Plekocheilus (Eurytus) coloratus ( Nyst, 1845 )

Borrero, Francisco J. & Breure, Abraham S. H., 2011, The Amphibulimidae (Mollusca: Gastropoda: Orthalicoidea) from Colombia and adjacent areas, Zootaxa 3054, pp. 1-59 : 32-37

Plekocheilus (Eurytus) coloratus ( Nyst, 1845)

( Figs 11 A–F, 12B)

Under this heading we are grouping all records available to us of a widely distributed but poorly understood taxon. Several varieties or subspecies have been recognized, and museum labels abound with subspecific names, including: “ P. coloratus coloratus ,” “ P. coloratus ampullaroides ,” “ P. coloratus lamarckianus ” and “ P. coloratus subplicatus .” These denominations are confusing as they mix elements of what may be three distinct species [i.e., P. ampullaroides ( Mousson, 1873) , P. coloratus ( Nyst 1845) and P. lamarckianus ( Pfeiffer, 1848a) ]. In addition, unpublished subspecific names are also found among these labels [i.e., “ sublamarckianus,” “ subplicatus ”]. As discussed below under Remarks, it is possible that at least one more recognized specific level taxon may be part of what has been referred to as “ Plekocheilus coloratus ” in museum labels. The resolution of the phylogeography of this group is a pressing need, but of a complexity beyond the scope of the present paper and needing better quality material than currently available to us. For these reasons, we treat here the forms coloratus , subplicatus and lamarckianus , as a single taxon, P. (E.) coloratus ( Nyst, 1845) . However, for now we choose to consider P. ampullaroides ( Mousson, 1873) as a separate species because of its distinct shape, but we are aware that further studies may lead to a different conclusion.

Bulimus coloratus Nyst 1845: 228 , pl. fig. 2. Type locality: “la province de Cumana, dans la Colombie ” [ Venezuela, Edo. Sucre]. Lectotype RBINS/MT2345 ( Breure 2011). Plekocheilus coloratus (Nyst) ; Richardson 1995: 307 (references, synonymy).

Bulimus lamarckianus Pfeiffer 1848a: 229 . Type locality: “Andes of New Granada, 8000 feet.” Lectotype BMNH 1975295 ( Breure 1979).

Bulimus coloratus var. ß Pfeiffer 1855? (1854–1860): 29, pl. 8 figs 4–5.

Bulimus (Dryptus) coloratus var. subplicata Pfeiffer 1881 [ Pfeiffer & Clessin 1879 –1881]: 226. Type locality: Prov. Ocana. Type material: not located. Plekocheilus coloratus var. subplicata (Pfeiffer) ; Pilsbry 1895 [ 1895 –1896]: 198–199; Richardson 1995: 307 (references, synonymy).

Plekocheilus coloratus ampullaroides ( Mousson, 1873) ; Breure 1976: 1, pl. 2 f. 8.

Material. Antioquia, Frontino: CMC C11047 (ex Fulton), DMNH 141721, UMMZ 147710 (ex Rolle, as P. coloratus sublamarckianus Mousson ), ZMH 7478, ZMUZ 511585 (as P. coloratus ampullaroides , not seen), ZMUZ 511593 (not seen); La Bodega, between Sonsón and Nariño, 6000 ft [~ 1800 m], Carriker leg./1951: USNM 598490 (as P. coloratus ampullaroides ); Porce, Bosque Normandía, [300–700 m], J.M. Daza leg./2001: UAM; Cauca, Isla de Gorgona, north portion, humid forest [island maximum elevation is 338 m], Medem leg./1961: FMNH 114095; Cesar, Sierra de Perijá, 25–30 km E of Becerril, Finca América, 1100 m, Malkin leg./1969: FMNH 163690; El Roncón, foothills of Sierra de Perijá, 10–12 km E of Becerril, 200–300 m, Malkin leg./1969: FMNH 163691; San Sebastián de Rábago [= Nabusimake]: FMNH 157551 (2000 m, Malkin leg./1968), CMC C14234, CMC C14235 (1950 m, Borrero & T.A. Pearce leg./2010), CMC C14249 (nearby hillside forests, 1943–1970 m, Borrero & Pearce leg./2010); Socorpa Mission, forest only, Becerril, Sierra de Perijá, Malkin leg./1968: FMNH 159206, FMNH 159208, FMNH 159209, FMNH 159210, FMNH 159211, FMNH 172861, FMNH 173013; Rio Guatapurí, Sierra Nevada de Santa Marta: USNM 599524 [partim] (1200 ft [~ 360 m], Carriker leg./1946), USNM 599529 (8000–9000 ft [~ 2400–2700 m]); San José [most likely San José de Oriente], Sierra Nevada de Santa Marta: USNM 488805 (5000–6000 ft [~ 1500–1800 m], Carriker leg./1945), USNM 59301; Sierra de Perijá, Airoca: USNM 488785 (4500 ft [~ 1350 m], Carriker leg./1942), USNM 488783 (4000–6000 ft [~ 1200–1800 m]); El Africa, Sierra de Perijá, 6500 ft [~ 1950 m], Carriker leg.: USNM 488804 (leg./1942), MCZ 183869; Pueblo Bello, canyon next to Rio Ariguaní, 1125 m, Borrero & Pearce leg./2010: CMC C14192, CMC C14250, CMC C14251); 5–6 km above Manaure [Balcón del Cesar], Magdalena [sic, Cesar], W side Perijá Range, Olsson leg.: ANSP 181847 (leg./1946, under rocks in woods), ANSP 186716 (leg./1950); Cundinamarca, Bogotá: USNM 316124, DMNH 76770 [partim], BMNH 1894.2.28.4 (Cuming Colln, partim), UMMZ 242806 (Marsh Colln, as P. coloratus ampullaroides ), UMMZ 147708 (ex Caziot & Hno. Apolinar María, as P. coloratus ampullaroides ), UMMZ 13839-13840 (Stearns Colln); La Guajira, San Juán [del Cesar], 200 m ( Götting 1978 [not seen]); below Pueblo Viejo, 300–1000 m ( Götting 1978 [not seen]); above Pueblo Viejo, 1400 m ( Götting 1978 [not seen]); Riohacha: ZMH 7477; hills of Carraipía, Haught leg./1943: USNM 599496; near Carraipía, Haught leg.: ANSP 388534 (as P. oligostylus Pils. ), USNM 543490; Sierra Nueva, Sierra Negra, Carriker leg.: FMNH 71056 (3700–5000 ft [~ 1100- 1500 m]), USNM 533874 (1100–1500 m, leg./1942); Los Gorros, trail Fonseca–Riohacha, NE spur of Sierra Nevada, 2500 ft [~ 750 m], Carriker leg./1945: USNM 488810; Potrero de Venancio, 1100 m ( Breure 1984 [not seen]); Magdalena, mountains near Santa Marta, 1219 m, H.H. Smith leg./1900: UF 109842 [not seen], UF 109843 [not seen]; Sierra Nevada de Santa Marta [= SNSM], Pittier leg.: USNM 192936a; SNSM, San Andrés, Rio Frio Basin, Pittier leg./1906: USNM 192949 (1200 m), USNM 192947 (1300 m); SNSM, western foothills, between the plains and Quebrada del Bollo, 0–300 m, Pittier leg./1906: USNM 192936; SNSM, near Rio Frio, between Quebrada del Bollo and Quebrada Rodríguez, 0–300 m, Pittier leg./1906: USNM 192940; SNSM, Rancho del Bollo, Pittier leg./1906: USNM 192946; SNSM, 50 km SE Santa Marta, just W of Rio Buritaca along trail to Ciudad Perdida, 11°02’N 73°55’W, D.K. Dorman leg./1989: UF 137920, UF 137933 [both not seen]); SNSM, 2550 m, near trail beside Rio Buritaca, 11°01’N 73°55’W: UF 137926, UF 137932 [both not seen]); SNSM, Alto Buritaca, 500 m, Linares leg./1991: ICNB 2601, ICNB 2602); SNSM, Buritaca 2000 [archaeological name given to Ciudad Perdida, a former religious site of the local natives], 1100 m, leg./1981: UVZ 81190–81199, UVZ 81206; Cerro San Lorenzo, SNSM, 7000 ft [~ 2100 m], Carriker leg./1945: FMNH 71055, USNM 599526; Finca “La Granja” Cerro San Lorenzo, SNSM, Medem et al. leg./1964: FMNH 187286, FMNH 187287; Finca La Victoria, Sierra de San Lorenzo, 996 m, Borrero & Pearce leg./2010: CMC C14242; Estación San Lorenzo, Borrero & Pearce leg./2010: CMC C14193 (trail above camp, 2329–2360 m), CMC C14248 (trail below camp, 2329– 2175 m); SNSM, between San Fernando and San Francisco, Creutzberg leg./1975: RMNH; near Minca, northern slope of Sierra, approx. 700 m, Götting leg./ 1976–1978: RMNH; Forests near Minca, and Minca lower slope of San Lorenzo: CM 106208 , CM 106209 (both H.H. Smith leg./1899), ANSP 46587 (J. Rehn & M. Hebard leg./1920); Minca, 2000 ft [~ 600 m], H.H. Smith leg.: ANSP 78243; Road Minca–Hacienda La Victoria, Sector Jaguar, 1100 m, S. Guataguira & L. Machado leg./2007: ICNB; El Libano forests, leg./1899: CM 100142 , CM 106217 ; Don Diego, Carriker leg./1914: ANSP 140086; Don Amo Estate, Sierra de Santa Marta, H.H. Smith leg./1900: CM 106222 (2000 ft [~ 600 m]), CM 106223 (3500 ft [~ 1060 m]); Onaca Estate, 2500 ft [~ 750 m], H.H. Smith leg./1900: CM 106210 , CM 106220 ; Valparaiso, H.H. Smith leg./1900: CM 106211 ; Magdalena River Valley, 2500 m ( Götting 1978 [not seen]); From Breure (1984): Cacaguelito (600 m), Jiracasaca (800 m), below La Horqueta (1300 m), Las Pantidas [= Las Partidas] (1300 m), Valparaiso (1500 m), El Líbano (2000–2325 m); El Líbano: DMNH 76787, ANSP 78244 (SE of Santa Marta, 6000 ft [~ 1800 m], H.H. Smith leg./1899); Manancanaca, 11000–12000 ft [~ 3300–3600 m], Carriker leg./1946: USNM 599523; Jiracasaca, 2800–3000 ft [~ 840–900 m], H.H. Smith leg./1898: CM 106214 , CM 106218 [not seen]; Las Partidas, 3500 ft [~ 1050 m], H.H. Smith leg./1899: CM 106215 [not seen]; near Pueblito, Sierra Nevada de Santa Marta region, 463 m, Borrero & Pearce leg./2010: CMC C14246, CMC C14247; “Magdalena:” ZMUZ 511597 [not seen]; Meta, Villavicencio: DMNH 141730 (ex Walton); Norte de Santander, “Provincia de Ocaña” [type locality of P. subplicatus Pfeiffer ]; Ocaña: ZMUZ 511596, ZMUZ 511598 [both not seen]; Mount Lucerno [= Alto de Lucero]: ZMUZ 511595 [not seen]; Buenos Aires, Carriker leg./1947: USNM 486844; Guamalito, 0–2000 ft [0–600 m], Carriker leg./1949: USNM 517804; Hacienda Villa Felisa, near Cúcuta, 1100 ft [330 m], Carriker leg./1947: USNM 590658; Petrolea, near sea level, Carriker leg.: USNM 517812; La Palmita, 6000–7500 ft [~ 1800–2250 m], Carriker leg./1916: ANSP 137828; Santander, Trail to El Salado, 1800 m, P.E. Acosta leg./1960: FMNH 114053; Molagavita, 2300 m, Acosta leg./1960: FMNH 114054; Los Santos, Creutzberg leg.: RMNH.?, Colombia, Linter leg.: UMMZ 147707; “ Columbia ” [= Colombia]: ANSP 66411, ANSP 174436, CM 42437, FMNH 31169, USNM 105227, UMMZ 41101 (ex Rolle, as P. coloratus lamarckianus ), ZMA, ZMUZ 511586 [not seen]; “ New Granada:” ZMUZ 511594, ZMUZ 544089, ZMUZ 511592 [all not seen], ANSP 66427 (ex Cuming), CMC C10721, C10722, BMNH Acc. no. 1563 (MacAndrew Colln); “Andes of New Granada:” BMNH Acc. no.1829 (ex Cuming), ( Baker 1926 [at 2438 m]); “Andes Colombia:” USNM 532632 (ex Fulton).?, Campo Alegre forest, ~ 1500 ft [~ 450 m], on a tree trunk, H.H. Smith leg./1898: CM 106219 ; above Agua Dulce, 3000 ft [~ 900 m], H.H. Smith leg./1899: CM 106216 .

Altitudinal range. The localities that can be precisely located range from 70–3600 m.

Remarks. Within this taxon, many specimens from several different nominal (sub)species are frequently found in museum collections, most commonly P. (E.) coloratus ( Nyst, 1845) , P. (E.) lamarckianus (Pfeiffer, 1848) and P. (E.) ampullaroides ( Mousson, 1873) . Breure (1978: 10) placed Bulimus lamarckianus Pfeiffer, 1848 in the synonymy of P. coloratus ( Nyst, 1845) . We are unaware of a formal synonymy between P. coloratus and P. (E.) ampullaroides . In addition, Pfeiffer (1855: 29, pl. 8, figs 4–5) described Bulimus coloratus var. ß and suggested that it may deserve the status of a separate species but preferred to conservatively maintain this form as a variety of P. coloratus . This form was later referred to by Pfeiffer and Clessin (1879 –1881: 226) as Bulimus (Dryptus) coloratus var. subplicata Pfr. Thus , specimens attributed to the variety subplicatus Pfeiffer, 1881 are also found together with the taxa mentioned above in museum collections, the labels with or without the subspecific epithet. During the course of our analysis, we have found that shells belonging to other species may also be mixed in with “ P. coloratus ,” including Plekocheilus (E.) nachiyacu Pilsbry 1939 , a poorly understood species described without a specific type locality ( Pilsbry 1939: 2). Sorting out the correct affiliation of the many specimens of “ P. coloratus ” available at museum collections has proven difficult, partly due to the lack of precise localities for some of the taxa involved, and the lack of material suitable for anatomical and molecular studies. For this reason, we have chosen to list under P. coloratus all records of the various taxa mentioned above, which often exhibit strong similarities. However, we have also found specimens that were attributed to P. coloratus , but clearly belong to other, well defined species (i.e., P. gibbonius , P. oligostylus ), and have listed those under the appropriate species names. We consider it beyond the scope of the current paper to carry out a more complete analysis of “ P. coloratus ” at this time; this will be part of our on-going research. Photographs of the various “forms of P. coloratus ” are presented in Figures 10 A-F, 13A, without indication of sub-specific or variety status. Likewise, Fig. 12B shows the distribution map for those specimens of “ P. coloratus ” associated with precise localities. As stated, we are aware that there may be more than one species involved in this map, which may be partially evident as a disjunct distribution.

Other than the type specimen of P. coloratus (Nyst) , we have not seen any specimens from northern Venezuela that seem to belong to this species. Nyst’s type locality is problematic, as it seems well outside the range of this species. Two Venezuelan records available (cf. Breure 2009) are here considered questionable and in need of additional, confirmatory material. The distribution given in Pilsbry (1895: 74) includes “near Quito, Ecuador.” This is the species of Plekocheilus with the broadest latitudinal and altitudinal distribution that we know of, and we currently question its veracity. We have not seen any material from Ecuador and this species was not listed by Breure and Borrero (2008) for that country. The record from Bogotá (DMNH 76770, partim) may not be credible as the lot consists of two specimens of two different species, and the presence of this species in Cundinamarca is under question. Campo Alegre and Agua Dulce may be localities near the Sierra Nevada de Santa Marta since most of H.H. Smith’s collections come from the Santa Marta area; however, places with those names exist in other Colombian Departments. Lots ANSP 181847 and 186716 are labeled as from Magdalena; this is in error and is probably due to the fact that two towns named Manaure exist is northern Colombia: Manaure, La Guajira (formerly part of Magdalena), and Manaure Balcón del Cesar, in Cesar; only the second locality is in proximity of the Perijá Range (on its west side) and thus corresponds to Olsson’s locality.

We have observed in part of the material (specifically from Sierra Nevada de Santa Marta) a variation in the color pattern (“yellow trait”). The axial undulating stripes are bordered by a yellowish “shadow” [cf. P. (E.) fusitorsus (Oberwimmer, 1931) from Venezuelan Guayana ( Breure 2009)]. Furthermore, some shells from La Guajira and Perijá regions tend to have a strong lip and a distinct parietal callus. These shells, all from low elevation localities where dry conditions prevail, are tentatively referred to this species.

During the revision of this paper the type material of this taxon was located in the Brussels museum; see Breure (2011).

Figures

FIGURE 10. Plekocheilus (Eurytus). A – C. P. (E.) episcopalis episcopalis (Pfeiffer, 1855), lectotype BMNH 1953.11. 30.1 (H 53.5 mm), photo P. Crabb. D – G. P. (E.) e. corticosus (Sowerby, 1895). D – F. Lectotype BMNH 1907.11. 21.110 (H 54 mm), photo P. Crabb. G. Ibidem, holotype of apolinari Pilsbry, 1935, ANSP 164566 (H 55.8 mm). H – K. P. (E.) e. auriformis (Da Costa, 1904). H – J. Holotype BMNH 1907.11. 21.112 (H 68.0 mm), photo P. Crabb. K. ANSP 174434 (H 71.0 mm). L – N. P. (E.) subglandiformis (Mousson, 1873), lectotype ZMUZ 511633 (H 40 mm). O – Q. P. (E.) plectostylus (Pfeiffer, 1848). O – P. ANSP 362089 (H 49.4 mm). Q. UVZ 83018 (H 54 mm). R – S. P. (E.) piperatoides Pilsbry, 1901, lectotype ANSP 263536 (H 29.1 mm). T – U. P. (E.) argenteus (Jousseaume, 1900), USNM 532637 (H 29 mm). V – W. P. (E.) dalmasi (Dautzenberg, 1900), lectotype RBINS / MT 668 (H 25 mm), photo R. Sablon.

FIGURE 11. Plekocheilus (Eurytus). A – F. P. (E.) coloratus (Nyst, 1845). A – B. CM 3493 (H 83 mm). C – D. USNM 543490 (H 70 mm). E – F. USNM 599523 (H 82 mm). G – H. P. (E.) pirriensis Dall, 1912, holotype USNM 214405 (H 82 mm).