Chondrocyclus alabastris – Connolly 1939: 539 Revision of Chondrocyclus s. l. (Mollusca: Cyclophoridae), with description of a new genus and twelve new species Cole, Mary L. European Journal of Taxonomy 2019 2019-10-22 569 1 92 7QNSL Cole, 2019 Cole 2019 [594,896,897,924] Gastropoda Cyclophoridae Chondrocyclus GBIF Animalia Architaenioglossa 25 26 Mollusca species amathole sp. nov.   Figs 9, 10D–E, 14      Chondrocyclus alabastris– Connolly 1939: 539.    Diagnosis Shell small, very depressed, discoidal to lenticular; protoconch mammillate; periostracum with dense transverse costae developing at periphery broad quadrangular flanges; lamellate axial costae with dense axial riblets, rendering upper edge of each costa scalloped along its length; operculum duplex, exterior portion very shallowly concave to almost flat, multispiral lamella with fringe of very long bristles, fused at their tips, below main fringe is a second shorter fringe of loose bristles; umbilicus wide, exposing all the whorls; radula with two large cusps on second lateral tooth.    Etymology Named after the Amathole Mountains, Eastern Cape, an area of natural and historic interest.    Type material examined   Holotype   SOUTH AFRICA–  Eastern Cape • Amathole Mountains, Hogsback, downstream of Madonnaand Child Waterfall, Southern Mistbelt forest; 32.6068°S, 26.9622°E;  1061 ma.s.l.;  6 Apr. 2014; M. Cole, D. Herbertand L. Davisleg.; NMSA P0640/T4157. ( Fig. 10D–E)   Paratypes   SOUTH AFRICA–  Eastern Cape • 18 specimens; same collection data as for holotype; ELM W3812/ T59•  21 specimens; Hogsback, downstream of Madonnaand Child Waterfall; 32.6068°S, 26.9622°E;  1061 ma.s.l.;  31 Dec. 2008; M. Burseyleg.; ELMD16944/T55•  9 specimens; same collection data as for preceding; ELM W3629/T56•  22 specimens; same collection data as for preceding; M. Coleleg.,  13 Sep. 2012; ELMD17014/T57•  6 specimens; same collection data as for preceding; ELM W3665/ T58•  4 specimens; same collection data as for preceding; NMSA P0601/T4115•  2 specimens; same collection data as for preceding; NHMUK 20120278•  3 specimens; same collection data as for preceding; NMW.Z.2012.065.00008•  3 specimens; same collection data as for preceding; RMNHMOL.33050•  3 specimens; Hogsback, forest above Hobbiton Camp, Southern Mistbelt forest; 32.5958°S, 26.9617°E;  1253 ma.s.l.;  1 Dec. 2005; M. Burseyand V. Ndiboleg.; ELMW02966/T54•  13 specimens; same collection data as for preceding;  13 Sep. 2012; ELMD17013/T60•  13 specimens; same collection data as for preceding; ELM W3662/T61•  28 specimens; same collection data as for preceding;  10 Jun. 2013; M. Coleleg.; ELMD17338/T63•  7 specimens; same collection data as for preceding;  6 Apr. 2014; M. Cole, D. Herbertand L. Davisleg.; ELM W3811/T62•  1 specimen; Kologha Forest, Evelyn valley, north side of Maden Dam, Murray’s Krantz; 32.7257°S, 27.3078°E;  25 Apr. 1998; M. Burseyand N. Smithleg.; alive under a log; ELMD14417/T76•  1 specimen; same collection data as for preceding; ELM W3693/T77•  13 specimens; Isidenge, SE of Mount Kemp, harvested site, south-facing forest; 32.6885° S, 27.2783°E;  1159 ma.s.l.;  5Apr. 2016; M. Coleleg.; ELMD18082/T198•  4 specimens; same collection data as for preceding; ELMW03915/T196•  23 specimens; Isidenge, Sandile’s Rest, small patch of forest near base of Mount Kemp; 32.6614° S, 27.3004° E;  900 ma.s.l.;  7 Apr. 2016; M. Coleleg.; ELMD18077/T200•  8 specimens; same collection data as for preceding; ELMW03914/T195•  3 specimens; Stutterheim, Kologha Forest, waterfall walk from picnic site; 32.5339°S, 27.4308°E;  18 Jan. 2009; M. Coleleg.; ELM W3633/T64•  5 specimens; same collection data as for preceding;  22 Dec. 2009; M. Coleand T. Pretoriusleg.; ELMD16942/T65•  1 specimen; same collection data as for preceding; ELMW03628/T66•  2 specimens; same collection data as for preceding;  11 Dec. 2010; M. Coleleg.; ELM W3634/T67•  4 specimens; Qacu forest; 32.4031°S, 27.4486° E;  6 Apr. 2016; M. Coleleg.; ELMD18126/T201•  5 specimens; same collection data as for preceding; ELMW03917/T197•  5 specimens; Patchwood Farm, forest at source of Quanti River; 32.3924°S, 27.4470°E;  1238 ma.s.l.;  18 Dec. 2012; M. and K. Coleleg.; ELMD17100/T80•  3 specimens; Patchwood Farm, forest below (north of) house; 32.3875° S, 27.4501° E;  1196 ma.s.l.;  6 Apr. 2016; M. Coleleg.; ELMD18140/T199•  8 specimens; Fort Fordyce, south facing slope near top of pass; 32.6843° S, 26.4956° E;  1110 ma.s.l.;  29 Dec. 2008; M. Coleleg.; ELMD16946/T68•  19 specimens; same collection data as for preceding; ELM W3630/T69•  36 specimens; same collection data as for preceding;  3 Jan. 2010; M. Coleleg.; ELMD16947/T70•  12 specimens; same collection data as for preceding; ELM W3632/T71•  1 specimen; same collection data as for preceding;  31 Dec. 2011; ELMD17005/T72•  7 specimens; same collection data as for preceding; ELM W3631/T73•  4 specimens; same collection data as for preceding; NHMUK 20120279•  4 specimens; same collection data as for preceding; NMSA P0600/T4111•  10 specimens; Fort Fordyce, south facing slope behind lookout; 32.6956°S, 26.4857°E;  1120 ma.s.l.;  30 Dec. 2008; M. Coleleg.; ELMD16950/T74•  5 specimens; same collection data as for preceding;  17 Jan. 2015; ELMD17860/T75•  10 specimens; same collection data as for preceding;  1 Apr. 2016; ELMD18116/ T202•  3 specimens; Kapp River, north bank, indigenous riverine forest on shady, south-facing slope; 33.4830° S, 27.0807°E;  30 Dec. 2002; M. Bursey; ELM W3694/T78•  2 specimens; Beggars’ Bush,  15 kmE of Grahamstown, south facing slope; 33.2871°E, 26.6885°E;  25 Feb. 2013; M. Cole, G. Godfreyand S. Ritcherleg.; ELMD17289/T79.   Other material examined   SOUTH AFRICA–  Eastern Cape • 5 specimens; Amathole Mountains: Hogsback, downstream of Madonnaand Child Waterfall, Southern Mistbelt forest; 32.6068° S, 26.9622°E;  1061 ma.s.l.;  25 Jan. 2002; M. Burseyleg.; ELMD13627•  5 specimens; same collection data as for preceding; ELMW02834•  6 specimens; Hogsback, forest above Hobbiton Camp, Southern Mistbelt forest; 32.5958° S, 26.9617° E,  1253 ma.s.l.;  31 Dec. 2008; M. Burseyleg.; ELMD16945•  14 specimens; Kologha Forest, near Stutterheim; 32.5333°S, 27.3667° E;  27 Nov. 1993; R. Scottleg.; ELMD14405•  5 specimens; Stutterheim, Kologha Forest, forest drive near Protea Hill; 32.5558° S, 27.3175°E;  26 Jan. 2010; M. Coleleg.; ELMD16943•  1 specimen; Stutterheim, Kologha Forest, waterfall walk from picnic site; 32.5339° S, 27.4308°E;  30 Dec. 2012; M. Coleleg.; ELMD17623•  3 specimens; Fort Fordyce, kloof with watercourse, running eastwards; 32.6704° S, 26.4851° E;  1137 ma.s.l.;  29 Dec. 2008; M. Coleleg.; ELMD16949•  3 specimens; Fort Fordyce, top of krantz near campsite; 32.6813° S, 26.4802°E;  1134 ma.s.l.;  30 Dec. 2011; M. Coleleg.; ELMD16948•  7 specimens; same collection data as for preceding; ELM W3631•  2 specimens; Patchwood Farm, forest at source of Quanti River; 32.3924° S, 27.4470° E,  1238 ma.s.l.;  18 Dec. 2012; M and K Coleleg.; ELMW03695•  3 specimens; Patchwood Farm, forest below (north of) house; 32.3875° S, 27.4501°E);  1196 ma.s.l.; M. Coleleg.;  18 Dec. 2012 ELMD17107•  5 specimens; same collection data as for preceding; ELMW03699.    Description SHELL ( Fig. 14A–C). Small, very depressed, discoidal to lenticular, adult diameter 5.09–7.4 mm, height 2.34–3.92 mm, diameter:height 1.52–2.54 (n = 68 measured in three populations spanning the Amathole Mountains). Spire little exserted, apex mammillate. Embryonic shell ( Fig. 14E) 2.25 whorls, microscopically malleate, junction between embryonic shell and teleoconch evident with development of costae on teleoconch. Teleoconch comprising 2.5 whorls, convex, very rapidly increasing, suture deeply impressed. Aperture circular, last whorl descending near aperture, peristome simple, continuous and free. Umbilicus wide, exposing all the whorls. Periostracum glossy, honey-brown and lacquerlike with dense lamellate costae at regular intervals, 73–167 (n = 58) on last whorl, expanded into broad quadrangular flanges at periphery; lamellar blades ridged with dense axial riblets, rendering upper edge of each blade scalloped along its length; intervals between costae with about 6 microscopic axial threads. Shell translucent glossy honey-brown when fresh. LIVING ANIMAL. Head, tentacles and snout dark grey, underside of foot creamy white. OPERCULUM ( Fig. 14D, G). Duplex, very shallowly concave to almost flat; outer portion consists of multispiral lamella with five–six whorls; each step-shaped; height of lamellar blade very low; fringe of very long bristles, fused at their tips but not along their entire length, below main fringe is a second shorter fringe of relatively loose bristles, long outer fringe of each whorl fused to lamella of following whorl, but due to its long length, fringe forms convex curve between one whorl and next, height of fringe of each whorl exceeds height of lamellar blade, outermost lamellar fringe very long and is reflexed over peristome in life, but operculum can be withdrawn into aperture. RADULA ( Fig. 14F). Rachidian with five cusps, central cusp approx. twice length of outer cusps; first lateral tooth usually with three large cusps, small fourth cusp and a vestigial fifth, third cusp (from centre) largest; second lateral with two large cusps, second cusp (from centre) longer and broader than first, a very small third cusp and a vestigial fourth. PENIS ( Fig. 14H–I). Shaft more or less cylindrical, slightly flattened, with prominent annular rugae, distal end smooth, intromittent organ short.    Distribution and habitat Throughout the Amathole Mountains and extending eastwards towards the Kei River in isolated forest patches; also recorded at Beggars’ Bush near Grahamstown and at Kap River Nature Reserve near the mouth of the Great Fish River ( Fig. 9). Amathole Mistbelt forest (Southern Mistbelt Forest group) ( von Maltitz et al.2003) and Great Fish Thicket (Kap River) ( Hoare et al.2006); in leaf-litter.    Remarks The operculum of  Chondrocyclus amathole sp. nov.is unique among all other  Chondrocyclusspecies in its flatness, its long terminal fringe and relatively long secondary fringe below it and the low height of the multispiral lamellar blade.  Chondrocyclus amathole sp. nov.was sister to  C. alabastrisand formed a well-supported clade (the Southern-Eastern Capeclade) with  C. herberti sp. nov.and  C. silvicolus sp. nov.( Fig. 1). The periostracal flanges of  C. amathole sp. nov.were broad and quadrangular while in the other three species in its clade, the flanges were pointed. In this feature,  C. amathole sp. nov.resembled species in the Eastern Clade, but differed from species in the latter clade in all other diagnostic morphological features, i.e., single row of flanges at periphery, unique operculum, radula with two large cusps on second lateral tooth, cyclindrical penis ( Cole et al.2019). The Amathole Range is an outlier of the southern end of the Drakensberg, isolated by the valleys of the Great Fish and Great Kei Rivers ( Stuckenberg 1962) and is known for endemicity of many low-vagility forest-dependent taxa, including molluscs ( Connolly 1939; Herbert & Moussalli 2010; unpublished data), forest-floor spiders ( Griswold 1985), velvet worms ( Daniels & Ruhberg 2010), harvestmen ( de Bivort & Giribet 2010) and two frogs. Furthermore, populations of velvet worms in close geographic proximity are genetically discrete, suggesting that the historic contractions and expansions of forests in this region left a significant and complex impact on the phylogeography of low-vagility organisms ( Daniels et al.2017).   Fig. 14.  Chondrocyclus amathole sp. nov. A. Aperture view (ELM W3629/T56). B. Dorsal view (ELM W3629/T56). C. Ventral view (ELM W2966/T54). D, G. Operculum (ELM W2966/T54). E. Protoconch (ELM W3812/T59). F. Portion of radula (ELM W3628/T66). H. Penis, dorsal view (ELM W3628/T66). I. Penis, lateral view (ELM W3632/T71). Scale bars: A–C = 2 mm; D, G–I = 1 mm; E = 500 µm; F = 50 µm.   Chondrocyclus amathole sp. nov.has been recorded in two localities geographically separated from the mistbelt forests of the Amatholes, indicative of past vegetation shifts. The population of  C. amathole sp. nov.in the outlying forest patch, Beggars’ Bush near Grahamstown, was separated from populations of the sister species,  C. alabastrison the opposite ridge, by a dry intervening valley. Beggars’ Bush is currently separated from the Amathole mountains by the dry Great Fish River basin although both regions are designated Amathole Mistbelt in certain classification systems and distinguished from other Southern Mistbelt forests ( von Maltitz et al.2003). The presence of  C. amathole sp. nov.near the mouth of the Great Fish River suggests a link between the coast and the Amathole Mountains, a pattern also demonstrated in other taxa (e.g.,  Streptocarpus( Hughes et al.2005)). The Great Fish River basin has been identified as an area of persistence of thicket during the contractions of the LGM ( Potts et al.2012). The distribution pattern in this eastern subclade of the Southern-Eastern Capeclade (i.e.,  C. alabastrisand  C. amathole sp. nov.) is mirrored by rhytidid snails ( Herbert & Moussalli 2010, 2016). 2429182615 2014-04-06 NMSA M. Cole & D. Herbert & L. Davis South Africa Amathole Mountains 1061 -32.6068 Madonna 6 26.9622 Hogsback 25 26 P0640 1 Eastern Cape holotype 2429182727 2014-04-06 ELM M. Cole & D. Herbert & L. Davis South Africa Amathole Mountains 1061 -32.6068 Madonna 6 26.9622 Hogsback 25 26 W3812 18 Eastern Cape paratype 2429182721 2008-12-31 ELM M. Bursey South Africa Hogsback 1061 -32.6068 Child Waterfall 6 26.9622 Madonna 25 26 21 Eastern Cape paratype 2429182682 2008-12-31 ELM M. Bursey South Africa Hogsback 1061 -32.6068 Child Waterfall 6 26.9622 Madonna 25 26 W3629 9 Eastern Cape paratype 2429182709 2012-09-13 ELM M. Cole South Africa Hogsback 1061 -32.6068 Child Waterfall 6 26.9622 Madonna 25 26 22 Eastern Cape paratype 2429182617 2012-09-13 ELM M. Cole South Africa Hogsback 1061 -32.6068 Child Waterfall 6 26.9622 Madonna 25 26 W3665 6 Eastern Cape paratype 2429182568 [265,1145,1930,1957] 2012-09-13 NMSA M. Cole South Africa Hogsback 1061 -32.6068 Child Waterfall 6 26.9622 Madonna 25 26 P0601 4 Eastern Cape paratype 2429182656 2012-09-13 NHMUK M. Cole South Africa Hogsback 1061 -32.6068 Child Waterfall 6 26.9622 Madonna 25 26 NHMUK 20120278 2 Eastern Cape paratype 2429182629 2012-09-13 NMW M. Cole South Africa Hogsback 1061 -32.6068 Child Waterfall 6 26.9622 Madonna 25 26 3 Eastern Cape paratype 2429182584 [513,1379,2000,2027] 2012-09-13 RMNH M. Cole South Africa Hogsback 1061 -32.6068 Child Waterfall 6 26.9622 Madonna 25 26 3 Eastern Cape paratype 2429182646 2005-12-01 ELM M. Bursey & V. Ndibo South Africa Hogsback 1253 -32.5958 Southern Mistbelt forest 6 26.9617 Hobbiton Camp 26 27 3 Eastern Cape paratype 2429182581 2012-09-13 ELM M. Bursey & V. Ndibo South Africa Hogsback 1253 -32.5958 Southern Mistbelt forest 6 26.9617 Hobbiton Camp 26 27 13 Eastern Cape paratype 2429182717 2012-09-13 ELM M. Bursey & V. Ndibo South Africa Hogsback 1253 -32.5958 Southern Mistbelt forest 6 26.9617 Hobbiton Camp 26 27 W3662 13 Eastern Cape paratype 2429182648 2013-06-10 ELM M. Cole South Africa Hogsback 1253 -32.5958 Southern Mistbelt forest 6 26.9617 Hobbiton Camp 26 27 28 Eastern Cape paratype 2429182705 2014-04-06 ELM M. Cole & D. Herbert & L. Davis South Africa Hogsback 1253 -32.5958 Southern Mistbelt forest 6 26.9617 Hobbiton Camp 26 27 W3811 7 Eastern Cape paratype 2429182562 1998-04-25 ELM M. Bursey & N. Smith South Africa Kologha Forest -32.7257 Maden Dam 6 27.3078 Evelyn valley 26 27 1 Eastern Cape paratype 2429182695 1998-04-25 ELM M. Bursey & N. Smith South Africa Kologha Forest -32.7257 Maden Dam 6 27.3078 Evelyn valley 26 27 W3693 1 Eastern Cape paratype 2429182673 2016-04-05 ELM M. Cole South Africa 1159 -32.6885 Isidenge 6 27.2783 26 27 13 Eastern Cape paratype 2429182660 2016-04-05 ELM M. Cole South Africa 1159 -32.6885 Isidenge 6 27.2783 26 27 4 Eastern Cape paratype 2429182616 2016-04-07 ELM M. Cole South Africa Isidenge 900 -32.6614 Mount Kemp 6 27.3004 Sandile's Rest 26 27 23 Eastern Cape paratype 2429182563 [513,1379,690,717] 2016-04-07 ELM M. Cole South Africa Isidenge 900 -32.6614 Mount Kemp 6 27.3004 Sandile's Rest 26 27 8 Eastern Cape paratype 2429182703 2009-01-18 ELM M. Cole South Africa -32.5339 Kologha Forest 6 27.4308 Stutterheim 26 27 W3633 3 Eastern Cape paratype 2429182606 2009-12-22 ELM M. Cole & T. Pretorius South Africa -32.5339 Kologha Forest 6 27.4308 Stutterheim 26 27 5 Eastern Cape paratype 2429182633 2009-12-22 ELM M. Cole & T. Pretorius South Africa -32.5339 Kologha Forest 6 27.4308 Stutterheim 26 27 1 Eastern Cape paratype 2429182674 2010-12-11 ELM M. Cole South Africa -32.5339 Kologha Forest 6 27.4308 Stutterheim 26 27 W3634 2 Eastern Cape paratype 2429182684 2016-04-06 ELM M. Cole South Africa -32.4031 Qacu forest 6 27.4486 26 27 4 Eastern Cape paratype 2429182607 [513,1379,902,929] 2016-04-06 ELM M. Cole South Africa -32.4031 Qacu forest 6 27.4486 26 27 5 Eastern Cape paratype 2429182567 2012-12-18 ELM M. and K. Cole South Africa 1238 -32.3924 Quanti River 6 27.447 Patchwood Farm 26 27 5 Eastern Cape paratype 2429182634 2016-04-06 ELM M. Cole South Africa 1196 -32.3875 Patchwood Farm 6 27.4501 26 27 3 Eastern Cape paratype 2429182576 2008-12-29 ELM M. Cole South Africa 1110 -32.6843 Fort Fordyce 6 26.4956 26 27 8 Eastern Cape paratype 2429182635 2008-12-29 ELM M. Cole South Africa 1110 -32.6843 Fort Fordyce 6 26.4956 26 27 W3630 19 Eastern Cape paratype 2429182715 2010-01-03 ELM M. Cole South Africa 1110 -32.6843 Fort Fordyce 6 26.4956 26 27 36 Eastern Cape paratype 2429182589 [368,1226,1148,1175] 2010-01-03 ELM M. Cole South Africa 1110 -32.6843 Fort Fordyce 6 26.4956 26 27 W3632 12 Eastern Cape paratype 2429182600 2011-12-31 ELM M. Cole South Africa 1110 -32.6843 Fort Fordyce 6 26.4956 26 27 1 Eastern Cape paratype 2429182573 2011-12-31 ELM M. Cole South Africa 1110 -32.6843 Fort Fordyce 6 26.4956 26 27 W3631 7 Eastern Cape paratype 2429182628 2011-12-31 NHMUK M. Cole South Africa 1110 -32.6843 Fort Fordyce 6 26.4956 26 27 NHMUK 20120279 4 Eastern Cape paratype 2429182565 [334,1203,1254,1281] 2011-12-31 NMSA M. Cole South Africa 1110 -32.6843 Fort Fordyce 6 26.4956 26 27 P0600 4 Eastern Cape paratype 2429182665 2008-12-30 ELM M. Cole South Africa 1120 -32.6956 Fort Fordyce 6 26.4857 26 27 10 Eastern Cape paratype 2429182718 2015-01-17 ELM M. Cole South Africa 1120 -32.6956 Fort Fordyce 6 26.4857 26 27 5 Eastern Cape paratype 2429182577 2016-04-01 ELM M. Cole South Africa 1120 -32.6956 Fort Fordyce 6 26.4857 26 27 10 Eastern Cape paratype 2429182758 2002-12-30 ELM M. Bursey South Africa -33.483 Kapp River 6 27.0807 26 27 W3694 3 Eastern Cape paratype 2429182611 2013-02-25 ELM M. Cole & G. Godfrey & S. Ritcher South Africa Beggars' Bush 26.6885 26 27 2 Eastern Cape paratype 2429182662 2002-01-25 ELM M. Bursey South Africa Amathole Mountains 1061 -32.6068 Madonna 6 26.9622 Hogsback 26 27 5 Eastern Cape 2429182799 2002-01-25 ELM M. Bursey South Africa Amathole Mountains 1061 -32.6068 Madonna 6 26.9622 Hogsback 26 27 5 Eastern Cape 2429182719 2008-12-31 ELM M. Bursey South Africa Hogsback 1253 -32.5958 Southern Mistbelt forest 6 26.9617 Hobbiton Camp 26 27 6 Eastern Cape 2429182730 1993-11-27 ELM R. Scott South Africa -32.5333 Stutterheim 6 27.3667 Kologha Forest 26 27 14 Eastern Cape 2429182666 2010-01-26 ELM M. Cole South Africa Stutterheim -32.5558 Protea Hill 6 27.3175 Kologha Forest 26 27 5 Eastern Cape 2429182557 2012-12-30 ELM M. Cole South Africa -32.5339 Kologha Forest 6 27.4308 Stutterheim 26 27 1 Eastern Cape 2429182678 2008-12-29 ELM M. Cole South Africa 1137 -32.6704 Fort Fordyce 6 26.4851 26 27 3 Eastern Cape 2429182687 2011-12-30 ELM M. Cole South Africa 1134 -32.6813 Fort Fordyce 6 26.4802 26 27 3 Eastern Cape 2429182699 2011-12-30 ELM M. Cole South Africa 1134 27 28 -32.6813 Fort Fordyce 6 26.4802 26 27 W3631 7 Eastern Cape 2429182765 2012-12-18 ELM M and K Cole South Africa 1238 -32.3924 Quanti River 6 27.447 Patchwood Farm 27 28 2 Eastern Cape 2429182590 2012-12-18 ELM M. Cole South Africa 1196 -32.3875 Patchwood Farm 6 27.4501 27 28 3 Eastern Cape 2429182612 [378,1162,374,401] 2012-12-18 ELM M. Cole South Africa 1196 -32.3875 Patchwood Farm 6 27.4501 27 28 5 Eastern Cape