Annotated type catalogue of the Orthalicoidea (Mollusca, Gastropoda) in the Royal Belgian Institute of Sciences, Brussels, with descriptions of two new species

Abstract The type status is described of 57 taxa from the superfamily Orthalicoidea in the collection of the Brussels museum. Two new species are described: Stenostylus perturbatus sp. n., and Suniellus adriani sp. n. New lectotypes are designated for Bulimulus (Naesiotus) amastroides Ancey, 1887; Bulimulus blanfordianus Ancey, 1903; Bulimulus montivagus chacoensis Ancey, 1897; Bulimus coloratus Nyst, 1845; Plecochilus dalmasi Dautzenberg, 1900; Placostylus porphyrostomus elata Dautzenberg, 1923; Bulimulus ephippium Ancey, 1904; Bulimus fulminans Nyst, 1843; Bulimus funckii Nyst, 1843; Orphnus thompsoni lutea Cousin, 1887; Bulimus melanocheilus Nyst, 1845; Orphnus thompsoni nigricans Cousin, 1887; Orphnus thompsoni olivacea Cousin, 1887; Bulimulus pollonerae Ancey, 1897; Orphnus thompsoni zebra Cousin, 1887. New combinations are: Bostryx borellii (Ancey, 1897); Bostryx carandaitiensis (Preston, 1907); Protoglyptus mazei (Crosse, 1874); Kuschelenia (Vermiculatus) sanborni (Haas, 1947). New synonymies are established for the following nominal taxa: Orphnus thompsoni var. lutea Cousin, 1887 = Kara thompsonii (Pfeiffer, 1845); Orphnus thompsoni var. nigricans Cousin, 1887 = Kara thompsonii (Pfeiffer, 1845); Thaumastus nystianus var. nigricans Cousin, 1887 = Drymaeus (Drymaeus) nystianus (Pfeiffer, 1853); Orphnus thompsoni var. olivacea Cousin, 1887 = Kara thompsonii (Pfeiffer, 1845); Orphnus thompsoni var. zebra Cousin, 1887 = Kara thompsonii (Pfeiffer, 1845).


Introduction
The Orthalicoidea is a dominant faunal element in the Neotropics (Breure and Mogollón 2010), but also has a number of genera with a Gondwanan distribution (Breure 1979;Herbert and Mitchell 2009;Neubert et al. 2009). The relationships within this group have predominantly been based on morphological data (Breure 1974b, 1979, Breure and Schouten 1985, but are being re-defined by ongoing molecular work Breure and Romero, in preparation). In this superfamily approximately 1750 taxon names are available, which calls for an inventory of as much type material as possible to enable revisionary work. This paper complements previous data on type material for this group in the museums of Paris (Breure 1975b), Zürich (Breure 1976), Frankfurt (Zilch 1971;Neubert and Janssen 2004), Berlin (Köhler 2007), and-forthcoming-in the London museum (Breure and Ablett, unpublished data). The aim of this paper is to present data on the types of Orthalicoidea in the collection of the Royal Belgian Institute of Sciences (hereafter RBINS), Brussels.

the collection
One of the interesting parts of the RBINS malacological collection is the former collection of Philippe Dautzenberg (1849Dautzenberg ( -1935see Lamy 1935 andDuchamps 1999 for a biography). This collection is a rich source for many groups, as Dautzenberg was very keen on the quality of his acquisitions (either by exchange or purchase), and always aimed to have species represented by larger series to allow study of the variation. He befriended many malacologists of his time and thanks to ample financial resources was able to acquire important shell collections whenever they came on the market. In this way, parts of the Ancey collection (see Wood and Gallichan 2008), and the Cousin collection (see below), are now housed in RBINS. Duchamps (1999: 3-19) published an extensive list of collaborators and sources of material for Dautzenberg. In the Dautzenberg archive, autographs were found of the following persons mentioned in this paper, which are relevant to compare handwriting on labels in the Dautzenberg collection: César-Marie-Félix Ancey, parts of whose collection Dautzenberg purchased (Wood and Gallichan 2008); Hugh Coomber Fulton, shell dealer who sold type material to Dautzenberg; Arthur Morelet, who donated several types; Hugh Berthon Preston, also a shell dealer, and well-known for the distribution of much material under manuscript names (see also Adam 1971); Hermann Rolle, another shell dealer of whom Dautzenberg was a client. Excerpts of these autographs are given in Figs 1, 2A-B; examples of labels in their label handwriting may be found in different figures, of taxa related to them. It may be noted, that after the receipt of the Dautzenberg collection by RBINS, some type material has been added through exchange of specimens originally in the Dautzenberg collection from other sources (Van Goethem, pers. commun.). Two other authors need mentioning, who were not referred to by Duchamps (1999). The first is Auguste Cousin, a Frenchman who lived for many years in Ecuador and travelled extensively throughout the country. Although he may be regarded as the "father of Ecuadorian malacology" nothing is known about his life, except that he was born in Ecuador in 1835 and died in Paris, France in 1899 (Correoso, pers. commun.). He published only one, extensive paper on the non-marine malacofauna of that country (Cousin 1887). Through his relationship with Jousseaume, some material was known to exist in the MNHN collection (Breure 1975b). To my surprise, there was ample mate-rial in the Dautzenberg collection. An inventory of the Cousin collection was found by Rose Sablon, RBINS technician, in the Dautzenberg archive. It appeared that Dautzenberg acquired this collection in 1913 (Fig. 2C), but it is not documented from whom.
The second author is Pierre-Henri Nyst, a Belgian paleontologist who in his youth described several species of non-marine molluscs from South America, supplied by Belgian explorers (Dupont 1882: 314). His type material, for many years thought to be lost, was now found in the RBINS collection.

Methods
The following criteria were applied to assess potential type material: (a) the locality fits with the original description (taking into account changes in geographical names due to political or administrative reasons); (b) alleged type material is in accordance with the established understanding of the taxon. In order to fulfill the requirements of article 74 of the International Code of Zoological Nomenclature (ICZN), any lectotype designations herein are to be understood as to have the sole purpose of fixing the status of these specimens as the sole name-bearing types of the nominal taxa, to ensure the name's proper and consistent application, even if this is not explicitly done in every single case but abbreviated as "lectotype designation". Lectotypes are designated herein using the following criteria, in order of preference: (1) the relevant specimen was figured in the original description, or in subsequent revisionary works; (2) if no original figure was published, a specimen was selected that matches as closely as possible the measurements given in the original description.
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. The name of the collector, if given in the original paper, is only mentioned (in italics) if it might give a clue about the type status of material present in the collection. The text of the original, or oldest, label is quoted, together with information from subsequent labels if containing information necessary for a correct interpretation. All labels have been photographed and are figured for future historic reference. The dimensions of the type are quoted, as given in the original paper. Dimensions of the type specimens have been taken with a digital caliper, using the methods figured by Breure (1974a: figs 2-3); measurements up to 10 mm have an accuracy of 0.1 mm, those above 10 mm are accurate to 0.5 mm. Due to improvements in accuracy of Vernier calipers, the measurements given herein are in several cases slightly different from those originally reported. In the case of syntypes, only the largest specimen has been measured. Under type material the RBINS-registration numbers are given; if specimens from different localities are present, the order of the lots corresponds with the information of the different labels. The number of specimens originally available, if quoted by the original author, is mentioned under remarks. Further remarks are 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 the generic scheme of Breure (1979) and the familial arrangement of  and Breure and Romero (in preparation

Systematic list of nominal taxa arranged in generic order
This systematic list follows the arrangement of families as proposed by  and Breure and Romero (in preparation), and the generic order from Breure (1979). Within the families, genera are presented in alphabetical order. Neubert and Janssen (2004) correctly stated that the system of Orthalicoidea still remains in an unsatisfactory state. This is partly due to the sensu lato approach followed for some genera (notably Bostryx and Naesiotus) by Breure (1979), which caused a considerable number of taxa to be affiliated with a relatively low number of genera. The phylogenetic studies of  and Breure and Romero (in preparation) resolve this only partly. As some revisionary work, including phylogenetic studies, related to the status of these genera is ongoing, their sensu lato status is here tentatively retained with the exception of the status of Protoglyptus. This taxon, considered a synonym of Naesiotus by Breure (1979), is herein treated generic status given its separate position in the analysis of Breure and Romero (in preparation). Also the status of Kara, treated as a subgenus of Thaumastus by Breure (1979), is now changed on the basis of their analysis; this taxon is now given generic status. No phylogenetic data have been obtained yet for some other genera treated in this paper (e.g. Dryptus); their familial relationship remains tentative until a more satisfactory arrangement can be presented. Finally, re-interpreting the results of Breure (1978Breure ( , 1979, the genus Scutalus is now restricted to Scutalus (Scutalus) sensu Breure (1979). Two other subgenera, Kuschelenia and Vermiculatus are now considered as Kuschelenia (K.) and K. (Vermiculatus) respectively; Suniellus is herein given generic rank.
At the family level the new scheme of Breure and Romero (in preparation) is followed. They give familial rank to the tribus Simpulopsini Schileyko, 1999, and showed that the Placostylidae Pilsbry, 1946sensu Neubert et al. (2009 (Dautzenberg coll.).

Remarks.
Rolle did not state on how many specimens his description was based upon. However, there is another specimen in the NHM collection which will be designated lectotype (Breure and Ablett, unpublished data). The locality is ambiguous, as there are several places called "Huancabamba" in Peru. There were, however, several other species described by Rolle from the same locality, among them Columbinia huancabambensis, which is regarded by Loosjes and Loosjes-van Bemmel (1984: 33) as occurring in northern Peru. Another species described in the same paper, Systrophia moellendorffii, was said by Haas (1955: 367) to have been rediscovered in the Chanchamayo valley [Dept. Pasco]. These, and the fact that a third Rolle species, Newboldius illustris is known to occur in the same region, makes Dept. Pasco, Huancabamba likely to be the locality where Rolle's taxa occur.

Thaumastus alausiensis
Remarks. The material which Breure (1975) used to select a lectotype, came from the Jousseaume collection and originated from Cousin. However, the locality was imprecise ("Équateur" [Ecuador]). At that time, no other material from Cousin was known. In RBINS, material originating from Cousin and exactly corresponding to the type locality has been found. Therefore it may now be questioned if the Jousseaume material was validly regarded as the prime type material. However, in the inventory of Cousin's collection, a total of 30 specimens is mentioned and it cannot be excluded that Jousseaume's material originated from the Cousin collection.
Type material. RBINS/MT2336, three paralectotypes, Baer leg. (Dautzenberg coll.). Remarks. Dautzenberg (1901a) states in his paper that his description is based on four specimens. The holotype designation of a specimen in the MNHN collection (Fischer-Piette 1950) has to be interpreted as lectotype designation (Art. 74.6 ICZN); the measurements of this specimen closely match those given by Dautzenberg (Breure 1975b). The type locality is probably Jocos in Dept. La Libertad [8°14' S 77°28' W].
Remarks. Ancey did not state on how many specimens his description was based. Therefore, the specimen is considered a syntype.
Remarks. Ancey did not state on how many specimens his description was based. Despite the fact that the specimen is slightly smaller than published by Ancey, there is enough evidence to consider this shell as from the original type series. It is now designated lectotype.
Remarks. Wood and Gallichan (2008) discussed the different labels of Fulton, Ancey and Dautzenberg. They argued for a syntypic status of this specimen, and for the material that is in the MNHN collection, which was regarded as the holotype by Fischer-Piette (1950); according to Art. 74.6 ICZN the MNHN specimen has to be regarded as a lectotype. The shell height of the MNHN-specimen is 22.0 mm, thus not corresponding to the original measurements given by Ancey. Breure (1975b) followed the designation as holotype by Fischer-Piette, being unaware of a second specimen in RBINS. The top of this specimen is damaged and it is likely that the original shell height was closer to Ancey's dimensions.
Remarks. The specimens are smaller than the measurements given by Ancey, although they are from the type locality. This taxon was hitherto classified as Drymaeus. However, the sculpture of the protoconch shows very fine spiral lines and dispersed granules, more or less axially arranged. It thus belongs to Bostryx (s.l.).
Current systematic position. Bulimulidae, Bostryx borellii (Ancey, 1897) (comb. n.). Remarks. This taxon was hitherto regarded a Drymaeus species, but the protoconch is smooth (not worn) in both specimens examined. It is here tentatively placed in Bostryx, despite the fact that the colour pattern of the specimen in RBINS is unlike the ones normally found in this genus.

Bulimulus montivagus chacoensis
Remarks. The RBINS material corresponds with the original measurements and is here designated lectotype. The shell shape, especially the flaring basal lip, and the number of whorls (8+) are not typical for Naesiotus, and this species is only tentatively placed here. Miquel (1989: 62) suggested a possible synonymy with N. rocayanus (d'Orbigny, 1835); this has to be ascertained by further comparison of type material.
Remarks. The specimen is not accompanied by an original Preston label. As Dautzenberg always accurately documented on his labels the source and date of his acquisition (in this case "Preston 14.xi.07"), there is hardly any doubt that this is an original type specimen. The protoconch is smooth, confirming that this taxon should be classified within Bostryx (cf. Breure 1979: 52).

Otostomus colimensis
Remarks. This material is not accompanied by an original Rolle label, but Dautzenberg documented that he received the shells from him on 16.vii.1907. The fact that Rolle was a dealer may account for the delay between the time of publication and the acquisitions by Dautzenberg. The material is from the type locality and is here considered as syntypes. Köhler (2007) selected a lectotype from the ZMB material, thus these specimens are paralectotypes.
Current systematic position. Bulimulidae, Drymaeus (Drymaeus) colimensis (Rolle, 1904). Remarks. The type material was found in the RBINS collection and is now figured for the first time since the original publication. Of the two specimens present, one shows the 'yellow shadow' which is characteristic for this species, and is here designated lectotype. The taxon is only known from confirmed localities in northern Colombia; the original locality, which is in Venezuela, Edo. Sucre, seems erroneous. The type locality is now restricted to Sierra Nevada de Santa Marta.

Plecochilus dalmasi
Remarks. Dautzenberg did not state on how many specimens his description was based, but one of the specimens corresponds to the original dimensions and agrees with his figure. This shell is here designated lectotype. The label states that it was collected on 15.iii.1896, which corresponds with the date of excursion to Sierra de Santa Marta mentioned in the station list (Dautzenberg 1900: 147).

Placostylus porphyrostomus elata
Current systematic position. Bulimulidae, Bostryx emaciatus (Morelet, 1863). Remarks. Breure (1978) mentioned that he had found syntypes in both the NHM and RBINS collections and figured the species for the first time. He redescribed the species but noted that its classification remained doubtful. Nevertheless he arranged it under Bulimulus in his 1979 revision. Upon restudying both specimens, it is clear that the protoconch sculpture is quite different from the pattern normal for that genus; especially the thin spiral lines give a strong hint that it should be separated. For that reason it cannot be classified with Rhinus either, although it bears resemblance in shell shape to species of that genus. It is now considered as Simpulopsis (Eudioptus), where it is the largest species. The RBINS specimen is here designated as lectotype.

Bulimulus ephippium
Current systematic position. Simpulopsidae, Simpulopsis (Eudioptus) ephippium (Ancey, 1904). Remarks. The specimen corresponds to the original measurements given by Nyst, whose label is dated 1874 and indicated "type". As there is no evidence that the description of Nyst was based on one specimen, it is here designated lectotype.

Bulimus fulminans
Current systematic position. Amphibulimidae, Plekocheilus (Plekocheilus) fulminans (Nyst, 1843).  (1843) did not indicate that he only had an unique type at hand. In his label dated 1874, he indicated this specimen as "type"; it is now designated as lectotype.

Bulimus funckii
Current systematic position. Amphibulimidae, Dryptus funckii (Nyst, 1843).   Pilsbry and Ferriss (in Ferris 1924) state they have selected a specimen as type, which according to H.B. Baker (1962) is holotype ANSP 84627a. As the label states that the material was collected by Ferriss in 1924, it probably belongs to the original type series. The locality-although more vaguely formulated on the labelcorresponds to the area mentioned by Ferriss (1924) for the type series.

Bulimulus (Ataxus) huayaboensis
Remarks. The holotype designation of a specimen in the MNHN collection (Fischer-Piette 1950) has to be interpreted as lectotype designation (Art. 74.6 ICZN); the measurements of this specimen closely match those given by Dautzenberg (Breure 1975b).
Remarks. Ancey (1892) states that he had seen two specimens. The whereabouts of the other syntype is unknown.
Remarks. The lectotype is in the MNHN collection (Breure 1975b). According to the inventory of Cousin's collection, there were originally 35 specimens present.

Orphnus thompsoni nigricans
Remarks. The original Cousin label mentions the taxon name only; the locality has been added by Dautzenberg, probably on the basis of Cousin's paper. One specimen was found with the Cousin label stuffed inside the aperture and is here designated lectotype. The taxon is considered a junior subjective synonym of Bulimus thompsonii Pfeiffer, 1845 (syn. n.). Current systematic position. Orthalicidae, Kara thompsonii (Pfeiffer, 1845) (comb. n.). Remarks. The original Cousin label mentions the taxon name only; the locality has been added by Dautzenberg, probably on the basis of Cousin's paper. One specimen was found with the Cousin label stuffed inside the aperture and is here designated lectotype. The taxon is considered a junior subjective synonym of Bulimus thompsonii Pfeiffer, 1845 (syn. n.). Current systematic position. Orthalicidae, Kara thompsonii (Pfeiffer, 1845) (comb. n.).  (Dautzenberg and Bernier, 1901).  (Breure 1979). These species are known from Peru and Bolivia (Breure 1978(Breure , 1979; the new taxon described herein thus extends the distribution range further north. In shell morphology the species may be confused with those of Stenostylus, but the protoconch sculpture separates the two taxa: a grating sculpture in Stenostylus; axial wrinkles, which may be partly anastomosing, in Suniellus. Description. Shell 32.5 mm, 1.86 times as long as wide, elongate-ovate, with hardly convex sides, imperforate; very thin. Colour yellowish-olivaceus with brownish streaks, especially on last whorl. Upper whorls lighter in colour. Surface rather shining, with irregularly spaced, thickened growth striae, in between with weaker incrassations and, especially at upper part of last whorl, some oblong granules. Protoconch eroded, but on the dorsal side traces visible of axial wrinkles, partly anastomosing. Whorls 5.7, hardly convex, the last 0.81 times total shell height, a little saccate; suture well impressed. Aperture elongate-ovate, 0.54 times shell height, approx. 1.4 times as long as wide, with a pearly lustre inside. Peristome thin and simple, columella thread-like, somewhat dilated above and appressed at the transition to the parietal wall, which bears a hardly noticeable callus.
Remarks. The lip is damaged at the upper palatal side and the insertion to the shell has been torn off, leaving some tears at the upper part of the last whorl, just behind the lip. The type locality could not be found in modern gazetteers; it is a mixture of Spanish and Quechua words, and may not be officially known under this name. It is likely to be in the páramo area, as Cousin on a second label has written "pie de la nieve / Frutillas por arriba" [at the foot of the snow / Strawberries above]; snow in the 19th century probably occurred at lower elevations than today, but an elevation for this locality of above 2750 m may be a safe guess (González, pers. commun.). There is only one specimen known.
Etymology. The epithet is a patronym in honour of Adrián González-a Cuban malacologist now living in Quito, Ecuador-in recognition for his contributions to Neotropical malacology by his photographic work, his books aimed at a general public, and his dedicated fieldwork.