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 Parachondria rubicundus ( Morelet, 1849 ) Figure 10 A– O , 12 A , 13 C, J Type material. Cyclostoma rubicundum Morelet, 1849 : NHMUK 93.2.4.689-91, holotype (fig. 10 A); NHMUK same (2), paratypes ; ANSP 14102 (3), paratypes (as syntypes in Richardson et al. , 1991 ); MNHN unnumbered (12), paratypes [ Chevallier, 1965, cites eight specimens ]; SMF unnumbered (1), paratype . Cyclostoma acerbulum Morelet, 1851 : NHMUK 1893.2.4.706–7 (2), probable syntypes (fig. 10 F); ZMB 65687 (1), possible syntype . Cistula subangulata Martens, 1886 : not located. Type locality. Cyclostoma rubicundum Morelet, 1849 : “prov. Peten et Vera-Paz, arborum super folis” [ Guatemala ]. Cyclostoma acerbulum Morelet, 1851 : “non vulgaris pr. Vera-Paz” [ Guatemala ]. Cistula subangulata Martens, 1886 : “Teleman, Guatemala .” Type figured. Cyclostoma rubicundum Morelet, 1849 : possibly Pfeiffer, 1854a : pl. 37, figs. 29, 30. Cyclostoma acerbulum Morelet, 1851 : unfigured. Cistula subangulata Martens, 1886 : unfigured. Cresonymy. Cyclostoma rubicundum Morelet, 1849 : 22 ; Petit de la Saussaye, 1850 : 46 ; Pfeiffer, 1854a : 280 , pl. 37, figs. 29, 30; Richardson et al. , 1991 : 53 ; Watters, 2006 : 448 –449. Cyclostoma acerbulum Morelet, 1851 : 19 ; Pfeiffer, 1852a : 420 ; Pfeiffer, 1852b : 154 ; Solem, 1961 : 205 –207, pl. 11, fig. 19 [as form or synonym of Cyclostoma rubicundum Morelet, 1849 ]; Watters, 2006 : 128 ; Thompson, 2011 : 44 [in synonymy of Cyclostoma rubicundum Morelet, 1849 ]. Chondropoma rubicundum ( Morelet, 1849 ) . Pfeiffer, 1851 : 173 ; Pfeiffer, 1852a : 291 –292; Pfeiffer, 1852c : 45 ; Adams & Adams, 1856 : 295 ; Tristram, 1861 : 232 ; Reeve, 1863a : pl. 6, fig. 47; Pfeiffer, 1865 : 154 ; Bland, 1866 : 61 ; Pfeiffer, 1876 : 196 ; Martens, 1890 : 12 , 16, 17; Fischer & Crosse, 1890 : 203 , 205–207, 209–210, pl. 41, fig. 5; Hinkley, 1920 : 42 , 44, 52; Goodrich & van der Schalie, 1937 : 12 , 15, 20, 31; Bequaert, 1957 : 212 ; Chevallier, 1965 : 29 . Cistula ? acerbula ( Morelet, 1851 ). Pfeiffer, 1858 : 133 ; Bland, 1866 : 61 ; Pfeiffer, 1876 : 188 . ? Chondropoma trochleare (Pfeiffer, 1852) . Reeve, 1863b : pl. 11, fig. 82 [misidentification]. Cistula subangulata Martens, 1886 : 162 ; Solem, 1961 : 205 –206, pl. 11, fig. 20 [as form or synonym of Cyclostoma rubicundum Morelet, 1849 ]. Chondropoma acerbulum ( Morelet, 1851 ) . Martens, 1890 : 15 , 17; Fischer & Crosse, 1890 : 204 , 210–211, pl. 41, figs. 4, 4a–d. Chondropoma subangulatum ( Martens, 1886 ) . Martens, 1890 : 15 , 17, pl. 1, fig. 5; Watters, 2006 : 493 [in synonymy of Cyclostoma rubicundum Morelet, 1849 ]. Ctenopoma acerbulum ( Morelet, 1851 ) . Fischer & Crosse, 1890 : 188 . Ctenopoma subangulata ( Martens, 1886 ) . Fischer & Crosse, 1890 : 188 . Chondropoma ( Chondropomium ) rubicundum ( Morelet, 1849 ) . Henderson & Bartsch, 1921 : 60 ; Baker , 1928 : 50 ; Thompson, 2011 : 44 , 278, 280, 282, 285, 293. Choanopoma ( Choanopomops ) rubicundum ( Morelet, 1849 ) . Solem, 1961 : 204 –207, pl. 11, figs. 18–20, pl. 12, figs. 24p, q, map 2. Parachondrops ? acerbulus ( Morelet, 1851 ). Watters, 2006 : 48 , 128. Gouldipoma rubicundum ( Morelet, 1849 ) . Watters, 2006 : 72 , 448–449. Distribution and habitat. The Maya Mountains from southern Belize into northwestern Honduras , northern Guatemala , and northern Chiapas State of México , usually from 75–1,200 m altitude. Morelet (1849) described the species as arboreal. It is more commonly found in limestone outcrops; often associated with Mayan ruins (Palenque, etc.). “Situated above fallen trees, base of rocks and under old banana leaves” ( Hinkley, 1920: 44 ). Often locally abundant. Conservation. Portions of the distribution are contained in forest reserves, such as the Columbia Forest Reserve. Other material (specimens examined: 397). México . Chiapas State: GTW 13723a (1), Grego coll. (3), UF 19146 (10), Palenque ruins; Grego coll. (1), Agua Chiara (locality not found). Guatemala . UF 119154 (5). Izabal Department: UF 216798 (1), near Livingston; GTW 13723d (3), entrance to small cave, Barra Lampara; GTW 13723b (3), UF 119158 (5), mountains at Cavech River, Livingston; UF 19147 (3), UF 35635 (2), rain forest at Puerto Matías de Gálvez; UF 190585 (2), Rio Escobas at Las Escobas, 300 m ; UF 190576 (3), 7 km WSW of Las Escobas, 750 m ; UF 190560 (2), Cerro San Gil 8.5 km WSW of Las Escobas, 910 m . Alta Verapaz Department: UF 216800 (1); UF 216799 (1), Cobán; GTW 13723e (1), road leaving Lanquín; UF 190107 (6), 6.5 km SE of Lanquín, 400 m ; UF 190044 (43), 9.0 km W of Lanquín, 690 m ; UF 190118 (22), 8.0 km SE of Lanquín, 690 m ; UF 190035 (3), 9.0 km W of Lanquín, 690 m ; UF 190143 (2), 11.0 km W of Lanquín, 1,000 m ; GTW 14640a (4), Sierra de Chama; UF 189974 (3), 11.0 km W of San Cristóbal Verapaz, 1,120 m ; UF 190005 (2), 2.0 km ESE of Cojaj, 1,250 m ; UF 189987 (12), 2.5 road km from Chixoy Bridge, 810 m ; UF 189906 (5), 15.0 road km N of Cobán, 1,050 m ; GTW 13723c (15), on the way to Candelarea Cave; GTW 13723f (21), B'onb'il Pek Cave area; GTW 13723H (1), Semuc Champey. Petén Department: UF 234126 (53), S of Sayaxché beyond Laguna Petexbatún; UF 155810 (1), 2.3 km N of Poptún; UF 155792 (4), on Modesto Mendez Fores Road 7.5 km S of Poptún; GTW 13723 g (1), km 333 off road from Rio Dulce to Flores . Hueheutenango Department: UF 190300 (7), UF 190381 (6), Finca Chiblac ca. 5 km W of San Rámon, 1,000 m ; UF 190360 (1), ridge above Finca Chiblac ca. 5 km W of San Rámon, 1,200 m ; UF 190254 (7), 4.5 km WSW of Finca Tres Ranchos, 1,300 m . Belize . Toledo District: UF 207633 (31), limestone hill 2.0 km S of Pueblo Viejo, 290 m ; UF 207732 (1), UF 207737 (2), limestone hill 3.0 km SE of Pueblo Viejo; UF 268054 (2), Jimmy Cut, Columbia River Forest Reserve; UF 268068 (8), Columbia River Forest Reserve; UF 268072 (8), hill W of quarry on road to Columbia River Forest Reserve; UF 268084 (6), Camp 2, Quartz Ridge; UF 268082 (3), Union Camp, Quartz Ridge; UF 207745 (2), limestone ridge 2.0 km N of Blue Creek, 75 m ; UF 207886 (5), limestone hill 3.2 km E of Blue Creek, 75 m ; UF 207687 (9), limestone hill 1.0 km W of San Felipe, 75 m ; UF 207781 (5), UF 207838 (7), limestone ridge 1.0 km S of Aguacate, 100 m ; Cayo District: UF 207461 (1), hill 1.0 km NE of Arenal; UF 207403 (3), Caracol ruins, 600 m ; UF 146304 (1), Xunantunich ruins, 110 m ; UF 207430 (2), limestone hill 12.5 km NE of Caracol, 510 m . Honduras . Santa Barbára Department: UF 213213 (40), Chiquila. FIGURE 10 A–O. Parachondria rubicundus (Morelet, 1849) . A. Cyclostoma rubicundum Morelet, 1849 : NHMUK 93.2.4.689-91, holotype, 15 mm. B. UF 207737, 13.8 mm. C. UF 216798, 12.8 mm. D. GTW 13723b, 11.7 mm. E. GTW 13723b, 14.0 mm. F. Cyclostoma acerbulum Morelet, 1851 : NHMUK 1893-2-4.706, possible syntype, ca. 9 mm, photo courtesy J. Ablett (NHMUK). G. UF 2077430, 9.3 mm. H. GTW 14640a, 8.9 mm. I. UF 190107, 10.4 mm. J. UF 190254, 11.4 mm. K. GTW 13723a, 8.7 mm. L. UF 19146, 10.3 mm. M. UF 207687, 12.9 mm. N–O. UF 207745, 11.1 mm. Description. Shell conical, moderately high-spired, rather thin. Smallest adult specimen seen 9.3 mm in length, largest 16.8 mm (exceptional), average 10.8 mm (non-decollate). Protoconch retained, 1.5 large, rounded, smooth whorls, dark brown or banded, earliest portion paler, clearly demarcated from teleoconch. Teleoconch of 4.5–5 rounded whorls. Umbilicus open, wide. Spiral sculpture absent except for 1–3 weak cords in the umbilicus. Axial sculpture of erect, regularly spaced, thin, blade-like lamellae, spacing varying from close-set to widely separated. Suture strongly indented. Tufts absent but the axial lamellae may render the suture serrate. Aperture nearly circular. Inner lip smooth, narrowly exserted. Outer lip lamellate, reflected perpendicular to whorl, more or less evenly expanded, rather narrow, somewhat narrower facing umbilicus, slightly auriculate posteriorly, adnate to previous whorl. Base color tan, ruby red, or pale orange, spire may be darker than final whorls. Unicolor, spotted, or banded, markings darker, bands rarely continue over adapertural face of peristome, pattern visible inside aperture. Lip white, orange, or red. Operculum paucispiral, flat, with a thin calcareous deposit. Radula and anatomy unknown. Animal pale yellowish tan, area behind head pink or orange (fig. 13 C); antennae pink or orange, tips abruptly pale tan; area between bases of antennae pale purplish brown, diffuse. Eyes black. Variation in specimens. Strength of and distance between axial lamellae vary. Color varies considerably, from unicolor to spotted (color form acerbulum ). The degree of expansion of the outer lip also varies but is usually narrow. Specimens from Belize are smaller than specimens from elsewhere. Comparison with other species. See under P. cordovanus . See also under Gouldipoma terecostatum , which differs primarily in opercular features. Remarks: See under P. cordovanus . The color form acerbulum (figs. 10 F–L) is most common in the southwestern portion of the range but it also occurs sporadically across the entire range of the species. It occurs in both pure and intergrading populations. It is not limited to any particular altitude as suspected by Solem (1961) . In Belize in the extreme eastern end of its range specimens may be shinier and more ruby colored than elsewhere (figs. 10 N– O ) but nevertheless intergrade into more typical examples. Original description (translated here from Latin). Cyclostoma rubicundum Morelet, 1849 . “Shell ovatepyramid, umbilicate, translucent, scarlet-red, towards the apex dark violet; six convex whorls, first ones with plicate ribs, last with delicate striations, base prominently striated, open umbilicus with three grooves; aperture round-oval; peristome double, inner narrow, exserted; outer narrowed, penultimate whorl emarginate.” 15 mm . Original description (translated here from Latin). Cyclostoma acerbulum Morelet, 1851 . “Shell umbilicate, ovate-conic, entire, distinctly costate, ribs on the last whorl distant and barely elevated, waxy horn-colored, banded with pale red interrupted zones; 6–7 convex whorls; aperture oblong; peristome double, inner lip white, shining; outer lamellate, a little expanded, upwards slightly produced.” 10 mm . Original description (translated here from Latin). Cistula subangulata Martens, 1886 . “Shell perforate, ovate-conical, all over thin, sculptured with crowded, vertical, delicate, costae, yellowish-red, with interrupted dark bands, on the penultimate whorl with five, on the last with six, marked; whorls 6.5, prior two smooth, first pale, depressed, second dark violet, following sculptured, middle slightly angulated, last rotund, base with some raised spirals. Aperture subvertical, ovate, top slightly angulate; peristome double, red, spotted, outer slightly expanded, top of penultimate whorl adnate.” 12 mm . Etymology. Cyclostoma rubicundum Morelet, 1849 : L. rubicundus , red. Cyclostoma acerbulum Morelet, 1851 : L. diminutive of acerbus , rough—a little rough. Cistula subangulata Martens, 1886 : L. sub -, somewhat + angulatus , angled—somewhat angled. Incertae sedis