A misinterpreted disjunction: the phylogenetic relationships of the North African land snail Gyrostomella (Gastropoda: Stylommatophora: Helicidae) Author Neiber, Marco T. Author Korábek, Ondřej Author Glaubrecht, Matthias Author Hausdorf, Bernhard text Zoological Journal of the Linnean Society 2022 2021-11-29 194 4 1236 1251 https://doi.org/10.1093/zoolinnean/zlab059 journal article 55998 10.1093/zoolinnean/zlab059 34d33ce1-7740-4dda-a596-56ee43072231 0024-4082 6459053 3ACD5252-F163-406B-ABCC-6B0F6457AA81 SYSTEMATICS OF THE HELICINAE Our phylogenetic analyses strongly support the monophyletic origin of the Maghreb radiation of Helicinae in the Oligocene in the Maghreb region ( Figs 3 , 4 ). However, our analyses do not provide a robust resolution of the relationships within the Maghreb radiation. Based on these results, we propose to include all groups of the Maghreb radiation (Thebini, Otalini and Ereminini) in a single tribe, for which the oldest available name is Thebini (see Table 1 for an overview of classifications and newly proposed synonyms of the Maghreb radiation of Helicinae based on molecular phylogenetic analyses). Holyoak et al. (2020) proposed subdividing the Maghreb groups into three tribes: Thebini (in the strict sense only including Theba ), Ereminini and Otalini (with a further subdivision into subtribes Otalina and Cantareusina Holyoak et al. , 2020 ). However, Otalini in the sense of Holyoak et al. (2020) is neither statistically supported in their nor in our analyses ( Figs 3 and 4 ). The western Alpine Macularia was weakly joined with the North African Eremina in the analyses of Holyoak et al. (2020) , but was not assigned to any tribe by these authors. Contrary to the analysis of Holyoak et al. (2020) , our analyses strongly support a sister-group relationship of Macularia and the Thebini (in the wide sense). Because of the wide geographic disjunction between the western Alpine Macularia and the primarily Maghrebian Thebini, we suggest a tribe Maculariini trib. nov. [ type genus: Macularia Albers, 1850 ; diagnosis: Helicinae supported by the following apomorphic nucleotide substitutions (based on the alignment deposited in TreeBase under the study accession number 27632): 16S (position 267: T → A; position 283: A → T); only positions considered with data for all taxa and only changes with a consistency index of 1.00 given] to adapt the Linnean systematics of Helicinae to the new findings (ZooBank registration: https://zoobank.org/ urn:lsid:zoobank. org:act: 71BA5313-1FA2-48AE-8A3F-8ED8791EE4EE ). Table 1. Overview of genera assigned to Thebini Wenz, 1923 and Maculariini trib. nov. as proposed here in comparison with previously published classifications based on molecular phylogenetic analyses
Taxon Razkin et al. (2015) Neiber and Hausdorf (2015) Bouaziz- Yahiatene et al. (2017) Bouaziz- Yahiatene et al. (2019) Kneubühler et al. (2019) Hausdorf et al. (2020) Holyoak et al. (2020) This study
Thebini Wenz, 1923 Thebini as Thebini + Otalini Thebini - Thebini Thebini Thebini Thebini
Theba Risso, 1826 Otalini Pfeffer, 1930 Theba Otalini Theba as Thebini + Otalini Theba Otalini - Otalini Theba Otalini Theba Otalini Theba Otalini Theba as Thebini
Otalina Pfeffer, 1930 as Otalini as Thebini + Otalini as Otalini as Otalini as Otalini as Otalini Otalina as Thebini
Otala Schumacher, 1817 Otala - Otala - Otala - Otala Otala
Dupotetia Kobelt, 1904 as subgenus of Otala - - - as Otala - as Otala as Otala
Tingitana Pallary, 1919 Tingitana - as Otala - as Otala - as Otala as Otala
Siretia Pallary, 1926 - - - - Siretia - as Otala as Otala
Loxana Pallary, 1899 as Otala ( Atlasica ) - - - - - Loxana Loxana
Atlasica Pallary, 1917 as Otala ( Atlasica ) - - - - - as Loxana as Loxana
Alabastrina Kobelt, 1904 Alabastrina - - - Alabastrina - as Loxana as Loxana
Maurohelix Hesse, 1917 Maurohelix - - - - - Maurohelix as subgenus of Gyrostomella
Gyrostomella Hesse, 1911 - - - - - - - Gyrostomella
Eobania Hesse, 1913 Eobania Eobania as Massylaea as Massylaea as Massylaea Eobania Eobania Eobania
Massylaea Möllendorff, 1898 - - Massylaea * - - - Massylaea Massylaea
Cantareusina Holyoak et al. 2020 as Otalini as Thebini + Otalini - as Otalini - as Otalini Cantareusina as Thebini
Cantareus Risso, 1826 Cantareus - - Cantareus - Cantareus Cantareus Cantareus
Table 1. Continued
Taxon Razkin et al. Neiber and Bouaziz- Bouaziz- Kneubühler Hausdorf et al. Holyoak et al. This study
(2015) Hausdorf (2015) Yahiatene et al. Yahiatene et al. (2019) (2020) (2020)
(2017) et al. (2019)
Cornu Born, 1778 Cornu Cornu - Cornu - Cornu Cornu Cornu
Erctella Erctella - - Erctella - as subgenus of Erctella as subgenus of
Monterosato, Cornu Cornu
1894
Rossmaessleria Rossmaessleria Rossmaessleria - - - Rossmaessleria Rossmaessleria Rossmaessleria
Hesse, 1907
Ereminini as Helicini† as Thebini + Otalini - - - as Otalini Ereminini as Thebini
Holyoak et al.
2020
Eremina Pfeiffer , Eremina Eremina - - - Eremina Eremina Eremina
1855
Maculariini - as Thebini + Otalini - - - as Otalini incertae sedis Maculariini
trib. nov. trib. nov.
Macularia Albers , - Macularia - - - Macularia Macularia Macularia
1850
*According to Holyoak et al. (2020) the specimen determined as Massylaea massylaea (Morelet, 1851) by Bouaziz-Yahiatene et al. (2017) is a misidentified Eobania constantina (Forbes, 1838) . † This placement was based on a misidentified sequence from GenBank attributed to Eremina desertorum (Forsskål, 1775) ( AY841335 ; in fact Levantina ). This proposed classification closely mirrors the biogeographic history of the Helicinae . The Eastern Mediterranean Helicini diverged from the Western Mediterranean clade in the Eocene ( Fig. 4 ). In the Late Eocene or Early Oligocene, the Western Mediterranean clade split into Allognathini in the Iberian Peninsula, Maculariini in the region of the future western Alps and Thebini on terranes probably including the Rif-Betic and Kabylies blocks that were at that time adjacent to south-western Europe, but later drifted south-westwards to form part of the Maghreb region ( Rosenbaum et al. , 2002 ). Further phylogenetic investigations are necessary to provide a better resolution within the tribes as a basis for more detailed biogeographic reconstructions.