Revision of Chondrocyclus s.l. (Mollusca: Cyclophoridae), with description of a new genus and twelve new species

Chondrocyclus Ancey, 1898 is a genus of nine species of African operculate land snails restricted to indigenous forest and mesic thicket. Worn specimens (i.e., without a periostracum or operculum), on which some species descriptions and records were based, appear to be indistinguishable morphologically. A comprehensive revision of Chondrocyclus s.l. is provided here based on comparative morphological examinations of the shell, protoconch, periostracum, operculum, radula and penis, and on mitochondrial genes cytochrome c oxidase subunit I and 16S rRNA. Two genus-level lineages are recognised, Chondrocyclus s.s. and Afrocyclus gen. nov. Revised species descriptions are given for seven species. Two species, C. meredithae Bruggen, 1983 and C. chirindae Bruggen, 1986 both from north of South Africa, are removed from Chondrocyclus. Twelve new species are described: C. herberti sp. nov., C. silvicolus sp. nov., C. amathole sp. nov., C. pondoensis sp. nov., C. devilliersi sp. nov., C. pulcherrimus sp. nov., C. cooperae sp. nov., C. langebergensis sp. nov., C. kevincolei sp. nov., A. oxygala gen. et sp. nov., A. potteri gen. et sp. nov. and A. bhaca gen. et sp. nov. This is the first detailed systematic revision of an Afrotropical cyclophorid group to include morphological and molecular data. This study complements research on other taxa of low-vagility forest-dwelling habitat specialists by providing comparative distribution data for an independent, widespread group. Such evidence is urgently needed for conservation of South Africa’s threatened forest biome.


Introduction
Madagascan cyclophorids were also classifi ed in Chondrocyclus (Fischer-Piette et al. 1993), but these have been reassigned to three other genera .
The fi rst molecular systematic study of an Afrotropical cyclophorid group presented a molecular phylogeny of Chondrocyclus s.l. (Fig. 1), and assessed the taxonomic value of a range of morphological characters, using fresh material and museum specimens from across the known range of the genus (Cole et al. 2019). Concordance of molecular and morphological data confi rmed the taxonomic status of the seven South African species and provided evidence for the existence of at least twelve undescribed species (Cole et al. 2019). These data show that Chondrocyclus s.l. underwent at least two major Fig. 1. Bayesian Inference majority consensus tree of the "taxa complete" concatenated CO1-16S sequence dataset for Chondrocyclus s.l. Ancey, 1898 (84 sequences) and seven other caenogastropods with the tree rooted on Pomacea insularum (d'Orbigny, 1837) (modifi ed from Cole et al. 2019). Colours indicate major clades of Chondrocyclus Ancey, 1898 and correspond with colours used in maps. Support values are given as posterior probabilities for Bayesian analysis above nodes (only values ≥ 0.95 are shown) and as bootstrap percentages for ML analysis below nodes (only values ≥ 65% are shown); maximal support (Bayesian = 1, ML = 100%) is indicated by an asterisk. Scale bar indicates substitutions per site.
European Journal of Taxonomy 569: 1-92 (2019) radiations, one mainly Afromontane and the other largely coastal. Chondrocyclus s.s. contained four major clades distributed coastwards of the Great Escarpment, between the Cape Peninsula and northern KwaZulu-Natal (Fig. 2). This radiation showed a primary dichotomy into an Eastern clade and a western group of three clades (Southwestern Cape, Overberg and Southern-Eastern Cape), in the region of the country where a transition between subtropical and temperate infl uences is well documented across other faunal and fl oral taxa (e.g., Branch & Branch 1981;Cowling & Pierce 2009). The other lineage was even more widely distributed in latitude and altitude, occupying isolated Afromontane forest patches along the Great Escarpment from northern South Africa (Limpopo province, 23° S) to the Eastern Cape and extending to the coast (Fig. 2). This lineage is recognised herein as a separate genus, Afrocyclus gen. nov., which requires further molecular and morphological data from throughout its range to unravel its systematics.
Cyclophoridae in the Afrotropical region, although not comparable in species and genus-level diversity to the Asian tropics, is taxonomically diverse and includes a few species-rich groups in Madagascar Emberton & Pearce 1999;Emberton et al. 2010). In mainland Africa, cyclophorids are widely distributed, and currently 20 species are recognised in three genera: Chondrocyclus (seven species in South Africa, one species in Zimbabwe and one in Malawi), Cyathopoma W. & H. Blanford, 1861 (nine species across East, Central and West Africa) and Elgonocyclus Verdcourt, 1982 (one species in East and Central Africa and one undescribed species in South Africa). Historically, South African Cyclophoridae were assigned to various genera (Pfeiffer 1855;Craven 1880;Sturany 1898;Kobelt 1902).
The taxonomy of some cyclophoroidean taxa is in a state of fl ux, including the family placement of Chondrocyclus. The Cyclophoridae Gray, 1847 was considered to be the most speciose For the anatomical study, living adult specimens were drowned in sealed containers and preserved in 70% ethanol for dissection. Cleaning and dissections were performed under a Euromex dissecting microscope. Shells and opercula were cleaned in an ultrasonic bath and then manually under the dissecting microscope to remove caked soil from within the delicate ornamentation of both structures. Radulae were extracted by maceration of the body anterior in dilute NaOH and the radula teased out from the surrounding tissue, and rinsed in distilled water. The radula was then placed in a drop of ethanol on a stub with double-sided carbon tape and manouvered into position as the ethanol dried. The teeth at the outer edge of each row were folded outwards to expose the seven teeth per row. Penes were prepared for examination by cutting away the mantle tissue to expose the penis where it lies on the dorsal surface behind the right tentacle. Samples were dehydrated by placing in a series of ethanol of increasing concentration, and dried in a Polaron Critical Point Drier. Shells, opercula, radulae and penes for Scanning Electron Microscope examination were coated with gold-palladium and examined at 5-20 kv accelerating voltage in a Tescan Vega SEM. Colour photographs of NHM types were supplied by the NHM. Photographs of holotypes were taken using a Zeiss Stemi 2000-C dissecting microscope with AxioCam ERc5s digital camera and stacked images were then combined using Helicon Focus Pro (Helicon Soft Ltd) to provide extended depth of fi eld. Figure 8 was taken using an Olympus SZX 16 microscope with the AnalySIS programme.
Shell diameter at widest point (D) and height (H) measurements were made with the shell held in apertural view with the axis of coiling vertical, using an eyepiece graticule. Shell diameter:height ratios (D:H) were calculated. The number of lamellate axial costae on the last whorl were counted. Immature shells (peristome not detached from last whorl) were excluded from the data used to calculate D:H ratios and densities of lamellate costae. The number of protoconch whorls were counted as shown in Fig. 3 although the precise number was often diffi cult to determine.
Systematic descriptions are provided for all taxa identifi ed during this study. Type material is listed with redescriptions and descriptions. COLE M.L., Revision of cyclophorid snails Chondrocyclus s.l. by three laterals in each row), all with well-developed, sharp cusps. The central cusp of the rachidian tooth was usually approx. twice as long as the fl anking cusps, but in two taxa in the Eastern clade all fi ve cusps were uniform in size. There were either two or three large cusps on the second lateral tooth, consistent within a clade. The penis lay on the dorsal surface immediately behind the right tentacle, and consisted of a muscular shaft with annular rugae and a shorter, tapering intromittent organ without fl agellum. The shaft was more or less cylindrical in most taxa, but dorso-ventrally fl attened and laterally expanded towards the distal end in the Eastern clade.
Several characters closely resembled those features described in other cyclophorids or species in related families in Africa, Asia and New Zealand (e.g., Morton 1952;Bruggen 1983Bruggen , 1985Bruggen , 1986Marshall & Barker 2007;Lee et al. 2008a;Rowson et al. 2010b;Páll-Gergely et al. 2017b). These include multiple spiral rows of hairs on the periostracum, taenioglossate radula and numbers of cusps on teeth, mammillation of the protoconch, and penis position and shape.

Diagnosis
Shell small, depressed, lenticular; protoconch mammillate and tilted; periostracum with axial costae developing at the periphery broad triangular fl anges each terminating in a long, hair-like spine; operculum fl attened, exterior portion shallowly concave, with step-shaped multispiral lamella terminating in a long solid fringe; radula with three large cusps on second lateral tooth.

Etymology
The specifi c name is derived from the Latin 'convex', meaning 'arched or vaulted'.

Fig. 8.
Close-up of periostracum of species of the Southern-Eastern Cape clade and Chondrocyclus convexiusculus (Pfeiffer, 1855), showing variation in detail of fl anges at periphery. A. C. alabastris (Craven, 1880)  OPERCULUM ( Fig. 7C-E). Duplex outer portion shallowly concave, and consisting of a multispiral lamella with fi ve whorls, at its growing edge the top edge of the lamellar blade is sloping, becoming perpendicular to the disc where it is attached; fringe of fused bristles emanates near top of vertical portion of lamellar blade on its outer surface and curves upwards and outwards forming a shallow furrow between fringe and lamellar blade, a second very short fringe emanates from lower down on the lamellar blade; top edge of lamella does not project above level of fringe; fringe of each whorl fused to lamella of following whorl. Fringe overlaps disc slightly, but operculum can be withdrawn into aperture.
RADULA (Fig. 7F). Rachidian with fi ve cusps, central approx. twice as long as two on either side of it; fi rst lateral tooth with four cusps and sometimes a vestigial fi fth, fi rst three cusps (from centre) gradually increasing in size and fourth considerably smaller; second lateral tooth with two large cusps and two small cusps towards outside. PENIS ( Fig. 7G-H). Shaft more or less straight-sided and slightly fl attened, with numerous annular rugae, intromittent organ short.

Distribution and habitat
Endemic to the Eastern Cape, centered around Port Elizabeth and including the Albany district ( Fig. 9). Found in a wider variety of vegetation types than other Chondrocyclus species: Algoa Dune Strandveld, Albany Coastal Forest, Southern Mistbelt Forest, Great Fish Thicket (von Maltitz et al. 2003;Hoare et al. 2006;; in leaf-litter.

Remarks
There is variation in size of shells between populations (Table 1), an observation also recorded by Connolly (1939). Shells in some populations are among the largest recorded in the genus (the only other species attaining a similar size is C. amathole sp. nov.). However, the types (Fig. 6A) from Cape Receife, populations at The Island Nature Reserve (40 km west of type locality) and at Gamtoos River mouth are smaller on average and the population at Gamtoos mouth has a distinctly less depressed shape than is typical for the species. Its diameter/height proportions resemble those of relatively less depressed species, e.g., C. convexiusculus, C. herberti sp. nov. and A. exsertus gen. et comb. nov.
There was a relatively large intraspecifi c divergence in C. alabastris (Cole et al. 2019) and the species contained several very well-supported subclades (Fig. 1). There was a distinct lineage in the mistbelt forest patches on the Rietberg ridge south of Grahamstown (Paradise Kloof and Coleridge in Fig. 1). This lineage exhibited a conspicuous variation from other populations of C. alabastris: the peripheral extensions of the axial lamellae were not serrated along the edges and each fl ange tapered gradually into a short spine tip (Fig. 8B). Specimens from these localities superfi cially resembled C. convexiusculus (Fig. 8C). However, the spines are longer in C. convexiusculus, emanating from a relatively short, broad triangular proximal portion while in C. alabastris the triangular fl anges taper gradually into the short spine tip (Fig. 8B).
The lamellar blade of the outer portion of the operculum of C. alabastris is relatively lower and hence the operculum is less concave than that of C. convexiusculus, and has a very short fringe below the long fringe, absent in C. convexiusculus.

Diagnosis
Shell very small, moderately depressed; periostracum with widely spaced axial costae bearing broad, distally rounded fl anges at the shell periphery; operculum duplex, exterior portion shallowly concave, with step-shaped multispiral lamella terminating in a long solid fringe refl exed over peristome; radula with two large cusps on second lateral tooth.

Etymology
Named for Dai Herbert, Malacologist at the KwaZulu-Natal Museum for 33 years, in recognition of his expertise in South African molluscs and in acknowledgement of his mentorship.

Type material examined
Holotype SOUTH AFRICA -Eastern Cape • Baviaanskloof, Western Poortjies area; 33.6536° S, 24 . 5161° E; 9 Oct. 2000; D. Herbert leg.; riverine thicket, in leaf-litter; NMSA P1043/T4277. (Fig. 10A 1.46-1.94 (n = 20). Spire moderately exserted, protoconch acutely mammillate and tilted (Fig. 12A). Embryonic shell (Fig. 12D) just over two whorls, microscopically malleate for approx. two whorls, axial costae begin to develop just after two whorls. Teleoconch comprising approx. 2.5 whorls, very convex, rapidly increasing, suture impressed. Aperture circular, last whorl descending near aperture, peristome simple, continuous and free. Umbilicus deep, exposing all the whorls. Periostracum glossy, light yellowish-brown and lacquer-like with relatively widely spaced lamellate axial costae at regular intervals, 39-51 (n = 22) on last whorl ( Fig. 12A-C), which produce broad fl anges with rounded apices at periphery (Fig. 8D, 12E); intervals between costae with six to nine distinct microscopic axial threads. Shell translucent and corneous when fresh. LIVING ANIMAL. Very dark grey. OPERCULUM ( Fig. 12G-H). Duplex, outer portion shallowly concave and consisting of multispiral lamella with 4.5 whorls, each step-shaped and increasing in height towards outside, sloping at growing edge; very long fringe of fused bristles emanates near top of lamellar blade with a slight groove between fringe and lamellar blade; lamellar blade does not project above level of fringe; fringe of each whorl fused to lamella of following whorl. RADULA (Fig. 12F). Rachidian with fi ve cusps, central one approx. twice length of outer two on each side; fi rst lateral tooth with four cusps and a vestigial fi fth, third cusp (from centre) the largest; second lateral tooth with two very large cusps and two small ones on the outer side (outermost sometimes vestigial). PENIS (Fig. 12I). Shaft more or less cylindrical, distal half slightly expanded on left, intromittent organ short.

Remarks
The shell is less depressed than in the majority of other species of Chondrocyclus. The population of C. alabastris at Gamtoos River mouth has similar proportions but the periostracal fl anges are serrated. The periostracum of C. herberti sp. nov. is distinctive; the fl anges have broadly rounded apices and an unserrated margin. C. convexiusculus also has similar proportions and widely spaced costae, but the shape of the fl anges as well as the radulae, distinguish the species. The operculum of C. herberti sp. nov. is more concave than that of C. alabastris and lacks the furrow between the vertical portion of lamellar blade and fringe. Chondrocyclus herberti sp. nov. is known only from the narrow valley of the Baviaanskloof, in isolated patches of relatively dry thicket and in thicket/fynbos transitional areas. C. herberti sp. nov. and C. alabastris have not been recorded sympatrically, although they occur in steep-sided river valleys in close proximity to each other (Fig. 9). Both of these valleys are tributaries of the Gamtoos River, a postulated dispersal corridor between the coast and the Baviaanskloof (Geldenhuys 1997). Watersheds may have been signifi cant barriers to gene fl ow for thicket plant and insect species during glacial periods as thicket retracted downward into catchment valleys and survived in isolated refugia (Price et al. 2010;Potts et al. 2012). Terrestrial fauna with poor dispersal ability were also bounded by watersheds during Pleistocene climate cycling (Herbert & Moussalli 2010).
When the animals were killed by drowning, they survived for a couple of weeks fully submerged with head and foot extended, which although not quantifi ed, was longer than any other species. Chondrocyclus herberti sp. nov. can be very abundant; D. Herbert (pers. comm.) who fi rst collected the species at Poortjies, found in excess of 1000 shells per m 2 in a leaf litter sample.

Diagnosis
Shell small, moderately depressed; protoconch mammillate and tilted; periostracum with widely spaced axial costae developing at the periphery broad fl anges with pointed tips, tips are rounded not spinetipped; operculum duplex, exterior portion shallowly concave, with step-shaped multispiral lamella terminating in a long solid fringe refl exed over peristome; radula with two large cusps on second lateral tooth.

Etymology
The specifi c name is derived from the Latin 'silvicolus', meaning 'an inhabitant of the woods' and referring to the occurrence of the species in the largest forest blocks in South Africa, in the Knysna area. Description SHELL ( Fig. 13A-C). Small, moderately depressed, adult diameter 3.76-6.10 mm, height 2.09-3.84 mm, diameter:height 1.5-1.97 (n = 49). Spire moderately exserted, protoconch mammillate and tilted. Embryonic shell (Fig. 13D) approx. 2.5 whorls, microscopically malleate, junction between embryonic shell and teleoconch evident with development of costae on teleoconch. Teleoconch comprising approx. 2.5 whorls, very convex, rapidly increasing, suture impressed. Aperture circular, last whorl descending near aperture, peristome simple, continuous and free. Umbilicus wide and deep, exposing all the whorls. Periostracum glossy, light yellowish-brown and lacquer-like with relatively widely spaced ridged, lamellate axial costae at regular intervals, 59-79 (n = 13) on last whorl, which produce broad, pointed fl anges with rounded tips at periphery (Fig. 13E); intervals between costae with six to nine distinct microscopic axial threads. Shell translucent golden brown or creamy white when fresh. LIVING ANIMAL. Head, tentacles and snout pigmented with grey; underside of foot creamy white. OPERCULUM ( Fig. 13G-H). Duplex, outer portion shallowly concave and consisting of multispiral lamella with 4.75 whorls, each step-shaped and increasing in height towards outside, sloping at growing edge; long fringe of fused bristles emanates near top of lamellar blade with a slight groove between fringe and lamellar blade; lamellar blade does not project above level of fringe; fringe of each whorl fused to lamella of following whorl. RADULA (Fig. 13F). Rachidian with fi ve cusps, central one approx. twice length of outer two on each side; fi rst lateral tooth with three large cusps, one small and a vestigial fi fth, third cusp (from centre) the largest; second lateral tooth with two very large cusps and two small ones on the outer side (outermost sometimes vestigial). PENIS (Fig. 13I). Shaft more or less cylindrical, intromittent organ short.

Distribution and habitat
Western Cape, from Storms River to Wilderness (Touw River) (Fig. 9). Indigenous Southern Cape Afrotemperate Forest and Western Cape Milkwood Forest (von Maltitz et al. 2003); in leaf-litter.

Remarks
The shell is less depressed than in the majority of other species of Chondrocyclus. Chondrocyclu. herberti sp. nov. has similar proportions, but the costae in C. silvicolus sp. nov. are not as widely spaced (Table 2) and have more pointed ends. C. silvicolus sp. nov. shells are also larger on average ( Table 2). The two species inhabit different environmental conditions. C. silvicolus sp. nov. is a species of the ancient Southern Cape Afrotemperate forests, while C. herberti sp. nov. is adapted to drier conditions. Forest patches between the Baviaanskloof with C. herberti sp. nov. and the Southern Cape Afrotemperate forests where C. silvicolus sp. nov. occurs, have not been sampled.
Chondrocyclus convexiusculus has similar proportions and relatively widely spaced costae, but the ends of the fl anges are produced into a narrow point. The operculum is less concave and the radula differs in having three rather than two large cusps on the second lateral tooth.

Diagnosis
Shell small, very depressed, discoidal to lenticular; protoconch mammillate; periostracum with dense transverse costae developing at periphery broad quadrangular fl anges; 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 fl at, 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. 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 fl anges 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 fl at; outer portion consists of multispiral lamella with fi ve-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 refl exed over peristome in life, but operculum can be withdrawn into aperture. RADULA (Fig. 14F). Rachidian with fi ve cusps, central cusp approx. twice length of outer cusps; fi rst lateral tooth usually with three large cusps, small fourth cusp and a vestigial fi fth, third cusp (from centre) largest; second lateral with two large cusps, second cusp (from centre) longer and broader than fi rst, a very small third cusp and a vestigial fourth. PENIS ( Fig. 14H-I). Shaft more or less cylindrical, slightly fl attened, 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).

Remarks
The operculum of Chondrocyclus amathole sp. nov. is unique among all other Chondrocyclus species in its fl atness, 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. alabastris and formed a well-supported clade (the Southern-Eastern Cape clade) with C. herberti sp. nov. and C. silvicolus sp. nov. (Fig. 1). The periostracal fl anges of C. amathole sp. nov. were broad and quadrangular while European Journal of Taxonomy 569: 1-92 (2019) in the other three species in its clade, the fl anges 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 fl anges 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-fl oor 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 signifi cant and complex impact on the phylogeography of low-vagility organisms (Daniels et al. 2017).
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. alabastris on 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 classifi cation 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 identifi ed 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 Cape clade (i.e., C. alabastris and C. amathole sp. nov.) is mirrored by rhytidid snails (Herbert & Moussalli 2010, 2016. Connolly, 1929 Figs 15, 16A

Diagnosis
Shell small, depressed, lenticular; periostracum with dense axial costae developing fi ve rows of fl anges on last whorl: just below suture, at periphery, around umbilicus and two weaker rows on either side of periphery; operculum duplex, exterior portion shallowly concave with step-shaped multispiral lamella terminating in a long solid fringe refl exed over peristome; radula with three large cusps on second lateral tooth, cusps of rachidian, fi rst and second lateral teeth fairly uniform in size; penis fl attened dorsoventrally and laterally expanded on left side from about midway down the shaft, intromittent organ relatively long.

Etymology
The specifi c name is derived from the Latin 'tres', meaning 'three', and 'fi mbriae', meaning 'fringe', with reference to the three spiral cords of compacted fl ange-like bristles on the last whorl.  (2019) data as for preceding; 13 Jan. 2010; ELM D17003 • 3 specimens; same collection data as for preceding; ELM W03661.
Description SHELL ( Fig. 15A-C). Small, depressed, lenticular, adult diameter 3.9-4.37 mm (4.37-5.32 mm Entumeni population), height 1.95-2.66 mm (2.39-2.86 mm Entumeni population), diameter:height 1.65-2.0 (1.62-2.02 mm Entumeni population) (n = 5 C. trifi mbriatus from type locality; n = 6 Entumeni population). Spire not much raised, protoconch sub-mammillate (Connolly 1929). Embryonic shell two whorls, microscopically malleate, junction between embryonic shell and teleoconch evident with development of costae on teleoconch. Teleoconch comprising 2.75 whorls, convex, rapidly increasing, suture deeply impressed. Aperture circular, last whorl descending near aperture, peristome simple, continuous and free. Umbilicus wide and deep, exposing all the whorls. Periostracum glossy, honeybrown and lacquer-like with dense lamellate axial costae at regular intervals, approx. 150-155 (n = 4) on last whorl in specimens from the type locality, but varying between populations (see Remarks below), expanded into quadrangularly-shaped fl anges at periphery, immediately below the suture and around the umbilicus, the row at the periphery the strongest, and with two less prominent rows of lower-standing fl anges on either side of periphery. Shell translucent, glossy, corneous yellow-brown when fresh. LIVING ANIMAL. Creamy white with slight pigmentation on tentacles. OPERCULUM (Fig. 15D, F). Duplex, lamella of outer multispiral portion with 4.25 whorls, step-shaped; on outer surface of lamellar blade a long, solid fringe curves upwards and then outwards forming a groove between fringe and lamellar blade, spanned by a very loose network of bristles; a very short solid horizontal fringe emanates just below main fringe; upper edge of lamella thin and projects above fringe in side view; outer surface of lamellar blade tuberculate at high magnifi cation. RADULA (Fig. 15E). Rachidian with fi ve cusps, approx. equivalent length; fi rst and second lateral teeth similar, each with fi ve cusps, fi rst three cusps approx. equivalent in size, the fourth very small and the fi fth (from centre) vestigial; cusps increase in size very slightly from central tooth outwards, but there is not a large difference in size between cusps. PENIS ( Fig. 15G). fl attened dorsoventrally and laterally expanded on left side from about midway down the shaft, with numerous annular rugae, smooth distal end narrows, intromittent organ relatively long.

Distribution and habitat
Originally known only from Karkloof River Valley, downstream of Karkloof Falls, but the species appears to have disappeared from the Karkloof vicinity (Fig. 16A). Specimens recently discovered at Entumeni and Nkandla forests are considered here to be C. trifi mbriatus based on comparison with type material.
No habitat data available for original specimens, but indigenous Eastern Mistbelt Forest (von Maltitz et al. 2003) occurs in the Karkloof vicinity. Recently collected specimens occur in Scarp forests (Entumeni) and Mistbelt/Scarp (Nkandla) in leaf litter.

Remarks
There are morphological differences between populations from different localities. Specimens from Entumeni ( Fig. 15B-C) lack the spiral row of fl anges just below the suture and the weak row below the periphery. Specimens from Nkandla also bear fewer spiral cords than C. trifi mbriatus: there is a spiral cord just below suture and one between periphery and umbilicus but the weak cords on either side of periphery are absent. Although not many specimens were available to measure (four from Karkloof, COLE M.L., Revision of cyclophorid snails Chondrocyclus s.l.  (2019) six from Entumeni and two from Nkandla), D:H ratio is similar throughout but the axial costae on last whorl are more dense in Entumeni specimens (approx. 200) and less dense in Nkandla specimens (117) compared to 150-155 in Karkloof specimens. The opercula of specimens from Entumeni and Nkandla are similar to the operculum of C. trifi mbriatus from the type locality (Fig. 15D). The radulae of all populations agree with the descriptions and fi gure of Connolly (1929Connolly ( , 1939. Few specimens were available for study since they seemed to be very rare in the two forests where they were found, and more sampling from these and other forests in north-central KwaZulu-Natal are required to resolve more conclusively whether C. trifi mbriatus occurs throughout the region or whether there may be additional narrow-range species. The only localities represented in the molecular study were Entumeni and Ngome (considered to contain a separate species, C. pulcherrimus sp. nov.). The highly fragmented forests of north-central Kwazulu-Natal contain many examples of species, in unrelated taxa, with extremely narrow distributions (e.g., Huber 2003;Herbert & Kilburn 2004;Tilbury & Tolley 2009) suggesting that the observed morphological differences between populations may indicate undescribed species.

Diagnosis
Shell small, depressed, lenticular; protoconch not mammillate; periostracum with dense, erect, transverse costae, expanded to form fi ve spiral rows of raised fl anges: a paddle-shaped row at periphery, a row of well-developed quadrangularly-shaped fl anges around umbilicus and above periphery and a row of lower semi-circular fl anges below suture; operculum duplex, lamella forming outer portion of operculum step shaped and terminating in a long solid fringe, portion above fringe consists of a lattice-like network of interwoven bristles some fused to fringe, projecting well above fringe and sloping inward towards centre; umbilicus wide and deep, exposing all the whorls; radula with three large cusps on second lateral tooth, cusps of rachidian, fi rst and second lateral teeth fairly uniform in size; penis fl attened dorsoventrally and laterally expanded about midway down the shaft, intromittent organ relatively long.

Etymology
Named for the distribution of the species, centered around the Pondoland region.  Fig. 17A-C). Small, depressed, lenticular, adult diameter 4.59-6.18 mm, height 2.42-3.42 mm, diameter:height 1.59-2.00 (n = 19). Spire not much raised, protoconch not mammillate. Embryonic shell just over 1.5 whorls, microscopically malleate, junction between embryonic shell and teleoconch not particularly distinct, initially costae weak, becoming well developed after about a quarter of a whorl (Fig. 17D). Teleoconch comprising 2.75 whorls, moderately convex, rapidly increasing, suture deeply impressed. Aperture circular, last whorl descending near aperture, peristome simple, continuous and free. Umbilicus wide and deep, exposing all the whorls. Periostracum glossy, honey-brown and European Journal of Taxonomy 569: 1-92 (2019) lacquer-like with dense lamellate costae at regular intervals, approx. 154-184 (n = 14) on last whorl, expanded into fi ve spiral rows of fl anges: a paddle-shaped row at periphery, a row of well-developed quadrangularly-shaped fl anges around umbilicus and above periphery and a row of lower semi-circular fl anges below suture; fl anges bear numerous axial riblets visible at very high magnifi cation (Fig. 17E); intervals between costae with microscopic axial threads. Shell translucent, glossy, corneous yellowbrown when fresh. LIVING ANIMAL. Creamy white with slight pigmentation on tentacles. OPERCULUM ( Fig. 17G-H). Duplex, outer portion consists of multispiral lamella with 4.5-5 whorls; lamellar blade high, stepped, long fringe emanates near top of vertical portion of blade; blade projects high above fringe and slopes inward towards centre, upper portion of lamellar blade consists of latticelike network of interwoven bristles, some bristles connected to main fringe so there is no groove between fringe and lamellar blade (Fig. 17G); at the growing edge the top portion of lamellar blade is more or less square, not sloping, becoming arrow shaped below fringe (Fig. 17H). Outer lamellar fringe refl exed over peristome preventing animal withdrawing into shell. RADULA (Fig. 17F). Rachidian with fi ve cusps, central one slightly longer; fi rst and second laterals each with three large cusps, a smaller fourth and a vestigial fi fth, cusps increase in size very slightly from central tooth outwards, but there is not a large difference in size between cusps. PENIS ( Fig. 17I-J). Expanded towards distal end, intromittent organ relatively long.

Distribution and habitat
Known primarily from near the coast between Xora in Transkei and Umtamvuna in southern Kwazulu-Natal, and also recorded inland in the Port Shepstone area (ca 200 m a.s.l.) (Fig. 16A). (The Transkei is the area of the Eastern Cape Province between the Kei and Umtamvuna Rivers.) Indigenous Scarp Forest of the Pondoland Gorge and Transkei Coastal Scarp subtypes (von Maltitz et al. 2003), adjacent to rivers, in leaf litter. Scarp forests are a forest type occurring on south-and east-facing hills and gorges of the fi rst plateau escarpment (300-1100 m) and unique to the east of South Africa.

Remarks
Morphologically, C. pondoensis sp. nov. resembles C. trifi mbriatus in features of the periostracum, operculum and radula. The position of spiral rows of fl anges is equivalent in C. pondoensis sp. nov. and C. trifi mbriatus specimens from the type locality. The radulae have teeth with cusps relatively uniform in size. The operculum of C. pondoensis sp. nov. is fl atter than that of coastal species of Chondrocyclus. However, it is distinguished from that of C. trifi mbriatus by the outer lamellar blade projecting high above the fringe and sloping towards the centre and a well-developed lattice of interwoven bristles joining the lamellar blade to the fringe (Fig. 17G). Other species bearing a lattice of interwoven bristles at the top edge of lamellar blade, fused with the main fringe are the coastal C.putealis Conolly, 1939, C. bathrolophodes Conolly, 1929 and C. cooperae sp. nov. but these species have a relatively deep operculum.
In the molecular study, C. pondoensis sp. nov. did not appear related to C. trifi mbriatus or to any other lineages (Cole et al. 2019).
In coastal areas of Transkei where C. pondoensis sp. nov. is sympatric with C. putealis and C. cooperae sp. nov. specimens appeared to be scarce and patchily distributed, while at localities in southern KwaZulu-Natal (Umtamvuna Nature Reserve and Hlokohloko valley inland of Port Shepstone) where C. pondoensis sp. nov. was the only Chondrocyclus species recorded, specimens appeared to be common.

Diagnosis
Shell small, depressed, lenticular; protoconch not mammillate; periostracum with widely spaced axial costae developing at the periphery broad rounded fl anges, semi-circular in shape; operculum duplex, exterior portion very shallowly concave to almost fl at, with step-shaped multispiral lamella terminating in a short fringe of fused bristles.

Etymology
Named for Div DeVilliers whose unwaivering dedication to nature conservation in the Eastern Cape province, and the Transkei in particular, has helped preserve the remaining forests and their biota. Description SHELL ( Fig. 18A-C). Small, depressed, lenticular, adult diameter 5.01-6.01 mm, height 2.59-3.17 mm, diameter:height 1.70-2.16 (n = 16). Spire not much raised, apex almost fl at. Embryonic shell (Fig. 18D-E) nearly 2.25 whorls, microscopically malleate, junction between embryonic shell and teleoconch evident with development of costae on teleoconch. Teleoconch comprising 2.75 whorls, convex, rapidly increasing, suture deeply impressed. Aperture circular, last whorl descending near aperture, peristome simple, continuous and free. Umbilicus wide and deep, exposing all the whorls. Periostracum glossy, COLE M.L., Revision of cyclophorid snails Chondrocyclus s.l.  (2019) honey-brown and lacquer-like with widely spaced lamellate axial costae at regular intervals, 65-77 (n = 4) on last whorl, expanded into a single row of semi-circular fl anges at periphery (Fig. 18D). Shell translucent, glossy, corneous yellow-brown when fresh. OPERCULUM ( Fig. 18F-G). Duplex, outer portion slightly concave; lamella of outer multispiral portion with 4.5 whorls, step-shaped; upper edge of lamella thin and projects slightly above fringe. Radula (Fig. 18H). Rachidian with fi ve cusps, central cusp approx. twice length of outer two on each side; fi rst and second lateral teeth similar but second is slightly larger, each with four cusps and a vestigial fi fth; fi rst three cusps gradually increasing in size from centre outwards and fourth very small. PENIS ( Fig. 18I-J). Shaft dorsoventrally fl attened, with lateral expansions of shaft on both sides towards distal end but more prominent on left, with numerous annular rugae, smooth distal end narrower, intromittent organ relatively long.

Distribution and habitat
Known only from two forests, Nqadu approx. 20 km north of Mthatha and Bele to the north-west. Transkei Mistbelt Forest (von Maltitz et al. 2003), in leaf litter (Fig. 16A).

Remarks
Chondrocyclus devilliersi sp. nov. has several unique morphological features and in the molecular analyses it was not closely related to other species of Chondrocyclus in the Eastern Clade. Although the majority of adult specimens are worn even when collected alive and display few features of the periostracum, the fresh juveniles reveal that there is only one row of expanded fl anges around the periphery of the shell (Fig. 18D) and not multiple spiral rows on the body whorl as in other members of the Eastern clade. This resembles the pattern in C. convexiusculus and species in the Southern-Eastern Cape clade.
The radula resembles that of the coastal taxa C. bathrolophodes, C. putealis and C. cooperae sp. nov. with varying sizes of cusps, while the fl at operculum is quite unlike the deeply concave operculum of these species. Diagnosis Shell very small, very depressed, discoidal; periostracum with very dense axial costae developing at the periphery a row of quadrangularly-shaped fl anges and several spiral rows of semicircular fl anges above and below this; operculum duplex, exterior saucer-shaped portion with step-shaped multispiral lamella terminating in a long solid fringe refl exed over peristome; radula with three large cusps on second lateral tooth, cusps on rachidian uniform in size; penis fl attened dorsoventrally and laterally expanded on left side from about midway down the shaft.

Etymology
The specifi c name is derived from the Latin 'pulcherrimus', meaning 'most beautiful', with reference to the ornate periostracum with very dense axial lamellae and a larger number of spiral rows of fl anges than any other species.  (Fig. 19A-C). Small, very depressed, discoidal, adult diameter 3.50-4.85 mm, height 1.38-2.56 mm, diameter:height 1.87-2.54 (n = 22). Spire almost fl at with sometimes only protoconch projecting (Fig. 19A), suture deeply impressed. Embryonic shell (Fig. 19D) just over two whorls, sculptured with pock marks, junction between embryonic shell and teleoconch not particularly distinct, with weak costae at fi rst, becoming stronger. Teleoconch comprising two whorls, rapidly increasing, suture deeply impressed. Aperture circular, last whorl descending near aperture, peristome simple, continuous and free. Umbilicus very wide, exposing all the whorls (Fig. 19C). Periostracum glossy, light yellowishbrown and lacquer-like, with very dense lamellate axial costae at regular intervals, approx. 300 on last whorl (300 and 340 in Fig. 19B and A, respectively), developing at the periphery quarangularly-shaped fl anges, their expanded distal ends fused to form a solid fringe (Fig. 19E); above periphery are four-fi ve spiral rows of fl anges and six-seven rows below it spiraling into umbilicus, height of fl anges forming spiral rows varies; row of fl anges below suture forms a channel (see Fig. 19B,D); intervals between costae smooth. Shell pale, corneous and translucent when fresh. OPERCULUM ( Fig. 19G-I). Duplex, outer portion consists of multispiral lamella with about 4.5 whorls, each step-shaped; at the growing edge the top of lamellar blade slopes down at an angle then curves under the step formed by the previous whorl and is thinner and more or less perpendicular where it is attached to the disc; very long fringe of fused bristles emanates from near base of vertical portion of step and curves upwards and then outwards forming a deep furrow between fringe and lamellar blade, spanned by a few bristles; a very short solid horizontal fringe emanates just below main fringe; top edge of lamella projects above level of fringe; fringe of each whorl fused to lamella of following whorl; outermost lamellar fringe very long and refl exed over peristome. RADULA (Fig. 19F). Rachidian with fi ve cusps of approx. equivalent length; fi rst lateral tooth with three cusps of approx. equivalent length and a very small fourth; second lateral tooth with three cusps of approx. equivalent length and two vestigial outer cusps; teeth do not differ much in size. PENIS (Fig. 19J-K). fl attened dorsoventrally and laterally expanded on left side from about midway down the shaft, with numerous annular rugae, smooth distal end occupies about one quarter of shaft.

Remarks
The very dense costae of the periostracum of C. pulcherrimus sp. nov. and large number of spiral rows of fl anges, as well as the fl at spire, immediately distinguish this species from all other species of Chondrocyclus. Chondrocyclus pulcherrimus sp. nov. resembles C. trifi mbriatus in features of the operculum, radula, penis and the rough texture of the protoconch. The operculum is similar to that of C. trifi mbriatus, but the fringe emanates from near the bottom of the vertical portion of the stepped lamellar blade forming a deep groove while in C. trifi mbriatus it emanates from near the top of the blade and there is a very shallow groove between the fringe and lamella.
The Ngome Forest is 3360 ha and much of the forest is protected in the Ntendeka Wilderness Area. Chondrocyclus pulcherrimus sp. nov. is locally very common unlike C. trifi mbriatus which appears to be rare in all populations sampled. Connolly, 1929 Figs 16B, 20-21 Chondrocyclus bathrolophodes Connolly, 1929: 239, pl

Diagnosis
Shell small, depressed, lenticular; periostracum with dense axial costae expanded into three spiral rows of fl anges on last whorl: at periphery, around umbilicus and below suture; costae and fl anges bear numerous tiny vertical ridges; operculum duplex, exterior portion with cup-shaped multispiral lamella with horizontal shelf of interwoven bristles spiralling up on inside of cup, fused to a very long, loose fringe refl exed over peristome; radula with three large cusps on second lateral tooth; penis fl attened dorsoventrally with lateral expansions of shaft towards distal end, more prominent on left side. Description SHELL (Fig. 20A-D). Small, depressed, lenticular, adult diameter 5.32-6.16 mm, height 2.76-3.45 mm, diameter:height 1.73-2.00 (n = 25). Spire depressed, each whorl just rising above the next, apex almost fl at (Fig. 20A-B). Embryonic shell (Fig. 21A) 1.75 whorls, microscopically malleate, roughest in centre, junction between embryonic shell and teleoconch evident with appearance of a few weak axial costae, but not sharply demarcated. Teleoconch comprising 2.75 whorls, moderately convex, rapidly increasing, suture deeply impressed. Aperture circular, last whorl descending near aperture, peristome simple, continuous and free. Umbilicus wide and deep, exposing all the whorls (Fig. 20D). Periostracum glossy, honey-brown and lacquer-like with dense lamellate costae at regular intervals, 92-127 (n = 24) on last whorl, expanded into quadrangularly-shaped fl anges at periphery, forming a distinct peripheral cord, as well as a row of shorter semicircular fl anges below the suture and around the umbilicus (Fig. 20A-D); fl anges bear numerous axial riblets visible at very high magnifi cation (Fig. 21B); intervals between costae with microscopic axial threads (Fig. 20B). Shell translucent honey-brown when fresh. LIVING ANIMAL. Head, tentacles and snout dark grey, underside of foot creamy white. OPERCULUM (Fig. 21C-E). Duplex, outer multispiral portion with 5.5 whorls forming a strongly concave cup, upper edge of lamellar blade thickened, forming a horizontal shelf of interwoven bristles spiralling up the inside of the cup; in the outermost whorl the latter is loosely connected to a very long fringe refl exed over peristome and preventing operculum being withdrawn into shell; in earlier whorls this fringe is not visible and appears fused with the lamella; surface of lamella of last whorl tuberculate at high magnifi cation. RADULA (Fig. 21F). Rachidian with fi ve cusps, central cusp approx. twice length of outer two on each side; fi rst and second lateral teeth similar but second is larger, each with four cusps and a vestigial fi fth; fi rst three cusps gradually increasing in size from centre outwards and fourth very small. PENIS (Fig. 21G-H). Shaft dorsoventrally fl attened, with lateral expansions of shaft on both sides towards distal end but more prominent on left, with numerous annular rugae, distal end bulbous and smooth, intromittent organ exserted, but not elongated. COLE M.L., Revision of cyclophorid snails Chondrocyclus s.l.

Distribution and habitat
Endemic to Eastern Cape coast with a disjunct distribution, occurring between Port Elizabeth and Great Fish River (recorded up to 9 km inland) and between Great Kei and Kobonqaba River (recorded up to 6 km inland) (Fig. 16B).

Remarks
Chondrocyclus bathrolophodes is similar to C. putealis (differences are discussed under the latter species) in periostracal ornamentation, operculum, radula and penis.
Specimens recorded in the vicinity of the Kei and Kobonqaba Rivers would have been diffi cult to identify on morphological grounds alone, but the molecular phylogeny shows unambiguously that the specimens sequenced are C. bathrolophodes (Fig. 1). The species was previously thought to occur only in the Port Elizabeth-Albany area, but is shown to have a disjunct distribution. In very close proximity to Kobonqaba to the east, in the Nxaxo forest, Chondrocyclus specimens are morphologically very similar to C. bathrolophodes and C. putealis, but are neither of these species on molecular grounds (Cole et al. 2019; Fig. 1). Chondrocyclus bathrolophodes exhibits very little genetic diversity at the end of a relatively long branch, suggesting recent geographical expansion and severe pruning of ancestral diversity by extinctions. All the coastal taxa of the Eastern clade show complex distribution patterns of sympatry, disjunctions and turnover within sharp contact zones. These could be the result of dynamic changes in vegetation over time, with forest contraction and expansion due to climatic oscillations (Partridge 1993;Partridge et al. 1999). These processes could have caused repeated periods of isolation of populations in shrinking refuges, followed by dispersal and contact. Connolly, 1939 Figs 16B, 22-23 Chondrocyclus putealis Connolly, 1939: 538, pl

Diagnosis
Shell small, depressed, lenticular; periostracum with dense axial costae expanded into three spiral rows of fl anges on last whorl. at periphery around umbilicus and below suture; operculum duplex, multispiral lamella of exterior portion deeply concave, with horizontal shelf of interwoven bristles spiralling up on inside of lamella and connected to a very long, loose fringe refl exed over peristome; radula with three large cusps on second lateral tooth; penis fl attened dorsoventrally with lateral expansions of shaft towards distal end, more prominent on left side.

Etymology
The specifi c name is derived from the Latin 'putealis', meaning 'of or relating to a well', with reference to the deep well-like operculum.

Type material examined
Syntypes SOUTH AFRICA -KwaZulu-Natal • 4 specimens; Natal, Southport;NHMUK 1937NHMUK .12.30.5087-1937.5090. (Fig. 22A (2019) Description SHELL (Fig. 22A-D). Small, depressed, lenticular, adult diameter 4.91-5.38 mm, height 2.49-3.26 mm, diameter:height 1.61-2.00 (n = 20). Spire not much raised, apex almost fl at (Fig. 22A, C). Embryonic shell (Fig. 23A) approx. 1.75 whorls, microscopically malleate, roughest in centre, junction between embryonic shell and teleoconch evident with appearance of a few weak axial costae, but not sharply demarcated. Teleoconch comprising 2.5-2.75 whorls, moderately convex, rapidly increasing, suture deeply impressed. Aperture circular, last whorl descending strongly near aperture, peristome simple, continuous and free. Umbilicus wid and deep, exposing all the whorls (Fig. 22D). Periostracum glossy, honey-brown and lacquer-like with dense lamellate costae at regular intervals, 107-150 (n = 15) on last whorl, expanded into paddle-shaped fl anges at periphery and umbilicus, as well as a row of shorter broadly rounded lamellae below suture (Fig. 22B-D); fl anges bear axial riblets visible at very high magnifi cation (Fig. 23B); intervals between costae with microscopic axial threads. OPERCULUM ( Fig. 23D-E, G-H). Duplex, outer multispiral portion with approx. fi ve whorls; lamellar blade high and steep sided, resulting in a deeply concave outer portion with more-or-less perpendicular sides towards the top and curving inwards towards the bottom; upper edge of lamellar blade thickened, forming a horizontal shelf of interwoven bristles which forms a spiral staircase on the inside of the lamellar blade; in the outermost whorl the latter is connected to a very long fringe refl exed over peristome and preventing operculum being withdrawn into shell. In earlier whorls this fringe is not visible and appears fused with the lamella; surface of lamella of last whorl tuberculate at high magnifi cation (Fig. 23H). Shell translucent, glossy, honey-brown when fresh. RADULA (Fig. 23C). Rachidian with fi ve cusps, central one approx. twice length of outer two on each side, the latter four approx. equivalent length; fi rst and second lateral teeth each with three large cusps, a smaller fourth and a vestigial fi fth, second lateral tooth is larger. PENIS (Fig. 23F, I). Shaft dorsoventrally fl attened, with lateral expansions towards the distal end on both sides but more prominent on left, with numerous annular rugae, distal end bulbous and smooth.

Distribution and habitat
Endemic to a narrow range primarily near the coast in southern KwaZulu-Natal and Pondoland (the northeastern region of the Eastern Cape province), between Mtentu in northern Transkei and Southport in southern Kwazulu-Natal, but also recorded inland in the Port Shepstone area (ca 200 m a.s.l.) (Fig. 16B).

Remarks
Morphological differences between C. putealis and C. bathrolophodes are slight. The shell of C. putealis resembles C. bathrolophodes in shape but attains slightly smaller size. The position of spiral rows of fl anges is similar in the two species, but costae are slightly more dense on average in C. putealis and the peripheral and umbilical cords stronger than those of C. bathrolophodes. The opercula of the two species are very similar although the operculum of C. putealis is deeper with perpendicular sides near the top and curving inward near the disc, while the lamellar blade of C. bathrolophodes widens evenly towards the top. Both species occupy a relatively narrow range, separated geographically by a wide intervening distance in which C. cooperae sp. nov. (below) and C. pondoensis sp. nov. occur. On morphological grounds alone, it was not clear whether C. putealis and C. bathrolophodes should indeed be considered distinct species, in spite of subtle differences. In the molecular analyses C. pondoensis sp. nov. is nested within this group (Fig. 1), adding weight to the evidence to treat these lineages as distinct species.
Chondrocyclus putealis appears to be a very rare species and there are only a few localities where recent specimens could be found and it was patchily distributed at these localities. Chondrocyclus putealis is replaced by the closely related C. cooperae sp. nov. westwards along the coast, while C. pondoensis sp. nov. also extends westwards from Pondoland and is sympatric with both these species. This region of the north-eastern coast of the Eastern Cape is an important centre of cladogenic events in rhytidid molluscs (Moussalli et al. 2009

Diagnosis
Shell small, depressed, lenticular; periostracum with dense axial costae expanded into seven spiral rows of fl anges on last whorl: one at periphery, and three each between periphery and suture and between periphery and umbilicus; costae and fl anges bear numerous tiny vertical ridges; operculum duplex, exterior concave portion shaped like a wide cup or bowl, multispiral lamella with horizontal shelf of interwoven bristles spiralling up on inside of bowl, fused to a very long, loose fringe refl exed over peristome; radula with three large cusps on second lateral tooth; penis fl attened dorsoventrally with prominent lateral expansion on left of shaft towards distal end, and with a characteristic long intromittent organ.

Etymology
Named for Janna Cooper, who helped collect specimens of this species and in recognition of her intimate knowledge of the Transkei coast.

Type material examined
Holotype SOUTH AFRICA -Eastern Cape • Umtiza Nature Reserve, east side of Buffalo Pass, indigenous scarp forest; 33.0144° S: 27.8081° E; 21 Apr. 2006; M. Bursey leg.; in leaf litter; NMSA W9271/T3074. (Fig. 10M-N) Paratypes SOUTH AFRICA -Eastern Cape • 2 specimens; same collection data as for holotype; ELM D14918/ T122 • 15 specimens; same collection data as for preceding; M. Cole leg., 18 Mar. 2011; ELM D16936/ T120 • 5 specimens; same collection data as for preceding; ELM W3623/T121 • 4 specimens; same collection data as for preceding; NHMUK 20120280 • 2 specimens; same collection data as for preceding; NMSA W9275/T3076 • 1 specimen; same collection data as for preceding; NMW.Z.2012.065.00009 • 3 specimens; same collection data as for preceding; RMNH MOL.330502 • 3 specimens; same collection data as for preceding; 20 May 2011; M. Cole  Description SHELL ( Fig. 24A-C). Small, depressed, lenticular, adult diameter 4.51-6.51 mm, height 2.25-4.01 mm, diameter:height 1.44-2.32 (n = 66, measured in several populations throughout the range). Spire not much raised, apex almost fl at (Fig. 24A). Embryonic shell (Fig. 23D) just under 1.75 whorls, microscopically malleate, roughest in centre, junction between embryonic shell and teleoconch evident with appearance of a few weak costae, but not sharply demarcated. Teleoconch comprising 2.75 whorls, moderately convex, rapidly increasing, suture deeply impressed. Aperture circular, last whorl descending strongly near aperture, peristome simple, continuous and free. Umbilicus wide and deep, exposing all the whorls (Fig. 24C). Periostracum glossy, honey-brown and lacquer-like with dense lamellate axial costae at regular intervals, density varying between populations, 105-179 (n = 27, counted in three separate populations), expanded into tall quadrangularly-shaped fl anges at periphery, and six additional spiral rows of lowerstanding fl anges on body whorl, three between suture and periphery and three rows between periphery and umbilicus, the row closest to umbilicus weaker than other rows (Fig. 24A-C); fl anges and portion of costae near umbilicus bear axial ridges visible at very high magnifi cation (Fig. 24E); intervals between costae with 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. 24G-H). Duplex, outer multispiral portion with 5.5 whorls forming a widely cupolaeform cup or bowl, upper edge of lamellar blade thickened, forming a horizontal shelf of interwoven bristles in a lattice-like pattern; in the outermost whorl the latter is continuous with a very long fringe refl exed over peristome and preventing operculum being withdrawn into shell; in earlier whorls this fringe is not visible and appears fused with the lamella; growing edge of lamella arrow shaped. Surface of lamella of last whorl tuberculate at high magnifi cation. COLE M.L., Revision of cyclophorid snails Chondrocyclus s.l. RADULA (Fig. 24F). Rachidian with fi ve cusps, central one approx. twice length of outer two on each side; fi rst and second lateral teeth similar but second is larger, each with four cusps and a vestigial fi fth; fi rst three cusps gradually increasing in size from centre outwards and fourth very small. PENIS (Fig. 24I-J). Shaft dorsoventrally fl attened, with large lateral expansion on left side, with numerous annular rugae, distal end smooth and not bulbous, intromittent organ very long.

Distribution and habitat
Endemic to Eastern Cape from East London to Mbotyi, primarily near the coast. The range of C. cooperae sp. nov. overlaps with that of C. bathrolophodes in the west, and C. pondoensis sp. nov. in the east (Fig.  16B).

Remarks
Chondrocyclus cooperae sp. nov. resembles C. bathrolophodes and C. putealis in general shape, operculum and radula, but differs consistently in the larger number of spiral rows of periostracal fl anges. The very long intromittent organ of the penis is unmistakable and immediately distinguishes the species.

Diagnosis
Shell small, very depressed, discoidal; periostracum with axial costae producing spiral rows of simple, robust hairs concentrated at and on either side of periphery; operculum duplex, exterior portion very shallowly concave, with thickened ridge on multispiral lamella from which emanates a fairly long solid fringe and a very short fringe below this; radula with three large cusps on second lateral tooth.

Etymology
Named after the Langeberg mountain range, part of the Cape Fold Mountains.

Type material examined
Holotype SOUTH AFRICA -Western Cape • Langeberge foothills, Pat Busch Nature Reserve, Karin Trail, riverine fynbos; 33.7551° S, 19.9947° E; 450 m a.s.l.; 7 Aug. 2014; M. Cole leg.; in leaf-litter beneath bushes; NMSA P0642/T4159. (Fig. 10O, R) Paratypes SOUTH AFRICA -Western Cape • 21 specimens; same collection data as for holotype; ELM D17981/ T98 • 1 specimen; same collection data as for holotype; ELM W3899/T99 • 14 specimens; same collection data as for holotype; 10 Oct. 2007; D. Herbert (Fig. 25A-C). Small, very depressed, discoidal, adult diameter 3.63-5.76 mm, height 1.42-2.76 mm, diameter:height 1.79-2.85 (n = 67, measured in four different populations; Table 3). Spire almost fl at (Fig. 25A), sometimes concave, usually with only the mammillate, tilted protoconch projecting. Embryonic shell (Fig. 25D) approx. 2-2.25 whorls, microscopically malleate, junction between embryonic shell and teleoconch evident with development of widely spaced axial costae on teleoconch. Teleoconch comprising 2.25 whorls, very rapidly increasing, convex, suture deeply impressed. Aperture circular, last whorl descending steeply nearing aperture, peristome simple, continuous and free. Umbilicus very wide, exposing all the whorls (Fig. 25C). Periostracum glossy and lacquer-like with lamellate axial costae at regular intervals, 47-63 (n = 14) on last whorl in Grootvadersbosch population but varies between populations (Table 3), which produce six spiral rows of simple, very long and robust hairs around the periphery; intervals between costae with six-eight microscopic axial threads. Shell translucent reddish brown, honey brown or yellowish-white when fresh. Living animal. Variable in colour between populations from creamy white with light brown pigmentation on tentacles to almost black, underside of foot creamy white. OPERCULUM (Fig. 25F, I). Duplex and shallowly concave; multispiral lamella of outer portion with fi ve low whorls, thickened horizontal ridge near base of lamellar blade runs parallel to disc surface, a long fringe of fused bristles and a second very short fringe below it emanate from this ridge; main fringe grows upwards (i.e., parallel to lamellar blade) and then downwards, leaving a deep, wide groove between fringe and blade of lamella; lamellar blade projects above level of fringe and is very thin; fringe of each whorl fused to lamellar blade of next whorl; fringe of outermost whorl overlaps disc slightly and is refl exed over peristome in life although operculum is retractile. RADULA (Fig. 25E). Rachidian with fi ve cusps, middle one longer than 2 cusps on either side of it; fi rst and second lateral teeth with fi ve cusps (5 th sometimes vestigial), the third cusp from centre the largest. PENIS (Fig. 25G-H). Shaft more or less cylindrical and slightly fl attened dorsoventrally, distal half slightly expanded on left side, numerous annular rugae, distal end smooth but not bulbous, intromittent organ short.

Distribution and habitat
Western Cape, evidently endemic to Langeberg mountain range in Cape Fold Mountain belt, southfacing slopes and on northern side of range in Montagu area (Fig. 5).
Diverse vegetation types: patches of Western Cape Afrotemperate Forest (von Maltitz et al. 2003) and riverine fynbos, in leaf litter.

Remarks
In terms of its hairy periostracum Chondrocyclus langebergensis sp. nov. resembles Afrocyclus isipingoensis gen. et comb. nov., but the molecular analyses placed C. langebergensis sp. nov. and C. kevincolei sp. nov. in a well-supported monophyletic clade, termed the Overberg clade, within the  Chondrocyclus s.s. radiation (Cole et al. 2019; Fig. 1). The major morphological feature distinguishing this clade from A. isipingoensis gen. et comb. nov. is that the second lateral tooth of the radula has three large cusps (Fig. 25E) as opposed to two in A. isipinoensis comb. nov. (Fig. 27E). Differences between C. langebergensis sp. nov. and C. kevincolei sp. nov. are discussed under the latter species.
The Overberg clade and the other taxon in the southwestern Cape, C. convexiusculus, have not been recorded sympatrically although they occur in close proximity inland in the upper Breede River valley and near the coast (Fig. 5). Other taxa also contain distinct clades in either the Hottentots-Holland Mountains or the Overberg which do not occur in the other region (e.g., Gouws et al. 2010;Herbert & Moussalli 2010;Daniels et al. 2013) and the Breede River valley is considered an important barrier to gene fl ow (Weimarck 1941;Linder 2003;

Diagnosis
Shell small, depressed, lenticular; periostracum with axial costae producing spiral rows of simple hairs; spiral grooves on shell corresponding with rows of hairs; operculum duplex, exterior portion very shallowly concave, with step-shaped multispiral lamella terminating in a short fringe with uneven edge, diameter of exterior portion less than diameter of inner disc; radula with three large cusps on second lateral tooth.

Etymology
Named for the author's husband, Kevin Cole, in acknowledgement of his indispensable assistance on fi eldtrips.  Moussalli and D. Stuart-Fox leg.; in leaf-litter; NMSA W3537/T4124. Description SHELL (Fig. 26A-F). Small, depressed, lenticular, adult diameter 3.59-5.34 mm, height 1.59-2.84 mm, diameter:height 1.68-2.37 (n = 87 from 4 populations; variations in dimensions between populations are given in Table 4). Spire little exserted, protoconch large and mammillate (Fig. 26A). Embryonic shell (Fig. 26G) approx. 2.25 whorls, almost smooth but microscopically malleate at tip, junction between embryonic shell and teleoconch evident with development of costae on teleoconch. Teleoconch comprising just over two whorls, very convex, rapidly increasing, suture deeply impressed. Aperture circular, last whorl descending near aperture, peristome simple, continuous and free. Umbilicus wide and deep, exposing all the whorls Fig. 26C, F). Periostracum glossy and lacquer-like with lamellate axial costae at regular intervals on last whorl, 51-75 (n = 22) in Platbos population, but vary between populations (Table 4), which produce spiral rows of simple hairs varying in different populations from approx. 4-14 on last whorl, longest at periphery and shortest around umbilicus; intervals between axial costae with approx. eight microscopic axial threads (Fig. 26H); shell bears spiral grooves corresponding with the rows of hairs (Fig. 26D-F). Shell translucent when fresh; two colour morphs present reddish brown and creamy white. LIVING ANIMAL. Varies in colour between populations from creamy white with slight pigmentation on tentacles to very dark grey (see below). OPERCULUM (Fig. 26J-K). Duplex, shallowly concave; multispiral lamella of outer portion with 4.5 whorls, each step-shaped, growing edge of lamella at angle, perpendicular where it is attached to disc; fringe very short and solid, with a frayed distal edge even in fresh specimens, fringe of each whorl not fused to lamellar blade of following whorl, diameter of outer multispiral portion smaller than diameter of inner disc due to very short fringe; operculum retractile. RADULA (Fig. 26I). Rachidian with fi ve cusps, middle one longer than two cusps on either side of it; fi rst and second lateral teeth each with four cusps and a vestigial 5 th , the third cusp from centre the largest. PENIS (Fig. 26L). Shaft more or less cylindrical and slightly fl attened dorsoventrally, distal half slightly expanded on left side, numerous annular rugae, distal end smooth, intromittent organ short.

Distribution and habitat
Western Cape, Agulhas Plain, between Gansbaai and Bredasdorp. Various forested or bushy habitats in patches of Western Cape Milkwood Forest (part of Southern Coastal forest group) (von Maltitz et al. 2003) with large trees, characterised by white milkwoods (Sideroxylon inerme), white stinkwoods (Celtis africana) and white pear (Apodytes dimidiata E. Mey. ex Arn.), including the southernmost forests in Africa (Platbos and Grootbos), dune scrub at Gansbaai and 'dry bush on hillside' (south-west of Bredasdorp); in leaf-litter (Fig. 5).

Remarks
The distinct spiral grooves on the shell of C. kevincolei sp. nov. (Fig. 26D-F) are unique among species of Chondrocyclus although there are sometimes faint traces of spiral grooves on shells of A. isipingoensis gen. et comb. nov. and C. langebergensis sp. nov. visible only at very high magnifi cation. Chondrocyclus langebergensis sp. nov. differs from C. kevincolei sp. nov. in its lack of prominent spiral grooves on the shell, the spiral rows of long hairs are concentrated around the periphery, the protoconch is more strongly sculptured and the opercula differ. In C. langebergensis sp. nov., the lamellar fringe of each spiral whorl of the operculum emanates from a ridge at the base of the lamellar blade and is fused to the lamellar blade of the following whorl. The terminal fringe is long and overlaps the base when viewed from above (Fig. 25F). In C. kevincolei sp. nov. the fringe emanates from near the top of the lamellar blade and is very short and appears distinct from the following lamellar whorl; the outer saucershaped portion is smaller than the polished base (Fig. 26J). These differences together with molecular evidence, are considered to warrant recognition of at least two distinct species, one in the Langeberg Mountains and the other on the relatively low-lying coastal region of the Agulhas Plain. The Langeberge and Agulhas Plain are separated by the Breede River valley and by a belt of relatively arid vegetation, consisting mainly of Ruens Shale Renosterveld and Agulhas Limestone Fynbos (Rebelo et al. 2006), which developed in the Pliocene (Cowling et al. 2008). Molecular studies in unrelated faunal taxa also provide evidence for separate lineages in the Langeberge and the Agulhas Plain (e.g., Price et al. 2007;Moussalli et al. 2009;Gouws et al. 2010; and for an absence of gene fl ow between these areas (Myburgh & Daniels 2015). Species of Chondrocyclus have not been recorded in the Riviersonderendberge where apparently suitable habitat exists.
There are morphological differences between populations of C. kevincolei sp. nov. (Table 4). The number of spiral rows of hairs is fewer in the Bredasdorp population (only 4-5 as opposed to 8-15 in the Platbos and Grootbos populations). The colour of the living animal is creamy white with only slight pigmentation of the tentacles in the Platbos population and dark grey to almost black at Bredasdorp. There were relatively large genetic divergences between populations (Cole et al. 2019). Their long branch lengths (Fig. 1) suggest isolation in shrinking refuges as arid-adapted vegetation replaced forest (see Tolley et al. 2006Tolley et al. , 2008. Recent radiations of faunal taxa in this region are associated with adaptation to open habitats (Tolley et al. 2006;Herbert & Moussalli 2010). Sea level changes have also had dramatic effects on vegetation of the Agulhas Plain (Linder 2003) and its biota with marine transgressions potentially restricting ancestral populations of Chondrocyclus to refuges at higher elevations and thus driving allopatric divergence or eliminating closely related taxa at lower elevations. Further studies including unsampled populations on the Agulhas Plain and additional markers may shed light on whether separate populations may warrant species status. It was, however, decided to treat these as one species pending further investigation. By contrast, C. langebergensis sp. nov. occurs in relatively stable relictual forest patches in the Langeberge and has undergone little change. Genus Afrocyclus gen. nov. urn:lsid:zoobank.org:act:0C542886-FB1A-4550-BBC2-80BBD28FA01E

Diagnosis
Shell dextral, small, depressed; aperture circular, last whorl descending near aperture, peristome not thickened, continuous and free; umbilicus wide and deep, exposing all the whorls; periostracum glossy and lacquer-like with lamellate axial costae at regular intervals, usually expanded into spiral rows of hairs; operculum duplex, corneous; inner portion consists of a thin disc which grows outwards in a tight spiral, smooth on inside where it attaches to foot; fused to disc on its outer surface is a multispiral, elevated horny blade-like lamella formed by fused bristles; a fringe of fused bristles emanates from outer surface of each lamellar whorl; radula taenioglossate, two large cusps on second lateral tooth; penis lies dorsally immediately behind right tentacle and consists of a stout, muscular, cylindrical shaft, wrinkled along most of its length due to annular rugae, and a terminal, tapering intromittent organ without fl agellum; seminal tube completely enclosed without seminal groove.

Etymology
The specifi c name is derived from 'Africa' and refers to its Afromontane occurrence.
Remarks mtDNA data clarifi ed that Afrocyclus gen. nov. constituted a radiation distinct from that of Chondrocyclus s.s. (Cole et al. 2019). Specimens of Afrocyclus gen. nov. can in most cases be recognised at a glance on morphological grounds, being very small, very depressed (the apex can even be concave), discoid shells with spiral rows of hairs projecting from the axial costae. This morphotype is widely distributed in latitude and altitude, occupying isolated Afromontane forest patches along the Great Escarpment from northern South Africa (Limpopo province, 23° S) to the Eastern Cape (32.7° S) and extending to the coast (Fig. 2). mtDNA data showed that what was previously lumped as Afrocyclus isipingoensis gen. et comb. nov. is a species complex, and within it, external morphology is relatively conserved, making it diffi cult to distinguish taxa by morphology alone. A. exsertus gen. et comb. nov., which occupies a narrow zone on the east coast, is strikingly different morphologically, bearing no hairs and with the axial lamellae only slightly expanded around the periphery. The operculum of the latter also deviates dramatically from the fl at, fragile operculum of most specimens of Afrocyclus gen. nov. and is more robust with a rigid fringe consisting of bunches of short bristles. Phylogenetic relationships within Afrocyclus gen. nov. are not fully resolved and further molecular and morphological data from throughout its range are required to revise this complex.

Diagnosis
Shell very small, very depressed, discoidal; periostracum with axial costae producing spiral rows of simple hairs; operculum very fragile and duplex, exterior portion very shallowly concave, with low multispiral lamella terminating in a solid fringe; rachidian tooth of radula with serrated upper edge, two large cusps on second lateral tooth.

Etymology
Named after the type locality, Isipingo, KwaZulu-Natal, South Africa.

Description
Specimens from Kenneth Stainbank Nature Reserve, Durban, approx. 9 km from the type locality Isipingo, are considered to represent the type in the following description. SHELL (Fig. 27A-C). Very small, very depressed, discoidal, adult diameter 2.24-3.67 mm, height 1.09-1.67 mm, diameter:height 1.9-2.31 (n = 16) Spire almost fl at, sometimes concave, usually with only the weakly mammillate, slightly tilted protoconch projecting (Fig. 27A). Embryonic shell (Fig. 28A) approx. 2.25 whorls, microscopically malleate for approx. two whorls, axial costae begin to develop at about 2.25 whorls. Teleoconch comprising just over two whorls, very rapidly increasing, convex, suture impressed. Aperture circular, last whorl descending steeply nearing aperture, peristome simple, continuous and free. Umbilicus very wide, exposing all the whorls (Fig. 27C). Periostracum glossy and lacquer-like with lamellate well-spaced axial costae at regular intervals, the average number on European Journal of Taxonomy 569: 1-92 (2019) last whorl varying between 55 and 92 (Table 5), which produce 10-12 spiral rows of simple hairs, longest at periphery (four rows) with approx. fi ve-seven rows between periphery and umbilicus, the latter becoming progressively shorter nearer the umbilicus, and one very short row immediately below the suture; intervals between costae usually with six to nine microscopic axial threads (Fig. 27D). Shell translucent when fresh. OPERCULUM (Fig. 27E-F). Very fragile and duplex, outer portion consists of multispiral lamella with fi ve-six whorls, height of lamellar blade very low thus operculum is very shallowly concave to almost fl at, thickened horizontal ridge on lamellar blade just above disc surface; long fringe of fused bristles emanates from this ridge, fused to blade and then curving outwards, leaving no furrow between fringe and vertical portion of blade, fringe of each whorl fused to lamella of following whorl, fringe of outer whorl fairly long and overlaps disc slightly; inner portion of operculum is a thin disc, in some populations side of disc facing body has a prominent tubercle or swelling in centre (Fig. 27F). RADULA (Fig. 27G). Rachidian with fi ve cusps, central one very long, outermost two very small to almost vestigial, cusps set a little distance below 'top' of tooth, top edge serrated; fi rst lateral tooth with four cusps and a vestigial fi fth, third cusp (from centre) largest; second lateral tooth with two large cusps, second cusp (from centre) largest, a third small cusp and a vestigial fourth. PENIS (Fig. 27H). Shaft more or less cylindrical and laterally expanded on left side about midway down the shaft.

Distribution and habitat
This species complex is widely distributed throughout the eastern region of South Africa, from Soutpansberg mountains in Limpopo Province to the Amathole mountains in the Eastern Cape; from the coast to the Drakensberg foothills (1800 m) (Fig. 29).

Remarks
Selected populations from across the range of this lineage were sampled for the molecular study and morphological examination and there is compelling evidence that A. isipingoensis gen. et comb. nov. is a species complex rather than one widespread species. Until further study is undertaken the name A. isipingoensis gen. et comb. nov. is applied to all populations with the exception of the three new species described below. Differences in the density of axial lamellae, the number and position of spiral rows of hairs, the length of the hairs, the density of microscopic axial threads between the lamellae and the strength of protoconch sculpture will need to be evaluated in conjunction with molecular evidence from more comprehensive coverage of the range of the complex.
The  The rachidian tooth of the radula is unusual in having a serrated upper edge and the cusps are set a little distance below the 'top'of tooth. Afrocyclus isipingoensis gen. et comb. nov. (Fig. 27G), A. exsertus gen. et comb. nov. (Fig. 33G), A. potteri gen. et sp. nov. (Fig. 31G) and A. bhaca gen. et sp. nov. (Fig. 32E) showed this feature while the upper edge of the rachidian tooth of A. oxygala gen. et sp. nov. (Fig. 30G)

Diagnosis
Shell very small, depressed, discoidal; periostracum with axial costae producing spiral rows of simple hairs; operculum very fragile and duplex, exterior portion very shallowly concave, with low multispiral lamella terminating in a solid fringe, radula with two large cusps on second lateral tooth and rachidian tooth without serrated upper edge.

Etymology
The specifi c name is derived from the Greek 'oxygala', meaning 'sour milk', a translation of the name of the type locality, Maasstrom.
Aperture circular, last whorl descending steeply nearing aperture, peristome simple, continuous and free. Umbilicus very wide, exposing all the whorls (Fig. 30C). Periostracum glossy and lacquer-like with lamellate axial costae at regular intervals (Fig. 30D), the average number on last whorl varying between 63 and 95 (Table 5), which produce three spiral rows of long, simple hairs at periphery; intervals between costae usually with approx. fi ve microscopic axial threads. Shell translucent when fresh. OPERCULUM (Fig. 30E-F). Very fragile and duplex, outer portion consists of multispiral lamella with fi ve-six whorls, height of lamellar blade very low thus operculum is very shallowly concave to almost fl at, thickened horizontal ridge on lamellar blade just above disc surface; long fringe of fused bristles emanates from this ridge, fused to blade and then curving outwards, leaving no furrow between fringe and vertical portion of blade, fringe of each whorl does not appear to be fused to lamella of following whorl, fringe of outer whorl fairly long and overlaps disc slightly; inner portion of operculum is a thin disc, without a prominent tubercle in centre. RADULA (Fig. 30G). Rachidian with fi ve cusps, central one very long, outermost two relatively short, fi rst lateral tooth with four cusps and a vestigial fi fth, third cusp (from centre) very long; second lateral tooth with two large cusps, second cusp (from centre) larger, a third small cusp and a vestigial fourth. PENIS ( Fig. 30H-I). Relatively short, shaft more or less cylindrical.

Distribution and habitat
Eastern Cape, southern end of Great Escarpment, in Eastern Cape Escarpment Thicket (Hoare et al. 2006) associated with river valleys (Fig. 29).

Remarks
The molecular phylogeny suggests that A. oxygala gen. et sp. nov. belongs to a lineage distinct from all other populations of Afrocyclus gen. nov. sampled ( Fig. 1) but further study is required to resolve phylogenetic relationships between A. oxygala gen. et sp. nov. and other species of Afrocyclus gen. nov.
The species is readily distinguished by having only three rows of long hairs on lamellate costae round the periphery (although in adults the hairs were usually worn off), while A. isipingoensis gen. et comb. nov. has four rows around the periphery. A. oxygala gen. et sp. nov. lacks the additional spiral rows of hairs present in A. isipingoensis gen. et comb. nov.: one row of very short hairs below the suture and approx. fi ve-seven rows of shorter hairs between the periphery and umbilicus. A. oxygala gen. et sp. nov. has a higher spire (and consequently deeper umbilicus) and a more strongly sculptured protoconch (Fig. 28B). The operculum has a smooth inner disc without a tubercle in its centre. The radula more closely resembles those of species in the Southern-Eastern Cape clade (smooth upper edge of rachidian and two large cusps on second lateral tooth) than those of other specimens of Afrocyclus gen. nov. examined. The penis of A. oxygala gen. et sp. nov. is relatively short.
This lineage occurs at the western extremity of the distribution of gen. nov. and the populations at Somerset East and Bedford are on opposite sides of the upper Great Fish River Valley (Fig. 29). Although considered an arid barrier for dispersal of montane taxa (Weimarck 1941;Stuckenberg 1962), this valley has not consistently been a major hindrance to connectivity along this region of the Great Escarpment (Clark et al. 2010) and may have been an area of persistence of thicket during the Last Glacial Maximum (24000 to 18000 years ago) (Potts et al. 2012) and perhaps prior maxima. Diagnosis Shell very small, depressed, discoidal; periostracum with widely-spaced axial costae producing spiral rows of simple hairs; protoconch strongly malleate; operculum very fragile and duplex, exterior portion very shallowly concave, with low multispiral lamella terminating in a solid fringe, radula with two large cusps on second lateral tooth and rachidian tooth with serrated upper edge.  Description SHELL (Fig. 31A-C). Very small, depressed, discoidal, adult diameter 2.17-3.51 mm, height 1.02-1.68 mm, diameter:height 1.7-2.41 (n = 45). Spire low, each whorl just rising above the next, apex mammillate and slightly tilted (Fig. 31A). Embryonic shell (Fig. 28C) just under 2.5 whorls, strongly malleate, junction between embryonic shell and teleoconch evident with development of axial costae on teleoconch. Teleoconch comprising two whorls, very rapidly increasing, convex, suture impressed. Aperture circular, last whorl descending steeply nearing aperture, peristome simple, continuous and free. Umbilicus very wide, exposing all the whorls (Fig. 31C). Periostracum glossy and lacquer-like with well-spaced lamellate axial costae at regular intervals, the number on last whorl varying between 31 and 67 (Table 5), which produce six-ten spiral rows of simple hairs; intervals between costae with approx. 15-20 microscopic axial threads. Shell translucent when fresh. OPERCULUM (Fig. 31F). Very fragile and duplex, outer portion consists of multispiral lamella with 5.25 whorls, height of lamellar blade very low and thus operculum is very shallowly concave to almost fl at, thickened horizontal ridge on lamellar blade just above disc surface; long fringe of fused bristles emanates from this ridge, fused to blade and then curving outwards, leaving no furrow between fringe and vertical portion of blade, fringe of each whorl does not appear to be fused to lamella of following whorl, fringe of outer whorl does not overlap the disc; inner portion of operculum is a thin disc, without a prominent tubercle in centre. RADULA (Fig. 31G). Rachidian with fi ve cusps set a little distance below upper edge of tooth, upper edge slightly serrated, central cusp very long; fi rst lateral tooth with four cusps and a vestigial fi fth, third cusp (from centre) very long; second lateral tooth with two large cusps, second cusp (from centre) larger, a third small cusp and a vestigial fourth. PENIS (Fig. 31H). Shaft more or less cylindrical.

Distribution and habitat
Medium altitude forests of the 'fi rst escarpment' in the interior of the Transkei region of the Eastern Cape and Kei River Valley. Transkei Mistbelt Forest (von Maltitz et al. 2003) (Fig. 29).

Remarks
Populations in the geographically disjunct areas of central Transkei and Kei River Valley formed a very well-supported clade, although populations in the two areas were morphologically distinct (Fig. 31D-E). Specimens from the type locality on the west side of the Kei River Valley are readily distinguished by very widely-spaced axial costae and relatively short periostracal hairs. The protoconchs of specimens from both areas were relatively large (approx. 750 μm) and more strongly mammillate than in any other species of Afrocyclus gen. nov.
It has been suggested that recent migration may have been facilitated by more extensive forest cover along the inland Transkei mistbelt during the Holocene altithermal (Hughes et al. 2005) and perhaps some of the more recent altithermals of the Quaternary climatic oscillations (Partridge 1993;Partridge et al. 1999).    (Fig. 32A-C). Spire almost fl at, sometimes concave, usually with only the weakly mammillate protoconch projecting (Fig. 32A). Embryonic shell (Fig. 28D) just over two whorls, microscopically malleate, junction between embryonic shell and teleoconch evident with development of axial costae on teleoconch. Teleoconch comprising two whorls, very rapidly increasing, convex, suture impressed. Aperture circular, last whorl descending steeply nearing aperture, peristome simple, continuous and free. Umbilicus very wide, exposing all the whorls. Periostracum glossy and lacquer-like with well-spaced lamellate axial costae at regular intervals, the number on last whorl varying between 46 and 65 (Table 5), which produce spiral rows of simple hairs varying from 4-15 on last whorl (4-5 and 14-15 respectively in the two populations examined) (Fig. 32B, D), longest at periphery and shortest around umbilicus; intervals between costae with approx. nine microscopic axial threads. Shell translucent when fresh. OPERCULUM (Fig. 32E). Very fragile and duplex, outer portion consists of multispiral lamella with fi ve whorls, height of lamellar blade very low thus operculum is very shallowly concave to almost fl at, thickened horizontal ridge on lamellar blade just above disc surface; starting just after whorl 3 a long fringe of fused bristles emanates from this ridge, fused to blade and then curving outwards, leaving a furrow between fringe and vertical portion of blade, fringe of each whorl is not fused to lamella of following whorl, fringe of outer whorl extends to edge of disc; inner portion of operculum is a thin disc, without a prominent tubercle in centre.  (Fig. 32F). Rachidian with fi ve cusps set a little distance below upper edge of tooth, upper edge mildly serrated, central cusp very long; fi rst lateral tooth with four cusps and a vestigial fi fth, third cusp (from centre) very long; second lateral tooth with two large cusps, second cusp (from centre) larger, a third small cusp and a vestigial fourth.
PENIS. Not examined.

Distribution and habitat
Medium altitude forests of the 'fi rst escarpment' in the interior of north-eastern Eastern Cape and southern KwaZulu-Natal (Fig. 29).

Remarks
Afrocyclus bhaca gen. et sp. nov. differed from Afrocyclus isipingoensis gen. et comb. nov. by lacking the spiral row of extremely short hairs immediately below the suture and the protoconch was more strongly sculptured, but less so than that of A. potteri gen. et sp. nov. and A. oxygala gen. et sp. nov. The protoconch was smaller than that of A. potteri gen. et sp. nov. (Fig. 28C-D). The two populations examined differed in the arrangement of the spiral rows of hairs: specimens from Ngele had four-fi ve rows concentrated around the periphery (Fig. 32A-B), while specimens from near Mount Frere/Buffalo Nek had 14-15 rows on the last whorl, more or less evenly distributed over the whorl (Fig. 32D). The latter were relatively more depressed than other populations (Table 5). Specimens from Mount Frere are tentatively included in A. bhaca gen. et sp. nov., but may prove to be a separate species upon further study. In the molecular phylogeny they were more closely related to specimens from Ngele than to any other specimens in the Afrocyclus isipingoensis gen. et comb. nov. species complex (Fig. 1), although relatively divergent compared to the genetic variation within the majority of species (see Cole et al. 2019).
Ngele and nearby forests appear to be a hot-spot of narrow range endemism in low-vagility, saproxylic invertebrates including other molluscan taxa (e.g., Gulella claustralis Connolly, 1939) and Onychophora (Daniels et al. 2016). The occurrence of two distinct lineages (A. potteri gen. et sp. nov. and A. bhaca gen. et sp. nov.) in close proximity has also been demonstrated in this region in Onychophora, which belong to ancient lineages that diverged in the Eocene or earlier (Daniels et al. 2016). This is in contrast to fi ndings in other molluscan taxa where populations across the region are considered to belong to the same lineage (Herbert 2017). (Melvill & Ponsonby, 1903)

Diagnosis
Shell very small, moderately depressed; periostracum with dense axial costae increasing slightly in height at periphery but not expanded into spines or fl anges; operculum duplex, almost fl at, multispiral lamellar blade low, loose fringe of bunches of bristles, fringe of each whorl with free ends not fused to lamella of following whorl; umbilicus deep and widely open exposing all the whorls; radula with two large cusps on second lateral tooth. COLE M.L., Revision of cyclophorid snails Chondrocyclus s.l.

Etymology
The specifi c name is derived from the Latin 'exsertus', meaning 'stretched or thrust out', and referring to the elevated spire.  33A-B, E-F). Spire moderately exserted, protoconch not mammillate (Fig. 33B). Embryonic shell (Fig. 33C) just over 2.5 whorls, microscopically malleate, roughest in centre; junction between embryonic shell and teleoconch evident with development of costae on teleoconch. Teleoconch comprising just under two whorls, very convex, rapidly increasing, suture impressed. Aperture circular, last whorl descending near aperture, peristome simple, continuous and free. Umbilicus wide and deep, exposing all the whorls (Fig. 33F). Periostracum pale yellowish-brown with lamellate axial costae at regular intervals, 84-110 (n = 14) on last whorl, each lamellar blade has numerous tiny vertical ridges visible at very high magnifi cation (Fig. 33D); lamellar blades increase in height at periphery but not expanded into fl anges or hairs; intervals between costae with microscopic axial threads. Shell translucent and corneous when fresh. OPERCULUM (Fig. 33H, J). Duplex, almost fl at, multispiral outer portion with a broad, thin nucleus and six whorls, height of lamellar blade low and not stepped, thickened horizontal ridge near bottom of blade just above disc surface, loose fringe of bristles emanates from ridge, fringe consists of bunches of short bristles, with free ends not fused to lamella of following whorl, fringe of outer whorl overlaps disc slightly. RADULA (Fig. 33G). Rachidian with fi ve cusps, central cusp approx. twice length of outer cusps, outermost cusps relatively weak, cusps of rachidian set a little distance below 'top' of tooth, top edge serrated; fi rst lateral tooth usually with three large cusps, third cusp (from centre) largest, a small fourth cusp and a vestigial fi fth; second lateral tooth with two large cusps, second cusp (from centre) longer and broader than fi rst, a very small third cusp and a vestigial fourth. PENIS (Fig. 33I, K). Shaft short, broad and more or less cylindrical, intromittent organ very short.

Distribution and habitat
Endemic to the coastal belt of southern Kwazulu-Natal and possibly extending into northern Pondoland. Recent specimens found in a few small patches of remaining indigenous forest inland of Umtentwini and Umzumbe (Fig. 29).

Remarks
The spire is more elevated than in the majority of other species of Afrocyclus and in the majority of species of Chondrocyclus s.s. barring C. herberti sp. nov., with similar shell proportions to the latter. The periostracum has simpler axial lamellae than other species, lacking fl anges or hairs. The operculum is unique in that the fringe consists of bunches of bristles with free ends which are not fused to the lamellar blade of the following whorl; fringe has a looser arrangement with fewer bristles than in all other species. The radula resembles that of A. isipingoensis gen. et comb. nov., including the unusual serrated top edge of the rachidian.
Afrocyclus exsertus gen. et comb. nov. is very rare and patchily distributed. Despite searching, it has not been found recently at any of the localities recorded by Connolly (1939) or in several other patches of forest on the Kwazulu-Natal south coast, excepting three very small patches among sugarcane fi elds, where it was presumably too steep for clearing of original forest. The vegetation on the Kwazulu-Natal south coast has undergone extensive modifi cation since 1939. (Bruggen, 1983) comb. nov. and Cyathopoma chirindae (Bruggen, 1986)

comb. nov.
Cyathopoma meredithae comb. nov. and Cyathopoma chirindae comb. nov. are removed from genus Chondrocyclus s.l. for three reasons, based on morphology of shell and operculum: i) shells have spiral keels while those of Chondrocyclus s.l. are smooth once the periostracum is removed, ii) opercula are calcareous, while those of Chondrocyclus s.l. are corneous, iii) periostracum is not glossy and lacquerlike as in all Chondrocyclus s.l. species, but is white and matt, suggesting a degree of calcifi cation. C. meredithae (Bruggen, 1983) comb. nov. and C. chirindae (Bruggen, 1986) comb. nov. resemble one another in general shape, possession of periostracal hairs at junctions between axial and spiral sculpture, and in the appearance of the operculum, but the latter has more dense axial lamellae and spiral keels (Fig. 34). Bruggen (1986) erroneously considered major spiral sculpture in the form of keels, ridges or cords to be a diagnostic feature of the Cyclophoridae in Africa. Shells of species of Chondrocyclus s.l. have spiral rows of periostracal projections, but are usually smooth once the periostracum is removed with only traces of axial sculpture. (Chondrocyclus kevincolei sp. nov. and to a lesser extent C. langebergensis sp. nov. from the Western Cape have spiral engraving.) Cyathopoma meredithae comb. nov. and C. chirindae comb. nov. both have a reticulate surface sculpture with spiral keels and axial lamellae more or less equal in density, with dark brown bristles on the points where the axial and spiral sculpture meet. In both species the periostracum is described as "corneous" (Bruggen 1983(Bruggen , 1986). However, the periostracum of C. chirindae (NM L4904) (Fig. 33B) does not appear to be "corneous", but is white and European Journal of Taxonomy 569: 1-92 (2019) matt, suggesting a degree of calcifi cation and this is also apparent to a lesser degree in C. meredithae comb. nov.
The descriptions of their opercula are also identical: duplex with exterior portion in the form of a large, concave, shallow saucer with six-eight spiral whorls (Bruggen 1983(Bruggen , 1986). Bruggen does not mention whether the opercula are corneous or calcareous. Examination of the operculum of C. chirindae comb. nov. (NM L4904) and the photograph of the operculum of C. meredithae comb. nov. (Fig. 34A) shows that both species have a calcareous operculum.
Attempts to assign the abovementioned two species to other genera are complicated by the fact that all other genera of operculate snails in mainland Africa and Madagascar are in need of revision ). Until recently, three of the African cyclophorids were placed in Afroditropis Bequaert & Clench, 1936, a genus characterised by widely spaced spiral keels on the shell, a thin peristome, not refl exed nor fl aring, and a thin, corneous, simple operculum. Afroditropis was synonymised with Cyathopoma by de Winter (2002), based on conchological similarity as well as similarities between the opercula of two specimens identifi ed as A. strongi Bequaert & Clench, 1936, and the opercular structure of Cyathopoma africanum Pilsbry, 1919, C. straeleni Adam, 1987 and C. camerunense de Winter, 2002.  (Bruggen, 1983) comb. nov., paratype and close-up of operculum (NHMUK 198338). B. Cyathopoma chirindae (Bruggen, 1986)  Whether these genera should be synonymised remains inconclusive since de Winter (2002) did not examine the operculum of the type specimen of A. strongi and he acknowledged that the two specimens on which he based his synonymy may actually represent an as yet undescribed species due to various differences between each of them and A. strongi. The other mainl and African genus in Cyclophoridae is Elgonocyclus Verdcourt, 1982 with two species, E. koptawaliensis (Germain, 1934) and an undescribed species from Ongoye forest, Zululand, both minute species with marked axial costulation and simple, corneous opercula.
Recently described African cyclophorids have been placed in Cyathopoma (e.g., C. camerunense de Winter, 2002;C. tres Bruggen, 2008and C. pembense Rowson, 2010in Rowson et al. 2010b and following this trend, Cyathopoma meredithae comb. nov. and C. chirindae comb. nov. were tentatively placed in Cyathopoma (Cole et al. 2019). The diagnosis in Blanford (1864) of Cyathopoma relies chiefl y on features of the operculum which is usually truncate-conoid, with the concave exterior portion formed by a calcareous, spiral lamella which usually curves inwards towards the centre and is sometimes beautifully sculpted and elevated. The only cyclophorid on mainland Africa with an operculum conforming to this pattern is Cyathopoma tres Bruggen, 2008, from Malawi, whereas in all the other mainland African species, including Cyathopoma meredithae comb. nov. and C. chirindae comb. nov., the operculum is concave since the erect spiral lamella is higher towards the periphery (Figs 34, 35A, C). There is considerable variation in the opercula of the 60 Malagasy species currently assigned to Cyathopoma s.l. so a future revision may lead to amendment of Blanford's diagnosis . At the time of the descriptions of Cyathopoma meredithae comb. nov. and C. chirindae comb. nov. the opercula of African Cyathopoma (except C. africanum Pilsbry, 1919) were not known because many of the current species had not yet been described (e.g., Cyathopoma straeleni Adam, 1987, C. camerunense de Winter, 2002, C. pembense Rowson, 2010 or the operculum was not known (C. azaniense Verdcourt, 1978). The opercula of both Chondrocyclus s.l. and Cyathopoma are duplex, with an erect spiral lamella forming the whorls of the exterior portion, but that of Chondrocyclus s.l. is uncalcifi ed (Kobelt 1902) while that of Cyathopoma is calcareous (Blanford 1864).
The reticulate surface sculpture of Cyathopoma meredithae comb. nov. and C. chirindae comb. nov. is unlike that of all other mainland African Cyathopoma which have prominent spiral keels and numerous closely set axial riblets. As far as is known, no other mainland African cyclophorids (except Chondrocyclus s.l.) have periostracal bristles. In some species the axial striae grow out into long processes of the periostracum, adhering in groups to the keels, e.g., Cyathopoma africanum Pilsbry, 1919 and C. pembense Rowson, 2010. It is not known whether such an elaborate periostracum occurs in the other African species because it is easily worn off. The type species of Cyathopoma, C. fi locinctum Benson, 1851, as well as a few Malagasy species , also have reticulate surface sculpture, and a few Malagasy species have periostracal hairs .
The reticulate appearance of Cyathopoma meredithae comb. nov. and C chirindae comb. nov. resembles several species of Malagasy Cyclotus   (Fig. 36A-B). In some Malagasy Cyclotus the axial lamellate costae elevate into triangular periostracal projections along the sutural edge and within the umbilicus ). In C. chirindae comb. nov. the sharp edged lamellae are raised into small triangular projections at the junctions with the spiral keels and these are particularly dense within the umbilicus (Fig. 34B). The operculum of the specimen of Cyclotus examined is fl at and bilayered ( Fig. 36C-D), and resembles that of the type species of Cyclotus, C. variegatus Swainson, 1840 (Fig. 36E), the outer surface concave, formed by an inwardly curving, spiral calcareous sheet that increases in elevation towards the periphery and without projections or appendages. Bruggen (1983) draws attention to the similarity between the opercula of C. meredithae comb. nov. and Cyclotus mamillaris Odhner, 1919 (which at the time of Bruggen's description was classifi ed as Chondrocyclus mamillaris (Fischer-Piette et al. 1993)).

European Journal of Taxonomy 569: 1-92 (2019)
Cyathopoma meredithae comb. nov. and C chirindae comb. nov. both have a thin peristome, practically free, but with a limited area touching the body whorl (Fig. 34), but it is not certain whether the specimens examined were adult. In adult Chondrocyclus s.l. the peristome is detached from the last whorl and droops downwards, but in subadults it is attached in a limited area. The peristome of C. fi locinctum Benson, 1851 is moderately thick with a broad basal fl are, and in several mainland African and Malagasy Cyathopoma it is expanded and may also be thickened.
The radulae and penes of mainland African and Malagasy Cyathopoma and Cyclotus have not been documented prior to this study and no bodies of Cyathopoma meredithae comb. nov. or C. chirindae comb. nov. were available for examination. These features have proved useful in diagnosis and taxonomy of species of Chondrocyclus s.l., so their examination in C. meredithae comb. nov. and C. chirindae comb. nov. may provide useful insights as to their affi nities, even though they would probably not assist with generic placement at present. The penis of Cyathopoma pembense is extremely long and narrow relative to that of Chondrocyclus s.l. and does not appear to have an intromittent organ, but tapers to a very fi ne point (Fig. 35D). The penis of Cyclotus sp. from Madagascar resembles that of C. pembense, but has a relatively broader shaft which also narrows to a fi ne point and no intromittent organ ( Fig. 36G-Fig. 35. Cyathopoma pembense Rowson, 2010 H). What appears to be a groove possibly corresponding to a seminal groove, runs down the ventral surface. In both these species the penis lies doubled back on itself.
The radulae of Cyathopoma pembense and Cyclotus sp. differ (Figs 35B and 36F). Cyathopoma pembense has a serrated upper edge to the rachidian tooth, also found in Afrocyclus gen. nov., but the fi rst and second lateral teeth have four large cusps (often with extras), unlike the radula of any Afrocyclus gen. nov. or species of Chondrocyclus s.s. The radula of Cyclotus sp. resembles the radulae of European Journal of Taxonomy 569: 1-92 (2019) several species of Chondrocyclus s.s., with three large cusps on the second lateral tooth. As noted in the Comparative morphological observations section, radulae of various Cyclophoridae look superfi cially alike, so similarities in the radula do not necessarily imply close relationship.

Discussion
This study is a signifi cant contribution to the knowledge of molluscs on the African continent and in South Africa in particular. Although considerable progress has been made in taxonomy and systematics of South African terrestrial molluscs in the past few decades, the majority of South African taxa still require revision and a host of species await description. Studies which have been undertaken consist of species descriptions across a small range of taxa (Bruggen 1965(Bruggen , 1966(Bruggen , 2006Herbert 2002Herbert , 2006Herbert , 2016Bursey & Herbert 2004;Cole & Herbert 2009), a few revisions (Herbert 2007;Herbert & Moussalli 2010, 2016 and a limited number of other treatments (Sirgel 1985;Herbert 1997Herbert , 2006Herbert , 2017Herbert & Kilburn 2004). In total, only a handful of molecular studies have been undertaken on African land snails (Herbert & Mitchell 2009;Rowson & Herbert 2016;Moussalli et al. 2009;Cole et al. 2019) and very few global studies have included African taxa (Rowson et al. 2010a;Fontanilla et al. 2017;Harl et al. 2017).
Chondrocyclus s.s. and, as far as is known, Afrocyclus gen. nov. are endemic to South Africa and the majority of species are narrow-range endemics. This study reveals a large proportion of undetected diversity in the family, both as cryptic species previously included under other names (i.e., overlooked because identifi cations were based on old/worn shell material) or as new discoveries. With a better understanding of the family's species composition and distribution, an assessments of species qualifying for Red Listing can now be made. Forests cover only 0.1% of South Africa's land surface and are highly fragmented (Mucina & Geldenhuys 2006). Forest degradation and further fragmentation through human activity are threatening the survival of the biome and leading to changes in species abundance and distribution (Eeley et al. 1999;Cooper et al. 2017). These habitats have been shown to contain high levels of undetected diversity and endemism in molluscs (Herbert 2016(Herbert , 2017Bursey & Herbert 2004;Cole & Herbert 2009;Herbert & Moussalli 2010, 2016 and other low-vagility invertebrate taxa (Huber 2003;Hamer & Slotow 2002;Tilbury & Tolley 2009;de Bivort & Giribet 2010;Ruhberg & Daniels 2013;Daniels et al. 2009Daniels et al. , 2013Daniels et al. , 2016. The contractions and expansions of forest habitat due to climatic oscillations (Mucina & Geldenhuys 2006;Scott et al. 1997;Partridge et al. 1999) appear to have resulted in the formation of complex phylogeographic patterning, potentially promoting cladogenesis among forest-dwelling habitat specialists Daniels et al. 2016Daniels et al. , 2017. Observations on biogeography of Chondrocyclus s.l., and parallel distribution patterns in a range of low-vagility forest-dwelling taxa, are discussed in Cole et al. (2019) but data on invertebrates remains patchy (McGeoch et al. 2011;Daniels et al. 2017). This study will complement other research by providing comparative material for an independent, widespread group and expand the scientifi c evidence available for biodiversity conservation in South Africa. Improved knowledge of the composition and distribution of this group of forest specialists with weak dispersal capabilities may contribute insights into the processes generating biodiversity in the region where common patterns across unrelated taxa are beginning to emerge. This is the fi rst detailed systematic revision of an Afrotropical cyclophorid group to include morphological and molecular data. It lays a foundation for further studies of the family elsewhere in Africa, including relationships between taxa. Molecular data and anatomical studies on the soft parts of other African cyclophoroidean taxa, including Cyathopoma meredithae comb. nov. and C. chirindae comb. nov., Elgonocyclus spp. and the mainland African endemic family Maizaniidae to supplement morphological observations of their shells and opercula would enable assessment of their affi nities to Chondrocyclus s.l. COLE M.L., Revision of cyclophorid snails Chondrocyclus s.l.