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January 2001 Phylogeny, biogeography, and revision of the subfamily Dallatorrellinae (, Liopteridae) Zhiwei Liu Eastern Illinois University, [email protected]

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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 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 . 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 making parasitoids of wood- or cone-boring 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 , Field Museum of Natural History, Chicago, IL 60605.

Copyright ᭧ 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 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 part of meta- rellinae is constrained accordingly in the coxa with a lateral ridge and an anterolateral, analysis. pubescent depression. Metatibial lobe pro- The resulted trees were subsequently sub- duced into distinct, slender, toothlike process jected to WinClada (Nixon, 1998) for char- (except D. maxima); petiolar annulus of fe- acter optimization. Bootstrap and Jackknife male with ventral hump. Abdominal T3±T5 values were calculated with NONA and covered with conspicuous, erect pubescence; WinClada (1000 replications, each with 100 T7 larger than T5 and T6 in lateral view and search replications). with a patch of recumbent pubescence ven- The study is entirely based on adult exter- trolaterally. nal morphology. SEM images were taken with a Hitachi S4700 ®eld emission scanning KEY TO GENERA AND SPECIES OF electron microscope (FE-SEM). Because of DALLATORRELLINAE the scarcity of available material, specimens 1. Body entirely yellow to red-brown. Flagellum were examined unprepared and uncoated long and slender, F1 about 4 times as long with emission current at 4500±7000 nA and as wide at midpoint; ¯agellomeres cylin- accelerating voltage at 1±1.5 kV. Morpholog- drical. Eyes distinctly shorter than height of ical terminology follows that proposed by malar space (®g. 1) and protruding lateral- Ronquist and Nordlander (1989) and Ron- ly, but not distinctly beyond temple (®g. 2). Clypeus only slightly emarginated anteri- quist (1995b). Terms for surface sculpturing orly (®g. 1). Foveate sculpture absent or al- follow Harris (1979). Antennal ¯agellomeres most absent from head, pronotum, and me- and abdominal terga are represented in the soscutum (®gs. 1±3). Lateral pronotal ca- text as, e.g., F1 and T3. rina meeting pronotal crest dorsally; dorsal Since detailed descriptions of Mesocynips pronotal area behind pronotal crest indis- 4 AMERICAN MUSEUM NOVITATES NO. 3353

Figs. 1±8. 1±5, 7. Mesocynips insignis. 1. Head in frontal view; 2. Head and pronotum in dorsal view; 3. Mesosoma in lateral view; 4. Posterior part of mesoscutum and mesoscutellum in dorsal view; 5. Metatibia in laterodorsal view; 7. Forewing. 6, 8: Dallatorrella albata. 6. Metatibia in laterodorsal view; 8. Forewing.

tinct (®g. 2). Parascutal carinae of meso- ward RϩSc medially (®g. 7). Procoxa lat- scutum posterolaterally not produced, erally without a vertical carina. T7±T8 smooth and broadly rounded, scutellar sul- fused ...... cus divided into two foveae by median ca- ...... Mesocynips insignis (Smith) rina, mesoscutellum sloped laterally and Ð Body usually entirely black, or at least me- posteriorly, dorsal surface transversely cos- sosoma black. Flagellum short and stout, tate and lacking foveate structure (®g. 4). F1 at most 1.7 times as long as wide at Metapleural sulcus abruptly bent at middle, midpoint; ¯agellomeres distinctly con- metepisternum subrectangular (®g. 3). stricted basally and apically (®g. 9). Eyes

MϩCu1 of forewing straight, diverging at least as long as height of malar space from RϩSc at the base and not curved to- (e.g., ®gs. 10, 11) and distinctly protruding 2001 LIU: DALLATORRELLINAE 5

Figs. 9±15. 9, 10, 12, and 14: Dallatorrella albata. 9. Female antenna. 10. Head in frontal view; 12, Mesoscutellum; 14. Metasoma in lateral view. 11, 13, 15: Dallatorrella maculata. 11. Head in frontal view; 13, Mesoscutellum; 15. Metasoma in lateral view.

laterally beyond temple (e.g., ®g. 20). area behind pronotal crest distinct (®gs. Clypeus anteriorly with a deep, broad tri- 17, 20). Parascutal carinae posterolaterally angular emargination (®gs. 10, 11, 22, 23, distinctly produced into a more or less tri- 35±37). Foveate sculpture present on low- angular process (e.g., ®gs. 12, 13, 24, 25). er face, lateral surfaces of pronotum, me- Scutellar sulcus divided into several fove- soscutum, and scutellum (e.g., ®gs. 10± ae by two or more longitudinal carinae. 13). Lateral pronotal carina not meeting Mesocutellum at least raised laterally and pronotal crest dorsally; dorsal pronotal sometimes also posteriorly; dorsal surface 6 AMERICAN MUSEUM NOVITATES NO. 3353

Figs. 16±21. 16, 18: Dallatorrella albata. 16. Vertex; 18. Head and mesosoma in lateral view. 17, 19: Dallatorrella maculata. 17. Vertex; 19. Head and mesosoma in lateral view. 20, 21: Dallatorrella maxima. 20. Head and pronotum in dorsal view; 21. Mesosoma in lateral view.

entirely foveate-reticulate or foveate-ru- Metepisternum with extra process laterad gose and without transverse costae (e.g., to metacoxal foramen, anterior to the ®gs. 12, 13). Metapleural sulcus slightly prominent process that extends from me- and evenly curved, metepisternum sub- tapleural carina (®g. 21). Apical process of

triangular (®gs. 18, 19, 21). MϩCu1 of metatibia absent or reduced, short and api- forewing run almost parallel to RϩSc ba- cally rounded (®gs. 26, 27) ...... 3 sally and curved abruptly from RϩSc api- Ð Median frontal carina not percurrent to clyp- cally (®g. 8). Procoxa laterally with a ver- eus, continuous at most to the middle of tical carina. T7±T8 not fused ...... lower face. Pronotum not expanded pos- ...... Dallatorrella,2 terolaterally. Posterolateral projection of 2. Median frontal carina distinctly percurrent to parascutal carina apically angular (®gs. 12, clypeus in lower face (®gs. 22, 23). Pron- 13, 30, 31). Mesosubpleuron in lateral otum strongly expanded posterolaterally view nearly straight, not ventrally project- (®g. 20). Posterolateral projection of para- ed into triangular process. Metepisternum scutal carina apically rounded (®gs. 24, without an extra process laterad to meta- 25). Mesosubpleuron in lateral view ven- coxal foramen, anterior to the prominent trally projected into a triangular process. process that extends from metapleural ca- 2001 LIU: DALLATORRELLINAE 7

Figs. 22±27. 22, 24, 26: Dallatorrella maxima. 22. Head in frontal view; 24. Mesoscutellum; 26. Metatibia in laterodorsal view. 23, 25, 27: Dallatorrella pulla. 23. Head in frontal view; 25. Mesoscu- tellum; 27. Metatibia in laterodorsal view.

rina. Apical process of metatibia long and 26)...... pointed apically (®gs. 32, 33) ...... 4 ..... Dallatorrella maxima, new species 3. Radiate-carinate sculpture present on almost 4. Forewing with large, contrasting macula to- entire lower face (®g. 23). Mesoscutellum ward tip. Female metasoma dorsally with transversely rugose and without distinct a band of dense hairs along posterior mar- foveae (®g. 25). Wings ferruginous. Apical gin of T6 (®gs. 14, 15) ...... 5 metatibial process present (®g. 27) . . . Ð Wings transparent or evenly ferruginous; ...... Dallatorrella pulla, new species forewing sometimes with darker areas, but Ð Radiate-carinate sculpture present only on they are never contrastingly so. Female ventral one-third of lower face (®g. 22). metasoma dorsally without a band of Mesoscutellum distinctly foveate and dense hairs along posterior margin of T6 without distinct transverse sculpture (®g. (®g. 30) ...... 6 24). Forewing with a wide band across 5. Vertex (®g. 15), lower face (®g. 9), and pro- middle and along outer margin, respective- notum (®g. 17) sparsely pubescent. Meso- ly, and hind wing with basal half transpar- scutellum distinctly narrower across scu- ent and distal half ferruginous except tip tellar sulcus than behind (®g. 12). Axillar pale. Apical metatibial process absent (®g. area without conspicuous hair tuft (®g. 8 AMERICAN MUSEUM NOVITATES NO. 3353

Figs. 28±33. 28, 30, 32: Dallatorrella ronquisti. 28. Head and mesosoma in lateral view; 30. Me- soscutellum; 32. Mesosoma and metatibia in lateral view. 29, 31, 33: Dallatorrella sinica. 29. Head and mesosoma in lateral view; 31. Mesoscutellum; 33. Metatibia in lateral view.

11). Female metasoma devoid of hairs on median line (®g. 12) ...... T3±T5, with only a band of dense hairs ...... Dallatorrella albata (Weld, 1944) dorsally along posterior margin of T6; T3 6. Face and gena orange. Lower face glabrate- only slightly longer than T4 along a lateral punctate with sparse pubescence (®g. 36). median line (®g. 13) ...... Scutellar sulcus subdivided by three or .... Dallatorrella maculata, new species more longitudinal carinae. Lateroposterior Ð Vertex (®g. 14), lower face (®g. 8), and pro- process of parascutal carina apically bent notum densely pubescent. Mesoscutellum ventrally. Metasoma bright yellow .... only slightly narrower across scutellar sul- Dallatorrella rubriventris Kieffer, 1911 cus than behind (®g. 13). Axillar area with Ð Face and gena black to dark brown. Lower conspicuous hair tuft (®g. 10). Female me- face foveate and punctate with dense pu- tasoma with more or less dense hairs on bescence (®gs. 28, 29, 35, 37). Scutellar all terga, dorsally with a band of dense sulcus subdivided by two submedian lon- hairs along posterior margin of T5±T6; T3 gitudinal carinae. Lateroposterior process distinctly longer than T4 along a lateral of parascutal carina apically not bent ven- 2001 LIU: DALLATORRELLINAE 9

Figs. 34±41. 34. Dallatorrella ronquisti, head in dorsal view. 35. Dallatorrella sinica, head in front view. 36, 38, 40: Dallatorrella rubriventris. 36. Head in frontal view; 38. Mesosoma in lateral view; 40. Metasoma in lateral view. 37, 39, 41: Dallatorrella carinifrons. 37. Head in front view; 39. Me- sosoma in lateral view; 41. Metasoma in lateral view.

trally. Metasoma brown to dark brown . . lar space (®gs. 35, 37). Mesoscutellum ...... 7 broadly rounded posteriorly (®g. 31). Me- 7. Eyes prominent, 1.5 times as long as malar tasoma dark brown to black ...... 8 space (®g. 28). Mesoscutellum nearly trun- 8. Genae in frontal view almost straight (®g. cate posteriorly (®g. 30). Metasoma light 37). Median frontal carina distinctly ex- brown ...... tended to middle of lower face. Meso- .... Dallatorrella ronquisti, new species scutellum distinctly narrower across scu- Ð Eyes less prominent, 1.2 times as long as ma- tellar sulcus than behind. Metasoma 10 AMERICAN MUSEUM NOVITATES NO. 3353

brown ...... Mesocynips insignis (Smith) .... Dallatorrella carinifrons (Cameron, Figures 1±5, 7 1910) Ð Genae in frontal view distinctly curved lat- Cynips insignis Smith, 1858: 117, &. Neotype erally (®g. 35). Median frontal carina ex- (designated below). tended to lower face, but rather weak, and Mesocynips insignis Cameron, 1903: 91, &. Ob- discontinuous beneath antennal sockets jective synonym of C. insignis Smith (through (®g. 35). Mesoscutellum across scutellar neotype designation: see Comments below). sulcus not distinctly narrower than behind Mesocynips insignis (Smith); Weld, 1952: 161. (®g. 31). Metasoma black ...... Ronquist, 1995b: 38±40...... Dallatorrella sinica, new species DESCRIPTION: Length: ( 9.0 mm, & 9.0± 14.0 mm. Body entirely yellow to yellowish Mesocynips Cameron brown. Basal half of wings clear and distal half ferruginous. The female specimen from Mesocynips Cameron, 1903: 91. Type species: the Philippines is darker on distal half of Mesocynips insignis Cameron, 1903 (ϭ Cynips insignis Smith, 1858) by monotypy. wings. Face, gena, and vertex glabrous and DIAGNOSIS: Body entirely yellowish sparsely punctate with hairs. Eyes only about brown. Foveate sculpture absent or almost two-thirds as long as height of malar space, absent from head and pronotum. Flagellum slightly protruding laterally beyond temple. long and slender, F1 about 4 times as long Median frontal carina present only as small as wide at midpoint; ¯agellomeres cylindri- triangular, lobular process between antennal cal, swollen distally. Eyes shorter than height sockets and completely lacking in lower face. of malar space and laterally protruding, but Vertex glabrous, posteriorly with a rim of not distinctly beyond temple. Lower face and sparse long hairs. Occiput glabrous, laterally lateral surface of pronotum almost entirely with sparse long hairs. Lateral pronotal ca- glabrous. Clypeus strongly projected into up- rinae dorsally meeting pronotal crest, thus turned trapzoid, anteriorly only slightly emarginated (®g. 1). Occipital carina present medially disconnecting lateral pronotal areas. only ventrally; dorsally carina reaches only Lateral surface of pronotum glabrous except lower margin of compound eye. Parascutal elevated dorsal part transversely costate. Me- carinae posterolaterally smoothly curved. soscutum sparsely punctate with apressed Scutellar sulcus divided into two foveae by long hair and strongly costulate, except in a median carina. Scutellum transversely cos- narrow glabrous anterolateral strip and a gla- tulate, falling gradually laterally as well as brous area laterad to parapsidal signum. Scu- posteriorly. Metapleural sulcus abruptly bent tellum transversely costulate laterally with at middle, metepisternum subrectangular. glabrate area. Mesopleuron glabrous; meso- pleural impression only vaguely visible an- MϩCu1 of forewing straight, not curved to- ward RϩSc (®g. 7). Procoxa laterally without teriorly and shallowly present posteriorly. a vertical carina. T7±T8 fused. Mesopleural triangle glabrous with sparse, COMMENTS: Ronquist (1995a) listed a short hairs. Process produced from meta- slightly and evenly curved metapleural sul- pleural carina laterad to metacoxal foramen cus and a subtriangular metepisternum as a short, slightly extended beyond ventroposter- synapomorphy for the family of Liopteridae. ior margin of metepisternum. Metepisternum I have examined the four specimens of the glabrous, sparsely punctate with hairs. Lat- only species of Mesocynips available to me eral propodeal carina raised dorsally into a and found, however, that the genus has a me- laminate structure ending abruptly posteri- dially abruptly bent metapleural sulcus and a orly. Apical metatibial process apically trun- subrectangular metepisternum, a feature cate, by far not reaching apex of metatibia. found only in the Ibaliidae among the Cy- DISTRIBUTION: Borneo and the Philippines nipoidea (Ronquist, 1995b). Since the mono- (Mindanao). phyly of the Liopteridae is well supported MATERIAL EXAMINED:3&1(. NHM: 1& (Ronquist, 1995a), this feature has apparently (Type, #7.45.), Borneo; USNM: 1&1(, Bor- developed independently in the two lineages. neo; ZLMU: 1&, Philippines: Mindanao. 2001 LIU: DALLATORRELLINAE 11

NEOTYPE DESIGNATION: First described by surfaces of pronotum, and scutellum. Anten- Smith (1858), the species was redescribed as na short in both sexes, shorter than the length Mesocynips insignis by Cameron (1903) of head and mesosoma combined; F1 at most based on specimens from the type locality. 1.7 times as long as wide at midpoint; ¯a- The latter author realized the possible syn- gellomeres distinctly swollen submedially. onymy between Smith's species and his Eyes distinctly protruding laterally beyond ``new species'', and thus gave his species the temple. Parascutal carinae posterolaterally same speci®c epithet as Smith's. The syn- produced into a triangular process. Scuto- onymy was formally established by Weld scutellar sulcus divided by two or more sub- (1950). median carinae into three or more foveae; The type of Cynips insignis Smith, 1858 median carina absent. Mesoscutellum at least could not be located and has apparently been raised laterally and sometimes also posteri- lost. It is not in the Oxford University Mu- orly; dorsal surface entirely foveate-reticu- seum (Ronquist, 1995b), nor in the Natural late or foveate-rugose and without transverse History Museum in London (Ronquist, costae. Metapleural sulcus straight to slightly 1995b, S. Lewis, personal commun.). There- bent at middle, metepisternum subtriangular. fore, the designation of a neotype becomes MϩCu1 of forewing run almost parallel ba- necessary according to the ICZN: Article sally to RϩSc before turning away abruptly 75.3.4. (ICZN, 1999). from the latter apically. Procoxa anterolater- The type specimen of Mesocynips insignis ally with a vertical carina (®gs. 18, 19, 28, Cameron, 1903 is hereby designated as the 29). Median, ventral hump on petiolar an- neotype of Cynips insignis Smith, 1858. It nulus present in females but absent in males. matches well with Smith's (1858) descrip- T7±T8 not fused with each other. tion, is from the same localities as Cynipis insignis, and is in good condition. The spec- Dallatorrella albata (Weld) imen is deposited in the Natural History Mu- Figures 6, 8±10, 12, 14, 16, 18 seum in London (Type, #7.45) and has labels reading ``Type (in red circle)'', ``B. M. Type Mesocynips albata Weld, 1944: 55, &. Dallatorrella albata (Weld); Ronquist, 1995b: 43. HYM 7.45'', ''Mesocynips insignis Cam. Type, Borneo [in Cameron's handwriting]'', DESCRIPTION: Length: & 9.0 mm. Body al- ``Cameron Coll. 1903±121'', and the neotype most entirely black, except antennae, legs, designation label of the current author (blue). and metasoma ventrally dark brown. Fore- COMMENT ON ADDITIONAL MATERIAL: The wing clear with a distal, ferruginous macula, male specimen from USNM is labeled as and hind wing entirely transparent. ``type'' (# 9939) and indicated as female. It Face between eyes and vertex densely bears Cameron's handwritten identi®cation punctate with long hairs; antennal scrobes label. Weld (1950) stated that this specimen glabrous; lower face glabrate, with sparse is a male ``cotype'' ``sent by the Sarawak hairs. Median frontal carina almost percur- Museum in July 1906''. However, in Cam- rent, indistinct only shortly at middle of low- eron's (1903) original description of M. in- er face, and strongly raised into a laminate, signis he did not describe the male, which apically round triangular process between was subsequently described two years later ocellar plate and antennal sockets. Eyes (Cameron, 1905). Therefore, the specimen in prominent, strongly protruding beyond tem- USNM cannot be a type. ple and 1.2 times as long as height of malar space. Gena and malar space glabrate and Genus Dallatorrella Kieffer sparsely punctate with hair; malar space with a vertical, almost linear shallow impression Dallatorrella Kieffer, 1911:244. Type species beneath eye. Clypeus laterally not curved Dallatorrella rubriventris Kieffer, 1911, by monotypy. and anteroventrally with a broad triangular emargination, with radiating striation contin- DIAGNOSIS: Body usually entirely black to ued to lower face. Lateral pronotal carina dark brown or at least so on mesosoma. Fo- slightly not reaching pronotal crest dorso- veate sculpture present on lower face, lateral medially. Lateral pronotal area mostly fove- 12 AMERICAN MUSEUM NOVITATES NO. 3353 ate-reticulate except ventral one-fourth or so Dallatorrella carinifrons (Cameron); Weld, 1952: glabrous; lateroventral margin roundly emar- 162. Ronquist, 1995b: 42. ginated. Mesoscutum strongly costulate DESCRIPTION: Length: & 9.0 mm. Head and transversely and sparsely foveate with long mesosoma including legs entirely black, me- hairs; median mesoscutal impression only tasoma brown. Wings ferruginous and some- present as a short impression posteriorly; no- what lighter toward base. tauli percurrent, diverging anteriorly; para- scutal carina conspicuously raised and pro- Face, most of gena, and vertex moderately duced posteriorly into a triangle. Lateral mar- punctate with long hairs; antennal scrobes gin of axilla raised. Lateral bar laterally glabrous; lower face less hairy, with distinct, curved inward, shortest distance between lat- extensive radiating striations that reach lat- eral margins of lateral bar distinctly narrower erally lower margin of eye. Median frontal than maximal width of scutellum behind. carina almost percurrent, discontinued only Scutellar sulcus divided by two submedian at middle of lower face, and strongly raised carinae into three large foveae. Dorsal sur- into a laminate, dorsally somewhat ¯attened face of foveate-reticulate with sparse long triangular process between ocellar plate and hairs. In dorsal view, scutellum broadly antennal sockets. Eyes prominent, strongly rounded posteriorly. Axillula with conspicu- protruding beyond temple and 1.2 times as ous white pubescence. Mesopleuron gla- long as height of malar space. Malar space brous; mesopleural triangle with conspicuous glabrate and sparsely punctate with hairs; white pubescence. Metepisternum glabrous, malar space with a distinct vertical, almost process ventrally projected from metapleural linear impression beneath eye. Clypeus lat- carina small. Lateral propodeal carina strong- erally distinctly incised, anteroventrally with ly raised anterodorsally into laminate pro- a broad triangular emargination, and with ra- cess. Metasomal T3 distinctly longer than T4 diating striation continued to lower face. Lat- along a dorsal median line; all postpetiolar eral pronotal carina laterally strong and crest- metasomal terga with more or less dense like, dorsomedially not quite reaching pron- hairs; T5±T6 dorsally with a band of dense otal crest. Lateral pronotal area mostly fo- hairs along posterior margin. Apical metati- veate-reticulate except a small area ventrally bial process apically tapering, reaching glabrate; lateroventral margin straight. Me- slightly beyond apex of metatibia. Pubes- soscutum strongly costulate transversely and cence sparse on femurs and conspicuous on sparsely foveate with long hairs; median me- tibiae and tarsomeres. soscutal impression only present as a short DIAGNOSIS: Dallatorrella albata is similar impression posteriorly; notauli percurrent, di- to D. maculata, can be distinguished from verging anteriorly; parascutal carina conspic- the latter by (1) female metasoma with more uously raised and produced posteriorly into or less dense hairs on all terga, (2) dorsally a triangle. Lateral margin of axilla raised. with a band of dense hairs along posterior Lateral bar broad, laterally almost straight, margin of T5±T6, and (3) T3 distinctly lon- shortest distance between lateral margins of ger than T4 along a lateral median line. In lateral bar as wide as maximal width of scu- fact, the species can be easily distinguished tellum behind. Scutellar sulcus subdivided by from all other species by its band of dense, two submedian carinae into three large fo- long hairs along posterior margin of meta- veae. Dorsal surface of scutellum foveate-re- somal T5±T6. ticulate with sparse long hairs. In dorsal DISTRIBUTION: Philippines: Mindanao. view, scutellum broadly rounded posteriorly. MATERIAL EXAMINED:2&. USNM: 1& Axillula with conspicuous white pubescence. (Type, #56810), Mindanao: Surigao; 1& Mesopleuron glabrous; mesopleural triangle (NHM), locality as type. with conspicuous white pubescence. Met- episternum punctate with hairs in upper part Dallatorrella carinifrons (Cameron) and strongly depressed with conspicuous pu- Figures 37, 39, 41 bescence in lower part, process ventrally pro- Paramblynotus carinifrons Cameron, 1910a: jected from metapleural carina well devel- 132, &. oped. Lateral propodeal carina evenly raised 2001 LIU: DALLATORRELLINAE 13 and dorsally with dense pubescence. Meta- neath eye. Clypeus laterally distinctly in- somal T3 distinctly longer than T4 along cised, anteroventrally with a broad triangular dorsal median line; T3-T6 with only sparse emargination, and with radiating striation. long hairs dorsally, T7±T8 with more or less Lateral pronotal carina distinct, but not crest- evenly distributed long hairs, and tend to be like, dorsomedially not reaching pronotal congregated into a medial band lateroven- crest. Lateral pronotal area mostly foveate- trally on T7. Apical metatibial process taper- reticulate except a small area ventrally gla- ing apically, reaching slightly beyond apex brate; lateroventral margin straight. Meso- of metatibia. Pubescence sparse on femurs scutum strongly costulate transversely within and conspicuous on tibiae and tarsomeres. area delimited by notauli; costulae in area DIAGNOSIS: D. carinifrons is very similar laterad to notauli diminutive; whole surface to D. sinica, and can be distinguished from of mesoscutum sparsely, weakly foveate with the latter by the following characters (1) gena long hairs; median mesoscutal impression in front view almost straight (®g. 37), (2) only present as a short impression posteri- median frontal carina continuous to middle orly; notauli percurrent, diverging anteriorly; of lower face, and (3) lateral margin of axilla parascutal carina conspicuously raised and laterally distinctly narrower than scutellum produced posteriorly into a nearly right an- behind. In addition, metasoma of D. carini- gle. Lateral margin of axilla raised. Lateral frons is light brown, differing from D. sinica, bar broad with foveae, laterally slightly which has black metasoma. The last ¯agel- curved inward, shortest distance between lat- lomere of female antennae has a more or less eral margins of lateral bars slightly smaller distinct segmentation. than maximal width of scutellum behind. DISTRIBUTION: Malaysia: Borneo; Singa- Scutellar sulcus subdivided by two subme- pore. dian carinae into three large foveae. Dorsal MATERIAL EXAMINED:4&:1& (holotype, surface of scutellum foveate-reticulate with NHM, #7.10.), Malaysia: Kuching; 1& sparse, long hair. In dorsal view, scutellum (AEI), Malaysia: Negri S., Pasoh Forest Re- broadly rounded posteriorly. Axillula with serve; 1& (NHM), Sarawak: Mt. Matang; 1& conspicuous white pubescence. Mesopleuron (NHM), Singapore. glabrous; mesopleural triangle with conspic- uous white pubescence. Metepisternum gla- Dallatorrella maculata Liu, new species brous; metapleural carina projected ventrally Figures 11, 13, 15, 17, 19 into a robust process and posterior to it pre- sent an additional, less prominent process. DESCRIPTION: Length: & 6.5±10.0 mm. Lateral propodeal carina slightly raised an- Body almost entirely black, except metasoma terodorsally. Metasomal T3 longer than T4 ventrally somewhat brownish; fore- and mid- along dorsal median line; T6 as large as T4 dle legs dark brown and hindlegs black. and T5 combined, T7 posteriorly oblique and Wings transparent, forewing with a distinctly broadly curved, about 1.25 times as long as contrasting, ferruginous macula distally. T6 and 2 times as long as T8. T3±T5 gla- Vertex, face, and gena glabrous; vertex brous; T6 largely glabrous with a conspicu- and gena sparsely punctate with long hairs ous band of pubescence dorsolaterally along along margins and lower face evenly punc- posterior margin; T7±T8 more or less evenly tate with hairs; antennal scrobes glabrous. punctate with long hair; T7 also with a con- Median frontal carina present only as a lin- spicuous patch of pubescence lateroventrally. ear, slight elevation in lower face and strong- Apical metatibial process tapering apically, ly raised into a prominent laminate, dorsally distinctly reaching beyond apex of metatibia. somewhat ¯attened triangular process be- Pubescence sparse on femurs and conspicu- tween ocellar plate and antennal sockets. ous on tibiae and tarsomeres. Eyes prominent, strongly protruding beyond DIAGNOSIS: This new species is similar to temple and 1.5 times as long as height of D. albata, differing from other species of the malar space. Malar space glabrate and genus in that both species have distally mac- sparsely punctate with hair, and with a dis- ulated forewing and T6 of female metasoma tinct vertical, almost linear impression be- dorsally with a band of dense pubescence 14 AMERICAN MUSEUM NOVITATES NO. 3353 along posterior margin. The new species can area. Lateral pronotal carina distinct, dorsally be further separated from D. albata by lack widened ventrally and raised dorsally into a of pubescence on T3±T5 of female metaso- submedial lobular structure, and dorsomedi- ma, and T3 being only slightly longer than ally nearly reaching pronotal crest. Lateral T4 along a lateral median line. pronotal area foveate-reticulate anterodorsal- Male unknown. ly and glabrous lateroposteriorly; lateroven- DISTRIBUTION: Borneo: Sulawesi. tral margin straight. Mesoscutum strongly MATERIAL EXAMINED: Holotype &, Sula- transversely costulate and foveate; foveae wesi: Tengah, Mt. Morowali, Ranu River less distinct posteriorly; median mesoscutal Area, 27 January±20 April 1980, M. Bren- impression only present as a short impression dell coll. (NHM, London). Paratypes 6&: posteriorly; notauli percurrent, moderately 5&, data as holotype (NHM, London); 1&, diverging anteriorly; parascutal carina con- Indonesia: Moluccas Bacan Island, Kam- spicuously raised and posteriorly rounded, pung, Wayamiga, VII-27±31 July 1981, A. not produced into an angle. Lateral margin C. Messer coll. (USNM, Washington, DC). of axilla not raised. Lateral bar broad and ADDITIONAL MATERIAL:1&, no collection glabrous, laterally converging posteriorly, data available. shortest distance between lateral margins of ETYMOLOGY: The name maculata is from lateral bars distinctly larger than maximal Latin, meaning spotted. width of scutellum. Scutellar sulcus subdi- vided by two longitudinal submedian carinae Dallatorrella maxima Liu, new species into three foveae. Dorsal surface of scutellum Figures 20±22, 24, 26 foveate-reticulate with sparse long hair; fo- veae more or less set in rows. In dorsal view, DESCRIPTION: Length: & 14.0 mm. Body scutellum rounded posteriorly. Axillula with- and all legs entirely black, eyes brown. out conspicuous pubescence. Mesopleuron Wings transparent, forewing with a broad glabrous; mesopleural triangle reduced into a distinct ferruginous band across the middle narrow strip, without conspicuous pubes- and along outer margin. cence. Metepisternum glabrous; metapleural Antenna slender in comparison with most carina projected ventrally into a robust pro- species of the genus; median ¯agellomeres cess and posterior to it present an additional, not distinctly constricted basally and apical- less prominent process. Lateral propodeal ca- ly. Vertex glabrous with shallow foveae be- rina dorsomedially strongly raised into a con- hind ocellar plate and a few less distinct fo- spicuous hump and densely pubescent. Me- veae along eye. Gena glabrous with evenly tasomal T3 distinctly longer than T4 along distributed shallow foveae, with or without dorsal median line; T6 shorter than T4 and hair. Upper face areolate-rugose except an- T5 combined, T7 posteriorly oblique and al- tennal scrobe glabrous. Lower face evenly most straight, twice as long as either T6 or foveate with dense pubescence in upper part T8. T3±T4 glabrous. T5 sparsely, ®nely and with radiating striation running from punctate. T6 densely, ®nely punctate with clypeus in lower one-third. Median frontal pubescence dorsolaterally in anterior two- carina percurrent and strongly raised into a thirds. T7 densely and ®nely punctate with prominent laminate, dorsally somewhat ¯at- pubescence anteriorly. T8 sparsely and tened triangular process between ocellar coarsely punctate with pubescence, latero- plate and antennal sockets. Eyes prominent, ventrally with a conspicuous patch of white strongly protruding beyond temple and pubescence. Apical metatibial process ab- slightly longer than height of malar space. sent, only indicated by a long, slightly ele- Malar space glabrate and sparsely punctate vated transverse carina. Pubescence sparse with hair, with a distinct vertical, almost lin- on femurs and conspicuous on tibiae and tar- ear impression beneath eye. Clypeus laterally someres. distinctly incised, anteroventrally with a DIAGNOSIS: This new species and D. pulla broad triangular emargination and with ra- are similar to each other and differ from all diating striation. Pronotum strongly expand- other species of the genus in having (1) me- ed posterolaterally to form a wide, ¯at dorsal dian frontal carina distinctly extended to 2001 LIU: DALLATORRELLINAE 15 clypeus in lower face (®g. 22), (2) pronotum cised, anteroventrally with a broad triangular strongly expanded posterolaterally (®g. 20), emargination and with radiating striation. (3) mesosubpleuron ventrally projected into Pronotum strongly expanded posterolaterally a triangular process, and (4) metepisternum to form a wide, ¯at dorsal area. Lateral pro- with an extra process laterad to metacoxal notal carina distinct, dorsomedially not foramen anterior to the prominent process reaching pronotal crest; dorsally raised into extended from metapleural carina. D. maxi- a submedial lobular structure and thickened ma differs from D. pulla by having (1) lower ventrally. Lateral pronotal area foveate-retic- face densely punctate with pubescence in up- ulate anterodorsally and glabrous lateropos- per half, (2) Mesoscutellum distinctly fove- teriorly; lateroventral margin straight. Me- ate-reticulate; without obvious transverse ru- soscutum strongly transversely costulate and gosity (®g. 24), (3) apical process of meta- mostly with foveae set in rows; median me- tibia absent (®g. 26), and (4) Forewing with soscutal impression only present as a short a wide band across middle and along outer impression posteriorly; notauli percurrent, margin, respectively, and hind wing with moderately diverging anteriorly; parascutal basal half transparent and distal half ferru- carina conspicuously raised and posteriorly ginous except tip pale. rounded, not produced into an angle. Lateral Male unknown. margin of axilla not raised. Lateral bar broad, DISTRIBUTION: Laos. with large fovea, laterally converging poste- MATERIAL EXAMINED: Holotype &, Laos: riorly, shortest distance between lateral mar- Ban Van Heue, 20 km east of Phou-kow- gins of lateral bars distinctly larger than max- kuei, 15±31 June 1965, native collector coll. imal width of scutellum. Scutellar sulcus (BPBM). subdivided by two submedian carinae into ETYMOLOGY: The name maxima is from three foveae. Dorsal surface of scutellum Latin, meaning largest. scabrous to foveate-reticulate with sparse long hairs; foveae more or less set in rows Dallatorrella pulla Liu, new species without distinct division. In dorsal view, scu- Figures 23, 25, 27 tellum rounded posteriorly. Axillula without conspicuous pubescence. Mesopleuron gla- DESCRIPTION: Length: ( 13.0 mm. Body brous; mesopleural triangle reduced into a and all legs almost entirely black, except me- narrow strip, without conspicuous pubes- tasoma slightly brownish ventrally. Wings cence. Metepisternum glabrous; metapleural evenly ferruginous. carina projected ventrally into a robust pro- Vertex mostly foveate, with small glabrous cess and posterior to it present an additional, areas submedially. Gena glabrous with even- less prominent process. Lateral propodeal ca- ly distributed shallow foveae, with or without rina evenly raised dorsally into a ridge and hair. Upper face areolate-rugose except an- pubescent along base. Apical metatibial pro- tennal scrobes glabrous. Lower face with cess present, but only as a blunt, short pro- dense pubescence in a triangular area be- cess. Pubescence sparse on femurs and con- neath antennal sockets and mostly with spicuous on tibiae and tarsomeres. strong radiating striation running from clyp- DIAGNOSIS: The new species is similar to eus. Median frontal carina percurrent and D. maxima (see above), but differs from the slightly weakened beneath antennal sockets; latter in (1) lower face nearly entirely radi- strongly raised into a distinct, dorsally round- ate-carinate, (2) forewing evenly ferrugine- ed process between ocellar plate and anten- ous, mesoscutellum distinctly foveate-retic- nal sockets; and raised beneath antennal ulate and not distinctly transversely rugose, sockets into a prominent ridge diminishing (3) metatibia with a short, distally rounded ventrally. Eyes prominent, only moderately apical process, and (4) dorsal surface of me- protruding beyond temple and slightly longer soscutellum scabrous (®g. 25). than height of malar space. Malar space gla- Female unknown. brate and sparsely punctate with hair, with a DISTRIBUTION: Malay. distinct, vertical, almost linear impression MATERIAL EXAMINED: Holotype (, Malay: beneath eye. Clypeus laterally distinctly in- Pahang, F.M.S. Fraser's Hill, 4000 ft, 6 16 AMERICAN MUSEUM NOVITATES NO. 3353

March 1946, no data on collector, with an surface of scutellum foveate-reticulate with extra label ``Ex F.M.S. Museum, B.M. 1955± long hair. In dorsal view, scutellum nearly 354'' (NHM, London). truncate posteriorly, only rounded posterola- ETYMOLOGY: The name pulla is from Lat- teally. Axillula with inconspicuous, whitish in, meaning dark-colored, blackish, grayish pubescence. Mesopleuron glabrous; meso- black. pleural triangle with inconspicuous whitish pubescence. Metepisternum glabrous; meta- Dallatorrella ronquisti Liu, new species pleural carina projected ventrally into a ro- Figures 28, 30, 32 bust, short process. Lateral propodeal carina raised dorsally throughout. Metasomal T3 DESCRIPTION: Length: & 6.0±9.0 mm. longer than T4 along dorsal median line. T6 Head and mesosoma black; legs brown to slightly shorter than T4 and T5 combined. T7 dark brown. Metasoma brown to dark brown. posteriorly oblique and nearly straight, Wings evenly ferruginous. slightly longer than T6 and about 1.4 times Vertex glabrous and sparsely punctate with as long as T8. T3±T6 sparsely punctate with hair. Gena glabrous and shallowly foveate to hair laterodorsally and glabrous laterovental- punctate with hair; face foveate with dense ly. T7±T8 evenly, sparsely punctate with pubescence; antennal scrobe glabrous and long hair. Apical metatibial process apically relatively densely punctate with hair close to tapering, not reaching apex of metatibia. Pu- base antennal socket. Median frontal carina bescence sparse on femurs and tibiae, con- strongly raised into a prominent laminate, spicuous on tarsomeres. dorsally slightly ¯attened and trapezoid (in DIAGNOSIS: The new species is similar to lateral view) process between ocellar plate D. carinifrons, D. rubriventris, and D. sini- and antennal sockets; indistinct in upper low- ca, but differs from them by having eyes er face and raised to a low ridge ventrally in prominent and distinctly longer than malar lower face. Lower face with conspicuous pu- space (®g. 28), lower part of lateral surface bescence, which makes sculpture not easily of pronotum glabrous (®g. 28), and meso- observable. Eyes prominent, strongly pro- scutellum truncate posteriorly (®g. 30). truding beyond temple and 1.5 times as long Male unknown. as height of malar space. Malar space gla- DISTRIBUTION: Papua New Guinea. brate and sparsely punctate with hair; vertical MATERIAL EXAMINED: Holotype &, Papua impression beneath eye absent. Clypeus lat- New Guinea, Bulolo, 21 October 1981, Man- erally distinctly incised, anteroventrally with ki L. A. coll. (NHM, London). Paratypes 4&: a broad triangular emargination and with ra- 2&, data as holotype (NHM, London); 1&, diating striation. Lateral pronotal carina dis- no collection data available, presumably tinct, crest like lateroventally, dorsomedially from Aru (Ronquist, 1995b); 1&, New Guin- not reaching pronotal crest. Lateral pronotal ea, NE Green River Post, 200 m, Wet Forest, area mostly foveate-reticulate except one 26 June 1963, S. Straatman coll. (BPBM). third lateroventrally glabrate; lateroventral COMMENTS: The paratype in BPBM differs margin straight. Mesoscutum strongly cos- from the other type specimens in that the tulate transversely and sparsely punctate with transverse sculptures on the lateral sides of long hair; median mesoscutal impression pronotum are more obvious and the abdomen only present as a short impression posteri- is darker. orly; notauli percurrent, strongly diverging ETYMOLOGY: This species is named after F. anteriorly; parascutal carina conspicuously Ronquist for his contributions to the system- raised and produced posteriorly into a trian- atic studies on the Liopteridae, which facili- gular process. Lateral margin of axilla not tated this present work and my other ongoing raised. Lateral bar moderately wide and lon- researches on the family. gitudinally costate, laterally straight; shortest distance between lateral margins of lateral Dallatorrella rubriventris Kieffer, 1911 bars as large as maximal width of scutellum. Figures 23±25, 38, 40 Scutellar sulcus subdivided by two subme- Dallatorrella rubriventris Kieffer, 1911: 244, (. dian carinae into three large foveae. Dorsal DESCRIPTION: Length: ( 5.5±8.0 mm; & 2001 LIU: DALLATORRELLINAE 17

5.0±8.0 mm. Head yellow, except vertex sparse pubescence. Metepisternum scabrous; dark brown. Mesosoma black; legs black to metapleural carina projected ventrally into a dark brown. Metasoma yellow. Wings trans- robust, short process. Lateral propodeal ca- parent, forewing with a very faint ferrugi- rina slightly raised dorsally throughout. Me- nous mark distally along anterior margin. tasomal T3 only slightly shorter than T4±T5 Vertex foveate-reticulate with hair. Gena combined. T6 slightly longer than T4 and T5 mostly glabrous and shallowly foveate with combined. T7 posteriorly oblique and nearly hair anteriorly and posteriorly; upper face ru- straight, 1.6 as long as T6 and about twice gose and sparsely pubescent; antennal scrobe as long as T8. All post petiolar terga dorso- glabrous and somewhat punctate with hair laterally punctate with hair; punctures coars- close to base antennal socket. Median frontal er and denser on T7 and T8. Apical metati- carina strongly raised into a prominent lam- bial process apically tapering, distinctly inate, dorsally slightly ¯attened and triangu- reaching beyond apex of metatibia. Pubes- lar (in lateral view) process between ocellar cence sparse on femurs and tibiae, denser plate and antennal sockets, indistinct in upper ventrally, conspicuous on tarsomeres. lower face and distinct in lower part of lower DIAGNOSIS: The species is most easily dis- face. Lower face mostly glabrate and rela- tinguished from other species of the genus tively densely punctate with hair in upper by its body color pattern and its glabrate- part and ventrally radiating striate. Eyes punctate lower face (®g. 36). prominent, strongly protruding, but not so DISTRIBUTION: Australia. much beyond temple as in the other conge- MATERIAL EXAMINED:3&5(:1( (holo- neric species, 1.2 times as long as height of type, NHM #7.12.), Australia: New South malar space. Malar space glabrate and Wales; 3(, Australia: New South Wales sparsely punctate with hair, with very faint (MCZ), 1(, Australia (NHM, London), 3&, vertical impression beneath eye. Clypeus lat- Australia: New South Wales and New erally distinctly incised, anteroventrally with Queensland (ANIC). a broad triangular emargination, with radi- ating striation. Lateral pronotal carina dis- Dallatorrella sinica Liu, new species tinct, but not crest like lateroventally, dor- Figures 29, 31, 33, 35 somedially not reaching pronotal crest. Lateral pronotal area almost entirely foveate- DESCRIPTION: Length: & 8.0 mm. Body reticulate, foveae less prominent lateroven- and legs almost entirely black, except meta- trally; lateroventral margin straight. Meso- soma dark brownish ventrally. Wings evenly scutum strongly costulate transversely and ferruginous. sparsely punctate with long hair; median me- Vertex densely foveate with hair, except soscutal impression only present as a short area continued posteriorly from antennal impression posteriorly; notauli percurrent, scrobe glabrous. Gena foveate-punctate with strongly diverging anteriorly; parascutal ca- hairs. Upper face more or less scrobiculate rina conspicuously raised and produced pos- with dense pubescence laterad to antennal teriorly into a triangular process ventrally scrobe. Antennal scrobe mostly glabrous and turned apically. Lateral margin of axilla not ventrally with dense pubescence. Median raised. Lateral bar moderately wide and fo- frontal carina extended in lower face to the veate, laterally slightly curved inward, short- level of ventral margin of eye, distinctly dis- est distance between lateral margins of lateral continuous beneath antennal socket, strongly bars distinctly smaller than maximal width of raised into a prominent laminate, dorsally scutellum. Scutellar sulcus subdivided by a slightly ¯attened and triangular (in lateral median and two or more submedian carinae view) process between ocellar plate and an- into multiple large foveae. Dorsal surface of tennal sockets. Lower face punctate-foveate scutellum foveate-reticulate with long hair, with dense pubescence in upper two-thirds strongly sloping posteriorly. In dorsal view, and ventrally radiating-striate. Eyes promi- scutellum rounded posteriorly. Axillula with nent, strongly protruding beyond temple, 1.2 inconspicuous whitish pubescence. Meso- times as long as height of malar space. Malar pleuron glabrous; mesopleural triangle with space punctate-foveate with hair, with dis- 18 AMERICAN MUSEUM NOVITATES NO. 3353 tinct vertical impression beneath eye. Clyp- Male unknown. eus laterally distinctly incised, anteroventral- DISTRIBUTION: China. ly with a broad triangular emargination, ra- MATERIAL EXAMINED: Holotype &, China: diating-striate. Lateral pronotal carina dis- Yunnan, Xishuangbanna, Meng'a, 1050± tinct, but not crest like lateroventally, almost 1080 m, 11 May 1958, S.-Y. Wang coll. reaching pronotal crest dorsomedially. Lat- (ZICA, Beijing). eral pronotal area almost entirely foveate-re- ETYMOLOGY: The name sinica is from Lat- ticulate, except a very small glabrous area in, meaning of China, referring to the north- lateroventrally; lateroventral margin straight. ernmost distribution of the genus in south- Mesoscutum strongly costulate transversely western China. and sparsely punctate with long hair; median mesoscutal impression only present as a PHYLOGENY AND HISTORICAL short impression posteriorly; notauli percur- BIOGEOGRAPHY rent, strongly diverging anteriorly; parascutal carina conspicuously raised and produced Cladistic analysis found three most parsi- posteriorly into a triangular process. Lateral monious trees with length of 63 steps (CI ϭ margin of axilla slightly raised. Lateral bar 74; RI ϭ 71). The topology of all trees is the moderately wide and foveate, laterally not same at the base but differs toward the top. curved inward, shortest distance between lat- The monophyly of Dallatorrella is well sup- eral margins of lateral bars as wide as max- ported by seven apomorphic character chang- imal width of scutellum. Scutellar sulcus es (®gs. 42, 43), and relatively high bootstrap subdivided by two submedian carinae into and jackknife values (®g. 42). three large foveae. Dorsal surface of scutel- Biogeographically, seven of the nine spe- lum foveate-reticulate with long hair. In dor- cies of Dallatorrellinae recognized in this pa- sal view, scutellum rounded posteriorly. Ax- per are distributed in southeast and eastern illula with conspicuous whitish pubescence. Asia. The two species of the subfamily that Mesopleuron glabrous; mesopleural triangle do not occur in this region are from Australia strongly depressed, with conspicuous white and Papua New Guinea, and they nest in an pubescence in anterior half and nearly nude unresolved terminal cluster with three others posteriorly. Metepisternum glabrate and from the Oriental region in the strict consen- sparsely punctate with hair; metapleural ca- sus tree. The cladogram suggests that Dal- rina projected ventrally into a robust, short latorrellinae originated in the Oriental region process. Lateral propodeal carina slightly and subsequently dispersed to the Australian raised dorsally throughout and densely pu- region, contrary to an earlier hypothesis by bescent. Metasomal T3 approximately as Ronquist (1995a). Ronquist suggested the long as T4 and T5 combined, as long as T6. Dallatorrellinae originated in the Australian T7 posteriorly oblique and nearly straight, region and subsequently dispersed to the Ori- 1.5 as long as T6 and about 2 twice as long ental region. However, Ronquist's hypothesis as T8. All post petiolar terga laterodorsally was not based on a phylogenetic analysis of punctate with hair, punctures coarser and relationships within the subfamily. denser on T7 and T8. Apical metatibial pro- Ronquist (1995b) also reconstructed the cess apically tapering, distinctly reaching be- historical biogeography of the Liopteridae yond apex of metatibia. Pubescence sparse based on his phylogenetic analysis of the on femurs and tibiae, being denser ventrally family. Since Ronquist (1995b), more work and conspicuous on tarsomeres. has been done on the phylogeny and histor- DIAGNOSIS: This new species is close to D. ical biogeography of several of the major lin- carinifrons, but with median frontal carina eages of Liopteridae. In addition to the pre- distinctly discontinuous beneath antennal sent study, a forthcoming study on the sub- sockets; genae, in front view, distinctly family Mayrellinae suggests that the Mayrel- curved medially (®g. 35); and lateral margin linae has originated in either the Nearctic or of axilla laterally as wide as scutellum be- eastern Palearctic, with a clade subsequently hind (®g. 31). In addition, metasoma of the dispersed to Africa (Liu et al., in prep.). new species is dark brown. Thus, both Mayrellinae and Dallatorellinae 2001 LIU: DALLATORRELLINAE 19

Fig. 42. Strict consensus of the three shortest trees (length 62, CI ϭ 78, RI ϭ 72) resulting from parsimony analysis using NONA (options: hold at maximum 1000 trees in memory, 100 replications of tree bisectionÐreconnection search). Branches marked with X indicate phylogenetic relationships con- strained during the search according to earlier studies (Ronquist, 1995a, 1995b). Numbers above clades are the number of unambiguous character changes on the clade; numbers below the clades are bootstrap and jackknife values (1000 replications, each with the same options as the original tree search).

Fig. 43. Strict consensus tree of the shortest trees with character changes that can be mapped un- ambiguously. Character changes that are uniquely derived on the branches of the strict consensus tree are indicated as ®lled circles and those are not uniquely derived are indicated as empty circles. 20 AMERICAN MUSEUM NOVITATES NO. 3353

Fig. 44. Reconstruction of the historical biogeography of Liopteridae. Geological events are sug- gested to have possibly led to the vicariances in the higher level phylogeny. ϽϾindicates a split between land masses. Related events in bold are based either on reconstruction of this study or on a concurrent study by Liu et al. (in prep.); and the events in lightface type were ®rst suggested by Ronquist (1995b). have a Laurasian origin. Consequently, the ably. Finally, Paul Goldstein has always been split of the Dallatorrellinae from the stem a pleasure to work with and talk to. species of the two Gondwanian subfamilies Liopterinae and Oberthuerellinae probably REFERENCES corresponds to the breakup of Pangea into Gondwana and Laurasia in the Middle to Cameron, P. Late Jurassic (180±145 mybp) (®g. 44). 1903. Descriptions of new genera and new species of Hymenoptera taken by Mr. ACKNOWLEDGMENTS Robert Shelford at Sarawak, Borneo. J. Straits. Brch. R. Asiat. Soc. 39: 89± The work was completed when I held a 181. Kalb¯eisch Postdoctoral Fellowship at the 1905. A third contribution to the knowledge AMNH. Fredrik Ronquist generously trans- of the Hymenoptera of Sarawak. J. Straits. Brch. R. Asiat. Soc. 42: 94±95. ferred to me the loans in his holding. Com- 1908. Descriptions of a new genus and two ments by James Carpenter, Michael Engel, new species of parasitic Cynipidae and Jerry Rozen signi®cantly improved the from Borneo. Entomologist 41: 299± manuscript. Molly Rightmyer kindly tested 300. my key against specimens. Eric Quinter 1910. On new species of parasitic Cynipidae kindly suggested linguistic corrections. Com- captured by Mr. John Hewitt, B.A., at ments from Bob Wharton and Lee Herman Kuching, Borneo. Entomologist 43: helped to improve the manuscript consider- 131±133. 2001 LIU: DALLATORRELLINAE 21

Goloboff, P. A. Ronquist, F. 1999. PIWE/NONAME version 1.5.1 (32 1995a. Phylogeny and early evolution of the bit). (available from Dr. J. Carpenter, Cynipoidea. Syst. Entomol. 20: 309± American Museum of Natural History.) 335. Harris, R. A. 1995b. Phylogeny and classi®cation of the 1979. A glossary of surface sculpturing. Oc- Liopteridae, an archaic group of cyni- cas. Pap. Lab. Serv./Entomology 28, 1± poid (Hymenoptera). Entomol. 31. Scand. Suppl. 46, 1±74. Phylogeny, classi®cation, and evolution Hedicke, H. 1999. of the Cynipoidea. Zool. Scr. 28: 139± 1922. BeitraÈge zur Kenntnis der Cynipiden 164. (Hym.). X. Eine neue Unterfamilie der Ronquist, F., and G. Nordlander Cynipiden. Dsch Entomol. Z. 1922: 1989. Skeletal morphology of an archaic cy- 190±192. nipoid, Ibalia ru®pes (Hymenoptera: Hedicke, H., and G. J. Kerrich Ibaliidae). Entomol. Scand. Suppl. 33, 1940. A revision of the family Liopteridae 1±60. (Hymenoptera, Cynipoidea). Trans. R. Smith, F. Entomol. Soc. London 90: 177±225. 1858. Catalogue of the hymenopterous in- International Commission on Zoological Nomen- sects collected at Sarawak, Borneo; clature Mount Ophir, Malacca; and at Singa- 1999. International Code of Zoological No- pore, by A. R. Wallace. J. Linn. Soc. menclature. London: The International London Zool. 2: 117. Trust for Zoological Nomenclature (c/o Weld L. H. The Natural History Museum). 1922. Notes on the Liopterinae with descrip- Kieffer, J. J. tions of new species from the oriental 1911 (1909). Nouveaux Cynipides exotiques region (Hymenoptera: Cynipidae). Phi- lipp. J. Sci. 21: 323±335. du British Museum de Londres. Boll. 1944. Descriptions of new Cynipidae includ- Soc. Enomolt. Ital. 41: 244±254. ing two new genera (Hymenoptera). Nixon, K. C. Proc. Entomol. Soc. Washington. 46: 1998. WinClada ver. 0.9.99tuc.12(beta), 55±66. shareware (download from the Willi 1952. Cynipoidea (Hym.) 1905±1950. Ann Hennig Society website http://www. Arbor, Michigan (privately printed). cladistics.com). 351 pp.

APPENDIX 1. LIST OF CHARACTERS

MAIN STRUCTURE riorly, not delimited by lateroposterior ex- tension of pronotal crest; 1, closed poste- riorly, delimited by lateroposterior exten- 1. Length of ¯agellum: 0, long and slender, sion of pronotal crest. F1 is about 4 times as long as wide at mid- 8. Posterior part of dorsal pronotal area:0, dle; 1, short and stout, F1 is at most 1.7 not expanded laterally; 1, prominently ex- times as long as wide at middle. panded laterally (®g. 20). 2. Shape of ¯agellomeres: 0, cylindrical; 1, 9. Foveate sculpture on lateral surface of constricted toward ends (®g. 9). pronotum: 0, absent; 1, present. 3. Shape of ocellar triangle: 0, not distinctly raised (®g. 2); 1, strongly raised (e.g., ®g. 10. Secondary sculpture on lateral surface 16). of pronotum: 0, more or less transversely 4. Prominence of eye: 0, prominent, distinct- ccostulate; 1, without transverse costula- ly wider than temple laterally (®gs. 35±37); tion. 1, not prominent, narrower than or as wide 11. Mesopleural triangle: 0, large (e.g., ®gs. as temple laterally (®g. 2). 3, 28, 29), 1, small (®g. 21). 5. Foveate sculpture on lower face:0,ab- 12. Shape of parascutal carinae: 0, parascutal sent; 1, present. carinae broadly not produced lateroposter- 6. Vertical impression beneath eye: 0, in- iorly (®g. 4); 1, parascutal carinae expand- distinct; 1, distinct (®gs. 22, 23). ed lateroposteriorly into a rounded triangle 7. Dorsal area of pronotum: 0, open poste- (®gs. 24, 25); 2, parascutal carinae expand- 22 AMERICAN MUSEUM NOVITATES NO. 3353

ed lateroposteriorly into an acute triangle. 25. Secondary sculpture on lower face:0,co- Ordered. riaceous; 1, foveate; 2, foveate and densely 13. Shape of mesoscutellum: 0, sloped later- punctate with pubescence. ally and posteriorly (®g. 4); 1, at least lat- 26. Glabrous area on lateral surface of erally distinctly raised (e.g.; ®g. 30). pronotum: 0, small, at most limited to 14. Scutellar sulcus: 0, divided by a longitu- ventral one-third; 1, large, occupying pos- dinal median carina (®g. 4); 1, divided by teroventral half (®g. 19); 2, extensive, cov- at least two equally strong longitudinal ca- ering most of lateral surface of pronotum rinae (e.g., ®g. 30). except upper one-®fth (®g. 4). Ordered. Surface sculpture of mesoscutellum:0, 27. Direction of vein Rs؉M and shape of .15 transversely carinate (®g. 4); 1, foveate-re- submarginal of forewing:0,RsϩM aris- ticulate or foveate and rugose (e.g., ®gs. ing proximally from middle or anterior one 25, 30). third of basalis, submarginal trapezoid (®g. 16. Shape of mesoscutellum posteriorly:0, 8); 1, RsϩM arising proximally from an- broadly rounded (®g. 31); 1, almost trun- terior end of basalis, submarginal subtrian- cate (®g. 30). gular (®g. 7).

Shape of lateral margin of axilla:0,not 28. Shape of M؉Cu1 of forewing: 0, straight .17 raised (®g. 4); 1, raised (e.g., ®g. 13). or slightly curved with basal half divergent 18. Shape of mesosubpleuron: 0, ventrally from RϩSc vein distally (®g. 7); 1, abrupt- straight or very slightly curved medially; ly curved in the middle with basal half par- 1, ventrally projected medially into a tri- allel with RϩSc vein distally (®g. 8). angular process. 19. Presence of additional process laterad to COLOR AND PUBESCENSE metacoxal foramen anterior to the prominent process extended from meta- pleural carina: 0, absent; 1, present (®g. 29. Color of head: 0, yellow; 1, black. 21). 30. Color of metasoma: 0, yellow to brown; 20. Presence of vertical carina on lateral 1, black. side of procoxa: 0, absent; 1, present. 31. Color pattern of forewings: 0, nearly 21. Shape and size of apical process of me- clear; 1, evenly ferruginous; 2, distally tatibia: 0, absent (®g. 26); 1, short and banded broadly; 3, distally maculate. truncate apically (®gs. 5, 27); 2, long and 32. Hair patch in mesopleural triangle:0, pointed apically (®g. 33). Ordered. sparse or absent; 1, conspicuous. 22. Abdominal T7±T8: 0, separate; 1, fused 33. Conspicuous pubescence in axillula:0, with each other. absent; 1, present. 34. Band of dense hairs in coarse punctures dorsally along posterior margin of ab- SURFACE SCULPTURE AND WING VENATION dominal T5 (female): 0, absent; 1, present. 35. Band of dense hairs in coarse punctures 23. Extension of median frontal carina in dorsally along posterior margin of ab- lower face: 0, absent; 1, to the middle or dominal T6 (female): 0, absent; 1, present at least broken medially; 2, complete to (®gs. 14, 15). clypeus. 36. Band of dense hairs in coarse punctures 24. Radiating striate sculpture on lower ventrolaterally along posterior margin of face: 0, present only on lower half; 1, pre- abdominal T7 (female): 0, absent; 1, pre- sent nearly on entire lower face. sent (®g. 14).