CSIRO PUBLISHING
Australian Systematic Botany, 2014, 27, 23–37
http://dx.doi.org/10.1071/SB13016
A revision of Chara sect. Protochara, comb. et stat. nov.
(Characeae: Charophyceae)
Michelle T. Casanova A,B,D and Kenneth G. Karol C
A
Royal Botanic Gardens, Melbourne, Birdwood Avenue, South Yarra, Vic. 3141, Australia.
Centre for Environmental Management, University of Ballarat, Mt Helen, Vic. 3350, Australia.
C
New York Botanical Garden, 2900 Southern Boulevard, Bronx, NY 10458-5126, USA.
D
Corresponding author. Present address: 273 Casanova Road, Westmere, Vic. 3351, Australia.
Email: amcnova@netconnect.com.au
B
Abstract. A revision of a group of ecorticate species of Chara is presented, on the basis of fresh, pressed and spiritpreserved material. The following seven species are recognised, characterised by a very simple morphology, with few or
inconspicuous accessory cells (cortication, stipulodes, bract cells, bracteoles) and large gametangia: Chara australis R.Br.,
C. lucida (A.Braun) Casanova & Karol comb et. stat. nov., C. porteri Casanova, sp. nov., C. protocharoides Casanova &
Karol, nom. nov. (=Protochara australis Womersley & Ophel) and C. stuartiana (Kütz.) Casanova & Karol comb. et. stat.
nov. from Australia, and C. corallina Klein ex Willd. and C. wallichii A.Braun from Asia. A new section, Chara subg.
Charopsis sect. Protochara (Womersley & Ophel) Casanova & Karol, comb. et stat. nov., is erected to accommodate these
taxa, formerly placed in sect. Charopsis.
Additional keywords: Chara australis, Chara stuartiana, Chara lucida, Chara protocharoides, Chara porteri,
Protochara, systematics, Tolypellopsis.
Received 2 April 2013, accepted 15 April 2014, published online 30 June 2014
Introduction
Charophytes (members of the extant family Characeae,
Charophyceae) are submerged water plants that grow in fresh,
brackish and saline water throughout the world. They are
distinguished from other aquatic macrophytes on the basis of
their overall simple morphology (consisting of essentially
uniseriate, branched filaments of cells), their unique
reproductive organs (antheridia producing motile sperm cells,
and oogonia or oosporangia containing a non-motile egg cell),
and distinctive oospores patterned with helical sutures. The genus
Chara L. is characterised by a five-celled coronula on the
oogonium (oosporangium), placement of the oogonium above
the antheridium where they occur together, whorls of accessory
cells below the branchlet whorls (stipulodes) and essentially
monopodial branchlet structure. The family Characeae was
revised for Australia by Wood (1971); however, Wood’s
revision resulted in an unusable taxonomy (Casanova 2005,
2009) and recent studies have shown that the lumping of
Australian species with those in the northern hemisphere, and
the delineation of numerous subtaxa (varieties, forms) do not
represent reality, or enhance utility. There are many more species
in Australia than were recognised by Wood (1971) (Casanova
2005, 2009, 2013; García and Chivas 2006) and current
advances in charophyte taxonomy, including investigation of
Journal compilation CSIRO 2014
oospore morphology with scanning electron microscopy, have
allowed a new approach to the systematics of this group.
Historical review
Traditionally (in what has been termed the Braun–Wood
taxonomy, Proctor 1980), the genus Chara has been divided
into two groups, defined on the basis of the number of stipulode
whorls. These groups have been variously treated as subgenera or
sections by different authors (see Kützing 1843; Nordstedt 1883;
Zaneveld 1940; Wood 1962; Wood and Imahori 1965; Proctor
1980). Current taxonomy recognises two subgenera, namely,
subg. Chara and subg. Charopsis (Kütz.) Leonh. Within subg.
Charopsis, the following two sections are currently recognised,
following Wood and Imahori (1965): sect. Charopsis (totally
ecorticate) and sect. Agardhia R.D.Wood (corticated axes,
ecorticate branchlets), with Chara braunii C.C.Gmel. being
the type of subg. and sect. Charopsis. However, phylogenetic
analysis places C. braunii close to C. muelleri A.Braun, which is
in sect. Agardhia (Meiers et al. 1999; McCourt et al. 2000). The
remaining totally ecorticate Chara species are morphologically
and phylogenetically (McCourt et al. 2000) more similar to each
other than to C. braunii; however, a group comprising these
ecorticate species, without C. braunii, has no name under the rules
of the ICN (McNeill et al. 2012). The members of this group have
www.publish.csiro.au/journals/asb
24
Australian Systematic Botany
ecorticate axes and branchlets, and have a relatively simple
morphology (compared with corticated species in subgenus
Chara). Some of these species have, in the past, been
allocated to different genera (e.g. Charopsis Kütz., Nitella
C.Agardh, Nitellopsis Hy, Tolypellopsis (Leonh.) Mig. and
Protochara Womersley & Ophel). The first member of the
group to be described was Chara corallina Klein ex Willd., a
monoecious species first collected in India (Willdenow 1805).
Soon after, a dioecious species was described from Australia,
C. australis R.Br. (Brown 1810). Kützing (1857) placed Brown’s
C. australis in the genus Nitella as N. australis (R.Br.) Kütz.,
described N. stuartiana Kütz. on the basis of more robust
material collected from Tasmania, and illustrated both species.
In a summary of all the taxa in the subgenus, presented by
Nordstedt (1883), Brown’s C. australis was retained without
further description, and N. stuartiana was placed in synonymy
of that species. From a relatively small number of specimens,
an additional six taxa at different ranks were also recognised,
including C. australis var. lucida A.Braun, C. australis var.
nobilis A.Braun, C. australis subsp. plebeja (R.Br. ex
A.Braun) A.Braun, C. australis var. vieillardii A.Braun,
C. corallina and C. wallichii A.Braun. The other ecorticate
Chara species included in this group by Nordstedt (1883),
C. braunii, will not be dealt with in the present study.
Zaneveld (1940) treated C. australis and C. australis var.
nobilis as synonymous, but distinguished northern Australian and
New Caledonian specimens as different forms of C. australis var.
lucida: i.e. f. typica Zaneveld, nom. inval., f. tenerior Zaneveld
and f. vieillardii (A.Braun) Zaneveld. Womersley and Ophel
(1947) erected the genus Protochara to accommodate a totally
ecorticate taxon from Western Australia, Protochara australis
Womersley & Ophel, and transferred Nitellopsis inflata Fil. &
G.O.Allen ex Fil. to that genus as Protochara inflata (Fil. &
G.O.Allen ex Fil.) Womersley & Ophel. Filarszky (1934) also
described a species with morphological similarities to
C. australis, from Indonesia, as Tolypellopsis simplicissima Fil.
In the first experimental breeding study in the Characeae,
Macdonald and Hotchkiss (1956) found that female Chara
australis and a specimen from New South Wales identified by
H. B. S. Womersley as male Protochara australis were interfertile to the point of oospore production, and so amalgamated
C. australis and P. australis, separating material that had
been called P. australis as a new subspecies, C. australis
subsp. estipulodica M.B.Macdon. & Hotchk. In amalgamating
Protochara australis with Chara, they also included P. inflata in
Chara, as C. inflata (Fil. et G.O.Allen ex Fil.) M.B.Macdon. &
Hotchk. Subsequently, on the basis of reproductive morphology
and nucleotide sequences, García and Karol (2004) recognised
Protochara inflata as a species of Lamprothamnium J.Groves,
L. inflatum (Fil. & G.O.Allen ex Fil.) Adr.García & Karol. Wood
(1962) amalgamated most of the members of subg. Charopsis into
a single species, C. corallina, and delineated several infraspecific
taxa, some of which were not validly published (cf. Art. 41.5 of
the ICN; McNeill et al. 2012). These were subsequently treated as
both forms and varieties of C. corallina, as well as being treated
at ‘microspecies’ rank (equivalent in rank to Wood’s forms as
well as his ‘monotypic’ varieties and species, i.e. those lacking
subordinate taxa) by Wood and Imahori (1965). Infraspecific taxa
are treated in the main body of the monograph, with alternative
M. T. Casanova and K. G. Karol
‘microspecies’ status presented in the Microspecies Appendix
at the end of the work. These names are here treated as
invalid, alternative names under Art. 36.2 of the ICN (McNeill
et al. 2012). Wood (1971) later revised his taxonomy for
Australian material, delineating C. corallina var. corallina as
those specimens with cylindrical branchlet segments, and
C. corallina var. nobilis for those specimens with inflated
branchlet segments. Within C. corallina var. nobilis, he
tentatively recognised the following four forms: nobilis,
inflata, simplicissima and stuartiana. These formae names are
here treated as invalid provisional names under Art. 36.1 of the
ICN (McNeill et al. 2012).
In the present study, the type material of all the Australian
species and their close relatives, as well as more than 200
specimens from field collections and herbaria, were examined
to determine the status and morphological limits of the species
previously included in Chara subg. Charopsis sect. Charopsis
(excluding C. braunii).
Morphology
Chara species grow as totally submerged water plants, rooted in
the sediment by colourless rhizoids, with a photosynthetic axis in
the water column. The axis consists of single-celled internodes
and whorls of multicellular branchlets, on which the reproductive
structures are borne. The axes look very similar to submerged
flowering plants, and can be confused with species of
Myriophyllum L. Species of Chara in this group are entirely
ecorticate, i.e. with no cortex on the axis or branchlets, and, in
general, have little in the way of ‘accessory cells’ (i.e. stipulodes,
bract cells or bracteoles). Thus, there are few morphological
characteristics on which to separate species, which has led to
the current taxonomic confusion. They are, in general, somewhat
large, with robust axes and spreading whorls of branchlets. The
antheridia and oogonia are typical of the genus, but large (up to
1 mm long) in comparison with some other species, and visible to
the naked eye. The antheridia are usually bright orange, and
the oogonia have few circumvolutions of the enveloping spiral
cells and very small, apiculate coronula cells. Their persistent
reproductive units (oospores) are among the largest recorded
for Australian charophytes, but with relatively few striae
(spiral sutures), and some species accumulate starch in the
lower and rhizoidal nodal cells (bulbils), which allows them to
persist vegetatively. The major characteristics used to separate
species in this group are sexuality (monoecious v. dioecious),
development and abundance of accessory cells, axis and
branchlet diameter and length, characteristics of the branchlet
nodes and oospore morphology. They are distinguished
from similar species of Lamprothamnium by the presence, in
Lamprothamnium, of gyrogonites (calcified covering on the
oospore), rounded coronula cells, downward pointing
stipulodes and verticillate bract cells.
Materials and methods
Approximately 200 fresh, pressed and spirit-preserved specimens
of charophytes in Australian and overseas herbaria and several
private collections were examined for the present study, including
15 specimens designated as type material of species, subspecies,
varieties or forms within the group. Each specimen was allocated
Chara section Protochara in Australia
a letter–number combination (p###, r###, t### or v###), so that
all data obtained and stored for that specimen (i.e. measurements,
photographs, chromosome counts, scanning electron
micrographs) could be related back to the specimen. Initial
examination involved measurements (size, number) directly
from the specimen (e.g. number of branchlets, number of
branchlet segments) and with the use of a microscope (e.g.
axis and branchlet diameter, number and morphology of
stipulodes and bract cells, arrangement of gametangia).
Oospore features were examined and measured with the aid of
light microscopy and scanning electron microscopy.
Fresh specimens were obtained in field surveys in all states of
Australia (following the methods of Casanova 2004), and from
culture of seed-bank material in a greenhouse (following the
methods of Casanova 2004). In the greenhouse, plastic containers
(185 mm 125 mm 50 mm) containing ~300 g of wetland
soil were inundated to a depth of 10–14 cm in large tanks and the
charophytes that germinated from them were examined. When
oospores were present they were removed from the specimens for
examination with a scanning electron microscope (SEM), then
plants were gathered and pressed, or preserved in 70% alcohol.
If oospores were not present, the plants were either returned to
culture to mature, or kept in jars on a windowsill until they
matured and oospores were freely released. If antheridia were
present, chromosome counts were attempted, following the
methods outlined in Casanova (1997).
Where possible and appropriate, oospores were removed from
the herbarium specimens or obtained as above from live material.
Approximately a quarter (50) of the specimens examined had
oospores. They were prepared for SEM examination by cleaning
(if required) with a detergent solution, using a modification
of the methods of Crawford et al. (2001). Sometimes the
enveloping cells were removed by hand, using fine needles.
For type specimens, very old material, or specimens with a
single oospore available for examination, the oospores were
handled with great care, with minimal manipulation. Oospores
were removed from the stubs after microscopy and are stored
in alcohol and deposited with the specimen for possible future
examination.
Australian Systematic Botany
25
Chara sect. Monosiphoniae Wallr., Fl. crypt. Germ.2 107 (1833),
p.p.; Chara hypogynae a. Monosiphoniae A.Braun, Ann. Sci. Nat.
ser. 2, 1: 353 (1834) p.p. (excluding C. braunii); Chara sect.
Charopsis (Kütz.) R.D.Wood, Taxon 11: 12 (1962), p.p.
(excluding C. braunii).
Type: Chara protocharoides Casanova & Karol.
Axes ecorticate. Stipulodes in a single tier when present,
sometimes obscure, up to 2 per branchlet. Branchlets
ecorticate, with a single-celled end segment, sometimes
surrounded by smaller bract cells. Bract cells 2 or 3, often
obscure. Bracteoles 2, often replaced by gametangia or
obscure. Gametangia sessile; solitary, geminate or aggregate,
on the branchlet nodes and at the base of the branchlets, conjoined
(monoecious) or on separate plants (dioecious). Oogonia
with few circumvolutions, coronula cells slightly apiculate.
Oospores with fewer than 8 striae, without a gyrogonite.
Annual or perennial.
Species in this section have been variously placed in Chara,
Nitella, Nitellopsis, Protochara and Tolypellopsis by previous
authors. All these names refer to currently recognised genera,
except Protochara and Tolypellopsis (=Nitellopsis). Thus, the
name Protochara appears to be an appropriate name for the new
section.
Key to the species in Chara subgen. Charopsis
sect. Protochara
1. Plants monoecious................................................................. C. corallina
Plants dioecious.......................................................................................2
2. Bract cells and stipulodes elongate and inflated .................... C. wallichii
Bract cells and stipulodes small or absent ..............................................3
3. Axis diameter <1 mm, branchlets thin....................................................4
Axis diameter usually >1.1 mm, branchlets somewhat inflated,
6–13 mm long....................................................................................5
4. Branchlets <20 mm long, branchlets incurved ......................... C. porteri
Branchlets 20–50 mm long, axes tangled and wiry ..................C. lucida
5. Stipulodes, bract cells and bracteoles clearly present .............................6
Stipulodes, bract cells or bracteoles absent..................C. protocharoides
6. Branchlets and internodes 2 mm in diameter .................. C. stuartiana
Branchlets and internodes 1.2–1.8 mm in diameter .............. C. australis
Chara australis R.Br., Prodr. 1: 346 (1810)
Results
(Fig. 1)
Chara L., Sp. Pl. 2: 1156 (1753)
Type: Chara vulgaris L.
Chara subgen. Charopsis (Kütz.) Leonh., Lotos 13: 73 (1863)
Type: Chara braunii C.C.Gmel.
Monoecious or dioecious. Plant axis corticated or ecorticate,
stipulodes in a single whorl (haplostephanous), bract cells
unilateral or verticillate, branchlets ecorticate and terminated
by a single cell, a cluster of bract cells or a group of three
equal-sized cells (corona).
Chara subgen. Charopsis sect. Protochara (Womersley &
Ophel) Casanova & Karol, comb. et stat. nov.
Protochara Womersley & Ophel, Trans. Roy. Soc. S. Australia 71:
311 (1947).
Nitella australis (R.Br.) Kütz. Tab. phycol. 7: 11 (1857)
Type: NEW SOUTH WALES: Port Jackson, 1804, R.Brown
277. Holo: BM; iso: LD.
Chara australis var. nobilis A.Braun in C.F.O.Nordstedt, Abh.
Königl. Akad. Wiss. Berlin 1882: 105 (1883); Chara corallina
var. nobilis R.D.Wood, nom. inval., Taxon 11: 12 (1962); Chara
corallina f. nobilis R.D.Wood, nom. inval., Taxon 11: 12 (1962);
Chara corallina var. nobilis (A.Braun) R.D.Wood, Rev.
Characeae 1: 274 (1965); Chara corallina f. nobilis (A.Braun)
R.D.Wood, Rev. Characeae 1: 274 (1965).
Type: NEW ZEALAND: Whangape [Whangapa?] Lake, T.K
[irk] s.n., v.-1870, Lecto: NY (fide Wood and Imahori (1965:
275)); ?isolecto: WELT A9062 n.v., CHR n.v.
Dioecious. Plants robust, translucent and turgid, up to 40 cm
high. Axes 0.9–1.75 mm wide, totally ecorticate, internodes
30–100 mm long. No spine cells. Stipulodes in a single row
26
Australian Systematic Botany
M. T. Casanova and K. G. Karol
D
A
E
B
G
F
C
I
H
K
J
L
Fig. 1. Chara australis R.Br. A. Habit of whole plant; scale bar: 1 cm. B. Fertile antheridial whorl; scale bar: 10 mm. C. Fertile oogonial
whorl; scale bar: 10 mm. D. Base of fertile oogonial whorl; scale bar: 1 mm. E. Branchlet node; scale bar: 0.5 mm. F. Base of sterile whorl;
scale bar: 0.5 mm. G. Geminate oogonia; scale bar: 0.5 mm. H. Base of sterile whorl; scale bar: 1 mm. I. Fertile oogonial node; scale bar:
0.5 mm. J. Oospore; scale bar: 200 mm. K. Detail of smooth oospore wall; scale bar: 2 mm. L. Chromosomes n = 14, 1000 magnification.
A–C and J from M.T.Casanova r186; D, E from M.T.Casanova r612; F, G from D.McNeil r692; H, I from M.T.Casanova r952 and K, L from
M.T.Casanova p788.
when present, 6–12 in a whorl, up to 0.38 mm long, but often
absent or obscure. Branchlets almost uniformly 6 in a whorl,
rarely 7, completely ecorticate, 3–5 cells long, cylindrical,
15–43 mm long, basal branchlet cell elongate, up to 15 mm
long, branchlet end segments usually a single mucronate cell,
sometimes with up to two subtending bract cells, bract cells 2–4
at a node, up to 0.2 mm long, frequently obscure or absent.
Branchlets are often strongly incurved at the apices, especially
in shallow water. Bracteoles obscure, possibly 2 per oogonium.
Gametangia solitary, geminate or aggregate at the lowest 1 or
2 branchlet nodes and inside the base of the whorl, rarely
antheridia sessile outside the base of the whorl. Oogonia
orange, up to 1 mm long, 0.8 mm wide, coronula cells obtuse
to slightly apiculate and usually very small on mature oogonia.
Oospores black, cylindrical to ovate, 700–800 mm long,
450–600 mm wide, with 5 or 6 striae, appearing smooth or
finely granulate, impression of the basal cell 180–200 mm in
diameter. Antheridia up to 800 mm in diameter, orange and often
very obvious. Vegetative reproduction by perennial basal
internodes and the deposition of starch in the lower and
rhizoid nodes. Chromosomes n = 14 (M.T.Casanova p426,
p788).
Recognition
Chara australis is the most common totally ecorticate species of
Chara in eastern mainland Australia. It can be distinguished as
dioecious, robust, ecorticate plants with large gametangia and
small, often inconspicuous, stipulodes, bract cells and bracteoles.
The internodes are usually as long as or longer than the branchlets
and the axes are usually greater than 1 mm in diameter.
Distribution
Occurring in cool, deep, still or slow-flowing water from east
of Brisbane in Queensland, south through New South Wales
(including the Australian Capital Territory) to Victoria, with an
Chara section Protochara in Australia
isolated collection from the Flinders Ranges in South Australia.
Also occurs in New Zealand.
Etymology
From the Latin ‘australis’, referring to the southern distribution
of the species.
Specimens examined
SOUTH AUSTRALIA: Mawson Plateau, Flinders Ranges, 20 May
2010, D.McNeil r692 (MEL). NEW SOUTH WALES: Dumaresq
Creek above Dumaresq Dam near Armidale, 13 May 1990,
M.T.Casanova p306 (MEL), p463 (NE), r396 (MEL); Dumaresq
Creek below the dam near Armidale, 13 May 1990, M.T.Casanova
p307 (MEL); Llangothlin Lagoon, seed-bank culture, 31 July 1992,
M.T.Casanova p426, r977 (MEL); Beardy Waters, outside Glen Innes,
M.T.Casanova, M.Feist & A. García r395 (MEL, NE); Little Manning
River, between Gloucester and Walcha, 31 Oct. 1996, M.T.Casanova
r397 (MEL); Deua River (Moruya River) Araluen Road, 9 Jan. 2009,
M.T.Casanova r510, r514 (BM, MEL, NY); Tuross River on Comerang
Road, Silo Farm Bridge, 10 Jan. 2009, M.T.Casanova r523 (BM, MEL,
NY); Tuross River at Eurobodalla, Tyrone Bridge on Nerrigundah
Road, 10 Jan. 2009, M.T.Casanova r528 (MEL); Dumaresq Dam, 17
Feb. 1988, M.T.Casanova r996 (MEL); Gledwood, Camden, cultured
in biophysics laboratory, University of Sydey, 5 Dec. 1960, R.D.Wood
60-12-5-1 (AD, PC); Duval Creek, near Armidale, 17 Nov. 1960,
R.D.Wood 60-11-17-14A and 15A (AD). AUSTRALIAN CAPITAL
TERRITORY: Paddys River, 5 Apr. 1964, N.Burbidge 7428 (CANB);
Point Hutt, Murrumbidgee River, 8. Jan. 1966, E.D’Arnay 466 (CANB,
MEL); Murrumbidgee River, Uriarra, 4 Apr. 1964, H.S.McKee 11431
(CANB). VICTORIA.: Fitzroy River bridge, Tyrrendarra,
A.C.Beauglehole 203 (MEL); Mount Emu Creek, Panmure Waterhole,
3 Mar. 2004, M.T.Casanova & R.L.A.Casanova p587 (MEL, NY); Lake
Fyans, 24 Oct. 2005, M.T.Casanova p788 (MEL); Woorndoo Olympic
Wetland, 24 Feb. 2008, M.T.Casanova r058 (MEL); Darlots Creek,
Tyrendarra Bridge at the Budj Bim IPA, 29 Mar. 2011, M.T.Casanova
r952 (MEL, NY); The Woolwash, Surry River, 2.5 miles SE of
Heathmere, 31 Mar. 1962, A.Fuhrer & A.C.Beauglehole 181 (MEL,
NY); Snowy River upstream of Sandy Creek confluence, 24 Jan. 1994,
N.G.Walsh 2417 (MEL); Wonga Park, Warrandyte State Park, 3 Apr.
1994, D.J.van Bockel 384 & P.Coupar (MEL). NEW ZEALAND:
Lake Oahu, Canterbury, South Island, 3 Mar. 2001, J.S.Clayton r402
(MEL).
Australian Systematic Botany
27
branchlet cell long, in diameter similar to the other branchlet
cells. Bract cells small, 2–4 per branchlet node,
200–600 mm long, bracteoles difficult to distinguish, 2,
similar to bract cells. Gametangia conjoined, clustered inside
and below the branchlet whorls, and solitary or geminate on the
lowest two branchlet nodes. Oogonia 0.8–1.0 mm long,
0.6–0.8 mm wide, coronula cells appressed, slightly apiculate.
Oospores black, cylindrical, 600–830 mm long (778 5 mm:
Mandal et al. 2008) and 450–600 mm wide, 6 or 7 striae of low
ridges, sometimes slightly flanged, the impression of the basal cell
165–180 mm in diameter. In a population from China (Lu &
Soulié-Märshe, 1996), a proportion of the oospores had a 6-sided
basal-cell impression, rather than a pentagonal basal-cell
impression. Antheridia below or adjacent to the oogonia where
they are at the same node, 500–750 mm in diameter. The
antheridia appear tetrascutate. Chromosomes n = 42 (Khatun
et al. 2009).
Nomenclatural notes
Living material of Chara corallina was not available for
examination and the above description is based on the type
material, the protologue, Nordstedt (1883), Imahori (1954) and
the description of Indian material by Wood and Imahori (1964,
icon 111). The status of C. corallina var. basilaris A.Braun, with
gametangia absent from the branchlets and restricted to the
base of the whorls, and C. corallina var. kyusyensis Imahori
characterised by an almost complete absence of bract cells and
stipulodes, cannot be commented on without reference to fresh
specimens. The holotype material of C. corallina is lodged in the
Willdenow collection in B, but oospores were not available for
examination, apparently having been removed (R. Jahn, pers.
comm.). Examination of material in the Sonder Herbarium at
MEL revealed two specimens with morphology identical to the
type material in B-W. One appears to have been used as the model
for Kützing’s (1857) tabulae 80. These have been identified as
isotypes of C. corallina, and the material is in good condition
compared with the material in B-W (which appears to have been
moistened, dissected and re-pressed).
Recognition
Chara corallina Klein ex Willd., Mém. Acad. Roy. Sci. Hist.
(Berlin): 89 (1805)
Chara corallina is the only monoecious member of Chara sect.
Protochara.
(Fig. 2)
Nitella corallina (Klein ex Willd.) C.Agardh, Syst. alg. 123 (1824).
Type: INDIA: Wöppanpasi, Tranquebar (Tharamgambadi),
near Madras (Chennai), Tamil Nadu, 7 Jan. 1799, Klein 510.
Holo: B-W 17105; iso: MEL 2315415 & 2367911.
Monoecious. Plants usually elongate with stout branchlets,
up to 40 cm tall, sometimes lightly encrusted with calcium
carbonate. Axes up to 1.5 mm in diameter (~2 mm in flattened,
preserved specimens), ecorticate, internodes up to 30 mm long,
as long as or longer than the branchlets. Stipulodes crowded
out by gametangia, 1 per branchlet, up to 380 mm long, often
difficult to discern on pressed specimens. Branchlets 6 or 7 in a
whorl, 20–60 mm long, somewhat pinched-in at the nodes, and
appearing quite wide on pressed material, ecorticate, 4 or 5 cells
long, branchlet end segment pointed, conical and acute, basal
Distribution
Chara corallina is apparently widespread in Asia, occurring in
India, Bangladesh, Malaysia, Japan and China, at least as far north
as Nanjing.
Etymology
The term ‘corallina’ is derived from Latin corallinus meaning
‘coral-red’, referring to the bright colour of the gametangia.
Specimens examined
INDIA: near Bombay, G.O.Allen 105 (BM). CHINA: Chengdu City,
Sichuan, 1996, Lu Hui-nan & I.Soulié-Märsche s.n.,(NY, MEL:
oospores only).
28
Australian Systematic Botany
A
M. T. Casanova and K. G. Karol
B
D
C
E
F
G
Fig. 2. Chara corallina Klein ex Willd. A. Habit of whole plant; scale bar: 1 cm, after Kützing (1857). B. Upper branchlet whorl; scale bar: 1 mm.
C. Lower branchlet whorl, two branchlets cut short; scale bar: 1 mm. D. Oogonium and branchlet node with bract cells; scale bar: 1 mm. E. Oospore;
scale bar: 200 mm. F. Detail of oospore wall; scale bar: 20 mm.; G. Basal-cell impression, atypically six-sided; scale bar: 50 mm. A–C from J.G.Klein
510; E, F from G.O.Allen 104 and G from Lu & Soulié-Märshe 1996.
Chara lucida (A.Braun) Casanova & Karol, comb. et stat. nov.
(Fig. 3)
Chara australis var. lucida A.Braun in Nordst., Abh. Königl. Akad.
Wiss. Berlin 1882: 106 (1883); Chara australis var. lucida f. typica
Zaneveld, nom. inval., Blumea 4: 126 (1940); Chara corallina f.
lucida R.D.Wood, nom. inval., Taxon 11: 13 (1962); Chara
corallina f. lucida R.D.Wood, nom. inval., nom. alt., Rev.
Characeae 1: 275 (1965); Chara lucida R.D.Wood, nom. inval.,
nom. alt., Rev. Characeae 1: 769 (1965).
Type: Northern Territory: Victoria River, s. dat., F.Mueller 5.
Lecto (fide Zaneveld 1940): LD; isolecto: L. Syn: Northern
Territory: Baines Creek, 1856, F.Mueller s.n.: LD.
Chara australis f. tenerior Zaneveld, Blumea 4 : 127 (1940).
Type: Gulf of Carpentaria, 1856, F.Mueller s.n. Holo: B
(destroyed; fide Wood and Imahori (1965: 276)); iso: LD, L.
Dioecious. Plants flexible, transparent, often in tangled
clumps, up to 200 mm tall. Axes 0.35–0.7 mm wide (when flat
and dried, narrower in fresh material), ecorticate, internodes
8–20 mm long, generally much shorter than the adjacent
branchlets. Stipulodes in a single row, 6–12 in total, usually
fewer apparent, up to 0.5 mm long. Branchlets 6 or 7 in a whorl,
ecorticate, 11–50 mm long, 0.3–0.6 mm wide, 4 or 5 cells long,
basal branchlet cell up to 10 mm long, branchlet end segments
small, conical and acute, sometimes subtended by 1 or 2 bract
cells, bract cells obscure or short. Bracteoles obscure or short,
occasionally up to 0.5 mm long. Fertile parts sometimes
somewhat contracted with shorter branchlets and internodes.
Chara section Protochara in Australia
Australian Systematic Botany
29
Fig. 3. Chara lucida (A.Braun) Casanova & Karol. A. Habit of whole plant; scale bar: 5 mm. B. Fertile oogonial branchlet whorl; scale bar: 5 mm.
C. Final two branchlet cells; scale bar: 0.5 mm. D. Oogonium; scale bar: 100 mm. E. Fertile antheridial branchlet whorl; scale bar: 5 mm. F. Scanning
electron micrograph (SEM) of oospore; scale bar: 200 mm. G. SEM of detail of oospore wall; scale bar: 50 mm. H. SEM of impression of basal cell on
the oospore; scale bar: 100 mm. A–D from M.T.Casanova r784; E from L.A.Craven 4782 and F–H from F.Mueller 5.
Specimens with only fertile branchlets were distinguished as
form tenerior but recent collections (e.g. M.T.Casanova r758)
have both long sterile and short fertile branchlets on the same
plant. Gametangia sessile inside the base of the whorl and
solitary, geminate or clustered at the lowest 1 or 2 branchlet
nodes. Oogonia 0.9 mm long, 0.5 mm wide, coronula cells very
short, obtuse. Oospores black, cylindrical, 600–670 mm long,
380–400 mm wide, 7 or 8 striae of low ridges, ornamentation
smooth to minutely granulate, basal-cell impression 120–150 mm
in diameter at the widest part, edges 95–100 mm long. Antheridia
from 400–850 mm in diameter. Vegetative reproduction not
known. Chromosomes not known.
Nomenclatural notes
Wood’s (1962) combination Chara corallina f. lucida is here
treated as invalid, under Art. 41.5 of the ICN (McNeill et al.
2012). Wood and Imahori (1965) subsequently simultaneously
published C. corallina f. lucida and C. lucida, the latter at the rank
of ‘microspecies’, equivalent in rank to Wood’s forms as well as
30
Australian Systematic Botany
his ‘monotypic’ varieties and species (i.e. those lacking
subordinate taxa). Both these names are here treated as invalid
alternative names, under Art. 36.2 of the ICN (McNeill et al.
2012).
Taxonomic notes
Zaneveld (1940) distinguished the following two formae in his
concept of C. australis var. lucida, on the basis of material from
New Guinea and northern Australia: f. typica, with somewhat
stout (~600 mm wide), long branchlets, and f. tenerior, with
shorter, narrower (~350 mm wide) branchlets. New collections
from the Northern Territory (M.T.Casanova r784, r758)
represent material with both long and short branchlets on
apical parts of the same plant, and branchlet width varies in
relation to branchlet length. Additionally, specimens from the
Northern Territory and the Gulf of Carpentaria have identical
oospores, and accordingly, no infraspecific taxa are here
recognised in this species.
Recognition
The species can be distinguished as a totally ecorticate Chara
species with narrow axes, distinguished from narrower morphs
of C. australis by the short internodes in comparison with the
branchlets. C. lucida forms tangled masses in warm waters,
compared to the upright, turgid stems of C. australis in colder,
deeper waters.
Distribution
Tropical and subtropical Queensland, Western Australia and the
Northern Territory in dams, ponds, still rivers and lagoons. Also
occurs in Papua New Guinea.
Etymology
From the Latin ‘lucidus’, meaning shining, clear, transparent.
Zaneveld (1940) and Wood (1971) thought this referred to a shiny
surface on the pressed specimens, but that feature is neither
obvious nor consistent. More likely, it refers to the clear and
transparent, almost colourless appearance (fide Nordstedt 1883)
of the pressed type material.
Specimens examined
WESTERN AUSTRALIA: Rudall River, 29 May 2004, M.N.Lyons &
D.A.Mickle 3057 (PERTH); Redrocks Granites, Pilbara Biological
Survey site 080, 25 Aug. 2005, N.M.Lyons & D.A.Mickle 3160
(PERTH); Kumina Creek, Hamersley Ranges, 13 Aug. 1991,
M.Trudgen & D.True MET10264 (CANB, AD, PERTH).
NORTHERN TERRITORY: Manton Dam recreation area,
M.T.Casanova r758 (MEL, NY); roadside borrow-pit on Arnhemland
Highway, M.T.Casanova r784 (B, MEL, NY); SW of Elizabeth Downs,
Daly River region, 23 June 1977, L.A.Craven 4375 (CANB); Oenpelli,
10 Oct. 1948, R.L.Specht A110 (AD, MEL, NY); Katherine, 14 Feb.
1961, H.S.McKee 8427 (CANB);. QUEENSLAND: Ross River Dam
overflow near Townsville, 12 May 1987, M.T.Casanova r997; in lagoon
at Magoura Station, 18 July 1977, L.A.Craven, 4782 (CANB); Pig
Swamp near Biboohra, N of Mareeba, 27 June 1962, R.D.Hoogland
8492 (CANB); Hastie Swamp, Atherton Tablelands, 6 Aug. 1970,
A.P.Kershaw 10109 (CANB); Breeza Plains, 12 Aug. 1978,
K.Paijmans 2871 (CANB); Enoggera Reservoir, 21 Nov. 1960,
R.D.Wood 60-11-21-25 (AD, NY); Palm Creek at Miriam Vale, 30
M. T. Casanova and K. G. Karol
Nov. 1960, R.D.Wood 60–11–30–6 (AD, NY); South Pine River,
Bunya, 22 Nov. 1960, R.D.Wood 60-11-22-20 (AD, NY). PAPUA
NEW GUINEA: Hisin Village, Kairuku, Central District, 16 Aug.
1962, P.J.Darbyshire 813 (CANB); Laloki River, Central District,
5 Sep. 1962, T.G.Hartley 10619 (CANB); Rouna Falls, Laoki River,
Central District, 7 Sep. 1962, R.Schodde 2946 (CANB).
Chara porteri Casanova, sp. nov.
(Fig. 4)
Type: Western Australia: Ethel Creek Claypan, Pilbara
Biological Survey site 041, T2, 1 June 2004, M.N.Lyons &
D.A.Mickle 3077. Holo: PERTH; iso: MEL.
Dioecious. Plants somewhat inflated and turgid, sometimes
calcified, up to 150 mm tall, somewhat densely branched, with incurved branchlets in shallow water. Axes 0.7 mm wide (up to
1 mm on pressed specimens), ecorticate, internodes 8–25 mm
long, shorter and somewhat contracted at the apices. Rarely
internodes appear 2 cells long. Stipulodes in a single row
when present, frequently obscure or absent. Branchlets 6 in a
whorl, sometimes made up of a few elongate branchlets and some
dwarf branchlets, particularly at the apices, ecorticate, 8–15 mm
long, 0.4–0.6 mm wide, 3 or 4 cells long, basal branchlet cell up
to 6 mm long, usually the longest cell in the branchlet, branchlet
end segments small, conical and acute, occasionally subtended
by bract cells, bract cells usually obscure, up to 0.2 mm long,
or absent. Bracteoles obscure, up to 0.2 mm long, or absent.
Gametangia on separate plants, sessile inside the base of the
whorl, and solitary, geminate or clustered at the lowest 1 or 2
branchlet nodes. Oogonia 0.6–0.75 mm long, 0.5 mm wide,
coronula cells short, somewhat spreading, slightly apiculate.
Oospores black, cylindrical, 470–525 mm long, 250–350 mm
wide, 5 or 6 striae of wide ridges (pachygyra), ornamentation
smooth to minutely granulate, basal-cell impression 100–120 mm
in diameter at the widest part, edges 70–76 mm long, rarely
6-sided. Antheridia octoscutate up to 850 mm in diameter.
Vegetative reproduction not known. Chromosomes n = 14
(M.T.Casanova r416).
Recognition
Specimens are distinctly turgid and brightly festooned with
gametangia, with incurved branchlets. The oospores have
markedly enlarged striae (pachygyra).
Distribution
Occurs in temporary wetlands (claypans and riparian overflows)
in the Paroo region of New South Wales, the Pilbara region of
Western Australia and in south-western Queensland.
Etymology
Named in honour of John L. Porter, who collected the first
herbarium material of this species, from the Paroo region in
New South Wales.
Specimens examined
WESTERN AUSTRALIA: Coondiner Pool, Pilbara Biological Survey
site 001, 15 Aug. 2006, M.T.Casanova r416 and r417 (B, BM, MEL);
Cooliarin Pool, M.N.Lyons & D.A.Mickle 3061 (paratype) (CANB, MEL,
PERTH); Myanore Creek Pool, Pilbara Biological Survey site 014,
Chara section Protochara in Australia
Australian Systematic Botany
Fig. 4. Chara porteri Casanova. A. Habit of whole plant; scale bar: 1 cm. B. Branchlet whorl from an oogonial plant; scale bar: 5 mm. C. Individual
oogonial branchlet; scale bar: 1 mm. D. Antheridial branchlet whorl; scale bar: 1 mm. E. Individual antheridial branchlet; scale bar: 1 mm.
F. Oogonium; scale bar: 500 mm. G. Axis and branchlet whorl, with a bicellulate axis; scale bar: 1 mm. H. Oogonial axis tip, with clustered oogonia at
the base of the branchlets; scale bar: 1 mm. I. Oospore; scale bar: 100 mm. J. Atypical six-sided basal-cell impression; scale bar: 50 mm. K. Oospore;
scale bar: 100 mm. A from M.N.Lyons & D.A.Mickle 3061; B–F, I–J from M.N.Lyons & D.A.Mickle 3045 and G–H and K from M.N.Lyons &
D.A.Mickle 3077.
31
32
Australian Systematic Botany
13 May 2006, D.A.Mickle & N.Y.Huang 3108, 3109 and 29-3108
(BM, MEL, PERTH); Watrara Creek Pool, Pilbara Biological Survey
site 028, 8 Sep. 2003, M.N.Lyons & S.D.Lyons 3045 (PERTH); Cooliarin
Pool, Pilbara Biological Survey site 033, 18 May 2004, M.N.Lyons &
D.A.Mickle, 3064 (CANB, MEL, PERTH); Cooliarin Pool, Pilbara
Biological Survey site 033, 20 Apr. 2006, M.N.Lyons & D.A.Mickle
3087 (AD); Cooliarin Pool, Pilbara Biological Survey site 033, 20 Apr.
2006, M.N.Lyons & D.A.Mickle 3089, 3090 and 13-3092 (PERTH);
Sweetwell Claypan, Pilbara Biological Survey site 038, 21 May 2004,
M.N.Lyons & D.A.Mickle 3069 (PERTH); Ethel Creek Claypan, Pilbara
Biological Survey site 041, 1 June 2004, M.N.Lyons & D.A.Mickle
3074 (NY, BM); Redrocks Granites, Pilbara Biological Survey site
080 T1, 25 Aug. 2005, M.N. Lyons & D.A. Mickle 3159 (PERTH).
QUEENSLAND: Cravens Peak site 4, Little Kanamuka, 17 Apr. 2007,
I.J.Powling r399 (MEL). NEW SOUTH WALES: Pied Stilt Swamp,
27 Jan. 2000, J.L.Porter 239 & 240 (MEL).
Chara protocharoides Casanova & Karol, nom. nov.
(Fig. 5)
Protochara australis Womersley & Ophel, Proc Roy. Soc.
S. Australia 71: 311 (1947); Chara australis subsp. estipulodica
M.B.Macdon. & Hotchk., Proc. Linn. Soc. New South Wales 80:
282 (1956) (non Chara australis R.Br., Prodr. 346 (1810)).
[Chara australis f. inflata auct. non R.D.Wood, nom. inval.: R.D.
Wood, Nova Hedwigia 22: 25 (1972), p.p.].
Type: Western Australia: In shallow swamp on peneplain of
breakaway country between Mingenew and Irwin River coal
seam, south-west of Geraldton, 28 Aug. 1947, H.B.S.Womersley
s.n. Holo: AD A5917a; Iso: AD A5917c, NY 01089127.
Dioecious. Plants up to 40 cm high, somewhat branched,
variably inflated. Axes ecorticate, internodes 10–100 mm long,
0.9–1.5 mm in diameter in life (up to 2 mm when flattened
in pressing). Stipulodes completely absent. Branchlets 6 or 7
in a whorl, entirely ecorticate, segments swollen or inflated in
shallow water, elongate and cylindrical in deep-water
populations, pinched in at the nodes, similar in diameter to the
axes, 14–30 mm long, 3 or 4 cells long including end segment,
basal branchlet cell variable in size, ~300 mm long in upper,
fertile branchlets, similar to second branchlet segments on sterile
branchlets, branchlet end segments unicellular, very small and
mucronate, up to 200 mm long, with nodal cells at the base, bract
cells and bracteoles completely absent from all branchlet nodes.
Upper axes somewhat contracted. Gametangia on separate
plants, singly and geminate on first and second branchlet
nodes, occasionally oogonia inside the base of the branchlet
whorl. Oogonia 670–900 mm long 600–780 mm wide with
6 or 7 convolutions, coronula cells connivent and blunt,
80 mm high. Oospores black, 490–560 mm long and
310–390 mm wide, almost rectangular in side view, with 4 or
5 striae, sometimes flanged, and impression of the basal cell
80 mm in diameter. Antheridia 800–1150 mm in diameter,
octoscutate. Vegetative reproduction not known; appears to be
an annual in temporary water bodies. Chromosomes not known.
Taxonomic notes
Originally placed in the segregate genus Protochara on the basis
of very simple, inflated axes and the lack of accessory cells (bract
cells, stipulodes or bracteoles) (Womersley and Ophel 1947).
Apart from the absence of any accessory cells, type material of
M. T. Casanova and K. G. Karol
Protochara australis is otherwise consistent with the morphology
of the genus Chara. The epithet australis is preoccupied in Chara
(C. australis R.Br., 1810), and a new name is required for this
taxon.
Recognition
The complete absence of stipulodes, bract cells and bracteoles are
the main characteristics that allow this species to be distinguished.
Distribution
Grows in permanent, seasonal and temporary water bodies in
Western Australia.
Etymology
From the Latin ‘protocharoides’ in reference to the genus
Protochara for which this species remains the type.
Specimens examined
WESTERN AUSTRALIA: Bluegum Lake, Brentwood, Perth, 3 Oct.
2002, M.T.Casanova r392 (MEL); Swan Lake, Bayswater, Perth, 3 Oct.
2010, M.T.Casanova r805 (B, BM, MEL, NY, PERTH); small lake near
inflow of Lake Leschenaultia, 5 Oct. 2010, M.T.Casanova r812 (MEL,
NY, PERTH); River pool 2 km NNW of Tindellara Well, 20 Sep. 1987,
R.J.Cranfield 6246 (PERTH); Hammersley River, in freshwater holes,
12 Oct. 1901, F.L.E.Diels 4913 (B); interior of western Australia, 1890,
M.Heal (LD, MEL); Cape Bertholet, Dampierland, N of Broome, 18 Apr.
1977, K.F.Kenneally 6009 (PERTH); Yaladinia Rock Hole, 18 Sep.
1980, K.R.Newbey 7462A (PERTH); pool in woods, immediately S of
Piesseville, 11 Oct. 1960, R.D.Wood 60-10-11-9a (AD); roadside pool,
3 miles (~4.83 km) S of Gnowangerup Road intersection, Tambellup,
10 Oct. 1960, R.D.Wood 60-10-10-11A (AD, CANB, PC).
Chara stuartiana (F.Muell. ex Kütz.) Casanova & Karol,
comb. et stat. nov.
(Fig. 6)
Nitella stuartiana F.Muell. ex Kütz., Tab. phycol. 7: 28 (1857);
Chara stuartiana A.Braun, Linnaea 25: 707 (1853), nom.
inval., pro syn.; Chara australis f. stuartiana (F.Muell. ex
Kütz.) Zaneveld, Blumea 4: 126 (1940); Chara corallina f.
stuartiana R.D.Wood, nom. inval., Taxon, 11: 13 (1962); Chara
corallina f. stuartiana R.D.Wood, nom. inval., nom. alt., Rev.
Characeae 1: 275 (1965); Chara stuartiana R.D.Wood, nom.
inval., nom. alt., Rev. Characeae 1: 769 (1965).
Type: Tasmania: South Esk River, R.C.Gunn 1565, 1852.
Holo: L; iso: PC.
[Chara australis var. nobilis auct. non A.Braun in C.F.O.Nordstedt,
Abh. Königl. Akad. Wiss. Berlin 1882: 105 (1883).]
Dioecious. Plants large and inflated, transparent and turgid,
up to 27 cm high, but fragile, fragmenting easily when handled.
Axes 1.5–2.8 mm wide, ecorticate, internodes 30–120 mm long.
Stipulodes in a single row, tiny, up to 12 per whorl, up to
400 mm long, alternate to the branchlets and pointing
upwards. Branchlets 6 in a whorl, 20–30 mm long,
1.6–2.8 mm wide, 3 or 4 cells long, all cells cylindrical, end
segment a single cell, or subtended by small bract cells, basal
branchlet cell elongate and cylindrical, the second branchlet cell
is usually shorter than the basal branchlet cell, abbreviated and
sometimes almost spherical, particularly on the upper parts of the
Chara section Protochara in Australia
Australian Systematic Botany
33
Fig. 5. Chara protocharoides Casanova & Karol. A. Habit of plant; scale bar: 1 cm. B. Whorl of antheridial branchlets; scale bar: 1 mm.
C. Branchlet tip with single end cell; scale bar: 1 mm. D. Branchlet node and antheridium; scale bar: 0.5 mm. E. Oogonial branchlet whorl;
scale bar: 1 mm. F. Oospore; scale bar: 200 mm. G. Detail of oospore wall; scale bar: 20 mm. H. Impression of end cell; scale bar: 100 mm.
A–D, F–H from H.B.S.Womersley 28-Aug-1947; E from M.T.Casanova r812.
axis; bract cells 2 or 3 at branchlet nodes, up to 400 mm long.
Bracteoles 2, up to 200 mm long. Gametangia on separate plants,
solitary or geminate at all branchlet nodes. Oogonia at the lowest
branchlet node, up to 1 mm long and 700 mm wide, with 8
convolutions, coronula very small and minutely apiculate.
Oospores black, cylindrical to ovate, 615–700 mm long,
450–550 mm wide, with 6 or 7 striae of low ridges, basal-cell
impression up to 140 mm in diameter at widest point, edges
85–115 mm long. Antheridia up to 500 mm in diameter, solitary,
geminate or aggregate at the lowest branchlet node and inside
the base of the whorl. Vegetative reproduction by starch
accumulations in the lower and rhizoidal nodes. Chromosomes
not known.
Nomenclatural notes
Wood’s (1962) combination Chara corallina f. stuartiana is here
treated as invalid, under Art. 41.5 of the ICN (McNeill et al.
2012). Wood and Imahori (1965) subsequently simultaneously
published C. corallina f. stuartiana and C. stuartiana, the latter at
the rank of ‘microspecies’, equivalent in rank to Wood’s forms as
well as his ‘monotypic’ varieties and species (i.e. those lacking
subordinate taxa). Both these names are here treated as invalid
alternative names, under Art. 36.2 of the ICN (McNeill et al.
2012).
Taxonomic notes
Braun (1852) included a specimen collected by C. Stuart in
Tasmania (Stuart 1565) in Chara australis. Kützing (1857)
named the species Nitella stuartiana based on Stuart 1565 and
provided an illustration (t. 28). Braun (1860) listed a collection
from Tasmania, Gunn 1565, in his concept of C. australis. This is
likely to refer to the same specimen as cited by Braun (1852) and
Kützing (1857), and was collected by Gunn rather than Stuart,
because the specimens attributed to Gunn have collection
numbers for charophytes in the vicinity of Launceston (and the
South Esk River) in the 1560s, and Stuart’s collection numbers
for the South Esk River are either very low (3, 4, 5), in the 210s
or in the 1020s. Only two specimens in sect. Protochara collected
34
Australian Systematic Botany
M. T. Casanova and K. G. Karol
Fig. 6. Chara stuartiana (F.Muell. ex Kütz.) Casanova & Karol, from specimen from the type locality, H. & A.Wapstra HAW044. A. Habit of
whole plant; scale bar: 5 cm. B. Base of a branchlet whorl; scale bar: 200 mm. C. Two branchlet nodes; scale bar: 1 mm. D. Oogonium and end cells of
branchlet; scale bar: 1 mm. E. Sterile branchlet whorl; scale bar: 1 cm. F. Branchlet end cells; scale bar: 0.5 mm. G. Scanning electron micrograph
(SEM) of oospore; scale bar: 200 mm. H. SEM of detail of oospore wall; scale bar: 50 mm. I. SEM of impression of the basal cell; scale bar: 100 mm.
from Tasmania in the 1800s are known, including material
used by Kützing for his illustration, almost certainly obtained
from Sonder’s herbarium (L), and a specimen labelled ‘Nitella
australis R.Br. Donné par Sir William Hooker, 1863; Ex Herb.
Hook. Chara australis Br, Tasmania, Coll. R.C.Gunn’ (PC).
It is possible that these are parts of the same gathering, and
therefore the PC specimen would constitute isotype material.
Braun, in Nordstedt (1883), retained N. stuartiana in synonymy
of C. australis, but cited another specimen from Tasmania
(Oldfield 283 n.v.) as C. australis var. nobilis. Zaneveld
(1940) treated N. stuartiana as a form of C. australis var.
nobilis. Proctor (1999) found that specimens identified as
C. australis and C. stuartiana could be crossed to produce
oospores, and explained C. stuartiana as a ‘genetically fixed
morphotype of C. australis’. Examination of the type material and
more recent collections indicated that most of specimens from
Tasmania in sect. Protochara can be clearly separated from
C. australis on the basis of more robust axes and branchlets,
complex axial and branchlet nodes, overall fragility and different
oospores. C. stuartiana is rarely collected as a whole plant,
because the heavy, turgid axes are fragile and tend to break as
they are removed from the water. The species still occurs in what
is presumed to be the type locality, namely, the first basin of
Cataract Gorge in Launceston. Specimens collected from
Kimberley Warm Springs (T.Dugdale & S.Talman 20, HO,
MEL, NY; H. & A.Wapstra HAW002, MEL, HO) appear to be
C. australis rather than C. stuartiana, but if so, that is the only
occurrence of C. australis known from Tasmania.
Recognition
Chara stuartiana has large, fragile, often inflated, ecorticate axes
and branchlets, with orange antheridia and oogonia. It can be
distinguished from C. australis by its wide axes and branchlets,
and by the abbreviated penultimate cell on the upper branchlet
whorls.
Distribution
Grows in Tasmania in rivers and streams, at depth.
Chara section Protochara in Australia
Etymology
Named for Charles Stuart (1802–1877) who collected
charophytes in Tasmania from 1842 to 1852, initially for R.C.
Gunn, later for F. von Mueller (Orchard 1999).
Specimens examined
TASMANIA: Apsley River at Coles Bay Road, 7 July 2008, H. &
A.Wapstra HAW009 (HO, MEL); Prosser River, Paradise Gorge, 1 Oct.
2008, H. & A.Wapstra HAW027 (HO, MEL); South Esk River, Glen Esk
Bridge, 21 Oct. 2008, H. & A.Wapstra HAW044 (B, HO, MEL, NY);
South Esk River, 22 Oct. 2008, H. & A.Wapstra HAW048 (HO, MEL,
NY); Meredith River at Swansea, 22 Apr. 2009, H. & A.Wapstra
HAW133 (HO, MEL); Curries River reservoir, 14 Oct. 2008,
M.Wapstra HAW034 (HO, MEL); farm dam, Bridport road between
Yarrow Creek and Pipers River, 15 Oct. 2008, M.Wapstra HAW036
(HO, MEL, NY); first basin of Cataract Gorge, 20 Feb. 2013, M.Wapstra
HAW145 (HO, MEL); Elizabeth River, Campbell Town, 19 Jan. 1961,
R.D.Wood 61-1-19-3 (AD, NY, PC); stream 6 miles (~9.66 km) S of
Sorrel, 1 Feb. 1961, R.D.Wood 61-2-1-2 (AD, NY, PC); 15 miles
(~24.14 km) S of Seymour, 2 Feb. 1961, R.D.Wood 61-2-2-6 (AD,
NY); St Pauls River, 3 Feb. 1961, R.D.Wood 61-2-3-6 (AD, NY);
South Esk River, 3 Feb. 1961, R.D.Wood 61-2-3-7 (AD); 2 miles
(~3.22 km) N of Longford, 4 Feb. 1961, R.D.Wood 61-2-4-14 (AD, NY).
Chara wallichii A.Braun in C.F.O. Abh. Königl. Akad. Wiss.
Berlin 1882: 107 (1883)
(Fig. 7)
Chara corallina var. wallichii R.D.Wood, nom. inval., Taxon 11: 13
(1962); Chara corallina var. wallichii (A.Braun) R.D.Wood, Rev.
Characeae 1: 278 (1965), nom. inval., nom. alt.
Australian Systematic Botany
35
Type: INDIA: Gangetic Plain, leg. ign., 9 Jan. 1809. Holo:
LINN, n.v.
Dioecious. Plants up to 25 cm high, prickly in appearance.
Axes ecorticate, 875–1100 mm wide, internodes somewhat
shorter than the branchlets. Stipulodes well developed and
somewhat inflated where present, up to 450 mm long.
Branchlets 6 in a whorl, ecorticate, up to 30 mm long, 4–6
cells long, basal branchlet cell longest, the end segment short,
conical and acute. Bract cells well developed, 1–4 at a node, up
to 2 mm long. Gametangia on separate plants, clustered at the
base of the whorl and singly or geminate at all branchlet
nodes. Oogonia up to 745 mm long, 700 mm wide, with 7 or 8
convolutions. Oospores 500–610 mm long, 330–440 mm wide
with 6 or 7 flanged ridges. Antheridia 750–950 mm in diameter,
tetrascutate (Wood and Imahori 1965). Chromosomes not
known.
Nomenclatural notes
Wood’s (1962) combination Chara corallina f. wallichii is here
treated as invalid, under Art. 41.5 of the ICN (McNeill et al.
2012). Wood and Imahori (1965) subsequently simultaneously
published C. corallina var. wallichii and C. wallichii, the latter at
the rank of ‘microspecies’, equivalent in rank to Wood’s forms as
well as his ‘monotypic’ varieties and species (i.e. those lacking
subordinate taxa)). C. corallina var. wallichii is here treated as
invalid, under Art. 36.2 of the ICN (McNeill et al. 2012).
Recognition
Chara wallichii is a dioecious species that, in contrast to most
other species in the group, is distinguished by well developed,
Fig. 7. Chara wallichii A.Braun, from Parsotam 230. A. Habit of whole plant; scale bar: 1 cm. B. Branchlet tip; scale bar: 1 mm. C. Branchlet
whorl from an antheridial plant; scale bar: 1 mm. D. Oospore; scale bar: 200 mm. E. Detail of oospore wall; scale bar: 20 mm.
36
Australian Systematic Botany
somewhat inflated, stipulodes and bract cells. Material seen in
BM and BP is relatively uniform.
Distribution
India and Bangladesh.
Etymology
Named in honour of Nathaniel Wallich (1786–1854), a Danish
botanist involved in the development of the Calcutta Botanic
Garden.
M. T. Casanova and K. G. Karol
Several other species have been recorded from the Indian
subcontinent (Chara fulgens Fil., C. pashanii S.C.Dixit and
C. nuda Pal) and oceanic islands of the southern hemisphere
(Chara socotrensis Nordst. from Socotra, C. corallina f.
vieillardii (A.Braun) R.D.Wood from New Caledonia,
C. corallina f. vitiensis Nordst. from Fiji and C. corallina f.
mascarensis J.Groves & Stephens, nom. inval., nom. prov. from
Mauritius); however, because specimens have not been
examined, no comment on their status can be made in the
present study.
Specimens examined
Discussion
INDIA: Sarmalt, Benares, Uttar Pradesh, Jan. 1934, Parsotam 230,
(BP, BM).
Species in sect. Protochara have few characters to aid
discrimination, a fact that has contributed to the confused
history of generic and species names in this group. Axis and
branchlet diameter and the expression of accessory cells
(stipulodes, bract cells) can allow discrimination in well grown
plants; however, the occurrence and expression of basal
gametangia and abundance of branchlet gametangia probably
vary in relation to plant nutrition and age and there are examples
of specimens that fail to conform to the ‘average’ morphology
(e.g. from cultured material, young plants growing in shallow
water, or etiolated plants from deep water). Reference to the
oospore morphology is necessary to distinguish such doubtful
specimens.
Separation of Chara sect. Protochara and the genus
Lamprothamnium (also ecorticate, sometimes inflated,
sometimes with few accessories) on the basis of vegetative
morphology will always be something of a challenge,
particularly with material lacking diagnostic characters. These
two taxa appear to be closely related on the basis of nucleotidesequencing studies (Meiers et al. 1999; McCourt et al. 2000). The
main characters to aid identification are that Lamprothamnium
species usually have more than six branchlets in a whorl,
their stipulodes are opposite and decumbent when present,
and the oospores possess gyrogonites (a calcified covering).
Additionally, most members of sect. Protochara occur in
freshwater, whereas Lamprothamnium species occur in
brackish to saline water.
Chara braunii (sect. Charopsis) can be separated from
members of sect. Protochara phylogenetically (Meiers et al.
1999; McCourt et al. 2000) and morphologically. C. braunii
has well developed stipulodes and bract cells, and the branchlets
are terminated by a ‘corona’ of approximately three equal-sized
cells, rather than a single cell, as in sect. Protochara. C. braunii
is a relatively ephemeral charophyte, and is rarely found in
Australia, with most specimens referred to C. braunii in
Australian herbaria being referable to C. muelleri.
Additional taxa of uncertain status
Tolypellopsis simplicissima Fil., Arch. Hydrobiol., Suppl. 12,
Tropische Binnengewässer 3: 716 (1934)
Known only from the type gathering from Lake Toba, a deep,
oligotrophic caldera in Sumatra, in 1929. The morphology of this
species is similar to that of C. australis; however, the oogonia are
much smaller and more spherical than those of any other species
in sect. Protochara. Because of its overwhelming morphological
similarity to other species in sect. Protochara, it is likely to be a
species of Chara. It does not appear to have many accessory cells
(stipulodes, bract cells); however, further collections of mature
female material is required for confirmation of the identity and
placement of this taxon.
Chara plebeja R.Br. ex A.Braun, Linnaea 17: 118 (1843)
Known only from the type gathering from ‘Carpentaria Point’,
possibly the ‘Carpentaria Main’ of Brown’s and Flinders’ diaries,
now known as Cape Shield (Groves and Moore 1989), in Arnhem
Land near Groote Eylandt in the Gulf of Carpentaria. Satellite
images of the locality of Cape Shield revealed a hind-dune
wetland area that could be where Brown collected this species.
Treated as a subspecies of C. australis by Braun in Nordstedt
(1883) and as a variety of C. australis by Zaneveld (1940), and
distinguished from C. australis sens. str. on the basis of obtuse
end cells on the branchlets. Detailed examination of the type
material revealed a few stipulodes and bract cells, and what appear
to be obtuse end cells are probably penultimate cells, with the
end cells dehiscent. Given the possible habitat and evidence
from the morphology (7 or 8 branchlets rather than 6,
stipulodes opposite the branchlet and somewhat decumbent), it
seems likely that C. plebeja is a species of Lamprothamnium.
If so, it would be the only known coastal Lamprothamnium
from the tropics. Further collections of fertile female material
are required to confirm this.
Chara sp. Woorndoo (M.T.Casanova p246)
This specimen is likely to represent a new species. Oospores are
similar in appearance to those of C. porteri, but are over
700 mm long
(c.f.
C.
porteri
with
oospores
470–525 mm long), and its distribution is disjunct from
C. porteri. However, there are insufficient gatherings to
confirm its status at the present time.
Acknowledgements
Thanks go to Professor Jim Ross, who, early in the project, said ‘if you don’t do
it, who will?’. For this study, access to type material in various European
(B, BM, BP, GFW, GOET, HBG, L, LD, PC and W) and Australian (AD,
BRI, CANB, DNA, HO, MEL, NSW, PERTH) herbaria was essential;
I thank the directors and staff of these institutions for their kind assistance
and permission to remove oospores for examination. Many people have
assisted with providing specimens, fresh and preserved, since 2000, some
of their names are noted in the specimen lists. This study would have
been impossible without their efforts. Mary Beth Williams provided notes,
Chara section Protochara in Australia
Australian Systematic Botany
translations and literature in an early part of this study. Professor
W. Prud’homme van Reine offered valuable references from Joop van
Raam’s literature collection. Funds for this work were provided by the
Australian Biological Resources Study Grant number 209-49, Bush Blitz
and Charophyte Services, and by the US National Science Foundation under
Grant numbers DEB-1020660 and DEB-1036466. We thank Joan Powling
for her meticulous proofreading and an anonymous referee for suggesting
improvements. The assistance of Brendan Lepschi, Peter Wilson and Nick
Turland with difficult nomenclatural issues is greatly appreciated.
References
Braun A (1852) Characeae. F. L. von Schlechtendal plantae Mullerianae.
Linnaea 25, 705–709.
Braun A (1860) Characeae. In ‘The Botany of the Antarctic Voyage III. Flora
Tasmaniae. Vol. 2’. (Ed. JD Hooker) pp. 159–160. (Lovell Reeve:
London)
Brown R (1810) ‘Prodromus florae novae hollandiae et insulare van diemen.
Vol. 1.’ (Richard Taylor and Sons: London)
Casanova MT (1997) Oospore variation in three species of Chara (Charales,
Chlorophyta). Phycologia 36, 274–280.
doi:10.2216/i0031-8884-36-4-274.1
Casanova MT (2004) A census of submerged plants of the Angas River and
Tookayerta Creek catchments. Consultants report to the River Murray
Catchment Management Board. (Charophyte Services: Lake Bolac, Vic.)
Casanova MT (2005) An overview of Chara L. in Australia (Characeae,
Charophyta). Australian Systematic Botany 18, 25–39.
doi:10.1071/SB04027
Casanova MT (2009) An overview of Nitella in Australia (Characeae,
Charophyta). Australian Systematic Botany 22, 193–218.
doi:10.1071/SB08039
Casanova MT (2013) Lamprothamnium in Australia (Characeae,
Charophyta). Australian Systematic Botany 26, 268–290.
doi:10.1071/SB13026
Crawford SA, Higgins SJ, Mulvaney P, Wetherbee R (2001) Nanostructure of
the diatom frustule as revealed by atomic force and scanning electron
microscopy. Journal of Phycology 37, 543–554.
doi:10.1046/j.1529-8817.2001.037004543.x
Filarszky F (1934) Die Characeen der Deutschen Limnologischen SundaExpedition. Archiv fuer Hydrobiologie 4(Supp. 12.), 705–726.
García A, Chivas AR (2006) Diversity and ecology of extant and Quaternary
Australian charophytes (Charales). Cryptogamie. Algologie 27, 323–340.
García A, Karol KG (2004) A paradigm in the taxonomy of charophytes:
the oospore and gyrogonite of Nitellopsis inflata (Fil. & G.O.Allen ex
Fil.) = Lamprothamnium inflatum comb. nov. In ‘Proceedings of the
Fourth International Congress on Extant and Fossil Charophytes’,
October 2000, Robertson, NSW, Australia. (Eds A García, A Chivas)
p. 42. (University of Wollongong: Wollongong, NSW)
Groves EW, Moore DT (1989) A list of the cryptogams and gymnospermous
plant specimens in the British Museum (Natural History) gathered by
Robert Brown in Australia 1801–5. Proceedings of the Linnean Society of
New South Wales 111, 65–102.
Imahori K (1954) ‘Ecology, Phytogeography and Taxonomy of the Japanese
Charophyta.’ (Kanazawa University: Kanazawa, Japan)
Khatun W, Ud-Deen MM, Kabir G (2009) Karyotype of some species of
Chara and Nitella from Bangladesh. Bangladesh Journal of Botany 38,
205–207.
Kützing FT (1843) ‘Phycologia generalis, oder Anatomie, Physiologie und
Systemkunde der Tange. Vol. 2’. (F.A. Brockhaus: Leipzig)
37
Kützing FT (1857) ‘Tabulae Phycologiae, oder Abbildungen der Tange.
Vol. 7’, pp. 10–32. (Nordhausen)
Macdonald MB, Hotchkiss AT (1956) An estipulodic form of Chara australis
R.Br. (= Protochara australis Woms. and Ophel). Proceedings of the
Linnean Society of New South Wales 80, 274–284.
Mandal DK, Ojha SN, Ray S (2008) Species-specificity of oospore
dimensions in three ecorticate species of Chara (Charales,
Charophyceae). Aquatic Botany 88, 181–183.
doi:10.1016/j.aquabot.2007.09.006
McCourt RM, Karol KG, Proctor V, Feist M, Delwiche CF (2000) Molecular
phylogeny of tribe Charae (Characeae) based on rbcL sequences, with
comments on classification of Chara sensu Wood. In ‘The Third
International Symposium on Extant and Fossil Charophytes’, October
2000, Nanjing, Peoples Republic of China. (Eds W Qifei, L Hui-nan, I
Soulié-Märche) p. 29. (Nanjing Institute of Geology and Palaeontology,
Academia Sinica: Nanjin, China)
McNeill J, Barrie FR, Buck WR, Demoulin V, Greuter W, Hawksworth D,
Herendeen PS, Knapp S, Marhold K, Prado J, Prud’homme van Reine WF,
Smith GF, Wiersema JH, Turland NJ (Eds) (2012) International
Code of Nomenclature for algae, fungi and plants (Melbourne Code),
adopted by the Eighteenth International Botanical Congress Melbourne,
Australia, July 2011. (International Association for Plant Taxonomy:
Bratislava, Slovakia) Available at http://www.iatp-taxon.org/nomen/
main.php [Verified September 2013]
Meiers ST, Proctor VW, Chapman RL (1999) Phylogeny and biogeography
of Chara (Charophyta) inferred from 18S rDNA sequences. Australian
Journal of Botany 47, 347–360. doi:10.1071/BT97089
Nordstedt CFO (1883) Fragmente einer Monographie der Characeen.
Abhandlungen der Königlichen Akademie der Wissenschaften in Berlin
1882, 1–211.
Orchard AE (1999) A history of systematic botany in Australia. In ‘Flora of
Australia. Vol. 1’. 2nd edn. (Ed. AE Orchard) pp. 11–103. (ABRS:
Canberra)
Proctor VW (1980) Historical biogeography of Chara (Charophyta): an
appraisal of the Braun–Wood classification plus a falsifiable alternative
for future consideration. Journal of Phycology 16, 218–233.
doi:10.1111/j.1529-8817.1980.tb03023.x
Proctor VW (1999) Charophytivory, Playas y Papalotes, a local paradigm of
global relevance. Australian Journal of Botany 47, 399–406.
doi:10.1071/BT97088
Willdenow CL (1805) Du genere nommé Chara. Mémoires de l’Académie
Royale des Sciences 1803, 79–90.
Womersley HBS, Ophel IL (1947) Protochara, a new genus of Characeae
from Western Australia. Transactions of the Royal Society of South
Australia 71, 311–317.
Wood RD (1962) New combinations and taxa in the revision of the
Characeae. Taxon 11, 7–25. doi:10.2307/1216853
Wood RD (1971) Characeae of Australia. Nova Hedwigia 22, 1–120.
Wood RD, Imahori KI (1964) Iconograph of the Characeae. In ‘A Revision of
the Characeae. Vol. 2’. (Eds RD Wood, KI Imahori) pp. 1–395. (Cramer:
Weinheim, Germany)
Wood RD, Imahori KI (1965) Monograph of the Characeae. In ‘A Revision of
the Characeae. Vol. 1’. (Eds RD Wood, KI Imahori) pp. 1–904. (Cramer:
Weinheim, Germany)
Zaneveld JS (1940) The Charophyta of Malaysia and adjacent countries.
Blumea 4, 1–223.
www.publish.csiro.au/journals/asb