Essay on the phylogeny of (: )

C. VAN ACHTERBERG

Achterberg, C. van.: Essay on the phylogeny of Braconidae (Hymenoptera: Ichneumo- noidea). [Studie

A cladistic analysis is given of the higher classification of the Braconidae (Hymenoptera). The analysis is based on characters of both adult and larval morphology, as well as on their general biology. The Aphidiinae are considered to be a subfamily of the Braconidae not ilosely related to any of the other main groups of the subfamilies. Three groups are given subfamily rank in this paper: Dirrhopinae subfam. nov. (based on the Holarctic genus Dirrlu4te Foersler, 1862), Histeromerinae Fahringer (based on the Holarctic genus Flisrer- omzrrrs Wesmael, 1838), and Trachypetinae Schulz (based on the Australian genus lra- chtpetrrs Gudrilrt.lS3l). The Cenocoeliinae are separated from the Helconinae, a subfamily to which they are not closely related. The Blacinae (including 1)y.rcrlztes Haliday, 1849) are treat€d as a group most closely related to the Euphorinae, and the Microtypini are included in the Homolobinae, New tribes are: Leptorhaconotini trib. nov. (position uncertain. provisionally in the Doryctinael based on the genus Leptorha

C. \'on Achterberg, Rijksmtseun vtn Nattrurlijke Historie, Postbts 9517, NL'2300 RA Lciden, t h c N etherlands.

The phylogeny of the Braconidae has been the was the research of Edson and Vinson ( 1979) on subject of only a limited number of papers. The the venom apparatus of female braconids. basis was laid by Griffiths (1964, only The main reason for the confusion in the sys- Alysiinae-Dacnusini), iapek (1965 & 1970), and tematics of the Braconidae is the occurrence of Tobias ( 1967). ln all these papers general biology many parallel evolutionary trends (leading to was of importance for the construction of a final " homeoplasies"); for a list see Van Achterberg, classification. Van Achterberg (1976) gave a hi- 1976. A good example is the formation of a storical review and tried to construct a division metasomal carapace (Tobias & Dudarenko, and key to the subfamilies of the Braconidae. 1974). Comparative morphology of the posterior mainly based on morphological characters of the side of the head (Tobias & Potapova, 1982) and final instar larvae and (some additional) features grouping on base of overall similarity, result in of the adults, especially of the lst metasomal one huge mixed group ( = type H II): inclrrding tergite 1V an Achterberg. 1974). part of the Agathidinae, Blacinae, Euphorinae, The (internal) relationship of some subfamilies Neoneurinae, Macrocentrinae. Microgastrinae, hav e been analy zed cladi sti cally : Amicrocentri- Ichneutinae, part of Opiinae, Gnaptodontinae, nae (Van Achterberg, 1979a), Xiphozelinae (id.. Sigalphinae, Orgilinae. Homolobinae, Acaelii- l979b). Homolobinae (id.. 1979c), Helconinae nae, Cheloninae, Helconinae, and Cenocoelii- (id., 1983a), Gnaptodontinae (id., 1983b), and nae. Additionally a large. more homogenous Microgastrinae (Mason, I98l), but the majority group (B II with Rogadinae, some Doryctinae, ofthe groups remain untreated. A major advance Braconinae, and large part of Opiinae), a smaller 42 C. van Achterberg

' -'- .a - -)' i<-+-1''\S:r-==--r ,( ) t I S ,y': il / 6 ': '; ----'r"-' I/,Q \&/ ^ } )-*< d/

5 --- --*- Figs. l-l l. Bracon subrrlosrts Sz6pligeti, holotype 9, Braconidae-Braconinae. - l. Habitus. latera.l aspect. - 2- Wings. - 3- Scapus and pedicellus, outeraspect. - 4. Outer middle claw. 5. Apex ofovipositor. - 6. Hind leg. - 7. Head, dorsal aspect. - E. Head, frontal aspect. - 9. Detail of vein l-SR of fore wing. - 10. Thorax. doisal aspect. - I l. lst 3rd metasomal tergites. dorsal aspect. - l. 2,6. Scale line : I mm. -3-5. Scale-line =5mm.- 7-l t. Scale-line = 2 mm. homogeneous group (B I with part of Doryctinae but no synapomorphy has yet been detected. For and Histeromerinae) and groups with specialized such a group the name "interim-group" is pro- head morphology (A ll with Alysiinae, Ag II posed because of its provisional nature and to containing part of Agathidinae with elongated indicate that it is "sandwiched" between groups head, H lll with Cltrvsopolthtlnnr s and with autapomorphies. At least some of the in- Wesmaelia (Euphorinae) and H IV with Cerco- terim-groups may be poly- or paraphyletic and barconinae). Only these latter groups possess will certainly be subdivided in the future. Candi- "positive apomorphous character-states" in the dates within e.9., the Doryctinae, are the tribes head morphology, opposed to "negative Evaniodini Fischer (Neotropics) and Leptorha- apomorphous character-states" or "reductional conotini (tribus nov., based on Leptorlruconotus character-states" (Van Achterberg, 1983b). The Granger, 1949 from Malagasy), which do not fit latler concerns reductions. which occur rather well within the Doryctinae: other groups lrequent[y and independently in different groups; assigned very provisionally to a subfamily are thus easily leading to incorrect conclusions. The the Microtypini (Homolobinae) and the Aneuro- chance of pafallelism in "positive apomorphous braconini (Orgilinae). The only possibility for a character-states" (especially when complicated more natural system is to find more (positive) struclures are involved) is less and is lound only apomorphous character-states by careful cladi- rarely in distantly related groups. but the sec- stic analysis of large collections. It is obvious ondary loss of a stntcture should be reckoned lhat the cladistic analysis as summarized in Figs. with. Another problem is the lack of synapomor- 12-17 is a very provisional one to stimulate fur- phous character-states for some groups. For in- ther research; e.g., larvae of too many subfami- stance, the Doryctinae and the Rogadinae seem lies (22 out of 35) are not or hardly morphologi- to be sister-groups because of overali similarity, cally known. Essay on the phylogeny of Braconidae (Hymenoptera: Ichneumonoidea) 43 New or less known characters used 1950), but the base of the "tail" is widened as in the Euphorinae (Euphorini: Jackson, 192E; analysis for the Brindley, 1939: Centistini: Loan & Holdaway, Despite the huge size of the Braconidae (t 1962: Meteorini: Simmonds, 1947), a condition 40.000 spp. or more) this paper attempts to gi\e dissimilar to the (plesiomorphous) condition in an overall analysis of the sublhmilies. Many early instar larvae ofgroup IL In the Aphidiinae characters have been reviewed in the material some Aphidiini may have a somewhat widened available and some characters found to be stable base ofthe "tail". a condition absent in the other enough within genera or tribes to be used for the tribes (e.g., Stari, 1966). The young larval in- cladistic analysis. However, several exceptions stars of many groups still need to be investigat- are present, but these are usually not mentioned ed, especially in such controversial groups as the because they appear to be secondary develop- Acaeliinae, Miracinae, Dirrhopinae, Gnapto- ments; e.g., the few Ichneumonidae-Cryptinae dontinae, Meteorideinae, Blacinae, and Sigal- with 2nd and 3rd metasomal tergites united. phinae. The function of the caudal vesicle is un- L Venom apperutus of fentale (Edson & Vin- certain; it may be an accessory respiratory or- son, 1979). Group II and a majority of the Ap- gan, used for the excretion or for the avoiding hidiinae posses a type I venom reservoir with a encapsulation by the host; an explanation is wall containing many longitudinal and circular needed to account for the disappearance of the muscles. The minority of the Aphidiinae and the organ in later instars of the endoparasitic . past groups lll and lV have type II venom reservoir: 3. Euo- verstrs e ndopttrasi tislt. ln the thin-walled with relatively few muscles. Out- conlusion has arisen about the ectoparasitism of group comparison reveals that type I is the several subfamilies in the groups III and IV. apomorphous state (as outgroup is used the sis- because (as was discovered later) numerous en- ter-group of the Braconidae as depicted in Fig. doparasites have a final (obligatorily) ectopara- l2)l Ihis state is probably lost independently in sitic phase. External feeding of endoparasites the tribe Praini of the Aphidiinae, a group with was observed in the Macrocentrinae (Daniel. several other (not related) apomorphous charac- 1932), Homolobinae (Allen, 1977). Helconinae ter-states. The venom apparatus of the following (Brachistini: Haeselbarth, 1962)' Cheloninae subfamilies has not examined by Edson & Vin- (Vance, 1932), and Agathidinae (Simmonds. son: Histeromerinae, Mesostoinae, Ypsistoceri- 19471 Nickels et al., 1950). Endoparasiles of nae, Gnaptodontinae, Acaeliinae, Miracinae, group I (e.g., Alysiinae: Caudri. l94l) and the Dirrhopinae, Meteorideinae. Cercobarconinae, Aphidiinae, also some parasites ofthe groups III Trachypetinae, Xiphozelinae, Sigalphinae, and IV ( Microgastrinae: Vance. l93l: Cardio- Khoikhoiinae, Betylobraconinae. and Amicro- chilinae: Chamberlin & Tenhet' 1926: Eup- cenlrinae. Mostly small subfamilies of which no horinae: Brindley, 1939; Loan and Holdaway, material in alcohol was available for study. I 1962) have no ectoparasitic final phase. It is have prepared the venom apparatus of Acaelius obvious from cladistic analysis that the switch sub.fhsciatus (Haliday) (Acaeliinae); it proved to from ecto- to endoparasitism has been made at be different from the venom apparatus of the least 4 or 5 times (first in the parent population of Microgastrinae, a group with which it is gener- the groups III and IV. later in the Aphidiinae, in ally associated, and fits best the diagnosis oftype the I chneutinae-group (nos. lO l4), in the I. Alysiinae-group (nos, 8 and 9), and finally in the 2. Cutt

I5. APHIDII}IAE

subalar depress ion soooEh

larval antenna disc-shaped 1arva1 spiracles without division into atrium and closing apparatus parasi Ees of aphids 2nd oetasomal spiracle in epipleuron III

larvae r^rithout caudal vesicle IV Type I venoo apparatus of g (secondari Iy lost in Praini) lst and 2nd larval instar with if endoparasi tic then caudal vesicle or "tail" s,idened basally pupation in host 1arva, Type II venom apparatus of g adult, pupa, cocoon or pupariun (secondarily endop aras i te s uith pupation outside hosl or its cocoon (except I'leteorideinae IGINEIII'TCIIIDAE outside hos ! in soDe genera Aphidiinae), and some Microgastiinae), and final ectopa- of phase ,.,ithout. tinal ecLopaEa- rasitic occurs in several subfamilies [(2 1 metasonal spiracle in epipleuron sl trc phase I larva€ wi th accessory longi- tudinal tracheal .oIIl- BRAMNIDAE I st me tasomal ml, s s ure s terni te ends near apex of PAXYLOMMATIDAE 2nd and 3rd metasomal I st tergir rgi tes united lst sternite lst sternite ends near beIoe,2od rergite middle of lst tergite partial amalSaoation of SR and M of fole wing 2r:-m of hind wing absent lst subdiscal ce11 of fore !,ing subquadrate associated aastigma + vith ants parated ectoParas i tes abselce of vein I-SR+M of fore wing of concealed J.iving coleo- vein lr-m of hird wing absent p terous larvae

Type lI venom apparatus of g spinning of a cocoon amalgarnation of veins C aod SC+R of fore wing head of larvae with prominent sclerotized bands hypostomal spur of l-arvae present

apotDorphous I. IG]NEU}.O{OIDEA o= cha!acter-state U= plesiomorphous character-state

Fig. 12. Cladogram of lchneumonoidea: families and subfamily groups within the Braconidae (Aphidiinae and goups Il-lv). Essay on the phylogeny of Braconid.ae (Hymenoptera: Ichneunanoidea) 45 an endoparasite of a (partly) concealed living and the statement of Mason ( l9tt I :3) that the host larva still allows a final ectoparasitic phase position of the lst spiracle in the epipleuron (e.g., in the Helconinae, Cheloninae, Homo- should be considered the plesiomorphous state lobinae, and Macrocentrinae). An ectoparasitic is, in my opinion, therefore incorrect. phase of an exposed ("phanerozoic") host larva 6. Derek4tment oJ' parastigma. The para- would involve a high risk for predation. This may stigma is usually much more differentiated from be why the final ectoparasitic phase is absent vein SC+R than in other Hymenoptera and (thus secondarily lost) in the Microgastrinae, seems to be an autapomorphy of the Braconidae. Cardiochilinae and Euphorinae. If this is correct, The vein connected to it (l-SR) is also subject to then it is to be predicted that the Orgilinae have change of position in many groups (part of the an ectoparasitic phase, although it has yet to be Braconinae, Microgastrinae, Neoneurinae) or observ ed. may be (largely) absent (Agathidinae). 4. Pupation in- or outside the host ttr its t'o- 7. Development ol dorsope - The development toon. Pupation site selection is an indication of ol'a dorsope (Van Achterberg, 1974) is an impor- the polyphyletic origin of the endoparasitism. tant apomorphous character-state. It is not known Most aberrant is pupation in the mummified cat- outside the Braconidae and occurs in two erpillar by the Rogadinae-Rogadini. groups: the group with Doryctinae and Alysiinae ln the Alysiinae-$oul the parasite pupate in- (large parts of nos. l, 2, 8 and 9, but absent in side the host-puparium (usually after the con- nos. 3-7) and in the Euphorinae-group (large part struction of a delicate cocoon) and in the of nos. l7-19). Within these groups the dorsope Alysiinae the specialized mandibles are used to may be secondarily lost or shallow. In the make a hole in the puparium. The subfamilies of Blacinae a shallow dorsope is still recognizable the Ichneutinae-group (nos. l0-14) are known to by the curvature of the dorsal carinae basally; in make a delicate cocoon within the host-cocoon. the Euphorinae as well in the Doryctinae the The Aphidiinae pupate within the mummified dorsope disappears because of the petiolization aphid skin mounted to the surface of the plant; of the lst metasomal tergite. y p o p e de p r e s s i only three genera (with several apomorphies) are t\. H cl -v al o tt. Usually referred known to pupate below the aphid. viz.. Praon to by the incorrect term "cyclostoma": the real Haliday (tribe Praini), D.yscrirrlrr.s Hincks and apomorphous state concerns a more or less con- Protapltidius Ashmead (both tribe Aphidiini). cave and glabrous labrum and a clypeus which is The remaining groups III and IV, which contain partly depressed and forms part of the hypo- endoparasites, pupate outside the host or its co- clypeal depression. The result is not a "round coon. Exceptions are the genus Dealedx-vs Ma- mouth" (=cyclostoma), because the mouth is son (pupation in host cocoon. parasite of situated more ventrally and the depression is Lyonetiidae, obviously a secondary develop- normally elliptical rather than round. The hypo- ment within the Microgastrinae) and the subfam- clypeal depression may have been evolved to ily Meteorideinae (gregarious parasites of provide the mandibles more working space, Gelechiidae and Tortricidae, spin cocoons inside especially to grip objects. The hypoclypeal de- the host pupai which is almost unique for the pression is distinctly development in almost all Braconidae, independently developed only in the spp. of the subfamilies nos. l-3; the nos. 5 and 6 genus B uc c u latriple"r Viereck (Rogadinae)). have the depression formed only by the back- 5. Sitrrtttion ol the spirat:le ql tlte se

Braconinae (except in a few tropical genera), sites of larvae of (wood- and bark-boring) Col- Histeromerinae. Mesostoinae, Ypsistocerinae, eoptera. Less commonly other hosts in plant Ichneutinae, Neoneurinae. Cardiochilinae, tissues are attacked. Khoikhoiinae. and Microgastrinae (with the ex- ception ol Philoplltrs Nixon). 2. Rogadinae Foerster. 1862 lO. Tru n sr ers e post e rior sc t I e llar de pre s s io n. This is a frequently narrow, transverse depres- Large and heterogenous group: in this subfamily sion medio-posteriorly at the scutellum and is an independent transition to endoparasitism has more or less crenulate. Main character-state fbr taken place. This accompanied with some pecul- group I I I. uniting mainly spp. with comparative- arities, such as the toothless mandibles ot'the ly complete venation ol the wings, if compared larvae and the mummification of the host- with group IV. Especially in the Microgastrinae caterpillar. Gregarious ectoparasitism occurs in and the Cardiochilinae the scutellum may be several genera, e.g., Hormitrs Nees, (Jnt.o- sculptured posteriorly, but only very rarely (e.g., 1rlrnnes Foerster, and Cltrentylus Haliday. Solif Ildrtemita Cameron) is it similarly depressed as ary endoparasitism is usual in the genus Rogns in group III. This may indicate the transverse Nees and its allied genera, but gregarious endo- posterior scutellar depression is secondarily lost parasitism is known. in group IV. ll. Separated crrspidal pro(?ss. As pointed 3. Braconinae Stephens, 1829 out by Tobias (1967), an unseparated cuspidal process of the male genitalia has to be consid- Very large subfamily, consisting of solitary or ered the plesiomorphus character-state, as pre- gregarious ectoparasites of larvae of holometa- sent in the sister-group. A narrow cuspidal pro- bolous . According to iapek (1970:862r cess is separated in the Homolobinae as treated the host larva is paralyzed at egg deposition, the in this paper (including the Microtypinae) as well parasite larva feeds on the paralyzed host and as in the Euphorinae and Blacinae. A dissimilar forms its delicate cocoon in a sheltered place. It separale cuspidal process is present in the Agat- is the only group wherein genera occur with two hidinae. In the cladistic analysis presented here fore tibial spurs (Rhumnura Enderlein and some the Agathidinae are referred to a group dill-erent new genera). from the group which includes the Homolobinae and the Euphorinae; so it appears that the sep- 4. Telengaiinae aration of the cuspidal process in these two Tobias, 1962 groups is a parallelism. The presence of a sepa- Small subfamily, containing only the South rate cuspidal process in the Homolobinae as well Palaearctic gents Telengaia Tobias. The biology as in the Blacinae and Euphorinae (p.p.) seems is unknown. Because of the 5-segmented to be another case of parallelism, although these maxillary palp, the absence of the occipital and groups are comparatively closely related. prepectal carinae, and of the dorsope it seems to 12. Development oJ pronop?. The develop- be related to the Braconinae. However. the lack ment of a pronope (a depression in the pro- of the synapomorphies of the hind wing, as well notum: Van Achterberg, 191t3: 301, Fig. 55) is the highly aberrant mesopleural depression seem considered to be an important character-state of to justity its separate position. several groups av Braconidae; e.9., of the Helconinae-group (nos 16-19). Some genera of 5. Ypsistocerinae 1923 these groups may have the pronope absent, c.g., Cushman, D),scoletes Haliday of the Blacinae, but usually Small subfamily, which contains the genera the pronope is more or less developed. Yltsist

Fischer. 1970 7. Histeromerinae Fahringer. 1930, stat. nov. 10. Gnaptodontinae genus Small subthmily, containing the Holarctic genus Small subfamily, the cosmopolitan primarily parasite ol Histerom?nts Wesmael; in the Netherlands Gnuptodon Haliday is a (Lepidoptera). lound to be a parasite of Cisidae boring in Nepticulidae It is still not certain if this genus contains ecto- or endoparasites (Van Polyponrs sp., in the Nearctic region reared fiom ven- Anobiidae and Lyctidae (Coleoptera). This Achterberg, 1983b); the morphology o[ the is unknown. group was included in the Braconinae by Van ome apparatus and of the larva Achterberg (1976:45) because of several syna- pomorphies. However, it does not fit well (as it did not in the Doryctinae before) because of the ll. Dirrhopinae subfam. nov. character-states: 2nd lollowing apomorphous Contains only one gents. Dirrlnltt Foerster, and 3rd metasomal spiracles in the epipleuron, known in the Northern Hemisphere, and a para- peculiar of head and antenna, the cluster shape site of Nepticulidae. Peculiar autapomorphy is of pegs on the flore tibia, and the long hind the llattened I st metasomal tergite with the basitarsus. Further plesiomorphus character- spiracle behind the middle of the tergite. states (absent in the Braconinae) such as the presence of the occipital carina and the long vein M+CU of the hind wing stress the isolated posi- tion of this group. 12. Ichneutinae Foerster, I tl62 Rather small subfamily, consisting of endopara- sites larvae sawflies (lchneutini Foerster 8. 0piinae Foerster, 1862 of of and Proteropini Van Achterberg, both with fairly Rather large subfamily, consisting of endopara- complete fore wing venation) and of lepidopte- sites of (especially mining) larvae cyclorraphous rous leaf-miners (Nepticulidae: Mtresebeckiini Diptera. exceptionally ovo-larval parasitism Mason with reduced wing venation). occurs. The pupation (as in the following sub- family) is in the puparium ofthe host. Frequently hypoclypeal depression has been de- a shallow 13. Miracinae Viereck, l9ltl veloped, indicating the relationship with the Braconinae-group, but the labrum is flat and fre- Small subfamily, parasites of Nepticulidae and quently bears some setae. Lyonetiidae, in the Palaearctic region only the genus Mirax Haliday. Additional apomorphous character-states are the strongly oblique vein 9. Alysiinae Stephens, 1829 cu-a of hind wing and the absence of the pre- One of the largest subfamilies of the Braconidae l pectal carina. 48 C. von Achterberg

IIJ 8. OPTTNAF 9. AIYSl INAE ?nd spi rac'l e in notum or epi- p I euron mandibles'rexodont" 2nd metasomal spi rac le in epipleuron MIRAClNAE lst metasomal spira- 1arva1 mandibles s looth c1e in membranous epipleural area endoparasites of cyclorraphous I4. ACAELIINAE Diptera scu!ellutrl lrith oblique antenrtal . Dosterior fovae E-cu of educt ion of pr:epectal carlna ' seg&ents fixated fore wing very short rooope often present I2. ICHNEUTINAX t.' l4 l-u of fore tendency to wing curved ovo-1arwa1 paras it ism lst tergite 'irmovably j oined l-M of fore to 2nd tergite winS curved I ]. DIRRHOPINAE propleu"al SRI fl ange of fore wing. stlong1y curved ab s ent dorsope preseot lr-m of hind rring 1st metasomal (sub)vertical 'spiracle behind t of fore wing couparatively middle of lst I LerSite anteriorly situated

spiracle n notullt 2nd cergite lst metasorltal spiracle in flont r{i th basal of middle of tergite e levat i on

2nd metasoma 2nd metasotual spiracle in epipleuron sp i rac le in notum

tendency to an open marginal ce11 of fore ving

tendency to flattened l6t EetasoDal tergite

prlmary endoparasites of Nepticulidae, further of mining or boring Lepidoptera (Lyonetiidae, Cracillariidae and Eucosmidae) mining, gall-forning or exposed living htEenopterous lalvae prepectal carina reduced margina'l cell of fore wing shortened or reduced dorsope i1b sent Essay on the phylogmy of Braconidae (Hymenoptera: Ichneumonoidea) 49

INAE 3. BRACONTNAE 4, TSLTNGAT 5. YPSISTOCER INAE l-M of hind wing aL least 2 x 6th tergite sociated rrith ternites M+CU and wide[ed near cu_a stroogly convex ventral part of clypeus depressed and wi rir lateral and part of hypoclypeal depression crease ovipos i tor issued submeci- cu-a of hind ir'ing short and + mcsopleuron r,r i th a1ly from me t as ona reclivous wide elliptical -SR of fore r,ring long depres s ion 2. ROCADINAE . 6. MESOSTOINA.E of hind wing arSr long and s l ender widened ovipositor l. DORYCTINAE upcurved developrEent of apical 2 segments mesoscutum Pro lru- endoparas i ti sm of naxi 11ary ding anteriorly of nore exposed palp uni ted l-SR of fore wing very short or gregari- hosEs occipital clypeus not depres sed ous ectoparasites pe 8s ca! i na occipital carioa absent fore t ibia absent s.ure11er sulcus absent 2nd meta- somaL 7. H ISTEROMERINAE spiracle hind basitarsus eolaiged in notum occipital carina present hypoc lypeal depression very large scutellar sulcus reduced propleural flange lsE tergi te strongly flattened PresenE 2nd spiracle io epipleuron fenora compressed dorsope usuatly Dre s ent tendeocy lo flattened sides of lst tergif,e dorsope absent propleulal flange absent

labrum + concave and gl abrous clypeus usuatly part of hypoclypeal depression, i !s ventral margin medially more dorsal ly s i tua ted than mandibular condylus (gregarious) ectopata- sites

Figs. l3-14. - 13. (left) Cladogram of subfamily group ll, excluding lla. - 14. (above) Cladogram of subfamily group lIa. 50 C. van Achterberg

14. Acaeliinae Viereck. l9l u However, the presence of a dorsope (if weatly developed still recognizable by Small subfamily, sometimes incorrectly spelled the curvature of the basal part of the dorsal as Adeliinae which is based on a "lapsus calami" carinae) separates the Blacinae from the Helconinae. Because si- in the original description. Primarily solitary en- the milarities concern negative (re- doparasites of larvae of Nepticulidae, but also apomorphies ductions) and the development reponed from Gracilariidae and Eucosmidae. of a dorsope is a positive apomorphy, I give mosr weight to the latter character. Therefore the and the 15. Aphidiinae Haliday. 1833 Dyscoletini are excluded from the Helconinae Medium-sized subfamily, exclusively solitary and form a separate subfamily. The secondary endoparasites of aphid nymphs and adults. Fre- loss of the dorsope by the Brachistini is unlikely quently treated as a separate family, but it shares because of the shape of the basal portion of the dorsal carinae, all the synapomorphies of the Braconidae, sev- further they differ by the short dorsal propodeum. eral ones with groLrp II (venom apparatusl), and face of the A paper (partly dealing with the phylogeny the cephalic structure of the final instar larva is of rhe Blacinae) will published similar to that of other Braconidae according to be in the near tuture. Capek (1970: 848). The occurrence of two addi- The Blacini are included in the Euphorinae by Tobias (1965: tional apomorphies: the venom appa"ratus type II s.l. 502-504, translation) on in the Praini and the obviously original pupation basis of a negative apomorphy: the loss of vein CU within the host of most spp. indicate the correct- lb of the fore wing, but it is also lost in the subgenus Calyptus of the genus ness of the inclusion - of lhe Helconinae. A further reason is the presence ofa separated cuspidal process and of a supposed 16. Helconinae F-oerster, I tl62 similar biology. The biology has not been well Rather large subfamily, all endoparasites of studied, but the best provisional conclusion is larvae of Coleoptera. The group is not well char- that the Blacini are parasites of coleopterous acterized by autapomorphous character-states, larvae. The Dyscoletini are parasites of larvae of both larvae and adults possess many plesiomor- Boreidae (Mecoptera). Further study of the phous character-states. This supports the hypo- larvae is necessary; study of the morphology of thesis that the Braconidae evolved from ectopar- the lst and 2nd larval instars, and the presence of asites of sheltered living larvae of Coleoptera a final ectoparasitic phase (absent in the Eupho- (e.g., Cerambycidae). rinae, present in the Helconinae) is essential. Inclusion of the Blacini in the Euphorinae is obstructed by differences in the mandibles ofthe 17. Meteorideinae ilapek. 1970 final larval instar, the biology and the morpho- Small subfamilyl gregarious endoparasites of Iogy of the lst metasomal tergite. larvae of Lepidoptcra and pupation inside the Peculiar to the Blacini is the development of host pupa. Contains lwo genera: Bariazrzr Nixon nuptial swarming, starting with swarming con- and Meleoridea Ashmead. and is resticted to the gregation of males in the evening and followed (sub)tropics (but unknown from the Australian by mainly nocturnal mating. Southwood (1957) region). observed two types of dance in a swarm of males, similar to a chironomid swarm: a slow vertical dance and a rapid horizontal dance lead- 18. Blacinae Foerster. 1862 ing to consolidation of the swarm, probably Medium-sized subfamily, containing the Blacini owing to the enlry ol a I'emalc. sensu Van Achterberg. 1976 and the Dyscoletini (tribus nov. based on the genus Dlscoletcs Hali- 19. Euphorinae Foerster, day). Frequently the Blacini are included in the lu62 Helconinae or in the Euphorinae. The venation Rather large and diverse subthmily, consisting of of the adults and the cephalic structure of the solitary or gregarious endoparasites ol lepidopte- final instar larvae of the Blacini is similar to rous and coleopterous larvae (Meteorini those of the tribe Brachistini of the Helconinae. Cresson; some spp. with the cocoon hanging Essay on the phylogeny of Braconid.ae (Hymenoptera: Ichneumonoidea) 5l

20. GFCOBARCC'.IIME 2l ' | 7 . MEIEORIDEII,AE r nd spurs ane 11i form scaPus pupation ovoid and propodea ins ide host I 8. BIACIME I concave apically (inc1, Dyscoletini) irac le 1i r- around s temmaticum 3rd sterni larval epistoma crenulate, dePressed enlarged reduced occiput incised gregarious 3rd s ternite norrnal parasi tes marginal cell of fole wing lepidop- of ( sub ) truncat e lalvae 2-CU of hind wing ptesent laterope developmenr 22. xrprrozE-rt{AE shal10t, or of parasita_ ab s eot tion of imagines !i c-shaped laEerope near sPiracle p1ical 1ob e Larval Earldibles rather large rnandib 1es gnooth toothed absent final ectoparasitic 2-CU of hind wing usually dorsope phase lost Ist metasomal terglte (near -ry, . 21. Hcr'lor,oBrNAE cuspidal process ' "(incl.',/ [icrotyPini ) + tube-shePed separated basal ly of hind wing vertical netasome insetted above hind coxae 2-Cu of hind uing absent

dorsope pres eot antescutal depres sion preseDt cusDidal process sepafated

pronoPe pronoPe present abs ent medi a 11y with subpronope alrd a posterior Pronope ovo-larval parasiEes of lepidop!erous larvae

pronope absent or deep and posteriorly cu-a of hind a,ing strongly inclivous

2-CU and r of hind wing often present parasites of lepidopterous larvae (uflknoirn of nos 20 & 2l)

transverse scuEe Llar depress ion present posEeriorly palas ites of coleopterous larvae

III Fig. 15. Cladogram of subfamily group ltl. 52 C. van Achterberg

from a thread), of solitary or gregarious en- Tratltypettts Gu6rin. ol'which the biology is un- doparasites of adult Coleoptera (and more sel- known. Only the type-species, T. clovttl,t dom of their larvae I Cosmophorini Muesebeck & Gu6rin, ltl3l is described and the t'emale is un- Walkley, Centistini Capek, and several genera ol known. l'raclt1, petrl.r was generally included in the Euphorini: Perilitu s -grotp, Roltalopltortrs the Helconinae, however, there are many dift'e- Haliday. lrypt

Homolobinae). According to Capek (1970: 859) 28. Amicrocentrinae Van Achterberg. 1979 Acampsini are ovolarval the Sigalphini and Small subfamily. containing only the Afrotropi of lepidopterous larvae. endoparasites cal genus Anti( n)tentn!m Schultz, u'ith solitary parasites of large boring lepidopterous larvae (Cossidae and Noctuidae). Despite the large 25. Cenocoeliinae Sz6pligeti, 1901 number of remarkable autapomorphous charact- er-states as listed by Van Achterberg (1979a Rather small subfamily. containing endopara- 2-3), and the lack of almost any synapomor- sites of larvae of wood- or bark-boring Coleo- phous character-states uniting lhe Macro- ptera, usually Cerambycidae, but sometimes centrinae and Amicrocentrinae, with the availa- parasitize Scolytidae or Buprestidae. Some spp. ble set of characters both groups seem to be the larvae (Curculionidae and Cerambycidae) most closely related subfamilies of the (Saffer, within herbaceous stems or fruits 1982). Braconidae, a conclusion in agreement with the The Cenocoeliinae are included in the morphology ot the larval cephalic structures re- Helconinae by iapek, (1970), because ol the (Short in Van Achterberg, 1979a). semblance in biology and larval slnlctures. However, the larval struclures are diff'erent from the Helconinae: the labial sclerite is more trans- Tobias, 1979 verse and the stipital sclerite is crooked (figs. 7-tt 29. Betylobraconinae versus figs. 9-ll + 14 in Capek, 1970). The Small subfamily. restricted to the Australian re- Cenocoeliinae form an isolated group because of gion and containing one genus: Belvlrthrnctttt the extremely high inserted metasoma and the Tobias. The biology is ttnknown, bul may be subpetiolate lst metasomal tergites of the im- peculiar considering the large number of apo- agines. The absence of a transverse scLrtellar de- morphous character-states. The Betylobraconinae pression, and the more or less developed post- are associated by Tobias ( 1979) with the Braconi- pectal carina (respectively, presenl and absent in nae-group, but I could not find any synapomor- the Helconinae) are other remarkable leatures. phous character-state to justify this placement. In The position of the Cenocoeliinae is uncertain the cladistic analysis the group comes near the and it is placed provisionally as an old group Orgilinae. but more synapomorphies arc needed basally in group lV. to be certain of this alignmcnt.

26. Agathidinae Blanchard, I tl45 30. Orgilinae Ashmead, I 900 Rather large and homogeneous subfamily, con- Rather small sublamily. containing three tribes: taining solitary or gregarious endoparasites ol Orgilini Ashmead (mainly Orgrlrrs Nees). Mima- lepidopterous larvae. Remarkable apomorphotrs gathidini Enderlein (mainly Staalrrrllrt Ash- character-states are the narrow marginal cell of mead), and Mesocoelini Viereck (provisionally lbre wing of adults and the paired fleshy ventral rncluded. contains Mesot oelus Schulz and processes in the lst larval instar (may be mod- Anettrobrucon Brues). This subfamily is often itied larval legs t. included in the Agathidinae because ofthe rather narrow marginal cell of fore wing. As shown in the cladogram the Orgilinae are not closely relat- ed. although both belong to group IV. The re- 27. Macrocentrinae Foerster. 1862 dLrction of the fore wing venation is most ex- Moderately sized subfamily, containing solitary treme in the Mesocoelini. which also have ex- or gregarious endoparasites of lepidopterous tremely long hind legs and spurs, and the scutel- larvae. Polyembryony is known of several spp. lum possesses a hemi-circular depression (even of a secondary solitary sp.) and the grega- medio-posteriorly. Especially the latter may in- rious specimens spin a common web. The most dicate that the Mesocoelini does not really be- striking apomorphy is the presence ol'small pegs long here and should be treated as a separate on the trochantellus of the adults. subfamily. 54 C. van Achterberg

30. ORGILINAE (inc1. Mesocoelini) 29. BETYIOBMCONINAE occipital carina very low 2nd metasomal spiracle in notum l-SR of fore wing clypeus not differ:- absent entiated 28. AUICROCXNTR]NAE narginal ce11 of frons !{ith deep lateral grooves fore wing slender and rather narrow maxillary fore coxae extreme- 27. MACROCENTRINAX or absent palp with 1y protruding pos- segment I ter i orly trochantell i lst tergite transverse Sroove hole letween toruli wi th sma11 with basal pegs plicat lobe reticu- middle lobe of mesoscutum pro- 2nd submarginal ce11 of fore truding wing smatl or absent hind coxa s trongly enlarged occipi ta1 carina absent metasoina inserted above epistomal sulure (part 1y) reduced bind coxae metasoma inserted nea! hind coxae slender lateral parts of 1ab iaI scleri te of final instar larva

26. ACATHIDTNAE

marginal ce 11 of fore larvae without fteshy appendages uing very narrou subpronope often present dorsal carinae of lst tergite often reduced lst insLar larvae lrith paired fleshy ventral cuspidal process oot separated appendages i-SR of fore wing reduced cuspidal process seParated 2-CU of hind \ring absent 2-CU of hiod i^,iog usually presgnt

25. CENOCOELIINAI parasites of lepidopterous larvae (except Neoneurinae) metasoma extremely high inserted postpectal carina absent postpectal carina present

transverse scutellar de?tession absent

Fig. I6. Cladog.ram of subfamily group IV, excluding IVa. Essay on the phylogeny of Braconi-dae (Hymenoptera: Ichneumonoidea) 55

34. I{ICROGAS?RINAE 35. KHOTKHOIINAX larval mandible lrith upcurved tlade latero-tergite of lst metasonal segment number of antennal not differentiated segment6 fixated ( l8) antennal segmelts variable, nore than 20 flagellar segment usually r^,ith 2 ranks of placodes 33. CARDIOCHILINAE

apical clypeal uargin s traight notutr! of lst metasomal te!8ite often not or weakly concave differentiated flom its epipleuron maxi l Iary palp with 5 segments base of larval mandible not differentiated from (nearly) straight blade

32. hypos tohal carina absent ovipositor curved do!.rrll,ards lst metasomal spiracle in epi- pleural area fore wing venation specialized

fore spur 0.7-1.0 times fore parasites of adults of ants bas i tarsus

marginal cell of fore ving oPen propleural flaoge and prepectal carina leduced lst metasonal tergite flattened

occipital carina atrsent

I -SR of fore wing subvertical 31. CHELONTNAE

CU Ib of fore wing absent ovo-1arval parasites ectoparas i tic phase lost of lepidopterous larvae metasomal carapace present postpectal carina present fietapleural f lange absent

marginal cell of fore wing short and wide or reduced eyes often setose spiracle of 7th metasomal segment absent

lr-o of hind lring short

Fig. 17. Cladogram of subfamily group IVa. 56 C. van Achterberg,

31. Cheloninae Nees. l8l6 morphies of the Neoneurinae are the un- or in- distinctly differentiated hind trochantellus, the Rather large subfamily. which consists of solit- usually strongly flattened and curved ovipositor, ary endoparasites of lepidopterous larvae. The the more or less incised hypopygium egg is deposited in the egg of the host; this may medio-posteriorly, and the spiracle of the lst be an adaptation Io the hosts (which live as larva tergite behind middle of the tergite. at less exposed places) as well as to avoid encap- sulation by the host. Easily recognizable group because of the metasomal carapace, combined with the presence of the postpectal carina. 33. Cardiochilinae Ashmead, 1900 Medium-sized sublhmily. containing solitary en- doparasites of lepidopterous larvae. The cep- 32. Neoneurinae Bengtsson, l9lt{ halic structures of the I'inal larval instar are simi- Small subfamily. containing endoparasites of Iar to those of several Microgastrinae. but the adults of worker ants: the development is in the mandibles lack the apomorphous condition of a metasoma of the host. Because of the biology differentiated base. The bifid apex of the larval (parasitism of adult insects) it has been inserted mandibles may be a synapomorphy with the in the Euphorinae by Tobias (1966). According Microgastrinae, although in the Microgastrinae to Tobias it is unlikely that the parasitism of several other (obviously apomorphous) condi- adult insects could develop twice within the tions occur. The adults are recognizable by the same family (Braconidae). However, this has vein SRI of the fbre wing strongly curved to- happened more than once. viz., in the Aphidii- wards vein l-Rl (straight or nearly so in nearly nae, in the Euphorinae (in three not closely re- all Microgastrinae, rather intermediate in the lated tribes), and, in my opinion, independenlly Khoikhoiinae and some genera of the Micro- also in the Neoneurinae. The downwards curvcd gastrinae, e.g.. Senioni.s Nixon) and the large ovipositor and the reduced wing venation are the 2nd submarginal cell of the fore wing (in- only characters which might allow inclusion in termediate in the Khoikhoiinae and in some gen- the Euphorinae. In the cladistic analysis pre- era of the Microgastrinae-Apantelini (Scnilorris sented in this paper it was not possible to insert Nixon and Pc,/lt opc Mason)). the Neoneurinae near or in the Euphorinae; actually numerous apomorphies allow an inser- tion near the Microgastrinae. The very long and 34. Microgastrinae Foerster, 1862 slender stipital sclerite of the final larval instar is trnique amo,ng the Braconidae so far as is known Very large and comparatively homogenous sub- (tig. 25 in Capek, 1970): the smooth mandibles tamily. consisting of solitary or (frequentlyt gre- is a parallelism (independently developed in the garious endoparasites of lepidopterous larvae. Microgastrinae as well in the Euphorinae) judg- Pupation is outside the host or its cocoon. Only ing from the other larval strrctures. in the small genus /)r,rrlcfl.rys Mason is the Besides lhe synapomorphies with the pupation in the host cocoon. apparently a M icrogastrinae-group mentioned in the clado- sccondary development. The gregarious speci- gram, several tendencies occur both in the mens spin a common web, sometimes with the Neoneurinae and in the M icrogastrinae-grortp. cocoons very regularly arranged. A cladistic e.g.. antennal segments of 9 with pseudodivi- analysis is given by Mason ( I9tl I & l9tl3l: as sion (most Microgastrinae and in the treated in this paper an easily recognizable group Neoneurinae: l'arc kt s rttosortttt Tobias & because both sexes have antennae with 18 seg- Yuldashev), d antenna sometimes compressed ments. The tribe Apantelini Viereck s.l. (includ- and widened. absence ol'the lateral carina of the ing the Microgastrini sensu Mason) contains the mesoscutum, vein SRI of the fbre wing some- species with the largest number of plesiomor- times with ramellus (as frequently in the Cardio- phous character-states. e.9., the morphology of chilinae), vein C+SC of fore wing more or less the lst metasomal tergite, the ovipositor sheath widened distally, and vein 3-CUl of fore wing of the adults and the mandibles of the final larval not angled with vein 2-CUl. Striking autapo- instar. Essay on the phylogeny of Braconid,ae (Hymenoptera: Ichneumonoidea) 57

35. Khoikhoiinae Mason, 1983 phora erythrocephala Meigen, p. I 91, figs. l-13, plates l-12.-Thesis, Leiden. Small subfamily, of which the biology is un- Chamberlin. F. S. & J. N. Tenhet, 1926. Cardiochiles parasite to- known. Known only from South Africa and diffi- nigriceps Vier., an important of the bacco bud worm Heliothis virescens Fab.-J. agr. to differentiate from the Microgastrinae with cult Res. 33: 2l-27. figs. 1 4. autapomorphous character-states. The only Daniel, D. M.. 1932. Macrocentrus ancylivorus autapomorphy seems to be the striated epi- Rohwer, a polyembryonic braconid parasite of the pleural membrane (lacking a more or less ditl'e- Oriental fruit moth.-Techn. Bull. N.Y. State Agr. l-101, figs. 1-18. present Micro- Exp. Srat. 187: rentiated latero-tergite as in the Edson, K. M. & S. B. Vinson. 1979. A comparative gastrinae) ofthe lst metasomal tergite. Unfort- morphology of the venom apparatus of female bra- unately the latero-tergite of the lst segment conids (Hym., Braconidae).-Can. Ent. I ll: (wherein the spiracle is situated) is oflen scarcely l0l3-1024, figs. l-2, table l. Griffiths, G. c. D., 1964. The Alysiinae (Hym., makes a differentiated in the Microgastrinae, it Braconidae), parasites of the Agromyzidae (Dip- difficult character to interprete. Nevertheless the tera).-Beitr. Ent. l4: 832-914, figs. l-38. Khoikhoiinae are easily recognizable because of Haeselbarth, E., 1 2. Zw Biologie, Entwicklungsge- the long vein 3-SR of the fore wing, the lst ter- schichte und Okologie von Brachistes atricornis Ratz. (Hym., Braconidae) als eines Parasiten von gite groove and with with a long medial antennae Pissodes piceae (lll.) (Cot., Curc.).-2. ang. Ent. more than [8 segments. 49: 233-289 , figs. l-26. Jackson, D. J., 1928. The biology of Dinocampus (Perilitus) rutilus Nees, a braconid parasite of L. (pt. 2).-Proc. zool. Soc. London: References Sitona lineata 597-630. Achterberg, C. van, 1974. The features of the petiolar Loan, C. C. & F. G. Holdaway, 1962. Pygostolus segment in some Braconidae (Hym.).-Ent. Ber., falcatus (Nees) (Hym., Braconidae), a parasite of Amst. 34: 213-214, figs. 14. Sitona species (Cot., Curculionidae).-Bull. ent. 1976. A preliminary key to the subfamilies of the Res. 52: 473-488, figs. l{. - Braconidae (Hym.).-Tijdschr. Ent. I l9: 33-78. Mason, W. R. M., 1981. The polyphyletic nature of figs. l-121, I table. Apanteles Foerster (Hym.: Braconidae): a 1979a. A revision of the species of Amicrocen- phylogeny and reclassification of Microgastri- - trinae, a new subfamily (Hym., Braconidae), with a nae.-Mem. ent. Soc. Canada I 15; l-147, figs. descriplion of the final larval instar of Amicrocent- l-t 10. rum curvinervis by J. R. T. Short.-Tijdschr. Ent. 1983. A new South African subfamily related to 122'. l-ZE. figs. l-68. - Cardiochilinae (Hym.: Braconidae).-Contr. Am. 1979b. A revision of the new subfamily Xip- ent. Inst. 20:49-62, figs. I 21. hozelinae (Hym-, Braconidae).-Tijdschr. Ent. Nickels, C. B., W. C. Pierce & C. C. Pinkney, 1950. l2Z:2946, figs. 145. Parasites of the Pecan Nut Casebearer in 1979c. A revision of the subfamily Zelinae auct. Texas.-Techn. Bull. U.S. Dept. Agr. l0ll; l-21, - (Hym., Braconidae).-Tijdschr. Enl- 122: 241'479, figs. l-7. figs. l-900. Oatman, E. R., G. R. Platner & P. D. Greany, 1969. 1983a. A revision ofthe new tribe Brulleiini (Hym.. The biology of Orgilus lepidus (Hym.: Braconidae), - Braconidae).-Contr. Am. ent. lnst. 20: 281-306, a primary parasite of the Potato Tuber- figs. I l0l. worm.-Ann. ent. Soc. Am. 62: 1407-1414, figs. 1983b. Revisionary notes on the subfamily t-6. - Gnaptodontinae, with the description ofeleven new Parker, D. L., 1935. Apanteles solitarius (Ratzeburg) species (Hym., Braconidae). Tijdschr. Ent. 126: an introduced braconid parasite of the Satin 25-57, figs. 1-131. Molh.-Techn. Bull. U.S. Dept. Agr. 477: l-17, Allen, A. A., 1977. Exhibits. Proc. Brit. ent. nat. figs. I -l't. hist. Soc.: lll ll2. Rosenberg, H. T., 1934. The biology and distribution Brindley, M. D., 1939. observations of the life-history in France of the larval parasites of Cydia pomonella of Euphorus pallipes (Curtis) (Hym., Braconidae) a L.-Bull. ent. Res. 25:232-256, figs. I 15. parasite of Hemiptera-Heteroptera.-Proc. R. ent. Saffer, 8., 1982. A systematic revision of the genus Soc. London 14: 5l-56, figs. l-4. Cenocoelius (Hym., Braconidae) in North Capek, M., 1965. The classification of Braconidae America. including Mexico.-Polskie Pismo ent. (Hym.) in relation to host specifity.-Proc. l2th 52:71-16'1, figs. l-53. Intern. Congr. Ent. London. p.98-99. Shaw, M. R., 1981. Some parasites (Hym.) of Epip 1970. A new classification of the Braconidae hyos postvittana (Walker) (Lep.: Tonricidae) in - (Hym.) based on the cephalic structures ofthe final Cornwall.-Entomologist's Gaz. 32: 3G38. instar larvae and biological evidence.-Can. Ent. Simmonds. F. J., 1 7. The biology of the parasite of 102: 846 875, figs. l-58. Loxostege sticticalis L. in North America- Caudri, L. W. D.. l%1. The braconid Alysia man- Meleorus loxostegei Vier (Brac.. Meteorinal]). 1-6. ducator Panzer in its relation to the bow-fly Calli- -Bull ent. Res.38: 373-379. figs. 58 C. van Achterberg

Southwood, T., 1957. Observations on swarming in pean corn borer.-Tech. Bull. U.S. Dept. Agr. 294: Braconidae (Hym.) and Coniopterygidae (Neurop- l-48, figs. l-l E. t€ra).-Proc- R. ent. Soc. London 32:6O-82. Starf, P.. I 6. Aphid parasites of Czechoslovakia. p. l-242. figs. l-112. photos l -19.-Academia. Sammanfattning Prague. Timberlake, P. H., 1916. Note on an interesring case of Parasitslekelfamiljen Braconidae omfattar minst two generations parasite of a reared from the same ca 40 000 individual host.-Can. Ent. 48: 89-91. arter. Utseendet hos en representant Tobias. V. I.. 1 5. Generic groupings and evolution of lor gruppen visas i figurerna l-ll. Viktiga parasiiic Hymenoptera of the subfamily Eup- karakfirer som skiljer familjen Braconidae frin horinae (Hym.. Braconidae), pt. l.-Ent. Obozr. dvriga lchneumonoidea er att 2:a och 3:e bak- 4l: 841-865. Russian (transl.: Ent. Rev. 44: 494-507). kroppssegmentens ryggplatar hr ftirenade hos 1966. ld., pt. 2 Ent. Obozr. 45: 612-633. Russian Braconidae och att vingnerven l-SR+M (skrivs - (transl.: Ent. Rev.45: 348,358, figs. l-14). oftast Rs +M) finns kvar hos Braconidae men ej l 7.A reviewof the classification, phylogeny and hos ovriga Ichneumonoidea (se Fig. - (Hym.).-Ent. l2). evolution of the family Braconidae presenterar Obozr.46: 645-669. Russian (transl.: Ent. Rev.46; Arbetet en kladistisk analys av 387-399, figs. l-40). huvudgruppema inom familjen Braconidae. Ana- - & G. P. Dudarenko. 1974. Genral trends in evolu- lysen resulterar ien klassificering diir familjen tion of Braconidae (Hym.) abdomen.-Vest. Zool.: indelas i35 underfamiljer grupperade i fyra 65-72, figs. l-2. Russian. stiirre avdelningar (Aphidiinae grupperna 1979. Two new and one little known subfamilies of samt - Braconidae (Hym.) from Australia.-Ent. Obozr. II-lV. se Fig. l3-17). Den kladistiska analysen 58: 128-142, figs. l-37. Russian. kan lAtt f