Integrative Taxonomy on the Fast Track
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Consequences of Evolutionary Transitions in Changing Photic Environments
bs_bs_banner Austral Entomology (2017) 56,23–46 Review Consequences of evolutionary transitions in changing photic environments Simon M Tierney,1* Markus Friedrich,2,3 William F Humphreys,1,4,5 Therésa M Jones,6 Eric J Warrant7 and William T Wcislo8 1School of Biological Sciences, The University of Adelaide, North Terrace, Adelaide, SA 5005, Australia. 2Department of Biological Sciences, Wayne State University, 5047 Gullen Mall, Detroit, MI 48202, USA. 3Department of Anatomy and Cell Biology, Wayne State University, School of Medicine, 540 East Canfield Avenue, Detroit, MI 48201, USA. 4Terrestrial Zoology, Western Australian Museum, Locked Bag 49, Welshpool DC, WA 6986, Australia. 5School of Animal Biology, University of Western Australia, Nedlands, WA 6907, Australia. 6Department of Zoology, The University of Melbourne, Melbourne, Vic. 3010, Australia. 7Department of Biology, Lund University, Sölvegatan 35, S-22362 Lund, Sweden. 8Smithsonian Tropical Research Institute, PO Box 0843-03092, Balboa, Ancón, Republic of Panamá. Abstract Light represents one of the most reliable environmental cues in the biological world. In this review we focus on the evolutionary consequences to changes in organismal photic environments, with a specific focus on the class Insecta. Particular emphasis is placed on transitional forms that can be used to track the evolution from (1) diurnal to nocturnal (dim-light) or (2) surface to subterranean (aphotic) environments, as well as (3) the ecological encroachment of anthropomorphic light on nocturnal habitats (artificial light at night). We explore the influence of the light environment in an integrated manner, highlighting the connections between phenotypic adaptations (behaviour, morphology, neurology and endocrinology), molecular genetics and their combined influence on organismal fitness. -
Opsin Transcripts of Predatory Diving Beetles: a Comparison of Surface
Downloaded from http://rsos.royalsocietypublishing.org/ on January 28, 2015 Opsin transcripts of predatory diving beetles: a rsos.royalsocietypublishing.org comparison of surface and Research subterranean photic niches Cite this article: Tierney SM, Cooper SJB, Simon M. Tierney1,StevenJ.B.Cooper1,2, Saint KM, Bertozzi T, Hyde J, Humphreys WF, 2 1,2 1 Austin AD. 2015 Opsin transcripts of predatory Kathleen M. Saint , Terry Bertozzi , Josephine Hyde , diving beetles: a comparison of surface and William F. Humphreys1,3,4 and Andrew D. Austin1 subterranean photic niches. R. Soc. open sci. 2: 140386. 1Australian Centre for Evolutionary Biology and Biodiversity and School of Biological http://dx.doi.org/10.1098/rsos.140386 Sciences, University of Adelaide, South Australia 5005, Australia 2Evolutionary Biology Unit, South Australian Museum, North Terrace, Adelaide, South Australia 5000, Australia 3Terrestrial Zoology, Western Australian Museum, Locked Bag 49, Welshpool DC, Received: 16 October 2014 Western Australia 6986, Australia Accepted: 22 December 2014 4School of Animal Biology, University of Western Australia, Nedlands, Western Australia 6907, Australia SMT, 0000-0002-8812-6753 Subject Category: 1. Summary Biology (whole organism) The regressive evolution of eyes has long intrigued biologists Subject Areas: yet the genetic underpinnings remain opaque. A system evolution/ecology/genomics of discrete aquifers in arid Australia provides a powerful comparative means to explore trait regression at the genomic level. Multiple surface ancestors from two tribes of diving Keywords: beetles (Dytiscidae) repeatedly invaded these calcrete aquifers regressive evolution, opsin, pleiotropy, and convergently evolved eye-less phenotypes. We use this pseudogene, Bidessini, Hydroporini system to assess transcription of opsin photoreceptor genes among the transcriptomes of two surface and three subterranean dytiscid species and test whether these genes have evolved under neutral predictions. -
Introduction of the Exocelina Ekari-Group with Descriptions of 22 New Species from New Guinea (Coleoptera, Dytiscidae, Copelatinae)
A peer-reviewed open-access journal ZooKeys 250: 1–76Introduction (2012) of the Exocelina ekari-group with descriptions of 22 new species... 1 doi: 10.3897/zookeys.250.3715 RESEARCH ARTICLE www.zookeys.org Launched to accelerate biodiversity research Introduction of the Exocelina ekari-group with descriptions of 22 new species from New Guinea (Coleoptera, Dytiscidae, Copelatinae) Helena V. Shaverdo1,¶, Suriani Surbakti2,‡, Lars Hendrich3,§, Michael Balke4,| 1 Naturhistorisches Museum, Burgring 7, A-1010 Vienna, Austria 2 Department of Biology, Universitas Cendrawasih, Jayapura, Papua, Indonesia 3 Zoologische Staatssammlung München, Münchhausenstraße 21, D-81247 Munich, Germany 4 Zoologische Staatssammlung München, Münchhausenstraße 21, D-81247 Mu- nich, Germany and GeoBioCenter, Ludwig-Maximilians-University, Munich, Germany ¶ urn:lsid:zoobank.org:author:262CB5BD-F998-4D4B-A4F4-BFA04806A42E ‡ urn:lsid:zoobank.org:author:0D87BE16-CB33-4372-8939-A0EFDCAA3FD3 § urn:lsid:zoobank.org:author:06907F16-4F27-44BA-953F-513457C85DBF | urn:lsid:zoobank.org:author:945480F8-C4E7-41F4-A637-7F43CCF84D40 Corresponding author: Helena V. Shaverdo ([email protected], [email protected]) Academic editor: M. Fikácek | Received 10 August 2012 | Accepted 8 November 2012 | Published 13 December 2012 urn:lsid:zoobank.org:pub:FC92592B-6861-4FE2-B5E8-81C50154AD2A Citation: Shaverdo HV, Surbakti S, Hendrich L, Balke M (2012) Introduction of the Exocelina ekari-group with descriptions of 22 new species from New Guinea (Coleoptera, Dytiscidae, Copelatinae). ZooKeys 250: 1–76. doi: 10.3897/zookeys.250.3715 Abstract The Exocelina ekari-group is here introduced and defined mainly on the basis of a discontinuous outline of the median lobe of the aedeagus. The group is known only from New Guinea (Indonesia and Papua New Guinea). -
Coleoptera, Dytiscidae, Copelatinae)
A peer-reviewed open-access journal ZooKeys 304:Exocelina 83–99 (2013) baliem sp. n., the only known pond species of New Guinea Exocelina... 83 doi: 10.3897/zookeys.304.4852 RESEARCH ARTICLE www.zookeys.org Launched to accelerate biodiversity research Exocelina baliem sp. n., the only known pond species of New Guinea Exocelina Broun, 1886 (Coleoptera, Dytiscidae, Copelatinae) Helena V. Shaverdo1,†, Lars Hendrich2,‡, Michael Balke3,§ 1 Naturhistorisches Museum, Burgring 7, A-1010 Vienna, Austria 2 Zoologische Staatssammlung München, Münchhausenstraße 21, D-81247 Munich, Germany 3 Zoologische Staatssammlung München, Münchhausen- straße 21, D-81247 Munich, Germany and GeoBioCenter, Ludwig-Maximilians-University, Munich, Germany † urn:lsid:zoobank.org:author:262CB5BD-F998-4D4B-A4F4-BFA04806A42E ‡ urn:lsid:zoobank.org:author:06907F16-4F27-44BA-953F-513457C85DBF § urn:lsid:zoobank.org:author:945480F8-C4E7-41F4-A637-7F43CCF84D40 Corresponding author: Helena V. Shaverdo ([email protected]; [email protected]) Academic editor: M. Fikácek | Received 11 February 2012 | Accepted 15 March 2013 | Published 28 May 2013 urn:lsid:zoobank.org:pub:39C569AF-2376-4FA2-9204-A6F0B81A5970 Citation: Shaverdo HV, Hendrich L, Michael Balke M (2013) Exocelina baliem sp. n., the only known pond species of New Guinea Exocelina Broun, 1886 (Coleoptera, Dytiscidae, Copelatinae). ZooKeys 304: 83–99. doi: 10.3897/ zookeys.304.4852 Abstract Exocelina baliem sp. n. is described from the Baliem Valley in the Central Mountain Range of New Guinea (Papua Province, Indonesia). Having striolate elytra, different structure and setation of the male and fe- male genitalia and tarsomeres, and inhabiting swampy ponds, the new species differs from all known New Guinea species, which have smooth elytra and are stream associated. -
Coleoptera: Dytiscidae) Yves Alarie, J
Tijdschrift voor Entomologie 156 (2013) 1–10 brill.com/tve Descriptions of larvae of the North American endemic stygobiontic Ereboporus naturaconservatus Miller, Gibson & Alarie and Haideoporus texanus Young & Longley (Coleoptera: Dytiscidae) Yves Alarie, J. Randy Gibson & Kelly B. Miller The larvae of the North American stygobiontic dytiscid species Ereboporus naturaconservatus Miller, Gibson & Alarie, 2009 and Haideoporus texanus Young & Longley, 1976 are described with an emphasis on chaetotaxy of the head capsule, head appendages, legs, last abdominal segment and urogomphi. Both of these species share the presence of a nasale and the absence of the primary pores MXd and LAc, which have been recognized as synapomorphies for members of the subfamily Hydroporinae. Out of the common convergent characteristics associated with hypogaeic living, no synapomorphies were found that could relate Haideoporus texanus and Ereboporus naturaconservatus, which reinforces the hypothesis that these species evolved independently within the subfamily Hydroporinae. In terms of morphological adaptations, E. naturaconservatus stands as a remarkable hydroporine in that its larvae evolved a truncated last abdominal segment and a very elongate urogomphomere 1 relative to urogomphomere 2. Keywords: Adephaga, Dytiscidae, Hydroporini, stygobiontic, larval chaetotaxy. Yves Alarie*, Department of Biology, Laurentian University, Ramsey Lake Road, Sudbury, ON, Canada P3E 2C6. [email protected] J. Randy Gibson, National Fish Hatchery and Technology Center, U.S. Fish and Wildlife Service, 500 East McCarty Lane, San Marcos, TX 78666, USA. [email protected] Kelly B. Miller, Department of Biology and Museum of Southwestern Biology, University of New Mexico, Albuquerque, NM 87131, USA. [email protected] Introduction tinct (Porter 2007). Most stygobiontic beetles are Stygobiontic aquatic Coleoptera represent a hetero- placed within the coleopteran suborder Adephaga, geneous and fascinating grouping of taxa associated and a great number of them in the dytiscid subfam- with underground waters. -
Mitochondrial Cox1 Sequence Data Reliably Uncover Patterns of Insect Diversity but Suffer from High Lineage- Idiosyncratic Error Rates
Mitochondrial Cox1 Sequence Data Reliably Uncover Patterns of Insect Diversity But Suffer from High Lineage- Idiosyncratic Error Rates Lars Hendrich1,2, Joan Pons3., Ignacio Ribera4, Michael Balke1,2*. 1 Department of Entomology, Zoological State Collection, Munich, Germany, 2 GeoBioCenter, Ludwig-Maximilians-University, Munich, Germany, 3 Departament de Biodiversitat i Conservacio´, Institut Mediterrani d’Estudis Avanc¸ats, Esporles, Spain, 4 Institute of Evolutionary Biology, Barcelona, Spain Abstract Background: The demand for scientific biodiversity data is increasing, but taxonomic expertise is often limited or not available. DNA sequencing is a potential remedy to overcome this taxonomic impediment. Mitochondrial DNA is most commonly used, e.g., for species identification (‘‘DNA barcoding’’). Here, we present the first study in arthropods based on a near-complete species sampling of a family-level taxon from the entire Australian region. We aimed to assess how reliably mtDNA data can capture species diversity when many sister species pairs are included. Then, we contrasted phylogenetic subsampling with the hitherto more commonly applied geographical subsampling, where sister species are not necessarily captured. Methodology/Principal Findings: We sequenced 800 bp cox1 for 1,439 individuals including 260 Australian species (78% species coverage). We used clustering with thresholds of 1 to 10% and general mixed Yule Coalescent (GMYC) analysis for the estimation of species richness. The performance metrics used were taxonomic accuracy and agreement between the morphological and molecular species richness estimation. Clustering (at the 3% level) and GMYC reliably estimated species diversity for single or multiple geographic regions, with an error for larger clades of lower than 10%, thus outperforming parataxonomy. -
Coleoptera: Dytiscidae) Yves Alarie, J
Tijdschrift voor Entomologie 156 (2013) 1–10 brill.com/tve Descriptions of larvae of the North American endemic stygobiontic Ereboporus naturaconservatus Miller, Gibson & Alarie and Haideoporus texanus Young & Longley (Coleoptera: Dytiscidae) Yves Alarie, J. Randy Gibson & Kelly B. Miller The larvae of the North American stygobiontic dytiscid species Ereboporus naturaconservatus Miller, Gibson & Alarie, 2009 and Haideoporus texanus Young & Longley, 1976 are described with an emphasis on chaetotaxy of the head capsule, head appendages, legs, last abdominal segment and urogomphi. Both of these species share the presence of a nasale and the absence of the primary pores MXd and LAc, which have been recognized as synapomorphies for members of the subfamily Hydroporinae. Out of the common convergent characteristics associated with hypogaeic living, no synapomorphies were found that could relate Haideoporus texanus and Ereboporus naturaconservatus, which reinforces the hypothesis that these species evolved independently within the subfamily Hydroporinae. In terms of morphological adaptations, E. naturaconservatus stands as a remarkable hydroporine in that its larvae evolved a truncated last abdominal segment and a very elongate urogomphomere 1 relative to urogomphomere 2. Keywords: Adephaga, Dytiscidae, Hydroporini, stygobiontic, larval chaetotaxy. Yves Alarie*, Department of Biology, Laurentian University, Ramsey Lake Road, Sudbury, ON, Canada P3E 2C6. [email protected] J. Randy Gibson, National Fish Hatchery and Technology Center, U.S. Fish and Wildlife Service, 500 East McCarty Lane, San Marcos, TX 78666, USA. [email protected] Kelly B. Miller, Department of Biology and Museum of Southwestern Biology, University of New Mexico, Albuquerque, NM 87131, USA. [email protected] Introduction tinct (Porter 2007). Most stygobiontic beetles are Stygobiontic aquatic Coleoptera represent a hetero- placed within the coleopteran suborder Adephaga, geneous and fascinating grouping of taxa associated and a great number of them in the dytiscid subfam- with underground waters. -
Wikis in Scholarly Publishing
Title Wikis in scholarly publishing Authors Daniel Mietchen[1], Gregor Hagedorn[2], Konrad U. Förstner [3] [4] , M Fabiana Kubke[5], Claudia Koltzenburg [6] [7] , Mark Hahnel[8] and Lyubomir Penev][9]. 1. ↑ Science 3.0 2. ↑ Julius Kühn-Institut, Königin-Luise-Straße 19, 14195 Berlin, Germany 3. ↑ Institute for Molecular Infection Biology, University of W ü rzburg, D-97080 W ü rzburg, Germany 4. ↑ Research Centre for Infectious Diseases, University of W ü rzburg, D-97080 W ü rzburg, Germany 5. ↑ School of Medical Sciences, University of Auckland, New Zealand 6. ↑ Cellular Therapy and Transplantation 7. ↑ Clinic for Stem Cell Transplantation, University Medical Center Hamburg-Eppendorf, Germany 8. ↑ Science 3.0 9. ↑ Pensoft Publishers, 13a Geo Milev Street, 1111 Sofia, Bulgaria Abstract Scientific research is a process concerned with the creation, collective accumulation, contextualization, updating and maintenance of knowledge. Wikis provide an environment that allows to collectively accumulate, contextualize, update and maintain knowledge in a coherent and transparent fashion. Here, we examine the potential of wikis as platforms for scholarly publishing. In the hope to stimulate further discussion, the article itself was drafted on Species-ID - http://species-id.net; a wiki that hosts a prototype for wiki-based scholarly publishing - where it can be updated, expanded or otherwise improved. Introduction "Science is based on building on, reusing and openly criticising the published body of scientific knowledge." --Panton Principles [1] At its inception, a research project is typically fueled by at least one of: new data, new methodological approaches, or new hypotheses. Each of these can contribute either to small incremental steps that further specific knowledge or have more profound effects on the theoretical framework of a field. -
On the Identity of the Fossil Aquatic Beetles from the Tertiary Localities in the Southern Part of the Upper Rhine Graben (Coleoptera, Hydrophilidae, Dytiscidae)
A peer-reviewed open-access journal ZooKeys 78: 15–25 (2011)Fossil aquatic beetles from the Tertiary localities in the southern part... 15 doi: 10.3897/zookeys.78.800 RESEARCH ARTICLE www.zookeys.org Launched to accelerate biodiversity research On the identity of the fossil aquatic beetles from the Tertiary localities in the southern part of the Upper Rhine Graben (Coleoptera, Hydrophilidae, Dytiscidae) Martin Fikáček1,2, Jiří Hájek1, Heiko Schmied3,4 1 Department of Entomology, National Museum, Kunratice 1, CZ-148 00 Praha 2, Czech Republic 2 De- partment of Zoology, Faculty of Science, Charles University in Prague, Viničná 7, CZ-128 44 Praha 2, Czech Republic 3 Steinmann Insitute of Geology, Mineralogy and Palaeontology, University of Bonn, Nußalle 8, 53115 Bonn, Germany 4 Institute of Crop Science and Resource Conservation, Animal Ecology, University of Bonn, Melbweg 42, 53127 Bonn, Germany Corresponding author : Martin Fikáček ( mfi [email protected] ) Academic editor: Jes Rust | Received 12 December 2010 | Accepted 25 January 2011 | Published 28 January 2011 Citation: Fikáček M, Hájek J, Schmied H (2011) On the identity of the fossil aquatic beetles from the Tertiary localities in the southern part of the Upper Rhine Graben (Coleoptera, Hydrophilidae, Dytiscidae). ZooKeys 78 : 15 – 25 . doi: 10.3897/zookeys.78.800 Abstract Th is study focuses on the fossil beetles assigned previously to the family Hydrophilidae described from the localities in the southern part of the Upper Rhine Graben: Brunstatt (France, Alsace) and Kleinkems (Ger- many, Baden-Württemberg) (both dated ca. to Eocene-Oligocene boundary, 34 Ma). Th e identity of Es- cheria convexa Förster, 1891 is fi xed by the designation of its neotype, the species is redescribed, illustrated, transferred from the hydrophilid genus Hydrobius Leach, 1815 to the genus Copelatus Erichson, 1832 (Co- leoptera: Adephaga: Dytiscidae) and compared with other fossil representatives of Copelatus. -
Mitochondrial Cox1 Sequence Data Reliably Uncover Patterns of Insect Diversity but Suffer from High Lineage- Idiosyncratic Error Rates
Mitochondrial Cox1 Sequence Data Reliably Uncover Patterns of Insect Diversity But Suffer from High Lineage- Idiosyncratic Error Rates Lars Hendrich1,2, Joan Pons3., Ignacio Ribera4, Michael Balke1,2*. 1 Department of Entomology, Zoological State Collection, Munich, Germany, 2 GeoBioCenter, Ludwig-Maximilians-University, Munich, Germany, 3 Departament de Biodiversitat i Conservacio´, Institut Mediterrani d’Estudis Avanc¸ats, Esporles, Spain, 4 Institute of Evolutionary Biology, Barcelona, Spain Abstract Background: The demand for scientific biodiversity data is increasing, but taxonomic expertise is often limited or not available. DNA sequencing is a potential remedy to overcome this taxonomic impediment. Mitochondrial DNA is most commonly used, e.g., for species identification (‘‘DNA barcoding’’). Here, we present the first study in arthropods based on a near-complete species sampling of a family-level taxon from the entire Australian region. We aimed to assess how reliably mtDNA data can capture species diversity when many sister species pairs are included. Then, we contrasted phylogenetic subsampling with the hitherto more commonly applied geographical subsampling, where sister species are not necessarily captured. Methodology/Principal Findings: We sequenced 800 bp cox1 for 1,439 individuals including 260 Australian species (78% species coverage). We used clustering with thresholds of 1 to 10% and general mixed Yule Coalescent (GMYC) analysis for the estimation of species richness. The performance metrics used were taxonomic accuracy and agreement between the morphological and molecular species richness estimation. Clustering (at the 3% level) and GMYC reliably estimated species diversity for single or multiple geographic regions, with an error for larger clades of lower than 10%, thus outperforming parataxonomy. -
A New, Widely Distributed Species of the Exocelina Ekari-Group From
A peer-reviewed open-access journal ZooKeys 554: A69–85 new, (2016) widely distributed species of the Exocelina ekari-group from West Papua... 69 doi: 10.3897/zookeys.554.6065 RESEARCH ARTICLE http://zookeys.pensoft.net Launched to accelerate biodiversity research A new, widely distributed species of the Exocelina ekari-group from West Papua (Coleoptera, Dytiscidae, Copelatinae) Helena Shaverdo1, Rawati Panjaitan2, Michael Balke3 1 Naturhistorisches Museum, Burgring 7, A-1010 Vienna, Austria 2 Department of Biology, Faculty of Scien- ces and Mathematics, State University of Papua (UNIPA), Jalan Gunung Salju Amban, Manokwari 98314, West Papua, Indonesia 3 Zoologische Staatssammlung München, Münchhausenstraße 21, D-81247 Munich, Germany and GeoBioCenter, Ludwig-Maximilians-University, Munich, Germany Corresponding author: Helena Shaverdo ([email protected]; [email protected]) Academic editor: Michael Ivie | Received 28 May 2015 | Accepted 22 November 2015 | Published 18 January 2016 http://zoobank.org/04FDA81C-59CB-4172-B3FB-FD57BFCD62E3 Citation: Shaverdo H, Panjaitan R, Balke M (2016) A new, widely distributed species of the Exocelina ekari-group from West Papua (Coleoptera, Dytiscidae, Copelatinae). ZooKeys 554: 69–85. doi: 10.3897/zookeys.554.6065 Abstract Exocelina manokwariensis sp. n. from West Papua is placed into the E. ekari-group based on the structure of its male genitalia. The new species is described, including its three subspecies, from the mainland of West Papua, Waigeo Island, Batanta and Salawati Islands, and Bomberai peninsula. An identification key to the subspecies as well as data on species distribution are provided. Keywords Coleoptera, Dytiscidae, Exocelina, new species, new subspecies, molecular phylogenetics, West Papua Introduction Of the 88 species of the genus Exocelina Broun, 1886 described from New Guinea, only eight species are known from West Papua, and all of them belong to the E. -
Papuadessus Baueri Spec. Nov. from Biak Island, Papua 283-288 ©Zoologische Staatssammlung München/Verlag Friedrich Pfeil; Download
ZOBODAT - www.zobodat.at Zoologisch-Botanische Datenbank/Zoological-Botanical Database Digitale Literatur/Digital Literature Zeitschrift/Journal: Spixiana, Zeitschrift für Zoologie Jahr/Year: 2013 Band/Volume: 036 Autor(en)/Author(s): Balke Michael, Warikar Evi, Toussaint Emmanuel F. A., Hendrich Lars Artikel/Article: Papuadessus baueri spec. nov. from Biak Island, Papua 283-288 ©Zoologische Staatssammlung München/Verlag Friedrich Pfeil; download www.pfeil-verlag.de SPIXIANA 36 2 283-288 München, Dezember 2013 ISSN 0341-8391 Papuadessus baueri spec. nov. from Biak Island, Papua (Coleoptera, Dytiscidae, Hydroporinae) Michael Balke, Evi Warikar, Emmanuel F. A. Toussaint & Lars Hendrich Balke, M., Warikar, E., Toussaint, E. F. A. & Hendrich, L. 2013. Papuadessus baueri spec. nov. from Biak Island, Papua (Coleoptera, Dytiscidae, Hydroporinae). Spixi- ana 36 (2): 283-288. Papuadessus baueri spec. nov. is decribed from Biak Island, Papua. The phyloge- netic analysis of DNA sequence data suggested placement in that genus which otherwise contains P. pakdjoko Balke, 2001, a species widespread across mainland New Guinea. The new species seems to be endemic to Biak where it was collected from a limestone sinkhole. Important species characters (habitus, median lobe and paramere) are illustrated, and the habitat of P. baueri spec. nov. and its water beetle coenosis are briefly outlined. Michael Balke (corresponding author), Lars Hendrich, Emmanuel F. A. Tous- saint, Zoologische Staatssammlung München, Münchhausenstraße 21, 81247 Mün- chen, Germany; email: [email protected] Evi Warikar, Department of Biology, Universitas Cendrawasih, Jayapura, Papua, Indonesia Introduction Material and methods Papuadessus was described by Balke (2001) for the The specimens included in this study are deposited in large and conspicuous New Guinea species P.