A Redescription of the Hemipenis of Hydromorphus concolor () with Comments on Its Tribal Allocation Author(s): Brian I. Crother Source: Copeia, Vol. 1989, No. 1 (Feb. 27, 1989), pp. 227-229 Published by: American Society of Ichthyologists and Herpetologists Stable URL: http://www.jstor.org/stable/1445634 Accessed: 16/09/2008 17:02

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http://www.jstor.org HERPETOLOGICAL NOTES 227

TAYLOR, E. H. 1920. Philippine turtles. Philippine everted specimen (CRE 3014) was used for the J. Sci. 16:130-133. following description (Fig. 1). The organ is 11 . 1944. Present location of certain herpeto- subcaudals long, moderately bilobed and fully and other Univ. KansasSci. logical type specimens. capitate; capitulum calyculate distally and spi- Bull. 30, Pt. 1, no. 11:160. nose proximally; reduced papillate micro-or- WERMUTH, H., AND R. MERTENS.1961. Schildkroe- namentation on four basal hooks; ten, Krokodile, Brueckenechsen. Gustav Fischer calyces; large cover the central of the Verlag, Jena, German Democratic Republic. accessory spines portion organ between the capitulum and the basal sulcus bifurcates within the JAMESR. BUSKIRK,La Clinica de la Raza, 1515 hooks; capitulum and is centrolineal. Fruitvale Ave., Oakland, California 94601. Ac- (unicapitate) The asulcate side has a row of cepted 30 March 1988. single large spines just below the capitular groove which increase in size towards the lateral sides of the organ. A central groove is present from the base of the capitulum to the mid-portion of the stalk. Copeia, 1989(1), pp. 227-229 This groove is lined on both sides by three spines ? 1989 by the American Society of which decrease in size towards the middle of Ichthyologists and Herpetologists the stalk. The basal region is nude except for A REDESCRIPTION OF THE HEMIPENIS scattered spinules and two of the basal hooks. OF HYDROMORPHUS CONCOLOR (COLU- Minor variation occurs in the numbers and BRIDAE) WITH COMMENTS ON ITS the arrangements of spines on the stalk and in TRIBAL ALLOCATION.-Hemipenes have the sulcus bifurcation point. In all organs the long been recognized as providing useful infor- sulcus bifurcates well within the capitulum (Fig. mation for systematists and the impor- I) except in a single individual (CRE 9000). In tance of hemipenes as systematic characters has this specimen the sulcus bifurcates just inside been re-emphasized repeatedly (Cope, 1893, the base of the capitulum. 1895; Dunn, 1928; Dowling and Savage, 1960; Dunn (1939) described the hemipenis of Dowling, 1967; Jenner and Dowling, 1985; mainland Tretanorhinusand suggested that Tre- Branch, 1986), even if with caution (Inger and tanorhinus may be closely related to Hydromor- Marx, 1962). phus based on the marked similarity in dentition Jenner (1981) attempted to categorize the and hemipenial morphology. Dunn's descrip- Xenodontinae (sensu Dowling and Duellman, tion is as follows: ". .. hemipenis single or slight- 1978) at the tribal level based on geographic ly bilobed; calyculate area capitate; sulcus forks distribution and hemipenial morphology, but within the calyculate area; four basal hooks; be- she was unable to place Hydromorphus(with three tween hooks and calyculate area about four cross nominal ) because no adequate descrip- rows (about 10 longitudinal rows) of rather uni- tion of its hemipenes was available. The only form smaller spines." This description is re- published description of Hydromorphusconcolor markably similar to the Hydromorphus hemi- (Villa, 1970), a retracted organ, is incomplete. penis, but I examined T. variabilis (a West Indian In the described specimen, Villa (loc. cit.) sug- species) and found it to be distinctly different: gested that the sulcus bifurcated near the base a single lobe; fully capitate; entire captitulum of the penis, at the level of the first or second calyculate with well developed spinulate micro- subcaudal. However, on a partially everted or- ornamentation and a large groove forming a gan from another specimen examined later, he naked pocket on the asulcate side of the capit- noted that the sulcus seemed to remain single ulum (see p. 112a.2 in Dowling and Duellman, into the capitulum. I quote his assessment of 1978 for an illustration). These differences do the sulcus condition: "Whether or not it bifur- not suggest a close relationship (i.e., sister group cates beyond this point I could not ascertain." status) between Tretanorhinus and Hydromor- Here I present a redescription of the hemi- phus, but neither do these differences discount penis of H. concolor following the terminology the possibility of a close relationship. However, of Dowling and Savage (1960), as modified by Dunn (1939) also recognized similarities among Myers (1974) and Myers and Campbell (1981). Rhadinaea, Coniophanes, Trimetopon, Ninia, Additionally, I comment on the uncertain tribal Amastridiumand Tretanorhinus, all of which (ex- affinities of Hydromorphus. cept Amastridium) are placed in the Dipsadini Eight individuals of H. concolor with everted (sensu Jenner, 1981, 1983). hemipenes were examined and the most fully Jenner defined the tribe Dipsadini by the pos- 228 COPEIA, 1989, NO. 1

session of a fully capitate hemipenis with prox- imal spines and a forked sulcus, and a geograph- ic distribution centered in Central America. If we follow these explicitly phenetic criteria, it seems clear that Hydromorphusis part of the dip- sadine radiation because it possesses both the hemipenial characters (Fig. 1) and the distri- butional character (Fig. 2). However, Cadle (1984) found little unity in Jenner's Dipsadini !3 4 <-lH POINT OF CAPITATION (i.e., not monophyletic) and could not place Tretanorhinus in any lineage (Hydromorphuswas not included in his study). My own preliminary cladistic analysis of xenodontines based exclu- sively on hemipenis characters suggests that the Dipsadini (sensu Jenner, 1981, 1983) is not monophyletic with respect to the other Neo- tropical xenodontine tribes. The uncertainty of Jenner's Dipsadini as monphyletic, and the in- conclusiveness of Cadle's (1984) work regard- ing Hydromorphus,makes the systematic place- ment of Hydromorphussubjective at this time.

Material examined.-Hydromorphus concolor:Cos- ta Rica: S bank Jorco, S base of Fila Cedral, Ignacio de Acosta, Depto. SanJose (CRE 3014); 1 cm Caribbean coast NW of Puerto Limon, Depto. Limon (CRE 9000); University of Costa Rica Fig. 1. Sulcate site of left hemipenis of Hydro- San Pedro, SanJose (CRE morphusconcolor (CRE 3014, SVL = 430 mm). Scale: campus, Depto. 1542); 10 cm = 1 cm. 6 km NE Rio Naranjo, Bijagua, Depto. Puntar- enas (CRE 3679); 4 km ESE Monteverde, Finca

Fig. 2. Geographic distribution of Hydromorphus.Dots represent specimens examined in this study and squaresrepresent other known collection localities. HERPETOLOGICAL NOTES 229

Penas Blancas, Depto. Alajuela (CRE 4611); N American xenodontine : the tribe Pseudo- bank Rio Grande de Terraba, 1.5 km SW Que- boini. Herpetologica 41(7):161-172. brada Drori or Puntarenas MYERS,C. W. 1974. The systematics of Rhadinaea Disciplina, Depto. a of New World snakes. Bull. (UCR 7440); Barrio Monte Rey, San Pedro, (Colubridae), Amer. Mus. Nat. Hist. 153(1):1-262. Depto. San Jose (UCR 6330); SW San Jose at , ANDJ. A. CAMPBELL.1981. A new genus and base of N of Fila Cedral, Escazu, slope Depto. species of colubrid snake from the Sierra Madre San Jose (UCR 9229). Tretanorhinus variabilis: del Sur of Guerrero, Mexico. Amer. Mus. Novit. Grand Cayman: 1 mile NE Georgetown (ASFS 2708:1-20. 16231). VILLA, J. 1970. The snake Hydromorphusin Nicara- gua, with a description of its hemipenis. Carib. J. Acknowledgments.-I would like to thank J. M. Sci. 10(3-4):119-121. Savage for sparking my interest in hemipenial BRIAN I. Texas Memorial 2400 morphology and commenting on this note. A. CROTHER, Museum, Texas 78705. 22 Schwartz kindly loaned me material and thanks Trinity St., Austin, Accepted 1988. also go to M. A. Donnelly, R. deSa, C. Myers April and J. B. Slowinski for their comments. M. P. Hayes, D. Krempels, and especially R. deSa are thanked for their expert advice on how to draw. Copeia, 1989(1), pp. 229-231 ? 1989 by the American Society of LITERATURE CITED Ichthyologists and Herpetologists IDENTITY OF A SUPPOSED SOUTH BRANCH, W. R. 1986. of Hemipenial morphology AMERICAN MICROHYLID CTE- African snakes: A taxonomic review. J. Herpetol. FROG, 20(3):285-299. NOPHRYNE MARMORATA.-Ahl (1935) de- CADLE, J. E. 1984. Molecular systematics of Neo- scribed Ctenophrynemarmorata from two speci- tropicalxenodontine snakes:I. South Americanxe- mens in the ZMB for which there are no data nodontines. Herpetologica 4(1):8-20. as to locality or collector. Ahl's reasons for as- E. Prodromusof a new of the COPE, D. 1893. system signing the species to the hitherto monotypic non-venomoussnakes. Amer. Natur. 1893:477-483. genus Ctenophryneare not stated, but the ab- 1895. The classification of the Ophidia. sence of clavicles and the con- Trans. American Phil. Soc. procoracoids, (N.S.) 28(art. 3):186- cealed and vertical in the new 219. tympana pupils are features shared with C. Moc- DOWLING,H.G. 1967. Hemipenes and other char- species, geayi and have influenced his de- acters in colubrid classification. Herpetologica 23(2): quard 1904, may 138-142. cision. Reasoning from the supposed congener- , ANDW. E. DUELLMAN.1978. Systematic her- ity, Ahl assumed that the locality for his new petology: a synopsis of families and higher cate- species differed little from that for C. geayi, gories. HISS Publications, New York, New York. known at the time only from its type locality in , ANDJ. M. SAVAGE. 1960. A guide to the snake Colombia. a of basic structure and hemipenis: survey system- The date of publication of C. marmorata was atic characters. Zoologica 45(2): 17-28. unfortunate in that it followed the E. R. 1928. A tentative and closely upon DUNN, key arrangement of Parker's of of the American of Colubridae. Bull. An- appearance (1934) monograph genera the the new did not tivenin Inst. America 2(1):18-24. Microhylidae. Thus, species receive the critical review it had it come . 1939. Mainland forms of the snake genus might Tretanorhinus.Copeia 1939(4):212-217. to Parker's attention in time, and Ahl evidently INGER, R. F., AND H. MARX 1962. Variation of hemi- did not have Parker's work at hand when pre- penis and cloaca in the colubrid snake Calamaria paring his description. Indeed, the species vir- lumbricoidea.Syst. Zool. (11)1:32-38. tually vanished from the literature as soon as V. 1981. A and the JENNER, J. zoogeographic study described. I find only two mentions subsequent of the xenodontine colubrid snakes. Un- to the Dunn called Ph.D. New York New York original description: (1949) publ. thesis, University, it "a dubious form" and that it New York. very suggested City, well be a form of or of 1983. Allocation of (Tab. X). In: Bio- "might Glyphoglossus genera of southeast Asia." Gorham in- chemical evaluation of colubrid snake taxonomy. Uperodon (1974) H. G. Dowling, R. Highton, G. C. Maha and L. R. cluded it, with question, in the synonymy of C. Maxon (eds.).J. Zool. (London) 201:309-329. geayi. , AND H. G. DOWLING.1985. Taxonomy of Apparently until now no herpetologist has