The mostly cavernicolous millipede genus Stygiiulus Verhoeff, 1929, stat. nov.: taxonomy, distribution and phylogenetic relationships (Diplopoda, Julida, Julidae)

Keywords: Typhloiulus, new combination, new synonymy, lectotype designation, modified mouthparts

Abstract

The blind and mostly endogean julid genus Stygiiulus Verhoeff, 1929 stat. nov. is reviewed to include ten species: S. ausugi (Manfredi, 1953) comb. nov., S. fimbriatus (Strasser, 1971) comb. et stat. nov., S. gentianae (Strasser, 1971) comb. et stat. nov., S. illyricus (Verhoeff, 1929) comb. nov., S. insularis (Strasser, 1938) comb. nov., S. maximus (Verhoeff, 1929) comb. nov., S. montellensis (Verhoeff, 1930) comb. nov., S. rotundatus (Strasser, 1962) comb. et stat. nov., S. seewaldi (Strasser, 1967) comb. nov., and S. tobias (Berlese, 1886) comb. nov. The distinctiveness of the genus from Typhloiulus is proven by both morphological and molecular data. The monotypic genus Alpityphlus Strasser, 1967 is here treated as a junior subjective synonym of Stygiiulus syn. nov. Stygiiulus fimbriatus comb. et stat. nov. and S. gentianae comb. et stat. nov., originally described as subspecies of S. ausugi comb. nov., are here erected to full species, while the subspecies S. illyricus stygis (Verhoeff, 1933) comb. et syn. nov. is considered to be a junior subjective synonym of the typical S. illyricus comb. nov. Lectotypes are formally designated for S. gentianae comb. et stat. nov. and S. rotundatus comb. et stat. nov., to stabilize the nomenclature of the two species under Article 74.1 of the ICZN. The phylogeny of the genus, its distribution patterns, and the modified mouthparts in some of its species are discussed.

References

Antić D.Ž. & Reip H.S. 2020. The millipede genus Leucogeorgia Verhoeff, 1930 in the Caucasus, with descriptions of eleven new species, erection of a new monotypic genus and notes on the tribe Leucogeorgiini (Diplopoda: Julida: Julidae). European Journal of Taxonomy 713: 1–106. https://doi.org/10.5852/ejt.2020.713

Antić D.Ž., Dudić B.D., Gajić M.R. & Lučić L.R. 2017. The first hydrophilous cave-dwelling millipede from Serbia – Typhloiulus balcanicus sp. nov. (Diplopoda, Julida, Julidae). Zootaxa 4226 (1): 137–143. https://doi.org/10.11646/zootaxa.4226.1.8

Antić D.Ž., Dražina T., Rađa T., Lučić L.R & Makarov S.E. 2018. Review of the genus Typhloiulus Latzel, 1884 in the Dinaric region, with a description of four new species and the first description of the male of Typhloiulus insularis Strasser, 1938 (Diplopoda: Julida: Julidae). Zootaxa 4455 (2): 258–294. https://doi.org/10.11646/zootaxa.4455.2.2

Attems C. 1927. Über palaearktische Diplopoden. Archiv für Naturgeschichte 92 (1–2): 1–256.

Attems C. 1949. Die Myriopodenfauna der Ostalpen. Sitzungsberichte, Österreichische Akademie der Wissenschaften, Mathematisch-Naturwissenschaftliche Klasse, Abteilung I 158 (1–2): 79–153.

Boldori L. 1936. Ricerche in caverne italiane (IV serie 1934–1935). Natura XXVII: 106–114.

Boldori L. 1937. Fauna cavernicola. In: Trevisiol G. & Boldori L. (eds) “Grotte del Vicentino”: 10–12. Gruppo Grotte C.A.I., Vicenza.

Canciani G. 2019. La fauna cavernicola del Foran di Landri. In: A.S. Forum Julii Speleo (ed.) Foran di Landri. Il Landri svelato: ricerche e approfondimenti su una grotta tra storia e folklore: 65–79.

Cognato A.I. & Vogler A.P. 2001. Exploring data interaction and nucleotide alignment in a multiple gene analysis of Ips (Coleoptera: Scolytinae). Systematic Biology 50 (6): 758–780. https://doi.org/10.1080/106351501753462803

Conci C. 1951. Contributo alla conoscenza della speleo-fauna della Venezia Tridentina. Memorie della Società Entomologica Italiana, XXX: 5–76.

Enghoff H. 1976. Parthenogenesis and bisexuality in the millipede, Nemasoma varicorne C. L. Koch, 1847 (Diplopoda: Blaniulidae). Morphological, ecological, and biogeographical aspects. Videnskabelige meddelelser fra Dansk Naturhistorisk Forening 139: 21–59.

Enghoff H. 1985. Modified mouthparts in hydrophilous cave millipedes (Diplopoda). In: Ellis W.N., Jeekel C.A.W. & Pieters F.F.J.M. (eds) Proceedings of the 6th International Congress of Myriapodology. Amsterdam, 12–17 April 1984. Bijdragen tot de Dierkunde 55 (1): 67–77.

Enghoff H. 1996. The penis as a phylogenetic character in the millipede family Julidae. In: Geoffroy J.J., Mauriès J.-P. & Nguyen Duy-Jacquemin M. (eds) Acta Myriapodologica. Mémoires du Muséum national d'histoire naturelle, N.S. 169: 313-–326.

Enghoff H., Petersen G. & Seberg O. 2011. Phylogenetic relationships in the millipede family Julidae. Cladistics 27 (6): 606–616. https://doi.org/10.1111/j.1096-0031.2011.00360.x

Enghoff H., Petersen G. & Seberg O. 2013. The aberrant millipede genus Pteridoiulus and its position in a revised molecular phylogeny of the family Julidae (Diplopoda: Julida). Invertebrate Systematics 27: 515–529. http://doi.org/10.1071/IS13016

Evsyukov A., Golovatch S. & Reip H.S. 2018. The millipede genus Julus Linnaeus, 1758 in the Caucasus (Diplopoda: Julida: Julidae). Zootaxa 4461 (1): 089–117. https://doi.org/10.11646/zootaxa.4461.1.7

Fritsch E. 1998. Die Höhlenfauna des Hirlatzmassivs. In: Buchegger G. & Greger W. (eds) Die Hirlatzhöhle im Dachstein 52: 136–192. ARGE Hirlatzhöhlenbuch, Hallstadt, Germany.

Golovatch S.I. 2009. Millipedes (Diplopoda) in extreme environments. In: Golovatch S.I., Makarova O.L., Babenko A.B. & Penev L.D. (eds) Species and Communities in Extreme Environments, Festschrift towards the 75th Anniversary and a Laudatio in Honour of Academician Yuri Ivanovich Chernov: 87–112. Pensoft Publishers & KMK Scientific Press, Sofia, Moscow.

Kime R.D. & Enghoff H. 2017. Atlas of European millipedes 2: Order Julida (Class Diplopoda). European Journal of Taxonomy 346: 1–299. https://doi.org/10.5852/ejt.2017.346

Kumar S., Stecher G., Li M., Knyaz C. & Tamura K. 2018. MEGA X: Molecular evolutionary genetics analysis across computing platforms. Molecular Biology and Evolution 35 (6): 1547–1549. https://doi.org/10.1093/molbev/msy096

Kutty S.N., Bernasconi M.V., Šifner F. & Meier R. 2007. Sensitivity analysis, molecular systematics and natural history evolution of Scathophagidae (Diptera: Cyclorrhapha: Calyptratae). Cladistics 23 (1): 64–83. https://doi.org/10.1111/j.1096-0031.2006.00131.x

Lanfear R., Frandsen P.B., Wright A.M., Senfeld T. & Calcott B. 2017. PartitionFinder 2: new methods for selecting partitioned models of evolution for molecular and morphological phylogenetic analyses. Molecular Biology and Evolution 34 (3): 772–773. https://doi.org/10.1093/molbev/msw260

Latzel R. 1884. Die Myriopoden der österreichisch-ungarischen Monarchie. Zweite Hälfte. Die Symphylen, Pauropoden und Diplopoden. Alfred Hölder, Wien.

Liu W., Golovatch S., Wesener T. & Tian, M. 2017. Convergent evolution of unique morphological adaptations to a subterranean environment in cave millipedes (Diplopoda). PLoS ONE 12 (2): 1–20. https://doi.org/10.1371/journal.pone.0170717

Makarov S.E., Bodner M., Reineke D., Vujisić Lj.V., Todosijević M.M., Antić D.Ž., Vagalinski B., Lučić L.R., Mitić B.M., Mitov P., Anđelković B.D., Pavković Lučić S., Vajs V., Tomić V.T. & Raspotnig G. 2017. Chemical ecology of cave-dwelling millipedes: defensive secretions of the Typhloiulini (Diplopoda, Julida, Julidae). Journal of Chemical Ecology 43 (4): 317–326. https://doi.org/10.1007/s10886-017-0832-1

Manfredi P. 1932. Contributo alla conoscenza della fauna cavernicola italiana. Miriapodi. Rivista di Scienze Naturali “Natura” 23: 71–96.

Manfredi P. 1953a. Nuovi miriapodi cavernicoli del Trentino. Studi Trentini di Scienze Naturali 30: 136–139.

Manfredi P. 1953b. Miriapodi italiani – VIII. contributo. Atti della Società italiana di Scienze naturali, e del Museo civile di Storia naturale 92: 117–122.

Miller M.A., Pfeiffer W. & Schwartz T. 2011. The CIPRES science gateway: a community resource for phylogenetic analyses. In: Proceedings of the 2011 TeraGrid Conference: Extreme Digital Discovery: 1–8. Association for Computing Machinery, New York. https://doi.org/10.1145/2016741.2016785

Minelli A. 1985. Catalogo dei Diplopodi e dei Chilopodi cavernicoli italiani. Memorie del Museo civico di Storia naturale di Verona, Serie 2 (Sezione scienze della Vita) 4: 1–50.

Muraji M. & Tachikawa S. 2000. Phylogenetic analysis of water striders (Hemiptera: Gerroidea) based on partial sequences of mitochondrial and nuclear ribosomal RNA genes. Entomological Science 3 (4): 615–626.

Ozimec R., Bedek J., Gottstein S., Jalžić B., Slapnik R., Štamol V., Bilandžija H., Dražina T., Kletečki E., Komerički A., Lukić M. & Pavlek M. 2009. Crvena knjiga špiljske faune Hrvatske. [Red book of Croatian cave dwelling fauna]. In: Ozimec R. & Katušić L. (eds) Ministarstvo kulture. Državni zavod za zaštitu prirode, Zagreb.

Paoletti M.G. 1978. Cenni sulla fauna ipogea delle Prealpi Bellunesi e Colli Subalpini. Le Grotte d’Italia 4 (7): 45–198.

Pretner E. & Strasser K. 1931. Die Fauna der nordfriauler Höhlen. Mitteilungen über Höhlen- und Karstforschung 1931 (3): 84–90.

Rambaut A., Drummond A.J., Xie D., Baele G. & Suchard M.A. 2018. Posterior summarization in Bayesian phylogenetics using Tracer 1.7. Systematic biology 67 (5): 901–904. https://doi.org/10.1093/sysbio/syy032

Ronquist F., Teslenko M., Van Der Mark P., Ayres D.L., Darling A., Höhna S., Larget B., Liu L., Suchard M.A. & Huelsenbeck J.P. 2012. MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61 (3): 539–542. https://doi.org/10.1093/sysbio/sys029

Simon C., Frati F., Beckenbach A., Crespi B., Liu H. & Flook P. 1994. Evolution, weighting, and phylogenetic utility of mitochondrial gene sequences and a compilation of conserved polymerase chain reaction primers. Annals of the Entomological Society of America 87 (6): 651–701. https://doi.org/10.1093/aesa/87.6.651

Stamatakis A. 2014. RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics 30 (9): 1312–1313. https://doi.org/10.1093/bioinformatics/btu033

Strasser K. 1938. Neue Diplopoden der Insel Cherso. Zoologische Jahrbücher, Abteilung für Systematik, Ökologie und Geographie der Tiere 71 (4–6): 389–407.

Strasser K. 1962. Die Typhloiulini (Diplopoda Symphyognatha). Atti del Museo civico di Storia naturale di Trieste 23 (1): 1–77.

Strasser K. 1967. Ein Typhloiuline aus den nördlichen Kalkalpen (Diplopoda Symphyognatha). Berichte des naturwissenschaftlich-medizinischen Vereins in Innsbruck 55: 145–153.

Strasser K. 1971a. Über italienische, besonders kavernikole Diplopoden. Memorie del Museo civico di Storia naturale di Verona 19: 1–21.

Strasser K. 1971b. Über Diplopoden Jugoslawiens. Senckenbergiana biologica 53: 313–345.

Vagalinski B. & Lazányi E. 2018. Revision of the millipede tribe Brachyiulini Verhoeff, 1909 (Diplopoda: Julida: Julidae), with descriptions of new taxa. Zootaxa 4421 (1): 001–142. https://doi.org/10.11646/zootaxa.4421.1.1

Vagalinski B. 2021. Rhodopotyphlus mitovi gen. et sp. nov., a new endogean millipede from the West Rhodope Mountains, Bulgaria (Diplopoda: Julida: Julidae). Zootaxa 5071: 415–426. https://doi.org/10.11646/zootaxa.5071.3.7

Vagalinski B., Stoev P. & Enghoff H. 2015. A review of the millipede genus Typhloiulus Latzel, 1884 (Diplopoda: Julida: Julidae), with a description of three new species from Bulgaria and Greece. Zootaxa 3999 (3): 334–362. https://doi.org/10.11646/zootaxa.3999.3.2

Verhoeff K. 1929. Arthropoden aus südostalpinen Höhlen (I). Gesammelt von Karl Strasser in Triest. Mitteilungen über Höhlen- und Karstforschung 1929: 14–35.

Verhoeff K. 1930. Arthropoden aus südostalpinen Höhlen, gesammelt von Karl Strasser. 5. Aufsatz. Mitteilungen über Höhlen- und Karstforschung 1930 (4): 97–113.

Verhoeff K.W. 1933. Arthropoden aus südostalpinen Höhlen, gesammelt von Karl Strasser. 7. Aufsatz. Mitteilungen über Höhlen- und Karstforschung 1933 (4): 1–21.

Wolf B. 1934–38. Animalium Cavernarus Catalogus. ‘sGravenhage, W. Junk, Vol. 3.

Xia X. 2018. DAMBE7: New and improved tools for data analysis in molecular biology and evolution. Molecular Biology and Evolution 35 (6): 1550–1552. https://doi.org/10.1093/molbev/msy073

Published
2022-02-28
How to Cite
Vagalinski, B., Borissov, S., Bobeva, A., Canciani, G., & Antić, D. Ž. (2022). The mostly cavernicolous millipede genus Stygiiulus Verhoeff, 1929, stat. nov.: taxonomy, distribution and phylogenetic relationships (Diplopoda, Julida, Julidae). European Journal of Taxonomy, 798(1), 30-69. https://doi.org/10.5852/ejt.2022.798.1669
Section
Research article