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) Author De Mattia, Willy https://orcid.org/0000-0002-0056-467X Central Research Laboratories, of Natural History Museum Vienna, Burgring 7, 1010 Vienna, Austria willy.demattia@icgeb.org Author Reier, Susanne Central Research Laboratories, of Natural History Museum Vienna, Burgring 7, 1010 Vienna, Austria Author Haring, Elisabeth Central Research Laboratories, of Natural History Museum Vienna, Burgring 7, 1010 Vienna, Austria text ZooKeys 2021 2021-12-14 1077 1 175 http://dx.doi.org/10.3897/zookeys.1077.67081 journal article http://dx.doi.org/10.3897/zookeys.1077.67081 1313-2970-1077-1 C28AD65A76F242CFBED7DFB3702CABCE 734088641608531C8E2CC69397B000ED Papillifera Hartmann, 1842 Remarks. The genus Papillifera was used as the outgroup genus in the molecular genetic analysis (Figs 4 - 6 ). The general features of the genus Papillifera were summarised by Nordsieck (2013a : 8) who provided two lists of synonyms without any explanation or justification. Papillifera includes two species: P. papillaris and P. solida ( Margelli 2014 ; MolluscaBase 2021). The current known distribution and status of its (sub)specific taxa is far of being satisfactorily assessed, due to the unknown or inadequately evaluated validity of many old names. Papillifera is clearly distinguishable from Siciliaria s. s., Sicania and Stigmatica by its G-type clausiliar apparatus and the general shell features (e.g., the sutural dark band, see Fig. 68 .1-68.7). In our study, we dissected the Papillifera samples that underwent molecular genetic investigations, which are shown in Fig. 67 .1-67.12. The genital anatomical drawings of Papillifera papillaris papillaris (O. F. Mueller , 1774) (Fig. 67 .1-67.2), Papillifera papillaris affinis (Philippi, 1836) (Fig. 67 .3-67.8), Papillifera papillaris transitans (Paulucci, 1878) (Figs 67 .9-67.10), and Papillifera solida solida (Draparnaud, 1805) (Fig. 67 .11-67.12) show a stable genital structure in dimensions, ratios among the main genital parts and internal features. Nordsieck (2013a : 8) highlights that the anatomy of the genital organs of Papillifera is the same as Siciliaria s. l. Following our anatomical investigations, this statement must be rejected, as the general internal architecture of the PC (P+E and transition) revealed to be substantially different from Siciliaria / Sicania and Stigmatica . In detail, Papillifera lacks the ET and the connecting longitudinal pleats between PP and the ELP. The epiphallar formula of Papillifera is: PP+ELP. The sculpturing of the internal walls of the penis are mainly smooth or with very soft longitudinal pleats. The pseudopapilla is elongated to rounded, smooth and unrooted, directly emerging from the P-E transition walls. The E shows two smooth ELP. Table 11. Examined taxa of Papillifera with information on availability of genetic data (DNA) and description of the variability penis-epiphallus transition (epiphallar formula).
Taxon Examined specimens Epiphallar formula DNA
Papillifera papillaris papillaris (O.F. Mueller , 1774) Vicovaro, Latina, Lazio, Italy. 320 m asl. 42°01'04.58"N , 12°54'19.30"E , H. Nordsieck leg. and det. PP+ELP Y
Papillifera papillaris affinis (Philippi, 1836) Jato Antica, Monreale, Sicily, Italy, 830 m asl, 37°58'2.36"N , 13°11'45.19"E , W. De Mattia and J. Macor leg. and det. PP+ELP Y
Papillifera papillaris affinis (Philippi, 1836) western part of Monte Kumeta toward Jato Antica, Monreale, Sicily, Italy, 630 m asl, 37°57'8.18"N , 13°13'14.57"E , W. De Mattia and J. Macor leg. and det. PP+ELP Y
Papillifera papillaris affinis (Philippi, 1836) Monte Bonifato, west side of the mountain, over the quarry, Alcamo, Sicily, Italy. 550 m asl, 37°57'16.92"N , 12°58'09.06"E , W. De Mattia and J. Macor leg. and det. PP+ELP Y
Papillifera papillaris transitans (Paulucci, 1878) road to the Norman Castle, Stilo, Reggio Calabria, Calabria, Italy. 320 m asl. 38°28'45.91"N , 16°27'48.05"E , H. Nordsieck leg. and det. PP+ELP Y
Table 12. New Siciliaria / Sicania checklist and comparison with lists provided by Nordsieck (2013b) and MolluscaBase eds. (2021) .
Nordsieck 2013b MolluscaBase 2021 De Mattia, Reier and Haring 2021
Siciliaria grohmanniana ( Rossmaessler , 1836) Charpentieria grohmanniana ( Rossmaessler , 1836) Siciliaria grohmanniana grohmanniana ( Rossmaessler , 1836)
- - Siciliaria grohmanniana addaurae ssp. nov. De Mattia, Reier and Haring
Siciliaria septemplicata (Philippi, 1836) Charpentieria septemplicata (Philippi, 1836) Siciliaria septemplicata (Philippi, 1836)
Siciliaria leucophryna (L. Pfeiffer, 1862) Charpentieria leucophryna (L. Pfeiffer, 1862) Siciliaria leucophryna (L. Pfeiffer, 1862)
Siciliaria calcarae ( Philippi 1844 ) Charpentieria calcarae (Philippi, 1844) Siciliaria calcarae calcarae ( Philippi 1844 )
Siciliaria calcarae belliemii (Brandt, 1961) Charpentieria calcarae belliemii (Brandt, 1961) Siciliaria calcarae belliemii (Brandt, 1961)
- - Siciliaria calcarae borgettensis ssp. nov. De Mattia, Reier and Haring
- - Siciliaria calcarae jatinensis ssp. nov. De Mattia, Reier and Haring
- - Siciliaria calcarae parajatinensis ssp. nov. De Mattia, Reier and Haring
- Charpentieria calcarae orlandoi Liberto, Reitano, Giglio, Colomba & Sparacio, 2016 Siciliaria calcarae orlandoi Liberto, Reitano, Giglio, Colomba & Sparacio, 2016
- - Siciliaria calcarae cruenta ssp. nov. De Mattia, Reier and Haring
Siciliaria ferrox (Brandt, 1961) Charpentieria ferrox (Brandt, 1961) Siciliaria ferrox (Brandt, 1961)
Siciliaria riberothi (Brandt, 1961) Charpentieria riberothi (Brandt, 1961) see Nordsieck (2013b)
Siciliaria tiberii tiberii (A. Schmidt, 1868) Charpentieria tiberii (A. Schmidt, 1868) Siciliaria tiberii tiberii (A. Schmidt, 1868)
- - Siciliaria tiberii armettensis ssp. nov. De Mattia, Reier and Haring
Siciliaria septemplicata alcamoensis Brandt, 1961 Siciliaria septemplicata alcamoensis Brandt, 1961 Siciliaria tiberii alcamoensis Brandt, 1961 comb. nov.
- Charpentieria septemplicata hemmeni Beckmann, 2004 -
Siciliaria tiberii scalettensis Beckmann, 2004 Charpentieria tiberii scalettensis Beckmann, 2004 Siciliaria tiberii scalettensis Beckmann, 2004
Siciliaria crassicostata (L. Pfeiffer, 1856) Siciliaria crassicostata (L. Pfeiffer, 1856) Sicania crassicostata (L. Pfeiffer, 1856) comb. nov.
Siciliaria eminens (A. Schmidt, 1868) Charpentieria eminens (A. Schmidt, 1868) Sicania eminens (A. Schmidt, 1868) comb. nov.
Siciliaria nobilis (L. Pfeiffer, 1848) Charpentieria nobilis (L. Pfeiffer, 1848) Sicania nobilis nobilis (L. Pfeiffer, 1848) comb. nov.
Siciliaria spezialensis (Nordsieck, 1984) Charpentieria spezialensis (Nordsieck, 1984) Sicania nobilis spezialensis (Nordsieck, 1984) comb. nov.; stat. nov.
Siciliaria scarificata (L. Pfeiffer 1856 ) Charpentieria scarificata (L. Pfeiffer 1856 ) Mauritanica scarificata (L. Pfeiffer 1856 ) comb. nov.
Figure 67. Papillifera papillaris papillaris (O.F. Mueller , 1774). Vicovaro, Latina 67.1 whole distal genital organs 67.2 internal distal part of genital organs. Papillifera papillaris affinis (Philippi, 1836). Jato Antica, San Giuseppe Jato 67.3 whole distal genital organs 67.4 internal distal part of genital organs. Monte Bonifato, Alcamo 67.5 whole distal genital organs 67.6 internal distal part of genital organs. Western slopes Monte Kumeta, Monreale 67.7 whole distal genital organs 67.8 internal distal part of genital organs. Papillifera papillaris transitans (Paulucci, 1878). Road to the Norman Castle, Stilo 67.9 whole distal genital organs 67.10 internal distal part of genital organs. Papillifera solida solida (Draparnaud, 1805), Matera 67.11 whole distal genital organs 67.12 internal distal part of genital organs. Figure 68. Papillifera papillaris papillaris (O.F. Mueller , 1774). Vicovaro, Latina 68.1 shell 68.2 clausiliar plate. Papillifera papillaris affinis (Philippi, 1836). Jato Antica, San Giuseppe Jato 68.3 shell. Monte Bonifato, Alcamo 68.4 shell. Western slopes Monte Kumeta, Monreale 68.5 shell. Papillifera papillaris transitans (Paulucci, 1878). Road to the Norman Castle, Stilo 68.6 shell. Papillifera solida solida (Draparnaud, 1805), Matera 68.7 shell. Figure 69. 69.1 Siciliaria grohmanniana grohmanniana ( Rossmaessler , 1836), Monte Pellegrino, Palermo 69.2 Siciliaria grohmanniana addaurae ssp. nov. Grotta dell'Addaura , Punta Priola 69.3 Limestone walls at Grotta dell'Addaura , Punta Priola 69.4 Siciliaria calcarae belliemii (Brandt, 1961), Castello Calatubo, Alcamo 69.5 Siciliaria calcarae parajatinensis ssp. nov., west side Monte Kumeta 69 . 6 Siciliaria tiberii armettensis ssp. nov., Grotta dei Puntali, Carini. Figure 70. 70.1 Siciliaria leucophryna (L. Pfeiffer, 1862), Grotta Conza, Palermo 70.2 Siciliaria tiberii scalettensis Beckmann, 2004 Portella Scaletta, Villagrazia 70.3 Limestone walls at Monte Cofano, Custonaci 70.4 Sicania crassicostata (L. Pfeiffer, 1856), comb. nov., Monte Cofano, Custonaci 70.5 Sicania nobilis spezialensis ( Nordsieck 1984 ), stat. nov., comb. nov. Macari. Figure 71. 71.1 Sicania nobilis nobilis (L. Pfeiffer, 1848), comb. nov., Monte Cofano, Tonnara Cofano 71.2 Limestone walls at Monte Cofano, Tonnara Cofano 71.3 Mauritanica scarificata (L. Pfeiffer, 1856), comb. nov., Marettimo.