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Pyrgulopsis madridensis Perez, sp. nov. urn:lsid:zoobank.org:act: F22BFC52-391E-41FB-877C-D27360C1FFE3 Figs. 4A–H, 5A–D, 6A–D. Diagnosis. A low-conical to trochoform snail with visible growth lines on teleoconch. Aperture ovate and usually separated from body whorl in adults with a distinct groove. Mantle tissue pigmented gray-black or unpigmented. Sharply pointed median cusp of lateral radular teeth. Penial lobe much shorter than filament, with large terminal gland and numerous large penial glands on the lobe. Types. Holotype ANSP A492826, paratypes ANSP A492828, all from Madrid Falls, Presidio County, Texas. Type locality. Madrid Falls, Big Bend Ranch State Park, 29.37975 W, -103.88411 N, Presidio County, TX Additional materials examined. ANSP A492830, South Fork, Crawford-Smith Canyon, Big Bend Ranch State Park, 29.39205 W, -103.87243 N, Presidio County, TX. Etymology. We use the specific epithet “ madridensis ” reflecting the type locality, a spectacular waterfall in desert west Texas. Remarks. We propose the common name “Madrid Falls springsnail”. Description. Shell low-conical to ovate-conic, pale to light brown in color (Fig. 4A–C; 5A–B). In life, the shell usually appears partially or completely black in color due to the shell being covered in dark-colored biofilm. The spire is relatively short with convex to slightly shouldered whorls. Whorls with a distinct suture. Protoconch with the initial portion lightly punctate (Fig. 5C–D). Teleoconch with strong growth lines. Aperture ovate, angled above, sometimes appressed to body whorl, but usually slightly separated with a distinct groove or gap between aperture and previous whorl. Peristome complete, inner lip sometimes nearly straight, slightly reflected in the columellar portion and outer lip. Outer lip simple, slightly prosocline. Umbilicus narrow. Average shell measurements for n = 15 adult individuals from Madrid Falls: shell height = 1.73 mm (SD = 0.20), shell width = 1.41 mm (SD = 0.12), body whorl width = 1.21 mm (SD = 0.11), aperture height = 0.95 mm (SD = 0.08), aperture width = 0.85 mm (SD = 0.10), number of whorls = 3.47 (SD = 0.27). Average shell measurements for n = 15 adult individuals from Crawford-Smith Canyon: shell height = 2.14 mm (SD = 0.27), shell width = 1.53 mm (SD = 0.13), body whorl width = 1.30 mm (SD = 0.13), aperture height = 1.05 mm (SD = 0.08), aperture width = 0.93 mm (SD = 0.10), number of whorls = 3.98 (SD = 0.06). Operculum (Fig. 4G, H) narrowly ovate, flat, light amber, except for pale narrow fringe; eccentric nucleus, paucispiral growth pattern.Attachment scar distinct and ending with gap before columellar edge. Attachment region callus moderately thick. Growth lines distinct, simple. Outer margin thin but with raised ridge along inner margin between nucleus and columellar edge. Pale, outer margin wider on the rest of the operculum. Central radular tooth (Fig. 6A–B, D) with moderately to deeply indented dorsal edge; lateral cusps 4–6; central cusp sharply pointed, considerably broader and 2X longer than lateral cusps, basal cusps narrowly elongate, straight, with slight buttress at base. Basal process medium width, broadly v-shaped; basal sockets deep. Lateral tooth face tall, rectangular, with broad neck and inner wing; central cusp large, wide, and blade-like, lateral cusps 3 (inner) and 3 (outer); on one individual, a few of the central cusps of the lateral tooth at the end of the radula were notched at tip; large, blunt basal projection and outer wing long and broad with distinct ridge along outer edge. Inner marginal teeth having 16-18 cusps; fourth cusp from outer edge enlarged. Outer marginal teeth (Fig. 6C) rounded and spoonlike, with a narrow neck, inner edge with a long wing and with 12-20 on the outer surface and ~20 small cusps on the inner surface pointed but small. Tentacles pale with scattered patches of pigmentation on dorsal surface. Snout dark gray-black, light colored on distal edge. Foot darkly pigmented on part of anterior surface, and light colored beneath. Opercular lobe black along anterior edge with pigment patches elsewhere. Neck pigmented along edges, pale below. In some individuals visceral coil and pallial roof both darkly pigmented, purplish-black. In others, dark purplish pigment occurs in large patch near entrance to mantle cavity, then sparsely on rest of body. Ctenidium occupies a small portion of the pallial cavity, at anterior end filaments triangular, broader than high, with central portion of filaments elongate, roughly triangular, both tall and wide at the base, ~15. Osphradium elongate, long, opposite posterior end to middle of ctenidium, anterior end simple. Digestive gland occupies most of first two whorls, with highly convoluted intestine leading to stomach. Stomach as expected for Pyrgulopsis with ~ equally sized chambers. Rectum straight with anus near edge of mantle. Testes sac-like lobe, elongate bean-shaped prostate. Penis large (Fig. 4E, F); filament medium length, fingerlike, rounded, tapered at distal tip; lobe shorter than filament, broad. Penial gland covers the length of the filament, distal to the body, does not appear bifurcate. Dorsal gland 1 (Dg1) robust, broad, almost separate from base of proximal end of the penis base, visible both dorsally and ventrally; Dg2 slightly curved, situated vertically, crossing penial lobe; Dg3 smaller positioned near inner base of lobe; several smaller glands run alongside Dg2. Terminal gland very large, appearing to cover terminal portion of penial lobe, visible on both dorsal and ventral sides of lobe. Ventral gland wide and prominent, near base of lobe. Filament with line of moderately dark internal pigment, appearing ventrally, and pigmental patches scattered on ventral surface. Capsule and albumen gland similar in size, albumen gland tapered, elongate. Most (2/3) of albumen gland ahead of pallial wall. Capsule gland divided into two sections. Genital aperture near terminal end of capsule gland with small vestibule. Coiled oviduct a long, large loop. Seminal receptacle small elongate sac that narrows near insertion with common duct of seminal receptacle and coiled oviduct.

Pyrgulopsis harrymilleri Perez, sp. nov. Figs. 5A–F, 6A–F, 7A–D. urn:lsid:zoobank.org:act: E2CA0B7D-0283-4C6F-8F42-3BE8BB36FF1B Diagnosis. A globose to low-conical snail with strong growth lines on teleoconch. Aperture ovate and usually slightly separated from body whorl. Mantle tissue pigmented gray-black. Sharply pointed median cusp of central radular teeth. Penial gland bifurcate at proximal end. Types. Holotype ANSP A483285, paratypes ANSP A483286, all from Vasquez Springs, Presidio County, Texas. Type locality. Vasquez Spring, N of Candelaria, 30.224070 W, - 104.583990 N, Presidio County, TX Etymology. We use the specific epithet “ harrymilleri ” in honor of Dr. Harry Miller III and Harry Miller IV who provided support and access to land, which allowed discovery of this Pyrgulopsis species. Remarks. We propose the common name “Millers’ springsnail” in honor of the Millers and aligning with common naming convention for honorific names in Pyrgulopsis. Description. Shell globose to broadly low-conical, light tan in color, relatively short spire with convex to slightly shouldered whorls. Whorls with a distinct suture. Protoconch with periostracum mostly eroded in some individuals. The initial portion lightly wrinkled with a few small spiral lines. Teleoconch with strong growth lines. Aperture ovate, angled above, adnate, or usually slightly separated from the body whorl (in larger shells). Peristome complete, inner lip slightly thickened and slightly reflected in the columellar portion. Outer lip slightly prosocline, sinuate adapically. Umbilicus narrow to rimate. Average shell measurements for n= 10 adult individuals: shell height= 2.40 mm (SD=0.21), spire height= 0.46 mm (SD=0.08), body whorl height= 1.95 mm (SD=0.15), shell width= 1.78 mm (SD=0.09), body whorl width= 1.47 mm (SD=0.08), aperture height= 1.24 mm (SD=0.07), aperture width= 1.03 mm (SD=0.06), number of whorls, 4.05 (SD=0.23). Operculum(Fig.6E,F)narrowly ovate,flat,light amber;eccentric nucleus,paucispiral growth pattern.Attachment scar slightly thickened midway between nucleus and columellar edge. Attachment region callus moderately thick. Growth lines distinct, simple. Outer margin thin (nearly colorless), but with raised ridge, particularly along outer margin but with narrow thicker edge along inner margin. Central radular tooth (Fig. 7) with moderately indented dorsal edge; lateral cusps 5–8; central cusp sharply pointed, considerably broader and longer than laterals, basal cusp 1, narrowly elongate, with strong dorsal support. Basal process medium width, broadly v-shaped; basal sockets deep. Lateral tooth face tall with broad neck and outer wing; central cusp shovel-like, sometimes flexed, lateral cusps 1–3 (inner) and 1–4 (outer); outer wing broad and about 200% length of cutting edge; basal tongue well developed. Inner marginal teeth having 20–25 cusps; fifth cusp from outer edge enlarged, sometimes buttressed with wide base including adjacent teeth. Outer marginal teeth having 31–34 cusps, inner edge sometimes having a short wing, near mid-length. Tentacles pale with darkly pigmented patch at base and one solid black line running the entire length of the tentacle. Snout dark gray-black, light colored on distal edge. Foot darkly pigmented on anterior surface, and light colored beneath. Opercular lobe black along anterior edge and with sides, pale in the center. Neck pigmented along edges, pale in center. Visceral coil and pallial roof both darkly pigmented, almost black. Ctenidium occupies a portion of the pallial cavity, filaments much broader than high, with filaments elongate, roughly triangular, wider at the base, lateral surface with elongate ridge, ~15–20. Middle and posterior filaments with widely elongated base. Osphradium elongate, short, opposite posterior end to middle of ctenidium, anterior end simple. Penis large (Fig. 5D,E); filament medium length, narrow; lobe slightly shorter than filament, very broad. Penial gland covers half the length of the filament, distal to the body, with a strong bifurcation proximally. Dorsal gland (Dg) 1 short, broad, situated at proximal end of filament; Dg2 slightly curved, situated on a swelling left of Dg1; Dg3 similar in size, positioned near inner edge of lobe. Terminal gland elongate, curved, and situated entirely on ventral side of lobe. Ventral gland weakly curved and prominent, on large subterminal swelling. Filament with moderately dark internal pigment. Capsule and albumen gland similar in size. Pallial portion of the albumen gland is large. Genital aperture near terminal end of capsule gland. Coiled oviduct a large loop, partially imbedded in albumen gland, with a light scattering of pigment. Bursa copulatrix rounded. Seminal receptacle elongate to pyriform, narrows near insertion with common duct of seminal receptacle and coiled oviduct. In many individuals, a line of oval fecal pellets fills intestine, passing along lateral edge of ctenidium. No fecal pellets observed in P. rubra sp. nov. or P. metcalfi individuals. Habitat and Distribution. This species’ known range is in Vasquez Spring on the western slopes of the centralsouthern portion of the Sierra Vieja mountains (~ 7 km SW of Capote Peak). The habitat is approximately the upper 10 m of a 20 m length of narrow, high-gradient spring run, 10 to 30 cm wide and 0.5 to 3 cm deep on the upstream. The snails occupy ~50% of a tiny spring run that discharges from below a boulder on a steep hillslope at a geologic contact between volcanic talus and an underlying lens of weathered volcanic ash. Substrate consisted of gravel and small rocks, with sand and fine organics, surrounded by spike rushes. Habitat is partly fenced but fence is broken, and stream is visibly impacted by animals; likely audad (Fig. 11A). At the source, water temperature was 26.81 °C, pH = 7.73, conductivity = 1297 (µS/cm), dissolved oxygen = 10.87 mg /L. Two adjacent seeps did not have snails.

Pyrgulopsis rubra Perez, sp. nov. Figs. 8A–F, 9A–F, 10A–E. urn:lsid:zoobank.org:act: C7481249-6B82-4C95-A08F-26E5AD4ACC7E Diagnosis. Shell ovate-conic, pale brown with slightly convex whorls. Aperture ovate, usually appressed to body whorl. Mantle tissue pigmented reddish. Rounded median cusp of central radular teeth. Penial gland does not bifurcate at proximal end, terminal gland of the penis extending to ventral and dorsal aspects. Types. HolotypeANSPA483288,paratypesANSPA483289, all from PaloAmarillo Springs, Presidio County,TX. Type locality. PaloAmarillo Springs, Big Bend Ranch State Park, 29.50925N,- 104.17158W, Presidio County,TX. Etymology. We use the specific epithet “rubra” to refer to both the reddish pigmentation on the tissues of this snail and the red volcanic rock faces in the high canyon walls and surrounding landscape. Remarks. Among the 30 individuals dissected from this population, only 1 male was observed. Therefore, features of the male anatomy were observed in one individual and should be considered accordingly. Description. Shell ovate-conic, very pale brown in color, taller spire with slightly convex and shouldered whorls. Protoconch uneroded, mostly smooth with scattered raised granules. Teleoconch with strong growth lines. Aperture ovate, angled above, nearly appressed to body whorl. Peristome complete, inner lip slightly thickened. Outer lip prosocline, simple. Narrowly umbilicate. Average shell measurements for n= 10 adult individuals: shell height= 2.49 mm (SD=0.17), spire height= 0.48 mm (SD=0.08), body whorl height= 2.01 mm (SD=0.13), shell width= 1.84 mm (SD=0.10), body whorl width= 1.34 mm (SD=0.15), aperture height= 1.30 mm (SD=0.05), aperture width= 1.06 mm (SD=0.09), number of whorls=4.20 (SD=0.16). Operculum (Fig. 9E, F) narrowly ovate, flat, light amber, nucleus eccentric and paucispiral. Attachment scar thickened midway to columellar edge. Attachment region callus concentrated near nucleus but with usual extent for the group. Growth lines distinct, but slight, simple. Wide band of thinned outer edges especially along outer margin, without a hint of a ridge, not elevated. Central radular tooth (Fig. 9) with moderately indented dorsal edge; lateral cusps 5; central cusp rounded, considerably broader, and longer than laterals, basal cusp 1, widely elongate, slightly curved, with moderate, wide dorsal support. Basal process medium width, narrowly v-shaped; basal sockets deep. Lateral tooth face wide; Central cusp narrowly shovel-like, lateral cusps bluntly pointed, 3–4 (inner) and 3–4 (outer); outer wing broad and about same length as cutting edge. Inner marginal teeth having 14–16 cusps which enlarge rapidly beginning with the 5th tooth from the outer edge. Outer marginal teeth having ~13-small, terminal cusps, outermost tooth usually shorter and wider than others, inner edge of tooth margin sometimes with a short wing, near mid-length. Cephalic tentacles tapered,rounded, nonexpanded, weakly developed eye lobes.Tentacles pale, except for weakly pigmented patch just distal to eye. Some individuals have patch extended into two lines of weak pigment running proximal to eyespot and ending halfway to distal end of tentacle. Snout with dark reddish-brown pigmentation, light colored on distal end. Foot pigmented reddish along anterior portion. Opercular lobe broadly pigmented along anterior portion, with spotty pigment throughout. Neck pale. Pallial roof uniformly dark reddish-brown. Mantle edge of pallial roof without coloration. Visceral coil consistent red pigment, with regular dark reddish-brown patches. Ctenidium occupies nearly the entire length of pallial cavity, filaments broader than high, with filaments elongate, roughly triangular in shape, wider at the base, slightly rounded at tip, ~15. Osphradium broadly ovate, short, opposite near middle of ctenidium, anterior end simple. Penis large (Fig. 8D,E); filament medium length, narrow; lobe half the length of filament, broad. Penial gland covering most of filament, not noticeably bifurcate. Dg1 short, slightly oblique, stretching from just behind penial gland to near base of penis; Dg2 positioned along far left edge, slightly curved proximally toward Dg1; Dg3 very short, positioned at conjunction of lobe and filament. Terminal gland long, transverse, reaching from midpoint of dorsal lobe across outer edge and to near midpoint of ventral lobe. Ventral gland short, vertically aligned. Filament with moderately dark internal pigment. Capsule gland similar in size to albumen gland. Coiled oviduct a large loop, as large as albumen gland and covering it, with a darkly pigmented stripe. Bursa copulatrix rounded at posterior end, same width as albumen gland, posterior to gland and nearly the same size. Seminal receptacle elongate, slightly curved, larger at the end and narrow near insertion with common duct of seminal receptacle and coiled oviduct, extends nearly to end of albumen gland. Habitat and Distribution. Palo Amarillo Spring, (89 km SSE of Vasquez Spring) is in a canyon on the SW side of the Big Bend Ranch State Park. The Palo Amarillo Spring population (~ 10 m 2) is found in high concentrations on or just below a vertical igneous bedrock face where a single small spring discharges from a horizontal fracture and flows into a small spring-fed creek in a canyon. Habitat is in a sheet of water flowing down a 2–3 m vertical rock face and through vegetation into an adjacent pool in the canyon. In lower densities on cattail, algae, and on fallen Opuntia sp. pads where spring flows into main pool (Fig. 11B). Water temperature was 22.45 °C, pH = 7.85, conductivity = 894.1 (µS/cm), dissolved oxygen = 7.83 mg /L.

Subgenus Neoxyloma Pilsbry, 1948 Oxyloma (Neoxyloma) Pilsbry, 1948: 775, 777. Type species Succinea effusa L. Pfeiffer, 1853 by original designation. Remarks The degree of coiling and folding of the epiphallus was used by Pilsbry (1948) as the sole anatomical character to separate Neoxyloma from Oxyloma s. str. The validity of this classification schema has yet to be properly evaluated with molecular phylogenetic and anatomical analyses., Published as part of Perez, Kathryn E., Martinez Cruz, Marco A., Steury, Brent W. & Barker, Gary M., 2021, A fresh start in ambersnail (Gastropoda: Succineidae) taxonomy: finding a foothold using a widespread species of Oxyloma, pp. 102-126 in European Journal of Taxonomy 757 on page 112, DOI: 10.5852/ejt.2021.757.1419, http://zenodo.org/record/5076100, {"references":["Pilsbry H. A. 1948. Land Mollusca of North America (North of Mexico). Vol. 2, Part 2. Academy of Natural Sciences of Philadelphia, Philadelphia, PA.","Pfeiffer L. 1853. Monographia Heliceorum Viventium. Sistens Descriptiones Systematicas et Criticas omnium huius Familiae Generum et Specierum hodie Cognitarum, Vol. 3. F. A. Brockhaus, Leipzig. https: // doi. org / 10.5962 / bhl. title. 10791"]}

Genus Oxyloma Westerlund, 1885 Succinea (Oxyloma) Westerlund, 1885: 1, 7. Type species Succinea dunkeri L. Pfeiffer, 1865, subsequent designated by Westerlund (1885). Gender: Neuter. Remarks Readily distinguished anatomically from other succineid genera by presence of a digiform appendix opening to the proximal apex of the penis., Published as part of Perez, Kathryn E., Martinez Cruz, Marco A., Steury, Brent W. & Barker, Gary M., 2021, A fresh start in ambersnail (Gastropoda: Succineidae) taxonomy: finding a foothold using a widespread species of Oxyloma, pp. 102-126 in European Journal of Taxonomy 757 on pages 111-112, DOI: 10.5852/ejt.2021.757.1419, http://zenodo.org/record/5076100, {"references":["Westerlund C. A. 1885. Fauna der in der Palaarctischen Region (Europa, Kaukasien, Sibirien, Turan, Persien, Kurdistan, Armenien, Mesopotamien, Kleinasien, Syrien, Arabien, Egypten, Tripolis, Tunesien, Algerien und Marocco) lebenden Binnenconchylien. Vol. V. Fam. Succineidae, Auriculidae, Limnaeidae, Cyclostomidae und Hydrocenidae. H. Ohlsson, Lund."]}