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      A species-group key and notes on phylogeny and character evolution in New Guinean Exocelina Broun, 1886 diving beetles (Coleoptera, Dytiscidae, Copelatinae)

      research-article
      1 , , 2 , 3
      ZooKeys
      Pensoft Publishers
      Morphology, New Guinea, phylogeny, water beetles

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          Abstract

          Detailed information about the known species groups of Exocelina Broun, 1886 from New Guinea is presented, including species numbers, distribution, and references of species-group diagnoses, keys to the species, and species descriptions. An identification key to all species groups is provided. Phylogeny and morphological character evolution are discussed.

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          The towering orogeny of New Guinea as a trigger for arthropod megadiversity.

          Early studies on Melanesian mountain systems provided insights for fundamental evolutionary and ecological concepts. These island-like systems are thought to provide opportunities in the form of newly formed, competition-free niches. Here we show that a hyperdiverse radiation of freshwater arthropods originated in the emerging central New Guinea orogen, out of Australia, about 10 million years ago. Further diversification was mainly allopatric, with repeated more recent colonization of lowlands as they emerged in the form of colliding oceanic island arcs, continental fragments and the Papuan Peninsula, as well as recolonization of the central orogen. We unveil a constant and ongoing process of lineage accumulation while the carrying capacity of the island is about to be reached, suggesting that lineage diversification speed now exceeds that of landmass/new ecological opportunity formation. Therefore, the central orogeny of New Guinea acts as a motor of diversification for the entire region.
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            MtDNA phylogeny and biogeography of Copelatinae, a highly diverse group of tropical diving beetles (Dytiscidae).

            Copelatinae is a diverse lineage of diving beetles (Dytiscidae) frequently encountered in wet tropical and subtropical forests, but phylogenetic relationships are very poorly understood. We performed a phylogenetic and biogeographic analysis of this worldwide distributed group based on 50 species including a representative sample of major taxonomic groups and biogeographical regions. DNA sequences were obtained for the mitochondrial genes cytochrome oxidase I, cytochrome b, and 16S rRNA, for a total of 1575 aligned nucleotide positions. We found Copelatinae to be monophyletic, placed in a derived position and not sister to all remaining dytiscids, as had been suggested by earlier authors. The largest genus, Copelatus with some 460 known species was paraphyletic with respect to the smaller genera Lacconectus and Aglymbus. Among the major lineages of Copelatus, the subgenus Papuadytes was consistently recovered as sister to all other species (including Lacconectus and Aglymbus) with the possible exception of two western Palearctic taxa. We propose that the subgenus Papuadytes is removed from Copelatus and assigned generic status. Likewise, the two western Palearctic Copelatus are removed from this genus, and assigned the available genus name Liopterus. Our best phylogenetic hypothesis retrieved Afrotropical and New Guinean plus Australian species of Copelatus as monophyletic. Asian species were paraphyletic with respect to a species from Sulawesi which grouped with the species from New Guinea. Asian species were also paraphyletic with respect to Oriental Lacconectus, which was grouped with a clade of Neotropical species. Neotropical Copelatus form at least two separate lineages. The biogeographical evolution of Papuadytes is consistent with the relative age of the landmasses in the Austral region. Basal species are Australian, and successively derived ones are from New Caledonia and New Guinea. One species apparently dispersed from New Caledonia to China. Assuming a molecular clock and using a standard calibration of 2% divergence/MY the origin of Copelatinae is estimated to be between 85 and 95 MY.
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              Infrequent and unidirectional colonization of hyperdiverse Papuadytes diving beetles in New Caledonia and New Guinea.

              We present a molecular phylogenetic analysis of 2808 aligned bp of rrnL, cox1, cob, H3 and 18S rRNA of all major morphological groups of Papuadytes diving beetles (Coleoptera: Dytiscidae) which are diverse in running water habitats throughout the Australian region. We focus on the origin of the fauna of the megadiverse islands of New Guinea and New Caledonia. Parsimony as well as Bayesian analyses suggest a basal position of Australian species in a paraphyletic series, with more recent nested radiations in New Caledonia and New Guinea. According to molecular clock analyses, both landmasses were colonized during the Miocene, which matches geological data and corroborates similar findings in other taxonomic groups. Our analyses suggest that dispersal played an important role in the formation of these large insular faunas, although successful colonization appears to be a rare event, and, in this case, is unidirectional. Whether or not a lineage is present on an island is due to chance: Papuadytes are absent from Fiji, where related Copelatus have radiated extensively in the same habitats occupied by Papuadytes in New Caledonia and New Guinea, while Copelatus are absent from New Caledonia. Lineages of Papuadytes apparently colonized New Caledonia twice, around 14 and 9 MYA according to the molecular calibration, and both lineages are derived from an Australian ancestor. The older clade is represented only by two apparently relictual mountain species (one morphologically strongly adapted to highly ephemeral habitats), while the younger clade contains at least 18 species exhibiting a great morphological diversity. The 150+ species in New Guinea are monophyletic, apparently derived from an Australian ancestor, and constitute a morphologically rather homogenous group. The tree backbone remains insufficiently supported under parsimony and Bayesian analyses, where shorter branches suggest a rapid sequence of major branching events.

                Author and article information

                Contributors
                Journal
                Zookeys
                Zookeys
                2
                urn:lsid:arphahub.com:pub:45048D35-BB1D-5CE8-9668-537E44BD4C7E
                urn:lsid:zoobank.org:pub:91BD42D4-90F1-4B45-9350-EEF175B1727A
                ZooKeys
                Pensoft Publishers
                1313-2989
                1313-2970
                2022
                22 November 2022
                : 1131
                : 31-58
                Affiliations
                [1 ] Naturhistorisches Museum, Burgring 7, A-1010 Vienna, Austria Naturhistorisches Museum Vienna Austria
                [2 ] SNSB-Zoologische Staatssammlung München, Münchhausenstraße 21, D-81247 Munich, Germany SNSB-Zoologische Staatssammlung München Munich Germany
                [3 ] GeoBioCenter, Ludwig-Maximilians-University, Munich, Germany Ludwig-Maximilians-University Munich Germany
                Author notes
                Corresponding author: Helena Shaverdo (shaverdo@mail.ru, helena.shaverdo@nhm-wien.ac.at)

                Academic editor: Mariano Michat

                Author information
                https://orcid.org/0000-0001-5034-7342
                https://orcid.org/0000-0002-3773-6586
                Article
                94205
                10.3897/zookeys.1131.94205
                9836653
                36761459
                e3af6b1c-4bf9-4737-9c92-55577574adc3
                Helena Shaverdo, Michael Balke

                This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

                History
                : 29 August 2022
                : 27 October 2022
                Funding
                Funded by: Austrian Science Fund 501100002428 http://doi.org/10.13039/501100002428
                Categories
                Research Article
                Adephaga
                Arthropoda
                Coleoptera
                Dytiscidae
                Hexapoda
                Insecta
                Invertebrata
                Identification Key
                Molecular Systematics
                Phylogeny
                Systematics
                Taxonomy
                Neogene
                Australasia
                Pacific
                Papua New Guinea

                Animal science & Zoology
                morphology,new guinea,phylogeny,water beetles
                Animal science & Zoology
                morphology, new guinea, phylogeny, water beetles

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