Èn.tt .Õò SYSTEMATICS AND BIOGEOGRAPHY OF HYPTIOGASTRINE WASPS (HYMENOPTERA: GASTERUPTIIDAE) Pseudoþenus spinitarsru (Westwood) comb. nov [Photo: G. V/eber] JOHN T. JENNINGS B. Sc. (Hons.) (Flinders) A thesis submitted for the Degree of Doctor of Philosophy in the Faculty of Agricultural and Natural Resource Sciences at The University of Adelaide September 1999 To my parents John and Erica qþ Table of contents Page Summary Y Declaration vll I Acknowledgments ix Chapter 1. Introduction 1 Chapter 2. Systematics and distribution of Evanioidea 5 2.1. Introduction 6 2.2. Systematics 6 2.3. Distribution T2 Chapter 3. Synopsis of aulacid and gasteruptiid biology L4 3.1. Oviposition 15 3.2. Mode of development T7 3.3. Larvae 19 3.4. Pupation and emergence I9 3.5. Adults 20 3.6. Phenology 24 3.7. Host relationships 25 3.1.1. Aulacidae 25 3.7.2. Gasteruptiidae 27 Figures 3.1-3.2 Chapter 4. Materials and Methods 31 4.1. Microscopy 32 4.2. Terminology 32 4.3. Abbreviations 32 4.4. Localities 34 4.5. Phylogenetic analyses 34 4.51. Software and computing 34 4.5.2. Treatment of characters 35 4.5.3. Options in parsimony analyses 35 4.5.4. Measures of tree fitness 38 Chapter 5. Morphology of Gasteruptiidae 40 5.1. Head 4t 5.2. Mesosoma 42 5.3. V/ings 42 5.4. Legs 43 5.5. Metasoma 43 5.6. Genitalia 43 Figures 5.1-5.1 1 Chapter 6. Phylogeny of the Hyptiogastrinae 44 6. 1. Introduction 45 6.2. Monophyly of the Hyptiogastrinae and outgroup determination 45 6.3. Character selection 46 6.4. Preliminary phylogenetic analyses 51 6.4.1. Test for the monophyly of families and subfamilies 51 6.4.2. Characters weighted 52 6.4.3. Discussion 53 6.5. Relationships within the Hyptiogastrinae 54 6.5,1. Additional characters included in the phylogenetic analyses 55 6.5.2. Results of analYses 58 6.5.2.I. Heuristic search 58 6.5.2.2. Bootstrap and Bremer support values 59 6.5.2.3. Successive weighting 59 6.5.3. Analyses with constraints 60 6.5.4. New classification for the Hyptiogastrinae 6û 6.5.4.I. Species-groups of Pseudoþenus sensu lato Ytteffer 6l Figures 6.1-6.4 Chapter 7. BiogeographY 64 7. 1. Biogeographical principles 65 7.2. Biogeography of the Hyptiogastrinae 68 7 .2.I. Distribution of hyptiogastrines in Australia 68 7 .2.2. Vicariance hypothesis 68 7 .2.2.1. Phylogenetic constraints 69 7 .2.2.2. Dispersal ability of Hyptiogastrinae 10 7 .2.2.3. Hosts of Hyptiogastrinae 7l 7.2.3. Discussion 72 Figures 7.1-7.6 Chapter 8. Synopsis of Gasteruptiidae and revision of Hyptiogastrinae 74 8. 1. Introduction 15 8.2. Families of Evanioidea 75 8.3. Subfamilies of Gasteruptiidae 76 8.3.1. Diagnosis of subfamilies 16 8.4. Treatment of genera and species of Hyptiogastrinae 78 8.4.1. Hyptiogaster Kieffer 79 il 8.4. 1. 1. Included species 82 8.4.2. Pseudoþenus sensu lato Kieff.er 83 8.4.2.1. Key to species-groups and subgroups and species of Pseudoþenus sensu lato \(teffer 87 8.4.2.2. Treatment of sPecies 95 8.4.2.2.1. crassipes-species group 95 8.4.2.2.2. fletcheri-species grotp 96 8.4.2.2.2.I. thoracicas-subgroup 96 8.4.2.2.2.2. mac ronyx-subgrouP 98 8.4.2.2.2.3. infumatus -sub group 98 8.4.2.2.3. Species not assigned to species groups rt2 8.5.2.2.4. Species incertae sedis 181 Figures 8.1 - 8.198 Chapter 9. General discussion 184 References 188 Appendices 209 A.1. Biology and host relationships A1 . 1. Recorded plant association s for P ristaulacus (Aulacidae) 2r0 AI.2. Records of flowers visited by adult Gasteruptiidae 2t2 41.3. Host records of Aulacidae 2t8 41.4. Host records of Gasteruptiidae 223 Phylogeny ^2.A2.1. Data matrix for preliminary analyses (Section 6.4) 229 A2.2. Extended data matrix used in Section 6.5.2. 231 A2.3. Segment coding for character 59 232 Segment coding for character 60 233 ^2.4.A2.5. Segment coding for character 62 234 A2.6. Segment coding for character 63 235 A2.7. Segment coding for character 64 236 A2.8. Segment coding for character 65 231 A2.9. Segment coding for cha¡acter 66 238 A2.10. Segment coding for character 67 239 ilt 43.-48. Publications. Included at the end of the thesis 43. Jennings, J. T., and Austin, A. D. 1994a. Revision of the genus Crassifoenus Crosskey (Hymenoptera: Gasteruptiidae: Hyptiogastrinae), with a description of a new species from'Western Australia. Records of the Western Australian Museum 16,575-91. 44. Jennings, J. T., and Austin, A. D. I994b. Revision of. Pseudofoenus Kieffer (Hymenoptera: Gasteruptiidae), a hyptiogastrine rwasp genus endemic to New Zealand. Invertebrate Taxonomy 8' 1289-1303. 45. Jennings, J. T., and Austin, A. D. 1997a. Revision of the Australian endemic genus Hyptiogaster Kieffer (Hymenoptera: Gasteruptiidae), with descriptions of seven new specie s. Journal of Natural History 31, 1533-1562. A6. Jennings, J. T., and Austin, A. D. L991b. Revision of. Aulacofoenus Kieff.er (Hymenoptera: Gasteruptiidae), hyptiogastrine wasps with a restricted Gondwanic distribution . Invertebrate Taxonomy ll, 943-97 6. A7. Austin, A. D., Jennings, J. T., and Harvey, M.S. 1996. Case 2950' Pseudoþer¿øs Kieffer, 1902 (Insecta, Hymenoptera): proposed designation of Foenus unguiculatas'Westwood, 1841 as the type species. The Bul.l.etin of Zoological Nomenclature 53, 26I-263' 48. Austin, A. D., Jennings, J. T., and Harvey, M. S. 1991. Comments on the proposed designation of Foenus unguiculatus'Westwood, 1841 as the type species of Pseudofoenus Kieffer, 1902 (Insecta, Hymenoptera) (Case 2950). The Bulletin of Zoological Nomenclature 55' 185-187' fV Summary The Hyptiogastrinae are a Gondwanan subfamily of small to medium sized wasps that are predator-inquilines of solitary ground-nesting bees that provision their nests with pollen. Along with the Gasteruptiinae they comprise the family Gasteruptiidae. Whereas Gasteruption L, (Gasteruptiinae) is almost cosmopolitan, the Hyptiogastrinae are found mostly in Australia but also in South America, New Zealand, New Guinea and on several south-west Pacific islands. Prior to this study, the latter subfamily comprised some 37 species in five genera; Aulacofoenus Kieffer, Crassifoenus Crosskey, Eufoenus Szépligeti, Hyptiogast¿r Kieffer and Pseudoþenus Kieffer. Recent extensive collecting using techniques such as malaise traps has greatly increased the number of specimens available which have provided the basis for the present study. Overall, the project aimed to investigate the phylogenetic relationships among genera and species of Hyptiogastrinae, examine whether the current distribution of taxa can be explained bv vicariance events, and revise the taxonomy of species. A brief literature review is first provided which describes the taxonomic history of the group, and in particular the Hyptiogastrinae. This is followed by an extensive synopsis o1 the biology of gasteruptiids and a comparison with that of its sister family, the Aulacidae. This latter section is mainly based on a review of the relevant literature but also includes new results based on observations made during this study. The materials and methods used in the phylogenetic analyses and taxonomic revision are then outlined. This chapter describes the techniques used in the examination of specimens, including field emission scanning electron microscopy, and a detailed discussion of the methods adopted for phylogenetic analyses using PAUP. The fourth chapter details the morphology of the Gasteruptiidae as a prelude to dealing with the phylogenetic analyses and taxonomic revision. Two series of phylogenetic analyses were conducted; one to test relationships among families and subfamilies and the other to examine relationships within the Hyptiogastrinae. The first set of analyses employed 56 taxa and used Schlettererius V cinctipes (Cresson) (Stephanidae) as the outgroup. Of the included taxa, six exemplar species of Aulacidae were selected, three each from Aulacøs and Pristaulacus, five from Gasteruption and 42 hyptiogastrines; nine species of Aulacoþenus, thtee Crassifoeruts, 23 Eufoenus, five Hyptiogaster andtwo Pseudoþenus. A data matrix of 58 characters was compiled using MacClade (version 3.7), and was analysed using an heuristic search of 100 random replicates using PAUP with random addition sequence and with all character states unordered. The resultant strict consensus tree showed that the Aulacidae, Gasteruptiidae and Hyptiogastrinae are all monophyletic. Little resolution was achieved within the Hyptiogastrinae, although Pseudoþenus s. str. and Crassifoenus came out as monophyletic. The second series of analyses examined relationships within the Hyptiogastrinae in more detail and was based on a separate matrix of 34 informative characters; the characters used in the above analyses were re-examined and redefined where necessary, and a number of additional characters and species were included to better represent the morphological and taxonomic diversity within the subfamily. The most basal Gasteruption species in the first analysis was used as the outgroup and 51 taxa included within the ingroup. A number of important outcomes were evident. The genera Crassifoenus, Hyptiogaster and Pseudofoenus s. sfr. were shown to be monophyletic and are well supported by a number of characters. However, Crassifoenus and Pseudoþenus s. str. were contained within larger clades which encompass Aulacoþenu.s and Eufoenøs, the latter genera being polyphyletic. A new phylogenetically-based classihcation is proposed for the Hyptiogastrinae reflecting the results of a successively weighting analysis. Two monophyletic groups are recognised: Hyptiogaster and Pseudofoenus s. 1., the latter genus comprising Aulacofoenus, Crassifoenus and Eufoen¿¿s which are synonymised with it. Two monophyletic species groups are designated for Pseudoþenus s.1., with a further 30 species not assigned to any groups. The results of the phylogenetic analyses are then used to examine the biogeography of the Hyptiogastrinae, in particular whether their current distribution can VI be explained by vicariance events alone.