Taprobanica (2011) Vol. 3. No. 1. Pages 01-49.

Published date: 30th, July 2011

TAPROBANICA the Journal of Asian Biodiversity ISSN 1800-427X - Volume 03, Number 01, pp. 1-49, Pls. 6.

© 2011, Taprobanica Private Limited, Jl. Kuricang 18 Gd.9 No.47, Ciputat 15412, Tangerang, Indonesia

EDITOR-IN-CHIEF THASUN AMARASINGHE

[email protected]

DEPUTY EDITORS SURANJAN KARUNARATHNE

[email protected] NIKI AMARASINGHE

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ASSOCIATE EDITORS MADHAVA BOTEJUE

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SANDY NURVI [email protected]

RIZKA ALNANDA [email protected]

SECTIONAL EDITORS MIGUEL ALONSO [email protected] UPALI AMARASINGHE [email protected] NATALIA ANANJEVA [email protected] MOHOMED BAHIR [email protected] AARON BAUER [email protected] BRUCE BEEHLER [email protected] FRANKY BOSSUYT [email protected] RAFE BROWN [email protected] BIJU DAS [email protected] INDRANEIL DAS [email protected] ANSLEM DE SILVA [email protected] REMA DEVI [email protected] SURATISSA DISSANAYAKE [email protected] ALAIN DUBOIS [email protected] ROHAN FERNANDO [email protected] COLIN GROVES [email protected] LEE HARDING [email protected]

S. HENKANATHTHEGEDARA [email protected] BRENDEN HOLLAND [email protected] KEVIN HYDE [email protected] DJOKO ISKANDAR [email protected] JAYANTHA JAYEWARDENE [email protected] H. KATHRIARACHCHI [email protected] ANDRE' KOCH [email protected] SARATH KOTAGAMA [email protected] SVEN KULLANDER [email protected] ENRIQUE LA MARCA [email protected] TZI MING LEONG [email protected] ARAVIND MADHYASTHA [email protected] K. MANAMENDRA-ARACHCHI [email protected] M. MEEGASKUMBURA [email protected] JEFFREY MILLER [email protected] MOHOMED NAJIM [email protected] ANNA NEKARIS [email protected]

VINCENT NIJMAN [email protected] HANS-DIETER PHILIPPEN [email protected] SUDHEERA RANWALA [email protected] DON REYNOLDS [email protected] JODI ROWLEY [email protected] JOHN RUDGE [email protected] PRASAD SENADHEERA [email protected] B. K. SHARMA [email protected] RALF SOMMERLAD [email protected] ROBERT STUEBING [email protected] JATNA SUPRIATNA [email protected] RAM VERMA [email protected] GERNOT VOGEL [email protected] RICHARD WAHLGREN [email protected] YEHUDAH WERNER [email protected] NIKHIL WHITAKER [email protected]

1 TAPROBANICA VOL. 03: NO. 01

EDITORIAL

A splitter’s systematics of writing:

scientific writing and writing English are separate issues and this has implications Publishing is an essential component of scientific activity and an increasing number of well-known forces, but including also editors, press us to publish much (Werner, 1978). Recently, refinement of some of these forces coerces at least some of us to publish not merely in peer-reviewed journals but in those that are hardest to penetrate. My personal opinion that this is to the detriment of science (Werner, 2009) does not help. Publishing well is difficult. Here I try to analyze part of the difficulty and to conclude partial remedies.

The English language, and especially “American” (USAian), has become, or is becoming, the world’s language of science. This is due to two successive historical situations: the size and geographical spread of the past British Empire, and the size and scientific productivity of the USA. Certainly, having a global language is advantageous for science. But that this language is English, is unfortunate. English is relatively difficult to learn, because there is no regular correspondence of spelling with pronunciation: “Now you read the red book that I read yesterday” – and this is a moderate example. This drawback is claimed to be offset by the famous richness of the English vocabulary. The latter is due in part to the language’s dual origin. Additionally, linguistic richness goes hand in hand with national culture; not everybody has horse-stallion-gelding-mare-foal-colt-filly. Piquantly, English has no word for the German verb gönnen, the opposite of grudge (=begrudge), i.e., feeling happy with somebody else’s good fortune. Anyway, we are stuck with writing our papers in English. Writing a scientific paper is a challenge, even after problems of contents have been settled. Help is available from many books (my favorite is by Trelease, 1969; but see comparative book review by Baker, 1969), guideline brochures, articles, websites of publishers and very detailed Instructions for Authors in some of the journals. In biology, many editors recommend, or even declare as adopted, the CBE (Council of Biology Editors) Style Manual (CBE Style Manual Committee, 1994) but few impose its simplest rules. Most of all this material radiates an attitude that scientific writing and writing English constitute one problem. In this spirit, a long scholarly editorial in an upper-class journal recently admonished, especially young Americans, to write more carefully (Heatwole, 2008). Rather than quote and repeat here that review of sloppy and faulty English in science papers (that is worth reading), I endeavor to base my argument below on additional points.

I consider scientific writing and writing English to be separate issues that sometimes clash. A notable example is, when X moved several times from A to B, the English teacher urges, “Don’t monotonously repeat yourself. You have at your disposal walk, go, march, step, pace, stride, rush and others”. In contrast, the scientific writing guide says, “If X did the same thing repeatedly, use the same verb. Keep the uniformity obvious, also in consideration of non-native readers.” Most of the specific characteristics of scientific writing have nothing to do with English; they apply in any language. And some counter the ideals of elegant writing, in any language. They are intended to enhance communication, also for non-native readers. Foremost among these are plain language (as exemplified above); avoidance of verbosity (drop any phrase or word that does not add information; “very quickly” says no more than “quickly”); and avoidance of redundancy (said once is enough). These function in part through promoting brevity. Brevity in itself is good for the reader and therefore for the author and it alleviates the costs of publication. Several rules of scientific writing are desirable everywhere although commonly taken lightly, but are absolutely necessary for the high-fidelity communication needed in science. These include clarity and lack of ambiguity. General words should be replaced with specific ones. Not “as he went” – either “when he went” or “because he went”. Not “while he went” – either “when he went” or “although he went” (This one may vary among languages.). In this context I particularly negate the use of three punctuation marks. The slash or diagonal has too many meanings: A/B can mean (1) A divided by B. (2) A, which is part of B. (3) B, which is part of A. (4) A and B. (e.g., males and females). (5) A or B. (6) In “and/or” (prohibited but not

TAPROBANICA, ISSN 1800-427X. April, 2011. Vol. 03, No. 01: pp. 1-4. © Taprobanica Private Limited, Jl. Kuricang 18 Gd.9 No.47, Ciputat 15412, Tangerang, Indonesia.


imposed) it stands for A or B or both. I advocate minimizing the use of the hyphen because it so easily mutates from separating to connecting and vice versa. Worst is its common use in hybrid expressions of ranges. “Temperature varied from 10-15 oC” is prohibited (but rarely prevented) because it could be the beginning of “Temperature varied from 10-15 oC at night to 30-35 oC by day.” Last, “bullets” to emphasize items in a list, come from advertizing. They are the uncivilized visual counterpart of raising the voice in vocal communication. Serial numbers would be more useful, can be referred to. The need for sentence structure to facilitate understanding is much more acute in scientific writing than in other literature. Snake-sentences should be avoided even by herpetologists. Greater attention is required for the concept of parallelism. Not “There were 20 chairs in classroom A but in B only 18.” Rather, “There were 20 chairs in classroom A but only 18 in B”.

Another problem of English, that looms larger in scientific writing, is the capitalization of vernacular names of animals and plants. Many editors want these names to augment the scientific names, ostensibly because they could be more stable, or to attract readers to whom they are more acceptable. Normally such names are nouns in the English language and not capitalized (University of Chicago Press, 1969). “Have you tasted the fried Grey Mullet?” Looks as strange as “Have you fed the Cat”. Until recently only (or mainly) the ornithologists have promoted their birds to capitalization, based on their global check-list. But for other groups there are no global lists and no general agreements. There is no reason to venerate (and capitalize) the English names given by enterprising Americans to exotic animals, the more so since many of these names violate the natural logic of vernacular names. Some issues arise uniquely, or at least mainly, in scientific writing. The aspiration to clarity begets the (admittedly debated) rule to write in active voice, first person, and moreover for a single author (now almost extinct), in singular. This ‘arrogance’ is briefer, clearer, accepts responsibility. Abbreviations should be explained (once per paper!). Interestingly editors plead to minimize the use of abbreviations, while they urge to keep the text short. Some editors want them to be explained at first mention, charging the reader with remembering; other request a concentrated look-up table. Units of assorted measures must be metric. (Isn’t it amusing how after the inch became metrically subdivided for precision purposes, now the geographical degrees are metrically subdivided. Of course, to the second it happened long ago.) Incidentally, sentences may not begin with an abbreviation or with digits.

The abstract is a major stumbling block. It should represent the contents, not define it. Thus “The average number of chairs per classroom was 18.5” and not “This project describes the furniture in classrooms” or “The number of chairs in classrooms was surveyed”. It may not contain anything not represented in the main text. Opinions differ whether it, too, should be in active voice or maybe in passive voice. In contrast, there exist pitfalls in writing English that do not really affect the fidelity of scientific communication. For some reason, we study at a university but in a department (e.g., of Zoology). If we ignorantly follow our logic and exchange the prepositions, everybody understands. If instead of “fewer readers” we write “less readers”, everybody understands. If we, like most people, err in selecting “which” or “that” (see your larger dictionary), everybody understands. Boorish English in itself, although not exactly an honor, does not necessarily impede communication. For decades, the Journal of Zoology, London, has tolerated the faulty English of authors, as long as there occurred no ambiguity or other doubt as to meaning. Communicating in science and writing English are separate issues, and this generates some practical conclusions. At another level, it is obviously important that authors would really say what they intend to say. (This differs from the issue whether what they intend to say is right or wrong.) Let me bring three examples of failure. The first is my own; the others are from much greater authors. (1) “The regenerated portion of each tail was distinguishable not only by radiography, but also by its ventral scales, which were smaller and paler than the original ones.” (Werner, 1967). Simple? But unfortunately, ventrally the regenerated part was darker, not paler, than the original, as shown in the accompanying photographs. (2) “Critical reexamination of Gadow’s theory of the composition of vertebrae from four pairs of arcualia is inapplicable to tetrapods…” (Williams, 1959). This was the opening sentence of the abstract! The author obviously intended to say, not that the reexamination was inapplicable, but that it showed that the theory was inapplicable (as shown in the text). (3) “A recent example of an uncertain object of a verb is: ‘Males consumed more prey than females.’ This implies that males consumed females to a greater extent than males consumed prey.” (see


Heatwole, 2008). But the commenting author obviously intended to say, “This implies that males consumed females to a lesser extent than males consumed prey.” One might well ask, so, whence all this sloppiness that has been lamented by Heatwole (2008) and indicated here? While a general review and analysis of sloppiness are outside my competence and Taprobanica’s scope, in this case I can suggest a hypothetical source: University professors, the people who educate heads of state, school teachers, and scientists. Based on my experience in Israel and the USA, and on evidence from Germany, I dare say that they too often refrain from correcting their students. Saving their own time is only a fringe benefit. The real purpose is to be attractive to the students, because nowadays research students are manpower producing papers for the teacher’s resume. They say, “No need to harass them with technicalities. They will learn them when needed.” They don’t consider that sloppiness is sloppiness and in some situations (the roads are famous) sloppiness is lethal. So what are we to do about all this? I see two practical conclusions. First, if we accept that scientific writing and writing English are separate issues, we should separate their teaching. Scientific writing should be taught everywhere in the local language, including all examples and exercises. This would make the teaching, including learning, more effective, and would lead to better understanding and acceptance of the logic. It would also increase the number of potential teachers. Separately, it would be nice to have the teaching of English improved (expanded and deepened) and this can be done (or guided) by teachers who are ignorant of science. The students should be able to merge the two inputs and would gain much more. Second, it is always prudent and productive to show our manuscript to a friend before we ejaculate it to an editor. But such preliminary peer review is doubly important when we suspect that the editor knows us and respects us and thus might be misguided to economize and take the reviewing lightly. Third, if we could revive the historical ethics that a research student’s project is the student’s project and is published by the student (the teacher is paid to teach the student how), we would restrain the incentive for the teacher to attract students. Students would still be wanted, to pass the flame to them, but would be educated without such fear of hurting their feelings. Acknowledgement I thank Harold Heatwole for constructive and encouraging reading of the manuscript. Literature cited Baker, S., 1969. Book reviews: Clarity. Science, 166: 365-366. CBE Style Manual Committee, 1994. Scientific style and format: The CBE manual for authors, editors, and publishers. 6th ed. Style Manual Committee, Council of Biology Editors, Cambridge and New York, Cambridge University Press: 825. Heatwole, H., 2008. Editorial—A plea for scholarly writing. Integrative and Comparative Biology, 48: 159–163. Trelease, S. F., 1969. How to write scientific and technical papers. Cambridge, Mass., M.I.T. Press: 39. University of Chicago Press, 1969. A manual of style. 12th ed. Chicago and London, The University of Chicago Press. Werner, Y. L., 1967. Regeneration of specialized scales in tails of Teratoscincus (Reptilia: Gekkonidae). Senckenbergiana. Biologia, 48:117-124. Werner, Y. L., 1978. How editors catalyze the publication explosion, in: Scientific information transfer: the editor’s role (Proceedings of 1st International Conference of Scientific Editors, Jerusalem, Israel, 1977) ed. M. Balaban. Dordrecht, Holland & Boston, Mass., USA, D. Reidel Publ. Co., 113-121. Werner, Y. L., 2009. The Aspiration to be Good is Bad: The ‘impact factor’ hurts both science and society. The International Journal of Science in Society, 1: 99-106.


Williams, E. E., 1959. Gadow's arcualia and the development of tetrapod vertebrae. Quarterly Review of Biology, 34: 1–32.

Yehudah L. Werner Sectional Editor: Taprobanica, the journal of Asian Biodiversity March 04th, 2011 The Alexander Silberman Institute of Life Sciences, The Hebrew University of Jerusalem, 91904 Jerusalem, ISRAEL and Museum of Zoology (Museum für Tierkunde), Senckenberg Dresden, A. B. Meyer Building, Königsbrücker Landstraße 159, D-01109 Dresden, GERMANY From Volume 3: Number 1, Taprobanica – The Journal of Asian Biodiversity (ISSN: 1800-427X) is published by Taprobanica Private Limited – Sri Lanka, which is registered as a private company limited, incorporated under the Companies Act No. 7 of the Democratic Socialist Republic of Sri Lanka (company registration no. PV 77205) as of 15th February 2011. The trade mark for the journal is illustrated here (slightly changed from the previous mark published).

This trade mark is registered under the Intellectual Property Act No. 36 of 2003 as an individual trade mark of A. A. Thasun Amarasinghe (trade mark registration no. LK/T/1/164048).

The editor-in-chief (Taprobanica journal) & Chairman (Taprobanica Pvt. Ltd.)


REPORT OF SOME NOTEWORTHY SPECIMENS AND SPECIES OF HERPETOFAUNA FROM SOUTH-EAST INDIA

Sectional Editor: Gernot Vogel Submitted: 24 March 2010, Accepted: 13 June 2011

S. R. Ganesh 1,2 and S. R. Chandramouli 1

1 Department of Zoology, Division of Wildlife Biology, A.V.C College, Mannampandal, Mayiladuthurai–609 305, Tamil Nadu, India 2 Chennai Snake Park, Rajbhavan post, Chennai - 600 020, Tamil Nadu, India E-mail: [email protected] Abstract We report abnormal individuals of Ramanella variegata, Lycodon aulicus (sensu lato), Bungarus caeruleus which exhibited variation from the ‘typical morphs’ of their respective species. Also we report a rarely recorded species Polypedates cf. leucomystax (from south India), from the Mannampandal area of Tamil Nadu. These observations based on voucher photographs are presented for the first time. Keywords: Morph, aberration, variation, phenotypic plasticity, polymorphism Introduction The Coromandel Coast of India was one of the first areas in south Asia where herpetological investigations began, dating back to Russell (1796, 1801). Several species have their type localities in ‘Tranquebar’ (now Tarangambadi), Pondicherry, ‘Madras’ (now Chennai) and ‘Vizagapatam’ (now Vishakapatnam), located in this part of India. The herpetofauna of Mannampandal village (11°09’N 079°68’E; 19 m a.s.l.) in Mayiladuthurai Taluk, Nagapattinam District, ca. 28 km west off the historical place Tranquebar has been briefly discussed (Kannan et al., 1994; Ganesh & Chandramouli, 2007). In this paper we report on certain unique specimens and species of

herpetofauna from this area, which are noteworthy in terms of some of their hitherto unknown natural history traits. Materials and Methods These observations were made by random and/or opportunistic sightings between July 2006 and November 2008 in and around Anbanadhapuram Vahaira Charity (A.V.C) College campus. Animals seen were diagnosed, measured and photographed in-situ using Canon Powershot A640 and Canon EOS 400D model cameras. Values of a character presented for more than one individual are separated by a comma. Altitude was determined by


SOME HERPETOFAUNA FROM SOUTH-EAST INDIA


Garmin 12 channel Global Positioning System readings taken at the locality. Syntopic conspecifics observed were also used for comparison, in addition to published keys, so as to determine any possible patterns of geographically correlated variance if present. However we omitted all data that did not deviate from literature, unless strongly needed, as a backup for species identity. Observations and Discussion Ramanella variegata (Stoliczka, 1872) An adult (Fig.1) sighted on 23/10/07 on tar road, during a rainy night. Dorsum plain grayish brown all over without any visible markings or patterns; supraocular area bluish, labials and gular bluish grey, dorsal parts of fore and hindlimbs pale white, with islets of dark grayish brown, best visible in femur and humerus; venter pinkish white, iris and pupil black and indistinguishable. The usual colouration of this species, which is the only Ramanella distributed in the southern Indian plains; is olive brown above, finely marbled with yellow or cream, underside white, sometimes marked with brown on the throat and sides (Biju, 2001; Daniels, 2005; Dutta, 1997; Dutta & Manamendra-Arachchi, 1996). The specific epithet ‘variegata’ and its common English name ‘marbled’ narrow mouthed frog are indicative of the variegated / marbled pattern of its dorsum. Uniform grey colouration in this species is hitherto unreported in the literature. Fig. 1: Ramanella variegata Polypedates cf. leucomystax (Gravenhorst, 1829) Two adults (Fig. 2) sighted on 29/10/07 and 28/9/08 on shrubs at night. Dorsum yellowish brown with four darker stripes extending from postnasal, the inner two stripes being paravertebral and the outer two being dorsolateral, the outer and the inner

stripes converge at the supraocular region, from where they divide posteriorly to pass through the temporal area and extend to the cloaca, the outer stripes being broadest at mid-torso, exactly at the articulation of the hindlimb with the trunk; postocular region bluish, infralabial and gular surfaces off-white, limbs dorsally cross-barred with darker shades, best visible on the proximal elements of the limbs; venter pinkish white, iris golden brown, pupil horizontal; snout-vent length 20.0, 50.0; axilla-groin distance: 8.3, 20.7; head length 9.0, 22.6; head width 7.0, 17.6; snout length 3.0, 7.6; eye diameter 1.3, 3.3; tympanum diameter 1.3, 3.3; internarial distance 1.9, 4.7; interorbital distance 3.0, 7.5. We observed several Polypedates maculatus (Gray, 1834) that matched the descriptions in literature (Dutta, 1997; Dutta & Manamendra-Arachchi, 1996) but these two individuals are certainly not P. maculatus as there is no dorsally ‘striped’ pattern present in P. maculatus, whose specific epithet means ‘spotted’ (Dutta & Manamendra-Arachchi, 1996). Dutta (1997) stated that P. leucomystax is widely distributed in most parts of Southeast Asia and different colour morphs led to the erection of subspecies from different geographic localities. It differs from its closely allied congener P. maculatus by an osteological character ‘parieto-squamosal arch bone’ which is evident at normal resting posture in P. lecomystax but not in P. maculatus (Daniel, 2002; Daniels, 2005; Dutta, 1997). Dutta (1997) remarked that some earlier authors considered P. maculatus and leucomystax to be subspecies and the occurrence of P. leucomystax in Sri Lanka is erroneous and in Karnataka, south India is doubtful. The report from the Western Ghats of Karnataka was once considered authentic and then ‘changed’ to doubtful (Daniels, 1997, 2000 & 2005). Biju (2001) and Daniel (2002) deny its presence in south India. Banerjee & Deuti (2006) give its English name as ‘four-lined’ tree frog (vs. ‘six-lined’ tree frog fide Daniels, 2005), which is consistent with its former, specific epithet ‘quadrilineata’ [(sensu Boie, 1835) see Dutta, 1997]. Soud & Das (2005) state P. leucomystax to be common in low lands and urban areas with water and prolific vegetation, in the Bongaigon District of Assam State. Hussain et al. (1999) mention its distribution in Northeast India up to an elevation of 300 m asl. Deuti (1997) reports a range extension of this species from Sikkim, West Bengal, Assam,

GANESH & CHANDRAMOULI, 2011


Meghalaya, Arunachal Pradesh, Manipur and also curiously from Gujarat and Madhya Pradesh. Rao et al. (2005) mentions its occurrence in Nallamalai hills, a part of the Eastern Ghats of Andhra Pradesh in south India, with photographic evidence. Given this scenario, it can no longer be considered as a mesic forest habitat specialist, but is rather a eurytopic species occurring in plains and anthropogenic habitats as well. Since P. leucomystax in itself is a species-complex containing sympatric morphotypes (Narins et al., 1998), we refer our specimens to Polypedates cf. leucomystax, based on evident parietosquamosal arch bone visible at rest and the ‘striped’ dorsum.

Fig. 02: Polypedates cf. leucomystax

Lycodon aulicus (Linnaeus, 1758) sensu lato Table 1: Comparison of meristic, morphologic and metric characters between ‘typical’ and ‘aberrant’ morphs of syntopic adult males of Lycodon aulicus from Mannampandal.

No Characters Morph 1 Morph 2 1 Scalerows (smooth) 17:17:15 17:17:15 2 Apical pits towards centre of tip towards upside of tip 3 Supralabials (those touching eye) 9 (3,4,5) 9, 11 (3,4,5) 4 Infralabials (those touching genials) 9 (6,7) 9, 10 (5) 5 Loreal (horizontally elongate) 1 1 6 Temporal 2+3 2+2 7 Preocular 1 1 8 Postocular 2 2 9 Preventrals 3 3

10 Linguals 5 4 11 Ventrals (strongly angulate laterally) 218 199 12 Anals 2 2 13 Subcaudals (divided) + terminal scale 68 pairs +1 68 pairs + 1 14 Nuchal mark inverted V shape V shape 15 Band structure parallel diverging 16 Band pattern patterned interiorly plain interiorly 17 Band extent visible dorsally only visible laterally also 18 Head length 21.3 25 19 Snout length 7 3.3 20 Head width (maximum) 13 14.5 21 Head width (eye level) 10.3 11 22 Neck width 10.6 10.6 23 Eye diameter 3.6 1.3 24 Loreal length 3 2.5 25 Lower eye margin–lip distance 2 1.3 26 Inter orbital distance 5.9 3.3 27 Position of first band (respect to ventrals) 9 0 28 Scales between parietal and first band 11 2 29 Head length: snout length 3.04 7.37



30 Head length: maximum head width 1.63 1.72 31 Max. head width: neck width 1.22 1.36 32 Head width (eye level): neck width 0.97 1.03 33 Eye diameter: lower eye margin-lip distance 1.8 1 34 Loreal length: eye diameter 1.2 1.92 35 Snout length: Interorbital distance 1.18 1

An adult male, (Figs. 3 & 4) one each of the two morphs, on tarred road and brick pile (respectively), at night. Comparison of characters 2, 14-17 and 27-35 in the above table reveals considerable differences in cephalic morphometry, general habitus and colouration between these two morphs. There is no literature report about this phenomenon (Sharma, 2003; Smith, 1943; Whitaker, 1978; Whitaker & Captain, 2004), except Daniel (2002) who gives drawings [from Wall] of its various colour morphs. Smith (1943) in his line drawings, Daniel (2002), Whitaker & Captain (2004) and Goonewardene et al. (2006) in their photographs depict morph 1 with thick head and first band in inverted-V shape just behind parietals (i.e., on the head and well before the neck). Das (2002) Das & de Silva (2005) and Whitaker (1978) depict morph 2, with the first band in V shape, well off the parietals, but near the neck. Rao et al. (2005) from Nallamalai hills depict morph 2 as L. aulicus and morph 1 (incorrectly) as L. travancoricus, which apart from our record, are another proof for syntopy between these two morphs. Though morph 2 superficially resembles L. osmanhilli (Taylor, 1950) of Sri Lanka, it differs from the latter by the character preocular contacting frontal (vs. not in contact, in L. osmanhilli) (de Silva, 1980). Therefore, we doubt that the Lycodon aulicus s. lat. complex is yet taxonomically unresolved. We have given the differences between merely a single representative from each of the two morphs, but this is evident enough to distinguish them. In the live individual of morph 1, we counted 218 ventrals excluding preventrals (vs. < 214 in Smith, 1943; Whitaker & Captain, 2004) which imply that literature defining this species cannot be considered as fully comprehensive. Considering the variations shown herein and the rich, subjective synonyms originating from places far and wide, we strongly suggest that more detailed studies needed to be undertaken to resolve the taxonomy of L. aulicus (sensu lato). Similar works involving subtle variation in colouration and morphometry led to the resurrection of Dendrophis chairecacos Boie, 1827 and Dipsas schokari Kuhl, 1820 from the synonymy of Dendrelaphis tristis (Daudin, 1803) sensu Smith (1943) (see Van Rooijen & Vogel,

2008 & ‘2009’ 2010). Fig. 3: Lycodon aulicus morph 1 Fig. 4: Lycodon cf. aulicus morph 2 Bungarus caeruleus (Schneider, 1802) An adult (Fig. 5), dead individual measuring 780 mm observed at night on 23/8/07 on a tarred road during a rainy night. Dorsally grayish black without any white cross bands and was without even a speck of white on the dorsum; supralabials, penultimate costals and venter white. Scalerows (smooth) 15:15:15; ventrals (not angulate laterally) 202; subcaudals (undivided) 40; anal 1; supralabials (touching eye) 7 (3, 4); preocular 1; postoculars 2; temporals 1+2. Scalation of our individual agrees with literature (Russell, 1796; Whitaker & Captain, 2004) accounts of Bungarus caeruleus. It is certainly not B. niger Wall, 1908 as the distribution records are well off the mark and B. niger has higher ventral

8

GANESH & CHANDRAMOULI, 2011


and subcaudal counts (ventrals: 216–231; subcaudals: 47-57) than B. caeruleus, (see Smith, 1943; Whitaker & Captain, 2004). Fig. 5: Bungarus caeruleus The aberrant individual was not observed to be in the cycle of ecdysis, which can render body patterns unclear (Whitaker & Captain, 2004). It was an adult, 780 mm long, which is noteworthy here as young snakes are known to have much more intense patterns than adults, as seen in Eryx johnii, Macrophisthodon plumbicolor, Argyrogena fasciolatus, Ophiophagus hannah and Bungarus caeruleus (see Smith, 1943; Whitaker & Captain, 2004). Full grown individuals of this species are known to become dull gray in colour without any banded pattern (Anslem de Silva pers. comm., March, 2010). The total length of our individual was 780 mm, which is close to the average length (1000 mm) and is nowhere near the maximum length (1750 mm) (Whitaker & Captain, 2004). Moreover, B. caeruleus exhibits a peculiar phenomenon of colour variation with respect to geography. In India, the south-west coast and the south-east coast populations of B. caeruleus differ in colouration, with the west coast kraits being more evidently banded than those of the east (Das, 2002). All conspecifics (both adults and juveniles) observed from the present area, were typically banded (pers. obs.). Dravidamani et al. (2006) examined up to 200 individuals, but failed to record any aberrations. Kuch (1991), Whitaker (1969) and Vogel & Chanhome (2006) report of banded snakes like Bungarus fasciatus and Boiga dendrophila melanota exhibiting aberrant pattern, notably stripes instead of the usual banded pattern (see Whitaker & Captain, 2004; Smith, 1943; Vogel & Chanhome, 2006). Kuch (1991) stated that he provisionally preferred to regard longitudinally striped aberrant snakes to be individual mutations rather than a geographically correlated phenotype. While Whitaker (1969) commented that the female and all six juveniles were striped, with just two bands on

the tail and in scalation they did not differ from Bungarus fasciatus. Vogel & Chanhome (2006) remarked that such phenomenon of parent and offspring exhibiting similar, consistent variation was associated with low incubation temperatures or a dominant recessive genetic disposition. Since we observed only one individual with this sort of aberration, we are presently unable to comment on the reason for the same. Remarks Our observations indicate the lacuna present in the herpetological community of this highly anthropogenic, non-forested alluvial plains country, which is no more than a matrix of plantations and rivulets, despite the fact that many of the species reported here are widespread in the country and are often encountered in the wild. Phenotypic plasticity has been a very interesting and often highly influential factor in contributing to polymorphism. Prince et al. (2003) state that “genotype + environment + random variation phenotype”, which are aberrant individuals that differ from normal conspecifics in any noticeable way including morphology, physiology and behaviour. Very little published information is available on phenotypic plasticity in Indian herpetofauna. Acknowledgements We thank our college’s Zoology Department and Men’s Hostel for providing the necessary infrastructure; Agumbe Rainforest Research Station, Chennai Snake Park and Madras Crocodile Bank for access to their libraries; herpetologists, Anslem de Silva, Biju Sathyabama Das, Aaron Bauer, Romulus Earl Whitaker and Ruchira Somaweera for their advice, suggestions and literature. Literature Cited Banerjee, A. and K. Deuti, 2006. Breeding record of four-lined tree frog (Polypedates leucomystax) (Gravenhorst, 1829) at Rajpur, South 24 Praganas District, West Bengal. Cobra, 64: 16-17. Biju, S. D., 2001. A Synopsis to the frog fauna of Western Ghats, India. Indian Society for Conservation Biology. Occasional Publication, (1): 1-24. Daniel, J. C., 2002. The Book of Indian Reptiles and Amphibians. Bombay Natural History Society, Oxford University Press, Mumbai: 238. Daniels, R. J. R. 1997. A Field Guide to the frogs and toads of the Western Ghats. Part III. Cobra, 29:1-13.

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Daniels, R. J. R., 2000. Reptiles and amphibians of Karnataka. Cobra, 42: 1-11. Daniels, R. J. R., 2005. Amphibians of Peninsular India. Universities Press, Hyderabad, India: 286. Das, I., 2002. A Photographic Guide to Snakes and other Reptiles of India. New Holland publications, London, UK: 144. Das, I. and A. de Silva, 2005. A Photographic Guide to Snakes and other Reptiles of Sri Lanka. New Holland Publications, London, UK 144. De Silva, P. H. D. H., 1980. Snake fauna of Sri Lanka, with special reference to skull, dentition and venoms in snakes. National Museum of Sri Lanka publication. Kandy, Sri Lanka: 472. Deuti, K., 1997. Range extension of some Indian Amphibians. Cobra, 29: 19-28. Dravidamani, S., P. Kannan, V. Kalaiarasan, R. Deepika, J. Gitanjali, J. R. Moss and S. Rajan, 2006. Studies on the size composition and morphometry of common krait Bungarus caeruleus (Schneider, 1801) at the Irula Snake Catchers’ Industrial Co-operative Society, Vadanemmeli, Kanchipuram district, Tamil Nadu. Cobra, 64: 1-7. Dutta, S. K. and K. Manamendra-Arachchi, 1996. The Amphibian fauna of Sri Lanka. Wildlife Heritage Trust, Sri Lanka: 230. Dutta, S. K., 1997. The Amphibians of India and Sri Lanka. Odyssey Book House, Orissa, India: 342. Ganesh, S. R. and S. R. Chandramouli, 2007. A study of herpetofaunal community in Mannampandal, Nagapatinam District, Tamil Nadu. Cobra, 1 (4): 33-43. Goonewardene, S., J. S. Drake and A. de Silva, 2006. The Herpetofauna of the Knuckles Range. Amphibian and Reptile Research Organization of Sri Lanka: 208. Hussain, B., N. Choudry and S. Senguptha, 1999. A note on the reproduction in Polypedates leucomystax (Gravenhorst, 1829). Hamadryad, 24 (1): 44-45. Kannan, P, C. Sankaravadivelu and V. Kalaiarasan, 1994. Herpetofaunal assemblage in Mayiladuthurai area: A conservational approach. Cobra, (18): 1-15. Kuch, U., 1991. Abnormal colouration in the banded krait Bungarus fasciatus (Schneider, 1801). The Snake, 23: 25-28. Narins, P. M, A. S. Feng, H. Yong and J. Christensen-Dalsgaard, 1998. Morphological, behavioral and genetic divergence of sympatric morphotypes of the tree frog Polypedates leucomystax in Peninsular Malaysia. Herpetologica, 54 (2): 129-142.

Prince, T. D., A. Quarnstrom and D. E. Irwin, 2003. The role of phenotypic plasticity in driving evolution. Proceedings of Biological Sciences, 270: 1433-1440. Rao, T., H. V. Ghate, M. Sudhakar, S. M. M. Javed, and S. R. Krishna, 2005. Herpetofauna of Nallamalai Hills, with eleven new records from the region, including ten new records from Andhra Pradesh. Zoo’s Print Journal, 20 (1): 1737-1740. Russell, P., 1796. An account of Indian serpents collected on the coast of Coromandel, containing descriptions and drawings of each species together with several experiments and remarks on their poisons. Shakespeare Press, London: 49+43 pls. Russell, P., 1801. Continuation of an account of Indian Serpents containing descriptions and figures from specimens and drawings, transmitted from various parts of India. Shakespeare Press, London: 45+39 pls. Sharma, R. C., 2003. Handbook – Indian Snakes. Zoological Survey of India, Kolkata, India: 292. Smith, M. A., 1943. Fauna of British India, including Ceylon and Burma. Vol-III Serpentes. Taylor & Francis publications, London : 583. Soud, R. and R. Das, 2005. Some observations on Polypedates leucomystax (Gravenhorst, 1829) (Anura: Rhacophoridae) from urban areas. Cobra, 62: 29-30. Van Rooijen, J. and G. Vogel, 2008. An investigation into the taxonomy of Dendrelaphis tristis (Daudin, 1803): revalidation of Dipsas schokari Khul, 1820 (Serpentes, Colubridae). Contributions to Zoology, 77: 33-43. Van Rooijen, J. and G. Vogel, 2009 ‘2010’. A multivariate investigation into the population systematics of Dendrelaphis tristis (Daudin, 1803) and Dendrelaphis schokari (Khul, 1820): revalidation of Dendrophis chairecacos Boie, 1827 (Srepentes, Colubridae). The Herpetological Journal, 19: 193-200. Vogel, G. and L. Chanhome, 2006. On two remarkable colour variants in Boiga dendrophila melanota (Boulenger, 1896) (Serpentes: Colubridae). Hamadryad, 30 (1&2): 199-203. Whitaker, R., 1969. Abnormal colouration in the Banded Krait (Bungarus fasciatus). Journal of the Bombay Natural History Society, 66 (1): 184-185. Whitaker, R., 1978. Common Indian Snakes – A Field Guide. MacMillan Press, New Delhi, India: 154. Whitaker, R. and A. Captain, 2004. Snakes of India – The Field Guide. Draco Books, Chengalpet, South India: 481.

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ON A RARE, SOUTH INDIAN BURROWING SNAKE Platyplectrurus trilineatus (BEDDOME, 1867) Sectional Editor: Gernot Vogel Submitted: 01 April 2011, Accepted: 13 June 2011

S. R. Ganesh*

* Chennai Snake Park, Rajbhavan post, Chennai - 600 020, Tamil Nadu, India; E-mail: [email protected] Abstract Examination of five juvenile preserved specimens of Platyplectrurus trilineatus, an endemic, poorly-known Uropeltid snake species from the Western Ghats Mountains of Southwestern India provided further insights into its taxonomy. The sample examined here agreed well with the existing descriptions in literature in colouration and most aspects of scalation but had larger range of ventral scale count and smaller supraocular relative to prefrontal. Character definition (in the case of ventrals) and ontogenic variation (in the case of supraocular size) might have possibly created these discrepancies. These differences indicate that a better sampling of both specimens and characters would throw more light on this species. Keywords: morphology, characterization, lepidosis, habitus, colouration Introduction The snake family Uropeltidae Müller, 1832 is one of the most poorly-understood families of small, burrowing snakes restricted to the Ceylonese-Malabar subregion of south Asia (Rajendran, 1985). Platyplectrurus Günther, 1868 is a species-poor genus endemic to the Western Ghats Mountains of southwestern India, containing two valid species viz. P. trilineatus (Beddome, 1867) and P. madurensis Beddome, 1877 (Smith, 1943). Beddome (1867) described Plectrurus trilineatus based on a specimen from “Anamallay forests; elevation 4000 feet”. He doubted its generic

allocation by writing “Plectrurus?” and “…..will perhaps have to be placed in a new genus.” A year later, Günther (1868) erected the new genus Platyplectrurus for this species, thus giving it the currently-valid name combination Platyplectrurus trilineatus. Subsequently, Beddome (1886) procured six more specimens from the same general locality and reported a range of variation in ventral and subcaudal counts for four females and three males (including the holotype). In the same work, i.e. Beddome (1886), Platyplectrurus bilineatus was described as a new species, based on two syntypes (fide Boulenger, 1893: 166) that were “probably not

TAPROBANICA, ISSN 1800-427X. April, 2011. Vol. 03, No. 01: pp. 11-14, 1 pl. © Taprobanica Private Limited, Jl. Kuricang 18 Gd.9 No.47, Ciputat 15412, Tangerang, Indonesia.

ON FURTHER SPECIMENS OF Platyplectrurus trilineatus (BEDDOME, 1867)


adults”, collected from “Madura Hills”. Later, P. bilineatus was synonymized with P. trilineatus, as Beddome’s purported specifically-distinct characters were considered as intraspecific variations (Boulenger, 1890). This view is still being followed (see Smith, 1943; Whitaker, 1978; Rajendran, 1985; Das, 2002; Whitaker and Captain, 2004). Platyplectrurus trilineatus has seldom been reportedly sighted / collected since then, as several surveys in southern Western Ghats did not yield this species (Ferguson, 1895; Hutton, 1949; Hutton and David, 2009; Inger et al., 1984; Ishwar et al., 2001; Malhotra and Davis, 1991; Wall, 1919, 1920). However, Rajendran (1985) collected this species and provided further morphological characterization based on his specimens. In spite of this, P. trilineatus still remains to be a little-known species, as the latest treatises on Indian snakes did not deal with this species (Daniel, 2002; Das, 2002; Whitaker and Captain, 2004). I discovered five specimens labeled as “Platyplectrurus madurensis”, which I identified as P. trilineatus (see Kalaiarasan et al., 1995; Ganesh, 2010), with unknown collection locality, in the museum of the Chennai Snake Park. Owing to the paucity of published accounts on this poorly-known species, this paper is presented to further improve its morphological characterization based on character-state data obtained from these apparently unpublished specimens. Materials and Methods Five, formalin-preserved, juvenile specimens were examined and their scale counts, measurements and colour pattern were recorded. Scalerows were counted around midbody. Scales from the mental up to the scale before the anal scale were counted as ventrals (Gower and Ablett, 2006). The terminal scale is excluded from the subcaudal scale count. Scales between rostral and the final scale bordering jaw angle were counted as supralabials, those touching eye given within parenthesis. Scales between mental and scale bordering last supralabial were counted as infralabials, those touching genials given within parenthesis. Scales surrounded by supralabials, postoculars and parietals were counted as temporals. Symmetrical head scalation character values were given in left / right order. Morphological data included colouration and pattern in formalin. Measurements were recorded using vernier callipers, except snout-vent and total lengths, which were measured with a standard measuring tape to the nearest millimeter. Head

length, width and depth were measured keeping the posteriormost corner of the mouth as the reference point. Body width was measured at the point at which the trunk appeared broadest (most often near the midbody), although some specimens appear a bit flattened due to preservation artifact. Taxonomy Platyplectrurus trilineatus (Beddome, 1867) Lepidosis: Rostral triangular, visible from above, not completely dividing the nasal; nasal five-sided, pierced by nostril; nasal smaller than prefrontal but as large as supraocular; supraocular smaller than prefrontal; frontal as large as distance between it and rostral; parietal subequal in size to that of frontal and prefrontal together; postocular pentagonal, small, smaller than the eye; supralabials 4/4; first supralabial triangular, smallest, in contact with nasal; second supralabial pentagonal, higher than broad, also in contact with nasal; third supralabial six-sided, twice as broad as high, completely bordering the lower rim of eye, in contact with postocular; fourth supralabial largest, a little broader than third, posteriorly twice as high as that of anterior side, not extending backward beyond parietal, but larger than temporal; temporal scale one, rectangular, horizontally elongate, not extending backward beyond the parietal and / or the fourth supralabial; infralabials 4/3-4; all horizontally elongate, first one more or less curved; mental small, subequal to infralabials; no mental groove; gular scales larger than infralabials, rhomboid, the median row of which progressively widens to appear like the proper ventral scales; ventrals 159-183, those in the first one-fifth of the body much less wider than those posteriorly, twice as broad as the adjacent row of costal scales; anal scales bifid; subcaudals 11-16, paired; tail tip bilaterally compressed, covered by somewhat larger scales, ending in a small transverse spur-like structure; overall scalation smooth, scales lacking apical pits, with slight overlap / imbrication, especially ventrally. Habitus: Snout rounded, not depressed, snout length (2.10) more than twice the eye-diameter (0.93); nostril closer to snout tip (0.34) than to eye (2.10); neck not distinct; head-width (3.57) smaller than body width (4.12), but greater than head depth (2.56); head long, 4.7% of total body length; body slender, its width 3.5% the length, subcylindrical, slightly flattened dorso-ventrally, especially in the posterior part; tail short, 6.54 % of total body length, sometimes evident even when viewed dorsally.

S. R. GANESH, 2011


Colouration in formalin: Dorsum light yellowish to brown with one dorsal and two lateral series of darker stripes extending from parietal region on to the tail tip; each series consisting of three lines, that is, a series of dots present on each consecutive scale, forming a dotted line / stripe; stripes feeble in some individuals; venter paler, unpatterned,

extending on to supralabials; dark brown along the edge of most ventral scales, especially in those places where scales overlap; a pair of crescent-shaped spots on nuchal region, bordering the parietals; eyes dark grayish black, pupil mildly visible.

Table 1: Summary of morphological characters of the five examined (preserved) specimens. Measurements in mm. Characters CSPT-S2a.1 CSPT-Sa.2 CSPT-Sa.3 CSPT-Sa.4 CSPT-Sa.5 Mean Total length 103.50 111.20 144.00 107.50 111.00 115.44

Snout-vent length 97.00 106.00 136.00 100.50 105.00 108.9 Tail length 6.50 5.20 8.00 7.00 6.00 6.54 Relative tail length 0.062 0.046 0.055 0.065 0.054 0.056 Head length 5.24 5.27 6.41 5.24 5.28 5.48 Head width 3.47 3.55 3.76 3.52 3.55 3.57 Head depth 2.43 2.45 2.69 2.93* 2.34 2.56 Head length: total length 0.050 0.047 0.044 0.048 0.047 0.047 Head length: snout-vent length 0.054 0.049 0.047 0.052 0.050 0.050 Body width 4.01 3.91 4.51 4.06 4.12 4.12 Body width: total length 0.038 0.035 0.031 0.037 0.037 0.035 Body width: snout-vent length 0.041 0.036 0.033 0.040 0.039 0.037 Eye diameter 0.92 0.95 0.99 0.90 0.92 0.93 Eye-lip distance 0.69 0.74 0.79 0.69 0.70 0.72 Eye-rostrum distance 2.05 2.16 2.21 2.12 1.99 2.10 Eye-nostril distance 1.73 1.75 1.93 1.63 1.77 1.76 Inter-ocular distance 2.15 2.32 2.70 2.59 2.42 2.43 Inter-narial distance 1.27 1.52 1.66 1.40 1.54 1.47 Prefrontal length 1.52 1.49 1.58 1.69 1.30 1.51 Supraocular length 1.40 1.40 1.47 1.49 1.19 1.39 Midbody Scalerows 15 15 15 15 15 15 Supralabial scales 4/4 4/4 4/4 4/4 4/4 4/4 Infralabial scales 4/4 4/4 4/4 4/3 4/4 4/3-5 Postocular scale 1 1 1 1 1 1 Temporal scale 1 1 1 1 1 1 Ventral scales (angulate) 162 159 183 174 170 170 Anal scales 2 2 2 2 2 2 Subcaudal scales (pairs) 11 11 16 16 13 13 Discussion This sample (n=5) has a ventral scale count range of 159-183 (vs. 163-175 in Smith, 1943; 173-177 in Rajendran, 1985 [n=7]). The fact that this sample, consisting of five specimens yielded the largest variation in the ventral count value is remarkable. This can be attributed possibly to the sex of the specimens, but sadly, since these specimens are juveniles, their sex could not be precisely determined. The revised scalation value presented here is largely a result of differences in character definitions in the method of counting ventral scales, which varied in previous works and mine (i.e., Gower & Ablett, 2006). Other characters like midbody scalerows, labials, anal scales and tail-

shield are consistent with previously reported features for this species (see Boulenger, 1890, 1893; Smith, 1943; Rajendran, 1985). However, the size of supraocular relative to prefrontal differed from literature (Boulenger, 1890; Smith, 1943; Rajendran, 1985). Although our specimens agree well with literature descriptions of P. trilineatus, in our specimens the supraocular was not longer than the prefrontal (vs. supraocular longer than prefrontal according to Boulenger, 1890; Smith, 1943; Rajendran, 1985). Boulenger (1890) was the first to synonymize P. bilineatus with P. trilineatus and also the first to distinguish Platyplectrurus species based on ratio of prefrontal and supraocular lengths. But, Beddome (1886) in his original

ON FURTHER SPECIMENS OF Platyplectrurus trilineatus (BEDDOME, 1867)


description of P. bilineatus mentioned “suprtaorbital (=supraocular) as in madurensis.” P. madurensis is a species having supraocular shorter than prefrontal, which is the also case with our specimens; but P. madurensis has a uniform unpatterned dorsum that is very different from the striped dorsum of our specimens. To add further to the confusion, Smith (1943) in his text on P. trilineatus has given figures showing head scalation of P. madurensis although with correct figure-captions. Obviously collection of fresh material and additional data on biology, distribution and species boundaries are needed for a better understanding of this taxon. Acknowledgements I thank B. Vijayaraghavan (Chairman), R. Rajarathinam (Director) and S. Sivakumar (Deputy Director / Environmental Education Officer) of the Chennai Snake Park for the facilities provided. Literature Cited Beddome, R. H., 1867. Descriptions and figures of five new snakes from the Madras Presidency. Madras Quarterly Journal of Medical Science, 21: 14-16+2pl. Beddome, R. H., 1886. An account of the earth snakes of the peninsula of India and Ceylon. The Annals and Magazine of Natural History, 17(5): 3-33. Boulenger, G. A., 1890. The fauna of British India, including Ceylon and Burma. Reptilia and Batrachia. Taylor and Francis, London, UK: 541. Boulenger, G. A., 1893. Catalogue of the snakes in the British Museum (Natural History). Taylor and Francis, London, UK: 485. Daniel, J. C., 2002. The Book on Indian Reptiles and Amphibians. Bombay Natural History Society. Mumbai, Oxford press: 252. Das, I., 2002. A photographic guide to snakes and other reptiles of India. New Holland, UK: 144. Ferguson, S. H., 1895. List of snakes taken in Travancore from 1888 to 1895. Journal of the Bombay Natural History Society, 10: 68-77. Ganesh, S. R., 2010. Catalogue of herpetological specimens in the Chennai Snake Park. Cobra, 4 (1):1-22. Gower, D. J. and J. D. Ablett, 2006. Counting ventral scales in anilioid snakes. Herpetological Journal, 16 (3): 259-263.

Günther, A. C. L., 1868. Sixth account of new species of snakes in the collection of the British Museum. The Annals and Magazine of Natural History, 1(4): 413-429. Hutton, A. F., 1949. Notes on the snakes and mammals of the High Wavy Mountains, Madura district, South India. Part I Snakes. Journal of the Bombay Natural History Society, 48 (3): 454-460. Hutton, A. F. and P. David, “2008” 2009. Notes on a collection of snakes from south India, with emphasis on the snake fauna of Meghamalai Hills (High Wavy Mountains). Journal of the Bombay Natural History Society, 105 (3): 299-316. Inger, R. F., H. B. Shaffer, M. Koshy and R. Brakde, 1984. A report on a collection of amphibians and reptiles from the Ponmudi, Kerala, South India. Journal of the Bombay Natural History Society, 81 (3): 551-570. Ishwar, N. M., R. Chellam and A. Kumar, 2001. Distribution of forest floor reptiles in the rainforest of Kalakkad-Mundanthurai Tiger Reserve, South India. Current Science, 80 (3): 413-418. Kalaiarasan, V., R. Rajarathinam and R. Aengals, 1995. A catalogue of herpetological specimens in Madras Snake Park-Part III: Snakes to Crocodiles. Cobra, 21: 12-15. Malhotra, A. and K. Davis, 1991. A report on the herpetological survey of the Srivilliputhur Reserve Forests, Tamil Nadu. Journal of the Bombay Natural History Society, 88 (2): 157-166. Rajendran, M. V., 1985. Studies in Uropeltid snakes. Madurai Kamaraj University, Publications Division, Madurai, India: 132. Smith, M. A., 1943. Fauna of British India including Ceylon and Burma. Vol-III Serpentes. Taylor & Francis, London: 583. Wall, F., 1919. Notes on a collection of snakes made in the Nilgiri Hills and the adjacent Wynaad. Journal of the Bombay Natural History Society, 26: 552-584. Wall, F. 1920. Notes on a collection of snakes from Shenbaganur, Palni Hills. Journal of the Bombay Natural History Society, 29: 388-398. Whitaker, R., 1978. Common Indian Snakes - A Field Guide. Macmillan press, New Delhi, India: 154. Whitaker, R. and A. Captain, 2004. Snakes of India - The Field Guide. Draco Books, Chengelpet, South India: 481.

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FOUR NEW ASTERINACEOUS MEMBERS FROM KERALA, INDIA Sectional Editor: R. K. Verma Submitted: 16 May 2011, Accepted: 13 June 2011

V. B. Hosagoudar1*, Jacob Thomas1 and D.K. Agarwal2 1 Tropical Botanic Garden & Research Institute, Palode - 695 562, Thiruvananthapuram, Kerala, India * E-mail: [email protected] 2 Plant Pathology Division, Indian Agricultural Research Institute, New Delhi 110 012, India Abstract This paper gives an account of four new members belonging to the genera Asterina and Cirsosia, namely, Asterina aristolochiae, Asterina phyllanthi-beddomei, Cirsosia hopeae and Asterina thunbergiicola Hansford var. indica are described and illustrated. Key words: black mildews, Asterina, Cirsosia, Kerala, India Introduction Peppara and Neyyar is twin and adjacent wildlife sanctuaries located on the western slope and in the penultimate end of the Western Ghats including the Agastyar peak in Thiruvananthapuram District of Kerala state, include the hotspot area of Agastyamala. These sanctuaries lie between 8o 7’ - 8o 53’ N and 70o 4’ - 77o 17’ E, surrounded by Kalakkad and Mundandurai wildlife sanctuaries in the East, Palode and Paruthipalli forest range in the North. Both sanctuaries together have an area of 181 km2, with an altitudinal range from 100–1864 m a.s.l., temperature from 16-35 oC, annual rain fall about 2800 mm. Agastiar peak (1864 m), Pongalapara (1500 m) and Chemunji (1000 m) are the continuous hill ranges, located towards the Eastern side of the study area and steeply descend towards western side. Because of the varied

topography, this area is rich in its plant diversity. We have been studying the foliicolous fungi of this region since 1996 and the present paper is the novelties of this study. Taxonomy

1. Asterina aristolochiae sp. nov. (Pl. 2, Fig. 1) Coloniae amphigenae, plerumque epiphyllae, tenues, ad 3 mm diam., confluentes et patentiae. Hyphae pallide brunneae, undulatae, oppositae vel irregulariter laxe ramosae, laxe reticulatae, cellulae 21-36 x 4-6 µm. Appressoria alternata vel unilateralis, unicellularis, ovata, subglobosa, integra vel sublobata, lata posita, sessilis, 4-12 x 7-12 µm. Thyriothecia laxe dispersa, orbicularis, saepe connata, ad 100 µm diam., stellatim dehiscentes ad centro, crenatae vel fimbriatae ad

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FOUR NEW ASTERINACEOUS MEMBERS FROM KERALA, INDIA


margine, hyphae fringiorae flexuosae; asci pauci vel numerosi, globosi, octospori, ad 43 µm diam.; ascosporae oblongae, conglobatae, brunnae, uniseptatae, constrictus ad septatae, 14-17 x 8-10 µm, parietus echinulatus.

Colonies amphigenous, mostly epiphyllous, thin, up to 3 mm in diameter, confluent and cover almost upper surface of the leaves. Hyphae pale brown, undulate, branching opposite to irregular at wide angles, loosely reticulate, cells 21-36 x 4-6 µm. Appressoria alternate to unilateral, unicellular, ovate, subglobose, entire to sublobate, broad based, sessile, 4-12 x 7-12 µm. Thyriothecia loosely scattered, orbicular, often connate, up to 100 µm in diameter, stellately dehisced at the centre, crenate to fimbriate at the margin, fringed hyphae flexuous; asci few to many, globose, octosporous, up to 43 µm in diameter; ascospores oblong, conglobate, brown, uniseptate, constricted at the septum, 14-17 x 8-10 µm, wall echinulate. Materials examined: type On leaves of Aristolochia tagala Cham. (Aristolochiaceae); Cat. no. HCIO 48252; Loc. Peppara Wildlife Sanctuary, Thiruvananthapuram, Kerala; Coll. Jacob Thomas & Vimalkumar; Date. 18-XI-2007. Isotype, Cat. no. TBGT 2991. Asterina heterotropae Nakamura on Heterotropa hirsutisepala, from Japan and Asterina thotteae Hosagoudar & Hanlin on Thottea spp. from India are reported on the family Aristolochiaceae (Katumoto, 1975; Hosagoudar & Hanlin, 1995). However, the present species differs from both in having unicellular appressoria. 2. Asterina phyllanthi-beddomei sp. nov. (Pl. 2,Fig.2) Coloniae epiphyllae, subdensae, ad 1 mm diam., confluentes. Hyphae flexuosae vel anfractuae, alternata vel irregulariter laxe ramosae, laxe reticulatae, cellulae 28-43 x 3-5 µm. Appressoria alternata vel unilateralis, bi-cellula, subantrorsa vel patentia, recta vel curvula, 9-15 µm longa; cellulae basilares cylindraceae vel cuneatae, 2-5 µm longae; cellulae apicales ovatae vel plerumque globosae, 3-5- stellatim lobatae, 7-10 x 9-12 µm. Thyriothecia dispersa, orbicularis, ad 140 µm diam., stellatim dehiscentes ad centro, margine crenatae; asci numerosi, globosi, octospori, ad 38 µm diam.; ascosporae oblongae, conglobatae, brunneae, uniseptatae, constrictus ad septae, 16-24 x 7-10 µm, parietus glabrus.

Colonies epiphyllous, subdense, up to 1 mm in diameter, confluent. Hyphae flexuous to crooked, branching alternate to irregular at wide

angles, loosely reticulate, cells 28-43 x 3-5 µm. Appressoria alternate to unilateral, two celled, subantrorse to spreading, straight to curved, 9-15 µm long; stalk cells cylindrical to cuneate, 2-5 µm long; head cells ovate to mostly globose, 3-5-times stellately lobate, 7-10 x 9-12 µm. Thyriothecia scattered, orbicular, up to 140 µm in diameter, stellately dehisced at the centre, margin crenate; asci many, globose, octosporous, up to 38 µm in diam.; ascospores oblong, conglobate, brown, uniseptate, constricted at the septum, 16-24 x 7-10 µm, wall smooth. Materials examined: type On leaves of Phyllanthus beddomei (Gamble) M. Mohanan (Euporbiaceae); Cat. no. HCIO 48869; Loc. Peppara Wildlife Sanctuary, Thiruvananthapuram, Kerala; Coll. Jacob Thomas; Date. 27-II-2008. Isotype, Cat. no. TBGT 3245. Two species of the genus Asterina, namely, A. phyllanthicola Sudama Singh and A. phyllanthigena Hosagoudar are known on the host genus Phyllanthus (Hosagoudar, 2004; Singh, 1980). However, the present new species differs from both in having typically lobate head cells of the appressoria. Key to the Asterina species known on the genus Phyllanthus 1. Appressoria lobate …… Asterina phyllanthi-beddomei Appressoria entire …………….……………………. 2 2. Form mycelial net …………...……. A. phyllanthigena Not so ……………..……………….. A. phyllanthicola

3. Asterina thunbergiicola Hansford indica var. nov. (Pl. 3, Fig. 3)

Differt a var. thunbergiicola appressoriis et ascosporis longioribus. Colonies hypophyllous, thin, crustose, up to 5 mm in diameter, confluent. Hyphae crooked, branching irregular at various angles, loosely reticulate to form a mycelial net, cells 21-34 x 2-5 µm. Appressoria alternate, two celled, straight to curved, plugged around stomata of the host leaf, 12-24 µm long, stalk cells cylindrical, 7-12 µm long; head cells ovate, globose to hamate, subangular, angular, narrowly to deeply lobate, 4-12 x 9-15 µm. Thyriothecia scattered, orbicular, often 1-2 connate, up to 180 µm in diameter, stellately dehisce at the centre and dissolved later, margin crenate; asci few to many, globose, octosporous, up to 30 µm in diam.; ascospores, conglobate, brown, uniseptate, constricted at the septum, 16-22 x 7-10 µm, wall smooth. Pycnothyria similar to thyriothecia, smaller; pycnothyriospores pyriform, brown,

HOSAGOUDAR ET AL., 2011


apiculate, broadly rounded at one end and, attenuated and truncate at the other, 16-22 x 9-15 µm, wall smooth. Materials examined: type On leaves of Thunbergia sp. (Thunbergiaceae); Cat. no. HCIO 48870; Loc. Peppara Wildlife Sanctuary, Thiruvananthapuram, Kerala; Coll. Jacob Thomas; Date. 28-II-2008. Isotype, Cat. no. TBGT 3246. Asterina thunbergiicola Hansford is known on Thunbergia chrysops from Sierra Leone, Uganda (Hansford, 1945). However, the new variety differs from the var. thunbergiicola in having longer appressoria and ascospores.

4. Cirsosia hopeae sp. nov. (Pl. 3, Fig. 4) Coloniae epiphyllae, subdensae, ad 2 mm diam. Hyphae rectae, plerumque oppositae acuteque ramosae, laxe reticulatae, cellulae 38-48 x 9-12 µm. Appressoria intercalaris, ovata, saepe leniter lateralis, 9-15 µm diam. Thyriothecia dispersa, ad initio rotundata vel ovata, elongata ad maturitata cum rima longitudinalis ad centre, 300-470 x 250-300 µm, margine crenatae vel fimbriatae, hyphae fringiorae rectae, arte aggregatae et parallel, non-appressoriatae; asci numerosi, globosi, octospori, 35-44 µm diam.; ascosporae obovatae, conglobatae, uniseptatae, fortiter constrictus ad septatae, cinnameo brunnneae, 22-25 x 11-13 µm, parietus echinulatus. Pycnothyria numerosa, thyriotheciis similis; pycnothyriosporae unicellularis, fortiter brunnneae, pyriformes, leniter papillatae, 18-20 x 11-13 µm.

Colonies epiphyllous, subdense, up to 2 mm in diameter. Hyphae straight, branching mostly opposite at acute angles, loosely reticulate, cells 38-48 x 9-12 µm. Appressoria intercalary, ovate, often slightly lateral, 9-15 µm in diam. Thyriothecia scattered, initially round to ovate, elongated at maturity with a longitudinal slit at the centre, 300-470 x 250-300 µm, margin crenate to fimbriate, fringed hyphae straight, closely aggregated and parallel, devoid of intercalary appressoria; asci many, globose, octosporous, 35-44 µm in diam.; ascospores obovate, conglobate, uniseptate, deeply constricted at the septum, cinnamon brown, 22-25 x 11-13 µm, wall echinulate. Pycnothyria many, similar to thyriothecia; pycnothyriospores unicellular, deep brown, pyriform, slightly papillate, 18-20 x 11-13 µm. Materials examined: type On the leaves of Hopea ponga (Dennst.) Mabb. (Dipterocarpaceae); Cat. no. HCIO 48846; Loc. near

Peppara Wildlife Sanctuary, Thiruvananthapuram, Kerala; Coll. Jacob Thomas and Vimalkumar; Date. 31-III-2007. Isotype, Cat. no. TBGT 3222.

Intercalary appressoria, elliptical to elongated thyriothecia with longitudinal dehiscence are the characteristic of the genus Cirsosia. There are five species of the genus Cirsosia are known. Of these, C. arecacearum Hosagoudar & Pillai, C. globuliferae (Pat.) Arx. and C. transversalis Bat. & Maia are known on Arecaceae (Hosagoudar & Pillai, 1993), while, C. irregularis (Sydow) Arx is known on Vatica obtusifolia from Philippines (Müller & Arx, 1962). Cirsosia hopeae differs from it in having epiphyllous colonies in contrast to the hypophyllous, straight mycelium in contrast to crooked, smaller thyriothecia against 750 x 200-300 µm and smaller ascospores 23-25 x 11-12 against 35-36 x 15-16 µm. Acknowledgements We thank Director (TBGRI) for the facilities. We are grateful to Ministry of Environment and Forest, New Delhi for the financial support and to the Forest Department, Govt. of Kerala for the forest permission. Literature cited Hansford, C. G., 1945. Contributions towards the fungus flora of Uganda VII. New records and revisions. Proceedings of Linnean Society London, 157: 20-41. Hosagoudar, V. B. and C. M. Pillai, 1993. Two interesting Cirsosia species on Calamus from India. Mycological Research, 95: 127-128. Hosagoudar, V. B., 2004. A new Asterina species from Kerala, India. Zoos´ Print Journal, 19: 1522. Hosagoudar, V. B. and R. T. Hanlin, 1995. New species of Asterina and Echidnodes from India. New Botanist, 22: 187-192. Katumoto, K., 1975. The Hemisphaeriales in Japan. Bulletin of the faculty of agriculture, Yamaguti University, 26: 45-122. Müller, E. and J. A. V. Arx, 1962. Die Gattungen der didymosporen Pyrenomyceten. Beitrage zur Kryptogamenflora der Schweiz, 11: 1-922. Singh, S., 1980. Asterina phyllanthicola sp. nov. from India. Transactions of the British Mycological Society, 74: 204-205.


BENTHIC MACRO-INVERTEBRATE FAUNA AND “MARINE ELEMENTS” SENSU ANNANDALE (1922) HIGHLIGHT THE VALUABLE DOLPHIN HABITAT OF RIVER GANGA IN BIHAR - INDIA

Sectional Editor: Remadevi Submitted: 28 February 2011, Accepted: 13 July 2011

Hasko Nesemann1, Gopal Sharma2 and Ravindra K. Sinha1

1 Centre for Environmental Science, Central University of Bihar, BIT Campus, Patna–800 014, India 2 Zoological Survey of India, Gangetic Plains Regional Centre, Road No. 11-D, Rajendra Nagar, Patna–800 016, India E-mail: [email protected] Abstract From the main channel of River Ganga 95 invertebrate taxa have been recorded in the endangered Gangetic Dolphin (Platanista gangetica) habitat over an observation period of ten years. Mollusks, Annelids and Arthropods are the dominating benthic groups that constitute the detritivores, filter-feeders and sediment feeders, scrapers/grazers and herbivores. The benthic sediment fauna is rich in diversity and high in abundance. This enables carnivores to occupy a large variety of specialized ecological niches. The qualitative faunal composition of Ganga resembles in general large European rivers with similar representation of taxa. Twelve taxa of marine-originated families were identified, but none of them can be classified as invasive or non-indigenous species. Only two taxa are certainly recognized as non-indigenous neozoans, whereas the remaining fauna shows pristine and stable ecological conditions. In this aspect River Ganga differs from regulated large rivers, where faunal change has largely replaced the original species inventory. Despite the heavy pollution in parts of the river, the original composition of biological diversity is still persisting in the middle reaches of the Ganga. This provides hope for the survival of the Gangetic Dolphin. Key words: Aquatic, invasive species, functional feeding groups, Gangetic Dolphin Introduction The Ganga is the largest river among the rivers originating from the Himalayan region in northern India. The river section in Bihar is one of the few natural and free-flowing large rivers in south Asia.

It is a water resource for one of the world’s most fertile plains with pristine river morphology. This river is an irreplaceable unique habitat for the endangered Gangetic dolphin, Platanista gangetica


BENTHIC MACRO-INVERTEBRATE FAUNA & “MARINE ELEMENTS” OF RIVER GANGA - INDIA


gangetica (Roxburgh 1801) and many other endemic and endangered species. The benthic fauna studied included all groups of taxonomic units to provide comparable results regarding diversity of different habitat-types. Reference data were available for several invertebrate groups only through Datta Munshi et al. (1988), Sharan & Sinha (1988), Sinha (1988) and Subba Rao (1989) from the localities near Patna. Systematic investigations have been conducted for several gastropods e.g. Stenothyridae and Physidae (Sinha & Sharma 2001; Sinha et al., 2003), bivalves (Nesemann et al., 2003, Nesemann et al., 2005) and Annelids (Nesemann et al., 2004). Although throughout most of its range the Gangetic dolphin is declining because of river developments, pollution, deliberate killing and entanglement in nets (Smith and Braulik, 2008) all along the study area there is a good habitat. Five to ten dolphins were regularly surfacing a few meters away from the sampling sites in the main current. Altogether up to 37 dolphins inhabit the River Ganga stretch around Patna (Sinha et al., 2010). Materials and Methods The macro-invertebrate fauna of the Ganga River was investigated frequently along the right bank in the city of Patna. Benthic samples were collected qualitatively using a hand net. Annelid specimens were preserved in 70% ethanol; leeches were usually relaxed in 15% ethanol, and then transferred into 70% ethanol for preservation. Molluscs and decapods were washed from the sediment samples at the spot and if necessary preserved in 4% formaldehyde. Usually only empty shells of large bivalves have been collected and living specimens were released. Study area: The main study area is the right (erosion-) bank of a 4km stretch of River Ganga along the city of Patna from Mahendrughat in the west (25° 37’ 19” N, 85° 09’ 18” E) downstream to Bhadraghat in the East (25° 36’ 40” N, 85° 12’ 35” E). The faunal collection were done from Mahendrughat, seeping springs, Mahendrughat downstream, Adalatghat, Periphyton, Krishnaghat upstream, Gandhighat, Old Palace, Lithal, artificial stone substrate, Old Palace, Phytal: Potamogeton crispus. The research was conducted from 31st January 2000 to 31st January 2011 including frequent field observations. Altogether eight sites have been

visited frequently and their exact results are shown in table 1-3. In addition the left (sedimentation-) bank opposite city of Patna was visited between October and March for collecting faunal samples from different habitats (Boulders, sand, silt and mud substrate). Results The benthic macro-invertebrate fauna of the main channel comprises 95 identified taxa with high diversity of 26 species of annelids (Table 1), 35 species of mollusks (Table 2) and 29 families, genera or species of arthropods (Table 3), The higher crustacean (Malacostraca) fauna includes 8 taxa of crabs, prawns, shrimps, mysid shrimps and one isopod. Aquatic insects are mainly represented with 21 identified taxa out of which nymphs of Dragonflies and Damselflies, larvae of Two-winged flies and adults of Water-Bugs are the most striking groups. Additionally the presence of Roundworms (Nematoda) and Ribbon worms (Nemertina) was generally noticed with small abundances. Functional Feeding Groups of Macro-invertebrates in the River Ganga: The habitat was classified according to longitudinal and lateral terminology described and defined by Illies (1961), Illies & Botosaneanu (1963) and Amoros & Roux (1988). The River Ganga at Patna is a heterotrophic Meta-Potamon system. Organic load is brought from upstream through river-continuum or it is introduced from surroundings along the banks and during flood. According to the commonly used classification of higher invertebrate taxa and field observations, their particular role in processing food can be roughly outlined at least at family level. Each taxon is assigned to a specific functional feeding group based on the definitions of Vannote et al. (1980), Williams & Feltmate (1992), Merritt & Cummins (1996). Functional feeding groups at family level (Tables 1-3) are summarized and figured for 51 taxa found in River Ganga at Patna, based on the qualitative composition of benthic fauna (Fig. 1). The detritivores include in part shredders (Polychaeta) and scrapers or grazers (Gastropoda) with all sediment- and filter feeders. Altogether 54 % of the families can be assigned to this group (Fig. 2). The true carnivores represent 34 % of the qualitative faunal composition, indicating high diversification and prey specialization (Fig. 3). Herbivores (minimum of 2 % or more) are the minor group in the turbid River Ganga, and this may well reflect the rare occurrence of vascular

NESEMANN ET AL., 2011


aquatic plants along the banks. Some scrapers among the gastropods, especially Lymnaeidae, and miners among the insects e.g. Pyralidae nymphs of moths feed mainly on living algae and macrophytes. Figure 1: Qualitative composition of Benthic Macro-Invertebrates of the River Ganga at Patna with 95 identified taxa. Figure 2: General Feeding Guilds of Benthic Macro-Invertebrates in River Ganga with 51 classified families of the present study: Dominance of Detritivores (54 %), Herbivores and others (12 %) over Carnivores (34 %). The “Marine Element” (Annandale 1922): Besides the Gangetic River dolphin Platanista gangetica gangetica, some unique freshwater species of predominantly or nearly exclusively marine invertebrate families have drawn early attention of scientists. Annandale (1922) has already distinguished two groups of marine origin characterizing these as “The Euryhaline Fauna of

the Delta” and “The Relict Fauna of the River”. He listed three bivalves Novaculina gangetica, Scaphula celox and Scaphula deltae as relict fauna. Their occurrence in River Ganga nowadays extends upstream to 1500 km away from coastal waters and the Gangetic delta region. According to Annandale (1922) they are possible marine relics of the former tertiary sea. Figure 3: Functional Feeding Groups of Benthic Macro-invertebrates in River Ganga with 51 classified families of the present study. Presently a total of twelve species of the macro-invertebrates occur at Patna belonging to marine-originated or primary brackish water families. These are Nereididae: Namalycastis indica, Nephthydae: Nephthys oligobranchia, Ozobranchidae: Ozobranchus shipleyi, Stenothyridae: Stenothyra ornata, Gangetia miliacea, Arcidae: Scaphula celox, S. deltae, Psammobiidae: Novaculina gangetica, Mysidae: Gangemysis assimilis, Corallanidae: Tachaea spongillicola, and Hymenosomatidae: Hymenicoides carteri, Neorhynchoplax spp. Recent observations of invertebrate invasion along large rivers used as waterways allow two alternative hypotheses: 1. The occurrence of the above-mentioned species in freshwater upstream from the upper tidal limit are true marine “Relict fauna of the River Ganges” according to Annandale (1922). 2. The occurrence of the above mentioned species in freshwater upstream from the upper tidal limit is based on both recent introduction by shipping and ongoing upstream range extension by active dispersal as a response to the increasing human impact with environmental changes of habitat.



Since the species are restricted to the Ganges River, their presence may be due to the use of the waterway during the last one hundred years. On other hand, some species of “marine origin” are widespread throughout the Gangetic plain. Here the artificial introduction by shipping is most unlikely. All species are native to the Indian subcontinent, eleven of them being originally described from the delta region of River Ganges with type localities somewhere in Hugli River floodplain or connected channels. Two examples will be described here in detail with data of their first collections (Nesemann et al., 2007) and additional records (Nesemann 2009): Tachaea spongillicola (Stebbing 1907) Material examined: Nepal, Rupandehi District, Ghagara Khola, February 1994, leg. S. Sharma & H. Nesemann; Nepal, Rautahat District, Lamaha Khola at Shivpur, February 1994, leg. S. Sharma & H. Nesemann, October 2005, leg. Sharma, Tachamo, Shah, Nesemann; Nepal, Rautahat District, Jhajh Nadi confluence into Bagmati River, October 2005, May 2006, leg. Timalsina, Tachamo, Shah, Nesemann; Nepal, Kailali District, Jagadishpur reservoir, January 2008, leg. Tachamo, Shah, Nesemann; Nepal, Sunsari District, Khantaha River at Kushaha and Haripur, December 1996, leg. S. Sharma, S. Khanal & H. Nesemann; India, Bihar, Ganga River right bank at Patna, Old Palace, March 2003, leg. H. Nesemann; India, Bihar, Pupun River at Fatuha, November 2002, leg. G. Sharma & H. Nesemann; India, Bihar, Ganga River right bank upstream from Buxar, April 2004, leg. D. Kedia, G. Sharma & H. Nesemann. This isopod has been originally described by Stebbing (1907) from a freshwater tank at Kolkata as a commensal of the freshwater sponge Spongilla carteri. The first record of T. spongillicola being collected as ectoparasites of freshwater prawns Macrobrachium spp. in southern India was published by Mariappan, Balasundaram & Trilles (2003). During the present survey T. spongillicola was regularly collected from benthic samples of lowland streams and small rivers of the Lower Gangetic Plain in Nepal and India as well as from River Ganges itself. The wide distribution range exceeds northwards to the Himalayan foothill streams in the Terai region. This may indicate the natural pattern of dispersal. Gangemysis assimilis (Tattersall, 1908) Material examined: Nepal, Rautahat District, Barahwa Nadi north of Gaur, December 2005, May

2006, leg. Sharma, Timalsina, Tachamo, Shah, Nesemann; Nepal, Rautahat District, Jhajh Nadi confluence into Bagmati River, December 2005, May 2006, leg. Sharma, Timalsina, Tachamo, Shah, Nesemann; Nepal, Rautahat District, Jhajh Nadi confluence into Bagmati River, December 2005, May 2006, leg. Timalsina, Tachamo, Shah, Nesemann; India, Bihar, Ganga River right bank at Patna, Old Palace, March 2003, leg. H. Nesemann; India, Bihar, Ganga River at Patna, Adalatghat, March 2008, leg. G. Sharma & H. Nesemann. This mysid shrimp has been originally described by Tattersall (1908) from a brackish water pond at Port Canning. The first records of G. assimilis were collected in small rivers and oxbow lakes of southern Nepal from December 2005 onwards (Nesemann et al., 2007). During the present survey G. assimilis was collected from periphyton-samples from River Ganges itself in March 2008. The wide distribution range northwards to the Himalayan foothill streams in the Terai region resembles the distribution pattern of the isopod T. spongillicola. It supports the natural pattern of dispersal of this species. Discussion How to identify invasive species (Neozoa) among benthic Macro-invertebrates in the River Ganga? Neozoa are numerously reported among the aquatic invertebrates of many large rivers all over the northern hemisphere. The neozoa are benefited from regulated rivers and ships often initially distribute them. The total amount of non-indigenous macro-invertebrates in rivers can be used to understand and describe the degree of anthropogenic changes of the potamocoenosis environment. Many neozoa tolerate higher salinity as well and originate from costal brackish waters. Therefore several neozoa (e.g. the Danubian Limnomysis benedeni Czerniavsky, 1882 of the ponto-caspian basin) have been erroneously regarded as marine relicts. This question is of great interest for the River Ganga fauna. It makes it necessary to review thoroughly the following four aspects for each species: 1. How long back does the knowledge of any

particular species dates? 2. From which country and watershed the species

has been described? 3. Is there any certain observation of the invasive

character of the species? 4. Does the analysis of the present-day distribution

pattern allow any conclusions about the probable impact of transportation by waterways?

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Many taxa of River Ganga that could be identified to species level are most likely a part of the indigenous fauna; based on the knowledge from their first observations. They could have been originally described from either northern Indian subcontinent or from Gangetic delta and certain literature records are known from nineteenth or early twentieth century. For some popular gastropods, the River Ganga is representing the “terra typica” without any precise location e.g. Brotia costula. Early observations of numerous aquatic molluscs have been already mentioned from Gangetic plains by Preston (1915) and Annandale (1922) and additional records were summarized by Subba Rao (1989). Several malacostracans (Mysida, Decapoda) have been originally described from Gangetic delta in Bengal. Similarly some of the wide spread oriental leeches have been already reported from at least few localities of Gangetic Plains by Harding & Moore (1927). Thus the presumed theory of recent faunal changes in River Ganga by invasion or introduction of euryhaline and pollution-tolerant neozoa is not supported with any certain observations. In contrast it has to be highlighted that all members of marine-originated families are autochthonous species of the river since their scientific discovery and description. Among all invertebrates found in the study area at Patna only two species can be certainly identified as non-native invaders, so-called neozoa (Fig. 1). The gastropod Haitia mexicana is of nearctic origin with rapid spreading during last fifteen years. This North American Physidae Haitia mexicana was invading the river system since the early nineties, starting from few introductions in 1994 in capitals like New Delhi and Kathmandu and in 1998 in Allahabad. Its rapid spreading was initiated by commercial distribution of aquaria material and aquatic plants. Shortly after colonization of River Ganga in Patna after 1998, H. mexicana was found in high abundance. Since that time the individual density is declining. Mass occurrence is nowadays restricted to few highly polluted places, where H. mexicana lives without competition with other gastropods (Sinha et al., 2003). The earthworm Perionyx excavatus originates from the Eastern Himalayan foothills (Gates, 1972). This species is helpful in agriculture, with successful early introductions to subtropical countries all over the world. Many pan-tropical localities have been already reported by Gates (1972) from the first half of twentieth century. In Gangetic Plain P. excavatus appears to be well established since long. Thus the

occurrence in semi-aquatic zone of River Ganga at Patna might be the result of natural invasion from agricultural terrestrial habitats. Comparison of River Ganga fauna (Oriental region) with large rivers of temperate Central Europe (Palearctic region): Although numerous faunal lists and inventories have been published for floodplains of large rivers (Obrdlik, Falkner & Castella 1995), detailed studies on the benthic invertebrates of main channels are very rare. Important reason is the different quality of taxa-lists caused by incomparable intensity of research and different levels of identification. Thus only a few studies of large rivers can be compared with the present results of River Ganga. The current identification level for microdrile tube worms (Oligochaeta: Tubificina) and aquatic insects, especially dipterans, varies largely in different studies and it needs a detailed comparison for each individual river. The Gangetic fauna of Patna is compared with some of the largest European rivers Rhine, Main and Oder flowing into the North and Baltic Seas for discussion of differences in Potamocoenosis of Eurasian subtropical and temperate zones. Comparable faunal lists of representative river sections in low altitudes and plains have been rarely published for all macro-zoo benthic organisms. The results investigated by Kinzelbach (1983, 1985), Sopp (1983), Ziese (1985, 1987), Schmid (1999) and Schleuter & Haybach (2003) provide benthic faunal particulars of the main river channels (Table 4). The available physico-chemical parameters show similar condition in the lowland rivers except of the temperature range. In biodiversity all large rivers have three main components of benthic macro-invertebrates: Mollusca (Class: Bivalvia, Gastropda), Arthropoda (Class: Malacostraca, Insecta) and Annelida. Molluscs are more diversified and most dominating in River Ganga with 38 % of identified taxa. The temperate aquatic malacofauna ranges from 21 – 28 %. Arthropods are leading groups in Rhein (Rhine), Main and Oder Rivers with 33 – 51%, but identification level cannot be compared directly due to different species-, genus- and family-level. In contrast the Gangetic insects (23 %) are insufficiently known from the preliminary list at family level. In Annelids all three major groups (Polychaeta, Oligochaeta, Hirudinida) cover 28 % of the identified taxa in River Ganga and the fauna has been carefully investigated and described. The

22



European rivers display similar amount of annelid taxa with 30 % for Order, largely based on the very rich Tubificidae diversity, up to 19 % for Rhine and 17 % for Main River. It is noteworthy that the true freshwater Polychaeta were originally absent in the river systems of North and Baltic Sea. Neozoa in River Ganga compared with the European Rivers (Fig. 4): In regulated large rivers of temperate zone the indigenous or native benthic invertebrates are becoming accompanied by aquatic invaders or invasive species. They are partially replacing the reduced original fauna or even occupy free ecological niches. Neozoa play an important role in benthic fauna of navigable rivers. Their increasing number of species is sufficiently documented in some European rivers and especially studied for River Rhine (Fig. 4). The relative amount of non-indigenous invasive species changes from estimated 2 - 6 % to approximately 12 - 18 % of the total number of identified taxa within the last hundred years (Kinzelbach 1983, Nehring 2003). Similar tendency in Rivers Main (21 Neozoa) and Oder (14 Neozoa) is showing the same range. Figure 4: Presence of Neozoa in River Ganga main channel in comparison with lowland-reaches of large river systems (main channels) in Central Europe, Germany. Discussion of possible reasons for faunal changes in River Ganga: Natural changes are well documented and redrawn from satellite photographs. The river bed permanently undergoes a high natural dynamic process. During the last decade the lateral erosion has shifted the main channel of River Ganga bed northwards. The river became diverted upstream Patna city and the location of the confluence of the Gandak River has changed to a northern direction. The most important visible change of environmental condition is the

fact that the former main channel of River Ganga along Patna has now become a southern branch with residual flow during low water season. Consequently deposition of silt from Ganges and deposition of sand from Gandak have reduced the maximum depth and water current along the right bank of the river. Anthropogenic changes took place along the Patna section during the same time. 1. The organic load has been reduced by sewage

treatment plants as a measurable result of the Ganga Action Plan.

2. The bank fixation originally constructed by standardized bricks was renewed and enlarged with large amount of natural boulders (> 30 cm) along the water’s edge for low water level.

Thus substrate compositions of the right bank are gradually changed. Loam and silt have been reduced or covered and replaced by hard substrate. The quality of hard substrate in form of large natural boulders is providing stable surface habitat for lithophilic species, especially leeches (Hirudinida) and snails (Gastropoda) largely supported by the interspaces and subsurface. These artificially created micro-habitats are stable against flood-disturbance during monsoon period. Lotic/rheophilic species are declining (Novaculina gangetica, Namalycastis indica, Hymenosomatidae), whereas lentic fauna is more supported by the residual flow with side-arm condition during low water period. Especially the increasing density and extension of large freshwater mussels is co related with the environmental changes. Dense mussel-beds were observed in May 2010 at Gandhighat, where ten years ago in January 2000 mostly Thiara scabra and Thiara lineata have been the dominating molluscs. Polychaetes and spider crabs disappeared at the same place. Conclusion Faunal change of benthic macro-invertebrates in River Ganga: Compared with initial studies (Sharan and Sinha 1988, Sinha 1988), the number of taxa is continually growing throughout the last years. In the following decades more detailed studies have been conducted and thorough sampling has brought to light the discovery of many species. On the other hand, the continuous field-observations have clearly documented some faunal change for several groups:

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1. Appearance of invasive non-indigenous species, which have not been recorded before 1998: Physidae: Haitia mexicana.

2. Appearance of indigenous species, which have been overlooked or miss-identified during previous studies: Examples are the Hymenosomatidae: Hymenicoides carteri, Neorhynchoplax sp., Tachaea spongillicola, Gangemysis assimilis.

3. Decline of indigenous species, which have been recorded with high abundance during previous studies: Psammobiidae: Novaculina gangetica.

4. Increase of indigenous species, which have been recorded with low abundance during previous studies: Leeches Hirudinida e.g. Alboglossiphonia sp., Salifa sp. and others.

5. Enlargement and downstream-shifting of the dense mussel-bed within the present study: Unionidae and Amblemidae: Lamellidens spp. and Radiatula spp.

The present scientific knowledge of macro-invertebrates from River Ganga before the twentieth century is very poor. It is limited mainly to collections of molluscs (Preston 1915, Annandale 1922). Many invertebrate species of marine and

brackish origin were described in the early twentieth century and no exact observations prior to their scientific descriptions are available. Thus “The Relict Fauna of the River” (Annandale 1922: 146) “that flourish in the Middle Reaches of Ganges” cannot be certainly assigned to invasive, non-indigenous species. Considering the high number and native character of the marine-originated families, they can be regarded as original members of the eco-region of Lower Gangetic Plains. There is neither proof for their rapid invasion within the last decades, nor for their absence. The few studies on benthic fauna do not allow comparison of the present-day situation with former conditions one hundred years ago. While comparing River Ganga with regulated European rivers the presumed analogous faunal change is not supported. In contrast Ganges fauna has preserved comparatively semi-natural conditions with original species inventory of most invertebrates. The benthic neozoa are actually represented with 2 % of total identified taxa, similar to large European rivers at the end of nineteenth century (Fig. 4). Thus the River Ganga around Patna displays a rich aquatic biodiversity, but the ecological status needs sustainable conservation and improvement of the ecosystem.

Table 1: Taxa-list for Annelida in River Ganga at Patna.

Phylum Annelida Taxon: Records and Abundance (2000-2010) Class Polychaeta

Family Nereididae Namalycastis indica (Southern, 1921) Common, frequent in winter

Family Nephthydae Nephthys oligobranchia Southern, 1921 Nephthys polybranchia Southern, 1921

Badarghat, March 2002 Earlier records: Sharan & Sinha (1988)

Class Oligochaeta Family Naididae

Chaetogaster limnaei bengalensis Annandale, 1905 Branchiodrilus semperi (Bourne, 1890) Allonais paraguayensis (Michaelsen, 1905) Pristina acuminata Liang, 1958 Pristina cf. biserrata Chen, 1940

Badarghat, October 2002 common Mahendrughat, October, November 2002 Mahendrughat, November 2002 Badarghat, March 2002

Dero pectinata Aiyer, 1930 Aulophorus indicus Naidu, 1963 Nais bretscheri Michaelsen, 1899 Nais spec.

Badarghat, March, April 2002 Badarghat, March 2002 Badarghat, February 2003 Mahendrughat, Adalatghat, March 2008

Family Tubificidae Limnodrilus hoffmeisteri Claparède, 1862 Aulodrilus pigueti Kowalewski, 1914 Branchiura sowerbyi Beddard, 1892

Locally very common common Gandhighat, March 2008

Family Microchaetidae Glyphidrilus gangeticus Gates, 1958 Locally very common

Family Megascolecidae Perionyx excavatus Perrier, 1872 Mahendrughat, locally common

Order Hirudinida

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Family Glossiphoniidae Alboglossiphonia weberi (Blanchard, 1896) Alboglossiphonia pahariensis Nesemann & Sharma, 2007 Placobdelloides fulvus (Harding, 1921)

common Gandhighat, May 2010 Widespread but not abundant

Family Ozobranchidae Ozobranchus shipleyi Harding, 1909 Gandak confluence, March 2001

Family Hirudinidae Asiaticobdella birmanica (Blanchard, 1894) Mahendrughat, January, November 2002

Family Salifidae Barbronia weberi (Blanchard, 1894) Salifa lateroculata (Kaburaki, 1921) Salifa biharensis Nesemann, Sharma & Sinha, 2004 Odontobdella krishna Nesemann & Sharma, 2011

Mahendrughat, locally common Widespread but not abundant Mahendrughat, Gandhighat Gandhighat, May 2010

Total number of identified taxa: 26 Table 2: Taxa-list for Mollusca in River Ganga at Patna.

Phylum Mollusca Taxon: Records and Abundance (2000-2010) Class Gastropoda

Family Bithyniidae Digoniostoma pulchella (Benson, 1836) Old Palace, March 2008

Family Stenothyridae

Stenothyra ornata Prashad, 1921 Gangetia miliacea (Nevill, 1880)

Gandak confluence 2001, PM Ghat 2004 Old Palace, March 2003

Family Thiaridae

Thiara (Tarebia) lineata Gray, 1828 Thiara (Thiara) scabra (O. F. Müller, 1774) Thiara (Tarebia) granifera (Lamarck, 1822) Melanoides tuberculatus (O. F. Müller, 1774)

Very common Very common Rare Common

Family Pleuroceridae Brotia costula costula (Rafinesque, 1833) Common

Family Viviparidae Bellamya (Filopaludina) bengalensis (Lamarck, 1822) Mekongia crassa (Benson, 1836) (=Bellamya crassa)

Common Very common

Family Lymnaeidae Lymnaea acuminata (Lamarck, 1822) Krishnaghat, 2003, Gandhighat, May 2010 Radix persica (Issel, 1865) Common

Family Planorbidae

Ferrissia verruca (Benson, 1855) Ferrissia baconi (Bourguignat, 1853) Indoplanorbis exustus (Deshayes, 1834) Gyraulus convexiusculus (Hutton, 1849)

Rare Krishnaghat, 2002 Rare Rare

Family Physidae Haitia mexicana (Phillipi, 1889) Common

Family Succineidae Quickia bensoni (Pfeiffer) Common: Mahendrughat

Class Bivalvia (=Pelecypoda) Family Arcidae

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Scaphula celox Benson, 1836 Scaphula deltae Blanford, 1867

Locally abundant on hard sustrate Rare: Mahendrughat, Old Palace

Family Psammobiidae Novaculina gangetica Benson, 1830 Silt, loam substrate (? declining population)

Family Corbiculidae

Corbicula striatella Deshayes, 185 Corbicula bensoni Deshayes, 1854 Corbicula assamensis Prashad, 1828 Corbicula aurea Nesemann & Sharma, 2007

Common Rare, common only at Badarghat Rare: Mahendrughat, Badarghat Rare: Mahendrughat, Badarghat

Family Sphaeriidae Pisidium (Afropisidium) clarkeanum G. & H. Nevill, 1871 Pisidium (Afropisidium) nevillianum Theobald, 1876

Rare: Gandhighat, common: Badarghat Rare: Badarghat

Family Unionidae Lamellidens corrianus (Lea, 1834) Lamellidens consobrinus (Lea, 1859)

Very common Fairly common

Family Amblemidae

Radiatula caerulea (Lea, 1831) Radiatula occata (Lea, 1860) Radiatula lima (Simpson, 1900) “Radiatula” olivaria (Lea, 1831) Parreysia favidens chrysis (Benson, 1862) Parreysia corrugata laevirostris (Benson, 1862)

Very common Common Rare Common, left bank opposite Patna + upstream Common Fairly common

Total number of identified taxa: 35 Table 3: Taxa-list for Arthropoda (Malacostraca, Insecta) in River Ganga at Patna

Phylum Crustacea Taxon: Records and Abundance (2000-2010)

Class Malacostraca: Order Mysida Family Mysidae

Gangemysis cf. assimilis (W.M. Tattersall, 1908) Adalat Ghat, March 2008 Order Isopoda: Family Corallanidae

Tachaea spongillicola Stebbing, 1907 Old Palace, March 2003 Order Decapoda: Family Palaemonidae

Macrobrachium spec. Old Palace, frequently observed Family Atyidae

“Caridina” spec. Common Family Hymenosomatidae

Hymenicoides carteri Kemp, 1917 Neorhynchoplax spec.

Common, winter saison 2000-2003, 2011 Widespread but not abundant, winter saison

Family Parathelphusidae Barythelphusa lugubris (Wood-Mason, 1871) Parathelphusa martensi (Wood-Mason, 1871)

Locally common Krishnaghat, March 2010 (Exuviae’s)

Class Insecta: Order Odonata Family Gomphidae:

Asiagomphus spec. Macrogomphus spec.

Common, adults March-May Ghajghat, January 2011

Family Libellulidae Gen spp. Gandhighat, May 2010 Family Protoneuridae Gen spp. Old Palace, aquatic plants, March 2008

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Family Coenagrionidae Gen spp. Left bank opposite Patna, Macrophytes Order Trichoptera

Family Hydropsychidae Gen spp. Very common, adults April-June Order Lepidoptera

Family Pyralidae Gen spp. Old Palace, aquatic plants, March 2008 Order Diptera

Family Chironomidae Gen spp. Common Family Ceratopogonidae Gen spp. Gandak confluence, sand substrate

Family Psychodidae Gen spp. Rare: Mahendrughat, March 2008 Family Culicidae Gen spp. Very common Family Muscidae Gen spp. Old Palace, hard substrate

Order Heteroptera: Family Belostomatidae Diplonychus annulatus ( Fabricius, 1781) Old Palace, aquatic plants, March 2008

Family Nepidae: Nepinae: Laccotrephes spec. Ranatrinae: Ranatra spec.

Old Palace, March 2008, Gandhighat, May 2010 Adalat Ghat, Old Palace, March 2008

Family Pleidae: Paraplea spec. Old Palace, aquatic plants, March 2008 Family Micronectidae: Micronecta spec. Very common

Family Corixidae: Sigara (Tropocorixa) spec. Adalat Ghat, Old Palace, March 2008 Family Mesoveliidae: Mesovelia spec. Old Palace, aquatic plants, March 2008

Order Coleoptera Family Gyrinidae Gen spp. Widespread and locally abundant Family Dytiscidae Gen spp. Fairly common

Family Hydrophilidae Gen spp. Fairly common Total number of identified taxa: 30

Table 4: Benthic Macro-Invertebrates of River Ganga main channel in comparison with lowland-reaches of large river systems (main channels) in Central Europe:

Phylum / Class

Subtropical River of Oriental Region (India)

Temperate Rivers of Palearctic Region (Germany)

Ganga Oder Rhein (a) Rhein (b) Main Porifera - - 2 2 3 Bryozoa 1 5 5 6 - Hydrozoa 1 2 2 2 1 Nemathelminthes 1 - 4 - 1 Nemertini 1 - 1 - - Turbellaria - 3 3 3 4 Gastropoda 18 14 14 7 14 Bivalvia 17 15 8 4 15 Polychaeta 3 - - - 1 Oligochaeta 14 28 6 4 5 Hirudinida 9 9 7 6 14 Malacostraca 8 7 7 7 10 Insecta 22 37 19 12 52 Total number of identified taxa 95 120 78 53 120

River Ganga, Lower Gangetic Plain, Patna (India), altitude ≈ 53-55 m a.s.l.; Oder River (Schmid 1999), Lower Oder Valley (Germany), altitude ≈ 28-33 m a.s.l.; Rhein/Rhine (a) River (Ziese 1985, 1987), (a) Mainz – Wiesbaden (Germany), altitude ≈ 83m a.s.l.; Rhein/Rhine (b) River (Sopp 1983), (b) Loreley – St. Goarshausen (Germany), altitude ≈ 75m a.s.l.; Main River (Schleuter and Haybach 2003), Lower Main Plain (Germany), altitude ≈ 85-115m a.s.l..

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Table 5: Physico-chemical parameters of River Ganga (India) compared with Oder, Rhine and Main Rivers (Germany)

Parameters Rivers

Ganga Oder Rhein (a) Main Water Temperature (°C) 14.0-31.2 0.0-26.7 -22.0 -27.0 Conductivity (µS/cm) 229-313 - 475-850 414-872 pH 8.3-8.7 7.1-9.3 7.3-9.0 - Dissolved Oxygen (mg/l) 6.7-9.4 3.3-15.7 5.0-10.0 5.45-13.35

Data sources: Ziese (1987), Sharma (1998), Trzebiatowski (1999), Bernerth et al. (2005), Brahmer & Teichmann, (2008). Acknowledgements Support of K. Prasad (Patna University, India), is duly acknowledged. We are very much thankful to Lee Harding for reviewing the manuscript and Dilip K. Kedia, Ajit K. Singh Patna University) and Samir K. Sinha (WTI) for help during the field work and providing materials. Special thanks are due to Jean-Paul Trilles (Montpellier) and Andreas Allspach (Frankfurt) for identification of Isopoda, and Dirk Brandis (Kiel) for identification of Parathelphusidae. Thanks are also due to the Director (ZSI), Kolkata for the encouragements. Literature Cited Amoros, C. and A. L. Roux, 1988. Interactions between water-bodies within the floodplain of large rivers: function and development of connectivity. Proceedings of the 2nd International Seminar of “International Association of Landscape Ecology”: 125-130. Annandale, N., 1922. The Marine Element in the Fauna of the Ganges. Bijdragen tot de Dierkunde: 143-154. Brahmer, G. and W. Teichmann, 2008. Ein Wärmesimulationsmodell für den hessischen Main. Jahresbericht 2007 des Hessischen Landesamtes für Umwelt und Geologie, Wiesbaden, 31-38. Bernerth, H., W. Tobias and S. Stein, 2005. Faunenwandel im Main zwischen 1997 und 2002 am Beispiel des Makrozoobenthos, Faunistisch-ökologische Untersuchungen des Forschungsinstitutes Senckenberg im hessischen Main. Hessisches Landesamt für Umwelt und Geologie, Wiesbaden, 15-87. Chapman, D., 1992. Water Quality Assessment. A guide to the use of biota, sediments and water in environmental monitoring, Chapman & Hall, London: 585. Datta Munshi, J. S., G. N. Singh and D. K. Singh, 1988. Ecology of Freshwater Polychaetes of River Ganga. Journal of Freshwater Biology, 1: 103-108.

Gates, G. E., 1972. Burmese Earthworms. Transactions of the American philosophical Society New Series, 62: 1-326. Harding, W. A. and J. P. Moore, 1927. Hirudinea. The fauna of British India, including Ceylon and Burma. London, Taylor & Francis: 302. Illies, J., 1961. Versuch einer allgemeinen biozönotischen Gliederung der Flieβgewässer. Internationale Revue der gesamten Hydrobiologie, 46: 205-213. Illies, J. & L. Botosaneanu, 1963. Problèmes et methods de la classification et de la zonation écologique des eaux courantes, considerées surtout du point de vue faunistique. Mitteilungen der internationalen Vereinigung für theoretische und angewandte Limnologie, 12: 1-57. Kinzelbach, R., 1983. Zur Dynamik der Zoobenthon-Biozönosen des Rheins. – Verhandlungen der Gesellschaft für Ökologie (Mainz 1981), Band 10: 263-271. Kinzelbach, R. (Ed.), 1985. Die Tierwelt des Rheins einst und jetzt. Mainzer Naturwissenschaftliches Archiv, Beiheft 6: 85-103. Mariappan, P., C. Balasundaram and J. -P. Trilles, 2003. Infection of Isopod Tachaea spongillicola on freshwater prawns Macrobrachium spp. in southern India. Diseases of Aquatic Organisms, 55: 259-260. Mason, C. F., 1981. Biology of Freshwater Pollution. Longman, Harlow: 250. Merritt, R. W. and K. W. Cummins, 1996. An Introduction to the Aquatic Insects of North America, Third Edition, Kendall/Hunt Publishing Company, Iowa: 862. Nehring, S., 2003. Gebietsfremde Arten in deutschen Gewässern – ein Risiko für die Biodiversität. Schriftenreihe des BMVEL “Angewandte

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Wissenschaft” Heft 498 “Bedrohung der biologischen Vielfalt durch invasive gebietsfremde Arten: 40-52. Nesemann, H., 2009. Aquatic Benthic Macroinvertebrate’ Biological Diversity and their use in Habitat Quality Assessment at the Himalayan Hot Spots of River Ganga Basin. Thesis submitted in partial fulfillment of the requirements for the degree Doctor of Philosophy (Ph.D.). Submitted to the School of Science Kathmandu University (KU), Dhulikhel, Nepal: 212+50 Pls. Nesemann, H., G. Sharma and R. K. Sinha, 2003. The Bivalvia species of the Ganga River and adjacent stagnant water bodies in Patna (Bihar, India) with special reference on Unionacea. Acta Conchyliorum, 7: 1-43. Nesemann, H., G. Sharma and R. K. Sinha, 2004. Aquatic Annelida (Polychaeta, Oligochaeta, Hirudinea) of the Ganga River and adjacent water bodies in Patna (India: Bihar), with description of a new leech species (Family Salifidae). Annalen des Naturhistorischen Museums in Wien 105 B: 139-187. Nesemann, H., S. Sharma, G. Sharma and R. K. Sinha, 2005. Illustrated Checklist of large Freshwater Bivalves of the Ganga River System (Mollusca: Bivalvia: Solecurtidae, Unionidae, Amblemidae). Nachrichtenblatt der Ersten Vorarlberger Malakologischen Gesellschaft, Rankweil, 13: 1-51, Nesemann, H., S. Sharma, G. Sharma, S. N.Khanal, B. Pradhan, D. N. Shah and R. D. Tachamo, 2007. Aquatic Invertebrates of the Ganga River System: Volume 1 – Mollusca, Annelida, Crustacea (in part), Chandi Press: 263. Obrdlik, P., G. Falkner and E. Castella, 1995. Biodiversity of Gastropoda in European floodplains. Archiv für Hydrobiologie, Suppl. 101, Large Rivers 9: 339-356. Preston, H. B., 1915. Freshwater Gastropoda + Pelecypoda. The Fauna of British India including Ceylon and Burma. Francis & Taylor, London: 244. Roxburgh, W., 1801. An account of a new species of Delphinus and inhabitant of the Ganges. Asiatic Researches-Transactions of the Asiatic Society (Calcutta edition), 7: 170+5pls. Schleuter, M. and A. Haybach, 2003. Das Makrozoobenthos des Mains in den Jahren 1992-2001 – Eine Artenliste. Lauterbornia, 48: 45-56.

Schmid, U., 1999. Das Makrozoobenthos des Unteren Odertales – Faunenzusammensetzung und Besiedlungsdynamik in einer Fluβaue. Limnologie aktuell, 9: 317-336. Sharan, R. K. and R. K. Sinha, 1988. Ganga Basin Research Project Buxar-Barh. Final Technical Report (July, 1985-June, 1988), Patna University, Patna. Ganga Project Directorate, New Delhi, Project No. J-13013/24/83-En-I: 88. Sharma, G., 1998. Ecological Study of Gangetic dolphin Platanista gangetica, Ph.D Patna University. Sinha, R. K., 1988. Benthic Macroinvertebrates. In: Sharan, R. K. and R. K. Sinha: Ganga Basin Research Project Buxar-Barh. Final Technical Report (July, 1985-June, 1988), Patna University, Patna: 34-44+2pls, Sinha, R. K. and G. Sharma, 2001. The gastropod Stenothyra ornata Annandale and Prashad, 1921, A New Record from River Ganga in Bihar. Journal of the Bombay Natural History Society, 98 (3): 485-487. Sinha, R. K., H. Nesemann and G. Sharma, 2003. New records of Physa (Gastropoda: Physidae) from Indian subcontinent. Club Conchylia Informationen, 34: 3-11. Sinha, R. K., S. K.Sinha, G. Sharma and D. K. Kedia, 2010. Surfacing and diving behaviour of free-ranging River Ganga dolphin, Platanista gangetica gangetica. Current Science, 98: 230-236. Smith, B. D. and G. T. Braulik. 2008. Platanista gangetica IUCN 2011. IUCN Red List of Threatened Species. Version 2011.1. www.iucnredlist.org Downloaded 12 July 2011. Sopp, E., 1983. Verteilung des Makrozoobenthons im Querprofil des Rheins bei der Loreley. Verhandlungen der Gesellschaft für Ökologie (Mainz 1981) 10: 279-285. Stebbing, T. R. R., 1907. A Freshwater Isopod from Calcutta. Zoological Journal of the Linnéan Society (London) 30: 39-41, Plate 6. Subba Rao N. V., 1989. Freshwater Molluscs of India. Calcutta: Zoological Survey of India: 289. Tattersall, W. M., 1908. The fauna of the brackish ponds at Port Canning, Lower Bengal. Records of the Indian Museum 2: 233-239,Pls 21, 22.

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Trzebiatowski, R., 1999. Occurrence, catches and protection policies of Ichtyofauna in the Lower Oder Valley Landscape Park waters in relation to environmental conditions in 1982-1996, a review. Limnologie aktuell, 9: 387-406. Vannote, R. L., G. W. Minshall, K. W. Cummins, J. R. Sedell and C. E. Cushing 1980: The River Continuum Concept. Canadian Journal of Fisheries and Aquatic Sciences, 37: 130-137. Wiliams, D. D. and B. W. Feltmate, 1992. Aquatic Insects. C. A. B International, Melksham. Ziese, M., 1985. Makrozoobenthon eines Querprofils des Rheins bei Wiesbaden. In: Kinzelbach, R. (Ed.). Die Tierwelt des Rheins einst und jetzt. Mainzer Naturwissenschaftliches Archiv, Beiheft, 6: 85-103. Ziese, M., 1987. Das Makrozoobenthos des Rheins im Bereich von Mainz und Wiesbaden. Mainzer Naturwissenschaftliches Archiv, Beiheft, 7: 1-138.

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PAL ET AL., 2011


REPRODUCTION AND SEXUAL DICHROMATISM IN Sitana ponticeriana (REPTILIA: DRACONINAE: AGAMIDAE) Sectional Editor: Indraneil Das Submitted: 07 June 2011, Accepted: 13 July 2011

Arttatrana Pal1, Mitali Madhusmita Swain1 and Swapnananda Rath2

1 School of Biotechnology, KIIT University, Bhubaneswar, Orissa751024, India; E-mail: [email protected] 2 B. J. B. College, Bhubaneswar 751 014, Orissa, India Abstract This paper presents the results of a two-year study of the sexual dichromatism, breeding, clutch size and development in a population of Sitana ponticeriana, conducted at Balukhand-Konark Wildlife Sanctuary, Orissa, India. Sexual dimorphism appears as a brighter overall colouration with red and dark blue gular pouch develops as outward projection at the neck. In the breeding season, mature males have crimson-red or brick-red colour at the base of the tail. Lizards start courtship in the early morning and afternoon, depending on the ambient temperature, during the early monsoons. Vitellogenic and gravid females occur simultaneously in early and mid-summer, and lay eggs during the onset of rains. Potentially, 2-3 clutches of 6- 23 eggs (mean 17.39 ± 4.39 eggs/clutch) are produced by each female each year. The smallest gravid female contained more eggs than larger ones. Eggs collected late in the reproductive season were significantly larger than those collected earlier. Freshly laid eggs are elliptical; during development, the shape, size and volume of the eggs were changed; hatchling success observed after 39 days of incubation.

Key words: ambient temperature, clutch size, development, Fan-throated lizard, India

Introduction Sexual dimorphism is a widespread feature among lizards, as in many other groups of animals (Andersson, 1994). Male and female lizards may differ in many traits, such as colouration, body shape and size, and these differences have been assigned to the effects of sexual selection (Andersson, 1994). However, other mechanisms,

such as ecological divergence (Schoener, 1967; Shine, 1991) and fecundity selection (Shine & Charnov, 1992) offer alternative explanations for sexual dimorphism (Shine, 1979). Asian agamid lizards of the genus Draco shows distinct morphological patterns of sexual dimorphism in body size and wing size investigated by Mori & Hikida (1992) and Shine et al. (1998). Many of the


REPRODUCTION AND SEXUAL DICHROMATISM IN Sitana ponticeriana


lizards in the Iguanidae, Lacertidae and Agamidae families have complex display behaviors and brightly colored ornaments (Cooper & Greenberg, 1992; LeBas & Marshall, 2000). Australian agamids are no exception with widespread sexual dichromatism (Cooper & Greenberg, 1992), territoriality and highly visually oriented behaviour (Greer, 1989). Information on the breeding biology of Indian Agamid lizards is sparse. Calotes versicolor breeds during the southwest monsoon from late May to October and as in other lizards, its reproductive sequence comprises renewal, breeding and post breeding phases (Shanbhag, 2002). The males are spermatogenetically active from April–September (Radder et al., 2001) and gravid lizards occur from May–October (Shanbhag, 2002). In individuals with variable clutch patterns, both size and frequency of the clutch may be different depending on the proximate climatic factors, food availability, fat reserves and its maternal body size (Ballinger, 1978). Clutch size in Sceloperous virgatus correlates with environmental factors (Smith et al. 1995). Some previous studies indicate that hatchling size is not always correlated with egg size, and that proximate factors such as nesting site and breeding timing may influence this relationship (Kraemer & Bennett, 1981; Packard, 1991; Radder et al., 2002). Prolonged oviductal egg retention occurs in several species of oviparous lizards (Brana et al., 1991). In some of them, embryonic development of eggs continues normally during prolonged egg retention and even continues to hatching (Packard et al., 1977). However, some others such as Urosaurus ornatus (Mathies & Andrews, 1999) arrest development until after oviposition. The garden lizard, Calotes versicolor retains eggs in the oviduct for as long as six months under unfavorable conditions such as a lack of proper oviposition sites or un-optimal rainfall (Shanbhag et al., 2003). The maximum incubation period is 62 days; whereas, under ideal conditions the minimum incubation period is 22 days (Muthukaruppan et al., 1970). Generally, reptilian eggs that incubate at high temperatures hatch earlier than those that incubate at low temperatures (Angilletta et al., 2000; Deeming & Ferguson 1991). Embryonic development of Calotes versicolor stops below 23º C (Radder et al., 2002; Shanbhag et al., 2003). Its eggs have a flexible eggshell and increase in volume after oviposition, with an encroachment in embryonic growth primarily due to infused moisture (Radder et al., 2002).

The fan-throated lizard, Sitana ponticeriana is a medium-sized, ground-dwelling agamid scattered throughout open xeric patches of forest in India (Pal et al., 2010). There are only few studies on the biology of Sitana ponticeriana (Pal et al., 2007; Rajkumar &Shanbhag, 2003, 2004; Rath & Pal, 2007) and its reproductive biology remains subject to speculation because of small sample sizes over limited periods of time. Here, we report on sexual dimorphism, breeding, clutch size and development of Sitana ponticeriana over a two-year period. Materials and methods Study area: This study took place on the Balukhand-Konark Wildlife Sanctuary (BKWS) in the District of Puri, Orissa, India (19º48’ to 19º54’ N, 85º52’ to 86º14’ E) (Pal et al., 2010a). The Sanctuary is a 71.7 km2 area established on a sandy tract along the coast between Puri and Konark. The canopy composition includes mainly Casuarinas sp., Tamarind (Tamarindus indica), Anacardium sp., Karanja (Pongamia glabra), Polanga (Calophyllum inophyllum), Neem (Azadirachta indica), Eucalyptus sp., and Acacia sp. The understory shrubs include Pandanus sp., Adhatoda (Adhatoda vasica), and various horticultural plantings. The lizard fauna includes Lygosoma punctata, Eutropis bibroni, E. macularia and Calotes versicolor. The maximum air temperature is 40ºC during May-June. Maximum surface soil temperature ranges from 40ºC in the shade to over 45ºC in areas with full sun. Minimum winter temperature is 10ºC. Maintenance in the laboratory: We collected lizards by hand from October 2001 through September 2003 and after an acclimatization period of about 2-7 days in the laboratory; we sexed the specimens by examining the gonads and secondary sexual characteristics. We categorized the lizards as juveniles (SVL = 15-26mm), immature (SVL = 27-35mm) and mature adults (SVL ≥ 36 mm above) including gravid females (Pal et al., 2010a,b). We fed the lizards by Isoptera, Hymenoptera, Orthoptera, Acrididae etc. (Pal et al., 2007) and provided water ad-libidum each day. We collected gravid females of S. ponticeriana during the months of May-August (2001 and 2003) exclusively for determining clutch size. We recorded snout-vent length (SVL), tail-length (TL) and total length (TOL) for each lizard. We confirmed oviposition following the methods adopted by Shanbhag et al. (2003). Each terrarium contained a few potted plants, dry sticks and leaves, hiding sites, water and a substratum of sandy soil from the capture site. Gravid females were found

PAL ET AL., 2011


from May to October, consequently, we classified them as early, mid- and late breeders depending on the captured date. After egg laying, we incubated the eggs in moist sandy soil at ambient temperature ~35 ± 2°C). We did not consider second or third clutches in our study because these developed in the female’s body while she was in captivity. After the first clutch, we returned the adult lizards to the sites where we originally captured them. Eggs hatched after 39 days of incubation and >10% of eggs hatched for each clutch. We recorded the SVL, TL and TOL of the hatchlings and then released them into the Sanctuary after 2-3 days. Statistical analysis: We used Chi-square to compare sexual dimorphism and reproductive status between the two years in our study. We identified the reproductive characteristics between early and late in the breeding season with regression analysis. Student’s t-tests were used when the covariant was not obvious. We presented all of the averages ± SE unless otherwise indicated. We used α = 0.01 for decision theory in all statistical tests. Results Sexual dimorphism and dichromatism: Both sexes were olive-brown above and whitish below neck. The dorsum of body is dark-brown with black-edged diamond-shaped rhomboidal patches in both sexes. The gular pouches of males turned brilliant red and dark blue during the breeding season. Males unfolded and folded their pouches several times to entice females during courtship. During the breeding season the tail of mature males are crimson-red or brick-red; whereas, the females have a brown coloured tail. Immature adult males show a white flap-like structure below the neck. However, the mean of SVL, TL and TOL of 256 mature males were 43.51mm ± 5.96, 90.22 mm ± 12.44 and 133.63 mm ± 17.95, respectively. Similarly, the mean of SVL, TL, and TOL of 156 mature females were 43.96 mm ± 6.82, 84.26 mm ± 12.44 and 128.21 mm ± 17.98, respectively. The t-values between SVL, TL and TOL of the males and females were 0.69, 4.69 and 2.95 respectively and were significant at 1 % level (Table 1). The regression equation of males and the females were TL = 1.637 SVL + 0.0072 and TOL = 2.5793 SVL + 0.0248 and TL = 1.7392 SVL + 0.8364 and TOL = 2.7864 SVL+ 0.6843 respectively (Fig. 1a,b,c,d) Breeding: Sitana ponticeriana breed from pre-monsoon through post-monsoon periods (May to September). The peak breeding season was from June through July, but occasionally, an early or late

monsoon stimulated breeding as early as the end of April or as late as September, was recorded. This species required some form of weather conditions such as minimum amount of rainfall and ambient temperature should be < 40ºC before breeding. The maximum number of mature males and the gravid females appeared in April through July. During an early monsoon several mating pairs of the species were active in the field from April through August, but the breeding males were more active than the females. Lizards start courtship in the early morning and afternoon, depending on the ambient temperature, and high ambient temperatures at midday appeared to preclude mating during that time. Interestingly, sometimes mating has been observed throughout the day, on cloudy days. Clutch size: We collected all the adult lizards (n = 86) during October 2001 through September 2003. Lizards were reproductively active as obvious by the presence of either vitellogenic follicles or oviductal eggs or both. The SVL of the smallest female with oviductal eggs was 37 mm, and that of the largest, 58 mm. Clutch size averaged 17.39 ± 4.39 (Table 2) and varied from 6-23 eggs (Fig. 2). Maternal SVL and clutch size was not positively correlated. There was no significant relationship between egg mass and clutch size. Early clutches were larger than those from later clutches. Development: Lizards exhibit prolonged oviductal egg retention when oviposition sites, certain proximate like high ambient temperature and lack of rain falls affect egg-laying behaviour. The egg retention was systematically studied under laboratory condition. The lizards held eggs in their oviducts for about three to four weeks before oviposition. However, their reproductive output was dependent on the climatic factors (e.g., rainfall) that they normally experienced in the wild. We observed in our terrarium, that oviposition was not fixed in a particular place rather it was here and there. It took 10-20 minutes to complete and some lizards (60%) even took 30-45 minutes (2%) even took 30-45 minutes. We found 6-8 eggs in the terrarium that we immediately measured (Table 2), and then placed them under incubation. The eggs were oval or elliptical with flexible shells and coloured white. We measured the length and breadth of those eggs that had not died after 25-32 days of oviposition. Egg volumes increased with embryonic growth up to 32 days. We removed some eggs after 32 days of incubation and observed the growth stage of the lizard. Successful hatching occurred after 39 days of incubation.



Fig. 1a: (A) Plot of TLagainst SVL of mature male Sitana ponticeriana at BKWS Fig. 1b: Regression curves of TOL against SVL of mature male Sitana ponticeriana at BKWS Fig. 1c: Regression curves of TL against SVL of mature female Sitana ponticeriana at BKWS Fig. 1d: Regression curves of TOL against SVL of mature female Sitana ponticeriana at BKWS

Fig. 2: Mature (M) and immature (IM) eggs of dissected gravid Sitana ponticeriana (SVL: 49mm) from BKWS Discussion The evidence for female choice in lizards is at best equivocal (Olsson & Madsen, 1998) and many taxa display remarkable sexual dichromatism with males of many species having ornaments such as rostral horns, dewlaps, gular sacs and crests. This is true of agamids, chamaeleons and their New World sister group, the iguanids. Colour patches may serve as badges of status among males and can affect the outcome of male contests in some species (Olsson & Madsen, 1998). Here, Sitana ponticeriana showed a distinctive red and dark blue gular pouch developed under the neck of male that they used to attract female lizards during the breeding season using the crimson red colour at the base of tail. For instance, there is evidence that size dimorphism may be associated with territoriality, social polygyny and male–male competition in lizards (Olsson & Madsen, 1998; Stamps, 1983). Size dimorphism in lizards can reflect ecological resource partitioning (Magurran, 1998; Perez-Mellado & de la Riva, 1993; Shine, 1979). Our study suggests a negative association between size dimorphism that may reflect mechanisms other than intersexual selection (female choice) in Sitana ponticeriana.

Many previous publications reported the clutch sizes of Calotes versicolor (Erdelen, 1986; Rout & Ghosh, 1984). The present study on the clutch size of S. ponticeriana confirms some previous reports, but contradicts others. Clutch size was larger in our study than in previous reports. Subba Rao & Rajabai (1972) reported that these lizards produce 8–13 eggs with a mean value of 11.2 eggs/clutch (n = 10 clutches, SVL range 36–52 mm), but Tikader & Sharma (1992) reported 11–14 eggs/clutch.

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According to Rajkumar & Shanbhag (2003) the clutch size varies from 7-19 with a mean value of 15.47. In our study, clutch size varied from 6–23 eggs per clutch, but a mean value of 17.39 (n = 88 clutches) was noted. The smaller clutch sizes reported in the previous studies may stem from smaller females, geographic variation, or other environmental factors that regulate clutch size. However, clutch size did not differ between the two years of our study. Clutch size of S. ponticeriana was not increased with maternal SVL, and there was no evidence of any relationship between maternal SVL and egg size. Further, the lack of an inverse correlation between clutch size and egg size suggests that there was no trade-off between these two reproductive traits, implying that there is optimization of egg size. This species produces at least two to three clutches in a breeding season. The early clutches are larger than the later ones as reported in other species of lizards (Shanbhag, 2002, 2003). Decrease in clutch size results from a trade-off between egg size and clutch size (i.e., larger eggs, but few in number) where food availability and modes of energy allocation exist in several species of lizards (Shanbhag, 2003).

The present study on development of S. ponticeriana occurred over 30-39 days in captivity. This species shows polyautochrony because it ovulates many eggs simultaneously from both of its ovaries as also seen in garden lizard (Radder et al., 2001). Eggs of S. ponticeriana increase in volume with embryonic growth following oviposition. The increase in the size of eggs is due to infused moisture (Shanbhag et al., 2003). Further, we speculate that arresting embryonic growth also lessens physiological demands for oxygen and moisture needed by the growing embryo. Shanbhag et al. (2003) demonstrated that embryonic diapauses during prolonged egg retention occurred in Calotes versicolor when its body temperature is lowered by 3–5°C.

Our investigation supports the ambient temperature and rainfall, both which influence the amount of food that increasing energy available for reproduction of this species. Further, low temperature, cloudy skies and wet conditions may interfere with lizard thermal ecology but interestingly, these characters strengthening the reproductive behaviour also resulting in complex interactions with life history of this subtropical lizard.

Table 1: Summary of the sexual dimorphism traits in mature male (N=256) and female (N=156) of Sitana ponticeriana at BKWS, Orissa, India from September 2001-October 2003.

# α = 0.01 Table 2: Summary of the recorded traits in Sitana ponticeriana at BKWS, Orissa, India from September 2001-October 2003

SVL ± SD (mm) TL ± SD (mm) TOL ± SD (mm) Male 43.51mm ± 5.9 90.22 ± 12.44 133.63 ± 17.95

Female 43.96 ± 6.82 84.26 ± 12.44 128.21 ± 17.98 t-Test# 0.69 4.69 2.95

Variable Mean ± SD Range Maternal SVL (mm) 46.68 ± 6.38 37-58 Clutch size (n) 17.39 ± 4.39 6-23 Egg mass (mg) 0.11 ± 0.01 0.08-0.13 Egg length (mm) 9.87 ±1.33 8-11 Egg width (mm) 5.04 ± 0.75 4-6 Offspring SVL (mm) 13.8 ± 2.47 13-15 Offspring TL (mm) 26.6± 1.06 25-28 Offspring TOL (mm) 40.4± 0.89 38-43

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Acknowledgements: We are grateful to Ulrich Manthey, Natalia Ananjeva and John Rudge for reviewing the manuscript and Ajay K Patra (Utkal University, Orissa) for providing laboratory facilities. Literature cited Anderson, R. A., 1994. Functional and population responses of the lizard, Cnemidophorus tigris to environmental fluctuations. American Zoologist, 34: 409-421. Angilletta, M. J. Jr., R. S. Winters, and A.E. Dunham, 2000. Thermal effects of the energetic of lizard embryos: implications for hatchling phenotypes. Ecology, 81: 2957–2968. Ballinger, R. E., 1978. The vertebrate ovary. In: Comparative Biology and Evolution. Jones, R.E. (Ed.). Plenum Press, New York, USA: 789–825. Brana, D., A. Bea, and J.M. Arrayago, 1991. Egg retention in lacertid lizards: relationship with reproductive ecology and the evolution of viviparity. Herpetologica, 47: 218–226. Cooper, W. E., and N. Greenberg, 1992. Reptilian coloration and behaviour. In: Biology of the Reptilia. Gans. C. (Ed.). University of Chicago Press, USA, Vol. 18: 298-422. Deeming, D. C., and M. W. J. Ferguson, 1991. Physiological effects of incubation temperature on embryonic development in reptiles and birds. In: Egg incubation, its effects on embryonic development in birds and reptiles. Deeming, D.C., and M.W.J. Ferguson (Eds.). Cambridge University Press, Cambridge: 147–171. Erdelen, W., 1986. The genus Calotes (Sauria: Agamidae) in Sri Lanka: clutch size and reproductive seasonality of Calotes versicolor- preliminary results. Spixiana, 9: 111-115. Greer, A. E., 1989. The biology and evolution of Australian lizards. Chipping Norton, Australia: Surrey Beatty & Sons. Kraemer, J. E., and S. H. Bennett, 1981. Utilization of posthatching yolk in loggerhead sea turtles, Caretta caretta. Copeia, 1981: 406 – 411. Magurran, A. E., 1998. Population differentiation without speciation. Philosophical Transactions of the Royal Society of London Series B 353: 275-286.

Mathies, T., and R. M. Andrews, 1999. Determinants of embryonic stage at oviposition in the lizard Urosaurus ornatus. Physiological and Biochemical Zoology, 72: 645–655. Mori, A., and T. Hikida, 1992. A preliminary study of sexual dimorphism in wing morphology of the species of the flying lizards, genus Draco. Japanese Journal of Herpetology, 14: 178-183. Muthukkaruppan, V., P. Kanakambika, V. Manickavel, and K. Veeraraghavan. 1970. Analysis of the development of the lizard, Calotes versicolor. A series of normal stages in the embryonic development. Journal of Morphology, 130: 479-490. LeBas N. R. and N. J. Marshall, 2000. The role of colour in signalling and male choice in the agamid lizard Ctenophorus ornatus. Proceedings of the Royal Society B 267: 445-452. Olsson, M., and T. Madsen, 1998. Sexual selection and sperm competition in reptiles. In: Sperm competition and sexual selection. Birkhead, T. R. and A. P., Moller (Eds.). London, Academic Press, London: 503-577 Packard, G. C., 1991. Physiological and ecological importance of water to embryos of oviparous reptiles. In: Egg incubation: its effects on embryonic development in birds and reptiles. Deeming, D. C., and M. W. J. Ferguson (Eds.). Cambridge, Cambridge University press, Cambridge: 213–228. Packard, G. C., R. C. Tracy, and J. J. Roth, 1977. The physiological ecology of reptilian eggs and embryos, and the evolution of viviparity within the class reptilia. Biological Reviews, 52: 71–105. Pal, A., M. M., Swain, and S. Rath, 2007. Seasonal variation in the diet of the fan-throated lizard, Sitana ponticeriana (Sauria: Agamidae). Herpetological Conservation and Biology, 2: 145-148. Pal, A., M. M., Swain, and S. Rath, 2010a. Growth and demography of the fan-throated Sitana ponticeriana (Sauria: Agamidae) from a tropical environment in India. Herpetological Bulletin, 111: 25-35. Pal, A., M. M., Swain, and S. Rath, 2010b. Observation on microhabitat use and activity patterns in Sitana ponticeriana (Sauria: Agamidae). Russian Journal of Herpetology, 17(1): 22-30. Perez-Mellado, V., and I. de la Riva, 1993. Sexual size, dimorphism and ecology: the case of a tropical

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lizard, Tropidurus melanopleurus (Sauria: Tropiduridae). Copeia, 1993: 969-976. Radder, R. S., B. A. Shanbhag, and S. K. Saidapur, 2001. Pattern of plasma sex steroid hormone levels during reproductive cycles of male and female tropical lizard, Calotes versicolor. General and Comparative Endocrinology, 124: 285–292. Radder, R. S., B. A. Shanbhag, and S. K. Saidapur, 2002. Influence of incubation temperature and substrate on eggs and embryos of the garden lizard, Calotes versicolor (Daud.). Amphibia-Reptilia, 23: 71–82. Rajkumar, S. R., and B. A. Shanbhag, 2003. Interrelationships among reproductive traits of female lizard, Sitana ponticeriana (Cuvier). Current Science, 85: 89-91. Rajkumar, S. R., and B. A. Shanbhag, 2004. Factors influencing offspring traits in the oviparous multi-clutched lizard, Calotes versicolor (Agamidae). Journal of Bioscience, 29: 105–110. Rath, S., and A. Pal, 2007. Age determination in Fan-throated Lizard, Sitana ponticeriana (Cuvier). Indian Journal of Gerontology, 21: 1-8. Rout, S. K., and K. C. Ghose, 1984. Nesting and egg laying behavior of the garden lizard (Calotes versicolor). Herpetological Review, 15: 108. Schoener, T. W., 1967. The ecological significance of sexual dimorphism in size in the lizard Anolis conspersus. Science, 155: 474-477. Shanbhag, B.A. 2002. Reproductive biology of Indian reptiles. Proceedings of the Indian National Science Academy, 68: 497–528. Shanbhag, B. A., 2003. Reproductive strategies in the lizard, Calotes versicolor. Current Scienc,e 84: 646-652. Shanbhag, B. A., R. S. Radder, and S. K Saidapur, 2002. Members of opposite sex mutually regulate gonadal recrudescence in the lizard Calotes versicolor (Agamidae). Journal of Bioscience, 27: 529–537. Shanbhag, B. A., K. Srinivas, S. Rajkumar, and S. Radder, 2003. Lowering body temperature induces embryonic diapause during prolonged egg retention in the lizard, Calotes versicolor. Naturwissenschaften, 90: 33–35.

Shine, R., 1979. Sexual selection and sexual dimorphism in the Amphibia. Copeia, 1979: 297-306. Shine, R., 1991. Australian Snakes: A Natural History. Cornell. University Press, Ithaca, New York: 223. Shine, R. and E. L. Charnov, 1992. Patterns of survival, growth, and maturation in snakes and lizards. American Naturalist, 139: 1257-1269. Shine, R., S. Keogh, P. Doughty, and H. Giragossyan, 1998. Costs of reproduction and the evolution of sexual dimorphism in a flying lizard Draco melanopogon (Agamidae). Journal of Zoology London, 246: 203-213. Smith G. R., R. E. Ballinger, and B. R. Rose, 1995. Reproduction in Sceloporus virgatus from the Chiricahua Mountains of Southeastern Arizona with emphasis on annual variation. Herpetologica, 51: 342-349. Stamps, J. A., 1983. Sexual selection, sexual dimorphism and territoriality. In: Lizard Ecology, Studies of a Model Organism. Huey, R. B., E. R Pianka and T. W. Schoener (Eds.). Harvard University Press, Cambridge, Massachusetts, USA: 169–204 Subba Rao, M. V., and B. S. Rajabai, 1972. Reproduction in the ground lizard, Sitana ponticeriana and the garden lizard, Calotes nemoricola. British Journal of Herpetology, 4: 245–251. Tikader, B. K., and R. C. Sharma, 1992. Handbook of Indian Lizards, Director, Zoological Survey of India, (Eds.). ZSI, Calcutta, India.

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MUGGER CROCODILE (Crocodylus palustris Lesson, 1831) PREYS ON A RADIATED TORTOISE IN SRI LANKA Sectional Editor: John Rudge Submitted: 28 June 2011, Accepted: 29 July 2011 Mahesh C. De Silva1, A. A. Thasun Amarasinghe2, Anslem de Silva3 and D. M. S. Suranjan Karunarathna4

1 The Young Zoologists’ Association, Department of National Zoological Gardens, Dehiwala 10350, Sri Lanka 2 Komunitas Konservasi Alam Tanah Timur, Jl. Kuricang 18 Gd.9 No.47, Ciputat 15412, Tangerang, Indonesia E-mail: [email protected] 3 Amphibia and Reptile Research Organization of Sri Lanka, 15/1, Dolosbage Road, Gampola, Sri Lanka E-mail: [email protected] 4 Nature Exploration & Education Team, No: B-1 / G-6, De Soysapura, Morauwa 10400, Sri Lanka E-mail: [email protected] Abstract The Mugger crocodile is the second largest crocodile in Sri Lanka. Recently we observed a Mugger attacking an unidentified radiated tortoise. The body morphology and colouration of this particular tortoise appeared different to the native Star tortoise widely distributed in the country. Thus, the possibility of an escaped or released exotic pet is discussed. Key words: Sri Lanka, Crocodylus palustris, Geochelone elegans, Stigmochelys pardalis, feeding Introduction The Mugger crocodile, Crocodylus palustris is the second largest reptile in Sri Lanka (Deraniyagala, 1953), living in freshwater streams, rivers, canals, tanks and even in hyper-saline estuaries (Deraniyagala, 1930; de Silva, 2010). Currently, the Mugger is categorized as a vulnerable species (IUCN, 2011) and protected by the Fauna and Flora protection ordinance (Act no: 44 of 1964) in Sri Lanka. The Mugger crocodile is also found in India, Pakistan, Bangladesh, Nepal and Iran (Das and de Silva, 2011). Muggers take a wide range of prey (Alderton, 2004; Santiapillai & de Silva, 2001;

Vyas, 2010). The principal diet of the species is insects, crabs, shrimps, fish, frogs, terrapins, other reptiles, small mammals, monkeys, deer, sambars, buffalos, and occasionally humans (Deraniyagala, 1939; Vyas, 1993; Wilson, 1971; de Silva, 2010). Here we report on the attack strategy of a Crocodylus palustris which took place at Bundala National Park (BNP) and details of the radiated tortoise that was attacked. Observation (see plate 4: figs. A-P) On 2nd May 2011, we observed an adult tortoise at

TAPROBANICA, ISSN 1800-427X. April, 2011. Vol. 03, No. 01: pp. 38-41, 1 pl. © Taprobanica Private Limited, Jl. Kuricang 18 Gd.9 No.47, Ciputat 15412, Tangerang, Indonesia.

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11:40 hr. in the Bundala National Park (BNP), southern Sri Lanka. The animal most closely fitted the description of the Star tortoise, Geochelone elegans and had a straight carapace length of around 30 cm. The weather was bright and sunny but the surrounding water bodies were full due to five earlier days of heavy rains. The soil was dry. The tortoise was first seen from a distance of 20 m. and it was about 2 meters from the tank’s edge. This particular tank is known as “Palugaswala (Pathiraja road)” (which is about 50 m long and 30 m wide and elliptic in shape). The tortoise was feeding on vegetation and was looking towards the tank. A Canon 7D camera with 400 mm. lens was used to take photographs. Eight people, including one official from the BNP, witnessed the episode. Around 11:50 hr. a Mugger (total length about 2.5 m) appeared at the water surface about 4 m away from the tortoise and 2 m away from the water’s edge. Within 3 minutes the crocodile had edged slowly to the land and suddenly grabbed the tortoise. The crocodile caught the tortoise from its anterior carapace dorso-ventrally. Just after catching the tortoise the crocodile went back to the water to around 10 m from the bank with the tortoise in its mouth (Pl. 4, Fig. A). Then the crocodile dived and disappeared. After about 30 minutes at 12:25 hr. the tortoise re-surfaced (Pl. 4, Fig. B). The tortoise appeared weak, possibly injured by the crocodile attack and moved slowly towards the land. This movement was more floating than active swimming. At 12:31 hr. the tortoise reached the bank (Pl. 4, Fig. C) without seemingly attracting the crocodile. Suddenly the crocodile re-appeared at the bank from underneath the water (Pl. 4, Fig. D). It approached the tortoise quickly and tried to grab virtually the entire tortoise a number of times. However it failed in all attempts, due to the movements of the tortoise and the shape of the carapace (Pl. 4, Fig. E-I). The tortoise was turned upside down and it tried to right itself but failed due to the mud on the bank (Pl. 4, Fig. J). Surprisingly, considering the crocodile’s huge jaws and powerful bites, we did not observe any damage to the carapace of the tortoise. The crocodile then moved back about 50 cm and remained still for a few seconds (Pl. 4, Fig. K). It then jumped forward and this time grabbed the tortoise from one end of the carapace (Pl. 4, Fig. L-N). After this the crocodile went back into the water about 10 m from the bank, waited for a few minutes on the surface (Pl. 4, Fig. O, P) and then dived and disappeared at 12:34 hr.

We waited there until 1:22 hr. but did not see either the crocodile or the tortoise again. Discussion There are several significant aspects to this episode. Firstly, we did not observe any damage to the carapace of the tortoise, even after the crushing force of the crocodile bite. Secondly, we were surprised how the tortoise survived around 30 minutes under water. According to Bonin et al. (2006), Das (1995) and Jayakar & Spurway (1996), Star Tortoises cannot swim and dive in water. Thirdly, the radiated tortoise in this instance coming towards the tank and feeding on the vegetation at noon in the hot climate of BNP is uncommon. The more usual active foraging times of the land tortoise are in the morning (06:00 to 08:00 hr.) and at dusk when the sun is not hot (de Silva, 2003). Possibly it could have come to drink water. Finally, the distinct carapace, body morphology and colouration of this particular tortoise was quite different to the common Star tortoise (Geochelone elegans), which is the native species widely distributed in the dry zone lowlands of Sri Lanka (Deraniyagla, 1939 & 1953; de Silva, 2003 & 2006) and common in the BNP. Based on the photograph of this animal (Fig. 1) it has been considered by some as Stigmochelys pardalis (personal comm., Indraneil Das and Andy Highfield, July, 2011). It is also possible that it may be an aberrant form of Geochelone elegans (personal comm., Uwe Fritz, July, 2011) or possibly a different foreign species. Fig. 1: The land tortoise consumed by the Mugger crocodile Scanning through the available literature we see an interesting report by Amyrald Haly (the first Director of the Colombo Museum) of purchasing a ‘stared tortoise’ from Wellawatte (Colombo) - the

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A B

A B

person who had collected it stated that it was not the first time he had seen specimens of this particular species. Haly (1894) assumes that this could have been a foreign species that had escaped from captivity or been released after keeping for some time. Haly (1892) also reports of securing a Star Tortoise from a grass field in Cinnamon Gardens (virtually central Colombo). This could have been an escaped pet or one that had been brought and released after some time. Furthermore, we could assume that Europeans who lived in Sri Lanka during the colonial period (up to 1948) would have brought pet tortoises and released them after some time. We have to consider another important fact. That is that many pet traders in the country have been importing the American Red-eared Slider (Trachemys scripta) which is considered as one of the top invaders into many ecosystems in the world and is now known from water bodies in Colombo and its suburbs (de Silva, 1996). Thus, there is a possibility that the same pet traders could have also imported other radiated tortoises. However, due to legal actions taken by the Department of Wildlife Conservation, many people have handed over their pet Star tortoise collections to the Department of National Zoological Gardens (de Silva, 2003) or released them in various parks or forests. It is highly possible that there could be some exotic radiated tortoises in the wild in Sri Lanka. Fig. 2: Geochelone cf. elegans from Puttalam; A. dorsal view, B. ventral view (Photo: Vimukthi Weeratunga) As regards G. elegans, recent molecular work on it from the Indian subcontinent has shown a strong genetic diversity (Gaur et al., 2006). Vyas (2009) has even suggested three discrete sub-populations

of G. elegans, two from mainland India and one from Sri Lanka. See the cover image of this journal and Fig. 2 for a tortoise in an isolated population in Puttalam (northwest Sri Lanka). For comparison, a typical morph of the Star Tortoise from Puttalam and a Leopard Tortoise from Botswana are photographed in Fig. 3 and Fig. 4 respectively; notice the difference in carapace morphology, as well as markings. Thus, we suggest that a molecular study of the Sri Lankan G. elegans from the south, east, north-central, north-west and the north of the country should be conducted. Fig. 3: Typical morph of Star tortoise, Geochelone elegans from Puttalam; A. dorsal view, B. ventral view (Photo: Sameera Karunarathna) Fig. 4: Leopard tortoise, Stigmochelys pardalis from Botswana (Photo: Lee Harding) Acknowledgements We would like to thank Lee Harding and Ralf Sommerlad for reviewing the manuscript. Indraneil Das, Andy Highfield and Uwe Fritz are acknowledged for personal communications. Also the first author thanks Ajith Nishantha and the staff of BNP. Finally Vimukthi Weeratunga and Lee Harding are thanked for photographs.

MUGGER CROCODILE PREYS ON A RADIATED TORTOISE IN SRI LANKA


Literature cited Alderton, D., 2004. Crocodiles and Alligators of the World. Fact on File, Inc and Cassell Illustrated, London: 190. Bonin, F., B. Devaux, and A. Dupre, 2006. Turtles of the world. The Johns Hopkins University Press, USA: 416. Das, I., 1995. Turtles and Tortoise of India. WWF India and the Oxford University Press, Bombay: 176. Das, I. and A. de Silva, 2011. A photographic guide to snakes and other reptiles of Sri Lanka. New Holland, UK: 144. Deraniyagala, P. E. P., 1930. The Crocodiles of Ceylon. Spolia Zeylanica, 16 (1): 89-95. Deraniyagala, P. E. P., 1939. The Tetrapod reptiles of Ceylon; Testudinates and Crocodilians – Vol 1, National Museums of Sri Lanka, Colombo: 412. Deraniyagala, P. E. P., 1953. A Colored Atlas of some vertebrates from Ceylon, Tetrapod Reptilia – Vol 2, National Museums of Sri Lanka, Colombo. de Silva, A., 1996. Proposed action plan: conservation, restoration and management of the Testudines and their habitats in Sri Lanka. Global Environmental Facility Programme: 28. de Silva, A., 2003. The biology and status of the star tortoise in Sri Lanka. Ministry of Environment and Natural Resources: 100 de Silva, A., 2006. Current status of the Reptiles of Sri Lanka, In: Bambaradeniya, C. N. B. (Ed.). Fauna of Sri Lanka: Status of Taxonomy, Research and Conservation. IUCN Sri Lanka: 134-163. de Silva, A., 2010. Crocodiles of Sri Lanka: preliminary assessment of their status and the human crocodile conflict situation. (Report submitted after fulfillment of the project to Mohamed Bin Zeyed Species Conservation Fund) Author, Gampola: 49. Gaur, A., A. Reddy, S. Annapoorni, B. Satyarebala, and S. Shivaji, 2006. The origin of Indian Star tortoises (Geochelone elegans) based on nuclear and mitochondrial DNA analysis: A story of rescue and repatriation. Conservation Genetics, 7 (2): 231–240. Haly, A., 1892. Reptiles. In: The Ceylon Administrative Reports for the year 1891. Government Press. Colombo: 15.

Haly, A., 1894. Reptiles. In: The Ceylon Administrative Reports for the year 1893. Government Press. Colombo: 13. IUCN, 2011. IUCN Red List of Threatened Species. www.iucnredlist.org. downloaded on 06th July 2011. Jayakar, S. K. and H. Spurway, 1996. Contribution to the biology of the Indian stared tortoise, Testudo elegans Schoepff. Journal of the Bombay Natural History Society, 63 (1): 83–114. Santiapillai, C. and M. de Silva, 2001. Status, distribution and conservation of crocodiles in Sri Lanka. Biological Conservation, 97 (3): 305-318. Vyas, R., 1993. Recent cases of man-eating by the mugger (Crocodylus palustris) in Gujarat State. Hamadryad, 18: 48-49. Vyas, R., 2009. The Status of North-Western Populations of Star Tortoise (Geochelone elegans). Chapter 11. IN: Vasudevan, K. (Ed.). Freshwater Turtles and Tortoises of India. ENVIS Bulletin: Wildlife and Protected Areas, Vol. 12 (1). Wildlife Institute of India, Dehradun, India. Vyas, R., 2010. Mugger (Crocodylus palustris) population in and around Vadodara City, Gujarat State, India. Russian Journal of Herpetology, 17 (1): 43-50. Wilson, C., 1971. Crocodiles!. Loris, 12 (3): 151-152.

41


Additions to black mildews of India Kerala is located in the south-west corner of peninsular India and towards the western side of the southern Western Ghats, harbouring rich vegetation. It is a treasure of biological wealth and it needs the specialized personalities to identify and name them. Senior author is being engaged in the study of this never completing treasure of microfungal wealth of Kerala state since three decades. This paper gives an account of four taxa, hitherto unrecorded from India, belonging to the genera Asterina and Meliola, namely, Asterina garciniicola, Meliola abrupta, Meliola bakeri and Meliola hugoniae are described and illustrated. Taxonomy 1. Asterina garciniicola Ouyang & Song, 1995 in Ouyang, Song & Hu, Acta Mycologiaca Sinica, 14: 244. (Pl. 5, Fig. 1) Colonies epiphyllous, dense, velvety, up to 5 mm in diameter, rarely confluent. Hyphae straight to substraight, branching opposite at acute to wide angles, loosely to closely reticulate, cells 14-29 x 7-10 µm. Appressoria alternate to 2% opposite, subopposite, unicellular, globose to cylindrical, straight to curved, antrorse to spreading, entire, 9-14 x 7-10 µm. Thyriothecia scattered, orbicular, often connate, up to 270 µm in diameter, stellately dehisced at the centre, margin crenate to fimbriate, fringed hyphae flexuous; asci globose to ovate, octosporous, up to 65 µm in diameter; ascospores conglobate, uniseptate, constricted at the septum, 31-36 x 14-17 µm. Material examined: On leaves of Garcinia sp. (Clusiaceae); Loc. Pandipathal, Peppara Wildlife Sanctuary; Coll. Jacob Thomas et al.; Date. 29-II-2008; Cat. no. HCIO 48847, TBGT 3223. Asterina garciniae Hansford and A. garciniicola Ouyang & Song are known on the host genus (Hansford, 1946; Ouyang et al., 1995). The latter

species was reported on G. multiflora from China and is reported for the first time from India. 2. Meliola abrupta Sydow & Sydow, 1917. Annales Mycologici, 15 (3-4): 181; Hansford, 1961. Sydowia Beih, 2: 292. (Pl. 5, Fig. 2) Meliola derridis Yates, 1918. Philippine Journal of Science, 13: 368.

Colonies amphigenous, mostly epiphyllous, dense, velvety, up to 2 mm in diameter, confluent and covering entire upper surface of the leaves. Hyphae straight to substraight, branching mostly opposite at acute to wide angles, loosely to closely reticulate, cells 14-38 x 4-7 µm. Appressoria alternate, about 5% opposite, antrorse, curved, 12-17 µm long; stalk cells cylindrical to cuneate, 2-5 µm long; head cells globose to subglobose, straight to curved, entire, 9-14 x 7-12 µm. Phialides mixed with appressoria, alternate to opposite, ampulliform, 16-29 x 6-12 µm. Mycelial setae numerous, scattered, straight, simple, acute to dentate at the tip, up to 520 µm long. Perithecia scattered, verrucose, globose, up to 160 µm in diameter; ascospores cylindrical to ellipsoidal, 4-septate, constricted, 36-43 x 12-17 µm. Materials examined: On leaves of Derris sp. (Fabaceae); Loc. near Peppara dam, Peppara Wildlife Sanctuary; Coll. Jacob Thomas; Date. 18-XI-2007; Cat. no. HCIO 49034, TBGT 3289. This species reported for the first time from India (Hosagoudar, 1996 & 2008). 3. Meliola bakeri Sydow, 1916. Annales Mycologici, 14: 335; Hansford, 1961. Sydowia Beih, 2: 374. (Pl. 6, Fig. 3)

Colonies epiphyllous, scattered, up to 2 mm in diameter. Hyphae straight to undulate, branching opposite at acute angles, closely reticulate, cells 17-24 x 4-7 µm. Appressoria alternate to opposite, antrorse to subantrorse, retrorse to spreading, 17-20 µm long; stalk cells cuneate, 4-7 µm long; head cells ovate, globose, entire, 13-16 x 8-11 µm.

TAPROBANICA, ISSN 1800-427X. April, 2011. Vol. 03, No. 01: pp. 42-43, 2 pls. © Taprobanica Private Limited, Jl. Kuricang 18 Gd.9 No.47, Ciputat 15412, Tangerang, Indonesia.

ADDITIONS TO BLACK MILDEWS OF INDIA


Phialides mixed with appressoria, alternate to opposite, ampulliform, 19-24 x 4-7 µm. Mycelial setae straight, simple, acute to obtuse at the tip, up to 580 µm long. Perithecia scattered, verrucose, up to 128 µm in diameter; ascospores obovoidal, 4-septate, constricted, 30-38 x 13-16 µm. Materials examined: On leaves of Cayrtia pedata (Lam.) A. L. Juss ex Gagnepain (Vitaceae); Loc. Thiruvalla, Pathanamthitta, Kerala; Coll. Jacob Thomas; Date. 19-XI-2006; Cat. no. HCIO 48261, TBGT 3000. This species reported for the first time from India (Hosagoudar, 1996 & 2008). 4. Meliola hugoniae Hanford & Deighton, 1948. Mycological Papers. 23: 5; Hansford, 1961. Sydowia Beih, 2: 91. (Pl. 6, Fig. 4)

Colonies epiphyllous, subdense to dense, velvety, up to 2 mm in diameter. Hyphae straight to substraight, branching mostly opposite at acute to wide angles, loosely to closely reticulate, cells 15-29 x 4-7 µm. Appressoria opposite, about 2% alternate to unilateral, straight to curved, antrorse to spreading, 13-20 µm long; stalk cells cylindrical to cuneate, 2-5 µm long; head cells ovate to oblong, entire, broadly rounded at the apex, 11-15 x 6-9 µm. Phialides mixed with appressoria, alternate to opposite, ampulliform, 17-26 x 5-7 µm. Mycelial setae scattered to grouped around perithecia, simple, straight, acute to obtuse at the tip, up to 1200 µm long. Perithecia scattered, up to 160 µm in diameter; ascospores obovoidal, 4-septate, constricted at the septa, 28-41 x 11-17 µm. Materials examined: On leaves of Hugonia belli Sedgwick (Linaceae); Loc. Aaralam Wildlife Sanctuary, Kannur, Kerala; Coll. Jacob Thomas et al.; Date. 28-II-2007; Cat. no. HCIO 49036, TBGT 3291. This species reported for the first time from India (Hosagoudar, 1996 & 2008). Acknowledgements We thank Director, (TBGRI) for providing facilities. We are grateful to Ministry of Environment and Forest, New Delhi for the financial support and to Forest Department of Kerala for the forest permission.

Literature cited Hansford, C. G., 1946. Contribution towards the fungus flora of Uganda - VVIII. New Records. Proceedings of the Linnean Society of London, 151: 138-212. Hansford, C. G., 1961. The Meliolineae. A Monograph. Sydowia Beih, 2: 1-806. Hosagoudar, V. B., 1996. Meliolales of India. Botanical Survey of India, Calcutta: 363. Hosagoudar, V. B., 2008. Meliolales of India. Vol. II. Botanical Survey of India, Calcutta: 390. Ouyang, Y., B. Song, and Y. Hu, 1995. Studies on the taxonomy of Asterina in China-I. Acta Mycologiaca Sinica, 14: 241-247.

Submitted: 16 May 2011, Accepted: 24 June 2011

Sectional Editor: R. K. Verma

V. B. Hosagoudar1,2, Jacob Thomas1 and D. K. Agarwal3

1Tropical Botanic Garden and Research Institute,

Palode - 695 562, Thiruvananthapuram, Kerala, India 2 [email protected]

3 Indian Agricultural Research Institute,

Plant Pathology Division, New Delhi, India


Two rare butterfly species observed from two isolated forest patches in Kalutara District, Sri Lanka Butterflies are a group of charismatic insects, which forms a major component of Sri Lankan biodiversity (D’abrera, 1998). The total butterfly species recorded in Sri Lanka consists of 244 species and among them 20 (8.1%) are endemic to the island (D’abrera, 1998; Perera & Bambaradeniya, 2006). Overall, 22 butterfly species are critically endangered, 29 endangered and 15 vulnerable and insufficient data are available for rating the status of 29 species (IUCN & MENR, 2007). The distribution and ecology of butterflies, as well as of other insects, has been underestimated or neglected over the last few decades in Sri Lanka. Here we extend the distribution of two endangered butterflies in Sri Lanka. The Southern Duffer, Discophora lepida Moore, 1857 (Lepidoptera: Nymphalidae) from Aduragala forest: The Southern Duffer, Discophora lepida, is the only member of its genus found in Sri Lanka and rated as endangered by IUCN. This rare butterfly inhabits the lowland wet zone forest of Sri Lanka and some areas of Southern India (Woodhouse, 1950) but the subspecies D. l. ceylonica is endemic to Sri Lanka. According to Woodhouse (1950) this species had been recorded from Awissawella, Balangoda, Galle, Labugama, Ratnapura and considered this butterfly a common insect in hilly areas and probably in all bamboo jungle habitats in the south-west of the island. Additionally, this species has been recorded from Madakada Mukalana, Dombagaskanda and Yagirala forest during the last decade by several researchers (Chamikara & Sumanarathna, 1998; Jayasinghe, 2004). Aduragala forest is situated in the low country wet zone of Sri Lanka, between 6º 44’ – 6º 46’ N and 80º 06’ – 80º 08’ E. It is located in an altitude range of 200 to 300 m approximately 10 km away from Horana town and Panadura-Ratnapura main road

crosses through the forest. Part of the Aduragala forest remains in a relatively undisturbed condition and is controlled by the Forest Department while another part is managed as a rubber plantation. The average annual rainfall is about 3660 mm (mainly during the South-west monsoon from June to September). The average annual temperature is 29ºC (IUCN/FAO, 1997). The Aduragala forest contains large trees including Schumacheria castaneifolia, Artocarpus nobilis, Calophyllum inophyllum, Mangifera zeylanica, Humboldtia laurifolia, Oncosperma fasciculatum, and Canarium zeylanicum. The undergrowth is occupied by bamboo trees. According to Gunatilleke & Gunatilleke (1990) the Aduragala forest vegetation can be categorized as a lowland evergreen rain forest. An observation on one individual of the Southern Duffer was made by the naked eye, while standing only 2 m away from the butterfly during 15:10 hr to 15:46 hr. During this period a male Southern Duffer (fig. 1) was observed on 13th May 2006 while it was perched on a leaf of a Schumacheria castaneifolia (kekiriwara in Sinhala) tree about 3 m above ground level. The temperature and the humidity at the time was 28.4 ºC and 68 %. The weather was sunny but inside of the forest was dark. During the 36-minute period the butterfly did not fly beyond the dark area of the forest and we never saw it land on a dead leaf, branch or leaf litter. It only landed on the upper side of leaves. During 20 days we only recorded one individual in the study area. The Southern Duffer has been recorded previously from eight small isolated forest patches in Sri Lanka. Many of these forest patches are also threatened by manmade fire, illegal logging, chena cultivations and using pesticides in surrounding areas. The Common Imperial, Cheritra freja Fabricius, 1793 (Lepidoptera: Lycaenidae) from Atwelthota forest: The members of Lycaenidae represent 34.8% (85 species) of all known butterfly species in Sri Lanka. But yet the ecology of their amazing relationships is not understood. The genus


TWO RARE BUTTERFLY SPECIES OBSERVED FROM TWO ISOLATED FOREST PATCHES


Cheritra in Sri Lanka is represented by a single species (Cheritra freja, Fabricius, 1793). Woodhouse (1952) caught 25 individuals during three week-ends in February and March, adjoining the low country wet zone, at Deniyaya and adjoining the central hills Madugoda and were not rare. According to the IUCN Red list (IUCN & MENR, 2007) this species is endangered. However, the Common Imperial is not common now and it has been recorded in only a few isolated forest patches. This species is also now absent from many areas recorded 50 years back. Atwelthota Proposed Forest Reserve (APFR) is a typical wet zone forest (8º 07’ 03.41” N and 79º 44’ 21.44” E). The APFR is approximately 3 km from the south of Atwelthota junction, Kalutara District in Western Province. The area is currently protected because of its ecological, hydrological and cultural importance. It covers an area of approximately 500 km2, with an average annual rainfall of about 3500 mm. The floral community of the APFR is classified as tropical lowland evergreen rainforest (Gunatilleke & Gunatilleke, 1990). Most of the APFR remains undisturbed by any large scale forest clearance. However, this important forest is threatened by manmade fire, encroachments, illegal logging and chena cultivations. On 19th February 2010 at 13:45 hr we spotted a male Common Imperial (Fig. 2), locally known as ‘Digu-penda Gas-nilaya’, perched on a Cinnamomum sp. leaf about 2 m above ground level. The butterfly was observed from a distance of about 3 m. Conservation Even though Sri Lanka harbors 244 butterfly species, very little knowledge exist regarding their ecology. Large-scale habitat deforestation and fragmentation has led to the decline of various butterfly species of the island, and many species were believed to be common during the early 19th century (see D’abrera, 1998; Ormiston, 1924; Woodhouse, 1950) but now are rare and endangered. Agricultural plantations have replaced indigenous vegetation with monoculture plantations and it has been found that butterfly species diversity is considerably lower than in natural forests. This is probably due to both destruction of natural habitat and extensive use of insecticide and other agrochemicals. Hence restoring of host plants and nectar sources in the boundary plantations and maintaining isolated natural forest patches would be very useful for butterfly conservation.

Fig. 1: Discophora lepida ceylonica male Fig. 2: Cheritra freja male Conservation efforts are hindered by a lack of academic research on butterflies and also there is no up-to-date available published information or baseline data on them. Although inventories exist for several faunal groups in protected areas there are no comprehensive checklists for butterflies. Moreover, only a handful of ecological studies on the butterflies of the island have been undertaken and thus very little technical information is available for conservation managers and policy makers to take steps for effective butterfly conservation. A new generation of research scientists should be trained and supported in the development of projects to protect the future of Sri Lanka’s butterflies and other apects of its natural heritage. Ultimately, conservation of butterflies will need to be accepted through awareness and participation of the general public, commercial interests, and government, in particular school children, undergraduates, farmers and villagers.

KARUNARATHNA ET AL., 2011


Acknowledgements The authors wish to thank Thasun Amarasinghe, Madhava Botejue (TNCS), Gayan Chathuranga, Gayan Pradeep and other members of YZA of Sri Lanka for their help in the field. Literature Cited Chamikara, S. S. and B. S. Sumanarathna, 1999. “Aduragala kanda hewath Mayura handu kanda” (text in sinhala). Parisara Sangrahaya (August – Desember), Society for Environmental Education. D' abrera, B., 1998. The butterflies of Ceylon. Wildlife Heritage Trust, Colombo, Sri Lanka: 224. Gunasekara, S., 2002. Over-exploitation of endangered species. Sri Lanka Naturalist. 5 (1): 11-15. Gunatilleke, I. A. U. N. and C. V. S. Gunatilleke, 1990. Distribution of floristic richness and its conservation in Sri Lanka. Conservation Biology, 4 (1): 21-31. IUCN/FAO, 1997. Designing an optimum protected areas system for Sri Lanka’s Natural Forest. Vol. 2, IUCN and FAO, Sri Lanka: 487. IUCN and MENR, 2007. The 2007 red list of threatened fauna and flora of Sri Lanka. Colombo, Sri Lanka: 148. Jayasinghe, H. D., 2004. Observation of endangered butterfly - Southern Duffer (Discophora lepida) at two wet zone lowland forests. Sri Lanka Naturalist, 6 (1&2): 24-25. Ormiston W., 1924. The butterflies of Ceylon. H. W. Cave & Company, Colombo: 143+VIII Pls. Perera, W. P. N. and C. N. B. Bambaradeniya, 2006. Species richness, distribution and conservation status of butterflies in Sri Lanka. In: Bambaradeniya, C. N. B (Ed.), Fauna of Sri Lanka: status of taxonomy, research and conservation, IUCN Sri Lanka: 53-64. Woodhouse, L. G. O., 1950. The butterfly fauna of Ceylon. Ceylon Government Press, Colombo: 133+XLVIII pls.

Submitted: 29 December 2010, Accepted: 29 June 2011

Sectional Editor: Jeffrey Miller

D. M. S. S. Karunarathna 1,2, R. G. A. T. S. Wickramarachchi1,

D. H. P. U. Silva1 and U. T. I. Abeywardena1

1 The Young Zoologists’ Association, Department of National Zoological Gardens, Sri Lanka; 2 E-mail: [email protected]


Red morph of silvered lutung (Trachypithecus cristatus) rediscovered in Borneo, Malaysia The red morph of Silvered Lutung (Trachypithecus cristatus) a leaf monkey that reportedly occurs or occurred in northeast Borneo, has heretofore been more mythological than real. As far as I can discover, the red morph was only actually observed and reported once, by Allen & Coolidge Jr. (1940). They recorded that seven out of fifteen adult males, but only three out of twenty-six adult females collected at Abai, were reddish. Abai is 47 miles/76 km from the mouth of the Kinabatangan River. This observation was repeated by others (e.g., Groves, 2001; Hooijer, 1962; Payne et al. 1985). Payne et al. (1985), who lived in Sabah, wrote that the red form occurs at the mouth of the Kinabatangan River. In writing the species account of T. cristatus for Mammalian Species, Harding (2010) did not describe the red form for two reasons. First, a pale or orange form of the ebony lutung, T. auratus occurs in southeast Java. It was formerly classified as a subspecies, T. a. pyrrhus (Brandon-Jones et al., 2004), which is not, however, supported by genetic data (Roos et al., 2008). Nevertheless, Brandon-Jones et al. (2004) synonymised T. cristatus with T. auratus on the basis of the similarity of pelage color of some Borneo specimens of T. cristatus with those of T. auratus from east Java. The second reason was that the red or maroon surili (Presbytis rubicunda), another leaf monkey, occurs in Sabah where it is in some locations sympatric with T. cristatus (Davies, 1991). These sources of possible confusion led Harding (2010) and others to doubt whether the red form of T. cristatus actually existed. On May 27, 2011, on a wildlife-watching boat ride on the middle Kinabatangan River at about 5o 25’ 58” N, 117o 44’ 54” E, I saw a troop of about 20 T. cristatus (a typical group size for this species-Amarasinghe et al., 2009), two of which were red.

This is about 250 river km, or 115 straight-line km, up the river. Two smaller groups of T. cristatus on the Kinabatangan River and one on the Klias River in north Sabah did not have any red morphs. The local guides and boat drivers on the Kinabatangan River trip were familiar with regular occurrence of the red morph, which they referred to as “albino” silvered langurs. The said that the maroon leaf monkeys did not occur on this stretch of the Kinabatangan River, but could be found about an hour’s boat drive up a nearby tributary. One of the red ones was a sub-adult: Figure 1 shows it to have the black face of T. cristatus, unlike the blue-grey face of P. rubicunda. It also shows the pointed crest and laterally-extending cheek whiskers typical of T. cristatus, but unlike P. rubicunda. Fig. 1: A sub adult red morph of silvered lutung Figure 2 is one of a series showing a red adult female T. cristatus with a black juvenile clinging to her belly. T. cristatus infants are orange and change to black adult colouration within 3-5 months (Bernstein, 1968), while P. rubicunda infants are whitish with black markings on the face and shoulders. Figure 3 shows a normally-coloured


RED MORPH OF SILVERED LUTUNG REDISCOVERED IN BORNEO, MALAYSIA


adult T. cristatus in the same group. These photos are grainy because of the poor evening light, but are sufficient to establish the presence of the red morph, in about the proportion originally reported by Allen & Coolidge Jr. (1940), but considerably farther upstream on the Kinabatangan River. Hooijer (1968), citing Allen & Coolidge Jr. (1940) called it an "erythristic mutant" and this seems the most likely explanation for the red morph. Fig. 2: An adult female red morph with a black juvenile Fig. 3: a normally-coloured adult silvered lutung

Literature Cited Allen, G. M. and H. J. Coolidge Jr., 1940. Mammal and bird collections of the Asiatic Primate Expedition. Mammals. Bulletin Museum of Comparative Zoology, 87: 131-166. Amarasinghe, A. A. T., W. M. S. Botejue and L. E. Harding, 2009. Social behaviours of captive Trachypithecus cristatus (Mammalia: Cercopithecidae) in the National Zoological Gardens of Sri Lanka. Taprobanica, 1: 66-73. Bernstein, I. S., 1968. The lutong of Kuala Selangor. Behaviour, 32: 1-16. Brandon-Jones, D., A. A. Eudey, T. Geissmann, C. P. Groves, D. J. Melnick, J. C. Morales, M. Shekelle and C. B. Stewart, 2004. Asian Primate Classification. International Journal of Primatology, 25: 97-164. Davies, G., 1991. Seed-eating by red leaf monkeys (Presbytis rubicunda) in dipterocarp forest of northern Borneo. International Journal of Primatology, 12: 119-144. Groves, C. P., 2001. Primate taxonomy. Smithsonian Institution Press, Washington, D.C. Harding, L. E., 2010. Trachypithecus cristatus. Mammalian Species, 42: 149-165. Hooijer, D. A., 1962. Quaternary langurs and macaques from the Malay Archipelago. Zoolologsche Verhandelingen. 55: 3-64. Payne, J. B., C. M. Francis and K. Phillipps, 1985. A field guide to the Mammals of Borneo. Sabah Society and World Wildlife Fund, Kuala Lumpur. Roos, C., T. Nadler and L. Walter, 2008. Mitochondrial phylogeny, taxonomy and biogeography of the silvered langur species group (Trachypithecus cristatus). Molecular Phylogenetics and Evolution, 47: 629-636.

Submitted: 20 July 2011, Accepted: 27 July 2011

Lee E. Harding SciWrite Environmental Sciences Ltd.,

2339 Sumpter Drive, Coquitlam, British Columbia, Canada

E-mail: [email protected]

LETTER TO THE EDITOR Date: 24th May, 2011


RESPONSE TO WARAKAGODA AND SIRIVARDANA, 2009 S.W. Kotagama1, Rex I. De Silva2, A. S. Wijayasinha1 and V. Abeygunawardana1 1 Field Ornithology Group of Sri Lanka, Department of Zoology, University of Colombo, Sri Lanka E-mail: [email protected] 2 Seabird Watch (Sri Lanka), 31, Dampe, Madapatha 10306, Sri Lanka We respond to the article by Deepal Warakagoda and Udaya Sirivardana entitled “The Avifauna of Sri Lanka: An Overview of the Current Status” (2009). In an appendix to this article the writers make several critical comments on the “Avifaunal List of Sri Lanka”, authored by us (Kotagama, et al., 2006). Warakagoda and Sirivardana contend that many species are included in our publication “Which had not been accepted by the CBCRRC [Ceylon Bird Club Records and Rarities Committee], and reports whose contents fall short of the level of authenticity required by a standard rarities committee”. It goes on to say that “A clearly inadequate criterion is used for accepting sight records of rare migrants”. A single example will serve to totally refute their allegations. In 1983 and again in 1990 White-cheeked Tern Sterns repressa were observed in Sri Lanka. The CBCRRC however did not accept that the species occurs in the Island and the Ceylon Bird Club (CBC) Checklist of the Birds of Sri Lanka (Wijesinghe, 1994) relegated the sightings to an appendix of rejected species considered to have been “Reported doubtfully”. In contrast the Field Ornithology Group of Sri Lanka (FOGSL) recognized that the White-cheeked Tern occurs in the Island and included the species in its publications. The writers subsequently included the White-cheeked Tern in their “Avifaunal List of Sri Lanka” (Kotagama et al., 2006). Twenty-five years after the first sighting, a White-cheeked Tern was captured in Bundala on 13 December 2008. The bird was measured, photographed, ringed and released. This clearly substantiated the reliability of our criteria. Images of the bird were disseminated widely in ornithological literature and the internet. Subsequently a second specimen was also captured and ringed. This confirms beyond any reasonable doubt that White-cheeked Terns do visit Sri Lanka. Despite this compelling evidence, the “CBC Sri Lanka List (Country List) of Bird Species” omits the White-cheeked Tern (see CBC website). This raises serious doubts relating to the reliability and validity of the methodology adopted by the CBCRRC where it appears that clearly inadequate criteria, which lack a

level of authenticity, are used for assessing and validating records of rare migrants. It is necessary to clarify that although Warakagoda and Sirivardana’s article may give some readers the (incorrect) impression that the CBCRRC is the ultimate authority on ornithological matters in Sri Lanka, this is emphatically not the case. The CBCRRC does not have any official status outside the CBC (c. 95 members). It does not represent the majority of Sri Lankan ornithologists and its decisions are not recognized by Sri Lanka’s largest ornithological organization, the Field Ornithology Group of Sri Lanka (membership c. 900+). We have outlined clearly the criteria adopted by us in the “Avifaunal List of Sri Lanka” (Kotagama et al., 2006); hence, there should really be no grounds for confusion or unwarranted criticism. The fact is that the FOGSL criteria and list are accepted by the majority of Sri Lankan ornithologists and birders, as well as by global organizations such as BirdLife International. Literature cited CBC web site: http://www.ceylonbirdclub.org/sri-lanka-bird-list.php (Accessed on 22 May 2011) Kotagama, S. W., R. I. De Silva, A. S. Wijayasinha and V. Abeygunawardane, 2006. Avifaunal List of Sri Lanka. In: Bambaradeniya, C. N. B. (Ed.) The Fauna of Sri Lanka: Status of Taxonomy, Research and Conservation. The World Conservation Union and Government of Sri Lanka, Colombo. 164-203. Warakagoda, D. and U. Sirivardana. 2009. The Avifauna of Sri Lanka: An Overview of the Current Status. Taprobanica, 1 (1): 28-35. Wijesinghe, D. P. 1994. Checklist of the Birds of Sri Lanka. Ceylon Bird Club, Colombo: 49.

Taprobanica (2011) Vol. 3. No. 1. Pages 01-49.

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