3 April 2002 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 115(1):171-188. 2002. A new and species of aquatic gastropods (: ) from the North American Southwest: phylogenetic relationships and biogeography

Robert Hershler, Hsiu-Ping Liu, and Craig A. Stockwell (RH) Department of Systematic Biology, National Museum of Natural History, Smithsonian Institution, Washington, DC. 20560-0118, U.S.A, e-mail: [email protected]; (HPL) Department of Biological Sciences, University of Denver, Denver Colorado 80208, U.S.A; (CAS) Department of Zoology, Stevens Hall, North Dakota State University, Fargo, North Dakota 58105, U.S.A.

Abstract.—Juturnia, a new genus composed of three species from the Rio Grande region of southwestern North America, is described. Two of these spe- cies were previously placed in the genus while the third, J. tularosae, is described herein. Juturnia is shown to be a monophyletic sub- unit of the gastropod subfamily Cochliopinae based on phylogenetic analysis of mitochondrial DNA sequences and an uniquely shared morphological char- acter (within the subfamily), the presence of a single non-glandular lobe on the inner edge of the penis. Our phylogenetic hypothesis indicates that Juturnia is most closely related to North American brackish-coastal taxa, and we con- jecture that progenitors of this genus penetrated inland along the Tertiary drain- ages which headed along eastern flanks of Laramide uplifts and flowed to the Gulf of Mexico. Subsequent vicariance of congeners in southeastern New Mex- ico is attributed to development of the Pecos depression, and inception of the drainage divide between the Rio Grande Rift and southern Great Plains.

While various hypotheses have been pro- liopinids whose distribution, habitat, and posed to explain the distributions of aquatic systematic relationships provide an addi- biota in the Rio Grande region of south- tional perspective on the biogeographic his- western North America, these have been tory of this region, hampered by a paucity of pertinent phylog- enies (Smith & Miller 1986). Cochliopinid Materials and Methods (= littoridinid) gastropods form a diverse component of this biota and also are of in- This report primarily is based on material terest because of their high level of local reposited in the National Museum of Nat- endemism (Taylor 1966, 1987). Recent ural History, Smithsonian Institution phylogenetic reconstructions based on anal- (USNM). Relevant type material from the ysis of mitochondrial DNA sequences sug- Natural History Museum, London (BMNH) gest that this fauna includes several lineages also was studied. Specimens of the new derived from marine or brackish water species were relaxed in the field with men- coastal progenitors (Liu et al. 2001), and thol crystals, and then fixed in dilute for- mat inland snail distributions reflect vicar- malin (10% of stock solution) and pre- iant events which preceded establishment of served in 70% ethanol. Prior to dissection, modern regional topography (Hershler et al. shells were removed from by soak- 1999, 2002). Herein, we describe a newly ing in dilute hydrochloric acid. Animals recognized monophyletic group of coch- were dissected using a WILD M8 research 172 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

microscope and drawings were made with Systematics the aid of a camera lucida attachment. Live Family Hydrobiidae Troschel, 1857 specimens were studied for description of Subfamily Cochliopinae Tryon, 1866 head-foot and penis. Methods of prepara- Juturnia, new genus tion of material for scanning electron mi- croscopy are of Hershler (1998). Counts of Type species.—Durangonella coahuilae radular cusps and tooth rows were deter- Taylor, 1966, by original designation. Other mined from scanning electron images. Ter- species included are Tryonia kosteri Taylor, minology of anatomical features follows 1987, and Juturnia tularosae, new species Hershler (2001) and character state delin- (described below). eations are from Hershler & Ponder (1998). Diagnosis.—Shell ovate-conic to turri- form, small to medium-sized (2.0-5.1 mm Institutional abbreviations follow Leviton et shell length), grey-white or clear, transpar- al. (1985). ent. Periostracum thin, light tan to brown. Partial sequences of the mitochondrial Protoconch of 0.7-1.0 whorl, planispiral or cytochrome c oxidase subunit I (mtCOI) depressed trochiform, surface corrugated gene were obtained from two specimens of but without microsculpture. Teleoconch of the new species using methods of Liu et al. 4.0—8.5 slightly to strongly convex whorls. (2001) (Genbank accession number Teleoconch sculpture of closely spaced or- AF474371). Sequences were utilized for thocline or weakly prosocline growth lines, phylogenetic analysis along with previously fine spiral striae sometimes present and analyzed mtCOI sequences of members of crossed by growth lines. Aperture ovate, in- cochliopine sub-clades III, IV and V (Liu ner lip thin to slightly thickened, broadly et al. 2001; also see Hershler et al. 1999), adnate to completely separated from parie- which represent the portion of the subfam- tal wall, outer lip thin or slightly thickened. ily topology that contained previously de- Umbilicus rimate to broadly open. Oper- scribed species which we allocate herein to culum thin, multispiral, with eccentric nu- Juturnia. In this earlier analysis, sub-clade cleus. V was sister to the lineage containing spe- Central radular teeth with weak to mod- cies of Juturnia and thus we used its most erately indented dorsal edge; 3-6 lateral basal member (Mexipyrgus carranzae) for cusps on each side; 1 pair of basal cusps, rooting in the current study. Analyses were basal process V-shaped. Lateral radular conducted using the maximum parsimony, teeth with straight to slightly indented dor- maximum likelihood, and minimal evolu- sal edge; lateral shaft much longer than dor- tion options of PAUP*4.0 (Swofford 2000) sal edge; 3—5 cusps on inner side; 4—7 following Liu et al. (2001). Bootstrapping cusps on outer side. Inner marginal teeth with 23-33 cusps, outer marginal teeth with (Felsenstein 1985) with 1000 replications 21—34 cusps. was performed to assess branch support on moderately pigmented, eyes pig- the resulting trees. Timing of branching mented. Cephalic tentacles with dorsal and events was estimated using the fossil-based ventral longitudinal cilial tracts and trans- calibration for mtDNA third position tran- verse tracts along outer edge of basal por- sitions (3—4% per My) and transversions tion of left tentacle. Ctenidium well devel- (0.42% per My) for the marine gastropod oped and wide, extending most of to entire genus Nucella (Collins et al. 1996) and may length of pallial cavity. Osphradium located be accurate only to the extent that evolu- near middle of ctenidium, narrow to ovate. tionary rates are comparable among mito- Renal organ with short pallial section; renal chondrial genes and among muricid and organ longitudinal, renal opening slightly cochliopinid gastropods. muscularized. Pericardium with short pal- VOLUME 115, NUMBER 1 173

Hal section; short efferent vessel between duct a little in front of posterior wall of pal- anterior end and posterior end of ctenidium lial cavity. Albumen gland very small, abut- or ctenidium overlapping anterior end of ting right side of bursa copulatrix. Female pericardium. Posterior stomach slightly ovoviviparous, with small to medium num- smaller than anterior chamber; single diges- ber of embryos brooded in large, thin- tive gland opening; small posterior caecum walled capsule gland. Brood pouch poste- present or absent; style sac about as long as riorly folded, folded section narrow, ven- remaining portion of stomach. tral; anterior opening terminal, musculari- Testis of vertical compound lobes, oc- zed, slightly recurved. Spermathecal duct cupying most of visceral coil behind stom- narrow to medium width, opening a little ach and partly overlapping posterior stom- behind brood pouch opening. ach chamber. Seminal vesicle beneath an- Remarks.—The two previously described terior portion of testis, coiled on digestive species that we allocate to Juturnia most gland behind stomach. Prostate gland ex- recently were placed in the genus Duran- tends into pallial roof for 33-67% length, gonella (Hershler 2001). Durangonella narrow-pyriform. Visceral vas defer ens Morrison, 1945 was erected for four species opens to ventral edge of prostate gland from from central Mexico which have smooth, near posterior edge to 33% anterior from elongate-conic shells. Morrison (1945) re- edge. Pallial vas deferens opens to ventral described the type species, Hydrobia see- edge of prostate gland slightly behind an- mani Fraunfeld, 1863, and described the terior edge, initial portion weakly recurved, other three congeners from empty shells. remainder straight or weakly undulating. Morrison presented but few details of soft Penis elongate-rectangular with little taper; parts, based on study of dried material of distal end blunt to evenly rounded, with the type species, and the only anatomical sub-terminal papilla, generously ciliated; criterion by which he distinguished Dur- inner edge with small narrow-triangular angonella from closely similar Tryonia was lobe positioned at about 67% of length its "different pattern" of penial ornament from base. Penial duct straight distally, near (Morrison 1945:18-19). A fifth species of straight to moderately undulating in basal Durangonella (D. coahuilae) subsequently portion. was described from Chihuahuan Desert Ovary simple, occupying very small part shells (Taylor 1966) and a sixth congener of visceral coil (< VA whorl). Proximal (D. kosteri), from the Pecos River basin, re- coiled oviduct narrow, thin-walled; distal cently was transferred to the genus (Her- portion broader, with posterior-oblique bend shler 2001). The two northern congeners or weak coil, usually containing sperm. (D. coahuilae, D. kosteri) have been shown Coiled oviduct and seminal receptacle duct to be members of the hydrobiid subfamily joined by very short duct at point where Cochliopinae, within which they form a former opens to posterior edge of albumen distinct group united by an unique suite of gland. Bursa copulatrix small, horizontally anatomical character-states (Hershler 2001). ovate or cylindrical, positioned well ante- They also have been shown to form a well rior to posterior edge of brood pouch; duct supported cochliopine sub-clade based on opening from anterior edge of pouch, nar- mtDNA sequences (Liu et al. 2001). How- row to medium width. Seminal receptacle ever, trenchant characterization of Duran- small to medium-sized, sub-globular, abut- gonella has been prevented by a paucity of ting left side of bursa copulatrix; duct open- information for southern congeners, none of ing from anterior edge of pouch, very short. which has been further studied since Mor- Sperm duct narrow, posterior section with rison's (1945) work. weak bend or kink, anterior portion straight, The type species of Durangonella, Hy- opening to ventral edge of spermathecal drobia seemani Frauenfeld, 1863, was de- 174 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

A

Fig. 1. Shells of Durangonella seemani. A. Durango City, Durango (USNM 251826). B. Syntype, Durango, N.W. Mexico (BMNH 20001099). Shell "A" is 3.02 mm tall; shell "B" is printed to the same scale. scribed from shells collected in "Durango, closely resemble syntypes of H. seemani N.W. Mexico" by Berthold Seemann dur- (Fig. lb) and are presumed by us to be con- ing his tenure as naturalist aboard the specific, although they are not assuredly H.M.S. Herald. Presumably Seeman made topotypes as Morrison (1945:19) claimed. this collection in 1849 or 1850 when he left Recently we examined rehydrated speci- the vessel, ventured inland from the Pacific mens from USNM 251826 and determined Coast, and visited the city of Durango and that Morrison's (1945, fig. 5) description nearby areas. However, neither his expedi- was in error as the male penes do not have tion narrative (Seeman 1853) nor his re- simple lobes but instead are ornamented sulting botanical monograph (Seeman with glandular papillae (Fig. 2). We also 1852-7) mention this collection. Frauenfeld observed in dissection that the female sper- (1863:1025; English translation in Morrison mathecal duct (= sperm tube) of these 1945:19) briefly described the shell of this snails opens a little in front of the posterior species, emphasizing its slender and conical wall of the pallial cavity which, together shape, deep suture, small aperture, and sim- with penial morphology, indicates that this ple lip; and subsequently illustrated a spec- snail belongs to the genus Tryonia (fide imen (Frauenfeld 1865, fig. 8). Morrison Hershler 2001). This is consistent with on- (1945) redescribed this species from speci- going field surveys in the Durango region mens that botanist Edward Palmer collected which have yielded snails with Tryonia-like (probably in 1896 fide USNM accession pa- anatomy (Hershler unpublished). pers; and field itinerary [McVaugh 1956]) Based on these new observations (made from a spring that then provided the do- subsequent to the recent synopsis of Dur- mestic water supply to Durango City. Shells angonella; Hershler 2001), we conclude of this sample (USNM 251826, Fig. la) that Durangonella Morrison, 1945 (type VOLUME 115, NUMBER 1 175

(Other similar-shelled regional cochliopines have a coiled sperm duct positioned entirely behind the posterior wall of the pallial cav- ity; Hershler 2001.) Juturnia differs from these two genera and all other members of subfamily in having a single, non-glandular lobe on the inner edge of the penis, which may be a synapomorphy for the genus. Ju- turnia is further distinguished from other regional cochliopines by its sub-globular seminal receptacle, weakly coiled renal ovi- duct, and storage of sperm in the coiled ovi- duct. Our phylogenetic analyses of mtCOI se- Fig. 2. Penis of Durangonella seemani, USNM quences (all of which resulted in the same 251826; scale equals 0.5 mm. Abbreviations—Pa, ter- tree topology) provided good (73%) sup- minal papilla; PI, penial lobe. port for monophyly of Juturnia (Fig. 3). Within this topology Juturnia was placed species, Hydrobia seemani) is a junior sub- within a poorly supported clade that also jective synonym of Tryonia Stimpson, contained two other genera which have an 1865. Accordingly, we have erected a new elongate sperm duct {, Spur- genus (Juturnia) for the two species from winkia) and , which has a the northern Chihuahuan Desert and Pecos coiled sperm duct. River basin which previously were assigned Distribution.—Lower Rio Grande region to Durangonella, but which are well-differ- (Fig. 4). Juturnia kosteri is distributed entiated from Tryonia and other cochliopine along the lower Pecos River, a major trib- genera. The three other species of Duran- utary to the lower Rio Grande. Juturnia gonella (all from central Mexico) cannot coahuilae lives in the Cuatro Cienegas ba- confidently be assigned to genus because sin (Coahuila, Mexico), whose northeast they are known only from shells and be- portion drains to the headwater region of cause the regional hydrobiid fauna remains the Rio Salado (Minckley 1969), which also poorly known. These species are "Duran- is tributary to the lower Rio Grande. Jutur- gonella" dugesiana Morrison, 1945 (type nia tularosae (described below) lives in a locality, Andocutin, Guanajuato; fide Taylor hydrographically closed basin in the south- 1966); "D." mariae Morrison, 1945 (one ern Rio Grande rift. meter below the present surface of the dry Habitat.—Spring-fed water bodies, typi- bed of the lake at Tlahuac, 20 kilometers cally consisting of headsprings and their east of Xochimilco, Distrito Federal); and stream outflows. Juturnia coahuilae also "D." pilsbryi Morrison, 1945 (Paso del lives in Laguna Grande, the terminus of a Rio, Colima). spring-fed drainage system (Hershler 1985). The shells of Juturnia broadly overlap in Waters inhabited by congeners range from size, shape, and sculptural features with near freshwater to hypersaline (e.g., Laguna those of several other epigean cochliopines Grande [=Laguna Salada], 309 g/1 total dis- of southwestern North America (Eremopyr- solved solids; Minckley & Cole 1968). gus, Littoridinops, Pseudotryonia, Tryon- Snails are benthic and typically live in soft ia). Among these taxa Juturnia shares with sediments. Littoridinops and Pseudotryonia an elon- Etymology.—From N. Latin Juturna, the gate sperm duct which opens to the sper- Roman goddess of wells and springs. Gen- mathecal duct in the posterior pallial cavity. der feminine. 176 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Eremopyrgus elegans 100

-Eremopyrgus eganensis

• Zetekina sp. 2 100

Zetekina sp. 1

Pseudotryonia hrevissima 95

96 Pseudotryonia alamosae

Aphaostracon sp. (AS) 100

1 Aphaostracon sp. (LP)

Littoridinops monroensis 99 57 — Littoridinops palustris

Pyrgophorus platyrachis

- Juturnia kosteri 97

73 - Juturnia tularosae

Juturnia coahuilae

• Spurwinkia salsa

• Tryonia clathrata 50

85 • Tryonia rowlandsi

Tryonia aequicostata

• Mexipyrgus carranzae 50 changes

Fig. 3. Single shortest length tree based on maximum parsimony analysis of mtCOI sequence data. CI, 0.57; TL, 1473. Bootstrap percentages are given when >50%. The two specimens of J. tularosae were identical in their sequences. Samples of Aphaostracon sp. were from Alexander Springs, Lake Co, FL (AS); and Lake Panasoffkee, Sumter Co., FL (LP). Data for species other than /. tularosae are from Hershler et al. (1999) and Liu et al. (2001).

Juturnia tularosae, new species conic, width/height 57—80%, with convex Tularosa juturnia spire sub-equal to shell width, 1.70-4.84 Diagnosis.—A small to medium-sized mm tall, whorls 4.0-6.0. Periostracum light Juturnia which differs from congeners in tan. Protoconch of about 0.75 whorl, sur- shell having more convex teleoconch face slightly roughened to corrugated (Fig. whorls (which frequently are shouldered); 5C, D). Teleoconch whorls highly convex, and female genitalia having smaller bursa evenly rounded or wider above than below, copulatrix and narrower spermathecal duct. smooth apart from well-developed growth Description.—Shell (Fig. 5A, B) ovate- lines and occasional weak spirals, shoulders VOLUME 115, NUMBER 1 177

Fig. 4. Drainage map of Rio Grande basin and adjacent areas showing distribution of Juturnia. often well-developed. Teleoconch some- (Fig. 7A). Attachment scar margin slightly times with weak demarcation at 1.25-1.75 thickened on inner side; outer margin very whorl whorls reflecting cessation of growth weakly rimmed (Fig. 7B). in brood pouch. Aperture medium-sized. Radula with about 53 well-formed rows Inner lip thin or slightly thickened, straight of teeth; ribbon length, 579 u.m; ribbon or weakly rounded, usually complete and width, 120 urn; central tooth width, 24.8 narrowly or broadly adnate across parietal u.m. Dorsal edge of central teeth moderately wall, sometimes narrowly disjunct; lip indented, basal process usually rounded be- sometimes narrowly reflected in columellar low, base even with lateral margins, median region. Outer lip orthocline or weakly pro- cusps slightly longer than laterals, pointed socline, straight or weakly sinuate. Larger or rounded distally; lateral cusps 4-5; basal specimens often having last 0.25-0.5 whorl cusps small (Fig. 8A). Lateral teeth having loosened from body whorl. Umbilicus ri- 3-4 cusps on inner side and 6-7 cusps on mate to broad. Females larger than males, outer side; outer wing well-flexed, curved, but having similarly shaped shells (Fig. 6, length 210-240% width of tooth face; cen- Table 1). Shell parameters are in Table 1. tral cusp pointed or rounded distally, dorsal Operculum flat or slightly concave, edge weakly concave; basal tongue mod- ovate, light amber. Outer side nearly erately developed (Fig. 8B). Inner marginal smooth, edges of later whorls weakly frilled teeth with 26—33 cusps, cutting edge on 178 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 5. Scanning electron micrographs of shells of J. tularosae, USNM 894803. A, B. Variation in shell shape; scales equal 1 mm. C, D. Variation in protoconch appearance; scales equal 100 u,m. outer side of tooth, occupying ca. 33% of teeth larger than those on outer marginals tooth length (Fig. 8D). Outer marginal teeth (Fig. 8C). with 26-33 cusps, cutting edge on inner Snout dark grey or black. Cephalic ten- side of tooth, occupying ca. 30% of tooth tacles light-medium grey, sometimes darker length (Fig. 8E). Cusps on inner marginal near bases and often having a black patch VOLUME 115, NUMBER 1 179

Fig. 6. Shells of /. tularosae. A female (holotype), USNM 894737. B. male, USNM 854956. Shell "A" is 3.77 mm tall; shell "B" is printed to the same scale. near distal tips. Yellow granules scattered tapered, with rounded tips. Base of tenta- along tentacle length, with dense white cles without pronounced eyelobe. Foot ca. patch of granules clustered on neck. Foot 250% longer than wide; lateral wings well- dark grey, with numerous embedded clear- developed; anterior edge slightly convex. white granules. Mantle, visceral coil grey- Ctenidium extending to near mantle edge black, pigment particularly dark over testis. and overlapping pericardium posteriorly. Distal penis having large grey-black patch Ctenidial filaments ca. 27, extending to near of melanic pigment; proximal portion pig- rectum, narrowly triangular with apices po- mented with numerous yellow granules. sitioned slightly to left of mid-line; lateral Snout rectangular, distal lips slightly ex- surfaces having well developed ridges. Hy- panded. Tentacles narrow, ca. 150% length pobranchial gland weakly developed along- of snout (while animal crawling), slightly side rectum in posterior pallial cavity. Os-

Table 1.—Shell measurements of holotype (USNM 894737) and paratypes (USNM 894741) of Juturnia tularosae. t values are given for comparisons of means among sexes (95% confidence level). *Sexual dimorphism significant (P < 0.01).

SL sw LBW WBW AL AW TW SW/SL

Holotype 3.14 1.87 2.26 1.75 1.27 1.15 4.75 0.59 Paratypes d (« = 22) X 2.44 1.61 1.77 1.41 1.03 0.97 4.55 0.66 SD 0.23 0.13 0.15 0.11 0.09 0.08 0.24 0.03 9 (n = 30) X 2.91 1.92 2.06 1.69 1.20 1.14 4.90 0.67 SD 036 0.14 0.22 0.15 0.12 0.09 0.25 0.04 t 5.81* 8.46* 5.63* 7.65* 6.17* 7.16* 5.17* 0.260 ISO PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

A B

• .v

Fig. 7. Scanning electron micrographs of opercula of J. tularosae, USNM 894803; scales equal 100 p.m. A. outer side. B. inner side. phradium ovate, positioned slightly poste- pulatrix. Sperm duct (Sdu) kinked posteri- rior to middle of ctenidial axis. Renal organ orly, otherwise straight. Duct to albumen narrow. Style sac about as long as remain- gland (Dag) issuing from oviduct slightly ing portion of stomach, posterior caecum of distal to coiled section. Spermathecal duct stomach small, triangular. (Sd) narrow, although broadening distally, Testis 1.0-1.25 whorl, slightly overlap- opening slightly raised. Albumen gland ping posterior stomach chamber. Seminal (Ag) abutting the ventro-right lateral sur- vesicle consisting of a few coils. Prostate face of the bursa copulatrix. Brood pouch gland small, yellow, bean-shaped, 50—67% (Bp) containing 2-5 shelled embryos; an of length in pallial roof, ovate in section, additional series of weakly developed em- walls thickened all around (Fig. 9). Visceral bryos often present in posteriorly folded vas deferens opening near posteroventral section. Anterior opening of brood pouch edge of pallial gland; pallial vas deferens (Obp) muscular, raised, positioned along- straight or weakly kinked proximally on side anus. columellar muscle. Penis medium-sized, Type material.—A live-collected series straight or slightly curved; distal portion obtained by Craig Stockwell on 4 Feb 2001 well expanded in living specimens (Fig. from Salt Creek at Range Road 316 cross- 10A), evenly rounded or weakly expanded ing, White Sands Missile Range, Tularosa in preserved material (Fig. 10B). Narrow basin, Sierra County, New Mexico, Town- penial duct straight, or weakly undulating ship 12S, Range 6E, NE 1/4 section 16. The in middle section. holotype (USNM 894737, Fig. 5A) is a Ovary positioned slightly behind stom- dried female from this series, while remain- ach. Distal genitalia shown in Fig. 11. Re- ing specimens (also dried) are reposited as nal oviduct loop (Ro) posterior-oblique, paratypes (USNM 894741). Three addition- bulging with sperm. Bursa copulatrix (Bu) al paratype series from the type locality ovate, sometimes weakly pigmented, posi- were collected at other times during 1999- tioned a little behind posterior pallial wall, 2001 (USNM 854956, USNM 894803, duct (Dbu) narrow. Seminal receptacle (Sr) USNM 905339). small, abutting anterior portion of bursa co- Other material examined.—Dried shells Fig. 8. Scanning electron micrographs of radula of J. tularosae, USNM 894803; scales equal 10 u,m. A. Central teeth. B. Lateral teeth. C. Portion of radula ribbon showing relative sizes of cusps on outer and inner marginal teeth. D. Inner marginal tooth. E. Outer marginal teeth. from Salt Creek, in a canyon ca. 6.4 km ranges and is supported on the basin floor above Range Road 316, Tularosa basin, Si- by the discharge of Salt Springs. A field erra County, New Mexico, Township 12S, survey during March, 2001 indicated that Range 6E, NE 1/4 section 3 (USNM snails were restricted to a central reach of 894738). Salt Creek, extending from a canyon ca. 6.4 Distribution and habitat.—Juturnia tu- km north of Range Road 316 to ca. 0.5 km larosae apparently is endemic to Salt Creek downfiow from this road. The northern end (Meinzer & Hare 1915, pi. VIIIC), a major of the snail range coincides with a waterfall drainage of the northern Tularosa basin (ca. 2 m high) at the head of this canyon. which heads in the San Andres and Oscura During recent years (1995—1999) dis- 182 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 9. Prostate gland (left side) of J. tularosae, USNM 894956; scale equals 0.5 mm. Abbreviations— Pr, prostate gland; Pvd, pallial vas deferens; Vvd, vis- ceral vas deferens. Fig. 10. Penes (left side) of J. tularosae; scale equals 250 u,m. A. Live specimen, USNM 894803. B. Formalin-fixed specimen, USNM 894956. Abbrevia- charge of Salt Creek in the vicinity of the tions—Pa, terminal papilla; Pd, penial duct; PI, penial type locality of J. tularosae averaged 32.28 lobe. liters/second (Ortiz & Lang 1996, Ortiz et al. 1997, 1998, 1999). During this time pe- formed a strongly supported (97%) sub- riod the stream has been subject to periodic clade. In addition to the characters listed flooding, usually during the summer, with a above (see Diagnosis) Juturnia tularosae maximum discharge of 1982.18 liters/sec- differs from J. kosteri by the more deeply ond. Water temperature varied from 7.0- indented dorsal edge of the central radular 33.5°C; dissolved oxygen was typically teeth (compare Fig. 8A with Hershler 2001, high, with an average of 9.29 mg/liter; and fig. 12A), squatter shape of prostate gland, pH was fairly constant, with an average of very short pallial extension of the female 8.07. Salt Creek is saline and both alkali- sperm duct, papilla-like opening of the (Na and K) and chloride-rich. These water brood pouch, and occasional pigment on quality data are similar to those of earlier the bursa copulatrix (compare Fig. 11 with studies of Salt Creek (Meinzer & Hare Hershler 2001, fig. 13). 1915, Miller & Echelle 1975, Stockwell & Mulvey 1998). Discussion Etymology.—A geographical name refer- ring to endemism of this snail in Tularosa The phylogenetic analysis of mtCOI se- basin, southeast New Mexico. quences (Fig. 3) indicates that monophylet- Remarks.—This species, together with J. ic Juturnia is more closely related to brack- kosteri, is distinguished from J. coahuilae ish water cochliopines confined to the west- by its much broader shell (compare Figs. 5, ern Atlantic and Gulf of Mexico margins 6 with Hershler 1985, fig. 33), less elongate (Littoridinops, Pyrgophorus, Spurwinkia) bursa copulatrix, and more anteriorly posi- than to other taxa of southwestern North tioned seminal receptacle (compare Fig. 11 America (Eremopyrgus, Mexipyrgus, Tryon- with Hershler 1985, fig. 36). Our mtCOI- ia) and is consistent with an origin associ- based tree (Fig. 3) was consistent with mor- ated with inland invasion by Gulf Coastal phology in that the former two species ancestors. These sequence data also suggest VOLUME 115, NUMBER 1 183

Fig. 11. Female genitalia (left side) of J. tularosae, USNM 894956; scale equals 500 (xm. A. Brood pouch (Bp) and associated structures. B. Bursa copulatrix (Bu) and associated structures. Abbreviations—Bp, brood pouch; Bu, bursa copulatrix; Dag, duct connecting oviduct and albumen gland; Obp, opening of brood pouch; Osd, opening of spermathecal duct; Pf, posterior folded section of brood pouch; Ro, renal oviduct; Sd, sper- mathecal duct; Sdu, sperm duct; Sr, seminal receptacle. that divergence of Juturnia occurred 4.3— Laramide uplifts (e.g., Rocky Mountains, 14.9 Ma (Miocene-early Pliocene) and thus Sierra Madre Orientate; Winker 1982, Gal- preceded assembly of the modern Rio loway et al. 1991) and flowed east and Grande drainage, which was not completed southeast to the Texas Gulf Coast during until its master stream breached the Lake the early-mid Tertiary (Thomas 1972, Bel- Cabeza de Vaca complex (sensu Strain cher 1975, Belcher & Galloway 1977, 1966) in northeastern Chihuahua and north- Gather 1991, Christiansen & Yeats 1992). western Texas ca. 2.5-2.0 Ma (Gustavson Note that the Rio Grande region has had a 1991, Mack 1997, 2001) and subsequently complex biogeographic history with respect penetrated to the sea. We conjecture that to the Gulf of Mexico and other venues of coastal progenitors did not penetrate inland inland transport of marine biota, which may through the Rio Grande but instead along have ranged from stranding of 'relicts' in the drainages which headed on the flanks of conjunction with recession of Cretaceous 184 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON seas (Bowman 1981, Holsinger 1986) to Gustavson & Winkler 1988). The closely Quaternary aerial dispersal on or in birds proximate occurrence of marine-like biota (Bachuber & McClellan 1977), also may in Great Plains drainage just east of the Pe- have occurred. cos divide (Creel 1964) is congruent with Progenitors may have been euryhaline this hypothesis. The timing of drainage di- and capable of penetrating into freshwater version has not been well established, but environments (fide Hershler et al. 2002). presumably occurred following deposition Alternatively, ancestral snails may have of the regionally widespread Ogallala For- been salt dependent in which case their in- mation, which ended ca. 3.5-5 Ma (Hawley land dispersal was more tightly constrained 1993 a). These dates conform well to our by water chemistry. Taylor (1985:318-319) mtDNA evidence, which suggests that di- speculated that the Pecos River, which vergence occurred 4.2-5.9 Ma. flows through a region having a long his- Taylor (1985) suggested that regional tory of evaporite dissolution (Bachman isolation of other marine-like mollusks was 1976) and is highly mineralized along por- effected by inception of the southern reach tions of its length (Gustavson & Finley of the freshwater Rio Grande in the Pleis- 1985), functioned as an early saline conduit tocene, which disrupted and diluted an an- for coastal molluscan colonists into this re- cestral saline Pecos River (through which gion. Although we disagree with aspects of marine colonists dispersed). This assumes this interpretation of Pecos River history that the Pecos River was through-flowing (see below), we note that the Rio Salado (to to the Gulf prior to its integration with the which J. coahuilae is endemic) also con- Rio Grande and is supported by provenance tributes a large amount of salt to the Rio interpretations of gravels on the Edwards Grande (Miyamoto et al. 1995) derived in Plateau and the lower Rio Grande (Thomas part from dissolution of (Cretaceous) evap- 1972, Belcher 1975). However, an analysis orites (Back et al. 1977, Lesser & Lesser of river meander patterns suggests that the 1988) and thus may have had a long brack- lower Pecos River early headed on the Ed- ish-saline history. (This river may date to wards Plateau and did not integrate with the the Paleocene; Galloway et al. 1991:302.) Pecos depression (where snails are endem- Note that , a member of a ic) until ca. 0.6 Ma, following incision of salt-adapted gastropod genus typically as- the Pecos Canyon through headward ero- sociated with coastal margins (Abbott sion (Amsbury & Dietrich 1987, Amsbury 1958), is distributed (together with Jutur- & Hayward 1999; also see Hills 1984:264). nia) in upper segments of the Pecos River The occurrence of a fossil herring in the and Rio Salado basins (Taylor 1987), which Gatuna Formation in southeastern New further implies a close historical relation- Mexico (Miller 1982), whose age is brack- ship between these areas and saline habi- eted at 13-0.5 Ma (Powers & Holt 1993), tats. suggests that marine biota may have been We attribute vicariance of the Juturnia present in the region well prior to integra- sub-clade in southeastern New Mexico (J. tion of the Pecos River drainage in confor- kosteri and J. tularosae) to the well docu- mance with our hypothesis. We agree with mented capture of segments of the old Taylor (1985) that marine-like Pleistocene drainage system which extended from the gastropods from Terrell County, Texas (An- southern Rocky Mountains across the drews 1977:92,98,180) could have dis- southern High Plains (Plummer 1933, fig. persed into this region along the ancestral 51 A) by the Pecos depression (fide Horberg lower Pecos River, however we attribute the 1949), which formed through dissolution of Pleistocene occurrence of the brackish-wa- bedded Permian salt followed by regional ter clam Rangia cuneata in the Pecos de- subsidence (Gustavson & Finley 1985, pression (Metcalf 1980, Taylor 1985, fig. VOLUME 115, NUMBER 1 185

30) to dispersal along the same drainage lected under the auspices of New Mexico system that was invaded by Juturnia pro- Game and Fish permit #2887. Hildegard genitors. Reiser, Julie Lessard, Amber Moore and Subsequent vicariance of J. tularosae es- David Rogowski assisted with field collec- timated at 2.2-3.0 Ma (based on third po- tions and habitat surveys, and Robert Myers sition transitions only as there were no facilitated access to White Sands Missile transversion changes at this position) is Range. Fred Naggs and Joan Pickering enigmatic in that geologic evidence sug- (BMNH) loaned specimens. Bill Cox gests that Tularosa basin was endorheic dur- (Smithsonian Institution Archive) assisted ing the Pleistocene and opened not to the with locating museum accession records of east but to the south during the late Plio- Edward Palmer's molluscan collections. cene (5-2 Ma), when it served as a periodic Karie Darrow prepared the anatomical il- distributary of the ancestral upper Rio lustrations, Molly K. Ryan drew the holo- Grande (Mack et al. 1996, 1997, Mack type, and Yolanda Villacampa prepared the 1999). (Although pluvial Lake Otero [fide scanning electron micrographs (all USNM). Herrick 1904] flooded Tularosa basin up to The first author thanks David Amsbury and 1250 m elevation during its late Pleistocene Dennis Powers for helpful discussion con- highstand, it never breached its basin [Kot- cerning geologic history of the Rio Grande tlowski 1958, Miller 1981, Hawley 1993b, region. Pittenger & Springer 1999].) Any prior his- torical drainage relationship between this Literature Cited area and the Pecos region presumably was disrupted by late Tertiary extensional fault- Abbott, R. T. 1958. The gastropod genus Assiminea in ing, but is not well constrained as Tularosa the Phillipines.—Proceedings of the Academy basin, which consists of two adjacent deep- of Natural Sciences of Philadelphia 110:213- 278. ly incised half-grabens (Lozinsky & Bauer Adams, D. C, & G. R. Keller. 1994. Crustal structure 1991, Adams & Keller 1994), is bounded and basin geometry in south-central New Mex- by mountain blocks which have had long ico.—Geological Society of America Special uplift histories (Kelley & Chapin 1997) ex- Paper 291:241-255. tending into the Quaternary (Machette Amsbury, D. L., & J. W. Dietrich. 1987. Some impli- cations of incised meanders in the southern Ed- 1987). The close relationship implied by wards Plateau.—Geological Society of America our data between this component of the Rio Abstracts with Programs 19(3): 115. Grande Rift and the southern Great Plains , & O. T. Hayward. 1999. Quaternary erosional is paralleled by allozymic evidence of re- landscapes of central and North Texas.—Amer- gional pupfishes (Echelle & Echelle 1992) ican Association of Petroleum Geologists An- nual Meeting Expanded Abstracts 1999:5. and suggests a need for further study of the Andrews, J. 1977. Shells and shores of Texas, 2nd edi- functionality of the regional drainage divide tion. University of Texas Press, Austin and Lon- during the late Neogene. don, 365 pp. Bachman, G. O. 1976. Cenozoic deposits of south- eastern New Mexico and an outline of the his- Acknowledgments tory of evaporite dissolution.—Journal of Re- search, U.S. Geological Survey 4:135-149. Research funding was provided, in part, Bachuber, E W., & W. A. McClellan. 1977. Paleo- by awards from the Smithsonian Institu- ecology of marine Foraminifera in the pluvial tion's Office of Biodiversity Programs and Estancia Valley, central New Mexico.—Quater- the Department of Defense Legacy Re- nary Research 7:254-267. source Program (Grant # DACA87-00-H- Back, W., J. M. Lesser, & B. B. Hanshaw. 1977. Struc- tural and stratigraphic occurrence of "bad wa- 0014 administered by Dr. M. Hildegard Re- ter" in Coahuila, Mexico.—Geological Society iser, 49 CES/CEV, Environmental Flight, of America Abstracts with Programs 9(7):885. Holloman Air Force Base). Snails were col- Belcher, R. C. 1975. The geomorphic evolution of the 186 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Rio Grande.—Baylor Geological Studies Bul- geomorphic evolution of the Texas Panhandle letin 29:1-64. and northeastern New Mexico—case studies of , & W. E. Galloway. 1977. Evidence for early structural controls on regional drainage devel- Tertiary Rio Grande drainage.—Geological opment.—Bureau of Economic Geology Report Society of America Abstracts with Programs of Investigations 148:1-42. 9(1):5. , & D. A. Winkler. 1988. Depositional fades of Bowman, T. E. 1981. Thermosphaeroma milleri and T. the Miocene-Pliocene Ogallala Formation, smithi, new sphaeromatid isopod northwestern Texas and eastern New Mexico.— from hot springs in Chihuahua, Mexico, with a Geology 16:203-206. review of the genus.—Journal of Bi- Hawley, J. W. 1993a. The Ogallala and Gatuna For- ology 1:105-122. mations in the southeastern New Mexico re- Gather, S. M. 1991. Late Laramide paleodrainage pat- gion, a progress report.—New Mexico Geolog- terns and timing of sedimentation in the south- ical Society Guidebook 44th Field Conference: ern Rocky Mountains and northwest Gulf of 261-269. Mexico.—Geological Society of America Ab- . 1993b. Geomorphic setting and late Quater- stracts with Programs 23(4):10. nary history of pluvial-lake basins in the south- Christiansen, R. L., & R. S. Yeats. 1992. Post-Lar- ern New Mexico region.—New Mexico Bureau amide geology of the U.S. Cordilleran region. of Mines and Mineral Resources Open-File Re- Pp. 261-406 in B. C. Burchfiel, P. W. Lipman, port 391:1-28. & M. L. Zoback, eds., The Cordilleran orogen, Herrick, C. L. 1904. Lake Otero, an ancient salt lake conterminous U.S. Geological Society of Amer- basin in south-eastern New Mexico.—American ica, Boulder, 724 pp. Geologist 34:174-189. Collins, T. M., K. Frazier, A. R. Palmer, G. J. Vermeij, Hershler, R. 1985. Systematic revision of the Hydro- & W. M. Brown. 1996. Evolutionary history of biidae (: Rissoacea) of the Cuatro northern hemisphere Nucella (Gastropoda, Mur- Cienegas Basin, Coahuila, Mexico.—Malacol- icidae): molecular, morphological, ecological, ogia 26:31-123. and paleontological evidence.—Evolution 50: . 1998. A systematic review of the hydrobiid 2287-2304. snails (Gastropoda: Rissooidea) of the Great Creel, G. C. 1964. Hemigrapsus estellinensis: a new Basin, western United States. Part I. Genus Pyr- grapsoid from north Texas.—Southwestern gulopsis.—The Veliger 41:1-132. Naturalist 8:236-241. . 2001. Systematics of the North and Central Echelle, A. A., & A. F. Echelle. 1992. Mode and pat- American aquatic snail genus Tryonia (Rissooi- tern of speciation in the evolution of inland dea: Hydrobiidae).—Smithsonian Contributions pupfishes in the Cyprinodon variegatus com- to Zoology 612:1-53. plex (Teleostei: Cyprinodontidae): an ancestor- , & W. F. Ponder. 1998. A review of morpho- descendant hypothesis. Pp. 691-709 in R. L. logical characters of hydrobioid snails.—Smith- Mayden, ed., Systematic^, historical ecology, sonian Contributions to Zoology 600:1-55. and North American freshwater fishes. Stanford , H.-P. Liu, & J. J. Landye. 2002. A new species University Press, Stanford, 969 pp. of aquatic snail of the genus Eremopyrgus (Hy- Felsenstein, J. 1985. Confidence limits on phylogenies: drobiidae: Cochliopinae) from the Chihuahuan an approach using the bootstrap.—Evolution Desert, Mexico: phylogenetic relationships and 39:783-791. biogeography.—Journal of Molluscan Studies Frauenfeld, G. R. von. 1863. Vorlaufige aufzahlung der (in press). arten der gattungen Hydrobia Htm. Und Amni- , , & M. Mulvey. 1999. Phylogenetic cola Gld. Hldm. In der kaiserlichen und in relationships within the aquatic snail genus Cuming's sammlung.—Verhandlungen der Tryonia: implications for biogeography of the Kaiserlich-Koniglichen Zoologisch-botanischen North American Southwest.—Molecular Phy- Gesellschaft in Wien 13:1017-1032. logenetics and Evolution 13:377-391. . 1865. Zoologische Miscellen.—Verhandlun- Hills, J. M. 1984. Sedimentation, tectonism, and hy- gen der Kaiserlich-Koniglichen Zoologisch-bo- drocarbon generation in Delaware Basin, west tanischen Gesellschaft in Wien 15:525-536. Texas and southeastern New Mexico.—Ameri- Galloway, W E., D. G. Bebout, W. L. Fisher, J. B. can Association of Petroleum Geologists Bul- Dunlap Jr., R. Cabrera-Castro, J. E. Lugo-Ri- letin 68:250-267. vera, & T. M. Scott. 1991. Cenozoic. Pp. 245- Holsinger, J. R. 1986. Zoogeographic patterns of North 432 in A. Salvador, ed., The Gulf of Mexico American subterranean amphipod crustaceans. Basin. Geological Society of America, Boulder, Pp. 85-106 in R. H. Gore & K. L. Heck, eds., 568 pp. Crustacean biogeography. A.A. Balkema, Rot- Gustavson, T. C, & R. J. Finley. 1985. Late Cenozoic terdam & Boston, 292 pp. VOLUME 115, NUMBER 1 187

Horberg, L. 1949. Geomorphic history of the Carlsbad the southern Rio Grande rift, USA.—Sedimen- Caverns area, New Mexico.—Journal of Geol- tology 44:637-652. ogy 57:464-476. Meinzer, O. E., & R. F. Hare. 1915. Geology and water Kelley, S. A., & C. E. Chapin. 1997. Cooling histories resources of Tularosa basin, New Mexico.— of mountain ranges in the southern Rio Grande United States Geological Survey Water-Supply rift based on apatite fission-track analysis-a re- Paper 343:1-317. connaissance survey.—New Mexico Geology Metcalf, A. L. 1980. Fossil Rangia cuneata (Mactri- 19(1):1-14. dae) in Eddy County, New Mexico.—Nautilus Kottlowski, F. E. 1958. Lake Otero-second phase in 94:2-3. formation of New Mexico's gypsum dunes.— Miller, R. R. 1981. Coevolution of deserts and pup- Bulletin of the Geological Society of America fishes (genus Cyprinodori) in the American 69:1733-1734. [abstract] Southwest. Pp. 39-94 in R. J. Naiman & D. L. Lesser, J. M., & G. Lesser. 1988. Region 9, Sierra Ma- Soltz, eds., Fishes in North American deserts. dre Orientale. Pp. 89-92 in W. Back, J. S. Ro- John Wiley & Sons, New York, 552 pp. senshein, & P. R. Seaber, eds., Hydrogeology. . 1982. First fossil record (Plio-Pleistocene) of Geological Society of America, Boulder, 524 Threadfin Shad, Dorosoma petenense, from the pp. Gatufia Formation of southeastern New Mexi- Leviton, A. E., R. H. Gibbs, Jr., E. Heal, & C. E. co.—Journal of Paleontology 56:423-425. Dawson. 1985. Standards in herpetology and , & A. A. Echelle. 1975. Cyprinodon tularosa, ichthyology, part I: standard symbolic codes for a new cyprinodontid fish from the Tularosa ba- institutional resource collections in herpetology sin, New Mexico.—Southwestern Naturalist 19: 365-377. and ichthyology.—Copeia 1985:802-832. Minckley, W L. 1969. Environments of the Bolson of Liu, H.-R, R. Hershler, & F. G. Thompson. 2001. Phy- Cuatro Cienegas, Coahuila, Mexico with special logenetic relationships of the Cochliopinae reference to the aquatic biota.—University of (Rissooidea: Hydrobiidae): an enigmatic group Texas at El Paso Science Series 2:1-65. of aquatic gastropods.—Molecular Phylogenet- , & G. A. Cole. 1968. Preliminary limnologic ics and Evolution 21(l):17-25. information on waters of the Cuatro Cienegas Lozinsky, R. P., & P. W. Bauer. 1991. Structure and basin, Coahuila, Mexico.—Southwestern Natu- basin-fill units of the Tularosa basin.—New ralist 13:421-431. Mexico Geological Society 42nd Annual Field Miyamoto, S., L. B. Fenn, & D. Swietlik. 1995. Flow, Conference:?-! 1 (+ references). salts, and trace elements in the Rio Grande: a McVaugh, R. 1956. Edward Palmer, plant explorer of review.—Texas Water Resources Institute Tech- the American West.—University of Oklahoma nical Completion Report TR-169:l-30. Press, Norman, 430 pp. Morrison, J. P. E. 1945. Durangonella, a new hydro- Machette, M. N. 1987. Preliminary assessment of pa- biine genus from Mexico, with three new spe- leoseismicity at White Sands Missile Range, cies.—The Nautilus 59:18-23. southern New Mexico; evidence for recency of Ortiz, D., & K. M. Lang. 1996. Water resources data: faulting, fault segmentation, and repeat intervals New Mexico Water Year 1996.—United States for major earthquakes in the region.—United Geological Survey Water-Data Report NM-95- States Geological Survey Open-file Report 87- 1:1-446. 0444:1-46. , , & L. Seal. 1997. Water resources Mack, G. H. 1999. Neogene evolution of lakes and data: New Mexico water year 1997.—United rivers in extensional basins of southwestern States Geological Survey Water-Data Report New Mexico.—Geological Society of America NM-97-1:1-574. Abstracts with Programs 31(7):444. , , & —. 1998. Water resources . 2001. Evolution of Cenozoic extensional data: New Mexico water year 1998, vol. 1.— block faulting and sedimentation in the southern United States Geological Survey Water-Data Rio Grande Rift, New Mexico.—Geological Report NM-98-1:1-404. Society of America Abstracts with Programs , , . 1999. Water resources data: 33(5):47. New Mexico water year 1999, vol. 1.—United , W. R. Seager, & W C. Mclntosh. 1996. Evo- States Geological Survey Water-Data Report lution of the southern Rio Grande Rift, south- NM-99-L1-402. central New Mexico.—Geological Society of Pittinger, J. S., & C. L. Springer. 1999. Native range America Abstracts with Programs 28(7):516. and conservation of the White Sands Pupfish , D. W Love, & W. R. Seager. 1997. Spillover (Cyprinodon tularosa).—Southwestern Natural- models for axial rivers in regions of continental ist 44:157-165. extension: the Rio Mimbres and Rio Grande in Plummer, F. B. 1932 [1933]. Cenozoic systems in Tex- IE PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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