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Camaenid Land Snails from Western and Central Australia (Mollusca: Pulmonata: Camaenidae) Iv Taxa from the Kimberley, Westrau'rachia Iredale, 1933 and Related Genera

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Camaenid Land Snails from Western and Central Australia (Mollusca: Pulmonata: Camaenidae) Iv Taxa from the Kimberley, Westrau'rachia Iredale, 1933 and Related Genera Rec. West. Aust. Mus., Suppl. no. 17,1984 CAMAENID LAND SNAILS FROM WESTERN AND CENTRAL AUSTRALIA (MOLLUSCA: PULMONATA: CAMAENIDAE) IV TAXA FROM THE KIMBERLEY, WESTRAU'RACHIA IREDALE, 1933 AND RELATED GENERA ALANSOLEM* [Received 9 February 1983. Accepted 22 July 1983. Published 30 June 1984.] INTRODUCTION This is the fourth report on the semi-arid zone dominant land snails of Western and central Australia, which belong to the Camaenidae, sensu lato. It reviews 29 species level taxa in five genera. Twenty-two species and three genera are new. Most belong to Westraltrachia Iredale, 1933, a genus restricted to the south fringes of the Kimberley in limestone hills between Hawkstone Creek, a western tributary of the Meda Riv:er, and the Lawford Ranges east ofFitzroy Crossing. Six of the 21 taxa were previously known and 15 are new. Several related genera from the East Kimberley and adjacent portions ofthe Northern Territory contain eight species. New taxa are Mouldingia (one species in the Napier Range, one near Lake Argyle); Ordtrachia (four species between Lake Argyle and Nicholson); and Prototrachia (one species from east of Timber Creek Police Station, Northern Territory). Exiligada negriensis (Iredale, 1933), lives on the southern edges of Lake Argyle and ranges south about to the level of Nicholson. A total of12,292 adult specimens were measured: 9,787 belong to Westraltrachia and 2,505 to the related genera. Several very large sets were not measured to save time, and thus more than 14,000 specimens were studied. Part I (Solem, 1979) ofthis series covered genera with trans-Australian distributions (Hadra Albers, 1860; Xanthomelon von Martens, 1860; Damochlora Iredale, 1938; Torresitrachia Iredale, 1939), plus related Chloritis-like genera from eastern Australia. Part II (Solem, 1981a) monographed 28 species-level taxa of Amplirhagada Iredale, 1933, a genus whose centre ofdiversity is in the north-west Kimberley, but with species as far south as the King Leopold and Napier Ranges, and as far east as near Wyndham. Part III (Solem, 1981b) described 19 new species-level taxa in four new genera (Ningbingia, Turgenitubulus, Cristilabrum, and Prymnbriareus) from the Ningbing Ranges north of Kununurra and El Questro Homestead south-west of Wyndham, Western Australia. *Field Museum of Natural History, Chicago, Illinois 60605-2496, U.S.A. 427 Only a few more genera from the Kimberley remain to be monographed. It is anticipated that Part V will complete this section of the study, and permit biogeographic analysis of the Kimberley land snail fauna. The recent account of Bishop (1981) on the biogeography of Australian land snails merely outlined problems for study and offered little beyond a summary of classic literature. Major financial sponsorship ofthis co-operative project between the Western Aust­ ralian Museum, Perth (hereafter WAM), and Field Museum of Natural History, Chicago (hereafter FMNH), has been provided by National Science Foundation grants DEB 75-20113, DEB 78-21444, and DEB 81-19208 to FMNH for fieldwork and subsequent study of collected materials, and National Science Foundation grant BMS 72-02149 that established a scanning electron microscope laboratory at FMNH. Grate­ ful acknowledgement is made of this support. Continuing contributions by Mrs. Arthur T. Moulding that permitted extending both field time and sampling into new areas have improved both the depth and breadth of coverage in these reports. These significant contributions are recognised in the dedication of the new genus Mouldingia. Nearly all anatomical and most ofthe shell line illustrations are by Linnea Lahlum, Illustrator, Division of Invertebrates. Additional illustrations are by Elizabeth A. Liebman and Marjorie M. Connors, formerly illustrators, Division of Invertebrates. Maps (Figs 159-162, 179) of collecting localities were drafted by Linnea Lahlum and Elizabeth Lizzio, volunteer Illustrator, Division of Invertebrates. For mounting and labeling the many plates and figures, we are deeply indebted to Dorothy Karall, Associate, Division of Invertebrates. Margaret Baker, Custodian of Collections, Div­ ision of Invertebrates, helped with both specimen measuring and data analysis, and later prepared specimens for return shipment to Perth. Manuscript typing, assistance with proof-reading, and aid with preparation of specimen lists was handled with great efficiency by first Valerie Connor-Jackson and then Patricia Johnson, Secretary, Division of Invertebrates. For field assistance during these surveys, I am indebted to Laurie Price, Field Associate, Division of Invert­ ebrates; Carl C. Christensen, formerly University of Arizona; Fred and Jan Aslin, Mt. Gambier, South Australia; Laurel Keller, peripatetic; and Barbara Duckworth, Australian Museum, Sydney. Data on the materials collected pre-1920 and studied by Iredale (1939) are presented below under 'Previous studies and nomenclature' of the genus Westraltrachia. Activi­ ties by the staff of WAM and my grant supported field surveys provided the bulk of studied materials. As is true for so many taxa ofWestern Australian land molluscs, the field efforts of WAM staff members George W. Kendrick and A.M. Douglas in the mid-1960's produced the first known specimens of several species. These were of infinite help in planning the main field surveys and offering evidence that such efforts would be productive. Barry R. Wilson and Shirley Slack-Smith made highly significant collections in both the Napier and Oscar Ranges during the mid-1970's. To save space in the specimen lists, dates ofcollecting generally have been omitted, but the following inclusive station numbers identify the month and year oftheir fieldwork in the Napier (August 1975 Stas. NRI-XXIV; May 1976-Stas. NRII-1-31) and Oscar Ranges (early September 1975 - Stas. OR I-IV). Activities ofFMNH representatives through­ out the range of Westraltrachia and related genera (Alan Solem, Laurie Price, Carl C. Christensen, Fred and Jan Aslin, Laurel E. Keller, and Barbara Duckworth) were 428 concentrated in two periods-October.1976 to mid-May 1977 (Stas. WA-189 through WA-402) and May to late June 1980 (Stas. WA-567 through WA-600, WA-707 through WA-717). Collecting dates are provided when necessary to discuss aspects of repro­ ductive seasonality and for illustrated specimens, but for space reasons they are omitted from the lists of specimens and types. The genera reviewed here form an apparently monophyletic assemblage character­ ised by complex specialisations primarily in the terminal male genitalia. Variation is at least partly mosaic, in that conchological specialisations do not correlate with anatom­ ical specialisations. The genus with the simplest genitalia, Mouldingia (Figs 164,165), has the most specialised shell (Figs 163a-f), whereas the genera with the most complex genitalia, Ordtrachia (Figs 167, 168, 170, 173) and Exiligada (Fig 175), range the gamut from specialised to very generalised shell features. Prototrachia, known from an isolated limestone outcrop in the Northern Territory, has a very generalised shell (Fig. 176), and genitalia (Figs 176, 177) that combine generalised and specialised features. The probable directions of character change and reasons for the suggested polarities are reviewed below, and their implications are summarised in the biogeographic discussion of this complex. LIST OFTHETAXA Genus Westraltrachia Iredale, 1933 (+ Parrhagada Iredale, 1938 and Zygotrachia Iredale, 1939): 431 W woodwardi (Fulton, 1902) ( + sedula Iredale, 1939 and detecta Iredale, 1939): 490 W commoda (Iredale, 1939) ( + ferrosa Iredale, 1939): 501 W turbinata sp. nov.: 511 W inopinata sp. nov.: 521 W froggatti complanata subsp. nov.: 531 W froggatti froggatti (Ancey, 1898): 535 W derbyi (Cox, 1892) (+ derbyana Smith, 1894 and orthocheila Ancey, 1898): 540 W alterna Iredale, 1939 ( + increta Iredale, 1939): 554 W oscarensis (Cox, 1892) (+ inconvicta Smith, 1894 and perca Iredale, 1939): 563 W cunicula sp. nov.: 570 W subtila sp. nov.: 575 W instita sp. nov.: 580 W lievreana sp. nov.: 584 W tropida sp. nov.: 588 W porcata sp. nov.: 592 429 W rotunda sp. nov.: 593 W limbana sp. nov.: 602 W recta sp. nov.: 608 W pillarana sp. nov.: 613 W ascita sp. nov.: 618 W ampla sp. nov.: 624 Genus Mouldingia new genus: 635 M. occidentalis sp. nov.: 638 M. orientalis sp. nov.: 644 Genus Ordtrachia new genus: 647 0. septentrionalis sp. nov.: 650 0. australis sp. nov.: 655 O. grandis sp. nov.: 658 O. intermedia sp. nov.: 666 Genus Exiligada Iredale, 1939: 670 E. negriensis Iredale, 1939 ( + qualis Iredale, 1939): 673 Genus Prototrachia new genus: 681 P. sedula sp. nov.: 687 SYSTEMATIC REVIEW The genera reviewed here show a coherent ecological pattern of association with exposed limestone, aestivation as free sealers on the soil surface (most genera) or burrowed into sandy soil (Exiligada), and apparent tendency to feed at least opportunistically on seepage face floral blooms. Mouldingia, with a disjunct distri­ bution consisting ofa few hills near the south-west corner ofLake Argyle and then near McSherry Gap in the Napier Range, combines relatively simple genital structures and highly complex shell features. The most speciose genus, Westraltrachia Iredale, 1933, has a unique genital structure: the epiphallus extends into a 'U'-shaped loop, and there is an extension ofthe penis retractor muscle that reaches the penis head to form a penis muscle. Usually the arms of the epiphallic loop are bound together by muscle fibers. In some species of Westraltrachia heavy
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