Bull. not. Hist. Mus. Lond. (Entomol.)64(2): 117-163 Issued30November 1995 Encyrtidae of Costa Rica (Hymenoptera: Chalcidoidea): the genus Aenasius Walker, parasitoids of mealybugs (Homoptera: Pseudococcidae) JOHN NOYES S. DepartmentofEntomology, The NaturalHistory Museum, Cromwell Road, London SW75BD H.REN Guangdong EntomologicalInstitute, Guangdong, Guangzhou, P.R. China CONTENTS Synopsis 117 Intreduction 117 Depositories 118 Acknowledgements 119 AenasiusWalker 119 Systematicnote 119 Diagnosticcharacters 119 Biology 119 Use in biocontrol 120 Distribution 120 Identificationofspecies 120 Abbreviations used in text 121 KeytoCosta RicanspeciesofAenasius 121 Reviewofspecies 123 References 144 Illustrations 146 Index toscientific names 163 Synopsis. ThespeciesofAenasiusfromCosta Ricaare revised. Ofthese, 16are named andafurther20recognised frommales, but notnamed. Sixspeciesaredescribedasnew and 9 new specific synonymies are proposed. A dichotomous key to all species is providedandeach speciesisfurthercharacterisedbyadiagnosisand notesare provided on theirdistribution and hosts. INTRODUCTION onomists in comparison with those of tem- perate areas such as Europe and North America. Early European taxonomists concentrated their Until recently, tropical parasitic Hymenoptera attention on their home ground because that is have received relatively little attention from tax- where it was easiest to obtain material on which ©TheNatural History Museum, 1995 118 J.S. NOYES ANDH. REN to work. The early work of North American including the possibility that many tropical taxonomists largely related to agriculture and the insects may be unsuitable to act as hosts for control of insect pests. Thus, it was not until ichneumonids because of an increase in toxins relatively recently, with the use of modern col- accruedbythemfromthe plantsupon which they lecting and preservation techniques, that the feed ('poison host hypothesis'; Gauld et al., focus of these studies changed significantly. In 1992). This work has been supported by the particular, the use of Malaise traps greatly extensive use of hundreds of Malaise traps to enhancedcollection ofparasiticHymenopteraon survey Ichneumonidae in Costa Rica since 1985. a scale never before achieved. For instance, the As a side-product of this survey, a vast quantity work of Henry Townes revolutionised the study of microhymenoptera has become available for of Ichneumonidae by making available huge study. numbers ofspecimens collected from all overthe This unique collection has now made it pos- worldusinghis lightweight versionofthe Malaise sible to survey in detail the microhymenopteran trap (Townes, 1972). He also perfected a rapid fauna ofa tropical country for the first time. The technique formountingspecimenson the sidesof resultsofthe surveycan be used forcomparisons pins. Althoughthe use ofa Malaisetrapwasseen of species richness with similarly well-known as a valuable method for collecting microhy- temperate areas such as Great Britain. The col- menoptera, there was still one serious problem lection can also be used for comparative studies with using material collected by this method for on distribution within Costa Rica. Perhaps most taxonomic purposes. Small, weakly sclerotised important, by providing a sound framework for specimens, such as Chalcidoidea, shrivelled future work on this poorly studied group, taxo- badly when air-dried from alcohol. This made nomicstudiescan ultimatelyprovide the basisfor taxonomic work very difficult. The problem was future work on this group over a much wider overcome when it was realised that the use of area, perhaps the whole ofSouth America. critical-point drying could prevent specimens The present paper results directly from the from collapsing (Gordh & Hall, 1976). This was above mentioned Malaise trap survey and from anextremelyimportantdevelopmentinthe study the collecting efforts of many other entomolo- ofweakly sclerotised microhymenoptera because gists too numerous to mention here. It is it allowed specialists to remove specimens from intended as the first contribution in a series of alcohol and dry them without the danger oftheir studiesthat will cover the taxonomy, distribution integument collapsing. This revolutionised the and known hosts of the 700 or so species of one study of smaller Chalcidoidea in tropical areas family of Chalcidoidea, the Encyrtidae, that are because it has allowed specialists to study a large known to occur in Costa Rica. It is hoped that amount of good-quality material from a wide these studies will encourage more detailed stud- area. ies of this family in Costa Rica and elsewhere in The useofMalaise trapsforcollectingparasitic order to learn more about their biology, in Hymenoptera has led to some surprising find- particular their host ranges. Such work may ings. Owen & Owen (1974) observed that Ich- facilitate the use of particular species in biologi- neumonidae were no more diverse in tropical cal control programmes in other parts of the Africa than in temperate areas such as USA world should that become necessary. (Michigan) and Sweden. Thiswascontrary to the This first study is of the genus Aenasius, the generally accepted view that species richness, or species of which are relatively easy to recognise diversity, increases with a decrease in latitude. (see Fig. 1). The genusincludesseveral speciesof Many plausible theories were put forward as to actual or potential economic importance since why at least some parasitic Hymenoptera in their hosts are mealybugs (Homoptera: Pseudo- generalfollowthisreverse trend. Yetfindingsfor coccidae), many of which are important pests of northern Sulawesi (Noyes, 1989) indicated agriculture in most parts ofthe world. strongly that whatever might be true for Ichneu- monidae did not seem to be the case for other groups of parasitic Hymenoptera, particularly DEPOSITORIES the Chalcidoidea. The view that there is not an increase in species richness in Ichneumonidae in lower latitudes now seems to be well supported AMNH American Museum ofNatural History. New by some extensive studies in Costa Rica and York, USA eisewhere (Gauld, 1986; Gauld et al, 1992). BMNH The Natural History Museum, London, UK These studies suggest that species richness in this CNC Canadian National Collection. Ottawa, group may be influenced by a number offactors Canada ENCYRTIDAE OFCOSTA RICA 119 IEE Instituto di Entomologfa Espariol, Madrid, can also be distinguised from all other genera of Spain Encyrtidae using the characters given in the INBio Instituto National de Biodiversidad, Costa followingdiagnosis. Rica MBA Museo Argentino de Ciencias Naturales. Diagnostic characters BuenosAires, Argentina PPRI Plant Protection Research Institute, Preto- Body generally squat and robust; mandibles TAMU Trieax,aSsouAth&AMfricUaniversity, College Station, bidentate with lower tooth short (Fig. 101), or USNM TUenxiatse,dUSStaAtes National Museum, Washing- wFiEthMAaLEs:horftr,onutpopveerr,texth(iFridgsto8o6t-h91)(Fwigi.th1c0o0n).- ton D.C., USA spicuous piliferous punctures giving it the ZISP Zoological Institute, Academy of Sciences, appearance of the surface of a thimble or golf St Petersburg, Russia ball; scrobes deep and frequently sharply mar- gined dorsally and laterally; scape varying from almost cylindrical (Fig. 18) tostronglybroadened Acknowledgements. We thank The Royal Society, and flattened and less than twice as longasbroad via the K.C. Wong fellowship, for allowing the junior (Figs 3, 6); funicle 7-segmented (Figs 2-19), the authortheopportunitytostudyatThe Natural History first segment verysmall and normallyvisible only Museum in London; his host institution, Guangdong at higher magnifications on slide-mounted mate- Entomological Institute, permitted him one year'ssab- rial; clava withsensory area enlarged, forming an baticalleave. Wealsothank DougWilliamsforhishelp oblique apical truncation which is longitudinally in checking mealybug names. We especially thank Sue divided by a straight to sinuate line (Figs 9, 11, Lewis for preparing the electron micrographs (Figs 92, 93); forewings infuscate (Figs 20-45); apexof 86-97). Also thanks to Dr M.E. Schauff (USNM), Dr postmarginal and stigmal veins frequently con- L. Masner (CNC) Dr J. Huber (CNC), Sa Izabel nected by a naked, hyaline streak (Figs 32-45); Izquierdo(IEE), DrA. Bachman (MBA) and Dr E.L. gasterwith hypopygium reachingapex; paraterg- Quinter (AMNH) for the loan orgift ofmaterial. Our ites sclerotised (Figs 46-49); last tergite more or special thanks to Pam Mitchell and Paul Hanson, for less U-shaped; third valvulae usually free (Figs without their painstaking sorting of Malaise trap 50, 52, 55). MALE: frontovertex either with catches this project would not have been possible. piliferous punctures (Figs 96, 97) or with more Finally we would like to thank Aldo de Oyarzabal for irregular, sometimes coarse, sculpture (Figs 94, the colour illustrationofAenasimfrontalis (Fig. 1). 95); antennae either with 6 distinct funicle seg- ments and a small, entire clava (Figs 69-75), or Aenasius Walker with 2-5 anelliform segments and a relatively long, clava (Figs 76-83); wings hyaline (Figs Aenasius Walker, 1846: 180. Type species Aena- 62-68), rarely distinctly infuscate (Fig 65); post- sius hyettus Walker, by monotypy. marginal vein conspicuously longer than stigmal; Neodiscodes Compere, 1931: 212-21A. Type spe- phallobase (Figs84, 85, 99) with a pairofdistinct ciesNeodiscodesmartiniiCompere, byoriginal digiti, each with two or three apical hooks, designation and monotypy. Synonymised with outside each digitus a pair of bristles; aedeagus Aenasius by Prinsloo, 1988: 1468. broad (Fig. 99) and about as long as mid tibial Pseudanasius Hayat, Alam & Agarwal, 1975: spur. 21-23. Type species Pseudanasius clavus Hayat, Alam & Agarwal, by original designa- Biology tion and monotypy. Synonymised with Aena- sius by Hayat, 1981: 17. Where their hosts are known, all species are solitary endoparasitoids of mealybugs Systematic note (Homoptera: Pseudococcidae). Little is known of their development but the egg and newly Within the Encyrtidae, Aenasius belongs to the hatched larva have been described for Aenasius subfamily Tetracneminae, tribe Aenasiini. The maplei Compere from North America (Maple, tribal classification of the Tetracneminae has 1947). In this species the deposited egg is typi- been reviewed by Noyes & Hayat (1994) with six cally encyrtiform with the main part of the egg tribes being recognised and diagnoses provided being attached to a smaller bulb by a narrow for the subfamily and each of the six included stalk. The deposited egg remains attached to the tribes. The genus has been diagnosed in a key to integument of the mealybug host via the stalk Neotropical encyrtid genera (Noyes, 1980), but which, together with the collapsed bulb, projects 120 J.S. NOYES AND H. REN to the exteriorofthe host. The young larva hasa Identification ofspecies single pair of functioning spiracles and remains attached to the remains of the egg shell and Several keys have been published to the species: utilises atmospheric air via the protruding egg Compere (1937) for New World species, Kerrich stalk and bulb. (1967) for all known species (Old World species as Neodiscodes), Kaul & Agarwal (1985) for Old World species (as Neodiscodes) and Prinsloo Use in biocontrol (1988) for Afrotropical species. Some of these keys are based on characters Although species of Aenasius must play an which may be unreliable, e.g. coloration of important role in the natural regulation ofpopu- antennal segments, or very small differences in lations of their mealybug hosts, there have been the relative width of the scape or frontovertex. few attempts to utilise the species in biocontrol During our study we found that some characters programmes (see Table 1). appear to be reliable in defining species or groups ofspecies. In the female these are: a) the Distribution presence or absence of a hyaline streak at the apex of the venation, b) the relative width and Of the 38 described species of Aenasius, 29 are shape of the scape, c) the size and depth of the knownfrom the NewWorld, eight are Afrotropi- antennalscrobes, d) the appearance anddistribu- cal and Oriental. The single remaining species, tion of the piliferous punctures, e) the shape of advena, is circumtropical, but of Neotropical the lower distal margin of the forewing, and f) origin. The Old World species form a group the shape of the posterior margin of the last which is distinguished by the frontovertex being tergite ofthe gaster. In the male, four characters notmore than one-sixth the head width, whilst in appear to be useful: a) the number and relative the New World species the frontovertex is con- sizeofthe funiclesegments, b) the relativesizeof spicuously wider, normally at least one-quarter the clava, c) the sculpture of the frontovertex, head width. We have examined many unde- and d) the presence or absence of particular scribed species from South America. sensilla on F6 or the clava (see Figs 102-105). Within Costa Rica, species ofAenasius gener- The male genitalia of Aenasius do not seem to ally tend to be most common in drier habitats at show a great deal ofvariation, although we have or below 500 m altitude, eg. Guanacaste, San noted some variation in the relative size of the Jose Province, and west of the central range. bristles at the bases of the digiti. However, we Malaise trap catches suggest that, although they have not examined enough slide mounted mate- are generally more common during the drier rialofanysingle speciestodetermine the amount parts of the year (January to March in Guana- ofintraspecific variation in this character. caste), they can also be common if there are Although the present work deals with 16 longish dry spells during the wetter parts (eg. described species, we have recognised a further September-October in Guanacaste). This either 20speciesfrom malescollected in Costa Rica. As reflects a real build-up in numbers due to the with most Encyrtidae, the taxonomy ofAenasius rapid population growth oftheirmealybug hosts, species is based entirely on female characters, or greater parasitoid activity during drier peri- and therefore these additional species are not ods, or perhaps a combination ofboth. being named in this paper, although some prob- Table 1 A summaryofthe useofAenasiusspp. in biological programmesworldwide (Abbreviations: E- established; NR-not released; ??-nosubsequent information; P-partialcontrol; R-released, but no further informationavailable; SC-successfulcontrol). Targetpestspecies Aenasiussp. Country Yearfirst used Result Source Phenacoccusherreni vexans Colombia 1994 SC J. Castillo (perscomm.) Dysmicoccusbrevipes cariocus Hawaii 1935 99 Swezeyelal. (1939) Ferrisia virgata advena Hawaii 1923. 1929. 1958 P Bartlett in Clausen (1978). Funasaki, etal. (1988) advena California 1966-1967 R DeBach& Warner (1969) Phenacoccusmanihoti phenacocci Africa 1978 NR? CIBC(1979, 1980) Pseudococcustnaritimus paulistus USA 9 E? Compere (1937) ENCYRTIDAE OFCOSTA RICA 121 ably represent undescribed species. In order to median incision (Figs46, 47), either tegulae yellow facilitate their future recognition we include or head and thorax with conspicuous silvery below short diagnoses ofthe males ofall unasso- setae 5 ciated species and include them in the key to - Apexoflast tergitewith a median incision (Fig. 48, species. also as in Fig. 49), tegulae dark brown, head and thorax without conspicuous, lamellate silvery Abbreviations used in text setae 6 CCLW MLeanxgitmhuomfcwoisdttalhcoefllcoosftaflorceelwlinogfforewing 5 dHaerakdbranodwnthorax clothed in silvery setae, tegduilvaees Fl,F2,etc First funicle segment, second funicle seg- - Head and thorax clothed in brown setae, tegulae ment, etc yellow vexans EL Maximumeye length EW Maximumeye width 6 Funicle completely dark brown without yellow seg- FV Minimum frontovertex width ments phenacocci FWL Forewinglength - Funiclewith several yellowsegments 7 FWW Forewingwidth GS Maximum length ofgonostylus (orthird val- 7 Scape mostly dark brown, piliferous punctures immediately above facial impression completely vula) HW smooth and veryshiny advena Headwidth HWL Hindwinglength - Scape completely yellow, piliferous punctures HWW Hindwingwidth immediatelyabove facial impression dull lua MS Malar space (the shortest distance from the eye to mouth margin) 8 Antennaalmostcompletelyyellow,onlythe pedicel MT Mid tibia length dark brown cirrha OL Ovipositorlength - Antenna with scape and clava variously marked OOL Ocular-ocellar line, or the shortest distance with brown 9 between eachposteriorocellusand the adja- 9 Large piliferous punctures of frontovertex not, or centeye margin POL Posterior ocellar line, or the shortest dis- hardly, extending between eyes and facial cavity; costal cell normally gradually taperingdistally (Fig. tance between the twoposteriorocelli 27) rarely abruptlyincised insularis SL Scape length (excludingradicle) SW Maximumscape width - Large piliferous punctures extending at least half way between top of facial cavity and malar sulcus Key to Costa Rican species ofAenasius (as in Figs ,N(>-90); costal cell subrectangular, its apex abruptly incised at the point where the sub- (Females and males) marginal vein and marginal veins meet (Figs 29. 31) 10 1 cCaltaivoant(hFriegse-2s-e5g,m8e-n1t9e,d9w8i)thneavneroblloinqgueanapdicsaalustarguen-- 10 Frontovertex more than one-third headwidth shaped). Females 2 kerrichi [Funicle 7-segmented, the first segment sometimes - Frontovertexless than one-third head width ... 11 verysmall, anelliform andeasilyoverlooked] 11 Marginal vein about as long as stigmal vein (Fig. - Clava entire, with apex rounded and frequently 29) pelops very long and sausage-shaped (Figs 69-83, - Marginal vein not more than two-thirds length of 102-105). Males 17 stigmalvein (Fig. 31) paulistus [Funicle 3- to6-segmented] 12 Scape notmorethan2 x aslongasbroad,orhardly 2 Forewingwithoutanaked, hyalinestreakatapexof so (Figs3, 4. 10) 13 venation (Figs20-31) 3 - Scape at least about 3 x as long as broad (Figs - Forewing with naked, hyaline streak present (Figs 16-19) 15 1,32-45) 12 13 Scape about twice as long as broad and widest in 3 Scapelessthan2.5 x aslongasbroad(Figs2,8, 12, middle with lower margin more or less evenly 98) 4 curved from apex to base (Fig. 18); funicle com- pletelyblackorwithsome yellowsegments - Scape more than 2.5 x as long as broad (Figs frontalis 3-15) 8 - Scape less than 1.9 x as long as broad and widest 4 Apex of last tergite broadly concave not with a beyond middle, with lower margin distally bulging 122 J.S. NOYES ANDH. REN beyond insertion of pedicel and thus not evenly - Forewing without a naked, hyaline streak at the curved to base (Figs 3, 4, 6); funicle never with apexofthevenation 21 yellowsegments, alwayscompletelyblack 14 21 Pedicel not largerthan Fl and usually shorter (Figs 14 Facial impression acutely margined dorsally and 73-75, 104) 22 relatively narrow and deep at this point, delimited by a concave or straight line (Fig. 88); dorsal - EitherpedicellongerthanFl (measureddorsallyor surface of costal cell with setae relatively sparse, medially, excluding any ventral process, ifpresent) arranged in twoorthree lines(Fig. 34) bolowi or pedicel clearly larger than Fl (Figs 71, 102, 105) 25 - Facialimpressionnotacutelymargineddorsallyand breylataivbeilycobnrcoaavdealnidnesh(aFlilgo.w8a9t);thidsoprosianlt,sudreflaicmeiteodf 22 Hdairnkdastarhsiinddatribkiaberoorwnneaorrlydsaork orange-browns,p.aCs costalcellwithsetae dense andarrangedin threeor - Hind tarsi yellow to yellow-brown, clearly paler fourlines (Fig. 38) caeruleus than hind tibiae 23 15 Lowerpart ofapical margin offorewingstraight or 23 Mesoscutum and scutellum with similar, fairly evenslightlyconvex (Fig. 40);sensoryareaofclava smooth and shallow imbricate-reticulate sculptu- not more than 2/3 as long as clava (both measured re sp. D along longest axis) (Fig. 16); apex of last tergite withouta median incision (as inFig. 47) - At least scutellum with rough, reticulate, or brasiliensis punctate-reticulatesculpture 24 - Lower part of apical margin of forewing slightly 24 Clava on external ventral surface near base, and emarginate (sometimes almost imperceptibly so) usuallyF6,with an oval patchofspecialisedsensilla (Figs42,44);sensoryareaofclavamore than2/3 as or setae which are clearly more dense than other long as clava (Fig. 19); apex of last tergite with a setaepresenton thesesegments (Fig. 104) .. sp. E distinct median incision (as in Fig. 49) 16 - NeitherclavanorF6withasimilarpatchofspecial- 16 Scape longer than maximum width of eye; dorsum isedsensillaorsetae sp. F of thorax with a moderately strong blue-green lus- 25 Costal cell or forewingwith at least three complete tre mitchellae linesofsetae dorsally (Fig. 64); clavawith twooval - Scape slightly shorter than maximum eye width; sensory areas basally on outerventral surface (Fig. dorsumofthoraxdull, blackish, with aweakgreen- 102) bolowi ishorpurple lustre longiscapus - Costalcellofforewingwithonlytwocomplete lines 17 Funicle composed of six distinct segments, none of setae dorsally, only occasionally with a few obscured by base of clava (Figs 69-75. scattered setaecomprisinga partial thirdline; clava 102-105) 18 withatmostonlyoneovalsensoryareabasally . 26 - Funicle with fewer than six segments, these fre- 26 Antenna with only a very short, inconspicuous quently hidden by base of clava, the latter some- perpendicular seta on ventral surface of F6, this times very long and sausage-shaped (Figs onlyabout as longasadjacent recumbentsetae and 76-83) 30 clearly less than one-third diameter of segment; forewings normally infuscate in basal half; larger 18 Funicle with both Fl and F2 relatively small, nei- species, generally >1 mm, smaller specimens ther with longitudinal sensilla and not wider than rare brasiliensis pedicel or hardly so; Fl not longer than pedicel (Figs70, 72) 19 - Antenna with a conspicuous perpendicular seta on ventral surface of F6 which is clearly much longer - Funicle with at least F2 clearly wider than pedicel than the adjacent recumbent setae and at least andusuallywith longitudinalsensilla; Fl frequently nearlyhalfaslongasdiameterofsegment(Figs 103, longerthan pedicel (Figs71, 73-75) 20 105); forewingsentirelyhyaline; smallerspecies <1 mm 27 19 Tegulaeandalaryscleritesdarkbrown,moreorless concolorouswith mesoscutum; forewingswithpost- 27 Frontovertex, below anterior ocellus, with regular, marginalvein slightlylongerthanstigmalvein distinctpiliferouspunctures (asin Figs96, 97) . 28 sp.A - Frontovertex, below anterior ocellus, without pilif- - Tegulaeandalaryscleritesorangebrown,conspicu- erous punctures or if present than they are very ously paler than mesoscutum; forewings with post- indistinct and obscured by irregular, rough sculp- marginalveinatleastabout 1.5 x aslongasstigmal ture (asinFigs94,95) 29 vein (Fig. 62) vexans 28 Antennawith ventral sensory areason both F6and 20 Forewing with a naked, hyaline streak at the apex clava; clavawith alineofpeg-like sensillaon inner ofthevenation sp. B surface extending from base to a little more than ENCYRTIDAE OF COSTA RICA 123 one-third toapex (Fig. 103) sp. G tovertex not more thanone-third headwidthand at most about as wide as distance of anterior ocellus - F6 and clava without any visible ventral sensory from facial impression sp. O areas; clava with one or two peg-like sensilla about halfway along ventral surface (Fig. 105) .... sp. H - Clava about 2.5 x as long as scape (Fig. 76); frontovertex slightly more than one-third head 29 Clava basally with a single oval, ventral sensory width and nearly 1.5 x as broad as distance from area (similarto those in Fig. 102) and with apairof anteriorocellus to facial impression paulistus basal peg-like sensilla on inner surface (similar to Fig. 105) sp. I 38 Forewingsstronglyinfuscate;clavawithaconspicu- ous, nakedventral ridgeextendingalongmostofits - Clava without an oval, ventral sensory area near length (Fig. 82) 39 base and without peg-like sensilla on inner sur- face sp.J - Forewings only weakly infuscate; clava without a naked, ventral area (Fig. 83) 40 30 Funicle with more than two segments, distal seg- ments always obscured by base of clava, these 39 Head about 2.5 x as wide as frontovertex; fron- sometimesonlyvisible on slide-mounted specimens tovertex about 1.5 x as wide as distance between (Figs77, 80, 81) 31 anteriorocellusand facial impression sp. Q - Funicle with two, clearly visible segments (if - Head about 2.25 x as wide as frontovertex; fron- obscured dorsally by base of clava, then clearly tovertex a little more than twice aswide asdistance visibleventrally) (Figs78, 79, 82, 83) 35 between anteriorocellus and facial impression sp.R 31 Frontovertex with distinct piliferous punctures which are smooth, or nearly so, and which give it 40 Antenna with Fl subquadrate, nearly as long as the appearance of the surface of a thimble or golf pedicel, clava about aslongasscape sp. S ball (Fig. 97) 32 - Antenna with Fl strongly transverse, clearly much - Frontovertex with piliferous punctures obscure, shorter than pedicel; clava about 2 x as long as sculpture irregular and rough in appearance, not scape (Fig. 83) sp. T like the surface of a thimble or golf ball (Figs 94, 95) 33 32 Facial impression shallow and clearly remote from REVIEW OF SPECIES eye margins; piliferous punctures extending to nearlylevel with topsofantennal toruli sp. K Aenasius dives sp. n. - Facial impression relatively steep and more or less touching eye margins; piliferous punctures not (Figs2,20,21,46,52,53,56) extending between facial impression and eyes (Fig. 97) phenacocciladvenal?longiscapus Diagnosis. Female: head, dorsum of thorax and sides of gaster basally clothed in very con- 33 Clavaat most about 1.5 x as longasscape .. sp. L spicuous whitish setae; antenna uniformly black; - Clava at least about 2 x as long as scape (Figs 80, scape about 1.5 x as long as broad; sensory area 81) 34 of clava obliquely divided; frontovertex about one-quarter head width; piliferous punctures 34 All tarsi dark brown and more or less concolorous below anterior ocellus very shiny; scrobes deep, with tibiae; funicle 5-segmented; clava about 4.5Mx delimited dorsally and laterally; forewings with aslongasbroad (Fig. 80) sp. apex hyaline but with no hyaline streak present - At least mid basitarsus yellow or testaceous-brown at apexofvenation; lowerpart ofoutermargin of and conspicuously paler than mid tibia; funicle forewing slightly convex; scutellum with a deep, 3-segmented; clava at least 7 x as long as broad longitudinal median groove in basal half; apex of N (Fig. 81) sp. last tergite of gaster evenly and shallowly con- 35 Forewinghyaline 36 cave. - Forewinginfuscate 38 Male. Unknown. 36 Clava about 3 x as long as scape and without a Female. Length: 1.42-1.65 mm (holotype 1.62 naked, ventral ridge (Fig. 79) sp. P mm). Frontovertex metallic green, slightly cop- pery in ocellar area, below anterior ocellus pilif- - Clavanotmorethan2.5 x aslongasscapeandwith erous punctures metallic green or blue-green, a naked, longitudinal, ventral ridge (Fig. 76, ridges between punctures purple, setae contrast- 78) 37 ing snow-white; temples and genae metallic 37 Clava about 2 x as long as scape (Fig. 78); fron- green or blue green; facial impression metallic 124 J.S. NOYES ANDH. REN green mixed coppery, interantennal prominence venation, lower part of apical margin slightly mixed with purple; antennae black-brown; dor- convex. Relative measurements (holotype): sum ofthorax dull dark blue, mixed with purple, FWL 81, FWW 36, CL 36, CW 3; HWL 60, HWW almost black and clothed in very conspicuous 20. pale brown, almost white setae; tegulae dark Gaster with last tergite apically slightly, but brown; propodeum medially blackish; mesopleu- evenly concave (Fig. 46); ovipositor (Fig. 52, 53, ron blue-green; sides of propodeum coppery 56) with outer apical part of second valvifer purple clothed in conspicuous pale setae; forew- strongly obliquely truncate and apically acute. ings (Fig. 20) more or less uniformly brown Relative measurements (paratype): OL37; GS9; basally, but past apex of venation becoming MT55. gradually paler so that apical one-eighth or so is Male. Unknown. hyaline; no hyaline streak joining apices of post- marginal and stigmal veins (Fig. 21); hindwing Variation. Very little ofnote in material avail- hyaline; all coxae dark brown, fore coxae dis- able. The frontovertex varies from slightly less tinctly metallic blue-green; femora and tibiae than to more than one-quarter head width and dark brown, but femora testaceous distally; fore the ovipositor varies from two-thirds to three- tarsi almost completely brown, mid and hind quarters as long as mid tibia. tarsi yellow-testaceous, pretarsi dark brown; mid Hosts. Unknown. tibial spurdark brown; gasterdark purple-brown with conspicuous pale setae laterally at base. Distribution. Costa Rica. Head below anterior ocellus with large, con- Materialexamined. spicuous, shiny piliferous punctures, each sepa- Type material. Holotype $: COSTA RICA, rated by a sharp ridge, punctures in ocellar area Guanacaste Prov., Santa Rosa NP, Hacienda- shallower less shiny, between antennal scrobes & samnadllaenrtedriiaomretoecreltlihanthaensteerpiuonrcotcuerlelsus;oftwsolilgihntelsy 3P-a0r,atyp2e9s.:xi-2C0.OxSiiT.A1986RIC(AJ,anze1n$, GuanGaacualsd)t.e Prov., Santa Rosa NP, Bosq. Hum.-ll-0, ofpuncturesextendingbetweenfacial impression 13.iv-4.v.l986 (Janzen & Gauld); 1$, Santa and eyes nearly to level of lowest eye margin; sslcohrnaoglbleoasws cddaeerelipin,ma;itehevdeenalddyoirncsauslrilvdyeedvainteodwtaaotpbosuoiftdestscwriobcbeyesa,as MR(JoeasnnagzeonNVPn&,CGaHacauacloid,e)nv;d.a12-9§38-,80(,GJuaan2n0z.eaxnc.a&1s9t8Ge6a-u1Nl0Pd.,i).;1E19s$8t,.7 sides of facial impression slightly bulging out- Guanacaste NP, near HQ, 2-10.iii.1990 (J.S. Noyes); 1$, San Jose, Ciudad Colon. Hda El wards and then abruptly angled inwards towards mapoeuxt;h;clsacvaapseli(gFhitgl.y2s)hosrltigehrtltyhabnulfguinnigcloeu,tiwtsarapdiscaalt RHooldoetoy,pe 1a6n.dii.p1a9r9a1typ(eHsyimn.BMPNarHa,taxpoanroamtiysptes)i.n INBio. sensory part about three times as longasventral, straight margin of clava and divided by an Comments. Aenasiusdives isdistinctive and can oblique conspicuouslysinuate line (asin Fig. 93); be separated from all other known species ofthe ocelli forming a distinctly acute. Relative mea- genus by the very conspicuous whitish setae on surements (holotype): HW43.5; FV 10.5;EL31; the head, dorsum of thorax and sides of propo- EW21; MS9; SL24.5; SW 16; otherproportions deum and gaster. ofantenna as in Fig. 2. Thorax without distinct piliferous punctures; Aenasius vexans Kerrich sculpture on mesoscutum relatively deep, regu- lar, imbricate-reticulate, almost polygonally (Figs22, 23, 47, 50, 51, 62, 70, 86, 98, 99, 101) reticulate; scutellum with similar but slightly Aenasius vexans Kerrich, 1967: 202-203. Holo- finer sculpture; scutellum about as long as broad type 9> Brazil, USNM, examined. and with a distinct marginal carina dorsally in Aenasius tvexans Kerrich; Williamsetal., 1981. apical one-fifth or so and with a deep, very Aenasiussp. nrvexans Kerrich; Lohretal., 1990. conspicuous, longitudinal groove inits basal half; mid tibial spur clearly shorter than mid basitar- Diagnosis. Female (length: 0.76-1.35 mm): sus; forewing with distribution of setae at base head metallic green, dorsum of thorax similar, and proportions of venation as in Figs 20, 21; but duller; antenna more or less uniformly dark submarginalveinwith adistinctsubapicalhyaline brown; tegulae orange basally; antenna (Fig. 98) break; costal cell almost rectangular and con- with scape about 1.5 x as long as broad, sensory spicuously incised at apex, dorsally with three or areaofclavadivided obliquely; head in side view four lines of setae; no hyaline streak at apex of about twice as long as deep and evenlycurved to ENCYRTIDAE OFCOSTA RICA 125 top of scrobes, sides of facial impression slightly (M.N. Beg); 5$, St George, various localities bulging outwards and then abruptly angled and dates, vi—vii.1976 (J.S. Noyes); 5$, Nariva, inwards towards mouth; frontovertex (Fig. 86) Cocos Bay, coastland and mangrove swamp, about one-third to one-quarter head width, pilif- 28.vii.1976 (J.S. Noyes); 1$, St. Augustine, ex erous punctures below anterior ocellus shiny, mealybug on Lantana montividensis, V.1982 scrobes moderately deep, not sharply delimited (M.W.J. Cock); 6$, Curepe, ex Phenacoccussp. laterally; forewings (Fig. 22) apically hyaline on tomato, ix.1983 (F.D. Bennett); COLOM- with hyaline streak absent at apex of venation; BIA, 339, 26cf, Cali, CIAT, lab culture from postmarginal vein slightly longer than stigmal Venezuela, Bolivia Sta (Upsta) and Sucre Sta (Fig. 23); lower part ofouter margin offorewing (Ayacucho and Cumana), ex Phenacoccus her- clearly convex; apex of last tergite of gaster renion cassava, xi.1991 (J. Castillo); VENEZU- almost straight, hardly concave (Fig. 47); ovi- ELA, 119. UO\ Sucre, ex Phenacoccus herreni positor as in Figs 50, 51. Male (length; 0.65-0.87 15.vi.1989 (J. Castillo); GUYANA, 19, 1Q\ mm): very similar to female but scape (Fig. 70) Enmore Est., x.1979, ex Phenacocci herreni (as about2.5 x as longasbroad, all funicle segments P. manihoti misident.) on cassava, x.1979 (F.D. transverse F4-6 subequal and each nearly twice Bennett); 19- Enmore Est., ex Phenacoccus on as broad as long, clava about as long as F3-6 and cassava, 11.xi.1977 (M. Yaseen); Id", Dalgin, ex without any conspicuous differentiated sensory Phenacocciherreni (as P. manihoti misident.) on areas or sensilla; forewings (Fig. 62) not or cassava, xi.1978 (M. Yaseen); ld\ Diamond, ex hardly infuscate; tarsi yellow; mid tibial spur Phenacocciherreni (as P. manihoti misident.) on yellow, occasionally pale brown; lateral bristle cassava, xi.1979 (M. Yaseen); FRENCH GUI- on phallobase (Fig. 99) about 0.25 x as long as ANA, 19. Km 30 Highway, ex Phenacocci her- aedeagus which is broad, spatulate and about reni (as P. manihoti misident.) on cassava, one-third as long as mid tibia. xi.1978 (M. Yaseen); 39- Sinnamary, ex Phen- acocci herreni (as P. manihoti misident.) on Hosts. Recorded as a parasitoidofPhenacoccus cassava, xi.1978 (M. Yaseen); 29. 5d\ Mana, ex herreniCox & Williams (Williams ettil., 1981, as Phenacocciherreni (as P. manihoti misident.) on Aenasius Ivexans from yellow mealybug; Lohret cassava, xi.1978 (M. Yaseen); ECUADOR, 19, al., 1990), a pest ofcassava in areasofBrazil and Pichincha, Tinlandia, 800m, 2.ii.l983 (Masner & a potential pest ofcassava in Colombia, Venezu- Sharkey); PERU, Cuaco, Quilambamba, ela and the Guyanas (J.A. Castillo L., pers 24-26.xii.l983(L. Huggert); 1$, Madrede Dios, comm.). The eggs are laid into the 2nd and 3rd Pto Maldonado, 3.i.1984 (L. Huggert); BRA- instar nymphs and adult females (Castillo L., ZIL, 39, 2d", Santa Lucia, xi.1977, ex Phenacoc- pers comm.) cus on cassava, xi,1977 (M. Yaseen); 49. 30\ Macapa, ex Phenacoccus on cassava, 22.xi.1977 Use in biocontrol. The species successfully controls Phenacoccusherreni Cox & Williams on (M. Yaseen); 29. Porto Grande,ex Phenacoccus on cassava, 24.xi.1979 (M. Yaseen); 69. 2o\ cassava in Colombia (J. Castillo, pers. comm.). Para, Alanquer, ex Phenacoccus herreni on cas- Distribution. Mexico, Costa Rica, Tobago, sava, 11.x.1985 (B. Lohr). Material in BMNH, Trinidad, Colombia, Venezuela, Guyana, CNC, USNM, TAMU, ZISP, PPRI, INBio. French Guiana, Ecuador, Peru, Brazil. Comments. Aenasius vexans can be mistaken Materialexamined. forphenacocci but is separated on the coloration Type material. Holotype $: BRAZIL, Sao of tegulae and relative length of postmarginal Paulo, ex Phenacoccus sp., xii.1935 (E. Hamble- vein (see comments underphenacocci). ton) (USNM). Paratypes: MEXICO, 2$, Magdalena Is., Tres Marias, v.1925 (H.H. Aenasiusphenacocci Bennett Kiefer); BRAZIL, 1$, Sao Paulo, ex Phenacoc- cus No 3, xii.1935 (E. Hambleton) (BMNH). (Figs 8, 9, 24, 25,48, 55,63) Non-type material: COSTA RICA, 6$, Gua- Aenasius phenacocci Bennett, 1957: 569-570. nacaste, Santa Rosa NP, various dates Holotype 9- Trinidad, USNM, examined. i.l986-iii.l987 (Janzen & Gauld); TOBAGO, AenasiusflandersiKerrich, 1967: 204, 221. Holo- 19, St Paul, Pamatuvier Valley, edge ofrainfor- type 9. USA, USNM, examined, syn. n. est, 20.vii.1976 (J.S. Noyes); TRINIDAD, 1$, I.C.T.A., ex Phenacoccus hibsici on Hibiscus, Diagnosis. Female (length: 0.88-1.84 mm): No 25, v.1953 (F.D. Bennett); 29, Curepe, head mostly metallic green or blue green; Santa Margarita Circular Road, 26.ix-26.x.l974 antenna uniformly black brown; pronotum and 126 J.S. NOYES ANDH. REN mesoscutum weakly metallic green or blue- 14.viii-6.ix.1986 (Janzen & Gauld); 1$, San green; scutellum with a weak purple or blue- Jose, Ciudad Colon, 800m, iii-iv.1990 (L. green lustre; tegulae dark brown; antenna (Fig. Fournier); CAYMAN ISLANDS, Grand Cay- 8) with scape about 1.6 x as long as broad; man, West Bay, Willie Farringdon Drive, ex sensory area of clava divided obliquely into Ferrisia virgata on Sida, 16.x.1987 (F.D. Ben- almost equal portions (Fig. 9); head in side view nett); ST VINCENT, 1$, 207, West Indies very slightly more than twice as long as deep; 99-331, Aenasius hyettus Wlk. How.(H.H. slightly curved above scrobes, below scrobes Smith); TRINIDAD, 4$, I.C.T.A., ex Phen- more strongly curved towards mouth; frontover- acoccus gossypii on Hibiscus, v-vi.1953 (F.D. tex about one-third (small specimens) to one- Bennett); 15$, St. Augustine, ex Phenacoccus fifth (large specimens) head width, piliferous gossypii on Acalypha, v.1955 (F.D. Bennett); punctures below anterior ocellus shiny, scrobes 2$, lcf, St. Augustine, ex mealybugs, hi.1961 moderately deep, not sharply delimited; forew- (F.D. Bennett); 2$, Curepe, Sta. Margarita, ings (Fig. 24) apically hyaline without a hyaline 26.ix-26.x.l974 (M.N. Berg); 1$, St. George, streak at apex ofvenation; postmarginal vein not San Juan, ex mealybug; 18.vii.1976 (F.D. Ben- quite reaching level with apex of stigmal (Fig. nett); 1$, St Augustine, Malaise trap; ix.1976 25); lower part of outer margin of forewing very (F.D. Bennett); 11$, St George, various locali- slightly convex; apex of last tergite of gaster ties, vi-viii.1976(J.S. Noyes, F.D. Bennett); 1$, inconspicuously medially incised (Fig. 48); ovi- St. Andrew, Oropuche, cocoa and banana plan- positor(Fig. 55) about aslongasmid tibia. Male: tations, 28.V.1976 (J.S. Noyes); 3$, CIBC lab indistinguishable from that of advena Compere culture ex Phenacoccus grenadensis, 1977 (M. (see diagnosis); forewing hyaline (Fig. 63). Yaseen); 1$, Maracas Valley, xii.1977 (F.D. Bennett); 3$, Curepe, ex Phenacoccus gossypii Hosts. Recorded asaparasitoid ofPhenacoccus on Lantana montividensis, iv.1977 (F.D. Ben- madeirensisGreen (asP. gossypii, probable misi- nett); 5$, Curepe, ex Phenacoccus grenadensis dentification, D. Williams, pers. comm.) (Ben- on Cordia curassavica, iii.1979 (F.D. Bennett); nett, 1957) and from the same host on Acalypha 2$, Curepe, CIBC lab culture on Phenacoccus sp., Hibiscus sp. and an unidentified mealybug grenadensis, iv.1979 (F.D. BEnnett); ld\ ex on Pittosporum and cotton (Kerrich, 1967). Also Phencacoccus grenadensis on Tussacia, x.1979 recorded from Phenacoccus herreni Cox & Will- (F.D. Bennett); 1$, St George, Arima Valley miP.oanmtgsiosv(siLydopehinrisieTstoawlP.nh,es1ne9an9c0do)c.&cTuhsCeogcrrkeeencraoedrledlnssoibnselGLoraweneftnraon&ma N(1osytesm)i;le)1,gyendgaendrofomroaripnhfo,reSstt,Ge3.ovrigi.e1,97S6t A(Ju.sS-. , gustine, malaise trap, 15.vii-13.viii.1976 (J.S. Laingon Cordiacurassavica and Tussacia sp. are Noyes); GUYANA, 1$, Georgetown, on orna- probably all misidentifications of P. madeirensis mentals; 15.x.1961 (F.D. Bennett);4$, Enmore, (D. Williams, pers. comm.). Alsonoted hereasa ex Phenacoccus Imanihoti on cassava, x.1979 parasitoid of Ferrisia virgata (Cockerell) on Sida (M. Yaseen); COLOMBIA, 2$, Cali, CIAT, ex sp. Phenacoccus grenadensis on cassava, xi.1977 Distribution. USA (California), Costa Rica, (F.D. Bennett); 1$, Palmira, ex Phenacoccus on Cayman Islands, St Vincent, Trinidad, Colom- Acaplypha, 14.xi.1979 (F.D. Bennett); ECUA- bia, Guyana, French Guiana, Ecuador, Peru, DOR, 1$, Rio Palenque, forest, 4.ii.l983 (L. Uruguay. Huggert). Material in BMNH, INBio. Materialexamined. Comments. We have been unable to find any Type material. Holotype $ of Aenasius phen- significant differences between the type material acocci, TRINIDAD, I.C.T.A., ex Phenacoccus of flandersi examined and the series of phen- gossypii on Acalypha, v.1955 (F.D. Bennett) acocci listed above. We therefore treat the two (USNM Type No 63501). Paratypes ofAenasius names as synonymous. A. phenacocci is phenacocci: TRINIDAD, 2$, I.C.T.A., St. extremely close to advena differing only in the Augustine, ex Phenacoccus gossypii on Aca- colour of the funicle segments, other differences lypha,vi.1955 (F.D. Bennett) (inBMNH). Holo- given byKerrich (1967) areseeminglyunreliable. type $ of Aenasius flandersi: USA, California, For the present, we continue to treat the two San Diego, Balboa Park, on Pittosporum, species as valid until further material becomes 15.viii.1958 (S. Flanders) (Type No 2122 available. USNM). Superficially,Aenasiusphenacoccimaybemis- Non-type material. COSTA RICA, 1$, Gua- taken for vexans, but is distinguished on the nacaste, Santa Rosa NP, Hacienda 1-0, completely brown tegulae and postmarginal vein