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Andrefrancia Solem, 1960 (Gastropoda: Pulmonata: Charopidae) – a Systematic Revision

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Andrefrancia Solem, 1960 (Gastropoda: Pulmonata: Charopidae) – a Systematic Revision Vol. 16(3): 101–195 ANDREFRANCIA SOLEM, 1960 (GASTROPODA: PULMONATA: CHAROPIDAE) – A SYSTEMATIC REVISION EWA PAW£OWSKA-BANASIAK Museum of Natural History, Wroc³aw University, Sienkiewicza 21, 50-335 Wroc³aw, Poland (e-mail: [email protected]) ABSTRACT: The revision of the endemic New Caledonian genus Andrefrancia Solem, 1960 is based on 1,167 specimens from 108 localities (collection Paris Museum of Natural History). Seventeen earlier described spe- cies are revised, and 33 new species are described; data on further six species, absent from the Paris collection but included in Andrefrancia, are cited. Identification keys, synonymy, conchological and anatomical descrip- tions and figures, as well as distribution maps are provided. Phylogeny reconstruction used 21 conchological characters with 64 character states, and 26 characters of the reproductive system with 62 character states. The characters were polarised based on out-group comparison (out-groups: Charopidae from the Pacific islands, Australia, New Zealand, Juan Fernandez and Chile; Endodontidae from the Pacific islands) and ontogenetic criterion. The aim of the study was to answer the following questions: (1) is the genus monophyletic? (2) are species groups within the genus monophyletic? (3) what are phylogenetic relationships among the species and their groups? Difficulties resulted from incompleteness of data (only shells available for some species), fre- quent instances of convergent evolution and consequent paucity of unequivocal apomorphies which would define monophyletic groups. As a result the phylogenies provided are only preliminary, and any formal classi- fication changes would be premature. Based on combination of characters the genus was divided into seven groups; most species were assigned to groups distinguished by earlier authors; one new group was proposed. The geographical distribution was analysed in terms of endemism, but the lack of unequivocal phylogeny made it impossible to analyse it in the light of evolutionary relationships. KEY WORDS: terrestrial snails, Charopidae, Andrefrancia, New Caledonia, revision, new species INTRODUCTION New Caledonia, a Melanesian Island in the structure still remain unclear. The literature reflects South-west Pacific, is well known for its high level of knowledge based on material collected mainly endemism and recognised as one of the “hot spots” in around 1850–1900, with scattered additional collect- tropical forest diversity. In regard to terrestrial snails ing until the 1930s. Extensive collections were made it also presents an important opportunity to study the by the staff of the Paris Museum of Natural History in very old part of Gondwanan fauna, that has been 1978–1994, in part within the context of the mu- evolving in increasing isolation since its separation seum’s programme “Evolution and Vicariance in New from the Northern Island of New Zealand. New Cale- Caledonia”. In contrast to the earlier 19th century donia lies at the centre of distribution of the large material, the newly collected material is well localised, family Charopidae which spreads as far east as the So- consists of larger series of specimens, and is mostly ciety Islands and as far north as Indonesia and parts of suited for anatomical work and detailed examination Micronesia. Despite the extensive work on the family, of shell microsculpture. The objective of this study done by A. SOLEM in 1960–1983 (SOLEM 1960, 1961, was a revision of the most speciose New Caledonian 1970, 1976, 1983), the phylogenetic relationships charopid genus – Andrefrancia. amongst some groups, as well as their taxonomic 102 Ewa Paw³owska-Banasiak CHARACTERISTICS OF CHAROPIDAE thin free oviduct section varies in length, the base of spermatheca and the upper section of vagina are nor- Shell mally swollen. The spermathecal container is located The Charopidae are a family of minute to large above the apex of pallial cavity, and ranges from short pulmonate land snails. Their most striking character to long oval. Its shaft tapers abruptly entering the free is a very diverse and often complex apical sculpture of oviduct. The vas deferens is slender, usually entering a the shell. For example, the Pacific island Charopidae clearly differentiated epiphallus, often through a have an apical sculpture primitively featuring broadly complex valvular pore. In many taxa the epiphallus is rounded spiral cords, usually 8–12 in number, without secondarily compressed into penis or absent. The pe- any radial elements. Frequently just before nis is usually stout, at least in its upper section, which teleoconch 1–3 low radial undulations will appear in only rarely is long and slender; the lower part is often the shell surface. In a few species, the number of spi- a thin tube. The interior of penis frequently bears a ral cords has increased to more than 20, and there are well developed verge, circular ridges, stimulatory pi- 10–20 radial undulations. Such a pattern is typical for lasters, and pocket stimulators. The penial retractor Pacific island species. In extralimital areas, however, muscle is usually very short, inserting in a simple or the pattern of apical sculpture is much more com- complex fashion on the penis/epiphallus junction, plex, with Charopidae from Australia and New Zea- the epiphallus or penis with vas deferens piercing the land, for example, showing a variety of types (e.g. muscle. The interior of lower female tract is simple, SOLEM 1970: pl. 58, Figs 3–6). Postapical whorls bear or with very large pilasters which may be simple or major radial ribs and a microsculpture of radial corrugated; the outlets of free oviduct and/or sper- riblets, combined with secondary spiral elements. In mathecal shaft vary from simple pores to apertures many taxa, however, the sculpture has been second- provided with complex valves. arily reduced or lost. The whorl counts and shell shape vary rather widely, the umbilicus ranges from Ecology and distribution widely open to closed. The shell colour is mono- Charopidae exploit the semiarboreal and arboreal chrome, flammulated or tessellated. In several lin- habitats, but are also well represented in the ground eages apertural barriers have evolved, probably inde- stratum. In terms of species, it is the dominant land pendently (SOLEM 1983), but most species have no snail group of New Zealand, New Caledonia, parts of trace of dentiture. Australia (Tasmania, Victoria, less abundant in New South Wales, South Australia, southwestern Western Reproductive system Australia, sparse in the Northern Territory, well repre- Like the shell, the genital system is highly variable. sented in the rain forest areas of Queensland), Lord The ovotestis consists typically of one or two clusters Howe Island, Norfolk Island and Kermadec Island. of long follicles that lie parallel to the outer wall of Approximately 300 species in 70 genera from Austra- the body cavity and occur sequentially. In some taxa lia and Pacific Islands have been described (SOLEM the ovotestis is broken up into a series of lobes that 1983, STANISIC 1990). The Charopidae are only are perpendicular to the outer wall of the body cavity sparsely represented in Indonesia, New Guinea, Solo- and widely separated from each other. The hermaph- mon Islands, and New Hebrides. roditic duct is either straight or coiled, usually an Extralimital to the Pacific Basin, species of irridescent tube entering laterally into the apex of charopids are common to dominant in southern Af- carrefour. The talon is a finger-like projection or swol- rica, and were common on St. Helena. One genus is len head on a short stalk. The albumen gland, of ir- found on the Subantarctic Islands. There is a modest regular shape, is deeply indented by loops of intestine Juan Fernandez radiation and many scattered de- and head of spermatheca. The prostate and uterus scriptions and records from Southern South America, are fused, with partial common lumen, the uterus Andes and Central America. Two species occur in with separate and sequential glandular sections. The western North America. STUDY AREA New Caledonia is a French territory in the Coral Grande Terre. It is a large (16,648 km2) elongated is- Sea in the southwestern Pacific Ocean. It incorpo- land located approximately equidistantly from north- rates the island of New Caledonia, along with numer- ern New Zealand, southeastern Papua New Guinea, ous small offshore islands and reefs, the Isle of Pines, and the east-central coast of Australia (BAUER & the Loyalty Islands, the d’Entrecasteaux Reefs, and SADLIER 2000). The topography of New Caledonia is most distantly the Chesterfield Reefs. The island of dominated by a chain of relatively high mountains New Caledonia itself is sometimes referred to as the that runs along its entire length, rising to over 1,600 m. Andrefrancia Solem, 1960 – a systematic revision 103 The high points are Mt. Panié (1,629 m) in the north diverse than any of the forest or maquis formations. and Mt. Humboldt (1,618 m) in the south but many The vegetation of the Isle of Pines has largely been peaks exceed 1,000 m. disrupted by plantations, but extensive forests remain The climate of New Caledonia is subtropical, with on the east coast. Stands of Araucaria columnaris also temperatures averaging 22–24°C. Trade winds from are patchily distributed throughout the island and the east yield the high orographic precipitation typi- give it its name. Typical maquis vegetation occurs on cal of the east-facing slopes of the island. As a result, the plateau. The vegetation of the Loyalty Islands is the windward eastern coast is mesic, with 2,500–4,000 depauperate relative to mainland New Caledonia. mm of rain per annum, whereas the leeward west Araucaria columnaris stands occur on the southern coast typically receives less than 1,200 mm. The north- coasts of the two largest islands and coastal mangroves western area, including the northern tip of the island, are also present. The island areas support humid for- averages less than 800 mm per annum.
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