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Limnephilid Taxa Revised by Speciation Traits: Rhadicoleptus, Isogamus, (Trichoptera: Limnephilidae)

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Limnephilid Taxa Revised by Speciation Traits: Rhadicoleptus, Isogamus, (Trichoptera: Limnephilidae) Opusc. Zool. Budapest, 2015, 46(1): 3–117 Limnephilid taxa revised by speciation traits: Rhadicoleptus, Isogamus, Melampophylax genera, Chaetopteryx rugulosa, Psilopteryx psorosa species groups, Drusus bolivari, Annitella kosciuszkii species complexes (Trichoptera: Limnephilidae)* 1 2 3 4,5 6 7 7 J. OLÁH , P. CHVOJKA , G. COPPA , R. J. GODUNKO , O. LODOVICI , K. MAJECKA , J. MAJECKI , 8 9,10 11 B. SZCZĘSNY , G. URBANIC & M. VALLE 1János Oláh, residence postal address: Tarján u. 28, H-4032 Debrecen, Hungary. E-mail: [email protected] 2Pavel Chvojka, National Museum, Natural History Museum, Department of Entomology, Cirkusová 1740, CZ-193 00, Praha 9, Czech Republic. E-mail: [email protected] 3 Gennaro Coppa, 1, rue du Courlis, F-08350 Villers-sur-Bar, France. E-mail: [email protected] 4Roman J. Godunko, State Museum of Natural History, National Academy of Sciences of Ukraine, Teatralna 18, 79008 Lviv, Ukraine 5Biology Centre, Czech Academy of Sciences, Institute of Entomology, Branišovská 31, 37005 České Budějovice, Czech Republic [email protected] 6Omar Lodovici, Museo Scienze Naturali “E. Caffi”, Piazza Cittadella 10, I-24129 Bergamo, Italy. E-mail: [email protected] 7Katarzyna Majecka and Janusz Majecki, Department of Experimental Zoology and Evolutionary Biology, University of Lódz, Banacha 12/16, 90-237 Lódz, Poland. E-mails: [email protected], [email protected] 8Bronislaw Szczęsny, Institute of Nature Conservation, Polish Academy of Sciences, Kraków, Poland. E-mail: [email protected] 9Gorazd Urbanic, Institute for Water of the Republic of Slovenia, Hajdrihova 28c,1000 Ljubljana, Slovenia Email: [email protected] 10University of Ljubljana, Biotechnical Faculty, Department of Biology, Večna pot 111, 1000 Ljubljana, Slovenia E-mail. [email protected] 11Marco Valle, Museo Scienze Naturali “E. Caffi”, Piazza Cittadella 10, I-24129 Bergamo, Italy. E-mail: [email protected] Abstract. Speciation traits of paramere, paraproct and aedeagus were applied to find initial split criteria with fine structure analysis in order to prepare diverged trait matrices for delimiting phylogenetic incipient species of unsettled limnephilid taxa in the early stages of reproductive isolation. A brief history is presented how this phenotypic taxonomic tool of the speciation traits was discovered and applied in caddisfly taxonomy. The theoretical basis was elaborated for the phenotypic speciation trait by reviewing several relevant topics in the sciences of taxonomy, molecular genetics and phylogenetics. Perspectives of integrative taxonomy is discussed in context of phenotype versus genotype, immensely complex phenotype versus phenomic challenge, taxonomic impediment versus genetic expedient, taxonomic adaptation of genetic vocabulary versus genetic sophistication and virtualization, New Systematics of Huxley and Mayr versus New Taxonomy of Wheeler. Debates on magic trait, speciation phenotype, speciation trait and super traits are discussed concluding that evolution works with phenotype and why the cryptic species concept is irrelevant. Briefly summarized how speciation traits evolve in sexual selection, through accelerated reproductive isolation with genital evolution through sex-limited speciation traits, including minor sex chromosomes. Why neutral molecular markers are blind compared to the adaptive speciation traits sensitized by fine structure analysis and backed by the potential of high-tech and high-throughput phenotyping and cyber-infrastructure broadly accessible and fed by computable phenotype descriptions. What sort of genetics could really help taxonomy to describe biodiversity of the over 100 million unknown taxa? Collecting new and re-examining old type materials deposited in various collections, the following taxonomic actions were elaborated by speciation traits. Drusus bolivari new species complex has been erected with redescription of Drusus bolivari (McLachlan, 1876), with species status resurrection of D. estrellensis (McLachlan, 1884) stat. restit., with description of five new species: D. carmenae Oláh, sp. nov., D. gonzalezi Oláh, sp. nov., D. grafi Oláh, sp. nov., D. gredosensis Oláh, sp. nov., D. jesusi Oláh, sp. nov., D. pyrenensis Oláh & Coppa, sp. nov. Genus Isogamus is revised with description of two new species: I. baloghi Oláh, sp. nov., I. balinti Oláh, sp. nov. Melampophylax genus revised with one new species cluster: M. nepos, with two new species descriptions: M. keses Coppa & _______________________________________________________________________________________________________ urn:lsid:zoobank.org:pub:DD21C9A2-2104-4EE6-8F49-ADBDE1AC81C6 HU ISSN 2063-1588 (online), HU ISSN 0237-5419 (print) http://dx.doi.org/10.18348/opzool.2015.1.3 *This article is dedicated to the memory of the late Prof. Dr. Sándor Mahunka, ordinary member of HAS. Oláh et al.: Limnephilid taxa revised by speciation traits Oláh, sp. nov. M. szczesnyorum Oláh & Chvojka, sp. nov., with three new species status: M. banaticus Botosaneanu, 1995 stat. nov., M. gutinicus Botosaneanu, 1995 stat. nov., M. triangulifera Botosaneanu, 1957 stat. nov. Rhadicoleptus genus revised with redescription of R. alpestris (Kolenati, 1848), with three new species status: R. macedonicus Botosaneanu & Riedel, 1965 stat. nov. R. meridiocarpaticus Botosaneanu & Riedel, 1965 stat. nov. R. sylvanocarpaticus Botosaneanu & Riedel, 1965 stat. nov., with one species status resurrection: R. spinifer (McLachlan, 1875) stat. restit. Based on paramere evolution the Rhadicoleptus genus is transferred from the tribe Limnephilini to Stenophylacini. Annitella kosciuszkii new species complex has been erected and revised with redescription of A. chomiacensis (Dziędzielewicz, 1908), A. lateroproducta (Botosaneanu, 1952), with one species status resurrection: A. kosciuszkii Klapálek, 1907 stat. restit., with description of a new species: A. wolosatka Oláh &Szczęsny, sp. nov., with two new synonyms: A. dziedzielewiczi Schmid, 1952 synonym of A. kosciuszkii. syn. nov., A. transylvanica Murgoci, 1957 synonym of A. kosciuszkii. syn. nov. Chaetopteryx rugulosa species group revised with description of five new species: C. balcanica Oláh, sp. nov., C. karima Oláh, sp. nov., C. kozarensis Oláh, sp. nov., C. psunjensis Oláh, sp. nov., C. tompa Oláh, sp. nov., with three species status resurrections: C. papukensis Oláh & Szivák, 2012 stat. restit., C. prealpensis Oláh, 2012 stat. restit., C. zalaensis Oláh, 2012 stat. restit. Psilopteryx psorosa new species group erected and revised with three new sibling species complexes: P. bohemosaxonica, P. carpathica, P. psorosa, with two new species descriptions: P. javorensis Oláh, sp. nov., P. harmas Oláh & Chvojka, sp. nov. with one species status resurrection: P. carpathica Schmid, 1952 stat. restit., with three new species status: P. bohemosaxonica Mey & Botosaneanu, 1985 stat. nov., P. retezatica Botosaneanu & Schneider, 1978 stat. nov., P. transylvanica Mey & Botosaneanu, 1985 stat. nov. Keywords. Speciation traits, neutral and adaptive molecular markers, neutral and adaptive traits, cryptic species, sibling species, incipient species, sexual selection, genital evolution, phenomics versus genomics, new species. INTRODUCTION the genotypes of neutral DNA markers? Why phenomics needs to get importance over genomics he central role of the parameres in the early in biodiversity research that is in taxonomy? Why T stages of speciation processes of certain cad- has the new taxonomy shifted the primary em- disfly taxa, driven by sexual selection, was recog- phasis in taxonomy from genomic basis to diverg- nised and information slowly accumulated in our ing phenotypes of incipient species? Why speci- previous studies on limnephilids (Oláh et al. ation phenotypes have new evolutionary perspec- 2012; Oláh et al. 2013a, b, c; Oláh et al. 2014). tives in sexual selection? Trials to answer all these Paramere and aedeagus diversity, under the term questions, taxonomists need to learn more how to of morphological polymorphism, has already been adapt in taxonomy the relevant part of the knowl- recognised earlier in the Potamophylax cingulatus edge, principles, theories, models, algorithms, and species complex (Szczęsny 1990). In this paper procedures accumulated in genetics. we give a historical overview how this phenotypic taxonomic tool was discovered. We review its For instance, why don’t we use Pst, an theoretical background and apply it to revise fur- approximated outlier analogue of Qst or Fst, to ther unsettled limnephilid taxa, all having closely compute the phenotypic divergence of parameres, related incipient species: Rhadicoleptus, Isoga- in case if not visible properly with the micro- mus, Melampophylax genera, Chaetopteryx rugu- scoped eye of the taxonomist? Why taxonomist losa, Psilopteryx psorosa species groups, Drusus and geneticist do not team for simple Pst – Fst or bolivari, Annitella kosciuszkii species complexes. for complex Qst – Fst comparisions? Both would Our reliance on fine structure analysis of re- greatly contribute to distinguish between natural productive traits is based on empirical diversity selection and genetic drift as causes of divergence evidences evolved by sexual selection. Therefore of speciation traits instead of giving jobs for tech- we instigate taxonomists for more sophistication nicians or students to measure more neutral mark- in the phenomics of alpha taxonomy. We discuss ers or barcoding (Leinonen et al. 2013). In return, several aspects of how parameres
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