A revision of the Annulariidae of Central America (Gastropoda: Littorinoidea) Author Watters, G. Thomas text Zootaxa 2014 3878 4 301 350 journal article 42318 10.11646/zootaxa.3878.4.1 e778cf07-b718-454a-98d1-1a9edbe071aa 1175-5326 252701 B6AF841A-2D56-4F76-847F-44E881DF38B5 Halotudora kuesteri (Pfeiffer, 1852) Figures 3 P–Z, 12 D, 13 E, F, I Type material. Cistula küsteri Pfeiffer, 1852 : JDC, figured type (but not in MNHN database); NHMUK unnumbered (3), specimens from Dyson bearing Pfeiffer’s handwriting may be syntypes . Cistula sargi Crosse & Fischer, 1883 : Chevallier (1965) gave the type as in the JDC; MNHN-IM-2000-5579 (15), paratypes , one of which may be the JDC specimen. Type locality. Cistula küsteri Pfeiffer, 1852 : “ Honduras .” Restricted by Thompson (2011) to Belize . Cistula sargi Crosse & Fischer, 1883 : “in regione Coban dicte, Guatemalae.” Type figured. Cistula küsteri Pfeiffer, 1852 : Pfeiffer, 1854a : pl. 41, figs. 9, 10. Cistula sargi Crosse & Fischer, 1883 : Fischer & Crosse, 1890 : pl. 42, figs. 4, 4a– 4i , fide Chevallier, 1965 : 29. Cresonymy. “ Cyclostoma radiosum Morelet, 1849 ” Petit de la Saussaye, 1850 : 45 ; Pfeiffer, 1854a : 275 –276, pl. 37, figs. 15, 16 [all misidentifications]. “ Cistula radiosa ( Morelet, 1849 ) ” Pfeiffer, 1851 : 169 ; Pfeiffer, 1852a : 263 ; Pfeiffer, 1852c : 41 ; Pfeiffer, 1853a : 184 ; Adams & Adams, 1856 : 294 ; Pfeiffer, 1858 : 131 ; Pfeiffer, 1865 : 142 ; Bland, 1866 : 61 ; Pfeiffer, 1876 : 187 ; Crosse & Fischer, 1883 : 103 ; Hinkley, 1920 : 44 ; Chevallier, 1965 : 29 [all misidentifications]. Cistula küsteri Pfeiffer, 1851 : 170 [ nomen nudum ]. Cistula küsteri Pfeiffer, 1852a : 265 ; Adams & Adams, 1856 : 294 ; Pfeiffer, 1858 : 131 ; Pfeiffer, 1865 : 142 ; Bland, 1866 : 61 ; Pfeiffer, 1876 : 187 ; Chevallier, 1965 : 29 [in synonymy of Cistula sargi Crosse & Fischer, 1883 ]; Watters, 2006 : 321 . Cistula ? küsteri Pfeiffer, 1852 . Pfeiffer, 1853a : 186 . Cyclostoma küsteri (Pfeiffer, 1852) . Pfeiffer, 1853b : 249 ; Pfeiffer, 1854a : 312 , pl. 41, figs. 9, 10; Pfeiffer, 1854b : 316 . Ctenopoma trochlearis var. Tristram, 1861 : 233 ; Tristram, 1863 : 412 . Chondropoma ottonis “Pfeiffer ” Tristram, 1863 : 412 [ non Pfeiffer, 1846 ]; Martens, 1890 : 18 [in synonymy of Cyclostoma radiosum Morelet, 1849 ]; Fischer & Crosse, 1890 : 189 , 191 [in synonymy of Cyclostoma radiosum Morelet, 1849 ]. “ Chondropoma radiosum ( Morelet, 1849 ) ” Reeve, 1863b : pl. 9, fig. 69; Martens, 1890 : 14 , 18; Pilsbry, 1900 : 140 [all misidentifications]. Chondropoma kusteri (Pfeiffer, 1852) . Reeve, 1863b : pl. 11, fig. 87; Martens, 1890 : 18 . Cistula sargi Crosse & Fischer, 1883 : 103 ; Solem, 1961 : 202 –204; Thompson, 2011 : 46 [both in synonymy of Cyclostoma radiosum Morelet, 1849 ]. “ Ctenopoma radiosa ( Morelet, 1849 ) ” Fischer & Crosse, 1890 : 188 –191, 193–195, pl. 42, figs. 1, 1a, 1b [misidentification]. Ctenopoma sargi ( Crosse & Fischer, 1883 ) . Fischer & Crosse, 1890 : 188 , 191–194, 195, pl. 42, figs. 4, 4a– 4i. Ctenopoma küsteri (Pfeiffer, 1852) . Fischer & Crosse, 1890 : 188 , 194–196, pl. 42, figs. 7, 7a, 7b. Chondropoma sargi ( Crosse & Fischer, 1883 ) . Martens, 1890 : 14 , 18. “ Choanopoma radiosum ( Morelet, 1849 ) ” Goodrich & van der Schalie, 1937 : 20 , 31; Harry, 1950 : 27 ; Thompson, 1966 : 28 [all misidentifications]. “ Licina radiosa ( Morelet, 1849 ) ” Basch, 1959 : 7 [misidentification]. “ Choanopoma ( Colobostylus ) radiosum ( Morelet, 1849 ) ” Haas & Solem, 1960 : 130 , pl. 13, figs. 5–7 [misidentification]. Choanopoma ( Choanopomops ) küsteri (Pfeiffer, 1852) . Solem, 1961 : 195 , 203–204. “ Choanopoma ( Choanopomops ) radiosum ( Morelet, 1849 )” Solem, 1961 : 197 –198, 202–203, pl. 11, figs. 13, 14, pl. 12, figs. 24l, m, map 1; Thompson, 2011 : 46 –47, 278, 280, 296 [all misidentifications]. Cistula küsteri ( Pfeiffer, 1851 ) . Chevallier, 1965 : 29 [in synonymy of Cistula sargi Crosse & Fischer, 1883 ]. Cistula grateloupi (Pfeiffer, 1852) . Chevallier, 1965 : 29 [in synonymy of Cistula sargi Crosse & Fischer, 1883 ]. Halotudora kuesteri (Pfeiffer, 1852) . Watters, 2006 : 73 , 321. “ Halotudora radiosa ( Morelet, 1849 ) ” Watters, 2006 : 73 , 430 [misidentification]. Chondropoma ( Chondropomium ) kuesteri (Pfeiffer, 1852) . Thompson, 2011 : 44 , 278. Distribution and habitat. Limestone outcrops and caves in the Maya Mountains and Sierra de los Cuchumatanes throughout most of Belize and portions of the Petén and Alta Verapaz departments in Guatemala , from 50–600 m altitude, more common at lower levels. “Found on limestone and dead wood, sometimes hanging by a thread. Some were in motion, but the larger part were attached to the rock or wood of similar color” ( Hinkley, 1920: 44 ). Commonly found among Mayan ruins. Locally abundant. There are historical records from several sites in Alta Verapaz Department, Guatemala , including the type locality of Cistula sargi Crosse & Fischer, 1883 . Solem (1961) questioned the validity of these records but collections in 2014 confirm its presence there. See “Remarks” below. Conservation. A significant portion of the range is contained in various protected areas (e.g., Tikal National Park, Mountain Pine Ridge Forest Reserve, Columbia Forest Reserve). Other material (specimens examined: 1,185). Belize . Belize District: UF 207513 (63), limestone hill on S side of Sibun River at Cedar Bank, 50 m ; UF 207564 (42), limestone hill on S shore of Soldier Creek, 4.0 km W of Gales Point. Orange Walk District: UF 41491 (34), 268517 (79), Mayan ruins 8.4 km NE of Gallon Jug; UF 207878 (14), limestone hills 3.2 km E of Blue Village, 75 m . Stann Creek District: UF 207591 (90), limestone hill 2.2 km S of Trio Village, 50 m ; UF 207916 (21), limestone hill 4.5 km S of Trio Village, 100 m . Toledo District: GTW 13724a (1), Grego coll. (8), Lubaantún ruins; UF 207686 (67), 1.0 km W of San Felipe, 75 m ; UF 207647 (1), limestone hill 2.0 km S of Pueblo Viejo, 290 m ; UF 207732 (44), limestone hill 3.0 km SE of Pueblo Viejo; UF 207746 (27), limestone hill 2.0 km N of Blue Creek, 75 m ; UF 207637 (57), UF 207775 (55), UF 207837 (63), limestone hill 1.0 km S of Aguacate, 100 m ; UF 268087 (6), Camp 2, Quartz Ridge; UF 371984 (24), summit of entrance to cave, Cerrito; UF 370064 (30), entrance to cave, Cerrito. Cayo District: GTW 13724b (2), Grego coll. (6), Mountain Pine Ridge Forest Reserve; Grego coll. (9), Rio Frío Cave; UF 207482 (160), limestone cliff 1.0 km N of Black Rock on E side of East Branch Macal River, 120 m ; UF 216383 (15), 1.6 km N of Benque Viejo Del Carmen; UF 207337 (58), limestone hill 14.0 km SSE of Benque Viejo Del Carmen, 300 m ; UF 207455 (55), limestone hill 1.0 km NE of Arenal, 150 m ; UF 207393 (4), Caracol ruins, 600 m ; UF 207423 (17), limestone hill 12.5 km NE of Caracol, 23 km S of Augustine, 510 m ; UF 268056 (2), Los Lirios Camp hill top on road to Grano de Oro; UF 20023 (9), rock pit near river at San Jose Succotz; UF 146234 (4), slope behind San Ignacio Hotel, 110 m ; UF 135069 (10), cow pasture 4.2 km W of San Ignacio; UF 146246 (12), 7.9 km SE of San Ignacio on Cristo Rey Rd., 110 m ; UF 146357 (1), Deep Hole 13.5 km SE of Belmopan; UF 135107 (21), vicinity of Herman’s Cave and Blue Hole National Park; GTW 13724k (1), Cahal Pech ruins. Guatemala . Izabal Department: GTW 13724d (2), Livingston; UF 119156 (2), UF 217908 (4), near Livingston; UF 119159 (3), mouth of Rio Cavech, Livingston. Petén Department: UF 155811 (1), Poptún; GTW 13724i (5), entrance to Poptún; UF 155791 (8), on Modesto Mendez Fores Road 7.5 km S of Poptún; UF 109157 (27), 213710 (9), limestone hill E of road to Santa Ana, 2 km S of Flores ; UF 155803 (6), Tikal; GTW 13724f (5), km 375 on road from Rio Dulce to Flores ; GTW 13724 g (6), km 317, road from Rio Dulce to Flores ; GTW 13724j (9), km 333, road from Rio Dulce to Flores ; GTW 13724h (7), road 2 km before Quebrada Seca; GTW 13724e (2), entrance to small cave, Barra Lampara. Description. Shell conical, moderately high-spired, compact, solid. Smallest adult specimen seen 10.7 mm in length, largest 21.2 mm , average 15.0 mm (decollate). Protoconch lost as adult, 1.5 prominent, rounded, pale whorls. Teleoconch of 3.5–4 rounded whorls. Umbilicus nearly occluded by outer lip. Spiral sculpture of ca. 30 low cords separated by their own width, slightly stronger in the umbilicus. Axial sculpture of numerous, thin threads that render the spiral cords minutely scalloped or serrate. Suture strongly indented, narrowly channeled. Tufts absent but the axial lamellae may render the suture serrate. Aperture nearly circular. Inner lip smooth, narrowly exserted. Outer lip strongly lamellate, widely reflected perpendicular to whorl, evenly expanded except much narrower facing umbilicus, auriculate posteriorly, widely adnate to previous whorl. In some specimens the outer lip may be concave, particularly the posterior auricle and outer border. Base color dingy white to tan, first teleoconch whorl reddish-orange in some specimens. Unicolor or with very faint spiral bands, bands most conspicuous on both sides of outer lip as radiating brown rays. Operculum paucispiral, with a thick reflected plate composed of numerous recurved lamellae; in un-eroded specimens the surface of the plate is smooth, eroded specimens reveal the underlying lamellae. Radula and anatomy unknown. Animal uniformly pale cream or tan, or sides darker than head (fig. 13 E); middle portion of antennae darker brown, tips abruptly paler; area between bases of antennae marked with a large, dark “V” which partially encircles the bases; between this marking is a smaller, inverted dark “V.” These markings may be much paler and more diffuse in some populations or reduced to a series of longitudinal stripes. Eyes black. Variation in specimens. Specimens vary in the shape of the spire, which is more inflated in some specimens than others. Sculpture varies from finely beaded to serrate. Adult size is quite variable, from 10 mm to over 21 mm . Comparison with other species. This species most closely resembles H. gruneri but that species is nearly smooth and translucent; H. kuesteri has a fine but obvious cancellate or minutely serrate sculpture and is much more solid. Halotudora gaigei has a higher spire (the protoconch is usually retained), smoother sculpture, and a less expanded outer lip that is usually solute. Choanopomops largillierti is immediately recognized as the only Central American species having sutural tufts. All of these species may found in Belize . Remarks. This species has been almost uniformly referred to as radiosa . However, Morelet’s (1849) description of Cyclostoma radiosum being “translucent,” with “weak striae,” “brown banded below the periphery,” and the size of the specimen leaves little doubt that the shell he had before him was the species known as gruneri rather than the radiosa of authors. Morelet’s types are often at NHMUK. NHMUK 93.2.4.686 consists of three specimens labeled Cyclostoma radiosum . One specimen, loose from the board, is marked with an “X.” This is a specimen of radiosa of authors and might be construed as the type . But in addition to the discrepances in shell form mentioned above, Morelet gave the length of his shell as 20 mm—the marked NHMUK specimen is only 13 mm in length. The locality information also does not match: although described from Petén Department, the accompanying label to NHMUK 93.2.4.686 reads “Vera Paz” (Alta Verapaz Department). It appears that the NHMUK specimens are not Morelet’s original lot but were added later. They do not represent the same species. Based on Morelet’s description, radiosa is a junior synonym of Cyclostoma gruneri Pfeiffer, 1846 . The radiosa of authors defaults to the next available name— Cistula küsteri Pfeiffer, 1852 . Cistula küsteri Pfeiffer, 1852 , is clearly recognizable from its description and three unnumbered specimens at NHMUK from Dyson bearing Pfeiffer’s handwriting are likely syntypes . In the chresonymy above, I have assumed that subsequent authors followed Pfeiffer’s 1854 depiction of “ radiosum ” rather than Morelet’s unillustrated description, and regard these subsequent citations as misidentifications. Cistula küsteri Pfeiffer, 1852 , actually refers to an occasionally seen sculptural form of this species. In this form the outer lip is concave and sinuous rather than flat (figs. 3 R–T); Pfeiffer described it as “ campanulatoexpansum .” This variation occurs throughout the range of the species. It does not appear to be related to sexual dimorphism. Cistula sargi Crosse & Fischer, 1883 , described from Cobán, Guatemala , was considered by Solem (1961) as a sculptural variation of “ radiosum .” It differs in having stronger sculpture and a strongly fimbriated outer lip facing the umbilicus (fig. 3 V ). It bears an uncanny resemblance to some Jamaican Annularia . The collection of additional specimens may reveal it to be distinct. Original description (translated here from Latin). Cistula küsteri Pfeiffer, 1852a . “Shell perforate, ovateturret, truncate, thin, spiral sulci and regularly granulate-reticulate crowded longitudinal ribs, almost rough, slightly shiny, diaphanous, yellowish-brown, painted with obsolete, interrupted red lines; spire convex-turret, widely truncate; suture deep, simple; remaining 4 whorls convex, last narrow, rounded; aperture nearly vertical, subcircular; peristome double: inner briefly expanded, adnate, outer expanded bell-shaped, concentric striations, forward concave, radiating red, top angled, penultimate whorl narrow.—Operculum.” 14 mm . Original description (translated here from Latin). Cistula sargi Crosse & Fischer, 1883 . “Shell distinctly perforate, oblong-turret, truncate, elevated transverse lirae and longitudinal lines, very crowded elegantly pricklydecussate, not shining, rough, thin, diaphanous, pale brown, interrupted lines, red bands; spire quite elevated, apex wanting; suture simple, impressed; remaining four whorls convex, farthest penultimate scarely produced; aperture vertical, nearly circular, suboval, interior pale brown; peristome double, inner continuous, scarely thickened, thin, pale brownish-white, outer wide and squarely expanded, reflexed, cut off by penultimate whorl; fringe of outer lip with concentric subimbricated striations and beautiful red rays.—Nucleus of operculum slightly eccentric, somewhat concave, outer margin of whorl free and outside of last whorl with oblique striations, especially strongly impressed.” 10 mm width. Etymology. Cistula küsteri Pfeiffer, 1852 : Heinrich Carl Küster ( 1807–1876 ), German conchologist. Cistula sargi Crosse & Fischer, 1883 : F. Sarg, collector of the types ; probably Franz Sarg, owner of the Finca Pancorral coffee plantation at San Cristóbal , Guatemala .