Systematics of the tribe Euderomphalini (Hymenoptera: Eulophidae): parasitoids of whiteflies...

Systematics of the tribe Euderornphalini (Hymenoptera: Eulophidae): parasitoids of whiteflies (Hornoptera: A I e y rod i d ae) J O H N LASALLE and M I C H A E L E. S C H A U F F " International Institute of Entomology, c/o The Natural History Museum, London, and .5ystemntic Entomology Lab/USDA. c /o U.S. National Muscum, Washington, D.C.. U.S.A

Abstract. Eulophid parasitoids of whiteflies are reviewed, with comments on their systematic placement, and a generic key. All eulophid whitefly parasitoids are placed in the tribe Euderomphalini in the subfamily Entedoninae, and reasons for this placement are discussed. Three new genera are described. The seven included genera can be divided into two distinct genus groups: the Eiidcromphalc group contains B(ieocwtcdm Gir:iult, ELiilcromphale Girault, Neopomphdc gen.n., and Pomnplicik Husain cJt id.; the Entcdononecrrrnnus group contains Ak~wroctoniis gen.n., D(isyonrpliti1c gen.n.. and Entedonon- ccrcmiiiis Girault. The new species Diisyomplicilc clii1ciisi.s is described.

introduction

'lhcrc ;ire over 1 IS0 spccics of whitcHics (Homoptcra: Alcyroclitlac) (Mound L" Hdscy , 107s). all o f which use wckirig mouthparts t o fccil on soft plant tissue. Because of thcir w;isy covcring. high fecundity, and ability to develop rcsistancc rapidly. they have proved to be extremely cliflicult to control through the use o f insecticides, and many spccics have rcached economic importance. Their small size and ability to survive transport has further allowed many species to increase their distribution through invading new areas of the world. In addition to direct damage t o the plant, whiteflies can cause indirect damage through the production of honeydew which induces sooty mould, mid through the transmission of plant viruses.

Although whiteHies are often resistant to pesticides, their sessile immature stages and wide range of natural cncmies make them highly amenable to biological control. Due to the various biological control successes against whiteflies, there is currently an increased interest in and awareness of their natural enemies.

The majority of whitefly parasitoids are in the family Aphelinidae. particularly in the genera Eticarsia and Erefmocerr~s; however, whitefly parasitoids are also known in the families Piatygasteridae (genus Amitus), Signi- phoridae (often as hyperparasites) and Eulophidae. This lattcr family contains a relatively small proportion of

Corrcspondcncc: Dr john LaSalle. Intcrnational Institutc of Entomology, c /o Thc Natural History Museum, South Kcn- \ington, London SW7 5BD.

whitefly parasitoicls. i ind thcsc parmitoitls have n o t gained attention as effcctivc biological control agents. For example. reviews of succcssful biological control programmes (Laing s( Hamai. 1Y7h: Clnuscn, IY7X) listed no species o f eulophids as having been importcd against whitcHy pests. However, hccausc this group is difficult and poorly known taxonomically, their v d u c might hnvc been overlooked t o some extent.

Most o f the genera of Eulophidae which are now known to attack whiteflies are not found in existing keys to genera. Eutlerornphale is the best known of these genera. and it has only been included in keys to genera for Europe (Peck ef al., 1964: Trjapitzin, 1Y78). India (Hayat, 1985) and Australasia (Boufek. 1988). Pomphale has only been included in a key to Indian genera (Hayat, 1YSS): however. at that time it was not known to be a whiteHy parasitoid. Baeoentedon was included in a key to Australasian genera (Boufek, 1988). although Bourek presumed that it was a parasitoid of Psylloidea (biological information for Baeoentedon is still unknown; however, we would suspect that it is a whitefly parasitoid based on its clear relationship with other genera of the Euderomphalini). Eiiredoiioii- eciemiius has never been included in any generic keys to the Eulophidae, and the remaining three genera of whitefly parasitoids are described i n this paper.

Perhaps because Euderomphule has often been the only eulophid genus mentioned to attack whiteflies, it has become a dumping ground for small eulophids reared from whiteflies. For example, two species originally described in Euderomphale (viftafu Dozier, aleurorhrixi Dozier) are placed in the genera Alritroctoriirs and Neopurnphalr

235

236 Johti LaSalle uird Michael E. Schaldff

respectively in this paper. The small size and confused taxonomy in this group has also produced mistakes at the family level. A.vifrafus was included in the genus Emorria (Aphelinidae) in a recent catalogue to Neotropical Chal- cidoidea (De Santis, 1979: 330). Another species described and since treated as an aphelinid, Pferoptrix australis Brttthes, is transferred to the genus Neopomphale in this paper.

This paper will enable workers to recognize more easily the eulophids associated with whiteflies, and in doing so the increased knowledge gathered about these parasitoids might show that they are more common than previously suspected. It is hoped that such knowledge may prove useful to future biological control attempts.

Thus the purpose of this work is varied. I t began as the description of two genera known from Florida (Akur- octomis, Neopomphalr), which were necessary to a key to genera of North American Eulophidae which is currently in preparation. However, because of the interest in natural enemies of whiteflies, and the general lack of knowledge of the eulophid whitefly parasitoids, all species of eulophid whitefly parasitoids are reviewed and a generic key is presented. Finally, the placement of these species within eulophid higher classification proved problematical, and these problems are discussed with our justification for the classification we employ.

Methods

Basic morphological terminology is taken from Schauff (1991). with the terminology for sulci o n t h e head somewhat modified. The scrohal sukcus (Figs 10- 12) is a longitudinal sulcus which is placed in the scrobal cavity. The Jronral ,suicu.~ is il transverse sulcus, which can be straight (Figs 10-12. 37) , curved (Fig. 42), or v-shaped (not found i n Euderomphalini, but frequently seen in other entedonines), which is ventral to the median ocellus, and often joins the scrobal sulcus. The vertexal sulcus is a transverse sulcus which is placed on the vertex, and extends from the eye margin either between the median and lateral ocellus (Figs 3. 4, 11, 12), or just posterior to the ocelli (Figs 23, 24). Schauff (1991) referred to a similar sulcus as the occipital suture (in reference to a group of genera related t o Ceranisus): however, this is more properly termed a vertexal sulcus as it is on the top of the lead (vertex) rather than the back (occiput). It should be noted that the terms scrobal, frontal and vertexal are used here in an adjectival sense simply to denote where a sulcus is found, and are not meant to imply homology with similar structures in other groups.

The distinction between types of antenna1 segments can cause some confusion, particularly between anelli and funicular segments. In some cases a segment can appear to be intermediate between a large anellus and a small funicular segment (for example, Figs 54, 58, 59). This is a problem which confuses specialists in the group as well as beginners. As a general rule, funicular segments will have linear sensilla (also termed rhinaria or multiporous plate

sensillae); and anelli. although they sometimes have small setae, will lack sensilla. Using the key and descriptions in conjunction with the illustrations should prcvent confusion.

The outer plates of the ovipositor are formed from the lateral parts of the ninth abdominal tergite (seventh gastral tergite) and have become involved in the ovipositor mechanism (Copland Sr King, 1972). This tergite is most easily recognized because i t bears the cerci.

I t should be noted that these are extremely small wasps. and difficult to study. Additionally, some of the characters, such as the sulci on the head, are not easily seen. We have tried to use characters in the key that are relatively easy to see, but have incorporated all characters into our attempts to generate a classification.

Taxonomy of whiteflies is taken from Mound Sr Halsey (1Y78). See this work also for any host records not listed in references to each species.

Abbreviations for institutions and collections: AEI, American Entomological Institute. Gainesvillc, Florida: AMU, Aligarh Muslim University, Aligarh, India: BMNH. The Natural History Museum, London. U.K. ; CNC, Canadian National Collcction of Insects a n d Arachnids. Ottawa: LAS, collection of 1. LaSallc; QMB. Queensland Museum, Brisbane, Australia; TAMU, Texas A&M University, Collcgc Station; UCR, University of California Riverside; UFG, University of Florida. Gainesvillc: UQ, University of Quecnsland, Australia: USNM. The U.S. National Museum o f Natural History.

Abbreviation used f o r morphology: PI , F2. funicular segments 1 and 2.

Tribe Euderomphalini

The tribe Euderomphalini was first proposed by Shafce ef oi. (1988) within thc subfamily Entcdoninae to contain the single genus Eurleromphule. They distinguished i t from other entedonines by the following characters: axilla entirely situated anterior to scuto-scutcllar suture; notauli completely absent: pronotum with a membranous area medially. Of these characters, only the position of the axilla is informative. The notauli are absent in almost all entedonines and many eulophines, and although the pronotum is strongly narrowed in Euderomphalini, it is not always membranous medially even within the genus Euderomphale.

BouEek ( 1988) gave the following additional characters for distinguishing Euderomphale from other Entedoninae: scutellum much broader than long; hind margin of scutellurn with an additional pair of setae; antenna clavate; funicle shorter than pedicel, two-segmented, with the first segment smaller than the second. Some of these characters serve to distinguish Euderomphale, although others can be used at a higher level within the tribe (see below).

Other unusual aspects of Euderomphale had already been mentioned by Compere Sr Annecke (1961). They pointed out that the sixth gastral tergite is mainly membranous, with the sclerotized portion reduced to a small plate which bears the spiracle (Fig. 9) (this is true only in

Sy vtettircfics 01 the Eriderompliulit~i 237

I t may be that these characters are linked as a general expansion of the dorsum of the mesothorax. Eprhopalotus. mentioned above, also displays this scutellar character, but otherwise does not seem particularly related to the Euderomphalini (possesses complete notauli. and more ‘typically entedonine’ antenna and clypeus).

Male scape with a ventral groove present iti apical hulf, but without an easily visible ridge or sensory pores (Figs 71-77). This type of male scape is only found in the Euderomphalini, and seems to be a modification of the type of male scape usually seen in Entedoninae. This character is discussed further below under synapomorphies for the Entedoninae.

Ciypeus delimited dorsally by a complete, distinct and smoothly curved sulcus (Figs 62-70). This character is somewhat equivocal, because there is variation within the tribe. However, the clypeal sulcus is distinct and smoothly curved in Euderomphale (Fig. 62), Entedononecremnus (Figs 67, 68) and Aleuroctonus (Fig. 69). It is not as distinct and clearly delimited in the other genera. How- ever, we would suspect that the complete and smoothly curved clypeal sulcus is a synapomorphy for the Euderom- phalini, and any deviations from this form are subsequent reductions.

Antenna reduced, with 2 or I funicular segments; funicle short, generally shorter than pedicel; club long, generally longer than the funicle, and with a long terminal spine (Figs 54-60). This type of antenna is found throughout the Euderomphalini, and is rare in other Eulophidae.

Axiila, when indicated, generally pluced entirely anterior to scuto-scutellar suture (Figs 7, 17, 21, 26, 39, 44). This is unusual for Entedoninae, and is found in all genera except Entedononecremnus, where some species have the axilla only partially advanced forward of the scuto-scutellar suture (Figs 31, 32). Hosts are whiteflies. All Euderomphalini whose biologies

are known are parasitoids of whiteflies. This biology is not known in any other Eulophidae. Any other classification would require the polyphyletic origin of whitefly parasitism within the Eulophidae. This cannot be entirely ruled out, because biology and morphology both seem to be equally plastic in the Eulophidae.

[he female. seems t o be restricted to Euderornphulu, ;ind may not appear in all members of the genus), and that the seventh and eighth tergites are fused (as in all culophids except Euderinae). They also discussed the unusual shape of the prepectus in this genus, which was reduced and fused to the mesopleuron (Fig. h), and stated that ‘the mesopectus is so greatly unlike that of typical tulophids as to raise a question as to whether they are placed here correctly’ (Compere & Annecke, 1961: 68). This character is found only in the genera Euderomphale and Neopomphale (Figs 6, 14).

This paper expands the definition of the Euderomphalini, and includes several genera which had not been previously placed here. Within the Euderomphalini are two distinct groups, which we are here treating with the informal categories of genus groups. The Euderomphale group contains Baeoeritedoti, Euderoniphale, Neopomphale and Pomphale; the Etrtedotiotiecremtius group contains /Ileuroctoti!is, Dasyomphale and Entedononecremnus.

Diagtmsis of the tribe Euderomphalini

Sniiill parasitoids, generally less than 1.5mm in length (sonic species of Ditedonotzecremtzus up to 1.8 mm). Pronotum (Figs 5 , 13, 29) reduced, vertically aligned and not visible in dorsal view, and strongly narrowed or memhronous medially. Scutellum (Figs 7, 8, 15, 16, 20, 15, 31, 38, 43) overhanging dorsellum; median area of dorscllum not visible in dorsal view. Notauli absent. Antenna (Figs 54-61) reduced, with 2 or 1 funicular segments; funicle short, generally shorter than pedicel; club long, generally longer than the funicle, and usually with a long terminal spine. Axilla, when indicated, generally placed entirely anterior to scuto-scutellar suture. Clypeus (Figs 62-70) generally delimited dorsally by a complete, distinct and smoothly curved sulcus, although this reduced or modified in some groups. Male scape (Figs 71-77) with a ventral groove present in apical half, but without a visible ridge or sensory pores present within groove.

Monophyly of the tribe Euderomphalini Discussion and relationships

Although there is a great deal of variation in the general appearance OE the two genus groups in the Euderomphalini, the following characters support the overall monophyly of the tribe.

Pronotum reduced, vertically aligned and not visible iti dorsal view, arid strongly narrowed or membranous medially (Figs 5, 13, 29). All Euderomphalini have the pronoturn of this form. It is rare in other Entedoninae, but present in certain genera such as Eprhopalotus (see Schauff, 1991, as Aabacharis).

Scittellum overhatrgirig dorsellum; median area of dor- Jellurn not visible in dorsal view (Figs 7, 8, 15, 16, 20, 25, 31, 38, 43). As with the previous character, it is present In all Euderomphalini, and rare in other Entedoninae.

There are two main problem areas with the treatment of the Euderomphalini presented in this paper: overall support for the monophyly of the Euderomphalini, and the relationship of the Euderomphalini to the rest of the Eulophidae.

Support for the monophyly of the Euderomphalini

Within the Euderomphalini, we are confident with the monophyly of each of the two genus groups. However, the placing of both groups within a single tribe Euderomphalini should be considered as preliminary, and future studies

238 John LaSullr uirrl Michuel E. SchulrJ””

Figs 1-9. Euderomphole spp.. P: 1, hcad: 2, malar sulcus: 3, vertex: 4, vcrtcxal sulcus: 5 . pronoturn; 6, mcsosoma, latcral: 7. mcsosorna, dorsal; 8. scutellurn: 9, apcx of gaster. a, axilla; rn, rncsoscutum: mp, rncsopleuron; rns. malar sulcus: p. pronoturn: pp. prcpcctus: S. scutellum; st, scutellar seta: T5. T6, gastral tergitcs five and six: vs, vcrtcxal sulcus.

may provide characters which will refute this classification. This is particularly true hecause the genus groups are sufficiently distinct that members have never previously been placed in the same subfamily.

Despite the differences between the groups, we have supplied several characters that s ~ p p o r t the monophyly of the tribe as a whole, and in the absence of further evidence we consider this classification as the most valid hypothesis. In addition to morphological characters. the utilization of

whiteflies as hosts is considered to be a derived character to support this group. Any other classification would require the polyphyletic origin of parasitism on whiteflies.

Syrznpomorphies for the Eirtedorzirrcre

Schauff (1991: 10- 13) listed and discussed several characters which he considered synapomorphies for the

Figs 10-18. Neopotnphale spp., 9 : 10, head; 11, vertcx: 12. hcad; 13, pronotum: 14. mesosoma, lateral: 15. mesosoma, dorsal: 16. mcsosoma. dorsal: 17, axilla. scutcllum. propodeum; 18, basc of gastcr. a, axilla: fs, frontal sulcus: m, mcsoscuttim: mg, membranous arca at basc of gastcr: mp, mcsoplcuron: rns, malar sulcus: p. pronoturn: pp, prepectus: s. scutellum: ss, scrobal sulcus: vs, vcrtexal sulcus.

Entedoninae: (1) the scutellum with a single pair of setae; (2) male scape with sensory pores restricted to the ventral edge: (3) submarginal vein with 2 dorsal setae; (4) mesoscutal midlobe with 2 pairs of setae; (5) position of frontal sulcus: (6) subspiracular propodeal tubercle: (7) marginal vein much longer than submarginal: (8) short stigma1 vein: ( 9 ) submarginal vein broken. Of these characters, Schauff (loc. cir.) felt that the most important were the scutellum with a single pair of setae, and the male

scape with the pores restricted to the ventral edge. He considered the last three characters to be particularly unreliable, and we will not consider them further.

(1) Scutellicm with a single pair of setae. The Entedoninae generally have only a single pair of setae on the scutellum, and this has been considered a good synapomorphy for the subfamily. The primitive condition in the family would be two to several pairs of setae, as found in the other subfamilies. There are exceptions to the presence of only a

240 J o h t i LiiSulle utiil Micliuel E. SchoiiJj

single pair of scutellar setac within the Entcdoninac, notably Parahorismenits and P1eirrolropi~"p.si.s (scc Boui-ek, 1988), and a few aberrant species of Pediohi1i.s such as .vetigrrits (Kerrich, 1970) and mu1ti.scrie.s (BouEck, 1976). so this character has arisen independently on more than one occasion within the Entedoninae, and an additional appearance would not in itself preclude a taxon from inclusion. Myrniuknra diaprioides BouEek has no setae on the scutellum (BouCek. 1072).

Only three of the seven genera (Pomphule, Neopom- phule, Baeoentedori) have a single pair of setae on the scutellum (Figs 15. 16, 20, 25). Euderomphale has two pairs, although the posterior pair is distinctly reduced in size compared to the anterior pair (Fig. 8). The other generu (Etireclorioriecremtius genus group) all have many scattered setac on the scutellum (Figs 31, 38, 43).

(2) Male sciipe with sensory pores restricted to the ventral edge. The distribution and location of the sensory pores on the male scape have been used to indicate the monophyly of the Entedoninac (Schauff. 199 1) and the Tetrastichinae (Graham, 1987). I n entedonincs the pores are restricted to ;I spucializcd structure on the ventral edge of the scape. which is in the form of a longitudinal ridge which is situated i n a groove which extends for nearly the entire lcngth of thc scape (Schauff, 19Yl). This structure is not found in other eulophids.

We find this to be one of the best characters to support thc placement of the Euderomphalini in the Entedoninae. Whcrc malcs wcre cxaniined (all genera except Bueoetrtedot~ and Dosyomphule), the ventral edge of thc scape has a distinct groovc along the vcntral margin (Figs 71-77). However, this groove is restricted to the anterior half of the scape, and the ridgc (and, indeed, the pores) are not distinctly visible within the groove. But, since no other eulophids have the groove, we consider this a synapomorphy with the rest of the entedonines. The unusual structure of the groove is further support for the monophyly of the Euderomphalini.

( 3 ) Submurgirral vein with two dorsalsetue. This character has often been used as one of the defining synapomorphies of the Entedoninae. All Eulophinae and Euderinae have 3 or more setae on the sumarginal vein: the Tetrastichinae are quite variable, and can have one to several setae. Within the Entedominae there are exceptions to this character: Myrmokata diaprioides Boufek has only a single seta on the submarginal vein.

Within the Euderomphalini there is also variation. Euderomphale and Neopomphule have two pairs of setae on the submarginal vein (Figs 46, 47); Pomphale and Baeoentedori have a single setae on the submarginal vein (Figs 48.49). Two pairs of setae seems to be the groundplan condition in the Euderomphale genus group, so would not exclude their inclusion in the Entedoninae. The Entedononecremtius genus group has 3 or more setae on the submarginal vein (Figs 50-53); no other Entedoninae are known which have more than 2 setae on the submarginal vein.

(4) Mesoscutal midlobe with two pairs of setae. Most entedonines have only two (or occasionally one) pairs of

setac on the niidlobc of the mcsoscutum, although more setae arc present in genera such as Pleierorroppopsis and Paruhori,smc.rius (see Boutek, 1988) and some aberrant Pediohiics. This character state also occurs occasionally (and quite independently) in other subfamilies (e.g. Hy.s.sopi4.s in the Eulophinae, Tumnrixia in the Tetra- stichinile). This is a rather weak character, as it is fairly homoplastic throughout the Eulophidae.

Within the Euderomphalini, the Euderomphule group has one or two pairs of setae on the mesoscutal midlobe (Figs 7, 15, 16, 20,25): the Eiitedoriotzecremtius group has many scattered setae (Figs 31, 38, 43).

( 5 ) Plucemetzt of frotitul silkus. Within the Entedoninae the frontal sulcus, which is generally transverse or v- shaped, is always separated from the median ocellus. It is generally placed about midway between the median ocellus and torulus, but if close to the median ocellus, it is still always separated by a distance greater than the diameter of the rncdian ocellus. This sulcus can occasionally be abscnt or represented merely be a changc in sculpture. particularly in the case of strongly sclerotized species, such as many species of Elitedon (Schauff, lY8X).

Within the E~',trrdorionecremrius group, the frontal sulcus is absent in Etitedotiotircrcmtiirs (Fig. 28), and present in Alei4roctotici.s and Dasyomphde. In these latter two genera it is separated from thc median ocellus by a distance distinctly greater than the diameter of the ocellus in A1euroctotirt.s (Fig. 37). and by a distance about equal to the diameter of the ocellus in Dusyottrphuk (Fig. 42). Within the Euderomphale group. t h e frontal sulcus (present in Neopornphule, faintly indicated in Pomphale) is separated from the median ocellus by a distance no greater than the diameter of the ocellus (Figs 10-12).

We consider the placement of the frontal sulcus as one of the better characters to support the monophyly of the Entedoninae, and thus the presence of a different character state in the Eucleronzphale group may be one of the major weaknesses of our proposed classification.

(6) Subspiraciclur propodeal tubercle. Most Entcdoninae possess a small bump or tubercle behind the propodeal spiracle near the posterior edge of the segment. With- in the Euderomphalini this tubercle is only distinct in Etitedottotzrcrem~zits; however, it is of a very typical entedonine form in this genus (Fig. 32). It is absent or indistinct in the remaining genera (Figs 8, 17, 21, 26, 39, 44). However, this character may also be absent or extremely reduced in many other small Entedoninae, such as Chrysotzotomyia, Ceranisus and its allies, and Asecodes. The absence of this tubercle in most euderomphalines would not necessarily exclude their placement in the Entedoninae, particularly as this is a character which seems to be frequently lost with a reduction in size.

Support for the placement of the Euderomphalitii in the Etitedoninae

The best character to support the placement of the Euderomphalini in the Entedoninae is the ventral groove

Systemalics of rlir Eudrmmphulirii 23 1

Figs 19-27. (1Y-22): Pomphale srripripennis, 9 : 19, head; 20, mesosoma; 21, axilla, scutellum, propodeum: 22, antenna. (23-27): Baeoenredon sp. P : 23, vcrtex; 24, vertexal sulcus; 25, mesosoma: 26, axilla, scutellum, propodeurn; 27, propodeum showing pit with sensilla. ps, pit on propodcum with sensilla; ss, scrobal sulcus; vs, vertexal sulcus.

on the male scape, which is present and consistent in form in both groups. Other important characters for defining the Entedoninae are the number of setae on the scutellum, submarginal vein and mesoscutum, and the shape and position of the frontal sulcus.

The character states found in the two genus groups are conflicting, as each group possesses character states that are both typical and atypical for the Entedoninae. The Euderomphalr group has setation (of both the meso-

soma and wing) which is typical of Entedoninae, but does not possess an entedonine style frontal sulcus. The Entedononecremnus group has an entedonine style frontal sulcus, but does not have typical entedonine setation.

Despite the fact that neither of the genus groups possesses all the characters which have previously been used to define the Entedoninae, the presence of the ventral groove on the male scape, as well as the fact that they seem to fit here better than in any other existing subfamily,

Table 1. Spccitic host assr>ci;rtioiis for Eudcromphalini.

Alcyrodidac species Eudcromphalini spccics

Alcyrodinac Aleriroclii!oti uccris (Modccr) A1ciirocyboiii.s oc.cidiiia Russcll Akirrolobiis burotlensis (Maskcll) Aleiirolobcrs sp. A/eriroiliri\-its ~Ynccosirs (Maskcll) A leyrodes uir rcocinciu Cockcrc I I

A Ieyrodes proletcllu Li nnacus ' Aleyrodes lotiicerue Walkcr

Aleyrodes prriitiosiis Bcmis Alqvroiles spirueoidcs Quairitancc Asrerobemisiu curpitii (Koch) Bemisiu uj?r (Pricsncr 19 Hosncy) Terrulerrrodrs sp.

Alcurodicinac Ahiirodicirs unrillctisis Dozicr Aleiirodicits cocoi.s (Curtis) A leiirodicia dispcrsiis R ussc I I Ceru1rirrodicri.s oliissirnris (Quaintaiicc) Eiitlialeirroilicirs borlkitii Quaintancc B Bnkcr Undctcrmincd Alcurodiciniic

Eirrleromplrule secrciu Euclerornphnle liyrilitiu Eirtlcrotnp Aale secitndirs Pornpliule srripripetitiir Neopotnplicile ulcicrorliriri Eiidcrotnplirile jlciviinediu Eirclrrotnplinle clielidonii Eitderotnplicile cerris Eirtleromp~idc dididotiii Endcrotnphole jin i-irncdiu Etrtlcrornpliirle jfu vitnetliu Errderotnplntle cerris Eiitlerr~rnplicrlc bertiisicrc Etrderotnplrtile qirrrcicnltr Nropornplrtrlc sp.

leads us to place the Eurleromphalini in thc Entcdoninac. We are well aware that further analysis is necdcd to fully

undcrstand higher relationships within thc Eulophiclac, and that future studies may provide information which will refute this classification.

Biology

Eulophidae are predominantly parasites of Lepidoptera, Diptera and Coleoptera, although a wide range of other hosts i s known, particularly in the Tetrastichinae and Entedoninae. These latter two subfamilies contain (either as primary or secondary parasitoids) the only eulophids which attack Homoptera (mainly Coccoidea and Psy- Iloidea). All species of Euderomphalini whose biology is known are parasitoids of whiteflies, and these are the only eulophids known to attack whiteflies.

Host relationships of the Euderomphalini are given in Table 1 (specific host relationships) and Table 2 (generic relationships and distribution). Although host records are sparse, it appears that the two genus groups have specialized on different subfamilies of Aleyrodinae; the Euderomphnle group on Aleyrodinae, the Efrtedonorlecremtiils group on Aleurodicinae. This host specialization is supported to some extent by distribution. The Aleurodicinae are almost entirely New World in distribution. although a few species from outside this region are known (Mound, 1984): and the Entedononecremtzus group is entirely New World in distribution. The Aleyrodinae are worldwide in distribution; the genus Euderumphale is almost cosmopolitan in distribution, Pomphule is from the Indian subcontinent,

Bwoet rkcluti from Sou t hcast Asia and Australia, and Neopotnplritle is Ncotropicill.

Biologicul cotztrol. Thc taxonomic confusion concerning this group may have concealed the presence of econonii- cally important species. The spiralling whitefly, Alcrtrotlictrs di.sper.sw, is a serious pest which attacks over forty spccies of plants (Mound & Halsey, 1978). A1euroctoriir.s Iirtutiis has bccn crcditcd with providing fortuitous biological control of spiralling whitefly in Florida (Bcnnctt Sr Noycs. 1989). and should be considered for importation into other areas where this whitefly is currently causing economic damage. Neopon?p/ruke ahrrotltrixi has been reared from the woolly whitefly. Al~~rrothrisirs~ucco.s~t.s, a pest species with a wide host range but which is particularly damaging to citrus. This whitefly entered California in the mid 1960s, where it quickly became a serious pest (DeBach & Rose, 1976). It was eventually controlled through the introduction

Table 2. Genus groups of Eudcromphalini, with distribution and host associations.

Distribution Host subkiinily Genus

Eiiderompliale group Bueoenredon S.E. Asia, Australiii Alcyrodinac Eirderomphale Cosmopolitan Alc yrodiniic Neopomplmle Ncotropics Alcyrodinac Potnpliule India. Pakistan Alcyrodiiiac

Alcuroclicinac Ncotropicb Akiirocroncis Alcurodicinac Eniedotintiecrerntiiis Ncotropics

Dasy ompliule Ncotropics Alcurodicinac

Eniedononecretnniib group

of a complex of parasitoids; howcver, N.u/eio-ofhrixi was not one of the species introduced. Another species of Neopornphale has been reared from a whitefly on coconut (see below).

Note on host records. We have included host records as we have received them (from label data or the literature), except when we feel that we can reasonably demonstrate

that thcse records arc wrong. Howevcr, certain records, particularly those from Aleyrodes sp., should be regarded as suspicious without further confirmation, because many workers seemed to regard any whitefly us belonging to Aleyrodrs. For example, the type series of Dasyomphalr chiletisis was recorded as ‘ex Aleurodes [sic] sp.’; however, examination of a host mummy associated with this series

Figs 28-36. Enredononecretnrzus sp. , P : 25, head; 29. pronotum; 30. mesosoma, lateral: 31, mesosoma, dorsal; 32, axilla, scutcllum. propodcum; 33, pstcr, dorsal: 34. gastcr. lateral; 35, gastcr, ventral; 36. apex of gaster, ventral. a, milla; c, cercus; hy, hypopygium; m. mcsoscutum; mp, mesoplcuron; op, outcr plate of ovipositor (P); p. pronotum: pp, prepectus: s, scutcllum; sp, subspiracular propodcal tubcrclc.

244 Johr~ LuSallc u d Micliiiel E. SchiirmrJ

leads us to believe that the host is an Aleurodicinac rathcr than Aleyrodes or any other Aleyrodinae (see below).

quadratc or loiigcr than widc (Figs 58. 5 9 ) . Dorsum or gaStcr cntircly metallic. the tirst tcrgitc morc >troiigly mct;illic 111381

following segments. without sctac (cxccpt a fcw latcral oncs). and usually smooth and polished in contrabt t o dktinctly wlpturcd rcmaining gastral tcrgitcs (Fig. 33). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Enredotrotler.rernrrllr Key to Genera of Euderomphalini

Scutellum with 1 or 2 pairs of sctac, and distinctly widcr than long (Figs 7, 15, 10. 20. 25). Midlobc of nicsoscutuin usunlly with oiily 1-2 pairs of sctiic (Figs 7, 15. I6.2U. 25). rarcly with 3-4 pairs. Submarginal win with 1 or 2 dorsal sctac (Figs 46-49). Mcsosoma somcwhat Rattcncd in latcral vicw, oftcn smooth or lightly sculpturcd (Figs 7, 15. 16. 20, 25). Vcrtcxal sulcus prcscnt, cithcr cxtcnding from thc cyc margin bctwccn thc occlli (Figs 3.4. 10-12) or bchind thc occlli (Figs 23, 24)

Scutcllum with many sctac scattcrcd over thc surfncc. thcsc not arranged in pairs, or (in onc spccics) with 3 to scvcral pairs: gcncrally not distinctly widcr than long (Figs 31, 38.43). Midlobc of mesoscutuin with many scattcrcd sctac (Figs 31, 38, 43). Submarginal vcin with morc than 2 dorsal sctac (Figs 50-53). Mcsosoma usually convcx (rarcly Rattcncd) in latcral vicw, and with distinct and rcgular raised rcticulation (Figs 31. 38) . Vcrtcxal sulcus abscnt. (Orfedonoriecrcmn

(Euderomphak gcnus group). . . . . . . .2

group). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Axilla largc, about as widc as long. and clcarly scparatcd from mcsoscutum by complctc suturcs (Fig. 7). Scutcllum with 2 pairs of sctac: a largc pair placcd mcdially and a smallcr pair (oftcn difticult to scc) placcd on thc postcrior margin (Fig. 8) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ermderomphule

Axilla fuscd to mcsoscutum (Figs 17. 21. 26) SO that thc suturc separating thcsc two sclcritcs is abscnt or indicatcd only an- tcriorly; if , rarcly. axilla is scparatcd by a complctc suturc, thcn it is distinctly longcr than widc. pair of sctac (Figs 15, 16.20.25). . . . . . . . . . . . . . . . . . .3

Dorsal surfacc of submarginal vein with a singlc scta (Figs 48, 4Y) (Notc: carc must bc rakcn not to includc a sccond strong scta which is placcd ncar thc basc of thc parastigma). Sculpture on mcsosoma cithcr: dccply inciscd and widc-mcshed (Fig. 25, Baeoenredon), or tincly raiscd rcticulatc (Fig. 20. Pomphale). Coloration oftcn mctallic or with mctallic tingc. Old World. .4

Dorsal surfacc of submarginal vcin with two sctac (Fig. 47). Sculpturc on mcsosoma usually smooth or dclicatcly inciscd (Figs 15-17). Coloration ycllow to brown or black, non- metallic. Ncw World. ...................... Neopomphale

Mcsosoma with sculpture on dorsal surface in thc form of fine raiscd rcticulations (Fig. 20). Scrobal sulcus absent or indistinct (Fig. 19). Vcrtcxal sulcus cxtcnding from cyc margin between thc mcdian and lateral occlli (Fig. 191. Prooodcum without

- Eycs with numcrous small sctac (although thcsc may not bc cabily sccn at low magnification) (Figs 37. 42). Oulcr phtcs of ovipositor normal, not widcned or Rattcncd: hypopygitim cxtcnding distinctly morc than 0.25 thc lcngth of the gilstcr. Fcmalc antcnna with F1 and F2 about equal in Icngth, both segments distinctly transvcrsc (Figs 60, 61). Dorsurn of gastcr not as abovc. eirher: first tcrgitc not contrasting in intcnsity of colour with remaining scgmcnts and with many small sctac (Figs. 45). or: gastcr with distinct non-mctallic porrions. .... .6

6 Gastcr uniformly dark i n colour: with many small sctac uniformly covcring all tcrgitcs cxccpt thc basal onc which is barc mcdially (Fig. 45). Forcwing (Fig. 53) dcnscly and uniformly covcrcd with drak sctac. spcculum small or abscnt: antcrior margin inciscd at apcx of costal ccll and marginal vcin; p s t - marginal vcin quitc long. as long as or longcr than stigma1 vcin.

- Gastcr with palc ycllow to wliitc markings at least basally. somctimcs morc cxtcnsivcly: not as sctosc as abovc with basal thrcc tcrgitcs barc mcdially (Fig. 11) . Forcwing (Fig. 5 2 ) not so dcnscly sctosc. and usually with light x t ac at least in proximal portion of wing. spcculum distinct: antcrior margin smooth, not incised: postmarginal vcin shorter than stigma1

Midtibial spur simplc. . . . . . . . . . . . . . . . . . . . . . . . Dasyomphule

vcin. Midtibial spur pcctiniitc (Fig. 40). . . . . . . . . Aleiirocfotius

Euderomphale genus group

lricluded geiiera. Baeoetrtedori, Euiieroniplzalr, Neo-

Distribution. Cosmopolitan. Hosts. Whiteflies in the subfamily Alcyrodinae. Diagtwsis. Head usually smooth to lightly sculptured

(Figs 1-4, 10- 12, 19, 23). Frontal sulcus, when present, placed just ventral t o median ocellus (Figs 10- 12). Ver- texal sulcus present, ei ther placed between the occlli (Figs 3, 4, 10-12) or behind the ocelli (Figs 13, 24). Malar sulcus usually present (Fig. 12), although sometimes incomplete (Fig. 2). Funicle with 1 or 2 segments (Figs 54-57). Dorsum of mesosoma more or less flattened, and usually smooth to lightly sculptured (Figs 7, 8. 15-17); generally when distinct sculpture is present it is incised (Fig. 25), although fine reticulate sculpture is present in

pomphale, Pomphult.

. - I

small pit with scnsilla. .................... :. .... Pomphale

- Mesosoma with sculpture on dorsal surfacc in thc form of wide-mcshcd. dcep incisions (Figs 25, 26). Scrobal sulcus

SU~CUS cxtcnding from eye margin behind the lateral ocelli whcre it joins the occipital edge (Figs 23,24). Propodeurn with small pit just medial to the spiracle which contains several sensilla (Figs 26,27). . . . . . . . . . . . . . . . . . . . . . . . . Bueoenfedon

Eyes bare (Fig. 28) . plates of ovipositor enlarged and flattcncd (Figs 31-36): hypopygium cxtcnding less than 0.25 the lcngth of thc gastcr (Fig. 35). Fcmalc antenna with distinctly longcr (at least twicc as long) than F1. and cithcr

one genus (Fig. 20). Midlobe of mesoscutum IlsualIY with only 1 or 2 pairs of setae (Figs 7, 15, 16,20,25). Scutellum distinctly wider than long, with 1 or 2 pairs of setae (Figs 7,

~ ~ X X o s c u t u ~ by a suture (Figs 77 8)* Or fused to meSOScutUm (Figs 15- 17, 21, 26). Submarginal vein with one or two setae on dorsal surface (Figs 46-49).

Monophyly. Two characters support the monophyly of this group. T h e fint is the presence of a transverse vertcxal Sulcus, which extends either between the median and lateral ocelli (Figs 3. 4. 10-12), or behind the ocelli (Figs 23, 24). A similar sulcus is present in another group of

present, y-shaped (can be partially %en i n Fig, 23). Vertexal 8, 15-16, 20, either completely separated from

Figs 37-45. (37-41): Aleurocfonus vitmriis P : 37. head; 38. mesosoma: 39, axilla. scutellum. propodeurn; 40, mid tibia1 spur; 41. gastcr. dorsal. (32-45): Dasyomplirrle chiletisis. 9 : 42. head: 43. mesosoma: 44. axilla. scutellum, propodeurn: 45. gaster. dorsal. a. axilla: fs. Frontal sulcus; mt. rnidtibial spur.

entedonine genera which parasitize thrips (Ceranisus, supporting the monophyly of this genus group: however, Enrt.tlanu.stichus, Coerlwrrrzn, Thripahiirs) (Schauff, 1991). its polarity is difficult to determine at the present time. Unlike this group of thrips parasitoicls, the frontal sulcus The second character which supports the monophyly of in the Eitderompliule genus group, when present or indi- this group is that the scutellum is distinctly wider than cated, is situated just below the median ocelli (Figs 1, long; this is quite rare in Entedoninae, and outside of the 10- 12. 19). This form of frontal sulcus is unknown in other Euderomphalini only seems to appear in i\ few unrelated Entedoninae. which might represent another character genera.

Euderomphale Girault (Figs 1-9, 46, 54, 62, 71, 72)

[ Cardiogasrer Motschulsky, 1863: 72. Type species Curdio- ~asterf irsr i~~entri .~ Motschulsky (monotypy). Misidentiti- cation (see comments below).]

Euderomphale Girault, 19 16: 410. Type species: Eictlerom- phale fu.scipenrii.s Cirault [ = E.Jlavimedia (Howard)] (original designation).

Aleurodiphagus Nowicki, 1929: 154- 155. Type species Pteropfrix [ = Eiideromphale] Jm.imedia Howard (subsequent designation by Peck, 1951: 432). [A.chelidonii Nowicki, designated as type species by Nowicki, 1929: 155, is a itomen niidum].

Diagriosis. Scutellum distinctly wider than long, and with two pairs of setae: a larger one near the middle of the scutellum and a smaller one at the posterior margin (Figs 7, 8). Midlobe of mesoscutum generally with two pairs of setae near the anterior margin (Fig. 7). Axilla large (as wide as long), and completely separated from mesoscutum by a suture (Fig. 7). Prepectus reduced and fused to the mcsopleuron (Fig. 6 ) . Dorsal surface of submarginal vcin with 2 setae (Fig. 46). Antenna with two funicular segments, although the first is reduced to not much larger than the size of an anellus (Fig. 54). Scrobal sulcus and frontal sulcus absent (Fig. l ) , although frontal sulcus occasionally very slightly indicated in a position just ventral to the median ocellus; vertexal sulcus extending from the eye margin between the median and lateral ocelli (Figs 3, 4). Mesosoma always black, non-metallic, more or less flattened in lateral view, sculpture on dorsal surface usually either finely incised or smooth. Malar sulcus present, although incomplete and extending away from the mouth margin (postero-laterally from the eye) (Fig. 2).

Monophyly und relationships. Euderomphale is best defined as monophyletic by two characters. The first is the presence of the large axilla, which is completely separated from the mesoscutum by a distinct suture (Fig. 7). This form of axilla is seen in no other Eulophidae. A second character is the shape of the malar sulcus, which is present, although incomplete and directed posteriorly away from the mouth margin (Fig. 2). Again, this i s a unique character which is not seen in other Eulophidae.

Euderomphale appears to be related to Neopomphale on the basis of the prepectus being reduced and fused to the mesopleuron (Fig. 6). This character is only seen in these two genera. It is also distinct from other members of the Euderomphale group because of the presence of a second pair of setae o n the scutellum, although the polarity of this character is equivocal and may not necessarily support the monophyly of the genus.

Distribution. Nearly cosmopolitan. although not known from Australia and SE Asia. BouEek (1988: 732) stated that this genus was from the Northern Hemisphere; however, we have seen specimens from Central America and Brazil, and a species from Madagascar has recently been transferred to Euderomphale (see E.phiIippiae below).

Hosts. Various whiteflies in the subfamily Aleyrodinae. Comments. This genus has been treated under the name

Curdiogaster (Burks, 1958: Conipere B Anncckc, 1Yhl; Peck, 1963), but Cardiogaster has since hecn shown to belong to the Pteromalidae, where it is a junior synonym (if Cephuleta (BouEek, 1965; Bou€ek et ul.. 1978).

De Santis (1979) listed four species in Eiideromphale, three of which do not belong in this genus: aleitrorhrixi (Dozier) (transferred to Neopomphale in this paper): f l . 1 ~

(Howard) (transferred to Omphale by LaSalle & Schauff, 1W2: 12): and metdlica (Ashmead) (transferred to Chryso- JlOlf~nlylU by Lasalle & Schauff, 1992: 8).

Thc vertexal sulcus in Euderomphale generally extends between the median and lateral ocelli, and the vertex posteriorly is without a sharp carina. In two undescribed species from the southeastern U.S.A. (CNC, Florida and Missouri) (one of which is brachypterous), t h e vertexal sulcus joins a sharp carina which separates the vertex from the occiput: however, the lateral ocelli are still posterior to the sulcus and are placed on the rear of the head (in Bneoeritedori, in which the vertexal sulcus also joins a posterior carina on the vertex. this carina is posterior to the lateral ocelli). These two undescribed species are unusual in anothcr character: the anterior margin of the axilla is not smoothly round as in most species, but sinuate and concave.

Irlclrtde~i .species

bemisiiie Viggiani. Euderomphule hemlviue Viggiani. 1977: 16- 18.

Hosts. Bemisia afer. Distribution. Europe. Notes. Recorded as a parasitoid of B.citricolu by Viggiani.

1977: however, this species was synonymized with B. 11uw cocki (Mound & Halsey, 197s). and B.haticocki was synonymized with B.afer by Bink-Moenen (lYS3).

callutiue Erdos. Eiideromphulr culliitiue Erdiis, 1966: 411-412.

Hosts. Unknown. Distribution. Europe . Refereirces. BouEek QL Askew, 1968: 139.

cerris Erd6s. Euderomphale cerris Erd6s. 1961: 482-453. Hosts. A leyrodes proletella. Asterobemisia carpirri. Distribution. Europe. References. Boufek & Askew, 1968: 139: Huldh,

20.

chelidoriii Erdos. Euderomphale chelidonii Erdijs. 412.

Hosts. Aleytodes proletella, A. lonicerae. Distribution. Europe.

986:

966:

References. BouEek Sr Askew, 1968: 139: Huldh , 19%: 20.

corfiriae Graham. Euderomphale cortinae Graham. 1986: 37.

Hosts. Unknown. Distribution. Madeira.

Figs 46 -53. Forcwing, ? : 46, Eirderomphule clielidonii: 47, Neopomphnle uletirothrixi: W, Pomplrule striptipentiis; 49, Baeocntedon sp.; 50-5 1, Etrtcdotrcitiecrernrrirs 5pp.: 52. Aleiirocronrrs virtutiis: 53, DusyornpAolc chilensis. Arrow points to whcrc anterior margin is incised at apex ot' costal ccl l and marginal w i n .

248 John LuSalle urid Michael E. Schuriff

Figs 54-61. Antenna, E : 54. Euderomphale chelidonii: 5 5 . Neopomplinle aleuroihrixi: 56. Pomplrale stripripetinis: 57. Bururtriedot~ sp.: 58-59. Eniedononecremnrcs spp.: 60. ALeurocionrts viitarus; 61, Dasyomphule chilensis. f l . f2. funicular scgmcnts 1. 2.

flulirnediu (Howard). Gyrolusia flrcvimediu Howard, 1881: 369.

Synonym. Euderomphale firscipeiziri.s G irault, 19 16: 4 10. Hosts. A1eyrode.s aureocirzcta, A .pruinosa, A.spiraeoides,

Aleyrodes sp. Distribution. U.S.A., Caribbean. Also listed from

Europe (BouEek & Askew, 1968: 139). Refererzces. Compere & Annecke, 1961: 65-63: Peck,

1963: 238-239 (both as Cardiogaster flavimedia): Burks, 1979: 1006: De Santis, 1979: 285.

hyalitia (Compere & Annecke). Cardiogaster hyaha Compere & Annecke, 1961: 68-69.

Hosts. Aleurocybotus occiduus. Distribution. U.S. A. References. Burks, 1979: 1006. Notes. Compere & Annecke (1961) record this species

from Aleyrodes sp. on milo (a variety of sorghum). We believe that the host species is Aleurocybotirs occiduus Russell. Poinar (1964) reported that E.hyalina (as Cardio- gaster) parasitized 10-15% of a whitefly that attacked numerous species of grasses, including milo and bermuda grass in the Coachella Valley of Southern California. He stated that it had been identified as an unidentified species of Aleurocybotus by L. Russell. About the same time, Russell (1964) described A.occiduus from a variety of desert locations (including Coachella Valley) in Southern California and Arizona, on a variety of grasses (including two species of Sorghum). This is the only species of whitefly known to attack grasses in the desert regions of Southern California (Gill, 1982).

lorzgipedicela Shafee et al. Euderomphale lortgipedicelus Shafee, Rizvi & Khan, 1988: 1-2.

Hosts. Unknown whitefly o n wild plant. Distributio~i. India.

philippine Risbec. Cascaphilippiae Risbec, 1957: 269-270. Hosts. Reported as reared from ‘cochenilles’ [scale

insects] on Phifippia [Ericaceae]. The rearing record needs verification.

Distribution. Madagascar. Notes. This species was described in the Aphelinidae.

and only recently transferred to Euderomphale by Polaszek (1993).

postmarginalis Shafee et al. Euderomphale postmarginalis Shafee, Rizvi & Khan, 1988: 2.

Hosts. Unknown whitefly on citrus. Distribution. India.

quercicola Dozier. Euderomphale quercicola Dozier, 1933: 85-86.

Hosts. Tetraleurodes sp. Distribution. U .S. A. References. Peck, 1963: 239 (as Cardiogaster quercicola);

Burks, 1979: 1006.

secrrtu Hulden. Euderonrphule wcretu HuldCn, IWh: 20 -2 1.

Hosts. A leu roc11 itorr aceris . Distribution . Europe.

secundu (Mani). Cardiogaster secundus Mani, 1939: 77. Hosts. Aleurolobirs buroderzsi.v. Distribution. India. References. Husain & Khan. 1986: 218: BouEek et a / . ,

1978: 460.

Neopomphale gen.n. (Figs 10-18,47,55,63-65,73,74)

Type species. Euderomphale aleurorhrixi Dozier. (Gender feminine).

Diagnosis. Scutellum distinctly wider than long, and with a single pair of setae (Figs 15, 16). Midlobe of mesoscutum generally with one pair of large setae near the anterior margin, and 1 or 2 smaller setae laterally on the scapula (Figs 15, 16). Axilla distinctly longer than wide, not clearly separated from the mesoscutum by a suture, although a suture may be partially indicated anteriorly (Fig. 17). Prepectus reduced and fused to the mesopleuron (Fig. 14). Dorsal surface of submarginal vein with 2 setae (Fig. 47). Antenna with 1 or 2 minute anelli, and a single large funicular segment (Fig. 55). Scrobal sulcus present, y-shaped (Figs 10- 12); frontal sulcus generally present just ventral to the median ocellus (Figs 10, l l) , occasionally only faintly indicated or absent (Fig. 12). Vertexal sulcus extending between the median and lateral ocelli (Figs 10-12). Malar sulcus present, complete (Fig. 12). Mesosoma from yellow to black, non-metallic, more or less flattened in lateral view, sculpture on dorsal surface usually either finely incised or smooth (Figs 15- 17).

Monophyly and relationships. At present we can find no single character to support the monophyly of Neopomphale, thus the group may eventually prove to be paraphyletic. It possesses a reduced prepectus which is fused to the mesopleuron. This is a derived character which it shares with Euderompkafe, and appears to support the rnonophyly of these two genera taken together. However, it is further characterized only by the absence of derived characters found in Euderomphale (shape of axilla, shape of malar sulcus). Neopomphale possesses a distinct scrobal sulcus, which is absent in Euderomphale, but the polarity of this sulcus is unknown, and it is also found in Baeoentedon. The further reduction of the antenna (two funicular segments in Euderomphale, although the first reduced; a single funicular segment in Neopomphale) might indicate a derived character, but since this form of antenna is also seen in Pomphule and Baeoentedon it remains ambiguous.

One character which might support the monophyly of this genus is the presence of a membranous area just following (or largely replacing) the basal gastral tergite (Fig. 18). This is generally present and distinct in Neo- pomphale species, although sometimes it is not very clear. Additionally, a small membranous area appears to be

250 Joliii LaSalle atid Michael E. .Tchuit ff

Figs 62-70. Clypcus. P : 62. Euderompliale sp.: 63-65, Neopornplrale spp.: 66, Pornp/iale sfriptiperritis: 67-68. Erifetlo~toiiecrcr~irius sp.: 69, Aletiroctortus viffufus; 10, Dasyornphale chilensis.

present in a few species in related genera (some species of Baeoentedon, and perhaps in a few Euderomphale species, although this character is often hard to interpret due to distortion in dried specimens).

Female. Colour yellow to brown or black, always non- metallic.

Head (Figs 10-12) smooth or with delicate incised sculpture. Scrobal sulcus present, y-shaped; frontal sulcus generally present just ventral to the median ocellus: occasionally only faintly indicated or absent. Vertexal sulcus extending from the eye margin between the median and lateral ocelli. Malar sulcus present.

Antenna (Fig. 55) with 1 or 2 minute anelli, and a single funicular segment, which is distinctly longer than wide. Club generally elongate, and with a long terminal spine.

Mesosoma (Figs 13-17) more or less flattened in lateral view, sculpture on dorsal surface usually either finely

incised or smooth. Scutellum distinctly wider than long, and with a single pair of setae. Midlobe of mesoscutum generally with one pair of large setae near the anterior margin, and 1 or 2 smaller setae laterally on the scapula. Axilla distinctly longer than wide, not clearly separated from the mesoscutum by a suture, although a suture may be partially indicated anteriorly: or, rarely, suture may appear to be complete, although faint. Prepectus reduced and fused to mesopleuron.

Forewing (Fig. 47) with dorsal surface of submarginal vein with 2 setae. Stigma1 vein short, straight or slightly curved. Speculum present and distinct. Disc often with infuscated areas, usually in the form of a partial or complete darker band placed near the middle of the marginal vein.

Caster (Fig. 18) short, ovate. without metallic coloration or distinct sculpture. Caster usually with a visible mem- braneous area between the first and second tergite, some-

times wi th thu dorsal surface of thc first tcrgitc largely replaced by a rncmbranous structure.

Mole. Similar to female except in genitalia. Antenna1 segments generally with distinct linear sensilla.

Distrihufiorr. New World tropics and subtropics. From the U.S.A. (Arizona, California, Florida) to Chile.

Hosts. Subfamily Aleyrodinae. Most specimens have been collected without host data, although hosts are known for the two described species. The record of Aleyroiles sp. for N.australi.s may be questionable.

Several specimens of a Neopomphale species (on a slide in the USNM) are recorded from whitefly nymphs on coconut, Piarco, Trinidae. They were determined by Gahan as 'Eiitleromphale australis Brkthes'; however, we are not convinced that Gahan properly understood the identity of this species. particularly as there is another specimen from the West lndies reared from citrus, also determined by Gahan to be N.au.str+ali.s, and which appears to be N.aleicrothrixi.

There is a series of an undescribed species from Sam Diego Co. , California, which was reared from a Trrraleicrodes sp. (TAMU ) .

C~~mt~icw/.s. Therc are presently only two spccics included in this genus; however. we are aware of several more species (at least six) from the New World tropics and subtropics (BMNH, USNM, CNC).

Species can range from almost entirely yellow in colour to black. One undescribed species (BMNH. Costa Rica) has a very long wing fringe (over a third the maximum width of wing).

iticluded species

aieurothriui (Dozier), c0mb.n. Euderomphale aleurothrixi Dozier, 1932: 120.

Hosts. Aleitrothrixics ~ o c c o s ~ i s ( = A . howardi). Distribicriorz. Caribbean, Argentina, U.S.A.: Florida. References. De Santis 1979: 285; 1989: 52.

australis (Brtthes), c0mb.n. Pteropfrix australis Brhthes, 1 Y 16: 24-25.

Hosts. A leyrodes spa Distribiction. Chile. References. De Santis 1979: 339.

Figs 71 -77. Antenna, d: 71-72, Euderornphale sp.; 73-74, Neopomphale sp.; 75-76, Eniedononecremnus sp.; 77, Aleurocronus vittaratus.

252 Johrz LaSalle arrd Michael E. Schautff

Notes. We have not examined types for this species; however, from the original description, with accompanying figure of the antenna, it is clear that this species belongs in Neopomphale. We have also seen material in the USNM which is determined as this species, and which belongs to Neopomphale.

Pomphale Husain, Rauf 81 Kudeshia (Figs 19-22, 48, 56, 66)

Hosts. Unknown. Distriburion. India.

striptipemis Husain et 01. Pomphuh .stripfipipeiinir Husain, Rauf & Kudeshia, 1983: 112.

Hosfs. Aleurolobus sp. Distribufion. India, Pakistan. References. Husain 8r Khan, 1986: 231. New record. Pakistan, Multan, 23.i.1981, ex. Aleurolobus

sp. on Zizyphus jujribu (Rhamnaceae) (29 I d . BMNH).

Pomphale Husain et al . , 1983: 112. Type species: Pomphale srriptipennis Husain et a / . (original designation).

Baeoentedon Girault (Figs 23-27, 49, 57) Diagnosis. Scutellum distinctly wider than long, and

with a single pair of setae (Fig. 20). Midlobe of mesoscutum generally with a single pair of setae (Fig. 20). Axilla distinctly longer than wide, not clearly separated from the mesoscutum by a suture, although a suture may be partially indicated anteriorly (Fig. 21). Dorsal surface of submarginal vein with a single seta (and base of parastigma with a strong seta) (Fig. 48). Antenna with a single transverse funicular segment (Figs 22, 56). Scrobal sulcus absent. frontal sulcus absent, although faintly indicated just ventral to the median ocellus (Fig. 19). Vertexal sulcus extending from eye margin between the median and lateral ocelli (Fig. 19). Eyes large; malar area relatively small and malar sulcus absent (Fig. 19). Mesosoma distinctly metallic (purple), more or less flattened in lateral view, sculpture on dorsal surface finely reticulate (Figs 20, 21). Colour distinctly metallic.

Monophyly and relationships. The monophyly of Pom- phale is best supported by a single character: the fine, reticulate sculpture on the dorsum of the mesosoma (not seen in other members of this genus group). This character is not extremely convincing; however, Pomphale does appear as quite distinct from other members of the group. Pomphale shares two derived characters with Bneoentedon: t he presence of only a single seta on the submarginal vein, and distinct metallic coloration (species of Bneoentedon have a slight metallic shine). Pomphule also differs from other members of the Euderomphale group in having relatively large eyes, so that the malar space is quite small, and the malar sulcus is absent (malar sulcus present in at least some form in the other genera). The polarity of these character states is questionable, however, as they resemble that seen in certain members of the Entedononecremnus genus group.

Distribution. Known only from India and Pakistan. Hosts. Reared from Aleurolobus sp. (Aleyrodinae). Comments. Mentioned by BouEek (1988: 732) as close

to Baeoentedon Girault, and as probably a parasite of Psylloidea. Since then a series of P.striptipennis has been reared from Aleurolobus sp.

Included species

setosipennis Hayat & Zeya. Pomphale serosipennis Hayat &k Zeya, 1992: 184-186.

Baeoeritedon Girault, 1915a: IYO. Type species Baeoeirtedoiz peculiconris Girault 1915 (original designation)

Diagnosis. Scutellum distinctly wider t h a n long, and with a single pair of setae (Fig. 25): midlobe of mesoscutuni with one or two pairs of setae (Fig. 25). Axilla distinctly longer than wide (Fig. 26). Dorsal surface of submarginal vein with a single seta (and base of parastigma with a strong seta) (Fig. 49). Antenna with a single funicular segment (Fig. 57). Scrobal sulcus y-shaped (can be partially seen in Fig. 23). Propodeum with a small pit just medial to spiracle which contains several sensilla (Figs 26, 27). Mesosoma black, with or without metallic shine, more or less flattened in lateral view, sculpture on dorsal surface in the form of wide-meshed, deep incisions (Figs 25, 26). Vertexal sulcus extending from eye margin behind the lateral ocellus and marking the occipital edge (Figs 23,24). Frontal sulcus absent. Malar sulcus present (although sometimes fine).

Monophyly and relutioruhips. Baeoerrtedori is best defined as monophyletic on the basis of two characters. The first is thc position of the vertexal sulcus, which extends posterior to all the ocelli (Figs 23, 24). This form of sulcus is not seen in other members of the Euderornphale group. The second is the presence of a small pit on the propodeum just medial to spiracle which contains several sensilla (Figs 26, 27); this pit was mentioned by BouEek (1988) and Hayat & Zeya (1992). It also is the only member of this group to have such deep, incised sculpture on the dorsum of the mesosoma, although some members of Neopomphale also have distinct incised sculpture.

It shares with Pomphale the presence of only a single seta on the submarginal vein, which is presumably a derived character, but one which could easily be the subject of convergence. As in Pomphale it has the prepectus free, and not'fused to the mesopleuron (as seen in Euderomphale and Neopomphale), but as this is presumably the primitive condition of this character it is not informative.

Distribution. Northeastern Australia, Indonesia, India and Southern China.

Hosts. Unknown (presumably Aleyrodidae). Comments. There is only a single described species in

this genus from Queensland, Australia. We have seen other specimens representing at least two additional species

Systernaiics of the Eudvrornphalini 253

from India (Karnataka), Indonesia (Sulawesi) and China (Hainan Island) (BMNH).

Iricluded species

peciilicoriiis Girault. Baeoentedon peculicornis Girault, 1915a: 191.

Hosts. Unknown. Distribution. Australia (Queensland). Refereiice. Boufek, 1988: 732.

Enfedononecremnus genus group

Included genera. Aleuroctonus, Dasyornphalr, Entedo-

Distribution. New World tropics and subtropics. Hosts. Whiteflies in the subfamily Aleurodicinae Diagnosis. Head with distinct reticulate sculpture (Figs

28. 37, 42). Frontal sulcus, variable: either absent (Fig. 28) ; present, transverse, and distinctly separated from median ocellus (Fig. 37); or present, curved, and only

tzoiiecrernrius.

separated from the median ocellus by a distance about equal to the diameter of the ocellus (Fig. 42). Vertex without any sulci. Malar sulcus generally absent, although may be somewhat indicated by a change in the pattern of the sculpture. Funicie always with 2 segments (Figs 58-61). Dorsum of mesosoma generally convex (somewhat flattened in one genus), and with distinct and strong reticulate sculpture (Figs 31, 38, 43). Midlobe of mesoscutum with many scattered setae (Figs 31,38, 43). Scutellum generally not distinctly wider than long, and with many setae scattered over the surface, these not arranged in pairs (Figs 31, 38, 43), or (in one’species) with three to several pairs. Axilla always separated from mesoscutum by a complete suture, although this suture may be very fine (Figs 32, 39, 44). Submarginal vein with 3-8 dorsal setae (Figs 50-53).

Monophyly. The best character to support the rnonophyly of this group is the presence of numerous, unpaired setae on the scutellum and rnesoscutum (Figs 31, 38, 43). This character is rare throughout the Eulophidae, and unknown in most other Entedoninae, although it does occasionally occur in genera such as Parahorismenus and

Figs 78-82. (78779): Encyrtomphale parvicorpus, 0 : 78, thorax (redrawn from Boucek, 1988, Fig. 1315); 79, antenna. ( 8 0 - a ) : Horismenoides sulfureivenrris, 9 : 80. rhorax (redrawn from Boucek, 1988, Fig. 1318); 81, face (redrawn from Boucek, 1988, Fig. 1317); 82. forewing. a, axilla: d , dorsellum; fs, frontal sulcus; m, mesoscutum; s, scutellum.

254 Johti LaSulle u t ~ t l Michael E. Schairfl

Plrurorroppopsis (BouEek, 1988: Schauff. 1991). o r in a few aberrant species of Pediohiic.v (e.g. .sefidqerit.s Kerrich, rnicltiserirs BouCek).

Entedononecremnus Girault (Figs 28-36,50.5 I , 58.59, 67, 68, 75, 76)

Eiitedotioiiecremnus Girault. 1915b: 278. Type species: Eiitedoriotiecrrmiiiis ienicirs Girault, I915 (original designation).

Diagnosis. Scutellum with many setae scattered over the surface, these not arranged in pairs (Figs 31,32), or in one species with three to several pairs. Forewing with submarginal vein with more than 2 dorsal setae (usuiilly 3, but occasionally 4-5) (Figs 50. 51): speculum present and usually distinct. Mesosoma convex in lateral view, and with distinct and regular reticulation (Figs 31, 32). Scutellum generally not distinctly wider than long. Antenna with F2 either quadrate or longer than wide, and distinctly longer (at least twice as long as) than F l (Figs 58 , 59). Outer plates of ovipositor enlarged and Hattencd (Figs 34-36). Eyes bare ( F i g 28). Entire gastcr metallic, the first segment more strongly n ~ t i i l l i ~ than following scg- rnents, and usually smooth and polished in contrast to

Midtibial spur usuitlly short, simple (however slightly pectinate in one undescribcd species).

Motrophyly atid relnfiorlships. Several characters support t h e monophyly of E/iredutiotzecremrlrl.s: the female gustcr has the hypopygium very short (Fig. 35); the outcr plates of t h e ovipositor are enlarged and flattened (Figs 34-36): t h e first gastral tergite is usually smooth (sculptured i n one undescribed species), and distinctly more metallic than the remaining tergites, which have at least some sculpture (Fig. 33). Otherwise it might be considered primitive in regard to other members of this tribe due to the position of the axilla, which is only partially advanced forward of the scuto-scutellar suture (however, this character is somewhat variable, and some species have the axillae quite advanced).

Distribution. New World tropics, from the Caribbean and Mexico south to Brazil.

Hosts. Associated with a variety of Aleurodicinae. Hosts from undescribed species include Ceruleitrodicus altissimus and Aleurodicus cocois.

E. unicus, the only described species, was originally recorded by Girault (1915b) from Aleurochiton sp. (Aleyro- dinae); however, there is reason to think that this record is wrong. Girault's type material was from near Georgetown, Demerara, British Guiana, March 1915, G . E. Bodkin, and was presumably part of a shipment of material sent for identification to the USNM. De Santis (1979) lists E. unicus from Aleurochiton sp. and Eudiufeurodicrts bodkini, although it is not clear where he got the record for E.bodkini. Mound. & Halsey (1975) similarly list Ebodkini as a host for E.unicus. E.bbdkini was described by Quaintance & Baker (1915) from material collected

d ' . istinctly . sculptured remaining gastral tcrgites (Fig. 33).

in Berbice, Guyana, March 191s. by (I. E. Bodkin. I t is highly possible that E.e~nicus was actually from the material which was descrihcil by Quaintarice & Baker as E. hotlkitii, rather than the A/eierochimz recorded by Girault.

A series of an undescribed species of Etite[loiioii~crern,Ilc.v from Colombia has label data which states that i t is reared from the nymph of Orthezici in.signi,s, but wc suspect that this record ikincorrect. Ortheziid nymphs, like Aleurodici- nac nymphs. have long, waxy filaments. and these two groups may be superficially similar to each other. There are other specimens of this same unclescrihetl species from Costa Rica and Trinidad which have label data indicating that they had been reared from aleyrodids. Without further rearings, we are considering that the Orrheziu record is simply the case of ;I misidentilied host

Cornmetiis. There is some variation in wing venation within this genus, with some species (including iinicits)

having the stigma1 vein long, straight. and with thc uncus well-separated from the apex of the vein (Fig. SO): other spccics have the stigma1 vein short and curved (Fig. 51). One undoscribed species (from Ecuador. CNC) has a pectinate mid-tibia1 spur similar to that s w n in ~ l ~ ~ , i l r ~ ~ ~ f f ~ i i f i . \ . only t h e hairs arc shorter.

A single undcscribetl spccies (BMNI-1. C h t n Rica) has the first gastral tergite with reticulate sculpture similar to that found on the rcmaining tergitcs; however. the lirst tergite is still more mctallic anti shiny t h a n the rcmaining tergitcs. This species also has ;I niorc seLosc t'orcwing. wi th the basal cell setosc, t h e speculum quite small, and 3-5 setae on the dorsal surface of the suhmarginal vcin.

There is only a single described species i n this genus, however we have seen an additional seven or eight unclc- scribed species from the Ncotropical Region (BMNH. USNM. CNC).

Inclirtled species

ienicies Girault. E/itc~do/Eoiiecreniizif.s iitiicrrs CirauIt, 1s) 1%: 278.

Hosts. Eudiuleiirudicirs bodki/ii (see discussion of biology above).

Disiribiitioti. Guyana. Reference. De Santis, 1979: 264.

Aleuroctonus gen.n. (Figs 37-41, 52. 60, 69. 77)

Type species. Euderomphak VlffAtri Dozier (Gender masculine.)

Diagnosis. Scutellum (and mesoscutum) with many setae scattered over the surface, these not arranged i n pairs (Fig. 38). Submarginal vein with more than 2 (generally 3) dorsal setae (Fig. 52). Mesosoma convex in lateral view, and with distinct and regular reticulation (Fig. 38). Female antenna with F1 and F2 wider than long and subequal in length, although F1 narrower than F2 (Fig. 60). Eyes with numerous small setae (Fig. 37). Caster without distinct sculpture: with pale yellow to white markings at

least basally sometimes more extensively. Micltibial spur long, pectinate (Fig. 40).

Moriophyly mid rehiomhips. A single character supports the monophyly of Aleurocforii4.s: the mid-tibia1 spur is long and pectinate (Fig. 40). This type of mid-tibia1 spur is not common i n the Chalcidoidea, although it has been reported in the Signiphoridae (Woolley, 1988). Unfortu- nately, a single undescribed species of Etifedoriotiecremtzus (CNC, Ecuador) and a single undescribed species of Nropotnphale (TAMU, Mexico) also have a slightly pectinate mid-tibia1 spur, although we presume that these are convergences. Aleuroctoiius shares the following characters with Dasyomphale: axilla situated completely anterior to the scuto-scutellar suture; eyes distinctly setose; both funicular segments transverse. The polarity of these characters might be considered as equivocal, and not necessarily indicative of recent common ancestry. How- ever, the position of the axilla in Etifedo~iotiecremriirs (only partially advanced) appears to be the most primitive within the tribe, as it is more typical of the state that is found in other Entedoninae and Eulophinae. Thus, the axillar placement in Aleurociottirs and Dasyomphale might represent a synapomorphy.

Femule. Head and mesosomn black, sometimes with slight metallic shine, particularly on head. Gaster generally brown apically, light yellow to white basally, the extent and intensity of the coloration somewhat variable. Legs and antenna light yellow to white, may have some dark markings.

Head (Fig. 37) with strong reticulate sculpture. Frontal sulcus present, well removed from the median ocellus. Malar sulcus absent, although somewhat indicated by a change in sculpture. Eyes large. with many setae.

Antenna (Fig. 60) with one anellus and two funicular segments. F1 and F2 both distinctly wider than long and subequal in length, although F1 narrower than F2.

Mesosoma (Figs 38, 39) convex in lateral view, and with distinct and regular reticulation. Pronotum strongly narrowed medially, although sclerotized throughout. Axilla present, narrow, placed entirely anterior to scuto-scutellar suture, and separated from mesoscutum by a suture. Mesoscutum and scutellum with many scattered setae. Midtibial spur long, pectinate (Fig. 40).

Forewing (Fig. 52) with more than 2 (usually 3) setae on dorsal surface of submarginal vein. Stigma1 vein straight, slightly elongated. Speculum present and distinct, although small. Setae on basal part of wing often distinctly lighter in colour than setae in distal part.

Gaster (Fig. 41) short, ovate, without metallic coloration or distinct sculpture.

Mufe. Similar to female except: antenna with F1 thin, anelliform (although wider than anellus and with setae); F2 distinctly longer than F1, quadrate to longer than wide.

Distribution. Neotropical: Florida, the Caribbean, Costa Rica, Colombia and Brazil.

Ho.si.s. A.viiturii.s has been recorded from two species of the genus Alritrodicus (see below).

Comments. Presently only a single described species, A.vittatus, which was described from Puerto Rico. This

species has dispersed to Florida, where it has been creditcd with the fortuitous biological control of the introduced Aleurodicus dixpersits (Bennett s( Noyes, 1989).

We have seen specimens of Aleuroctorzus from Costa Rica, Colombia and Brazil (BMNH, USNM, CNC), which represent at least one undescribed species.

iticluded species

viftutus (Dozier), c0mb.n. Euderornphule vitfatu Dozier, 1993: 86-87.

Hosts. Aleurotiiciis ailtillerisis, A. dispersus. Distribution. Florida, Puerto Rico, Cayman Islands. New records. CAYMAN ISLANDS: Grand Cayman,

7 Mile Beach, 26.vi.1986, D. M. LaSalle (19, LAS): Grand Cayman, 6.xii.1986, F. D. Bennett, ex. Aleurodicus dispersus on coconut (36 , USNM); Cayman Brac, Brac Reef, 25.vi.1986, D. M. LaSalle (1 6 , LAS).

Refereuces. De Santis, 1979: 330 (as Encarsia virfatu); Bennett s( Noyes, 198Y: 371-432 (as Euderomphale viitaiu).

Dasyomphale gen.n. (Figs 42-45, 53, 61, 70)

Type species. Dusyomphnle chiletisis sp.n. (Gender feminine.)

Diugriosis. Scutellum (and mesoscutum) densely covered with short, decumbent setae (Fig. 43), short setae also present medially on propodeum (Fig. 44). Submarginal vein with several (6-8) dorsal setae (Fig. 53). Mesosoma somewhat flattened in lateral view, with distinct and regular (although fine) reticulation (Fig. 43). F1 and FI! wider than long, F2 slightly longer and distinctly wider than F1 (Fig. 61). Eyes with numerous small setae (Fig. 42). Gaster uniformly dark in colour; first gastral tergite smooth, with many small setae; remaining gastral tergites lightly sculptured and setose (Fig. 45). Forewing (Fig. 53) densely and uniformly covered with dark setae; basal cell densely setose, and speculum virtually obliterated; anterior margin incised at apex of costal cell and marginal vein; stigma1 vein straight, distinctly elongate, as long as or longer than the elongated postmarginal vein; submarginal vein with several (6-8) dorsal setae. Midtibial spur short, simple.

Monophyly and relationships. The monophyly of Dasy- omphde is supported by several characters: the anterior margin of the forewing is incised at the end of the costal cell (Fig. 53); the speculum is much reduced or absent; scutellum and mesoscutum (and gaster) are extremely densely hairy (Figs 43-45).

Dasyomphale also shares several characters with Aleur- ocionus (see above).

Female. Entire body dark, sometimes with slight metallic shine. particularly on head.

Head (Fig. 42) with fine reticulate sculpture. Frontal sulcus present, curved, and only separated from the median ocellus by a distance about equal to the diameter of the ocellus. Malar sulcus absent, although somewhat indicated by a change in sculpture. Eyes with numerous small setae.

Antenna (Fig. 61) with one anellus and two funicular segments. FI and F2 wider than long, F2 slightly longer and distinctly wider than F2.

Mesosoma (Figs 43, 44) somewhat flattened in lateral view, with distinct and regular (although fine) reticulation. Pronotum strongly narrowed medially, although sclerotized throughout. Axilla present, narrow, placed entirely anterior to scuto-scutcllar suture, and separated from mesoscutum by a suture. Mesoscutum and scutellum densely covered with short, decumbent setae. Propodeum with setae median to the spiracle. Mid-tibia1 spur short, simple.

Forewing (Fig. 53) densely and uniformly covered with dark setae; basal cell denselysetose, and speculum virtually obliterated. Dorsal surface of submarginal vein with several (6-8) setae. Anterior margin incised near junction of costal cell and marginal vein. Stigma1 vein straight, distinctly elongate, as long as or longer than the elongated postmarginal vein. Forewing lightly infuscated, with a darker area of infuscation associated with the stigmal vein.

Caster (Fig. 45) short, ovate, entirely dark, with a slight metallic shine. Tergites lightly sculptured, and with many scattered setae.

~Mulr. Unknown. Di.srrib~itiotr. Chile. Hosts. Unknown Aleurodicinae (see discussion under

D. chilensis). Cornrnetifs. This genus is known from a single species,

which is described below.

It1 clu drd species chiletisis sp.n.

Dasyomphale chilensis sp.n. (Figs 42-45, 53. 61, 70)

Fernufr. Length 1.0-1.35mm. Head and body black, with faint metallic shine; head slightly more metallic than mesosoma or gaster. Antenna brown. Coxae and femora dark brown to black: tibiae and tarsi yellow, except hind tibia which is brown basally and yellow apically. Forewing lightly and evenly infuscated over entire surface, with an area of darker infuscation near the stigmal vein. Veins brown; parastigma with a hyaline area.

Antenna (Fig. 61). Proportions as follows; scape, 23; pedicel, 11: anellus and funicle together, 7.5: club (including terminal spine), 16.

Forewing (Fig. 53). Proportions as follows: costal cell, 40; marginal vein, 42: stigmal vein, 19; postmarginal vein, 21.

Other characters as in the generic description. Male. Unknown. Hosts and biology. This species has been reared from an

unknown species of Aleurodicinae on Quiflaja saponaria (Rosaceae). Label data for the type series .states ‘ex Aleurodes [sic] on Quiflaya [sic] saponaria’, however, a single whitefly mummy was mounted on one of the cards. Although badly damaged, this mummy appears to be an aleurodicine (J. Martin, pers. comm.).

Material e-urrnirzrd. Holotype E: CHILE. Illapel. IYhX. ex undetermined Aleurodicinae on Quilluju .supotiuriu ( B M N H ) .

4? paratypes. Same data as holotype (3? BMNH, I9 U S N M ) .

Genera possibly related to the Euderomphalini

Two Australian genera are possibly related to the Euder- omphalini. They each display some, but not all, of the characters which we consider as defining the tribe.

We are not presently treating them as belonging to the Euderomphalini. Future studies, including the examination of more morphological character3 and preferably biological information, might necessitate their placement in the Euderomphalini, which would result in a modified definition of the tribe from that presented here.

These genera were treated by Boui-ek (1988) who mentioned in both cases the possibility of a relationship with Euderornphalr. and included both genera in a key to Australasian Eulophidae.

Encyrfornphale Girault, 191Sa: 210. Type spccies Encyrtorn- phalr parrwlicorpits Girnult, I9 15 (original dcsignation).

Eticyrfomphule (based o n the singlc species E.purvuli- corpus) is similar to the Euderomphalini in having the pronotum concealed in dorsal view and the axilla strongly advanced and clearly separated from the mesoscuturn by ;I

suture (Fig. 78). However, it differs from Euderomphalini in having the dorsellum projecting past the apex of the scutellum and visible in dorsal view, and in the shape of the antenna (Fig. 79) which has a single anellus. four transverse to quadrate funicular segments. and an unseg- mented club which is very long (longer than pedicel and funicle together).

Additional characters are: scutellum with a longitudinal median line (not seen in any Euderomphalini); submarginal vein with a single seta: postmarginal vein distinctly developed.

This species is known only from two poor slide mounts (QMB, USNb1: both examined), which prevent proper assessment of all characters, particularly those on the head.

Horisrnenoides Girault. 1913; 153; 1915a: 1S1- 181. Type species Horisrnenoides sulfureiventris Girault, 1Y 13 (original designation).

Horisrnenoides (based on the single species Hmlfrtr- eiventris) is similar to the Euderomphalini in having the pronotum and dorsellum concealed in dorsal view: and the axilla strongly advanced and clearly separated from the mesoscutum by a suture (Fig. 80). The main differences are in the unusual shape of the frontal sulcus (Fig. 81), which is curved and placed about halfway between the median ocellus and the torulus: the venation in the fore wing (Fig. 82), which has the marginal vein distinctly (over

S,v.sirrtiudcv of thr Euclrrornphuliru' 257

3 times) longer than the submarginal vein; and the clypeal margin which is straight (Fig. 81).

Additional characters are: mesoscutum with a longitudinal median line (not seen in any Euderomphalini), and short, deep notauli present a t anterior margin: entire dorsal surface of the head and mesosoma is quite smooth and shiny; head without vertexal sulcus, without malar sulcus, and with 3 strong occipital carina; propodeal spira- cles placed on distinct tubercles; head and mesosoma dark, with slight metallic shine, gaster yellow; antenna with two funicular and three club segments.

This species is known only from a poor slide, one complete point mounted specimen, and three incomplete point mounted specimens (all in QMB: examined).

Acknowledgments

Several people have helped us in various ways in the preparation of the manuscript. For critical comments on the manuscript: J . Heraty, L. A. Mound, S. Nakahara, D. Nickle, A. Wijesekara. For access to information from his world catalogue of Chalcidoidea: John Noyes. For the loan of specimens: Z. BouEek (IIE), G. Evans (UFG), G. Gordh (UCRKJQ), M. Hayat (AMU), J. Huber (CNC), K. Lambkin (QMB), J. S. Noyes (BMNH), D. B. Wahl (AEI) , J. B. Woolley (TAMU). For information concerning hosts: R. J. Gill, J. Martin, L. A. Mound. For making specimens which they collected available to us: Tom Bellows, my brother Mark LaSalle, Lubomir Masner. Linda Millar prepared the line drawings in Figs 46-61. Ian Boler and Georgina Godwin were essential to the production of the photomicrographs.

This work was partially funded through NSF grant no. BSR-9020206. Additional funding for the senior author to travel and study in their institutions came from a Smithsonian Institution Short Term Visitor Fellowship (USNM) and a CanaColl Grant (CNC).

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Accepted 19 May 1994

Jo11 r n d of' E(r~,lornic. En 1fN?1 OiOgJ. 57, 37') - 3x3.

OJ / I i r I:',fior,toli,Sic.cil So<.iL'fy i/' k V l t . \ / / i I t ~ i i m , 66. 10 I - 1112.

Systematics of the tribe Euderomphalini (Hymenoptera: Eulophidae): parasitoids of whiteflies...

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Transcript of Systematics of the tribe Euderomphalini (Hymenoptera: Eulophidae): parasitoids of whiteflies...