Check-list of the Neniinae

Hartmut Nordsieck (VI.2012)

General remarks

In the present article a check-list of the clausiliid subfamily Neniinae is given. The range of this subfamily comprises a part of the West Indies (Hispaniola, Puerto Rico) and northern and central Andine South America. The list contains 153 species with 198 subspecies. Six new species taxa which need further examination are not included.
The publication was necessary, because the most recent lists, that of the Peruvian Neniinae in the last revision of the subfamily (Nordsieck 2005) and that of the whole subfamily in my book (Nordsieck 2007), are outdated. This is due to new results concerning the tribe classification and the description of new species taxa (Nordsieck 2010).
The main problem of Neniinae systematics is the imbalance of knowledge of the different groups. While that of the Peruvian Neniinae (plus one species occurring in Argentina) is quite satisfactory the Neniinae fauna of the other countries (Columbia, Venezuela, Ecuador, Bolivia) is nearly unknown. The shell material from these countries is scarce and comes exclusively from collections of the nineteenth century; except for some species from Columbia material for anatomical examinations is not available. Because the borderline between the two tribes, Neniini and Peruiniini, must be expected within the northern countries, the tribe classification remains uncertain.
Another problem, the insufficient knowledge of the localities and distribution of the species, hampers severely the systematic and zoogeographical work on the Neniinae. For many species described in the nineteenth and early twentieth century only inexact or wrong localities were given. For example, the localities of nearly all species collected by Jelski in Peru (and described by Lubomirski or Polinski), if given at all, are wrong. So, one has to do with several species with unknown origin, some belonging apparently to genera of its own, which in the check-list appear as genera incertae sedis.
The shell morphology of the different Neniinae groups is rather uniform, while that of the genitalia is more or less different. Therefore, the relationships of the genera within the subfamily cannot be ascertained without anatomical investigations. The proposed system of the subfamily is mainly based on genital morphology; on the grounds mentioned above only that of the Peruvian species is well-founded. Loosjes & Loosjes-van Bemmel (1966, 1984) examined the genitalia of 32 species, nearly all from Peru. Before, Zilch (1953, 1959) had figured those of two Peruvian species, Hylton Scott (1954) those of the only species from Argentina; afterwards, Thompson (1985, 1998, 2008) described those of the two species from the Caribbean and of two species from other countries of South America . Until now, my own anatomical examinations refer to 38 species, except for seven all from Peru. Altogether, the genital morphology of 61 Neniinae species is known (some of them, however, insufficiently, because subadult specimens or some with atrophied male genitalia have been examined).

Systematical remarks

The subfamily Neniinae is subdivided into two tribes, Neniini and Peruiniini (Nordsieck 2005: 204). The main difference concerns the diverticulum of the bursa copulatrix. In the Neniini it has the same structure as the remaining parts of the organ, in the Peruiniini it is transformed into a glandular tube or missing. The other character in which the tribes were said to differ, the presence or absence of a penial sheath, is not suitable, because the Zilchiella group of the Peruiniini has a penial sheath like the Neniini (result of further investigation, contradicting Nordsieck 2005: 204-205).
As the examination of the genitalia has shown, not only the Caribbean genera Nenia and Nenisca, but also „Nenia“ bequaerti from Sierra de Perija, Columbia / Venezuela frontier region, belongs to the tribe Neniini (Grego & Szekeres 2008, Nordsieck 2010: 49-50). This species is closely related to „Clausilia“ smithiae from Sierra de Santa Marta, Columbia; because of shell morphology both species have been affiliated to the genus Neniops (Nordsieck 2007). Without knowledge of the genitalia, the other Neniops species (karsteniana group) and the related genus Gonionenia (dohrni) are tentatively classified with the Neniini.
The tribe Peruiniini in which the remaining genera of South America are united has been subdivided into several genus groups (Loosjes & Loosjes-van Bemmel 1966, 1984, Nordsieck 2005). The anatomical examination of further Neniinae species since then has shown that the last proposal needs revision. The revised genus groups are the following (for particulars of the genitalia see Nordsieck 2007: 75-77):

Columbinia group:
Shell entire; posterior lower palatal plica mostly present; diverticulum inserted on pedunculus (in Columbinia columbiana on free oviduct) or missing.
Only with plesiomorphic characters (for character evaluation see Nordsieck 2005: 199-200), therefore the grouping may not be natural.
Notes: Neniaptyx has an uncertain systematic position (clausilial apparatus reduced, genitalia unknown).
Columbinia exul from the Venezuela / Brazil frontier region differs from the other examined species by the development of the male genitalia (proximal part of penis strikingly thin; penial retractor inserted far distally) and is therefore placed in a subgenus of its own. Its occurrence is remote from the range of the genus.

Zilchiella group:
Shell entire or decollated; if lunellar fully developed, posterior lower palatal plica present; diverticulum inserted on pedunculus; in contrast to the other groups penial sheath present.

Temesa group:
Shell entire; posterior lower palatal plica missing; diverticulum inserted on pedunculus, basally dilated, or missing.
Note: Because of the reduced clausilial apparatus and the lack of diverticulum of Temesa it is not clear if that genus is in fact closely related to Parabalea.

Incaglaia group:
Shell entire; posterior lower palatal plica missing; diverticulum inserted on pedunculus or missing.
Incaglaia, n. gen. (argentina).
Note: For „Nenia“ argentina a new genus is proposed, because it differs from Incaglaia by the following characters (Hylton Scott 1954, Loosjes & Loosjes-van Bemmel 1966): Diverticulum present; epiphallus not reduced; penial caecum missing. Its range is remote from that of the Incaglaia species.

Ehrmanniella group:
Shell decollated; posterior lower palatal plica missing; diverticulum inserted on pedunculus, in part basally dilated.

Peruinia group:
Shell decollated; posterior lower palatal plica missing; diverticulum inserted on pedunculus or missing. In contrast to the other groups, radula with crescent-shaped teeth.
Note: Symptychiella is possibly related to the Peruinia group. Like in Gracilinenia its diverticulum is missing; a penial caecum is sometimes present. In one species crescent-shaped teeth have been traced (Nordsieck 2007: 77). It differs, however, from the other genera by the entire shell and the G-type-like clausilial apparatus.

Steeriana group:
Shell entire or decollated; posterior lower palatal plica present to missing; diverticulum inserted on free oviduct.
With entire shell: NeniatractaHemicenaLeuconenia; with decollated shell: CylindroneniaAndiniaAndiniastraSteerianaNeniella.


The following list includes all known species and subspecies, with indication of the origin and (as far as known) distribution of the species (in general countries, in Peru also regions = departamentos given).

Neniinae Wenz 1923


Nenia H. & A. Adams 1855:
tridens (Schweigger 1820): Puerto Rico.

Nenisca Rehder 1939:
bartschi (Rehder 1939): Haiti;
franzi Thompson 1998 (fig. 1): Haiti.

Paranenia Rehder 1939:
perarata (E. Martens 1873): Columbia.

Neniops Pilsbry 1926:
archidona (Jousseaume 1900): Ecuador;
barcrofti (Pilsbry 1957): Columbia;
bequaerti (Arias 1953): Columbia, Venezuela;
bogotensis (O. Boettger 1879): Columbia;
geayi (Jousseaume 1900): Venezuela;
karsteniana (Dohrn 1860) (fig. 2): Columbia;
smithiae (Pilsbry 1902): Columbia.

Gonionenia Pilsbry 1926:
dohrni (L. Pfeiffer 1861): Columbia, Venezuela,
rochebrunii (Jousseaume 1900): Columbia.

Peruiniini H. Nordsieck 2005:

Columbinia Polinski 1924:

Columbinia (Columbinia):
adamsiana (L. Pfeiffer 1861): Peru (Junin);
atracta (Pilsbry 1949): Peru (Loreto);
blandiana (L. Pfeiffer 1856): Columbia;
bryantwalkeri (Pilsbry 1922): Peru (Huanuco, Junin);
callangana (Ehrmann 1905): Peru (Junin?);
cocaensis (Jousseaume 1900): Columbia, Ecuador;
columbiana (Polinski 1924) (fig. 3): Columbia;
convexivolvis H. Nordsieck 2005: Peru (Pasco);
elegans H. Nordsieck 2010: Peru (San Martin);
elegantula H. Nordsieck 2010: Peru (San Martin);
epistomium (Küster 1847): Columbia;
gracilis (Pilsbry 1949): Peru (Ucayali?);
marcapatensis H. Nordsieck 2010: Peru (Cuzco);
marshalli (Pilsbry 1926): Bolivia;
perezi (Jousseaume 1887): Ecuador;
riedeli Grego & Szekeres 2008: Columbia;
stylina (Ancey 1887): Columbia;
sublutea (O. Boettger 1909): Peru (Junin).

Columbinia (Steatonenia) Pilsbry 1926:
bartletti (H. Adams 1866): Peru (Ucayali?);
binkiae (Pilsbry 1949): Peru (Cuzco);
cooki (Pilsbry 1919): Peru (Cuzco);
cousini (Jousseaume 1900): Ecuador;
ehrmanni H. Nordsieck 2005: Peru (Junin?);
evae (Sykes 1896): ?;
hemmeni H. Nordsieck 2010: Peru (Pasco);
huancabambensis (Rolle 1904): Peru (Piura?);
juninensis (M. Smith 1943): Peru (Junin);
obesa (Haas 1949): Peru (Loreto);
orbignyi (Ancey 1892): Bolivia;
pachygastris Neubert & Nordsieck 2005: Peru (Piura);
reyrei (Jousseaume 1887): Ecuador;
vasquezi Thompson 1985: Bolivia;
zischkai (Weyrauch 1956): Bolivia.

Columbinia (subgen.):
exul Thompson 2008: Venezuela.

Neniaptyx H. Nordsieck 2007:
buckleyi (Higgins 1872): Ecuador.

Incania Polinski 1922:
anoecia (Ehrmann 1949): Ecuador;
auriculina (Jousseaume 1900): Ecuador;
bourcieri (Küster 1853): Ecuador;
chacaensis (Lubomirski 1879) (fig. 4): Peru (Junin);
crossei (Hidalgo 1869): Ecuador;
florezi Weyrauch 1964: Peru (Cuzco);
jelskii (Polinski 1922): Peru (?);
mariae (Zilch 1954): Peru (Junin);
papillosa Neubert & Nordsieck 2005: ppapillosapimbecilla Neubert & Nordsieck 2005: Peru (Amazonas);
pilsbryi (Sykes 1901): Peru (Junin);
platystoma Neubert & Nordsieck 2005: Peru (Amazonas);
trigonostoma (O. Boettger 1880): Peru (?);
warszewiczi (Polinski 1924): Peru (Cuzco?).

Cyclonenia H. Nordsieck 1999:
albosuturalis Neubert & Nordsieck 2005: Peru (Cajamarca);
alpina H. Nordsieck 1999: Columbia;
boliviana (O. Boettger 1893): Bolivia;
cyclostoma (L. Pfeiffer 1850): Columbia, Venezuela, Ecuador;
geertsi Grego & Szekeres 2004: Peru (Amazonas);
gibber Neubert & Nordsieck 2005: ggibbergsagasteguii H. Nordsieck 2005: Peru (Amazonas, La Libertad);
hemmeni H. Nordsieck 2005: Peru (Cajamarca);
sanmarcos Grego & Szekeres 2004: Peru (Amazonas);
violetta H. Nordsieck 1999: Columbia.

Pfeifferiella Weyrauch 1957:
haasi (Weyrauch 1957): hhaasihmagnifica H. Nordsieck 1999: Peru (Cajamarca);
koepckei (Zilch 1953): Peru (Cajamarca);
subterranea (Weyrauch 1957): Peru (Cajamarca).

Bequaertinenia Weyrauch 1964:

Bequaertinenia (Bequaertinenia):
bequaerti (Weyrauch 1957): Peru (Cajamarca);
delicata Neubert & Nordsieck 2005: Peru (Cajamarca).

Bequaertinenia (Miranenia) Grego & Szekeres 2004:
admirabilis (Loosjes & Loosjes-van Bemmel 1989): Peru (Amazonas).

Zilchiella Weyrauch 1957:
grandiportus Weyrauch 1957(fig. 5): Peru (Cajamarca);
palatalis Neubert & Nordsieck 2005: Peru (Cajamarca);
scala Neubert & Nordsieck 2005: Peru (Cajamarca).

Temesa H. & A. Adams 1855:
clausilioides (Reeve 1849): Peru (Cajamarca);
gradata Neubert & Nordsieck 2005: Peru (Cajamarca).

Parabalea Ancey 1882:
albocostata (Weyrauch 1964): aalbocostataapygmaea (Weyrauch 1964): Peru (Lima);
balnearum (Crawford 1939): Peru (Cuzco);
bicolor (Pilsbry 1949): bbicolorbundaticosta H. Nordsieck 2010: Peru (Junin, Lima);
decimvolvis (Weyrauch 1957): dcrassicostata (Weyrauch 1958), ddecimvolvisdminor (Weyrauch 1964): Peru (Junin);
dohrniana (Nevill 1881 sensu Pilsbry 1949): Peru (Junin);
eka (Pilsbry 1945): Peru (Junin);
gibbosula (H. Nordsieck 2005): ggibbosulaggrisea H. Nordsieck 2010: Peru (Junin);
incarum (Pilsbry 1926): ibreurei (Loosjes & Loosjes-van Bemmel 1984), iincarum: Peru (Junin);
kalinowskii (Haas 1955): Peru (Ayacucho);
latestriata (Weyrauch 1958): l. latestriataldensestriata (H. Nordsieck 2005), lmantaroensis (Weyrauch 1964), lqueroensis H. Nordsieck 2010: Peru (Junin);
omissa (Weyrauch 1957): Peru (Junin);
parcecostata (Polinski 1922): pdulacki Grego & Szekeres 2008, pmeridionalis H. Nordsieck 2010, pparcecostata (fig. 6): Peru (Huanuco, Junin, Huancavelica);
peruviana (L. Pfeiffer 1867): Peru (Puno, Cuzco), Bolivia;
pilsbryi (Weyrauch 1956): plaraosensis (Weyrauch 1960), pperfectecostata (Neubert & Nordsieck 2005), ppilsbryi: Peru (Lima, Ancash);
primigenia (Weyrauch 1960): pprimigeniapzilchi (Weyrauch 1964): Peru (Lima).

Incaglaia Pilsbry 1949:

Incaglaia (Incaglaia) (= Weyrauchiella Loosjes & Loosjes-van Bemmel 1966):
adusta (O. Boettger 1880): aadusta sensu Haas 1955, acallistoglypta (Pilsbry 1949), acuencaensis (Weyrauch 1964), atumens (Haas 1955): Peru (Huancavelica);
andecola (Morelet 1863): Peru (Cuzco);
angrandi (Morelet 1863): aangrandiasoukoupi H. Nordsieck 2010, avariegata H. Nordsieck 2010 (fig. 7): Peru (Cuzco);
dextroversa (Pilsbry 1949): Peru (Huancavelica);
huanucensis (Pilsbry 1949): Peru (Huanuco);
olssoni (Pilsbry 1949): Peru (Huancavelica);
pampasensis (Pilsbry 1910): Peru (Ayacucho, Apurimac).

Incaglaia (Gibbonenia) Zilch 1954:
raimondii (Philippi 1867): Peru (Huancavelica).

n. gen.:
argentina (Hylton Scott 1954): Argentina.

Ehrmanniella Zilch 1949:
boettgeri (Pilsbry 1945): Peru (Junin);
quadrata (O. Boettger 1880): Peru (Junin).

Brevinenia Neubert & Nordsieck 2005:
dedicata (Weyrauch & Zilch 1954): Peru (Huanuco);
richardsi (Grego & Szekeres 2004): Peru (Amazonas).

Andiniella Weyrauch 1958:
flammulata (Loosjes 1957) (fig. 8): fflammulatafunicolor H. Nordsieck 2005: Peru (Junin);
sztolcmani (Polinski 1922): Peru (Junin).

Peruinia Polinski 1922:
peruana (Troschel 1847): pbradina Pilsbry 1945, perythrostoma H. Nordsieck 2010, pflachi (O. Boettger 1889), pperuanaprosenbergi (Preston 1907), psuperba Weyrauch 1960 (fig. 9): Peru (Junin, Pasco);
slosarskii (Lubomirski 1879): Peru (Apurimac?);
tingamariae (Pilsbry 1922): Peru (Huanuco, San Martin).

Pseudogracilinenia Loosjes & Loosjes-van Bemmel 1984:
huallagana (Pilsbry 1949): hamoena Neubert & Nordsieck 2005, hhuallagana: Peru (Huanuco);
?jolyi (O. Boettger 1880): Peru (?);
pulchricosta H. Nordsieck 2010: plamellicosta H. Nordsieck 2010, ppulchricosta: Peru (Pasco).

Gracilinenia Polinski 1922:
eugeniae (Polinski 1922): Peru (Junin);
filocostulata (Lubomirski 1879): faequistriata Weyrauch 1956, ffilocostulata: Peru (Junin);
nitens H. Nordsieck 2005: Peru (Huanuco).

Symptychiella H. Nordsieck 1999:

Symptychiella (Symptychiella):
acuminata H. Nordsieck 2010: Peru (San Martin);
annae H. Nordsieck 1999: Peru (Amazonas);
bilamellata H. Nordsieck 2005: bbilamellatabcostulata H. Nordsieck 2010, bgracilicosta H. Nordsieck 2010, blaevigata H. Nordsieck 2010: Peru (San Martin);
fratermajor H. Nordsieck 2010 (fig. 10): Peru (San Martin).

Symptychiella (Divanenia) H. Nordsieck 2005:
christae H. Nordsieck 2007: Peru (San Martin);
elegantissima H. Nordsieck 2005: Peru (San Martin).

Neniatracta Pilsbry 1926:
belahubbardi (Pilsbry 1922): Peru (San Martin);
exoptata H. Nordsieck 1999: Peru (Junin).

Hemicena Pilsbry 1949:
cerrateae Weyrauch 1958: ccerrateaeccolcabambensis Zilch 1959, cdamianensis Zilch 1959: Peru (Ancash);
polinskiana (Pilsbry 1949): pancashensis Neubert & Nordsieck 2005, ppolinskiana: Peru (Lima, Ancash).

Leuconenia H. Nordsieck 2005:
leucostoma (Neubert & Nordsieck 2005): Peru (Cajamarca).

Cylindronenia Ehrmann (in Pilsbry) 1949:

Cylindronenia (Cylindronenia):
canescens (Polinski 1922): Peru (Cajamarca);
huarangoensis (Zilch 1949): Peru (Cajamarca);
maranhonensis (Albers 1854): mmaranhonensismterrestris (Weyrauch 1964): Peru (Cajamarca);
pangamitoensis (Loosjes & Loosjes-van Bemmel 1989): p.pangamitoensisp.pongoensis H. Nordsieck 2005: Peru (Amazonas).

Cylindronenia (Cylindroneniella) H. Nordsieck 2007:
cicatricosa (Loosjes & Loosjes-van Bemmel 1989): ccicatricosacleimebambensis H. Nordsieck 1999: Peru (Amazonas);
cumulloana (Pilsbry 1949): Peru (Cajamarca);
violacea Neubert & Nordsieck 2005: Peru (Huanuco).

Andiniastra H. Nordsieck 2005:
violascens H. Nordsieck 2005: Peru (Amazonas).

Andinia Polinski 1922:
taczanowskii (Lubomirski 1879): Peru (Cajamarca).

Steeriana Jousseaume 1900:
cajamarcana Weyrauch & Zilch 1954: ccajamarcanacpomabambensis (Loosjes & Loosjes-van Bemmel 1989), csolutilabrum H. Nordsieck 2005, csororminor Neubert & Nordsieck 2005: Peru (Cajamarca);
celendinensis Weyrauch & Zilch 1954: ccelendinensiscisidroensis Weyrauch & Zilch 1954, cminor Weyrauch 1958: Peru (Cajamarca);
malleolata (Philippi 1867) (fig. 11): Peru (Cajamarca);
nivea H. Nordsieck 2005: Peru (La Libertad);
sorormajor Neubert & Nordsieck 2005: Peru (Cajamarca).

Neniella Grego & Szekeres 2004:
gutierrezi Grego & Szekeres 2004: Peru (La Libertad);
macrosoma H. Nordsieck 2010 (fig. 12): Peru (Amazonas);
strictecostata H. Nordsieck 2005: Peru (Amazonas).

Incertae sedis:

gen. 1:
funcki (L. Pfeiffer 1848): Venezuela.

gen. 2:
pusilla (Polinski 1922): Peru (?).

gen. 3:
wagneri (Polinski 1922): Peru (?).

gen. 4:
dichroa (Haas 1929): Bolivia;
magnifica (Sykes 1901): Bolivia.

Figures 1.-12. Selected species of Neniinae. © SGN
Shell height (mm) = H.

1. Nenisca franzi, Haiti, Dept. du Sud, 3 km W Formón (1250 m), paratype, SMF 256445, H = 21.65.
2. Neniops karsteniana, Columbia, Santa Fé de Bogota, NMBE, H = 35.9.
3. Columbinia (C.) columbiana, Columbia, Aguacadal (2000 ft), lectotype, IZPAN, H = 24.6.
4. Incania chacaensis, Peru, Dept. Junin, near Puente Victoria near San Ramon (830 m), SMF 156211, H = 13.85.
5. Zilchiella grandiportus, Peru, Dept. Cajamarca, Peña Rota 8 km NE Bambamarca (2700 m), holotype, SMF 155710, H = 24.7.
6. Parabalea parcecostata meridionalis, Peru, Dept. Huancavelica, Acostambo (km 172 road La Oroya -Huancavelica) (3250 m), holotype, SMF 329360, H = 13.55.

7. Incaglaia (I.) angrandi variegata, Peru, Dept. Cuzco, Valle de Mandor near Aguas Calientes (1930 m), holotype, SMF 329364, H = 15.55.
8. Andiniella flammulata flammulata, Peru, Dept. Junin, Cerro Huilcashpata near Palca, Rio Tarma (3100 m), paratype, SMF 156337a, H = 11.3.
9. Peruinia peruana superba, Peru, Dept. Junin, 2 km from Mina Pichita Caluga near San Ramon (2200 m), lectotype, SMF 156235, H = 32.95.
10. Symptychiella fratermajor, Peru, Dept. San Martin, from Aguas Claras to Cuevas de Aguas Claras (950 m), holotype, SMF 329324, H = 16.2.
11. Steeriana malleolata, Peru, Dept. Cajamarca, Cajamarca, SMF 89483, H = 20.1
12. Neniella macrosoma, Peru, Dept. Amazonas, 13 km from Balsas to Leimebamba (1515 m), holotype, SMF 329317, H = 18.7.

Abbreviations for collections:
IZPAN = Institute of Zoology of the Polish Academy of Sciences Warszawa;
NMBE = Naturhistorisches Museum Bern;
SMF = Forschungsinstitut Senckenberg Frankfurt / Main.


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