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Phylogeny, Biogeography, and Revision of the Subfamily Dallatorrellinae (Hymenoptera, Liopteridae) Zhiwei Liu Eastern Illinois University, [email protected]

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Phylogeny, Biogeography, and Revision of the Subfamily Dallatorrellinae (Hymenoptera, Liopteridae) Zhiwei Liu Eastern Illinois University, Zliu@Eiu.Edu View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by Crossref Eastern Illinois University The Keep Faculty Research & Creative Activity Biological Sciences January 2001 Phylogeny, biogeography, and revision of the subfamily Dallatorrellinae (Hymenoptera, Liopteridae) Zhiwei Liu Eastern Illinois University, [email protected] Follow this and additional works at: http://thekeep.eiu.edu/bio_fac Part of the Biology Commons Recommended Citation Liu, Zhiwei, "Phylogeny, biogeography, and revision of the subfamily Dallatorrellinae (Hymenoptera, Liopteridae)" (2001). Faculty Research & Creative Activity. 200. http://thekeep.eiu.edu/bio_fac/200 This Article is brought to you for free and open access by the Biological Sciences at The Keep. It has been accepted for inclusion in Faculty Research & Creative Activity by an authorized administrator of The Keep. For more information, please contact [email protected]. PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 3353, 23 pp., 44 ®gures December 31, 2001 Phylogeny, Biogeography, and Revision of the Subfamily Dallatorrellinae (Hymenoptera: Liopteridae) ZHIWEI LIU1,2 ABSTRACT A recent cladistic study suggested that the parasitoid wasp subfamily Dallatorrellinae is a monophyletic clade within the cynipoid family Liopteridae. The subfamily currently includes only four known species belonging to two genera, i.e., the monotypic genus Mesocynips Cam- eron, 1903 and the genus Dallatorrella Kieffer, 1911. The present study presents a revision of the subfamily, including descriptions of ®ve new species in the genus Dallatorrella: D. maculata, D. maxima, D. pulla, D. ronquisti, and D. sinica. A key to the species of the subfamily is provided. Cladistic analysis of the subfamily at species level is undertaken and the biogeographic history of the subfamily Dallatorrellinae is reconstructed with respect to phylogeny. Contrary to an earlier assumption that the Dallatorrellinae originated in the Aus- tralian region, the phylogeny indicates that the Dallatorrellinae originated in southeastern Asia and dispersed to Australia at a later date. Consequently, the divergence between the Dallator- rellinae and the stem species of two other liopterid subfamilies, Liopterinae and Oberthuer- ellinae, corresponds with the separation between Laurasia and Gondwanaland. INTRODUCTION letic basal branches of the Cynipoidea. Their The subfamily Dallatorrellinae belongs to species are typically large in size. Although the cynipoid family Liopteridae, together the macrocynipoids are generally known as with Austrocynipidae and Ibaliidae making parasitoids of wood- or cone-boring insect up the macrocynipoids (Ronquist, 1995a). larvae (Ronquist, 1999), the biology of the The macrocynipoid families are the paraphy- Dallatorrellinae is virtually unknown. Fe- 1 Division of Invertebrate Zoology, American Museum of Natural History. 2 Present address: Division of Insects, Field Museum of Natural History, Chicago, IL 60605. Copyright q American Museum of Natural History 2001 ISSN 0003-0082 2 AMERICAN MUSEUM NOVITATES NO. 3353 males of one species of Dallatorrella have ies. Ronquist (1995b) revised the liopterid been collected on logs of Syzygium (Myrta- genera and studied phylogenetic relation- ceae) in Papua New Guinea, suggesting an ships among them. Despite this encouraging association with wood-boring insects on development in liopterid systematics, revi- broadleaved trees. Such an association is sions at the species level are still lacking, a characteristic of all other liopterids with situation common for cynipoid groups. known hosts (Ronquist, 1995a; Liu, personal The genus Dallatorrella currently includes observation). three named species. Ronquist (1995b) re- Dallatorrellinae as currently understood ferred to ®ve additional undescribed species, includes two genera: Dallatorrella Kieffer, of which two are considered as conspeci®c 1911 and Mesocynips Cameron, 1903 (Ron- here. An undescribed species was also sub- quist, 1995b). The subfamily was ®rst estab- sequently discovered from China. The genus lished by Kieffer (1911) for his new genus Mesocynips is monotypic. Dallatorrella and was originally assigned to The present study provides a revision of the family Cynipidae as then understood. Dallatorrellinae, including a key to and di- Hedicke and Kerrich (1940) placed Dallator- agnoses of all known species, analyzes the rellinae and several other groups previously phylogenetic relationships among these spe- considered as subfamilies within the Cyni- cies, and attempts to reconstruct the histori- pidae or Figitidae in a separate family, Liop- cal biogeography of the subfamily. The con- teridae. In that paper, Kerrich included Dal- cepts of the subfamily and the genera studied latorrellinae and the previously recognized here follow that of Ronquist (1995b), which Mayrellinae Hedicke, 1922 in a new subfam- should be consulted for detailed descriptions ily, Mesocynipinae, consisting of ®ve genera, of the higher taxa. namely Mesocynips Cameron, 1903, Par- amblynotus Cameron, 1908, Dallatorrella MATERIAL AND METHODS Kieffer, 1911, Mayrella Hedicke, 1922, and Paribalia Weld, 1922. The classi®cation re- SPECIMENS mained unchanged until recently. Based on a The present paper is based on 34 speci- phylogenetic analysis of the family, Ronquist mens of Dallatorrellinae borrowed from the (1995b) revised the Liopteridae and split the world's major insect collections given below Mesocynipinae once again into the two sub- through the kindness of persons listed. Ho- families recognized prior to Hedicke and lotypes of all known species were studied ex- Kerrich (1940), i.e., Mayrellinae and Dalla- cept that of Cynips insignis Smith, 1903, torrellinae. Phylogenetically, Mayrellinae is which is believed to be lost (see below). the basal clade of Liopteridae and is the sister group to all other liopterids, whereas Dalla- LIST OF DEPOSITORIES torrellinae is the sister clade of Oberthuerel- linae and Liopterinae (®g. 44). AEI American Entomological Institute, Biogeographically, Mesocynips has been Gainesville, FL, USA (D. Wahl) reported only from Southeast Asia while spe- ANIC Australian National Insect Collection, cies of Dallatorrella are distributed in both Canberra, Australia (I. Naumann) the Australian and Oriental regions. Recon- BPMB Bernice P. Bishop Museum, Honolulu, structions of historical biogeography for the HI, USA (K. Arakaki) Liopteridae suggested that the Dallatorrelli- MCZ Museum of Comparative Zoology, Har- nae originated in southern Gondwanaland vard University, Cambridge, MA, USA and subsequently dispersed into the Orient (P. Perkins) (Ronquist, 1995b). However, this hypothesis NHM The Natural History Museum, London, has not been tested by a phylogenetic recon- UK (T. Huddleston, S. Lewis) USNM Smithsonian Institution, Washington, struction of relationships within the subfam- DC, USA (D. Smith) ily. ZICA Zoological Institute, Chinese Academy Taxonomic work on Liopteridae in general of Sciences, Beijing (X. Yang) and on Dallatorrellinae in particular has been ZLMU Zoological Laboratory, Meijo Univer- almost entirely restricted to alpha-level stud- sity, Nagoya, Japan (K. Yamagishi) 2001 LIU: DALLATORRELLINAE 3 METHODS and Dallatorrella have recently been pre- sented by Ronquist (1995b), I limit the de- Phylogenetic analysis was carried out us- scriptions of taxa, including the new species, ing parsimony as implemented in the com- to the minimum. Instead, I provide a detailed puter package NONA 2.0 (Goloboff, 1999). key to species along with species diagnoses, Search options were: hold maximum number supplemented by ®gures and SEM images. of trees, i.e., 100,000 trees, and mult*1000. Characters with multiple states are treated as SYSTEMATICS ordered when a trend is obvious, and they are so indicated in the character list. All char- SUBFAMILY DALLATORRELLINAE acters were weighted equally. Liopteron and KIEFFER Paramblynotus, whose phylogenetic relation- ship with Dallatorrellinae are shown in ®gure Dallatorrellinae Kieffer, 1911: 244. Type genus: 44, were chosen as outgroups and coded as Dallatorrella Kieffer, 1911. RESTABLISHED by collective units. The coding for Parambly- Ronquist (1995b: 38). notus is based on an ongoing study on the Mesocynipinae Kerrich in Hedicke and Kerrich, 1940: 178. Type genus: Mesocynips Cameron, subfamily Mayrellinae (Liu et al., in prep.), 1903. whereas coding for Liopteron is based on published information (Ronquist, 1995b) as DIAGNOSIS: Median ¯agellomeres distinct- well as on observations of three unidenti®ed ly swollen submedially to somewhat swollen species in the Cynipoidea collection of the subapically, especially ventrally. First ¯agel- American Museum of Natural History and in lomere of female with placodeal sensilla. the author's own collection. Since the rela- Posterolateral pronotal margin distinctly in- tionship among the outgroups, Liopteron and cised in front of mesopleural triangle, the lat- Paramblynotus, and Dallatorrellinae is very ter deeply impressed anteriorly. Scutellum strongly supported by previous studies (Ron- laterally without impressed semilunar area. quist, 1995a, 1995b), and, because the pre- Ventral end of metapleural carina produced sent analysis focuses on the relationships into a prominent process laterad to metacoxal within the subfamily, the higher level rela- foramen. Pro- and mesotarsal claws with bas- tionship among the outgroups and Dallator- al, lamellate lobe. Laterobasal
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