An annotated type catalogue of seven genera of operculate land snails (Caenogastropoda, Cyclophoridae) in the Natural History Museum, London

Abstract The collection of the seven cyclophorid snail genera housed in the Natural History Museum, London (NHM), includes 95 available species-level names belonging to the genera Pterocyclos Benson, 1832, Cyclotus Swainson, 1840, Myxostoma Troschel, 1847, Rhiostoma Benson, 1860, Scabrina Blanford, 1863, Crossopoma Martens, 1891, and Pearsonia Kobelt, 1902. Lectotypes are here designated for twelve available species-level names to stabilise existing the nomenclature. A complete catalogue of these types, including colour photographs, is provided for the first time. After examining these type specimens, an unpublished manuscript name was found and is described herein as Pterocyclosanamullayensis Sutcharit & Panha, sp. n.


Introduction
Cyclophoridae Gray, 1847 is a major group of terrestrial operculate snails found across southern Europe, Central America, Asia, Africa, and Australia (Kobelt 1902, Solem 1959, Stanisic 1998, Nantarat et al. 2014a. Many cyclophorid groups are common (1870)(1871)(1872)(1873)(1874)(1875)(1876). The type locality is as stated in the original publication in the original wording and language. Additional locality data from original labels, with respect to current political boundaries or subsequently published localities is given in square brackets. Under the type materials, primary type specimens with the Natural History Museum registration number (hereafter NHMUK), number of specimen(s), and the figures as listed in this publication are given. In addition, if the paratype(s) of that taxa are present; the registration number, number of specimens, and figures of representative specimen are also given. The history and type evidence is summarised under each taxon. Full bibliographic references are provided at the end of this paper.
Type material. Holotype NHMUK 20160307 (Fig. 1A), paratype ANSP 467381 (1 shell). Remarks. Godwin-Austen's description was based on three lots of specimens. The original description includes an illustration and one set of measurements. The original description stated "Type no. 3104 Brit. Mus." There are three specimens from Godwin-Austen type collection that relate to this registration number. The specimen figured in the original description which corresponds to the measurements given is figured herein (Fig. 1B).
Remarks. The original description did not include an illustration and only one set of shell measurements was given. The author stated "sometime zigzag", which implied that this taxon was based on more than one specimen. The NHM collections contain a lot com-prised of a single specimen purchased from Sowerby and Fulton with a label stating "Type" and with a collection locality written on the original label, this specimen is figured herein. (Pfeiffer, (Fig. 1E), paralectotypes NHMUK 20170352/2-3 (2 shells; Fig. 1F).

amboinensis
Remarks. The original description lacked mention of an operculum and did not include an illustration, and only one set of shell measurements was given. Pfeiffer (1854a) re-published the description and figured the species which no operculum. There is a lot of four shells from the Cuming collection with two Pfeiffer handwritten labels. The larger label states "C. Amboinensis Pfr.", "The operculum shows that…" and the collection locality from "Isle of Lobok". We presume that this lot contains a subsequently introduced specimen that has an operculum. Therefore, the specimen with an operculum attached (NHMUK 20070352/4) is excluded from the type series. The remaining three shells have no operculum and the smaller label states "Cycl. Amboinense Pfr." and gives the collection locality as "Amboyana". The specimen that most closely matches with the measurements given in the original description and the illustration in Pfeiffer (1854a) is here designated as the lectotype to stabilise the name. Fig. 1G, H Spiraculum andersoni Blanford, 1869b: 447. Pterocyclos andersoni -Hanley andTheobald 1872: 23, pl. 49, figs 3, 4. Pearsonia andersoni -Kobelt 1902: 172. Kobelt 1911. Current generic position. Pearsonia Kobelt, 1902 Type locality. ad Bhamo cum praecedente [Bhamo District, Kachin State, Myanmar]. Type material. Syntype NHMUK 1906.5.5.77 (3 shells; Fig. 1G, H).
Current generic position. Pterocyclos Benson, 1832 Type locality. Les environs de Touranne (Cochinchine) [Da Nang City, Vietnam]. Type material. Syntype NHMUK 1854.7.24.365 (2 shells; Fig. 1I, J). Remarks. The original description included one set of measurements. Later, Eydoux and Souleyet (1852) re-published the description and figured this species. Gray (1855: 19) made a list of type specimens of molluscs described by the late M Souleyet and housed in the NHM collections. There are two specimens with an original label stating the species name and collection locality. The specimen that closely matches the figure in Eydoux and Souleyet (1852) and is closest to the measurements given in the original description is illustrated in Figure 1I. Bullen, 1906Fig. 1K, L Pterocyclos aspersus Bullen, 1906: 129, with text figure. Kobelt 1911 Fig. 1K, L). Remarks. The original description is clearly based on more than one specimen, but only one set of measurements and illustrations were given. The NHM type collections contain a lot of six shells with the label stating the taxon name, collection locality, and "Type". The specimen figured in the original description, which is closest to the shell dimensions given in the original description, is illustrated in Figure 1K. (Fulton, 1900) Fig. 2A, B
Remarks. The species was clearly based on two specimens, one dead and one alive. The original description did not include an illustration and only one set of measurements was given. Subsequently, Hanley and Theobald (1875) figured a specimen of this species. The single specimen in the NHM from the Godwin-Austen collection, ex. WT Blanford collection and figured in Hanley and Theobald (1875) is figured herein.
Remarks. The original description by Pfeiffer includes an illustration and one set of shell measurements. The type lot in the NHM collections was collected by "Mr. Wallace" and is from the Cuming collection as stated in the original description. It has an original label in Pfeiffer's handwritings giving the species name and collection locality. The adult specimen that closely matches the measurements and the illustration shown in the original description is figured herein.
Remarks. Smith clearly stated that this taxon was described based on a single specimen from the GB Sowerby I collection. The species description included an illustration and a set of shell dimensions. The NHM collections contain a type lot that has an original label stating "Type", subsequently re-written as "Holotype". Therefore, we recognise this single shell as the holotype fixed by monotypy.

Fig. 2F
Spiraculum beddomei Blanford, 1866: 31, 32. Pterocyclos beddomei -Hanley and Theobald 1875: 54, pl. 134, figs 5, 6. Pearsonia beddomei -Kobelt 1902: 172, 173, fig. 36. Kobelt 1911 2F). Remarks. The species description was based on more than one specimen. The original description did not include an illustration, and only one set of measurements was given. Hanley and Theobald (1875) subsequently figured this species. There is a specimen in the NHM from the Blanford collection figured in Hanley and Theobald (1875) that closely matches the dimensions given in the original description, it is figured herein.
Remarks. The original description did not include an illustration, and only one set of measurements was given. Gude (1921: 116, 117, fig. 20) re-published the description and figured a type specimen from Theobald's collection. There is one shell in the NHM collection purchased from W Theobald, with an original label stating "type" and the collection locality "Bhamo". The specimen figured herein closely matches the measurements given in the original description.  Fig. 2H, I).

bifrons
Remarks. The species was described based on specimens from the Cuming collection. The original description did not include illustrations, and only one set of measurements was given. Reeve (1863) re-described and illustrated a single specimen from the Cuming collection. There are three specimens from the Cuming collection in the NHM type lot with an original label in Pfeiffer's handwriting stating the species name and collection locality. The specimen figured in Reeve (1863) closely matches the measurements given in the original description is illustrated here in Figure 2H. Remarks. This species was described based on material from the Cuming collection, and only one set of shell measurements was given. Later, Reeve (1863) re-described the species and illustrated a shell from the Cuming collection. There are three specimens in the NHM collections with an original label stating "Rev C. I. f. 18a-b".

birostris
The specimen figured herein (Fig. 2J) is closest to the illustration in Reeve (1863) and the shell dimensions given in the original description.
Remarks. The original description did not include an illustration, and only one set of measurements was given. Gude (1921: 117, 118, fig. 21) re-published the description and figured a shell from Theobald's collection. There is one shell in the NHM collection purchased from W Theobald, with an original label stating "type" and the collection locality "Bhamo". This specimen closely matches the measurements given in the original description and is figured herein. Remarks. This species was described from the Hungerford collection. An illustration and a set of shell dimensions were given in the original description. Godwin-Austen (1889) does not explicitly indicate the number of specimens he has available to him in the original description. There is a single shell in the NHM collections from the R Hungerford collection with an original label stating "Type", the species name, and collection locality. This specimen closely matches with the illustration and the shell measurements given in the original description, and it is figured herein.   Fig. 3C).

boxalli Godwin-Austen, 1893
Remarks. The original description gives a set of shell measurements and figures of two specimens, so this species is clearly based on more than one specimen. There is a type lot (two shells) with differing registration numbers, one specimen NHMUK 1894.5.23.1 is from R Hungerford ex. Mr. Boxall from Kina Balu and the NHM registration book states "Type". The other shell, NHMUK 1895.12.5.34, in the same box was collected by Mr. Whitehead from Palawan and agrees well with Godwin-Austen (1893: fig. 1b, c). These two shells form part of the type series and are considered as syntypes. Remarks. Godwin-Austen clearly stated that the original description was based on a lot of three shells "Type No. 713 B.M.". The NHM type collections contain a lot of three shells from the Godwin-Austen collection and original label states "TYPE". The specimen with red wool inside the aperture that corresponds to the illustrations and measurements given in the original description, and it is figured herein (Fig. 3D).  3F). Remarks. Sykes clearly stated that this taxon was described based on a single specimen collected by Mrs. Broun. The original description included an illustration and a set of shell measurements. The NHM collections contain a Sykes type lot that has an original label stating "Type", and so we recognise this single illustrated specimen as the holotype fixed by monotypy.  Fig. 3G, H).

calyx
Remarks. The original description did not include an illustration or state the number of specimens examined and there is no information concerning the operculum, however one set of shell measurements were given. Later, Pfeiffer (1860b) and Reeve (1861) re-published the description and figured the species from specimens in the Benson collection. There is a type lot in the NHM ex. Hawkins collection containing three shells (1 with and 2 without periostracum). The specimen without periostracum that most closely matches with the measurements given in the original description and the illustrations in Pfeiffer (1860b: pl. 37, figs 25-27) and Reeve (1861: pl. 20 Fig. 3I).
Remarks. The original description includes an illustration and one set of shell measurements is given. However, the species description was not clearly based on a single specimen. Only one shell was found in the NHM collections with an original label stating "Type" and giving the reference of the original description and collection locality. This figured specimen exactly matches with the illustration and shell measurements given in the original description.  Fig. 3J, K).

Remarks.
There are two specimens in the type lot with original labels in Smith's handwriting stating "Type". The original description gives one set of measurements and illustrates one specimen. The specimen that corresponds to the illustrations in Smith (1896b: figs 4-6) and the given shell measurements in the original description and has a red spot on the last whorl is figured herein (Fig. 3J). Remarks. The original description did not include an illustration, and one set of shell measurements was given. Later, Reeve (1862) re-published the description and figured a shell from the Cuming collection. There is one lot from the Cuming collection containing three specimens collected by Mr. Fortune with an original label in Pfeiffer's handwriting stating the species name and collection locality. The specimen that most closely matches the measurements given in the original description and the illustration in Reeve (1862) is figured herein (Fig. 4A).   Remarks. The original description was clearly based on more than one specimen, and included an illustration and one set of shell measurements. There is only one specimen in the NHM type collections with Tomlin's hand written label stating "Type" and the collection locality. This specimen exactly matches with the illustration (not full adult stage) and the measurements given in the original description and is here designated as the lectotype to stabilise the name. The other two lots of Tomlin collections in the NMW with the label stating "paratype" are therefore considered as paralectotypes. Remarks. This species was described from specimens in the Cuming collection and only one set of shell measurements was given in the original description. The NHM collections contains a lot of a single shell from the Cuming collection with an original label, probably in Pfeiffer's handwriting, stating the species name, however this has subsequently been overwritten. Reeve (1863) illustrated a single specimen from the Cuming collection. This single specimen closely matches the illustration in Reeve (1863) but is slightly larger than the shell dimensions given in the original description. Therefore, we consider this specimen to be a probable syntype.
Remarks. The species description by Pfeiffer did not indicate the number of available specimens and did not include an illustration. Only one set of shell measurements was given in the original description. Reeve (1861) re-published the description and figured one specimen from the Cuming collection. The NHM collections contain a lot containing one shell with Pfeiffer's handwritten label giving the species name and collection locality. This figured shell matches well with the measurements given in the original description and the figure in Reeve (1861).  Fig. 4G).

cucullus
Remarks. The original description included an illustration and one set of shell measurements. Only one specimen is housed in the NHM type collections, with an original label stating "Type". This shell closely matches the illustration and the measurements given in the original description. Remarks. This species was described from specimens collected by Mr. WM. Daly from Thailand. The original description included an illustration and a set of shell dimensions. The NHM collections contain a lot of three specimens collected by Mr. Daly and have an original label stating the species name and collection locality. The specimen which is identical to the illustration, particularly in respect to the breathing device, and the shell measurements given in the original description is figured herein (Fig. 4I). Remarks. This species was described from material in the Cuming collection, and the original description included only one set of shell dimensions. Reeve (1863) redescribed the species and illustrated a shell from the Cuming collection. The NHM collections contain a lot of two specimens from the Cuming collection with Pfeiffer's hand written label giving the species name and collection locality. The specimen that corresponds to the shell measurements in the original description and the illustration in Reeve (1863) is figured herein (Fig. 4K). Fig. 5A, B

dautzenbergi (Sykes, 1902)
Opisthophorus dautzenbergi Sykes, 1902a: 23. Sykes 1902b Fig. 5A, B). Remarks. The species description does not include an illustration, but later, Sykes (1902b) cited the species name and illustrated a shell of this taxon. The NHM collections contain a lot of three specimens from the Sykes collection. There is a specimen that exactly matches with the illustration in regard to the brownish zigzag streaks on the last whorl, and with the shell measurements given in the original description plus it has "Type" written on the shell. It is figured herein (Fig. 5A).

Current generic position. Cyclotus Swainson, 1840
Type locality. Demerara. Type material. Syntype NHMUK 20170358 (2 shells; Fig. 5D, E). Remarks. The original description and illustration in Sowerby I (1843a) as well as those in Pfeiffer (1849) and Reeve (1863) are particularly accurate, both showing the varices on the last whorl, which suggests that these figures are from the same specimen. The NHM collections contain a lot of two shells from the Cuming collection with original labels stating the taxon name and collection locality. The specimen that corresponds to the illustrations in Sowerby I (1843a), Pfeiffer (1849) and Reeve (1863) is figured herein (Fig. 5D).  Fig. 5F, G).

enganoense
Remarks. The original description included an illustration and a set of shell measurements. The species description was based on more than one specimen. There are two shells in the NHM type collections with an original label stating "Type lot", taxon name, and collection locality. The specimen that matches well with the illustration and the shell dimensions given in the original description is figured herein (Fig. 5F).  Fig. 5H, I).

euryomphalus
Remarks. This species was described based on specimens from the Cuming collection, and was not illustrated in the original description. Reeve (1863) re-published the description and figured a specimen from the Cuming collection. The NHM collections contain a lot of three shells from the Cuming collection with a label in Pfeiffer's hand written label stating the taxon name and collection locality. The specimen which closely matches the illustration in Reeve (1863) and the shell measurements given in the original description is figured herein (Fig. 5H).  Fig. 6A).

fairbanki
Remarks. The original description did not clearly state how many specimens were available to the author, although only one set of measurements was given. The NHM collections contain a lot comprising one specimen and an original label stating that the specimens were figured in the Conchologia Indica. This specimen matches well with the figures in Hanley and Theobald (1875: pl. 5, fig. 9, pl. 134, fig. 1), and the measurements given in the original description. 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) re-published the description and figured a specimen from the Cuming collection. The NHM collections contain a lot of two specimens with a label stating "Type". The adult specimen that matches well with the illustrations in Pfeiffer (1854a) and Reeve (1863), and the shell dimensions given in the original description, is figured herein (Fig. 5K). Type material. Holotype NHMUK 1889.12.7.9 (Fig. 6C).

frednaggsi Sutcharit & Panha, 2014
Remarks. Godwin-Austen clearly stated this taxon was described based on only a single specimen collected by A Everett. The original description included an illustration and a set of measurements. The NHM collections contain a Godwin-Austen type lot with an original label stating "Type", and so we recognise this illustrated shell as the holotype fixed by monotypy. Fig. 6D Fig. 6D, E). Remarks. Adams's description was based on specimens collected by Mr. Swinhoe from Hainan. The original description gives both a set of shell measurements and an illustration of a specimen. The NHM collections contain a lot of three specimens from the H Adams collection with an original label stating the taxon name and collection locality. The specimen marked with an "x" on the shell matches well with the shell dimensions and illustration given in the original description and it is figured herein (Fig. 6D).  Fig. 6F, G).

hainesi
Remarks. This species was described based on material in the Cuming collection, and Pfeiffer provided an illustration and a set of measurements. The NHM type lot contains two shells collected by H Mouhot and is from the Cuming collection. It has an original label in Pfeiffer's handwriting giving the taxon name, the collector as "Mr. Mouhot" and collection locality. The specimen that most closely matches the illustration and shell measurements given in Pfeiffers (1862) description is figured herein (Fig. 6F).  (Fig. 6H).

harryleei
Remarks. The shell characters of an expanded apertural lip and a small accessory breathing device (sutural tube) located in the suture closest to the aperture suggest that it is a member of the genus Cyclotus. It is very closely resembles the widespread species Cyclotus setosus (Möllendorff, 1894) in the southern Thailand. Fig. 6I Fig. 6I, J).

hispidula (Blanford, 1863)
Remarks. The original description did not include an illustration, and only one set of shell measurements was given. Subsequently, Hanley and Theobald (1872) figured a specimen of this species. There is a specimen in the NHM collections from the Blanford collection which was figured in Hanley and Theobald (1872) and matches well with the dimensions given in the original description, and it is figured herein (Fig. 6I).

hungerfordi (Godwin-Austen
Remarks. Godwin-Austen clearly stated that this taxon was described based on only one specimen from the R Hungerford collection. There is one specimen in the NHM collections that has an original label stating "Type". We recognise this specimen as the holotype fixed by monotypy.  ; Fig. 7A).

inglisianus
Remarks. The species description included an illustration and one set of shell measurements. There is a shell in the NHM collections from the Blanford collection, with an original label stating "figd. Conch. Ind. 143, f. 8, 9" and the collection locality "Damatha, Molmein". This specimen matches with the shell dimensions given in the original description but lacks an operculum. Therefore, we consider this lot to be a possible syntype.  (Fig. 7B).

insignis
Remarks. The species description was very brief and only one set of measurements was given. The NHM collections contain a mixed-species lot consisting of two specimens with an original label stating "Type". The specimen that matches with the figures in Hanley and Theobald (1870: pl. 5, figs 6, 7), the measurements (width 29.6 mm) given in the original description and marked with an "x" is here designated as the lectotype to stabilise the name.
The other shell, NHMUK 1888.12.4.1977, differs from the lectotype in having a smaller shell width (23.6 mm), shallow suture without a canal-like structure and an expanded apertural lip without a canal-like accessory respiratory structure. We considered this shell as a distinct species, and so it excludes it from the type series.   ; Fig. 7C).
Remarks. The original description gave the measurements for only one shell and did not include an illustration. The NHM collections contain a lot of one specimen from the R Hungerford collection with an original label stating "Type" and giving the collection locality as "Borneo". This specimen matches well with the original description and shell dimensions given. Fig. 7D Fig. 7E).

kempi (Godwin-Austen, 1915)
Remarks. Godwin-Austen's description clearly indicated that the type series was from two collection localities "Abor Hills" and "Pongping", and he provided illustrations of two shells from each different specimen lot. These two specimen lots were listed as being housed in the NHM collections. These two type lots have an original label in Godwin-Austen's handwriting stating species name, collection locality and giving his own catalogue numbers. The figured specimen (Godwin-Austen 1915: figs 4, 4a) labelled as "Type", is figured herein (Fig. 7D). Another figured shell (Godwin-Austen 1915: figs 5, 5a) labelled as "Typic." is also figured herein (Fig. 7E). In addition, the original description states 'Two specimens to Indian Museum'.  dimensions was given. Pfeiffer (1869) re-published the description and illustrated a specimen. The NHM collections contain a lot of three shells from the Cuming collection with a label stated "Type". The specimen that closely matches with the shell measurements, the illustration in Pfeiffer (1869) and has an "X" written on the shell, is figured herein (Fig. 7F). Remarks. Smith based this taxon on more than one specimen since the author states "most of the specimens". The original description includes an illustration but no shell measurements are given. The type lot in the NHM collections contain three specimens with an original label in Smith's handwriting. The specimen that most closely matches the illustrations in the original description is figured herein (Fig. 7H).   Fig. 7K, L). Remarks. The species description did not include an illustration, and only one set of shell measurements were given. Reeve (1863) re-published the description and illustrated an adult specimen from the Cuming collection. The NHM collections contain a lot of three shells collected by FW Lindstedt from the Cuming collection with a label in Pfeiffer's handwritten stating the taxon name and collection locality. The adult specimen matches well with the shell dimensions given in the original description, the illustration in Reeve (1863), and is figured herein (Fig. 7K).
Remarks. The species description was not explicitly based on one specimen, but only a single shell was illustrated and just one set of shell measurements was given in the original description. The NHM type lot contains one specimen with an original label in Smith's handwriting and a label stating "Type", subsequently changed to "Holotype". This is not a valid holotype designation (ICZN 1999: Art. 73.1 and Recommendation 73F). This specimen matches well with the illustration and the shell measurements given in the original description, and is designated here as the lectotype to stabilise the name. Fig. 8B Fig. 8B, C).

lowianus (Pfeiffer, 1864)
Remarks. This species was described based on specimens collected by H Low from the Cuming collection. In the original description, only one set of shell measurements was given. In 1869, Pfeiffer re-described and illustrated a single specimen from the Cuming collection. The NHM collections contain a lot of three shells from the Cuming collection with Pfeiffer's handwritten label stating the taxon name, collector and collection locality. The specimen that matches well with the shell dimensions in the original description and the illustrations in Pfeiffer (1869) is figured herein (Fig. 8B). Fig. 8D Spiraculum luyorensis Godwin-Austen, 1915: 500, pl. 40, figs 5, 5a, b. Pearsonia luyorensis -Gude 1921: 119, 120. Kobelt, 1902 Type locality.  Fig. 8D).

Remarks.
Godwin-Austen's description was not clearly based on only one specimen. The original description included an illustration and one set of measurements were given. The author stated that one lot of the type series was housed in the NHM collections. The specimen NHMUK 1903.7.1.3530 has a label in Godwin-Austen's handwriting stating "Type" and exactly matches the measurements and illustrations given in the original description. Fig. 8E Fig. 8F).

lychnus (Morelet, 1862)
Remarks. The original description by Morelet (1862: 478) did not give an illustration of the species, although a set of shell dimensions was provided. The NHM register show that a lot of three specimens was purchased from Fulton from the A Morelet collection. The original label states the taxon name and gives the collection locality as "C. breve Martyn-lychnus Moret. I. Pulo Condor". The words "All too small for types" was added at a later date, however, our measurements of all specimens are close to those shell dimensions given in the original description (width 36 mm, height 25 mm). The specimen that matches well with the shell dimensions given in the original description is here designated as the lectotype to stablise the name.
Remarks. The original description does not clearly state how many specimens were available to the author. Laidlaw's use of the term "Type specimen in my collection" does not constitute a valid holotype designation (ICZN 1999: Art. 73.1.1, 73.2 and Recommendation 73F). The NHM collections hold a lot containing one specimen, NHMUK 1939.4.13.23, with a label in Laidlaw's handwriting stating the taxon name, collection locality and "From type series". It has subsequent been incorrectly labeled as a "Paratype". To avoid the assumption of the existence of a holotype (ICZN 1999: Recommendation 73F), this specimen is here designated as the lectotype to stabilise the name.
Remarks. This species was clearly described based on more than one specimen since the author stated "largest example". The NHM collections contain a lot containing a single specimen with an original label in Godwin-Austen's handwriting stating "Type". This specimen matches well with the illustration and the measurements given in the original description. Current generic position. Pterocyclos Benson, 1832 Type locality. Naga Hills [Assam, Arunachal Pradesh and Nagaland states, India].
Remarks. A unique name bearing type was not explicitly designated and the species description was not clearly based on one specimen. The original description included a single illustration and one set of shell measurements. The RBINS collections contain a lot with one specimen from the collection of P Dautzenberg with an original label stating "Type" and " fig.". This specimen matches well with the shell measurements given in the original description and is figured herein (Fig. 8I). The NHM collections also contain another lot of two specimens with an original label stated "cotypes", and are also considered to be syntypes.  Blanford, 1877: 313, 314. Pearsonia mastersi -Kobelt 1902: 174. Kobelt 1913. Kobelt, 1902 Type locality. in montibus Naga dictis, ad latus meridionale province Assam, haud procul a Golaghat [Naga Hills, Golaghat District, southern of Assam State, India].
Remarks. This taxon was first published in Hanley and Theobald (1870), who attributed the name to WT Blanford as a manuscript name. However, Hanley and Theobald (1870: 3) did not provide any description or definition of the taxon, only a figure was provided, which does not meet the requirements of the ICZN (1999: Art. 12.2). Therefore, this taxon name was not made available. Pfeiffer (1876: 386) mentioned the taxon name without further description or definition, and so again the was not made available. Blanford (1877) published a complete description of this taxon under the same name and so making it available. Therefore, the type series is made up of the specimens sent by Blanford and cited and figured in Hanley and Theobald (1870: pl. 5, fig. 1), along with the specimens mentioned by Blanford (1870). Coan and Kabat (2012: 326) could not trace the specimen figured in Hanley and Theobald (1870: pl. 5, fig. 1). However, the NHM collections hold a lot containing a single shell from the WT Blanford collection with a handwritten label stating the taxon name, and the collection locality "Gholaghat, Assam". This specimen matches well with the original description and the set of shell dimensions, and is considered as the syntype and figured herein.  Fig. 8K, L Pterocyclos mindaiensis Bock, 1881: 634, pl. 55, figs 8, 8a, b. Cyclotus mindaiensis -Kobelt 1902 Current generic position. Cyclotus Swainson, 1840 Type locality. Mindai (Amontai district) [Amuntai, Hulu Sungai Utara, South Kalimantan Province, Indonesia].
Remarks. In the original description, Bock illustrates two specimens (an adult and a juvenile), although only one set of shell measurements were given. There are four specimens in the NHM type lot from the Bock collection with an original label stating "Types". The adult specimen, marked with an "x" matches well with the shell dimensions and illustration given in the original description (Bock 1881: fig. 8) and is figured herein (Fig. 8K).

minima (Godwin-Austen, 1915) Fig. 9B, C
Spiraculum minimum Godwin-Austen, 1915: 501, 502, pl. 40, figs 2, 2a-c. Pearsonia minima -Gude 1921. Kobelt, 1902 Type locality. Remarks. Godwin-Austen (1915) clearly states that this taxon was based on at least four lots of specimens from two different localities. The original description included illustrations and one set of measurements. In addition, Godwin-Austen (1915) stated that two lots were housed in the Indian Museum, and the other two lots were housed in the NHM. The specimen lot NHMUK 1903.7.1.3145 consisting of three shells has a label in Godwin-Austen's handwriting stating "Typical", and gives the collection locality "Sibbum", and one of the specimens closely matches the original description. This specimen is here designated as the lectotype to stabilise the name.

mucronatus (Sowerby I, 1843)
Remarks. The species name was made available in volume 1 of the Thesaurus Conchyliorum, (Sowerby I 1843a; for date of publication see Petit (2009)). The original description includes an illustration, and the author clearly stated the taxon was described based on specimens collected by H Cuming. The NHM collections contain a lot of nine shells that have a label stating the type collection locality 'Calauang, Luzon'. The NHM registration records show that this lot was purchased from the H Cuming collection. The specimen that closely matches with the original description and illustration in Sowerby I (1843a) is figured herein (Fig. 9D). Pearsonia nagaensis -Kobelt 1902: 174. Kobelt 1913. Kobelt, 1902 Type locality. Maokokchung, Naga Hills [Mokokchung District, Nagaland State, India].
Remarks. The original description did not contain any illustrations, and only one set of measurements was given. Godwin-Austen stated that the type series was from the Beddome collection. The NHM collections contain two lots that are considered to constitute the type series. Lot NHMUK 1903.7.1.2783 consists of a single specimen, collected by Muspratt, from the Godwin-Austen collection and has original labels giving the species name "nagaensis", type collection locality and stating "TYPE". This specimen is here designated as the lectotype to stabilise the name.

natunensis Smith, 1894
Remarks. The original description included an illustration and one set of shell measurements. The NHM collections contain a lot of four shells collected by A Everett from Smith's collection with an original label stating taxon name, type collection locality, and "Type". The specimen that matches well with the illustration and shell dimensions given in the original description is figured herein (Fig. 9H).
Remarks. The species description clearly stated that the taxon was based on two specimens from Dihiri Parbat. An illustration and one set of measurements were given in the original description. The NHM type collection contains one lot, consisting of two shells with Godwin-Austen's handwritten label stating "Type". The specimen figured in the original description that is closest to the given measurements is here designated as the lectotype to stabilise the name. Godwin-Austen (1915: 497, 498) also recognised two varieties as "var." and "var. large". Nevertheless, these two varietal names have never been made available. There are two lots of specimens labeled as "var." and "var. large" housed in the NHM general collections, NHMUK 1903.7.1.3531 (3 shells) and NHMUK 1903.7.1.3089 (4 shells) respectively. They are excluded from the type series of this nominal species (ICZN 1999: Art. 72.4.1). Fig. 10B ; Fig. 10C).

niahensis Godwin-Austen, 1889
Remarks. Godwin-Austen described this species based on material from A Everett. The NHM collections contain a lot of two shells from the A Everett collection with an original label stating "Type". The specimen which corresponds to the illustrations and shell dimensions given in the original description, and marked with an "X", is figured herein (Fig. 10B).
Godwin-Austen also described a varietal form "var. depressa" from "Molu Hills" based on specimens collected by Mr. Boxall, ex. R Hungerford collection. The type series of the taxa could not be traced in the NHM collections.

Current generic position. Cyclotus Swainson, 1840
Type locality. Nias Island, Sumatra [Nias Island, north Sumatra Province, Indonesia]. Type material. Syntype NHMUK 1907.5.3.131-133 (3 shells; Figs 9L, 10A). Remarks. The original description included an illustration and gave one set of shell measurements. There are three specimens in the type lot with an original label in Fulton's handwriting stating "Type". The specimen that has a small label stating "Type." glued on top of the shell matches well with the measurements and figures given in the original description, is figured herein (Fig. 9L). Fig. 10D Fig.  10D, E).

oakesi (Godwin-Austen, 1915)
Remarks. The author indicated that five specimens were examined, and the original description included an illustration and one set of measurements. The NHM type collections contain a lot of five specimens with a label in Godwin-Austen's handwritingstating "Type". The specimen with red wool inside the aperture, illustrated in the original description and very close to the given measurements, is figured herein (Fig. 10D). The other two specimens housed in the Zoological Survey of India (no. 3081) are also considered as syntypes.  Sowerby I, 1843a: 97, pl. 23, fig. 17. Pfeiffer 1849: 155, pl. 21, figs 13, 14. Reeve 1862: volume 13, Cyclostoma, pl. 19, species 125. Cyclotus politus -Kobelt 1902 Current generic position. Cyclotus Swainson, 1840 Type locality. Unknown. Type material. Possible syntype NHMUK 20170360 (4 shells; Fig. 10F, G). Remarks. The original description was published by Sowerby I in 1843 andincluded andillustation. Later, Pfeiffer (1849) and Reeve (1862) re-published the description and figured this species based on material from the Cuming collection. The NHM collections contain a lot of four shells from the Cuming collection with original labels giving only the taxon name. However, the collection locality "Flores (Martens in litt.)" has been subsequently added. The specimens in the Cuming collection matches well with the illustration in the original description, especially in regard to the dark brown reticulated pattern and dark spiral band on the periphery of last whorl. However, Sowerby I (1843a) did not explicitly state that the species description was based on specimens from the Cuming collection. Therefore, we consider this lot to be possible syntypes.
Remarks. Reeve (1863) erroneously replaced the previously available name "pterocycloides Pfeiffer, 1855" with "anomalus Reeve, 1863". This unnecessary substitution is therefore made available with it own authorship and date and is considered to be a junior objective synonym (ICZN 1999: Arts. 19.1, 33.2, 50.5 and 60.1).
Pfeiffer (1855a) stated that this species was described based on specimens from the Cuming collection. The original description did not include an illustration or collection locality, and only one set of shell measurements were given. The type lotcontains four specimens of the same species, however an original label records that this is a mixed lot containing three specimens from the Cuming collection and 1 specimen with an operculum presented by GB Sowerby I. Specimen NHMUK 20170361/1 has no operculum, is without collection locality but has an original label in Pfeiffer's handwriting stating "Cyclost. pterocycloides Pfr.". This specimen, that matches with the description and shell measurements given in the original description, and the illustration in Reeve (1863), is here designated as the lectotype to stabilise the name. The other two shells, NHMUK 20170361/2-3, from the Cuming collection lot therefore become the paralectotypes.
The remaining a single shell with an operculum, NHMUK 1886.9.10.1, which was presented by GB Sowerby I and has the collection locality "Borneo"is excluded from the type series.  Fig. 11A) from Puri, Orissa.

puriensis
Remarks. The author clearly indicated that this taxon was described based on material from Pooree (=Puri) and Chandbally. The original description does not include any illustrations or shell measurements, and the species description is very brief, simply indicating that this taxon is a "dwarf form with raised spire…". The NHM collections contain a lot containing one shell from the Beddome collection with an original label stating collection locality as "Puri, Orissa". Since this lot was registered as "Nev. Hnd. List. p. 260", we consider this shell to be a possible syntype. Fig. 10J, K Spiraculum putaoensis Godwin-Austen, 1915: 500, 501, pl. 40, figs 3, 3a, b. Pearsonia putaoensis -Gude 1921 Current generic position. Pearsonia Kobelt, 1902 Type locality.  Fig. 10J, K). Remarks. The original description included an illustration, one set of measurements, and Godwin-Austen (1915) indicated that he examined three specimens. The NHM type collections contain a lot of three specimens with a label in Godwin-Austen's handwriting stating "Type". The specimen closest to the given measurements and illustrations in the original description is figured herein (Fig. 10J).  Fig. 11B), NHMUK 1896.5.1.1-2 (2 shells; Fig. 11C).

pyrostoma
Remarks. There are three specimens in the NHM type lot with a label in Smith's handwriting stating the species name and collection locality. One specimen has a small label "Type" attached under the shell. This specimen matches well to the illustration and shell dimensions given in the original description and is figured herein (Fig. 11B). Fig. 11D Fig. 11D, E). Remarks. Martens (1860) states that this species was described based on material in the British Museum [= NHM] collected by JC Bowring from Siam. The original description did not include an illustration and only one set of shell measurements was given. The NHM collections contain a lot of eight specimens with an original label stating "typical specimens", and with the collection locality "Siam". The NHM registration book shows that this specimen lot was purchased from JC Bowring. The specimen that matches most closely with the shell measurements in the original description is figured herein (Fig. 11D).  Fig. 11F, G).

simplicilabre
Remarks. This species was described based on specimens collected by H Mouhot from the Cuming collection. The original description included an illustration and one set of shell measurements. There are four specimens 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 specimen that closely matches the illustration ans has a small label stating "Type" glued inside the umbilicus is figured herein (Fig. 10F).  Fig. 11H, I).

spiniferus
Remarks. The original description did not include an illustration and only one set of shell measurements was given. There are three specimens in the NHM collections purchased from A Morelet with the original label stating "type of spiniferum on left", and with the collection locality "Borneo". The specimen with an attached operculum, that most closely matched with the measurements given in the original description, and is marked with "X" under the shell, is figured herein (Fig. 10H).
Remarks. The original description clearly stated that this taxon was described based on only one specimen. The NHM collections contain a lot consisting of a single specimen with a label in Godwin-Austen's handwriting stating "Type". This specimen matches well with the illustrations and measurements given in the original description, and so we consider this specimen as the holotype fixed by monotypy. Fig. 12A Cyclotus subflammulatus Pfeiffer, 1861: 28. Reeve 1863: volume 14, Cyclotus, pl. 8, species 43. Kobelt 1902.  ; Fig. 12A).

subflammulatus Pfeiffer, 1861
Remarks. The original description did not include an illustration and only one set of shell measurements was given. Pfeiffer (1861) stated that this taxon was described from a specimen collected by AR Wallace in the Cuming collection. Later Reeve (1863) re-published the description and figured a specimen from the Cuming collection. The NHM collections contain a lot consisting of only one specimen from the Cuming collection with a label in Pfeiffer's handwriting stating the species name and collection locality, however this has been subsequently overwritten. This shell matches well with the measurements given in the original description and the illustration in Reeve (1863).  Fig. 11K, L). Remarks. The species description was clearly based on two specimens collected by R Hungerford and housed in the British Museum [= NHM]. The original description included an illustration and gave one set of shell measurements. The NHM collections contain a lot of two specimens with a label stating "Types". The specimen that matches well with the illustration and shell dimensions given in the original description is figured herein (Fig. 10K).
Remarks. The original description included an illustration, one set of shell measurements, and was clearly based on more than one specimen. The author indicated that the 'Type' was kept in the JR le B Tomlin collection and 'Paratypes' were housed in the NHM collections. The holotype was clearly designated and is housed in the JR le B Tomlin collection (later transferred to the NMW in Cardiff). The NHM collections contain two lots, consisting of three shells in total, both containing an original label stating "Paratype". The collection locality matches with the original description. Therefore, we consider these specimens as paratypes. Remarks. The original description by Pfeiffer includes the illustration of a shell and gives one set of shell measurements. The type lot in the NHM collections was collected by "Dr. Trail" and is from the Cuming collection as stated in the original description. It has an original label in Pfeiffer's handwriting giving the species name and collection locality. The largest specimen, marked with an "x" which most closely matches the measurements and the illustration shown in the original description is figured herein (Fig. 12E). Fig. 12G Spiraculum travancoricum Blanford, 1880: 212, 213, pl. 3, figs 6, 6a. Pearsonia travancornica -Kobelt 1902: 175. Raheem et al. 2014.  Fig. 12H). Remarks. The species description was clearly based on more than one specimen. The original description included an illustration of a single specimen and gave two sets of shell measurements. The NHM collections contain a lot consisting of a single shell with an original label stating "type". This shell matches with the illustration and the shell sizes correspond well with the measurements of the specimen cited as "Size of shell drawn", in the original description.  Swainson, 1840 Type locality. unknown. Type material. Syntype NHMUK 20170370 (3 shells; Fig. 12I, J). Remarks. The original description did not include an illustration or collection locality. Pfeiffer stated that this species was described based on material from the Cuming collection, and only one set of shell dimensions was given. Later, Reeve (1863) re-published the description and figured a specimen from the Cuming collection. A specimen lot containg three shells from the Cuming collection with an original label in Pfeiffer's handwriting giving the species name is housed in the NHM collections. The specimen that most closely matches the measurements given in the original description and the illustration in Reeve (1863) is figured herein (Fig. 12I).  ; Fig. 12L).

vanbuensis
Remarks. The original description included only one set of shell measurements; however, the species description was not explicitly based on one specimen. There are two shells in the NHM type lot with an original label stating "Types", subsequently changed to read "holotype red spot". The shell that matches the measurements given in the original description is here designated as the lectotype to stabilise the name. Fig. 13A, B Smith, 1896c: 150, fig. 13. Kobelt 1902 Current generic position. Cyclotus Swainson, 1840 Type locality. Jampea Island [Pulau Jampea, Selayar Islands Regency, south Sulawesi Province, Indonesia].
Remarks. Smith stated in the introduction to his paper that he had received specimens from A Everett. There are six specimens in the NHM type lot with original labels in Smith's handwriting. The original description gives one set of measurements and an illustration of one specimen. The specimen that corresponds to the figure and measurements given in the original description, and with a red spot on the shell is figured herein (Fig. 13A). Fig. 13C, D Cyclostoma volvuloides Sowerby I, 1850: 162*, pl. 31b, figs 312, 313. Pfeiffer 1853b: 249, 250, pl. 33, figs 8, 9. Reeve 1863, volume 14, Cyclotus, pl. 4, species 19. Cyclotus volvuloides -Kobelt 1902.

Current generic position. Cyclotus Swainson, 1840
Type locality. Unknown. Type material. Possible syntype NHMUK 20160354 (3 shells; Fig. 13C, D). Remarks. The original description of this species included an illustration. Later, Pfeiffer (1853b) and Reeve (1863) re-published the description and figured this species based on material in the Cuming collection. The NHM collections contain a lot of three shells from the Cuming collection with original labels giving the taxon name and citing the illustration "f. 312, 313". The specimen in the Cuming collection matches well with the illustration in the original description, Pfeiffer (1853b) and Reeve (1863). However, Sowerby I did not clearly state that the species description was based on specimens from the Cuming collection. Therefore, we consider this lot to be possible syntypes.  13E). Paratypes NHMUK 1912.4.16.629/2-4 (3 shells; Fig. 13F) from the type locality.
Diagnosis. The differences between Pterocyclos anamullayensis new species, P. comatus Beddome 1881 and P. cyclophoroideus Nevill, 1881 are the elevated spire, expanded lip, stout last whorl, absence of a spiral band on periphery of the last whorl, the wingshape of upper peripheral lip near the suture and a canal-like accessory respiratory structure. In comparison, P. comatus and P. cyclophoroideus both have a depressed spire, narrow dark brown peripheral band, wide umbilicus, and slight expansion of the upper peripheral lip. The accessory respiratory structure of P. comatus forms a nearly closed tubular structure (see Raheem et al. 2014: fig. 23d, e), while P. cyclophoroideus has a nearly closed tubular to canal shaped structure (see Raheem et al. 2014: fig. 24b, c).
Description. Shell small to medium, elevated conic, thickened, and widely umbilicated. Apex acute; spire elevated; suture wide and depressed; whorl 4 to 5 convex and becoming increasingly regular. Shell surface nearly smooth with thin growth lines; periostracum thin, corneous to brownish colour. Last whorl rounded and stout. Shell colour monochrome with a white or brown zigzag pattern. Aperture rounded with white lip; upper peripheral-lip widely expanded with wing shaped near suture; lower part of apertural lip thickened and absent to little expanded. Accessory respiratory structure canallike or notch shape. Operculum calcareous, concave inside and multi-lamellae outside.
Etymology. The specific epithet is derived from the type locality. Distribution. This new species known only from the type locality.
Remarks. This new species was described based on the historical collections of RH Beddome from "Anamalis". The NHMUK 1912.4.16.629 lot consists of four shells labelled as "Pterocyclos anamullayensis Bedd." and "Type", but this taxon name has previously never been published.