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Turridae (Mollusca: Gastropoda) of Southern Africa and Mozambique

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Turridae (Mollusca: Gastropoda) of Southern Africa and Mozambique Ann. Natal Mus. Vol. 29(1) Pages 167-320 Pietermaritzburg May, 1988 Turridae (Mollusca: Gastropoda) of southern Africa and Mozambique. Part 4. Subfamilies Drilliinae, Crassispirinae and Strictispirinae by R. N. Kilburn (Natal Museum, Pietermaritzburg) SYNOPSIS 71 species (42 previously undescribed) are covered: Drilliinae (33 species), Strictispirinae (3), Crassispirinae (35). New genera: Orrmaesia, Acinodrillia (Drilliinae); Inkinga (Strictispirinae); Naudedrillia, Nquma, Psittacodrillia, Calcatodrillia, Funa (Crassispirinae). New species: Drilliinae: Acinodrillia viscum, A. amazimba; Clavus groschi; Tylotiella isibopho, T. basipunctata, T. papi/io, T. herberti, T. sulekile, T. quadrata, Agladrillia ukuminxa, A. piscorum; Drillia (Drillia) spirostachys, D. (Clathrodrillia) connelli; Orrmaesia dorsicosta, O. nucella; Splendrillia mikrokamelos, S. kylix, S. alabastrum, S. skambos, S. sarda, S. daviesi. Crassispirinae: Crassiclava balteata; Inquisitor nodicostatus, /. arctatus, I. latiriformis, I. isabella, Funa fraterculus, F. asra; Naudedrillia filosa, N. perardua, N. cerea, N. angulata, N. nealyoungi, N. mitromorpha; Pseudexomilus fenestratus; Ceritoturris nataliae; Nquma scalpta; Haedropleura summa; Mauidrillia felina; Calcatodrillia chamaeleon, C. hololeukos; Buchema dichroma. New generic records: Clavus Montfort, 1810, Tylotiella Habe, 1958, Iredalea Oliver, 1915, Splendrillia Hedley, 1922, Agladrillia Woodring, 1928 (Drilliinae); Paradrillia Makiyama, 1940 (Strictispirinae); Crassiclava McLean, 1971, Pseudexomilus Powell, 1944, Haedropleura Monterosato in Bucquoy, Dautzenberg & Dollfus, 1883, Mauidrillia Powell, 1942, Buchema Corea, 1934, Ceritoturris Dall, 1924 (Crassispirinae) . New species records: Iredalea exilis (Pease, 1868), Clavus unizonalis (Lamarck, 1822) (DriIliinae); Paradrillia melvilli Powell, 1969 (Strictispirinae); Funa layloriana (Reeve, 1846), Turridrupa cincta (Lamarck, 1822), T. acutigemmata (E. A. Smith, 1877) (Crassispirinae). Reclassification: Paradrillia is transferred from the Clavinae to the Strictispirinae, Turridrupa, Ceritoturris, Haedropleura and Mauidrillia from the Clavinae to the Crassispirinae, Pseudexomilus from the Daphnellinae to the Crassispirinae; Drillia signa Bartsch, 1915 (based on a teratological Clionella or Clavatula) to the Clavatulinae. New combinations: Pleurotoma inclinata Sowerby, 1893, to Iredalea; P. burnupi Sowerby, 1897, P. hottentota E. A. Smith, 1882, and Drilliafalcicosta Barnard, 1958, to Tylotiella; Drilliafalsa Barnard, 1958, and D. eva Thiele, 1925, to Splendrillia, Pleurotoma? paula Thiele, 1925, to Acinodrillia (Drilliinae); Drillia laterculoides Barnard, 1958, and Pleurotoma layloriana Reeve, 1846, to Funa; Pleurotoma layardi Sowerby, 1897, Drillia omia Barnard, 1958, and Clavatula halislrepta Bartsch, 1915, to Crassiclava; Drillia praetermissa E. A. Smith, 1904, to Naudedrillia; Pleurotoma rousi Sowerby, 1886, to Nquma; Cythara ima Bartsch, 1915, to Haedropleura; Pleurotoma diversa E. A. Smith, 1882, P. bairstowi Sowerby, 1886, and Dril/ia albonodulosa E. A. Smith, 1904, to Psittacodrillia (Crassispirinae). New synonyms: Drillia distincta Thiele, 1925 = Drillia (Dri/lia) ligna ria (Sowerby, 1903); Drillia sowerbyi Turton, 1932 (non Reeve, 1843) = Crassiclava hottentota (Sowerby, 1897); Haedropleura dora Thiele, 1925 = H. ima (Bartsch, 1915); Vexitomina Powell, 1942 = Paradrillia Makiyama, 1940. Types figured: Holotypes of Drilliafalsa Barnard, 1958, Pleurotoma castanea Reeve, 1845, P. layloriana Reeve, 1846, P. hottentota Smith, 1882, P. variabilis Smith, 1877, Surcula macilenta Melvill, 1923 (non Solander, 1766) [= Turris macella Melvill, 1923]; Drillia signa Bartsch, 1915, Mangilia benjamini Bartsch, 1915. Lectotypes of Drillia omia Barnard, 1958, D. falcicosta Barnard, 1958, D. laterculoides Barnard, 1958, D. pusilla Garrett, 1873, Pleurotoma diversa Smith, 1882, P. /ignaria Sowerby, 1903. Syntypes of Drillia exilis Pease, 1868, Pleurotoma inclinata Sowerby, 1893, Mangilia decaryi Dautzenberg, 1932. Nomina dubia: Dril/ia pecus Barnard, 1969; D. signa Bartsch, 1915; D. neptuni Turton, 1932. 167 168 ANNALS OF THE NATAL MUSEUM, VOL. 29(1) 1988 Radulae figured: Tyloliella basipuncta/a, Clavus unizonalis, Iredalea exilis, Drillia (C1athrodrillia) connelli, Orrmaesia dorsieosta lDrilliinae); Paradrillia melvilli (Strictispirinae); TlIrridrupa cillCla, T. bijllbata, Inquisitor areta/us, I. nodieostatllS, I. isabella, I. latiri/ormis, FUfla latercllioides, F. taylorianum, F. asra, Naudedrillia praetermissa, N. filosa, N. flealyoullgi, N. milromorpha, Crassiclava halistrep/a, Nquma sealpta, N. rousi, Caleatodrillia ehamaeleon, Mauidrillia /elina, Haedropleura ima, H. summa, Crassopleura maravignae. INTRODUCTION The present study deals for the most part with genera united by Powell (1966) in the subfamily Clavinae. These share a similar shell form, in that they all exhibit a high spire, a relatively short, truncate base, and (with few exceptions) a parietal pad or nodule that serves to buttress the extreme posterior end of the labrum (which would otherwise be weakened by the anal sinus). This 'clavine facies' seems to have evolved as an adaptation to a reef or under-rock existence, as opposed to the produced siphonal canal and non-reinforced labral extremity (the 'turrine facies') of the predominantly sand-dwelling Turrinae and Cochlespirinae. Within the Clavinae of Powell occur a number of radula types that McLean (1971a) demonstrated to be indicative of polyphyly. One group, with true toxoglossate dentition, has already been dealt with (Kilburn 1986) under the Borsoniinae. The remainder of the Clavinae were divided by McLean into five subfamilies, three of which prove to occur in southern Africa and Mozambique. Subsequently Cernohorsky (1985) showed that the name Clavinae was invalidated by homonymy, the subfamilial name Drilliinae being available in its stead; consequently the term 'clavine' or 'claviform' will be used here as a convenient descriptive term for the shell facies described above. In this sense, 'clavine' is not intended to convey any taxonomic implications except in so far as the term applies loosely to all three subfamilies covered here. A valuable account of the structure and functional morphology of the different radula-types found in the Turridae was given by Shimek & Kohn (1981). The observations of Maes (1983) have modified some of their conclusions. Of the three 'clavine' subfamilies occurring in the southern African region, the Drilliinae is characterised by its broad, pectinate lateral plates (Figs 6-11). Typically with a radula formula of 1-1-R-1-1, this is certainly the most primitive subfamily, although the term 'prototypic', applied to its radula by Powell (1966), does involve an untestable assumption. This style of radula is here termed 'drilliine'. Although hypothetically considered to be adapted for 'slicing-rasping' by Shimek & Kohn, Maes (1983) demonstrated that it was used solely for gripping (and/or pricking) the prey, which is swallowed intact. In the two other subfamilies, the radula is (with few exceptions), reduced to two rows of marginal plates. In the Crassispirinae (Figs 39-56) each plate is buttressed by an accessory limb, enabling the radula to be used in pricking and slashing ingested polychaete prey for the penetration of venom (and possibly lytic enzymes). In the third subfamily, the Strictispirinae, an accessory limb is fused to each marginal tooth, its distal end being recurved to form a median flange or collar (Fig. 17). This group is also characterised by the loss of the venom gland and bulb (Maes 1983), but the provision of strong circular muscles around the foregut and a large odontophore indicate that the ingested prey is ripped apart with the aid of the solid, hooked radula teeth. KILBURN: TURRJDAE PART 4 169 The 'clavine' genera are the most difficult of the Turridae to classify and characterise, particularly as some genera are still known only from shell characters. The latter taxa can only be classified by an estimate of their apparent affinities to genera whose radula has been examined. This also applies to the large number of southern African species that have not yet been collected alive. Reassessment of relationships will prove inevitable as material becomes available. A master-key to all the c1avine genera based on shell characters has not proved feasible. Within subfamilies I have given keys to genera, although these are of doubtful value, as routine examination of radulae for identification purposes is not a practical option. Consequently, figures are grouped together as an additional, pictorial aid to identification. Fossil species Two species of 'c1avines' are on record from the Miocene of Zululand, 'Drillia' curiosa King (1953: 84, text fig. 10) and Crassispira (Inquisitor) coxana King (1953: 85, text fig. 5). From the sketchy figures provided it is impossible to judge whether these are even Turridae. The types (now in the Geological Survey, Pretoria, pers. comm. Dr M. Cooper) will have to be studied afresh. Drillia tempestae Kensley & Pether, 1986: 195, fig. 31) from the early Pliocene of Namaqualand, lacks a c1avine anal sinus, and is probably a Clionella (subfamily Clavatulinae). Biogeography in southern Africa The subfamilies Drilliinae and Crassispirinae are too poorly known to permit anything but a simplistic analysis of their biogeography in southern Africa and Mozambique. Furthermore, such results will be heavily biased, as the samples available from the east coast (and to a lesser
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