the warm southern
flowing Agulhas current. Titgen (1986) divided the western Indian
Ocean into six basic faunal groups as provinces corresponding in
part with Briggs (1974) analysis. Pakistan comes in the West
Indian province more related to western Indian Ocean group than
to Malaysian Peninsula. Before this Montgomery (1930) divided
the western Indian Ocean arbitrarily in to five divisions for the
study of marine crabs.
According to Wyrtki (1971), the seasonally changing
monsoon gyre is not found anywhere else in the world oceans.
Other characteristics are the deepwater characterized by a weak
salinity maximum, the O2 content in both the bottom water and
deep being high but decreasing northward, nutrient concentration
generally increasing to the north. The Arabian Sea bordering
Pakistan has highly saline water, intensified by the outflow of high
salinity water from the Persian Gulf and the Red Sea. These water
masses of high salinity layer in the Arabian Sea prevent water of
402
southern hemisphere from penetrating efficiently into the
northern Indian Ocean.
Planktonic species of the Arabian Sea are affected by the
seasonal changes in the direction of the flow, the flow from the
Persian Gulf does not allow euryhaline species to survive
otherwise the mixing of epipelagic species with the Gulf area is
more obvious than the bathypelagic species, which are barred
due to shallow sills (Banse, 1968).
Achuthankutty et al (1991) working on neuston caridean
distribution in southwest monsoon found different families at
different layers. Without discriminating between caridean and
penaeoidean shrimps Hida & Pereyra (1966) informed that during
bottom trawling mostly the shrimp were taken in the 184-366m
depth range. The effects of intermitting upwelling and subsequent
fluctuations in temperature in the Arabian Sea is obvious and the
productivity is very high in the South West monsoon on the
eastern side (Saudi Arabian Coast) in the region of upwelling
consequently Hogarth (1987) analyzing the Omanis species found
70% increase in number of species in period following monsoon in
the western Arabian sea.
At our coast, the backwaters and their associated river
system form a large part of the inshore waters and influence the
hydrography of the coastal water considerably. These waters refer
to a system of shallow brackish water lagoon and swamps
behaving like estuaries, having a permanent connection with the
sea and so influenced by regular tidal rhythm. The above given
oceanographic features have an impact on ecology of Pakistani
shores and control the fauna. This can be seen by comparing the
low crustacean abundance in the Eastern Indian Ocean (Prasad,
1969).
The term pelagic used includes the species that are
understood to be inhabiting the main water body in sea. The
403
ecology of mesopelagic realm is very different from that of the
epipelagic.
a. Pelagic (Coastal and offshore).There are old records
and new collection from the pelagic and offshore realm via
expedition reports such as the John Murray and Fridjtof Nansen
sampling in the Arabian Sea ,the influx of the caridean fauna from
the Persian (Arabian) Gulf needs much more detailed study.
Presently adjacent gulfs Omanis Gulf, Persian (Arabian) Gulf and
Gulf of Aden faunas are known through Gurney (1937), Bruce
(1971), Motoh, (1975), Banner & Banner (1981), Titgen (1986),
Hogarth (1987) ,Holthius (1986, 1987), De Grave (2007) and
Anker & De Grave (2009). We are working on Discovery Cruise
(1997) samples and family Anchistioitidae is being added to the
Omanis Gulf fauna.
.
.
Family Pasipheidae: The small epiplegic pasipheid
shrimp Leptochela that is abundantly found near surface or near
the bottom during the day in Indo-west Pacific, eastern Pacific,
Atlantic Oceans, Red Sea, and Mediterranean .Nayeem et
al(1991) found adults and juveniles at near shore station(Manora
Channel) throughout the year but the offshore samples of Fridtjof
Nansen cruise had only subadults and adult females in less
number in front of Baluchistan coast than on Sindh coast.It as
been a source of attention to carcinologists for having a “disjunct”
distribution (Chace, 1976). The present studies include such
species which are up till now purely Atlantic, this is most unusual
that of a well studied genus like Leptochela (late L.B. Holthius
pers. comm. QBK) ,these records should be attributed to the
genuine penetration into the Arabian Sea or merely our lack of
knowledge concerning the distribution of pelagic species is
difficult to assess. The theoretical explanation of the
discontinuous distribution is possible though hardly convincing.
404
The periodic and predictable phenomena like production
maxima, production levels, diurnal migration etc. in marine
ecology and the marine ecosystem specifically is related to such
phenomena. This indicates that at different geographical localities
the dominant processes vary, different ecosystem may be present
and congruency in sometimes-disjunct distributions of many
species proves that homologous ecosystems are found in
different oceans (Spoel, 1994). Rios & Carvacho (1982) believe
that disjunct populations that were isolated for short geological
time belong to the same species, as the morphological characters
are not reliable.
The other Arabian Sea genera Psathyrocaris,
Eupasiphea and Pasiphaea are mesopelagic (Foxton, 1970)
undergoing vertical migration. The former genus and one species
of the latter (P. sivado), mainly living near the bottom are found in
the Indian and N. E. Atlantic, this invasion may be due to the
pelagic larval forms assisted by movement of Atlantic water
around and into south west Indian ocean or it may be due to influx
in the north through the Suez Canal.
Family
Oplophoridae:
Genera
Acanthephyra,
Hymenodora and Ephyrina, Systellapsis, Notostomus and
Janicella have their distribution in the Arabian Sea. Acanthephyra
(Scarlet Red shrimp) is represented by at least six species in the
Arabian Sea (Calman, 1939) particularly abundant in the upper
1000m, only one A. eximia is horizontally distributed in Atlantic
and Indian Ocean. It is a vertical migrant. No data is available on
the population in the Arabian sea but in the Mediterranean north
Atlantic and S. Africa other species of Acanthephyra tend to
execute considerable migration and are associated with certain
water masses (Foxton, 1971, Kensley, 1981, Chace, 1986,
Carters, 1993) .The analyses made by these authors may help
yielding information on population of Acanthephyra occurring in
respective regions. According to Chace (1986), the juveniles live
in mid water; adults probably live in or near the bottom of depth of
405
200m, to more than 4700m. Since the physical and biological
factors, acting alone or in combination are correlated to the range
of vertical migration the situation would be different in the Arabian
Sea.
Family Thalassocarididae: The Indo West Pacific small
family is represented in the Arabian Sea so far from coast of North
West India by Thalassocaris crinita (Menon & Williamson, 1971)
and T. obscura from tuna stomachs (George & George, 1964 as
T. lucida). The former is limited in shallow water. The adults and
juveniles of T. crinita were taken from 95m and larvae between
coast and edge of continental shelf. The latter is spread in the
northern Indian Ocean associated with deep water. The larvae
show a much wider distribution than the adults do.
b.Benthic: Normally species having a depth distribution
below 200m are considered benthic. Of the benthic groups three
dominating families are Palaemonidae, Alpheidae and
Hippolytidae; smaller families like Rhynchocinetidae and
Gnathophyllidae are well represented and to a lesser extent
Pasiphaeidae, Thalassocaridae, Processidae, Pandalidae and
others. A very small number of the benthic species have a
circumtropical distribution.
Family Palaemonidae: The subfamily Pontoniinae of
Palaemonidae is purely marine and serves to illustrate in
someway the evolution problem occurring in marine animals. The
present authors have not yet encountered its coral associated
species as corals in Pakistan generally limited to sparse and
stunted growth with no extensive reef formation. The majority of
species free living belongs to the genus Kemponia; the two
Pakistani Kemponia species are included in this group. The genus
has different composition in the western Indian ocean and Arabian
Sea (Calman, 1939, Bruce, 1971, Holthius, 1986), except that of
K. seychellensis which is common on both eastern and western
part of the Arabian Sea in the brackish and marine waters of
406
Karachi and creeks; the second pontonine genus Anchistus
(A. custos) live in association with pinnid shells. A third genus
Periclimenaeus occurs in the Omanis area on the surface of the
sponge (Calman, 1939).
This family is found in the pelagic zone as well as in marine
waters exceeding 100m depth and clearly indicates that these
shrimps are quite well represented in deeper tropical seas. Nearly
60 deep-sea species are known from Indo-west Pacific (Bruce,
1991). None has been reported in our waters.
Palaemon pacificus: In genus Palaemon P. pacificus as the
name indicates is distributed throughout the Pacific and Indian
Ocean. It is a littoral form and the only palaemonine of true marine
water
Family Pandalidae: Genera Heterocarpoides, Heterocarpus,
Dordotes, Procletes, and Pleisonika have been reported from the
Arabian Sea. A single species has been reported from Pakistan in
a world list (Frimbodt & Dore, 1986) and described here. The
family is more diverse in coldwater although numerous pandalid
species occur in tropical region but only in colder waters of deep
shelf and slope depth, the family is mainly benthic at 300 to
1150m, but some have epi-mesopelagic life history, the tropical
deeper genera are dioecious (King & Moffit, 1984). Adaptive or
phylogenetic explanation of this geographical variation in sexual
system is unknown (Bauer, 2004).
Family Crangonidae: Majority of members of this family
are tropical, they are small, in faunal shrimps generally taken from
the deeper waters of the continental shelf, consequently
crangonid population are often inadequately sampled by
conventional collecting gear and apparent scarcity of some
species is probably deceptive. This may also be found at some
exceptional sites influenced by the bottom conditions. The genus
Pontocaris at hand has the largest number of known species;
seem to be concentrated in the Indian Ocean. Another genus at
407
hand Philocheras includes species P. gracilis almost
cosmopolitan, not reported from here with the majority reported
from tropical region.
Family Processidae: The Processidae are pan tropical
and subtropical occasionally temperate, many are confined to
shallow grass flats and tide pools, but others form a component of
off shore fauna. The only genus recorded to date from here
Processa
has its eastern and western Arabian Sea
representative species different i.e. P. compacta (=P. barnardi) on
the eastern side and P. sulcata, P. coutiere, P. australiensis and
P. compacta on the western side on Southern Oman (Hogarth,
1989), P. australiensis is likely to occur here because of its range
extending from Australia to the Arabian Gulf. P. compacta occurs
in the Eastern Atlantic and the Indo-West Pacific that is a most
unusual distribution (Kensley, 1981).
Family Hippolytidae: Hippolytids are more diverse in
cooler water of the northern hemisphere and the tropics are not
quite hippolytid hot spots, the Pakistani genera Hippolyte , Saron,
Heptacarpus, Lysmata, Exhippolysmata and Latreutes
are
certainly important members of tropical shallow waters from littoral
zone through shelf depth. Not all Latreutes species reported from
the Arabian Sea were encountered from Pakistan.
In spite of presence of scattered canopies of
mangrove trees here we were not able to collect any species of
the genus Merguia which is the only Hippolytid genus having semi
terrestrial species crawling and jumping around the exposed roots
and trunks of mangrove trees in Africa (Bruce, 1993). One can
assume here that with the death of Rhizophora plants in our
mangrove forest the caridean M. rhizophorae if present is gone.
More surveys could be rewarding. The genus Saron forms the
natural community in the corals, Kemp (1914) did indicate species
niches from India: as coral reef and deltaic region but all the
specimens included in the present studies were collected from the
408
rock pools. Heptacarpus pendaloides has a vast gap in the
distribution occurring in Pakistan then in China and Japan.
Family Alpheidae: The family is the most common and
most diversified caridean family here. Alpheids are among the
smallest known caridean shrimp therefore may be easily
overlooked or mistaken for juveniles. Some species are new,
records and some fall within their established range, for example
Synalpheus thai records from Pakistan fill the gap in its known
range. At least eight of the Pakistani Alpheus species reach up to
the Australian waters and four Alpheid share with the Red Sea
and Gulfs. Alpheus lobidens has moved in to the Mediterranean.
(Lewinsohn & Galil, 1992); Alpheus lobidens is divided in to two
populations: one Western Pacific and another Indian Ocean
growing large (36mm in TL) and central Pacific which never
attained this size, indicating a slight difference in gene pools of
the two groups (Banner & Banner, 1974).At present the status of
some of the Pakistani species, such as A.lobidens, A.edwardsii,
remain uncertain. Several species found in Gulf of Oman and
South West India are not collected from Pakistan.
The older and the only record of Synalpheus
neptunus from Karachi by Henderson (1893) as Alpheus
neptunus was not relied upon by Banner & Banner (1972, 1975).
It is a coral associated species and in any case Henderson‟s
report was correct with the death of corals at Karachi the species
is likely to become rare.
The genus Athanas is generally believed to live in most
tropical and some temperate seas except off American continents.
The western side of the Arabian Sea including the Arabian Gulf
has different species composition when compared with the
Pakistani side.
The data on the collection of the various alpheid species
does not lend itself to zoogeographical division of the alpheid
fauna within the area studied. Anyhow one thing is true that
409
present collection does not represent the full picture of
alpheids when we compare western side of the Arabian Sea (Gulf
of Aden) where 48 species and from the Persian Gulf 15 species
are recorded (Banner & Banner, 1981). Indo West Pacific species
do not enter the Eastern Pacific is due to barrier of unbroken deep
expanse of water acting effectively to the pan tropical distribution.
Only S. tumidomanus, Alpheus lobidens, A. strenuus strenuus
and A. pacificus have reached the Pacific Ocean. Certainly more
collection is required. As we are loosing our corals and sponges,
consequently associated fauna is lost.
Family Ogyrididae: Ogyrididae are small shallow water
shrimps with tropical warm and temperate marine distribution.
Ogyrides saldanhae was so far supposed to be endemic to west
coast of South Africa (Kensley, 1981) and its presence from
Pakistan shows its eastern and northern extension in to the Indian
Ocean.
Family Gnathophyllidae: Gnathophyllidae is a small
caridean family commonly said to be associated with echinoderms
but well-established echinoderm association is not known for the
genus Gnathophyllum. G. americanum is circum-tropical intertidal,
shallow infratidal in distribution; its worldwide distribution implies
that it may occur in our waters (see Ali, 2006).
Family Rhynchocinetidae: Rhynocinetes uritai’s report
based on a under water photograph by Ali (2006) can be a
misidentification by him, the species is limited to S. Korea and
Southern Japan (Chace, 1997), Ali‟s collection could be the Indian
Ocean species R. (Cinetorhynchus) hendersonii Kemp, 1925, if
the generic identification is believed to be correct.
Family Hymenoceridae: Hymenocera picta is associated
with asteroids and feeds only on starfish. This striking attractive
shrimp is immediately recognizable.
410
3.2.2.3. Discussion& Conclusion:
A detailed analysis of the distribution of neither the
Pakistani marine shallow water nor deep-water caridean shrimp
families is possible at present time. For example, the Alpheidae
several of the known species from unrestricted marine waters of
the Arabian Sea are still known only from the type localities and
others are unavailable nomenclaturally.
The available data is meager but it tends to confirm
the belief that no endemism occurs in marine species, as the
barriers to the distribution of species with pelagic larvae are
virtually non-existent. Knowledge on regional endemism depends
on the state of knowledge of the geographic and depth distribution
of the species in question. Our intertidal species are relatively
known, for infratidal gaps of knowledge exist due to lack of
sampling. However, two faunal provinces (Sindh and Makran)
limited by salinity, topography, temperature with areas of
overlapping are marked. The different composition between the
two provinces indicates a transition reflecting the change in
salinity regime of the water of the areas. The Sindh area has a
diversified and abundant fauna since the habitats are also
diversified.
Our data are certainly insufficient to justify any
conclusion but the scarcity of species common to the western
Atlantic and eastern Pacific tends to support the idea earlier
mentioned. Factor controlling the presence or absence of several
species is the presence or absence of specialized habitats-the
three most notable of these are: coral reef not abundantly found in
our waters and those carideans always associated with corals
generally do not occur here and mangroves that occur in
estuarine, embayment or sheltered areas are noted for their
associate species and mudflats are colonized by plants, sea grass
beds, in the shelter of which genera such as Hippolyte,
Gnathophyllum and Kemponia flourish. We observed that the Indo
411
West Pacific component constituting the major section of this
part of the caridean fauna.
The relative homogeneity is unexpected as the dispersal of
pelagic larvae is expected to be disturbed by a rather complicated
and varying current system in the Indian Ocean..
Considering the worldwide geographical distribution, 56
marine species in present study are distributed as follows.
Table 3: Zoogeographical components of the
Pakistani marine caridean shrimps (including also single records, whether
the fore going figures indicate genuine penetration into neighboring areas is known from the
literature).
Atlantic
7.5%
Indian including Gulfs &
Red Sea
90%
Pacific
50%
Mediterranean
2.5%
Endemic (local endemicity)
0%
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Table 3 indicates the geographical range of carideans
recorded in the of course present study- 50% are widely spread,
in the west Pacific some up to eastern Pacific. Virtually all the
species are also found in the Arabian Sea.
Some species are distributed on both sides of the
subcontinent India, i.e. in the Arabian Sea as well as Bay of
Bengal. We see some geographical isolation and disjunct
population of few species; it could be due to isolation for a
geological time or the radiation of carideans not explosive.
However, the zoogeographical studies of the area cannot be
termed as complete as the poorly sampled areas are more
common than well-sampled areas.
Mirza (1989) has divided the freshwater regime of Pakistan
for the study of freshwater fish as five ichthyogeographic
provinces but no such studies are available for freshwater
shrimps. The same division may be applied here. The
temperature, land barriers, salinity nature of substratum and
latitude seem to determine the dispersal.
Overall, for fresh water species we can say that almost all
of the freshwater species may have localized distribution within
Pakistan for example M. semmelinkii, M. naraensis, M.
malcomsomi, and M. naso confined to a few localities in lower
Indus Basin in Sindh and Balochistan. All the subspecies and new
species show endemicity. The Sindh province is the most
important one as regarding the carideans.
In an attempt to analyze, the Pakistani carideans, which
form a heterogeneous group of animals it, was very difficult to
draw some sensical conclusion on biogeography without any
information of the modes of life histories of majority of the species
at hand.
413
The caridean fauna of Pakistani waters represents a
considerable variety of species that are dominated by the
members of three taxa, the Palaemonidae, Alpheidae, and
Hippolytidae. The Alpheidae and Palaemonidae are among the
most complex and taxonomically difficult groups. In order of
diversity 27 Palaemonids, 23 Alpheids, 7 Hippolytids, 7 Atyids, 6
Pasiphaeids, 2 Crangonids, 2 Ogyrids, 1 Rhynconetid, 1
Hymenocerid, 1 Oplophorid, 1 Gnathophylid, 1 Processid, and 1
Pandalid have been collected until to date.
Since the caridean fauna of Pakistan is still very
much in survey stage it is hoped that in future if systematic
exploratory surveys using high tech gears are made more taxa
are likely to be added. The high figures of species in the overall
families occurring in the Arabian Sea certainly represent a more
extensive effort at collecting.
It can be suggested that human activities and natural
disasters have adversely affected the abundance and distribution
of the species and we still are far from understanding the reasons
for limited distribution of freshwater various species, because lack
of knowledge in this field.
414
Chapter- 4
Species of interest to fisheries and aquaculture
4.1 Water Resources-Position, threats and
Potentials
The Islamic Republic of Pakistan is located north of Tropic
of Cancer extending from 24oto 37o N latitude and from 61o to 75o
longitude. The country covers an area of about 79.6 million
hectares of land, which is of diversified relief, having vast plains in
the middle as Indus Basin, a rock expanses of plateaus in the
southwestern part and a series of high mountains in the north.
From administrative point of view, the country is divided as
Federal Capital Area, Sindh, NWF, Punjab, Balochistan, GilgitBaltistan Provinces, Federally Administrative Tribal Area (FATA),
and Azad Kashmir Area. The water resources (Fig.181) for
fisheries purpose can be classified as fresh water, brackish water
and marine water.
4.1.1: Fresh water:
From hydrological point, the country is divisible into three
hydrological zones, viz, the Indus Basin, the closed Basin of the
arid desert of Kharan, and the Makran coastal Basin (Khurshid,
415
2000). The mighty Indus River with its tributaries, which are
spread in catchment area (561,250 sq km) in Pakistan and are
responsible for most of the country‟s large number of natural
water bodies as a complex system. The principal rivers of the
Indus River system are Kabul, Ravi, Sutlej, Jhelum, Chenab, and
Beas. Major tributaries in Himalayan area are the Rivers Kala
pani, Dras, Shyok, Shigar, Hunza, Gilgit, Astor, Tangir, Darel,
Poonch, Swat, Gomel, Neelum, Kunhar, Ashu, Utrot, Kandia,
Dubouri, Chore, Panjkora, Khiali, Soan, Drabi and others. They
are all glacier and snowfed. Some water is added during the
monsoon months. Surface runoff resulting from rains and
snowmelt is approximately 157MAF. From fisheries point of view
the major constrains, because of the multiple use of water Akhtar
(1995) has highlighted bodies and environmental
mismanagement.
416
Figure 181: Major Water bodies of Pakistan
417
As we know that Pakistan‟s vast
inland water
resources, not only comprise numerous large and small rivers,
reservoirs, but also barrages/head works, canals, lakes, **Dhoors,
**dhands, village ponds, man-made dams, and water logged
areas. Qadri (1960) mentioned of subterranean rivers in Thatta
and Karachi districts.
**The main course and branch of river Indus have moved
over a long period to the area now lying to the west of its present
course. Deeper areas of the old channels have become either
seasonal or perennial natural water bodies; these represent the type
of area locally characterized as *dhands. ** Dhoors, which generally
refer to, roughly circular open water lakes or ponds, seem either to
relate deeper, open water areas within a large Dora system to be
separate areas formed in distinct depression. (Lagoons).
Most of the natural lakes have disappeared over the last
fifty years, several new lakes have been created upstream of
dams, and barrages of the Indus, several micro reservoirs are
observed particularly in Potohar region.
The inland fisheries resources in Sindh comprise nearly
1209 water bodies covering about 8.6 million hectares. There are
some 400 dhands in Sindh province, at least 12 major ones
(Pateji, Cholri, Sanhro, Mehro (Nurri Lagoon) spreading over an
area about 370 sq km. Most of the dhands are mesotrophic –
eutrophic. All the dhands are interconnected through narrow open
interlinking natural channels. The average depth of these dhands
is from 1.5 to 6 ft. After the construction of tidal link canal all, the
associated dhands have become saline with an increase of 40%
salinity due to seawater intrusion from Shah Samandar creek.
418
These dhands and side channels were producing fresh water
shrimps in the late eighties.
The Punjab Fisheries Department provided a list of water
areas to be utilized for fisheries in Punjab during 2007-2008.
Although these demarcations (10500-hac fishpond area) at
present are meant for fresh water fishes but have equal potentials
for fresh water shrimps fisheries and aquaculture. The district wise
details show that the district Gujarat has the maximum potential
having water bodies of river Chenab and Jhelum, several nullahs,
canals, ponds areas, head works, drains, escape channels
whereas the district Attock has the minimum water areas for
fisheries.
In NWFP there are slow flowing rivers, streams, lower
perennial in Kurram River valley and fast flowing upper perennial
rivers and streams for example the River Swat. Riverine marshes
in D.I. Khan water storage reservoirs / dams are important areas
of NWFP. (500 ha pond area. The cold water of Kaghan, Swat,
Diral, Chitral vallies have no potentials for fresh water palaemonid
shrimps fishery.
In Balochistan province, there is only scant inland fisheries
development. A few reservoirs, like Akra Kaur Dam, Sahigee,
Balar, and Shadi Kaur Dam have been built for drinking water and
irrigation. The water level in the reservoirs fluctuates greatly, most
of them going dry in summer. There are a few inconsistent
perennial streams also, fluctuating from ragging in the rainy
season to a very low flow in dry season. The region receives an
average rainfall of 130mm per year, which makes Rivers Dasht,
Hingol, Purali, and Hub, intermittent that flood once in 2-3 years.
Other rivers are endothermic. Most of the water sinks to the gravel
419
of riverbeds. The newly built reservoirs have some scope
otherwise the Balochistan province has limited possibility for
inland fisheries development, mainly due to its climate and paucity
of water resources.
4.1.2. Marine and Brackish.
The coastal area of Pakistan is divisible into Sindh (300 km) and
Mekran region (750 km) for fisheries purpose. Near Thatta, the
Indus begins its deltaic stage and breaks into distributaries that
join the sea at south- east of Karachi. The delta covers an area of
600,000 ha. The Indus delta consists of creeks, inshore waters
with intertidal mud flats, and mangrove swamps. The latter
provide shelter and serve as nursery and breeding ground for
prawns both penaeideans and carideans and indicate their
productivity higher than the open sea, though not critically
understood .In Sindh the important fishing areas are Korangi
Creek area, Garho area, Gharo creek, Keti Bunder area, Karo
Chan area and Shah Bunder area. The estuary of present active
delta is located at Keti Bunder has now been practically
obliterated since no fresh water is allowed to reach it for ten
months of the year. So the estuary is replaced by the sea and
sand dunes are appearing in the Keti Bunder area owing to the
blocking of the Indus flow to sea (Kazmi, 1984) and badly affected
the shrimp‟s resources of the area. Another out come of the
reduction in flow of water from the Indus in the sea was the
discontinuation of red rice cultivation (Ahmed, 1997); this has
almost wiped out the rice shrimps from the area. Along with the
estuaries, the backwaters are very important. They are associated
with the river system and form a large part of the inshore waters.
The term backwater refers to a system of shallow brackish
lagoons and swampy areas. Our brackish waters are referred to
as tropical estuaries.
420
The Mekran coast is characterized by the presence of
number of bays and absence of any major river system. Important
areas are Gwadar, Jiwani coast, Pasni Bay, Miani Hor, Jiwani
brackish Lagoon and Kalmat Khor. The latter is a coastal plain (55
x 19 km) having a tidal lagoon consisting of mud and salt flats and
tidal channels. It receives fresh water from Basol River. The
Hingol River also makes an estuary on the Mekran coast.
In addition to this we have rights to the Exclusive
Economic zone of 240,000 sq km and 200nm and if ratified to
350nm then the Arabian Sea with the additional continental shelf
area of about 50,000 sq km will make a large Marine Ecosystem
(LME32, total 290, 270, sqkm), our maritime zone is over 30% of
the land area where the average biomass is 15 tons/ km 2 and
demersal zone at 0-200m depth (Brandhorst, 1986), with a deep
sea fishing venture initiated.
4.2. Shrimp Resources-position and potentials:
Fishermen from 100 different countries catch more than
346 different commercial species of shrimps, some of the shrimps
form the most valuable food commodity in the world although by
weight only a small fraction of world fisheries but price per unit
weight of the shrimp products is high. Shrimps are regarded as
“fish” by our common people and called as “Jhinga Machhi”
meaning “prawn fish” Out of the 346 commercial species 189
species belong to the Infraorders Caridea, 80 of them are
421
freshwater inhabitants. The marine species include the shallow
marine included in the Kalri category and deep-sea species of
large size and concentration also thought to be large enough to
support commercial fishery. The deep-sea shrimp live at such
depth that they are hard to catch, little is known about them, and
their potentials have yet to be established. The caridean shrimps
beat their penaeoidean counterpart in diversity and occur from the
tropics to Polar Regions but the latter by far outdo the caridean
fisheries making up only about a fifth by weight of the worlds
decapod shrimp catch (Bauer, 2004). Still the characteristics are
good for aquaculture but not for fisheries, compared to penaeids
its fisheries yield is low on global scale (7,705 mt in 2000, FAO,
2002). They are caught as target species, some species though
edible are caught as an admixture in the main catch, or as
Nematopalaemon tenuipes makes 29% in Gujarat, India in by
catch (Zynudheen et al, 2004). Shrimps are fished by many
methods. Large individuals of Macrobrachium are taken by
hooked line or even by hand. Other more productive methods are
traps, dip nets, sieves and hand nets; trawls are swept over large
areas of ocean bottom, accounting for the vast production of
shrimps in world.
In Pakistan, there are some minor fisheries in operation at
least in a world perspective for marine and brackish water
caridean species. Freshwater ranging into brackish water
palaemonids are the most productive in terms of fisheries than
their higher salinity counterpart. No reliable data on the caridean
catch is available although the shrimps land at the Karachi Fish
Harbour. There are early survey reports of marine shrimps like
that of Zupanovic & Mohiuddin (1973) but without mention of any
caridean. The problem is at the species encountered are not
identified, and no reference material is available for study except
for few species housed in Marine Reference Collection &
Resource Center, Karachi University, whatever is identified needs
422
verification, all these may cause confusion and lead to the use
of scientific names that are incorrect by modern standard or apply
to more than one species. All we can say that a subsistence
fishery of Macrobrachium in inland waters is operating.
Information on economic importance being rather scarce and of a
very general nature, relevant fisheries information are very
preliminary (Table 4)and extent of commercial importance in
Pakistan as staple market article is not known
Table 4:Statistical Data(Macrobrachium spp)
Place
Thatta FishMarket
Average
Production
25 kg/day-90 kg/day
50 gm -125
gm/individual
Hyderabad and
Jamshoro Fish
Market
4 kg/day – 7 kg/day
50 gm-125
gm/individual
Duration
December 2005May 2006
December 2005May 2006
Source:Mastoi, 2007, unpublished
4.2.1. Important Species:
According to Qureshi (1956) two species of fresh water shrimps
(M.carcinus = dacqueti & M.lamarrei) are brought to the market
and Qadri (1960) that the lakes, canals and reservoirs near
Ghulam Mohammad Barrage, Kotri Barrage, Manchar and
Chacher lakes were the places for the prawn (M. lamarrei)
fisheries in Sindh; a brackish water species Exopalaemon
423
styliferus could be easily collected from the brackish and fresh
water of Indus estuary, another species with doubtful identification
was extremely abundant in Nara Canal and M.dacqueti was
present from Sukkur to the mouth of Indus estuary.
Holthuis & Rosa Jr. (1965) published a world list of species
of shrimp and prawns of economic value which included eleven
Pakistani species (Caridina nilotica , Macrobrachium equidens,
M. idella, M. malcolmsonii, M. dacqueti, M.rude, M .scabriculum,
M. lamarrei, Palaeomon pacificus, Exopalaemon styliferus and
Nematopalaemon tenuipes) both from FAO water area (ISW)
and land area (421). The medium sized prawns (Table5)such as
M.idella, M. lammarei and Exopalaemon styliferus are found in
brackish water are quite popular locally.
Table 5:Categories of size
Size
Species
Large – 5 to10 whole
number/kg
M.daqcuiti’ M.malcomsonii
Medium - 200 to 300 whole
number/kg
M. idella
Small – 500 and more whole
number/kg
M. lamarrei
Shakoor (1968) collected Exopalaemon styliferus, M.
dacqueti and M. lamarrei from Hyderabad, Kalmat, Khairpur,
Lahore and Rawalpindi, M. dayanus from Sanghar, Lahore,
Sheikhupura and Rawal dam, M. scabriculum from Kalmat and
424
Kortri and M. malcolmsonii from D.I. Khan, Nowshera and
Sajawal and the former from seepage water around Haleji Lake.
Siddiqui et al (1973) working on fish of Kinjhar lake
mention of Palaemonidae and Atyidae in this lake. Siddiqui
(1976) discussing the fishery resources of River Indus in the
province Sindh indicated at least seven shrimp species having
commercial value over the Kotri Barrage, the first anicule of the
River Indus and which provides a source of seedlings (Yaqoob,
1987) during the monsoon.
In another list of world commercial caridean species
(Holthuis, 1980) another 8 species were added to Pakistani
shrimp species are of interest to fisheries making total 19:
Caridina
nilotica , Palaemon
pacificus, Nematopalaemon
tenuipes, Macrobrachium scabriculus, M. rude, M. dacqueti, M.
malcolmsonii, M. lamarrei, M.idella, M. equidens, M. dayanum,
Exopalaemon styliferus, Ogyrides orientalis, Exhippolysmata
ensirostris, Heptacarpus pandaloides, Latreutes
anoplonyx,
Lysmata vittata, Heterocarpus laevigata and Pontocaris pennata.
Ahmed (1985) while discussing the abundance of the
caridean shrimps in the Indus Delta found Exopalaemon styliferus
quite abundant on the west beach of Keti Bunder. According to
him, a 5-minute trawl haul in a 1.7m wide inlet yielded 96
specimens in February while an unspecified Palaemon in May
dominated. Ali & Jafri (1986) while discussing the status of inland
fisheries in Sindh in late eighties mentioned of M. malcolmsomi as
commercial species 15cm long fetching an extraordinary high
price. In 1980 25,943 tones worth 350 million rupees of foreign
exchange were produced (Lone, 1983). The drought condition
during the last two decades has badly tempered the catch of M.
malcolmsonii (Yaqoob, 1980),
425
Dore & Frimbodt (1986) in their list of world's
commercial species included M. rosenbergii , Heterocarpus
laevigata, and Exopalaemon styliferus from Pakistan. Ahmed
(1992) observed M. dacqueti and M. malcolmsonii from Sindh.
Akhter (1995) mentions of an in-stream fisheries of M.
dacqueti in Pakistan while Ayub (2007 pers.comm) does not
recognize any existing fishery in Punjab presently.
According to Jayachandran (2001), the important species
of Pakistan are M. malcolmsonii kotreeanum, M. malcolmsonii
malcolmsonii, M. dacqueti and M. rude.
Studying in 2007 Kazmi & Kazmi focused on the food
value of non-penaied shrimps. They found Macrobrachium
malcomsonii most important species previously reported from
Sindh and now found in river Chenab at Multan and River Ravi.
Yaqoob (1991) highlighted importance of this species, which
grows to 30 cm in length and weighs 250 g. The breeding season
starts from April and continues to October to November and a
production of 599/ha/crop could be achieved. The species
migrates downward in Indus to estuary for breeding. There is an
average annual production of about 100mt mostly exported with
some local consumption, one kg of these prawns costs US $ 5 or
Rs. 150 in the local market (Yaqoob, 1994) .
M. lammarei grows to a maximum length of about 80mm;
these prawns are continuous breeders with two breeding peaksOne from January to April and the other from September to
October. Exopalaemon styliferus and Palaemon semmelinkii
were collected by Kazmi et al (2002) in the deltaic region (Keti
Bunder, Ambro and Dubba creek). P.semmelinkii is very common
during February and March (Kazmi & Sultana, 2008 unpublished
report). In the Gulf of Kutch it is listed among vulnerable species
(Dholakia, 2004)
426
Kemponia seychellensis and K.elegans also found by
the same authors prominent and sometimes outnumbering
penaeid shrimp in the backwater near Karachi. Hippolytid
Latreutes spp. was frequently sampled from Gharo along with
Lysmata vittata and Exhippolysmata ensirostris.
Proximate composition studies on Alpheids (pistol shrimps)
by Kazmi et al (2006) show high caloric value but the commercial
value is quite minor, in spite being a common group at places they
are not consumed only offered for sale to be used as bait. The
nutritional profile of species studied also suggests that they can
be used as low cost feed for successful aquaculture enriched with
brine shrimp nauplii for fish and penaeid shrimps (Kazmi &
Sultana, 2008).
Aquaculture. Although scientists have been suggesting
since start for fresh water prawn fishery and culture (Qadri, 1960,
Shakoor, 1968). In spite of obstacle in the Macrobrachium shrimp
culture-the nonavailablity of seed, scientists are optimistic for the
future of Macrobrachium farming. (Yaqoob, 1986, Kazmi, 2007).
The fishery and aquaculture of none of the species has been
organized and established yet in our country (Ling, 1970). Few
related studies were undertaken in this region. This indigenous
resource utilization was limited to pilot projects
starting as
experimental culture at Jamshoro pond area in 1963-1964, on
food in relation to growth in 1956-58, on bionomics during 1965 69, a third on experimental rearing techniques of fresh water
prawns for commercial utility during 1977-1983 by the Directorate
of Fisheries, Government of Sindh on rearing fresh water
commercial prawns M. malcolmsoni of River Indus and its known
khories.(Ansari & Yaqoob, 1984), a project “Polyculture of
freshwater prawn Macrobrachium malcolmsoni with Indian Major
and Chinese carps at farmer's ponds in Pakistan” was taken up by
PARC, ALP (2006). The HEC Pakistan sponsored project on the
427
exploitation of non-penaeid shrimps (2007-2008)is the most
recent one that included laboratory culture.
After palaemonids caridean family Atyidae has aquaculture
potential, these shrimps are low on the food chain so that the
expensive high protein feeds needed for Macrobrachium and
penaeid shrimps do not seem necessary. These shrimps are good
candidates for intensive culture. The Marine Reference Collection
and Resource Center, University of Karachi successfully reared at
least a dozen caridean species under the laboratory conditions.
Here one thing has to be remembered that, there are
environmental impacts of shrimp culture, such as chemical and
biological pollution of natural waters. Now that the freshwater
hatcheries are on cards, it is suggested here that any expansion
of carideans or other shrimp culture should be closely monitored
and controlled to protect native habitats and species. We must
learn lesson from Singapore where M.rosenbergii is extinct. (Ng,
1997). The worldwide stocks of Macrobrachium have seen rapid
declines in recent years; because of overharvesting, habitat loss
and increased pollution requiring conservation .We have no ban
on wild collection of fresh water prawn during breeding season.
The berried females collected, carrying 30, 000 eggs for prawn
production can adversely effect leading to stock depletion in
natural water of River Indus where dam ill maintained design is
already hampering the migration of these prawns for breeding
purpose.
Both
atyid
and
palamonid
members
are
amphidromphous shrimps, which migrate to upstream by walking
at night. There is a need to modify and improve existing fish way
for preserving shrimp populations. Other threats our freshwater
bodies facing are habitat destruction and water scarcity due to
river diversion, urbanization, industrial pollution, see level rise and
seawater intrusion. In general various agro farming projects,
range management and other quarrying activities in and around
are affecting these water bodies, changing their water quality and
428
water regime, which has direct reflection on fisheries that
ensures food security for masses.
Our experts have been warning of the biological impacts of
fisheries development like introduction of hatcheries and increase
in the disturbance of wildlife, if suitable amelioration measures are
not adopted (Ahmed, 1993). There is a need of accurate maps
delineating the various types of coastal and inland shrimp‟s
resources, geographical distribution, breeding seasons, reserve
areas, commercial fishing areas. It is imperative to develop the
precious potentials of lake shrimp or fish-shrimp combined
fisheries on war footing particularly in the man made lakes as
these are now the biggest sources of water in Pakistan where
prawn fries can be released, the stocks increased and the water
conditions monitored, we have approximately 102, 000 ha of fresh
water reservoirs connected with dams or barrages and about
110,000 ha of natural or man made lakes (Hussain, 2007). It is
true that the stocking rates of prawns would be low but due to the
high value of the product, total revenues could be significantly
increased.
We are among the 113 countries in the world growing rice.
These paddy fields were found also suitable for stocking of young
Macrobrachium prawns of 6-7cm, which are able to grow almost
to maturity by the end of the paddy-growing period (Menon,
1954). The fish and shrimp culture offer opportunities, which can
play for food security and poverty alleviation in rural livelihood, but
need some modification in rice fields necessary for agronomic,
and aquaculture management (Hallwart & Gupta, 2004).
4.2.1. Other uses:
The other uses of caridean species besides for human
consumption, are as aquaculture species, known to be sold for
bait, as bioassay material, and as sub products especially in
429
chitin- a naturally occurring biopolymer which is approximately
75%, the total weight of shrimp to be bioconverted, as ornamental
shrimps popular with aquarists and sold in pet shops, though not
yet successfully cultured worldwide as large scale, nevertheless
the situation may change as research is improving. One report
informs that in marine aquarium trade of 18 species generates
$200-330 million worldwide each year (Wabnitz et al,
2003).Freshwater prawn not only have economic importance in
hydroponics and fisheries for food purpose but haverecently
received attentionin aquarium trade. M. dayanus generally known
as Kaira river prawn/ red claw shrimp is sold as a group of
juveniles for $15 in USA pet shops and Lysmata vittata (
Peppermint shrimp )for $18.
We need to change peoples taste. More distant are not the
days when proper use of these shrimps will be made in Pakistan
in view of present population trends and these shrimp being good
source of first class protein. To achieve this, non-morphological
studies like karyology, hybridization experiment breeding session
and biochemical studies may help. We cannot dismiss the future
possibilities of utilization of the shrimps to minimize the pressure
on our penaeid shrimps, and to strengthen the stagnating capture
prawn fisheries harvest from increase coming from aquaculture of
caridean shrimps-Insha’Allah
430
ADDENDUM
Some species were reserved by the authours to form part
of their large publication but ultimately did not materialize till them
going to press it was thought out most practical to publish them as
addendum in the present monograph so clean our tables from
several of the items that have cluttered them for so long.
Macrobrachium lehiai sp.nov.
(Pl.3
B)
Type material: Holotype ovigerous female CL 20mm.
Paratypes one male, three females (two ovigerous)
Type locality: Leh Nullah 25 January, 1980. Zoological
Survey Department
Description: The rostrum is moderately long extending
beyond tip of antennal scale; upper margin is sinuous armed with
10-12 teeth of which three are post orbital. The dorsal teeth are
arranged in three groups: the proximal groups is of 5-6 equidistant
teeth, this is followed by a second group of 2-4 equidistant teeth
and a distal group of 1-2 teeth, all these groups are separated by
wider gaps; the ventral margin has 6 teeth.
The carapace is smooth longer than rostrum in large
specimens smaller to equal in small individuals, both antennal and
hepatic spines are present, the latter is situated bellow and
behind of level of former.
The telson is slender extending slightly beyond level of
outer lateral spine of the uropodal exopod except in the largest
paratype in which extends visibly beyond level of outer lateral
spine; the dorsal surface is armed with two pairs of spines in distal
431
half (excluding the distal pair). The distal end is very sharply
pointed also has two pairs of spines of which outer pair is smaller
and inner pair is longer but does not reach tip of telson; there are
4-6 ventral plumose setae on each side of the pointed tip and 3-4
dorsal simple setae.
The first pair of chelate legs exceeds the antennal scale by
half chela two distal part of carpus when extending; the carpus is
the longest segment. The second pereopods are long provided
with spinules; the merus is longer than the palm; the carpus is the
longest segment, slender and cylindrical, some times equal to
chela; the movable finger is pubescent; the im movable finger is
also sparsely pubescent; one dentition is present on each of the
cutting adges of chela.
The nonchelate legsare slender. The uropods are
characteristics, by the presence of accessory subapical spine on
outer margin of exopod
Remarks: The new species shows affinity with M.
villosimanus (Tiwari, 1947) and M. rosenbergii in the nature of
telson, second legs being densely beset with spinules, in the
nature of rostrum with M. tiwari and M. naso, the first and second
pereopods resemble M. idella idella
Etymology: The species is named after the type locality
Leh Nullah near Rawal Pindi
Macrobrachium sp
(Fig.182)
Material examined: One female, CL 22mm
Description: The body is robust. The rostrum is long
exceeding the antennal peduncle by one fourth of its length but
432
not exceeding the length of antennal scale. The specimen is
from the group which includes M. idella and M. novaehollandiae
etc. The rostral teeth are present on both the margin, of dorsal
teeth three are postorbital like that of M.idae,and M.indicum etc.
There are 5 ventral teeth like those of M. equidens and M.
rosenbergii . There is one apical spine at distal margin and below
it on distal end, both separated by long setae. This gives the tip a
trifid appearance like M. siwalikensis. Setae are present between
teeth of both dorsal and ventral margin. Adrostral carina is thick
dividing rostrum into dorsal and ventral half. The shape of rostrum
is somewhat styliform or sigmoid as in M. rosenbergii. The
carapace is smooth. The antennal and hepatic spines are present;
the former is slightly below rounded orbital and latter behind and
below level of former. The abdominal pleura first to third are
broadly rounded, of second fully overlapping one and third, of
fourth and fifth narrower and tapering towards apex, which is
directed posterior. Pleuron of sixth somite is very small, ending in
point which overhangs articulation with telson. Relative size of
somites decreases from second to fifth. The telson is long narrows
distally does not end in a sharp median point, since its tip is wornout. Group of stiff setae are arranged on proximal dorsal aspect,
they are curved.
433
Figure 182: Macrobrachium sp. A-Rostrum and carapace, anterior part;
B- Telson and uropod; B’- Telson, tip; C- First leg; D- Third leg
The generic character of two dorsal pairs of telsonic spines
is not seen here, only a single pair is present at mid length of
telson. The apex is flanked by two pairs of spines, inner pair is
thicker and curved not overreaching tip of telson, numerous
plumose setae are present between inner pair of spines.
434
The stylocerite overreaches anterior lobe of first
segment. Fused portion of the two ramii is completely fused as in
M. carcinus. Mandibular palp is 3 segmented. First chelate leg is
fragile and overreaching antennular scale by distal one fourth of
carpus which is the longest segment of the limb. Non chelate legs
are simple. Since second chelate legs are missing, we can not
determine even up to the group of the species so the specimen
which is a female can only be named at present as an abnormal
Macrobrachium since of M.siwalikensis is found in northern areas
of subcontinent India. So there are bleak chances that the present
specimen belongs to the same.
Locality: Nothern area of Pakistan.
Alpheus sp
(Fig. 183 Pl. 6A)
Material: 3 specimens including one ovigerous female
Description: The present specimens could not be named,
however they are included in edwardsii group. Some of the salient
features of the specimens are: small rostrum, carinate posteriorly,
ocular hood inflated. Antennule rather stout, scaphocerite with
lateral margin concave in the middle, distal spine over reaching
distal end of antennular peduncle. Major chela over two time as
long as broad, fingers slightly narrower than palm, immovable
finger compressed laterally, almost right angular on superior distal
margin, tip triangular over reaching tip of immovable finger,
superior transverse groove of palm shallow with proximal shoulder
not over hanging groove, inferior transverse groove broad with
435
proximal shoulder rounded, groove extending upward to whole
of the fix finger. Major chela very elongate. Dactylus of third leg
subspatulate. Telson longer than broad, armed with two pairs of
dorsal spines, posterior margin convex, bearing setae and armed
with a pair of unequal spines laterally. Uropodal exopod bearing
an acute immovable tooth and internally a long curved tooth,
transverse suture straight.
436
C
A
B
D
E
Figure 183: Alpheus sp. A-Frontt; B- Telson and uropod; C- Large
cheliped; D- Dactylus of first leg; E- Same, third leg
437
Locality: Port Qasim
Size: 405 to 13mm in CL
Remarks: this species is related to all the species found in
mangroves of South East Asia i.e . A.paludicola, A.crassimanus,
A.euphrosyne, A.malabaricus and A.rapax.Of these all are
estuarine except A.euphrosyne which is an obligate mangrove
species. (Yeo&Ng,1996).
Alpheid sp.
(Fig.184)
Material Examined:One male -5mm inCL
Locality:Gwadar
Remarks: This specimen looks an unusual
Leptalpheus(Marin,pers.comm.QBK).The three-segmented
carpus of second leg is unusual however, it is being
included here as Leptalpheus sp until we obtain more
material.
438
Figure 184: Alpheid sp. A- Ffront; B- Telson; C- Abdominal lobe; DAntennule; E- Mandible; F- Small cheliped; G- Large cheliped; HSecond leg; I- Dactylus of third leg; J- Appendix masuclina
439
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500
Glossary
abdomen:
posterior-most portion of the body, consisting
of 7 segments, inclusive of the telson.
accessory:
secondary or smaller.
angle: aspect, corner, point or edge.
antenna or second antenna (pl. antennae). appendage
situated between the antennules and the orbits;
antennal flagellum: A long multi-articulate tactile
appendage arising from the peduncle or stalk of antenna.
antennular or first antenna: mobile sensory appendage,
normally biramous and flagellate, arising anteriorly just
below the eye. In some members of Palaemonidae and
Hippolytide the antennules are trimarous.
antennular peduncle: first 3 segments of the antennule
.anterior:
nearer the head end.
apex: tip.
appendix interna: a small tubular structure on the inner
border of the endopods of pleopods 2-5 in most Caridea.
arcuate:
curved or shaped like a bow.
armed:possessing spines or teeth.
arthrobranch: one of the three types found in natant
decapods, arising from arthrodial membrane joining the
limb and body.
arthrodial membranes: membranes between the joints of
the legs, in particular those at the bases of the legs.
article:
general term for segment of appendage.
501
basal: at or near the base; near the most proximal
portion of an appendage.
basis: second most proximal segmtent of a thoracic
appendage.
bearing:
possessing or having attached to it.
benthic:
related to bottom life.
bifurcate:
forked, having 2 prongs attached at the
proximal most part.
bilobed:
with 2 lobes.
biramous:
consisting of 2 parts.
blunt: not sharp or pointed.
brackish:
pertaining to water of salinity intermediate
between fresh water and sea water.
branchia:
a gill or respiratory projection.
branchial region of the carapace. Ventral portion of the
carapace that covers the gills.
carapace:
the covering of the dorsal and often lateral
surfaces of the head and thorax, which is usually
hardened.
carcinology: study of Crustacea.
cardiac region. Median region of the carapace behind the
cervical suture.
carina (pl. carinae). A keel-like ridge.
carpus:
third segment from the distal end of a
thoracic appendage.
caudal:
of or pertaining to the tail.
502
cervical groove. A complex groove or series of grooves
running transversely across the carapace separating the
hepatic and gastric regions from the branchial and cardiac
regions.
cephalon:
head region of body.
cephalothorax: anterior portion of the body formed by the
fusing of the head with one or more thoracic segment.
chela: a claw or pincer used for grasping, formed by the
last 2 segments of a cheliped such that the last segment
(dactylus) opposes the next to last (propodus).
chelate:
possessing a true chela, where the fixed
finger extends distally to oppose the movable finger.
cheliped:
any pereopod that bears a chela.
claw: short, curved process on the appendages.
cleft:
divided or partially divided; split.
commensal: an organism living with another where both
species generally benefit by the association.
compressed: flattened from side to side.
concave:
with lateral edges curving inwards toward the
center of the body.
conical:
shaped like a cone; round at the base,
tapering to a point at the tip.
convergent: turning toward each other; coming together
at one point.
convex:
with lateral edges bulging outward away
from the center of the body.
cornea:
pigmented,
compound eye.
light-receptive
portion
of
a
503
coxa: the most proximal segment of a thoracic leg.
coxal: pertaining to or attached to the coxa.
dactylus (pl. dactyli): the most distal or terminal segment of
a thoracic leg.
deflexed:
bent or curved downwards.
denuded:
with all debris or hairs removed.
depressed:
flattened from top to bottom.
depth: maximum measurement of body or appendage from
the dorsal to the ventral surface or edge.
detritus:
particles of decaying animal and plant
material, which with the associated micro-organisms,
provides food for a variety of animals.
developed:
formed into; present, readily apparent.
diaresis:
transverse suture on exopod of uropod
directed:
turned or bent.
distal: farthest from the center of the body or from the
point of attachment; opposite of proximal.
divergent (diverging): separated or bending away from
each other.
divided:
not entire; with a small notch; formed into
teeth or lobes.
dorsal: upper surface.
ecotone:
oceanic
defined boundaries.
ecosystem
divided
by
clearly
elevated:
surfaces.
raised higher than surrounding structures or
elongate:
lengthened or extended.
504
emarginated: having a notch at the apex or having a
notched margin.
endopod:
inner branch of a biramous appendage that
articulates on the basis.
entire: not toothed, notched, or divided; with a continuous
margin.
epigastric spine: spine on dorsal margin of carapace
behind rostrum
epipod:
a branch of an appendage that articulates on
the coxa or precoxa; a segment or filament attached to the
coxa or precoxa of a pereopod.
erect: upright or perpendicular; raised.
estuary:
that part of a river where there is a mixture of
fresh-water with the sea.
exopod:
an outer branch of a biramous appendage
that articulates on the basis; a small segment or filament
attached to the basis of a pereopod.
expanded:
spread out increased in size, enlarged.
eyestalk:
the movable stalk bearing the pigmented,
light receptive portion (cornea) of the compound eye;
length of the eyestalk includes the cornea (entire eye).
filiform:
thread-like.
finger: propodus or dactylus of a chela.
fixed finger – of a chela – propodus of a chela.
fixed spine: spine that is fused with the integument so
that it is not movable.
505
flagellum (pl. flagella): long, narrow terminal portion of
an appendage (eg., the antenna), consisting of many short
segments.
flank: placed or located to the side of.
foliaceous:
leaf-like.
front of carapace: anterior margin of carapace between the
orbits.
furrow: deep, narrow groove or channel.
gape: wide opening or space left between two cutting
surfaces of a chela when the fingers are closed tightly; not
closing tightly.
genus: a category of the animal kingdom whose members
have certain characters in common, a genus is subdivided
into subgenera and species.
globular or globose: spherical, ball-shaped.
gnathopod: any appendage in the oral region modified to
assist with food handling.
gonopore:
genital opening.
granulate:
granules.
appearing as if composed of or covered with
groove:
any channel, furrow, or depression.
hand or manus: the broad proximal part of the propodus
(second most distal segment) of a chela.
herbivorous: taking plant food.
incisor process: process on the mandible adapted for
cutting.
inflated:
swollen, larger than normal.
506
infra-orbital margin: margin below the orbit.
in situ:
in place.
intertidal:
between tide-marks.
ischium:
fifth segment from the distal end of a thoracic
appendage; usually the first large segment of a maxilliped.
joint: place of union or articulation; separation between 2
segments.
labrum:
the upper or anterior lip.
littoral: living at or near the sea shore between the high and
low water marks.
lobe or lobule: any rounded projection.
longitudinal: running lengthwise, along the long axis.
mandibles: the anterior-most or inner-most of the mouth
appendages.
margin:
edge or border.
maxilla:
the second and third feeding appendage
after the mandible.
maxilliped:
the most anterior
appendages, used for feeding,
of
the
medial:
towards the mid line.
median:
lying or running down the middle.
thoracic
membranous: resembling or consisting of amembrane;
pliable and semitransparent.
merus: fourth article from distal end of leg or maxilliped.
molar process: process on the mandible modified for
grinding food.
507
mottled:
marked with irregular blotches or spots of
different colours or shades.
moult: change of outer skin.
naked: without hairs or setae.
notched:
surface.
with a v-shaped cut on the edge or across a
oblique:
slanting.
ocular: near to or pertaining to the eye.
omnivorous: taking plant and animal food indiscriminately.
orbit: eye socket; cavity in the carapace to contain the
eye and eyestalk.
ovate: egg-shaped.
ovigerous:
carrying the eggs. 2 gill arisi
palm: proximal portion of propodus of chela.
parasite:
an organism which lives on or in other
organisms, called its hosts
pereopod:
a thoracic limb .
pectinate:
comb like.
pediform:
shaped like a foot; slender and pillar-like
(opposite of operculiform).
peduncle:
stalk supporting the antenna or antennule.
pelagic:
inhabiting the open seas.
pereopod:
term applied to the last 5 appendages of the
thorax; the cheliped and the walking legs.
508
petasma:
special extension of the first pleopod in
the males of certain crustacea, used to transfer sperm to
the female.
phyllobranchiate gill: gill consisting of a central axis on
each side of which there is a row of plates set closely
together like the pages of a book.
phylum:
a major grouping is the classification of the
animal kingdom, divided into classes, orders etc.
plankton:
the drifling life of water.
pleopod:
paired swimming appendages found on
some or all of the first 5 abdominal segments.
pleura: (sing. pleuron): lateral plates on the body segments
of Crustacea.
podobranch: any gill attached to the coxa of a pereopod.
post-orbital margin: margin of the carapace immediately
behind the orbit.
process:
any marked prominence or projecting part.
produced:
brought forward; shaped or formed into.
propodus:
pereopod.
the second segment from the distal end of a
protuberance: swelling, bulge, projection.
proximal:
nearest to the center of the body or nearest
to the point of attachment; opposite of distal.
pterygostomian spine: spine at anterolateral angle of
carapace.
punctate:
marked with very small depression.
rami (sing. ramus): branch-like structure.
509
rostrum:
that part of the carapace which projects
forward from between the bases of the eyestalks.
rudimentary: greatly
developed.
reduced
in
size;
rugose:
rough.
serrate:
notched on the edge like a saw.
imperfectly
setae (sing. seta): hair-like projections, usually stiff hairs or
bristles.
setose:
covered with stiff hairs or bristles.
simple leg:
not chelate.
spine: a sharply pointed process.
species:
a category lower than a genus.
spinule:
a very small spine or sharp granule.
spinulose:
covered with spinules.
stalked eyes: eyes resting on a stalk or peduncle.
statocyst:
a globular vesicle which functions
perceiving the position of the body in space.
sternite:
in
ventral plates of the thorax.
sternum (pl. sterna): ventral plates or ventral segmented
wall of the thorax.
stridulate:
to produce a rasping noise by rubbing two
roughened surface together.
stout: strong, sturdy, robust.
subchelate: possessing an imperfect chela in which the
pincer is formed by the dactylus folding back against a
broadened propodus, rather than opposing a distally
produced fixed finger.
510
supra-orbital spine: a spine on the anterior border of the
carapace above the orbit.
suture: a non-flexible or slightly flexible joint or seam.
tail fan:
terminal portion of the abdomen composed
of uropods and telson, forming a swimming structure.
tapering:
gradually getting smaller toward one end.
taxonomic:
pertaining to the classification of organisms.
teeth: hardened, projecting processes, usually somewhat
flattened and stouter than spines.
telson: terminal segment of the abdomen.
tergum (pl. terga): dorsal surface of body segments; dorsal
plate of abdominal segments.
terminal:
situated at or pertaining to the end or tip.
thelycum:
specialized receptacle for sperm located
between the coxae of pereopods IV and V in females of
some Crustacea.
third maxilliped: appendages preceding first leg.
thorax: body region between the head and the abdomen.
thumb: projection of the chela or hand of a decapod which
is opposed to the finger or seventh segment.
toothed:
bearing teeth.
transverse:
lying across; from side to side.
trilobed:
with 3 lobes.
truncate:
cut off abruptly.
trunk: body exclusive of head and extremeties.
tubercle:
a small conical prominence.
511
tubicolous:
living in a tube.
tuft: a cluster or bunch of hairs or setae located closely
together or attached at the same point.
unarmed:
lacking spines or teeth.
united: connected with or fused to.
uropod:
a pair of biramous abdominal appendages
that form part of the tail fan.
ventral:
near the lower surface of the organism or
appendage.
vestigial:
very small imperfectly developed; present
only as a very small spine or lobe.
well-developed: very prominent, readily apparent; fully
formed.
wrist: fifth segment of a cheliped of a shrimp which is
subdivided into joint lets.
512
Taxonomic Index
Acanthephyra armata, 41
Acanthephyra eximia, 16, 41,
42, 319, 329
Aegeon obsoletum, 259
Aegeon pennata, 259
Agostocaris, 3
Alberta banneri, 19, 158
Allobrachium, 79
Alpheus alberti, 19
Alpheus audouini, 170
Alpheus barbatus, 164
Alpheus bisincisus, 19, 167,
335
Alpheus brevirostris, 186
Alpheus cf barbatus, 19, 161
Alpheus chiragricus, 19, 161,
169
Alpheus crassimanus, 174,
333
Alpheus edwardsii, 19, 161,
170, 264
Alpheus gracilidigitus, 181
Alpheus inopinatus, 174, 301,
332
Alpheus isodactylus, 19, 161,
173
Alpheus lobidens, 19, 161, 174,
265, 276, 277
Alpheus malabaricus, 186
Alpheus manorensis, 19, 161,
180
Alpheus neptunus, 207, 208,
276
Alpheus pacificus, 19, 161,
181, 335
Alpheus pomatoceros, 188
Alpheus pseudoedwardsii, 19,
161, 185
Alpheus sivado, 39
Alpheus splendidus, 19, 161,
187, 265
Alpheus strenuus strenuus,
19, 161, 190
Alpheus strenuus var.
angulatus, 190
Alpheus zulfaquiri, 19, 161,
192, 319
Anchistia elegans, 134
Anchistia grandis, 134
Anchistioides, 129
Anchistus custos, 18, 131, 265,
318, 333
Anchistus inermis, 131
Athanas, 157, 197
Athanas arabicus, 19, 195
Athanas dimorphus, 19, 197
513
Automate dolichognatha, 19,
201, 265
Automate Gardiner, 201
Automate haightae, 201
Automate johnsoni, 201
Cancer (Astacus )carcinus, 82
Caridella sp, 17
Caridina, 231, 298
Caridina typus, 46
Caridina babaulti babaulti, 16,
46
Caridina nilotica, 10, 16, 48,
265, 286, 287, 312, 317, 330
Caridina nilotica var
brevidactyla, 48
Caridina sumatrensis, 50
Caridina weberi, 16, 49, 50, 51,
266, 320
Caridina weberi var.
sumatrensis, 49
Crangon pacifica, 181
Cuapetes elegans, 134, 135,
136, 137, 138
Cuapetes longirostris, 139
Cuapetes seychellensis, 141,
142, 143
Dordotes, 275
Ephyrina, 274
Eupasiphae, 26, 27, 265, 308,
332
Eupasiphae gilesi, 16
Exhippolysmata ensirostris
ensirostris, 19, 224, 225
Exopalaemon styliferus,
17, 71, 72, 264, 271, 286, 287,
288, 297, 330
Gnathophyllum americanum,
18
Harpilius Miersi, 131
Heptacarpus pandaloides, 19,
229, 264, 287
Heterocarpoides, 275
Heterocarpus ensifer, 253
Heterocarpus laevigatus, 20,
254
Hippolite Krausii, 246
Hippolysmata (Hippolysmata)
vittata, 241
Hippolyte, 223, 228, 229, 231,
246, 278
Hippolyte ventricosa, 20, 230,
231, 265, 333
Hymenocera picta, 12, 18, 266,
277, 330
Hymenodora, 274
Kalriana anissi, 17, 52
Kalriana jhimpirensis, 17
Kalriana karachi, 17
Kalriana sunahrensis, 17, 69
Kemponia elegans, 18, 135,
141
Kemponia longirostris, 18, 139
Kemponia seychellensis, 18,
140, 141, 265
Latreutes, 265
514
Latreutes anoplonyx, 20, 234,
287, 335
Latreutes mucronatus, 20, 238,
312
Leander celebensis, 73, 74
Leander fluminicola, 76
Leander longirostris, 71, 264
Leander modestus, 125
Leander pacificus, 123
Leander potamiscus, 76
Leander semmelinkii, 125
Leander sewelli, 128
Leander styliferus, 72, 322
Leander tenuipes, 120
Leandrites celebensis, 17, 74,
266, 321, 328
Leptocarpus potamiscus, 17,
76, 266, 271, 320, 328
Leptochela cf sydniensis, 16,
36
Leptochela hainanensis, 37
Leptochela irrobusta, 16, 31
Leptochela nasimae, 16, 33
Leptochela oculeocaudata, 37
Leptochela pugnax, 16, 35
Leptochela robusta, 31, 35
Lysmata amboiensis, 266
Lysmata vittata, 20, 264, 287,
288, 290
M. rosenbergii, 270
Macrobrachium, 264, 268, 270,
289, 322, 328, 334
Macrobrachium altifrons
ranjhai, 17, 81, 270
Macrobrachium dacqueti, 17,
82, 83, 264, 270
Macrobrachium dayanum, 17,
85, 270
Macrobrachium equidens, 17,
86, 111, 266, 270, 286, 320,
327
Macrobrachium idella idella,
17, 88
Macrobrachium lamarrei, 10,
17, 89, 91, 264, 271, 331, 335
Macrobrachium lamarrei
korangii, 90
Macrobrachium lehiai, 17, 291
Macrobrachium ltifrons
ranjhai, 17
Macrobrachium malcolmsonii
malcolmsonii, 17, 108
Macrobrachium malcomsonii
kotreeanum, 17, 100
Macrobrachium naraensis, 17,
269
Macrobrachium naso, 17, 109,
265, 271
Macrobrachium rosenbergii,
12, 83, 309, 314, 316, 318,
324, 334
Macrobrachium rude, 18, 111
Macrobrachium scabriculum,
18, 113, 265, 270
515
Macrobrachium shahpuri, 18,
114, 116, 271
Macrobrachium sintangense,
85
Macrobrachium sp, 18, 116,
292, 327
Macrobrachium taunsii, 18,
116
Macrobrachium tirmiziae, 18,
119
Macrobrachium yui, 119
Nauticaris unirecedens, 241
Nematopalaemon, 271
Nematopalaemon tenuipes,
18, 120, 266, 271, 285, 286,
287, 323, 333
Ogyrides orientalis, 18, 152,
265, 287
Ogyrides saldanhae, 18, 155,
277
Ogyrides sibogae, 152
Ogyris orientalis, 151, 152
Palaemon, 287, 307, 325, 334
Palaemon ( Eupalaemon)
sundaicus, 111
Palaemon (Eupalaemon)
carcinus, 82
Palaemon (Exopalaemon)
styliferus, 72
Palaemon (Nematopalaemon)
tenuipes, 120
Palaemon (Palaeander)
semmelinkii, 125
Palaemon carcinus, 82
Palaemon equidens, 86
Palaemon lamarrei, 91
Palaemon longirostris, 71
Palaemon luzonensis, 120
Palaemon mossambicus, 111
Palaemon multidens, 88
Palaemon pacificus, 18, 123,
264, 275, 287, 333
Palaemon semmelinkii, 18,
125, 288
Palaemon sewelli, 18, 128,
266, 271, 312
Palaemon styliferus, 71
Palaemonella longirostris, 139
Palaeomn marmoratus, 246
Pandalus, 9
Parabopyrella indica, 13
Parapasiphaea
(Eupasiphaea) gilesii, 27
Parapasipheae alcocki, 39
Pasiphaea alcocki, 16, 39, 264,
314
Pasiphaea sivado, 16, 39, 264
Pelias nilotica, 48
Periclimenes. digitalis, 13
Periclimenes (Ancylocaris)
elegans, 135
Periclimenes (Ancylocaris)
proximus, 139
Periclimenes (Falciger) affinis,
139
516
Periclimenes (Falciger)
dubius, 134
Periclimenes (Harpilius
elegans, 135
Periclimenes (Harpilius)
longirostris, 139
Periclimenes elegans, 134,
135
Periclimenes longirostris, 139
Periclimenes proximus, 139
Periclimenes seychellensis,
140
Philocheras parvirostris, 20,
257, 265, 317
Pleisonika, 275
Pontocaris pennata, 20, 258,
259, 287
Pontocaris propensalata, 258
Pontophilus parvirostris, 257
Probopyrus pica, 13
Procaris, 3
Processa, 145
Processa compacta, 18, 146,
265
Procletes, 275
Psathyrocaris, 274
R. (Cinetorhynchus
hendersonii, 277
Rhizophora, 276
Rhynchocinetes uritai, 17
Salmoneus, 157, 203, 299, 305
Salmoneus brevirostris, 19,
203, 204, 265, 319
Salmoneus cristatus, 19, 205
Saron gibberosus, 246
Saron marmoratus, 20, 246,
264, 302, 312, 320, 332
Synalpheus, 157, 210
Synalpheus neptunus, 19, 207,
210, 264, 276
Synalpheus thai, 19, 208, 266,
276, 321
Synalpheus tumidomanus, 19,
207, 210, 214, 265, 312
Tenualosa illisha, 267
Virbius australiensis, 231