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Curaçao and Other STUDIES ON THE FAUNA OF CURAÇAO AND OTHER CARIBBEAN ISLANDS: No. 145. Caribbean Land Molluscs: Bulimulidae I. Bulimulus by A.S.H. Breure (Zoologisch Laboratorium, Utrecht) Page Figure Table Plate INTRODUCTION 2 1 Description 5 1,4, 5b Methods 7 2-3,5a Material 8 Acknowledgments 9 SYSTEMATIcs 10 Genus Bulimulus Leach 10 2 B. limnoides ( 12 3 I, 1-6; VI, 6 B. guadalupensis (Brugui&re) 15 6-60 4 II B. 27 gittenbergerisp. n. 5 V, 10-13;VII, 3 B. hummelincki 28 61-71 6 sp. n. 111,1-5 B. diaphanus (Pfeiffer) 30 — diaphanus(Pfeiffer) 30 7 III, 11-14; VI,2; VII, 2 —fraterculus (Pot. & Mich.) 32 8 III, 6-10; VII, 1 B. lehmanni (Pfeiffer) 35 72-78 9 IV, 9-11; VI, 5 B. Iherminieri (Fischer) 36 79-80 10 V, 5-7 B. fuscus: Guilding 38 11 V, 1-4; VII, 5 B. riisei (Pfeiffer) 39 81-94 12 1,7-13;III, 15-17;VI,3-4;VII,4 B. erectus (Reeve) 42 13 IV, 6-8 B. cacticolus (Reeve) 44 14 IV, 12-15 B. eyriesii (Drouet) 46 15 IV, 1-3 B. dysoni (Pfeiffer) 47 IV, 4-5 B. stenogyroides (Guppy) 48 SubgenusRhinus Albers 49 B. constrictus (Pfeiffer) 49 16 V, 8-9; VI, 1; VII, 6 Zoogeographicalremarks 51 17 REFERENCES 53 TABLES 61 2 INTRODUCTION "The species of this subgenus [Bulimulus s.s.] are the difficult Bulimuli among most to distinguish, being in most and founded cases on slight inconspicuous differences." PILSBRY 36. 1897-8, p. This series of based the papers is on Bulimulidae collected by dr. P. WAGENAAR HUMMELINCK during several trips to the West Indies, and additional from material several museum collections. The species dealt with in this series have previously been treated in part by several authors and in particular monographically by PILSBRY in the Manual of Conchology (second series, vol. 10-12, 14). additionalinformation Thanks to and access to previously unknown it is type material, now possible to present a revision of this family in the Lesser Antilles and Puerto Rico. Sixteen (sub)species are treated, including two new ones: Bulimu- lus gittenbergeri and B. hummelincki. References, (re) description, for each measurements, distribution, as well as figures, are given species. The following new synonymies are proposed: Bulimus nichollsi limnoides Angas = Bulimulus (Ferussac); Thaumastus exilis var. concolor Piaget = Bulimulus guadalupensis (Bruguiere); Bulimulus Houelmontensis Crosse = Bulimulus diaphanus fraterculus (Potiez & Michaud); Bulimulus semicinctus Pilsbry = Bulimulus Iherminieri Pfeiffer Bulimulus (Fischer); Bulimus Barbadensis = fuscus Guil- ding. Finally some zoogeographical remarks are given. The Bulimulidaeform a relatively large family, mainly confined South At to America. present the family includes 144 genera and The of subgenera. number specific and subspecific names available is estimated at about 3000. According to BAKER (1956) the family name should be Orthalici- dae, ifpriority rules are applied. No attention has been paid to this proposal because of the unstable state of taxonomy of this family. At in which present several genera are included this family, may 3 better be elsewhere. The in placed genus Aillya Odhner, occurring Cameroon (West Africa), is placed in the Amphibuliminae by ODHNER (1928) on account of the anatomy. BAKER (1955) placed the in Aillyidae the Heterurethra*, near the Succineidae. Re-examination of the type species, Aillya camerunensis Odhner, should clarify the status of this genus. African Another genus included in the Amphibuliminae is Presto- nella Connolly. It occurs in South Africa and its anatomy is un- known. BAKER that it also be heterurethroussnail suggests may a (SOLEM 1959a, p. 123). WEYRAUCH (1967) described a monotypical genus from Peru, Pseudoglandina. The morphology of the shell suggests that it belongs to the Bulimulidae, but from a note of the collector (in WEYRAUCH), mr. J. M. SCHUNKE, it might be concluded that the species is a predator, and perhaps should be included in the Oleacinidae. The anatomy of Pseudoglandina agitata is unknown and the present status of the genus uncertain. On conchological, ecological and zoogeographical grounds WEY- Geoceras RAUCH (1958) removed the genus Pilsbry from the Bulimu- lidae the As the taken into to Urocoptidae. anatomy was not con- sideration, a study on this subject may be worthwhile. The Bulimulidae s.l. (see below) are here considered to be a family, be although it still has to proved that they form a monophyletic unit. The family is not easily characterized because of the great variationin shell form (see ZILCH 1959-1960, fig. 1668-1814); usual- the shells ovate-conic The be ly are to cylindric. genitalia seem to simple, but VAN MOL (1971) has shown that the internal structure, especially of the penis and epiphallus, is quite complex. The subdivision of the family into Bulimulinae, Amphibuliminae, Placostylinae, Odontostominae, and Orthalicinae is mainly based on shell features. This sensu lato conception of the Bulimulidae was already held by PILSBRY {Man. Conch., 1895-1902) and THIELE (1931), and is also favoured by the present author. More recent authors, e.g. ZILCH (1960), gave family rank to the subfamilies, the Placostylinae excepted. However, the differences between the sub- families the differ from the are slight, e.g. Amphibuliminae seem to 4 Bulimulinae in the and the free muscle only palleal organs retractor system (see VAN MOL 1971). VAN MOL (pers. comm.) is of the opinion be because that they should not even separated on subfamily level, the organ systems show progressive transformations. The distribution of the Bulimulidae is discussed and figured by PILSBRY 1946, SOLEM 1959b (fig. 15), and PARODIZ 1969 (map 6). In 1 number of in Table the (sub)genera the various areas is sum- marized for the Bulimulinaeand Placostylinae [= Bulimulidaes.s.]. The latter subfamily is entirely confined to the Australian Region. Regarding the Bulimulinae, the marked endemism in Peru, Brasil and Ecuador is endemic very striking. Comparative many (sub)genera occur in Mexico. Fossil records of Bulimulidae s.l. are available from Patagonia, Argentina (Paleocene and Eocene, PARODIZ 1946), Brasil (Miocene, PARODIZ 1969, or Early Paleocene - Late Cretaceous, FERREIRA & SANTOS COELHO 1971), and Florida (Miocene, DALL 1890). The Buli- mulidae, originally autochthonous in South America have migrated since the Miocene (or Pliocene?) towards Central America, the West Indies and North America. [For details on migration of Tertiairy mollusk faunas, see PARODIZ 1969, p. 189 ff.] Shells vary from approx. 10-150 mm, the largest to be found in the the of this the genus Placostylus. In taxonomy family sculpture of the protoconch is used mainly at (sub)generic level; other types of axial fine knobs sculpture (spiral striae, riblets, or costulae) mainly at species level. The majority of the species have no apertural den- tition, but most Odontostominae and some species of Bulimulidae have denticles lamellae in the s.s. one or more and/or aperture. Anatomical data are few. The majorfeatures in the anatomy of the Bulimulinae male MOL are: a long tract, simple externally, internally complicated (VAN 1971, fig. 13-17); a moderately long spermathecal duct, which is unbranched; the free muscles the columella. retractor are united near junctionon Authors in the past generally only considered jaw, radula and gross anatomy of the genitalia (e.g. BINNEY 1884, FISCHER & CROSSE 1870-1902, SEMPER 1874). More recently additional features have been taken into account (e.g. LANZIERI & REZENDE 1965, PILSBRY 1901-1902, VAN MOL 1971). The Bulimulidae are adapted to various kinds of habitat (e.g. trees, rock-faces, 5 shrubs and which litter), even can be occupied by species of the same genus (cf. the radiation of adaptive Naesiotus on the Galapagos). and of Data on the taxonomy zoogeography the family are widely scattered in literature. Only PILSBRY (1895-1896 . 1901-1902) has made a monographic attempt, but this great work is badly in need of revision. Summarizing publications of parts of the family, in a taxonomical or geographical sense, are those of MARTENS 1873, 1890-1901, PARODIZ 1942, 1943, SIMPSON 1929, and STREBEL 1909. DESCRIPTION Although PILSBRY often presented valuable descriptions in his Manual of Conchology, the present author preferred giving a re- description of each species in order to facilitate comparison. All descriptions are based on material examined, unless otherwise stated. References but are as complete as possible, generic synonymy has been omitted. For each species localities only have been men- tioned if they are new to science. Misidentifications are only indicated by a question-mark if the reference obviously falls outside the distribution of the species, as in most cases it is impossible to trace the materialwhich the authorhad at hand. References without that the of the further details mean only name species was given. In the distributionalrange following each description new locali- ties are followed by an exclamation-mark. If a locality is followed by reference material from a no that particular island was examined. Among the terms used in this series, some need definition, while others are already well defined ( fide Cox 1955, BROOKES KNIGHT al. et 1960). Highly subjective and indefinableones remain e.g. form of the aperture and colour. PROTOCONCH: The initialwhorls formedbefore hatching. - In this family these whorls are smooth or variously sculptured, a feature used for identification the level PILSBRY commonly on genus (cf. but in the it also be useful the 1896), present study proved to on species level. LOWER SIDE OF THE PROTOCONCH: The abapical part of each protoconch whorl (Fig. 4). 6 Fig. 1. Sketch map of the Caribbean, showing localities of West Indian Bulimulus treated in this paper. UPPER The of SIDE OF THE PROTOCONCH : adapical part each pro- toconch whorl (Fig. 4). COLUMELLAR MARGIN (c.m. in Fig. 5b): The adaxial part of the peristome, running from the base of the columella to the junction at the ultimatewhorl.
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