Synopsis of Central Andean Orthalicoid land snails (Gastropoda, Stylommatophora), excluding Bulimulidae

Abstract A faunal overview is presented of the molluscan families Amphibulimidae, Megaspiridae, Odontostomidae, Orthalicidae, Simpulopsidae in Bolivia, Ecuador, and Peru. These Central Andean countries are known for their biodiverse malacofauna, of which the superfamily Orthalicoidea takes relatively a large share. In this paper the five families containing 103 (sub)species, for which systematic information (original publication, type locality, type depository, summarizing literature) and distributional records are presented. All species are illustrated by photographs of the type material or, if this could not be located, by a reproduction of the original figure. The following new taxon is introduced: Thaumastus (Thaumastus) sumaqwayqu sp. n. Junior subjective synonyms are established for: Plekocheilus (Sparnotion) Pilsbry, 1944 = Plekocheilus (Eudolichotis) Pilsbry, 1896; Scholvienia (Thomsenia) Strebel, 1910 = Scholvienia Strebel, 1910; Sultana (Trachyorthalicus) Strebel, 1909 = Sultana (Metorthalicus) Pilsbry, 1899; Plekocheilus (Eurytus) conspicuus Pilsbry, 1932 = Thaumastus (Thaumastus) hartwegi (Pfeiffer in Philippi, 1846); Zebra gruneri Strebel, 1909 = Orthalicus maracaibensis (Pfeiffer, 1856); Scholvienia jaspidea minor Strebel, 1910 = Scholvienia alutacea (Reeve, 1850); Bulimus bifasciatus unicolor Philippi, 1869 = Scholvienia brephoides (d’Orbigny, 1835). A new status is given to Plekocheilus mcgintyi ‘Pilsbry’ H.B. Baker, 1963 (subspecies of Bulinus piperitus Sowerby I, 1837); Strophocheilus superstriatus var. prodeflexus Pilsbry, 1895 (subspecies of Bulinus piperitus Sowerby I, 1837); Thaumastus (Quechua) salteri maximus Weyrauch, 1967 (subspecies of Thaumastus (Quechua) olmosensis Zilch, 1954); Pseudoglandina agitata Weyrauch, 1967 (nomen inquirendum). New combinations are: Clathrorthalicus corydon (Crosse, 1869), and Cyclodontina chuquisacana (Marshall, 1930). Lectotypes are now designated for Bulimus incisus Hupé, 1857 and Bulinus piperitus Sowerby I, 1837.


Introduction
Faunal overviews are the keystones of modern biodiversity research, and while for many countries an overview is now available, this remains very fragmentary for most of the Neotropical realm.
The study area for this paper comprises the countries from the Central Andean area, i.e. Ecuador, Peru and Bolivia ( Figure 1A-B). Although some dispersed species descriptions appeared in publications during the early 19th century, the first major contribution was made by Alcide d'Orbigny (1834)(1835)(1836)(1837)(1838)(1839)(1840)(1841)(1842)(1843)(1844)(1845)(1846)(1847)1835) who travelled extensively in Brazil, Bolivia and Peru; details of his itinerary can be found in Papavero (1971), the localities have been transformed to modern geography by Breure (1973). The collections made by Hugh Cuming during his travels in the same era has been elaborated on elsewhere (Breure and Ablett 2011: 4-5). During the mid-19th century other expeditions followed, e.g., a French expedition during 1843-1847 (Hupé 1857) and a Spanish one during 1862-1865 (Hidalgo 1870, 1893a, 1893b, Breure and Araujo 2015. At the same time many individual travellers (e.g., Jørgensen 2015 for travellers in Ecuador) during this era brought smaller or larger collections of shells, from which new taxa were described by various European malacologists; e.g., Albers (1854aAlbers ( , 1854b described new species on the basis of specimens collected by Warszewicz, Morelet (1860Morelet ( , 1863 used collections made by Léoncide Angrand, and Lubomirski (1880) based his descriptions on material from Jelski and Sztolcman. The 161 taxa mentioned in this paper are plotted against publication date in Fig. 1C. The period 1851-1865 stands out with 32 newly introduced taxa.
The aim of this paper is to compile data for part of the Orthalicoidea occurring in the study area, giving systematic information (original publication, type locality, type depository, summarizing literature) and distributional records. A photograph of a type specimen, if located, or an identified specimen by us are presented; if these were not available a copy of a picture from literature is provided.

Methods
This compilation is based on literature (listed in Breure 1979, Breure andSchouten 1985, and recent papers; see Ramírez et al. 2003 for a name list of Peruvian Mollusca species), and distribution records with precise localities only from (unfortunately a limited number of ) verified museum collections (Brussels, Leiden, London). These are listed for each taxon under distribution, the countries of the study area are indicated in bold. We have been reluctant in using unverified data (marked in the text with *) from online databases as misidentifications are possible for many taxa; only if relatively little doubt existed about (easily identifiable) species have these data been used as distribution records in the maps. Altitudes, if not given in the literature for the species, have been taken from Google Earth but should be treated as an estimate. Localities have been mapped with SimpleMappr (Shorthouse 2010), with terrestrial ecoregions layer (see also WWF 2015). The systematic part herein generally follows current understanding and is not to be considered a major revisionary work. Type localities are quoted from the original publication, unless stated otherwise. Diagnoses of supra-specific taxa are tailored to the study area; sizes mentioned (small/medium/large) are relative to the variation within the supra-specific taxon in this area.
Photographs are presented of at least the ventral view of a species and, only if they were available, of other views of the shell. In the legends the shell height (H) is given in millimeters. If available, living specimens are figured to facilitate recognition in the field, but generally pictures of living snails with verified identifications are very scarce. Habitat. May be found in montane and cloud forest, and in páramos; occasionally in pockets of arid vegetation (e.g., Opuntia sp.). The vertical distribution is 1000-4000 m, with an emphasis on 2500-3000 m. Weyrauch, 1967 Figs 2A-C, 14 Plekocheilus (Orcesiellus) tenuissimus Weyrauch 1967: 469, figs 23, 50. Plekocheilus tenuissimus;Richardson 1995: 323 (references). Plekocheilus (Aeropictus) tenuissimus; Breure and Borrero 2008: 7;Borrero and Breure 2011: 15, figs 5A-C;Breure 2012a: 12. Type locality. "Ecuador, Tandayapa, en la vertiente oriental del cerro Pichincha, approximadamento 2500 m". Type material. FML 3364, holotype. Diagnosis. Shell relatively small, with hardly convex whorls, the height of the aperture 0.72 total shell height, suture descending in front, but sharply ascending behind the lip, parietal callus pale greenish-brown.
The loss of the holotype of Bulimus hauxwelli makes it necessary to judge this taxon-and the subgenus Sparnotion-largely on the basis of the figures provided by Pilsbry and the remaining paratype in MCZ. As far as we know there is no material with proven locality data present in other museum collections. However, we recently had the opportunity to re-study the specimen in MCZ on the basis of high resolution pictures supplied by Adam Baldinger. As noted earlier (Breure 1978: 22), the paratype does not show the "longitudinal groups of crowded, finely zigzag hydrophanous lines" very clearly and this could hardly be compared to the subcuticular cavities filled with air characteristic for Plekocheilus (Aeropictus); see also Borrero and Breure 2011: fig. 6 for shell sculptures of several Plekocheilus species. While the paratype shell shows a papillose sculpture of the last whorl (unfortunately not clearly shown on the picture), we think this sculpture is not atypical compared to the known species of Plekocheilus (Eudolichotis). The sprout in the basal lip seems stronger than in Crosse's or Pilsbry's figures; this may be a sign for intra-specific variation. Finally, the narrow and 'not calloused' lip reminds of several Plekocheilus (Eurytus) species and we hardly doubt if this characteristic alone may be sufficient for a subgeneric separation of this species.
When this manuscript was being finalized, we received information about a specimen with locality "Peru" in the RAMM collection. This specimen originates from the collection of Miss J.E. Linter  and is the sole specimen we have been able to trace apart from the type material. This specimen (Figs 12D-F) does show the characteristics that Pilsbry mentioned for the lost holotype. And although there seems some mixing in of a characteristic-zigzag hydrophanous lines-from Plekocheilus (Aeropictus), based on the shell morphology alone we conclude that this species may be best classified as P. (Eudolichotis) hauxwelli untill more material, hopefully allowing for anatomical and molecular studies, becomes available.

Remarks.
A full re-description, based on the lectotype, was given by Breure (1978: 11). The first precise locality for this species is based on two specimens found in the MIZW collection; the material is accompanied by an original label "Chaguarpata (5800´) [1768 m]", and was collected in 1883. Modern gazetteers do not provide any exact name like this for Ecuador; the closest match is Chaguarpamba in Prov. Loja, where altitudes in that range do occur. The (more recent, second) label "Chaguaspata, Peru" seems to be in error as there is no such place in Peru. The specimens have a white lip instead of the pink one in the type specimen. Remarks. The sentence "sur les bords de la rivière de l'Ucuyali" leaves little doubt about the type locality, although there is also a locality named Sarayacu in Ecuador, Prov. Pastaza. We have been unable to locate the type specimens of Deville and Hupé, but close examination of their original figure leads us to believe that the shell exhibits the same longitudinal plication as seen on the holotype of jacksoni Pilsbry. We now tentatively consider the specimens figured by Borrero and Breure (2011: figs 17G-J, as Plekocheilus (Eurytus) piperitus) to be conspecific with Deville and Hupé's species. Pilsbry, 1939Figs 11D-F, 15 Plekocheilus nocturnus Pilsbry 1939H.B. Baker 1963: 229;Richardson 1995: 317 (references Type locality. "Colombia", see remarks.
Ecoregion. Eastern Cordillera real montane forests [NT0121]. Remarks. Clench and Turner (1962: 109) have pointed out that the original locality was erroneous; the type locality should be Nachiyacu. This taxon has been synonymized by Richardson (1995) with Plekocheilus (Eurytus) jimenezi, but without any comments. The shape of the aperture in the type specimen may not be entirely typical; it does not show the ascending suture behind the lip typical of P. (E.) jimenezi, and the aperture is different in shape. Tentatively we have retained this taxon as a separate species, as it may be sympatric with P. (E.) jimenezi, awaiting further studies in the area.

Ecoregion. Ucayali moist forests [NT0174].
Remarks. Borrero and Breure (2011) identified Ecuadorian material as this species, without having seen the type material. However, the type material of Sowerby is somewhat tapering at base and has the suture descending in front ( Fig. 4B-C); also the shells figured by Borrero and Breure seem slightly smaller and slenderer. Therefore we are of the opinion that this taxon is best restricted to Peruvian material, and therefore the specimen figured by Breure and Ablett (2011: fig. 20A-C) is now designated lectotype (design. n.). Bulimus pseudopiperatus J. Moricand, 1858 is considered a junior subjective synonym of Bulinus piperitus Sowerby I, 1837. This was also the opinion of Weyrauch (1967), who provided Pucallpa as the first precise locality for this species. Remarks. This taxon has been considered as a separate species. Upon comparing the type with material of Plekocheilus (Eurytus) piperitus (Sowerby I, 1837), we conclude that mcgyntyi is very similar and consider it herein as subspecies of Sowerby's taxon (stat. n.). The differences seem to consist mainly in P. (E.) piperitus mcgyntyi having a more expanding lip, and a somewhat slenderer shell. The Ecuadorian material mentioned by Borrero and Breure (2011) is now tentatively considered to be this subspecies.  (Baker 1963: 230). Diagnosis. Shell relatively medium-sized, with irregularly spaced reddish-brown dots, sometimes forming longitudal streaks, sculptured with a regular pattern of granules (Fig. 7H), suture descending in front, rapidly descending behind the lip, aperture ovate, peristome simple.
Remarks. This taxon was described as a variety of Plekocheilus (Eurytus) superstriatus (Sowerby III, 1890). Upon comparing the type specimens we see differences in the shell shape, the sculpture of the last whorl at dorsal side (recognizing that Pilsbry's shell is somewhat worn), and the dimensions. Moreover prodeflexus Pilsbry has a descending suture in front. This taxon appears related to both P. (E.) piperitus piperitus (Sowerby I, 1837) and to P. (E.) p. mcgintyi 'Pilsbry' H.B. Baker, 1963, sharing characteristics with both; the differences are but slight and seem to lie mainly in the sculpture of the last whorl. However, since the type specimen is worn, additional material from that area should clarify the possible variation. Tentatively we give it a subspecific status as P. (E.) piperitus prodeflexus (Pilsbry, 1895)  Remarks. So far this species has not been re-found after its original description. The record from Prov. Zamora-Chinchipe, Tapichalaca (Breure and Borrero 2008) refers to a similar but as yet undescribed species.
Dimensions  (Breure 1978: 16). Diagnosis. Shell relatively large, with reddish-brown oblique zigzags on the penultimate whorl, becoming irregularly spaced dots on the last whorl which is sculptured with fine granulation (Fig. 6F Remarks. Pilsbry 1895Pilsbry [1895Pilsbry -1896: 88 regarded this species as closely resembling Plekocheilus (Eurytus) piperitus (Sowerby I, 1837), with which we concur. Also P. (E.) roseolabrum (E.A. Smith, 1877) may be added to this group. P. (E.) taylorianus differs mainly in its larger size and the fine granulation on the last whorl (Fig. 5F). Remarks. Of the two taxa mentioned only type material of the junior subjective synonym Bulimus semipictus Hidalgo has been located.

Subgenus Plekocheilus (Plekocheilus)
Diagnosis. Shell relatively medium-sized, with axial colour streaks of reddish-brown, partly oblique and zigzag, sculptured with axial riblets, becoming malleated on the last whorl and with a dense pattern of oblong granules behind the lip; aperture ovate, peristome expanded and reflexed.
Habitat. The species live in montane and cloud forest in leaf litter, at altitudes of ca. 900-3350 m; the ecology within the study area is unknown. Remarks. This species was described on the basis of material collected by the Comisión Científica del Pacífico with an imprecise locality. While more precise records are awaited, it is suggested that the eastern Cordillera could be a possible location where this species might occur.

Remarks.
During ongoing phylogenetic research Breure and Romero (2012) showed that species attributed to Thaumastus belonged to different monophyletic clades. Consequently they have been classified accordingly. The Brazilian species Thaumastus (Thaumastus) achilles (Pfeiffer, 1853) and T. (T.) largillierti (Philippi, 1845) grouped with Megaspira species and are thus without much doubt placed in the Megaspiridae. As material of the type species Thaumastus (T.) hartwegi (Pfeiffer in Philippi, 1846)-occurring in Ecuador-has not been sequenced, the classification of the Andean species of this group remains tentative. Also for Thaumastus (Thaumastiella) no material could be analysed as yet, and its classification with this family is only provisional. The genus Paeniscutalus is also provisionally arranged under this family, awaiting a further clarification of the findings of Breure and Romero (2012). They found that the sole species classified with this genus, which they suggested to be a relict of an older group, appeared at the very basis of the phylogenetic tree of the Orthalicoidea. Remarks. This species has been considered as a member of the Strophocheilidae due to its general shell shape, with a relatively low spire (Pilsbry 1944c, Haas 1955b. The anatomy and phylogenetic data, however, clearly shows it belongs to the superfamily Orthalicoidea. The mentioning by Schileyko (1999: 282) of "2 spp." within this genus is erroneous as all described taxa are synonymous.
Habitat. Occurring generally in evergreen forest up to ca. 3000 m, where the species live in the leaf litter layer.
Habitat. As far as ecological data are available, the species live in cloud and montane forest, mainly near rocky outcrop. The altitudinal distribution is 0-2300 m, but likely the species are mainly restricted to the upper half of this range in the area treated.
Dimensions Remarks. This species is only known from the type locality and is possibly a short-range endemic. The material referred to by Strebel (1909: 138) must be considered lost.

Ecoregion. Northwestern Andean montane forests [NT0145].
Remarks. This is a quite variable species which Breure and Mogollón (2010) considered identical with Plekocheilus (Eurytus) conspicuus Pilsbry, 1932. They also suggested Thaumastus (T.) flori (Jousseaume, 1897) to be closely related to Thaumastus (T.) hartwegi (Pfeiffer, 1846), which occurs in the same general area. Upon comparison of the type specimens, however, we are now of the opinion that Pilsbry's taxon is a junior subjective synonym of T. (T.) hartwegi, and Jousseaume's taxon is a related but distinct species. The record from Nanegal needs further confirmation.  (1), paralectotype. Dimensions. Shell height 71.5, diameter 37.0 mm. Diagnosis. Shell relatively medium-sized, uniformly brownish with a slightly darker spiral band at the periphery and a yellowish one below the suture, sculptured with spiral rows of oblong granules, suture crenulate, ascending in front, aperture subovate, columellar margin curved and dilated above, peristome white, hardly expanded below, and very narrowly reflexed.
Remarks. Pfeiffer (1844in Küster and Pfeiffer 1840-1865) figured a species clearly unlike the original figure by Sowerby, which he considered a Chilean species; Pfeiffer said his figured specimen was from "Chili und Peru", only the latter locality seems plausible for this species. Reeve (1849Reeve ( [1848Reeve ( -1850) considered his taxon identical to the species figured by Pfeiffer. The name "Vitoe" might be a misspelling for Mito, which is ca. 70 km SE Tarma at ca. 3450 m elevation.
Distribution. Ecuador, Prov. Loja, near Catamayo. Peru, Dept. Piura, Inia (Breure and Mogollón 2010); near Huasimal (Pilsbry 1932 Remarks. This species has only been recorded from the type locality, for which Totolima is now the current name. This is a high-altitude locality (4500 m), which makes it more likely that the species may occur 20-40 km (N)NE where elevations of 2000-2500 m occur; this is the Parque Nacional Isiboro Secure. The synonymization of the Ecuadorian Thaumastus (Atahualpa) brunneus by Richardson (1995 is evidently based upon the opinion of Pilsbry (1932: 391); as Strebel's material was destroyed during World War 2 there is no longer an opportunity for comparison.
Dimensions. Shell height 81.5, diameter 42.0 mm. Distribution. Ecuador, without precise locality. Remarks. The material described by Pfeiffer originated possibly from southern Ecuador. The figured specimen by Strebel (1909) was based on material without locality data. This species is related to Thaumastus (T.) hartwegi (Pfeiffer in Philippi, 1846 Type locality. "in Andibus Novae Granadae". Type material. NHMUK 1975125, one syntype. Diagnosis. Shell relatively medium-sized, with creamy ground colour and brownish axial streaks and blotches at irregular distances, suture crenulate, descending in front, ascending at the insertion of the peristome, which is thickened, hardly expanded below and hardly reflexed.
Remarks. This species has not been found since its original publication. Breure and Borrero (2008) assumed this species to be distributed in southern Ecuador, while Linares and Vera (2012) NHMUK 1907.11.22.24, lectotype (Breure 1978. Diagnosis. Shell relatively large, brownish with a small, somewhat lighter girdle at the periphery, sculptured with incrassate growth striae and spiral striation, most noteable on the upper whorls, suture slightly ascending in front, peristome thin, sinuous, simple. Dimensions. Shell height 78.0, diameter 36.0 mm. Distribution. ?Peru (see remarks). Brazil (Simone 2006). Remarks. This species has been recorded from eastern Brazil by Simone (2006), and its presence in Peru, for which we have not seen any verified material or record, remains doubtful at best.  (Breure 1978).
Diagnosis. A relatively small species of Thaumastus (Thaumastus), characterized, when freshly collected, by the deep brown colour on the last whorl, with a golden hue, with two small brown spiral bands on the upper whorls, one of which is subsutural, the lower one becomes peripheral on last whorls, which have on the upper side a zone of axial bands, the interstices twice as broad.
Description. Shell up to 52.5 mm, 2.0 times as long as wide, imperforate, rather thin, elongate-ovate, with hardly convex sides, with (when fresh) a deep brown colour on the last whorl, with a golden hue, with two small brown spiral bands on the upper whorls, one of which is subsutural, the lower one becomes peripheral on last whorls, which have on the upper side a zone of axial bands, the interstices twice as broad. Protoconch sculptured with fine axial wrinkles, partly bifurcating or anostomsing on lower part of whorl, on the second whorl partly broken up in oblong granules; teleoconch sculptured with incrassate growth striae and very shallow, more or less interrupted, spiral depressions. Whorls up to 5, hardly convex, suture slightly impressed, somewhat crenulate. Aperture narrowly elongate-ovate, pale brown with a whitish lustre inside, 1.6 times longer than wide, 0.5 times the total height, peristome thin and simple, columellar margin slightly curved, receding above, threadlike entering the aperture, parietal callus transparent and thin.

Remarks.
The holotype has lost the outer shell layer on the last whorls during conservation. Also some of the other specimens in the material examined have partially lost this layer.
Etymology. The specific epithet is formed from the Quechua words sumaq (good, beautiful) and wayqu (ravine), referring to the type locality, which is along the river at the basis of Machu Picchu. The epithet is used as a noun in apposition.  (Breure 1978: 32); NHMUK 1975269 (2), paralectotypes.

Thaumastus
Diagnosis. Shell relatively medium-sized, tawny coloured with some axial streaks of (purplish-to reddish-)brown, a light girdle at the periphery, sculptured with growth striae and fine, somewhat undulating, spiral, incised lines, aperture subovate, peristome somewhat thickened and hardly expanded at basal margin.
Dimensions. Shell height 58.0, diameter 25.5 mm. Distribution. Brazil (Simone 2006). Remarks. The record for Peru by Ramírez et al. (2003) of this eastern Brazilian species may be due to a misidentification and needs further confirmation.
Dimensions. Shell height 17.5, diameter 4.75 mm. Distribution. Bolivia, Dept. Chuquisaca. Remarks. In his description Marshall mentioned the protoconch sculpture as "apical and first three whorls are confusedly vertically costulate, malleate and spirally striate". The latter description hints to a protoconch sculpture which is classified by  as Cyclodontina Beck, 1837. Also other characteristics place this species in the vicinity of C. lemoinei (Ancey, 1892). This taxon needs further anatomical and molecular studies to clarify its systematic position. Type locality. "Santa Cruz de la Sierra, Bolivia". Type material. NMW 1955.158.24077 (1), possible syntype. Diagnosis. Shell tawny with whitish, oblique riblets, on lower whorls vermiculate or wrinkled, last whorl tapering, angular around the umbilicus, a deep pit just behind the outer lip, aperture oblique, oblong, with four teeth (moderate parietal lamella, prominent columellar lamella, indistinct basal lamella, large palatal lamella), peristome angular above and at base, expanded (modified after Ancey 1892).

Ecoregion. Chiquitano dry forests [NT0212].
Remarks. There is material in the UF collection (not seen), which is supposedly this species according to their datebase; this material was collected in Dept. Santa Cruz, Prov. Nuflo de Chavez, 32 km W Santa Rosa de la Roca at 545 m elevation (UF 212848). This is the only precise record known to us for this taxon.
Diagnosis. Shell slender and elongate, rather thin, broadly perforate, grayish-tawny coloured, sculptured with growth striae and a faint indication of spiral lines, suture abruptly ascending behind the lip, aperture oblique-ovate, with five teeth (small suprapalatal lamella, large palatal lamella, small basal lamella, prominent columellar lamella entering the aperture, large but relatively thin parietal lamella), peristome thickened, expanded.

Ecoregion. Chiquitano dry forests [NT0212].
Remarks. Breure (2013: 14) discussed the need for an in-depth study of the variation of this wide-ranging species; preferably with anatomical and molecular research. The Bolivian record based on d'Orbigny (1838) "frontières nord de la province de Chiquitos" needs further confirmation. The Bolivian material which is found as Spixia striata in museum collections may need re-identification in the light of the recent split of the two varieties of d'Orbigny.

Type species. Orthalicus wallisi
Diagnosis. Shell ovate-conic, thin, whorls slightly convex, apex rather blunt, height up to ca. 30-45 mm (study area), colour of early whorls uniformly pink, yellowish or greyish-brown, the last whorls with dark radial streaks interrupted by 2-3 light bands, typically on the penultimate whorl with a subsutural band on a lighter ground colour, protoconch pitted, teleoconch with growth striae and delicate spiral lines, aperture ovate, peristome expanded, parietal wall brown (modified after .
Habitat. Probably living in trees, as far as known in cloud forests (Figs 85E-F, 86D-F).
Remarks. This genus is scarcely represented in (historical) collections, and only recently some of its taxa were transferred to it and Strebel's taxon given generic status (Breure and Ablett 2015). Further morphological and molecular studies should clarify its systematic position.
Key to species in the study area Remarks. Crosse did not state on how many specimens his description was based. None of the syntypes found in MNCN correspond exactly with the measurements given by Crosse. However, since no specimens have been located in the MNHN collection, it is assumed that all material was returned by Crosse and is now preserved in Madrid. Breure and Ablett (2015: 39, 45) suggested that this taxon belongs to Clatrorthalicus, and inspection of the MNCN material corroborates this point of view. It may be noted that this species strongly resembles C. phoebus (Pfeiffer, 1863), and further studies of the variation and distribution of both species are needed to fully assess their taxonomic positions as a synonymy might be involved.  (Breure 1979: 30). Diagnosis. Shell ovate, creamy ground colour with few axial streaks and spots of russet-brown, on the last whorls a lighter subsutural band is visible, indistinctly sculptured with growth striae, suture hardly ascending behind the lip, aperture with well expanded and reflexed peristome.
Remarks. This species strongly resembles Clathrorthalicus corydon (Crosse, 1869), being only slightly smaller. Upon further studies both taxa may prove to be synonyms.

Diagnosis.
Shell dextral or sinistral (eniantomorphy), elongate-ovate, solid, shining, height up to ca. 80 mm (study area), corneous or pinkish ground colour, uniformly or (usually) with a dark or light peripheral band (mostly with arrow shaped markings) and axial streaks of reddish-brown, sculptured with growth striae, aperture (narrowly) subovate, peristome simple, parietal and columellar walls dark-brown to blackish.
Habitat. Species of this genus live supposedly most of the time at canopy level in lowland tropical rainforest (W.J.M. Maassen, unpublished data); at occasions they descent downwards and may be found on tree stems or near the ground.
Phylogenetic data. : Corona pfeifferi (Hidalgo, 1869. Remarks. Species of this genus show quite some variation and their distinction is, with some exceptions, difficult as they are often found in low numbers (one or a few shells at most) at a specific locality. Moreover, several species show enantiomorphy, which may add to taxonomic confusion. Distributional records for species in this group thus need to be viewed in this context. Due to their hidden habitat at the canopy level their distribution records probably do not reflect their true occurrence.
The taxonomy of this group is hampered by the fact that a) most species described are morphologically very similar; b) the type material of some species has either not been located or is worn, thus making comparative research difficult; c) intraspecific variation is insufficiently known, and anatomical and molecular data is rare; and d) many records in museum collections often have imprecise localities. Moreover, the distribution of these species over the larger part of the vast continent of South America, with the same species in unverified museum collections reportedly occurring at locations ca. 2500 km apart (e.g., central Bolivia and French Guiana), is puzzling. We regard it as suspicious for two species to occur sympatrically at such distances without distinct differences. For the time being, as many lots in museum collections may have been misidentified, it is here suggested that 1) Corona incisa (Hupé, 1857) is used for occurrences in the southern distribution range (Bolivia, adjacent areas of Peru and Brazil), 2) Corona regalis (Hupé, 1857) for specimens from western Brazil, central and northern Peru, Ecuador and southeastern Colombia, and 3) Corona regina (Férussac, 1823) for records from the northwestern distribution range (Brazil, French Guiana, Suriname). Unverified records from these areas have been plotted as Corona sp. in the distribution maps. Corona pfeifferi (Hidalgo, 1869) is a species that, within the study area, may be unambiguously recognized. The taxonomy of this group thus urgently needs further revision, preferably with molecular research from samples throughout the distribution range.  (4), paralectotypes.
Diagnosis. Shell sinistral or dextral, conic-ovate, solid, changing in ground colour from creamy (top) to tawny (last whorl) with a narrow girdle at the periphery of yellowish, arrow-like markings (>>) and darker sections in between, numerous narrow axial streaks, overlying few broader ones in the basic pattern.
Remarks. This species shows enantiomorphy and its geographic variation needs more study. The morphological differences with Corona regina (Férussac, 1823) seem but marginal, and only a thorough revision may shed further light on the taxonomy of this group.
Habitat. Presumably living in leaf litter in (secondary) forests. Phylogenetic data. Breure and Romero 2012: Kara thompsonii (Pfeiffer, 1845). Remarks. This taxon was given generic status by Breure (2011).  (2015) remarked, this species may be closely allied to Kara thompsonii (Pfeiffer, 1845) and K. yanamensis (Morelet, 1863), and upon further studies may prove to be a synonym of either of these species.

Kara ortiziana (Haas, 1955) Figs 46D-F, 47
Plecocheilus ( Remarks. Haas (1955) characterized this species "by the gloss of its shell, which is without any trace of bands or spots". Remarks. The type locality is likely in the Dept. Cuzco, given the addition of "la vallée de Santa-Anna"; however, it is not mentioned in modern gazetteers. The species was described from specimens denuded of the periostracum. The figured specimen has a trace of it remaining, which suggests the colour pattern described above. Size and shape are very similar to Kara yanamensis (Morelet, 1863), and the variation and distribution of these two taxa need further study. ( Remarks. There are different places called Yanama in Peru, but probably the type locality is found between Abancay and Andahaylas, as L. Angrand, the collector, travelled along this route. The relationship of this species with Kara viriata (Morelet, 1863) needs more study as the differences are but slight.
Description. Shell ovate-conical, imperforate, rather thin, shell height up to ca. 45-75 mm (study area), colour whitish with usually longitudinal or zigzag stripes, and more or less modified by three equidistant spiral bands, surface with incrassate growth lines, sometimes with spiral lines or rarely with weak malleation, protoconch smooth, whorls hardly convex, suture well impressed, aperture (elongate-)ovate, skewed in side view, peristome thin and simple. Remarks. Rehder (1945: 29-31) has elucidated the status of Buccinum zebra Müller, which had obscured the taxonomy of this group due to the high variation and many contradicting interpretations in literature. Nevertheless this genus urgently needs a thorough revision using morphological, anatomical and molecular data from specimens throughout the vaste distribution range. Additional to the species listed below, in some museum collections unidentified material of this genus has been listed. Some of these unverified records, collected from precise localities, are plotted as Orthalicus sp. in Figure 52.  (Reeve, 1849). The Peruvian record is based on the specimens described by Martens as Orthalicus isabellinus, which were collected by Tschudi at an unspecified locality (Breure 2013). Strebel (1909: 29) remarked that these subadult shells were not quite typical; however, we are of the opinion that Martens' and Reeve's taxa may be synonymous, although some doubt remains. The species is evidently widely distributed within the Amazon river basin, although molecular research may show that the Peruvian population is distinct from the eastern forms; in that case Martens' taxon should be resurrected. Remarks. Breure and Ablett (2015) noted that the lectotype, for which the dimensions are given above, is probably not full-grown. Their figures L2i-ii illustrate Orthalicus bensoni (Reeve, 1849); this error is now redressed by figuring the correct shell. Also the specimen of Zebra fulgur Miller, 1878 was subadult as Miller gave the shell height as 50 mm. Cousin (1887) mentioned a specimen from San Nicolás, which was located in the RBINS collection. In the same collection a specimen was found with locality "Cabiloña, montaña / près Ambato"; we have been unable to find this locality in modern gazetteers.  (Simone 2006). Brazil (Simone 2006). French Guiana (Massemin et al. 2009). Suriname (Pilsbry 1899: 136; not in Altena 1975). ?Venezuela (Simone 2006).

Ecoregion. Chiquitano dry forests [NT0212].
Remarks. This Brazilian species has an enormous distribution range given the records mentioned above. Some of these, especially of Venezuela and Colombia, need to be viewed with much suspicion and further evidence is needed as misidentifications are likely. The shell figured by Massemin et al. 2009: pl. 5 fig. D has the stripes more waving and partly confluenced.

Type species. Bulimus iostomus Sowerby I, 1824, by subsequent designation (Martens in Albers, 1860: 227).
Description. Shell ovate-conical, imperforate, rather solid, up to ca. 60-80 mm (study area), groundcolour yellowish to tawny, with longitudinal streaks (or with an irregular zigzag pattern or with irregular spots), protoconch smooth, teleoconch sculptured with growth striae and spiral impressions, which may be either closely set or at larger intervals, aperture ovate, columellar margin straight, peristome expanded and narrowly reflexed.
Distribution. Colombia, Ecuador, Peru. Habitat. As far as known mainly found in leaf litter, but some species are (also) tree-inhabiting.
Diagnosis. Shell upper whorls pointed, spiral sculpture rather strong, cutting the growth striae into oblong granules, height of last whorl ca. 0.8 shell height, aperture without columellar fold.
Distribution. Colombia, Ecuador. Remarks. The Colombian records by Linares and Vera (2012) are based on unverified material and need confirmation as far as they are not adjacent to the distribution range in Ecuador (i.e. their record from Antioquia is likely a misidentification).  Breure and Borrero 2008: 29. Type locality. "Paramba, Ecuador".
Diagnosis. Shell with an irregular pattern of lighter and darker brown stripes, aperture ear-shaped, light coloured inside, parietal callus whitish-transparent.

Quechua olmosensis maxima
Remarks. This taxon was described as a subspecies of Quechua salteri (Sowerby III, 1890); Neubert and Janssen (2004) considered it as a distinct species. Upon comparison with both the types from Quechua salteri and Q. olmosensis (Zilch, 1954), it appears very similar in its external morphology to the latter, and quite distinct to the former. Given the difference in size and the dislocation of the type locality, Weyrauch's taxon is now considered as Quechua olmosensis maxima (stat. n.).

Ecoregion. Peruvian Yungas [NT0153].
Remarks. Sowerby also mentioned a variety which was hardly malleated and reached a larger size. The variation and distribution of this species needs further studies.  Remarks. Zilch compared this species with Thaumastus (Quechua) salteri, but said it differs by having a smaller and slenderer shell, which is less sculptured and with a relatively smaller aperture. Scholvienia Strebel 1910: 20. Scholvienia (Thomsenia) Strebel 1910 n. Type species. Bulimus bitaeniatus Nyst, 1845, by subsequent designation (Pilsbry 1932: 391).

Genus Scholvienia
Description. Shell elongate-ovate, rimate, rather solid, shell height up to ca. 35-62 mm, colour uniformly (chestnut-)brown, in some species with few spiral bands, protoconch with axial, waving riblets, on the lower part becoming split or broken up in wrinkles, teleoconch with incrassate growth striae, in some species becoming thickened at irregular distances, in some species (additionally) crossed by spiral striation, aperture ovate, relatively small, peristome thin and simple, slightly sinuate in side view.
Remarks. Weyrauch (1964: 46) argued that the type locality is probably in error; this species has not been re-found in the Cuzco area nor in Bolivia, and neither has any similar species. The taxa from Philippi and Pilsbry were described from "ad Oroya haud procul ab oppido Tarma", respectively [La] Oroya. The specimens on which Strebel based his Scholvienia jaspidea forma minor were collected at "Quimia", which might be a misspelling for Quenua at ca. 4000 m in the same general area. The shells figured fit into the variation shown by Scholvienia alutacea (Reeve). We found a lot from Yambrasbamba, north of Chachapoyas, which we tentatively refer to this species. Further studies should clarify the distribution and systematic position of Reeve's taxon.
Type material. Not located. Additional material. FMNH 30920, holotype of Thaumastus (Quechua) tetricus Haas, 1951. Diagnosis. Shell with straight sides, russet-brown with two yellowish bands (on the last whorl one subsutural, one peripheral), height of aperture 0.4 times shell height or less.

Scholvienia brephoides (d'Orbigny, 1835) Figs 60E-G, 66
Helix brephoides d' Remarks. Phillipi's variety unicolor was described as an unbanded form from Huaribamba, which is found at ca. 3100 m elevation in Dept. Huancavelica or at ca. 4900 m in Dept. Junín. The former is adjacent to Prov. Huancayo mentioned by Pilsbry (1895Pilsbry ( [1895Pilsbry ( -1896). Philippi compared his taxon to Scholvienia brephoides (d'Orbigny), which is also unicoloured. Philippi's unfigured form-for which no dimensions were given and of which the type has not been located-has been treated as subspecies of the nominate form, but is herein considered as junior subjective synonym of d'Orbigny's species.
Type material. Not located, see remarks.
Dimensions Remarks. Strebel described his species on the basis of material supplied by Rolle. There are several places with the name Huancabamba throughout Peru, but Rolle supplied more often material from the Chanchamayo region. Therefore it is assumed this material originated from (Tingo de) Huanacabamba in Dept. Pasco, which is at ca. 1870 m altitude in the Chanchamayo region. Strebel (1910: 26) remarked that, when the shells were held against bright backlight, one sees one, or more often two, spiral bands that are lighter than the groundcolour. This hints at a possible close relationship of this taxon with Scholvienia bifasciata (Philippi, 1845) or S. iserni (Philippi, 1867). The spiral banding visible in Strebel's original figure (Fig. 65I) may be due to a growth anomaly.
Diagnosis. Shell with hardly convex sides, tawny coloured with some darker patches, sculptured with incrassate growth striae, especially on the last whorl thickened at irregular distances, crossed by spiral lines resulting in oblong granulation, aperture with columellar margin well dilated above.
Ecoregion. Peruvian Yungas [NT0153]. Remarks. Breure (1978: 46) has argued why the type locality might be based on a labelling error. Morelet (1863: 173) attributed juvenile specimens from Andahuaylas to this species. This species has not been recognised in material from southern Peru; the presumed occurrence in Bolivia remains problematic as we have not seen any trusted material from that country that could be assigned to this species.
Habitat Remarks. We follow herein the classification of , awaiting further morphological and molecular studies to ascertain the systematic position of this genus. The record for Panama is based on unverified museum collections. Contrary to Schileyko (1999: 362) we are not aware of any sinistral Sultana species.
Habitat. Not known.
Remarks. The main distinction between Sultana (Metorthalicus) and S. (Trachyorthalicus)-type species Bulimus fraseri Pfeiffer, 1858, by original designation (Strebel 1909: 103)-is a slight difference in the protoconch scultpture, which in the latter subgenus consists of "schräge sich kreuzenden Reihen von Grübchen" (Strebel 1909: 151). The two subgenera are here synonymized after examination of the protoconch sculpture in the type specimens of the two type species; this sculpture proved to be nearly identical.
Most species in this group are represented in museum collections by a low number of specimens, which hampers an in-depth study of their variation. Also the lack of anatomical and phylogenetical data is currently a bottle-neck to fully understand their systematic position.  (NHMUK 1928.12.6.15).

Ecoregion. Napo moist forests [NT0142], Peruvian Yungas [NT0153].
Remarks. Pfeiffer based his description on material from the Cuming collection; Shuttleworth described his material, which he received from Cuming, from "in Andibus Columbiae". At the time of collection of the material, both type localities extended beyond the present-day administrative boundaries of Colombia. The Ecuadorian record is on authority of Martens, and needs confirmation of the material to be conspecific with the Colombian specimens. We found a lot in NHMUK corresponding to this species, with a precise locality in Peru. Distribution. Ecuador, Prov. Azuay, Quebrada Machai (Pilsbry, 1899: 195); ?Prov. El Oro, Mirador (RBINS); Prov. Pastaza, Mera; ibid., Puyo; Prov. Tungurahua, Topo (Breure and Borrero 2008).

Sultana
Diagnosis. Shell with broad dark stripes separated by light coloured, narrow zigzag stripes, a wide umbilical zone paler, two narrow dark bands at the periphery and around the umbilical area, both interrupted by light spots.
Type material. Not located (see Pain 1959). Diagnosis. Shell light to dark brown, with three (indistinct) spiral bands on the last whorl, interrupted by a few, oblique, light-coloured streaks, apex blunt, aperture with a calloused peristome, flesh-to dark-brown coloured on the front side, dirty whitish on the dorsal side.

Ecoregion. Peruvian Yungas [NT0153].
Remarks. Broderip (1828: 223) mentioned in his text that the type material originated from Lieut. Maw, who obtained it at "Toulea, about nine leagues to the eastward of Chachapoyas"; this is evidently the locality presently known as Taulia [-06.1200 S, -077.3700 W].
Dimensions. Shell height 70, diameter 30 mm. Distribution. Peru, Dept. La Libertad, Prov. Santiago de Chuco (?, see remarks). Remarks. The type locality mentioned by Sowerby could not been found in modern gazetteers. Since he was a shell dealer and obtained his specimens through third persons, there might have been a mistake in labeling. In case this assumption is correct, the province of Santiago de Chuco might have been meant; this province is located west of and adjacent to the Marañon river. This species was regarded so far as Orthalicus macandrewi, but is unlike other Orthalicus species in shell shape and colouration. It is now tentatively placed in Sultana (Metorthalicus), but future studies are needed to confirm this.
Type material. ZMB 101825, lectotype (Breure 2013: 31). Additional material. ZMB 111927 (1) (Pfeiffer, 1855) by Richardson (1993) without further comments. Breure and Ablett (2015) followed this opinion, but doubt remained. In the context of this study we prefer to treat Fulton's taxon tentatively as a separate species as it seems to differ by the stouter shell, the more inflated last whorl, and the uniform colour pattern. Remarks. The type material was located in the Lubomirski collection (D. Mierzwa-Szymkowiak, pers. commun., 2012). This species has not been re-collected after its description.
Diagnosis. Shell elongate-ovate, ground colour varying from yellowish to reddish-brown and purplish, with more or less conspicuous longitudinal sinuous streaks and crossed by up to four spiral bands, peristome and parietal callus whitish or pinkish.
Remarks. This taxon is known by the type material only. According to Wood and Gallichan (2008: 46), the subsequent record from Peru by Ancey (1903: 89) was a mistake; there is no verified material to prove this record.
Type material. NHMUK 20100505 (2) Remarks. Cuming's material might have originated in the Province of Moyobamba rather than near the locality of the same name. This taxon was regarded by Richardson (1993) as a junior subjective synonym of Sultana sultana Dillwyn, 1817, possibly reflecting the opinions of Parodiz (1962: 456), who wrote "I am inclined to think that Orthalicus meobambensis Pfeiffer is a synonym", and of Pilsbry (1899: 191) stating "from the description (…) I would think meobambensis identical with the upper Amazonian variety of O. sultana". We agree that both species are closely related, but refrain from this conclusion until the variation of both taxa is better documented.
Description. Shell elongate-ovate to globose, rimate or imperforate, thin, protoconch with fine spiral lines that more or less cut the low, oblique riblets or wrinkles into granules, teleoconch smooth or corrugate, last whorl prominent, suture well impressed, aperture oblique to ovate, peristome thin and simple.

Ecoregion. Northwestern Andean montane forests [NT0145].
Remarks. This taxon has been reported from disjunct localities that are widely separate, at altitudes ranging ca. 700-1500 m. The external morphology is, however, very similar. Miquel (1998: 186) remarked that Simpulopsis (Eudioptus) willineri (Hylton Scott, 1967)-known from Paraguay and northern Argentina-strongly resembles this species. Cuezzo et al. (2013: 184) had it as synonym of Moricand's taxon. The species might also be expected in suitable habitats in Bolivia. Breure (1978: 235) already pointed out that this disjunct distribution needs further investigation and molecular research may show either convergent evolution or a species complex.

Figs 82C, 83
Pseudoglandina agitata Weyrauch, 1967: 486, fig. 53;Neubert and Janssen 2004: 197, pl. 17 fig. 212; Barbosa et al. 2008 Remarks. This species has been synonymized with Simpulopsis (Eudioptus) citrinovitrea by Breure (1978: 235). Upon re-study of the type specimens of Weyrauch's taxon we notice that the holotype seems to be the most adult shell and is different from Moricand's species. The paratypes in SMF (Fig. 82C) and FML are subadult and similar to S. (E.) citrinovitrea. It is possible that this taxon may prove to be a related species, for which anatomical and molecular studies are needed. Weyrauch's taxon is now re-surrected and considered as nomen inquirendum until further studies have clarified its systematic position. Remarks. Férussac did not mention a type locality for this species, which is generally considered to occur in northeastern South America. Gargominy in Massemin et al. (2009: 404) reported it from "Guyane et Brésil (?)", and noticed that it has not been collected alive for more than a century in French Guiana. The lectotype, designated by Tillier (1980: 71, pl. 4 fig. 13), is MNHN 21881 (Figs 40E-G). Six specimens (MNHN 21883), labelled as "var. B com. Howe / Cayenne, haut de Raouza", are labeled as paralectotypes; one specimen proved to be Orthalicus bensoni (Reeve, 1849). The presence of this species in Colombia (Linares and Vera 2012: 156) and Ecuador (Breure and Borrero 2008) are based on museum records which need further confirmation. See below for a record from Peru, and also the remarks above at the genus level. Remarks. This species was described from "l'Amérique méridionale" and subsequent authors have never been able to pinpoint a more precise locality. Pilsbry (1895Pilsbry ( [1895Pilsbry ( -1896: 57) wrote "Compare S[trophocheilus]. brephoides Orb.; S. spixii Reeve; S. spixii Wagner, etc.". As the type material has not been located, this taxon is best treated as nomen inquirendum.

Taxa excluded from the Peruvian malacofauna
The following taxa, arranged alphabetically by species name, are now excluded from the land snail fauna of Peru: Cyclodontina angulata (Wagner, 1827) [Ramiréz et al. 2003: 282, species 658] (type locality "in sylvis ad Solimoès et Purù fluvios"; type material not located); no evidence found for Peruvian material. This Brazilian species is listed as Moricandia angulata in Simone (2006: 171).

Discussion
The frequent papers by Simone and collaborators on the relatively well-known malacofauna of Brazil, regularly describing new species (e.g. Simone 2012, Salvador and Cavallari 2014, Salvador and Simone 2014, Simone 2015, demonstrates that the Neotropical malacofauna is still insufficiently known. This is especially alarming in the light of the ongoing biodiversity loss due to habitat destruction which hampers conservation efforts. In this context the poor knowledge of the malacofauna of Bolivia as shown in this paper makes it relatively a 'white spot' in South America; this calls for an urgent prioritisation for field work being undertaken throughout this country, and subsequent taxonomical studies to bring our knowledge of this fauna up to date. The same applies, although this cannot be detailed in the present paper, for the malacofauna of Paraguay. Although the data for Ecuador and Peru are seemingly more substantial, it may be noted that many taxa are still poorly known, with (very) limited verified distribution records, and their anatomical morphology unknown in most cases. This compilation of data on these families thus can only be seen as a necessarily incomplete attempt to give an overview of this malacofauna, but hopefully will also act as a stimulus for (local) malacologists to further our knowledge.
Given the limited verified distribution records presented in this paper, we refrain from an analysis of ecoregions and endemism comparable to Breure and Borrero (2008: 30-32). It is hoped that the data in this paper will stimulate curators to have their collections checked for additional distribution records, as well as Neotropical malacologists to gather new records, which together will allow a future analysis.

Terrestrial ecoregions
The following ecoregions (see Figure 91)-as currently defined by the World Wildlife Fund, and occurring in the study area-have been used in the text as far as distribution records can be assigned to any of them. It should be noted that the Ecuadorian ones as used by Breure and Borrero (2008) have been partly re-defined or iserni Philippi, 1867-69