A peer-reviewed open-access journal Zookeys 732: |—125 (2018) doi: 10.3897/zookeys.732.2 1677 #7Zo00Ke y S http:/ / ZOO keys -pen soft. net Launched to accelerate biodiversity research Revision of the genus-group Hystricella R. T. Lowe, 1855 from Porto Santo (Madeira Archipelago), with descriptions of new recent and fossil taxa (Gastropoda, Helicoidea, Geomitridae) Willy De Mattia'’, Marco T. Neiber?, Klaus Groh* | Natural History Museum Vienna, Burgring 7, 1010 Vienna, Austria 2 International Centre for Genetic Engineering and Biotechnology, Padriciano 99, 34149, Trieste, Italy 3 Centre of Natural History, Zoological Museum, University of Hamburg, Martin-Luther-King-Platz 3, D-20146 Hamburg, Germany 4 Hinter- bergstraffe 15, D-67098 Bad Diirkheim, Germany Corresponding author: Willy De Mattia (demattia@icgeb.org) Academic editor: /. Haase | Received 16 October 2017 | Accepted 29 November 2017 | Published 24 January 2018 http-//zoobank. ore/9995 702B-6 146-4BA 1-BB53-23DC9BA9650F Citation: De Mattia W, Neiber MT, Groh K (2018) Revision of the genus-group Hystricella R. T. Lowe, 1855 from Porto Santo (Madeira Archipelago), with descriptions of new recent and fossil taxa (Gastropoda, Helicoidea, Geomitridae). ZooKeys 732: 1-125. https://doi.org/10.3897/zookeys.732.21677 Abstract The genus-group Hystricella R. T. Lowe, 1855 is revised on the basis of conchological, anatomical and genetic characteristics. A new genus Wollastonia gen. n., two recent species, W. jessicae sp. n. and W. klaus- grohi sp. n., and one recent subspecies, W. jessicae monticola ssp. n. are described as new to science, as well as five fossil taxa, H. microcarinata sp. n., W. beckmanni sp. n., W. falknerorum sp. n., W. ripkeni sp. n., and W. inexpectata sp. n. For Helix vermetiformis R. T. Lowe, 1855, H. leacockiana Wollaston, 1878, H. oxytropis R. T. Lowe, 1831, H. duplicata R. T. Lowe, 1831 and H. oxytropis var. 8 subcarinulata Wollaston, 1878 lectotypes are designated. For the taxa Helix bicarinata G. B. Sowerby I, 1824, Helix bicarinata var. {§ aucta Wollaston, 1878 and Discula bulverii W. Wood, 1828 neotypes are selected. The taxa aucta and subcarinulata are elevated to specific rank. For the hitherto monospecific (sub-) genus Callina R. T. Lowe, 1855 it is shown that it is not closely related to the genus Discu/a but to the Aystricella-group and its ge- neric rank is confirmed. The taxon D. bulverii W. Wood, 1828 is transferred from the genus Discula s. str. to the genus Callina. A further fossil taxon C. waldeni sp. n. is described as new to science. Copyright Willy De Mattia et al. 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. 2 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) Kurzfassung Die Gattungsgruppe Hystricella R. T. Lowe, 1855 wird anhand von conchologischen, anatomischen und genetischen Merkmalen revidiert. Eine neue Gattung Wollastonia gen. n., zwei rezente Arten, W. jessicae sp. n. und W. klausgrohi sp. n. sowie eine rezente Unterart, W. jessicae monticola ssp. n., werden neu be- schrieben, wie auch ftinf fossile Taxa, H. microcarinata sp. n., W. beckmanni sp. n., W. falknerorum sp. n., W. ripkeni sp. n., und W. inexpectata sp. n. Fiir Helix vermetiformis R. T. Lowe, 1855, H. leacockiana Wollaston, 1878, H. oxytropis R. T. Lowe, 1831, H. duplicata R. T. Lowe, 1831 und H. oxytropis var. & subcarinulata Wollaston, 1878 werden Lectotypen designiert, fiir die Taxa Helix bicarinata G. B. Sowerby I, 1821, Helix bicarinata var. 8 aucta Wollaston, 1878 und Discula bulverii W. Wood, 1828 Neotypen festgelegt. Die Taxa aucta und subcarinulata werden in einen spezifischen Rang erhoben. Es wird gezeigt, dass die bisher monospezifische (Unter-) Gattung Callina R. T. Lowe, 1855 nicht mit der Gattung Dis- cula, sondern mit der Hystricella-Gruppe naher verwandt ist und im Gattungsrang bestatigt. Das Taxon D. bulverii W. Wood, 1828 wird aus der Gattung Discula s. str. zu Callina gestellt und ein weiteres fossiles Taxon C. waldeni sp. n. in dieser Gattung neu beschrieben. Resumo No presente trabalho o grupo genérico Hystricella R. T. Lowe, 1855 é revisto com base em critérios mor- folégicos, anatémicos e genéticos. E descrito um novo género, Wollastonia gen. n., duas novas espécies atuais, W. jessicae sp. n. e W. klausgrohi sp. n. e uma nova subespécie também atual, W. jessicae monticola ssp. n. Sao também descritas neste novo género cinco novas espécies fésseis: H. microcarinata sp. n., W. beckmanni sp. n., W. falknerorum sp. n., W. ripkeni sp. n. e H. inexpectata sp. n. Sao designados lectétipos para as espécies, Helix vermetiformis R. T. Lowe, 1855, H. leacockiana Wollaston, 1878, H. oxytropis R. T. Lowe, 1831, H. duplicata R. T. Lowe, 1831 e H. oxytropis var. & subcarinulata Wollaston, 1878. Sao designados nedtipos para as taxones Helix bicarinata G. B. Sowerby I, 1824, Helix bicarinata var. & aucta Wollaston, 1878 e Discula bulverii W. Wood, 1828. Os taxa aucta e subcarinulata sao elevados a categoria de espécie. E demonstrado que o (sub) género, até agora monotipico, Callina R. T. Lowe, 1855, nao esta relacionado com o grupo genérico Discula, mas sim com o grupo genérico Hystricella e o seu estatuto de género é confirmado. O taxon D. bulverii W. Wood, 1828 é transferido do género Discula s. str. para o género Callina e é descrita uma nova espécie fossil adicional, C. waldeni sp. n. Keywords Callina, Discula, Hystricella, Wollastonia gen. n., Macaronesia, Porto Santo (Madeira Archipelago), recent, Quaternary, phylogeny Contents Tito Gh We OIE ort eee cent sears et esr Doe li Aes ck Bieler at Aree rea Rc eteatee ne Reca 3 IVigreiialls are hm Gl Gels, 04.2.0 stehag saath Peli eeu Tote A Moa Nett et ALE Sug alia ae 6 Gallectine- and. preseiyationcor Sar plete 45 5..0M ible as AOR ihe eda Sakaies cl 6 Morphologicalnivesti SattOnis. iat. cchet dean sc het ne atanucdptaaninis BosBoan con egecatsivnenistsSecsse 4 Molecular data preparation and phylogenetic analyses.........eeeeeseeseeeeseeeeeeeees 7 Cr GME STIOTT A Cre My AMIS: 9 We «cide aul wetleSenesn «ibaitaneton ee ceaseeaiaaseanieatsaueiee Bensadeer dee 13 Morpholoeical abbreviation sa iatecessxsarcanrosetsursepn sap onesane ncunticowndae' sede tat asesantlounnstiene 13 Revision of the genus-group Hystricella R. 77 Lowe, 1855 from Porto Santo... 2 REGU ESS AIG CLISCIESSTOLIT, ot eax ces ate aes ee ee ete ee see etre Es adh Aenactcamte ses 14 Phir lOgeneticra ma lySe92 xchteidadbescastes dees eseaia wiced bonnaleoueinctTbau ens ame ienan aman easels 14 SYSTEMATIC. BECOUMM oe. - eve cuyvendaenn s+ cevnmecaevavgcn vane ewes cacneynewrae eRe TRE. SAO eS 19 PIVSERICCH ANS. “LAGE WCDI ackls ches bihovccsaboa tact dstus phoct ss doxhluahous ta AduMig vkve tans 20 Hystricella bicarinata (G. B. Sowerby 1, 1824)... .eceeceeeeeseeeseeseeeeeeees 27 + Alystricella aucta (Wollaston, 1878), stat. 1. ....s cesses ceesseeseeeseeaeeees 38 + Alystricella microcarinata De Mattia & Groh, sp. nu eeeeeeseseeseeseeeee 4] Elystriceliavecninwlata ds: EAWOwe VEST. mnasessvuldasnniennenniosronseritsn bese: 43 + Aystricella echinoderma (Wollaston, 1878)......scssseceeseeseeseeseceseeseeees 48 WOMASLONUE SONY ca ches vas cxascstet tia Ubasdocksislebadaxscntaauasaende. sigceateaet uses SatcGs 50 Wollastonia turricula (R. T. Lowe, 1831), comb. niece eeeeeeeeees 56 Wollastonia vermetiformis (R. T. Lowe, 1855), comb. 1. wee eeeeeeseeeeeees 63 + Wollastonia ripkeni De Mattia & Groh, sp. 0. .i.ceeeeeesecseeseeseeeseeaeeeee 66 + Wollastonia falknerorum Groh, Neiber & De Mattia, sp. no... 69 Wollastonia leacockiana (Wollaston, 1878), comb. novice eee rip) | Wollastonia beckmanni De Mattia & Groh, sp. 0... eeeeeeeeereeereeeeeeeee Ta Wollastonia jessicae jessicae De Mattia, Neiber & Groh, sp. nu... 79 Wollastonia jessicae monticola De Mattia, Neiber & Groh, ssp. n. ......... 84 Wollastonia klausgrohi De Mattia & Neiber, sp. ne vice eeeeseeeeseeseeeee 87 Wollastonia oxytropis (R. T. Lowe, 1831), comb. Dh... .eceeeeeseseeseeeeees 92 + Wollastonia subcarinulata (Wollaston, 1878), comb. n., stat. n.......... 98 + Wollastonia inexpectata De Mattia & Groh, sp. n. ..eeeeeeeseeeeseeeees 100 CAE GR PM MG WE MS Tt seks oh nckae hike pe ede dauk Deh aehu pM ave daakt a tacste stake eae 103 Cai lana rotilie UR A MeO WE AVS Be ohn geste oboe benonptbon- oftltinasenedesbonweeal 106 {, Callind waldeni Grol See Mattia, sp. a1 2.. LAS. -2e.2. di enkecee ont Ala Callina-bulverit: OW: WOOd.. 138.2 BY iCOMDS Merc sine tecceaset Mieco owls: 113 PROKTIOWG EU CORACINES.. sues sien ote cansaene Sadi enileshteteadett aruoeiateithn che euesiaedeaiisRneapttdhbtee 120 FREER EIIGeS 5, nce Mace ered deiner tea vinsetie Sel tt etme a ees Rates PRs gy AR Pa ER od So Oe ly de 121 Introduction The island of Porto Santo (42.17 km?) is the northernmost island of the Madeiran Archipelago (Fig. 1), situated in the eastern Atlantic Ocean off the northwest African coast. As with the other islands of the archipelago, i.e., the island of Madeira and the Ilhas Desertas, Porto Santo is of volcanic origin, with its oldest subaerial deposits dated at 14.2 to 13.1 Ma before present to the Middle Miocene (Geldmacher et al. 2000; Ribeiro and Ramalho 2010). Thus, there was nearly three times as much time on Porto Santo for the evolution of an endemic land snail fauna compared to the island of Madeira with an estimated age of 5.2 to 4.6 Ma before present (Geldmacher et al. 2000; Ribeiro and Ramalho 2010) or to the Ilhas Desertas which are with approximately 3.6 Ma 4 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) (Geldmacher et al. 2000), even younger than Madeira. The finding of a land snail from 20 to 13 Ma old Neogene deposits of Ilhéu de Cima, a satellite islet off southwestern Porto Santo, which Groh (1984) referred to Caseolus (Leptostictea) sp., suggests a relatively early colonization of Porto Santo by terrestrial gastropods belonging to lineages which are still present on the island today. Eleven genera and subgenera are currently considered to be endemic to Porto Santo incl. offshore islets: Leiostyla (Craticula) R. T. Lowe, 1852 (Lauriidae), Amphorella (Fusillus) R. T. Lowe, 1852 (Ferussaciidae), Serratorotula Groh & Hemmen, 1986, Lemniscia R. T. Lowe, 1855, Pseudocampylaea L. Pfeiffer, 1877, Callina R. T. Lowe, 1855, Discula (Mandahlia) Forcart, 1965 and Aystricella R. 'T. Lowe, 1855 (Geomitridae), Leptaxis (Katostoma) R. 'T. Lowe, 1855 (Hygromiidae), Lampadia Albers, 1854 and Idiomela 'T. D. A. Cockerell, 1921 (Helicidae). In total, 91 of the hitherto 102 known species and subspecies (c. 89%) recorded from Porto Santo are endemic (67 recent and 24 fossil taxa) resulting in a density of endemism of 2.13 taxa per square kilometre. Taking into consideration that most (sub-) species today only exist in the higher eastern part of the island, this value increases to 4.7 taxa per square kilometre, a value that is unique when measured on a global scale. ‘This is especially noteworthy, since on Porto Santo, contrary to Madeira, the vertical differentiation of habitats was greatly reduced due to massive erosion in the past million years as evidenced by deeply cut ravines, relatively thick Aeolian deposits in the island’s central part (Ribeiro and Ramalho 2010) and a large marine abrasion platform surrounding the island, with abrasion values of 12 cm per year (Lietz and Schwarzbach 1971). The documentation of the land snail diversity of Porto Santo started in 1824 when George Brettingham Sowerby I described the first four land snails from that island, among them Helix bicarinata, the type species of the genus Hystricella. The first more in-depth investigations of the terrestrial mollusc fauna of the Madeiran Archipelago were carried out by the clergyman Richard Thomas Lowe who visited the archipelago in 1828 for the first time. The results of his colleting efforts in that year were published in 1831 in his important synopsis of the fauna “Primitiz Faune et Flore Maderz et Portus Sancti”, in which many of the land snails from Porto Santo considered as valid taxa today were named and described. From 1832 until 1854 R. T. Lowe served as Anglican chaplain in the archipelago and the material that he collected during these years formed the basis for twelve additional publications on the malacofauna of the archipelago (see Bank et al. 2002). During this period only few additional taxa had been described, e.g., by Férussac (1832) and Deshayes (1835, 1850) in Férussac and Deshayes (1819-1851), Férus- sac (1835), Pfeiffer (1846), and Albers (1854). Anténio da Costa de Paiva (Bardo de Castelo de Paiva) described a number of species in 1866 and published a comprehen- sive thesis on the Madeiran land snail fauna in 1867, presenting an overview of the knowledge of the time. Another important contribution towards an inventory of the malacofauna of the Madeira Archipelago was published by Thomas Vernon Wollaston under the title “Testacea Atlantica” in 1878, in which he summarised all the knowledge of the time on the non-marine molluscs of the Mid-Atlantic islands including the results of his five Revision of the genus-group Hystricella R. T) Lowe, 1855 from Porto Santo... 5 & lIhéu de Fora _ : & Baixa do Meio Baixa dos Barbeiros Q <> Ilhéu das Cenouras lIhéu de Cima llhéu de Ferro oe IIhéu de Baixo ou da Cal Figure |. Map of Porto Santo with offshore islets and map of the Madeiran Archipelago in the Atlantic Ocean. extended expeditions to the Madeiran Archipelago between 1847 and 1855. In this book, T. V. Wollaston listed ten taxa under the section Hystricella of the genus Helix Linnaeus, 1758, namely H. bicarinata var. aucta Wollaston, 1878, H. bicarinata, H. echinoderma Wollaston, 1878, H. echinulata R. 'T. Lowe, 1831, H. leacockiana Wollas- ton, 1878, H. turricula R. T. Lowe, 1831, H. oxytropis R. T. Lowe, 1831, H. turricula var. pererosa Wollaston, 1878, H. oxytropis var. subcarinulata Wollaston, 1878, and H. vermetiformis R. T. Lowe, 1855. Subsequent researchers have not added any new taxa to the Aystricella-group, but evaluated its constituent taxa supra-specifically differently, without any substantial taxonomic basis, mostly placing them at a sub-generic level in the genus Discula R. T. Lowe, 1852. Furthermore, the status of many nominal taxa was, and still remains, controversial, ranging from treatments as mere varieties to subspecies or species. The two most up-to-date publications listed nine (Bank et al. 2002), viz. seven (Seddon 2008) specific and subspecific taxa in the genus Aystricella, the latter ignoring (except two) described fossil taxa from Quaternary deposits. Groh et al. (2009) published ad- ditions and corrections to Seddon (2008), including also three fossil taxa belonging to the genus Hystricella. Anatomical investigations are currently lacking for most species currently placed in the genus Hystricella. The only exception is . turricula, the anatomy of which was described by Watson (1923) in detail. Mandahl-Barth (1950) provided additional anatomical data for many geometrid taxa, but only repeated the results of Watson (1923) with regard to 6 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) Fystricella because of the lack of preserved soft parts. Although Seddon (2008) provided distribution maps and added comments on the known distribution in the IUCN Red List (Seddon 201 1a-e) for five recent taxa currently classified in Hystricella, some of the records presented in those publications seem to be erroneous or highly unlikely and ought to be critically revised. In the present contribution, we present a comprehensive revision of the species currently assigned to the genus Aystricella including, aside from conchological com- parisons, anatomical descriptions of all recent taxa (except for W. vermetiformis) as well as molecular phylogenetic analyses on the basis of mitochondrial as well as nuclear DNA sequences. Distribution data for all taxa are critically revised and stratigraphic as well as biogeographic relations are discussed. Materials and methods Collecting and preservation of samples Recent, living snails were collected by hand. Although specimens were quite abun- dant in many populations (often reaching densities of more than 50 specimens per square meter), we limited sampling to a maximum of 15—20 specimens per site for ethical and conservation reasons. The developmental stage of each specimen was carefully checked during the collecting process and all juvenile and subadult speci- mens were left in the field. Half of the selected specimens were immediately fixed in absolute ethanol after collection in order to preserve the DNA for molecular investigations. [he remaining specimens were drowned in water for 6-10 hours and subsequently fixed in 80% buffered ethanol. This was done to ensure that most of the distal parts of the snail’s body (foot, head, and distal genitalia) were stretched out from the shell, which facilitates dissection for morphological investigations. Two weeks after the first fixation, the ethanol solution was changed to remove the pre- cipitate that mostly consists of mucus and denatured proteins. In addition to living specimens, empty shells from recent populations were collected to obtain a better impression of the shell morphology and its intra-specific as well as intra-population variability. Most of the subfossil to fossil material was collected by hand. Fossiliferous depos- its were identified directly in the field or with the help of a geological map. At some sites with aeolinite deposits sediment was collected and immediately sieved in the field in order to separate the sandy part from the coarser portion of the sample that was expected to contain fossil shells. Larger samples of strongly cemented material were carefully crushed in the field. The smaller pieces were brought to the laboratory where they were soaked in a 2% solution of hydrogen peroxide (H,O,) and then rinsed with boiling water to separate the shells and to remove any agglutinations and encrustations adhering to them. ‘The shells cleaned in this way were again washed with water and finally dried. Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... 7 Morphological investigations Shells were photographed with a Canon EOS camera equipped with 60 mm macro lenses mounted on a Kaiser microslide frame for multi-image stacking. The shells of all the dissected specimens and additional empty shells were photographed in front, bot- tom, side, and top view. Maximum shell diameter (D) and maximum shell height (H) were measured using a calliper. Additional measurement, i.e., height of the first (body) whorl (FW), peristomal angle (PA), diameter of umbilicus (DU), standardised number of tubercles on the shell surface (see Fig. 2), and the number of whorls of the protoconch, the teleoconch and their sum were taken using an image editing software. Mean values, standard deviations as well as D/H and FW/H ratios were calculated using MS Excel. Dissections and examination of the anatomy were performed under a Zeiss stereo- scope with a ring LED illumination apparatus, connected to a digital, high-resolution camera and a camera lucida. The genitalia were isolated from the rest of the body, usually after crushing the shell, using very fine and pointed micro-tweezers (Dumostar Biology 55) in a Petri dish with the bottom filled with black parafhin. The genitalia were fixed with very fine steel micro pins (commonly used for the preparation of microlepidopteran specimens in entomology). The internal features of genitalia (including cross and longi- tudinal sections) were examined after dissection with micro tweezers or by a pair of bio- logical micro scissors (Aesculap OC series). Measurements were taken using a millimetric measurement scale. All examined genitalia were photographed in different positions (40 to 50 high-resolution images depicting all relevant anatomical features) in order to create an image database. Drawings were prepared by accurately tracing the most representative digital images after contrast enhancement with a picture editing software. The drawings, instead of digital photos, were used here because photos of the genitalia do not usually al- low the reader to easily observe finer anatomical/morphological details of the soft tissues (such as small, fine pleats, striae, folds, papillae, ornamentations and diverticula). The drawings were performed when the samples were still displayed under the stereoscope in order to constantly check their morphology in case of doubtful situations. For the preparation of jaws and radulae individual buccal masses were removed and immersed in a 2 M potassium hydroxide (KOH) solution for 5 h before extracting the jaw and the radula, which were then preserved in 70% ethanol for further investigations. Jaws and radulae were mounted and observed directly, i.e., without sputter coating, under a low vacuum SEM (Miniscope TM-1000, Hitachi High-Technologies, Tokyo). Molecular data preparation and phylogenetic analyses For molecular genetic analyses samples of foot muscle tissue stored in 70-96% ethanol from representatives of all recent (sub-) species (except 1. vermetiformis) and of species potentially belonging to the Aystricella group existing on the island of Porto Santo and of additional specimens of geomitrid species belonging to the nominal genera Discula, Caseolus R. T. Lowe, Callina and Serratorotula Groh & Hemmen, 1986 were used (see 8 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) whorls measurements peristomal angle Figure 2. Standardisation method for counting the tubercles on the shell surface. AB Maximum diam- eter CD Maximum height EF height of last whorl HG height of second whorl IC height of third whorl; peristoma angle, measured as the angle between the peristomal plane and the horizontal axis of the shell JK width of the last whorl measured along the maximum diameter (JK/2)? square on the lower surface of the last whorl delimiting the area for the count of tubercles. The centre of the square corresponds to the mid point between J and K and the length of its sides is JK/2. This method allows to compare different speci- mens and taxa as the area of the square depends on the shell’s overall dimensions and is always proportional (the tubercles that partially traverse the contour of the square were included in the count of tubercles). Table 1 for details). To root phylogenetic trees Cochlicella conoidea (Draparnaud, 1801) (Geomitridae, Geomitrinae, Cochlicellini) was included as outgroup (see Table 1). To- tal genomic DNA was extracted following a slightly modified version of the protocol of Sokolov (2000) as detailed in Scheel and Hausdorf (2012). Partial sequences of the mi- tochondrial cytochrome c oxidase subunit 1 (cox/) and (for a subset of samples repre- senting all genus-group taxa, see Table 1) of the nuclear rRNA gene cluster (including partial sequences of the 18S rDNA, the complete internal transcribed spacer 1 (ITS1), 5.8S rDNA and internal transcribed spacer 2 (ITS2) as well as partial sequences of the 28S rDNA) were amplified by polymerase chain reaction (PCR) using the primer pairs LCO1490 plus HCO2198 (Folmer et al. 1994) and Hyg18S_Fw (5’-GCT ACT ATC GAT TGA GCG GTT CAG-3’, J. Harl, pers. comm.) and Hyg28S_Rv: 5’°-CGT CCC ACA CAC CAC AGT T-3’, J. Harl, pers. comm.), respectively, following the protocols described in Neiber and Hausdorf (2015) and Harl et al. (2017), respec- tively. Both strands of the amplified products were sequenced at Macrogen Europe Revision of the genus-group Hystricella R. 77 Lowe, 1855 from Porto Santo... Table |. Vouchers, locality data and GenBank accession numbers of Geomitrinae samples used for mo- lecular analyses. Taxon Hystricella bicarinata Voucher Latitude number HY03_2 eae 16"N GenBank accession number Longitude coxl 16°18' SUSE MG575049 18S rDNA + ITS1 + 5.8S rDNA + ITS2 + 28S rDNA MG575189 Hystricella bicarinata Hystricella bicarinata Hystricella bicarinata Hystricella bicarinata TY03=.3 eae 16"N 16°18' SUSE MG575050 HY04_1 | 33°04'42"N | 16°20'00"W | MG575051 HY10_1 HY10_2 | 33°04'51"N 33°04'51"N 16°18'41"W | MG575052 16°18'41"W | MG575053 MG575190 Hystricella bicarinata HY10_3 | 33°04'51"N 16°18'41"W | MG575054 Hystricella bicarinata Hystricella bicarinata Hystricella bicarinata Hystricella bicarinata Ty a2eu 33°04'38"N 16°19'48"W | MG575055 HY32_2 | 33°04'38"N | 16°19'48"W | MG575056 HY33_1 HY33_2 | 33°04'52"N 33°04'52"N 16°19'26"W | MG575057 16°19'26"W | MG575058 Hystricella bicarinata HY33_3 | 33°04'52"N 16°19'26"W | MG575059 MG575191 Hystricella bicarinata Hystricella bicarinata Hystricella bicarinata HY41_1 33°05'04"N 16°19'21"W | MG575060 HY42_1 | 33°04'48"N | 16°18'31"W | MG575061 HY42_2 | 33°04'48"N 16°18'31"W | MG575062 Hystricella bicarinata HY42_3 | 33°04'48"N 16°18'31"W | MG575063 Hystricella bicarinata Hystricella bicarinata Hystricella bicarinata Hystricella bicarinata HY43_1 33°04'55"N 16°20'07"W | MG575064 HY43_2 | 33°04'55"N | 16°20'07"W | MG575065 HY45_2 | 33°04'42"N | 16°18'01"W | MG575066 HY45_3 | 33°04'42"N 16°18'01"W | MG575067 MG575192 Hystricella bicarinata HY47_1 33°05'01"N 16°19'14"W | MG575068 MG575193 Hystricella bicarinata Hystricella bicarinata Hystricella bicarinata Hystricella bicarinata HY47_2 | 33°05'01"N HY47_3 | 33°05'01"N vA N3005 | 33°04'44"N 13003 | 33°05'02"N 16°19'14"W | MG575069 16°19'14"W | MG575070 16°19'14"W | MG575071 16°17'59"W | MG575072 Hystricella bicarinata 3008 | 33°04'15"N 16°18'55"W | MG575073 Hystricella bicarinata Hystricella bicarinata Hystricella bicarinata |Z T 7, 13016 | 33°04'42"N 3009 | 33°05'04"N 3014 | 33°04'52"N 16°19'21"W | MG575074 16°19'26"W | MG575075 16°20'00"W | MG575076 Hystricella bicarinata 3018 | 33°04'32"N 16°19'38"W | MG575077 Hystricella bicarinata Hystricella bicarinata Hystricella bicarinata Hystricella bicarinata 4|Z 3019 | 33°04'42"N 16°18'01"W | MG575078 13020 | 33°04'55"N | 16°20'07"W | MG575079 3022 | 33°05'01"N | 16°19'14"W | MG575080 3023 | 33°04'51"N 16°18'41"W | MG575081 Hystricella bicarinata N 3029 | 33°04'48"N 16°18'31"W | MG575082 Hystricella bicarinata Hystricella bicarinata Sis SS SiS SiS Sis sss |S Z| zen ee | Ly N Z, 3047 | 33°04'44"N 16°17'59"W | MG575083 3049 | 33°05'34"N | 16°19'11"W | MG575084 Hystricella echinulata HY39_1 | 33°05'29"N | 16°18'14"W | MG575085 = Hystricella echinulata HY39_2 | 33°05'29"N | 16°18'14"W | MG575086 = Hystricella echinulata HY39_3 | 33°05'29"N | 16°18'14"W | MG575087 - Fiystricella echinulata HY40_1 | 33°05'38"N | 16°18'01"W | MG575088 = Hystricella echinulata HY40_2 16°18'01"W | MG575089 - Hystricella echinulata HY40_3 | 33°05'38"N 16°18'01"W | MG575090 10 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) GenBank accession number Taxon Voucher Tyusade+|| “oagierde 18S rDNA + ITS1 + number cox1 5.8S rDNA + ITS2 + 28S rDNA Hystricella echinulata MN3004 | 33°05'36"N MG575091 = Hystricella echinulata MN3006 | 33°05'29"N MG575092 - Fiystricella echinulata MN3010 | 33°05'38"N | 16°18'01"W | MG575093 - Hystricella bicarinata x echinulata | HY06_1 | 33°05'42"N MG575094 = Fiystricella bicarinata x echinulata | HY35_1 | 33°05'35"N | 16°18'25"W | MG575095 - Hystricella bicarinata x echinulata | HY35_2 | 33°05'35"N MG575096 7 Fiystricella bicarinata x echinulata | HY35_3 | 33°05'35"N | 16°18'25"W | MG575097 - Hystricella bicarinata x echinulata | HY36_1 | 33°05'57"N MG575098 - Fiystricella bicarinata x echinulata | HY36_2 | 33°05'57"N | 16°19'32"W | MG575099 - Hystricella bicarinata x echinulata | HY36_3 | 33°05'57"N MG575100 - Hystricella bicarinata x echinulata | HY37_1 | 33°06'09"N | 16°19'25"W | MG575101 - Hystricella bicarinata x echinulata | HY37_2 MG575102 MG575194 Hystricella bicarinata x echinulata | MN3007 | 33°05'35"N | 16°18'25"W | MG575103 - Hiystricella bicarinata x echinulata | MN3011 | 33°06'09"N MG575104 - Fiystricella bicarinata x echinulata | MN3015 | 33°05'57"N | 16°19'32"W | MG575105 - Fiystricella bicarinata x echinulata | MN3026 | 33°05'42"N MG575106 = Hystricella bicarinata x echinulata | MN3027 | 33°04'16"N | 16°18'53"W | MG575107 = Wollastonia jessicae jessicae HY05_1 | 33°03'44"N MG575108 MG575195 Wollastonia jessicae jessicae HY05_2 | 33°03'44"N | 16°19'35"W | MG575109 - Wollastonia jessicae jessicae EINO529:1/253,03. 44 N MG575110 - Wollastonia jessicae jessicae HY34_1 | 33°03'47"N | 16°19'41"W | MG575111 - Wollastonia jessicae jessicae HY34_2 | 33°03'47"N MG575112 - Wollastonia jessicae jessicae HY34_3 | 33°03'47"N | 16°19'41"W | MG575113 - Wollastonia jessicae jessicae MN3017 MG575114 - Wollastonia jessicae jessicae MN3021 | 33°03'44"N | 16°19'35"W | MG575115 - Wollastonia jessicae monticola HY03_1 | 33°04'16"N MG575116 - Wollastonia klausgrohi HY31_1 | 33°04'06"N | 16°18'52"W | MG575117 MG575196 Wollastonia klausgrohi HY31_2 | 33°04'06"N MG575118 = Wollastonia klausgrohi HY31_3 | 33°04'06"N | 16°18'52"W | MG575119 - Wollastonia klausgrohi MN3028 | 33°04'06"N MG575120 Bs Wollastonia leacockiana HYO1_1 | 33°02'57"N | 16°22'04"W | MG575121 = Wollastonia leacockiana HY01_2 MG575122| ___MG575197 Wollastonia leacockiana MN3002 | 33°02'39"N | 16°22'11"W | MG575123 = Wollastonia leacockiana MN3024 | 33°02'57"N MG575124 = Wollastonia oxytropis HY38_1 | 33°04'42"N | 16°17'56"W | MG575125 MG575198 Wollastonia oxytropis MN3001 | 33°04'43"N MG575126 = Wollastonia oxytropis MN3013 | 33°04'42"N | 16°17'56"W | MG575127 = Wollastonia oxytropis MN3048 | 33°04'41"N MG575128 = Wollastonia turricula MN3000 | 33°03'13"N | 16°16'48"W | MG575129 = Callina bulverii DI04_1 | 33°04'51"N MG575130 = Callina bulverii DI15_2 | 33°04'16"N | 16°18'53"W | MG575131 MG575199 Callina bulverii DI15_3 | 33°04'16"N MG575132 = Callina bulverii DI27_1 | 33°04'06"N | 16°18'52"W | MG575133 = Callina bulverii DI28_1 | 33°04'38"N | 16°19'48"W | MG575134 - Revision of the genus-group Hystricella R. 7’ Lowe, 1855 from Porto Santo... 11 GenBank accession number 18S rDNA + ITS1 + Voucher ‘ ; foee number cox 5.88 rDNA + ITS2 + 28S rDNA Callina rotula DI23_1 16°18'25"W | MG575135 - Callina rotula DI25_1 MG575136 = Callina rotula DI26_1 | 33°05'57"N | 16°19'32"W | MG575137 F Callina rotula DI31_1 | 33°06'09"N MG575138 = Callina rotula DI31_2 | 33°06'09"N | 16°19'25"W | MG575139 A Callina rotula DI35_1 | 33°04'48"N | 16°18'31"W | MG575140 7 Callina rotula DI35_2 MG575141 7 Callina rotula DI37_1 | 33°05'29"N | 16°18'14"W | MG575142 7 Callina rotula DI37_2 | 33°05'29"N MG575143 = Callina rotula DI44_1 | 33°05'38"N | 16°18'01"W | MG575144 = Callina rotula DI44.2 MG575145 MG575200 Callina rotula MN3051 | 33°04'48"N | 16°18'31"W | MG575146 v Discula (Discula) attrita DI05_1 | 33°04'44"N | 16°18'37"W | MG575147 MG575201 Discula (Discula) attrita DI05_2 | 33°04'44"N MG575148 " Discula (Discula) attrita DI05_3 | 33°04'44"N | 16°18'37"W | MG575149 x Discula (Discula) attrita DI12_1 | 33°02'57"N MG575150 5 Discula (Discula) attrita DI12_2 | 33°02'57"N | 16°22'04"W | MG575151 an Discula (Discula) attrita DI21_2 | 33°03'37"N MGS575152 Z Discula (Discula) attrita DI29_1 | 33°03'45"N | 16°17'57"W | MG575153 E Discula (Discula) attrita DI29_2 16°17'57"W | MG575154 e Discula (Discula) attrita DI29_3 | 33°03'45"N MG575155 = Discula (Discula) attrita DI29_4 | 33°03'45"N | 16°17'57"W | MG575156 = Discula (Discula) attrita DI29_5 | 33°03'45"N MG575157 2 Discula (Discula) attrita DI34_1 | 33°04'48"N | 16°18'31"W | MG575158 — Discula (Discula) attrita DI38_1 | 33°02'38"N MGS575159 a Discula (Discula) attrita DI38_2 | 33°02'38"N | 16°22'56"W | MG575160 = Discula (Discula) attrita DI43_2 | 33°04'01"N | 16°18'26"W | MG575161 = Discula (Discula) attrita MN3053 MG575162 Z Discula (Discula) calcigena DI42_1 | 33°04'16"N | 16°17'58"W | MG575163 MG575202 Discula (Discula) discina D108_1 | 33°05'48"N MG575164 A Discula (Discula) discina DI08_2 | 33°05'48"N | 16°18'54"W | MG575165 — Discula (Discula) discina D108_3 | 33°05'48"N MG575166 is Discula (Discula) discina DI08_4 | 33°05'48"N | 16°18'54"W | MG575167 . Discula (Discula) discina DIU6_1 MG575168 = Discula (Discula) discina DI16_2 MG575169 z Discula (Discula) discina DI16_3 | 33°05'25"N | 16°20'58"W | MG575170 2 Discula (Discula) discina DI16_4 | 33°05'25"N MGS575171 Z Discula (Discula) discina DI22_1 | 33°04'44"N | 16°18'37"W | MG575172 = Discula (Discula) discina DI22_2 | 33°04'44"N MG575173 = Discula (Discula) discina DI22_3 | 33°04'44"N | 16°18'37"W | MG575174 = Discula (Discula) discina DI22_4 | 33°04'44"N MG575175 = Discula (Discula) discina DI24_1 | 33°05'48"N | 16°18'54"W | MG575176 — Discula (Discula) discina DI24_2 16°18'54"W | MG575177 MG575203 Discula (Discula) polymorpha | \rnizq59 | 3°52'03"N | 17°09'56"W | MGS75178 7 alleniana 12 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) GenBank accession number Taxon Voucher ee Lpnsinide 18S rDNA + ITS1 + cox1 5.8S rDNA + ITS2 + number 28S rDNA Dee ee DI06_1 | 32°44' soraaan'N) 16°41' jiear'serw MGS575179 MGS575204 meee aie wea DI06.2 | 32°44'41"N | 16°41'58"w | MG575180 MGS575205 Discula (Discula) pulvinata DI30_1 | 33°05'07"N MG575181 MG575206 Discula (Discula) pulvinata DI30_2 | 33°05'07"N | 16°21'21"W | MG575182 = Discula (Discula) pulvinata DI30_3 | 33°05'07"N | 16°21'21"W | MG575183 - Discula (Discula) tabellata DI03_1 | 32°38'21"N | 16°51'03"W | MG575184 = Discula (Discula) tabellata DI03_2 | 32°38'21"N | 16°51'03"W | MG575185 — ‘Discula’ testudinalis MN3077 | 33°06'14"N | 16°19'18"W | MG575186 MG575207 aot cane ES MTG Ta MG575187 MG575208 Caseolus (Leptostictea) hartungi CA10_1 MG575188 MG575209 Serratorotula juliformis MN3036 16°17'57"W | MG575210 E Cochlicella (Cochlicella) conoidea SP165_ | 41°08'15"N | 08°40'00"W | KY818425! KY818623' ': Neiber et al. (2017) Laboratory (Amsterdam, The Netherlands). Information on vouchers and GenBank accession numbers are compiled in Table 1. ChromasPro 1.7.1 (Technelysium) software was used to assemble forward and re- verse sequence reads. Nuclear rDNA sequences were aligned with MAFFT 7 (Katoh and Standley 2013) using the Q-INS-i iterative refinement algorithm, whereas cox] sequences were aligned with MUSCLE (Edgar 2004) as implemented in MEGA 6.0 (Tamura et al. 2013). We used maximum likelihood (ML), Bayesian Inference (BI) and maximum parsimony (MP) approaches to reconstruct the phylogenetic relationships. Appropriate evolutionary models for the mitochondrial and nuclear sequence data sets were selected with PartitionFinder 1.1.1 (Lanfear et al. 2012) conducting an exhaustive search with separate estimation of branch lengths for each partition and with the Bayes- ian information criterion (as recommended by Luo et al. 2010) as criterion to select among models. As the nuclear ribosomal DNA sequence were hardly variable among ingroup and outgroup taxa, nuclear sequence data were not divided further into parti- tions. The models to choose from were restricted to those available in MrBayes 3.2.6 (Ronquist et al. 2012) as well as in Garli 2.1 (Zwickl, 2006). As best-fit partitioning scheme, the PartitionFinder analysis suggested to use the HKY + I + G model for the mitochondrial sequences and the GTR + I + G model for the nuclear sequences. BI analysis was performed using MrBayes. Metropolis-coupled Monte Carlo Mark- ov chain (MC’) searches in MrBayes were run with four chains in two separate runs for 50,000,000 generations with default priors, trees sampled every 1,000 generations un- der default heating using the best-fit model as suggested by PartitionFinder. Diagnostic tools in MrBayes, including Effective Sample Size (ESS) values > 200, Potential Scale Reduction Factors ~ 1.000 and an average standard deviation of split frequencies < 0.01, Revision of the genus-group Hystricella R. 7’ Lowe, 1855 from Porto Santo... 13 were used to ensure that the MC? searches had reached stationarity and convergence. The first 5,000,000 generations of each run were discarded as burn-in. ML analyses were performed with Garli and igtree 1.5.3 (Nguyen et al. 2015) us- ing the best-fit models as suggested by PartitionFinder and otherwise default settings. Support values were computed by bootstrapping (BS) with 1,000 replications. MP searches were carried out with PAUP* 4.0b10 (Swofford 2002) using 1,000 random-addition-sequence replicates and TBR branch swapping. Support values were computed by bootstrapping with 1,000 replications. BS values above 70% are interpreted as meaningful node support and PP values above 0.95 as statistically significant support. Collection acronyms ANSP Academy of Natural Sciences of Drexel University, Philadelphia, PA, USA CFW private collection of Frank Walther, Essen, Germany CJG private collection of Jochen Gerber, Chicago, USA CKG private collection of Klaus Groh, Bad Diirkheim, Germany CMN private collection of Marco T. Neiber, Sehnde, Germany CWDM private collection of Willy De Mattia, Trieste, Italy DRAM Direccao Regional do Ambiente da Madeira, Funchal, Madeira, Portugal GNM Goteborgs Naturhistorika Museum, Goteborg, Sweden HNHM ~— Hungarian National History Museum, Budapest, Hungary NHC Natural History Collections, University of Edinburgh, Edinburgh, Great Britain NHM Natural History Museum, London, Great Britain NHMW Natural History Museum Wien, Austria, Malacological collection NMS National Museum of Scotland, Edinburgh, Great Britain MMF Museu de Historia Natural do Funchal (formerly Museo Municipal do Funchal), Funchal, Madeira, Portugal MMUE = Manchester Museum, University of Manchester, Great Britain NMW National Museum of Wales, Cardiff, Great Britain OUMNH Oxford University Museum of Natural History, Oxford, Great Britain RAM Royal Albert Memorial Museum, Exeter, Great Britain SMF Forschungsinstitut und Naturmuseum Senckenberg, Frankfurt/M., Germany ZMH Zoological Museum Hamburg, Hamburg, Germany Morphological abbreviations Genital anatomy A atrium AG albumen gland AF atrial folds BC bursa copulatrix 14 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) DBC | duct of bursa copulatrix PN __ pneumostoma DP distal penis PP penial papilla F flagellum PW _penial wall FAP accessory papilla of the flagellum SL subpneumostomal lobe FO free oviduct RLL right lateral lobe FMP main papilla of the flagellum SP spongy pilaster of the distal penis LDL _ left dorsal lobe SVA _ single vaginal appendage (dartless LSPP longitudinal section of penial papilla stylphore) MR retractor muscle TSPP transversal section of penial papilla OSD _ ovispermiduct V vagina PC penial channel VD __ vas deferens PCE _penial channel exit VG vaginal glands PM _penial matrix VS vaginal section Shell D maximum shell diameter NW © number of whorls H maximum shell height PA peristomal angle FW height of the first (body) whorl DU diameter of umbilicus NT number of tubercles Taxa exclusively known from fossil material are indicated by a dagger symbol (f). Results and discussion Phylogenetic analyses The phylogenetic analyses, both based on the mitochondrial cox] data alone and the concat- enated mitochondrial and nuclear data recovered the included Geomitrini as a mostly well- supported monophyletic group (Fig. 3). Within this clade, several groups, here interpreted as genus-group taxa, were mostly significantly/meaningfully supported (Fig. 3), albeit rela- tionships among these groups did not receive significant/meaningful support in all analyses. Neither the genus Aystricella nor the genus Discula in the sense of Bank et al. (2002), Sed- don (2008), Groh et al. (2009) or Bank (2009) were recovered as monophyletic groups. Species formerly included in Hystricella were assigned to two different clades (Fig. 3). The clade including the type species of Hystricella, H. bicarinata, also included H. echinulata and received significant/meaningful support in all phylogenetic analyses ex- cept in the ML analysis with Garli based on cox/ data alone (Fig. 4). Within the Aystri- cella clade two well-separated clades which largely correspond to specimens morphologi- cally assigned to either . bicarinata or H. echinulata were recovered (Fig. 4). Hystricella bicarinata has its distribution centre in the eastern central to southeastern part of Porto Santo, whereas 1. echinulata is restricted to the northeastern part of the island (Fig. 4). Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... 15 1.00/79/89/100 Hystricella 0.99/57/66/74 Callina 0.79/-/-I79 | 1.00/94/94/ 100 1.00/54/57/71 Wollastonia n. gen cf. Discula 1,00/100/100/100 Discula 0.93/64/69/99 Caseolus 1.00/91/94/100 Cochlicella Hystricella Callina 0.84/-/-/81 1.00/76/85/100 1.00/-/-/57 Wollastonia n. gen cf. Discula 1.00/91/90/100 Caseolus 1,00/60/100/100 one Serratorotula Discula 1.00/72/81/100 Cochlicella B oo Figure 3. Bayesian 50% majority rule consensus tree based (A) on the concatenated sequences of cox] and 18S rDNA + ITS1 + 5.8S rDNA + ITS2 + 28S rDNA and (B) on the cox1 sequences alone. Posterior probability values (first value), bootstrap support values from the maximum likelihood analysis with Garli (second value) and igtree (third value) and maximum parsimony analysis with PAUP* (fourth value) are indicated at the nodes. Only posterior probability values > 0.5 and bootstrap support values > 50% are shown. For phylogenetic relationships of Hystricella, Wollastonia gen. n., Callina, and Discula species included in the analyses, see Figure 4-6. 16 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) 16°|25° W 16°|20'W 16°|15' W | & HY10_1 0.84/61/62/64fF HY10_2 HY45 3 MN3026 HY42_1 , 1.00/87/87/88.] fb riy42 2 HY42_ 3 MN3023 HY45_2 0.58/-/-/- b& MN3020 MN3015 HY36_1 HY36_2 MN3011 aa HY37_2 MN3009 i L. vn3019 HY10_3 HY47_1 HY47_2 HY47_3 MN3003 MN3005 MN3022 H. bicarinata HY35_1 HY36_3 MN3007 Bar? HY33_2 [| H. bicannata 0.94/59/65/62) jiy33 3 1.00/85/9494] L jinisor4 [| H. echinulata 1.00/78/92/95 MIN3049 HY33_1 [| Possible hybrids 0.98/63/78/89} 0.51/-/-/-& HY43_2 HY37_1 33705°N f a 33° 00'N 1.00/90/89/81 Hystricella 0.96/55/54/84 0.73/-/-/- HY03_2 HY03_3 HY04_1 HY32_1 pte HY32_2 0.91/60/60/55 fi TNs} MN3008 | & MN3016 MN3018 MN3029 HY41_1 0.90/83/100/92 & IN3047 0.99/61/79/70 HY39_1 HY39_3 HY40_2 HY40_3 MN3010 0.67/-/-/- HY35_2 HY35_3 H. echinulata MN3006 0.96/-/48/61 ee 1.00/77/84/100] pp HY39_2 MN3004 HY40_1 HY06_1 1.00/80/96/100 0.95/67/85/78 0.99/69/72/68 0.95/63/87/63 0.01 Figure 4. Bayesian 50% majority rule consensus tree based on cox] sequences of Hystricella species. Posterior probability values (first value), bootstrap support values from the maximum likelihood analysis with iqtree (second value) and Garli (third value) and maximum parsimony analysis with PAUP* (fourth value) are indicated at the nodes. Only posterior probability values = 0.5 and bootstrap support values > 50% are shown. The inset map shows the distribution of 1. bicarinata (blue), H. echinulata (green) and putative hybrids of both species (red) on Porto Santo. The tip labels on the tree are coloured accordingly. For information on vouchers, see Table 1 and the corresponding material lists under the species’ sections. ‘The inset tree shows the position of Hystricella in relation to the other investigated genus-group taxa based on the analysis of concatenated mitochondrial and nuclear sequences as shown in Figure 3. Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... 17 1.00/-/70/62 f HYOS_1 HY05_2 HY34_1 HY34_2 ‘ A io r denis W. jessicae jessicae n. ssp. Ss MN3017 MN3021 HY05 3 HY03_1 W. jessicae monticola n. ssp. ES HY01_2 HY01_3 . ~. | W. leacockiana & MN3002 P MN3024 HY38_1 MN3001 } ee W. oxytropis MN3048 1.00/73/90/80 Wollastonia n. gen. 0.66/-/51/61 0.99/62/66/91 1.00/56/67/100 0.67/-/-/64 1.00/92/92/100 1.00/93/95/95 1.00/94/98/98 0.99/-/-/87 0.74/-/50/- 0.77/-/50/- MN3028 HY31_2 HY31_4 W. klausgrohi n. sp. HY31_3 mnzooo )§=~VW. furricula 1.00/94/98/100 0.99/63/67/86 0.01 Figure 5. Bayesian 50% majority rule consensus tree based on cox] sequences of Wollastonia gen. n. species. Posterior probability values (first value), bootstrap support values from the maximum likelihood analysis with Garli (second value) and igtree (third value) and maximum parsimony analysis with PAUP* (fourth value) are indicated at the nodes. Only posterior probability values > 0.5 and bootstrap support values > 50% are shown. For information on vouchers, see Table 1 and the corresponding material lists under the species’ sections. The inset tree shows the position of Wollastonia gen. n. in relation to the other investigated genus-group taxa based on the analysis of concatenated mitochondrial and nuclear sequences as shown in Figure 3. In a small area in the north of Porto Santo populations with intermediate or deviating morphologies (i.e., presence/absence of a double keel on the body whorl and presence/ absence of papilla on the inner wall of the terminal part of the flagellum) were found. In the cox] phylogenies these specimens, either clustered with H. bicarinata or H. echinulata (Fig. 4) and are here interpreted as putative hybrids between the two species. The other clade including species hitherto assigned to Hystricella, although only supported in the BI and MP analyses (Figs 3, 5), will be formally described as Wol- lastonia gen. n. below. This clade includes W. leacockiana comb. n., W. turricula comb. n., and W. oxytropis comb. n., as well as three additional recent taxa that will be for- mally described below: W. klausgrohi sp. n., W. jessicae sp. n., and W. jessicae monticola ssp. n. (Fig. 5). The relationships of most of the taxa within Wollastonia gen. n. could not be well resolved (Fig. 5) on the basis of the phylogenetic analyses of the cox] data. Specifically, although the monophyly of W. oxytropis comb. n. was well-supported in all analyses, the phylogenetic analyses failed to recover the monophyly of W. klaus- grohi sp. n. (Fig. 5). However, the very distinct shell morphology and genital anatomy 18 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) DI35_1 DiI44_1 0.88/55/59/- 1.00/81/84/84 DI35_2 0.98/70/73/63 MINSOS' DI23.1 0.99/-79/83/78] b DI37_2 Di44_2 C. rotula 0.57/-/-I- DI25_1 DI29_2 .73/-167/- _ DI37_1 RAS DI26_1 Die 1.00/91/100/99 z pM3.2 DI31_1 0.55/55/63/52 DI21_2 DI31_2 1.00/76/85/100 ~ DI29.1 Dl04_1 0.69/66/70/75 0.95/59/63/63) f DI29_4 DIN5 3 —— es . DI29_5 0.01 bi271 | C. bulverii Crise 0.63/74/76/92 DI15_2 7 D. attrita ae 0.95/50/69/65 MN3053 A = DI05_1 1.00/82/83/99 0.74/67 | nnmmmmf P!95- DI05_3 ; DI34_1 Callina 1.00/74/80/100 i Di121 DI38_1 DI38_2 Pirie 1.00/80/83/85 os7iexieae2f 242-1 D. calcigena DI30_1 1.00/100/100/100 | | zi ; 0.99/62/65/63 DI30_2 D. pulvinata 30.3 0.97/70/67/53 0.71/60/65/55 MN3052 1.00/96/97/100 DI06_2 D polymorpha Di06_1 Di16_1 DI221 0.80/85/96/91 Di22_2 DI22 3 DI22_ 4 0.62/78/82/88 1.00/72/81/100 DI08_3 DI16_3 Diaiee Di244 . aiscina 1.00/83/90/100 Dlos_1 Di24.2 1.00/73/92/87 Dlos_2 0.73/-I51/- | DI08_4 Di16 2 0.01 DI16_4 DI03_1 B 1.00/78/100/100 L p93 2 D. tabellata Figure 6. Bayesian 50% majority rule consensus tree based on cox] sequences of (A) Callina and (B) Discula species. Posterior probability values (first value), bootstrap support values from the maximum like- lihood analysis with Garli (second value) and igtree (third value) and maximum parsimony analysis with PAUP* (fourth value) are indicated at the nodes. Only posterior probability values > 0.5 and bootstrap support values > 50% are shown. For information on vouchers, see Table 1 and the corresponding mate- rial lists under the species’ sections. The inset tree shows the position of Callina and Discula in relation to the other investigated genus-group taxa based on the analysis of concatenated mitochondrial and nuclear sequences as shown in Figure 3. Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... 19 of W. oxytropis comb. n. (see below) clearly supports the recognition of two distinct, though closely related species. Interspersed between the Hystricella and Wollastonia gen. n. clades, a clade con- taining Callina rotula (R. T. Lowe, 1831), the type species of Callina R. T. Lowe, 1855, was placed as the sister group of Hystricella with significant/meaningful support in the BI and MP analyses (Figs 3, 6A). Callina was up till now either regarded as a monospecific genus related to the genus Discula (Waldén 1983; Schileyko 2005), or included as a subgenus in Discula, but the phylogenetic analyses presented here do not support the monophyly of Callina plus Discula s. str. (i.e., the clade including the type species of the genus D. discina (R. T. Lowe, 1852), see Figs 3, 6A). Interestingly, the phylogenetic analyses recovered a close relationship of C. rotula and C. bulverii (Wood, 1828), comb. n., a taxon up till now included in Discula s. str. (Bank et al. 2002; Seddon 2008; Bank 2009). In the phylogenetic analyses, both based on the concatenated and the cox] data set, a specimen of Discula testudinalis (R. T. Lowe, 1852) from Porto Santo was placed in a sister group relationship to the clade including Hystricella, Wollastonia gen. n. and Callina, albeit not supported in all analyses (Fig. 3). The remaining Discula species were grouped together with statistical support in the BI and MP analyses based on the concatenated mitochondrial and nuclear data and in all analyses based on the cox/ data alone (Fig. 6B). Within the Discula clade three well-supported clades were placed in a polytomy (Fig. 6B): the first of these clades grouped two specimens of D. tabellata (R. T. Lowe, 1852) from Madeira together, the second group included specimens of D. discina (R. T. Lowe, 1852) from Porto Santo and the third clade included specimens of D. attrita (R. T. Lowe, 1831), D. pulvinata (R. T. Lowe, 1831) and D. calcigena (R. T. Lowe, 1831) from Porto Santo as well as D. polymorpha (R. T. Lowe, 1831) from Ma- deira. Within the latter clade, D. attrita was recovered as a mostly well-supported sister group of the remaining three taxa, and D. polymorpha as the sister group of a clade containing D. calcigena and D. pulvinata (Fig. 6B). Discula pulvinata was recovered as a paraphyletic group with respect to D. pulvinata, however, divergences within this clade were rather low overall and the branching patterns not well-resolved. Systematic account The suprageneric classification follows Razkin et al. (2015) and Neiber et al. (2017), elevating the Geomitridae from subfamily to family rank with Geomitridae plus Ca- nariellidae Schileyko, 1991 as the sister group of Hygromiidae Tryon, 1866. Helicoidea Rafinesque, 1815 Geomitridae C. R. Boettger, 1909 Geomitrinae C. R. Boettger, 1909 Syn. Ochthephilinae Zilch, 1960 in Wenz and Zilch (1959-1960) Geomitrini C. R. Boettger, 1909 20 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) Hystricella R. T. Lowe, 1855 Discula auct., partim [non R. T. Lowe, 1852]. Helix (Ochthephila) sensu Albers (1850) and Albers (1854), partim. Helix (Octephila), Paiva (1867) [incorrect subsequent spelling Ochthephila Albers, 1850], partim. Geomitra (Actinella (Hystricella)), Pilsbry (1893-1895). Geomitra sensu Bank et al. (2002). Type species, by original designation in R. T. Lowe (1855). Helix bicarinata G. B. Sowerby I, 1824 = Aystricella bicarinata (G. B. Sowerby I, 1824). Lowe (1855: 186-187) describes the new genus Hystricella in the section § 24, basing his description upon shell features only, as follows: (Typ. H. bicarinata, Sow.).'T. perf. v. angustae umbil. conuloidea v. trochiformis, aliquando turrito-pyramidata acute v. distincte 1—2-carinata solidiuscula aspero-granu- lata v. echinulata subfasciata. Anfr. 6—9 lente crescentes planiusculi, ult. valde carinato subtus planato, antice deflexo. Umbil. parvus anguste cylindricus v. subspiralis con- strictus. Apert. circularis circinata labris connexis; perist. continuum solutum expanso- reflexiusculum tenue acutum. Lowe (1855: 186-187) cites for the genus (under section § 24) the following species: H. bicarinata G. B. Sowerby I, 1824 H. echinulata R. T. Lowe, 1831 FH. oxytropis R.'T. Lowe, 1831 H. turricula R. 'T. Lowe, 1831 H. vermetiformis R.'T. Lowe, 1855 Hystricella mustelina Reeve, 1854 [under authorship R. T. Lowe, 1855], currently considered as a subspecies of Discula (Discula) cheiranthicola Reeve, 1854 [even though the description was prepared by R. T. Lowe in 1854 the work was only published in 1855 and therefore the earlier publication of that name in Reeve in 1854 has priority]. Hystricella cheiranticola R. 'T. Lowe, 1831 and its morphae (later put in the mono- specific subgenus [as section] Turritella by Wollaston 1878 [non Lamarck 1799]). Later, H. mustelina (currently Discula (Discula) cheiranthicola mustelina (Reeve, 1854)) and H. cheiranticola (currently Discula (Discula) cheiranthicola cheiranthicola (R. 'T. Lowe, 1831)) were assigned to the genus Discula R. T. Lowe, 1852 (Mandahl-Barth 1950). Description of the genus. Shell. ‘The shell is dextral, hairless and it is usually conical and scalariform. The protoconch is whitish to dark brown, with 1.3 to 2.2 whorls. It is almost smooth along the first whorl and shows fine radial striae and extremely small, scat- tered tubercles along its remaining portion. The teleoconch has from 4.2 to 5.0 rapidly increasing whorls. It is usually dark brown with brick red and/or dark violet colour shades. The dark areas of the shell are mottled with more or less light brown to whitish areas, usu- ally placed longitudinally and slightly slanting. No band pattern is visible along the upper whorls. On the lower part of the last whorl two well-defined, dark bands are visible that can differ in width. The area around the umbilicus is usually the lightest in colour. The Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... 21 spire is slightly variable in height, ranging in shape from compressed to somewhat more elevated. Along the last and the penultimate whorls one to two evident keels are present. The external upper surface has very fine but clearly visible, irregularly spaced growth lines. Along the last whorls the growth lines usually disappear along the lateral area and reappear on the lower part. Irregularly disposed tubercles are found all over the teleoconch. The dimensions of the tubercles tend to increase slightly from the first toward the last whorl, the density remaining, however, approximately the same. The tubercles are somewhat denser along the keels of the penultimate and last whorls, letting the keel/keels appear like a rough chord. On the lower part of the last whorl the tubercles are usually bigger and slightly less dense than in the remaining parts of the shell. The last whorl is large, with a contribution of 60% of the total shell height and descending towards the aperture. The umbilicus is open but very narrow, concentric and has a diameter of approximately 10% of the maximum shell diameter. The aperture is elliptical with a faint thickening along the inner side of the last whorl. The peristome is continuous, slightly reflected with the columellar margin somewhat thicker and more reflected. Body. Head and neck are grey to pale grey, slightly transparent. The sides and pos- terior upper section of the foot are whitish. The sole of the foot is whitish and longitu- dinally divided into three areas. ‘The central area is smooth whereas the two lateral areas are equipped with bands of muscles roughly arranged in a chevron pattern. The mantle border is grey to dark grey with five more or less developed lobes. The ratio among lateral and dorsal lobes varies from specimen to specimen also in the same population. In some specimens, particular lobes (regardless if lateral or dorsal) may be completely missing. The walls of the pallial cavity are colourless, without any stripes or spots. A strong pulmonary vein is visible. Genitalia. The general arrangement of the genitalia is semi-diaulic monotrematic. A convoluted to almost straight first hermaphroditic duct arises from a multi-lobated gonad. The albumen gland is long and moderately thin and connected to an equally long sperm-oviduct consisting of a prostatic and a uterine portion. Distally, the pros- tatic part extends into a thin vas deferens, twice as long as the sperm-oviduct, termi- nating in the penial complex. The distal portion of the uterine part inserts into the free oviduct, then transforming into a vagina at the level of the insertion of the duct of the bursa copulatrix. The free oviduct is usually three to four times shorter than the vagina. The duct of the bursa copulatrix is usually wide, approximately as long as the penis and uniform in diameter. This duct terminates in a variable, oval to roundish, small bursa copulatrix. The spermatophore is unknown. One tuft of digitiform glands arises from the proximal part of the vagina. There are usually two or three, equally long and very rarely branched glands. A short and thin vaginal appendix arises from the vagina’s wall, immediately distal of the glandular tuft. Very smooth, little elevated and spaced pleats run longitudinally along the inner surface of the vagina, reaching into the genital atrium as far as the genital orifice. The atrium can be short or moderately long. The penial complex consists of a flagellum, an epiphallus (which extends from insertion of the vas deferens to the penial retractor muscle) and a penis that inserts into the genital atrium. The penial flagellum is short, remarkably cylindrical and with 22 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) a blunt apex. It is usually twice as long as the epiphallus. Its internal walls can be either completely smooth or ornamented with very small papillae distributed mainly close to the blunt end. The epiphallus is usually extremely short and its internal walls are smooth. The retractor muscle is well developed, strong and is variable in length. The penis lacks a muscular or glandular sheath. It is thick-walled and approximately four times longer than the flagellum. It is usually cylindrical to sometimes slightly swollen in its distal part. The inner walls of the penis are usually smooth or with very smooth, little elevated and spaced pleats which run longitudinally and reach the genital atrium. The section where the penial papilla is located is usually detectable from the outside by virtue of a fine circular swelling corresponding to the origin of the papilla itself. The penial papilla is very variable in size, ranging from 1/6 to 1/2 of the total penial length and is conical to subcylindrical in shape. The inner lumen of the penial papilla is filled with a spongy and sturdy tissue which directly connects with the walls of the epiphal- lus. It has smooth external walls with the opening emerging apically. The channel of the penial papilla is thin and narrow. The longitudinal section of the penial papilla shows that its walls are the continuation of the penial walls that abruptly bend inward. Jaw and radula. The jaw and radular apparatus of Hystricella is depicted in Fig. 8. No notable variability was found among the species of the genus. The jaw is odonto- enathous, almost straight to markedly arched, with 13 to 15 smooth ridges. The radula ribbon is typical helicoid, it is elongated but not very slender. Central tooth present, tricuspid, the main cusp (endocone) is rhomboid, pointed; the ectocones are much smaller than the endocone; they are triangular, pointed. There are 19-20 laterals and marginals which do not distinctly differ from each other, i.e., their shape changes gradually from the first laterals towards the marginals. Laterals are bicuspid, endocones are rhomboid or triangular and pointed. The ectocones are much smaller, pointed, and triangular. The endocones of the central tooth and the first laterals are approximately of the same size. Both, the endocone and the ectocone of the laterals gradually become bifurcated towards the marginal teeth, but the ectocones may occasionally have three cusps as well. The cusps of the marginals are gradually decreasing in size; therefore, the outermost marginals appear nearly serrated. Distribution. The genus Hystricella is endemic to the island of Porto Santo (Ma- deiran Archipelago, Portugal). The genus is restricted to the eastern, mountainous part of the island and occurs there only in the central-northern section of this area (Fig. 7). Species assigned to the genus are commonly found on the slopes of Pico do Castelo, Pico do Facho, Pico de Juliana, Pico da Gandaia, Rocha de Nossa Senhora, the western slopes of Pico do Concelho, and those of the Pico Branco and Terra Cha complex. It is also commonly found at lower elevations of Serra de Dentro, Barranco Branco, and Lombo de Paredes. Recent Hystricella are absent from the southeastern part of the island, i.e., from Vale do Touro, Portela, Pico do Macarico, and Pico do Baixo. It is likewise not present on the small islets surrounding the main island, namely Ilhéu de Cima, Ilhéu de Baixo, and Ilhéu de Ferro. Subfossil representatives of the genus are found mainly in the mud and aeolinite deposits along the southeastern coast, namely Vale do Touro, Ponta do Passo, Barbinha, and Calhau da Serra de Fora. Revision of the genus-group Hystricella R. 77 Lowe, 1855 from Porto Santo... 23 Figure 7. Distribution of the genus Aystricella. Filled circles refer to recent and open circles to fossil records. Tce MEV Vy Leet b Mhiht. 1 2016/08/24 LD39 x18k 50um 2016/08/24 ppp ANY “7a y , 2016/08/24 LD3.9 x1.8k 50um Figure 8. Radulae and jaws of Hystricella species. A Hystricella bicarinata, Ribeira da Areia B Aystricella echinulata, path to Pico Branco. Ecology. Representatives of the genus Hystricella are commonly found under vol- canic rocks scattered on grassland in open fields that are more or less strongly sloping. They have been found under stones in pine woods (Pico do Castelo) or in cracks and 24 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) crevices of rocky walls (Pico da Gandaia). Disturbed or anthropogenic habitats have also been colonised by representatives of the genus, such as stone walls (Pedregal de Dentro) or terraced areas (southern slopes of Pico do Castelo). Specimens aestivate on the lower surfaces of stones or rocks, frequently forming large clusters of individuals attached to one another, reaching more than 40 to 50 in number. Under a single stone of roughly 60 x 40 cm approximately 200 individuals were counted (southern slopes of Pico do Facho). Nomenclatural and taxonomic remarks. Hystricella was considered a subgenus of Helix for a long time (until Wollaston 1878) or was replaced by Ochthephila Beck, 1837 (Albers 1850) or Octephila Paiva, 1867 (Paiva 1867). Later it was considered as a subgenus of Discula R. T. Lowe, 1852 and only since 2002 (Bank et al. 2002) accepted as a distinct genus. Bank et al. (2002) used the generic name Geomitra Swainson, 1840 despite Herr- mannsen’s (1847: 470) selection of the nominal species Helix tiarella Webb & Berthe- lot, 1833 as the type species of that genus. The decision of Herrmannsen was followed by all subsequent authors (e.g., Wollaston 1878; MandahI-Barth 1950; Waldén 1983). ‘The reason to contradict that type selection was the fact that it was wrong with respect to the rules of the International Commission on Zoological Nomenclature’s (ICZN) Code of Zoological Nomenclature (hereafter ‘the Code’) Art. 67.2.1: Swainson intro- duced Geomitra describing and depicting tiarella, namely on p. 166 and 332, but he did not mention that name, but instead only bicarinata, which therefore would be the type species by monotypy. In consequence Hystricella R. T. Lowe, 1855 would become a synonym of Geomitra and Geomitra sensu auct. has to be referred to Craspedaria R. T. Lowe, 1852. Nevertheless, this decision was much criticised (Seddon 2008) and a new study of the Code revealed a solution which permits the stabilisation of the former use of Geomitra (Groh et al. 2009). Following Art. 70.3 of the Code the combination of the figure of tiarella with the name bicarinata can be interpreted as a misidentification by Swainson and therefore following Art. 70.3.2, the name tiarella becomes available for type selection under the synonymous name Geomitra bicarinata Swainson, 1840. This was already recognised by Pfeiffer (1847: 191) who listed Swainson’s combination as a synonym of Helix tiarella (cf. Groh et al. 2009). Comparison and comments. [ystricella shows a number of morphological char- acters that clearly distinguishes it from the other native geomitrid genera from Porto Santo, in particular with regard to the genus Discula s. lat., into which the Hystricella species were previously placed (Mandahl-Barth 1950; Waldén 1983). On the basis of the shell, the main distinguishing feature of Hystricella from Discula s. lat. is the continuous and detached peristome. On the contrary, in Discula s. lat. and Callina (Figs 10—12) the peristome is always interrupted, forming only a thickening or a cal- lous along the parietal region but not a real and proper detached lip. Other minor differences can be found in the ornamentation of the shell’s surface. Hystricella always possesses round, somewhat spaced tubercles, whereas Discula s. lat. and Callina shows mainly elongated, sometimes drop-like tubercles arranged in a regular pattern. In Ays- tricella the size of the tubercles can reach large dimensions in comparison to the overall Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... 23 Figures 9-19. Shells of Portosanctan and Madeiran Geomitridae. 9 Hystricella bicarinata, Pico do Castelo, S slope around the Miradouro 10 Discula discina, Porto da Morena I Discula (Mandablia) tectiformis, Pico do Baixo 12 Callina rotula, Cabeco dos Bades 13 Actinella (Plebecula) littorinella, E of Pico do Castelo 14 Caseolus (C.) innominatus portosanctanus, Ribeira da Areia 15 Caseolus (Helicomela) punctulatus punctulatus, Fonte da Areia 16 Caseolus (Leptostictea) leptosticus, Ponta do Garajau, Madeira 17 Serratorotula juliformis, Pico de Ana Ferreira 18 Spirorbula depauperata, Pico de Ana Ferreira 19 Lem- niscia michaudi, Terra Cha. Scale bar: 1 cm. size of the shell and their size can remarkably vary on the same shell. Actinelia s. lat. and Caseolus s. lat. (see Figs 13-16) also possess an interrupted peristome. Most Caseolus species have a more or less granulated shell’s surface and, at first glance, could therefore be confused with Hystricella. However, a closer look reveals the different peristome and the overall different shape, with the whorls always rounded and usually without any prominent keel. Serratorotula (Fig. 17) has an extremely ornamented shell that is easily 26 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) Figures 20-30. Outer distal genitalia of Portosanctan and Madeiran Geomitridae. 20 Aystricella bicari- nata, Capela da Graga 21 Discula polymorpha alleniana, Porto Moniz, Madeira 22 Discula (Mandahlia) tectiformis, Pico do Baixo 23 Callina rotula, Cabeco dos Bades 24 Serratorotula juliformis, Pico de Ana Ferreira 25 Lemniscia michaudi, Terra Cha 26 Actinella (Plebecula) littorinella E of Pico do Castelo 27 Ca- seolus (C.) innominatus portosanctanus, Ribeira da Areia 28 Caseolus (Helicomela) punctulatus punctulatus, Fonte da Areia 29 Caseolus (Leptostictea) leptosticus, Ponta do Garajau, Madeira 30 Spirorbula depauperata, Pico de Ana Ferreira. Scale bars 1 mm. distinguishable from Hystricella. The shell form of Heterostoma is very different from that of Hystricella and these two genera are therefore easily distinguishable (see Seddon 2008: 125). Some species of the genus Spirorbula sometimes have shells with a con- Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... LF tinuous peristome with detached lips along the parietal side. The very depressed shell shape and different surface ornamentation of Spirorbula however, readily distinguishes the two genera (Fig. 18). Lemmniscia usually possesses a smooth and rather glossy shell without tubercles or papillae (Fig. 19). The shell morphology of Wollastonia gen. n. and its differences with regard to Hystricella will be discussed in the section on Wol- lastonia gen. n. below. With regard to genital morphology, Hystricella shows a unique feature that allows the separation of the genus from all native Portosanctan Geomitridae, except from Wollastonia gen. n. The main difference is the shape of the penial flagellum which is al- ways short and has a remarkably blunt apex (Fig. 20). Discula s. lat. and Callina, Serra- torotula, and Lemniscia (Figs 21—25) have a pointed, more or less elongated flagellum. Hystricella is easily distinguishable from Caseolus s. lat. and Actinella s. lat. by virtue of the single vaginal appendages instead of the two that are present in these genera (see Figs 26-29). Spirorbula is also easily distinguishable from Hystricella because members of this genus possess two extremely long, and sometimes branched, vaginal appendages and a genital atrium with structures on the inner wall consisting of large and partially fringed folds (see Fig. 30). Hystricella bicarinata (G. B. Sowerby I, 1824) Figs 31-54 List of synonyms. 1824 Helix bicarinata G. B. Sowerby 1: 58, pl. 3 fig. 7. 1828 Helix bicarinata — Wood: pl. 8 fig. 85. 1831 Helix duplicata BR. T. Lowe: 58, pl. 6 fig. 30 [unjustified replacement name for Helix bicarinata Sowerby I, 1824 [non Helix C.[ochlitoma] bicarinata J. Férus- sac, 1821, a secondary homonym of Bulimus bicarinatus Bruguiére, 1792 = Archachatina bicarinata (Bruguiére, 1792). See Taxonomic Remarks below]. 1846 Helix bicarinata — L. Pfeiffer: 141, pl. 91 figs 8-11. 1847 Helix bicarinata — L. Pfeiffer in L. Pfeiffer 1847-1848: 190. 1854 Helix bicarinata — Reeve in Reeve 1851-1854: pl. 142 figs 908-909. 1854 Helix (Ochthephila) bicarinata — Albers: pl. 9 figs 1-4. 1855 Helix (Aystricella) bicarinata — R.'T. Lowe: 186. 1867 Helix (Octephila) bicarinata — Paiva: 45. 1878 Helix (Hystricella) bicarinata — Wollaston: 161-163. 1888 Helix bicarinata —Tryon in Tryon and [Pilsbry] 1888: 33, pl. 7 fig. 90. 1894 Geomitra bicarinata — Pilsbry in Pilsbry 1893-1895: 242. 1931 Geomitra (Actinella) bicarnata — Nobre: fig. 37. 1950 Discula (Hystricella) bicarinata bicarinata — Mandahl-Barth 1950: 31, 55. 1966 Discula (Hystricella) bicarinata —S. G. A. Jaeckel: 53. 1983 Discula (Hystricella) bicarinata bicarinata — Waldén: 267. 2002 Geomitra bicarinata bicarinata — Bank et al.: 124. 28 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) 2008 Adystricella bicarinata — Seddon: 79, pl. 29 fig. E, map 178. 2011 Adystricella bicarinata — Seddon: e.16724A12800659. Type material. Despite intensive research in multiple museum collections (NHM, NMW, MMUE, ANSP, NHC, NMS, OUMNH, RAM, SMF) that could have held the type material of the taxon, no such material could be traced and therefore we deem it reasonable to assume that the type material is lost. To stabilise the present interpreta- tion of Helix bicarinata G. B. Sowerby I, 1824 and to clarify its taxonomic status we designate a neotype here, which is deposited in the collection of the SMF under the No. SMF 348936 (see Fig. 33). The original figures of Helix bicarinata G. B. Sowerby I, 1824 from Sowerby (1824: pl. 3 fig. 7) 32 and Helix duplicata R. T. Lowe, 1831 from R. T. Lowe (1831: pl. 6 fig. 20) are depicted in Figs 31-32. The neotype (Fig. 33) is consistent with the figure in Sowerby (1824: pl. 3 fig. 7) and the original description (see below), especially with regard to size, shape and the presence of two keels on the body whorl. The taxon was originally described from Porto Santo without more de- tailed locality data, which is consistent with origin of the specimen selected as neotype. Locus typicus. Pico do Castelo, S slope around the Miradouro, under stones in pine wood, 33°04'42"N/16°20'00"W, 270 m (through the designation of the neotype). Further material examined. All from Porto Santo, ANSP H 20185/13, NW slope of Pico Branco, 33°05'50"N/16°18'57"W to 33°05'52"N/16°18'49"W, 150- 200 m, leg. J. & C. Hemmen, Jan. 8 1981; ANSP H 11776/2, NW slope of Pico de Juliana, 33°05'34"N/16°19'25"W, up to 450 m, leg. J. & C. Hemmen, Jan. 6 1981; CKG/12, Pico de Juliana, under stones, 33°05'31"N/16°19'20"W, 365 m, leg. K. Groh, Jun. 23 1983; ANSP H 11788/11, CMN/3, SW slope of Pico do Juliana, 33°05'30"N/16°19'23"W, 360 m, leg. J. Gerber, K. Groh & J. Hemmen, Aug. 13 1985; CFW 11149/1, Pico Juliana, old quarry on the southern slope, 33°05'28"N/16°19'17"W, 330 m, leg. FE Walther & E. M. Gryl, Apr. 3 2017; ANSP H 11769/3, Ribeiro das Esmoitadas (E Camacha), 33°05'27"N/16°20'17"W, 130 m, leg. J. Gerber, K. Groh & J. Hemmen, Aug. 13 1985; CKG/2, SW of Pico de Juliana, 33°05'14"N/16°19'32"W, 360 m, leg. K. & C. Groh, Oct. 27 1980; CFW 11158/10, Pico do Facho, NW slope, 33°05'12"N, 16°19'38"W, 370 m, leg. RF Walther, Apr. 3 2017; ANSP H 20186/1, NE slope of Pico do Facho, 33°05'07"N/16°19'23"W, 430— 450 m, leg. K. Groh & J. Hemmen, Jun. 28 1983; CWDM/6, CMN/ 3, ridge between Pico do Facho and Pico da Gandaia, stone walls, 33°05'04"N/16°19'21"W, 460 m, leg. W. De Mattia & J. Macor, May 25 2015; CWDM/15, N slope of Pico do Facho in Tamarix wood, under stones, 33°05'04"N/16°19'25"W, 475 m, leg. W. De Mat- tia & J. Macor, May 25 2015; CKG/44, CMN/6, W of Pico do Facho, under stones, 33°05'04"N/16°19'33"W, 450 m, leg. K. Groh, Jun. 29 1983; ANSP H 11778/c. 40, SW slope of Pico do Facho, 33°05'03"N/16°19'31" W, 460-480 m, leg. K. Groh & J. Hemmen, Jun. 28 1983; CKG/24, CMN/44, upper Ribeira Formosa between Covau and Serra de Fora, under stones, 33°05'02"N/16°19'14"W, 140 m, leg. K. Groh, Jun. 19 1983; ANSP H 11775/13, W slope of Pico do Facho, 33°05'02"N/16°19'38"W, leg. J. & C. Hemmen, Jan. 6 1981; ANSP H 11834/19 [sub H. echinulata], summit of Pico da Gandaia, 33°05'01"N/16°19'14"W, 440 m, leg. J. & C. Hemmen, Jan. 6 Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... 29 Figures 31-32. 31 Original figure of Helix bicarinata G. B. Sowerby I, 1824 from Sowerby (1824: pl. 3 fig. 7) 32 Original figure of Helix duplicata R. T. Lowe, 1831 from R. T. Lowe (1831: pl. 6 fig. 20). 1981; CWDM/11, CMN/7, Pico da Gandaia, southern rocky cliffs close to the top, under stones, 33°05'01"N/16°19'14"W, 440 m, leg. W. De Mattia & J. Macor, May 25 2015; ANSP H 11779/24, SW slope of Pico do Facho, 33°04'56"N/16°19'39"W to 33°05'02"N/16°19'30"W, 300-450 m, leg. K. Groh & J. Hemmen, Jul. 1 1983; CWDM/15, CMN/3, N and E slopes and summit of Pico do Castelo, under stones in pine wood, 33°04'55"N/16°20'07"W, 350 m, leg. W. De Mattia & J. Macor, May 22 2015; CWDM/25, CMN/5, S slope of Pico do Facho, along path at the S border of the pine wood, under stones, 33°04'52"N/16°19'26"W, 355 m, leg. W. De Mattia & J. Macor, May 17 2015; CKG/8, Pico do Castelo, S slope around the Miradouro, under stones in pine wood, 33°04'5 1" N/16°20'03"W, 420 m, leg. K. & C. Groh, Oct. 27 1980; CWDM/22, CMN/4, Ribeira da Areia, serpentine 240 m NNW the quarry, under stones, 33°04'51"N/16°18'41"W, 140 m, leg. W. De Mattia & J. Macor, May 21 2014; ANSP H 11787/c. 30, approx. 1 km NE Capela da Graga, 33°04'49"N/16°19'04"W, 250 m, leg. J. & C. Hemmen, Jan. 4 1981; CWDM/12, CMN/4, lake shore S of Ribeira da Serra de Dentro, under stones and in stone walls, 33°04'48" N/16°18'31" W, 70 m, leg. W. De Mattia & J. Macor, May 22 2015; ANSP H 20187/1, NW slope of Pico do Concelho, 33°04'44"N/16°18'04"W to 33°04'43"N/16°18'01"W, 250-300 m, leg. K. Groh & J. Hemmen, Jul. 29 1983; CKG/8, CMN/8, Pico do Concelho, NW of the top, under stones, 33°04'43"N/16°17'59"W, 260 m, leg. K. Groh, Jun. 29 1983; ANSP H 11835/12 [sub H. echinulata], NW slope of Pico do Concelho, 33°04'43"N/16°18'01" W, approx. 280 m, leg. J. & C. Hemmen, Jan. 6 1981; ANSP H 12780/32, NW slope of Pico do Concelho, 33°04'43"N/16°18'01"W, 250-300 m, leg. K. Groh & J. Hemmen, Jun. 6 1983; CWDM/15, CMN/2, summit of Pico do Concelho, under stones, 33°04'42"N/16°18'01"W, 280 m, leg. W. De Mattia & J. Macor, May 18 2015; CWDM/18,CMN/4, Pico do Castelo, S slope around the Miradouro, under stones in pine wood, 33°04'42"N/16°20'00"W, 270 m, leg. 30 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) Figures 33-37. Shells of Hystricella bicarinata. 33 neotype of Hystricella bicarinata, Pico do Caste- lo, S slope around the Miradouro, SMF 348936 34 syntype of H. duplicata R. T. Lowe 1831, NHM 1948.7.8.34; Shells of Hystricella bicarinata 35 Capela da Graca 36 Pico do Castelo, S slope around the Miradouro 37 Pico da Gandaia. Scale bar 1 mm. W. De Mattia & J. Macor, Feb. 12 2012; CMN/1, Pico do Castelo, S slope around the Miradouro, under stones in pine wood, N slope, 33°04'42"N/16°20'01"W to 33°04'51"N/16°20'03"W, 300-430 m, leg. K. Groh Jul. 1 1983; ANSP H 11783/24, Pico do Castelo, 33°04'42"N/16°20'01" W to 33°04'51"N/16°20'03"W, 350-430 m, leg. K. Groh & J. Hemmen, Jul. 1 1983; CWDM/8, CMN/4, path from Capela da Graga to Pico do Castelo along the Levada, under stones, 33°04'38"N/16°19'48"W, 165 m, leg. W. De Mattia & J. Macor, May 17 2015; CKG/3, CMN/68, Capela da Graca up to Pico de Facho-Motos de Fora, under stones, 33°04'38"N/16°19'33"W, 200 m, leg. K. Groh, Jun. 23 1983; ANSP H 11785/9 spms, Ribeiro do Formoso, 33°04'36"N/16°19'04"W, 130 m, leg. J. Gerber, K. Groh & J. Hemmen, Aug. 13 1985; ANSP H 11777/12, approx. 1 km S Serra de Dentro, 33°04'33"N/16°18'25"W, 160 m, leg. J. & C. Hemmen, Jan. 8 1981; ANSP H 11782/c. 30, slope between the SE slope of Pico do Castelo and Capela da Graga, 33°04'32"N/16°19'38"W, 160 m, Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... oa leg. J. & C. Hemmen, Jan. 4 1981; CFW 11161/7, Casinhas, N of Capela da Graga, 33°04'27"N/16°19'27"W, 150 m, leg. E Walther, Apr. 4 2017; CKG/11, CMN/1, Pico do Baixo, 33°04'26"N/16°19'44"W to 33°04'46"N/16°19'44"W, 100-250 m, leg. K. Groh, Jul. 11, 1983; CWDM/23, Engula, under stones, 33°04'19"N/16°19'01" W, 150 m, leg. W. De Mattia & J. Macor, May 18 2015; CFW 11159/>10, E slope of Rocha de Nossa Senhora, 33°04'19"N/16°19'01"W, 150 m, leg. E Walther, Apr. 3 2017 ANSP H 11771/18, N of Portela (E of Vila Baleira), 33°04'07"N/16°19'17"W, 120 m, leg. J. & C. Hemmen, Jan. 4 1981; CKG/2, CMN/1, Capela da Graca up to Pico do Fa- cho-Motos de Fora, N slope, 33°04'38"N/16°19'33"W to 33°05'04"N/16°19'33"W, approx. 200-450 m, leg. J. Gerber, K. Groh & J. Hemmen, Jun. 28 1985; ZMH 120610/2, foot of Pico do Castelo, c. 33°04'37"N/16°19'49"W, 170 m, ex coll. W. Fauer, leg. J. & C. Hemmen, Jul. 11 1983; ZMH 110165/68, slopes of Pico do Facho, under stones; c. 33°04'59"N/16°19'25"W, 430 m a.s.L., leg. E. Clauss, Sep. 22 1992; ZMH 24288/2, Madeira Archipelago, without exact locality data, ex coll. Altonaer Museum; ZMH, 24289/1, Porto Santo, without exact locality data, ex coll. Museum Klagenfurt; ZMH 24290/3, Porto Santo, without exact locality data, ex coll. Altonaer Museum, ex coll. O. Semper, ex coll. Dohrn. Original description. From Sowerby (1824): testa subglobosa, spira breviuscula, subconica; anfractibus quinque quadratis, mediane carinis duabus, superiore obtu- siuscula: apertura integra, rotunda, peristomate distincto: umbilico parvo. Axis 4, diam. 14 unc. Diagnostic features of the shell. Shell as in the genus description. The main di- agnostic feature is the presence of two well-developed keels along the penultimate and body whorl. The lower keel is usually slightly stronger and more evident than the upper one along both whorls. The overall shape of the shell of Hystricella bicarinata is always conical and scalariform by virtue of a “shoulder” that lets the contour of the whorls appear markedly angular. The sutures are deep and well-marked (see Figs 33-37). Measurements. D 4.9 + 0.2 mm (range 4.4-5.3 mm); H 3.8 + 0.2 mm (range 3.4-4.2 mm); FW 2.4 + 0.2 mm; PA 53.5 + 2.4°; DU 0.4 + 0.09 mm; NT 21 + 6; NW 5.6 + 0.1 (range 5.1-5.75) (7 = 40). Ratio D/H 1.3; ratio FW/H 0.6. Body. Body as in the genus description. Genital anatomy. As in the genus description. The albumen gland is long and connected to an equally long sperm-oviduct. The prostatic part extends into a thin vas deferens that is twice as long as the sperm-oviduct. The free oviduct is somewhat vari- able in length, but is usually three to four times shorter than the vagina. The duct of the bursa copulatrix is usually wide, slightly shorter than the penis, opening out into a roundish, small bursa copulatrix. One tuft of digitiform glands consisting of two, more rarely three never branching glands. ‘The inner ornamentation of the vagina consists of very wide, low, and smooth pleats running longitudinally as far as the genital atrium. The penial flagellum is short, remarkably cylindrical and with a blunt apex. It is usu- ally as long as the epiphallus. Its internal walls are completely smooth. ‘The epiphallus is short and its internal walls are smooth. ‘The penis lacks any muscular or glandular sheath. It is thick-walled and approximately four times longer than the flagellum. It 32 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) is cylindrical and sometimes slightly swollen in its distal part. The inner walls of the penis are usually smooth or with very smooth, little elevated and spaced pleats which run longitudinally and reach the genital atrium (see Figs 38-53). Distribution. Hystricella bicarinata is endemic to the island of Porto Santo (Ma- deiran Archipelago, Portugal) (Fig. 54). The species is restricted to the eastern, moun- tainous part of the island and is present only in the central section of this area. It is commonly found on the slopes of Pico do Castelo, Pico do Facho, Pico de Juliana, Pico da Gandaia, Rocha de Nossa Senhora, and the western slopes of Pico do Concelho. It is also commonly found at lower elevations of the Serra de Dentro, Barranco Branco, and Lombo de Paredes. It is not present on the small islets surrounding the main island, namely Ilhéu de Cima, Ilhéu de Baixo, and Ilhéu de Ferro. Subfossil representatives of Aystricella bicarinata are not known. Seddon (2008: 181) depicts the distribution of H. bicarinata. The four southernmost points indicate the presence of the species also for the southern ridge of eastern Porto Santo. We here infer that these localities are based upon misidentifications with other species, probably some representatives of Wollastonia, i.e., W. jessicae sp. n. or W. klausgrohi sp. n. Ecology. Hystricella bicarinata is commonly found under volcanic rocks scattered on grassland in open fields that are more or less sloping. It has also been found under stones in pine woods (Pico do Castelo) or in cracks and crevices on rocky walls (Pico da Gandaia) or terraced areas (southern slopes of Pico do Castelo). The specimens aestivate on the lower surfaces of stones or rocks, frequently forming large clusters of 40 to 50 individuals attached one to another. Under a single stone of roughly 60 x 40 cm approximately 200 individuals were observed (southern slopes of Pico do Facho). Comparison and comments. The presence of two keels along the body whorl eas- ily distinguishes H. bicarinata from H. echinulata. Where both taxa come into contact in the northern to northeastern part of Porto Santo (Fig. 4) intermediate populations exist however, suggesting a possible hybrid zone (see below). At first glance, and due to the small dimensions, H. bicarinata can be confused with the fossil species H. aucta and H. microcarinata sp. n. (see Fig 74). However, a closer analysis reveals major differences in many features of the shell. Hystricella aucta has a more solid shell with two, almost equally developed keels along the body whorl. The penultimate and the third whorl have a single strong upper keel only. The tubercles in 1. aucta are much smaller and finer than in 1. bicarinata (see Figs 71-73). Fystricella microcarinata sp. n. resembles H. bicarinata with regard to dimensions and overall shape; however, the weak keels along the penultimate and body whorl and the well- developed and rather elevated growth lines allow the separation of the two species. Moreover, the tubercles in H. microcarinata sp. n. are somewhat smaller and more scattered. Hystricella bicarinata can also resemble some of the species included in the newly described genus Wollastonia gen. n. Species such as W. turricula, W. leackociana and W. oxy- tropis differ in the very fine granulation present on the shell’s surface. These species do not possess evident tubercles but only small granules that are remarkably smaller in dimension than the tubercles of H. bicarinata. Moreover, the overall shape of all three above-men- tioned species is different from that of H. bicarinata (see e.g. Figs 100, 119, 130). Other Revision of the genus-group Hystricella R. T) Lowe, 1855 from Porto Santo... 33 MB Figures 38-45. Genitalia and anatomy of Hystricella bicarinata. Pico do Castelo, S slope around the Miradouro: 38 whole genitalia excluding part of OSD, AG and gonads 39 ornamentation of the inner walls of the distal penis, the vagina and the genital atrium 40 transverse section of penial papilla. Pico da Gandaia: 41 whole genitalia excluding gonads 42 ornamentation of the inner walls of the flagellum, the penial complex, the vagina and the genital atrium. Ribeira de Serra de Dentro: 43 jaw. Pico da Gandaia: 44 mantle border. Ribeira de Serra de Dentro: 45 whole genitalia excluding part of OSD, AG and gonads. Scale bars 1 mm. 34 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) Figures 46-53. Genitalia and anatomy of Hystricella bicarinata. Capela de Graca: 46 whole genitalia excluding part of OSD, AG and gonads 47 penial papilla 48 longitudinal section of the penial papilla 49 ornamentation of the inner walls of the flagellum, the penial complex, the vagina and the genital atrium. Ridge between Pico do Facho and Pico da Gandaia: 50 whole genitalia excluding part of OSD, AG and gonads. NE slopes of Pico do Castelo: 51 section of vagina. Capela da Graca: 52 jaw. S slopes Pico do Facho: 53 mantle border. Scale bars 1 mm. (sub-) species assigned to Wollastonia gen. n. such as W. vermetiformis, W. jessicae jessicae sp. n., W. jessicae monticola ssp. n., and W. klausgrohi sp. n. (see Figs 118-120, 147-149, 159- 161, 168-170) are somewhat similar to and could be confused with H. bicarinata. These Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... eB) Figure 54. Distribution of Hystricella bicarinata. taxa can be readily distinguished, however, by their more scalariform shape, somewhat stronger keels, and the sparser arrangement of the tubercles. Moreover, the distributional ranges of Hystricella and Wollastonia gen. n. do not significantly overlap and molecular data strongly supports differentiation at the genus-level (thus also at species level). Nomenclatural and taxonomic remarks. We report below a statement sent to us by Ruud Bank referring to the erroneously introduced name Helix duplicata R. T. Lowe, 1831. “The name Helix [Cochlitoma] bicarinata A. Férussac, 1821 does not exist! Férus- sac simply reclassified Bulimus bicarinatus Bruguiére, 1792 under Helix (Cochlitoma). Thus, Helix bicarinata G. B. Sowerby I, 1824 was at that time (and only for a very short period of time) a secondary homonym of Helix bicarinata (Bruguiére, 1792). Helix duplicata R. T. Lowe, 1831 was introduced to replace the name of Sowerby due to homonymy with the Férussac name, but as stated above, Férussac did not introduce such a name, it was simply a generic change for Bulimus bicarinatus Bruguiére. In my opinion, the name duplicata cannot be considered a nomen novum. It would have been only a nomen novum when R. T. Lowe mentioned Bruguiére as the author of the senior taxon — what he didn‘. Mandahl-Barth (1950), Jaeckel (1966) and Waldén (1983) placed this taxon in Discula and considered Hystricella as a subgenus. Bank et al. (2002) considered the species to belong to a distinct genus (under the name Geomitra). Only recently Seddon (2008) appointed this species to the genus Hystricella and emphasised anatomical dif- ferences with the genus Discula. 36 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) Status and conservation. The opinion of Seddon (201 1a) “[Aystricella bicarinata| has a total extent of occurrence of 10 km? but is present over the eastern part of the island and in these sites it is abundant” has been confirmed by our field researches. For this reason, we think that the assessment as Near Threatened (NT) is appropriate at present. Putative hybrid populations of H. bicarinata and H. echinulata Material examined. All from Porto Santo, CWDM/8, Porto de Pedregal, under stones. 33°06'14"N/16°19'18"W, 20 m, leg. W. De Mattia & J. Macor, May 2014 and May 2015; CWDM/22, CKG/3, CMN/8, SMF 348933/4, NMW Z.2016.013.00005/1 and NMW Z.2016.013.00006/19, Cebecos dos Bades at confluence of two tempo- rary creeks, under stones, 33°06'09"N/16°19'25"W, 34 m, leg. W. De Mattia & J. Macor, May 19 2015; CFW/11147/>10, Ribeira do Pedregal, lower part of the valley near confluence with the tributaries, 33°06'08"N/16°19'32"W, 50 m, leg. FR Walther & E.M. Gryl, Apr. 2 2017; CFW 11150/>10, Ribeiro do Pedregal, upper part, down- stream of the abandoned houses, 33°05'57"N/16°19'32"W, 120 m, leg. E Walther, Apr. 2 2017; CWDM/15, Pico Branco, Faja Pequena at the beginning of the path to Terra Cha, under stones, 33°05'50"N/16°18'44"W, 180 m, leg. W. De Mattia & J. Macor, May 2014; CWDM/10, CKG/1, CMN/5, Pedregal de Dentro near abandoned houses, under stones, 33°05'46"N/16°19'27"W, 195 m, leg. W. De Mattia & J. Ma- cor, May 2015; CWDM/16, Pico de Cabrita, northern slope close to the road, under stones, 33°05'42"N/16°19'23"W, 240 m, leg. W. De Mattia & J. Macor, May 2014; CWDM/11, CMN/5, path to Pico Branco, terraced S slope of Pico Branco, under stones. 33°05'35"N/16°18'25"W, 270 m, leg. W. De Mattia & J. Macor, May 2015. Comments. The genetic analyses revealed an area in the northeastern part of Porto Santo where both, haplotypes assigned to 1. bicarinata and H. echinulata are present (Fig. 5). Some of the specimens are morphologically closer to H. echinulata with regard to the shell, but possess a H. bicarinata mitochondrial haplotype or vice versa. Some specimens may also be conchologically intermediate. Investigation of the genital system showed that much of the genital anatomy is, in fact, identical with that of either H. bicarinata or H. echinulata. However, numerous small papillae that cover the inner walls of the tip of the penial flagellum were observed in several specimens that are lacking in H. bicarinata or H. echinulata (Figs 55—70). Sometimes these papillae can slightly extend distally, towards the vas deferens opening, but never exceed to more than one third of the total flagellum length. Populations that exhibit these features usually occur at lower altitudes compared to 1. echinulata or H. bicarinata, from almost sea level (Porto de Pedregal) to 240 ma.s.l. across the southern slope of Pico Branco and are limited to the northeastern part of the island, where they are distributed along the southern slopes of the Pico Branco massif, Lombo Celado, Cabecos dos Bades with Porto de Pedregal, Pedregal de Dentro, and the northern slopes of Pico de Cabrita. Because of the presence of mitochondrial haplotypes from both H. bicarinata and H. echinulata, these populations are here tentatively inter- preted as hybrid populations. Further investigations that allow an estimation of gene flow Revision of the genus-group Hystricella R. 7 Lowe, 1855 from Porto Santo... a7 | Wf 70 S+DG Figures 55-70. Shells, anatomy and genitalia of the putative hybrid populations of H. bicarinata and H. echinulata. Shells: 55 Pedregal de Dentro 56 Ribeira Formosa 57 Pico de Cabrita 58-59 Cabecos dos Bades 60 Pedregal de Fora. Genitalia and anatomy. Cabecos dos Bades: 61 whole genitalia excluding part of OSD, AG and gonads 62 section of penial papilla 63 penial papilla 64 inner walls of atrium and penis 65 mantle border. Pedregal de Dentro: 66 whole genitalia excluding part of OSD, AG and gonads 67 inner walls of atrium and penis 68 jaw 69 section of penial papilla 70 vaginal glands and vaginal ap- pendage. Scale bars 1 mm. among these populations will, however, be necessary to decide on their definite status. In case that these populations really are of hybrid origin, the specimens with papillae in the penial flagellum may represent an example of transgressive segregation. 38 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) + Hystricella aucta (Wollaston, 1878), stat. n. List of synonyms. 1867 Helix (Octephila) vermetiformis va. « minor Paiva: 48 [non Helix minor mult. auct.]. 1878 Helix (Aystricella) bicarinata var. {8 aucta Wollaston: 162—163. 1950 Discula (Hystricella) bicarinata aucta — Mandahl-Barth: 31, 55. 1983 Discula (Hystricella) bicarinata aucta — Waldén: 267. 2002 Geomitra bicarinata aucta — Bank et al.: 124. 2006 = Discula bicarinata aucta — Cameron et al.: 31. 2008 Aystricella bicarinata aucta — Seddon: 79. 2009 Aystricella bicarinata aucta — Groh et al.: 21 fig. 25. Type material. Despite intensive research in multiple museum collections (NHM, NMW, MMUE, ANSP, NHC, NMS, RAM, SMP) that could have held the type ma- terial of the taxon, no such material could be traced and therefore we deem it reason- able to assume that the type material is lost. To stabilise the present interpretation of Helix (Hystricella) bicarinata var. % aucta Wollaston, 1878 and to clarify its taxonomic status we designate a neotype here, which is deposited in the collection of the SME under the No. SMF 348937 (see Fig. 71). The neotype is consistent with the original description (see below), especially with regard to size, shape and the presence and de- velopment of the two keels on the body whorl. The taxon was originally described from Porto Santo, which is consistent with the origin of the specimen selected as neotype. Loci typici. [aucta] Barbinha, Quaternary slope deposits, "grey layer", 33°04'04"N/16°17'49"W, 8 m, (through the designation of the neotype); [var. minor]: ... rara ad Zimbral d'Aréa. Further material examined. All from Porto Santo, CKG/1, S slope of Pico do Concelho, Quaternary slope deposit, 33°04'35"N/16°18'03"W, 200-230 m, leg. K. & C. Groh, Jun. 29 1980; CKG/2, Barbinha, Quaternary slope deposits, "grey layer", 33°04'04"N/16°17'49"W, 8 m, leg. K. & C. Groh & J. & C. Hemmen, Jul. 4 1983; ANSP H 11789/9 [sub H. bicarinata|, Barbinha, Quaternary slope deposits, "red layer", 33°04'04"N/16°17'49"W, 8 m, leg. J. Gerber, K. Groh & J. Hemmen, Aug. 12 1985. CKG/1, first bay S of Barbinha, Quaternary slope deposit, 33°03'51"N/16°17'46"W, 18 m, leg. K. & C. Groh & J. & C. Hemmen, Jul. 5 1983. CGK/3, Ponta da Ca- naveira, Quaternary aeolinites, 33°02'30"N/16°23'35"W, 50 m, Jun. 24 1983, leg. K. Groh & J. Hemmen; CGK/3, S of Ponta da Canaveira, Quaternary aeolinites, 33°02'30"N 16°23'34"W, 55 m, Oct. 26 1980, leg. K. Groh & J. Hemmen; CKG/2, Vale do Touro, Quaternary slope deposit cut by water erosion in the banks of the val- ley, 33°03'48"N/16°19'17"W, 15 m, leg. K. & C. Groh, Aug. 16 1980; CWDM/14, S slope of Vale do Touro hill, 50 m W of the oil tanks, fossils beds in mixed gravel and mud deposits, 33°03'47"N/16°19'26"W, 15 m, leg. W. De Mattia & J. Macor, May 2015; CFW 12172/1 fragm. [cf. aucta], Ponta da Galé, E end of tunnel, lower level [of slope deposits], coarse, red coloured gravel with large stones, 33°03'44"N/ 16°17'45"W, 30 m, leg. F Walther, Apr. 4 2017; CFW 12173/16, CKG/3, S of Ponta da Canaveira, Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... 39 Figures 71-74. Shells of Aystricella aucta. T\ neotype, SMF 348937 from Barbinha 72 Barbinha 73 Ponta da Canaveira 74 Hystricella microcarinata sp. n., holotype, SMF 348924. Scale bar: 1 mm. (sub-)fossil [slope-]deposits, coarse, red coloured gravel, 33°02'25"N/16°23'41"W, 30 m, leg. FE Walther, Apr. 2 2017; CFW 12175/8, E of Vila Baleira, end of Vale do Touro, (sub-)fossil [slope-] deposits, coarse, black gravel, 33°03'48"N/16°19'15"W, 20 m, leg. F Walther, Apr. 5 2017; CFW 12174/13, CKG/2, E of Vila Baleira, S slope of the hill above Vale do Touro, W of the oil tanks, [(sub-)fossil slope-deposits of] red gravel, 33°03'47"N/16°19'26"W, 25 m, leg. E Walther, Apr. 5 2017. Original descriptions. [aucta]: From Wollaston 1878: There is however an ap- preciably larger form of this species [bicarinata] (cited in the present catalogue as the 'var. {$. Aucta’) to which the subfossil examples might perhaps be better referred, — in which the upper (or medial) keel is a trifle more horizontal and prominent, and the shell is full 3 lines (instead of only approximately 212) across its broadest part — which 40 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) Figure 75. Distribution of Hystricella aucta. was found in Porto Santo by Mr. Watson, and which I have received from him as the 'recent state of the H. vermetiformis, Lowe.’ the 'var. 8. Aucta' of the H. bicarinata in its very much larger size, and in its volutions (the ultimate one of which is not quite so deflected at the aperture) being 7 in number, instead of only 6 or 6%; [var. minor]: From Paiva 1867: Variet. adest « minor, testa minore. Redescription of the shell. Shell small for the genus, with 5.7 regularly increasing whorls, the protoconch with 1.9 whorls. The form of the shell is conical, the concave teleoconch whorls, with two prominent keels situated at the upper 7, respectively 7s of the total height of the last whorl. The last whorl measures 62%, the penultimate whorl 19% of the total shell height. The lower 60% of the body whorl beneath the second keel are rather straight, only a little convex in the middle. The suture is simple, only slightly sunken beneath the second keel of the preceding whorl. The aperture, which is inclined to the vertical axis of the shell in an angle of 49° and is descending in the last 5% of the last whorl in an angle of 38° to the horizontal axis, has an elliptical-ovate form, its width is 42% of the total shell width and its height 27% of the total shell height. It has an only slightly reflected lip, which is completely detached from the body whorl. The eccentric umbilicus, which is approximately 12% as wide as the shell, is in the upper whorls needle stitch-like. The protoconch is smooth. The teleoconch is equipped with few oblique radial ribs, seven in the penultimate quadrant of the body whorl and is additionally covered by numerous coarse tubercles. The number of tubercles in the standard quadrate of the base is 31; the tubercles are of approximately equal size. There are no traces of colouration (Figs 71-73). Revision of the genus-group Hystricella R. 7’ Lowe, 1855 from Porto Santo... 4] Measurements. D 5.5 + 0.5 mm (range 5.1-6.4 mm); H 4.6 + 0.4 mm (range 4.1-5.3 mm); FW 2.8 + 0.2 mm; PA 51 + 2.1; DU 0.5 + 0.03 mm; NT 18 + 6; NW 5.5 + 0.4 (vm = 22). Ratio D/H 1.2; ratio FW/H 0.6. Distribution. ystricella aucta is known only from the southeastern coast of Porto Santo: from the hill immediately east of Vila Baleira (mud and slope deposits at Vale do Touro) to the mud deposits and aeolinites along the southeastern coast (Barbinha, Zimbral da Areia, and Porto dos Frades) (Fig. 75). Comparison and comments. Because of its small size, the species can only be con- fused with the similar sized recent H. bicarinata which, however, is somewhat larger on average, has less developed keels, especially the upper one is less prominent and has coarser and denser tubercles on its shell surface, a narrower umbilicus and a more distinctly descending aperture. The smaller H. microcarinata sp. n., which has a much more rounded globular form, bears less prominent keels on the whorls, has finer and more densely arranged tubercles, has a much narrower umbilicus and a different shape as well as differently positioned angles of the aperture. The similar sized Wollastonia beckmanni sp. n., which has a much flatter shape, has no keels on the whorls, a much wider umbilicus, much more and finer tubercles and differently positioned angles of the aperture. Nomenclatural and taxonomic remarks. Considered by Wollaston (1878) as the fossil variation of H. bicarinata. We here consider it as a distinct species because of the shell differences. Like H. bicarinata, H. aucta has previously (Mandahl-Barth 1950, Wal- dén 1983, Bank et al. 2002) been listed as belonging to Discula and later on to Geomitra. Status and conservation. Extinct before the islands’ scientific exploration in the 19" century, possibly already before human settlement. | Hystricella microcarinata De Mattia & Groh, sp. n. http://zoobank.org/EE848F87-CEBC-4430-9D7F-4D00D2EE2FF9 Figs 74, 76 Type material. SMF 348924, holotype, from loc. typ. (locus typicus), leg. W. De Mattia & J. Macor, May 2016. Locus typicus. Porto Santo, E of Vila Baleira, S slope of the hill above Vale do Touro, 50 m W of the oil tanks, excavated Quaternary mixed gravel, 33°03'47"N/16°19'26"W, 24 m. Diagnosis. Hystricella species with two keels on the body whorl, the upper less developed than the lower one; growth lines rather coarse, giving the impression of an irregular ribbing; tubercles on the base of the shell rather scattered and not as promi- nent as in H. bicarinata. Description of the shell of the holotype. Shell very small for the genus, with 543 regularly increasing whorls, the protoconch with 1.6 whorls. The form of the shell is rounded conical, the convex teleoconch whorls exhibit two flat keels that become more and more expressed; the upper keel is much less developed than the 42 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) Figure 76. Distribution of Hystricella microcarinata sp. n. lower one; keels on the body whorl located in its upper half. The last whorl meas- ures 69%, the penultimate whorl 15% of the total shell height. The lower half of the body whorl is, beneath the distinctly angled periphery, in frontal view nearly straight, only very slightly convex. The suture between the whorls is simple, not sunken. The aperture, which is inclined to the vertical axis of the shell in an angle of 50° and is descending in the last 5% of the last whorl in an angle of 36° to the hori- zontal axis, has an oblique-ovate form, its width is 43% of the shell width, its height 34% of the shell height. It has a slightly reflected lip, which is completely detached from the body whorl. ‘The eccentric umbilicus, which is approximately 11% as wide as the shell, is shaped in the upper whorls like a pinhole. The protoconch is smooth. The teleoconch exhibits a number of oblique radial ribs, 16 in the penultimate quad- rant of the body whorl and is additionally covered by numerous rough tubercles. The number of tubercles in the standard-quadrate of the base is 43. There are no traces of colouration (Fig. 74). Measurements. D 4.1; H 4.2 mm; FW 2.7 mm; PA 49°; DU 0.3 mm; NT 12; NW 4.8 (z = 1). Ratio D/H 0.9; ratio FW/H 0.6. Distribution. The species is only known from the type locality where it appears to be extremely rare (Fig. 76). Its close resemblance to other (sub-) fossil Hystricella taxa may have led to misidentifications during previous collecting and sorting, thus the distribution area of the species may possibly become larger after further investigations. Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... 43 Etymology. The name is a combination of the Greek name for small (tuxe@d¢ = mikrdés) and the Latin name for keeled (carinatus) and alludes to the small size of the shell with a keel at its periphery. Comparison and comments. Being a (sub-) fossil species, only the shell features can be taken into account for comparisons. The globose shape is similar to that of some forms of H. echinulata; nevertheless the overall dimensions and surface sculpture of the shell clearly distinguish H. microcarinata sp. n. Because of its small size, the new species can also be confused with small specimens of H. aucta Wollaston, 1878 which, how- ever, possesses two well-developed keels, has a wider umbilicus, a different shape and differently positioned angles of the aperture, or with Wollastonia beckmanni gen. et sp. n., which has a much flatter spire, has no keels on the whorls, a much wider umbilicus and also a different shape and differently positioned angles of the aperture. Taxonomic remarks. The generic afhliation of this species is based exclusively on the shape of the shell and the presence of keels on the body whorl. A similar arrange- ment of the tubercles and keels can be found also in some Wollastonia gen. n. species that will be described below. Status and conservation. Extinct before the islands’ scientific exploration in the 19" century, possibly already before human settlement. Hystricella echinulata (R. T. Lowe, 1831) Figs 77-94 List of synonyms. 1831 Helix echinulata R. 'T. Lowe: 57, pl. 6 fig. 19. 1846 Helix echinulata — L. Pfeiffer: 140, pl. 91 figs 1-4. 1847 Helix echinulata —L. Pfeiffer in L. Pfeiffer 1847-1848: 190. 1854 Helix echinulata — Reeve in Reeve 1851-1854: pl. 142 figs 9-10. 1854 Helix echinulata — Albers: 36, pl. 9 figs 1-4. 1855 Helix (Hystricella) echinulata — R. 'T. Lowe: 186. 1867 Helix (Octephila) bicarinata var. « echinulata — Paiva: 45. 1878 Helix (Hystricella) echinulata — Wollaston: 160-161. 1950 Discula (Hystricella) echinulata — Mandahl-Barth: 31, 55. 1983 Discula (Hystricella) echinulata — Waldén: 267. 2002 Geomitra echinulata — Bank et al.: 124. 2008 Aystricella echinulata — Seddon: 79, pl. 29 fig. D (and cf. E [under H. bicarinata)), map 178. 2011 Aystricella echinulata — Seddon: e.16727A12801253. Type material. NHM 1968.586, lectotype (herewith designated) of Helix echinu- lata R. 'T. Lowe, 1831 ex coll. R. T. Lowe. For the original figure of Helix echinulata R. T. Lowe, 1831 from R. T. Lowe (1831: pl. 6 fig. 19) and the lectotype of H. echinulata R. T. Lowe, 1831, W = 4.9 mm, Phot. B. Faria, DRAM, see Figs 77 and 78. 44 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) Figures 77-78. 77 original figure of Helix echinulata R. T. Lowe, 1831 from R. T. Lowe (1831: pl. 6 fig. 19) and 78 lectotype of Helix echinulata R. T. Lowe, 1831, NMH 1968.586 ex coll. R. T. Lowe. Scale bar: 1 mm. Further material examined. All from Porto Santo, ANSP H 11829/1, Ribeiro do Pedregal, 33°06'10"N/16°19'25"W, 35 m, leg. K. Groh & J. Hemmen, Jul. 11 1983; ANSP H 11828/16, CKG/5, Pico do Ninho do Guincho, 33°06'07"N/16°18'51" W, 100 m, leg. K. Groh & J. Hemmen, Jul. 11 1983; CKG/1, Ribeiro do Golfeiras E Camacha, 33°05'47"N/16°19'33"W, 170 m, leg. K. & C. Groh, Oct. 25 1980; ANSP H 11831/3, CKG/1, CMN/1, SW slope of Lombo Branco, under stones, 33°05'46"N/16°18'46"W to 33°05'38"N/16°18'13"W, 150-400 m, leg. K. Groh & J. Hemmen, Jul. 8 1983; ANSP H_ 11830/4, Terra Cha, top, 33°05'46"N/16°18'14"W, 360 m, leg. J. Ger- ber, K. Groh & J. Hemmen, Aug. 14 1985; CFW 11162/5, Pico Branco, Terra Cha (350 m E of Pico Branco), 33°05'39"N/16°17'58"W, 340 m, leg. E Walther, Mar. 31 2017; CWDM/19, CMN/5, Terra Cha, near the "Casa Forestal", under stones in pine wood. 33°05"39"N/16°20'00"W, 340 m, leg. W. De Mattia & J. Macor, May 2014; CWDM/6, Terra Cha-Pico Branco ridge, under stones, 33°05'38"N/16°18'01"W, 335 m, leg. W. De Mattia & J. Macor, May 2015; ANSP H 11833/3, summit of Pico Bran- co, 33°05'38"N/16°18'14"W, 450 m, leg. K. Groh & J. Hemmen, Jul. 8 1983; ANSP H 11836/15 [sub A. leacockiana], Pico Branco, 33°05'38"N/16°18'14"W, above 350 m, leg. J. Gerber, K. Groh & J. Hemmen, Aug. 14 1985; CKG/16, Terra Cha, top, 33°05'36"N/16°18'14"W, 360 m, leg. J. Gerber, K. Groh & J. Hemmen, Aug. 14 1985; CKG/2, Terra Cha, 33°05'31"N/16°18'16"W to 33°05'38"N/16°18'13"W, 320—400 m, leg. K. Groh & J. Hemmen, Jul. 8 1983; CWDM/14, path to Pico Branco, eastern steep part of path to Pico Branco, under stones, 33°05'29"N/16°18'14" W, 320 m, leg. W. De Mattia & J. Macor, May 2014; CFW 11148/<10, SW slope of Pico Branco ca. 250 m SW of the top, 33°05'29"N/16°18'13" W, 310 m, leg. E Walther & E. M. Gryl, Mar. 31 2017; ANSP H 11826/7, approx. 1 km N Serra de Dentro, 33°05'24"N/16°18'27"W, 165 m, leg. J. & C. Hemmen, Jan. 6 1981; ZMH 24292/1, Madeira Archipelago, with- out exact locality data, ex coll. Altonaer Museum, ex coll. O. Semper, ex coll. Dohrn; ZMH 24291/1, Porto Santo, without exact locality data, ex coll. Museum Klagenfurt. Revision of the genus-group Hystricella R. 7 Lowe, 1855 from Porto Santo... 45 Figures 79-82. Shells of Hystricella echinulata. 19-80 Terra Cha 81 S slope of Pico Banco 82 steep path from Pico Branco to Terra Cha. Scale bar 1 mm. Locus typicus. Hab. in monte “Pico Branco” dicto Insulae Portus S". Original description. From Lowe 1831: H. testa parvula, conoidea, sub-pyrami- data, depressiuscula, supra planulata, perforata, carinata, tota scaberrima, fusca, supra fasciata: spira pyramidata elevata; sutura distincta, impressa; anfractibus convexis; ul- timi carina acuta, distincta, supra marginata sc. sulco expressa vel exarata; omnibus granulis distinctissimis, conferitis, asperrimis scobinatis et quasi echinulatis; umbilicus parvo, sub-spirali, aperto; apertura rotundata; peristomate continuo, circinato, dis- juncto, reflexo. Axis 2 lin. Diam. 2%. Anfr. 6. Diagnostic features of the shell. Shell as in the genus description. The main diag- nostic feature is the presence of only one keel along the body whorl. The overall shape of the shell of H. echinulata is always conical but not scalariform as in H. bicarinata. The whorls are rounded and lack the typical “shoulder”. The sutures are deep, but less deeply marked than in H. bicarinata (see Figs 79-82). 46 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) Figures 83-93. Anatomy and genitalia of Hystricella echinulata. Terra Cha: 83 whole genitalia excluding gonads 84 ornamentation of the inner walls of the distal penis, the distal vagina and the genital atrium 85 ornamentation of the inner walls of the flagellum, the penial complex, the vagina and the genital atrium 86 penial papilla 87 whole genitalia excluding part of OSD, AG and gonads 89 penial papilla and ornamentation of the inner walls of the flagellum 91 mantle border 92 section of penial papilla. Faja Pequena: 88 section of vagina 90 jaw 93 whole genitalia excluding gonads. Scale bars 1 mm. Measurements. D 5.2 + 0.3 mm (range 4.6-5.8 mm); H 4.8 + 0.5 mm (range 4,.3—5.0 mm); FW 2.5 + 0.2 mm; PA 42.6 + 8.3; DU 0.5 + 0.08 mm; NT 23 + 9; NW 5.6 + 0.1 (7 = 40). Ratio D/H 1.1; ratio FW/H 0.5. Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... 47 Figure 94. Distribution of Hystricella echinulata. Body. Body as in the genus description. The colour of the head and the neck is somewhat darker than in H. bicarinata. Genital anatomy. As in the genus description. Main diagnostic features are as follows. The free oviduct is usually very short and four times shorter than the vagina. The penial flagellum is usually as long as the epiphallus. It has a remarkably cylindrical shape and a blunt apex. Its internal walls are completely smooth, without any papillae. The genital atrium is always elongated. See Figs 83-93. Ecology. Hystricella echinulata is recorded from low (20 m a.s.l.) to relatively high altitudes (450 m a.s.l.). Except for the slightly higher altitudinal range, this species shares the same ecology as H. bicarinata; it is commonly found under volcanic rocks scattered on grassland in open fields that are usually sloping. It has also been found in cracks and crevices or among scree on steep slopes. Distribution. Hystricella echinulata is endemic to the island of Porto Santo (Ma- deiran Archipelago, Portugal). The species is restricted to the northeastern, mountain- ous part of the island and present only in the Pico Branco-Terra Cha massif area. It is commonly found along Faja Pequena and the ridge and the south-exposed slope eastwards of Terra Cha (Fig. 94). No populations are known from the offshore islets. Comparison and comments. See under H. bicarinata and/or the section on puta- tive hybrids of H. bicarinata and H. echinulata above. Status and conservation. According to Seddon (2011b) the species is Least Concern (LC), but in our opinion, the species should be considered as Endangered 48 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) (EN Bla, b(i, ii, iv), 2a, b(i, ii, iv)) because its area of occupancy and extent of occur- rence is less than 8 km’ and because it is mostly restricted to the highest elevations in the north-eastern part of Porto Santo and only occurs at scattered and isolated places with suitably humid microclimates. Mayor threats to the species include an observed decline of suitable habitats, extent of occurrence and area of occupancy over the past 30 years, i.e. as the result of longer drought periods which are possibly associated with effects of climate change, an increase of fire incidents, grazing by goats and to a lesser extent also hiking as tourism is increasing on Porto Santo. As most of the known localities are concentrated in the Pico Branco-Terra Cha area in the north- easternmost part of Porto Santo (Fig. 94), we consider that the species is not present at more than five locations. + Hystricella echinoderma (Wollaston, 1878) Figs 95-96 List of synonyms. 1878 Helix (Hystricella) echinoderma Wollaston: 159-160. 1894 Geomitra echinoderma — Pilsbry in Pilsbry 1893-1895: 242. 1931 Geomitra (Actinella) echinoderma —Nobtre: 88. 1950 Discula (Hystricella) echinoderma — Mandahl-Barth 1950: 31, 55. 1983 Discula (Hystricella) echinoderma — Waldén: 267. 2002 Geomitra echinoderma — Bank et al.: 124. 2008 Aystricella echinoderma — Seddon: 80. Type material. NHM 1875.02.02-52, lectotype (herewith designated) from loc. typ.; NHM 1875.02.02-53 to -54, 2 paralectotypes, from loc. typ. Locus typicus. Portum Sanctum, semifossilis; recens haud observata. Original description. From Wollaston 1878: T. trochiformis, subtus subplanulata perforata, undiquegranulis magnis obtusis sat dense obsita; spira elevata; anfractibus convexis, subgibbosis, ultimo subtectiformi acute carinato (carina simplici, solum an- tice gradatim obsolete subduplici); umbilico punctiformi, aperto; apertura subovali-ro- tundata, labris continuis conjunctis, peristomate simplici, expanso, subrecurvo, tenui, relevato. — Long. axis 2 ¥2 lin.; diam. 3 2. Description of the lectotype. Shell large for the genus, with 6%4 regularly increas- ing whorls, the protoconch with 1.9 whorls. The form of the shell is high conical, the convex to quadrangular teleoconch whorls showing two more and more expressed angulations which form in the last whorl two rounded keels, the upper less developed than the lower one. ‘The last whorl measures 59%, the penultimate whorl 14% of the total shell height. The lower 70% of the body whorl are situated below the peripheral keel which is slightly constricted by a concavity below the periphery in frontal view. The base is convex, slightly straightening towards the umbilicus. The two keels of the body whorl are located in the upper % and % of the total height of the body whorl. The suture between the whorls is simple but deeply sunken. The aperture, which is Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... 49 Figure 95. Lectotype of Hystricella echinoderma, NHM 1875.02.02-52. Scale bar 1 mm. inclined to the vertical axis of the shell in an angle of 48° and descending in the last 5% of the last whorl in an angle of 38° to the horizontal axis, has an oblique-ovate form, its width amounts to 43% of the total shell width, its height to 34% of the total shell height. It has a slightly reflected lip, which is completely detached from the body whorl. The centered umbilicus, which measures 10% of the shell’s total width, is in the last whorls circular, but completely closed deeper inside. The protoconch is smooth, the teleoconch shows a number of oblique radial ribs, 15 in the penultimate quadrant of the body whorl, and is additionally covered by numerous irregularly arranged, elon- gate, rough tubercles. The number of tubercles in the standard-quadrate of the base is 81. There are no traces of colouration. See Fig. 95. Measurements. D 7.0 mm; H 5.9 mm; FW 3.5 mm; PA 42.2°; DU 0.5 mm; NT = 35; NW 6.6 (7 = 1). Ratio D/H 1.2; ratio FW/H 0.5. Distribution. Hystricella echinoderma is only known from Fonte da Areia in the northern part of Porto Santo (Fig. 96). Comparison and comments. Hystricella echinoderma can be confused on first glance with comparatively large-sized species of the genus Wollastonia like W. vermeti- formis, W. ripkeni sp. n., and W. falknerorum sp. n.; from these it is distinguishable by the lack of one or two sharply pointed keels, respectively, a much rougher sculpture, a non-eccentric umbilicus, and the stepped, high conical form. From the similarly sized W. subcarinulata and W. inexpectata sp. n. it is separated by a wider or non-eccentric umbilicus, a relatively higher shell, the quadrangular instead of convex rounded whorls, a coarser granulation and the oblique-ovate instead of straight horizontally elliptical aperture. From the similar shaped H. echinulata and H. microcarinata sp. n. it is easily distinguishable by the much larger size. Taxonomic remarks. Wollaston (1878) compares H. echinoderma with the rather sim- ilar H. echinulata and calls it a monstrous sister species of that species, citing as examples of 50 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) Figure 96. Distribution of Hystricella echinoderma. pairs of nominal species with a distinct size difference Helix vermetiformis and H. bicarinata, H. Lowei and H. portosanctana as well as H. bowdichiana and H. punctulata. Status and conservation. Extinct before the islands’ scientific exploration in the 19" century, possibly already before human settlement. Wollastonia gen. n. http://zoobank.org/46BC6BCC-D9E1-4753-B9A2-062537F0B077 Remarks. ‘The synonymy is the same as for the genus Hystricella (see above). Type species. Helix |Helicella] turricula R. T. Lowe, 1831 — herewith by original designation. Included taxa. Wollastonia turricula (R. T. Lowe, 1831), comb. n., W. vermeti- formis (R. T. Lowe, 1855), comb. n., W. ripkeni De Mattia & Groh, sp. n., W. falknero- rum Groh, Neiber & De Mattia, sp. n., W. leacockiana (Wollaston, 1878), comb. n., W beckmanni De Mattia & Groh, sp. n., W. jessicae jessicae De Mattia, Neiber & Groh, sp. n., W. jessicae monticola De Mattia, Neiber & Groh, ssp. n., W. klausgrohi De Mattia & Neiber, sp. n., W. oxytropis (R. T. Lowe, 1831), comb. n., W subcarinulata (Wollaston, 1878), comb. n., W. inexpectata De Mattia & Groh, sp. n. Description of the genus. Shell. The shell is dextral and hairless. Its shape can be very variable, from conical, elongated and scalariform with deep sutures to rather Revision of the genus-group Hystricella R. 7’ Lowe, 1855 from Porto Santo... 51 flattened, with very shallow sutures. The protoconch is from whitish to completely dark brown with 1.5 to 2.5 whorls. It is almost smooth along the first whorl and shows fine radial striae and extremely small, scattered tubercles along its remaining portion. The teleoconch has from 4.3 to 7.0 rapidly increasing whorls. It is usually dark brown with brick-red and/or dark violet shades in colour. The dark areas of the shells are mottled with more or less light brown to whitish areas, usually placed longitudinally and slightly slanting. In some species, the lighter areas tend to be more evident along the keel. No band pattern is visible along the upper whorls. On the lower part of the last whorl two dark, more or less broad bands may be present. In some specimens, the two bands merge together forming a single wide, dark band. Sometimes the bands are interrupted by yellowish to whitish sections. The area around the umbilicus is usually the lightest in colour. Some species are usually covered entirely with soil or sand grains which probably serve as a camouflage. The spire is very variable in height, ranging from flattened, almost discoidal to slender and remarkably conical in shape. The number and the extent of the keels are variable from species to species. Some species have one or two distinct keels starting already from the second whorl of the teleoconch, while others have a single, more or less strong keel only along the last whorl. Some species have a lower, distinct principal keel somewhat protruding the body whorl, with an upper secondary and much less evident keel. The whorls can be either convex or flat. In some species, the whorls form a “shoulder”, giving the whorls and angular contour. The sutures are usually deep, even if in some species they may also be rather shallow. The external surface has from very fine to strong, clearly visible, irregularly spaced, growth lines. Irregularly disposed tubercles are found all over the teleoconch. ‘The di- mensions and the arrangement of these tubercles varies considerably from species to species, from large and scattered tubercles to a very dense pattern of small papilla-like tubercles all over the shell. In the species with larger tubercles, these are somewhat dens- er along the keels of the penultimate and last whorls, giving to the keel(s) the appearance of (a) rough chord(s). The last whorl is usually large, descending near the aperture with a contribution ranging from 40% to 60% of the total shell height. The umbilicus is open but very narrow, either concentric or eccentric, and measures approximately 10% of the maximum shell diameter. The aperture is elliptical with a faint thickening along the columellar portion of the aperture. Sometimes this thickening can also extend as far the parietal side of the aperture. The peristome is continuous, slightly to evidently reflected, with the columellar margin somewhat thicker and more reflected. The shells of the species belonging to Wollastonia gen. n. show a remarkable plas- ticity, with regard to dimensions, shape and ornamentation. This variability undoubt- edly exceeds that of the genus Hystricella. Thus, for each taxon, the shell’s features will be described in detail. Body. The head and neck are usually dark grey to grey. The sides and the posterior upper section of the foot are whitish. In some species, the pigmented ommatophoral retractor muscles are visible through the skin of the back of the cephalic area. The foot is white and the sole is longitudinally divided into three areas. The central area 52 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) is smooth, whereas the two lateral areas are equipped with bands of muscles roughly arranged in a chevron pattern. The mantle border is grey to dark grey with five more or less developed lobes. The ratio of lateral to the dorsal lobes varies from specimen to specimen, also in the same population. In some specimens one of these lobes (either lateral or dorsal) may be totally missing. The walls of the pallial cavity are colourless, without any stripes or spots. A strong pulmonary vein is visible. The jaw is odonto- enathous and very variable in shape, from almost straight to markedly arched. There are many transversal, smooth ridges, ranging from eight to 25 in number. The right ommatophoral retractor is independent from both penis and vagina. Genitalia. The general arrangement of the genitalia is semi-diaulic monotrem- atic. A convoluted to almost straight hermaphroditic duct arises from a multi-lobated gonad. The albumen gland is long and thin and connected to a variably long sperm- oviduct consisting of a prostatic and a uterine portion. The prostatic part extends into a thin vas deferens, roughly as long as the sperm-oviduct, and terminating in the penial complex. The distal portion of the uterine part extends into the free oviduct and turns into a vagina along its course at the level of the duct of the bursa copulatrix. The free oviduct can be as long as the vagina or also three to four times longer. The duct of the bursa copulatrix is usually wide, approximately as long as the penis and uniform in diameter. It extends into a variable, oval to roundish bursa copulatrix. In some species, the transition area between the duct and the bursa itself is not very distinctly delimited; the duct more or less abruptly widens and transforms into the bursa. The spermatophore is unknown. One tuft of digitiform glands arises from the proximal part of the vagina. The glands have usually two or three, approximately equally long and very rarely bifurcated branches. A short and thin vaginal appendix arises from the vagina’s wall, immediately distal of the glandular tuft. Very smooth, rather wide, and little elevated, irregularly spaced pleats run longitudinally along the inner surface of the vagina, reaching into the genital atrium as far as the genital ori- fice. The atrium can be short and wide or long and thin. Its internal walls can either be smooth or with large and soft pleats running longitudinally as far the genital orifice. The penial complex consists of a flagellum, an epiphallus (which extends from the insertion of the vas deferens to the penial retractor muscle) and a penis that inserts into the atrium. The penial flagellum is short, remarkably cylindrical and with a blunt apex. It is usually as long as the epiphallus. In one species, W. oxytropis, the flagellum is very short (4 of the epiphallus) but remarkably pointed. The internal walls of the flagellum can be either completely smooth or, in one species, can have a digitiform, pointed papilla originating from the proximal end of the flagellum and orientated to- ward the penial papilla. The epiphallus is usually short to moderately long. Its internal walls have a variable number of longitudinal pleats that can be more or less developed and elevated. The retractor muscle is large, strong and is of a variable length. The penis lacks a muscular or glandular sheath. It is thick-walled and approximately four times longer than the flagellum. It is usually cylindrical to sometimes slightly swollen in its distal part. Sometimes, a thin sheath consisting of connective tissue envelops the distal penis, causing a partial and longitudinal minor compression. The inner walls Revision of the genus-group Hystricella R. 77 Lowe, 1855 from Porto Santo... 53 of the penis are smooth or with irregular and spaced pleats, which run longitudinally and reach the genital atrium. The section where the penial papilla is located is usually detectable from the outside by virtue of a fine circular swelling corresponding to the origin of the papilla. The penial papilla is usually small, reaching maximally /% of the total penial length and is conical to subcylindrical in shape. It has smooth external walls with the opening emerging apically. The penial papilla channel is thin and nar- row. The inner lumen of the penial papilla is occupied by a spongy and sturdy tissue, which directly connects with the walls of the epiphallus. The longitudinal section of the penial papilla shows that its walls are the continuation of the penial walls that abruptly bend inwards. Jaw and radula. As for the genus Hystricella, no notable variability was found among the species of the genus Wollastonia gen. n. and the two genera share almost an identical jaw-radular apparatus. The jaw is odontognathous, arched and with rough wrinkles (irregular ribs). The radula ribbon is typical helicoid, it is elongated but not very slender. A central tooth is present, tricuspid, the main cusp (endocone) is rhom- boid, pointed; the ectocones are much smaller than the endocone, they are triangular, pointed. There are 19-20 laterals and marginals, which do not distinctly differ from each other. Their shapes rather change gradually from the first laterals towards the marginals. Laterals are bicuspid, with rhomboid or triangular and pointed endocones. The ectocones are much smaller, pointed, and triangular. The endocones of the cen- tral and first laterals are approximately of the same size. Both the endocone and the ectocone of the laterals gradually become bifurcated towards the marginal teeth, but the ectocones occasionally might have three cusps as well. The cusps of the marginals are gradually decreasing in size; therefore, the outermost marginals appear serrated. ‘The jaw is variable in shape: from almost straight to markedly arched. There are many transversal and smooth ridges, ranging from 13 to 20. For Wollastonia jaws and radu- lae, ‘see ‘Fig. 98. Distribution. The genus Wollastonia is endemic to the island of Porto Santo (Ma- deiran Archipelago, Portugal) and some surrounding islets (Fig. 97). Along the eastern part of the main island, the genus is restricted to the southern mountainous part. It is patchily distributed from the hilly area east of Vila Baleira towards the east to Por- tela, Zimbreiro and along the highest southern peaks of Porto Santo, namely Pico do Magarico and Pico do Baixo as far as Ponta da Galé. It is also present along the eastern coast area of Pico do Concelho. Along the western part of Porto Santo, Wollastonia is found on Pico de Ana Ferreira. With regard to the surrounding offshore islet, Wollas- tonia is present on Ilhéu de Cima and Ilhéu de Cenouras at the eastern and Ilhéu de Ferro at the western end of the island. Subfossil representatives of the genus are found mainly in the mud and aeolinite deposits along the southeastern (Vale do Touro, Ponta do Passo, Barbinha, Calhau da Serra de Fora) and northern (Fonte da Areia) coasts. Wollastonia may be older than Aystricella as suggested by the somewhat deeper diver- gences in the molecular trees (Fig. 3). Ecology. Representatives of the genus Wol/lastonia are commonly found under volcanic rocks scattered on grassland in open fields and more or less steep mountain 54 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) Figure 97. Distribution of the genus Wollastonia gen. n. Filled circles refer to recent and open circles to fossil records. slopes. Specimens aestivate on the lower surfaces of the rocks, frequently forming clus- ters of individuals attached to one another. Etymology. Named to honour the late British malacologist and entomologist Thomas Vernon Wollaston for his indispensable contributions to the taxonomy and nomenclature of the terrestrial snails of the mid-Atlantic islands. Taxonomic remarks. The genus Wollastonia gen. n. is separated here from Hystri- cella both upon molecular and morphological features. Moreover, the analysis of the distribution of its species supports this view. The phylogenetic analyses recovered three clades, in which morphologically mostly similar species were grouped together and that were supported in at least two of the analyses. The clade including recent species hitherto assigned to Hystricella (except H. bicarinata and H. echinulata), supported in the BI and MP analyses (Figs 3, 5), are here regarded as representing a new genus, Wollastonia gen. n., because representatives of the morphologically distinct Callina species are interspersed between Wollastonia gen. n. and Aystricella in the strict sense receiving significant/meaningful support in the BI and MP analyses of the cox] data alone and the concatenated mitochondrial and nuclear data. This clade includes the recent species W. leacockiana comb. n., W- turricula comb. n., and W. oxytropis comb. n. as well as three additional recent taxa that will be formally described below: W. klausgrohi sp. n., W. jessicae sp. n., and W. jessicae Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... 55 led rep vane ve 2015/03/22 LD27 x15k 50um Figure 98. Jaws and radulae Wollastonia gen. n. A W. leacockiana Pico de Ana Ferreira B W. oxytropis Pico do Concelho C W. jessicae jessicae sp. n. Vale do Touro. monticola ssp. n. (Fig. 5). The relationships of most of the taxa within Wollastonia gen. n. could, however, not be well resolved (Fig. 5) on the basis of the phylogenetic analyses of the cox/ data. Shell size and shape are remarkably more variable in the genus Wollastonia as com- pared to Aystricella. Most of the Wollastonia gen. n. species have the shell surface cov- ered with small and very dense tubercles that can be compared to a very fine granu- lation. The typical Hystricella’s “spiny” contour (Lat.: hystrix = spiny, deriving form Greek vorpié (hustrix = porcupine) is replaced by a smoother one. The genital morphology of the two genera shows no substantial differences. Nev- ertheless, some Wollastonia gen. n. species present a higher degree of complexity with regard to the inner ornamentation of the atrium and the flagellum. Structures such as more or less developed pleats, fleshy pads and/or fringes are found in Wollastonia gen. n. that are never present in Aystricella. The distribution ranges of Hystricella and Wollastonia gen. n. are separated and not considerably overlapping. Wollastonia colonises the southern and southwestern parts of Porto Santo, whereas Hystricella is currently found exclusively in the central and northern parts. Most of the morphological characters that clearly distinguish Hystricella from the other native Geomitridae of Porto Santo are also valid for Wollastonia. gen. n., e.g. the continuous and detached peristome and the shape of the penial flagellum that is always short and with a remarkably blunt apex (except in W. oxytropis stat. n.). For remarks on the differentiating features, see the respective section in the re-description of the genus Hystricella above. 56 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) Wollastonia turricula (R. T. Lowe, 1831), comb. n. Figs 99-112 List of synonyms. 1831 Helix turricula R. T. Lowe: 58, pl. 6 fig. 21. 1846 Helix turricula — L. Pfeiffer: 141, pl. 91 figs 5—7. 1847 Helix turricula — L. Pfeiffer in L. Pfeiffer 1847-1848: 190. 1854 Helix turricula— Reeve in Reeve 1851-1854: pl. 138 fig. 867. 1854 Helix (Ochthephila) turricula — Albers: 37, pl. 9 figs 11-13. 1855 Helix (Hystricella) turricula — R. T. Lowe: 186. 1867 Helix (Octephila) turricula — Paiva: 47. 1878 Helix (Aystricella) turricula — Wollaston: 163-165. 1878 Helix (Hystricella) turricula var. { pererosa Wollaston: 165. 1888 Helix turricula—Tryon in Tryon and [Pilsbry] 1888: 33, pl. 7 fig. 91. 1894 Geomitra bicarinata — Pilsbry in Pilsbry 1893-1895: 242. 1923 Ochthephila turricula — Watson: 283-293, pl. 6 figs 1-10. 1931 Geomitra (Actinella) bicarnata — Nobre: 88, fig. 38. 1950 Discula (Hystricella) turricula— MandahI-Barth: 31, 55. 1950 Discula (Hystricella) turricula pererosa — Mandahl-Barth: 31, 55. 1977 Discula (Hystricella) turricula — Pettitt: 147-150. 1983 Discula (Hystricella) turricula — Waldén: 267. 2002 Geomitra turricula — Bank et al.: 124. 2008 Aystricella turricula — Seddon: 79, pl. 29 fig. b, map 181. 2009 Adystricella turricula £. pererosa — Groh et al.: 21 fig. 28. 2011 Aystricella turricula — Seddon: e.16723A12800477. Type material. [turricula], NMH 1968.578, lectotype (herewith designated), from loc. typ., ex coll. R. T. Lowe; NMH 1948.7.8.35, 1 paralectotype, from loc. typ., ex coll. R. T. Lowe. The original figure of Helix turricula R. T. Lowe, 1831 (from Lowe 1831: pl. 6 fig. 21) is depicted in Fig. 99, the lectotype (Phot. P. Crabb, NHM) in Fig. 100; [pererosa], RAM EXEMS-1720-1909-d39-74a, lectotype (herewith designated), from loc. typ., ex coll. Linter, ex coll. T. V. Wollaston; RAM EXEMS-1720-1909- d39-74b, 1 paralectotype, from loc. typ., ex coll. Linter, ex coll. T. V. Wollaston (see Figs 113-115). Loci typici. [turricula], Hab. in Insula quadam «Ilheo de Cima» dicta, juxta Insulam Portum S"; [pererosa], ‘Ilheo de Cima’. Further material examined. All from Porto Santo. Fossil: CKG/10, CWDM/2, ZMH 116090/1 fragm [ex coll. E. Clauss], Quaternary slope deposits at the SW coast of the Ilhéu de Cima, 33°03'15"N/16°17'02"W, 40 m, leg. K. & C. Groh & J. & C. Hemmen, Jun. 26 1983 and leg. J. Gerber, K. Groh & J. Hemmen, Aug. 17 1985. Recent: CKG/9, Ilhéu de Cima, top, under stones, 33°03'13"N/16°16'48"W, approx. 100 m, leg. K. & C. Groh, Oct. 25 1980; CKG/7, CMN/29, ANSP H 11918/42, ZMH 120611/1 [ex coll. W Fauer], I[héu de Cima, top, under stones, 33°3'13"N/16°16'48" W, approx. 100 m, leg. K. & C. Groh & J. & C. Hemmen, Jun. 26 Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... 57 Figures 99-102. 99 Wollastonia turricula original figure as in R. T. Lowe, 1831 (from Lowe 1831: pl. 6 fig. 21). Shell of Wollastonia turricula \00 shell of the lectotype from loc. typ., ex coll. R. T: Lowe NMH 1948.7.8.35 101, 102 shells from Ilhéu de Cima. Scale bar 1 mm. 1983; ANSP H 11918/17, Ilhéu de Cima, top, under stones, 33°3'13"N/16°16'48"W, approx. 100 m, leg. J. Gerber, K. Groh & J. Hemmen, Aug. 17 1985; ZMH 24296/1, Porto Santo, without exact locality data, ex coll. Altonaer Museum, ex coll. O. Sem- per, ex coll. Dohrn; ZMH 24297/1, Porto Santo, without exact locality data, ex coll. Museum Klagenfurt; ZMH 24298/1, Porto Santo, without exact locality data, ex coll. Altonaer Museum. Original descriptions. [turricula]: From Lowe 1831: H. testa turrita, pyramidata, sub-cylindrica, bicarinata, perforata, tota minute et confertissime granulata, fusca, fere unicolore, vel supra obsolete fasciata: spira valde elevata, obtusissima; sutura distincta; anfractibus bicarinatis, carinis equalibus, prominentibus, distinctis, sulco divisis: ap- ertura rotunda; peristomate continuo, circinato, disjuncto, tenui, reflexo. Axis 4 lin. Diam. 3. Anfr. 8-8'; [pererosa]: From Wollaston 1878: ... a keel, very largely devel- 58 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) oped in the ‘«. pererosa, in the centre of each, causing the basal volution to be strongly bicarinated ..... var. {8. pererosa. — Plerumque obscurior, spira breviore, anfractibus in medio multo grossius carinatis (carina altissima), ultimo sensim latiore necnon antice obsolete subtortuoso, fere quasi superimposito, apertura submajore. Redescription of the shell. The shell is dextral and hairless. It is remarkably elon- gated and scalariform with deep sutures. The protoconch is from whitish to completely dark brown with 1.5 to 2 whorls. It is finely granulated along the first whorl and shows also fine radial striae and extremely small, scattered tubercles along its remaining por- tion. The teleoconch has from 6.9 to 7.2 rapidly increasing whorls. It is usually reddish brown, with brick red and/or dark violet shades in colour. The dark areas of the shell are mottled with more or less light brown to whitish areas, usually placed longitudi- nally and slightly slanting. In some specimens, the lighter areas tend to be more evident along the keels. No band pattern is visible along the upper whorls. On the lower part of the last whorl there is sometimes one dark, indistinct band that is usually very nar- row. [he area around the umbilicus is usually the lightest in colour. The shell is usually covered by fine debris serving as a camouflage. The spire is very high and remarkably conical in shape. Two evident keels start already from the second whorl of the teleoconch. The keels gradually strengthen, reaching their maximum extent along the body whorl. The lower and the upper keel are more or less equally developed, the lower only sometimes slightly more pronounced. The lower whorls (especially the fourth and fifth) form a “shoulder” giving the whorls an angular contour. ‘The sutures are deep and well marked. The external surface does usually not show fine growth lines, except for some strong, more or less regularly spaced ribs along the top and bottom of the body whorl. Irregularly arranged, fine tubercles are present all over the teleoconch, also along the edges of the keels the tubercles remain well-separated not forming a peripheral chord. The last whorl is not distinctly wider than the penultimate whorl, nevertheless abruptly descending near the aperture. The umbilicus is very narrow, almost closed, and somewhat eccentric. The aperture is elliptical with a faint thickening along the columellar portion of the stoma. Sometimes this thickening can also extend as far as the parietal side of the aperture. The peristome is continuous and detached from the body whorl, reflected, with the columellar margin somewhat thicker and more dis- tinctly reflected (Figs 101-102). Measurements. D 6.6 + 0.4 mm (range 6.1-7.0 mm); H 8.8 + 0.2 mm (range 6.1-9.0 mm); FW 4.3 + 0.1 mm; PA angle 32.1 + 2.1°; NT > 100; NW 7.0 + 0.1 (z = 8). Ratio D/H 0.8; ratio FW/H 0.5. Body. The head and the neck are usually dark grey to grey. The sides and the pos- terior upper section of the foot are whitish. The foot is white and the sole is longitudi- nally divided into three areas. The central area is smooth, whereas the two lateral areas are equipped with bands of muscles that are roughly arranged in a chevron pattern. The mantle border is dark grey with five more or less developed lobes. ‘The walls of the pallial cavity are colourless, without any stripes or spots. A strong pulmonary vein is visible. The jaw is odonthognatous and its shape is markedly arched. There are up to Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... ay) 25 smooth transverse ridges. The right ommatophoral retractor is independent from both penis and vagina. Genital anatomy. The albumen gland is short and as long as the sperm-oviduct. The prostatic part of the sperm-oviduct extends into a vas deferens that is approxi- mately as long as the sperm-oviduct, which is inserting into the penial complex. The free oviduct is two times longer than the vagina. The duct of the bursa copulatrix is rather wide, slightly shorter than the penis, and uniform in diameter. It terminates in a roundish bursa copulatrix. The transition area between the duct and the bursa is very sharply delimited, abruptly widening and turning into the bursa. The spermatophore is unknown. One tuft of digitiform glands arises from the proximal part of the vagina. There are usually two rather wide glands that are approximately equally long and very rarely branched. A short and thin vaginal appendix arises from the wall of the vagina, just distal of the glandular tuft. Very smooth, rather widely and irregularly spaced and little elevated pleats run lon- gitudinally along the inner surface of the vagina, reaching into the genital atrium as far as the genital orifice. The atrium is rather wide. Its internal walls are equipped with large and soft pleats, running longitudinally towards the genital orifice. The penial flagellum is short, remarkably cylindrical and with a blunt apex. It is usually as long as the epiphallus. Its internal walls are equipped with a digitiform papilla that originates at the proximal end of the flagellum and which is orientated towards the penial papilla. The epiphallus is approximately half as long as the penis. Its internal walls are usually equipped with two to three longitudinal pleats. These pleats are arranged more or less “metameric”. The retractor muscle is large, strong and is of variable length. The penis lacks any muscular or glandular sheath. It is thick-walled and approximately four times longer than the flagellum. It is cylindrical and slightly swollen in its proximal portion. The in- ner walls of the penis are equipped with a large longitudinal, fleshy, and smooth pleat, running from the penial papilla almost as far as the atrium. Some minor, smooth, longitudinal pleats are usually also present. The penial papilla is short but somewhat bulky. Its surface is smooth, with the opening emerging apically; its channel is rather narrow. The inner lumen of the penial papilla is filled with a spongy and sturdy tis- sue, which directly connects with the walls of the epiphallus. The longitudinal section of the penial papilla shows that its walls are the continuation of the penial walls that abruptly bend inward. See Figs 103-111. Ecology. Wollastonia turricula is found under volcanic rocks scattered on grassland in open fields. Distribution. Restricted to the slopes and plateau of the islet Ilhéu de Cima, off the southeastern coast of Porto Santo. The species occupies an area of less than two square kilometres (Fig. 112). The form pererosa of W. turricula is known to occur on the steep slopes at lower elevations of the islet Ilhéu de Cima. Comparison and comments. Wollastonia turricula is easily distinguishable from all the species belonging to either Hystricella or Wollastonia gen. n. by its peculiarly turreted shell and genital features. The anatomical and genital features of the species, under the name Ochtephila turricula, have previously been thoroughly described by Watson (1923), 60 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) Figures 103-111. Anatomy and genitalia of Wollastonia turricula, \lheu de Cima. 103 whole genitalia excluding gonads 104 mantle border 105 section of vagina 106 whole genitalia excluding part of OSD, AG and gonads 107 ornamentation of the inner walls of the flagellum 108 ornamentation of the inner walls of the flagellum, the penial complex, the vagina and the genital atrium 109 longitudinal section of penial papilla 110 section of penial papilla I 11 tip of penial papilla. Scale bars 1 mm. nevertheless Watson's study limited its observations mostly to the external features of the genital system. Unfortunately, except for some drawings presented by Mandahl-Barth (1950), no additional information on the genital anatomy was published until now. Wat- Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... 61 son's (1923) description, despite being thorough and accurate, sometimes gives a wrong impression of what the author observed. On p. 289 the author describes “three small finger-shaped processes” pointing out that “the shortest of the three is usually less than half the length of the others”. He considered all these three processes “homologous with the so called mucous glands found in so many of the Helicidae”. Watson considered also the vaginal appendix as a gland that is probably an apomorphic state of the stylommato- phores. He also wrongly considered the “conspicuous hemispherical swelling” on the outer side of the vagina to be “doubtless a degenerate dart-sac”. It is known that the dart sac can assume such a “swollen” appearance, as for example in the genus Cernuella, but its inner structure shows a totally different arrangement compared to that found in W turricula (see Giusti et al. 1995: 445). Watson also described the internal ornamentation of the penis as “smooth”, not detecting the main fleshy pleat and its surrounding second- ary small pleats. Nevertheless, Watsons’ (1923) description as a whole is thorough and accurate, including also interesting notes about the histology of the tissues. This kind of anatomical description was very rare at the time, where conchological features were still considered as the most important taxonomical characters in land snail systematics. Taxonomic remarks. After its description, the taxon Helix turricula var. pererosa Wol- laston, 1878 was mostly ignored by subsequent authors, except by Mandahl-Barth (1950) who treated it as a subspecies of W. turricula. Three syntypes of Helix turricula vat. pererosa Wollaston (see Figs 112-114) could be traced at the Exeter Museum (coll n°EXEMS- 1720-1909-d39-74). The preservation of the specimens clearly indicates that they were collected alive and not as subfossils. Following a careful morphological evaluation and a comparison with recent and subfossil specimens (Fig. 115), no substantial morphologi- cal differences could be detected with regard to the shell morphology, both concerning its micro- and macroscopic features (microsculpture, overall shape, keels, dimensions). Therefore, we treat it here as just a form of Wollastonia turricula and propose the name Helix turricula vat. pererosa Wollaston, 1878 as a junior synonym of Wollastonia turricula. Pettitt (1977) reports on significant changes in the shell shape of W. turricula after the construction of the lighthouse on the islet IIhé¢u de Cima after 1900. However, that obser- vation may be based on material collected prior to the construction works and therefore may have contained more specimens of the form pererosa because the top of the islet was only easily accessible after the construction of the stairs to the lighthouse and therefore col- lecting probably was carried out nearer to the coast where the pererosa form predominates. Status and conservation. According to Seddon (2011e) the species is Vulnerable (VU). In our opinion, however, it should be regarded as Critically Endangered (CR Bla, b(ii, v), 2a, b(ii, v)) because the extent of occurrence and the area of occupancy of the species is less than 1 km? and although Watson (1923) reported the species to “oc- cur in considerable numbers” on Ilheu de Cima, observations made during the 1980s (KG) showed that the species is considerably less frequent than in the past, probably as the result of a decline of habitat quality as a consequence of grazing by goats or other unknown reasons (see also Seddon (2008: 80)). Moreover, living specimens were only reported from the area around the top plateau of Ilheu de Cima, i.e., at a single loca- tion (Fig. 117). 62 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) Figures 112-116. 112-114 shells of Helix turricula var. pererosa RAM EXEMS-1720-1909-d39-74 115 Ilheu de Cima I 16a, 1 16b labels from RAM. Scale bar 1 mm. Figure 117. Distribution of Wollastonia turricula. Filled circles refer to recent and open circles to fossil records. Revision of the genus-group Hystricella R. 77 Lowe, 1855 from Porto Santo... 63 Wollastonia vermetiformis (R. T. Lowe, 1855), comb. n. Figs 118-120 List of synonyms. 1855 Helix (Hystricella) vermetiformis R. T. Lowe: 186. 1867 Helix (Octephila) vermetiformis — Paiva: 47-48. 1867 Helix (Octephila) vermetiformis var. « minor Paiva: 48. 1878 Helix (Hystricella) vermetiformis — Wollaston: 163. 1894 Geomitra vermetiformis — Pilsbry in Pilsbry 1893-1895: 242. 1931 Geomitra (Actinella) bicarinata var. vermetiformis — Nobre: 87. 1950 Discula (Hystricella) vermetiformis — Mandahl-Barth: 31, 55. 1983 Discula (Hystricella) oxytropis vermetiformis — Waldén: 267. 2002 Geomitra vermetiformis — Bank et al.: 124. 2006 Discula oxytropis — Cameron et al.: 40 [partim]. 2008 Aystricella oxytropis — Seddon: pl. 29 fig. C, map 177 [partim; in her “Cor- rigenda” Seddon (2009) mentions for pl. 29 fig. C in Seddon (2008) that the Shell of Aystricella oxytropis is normally approx. 8 mm wide, not 5 mm, and does not always have a double keel”]. 2009 Aystricella oxytropis — Seddon: 80 [partim]. 2009 Aystricella vermetiformis vermetiformis — Groh et al.: 21, fig. 29. 2017 Adystricella vermetiformis — Groh: e.T107396913A107396917. Type material. NHM 1968.588, lectotype (herewith designated), from loc. typ. ex coll. R. T. Lowe. The lectotype is depicted in the Fig. 118. Locus typicus. [vermetiformis] “Hab. fossilis in Portu S®” (= Porto Santo, fossil); [minor] rara ad Zimbral d'Aréa [= Zimbral da Areia (at the SE coast of Porto Santo, N of Barbinha-Porto dos Frades), 33°04'25"N/ 16°17'46"W]. Additional material. All from Porto Santo. Fossil: CWDM/4, Barbinha, a little distance south along the old rough road, mud fossil deposits, 33°03'56"N/16°17'49" W, 10 m, leg. W. De Mattia & J. Macor, May 2015; CKG/3, W coast, Ribeiro de Agua, 33°02'11"N/ 16°23'34"W, 50 m, leg. C. Groh, Jul. 27 1983; CKG/1, approx. 800 m E Vila Baleira, approx. 60 m N of oil tanks at bridge over Vale do Touro, from Quaternary tuffite layers, 33°03'51"N/16°19'20"W, 30 m, leg. K. Groh & J. Hemmen, Jul. 27 1983; CKG/1, Ponta da Canaveira, Quaternary aeolinites, 33°02'30"N/16°23'35"W, 50 m, leg. K. Groh & J. Hemmen, Jun. 24 1983; CKG/10, between Porto dos Frades and Ferreira Grande, approx. 300 m N sand pit, 33°04'28"N/16°17'42"W, 25 m, leg. K. Groh & J. Hemmen, Jun. 24 1983; CKG/1, Barbinha, uppermost light Quaternary aeolinite layer, 33°04'04"N/16°17'49"W, 8 m, leg. K. & C. Groh & J. & C. Hemmen, Jun. 24 1983; ANSP H 11919/21, Barbinha, Quaternary aeolinites, 33°04'04"N/16°17'49"W, 8 m, leg. K. & C. Groh & J. & C. Hemmen, Jul. 7 1983; ZMH 110125/34, coastal slopes at road from harbour to Porto Santo [Vila Baleira], 33°03'48"N/16°19'17"W, c. 10-50 m, leg. E. Clauss, Sep. 22 1992; ZMH 110139/23, coastal slopes at road from harbour to Vila Baleira, 33°03'48"N/16°19'17"W, c. 10-50 m, leg. E. Clauss, Sep. 22 1992; ZMH 110104/5, coastal slopes in the SE to E coast of Porto Santo, 33°03'46"N/16°18'21"W, 64 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) Figures | 18-120. Shells of Wollastonia vermetiformis. 118 lectotype of Helix vermetiformis R. T. Lowe, 1855, NMH 1968.588 ex coll. Lowe 119 recent specimen from Pico do Baixo 120 subfossil specimen from Porto dos Frades. Scale bar 1 mm. c. 10-50 m, leg. E. Clauss, Jun. 1 1996; ZMH 24295/1, Madeira archipelago, with- out exact locality data, ex coll. Altonaer Museum, ex coll. O. Semper, ex coll. Dohrn. Recent: CKG/1, W slope of Pico do Baixo, in a rock crack, 33°03'47"N/16°17'58"W, approx. 125 m, leg. K. Groh & J. Hemmen, Jun. 9 1983; CKG/2, Ilhéu das Cenouras, 33°06'12"N/16°17'20"W, 20 m, leg. K. Groh & J. Hemmen, Jul. 2 1983. Original description. [vermetiformis]: From Lowe 1855: T. anguste umbilicata distincte bicarinata pyramidato-conoidea solidula crassiuscula utrinque granulata; spira elevata anfractui ultimo quasi superimposita, carina inferiore suturae distinct superin- cumbente; anfr. 7—7¥% planiusculis conspicue bicarinatis, carina inf. prominente sulco infra exarata, ult. antice valde deflexo; umbil. parvo; apert. ovali-rotundata circinata, labris continuis conjunctis; perist. undique soluto relevato tenui acuto. Diam. major 84-9, min. 814-834, alt. 7-8% mill. Anfr. 7—7%; [var. minor]: From Paiva 1867: Va- riet. adest « minor, testa minore. Diagnosis. Shell large for the genus, conical, scalariform. Whorls not rounded, vertical. Surface of the shell covered by small and relatively densely set tubercles. Sec- ond whorl and body whorl bicarinated, with upper keel weaker than the lower one. Umbilicus narrow. Last whorl descending towards the aperture. Aperture oval with continuous peristome. Redescription of the shell. Shell medium to large for the genus, with 6/2 regularly increasing whorls, the protoconch with 2 whorls. The form of the shell is conical, the convex teleoconch whorls with two sharp keels. The last whorl measures 60%, the penultimate Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... 65 whorl 17% of the total shell height. The lower 74 of the body whorl are beneath the lower peripheral keel, which is set off by a relatively deep constriction above and below the keel; outline of body whorl below the lower constriction regularly convex in frontal view. The keels of the body whorl are placed in the upper twelfth and third, respectively, of its total height. ‘The suture between the whorls is simple and slightly sunken. The aperture, which is inclined to the vertical axis of the shell in an angle of 40° and is descending in its last 5% in an angle of 31° to the horizontal axis, has an elliptic form; its width is 42% of the total shell width and its height 26% of the total shell height. It has a slightly reflected, narrow lip, which is completely detached from the body whorl. The eccentric umbilicus, which measures 11% of the shell's total width, is circular in the last whorls and completely closed in earlier whorls. The protoconch is smooth, the teleoconch shows a number of oblique radial ribs, 19 in the penultimate quadrant of the body-whorl and is additionally covered by numerous small, ovate and rough tubercles. The number of tubercles in the standard-quadrate of the base is estimated to approximately 70. There are no traces of colouration in subfossil shells. In recent specimens, the colour pattern of the shell is the same as in 1. bicarinata. Figs 118-120. Variation of shell. The spire can be more or less elevated but always letting the shell appear remarkably conical and scalariform in shape. Subfossil and recent material does not show much variation. Measurements. D 6.3 + 0.4 mm (range 5.6-7.0 mm); H 6.1 + 0.6 mm (range 5.8—6.6 mm); FW 3.7 + 0.2 mm; PA 44.0 + 3.2°; DU 0.6 + 0.07 mm; NT 70 + 16; NW 6.75 + 0.2 (” = 7). Ratio D/H 1.0; ratio FW/H 0.6. Distribution. Subfossil material is known, except for one location in the west- ernmost part of the island, exclusively from the southeastern coast of Porto Santo, i.e. the mud deposits and aeolinites along Barbinha, Zimbral da Areia and Porto dos Frades. Recent, living specimens are only known from Pico do Baixo and Ilhéu das Cenouras, collected during the 1980’s. Other recent material is kept at the NUWC 80.202, acc. 55.158 (ex coll. Melvill-Tomlin, ex coll. T. V. Wollaston) and NMWC 2004.045.00055 (ex coll. Coles) (see Seddon 2008: pl. 29 fig. C), but without exact locality data (just: Porto Santo). The distribution is shown in Fig. 121. Ecology. Living specimens were collected in the 1980s on slopes under stones covered with lichens and in a rock crack. More field research is necessary for a better understanding of the ecology and actual distribution of this species. Comparison and comments. Wollastonia vermetiformis is superficially similar to H. echinoderma, W. falknerorum sp. n. and W. ripkeni sp. n. From the first two it differs in the development of the two keels on the body whorl, and from the second also in the much coarser shell sculpture, the more pronounced upper keel, and the narrower aperture. From the quite similar H. ripkeni sp. n. it differs in the lower last whorl, the oblique ovate aperture, the coarser granulation, and finer ribbing. From similar-sized W. subcarinulata and W. inexpectata sp. n. it can be separated by the coarser granulation and the presence of two keels. ‘The last known recent specimens were collected during the 1980s. Unfortunately, this material was not stored in ethanol. Therefore, an ana- tomical and molecular investigation was not possible. Recent intensive field research (on the main island Porto Santo and its outlying islets, D. Teixeira pers. comm. 2016) 66 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) Figure 121. Distribution of Wollastonia vermetiformis. Filled circles refer to recent and open circles to fossil records. failed to find any living specimen of W. vermetiformis on the Pico do Baixo. ‘The species may possibly have gone extinct in the 1980s on the main island. Status and conservation. Only three recent specimens of the species collected in the 1980s (KG) from two isolated localities (Porto Santo, W slope of Pico do Baixo, in a rock crack and Ilhéu das Cenouras; Fig. 121) are known. Despite intensive recent field research, the species has not been refound and is probably extinct on the main island of Porto Santo. Although recent data are missing for Ilhéu das Cenouras, we cannot exclude the existence of extant living populations on that small offshore islet. We regard the species as Critically Endangered (CR Bla, b(i, ii, iv), 2a, b(i, ii, iv)) (see also Groh 2017) because the extent of occurrence and the area of occupancy of the species is less than 1 km* and because the species is arguably extinct at one of the two known locations (Fig. 121). + Wollastonia ripkeni De Mattia & Groh, sp. n. http://zoobank.org/A2E9 1669-0659-447 D-9EB3-2F02174D4E87 Figs 122-125 Type material. SMF 348929, holotype, from loc. typ., leg. W. De Mattia & J. Macor, May 2015, NMWC Z.2016.013.00008, 2 PT (paratypes), NHMW 112144/1 PT, CKG, 2 PT, CWDM, 11 PT, from loc. typ., leg. W. De Mattia & J. Macor, May 2015. Revision of the genus-group Hystricella R. 7 Lowe, 1855 from Porto Santo... 67 Locus typicus. Porto Santo, excavated mud walls behind the cart speedway E of the new harbour of Porto Santo, 33°03'48"N/16°18'22"W, 30 m. Diagnosis. Shell large for the genus, solid and conical. Whorls not rounded, not vertical but somewhat slanting. Surface of the shell covered by small, and relatively densely set tubercles. Body whorl bicarinated, with upper keel weaker than the lower one. Remaining teleoconch whorls only with the upper keel visible. Umbilicus narrow. Last whorl descending toward the aperture. Aperture oval with continuous peristome. Description of the holotype. Shell large for the genus, with 674 regularly growing whorls, the protoconch with 1.85 whorls. The form of the shell is conical, the convex tel- eoconch whorls showing two rounded keels, the upper one less developed than the lower one. The last whorl measures 66%, the penultimate whorl 12% of the total shell height. The lower %4 of the body whorl are beneath the lower peripheral keel, which is set off by a constriction above and below the keel; outline of body whorl below the lower constriction regularly convex in frontal view. The two keels of the body whorl are placed in the upper Y,, and 4 of its total height. The suture between the whorls is simple, not sunken. The aperture, which is inclined to the vertical axis of the shell in an angle of 50° and is descend- ing in the last 5% of the last whorl in an angle of 35° to the horizontal axis, has an elliptic form; its width is 46%, of the total shell width, its height 31% of the total shell height. It has a slightly reflected lip, which is completely detached from the body whorl. ‘The eccen- tric umbilicus, which measures 10% of the total shell width, is in the last whorls circular and completely closed in the earlier whorls. The protoconch is smooth, the teleoconch shows a number of oblique radial ribs, 13 in the penultimate quadrant of the body-whorl and is additionally covered by numerous rounded tubercles. The number of tubercles in the standard-quadrate of the base is 89. There are no traces of colouration (Figs 122-124). Variation of the paratypes. Although the shell size is slightly variable, its shape is remarkably stable in all specimens. Measurements. D 8.0 + 0.4 mm (range 7.2-8.6 mm); H 6.2 + 0.4 mm (range 5.4—6.8 mm); FW 4.1 + 0.2 mm; PA 42 + 4.1°; DU 0.9 + 0.08 mm; NT 89 + 23; NW 6.7 + 0.2 (nm = 16). Ratio D/H 1.3; ratio FW/H 0.7. Distribution. Known only from the locus typicus. The species was hitherto found only along a 50 m stretch of mud deposits close to the cart speedway. Additional de- posits in the vicinity could not be checked for the presence of the species because of difficult accessibility of the steep slopes and associated risks of stone fall and landslides. The currently known distribution is shown in Fig. 125. Etymology. Named for the Dutch malacologist Theodor (“Theo”) J. E. Ripken from Delft, The Netherlands, to honour his valuable contributions to the malacofauna of the Macaronesian Islands. Comparison and comments. Wollastonia ripkeni sp. n. can be confused on first glance with other large-sized species such as H. echinoderma, W. falknerorum sp. n. or W. vermetiformis. From the first it is separated by the development of two distinct keels, from the second by a much finer sculpture, a narrower umbilicus and narrower aperture. From the quite similar H. vermetiformis it differs by a much more solid shell, a much higher last whorl, a larger aperture, finer granulation and coarser ribbing of the 68 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) Figures 122-124. Shells of Wollastonia ripkeni sp. n. 122 holotype, SMF 348929 123, 124 paratypes from the loc. typ. Scale bar 1 mm. Figure 125. Distribution of Wollastonia ripkeni sp. n. body whorl. From similar sized W. subcarinulata and W. inexpectata sp. n. it is separated by a coarser granulation and the presence of two keels. Taxonomic remarks. Wollastonia ripkeni sp. n. is included in the genus Wollas- tonia because it is similar to W. oxytropis in size and surface sculpture, considerably Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... 69 differing from Hystricella s. str. where the tubercles are bigger and less densely set. We are well aware that, due to the lack of anatomical and molecular data, the generic placement of this subfossil species is exclusively based upon shell features, with all the taxonomic limits that this might implicate. Status and conservation. Extinct before the islands’ scientific exploration in the 19" century; possibly already before human settlement. + Wollastonia falknerorum Groh, Neiber & De Mattia, sp. n. http://zoobank.org/644C84 1 3-OAF0-41E8-B286-2E7C413FA043 Figs 126-129 Type material. All from Porto Santo, holotype, SMF 348925, from loc. typ., leg. W. De Mattia & J. Macor, May 24 2015; SMF 348926/2 PT, NMW.Z.2016.013.00007/2 PT, NHMW 112142/1 PT, CKG/2 PT, CWDM/22 PT, from loc. typ., leg. W. De Mattia & J. Macor, May 24 2015; NMG No. 78-14764/2 PT [as H. vermetiformis, det. H. Waldén], Prainha & Serra de Fora, 33°03'57"N/16°17'49"W, 20 m, leg. A. De Noronha, before 1978; NMG No. 86-17153/4 PT [as H. vermetiformis, det. H. Waldén], Penedo, 33°03'38"N/16°19'50"W, 10 m, leg. J. J. De Sousa, before 1986; CWDM/9 PT, Barbinha, slightly S towards the tunnel, Quaternary mixed gravel slope deposit, 33°3'56"N/ 16°17'49"W, 20 m, leg. W. De Mattia & J. Macor, May 24 2015; CKG/9 PT; Vale do Touro, 800 m E of Vila Baleira, 60 m NW of oil tanks, Quater- nary slope deposit from the 3“ of 11 layers of tuffites with approx. 20% of stones, 33°03'49"N/16°19'21"W, 12-14 m, Jun. 18 1983, leg. K. & C. Groh; CJG/3 PT, Vale do Touro, 800 m E of Vila Baleira, 60 m NW of oiltanks, Quaternary slope deposit from the 3" of 11 layers of tuffites with approx. 20% of stones, 33°03'50"N/16°19'21"W, 12-14 m, leg. J. Gerber, K. Groh & J. Hemmen, Aug. 16 1985; CKG/3 PT + 2 juv. (no PT), Pico dos Magarico, opposite to Porto de Abrigo, leg. K.-H. Beckmann, Jan. 17 1998; CFW 12179/1 PT, Ponta da Galé, W end of tunnel, 300 m towards port, 33°03'36"N/16°17'58" W, 25 m, leg. E Walther, Apr. 1 2017; CFW 12178/10 PT + 4 juv. (no PT) + fragm. (no PT), Ponta da Galé, W end of tunnel, 100 m towards port, 33°03'38"N/16°17'51" W, 35 m, leg. EF Walther & E. M. Gryl, Apr. 1 2017; CFW 12177/13 PT + 6 juv. (no PT) + 1 fragm. (no PT), CKG/3 PT, Ponta da Galé, E end of tunnel, lower level [of slope deposits], coarse, red coloured gravel with large stones, 33°03'47"N/16°17'45"W, 30 m, leg. F Walther, Apr. 1 2017; CFW 12180/34 PT + 11 juv. (no PT) + fragms (no PT), CKG/5 PT, E of Vila Baleira, end of Vale do Touro, (sub-) fossil [slope-]deposits, coarse, black gravel, 33°03'48"N/16°19'15"W, 20 m, leg. F Walther, Apr. 5 2017; CFW 12181/21 PT + 4 juv. (no PT) + fragms (no PT), E of Vila Baleira, S-slope of the hill above Vale do Touro, W of the oil tanks, [(sub-)fossil slope-deposits of] red gravel, 33°03'47"N/16°19'26"W, 25 m, leg. E Walther, Apr. 5 2017; ANSP H 11854/9 PT [partim, together with 7 H. subcarinulata and 2 Discula cheiranticola], Barbinha, 33°04'04"N/16°17'49"W, 8 m, leg. J. & C. Hemmen & K. & C. Groh, 1983. 70 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) Locus typicus. Porto Santo, Pico do Baixo, E entrance of the tunnel, Quaternary mud deposit, 33°3'44"N/16°17'45"W, 20 m. Diagnosis. Wollastonia species with two keels on the body whorl, the upper keel only very weakly developed; suture overlapping; granulation very fine; ribbing of the body whorl rather coarse. Description of the holotype. Shell large for the genus, with 6.5 regularly increas- ing whorls, the protoconch with 2.1 whorls. The form of the shell is conical, the convex teleoconch whorls show an upper, rounded angulation and a lower distinctly developed keel. The last whorl measures 60%, the penultimate whorl 10%, of the total shell height. The lower 34 of the body whorl are beneath the keel, which is set off by a constriction above and below; shell below the lower constriction regularly convex in frontal view. The angulation of the body whorl is placed in the upper seventh of its total height and the keel in the upper fourth of its total height. The suture between the whorls is simple, but slightly overlapping the previous whorl. The aperture, which is inclined to the vertical axis of the shell in an angle of 54° and is descending in the last 5% of the last whorl in an angle of 42° to the horizontal axis, has an elliptic form, which is slightly narrowed towards the columella. Its width is 45%, of the total shell width and its height 31% of the total shell height. It has a slightly reflected lip, which is completely detached from the body whorl. The eccentric umbilicus, which measures 14% of the shell's total width, is in the last whorls circular and completely closed in earlier whorls. The protoconch is smooth, the teleoconch shows a number of oblique radial ribs, 9 in the penultimate quadrant of the body whorl and is additionally covered by numerous small, rounded tubercles. The number of tubercles in the standard-quadrate of the base is 135. There are traces of colouration in form of large blotches along the last two whorls (see Fig. 126). Variation of the paratypes. The shell shape is quite stable in all examined speci- mens (Figs 127-128). Measurements. D 8.4 + 0.1 mm (range 8.2-8.5 mm); H 7.0 + 0.3 mm (range 6.8—7.4 mm); FW 4.3 + 0.2 mm; PA 42 + 5.3°; DU 0.9 + 0.09 mm; NT 68 + 18; NW 6.5 + 0.2 (nm = 20). Ratio D/H 1.2; ratio FW/H 0.6. Distribution. Wollastonia falknerorum sp. n. is found in the Quaternary mud and slope deposits along the southeastern coast of Porto Santo. Its range extends from the hills east of Vila Baleira to Barbinha, S of Porto dos Frades. The distribution is shown ih Bio eh 2.9" Etymology. Named for the German malacologists Margrit & Gerhard Falkner, Horlkofen, to honour their highly valuable contributions to European malacology. Comparison and comments. Wollastonia falknerorum sp. n. can be confused on first glance with other comparatively large-sized geomitrid species like, H. echinoder- ma, W. ripkeni sp. n. and W. vermetiformis. From the first it is distinguishable by the presence of two distinct keels and from all three by a much finer sculpture. From the most similar W. vermetiformis and W. ripkeni sp. n. it is differentiated by the weakly developed upper keel, an overlapping suture, a much finer granulation and coarser rib- bing of the body whorl. From similar-sized W. subcarinulata and W. inexpectata sp. n. it is separated by the two keels and the differently developed suture. Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... Fl Figures | 26-128. Shells of Wollastonia falknerorum sp. n. 126 holotype, SMF 348925127 127, 128 para- types from the loc. typ. Scale bar 1 mm. Figure 129. Distribution of Wollastonia falknerorum sp. n. Taxonomic remarks. Wollastonia falknerorum sp. n., as with other subfossil Wo/l- lastonia species, is included in this genus because it is similar to W. oxytropis in shell shape, size and surface sculpture, which differ considerably from Hystricella species. We are well-aware that, due to the lack of anatomical and molecular data, the generic position of this subfossil species is exclusively based upon shell features, with all the taxonomic limits that this might implicate. ip Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) Status and conservation. Extinct before the islands’ scientific exploration in the 19" century, possibly already before human settlement. Wollastonia leacockiana (Wollaston, 1878), comb. n. Figs 130-142 List of synonyms. 1878 Helix (Hystricella) leacockiana Wollaston: 165-167. 1894 Geomitra leacockiana — Pilsbry in Pilsbry 1893-1895: 34. 1931 Geomitra (Actinella) bicarinata var. leacockiana — Nobre: 87. 1950 Discula (Aystricella) leacockiana — Mandahl-Barth: 31, 55. 1983 Discula (Hystricella) leacockiana — Waldén: 20: 267. 2002 Geomitra leacockiana — Bank et al.: 124. 2008 Aystricella leacockiana — Seddon: 79-80, pl. 29 fig. A, map 179. 2011 Aystricella leacockiana — Seddon: e.16720A12799605. Type material. NHM 1875.12.31.137-d, lectotype (herewith designated), from loc. typ., ex coll. T. V. Wollaston; NHM 1875.12.31.137-a to -c and -e, 4 paralecto- types, from loc. typ., ex coll. T. V. Wollaston. Further material examined. All from Porto Santo, CWDM/14, CMN/3, Pico de Ana Ferreira, northern slopes near the ‘lava columns’, under stones scattered in open grassy fields, 33°02'57"N/16°22'04"W, 125 m, leg. W. De Mattia & J. Macor, May 2014; CKG/32, CMN/9, Pico de Ana Ferreira, top of the hill, under stones scat- tered in open grassy fields, 33°02'39"N/16°22'11"W, 230 m, leg. K. & C. Groh & J. & C. Hemmen, Jul. 6 1983; CKG/20, CMN/10, E slope of Pico de Ana Ferreira, 33°02'37"N/ 16°22'05"W, 100-200 m, leg. K. & C. Groh, Oct. 26 1980; CKG/1, Ilhéu de Ferro, east side of the island plateau, under stones scattered in open grassy fields, 33°02'19"N/16°24'26"W, 75 m, leg. K. & C. Groh & J. & C. Hemmen, Jul. 2 1983; CFW 11151/<10, N slope of Pico de Ana Ferreira, near the basalt columns, 33°02'57"N/16°22'03"W, 125 m as.L., leg. FR Walther & E. M. Gryl, Mar. 30 2017; CFW 11152/<10 spms, Pico de Ana Ferreira, summit, 33°02'38"N/16°22'13"W, 230 m, leg. F Walther, Apr. 4 2017; ANSP H 11838/c. 40, NW slope of the Pico de Ana Ferreira, 33°02'48"N/ 16°22'11"W, 190 m, leg. K. & C. Groh & J. & C. Hemmen, Jul. 6 1983; ANSP H 11770/ c. 30 [sub H. bicarinata], E slope of Pico de Ana Ferreira, 33°02'37"N/ 16°22'05"W, 100-200 m, leg. J. & C. Hemmen, Jan. 5 1981. Locus typicus. Portum Sanctum; in monte ‘Pico d’Anna Ferreira’ dicto sat copiose reperta. Necnon in statu semifossili (cum exemplaribus recentibus vix omnino congru- ens) parcissime occurrit. Original description. From Wollaston, 1878: T. trochiformis, subtus planata per- forata, undique granulis obtusis densissime obsita, pallide brunneo-subflavescens sed fasciis (presertim subtus) nebulisque irregiularibus (presertim supra) rufo-brunneis hinc inde suffuse marmorata; spira sat elevata; anfractibus convexis, bicarinatis, ultimi (subtectiformis) carina exteriore acutissima valde exstanti, interior obtusa rotundata Revision of the genus-group Hystricella R. 77 Lowe, 1855 from Porto Santo... Ye) Figures 130-132. Shells of Wollastonia leacockiana. 130 lectotype of Helix leacockiana Wollaston, 1878, NMH 1875.12.31.137d, ex coll. Wollaston 131 Pico de Ana Ferreira 132 Ilheu de Ferro. Scale bar 1 mm. recedente rarius obsoleta; umbilico punctiformi; apertura subovali-rotundata, labris continuis conjunctis, peristomate simplici expanso subrecurvo tenui relevato. — Long. axis 174 lin.; diam. 2%. Redescription of the shell. The shell is dextral, hairless and it is usually conical and scalariform. The protoconch is from whitish to brown with 1.6 to 1.9 whorls. It is almost smooth along the first half whorl and shows fine radial striae and extremely small, scattered tubercles along its remaining portion. The teleoconch has from 3.9 to 4.4 rapidly increasing whorls. It is variable in colour, from very light brown to brown. The background colour is mottled with brownish to dark-reddish areas, irregularly ar- ranged along the whorls. No band pattern is visible along the upper whorls. On the lower part of the last whorl two more or less indistinct, dark bands are visible. The peripheral band is usually the thinnest and often very indistinct. The spire is usually pyramidal and slightly scalariform. Along the last and partially along the penultimate whorl one evident keel is present. This keel extends along the lower part of the whorls and is usually lighter in colour, sometimes whitish. The external upper surface has very fine but clearly visible, irregularly spaced, growth lines. Irregularly disposed tubercles are present all over the teleoconch. The dimensions of the tubercles tend to slightly increase from the first towards the last whorl, but keeping approximately the same density. The tubercles are somewhat denser along the keel, letting the keel appear like a rough chord. The tubercles usually also concentrate somewhat along the growth lines. 74 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) On the lower part, the tubercles are much denser along the periphery and gradually become scarcer towards the umbilicus. The last whorl is relatively large and descending near the aperture. The umbilicus is open but very narrow, concentric and measures approximately 10% of the maximum shell diameter. The aperture is elliptical with a quite strong thickening along the inner side of the last whorl. The peristome is con- tinuous, slightly reflected with the columellar margin being somewhat thicker (see Figs 130-132). Measurements. D 5.3 + 0.2 mm (range 5.0-5.6 mm); H 4.0 + 0.3 mm (range 3.7-4.8 mm); FW 2.6 + 0.2 mm (range 2.4—2.6); PA 58.8 + 2.1° (range 55.0—60.0°); DU 0.5 + 0.1 mm (range 0.4-0.6); NT 49 + 5 (range 44-56); NW 5.4 + 0.1 (range 5.3-5.5) (2 = 40). Ratio D/H 1.3; ratio FW/H 0.6. Body. ‘The head and neck are usually light grey, as is the posterior upper section of the foot. The foot is white and the sole is longitudinally divided into three areas. The central area is smooth, whereas the two lateral areas are equipped with bands of mus- cles roughly arranged in a chevron pattern. The mantle border is light grey, with five more or less developed lobes. The walls of the pallial cavity are colourless, without any stripes or spots. A strong pulmonary vein is visible. The jaw is odonthognatous and its shape is arched. There are 8 to 10 smooth transverse ridges. The right ommatophoral retractor is independent from both penis and vagina. Genital anatomy. The albumen gland is long and it is connected to an approxi- mately equally long sperm-oviduct. The thin vas deferens is approximately as long as the sperm-oviduct. The free oviduct is shorter than the vagina. The duct of the bursa copulatrix is usually thin, approximately as long as the penis and uniform in diameter. The bursa copulatrix is roundish. The transition area between the duct and the bursa itself is very sharply delimited, with the duct abruptly widening and turning into the bursa. The spermatophore is unknown. One tuft of a digitiform glands arises from the proximal part of the vagina. There are two or three glands that are equally long and never branched. A short and thin vaginal appendix arises from the vagina’s wall, just distal of the glandular tuft. The inner surface of the vagina is smooth. The atrium is usually long (% of the vagina) and relatively thin. Its internal walls are also smooth. ‘The penial flagellum is short, remarkably cylindrical and with a blunt apex. It is usually as long as the epiphallus and its internal walls are complete- ly smooth. The internal walls of the epiphallus are completely smooth as well. The retractor muscle is moderately long, strong and is of variable length. The penis lacks any muscular or glandular sheath. It is thick-walled and approximately three to four times longer than the flagellum. It is usually cylindrical, sometimes slightly swollen and partially folded up. The inner walls of the penis are smooth. The penial papilla is small and bulky. It has smooth external walls with the opening emerging apically but somewhat curving inwards. The channel of the penial papilla is thin and narrow and its cross section is ‘half-moon’-shaped. The inner lumen of the penial papilla is filled with a spongy and sturdy tissue, which directly connects with the walls of the epiphallus. As all the previous species, the longitudinal section of the penial papilla shows that its walls are the continuation of the penial walls that abruptly bend in- ward (see Figs 133-141). Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... fis) Figures 133-141. Genitalia and anatomy of Wollastonia leacockiana. Pico de Ana Ferreira, N slopes: 133 section of penial papilla 134 section of vagina 135 whole genitalia excluding gonads 136 tip of the penial papilla 137 longitudinal section of penial papilla 138 ornamentation of the inner walls of the distal penis, the distal vagina and the genital atrium 139 whole genitalia excluding gonads 140 jaw 141 mantle border. Scale bars 1 mm. Ecology. Wollastonia leacockiana is commonly found under volcanic rocks and boulders scattered on grassland in open fields that are more or less sloping. The speci- mens aestivate on the lower surfaces of the rocks, frequently forming clusters of indi- viduals attached one to another. 76 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) Figure 142. Distribution of Wollastonia leacockiana. Distribution. Wol/astonia leacockiana is restricted to Pico de Ana Ferreira and Ca- beco da Ponta at the western end of Porto Santo, with records also from Ilhéu de Ferro (CGk), although surveys are required to confirm if the species is still extant on that is- let. Seddon (2008: 79, 182) reported the species from a spot at the eastern end of Porto Santo (Ponta do Passo area?). Despite intensive recent field research (WDM 2012, 2014, 2015) the species has not been confirmed in this area. Seddon’s (2008) record could represent a misidentification of a different species (cf. W. klausgrohi sp. n.). The currently known distribution is shown in Fig. 142. Taxonomic remarks. As reported by Seddon (2008: 79), Wollaston (1878: 161) separated W. leacockiana from H. bicarinata on the basis of its shell sculpture. This study confirmed Wollaston’s (1878) opinion, as do our molecular analyses and the investigation of the anatomy of the species. Status and conservation. The species is currently not significantly impacted by the construction of touristic/residential facilities in its distributional range; only in the Cabeco da Ponta area some new buildings were recently constructed. Recent surveys (WDM, 2012, 2014, 2015) showed that the species is currently widely distributed and common within its range. According to Seddon (201 1c) the species is Vulnerable (VU D2), an assessment we regard as appropriate here because of the small range of the spe- cies and the potential threat of future construction works in the area. Further research is required to confirm the distribution of the species. Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... Ti + Wollastonia beckmanni De Mattia & Groh, sp. n. http://zoobank.org/EOAOF760-57EG6-4629-B09B-5FCCA78CCB 13 Figs 143-146 Type material. All from Porto Santo, SMF 348927, holotype, from loc. typ., leg. K. & C. Groh, June 6 1983; SMF 348928/5 PT, MMF 24956/1 PT, MMF 46281/1 PT, MME 46282/1 PT, NHMW 112143/1 PT, CKG/62 PT, NMWZ 2016.013.00009/5 PT, CJG/5 PT, from loc. typ., leg. K. & C. Groh, June 6 1983; CWDM/12 PT, E of Vila Baleira, S slope of the hill above Vale do Touro, 50 m W of the oil tanks, exca- vated Quaternary mixed gravel, 33°03'47"N/16°19'26"W, 24 m, leg. W. De Mattia & J. Macor, May 24 2015; CFW 12176/5 PT, E of Vila Baleira, S slope of the hill above Vale do Touro, W of the oil tanks, [(sub-)fossil slope-deposits of] red gravel, 33°37'52"N/16°19'26"W, 25 m, leg. F Walther, Apr. 5 2017; ANSP H 11843/2 PT [sub H. leacockiana], Vale do Touro, 33°03'46"N/16°19'29"W, 15 ma, leg. J. Gerber, K. Groh & J. Hemmen, Aug. 16 1985. Locus typicus. Porto Santo, south coast, ditch near the harbour built in 1985 at Vila Baleira, lens of quaternary calcareous aeolinite within a marine sandy beach ter- race (nowadays beyond the quay of the harbour), 33°03'50"N/16°18'56"W, 0-1.5 m. Diagnosis. Small Wollastonia species with a very depressed body whorl that is fine- ly granulated; umbilicus very wide in relation to maximum diameter, rather eccentric. Description of the shell of the holotype. Shell small for the genus, with 5.15 rap- idly increasing whorls, the protoconch measures 1.5 whorls. The form of the shell is flat conical, the convex teleoconch whorls show a strongly keeled periphery. The last whorl measures 65%, the penultimate whorl 14% of the total shell height. The lower half of the body whorl is slightly concave beneath the keel in frontal view near the periphery and oth- erwise regularly convex. The suture between the whorls is simple and slightly impressed. The aperture, which is inclined to the vertical axis of the shell in an angle of 58° and de- scending in the last 5% of the last whorl in an angle of 43° to the horizontal axis, has an elongate-ovate form, its width measures 42% of the total shell width, its height 27% of the total shell height. It has a distinctly reflected lip, which is completely detached from the body whorl. The very eccentric umbilicus, which measures 17% of the total shell width, is in the upper whorls narrowly perspective. The protoconch is smooth, the teleoconch shows a low number of oblique radial ribs, nine in the penultimate quadrant of the body whorl and is additionally covered by many fine tubercles. The number of tubercles in the standard quadrate of the base is 102; the tubercles become coarser towards the centre of the base and finer towards the periphery. There are no traces of colouration (see Fig. 143). Variation in paratypes. ‘The size varies slightly, and the density of the tubercles in the standard quadrate of the base may vary by + 10%. Exceptionally, the um- bilicus may be narrower and the angles of the aperture to the vertical axis and the descending of the last part of the body whorl to the vertical axis also slightly vary (see Measurements section below). Nevertheless, the overall shape and sculpture of all paratypes is very close to that of the holotype. In a few shells traces of a brownish 78 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) Figures 143-145. Shells of Wollastonia beckmanni sp. n. 143 holotype, SMF 348927 144 paratype from the loc. typ. 145 Vale do Touro. Scale bar 1 mm. colouration are perceptible, however there are no traces of a banding pattern (see Figs 144-145). Measurements. D 5.3 + 0.2 mm (range 5.0—5.6 mm); H 3.3 + 0.2 mm (range 3.2— 3.5 mm); FW 2.2 + 0.1 mm (range 2.1—-2.3 mm); PA 72.0 + 3.0° (range 66.0—76.0°); DU 0.4 + 0.05 mm (range 0.3-0.4 mm); NW 6.2 + 0.2 (range 6.0-6.4) (7 = 20). Ratio D/H 1.6; ratio FW/H 0.7. Distribution. Wollastonia beckmanni sp. n. is known only from the southeastern coast of Porto Santo, from the hill immediately east of Vila Baleira (mud deposits at Vale do Touro) to the aeolinite deposits behind the new harbour. See map in Fig. 146. Etymology. Named after the late German entrepreneur and self-taught malacologist Dr. Karl-Heinz Beckmann (1948-2007) to honour his valuable contributions to the malacofauna of different islands and archipelagos in the Mediterranean and the Atlantic. Comparison and comments. Because of its small size the new species can only be confused with small H. aucta Wollaston, 1878, which have, however, a higher and more distinctly stepped spire, have two well-developed keels and much coarser and more sparsely set tubercles on the shell surface. Hystricella microcarinata sp. n. has a much higher, rounded conical form, two keels on the body whorl, coarser and less densely set tubercles, a much narrower umbilicus, and a different shape and differently positioned angles of the aperture. Taxonomic remarks. As with other subfossil Wollastonia species, W. beckmanni sp. n. is included in this genus because it considerably differs from Hystricella and is similar to W. oxytropis in surface sculpture and overall shell shape. Status and conservation. Extinct before the islands’ scientific exploration in the 19" century, possibly already before human settlement. Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... 79 Figure | 46. Distribution of Wollastonia beckmanni sp. n. Wollastonia jessicae jessicae De Mattia, Neiber & Groh, sp. n. http://zoobank.org/ADE24AD8-A30A-4171-992B-C7671B10902A Figs 147-149 Type material. NUWC Z.2016.013.00001, holotype, from loc. typ., leg. W. De Mat- tia & J. Macor, May 25 2015, NUWC Z.2016.013.00002/4 PT, SMF 348934/4 PT, NHMW 112140/3 PT, CKG/3 PT, CMN/7 PT, CWDM/23 PT, from loc. typ., leg. W. De Mattia & J. Macor, May 24 2015; FW 11154/10 PT, CMN/10 PT, ZMH 131207, 10 PT, hill E of Ribeira Santo Antonio and W of Vale do Touro, upper edge of the S slope, 33°03'47"N/16°19'41"W, 40 m, leg. E Walther, Apr. 5 2017; CKG/5 PT, CJG/2 PT, Vale do Touro, under stones, 33°03'51"N/16°19'26"W, 45 m, leg. J. Gerber, K. Groh & J. Hemmen, Aug. 16 1985. Fossil: CKG/2 PT, Vale do Touro, Quaternary slope deposits, 33°03'47"N/16°19'26"W, 25 m, leg. J. Gerber, K. Groh & J. Hemmen, Aug. 16 1985. GNM 72-13.386/1 PT, Portela, 500 m SW, in the road cutting, Quaternary muds, 33°03'54"N/ 16°19'12"W, 95 m, leg. H. W. Waldén, Feb. 13 1972; ANSP H 11920/7 PT [with 8 spms of H. aucta], Vale do Touro, Quaternary slope deposits, 33°03'47"N/16°19'26"W, 25 m, leg. J. Gerber, K. Groh & J. Hem- men, Aug. 16 1985; ANSP H 11773/2 PT [with 3 Caseolus compactus], Vale do Touro, under stones, 33°03'51"N/16°19'26"W, 45 m, leg. J. Gerber, K. Groh & J. Hemmen, Aug. 16 1985. 80 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) Figures 147-149. Shells of Wollastonia jessicae jessicae sp. n. 147 holotype, NMWC Z.2016.013.00001 148 paratype from the loc. typ. 149 subfossil specimen from Vale do Touro. Scale bars 1 mm. Locus typicus. Upper edge of the S slope of hill E of Ribeira Santo Antonio and W of Vale do Touro, under stones in grassland, 33°03'47"N/16°19'41"W, 40 m. Diagnosis. Small Wollastonia species, with a conical shell and two keels on the body whorl, the lower more distinct than the upper; granulation relatively fine and scattered; internal walls of penis with one to three well-developed, irregular pleats. Etymology. Named after the wife of the first author, Mrs. Jessica Macor from Trieste, Italy, as a token of gratitude for her assistance and companionship during the collecting activities. Description of shell. The shell is dextral and hairless. Its shape is scalariform, with deep sutures and rather flattened whorls. The protoconch is from yellowish to dark brown with 1.5 to 2.5 whorls. It is almost smooth along the first whorl and shows fine radial striae in its remaining whorls. The teleoconch has from 3.3 to 3.8 rapidly increasing whorls. It is usually dark brown with brick red and/or dark violet shades in colour, but also yellowish specimens with light reddish shades are found. The darker areas of the shell are mottled with more or less light brown to whitish areas, usually placed longitudinally and slightly slanting. The body whorl has two well-developed keels. The lower one is stronger and more distinct than the upper one. Both keels are usually whitish and contrasting from the rest of the body whorl. No banding pattern is visible along the upper whorls. On the lower part of the last whorl one principal, but rather narrow, dark (reddish to dark brown) band is usually present. A second light Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... 81 and indistinct band is sometimes present just below the keel. The area around the um- bilicus is usually the lightest in colour. The external surface has strong, clearly visible, irregularly spaced, growth lines. Tubercles are present all over the teleoconch. They are usually large and rather widely scattered, whitish in colour and arranged somewhat obliquely, following the course of the growth lines. The larger tubercles are somewhat denser along the keel(s) of the last whorls, letting the keel(s) appear like a rough chord. The last whorl is usually large, with a contribution of 60% of the total shell height and descending near the aperture. The umbilicus is open but very narrow, either concentric or eccentric, and it measures approximately 10% of the maximum shell diameter. The aperture is elliptical with a faint thickening along the columellar portion of the stoma. Sometimes this thickening can also extend as far the parietal side of the aperture. The peristome is continuous and reflected (see Figs 147-149). Measurements. D 4.8 + 0.3 mm (range 4.4-5.2 mm); H 3.3 + 0.3 mm (range 3.0-3.9 mm); FW 2.3 + 0.1 mm (range 2.2—2.4 mm); PA 58.2 + 7.4° (range 50.0— 68.0°); DU 0.7 + 0.05 mm (range 0.6-0.8 mm); NT 23 + 12 (range 9-37); NW 5.6 + 0.4 (range 5.0-5.9) (7 = 40). Ratio D/H 1.5; ratio FW/H 0.7. Body. The head and the neck are usually grey. The sides and the posterior upper section of the foot are whitish. The pigmented ommatophoral retractor muscles are visible through the skin of the back of the cephalic area. ‘The foot is white and the sole is longitudinally divided into three areas. The central area is smooth, whereas the two lateral areas are equipped with bands of muscles that are roughly arranged in a chevron pattern. The mantle border is grey to dark grey, with five more or less developed lobes. The ratio of the lateral to the dorsal lobes varies from specimen to specimen, also in the same population. In some specimens, one of these lobes (either lateral or dorsal) may be totally missing. The walls of the pallial cavity are colourless, without any stripes or spots. A strong pulmonary vein is visible. The jaw is odonthognatous and is very variable in shape, ranging from almost straight to markedly arched. There are many smooth transverse ridges, ranging from 18 to 22 in number. The right ommatophoral retractor is independent from both penis and vagina. Genital anatomy. ‘The general arrangement of the genitalia is semi-diaulic mono- trematic. A convoluted to almost straight hermaphrodite duct arises from a multi- lobated gonad. The albumen gland is long and thin and is connected to an approxi- mately twice as long sperm-oviduct that consists of a prostatic and a uterine portion. The prostatic part extends into a thin vas deferens, which is approximately as long as the sperm-oviduct and terminates in the penial complex. The distal portion of the uterine part extends into the free oviduct, turning into a vagina at the level of the duct of the bursa copulatrix. The free oviduct can be as long as, or slightly shorter than the vagina. The duct of the bursa copulatrix is usually thin, approximately as long as the penis and uniform in diameter. It ends into a variable, oval to roundish, bursa copulatrix. The transition area between the duct and the bursa itself is usually sharply delimited, with the duct abruptly widening and turning into the bursa. The spermato- phore is unknown. One tuft of digitiform glands arises from the proximal part of the vagina. There are usually two glands that are approximately equally long and very rarely 82 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) branched. A short and thin vaginal appendix arises from the vagina’s wall, just distal of the glandular tuft. Very smooth, rather wide and little elevated, irregularly spaced pleats run longitudinally along the inner surface of the vagina, reaching into the genital atrium as far as the genital orifice. The atrium is usually long and thin. Its internal walls have large and soft pleats running longitudinally as far the genital orifice. The penial complex consists of a flagellum, an epiphallus (which extends from the insertion of the vas deferens to the penial retractor muscle) and a penis, which inserts into the atrium. The penial flagellum is short, remarkably cylindrical and with a blunt apex. It is usu- ally as long as the epiphallus. Its internal walls are completely smooth. ‘The epiphallus is usually short and its internal walls present 2 to 4 longitudinal, smooth pleats that are only slightly elevated. The retractor muscle is not very large, but strong, usually as long as the flagellum + epiphallus together. The penis lacks any muscular or glandular sheath. It is thick-walled and approximately three times longer than the flagellum. It is usually slightly swollen in its distal part near the genital atrium. The inner walls of the penis usually have one to three irregular, spaced pleats, which run longitudinally and reach the genital atrium. These pleats can be connected by small “bridge-like” pleats. In some specimens, the section where the penial papilla is located is detectable from the outside by virtue of a fine circular swelling corresponding to the origin of the papilla itself. The penial papilla is usually very variable in dimensions (measuring from 10% to 50% of the total penial length) and conical in shape. It has smooth external walls, with the opening emerging apically. The channel of the penial papilla is thin and narrow. The inner lumen of the penial papilla is occupied by a spongy and sturdy tis- sue, which directly connects with the walls of the epiphallus. The longitudinal section of the penial papilla shows that its walls are the continuation of the penial walls that abruptly bend inward (see Figs 150-157). Distribution. Wollastonia jessicae jessicae sp. n. is found only along the southern slope of a hill east of Ribeira Santo Antonio and west of Vale do Touro, just east of the town of Vila Baleira, along the road leading to the new harbour. The distribution is shown in Fig. 158. Ecology. Wollastonia jessicae jessicae sp. n. is commonly found under volcanic rocks scattered on grassland in open fields that are more or less sloping. The specimens aes- tivate on the lower surfaces of the rocks, frequently forming clusters of individuals attached one to another. Comparison and comments. At first glance W. jessicae jessicae sp. n. can be con- fused with H. bicarinata because of the overall similarity of their shell’s shape (i.e. granulated surface with the last whorl with two keels). This is probably the reason why the new species has been overlooked until now. Nevertheless, a closer look reveals dif- ferences such as the much coarser granulation on the surface of H. bicarinata. W. jessicae jessicae sp. n. (as for W. jessicae monticola sp. n. described in the following section) always shows a finer granulation, having smaller and more scattered tubercles. With regard to genital anatomy, W. jessicae jessicae sp. n. has internal walls of the penis with one to three irregular and well-spaced strong pleats, which run longitudinally and reach the genital atrium, whereas 1. bicarinata has at least some large and smooth Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... 83 Figures 150-157. Genitalia and anatomy of Wollastonia jessicae jessicae sp. n. S slope of hill E of Ribeira Santo Antonio and W of Vale do Touro: 150 whole genitalia excluding gonads 151 section of proximal penis at the intersection with vas deferens 152 whole genitalia excluding gonads 153 ornamentation of the inner walls of the distal penis, the distal vagina and the genital atrium 154 jaw 155 mantle border 156 whole genitalia excluding part of OSD, AG and gonads 157 ornamentation of the inner walls of the distal penis, the distal vagina and the genital atrium. Scale bars 1 mm. folds. Our phylogenetic analysis clearly show that the two species are not closely relat- ed, i.e. they were placed in two well-separated clades. This further corroborates, aside from morphology, its status as a distinct species. 84 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) Figure 158. Distribution of Wollastonia jessicae jessicae sp. n. Filled circles refer to recent and open circles to fossil records. Status and conservation. The subspecies has a very limited distribution area of less than | km? (Fig. 158) close to a village and population size is probably rather low. Habi- tat quality is inferred to be declining and potential and ongoing threats to the subspecies include, in our opinion, urbanization, tourism, goat grazing and quarrying. Therefore, the species is considered here to be Critically Endangered (CR Bla, b(iii), 2a, b(iii)). Wollastonia jessicae monticola De Mattia, Neiber & Groh, ssp. n. http://zoobank.org/98F7562E-7017-4CAA-8 DDA-91EA11C3DB54 Figs 159-167 Type material. NUWC Z.2016.013.00010, holotype, from loc. typ., leg. W. De Mat- tia & J. Macor, May 21 2015, NMWC Z.2016.013.00011/4 PT, SMF 348931/4 PT, NHMW 1121341/3 PT, CKG/3 PT, CMN/3 PT, CWDM/6 PT, from loc. typ., leg. W. De Mattia & J. Macor, May 21 2015; FW 11155/5 PT, CMN/5 PT, ZMH 131208/4 PT, 200 m SW of the Zimbreiro near the road turn serpentine, 33°04'16"N/16°18'53"W, 100 m, leg. E Walther, Mar. 31 2017; FW 11157/>10 PT, ridge between Zimbreiro and the quarry, 33°04'13"N/16°18'49"W, 110 m, leg. E Walther, Apr. 3 2017. Locus typicus. Porto Santo, Zimbreiro, 200 m SW of the village, near the road turn serpentine, 33°04'16"N/16°18'53"W, 85 m. Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... 85 Figures 159-161. Shells of Wollastonia jessicae monticolassp.n 159 holotype, NMWC Z.2016.013.00010 160, 161 paratypes from the loc. typ. Scale bar 1 mm. Diagnosis. Subspecies of Wollastonia jessicae with two keels on the body whorl, the lower distinct, the upper one very weakly developed to lacking; granulation relatively fine and scattered; internal walls of penis smooth, without pleats or prominent folds. Etymology. Named for the restriction of its habitat to the mountainous region (Lat. mons, montis = mountain(s) and -cola = inhabitor) of the island of Porto Santo. Shell. Similar to the nominate subspecies, except for the ornamentation of the body whorl. In W. jessicae monticola ssp. n. the upper keel is missing and the lower keel is somewhat less strongly developed and less evident compared to W. jessicae jessicae sp. n. (see Figs 159-161). Measurements. D 4.6 + 0.2 mm (range 4.2-4.9 mm); H 3.1 + 0.1 mm (range 2.8-3.6 mm); FW 2.0 + 0.1 mm (range 1.9—2.3 mm); PA 58,6 + 6.1° (range 51.3— 65.7°); DU 0.7 + 0.04 mm (range 0.6—0.8 mm); NT 25 + 14 (range 8-39); NW 5.5 + 0.3 (range 4.9-5.9) (7 = 30). Ratio D/H 1.5; ratio FW/H 0.6. Body. The overall body colouration (i.e. head, neck and sides) of W. jessicae monticola ssp. n. tends to be somewhat darker than in the nominate subspecies. Genital anatomy. The distal genitalia of W. jessicae monticola ssp. n. are similar to those of the nominate subspecies, except for the inner ornamentation of the genital atrium and penis that is, contrary to W. jessicae jessicae sp. n., completely smooth, without any pleats or folds. Minor differences can also be found in the length of the 86 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) Figures 162-166. Genitalia and anatomy of Wollastonia jessicae monticola ssp. n. Zimbreiro, 200 m SW of the village: 162, 163 whole genitalia excluding gonads 164 ornamentation of the inner walls of the distal penis, the distal vagina and the genital atrium 165 penial papilla and inner ornamentation of proximal penis 166 mantle border. Scale bars 1 mm. vaginal digitiform glands that are, on average, longer in W. jessicae monticola ssp. n. (see Figs 162-166). Ecology. W. jessicae monticola ssp. n. lives in a sloping hollow, in a relative hu- mid spot along a temporary creek. The new taxon has been found exclusively under small volcanic stones in shady places under small shrubs. Its ecology differs somewhat from all the other Aystricella and Wollastonia species on Porto Santo, which are usually found under stones in open, dry grassland areas or on exposed rocky cliffs. W. jessicae monticola sp. n. has been found syntopically with Callina bulverii. Distribution. W. jessicae monticola ssp. n. is only known from the area in the vicin- ity of the locus typicus. Despite extensive field research in the area, the subspecies seems to occupy an area not larger than 100 m/” (see Fig. 167 for the distribution map). Comparison and comments. The new taxon differs from the nominate subspe- cies 1. jessicae jessicae sp. n. by the almost complete lack of the upper keel on the body whorl and by the smooth internal genital atrium and penis. There is also a significant distributional gap between the two conspecific taxa and the habitats and altitudinal ranges also differ. In the phylogenetic analyses, W. jessicae monticola ssp. n. represents the sister group of W. jessicae jessicae inside the W. jessicae s. lat. clade, but divergence is rather low which speaks for a relatively recent separation and may be taken as justifica- tion for the subspecific status of the two taxa. Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... 87 Figure 167. Distribution of Wollastonia jessicae monticola ssp. n. Status and conservation. ‘The subspecies has a very limited distribution area of less than | km? (Fig. 167) close to a village and population size is probably rather low. Habi- tat quality is inferred to be declining and potential and ongoing threats to the species include, in our opinion, urbanization, tourism, goat grazing and quarrying. Therefore, the subspecies is considered here to be Critically Endangered (CR Bla, b(iii), 2a, b(iii)). Wollastonia klausgrohi De Mattia & Neiber, sp. n. http://zoobank.org/D4494EC4-F56E-4E01-BD51-FA18D1BEEB58 Figs 168-182 Type material. NMWC Z.2016.013.00003, holotype, from loc. typ., leg. W. De Mat- tia & J. Macor, May 27 2015; NMWC Z.2016.013.00004/4 PT, SMF 348932/4 PT, NHMW 112139/3 PT, CKG/3 PT, CWDM/8 PT, CMN/4 PT, from loc. typ., leg. W. De Mattia & J. Macor, May 24 2015; FW 11156/25 PT, CMN/10PT, ZMH 131209/10 PT, Casa Velhas, next to the W edge of the old quarry, 33°04'06"N/16°18'51"W, 125 m, leg. F Walther, Apr. 3 2017; CGK/3 PT, Portela, 33°03'58"N/16°19'03"W, 146 m, leg. K. & C. Groh, Oct. 25 1980; ANSP H 11784/8 PT, approx. 200 m SE of Casas Velhas, 33°04'04"N/16°18'47"W, 120 m, leg. J. & C. Hemmen & K. & C. Groh, Jul. 8 1983; ANSP H 11772/14 PT, approx. 50 m above Capela da Graca, 33°04'28"N 16°19'26"W, 165 m, leg. J. & C. Hemmen, Jan. 4 1981. 88 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) Figures 168-170. Shells of Wollastonia klausgrohi sp. n. 168 holotype, NUWC Z.2016.013.00003 169, 170 paratypes from the loc. typ. Scale bar 1 mm. Locus typicus. Casa Velhas, next to the W edge of the old quarry, under stones, 33°04'06"N/16°18'52"W, 124 m. Diagnosis. Small Wollastonia species, with two well-developed keels on the body whorl, the upper slightly less distinct than the lower; shell relatively dark coloured (in comparison with W. jessicae); internal walls of vagina, penis and genital atrium smooth, without folds. Description of shell. The shell is dextral and hairless. Its shape is conical and mark- edly scalariform, with deep sutures. The protoconch is completely dark brown with 1.5 to 2.5 whorls. It has fine radial striae starting from the first protoconch whorl; very few and scattered small tubercles may also be present on its last portion. The teleoconch has from 3.4 to 3.9 rapidly increasing whorls. It is dark brown, sometimes with dark violet shades. The dark areas of the shells are mottled with more or less light brown to whitish areas, usually arranged longitudinally and slightly slanting. In most specimens, the lighter areas tend to be more evident along the keel of the body whorl. No band pattern is visible along the upper whorls. On the lower part of the last whorl two dark bands are present. The inner band is usually thinner but more evident, whereas the outer band is usually weaker but broader. ‘The area around the umbilicus is the light- est in colour. Some specimens clearly have two evident keels starting already from the second whorl of the teleoconch, with the upper keel being more distinct than the lower one. In other specimens only the upper keel is visible. The body whorl is equipped with two keels, with the lower, principal keel somewhat protruding and the upper much less Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... 89 distinct. The whorls are flat and form a ‘shoulder’ giving them an angular contour. The external surface has strong, clearly visible, irregularly spaced, growth lines. Irregularly disposed tubercles are present all over the teleoconch. The tubercles are usually small and scattered, usually whitish in colour. The tubercles are somewhat denser along the keels of the penultimate and last whorl, letting the keel(s) appear like a rough, whitish chord. The last whorl is rather large, contributing 60% to the total shell height and descending near the aperture. The umbilicus is open but very narrow, eccentric, and measures approximately 10% of the maximum shell diameter. The aperture is ellipti- cal, with a faint thickening along its columellar portion. This thickening also extends as far as the parietal side of the aperture. The peristome is continuous and distinctly reflected (see Figs 168-170). Measurements. D 5.3 + 0.2 mm (range 5.1-5.5 mm); H 4.0 + 0.2 mm (range 3.8-4.2 mm); FW 2.5 + 0.1 mm (range 2.4—2.6 mm); PA 58.0 + 3.4° (range 52.8— 60.4°); DU 0.5 + 0.07 mm (range 0.4—0.6 mm); NT 57 + 17 (range 46-77); NW 5.6 + 0.1 (range 5.5—5.7) (7 = 25). Ratio D/H 1.3; ratio FW/H 0.6. Body. The head and neck are usually dark grey. The sides and the posterior upper section of the foot are grey. The foot is light grey and the sole is longitudinally divided into three areas. The central area is smooth, whereas the two lateral areas are equipped with bands of muscles that are roughly arranged in a chevron pattern. The mantle bor- der is dark grey, with five more or less developed lobes. ‘The ratio of the lateral and the dorsal lobes varies from specimen to specimen, also in the same population. In some specimens, one of these lobes (either lateral or dorsal) may be totally missing. ‘The walls of the pallial cavity are colourless, without any stripes or spots. A strong pulmonary vein is visible. The jaw is odonthognatous and is very variable in shape, from almost straight to markedly arched. There are many smooth transverse ridges, ranging from 18 to 22 in number. The right ommatophoral retractor is independent from both penis and vagina. Genitalia. The albumen gland is long and connected to an approximately equally long sperm-oviduct that consists of a prostatic and a uterine portion. The prostatic part extends into a thin vas deferens that is approximately as long as the sperm-oviduct and inserts into the penial complex. The distal portion of the uterine part extends into the free oviduct, turning into a vagina at the level of the duct of the bursa copulatrix. The free oviduct is variable in length and can be as long as the vagina or also three times longer. The duct of the bursa copulatrix is usually thin, approximately as long as the penis and uniform in diameter. It terminates into a roundish bursa copulatrix. The transition area between the duct and the bursa is very sharply delimited, with the duct abruptly widening and turning into the bursa. The spermatophore is unknown. One tuft of digitiform glands arises from the proximal part of the vagina. There are usually two glands that are approximately equally long and are never branched. A short and thin vaginal appendix arises from the wall of the vagina, just distal of the glandular tuft. The inner surface of the vagina is almost smooth. The atrium is usually moderately long and thin. Its internal walls are smooth. The penial complex consists of a flagellum, an epiphallus (which extends from insertion of the vas deferens to penial 90 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) retractor muscle) and a penis that inserts into the atrium. The penial flagellum is short, remarkably cylindrical and with a blunt apex. It is usually as long as the epiphallus. Its internal walls are completely smooth. The epiphallus is usually short. Its internal walls are also completely smooth. The retractor muscle is moderately large, strong and vari- able in length. The penis lacks any muscular or glandular sheath. It is thick-walled and approximately three to four times longer than the flagellum. It is usually cylindrical but sometimes slightly swollen and partially folded up. Sometimes, a thin sheath made of light connective tissue envelopes the distal folded part of the penis. The inner walls of the penis are smooth. ‘The penial papilla is usually small, sometimes swollen at its base, reaching 1/8 to 1/10 of the total length of the penis and is conical in shape. It has smooth external walls, with the opening emerging apically. The channel of the penial papilla is thin and narrow, with somewhat fringed internal walls. The inner lumen of the penial papilla is occupied by a spongy and sturdy tissue, which directly connects with the walls of the epiphallus. As all the previous species, the longitudinal section of the penial papilla shows that its walls are the continuation of the penial walls that abruptly bend inward (see Figs 171-181). Distribution. Wollastonia klausgrohi sp. n. is found along the Casa Velhas area, just W of an abandoned quarry. The W tip of the quarry (currently not exploited at this part) is located only 10 meters from the type locality. Casas Velhas is located along the road n° 233 in direction of Zimbreiro, 340 m ENE from the Miradouro de Portela, E of Vila Baleira. The species has also been collected at Portela, ca. 300 m WSW of the loc. typ., Capela da Graga and 200 m SE of the type locality. The species seems to oc- cupy less than 2,000 m’. For the distribution see Fig. 182. Ecology. Wollastonia klausgrohi sp. n. is commonly found under volcanic rocks of a low stone wall built in an open field in a sloping grassland. The specimens directly aestivate on the lower surfaces of the rocks, frequently forming clusters of individuals attached one to another. Etymology. Named for our co-author, the German malacologist Klaus Groh from Bad Diirkheim to honour his contributions to the malacology of continental snails on the Mid-Atlantic islands. Comparison and comments. Wollastonia klausgrohi sp. n. has a shell similar to W jessicae jessicae sp. n. but differs from that species by a last whorl that is always bicarinate, with the upper keel well-visible. The overall colour of the shell is constantly darker in W. klausgrohi sp. n. More important differences are found in the genital anatomy. The internal walls of the penis, atrium and vagina of W klausgrohi sp. n. are completely smooth, without any pleating or folding. It also resembles H. bicarinata, which, howev- er, always has a wider and shorter atrium and a shorter vagina. Molecular investigations revealed that W. klausgrohi sp. n. is rather closely related to the morphologically very distinct W. oxytropis, whereas W. jessicae sp. n. is closely related to W. leacockiana that occurs in the western part of Porto Santo. Altough W. klausgrohi sp. n. was not recovered as a monophyletic group in the phylogenetic analyses (Fig. 5), statistical support values are rather low, so that monophyly can also not be excluded. We assume that the resolu- tion power of the used molecular markers is insufficient to separate W. klausgrohi sp. n. Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... 91 Figures 171-181. Genitalia and anatomy of Wollastonia klausgrohi sp. n. Casa Velhas, next to the W edge of the old quarry: 171 section of penial papilla 172 ornamentation of the inner walls of the flagel- lum, the penial complex, the vagina and the genital atrium 173 penial papilla 174, 175 whole genitalia excluding part of OSD, AG and gonads 176 ornamentation of the inner walls of the flagellum, the penial complex, the vagina and the genital atrium 177 section of vagina 178 jaw 179 mantle border 180 penial papilla 181 whole genitalia excluding gonads. Scale bars: 1 mm. and W. oxytropis. However, both taxa can easily distinguished by size, shell shape, orna- mentation and genital anatomy (Figs 168-181, 183-194) and therefore are separated as distinct species here. Their distribution areas partly overlap, so that we can currently 92 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) Figure 182. Distribution of Wollastonia klausgrohi sp. n. not exclude mitochondrial introgression as an explanation for the observed pattern in the phylogenetic tree in Fig. 5. Another explanation could be incomplete lineage sort- ing/ancestral polymorphism assuming a relatively recent divergence of the two taxa. An approach using population genetic markers could help to resolve these issues, such an approach is, however, beyond the scope of the present investigation. Status and conservation. Because of the very restricted distribution area, the low number of known subpopulations (Fig. 182) and the probably very small population size as well as potential threats through tourism, goat grazing and especially ongoing quar- rying, the species is regarded as Critically Endangered (CR Bla, b(iii), 2a, b(iii)) here. Wollastonia oxytropis (R. T. Lowe, 1831), comb. n. Figs 183-195 List of synonyms. 1831 Helix oxytropis R. T. Lowe: 57, pl. 6 fig. 18. 1846 Helix oxytropis — L. Pfeiffer: 142, pl. 91 figs 12-13. 1847 Helix oxytropis — L. Pfeiffer in L. Pfeiffer 1847-1848: 190. 1854 Helix oxytropis — Reeve in Reeve 1851-1854: pl. 138 fig. 868. 1854 Helix (Ochthphila) oxytropis — Albers: 37, pl. 9 figs 8-10. 1855 Helix (Hystricella) oxytropis — R. T. Lowe: 186. Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... 93 1867 Helix (Octephila) oxytropis — Paiva: 46. 1878 Helix (Hystricella) oxytropis — Wollaston: 167-168. 1888 Helix oxytropis —Tryon in Tryon and [Pilsbry] 1888: 33, pl. 7 fig. 92. 1894 Geomitra oxytropis — Pilsbry in Pilsbry 1893-1895: 242. 1931 Geomitra (Actinella) oxytropis — Nobre: pl. 2 fig. 3. 1950 Discula (Hystricella) oxytropis —- Mandahl-Barth: 31, 55. 1983 Discula (Hystricella) oxytropis oxytropis — Waldén: 267. 2002 Geomitra oxytropis oxytropis — Bank et al.: 124. 2006 Discula oxytropis — Cameron et al.: 40 [partim]. 2008 Adystricella oxytropis — Seddon: 79, map 180. 2009 = Aystricella oxytropis oxytropis — Groh et al.: 21, fig. 26. 2011 Adystricella oxytropis — Seddon: e.T6728A12801442. Type material. NHM 1968.546, lectotype, (herewith designated), from loc. typ., ex coll. R. T. Lowe; NHM 1948.7.8.12/1 paralectotype, from loc. typ., ex coll. R. T- Lowe. See Fig. 183 for the original figure of Helix oxytropis R. T. Lowe, 1831 from Lowe (1831: pl. 6 fig. 18) and Figs 184-185 for the lectotype/paralectotype of Helix oxytropis R. T. Lowe, 1831 (Photo: P. Crabb, NHM). Further material examined. All from Porto Santo, CGK/1, CMN/1, Pico do Magarico, under stones close to the top, 33°04'00"N/16°18'17"W, 200 m, leg. K. Groh & J. Hemmen, Jul. 10 1983; CKG/2, Pico do Magarico, under stones close to the top, 33°04'00"N/16°18'17"W, 250-285 m, leg. K. Groh & J. Hemmen, Jul. 10 1983; CGK/2, CMN/1, Pico Novalido, W of Pico do Concelho, under stones, 33°04'43"N/16°18'19"W, 185 m, leg. K. Groh & J. Hemmen, Jun. 29 1983; CGK/5, Pico do Concelho, SW slopes, under stones, 33°04'37"N/16°17'47"W, 200-230 m, leg. K. Groh & J. Hemmen, Jun. 29 1983; CKG/2, slope of the Pico do Magarico from Serra de Fora via Casas Velhas, 33°04'05"N/16°18'35"W, 100-200 m, leg. K. Groh & J. Hemmen, Jul. 10 1983; CKG/5, Pico do Baixo, W slope, 33°03'45"N/16°17'58"W, 150-210 m, leg. K. Groh & J. Hemmen, Jun. 9 1983; CWDM/25, ridge of Pico do Concelho towards E, under stones in grassland, 33°04'42" N/16°17'56"W, 270 m, leg. W. De Mattia & J. Macor, May 18 2015; CFW 11153/<10, Pico do Concelho, ridge W of the summit, 33°04'43"N/16°18'07"W, 220 m, leg. F Walther, Apr. 3 2017; ANSP H 11846/10, CMN/2, Pico do Concelho, S slope, 33°04'41"N/16°17'57" W, 280 m, leg. J. Gerber, K. Groh & J. Hemmen, Aug. 12 1985; ANSP H 11845/3, Pico do Con- celho, W slope, 33°04'43"N/16°18'04"W, 250 m, leg. K. Groh & J. Hemmen, Jun. 29 1983; ANSP H 11848/12, Pico Malhada, NW slope, 33°04'00"N/16°18'02"W, 120 m, leg. K. Groh & J. Hemmen, Jul. 7 1983; ANSP H 11851/12, Pico do Ma- caricos, 33°03'58"N/16°18'13"W, 230 m, leg. K. Groh & J. Hemmen, Jul. 10 1983; ANSP H _ 11850/3, Pico do Magarico, SE slope, 33°03'57"N/16°18'11"W, 220 m, leg. J. & C. Hemmen, Jan. 8 1981; ANSP H 11849/7, Pico do Concelho, SW slope, 33°04'42"N/16°17'51"W, 200 m, leg. K. Groh & J. Hemmen, Jun. 29 1983; ANSP H 11849/14, approx. 0.5 km S Serra de Dentro, 33°04'44"N/16°18'26"W, 125 m, leg. J. & C. Hemmen, Jan. 6 1981; ANSP H 11852/9, Pico do Magarico, NE slope, 33°03'58"N/16°18'09"W, 210 m, leg. K. Groh & J. Hemmen, Jul. 10 1983; ANSP H 94 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) Figures |83-187. Shells of Wollastonia oxytropis. 183 original figure of Helix oxytropis R. T. Lowe, 1831 from Lowe (1831: pl. 6 fig. 18) 184 lectotype of Helix oxytropis R. T. Lowe, 1831, NMH 1968.546 ex coll. Lowe 185 paralectotype 186, 187 recent specimens from Pico do Concelho. Scale bar 1 mm. 11847/11, Pico do Baixo, 33°03'45"N/16°17'57"W, 150 m, leg. K. Groh & J. Hem- men, Jul. 9 1983; ZMH 24293/2, Porto Santo, without exact locality data, ex coll. Altonaer Museum; ZMH 24294/1, Porto Santo, without exact locality data, ex coll. Altonaer Museum, ex coll. O. Semper, ex coll. Dohrn. Locus typicus. Hab. in collibus maritimis Portus S". Original description. From Lowe 1831: H. testa depresso-conoidea, supra planu- lata, perforata, carinata, tota scabra, fusca, sub-fasciata: spira depresso-conica; sutura distincta; anfractibus planiusculis; ultimi carina acuta, distinctissima, supra marginata sc. exarata vel sulco expressa; omnibus distinctissime confertim granulatis, asperis: um- bilico minimo, sub-spirali, aperto: apertura rotundata; peristomate continuo, circi- nato, disjuncto, reflexo. Axis 2 %4 lin. Diam. 4. Anfr. 6 Y%. Shell. The shell is dextral and hairless. Its shape is rather flattened, almost discoi- dal, with shallow sutures. The protoconch is brownish with 2—2.3 whorls. It is almost Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... 95 smooth along the first whorl and shows fine radial striae along its remaining portion. The teleoconch has from 3.3 to 3.8 rapidly increasing whorls. It is violet-dark brown with brick red and/or dark violet shades. The dark areas of the shells are mottled with more or less brown to whitish areas, usually arranged longitudinally and slightly slant- ing. No band pattern is visible along the upper whorls. On the lower part of the last whorl two dark bands are visible that can be more or less broad. The peripheral band is usually broader, even if often rather blurred. In some specimens, the two bands merge together and form a single broad, dark band. The area around the umbilicus is the lightest in colour. The teleoconch whorls are rather flat, with shallow sutures and without a visible keel. The body whorl shows a strong, single keel that sometimes even bends slightly downwards. This keel is of the same colour as the remaining shell and its ornamentation pattern is also not markedly different from that of the rest of the shell’s surface. Contrary to most other Hystricella and Wollastonia species, the keel does not have the appearance of a rough chord. ‘The surface of the teleoconch is ornamented with very fine, irregularly spaced growth lines and regularly arranged, very densely set small tubercles. The last whorl is usually large, contributing approximately 50% to the total shell height and descending near the aperture. The umbilicus is open, eccen- tric, and measures approximately 10% of the maximum shell diameter. The aperture is elliptical, with a faint thickening along the columellar portion of the stoma. This thickening can also extend as far the parietal side of the aperture. The peristome is continuous, thin, slightly reflected, with the columellar margin somewhat thicker and more reflected (see also Figs 186-187). Measurements. D 6.8 + 0.2 mm (range 7.5—-8.0 mm); H 4.9 + 0.4 mm (range 4.4-5.5 mm); FW 2.7 + 0.2 mm (range 2.3-3.0 mm); PA 35.8 + 6.9° (range 33.6— 39.3°); DU 0.5 + 0.05 mm (range 0.4-0.6 mm); NT 89 + 14 (range 96-61); NW 5.6 + 0.3 (range 5.3-5.9) (” = 25). Ratio D/H 1.4; ratio FW/H 0.6. Body. As in the genus description. Wollastonia oxytropis tends however, to have an overall darker body colouration than the other Wollastonia species. Genital anatomy. The albumen gland is long and thin and is connected to an ap- proximately twice as long sperm-oviduct. The thin vas deferens is roughly 1.5 times longer than the penial complex. The free oviduct is as long as the vagina. The duct of the bursa copulatrix is thin, approximately as long as the penial complex and uniform in diameter. It ends in a roundish bursa copulatrix. The transition area between the duct and the bursa itself is sharply delimited, with the duct abruptly widening and turning into the bursa. The spermatophore is unknown. One tuft of digitiform glands arises from the proximal part of the vagina. There are usually three glands that are ap- proximately equally long and very rarely branched. A short and thin vaginal appendix arises from the wall of the vagina, just distal of the glandular tuft. The internal walls of the vagina are smooth, as are those of the atrium. The atrium is relatively long and thin. The penial flagellum is very short, cylindrical, has a somewhat blunt to slightly pointed apex, and is usually much shorter than the epiphallus. The epiphallus is ap- proximately as long to slightly longer than the penis. Its internal wall is equipped with irregular longitudinal pleats. The retractor muscle is rather large, strong and of variable 96 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) Figures 188-194. Genitalia and anatomy of Wollastonia oxytropis from Pico do Concelho: 188 whole genita- lia excluding part of OSD, AG and gonads. Pico do Concelho: 189 section of vagina 190 tip of penial papilla 191 whole genitalia excluding part of OSD, AG and gonads 192 mantle border 193 ornamentation of the inner walls of the flagellum, the penial complex, the vagina and the genital atrium 194 jaw. Scale bars 1 mm. length. The penis misses any muscular or glandular sheath. It is thick-walled, cylindri- cal, and slightly swollen at the level of the penial papilla. The inner walls of the penis are smooth. ‘The penial papilla is small and it usually has a blunt shape. It has smooth external walls with the opening emerging apically. The channel of the penial papilla is thin and narrow. The inner lumen of the penial papilla is occupied by a spongy and sturdy tissue which directly connects with the walls of the epiphallus. The longitudinal section of the penial papilla shows that its walls are the continuation of the penial walls that abruptly bend inward. See Figs 188-194. Ecology. Wollastonia oxytropis is found under volcanic stones and rocks, in rock crevices and stone walls built in open, stony fields in sloping grasslands. Distribution. Wollastonia oxytropis has a quite small distributional range. It is known to live exclusively along the SE part of the main island Porto Santo: Pico do Baixo, Pico do Magarico, Pico do Concelho and Pico do Novalido. These localities were confirmed by survey performed during the 1980’s and 1990’s (CKG, ANSP, Seddon 2008). During recent field surveys (WDM 2012, 2014, 2015, FW 2017) the species has been found living only along the ridge of Pico do Concelho despite careful searches at the above-mentioned localities. The known distribution is shown in Fig. 195. Revision of the genus-group Hystricella R. 77 Lowe, 1855 from Porto Santo... 97 Figure 195. Distribution of Wollastonia oxytropis. Filled circles refer to recent and open circles to fossil records. Comparison and comments. Wo/lastonia oxytropis is clearly distinguishable from all other species belonging to Hystricella and Wollastonia by the presence of only a sin- gle keel, the rather flattened, almost discoidal shell with very shallow sutures and the extremely short flagellum in relation to the epiphallus. Taxonomic remarks. Wollastonia oxytropis is somewhat unusual among the Wol- lastonia gen. n. species as its genital anatomy is more similar to that of the investigated Discula s. str. species than to that of the remaining Wollastonia gen. n. species. How- ever, in the cox] phylogeny (Fig. 5) the species is clearly embedded within the Wol/as- tonia gen. n. clade indicating that the similarity in genital anatomy may be the result of convergence. We therefore include the species in Wollastonia gen. n. until more in depth phylogenetic analyses and morphological comparisons of the remaining geomi- trinid genera will become available. Status and conservation. According to the current IUCN assessment (Seddon 2011d) the species is considered Near Threatened (NT). Recent surveys indicate how- ever that the species is clearly declining, both with regard to the distributional range and population size, probably as a result of a decline in habitat quality. The area of occupancy and extent of occurrence is approximately 4 km? and the species occurs at relatively few localities clustered in the upper parts of two hills in the south-eastern part of Porto Santo. Therefore, the species should be considered as Endangered (EN Bla, b(iii), 2a, b(iii)). 98 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) + Wollastonia subcarinulata (Wollaston, 1878), comb. n., stat. n. Figs 196-199 List of synonyms. 1878 Helix (Hystricella) oxytropis var. & subcarinulata Wollaston: 168. Discula (Hystricella) oxytropis subcarinulata — Waldén: 267. 2002 Geomitra oxytropis subcarinulata — Bank et al.: 124. 2006 Discula oxytropis — Cameron et al.: 49 (1): 40 [partim]. 2009 Aystricella oxytropis subcarinulata — Groh et al.: 21, fig. 27. Type material. NMWC 80.202, Acc 55.158, lectotype (herewith designated), from loc. typ., ex coll. Melvill-Tomlin, ex coll. T. V. Wollaston, Porto Santo, Madeira (Fig. 196). Further material examined. All from Porto Santo, CKG/2, Barbinha, Quaternary aeolinites, layer 3, 33°04'04"N/16°17'49" W, 8 m, leg. K. & C. Groh, Jul. 4 1983 and Jul. 5 1983; CKG/3, Barbinha, Quaternary aeolinites, layer 2, 33°04'04"N/16°17'49" W, 8 m, leg. K. & C. Groh, Jul. 4 1983 and Jul. 5 1983; CKG/8, Barbinha, Quaternary aeolinites, layer 4, 33°04'04"N/16°17'49"W, 8 m, leg. K. & C. Groh, Jul. 4 1983 and Jul. 5 1983; CKG/2, south coast, ditch at the in 1985 new built harbour E Vila Baleira, lens of Quaternary calcareous aeolinite within a marine sandy beach terrace (nowadays beyond the quay of the harbour), 33°03'50"N/16°18'56"W, 0-1.5 m, leg. J. Gerber, K. Groh & J. Hemmen, Aug. 1985; CWDM_/8, Porto Santo, excavated mud walls behind the cart speedway E of the new harbour of Porto Santo, 33°03'48"N/16°18'22"W, 30 m, leg. W. De Mattia & J. Macor, May 24 2015; CWDM/9, E of Vila Baleira, S slope of the hill above Vale do Touro, 50 m W of the oil tanks, excavated Quaternary mixed gravel, 33°03'47"N/16°19'26"W, 24 m, leg. W. De Mattia & J. Macor, May 24 2015; CWDM/20, Pico do Baixo, E entrance of the tunnel, Quaternary mud deposit, 33°03'44"N/16°17'45"W, 20 m, leg. W. De Mattia & J. Macor, May 24. 2015. Locus typicus. Porto Santo, Madeira. Original description. From Wollaston 1878: var. 8. subcarinulata — Major, spira magis elevata, ad apicem paulo magis acuta, anfractibus in medio obsolete subcarinu- lato. — Long. axis 2% lin.; diam. 3%. Redescription of shell. Shell large for the genus, with 5.9 regularly increasing whorls, the protoconch with 2.5 whorls. ‘The form of the shell is flat conical, the slight- ly convex teleoconch whorls separated by a rather shallow but distinct, simple suture. The body whorl shows a distinct, rounded keel, which is enforced by a concave impres- sion below the periphery. ‘The last whorl measures 70%, the penultimate whorl 10% of the total shell height. The lower part of the body whorl is beneath the significantly marked periphery in frontal view first significantly concave, later moderately convex. The keel of the body whorl is located in the upper part of its total height. The aperture, which is inclined to the vertical axis of the shell in an angle of 69° and is descending in its last 5% in an angle of 36° to the horizontal axis, is horizontally elliptic; towards the umbilical region a bit flattened. Its width measures 45% of the total shell width and its height 44% of the total shell height. It has a peristome that is completely detached Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... 99 This tad bEWellash eché no.b7 Helix oxycrepesLowe wena the Gromnule haw F v Fy i ele x XY th ee laure pace dh those Wether ca 03362, Porlés San lé ./Nadeira Figures 196-198. Shells of Wollastonia subcarinulata. 196 \ectotype, NMWC 80.202, Acc 55.158; 197-198 specimens from Barbinha, Quaternary aeolinites. Scale bar 1 mm. from the body whorl. The lip is distinctly reflected and very wide at its basal and colu- mellar sides. The eccentric umbilicus, which measures 13% of the total shell width, is shaped like a pinhole in the penultimate whorl and is completely closed in the earlier whorls. The protoconch is smooth, the teleoconch shows a number of oblique radial ribs, 10 in the penultimate quadrant of the body-whorl and is additionally covered by numerous small, round tubercles. The number of tubercles in the standard-quadrate of the base is 108. There is a yellowish hue of colouration in the teleoconch; the keel usually distinctly lighter in colour. See Figs 197-198. Measurements. D 8.3 + 0.3 mm (range 8.0-8.8 mm); H 6.1 + 0.3 mm (range 5.8-6.4 mm); FW 4.1 + 0.2 mm (range 3.9-4.2 mm); PA 48.2 + 5.4° (range 42.6— 54.4°); DU 0.6 + 0,06 mm (range 0.5-0.7 mm); NT > 100; NW 5.6 + 0.1 (range 5.5—5.7) (” = 25). Ratio D/H 1.4; ratio FW/H 0.5. Distribution. Wollastonia subcarinulata is known from the Quaternary mud and slope deposits along the southern slopes of the eastern part of Porto Santo, from Vale do Touro to the area around the tunnel of Ponta da Galé beneath Pico do Baixo. See map in Fig. 199. Comparison and comments. Wollastonia subcarinulata can be confused with other, rather large-sized species of the genera Hystricella and Wollastonia like H. echinoderma, W. vermetiformis and W. falknerorum sp. n., from which it can be distinguished, however, by the lack of a second keel, the rounded shape of the keel and a much finer sculpture as 100 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) Figure 199. Map of the distribution of Wollastonia subcarinulata. well as a more depressed conical form. From similar sized W. inexpectata it is distinguish- able by the wider umbilicus, a flatter form, the keeled instead of angled periphery and a coarser granulation. From the recent W. oxytropis it can be distinguished by the larger size, the wider umbilicus, the more convex teleoconch whorls that are separated by a more distinctly marked suture, the finer granulation, and the wider aperture. Taxonomic remarks. ‘The species is included in the genus Wollastonia because of its size and surface sculpture that is similar to W. oxytropis, as a variety of which it was described by Wollaston (1878). Subsequent authors, except Cameron et al. (2006), recognised the taxon as a subspecies of W. oxytropis. Status and conservation. Extinct before the islands’ scientific exploration in the 19" century, possibly already before human settlement. + Wollastonia inexpectata De Mattia & Groh, sp. n. http://zoobank.org/AD8D733D-8248-47EF-B9D8-CF9A3F83 1437 Figs 200-203 Type material. SMF 348930, holotype, from loc. typ., leg. W. De Mattia, May 23 2015 (see Fig. 200); NMWC 80.202 Acc. 55 158/1 PT as “Geomitra oxytropis Lowe var. subcarinulata Woll. Porto Santo” (Figs 201-202). Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... 101 » Hystricella oxykro pis. e, vo. pubeatmubllgl Porte santo, madeira [Wollaston] Fo-208 __ Ace 86 188 oxytropis, Loire. Figures 200-202. Shells of Wollastonia inexpectata sp. n. 200 holotype, SMF 348930 201 paratype NMWC 80.202 Acc. 55 158 202 label of the lot NMWC 80.202 Acc. 55 158. Scale bar 1 mm. Locus typicus. N of airport, end of the runway towards Fonte de Areia, Quater- nary calcareous sand deposit, 33°05'25"N/17°20'58"W, 99 m. Etymology. Wollastonia inexpectata sp. n. was unexpectedly found during inten- sive field researches at Fonte da Areia when looking for H. echinoderma. Diagnosis. Shell large for the genus, solid and conical. Whorls rounded, with deep sutures. The shell’s surface covered with small and densely set tubercles. Body whorl with a peripheral, rounded keel. Other teleoconch whorls without visible keel. Umbilicus narrow but open. Last whorl descending toward the aperture. Aperture oval with continuous peristome. Description of the shell. The shell is large for the genus and rather conical, with 6.3 regularly increasing whorls. The protoconch has 1.7 smooth whorls. The teleoconch whorls are distinctly convex and separated by a slightly impressed but simple suture. The body whorl measures 64% and the penultimate whorl 14% of total shell height and is descend- ing towards the aperture in its last 5% in an angle of 36°. It has a rounded angulation in its upper third (in relation to total shell height) that is emphasised by a narrow, only slightly concave impression below the periphery. The lower part of the body whorl (beneath the pe- riphery in frontal view) is otherwise rather convex. The aperture is regular elliptical, meas- uring 49% of the total shell width and 29% of the total shell height. It is inclined to the vertical axis of the shell in an angle of 57°. The peristome is completely detached from the body whorl, expanded and distinctly reflected, especially in its basal and columellar part. The umbilicus is eccentric, closed in the early whorls, but pinhole-like in the body whorl, measuring 5% of the maximum shell diameter. The sculpture of the teleoconch consists of oblique radial ribs (10 in the penultimate quadrant of the body whorl) and numerous, small roundish tubercles (132 in the standard square basal surface of the shell). The colour 102 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) Figure 203. Distribution of Wollastonia inexpectata sp. n. is only preserved as a yellowish hue on the teleoconch, a slightly darker yellowish, narrow spiral band in the middle of the base and a lighter marked keel. See Figs 200-202. Measurements. D 8.6 mm; H 6.7 mm; FW 4.6 mm; PA 41.2°; DU 0.6 mm; NT 132; NW 6.6 (z = 1). Ratio D/H 1.3; ratio FW/H 0.7. Distribution. The species is only known from the type locality. A second specimen is known from another unidentified locality in Porto Santo and housed at the NMW. The distribution is shown in Fig. 203. Comparison and comments. Wollastonia inexpectata sp. n. is superficially similar to H. echinoderma, or W. vermetiformis and W. falknerorum sp. n. but is readily dis- tinguishable from these species by the lack of a second keel, its much finer sculpture, narrower umbilicus and regular conical form. From H. echinoderma it can also be distinguished by its regular convex and not stepped whorls and the regular elliptical rather than oblique ovate aperture. From the similarly sized W. subcarinulata it differs in the narrower umbilicus, the presence of a rounded keel, the finer granulation and the higher shell in relation to its width. Taxonomic remarks. Wollastonia inexpectata sp. n. is included in the genus Wol- lastonia because it is similar to W. oxytropis in size and surface sculpture. It is noteworthy that the similarly large-sized Hystricella echinoderma most probably also originates from the Quaternary deposits in the north of the island, and likewise is extremely rare there. Status and conservation. Extinct before the islands’ scientific exploration in the 19" century, possibly already before human settlement. Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... 103 Callina R. T. Lowe, 1855 Helix (Callina), Wollaston (1878). Helix (Tectula), sensu Albers (1854) and Paiva (1867), partim. Geomitra (Actinella (Callina)), Pilsbry (1893-1895), partim. Discula (Callina), Mandahl-Barth (1950), Seddon (2008) and Bank et al. (2002). Callina, Waldén (1983) and Schileyko (2005). Type species. Helix [Helicella| rotula R. T. Lowe, 1831 by monotypy. Description of the genus. Shell. ‘The shell is dextral, solid, hairless, and it is usually discoidal to tectiform in shape. The protoconch is dark brown, with 1.4 to 1.9 whorls. It is almost smooth along the first whorl and shows radial striae along its remaining portion. The teleoconch has from 5.3 to 6.6 rapidly increasing whorls. It is corneous brown in colour on the upper side, mottled with scattered, light-brown small areas. One chestnut band is visible along the last three whorls. On the underside of the last whorl two brown bands are visible. The extent of these bands can be variable, either both of them very thin or the peripheral band very broad. Sometimes a number of thinner ad- ditional bands are present next to the main peripheral band or, less frequently, no bands are present at all on the underside of the last whorl. The peri-umbilical area is usually the lightest in colour. The spire is broadly conical, making the shell appear discoidal to tectiform. The periphery of the last whorl is either keeled, angled or rounded. The keel is often very distinct and may be slightly bend downwards. ‘The keel is usually lighter in colour than the remaining surface of the whorls, being light brown to whitish. The external upper surface has very fine but clearly visible, relatively regularly spaced growth lines. Regularly disposed, very small, drop-like tubercles are present all over the teleoconch, especially along the growth lines. The dimensions of the tubercles remain quite stable along the whorls and also their density is approximately the same on all teleoconch whorls. The underside of the last whorl is also equipped with tubercles. The last whorl is only slightly wider than the penultimate whorl, and only slightly descend- ing near the aperture. The umbilicus can either be completely closed or widely open (up to '/, of the maximum shell diameter). The aperture is elliptical. The lower palatal side of the last whorl shows a strong callous immediately behind the aperture or none. The peristome is interrupted along the palatal area, and it is only slightly reflected along the lower section. The palatal area never shows any kind callosities or thickenings. Body. Very similar to Wollastonia gen. n. Head and neck are usually grey to dark grey. [he sides and the posterior upper section of the foot are whitish to grey. The foot is whitish and the sole is longitudinally divided into three areas. The central area is smooth, whereas the two lateral portions are equipped with bands of muscles that are roughly arranged in a chevron pattern. The mantle border is dark grey, with five more or less well-developed lobes. In some specimens, one of these lobes (either lateral or dorsal) may be totally missing. The walls of the pallial cavity are colourless without any stripes or spots. A strong pulmonary vein is visible. The right ommatophoral retractor is independent from both penis and vagina. 104 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) Genitalia. The general arrangement of the genitalia is semi-diaulic monotrematic. A convoluted hermaphroditic duct arises from the gonad. The albumen gland is long and thin and connected to an approximately twice as long sperm-oviduct that consists of a prostatic and a uterine portion. The prostatic part extends to a thin vas deferens, which is approximately twice as long as the sperm-oviduct and which is terminating in the penial complex. ‘The distal portion of the uterine part extends into the free oviduct, which transforms into a vagina at the level of the duct of the bursa copulatrix. The free oviduct is approximately as long as the vagina. The duct of the bursa copulatrix is very wide, approximately as long as the penial complex and usually uniform in diameter. It ends with an oval bursa copulatrix. The transition area between the duct and the bursa itself is not sharply delimited but rather gradually widens. ‘The spermatophore is unknown. One tuft of digitiform glands arises from the proximal part of the vagina. There are usually two, equally long and very rarely branched glands present. A vaginal appendix arises from the vagina’s wall, just distal of the glandular tuft. Very smooth, rather wide, and little elevated, irregularly spaced pleats run longitudinally along the inner surface of the vagina, which reach into the genital atrium but not as far as the genital orifice. The atrium is short and wide. Its internal walls are smooth. The penial complex consists of a flagellum, an epiphallus (which extends from insertion of the vas deferens to the penial retractor muscle) and a penis that inserts into the genital atrium. The penial flagellum is short, cylindrical and with a pointed apex. It is usually Y as long as the epiphallus. Its internal walls are completely smooth. ‘The epiphallus is usually 14 longer than the penis. Its internal walls are equipped with 20-25 very fine, elevated, longitudinal pleats. The retractor muscle is strong and approximately half as long as the epiphallus. The penis lacks any muscular or glandular sheath. It is extremely thick-walled and approximately 74 as long as the epiphallus. It is cylindrical and slightly swollen in its distal portion. The inner walls of the penis are smooth. The section where the rather large penial papilla is located is usually detectable from outside by virtue of a circular swelling corresponding to the origin of the papilla itself. The penial papilla is approximately half as long as the entire penis. It is conical to subcylindrical in shape and has smooth external walls, with an apically emerging opening. The channel of the penial papilla is thin and narrow. The inner lumen of the penial papilla is occupied by a spongy and sturdy tissue which directly connects with the walls of the epiphallus. The longitudinal section of the penial papilla (Fig. 226) shows that its walls are the continu- ation of the penial walls that abruptly bend inwards. See also Schileyko (2005). Jaw and radula. The jaw and the radula of Callina are very similar to those of Hystricella and Wollastonia. Only the number of the lateral and marginal teeth may be slightly higher: 21 to 29. They do not differ markedly from each other, i.e. their shape gradually changes from the first laterals towards the marginals. The jaw is odonto- enathous and very variable in shape, from almost straight to markedly arched. There are many smooth transverse ridges, ranging from 9 to 10 in number. For jaws and radulae of Callina species, see Figs 205-207. Distribution. The genus Callina is endemic to the Island of Porto Santo (Ma- deiran Archipelago, Portugal). It is distributed along the central-eastern area of the main island. It is not present on the offshore islets. It is only present east of the line Revision of the genus-group Hystricella R. T) Lowe, 1855 from Porto Santo... 105 Figure 204. Distribution of the genus Callina. Filled circles refer to recent and open circles to fossil records. Vila Baleira-Dragoal-Camacha-Porto de Salemas. For distributional maps of Callina species, see Figs 204, 214, 217 and 224. Taxonomic remarks. The results of the phylogenetic analyses indicate that Cal lina is not closely related to Discula R. T. Lowe, 1852 s. lat. as previously supposed but represents a well-supported clade that is closely related to Hystricella and Wol- lastonia gen. n. ‘The taxon was already elevated to generic rank by Waldén (1995) and Schileyko (2005), although without providing strong morphological evidence for this decision. An isolated position of C. rotula within Discula s. lat. was supposed by these authors on the basis of a closed (or “dot-like” following Schileyko, 2005: 2018) umbilicus. However, our morphological (genitalia) and genetic investigations showed that the closed umbilicus of C. rotula cannot be used as a discriminating feature at the genus-level as the widely umbilicated species C. bulverii also belongs to Callina as supported by the phylogenetic analyses and the peculiar arrangement of pleats on the inner wall of the epiphallus that is equipped with 20-25 very fine, elevated longitu- dinal pleats (Fig. 211). Contrarily, both Discula s. str. and Discula (Mandahlia) have an epiphallus with 5—7 strong and rather elevated, longitudinal pleats (Figs 228-230). The penial papilla of Callina (Fig. 211 and Fig. 225) always reaches the genital atrium, whereas in Discula s. str. and Discula (Mandahlia) the penial papilla is much shorter and never reaches the genital atrium (Figs 228-229). Discula (Mandahlia) has a very long epiphallus, at least three times longer than the penis (Fig. 230). 106 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) Callina rotula (R. T. Lowe, 1831) Figs 206-214 List of synonyms. 1831 Helix rotula R. T. Lowe: 53, pl. 6 fig. 10. 1847 Helix rotula —L. Pfeiffer in L. Pfeiffer 1847-1848: 216. 1854 Helix rotula— Reeve in Reeve 1851-1854: pl. 137 fig. 854. 1854 Helix rotula — Albers: 28, pl. 6 figs 16-18. 1855 Helix (Callina) rotula — R. T. Lowe: 183. 1867 Helix (Tectula) rotula — Paiva: 82. 1867 Helix (Tectula) rotula var. « major Paiva: 83. 1867 Helix (Tectula) rotula var. 8 minor Paiva: 83. 1867 Helix (Tectula) rotula var. y cerina Paiva: 83. 1867 Helix (Tectula) rotula var. § monstrosa Paiva: 83. 1878 Helix (Callina) rotula — Wollaston: 151-152. 1888 Helix rotula —Tryon in Tryon [and Pilsbry] 1888: 46, pl. 9 fig. 4. 1894 Geomitra rotula — Pilsbry in Pilsbry 1893-1895: 241. 1922 Ochthephila (Callina) rotula mut. grisea 'T. D. A. Cockerell: 44—46[ unavailable, infrasubspecific]. 1931 Geomitra (Actinella) rotula — Nobre: 81, figs. 30 + 31. 1950 Discula (Callina) rotula — Mandahl-Barth: 36. 1983 Discula (Callina) rotula — Waldén: 267, note p. 273 [partim]. 2002 Discula (Callina) rotula — Bank et al.: 124. 2005 Callina rotula — Schileyko: 218-219. 2008 Discula (Callina) rotula — Seddon: 78, pl. 28 fig. E, map 175. 2011 Discula rotula — Seddon: e.T156384A4936221. Type material. ANSP 97116, lectotype (herewith designated), from loc. typ., ex coll. R. T. Lowe-T. V. Wollaston. For the original figure of Helix rotula R. T. Lowe, 1831 (from Lowe 1831: pl. 6 fig. 10), see Fig. 206. The lectotype of Helix rotula R. T. Lowe, 1831, W = 11.2 mm, is depicted in Fig. 207. Further material examined. All from Porto Santo. Fossil: CKG/1 juv., SE slope of the Pico de Ana Ferreira, 33°02'36"N/16°22'11"W, 220 m, leg. K. & C. Groh & J. & C. Hemmen, Jul. 6 1983; CKG/15, Barbinha, Quaternary aeolinites, 33°04'04"N/16°17'49"W, 8 m, leg. K. & C. Groh & J. & C. Hemmen, Jul. 4 1983; CFW 11227/1, Ponta da Galé, E end of the tunnel, 33°03'44"N/16°17'45"W, 30 m, leg. F Walther & E. M. Gryl, Apr. 1 2017. Recent: CKG/20, NW slope of Pico Juli- ana, along the upper Ribeira do Pedregal, 33°05'32"N/16°19'25"W, 340 m, leg. K. & C. Groh & J. & C. Hemmen, Jul. 3 1983; CKG/1, Terra Cha, path to forestry house, 33°05'37"N/16°18'03"W, 350 m, leg. K. Groh & J. Hemmen, Jul. 8 1983; CKG/2, region NW of the summit of Pico do Concelho, 33°04'43"N/16°18'01" W, 200-230 m, leg. K. Groh & J. Hemmen, Jun. 29 1983; CKG/1, SW slope of Lombo Branco, 33°05'46"N/16°18'46"W to 33°05'38"N/16°18'13"W, 150-400 m, leg. K. Groh & J. Hemmen, Jul. 8 1983; CKG/5, CMN/5, W slope and crest of the hill W of Pico do Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... 107 2016/08/23 LD38 x1.8k 50 um Figures 205-209. 205, 206 radula and jaw of Callina rotula from Ribeira da Areia 207 radula and jaw of Callina bulverii 208 original figure of Helix rotula R. T. Lowe, 1831 (from Lowe 1831: pl. 6 fig. 10) 209 lectotype of Callina rotula, ANSP 97116. Scale bar 1 mm. Concelho (Pico Novalido?), 33°04'50"N/16°19'37"W, 100-250 m, leg. K. Groh & J. Hemmen, Jun. 29 1983; CKG/10, N slope of Pico do Facho, 33°05'04"N/16°19'25"W, approx. 450 m, Zamarix- and Pinus-forest, leg. K. Groh & J. Hemmen, Jun. 28 1983; CMN/1, Pico de Juliana towards Cabo de Grafa, 33°05'34"N/16°19'11" W, 300 m, leg. K. & C. Groh, Aug. 13 1985; CWDM/5, Ribeira da Areia, serpentine 240 m NNW the quarry, under stones, 33°04'51"N/16°18'41"W, 120 m, leg. W. De Mattia & J. Macor, May 15 2014; CWDM/4, beginning of path to Pico Branco, 33°05'48"N/16°18'54" W, 180 m, leg. W. De Mattia & J. Macor, May 15 2015; CWDM/5, Pico de Cabrita, Pedregal de Dentro, northern slope close to the road, under stones, 33°05'42"N/16°19'23"W, 240 m, leg. W. De Mattia & J. Macor, May 15 2015; CWDM/5, path to Pico Bran- co, terraced S slope of Pico Branco, under stones, 33°05'35"N/16°18'25"W, 270 m, leg. W. De Mattia & J. Macor, May 16, 2015; CWDM/4, Cabeco dos Bades at 108 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) Figures 210-213. Genitalia and anatomy of Callina rotula. Ribeira da Areia: 210 whole genitalia ex- cluding part of OSD, AG and gonads 211 ornamentation of the inner walls of the flagellum, the penial complex and the genital atrium 212 penial papilla 213 cross section of penial papilla. Scale bars 1 mm. the confluence of the two streams, under stones, 33°06'09"N/16°19'25"W, 30 m, leg. W. De Mattia & J. Macor, May 19 2015; CWDM/4, lake S of Ribeira da Serra de Dentro, under stones, 33°04'48"N/16°18'31"W, 70 m, leg. W. De Mattia & J. Macor, May 22 2014; CWDM/5, path to Pico Branco, eastern steep part of path to Pico Branco, under stones, 33°05'29"N/16°18'14"W, 320 m, leg. W. De Mattia & J. Macor, May 22 2014; CWDM/4, Terra Cha-Pico Branco ridge, under stones, 33°05'38"N/16°18'01" W, 335 m, leg. W. De Mattia & J. Macor, May 16 2014; ZMH 120617/2, Pico de Cabrita, 33°05'02"N/16°19'16"W, 460 m, ex coll. W. Fauer, leg. M. Vilella, Mar. 20 1964; ZMH 120616/3, Pico de Juliana, 33°05'32"N/16°19'18"W, c. 360 m, ex coll. W. Fauer, leg. J. & C. Hemmen, Jun. 29 1983; ZMH 110140/15, slopes of Pico do Facho, under stones, c. 33°04'59"N/16°19'25"W, 430 m, leg. E. Clauss, Sep. 22 1992; CFW 10997/11, ZMH 92904/4, southwestern slope of Pico Branco ca. 250 m SW of the top, 33°05'29"N/16°18'13"W, 310 m, leg. EF Walther & E. M. Gryl, Mar. 31 2017; CFW 11107/2, S slope of Pico do Castelo, near parking area, 33°04'42"N/16°20'06"W, 260 m, leg. F. Walther & E. M. Gryl, Apr. 1 2017; CFW 10889/4, ZMH 92889/1, Ribeiro do Pedregal, upper part, downstream of the abandoned houses, 33°05'59"N/16°19'32"W, 120 m, leg. F Walther, Apr. 2 2017; CFW 10878/4, Pico do Facho, northwestern slope, 33°05'13"N/16°19'38"W, 370 m, leg. E Walther & E. M. Gryl, Apr. 3 2017; CFW 10835/4, ZMH 92811/1, Casinhas, N of Capela da Graga, 33°04'27"N/16°19'27"W, 150 m, leg. EF Walther, Apr. 4 2017; ZMH 24259/2, Madeira Archipelago, without locality data, ex coll. Altonaer Museum; ZMH 24260/1, Porto Santo, without exact locality data, ex coll. Altonaer Museum, ex coll. O. Semper, ex coll. Dohrn. Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... 109 Loci typici. [votu/a] Hab. in montibus Portus S".; not given for the var. « to 6 of Paiva. [grisea] Porto Santo, main island. Original descriptions. [votu/a] from Lowe, 1831: H. testa rotundata, conoideo- depressa, supra sub-planulata, subperforata, carinata, scabra, nitidiuscula, fasciata: spira conoidea, obtusissima; sutura obsoleta; anfractibus planis, transverse striatis et granulatis; ultimo acute carinato, carina ad peristoma obsoleta: aperture lunata, ex- trorsum dilatata; peristomate intiis incrassato, acuto, sub-expanso; ad angulum in- ternum reflexo, calloso, perforationem obtegente. Axis 3 lin. Diam. 6. Anfr. 8; from Paiva 1867: [var. « major] testa distincto fasciata, ampliore; [var. 8 minor] testa du- plo minore, fasciis angustioribus; [var. y cerina] testa albida, omnio efasciata; [var. 6 monstrosa] testa fasciata, anfractibus subsolutis, carina supra suturam valde prominent, rarissima; [grisea]; from Cockerell 1922: Shell pale gray, flecked with creamy white. Redescription of shell. The shell is dextral, solid, hairless, and it is usually discoi- dal to tectiform. The protoconch is dark brown with 1.5 to 1.7 whorls. It is almost smooth along the first whorl and shows radial striae along its remaining portion. The teleoconch has from 5.3 to 5.7 rapidly increasing whorls. It is more or less horn brown in colour on the upper side, mottled with small light brown scattered areas. One more or less blurred chestnut band is sometimes visible along the last three whorls. On the underside of the last whorl two brown bands are present that are usually not very broad. The peri-umbilical area is usually the lightest in colour. The spire has a rather conical shape, with the rather flat whorls being separated by a shallow suture. ‘The last whorl is not equipped with a real keel but is rather distinctly angled. ‘The peripheral angulation is often lighter in colour compared to the remaining shell surface, usually whitish to light brown. The surface of the shell has very fine but clearly visible, rather regularly spaced, growth lines. Small tubercles are present on surface of the teleoconch. These tubercles are concentrated along the growth lines, and especially on the un- derside quite dense. The last whorl is only slightly wider than the penultimate whorl and slightly descending towards the aperture. The umbilicus is completely closed. The aperture is elliptical and the peristome is white and rather solid. A strong callous along the lower palatal side is present, just behind the aperture. The peristome is interrupted along the palatal area and partially reflected along its basal side. ‘The palatal area does not show any callouses or thickenings. See Figs 208-209. Measurements. D 11.1 + 0.6 mm (range 10.4-11.8 mm); H 7.2 + 0.7 mm (range 6.3-8.1 mm); FW 5.1 + 0.2 mm (range 5.0—5.6 mm); PA 44.0 + 2.1° (range 42.3- 46.4°); NT > 100; NW 7.4 + 0.3 (range 7.0-7.6) (” = 25). Ratio D/H 1.6; ratio FW/H 0.6. Body. As in the genus description. Genital anatomy. A convoluted hermaphroditic duct arises from the gonad. The albumen gland is long and thin and is connected to an approximately twice as long sperm-oviduct that consists of a prostatic and a uterine portion. The prostatic part ex- tends into a thin vas deferens which is approximately twice as long as the sperm-oviduct and which inserts into the penial complex. The distal portion of the uterine part ex- 110 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) tends into the free oviduct, turning into a vagina at the level of the duct of the bursa copulatrix. The free oviduct is approximately as long as the vagina. The duct of the bursa copulatrix is very wide, approximately as long as the penial complex and usually uniform in diameter. It ends in an oval bursa copulatrix. The transition area between the duct and the bursa itself is not sharply delimited but rather gradually widens. The spermatophore is unknown. One tuft of digitiform glands arises from the proximal part of the vagina. There are usually two, equally long and very rarely branched glands pre- sent. A vaginal appendix arises from the vagina’s wall, just distal of the glandular tuft. Very smooth, rather widely separated, and little elevated, irregularly spaced pleats run longitudinally along the inner surface of the vagina, reaching the genital atrium but not as far as the genital orifice. The atrium is short and wide. Its internal walls are smooth. The penial complex consists of a flagellum, an epiphallus (which extends from inser- tion of vas deferens to penial retractor muscle) and a penis that inserts into the genital atrium. The penial flagellum is short, cylindrical and with a pointed apex. It is usually Y4 as long as the epiphallus. Its internal walls are completely smooth. ‘The epiphallus is usually 13 longer than the penis. Its internal walls are equipped with 20-25 very fine, and elevated longitudinal pleats. The retractor muscle is strong and approximately half as long as the epiphallus. The penis lacks any muscular or glandular sheath. It is ex- tremely thick-walled and approximately %4 as long as the epiphallus. It is cylindrical and slightly swollen in its distal part. The inner walls of the penis are smooth. ‘The section where the large penial papilla is located is usually detectable from the outside by virtue of a circular swelling corresponding to the origin of the papilla itself. The penial papilla reaches half the length of the penis. It is conical to subcylindrical in shape and has smooth external walls. Its opening emerges apically. The channel of the penial papilla is thin and narrow. The inner lumen of the penial papilla is occupied by a spongy and sturdy tissue which directly connects with the walls of the epiphallus. The longitudinal section of the penial papilla shows that its walls are the continuation of the penial walls that abruptly bend inwards (See Schileyko (2005) and Figs 210-213). Ecology. Callina rotula is commonly found under volcanic rocks scattered on grassland in open fields that are more or less sloping. The specimens aestivate among the soil and are only rarely attached to the lower surface of the rocks. Distribution. Callina rotula is endemic to the island of Porto Santo (Madeiran Ar- chipelago, Portugal). It is commonly found in the central-northern areas, north of the line Farrobo-Calhau da Serra de Fora. It is not present on the small islets surrounding the eastern side of the main island, namely Ilhéu de Cima and Ilhéu de Cenouras. See the distributional map in Fig. 214. Taxonomic remarks. Waldén (1983: 273) referred to a fossil specimen resembling C. rotula under his remark 72 from the MMF collection. After checking Waldén's specimen in the collection of the MMF and a recently collected shell, this fossil form apparently belongs to a new species that is described below. Status and conservation. According to Seddon (2011f) the species is considered Least Concern (LC). Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... 111 Figure 214. Distribution of Callina rotula. Filled circles refer to recent and open circles to fossil records. + Callina waldeni Groh & De Mattia, sp. n. http://zoobank.org/033 EFDF1-F522-4652-B259-57C8E313878D Figs 215-217 List of synonyms. 1983 Discula (Callina) rotula — Waldén: 267, note p. 273 [partim]. Type material. MMF 46276, holotype, from loc. typ.; CWDM/1 PT, Porto San- to, E of Vila Baleira, S slope of the hill above Vale do Touro, 50 m W of the oil tanks, excavated Quaternary mixed gravel, 33°03'47"N/16°19'26"W, 24 m, leg. W. De Mat- tia & J. Macor, May 24 2015. Locus typicus. Porto Santo, Quaternary slope deposits in Vale do Touro, Quater- nary mixed gravel, 33°03'51"N/16°19'20"W, 30 m. Diagnosis. Callina species with closed umbilicus and well-rounded rather than angulated or keeled last whorl; granulation arranged along the growth lines and not evenly scattered across the entire shell surface. Description of the holotype. The shell is dextral, very solid, hairless, and discoidal to conical in shape. The protoconch is dark brown, with 1.3 to 1.6 whorls and almost smooth except for very fine radial striae. The teleoconch has from 6.4 to 6.7 rapidly in- creasing whorls. It is whitish in colour, probably bleached. The whorls are slightly con- 112 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) Figures 215-216. Shells of Callina waldeni sp. n. 215 holotype, MMF 46276 216 paratype from E of Vila Baleira, S slope of the hill above Vale do Touro. Scale bar 1 mm. vex. The last whorl is rather rounded, neither distinctly angled nor keeled. ‘The external upper surface has very fine but clearly visible, irregularly spaced, growth lines. Small scattered tubercles are present on the upper surface of the teleoconch. These tubercles are arranged along the growth lines. Small and rather densely set (exclusively along the growth lines) tubercles are also present on the underside of the last whorl. ‘The last whorl is only slightly wider than the penultimate whorl and only slightly descending towards the aperture. The umbilicus is closed. The aperture is more or less elliptical and the peristome is white and very solid. A strong callous is present along the lower palatal side just behind the aperture. The peristome is interrupted along the palatal area and partially reflected along its basal portion. The palatal area does not show any callouses or thickenings. Measurements. D 15.7 + 0.1 mm (range 15.6—-15.8 mm); H 9.1 + 0.3 mm (range 8.9-9.2 mm); FW 5.6 + 0.2 mm (range 5.5—5.7); PA 52.4 + 2.4° (range 51.2—54.4°); NW 7.9 + 0.3 (range 7.7-8.1) (m7 = 2). Ratio D/H 1.7; ratio FW/H 0.6. Distribution. Callina waldeni sp. n. is only known from the locus typicus. See map in Fig. 217. Taxonomic remarks. Waldén (1983: 273) reported in note 72: “A single giant, subfossil specimen (diam. maj. = 15.8 mm, H = 8.9 mm, whorls: 7.9) has been found in Vale do Touro, Porto Santo (Voucher specimen in Funchal Museum)”. Waldén clearly introduced the fossil specimen of the MMF collection (n° 46276) as belonging Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... 113 Figure 217. Distribution of Callina waldeni sp. n. to Callina rotula as also reported in the original museum label depicted in Fig. 215. Callina waldeni sp. n. differs from C. rotula by its larger dimensions and a more conical shape, with slightly more convex whorls. The body whorl is not distinctly angled but rather well rounded. The tubercles are only arranged along the growth lines and not scattered on the entire surface of the shell. The aperture is also less elliptical. Callina waldeni sp. n. is currently the only known fossil representative of the genus. Status and conservation. Extinct before the islands’ scientific exploration in the 19" century, possibly already before human settlement. Callina bulverii (W. Wood, 1828), comb. n. Figs 218-227 List of synonyms. 1828 Helix bulverii W. Wood: 25, pl. 8 fig. 82. 1831 Helix Bulveriana R. 'T. Lowe: 44-45, pl. 5 fig. 11. 1838 Helix rota Potiez & Michaud: 106-107. 1847 Helix Bulweriana — L. Pfeiffer in L. Pfeiffer 1847-1848: 208. 1852 Helix |Discula| Albersii R. T. Lowe: 117. 1853 Helix Bulverii — L. Pfeiffer: 161. 1853 Helix Bulverii var 8 — L. Pfeiffer: 161. 114 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) 1854 Helix Bulveriana — Reeve in Reeve 1851-1854: pl. 136 fig. 849. 1854 Helix | Tectula| Bulwerii — Albers: 24, pl. 4 figs 12+13, 16-18, pl. 5 figs 1-3. 1854 Helix | Tectula| Bulwerii var. y — Albers: 24, pl. 4 figs 16-18. 1854 Helix | Tectula| Bulwerii var. pallidior Albers: 91, pl. 4 figs 14-15. 1854 Helix |Discula] Bulveriana —R.'T. Lowe: 192. 1854 Helix |Discula] Albersii — R.'T. Lowe: 192. 1867 Helix | Tectula| Bulweriana — Paiva: 95. 1867 Helix | Tectula] Bulweriana var. 8 major Paiva: 95. 1888 Helix Bulweri —Tryon in Tryon and Pilsbry 1888: 42, pl. 9 fig. 88. 1894 Geomitra Bulweri — Pilsbry in Pilsbry 1893-1895: 243. 1922 Ochthephila (Tectula) bulverii mut. albescens'T. D. A. Cockerell: 45 [unavailable, infrasubspecific]. 1931 Geomitra (Actinella) Bulweri — Nobre: 100-102, figs. 47, 48. 1950 Discula (Discula) bulweri bulweri — Mandahl-Barth: 469: 36. 1950 Discula (Discula) bulweri albersi — Mandahl|-Barth: 36, pl. 2 fig. 5, pl. 4 fig. 4, pl. 9 fig. 7. 1983 Discula (Discula) bulweri — Waldén: 20: 267. 1983 Discula (Discula) albersi — Waldén: 20: 267. 2002 Discula (Discula) bulverii — Bank et al.: 123. 2008 Discula (Discula) bulverii — Seddon: 78, pl. 29 fig. G, map 170. 2011 Daiscula bulverii — Seddon: e.16725A12800858. Type material. Despite intensive research in multiple museum collections (NHM, NMW, MMUE, ANSP, NHC, NMS, OUMNH, RAM, SMF) that could have held the type material of the taxon, no such material could be traced and there- fore we deem it reasonable to assume that the type material is lost. To stabilise the present interpretation of Helix bulverii WW. Wood, 1828 and to clarify its taxonomic status we designate a neotype here, which is deposited in the collection of the SMF, Frankfurt a. M. under SMF 348938, W. De Mattia & J. Macor, May 15 2014. See Fig. 218. The neotype is consistent with the figure in Wood (1828: pl. 8 fig. 82) in shape, the development of the aperture and the umbilicus and colouration. The taxon was originally described from the Madeiras (= Madeira Archipelago) without more detailed locality data, which is consistent with origin from Porto Santo of the specimen selected as neotype. Loci typici. [bulverii] Porto Santo, S slope of Pico do Facho, along path at the S border of the pine wood, under stones, 33°04'52.47"N/16°19'25.95"W, 355 m a.s.l. (through the designation of the neotype); [Bulveriana] Hab. in montibus Insu- lae Portus S"; [vota] Hab. Porto-Santo (Afrique); [Albersii] Hab. in Portu S*°.; [Bul werii| Habitat in Portosanto, in locis siccis, aridis; [Bu/werii var. y] unknown, as not mentioned; [pallidior| unknown, as not mentioned; [Bulweriana]: Habitat copiose in insulae Portosancti aridis, saxosis sub lapidibus ad montes Pico do Facho, Pico dos Macaricos, prope N.[ossa] Senhora da Graca, Casas Velhas, Cabeco das Fontainhas, fossil ad Zimbral d’Aréa rarior; [Bulweriana var. 8 major]: prope N.[ossa] Senhora da Graga, rarissima; [albescens] Slopes of Pico do Facho, Porto Santo. Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... 1 Further material examined. All from Porto Santo. Fossil: ZMH 110291/5, coast- al slopes at street between harbor and SE coast, 33°03'46"N/16°18'21"W, c. 10-50 m, Sep. 22 1992 and Jul. 1 1996, leg. E. Clauss; ZMH 110292/1, coastal slopes at street from harbor towards the east, 33°03'46"N/16°18'21"W, c. 10-50 m, Sep. 22 1992, leg. E. Clauss. Recent: CWDM/12, S slope of Pico do Facho, along path at the S border of the pine wood, under stones, 33°04'52"N/16°19'26"W, 355 m, leg. W. De Mattia & J. Macor, May 15 2014; CWDM/8, Pico de Juliana, under stones, 33°05'31"N/16°19'20"W, 365 m, leg. K. & C. Groh, Oct. 27 1980; CKG/10, Zimbreiro, 200 m SW of the vil- lage, near the road turn serpentine, under stones, 33°04'16"N/16°18'53"W, 90 m, leg. W. De Mattia & J. Macor, May 2014; CKG/2, summit of smaller hill and saddle be- low the Pico do Magarico, 33°03'58"N/16°18'29"W, approx. 160 m, leg. K. Groh & J. Hemmen, Jul. 2 1983; CKG/2, upper Ribeira do Formosa between Covao and Serra de Fora, 33°05'02"N/16°19'14"W, 140 m, leg. K. & C. Groh & J. & C. Hemmen, Jun. 19 1983; CKG/9, bank of the road above the quarry above the Miradouro da Portela; walls and pasture above Capela da Graca towards the rocky slopes of the Pico do Facho (Motos de Fora), 33°04'42"N/16°20'01"W, 250-400 m, leg. K. & C. Groh & J. & C. Hemmen, Jun. 23 1983; ZMH 120607/4, Pico do Castelo, 33°04'42"N/16°20'01"W to 33°04'51"N/16°20'03"W, 350-430 m, leg. J. & C. Hemmen, ex coll. W. Fauer, Jul. 11 1983; ZMH 120608/2, Graca, 33°04'27"N/16°19'24"W, 190 m, leg. Balka, ex coll. W. Fauer, Jun. 25 1964; CKG/3, Pico Juliana towards Cabo de Grafa, 33°05'34"N/16°19'11" W, 300 m, leg. C. & K. Groh, Aug. 13 1985; CKG/1, Pico do Castelo, 33°04'42"N/16°20'01"W to 33°04'51"N/16°20'03"W, 350-430 m, leg. C. &. K. Groh, Aug. 13 1985; CFW 10782/<10, Casinhas, N of Capela da Graga, 33°04'27"N/16°19'29"W, 150 m, leg. E Walther, Apr. 4 2017; CFW 10783/<10, CMN/3, ZMH 92919/>10 (tissue only), ridge between Zimbreiro and the quarry, 33°04'13"N/16°18'49"W, 110 m, leg. E Walther, Apr. 3 2017; CFW 10847/<10, ZMH 92830/11, Casa Velhas, next to the W edge of the old quarry, 33°04'06"N/16°18'52"W, 125 m, leg. E Walther, Apr. 3 2017; CFW 10918/7, Pico do Concelho, NW slope above water reservoir, 33°04'46"N/16°18'19"W, 170 m, leg. E Walther, Apr. 4 2017; CFW 11126/<10, ZMH 92905/4, 200 m SW of the Zim- breiro near the road turn serpentine, 33°04'15"N/16°18'54"W, 100 m, leg. E Walther, Mar. 31 2017; ZMH 92812/2, Pico do Macarico, small saddle at the NW ridge, leg. 33°03'58"N/16°18'15"W, 220 m, E Walther, Apr. 3 2017; ZMH 24261/2 [as albersi], Porto Santo, without exact locality data, ex coll. Altonaer Museum, ex coll. O. Semper, leg. Wessel; ZMH 24264/2, Porto Santo, without exact locality data, ex coll. Altonaer Museum. Original descriptions. [bu/verii] only name and figure; [Bulveriana]| from Lowe (1831): H. testa rotundato-depressa, hemisphaerica, rotata, supra planulata, acutissime carinata, tenui, nitidiuscula, tota minutissime et confertim granulata, fusco-castanea, supra fasciata; spira convexo-depressa, plus minus elevata, obtusissima; sutura obsoleta; anfractibus planis, acquis, quasi attritis vel confluentibus, ultimi carina acutissima, tenui, supra sulco exarata, limbata; umbilico patulo, spirali, profundo; apertura ro- tundato-lunata; peristomate interrupto, ad umbilicum incrassato, reflexo. Axis 3-212 lin. Diam. 7-8. Anfr. 8-7; [rota] from Potiez & Michaud (1838): Cette coquille est plus petite que lel. lapicida a laquelle elle peut étre comparée par sa forme générale; 116 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) elle est carénée et ombiliquée comme elle, sa couleur est 4 peu prés la méme, mais elle différe par la disposition des taches brunes; le péristome mest ni continu, ni évasé , et l'avant-dernier tour fait une saillie dans ouverture. Toute la coquille est lisse et luisante a la premiére vue, mais a la loupe, elle est finement chagrinée et striée; la caréne est trés tranchante. Hab. Porto-Santo (Afrique.); [Bulverii var. 8] from Pfeiffer (1847): Paulo minor, corneo-lutea , subtiliter marmorata et fasciata, anfractu ultimo subtus convexiore, carina minus compressa; [A/bersii] from R. T. Lowe, 1852: Species eximia inter insigniores, in honorem cl. J. C. Albers, M. D., Helicosophi peritissimi necnon taxophilorum omnium optime meriti dicata. H. Bulveriana similis differt testa submi- nore solidiore trochiformi colore cerino-corneo magis opaco, minus (praesertim jun- iore) planato-depressa, subtus convexiore, carina media minus limbato-prominente, apertura ad carinam haud angulata; [Bulwerii] from Albers 1854: Testa umbilicata, depresse conoidea, solidula, leviter striata et eleganter dense granulate, badia, maculis pallidioribus variegate; spira parum elevate, apice obtuse; anfractus 8 regulariter ac- crescentes, sutura obsolete juncti, ultimus acute limbato-carinatus, subtus planulatus, saturate flavus 1-3 fasciatus, umbilicus mediocris, perspectivus; aperture oblique, el- liptico-rotundata, ad carinam angulata; peristoma simplex, marginibus conniventibus, supero recto, basali crassiori, sinuato, ad umbilicus reflexiusculo. Diam. maj. 18, min. 16 %, al. 8 % millim.; [Bulwerii var. y] from Albers 1854: Testa pallida cornea, striata, sparsim granulate, anfractus ultimus subtus convexus, antice parum descendens, ap- erture subcircularis. Diam maj. 16, min. 15, alt. 7 millim.; [pallidior] from Albers 1854: only name and figure; [Bulveriana var. 8 major| from Paiva 1867: 6 major, spira altissima; [albescens] from Cockerell 1922: Shell greenish-white. Redescription of the shell. The shell is dextral, hairless, and discoidal to tecti- form in shape. The protoconch is whitish to light brown with 2.1 to 2.5 whorls. It is almost smooth along the first whorl and shows fine radial striae along its remain- ing portion. The teleoconch has from 5.2 to 5.6 rapidly increasing whorls. It is horn brown in colour on the upper side, mottled with scattered, dark or light brown small areas. The surface of the shell is somewhat shiny. One chestnut band is present on the upper side of the teleoconch that is gradually widening and becoming darker towards the body whorl. On the underside of the last whorl one main brown band is present. Sometimes, a number of thinner additional bands are present next to the main band. The peri-umbilical area is usually the lightest in colour. The spire is broadly conical, letting the shell appear tectiform. Along the last whorl a single, distinct keel is present which may be slightly bend downwards. ‘The keel is usually lighter in colour than the remaining surface of the whorls, being whitish to light brown. The upper surface is equipped with very fine but clearly visible, rather regularly spaced, growth lines. Small, drop-like tubercles are scattered all over the teleoconch, but concentrated along the growth lines. The last whorl is only slightly wider than the penultimate whorl and is only slightly descending towards the aperture. The umbilicus is widely open (measur- ing approximately 10% of the maximum shell diameter). The aperture is elliptical. The lower palatal side of the last whorl may be equipped with a strong callous just behind the aperture. The peristome is interrupted along the palatal area, and it is only slightly Revision of the genus-group Hystricella R. 7’ Lowe, 1855 from Porto Santo... 117 Figures 218-219. Shells of Callina bulverii. 218 neotype, SMF 348938 219 specimen from Zimbreiro. Scale bar 1 mm. reflected along its basal portion. The palatal area never shows any callosities or thicken- ings. See Fig. 219. Measurements. D 14.2 + 1.0 mm (range 13.3-15.6 mm); H 7.5 + 0.3 mm (range 7.1-7.9 mm); FW 6.2 + 0.2 mm (range 5.9—6.5); PA 48.0 + 7.3° (range 44.6—-49.4°); NT > 100; NW 7.4 + 0.2 (range 7.2—7.6) (” = 30). Ratio D/H 1.9; ratio FW/H 0.8. Body. As in the genus description. Callina bulverii tends to have an overall slightly darker body colouration than C. rotula. Genital anatomy. The albumen gland is long and thin and is connected to an ap- proximately twice as long sperm-oviduct that consists of a prostatic and a uterine por- tion. The prostatic part extends into a thin vas deferens which is approximately twice as long as the sperm-oviduct and which inserts into the penial complex. The distal portion of the uterine part extends into the free oviduct and transforms into a vagina at the level of the duct of the bursa copulatrix. The free oviduct is 7 as long as the vagina. The duct of the bursa copulatrix is very wide, approximately 2 as long as the penial complex and uniform in diameter. It ends into an oval bursa copulatrix. The transition area between the duct and the bursa itself is not sharply delimited but rather gradually widens. The spermatophore is unknown. One tuft of digitiform glands arises from the proximal part of the vagina. There are two to three, approximately equally long and very rarely branched glands. A vaginal appendix arises from the vagina’s wall just distal of the 118 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) Figures 220-226. Genitalia and anatomy of Callina bulverii. S slope of Pico do Facho: 220 whole genitalia excluding part of OSD, AG and gonads 221 ornamentation of the inner walls of the vagina, the penis and the genital atrium 222 cross section of the duct of the bursa copulatrix 223 penial papilla 224 inner walls of epiphallus 225 cross section of penial papilla 226 longitudinal section of penial pa- pilla. Scale bars 1 mm. glandular tuft. Irregularly spaced pleats run longitudinally along the inner surface of the vagina, reaching the genital atrium but extending not as far as the genital orifice. The atrium is short and wide. Its internal walls are smooth. The penial complex consists of a flagellum, an epiphallus, and a penis that inserts into the genital atrium. The penial flagellum is short, cylindrical and has a pointed apex. It is /s as long as the epiphallus. Its internal walls are completely smooth. The epiphallus is twice as long as the penis. Its internal walls are equipped with 20-25 very fine and elevated, longitudinal pleats. The retractor muscle is strong and approximately 2 as long as the epiphallus. The penis is cylindrical, it lacks any muscular or glandular sheath, and it is extremely thick-walled. The internal walls of the penis are smooth. ‘The section where the large penial papilla is located is usually detectable from outside by virtue of a circular swelling corresponding to the origin of the papilla itself. The penial papilla almost reaches the genital atrium and is approximately as long as the penis. It is conical to subcylindrical in shape and has smooth external walls, with the opening emerging apically. The channel of the penial papilla is thin and narrow. The inner lumen of the penial papilla is occupied by Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... 119 Figure 227. Distribution of Callina bulverii. a spongy and sturdy tissue which directly connects with the walls of the epiphallus. The longitudinal section of the penial papilla (Fig. 226) shows that its walls are the continu- ation of the penial walls that abruptly bend inwards. See Figs 220-226. Schileyko (2005: 2018, fig. 2549) depicted the genital anatomy of Callina bulverii, but, except for the penis papilla, only dealt with the outer appearance of the genitalia. Mandahl-Barth (1950) also depicted some anatomical details of C. bulverii, i.e. the foot pedal gland, the digitiform glands, and the vaginal appendix. Jaw and radula. The jaw has been depicted by Mandahl-Barth (1950). Distribution. Callina bulverii is endemic to the island of Porto Santo (Madeiran Archipelago, Portugal). It is restricted to the area delimited by the Pico do Facho, Cal- hau de Serra de Dentro, Portela and Porto dos Frades. It is not present on the small islets surrounding the eastern side of the main island, namely Ilhéu de Cima and Ilhéu de Cenouras. See distribution map in Fig. 227. Taxonomic remarks. Although the shell morphology is rather different from the two other Callina species, the very similar genital anatomy and its position in the cox1 tree clearly support the inclusion of the species in the genus Callina. Seddon (2008: 78) thoroughly summarised the nomenclatural vicissitudes of the synonyms of C. bulverii. Status and conservation. Seddon (2011g) assessed the species as Critical Endan- gered (CR B2a, b(iii, v)) but because of the wide distribution and high frequency the species should be considered Least Concern (LC). 120 Willy De Mattia et al. / ZooKeys 732: 1-125 (2018) Figures 228-232. Comparison of the inner genitalia of Discula (Discula) and Discula (Mandahlia) spe- cies. 228-230 Discula (Mandahlia) tectiformis from summit slopes of Pico do Baixo: 228 cross sections of the penial complex 229 inner walls of penis, atrium and distal vagina 230 inner walls of epiphallus 231-232 Discula (Discula) polymorpha polymorpha from Porto Moniz, Madeira: 231 inner walls of epiphallus, penis, and atrium 232 cross section of penial papilla. Acknowledgements We would like to thank Jon Ablett (NHM) for the search for type material and Phil Crabb (NHM) for photographs of syntypes from the Lowe-Wollaston collection, Ruud Bank (Hoogezand, The Netherlands) for advise on nomenclatural problems, Manuel Biscoito (MMF) for the Portoguese translation of the Abstract and photo- graphs of a Callina-specimen, Paul Callomon (ANSP) and Gary Rosenberg (ANSP) for the loan of material from the Hemmen-collection, Jochen Gerber (Chicago, USA) for the images of specimens from his private collection, Holly Morgenroth (RAM) for the search of type material and photographs of possible Wollaston-types in the Linter collection, Barna Pall-Gergely (HNHM), Takahiro Asami (Shinshu University, Japan) and Josef Harl (Vienna, Austria) for assistance in the lab, sequencing and the prepara- tion and SEM images of jaws and radulae, Ted von Proschwitz (NMG) for the loan of material from the Waldén-collection, Frank Walther (Essen, Germany) for making available parts of his collection for investigation, Harriet Wood (NMW) for the search for type material, the access to the collections and photographs of material from the Melvill-Tomlin collection including possible syntypes from the Wollaston collection, Revision of the genus-group Hystricella R. 7) Lowe, 1855 from Porto Santo... 24 and several people for the search for type material, like Mark Blaxter (NHC, UK), Sankurie Pye (NMS, UK), Mark Carnall (QUMNH, UK), Henry McGhie (MMUE, UK). 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