Genus Kermia (Mollusca: Gastropoda: Conoidea: Conidae: Raphitominae) in South African waters, with observations on the identities of related extralimital species.
The genus Kermia Oliver, 1915, is recorded for the first time from
South Africa. New species described are: Kermia geraldsmithi sp. n., K.
clathurelloides sp. n., K. drupelloides sp. n. and K. eugenei sp. n. New
records for southern Africa are: Kermia melanoxytum (Hervier, 1896), K.
edychroa (Hervier, 1897), K. pumila (Mighels, 1845), K. daedalea
(Garrett, 1873) and K. producta (Pease, 1860). New synonymies
established are: Clathurella birtsi Preston, 1908 and Mangelia
chrysolitha Melvill & Standen, 1896 under Kermia punctifera Garrett,
1873; Pleurotoma gracilis Folin, 1871 and Mangelia anxia Hedley, 1909
under Kermia edychroa (Hervier, 1897); Pseudodaphnella punicea Hedley,
1922 under Clathurella blanfordi pullula Hervier, 1897 = Pseudodaphnella
pullula; Defrancia mauritiana Sowerby, 1893 and Clathurina foraminata
pyrgodea Melvill, 1917 under Clathurella producta Pease, 1860;
Clathurella spelaeodea Hervier, 1897 under Clathurella infrasulcata
Garrett, 1873 (currently Pseudodaphnella subgenus Qii). Type specimens
of the following species are also illustrated: Pleurotoma apicalis
Montrouzier in Souverbie, 1861; Pleurotoma cavernosa Reeve, 1845;
Pleurotoma margaritifera Reeve, 1846; Glyphostoma melanoxytum Hervier,
1896; Clavatula tessellata Hinds, 1843. New combinations are
established: Clathurella blanfordi G. & H. Nevill, 1875 transferred
to the genus Paramontana Laseron, 1954, and Clathurella martensi G.
& H. Nevill, 1875 to Pseudodaphnella Boettger, 1895. Pleurotoma
pustulosa Folin, 1867 and Pleurotoma foraminata Reeve, 1845 are
considered nomina dubia. A lectotype is designated for Clathurella
daedalea Garrett, 1873.
KEY WORDS: Conidae, Raphitominae, Kermia, South Africa, Indo-West Pacific, new species, new synonymy.
Cone snails (Physiological aspects)
Cone snails (Identification and classification)
|Author:||Kilburn, Richard Neil|
|Publication:||Name: African Invertebrates Publisher: The Council of Natal Museum Audience: Academic Format: Magazine/Journal Subject: Zoology and wildlife conservation Copyright: COPYRIGHT 2009 The Council of Natal Museum ISSN: 1681-5556|
|Issue:||Date: Dec, 2009 Source Volume: 50 Source Issue: 2|
|Geographic:||Geographic Scope: South Africa Geographic Code: 6SOUT South Africa|
The Indo-Pacific genus Kermia Oliver, 1915, has not previously been reported from South Africa. However, at least nine species occur, four of them here described as new.
Like most raphitomine genera, Kermia has been very inadequately studied, and its limits are unclear. Characteristics include the presence of cancellate teleoconch sculpture, one or more diagonally cancellate protoconch whorls (initially smooth or spirally striate), and a linguiform to lacrimiform anal sinus, its upper margin drawn forward at suture by a callous node; outer lip with a strong varix, denticulate to dentate within.
The Mediterranean genus Philbertia Monterosato, 1884, sometimes used for Indo-Pacific taxa, is more applicable to species with a shallow, reversed L-shaped anal sinus. Using this definition, Kermia is a large genus of approximately 36 described species in the Indo-West Pacific. However, observed variation in protoconch characters is probably indicative of polyphyly. Borderline cases include the type species, Kermia benhami, which is atypical in its protoconch possessing arcuate axial riblets, with only their bases diagonally cancellated. The only Western Hemisphere species referred by authors to Kermia is K. informa McLean & Poorman, 1971, from Ecuador; however, this taxon evidently lacks diagonally cancellate protoconch sculpture.
A number of similar species with terminal protoconch sculpture of orthocline or arcuate axial riblets, with or without spiral threads, are provisionally referred to the genus Pseudodaphnella Boettger, 1895. As noted above, in Kermia the protoconch is initially smooth or spirally striated, but later whorls are diagonally cancellate. Nevertheless, it is possible that the two taxa may intergrade, as diagonally cancellate sculpture may only develop towards the termination of the protoconch. It should also be noted that specimens of some Pseudodaphnella species in which obliquely arcuate axial riblets cross spiral threads may confusingly resemble Kermia species, particularly when slightly eroded.
Pseudodaphnella auct. is probably polyphyletic, and contains several mostly unnamed (sub)groups. In the group containing the type species (Pleurotoma philippinensis Reeve, 1843) the aperture is wide and the outer lip is strongly arched and bears relatively numerous small pleats or denticles. However, although Hedley (1922: 349) described the last whorl of the protoconch as bearing fine axial threads in P. philippinensis, Shuto (1971: 18) claimed that the sculpture was diagonally cancellate. Species resembling P. philippinensis (at least in teleoconch characters) include Pleurotoma nexa Reeve, 1845, Clathurella pulchella Pease, 1860, and Pleurotoma granicostata Reeve, 1846. A second, larger, group, which includes species such as Pleurotoma tincta Reeve, 1846, Clathurella rubroguttata H. Adams, 1872, Pleurotoma lemniscata G. & H. Nevill, 1869, Clathurella cnephaea Melvill & Standen, 1896, and Clathurella ramsayi Brazier, 1876, differs from the philippinensis complex in its constricted base and in the presence of four or five strong labral denticles. However, several species, such as Pleurotoma (Drillia) hadfieldi Melvill & Standen, 1895 (= Clathurella ephela Hervier, 1897) and Clathurella rufolirata Hervier, 1897, are somewhat intermediate.
Reference may also be made to Pseudodaphnella subgenus Qii Zhang, 1995, whose type species (o.d.) Clathurella spelaeodea Hervier, 1897, is a synonym of Clathurella infrasulcata Garrett, 1873, according to comparison of their syntypes in the MNHN and ANSP respectively. Zhang (1995) characterised Qii by its strong basal furrow (waist) which is demarcated adapically by a somewhat strengthened spiral ridge. According to fresh examples of this species (P infrasulcata occurs as far south as Durban), the protoconch bears cancellate sculpture of fine axial ribs crossed by subequal spiral threads (Fig. 56). Other species presumably referrable to Qii are Pseudodaphnella oligoina Hedley, 1922, Pleurotoma (Defrancia?) alternans E.A. Smith, 1882, and perhaps Clathurella spyridula Melvill & Standen, 1896. However, some other species with a similar sunken waist and bordering ridge have a Kermia-type protoconch, so these characters presumably cannot be regarded as apomorphic.
Figures of radulae given by Orr-Maes (1967) appear to show no obvious differences between the two genera (figs 4G, H and I, representing respectively Pseudodaphnella granicostata (Reeve, 1846) (as Philbertia (Pseudodaphnella) granicostata, Pseudodaphnella rubroguttata (H. Adams, 1872), as Philbertia (Pseudodaphnella) tincta (non Reeve, 1846), and Kermia barnardi (Brazier, 1876), as Philbertia (Kermia) barnardi). However, no radula was present in material of P. granicostata examined by Kantor and Taylor (2002).
Melvill (1917) proposed the name Clathurina for a polyphyletic group of Kermialike species, with Pleurotoma foraminata Reeve, 1845, as the type species. Clathurina was synonymised with Kermia by Powell (1966: 134), but its type species should be regarded as a nomen dubium, as no types of P. foraminata are present in the BMNH, and Reeve's tiny figure and brief description (1845, pl. 33, sp. 301) are of little value in positive recognition of the species. However, specimens labelled as P. foraminata in the Melvill-Tomlin collection (NMWC) are actually Kermia margaritifera (Reeve, 1846) from India and the Arabian Gulf. When compared with Reeve's figure of foraminata, K. margaritifera (Fig. 49) has a shorter, more truncate base, a wider aperture and stronger, more numerous spiral ridges. The protoconch sculpture of Indian examples (Fig. 55) appears to be atypical for Kermia in having oblique axial riblets crossed by interrupted spiral threads.
MATERIAL AND METHODS
The studied material is kept in the following institutions:
AMSA--The Australian Museum, Sydney, Australia;
ANSP--Academy of Natural Sciences, Philadelphia, USA;
BMNH--The Natural History Museum, London, UK;
MCZH--Museum of Comparative Zoology, Harvard, USA;
MHNB--Museum d'Histoire Naturelle, Bordeaux, France;
MMUE--Manchester Museum University, Manchester, UK;
MNHN--Museum National d'Histoire Naturelle, Paris, France;
NMSA--Natal Museum, Pietermaritzburg, South Africa;
NMWC--National Museum of Wales, Cardiff, UK;
SML--Societe de Marie, Lyon, France;
USNM--National Museum of Natural History, Washington DC, USA;
ZSIC--Zoological Survey of India, Kolkata (Calcutta), India.
The following abbreviations and acronyms are used in this paper:
a/l--ratio of aperture length (measured along main shell axis) to total shell length;
b/l--ratio of breadth to total length;
ESEM--environmental scanning electron microscope; loc.--locality;
t/s--transverse section (i.e. as seen in apical view).
Family Conidae Fleming, 1822
Subfamily Raphitominae Bellardi, 1875
Genus Kermia Oliver, 1915
Kermia: Oliver 1915: 539. Type species: Kermia benhami Oliver, 1915, by original designation.
Kermia geraldsmithi sp. n.
?Philbertia (Pseudodaphnella) sp.: Oyama & Takemura 1957: Philbertia (2), figs 7, 8.
Etymology: Named in honour of its collector, Mr Gerald Smith of Durban. Description: Shell fusiform with truncated base; b/l 0.40-0.46, a/l 0.36-0.46; whorls slightly convex with angular, sloping shoulder, subsutural region wide and concave, suture undulating, base of body whorl constricted above rostrum into a slight but distinct waist; rostrum thick and convex; siphonal canal strongly contracted but expanded terminally, end truncate, level, in dorsal view medially convex, not notched. Outer lip flat in side view, with 4 serrations, caused by ridge terminations; inner edge with 3 equal-sized teeth and sometimes a weak 4th; inner lip smooth and flattened, posterior end of aperture with a parietal pad. Anal sinus shallow, U-shaped.
[FIGURES 1-7 OMITTED]
Axial ribs slightly prosocline, fairly straight, weaker below suture, on base extending onto rostrum as rounded nodules; in t/s high, asymmetrically rounded, with steeply sloping sides, ribs wider than their intervals, 9 or 10 per whorl. Spiral ridges thinner than axials in intervals, expanding on ribs to form angular nodules, ridges nearly equal in strength, adapical row slightly the stronger, abapical one slightly weaker; first two whorls with two rows of nodules plus a third usually beginning to develop above suture, later whorls with three main rows of nodules, plus a weak ridge immediately below suture; last whorl with a shallow waist-like furrow at base of rostrum containing a thin ridge, strengthening and bearing angular nodules on dorsum; rostrum forming a moderate fasciole, which bears 3 ridges weakening anteriorly, the upper 2 with angular nodules. Interstices with extremely fine and close collabral striae.
Protoconch (Fig. 3) bluntly conical, of 3.3 whorls, last two whorls with prosocyrt axial riblets, crossed diagonally by oblique threads; 1st whorl with a few minutely pliculate spiral threads above its suture; dark brown; breadth 0.46 mm.
White, alternate ribs with axially elongated orange-brown blotches, basal constriction dark to medium orange-brown; protoconch dark orange-brown.
Dimensions: 4.6x2.1 mm (holotype), largest paratype 5.5x2.3 mm.
Comparison and remarks: This appears to be the species figured by Oyama and Takemura (1957) as 'Philbertia (Pseudodaphnella) sp.', based on a specimen from southern Japan (Amami-Oshima Is). The most similar described species is the Japanese Clathurella excellens Sowerby, 1913. The holotype of that (Fig. 4) differs from K. geraldsmithi in its angular shoulder, darker brown blotches, fewer axial ribs (8 per whorl) and more numerous spiral ridges (5 on penultimate whorl). Japanese authors (e.g. Oyama & Takemura (1957: Philbertia (3), figs 7, 8); Fukuda (1995: 24, pl. 67, fig. 986); Higo et al. (2001: 109, fig. G3809, holotype); Hasegawa et al. (2000: 659, pl. 328, fig. 194) cite excellens as Philbertia (Pseudodaphnella) excellens. Unfortunately, the holotype of that species lacks its protoconch, and no details of it are available to me, so that its generic position remains to be confirmed.
Holotype: NMSA W6533/T2354. SOUTH AFRICA: KwaZulu-Natal: off Park Rynie (30[degrees]19'S:30[degrees]44'E), depth ca -35 m, in shell grit, dived G. Smith, 1996.
Paratype 1: same data as holotype, to be deposited in BMNH. Paratypes 2-9: NMSA S2999/T 2377, between Bhanga Neck and Kosi Bay, depth ca -13 m, D. Herbert.
Other material examined: SOUTH AFRICA: KwaZulu-Natal: Kosi Bay, 14 km S of estuary mouth, ca -20 m (D. Herbert, R. Broker; NMSA S2579), 1-4 km S of estuary mouth, ca -18 m (D. Herbert, K. Bloem; NMSA S2311) and -23-24 m (D. Herbert, K. Bloem; NMSA S2900); Park Rynie, ca -16 m, in shell grit (D. Herbert; NMSA S5920); Aliwal Shoal, off Scottburgh, ca -16 m, sand (D. Herbert; NMSA S5920).
Distribution: Inner continental shelf of KZN and probably southern Japan, in -13-35 m.
Kermia clathurelloides sp. n.
Etymology: From the genus name Clathurella Carpenter, 1857; it refers to the teleoconch being superficially similar to some species of latter genus.
Description: Shell fusiform with truncated base; b/l 0.35-0.38, a/l 0.38-0.41; whorls strongly convex with narrow angular shoulder, and undulant, fairly deep suture, base of last whorl without a waist-like constriction; rostrum narrow, without a fasciole; aperture narrowly elliptical; siphonal canal strongly contracted but slightly expanded terminally, end truncate, oblique in dorsal view, not notched. Outer lip evenly convex in side view, not serrated, anal sinus moderately deep, linguiform, its opening constricted, margin raised; inner edge of lip with 9-11 weak, ridge-like teeth, one at each end stronger than others, but in some specimens very irregular; inner lip smooth, shallowly sinuous, thinly calloused, parietal pad weak.
[FIGURES 8-9 OMITTED]
Sculpture cancellate, with spiral ridges finer than axial ribs, forming weak compressed nodules where they cross them. Axial ribs slightly opisthocline, almost straight, weakly sigmoid on last whorl, terminating (or weakening) above, slightly short of suture, on base extending to top of rostrum; on last whorl continuing to outer lip; in t/s constricted, steep-sided, sharply rounded, equal to or slightly narrower than their flattened intervals, 10 or 11 on 1st teleoconch whorl (first few ribs weak), 15 or 16 on penultimate whorl. Spiral ridges narrower than their intervals, nearly equal in strength, except for those on base of last whorl which are uneven; 1st whorl with 3 spiral ridges, the upper two close together; penultimate whorl with 8-12 ridges, base with 6-8, plus 4-7 smooth, slightly angular ridges on rostrum (which may become obsolete distally). Interstices with fine, close axial striae.
Protoconch (Fig. 7) bluntly conical, of ca 3.0 whorls, beginning of 1st whorl strongly tilted, next whorl finely cancellate, last whorls with obliquely cancellate sculpture, of sigmoid, opisthocyrt riblets, crossed diagonally by oblique threads; golden brown; breadth 0.41 mm.
Slightly glossy, medium or golden brown, with obscure darker and lighter bands, spiral ridges more or less darker than intervals, last whorl with a terminal brown band basally; interior of outer lip white, protoconch orange-brown.
Dimensions: 11.3 x 4.0 mm (holotype); 12.0x4.3 mm (paratype).
Comparison and remarks: Within the genus Kermia, the new species shows some resemblance to K. producta (Pease, 1860), but that species is only about half the size of K. clathurelloides (adults of both species having a maximum of about six teleoconch whorls), axial ribs are much finer in producta (20-22 on the penultimate whorl against 15 or 16) and markedly straighter, and the suture is crenulated by the rib terminations in clathurelloides, but not in producta. There also appears to be some resemblance to K. thorssoni Chang, 2001, from Taiwan (types not seen), but K. clathurelloides differs in its later whorls being distinctly shouldered, the last whorl lacking a submedian angle and the spiral ridges are much finer (in thorssoni these are subequal in strength to the axial ribs and fewer on the penultimate whorl (five according to Chang), whereas these number 8-12 in K. clathurelloides).
A relationship with Clathurella exilis Pease, 1860, from Hawaii is also possible. Unfortunately, the lectotype of that (BMNH 1962766, designated by Kay (1965: 21, pl. 2, figs 17, 18)) lacks its protoconch. Kay (1979: 356, fig. 116J) referred exilis to the genus Paramontana (a probable clathurelline group with smooth protoconch), but her illustration appears to show finer sculpture and a more elliptical aperture than the lecto-type (Figs 8, 9), so that confirmation of the identity of her material is needed. Compared with the lectotype of exilis, K. clathurelloides has weaker lip denticles, a distinct shoulder angle to the ribs, and a much more constricted base.
K. clathurelloides shows a superficial resemblance to several species generally referred to the genus Lienardia Jousseaume, 1884. Of these, the most similar species (apart from colouration) appear to be L. roseotincta (Montrouzier in Souverbie & Montrouzier, 1872), which also occurs in KZN, and L. montrouzieri (Souverbie, 1860). However, these two species are typical Lienardia in possessing a keeled protoconch, whereas that of K. clathurelloides not only lacks a keel but has the typical diagonally cancellate sculpture of Kermia.
Holotype: NMSA W6533/T2360. SOUTH AFRICA: KwaZulu-Natal: off Reunion Rocks, Durban (29[degrees]58'S: 30[degrees]56'E), ca -130 m, sand and rubble bottom, dredged G. Smith.
Paratype 1: same data as holotype, to be deposited in BMNH. Paratype 2: KwaZulu-Natal: Leadsman Shoal, -24-26 m, stone washings (D. Herbert; NMSA E2634/T2343). Eastern Cape: Off Port Grosvenor, -80 m, calcareous nodules (R. Kilburn et al.; NMSA C7367/T2342).
Other material examined comes from the type locality and is in G. Smith's collection.
Distribution: Eastern South Africa, from northern KZN to Eastern Cape, inner continental shelf.
Kermia drupelloides sp. n.
Etymology: From the genus name Drupella Thiele, 1925.
Description: Shell fusiform with high, orthoconoid spire and truncated base; b/l 0.380.43, a/l 0.33-0.39; whorls convex, not shouldered, subsutural region concave and sloping, suture not undulating, base of last whorl constricted above rostrum into a deeply concave "waist"; rostrum forming a strongly convex fasciole; aperture narrowly elliptical; siphonal canal strongly contracted but expanded terminally, end truncate, in dorsal view concave, with a slight projection medially. Outer lip almost flat in side view, with 4 slight, rounded serrations at terminations of spiral ridges; inner edge with 3 equal-sized, rounded teeth and sometimes a weak 4th; inner lip sigmoid, with a flattened callus deposit with slightly free edge, sometimes with a few feeble plicae, posterior end of aperture with a parietal pad. Anal sinus shallowly linguiform, directed slightly posteriorly, situated on subsutural ridge.
Sculptured by spiral cords that are narrower than the axial ridges, but expand to form angular nodules where they cross them. Axial ribs straight, slightly prosocline, moderately weak below suture, evanescing below "waist", fasciole with strong nodules; ribs in t/s more or less angularly rounded, with sloping sides, ribs wider than their intervals, 8 per whorl; bases of ribs narrowing where they cross basal "waist", where they are separated by foveolate interstices. First teleoconch whorl cancellate, with 2 spiral ridges, increasing to 4 from 2nd whorl, that below suture remaining weak, that above suture sometimes as strong as the median two (i.e. with three main rows of nodules); last whorl with 5 rows of nodules, of which subsutural one is weak, fasciole bearing 3-5 rows of strongly rounded tubercles, progressively weakening towards base. Interstices with extremely fine and close collabral striae.
Protoconch bluntly conical, of about 2.7 whorls, 1st whorl depressed and tilted, 2nd whorl with cancellate sculpture (about 9 spiral threads, crossed by orthocline axial threads); last whorls with arcuate axial riblets, crossed by short, opisthocline axial threads, interrupted in the intervals; breadth 0.43 mm.
[FIGURES 10-16 OMITTED]
Light brown, nodules, rostrum and early teleoconch whorls paler, aperture and its margin white; protoconch light brown.
Dimensions: 4.4 x 1.9 mm (holotype), 6.3 x 2.4 mm (paratype).
Comparison and remarks: This distinctive species shows some resemblance to the widely distributed Indo-Pacific K. punctifera (Garrett, 1873). The new species differs from K. punctifera in lacking characteristic brown marks on its tubercles, which are also markedly stronger and more angular, and its proportions are broader. Based on their types in the MMUE, BMNH and ANSP collections respectively (Figs 13-16), it is probable that Mangelia chrysolitha Melvill & Standen, 1896, and Clathurella birtsi Preston, 1908, are synonyms of K. punctifera, although they have fewer spiral cords and lip denticles. K. drupelloides also appears to show some resemblance to the figure of Pleurotoma microcerata type (Folin 1879: 248, pl. 8, fig. 2), described from the Andaman Islands, but that shows sharper nodules, which are spinose on the base, interstices that appear foveolate, an internally smooth outer lip, and its colour is white. Unfortunately, the holotype of P. microcerata is absent from the Folin turrid types preserved in the BMNH, and must be considered lost.
Sometimes the name Pleurotoma pustulosa Folin, 1867, has been used for specimens of Kermia punctifera, particularly in the Japanese and Chinese literature. However the figure of the P. pustulosa type (Folin 1867, pl. 5, fig. 14) illustrates finer, more numerous nodules, particularly so on the base; furthermore, the type material, which is evidently lost, might have originated from either the Philippines or the Bay of Panama. For the present I regard P. pustulosa as a nomen dubium.
Several species of Pseudodaphnella Boettger, 1895, and Paramontana Laseron, 1954, are essentially similar to K. drupelloides, but have a different protoconch. In the genus Pseudodaphnella this is generally paucispiral and papilliform with more or less orthocline axial riblets and weak spiral threads. A superficially comparable species of Pseudodaphnella is P. pullula (Hervier, 1897) from New Caledonia, which has a narrower indentation (waist) on the base and a narrower aperture than K. drupelloides. A synonym of Clathurella blanfordi var. pullula is Pseudodaphnella punicea Hedley, 1922 (Figs 17, 18). Another somewhat similar species is Clathurella martensi G. & H. Nevill, 1875 (Fig. 19), distributed from Sri Lanka to New Caledonia and southern Japan, which appears to have a Pseudodaphnella-type protoconch, similar to that of P. pullula.
Confusingly, Clathurella blanfordi G. & H. Nevill, 1875, of the Red Sea, has similar teleoconch sculpture to P. pullula and P. martensi comb. n. (Figs 20, 21), but the protoconch (in fresh condition) appears smooth, indicating a position in the genus Paramontana. This genus has been referred to the Mangeliinae (Sysoev 1993: 167), presumably on account of the smooth protoconch, but teleoconch characters are perhaps suggestive of the Clathurellinae or even Raphitominae. The teleoconch in Paramontana blanfordi comb. n. differs from that of K. drupelloides and P. martensi in its flattened lip profile, with a distinctive angle anteriorly.
Apart from its raphitomine protoconch and small adult size, K. drupelloides shows superficial resemblance to some members of genus Drupella (Muricidae).
Holotype: NMSA W6532/T2358. SOUTH AFRICA: KwaZulu-Natal: off Scottburgh (30[degrees]17'S:30[degrees]45'E), ca -35 m, in shell grit, 1995. Donated by G. Smith.
Paratype 1: same data, to be lodged in BMNH. Paratype 2: KwaZulu-Natal: Park Rynie, -35-45 m, dived (NMSA S9826/T2350).
Other material in G. Smith collection.
Distribution: KZN, in Park Rynie--Scottburgh area, inner continental shelf.
Kermia eugenei sp. n.
Figs 22, 23
Etymology: Named in honour of my son, Eugene, for his help and encouragement. Description: Shell globose-fusiform with a tapering base; b/l 0.43-0.47, a/l 0.47-0.56; whorls strongly convex with a slight indentation below suture, and a fairly deep suture, crenulated by rib terminations, base of last whorl only slightly constricted at top of rostrum; rostrum narrow, flat-sided, without a fasciole; aperture narrowly elliptical, more or less linear, constricted at siphonal canal, which is more or less parallel-sided, its termination oblique, shallowly indented in dorsal view; columella straight. Outer lip flattened in side view, slightly serrated at ridge terminations, edge incurved; anal sinus moderately deep, linguiform, abapically directed, its opening constricted, its margin not raised; interior of lip with 7 ridges or small teeth, the strongest one at entrance to siphonal canal; inner lip smooth, with a flattened callus, parietal pad weak.
Sculpture nodose-cancellate, of axial ribs crossed by narrower spiral ridges, forming compressed nodules, interstices rather deep and quadrate, a slightly deeper interval demarcating rostrum. Axial ribs somewhat arcuate, slightly opisthocyrt on last whorl, reaching and crenulating suture, evanescing on upper part of rostrum, fasciole with strong nodules; ribs in t/s strongly rounded and slightly compressed on spire whorls, lower with sloping sides on last whorl, ribs slightly wider than their intervals, one behind lip high and varicoid, 12 or 13 axials per whorl. Spiral ridges more or less evenly sized, first teleoconch whorl with 2, increasing to 3, spiral ridges, 2nd whorl with 4; penultimate whorl with 5-7 ridges, subsutural one or two usually weaker than others; base of last whorl with an additional 2 ridges, rostrum bearing 7 or 8 close, angular spiral ridges, upper 2 or 3 somewhat crenulate, progressively weakening towards base. Prelabral varix crossed by 8-10 ridges. Interstices with extremely fine and close collabral striae (coarsest behind lip).
[FIGURES 22-26 OMITTED]
Protoconch conical, of ca 3.3 convex whorls, 1st one tilted, last one very finely diagonally cancellate, penultimate one punctate; breadth of last whorl 0.37 mm.
Uniform pale to medium brown, early spire whorls sometimes faintly tinted with orange, protoconch pale brown; columella and inner edge of outer lip orange-brown.
Dimensions: 3.5x1.4 mm (holotype); 4.2x1.8 mm (largest paratype).
Comparisons and remarks: K. eugenei appears closely comparable only with K. cavernosa (Reeve, 1845) from the Philippines. The freshest of five syntypes of Pleurotoma cavernosa (BMNH 1963801), which retains its protoconch, is here illustrated (Figs 25, 26). Compared with K. eugenei, K. cavernosa has flatter whorls, only two spiral ridges on the early teleoconch whorls, a distinct waist-like basal furrow demarcating four or five nodular cords on the rostrum, and lacks an orange inner margin to the aperture. K. aniani Kay, 1979, of Hawaii, is markedly more cylindrical in shape, with a narrower aperture, stronger labral denticles and more numerous spiral ridges (Fig. 24).
Holotype: NMSA V23411/T2331. SOUTH AFRICA: KwaZulu-Natal: 1-4 km S of Kosi Bay (26[degrees]54'S: 32[degrees]52'E), -20-22 m, reef, D. Herbert & K. Bloem.
Paratypes: SOUTH AFRICA: KwaZulu-Natal: between Bhanga Neck and Kosi Bay, reef near pinnacles, -10-12 m, hand-dredged sand (D. Herbert; NMSA S2371/T2334); Leadsman Shoal, coral reef, -7-11 m, 1 specimen (D. Herbert & Natal Parks Board; NMSA E6778/T2332); off Durban Bluff, -20-22 m, sand (reclamation dump), 5 specimens (R. Kilburn & R. Fregona; NMSA B5467/T2333).
Distribution: From northern KZN to Durban, in ca -10-22 m.
Kermia melanoxytum (Hervier, 1896)
Glyphostoma melanoxytum: Hervier 1896a: 150; 1896b: 78, pl. 3, fig. 19. Type locality: Lifou. Kermia melanoxytum (Hervier): Kay 1979: 361, figs 117F (protoconch), 118J; Zhang 1995: 290, pl. 4, figs 23-24; Chang 2001: 72, figs 65, 65b, and p. 212, figs 19-16a, b.
Type material: Originally ten syntypes from Hervier collection in SML; three probable syntypes now in MNHN, one labelled by Fischer-Piette as "Type" 4.6x2.0 mm, early part of protoconch missing; unconfirmed syntypes AMSA C.7019.
Regional locality data: SOUTH AFRICA: KwaZulu-Natal: E of Kosi Bay, -50 m, fine sand (Kilburn et al.; NMSA E5348); Kosi Bay, 1-4 km S of mouth, -20-22 m (D. Herbert & K. Bloem; NMSA S775); off Island Rock, -62 m, sandstone, coral (Kilburn et al.; NMSA E5551); Leadsman Shoal, 1-2 km N of Leven Point, -9-14 m, mixed algal and coral reef (D. Herbert; NMSA E5375); off Sodwana Bay, talus slope, -46 m (Marine Geoscience Unit, University of KwaZulu-Natal, Durban; NMSA V5072); Aliwal Shoal, off Scottburgh, -10 m, reef debris (D. Herbert; NMSA S9237); same data but -25-27 m (D. Herbert; NMSA S7131); off Park Rynie, -35-40 m, offshore reef (G. Smith; NMSA S9857). Beach-worn shells also seen in private collection from Mzamba, Eastern Cape.
Distribution: New Caledonia to Hawaii, and west to south-eastern Africa, as far south as Eastern Cape.
Remarks: Specimen NMSA S775 (Fig. 29) has been compared with one of the "syntypes" (Figs 27, 28) in the MNHN.
Kermia edychroa (Hervier, 1897)
Clathurella edychroa: Hervier 1897a: 146; 1897b: 109, pl. 3, fig. 2; Bouge & Dautzenberg 1914: 195. Type locality: Lifou.
Clathurella edychroa var. violacea Hervier, 1897b (non Bucquoy, Dautzenberg & Dollfus, 1883): 110. Type locality: Lifou.
Kermia edychroa (Hervier): Kay & S. Johnson 1987: 122, fig. 4U.
Pleurotoma gracilis Folin, 1879 (non Conrad, 1830, nec Scacchi, 1836): 251, pl. 8, fig. 6. Syn. n. Type locality: Andaman Is.
Mangelia anxia Hedley, 1909: 455, pl. 43, fig. 89. Syn. n. Type locality: Hope I., Great Barrier Reef, Queensland, 5-10 fath. [9-18 m].
Exomilus anxius (Hedley): Hedley 1922: 333; Powell 1966: 137.
[FIGURES 27-33 OMITTED]
Type material: C. edychroa: originally two syntypes in SML, one possible syntype now in MNHN (labelled "Holotype"), worn and lacking its apex (Fig. 30); very doubtful syntypes AMSA C.6988 (not seen). C. edychroa var. violacea: not located. P. gracilis: Holotype in BMNH (ex Macandrew coll.), 3.0x1.1 mm. M. anxia: syntypes AMSA C.27446, 1 syntype NMWC 1955.158.912 (not seen).
Regional locality data: SOUTH AFRICA: KwaZulu-Natal: Kosi Bay, 1-4 km S of estuary, -20-22 m (D. Herbert & K. Bloem; NMSA V2284); off Kosi Bay, -23-24 m (D. Herbert & K. Bloem; NMSA S2936); SE of Kosi Bay, -50 m, fine sand (Kilburn et al.; NMSA E5349); off Kosi Bay, -9-17 m, stone washings (D. Herbert; NMSA D9928); between Kosi Bay and Bhanga Neck, -13 m, sand (D. Herbert; NMSA S2995); same data but -10-12 m, sand (D. Herbert; NMSA S2373); off Boteler Point, -70 m, coral rubble (Kilburn et al.; NMSA E1574); off Island Rock, -62 m, sand and coral debris (Kilburn et al.; NMSA E5562); off Leadsman Shoal, -24-26 m, stone washings (D. Herbert & Natal Parks Board; NMSA E2636); Sodwana Bay, Two-Mile Reef, -10-15 m, stone washings (D. Herbert & R. Broker; NMSA E734); off Richards Bay, -50 m, fine sand, broken shell (CSIR Water Research; NMSA A6102); off Cape St Lucia,-76-80 m, coarse sand (Kilburn et al.; NMSA E5692); SE of Mission Rocks, -50 m, old coral rubble (Kilburn et al.; NMSA E6297); off Durban Bluff, -20-22 m, sand (R. Kilburn & R. Fregona; NMSA B5468); Aliwal Shoal, off Umkomaas, -27 m, silt between rocks (G. Smith; NMSA E1677). Eastern Cape: Port Grosvenor, -82 m, worn coral nodules (Kilburn et al.; NMSA S9578); off Cape Morgan Lighthouse, east of East London, approx. -100 m, broken coral and shell (A. Connell; NMSA B7122).
Distribution: New Caledonia to Queensland, west to Reunion I. and south-eastern Africa, south as far as the East London area, in shallow water to about -100 m.
Remarks: The types listed above have been examined, except for M. anxia, photographs of a syntype of which (Fig. 31) were kindly provided by D. Beechey. The protoconch of a fresh Reunion I. specimen, donated by Dr Maurice Jay, is illustrated here (Fig. 33).
K. edychroa shows some resemblance in teleoconch form to Mangelia lutaria Hedley, 1907, of New South Wales, type species of Exomilus Hedley, 1918. In fact, M. anxia, here synonymised with K. edychroa, was included in Exomilus by Powell (1966). However, the protoconch in Exomilus lutarius, as figured by Laseron (1954: figs 251, 252), has a distinctive, cylindrical, truncate shape, without trace of diagonally cancellate sculpture.
Kermia pumila (Mighels, 1845)
Pleurotoma pumila: Mighels 1845: 23; Johnson 1949: 228, pl. 27, fig. 11 ('neoholotype' = lectotype). Type locality: Oahu [= Kauai, fide Johnson 1949: 217], Hawaii.
Philbertia (Kermia) pumila: Cernohorsky 1978: 162, pl. 57, fig. 14.
Kermia pumila (Mighels): Kay 1979: 362, fig. 118L.
Mangelia digitale Reeve, 1846: pl. 8, sp. 70. Type locality: unknown.
Pleurotoma (Mangelia) reticulata Garrett, 1857: 102 (non T. Brown, 1827, nec Bronn, 1831). Type locality: Hilo Bay, Hawaii. [= Pleurotoma pumila fide Pease 1868; Tryon 1884; Kay 1979]
Pleurotoma clandestina Deshayes, 1863: 110, pl. 12, figs 15, 16. Type locality: Reunion I.
Clathurella violacea Pease, 1868: 218, pl. 15, fig. 15; Johnson 1994: 27, pl. 6, fig. 15 (lectotype). Type locality: Paumotus [Tuamotus], Polynesia.
Type material (examined unless otherwise indicated): P. pumila: lectotype (designated by Johnson 1949) MCZH 176993 (Fig. 34). M. digitale: types probably lost (originally in Stainforth coll.). P. reticulata: not seen. P. clandestina: one syntype in MNHN (a juvenile, badly encrusted by "Byne's disease"). C. violacea: lectotype (Fig. 35), worn (designated by Johnson 1994) MCZH 231971, 4.6x1.8 mm, paralectotypes MCZH 231971 (not seen).
Regional locality data: SOUTH AFRICA: KwaZulu-Natal: between Bhanga Neck and Kosi Bay, near pinnacles of reef, -10-12 m, hand-dredged sand (D. Herbert; NMSA S2391).
Distribution: Polynesia, Hawaii, Fiji and Okinawa to the Mascarenes, Seychelles, and northern KZN, intertidal to about -20 m, under coral and stones.
Remarks: The single South African specimen, in fresh (probably live-collected) condition, appears to be the first record of K. pumila from the African mainland. Only the most important references are cited above. The protoconch of a specimen from Reunion I., collected by Dr M. Jay, is illustrated here (Fig. 38).
[FIGURES 34-38 OMITTED]
Kermia producta (Pease, 1860)
Clathurella producta: Pease 1860: 143; Von Martens & Langkavel 1871: 2, pl. 1, fig. 3; Tryon 1884: 298, pl. 19, fig. 63; Kay 1965: 19, pl. 2, figs 3, 4 (lectotype). Type locality: Sandwich Is [Hawaii]. Kermia producta (Pease): Kay 1979: 361, fig. 118M.
Defrancia mauritiana Sowerby, 1893: 491, pl. 38, figs 23, 24. Type locality: Mauritius. Syn. n.
Clathurina foraminata pyrgodea Melvill, 1917: 186, pl. 10, fig. 13; Trew 1987: 60. Type locality: Mussandam, Persian Gulf, 55 fath. [-101 m]. Syn. n.
Kermia foraminata pyrgodea (Melvill): Powell 1966, pl. 22, fig. 2 (after Melvill 1917).
Type material: Clathurella producta: lectotype (designated by Kay 1965) BMNH 1962761. D. mauritiana: holotype BMNH 18184.108.40.206, don. G.B. Sowerby. C. foraminata pyrgodea: holotype BMNH 19220.127.116.11.
[FIGURES 39-45 OMITTED]
Regional locality data: SOUTH AFRICA: KwaZulu-Natal: between Kosi Bay and Bhanga Neck, -13 m, sand (D. Herbert; NMSA S3006), same locality, ca -8 m (D. Herbert & K. Bloem; NMSA S771); off Boteler Point, -70 m, coral rubble (R. Kilburn et al.; NMSA E5727); Aliwal Shoal, off Umkomaas, -25-28 m (D. Herbert; NMSA S9894). Eastern Cape: Mzamba, beach drift (J.P. Marais; NMSA A5295).
Distribution: Hawaii and Polynesia to Red Sea, Mauritius, Mozambique and eastern South Africa.
Remarks: K. producta belongs to a highly problematic complex of taxa, much confused in the literature. Examination of the relevant types and fresher samples from various localities leads me to propose the synonymy given above. Comparison of types of D. mauritiana (Fig. 40) and C. foraminata var. pyrgodea (Fig. 41) shows them to be conspecific, and Clathurella producta (Fig. 39) to have been based on a slightly worn specimen of the same species. Although D. mauritiana was synonymised with Kermia tessellata (Hinds, 1843) by Hedley (1922), Powell (1966) and Cernohorsky (1978), the extant types differ as follows: those of D. mauritiana (and C. foraminata pyrgodea) differ from K. tessellata (Clavatula tessellata Hinds, 1843: 44, syntype BMNH 1818.104.22.168 ex Lombe Taylor coll.--presumably not the figured type (Hinds 1844: 23, pl. 7, fig. 11) as it is worn and broken) in their shouldered whorls, finer sculpture, flatter outer lip and strongly contracted base (Fig. 48). Two similar but evidently distinct species are Kermia apicalis (Montrouzier in Souverbie, 1861) and Kermia daedalea (Garrett, 1873). Of these, the former, according to the holotype of Pleurotoma apicalis in MHNB (Figs 46, 47), has non-shouldered whorls and a less contracted base than K. producta. K. daedalea is discussed below.
Kermia daedalea (Garrett, 1873)
Clathurella daedalea: Garrett 1873: 219, pl. 2, fig. 33; Tryon 1884: 294, pl. 19, fig. 66. Type locality: Viti Is [= Fiji].
Clathurella phaedra Hervier, 1897a: 141; 1897c: 96, pl. 2, fig. 3. Type locality: Lifou.
Philbertia phaedra (Hervier): Oyama & Takemura 1957: Philbertia (2), figs 13, 14.
Type material: C. daedalea: 3 syntypes from Viti [Fiji] Is in ANSP; of these, ANSP 15821 (Figs 50, 51) resembles the type figure and is here designated as lectotype; it is evidently a specimen of the species later described as C. phaedra; ANSP 15820 comprises two syntypes, one a chalky example of C. phaedra, the other a specimen of Kermia felina (Reeve, 1843). C. phaedra: originally 6 syntypes in SML; one presumed syntype (labelled "paratype", although it does not agree well with the type figure) now in MNHN (Figs 53, 54); another possible syntype AMSA C.7081 (not seen).
Regional locality data: SOUTH AFRICA: KwaZulu-Natal: 1-4 km S of Kosi Bay, ca -20 m (D. Herbert & R. Broker; NMSA S2580); between Bhanga Neck and Kosi Bay, -13 m (D. Herbert; NMSA S3006); Leadsman Shoal, Raggie Reef, -8-14 m, mixed algal and coral reef (D. Herbert & Natal Parks Board; NMSA E5397); off Durban Bluff, -20-22 m, reclamation sand (R. Kilburn & R. Fregona; NMSA B5471); Aliwal Shoal, off Scottburgh, -10 m, sand and reef debris (D. Herbert; NMSA S8227) and off Umkomaas, -25-28 m (D. Herbert; NMSA S9894).
[FIGURES 46-49 OMITTED]
[FIGURES 50-54 OMITTED]
Distribution: New Caledonia to the Gulf of Suez and south to southern KZN.
Remarks: The status of K. daedalea remains to be finally resolved and it may prove to be an extreme variant of K. producta, from which it appears to differ only in its coarser sculpture and sharper lip denticles.
I am indebted to the following curators for providing access to turrid type and other material in their care, or for supplying photographs of these: Dr D.G. Herbert (NMSA); Ms K. Way and Dr D. Reid (BMNH); Dr Ph. Bouchet and Ms V. Heros (MNHN); D. Beechey and I. Loch (digital photographs of types in the AMSA); C.W. Pettit (MMUE); Dr H. Subba Rao (ZSIC); P. Callomon (ANSP), Dr M.G. Harasewych (USNM); A. Baldinger (MCZH). I also thank Mr Tony Bruton of the Electron Microscope Unit, University of KwaZulu-Natal, for the use of the ESEM and Automontage system. Mr Gerald Smith provided material of small turrids dived off KZN. The late Dr Maurice Jay of Reunion Island kindly provided an extensive series of fresh specimens from that island. Finally, I thank referees for useful comments on the manuscript.
[FIGURES 55-56 OMITTED]
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Richard Neil Kilburn
Natal Museum, P. Bag 9070, Pietermaritzburg, 3200 South Africa, and School of Biological &
Conservation Sciences, University of KwaZulu-Natal, P. Bag X01, Scottsville, 3209 South Africa;
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