identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
03AD87E5FF82FF8A2D1AFAD2A9B8DA7A.text	03AD87E5FF82FF8A2D1AFAD2A9B8DA7A.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Vitrinidae Fitzinger 1833	<div><p>CHARACTERS</p><p>The characters of the genitalia are shown in Figure 2. The numbering refers to Table 1 and the cladogram (Fig. 3).</p><p>1 The so-called stimulator, which is apparently an adaptation to reciprocal copulation, is one of the most important characters for the systematics of the  Stylommatophora (Hausdorf 1998a) . The glandula amatoria is considered to be homologous to the stimulator. A stimulator is present in most vitrinids except  Vitrina,  Calidivitrina and some species of  Plutonia, namely  P. media (Lowe, 1854) (see Appendix) and  P. brevispira (Morelet, 1860) (Mordan &amp; Martins, 2001) .</p><p>2 In the outgroups,  Semilimax and  Vitrinobrachium, the stimulator inserts at the atrium. The stimulator of  Semilimacella cephalonica (Rähle) is basally fused with the vagina (Rähle, 1980). In  Semilimacella carniolica (O. Boettger) the stimulator apparently inserts at the vagina (Mildner, 1982) and in  Semilimacella bonelli (Targioni Tozzetti) it inserts at the atrium (Forcart, 1960; re-examined).  Semilimacella carniolica and  Semilimacella bonelli are probably sister species. Both species are characterized by duplicated stimulator papillae. In  Semilimacella cephalonica the gland of the stimulator is bilobed, but there is only a single stimulator papilla. It is assumed that the stimulator of  Semilimacella cephalonica represents the plesiomorphic state in  Semilimacella . The situation in  Semilimacella carniolica can easily be derived from that state by a further fusion of the distal sections of vagina and stimulator. If this is true, the atrial insertion of the stimulator of  Semilimacella bonelli must be due to a shortening of the fused section and is therefore a secondarily derived state. On the basis of this interpretation, the stimulator of  Semilimacella might represent an intermediate state between the atrial stimulator and the glandula amatoria, where the stimulator is completely fused with the vagina. In  Vitrina there is a vestige of a stimulator at the atrium of juvenile specimens (Umiński, 1968). However, it cannot be excluded that this is a rudiment of a stimulator which was basally fused with the vagina, because the vagina is missing in  Vitrina . In  Eucobresia,  Phenacolimax,  Plutonia,  Oligolimax and  Arabivitrina the stimulator is completely fused with the vagina and forms the glandula amatoria.</p><p>3  Eucobresia and some  Plutonia and  Oligolimax species have no external stimulator gland.</p><p>4 The external gland of the stimulator is distinctly longer than it is wide in the outgroups and in  Semilimax, whereas it is about as long as or shorther than it is wide in the other vitrinids.</p><p>5 The external gland of the stimulator forms a uniform sheath around the proximal stimulator section in the outgroups and in  Semilimax,  Vitrinobrachium,  Arabivitrina, and some  Plutonia species. In the other vitrinids, it is divided into lobes.</p><p>6 In  Semilimax and  Vitrinobrachium the stimulator is used to fix the copulation partner (Künkel, 1933), whereas the stimulator contacts the partner only temporarily during copulation in the other  Stylommatophora for which the mating behaviour is known. 7 The penis is inserted into the vagina or the bursa copulatrix of the copulation partner in most  Stylommatophora: however, it transfers the sperm on external parts of the everted genitalia in  Vitrinobrachium,  Phenacolimax,  Plutonia and, probably, all other genera with a well-developed glandula amatoria.</p><p>8 A penial tunica is present in the outgroups and in most genera of the  Vitrinidae . It is missing in  Plutonia,  Oligolimax,  Arabivitrina and  Calidivitrina . 9 The penial tunica usually covers most of the distal part of the penis, except in  Eucobresia and  Phenacolimax .</p><p>10 There is at least one distinct pilaster in the penis of the vitrinids which is partly formed by the penial gland. In  Arabivitrina and  Calidivitrina the pilaster is partitioned into a proximal glandular section, a short median section with a distinctly lamellated structure (Neubert, 1998: figs 97 and 106), and an undifferentiated distal section. The penial pilaster of the two Azorean  Plutonia species P. pelagica (Morelet, 1860) and  P. laxata (Morelet, 1860) is also partitioned into a proximal glandular section and a median section with a distinctly lamellated structure (Mordan &amp; Martins, 2001). However, in contrast to the pilaster of  Arabivitrina and  Calidivitrina, the distal section is divided into two branches. Nevertheless, it should be checked more thoroughly whether the lamellated median section of  Arabivitrina and  Calidivitrina might be homologous to that of the two Azorean  Plutonia species.</p><p>11 A more or less well-developed penial gland is present in all vitrinids, but is missing in the outgroups. The secretions of this gland are probably involved in sperm transfer and thus undertake some of the functions of the spermatophore, which is missing in the  Vitrinidae .</p><p>12 The right ommatophoral retractor muscle passes between the penis and the female genitalia in  Cryptozona,  Eucobresia,  Phenacolimax,  Arabivitrina and some  Semilimax and  Calidivitrina species.</p><p>13 The right ommatophoral retractor muscle runs left of or below the penis retractor muscle in  Troglaegopis,  Semilimacella,  Vitrina and  Oligolimax, but it runs right of or above the penis retractor muscle in all other vitrinids.</p><p>14 The penial retractor inserts at the diaphragma in the outgroups and in most groups of the  Vitrinidae . It inserts at the columellar muscle only in  Vitrinobrachium .</p><p>15 The vas deferens enters the penis terminally in  Semilimacella,  Vitrina, and  Plutonia finitima (Morelet, 1860), whereas it inserts at the penis subterminally or laterally in all other groups of the  Vitrinidae .</p><p>16 The bursa copulatrix inserts at the vagina or at the atrium in the outgroups and in all groups of the  Vitrinidae except in  Vitrinobrachium in which it inserts at the penis.</p><p>17 The radular marginals are unicuspid or have at most one ectoconus in the outgroups and in most  Vitrinidae . They are multicuspid in  Vitrina,  Semilimacella and in a few species of  Plutonia and  Oligolimax .</p><p>PHYLOGENY</p><p>Two equally and maximally parsimonious trees (length 25 steps, consistency index excluding uninformative characters = 0.636) have been found in a branch-and-bound search with the program PAUP and the character matrix (Table 1). The two trees differ only in the relationships of  Semilimax and  Vitrinobrachium . A consensus tree of the two trees is shown in Figure 3.</p><p>The unresolved trichotomy is due to two conflicting characters, the function of the stimulator vs. the length of the stimulator gland. Among the  Stylommatophora for which the mating behaviour has been studied,  Semilimax and  Vitrinobrachium are the only ones which use the stimulator to fix the copulation partner (Künkel, 1933). In the other  Stylommatophora the stimulator contacts the mating partner only temporarily during copulation. Therefore the use of the stimulator to fix the mating partner might be a synapomorphy of  Semilimax and  Vitrinobrachium . Unfortunately, the mating behaviour of  Troglaegopis (Zonitidae) (the stimulator of which resembles that of  Semilimax) as well as the mating behaviour of several other vitrinids, e.g.  Semilimacella, is unknown. Therefore, it cannot be excluded that the unusual function of the stimulator is an autapomorphy of the  Vitrinidae or originated even earlier.</p><p>If this insufficiently known character is excluded from the cladistic analysis only a single mostparsimonious tree is found, in which  Semilimax is the sister group of the remaining vitrinids, as in one of the two maximally parsimonious trees based on all characters. The clade including all vitrinids except  Semilimax is supported by the shortening of the stimulator gland.</p><p>The result of the present cladistic analysis differs fundamentally from the hypothetical scheme proposed by Schileyko (1986). The differences are primarily due to the assumption of Schileyko (1986) that the glandula amatoria is the plesiomorphic character state of the stimulator. This assumption is based on the presupposed homology of the stimulator of the  Vitrinidae with the capsular gland of other Limacoidea s. l.. However, Hausdorf (1998a) has shown that the capsular gland is not homologous with the stimulator. An outgroup comparison with the stimulator of the  Zonitidae or the Helicarionoidea demonstrates that the atrial stimulator represents the plesiomorphic character state and that the glandula amatoria is apomorphic.</p><p>Moreover, Schileyko (1986) ignored the findings of Umiński (1968), who discovered a rudiment of an atrial stimulator in  Vitrina . Therefore, Schileyko (1986) derived  Vitrina directly from his hypothetical stem form, which has only a capsular gland but no stimulator. Actually, the stem species of the  Vitrinidae did not have a capsular gland, because this organ is missing in all vitrinids as well as in their sister group, the  Boettgerillidae –  Limacidae –  Agriolimacidae . Schileyko (1986) overlooked the fact that  Vitrina only differs from  Semilimacella in that the atrial stimulator is reduced.</p><p>Finally, Schileyko (1986) did not realize the homology of the vaginal papilla of  Eucobresia with the papilla of the glandula amatoria. This homology is corroborated by the fact that there is a well-developed muscular section of the stimulator in  Eucobresia glacialis (Forbes), which has to be included into  Eucobresia (see Appendix). Moreover, the relationship of  Eucobresia to  Phenacolimax and the other groups with a glandula amatoria is corroborated by the development of the penial tunica, which surrounds only the proximal section of the penis in  Eucobresia and  Phenacolimax, whereas it also surrounds parts of the distal section in all groups with an atrial stimulator.</p><p>The subdivision of the  Vitrinidae proposed by Schileyko (1986) cannot be maintained. The Vitrininae Fitzinger, 1833 sensu Schileyko include the groups in which the stimulator has been lost, namely  Vitrina and  Calidivitrina, and are polyphyletic. The  Semilimacinae Schileyko, 1986 include the groups with an atrial stimulator,  Semilimacella,  Semilimax and  Vitrinobrachium, as well as  Eucobresia and are paraphyletic. Only the  Phenacolimacinae Schileyko, 1986; which are characterized by the glandula amatoria are monophyletic. However,  Plutoniinae Cockerell, 1893 is an older name for this group. Because of the low number of genera, a formal division of the  Vitrinidae into subfamilies is not necessary.</p><p>BIOGEOGRAPHY</p><p>The distribution of the genera of the  Vitrinidae is summarized in Table 2 and Figure 4. The range of the  Vitrinidae largely overlaps with the range of its sister group, the limacoid slugs  Boettgerillidae –  Limacidae –  Agriolimacidae . If one assumes that the ancestors of the two sister groups originated by allopatric speciation, their original ranges were smaller than those of the two sister groups are today. For the estimation of the ancestral area of the  Vitrinidae and of the limacoid slugs a weighted ancestral area analysis (Hausdorf, 1998b) has been applied.</p><p>The PI values of the  Vitrinidae (Table 2), which indicate the relative probability that an individual area was part of the ancestral area, are maximal for the Alps followed by West and Central Europe. The PI value of the  Vitrinidae for the Near East (including the Caucasus region) is distinctly lower.</p><p>The PI values of the limacoid slugs  Boettgerillidae –  Limacidae –  Agriolimacidae (Table 3) are maximal for the Near East, because the positionally plesiomorphic lineages, namely the  Boettgerillidae, Eumilacinae and Mesolimacinae are restricted to that area. Therefore, this region is probably the ancestral area of the  Boettgerillidae –  Limacidae –  Agriolimacidae . Consequently, the  Vitrinidae and the  Boettgerillidae –  Limacidae –  Agriolimacidae might have originated by a vicariance event between Central Europe and the Near East.</p><p>Some lineages of the  Vitrinidae spread from the European mainland.  Vitrina dispersed into Asia and North America (and one species even reached Hawaii),  Oligolimax also dispersed into Asia and North Africa,  Plutonia colonized the Macaronesian Islands (Azores, Madeira, Canary Islands and Cap Verde) and  Arabivitrina and  Calidivitrina colonized Arabia and East Africa.</p><p>Concerning species number, the most important radiation of the  Vitrinidae occurred on the Macaronesian Islands. There are more species on the Macaronesian Islands than on the European mainland. The apomorphic position of  Plutonia in the cladogram of the  Vitrinidae (Fig. 3) and the uniform bauplan of the genitalia of the Macaronesian species indicate that this radiation is younger than the radiation on the European mainland. On the other hand, the Macaronesian radiation resulted in some of the most extreme forms concerning the body bauplan, namely the only vitrinid slug,  Plutonia (Plutonia) atlantica (Morelet 1860), and  Plutonia (Guerrina), which can entirely withdraw into their ribbed and keeled shells. The ecology of the Macaronesian vitrinids differs from that of the European vitrinids. Whereas the highest diversity of European vitrinids can be found above 1000 m altitude, the highest diversity of vitrinids is below 500 m altitude on the Canary Islands. These biogeographical and ecological patterns are very unusual. The reasons for these patterns have been discussed elsewhere (Hausdorf, 2001).</p></div>	https://treatment.plazi.org/id/03AD87E5FF82FF8A2D1AFAD2A9B8DA7A	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Hausdorf, Bernhard	Hausdorf, Bernhard (2002): Phylogeny and biogeography of the Vitrinidae (Gastropoda: Stylommatophora). Zoological Journal of the Linnean Society 134 (3): 347-358, DOI: 10.1046/j.1096-3642.2002.00010.x, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1046/j.1096-3642.2002.00010.x
