Phytotaxa 275 (3): 243–262
http://www.mapress.com/j/pt/
Copyright © 2016 Magnolia Press
ISSN 1179-3155 (print edition)
Article
PHYTOTAXA
ISSN 1179-3163 (online edition)
http://dx.doi.org/10.11646/phytotaxa.275.3.3
Morphometric analysis reveals a new species of Aponogeton (Aponogetonaceae) in
Sri Lanka
CHAPA G. MANAWADUGE1, DEEPTHI YAKANDAWALA1* & DONALD H. LES2
1
Department of Botany, University of Peradeniya, Peradeniya, Sri Lanka
Department of Ecology & Evolutionary Biology, University of Connecticut, Storrs, CT, USA
*
Corresponding author: deepthiy@pdn.ac.lk, deepthiyakandawala@gmail.com
2
Abstract
A morphometric analysis of Sri Lankan Aponogeton was performed in order to clarify species delimitations and to facilitate
their identifications by comparing states coded for 61 vegetative and reproductive characters. Both cluster analysis and
principal coordinate analysis resolved five discrete phenetic groups within the genus in the island. Four of these clusters corresponded well with the four previously reported species (A. rigidifolius, A. jacobsenii, A. crispus and A. natans) whereas a
fifth cluster delimited a new Aponogeton species, A. dassanayakei, from the wet lowland regions of Sri Lanka. These analyses have disclosed more useful characters for field identification of the species.
Key Words: Aponogeton dassanayakei, Aquatic plants, Cluster analysis, Morphology, Phenetics
Introduction
Despite its small size (65,610 km2), Sri Lanka contains a rich and diverse flora. The island is known to harbor 3,154
flowering plant species (MOE 2012) but potentially contains many that remain undescribed. About 370 species
(<12% of total) represent aquatic or wetland plants, wherein 205 (6% of total) are monocots, which include the genus
Aponogeton Linnaeus f. (1781[1782]: 32) (Yakandawala 2012).
All species of Aponogeton (Alismatales: Aponogetonaceae) are obligate, aquatic (freshwater) plants, which are
distributed mainly in the tropical or subtropical regions of the Old World (Les et al. 2005). In Sri Lanka, the genus is
represented by four indigenous species; Aponogeton rigidifolius Bruggen (1962: 91), A. jacobsenii Bruggen (1983:
120), A. crispus Thunberg (1784: 73), and A. natans (Linnaeus 1767: 227) Engler & Krause (1906: 11). Of these, the
former two are both endemic and endangered (Bruggen 1987, Yakandawala 2012). All of the species are known by the
vernacular name “Kekatiya” (in Sinhala) and are popular aquarium plants.
Since the genus Aponogeton was first erected by Linnaeus fil. in 1781 (Yadav & Gaikwad 2003), a comprehensive
taxonomic treatment of the group was wanting for more than two centuries until Bruggen (1985) recognized 43 species
in his pioneering monograph. Subsequent workers added additional taxa and about 58 species are presently recorded
worldwide (Guillarmod 1978, Sundararaghavan et al. 1982, Hellquist & Jacobs 1998, Yadav & Gaikwad 2003,
Jacobs et al. 2006, Ali 2008, Chen et al. 2015, Yadav et al. 2015). The only major taxonomic study to include Sri
Lankan Aponogeton was that of Bruggen (1985 &1987) and to date there has not been a major revision of the genus.
During the course of recent fieldwork on the aquatic flora of Sri Lanka, a number of Aponogeton populations were
encountered, which contained individuals that were difficult to identify because of inconsistencies with the patterns
of morphological variation described in prior taxonomic studies. In an effort to better understand the taxonomy of Sri
Lankan Aponogeton, a morphometric analysis was conducted from material collected throughout the island as a means
of elucidating species delimitations and to develop a more competent approach for their identification.
Accepted by Samuli Lehtonen: 26 Aug. 2016; published: 21 Sept. 2016
243
Materials and Methods
Sample collection
Seventy Aponogeton plants from 28 Sri Lankan populations (collected from all suitable habitats located throughout
the island) comprised the operational taxonomic units (OTUs) included in this study; each population was denoted by
a unique acronym to facilitate its reference (Fig. 1, Table 1). These collections included material assigned to all of the
currently recognized Aponogeton species on the island as well as providing a comprehensive sample of morphological
variation with respect to both their flowering and fruiting stages. Representative samples of each population also were
grown and maintained at the Department of Botany, University of Peradeniya, Sri Lanka to provide living reference
material for consultation during the course of the study. Voucher specimens for all collections were deposited at The
National Herbarium, Peradeniya, Sri Lanka (Appendix 1)
TABLE 1. Field localities and acronym assigned to Sri Lankan populations sampled in this study.
Locality
Acronym
Locality
Acronym
Naula (Matale)
A1–A2
Wanduramba (Galle)
O43–O44
Pidurangala (Matale)
B3–B4
Ittapane (Kalutara)
P45–P46
Pidurangala (Matale)
C5–C6
Lunuganga (Kalutara)
Q47–Q48
Batticaloa (Batticaloa)
D7–D9
Tirappane (Anuradhapura)
R49–R50
Hanwella (Colombo)
E10–E11
Kebithigollewa(Anuradhapura)
S51–S52
Kottawa (Galle)
F12–F14
Tittagonewewa(Anuradhapura)
T53–T54
Kanneliya (Galle)
G15–G17
Muliyarkulam (Vavunia)
U55–U56
Meegama (Kalutara)
H1–H20
Murunkan (Mannar)
V57–V58
Lunuganga (Kalutara)
I21–I23
Murunkan (Mannar)
W59–W60
Delgoda (Kalutara)
J24–J26
Horton plains (N’ Eliya)
X61–X62
Royal Botanical Garden
K27
Ganemulla (Gampaha)
Y63–Y64
Horton plains (N’ Eliya)
L28–L30
Yagoda (Gampaha)
Z65–Z66
Somawathiya (Polonnaruwa)
M31–M40
Batticaloa (Batticaloa)
AA67–AA68
Sinharaja (Galle)
N41–N42
Kuruwita (Ratnapura)
AB69–AB70
Morphometric Analysis
Sixty-one morphological characters, representing 29 vegetative and 32 reproductive features, including those
emphasized as important taxonomically by Bruggen (1985 &1987), were scored for analysis (Table 2) (Appendix 2).
Characters were scored from 5–10 individuals in each population with five measurements taken per character in the
case of quantitative characters; in these instances, the mean values represented the character states evaluated.
Although scoring for most of the qualitative characters is self-explanatory, two are clarified further. For floret
spacing (i.e., the space between florets in the inflorescence), the character states were defined as: ‘loose’ = >5 mm;
‘slightly dense’ = <5 mm but >0 mm; or ‘dense’ = 0 mm. For stamen filament shape: ‘linear’= filament width equal
from top to bottom; ‘slightly dilated’ = filament width uniform in upper half, broadened slightly in lower half; and
‘broadly dilated’ = filament width increasing from top to bottom into a broadly expanded base. All characters and their
corresponding character states were assembled into a data matrix, which was analyzed (as follows) using version 2.15
of the PAST software program (Hammer et al. 2001).
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MANAWADUGE ET AL.
FIGURE 1. Map of Sri Lanka indicating the sample collection locations (N 5º55’–9º55’ / E 79º42’–81º52’).
Two analyses were performed using PAST. First, the Gower similarity measure option was selected to generate
a range-normalized Manhattan distance matrix. We input this matrix into a hierarchical cluster analysis (CA) by
NEW SPECIES OF APonogETon
Phytotaxa 275 (3) © 2016 Magnolia Press • 245
selecting the ‘paired group’ (UPGMA) option to depict the phenetic relationships among all OTUs as a dendrogram.
We then performed an ordination analysis using the Gower measure option (transformation exponent C=2) to generate
a distance matrix for use in a principal coordinates analysis (Davis 1986). Following the results of these analyses,
each major, consistently recovered cluster was identified and where possible, associated with a specific taxon using
taxonomic literature references (Bruggen 1985 & 1987).
TABLE 2. Morphological characters and character states evaluated in the study (sub = submersed; flo = floating; NA = not applicable).
Vegetative characters:
1. sub leaves (absent=0, present=1); 2. flo leaves (absent=0, present=1); 3. flo leaf habit (NA=0, never erect=1, sometimes
erect=2); 4. root stock (rhizome=1, tuber=2); 5. tuber shape (NA=0, globular=1, cylindrical=2); 6. root stock length (cm); 7. root stock
width (cm); 8. sub petiole length (cm); 9. sub petiole width (cm); 10. sub petiole shape (triangular=0, grooved=1 , NA= 2 ); 11. sub
leaf length (cm); 12. sub leaf width (cm); 13. sub leaf margin (flat=0, undulate=1, undulate and/or flat=2, NA=3); 14. undulation type
(type 1=0, type 2=1, type 3=2, type 4=3, NA=4); 15. flo leaves twisted around midrib (no=0, yes=1, NA=2); 16. sub leaf tip (narrowly
acute=0, acute =1, retuse=2, NA=3); 17. sub leaf base (acute=0, cordate=1,NA=2); 18. sub leaf texture (type 1=0, type 2=1, type 3=2,
NA=3); 19. sub leaf primary vein number (#); 20. flo leaf petiole length (cm); 21. flo leaf petiole width (cm); 22. flo leaf petiole cross
section (triangular=0, V-shaped=1, NA=2); 23. flo leaf length (cm); 24. flo leaf width (cm); 25. flo leaf margin (flat =0, undulate=1,
NA=2); 26. flo leaf base (cordate type 1=0, cordate type 2=1, NA=2); 27. flo leaf texture (type 4=0, type 5=1, NA=2); 28. flo leaf
primary vein number (#); 29. flo leaf midrib (flat=0, protruding=1, grooved=2, NA=3).
Reproductive characters:
30. peduncle (proliferous=0, not proliferous=1); 31. peduncle shape (widened apically=0, width uniform=1); 32. peduncle length
(cm); 33. spathe duration (caducous=0, persistent=1); 34. inflorescence length (cm); 35. inflorescence width (mm); 36. floret spacing
(loose=0, slightly dense=1, dense=2); 37. tepal number (#); 38. tepal colour (white=0, pinkish white=1, purple=2); 39. tepal length
(mm); 40. tepal width (mm); 41. tepal length: width (ratio); 42. nerve number (#); 43. stamen number (#); 44. filament shape (linear
=0, slightly dilated=1, broadly dilated=2); 45. stamen length (mm); 46. anther colour (light purple=0, dark purple=1, grayish green=2,
brown=3); 47. carpel number (#); 48. gynoecium length (mm); 49. gynoecium width (mm); 50. mature gynoecium colour (white=0,
light pink=1, dark pink=2, purple=3); 51. ovules per carpel (#); 52. fruit texture (rough=0, smooth=1); 53. fruit shape (elongate=0,
broadly triangular =1, globular=2); 54. testa number (#); 55. outer-testa thickness (thick=0, thin=1) ; 56. outer-testa texture (smooth=0,
ridged=1); 57. inner-testa (absent=0, present=1); 58. seed length (mm); 59. seed width (mm); 60. plumule (absent=0, present=1); 61.
embryo attachment (at base=0, at or above middle=1, NA=2).
Results
The UPGMA dendrogram (cophenetic correlation coefficient=0.9848) resolved five discrete clusters of OTUs (hereafter
clusters 1–5), which separated respectively at approximately 0.16, 0.20, 0.30, and 0.53 distance units (Fig. 2). The
OTUs within each cluster grouped together closely, with none of them exceeding a distance of more than 0.07 units
within any given cluster (Fig. 2). The first four (principal) eigen values recovered from the PCoA (3.9149, 0.6801,
0.1606, and 0.1180) accounted for 91% of the total variance (73%, 13%, 3%, and 2% respectively). A plot of the first
and third coordinates (which provided the greatest separation of OTUs) returned a result similar to that obtained by the
CA. Here the PCoA also resolved five discrete clusters (Fig. 3), with each corresponding exactly to one of the clusters
indicated by the UPGMA dendrogram (Fig. 2). Although cluster 1 overlapped with cluster 2 along the first coordinate
axis, there was no overlap between any of clusters 2–5; however, cluster 1 separated widely from cluster 2 and all other
clusters along coordinate 3 (Fig. 3).
Taxonomic descriptions (Bruggen 1985 & 1987) enabled us to explicitly associate three Aponogeton species with
three of the consistently recovered clusters: A. crispus (cluster 4), A. jacobsenii (cluster 3), and A. rigidifolius (cluster
1). Cluster 2 contained OTUs similar to A. rigidifolius, and the most distant cluster (cluster 5) did not precisely match
existing descriptions for A. natans, a fourth species attributed to Sri Lanka.
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MANAWADUGE ET AL.
FIGURE 2. A dendrogram resulting from CA resolves five distinct groups of OTUs among the Aponogeton material evaluated in this
study.
NEW SPECIES OF APonogETon
Phytotaxa 275 (3) © 2016 Magnolia Press • 247
FIGURE 3. Scatter plot resulting from PCoA showing the five distinct clusters of Aponogeton OTUs generated from the study material
evaluated.
Discussion
Results of the CA and PCoA analyses consistently indicated five discrete phenetic groups of Aponogeton OTUs in the
Sri Lankan flora. Three of these clusters (1, 3 and 4) could be assigned readily to three species attributed previously to
Sri Lanka, namely A. rigidifolius, A. jacobsenii, and A. crispus respectively.
The OTUs associated with cluster 5 did not completely agree with the morphological descriptions of A. natans,
which has been described and illustrated as a plant having both floating and submersed leaves (Bruggen 1985 & 1987,
Cook 1996, Yadav & Gaikwad 2003, Wijesundara & Shantha siri 2004, Ali 2008). None of the OTUs in cluster 5
possessed submersed leaves but they were all erect or emergent, which apparently developed into floating leaves when
the plants became inundated during the rainy seasons. We have recently clarified this discrepancy by demonstrating
that A. natans actually is a species with no submersed leaves (Manawaduage et al. 2016).
The identity of cluster 2 OTUs represented a more difficult challenge, given that all of the known Sri Lankan
Aponogeton species already had been accounted for by OTUs in clusters 1, 3, 4 and 5. The close phenetic association
of cluster 1 and 2 OTUs by morphometric analysis (Fig. 2; 0.16 distance units) closely grouped the latter cluster
with A. rigidifolius. Comparatively, the OTUs in both clusters were more similar morphologically to each other than
to any of the OTUs assigned to the other species. By their fully submerged, undulate leaves and emergent, whiteflowered inflorescences, one might readily assign cluster 2 OTUs to A. rigidifolius. Yet upon closer examination, the
OTUs in cluster 2 differed from A. rigidifolius by the presence of a tuber instead of a rhizome, and by their longer,
highly undulated, and brownish green coloured leaves as opposed to the shorter, less undulated or flattened, and dark
purplish-green leaves of A. rigidifolius. These differences provide practical characters for separating the two taxa in
the field.
Although this undescribed species bears some resemblance to A. rigidifolius, it also displays a number of
differences. The endemic A. rigidifolius is confined to few locations in the low land wet zone of Sri Lanka. Although
cluster 2 OTUs also occur in the low land wet zone, they exhibit a wider distribution. These taxa also differ ecologically,
with A. rigidifolius occurring in stony, rapidly flowing fresh water streams, while the members of cluster 2 OTUs occur
in slow flowing rivulets or rivers in much deeper water (below 1–2 m in some cases). The cluster 2 plants also occur
nearer to the lagoons, where we have never encountered A. rigidifolius.
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FIGURE 4. Proliferous peduncle of Aponogeton dassanayakei Manawaduge & Yakandawala (specimen collected on 11/08/2014 from
Lunuganga (Kalutara), GPS location: N 06° 24. 254 ′ / E 80° 01.121′).
NEW SPECIES OF APonogETon
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Further, we observed that some cluster 2 OTUs uniquely possessed a proliferous peduncle (Fig. 4), which is a
character that never had been recorded previously for any Sri Lankan Aponogeton, including A. rigidifolius. In cluster 2
plants, vegetative propagation can occur by the formation of young plantlets/propagules (rather than an inflorescence),
which develop at the tip of a peduncle-like axis. Further, the proliferous peduncle character was always associated with
individuals growing in close proximity to more saline waters. However, none of the plants with proliferous peduncles
considered during the study bore an inflorescence on the same individual.
Proliferous peduncles occur rarely in Aponogeton and have been reported previously in only two species: A.
proliferous C.B. Hellquist & S.W.L. Jacobs (1998: 11), an Australian species with yellow-flowered inflorescences and
A. undulatus Roxburgh (1832: 211), an Asian species with white-flowered inflorescences (Bruggen 1985, Hellquist &
Jacobs 1998, Les et al. 2005). Of these, A. proliferous is distinct from cluster 2 OTUs by its yellow tepals. In a nontaxonomic study, Thabrew & Thabrew (1983) recorded “A. undulatus” from Sri Lanka, but provided no distinguishing
features for its identification. Morphologically, A. undulatus appears to be much closer to members of cluster 2 than to
cluster 1. However, unlike the strictly submersed foliage of cluster 2 OTUs, A. undulatus possesses both floating and
submerged leaves (Bruggen 1985, Yadav & Gaikwad 2003). Cluster 2 OTUs also possess much longer, finer, and more
heavily undulated submerged leaves, which (unlike A. undulatus) sometimes twist around their mid rib (often in two to
three spirals). Their submerged leaves also lack the alternating transparent and opaque areas that are characteristic of
A. undulatus. Their caducous spathe also contrasts with the persistent spathe ascribed to A. undulatus (Bruggen 1985,
Yadav & Gaikwad 2003). The tepals of A. undulatus are elongate; whereas, those of cluster 2 OUTs are more spherical
(Bruggen 1985, Yadav & Gaikwad 2003), and are considerably wider than long. Further, the tepals are persistent and
remain on the infructescence below the developing ovaries; whereas, the tepals of A. undulatus are caducous (Bruggen
1985, Yadav & Gaikwad 2003) (Fig. 5). From our observations, all of these characters are stable and are not altered
substantially by environmental factors.
FIGURE 5. Persistent tepals in a mature inflorescence from Aponogeton dassanayakei Manawaduge & Yakandawala (specimen collected
on 11/08/2014 from Lunuganga (Kalutara), GPS location: N 06° 24. 254 ′ / E 80° 01.121′).
Thus, the particular combination of traits specific to cluster 2 OTUs is not found in any previously described
Aponogeton species. For this reason we conclude that these OTUs represent a new, undescribed Aponogeton species.
A formal description with the illustration (Fig. 6) to the new species is given below. In summary, our morphometric
analysis of Sri Lankan Aponogeton supports the recognition of five distinct species in the country, which include
four previously reported species along with a novel species. Combined with field observations, these analyses also
have disclosed errors in the morphological description of A. natans, to which submersed leaves have been associated
erroneously. Further studies of Sri Lankan Aponogeton using molecular data currently are underway and hopefully will
shed additional light on the diversity, origins and relationships of the genus within this small island country.
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MANAWADUGE ET AL.
Taxonomy
Aponogeton dassanayakei Manawaduge & Yakandawala, sp. nov. (Fig. 6)
Aponogeton dassanayakei is distinct from the other two proliferous species known in the genus by having the combination of characters;
highly undulated long strictly submersed leaves with no alternately transparent or opaque patterns, broad white coloured persistent
tepals and caducous spathe.
Type:—SRI LANKA. Western Province: Kalutara, Ittapana Thotupola-South End, (Ittapane–Horawala Thotupola Road), elev. 3 m, 08°
23.910′ N / 80° 4.067′ E, 15 February 2015, Manawaduge & Yakandawala 215. (Holotype: PDN!).
Tuber cylindrical and elongate; 9–20 mm Ø. Submerged leaves lamina; 17–56 × 1.4–5.2 cm; linear and slightly
curved, apex acute and base cuneate, margin finely undulate, midrib with 6–8 parallel nerves; Petioles up to 56 cm
long, brittle, bluntly triangular. Floating leaves absent. Peduncle up to 178 cm, widening towards the inflorescence.
Spathe up to 35 mm, caduceus. Inflorescence with 1 spike; spike up to 15 cm, densely flowered; flowers turned
towards all directions. Tepals 2, 1.2–2.3 × 1.3–2.5 mm, persistent, 1 nerved, white, broad. Stamens 6, 1–2 mm;
filaments slightly widening towards the base, white, anthers grayish green. Carpels 3, white, 1.0–1.5 × 0.4–1 mm;
ovules 2. Infructescence up to 31 cm. Folicals up to 10 × 6 mm, elongated, triangular, with rough coat and a short,
terminal, curved beak. Seeds with a simple testa; embryo up to 8–9 × 3–6 mm, plumule attached near the base of the
embryo and in a very wide groove. Proliferous peduncles ascending up to 45 cm, slightly thickened towards the tip.
FIGURE 6. Aponogeton dassanayakei Manawaduge & Yakandawala (a) habit, (b) inflorescence, (c) floret, (d) tepal, (e) stamen, (f)
gynoecium and (g) infructescence.
Phenology:—Throughout the year.
Distribution:—Aponogeton dassanayakei occurs in the low land wet zone, in slow flowing rivulets and rivers in
much deeper water (below 1–2 m in some cases) and also in saline water towards the lagoons.
Eponymy:—The new species has been named after Professor M. D. Dassanayake, an eminent Plant Taxonomist
in Sri Lanka and the General Editor of “A Revised Handbook to the Flora of Ceylon” (Vols. I–XV).
NEW SPECIES OF APonogETon
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Additional specimens examined (Paratypes):—SRI LANKA. Western Province: Colombo, Hanwella, 06°54.109′
N, 80°05.613′ E, 13 m, 28 July 2014, Manawaduge & Yakandawala 105 (PDN!); Kalutara, Meegama, near Meegama
Thotupola bridge, 06°24.825′N, 80°04.005′ E, 4 m , 11 August 2014, Manawaduge & Yakandawala 408, 409 (PDN!);
Lunuganga, Bawa Gardens, 06°24. 254′N, 80°01.121′ E, 18 m, 11 August 2014, Manawaduge & Yakandawala 109
(PDN!); same locality, 06°24.30′N, 80° 1.211′E, 18 m, 15 February 2015, Manawaduge & Yakandawala 117 (PDN!);
Bandaragama,Waskaduwa—Bandaragama road, 06°41.155 ′ N 79° 58. 857 ′E, 2 m, 11August 2014, Manawaduge &
Yakandawala 110 (PDN!); Gampaha, Kirindiwita, Kirindiwita–Ganemulla road, 07° 5.370′N 79°58.156′E, 10 m, 24
September 2015, Manawaduge & Yakandawala 126 (PDN!); Yagoda, 07°4.390′N, 79°58.188′E, 10 m, 24 September
2015, Manawaduge & Yakandawala 127 (PDN!); Sabaragamuwa Province: Ratnapura, Kuruwuta, Colombo–Batticaloa
highway, 06°46.982′N, 80°21.579′E, 27 m, 17 December 2015, Manawaduge & Yakandawala 128 (PDN!).
Acknowledgements
Financial assistance provided by the University of Peradeniya, Sri Lanka (University Research Grant RG/2014/38/S)
to DY is gratefully acknowledged. Authors wish to thank The Department of Wildlife Conservation of Sri Lanka for
granting permission for collecting of material, National Herbarium, Royal Botanical Gardens, Peradeniya, Sri Lanka
and Menaka Ariyarathne and U. B. Priyadharshana for their assistance in the field.
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Wijesundara, D.S.A. & Shantha Siri, J.G. (2004) Some selected aquatic ornamental plants of Sri Lanka: a simplified guide to identification
of Aponogeton (Kekatiya), Lagenandra (Ketala) and Cryptocoryne (Atiudayan). Technology Watch Centre, National Science
Foundation of Sri Lanka, Colombo, Sri Lanka, pp. 4–14.
Wit, H.C.D. de (1983) Aquariumplanten, ed. 4. Stuttgart, Germany, 120 pp.
Yadav, S.R. & Gaikwad, S.P. (2003) A revision of the Indian Aponogetonaceae. Bulletin of the Botanical Survey of India 45: 39–76.
Yadav, S.R., Patil, V.S., Gholave, A.R., Chandore, A.N., Yadav, U.S. & Kambale, S.S. (2015) Aponogeton nateshii (Aponogetonaceae): a
new species from India. Rheedea 25: 9–13.
Yakandawala, D. (2012) Present status of fresh water aquatic flora in Sri Lanka. In: Weerakoon, D.K. & Wijesundara, S. (Eds.) The
national Red List 2012 of Sri Lanka; Conservation Status of the Fauna and Flora. Ministry of Environment, Colombo, Sri Lanka,
pp. 186–196.
NEW SPECIES OF APonogETon
Phytotaxa 275 (3) © 2016 Magnolia Press • 253
APPENDIX 1. List of Aponogeton taxa investigated in the study, their collection dates, localities, and voucher specimen information (all
specimens deposited at The National Herbarium, Peradeniya, Sri Lanka).
Species
A. crispus
A. natans
Collection Date
Province
Locality
Voucher Specimen
25/05/2014
Central
Naula (Matale)
Manawaduge & Yakandawala 101
25/05/2014
Central
Pidurangala (Matale)
Manawaduge & Yakandawala 102
25/05/2014
Central
Pidurangala (Matale)
Manawaduge & Yakandawala 103
26/09/2014
Central
RBG (Peradeniya)
Manawaduge & Yakandawala 111
13/06/2015
North Central
Tirappane (Anuradhapura)
Manawaduge & Yakandawala 118
13/06/2015
North Central
Kebithigollewa (Anuradhapura)
Manawaduge & Yakandawala 119
13/06/2015
North Central
TittagoneWewa (Anuradhapura)
Manawaduge & Yakandawala 120
25/05/2014
Eastern
Batticaloa (Batticaloa)
Manawaduge & Yakandawala
704,705,706
25/10/2014
North Central
Somawathiya (Polonnaruwa)
14/06/2015
Northern
Muliyarkulam (Vavunia)
14/06/2015
Northern
Murunkan (Mannar)
14/06/2015
Northern
Murunkan (Mannar)
06/12/2015
Eastern
Batticaloa (Batticaloa)
28/07/2014
Southern
Kottawa (Galle)
28/07/2014
Southern
Kanneliya (Galle)
07/0/2015
southern
Sinharaja (Galle)
15/02/2015
Southern
Wanduramba (Galle)
13/10/2014
Central
Horton plains (N’ Eliya)
23/07/2015
Central
Horton plains (N’ Eliya)
28/07/2014
Western
Hanwella (Colombo)
11/08/2014
Western
Meegama (Kalutara)
11/08/2014
Western
Lunuganga (Kalutara)
11/08/2014
Western
Delgoda (Kalutara)
15/02/2015
Western
Ittapane (Kalutara)
15/02/2015
Western
Lunuganga (Kalutara)
Manawaduge & Yakandawala
215,216,217
24/09/2015
Western
Ganemulla (Gampaha)
Manawaduge &Yakandawala 117
24/09/2015
Western
Yagoda (Gampaha)
Manawaduge & Yakandawala 126
17/12/2015
Sabaragamuwa
Kuruwita (Ratnapura)
Manawaduge & Yakandawala 127
Manawaduge & Yakandawala 113
Manawaduge & Yakandawala 121
Manawaduge & Yakandawala 122
Manawaduge & Yakandawala 123
Manawaduge & Yakandawala 627
A. rigidifolius
Manawaduge & Yakandawala 506,507
Manawaduge & Yakandawala 107
Manawaduge & Yakandawala 114
Manawaduge & Yakandawala 315,316
A. jacobsenii
Manawaduge & Yakandawala 112
Manawaduge & Yakandawala 124
A. dassanayakei
Manawaduge & Yakandawala 105
Manawaduge & Yakandawala 408,409
Manawaduge & Yakandawala 109
Manawaduge & Yakandawala 110
Manawaduge & Yakandawala 128
254 • Phytotaxa 275 (3) © 2016 Magnolia Press
MANAWADUGE ET AL.
NEW SPECIES OF APonogETon
APPENDIX 2. Morphological data matrix used in the morphometric analysis. Character numbers correspond to that of Table 2. Acronyms assigned to OTUs correspond to that of Table 1. ? =
missing data.
oOTUs
Character
Phytotaxa 275 (3) © 2016 Magnolia Press • 255
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
A1
1
1
1
2
1
2.8
1.8
13.0
0.3
0
12.0
3.8
1
0
0
2
1
0
4
63.0
0.2
0
15.0
3.0
0
0
0
4
1
1
A2
1
1
1
2
1
3.5
1.2
22.0
0.5
0
20.0
5.7
1
0
0
2
1
0
4
71.0
0.3
0
17.0
4.0
0
0
0
4
1
1
B3
1
1
1
2
1
1.7
1.0
35.0
0.3
0
11.5
3.0
1
0
0
2
1
0
4
?
?
?
?
?
0
0
0
4
1
1
B4
1
1
1
2
1
1.3
0.7
38.0
0.4
0
18.0
4.0
1
0
0
2
1
0
4
17.0
0.2
0
6.0
2.0
0
0
0
4
1
1
C5
1
1
1
2
1
2.8
1.8
11.0
0.5
0
18.0
5.0
1
0
0
2
1
0
5
32.0
0.3
0
15.0
2.0
0
0
0
4
1
1
C6
1
1
1
2
1
1.0
0.8
9.8
0.6
0
21.0
6.1
1
0
0
2
1
0
5
35.0
0.3
0
18.0
2.0
0
0
0
4
1
1
D7
0
1
2
2
1
2.5
1.5
0.0
0.0
2
0.0
0.0
3
4
2
3
2
3
0
32.0
0.6
1
14.0
3.0
0
1
1
4
2
1
D8
0
1
2
2
1
2.0
1.5
0.0
0.0
2
0.0
0.0
3
4
2
3
2
3
0
33.0
0.6
1
13.0
3.0
0
1
1
4
2
1
D9
0
1
2
2
1
1.5
1.0
0.0
0.0
2
0.0
0.0
3
4
2
3
2
3
0
26.0
0.5
1
11.0
3.0
0
1
1
4
2
1
E10
1
0
0
2
2
4.0
0.9
23.0
0.4
0
31.0
1.4
1
3
1
0
0
0
4
0.0
0.0
2
0.0
0.0
2
3
3
0
3
1
E11
1
0
0
2
2
10.0
1.2
25.0
0.5
0
43.0
2.4
1
3
1
0
0
0
4
0.0
0.0
2
0.0
0.0
2
3
3
0
3
1
F12
1
0
0
1
0
6.0
0.8
56.0
0.3
0
59.0
4.1
2
2
0
0
0
2
5
0.0
0.0
2
0.0
0.0
2
3
3
0
3
1
F13
1
0
0
1
0
6.0
0.8
42.0
0.4
0
51.0
5.0
2
2
0
0
0
2
5
0.0
0.0
2
0.0
0.0
2
3
3
0
3
1
F14
1
0
0
1
0
4.0
0.3
13.0
0.3
0
28.0
3.0
2
2
0
0
0
2
5
0.0
0.0
2
0.0
0.0
2
3
3
0
3
1
G15
1
0
0
1
0
10.0
0.8
18.0
0.4
0
33.0
2.0
2
2
0
0
0
2
4
0.0
0.0
2
0.0
0.0
2
3
3
0
3
1
G16
1
0
0
1
0
5.0
0.6
11.0
0.3
0
19.0
3.0
2
2
0
0
0
2
4
0.0
0.0
2
0.0
0.0
2
3
3
0
3
1
...Continued on next page
256 • Phytotaxa 275 (3) © 2016 Magnolia Press
APPENDIX 2. (Continued)
oOTUs
Character
MANAWADUGE ET AL.
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
G17
1
0
0
1
0
6.0
0.8
16.0
0.3
0
30.0
3.0
2
2
0
0
0
2
4
0.0
0.0
2
0.0
0.0
2
3
3
0
3
1
H18
1
0
0
2
2
6.0
2.0
20.0
0.6
0
43.0
3.7
1
3
1
0
0
0
3
0.0
5.0
2
0.0
0.0
2
3
3
0
3
0
H19
1
0
0
2
2
6.0
2.0
32.0
0.8
0
56.0
4.9
1
3
1
0
0
0
4
0.0
0.0
2
0.0
0.0
2
3
3
0
3
0
H20
1
0
0
2
2
7.0
2.0
21.0
0.6
0
55.0
5.2
1
3
1
0
0
0
4
0.0
0.0
2
0.0
0.0
2
3
3
0
3
0
I21
1
0
0
2
2
4.0
2.0
13.0
0.5
0
25.0
3.2
1
3
1
0
0
0
3
0.0
0.0
2
0.0
0.0
2
3
3
0
3
0
I22
1
0
0
2
2
3.0
1.0
8.0
0.4
0
17.0
2.5
1
3
1
0
0
0
3
0.0
0.0
2
0.0
0.0
2
3
3
0
3
0
I23
1
0
0
2
2
5.0
2.0
15.0
0.7
0
28.0
4.6
1
3
1
0
0
0
4
0.0
0.0
2
0.0
0.0
2
3
3
0
3
0
J24
1
0
0
2
2
2.0
2.0
50.0
0.5
0
51.0
4.7
1
3
1
0
0
0
3
0.0
0.0
2
0.0
0.0
2
3
3
0
3
1
J25
1
0
0
2
2
3.0
1.0
40.0
0.7
0
59.0
4.5
1
3
1
0
0
0
4
0.0
0.0
2
0.0
0.0
2
3
3
0
3
1
J26
1
0
0
2
2
3.0
2.0
25.0
0.4
0
39.0
2.5
1
3
1
0
0
0
4
0.0
0.0
2
0.0
0.0
2
3
3
0
3
0
K27
1
1
1
2
1
?
?
33.3
0.5
0
25.0
9.0
1
0
0
2
1
0
4
71.0
0.3
0
13.0
3.8
0
0
0
5
1
1
L28
1
0
0
2
2
5.0
2.0
18.0
0.5
0
19.0
7.0
1
1
0
1
1
1
4
0.0
0.0
2
0.0
0.0
2
3
3
0
3
1
L29
1
0
0
2
2
4.0
1.9
10.0
0.5
0
8.0
3.6
1
1
0
1
1
1
4
0.0
0.0
2
0.0
0.0
2
3
3
0
3
1
L30
1
0
0
2
2
5.0
4.0
5.3
0.5
0
7.0
3.1
1
1
0
1
1
1
4
0.0
0.0
2
0.0
0.0
2
3
3
0
3
1
M31
0
1
2
2
1
4.5
0.9
0.0
0.0
2
0.0
0.0
3
4
2
3
2
3
0
4.8
0.2
1
5.0
0.6
0
1
1
4
2
1
M32
0
1
2
2
1
4.0
1.5
0.0
0.0
2
0.0
0.0
3
4
2
3
2
3
0
4.0
0.2
1
4.0
0.5
0
1
1
4
2
1
M33
0
1
2
2
1
4.8
1.5
0.0
0.0
2
0.0
0.0
3
4
2
3
2
3
0
5.7
0.3
1
5.5
0.5
0
1
1
4
2
1
M34
0
1
2
2
1
1.5
0.8
0.0
0.0
2
0.0
0.0
3
4
2
3
2
3
0
12.3
0.3
1
7.6
0.8
0
1
1
4
2
1
M35
0
1
2
2
1
2.5
1.5
0.0
0.0
2
0.0
0.0
3
4
2
3
2
3
0
12.5
0.2
1
7.5
1.3
0
1
1
4
2
1
...Continued on next page
NEW SPECIES OF APonogETon
APPENDIX 2. (Continued)
oOTUs
Character
Phytotaxa 275 (3) © 2016 Magnolia Press • 257
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
M36
0
1
2
2
1
3.0
1.2
0.0
0.0
2
0.0
0.0
3
4
2
3
2
3
0
12.3
0.2
1
7.3
1.0
0
1
1
4
2
1
M37
0
1
2
2
1
3.5
1.8
0.0
0.0
2
0.0
0.0
3
4
2
3
2
3
0
21.7
0.4
1
9.2
1.5
0
1
1
4
2
1
M38
0
1
2
2
1
4.0
1.2
0.0
0.0
2
0.0
0.0
3
4
2
3
2
3
0
22.0
0.3
1
9.6
1.2
0
1
1
4
2
1
M39
0
1
2
2
1
1.5
0.8
0.0
0.0
2
0.0
0.0
3
4
2
3
2
3
0
28.8
0.3
1
11.2
1.7
0
1
1
4
2
1
M40
0
1
2
2
1
2.0
1.2
0.0
0.0
2
0.0
0.0
3
4
2
3
2
3
0
28.7
0.3
1
10.2
2.0
0
1
1
4
2
1
N41
1
0
0
1
0
14.0
0.8
22.0
0.4
0
38.0
3.1
2
2
0
0
0
2
4
0.0
0.0
2
0.0
0.0
2
3
3
0
3
1
N42
1
0
0
1
0
10.0
0.7
25.0
0.3
0
40.0
3.5
2
2
0
0
0
2
4
0.0
0.0
2
0.0
0.0
2
3
3
0
3
1
O43
1
0
0
1
0
6.0
0.8
15.0
0.3
0
29.0
2.5
2
2
0
0
0
2
4
0.0
0.0
2
0.0
0.0
2
3
3
0
3
1
O44
1
0
0
1
0
8.0
0.6
17.0
0.3
0
32.0
3.0
2
2
0
0
0
2
4
0.0
0.0
2
0.0
0.0
2
3
3
0
3
1
P45
1
0
0
2
2
4.0
1.0
40.0
0.4
0
25.0
4.5
1
3
1
0
0
0
3
0.0
0.0
2
0.0
0.0
2
3
3
0
3
0
P46
1
0
0
2
2
4.0
2.0
35.0
0.6
0
39.0
4.0
1
3
1
0
0
0
4
0.0
0.0
2
0.0
0.0
2
3
3
0
3
0
Q47
1
0
0
2
2
3.0
1.0
31.0
0.5
0
40.0
3.5
1
3
1
0
0
0
3
0.0
0.0
2
0.0
0.0
2
3
3
0
3
0
Q48
1
0
0
2
2
3.0
2.0
25.0
0.5
0
45.0
3.0
1
3
1
0
0
0
4
0.0
0.0
2
0.0
0.0
2
3
3
0
3
0
R49
1
1
1
2
1
1.7
1.5
25.0
0.3
0
26.0
4.5
1
0
0
2
1
0
4
50.0
0.2
0
16.0
3.0
0
0
0
4
1
1
R50
1
1
1
2
1
3.0
2.8
22.0
0.4
0
21.0
4.0
1
0
0
2
1
0
4
45.0
0.3
0
15.0
2.8
0
0
0
4
1
1
S51
1
1
1
2
1
1.3
1.1
11.0
0.5
0
10.0
5.0
1
0
0
2
1
0
4
35.0
0.3
0
14.0
2.9
0
0
0
4
1
1
S52
1
1
1
2
1
2.8
2.5
19.0
0.3
0
15.0
5.7
1
0
0
2
1
0
4
39.0
0.2
0
13.0
2.2
0
0
0
4
1
1
T53
1
1
1
2
1
1.4
1.3
19.0
0.3
0
12.0
3.5
1
0
0
2
1
0
5
28.0
0.2
0
11.0
2.5
0
0
0
4
1
1
T54
1
1
1
2
1
1.7
1.5
21.0
0.4
0
19.0
5.5
1
0
0
2
1
0
5
29.0
0.3
0
10.0
3.0
0
0
0
4
1
1
...Continued on next page
258 • Phytotaxa 275 (3) © 2016 Magnolia Press
APPENDIX 2. (Continued)
oOTUs
Character
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
U55
0
1
2
2
1
3.0
2.3
0.0
0.0
2
0.0
0.0
3
4
2
3
2
3
0
34.0
0.6
1
12.0
3.0
0
1
1
4
2
1
U56
0
1
2
2
1
2.0
1.5
0.0
0.0
2
0.0
0.0
3
4
2
3
2
3
0
25.0
0.6
1
16.0
2.0
0
1
1
4
2
1
V57
0
1
2
2
1
1.9
1.5
0.0
0.0
2
0.0
0.0
3
4
2
3
2
3
0
28.0
0.3
1
15.0
2.0
0
1
1
4
2
1
V58
0
1
2
2
1
2.2
1.8
0.0
0.0
2
0.0
0.0
3
4
2
3
2
3
0
31.0
0.5
1
13.0
3.0
0
1
1
4
2
1
W59
0
1
2
2
1
3.5
2.0
0.0
0.0
2
0.0
0.0
3
4
2
3
2
3
0
34.0
0.6
1
15.0
3.0
0
1
1
4
2
1
W60
0
1
2
2
1
1.8
1.2
0.0
0.0
2
0.0
0.0
3
4
2
3
2
3
0
33.0
0.6
1
10.0
3.0
0
1
1
4
2
1
X61
1
0
0
2
2
5.0
2.0
10.0
0.6
0
15.0
5.0
1
1
0
1
1
1
4
0.0
0.0
2
0.0
0.0
2
3
3
0
3
1
X62
1
0
0
2
2
4.0
3.0
15.0
0.5
0
10.0
6.0
1
1
0
1
1
1
4
0.0
0.0
2
0.0
0.0
2
3
3
0
3
1
Y63
1
0
0
2
2
6.0
2.0
50.0
0.5
0
49.0
3.6
1
3
1
0
0
0
4
0.0
0.0
2
0.0
0.0
2
3
3
0
3
1
Y64
1
0
0
2
2
7.0
1.0
41.0
0.6
0
54.0
2.9
1
3
1
0
0
0
4
0.0
0.0
2
0.0
0.0
2
3
3
0
3
1
Z65
1
0
0
2
2
5.0
2.0
30.0
0.5
0
41.0
3.5
1
3
1
0
0
0
3
0.0
0.0
2
0.0
0.0
2
3
3
0
3
1
Z66
1
0
0
2
2
4.0
1.5
32.0
0.5
0
39.0
3.5
1
3
1
0
0
0
4
0.0
0.0
2
0.0
0.0
2
3
3
0
3
1
AA67
0
1
2
2
1
2.0
1.8
0.0
0.0
2
0.0
0.0
3
4
2
3
2
3
0
28.0
0.4
1
15.0
3.0
0
1
1
4
2
1
AA68
0
1
2
2
1
3.0
1.5
0.0
0.0
2
0.0
0.0
3
4
2
3
2
3
0
27.0
0.5
1
12.0
3.0
0
1
1
4
2
1
AB69
1
0
0
2
2
4.0
2.0
30.0
0.5
0
39.0
3.5
1
3
1
0
0
0
4
0.0
0.0
2
0.0
0.0
2
3
3
0
3
1
AB70
1
0
0
2
2
3.0
1.5
25.0
0.5
0
40.0
3.5
1
3
1
0
0
0
4
0.0
0.0
2
0.0
0.0
2
3
3
0
3
1
MANAWADUGE ET AL.
...Continued on next page
NEW SPECIES OF APonogETon
APPENDIX 2. (Continued)
oOTUs
Character
Phytotaxa 275 (3) © 2016 Magnolia Press • 259
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
A1
0
68
0
17
0.7
2
2
1
1.75
1.55
1.13
1
6
1
1.75
0
3
1.70
1.20
1
2
0
0
1
0
0
0
8.30
3.32
1
0
A2
0
77
0
15
0.6
2
2
1
1.55
1.25
1.24
1
6
1
1.50
0
3
1.70
1.25
1
2
0
0
1
0
0
0
8.72
3.74
1
0
B3
0
59
0
11
0.5
0
2
1
1.25
1.05
1.19
1
6
1
1.30
0
3
1.00
0.50
1
2
0
0
1
0
0
0
7.72
3.40
1
0
B4
0
70
0
10
0.4
0
2
1
1.60
1.50
1.07
1
6
1
1.65
0
3
1.10
0.50
1
2
0
0
1
0
0
0
7.55
3.49
1
0
C5
0
45
0
8
0.5
0
2
1
2.10
2.00
1.05
1
6
1
2.15
0
3
1.50
1.10
1
2
0
0
1
0
0
0
7.89
3.57
1
0
C6
0
40
0
11
0.5
0
2
1
2.00
1.90
1.05
1
6
1
2.05
0
3
1.60
1.20
1
2
0
0
1
0
0
0
7.64
3.49
1
0
D7
1
20
0
10
1.0
2
2
2
3.00
1.90
1.58
1
6
0
4.15
1
3
4.00
1.55
3
8
1
1
2
1
1
1
1.80
0.55
0
2
D8
1
30
0
11
0.8
2
2
2
2.85
1.60
1.78
1
6
0
3.00
1
3
3.60
1.50
3
8
1
1
2
1
1
1
1.60
0.55
0
2
D9
1
15
0
15
0.7
2
2
2
2.90
1.75
1.66
1
6
0
2.60
1
3
3.50
1.50
3
8
1
1
2
1
1
1
1.50
0.50
0
2
E10
0
63
0
16
0.5
0
2
0
1.25
1.50
0.83
1
6
1
1.08
2
3
1.05
0.63
0
2
0
0
1
0
0
1
6.64
2.99
1
0
E11
0
64
0
15
0.4
0
2
0
1.75
1.88
0.93
1
6
1
1.48
2
3
1.45
0.08
0
2
0
0
1
0
0
1
8.00
2.00
1
0
F12
0
40
0
18
0.5
2
2
0
1.50
1.38
1.09
1
6
1
2.00
0
3
1.50
0.08
0
2
0
0
1
0
0
1
10.00
3.00
1
0
F13
0
42
0
11
0.6
2
2
0
2.50
1.38
1.81
1
6
1
2.30
0
3
1.63
0.88
0
2
0
0
1
0
0
1
12.00
4.00
1
0
F14
0
34
0
8
0.5
2
2
0
2.55
1.40
1.82
1
6
1
2.25
0
3
1.70
0.90
0
2
0
0
1
0
0
1
9.00
3.00
1
0
G15
0
70
0
13
0.9
1
2
0
1.65
1.13
1.46
1
6
1
1.75
3
3
2.50
1.00
0
2
0
0
1
0
0
1
12.00
4.00
1
0
G16
0
93
0
43
0.9
1
2
0
1.70
1.20
1.42
1
6
1
1.98
3
3
2.55
1.05
0
2
0
0
1
0
0
1
12.00
4.00
1
0
G17
0
62
0
17
0.7
1
2
0
1.65
1.15
1.43
1
6
1
1.88
3
3
2.50
1.05
0
2
0
0
1
0
0
1
10.00
4.00
1
0
H18
0
178
0
14
0.8
2
2
0
1.33
1.40
0.95
1
6
1
1.50
2
3
1.10
0.63
0
2
0
0
1
0
0
1
9.96
0.87
1
0
...Continued on next page
260 • Phytotaxa 275 (3) © 2016 Magnolia Press
APPENDIX 2. (Continued)
oOTUs
31 32
33 34
H19
0
138 0
17
35
36
37
38
39
40
41
42
43
44
Character
45
46 47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
0
0.8
2
2
0
1.33
1.43
0.93
1
6
1
1.48 2
3
1.15 0.65
0
2
0
0
1
0
0
1
8.50
0.78
1
H20
0
163
0
31
0.5
2
2
0
1.25
1.33
0.94
1
6
1
1.38 2
3
1.08 0.63
0
2
0
0
1
0
0
1
0.95
0.89
1
0
I21
0
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
2
?
?
1
0
?
1
?
?
?
?
I22
0
53
0
6
0.5
2
2
0
1.53
1.88
0.81
1
6
1
1.78 2
3
1.43 0.83
0
2
0
0
1
0
0
1
7.50
0.68
1
0
I23
0
99
1
4
0.5
2
2
0
2.33
2.50
0.93
1
6
1
2.03 2
3
1.50 1.00
0
2
0
0
1
0
0
1
0.83
0.75
1
0
J24
0
136
0
14
0.9
2
2
0
1.13
1.30
0.87
1
6
1
1.23 2
3
0.98 0.38
0
2
0
0
1
0
0
1
7.14
0.65
1
0
J25
0
135
0
27
0.8
2
2
0
1.40
1.45
0.97
1
6
1
1.43 2
3
1.20 0.50
0
2
0
0
1
0
0
1
7.74
0.68
1
0
J26
0
156
0
29
0.8
2
2
0
1.25
1.28
0.98
1
6
1
1.30 2
3
1.00 0.45
0
2
0
0
1
0
0
1
8.50
0.75
1
0
K27
0
80
0
11
0.6
1
2
1
1.60
1.50
1.07
1
6
1
1.00 0
3
1.50 0.80
1
2
0
0
1
0
0
0
8.80
4.00
1
0
L28
1
53
0
11
0.7
2
2
0
1.75
2.50
0.70
1
6
2
1.40 1
3
2.25 1.00
2
2
2
0
1
0
0
1
13.00
11.00
1
1
L29
1
55
0
8
0.6
2
2
0
1.75
2.65
0.66
1
6
2
1.40 1
3
2.15 0.95
2
2
2
0
1
0
0
1
12.00
10.00
1
1
L30
1
50
0
5
8.0
2
2
0
1.70
2.25
0.76
1
6
2
1.35 1
3
2.00 0.85
2
2
2
0
1
0
0
1
10.00
8.00
1
1
M31
1
12.5
0
4
0.5
2
2
2
1.75
1.05
1.67
1
6
0
1.75 1
3
2.50 1.25
3
8
1
1
2
1
1
1
1.15
0.50
0
2
M32
1
12.8
0
3.3
0.5
2
2
2
1.80
1.20
1.50
1
6
0
2.20 1
3
2.55 1.30
3
8
1
1
2
1
1
1
1.10
0.50
0
2
M33
1
14
0
5.25
0.5
2
2
2
1.70
1.10
1.55
1
6
0
2.00 1
3
2.50 1.20
3
8
1
1
2
1
1
1
1.15
0.50
0
2
M34
1
23
0
4.5
0.5
2
2
2
1.90
1.00
1.90
1
6
0
2.10 1
3
2.00 1.20
3
8
1
1
2
1
1
1
1.60
0.50
0
2
M35
1
23
0
12
0.6
2
2
2
1.95
1.10
1.77
1
6
0
2.20 1
3
2.50 12.00
3
8
1
1
2
1
1
1
1.65
0.50
0
2
?
...Continued on next page
MANAWADUGE ET AL.
NEW SPECIES OF APonogETon
APPENDIX 2. (Continued)
oOTUs
Character
Phytotaxa 275 (3) © 2016 Magnolia Press • 261
31 32
33 34
35
36 37 38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
M36
1
22.5
0
7.5
0.6
2
2
2
1.90
1.15
1.65
1
6
0
2.15
1
3
2.00
1.00
3
8
1
1
2
1
1
1
1.65
0.60
0
2
M37
1
34
0
12
0.8
2
2
2
2.15
1.25
1.72
1
6
0
2.85
1
3
2.50
1.20
3
8
1
1
2
1
1
1
1.75
0.60
0
2
M38
1
27
0
9
0.8
2
2
2
2.20
1.30
1.69
1
6
0
2.90
1
3
2.55
1.30
3
8
1
1
2
1
1
1
1.70
0.65
0
2
M39
1
28
0
9.5
0.6
2
2
2
2.30
1.30
1.77
1
6
0
3.05
1
3
1.20
1.25
3
8
1
1
2
1
1
1
1.80
0.60
0
2
M40
1
38
0
8
0.6
2
2
2
2.20
1.20
1.83
1
6
0
2.90
1
3
2.50
1.25
3
8
1
1
2
1
1
1
1.75
0.60
0
2
N41
0
56
0
15
0.7
1
2
0
1.65
1.13
1.46
1
6
1
1.88
0
3
1.63
0.88
0
2
0
0
1
0
0
1
9.00
3.00
1
0
N42
0
50
0
10
0.5
2
2
0
2.50
1.40
1.79
1
6
1
2.25
0
3
1.70
0.88
0
2
0
0
1
0
0
1
10.00
4.00
1
0
O43
0
45
0
12
0.6
1
2
0
1.70
1.30
1.31
1
6
1
1.75
3
3
2.55
1.05
0
2
0
0
1
0
0
1
12.00
4.00
1
0
O44
0
42
0
10
0.5
1
2
0
1.65
1.20
1.38
1
6
1
1.98
3
3
2.50
1.05
0
2
0
0
1
0
0
1
10.00
4.00
1
0
P45
0
150
0
20
0.8
2
2
0
1.55
1.60
0.97
1
6
1
1.60
2
3
1.20
0.38
0
2
0
0
1
0
0
1
9.00
0.60
1
0
P46
0
145
0
11
0.6
2
2
0
1.33
1.40
0.95
1
6
1
1.40
2
3
1.50
0.40
0
2
0
0
1
0
0
1
7.50
0.60
1
0
Q47
0
155
0
16
0.7
2
2
0
2.33
2.40
0.97
1
6
1
2.40
2
3
1.43
0.60
0
2
0
0
1
0
0
1
8.80
0.68
1
0
Q48
0
146
0
15
0.6
2
2
0
1.55
1.60
0.97
1
6
1
1.60
2
3
1.08
0.50
0
2
0
0
1
0
0
1
8.50
0.70
1
0
R49
0
65
0
15
0.5
0
2
0
1.55
1.25
1.24
1
6
1
1.55
0
3
1.70
1.20
1
2
0
0
1
0
0
0
8.70
3.50
1
0
R50
0
56
0
16
0.6
0
2
0
1.75
1.55
1.13
1
6
1
1.75
0
3
1.70
1.20
1
2
0
0
1
0
0
0
7.80
3.20
1
0
S51
0
40
0
18
0.7
0
2
1
1.50
1.20
1.25
1
6
1
1.55
0
3
1.50
0.50
1
2
0
0
1
0
0
0
8.80
3.75
1
0
S52
0
46
0
13
0.5
0
2
1
1.60
1.30
1.23
1
6
1
1.65
0
3
1.60
0.80
1
2
0
0
1
0
0
0
7.50
3.40
1
0
...Continued on next page
262 • Phytotaxa 275 (3) © 2016 Magnolia Press
APPENDIX 2. (Continued)
oOTUs
Character
MANAWADUGE ET AL.
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
T53
0
58
0
12
0.5
0
2
1
1.70
1.50
1.13
1
6
1
1.75
0
3
1.50
0.90
1
2
0
0
1
0
0
0
7.09
3.50
1
0
T54
0
43
0
11
0.6
0
2
1
1.50
1.40
1.07
1
6
1
1.55
0
3
1.50
0.80
1
2
0
0
1
0
0
0
8.20
3.50
1
0
U55
1
25
0
15
0.8
2
2
2
2.50
1.50
1.67
1
6
0
3.00
1
3
3.50
1.55
3
8
1
1
2
1
1
1
1.80
0.55
0
2
U56
1
30
0
11
0.5
2
2
2
2.90
1.80
1.61
1
6
0
2.90
1
3
3.40
1.60
3
8
1
1
2
1
1
1
1.50
0.50
0
2
V57
1
15
0
9
0.6
2
2
2
2.10
1.30
1.62
1
6
0
3.40
1
3
3.80
1.50
3
8
1
1
2
1
1
1
1.70
0.55
0
2
V58
1
20
0
8
0.6
2
2
2
2.60
1.60
1.63
1
6
0
4.00
1
3
3.90
1.60
3
8
1
1
2
1
1
1
1.90
0.55
0
2
W59
1
28
0
13
0.7
2
2
2
1.90
1.10
1.73
1
6
0
3.10
1
3
3.50
1.50
3
8
1
1
2
1
1
1
1.70
0.55
0
2
W60
1
35
0
10
0.8
2
2
2
2.50
1.50
1.67
1
6
0
4.00
1
3
3.80
1.50
3
8
1
1
2
1
1
1
1.60
0.50
0
2
X61
1
55
0
7
0.7
2
2
0
1.75
2.25
0.78
1
6
2
1.40
1
3
2.25
0.95
2
2
2
0
1
0
0
1
12.00
10.00
1
1
X62
1
53
0
6
0.8
2
2
0
1.70
2.50
0.68
1
6
2
1.35
1
3
2.10
0.85
2
2
2
0
1
0
0
1
10.00
9.00
1
1
Y63
0
98
0
14
0.8
2
2
0
1.33
1.43
0.93
1
6
1
1.50
2
3
1.50
0.60
0
2
0
0
1
0
0
1
9.00
0.60
1
0
Y64
0
80
0
10
0.9
2
2
0
1.40
1.55
0.90
1
6
1
1.40
2
3
1.20
0.60
0
2
0
0
1
0
0
1
6.20
0.50
1
0
Z65
0
59
0
16
0.7
2
2
0
1.30
1.50
0.87
1
6
1
1.70
2
3
1.10
0.50
0
2
0
0
1
0
0
1
7.10
0.60
1
0
Z66
0
151
0
15
0.8
2
2
0
1.50
1.65
0.91
1
6
1
1.60
2
3
1.00
0.50
0
2
0
0
1
0
0
1
5.80
0.50
1
0
AA67
1
28
0
15
0.8
2
2
2
1.90
1.10
1.73
1
6
0
2.10
1
3
3.30
1.30
3
8
1
1
2
1
1
1
1.50
0.50
0
2
AA68
1
30
0
13
0.7
2
2
2
2.00
1.30
1.54
1
6
0
2.00
1
3
3.50
1.50
3
8
1
1
2
1
1
1
1.70
0.60
0
2
AB69
0
80
0
10
0.8
2
2
0
1.50
1.70
0.88
1
6
1
1.60
2
3
1.30
0.50
0
2
0
0
1
0
0
1
6.20
0.50
1
0
AB70
0
75
0
15
0.6
2
2
0
1.30
1.40
0.93
1
6
1
1.70
2
3
1.20
0.60
0
2
0
0
1
0
0
1
7.70
0.60
1
0