{"id":213,"date":"2021-05-22T21:43:04","date_gmt":"2021-05-22T21:43:04","guid":{"rendered":"https:\/\/hnords.de\/wordpress\/?page_id=213"},"modified":"2021-12-10T09:15:56","modified_gmt":"2021-12-10T08:15:56","slug":"system-phaedusinae","status":"publish","type":"page","link":"https:\/\/hnords.de\/wordpress\/system-phaedusinae\/","title":{"rendered":"System Phaedusinae"},"content":{"rendered":"

System changes in Phaedusinae<\/strong><\/p>\n

Hartmut Nordsieck (II.2021)<\/p>\n

Introduction<\/strong><\/h4>\n

Two recent papers (Motochin et al. 2017, Mamos et al. 2020), presenting DNA studies of the Phaedusinae, require a revision of the shell-based system of that subfamily used until now (Nordsieck 2007). In parts I and II these papers are reported and commented. In part III the shell-based system is summarized. In part IV the evaluation of morphology is discussed. In part V the conclusions are given.
\nAbbreviation: sbs = of shell-based system.<\/p>\n

I. Motochin et al.<\/h4>\n

Results of gene trees:<\/p>\n

Synprosphymini (Synprosphyma<\/i>) basal.
\nSeparation of tribe of its own (Nordsieck 2007) confirmed.<\/p>\n

Phaedusini:
\n[In brackets representative species. Groups with G type CA = G].<\/p>\n

Groups from Japan and Taiwan (new system used, clades denoted with capitals):
\nA =\u00a0Stereophaedusa<\/i>:
\nBreviphaedusa\u00a0<\/i>(gouldi\u00a0<\/i>=\u00a0Stereophaedusa\u00a0<\/i>sbs);
\nStereophaedusa<\/i>\u00a0(valida<\/i>);
\nMesophaedusa<\/i>\u00a0(hickonis<\/i>,\u00a0japonica<\/i>\u00a0=\u00a0Stereophaedusa\u00a0<\/i>sbs,\u00a0koshikijimana<\/i>\u00a0=\u00a0Pliciphaedusa\u00a0<\/i>sbs);
\nLuchuphaedusa<\/i>\u00a0(callistochila<\/i>,\u00a0bernardii<\/i>\u00a0=\u00a0Nesiophaedusa<\/i>\u00a0sbs);
\nPliciphaedusa<\/i>\u00a0(tosana<\/i>);
\nOophaedusa<\/i>\u00a0(ophidoon<\/i>\u00a0=\u00a0Luchuphaedusa\u00a0<\/i>sbs);
\nunassigned\u00a0neniopsis<\/i>\u00a0=\u00a0Pseudophaedusa\u00a0<\/i>sbs,\u00a0hemileuca<\/i>\u00a0=\u00a0Pliciphaedusa\u00a0<\/i>sbs,\u00a0okimodoki<\/i>\u00a0=\u00a0Mesophaedusa\u00a0<\/i>sbs,\u00a0caloptyx<\/i>\u00a0=\u00a0Hemizaptyx\u00a0<\/i>sbs.
\nB =\u00a0Megalophaedusa<\/i>:
\nPinguiphaedusa<\/i>\u00a0(pinguis<\/i>,\u00a0awajiensis<\/i>\u00a0=\u00a0Placeophaedusa<\/i>\u00a0sbs);
\nVentriphaedusa\u00a0<\/i>(platyderula\u00a0<\/i>=\u00a0Tyrannophaedusa\u00a0<\/i>sbs);
\nVitriphaedusa\u00a0<\/i>(subulina<\/i>,\u00a0hosayaka<\/i>\u00a0=\u00a0Mundiphaedusa\u00a0<\/i>sbs);
\nAulacophaedusa<\/i>\u00a0(gracilispira<\/i>,\u00a0aratorum<\/i>\u00a0=\u00a0Mundiphaedusa\u00a0<\/i>sbs);
\nMegalophaedusa\u00a0<\/i>(martensi<\/i>,\u00a0ducalis<\/i>\u00a0=\u00a0Mundiphaedusa\u00a0<\/i>sbs);
\nDimphaedusa\u00a0<\/i>(decapitata\u00a0<\/i>=\u00a0Mundiphaedusa\u00a0<\/i>sbs);
\nTyrannophaedusa<\/i>\u00a0(mikado<\/i>,\u00a0bilabrata<\/i>);
\nTyrannophaedusoides\u00a0<\/i>(kawamotoi\u00a0<\/i>=\u00a0Tyrannophaedusa\u00a0<\/i>sbs);
\nNeophaedusa<\/i>\u00a0(ishikawai<\/i>\u00a0=\u00a0Mesozaptyx<\/i>\u00a0sbs, other sp. =\u00a0Mundiphaedusa<\/i>\u00a0sbs);
\nPauciphaedusa<\/i>\u00a0(toshiyukii<\/i>\u00a0=\u00a0Mundiphaedusa<\/i>\u00a0sbs);
\nunassigned\u00a0aenea<\/i>\u00a0=\u00a0Pliciphaedusa\u00a0<\/i>sbs,\u00a0rex<\/i>\u00a0=\u00a0Mundiphaedusa\u00a0<\/i>sbs,\u00a0ikenoi<\/i>\u00a0=\u00a0Mesophaedusa\u00a0<\/i>sbs.
\nC =\u00a0Tosaphaedusa<\/i>:
\n(cincticollis<\/i>,\u00a0akiratadai<\/i>\u00a0=\u00a0Mundiphaedusa\u00a0<\/i>sbs).
\nD =\u00a0Zaptyx<\/i>:
\nTaiwanphaedusa<\/i>\u00a0(similaris<\/i>\u00a0=\u00a0Hemizaptyx\u00a0<\/i>sbs);
\nSelenoptyx<\/i>;
\nTyrannozaptyx<\/i>\u00a0(+\u00a0polita<\/i>\u00a0=\u00a0Hemizaptyx\u00a0<\/i>sbs);
\nHemizaptyx<\/i>\u00a0(pinto<\/i>, + probably\u00a0stimpsoni<\/i>\u00a0=\u00a0Hemizaptyx\u00a0<\/i>sbs);
\nParazaptyx<\/i>;
\nPulchraptyx<\/i>;
\nStereozaptyx<\/i>\u00a0(+\u00a0Oligozaptyx<\/i>\u00a0G);
\nZaptyx<\/i>\u00a0G (+\u00a0Metazaptyx<\/i>\u00a0G,\u00a0Diceratoptyx<\/i>\u00a0G);
\nZaptychopsis<\/i>\u00a0G;
\nThaumatoptyx<\/i>\u00a0(+\u00a0kotoshoensis<\/i>\u00a0=\u00a0Zaptyx\u00a0<\/i>sbs, +\u00a0Hemizaptyx<\/i>\u00a0sp. from Taiwan, but without examination);
\nunassigned\u00a0oxypomatica<\/i>\u00a0=\u00a0Heterozaptyx<\/i>,\u00a0asperata<\/i>\u00a0=\u00a0Hemizaptyx\u00a0<\/i>sbs,\u00a0ptychocyma<\/i>\u00a0=\u00a0Hemizaptyx\u00a0<\/i>sbs,\u00a0pseudosheridani<\/i>\u00a0=\u00a0Hemizaptyx\u00a0<\/i>sbs.
\nE =\u00a0Tauphaedusa<\/i>.
\nF =\u00a0Reinia<\/i>\u00a0(+\u00a0Pictophaedusa<\/i>,\u00a0Parareinia<\/i>,\u00a0sieboldtii<\/i>\u00a0=\u00a0Pseudophaedusa\u00a0<\/i>sbs).
\nG =\u00a0Changphaedusa\u00a0<\/i>(horikawai\u00a0<\/i>=\u00a0Pseudophaedusa\u00a0<\/i>sbs).
\nBasal\u00a0Solitariphaedusa<\/i>\u00a0(miyoshii <\/i>=\u00a0Mundiphaedusa\u00a0<\/i>sbs).<\/p>\n

Groups from foreign regions (shell-based system used):
\nBasal to groups from Japan and Taiwan\u00a0Selenophaedusa bavayi<\/i>\u00a0+\u00a0Hemizaptyx minuta<\/i>;
\nClade of remaining groups:
\nbasal\u00a0Atractophaedusa antibouddah<\/i>;
\nSerriphaedusa violacea<\/i>\u00a0=\u00a0serrata<\/i>;
\nFormosana swinhoei<\/i>\u00a0+\u00a0F<\/i>.\u00a0splendens<\/i>;
\nPhaedusa\u00a0<\/i>subclade:
\nPapilliphaedusa porphyrea<\/i>\u00a0+\u00a0Margaritiphaedusa ziyuanensis<\/i>;
\nEuphaedusa aculus\u00a0<\/i>+\u00a0Dentiphaedusa spinula<\/i>;
\nPseudonenia aenigmatica<\/i>\u00a0=\u00a0salacana<\/i>\u00a0(=\u00a0Oospira\u00a0<\/i>sbs) +\u00a0Acrophaedusa<\/i>\u00a0sp. (=\u00a0Oospira\u00a0<\/i>sbs), basal\u00a0Margaritiphaedusa amoena<\/i>\u00a0=\u00a0rusticana<\/i>;
\nFormosanella bensoni<\/i>,\u00a0F<\/i>.\u00a0recedens\u00a0<\/i>+\u00a0Oospira ootayoshinarii<\/i>;
\nPhaedusa dichroa<\/i>,\u00a0P<\/i>.\u00a0sorella<\/i>,\u00a0P<\/i>.\u00a0filicostata<\/i>,\u00a0P<\/i>.\u00a0kelantanensis\u00a0<\/i>+\u00a0Papilliphaedusa kunmingensis<\/i>.<\/p>\n

Morphological characters of following structures considered:
\nShell:
\n1 superior lamella;
\n2 inferior lamella;
\n3 clausilium plate;
\n4 lunellar (mistake: many Phaedusini species have a dorsolateral lunellar!);
\n5 lunella;
\n6 + 7 upper palatal plica;
\n8 middle palatal plica;
\n9 lower palatal plica;
\n10 sutural plica(e);
\n11 shell apex;
\nGenitalia:
\n12 diverticulum of bursa copulatrix.<\/p>\n

Interspecies hybridization:
\nFor\u00a0stereoma<\/i>\u00a0(=\u00a0Stereophaedusa\u00a0<\/i>sbs) and\u00a0cymatodes<\/i>\u00a0(=\u00a0Mesophaedusa\u00a0<\/i>sbs) incongruity between nDNA and mtDNA trees stated: introgression concluded.<\/p>\n

Results:
\nMorphology discordant with phylogeny.
\nPhylogeny concordant with biogeography.<\/p>\n

Comment:<\/p>\n

Gene trees thought to represent one to one phylogenies of taxa. Clades of gene trees regarded as genera; this is an overlumping. Species not belonging to certain clades remain unassigned.<\/p>\n

Imbalance of ranks: In the new system for Japanese and Taiwanese groups former genera downgraded to subgenera, in the system for foreign groups subgenera elevated to genera.<\/p>\n

Characters of shell morphology, which are of decisive importance for species definitions, obviously without any importance for genus definitions. At least difficult to understand.<\/p>\n

Several characters of shell morphology not considered:
\n1 peristome;
\n2 basal keel;
\n3 relationship of spiral lamella and superior lamella;
\n4 subcolumellar lamella;
\n5 lamellulae of G type CA;
\n6 relationship of inner endings of spiral lamella and inferior lamella;
\n7 inner part of subcolumellar lamella.<\/p>\n

Characters of genital morphology (except of presence or absence of diverticulum) not considered.<\/p>\n

So-called diagnoses are simply enumerations of morphological characteristics of the included species or species groups.<\/p>\n

Similar genital organs of Phaedusini make possible interspecies hybridization. Possibility of interspecies hybridization proved by case\u00a0stereoma<\/i>–cymatodes<\/i>. Phaedusini comparable to\u00a0Albinaria<\/i> group of Alopiinae (shell morphology diverse, genital organs similar), in which interspecies hybridization is a frequent phenomenon (see website article on hybridization).
\nPossible past interspecies hybridization in Phaedusini a long time ago not taken into consideration.
\nSeveral remarkable cases of incongruity of shell morphology and gene sequences would be explicable by interspecies hybridization:
\nHemizaptychoids scattered over many clades;
\nstereophaedusoids scattered over several clades,
\nluchuphaedusoids distributed among two clades;
\nstriking similarity of\u00a0Oligozaptyx<\/i>\u00a0with luchuphaedusoids;
\ninclusion of\u00a0sieboldtii<\/i>\u00a0(Pseudophaedusa\u00a0<\/i>sbs) in the\u00a0Reinia<\/i>\u00a0clade.<\/p>\n

II. Mamos et al.<\/h4>\n

Results:<\/p>\n

Synprosphymini (Synprosphyma<\/i>) sister group of Garnieriinae (not well-supported).
\nSee part I.<\/p>\n

Phaedusini (for groups from Japan and Taiwan new system used, for other groups shell-based system, clades denoted with PH):
\n1: Phaedusa <\/i>sp. Timor,\u00a0P<\/i>.\u00a0dichroa<\/i>; Oospira vanbuensis\u00a0<\/i>(Oospira<\/i>),\u00a0Phaedusa paviei<\/i>; Euphaedusa aculus<\/i>,\u00a0Phaedusa lypra<\/i>; Euphaedusa porphyrea<\/i>\u00a0(Papilliphaedusa<\/i>).
\n2: Acanthophaedusa ookuboi<\/i>.
\n3: Hemiphaedusa polydona<\/i>\u00a0(Notoptychia<\/i>),\u00a0H<\/i>.\u00a0billeti<\/i>\u00a0(Selenophaedusa<\/i>).
\n4: Leptacme\u00a0<\/i>(=\u00a0Leptocochlea<\/i>)\u00a0sykesi<\/i>; Liparophaedusa auregani<\/i>,\u00a0Oospira grangeri<\/i>\u00a0(Siphonophaedusa<\/i>); Dautzenbergiella duella<\/i>,\u00a0Oospira miranda\u00a0<\/i>(Oospira<\/i>).
\n5: Phaedusa stenothyra\u00a0<\/i>(Phaedusa<\/i>).
\n6: Oospira antibouddah\u00a0<\/i>(Atractophaedusa<\/i>).
\n7: Oospira swinhoei<\/i>,\u00a0O<\/i>.\u00a0formosensis<\/i>\u00a0(Formosana<\/i>).
\n8: (subclades denoted with J):
\nTauphaedusa tau<\/i>,\u00a0T<\/i>.\u00a0sheridani<\/i>;
\nMessageriella gargominyi<\/i>,\u00a0Zaptyx hachijoensis<\/i>\u00a0(Metazaptyx<\/i>),\u00a0Z<\/i>.\u00a0kikaiensis<\/i>,\u00a0Z<\/i>.\u00a0buschii<\/i>\u00a0(Zaptychopsis<\/i>);
\nReinia ashizuriensis<\/i>,\u00a0R<\/i>.\u00a0variegata<\/i>,\u00a0R<\/i>.\u00a0eastlakeana<\/i>;
\nChangphaedusa horikawai <\/i>(=\u00a0Pseudophaedusa\u00a0<\/i>sbs);
\nStereophaedusa japonica<\/i>,\u00a0hickonis<\/i>\u00a0(=\u00a0Mesophaedusa\u00a0<\/i>sbs),\u00a0jacobiana<\/i>,\u00a0nesiothauma<\/i>\u00a0(=\u00a0Nesiophaedusa\u00a0<\/i>sbs); Megalophaedusa martensi<\/i>,\u00a0M<\/i>.\u00a0hosayaka\u00a0<\/i>(=\u00a0Mundiphaedusa\u00a0<\/i>sbs),\u00a0M<\/i>.\u00a0attrita<\/i>\u00a0(=\u00a0Pinguiphaedusa\u00a0<\/i>sbs),\u00a0M<\/i>.\u00a0pinguis<\/i>\u00a0(=\u00a0Pinguiphaedusa\u00a0<\/i>sbs),\u00a0M<\/i>.\u00a0mikado<\/i>\u00a0(=\u00a0Tyrannophaedusa\u00a0<\/i>sbs),\u00a0M<\/i>.\u00a0iotaptyx<\/i>\u00a0(=\u00a0Tyrannophaedusa\u00a0<\/i>sbs).<\/p>\n

Comment:<\/p>\n

No morphology considered.<\/p>\n

Genus system used: For groups from Japan and Taiwan system resulting from gene trees (former genera downgraded to subgenera, following Motochin et al.), for other groups system based on shell morphology (former genera used, not subgenera, in contrast to Motochin et al.).<\/p>\n

III. System based on shell morphology<\/h4>\n

System of Phaedusini (Nordsieck 2007) based exclusively on shell characters as follows:
\n[Evolution from … to …: genera]
\n1. Shell type:
\nfrom normal = Oospira<\/i>\u00a0type (inferior lamella steeply ascending, clausilium plate narrow) to\u00a0Phaedusa<\/i>\u00a0type (inferior lamella spirally ascending , clausilium plate broad), in connection with ovipary to (ovo)vivipary:
\nmegagenera\u00a0Oospira<\/i>\u00a0and\u00a0Phaedusa \/ Euphaedusa<\/i>.
\n2. Lunellar type:
\nfrom plicae = Oospira<\/i>\u00a0type (free palatal plicae) to lunella =\u00a0Hemiphaedusa<\/i>\u00a0type (instead of plicae lunella):
\nmegagenera\u00a0Oospira<\/i>\u00a0and\u00a0Hemiphaedusa<\/i>.
\nIn all regions from\u00a0Oospira<\/i>\u00a0type
\nto different types with modified plicae in direction to formation of lunella:
\nseveral genera with modified\u00a0Oospira<\/i>\u00a0type.
\nIn Japanese regions from\u00a0Hemiphaedusa<\/i>\u00a0type
\nto\u00a0Tyrannophaedusa<\/i> type (inferior lamella straightly ascending, clausilium plate angular to notched), and
\nto\u00a0Luchuphaedusa<\/i>\u00a0type (inferior lamella spirally ascending, clausilium plate chanelled to notched):
\ngenera\u00a0Tyrannophaedusa<\/i>\u00a0and\u00a0Luchuphaedusa<\/i>.
\nIn Japanese and Taiwanese regions shell becoming smaller to dwarf insular = zaptychoid forms:
\nseveral zaptychoid genera.
\n3. Clausiliar type:
\nfrom normal = N type to special G type of zaptychoids (parallel lamellae and sutural plicae distinct, clausilium plate enlarged into sutural angle):
\nzaptychoid genera with G type.<\/p>\n

IV. Evaluation of morphology<\/h4>\n

Published shell morphology is often incomplete, its knowledge is insufficiently used (see Nordsieck 2021).
\nThe genus taxa have been diagnosed among others by the characters of the inner lamellae endings. These characters have not been considered for the genus taxa out of Japan and Taiwan described in the last decade (except of two); thus, the morphological analysis of these taxa is incomplete. This is of importance, if one tries to classify species with these genus taxa. A good example is\u00a0Oospira ootayoshinarii<\/i>. This species has been classified with nominate\u00a0Oospira<\/i>, but clusters in DNA analysis with\u00a0Formosanella<\/i>\u00a0(see part I). The examination of the inner lamellae endings could give information, where it belongs. Thus, the systematic position of many species described in the last years is uncertain.
\nIt should be considered that the genus taxa out of Japan and Taiwan can include several groups differing in shell characters, which might be not closely related. Atractophaedusa<\/i>\u00a0consists of two groups, the type group with decollate shell and another group with entire shell.\u00a0Formosana<\/i>\u00a0includes at least three groups: type group, group of\u00a0artifina<\/i>, and group of\u00a0moschina<\/i>\u00a0(small dextral species).\u00a0Serriphaedusa<\/i>\u00a0includes two groups, type group and group of\u00a0draesekei<\/i>; two further groups (Gibbophaedusa<\/i>,\u00a0Altiplica)<\/i>\u00a0have recently been added. Within\u00a0Bathyptychia<\/i>\u00a0three subgenera have been described. Selenophaedusa<\/i>\u00a0consists of five groups: type group, groups of\u00a0diplochilus<\/i>,\u00a0billeti<\/i>,\u00a0bavayi,<\/i>\u00a0and\u00a0spinifera<\/i>.\u00a0Margaritiphaedusa\u00a0<\/i>includes two groups: type group and group of\u00a0ziyuanensis<\/i>,\u00a0Euphaedusa<\/i>\u00a0also two groups: type group and group of\u00a0cylindrella<\/i>. One may not be surprised if these genus taxa do not come out in DNA analysis as monophyletic.
\nMost publications on genital morphology do not give measurements and exact subdivisions of the male end ducts. So it must be stated that genital morphology of most species is insufficiently known.<\/p>\n

V. Conclusions<\/h4>\n

From DNA studies (I, II) it is concluded:
\nCharacter changes used for the shell-based system (III) frequently evolved in parallel.
\nThus groups with lunella type (included in megagenus\u00a0Hemiphaedusa<\/i>) and groups with\u00a0Phaedusa<\/i>\u00a0type = (ovo)viviparous groups (included in megagenera\u00a0Phaedusa<\/i>\u00a0and\u00a0Euphaedusa<\/i>) comprise non-related evolutionary units.
\nNot only groups with modified plicae types are different evolutionary units, but also several groups with normal plicae type included in megagenus Oospira<\/i>.
\nZaptychoids did not evolve separately from similar groups with normal shell size (as assumed in Nordsieck 1998), but evolved within one (Zaptyx<\/i>) clade.
\nBut \u2013 morphological characters should be evaluated more thoroughly (IV). This should be considered in future DNA studies.
\nBecause shell characters are often homoplasies and genital characters are similar, higher taxonomic units within Phaedusini can only be characterized by gene sequences. Species or species groups with unknown sequences (at time nearly two thirds of the species, among them the vast majority of species out of Japan and Taiwan) must be provisionally assigned according to morphology and distribution (see Nordsieck 2021).<\/p>\n

References<\/h4>\n

Motochin, R., Wang, M. & Ueshima, R . (2017): Molecular phylogeny, frequent parallel evolution and new system of Japanese clausiliid land snails (Gastropoda: Stylommatophora). \u2013 Zoological Journal of the Linnean Society, 181<\/b>\u00a0(4): 795-845.<\/p>\n

Mamos, T., Uit de Weerd, D., von Oheimb, P. V. & Sulikowska-Drozd, A. (2020): Evolution of reproductive strategies in the species-rich land snail subfamily Phaedusinae (Stylommatophora: Clausiliidae). \u2013 Molecular Phylogenetics and Evolution. https:\/\/doi.org\/10.1016\/j.ympev.2020.107060.<\/p>\n

Nordsieck, H. (1998): Critical revision of the system of the Japanese Phaedusinae, proposed by Minato (1994) (Gastropoda: Stylommatophora: Clausiliidae). \u2013 Archiv f\u00fcr Molluskenkunde,\u00a0127<\/b>\u00a0(1\/2): 21-32.<\/p>\n

Nordsieck, H. (2007): Worldwide Door Snails (Clausiliidae), recent and fossil. \u2013 214 pp., 20 pls. Hackenheim (ConchBooks).<\/p>\n

Nordsieck, H. (2021): Taxonomic important shell characters of Asiatic Phaedusinae (Gastropoda, Stylommatophora, Clausiliidae). \u2013 Acta Conchyliorum, 20<\/strong>: 56 pp., 19 pls.<\/p>\n

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System changes in Phaedusinae Hartmut Nordsieck (II.2021) Introduction Two recent papers (Motochin et al. 2017, Mamos et al. 2020), presenting DNA studies of the Phaedusinae, require a revision of the shell-based system of that subfamily used until now (Nordsieck 2007). In parts I and II these papers are reported and commented. In part III the shell-based system is summarized. In part IV the evaluation of morphology is discussed. In part V the conclusions are given. Abbreviation: sbs = of shell-based…<\/p>\n

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