Garnieriinae, Neniinae

Garnieriinae, Neniinae

Garnieriinae, Neniinae

Hartmut Nordsieck (VI. 2022)

Check-list of the Garnieriinae

In the present chapter a check-list of the clausiliid subfamily Garnieriinae is given, together with diagnoses of the tribe and genus taxa and remarks to distribution and phylogeny. The range of the subfamily comprises a part of South East Asia (Myanmar, Laos, Vietnam, southern China).
The publication was necessary, because the most recent list, that of the subfamily in my book (Nordsieck 2007), is outdated. This is due to new results concerning genital anatomy and the description of new species taxa.
Until now, my own anatomical examinations refer to 15 species. Altogether, the genital morphology of 18 Garnieriinae species is known

Tribe and genus diagnoses

In the following shell and genital diagnoses of the tribes and genera of the subfamily are given.
For the genital diagnoses of the genera the examined species are listed. Most species have been examined personally. For an example of preparation see Fig. 1. All preparations were measured as exactly as possible; the information on lengths of genital organs in the diagnoses are based on these measurements. If the species have not been examined by me, the respective publications are given.
Note: In Nordsieck (2007: 73, table 1) the transitional part epiphallus-penis is named most proximal part of penis.

Fig. 1: Neniauchenia (Ptychauchenia) panhai, Vietnam, Quang Binh, genital apparatus.

Abbreviations: adb = apical part of diverticulum; ag = albumen gland; at = genital atrium; bb = bursa; bdb = basal part of diverticulum; dp = distal part of penis; ep = epiphallus; fod = free oviduct; hd = hermaphrodite duct; pb = pedunculus; pc = penial caecum; pp = proximal part of penis; rp = penial retractor; spo (od) = spermoviduct (oviduct); spo (pro) = spermoviduct (prostate); v = vagina; vd = vas deferens.

Garnieriinae C. Boettger 1926


Shell semiapostrophic; superior lamella separated from spiral lamella.
Penial retractor inserting on epiphallus; penis not bipartite.

Garnieria Bourguignat 1877:

Inferior lamella near to superior lamella;  posterior lower palatal plica present, anterior lower palatal plica short or absent; clausilium plate normally developed or transformed (creased and bent up to inferior lamella).
Bursa not bent off spermoviduct; diverticulum inserting on pedunculus, bipartite, terminal part bent off spermoviduct, proximal part of pedunculus moderately long, distal part short; penial caecum absent; penial retractor inserting on epiphallus nearly at its proximal end; flagellum absent.
Garnieria (G.) mouhoti.
Páll-Gergely & Szekeres 2017 (: fig. 4 c-d): Garnieria (Doducsangia) nhuongi.

Note: The descriptions and figures given by Páll-Gergely & Szekeres (: fig. 4) do not fit the results of my examination. In my preparations of animals in normal condition the diverticulum is longer than the proximal part of pedunculus and much longer than its distal part. Besides the terminal part of diverticulum is bent off spermoviduct. Vagina and epiphallus are shorter than the penis (see also Nordsieck 2007: 73, table 1).

Tropidaucheniini H. Nordsieck 2002:

Shell apostrophic; superior lamella continuous with spiral lamella, in several groups superior lamella and inferior lamella fused.
Penial retractor inserting on penis; penis bipartite.

Megalauchenia H. Nordsieck 2007:

Inferior lamella near to superior-spiral lamella; upper palatal plica connected with lunella by a long arch, in part with posterior lower palatal plica, anterior lower palatal plica absent; clausilum plate ± broad.
Bursa not bent off spermoviduct; diverticulum inserting on pedunculus, with same structure as pedunculus, proximal part of pedunculus short, distal part very short; transitional part epiphallus-penis absent, penial caecum present; penial retractor inserting proximally on penis; flagellum present.
Megalauchenia proctostoma.

Euryauchenia H. Nordsieck 2007:

Superior lamella fused with inferior lamella, spiral lamella distant from superior-inferior lamella; posterior lower palatal plica short, anterior lower palatal plica absent, rarely rudimentary; clausilium plate ± broad.
Bursa not bent off spermoviduct; diverticulum inserting on pedunculus near to its distal end, divided in basal and apical part, apical part thick, proximal part of pedunculus long, distal part nearly absent; transitional part epiphallus-penis present, indistinctly delimited, penial caecum present; penial retractor inserting proximally on penis; flagellum present.
Euryauchenia demangei, E. fischeri.

Tropidauchenia Lindholm 1924:

Inferior lamella near to superior-spiral lamella or superior lamella and inferior lamella fused, spiral lamella near to superior-inferior lamella; posterior lower palatal plica short or absent, in part separated from lunella, anterior lower palatal plica present or absent; clausilium plate not broad.
Bursa bent off spermoviduct; diverticulum inserting on pedunculus near to or at its distal end, divided into basal and apical part, apical part narrow tube, proximal part of pedunculus long, distal part very short or absent; transitional part epiphallus-penis present, penial caecum present; penial retractor inserting proximally on penis; flagellum present or absent.
Tropidauchenia bavayi, T. dorri, T. giardi, T. orientalis.

Indonenia Ehrmann 1927:

Inferior lamella near to superior-spiral lamella; posterior lower palatal plica present, anterior lower palatal plica absent; clausilium plate ± broad.
Bursa bent off spermoviduct?; diverticulum absent, pedunculus long; transitional part epiphallus-penis present?, penial caecum present; penial retractor inserting on middle of penis; flagellum?
Grego et al. 2021 (: fig. 3): I. admirabilis. The characters given with question marks are not recognizable in the figure.

 Neniauchenia H. Nordsieck 2002:


Inferior lamella ± distant from superior-spiral lamella; posterior lower palatal plica and anterior lower palatal plica present, often ± separated from lunella, posterior lower palatal plica rarely rudimentary; clausilium plate not broad.
Genitalia like Ptychauchenia.
Grego et al. 2014 (: fig. 1): N. (N.) steffeki.

(Ptychauchenia) H. Nordsieck 2010:

Superior lamella and inferior lamella fused, spiral lamella distant from superior-inferior lamella; posterior lower palatal plica connected with lunella, anterior lower palatal plica long; clausilium plate not broad.
Bursa bent off spermoviduct; diverticulum inserting on free oviduct, divided into basal and apical part, apical part narrow tube, pedunculus long; distal part of epiphallus forming a loop, transitional part epiphallus-penis indistinct, penial caecum present; penial retractor inserting proximally on penis; flagellum absent.
N. (Ptychauchenia) panhai.

Note: Neniauchenia differs from Grandinenia by the shorter penis and longer epiphallus (longer than penis). All other genera of Tropidaucheniini inclusive of Grandinenia have a shorter epiphallus (shorter than penis). Therefore, the union of Neniauchenia and Grandinenia to one genus, as proposed by Grego et al. (2014), is inappropriate. Also, shell characters (palatal plicae in Neniauchenia more plica-like, see Figs. 2-4) and distribution (separated by a gap) speak for a separation as genera.

Grandinenia Minato & Chen 1984:

Inferior lamella ± distant from superior-spiral lamella; posterior lower palatal plica short or absent, anterior lower palatal plica present or absent; clausilium plate not broad.
Bursa bent off spermoviduct; diverticulum inserting on free oviduct, divided into basal and apical part, apical part narrow tube, pedunculus long; transitional part epiphallus-penis present, penial caecum present or reduced; penial retractor inserting proximally on penis (G. ardouiniana) or on middle of penis (Chinese Grandinenia species); flagellum present or absent.
Grandinenia ardouiniana, G. fuchsi, G. gastrum, G. mirifica, G. pseudofuchsi, G. takagii.

Figs. 2-4: Grandinenia and Neniauchenia species.

Shell height (mm) = H.

2. Grandinenia g. gastrum, China, Guangxi, 5 km NW of Gu Zhang, holotype, SMF 326120, H 24.8.
3. Neniauchenia (N.) tonkinensis, Vietnam, Quang Binh, holotype, SMF 331370, H 32.95.
4. Neniauchenia (Ptychauchenia) panhai euclista, Vietnam, Quang Binh, 14 km from Deo Da Deo to Pheo, holotype, SMF 339699, H 37.3.

Distribution and phylogeny

The tribe Garnieriini with its only genus Garnieria is distributed in N. Laos with adjacent Yunnan and S. Tonkin (Son La). The tribe Tropidaucheniini covers the whole range of the subfamily. Megalauchenia and Euryauchenia are both restricted to a range in the frontier region of Tonkin and Annam (mainly Hoa Binh, Ninh Binh, Thanh Hoa). Tropidauchenia is distributed from N. E. Tonkin (borderline Ha Giang – Ha Noi) to  Yunnan, Guangxi and adjacent Guangdong. Indonenia occurs in the Shan region of Myanmar. Neniauchenia (with Ptychauchenia) is distributed in S. Laos and N. Annam (Nghe An, Quang Binh), Grandinenia in coastal E. Tonkin (from Hai Phong), Hainan and with a striking diversity in Guangxi. Both genera are vicariant; they are separated by a considerable gap in Tonkin.
In the DNA analysis of Uit de Weerd & Gittenberger (2013) only three Garnieriinae species are included, one Garnieria species and two Grandinenia species. In their trees (except of the H3 tree) they form a strongly supported clade; in the H4 and the consensus tree (: fig. 3) Garnieria is sister group of the subclade including the Grandinenia species. According to the chronogram of the analysis (: fig. 3) the sister groups separated in the Miocene; thus, the rank as tribes Garnieriini and Tropidaucheniini is confirmed.
Until further DNA studies, for a reconstruction of the phylogeny within the Tropidaucheniini the genital characters are decisive. Megalauchenia is regarded as basal, because its diverticulum has the same structure as the other parts of the bursa copulatrix, which is certainly the plesiomorphic condition. The other genera have a transformed (bipartite) diverticulum which is apomorphic. In Euryauchenia both parts are thick, while in Tropidauchenia and the Grandinenia group (Neniauchenia, Grandinenia) the apical part is transformed to a narrow tube. In Tropidauchenia the diverticulum is inserting on the pedunculus, in the Grandinenia group on the free oviduct, which is the more apomorphic condition. Within that group both genera are vicariant and can therefore be regarded as sister groups. Indonenia lacks the diverticulum, so that no conclusion based on the genitalia is possible.


The following list includes all known species and subspecies, with indication of distribution of the species.

Garnieriinae C. Boettger 1926


Garnieria Bourguignat 1877:

Garnieria (Garnieria):
mouhoti (L. Pfeiffer 1862): m. moellendorffi H. Nordsieck 2002, m. mouhoti, m. yunnancola H. Nordsieck 2012: Laos, China (Yunnan);
saurini H. Nordsieck 2002: Laos.

Garnieria (Doducsangia) Páll-Gergely & Szekeres 2017:
nhuongi  Do 2015: Vietnam.

 Garnieria (Progarnieria) H. Nordsieck 2012:
huleschheliae Grego & Szekeres 2011: Laos.

Tropidaucheniini H. Nordsieck 2002:

Megalauchenia H. Nordsieck 2007:
proctostoma (Mabille 1889): p. donggiaoensis (H. Nordsieck 2002), p. forceps (Loosjes & Loosjes-van Bemmel 1973), p. proctostoma: Vietnam;
soror H. Nordsieck 2011: Vietnam;
yongshuae H. Nordsieck 2007: Vietnam.

Euryauchenia H. Nordsieck 2007:
demangei (Bavay & Dautzenberg 1909): d. demangei, d. dextroversa H. Nordsieck 2014: Vietnam;
fischeri (H. Nordsieck 2002): f. fischeri, f. reticulata (H. Nordsieck 2002): Vietnam.

Tropidauchenia Lindholm 1924:
bavayi (Lindholm 1924): b. bavayi, b. senescens H. Nordsieck 2011: Vietnam;
danjuan Qiu 2021: China (Guangxi);
dorri (Bavay & Dautzenberg 1899): d. adusta H. Nordsieck 2012, d. adustella H. Nordsieck 2012, d. dorri: Vietnam, China (Guangxi);
giardi (Fischer 1898): g. brunnea H. Nordsieck 2002, g. citrina H. Nordsieck 2002, g. giardi: Vietnam, China (Guangxi);
hitomiae H. Nordsieck 2007: h. hitomiae, h. rufescens H. Nordsieck 2007: China (Guangxi);
lucida H. Nordsieck 2007: l. gracillima H. Nordsieck 2007, l. lucida: China (Guangxi);
nakaharai H. Nordsieck 2007: China (Guangxi);
napoensis H. Nordsieck 2007: China (Guangxi);
ootanii H. Nordsieck 2007: o. longicollis H. Nordsieck 2011, o. ootanii: Vietnam, China (Yunnan);
orientalis (Mabille 1887): o. messageri (Bavay & Dautzenberg 1899), o. orientalis, o. rufocincta H. Nordsieck 2007: Vietnam, China (Guangxi);
palatalis H. Nordsieck 2011: Vietnam;
parasulcicollis Qiu 2021: Cina (Guangxi);
sulcicollis Grego & Szekeres 2017: China (Guangxi);
yanghaoi Grego & Szekeres 2017: China (Guangdong).

Indonenia Ehrmann 1927:
admirabilis Grego & Szekeres 2021: Myanmar;
excellens (H. Nordsieck 2002): Myanmar;
masoni (Theobald 1864): Myanmar;
tuba (Hanley 1868): Myanmar.

Neniauchenia H. Nordsieck 2002:

 Neniauchenia (Neniauchenia):
amoena (H. Nordsieck 2002): Laos;
dautzenbergi (Morlet 1893):d. abdoui (Grego & Szekeres 2019),  d. dautzenbergi, d. decollata (H. Nordsieck 2002): Laos;
gabijakabi (Grego & Szekeres 2014): Vietnam;
muratovi (Grego & Szekeres 2019): Laos;
rugifera (Moellendorff 1898): Laos;
steffeki (Grego & Szekeres 2014): Vietnam;
tonkinensis H. Nordsieck 2010: Vietnam, Laos.

Neniauchenia (Ptychauchenia) H. Nordsieck 2010:
panhai (H. Nordsieck 2010): p. euclista (H. Nordsieck 2016), p. panhai: Vietnam.

Grandinenia Minato & Chen 1984:
ardouiniana (Heude 1885): Vietnam;
crassilabris H. Nordsieck 2016: China (Guangxi);
fuchsi (Gredler 1883): f. cinderella (H. Nordsieck 2002), f. costicollis H. Nordsieck 2007, f. fuchsi, f. pseudotigris H. Nordsieck 2012, f. pygmaea H. Nordsieck 2007, f. tigris (Chang 2005): China (Guangxi);
gastrum (H. Nordsieck 2005): g. costigera (H. Nordsieck 2005), g. gastrum, g. laticosta (H. Nordsieck 2005), g. rubens (H. Nordsieck 2005): China (Guangxi);
ignea H. Nordsieck 2012: China (Guangxi);
magnilabris H. Nordsieck 2012: China (Guangxi);
maroskoi Grego & Szekeres 2011: m. gigantea H. Nordsieck 2012, m. maroskoi: China (Guangxi);
mirifica (Chen & Gao 1982): China (Guangxi);
ookuboi (H. Nordsieck 2005): o. ookuboi, o. pulchricosta H. Nordsieck 2007: China (Guangxi);
pallidissima H. Nordsieck 2010: p. albescens H. Nordsieck 2012, p. ooharai Hunyadi & Szekeres 2016,  p. pallidissima: China (Guangxi);
pseudofuchsi (H. Nordsieck 2005): p. amabilis (H. Nordsieck 2005), p. nigrofasciata H. Nordsieck 2012, p. pseudofuchsi, p. remota H. Nordsieck 2012, p. varicolor H. Nordsieck 2012: China (Guangxi);
puella H. Nordsieck 2012: China (Guangxi);
rutila H. Nordsieck 2016: China (Guangxi);
schomburgi (Schmacker & Boettger 1890): China (Hainan);
takagii (Chang 2004): t. capreolus (H. Nordsieck 2005), t. gigas H. Nordsieck 2007, t. takagii: China (Guangxi);
umbra (Chang 2004): u. rex H. Nordsieck 2007, u. umbra: China (Guangxi);
unicolor H. Nordsieck 2012: China (Guangxi);
yulinensis H. Nordsieck 2012: China (Guangxi).


Grego, J., Luong, H. V., Pham, S. V. & Szekeres, M. (2014): Vietnamese Clausiliidae (Gastropoda: Pulmonata): new taxa and novel distribution data. – Journal of Conchology, 41: 749–757.

Páll-Gergely, B. & Szekeres, M. (2017): New and little-known Clausiliidae (Gastropoda: Pulmonata) from Laos and southern Vietnam. – Journal of Conchology, 42: 507-521.

Grego, J., Hunyadi, A. & Szekeres, M. (2021): New and little-known Clausiliidae of continental South East Asia (Gastropoda: Pulmonata). – Journal of Conchology, 44: 21-29.

 Nordsieck, H. (2007): Worldwide Door Snails (Clausiliidae), recent and fossil. – 214 pp., 20 pls. Hackenheim (ConchBooks).

Uit de Weerd, D. R. & Gittenberger, E. (2013): Phylogeny of the land snail family Clausiliidae (Gastropoda: Pulmonata). – Molecular Phylogenetics and Evolution, 67: 201-216.

Check-list of the Neniinae

In the present chapter a check-list of the clausiliid subfamily Neniinae is given, together with a revised system and remarks on distribution and phylogeny. The range of the subfamily comprises a part of the West Indies (Hispaniola, Puerto Rico) and northern and central Andean South America.
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 (Colombia, 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 Colombia material for anatomical examinations is not available. Because the borderline between the two tribes, which have been erected, Neniini and Peruiniini, must be expected within the northern countries, the 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.
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. Zilch (1953, 1959) had figured those of two Peruvian species, Hylton Scott (1954) those of the only species from Argentina; later on, 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.


The subfamily Neniinae has been subdivided into two tribes, Neniini and Peruiniini (Nordsieck 2005). 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 narrow tube or absent. The other character, in which the tribes were said to differ, the presence or absence of a penial sheath, is not so suitable, because the Zilchiella group of the Peruiniini has a penial sheath similar to that of the Neniini.
As the examination of the genitalia has shown, not only the Caribbean genera Nenia and Nenisca, but also „Nenia“ bequaerti from Sierra de Perija, Colombia / Venezuela frontier region, belongs to the tribe Neniini (Grego & Szekeres 2008, Nordsieck 2010: 49-50). This species has 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.
In a DNA analysis carried out by Uit de Weerd & Gittenberger (2013) the Neniini (2 species from the Caribbean) and Peruiniini (9 species from Peru) come out as different clades, distant from one another, the Neniini basal to all other Clausiliidae and the Peruiniini as sister group of Phaedusinae + Garnieriinae. Thus, the separation as tribes is confirmed. The conclusion, however, that the tribes should be regarded as subfamilies (e. g., Molluscabase) is premature, as long as it is not clear, how these subfamilies are defined and which genus belongs to which subfamily. Besides, there are beyond the common apostrophy similarities in the shell (subcolumellaris not emerging) and genitalia (penial sheath also in the Zilchiella group), which speak against an independent origin of the two groups.
The tribe Peruiniini, in which most 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 inserting on pedunculus (in Columbinia columbiana on free oviduct) or absent.
Only with plesiomorphic characters (for character evaluation see Nordsieck 2005: 199-200), therefore the grouping may be not natural.
Note: 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 genus 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 inserting on pedunculus; in contrast to the other groups penial sheath present.

Temesa group:
Shell entire; posterior lower palatal plica absent; diverticulum inserting on pedunculus, basally dilated (Parabalea), or absent (Temesa).
Note: Both genera were formerly united in the genus Temesa. Because of the lack of diverticulum in Temesa it is questionable if that genus is in fact related to Parabalea.

Incaglaia group:
Shell entire; posterior lower palatal plica absent; diverticulum inserting on pedunculus or absent.
Incaglaia, n. gen. (argentina).
Note: For I. 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 absent. Its range is remote from that of the Incaglaia species.

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

Peruinia group:
Shell decollated; posterior lower palatal plica absent; diverticulum inserting on pedunculus or absent. 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 absent; 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 clausilial apparatus.

Steeriana group:
Shell entire or decollated; posterior lower palatal plica present to absent; diverticulum inserting on free oviduct.
With entire shell: NeniatractaHemicenaLeuconenia; with decollated shell = Steeriana group sensu stricto: CylindroneniaAndiniaAndiniastraSteerianaNeniella.

Distribution and phylogeny

The Caribbean genera of the Neniini occur on the islands Hispaniola (Nenisca) and Puerto Rico (Nenia). The genera from South America, which are provisionally classified with the Neniini (Neniops, Paranenia, Gonionenia), are distributed in Colombia; some species are said to come from Venezuela and Ecuador.
The genera of the Peruiniini are distributed in South America. Columbinia, Incania and Cyclonenia species have been found across the whole range of the tribe from Colombia to Bolivia, especially Incania and Cyclonenia (that genus only in N. Andean Peru) also in higher altitudes; Pfeifferiella occurs in higher N. W. Andean Peru. The Zilchiella group is restricted to a small range in N. Andean Peru. Temesa occurs in higher N. W. Andean Peru, Parabalea in higher central and S. Andean Peru. Incaglaia is distributed in higher central and S. Andean Peru, the different I. argentina in N. Argentina in a lower altitude. The Ehrmanniella group is known from higher N. and central Andean Peru. The Peruinia group occurs in central Andean Peru, at the slope towards the Amazon Basin, Symptychiella at this slope in the northeast. The Steeriana group in the strict sense is restricted to higher N. Andean Peru; the other genera provisionally attached to the Steeriana group occur in N. and central Andean Peru, Neniatracta at the slope towards the Amazon Basin, Hemicena and Leuconenia in higher altitudes.
The DNA analysis of Uit de Weerd & Gittenberger (2013) shows that Neniini and Peruiniini are  independent phylogenetic units; therefore, it is doubted by the authors that the Neniinae are monophyletic. According to their chronogram (: fig. 3) the Neniini diverged from the other Clausiliidae already in the Cretaceous.
In the DNA analysis the included species of the Peruiniini form in all trees a well-supported clade, i. e. the  tribe is probably monophyletic. Within the tribe Zilchiella and Peruinia are basal to other clades, Zilchiella more basal. In their chronogram both are separated from the remaining groups already since Eocene. The other species form only two well-supported subclades. The Steeriana group sensu stricto (Cylindronenia (Cylindroneniella), Andinia, Steeriana) is a well-supported clade in nearly all trees. Temesa and the neighbouring Cyclonenia species come out within a well-supported clade in the 28S and the consensus tree. The included Incania and Incaglaia species form also a clade, which is, however, well-supported only in the consensus tree.
Thus from the DNA analysis the following is concluded: The Zilchiella and the Peruinia groups are basal within the phylogeny of the Peruiniini. The basal position of the Zilchiella group is also proved by a plesiomorphic genital character (with penial sheath like the Neniini). The Steeriana group sensu stricto is obviously a monophyletic group, characterized by the apomorphic characters decollate shell and diverticulum inserting on free oviduct. Temesa and Cyclonenia are closely related; this increases the doubt if Temesa and Parabalea belong in fact to one group. A close relationship of Incania and Incaglaia, which could be concluded from the consensus tree, is improbable, because  they are very different in shell and genital morphology.
A DNA analysis carried out by Morín Lagos (2018) worked into the analysis of Uit de Weerd & Gittenberger includes species of Andiniastra (but one of them belonging to Cylindronenia (C.)) and Symptychiella. The result is that Andiniastra, Cylindronenia and the once more included Cylindronenia (Cylindroneniella) are within the clade of the Steeriana group sensu stricto, while Symptychiella clusters with Zilchiella. The latter shows only that Symptychiella is like Zilchiella and Peruinia a basal phylogenetic group.
The species named Andiniastra aff. violascens by Morín Lagos, which comes from Cajamarca departamento near Cutervo, is very similar to Cylindrophaedusa (C.) canescens, which occurs in the same region. The figures (figs. 19, 54-55) show that in contrast to Andiniastra violascens it has a weak dorsal keel, a spur-like anterior lower palatal plica and a superior lamella separated from the spiral lamella just in the same manner as in C. canescens. Its shell is, however, somewhat larger and less decollated. It is interesting that according to the author it is less related to Andiniastra than the genus taxa Steeriana, Andinia and Cylindronenia (Cylindroneniella) to one another and has diverged earlier from it than these taxa from one another. This speaks for a ranking of Cylindroneniella as genus of its own.
Further DNA analyses are necessary to get deeper insights in the phylogeny of the Peruiniini.


The following list includes all known species and subspecies, with indication of 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): Colombia.

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

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

Peruiniini H. Nordsieck 2005:

Columbinia Polinski 1924:

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

n. gen.:
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 1880) (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 (Junin?);
warszewiczi (Polinski 1924): Peru (Cuzco?).

Cyclonenia H. Nordsieck 1999:
albosuturalis Neubert & Nordsieck 2005: Peru (Cajamarca);
alpina H. Nordsieck 1999: Colombia;
boliviana (O. Boettger 1893): Bolivia;
cyclostoma (L. Pfeiffer 1850): Colombia, 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: Colombia.

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. Adams & 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 1880): Peru (Cuzco?);
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 1880): 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 1880): 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:

funcki (L. Pfeiffer 1848): Venezuela.

gen. 1:
pusilla (Polinski 1922): Peru (?).

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

gen. 3:
dichroa (Haas 1929): Bolivia;
magnifica (Sykes 1901): Bolivia.

Figs. 1.-12. Selected species of Neniinae. 

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, Colombia, Santa Fé de Bogota, NMBE, H = 35.9.
3. Columbinia (C.) columbiana, Colombia, 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 f. 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 of 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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