Tepalcatia, a New Genus of Hydrobiid Snails (Prosobranchia: Rissooidea) from the Rio Balsas Basin, Central Mexico
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3 April 2002 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 115(1): 189-204. 2002. Tepalcatia, a new genus of hydrobiid snails (Prosobranchia: Rissooidea) from the Rio Balsas basin, central Mexico Fred G. Thompson and Robert Hershler (FGT) Division of Malacology, Florida Museum of Natural History, University of Florida, Gainesville, Florida 32611-7800, U.S.A.; (RH) Department of Systematic Biology, National Museum of Natural History, Smithsonian Institution, Washington, DC. 20560-0118, U.S.A. Abstract.—Tepalcatia, a new genus of cochliopine snail from the Rio Balsas basin of central Mexico, is composed of Tepalcatia tela, new species, described herein; and Tepalcatia polia (Thompson & Hershler, 1991), previously assigned to Aroapyrgus. A third species, Tepalcatia bakeri (Pilsbry, 1891a), is tenta- tively referred to the genus. This snail, known only as a subfossil, most recently was assigned to Tryonia. Tepalcatia is placed in the informal "Cochliopa group" based on the absence of penial glands; division of penis into a folded base and smooth, tapered filament; and the very short female sperm duct. Te- palcatia is distinguished from other members of this group by its ovate-conic to turriform shell; long ducts of the female sperm pouches; and large circular- horizontal loop of the coiled oviduct. Tepalcatia tela differs from T. polia by its smaller size, more elongate shell, longer and broader penial filament, and several additional anatomical features. Tepalcatia bakeri is a large, slender- shelled species with stronger spiral sculpture than its congeners. The freshwater snail fauna of the Amer- whose drainage basin is much larger than ican tropics remains poorly known that of the Rio Panuco and is comparable (Thompson & Hanley 1982). The only fam- to that of the Rio Usumacinta (Tomayo ilies that have been studied within the con- 1964, table 1), is by contrast almost com- text of detailed anatomical investigations pletely unexplored. Pilsbry (1891b) record- are the biomedically important Planorbidae, ed a planorbid, Gyraulus parvus, from near and the Hydrobiidae. Those Planorbidae Yautepec (Morelos) and also described a that are not biomedically important have hydrobiid from this general area (Pilsbry been little treated in contemporary litera- 1891a), while we described a second hy- ture. Although the Hydrobiidae may be the drobiid species, also from the eastern por- best known family of freshwater snails, its tion of the Balsas basin (Thompson & Her- tropical American fauna has been little col- shler 1991). All three snails are known lected, and cursory field investigations rou- from single localities. Herein, we describe tinely yield new species. One such species another new hydrobiid from the Rio Balsas is the basis of this report. basin and propose a new genus for this nov- The only drainages in Mexico and Cen- elty, together with the two earlier recorded tral America that have been surveyed in hydrobiids from this drainage. some detail for aquatic gastropods are those of the Rio Panuco (Pilsbry 1909, 1910; Material and Methods Pilsbry & Hinkley 1910, Pilsbry 1956) and the Rio de la Pasidn (Goodrich & Schalie Specimens used for anatomical study 1937), a tributary of the Rio Usumacinta. were anaesthetized with menthol crystals The Rio Balsas of central-Pacific Mexico, overnight and subsequently fixed in Bouin's 190 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON solution and preserved in 70% ethanol. bryonic shell of 1.6-1.75 whorls, with two Shell whorl counts and measurements were distinct sections; initial portion (ca. 0.75 made using the methods of Hershler (1989). whorl) irregularly roughened and/or lined Counts of cusps on radular teeth were made with a few spiral elements, sculpture dimin- by inspection of mounted material using a ishing to near smooth on later portion. Te- scanning electron microscope; other com- leoconch of 2.5-4.25 convex whorls sculp- ponents of radular description were gener- tured with finely spaced growth lines and ated by study of scanning electron micro- occasional, narrowly spaced spiral striae; a graphs. Morphological terminology follows few spiral ridges and weak collabral some- Hershler & Ponder (1998). Specimens of times present in one species. Aperture the new species were deposited in the Flor- ovate, small, inner lip thin, adnate to and ida Museum of Natural History, Gainesville usually complete across parietal wall; outer (UF); and the National Museum of Natural lip thin, straight or weakly sinuate. Umbi- History, Smithsonian Institution, Washing- licus absent or rimate to well developed. ton, DC. (USNM). Other material exam- Operculum thin, transparent, amber col- ined was from the above collections and ored, multispiral, with eccentric nucleus. that of the Academy of Natural Sciences of Outer side of operculum smooth or frilled Philadelphia (ANSP). along edge of last whorl; attachment scar margin smooth. Family Hydrobiidae Radula small, with about 48 well-formed Subfamily Cochliopinae rows of teeth. Dorsal edge of central teeth indented; basal process angled, rounded be- Tepalcatia, new genus low, even with lateral margins; median Type species.—Tepalcatia tela, new spe- cusps longer than laterals, dagger-like; large cies. Also included are Aroapyrgus polius basal cusp sometimes flanked by 1—2 small- Hershler & Thompson, 1991; and Potamo- er cusps. Lateral teeth with rounded or pyrgus bakeri Pilsbry, 1891a. weakly pointed central cusp, weakly or Etymology.—From the Nahuatl (Aztec) moderately developed basal tongue, and tepalcate, meaning shard, which is used as long, straight, flexed outer wing of medium a linguistic stem for various regional fea- width; cutting edge weakly indented dor- tures associated with the southern edge of sally. Cusps on outer marginal teeth larger the Mesa Central (e.g., Rio Tepalcatepec). than those on inner marginals. The genus name refers to the distribution of Head-foot pigment well-developed, gray the three included species along this escarp- or black. Snout darker than cephalic tenta- ment. Gender feminine. cles, although distal lips pale. Snout weakly Diagnosis.—A central Mexican coch- tapered, distal lips expanded. Cephalic ten- liopine genus in the informal "Cochliopa tacles narrow, a little longer than snout, par- group" (of Hershler & Thompson 1992) allel-sided, with rounded tips. Eyes small, which differs from other group members in black, at outer bases of tentacles. Bases of having a more slender (versus planispiral to tentacles without pronounced eyelobe. low conical) shell; elongate female sperm Snout with scattered ciliary tufts. Cephalic pouch ducts; and a large, circular or pos- tentacles with well developed longitudinal teriorly oriented (versus a small, narrowly ciliary tracts on dorsal and ventral surfaces, vertical) loop of the coiled oviduct. and along inner edges. Mantle weakly pig- Description.—Shell ovate- to elongate- mented; pigment concentrated along edges conic, with convex spire, minute to small of genital ducts and ctenidium. Pigment (maximum length 1.6-4.0 mm). Shell clear well developed on testis and stomach; di- and transparent. Periostracum thin, tan or gestive gland pigmented with scattered gray. Apex blunt, often slightly tilted. Em- black granules. VOLUME 115, NUMBER 1 191 Ctenidium well developed, wide, extends a terminal papilla. Sperm pouches small, entire length of pallial cavity. Ctenidial fil- with pink sheen, horizontal, positioned well aments 25-27, triangular, lateral surfaces behind pallial wall and extending to near ridged. Osphradium located near middle of posterior edge of albumen gland; ducts ctenidium, short, narrow. Rectum nearly broadly overlapping, opening from anterior straight, packed with oval fecal pellets. Re- edge of pouches. Bursa copulatrix narrow nal organ extends ca. 33% of length of renal or oval, duct narrow or broad. Seminal re- gland into pallial roof, renal gland oriented ceptacle oval or sub-globular, duct narrow. longitudinally, opening of gland with sim- Coiled oviduct and seminal receptacle join ple lips. Pericardium with a little less than a little behind posterior pallial wall at point 50% of length in pallial roof, ctenidium where former opens to posterior edge of al- overlapping anterior end of pericardium. bumen gland. Sperm duct opening to distal Stomach with similar-sized anterior and section of bursal duct, very short. Sperma- posterior chambers, single digestive gland thecal duct narrow, opening a little in front opening, very small posterior caecum pre- of posterior wall of pallial cavity. sent or absent; style sac about as long as Distribution.—Tepalcatia is distributed remainder of stomach. in widely separated headwater regions of Testis of numerous simple vertically the Rio Balsas and its western tributary, the stalked lobes occupying most of visceral Rio Tepalcatepec (Fig. 1). coil behind stomach and overlapping pos- Remarks.—The anatomical groundplan terior chamber of stomach. Seminal vesicle of Tepalcatia well conforms to that of the a tightly coiled mass of numerous narrow "Cochliopa group" (sensu Hershler & coils, positioned beneath about 0.25 whorl Thompson 1992), an informally recognized of testis behind stomach. Prostate gland al- assemblage of five Neotropical cochliopine most entirely visceral, sub-globular or oval, genera (Aroapyrgus, Cochliopa, Cochliop- thin-walled. Visceral vas deferens opens ina, Mexithauma, Subcochliopa) which ventrally in front of posterior edge of pros- share the following characters: absence of tate gland; pallial