In Table 3, the most important character states of the shells and the genital organs of the Otalini subgroup according to Razkin et al. (2015: 108, Fig. 2) including the genera Cantareus, Cornu, and Rossmaessleria are shown. Within the Otalini, Cantareus, Cornu, and Erctella share the synapomorphy of a globular to slightly conical shell, other genera in the tribe tend to have more flattened shells (character 1). In all other shell traits, there is no apomorphy that discriminates between Cantareus, Cornu, and Erctella on generic level. On the level of the genital organs, the three genera share the synapomophy of the simple pore connecting epiphallus and penial chamber (character 8), while Rossmaessleria shows the plesiomophic state with two functional penial papillae. In Cornu, the flagellum is much longer than in the other genera. The phylogenetic value of this character state is not clear within the Otalini, within the Helicini, it is considered a plesiomorphy (Neubert 2014). Other character states like ratios in the bursa copulatrix complex (character 10). A massive atrial stimulator can be found in Cantareus and Cornu, while in Erctella, it is relatively small (character 11). Large and massive stimulators are found in many taxa of Helicidae, so a reduction of this system is here interpreted as an apomorphic character state.

Remarks. The change of the status of Cantareus from a monotypic to a polytypic genus causes some nomenclatorial problems. The type species of the genus is Helix naticoides Draparnaud, 1801 from France, which so far has been considered a synonym of Helix aperta Born, 1778, with the specimen preserved in the Born collection in the NHMW as the name bearing syntype of aperta (Fig. 2).The origin of Born’s specimen is unknown, and there are almost no shell morphological differences to the Algerian lineage, which proved to represent a separate species (Fig. 1). The correct origin of Born’s specimen could probably be clarified genetically by applying NGS methods using shell fragments of the syntype specimen, but this is beyond the scope of this paper.

Figure 2. 

Cantareus apertus. Syntype of Helix aperta NHMW-MO 14005, shell diameter 28.75 mm. Shell in A frontal B dorsal C lateral and D apical views E labels of the syntype lot. Photographs NHMW, × 1.5.

Anticipating a north African origin of the syntype NHMW-MO 14005 by fixing its type locality in Algeria ends up in a chaotic rearrangement of species names in the group. For Europe, the name naticoides would be reactivated with its last use as an accepted species in 1850 (!). The north African species would then be named apertus contradicting 170 years of permanent use. By fixing the use of the name Helix koraegaelia Bourguignat in Locard, 1882, to the north African lineage, this problem is resolved, and the stability or universality of names used in zoology is guaranteed.

Cantareus apertus is well known for its protective behaviour, which gave the genus its name “Cantareus: the singer”. Once disturbed (Fig. 23), the species is able to press the air in its lung cavity through the pneumostome producing a series of tweeking sounds (https://youtu.be/CWOhZWLkd4o).

Syntype aperta: NHMW-MO 14005.

Italy: Foggia, Ordona, 41.313889N, 15.622222E, 12.10.2018, leg. G. Martucci (ex coll. Sparacio 5031/9), coll. Liberto (Fig. 3); Calabria, Amantea, Marincola, 39.112778N, 16.079722E, 7.10.2018, leg. W. Renda, NMBE 560941/2 (preserved), ex coll. Liberto (Fig. 4).

Figure 3. 

Cantareus apertus. Italy, Foggia, Ordona A shell in frontal (left) and B lateral (right) view; genital anatomy: situs (C) and a section showing the male genital tract and the atrium (D). Photographs F. Liberto.

Figure 4. 

Cantareus apertus. Italy, Calabria, Amantea, Marincola, NMBE 560941 A shell in frontal and B lateral views C genital anatomy, situs D, E details showing the male genital tract and the atrium. Photographs E. Neubert, shell × 1.5.

Shell thick, medium sized if compared to other helicid species, with a depressed spire and a large last whorl occupying more than two thirds of the complete height of the shell; protoconch small, consisting of 1.5 smooth whorls; teleoconch consisting of approximately 4 whorls, separated by a deep, sometimes crenulated suture; basic colour of teleoconch greenish-brownish, often with longitudinal yellow streaks and a few scattered zig-zag markings; surface of teleoconch smooth, but also often covered by low longitudinal riblets; aperture almost perfectly rounded, enormously large, old specimens with an inconspicuous whitish lip; umbilicus always completely closed.

Genital organs: penis short, club-shaped, epiphallus short, of the same length as penis, mrp attaching in the distal third of epiphallus or even closer to penis; flagellum twice the length of the epiphallus; atrial and penial lumen with numerous crests, penial chamber lumen is wrinkled, pp2 a short broad papilla with a central perforation structured by thick annuli to almost completely reduced forming a septum; pp1 a blind papilla, in a central position inside the penial chamber, elongate, sometimes with a broadened tip; epiphallial pore in a lateral position; distal epiphallial lumen with six broad pilasters, the proximal lumen with elongated ridges.

Vagina short, stem of pedunculus thickened and short, diverticulum slightly longer than the vesicle stem + vesicle, longer than the flagellum; glandulae mucosae longer than the dart sac, with a thickened basal part and two subsequent ramifications, tubules thin and weak, less than 10 tubules per stem; atrium with a medium sized stimulator flap.

Syntype NHMW: H = 28.25 mm; D = 28.75 mm; PH = 22.3 mm; PD = 19.2 mm.

South-eastern France including Corsica, Italy, Sicily, south-eastern Adriatic coast, Albania, western Greece; scattered found introduced on some Aegean Islands, and in Turkey, Muğla, Gökçebel (Örstan et al. 2005: 7).

The anatomy of the genital organs of C. apertus has been investigated by several authors, for example Hesse (1919), Germain (1930) and Giusti et al. (1995). Schileyko (2006: 1801, fi.g. 2308) presented also details of the penial lumina. In his picture of the genital organs of an animal collected in the surroundings of Pisa, Italy, he misinterpreted the morphology of the epiphallial papilla (pp1) suggesting that it was a functional papilla as in many other helicid genera (the illustrated shell comes from Arles, France, and thus does not belong to the dissected specimen). Secondly, in his specimen, the penial papilla (pp2) was completely reduced, so only the perpendicular wall forming the basis of pp2 was left. This led to the misapprehension that in the genus Cantareus, this papilla is reduced, and only a “septum” is left in the place of the papilla.

Mazzulopsis: lectotype ANSP 63133, paralectotype ANSP 459220. subaperta: 3 syntypes, NHMW 7861, NHMW 7862, NHMW 7863; paratypes SMF 75256/8, coll. Nägele ex Ancey, the original label of Ancey contains the additional information “Dra-el-Mizan, 1893”.

Algeria, Kabylie: Tiguemounine (Ouacif), 1100 m alt. coll. Bouaziz; Ighil Bourmi (Ait Bouaddou), 950 m alt. NMBE 550458; ditto, le. F. Medjoub, NMBE 555649; Ait Houari (Assi Youcef), 1000 m alt. coll. Bouaziz; Tizi Guefres (Iferhounene), 1100 m alt coll. Bouaziz. The Senckenberg Research Institute houses > 30 shells of this species, all of them from “Kabylie” and/or “Djudjura”.

Shell medium sized to large, thin, globose with a broad to relatively acute conical spire; protoconch whitish, large, with a diameter of up to 6 mm and 2.5 smooth whorls; basic shell colour olive yellowish with up to five separate brown spiral bands; teleoconch covered by a dense granulation, sometimes accompanied by very fine, deep spirals; teleoconch usually covered by irregularly arranged riblets of even ribs, usually stronger around the umbilical area; periostracum thick, often preserved on the shell in small patches; in eroded shells, ribs and riblets whitish; aperture large, elongate oval, slightly thickened forming a lip callus, with a parietal callus in fully adult specimens; aperture whitish inside, with the spiral bands shining through the thin shell; peristome sharp; umbilicus closed, periomphalum covered by a thickened calcareous layer.

Figures 5–9. 

Cantareus subapertus (Ancey, 1893). 5–7 syntypes Helix subaperta NHMW, Djurdjura, Kabylie ex Ancey 5 NHMW 7861, D = 23.44 mm 6 NHMW 7862, D = 23.51 mm 7 NHMW 7863, D = 29.44 mm 8 Helix mazzulopsis lectotype ANSP 63133, Jurjura Mts. Shell in frontal (A) lateral (B) and dorsal (C) view (D, E) labels 9 “Helix aspersa“, original specimen of Iconographie (2) 3, pl. 69, fig. 359. Shell in frontal (A) lateral (B) ventral (C) and apical (D) views. Photographers 5–7 H. Wood, NHMW; photograph 8 E. Wildner, ANSP; photograph 9 E. Bochud, NMBE; all shells × 1.5.

Penis short, epiphallus reaching at least three times the length of penis, mrp attaching in the distal third of epiphallus; flagellum twice the length of the epiphallus; penial lumen smooth, pp2 a short broad papilla with a central perforation structured by thick annuli, pp1 a blind papilla, the epiphallial pore in a lateral position; distal epiphallial lumen with broad pilasters, the proximal lumen with elongated ridges.

Vagina short, stem of pedunculus thickened and short, diverticulum longer than the vesicle stem + vesicle, longer than the flagellum; glandulae mucosae longer than the dart sac, with a thickened basal part and two subsequent ramifications, tubules thin and weak, less than 10 tubules per stem; atrium dominated by a massive stimulator.

Syntypes figured (n = 4): H = 26 mm; D = 27.5 mm; PH = 17.7 mm; PD = 15.4 mm.

As far as known, this species is restricted to the Djudjura Mts., where it inhabits quite high altitudes. It also occurs in the northern promontory of this mountain ridge.

In the description of Helix subaperta, Ancey (1893) mentioned in a footnote that he already shared this species under the name H. mazzuliopsis with his correspondents. Pilsbry’s shells (1893) were purchased from a shell dealer (see label in Fig. 8E) bearing the name H. mazzulopsis, which he consequently used! Moreover, Pilsbry remarks: “I have been unable to find any description or mention of this form in the literature”; thus, H. mazzulopsis cannot be considered an emendation or an incorrect subsequent spelling of H. mazzuliopsis. Strictly speaking, he introduced a new name, and corrected his error two years later in the “Index to Helices” (Pilsbry 1895: 120) with the note “For H. mazzulopsis read H. subaperta. Ancey’s description appeared June 20; that in Man. Conch., July 1”.

Globulosa: lost. koraegaelia: lectotype [sic!] MHNG-MOLL 117907 from Algeria; type locality: “Djemaa N’Saharidj” (= Djama-N-Saharidj) [Djemaa Saharidj, Mekla, 36.683484° 4.288257°].

Cantareus koraegaelius is a species that is almost inseparable from its congener C. apertus. This also explains why Bourguignat recorded this species from the complete distribution area of the latter species (and including the Algerian lineage). All specimens left in Bourguignat’s collection originating from the localities mentioned are syntypes of Helix koraegaelia. Thus, the type lots contain two different species. To unambiguously fix the use of this specific name, we herewith select the single specimen MHNG-MOLL 117907 from “Djemaa N’Saharidj” in Algeria as lectotype. This locality in Tizi Ouzou is very close to the places, where the anatomically and genetically well-known specimens (see below) have been recorded. The application of the name H. koraegaelia is herewith restricted to specimens exhibiting the character states as explained in this paper forming a new species.

Algeria: Tigzirt/ Tizi Ouzou/ Kabylie, NMBE 534199/1 (specimen preserved and sequenced); Draa Ben Khedaa/ Tizi Ouzou/ Kabylie, NMBE 519923/1 (preserved and sequenced specimen).

Shell thin, medium sized, with a relatively elevated spire and a large last whorl occupying more than half of the complete height of the shell; protoconch medium-sized, consisting of 1.5 smooth whorls; teleoconch with approximately four whorls, separated by a deep, crenulated suture; colour of teleoconch brownish, surface of teleoconch covered by low longitudinal riblets, which are more prominent below the suture, disappearing on the last whorl; aperture rounded, very large, with an inconspicuous whitish lip; umbilicus always completely closed.

penis elongate, club-shaped, epiphallus as long as penis, mrp attaching in the distal third of epiphallus or even closer to penis; flagellum twice the length of the epiphallus; penial lumen smooth; pp2 a broad acute conical papilla with a central perforation structured by thick annuli; pp1 a blind papilla with a slightly broadened apex, the epiphallial pore in a lateral position; atrial and penial lumen with numerous strong crests; distal epiphallial lumen with six broad pilasters, the proximal lumen with elongated ridges.

Vagina short, stem of pedunculus thickened and short, diverticulum extremely longer than the vesicle stem + vesicle, and almost three times longer than the flagellum; glandulae mucosae longer than the dart sac, with a thickened basal part and two subsequent ramifications, tubules thin and weak, less than 10 tubules per stem; atrium dominated by a massive stimulator flap.

(of lectotype): H = 26.3 mm; D = 27.0 mm; PH = 21.8 mm; PD = 17.7 mm.

the two genetically identified specimens originate from Eastern Algeria.

The description of the genital organs is based on the specimen NMBE 534199 from Tigzirt; unfortunately, the other specimen from Draa Ben Khedaa was subadult with only partially developed genital organs.

It is almost impossible to define differences in shell morphology between this new species and C. apertus. In the two genetically identified specimens, the protoconch of C. koraegaelius seems to be larger than in C. apertus, and the shell colour is more or less uniformly brown without any yellowish or greenish streaks. However, the morphology of the genital organs is in fact different: the large triangular pp2 is strikingly different to all what is known so far from the Italian C. apertus, where pp2 is very short to almost completely reduced, so that a “septum” is left.

This specimen was collected by Kobelt in the Gorge d’Isser; it lacks the malleation typical for Cornu aspersum, and thus is here considered to rather constitute a species in Cantareus than in Cornu. However, it also lacks the riblets on the teleoconch, but also has the typical granulation on the whorl exactly like in the specimens from the Djudjura Mts. This form might represent another species close to C. subapertus, but preserved specimens from the canyon are needed to decide about its status. This form might be a separate species endemic to the Gorge d’Isser.

Figures 10–14. 

Cantareus subapertus. Anatomical details of the genital organs; specimen collected at Ighil Bourmi, leg. H. Bouaziz- Yahiatene, NMBE 550458 10 situs of the genital organs, 46 mm total length 11 partly everted genital atrium with the atrial stimulator 12 distal penial tube with pp2 13 penis and epiphallus completely opened showing both papillae, and the internal structure of the penial chamber and the epiphallus 14 detail of the penial lumen; note: the needle represents the epiphallial canal, with pp2 bent upwards to show the ending of the canal. Abbreviations: ag = albumen gland; as = atrial stimulator; div = diverticulum; ds = dart sac; ep = epiphallus; f = flagellum; gm = glandulae mucosae; hd = hermaphroditic duct; mrp = musculus retractor penis; p = penis; pp1 = penial papilla 1; pp2 = penial papilla 2; spo = spermoviduct. All figures not to scale.

Figure 15. 

Cantareus koraegaelius. Lectotype of Helix koraegaelia MHNG-MOLL 117907, shell diameter 27.0 mm. Shell in A frontal B dorsal C lateral D ventral and E apical views F label. Photographs T. Inäbnit, NMBE, × 1.5.

Figures 16–20. 

Cantareus koraegaelius. Shell and anatomical details of the genital organs of dissected and sequenced specimen NMBE 534199; specimen collected at Tigzirt, Tizi Ouzou, Kabylie 16 shell; shell diameter 28.1 mm 17 situs; situs length 57.5 mm 18 lumina of epiphallus, penial chamber, penial papillae and atrium 19 penial papillae 20 atrium with atrial stimulator. Photographs E. Neubert, shell × 1.5.

Figures 21–25. 

Pictures of living specimens of Cantareus species 21 Cantareus apertus: on the left a black specimen with brown shell from Roccapalumba, Sicily, Italy, 15.XI.2009 (Coll. F. Liberto 5532); on the right a yellow specimen with green shell from Prizzi, Sicily, Italy, 15.XI.2009 (Coll. F. Liberto 5545) 22 Cantareus apertus, Niscemi, Sicily Italy, 22.X.2016 23 Cantareus apertus, Niscemi, Sicily, Italy, 22.X.2016, specimens defending themselves by emitting bubbles of slime and a series of tweeting sounds 24 Cantareus apertus, Foggia, Ordona, 41.313889N, 15.622222E, 12.10.2018, leg. G. Martucci (photo/collection I. Sparacio 5031/9) 25 Cantareus subapertus, Algeria, Parc National du Djudjura, 1700 m, 11.X.2008 (Photographs Vela Errol).