School of Science and Engineering Department of Environmental and Aquatic Sciences The Systematics of the Monoscutidae (Arachnida: Opiliones) Christopher Kenneth Taylor This thesis is presented for the degree of Doctor of Philosophy of Curtin University of Technology February 2010 To the best of my knowledge and belief this thesis contains no material previously published by any other person except where due acknowledgement has been made. This thesis contains no material which has been accepted for the award of any other degree or diploma in any university. Signed Christopher Taylor Date 2 Contents Abstract .......................................................................................................................... 1 Introduction .................................................................................................................... 3 Materials and Methods ................................................................................................... 8 Dorsal appearance of the enantiobunine prosoma ....................................................... 12 Phylogenetic Analysis .................................................................................................. 14 Characters used in analysis .......................................................................................... 16 Results and Discussion ................................................................................................. 37 Key to Genera of Enantiobuninae ................................................................................ 47 Taxonomic Descriptions .............................................................................................. 50 Acihasta Forster 1948a ............................................................................................. 51 Australiscutum new genus ....................................................................................... 52 Forsteropsalis new genus ........................................................................................ 66 Hypomegalopsalis new genus .................................................................................. 67 Intutoportula new genus .......................................................................................... 70 Megalopsalis Roewer 1923 ...................................................................................... 83 Monoscutum Forster 1948a .................................................................................... 103 Neopantopsalis new genus ..................................................................................... 104 Scissorestis new genus ........................................................................................... 124 Spinibunus new genus ............................................................................................ 125 Spinicrurellum new genus ...................................................................................... 129 Spinicruroides new genus ...................................................................................... 134 Spinicrus Forster, 1949a ........................................................................................ 138 Templar new genus ................................................................................................ 145 Tercentenarium new genus .................................................................................... 147 Thrasychiroides Soares & Soares 1947 ................................................................. 153 Thrasychirus Simon 1884……...…………………………………………………154 Appendix I: Revision of the Australian Gagrellinae (Arachnida: Opiliones: Sclerosomatidae), with a description of a new species .............................................. 155 Abstract .................................................................................................................. 155 Introduction ............................................................................................................ 155 Methods .................................................................................................................. 157 Taxonomic Description .......................................................................................... 158 Acknowledgements .................................................................................................... 165 References .................................................................................................................. 166 i Tables 1. Character state matrix for phylogenetic analysis……………………………. 32–36 Figures 1. Dorsal view of Enantiobuninae exemplars, showing division of propeltidium into three areas…...……………………………………………………………………... 13 2–7. Genitalia of Enantiobuninae and Dyspnoi, SEM……………………………. 18 8–11. Close-up images of enantiobunine glans to show differing pore morphologies……………………………………………………………………….. 19 12–15. Spiracles of outgroup taxa…………………………………………………. 21 16–19. Neopilionid spiracles………………….…………………………………… 22 20–23. Spiracles of Thrasychirus and Pantopsalis………………………………… 23 24–27. Spiracles of Spinicrus species……………………………………………… 24 28–31. Spiracles of Neopantopsalis and New Zealand Megalopsalis…………… 25 32–35. Spiracles of Western Australian enantiobunines………………………… 26 36–39. Spiracles of Megalopsalis species…………………………………………. 26 40–41. Modified distitarsi in Megalopsalis species……………………………… 30 42. Strict consensus tree from parsimony analysis………………………………… 38 43. Strict consensus, with mapped synapomorphies……………………………… 39 44. Strict consensus of eight most parsimonious trees resulting from analysis constrained for monophyly of Monoscutidae……………………………………… 40 45. Distribution map for Australiscutum species………………………………… 53 46–49. Australiscutum hunti new species ………………………………………… 56 50. Male chelicerae of Australiscutum species…………………………………… 59 51–57. Genitalia of Australiscutum species……………………………………… 61 58–63. Hypomegalopsalis tanisphyros new species, male………………………… 68 64. Locality map for Intutoportula species in southern Western Australia……… 71 65–68. Spiracles of Intutoportula species………………………………………….. 73 69–74. Intutoportula minima, major male…………………………………………. 74 75–79. Intutoportula porongorupensis, male……………………………………… 76 80–83. Intutoportula suffugiens new species, male………………………………... 78 84-86. Intutoportula walpolensis new species, male………………………………. 81 87. Distribution map for Megalopsalis and Hypomegalopsalis……………………. 83 88–94. Megalopsalis serritarsus, male…………………………………………….. 86 95–100. Megalopsalis epizephyros new species, male…………………………… 89 101–107. Megalopsalis eremiotis new species…………………………………… 93 108–116. Megalopsalis hoggi, male……………………………………………… 96 117–122. Megalopsalis leptekes new species, male……………………………… 99 123–128. Megalopsalis pilliga new species, male………………………………... 102 129. Localities of Neopantopsalis in north-eastern Australia…………………….. 104 130–137. Neopantopsalis quasimodo new species, male……………………........ 109 138–139. Neopantopsalis quasimodo new species, male……………………........ 110 140–143. Neopantopsalis camelus………………………………………………... 112 144–150. Neopantopsalis pentheter new species………………………………. 116 151–156. Neopantopsalis psile new species, male……………………………….. 119 157–158. Neopantopsalis psile new species, major male, SEM images…………. 120 159. Neopantopsalis psile new species, body of female, dorsal view……………. 121 160–163. Neopantopsalis thaumatopoios new species, male…………………….. 123 ii 164. Locality map for Spinicrus and other Enantiobuninae in eastern Australia… 126 165–169. Spinibunus atrocidiana new species………………………………........ 127 170–173. Spinibunus atrocidiana new species and Spinicrurellum coronatum new species, SEM images……………………………………………………………... 128 174-178. Spinicrurellum coronatum new species, male………………………….. 131 179-182. Spiracles of Spinicrurellum species…………………………………….. 132 183-186. Spinicrurellum puerile new species, male…………………………........ 133 187–192. Spinicruroides caeruleomontium new species…………………………. 136 193-194. Spinicruroides caeruleomontium new species, spiracle………………... 138 195–197. Spinicrus stewarti, male genitalia…………………………………........ 141 198–200. Spiracles of Spinicrus species………………………………………… 142 201–204. Spinicrus sublucens new species, male………………………………… 143 205–212. Templar incongruens new species…………………………………….. 146 213–219. Tercentenarium linnaei new species…………………………………… 149 220–221. Tercentenarium linnaei new species, right male pedipalp…………….. 150 222–224. Tercentenarium linnaei new species, genitalia………………………… 151 225-226. Tercentenarium linnaei new species, penis…………………………….. 152 227-231. Gagrella cauricrepa new species………………………………………. 160 iii Abstract The taxonomy of the Australian species of the family Monoscutidae is reviewed, and a phylogenetic parsimony analysis is conducted of Monoscutidae and related taxa. As a result of this analysis, Monoscutidae (including subfamilies Monoscutinae and Megalopsalidinae) is synonymised with the subfamily Enantiobuninae of the family Neopilionidae. All Australian species of Phalangioidea now belong to Neopilionidae with the exception of Gagrella cauricrepa new species (Sclerosomatidae) which is described from Cape York Peninsula, Queensland. Ten new genera and twenty-one new species of Enantiobuninae are described, including Scissorestis nigricans (Hickman 1957) new genus and combination, Templar incongruens new genus and species (New Zealand), Tercentenarium linnaei new genus and species, Hypomegalopsalis tanisphyros new genus and species, Megalopsalis epizephyros new species,