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Review of the Indo-Pacific Doryrhamphine Pipefish Genus Doryichthys

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Review of the Indo-Pacific Doryrhamphine Pipefish Genus Doryichthys Japanese Journal of Ichthyology 魚 類 学 雑 誌 Vol.28,No.11981 28巻1号1981年 Review of the Indo-Pacific Doryrhamphine Pipefish Genus Doryichthys C.E.Dawson (Received November 6,1980) Abstract The tropical Indo-Pacific trunk-pouch pipefish genus Doryichthys Kaup is reviewed. The genus is,in part,characterized by the presence of a single longitudinal ridge on the opercle,by typically failing to have the lateral trunk ridge confluent with the inferior tail ridge and by having a modal count of 9 caudal-fin rays.Included species(D.deokhatoides,D.martensii,D.boaja and D.heterosoma)are presently known from freshwater habitats at insular and mainland localities from Indonesia to Vietnam.A key,synonymies,diagnoses and illustrations are provided for all species . Kaup(1856)included two morphological confluent with the inferior tail ridge(e.g. lineages in his diagnosis of the doryrhamphine Microphis,Oostethus,etc.).Although available (trunk-pouch)pipefish genus Doryichthys,and museum specimens have been examined,study failed to clearly differentiate this taxon from the material is limited for some species.Never- genus Microphis Kaup.Duncker(1910,1912, theless,the present treatment should provide a 1915)discussed these genera but failed to examine basis for identification of included species as well critical type material,perpetuated some of Kaup's as for future studies on variation and distribu- original errors,and added further confusion by tion. introducing major nomenclatural problems. Some species have entered the international Duncker initially(1910)referred these genera to aquarium trade(Sterba,1959;Wheeler,1975) his nominal subfamily Doryichthyina(=Dory- and introduction of species of Doryichthys to ichthyinae),but later(1915)changed this name extralimital freshwater habitats is to be expected. to Doryrhamphina(=Doryrhamphinae).As a result,subsequent authors have,with little con- Methods and materials sistency,referred some 15 or more nominal Methods are those of Dawson(1977).Meas- species of predominantly freshwater or estuarine urements are in millimeters(mm);proportional Indo-Pacific pipefishes to one or the other of data are referred to standard length(SL)or head t hese genera.In addition,species diagnoses length(HL);some measurements are given for are largely inadequate,some species have never total length(TL),whereas those given for damag- been illustrated,and available literature generally ed specimens in Materials Examined refer to fails to allow identification of individual speci- present overall length;color descriptions are mens.In continuation of studies on Indo- from specimens preserved in alcohol.As used Pacific pipefishes,I here review the genus Dory- here,the term "venter" refers to the ventral ichythys Kaup and include therein four species surface of head or body.Distribution is based which typically fail to have the lateral trunk ridge largely on specimens examined. confluent with the inferior tail ridge.This Abbreviations for repositories of material treatment differs from those of Kaup(1856) examined follow: AMS•\Australian Museum, and Duncker(1915)in that one,rather than two, Sydney;ANSP•\Academy of Natural Sciences lateral ridge configurations are included in the of Philadelphia;BMNH—British Museum generic diagnosis.Although there is some in- (Natural History),London;CAS-SU•\former dividual variation,the typical ridge configura- Stanford University specimens,housed at Cali- tion is here considered a primary character dis- fornia Academy of Sciences,San Francisco; tinguishing Doryichthys from nominal genera GCRL•\Gulf Coast Research Laboratory characterized by having the lateral trunk ridge Museum;MCZ•\Museum of Comparative ―1― 魚 類 学 雑 誌Japan.J.Ichthyol.28(1),1981 Zoology,Harvard University;MNHN•\Mu- plemental ridges;pectoral-fin base not protruding seum National d'Histoire Naturelle,Paris; strongly laterad,the superior and inferior ridges MZB•\Museum Zoologicum Bogoriense,Bogor; usually distinct;scutella without longitudinal NMW•\NaturhistorischesMuseum Fischsam- keels;principal body ridges low to somewhat mlung,Wien;RMNH•\Rijksmuseum van elevated,notched or indented between rings,the Natuurlijke Historie,Leiden;SMF•\Natur- margins entire,denticulate or serrate under x 30 Museum und Forschungs-Institut Senckenberg, magnification;rings with or without a prominent Frankfurt am Main;UMMZ•\niversity of spine on posterior third of each principal ridge; Michigan Museum of Zoology;USNM•\ head and body without dermal flaps or papil- National Museum of Natural History,Smith- lae;dorsal fin originates on trunk,its base not sonian Institution;WAM—Western Australian elevated;pectoral fins emarginate(unknown in D. Museum,Perth;ZMA•\ZoOlogisch Museum, heterosoma),the median rays shorter than those Amsterdam;ZMB•\Zoologisches Museum, above and below.Head length 4.6•`9.2 in Museum far Naturkunde der Humboldt Univer- SL;snout length 1.4•`2.4 in HL;length of dorsal- sitat,Berlin;ZSI•\Zoological Survey of India, fin base 1.0•`2.5 in HL;trunk rings 15•`26; Calcutta. total rings 45•`62;total subdorsal rings 5.0•` 12.5,mostly on tail ;dorsal-fin rays 27•`69; Doryichthys Kaup 1856 pectoral-fin rays 16•`27;anal-fin rays usually Doryichthys Kaup,1853:233(nomen nudum). 4;caudal-fin rays typically 9(unknown in D. Doryichthys Kaup,1856:56(type-species Dory- heterosoma).Brood pouch originates on 1st•` ichthys bilineatus Kaup 1856(=Syngnathus 3rd trunk ring;pouch plates well developed,with deokhatoides Bleeker 1853),by subsequent or without a narrow marginal membranous fold; designation of Jordan and Evermann 1896). pouch plates vertical to somewhat convergent in Doryrhamphinarum Kaup,1856 62(nomen brooding fish,fail to completely enclose develop- nudum). ing eggs and do not meet on ventral midline;eggs Kaupia Smith,1963:533(type-species:Syn- deposited in 1 5 transverse rows within mem- gnathus boaja Bleeker 1851,by original branous compartments lining dorsum of pouch designation). (Fig.4),evidently in single layer(unknown in D. Diagnosis.Superior trunk and tail ridges boaja and D.heterosoma).Without odontoid discontinous near vertical through rear of dorsal- processes in jaws(Dawson and Fritzsche,1975) fin base,not arched dorsad below dorsal-fin base; or bony inclusions in gill membranes(Dawson, lateral tail ridge ends near anal ring;lateral 1978).Maximum size at least 470 mm TL. trunk ridge ends with or without deflection Indonesia,Malay Peninsula,Cambodia,Thai- near anal ring,typically fails to unite with con- land and(questionably)Taiwan;presumably fluent inferior trunk and tail ridges(Figs.2,4); freshwater,not definitely recorded from marine venter of trunk more or less V-shaped in adult or estuarine habitats. females ;median ventral trunk ridge begins on Comparisons.Among trunk-pouch pipefishes, pectoral ring,typically distinct,sometimes keel- the Doryichthys configuration of principal body like in large females;median dorsal snout ridges is shared only with the endemic Austral- ridge entire,mainly low,usually ends near ver- ian monotypic,marine,genus Leptoichthys tical through rear of nares but may continue on Kaup.This genus lacks the opercular ridge and anterior part of interorbital;lateral snout ridge pouch protective plates present in Doryichthys not arched strongly dorsad;anterior supraorbital and typically has 5 anal-fin rays and 11 caudal- ridges end at or before vertical through middle fin rays(respectively,4 and 9 in Doryichthys). of orbit,infrequently confluent with posterior Species of several nominal doryrhamphine genera supraorbital ridges;median dorsal head ridges (e.g.Coelonotus Peters,Microphis Kaup,0o- low to somewhat elevated;typically with 1 3 stethus Hubbs,Paramicrophis Klausewitz)may supraopercular ridges and a ridge above gill also occupy Indo-Pacific freshwater or estuarine opening;opercle with a complete or incomplete habitats and may be superficially similar to those median longitudinal ridge,otherwise ornamented of Doryichthys.Unlike Doryichthys,these with minute striae but without prominent sup- genera are,in part,characterized by typically ―2― Dawson:Indo-Pacific Doryichthys A B Fig.1.Lateral and cross-sectional aspects of trunk and anterior tail rings,illustrating location of enlarged spines on principal ridges in Doryichthys boaja(A)and D.heterosoma(B) . having the lateral trunk ridge confluent with the trunk ridge confluent with inferior tail ridge) inferior tail ridge.The Doryichthys ridge con- included D.pristipeltis(a nomen dubium,pro- figuration is shared with a number of syngnath- bably conspecific with Syngnathus brachyurus ine(tail-pouch)genera(Syngnathus Linnaeus, Bleeker 1853)and six nominal species subse- Corythoichthys Kaup,Bhanotia Hora,etc.) quently shown(Dawson,1979)to be conspecific but these have modally 10 rather than the 9 with Syngnathus brachyurus.Kaup diagnosed caudal-fin rays typical of Doryichthys. Doryichthys inadequately and it was synony- Remarks.The name Doryichthys was intro- mized with Microphis Kaup 1853 by Dumeril duced by Kaup(1853)but the ten specific names (1870).In contrast,Gunther(1870)synony- then listed for the genus were nomina nuda. mized Microphis and three other doryrhamphine Although the type-species was not indicated,the genera(Doryrhamphus Kaup,Choeroichthys generic name was validated by Kaup(1856)with Kaup,Belonichthys Peters)with Doryichthys. the description of nine species in two groups Duncker(1910)incorrectly employed Dory- ("A"and"B")with different principal body ichthys as the name for a new genus and later ridge configurations.Within the A-group (1912)designated
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