Annotated type catalogue of Bothriembryon (Mollusca, Gastropoda, Orthalicoidea) in Australian museums, with a compilation of types in other museums

Abstract Type material of 41 Australian Bothriembryon taxa present in Australian museums is critically listed, indicating systematic issues that need to be resolved in further studies. Information on additional type material of 22 taxa in non-Australian museums is compiled. The seven fossil taxa known so far are included in this catalogue. Based on the current systematic position, 38 species are treated in this paper. Bothriembryon jacksoni Iredale, 1939, Bothriembryon notatus Iredale, 1939, Bothriembryon praecelsus Iredale, 1939 and Bothriembryon serpentinus Iredale, 1939 are elevated to species level. Bothriembryon gratwicki (Cox, 1899) is listed as status to be determined.


Introduction
The land snail genus Bothriembryon Pilsbry, 1894, is endemic to Australia but forms part of the Gondwanan element in the superfamily Orthalicoidea. This genus has received little in-depth attention, the last review being B.J. Smith (1992) summarizing all known data. Given the fact that Bothriembryon species are mostly patchy in their distribution, and in some areas species complexes have been identified, a revision of the genus is needed using modern techniques (Breure and Whisson, in preparation). An inventory of as much type material as possible would facilitate revisionary work; data of primary types for 63 taxa are presented herein. The aim of this paper is (1) to present additional data on the types of Bothriembryon in Australian museums (see also Kendrick and Wilson 1975, Wells 1977, B.J. Smith 1992; (2) to summarize recently published data for some European museums (Neubert and Janssen 2004, Köhler 2007, Breure and Ablett 2012, and add additional data from American and European museums; (3) to indicate systematic issues for several species to be clarified by further studies.

Methods
For each taxon the original publication-in which the taxon was proposed-is mentioned, as well as papers in which reference is made to the type material. The type locality is quoted from the original publication in the original wording and language, with clarifying notes between square brackets. These localities have been mapped using SimpleMappr (Shorthouse 2010). The text of the original, or oldest, label is quoted, together with information from subsequent labels if containing information necessary for a correct interpretation. It should be noted that Iredale is considered to be somewhat remiss about marking his type material and that the designations of Iredale's material in the Sydney museum has been done at a later stage by McMichael, presumably together with Iredale (A. Miller, pers. commun.). As a consequence this material has incorrectly been considered as holo-or paratypes by later authors (Wells 1975, B.J. Smith 1992. The original dimensions are quoted, as well as the dimensions of the type specimens; these have been taken with a digital calliper, using the methods figured by Breure (1974: fig. 2) for shell dimensions and Kendrick and Wilson (1975: fig. 1) for whorl counts; measurements up to 10 mm have an accuracy of 0.1 mm, those above 10 mm are accurate to 0.5 mm. Comparing the current measurements to those quoted from the original publication, one should be aware that the diameter especially may have been measured differently. In the case of syntypes, only the largest specimen has been measured. The number of specimens originally available, if quoted by the original author, is mentioned under remarks. Remarks are further given to describe any individual characteristics of the type specimens or any other details of the type lot. The current systematic position is given, following Breure and Romero (in press) for the classification at the family level, and B.J. Smith (1992) for the species level unless otherwise stated.

Systematic list of nominal taxa of Bothriembryon in Australian museums
Fossil taxa are indicated with an asterisk (*).
Remarks. Although the formulation differs, this is the same locality where Bothriembryon gardneri Kendrick, 1978  Remarks. We tentatively follow the opinion of B.J. Smith (1992) that this taxon is synonymous with Bothriembryon mastersi (Cox, 1867). However, a more detailed study should corroborate this view.

Bothriembryon leeuwinensis eventus
Remarks. Iredale (1939) refers to "Another series from Margaret River"; this may refer to the village "Margaret River", or to a location along the Margaret River watercourse. In the latter case it is more likely to be in the coastal area.

Bothriembryon franki
Remarks. The text in Iredale (1939) has led to some confusion with later authors. Under the heading "Bothriembryon fuscus Thiele 1930" Iredale described shells ("a very large series") from Denmark as Bothriembryon franki (see the legend of his pl. 2; not B. fuscus franki as Smith (1992) suggested). The type locality Deep River is ca. 100 km W Denmark in the Walpole area. The shells he referred to as collected by S.W. Jackson at "Deep River, Nornalup Inlet" were suggested to be identical with B. fuscus, which was described by Thiele from Torbay. Wells (1977) erroneously listed the Deep River specimens as types of fuscus Thiele. Smith (1992)  Remarks. Although the formulation differs, this is the same locality where Bothriembryon consors Kendrick, 1978 has been found.
Remarks. According to the old WAM registration book (No. 10127) this material was collected by F.R. Bradshaw at Bluff Knoll. This more specific locality was not mentioned by Iredale (1939); the type locality is now restricted to Bluff Knoll. Iredale did not mention on how many specimens his description was based. The relationships between this taxon and Bothriembryon fuscus Thiele, 1930, B. franki Iredale, 1939 andB. indutus (Menke, 1843)  Remarks. The AM label indicates that this is the specimen figured by Iredale, but shows doubt about the status of the specimen. Iredale (1939) gives several sets of dimensions, but the ones quoted above are for the "type"; see introduction why we regard Iredale's specimens generally as syntypes. The possible subspecific status of this taxon needs further confirmation.
Type material. AM C6376, two syntypes; AM C127559, eight syntypes (ex Cox coll.). Remarks. According to B.J. Smith (1992) the two specimens in lot C6376 are syntypes; they are accompanied by a type written label stating "Type figd P.L.S.N.S.W. XXIV, p. 435 / Isrealite [sic] Bay, W.A. / Pres. Dr. J. C. Cox". These specimens are, however, smaller than Cox stated and both are unlikely to have been figured. The largest specimen of lot C127599 fits the dimensions given by Cox better and matches the figure. These specimens are accompanied by the label mentioned above with Cox' handwriting; a second label read reads "fr: near Israelite Bay / W.A." and also bears the taxon name in the handwriting of Cox. The status of this taxon needs further clarification. The type locality is well known for fossil marine deposits; all museum material was collected as shells only, and the shape of this taxon is aberrant within the genus Bothriembryon, leaving doubt about its systematic classification.
Current systematic position. Status to be determined. Remarks. The AM specimen is labelled "holotype" (not in Iredale's but in McMichael's hand), but does not confirm with the original measurements; moreover, Iredale (1939) states he had more specimens ("shells"). Therefore we consider it a syntype. The possible subspecific status of this taxon needs further confirmation.

Bothriembryon physoides humilis
Remarks. Iredale (1939) based himself on "a series" of shells; we consider therefore all material as syntypes.
Current systematic position. Bothriembryontidae, Bothriembryon irvineanus Iredale, 1939. Remarks. Iredale (1939) did not explicitly state on how many specimens his description was based. This taxon is part of the Bothriembryon kingii species complex that extends from the Walpole area in the west to near Hopetoun in the east. Smith (1992) placed this species in the synonymy of B. kingii (Gray, 1825). However, we are of the opinion that this species group needs further study to ascertain the systematic position of this taxon.
Remarks. The paratypes are-if not from the type locality-from several localities in and around Perth; see Hill et al. 1983 Type material. SAMA D11341, one possible syntype.
Remarks. The specimen is slightly smaller than Cox' original dimensions, and the label is different from that given by Cox (1867). It is considered a possible syntype as there remains some doubt if this shell was part of the original series.

Bothriembryon multispirus
Remarks. This taxon is part of the Bothriembryon kingii species complex that extends from the Walpole area in the west to near Hopetoun in the east. Smith (1992) placed this species in the synonymy of B. kingii (Gray, 1825). However, we are of the opinion that this species group needs further study.
Current systematic position. Bothriembryontidae, Bothriembryon notatus Iredale, 1939. Remarks. Kendrick and Wilson (1975) have discussed the type specimen, clarifying that the original label has been lost and that "the dimensions differ slightly from those given by Cox". Current systematic position. Bothriembryontidae, Bothriembryon onslowi (Cox, 1864). Tate Type locality. [Western Australia] "the mouth of the Moore River". Label. The material is accompanied by three labels. A very thin fragile old paper label with fine ink reads "Moore River, WA"; an old thicker card label with thicker black ink says "mouth of Moore River" and a lined paper label with pencil says "Moore River (mouth of)".

Bulimus indutus pallidus
Dimensions. "The height of the shell is 25 mm., while its breadth is 19 mm."; holotype H 24.2, D 17.9, W 4.9 Type material. WAM S14467, holotype. Remarks. Iredale (1939) mentioned to have "One specimen" available; therefore we consider this as the holotype. The current taxonomic status of the species needs confirmation.

Bothriembryon rhodostomus perspectus
Remarks. Iredale (1939) gave two sets of measurements. The specimen in the AM collection does not confirm to these measurements, but bears an original label with the indication "type"; we consider it a syntype but not the holotype. The possible subspecific status of this taxon needs further confirmation.
Current systematic position. Bothriembryontidae, Bothriembryon rhodostomus (Gray, 1834). Type material. WAM S14468, holotype. Remarks. Iredale (1939) mentioned to have "One specimen" available; therefore we consider this as the holotype. This species, which was synonymized by Smith (1992) with Bothriembryon bulla (Menke, 1843), is now thought to be extinct (DEC 2010 Remarks. This is a fossil species from Tertiary age; no further details are known about the dating of the horizon where the shells were found. "The holotype is an isolated steinkern of an immature shell" (McMichael 1968); the top of the shell is damaged. The paratypes are larger, up to 20.0 mm. The specimens are not accompanied with labels, but Crespin (1974) fig. 12; B.J. Smith 1992: 107.
Remarks. Iredale (1939) had "a series including juveniles" available for his description.

Remarks.
As it is clear that this material was seen by Pilsbry, this material is herein considered as syntypes. It is not clear whether Iredale referred to "Margaret River" as a place or as geographical feature. The latter Margaret River flows near Cape Mentelle into the Indian Ocean; Cape Freychinet-type locality of Bothriembryon sayi (Pfeiffer, 1847)-is ca. 20 km to the south. Current systematic position. Bothriembryontidae, Bothriembryon sayi (Pfeiffer, 1847). Tate Remarks. This material was collected during the Horn Scientific Expedition to Central Australia in 1894. Further type material is present in the Natural History Museum in London (Breure and Ablett 2012).
Remarks. The shell in Fig. 7J is somewhat smaller than Iredale (1939) stated, but agrees with the specimen figured by him; the description is based on "A nice series". This species is now thought to be extinct (DEC 2010).

Bothriembryon rhodostomus wrightianus
Remarks. The top of the specimen figured by Iredale (1939) is damaged; it may be noted that the measurements deviate from the published ones and the description was based on "a fine series". The possible subspecific status of this taxon needs further confirmation.

Compilation of Bothriembryon type material in non-Australian museums
Additional to the data presented above, the following type specimens have been located in non-Australian museums. References to recently figured type specimens have been added; some other taxa are (re-)figured. Taxa are listed as originally published, arranged alphabetically on species name. Remarks. This taxon was placed in the synonymy of Bothriembryon tasmanicus (Pfeiffer, 1853) by Iredale (1937: 313). We tentatively follow his judgement, but further study should clarify the status of this taxon.
Remarks. This taxon was considered a synonym of Bothriembryon bulla (Menke, 1843) by B.J. Smith (1992: 102); it was grouped by Richardson (1995) with B. melo (Quoy and Gaimard, 1832). We here tentatively follow the latter opinion; however, further study is needed to clarify the status of this taxon. Current systematic position. Bothriembryontidae, Bothriembryon melo (Quoy and Gaimard, 1832 Remarks. This taxon is probably of Pliocene age, found at Hobart, Tasmania in Yellow Limestone (Travertine). Kershaw (1987) copied Sowerby's figure and discussed the relationship between this taxon and Bothriembryon tasmanicus (Pfeiffer, 1853). A photograph of the holotype of B. gunnii is here presented for the first time since its original publication.