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Lineage Sorting by Parameres in Limnephilinae Subfamily (Trichoptera): with Description of a New Tribe, New Genera and New Species

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Lineage Sorting by Parameres in Limnephilinae Subfamily (Trichoptera): with Description of a New Tribe, New Genera and New Species Opusc. Zool. Budapest, 2019, 50(Supplementum 1): 03–98 Lineage sorting by parameres in Limnephilinae subfamily (Trichoptera): with description of a new tribe, new genera and new species 1 2 3 4 5 6 J. OLÁH , T. ANDERSEN , S. BESHKOV , A. BILALLI , G. COPPA , H. IBRAHIMI , K.A. 7 8 9 10 11 12 JOHANSON , T. KOVÁCS , W. MEY , M. MUSLIU J. OLÁH JR & A. RUIZ-GARCIA 1János Oláh, Residence postal address: Tarján u. 28, H-4032 Debrecen, Hungary, [email protected] 2Trond Andersen, Department of Natural History, University Museum of Bergen, University of Bergen, P.O. Box 7800, N-5020 Bergen, Norway, [email protected] 3Stoyan Beshkov, National Museum of Natural History, 1 Tsar Osvoboditel Blvd. 1000 Sofia, Bulgaria, [email protected] 4Astrit Bilalli, University of Peja "Haxhi Zeka", Faculty of Agribusiness, “UÇK” street, 30000 Pejë, Republic of Kosovo, [email protected] 5Gennaro Coppa, 1, rue du Courlis, F-08350 Villers-sur-Bar, France, [email protected] 6Halil Ibrahimi, University of Prishtina, Faculty of Mathematics and Natural Sciences, Department of Biology, Mother Teresa p.n., 10000 Prishtina, Kosovo, [email protected] 7Kjell Arne Johanson, Swedish Museum of Natural History, Department of Zoology, Box 50007, SE-10405 Stockholm, Sweden, [email protected] 8Tibor Kovács, Mátra Museum of Hungarian Natural History Museum, Kossuth Lajos u. 40, H-3200 Gyöngyös, Hungary, [email protected] 9Wolfram Mey, Museum für Naturkunde, Humboldt-Universität, Invalidenstr. 43, D-10115 Berlin, Germany, [email protected] 10Milaim Musliu, University of Peja "Haxhi Zeka", Faculty of Agribusiness, “UÇK” street, 30000 Pejë, Republic of Kosovo, [email protected] 11János Oláh jr., Residence postal address: Tarján u. 6, H-4032 Debrecen, Hungary, [email protected] 12Antonio Ruiz-Garcia, Departamento de Sistemas Físicos, Químicos y Naturales (Zoología), Universidad Pablo de Olavide, A-376, Km 1, 41013 Sevilla, Spain, [email protected] Abstract. The discovery of the new Agaphylax genus with unique paramere organisation has initiated our lineage sorting of tribes by parameres in the Limnephilinae subfamily applying the principles and procedures of fine phenomics in order to establish transformation series of the polarized plesiomorphy-apomorphy character states for each limnephiline genera. According to the extraordinary high diversity the paramere that is the stimulatory and titillating structure of the phallic organ is a speciation supertrait. This adaptive trait is directly involved in the processes of reproductive isolation and diverging as subtle initial split of lineages producing the incipient sibling species in the recent past of contemporary speciation processes. Contrary, the drastic divergence of the Agaphylax plated paramere is much older, similarly to the many-spined parameres of the Hesperophylacini tribe. It has been initiated by drastic combined and synchronous external and internal stochastic effects, processed in ancestral sexual integrative adaptation as well as organised and fixed in older and deeper coalescence events and appears as a character with tribe ranking potential. To open a wider perspective, a systemic relational analysis is required in the future including other adaptive or neutral character transformation series, due to the burden of taxonomic incongruences grounded by chimerism in stochastic genetic reticulation. Traits of species are mixed products coming from various sources. Only character combinations can and ought to be analysed in terms how to classify taxa. We have polarized eight genitalic characters additional to parameres for a future analysis of the potential of character combinations. Limnephilinae subfamily is composed of Limnephilini, Chilostigmatini, Chaetopterygini, Stenophylacini and Hespero- phylacini tribes and here we established the new Agaphylacini tribe. Based on parameres we have delineated taxa in lineage sorting and described two new genera: Fogophylax gen. nov., Simaphylax gen. nov. and fourteen new species: Anabolia alsoja, A. hepehupa, Asynarchus kimaros, Limnephilus kerekes, L. maghrebensis, L. oblos, Homophylax beges, H. coros, Chaetopteroides plackovicensis, C. rilaensis, Allogamus ketpar, Platyphylax beshkovi, Pycnopsyche letova and P. telea spp. nov. The by-product of this survey is a world atlas of paramere drawings for the entire Limnephilinae subfamily. Keywords. Paramere organisation, paramere polarization, paramere atlas, lineage sorting. _______________________________________________________________________________________________________ urn: lsid:zoobank.org:pub:31329FBB-5B10-4B5E-8F7D-56DA267CAD78 published: 15 November 2019 HU ISSN 2063-1588 (online) http://dx.doi.org/10.18348/opzool.2019.S1.3 Oláh et al.: Lineage sorting by parameres in Limnephilinae INTRODUCTION and fixed probably in older and deeper coales- cence events and processed by ancestral sexual his paramere survey enriched with other cha- integrative adaptation (Oláh et al. 2018). Sexually T racter transformation series was initiated by adapted structures diverged in contemporary re- the discovery of the unique Agaphylax genus. cent past exhibit usually more subtle, but never- Recently we have described a new European lim- theless stable shape divergences of the initial split. nephiline genus Agaphylax characterized by sur- They are particularly suitable to delineate the prisingly unique parameres of drastic shape diver- recent lineages of the new incipient sibling spe- gence, almost incongruent to the known paramere cies. These speciation traits are useful and prac- patterns in the subfamily. This unusual paramere tical for taxonomist in converting the natural kind structure, the product of drastic divergence in in- of divergences into nominal kind of taxonomic tegrative organisation could be a character with rank in early stages of speciation processes. The tribe ranking potential. Therefore it was suggested organisation of these adaptive structures is pro- to conduct a comprehensive paramere revision on cessed frequently in post-mating prezygotic copu- the entire subfamily (Oláh et al. 2018). latory mechanisms like cryptic female choice with over 25 known mechanisms (Eberhard 2010) Here we survey the paramere organisation in including gametic incompatibilities. The diver- each tribe of the Limnephilinae subfamily along gence of speciation traits, like any kind of diver- the generic lineages by the principles and proce- gences, is initiated stochastically by random envi- dures of fine phenomics. According to the extra- ronmental effects, but stabilised on molecular ordinary high diversity the paramere is a speci- level by complex adaptive and integrative mecha- ation supertrait. How does the diversity of para- nisms (Oláh et al. 2017, Oláh & Oláh 2017). mere, the stimulatory and titillating substructure of the phallic organ, influences the speciation If the adaptive paramere shape divergence is processes and results in lineage sorting? Based on drastic and the morphological differences are parameres we delineate taxa in lineage sorting and great, as we have detected in Agaphylax genus, it as a by-product we describe a new tribe, new was probably produced by deeper coalescence genera and species accordingly, applying the events during older and drastic stochastic acci- shape organisation of this adaptive trait that is dents and fixed by ancestral integration and not by directly involved in the processes of reproductive contemporary one. The combination of gross and isolation. To open a wider perspective, a systemic fine paramere shape divergences may have real relational analysis is required on other adaptive or potential and unexplored capacity to utilise in the neutral character transformation series in the lineage sorting of multiple nodes by the gross future, due to the burden of taxonomic incon- phenomics of ancestral traits and by the fine gruences grounded by chimerism in stochastic phenomics of contemporary more subtle speci- genetic reticulation. The analytic possibility of ation traits. This possibility inspired us to ela- character combinations is established here by borate an outline of working hypothesis for a surveying the plesiomorphic or apomorphic states future comprehensive and systemic revision of of eight basic genital substructures besides the paramere organisation in the Limnephilinae sub- parameres in each genus. The by-product of this family (Oláh et al. 2018). survey is a world atlas of paramere drawings for the entire Limnephilinae subfamily. But what we can really gain by elaborating paramere trees of limnephiline genera if clouds of endless trait trees exist inside each species trees Theorethical grounding frequently contradicting to each other (Oláh et al. 2019)? Can we sort lineages and delineate phylo- Such a large shape divergence must have been genetic species directly by the trees of the adap- initiated by combined and synchronous external tive paramere divergences? Yes, if these particu- and internal stochastic effect, as well as organised lar traits are supertraits representing the repro- 4 Oláh et al.: Lineage sorting by parameres in Limnephilinae ductive barrier building by their shape diver- species trees of many other morphological traits gences either in sympatry or more frequently in remains ambiguous. allopatry under strong genetic drift of small isolated populations. Recombination, genetic drift, genetic draft, gene flow, lateral transfer, introgression and many These questions lead us to the basic problem more, innumerable,
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