Free Standard AU & NZ Shipping For All Book Orders Over $80!
Register      Login
Invertebrate Systematics Invertebrate Systematics Society
Systematics, phylogeny and biogeography
RESEARCH ARTICLE

Systematics of the Apteropanorpidae (Insecta : Mecoptera) based on morphological and molecular evidence

Christopher M. Palmer A B C D , John W. H. Trueman A and David K. Yeates B
+ Author Affiliations
- Author Affiliations

A School of Botany and Zoology, The Australian National University, Canberra ACT 0200, Australia.

B CSIRO Entomology, GPO Box 1700, Canberra ACT 2601, Australia.

C Present address: Biodiversity Conservation South, Parks and Wildlife Service, Department of Natural Resources, Environment, and the Arts, PO Box 1120, Alice Springs NT 0871, Australia.

D Corresponding author. Email: christopher.palmer@nt.gov.au

Invertebrate Systematics 21(6) 589-612 https://doi.org/10.1071/IS07014
Submitted: 20 April 2007  Accepted: 9 November 2007   Published: 2007

Abstract

Apteropanorpa Carpenter, 1941 is a genus of scorpionflies endemic to Tasmania. The genus comprised two described species before the current study. However, many anecdotal reports suggested that Apteropanorpa was more widespread in Tasmania than had previously been thought, and that more species awaited discovery and description. Intensive field surveys for the Apteropanorpidae were conducted from 2001 to 2003 in a range of altitudes and habitats all over Tasmania. These surveys yielded a large number of adult specimens, collected at many new localities and at a range of elevations. Cladistic analyses of COI molecular and morphological data were congruent in their inferred species composition of the family. Phylogenetic analyses with evaluation of species concepts provided evidence for two new species: Apteropanorpa warra, sp. nov. and A. hartzi, sp. nov. The distribution of the Apteropanorpidae is much more extensive than was previously thought; many populations discovered as part of this research represent extensions of the known ranges of A. evansi Byers & Yeates and A. tasmanica Carpenter. A key to species is presented. The altitudinal range of each species is closely associated with the local climatic treeline. Hypotheses concerning the biogeography of the family are consistent with the glacial history of Tasmania.

Additional keywords: scorpionflies, Apteropanorpa, taxonomy, new species, population, distribution, phylogeny, biogeography, glaciation, altitude, speciation, Tasmania, Australia, morphology, mtDNA.


Acknowledgements

We thank Forestry Tasmania, the Australian National University, and the Australian Biological Resources Study for funding to assist with fieldwork, and the Biodiversity Conservation Branch of the Tasmanian Department of Primary Industries and Water (DPIW) for permits to enter and conduct research in National Parks. Curators at the following institutions provided loans of insects in their care: the Australian Museum (Sydney), DPIW (Hobart), Forestry Tasmania (Hobart), and the Queen Victoria Museum and Art Gallery (Launceston). Christine Lambkin and Sue Baker kindly loaned specimens from their own field collections. Dick Bashford (Forestry Tasmania) and Niall Doran (DPIW) allowed unlimited access to the pitfall trapped specimens collected on Mt Weld and at the Warra Long-term Ecological Research (LTER) site. Dick Bashford also provided field equipment and laboratory space in Hobart, and facilitated access to the Warra LTER site. One of us (CMP) would especially like to thank Owen Seeman and Helen Nahrung for assistance with accommodation whilst in Hobart, and for assistance with fieldwork. The following people also assisted with accommodation: Sue Baker, Lisa Boutin, and Bryan and Vicki Evans. Thanks are also due to David Morris for kind assistance in the genetics laboratory, and Christine Lambkin assisted with the computational aspects of phylogenetic analyses. Peter McQuillan (University of Tasmania) provided very helpful comments on an earlier draft of this manuscript. This investigation was conducted in partial fulfilment of a PhD for CMP, whilst in receipt of an Australian Postgraduate Award.


References


Avise J. C. (1994). ‘Molecular Markers, Natural History and Evolution.’ (Chapman and Hall: New York, USA.)

Avise J. C. (2000). ‘Phylogeography: the History and Formation of Species.’ (Harvard University Press: Cambridge, UK.)

Barrows T. T., Stone J. O., Fifield L. K., Cresswell R. G. (2002) The timing of the last glacial maximum in Australia. Quaternary Science Reviews 21, 159–173.
Crossref | GoogleScholarGoogle Scholar | open url image1

Bremer K. (1988) The limits of amino acid sequence data in angiosperm phylogenetic reconstruction. Evolution 42, 795–803.
Crossref | GoogleScholarGoogle Scholar | open url image1

Bremer K. (1994) Branch support and tree stability. Cladistics 10, 295–304.
Crossref | GoogleScholarGoogle Scholar | open url image1

Brühl C. A. (1997) Flightless insects: a test case for historical relationships of African mountains. Journal of Biogeography 24, 233–250.
Crossref | GoogleScholarGoogle Scholar | open url image1

Byers G. W. (1965). Families and genera of Mecoptera. In ‘Proceedings of the XII International Congress of Entomology, 8–16 July 1964’. (Ed. P. Freeman.) p. 123. (Royal Entomological Society of London, London, UK.)

Byers G. W. (1991). Mecoptera (scorpion flies, hanging-flies). In ‘The Insects of Australia: a Textbook for Students and Research Workers, Vol. 2’. 2nd edn. (Ed. CSIRO.) pp. 696–704. (Melbourne University Press: Melbourne, Australia.)

Byers G. W., Yeates D. K. (1999) A second species of Apteropanorpa Carpenter from Tasmania (Mecoptera: Apteropanorpidae). Australian Journal of Entomology 38, 60–65.
Crossref | GoogleScholarGoogle Scholar | open url image1

Carpenter F. M. (1941). new genus of Mecoptera from Tasmania. Papers and Proceedings of the Royal Society of Tasmania 1940, 51–53.

Carver M. , Gross G. F. , and Woodward T. E. (1991). Hemiptera (bugs, leafhoppers, cicadas, aphids, scale insects etc.). In ‘The Insects of Australia: a Textbook for Students and Research Workers, Vol. 1’. 2nd edn. (Ed. CSIRO.) pp. 429–509. (Melbourne University Press: Melbourne, Australia.)

Colhoun E. A. (1985) Glaciations of the West Coast Range, Tasmania. Quaternary Research 24, 39–59.
Crossref | GoogleScholarGoogle Scholar | open url image1

Colhoun E. A., Hannan D., Kiernan K. (1996) Late Wisconsin glaciation of Tasmania. Papers and Proceedings of the Royal Society of Tasmania 130(2), 33–45. open url image1

Cracraft J. (1983) Species concepts and speciation analysis. Current Ornithology 1, 159–187. open url image1

de Queiroz K. (1998). The general lineage concept of species, species criteria, and the process of speciation: a conceptual unification and terminological recommendations. In ‘Endless Forms: Species and Speciation’. (Eds D. J. Howard and S. H. Berlocher.) pp. 57–75. (Oxford University Press: New York, USA.)

Derbyshire E. (1972) Pleistocene glaciation of Tasmania: review and speculations. Australian Geographical Studies 10, 79–84. open url image1

Donoghue M. J. (1985) A critique of the biological species concept and recommendations for a phylogenetic alternative. The Bryologist 88(3), 172–181.
Crossref | GoogleScholarGoogle Scholar | open url image1

Doran N. E., Balmer J., Driessen M., Bashford R., Grove S., Richardson A. M. M., Griggs J., Ziegeler D. (2003) Moving with the times: baseline data to gauge future shifts in vegetation and invertebrate altitudinal assemblages due to environmental change. Organisms, Diversity and Evolution 3, 127–149.
Crossref | GoogleScholarGoogle Scholar | open url image1

Eldredge N. , and Cracraft J. (1980). ‘Phylogenetic Patterns and the Evolutionary Process.’ (Columbia University Press: New York, USA.)

Evans J. W. (1942) A mecopterous larva from Tasmania and notes on the morphology of the insect head. Papers and Proceedings of the Royal Society of Tasmania 1941, 31–35. open url image1

Felsenstein J. (1985) Confidence limits on phylogenies: an approach using the bootstrap. Evolution 39(4), 783–791.
Crossref | GoogleScholarGoogle Scholar | open url image1

Folland C. K. , Karl T. R. , Christy J. R. , Clarke R. A. , Gruza G. V. , Jouzel J. , Mann M. E. , Oerlemans J. , Salinger M. J. , and Wang S.-W. (2001). Observed climate variability and change. In ‘Climate Change 2001: The Scientific Basis. Contribution of Working Group I to the Third Assessment Report of the Intergovernmental Panel on Climate Change’. (Eds J. T. Houghton, Y. Ding, D. J. Griggs, M. Noguer, P. J. van der Linden, X. Dai, K. Maskell and C. A. Johnson.) pp. 93–181. (Cambridge University Press: Cambridge, UK.)

Greenslade P. J. (1991). Collembola (springtails). In ‘The Insects of Australia: a Textbook for Students and Research Workers, Vol. 1’. 2nd edn. (Ed. CSIRO.) pp. 252–264. (Melbourne University Press: Melbourne, Australia.)

Hansen B., Richardson A. M. M. (2002) Geographic ranges, sympatry and the influence of environmental factors on the distribution of species of an endemic Tasmanian freshwater crayfish. Invertebrate Systematics 16, 621–629.
Crossref | GoogleScholarGoogle Scholar | open url image1

Johns G., Avise J. C. (1998) A comparative summary of genetic distances in the vertebrates from the mitochondrial cytochrome b gene. Molecular Biology and Evolution 15, 1481–1490.
PubMed |
open url image1

Key K. H. L. (1991) On four endemic genera of Tasmanian Acrididae (Orthoptera). Invertebrate Taxonomy 5(2), 241–288.
Crossref | GoogleScholarGoogle Scholar | open url image1

Kiernan K. (1990) The extent of Late Cenozoic glaciation in the Central Highlands of Tasmania, Australia. Arctic and Alpine Research 22(4), 341–354.
Crossref | GoogleScholarGoogle Scholar | open url image1

Kiernan K. (1999) The southern margin of the Late Cainozoic ice cap on the Central Plateau of Tasmania. The Australian Geographer 30(1), 5–33.
Crossref | GoogleScholarGoogle Scholar | open url image1

Kiernan K., Lauritzen S.-E., Duhig N. (2001) Glaciation and cave sediment aggradation around the margins of the Mt Field Plateau, Tasmania. Australian Journal of Earth Sciences 48, 251–263.
Crossref | GoogleScholarGoogle Scholar | open url image1

Kirkpatrick J. B., Fowler M. (1998) Locating likely glacial forest refugia in Tasmania using palynological and ecological information to test alternative climatic models. Biological Conservation 85, 171–182.
Crossref | GoogleScholarGoogle Scholar | open url image1

Maddison W. P. , and Maddison D. R. (1999). ‘MacClade: Analysis of Phylogeny and Character Evolution. Version 3.08a.’ (Sinauer Associates: Sunderland, MA, USA.)

Mallet J. (1995) A species definition for the modern synthesis. Trends in Ecology and Evolution 10(7), 294–299.
Crossref | GoogleScholarGoogle Scholar | open url image1

Mani M. S. (1968). ‘Ecology and Biogeography of High Altitude Insects.’ (Dr W. Junk N. V.: The Hague, The Netherlands.)

McQuillan P. B. (1986). Trans-Tasman relationships in the highland moth (Lepidoptera) fauna. In ‘Flora and Fauna of Alpine Australia’. (Ed. B. A. Barlow.) pp. 263–276. (CSIRO Publishing: Melbourne, Australia.)

Mesibov R. (1994) Faunal breaks in Tasmania and their significance for invertebrate conservation. Memoirs of the Queensland Museum 36(1), 133–136. open url image1

Mesibov R. (1999). The Mersey Break: an unexplained faunal boundary on the north coast of Tasmania. In ‘The Other 99%: The Conservation and Biodiversity of Invertebrates’. (Eds W. Ponder and D. Lunney.) pp. 246–252. (Royal Zoological Society of New South Wales: Sydney, Australia.)

Mickoleit G. (1975) Die genital- und postgenitalsegmente der Mecoptera - Weibchen (Insecta, Holometabola). I. Das exoskelet. Zeitschrift fuer Morphologie der Tiere 80, 97–135. open url image1

Mickoleit G. (1976) Die genital- und postgenitalsegmente der Mecoptera - Weibchen (Insecta, Holometabola). II. Das dach der ganitalkammer. Zoomorphologie 85, 133–156.
Crossref | GoogleScholarGoogle Scholar | open url image1

Morris D. C., Mound L. A. (2004) Molecular relationships between populations of South African citrus thrips (Scirtothrips aurantii Faure) in South Africa and Queensland, Australia. Australian Journal of Entomology 43, 353–358.
Crossref | GoogleScholarGoogle Scholar | open url image1

Morris D. C., Schwarz M. P., Crespi B. J., Cooper S. J. B. (2001) Phylogenetics of gall-inducing thrips on Australian Acacia.  Biological Journal of the Linnean Society 74, 73–86.
Crossref | GoogleScholarGoogle Scholar | open url image1

Nielsen E. S., McQuillan P. B., Common I. F. B. (1992) The Tasmanian cushion plant moth Nemotyla oribates gen. n., sp. n.: systematics and biology (Lepidoptera: Oecophoridae: Xyloryctinae). Journal of the Australian Entomological Society 31, 47–56.
Crossref | GoogleScholarGoogle Scholar | open url image1

Nixon K. C., Wheeler Q. D. (1990) An amplification of the phylogenetic species concept. Cladistics 6, 211–223. open url image1

Nunez M. (1988). regional lapse rate for Tasmania. Papers and Proceedings of the Royal Society of Tasmania 122 (2), 53–57.

Ollier C. D. (1986). The origin of alpine landforms in Australasia. In ‘Flora and Fauna of Alpine Australia’. (Ed. B. A. Barlow.) pp. 3–26. (CSIRO Publishing: Melbourne, Australia.)

Penny N. D. (1975) Evolution of the extant Mecoptera. Journal of the Kansas Entomological Society 48(3), 331–350. open url image1

Penny N. D., Byers G. W. (1979) A Checklist of the Mecoptera of the world. Acta Amazonica 9(2), 365–388. open url image1

Peñuelas J., Filella I. (2001) Responses to a warming world. Science 294, 793–795.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Peterson A. T., Soberón J., Sanchez-Cordero V. (1999) Conservatism of ecological niches in evolutionary time. Science 285, 1265–1267.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Ponder W. F., Colgan D. J. (2002) What makes a narrow-range taxon? Insights from Australian freshwater snails. Invertebrate Systematics 16, 571–582.
Crossref | GoogleScholarGoogle Scholar | open url image1

Pounds J. A., Fogden M. P. L., Campbell J. H. (1999) Biological response to climate change on a tropical mountain. Nature 398, 611–615.
Crossref | GoogleScholarGoogle Scholar | open url image1

Ricklefs R. E., Latham R. E. (1992) Intercontinental correlation of geographical ranges suggests stasis in ecological traits of relict genera of temperate herbs. American Naturalist 139, 1305–1321.
Crossref | GoogleScholarGoogle Scholar | open url image1

Riek E. F. (1973) A revision of Australian scorpion flies of the family Choristidae (Mecoptera). Journal of the Australian Entomological Society 12, 103–112.
Crossref | GoogleScholarGoogle Scholar | open url image1

Roehrdanz R. L. (1993) An improved primer for PCR amplification of mitochondrial DNA in a variety of insect species. Insect Molecular Biology 2, 89–91.
PubMed |
open url image1

Root T. L., Price J. T., Hall K. R., Schneider S. H., Rosenzweig C., Pounds J. A. (2003) Fingerprints of global warming on wild animals and plants. Nature 421, 57–60.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Shiel R. J., Koste W., Tan L. W. (1989) Tasmania revisited: rotifer communities and habitat heterogeneity. Hydrobiologia 186/187, 239–245.
Crossref | GoogleScholarGoogle Scholar | open url image1

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. open url image1

Sorenson M. D. (1999). ‘TreeRot. Version 2.’ (Boston University: Boston, MA, USA.)

Sunnucks P., Hales D. F. (1996) Numerous transposed sequences of mitochondrial cytochrome oxidase I–II in aphids of the genus Sitobion (Hemiptera: Aphididae). Molecular Biology and Evolution 13(3), 510–524.
PubMed |
open url image1

Swofford D. (2002). ‘PAUP*. Phylogenetic Analysis Using Parsimony. Version 4.0b10.’ (Sinauer Associates: Sunderland, MA, USA.)

Thomas C. D., Cameron A., Green R. E., Bakkenes M., Beaumont L. J. , et al. . (2004) Extinction risk from climate change. Nature 427, 145–148.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Thomas I., Hope G. (1994) An example of Holocene vegetation stability from Camerons Lagoon, a near treeline site on the Central Plateau, Tasmania. Australian Journal of Ecology 19, 150–158. open url image1

Thompson J. D., Gibson T. J., Plewniak F., Jeanmougin F., Higgins D. G. (1997) The Clustal_X windows interface: flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Research 25(24), 4876–4882.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Tyler P. A. (1992) A lakeland from the Dreamtime: the second Founders’ Lecture. British Phycological Journal 27, 353–368.
Crossref | GoogleScholarGoogle Scholar | open url image1

Van Valen L. (1976) Ecological species, multispecies, and oaks. Taxon 25(2/3), 233–239.
Crossref | GoogleScholarGoogle Scholar | open url image1

Walther G.-R., Post E., Convey P., Menzel A., Parmesan C., Beebee T. J. C., Fromentin J.-M., Hoegh-Guldberg O., Bairlein F. (2002) Ecological responses to recent climate change. Nature 416, 389–395.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Watson J. A. L. , and O’Farrell A. F. (1991). Odonata (dragonflies and damselflies). In ‘The Insects of Australia: a Textbook for Students and Research Workers. Vol. 1’. 2nd edn. (Ed. CSIRO.) pp. 294–310. (Melbourne University Press: Melbourne, Australia.)

Whiting M. F. (2002) Mecoptera is paraphyletic: multiple genes and phylogeny of Mecoptera and Siphonaptera. Zoologica Scripta 31, 93–104.
Crossref | GoogleScholarGoogle Scholar | open url image1

Wiens J. J. (2004) Speciation and ecology revisited: phylogenetic niche conservatism and the origin of species. Evolution 58(1), 193–197.
PubMed |
open url image1

Willmann R. (1981) Das exoskelett der männlichen genitalien der Mecoptera (Insecta) I. Morphologie. Zeitschrift fur Zoologische Systematik und Evolutionsforschung 19(2), 96–150. open url image1










Appendix 1.  Data matrix of 36 morphological characters coded for 21 populations of Apteropanorpidae
A1



Appendix 2.  Morphological characters used in the phylogenetic analysis
Characters 5, 6 and 36 are parsimony uninformative
Click to zoom



Appendix 3.  Uncorrected pairwise genetic distance matrix of 13 populations of Apteropanorpidae and two outgroup taxa (Harpobittacus australis and Chorista australis)
Click to zoom