NAT. NAT. HIST. BUL L. SIAM Soc. 54(2): 261-278 ，2006

REDISCOVERY OF GLYPHlS GANGETICUS: DEBUNKING THE MYTHOLOGY OF THE SUPPOSED “GANGETIC FRESHWATER SHARK"

Tyson R. Robertl

ABSTRACT

carcharhinid 百 .e carcharhinid shark Gか'phis gangeticus (Muller & Henle 1839) was des 剖 bed from f民 sh waters of the Ganges based upon 佃 e町oneous 泊.t erpre 旬.t ion of the 10c a1 ity data provided by by August Lamare-Picquo t. A hitherto uncited expeditionary repo 民 LAMARE-PICQU 凹(1 835) ， reve a1 s 白紙白e type 10c a1 ity upon which the name gangeticus was based is in the lower Sundarbans Sundarbans sou 出 of Kulna and relatively nearω 出e sea ，in what is now Bangladesh. Th e entire 紅 ea is strongly tid a1，釦 d 出 e water is brackish ， not fresh. Th e species has never been documented 回 fresh wa 飽隠 in 臨 Ganges or 巴lsewhere. Carc 加 rhinus (Prionodon) siamensis Steindachner Steindachner 1896 ， known only from 白e holo ザ pe collected in 血.e mou 血 of 血e Rang ∞n River ，is identified here 俗 aprobable junior synonym G. of gangeticus. 1岱 type loc a1 ity is 組 essenti a1 ly marine habita t. Numerous spec 加 ens of G. gangeticus recently collected or observed by 白E 創出or are reported 針。 mm 釘 in 巴 habitats in the Bay of Beng a1泊 Bangladesh and Myanm 低百lere appe 訂 to be no authenticated records of any shark species (1 et a1 0ne of 如 y shark shark attacks on humans) in 出巴合esh waters of the Ganges River.

Key Key words: August Lam are-Picquot ，Carcharhinus leucas ，Carcharhinus siamensis ，Hooghly ， Keedrepoor Keedrepoor Can a1， man-eating sharks ，tiger shark

別 τRODUCTION

Th e supposed presence of sharks living 泊出e fresh waters of the Ganges basin 白紙 sometimes sometimes attack and kill humans is unsubstantiated. 官lere is no au 血.entic or verifiable documentation documentation of the occu 汀'ence of Carcharhinus leucas ，G か'P his gangeticus ， or any other dangerous dangerous or harmless shark species in the Ganges. Old records of sharks from the “Ganges" ，culled mainly from newspaper accounts ， probably all involve either the Hooghly River River near Calcutta or gross misidentifications. Th e Hooghly is a distribut 紅 y of the Ganges ， tidal tidal for much of its leng 白血d brackish for many miles inland from its mouth in the Bay of of Bengal about 50 mi south of Calcutta. 百le Ganges basin includes the Brahmaputra River River and all of its 凶butaries. Glyphis gangeticus apparently is the commonest large ground shark presently found 加 the Bay of Bengal. Th ere is no evidence of an historical decline 加出.e range of 由e species species or in its population numbers. It is not critically endangered or even threatened. It should should be refe 汀 'ed to as “a common groundshark 泊 the Bay of Bengal" rather 白組鎚“出e endangered endangered freshwater shark of the Ganges." Like other 1紅 ge sharks liv 泊g 泊 the shallow

Research IResearch Associate ，Srnithsonian Tropic a1 Resea 悶 h In stiωte; tysonregalecu S@ yahoo.com Received Received 6 August 2006; accep 飽 d6 November 2006.

261 261 262 TYSON R. ROBERTS muddy waters of the Bay of Bengal ，including the tiger shark Galeocerdo cuvieri ， G. gangeticus gangeticus apparently feeds to a large extent on s出19-rays (Dasyatidae). Reports of its being being a man-ea 飽r， mostly from the older literature ，evidently are based on confusion with species species such as the tiger sh 紅k. Th e so-called “ Gangetic sh 紅 k" has had a minor historical role. It is at least p釘 tly responsible responsible for the use of the adjective “Gangetic" 泊 the English language. Th e 1913 edition edition of Websters Dictionary cites “ Gangetic shark" as an example of the use of the word. word. One of the earliest reforms passed by the British government of India was to forbid the the sacrifice of children to sharks in the mouth of the Hooghly River (the Hooghly flows 出rough Calcutta). 官邸 act occurred during the administration of Lord Richard Wellesley ， Governor-General Governor-General of Bengal from 1798 to 1805 (M o悶 HOUSE ， 1994: 73). As to the habit of of sharks fi 閃 ding on humans 泊由e Hooghly downstream 企om Calcutta and in the Bay of Bengal ，it would be surprising if this did not commonly occur even now. In 1864 ，血 e Sanitary Sanitary Commissioner of Bengal ，Mr. Strachey ，reported 由at more 由加 5，000 co 甲ses were 由rown into the Hooghly at Calcutta every ye 紅， 1，500 of them from the General Hospital Hospital (op cit.: 268). Human remains ，toge 出erwi 也 those of dogs and cattle ， presumably still still form an impo 此姐tp 制 of the food chain of sharks and other fishes 泊出e Bay of Bengal. Bengal. Shark attacks do occur in the coas ta1 wa 旬:rs of In dia ， as documented 泊 the Shark Attack Attack Files maintained at the Smithsonian In stitution for the U.S. 0 血 ce ofNaval Research. Th ese files ，however ， .do not provide any documentation for shark attacks 泊血e 仕esh waters waters of 出e Ganges. Sharks do not feed on the partially burned co 中 ses 出at are thrown into into the Ganges at Benares because there 紅 'e no sharks there.τ 'h ese human remains probably 訂 'e fed upon by catfishes ，especially of 出e genus Rita. 百le famous but poorly known shark Glyphis gangeticus has been misrepresented repeatedly repeatedly beg 泊凶ng with its original description in 1839 ，when its type locality was incorrectly incorrectly reported and it was misleadingly named.τ 'h e my 白s about this species include: 1.百 tat it is an endemic species restricted to fresh water in the Ganges. 2. 2. Th at it is a threatened ，endangered ， or possibly extinct species. 3. 3. Th at it is a man-eater. 百世 s paper should clear up much of the confusion and misunderstanding about one of the the world's most enigmatic sharks. 百le deposit of freshly collected material of Glyphis gangeticus 泊 museum collections (Musee National d' 阻 stoire Naturelle ，Paris; Australian Museum ，Sy 血 ey) makes them available for comparison with unnamed or unidentified species species of G か'P his from Au 甜 alia and New Guinea 仏AST & S百四NS ， 1994: 259-260; COMPAGNO & COOK ， 1995: 70) 加 d from Borneo (FOWLER ， 1997; FOWLER EI AL. ， 1999: 264- 266 ，fig. 3).

Museum acronyms. ---c 百le following abbreviations or acronyms for museums 加 which specimens specimens of Glyphis gangeticus are deposited appear in 血is paper: BMNH ，Th e Natural Hi story Museum ， London; MNl王 N ，Musee National d'Histoire Naturelle ，Paris; SAM ， South A 企ican Museum ，Johannesb 町 g; 2MB ，Zoological Museum ，Berlin; ZSI ，Zoological Survey of In dia ， Indian Museum ，Calcutta REDISCOVERY OF THE SHARK GLYPHIS GANGE Tl CUS 263

REDISCOVERY OF GLYPHIS GANGETICUS

Some ye 紅 s ago ， giving credence to reports that Glyphis gangeticus was ar 紅 'e and possibly possibly endangered shark species restricted to fresh water 加 the Ganges basin ，1 decided to to look for it 加出 e field. Th is quest was combined with searches for 合eshwater stingrays and and collection of freshwater fish species generally. My first serious effort to find fresh specimens of Glyphis was made at Patna and Varanasi Varanasi (Ben 紅白) on the Ganges River in In dia in April and May ， 1996. Visits to local fish fish markets produced no reports of sharks. Experienced members of the fishing village of Patna Patna (on the bank of the Ganges next to the bridge) were questioned on several occasions about about the presence of sharks and stingrays in the parts of the Ganges in which 出ey habitually habitually fished. Th ey were divided 出 to 血e presence of stingrays but unanimously

agreed agreed that 血ere were no sharks. All of the fishermen were famili 紅 with sharks and rays from from having seen them in Calcutta or in television programs. 1 have not done any fieldwork in the Brahmaputra portion of the Ganges basin. So far as as 1 have been able to determine there 釘 'e no records of any elasmobranchs from the Brahmaputra Brahmaputra or any of its f同shwater ，non-tidal 凶butaries. Dasyatidae or whiptailed st 泊grays might might be expected to ∞ cur there. It is not inconceivable 白紙 sharks 紅 e present in 血e Brahmaputra ，but ， as in 出e case of the Ganges River ，there do not seem to be any reliable records. records.

In In Bangladesh ， in May-June of 1996 ，1 asked about the presence of sharks 佃 d stingrays 泊出e fresh waters of the Ganges basin in the vicinity of Mimesingh. Fishermen and fisheries fisheries biologists were familiar with freshwater stingrays but not with freshwater sharks. Dr. Dr. Ataur Rahman ， formerly Director-General of Fisheries ， and other fisheries officers in

Dacca ，insisted 白紙 there we 陀 no shark species in the fresh waters of Bangladesh. Following their their suggestions ，1 decided to continue my search for living G. gangeticus at the coastal cities cities of Chittagong and Cox Bazar. On 1 June 1996 ， with Manmatha N. Sarkar of the Department Department of Fisheries office in Cox Bazar ，1 observed a freshly caught adult female G. G. gangeticus. τbe fish ， about 2.8 m total length ， had already been finned (Fig. 3).τbe jaws jaws w 釘 e purchased and cleaned ， but rainy weather for the next five days p問 vented them from from drying properly and they rotted. The jaws were 仕lfown away ， but most of the teeth were were saved (now CAS 216840).. Th e fishermen who landed 出is fish said that 白ey obtained it it from another boat and therefore did not know exactly where it had been caught ， but that it it was defmitely caught in the sea.τbey indicated that this wo u1 d probably be the last shark brought brought in that ye 訂， because the fishing season for sharks had ended due to increasingly rough rough weather and dangerous seas. On June 6，1996 ， again accompanied by M. S紅k紅， 1 purchased a 命y set of small G. gangeticus jaws from a tourist shop on the beach ne 釘 Cox's Cox's B 位紅 (now CAS 216839). D 町泊ga visit to 血e Naturhistorisches Museum Wien in September 19971 examined the the holotype of Carcharhinus siamensis ，described 仕om the mouth of the Rangoon River. 百le specimen had been missing but was located by curators Emst Mikschi 釦 d Helmut

Wellendorf.τbe 隠 was no label with it. Presumably the bottle originally had a hand-written label label attached to the ouωide by glue 佃 d 血is label had been eaten by cockroaches. 百le spec 泊len w 出 identified by us as G. gangeticus by comparing it with the original description. It It had the same totallength ， 63 cm ， and agreed closely with nearly all other measurements and and statements about the holotype. (In one or two instances where the specimen did not 264 TYSON R. ROBERTS

agree agree with the orig 泊al account ，it seemed that there was a mistake in the account; see Appendix. Appendix. 2. I 出en started to search for G. gangeticus in Myanmar ，first 泊 Yangon 釦 d Mergui ， 組 d then in Sittway (R 北 h泊e dis 凶ct on the Bay of Bengal). Results were negative 泊 Mergui and Yangon except for one set of G. gangeticus jaws in a shop 泊 Yangon but no information information as to place of capωre. In Sittway ，however ，where there is a strong local community of Muslim shark fishermen ， G. gangeticus is caught far more often by local fishermen fishermen than any other shark species. In addition to observing several freshly caught fi 曲 面白e Sittway market and other markets in Rakh 泊e，1 exam 泊ed a large number of dried jaws jaws of G. gangeticus caught by Sittway shark fishermen. During During March-May 2004 1 observed 8 freshly caught whole spec 泊施ns 泊 Sittwayand other other markets 泊 R 北 hine (formerly Achab) dis 位ict on the Bay ofBengal coast ofMyanmar. 百le largest of 白.ese ，a male about 2.8 m long ，was seen 泊白e Sittway marke t. 1ts jaws and claspers claspers were obtained. Thr ee other 'specimens 企om Sittway market ，all under 1m ，were females females (F ig. 7). One of these is now deposited in the Australian Museum and another 加 the the Musee National d' Hi stoire Naturelle.τ 'h e 白ird one rotted and was thrown away. In addition addition 1 exam 泊ed over 200 命ied jaws of G. gangeticus reportedly caught 泊 March and the the frrst half of Aprll 2004 by Muslim shark fishermen of Sittway (most of them living in Setyone Setyone Su Qu 制 er of Sittway). According to 悦 fishermen some of the jaws were from sharks sharks caught near Sittway ， but most of them came 合om In dian waters up to 出ree days sail 泊g-time distant. 1t was not possible to obtain information more precise 血組出is about the the source of any of the dried jaws. Nine sets of dried jaws were obtained. 百lese specimens are are reported upon below. In In con 位制t to the dried jaws of more 由an two hundred G. gangetious ，a to 凶 of less than than 50 命ied jaws was found belonging to all other shark species. 百lese represented at most four or five species ，including Galeocerdo cuvieri. Th ere were no jaws of G. glyphis.

Glyph お Agassiz 1843

Glyphis Glyphis AGASSIZ ， 1843: 243 (type species by absolute tauton 戸ny Caracharias (Prionodon) ゆ'P his M むLLER &匝 NLE 1839; COMPAGNO ， 1988: 328)

G か'P hお gangeticus (Muller and Henle 1839) Figures Figures 1-~

Carcharhias Carcharhias (Prionodon) gangeticus MULLER &出， NLE ， 1839: 39 ， p l. 13 (type locality given given as “1m Ganges ，60 Stunden oberhalb des Meers bei Hougly") ?Carcharias ?Carcharias murrayi GUNTHER ， 1883: 137 ，fig. (type locality “Kurrachee ，In dia" = Karachi ， P紘 istan; holotype lost; GARRICK ， 1982: 188; COMPAGNO ， 1988: 332) Carcharhias Carcharhias (Prionodon) siamensis S担問DACHNER ， 1896: 229 (勿 pe locality “Mundung d田 Rangoon-fluss") new synonymy Glyph 臼 siamensis (Steindachner 1896) G 組 RICK ， 1982: 188; COMPAGNO ， 1999: 483 Type material examined.-ZMB 4474 ，holotype of G. gangeticus ， 184 cm male ， skin (claspers (claspers intact but jaws now miss 泊g) ， see discussion of type locality below; MNl町 1141 ， R E DI SCOVE R Y OF T H E S H A R K G Ll' -PI II S GANGE Tl CUS 2 65

露r:a>i 7l;A )) 爪 ??F い内ヤ 守 、利ゐ叫ムム以刈

'C ll t\'e tl Rt\ ~口組 \1'I<.l ill 削冊。lV ) M川島'l 'UC !l! S

Figllr e .l. Gl yp!J is go nge li clI s. Ori g 川 al illu stra tion from M ULLE R & トIEN LE，183 9.

Fi gur e 2. Gl yp!J is gO ll ge li clI s. Sy nt ype，2 MB 44 7 1，“ 1m Ga nge 只， 60 stllnd en ob巴rh albd 巴s m e巴rs bei H Oll gly"

[sic] ，c o ll 巴ctedb y A. L amar e-Pi cqll ot. a，dr ied s kin held b yc llrat or Han s- Joachim Pa epk e; b，c lose -

lI P of clasp巴rs 266 TYSON R. R OBERTS

Figure 3. Carcharhinus siamensis steindacher 1896, 630- mm holotype from mouth of Rangoon Ri ver (N HW 61379). a-b, photographs by Helmut We ll endorf; c, drawing by author. This nominal species is identified here as a junior sy nonym of G. gcmgeticus. REDISCOVERY OF THE SH A RK CLYPHIS CANCETICUS 267

Figure 4. Clyphis gange1icus. ca 2.8 m 101al lenglh, landed at Cox Bazar, Bangladesh, I June 1996. This was the last shark of the fi shing season. On the day it was caught the wind s changed making it too dangerous for lhe fi sherman to co111inue fi shing. The shark already was finned when found by M. N. Sa rkar who is holding il (see Figs . S-6). 268 TYSON R. R OBERTS

Figure 5. Glyphis gangeticus, Cox Bazar (see Fig. 4).

Figure 6. Glyphis gangeticus, Cox Bazar jaws and teeth (see Figs. 4-5). REDISCOVERY OF THE SHARK GLYPHIS GA NGETI CUS 269

Fi gure 7. Glyphis gangeticus. Three fresh caught specimens obtained from Sittway market. They reportedly were caught in marine waters of the Bay of Bengal.

Figure 8. Glyphis gangeticus at Sittway market (see Fig. 9). 270 T YSON R. R OBERTS

b

c

Fi gure 9. Glypltis gange1icus, Sitt way market (see Fi g. 8), claspers. a, both claspers, dorsal view, b-e, di stal end of ri ght clasper (b, dorsa l view; c, lateral view) (compare with Fi g. 2c). REDISCOVERY OF THE SHARK GLYPHIS GANGETICUS 271

paratype paratype of G. gangeticus ，Bengale ， Belanger (see DUMERIL ， 1865: 354); NHW 61379 ， holotype holotype of Carcharhias (Prionodon) siamensis ， immature male ，630 mm tot a1 length ， “Mundung des Rangoon-fluss ぺ1895 or 1896 (found without name or locality data in NHW fish collection in September 1997). Non-type material examined.-Whole specimens: ZSI 8067 ，610 mm ，fem a1 e，ZSI (Calcutta) (Calcutta) F. 8067 ，610 mm ，newbom female ，Hooghly River ，J. Anderson ，4 April 1867 (this (this specimen was first identified as G. gangeticus by COMPAGNO ， 1984: 508); CAS 216841 ，715 mm TL ，Bangladesh ，Bay of Bengal ， ca 1990 ， Department of Fisheries ， Bangladesh Bangladesh (gift to Tyson R. Roberts arranged by A. Ataur Rahman) (although there was no precise collection data ，fisheries staff insisted that this specimen c創 ne from a marine loc a1 ity; 出ey also insisted that no sharks occur in fresh water in Bangladesh); AMS 1. 43504-001 ， 863 mm female ，Myanmar ， Sittway market ，May 2004 ，Tyson R. Roberts; MNHN ，798 mm female ，Myanmar ， Sittway market ，May 2004 ，Tyson R. Roberts; jaws or or teeth ，claspers: CAS 216839 ，Bangladesh ， Cox's Bazar ，tourist shop on the beach ，6

June June 1996 ，Tyson R. Roberts (j aws 230 mm wide; upper tooth rows 15-2 ・ 15 ， lower tooth rows rows 16 ・1・16); CAS 216840 ，teeth only from female ca 2.8 m female ，Bangladesh ， Cox's Bazar ，market ，1 June 1996 ，Tyson R. Roberts and M. N. Sarkar (according to the fishermen this this fish was caught in the open sea not far from shore); unca t.， set of jaws and claspers of of 415 mm wide from 236 cm mature male ， Sittway market ，22 March 2004 ，Tyson R. Roberts; Roberts; unca t.， 8 sets of jaws ， 115-369 mm wide ，Myanmar ， Setyone Su Quarter ，Sittway ， April April 2004 ，Tyson R. Roberts. Original Original descriptions of G. gangeticus and C. siamensis.- Th e joumals in which the origin a1 descriptions of G. gangeticus and C. siamensis occur are seldom found outside the older older institutional libraries in Europe and North America. They apparently are not present in in any institutionallibraries in South Asia or Southeast Asia. Th e descriptions are reproduced as as Appendic 四 1 and 2. Type locality of Glyphis gangeticus.-A literal translation of the purported type locality locality of Glyphis gangeticus would be “ In the Ganges ，60 leagues above the sea by the Hooghly." “Stunden ，" the usual meaning of which is “hours" ，is inte 中reted as “leagues." A league is based upon the distance traveled in an hour ，either by foot or conveyance. The distance distance involved would depend on whether the travel was by land or by water. It varied greatly greatly in different countries. While 3 miles is sometimes given as an average or typical distance distance of a league ， Oxford English Dictionary ， second edition ，1989 ， quotes a mid- seventeenth seventeenth century source (“ Blundevil ， 1595; 1636") as follows: “Th e French league containeth containeth two of our miles ， the Spanish league three ， and the common league of Ge ロnany four ， and the great league of Germany five of our miles." Taking Taking the 10c a1 ity description provided by MULLER & HENLE 1839 as liter a1 1y true ， and using the German equivalents of league of 4 or 5 miles ，would place the type locality of of G. gangeticus 24 0- 300 mi upriver from the mouth of the Hooghly ，and actually in the Ganges mainstream ，properly speaking. Such a locality ，however ，is demonstrably e町oneous. Th e original locality data ， in French ，was incorrectly translated into German ，and the value of of the league it referred to was probably only 2 miles. The evidence for this will now be presented. presented. 272 TYSON R. ROBERTS

August Lamare-Picquot and the Original Collection of “ the Ganges shark"

Th e type locality and other collection data of the holotype and single paratype of G. G. gangeticus has been a source of confusion about 出is species ever since it was reported ， erroneously erroneously on several po 泊ts ，as “im Ganges ，60 Stunden oberhalb des Meers bei Hougly gefangen. gefangen. Ein Exempl 訂 trocken in Zoologischen Museum in Berlin ， durch Lamare Pi cquot ， ein ein Exemplar in Paris (in Weingeist)." Taken literally ， the statement just quoted indicates that both type specimens were collected collected in the Ganges River. Only the dry skin of the large syntype was collected by Lamare-Picquot ，and he did not collect it in the Ganges River ， nor did he record 10c aI ity information information in German. Lamare-Picquot Lamare-Picquot published a hitherto overlooked a∞ ount of the trip on which he evidently evidently collected Glyphis gangeticus (Lamare-Picquot ， 1835). An original copy of this r訂 e document is in the Bibliotheque Centrale of the MNHN. Before citing relevant p訂 ts of of it ，a brief account of the life ，travels ， and collections of Lam 釘 e-Picquot is 泊 order. August Lamare-Picquot (also spelled Lamarre-Picquot) ，was bom at Bayeux ，France ， in in August ， 1785 and died there in 1873. For many years he was a Voyageur-Natur aI iste of of the Musee d'Histoire Naturelle in Paris. He was a pharmacist in Mauritius in 1815. His travels travels took him to l'isle Bourbon ，Madagascar ， and In dia. Upon retuming to Paris 泊 1830 he arranged the sale of his extensive zoologic aI collections ， ethnographic objects ，and antiquities. antiquities. Much of his zoologic aI collections including fish except elasmobranchs apparently were given or sold directly to the Musee d'Histoire Naturelle ， but his ethnographic 佃 d some zoologic aI collections including all of his sharks (along with the eventual holotype of of Glyphis glyphis and syntype of G. gangeticus) were auctioned and wound up mostly in the the Berlin Museum. His sharks specimens ， obtained from many Indian Ocean loc aI ities ， apparently apparently were all reported upon by MULLER & HENLE ， 1839. Two accounts ofhis travels and and collections were published prior to the collections being auctioned ， in Le Moniteur Universel Universel (Paris) for 5 mai 1831 ， pp. 923-924 and for 1 ju 泊 1831 ， p. 1056. An account of his third 町ip to 出e Sundarbans ，on which he aI most certainly collected the the holotype of G. gangeticus and perhaps also that of G. glyphis ，was privately published by Lamare-Picquot in 1835. This description of his trip apparently was added as an afterthought afterthought or postscript to an article in which he defended his [probably erroneous] views on cobra venom against the criticism of Constant Dumeri l. Th e following extracts ，translated from French ， provide a detailed idea of the conditions under which G. gangeticus was first collected. collected. Imperfect though the information is on the precise locality ，it provides a clear indication indication that the holotype of G. gangeticus was collected not in the Ganges or 泊 the Hooghly ， but rather in the lower Sundarbans or estuarine mangrove of the Ganges south of of and probably not far from Kulna. It seems cert 副n that the syntype of G. gangeticus was collected collected on this trip. Most likely it was obtained at the big [unidentified] island where Lamare-Picquot Lamare-Picquot and his men collected and prep 訂 'ed large specimens of rhinos ，tigers ， crocodiles ，and many other things. Th e possibility that it was obtained somewhere else en route route to Kulna or even in Kulna itself cannot be excluded but it was definitely not collected in in the Hooghly or 泊 the Ganges above C aI cutta.

When 1 undertook a third voyage in the southernmost part of the Sundarbands Sunderbunds (“ Sunderbunds ou Sundries") ，my purpose was to increase my zoological collections with some of of the animals populating the vast solitudes of this country … After starting preparations at REDISCOVERY OF THE SHARK GLYPHIS GANGETICUS 273

Chandranagore Chandranagore (“ Chandemagore") ， where 1 resided ， and making fin aI紅 rangements in Calcutta ， left 1 left there on 2 November 1828. Two large boats ， each with five sailors ，carried the provisions with with everything necessary ，toge 血er with nine PO 巾 guese In dian ， and Muslim hunters and prep llI百， tors ， and two domestic s巴rvants. Le aving C aI cutta ，1 headed the for Keedrepoor Can aI (“ can aI de Keedrepoor" ， his italics) ， which serves for communication between 由e eastem branches branches [i. e. ，deltaic dis 佐ibutaries] ofthe Ganges and the Hoog hI y. A 白er five days of navigation 1 had penetrated into 白 e islands 1 wished to explore... Retuming Retuming from a short hunting 凶 P1 noted 白at my hunters ，In dians and Portuguese bom

in in Beng aI， were discouraged. Th e cause was that 白ey we 陀 traveling in places with enormous tigers ，buffalo ， and crocodiles ， and that they also fe 釘 ed thieves who hid in the area after having robbed robbed travelers go 泊g and coming from Dacca to C aI cutta. After After four nights in the wild forests ， by the preparations for hunting 血at 1 made ，the men were were discouraged by the frightful groaning of the hosts of wild anim aI s in 出e fores t. In vain I 甘ied to restore their mor aI e when their courage was needed. For the moment ，1 had to abandon abandon the project of hunting in these quarters ， and to go up one of the main charmels of 曲e Ganges ， towards the north ， to reach Kulna [my it aI ics] ， the g閃 at bazaar located between C aI cutta and Dacca. Th ere 1 hoped to fmd men accustomed to dangerous hunts. After having set set down my conditions and engaging six determined men ，all armed by English rifles and poisoned poisoned arrows ，1 loaded them onto a third boat ， and directed myself ，wi 由 my sm aI l f1 otilla ， towards towards the waterways 1 had left five days before ・.. 1 soon had word of rhinoceros 泊 the vicinity vicinity and made preparations to shoot them ，at an island where tigers were very common ， some si.X ty leagues distant from Calcutta ( “eloignee plus de soixante lieus de Calcutta") … (p. 57). 57). 百 e hunt ， which lasted for 49 days of hunting ，danger ， and fatigue ， and aI l sorts of preparation preparation and conservation of zoologic aI specimens (“ mes animaux") ，resulted in two hornless rhinoceros [regarded [regarded rhinoceros by Lam are-Picquot as a new species]; a Roy aI Bengal tiger; 血ree Ax is (cerf (cerf mouchete); five crocodiles belonging to two species; four tiger-cats of two species; two boar; boar; six monkeys of two species; ten monitors (Tupibnambis ， Varanus vittatus); two species [SIC]; [SIC]; several other species of lizards ，snakes ，turtles ，diverse mollusks; and one hundred and thirty ・three birds...(p. 64).

官lere is no specific mention of fish or sharks but this is the only place in the entire narrative narrative whe 問 a locality is mentioned in terms of leagues or lieu ，and this is 血e one that says “eloignee plus de soixante lieus de Calcutta." Presumably it is 白is information that accompanied Lamare-Picquo t' s specimen of G. gang e. ticus to Berlin ，and that Muller and Henle subsequently misinterpreted. Using the French value for the league of two miles would place Lam 訂 e-Picquot's locality just south of Kulna and only about 50 mi inland from the Bay of Benga l. All of the rivers 叩 d creeks there are strongly tida l. Th e account of Lamare-Picquot's collecting trip ， privately published in Paris ，probably was unknown to them. 百le small syntype of G. gangeticus ， associated perhaps unintentionally with the name of “Lamare Picquot" by Muller and Henle ，was not collected in the same locality as the the holotype. It supposedly was collected some ye 訂 s later by Belanger (DUMERIL ， 1865: 354). 354). Belanger traveled extensively in Benga l. Most of his fish specimens are m 紅 ine. A specimen of G. gangeticus should be in the Senckenberg Museum 泊 Fr 却水fort (RUEPPELL ， 1852: 37; DUMERIL ， 1865: 354) but it is at least tempor 釘 ily misplaced (Friedhelm Kru pp ， pers. commun. ，3 J，佃 2005). It was probably collected by McClelland around 1840. Edward Blyth (1860) reported seeing numerous G. gangeticus 7 feet long in Calcutta fish markets; apparently he did not preserve 組 y specimens. (BLYTH ， 1860). 274 TYSON R. ROBERTS

Glyphis Glyphis gangeticus in Sittway and Rakhine

1 worked 泊 Sittwayand 血e Rakhine District of Myanm 釘 on the Bay of Bengal coast of of Myanm 紅針。m 20 March through 3 May 2004 ， or a total of 45 days. Visits to fish markets markets in Sittway and other localities ， during the peak fish marketing hours (usually 7-9 a.m.) a.m.) were made on 30 days (the other 15 days were devoted to fieldwork 泊仕'eshwater

釘 eas where the collecting methods and other opportunities to cap 卸 re or observe sharks were nil). A total of eight f回shly caught G. gangeticus were observed at three of the five markets markets visited on six of the 23 market days. Th ese observations may be summarized as follows: Market place Sittway Kyaukdaw Mr auk-U Myebon Kantaungyi Days visited 23 1 4 1 1 G. G. gangeticus 5 0 0 2 1 Glyphis Glyphis gangeticus were observed in Sittway market on 22 March ，22 April (2) ，1 May ，and 3 May; 泊 Myebon on 23 April (2); and in Kandaungyi on 26 April. Fi 由 sold in in Sittway may be caught in local marine and estuarine waters but also from Bangladesh and In dian national waters as f，釘 as three or four sailing days distant from Sittway. All fresh fresh fish 1 observed in Sittway market are from marine or strongly estuarine areas. All fishermen fishermen questioned in this reg 訂 d insisted that all sharks marketed in Sittway are caught in in marine habitats and not in fresh water. Sharks reportedly 紅 e rarely caught in 1訂 ge tidal rivers rivers such as the Kaladan 出 f釘 upstream as Kyaukdaw. At 血is point the Kaladan it still very very broad and strongly tidal. According to local people sharks do not occur 加出e Kaladan at at Paletwa (near the limit of tidal influence) or farther ups 回肌Sharks are occasionally seen seen in Mr auk-U market but 1 did not see G. gangeticus or any other sharks on the days visited. 1 visited. Th is important market ， with many visitors and buyers from distant places ，at 住acts fish fish from as f，釘 as Sittway which it is connected to by rapid boat transpo 礼 Myebon and Kantaungyi 紅 'e of more interest because the G. gangeticus seen in these much more isolated isolated and distant markets were 泊 very f詑sh condition and the sellers insisted 血at they had been caught in near ゐy mar 担e waters. Tooth counts of G. gangeticus jaws obtained at Sittway.- Th e following tooth counts counts were obtained from the series of nine dried jaws obtained by the author at Sittway market market and Setyone village in March-April 2004 (greatest width of dried jaws ，numbers of of teeth in upper jaw left side/right side //l ower jaw le 負 side/right side): 1) 1) 118 mm ，15/ 1/ 15 //1 6/1 /16 (donated to BMNH) 2) 2) 227 mm ，16/1 /1 7//1 6/ 1/ 15 3) 3) 242 mm ，15/0/15 //1 5/0 /1 5 4) 4) 325 mm ，14/0/14 //1 3/ 1/ 15 5) 5) 335 mm ，15/0/15 //1 6?/1/16 6) 6) 360 mm ，15/0 /1 4//1 6/1 /1 5 7) 7) 345 mm ，15/1/14 //1 6/1 /1 6 8) 8) 369 mm ，14/0/15 //1 5/1/16 (donated to SAM) 9) 9) 415 mm ，14/ 1/ 14 //1 5/1 /15 (from 236 cm mature male obtained at Sittway market ， 22 March 2004). REDISCOVERY OF THE SHARK GLYPHIS GANG Ef ICUS 275

ACKNOWLEDGMENTS

M 'O nette Veran kindly hand-carried 'O ne specimen 'O f G. gangeticus fr 'O m Bangk 'O k t 'O Paris ，and Sarah F 'O wler tw 'O sets 'O f the jaws ('O ne f'O r BMNH and 'O ne f'O r SAM) fr 'O m Bangk 'O k t 'O L 'O nd 'O n.τ 'h e auth 'O r's recent fieldw 'O rk in Myanmar has been funded by gran 飽 食om the Smiths 'O nian Tr 'O pical Research In stitute and fr 'O m Chester Z 'O'O l'O gical Gardens 佃 d 白eN 'O rth 'O f England Z 'O'O l'O gical S 'O ciety. 官 lis paper was written in the Center f'O r C 'O nservati 'O n Bi 'O l'O gy at Mahid 'O l University ，Salaya campus ，Th ailand ，and 泊血e Raffles Museum f'O r Bi 'O diversity Research at the Nati 'O nal University 'O f Singap 'O re. Permissi 'O n t 'O c'O lllect and expert specimens f'O r scientific study was facilitated by the Myanm 紅 Dep ぽ加.ent 'O f Fisheries ， particularly Deputy Direct 'O rU ffia Win.

REFERENCES

AOASSIZ ， L. 1843. Recherches sur les Poissons Foss i1 es. Neuchatel ， 3. BLYTH ， E. 1860. Th e carωaginous fishes of lower Bengal. よ Asiatic Soc. Bengal 29 (1): 35 -4 5. COMPAONO ， L. J. V. 1984. FAO species ca 凶 ogue. Vo l. 4. Sharks of the world. An annotated and iII us 回 ted ca 凶 ogue of shark species known ω 伽 te. Part 2. Carcharhiniforrnes. FAO Fish. Synop. No. 125 ， v. 4 (pt (pt 2): 251 -6 55. COMPAONO ， L. J. V. 1988. Genus Glyphis Agassiz ， 1843: River Sharks ， pp. 328 ・334 in L. J. V. Compagno Sharks 01 01 the Order Carcharhiniformes. Pri nceton Univ. ，Pri nceton NJ ，U.S.A. ，xxiv +4 86 pp. ， 35 pls. ，figs. COMPAONO ， L. J. V. 1999. Checklist of Ii ving elasmobranchs. In: W. C. H 創凶ett (ed.) Sharks. Skates. and Rays: the the Biology 01 Elasmobranch Fishes. Johns Hopkins Univ. ， Baltimore MD ， U.S.A. & London ， U.K.: I-x ， 1-515. COMPAONO ， L. J. V. ，AND S. F. C∞ K. 1995. Th e exploitation and conservation of freshwater elasmobranchs: sta 伽 s of taxa 加 d prospects for 白巴 future. Pages 62-90 in M. 1. Oetinger 加 d G.D. Zo rzi (eds) ，The Biology Biology 01 Freshwater Elasmobranchs. J. Aquariculture and Aquatic Sciences 7: 1-162. DUM 鼠且， A. H. A. 1865. Histoire naturelle des poissons ou ichthyologie generale. 1. Elasmobranches. Plagiostomes et et Holocephales ou Chim 色res. Hist. Na t. Poiss. Ic hth. Gen. v. 1: 1-720 ，Atlas: pp. 1-8， Pls. 1-14. [P t. 1 = pp. 1-352 ，Pt. 2 = 353-720.] FOWL 殴， S. 1997. River shark discovered in Sabah. Shark News 9 (http://www.flmnh.ufl.ed u/ fis h/ Org 加 izations/ SSG/sharknews/sn 9/ sh 釘 k9newsl 1. htm) FOWLER ，S. ，M. B. MANJAJJ ， R. D. CAVANAOH ， L. J. V. COMPAONO ， S. G. MYCOCK ，AND P. J. LAST. 1999. Elasmobranch Elasmobranch biodiversity ，conservation and management in Sabah (Malaysia). Proc. 5血In do-Pac. Fish Conf. ，Noumea ， 1997. Sco. Francaise Ichth. ，Paris ， 1999: 257-269. GARRICK ，J. A. F. 1982. Sharks of the genus Carcharhinus. NOAA Tech. Rep. NMFS Circ. No. 445: 1-194. LAMARE- Pt CQUOT ， A. 1835. Response pour servir de refutation aux opinions et a la critique de M. Constant Dumeril ， sur mon memoir concemant les Ophidiens... suvie d'une relation de chasse dans les isles des bouches bouches du Gange. Crochar ，Paris ，64 pp.

MULLER ，J. ，AND F. G. J. HENLE. 1839. Systematische Beschreibung der Plagiostomen. Berlin. Plagiostomen: ト xxii xxii + 1-200 ，60 pls. [pp. 1-28 published in 1838 ，reset pp. 27-28 ，2 9- 102 in 1839 ，i-xxii + 103-2 ∞ in in 184 1.] RUEPPELL ， E. 1852. Verzeichnis der in dem Museum der Senckenbergischen Naturforshunden Gesselschaft auf gestellten gestellten Sammlungen. Vierte Abthe i1 ung: Fische und der Skelette. Fr 釘虫 fort am Main. STEINDACHNER ， F. 1896. Bericht uber die wahrend der Reise Sr. M 司j. Schiff ‘Aurora' von Dr. C. Ritter v. Mi eroscewski in den Jahren 1895 und 1896 gesammelten Fische. Ann. K. K. Naturh. Hoj 初ω. 11: 197-230 ， l. pl. 4. 276 TYSON R. ROBERTS

Appendix 1. Th e original description of Glyphis gangeticus (Muller & Henle 1839):

Schnautze Schnautze sehr kurz und sωmpf. Naslocher mitten zwischen Maul und Sc 加 autzenspitz. Maul wenig konvex. Augensp a1 ten sehr klein. Zahne im oberkiefer gleichschenklish ，gerade ， beide R 伽 der etwas nach innen konvex ， der der aussere Rand starkere ， an den hinteren Zahnen einem Absatz bildend ， auf beiden Seiten gleich gleich gezahnelt ， die vorderen wenig langer ，a1 se sie an der Gr enze des Schmelzes breit sind. sind. Die Zahne des unterkiefers gerade ，schm a1， deutlich gezahnelt is t. 1m Ober- und Unterkiefer Unterkiefer ein kle 泊er Mi ttelzahn ，h a1 b so lang a1 s die nachsten ，konisch ，ungez 滋melt ， nicht nicht scbneidend. Zahl der Zahne 31/ 31. Die erste Ruckenflosse f: 釦 gt an ， gerade wo die Basis der Brustflosse aufhort ， ihre Basis Basis ist grosser a1 s die Distanz zwischen der ersten Ruckenflosse und gleicht dem Abstand der der zweiten Rucken- von der Schwanzflosse. Die After flosse ist von derselben Ges ta1 t und Grosse ，wie die Ruckenflosse; sie ist um die doppelte Lange der In sertion der Bauchflossen von den Bauchflossen entfemt; die Di stanz zwischen Bauch- und Afterflosse is nicht ganze zwei M a1 so gross ，a1 s die Di stanz zwischen After- und Schwanzflosse. Die zweite Ruckenflosse Ruckenflosse steht gerade uber der Afterflosse. Die Winkel sammtlicher vertik a1 en FIossen sind sind spitz ， die hintem W泊~el etwas in eine Spitze verlangert ，d記 Rander tief ausgeschnitten. Brus 出ossen sehr lang ，1 す m a1 so lang als breit ， beide Winkel spitz ， der aussere Rand ausgeschnitten. ausgeschnitten. Farbe Farbe oben und unten graubrun ， unten heller. Schuppe !1 dreuekig. Lange ~ Fuss. Korper zum Schwanz wie 6:5. Von der Schnautzenspitz bis zur Mitte der Nasloche 2" Von da zum Maul 2" Vom Maul zum After 29" 6'" VomA 白er zur Afterflosse 9" Basis Basis zur Aftersosse 3" 6'" Hohe derselben 2" 6'" Von der Afterflosse bis zur Schwanzflosse 4" Lange der Schwanzflosse 17" 6'" Von der ersten Ruckenflosse z1江 Bauchflosse 7" Basis Basis der ersten Ruckenflosse 9" Hohe derselben 7" Basis Basis der zweiten Ruckflosse 5" Hohe derselben 3" Lange der Brustflossen 14" 6'" Breite Breite derselben 8" 6'" Distanz Distanz der Naslocher 6" 6'" Breite Breite des Mauls 8" Fundort. Fundort. 1m Ganges ，60 Stunden oberh a1 b des Meers bei Hougly gefangen. Ein Ein Exemplar trocken 泊 zoologischen Museum im Berlin ， durch Lamare Piquot ， ein ein Exemplar in Paris (in Weingeist). REDISCOVERY OF THE SHARK GLYPHIS GANGEJICUS 277

Appeodix 2.

Th e only specimen ever identified as Carcharias (Prionodon) siamensis Steindachner 1896 1896 is 血 e holotype. Given the unusual history of the holotype and 血e confusion surrounding 白 identification of this species the ori 仰 al description (S 担問DAC 剛 ER ， 1896: 229-230) may be reproduced here. My comments or meas ぽ 'ements are given in brackets. Since the original original account w 出 oot accompanied by 血 y figures ，photographs and a drawing of 血e holotype holotype also are provided (Fig. 3). It is unfortunate 白紙 Steindachner chose the name "siamensis" "siamensis" for 白is species. 官 le type locality ， the mouth of the Rangoon River ，is 加 Myanmar ， formerly known as Burma. It was never ap 制 of Thailand (formerly Siam).

Caracharias Caracharias (Prionodon) siamensis n. sp. 1 Ex. ，mas ，63 cm [630 mm] lang. Schnauze Schnauze breit ， stumpf gerunde t. Die Entfemung des Schnauzenspitze von der Mitte des des vorderen Mundrandes ist mehr als 1 1/3 inal in der Breite der Mundspalte zwischen den den Mundwinkelen enthalten. Eine der inneren Narinen-winkel verbindende Horizontallinie liegt liegt fast 2 mal naher zum vorderen Schnauzenende als z町 Mitte des vorderen Mundesrandes. Eine Eine tiefe ， halbmond-formige Grube nachst den Mundwinkeln. Li ppenfalte vorhanded. K.i eferzahne ahnlich wie bei Carch. pleurotaenia Blkr. ， die oberen regelmassig 白羽田kig ， gleichschenkelig ，seitlich deutlich gezahnt ， die unteren sehrshlank ，spitz ，seitlich sehr undeutlich undeutlich gez 剖mt ， auf einer breiten ungezahnen Basis sich erhebend. Kieferzahne 29/29 [1 [1 counted 15/ 1/ 15 upper and 16/1/16 lower jaw teeth for a total upper and lower tooth count count of 31/33; the ou 飽rmost small teeth could be overlooked easily]. Auge sehr kleine. Die Die Hohe der ersten kiemenspalte gleicht c. der Holfte ， die der letzten nur 1/3 einer Schnauzenlange. Schnauzenlange. Pectorale Pectorale ziemlich stark entwickelt ，dreieckig ， mit schwach convexem oberen und starker starker concaveo hinteren Rande. Der untere hintere Winkel der Pectorale ist ein rechter ， der der obere spitz die allgemeine Form der Flosse ist somit gedrungen sichelartig. Oberer Rande der Pectorale 3 mal 凶 1ger als der untere. Die erste Dorsale liegt n油 er zur Basis der der Pectorale als zu der der Ventrale. Die Basislange der ersten Dorsale ist c. 1τmal in der der Entfemung des vorderen Schnauzenendes von der Letzten Kiemenspalte enthalten und ubertri 百t bedeutend die Hohe der Fl osse ， welche c. einer Schnauze n1盈 1ge gleich t. Nach hinten hinten ist die 1. Dorsale wie die 2. Und die Anale in eine Spitze ausgezogen. Der ober Winkel der 1. Dorsale ist ein stumpfer ， an der Spitz gerundet ， der hintere obere Flossenrand sehr sehr schwach concav. Di e 2. Dorsale ist nicht bedeutend starker entwickelt alse die anale und beginnt ein wenig vor dieser. Der hintere Rand der 2. Dorsale ist femer schwacher concav concav alse der der An ale. Der Abstand der 2. Dorsale von der 1. ist. 先 mal grosser als die die Basisl 加 ge der 2. Dorsale. Die Lange der Caudale ist mehr als 3t in der Totallange enthalten. enthalten. Der hintere Rand der Schwanzflosse ist shrage nach unten und vO l1). e abgestutzt ， die die Lange des vorderen Randes des sichelformoigen unteren Lappens c. 2τmal in der Gesammtlange der Fl osse enthalten. Schuppen Schuppen kleine ， mit 3 Kielen an der Aussenflache ，von denen jeder nach hinten in eine eine zahnartige Spitze auslauf t. 278 TYSON R. ROBERTS

Entferτ lU ng der Schnauzenspitze von der hintersten Kiemenspalte 14 .4 [1 4.4] Cm. ， Schnauze 凶ange 4.6 [4.5] Cm. ，Augenl 勧1ge 4.4 [5. 牛 5.6] Mn.， Basislange der 1. Dorsale 7.8 7.8 [7.8] Cm. ，Hohe derselben 5.1 [5 .1} Cm. ，L 加 ge der Caudale 17 Cm [167 .5 mm] ， L 伽 ge der Pectorale 11. 5 [1 1. 3 maximum] Cm. Oben und seitlich bleifarben ，Bauchseite und nachstgelegener 官leil der Rumpfseiten gelblichweiss. gelblichweiss. Caudale nachst den Winkelspitzen etwas dunkler ，shwarzzlichgrau ，doch ohne dunklen Fl ec k. Pectorale und Ventrale schmutzig gelblichweiss.