Corresponding author: Willy De Mattia (
Academic editor: Eike Neubert
De Mattia W, Reier S, Haring E (2021) Morphological investigation of genital organs and first insights into the phylogeny of the genus
Later,
The name
Wagner’s (1913,
More recently,
Qualitative shell-based tree depicting
According to the currently accepted system (MolluscaBase 2021),
In the present study a comprehensive investigation of the genus
Most of the Sicilian material (mainly
The non-Sicilian specimens were also collected mostly by the first author (WDM) during collecting trips throughout the central and eastern southern Alps, Italian peninsula, and western Balkans from 1992 to date, though interesting specimens belonging to
Ninety-four specimens comprising
Specimens used in the molecular genetic analyses. Collection numbers, laboratory IDs, GenBank accession numbers, and geographical coordinates are listed. More than one
Taxon | Collection No. | Lab ID | GenBank Accession No. | Coordinates | |
---|---|---|---|---|---|
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WDM |
1_1 |
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WDM |
1_4 |
|
no |
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WDM |
2_1 |
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||
WDM |
2_2 |
|
no |
|
|
WDM |
5_1 |
|
|
||
WDM |
14_1 |
|
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||
WDM |
17_2 |
|
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||
WDM |
18_1 |
|
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||
WDM |
19_1 |
|
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||
WDM |
19_2 |
|
no |
|
|
WDM |
Sf_1 |
|
|
|
|
WDM |
Sf_2 |
|
no |
|
|
WDM |
33_1 |
|
no |
|
|
WDM |
33_2 |
|
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||
WDM |
34_1 |
|
no |
|
|
WDM |
34_2 |
|
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||
WDM |
34_3 |
|
no |
|
|
WDM |
34_4 |
|
no |
|
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|
WDM |
36_1 |
|
|
|
WDM |
37_1 |
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||
WDM |
38_1 |
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||
WDM |
38_2 |
|
no |
|
|
WDM |
39_1 |
|
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||
WDM |
40_1 | no | no |
|
|
WDM Siciiiaria 41 | 41_1 |
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WDM Siciiiaria 41 | 41_2 |
|
no |
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|
WDM Siciiiaria 41 | 41_3 |
|
no |
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|
WDM |
42_1 |
|
no |
|
|
WDM |
42_2 |
|
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||
WDM |
42_3 |
|
no |
|
|
WDM |
43_1 |
|
no |
|
|
WDM |
44_1 |
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||
WDM |
44_2 |
|
no |
|
|
WDM |
44_3 |
|
no |
|
|
WDM |
44_4 |
|
no |
|
|
WDM |
46_1 |
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||
WDM |
46_2 |
|
no |
|
|
WDM |
46_3 |
|
no |
|
|
WDM |
47_1 |
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||
WDM |
47_2 |
|
|
||
WDM |
47_3 |
|
no |
|
|
WDM |
47_4 |
|
no |
|
|
WDM |
49_1 |
|
no |
|
|
WDM |
49_2 |
|
no |
|
|
WDM |
50_1 |
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||
WDM |
50_2 |
|
no |
|
|
WDM |
51_1 |
|
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|
|
WDM |
51_2 |
|
no |
|
|
WDM |
52_1 |
|
no |
|
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WDM |
52_2 |
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||
WDM |
54_1 |
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||
WDM |
54_2 |
|
no |
|
|
WDM |
55_1 |
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|
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|
WDM |
55_2 |
|
no |
|
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WDM |
56_1 |
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||
WDM |
57_1 |
|
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|
|
WDM |
59_1 |
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WDM |
60_1 |
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WDM |
60_2 |
|
no |
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WDM |
60_4 |
|
no |
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WDM |
60_5 |
|
no |
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WDM |
61_1 |
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||
WDM |
61_2 |
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WDM |
62_2 |
|
no |
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WDM |
62_3 |
|
no |
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WDM |
63_1 |
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||
WDM |
63_2 |
|
no |
|
|
WDM |
C3_2 |
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no |
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WDM |
C2_2 |
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||
WDM |
C1_1 |
|
no |
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WDM |
C19_1 |
|
no |
|
|
WDM |
C20_1 |
|
no |
|
|
WDM |
C44_1 |
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||
WDM |
C44_3 |
|
no |
|
|
Nordsieck 12167 | 12167_1 |
|
no |
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|
Nordsieck 12167 | 12167_2 |
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WDM |
G64_1 |
|
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WDM |
G64_2 |
|
no |
|
|
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||||
Nordsieck 11786 | 11786_1 |
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Nordsieck 11786 | 11786_2 |
|
no |
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Nordsieck 12100 | 12100_1 |
|
no |
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|
Nordsieck 12147 | 12147_1 |
|
no |
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Nordsieck 12147 | 12147_2 |
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WDM |
P2_1 |
|
no |
|
|
WDM |
P2_2 |
|
no |
|
|
WDM |
P3_1 |
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||
WDM |
P4_1 |
|
no |
|
|
Nordsieck 12194 | 12194_1 |
|
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Nordsieck 12194 | 12194_2 |
|
no |
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Nordsieck 12068 | 12068_2 |
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WDM |
St11_1 |
|
no |
|
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WDM |
St9_2 |
|
|
The DNA extraction was performed in a sterile room using the QIAmp DNeasy Blood and Tissue Kit (
Using the primers 5.8S_LSU-1fw (5'-CTAGCTGCGAGAATTAATGTGA-3') and 28S_LSU-3rv (5'-ACTTTCCCTCACGGTACTTG-3') (
The
Sequencing was performed at Microsynth (Balgach, Switzerland) in both directions using the original PCR primers or, for cloned PCR products, using M13 universal primers. Sequences were processed in Geneious 2.10.3 (
The shells of all dissected specimens were photographed in multiple views (front, back, lateral and internal mouth), including the clausilium distal part, with a Canon EOS camera equipped with 60 mm macro lenses mounted on a Kaiser microslide frame for multi-image stacking. Maximum shell height was measured with a digital caliper (accuracy ± 0.05 mm). For each population at least two specimens were dissected (except for
Anatomical examinations and dissections were performed under a Zeiss stereoscope with a ring LED illumination apparatus, connected to a digital, high-resolution camera and a camera lucida. The genital organs were separated from the rest of the body after a careful crushing of the shell. Dissections were made using a pair of very fine and pointed micro-tweezers (Dumostar Biology 55) and a micro-scissor (FST 150000, Aesculap OC series) in a Petri dish with black paraffin on the bottom and filled with 70% ethanol. The genital organs were fixed with very fine steel micro-pins (commonly used for the preparation of microlepidopteran specimens in entomology). Internal characters of the genital organs (based on cross and longitudinal sections) were examined after dissection with micro-tweezers or a pair of micro-scissors. Measurements were taken using a millimetric measurement scale. The genital organs were photographed in different positions (40–50 high-resolution images) to create an image database. Drawings were prepared by tracing the most representative digital images after contrast enhancement with picture editing software.
Measurements of anatomical features (e.g., length of P, V,
In order to highlight the proportions among the genital parts, the measurements were converted into relative values (ratios). All the ratios are based on measures of at least two specimens. The ratios are
In the first part of the Results we describe the molecular genetic data. For clarity, and in view of the various taxonomic changes in the past, a comprehensive table is presented at the end of the taxonomic part including the new full checklist of the taxa so far considered as
The taxonomic part of the results comprises the descriptions of the taxa with a detailed list of examined specimens and the principal systematic and/or faunistic publications. For the specimens investigated genetically, the identifier is listed together with the GenBank accession numbers (in square brackets) for sequences of the
The collection numbers, voucher numbers, GenBank accession numbers and geographical coordinates for the specimens used in the molecular genetic analyses are listed in Table
BI tree of based on the
The taxa comprising the two main clades I and II cluster in a geographic pattern (maps in Fig.
Mean genetic distances (p-distances in %) for the
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13.4 | |||||||||
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19.6 | 11.4 | ||||||||
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20.5 | 17.0 | ||||||||
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21.5 | 18.3 | 17.1 | |||||||
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21.3 | 21.3 | 20.7 | 18.6 | ||||||
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22.1 | 19.9 | 19.8 | 19.5 | 22.4 | |||||
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21.7 | 21.4 | 20.8 | 20.7 | 22.5 | 21.9 | ||||
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21.3 | 21.8 | 20.2 | 18.6 | 21.0 | 21.9 | 20.7 | |||
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21.0 | 21.4 | 19.4 | 19.4 | 20.7 | 22.8 | 22.3 | 20.1 | |||
|
23.1 | 23.1 | 22.2 | 20.7 | 22.4 | 23.8 | 22.3 | 21.9 | 20.5 | ||
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|
24.5 | 24. 5 | 24.0 | 24.1 | 25.7 | 23.9 | 25.6 | 24.4 | 23.8 | 25.3 |
Intra- and interspecific mean genetic distances (p-distances in %) among the main clades of
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11.3 |
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14.3 | 14.4 |
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17.0 | 16.6 | 17.8 |
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16.9 | 17.3 | 17.3 | 16.4 |
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20.8 | 21.3 | 20.8 | 20.8 | 19.7 |
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21.4 | 20.3 | 21.3 | 20.2 | 20.2 | 12.2 |
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20.9 | 20.5 | 21.6 | 19.7 | 18.9 | 13.3 | 12.5 |
|
The
BI tree of based on the
The tree based on the combined data set (
BI tree of based on the concatenated
Specimens from the same locality delivered very similar or identical sequences. Intraspecific distances (summarised in Table
The taxa comprising the two main clades I (
The high number of
No seasonal variations were detected concerning shape or internal sculpturing of genital organs. The only clear distinction was between adult specimens with full-developed genital organs and immature, subadult specimens with extremely reduced, vestigial genital organs.
The general external morphology of the genital organs is the same in all taxa of
The hermaphroditic gland (
In contrast to the external morphology of the genital organs, the internal sculpturing differs between the various taxa of
Two specimens were found with the spermatophore intact and embedded into the diverticulum of the bursa copulatrix complex or in the vagina and partially inside the first duct of the complex of the bursa copulatrix. The spermatophore of
All the genera
In
In the Southern Alpine
The alpha-taxonomy of
The external shape of the genital organs was found to be of low taxonomic value, because no particular configuration is exclusive and distinctive of one taxon or a group of taxa. On the other side, the internal sculpturing is highly variable, but stable at the population level and the morphology of the genital organs delimits (sub)species, similar to what was found in the genus
The principal rationale of our integrative strategy was whether phylogenetic relationships confirm morphogroups found in a specific area and whether or not they correspond with current taxonomy. We deliberately refrained from applying automated species delimitation methods, for which larger sample sizes would have been necessary. Furthermore, in various land snail species as it was shown such methods may deliver discordant results (e.g.,
So far, mostly mitochondrial marker sequences were used successfully in phylogenetic studies of land snails. The quite frequently used nuclear encoded histone genes H3 and H4 (including the spacer region in-between) often did not add much information or even resulted in confusing trees, probably due to multiple paralogous copies of these genes that escaped homogenisation and/or recombination events after hybridisation (e.g., Harl et al. 2019;
In summary, in the
Finally,
The status of
The concept of interspecies hybridisation (or geographic forms) introduced by Nordsieck (
The list of the examined specimens of
We isolated and depicted the clausiliar plates in order to verify the morphology described by
Taxon | Locality | DNA | Epiphallar formula |
---|---|---|---|
Italy, Sicily, Palermo, Monte Pellegrino, N side of the top plateau, 380 m asl, |
Y | 1ER( |
|
Italy, Sicily, Palermo, Monte Pellegrino, Santuario Santa Rosalia, 420 m asl, |
Y | 1ER( |
|
Italy, Sicily, Palermo, N side of Monte Pellegrino, Punta Priola, Grotte dell’Addaura, 115 m asl, |
Y | 1ER( |
|
Italy, Sicily, Palermo, N side of Monte Gallo near Sferracavallo, 40 m asl, |
Y | 1ER+2ER( |
|
Italy, Sicily, Palermo, E side of Monte Gallo, 140 m asl, |
Y | 1ER+2ER( |
|
Italy, Sicily, Palermo, Grotta Conza, 150 m asl, |
Y | 1ER+2ER( |
|
Italy, Sicily, Palermo, Sferracavallo, via Plauto, 50 m asl, |
Y | 1ER( |
|
Italy, Sicily, Alcamo, Monte Bonifato, top of the mountain, 640 m asl, |
N | 1ER( |
|
Italy, Sicily, Alcamo, Monte Bonifato, top of the mountain, 640 m asl, |
Y | 1ER( |
|
Italy, Sicily, Alcamo, Monte Bonifato, west side of the mountain, over the quarry, 550 m asl, |
N | 1ER( |
|
Italy, Sicily, Piana degli Albanesi, 500 m south of Portella Ginestra, northern cliffs of Monte Kumeta, 970 m asl, |
N | 1ER( |
|
Italy, Sicily, Castellammare del Golfo, Castello di Baida, W of the town along the road to Visicari, 300 m asl, |
N | 1ER( |
|
Italy, Sicily, San Vito lo Capo, Castelluzzo, west cliffs E of town, 120 m asl, |
Y | 1ER( |
|
Italy, Sicily, Castellammare del Golfo, Visicari, 395 m asl, |
Y | 1ER( |
|
Italy, Sicily, Calatafimi, Castello Eufemio, 395 m asl, |
N | 1ER+2ER( |
|
Italy, Sicily, Partinico, W side of the Mount Belliemi, 440 m asl, |
Y | 1ER+2ER( |
|
Italy, Sicily, Alcamo, E side of the Calatubo Castle, 75 m asl, |
Y | 1ER+2ER( |
|
Italy, Sicily, Municipality of Borghetto, road to Romitello, 400 m asl, |
Y | 1ER( |
|
Italy, Sicily, San Giuseppe Jato, quarry E of the town, 630 m asl, |
Y | 1ER+2ER( |
|
Italy, Sicily, Monreale, W part of Monte Kumeta toward Jato Antica, 630 m asl, |
Y | 1ER+2ER( |
|
Italy, Sicily, Monreale, Bosco Ficuzza, Ponte Arcere, 470 m asl, |
Y | 1ER+2ER( |
|
Italy, Sicily, Monreale, N side of Monte Gibilmesi, 890 m asl, |
Y | 1ER+2ER( |
|
Italy, Sicily, Trabia, Contrada Sant’Onofrio, 170 m asl, |
Y | ? |
|
Italy, Sicily, Terrasini, Capo Rama, 30 m asl, |
Y | ||
Italy, Sicily, Cinisi, Piano Margi, 670 m asl, |
Y | 1ER( |
|
Italy, Sicily, Carini, Grotta dei Puntali o dell’Armetta, 90 m asl, |
Y | 1ER+2ER+3ER( |
|
Italy, Sicily, Cinisi, Portella Scaletta [locus typicus], 90 m asl, |
Y | 1ER+ |
|
Italy, Sicily, Cinisi, along the SS113 road from Villagrazia di Carini to Cinisi, 70 m asl, |
Y | 1ER+ |
|
Italy, Sicily, Castelluzzo, cliffs W of the Tonnara di Monte Cofano, 50 m asl, |
Y | 1ER+ |
|
Italy, Sicily, Custonaci, cliffs N of the Mangiapane cave, 63 m asl, |
Y | 1ER( |
|
Italy, Sicily, Custonaci, Baglio Messina, 340 m asl, |
Y | 1ER+2ER( |
|
Italy, Sicily, Custonaci, Buffara, NE side of the hollow, 150 m asl, |
Y | 1ER( |
|
Italy, Sicily, Custonaci, contrada Scurati, crossing with Strada Provinciale 18, 65 m asl, |
Y | 1ER( |
|
Italy, Sicily, Custonaci, contrada Scurati, 60 m asl, |
Y | 1ER( |
|
Italy, Sicily, Custonaci, contrada Scurati, limestone cliffs S of the village, 60 m asl, |
Y | 1ER( |
|
Italy, Sicily, Castelluzzo, Monte Cofano E of Tonnara di Monte Cofano, 80 m asl, |
N | 1ER+2ER+3ER( |
|
Italy, Sicily, San Vito lo Capo, boulders W of the town, 50 m asl, 38°10'51.85"N12°43'37.70"E, W. De Mattia and J. Macor leg. | Y | 1ER+2ER+3ER( |
|
Italy, Sicily, San Vito lo Capo, Castelluzzo, west cliffs E of town, 120 m asl, |
Y | 1ER+2ER( |
|
Italy, Sicily, San Vito lo Capo, cliffs S of “El-Bahira” camping, 60 m asl, 38°08'48.81"N12°44'06.61"E, W. De Mattia and J. Macor leg. | Y | 1ER+2ER( |
|
Italy, Sicily, San Vito lo Capo, Torre delle Usciere, 5 m asl, |
Y | 1ER+2ER+3ER( |
|
Italy, Sicily, San Vito Lo Capo, Macari, E side of Monte Speziale, 75 m asl, |
Y | 1ER+2ER( |
Distribution of the clausilium character states within the
Taxon | n | Boettger’s groups: Gruppe der |
Clausiliar plate distally receding/not receding | Clausiliar plate gutter-like/not gutter like | Clausiliar plate bent upwards/not bent upwards |
---|---|---|---|---|---|
22 |
|
receding | gutter-like | bent | |
18 |
|
receding | gutter-like | bent | |
15 |
|
receding | gutter-like | bent | |
20 |
|
not receding | not gutter-like | not bent | |
36 |
|
not receding | not gutter-like | bent | |
25 |
|
not receding | not gutter-like | bent | |
12 |
|
not receding | not gutter-like | bent | |
21 |
|
not receding | not gutter-like | bent | |
15 |
|
not receding | not gutter-like | bent | |
9 |
|
partially receding | gutter-like | bent | |
7 |
|
not receding | not gutter-like | bent | |
11 |
|
partially receding | partially gutter-like | not bent | |
31 |
|
not receding | not gutter-like | bent | |
33 |
|
not receding | not gutter-like | bent | |
6 |
|
not receding | not gutter-like | not bent | |
27 |
|
not receding | not gutter-like | not bent | |
19 |
|
not receding | not gutter-like | not bent | |
29 |
|
not receding | not gutter-like | not bent | |
35 |
|
not receding | not gutter-like | not bent | |
21 |
|
not receding | not gutter-like | not bent |
Main morphological differences between
|
|
|
Shell – waxy surface | not present | only in |
Shell – percostate | not present | only in |
Shell – clausiliar plate gutter-like | only in |
not present |
Shell – clausiliar plate receding | only in |
not present |
Ganitalia – penial internal papillose sculpturing | not present | only in |
The shell of
Italy, Sicily, Palermo, Monte Pellegrino, N side of the top plateau, 380 m asl,
Shell mostly decollate; 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 upwards and more or less pointed (as in
(n = 25, decollate): shell height 19.4 ± 0.8, whorl width 5.1 ± 0.2, aperture height 3.9 ± 0.2, aperture width 2.7 ± 0.2.
The
The A is smooth or with weak traces of the distal penial pleats. The P presents 4–6 longitudinal and very irregular pleats. These pleats are very variable in thickness and sculpture, being both smooth or segmented along the same pleat. These pleats often split (proximally, distally or both) into smaller pleats. The fine structure of the wall is smooth. The
The taxon is limited to the calcareous mountains northern of Palermo: Monte Pellegrino and the southern slopes of Monte Gallo.
Italy, Sicily, Palermo, N side of Monte Pellegrino, Punta Priola, Grotte dell’Addaura, 115 m asl,
1
Shell decollate; whorls ribbed; dorsal keel weak but distinguishable; inferior lamella very high; two anterior upper palatal plicae present, both of them separated from the lunella; parietalis short ends before the beginning of the spiralis; palatal edge of clausilium plate distally receding, plate gutter-like narrowed, palatal edge against distal end bent upwards and more or less pointed.
The shell is elongated, fusiform, sinistral and decollate, rarely not decollate. It is brown to reddish brown in colour. The external surface has small raised ribs almost equally arranged in all whorls of the teleoconch. The spire slowly and regularly grows with eight or nine whorls that are only slightly convex. The sutures are moderately shallow with very rare whitish papillae present towards the apex. The basal and the cervical keels are weak but distinguishable. The umbilicus is closed. The aperture width is ~ 1⁄4 of shell height and subovoid in shape. The
The
The A is smooth. The P presents very smooth and scarcely elevated weak transverse pleats. The fine structure of the wall is smooth. The
The taxon is named after the Addaura caves, a complex of three natural caverns where wall engravings dated to the Paleolithic and the Mesolithic were discovered.
The case of
Italy, Sicily, Palermo, N side of Monte Gallo near Sferracavallo, 40 m asl,
(n = 18, not decollate): shell height 17.9 ± 0.6, whorl width 4.2 ± 0.2, aperture height 3.9 ± 0.2, aperture width 2.9 ± 0.4.
The
The A shows a set of longitudinal irregular pleats. The P presents four to five smooth longitudinal or slightly fringed pleats. The fine structure of the wall is smooth. The
Following the new taxonomical results, the distribution of
The population from Via Plauto, concerning the shell, matches with the “Sferracavallo form” described by
Italy, Sicily, Palermo, Grotta Conza, 150 m asl,
(
(n = 35, decollate): shell height 18.2 ± 2.1, whorl width 4.3 ± 0.2, aperture height 4.5 ± 0.2, aperture width 3.4 ± 0.2.
The
The A shows a set of irregular fleshy folds. The P presents two big smooth longitudinal pleats that occupy almost the whole internal penial space. The fine structure of the wall is smooth. The
In the
Two main anatomical types of male (penial) internal sculpturing are found. The first type presents clear longitudinal pleats that run from the origin of penial pseudopapilla down to (or almost to) the atrium. These longitudinal pleats can be either (almost) completely smooth (Monte Belliemi) or more or less markedly fringed (Calatubo, Romitello, Ficuzza, Jato city park, Monte Kumeta, and Castelluzzo). The second type presents transverse and strong smooth pleats (Bonifato, Gibilmesi, and Visicari).
The first type corresponds to
Throughout the whole
The populations belonging to this subclade were collected in two main ecological niches. Most of them (type 1) were found on limestone rocks and cliffs (Jato city park, Bonifato, Calatubo, Gibilmesi, Monte Kumeta, Romitello, Monte Belliemi and Visicari), whereas the other populations (type 2), from Ficuzza and Castelluzzo, were exclusively found on tree trunks.
Concerning the haplogroups of the
The position of
Two additional new subspecies of
Italy, Sicily, San Vito lo Capo, Castelluzzo, west cliffs E of town, 120 m asl,
The E internal sculpturing presents a variety of arrangements, as: two main large smooth or fringed longitudinal pleats that gradually disappear toward the distal origin of the vas deferens or 4 thin smooth longitudinal pleats that abruptly disappear turning into an irregular texture of small dense papillae. The V can be either smooth and showing a very fine granulation or showing a coarse chevron pattern made of large fleshy pleats, merging together along the median longitudinal axis.
Italy, Sicily, Partinico, W side of the Mount Belliemi, 440 m asl,
(n = 40, not decollate, Monte Belliemii): shell height 17.6 ± 1.2, whorl width 3.1 ± 0.1, aperture height 2.9 ± 0.2, aperture width 2.0 ± 0.1. (n = 22, not decollate, Calatubo): shell height 19.6 ± 1.4, whorl width 3.9 ± 0.2, aperture height 3.4 ± 0.2, aperture width 3.0 ± 0.2.
The subspecies inhabits both scattered isolated boulders (Monte Belliemi) or limestone cliffs (Calatubo Castle). This subspecies is known only from two localities with very limited size, although it is abundant. Both localities are not included in protected areas.
The Calatubo population is bigger than the population from the type locality (Monte Belliemi). The average shell heights are 19.6 ± 1.4 vs. 17.6 ± 1.2. The two populations occupy remarkably different ecological niches: high cliffs vs isolated small boulders scattered throughout open fields. Could the different ecology of the two populations be the origin or cause of such different dimensions of the shell? This interesting hypothesis could be tested also including other
Italy, Sicily, Municipality of Borgetto, Anime Sante hill, road to Romitello, 400 m asl,
1
Shell not decollate; whorls rib-striated; dorsal keel not distinguishable; inferior lamella very high; anterior upper palatal plicae present detached from the lunella; parietalis short partially overlapping of the spiralis; palatal edge of clausilium plate distally receding, plate gutter-like narrowed, palatal edge against distal end bent upwards and more or less pointed.
The
The subspecies is found on limestone boulders and under scattered rocks.
Italy, Sicily, San Giuseppe Jato, quarry E of the town, 630 m asl,
1
Shell decollate, rarely not decollate; whorls ribbed; dorsal keel weak but distinguishable; inferior lamella very high; anterior upper palatal plicae present and detached from the lunella; parietalis long; palatal edge of clausilium plate distally receding, plate gutter-like, narrowed, palatal edge against distal end bent upwards and more or less pointed.
The shell of
Moreover, in the text Nordsieck cited samples
Monte Kumeta was also deemed by Nordsieck as a spelling mistake for Monte della Fiera (pers. comm., April 2021) and stating that “form from Monte della Fiera belongs to the northern subspecies (of
In the
We agree that the sample depicted in Fig.
Regarding shell morphology, we carefully browsed Schmidt’s description (1868: 41–43). Few important details better fit with the shell morphology of the “northern” type form of
The new subspecies is an obliged limestone rock-dweller, exclusively collected on walls and cliffs, both natural or resulting from the previous quarrying activity.
Italy, Sicily, Monreale, W part of Monte Kumeta toward Jato Antica, 630 m asl,
1
Shell not decollate; whorls ribbed; dorsal keel weak but distinguishable; inferior lamella high or very high; anterior upper palatal plicae present and detached from the lunella; parietalis long; palatal edge of clausilium plate distally receding, plate gutter-like narrowed, palatal edge against distal end bent upwards and more or less pointed.
Italy, Sicily, Monreale, Bosco Ficuzza, Ponte Arcere, 470 m asl,
Following
Italy, Sicily, Monreale, N side of Monte Gibilmesi, 890 m asl,
1
Shell not decollate; whorls finely striated; dorsal keel absent or barely distinguishable; inferior lamella high or very high; anterior upper palatal plicae present but weak, detached from the lunella; parietalis very long; palatal edge of clausilium plate distally receding, plate somehow cylindrical, palatal edge against distal end bent upwards and more or less pointed.
The morphologically most similar taxon to
This subspecies was found climbing on limestone cliffs hiding among moss and rocks crevices.
In the
The status of
Italy, Sicily, Trabia, Contrada Sant’Onofrio, 170 m asl,
Italy, Sicily, Terrasini, Capo Rama, 30 m asl,
(n = 40, decollate): shell height 16.3 ± 1.2, whorl width 4.0 ± 0.1, aperture height 3.1 ± 0.2, aperture width 2.3 ± 0.2.
As a result, the actual distributional range of
Monte San Calogero in the east (cited by Nordsieck as also included in the
Italy, Sicily, Cinisi, Piano Margi, 670 m asl,
(n = 18, decollate): shell height 21.5 ± 1.3, whorl width 5.2 ± 0.3, aperture height 4.6 ± 0.2, aperture width 3.4 ± 0.4.
The
Italy, Sicily, Carini, Grotta dei Puntali o dell’Armetta, 90 m asl,
1
Shell decollate; whorls moderately ribbed; dorsal keel prominent; inferior lamella high; anterior upper palatal plica present, knob present at distal end of the upper palatal plica, detached from the lunella; lower anterior palatal plica (
The taxon is named after the Grotte dell’Armetta, a complex of natural caves where wall engravings and stone handcrafted tools, dated from the Paleolithic to the Bronze Age, were discovered (
Italy, Sicily, Cinisi, Portella Scaletta, 90 m asl,
The P of the population from the road to Cinisi (300 from NNE of the type locality), proximally shows pleats that are longitudinal and progressively turn into a transverse pattern as proceeding towards the A. The
90% of the specimens (n = 45) of
In Table
Main morphological differences (shell and genital organs) among the
|
|
|
|
|
Whorls | ribbed, percostate | ribbed | weakly rib-striated | densely ribbed |
Surface | opaque | opaque | white surface layer | white surface layer |
Dorsal keel | weak | weak or missing | missing | missing |
|
missing | present, visible from outside | present, mostly not visible from outside | mostly missing |
|
very short or mostly missing | present, visible from outside | present, visible from outside | very short to knob-like, not visible from outside |
Spiralis | not overlapping parietalis | overlapping parietalis | slightly overlapping parietalis | mostly not overlapping parietalis |
Sculpturing of atrium | fleshy pleats | large fleshy pleats | irregular folds | irregular fleshy folds. |
Sculpturing of vagina | smooth to an irregular | smooth to an irregular | smooth to an irregular | distally irregular pleats, proximally oblique pleats |
Sculpturing of penis | longitudinal fleshy smooth pleats | longitudinal fleshy pleats | smooth, tubercles, longitudinal, segmented pleats | longitudinal fleshy smooth pleats |
Pseudopapilla | short and conical | small and rounded | big and elongate to small and round | little, rhombus-shaped and smooth |
Epiphallar ring | single | single | two or three | two |
Sculpturing of epiphallus | two main longitudinal pleats | two main longitudinal pleats | two main longitudinal pleats | three or four irregular longitudinal fringed pleats |
. The morphology of the genital organs of
The shell is remarkably ribbed to percostated. It shares few characters with
Although
Italy, Sicily, Castelluzzo, cliffs W of the Tonnara di Monte Cofano, 50 m asl,
(
This species has a very limited distribution range, restricted to the western part of Monte Cofano north of Custonaci (Tonnara di Cofano and Cornino). A (probably) introduced population is known to occur on the Island of Favignana (Trapani) (
The general genital outline is stable for all the dissected populations. The internal sculpturing of the penis reveals a basic pattern of one to four longitudinal pleats. The remaining main genital internal parts (penial pseudopapilla, epiphallus and atrium) show good stability despite slight variations, which will be described below. Only the internal wall of the vagina could be either smooth or with a strong sculpturing.
The shell is remarkably ribbed although the number of the ribs (density) progressively decreases along a geographical S-N axis, whereby simultaneously the thickness of the ribs increases. Southern populations, e.g., from Dolina Buffara and Baglio Messina, present a dense ribbing (with an average of 70 ribs along the first whorl) whereby the northernmost populations from the environs of Mangiapane Caves and Scurati show much less rib density (with an average of 32 ribs along the first whorl). The northern populations are found a few hundred meters from the southernmost population of
Italy, Sicily, Custonaci, contrada Scurati, crossing with Strada Procinciale 19, 60 m asl,
(n = 45, decollate): shell height 20.1 ± 0.7, whorl width 4.7 ± 0.3, aperture height 4.7 ± 0.4, aperture width 3.1 ± 0.1.
The distribution range of
Italy, Sicily, Castelluzzo, Monte Cofano E of Tonnara di Monte Cofano, 80 m asl,
(
Italy, Sicily, San Vito Lo Capo, Macari, E side of Monte Speziale, 75 m asl,
The genus
In our molecular genetic analysis (Figs
The separation of
Both
Following our genital anatomical investigations, the common genital anatomical traits deemed by
Italy, Sicily, Marettimo, W of the town, path to Pizzo Falcone. 80 m asl,
(n = 35, decollate): shell height 18.5 ± 1.0, whorl width 4.6 ± 0.2, aperture height 4.0 ± 0.3, aperture width 2.9 ± 0.2.
Tunisia, above Zaghouan (road to summit), 789 m asl,
(
The separation of
The taxa included in the alpine
Despite the low support values, the
The results presented by Scheel and Hausdorf (2012: 3799, 3801) and
The stenzioid subspecies of
The list of the dissected material (at least two specimens per population) is found in the Table
Examined taxa of
Taxon | Specimens examined | Shell stenzioid/ nonstenzioid | DNA | Epiphallar formula |
---|---|---|---|---|
Trasquera di Iselle, Verbano, Piedmont, Italy. 750 m asl. |
- | Y | ||
Margone, Val di Lanzo, Usseglio, Torino, Piedmont, Italy. 1500 m asl, |
- | N | ||
Sentiero per Prà del Vecia, Piedicavallo, Biella, Piedmont, Italy. 1200 m asl. |
- | N | ||
Santuario di Oropa, Biella, Piedmont, Italy. 1100 m asl |
- | N | ||
Ogulin city centre, Croatia. 320 m asl. |
- | N | ||
San Donato, Barbarano Vicentino, Vicenza, Veneto, Italy. 300 m asl. |
nonstenzioid | Y | ||
Brescia, city castle walls, Lombardia, Italy. 200 m asl. |
nonstenzioid | N | ||
Albergo alla Tolla, Val Ampola, Storo, Trentino-Alto Adige, Italy. 720 m asl. |
nonstenzioid | N | ||
Naole, San Zeno di Montagna, Verona, Veneto, Italy. 900 m asl. |
nonstenzioid | Y | ||
Loc. Galleria, Bracca, Val Serina, Bergamo, Lombardia, Italy. |
stenzioid | N | ||
Ballabio, Lecco, Lombardia, Italy. 710 m asl. |
stenzioid | N | ||
Bedulita, Bergamo, Lombardia, Italy. 625 m asl. |
nonstenzioid | N | ||
Val di Lorina, Storo, Brescia, Lombardia, Italy. 600 m asl. |
stenzioid | Y | ||
Udine, Friuli-Venezia Giulia, Italy. 140 m asl. |
nonstenzioid | N | ||
Udine, Friuli-Venezia Giulia, Italy. 140 m asl. |
nonstenzioid | N | ||
Serravalle near Vittorio Veneto, Veneto, Italy. 500 m asl. |
nonstenzioid | nonstenzioid | ||
Forte Cima Ora, Anfo, Brescia, Lombardia, Italy. 1500 m asl. |
stenzioid | N | ||
Cima Caldoline, Lavenone, Brescia, Lombardia, Italy. 1700 m asl. |
stenzioid | N | ||
Lavina, Val Taleggio, Bergamo, Lombardia, Italy. 650 m asl. |
stenzioid | N | ||
San Romedio, Trento, Italy. 730 m asl. |
stenzioid | N | EP( |
|
Fai della Paganella, Trento, Trentino-Alto Adige, Italy. 600 m asl. |
stenzioid | N | EP( |
|
Bosplans, Pordenone, Friuli-Venezia Giulia, Italy. 500 m asl. |
stenzioid | Y | EP( |
|
Val Resia, sella di Carnizza, Tarvisio, Friuli-Venezia Giulia, Italy. 400 m asl. |
stenzioid | N | EP( |
|
Grotte di Oliero Valstagna, Vicenza, Veneto, Italy. 800 m asl. |
stenzioid | N | EP( |
|
Val Fonda, Misurina, Belluno, Veneto, Italy. 1500 m asl. |
stenzioid | Y | EP( |
|
Cison di Val Marino, Treviso, Veneto, Italy. 250 m asl. |
stenzioid | N | no pseudopapilla | |
Loc. La Goccia, Foza, Valstagna, Vicenza, Veneto, Italy. 800 m asl. |
stenzioid | N | EP( |
|
Val Fiscalina, Croda Rossa, Sesto, Bolzano, Trentino-Alto Adige, Italy. 1500 m asl. |
stenzioid | N | EP( |
The present anatomical investigation indeed generated consistent results that are congruent with the molecular genetic outcomes.
The genus
Boettger, 1877, introduced the genus
He introduced a new subspecific taxon,
Nordsieck cites
In our
Three out of five subspecies of
The four populations of
This parapseudopapilla represents an intermediate form between the simple structure of
Italy, Piedmont, Trasquera di Iselle, Verbano. 750 m asl,
The
The A and the P are smooth, with a very fine granulated sculpturing. The V shows many smooth longitudinal pleats. The smooth
Italy, Piedmont, Torino, Margone, Val di Lanzo, Usseglio. 1500 m asl,
Italy, Piedmont, Biella, Santuario di Oropa. 1100 m asl
The Southern Alpine
Its shell is weakly striated to frankly ribbed, with evident sutural papillae and a strong lunella, which is remarkably curved. Twelve out of the seventeen currently valid subspecies of
Croatia, Ogulin city centre, 320 m asl,
The actual distribution of
Italy, Veneto, Vicenza, San Donato, Barbarano Vicentino. 300 m asl,
Italy, Lombardia, Brescia, city castle walls. 200 m asl,
This taxon is known to occur in Val Ampola and Val Lorina (Province of Trento, Trentino-Alto Adige, Italy). It is also reported by
Italy, Trentino-Alto Adige, Trento, Storo, Albergo alla Tolla, Val Ampola. 720 m asl,
This taxon, previously considered as a junior synonym of
Italy, Veneto, Verona, Naole, San Zeno di Montagna. 900 m asl,
This small subspecies, found exclusively at higher altitudes, was previously considered as a transitional form between
This subspecies is known from Val Serina (province of Bergamo, Lombardia), Italy (
Italy, Lombardia, Bergamo, Val Serina, Loc. Galleria, Bracca.
This subspecies is known to occur in central-western Lombardia, in the provinces of Bergamo and Lecco (
Italy, Lombardia, Lecco, Ballabio. 710 m asl,
Known to occur in central Lombardia, Italy (
Italy, Lombardia, Bergamo, Bedulita. 625 m asl,
The distribution of this subspecies is limited to Val Lorina and surrounding peaks, across the border of Trento and Brescia provinces (Lombardia and Trentino-Alto Adige), Italy (
Italy, Lombardia, Brescia, Val di Lorina, Storo. 600 m asl,
During field collecting (WDM), five populations belonging to
The type locality of the nominate subspecies is less than 50 km west of Portogruaro. Thus, following
Despite
The Friulian populations with translucent shell and reduced shell dimensions are here named
Italy, Friuli-Venezia Giulia, Udine, castle hill. 140 m asl,
This taxon is widespread throughout the southeastern Alps, from Val Sugana in the west, Alpi Venete, Alps of Belluno and the axis Ampezzo-Pordenone in the east (Italy) (
Italy, Veneto, Serravalle (L.T.) near Vittorio Veneto. 500 m asl,
The actual status and validity of
The taxon is known for several stations around the Idro Lake, at the boundary between the provinces of Brescia and Trento (
Forte Cima Ora, Anfo, Brescia, Lombardia, Italy. 1500 m asl,
The V is 3 × longer than the
The A is smooth. The V presents smooth longitudinal pleats. The P is completely smooth. The
In the surroundings of Forte Cima Ora (Anfo) we collected this costate form that was wrongly identified as
This subspecific taxon of
Italy, Lombardia, Brescia, Lavenone, Cima Caldoline. 1700 m asl,
This taxon is known only from the Val Brembana and Val Taleggio (Bergamo, Lombardia, Italy) (
Italy, Lombardia, Bergamo, Lavina, Val Taleggio. 650 m asl,
Eight of nine subspecies of
Italy, Trentino-Alto Adige, including westernmost Dolomites in the surroundings of Schlern mountain, Val di Non and Passo della Mendola (
Italy, Trentino-Alto Adige, Trento, San Romedio. 730 m asl,
Italy, Trentino-Alto Adige, Trento, Fai della Paganella. 600 m asl,
Massif of Mount Paganella, Trentino-Alto Adige, Italy (
Italy, Friuli-Venezia Giulia, Pordenone, Bosplans. 500 m asl,
The V is much shorter than the
The A, the V and the P show many more or less irregular (longitudinal to oblique) smooth pleats. The hemipapilla is smooth to moderately folded and usually short. The outlet of the hemipapilla is slit-like. The E shows two fringed
Italy, Veneto, Vicenza, Grotte di Oliero near Valstagna. 800 m asl,
Italy, Veneto, Belluno, Misurina, Val Fonda. 1500 m asl,
Italy, Veneto, Treviso, Cison di Val Marino. 250 m asl,
Italy, Veneto, Vicenza, Valstagna, Foza, Loc. La Goccia. 800 m asl,
Italy, Trentino-Alto Adige, Bolzano, Sesto, Val Fiscalina (Fischleintal), Croda Rossa. 1500 m asl,
The V is smooth. The P shows three to four irregularly fringed pleats that continue as far as the A. The hemipapilla is very short, conical and moderately folded. The outlet of the hemipapilla is triangular. The E shows two fringed
The original name of this “Form vom Fischleintal in Südtirol” (
In our study, two populations of
Examined taxa of
Taxon | Examined specimens | Epiphallar formula | DNA |
---|---|---|---|
Muggia, Europa Gardens, Trieste, Friuli-Venezia Giulia, Italy. 30 m asl. |
|
Y | |
Apricena, Foggia, Puglia, Italy. 75 m asl. |
Y | ||
Mattinata close to Vieste, Puglia, Italy. 500 m asl. |
N | ||
Reka Dubrovačka, Sustjepan, Dubrovnik, Croatia. 5 m asl. |
N | ||
Rijeka Reževići, on the Budva–Petrovac road, 100 m asl. |
N | ||
Orjen Mts, Podi and Kameno, Herceg Novi-Trebinije Road, Montenegro. 325 m asl. |
N | ||
Punta Penne next to the Airport, Brindisi, Puglia, Italy. 5 m asl. |
|
Y | |
San Rufo, Passo della Sentinella, Salerno, Campania, Italy. 850 m asl. |
N | ||
Serra San Bruno, Vibo Valentia, Calabria, Italy. 945 m asl. |
N | ||
Altilia, Cosenza, Calabria, Italy. 460 m asl. |
N | ||
Gerace, Strada Provinciale 1, 3.5 km NW of the town, Reggio Calabria, Calabria, Italy. 400 m asl. |
N | ||
Gardiki fortress, Corfu, Greece. 40 m asl. |
N | ||
Monte Telegrafo, Argentario, Toscana, Italy. 600 m asl. |
N | ||
Balvano, Potenza, Basilicata; Italy. 450 m asl. |
N | ||
Balvano, near the castle, Potenza, Basilicata; Italy. 450 m asl. |
N | ||
Lafki, Corfu, Greece. 420 m asl. |
N | ||
Vlorë district, Qeparo, macchia with tree spurges W of the village, 20 m, 50 m asl. |
|
N | |
Delvinë District, Krongj, rocky grassland at the peak region of Mt Koqinolithar, Albania. 890 m asl. |
N | ||
Cosenza, Via Porta Piana, Calabria, Italy. 300 m asl. |
Y | ||
Castello di San Lucido, Paola, Calabria, Italy. 120 m asl. |
Y | ||
Amantea, Cosenza, Calabria, Italy. 30 m asl. |
N | ||
Contrada Piscitello, Belpasso, Catania, Sicily, Italy. 650 m asl. |
N | ||
Contrada Rizzuto, Cosenza, Calabria, Italy. 770 m asl. |
N | ||
Contrada Rizzuto road to San Bartolo, Cosenza, Calabria, Italy. 740 m asl. |
N |
The actual distribution of the genus
Italy, Friuli-Venezia Giulia, Muggia, Trieste, Europa Gardens. 20 m asl,
Italy, Puglia, Foggia, Apricena. 75 m asl,
The taxon is exclusively known from the southern slopes of Monte Sant’Angelo, Gargano Peninsula (Puglia, Italy) (
Italy, Puglia, Mattinata close to Vieste, southern slopes of the Gargano Peninsula. 500 m asl,
As a shell,
The nominate subspecies is distributed from southern Dalmatia (from the Neretva River southward), Herzegovina, Montenegro to northern Albania.
Croatia, Dubrovnik, Sustjepan, Reka Dubrovačka. 5 m asl,
The V is much shorter than the
This taxon is restricted to the Salento (Puglia, Italy) and central Albania with the type form (
Italy, Puglia, Brindisi, Punta Penne next to the Airport. 5 m asl,
The morphology of the genital organs of this population is remarkably different from all other
This species is restricted to the eastern side of the Monti Alburni, Campania, Italy.
Italy, Campania, Salerno, San Rufo, Passo della Sentinella. 850 m asl,
Italy, Calabria, Vibo Valentia, Serra San Bruno. 945 m asl,
This species is restricted to higher altitudes from 800 m asl up. It is found in mixed
Calabria, Italy (
Italy, Calabria, Cosenza, Altilia. 460 m asl,
The species is distributed in southwestern Albania and Corfu, Greece (
Greece, Corfu, Gardiki fortress. 40 m asl,
The nominate subspecies is widespread in central and southern peninsular Italy [from Tuscany (Monte Argentario) and S. Abruzzo to N. Calabria] (
Italy, Toscana, Grosseto, Porto Ercole, Monte Telegrafo. 600 m asl,
This taxon is present in Basilicata, known from the area surrounding Balvano (Italy) (
Italy, Basilicata, Potenza, Balvano, near the castle. 450 m asl,
The nominate subspecies is restricted to the Island of Corfu, Greece.
Greece, Corfu, Lafki. 420 m asl,
Albania, Delvinë District, Krongj, rocky grassland at the peak region of Mt Koqinolithar. 890 m asl,
Mainland populations of
This taxon is restricted to southern Albania from Tepelenë area near Lum i Bençës (
Albania, Vlorë district, Qeparo, macchia with tree spurges W of the village, 20 m, 50 m asl,
The lack of the pseudopapilla represents a possible apomorphic state of the epiphallus as already seen in other genera (e.g.,
Italy, Calabria, Cosenza, Via Porta Piana. 300 m asl,
Cosenza and Paola, Calabria (Figs
Unfortunately, genetic molecular data are not yet available from topotypic populations from the Mongibello-Etna (Sicily) thus, although the anatomical investigations highlight reliable differences, the relationships with and the status of the Calabrian populations still have to be properly assessed.
Italy, Calabria, Cosenza, Contrada Rizzuto. 770 m asl,
The genus
Examined taxa of
Taxon | Examined specimens | Epiphallar formula | DNA |
---|---|---|---|
Vicovaro, Latina, Lazio, Italy. 320 m asl. |
Y | ||
Jato Antica, Monreale, Sicily, Italy, 830 m asl, |
Y | ||
western part of Monte Kumeta toward Jato Antica, Monreale, Sicily, Italy, 630 m asl, |
Y | ||
Monte Bonifato, west side of the mountain, over the quarry, Alcamo, Sicily, Italy. 550 m asl, |
Y | ||
road to the Norman Castle, Stilo, Reggio Calabria, Calabria, Italy. 320 m asl. |
Y |
New
|
MolluscaBase 2021 | De Mattia, Reier and Haring 2021 |
---|---|---|
see |
||
The system of
For the genus
The authors are very much obliged to Alessandro Margelli, Gianbattista Nardi, Ivano Niero, and Hartmut Nordsieck for providing alcohol-preserved specimens for study. We thank Hartmut Nordsieck for species identification and taxonomic remarks. The first author is grateful to Claude and Amandine Evanno who authorised the use of their photograph of