Revisory remarks on the species of Siciliaria Vest from N. W. Sicily
Hartmut Nordsieck (VI.2013)
A. Schmidt (1868: 40-43) was the first who proposed a systematic arrangement of the seven Clausilia species from N. W. Sicily (= Siciliaria (S.)) known so far. He classified the species with two groups („Formenkreise“) which mainly differ by the formation of the clausilium plate. Other characters which he regarded as important were sculpture, development of lamellae (inserta, inferior lamella) and palatal plicae. His system was the following: Formenkreis of septemplicata: Clausilia grohmanniana Rossmässler, C. septemplicata Philippi, C. calcarae Philippi, C. confinata = scarificata L. Pfeiffer, C. tiberii A. Schmidt; Formenkreis of crassicostata: C. crassicostata L. Pfeiffer, C. nobilis L. Pfeiffer.
O. Boettger (1877: 33-34) named the groups of Schmidt (F. of septemplicata = Siciliaria Vest sensu stricto, F. of crassicostata = Trinacria O. Boettger). Later on, he (1879: 85-91) gave descriptions of several species taxa and listed eight species (C. leucophryne L. Pfeiffer added to those of Schmidt). His distribution data were in part based on wrong localities on the labels from Benoit and others.
Benoit (1882: 101-120 (sep.)) who had distributed most of the material in the collections of his contemporaries listed 13 species, several of them regarded as varieties or synonyms by Boettger. Like those of Boettger, his distribution data were partially wrong.
Westerlund (1892: 47-48) published three of the numerous taxa names of (Allery di) Monterosato and gave a determination key for the species.
A. J. Wagner (1913, 1925) figured the genital organs of three Siciliaria (S.) species (S. septemplicata: 1913: pl. 572, fig. 14; S. grohmanniana: 1925: pl. 1, fig. 8, S. calcarae: pl. 3, fig. 25). He (1924: 123-125) gave a list of the species which is nearly identical with that of Boettger.
Brandt, after his collection trip in Sicily in 1949, prepared a manuscript with the description of 24 new taxa based on „species“ of Monterosato or collected by himself. This manuscript remained unpublished; later on, only five of the species taxa were regarded by Brandt (1961: 6-9) as different enough to be described as new (among them two species: S. ferrox, with figure of genitalia fig. 1, S. alcamoensis). Besides, he (: 8) united S. leucophryne and S. tiberii (and S. eminens which later on was recognized as species) to one species under the name S. leucophryne, with which he also affiliated the newly described S. riberothi as subspecies. Many localities given by him cannot be identified or are obviously wrong.
Alzona (1971: 90-92) listed the same species as Boettger. Brandt's new taxa were not considered, although his paper was cited.
Nordsieck (1979: 259-260) listed also the same species as Boettger and (: 269) united his species groups (Siciliaria s. str., Trinacria (preoccupied) = Sicania Tomlin) because of the intermediate position of S. calcarae. Later on, he (1984: 202) described the new species S. spezialensis based on material collected by K. L. Pfeiffer.
Manganelli et al. (1995: 25, 47-48), in the checklist of Italian pulmonate gastropods, enumerated 10 species taxa, proposed as such by Nordsieck (in litteris, S. ferrox and S. spezialensis added to those of Nordsieck 1979). At the same time, Giusti et al. (1995: 327-330) evaded a distinction of species within the group which they gave the oldest name S. septemplicata. Therefore, they determined S. calcarae from Quaternary deposits of Malta as S. cf. septemplicata.
Nordsieck (2002: 30, 34) recognized C. leucophryna and C. eminens as different species. He (29, 34) listed 12 species of Charpentieria (Siciliaria) (= Siciliaria (S.)) and classified them with three species groups, based on some shell characters regarded as informative: 1 nobilis-calcarae group, 1a nobilis subgroup (C. nobilis, C. spezialensis, C. crassicostata, C. eminens), 1b calcarae subgroup (C. calcarae, C. tiberii, C. leucophryna), 2 grohmanniana group (C. grohmanniana, C. septemplicata, C. scarificata). Because of their ambiguous character combinations, C. ferrox and C. riberothi were not included in one of these groups. Several forms assumed to have originated by interspecies hybridization (: 34) were added.
Beckmann (2004: 188, maps 1 and 2) published detailed informations on the distribution of the species and described (: 188-190) two new subspecies, both assumed to be hybrid forms.
Reitano et al. (2007: 326-328) gave further informations on the distribution of S. ferrox.
The material on which the following system is based comprises about 230 samples of Siciliaria (S.) species from the collection of the Forschungsinstitut Senckenberg (abbreviation: SMF), containing the types of several species taxa, and 80 samples of my own collection, deposited in the Staatliches Museum für Naturkunde Stuttgart (abbreviation: SMNS) and the Forschungsinstitut Senckenberg. In the Senckenberg collection material collected by Kobelt, K. L. Pfeiffer, Brandt and Beckmann is kept, in my own collection an important part of the Brandt collection (SMNS) and material collected by C. and J. Hemmen in 2009 (SMF). 80 samples of all taxa have been subjected to a more thorough morphological examination.
In collections from the nineteenth century the localities given on the labels have often turned out to be wrong. In this article only locality data given by collectors, who in my opinion are reliable, are used. In any case, I had to rely on those data, because until now I had not the opportunity to collect in this region.
Siciliaria Vest and Charpentieria Stabile are separated as genera; for the arguments see Nordsieck (2013) and the website art. Delimini Italy. The DNA analyses of Uit de Weerd & Gittenberger (2013: tree fig. 3) and Hausdorf (unpubl.: tree based on 16S mtDNA analysis) confirm this separation; in both trees Charpentieria appears as sister group of a clade including Siciliaria and Delima Hartmann. In Hausdorf's tree Siciliaria (S.) species come out in two separate subclades, one with S. septemplicata and S. leucophryna, the other with three species of the nobilis subgroup.
The examination of the genital morphology of several Siciliaria species (listed in Nordsieck 2002: 27) had the result that the genitalia of all species are similar. Compared with the species of the other Siciliaria subgenera (except S. (Mauritanica) O. Boettger), the three examined species of Siciliaria (S.) (S. grohmanniana, S. calcarae, S. nobilis) have a relatively short diverticulum of bursa copulatrix in common. A character which with regard to the results of Hausdorf's analysis gains in importance is the development of the allospermiduct (a similar case as in Cochlodina, see website art. Cochlodina diversity). In S. calcarae and S. nobilis (also in the examined S. (Mauritanica) species) the allospermiduct is not widened along the spermoviduct, in S. grohmanniana , like in all other examined Siciliaria species, it is widened along the spermoviduct.
1. Diagnoses and distribution
In the following the diagnoses of the species of Siciliaria (S.) and their distribution, as far as known from the examined material, are given. In notes, geographic forms which could have subspecific rank are characterized. Among them, there are several forms which have probably originated by interspecies hybridization (see chapter 2.).
The following shell characters have turned out to be informative for species diagnoses (see figs. 1-7):
1. Decollation: Shell entire / decollated.
2. Sculpture: Whorls rib-striated, with sutural papillae / weakly ribbed, papillae still recognizable / strongly ribbed.
3. Cervix of body whorl: Only with basal keel, without dorsal keel / with more or less pronounced dorsal keel.
4. Inferior lamella: Lamella more or less protruding into the lumen of the body whorl, described as: lamella high / moderately high / low.
5. Anterior upper palatal plica: Only one plica present / two plicae present, upper and lower one / plica missing.
6. Clausilium plate: Palatal edge of plate distally receding, thus distal part narrowed, forming a gutter (figs. 1-2) / not receding, palatal and columellar edges of plate nearly parallel (figs. 3-7). Palatal edge distally much bent up (figs. 3-4) / bent up (figs. 1-2) / somewhat or not bent up (figs. 5-7).
The distribution of the subgenus is restricted to N. W. Sicily: Egadi islands, Sicilian mainland from Trapani in the west to the western boundary of Madonie in the east, in the south delimited by the imaginary line Trapani–Calatafimi–San Cipirello–Marineo–Monte San Calogero.
In a larger part of N. W. Sicily two species pairs with identical clausilium plate, but different appearance (shell entire, nearly smooth and papillate / shell decollated, ribbed) occur (figs. 1-4). The smooth papillate species is adapted to more shady habitats, while the ribbed one lives in more open rock habitats. Decollation is possibly an adaptation to a life under stones. One of these species pairs, S. septemplicata / S. grohmanniana (figs. 1-2), is distributed in the east, the other one, S. calcarae / S. tiberii (figs. 3-4), in a larger part of the region, S. calcarae in the whole region, S. tiberii only in the central and eastern part. The range restriction of the latter is probably caused by the occurrence of other rock-dwelling species, those of the nobilis subgroup, in the west.
On the San Vito peninsula, there is a species pair of the nobilis subgroup, which lives in open rock habitats, S. nobilis / S. spezialensis. Both are adapted to this habitat by a well-developed white surface layer of the shell (fig. 5), looking like Albinaria species, both are decollated. S. spezialensis exhibits a special closing apparatus (clausilium plate fit into a frame of palatal plicae, anterior palatal plicae missing), which may better protect it from desiccation. On one mountain of this peninsula, Monte Cofano, a species with percostate shell,
C. crassicostata (fig. 6), occurs. This sculpture, which is found also in certain Albinaria species and other rock-dwelling clausiliids of meridional Europe, may be another adaptation to a life in open rock habitats.
The occurrences of Siciliaria (S.) species on the Egadi islands are of special interest. On Marettimo, only S. scarificata occurs. This endemic species is closely related to the other species of the grohmanniana group distributed in the east, not to the neighbouring ones of the nobilis-calcarae group. On Favignana, originally only S. calcarae occurred. C. crassicostata, which has been collected by several authors on the island (e. g., by Brandt on garden walls, fig. 6) has obviously been introduced (by man?) from Monte Cofano nearby.
Siciliaria Vest 1867
Nomenclatural note: Clausilia decollata and C. deshayesii Potiez & Michaud 1838 are unidentified species. The first may be S. grohmanniana, the second S. septemplicata or S. calcarae. Three „species“ of Benoit (1882) are also unidentified: C. trinacliana (misprint for trinacriana) (= S. calcarae?), C. rubra (= S. septemplicata?) and C. subdiaphana (= S. grohmanniana?). The variety names published by Monterosato, in fact infrasubspecific, are not considered in this article.
S. grohmanniana group
S. grohmanniana (Rossmässler 1836)
Diagnosis: Shell mostly decollated; whorls ribbed; dorsal keel indistinct or missing; inferior lamella very high; two anterior upper palatal plicae present, upper one mostly separated from upper palatal plica; palatal edge of clausilium plate distally receding, plate gutter-like narrowed, palatal edge against distal end bent up and more or less pointed.
Distribution: Monte Pellegrino, Monte Gallo.
Synonyms: C. grohmanniana var. minor A. Schmidt 1868 [non Rossmässler], C. grohmanniana var. panormitana Paulucci 1878.
Priola form, Priola and S. Lorenzo near Palermo: Shell not decollated; whorls more weakly ribbed.
S. septemplicata (Philippi 1836)
Diagnosis: Shell mostly not decollated; whorls rib-striated, with sutural papillae; dorsal keel indistinct or missing; inferior lamella very high; two anterior upper palatal plicae present, upper one mostly separated from upper palatal plica; palatal edge of clausilium plate distally receding, plate gutter-like narrowed, palatal edge against distal end bent up and more or less pointed.
Distribution: From Cinisi in the west and Monte Gallo in the north to Piana degli Albanesi in the south and Monte Catalfano in the east.
Synonyms: C. sericina Rossmässler 1836, C. septemplicata var. prasina A. Schmidt 1868. The name C. prasina is a copying-mistake of C. proxima, caused by illegible labels of Benoit.
Parco form, Parco near Palermo: Inferior lamella in part lower; lower anterior upper palatal plica missing; palatal edge of clausilium plate less or not bent up.
s. alcamoensis Brandt 1961, „Alcamo“: Cervix more strongly sculptured; dorsal keel more pronounced; inferior lamella in part lower; lower anterior upper palatal plica more or less reduced; clausilium plate distally less narrowed, palatal edge less upbent.
s. hemmeni (Beckmann 2004), Piano Margi near Cinisi: Same characters as the foregoing, slight differences in the development of subcolumellar lamella and position of lunellar. S. s. hemmeni is therefore considered a synonym of S. s. alcamoensis.
S. scarificata (L. Pfeiffer 1856)
Diagnosis: Shell mostly decollated; whorls ribbed, sutural papillae recognizable; dorsal keel indistinct ore missing; inferior lamella high; anterior upper palatal plica present, separated from or connected with upper palatal plica; palatal edge of clausilium plate distally receding, palatal edge distally somewhat or not bent up.
Distribution: Island of Marettimo.
Synonyms: The names C. scarificata and C. sacrificata are copying-mistakes of C. confinata, caused by illegible labels of Benoit.
Species with intermediate position
S. leucophryna (L. Pfeiffer 1862)
Diagnosis: Shell decollated; whorls ribbed; dorsal keel mostly indistinct or missing; inferior lamella moderately high; anterior upper palatal plica present, separated from or connected with upper palatal plica; palatal edge of clausilium plate distally slightly or not receding, palatal edge distally somewhat or not bent up.
Distribution: Isola delle Femmine and Grotta Conza to Monte Gallo.
Synonyms: C. leucophryne var. laudabilis O. Boettger 1879, C. confinata var. merens Westerlund 1892.
Sferracavallo form, Sferracavallo: Inferior lamella lower; palatal edge of clausilium plate more strongy bent up.
S. nobilis-calcarae group, calcarae subgroup
S. calcarae (Philippi 1844)
Diagnosis: Shell not decollated; whorls rib-striated, with sutural papillae; dorsal keel indistinct or missing; inferior lamella moderately high or low; anterior upper palatal plica present, mostly separated from upper palatal plica, rarely lower anterior upper palatal plica present; palatal edge of clausilium plate distally not receding, palatal edge distally more or less strongly bent up.
Distribution: Island of Favignana, whole mainland of N. W. Sicily from Erice in the west to Montagna Grande in the east; in the south to Calatafimi and to south of Piana degli Albanesi.
Synonyms: C. adelinae Küster 1860, C. brugnoneana Pini 1884, C. calcarae var. nodosa Westerlund 1892.
Sagana form, Sagana near Montelepre: Inferior lamella higher; lower anterior upper palatal plica more or less developed; palatal edge of clausilium plate less strongly bent up.
c. belliemii Brandt 1961, Monte Belliemi near Partinico: Whorls ribbed.
S. tiberii (A. Schmidt 1868)
Diagnosis: Shell not decollated or decollated; whorls ribbed; dorsal keel of different prominence; inferior lamella moderately high or low; anterior upper palatal plica present, mostly separated from upper palatal plica, rarely lower anterior upper palatal plica present; palatal edge of clausilium plate distally not receding, palatal edge distally more or less strongly bent up.
Distribution: From north of Cinisi in the north to San Cipirello in the south, Monte San Calogero.
Synonyms: C. tiberiana Paulucci 1878.
Northern form from Monte Palmeto and Monte della Fiera: Shell decollated; dorsal keel strong.
Southern and eastern form from environs of San Cipirello and Monte San Calogero = type form: Shell not or only in part decollated; dorsal keel less strong.
Form from Monte della Scala: Shell not decollated; dorsal keel indistinct.
Armetta form, Grotta Armetta near Carini: Shell decollated; more densely ribbed; dorsal keel weak; inferior lamella higher.
t. scalettensis (Beckmann 2004), Port la Scaletti near Cinisi: Whorls rib-striated; inferior lamella higher; palatal edge of clausilium plate somewhat or not bent up.
S. nobilis-calcarae group, nobilis subgroup
S. nobilis (L. Pfeiffer 1848)
Diagnosis: Shell decollated; with well-developed white surface layer; whorls mostly weakly rib-striated, with sutural papillae; dorsal keel missing; inferior lamella moderately high or high; anterior upper palatal plica present, mostly separated from upper palatal plica, or missing, basalis tending to be shortened, posterior lower palatal plica more or less reduced; palatal edge of clausilium plate distally not receding, palatal edge distally somewhat or not bent up.
Distribution: San Vito peninsula from Monte Cofano in the west to Monte Monaco und Monte Acci in the east.
Synonyms: C. cantrainei Deshayes 1851, C. sicula Rossmässler 1856 (validated by Paulucci 1878).
Type form from „Ridicofano“ = San Vito: Anterior upper palatal plica mostly present.
Form from remaining species range: Anterior upper palatal plica mostly missing.
n. episoma Brandt 1961, coast near S. Vito: Shell smaller; whorls more distinctly rib-striated; anterior upper palatal plica missing.
S. spezialensis H. Nordsieck 1984
Diagnosis: Shell decollated; with well-developed white surface layer; whorls ribbed or weakly rib-striated, with sutural papillae; dorsal keel missing; inferior lamella mostly high; anterior upper palatal plica missing, basalis missing, posterior lower palatal plica connected with lunella; clausilium plate fit in the frame of lunella and posterior lower palatal plica; palatal edge distally not bent up.
Distribution: San Vito peninsula from Macari and Castelluzzo to Monte Acci and Monte Speziale.
Type form is ribbed.
Form from remaining species range: Whorls weakly rib-striated.
S. eminens (A. Schmidt 1868)
Diagnosis: Shell decollated; whorls ribbed; dorsal keel indistinct or missing; inferior lamella mostly high; anterior upper palatal plica present, mostly separated from upper palatal plica, basalis tending to be shortened; palatal edge of clausilium plate distally not receding, palatal edge distally somewhat or not bent up.
Distribution: From Custonaci in the west to Castellammare in the east, environs of Calatafimi, M. Bonifato near Alcamo.
S. crassicostata (L. Pfeiffer 1856)
Diagnosis: Shell decollated; whorls ribbed, in part percostate; dorsal keel distinct or indistinct; inferior lamella moderately high or high; anterior upper palatal plica present, separated from upper palatal plica, in part missing, basalis tending to be shortened; palatal edge of clausilium plate distally not receding, palatal edge distally somewhat or not bent up.
Distribution:: Monte Cofano, (probably introduced on) island of Favignana.
Type form is percostate.
Form from near Monte Cofano and part of Monte Cofano: Whorls not percostate = normally ribbed.
S. nobilis-calcarae group, no subgroup
S. ferrox Brandt 1961
Diagnosis: Shell mostly not decollated; whorls ribbed, sutural papillae recognizable; dorsal keel distinct or indistinct; inferior lamella moderately high or high; two anterior upper palatal plicae present, upper one mostly separated from upper palatal plica; palatal edge of clausilium plate distally less or not receding, palatal edge distally not bent up.
Distribution: From Altavilla Milicia to Termini Imerese.
Synonyms: C. confinata var. commeata Westerlund 1892 could be an older synonym, but the description is too poor (lower anterior upper palatal plica not mentioned) and types are not available (Reitano et al. 2007: 328).
S. riberothi Brandt 1961
Diagnosis: Shell decollated; whorls ribbed; dorsal keel prominent; inferior lamella high; anterior upper palatal plica present, mostly separated from upper palatal plica; palatal edge of clausilium plate distally not receding, palatal edge distally not bent up.
Distribution: Unknown (Trapani province?) (see chapter 3.).
2. Interspecies hybridization
The forms of Siciliaria (S.) assumed to be hybrid ones, which are enumerated in the following, are forms of well-defined species restricted to certain localities or regions differing from the nominotypical forms by showing characters of another species. This other species is clearly different from that species and is distributed in the same region. The assumed hybrid forms occur in the neighbourhood of or are even surrounded by populations of the nominotypical form of the species to which they belong. In all probability, they are the products of a limited gene flow from the other species restricted to a limited locality or region. This may be a similar case of interspecies hybridization as, e. g., in the genus Delima in central Dalmatia (Nordsieck 2007: chapter VI, 99-101).
The assumed hybrid forms are the following:
1. S. grohmanniana, Priola form (fig. 8), with characters of S. septemplicata, occurring in the environs of Palermo, where both species are distributed.
2. S. septemplicata, Parco form, with characters of S. calcarae, from a locality near Palermo in a region, where both species have been traced in localities nearby.
3. S. septemplicata alcamoensis (fig. 9) = S. s. hemmeni, with characters of S. tiberii, the latter occurring near Cinisi in a region, where both species are distributed. The locality Alcamo for S. s. alcamoensis is probably wrong, also because neither S. septemplicata nor S. tiberii have been found in that region. The shell from Alcamo figured under the name S. s. alcamoensis in the Internet forum Natura Mediterraneo (XII.2009) belongs to S. calcarae.
4. S. calcarae, Sagana form, with characters of S. septemplicata, from a locality near Montelepre in a region, where both species have been traced in localities nearby.
5. S. calcarae belliemii from Monte Belliemi, with the ribbing of S. tiberii, which occurs in the Monte della Fiera nearby.
6. S. tiberii from Monte della Scala (fig. 10), with characters of S. calcarae.
7. S. tiberii scalettensis, with characters of S. septemplicata, from a locality near Cinisi where both species are distributed.
8. S. leucophryna, Sferracavallo form, with characters of S. calcarae, from Sferracavallo where both species have been found.
9. S. crassicostata with normally ribbed shell from near Monte Cofano and part of Monte Cofano, ribbing like S. eminens.
3. The riberothi problem
The taxon which has been described by Brandt (1961: 8) as S. leucophryne riberothi is a special problem within the systematics of Siciliaria (S.). The locality of the type form „valley north of Ribera“ given by Monterosato, who provided it as Clausilia riberothi, is probably wrong. The species cannot be found in the environs of Ribera (Agrigento province) (Beckmann 2004: 188); besides, this locality is outside the range of Siciliaria (S.). Another form (fig. 7), also coming from Monterosato and named C. remota (this name, however, has been used by him also for the northern form of S. tiberii, Brandt 1961: 8), is said to come from the mountains „della Terrazza“ (or „Torrazza“) in the Trapani province (two samples, already mentioned by Beckmann: 188). This locality, within the range of Siciliaria (S.), could not be traced until now. In contrast to all other Siciliaria (S.) species, S. riberothi does not appear in the Internet forum Natura Mediterraneo.
S. riberothi differs from S. leucophryna by the prominent dorsal keel, which is like that of the northern form of S. tiberii, and from S. leucophryna and S. tiberii by the clausilium plate (fig. 7), which is like that of the nobilis subgroup (and S. ferrox). Therefore, it is provisionally regarded as an independent species.
It is a task for malacologists in Sicily to rediscover this questionable species with unknown origin.
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