An annotated catalogue of type specimens of the land snail genus Cyclophorus Monfort, 1810 (Caenogastropoda, Cyclophoridae) in the Natural History Museum, London

Abstract The collection of land caenogastropod snails in the genus Cyclophorus Monfort, 1810 housed in the Natural History Museum, London (NHM), includes 52 type lots. Lectotypes have been designated for 43 available species-level names to stabilize existing nomenclature, two previously designated lectotype, two holotypes, one paratype, one syntype, one possible syntype and two paralectotypes are also listed. A complete catalogue of the Cyclophorus types in NHM, London is provided for the first time.


Introduction
The Cyclophoridae Gray, 1847 are a family of caenogastropod land snails with a fossil history dating back to the Early Tertiary (Gordon and Olson 1995). Extant cyclophorids are distributed in Africa, Asia, Australia, Southern Europe and various Pacific islands (Ko-Type material. Lectotype (design. n.), NHMUK 20130115 ( Fig. 4D; D=42.8 mm, H=35.5 mm, W=5).
Remarks. This species was described based on specimens from the Cuming collection. In the original description only one set of specimen measurements was given. In 1853, Pfeiffer (1853a: 244, pl. 32, figs 11-13) re-described and illustrated a single specimen from the Cuming collection. There is a single specimen from the Cuming collection with an original label in Pfeiffer's handwriting stating the species name. The figured specimen in Pfeiffer (1853a) closely matches the measurements given in the original description and is here designated as the lectotype.

Cyclophorus excellens
Remarks. This species was described based on material from the Cuming collection, and only one set of shell measurement was given. Later, Pfeiffer (1854e: pl. 4, figs 1, 2) re-published the description and figured a shell from the Cuming collection. Two shells from Cuming's collection with an original label in Pfeiffer's handwriting giving the species name are in the NHM collections. The specimen which most closely matches the measurements given in the original description and the illustration in Pfeiffer (1854e) is here designated as the lectotype.
Cyclophorus expansus (Pfeiffer, 1853) http://species-id.net/wiki/Cyclophorus_expansus Remarks. The original description did not include an illustration or collection locality. Pfeiffer subsequently (1854a: 293, pl. 39, figs 20, 21) re-published the description and figured the species. There are three shells from Cuming's collection with an original label in Pfeiffer's handwriting. The specimen which most closely matches with the measurements given in the original description and the illustration in Pfeiffer (1854a) is here designated as the lectotype. Remarks. The original description by Pfeiffer did not give an illustration of the species or a collection locality. Pfeiffer (1854a: 354, pl. 45, figs 9, 10) re-published the description and figured the species. Three shells from Cuming's collection have an original Pfeiffer label giving the species name and a collection locality of "Arva", that could perhaps be in Pfeiffer's handwriting, although a later label states: "Arva'' added to label not by Pfeiffer? In addition there is a separate label with the specimens, also possibly in Pfeiffer's handwriting, stating: 'Prame Pegu'. The figured shell from Pfeiffer (1854a: pl. 45, figs 9, 10) with an "x" written in the aperture is here designated as the lectotype but the type locality remains uncertain. Remarks. Godwin-Austen stated that he received three specimens from Mr. Fulton. Only one specimen from Hugh Fulton could be located in the NHM collections. The specimen has an original Godwin-Austen label stating "Type" and the shell closely matches with the measurements given in the original description. It is here designated as the lectotype. Remarks. The original description stated "in some specimens", which implied that this taxon was based on more than one specimen. There are two specimens from the Godwin-Austen collection with "Type" written on the original label. The specimen figured in the original description is here designated as the lectotype. Remarks. This species was based on more than one specimen, from Theobald's collection but only one set of measurement was given. The original description did not include an illustration but subsequently, Hanley and Theobald (1870) figured three illustrations of this species. There are two shells in the NHM collections purchased from W. Theobald, with an original label stating "typical", and with the collection locality "Moulmein". The specimen that most closely matches the measurements given in the original description, and the illustration in Theobald (1870: pl. 1, fig. 3  Remarks. This species was described from specimens in the Cuming collection and only one set of shell measurements was given in the original description. Pfeiffer (1853a: 247, pl. 33, figs 10, 11) republished the description and figured a shell from Cuming's collection. There is a single shell in the NHM collections from the Cuming collection with an original label stating "the type" which exactly matches Pfeiffer illustration (1853a) and is here designated as the lectotype. Type material. Lectotype (design. n.), NHMUK 20130081/1 ( Fig. 11A; D=23.0 mm, H=17.7 mm, W=5), paralectotype NHMUK 20130081/2 (1 shell; Fig. 11B; D=22.0 mm, H=17.9 mm, W=5).

Cyclophorus haughtoni
Remarks. This species was described from specimens in the Cuming collection and only one set of shell measurements was given in the original description. Pfeiffer (1854a: 349, pl. 45, figs 19, 20) republished the description and figured a shell from Cuming's collection. There are two shells in the NHM collections with an original label in Pfeiffer's handwriting giving the species name and original collection locality. The specimen which matches the illustration in Pfeiffer (1854a) and the dimensions given in the original description is here designated as the lectotype. Remarks. The original description does not include an illustration but later, Bavay and Dautzenberg (1909: 285, 286, pl. 9, figs 5-7) republished the description and included illustrations of the species. Fischer-Piette (1950: 176) wrote the "holotype, 37 mm" which we consider to be an inadvertent lectotype designation (ICZN 1999: Art. 74.5). The lectotype is housed in the Muséum National ďHistoire Naturelle, Paris. The NHM specimen from the R.B. Lucas collection (purchased from Dautzenberg) has an original label stating "Type" and giving the collection locality "Muong-Bo" and is considered to be a paralectotype. Further paralectotypes are housed in the Royal Belgian Institute of Natural Sciences, Brussels.
Remarks. There are two shells in the NHM collections with Smith's handwriting on the original label. One specimen has a small label with "Type" written on it attached inside the aperture. This specimen corresponds to the figured specimen and the measurements given in the original description and is here designated as the lectotype. Remarks. Godwin-Austen's description was based on five specimen lots with figures of shells from different lots provided in the original description. Three lots were listed as being housed in the NHM, and two lots in the Indian Museum, Calcutta. Each of the three NHM has original labels in Godwin-Austen's handwriting stating species name, collection locality and catalogue numbers. The figured specimen from the lot 1903.7.1.3579 labelled "cotype" is here designated as the lectotype. ( Remarks. This species was described from material in the Cuming collection, and the original description does not include an illustration. Later, Reeve (1861: sp. 32) re-described the species and illustrated a shell from the Cuming collection. There is a single shell in the NHM collections from the Cuming collection with an original label in Pfeiffer's handwriting. This shell is matches the measurements given in the original description and figured in Reeve and is here designated as the lectotype. Remarks. The original description included three varieties indicated with "var. a", "var. b" and "var. c". Sowerby I subsequently published Thesaurus Conchyliorum (Sowerby I 1843c) with Latin and English descriptions and associated illustrations. Pfeiffer (1849: pl. 23, figs 1-3) and Reeve (1861: sp. 23a, b) published illustrations of the species, however, neither author recognized or used the three varietal forms. These subsequent illustrations are matched with the specimens in the Cuming collection labelled as "var. a". Therefore we believe this implies that "var. a" is the type series of Cyclostoma linguiferum s.s. and the specimens labelled as "var. b" and "var. c" are distinct variants and are therefore excluded from the type series of this nominal species (ICZN 1999: Art. 72.4.1). The specimen of "var. a" illustrated in Sowerby I (1843c: pl. 29, fig. 198), and closest to the dimensions given in the original description is here designated as the lectotype.
Remarks. The original description included eight varieties indicated as "var. a" to "var. h.", based on samples from various localities sampled by H. Cuming and cites an illustration in "Thesaurus Conchyliorum part 3, pl. 30, fig. 208". This illustration matches the specimens in Cuming collection labelled as "var. a". Therefore we believe this implies that "var. a" is the type series of Cyclostoma linguiferum s.s. and the specimens labelled as "var. b" to "var. h" are distinct variants and are therefore excluded from the type series of this nominal species (ICZN 1999: Art. 72.4.1). The specimen of "var. a" illustrated by Sowerby I (1843c: pl. 29, fig. 208) is here designated as the lectotype.
Measurements of specimens in the lots previously recognized as "var. b" to "var. h" are given here for future reference: Remarks. The original description did not include an illustration but Reeve (1861: sp. 2) subsequently re-published the description with illustrations of a specimen from the Cuming collection. The NHM collection contains a lot of three specimens' from the Cuming collection labelled "Malay Peninsula". A label reads "Mr. Benson has also sent me his Malayanum and the true volvulus for comparison….". Two of the three shells are close to the measurements and description in the original description and the label, presumably written by Cuming, indicates that these are the three specimens sent to Cuming by Benson, of which two without opercula are syntypes. The third specimen with an operculum being what Benson considered to be 'volvulus' and not a member of the type series of 'malayanus'. There is a further lot from "Pulo Penang" housed in the University Museum of Zoology Cambridge with original Benson labels including one specimen labelled by Benson as 'Type'. ( Remarks. The original description did not include an illustration and only one set of shell measurements was given. There are two specimens in the NHM collections with an original label stating "Type" and giving the reference of the original description. The shell that most closely matches with the measurement in the original description and with an "x" written in the aperture is here designated as the lectotype. Remarks. The original description included two sets of shell measurements (the type and the largest specimen), and Godwin-Austen explicitly stated there to be a unique name-bearing type. The NHM collections contain a lot of four shells from the Godwin-Austen collection (ex. Doherty collection) and have his original handwritten label stating "Type". The specimen with 'type' written on the shell most closely matches with the 'type' shell dimensions given in the original description and is here considered to be the holotype, the remaining three shells being paratypes.  Fig. 15B; D=43.9 mm, H=34.1 mm, W=5; D=44.7 mm, H=35.7 mm, W=5; D=42.3 mm, H=32.7 mm, W=5).

Cyclophorus niahensis
Remarks. The use of the term "holotype" in Vermeulen (1999: 144) does not constitute a valid lectotype designation, since there was no explicit indication to a particular specimen (ICZN 1999: Art. 74.5). The specimen which most closely matches the figure in the original description (especially in respect to the position of interrupted growth lines on last whorl) and marked with an "X" on the inside of the aperture is here designated as the lectotype. Remarks. The original description clearly stated the taxon was based on two specimens from Busan Hills, and the unique name-bearing type was not stated. However, there is only one remaining specimen from the Godwin-Austen type lot in the NHM collections. Subsequent use of the term "holotype" in Vermeulen (1999: 144) is seemed an unambiguously selected a particular specimen as the name-bearing type. This constitutes a valid lectotype designation (ICZN 1999: Art. 74.5).  Remarks. This species was described based on specimens from the Cuming collection. Pfeiffer (1854a: pl. 45, figs 13, 14.) re-published the description and figured this species. The NHM collections contain a lot of three shells from the Cuming collection with original labels in Pfeiffer's handwriting stating the species name. None of these shells exactly match with the illustration in Pfeiffer (1854a). However, the specimen mostly similar to the illustration in Reeve (1861: sp. 22) is illustrated here and is designated as the lectotype.  Fig. 16F; D=27.3 mm, H=20.2 mm, W=5).

Cyclophorus poeciloneurus
Remarks. The authors indicated that four lots of specimens were examined in the original description (from the Godwin-Austen, Ogel, Doherty and Beddome collections). In addition, the authors stated "Type" in relation to the specimens from the Godwin-Austen collection, which consist of two shells. The original description did not include an illustration, and only one set of shell measurement was given. The specimen that has a small label stating "Type" glued inside the aperture, and which matched the measurements given in original description is here designated as the lectotype. The paralectotypes from the Ogel, Doherty, and Beddome collections were not found. Kobelt, 1908 Type material. Lectotype (design. n.), NHMUK 20130119/1 (Fig. 18A; D=63.1 mm, H=46.5 mm, W=5½), paralectotypes NHMUK 20130119/2-3 (2 shells; Fig.  18B; D=57.7 mm, H=43.2 mm, W=5½; D=60.0 mm, H=46.8 mm, W=5½).

Cyclophorus fulguratus rangunensis
Remarks. This species was described based on a specimen collected by M. Mouhot from the Cuming collection, and only one set of shell measurements and one specimen was illustrated in the original description. There are three shells in the NHM collections from the Cuming collection with an original label in Pfeiffer's handwriting stating the taxon name, collector and collection locality. The figured specimen with an "x" written inside the aperture is here designated as the lectotype. The type locality of Cambodia applied to contemporaneous boundaries but Mouhot also collected in an area that now comes within the boundary of southern Vietnam.
Remarks. The authors indicated that four lots of specimens were examined in the original description. The original description did not include an illustration, and two set of shell measurement were given. However, the holotype was clearly designated and is housed in the Leiden Museum, Netherlands (now Naturalis Biodiversity Centre). The NHM registration records show that this specimen was purchased from Laidlaw, ex. Dr. Jacobson collection, and the original label states 'paratype'. The locality given by Laidlaw was "2 ex. July; Sinabang" match with the specimen. We therefore consider the single specimen as paratype.
Remarks. This taxon was described based on specimens from the Theobald collection. Coan and Kabat (2012: 326) stated that the types could not be located in either the NHM or the Leeds Museum. We located 3 specimens in the NHM general collection with a label stating that they were purchased from Theobald, "type figd in C. I.", but giving Moulmein (currently Mawlamyine), farm (=from?) caves, as the locality, which is at the mouth of the Salween. It seems likely that there is a mix up in the documentation of the locality but we cannot determine if the error is in the locality given in the original description, an error in the labelling with the specimens or if this lot is simply not type material. We therefore treat the material as possible syntypes.
The locality given by Hanley and Theobald was Upper Salwen (Salween or currently Thanlwin). However, the material identified as the syntype series carries labels giving Moulmein (currently Mawlamyine), farm (=from?) caves, as the locality, which is at the mouth of the Salween. As the Salween is close to 3,000 km in length and the river originates from the Tibetan Plateau, it is clear that 'Upper Salwen' is inaccurate. It is conceivable that the intended record was Upper Myanmar but on current evidence we conclude that the Mawlamyine is the type locality.

Cyclophorus siamensis (Sowerby I, 1850) http://species-id.net/wiki/Cyclophorus_siamensis
Cyclostoma siamense Sowerby I, 1850: 158, pl. 31a, figs 292, 293. Pfeiffer 1854a. Remarks. The original description as well as those in Pfeiffer (1854a: 323) are particularly accurate, both showing the dark banding pattern and varix on the last whorl, and both figures appear to be from the same specimen. The NHM collections contain two shells from the Cuming collection with original labels stated the taxon name, type locality and "f. 292, 293". The specimen which corresponds to the illustrations in Sowerby I (1850) and Pfeiffer (1854a) is here designated as the lectotype. Nevill (1878: 273) stated that Cyclostoma siamensis Sowerby I, 1850 occur in Khasi Hills, India not in Siam. Nevill considered this is an inappropriate taxon name, and Cyclophorus khasiensis Nevill, 1878 was nominated as a new replacement name based on specimens from the Godwin-Austen collection. This is however an unjustified replacement name, and therefore a junior objective synonym of Cyclostoma siamensis Sowerby I, 1850(ICZN 1999. The specimens of Cyclophorus khasiensis in the Godwin-Austen collection have no nomenclatural status. It should be noted that Kobelt (1902) attributed the date of publication of this species in error as "1843". For the correct dates of publication for "Thesaurus Conchyliorum" see Petit (2009: 32). ( Type material. Lectotypes (design. n.), NHMUK 20130083/1 (Fig. 21A; D=27.9 mm, H=19.8 mm, W=4), paralectotypes NHMUK 20130083/2-3 (2 shells; Fig. 21B; D=26.8 mm, H=21.4 mm, W=4; D=28.2 mm, H=20.9 mm, W=4).

Cyclophorus spironema
Remarks. This species was described based on specimens from the Cuming collection. The NHM collections contain three shells from the Cuming collection with an original label in Pfeiffer's handwriting giving the taxon name and collection locality. The specimen figured in Gude (1921: 55, 56, fig. 13) does not constitute a lectotype designation; as Gude did not select a particular syntype to be the unique name-bearing type (ICZN 1999: Art. 74.3). We here designate the specimen figured in Gude (1921: fig. 13) as the lectotype. Remarks. The term "nonnunquam" ("sometimes") in the original description of shell shape appears to imply that this taxon was based on more than one specimen although only one set of measurements was given in the original description. The use of "the type" in Hanley and Theobald (1870: 16) may not to constitute a valid lectotype designation because a label stating that it is the figured specimen is not in Hanley or Theobald's hand and it is not clear if only one specimen was available to Hanley and Theobald. We therefore treat "the type" attribution as an invalid, inadvertent lectotype designation (ICZN 1999: Art. 74.5). The single specimen in the NHM from the Blanford collection and figured in Hanley and Theobald (1870: pl. 34, fig. 7), is here designated as the lectotype. Remarks. The original description of this species included seven varieties indicated with "var. a" to "var. g" from the Cuming Collection. Latin and English descriptions associated with illustrations were then published in the "Thesaurus Conchyliorum" (Sowerby I 1843c). None of the subsequent authors recognized or used these seven varietal names. Two further works provided illustrations of the species from the Cuming coll. (Pfeiffer 1848: pl. 8, figs 13-16;pl. 16, figs 17-20;Reeve 1861: sp. 25a, b;8 fig. 30). These subsequently published illustrations match the specimens in the Cuming collection labelled as "var. a". Therefore we believe this implies that "var. a" is the type series of Cyclostoma tigrinum s.s. and the specimens labelled as "var. b" are distinct variants and are therefore excluded from the type series of this nominal species (ICZN 1999: Art. 72.4.1). The specimen of "var. a" illustrated by Sowerby I (1843c: pl. 29, figs 201, 202) is here designated as the lectotype.

Cyclophorus subblaevigatus
Measurements of specimens in "var. b" are given for future reference: Remarks. The original description of this species included two un-named varieties from the Cuming Collection. The NHM collection contain two lots from the Cuming collection with the original labels giving the taxon name, type locality and varietal names stated as "var. a" and "var. b". Latin and English descriptions associated with illustrations were then published in the "Thesaurus Conchyliorum" (Sowerby I 1843c). Two further works provided illustrations of the species from Cuming coll. (Pfeiffer 1849: 169, pl. 23, figs 10, 11;Reeve 1861: sp. 9). These subsequently published illustrations match specimens in the Cuming collection labelled as "var. a". We consider that this implies that "var. a" is the type series of Cyclostoma tuba s.s. and the specimens labelled as "var. b" are distinct variants and are therefore excluded from the type series (ICZN 1999: Art. 72.4.1). The specimen of "var. a" illustrated by Sowerby I (1843c: 122, pl. 27, figs 129, 130) is here designated as the lectotype.
Remarks. The name Cyclostoma turgidum Pfeiffer, 1851was presented as a replacement name for Cyclostoma crassum Pfeiffer, 1853, a junior homonym. However since the "crassum" description was not published until 1853 (in the Proceedings of the Zoological Society for 1851 volume, see Duncan 1937) Cyclostoma turgidum is the valid original description. This taxon was described and illustrated based on specimens from the Cuming collection. The NHM collections contain two lots from the Cuming collection that have original labels in Pfeiffer's handwriting with a striking-through of the taxon name "crassum", replaced with "turgidum". One lot of 3 specimens, NHMUK 20040591, has the collection locality "Liew Kiew" which matches with that given in the original description. The specimen figured in Pfeiffer (1853a: pl. 35, figs 15, 16) which matches with the measurements given in the original description is here designated as the lectotype. The second lot of three shells, NHMUK 20040590, has the collection locality "Ibyat, Bashee Islands" which corresponds to "var. minor in insula Ibyat (Bashee group)" from the Pfeiffer's (1853b) "crassum" description and is excluded from the type series of this nominal species (ICZN 1999: Art. 72.4.1).
Remarks. The original description of this species included four varieties indicated "var. a" to "var. d", without illustration from the Cuming Collection. Subsequent authors (including Sowerby I 1843b) did not recognise or use these varietal names. The NHM collections contain four lots of Cuming collection material with original labels giving the taxon and varietal names "var. a" to "var. d". The description and illustration in Sowerby I (1843b) match the specimens in the Cuming collection labelled as "var. a". We consider that this implies that "var. a" is the type series of Cyclostoma validum s.s. and the specimens labelled as "var. b" , "var. c" and "var. d" are distinct variants and are therefore excluded from the type series (ICZN 1999: Art. 72.4.1). The specimen of "var. a" illustrated by Sowerby I (1843c: pl. 27, figs 132, 133), is here designated as the lectotype.
Measurements of specimens in "var. b" to "var. d." are given for future reference:  Note that the strikethrough on the "type" and "Holotype red spot" with blue pen are possibly added later by the NHM assistant C The original label of Cyclophorus cucullatus marked with "Type" is not frequently occurred in Cuming collection, which the possibly indicates specimen received from A.A. Gould D The small glued-label written with blue ink on "excellens Pfr" and "allied to pirrieanum" are Pfeiffer's handwritten.