Research Article |
Corresponding author: Willy De Mattia ( willy.demattia@icgeb.org ) Academic editor: Eike Neubert
© 2021 Willy De Mattia, Susanne Reier, Elisabeth Haring.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
De Mattia W, Reier S, Haring E (2021) Morphological investigation of genital organs and first insights into the phylogeny of the genus Siciliaria Vest, 1867 as a basis for a taxonomic revision (Mollusca, Gastropoda, Clausiliidae). ZooKeys 1077: 1-175. https://doi.org/10.3897/zookeys.1077.67081
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The taxonomy and systematics of the door snail genus Siciliaria was revised based on an integrative approach including a comprehensive genital anatomical investigation, which was combined with shell morphology and DNA sequence data (mitochondrial COI, nuclear ITS2 sequences). The genital morphology of 120 specimens of 22 taxa from 44 populations was investigated, and a new general description of the genital morphology of the genus is provided. Additionally, 26 specimens of 14 taxa of five genera (MolluscaBase 2021) of Alopiinae (Mauritanica, Charpentieria, Stigmatica, Gibbularia, Papillifera) were included in the genetic analyses. New anatomical structures are described: the parapseudopapilla for Charpentieria dyodon and the hemipapilla for Charpentieria stenzii. In the phylogenetic tree based on COI sequences, the species of the genus Siciliaria s. l. from northwestern Sicily were found within two separate highly supported main clades. In the tree based on the nuclear ITS2 marker sequence, resolution was considerably lower but it partially confirmed the mitochondrial tree. The genus Sicania Tomlin, 1929 (corresponding to main clade II in the trees) is re-introduced. Siciliaria scarificata did not appear in one of the two main clades but clustered together with Mauritanica perinni polygyra. Concerning monophyly of species, only the widely distributed S. calcarae was paraphyletic in the COI tree, a finding that has to be investigated further with multiple marker sequences. For the other genera (Mauritanica, Charpentieria, Stigmatica, Gibbularia, and Papillifera) detailed descriptions of the anatomy of the genital organs of 46 taxa for a total amount of 133 dissected specimens are also provided here. Some of these taxa could be included in the phylogenetic analysis. Although the taxon sampling of these taxa was far from being complete, the comprehensive data provided here (concerning morphology, genetics, and distribution) provide first insights into the phylogenetic relationships of this diverse group of clausiliid taxa. The following six taxa new to science are described: Siciliaria grohmanniana addaurae ssp. nov., Siciliaria calcarae borgettensis ssp. nov., Siciliaria calcarae jatinensis ssp. nov., Siciliaria calcarae parajatinensis ssp. nov., Siciliaria calcarae cruenta ssp. nov., and Siciliaria tiberii armettensis ssp. nov. Anatomy of genital organs, Clausiliidae, phylogeny, Siciliaria, systematics, taxonomy
Siciliaria Vest, 1867 is a genus of the family Clausiliidae that is distributed along the calcareous ranges of northwestern Sicily (Italy). It is distributed from Egadi Islands and Trapani in the west to western Madonie in the east. The southern boundaries are represented by the line (west to east) Trapani, Calatafimi, Bosco Ficuzza and Monte San Calogero.
Later,
The name Sicania had been introduced in order to replace the pre-occupied name Trinacria
Wagner’s (1913,
More recently,
According to the currently accepted system (MolluscaBase 2021), Siciliaria is deemed as a subgenus of the all-inclusive genus Charpentieria, with 12 species and 17 subspecies. It currently comprises six subgenera with 29 taxa and ~ 70 subspecific taxa. The phylogenetic relationships among Siciliaria, Charpentieria, Stigmatica, Gibbularia and Mauritanica are still unclear. Scheel and Hausdorf (2012),
In the present study a comprehensive investigation of the genus Siciliaria Vest, 1867 (type species: Clausilia grohamnniana Rossmässler, 1836) was performed, including morphology of the genital organs and DNA sequence analyses, which were interpreted in the light of the current taxonomic system (
Most of the Sicilian material (mainly Siciliaria and Papillifera), unless differently stated, was collected during four field collecting trips by the first author (WDM) and currently kept in the Willy De Mattia collection (Syracuse, Italy). The type material of the newly described taxa was deposited in the Natural History Museum in Vienna. Part of the material was made available by other private collections. In order to increase the sample (n) for shell measurements, also dead material was extensively collected. In this study we investigated the anatomy of the genital organs of 120 specimens from 44 populations representing 16 of the 17 taxa presently listed in
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 Stigmatica and Papillifera were provided by other collectors. As regards the other genera (Mauritanica, Charpentieria, Stigmatica, Gibbularia, and Papillifera), 133 specimens of 46 taxa were anatomically investigated.
Ninety-four specimens comprising Siciliaria, Charpentieria, Gibbularia, Stigmatica, Mauritanica, and Papillifera were genetically analyzed (see Table
Specimens used in the molecular genetic analyses. Collection numbers, laboratory IDs, GenBank accession numbers, and geographical coordinates are listed. More than one ITS2 sequence was obtained from some individuals.
Taxon | Collection No. | Lab ID | GenBank Accession No. | Coordinates | |
---|---|---|---|---|---|
COI | ITS2 | ||||
Sicania crassicostata (L. Pfeiffer, 1856) | WDM Siciliaria 1 | 1_1 | MW758912 | MW757114, MW757113 | 38°05'40.91"N, 12°40'10.98"E |
WDM Siciliaria 1 | 1_4 | MW758924 | no | 38°05'40.91"N, 12°40'10.98"E | |
Mauritanica scarificata (L. |
WDM Siciliaria 2 | 2_1 | MW758915 | MW757112, MW757111 | 37°58'11.08"N, 12°03'59.36"E |
WDM Siciliaria 2 | 2_2 | MW758916 | no | 37°58'11.08"N, 12°03'59.36"E | |
Siciliaria calcarae calcarae ( |
WDM Siciliaria 5 | 5_1 | MW758925 | MW757125, MW757126, MW757127 | 37°57'29.40"N, 12°57'33.64"E |
Sicania nobilis nobilis (L. Pfeiffer, 1848) | WDM Siciliaria 14 | 14_1 | MW758875 | MW757128, MW757129, MW757130 | 38°10'51.85"N, 12°43'37.70"E |
Siciliaria tiberii scalettensis Beckmann, 2004 | WDM Siciliaria 17 | 17_2 | MW758923 | MW757110, MW757109, MW757108 | 38°10'35.53"N, 13°07'27.22"E |
Sicania nobilis nobilis (L. Pfeiffer, 1848) | WDM Siciliaria 18 | 18_1 | MW758918 | MW757107, MW757106 | 38°10'30.07"N, 12°46'14.09"E |
Siciliaria tiberii tiberii (A. Schmidt, 1868) | WDM Siciliaria 19 | 19_1 | MW758919 | MW757117, MW757116 | 38°08'19.06"N, 13°03'14.01"E |
WDM Siciliaria 19 | 19_2 | MW758920 | no | 38°08'19.06"N, 13°03'14.01"E | |
Siciliaria ferrox (Brandt, 1961) | WDM Siciliaria 21 | Sf_1 | MW758922 | MW757115 | 37°59'17.62"N, 13°36'51.11"E |
WDM Siciliaria 21 | Sf_2 | MW758917 | no | 37°59'17.62"N, 13°36'51.11"E | |
Sicania crassicostata (L. Pfeiffer, 1856) | WDM Siciliaria 33 | 33_1 | MW758876 | no | 38°06'44.89"N, 12°40'34.27"E |
WDM Siciliaria 33 | 33_2 | MW758877 | MW757105, MW757104, MW757131, MW757103 | 38°06'44.89"N, 12°40'34.27"E | |
Siciliaria calcarae jatinensis ssp. nov. | WDM Siciliaria 34 | 34_1 | MW758878 | no | 37°58'15.07"N, 13°12'2.23"E |
WDM Siciliaria 34 | 34_2 | MW758879 | MW757118, MW757120, MW757119, MW757121 | 37°58'15.07"N, 13°12'2.23"E | |
WDM Siciliaria 34 | 34_3 | MW758926 | no | 37°58'15.07"N, 13°12'2.23"E | |
WDM Siciliaria 34 | 34_4 | MW758927 | no | 37°58'15.07"N, 13°12'2.23"E | |
Siciliaria calcarae orlandoi |
WDM Siciliaria 36 | 36_1 | MW758921 | MW757132, MW757133 | 37°55'46.42"N, 13°23'5.22"E |
Siciliaria tiberii alcamoensis Brandt, 1961 | WDM Siciliaria 37 | 37_1 | MW758928 | MW757102, MW757101 | 38°07'48.60"N, 13°08'26.94"E |
Siciliaria tiberii armettensis ssp. nov. | WDM Siciliaria 38 | 38_1 | MW758880 | MW757100, MW757099, MW757098 | 38°08'58.56"N, 13°09'25.83"E |
WDM Siciliaria 38 | 38_2 | MW758881 | no | 38°08'58.56"N, 13°09'25.83"E | |
Siciliaria tiberii scalettensis Beckmann, 2004 | WDM Siciliaria 39 | 39_1 | MW758882 | MW757097, MW757096, MW757095 | 38°10'42.45"N, 13°07'34.50"E |
WDM Siciliaria 40 | 40_1 | no | no | 38°10'35.53"N, 13°07'27.22"E | |
Siciliaria grohmanniana addaurae ssp. nov. | WDM Siciiiaria 41 | 41_1 | MW758883 | MW757094, MW757093 | 38°11'13.01"N, 13°21'6.91"E |
WDM Siciiiaria 41 | 41_2 | MW758884 | no | 38°11'13.01"N, 13°21'6.91"E | |
WDM Siciiiaria 41 | 41_3 | MW758929 | no | 38°11'13.01"N, 13°21'6.91"E | |
Siciliaria grohmanniana grohmanniana (Rossmässler, 1836) | WDM Siciliaria 42 | 42_1 | MW758886 | no | 38°10'4.41"N, 13°21'2.60"E |
WDM Siciliaria 42 | 42_2 | MW758887 | MW757092, MW757091 | 38°10'4.41"N, 13°21'2.60"E | |
WDM Siciliaria 42 | 42_3 | MW758885 | no | 38°10'4.41"N, 13°21'2.60"E | |
Siciliaria leucophryna (L. Pfeiffer, 1862) | WDM Siciliaria 43 | 43_1 | MW758888 | no | 38°12'1.32"N, 13°16'3.05"E |
Siciliaria calcarae belliemii (Brandt, 1961) | WDM Siciliaria 44 | 44_1 | MW758889 | MW757122, MW757123 | 38°00'54.80"N, 12°59'13.35"E |
WDM Siciliaria 44 | 44_2 | MW758913 | no | 38°00'54.80"N, 12°59'13.35"E | |
WDM Siciliaria 44 | 44_3 | MW758930 | no | 38°00'54.80"N, 12°59'13.35"E | |
WDM Siciliaria 44 | 44_4 | MW758931 | no | 38°00'54.80"N, 12°59'13.35"E | |
Siciliaria calcarae borgettensis ssp. nov. | WDM Siciliaria 46 | 46_1 | MW758890 | MW757090, MW757089, MW757080 | 38°02'59.53"N, 13°08'58.55"E |
WDM Siciliaria 46 | 46_2 | MW758891 | no | 38°02'59.53"N, 13°08'58.55"E | |
WDM Siciliaria 46 | 46_3 | MW758932 | no | 38°02'59.53"N, 13°08'58.55"E | |
Siciliaria calcarae cruenta ssp. nov. | WDM Siciliaria 47 | 47_1 | MW758933 | MW757079, MW757078 | 38°03'37.03"N, 13°12'38.53"E |
WDM Siciliaria 47 | 47_2 | MW758934 | MW757088, MW757087, MW757077 | 38°03'37.03"N, 13°12'38.53"E | |
WDM Siciliaria 47 | 47_3 | MW758935 | no | 38°03'37.03"N, 13°12'38.53"E | |
WDM Siciliaria 47 | 47_4 | MW758936 | no | 38°03'37.03"N, 13°12'38.53"E | |
Sicania eminens (A. Schmidt, 1868) | WDM Siciliaria 49 | 49_1 | MW758892 | no | 38°04'0.52"N, 12°41'1.34"E |
WDM Siciliaria 49 | 49_2 | MW758893 | no | 38°04'0.52"N, 12°41'1.34"E | |
WDM Siciliaria 50 | 50_1 | MW758894 | MW757076, MW757075, MW757074, MW757073 | 38°04'6.38"N, 12°41'48.68"E | |
WDM Siciliaria 50 | 50_2 | MW758895 | no | 38°04'6.38"N, 12°41'48.68"E | |
WDM Siciliaria 51 | 51_1 | MW758896 | MW757072 | 38°04'55.97"N, 12°40'9.91"E | |
WDM Siciliaria 51 | 51_2 | MW758897 | no | 38°04'55.97"N, 12°40'9.91"E | |
WDM Siciliaria 52 | 52_1 | MW758898 | no | 38°05'18.10"N, 12°40'17.18"E | |
WDM Siciliaria 52 | 52_2 | MW758899 | MW757086, MW757071, MW757085 | 38°05'18.10"N, 12°40'17.18"E | |
Siciliaria calcarae belliemii (Brandt, 1961) | WDM Siciliaria 54 | 54_1 | MW758900 | MW757134, MW757135, MW757136 | 38°00'22.47"N, 13°06'37.79"E |
WDM Siciliaria 54 | 54_2 | MW758901 | no | 38°00'22.47"N, 13°06'37.79"E | |
Siciliaria leucophryna (L. Pfeiffer, 1862) | WDM Siciliaria 55 | 55_1 | MW758902 | MW757124 | 38°11'13.61"N, 13°16'44.68"E |
WDM Siciliaria 55 | 55_2 | MW758903 | no | 38°11'13.61"N, 13°16'44.68"E | |
Siciliaria septemplicata (Philippi, 1836) | WDM Siciliaria 56 | 56_1 | MW758904 | MW757137, MW757138 | 38°12'39.27"N, 13°17'21.26"E |
WDM Siciliaria 57 | 57_1 | MW758937 | MW757070 | 38°13'5.61"N, 13°18'4.76"E | |
Siciliaria calcarae calcarae ( |
WDM Siciliaria 59 | 59_1 | MW758905 | MW757084, MW757069, MW757068 | 38°02'54.18"N, 12°48'26.61"E |
Sicania nobilis nobilis (L. Pfeiffer, 1848) | WDM Siciliaria 60 | 60_1 | MW758906 | MW757067, MW757066, MW757139, MW757065 | 38°06'25.71"N, 12°44'33.37"E |
WDM Siciliaria 60 | 60_2 | MW758907 | no | 38°06'25.71"N, 12°44'33.37"E | |
WDM Siciliaria 60 | 60_4 | MW758938 | no | 38°06'25.71"N, 12°44'33.37"E | |
WDM Siciliaria 60 | 60_5 | MW758939 | no | 38°06'25.71"N, 12°44'33.37"E | |
Siciliaria calcarae calcarae ( |
WDM Siciliaria 61 | 61_1 | MW758908 | MW757140, MW757141 | 38°06'25.71"N, 12°44'33.37"E |
WDM Siciliaria 61 | 61_2 | MW758909 | MW757142, MW757143, MW757144 | 38°06'25.71"N, 12°44'33.37"E | |
Sicania nobilis nobilis (L. Pfeiffer, 1848) | WDM Siciliaria 62 | 62_2 | MW758914 | no | 38°08'48.81"N, 12°44'6.61"E |
WDM Siciliaria 62 | 62_3 | MW758940 | no | 38°08'48.81"N, 12°44'6.61"E | |
Sicania nobilis spezialensis (Nordsieck, 1984) | WDM Siciliaria 63 | 63_1 | MW758910 | MW757040, MW757039 | 38°07'41.78"N, 12°44'9.24"E |
WDM Siciliaria 63 | 63_2 | MW758911 | no | 38°07'41.78"N, 12°44'9.24"E | |
Charpentieria itala lorinae (Gredler, 1869) | WDM Charpentieria 3 | C3_2 | MW758954 | no | 45°50'23.18"N, 10°36'49.06"E |
Charpentieria stenzii cincta (Brumati, 1838) | WDM Charpentieria 2 | C2_2 | MW758953 | MW757050, MW757049 | 46°11'40.51"N, 12°38'14.26"E |
Charpentieria itala itala (G. von Martens, 1824) | WDM Charpentieria 1 | C1_1 | MW758952 | no | 45°24'22.30"N, 11°31'11.93"E |
Charpentieria stenzii letochana (Gredler, 1874) | WDM Charpentieria 19 | C19_1 | MW758957 | no | 46°36'27.42"N, 12°12'20.01"E |
Charpentieria itala baldensis (Strobel, 1851) | WDM Charpentieria 20 | C20_1 | MW758955 | no | 45°44'25.95"N, 10°51'13.69"E |
Charpentieria dyodon dyodon (S. Studer, 1820) | WDM Charpentieria 44 | C44_1 | MW758956 | MW757052, MW757051 | 46°12'29.53"N, 8°12'46.18"E |
WDM Charpentieria 44 | C44_3 | MW758951 | no | 46°12'29.53"N, 8°12'46.18"E | |
Gibbularia gibbula gibbula (Rossmässler, 1836) | Nordsieck 12167 | 12167_1 | MW758947 | no | 41°47'1.22"N, 15°26'39.92"E |
Nordsieck 12167 | 12167_2 | MW758948 | MW757057, MW757056, MW757055 | 41°47'1.22"N, 15°26'39.92"E | |
WDM Gibbularia 64 | G64_1 | MW758958 | MW757048, MW757083, MW757082 | 45°36'8.91"N, 13°46'0.72"E | |
WDM Gibbularia 64 | G64_2 | MW758959 | no | 45°36'8.91"N, 13°46'0.72"E | |
Mauritanica perinni polygyra (O. Boettger, 1879 | SMF335011/1 | SMF 335011_1 | MW758966 | MW757045, MW757044 | 36°22'2.45"N, 10°07'11.81"E |
Papillifera papillaris papillaris (O.F. Müller, 1774) | Nordsieck 11786 | 11786_1 | MW758941 | MW757063, MW757062, MW757064 | 42°01'4.58"N, 12°54'19.30"E |
Nordsieck 11786 | 11786_2 | MW758942 | no | 42°01'4.58"N, 12°54'19.30"E | |
Papillifera solida solida (Draparnaud, 1805) | Nordsieck 12100 | 12100_1 | MW758944 | no | 40°40'1.62"N, 16°37'16.08"E |
Papillifera papillaris transitans (Paulucci, 1878) | Nordsieck 12147 | 12147_1 | MW758945 | no | 38°28'45.91"N, 16°27'48.05"E |
Nordsieck 12147 | 12147_2 | MW758946 | MW757059, MW757058 | 38°28'45.91"N, 16°27'48.05"E | |
Papillifera papillaris affinis (Philippi, 1836) | WDM Papillifera 2 | P2_1 | MW758960 | no | 37°58'2.36"N, 13°11'45.19"E |
WDM Papillifera 2 | P2_2 | MW758961 | no | 37°58'2.36"N, 13°11'45.19"E | |
WDM Papillifera 3 | P3_1 | MW758962 | MW757081, MW757047, MW757046 | 37°57'16.92"N, 12°58'9.06"E | |
WDM Papillifera 4 | P4_1 | MW758963 | no | 37°57'58.18"N, 13°13'14.57"E | |
Stigmatica stigmatica sturmii (L. Pfeiffer, 1848) | Nordsieck 12194 | 12194_1 | MW758949 | MW757054, MW757053 | 40°40'31.60"N, 17°56'40.44"E |
Nordsieck 12194 | 12194_2 | MW758950 | no | 40°40'31.60"N, 17°56'40.44"E | |
Stigmatica vulcanica vulcanica (Benoit, 1860) | Nordsieck 12068 | 12068_2 | MW758943 | MW757061, MW757060 | 39°17'2.08"N, 16°15'30.22"E |
Stigmatica kobeltiana (Küster, 1876) | WDM Stigmatica 11 | St11_1 | MW758965 | no | 39°07'58.01"N, 16°15'01.92"E |
Stigmatica pantocratoris pantocratoris (O. Boettger, 1889) | WDM Stigmatica 9 | St9_2 | MW758964 | MW757042, MW757043, MW757041 | 39°46'18.23"N, 19°50'47.23"E |
The DNA extraction was performed in a sterile room using the QIAmp DNeasy Blood and Tissue Kit (QIAGEN, Hilden, Germany) following the protocol of the manufacturer. A 655 bp long fragment of the mitochondrial (mt) cytochrome c oxidase subunit 1 gene (COI) was amplified using the primers LCO1490_ABOL_Moll_1 (5'-TCAACAAAYCATAARGAYATTGG-3') and HCO2198_ABOL_Moll_1 (5'-TAAACTTCTGGRTGACCAAAAAAYCA-3') (
Using the primers 5.8S_LSU-1fw (5'-CTAGCTGCGAGAATTAATGTGA-3') and 28S_LSU-3rv (5'-ACTTTCCCTCACGGTACTTG-3') (
The ITS2 PCR fragments proved to be length-variable due to insertions/deletion, and most individuals were heterozygous. To avoid unreadable sequences, PCR products were cloned. For this task, additional PCRs were performed using the proofreading Platinum® Taq DNA Polymerase High Fidelity (Invitrogen, Carlsbad, CA, USA). These PCRs were conducted with different PCR conditions: 95 °C for 2 min, 5 cycles (94 °C for 15 s, 60 °C for 30 s, 72 °C for 1 min), 35 cycles (94 °C for 15 s, 50 °C for 30 s, 72 °C for 1 min) and without final elongation. PCR fragments were purified with the QIAquick Gel Extraction Kit (QIAGEN, Hilden, Germany) and cloned with the TOPO-TA© cloning kit (Invitrogen, Carlsbad, CA, USA). Three to five clones per individual were sequenced.
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 (https://www.geneious.com). COI sequences were unambiguously aligned (605 sites, no missing data), while the alignments of ITS2 sequences was performed in Geneious 2.10.3 using the MAFFT algorithm (
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 Mauritanica perinni polygyra) in order to reduce the possibility of being misled by abnormities or single freak specimens (
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, BC+SDBC) were taken using image editing software directly on the high-resolution photographs. The anatomical nomenclature partially follows Nordsieck (1969a, 1972, 2009) and
3.1, 2 Siciliaria/Sicania clausilial apparatus (from hnords.de, modified): 3.1 plicae 3.2 lamellae 3.3, 4 Siciliaria/Sicania anatomy of the genital organs general scheme and (3.5–9) variability of the pseudopapilla within genera Siciliaria, Sicania, Mauritanica, Charpentieria, Stigmatica, Gibbularia and Papillifera 3.5 no (pseudo)papilla 3.6 pseudopapilla 3.7 parapseudopapilla 3.8 hemipapilla 3.9 eupapilla (true penial papilla).
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 FO/V, D/BC+SDBC, BC+SDBC/V, D/BC+SDBC, P+E/V, and E/P. The measurements of both shell and genital organs are always given in mm. The proximal/distal indications refer as seen from the gonads. The subdivision of the genital apparatus in anatomical parts follows
A atrium;
ALPP anterior lower palatal plica;
AUPP anterior upper palatal plica;
BC actual bursa copulatrix;
CP clausiliar plate;
CS clausiliar stalk;
CSP cross section of the penial pseudopapilla;
CSE cross section of the epiphallus;
D diverticulum of the bursa copulatrix;
E epiphallus;
EF epiphallar funnel;
ELP epiphallar longitudinal pleats;
ER epiphallar ring;
ETLP epiphallar transitional longitudinal pleats;
EO epiphallar outlet of the hemipapilla;
ET epiphallar thickening;
FDBC first duct of the bursa copulatrix;
FELP free epiphallar longitudinal plate;
FO free oviduct;
H a aperture height;
H s shell height;
HG hermaphroditic gland;
H s shell height;
IOP internal sculpturing of penis;
IL columellaris or inferior lamella;
L lunella;
OELP occupied epiphallar longitudinal plate;
OS ovispermiduct (second hermaphroditic duct);
P penis;
PC penial complex (P+E);
PCL penial complex length;
PG prostatic gland;
PLL parallelis;
PP penial pseudopapilla;
PPP parapseudopapilla;
PR penial retractor muscle;
PRI principalis;
PUPP posterior upper palatal plica;
SCL subclaustralis;
SCOL subcolumellatis;
SDBC second duct of the bursa copulatrix;
SG sperm groove;
SPL spiralis;
spm specimen(s);
SUL sulcalis or parietalis;
SUT suturalis;
TP transition passage;
V vagina;
VD vas deferens;
VR vaginal retractor;
W a aperture width;
W s shell last whorl width (body whorl).
CWDM malacological collection of Willy De Mattia;
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 Siciliaria (Table
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 mt cytochrome c oxidase subunit 1 gene (COI) and where available for the internal transcribed spacer 2 (ITS2). The descriptions of the shells and the clausiliar apparatus of those taxa that are currently valid (MolluscaBase 2021) were already provided and discussed by
The collection numbers, voucher numbers, GenBank accession numbers and geographical coordinates for the specimens used in the molecular genetic analyses are listed in Table
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 COI dataset between the genera and/or lineages analysed. Mean genetic distances within the genera Siciliaria and Sicania are in bold. Gibbularia 1 = Gibbularia gibbula gibbula from Apricena; Gibbularia 2 = Gibbularia gibbula gibbula from Muggia.
1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | ||
1 | Siciliaria | 13.4 | |||||||||
2 | Sicania | 19.6 | 11.4 | ||||||||
3 | Mauritanica | 20.5 | 17.0 | ||||||||
4 | Stigmatica pantocratis | 21.5 | 18.3 | 17.1 | |||||||
5 | Stigmatica vulcanica | 21.3 | 21.3 | 20.7 | 18.6 | ||||||
6 | Stigmatica sturmii | 22.1 | 19.9 | 19.8 | 19.5 | 22.4 | |||||
7 | Charpentieria itala | 21.7 | 21.4 | 20.8 | 20.7 | 22.5 | 21.9 | ||||
8 | Charpentieria stenzi | 21.3 | 21.8 | 20.2 | 18.6 | 21.0 | 21.9 | 20.7 | |||
9 | Gibbularia 1 | 21.0 | 21.4 | 19.4 | 19.4 | 20.7 | 22.8 | 22.3 | 20.1 | ||
10 | Gibbularia 2 | 23.1 | 23.1 | 22.2 | 20.7 | 22.4 | 23.8 | 22.3 | 21.9 | 20.5 | |
11 | Papillifera | 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 Siciliaria and Sicania calculated from the COI sequences. Intraspecific mean distances are in bold.
1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | ||
1 | Siciliaria calcarae | 7.0 | |||||||
2 | Siciliaria leucophryna | 11.3 | 0.4 | ||||||
3 | Siciliaria tiberii | 14.3 | 14.4 | 0.5 | |||||
4 | Siciliaria ferrox | 17.0 | 16.6 | 17.8 | 0.4 | ||||
5 | Siciliaria septemplicata / grohmanniana | 16.9 | 17.3 | 17.3 | 16.4 | 5.1 | |||
6 | Sicania eminens | 20.8 | 21.3 | 20.8 | 20.8 | 19.7 | 2.8 | ||
7 | Sicania crassicostata | 21.4 | 20.3 | 21.3 | 20.2 | 20.2 | 12.2 | 2.5 | |
8 | Sicania nobilis | 20.9 | 20.5 | 21.6 | 19.7 | 18.9 | 13.3 | 12.5 | 4.3 |
The ITS2 sequences were obtained from 42 individuals (Table
The tree based on the combined data set (COI plus ITS2; 45 individuals) is shown in Fig.
Specimens from the same locality delivered very similar or identical sequences. Intraspecific distances (summarised in Table
The taxa comprising the two main clades I (Siciliaria) and II (Sicania) cluster in a geographic pattern (maps in Fig.
Siciliaria leucophryna shows a narrow distribution, limited to Isola delle Femmine, Pizzo Mollica, and Pizzo Manolfo. Siciliaria tiberii is restricted to the calcareous massif of Monte Palmeto, Pizzo di Mezzo, Monte Pecoraro and Montagna Longa and its northwestern coastline. Siciliaria ferrox occurs in the Trabia-S. Onofrio area. The closely related Siciliaria grohmanniana and Siciliaria septemplicata are indeed geographically close, the former occurring exclusively along the southern slopes of Monte Gallo and on Monte Pellegrino and the latter found along the northern slopes of Monte Gallo and scattered localities south of Palermo (
Siciliaria/Sicania taxa show a remarkable adaptation and capability to colonise not only dry, open and exposed limestone walls and cliffs but also a huge variety of habitats and ecological niches. The most stenoecious species are, in order, Sicania nobilis, Siciliaria ferrox, and Siciliaria septemplicata, which are found exclusively on limestone walls or limestone boulders. Sicania crassicostata is found on limestone walls or under rocks. More euryoecious habits were recorded for Siciliaria grohmanniana grohmanniana, which is found on exposed limestone walls hiding in cracks and under crags. It is also found in Pinus sp. forest on decaying woods and tree trunks, including shady and humid spots on mosses and ferns. Interestingly, its subspecies Siciliaria grohmanniana addaurae ssp. nov. is found exclusively on limestone walls. Siciliaria leucophryna and Sicania eminens colonise limestone cliffs hiding in cracks or under stones. They are very common on isolated boulders and tree trunks and barks in the xerophilic scrub. The most euryoecious species is Siciliaria calcarae, which colonises all the habitats until now described but also is commonly found in anthropogenic environments as stone walls, road walls, ruins and disturbed habitats.
The high number of Siciliaria/Sicania specimens dissected during this study (n = 120) provided an accurate overall description of the genital organs of these genera.
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 Siciliaria/Sicania. The relative dimensions (ratio between the shell height and the length of the penial complex Hs/PCL×100) do not substantially differ among taxa, ranging from 25% to 35%, revealing a great dimension stability of genital organs in relationship to the shell dimensions. The genus Montenegrina (
The hermaphroditic gland (HG) has many consolidated acini and it is found along the two apical whorls of the animal’s body. The first hermaphroditic duct (FHD) is very thin along its proximal portion but abruptly turns into a highly coiled, skin-like structure. The diameter of the FHD is much larger along the coiled part, and the number of whorls ranges from six to eleven. The distal uncoiled part of the FHD is very thin, straight and ends directly in the fertilisation chamber-spermatheca complex. The albumen gland (AG) is slender. The general scheme of the FHD is stable among all the Siciliaria taxa. The AG is very variable in size compared to the ovispermiduct (OS). In full-grown adult specimens, it can be larger or as long as the OS or barely reaching 1/4 of its length. The OS consists of a female and a male part including the seminal groove, which is not visible from outside. Externally, it is irregularly spaced by smooth constrictions and it is whitish to cream in colour. The male, prostatic part consists of the visible prostatic gland (PG) and the sperm groove (SG), light to dark grey in colour and clearly distinguishable from the lighter female part. The overall size of the OS is very stable among the Siciliaria/Sicania taxa. The free oviduct (FO) is usually thin and long and its length is stable in relation to the length of the vagina (V), and its average ratio (FO/V) is ~ 1. The V is simple, usually cylindrical, without any appendix or caecum. The vaginal retractor (VR) is strong; it is always present and attached to the second duct of the bursa copulatrix (SDBC). It was not depicted in the anatomical drawing due to its low taxonomic value. The V is stable in length in relation both to shell height and penis length. The ratio between the length of the penial complex (PC) and the length of the V (PC/V) is also stable, ranging from two to three. The BC+SDBC complex (CBC) consists of the FDBC, a second duct (SDBC), a well-developed D (diverticulum) and the actual BC+SDBC. The D is always longer than the SDBC+BC. The transitional section between the BC and the SDBC can range from clearly visible to completely missing (with all degrees of transitional states). The PC comprises the penis (P) and the epiphallus (E). The transition between P and E does not always show the epiphallar thickening (ET), as stated in
In contrast to the external morphology of the genital organs, the internal sculpturing differs between the various taxa of Siciliaria and Sicania, but is stable within the taxa, with only minor population/individual variability. The penis shows the highest variability, which involves the combination (or the absence) of pleats arranged in a longitudinal, transversal, or oblique direction or creating irregular patterns. The epiphallus shows different types of sculpturing, with longitudinal pleats or without any structure. The internal transition between the epiphallus and the vas deferens is very clear due to the clear change of the sculpturing. The structure and the variability of the penial pseudopapilla and the morphology of the internal transition between penis and epiphallus will be discussed in detail in the following. The atrium can exhibit different configurations, from completely smooth to variable pleats. In contrast to what was found in Montenegrina (
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 Siciliaria calcarae calcarae (Philippi, 1844) and Mauritanica perinni polygyra (O. Boettger, 1879) are described for the first time. The spermatophores are slender and slightly longer than the diverticulum. The overall shape is slightly to markedly bent. These preliminary observations revealed a difference between Siciliaria with both upper and lower keel and Mauritanica with only the upper keel.
All the genera Siciliaria, Sicania, Mauritanica, Stigmatica, Gibbularia, and Papillifera exhibit a pseudopapilla. The pseudopapilla (PP) is a “false” penial papilla, found at the transition between the penis and the epiphallus, slightly distally to the insertion of the retractor muscle. The pseudopapilla, conversely to the actual penial papilla [as in the genus Montenegrina, depicted in
In Siciliaria/Sicania one to three strong epiphallar rings are always present (only one being the origin of the pseudopapilla). Charpentieria and Stigmatica present only one epiphallar ring which is the origin of the pseudopapilla. In Gibbularia the epiphallar ring was found only in Gibbularia gibbula cf. sanctangeli (Fig.
In the Southern Alpine Charpentieria the genital investigations revealed two undescribed structures of the transition between P and E: in C. dyodon s. l. it is named “parapseudopapilla” (PPP) and in C. stenzii s. l. it is named “hemipapilla” (HP) and are described in the following sections. In order to describe and classify the huge variety of PP-ER-ELP combinations found in all the taxa, we created a pseudopapillar formula. This formula will be provided for each taxon and population dissected, in order to highlight the extreme variable situation of this part of the genital organs. In this formula the parentheses () mean direct connection (continuity/contiguity of anatomical traits), whereas the plus + means interruption of the anatomical continuity. For example, the formula 1ER(PP)+2ER(ELP) means that PP originates from the distal (first) ER and the proximal ER (second ER) is directly connected with the ELP.
Shell. Shell elongated, fusiform, sinistral, can be either decollate or non-decollate, with or without well-developed white surface layer. Apex subacute when present, whitish to dark brick red in colour. Protoconch smooth. Spire slowly and regularly growing with 9.5–11.5 whorls. Umbilicus closed, sutures shallow. Aperture subovoid to subquadrangular. Peristome continues to be interrupted. Average shell height 19.9 ± 2.1 mm (n = 630, decollate and not decollate shells), average last whorl width 4.5 ± 0.4 mm (n = 630), average aperture height 4.6 ± 0.5 mm (n = 630), average aperture width 2.9 ± 0.4 mm (n = 630). Whorls weakly rib-striated, rib-striated, ribbed to percostate without or with evident to barely recognisable sutural papillae. Dorsal keel missing, mostly indistinct to prominent. Lunella always present, dorsal to dorso-lateral, more or less developed. Inferior lamella low, moderately high, high to very high. Missing, one or two anterior upper palatal plica; when two are present: upper one mostly separated from upper palatal plica; when one is present: separated from or connected with upper posterior palatal plica. Rarely, lower anterior upper palatal plica is present, separated from or connected with upper palatal plica. ALPP tending to be shortened to almost missing. Spiralis overlapping or not overlapping parietalis. Posterior lower palatal plica missing more or less reduced. Palatal edge of clausilium plate distally receding, slightly or not receding. Clausilium palatal edge against distal end not bent upwards, or bent upwards to strongly bent upwards.
External distal part of the genital organs. Free oviduct slightly shorter to remarkably longer than the vagina. Vas deferens thin. First duct of the bursa copulatrix slightly shorter to much longer than second duct of bursa copulatrix + the actual bursa. Distinction between the actual bursa and second duct of bursa copulatrix clear to missing. Diverticulum of the bursa copulatrix longer to much longer than the vagina and second duct of bursa copulatrix + the actual bursa. The vagina is long or short and the atrium usually large. Penial complex always longer to much longer than the vagina. Retractor muscle short to long, usually robust. Epiphallar thickening present or missing. Epiphallus slightly shorter to much longer than the penis. Transition between epiphallus and vas deferens clearly visible to totally indistinguishable.
Internal distal part of the genital organs. Vagina and penis with internal sculpturing comprising a combination (or also absence) of pleats of different size and texture, arranged in a longitudinal, transversal or oblique direction or creating particular net-like pattern. Atrium simple or pleated. Pseudopapilla originating from epiphallar wall or epiphallar ring, elongated to roundish, with smooth or coarse surface. Epiphallar rings up to three to completely missing. Proximal epiphallar ring connected or not with the epiphallar pleats. Epiphallus with or without longitudinal pleats, both smooth or fringed. Epiphallar walls almost smooth, finely granulated of with small transverse papillae.
The alpha-taxonomy of Siciliaria/Sicania was based on shell morphology. However, the variability of the shell characters reflects the complicate nomenclatural history (
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 Montenegrina (
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 COI tree the species were monophyletic except the widely distributed S. calcarae. Clustering of subspecies is mentioned in the taxonomic part, albeit this aspect is of minor relevance, (1) because subspecies would not necessarily be expected to be monophyletic and (2) because they were represented by too small samples.
Finally,
Clausilia subdiaphana was introduced by
The status of Siciliaria riberothi, introduced by
The concept of interspecies hybridisation (or geographic forms) introduced by Nordsieck (
The list of the examined specimens of Siciliaria and Sicania is depicted in Table
We isolated and depicted the clausiliar plates in order to verify the morphology described by
Siciliaria/Sicania taxa examined (anatomy and/or molecular genetic data) with description of the variable penis-epiphallus transition: epiphallar formula. DNA = molecular genetic data available (Yes/No).
Taxon | Locality | DNA | Epiphallar formula |
---|---|---|---|
Siciliaria grohmanniana grohmanniana (Rossmässler, 1836) | Italy, Sicily, Palermo, Monte Pellegrino, N side of the top plateau, 380 m asl, 38°10'55.26"N, 13°21'1.66"E, leg. W. De Mattia and J. Macor leg. | Y | 1ER(PP+ELP) |
Italy, Sicily, Palermo, Monte Pellegrino, Santuario Santa Rosalia, 420 m asl, 38°10'4.41"N, 13°21'2.60"E, leg. W. De Mattia and J. Macor leg. | Y | 1ER(PP+ELP) | |
Siciliaria grohmanniana addaurae ssp. nov. De Mattia, Reier & Haring | Italy, Sicily, Palermo, N side of Monte Pellegrino, Punta Priola, Grotte dell’Addaura, 115 m asl, 38°11'13.01"N, 13°21'6.91"E, W. De Mattia and J. Macor leg. | Y | 1ER(PP)+ELP |
Siciliaria septemplicata (Philippi, 1836) | Italy, Sicily, Palermo, N side of Monte Gallo near Sferracavallo, 40 m asl, 38°12'39.27"N, 13°17'21.26"E, W. De Mattia and J. Macor leg. | Y | 1ER+2ER(PP+ELP) |
Italy, Sicily, Palermo, E side of Monte Gallo, 140 m asl, 38°13'5.61"N, 13°18'4.76"E, W. De Mattia and J. Macor leg. | Y | 1ER+2ER(PP+ELP) | |
Siciliaria leucophryna (L. Pfeiffer, 1862) | Italy, Sicily, Palermo, Grotta Conza, 150 m asl, 38°11'13.61"N, 13°16'44.68"E, W. De Mattia and J. Macor leg. | Y | 1ER+2ER(PP+ELP) |
Italy, Sicily, Palermo, Sferracavallo, via Plauto, 50 m asl, 38°11'13.01"N, 13°21'6.91"E, W. De Mattia and J. Macor leg. | Y | 1ER(PP)+ELP | |
Siciliaria calcarae calcarae (Philippi, 1844) | Italy, Sicily, Alcamo, Monte Bonifato, top of the mountain, 640 m asl, 37°57'29.40"N, 12°57'33.64"E, A. Margelli leg. | N | 1ER(PP)+ELP |
Italy, Sicily, Alcamo, Monte Bonifato, top of the mountain, 640 m asl, 37°57'29.40"N, 12°57'33.64"E, I. Niero leg. | Y | 1ER(PP+ELP) | |
Italy, Sicily, Alcamo, Monte Bonifato, west side of the mountain, over the quarry, 550 m asl, 37°57'16.92"N, 12°58'9.06"E, W. De Mattia and J. Macor leg. | N | 1ER(PP+ELP) | |
Italy, Sicily, Piana degli Albanesi, 500 m south of Portella Ginestra, northern cliffs of Monte Kumeta, 970 m asl, 37°58'13.35"N, 13°15'22.06"E, W. De Mattia and J. Macor leg. | N | 1ER(PP)+2ER(ELP) | |
Italy, Sicily, Castellammare del Golfo, Castello di Baida, W of the town along the road to Visicari, 300 m asl, 38° 02'41.64"N, 12°48'14.34"E, W. De Mattia and J. Macor leg. | N | 1ER(PP)+2ER(ELP) | |
Italy, Sicily, San Vito lo Capo, Castelluzzo, west cliffs E of town, 120 m asl, 38°06'25.71"N, 12°44'33.37"E, W. De Mattia and J. Macor leg. | Y | 1ER(PP)+ELP | |
Italy, Sicily, Castellammare del Golfo, Visicari, 395 m asl, 38°02'54.18"N, 12°48'26.61"E, W. De Mattia and J. Macor leg. | Y | 1ER(PP+ELP) | |
Siciliaria calcarae calcarae (Philippi, 1844) (= adelina Küster, 1847) | Italy, Sicily, Calatafimi, Castello Eufemio, 395 m asl, 37°57'45.67"N, 12°51'21.13"E, W. De Mattia and J. Macor leg. | N | 1ER+2ER(PP)+ELP |
Siciliaria calcarae belliemii (Brandt, 1961) | Italy, Sicily, Partinico, W side of the Mount Belliemi, 440 m asl, 38°00'22.47"N, 13°06'37.79"E, W. De Mattia and J. Macor leg. | Y | 1ER+2ER(PP)+ELP |
Italy, Sicily, Alcamo, E side of the Calatubo Castle, 75 m asl, 38°00'54.80"N, 12°59'13.35"E, W. De Mattia and J. Macor leg. | Y | 1ER+2ER(PP+ELP) | |
Siciliaria calcarae borgettensis ssp. nov. De Mattia, Reier & Haring | Italy, Sicily, Municipality of Borghetto, road to Romitello, 400 m asl, 38°02'59.53"N, 13°08'58.55"E, W. De Mattia and J. Macor leg. | Y | 1ER(PP)+ELP |
Italy, Sicily, San Giuseppe Jato, quarry E of the town, 630 m asl, 37°58'15.07"N, 13°12'2.23"E, W. De Mattia and J. Macor leg. | Y | 1ER+2ER(PP+ELP) | |
Siciliaria calcarae parajatinensis ssp. nov. De Mattia, Reier & Haring | Italy, Sicily, Monreale, W part of Monte Kumeta toward Jato Antica, 630 m asl, 37°57'8.18"N, 13°13'14.57"E, W. De Mattia and J. Macor leg., | Y | 1ER+2ER(PP+ELP) |
Siciliaria calcarae orlandoi Liberto, Reitano, Giglio, Colomba & Sparacio, 2016 | Italy, Sicily, Monreale, Bosco Ficuzza, Ponte Arcere, 470 m asl, 37°55'46.42"N, 13°23'5.22"E, W. De Mattia and J. Macor leg. | Y | 1ER+2ER(PP+ELP) |
Siciliaria calcarae cruenta ssp. nov. De Mattia, Reier & Haring | Italy, Sicily, Monreale, N side of Monte Gibilmesi, 890 m asl, 38°03'37.03"N, 13°12'38.53"E, W. De Mattia and J. Macor leg. | Y | 1ER+2ER(PP)+ELP |
Siciliaria ferrox Brandt, 1961 | Italy, Sicily, Trabia, Contrada Sant’Onofrio, 170 m asl, 37°59'17.62"N, 13°36'51.11"E, W. De Mattia and J. Macor leg. | Y | ?ER+PP(ELP) |
Siliciaria tiberi tiberi (A. Schmidt, 1868) | Italy, Sicily, Terrasini, Capo Rama, 30 m asl, 38°08'19.06"N, 13°03'14.01"E, W. De Mattia and J. Macor leg. | Y | ER+PP(ELP) |
Siciliaria tiberii alcamoensis Brandt, 1961 comb. nov. | Italy, Sicily, Cinisi, Piano Margi, 670 m asl, 38°08'58.56"N, 13°09'25.83"E, W. De Mattia and J. Macor leg. | Y | 1ER(PP+ELP) |
Siciliaria tiberii armettensis ssp. nov. De Mattia, Reier and Haring | Italy, Sicily, Carini, Grotta dei Puntali o dell’Armetta, 90 m asl, 38°08'58.56"N, 13°09'25.83"E, W. De Mattia and J. Macor leg. | Y | 1ER+2ER+3ER(PP+ELP) |
Siciliaria tiberii scalettensis Beckmann, 2004 | Italy, Sicily, Cinisi, Portella Scaletta [locus typicus], 90 m asl, 38°10'35.53"N, 13°07'27.22"E, W. De Mattia and J. Macor leg. | Y | 1ER+PP(ELP) |
Italy, Sicily, Cinisi, along the SS113 road from Villagrazia di Carini to Cinisi, 70 m asl, 38°10'42.45"N, 13°07'34.50"E, W. De Mattia and J. Macor leg. | Y | 1ER+PP(ELP) | |
Sicania crassicostata (Pfeiffer, 1856) | Italy, Sicily, Castelluzzo, cliffs W of the Tonnara di Monte Cofano, 50 m asl, 38°6'44.89"N, 12°40'34.27"E, W. De Mattia and J. Macor leg. | Y | 1ER+PP(ELP) |
Italy, Sicily, Custonaci, cliffs N of the Mangiapane cave, 63 m asl, 38°05'40.91"N, 12°40'10.98"E, W. De Mattia and J. Macor leg. | Y | 1ER(PP)+ELP | |
Sicania eminens (A. Schmidt, 1868), comb. nov. | Italy, Sicily, Custonaci, Baglio Messina, 340 m asl, 38° 4'06.38"N, 12°41'48.68"E, W. De Mattia and J. Macor leg., | Y | 1ER+2ER(PP+ELP) |
Italy, Sicily, Custonaci, Buffara, NE side of the hollow, 150 m asl, 38°04'0.52"N, 12°41'1.34"E, W. De Mattia and J. Macor leg. | Y | 1ER(PP+ELP) | |
Italy, Sicily, Custonaci, contrada Scurati, crossing with Strada Provinciale 18, 65 m asl, 38°04'56.04"N, 12°40'9.86"E, W. De Mattia and J. Macor leg. | Y | 1ER(PP)+2ER(ELP) | |
Italy, Sicily, Custonaci, contrada Scurati, 60 m asl, 38° 05'18.10"N, 12°40'17.18"E, W. De Mattia and J. Macor leg. | Y | 1ER(PP)+2ER(ELP) | |
Italy, Sicily, Custonaci, contrada Scurati, limestone cliffs S of the village, 60 m asl, 38°06'44.89"N, 12°40'34.27"E, W. De Mattia and J. Macor leg. | Y | 1ER(PP)+2ER(ELP) | |
Sicania nobilis nobilis (L. Pfeiffer, 1848), comb. nov. | Italy, Sicily, Castelluzzo, Monte Cofano E of Tonnara di Monte Cofano, 80 m asl, 38°06'22.04"N, 12°40'59.77"E, W. De Mattia and J. Macor leg. | N | 1ER+2ER+3ER(PP+ELP) |
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(PP+ELP) | |
Italy, Sicily, San Vito lo Capo, Castelluzzo, west cliffs E of town, 120 m asl, 38°06'25.71"N, 12°44'33.37"E, W. De Mattia and J. Macor leg. | Y | 1ER+2ER(PP)+ELP | |
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(PP)+ELP | |
Sicania nobilis nobilis (L. Pfeiffer, 1848), comb. nov. (= episoma Brandt, 1961) | Italy, Sicily, San Vito lo Capo, Torre delle Usciere, 5 m asl, 38°10'30.07"N, 12°46'14.09"E, W. De Mattia and J. Macor leg. | Y | 1ER+2ER+3ER(PP+ELP) |
Sicania nobilis spezialensis ( |
Italy, Sicily, San Vito Lo Capo, Macari, E side of Monte Speziale, 75 m asl, 38°07'41.78"N, 12°44'09.24"E, W. De Mattia and J. Macor leg. | Y | 1ER+2ER(PP+ELP) |
Distribution of the clausilium character states within the Siciliaria/Sicania taxa. n = number of specimens analysed.
Taxon | n | Boettger’s groups: Gruppe der septemplicata / Gruppe der crassicostata | Clausiliar plate distally receding/not receding | Clausiliar plate gutter-like/not gutter like | Clausiliar plate bent upwards/not bent upwards |
---|---|---|---|---|---|
Siciliaria grohmanniana grohmanniana (Rossmässler, 1836) | 22 | septemplicata | receding | gutter-like | bent |
Siciliaria grohmanniana addaurae ssp. nov. | 18 | septemplicata | receding | gutter-like | bent |
Siciliaria septemplicata (Philippi, 1836) | 15 | septemplicata | receding | gutter-like | bent |
Siciliaria leucophryna (L. Pfeiffer, 1862) | 20 | septemplicata | not receding | not gutter-like | not bent |
Siciliaria calcarae calcarae ( |
36 | septemplicata | not receding | not gutter-like | bent |
Siciliaria calcarae belliemii (Brandt, 1961) | 25 | septemplicata | not receding | not gutter-like | bent |
Siciliaria calcarae borgettensis ssp. nov. | 12 | septemplicata | not receding | not gutter-like | bent |
Siciliaria calcarae jatinensis ssp. nov. | 21 | septemplicata | not receding | not gutter-like | bent |
Siciliaria calcarae parajatinensis ssp. nov. | 15 | septemplicata | not receding | not gutter-like | bent |
Siciliaria calcarae orlandoi Liberto, Reitano, Giglio, Colomba & Sparacio, 2016 | 9 | septemplicata | partially receding | gutter-like | bent |
Siciliaria calcarae cruenta ssp. nov. | 7 | septemplicata | not receding | not gutter-like | bent |
Siciliaria ferrox (Brandt, 1961) | 11 | septemplicata | partially receding | partially gutter-like | not bent |
Siciliaria tiberii tiberii (A. Schmidt, 1868) | 31 | septemplicata | not receding | not gutter-like | bent |
Siciliaria tiberii armettensis ssp. nov. | 33 | septemplicata | not receding | not gutter-like | bent |
Siciliaria tiberii hemmeni (Beckmann, 2004), comb. nov. | 6 | septemplicata | not receding | not gutter-like | not bent |
Siciliaria tiberii scalettensis Beckmann, 2004 | 27 | septemplicata | not receding | not gutter-like | not bent |
Sicania crassicostata (L. Pfeiffer, 1856), comb. nov. | 19 | crassicostata | not receding | not gutter-like | not bent |
Sicania eminens (A. Schmidt, 1868), comb. nov. | 29 | crassicostata | not receding | not gutter-like | not bent |
Sicania nobilis nobilis (L. Pfeiffer, 1848), comb. nov. | 35 | crassicostata | not receding | not gutter-like | not bent |
Sicania nobilis spezialensis (Nordsieck, 1984), comb. nov., stat. nov. | 21 | crassicostata | not receding | not gutter-like | not bent |
Character | Siciliaria | Sicania |
Shell – waxy surface | not present | only in Sicania nobilis nobilis |
Shell – percostate | not present | only in Sicania crassicostata |
Shell – clausiliar plate gutter-like | only in S. grohmanniana s. l. and S. septemplicata | not present |
Shell – clausiliar plate receding | only in S. grohmanniana s. l. and S. septemplicata | not present |
Ganitalia – penial internal papillose sculpturing | not present | only in Sicania nobilis nobilis |
Infraordo Clausilioidei Gray, 1855
Superfamilia Clausilioidea Gray, 1855
Familia Clausiliidae Gray, 1855
Subfamilia Alopiinae A.J. Wagner, 1913
Siciliaria grohmanniana forms a subclade with its sister group Siciliaria septemplicata (mean p distance 9.0%). The two subspecies of S. grohmanniana appear monophyletic in the mt tree (Fig.
The shell of Siciliaria grohmanniana addaurae ssp. nov. was at first considered by
Clausilia grohmanniana Rossmässler 1836: 7.
Clausilia grohmanniana var. minor
A.
Clausilia grohmanuiana
[sic!] –
Clausilia grohmaniana
[sic!] –
Clausilia (Siciliaria) grohmanniana
–
Delima (Siciliaria) grohmanniana
–
Siciliaria grohmanniana
–
Charpentieria grohmanniana
–
Siciliaria grohmanniana
–
Siciliaria grohmanniana
–
Siciliaria grohmanniana
–
Charpentieria grohmanniana
–
Italy, Sicily, Palermo, Monte Pellegrino, N side of the top plateau, 380 m asl, 38°10'55.26"N, 13°21'1.66"E, W. De Mattia and J. Macor leg., 20.xii.2003. 15 live spm, 3 dissected spm. Italy, Sicily, Palermo, Monte Pellegrino, Santuario Santa Rosalia, 420 m asl, 38°10'4.41"N, 13°21'2.60"E, [Lab ID 42_1, COI: MW758886; Lab ID 42_2, COI: MW758887, ITS2: MW757091 MW757092; Lab ID 42_3, COI: MW758885], W. De Mattia and J. Macor leg., 15.iv.2017. 12 live spm, 3 dissected spm.
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 FO is longer than the V (FO/V range from 1.6 to 1.8). The VD is thin along its whole course. The FDBC is shorter than the BC+SDBC (FDBC/BC+SDBC range from 0.7 to 0.8). The BC+SDBC is cylindrical or club-like and slightly longer than the V (BC+SDBC/V range 1.3–1.4), with no clear distinction between the SDBC and the BC. The apex is wide and round or pointed. The D is longer than the V (D/V range 2.0–2.1) and longer that the BC+SDBC (D/BC+SDBC range 1.4–1.7), thinner than the BC+SDBC and with a pointed apex. The V is short and cylindrical. The A is large. The PC is longer than the V (P+E/V range 2.4–2.6). The PR is long and robust. There is a clear distinction between P and E as there is a visible ER and a proximal narrowing. The E is thinner but longer than the P (E/P range 1.1–1.3), almost gradually shrinking and turning into the VD.
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 PP is big, irregular, wrinkly and pointed. It can be smooth or with small tubercles. The P-E transition presents one ER with the PP and ELP originating from the ER. The epiphallar formula is: 1ER(PP+ELP). The E shows 4 to 6 main longitudinal finely fringed pleats. The V is almost completely smooth. The wall is finely granulated.
Siciliaria grohmanniana grohmanniana is widespread and common throughout its range. It is found to live on exposed limestone walls hiding in cracks and under crags. It is also found in Pinus sp. forest on decaying woods and tree trunks, including shady and humid spots on mosses and ferns. According to
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, 38°11'13.01"N, 13°21'6.91"E.
1 Holotype (
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 PRI is long and raised, wider along its posterior part and not fused with the L. The PRI is not visible from the aperture. The L is antero-lateral, with a reduced, knob-like PUPP connected to it. There are two AUPP, both of them detached from the L and barely visible from the aperture. The upper AUPP is stronger and longer than the lower one. The ALPP (BAS) starts directly from the L and it is long and strong, clearly visible from the aperture. The SCL is present and robust. It is connected to the L. The IL is high to very high. The SUL is short, tooth-like and ends before the beginning of the SPL that is wider along its posterior part. The SCOL is not emergent. The peristome is continuous, markedly thickened and reflected. It is not superiorly fused to the wall of the first whorl. The palatal edge of the clausilium is distally receding and bent upwards. The plate is narrow and gutter-like.
Siciliaria grohmanniana grohmanniana (Rossmässler, 1836), Monte Pellegrino, Palermo 7.1 whole distal genital organs 7.2 internal distal part of genital organs 7.3 penial pseudopapilla detail. Siciliaria grohmanniana addaurae ssp. nov. Grotta dell’Addaura, Punta Priola 7.4 whole distal genital organs 7.5 internal distal part of genital organs 7.6 penial pseudopapilla detail.
Siciliaria grohmanniana grohmanniana (Rossmässler, 1836), Monte Pellegrino, Palermo 8.1 shell 8.2 shell 8.3 detail of the aperture 8.4 clausiliar plate double side. Siciliaria grohmanniana addaurae ssp. nov. Grotta dell’Addaura, Punta Priola 8.5 shell 8.6 shell 8.7 detail of the aperture 8.8 detail of the columellar side of last whorl 8.9 clausiliar plate double side.
Holotype : decollate shell height 17.3, whorl width 4.5, aperture height 4, aperture width 2.5. Paratypes (n = 20, decollate): shell height 17.2 ± 0.6, whorl width 4.4 ± 0.1, aperture height 3.9 ± 0.3, aperture width 2.9 ± 0.3.
The FO is longer than the V (FO/V range 1.5–1.7). The VD is thin along its whole course. The FDBC is longer than the BC+SDBC (FDBC/BC+SDBC range 1.0–1.3). The BC+SDBC is cylindrical or club-like and slightly longer than the V (BC+SDBC/V = 1.1), with no clear distinction between the SDBC and the BC. The apex is wide and round. The D is longer than the V (D/V range 1.3–1.5) and longer that the BC+SDBC (D/BC+SDBC range 1.1–1.3), thinner than the BC+SDBC and with a small apex. The V is long and cylindrical and thin in diameter. The A is small. The PC is longer than the V (P+E/V range 1.6–1.8). The PR is long and thin. There is a clear distinction between P and E as there is a visible ER and a proximal narrowing. The E is much thinner but longer than the P (E/P range 1.2–1.5), gradually shrinking and turning into the vas deferens.
The A is smooth. The P presents very smooth and scarcely elevated weak transverse pleats. The fine structure of the wall is smooth. The PP is big, rounded and smooth. It originates from the big epiphallar ring. The P-E transition presents one ER with the PP originating from it. The ELP are not connected with the ER. The epiphallar formula is: 1ER(PP)+ELP. The E shows three or four main smooth longitudinal pleats. The proximal V shows an irregular set of smooth longitudinal pleats that, distally, turn into a smooth fold that goes as far as the atrium.
Siciliaria grohmanniana addaurae ssp. nov. differs from Siciliaria grohmanniana grohmanniana by its slender shape and darker shell colour and its denser ribbing (Fig.
Siciliaria grohmanniana addaurae ssp. nov. is exclusively known from the type locality, along the limestone cliffs around the Addaura caves near Punta Priola, Palermo. It is likely to be present in other spots along the northeastern limestone cliffs of the Monte Pellegrino, but more research is needed to define its distribution.
Siciliaria grohmanniana addaurae ssp. nov. is an obligate rock-dweller and inhabits the limestone cliffs, on open walls or hiding in cracks in humid spots, around the Addaura caves. This subspecies has a very limited distribution range with an area of much less than 1 km2. Although, the area is included in the Riserva Naturale Orientata Monte Pellegrino and the Addaura caves are fenced, the habitat quality is inferred to be declining due to no access regulation nor restriction, resulting in dumping, littering and murals on the limestone walls (
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.
Clausilia septemplicata
Clausilia sericina Rossmässler 1836: 7.
Clausilia septemplicata
–
Clausilia septemplicata var. prasina
A.
Clausilia septemplicata
–
Clausilia prasina
Clausilia septemplicata – Monterosato 1882: 102.
Clausilia (Siciliaria) septemplicata
–
Delima (Siciliaria) septemplicata prasina – Wagner 1913: plate 572, fig. 14.
Charpentieria septemplicata
–
Siciliaria septemplicata
–
Siciliaria septemplicata
–
Charpentieria septemplicata
–
The case of Siciliaria septemplicata and its alleged subspecies (“geographic forms” following
Italy, Sicily, Palermo, N side of Monte Gallo near Sferracavallo, 40 m asl, 38°12'39.27"N, 13°17'21.26"E, [Lab ID 56_1, COI: MW758904, ITS2: MW757137, MW757138], W. De Mattia and J. Macor leg., 16.iv.2017. 5 live spm, 2 dissected spm. Italy, Sicily, Palermo, E side of Monte Gallo, 140 m asl, 38°13'5.61"N, 13°18'4.76"E, [Lab ID 57_1, COI: MW758937, ITS2: MW757070], W. De Mattia and J. Macor leg., 16.iv.2017. 4 live spm, 2 dissected spm.
(Figs
(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 FO is longer than the V (FO/V range from 1.1 to 3.9). The VD is thin along its whole course. The FDBC is slightly longer than the BC+SDBC (FDBC/BC+SDBC range 1.1–1.9). The BC+SDBC is cylindrical to spindle-like and longer than the V (BC+SDBC/V range 1.1–1.3), with no clear distinction between the SDBC and the actual BC. The apex is big and pointed. The D is longer than the V (D/V = 1.9) and longer that the BC+SDBC (D/BC+SDBC range 1.6–1.7), thinner than the BC+SDBC and with a small and round apex. The V is cylindrical. The A is large. The PC is longer than the V (P+E/V range 2.0–2.4). The PR is long and thin or short and strong. The E is as long as the P (E/P = 1.0), gradually shrinking and turning into the VD.
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 PP is small to medium in dimensions and elongated. The P-E transition presents a first distal ER, PP and ELP originate from the second proximal ER. The epiphallar formula is: 1ER+2ER (PP+ELP). The E shows one main longitudinal slightly segmented pleat that fades out before the VD. Few additional small, weak, elevated pleats can also be present. The wall of the E is coarsely granulated. The V presents a set of irregular fleshy folds.
Siciliaria septemplicata was found dwelling on limestone cliffs, hiding in cracks in exposed or shady spots. According to
Following the new taxonomical results, the distribution of Siciliaria septemplicata, must be reconsidered.
Clausilia leucophryna
Siciliaria leucophryne
[sic!] –
Clausilia leucophryne var. laudabilis
[sic!]
Clausilia lencophryna
[sic!] –
Clausilia confinata merens
Clausilia leucophryne laudabilis
[sic!] –
Charpentieria leucophryna
–
Siciliaria leucophryna
–
Siciliaria leucophryna – Liberto et al. 2012: 560.
Siciliaria leucophryna
–
Siciliaria leucophryna
–
Charpentieria leucophryna
–
Siciliaria leucophryna microinsularis
Siciliaria leucophryna in the mt tree forms a separate subclade within the big S. calcarae clade (mean p distance 11.3%). As depicted in the combined tree (Fig.
The population from Via Plauto, concerning the shell, matches with the “Sferracavallo form” described by
Italy, Sicily, Palermo, Grotta Conza, 150 m asl, 38°11'13.61"N, 13°16'44.68"E, [Lab ID 55_1, COI: MW758902, ITS2: MW757124; Lab ID 55_2, COI: MW758903], W. De Mattia and J. Macor leg., 15.iv.2017. 15 live spm, 3 dissected spm. Italy, Sicily, Palermo, Sferracavallo, via Plauto, 50 m asl, 38°12'1.32"N, 13°16'3.05"E, [Lab ID 43_1, COI: MW758888], W. De Mattia and J. Macor leg., 15.iv.2017. 7 live spm, 2 dissected spm.
(Figs
(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 FO is longer than the V (FO/V range 2.1–2.4). The VD is thin along its whole course. The FDBC of the BC+SDBC is longer than the BC+SDBC (D/BC+SDBC = 2.1). The BC+SDBC is club-like to cylindrical and longer than the V (BC+SDBC/V range 1.4–1.6), with no clear distinction between the SDBC and the BC. The apex is wide and rounded. The D is longer than the V (D/V range 2.7–2.8) and longer that the BC+SDBC (D/BC+SDBC range 1.9–2.4), thinner than the BC+SDBC and with a pointed apex. The V is short and cylindrical. The A is very large. The PC is much longer than the V (P+E/V range 2.4–3.9). The PR is long and thin. There is a clear distinction between P and E as there is a visible ER and a proximal narrowing. The E is thinner but longer than the P (E/P range 1.4–1.8), almost gradually shrinking and turning into the VD.
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 PP is very elongated and smooth with a pointed tip. The P-E transition presents two slightly different structures. The population from the L.T. (Grotta Conza) presents a first distal ER, the PP and ELP originate from the second proximal ER. The epiphallar formula is: 1ER+2ER(PP+ELP). The V presents irregular, transverse to slightly longitudinal pleats. The population from Sferracavallo presents one ER with the PP originating from it. The ELP are not connected with the ER. The epiphallar formula is: 1ER(PP)+ELP. Distally, the E shows two main ELP that, at its mid length abruptly become finely fringed. The V presents a set of transverse-oblique smooth pleats that merge together one into another forming a kind of chevron or irregular pattern.
Siciliaria grohmanniana grohmanniana (Rossmässler, 1836), Monte Pellegrino, Palermo. 13.1–13.3 palatal plicae 13.4 parietal lamellae Siciliaria grohmanniana addaurae ssp. nov., Grotta dell’Addaura, Punta Priola 13.5–13.7 palatal plicae 13.8 parietal lamellae. Siciliaria septemplicata (Philippi, 1836), Monte Gallo, Sferracavallo 13.9–13.10 palatal plicae 13.11 parietal lamellae. Siciliaria leucophryna (L. Pfeiffer, 1862), Grotta Conza, Palermo 13.12–13.13 palatal plicae 13.14 parietal lamellae.
Siciliaria leucophryna colonises limestone cliffs hiding in cracks or under stones. It is very common on isolated boulders and tree trunks and barks in the xerophilic scrub around Grotta Conza. The species is quite abundant where found. According to
Siciliaria leucophryna is known to occur from a small area from Isola delle Femmine to Sferracavallo, including the northernmost sides of Pizzo Mollica and Pizzo Manolfo (where Grotta Conza is located) [(
In the COI tree two subclades of Siciliaria calcarae s. l. appear together with S. leucophryna in an unresolved trichotomy. The subspecies of Siciliaria calcarae are distributed within two haplogroups, one (haplogroup 1) comprising the nominate subspecies as well as S. c. belliemii, S. c. borgettensis ssp. nov., S. c. orlandoi, S. c. parajatinensis ssp. nov. and the other one (haplogroup 2) consisting of S. c. cruenta ssp. nov. and S. c. jatinensis ssp. nov. Unfortunately, S. parajatinensis could not be included into the ITS2 analysis. Yet, given the low distances among all representatives of S. calcarae in the ITS2 tree (and the resulting bad resolution), this data set does not tell us much concerning these relationships. Thus, presently the close relationship between S. c. orlandoi and S. c. parajatinensis hast to rely on mt data only. Concerning the anatomy of the genital organs, shell morphology, distribution, ecology and haplogroup of Siciliaria calcarae, two general types can be distinguished for each category.
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 S. calcarae calcarae (Philippi, 1844) morphotype, which is more or less striated (Bonifato, Ficuzza, Romitello, Visicari, Gibilmesi, and Castelluzzo), whereas the second type presents a markedly ribbed shell (Calatubo, Jato city park, Monte Kumeta, and Monte Belliemi).
Throughout the whole S. calcarae distribution, the nominate subspecies S. calcarae calcarae has the widest distribution, whereas all the remaining subspecies have an isolated, punctiform distribution. The populations from Alcamo, Visicari and Castelluzzo belong to the widely distributed nominate subspecies (1)m whereas the remaining populations (Ficuzza, Calatubo, Monte Kumeta, Jato city park, Romitello, Gibilmesi and Monte Belliemi) belong to limited, punctiform populations (2).
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 calcarae clade, eight populations fall into subclade 1 (Bonifato, Calatubo, Romitello, Monte Belliemi, Visicari and Catselluzzo, Monte Kumeta, Ficuzza), whereas two populations (Gibilmesi, and Jato city park) fall in to the subclade 2. Siciliaria calcarae orlandoi
The position of S. calcarae orlandoi in the COI tree within haplogroup 1 is close to sequences of a population of Siciliaria from the western slopes of Monte Kumeta. Despite the close phylogentic relationship of this population with S. calcarae orlandoi, the differences in the arrangement of the internal male genital organs (a set of transverse, interrupted fleshy smooth pleats and epiphallus with two main fringed chords vs. smooth longitudinal pleats and smooth proximal epiphallus) as well as the remarkable differences concerning shell morphology (see also
Two additional new subspecies of Siciliaria calcarae will be described in the following sections: S. calcarae borgettensis ssp. nov. and S. calcarae cruenta ssp. nov. Both taxa, despite showing close phylogenetic relationships with the other Siciliaria calcarae taxa, present distinctive shell and genital characters strong enough to propose their description as new subspecies.
Clausilia calcarae
Clausilia adelina
Clausilia adelina
–
Clausilia brugnoneana Pini 1884: 379.
Clausilia calcarae var. nodosa
Clausilia adelina var. subsolida
Delima (Siciliaria) calcarae
–
Charpentieria calcarae
–
Siciliaria calcarae
–
Siciliaria calcarae
–
Siciliaria (Siciliaria) calcarae calcarae
–
Siciliaria (Siciliaria) calcarae calcarae
–
Charpentieria calcarae
–
Italy, Sicily, San Vito lo Capo, Castelluzzo, west cliffs E of town, 120 m asl, 38°6'25.71"N, 12°44'33.37"E, [Lab ID 61_1, COI: MW758908, ITS2: MW757140 MW757141; Lab ID 61_2, COI: MW758909, ITS2: MW757142 MW757143 MW757144], W. De Mattia and J. Macor leg., 14.iv.2017. 2 dissected spm. Italy, Sicily, Castellammare del Golfo, Visicari, 395 m asl, 38°02'54.18"N, 12°48'26.61"E, [Lab ID 59_1, COI: MW758905, ITS2: MW757068 MW757069 MW757085], W. De Mattia and J. Macor leg., 15.iv.2017. 2 dissected spm. Italy, Sicily, Alcamo, Monte Bonifato, top of the mountain, 640 m asl, 37°57'29.40"N, 12°57'33.64"E, [Lab ID 5_1, COI: MW758925, ITS2: MW757125, MW757126, MW757127], I. Niero leg., 15.vi.2010. 2 dissected spm. Italy, Sicily, Alcamo, Monte Bonifato, top of the mountain, 640 m asl, 37°57'29.40"N, 12°57'33.64"E, A. Margelli leg., 15.vi.2010. 2 dissected spm. Italy, Sicily, Alcamo, Monte Bonifato, west side of the mountain, over the quarry, 550 m asl, 37°57'16.92"N, 12°58'9.06"E, W. De Mattia and J. Macor leg., 10.iv.2017. 2 dissected spm. Italy, Sicily, Castellammare del Golfo, Castello di Baida, W of the town along the road to Visicari, 300 m asl, 38°2'41.64"N, 12°48'14.34"E, W. De Mattia and J. Macor leg., 15.iv.2017. 2 dissected spm. Italy, Sicily, Piana degli Albanesi, 500 m south of Portella Ginestra, northern cliffs of Monte Kumeta, 970 m asl, 37°58'13.35"N, 13°15'22.06"E, W. De Mattia and J. Macor leg., 18.iv.2017. 2 dissected spm. Italy, Sicily, Calatafimi, Castello Eufemio, 395 m asl, 37°57'45.67"N, 12°51'21.13"E, W. De Mattia and J. Macor leg., 18.vi.2020. 2 dissected spm.
(Figs
(n = 50, not decollate). shell height 19.7 ± 0.8, whorl width 4.3 ± 0.3, aperture height 4.1 ± 0.2, aperture width 2.6 ± 0.1.
(Figs
(Figs
Siciliaria calcarae calcarae (Philippi, 1844), Visicari, Castellammare del Golfo 14.1 whole distal genital organs 14.2 internal distal part of genital organs. Castelluzzo, San Vito Lo Capo 14.3 whole distal genital organs 14.4 internal distal part of genital organs. Castello di Baida, Castellammare del Golfo 14.5 whole distal genital organs 14.6 internal distal part of genital organs. Piana delle Ginestre, Monte Kumeta 14.7 whole distal genital organs 14.8 internal distal part of genital organs. Calatafimi, Castello Eufemio. (= adelina Küster, 1847) 14.9 whole distal genital organs 14.10 internal distal part of genital organs.
Siciliaria calcarae calcarae (Philippi, 1844), Visicari, Castellammare del Golfo 15.1 shell 15.2 detail of the aperture 15.3 clausiliar plate double side. Castelluzzo, San Vito Lo Capo 15.4 shell 15.5 detail of the aperture 15.6 clausiliar plate double side. Castello di Baida, Castellammare del Golfo 15.7 shell 15.8 detail of the aperture 15.9 detail of the columellar side of last whorl. Monte Bonifato, Alcamo 15.10–15.12 shells. Calatafimi, Castello Eufemio, (= adelina Küster, 1847) 15.13 shell. Piana delle Ginestre, Monte Kumeta 15.14 shell 15.15 detail of the aperture. Monte Erice, western slopes 15.16 shell. Scopello 15.17 shell.
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.
(Figs
Siciliaria calcarae calcarae was found on south exposed limestone walls (Alcamo, Monte Bonifato), shady and humid north exposed limestone walls (Piana degli Albanesi, 500 m south of Portella Ginestra), shady habitats on decaying woods and tree trunks of Quercus ilex (Castelluzzo, west cliffs E of town) or among grass and shrub nearby a cave entrance (Castellammare del Golfo, Visicari). Siciliaria calcarae calcarae appeared to be adapted to many different habitats and niches and it is not an obliged rock-dwelling taxon. According to
Siciliaria calcarae calcarae presents the widest distributional area among the species of the genus Siciliaria. It is found from Trapani-Erice in the west to Montagna Grande and Bagheria in the east and Calatafimi-Piana degli Albanesi-Castelvetrano in the south. It is also known from the islands of Favignana and Levanzo (
Siciliaria calcarae calcarae shows minor shell differences among the populations which were described by the introduction of new names, such as a well-developed whole UPP (in Clausilia adelina var. subsolida), a well-developed AUPP in Clausilia adelina) or the presence of a knob as a second AUPP (Clausilia calcarae var. nodosa) (
Siciliaria (Siciliaria) calcarae belliemii
Charpentieria calcarae belliemii
–
Siciliaria calcarae belliemii
–
Siciliaria (Siciliaria) calcarae belliemii
–
Italy, Sicily, Partinico, W side of the Mount Belliemi, 440 m asl, 38°00'22.47"N, 13°6'37.79"E, [Lab ID 54_1, COI: MW758900, ITS2: MW757134, MW757135, MW757136; Lab ID 54_2, COI: MW758901], W. De Mattia and J. Macor leg., 12.iv.2017. 15 live spm, 3 dissected spm. Italy, Sicily, Alcamo, E side of the Calatubo Castle, 75 m asl, 38°00'54.80"N, 12°59'13.35"E, [Lab ID 44_1, COI: MW758889, ITS2: MW757122, MW757123; Lab ID 44_2, COI: MW758913; Lab ID 44_3, COI: MW758930; Lab ID 44_4, COI: MW758931], W. De Mattia and J. Macor leg., 14.iv.2017. 18 live spm, 2 dissected spm.
(Figs
(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.
(Figs
(Figs
Siciliaria calcarae calcarae (Philippi, 1844), Monte Bonifato 16.1 whole distal genital organs 16.2 internal distal part of genital organs 16.3 cross section of the spermatophore 16.4 spermatophore 16.5 whole distal genital organs 16.6 internal distal part of genital organs. Siciliaria calcarae belliemii (Brandt, 1961), Monte Belliemi, Partinico 16.7 whole distal genital organs 16.8 internal distal part of genital organs. Siciliaria calcarae belliemii (Brandt, 1961), Castello Calatubo, Alcamo 16.9 whole distal genital organs 16.10 internal distal part of genital organs 16.11 penial pseudopapilla.
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.
Siciliaria calcarae belliemii is known from Monte Belliemi near Partinico and from the limestone cliffs of the Calatubo Castle near Alcamo.
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 Siciliaria calcarae ssp. populations found both on limestone cliffs or different niches as tree’s barks (Visicari, Castelluzzo and Ficuzza).
Italy, Sicily, Municipality of Borgetto, Anime Sante hill, road to Romitello, 400 m asl, 38°02'59.53"N, 13°08'58.55"E.
1 Holotype (
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.
(Figs
Holotype : not decollate shell height 18.7, whorl width 4.2, aperture height 3.9, aperture width 2.8. Paratypes (n = 20, not decollate): shell height 19.0 ± 0.3, whorl width 4.3 ± 0.1, aperture height 3.8 ± 0.1, aperture width 2.9 ± 0.1.
The FO is as long as or slightly longer than the V (FO/V range 1.0–1.1). The VD is thin along its whole course. The FDBC is longer than the BC+SDBC (FDBC/BC+SDBC range 1.5–1.8). The BC+SDBC is club-like to cylindrical and as long as the V (BC+SDBC/V = 1.0), with no clear distinction between the SDBC and the BC. The apex is big and rounded. The D is longer than the V (D/V range 2.2–2.7) and longer that the BC+SDBC (D/BC+SDBC range 2.1–2.5), slightly thinner than BC+SDBC and with a small and round apex. The V is cylindrical. The A is very large. The PC is longer than the V (PC/V range 2.1–2.2). The PR is long and robust. The transition P-E does not show any ET. The E is almost as long as the P (E/P range 0.8–0.9), gradually shrinking and turning into the VD. The transition area between the E and the VD is not clearly visible outside.
(Fig.
Siciliaria calcarae borgettensis ssp. nov. was reported as “Sagana form” in
Siciliaria calcarae borgettensis ssp. nov. is exclusively known from the type locality. It is likely to be present along the whole northern limestone slopes of the Anime Sante hill (Romitello) as far as Sagana, thus further field investigations are needed in order to clearly define its actual distribution range.
The subspecies is found on limestone boulders and under scattered rocks.
Siciliaria calcarae borgettensis ssp. nov. is named after the nearby Borgetto town (Borgetto-Partinico), where the new subspecies was discovered.
Italy, Sicily, San Giuseppe Jato, quarry E of the town, 630 m asl, 37°58'15.07"N, 13°12'2.23"E.
1 Holotype (
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.
(Figs
Holotype : not decollate shell height 20.3, whorl width 4.3, aperture height 3.8, aperture width 2.4. Paratypes (n = 25, decollate): shell height 20.9 ± 1.0, whorl width 4.4 ± 0.1, aperture height 3.9 ± 0.2, aperture width 2.4 ± 0.2.
(Figs
(Figs
The shell of Siciliaria calcarae jatinensis ssp. nov. is very similar to the shells of Siciliaria calcarae belliemii and Siciliaria calcarae parajatinensis spp. nov. The three taxa have markedly ribbed shells. The density of the ribs along the body whorl is similar, with overlapping ranges: 32 ± 5 for Siciliaria calcarae belliemii (n = 25), 35 ± 6 for Siciliaria calcarae jatinensis ssp. nov. (n = 21) and 34 ± 5 for Siciliaria calcarae parajatinensis spp. nov. (n = 15). The PRI are short and stop at the level of the L, whereas the AUPP is strong to moderately weak and detached from the L. Slight differences are observed concerning the dimensions and the colour of the shell: Siciliaria calcarae jatinensis ssp. nov. is dark reddish to dark brown with an average height of 20.9 ± 1.0 whereas Siciliaria calcarae parajatinensis ssp. nov. (average height: 21.7 ± 1.4) and Siciliaria calcarae belliemii (average height: 17.6 ± 1.2, type locality) are light yellowish to light brown. The three subspecies show just slight differences in their anatomy of the genital organs, which fit into the intraspecific variability of S. calcarae and do not allow a certain separation. The different subspecific status of these populations is mainly supported by the phylogenetic results in connection to their patchy and isolated distribution. Siciliaria calcarae jatinensis ssp. nov. is included in the haplogroup 2 whereas Siciliaria calcarae parajatinensis n. spp. and Siciliaria calcarae belliemii both belong to haplogroup 1.
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 S. tiberii), based on
In the COI and combined tree (Fig.
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 Siciliaria tiberii (Figs
Siciliaria calcarae jatinensis ssp. nov. is known only from the type locality: the abandoned quarry north of the town of San Giuseppe Jato. Its actual distribution range must be further investigated as it is probably present along the northwestern cliffs of the Monte Jato, E of the town of San Giuseppe Jato.
The new subspecies is an obliged limestone rock-dweller, exclusively collected on walls and cliffs, both natural or resulting from the previous quarrying activity.
Siciliaria calcarae jatinensis ssp. nov. is named after Monte Jato (San Giuseppe Jato), where the new subspecies was discovered.
Italy, Sicily, Monreale, W part of Monte Kumeta toward Jato Antica, 630 m asl, 37°57'8.18"N, 13°13'14.57"E.
1 Holotype (
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.
(Figs
Siciliaria calcarae borgettensis ssp. nov., road to Romitello, Borghetto 18.1 whole distal genital organs 18.2 internal distal part of genital organs. Siciliaria calcarae jatinensis ssp. nov. San Giuseppe Jato 18.3 whole distal genital organs 18.4 internal distal part of genital organs 18.5 detail of the pseusopapilla 18.6 whole distal genital organs 18.7 internal distal part of genital organs. Siciliaria calcarae parajatinensis ssp. nov., west side Monte Kumeta 18.8 whole distal genital organs 18.9 internal distal part of genital organs.
Holotype : not decollate shell height 21.7, whorl width 4.2, aperture height 4.1, aperture width 3.3. Paratypes (n = 30, not decollate): shell height 21.7 ± 1.4, whorl width 4.4 ± 0.2, aperture height 4.2 ± 0.2, aperture width 3.1 ± 0.4.
(Fig.
(Fig.
Siciliaria calcarae parajatinensis ssp. nov. in the COI tree is well embedded within the calcarae subclade and closely related to the nominate subspecies (Fig.
Siciliaria calcarae parajatinensis ssp. nov. is exclusively known from the type locality, the western slopes of the Monte Kumeta. Further field research could extend its distributional range to the whole western side of the Monte Kumeta (Monreale) and surrounding mountains. Following
Siciliaria calcarae parajatinensis ssp. nov. is an obliged rock-dweller and inhabits limestone walls and boulders. The type locality, that is not included in any protected area, comprises a rich population.
Para = next to, considering strong morphological, genetic and distribution affinities with the newly described Siciliaria calcarae jatinensis ssp. nov.
Siciliaria calcarae borgettensis ssp. nov., road to Romitello, Borghetto 19.1 shell 19.2 detail of the aperture 19.3 clausiliar plate double side 19.4 shell 19.5 detail of the aperture. Siciliaria calcarae jatinensis ssp. nov. San Giuseppe Jato 19.6 shell 19.7 detail of the aperture 19.8 clausiliar plate double side. Siciliaria calcarae parajatinensis ssp. nov., west side Monte Kumeta 19.9 shell 19.10 detail of the aperture 19.11 detail of the columellar side of last whorl 19.12 clausiliar plate double side.
Siciliaria (Siciliaria) calcarae orlandoi Liberto, Reitano, Giglio Colomba and Sparacio 2016: 371.
(Figs
(n = 12, not decollate). shell height 18.0 ± 0.7, whorl width 4.1 ± 0.1, aperture height 3.8 ± 0.2, aperture width 2.7 ± 0.1.
(Fig.
(Fig.
Following
Siciliaria calcarae orlandoi in known to occur in the Nature Reserve of the Bosco della Ficuzza, Rocca della Busambra, Bosco del Cappelliere e Gorgo del Drago that is included in the wider Sicani Mountains Regional Nature Park (
Italy, Sicily, Monreale, N side of Monte Gibilmesi, 890 m asl, 38°03'37.03"N, 13°12'38.53"E.
1 Holotype (
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.
(Figs
Siciliaria calcarae orlandoi Liberto, Reitano, Giglio, Colomba & Sparacio, 2016, Bosco Ficuzza 20.1 whole distal genital organs 20.2 internal distal part of genital organs. Siciliaria calcarae cruenta ssp. nov., Monte Gibilmesi 20.3 whole distal genital organs 20.4 internal distal part of genital organs.
Siciliaria calcarae orlandoi Liberto, Reitano, Giglio, Colomba & Sparacio, 2016, Bosco Ficuzza 21.1 shell 21.2 detail of the aperture 21.3 clausiliar plate double side 21.4 shell 21.5 detail of the aperture. Siciliaria calcarae cruenta ssp. nov., Monte Gibilmesi 21.6 shell 21.7 detail of the aperture 21.8 shell 21.9 detail of the aperture 21.10 view of the clausiliar plate.
Holotype : not decollate shell height 19.2, whorl width 4.4, aperture height 4.3, aperture width 3.2. Paratypes (n = 11, not decollate): shell height 19.4 ± 1.0, whorl width 4.5 ± 0.3, aperture height 4.3 ± 0.3, aperture width 3.1 ± 0.2.
(Fig.
(Fig.
The morphologically most similar taxon to Siciliaria calcarae cruenta ssp. nov. is Siciliaria calcarae borgettensis ssp. nov., but it differs from the latter by its very dark and remarkably smoother shell and the AUPP is not sharp but always suffused and almost not visible from the aperture. As regards the genital organs, contrary to Siciliaria calcarae borgettensis ssp. nov., Siciliaria calcarae cruenta ssp. nov. presents a very long SDBC+BC and FO. The sculpturing of the internal penis differs by the less fringed longitudinal pleats and the EPLs that reach proximally back as far as the beginning of the VD. Siciliaria calcarae cruenta ssp. nov. is similar to Siciliaria calcarae orlandoi, anyhow the latter presents a more finely striated shell, its AUPP is stronger and usually closer to the lunella. It was found in haplogroup 1, while Siciliaria calcarae cruenta ssp. nov. is in the haplogroup 2.
Siciliaria calcarae cruenta ssp. nov. in known only from the type locality: northern slopes of Monte Gibilmesi near Sagana (Montelepre). Further field investigation is needed in order to determine the actual distribution of the taxon.
This subspecies was found climbing on limestone cliffs hiding among moss and rocks crevices.
Siciliaria calcarae cruenta ssp. nov. was named after its intense dark red colour of the shell (cruentus = bloody).
(?) Clausilia confinata commeata –
Siciliaria ferrox
Siciliaria (Siciliaria) ferrox
–
Siciliaria ferrox
–
Siciliaria ferrox
–
Siciliaria ferrox
–
Charpentieria ferrox
–
In the COI tree (Fig.
The status of Clausilia confinata commeata Westerlund, 1892 (
(Figs
(n = 32, decollate). shell height 17.7 ± 0.7, whorl width 3.8 ± 0.1, aperture height 3.6 ± 0.2, aperture width 2.4 ± 0.2.
(Fig.
(Figs
Siciliaria ferrox inhabits limestone walls, hiding in rocks cracks and crevices. According to
Siciliaria ferrox occupies the easternmost part of the distribution range of the northwestern Sicilian Siciliaria. It is found from Altavilla Milicia, Sant’Onofrio to Termini Imerese.
Clausilia tiberii
Clausilia tiberiana
Siciliaria tiberii
–
Siciliaria (Siciliaria) tiberii
–
Charpentieria tiberii
–
Charpentieria (Siciliaria) tiberii tiberii
–
Siciliaria tiberii
–
Siciliaria tiberii
–
Charpentieria tiberii
–
(Figs
(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.
(Fig.
(Fig.
Siciliaria calcarae calcarae (Philippi, 1844), Visicari, Castellammare del Golfo 24.1 palatal plicae 24.2 parietal lamellae. Siciliaria calcarae belliemii (Brandt, 1961), Monte Belliemi, Partinico 24.3 palatal plicae. Siciliaria calcarae belliemii (Brandt, 1961), Castello di Calatubo, Alcamo 24.4 palatal plicae. Siciliaria calcarae borgettensis ssp. nov., road to Romitello, Borghetto 24.5–24.6 palatal plicae. Siciliaria calcarae jatinensis ssp. nov. San Giuseppe Jato 24.7 palatal plicae 24.8 parietal lamellae. Siciliaria calcarae parajatinensis ssp. nov., west side Monte Kumeta 24.9–24.10 palatal plicae. Siciliaria calcarae orlandoi Liberto, Reitano, Giglio, Colomba & Sparacio, 2016, Bosco Ficuzza 24.11 palatal plicae 24.12 parietal lamellae. Siciliaria calcarae cruenta ssp. nov., Monte Gibilmesi 24.13–24.14 palatal plicae. Siciliaria ferrox (Brandt, 1961). Trabia Sant’Onofrio 24.15 palatal plicae 24.16 parietal lamellae.
Siciliaria tiberii tiberii is found under stones, among scree, rocky debris and on dry stonewalls along the coastline, on a limestone plateau. According to
As a result, the actual distributional range of Siciliaria tiberii tiberii seems to be limited to the area from San Cataldo (Partinico) in the south to Monte Palmeto and Capo Rama (Terrasini) in the north to the northern slopes of Monte Pecoraro. Further field research is needed in order to define its actual distribution. Recent findings of Diego Viola (Trieste, Italy) from a locality between San Giuseppe Jato and Piana degli Albanesi revealed the presence of a population of Siciliaria cfr. tiberii that clearly differ, as a shell, from S. calcarae jatinensis ssp. nov. and S. calcarae parajatinensis ssp. nov.(personal communication). The poor samples available (few damaged shells) did not allow us a thorough investigation.
Monte San Calogero in the east (cited by Nordsieck as also included in the S. tiberii distribution), is an isolate limestone massif 10 km SE of Termini Imerese, 50 km east of San Cipirello and 65 km SSE of Capo Rama. The presence of S. tiberii tiberii in this locality, due to its remarkable distance, should be confirmed by further field research and not only inferred from museum samples. Mislabelling or wrong locality assignments could have played a role in this regard (as for Siciliaria septemplicata alcamoensis, see below). The diagnosis provided by
Siciliaria (Siciliaria) alcamoensis
Siciliaria (Siciliaria) septemplicata alcamoensis
–
Siciliaria septemplicata hemmeni
Charpentieria (Siciliaria) septemplicata alcamoensis
–
Siciliaria septemplicata alcamoensis
–
(Figs
(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.
(Fig.
(Figs
The PP is large, elongated, and heavily wrinkled. The P-E transition presents one ER with both the PP and ELP originating from the ER. The epiphallar formula is: 1ER(PP+ELP). The E shows a set of three or four smooth longitudinal pleats. The V is almost completely smooth. The wall shows a fine granulation.
Siciliaria tiberii alcamoensis comb. nov. inhabits scattered isolated limestone boulders, and is hiding in rock cracks and crevices. After careful field research in a wide area around Piano Margi, the taxon revealed to be remarkably rare and scattered.
Siciliaria tiberii alcamoensis comb. nov. is known only from a small area of less than 1 km2 in the surrounding of the type locality: Piano Margi south of Cinisi on the Montagna Longa. Further field research is needed to define its actual distribution range.
Siciliaria tiberii alcamoensis comb. nov. (Figs
Siciliaria tiberii tiberii (A. Schmidt, 1868), Capo Rama, Terrasini 25.1 whole distal genital organs 25.2 internal distal part of genital organs. Siciliaria tiberii alcamoensis Brandt, 1961 comb. nov., Piano Margi, Carini 25.3 whole distal genital organs 25.4 internal distal part of genital organs 25.5 detail of the pseudopapilla.
Siciliaria tiberii tiberii (A. Schmidt, 1868), Capo Rama, Terrasini 26.1 shell 26.2 clausiliar plate double side 26.3 shell 26.4 detail of the aperture 26.5 clausiliar plate double side. Siciliaria tiberii alcamoensis Brandt, 1961 comb. nov., Piano Margi, Carini 26.6 shell 26.7 clausiliar plate double side 26.8 detail of the aperture 26.9 shell.
Italy, Sicily, Carini, Grotta dei Puntali o dell’Armetta, 90 m asl, 38°8'58.56"N, 13°9'25.83"E.
1 Holotype (
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 (ALPP) present; palatal edge of clausilium plate distally not receding, palatal edge distally strongly bent upwards.
(Figs
Holotype : decollate shell height 20.1, whorl width 4.8, aperture height 4.6, aperture width 3.1. Paratypes (n = 35, decollate): shell height 19.7 ± 1.6, whorl width 4.6 ± 0.2, aperture height 4.2 ± 0.3, aperture width 2.8 ± 0.3.
(Fig.
(Figs
Siciliaria tiberii armettensis ssp. nov., like the nominate subspecies, has a ribbed shell and it is clearly distinguishable from all the remaining subspecies that present a more or less striated but ribless shell surface. Nonetheless, the nominate subspecies shows a stronger ribbing, with elevated and robust ribs. The distal end of the AUPP always presents a knob that is well visible from the frontal view (Figs
Siciliaria tiberii armettensis ssp. nov. inhabits east-exposed limestone walls and cliffs, hiding among tuft of vegetation, rocks cracks and crevices.
Siciliaria tiberii armettensis ssp. nov. is exclusively known from the type locality, the surroundings of the Grotta dei Puntali o dell’Armetta. Further field investigation along the Monte Pecoraro eastern slopes is needed in order to determine the actual distribution of the taxon. Grotta dei Puntali o dell’Armetta are included in the protected area Riserva Naturale Grotta dei Puntali.
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 (
Siciliaria tiberii armettensis ssp. nov., Grotta dei Puntali, Carini 27.1 whole distal genital organs 27.2 internal distal part of genital organs 27.3 detail of the pseudopapilla. Siciliaria tiberii scalettensis Beckmann, 2004 Portella Scaletta, Villagrazia 27.4 whole distal genital organs 27.5 internal distal part of genital organs 27.6 detail of the pseudopapilla.
Charpentieria (Siciliaria) tiberii scalettensis
Siciliaria tiberii scalettensis
–
Siciliaria tiberii scalettensis
–
Italy, Sicily, Cinisi, Portella Scaletta, 90 m asl, 38°10'35.53"N, 13°07'27.22"E, [Lab ID 17_2, COI: MW758923, ITS2: MW757108, MW757109, MW757110], W. De Mattia and J. Macor leg., 18.iv.2017. 3 dissected spm. Italy, Sicily, Cinisi, along the SS113 road from Villagrazia di Carini to Cinisi, 70 m asl, 38°10'42.45"N, 13°07'34.50"E, [Lab ID 39_1, COI: MW758882, ITS2: MW757095, MW757096, MW757097], W. De Mattia and J. Macor leg., 18.iv.2017. 2 dissected spm.
(Figs
Siciliaria tiberii armettensis ssp. nov., Grotta dei Puntali, Carini 28.1 shell 28.2 shell clausiliar plate double side 28.3 detail of the aperture 28.4 clausiliar plate double side. Siciliaria tiberii scalettensis Beckmann, 2004 Portella Scaletta, Villagrazia 28.5 shell 28.6 detail of the aperture 28.7 shell 28.8 detail of the aperture 28.9 clausiliar plate double side.
(n = 45, decollate). shell height 18.5 ± 1.3, whorl width 4.6 ± 0.2, aperture height 4.0 ± 0.2, aperture width 2.9 ± 0.2.
(Fig.
Siciliaria tiberii tiberii (A. Schmidt, 1868), Capo Rama, Terrasini 29.1–29.2 palatal plicae 29.3 parietal lamellae. Siciliaria tiberii alcamoensis Brandt, 1961 comb. nov., Piano Margi, Carini 29.4 parietal lamellae 29.5 palatal plicae. Siciliaria tiberii armettensis ssp. nov., Grotta dei Puntali, Carini 29.6 parietal lamellae 29.7–29.8 palatal plicae. Siciliaria tiberii scalettensis Beckmann, 2004 Portella Scaletta, Villagrazia 29.9–29.10 palatal plicae 29.11 parietal lamellae.
(Figs
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 PP is also markedly different, being heavily wrinkled but the epiphallar formula is the same as the LT. The E shows a pattern of three or four smooth longitudinal pleats. These pleats merge one into another forming two main fringed pleats that run as far as the VD. The V is almost smooth with a very fine granulation.
Siciliaria tiberii scalettensis inhabits limestone walls and cliffs, hiding among tuft of vegetation, rocks cracks and crevices. The taxon is common at the type locality (Villa Paradiso) with a rich population and few more scattered, smaller populations among the limestone spur.
Siciliaria tiberii scalettensis is known only from a small area of less than 0.5 km2 around the of the type locality: Portella Scaletta along the SS113 road, from Villagrazia di Carini to Cinisi (Palermo). It was collected along a limestone spur heading NNE.
90% of the specimens (n = 45) of Siciliaria tiberii scalettensis present a second, lower AUPP, shorter than the superior AUPP and detached both from the AUPP and the L. This feature is shared only with the nominate subspecies.
In Table
Main morphological differences (shell and genital organs) among the Sicania taxa.
Sicania crassicostata comb. nov. | Sicania eminens comb. nov. | Sicania nobilis nobilis comb. nov. | Sicania nobilis spezialensis comb. nov., stat. nov. | |
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 |
AUPP | missing | present, visible from outside | present, mostly not visible from outside | mostly missing |
ALPP | 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 |
Infraordo Clausilioidei Gray, 1855
Superfamilia Clausilioidea Gray, 1855
Familia Clausiliidae Gray, 1855
Subfamilia Alopiinae A.J. Wagner, 1913
Tribus Delimini R.A. Brandt, 1956
Clausilia crassicostata
L.
Clausilia crassicostata
–
Siciliaria crassicostata
–
Clausilia crassicostata
–
Siciliaria (Siciliaria) crassicostata
–
Charpentieria crassicostata
–
Charpentieria crassicostata
–
Siciliaria crassicostata
–
Charpentieria crassicostata
–
. The morphology of the genital organs of Sicania crassicostata comb. nov. revealed to be very stable in both of the investigated populations (Fig.
The shell is remarkably ribbed to percostated. It shares few characters with Sicania nobilis spezialensis comb. nov. stat. nov., e.g., the (mostly) missing AUPP, both the very short (or missing) ALPP and the spiralis (mostly) not overlapping the parietalis (Fig.
Although Sicania eminens comb. nov. was initially introduced by
Italy, Sicily, Castelluzzo, cliffs W of the Tonnara di Monte Cofano, 50 m asl, 38°06'44.89"N, 12°40'34.27"E, [Lab ID 33_1, COI: MW758876; Lab ID 33_2, COI: MW758877, ITS2: MW757103, MW757104, MW757105 MW757131], W. De Mattia and J. Macor leg., 21.xii.2007. 3 dissected spm. Italy, Sicily, Custonaci, cliffs N of the Mangiapane cave, 63 m asl, 38°05'40.91"N, 12°40'10.98"E, [Lab ID 1_1, COI: MW758912, ITS2: MW757114, MW757113; Lab ID 1_4, COI: MW758924] W. De Mattia and J. Macor leg., 21.xii.2007. 2 dissected spm.
(Figs
(n = 24, decollate). shell height 24.6 ± 1.2, whorl width 4.9 ± 0.1, aperture height 4.8 ± 0.3, aperture width 3.0 ± 0.1.
(Figs
(Figs
Sicania crassicostata comb. nov. was collected on limestone cliffs or under rocks and boulders. According to
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) (
Clausilia crassicostata var. eminens
Clausilia crassicostata var. eminens
–
Siciliaria crassicostata eminens
–
Charpentieria eminens
–
Charpentieria eminens
–
Siciliaria crassicostata var. eminens
–
Siciliaria eminens
–
Charpentieria eminens
–
Sicania eminens comb. nov. inhabits a restricted range that approximately includes the surroundings of Custonaci and the S slopes of Monte Cofano in the east as far as Castellammare del Golfo in the west.
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 Sicania crassicostata comb. nov. and could be, by inexperienced eyes, easily confused with it. Probably this is the reason why the taxon was for long time considered as a subspecies of Sicania crassicostata comb. nov. (Alzona, 1971: 92). Nevertheless, Sicania eminens comb. nov. can be distinguished by the markedly pyriform shape of its shell and the presence of a strong anterior upper palatal plica and the ALPP, both of them clearly visible from outside of the shell’s aperture and missing in Sicania crassicostata comb. nov. (Figs
Sicania eminens comb. nov. is monophyletic in both the COI and in the ITS2 tree (both well supported). The ITS2 sequences were almost identical (Fig.
Italy, Sicily, Custonaci, contrada Scurati, crossing with Strada Procinciale 19, 60 m asl, 38°4'55.97"N, 12°40'9.91"E, [Lab ID 51_1, COI: MW758896, ITS2 MW757072; Lab ID 51_2, COI: MW758897], W. De Mattia and J. Macor leg., 12.iv.2017. 2 dissected specimens. Italy, Sicily, Custonaci, contrada Scurati, 60 m asl, 38°05'18.10"N, 12°40'17.18"E, [Lab ID 52_1, COI: MW758898; Lab ID 52_2, COI: MW758899, ITS2 MW757071, MW757085, MW757086], W. De Mattia and J. Macor leg., 12.iv.2017. 2 dissected spm. Italy, Sicily, Custonaci, contrada Scurati, crossing with Strada Provinciale 18, 65 m asl, 38°4'56.04"N, 12°40'9.86"E, W. De Mattia and J. Macor leg., 12.iv.2017. 2 dissected spm. Italy, Sicily, Custonaci, contrada Scurati, limestone cliffs S of the village, 60 m asl, 38°06'44.89"N, 12°40'34.27"E, W. De Mattia and J. Macor leg., 12.iv.2017. 2 dissected spm. Italy, Sicily, Custonaci, Buffara, NE side of the hollow, 150 m asl, 38°04'0.52"N, 12°41'1.34"E, [Lab ID 49_1, COI: MW758892; Lab ID 49_2, COI: MW758893], W. De Mattia and J. Macor leg., 12.iv.2017. 2 dissected spm. Italy, Sicily, Custonaci, Baglio Messina, 340 m asl, 38°04'6.38"N, 12°41'48.68"E, [Lab ID 50_1, COI: MW758894, ITS2 MW757073, MW757074, MW757075, MW757076; Lab ID 50_2, COI: MW758895], W. De Mattia and J. Macor leg., 12.iv.2017. 2 dissected spm.
(Figs
(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.
(Figs
(Figs
Sicania eminens (A. Schmidt, 1868), comb. nov., Custonaci, Baglio Messina 32.1 whole distal genital organs 32.2 internal distal part of genital organs 32.3 detail of the pseudopapilla. Dolina Buffara 32.4 whole distal genital organs 32.5 internal distal part of genital organs. Contrada Scurati, Custonaci 32.6 whole distal genital organs 32.7 internal distal part of genital organs. Contrada Scurati toward Mangiapane 32.8 whole distal genital organs 32.9 internal distal part of genital organs.
Sicania eminens comb. nov. inhabits limestone cliffs and boulders, hiding in rock cracks and crevices. It is also found climbing on tree barks and decaying woods (Custonaci, Baglio Messina). This taxon has never been found syntopic with other Sicania or Siciliaria taxa. Along its eastern distributional range, it is sympatric with Siciliaria calcarae calcarae. According to
The distribution range of Sicania eminens comb. nov. includes the southern slopes of Monte Cofano and Monte Palatimone in the north and, in the south, the line from Monte Sparagio-Castellammare del Golfo (east) and Monte Buffara (west).
Sicania nobilis comb. nov. s. l. has a big, weakly striated shell, mostly decollate and whitish in colour. Sicania nobilis comb. nov. forms a well-supported clade in the COI tree. Even in the ITS2 tree almost all sequences cluster, except those of population 14 (SanVito Lo Capo, Torazza, 3 clones), which are found within the S. eminens + S. nobilis cluster. This might point towards hybridisation between those taxa. Such an assumption would deserve further investigation.
Sicania eminens (A. Schmidt, 1868), comb. nov., Custonaci, Baglio Messina 33.1 shell. 33.2 detail of the aperture 33.3 clausiliar plate double side. Dolina Buffara 33.4 shell 33.5 detail of the aperture 33.6 clausiliar plate double side. Contrada Scurati, Custonaci 33.7 shell 33.8 detail of the aperture 33.9 clausiliar plate double side. Contrada Scurati toward Mangiapane 33.10 shell 33.11 detail of the aperture 33.12 clausiliar plate double side.
Sicania nobilis comb. nov. cannot be framed within a precise genital morphological outline. The anatomy of the genital organs of Sicania nobilis comb. nov. exhibits significant differences among the populations (albeit stable within a population), especially concerning the features of the internal sculpturing of V (smooth to transversal pleats), penis (smooth, with tubercles of fringed longitudinal pleats) and the shape of the penial pseudopapilla (small and roundish to big and elongate). As seen in the genital descriptions, also ratios among principal genital parts show great variability. Sicania nobilis spezialensis stat. nov. comb. nov. shows a further different genital outline (Fig.
Sicania nobilis comb. nov. s. l. is clearly differentiated from the remaining Sicania/Siciliaria taxa by shell morphology, especially the waxy whitish surface of the whorls and the arrangement of the plicae and lamellae, as repeatedly reported in literature with detailed descriptions and differential diagnoses (
Clausilia nobilis L. Pfeiffer 1848: 434.
Clausilia sicula
Siciliaria (Siciliaria) nobilis episoma
Siciliaria (Siciliaria) nobilis
–
Siciliaria (Siciliaria) nobilis
(= episoma) –
Charpentieria nobilis
–
Siciliaria nobilis
–
Siciliaria nobilis
–
Charpentieria nobilis
–
Italy, Sicily, Castelluzzo, Monte Cofano E of Tonnara di Monte Cofano, 80 m asl, 38°06'22.04"N, 12°40'59.77"E, W. De Mattia and J. Macor leg., 19.xii.2007. 2 dissected spm. I.D. 7. Italy, Sicily, San Vito lo Capo, Castelluzzo, west cliffs E of town, 120 m asl, 38°06'25.71"N, 12°44'33.37"E, [Lab ID 60_1, COI: MW758906, ITS2: MW757065, MW757066, MW757067, MW757139; Lab ID 60_2, COI: MW758907; Lab ID 60_4, COI: MW758938; Lab ID 60_5, COI: MW758939], W. De Mattia and J. Macor leg., 14.iv.2017. 2 dissected spm. Italy, Sicily, San Vito lo Capo, cliffs S of “El-Bahira” camping, 60 m asl, 38°08'48.81"N, 12°44'6.61"E, [Lab ID 62_2, COI: MW758914; Lab ID 62_3, COI MW758940], W. De Mattia and J. Macor leg., 14.iv.2017. 2 dissected spm. Italy, Sicily, San Vito lo Capo, Torre delle Usciere, 5 m asl, 38°10'30.07"N, 12°46'14.09"E, [Lab ID 18_1, COI: MW758918, ITS2: MW757106, MW757107], W. De Mattia and J. Macor leg., 19.xii.2007. 2 dissected spm. “episoma” Italy, Sicily, San Vito lo Capo, boulders W of the town, 50 m asl, 38°10'51.85"N, 12°43'37.70"E, [Lab ID 14_1, COI: MW758875, ITS2: MW757128, MW757129, MW757130], W. De Mattia and J. Macor leg., 19.xii.2007. 2 dissected spm.
(Figs
(n = 45, decollate). shell height 22.9 ± 1.6, whorl width 5.0 ± 0.2, aperture height 4.8 ± 0.3, aperture width 3.6 ± 0.4.
(Figs
(Figs
Sicania nobilis nobilis comb. nov. is a strictly limestone dweller and inhabits cliffs and big isolated boulders, hiding among rock cracks and crevices. It is found both in dry, sun exposed cliffs (i.e., San Vito lo Capo, cliffs S of ”El-Bahira“ camping) or shady walls under dense tree cover (San Vito lo Capo, Castelluzzo, west cliffs E of town). According to
Sicania nobilis nobilis comb. nov. is known to occur in two limestone massifs: Monte Cofano and in San Vito lo Capo Peninsula, from Monte Acci to Monte Monaco.
Siciliaria nobilis episoma hwas described by
Siciliaria (Siciliaria) spezialensis
Siciliaria (Siciliaria) spezialensis
–
Charpentieria spezialensis
–
Siciliaria spezialensis
–
Siciliaria spezialensis
–
Charpentieria spezialensis
–
(Figs
Sicania nobilis nobilis (A. Schmidt, 1868), comb. nov., Castelluzzo 34.1 whole distal genital organs 34.2 internal distal part of genital organs. San Vito Lo capo, camping 34.3 whole distal genital organs 34.4 internal distal part of genital organs. San Vito Lo Capo, boulders 34.5 whole distal genital organs 34.6 internal distal part of genital organs. Torre delle Usciere (ex. forma episoma) 34.7 whole distal genital organs 34.8 internal distal part of genital organs.
(n = 24, decollate). shell height 22.2 ± 1.3, whorl width 5.1 ± 0.2, aperture height 4.1 ± 0.3, aperture width 2.8 ± 0.3.
(Figs
(Figs
Sicania nobilis spezialensis stat. nov., comb. nov. is a strictly limestone dweller and inhabits cliffs, hiding among rock cracks and crevices. As previously deemed as a species, according to
Sicania nobilis spezialensis stat. nov., comb. nov. is known only from the type locality: along the western cliffs of Monte Speziale in the surroundings of Macari (San Vito lo Capo). No overlapping zone with Sicania nobilis nobilis stat. nov., comb. nov. is known until now. Although, in its original description, the taxon was reported for one locality only (
Sicania nobilis spezialensis stat. nov., comb. nov. is now considered as a subspecies of S. nobilis. It is well embedded within the clade of Sicania nobilis in both COI and combined trees. Sicania nobilis spezialensis stat. nov., comb. nov. is a local isolated, spatially restricted ribbed form of the more widespread Sicania nobilis comb. nov. with few different features of the shell, viz. the lack of the anterior upper palatal plica and a clausilium plate that fits into a frame of the palatal plicae.
The genus Mauritanica comprises four species that are distributed over northeastern Algeria and central-northern Tunisia (
35. Sicania nobilis spezialensis (
In our molecular genetic analysis (Figs
Sicania nobilis nobilis (A. Schmidt, 1868), comb. nov., Castelluzzo 36.1 shell 36.2 detail of the aperture 36.3 clausiliar plate double side. San Vito Lo capo, camping 36.4 shell 36.5 detail of the aperture 36.6 clausiliar plate double side. San Vito Lo Capo, boulders 36.7 shell 36.8 detail of the aperture 36.9 clausiliar plate double side. Torre delle Usciere (ex. forma episoma) 36.10 shell 36.11 detail of the aperture 36.12 clausiliar plate double side. Monte Cofano, Tonnara Cofano 36.13 shell. Sicania nobilis spezialensis (
The separation of Mauritanica from Siciliaria/Sicania is consistent through all the trees, thus we provisionally keep Mauritanica as a distinct genus, although the type species Mauritanica tristrami (L. Pfeiffer,1861) was not available to us. In order to fully understand the phylogenetic relationships of Mauritanica with Siciliaria/Sicania more genetic markers and specimens are essential.
Both M. perinni polygyra and M. scarificata underwent anatomical investigation. A similar external genital morphology was detected. The only noticeable difference is the relative length of the FO, with a V/FO ratio of 1.1 for M. scarificata and 0.6 for M. perinni polygyra. Remarkable differences were detected regarding the internal details of the penial complex. Mauritanica scarificata presents 3 ERs with a very short and globose PP that makes it fall into the general Siciliaria/Sicania genital anatomical arrangement. Mauritanica perinni polygyra lacks any ER and the PP is irregular with a convolute surface pattern. It is directly connected to the OELP. Unfortunately, the genital descriptions and the pictures provided by
Following our genital anatomical investigations, the common genital anatomical traits deemed by
Clausilia scarificata
L.
Clausilia scarificata
– L.
Clausilia sacrificata
–
Clausilia confinata
–
Clausilia (Siciliaria) confinata
Siciliaria scarificata
–
Charpentieria scarificata
–
Siciliaria scarificata
–
Siciliaria scarificata
–
Siciliaria scarificata
–
Charpentieria scarificata
–
Charpentieria scarificata
–
(Figs
(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.
(Fig.
(Figs
Mauritanica scarificata comb. nov. is found in a variety of habitats: limestone walls and cliffs, isolated boulders, under rocks and among scree and rocky debris. The species is common and abundant throughout the Island of Marettimo. According to
Mauritanica scarificata comb. nov. is endemic of the island of Marettimo.
Clausilia (Mauritanica) polygyra Boettger in Kobelt 1879: 153.
Charpenteria (Mauritanica) perinni polygyra
–
Charpenteria (Mauritanica) perinni polygyra
–
Mauritanica perinni polygyra
–
Mauritanica perinni polygyra was recently revised by
Mauritanica scarificata (L. Pfeiffer, 1856), comb. nov., Marettimo 37.1 whole distal genital organs 37.2 internal distal part of genital organs 37.3 whole distal genital organs 37.4 internal distal part of genital organs 37.5 detail of the pseudopapilla. Mauritanica perinni polygyra (O. Boettger, 1879), Tunisia, Zaghouan 37.6 whole distal genital organs 37.7 internal distal part of genital organs 37.8–37.9 spermatophore 37.10 cross section of the spermatophore.
(Figs
(n = 1, decollate). shell height 22.4, whorl width 4.1, aperture height 3.3, aperture width 2.4.
(Fig.
(Fig.
(Figs
Mauritanica perinni polygyra is a limestone dweller, on cliffs, crevices and under stones (
Mauritanica perinni polygyra is known only from Djebel Zaghouan area (
Mauritanica scarificata (L. Pfeiffer, 1856), comb. nov., Marettimo 38.1 shell 38.2 shell 38.3 detail of the aperture 38.4 clausiliar plate double side. Mauritanica perinni polygyra (O. Boettger, 1879), Tunisia, Zaghouan 38.5 shell 38.6 shell (picture courtesy of Claude and Amandine Evanno).
The separation of Siciliaria and Sicania (and provisionally Mauritanica) from the other genera is well supported, but the relationships among Charpentieria, Stigmatica and Gibbularia remain unresolved and appear as polytomy in the trees. Charpentieria (four species included) forms one (poorly supported) clade but, the relationships within this clade received no considerable support. The genera Stigmatica and Gibbularia are not monophyletic and populations from distant areas show high genetic distances (Table
Sicania crassicostata (L. Pfeiffer, 1856), comb. nov., Monte Cofano 39.1 palatal plicae 39.2 parietal lamellae. Sicania eminens (A. Schmidt, 1868), comb. nov., Custonaci, Baglio Messina 39.3–39.4 palatal plicae. Sicania nobilis nobilis (L. Pfeiffer, 1848), comb. nov., Castelluzzo 39.5–39.6 palatal plicae 39.7 parietal lamellae. Mauritanica scarificata (L. Pfeiffer, 1856), comb. nov., Marettimo 39.8 palatal plicae 39.9 parietal lamellae.
Papillifera, which was used as outgroup, appeared monophyletic in our trees, even though the distances within this genus are in the same range as among different genera.
The taxa included in the alpine Charpentieria itala–ornata–dyodon clade show a pan-Southern Alpine distribution, with C. dyodon isolated in Piemonte and Ticino, C. itala along central and eastern Southern Alps whereas C. ornata is present along the easternmost Southern Alps and northwestern Balkans. The taxa included in the C. stenzii subclade occupy the eastern Southern Alps, from eastern Trentino-Alto Adige, Friuli-Venezia Giulia, W Slovenia and SE Austria.
Despite the low support values, the COI and the ITS2 phylogenetic trees (Figs
The results presented by Scheel and Hausdorf (2012: 3799, 3801) and
The stenzioid subspecies of C. itala included in our genital anatomical investigations (clavata, variscoi, balsamoi, and lorinae) were first assigned to C. stenzii (Käufel, 1928) but later transferred to C. itala by
The list of the dissected material (at least two specimens per population) is found in the Table
Examined taxa of Charpentieria with information on shell type (stenzioid/nonstenzioid), availability of genetic data (DNA) and epiphallar formula.
Taxon | Specimens examined | Shell stenzioid/ nonstenzioid | DNA | Epiphallar formula |
---|---|---|---|---|
Charpentieria dyodon dyodon (S. Studer, 1820) | Trasquera di Iselle, Verbano, Piedmont, Italy. 750 m asl. 46°12'29.53"N, 08°12'46.18"E, I. Niero leg. and det. | - | Y | PP(ELP) |
Charpentieria dyodon alpina (Stabile, 1859) | Margone, Val di Lanzo, Usseglio, Torino, Piedmont, Italy. 1500 m asl, 45°14'04.89"N, 07°12'18.90"E, leg. and det. | - | N | PP(ELP) |
Charpentieria dyodon thomasiana (Küster, 1850) | Sentiero per Prà del Vecia, Piedicavallo, Biella, Piedmont, Italy. 1200 m asl. 45°41'32.37"N, 07°58'02.37"E, leg. and det. | - | N | PP(ELP) |
Charpentieria dyodon thomasiana (Küster, 1850) | Santuario di Oropa, Biella, Piedmont, Italy. 1100 m asl 45°37'49.01"N, 07°58'42.07"E, W. De Mattia and J. Macor leg. and det. | - | N | PP(ELP) |
Charpentieria ornata (Rossmässler, 1836) | Ogulin city centre, Croatia. 320 m asl. 45°16'2.55"N, 15°13'33.34"E, W. De Mattia and J. Macor leg. and det. | - | N | PP+ELP |
Charpentieria itala itala (G. von Martens, 1824) | San Donato, Barbarano Vicentino, Vicenza, Veneto, Italy. 300 m asl. 45°24'22.30"N, 11°31'11.93"E, W. De Mattia and J. Macor leg. and det. | nonstenzioid | Y | PP(ELP) |
Charpentieria itala albopustolata (De Cristofori & Jan, 1832) | Brescia, city castle walls, Lombardia, Italy. 200 m asl. 45°32'29.87"N, 10°13'28.37"E, W. De Mattia and J. Macor leg. and det. | nonstenzioid | N | PP(ER+ELP) |
Charpentieria itala allatollae (Käufel, 1928) | Albergo alla Tolla, Val Ampola, Storo, Trentino-Alto Adige, Italy. 720 m asl. 45°51'7.62"N, 10°38'04.51"E, W. De Mattia and J. Macor leg. and det. | nonstenzioid | N | PP(ELP) |
Charpentieria itala baldensis (Strobel, 1851) | Naole, San Zeno di Montagna, Verona, Veneto, Italy. 900 m asl. 45°38'43.67"N, 10°44'15.86"E, I. Niero leg. and det. | nonstenzioid | Y | ER+PP+ELP |
Charpentieria itala balsamoi (Strobel, 1850) | Loc. Galleria, Bracca, Val Serina, Bergamo, Lombardia, Italy. 45°49'24.49"N, 09°43'06.73"E, W. De Mattia and J. Macor leg. and det. | stenzioid | N | PP(ER(ELP)) |
Charpentieria itala clavata (Rossmässler, 1836) | Ballabio, Lecco, Lombardia, Italy. 710 m asl. 45°54'25.96"N, 09°25'55.16"E, W. De Mattia and J. Macor leg. and det. | stenzioid | N | PP(ELP) |
Charpentieria itala latestriata (Küster, 1850) | Bedulita, Bergamo, Lombardia, Italy. 625 m asl. 45°47'16.15"N, 09°32'42.26"E, W. De Mattia and J. Macor leg. and det. | nonstenzioid | N | PP(ER+ELP) |
Charpentieria itala lorinae (Gredler, 1869) | Val di Lorina, Storo, Brescia, Lombardia, Italy. 600 m asl. 45°50'35.28"N, 10°37'12.47"E, W. De Mattia and J. Macor leg. and det. | stenzioid | Y | ER+PP(ELP) |
Charpentieria itala zalloti nom. nov. | Udine, Friuli-Venezia Giulia, Italy. 140 m asl. 46°03'53.37"N, 13°14'12.35"E, L. Anzil leg., W. De Mattia det. | nonstenzioid | N | PP(ER(ELP)) |
Charpentieria itala zalloti nom. nov. | Udine, Friuli-Venezia Giulia, Italy. 140 m asl. 46°03'53.37"N, 13°14'12.35"E, L. Anzil leg., W. De Mattia det. | nonstenzioid | N | PP(ER(ELP)) |
Charpentieria itala serravalensis (Nordsieck, 1963) | Serravalle near Vittorio Veneto, Veneto, Italy. 500 m asl. 46°00'04.46"N, 12°17'11.13"E, W. De Mattia and J. Macor leg. and det. | nonstenzioid | nonstenzioid | PP(ER(ELP)) |
Charpentieria itala trepida (Käufel, 1928) | Forte Cima Ora, Anfo, Brescia, Lombardia, Italy. 1500 m asl. 45°47'50.57"N, 10°28'03.60"E, W. De Mattia and J. Macor leg. and det. | stenzioid | N | PP(ELP) |
Charpentieria itala triumplinae Nardi, 2011 | Cima Caldoline, Lavenone, Brescia, Lombardia, Italy. 1700 m asl. 45°48'03.54"N, 10°24'20.12"E, G. Nardi leg. and det. | stenzioid | N | PP(ER(ELP)) |
Charpentieria itala variscoi (Pini, 1883) | Lavina, Val Taleggio, Bergamo, Lombardia, Italy. 650 m asl. 45°53'11.73"N, 09°33'55.46"E, W. De Mattia and J. Macor leg. and det. | stenzioid | N | PP(ER+ELP) |
Charpentieria stenzii stenzii (Rossmässler, 1836) | San Romedio, Trento, Italy. 730 m asl. 46°22'03.83"N, 11°06'31.84"E, W. De Mattia and J. Macor leg. and det. | stenzioid | N | EP(ELP) |
Charpentieria stenzii butoti (Bank, 1987) | Fai della Paganella, Trento, Trentino-Alto Adige, Italy. 600 m asl. 46°12'57.04"N, 11°04'22.88"E, W. De Mattia and J. Macor leg. and det. | stenzioid | N | EP(ELP) |
Charpentieria stenzii cincta (Brumati, 1838) | Bosplans, Pordenone, Friuli-Venezia Giulia, Italy. 500 m asl. 46°11'58.11"N, 12°38'5.24"E, W. De Mattia and J. Macor leg. and det. | stenzioid | Y | EP(ELP) |
Charpentieria stenzii cincta (Brumati, 1838) | Val Resia, sella di Carnizza, Tarvisio, Friuli-Venezia Giulia, Italy. 400 m asl. 46°20'15.55"N, 13°19'17.11"E, W. De Mattia and J. Macor leg. and det. Mattia leg. and det. | stenzioid | N | EP(ELP) |
Charpentieria stenzii faueri (Bank, 1987) | Grotte di Oliero Valstagna, Vicenza, Veneto, Italy. 800 m asl. 45°50'47.79"N, 11°40'04.24"E, W. De Mattia and J. Macor leg. and det. | stenzioid | N | EP(ELP) |
Charpentieria stenzii letochana (Gredler, 1874) | Val Fonda, Misurina, Belluno, Veneto, Italy. 1500 m asl. 46°36'27.42"N, 12°12'20.01"E, W. De Mattia and J. Macor leg. and det. | stenzioid | Y | EP(ELP) |
Charpentieria stenzii nordsiecki Fauer, 1991 | Cison di Val Marino, Treviso, Veneto, Italy. 250 m asl. 45°58'05.53"N, 12°08'52.60"E, W. De Mattia and J. Macor leg. and det. | stenzioid | N | no pseudopapilla |
Charpentieria stenzii paroliniana (De Betta & Martinati, 1855) | Loc. La Goccia, Foza, Valstagna, Vicenza, Veneto, Italy. 800 m asl. 45°52'29.20"N, 11°38'59.73"E, W. De Mattia and J. Macor leg. and det. | stenzioid | N | EP(ELP) |
Charpentieria stenzii westerlundi (Nordsieck, 1993) | Val Fiscalina, Croda Rossa, Sesto, Bolzano, Trentino-Alto Adige, Italy. 1500 m asl. 46°38'02.83"N, 12°21'42.79"E, I. Niero leg. and det. | stenzioid | N | EP(ELP) |
The present anatomical investigation indeed generated consistent results that are congruent with the molecular genetic outcomes.
Charpentieria itala–ornata–dyodon and C. stenzii groups of taxa show the same external overall shape of the distal genital organs (Figs
The genus Charpentieria was introduced by
Boettger, 1877, introduced the genus Itala as “Gruppe der itala” (type species: Delima itala = Charpentieria itala), including all the alpine non-stenzioid taxa, excluding C. dyodon but including two stenzioid taxa: C. stenzii paroliniana and C. itala clavata. He also included two taxa from the Balkans: Delima conspersa [= Strigilodelima conspersa (L. Pfeiffer, 1848)] and its synonym Delima platystoma Küster, 1850. As for Tirolica, the recent results by
He introduced a new subspecific taxon, C. itala serravalensis (1963a: 175), but depicted its external anatomy of the genital organs [Nordsieck, 1963b: 184–185, Abb. 47 itala serravalensis, Serravalle (präp. 109)] with the same genital drawing provided for C. itala itala in Nordsieck, (1963a: 97, fig. 11a, b). The shell description, genital organs and measures provided for C. itala serravalensis (
Nordsieck cites C. itala itala (1963a) but unexpectedly,
Charpentieria itala was described by G. von
Charpentieria dyodon is the name-bearing species of the genus Charpentieria. It comprises five subspecies [C. dyodon dyodon (Studer, 1820; C. dyodon alpina (Stabile, 1859); C. dyodon paulucciana (Pollonera, 1885); C. dyodon studeri (Pini, 1885) and C. dyodon thomasiana (Küster, 1850)]. These taxa are distributed over an area that includes the western Southern Alps of Piemonte, Italy (from Dora Riparia River to Lago Maggiore) and the South Switzerland (Ticino and Valais). This species is not sympatric with any other Charpentieria species.
In our COI tree only the nominate subspecies Charpentieria d. dyodon was included (Figs
Three out of five subspecies of C. dyodon were anatomically investigated (Table
The four populations of Charpentieria dyodon included for dissection, revealed a very stable morphology of the genital organs (Figs
This parapseudopapilla represents an intermediate form between the simple structure of Charpentieria itala and the hemipapilla (HP) of Charpentieria stenzii. In the normal PP the transition stretch E is relatively wide and the counter wall opposite of the PP is represented by the simple P-E transition wall.
Charpentieria dyodon s. l. has a limited distribution, restricted to northern Piedmont and adjacent Swiss territories. The nominate subspecies is restricted to the surroundings of Iselle (Piedmont) and adjacent Swiss localities.
The FO is slightly longer than the V. The FDBC is shorter the BC+SDBC (SDBC+BC). The BC+SDBC is club-like and longer than the V with no distinction between the SDBC and the BC. The apex is round. The D is slightly longer than the BC+SDBC, thinner with a small but round apex. The V is short, cylindrical and large in diameter. The PC is much longer than the V. The PR is short and robust. The E is shorter than the P and very thin.
The A and the P are smooth, with a very fine granulated sculpturing. The V shows many smooth longitudinal pleats. The smooth PP is simple, big, conical with a pointed apex. Its base is partially connected to the ELP and transversally extends along the transition wall, narrowing the transition passage. The epiphallar formula is: PP(ELP). The E shows two main longitudinal moderately fringed pleats that proximally fade before the VD.
Charpentieria dyodon alpina is found in scattered populations along the Val di Lanzo, NW of Torino (Piedmont).
Italy, Piedmont, Torino, Margone, Val di Lanzo, Usseglio. 1500 m asl, 45°14'4.89"N, 07°12'18.90"E, leg. and det., 2 dissected spm.
(Fig.
(Fig.
Italy, Piedmont, Biella, Santuario di Oropa. 1100 m asl 45°37'49.01"N, 07°58'42.07"E, W. De Mattia and J. Macor leg. and det., 3 dissected spm. Italy, Piedmont, Biella, Sentiero per Prà del Vecia, Piedicavallo. 1200 m asl, 45°41'32.37"N, 07°58'2.37"E, leg. and det., 2 dissected spm.
Charpentieria dyodon thomasiana is found in the surroundings of Oropa and Piedicavallo, surroundings of Biella (Piedmont).
(Fig.
(Fig.
The Southern Alpine C. itala is distributed from Como Lake to the Garda Lake, Trentino, Veneto (including Colli Berici and Colli Euganei) and the Pre-Alps of Friuli.
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 C. itala were anatomically investigated, revealing a common overall arrangement of the genital organs. The pseudopapilla can be either smooth or markedly wrinkled and occupies almost the whole penial volume.
Charpentieria ornata has a wide distribution that includes Sudetes, southeastern Alps (Slovenia) and the western Dinarids (
Croatia, Ogulin city centre, 320 m asl, 45°16'2.55"N, 15°13'33.34"E, W. De Mattia and J. Macor leg. and det. 3 dissected spm.
(Fig.