Encyrtidae (Hymenoptera: Chalcidoidea) of Costa Rica: the genera and species associated with jumping plant-lice (Homoptera: Psylloidea)

60 Pages·1996·20.3 MB·

by  John S Noyes

8
Bull. nut. Hist. Mus. Land. (Ent.) 65(2): 105-164 Issued 28 November 1996
THE NATURAL
Encyrtidae (Hymenoptera:
Chalcidoidea) of Costa Rica: the 1 1 L-° 1998
genera and species associated with
jumping plant-lice (Homoptera:
Psylloidea).
JOHNS. NOYES
k
DepartmentofEntomology, TheNatural HistoryMuseum. London SW75BD,
London, England
PAUL HANSON
DepartmentofBiology, UniversityofCostaRica, San Pedro, SanJose, Costa Rica
CONTENTS
Synopsis 105
Introduction 105
Depositories 106
Acknowledgements 106
Abbreviations used in text 106
Review ofgenera and species 107
Key to genera 107
Revision of species 107
Cercobelus Walker 107
Trechnites Thomson 1 12
Homalotylus Mayr 113
Caldencyrtus gen. nov 114
Metaphycus Mercet
11
Sectiliclava Hoffer 121
Prionomitus Mayr 123
Psyllaephagus Ashmead 125
References 138
Illustrations 142
Index to scientific names 163
SYNOPSIS. Eightgeneraand 26 species ofencyrtids found in association withjumpingplant
lice(Psylloidea)inCostaRicaarereviewed.Threegeneraarerecordedforthefirsttimefromthe
Neotropicalregion,andonegenusand21 speciesaredescribedasnew.Dichotomouskeystoall
genera andspecies are provided,each species is furthercharacterisedbyadiagnosisandnotes
are provided on distribution and host associations where known.
INTRODUCTION Such projects normally devote large areas to rela-
tively few tree species in order to make plantations
easier to manage. The resulting increased density of
Alarming rates ofdeforestation in tropical countries individual tree species will inevitably heighten the
have generated considerable interest in reforestation risk of plantation trees being severely damaged by
projects utilising native trees (Butterfield, 1994). outbreaks of indigenous tree-feeding insect groups.
©TheNaturalHistoryMuseum, 1996 /2(?aJ ? «0 I %. (
106 J.S. NOYES ANDP. HANSON
One suchgroupofpotential forestry pests, thejump- DEPOSITORIES
ing plant-lice or Psylloidea, is particularly species
rich in Neotropical forests and in CostaRica species
are known to feed on several important native tree CNC Canadian National Collection, Ottawa, Canada
genera, eg. Alaroa, Cedrela, Haematoxylum, BMNH The Natural History Museum, London, UK
Hymenaea, Inga, Lonchocarpus Pentaclethra and INBio Instituto National de Biodiversidad. Costa Rica
Virola (D. Hollis, pers. comm.). The use of pesti- NMHS Naturhistoriska Riksmuseet, Stockholm,
Sweden
cides to control new forest pests may prove to be
ineffective, uneconomic orenvironmentally unsound NUMCIRD NMautsieonoaldeMIunsseecutoms,ofUnIirveelrasndi.tyDuobflCion,stEairReica,
and therefore other control measures, such as bio-
logical control, will be required. A taxonomic study USNM CUonsittaedRiStcaates National Museum, Washington
of the existing parasitoids of psyllids will provide D.C., USA
the necessary foundation for biological control of
these potential pests. One group of parasitoids of
particular importance is the chalcidoid family Acknowledgements. We thank Dr ME. Schauff
Einncclyurdteisdamea(nHyymsepneocipetserwah;icChhaalrceidsopiedceiaal)isbtecparuismeariyt t(hUeSlNoMa)n,orDrgiLf.t Mofasmnateerri(alC.NCO)uransdpeDciralJ.tHhuabnkesrg(oCNtCo)Pafomr
parasitoids ofjumping plant-lice. At least one Costa
Mitchell forherpainstaking sorting ofMalaisetrapcatches
Rican species, Psyllaephagus yaseeni Noyes, has fromSantaRosaNational Park. DaveHollis (NHM)helped
been introduced into various countries from Hawaii us by checking the psyllid names and several friends and
to the Indian subcontinent in order to aid in the colleaguesprovideduswithinformationrelatingtomultiple
control of Heteropsylla cubana Crawford, a pest of speciesrearings froma single hostspecies, although notall
Leucaena leucocephala (see below). This plant is examples were used (Jacques vanAlphen, Lex Kraaijeveld,
important as cattle fodder, green manure, a shade John LaSalle,AndrewPolaszek, David Rosen,JimWoolley
plant for coffee and cocoa and as a nitrogen fixing andBobZuparko).WewouldliketothankAldodeOyarzabal
plant in alley cropping. for the colour illustration of Psyllaephagus trioziphagus,
Inadditiontotheiruseinbiologicalcontrol,psyllid SectiliclavaisisandMetaphycuselectra(Figs 1-3).Finally,
parasitoids may be useful forstudies in host speciali- we would like to acknowledge financial support for this
sation orperhaps even co-evolution. Species in some work from the Vicerrectoria de Investigaciones of the
taxa, such as Psyllaephagus, appear to have special- Universidad de Costa Rica.
ised and diversified on psyllids whereas others, such
asafewspeciesofMetaphycus,appeartohaveswitched
from their normal scale-insect hosts (Homoptera: ABBREVIATIONS USED IN TEXT
Coccoidea) to psyllids. Further to this, psyllids that
inducegallformationseemtobeattackedprimarilyby
twosetsofparasitoids: thosethatspecialiseonpsyllid AL length ofaedeagus
hosts and those that generally attack gall-forming CCLW lmeanxgtihmoufmcowsitdatlhceolflcoofstfaolrceewlilnogfforewing
hosts.
AsaresultofrecentfieldworkinCostaRicaseveral F1,F2,etc. firstfuniclesegment, secondfuniclesegment.
etc.
species ofencyrtid parasitoids have been reared from
EL maximum eye length
jumping plant-lice and their galls. Most are primary EW maximum eye width
parasitoids ofpsyllids, whilst some areparasitoids of FV minimum frontovertex width
insects otherwise associated with psyllid galls, eg. FWL forewing length
parasitoidsofcoccinellidlarvaefeedingongall-form- FWW forewing width
ing psyllids. Additional material, belonging to these GL maximum length ofgonostylus (orthird
encyrtidgroups,hasalsobeenmadeavailablethrough valvula)
theextensiveMalaisetrappingprogrammeundertaken HI depth ofincised partofposteriormargin of
in Costa Rica since 1985 (see Noyes & Ren Hui, hypopygium
1995).Thepresentstudythusdirectlyresultsfromthis HW head width
work and is intended to facilitate the identification of HWL hindwing length
Encyrtidae that may be reared from psyllids or their HWW hindwing width
galls inCostaRica, oradjacentareas. Itisthe second MS malarspace (the shortestdistance from the
inaseriesthatisaimedateventuallyrevisingallCosta eye to mouth margin)
Rican Encyrtidae. MT mid tibia length
MTS mid tibial spur length
OL ovipositor length
OOL ocular-ocellar line, orthe shortest distance
COSTA RICAN ENCYRTIDAE ASSOCIATED WITH PSYLLOIDEA 107
between each posteriorocellus and the CERCOBELUS Walker
adjacent eye margin
POL posteriorocellar line, orthe shortestdistance CercobelusWalker, 1842: vi.TypespeciesCercobelus
between the two posteriorocelli jugaeusWalker, by monotypy.
SL scape length (excluding radicle)
SW maximum scape width Diagnosticcharacters. Female.Antennainserted
close to mouth margin and with funicle 4-segmented
Key to Costa Rican genera of andclava3-segmented;headovalorsphericalinfacial
Encyrtidae associated with psylloids or view witheyesrelatively small, notlongerthan mini-
their galls mum width of frontovertex, or hardly so; inner eye
margins slightly emarginate; occipital margin sharp;
mandibles with one very small tooth and a broad
1 Antennae with fewerthan 6 funicle segments 2 truncation or three or four very short, obtuse teeth;
—
Antennae with 6 funicle segments 3 forewingswithmarginalveinvaryingfromveryshort,
almost absent, to at least twice as long as broad;
2 Antennae with 4 funicle segmentsCercobelus (p. 107)
— postmarginal vein almost absent; setae in basal cell
Antennae with 5 funicle segments Trechnites (p. 112) much less dense than in disc; gaster, somewhat elon-
3 Mesoscutum with notaular lines complete; forewing gateandwithsegmentstelescopic,theovipositorshort
with stigmal vein abruptly bent below marginal vein and notmorethan0.5 xaslongasmidtibia,orhardly
(Fig. 28) so; ovipositor with a unique, highly modified struc-
Homalotylus (p. 113) ture.Male.Similartofemale,butantennausuallywith
— segmentsrelativelyslightlylongerandclothedinsetae
Notaular lines absent or not reaching more than half
way across mesoscutum; forewing with stigmal vein which are longer than diameter of segments; clava
straight or slightly curved, not abruptly bent below entire;genitaliawithaedeagusapically veryacuteand
marginal vein digitidistinctbutrelativelyslenderandapicallyarmed
4 with a pairofhooks.
4 Forewings with marginal vein at least 3 x as long as BIOLOGY. The only previously known species, the
broad (Fig. 31) Caldencyrtiis (p. I 14) EuropeanC.jugaeus(Walker),hasbeenrecordedasa
— Marginal vein not more than 2 x as long as broad, parasitoid of Psyilopsisfraximcola Forster, Pfraxini
mostly punctiform or absent (Figs 37, 39, 40, 43, 47, Linnaeus and Psyllopsis proprius Loginova
etc.) 5 (Homoptera: Psyllidae) (see Trjapitzin, 1989).
5 Mandiblestridentate(Fig. 132);headandthoraxyellow Distribution. Althoughthegenusisprobablycos-
to dark brown without metallic blue, green or purple mopolitan, intheNewWorldithasonlybeenfoundin
lustre Metaphycus (p. 118) MexicoandCostaRicaand hasyet tobe found in the
—
Mandibles withoneortwoteeth and atruncation (Figs eastern Palaearctic.
138, 139, 143-146), four teeth (Fig. 133), or more or Identificationofspecies. Otherthanthefivespe-
lessedentate (Fig. 137); headandthorax withat leasta
slight metallic green, blue orpurple lustre 6 cies described below, there is only a single described
species from Europe (jugaeus Walker) and several
6 Apicalsegmentofforetarsusenlarged,nearlyaslongas undescribed species from Mexico, Africa, India,
remainingsegmentstogetherandatleasttwiceas wide Sarawak and Australia (material in BMNH, TAMU,
(Fig. 2) Sectiliclava (p. 121) see also Noyes & Hayat, 1984).
—
Apicalsegmentofforetarsusnotenlarged,orhardlyso,
less than halfas long as remaining segments oftarsus
and not more than 1.5 x as wide 7 Key to Costa Rican species ofCercobelus
7 Postero-lateral margin ofmesoscutum, anteriortoeach
tegula, with a deep, distinct submarginal indentation 1 Clava 3-segmented (females) 2
which is about as wide as the tegula. FEMALE: — Clava entire (males) 6
hypopygium reachingpastapexoflasttergiteofgaster
and enclosing most ofovipositor FEMALES
Prionomitiis (p. 123)
— 2 F2-F4 transverse orquadrate 3
Mesoscutum without an indentation anterior to the —
tegulae, sometimes with a shallow impression adjacent F2-F4 clearly longerthan broad 4
to the antero-lateral margins. FEMALE: hypopygium 3 Forewings only lightly infuscate below apex of vena-
not reaching apex ofgaster Psyllaephagus (p. 125) tion (Fig. 5); Fl transverse, clava about as long as
funicle (Fig. 4) isara (p. 108)
108 J.S. NOYES AND P. HANSON
—
Forewings with a conspicuous interrupted transverse infuscate; coxae and femora very dark brown; fore
fascia from apex of venation and an elongate, dark tibia dark brown, except extreme apex which is yel-
brown,wedge-shapedmarktoapex(Fig. 10);Flslightly lowish;foretarsuswithbasalthreesegmentsyellowish
longer than broad, clava about as long as F2-F4 com- apicalsegmentsbrown;midtibiawithbasaltwo-thirds
bined (Fig. 9) sithon (p. 109) dark brown, apical one-third yellow; mid tibial spur
4 Pronotum very long and clearly longer than meso- pale yellow; mid tarsus yellow with apical segment
scutummedially;marginalveinofforewingatleast3x brown; hind tibia dark brown with basal one-third
aslongas broad (Fig. 16) ulixes (p. 110) yellow; hind tarsus dark brown; gaster dark brown,
—
Pronotum moderately long, but clearly shorter than basal tergite with aslight purple andgreen sheen.
mesoscutum medially; marginal vein of forewing not Headwithfairlyregular,raisedreticulate sculpture
longerthan broad (Fig. 19) 5 on frontovertex of mesh slightly larger than an eye
facet,sculpturebecomingmoreelongateandlessregu-
5 Propodeum with two, posteriorly diverging, median
carinae; mid legs extensively marked with brown lar on lower parts offace and genae; mandibles with
godoyae (p. 110) one very small lower tooth and a very broad, finely
— serrated truncation (Fig. 101); inner margins ofeyes
Propodeum with only asingle mediancarina; mid legs slightly sinuate;eyesinconspicuouslyhairyandsepa-
yellow, with mid tibiae hardly brownish basally ratedfrom occipital marginby about the longitudinal
daphne (p. Ill) diameter of a posterior ocellus; ocelli forming a
MALES strongly obtuse angle; occipital margin sharply cari-
6 Forewings hyaline without even an inconspicuous nate; transverse midline ofantennal toruli level with
— infuscate pattern isara (p. 108) 4l.owReesltateiyveemmaeragsiunrs;empernotpso:rtHioWns7o2f;anFtVen4n4a;ePasOiLn F1i9g;
Forewings withan infuscate pattern in apical half ....7 OOL 10;OCL5;EL38;EW25;MS22;SL28;SW8.
7 Setae on funicle at least twice as long as diameter of Mesoscutum with fairly regular raised, reticulate
funicle segments (Fig. 21) sithon (p. 109) sculpture, slightly deeper than that on frontovertex,
— anteriorly tending to imbricate-reticulate; scutellum
Setae on funicle at most about as long as diameter of
funicle segments ulixes (p. 110) withsimilar,butmore shallowsculpture,especiallyat
apexwhichisalmostsmooth;mesoscutumabout 1.5x
Cercobelus isara sp. nov. as broad as long and at least three times as long as
pronotummediallyindorsalview;venationanddistri-
(Figs 4-8, 100-102) bution ofsetae atbaseofforewingas in Figs 6and7;
Diagnosis. Female(length 1.30-1.41mm):allfuni- costalcellabout 10xaslongasbroad;midtibial spur
cle segments transverse and clava about as long as long and slender, nearly 10 x as long as broad; hind
ftuonoitchlaen(dFiag.ve4r)y;bmraonaddi,blfienselwyitsehrroanteedvetrryunscmaatliolnl(oFwiegr. FtliobWnigaWsalsighbtrlHoyaWdb.LroRaedleanteiHdveWanmWdeafsluatrteemneedn,tsa:bouFtW4L.5 2x09a,s
101); mesoscutum at least 3 x as long as pronotum 80; 133, 35.
medially in dorsal view; forewings slightly infuscate Gaster slightly shorter than the thorax; ovipositor
below stigmal vein and with apical one-third incon- (Fig. 100)slightlyexsertedatapex;lasttergiteclearly
spicuously infuscate (Fig. 5); marginal vein very constricted subapically and with apex broadly con-
slightly longerthanbroadandtouchinganteriorwing cave andtruncate. Relative measurements (paratype):
OL 19;GL5[MT59].
margin(Fig.7);propodeumaboutone-fifthaslongas
scutellum;gasterslightlyshorterthanthorax;oviposi- Male. Very similar to female except for antenna
tor slightly more than one-third length of mid tibia. (Fig. 8) and genitalia (Fig. 102). Relative measure-
Male(lengthabout 1.90mm): similartofemale,butall ments: FV 30, SL 18; MT59,AL 29.
funicle segments longer than broad and clothed in
setae that are longer than diameterofsegments (Fig. Variation. Very little in available material.
8); aedeagus about halflength ofmidtibia. HOSTS. Unknown.
Female(holotype). Length 1.35mm. Headgener- Distribution. Costa Rica.
allyblackishwithpurplelustre,ocellarareagreenish;
Material examined.
mesoscutum similar but with a green lustre, margins
purplish; scutellum and axillae mostly with coppery Holotype 9, Costa Rica, Puntarenas, San Vito, Est.
Las Alturas, 1500m, i.1992 (P. Hanson) (BMNH).
purplelustre; antennawith scapedarkbrown,pedicel
darkbrownwithapextestaceous,flagellumtestaceous- Paratypes: lc?, Guanacaste, Volcan Cacao, Cerro
yellow, clava slightly darker apically; wings mostly Pedregal, 1000m, ii—iv.1989 (I.D. Gauld); lc?, same
hyaline but with a weakly infuscate area below the data as holotype but xi.1991 (P. Hanson, C. Godoy);
Id1 same data as holotype; 2?, San Jose, Zurqui de
stigmal vein and apical one-third or so very weakly ,
COSTA RICAN ENCYRTIDAE ASSOCIATED WITH PSYLLOIDEA 109
Moravia, 1600m, iii.1992 and v.1994 (P. Hanson). separatedfromoccipital margin byaboutthe longitu-
Paratypes in BMNH, INBio. dinal diameterofaposteriorocellus; ocelli forminga
Comments. Cercobelusisaracanbeseparatedfrom strongly obtuse angle; occipital margin sharply cari-
nate; transverse midlineofantennal toruli about level
otherdescribed species ofthe genus by the relatively
withlowesteyemargins;proportionsofantennaeasin
short funicle segments, all other described species Fig 9. Relative measurements: HW 90; FV 57; POL
having the funicle segments distinctly longer than 29; OOL 13; OCL 8; EL46; EW 30; MS 21; SL 37;
broad.Itisverysimilartoanundescribedspeciesfrom SW
Mexico (BMNH, TAMU) which also has relatively 8.5.
Mesoscutum with fairly regular raised, reticulate
short funicle segments and similar mandibular struc-
sculpture,similartofrontovertex;scutellumwithsimi-
ture, butinthat speciesthe segmentsare very slightly
lar, but more shallow sculpture, especially at apex
longerthan broadandtheantennaareclearly inserted
which is almost smooth; mesoscutum about 1.5 x as
below the level ofthe lowest eye margins.
broadaslongandaboutone-thirdlongerthanpronotum
medially in dorsal view; venation and distribution of
Cercobelus sithon sp. nov.
setae at base offorewing as in Figs 1 1 and 12; costal
(Figs 9-12,20,21,105,106) cellabout8xaslongasbroad;midtibialspurlongand
slender,about 1 xaslongasbroad;hindtibiaslightly
Diagnosis. Female (length 1.90mm): Fl slightly broadenedandflattened,about4.9xaslongasbroad;
longer than broad, F2-F4 quadrate; clava about as propodeum with a single, inconspicuous median ca-
long as F2-F4combined (Fig. 9); mesoscutum about rina in is posterior half. Relative measurements
one-third longer than pronotum medially in dorsal (holotype): FWL 264. FWW 102; HWL 180, HWW
view; forewings with an interrupted, dark brown fas- 46.
ciafrom stigmal vein toposteriormargin and with an Gaster slightly shorter than the thorax; ovipositor
elongate wedge-shapeddark brown mark beyond this hardly exserted at apex.
towingapex (Fig. 10): marginal vein punctiform and
touching anterior wing margin (Fig. 12); propodeum Male. The males that we tentatively associate with
about one-third as long as scutellum; gaster slightly the female are generally similar to female other than
shorter than thorax. Male (length 1.80-2.00mm): an- slightly darkercoloration oflegs, palerinfuscationof
tennawith all funicle segments longerthan broadand forewings, relatively slightly narrower frontovertex
clothed in setae with are at least twice as long as and structureofantennae (Fig. 21)andgenitalia(Fig.
diameter of segments (Fig. 21); otherwise similar to 106). Fore and mid coxae dark brown, hind coxae
female. varying from yellow to dark brown; mid tibiae with
apical twothirds mixeddark brown,especially proxi-
Female (holotype). Head, pronotum and meso-
mally; hind tarsi dark brown; eyes separated from
scutum generally blackish with a green lustre, mixed occipital marginbyaboutone-thirddiameterofposte-
slightly brassy and coppery, occipital margin purple; rior ocellus; mandibles with two very short ventral
scutellum and axillae mostly with coppery purple
teethandabroad, minutelydenticulate,dorsaltrunca-
lustre; antenna with scape pale amber, pedicel dark tion (Fig. 105). Relative measurements: (paratype 1):
brown with apex testaceous, flagellum testaceous- HW 95, FV 55; EL 43, EW 33, MS 29; SL 27, SW
brown; forewing (Fig. 10) with base hyaline todistal 10.5; (paratype 2)AL 28.5, MT 74.
marginoflineacalva,an interruptedtransversebrown
fascia from postmarginal and stigmal veins to hind Variation. There is some variation in the colora-
wing margin and an elongate, dark brown, wedge tionofthehindcoxaeinthemalesasnotedabove.The
shaped mark from centre of wing to apex; fore legs male figured (Figs 20 and 21) has Fl significantly
pale amber, fore coxae with a small, external dark longerthaninothermalesbeingabout 1.5xaslongas
brownmark;midcoxaedarkbrown,remainderofmid F6 and the stigmal vein curved whereas in the other
legs pale amberbut forextreme bases oftibiae which malesFl issubequaltoF6andthestigmalveinismore
arebrownandbeyondthiswhite;hindlegswithcoxae orless straight (see alsocomments below).
and basal two-thirds offemora pale amber, apices of
femoraandapicalthree-quartersoftibiaedarkbrown, Hosts. Unknown.
basalpartoftibiaeandmostoftarsiwhite,apicaltarsal Distribution. Costa Rica.
segmentdark brown; gasterdark purple-brown.
Head with fairly regular, raised reticulate sculpture Material examined.
on frontovertex of mesh slightly larger than an eye Holotype 9, Costa Rica, Guanacaste P., Guanacaste
facet,sculpturebecomingmoreelongateandlessregu- NP, Cacao, 950m, 13.ii.1995 (J.S. Noyes) (BMNH).
laron lowerparts offace andgenae; innermarginsof Paratypes: Costa Rica, Id1 Puntarenas, Monteverde
,
eyes slightly sinuate; eyes inconspicuously hairy and Reserve, 1550m, 16.viii.1986 (L. Masner); lcJ,
110 J.S. NOYES ANDP. HANSON
Puntarenas, Monteverde, St Luis Valley, 17.viii.1986 onfrontovertexofmeshsubequaltothediameterofan
(L. Masner); 3d1, Guanacaste P., Guanacaste NP, Ca- eyefacet, sculpturebecoming moreelongate and less
cao. 950-1200m, 13.ii.1995 (L. Masner). Holotype regularbelow topofantennal toruli; innermarginsof
andparatypesinBMNH,paratypesinBMNH,INBio, eyes slightly sinuate; eyes inconspicuously hairy and
CNC. separated from occipital margin by slightly less than
Comments. Althoughthemalesdifferslightlyfrom the diameter ofan ocellus; ocelli forming a strongly
obtuse angle; occipital margin sharply carinate; pro-
the holotype female in having a relatively slightly
nbaraeercracouowsneefridfterhnoetntmotavhneatrdtietbxhl,eeysla,bregevlreonneagyteistoona,nthdewdiasnragkmeerselstepagetsciiowene,s pm5o0er;nttEiso:WnsH3oW2f;a8Mn7St;eFn2n4Va;e4S6aL;sP4i4nO;LFSig2W1;173..OOReLla9t;ivOeCmLeas9;urEeL-
Mesoscutum with fairly regular raised, reticulate
infuscate pattern on the forewings and general struc-
sculpture, similartothat on frontovertex, butperhaps
tureofthethoraxandpropodeumarevirtuallyidentical.
ThevariationnotedaboveintherelativelengthsofFl alittleshallowerandfiner;scutellumwithsimilar,but
and F6 and shape ofstigmal vein may suggest that a more shallow sculpture, especially at apex which is
almost smooth; mesoscutum about 2 x as broad as
sixth CostaRican species isrepresented in this mate-
rial,butforthepresentweareacceptingthisasnormal long and slightly shorter than pronotum medially in
dorsal view; scutellum clothed in very conspicuous,
variation within the species.
longsetae;venationanddistributionofsetaeatbaseof
C. sithon is avery distinctive species easily recog-
nisable by the infuscate pattern ofthe forewings. forewingasinFigs 15and 16;costalcellabout 12xas
long as broad; mid tibia with a lineofvery conspicu-
ousbristlesalongitslengthexternally;midtibialspur
Cercobelus ulixes sp. nov. longandslender,about8xaslongasbroad;hindtibia
(Figs 13-16) slightlybroadenedandflattened,onlyalittleover6x
as long as broad. Relative measurements: FWL 305,
Diagnosis. Female (length 2.65mm): antenna(Fig. FWW 109; HWL 225, HWW48.
13) with funicle segments subequal and much longer Gaster very slender and elongate, excluding the
thanbroad; mandibleswithone shorttoothandavery ovipositorabout3xaslongasbroadandaboutaslong
broad, slightlyconcave, smoothtruncation; pronotum as head and thorax together; cereal plates in apical
medially in dorsal view at least a little longer than one-third; ovipositor slightly exserted at apex, the
mesoscutum; forewings (Fig. 14) conspicuously exserted partnearly halfas long as midtibial spur.
infuscatefromlineacalvatowingapex;marginalvein
aboutfourorfivetimesaslongasbroadandtouching Male. Length 1.84mm. Generally very similar to
anteriorwingmargin(Fig. 16);propodeumlongmedi- female but frontovertex slightly wider, antenna with
ally,abouttwo-thirdsaslongasscutellum;gastervery Fl nearly 2 x as long as pedicel and other flagellar
elongate, clearly longer than head and thorax com- segmentsrelativelymoreelongatesothatflagellumis
bined;ovipositordistinctlyexsertedwithexsertedpart about 1.4 x as long as width ofhead, clava solid and
nearly half as long as mid tibial spur. Male (length gaster about two-thirds as long as thorax. Setae on
1.84mm): very similarto female except forrelatively funicle not longerthan width ofsegments.
longer and broader flagellar segments and shorter Hosts. Unknown.
gaster.
Distribution. Costa Rica.
Female(holotype). Body generally blackishwith
Material examined.
a weak brassy lustre on head and dorsum ofthorax;
Holotype?,CostaRica,Cartago,LaCangreja, 1950m,
scape dark brown, pedicel and flagellum brown;
vii.1991 (P.Hanson,C.Godoy).Paratype,CostaRica.
forewings (Fig. 14)infuscatefrom lineacalvatoapex
Id", Heredia, UvitadeSanRafael, 1700m, 10.iii.1991
with a pair of opposite, longitudinal hyaline streaks (P. Hanson). Material in BMNH.
distad of venation, the lower one extending to linea
calva; all coxae blackish; fore femora dark brown in Comments. Cercobelusulixesfemalescanbesepa-
basalhalforso,apexamber;foretibaeandtarsiamber; rated from those ofall other species ofCercobelusby
mid femur dark brown with basal one-third whitish; the extremely elongate pronotum. propodeum and
midtibiawithanarrow,darkbrownbasalring,awhite gaster. The male differs from other Costa Rican spe-
subbasal ring followed by a dark brown ring, distal cies in having setae on the flagellum which are not
halfyellowish; mid tibial spur and tarsi yellow; hind longerthan the diameterofthe segments.
femur dark brown proximally amber; hind tibia dark
brown, slightly less than proximal one-third white;
Cercobelusgodoyae sp. nov.
hind tarsus darkbrown.
Headwith fairlyregular, raisedreticulate sculpture (Figs 17-19, 103 and 104)
COSTA RICAN ENCYRTIDAE ASSOCIATED WITH PSYLLOIDEA 111
Diagnosis. Female(length2.01-2.21mm): antenna HOSTS. RearedfromKatacephalasp.nov.(D.Hollis,
(Fig. 17) with Fl and F2 subequal and longer than pers. comm.) (Psyllidae) on Eugenia carthagensis
broad; mandibles (Fig. 103) with three, short, obtuse (Myrtaceae).
teeth; mesoscutum about 2.5 x as long as pronotum Distribution. Costa Rica (Cartago).
medially in dorsal view; forewings conspicuously
Material examined.
infuscate below apex ofvenation (Fig. 18); marginal
vein punctiform, not quite touching anterior wing Holotype 9, Costa Rica, Cartago, Tobasi, 1500m, ex
margin (Fig. 19); propodeum long medially, nearly Katacephala (D. Hollis, sp. nov. #2, pers. comm.) on
one-third length of scutellum and with a pair of Myrtaceae (Eugenia carthagensis, D. Hollis, pers.
posteriorlydiverging,mediancarinae;gasterelongate, comm.), 6.iv.1993 (P. Hanson, D. Hollis). Paratypes:
aboutas longasthorax;ovipositordistinctlylessthan Costa Rica, 29, samedataas holotype. Holotype and
0.5 x as long as mid tibia. Male: unknown. paratype in BMNH, paratype in INBio.
Female(holotype). Length2.21mm. Bodygener- Comments. Cercobelus godoyae can be separated
ally blackishwithaweakbrassylustreonhead,green fromtheotherspeciesofCercobelusfromCostaRica
lustreonmesoscutumandpurplishlustreonscutellum; bythecharactersgiven inthe key (seealsocomments
antennae amber-testaceous; wings mostly hyaline but under daphne) and from the only other described
withaconspicuoustriangularinfuscateareaabouthalf species (jugaeusWalker from Europe) by the relative
wayacrosswingfromstigmalvein(Fig. 18)extending lengthofthefunicie segments, forewingvenationand
as a narrow median streak towards wing apex; all the infuscate forewing. C. jugaeus has Fl distinctly
coxae blackish; fore femoradark brown in basal half longerthanF2andthemarginalveinoftheforewingat
orso,apex amber; foretibae andtarsi amber; midand leastabouttwiceaslongasbroadandtheforewingnot
hindfemoramostlydarkbrownbutamberbasally,the infuscate orhardly so.
hind femora distinctly so; mid tibiae basally dark
brown; apical half, and spuramber, tarsi amber; hind Cercobelus daphne sp. nov.
tibiae dark brown, almost black, tarsi dark brown.
Headwithfairlyregular,raisedreticulatesculptureon Diagnosis. Female (length 2.79mm): antenna with
frontovertex, this becoming more elongate and less all funicie segments longer than broad; mandibles
regular towards eyes and on lower parts of face and with a very short, obtuse lower tooth and a broad
genae; inner margins of eyes slightly sinuate; eyes truncation; mesoscutum slightly more than twice as
moderately hairy; ocelli forming a strongly obtuse long as pronotum medially in dorsal view; forewings
angle; occipital margin sharply carinate; proportions conspicuously infuscatebelowapexofvenation;mar-
of antennae as in Fig. 17, pedicel and flagellum to- ginal vein punctiform, not quite touching anterior
gether equal in length to head width. Relative wing margin; propodeum long medially, about one-
measurements:HW90;FV54;POL32;OOL9;OCL third length of scutellum; gaster elongate, nearly as
8; EL46; EW 32; MS 28; SL 39; SW 7.5. long as head and thorax together. Male: unknown.
Mesoscutum with fairly regular raised, reticulate Female. Bodygenerally blackish with aweakpurple
sculpture, slightly deeper than that on frontovertex, and bluish lustre on head and mesoscutum and blue,
anteriorly tending to imbricate-reticulate; scutellum purple and brassy lustre on scutellum; antennae with
withsimilar,butmoreshallow sculpture,especiallyat scapeyellowandflagellumbrown;wingsmostlyhya-
apexwhichisalmostsmooth;mesoscutumabout 1.5x linebutwithaconspicuoustriangularinfuscateareato
asbroadas long and about 2.5 xas longas pronotum abouthalfwayacrosswingfromstigmal veinextend-
medially in dorsal view; venation and distribution of ingasaninconspicuousnarrowmedianstreaktowards
setae at base offorewing as in Figs 18 and 19; costal wingapex;allcoxaeblackish;forefemoradarkbrown
cell about 10 x as long as broad; mid tibial spur long in basal half or so, apex amber; foretibae and tarsi
and slender, nearly 10 x as long as broad; hind tibia amber;foreand mid legs mostly yellow,the midtibia
slightlybroadenedandflattened,only alittleover4x marked a small area of dark brown basally, hind
as long as broad; propodeum with two, posteriorly femora basally yellow, remainder ofleg dark brown;
dFiWveLrging, FmeWdiWan caHriWnaLe. RelatHivWeWmeasurements: gasterdarkbrownwithsomepurpleandbrassyreflec-
253, 98; 150, 50.
tions.
Gasteraboutaslongasthorax;ovipositor(Fig. 104) Head withfairlyregular, raisedreticulatesculpture
slightlyexsertedatapex;lasttergiteclearlyconstricted onfrontovertex,thisbecomingmoreelongateandless
level withapicalgastralspiracleandbroadeningagain regular towards eyes and on lower parts offace and
subapically, its apex broadly truncate. Relative meas- genae; inner margins of eyes slightly sinuate; eyes
urements (paratype): OL 24; GL9 [MT 55]. moderately hairy; ocelli forming a strongly obtuse
Male. Unknown. angle;occipital marginsharplycarinate;antennawith
flagellum a little longer than head width with all
;
112 J.S. NOYES AND P. HANSON
funicle segmentsclearly longerthanbroad, Fl longer hypopygiumreachingapexofgaster,frequentlyreach-
than pedicel and nearly 2.5 x as long as broad, F4 ingpastapex oflast tergite andcompletely enclosing
about0.75 x as long as Fl and about 1.5 x as long as theovipositor;ovipositorslightlytostronglymodified
broad. Relativemeasurements: HW 104;FV62; POL with gonostyli varying fromfree to completely fused
35; OOL 11; OCL 9; EL 54; EW 38; MS 34; SL 51 with the second valvifers. Male hardly differing from
SW9. female, clava entire, pedicel sometimes enlarged;
Mesoscutum with fairly regular raised, reticulate parameres not developed, digiti elongate, each with
sculpture, slightly deeper than that on frontovertex, one ortwo apical teeth.
anteriorly tending to imbricate-reticulate; scutellum
Biology. Parasitoids of psyllid nymphs. The de-
with similar sculpture anteriorly, but more shallow
scribed species have been recorded from Caillardia
sculpturetowardsapexwhichisalmostsmooth;meso-
spp. and Psylla spp.
scutum about 1.7 x asbroad as long and about2 x as
long as pronotum medially in dorsal view; venation USE in BIOCONTROL. One species, T. psyllae
anddistributionofsetaeatbaseofforewingsimilarto (Ruschka), was been imported into California(USA)
thatofgodoyae(Fig. 18);costalcellabout 10xaslong from Switzerland for the control of Psylla pyricola
asbroad; midtibialspurlongandslender, nearly 10x Forsterin 1965 (Clausen, 1978). No furtherinforma-
as long as broad; hind tibia slightly broadened and tion is available. The same species is accredited with
flattened, nearly 5 x as long as broad. Relative meas- effective control of Psylla pyri Linnaeus in the pear
urements: FWL 320, FWW cl23; HWL 219, HWW orchardsofGeorgia(Chkhaidze& Khlopunov, 1971)
66.
Distribution. Cosmopolitan.
Gaster nearly as long as head and thorax together;
ovipositor slightly exserted at apex. Identification of species. Of the 17 described
species,fiveold world species arekeyedby Hayat,et
Male. Unknown.
al.(1975),sixPalaearcticspeciesbyTrjapitzin(1989)
Hosts. Unknown. and four southern African species are reviewed by
Prinsloo (1981).
Distribution. Costa Rica (Puntarenas).
Comments. Trechnites is probably closest to
Material examined.
Cercobelus,bothgeneraincludingspeciesparasiticon
H25o.lVo.t1y9p9e3 9(,MicChoasltsaki)R.icIan,BPMuNntHa.renas, Monteverde, psyllids and having reduced antennal segmentation
and similar modified ovipositor structure. The two
COMMENTS. Cercobelusdaphneisclosesttogodoyae generacanbe separatedfromeachotherby anumber
and can be separated from this species by its larger ofcharacters,butmostnotablythedifferenceinnumber
size, the pedicel and flagellum together being longer of funicle segments (see key), the complete lack of
than head width (equal in godoyae), relatively longer notaularlines in Cercobelus, andlessmodifiedgaster
gaster, paler fore and mid legs and presence ofonly in Trechnites (strongly telescopic in Cercobelus).
one mediancarinaon propodeum (two in godoyae).
Trechnites merops sp. nov.
TRECHNITES Thomson (Figs. 22-25, 107-110)
Trechnites Thomson, 1876: 118. Type species Diagnosis. Female (0.94-0.97mm): hind tarsi
TrechnitesfuscitarsisThomson by monotypy. brown,contrastingwiththeyellowforeandmidtarsi;
Psylledontus Crawford, 1910: 88. Type species mesoscutumshiningmetallicgreenwithnotaularlines
Psylledontus insidiosusCrawford, by original des- relatively conspicuous and meeting medially at the
ignation. Synonymy with Trechnites by Ferriere, posterior margin ofthe mesoscutum; scutellum rela-
1961. tivelydullwithveryfinepunctate-reticulatesculpture
whichcontrasts stronglywiththe shallowersculpture
Metallonella Girault, 1915: 77. Type species Metal-
lonellaaustraliensisGirault,byoriginaldesignation. ofthe mesoscutum; last tergite with a pair ofapical
Synonymy with Psylledontus by Peck, 1951 and lateralinvaginations;hypopygiumwithamedian,api-
with Trechnitesby Ferriere, 1961. cal extension which reaches past apex oflast tergite
and completely encloses the ovipositor. Male (0.87-
Diagnostic characters. Head and thorax gener- 0.90mm): similartofemalebutforgenitaliaandsolid
ally metallic green; funicle 5-segmented; clava clava.
3-segmented;mesoscutumwithnotaularlinespresent,
but superficial and inconspicuous, occasionally ab- Female (holotype). Length 0.97mm. Head black
sent;forewingwithmarginalveinpunctiform,virtually withastrongmetalliccoppery orpurplesheen mixed
absent; postmarginal vein virtually absent; withgreen, scrobalareadistinctlypurple; scapeblack
;
COSTA RICAN ENCYRTIDAE ASSOCIATED WITH PSYLLOIDEA 113
with apex yellow; flagellum dark brown; pronotum (I.D. Gauld). Holotype and paratypes in BMNH.
purple-brown;mesoscutumbrightmetallicgreen,pur-
Comments. T. meropscan be separated from other
plishtowardsposteriormargin;axillaecoppery-purple,
species ofthe genus by the relatively dull scutellum
dull;scutellumrelativelydullbutofsilkyappearance,
which has very finely punctate-reticulate sculpture
medially coppery-purple, towards sidesandapex me-
which is clearly much deeper than the more coarse
tallicgreen;coxaeandfemorablackish,basesoffemora
reticulate sculpture ofthe mesoscutum. In other spe-
pale yellow; tibiae blackish, apical one third of fore
cies the scutellum is generally at least as shiny as the
tibiae, mid tibial spur and slightly more than apical
halfofmidtibiapaleyellow;foreandmidtarsiyellow, mesoscutum and the sculpture is relatively shallow
and rather elongate, almost striate. There are also
hind tarsi dark brown; wings hyaline; gasterblackish
notabledifferencesinthe shapeoftheapexofthe last
with apurple and green sheen.
tergite and hypopygium, many species having the
Frontovertex with shallow, fairly regular reticulate
apical partofthe lasttergitemuch more highly modi-
sculptureofmesh a little largerthaneye facet; sculp- fiedthan inmeropsandthe hypopygium less strongly
ture in scrobal area irregular and longitudinally
extended posteriorly.
elongate;interantennalprominencewithveryshallow,
regular,reticulatesculpture;scrobeswideandmoreor
lesstouchingeyes;ocelliforminganacuteangle;eyes HOMALOTYLUS Mayr
not conspicuously hairy; mandibles (Fig. 107) with
two teeth and a broad truncation, the second tooth
HomalotylusMayr, 1876: 752.TypespeciesEncyrtus
obtuse, very short and almost absent; proportions of
HW flaminius Dalman, by subsequent designation of
antennaasinFig.22.Relativemeasurements: 64;
F1V8;2S1L;O30O;LSW108;.POL2;OCL3.5;EL43;EW32;MS NobArsihmmuesadT,ho1m9s0o0n., 1876: 1 16.Typespecies£>?cvrfMs
flaminius Dalman, by subsequent designation of
Mesoscutum with notaular lines well marked and
Timberlake, 1919. SynonymywithHomalotylusby
more or less meeting medially at posterior margin;
Timberlake, 1919.
mesoscutum with shallow, imbricate-reticulate sculp-
Mendozaniella Brethes, 1913: 97. Type species
ture,themeshfinermediallyandnearposteriormargin
Mendozaniella mirabilis Brethes, by monotypy.
axillae and scutellum with conspicuously deeper and
Synonymy withHomalotylusbyTimberlake, 1919.
finer,punctate-reticulatesculpture,extremeapexquite
Hemaenasoidea Girault, 1916: 307. Type species
smoothandpolished; venation and setation atbaseof
HemaenasoideaoculataGirault,bymonotypy.Syn-
forewing as in Figs 23 and 24; propodeum outside
onymy with Homalotylus by Timberlake, 1919.
spiracle with only one or two inconspicuous setae.
Relative measurements: FWL 153, FWW 63; HWL Anisotylus Timberlake, 1919: 170. Type species
HWW HomalotylussimilisAshmead,byoriginaldesigna-
90, 22. tion. Synonymy with Homalotylus by Noyes &
Gaster about as long as thorax; hypopygium (Fig.
Hayat, 1984.
109) medially produced at apex, extending past apex
Lepidaphycus Blanchard, 1936: 13. Type species
oflasttergiteandcompletelyenclosingtheovipositor;
LepidaphycusbosqiBlanchard,bymonotypy. Syn-
ovipositor(Fig. 108) with gonostyli partially fusedto onymy with Homalotylus by Blanchard, 1941.
second valvifers. Relative measurements (paratype):
OL21.5;GL7 [MT 28]. Neoaenasioidea Agarwal, 1966: 71. Type species
NeoaenasioideaindicaAgarwal, by original desig-
Male. Generally almostidentical tofemalebutdif- nation.SynonymywithHomalotylusbyHayat,Alam
&
fering in antenna(Fig. 25) with clavasolidand inthe Agarwal, 1975.
structure ofthe genitalia (Fig. 110). Diagnosticcharacters. Length about 1.5-2mm;
HOSTS. RearedfromTriozasp.nov.(D.Hollis,pers. males and females extremely similar and often diffi-
comm.) (Triozidae) on Clethra sp. (Clethraceae). culttoseparate;head(Fig. Ill)aboutaslongasbroad
orlonger;eyesmoreorlesskidney-shapedandatleast
Distribution. Costa Rica (700-1700m).
about 3 x as long as malarspace; mandible tridentate
Material examined. (Fig. 112);antenna(Fig.26)withscapesubcylindrical,
at least about 6 x as long as broad; funicle 6-seg-
Holotype 9, Costa Rica, Heredia, San Rafael, Uvita, mented with segments subequal in width and
1700m, ex psyllid on Clethra, iii.1992 (P. Hanson). subquadrateorconspicuouslylongerthanbroad;clava
Paratypes: Costa Rica, Id1, Puntarenas, Monteverde, unsegmented or with partial segmentation, with a
15.xii.l985-5.i.l986 (A. Forsyth); Id, Puntarenas,
conspicuous, oblique apical truncation; mesoscutum
Monteverde, Finca Canada, 1500-1600m, 23-
withnotaularlinescompleteandmoreorlessmeeting
2N7P.,ii9.1K9m91S(.BS.Ja.nStiancCleaciirl)i;a,lcE?s,tG.uPaintailclaas,t7e0,0Gmu,anvaic.a1s9t8e9 2at7)popsatretrliyorinmfaurscgaitneoafndmewsiotshcusttiugmm;alfovreeinwignegnser(aFlilgy.
114 J.S. NOYES ANDP. HANSON
abruptly bent immediately below marginal vein (Fig. about half way along gaster; ovipositor distinctly
28); hypopygium reaching apex ofgaster; last tergite exserted.Male: antennaltoruliaboutlevelwithlowest
short so that cereal plates are in apical one-third of eye margins; funicle 6-segmented, clothed in short
gaster; ovipositor(Fig. 113)frequently wellexserted. setae; clava 2-segmented (Fig. 32); phallobase with
parameres developed but short and inconspicuous,
Biology. Gregariousendoparasitoidsofcoccinellid
larvae (Coleoptera, Coccinellidae) feeding on sterno- digiti moderately long andeach with an apical hook.
rrhynch homopterans (Coccoidea, Aphidoidea and Female. Head in facial view about 1.2 x asbroadas
Psylloidea).Nothinghasbeenpublishedontheimma- long,inprofileabout2xaslongasdeepandanteriorly
ture stages. The adult parasitoids emerge from the more or less gradually and evenly curved, but most
prepupal stage of the hosts. There are one or two strongly so at level with top ofantennal scrobes. Eye
records of individuals having been reared from the with posterior margin more or less straight, about
larvae ofChrysomelidae and Bruchidae but these re- 1.5 x as long as broad, slightly longer than width of
quire confirmation. Records of species of the genus frontovertex, reaching or nearly reaching occipital
being parasitoids of scale insects (Homoptera, margin which is sharply carinate at least medially.
Coccoidea)arealmostcertainlyincorrect,thetruehost Malar space about 2/3 as long as an eye and with
probablybeingcoccinellidlarvaefeedingonthescale sulcusabsentorindicatedbyaslightchangeinsculp-
insects. ture.Frontovertexabout2/5headwidth;ocelliforming
Distribution. Cosmopolitan. an angle ofabout 90°, posterior ones much closerto
eyes than to occipital margin. Antennal scrobes very
Identification of species. The most comprehen- shallowandhardlymeetingdorsally,notreachinghalf
sive work revising species of the genus is that of way from toruli to anterior ocellus; antennal toruli
Timberlake (1919). A number of species have been separated from mouth margin by slightly more than
described more recently, mostly fromthe Old World, their own lengths and from each other by nearly the
and therefore the following works may be of use: same distance, their upper most margins about level
Trjapitzin (1989; Palaearctic); Noyes& Hayat (1984; with, or a little above lowest eye margins; clypeal
Indo-Pacific). marginwithatruncatemedianlobe(Fig. 114);mouth
Comments. Homalotylus is included here because opening about as wide as frontovertex or wider.
onespecies{hyperaspidisTimberlake)hasbeenreared Antennal scape subcylindrical to slightly flattened;
from larvae ofHyperaspis sp. feeding on triozids, or about 3 x as long as broad and not longerthan mini-
coccids inside the leafgall producedby Trioza sp. nr mumfrontovertexwidth;pedicelconical,about 1/3as
maritimaonAvicenniagerminans (Guanacaste Prov., long as scape and subequal in length to any funicle
GuanacasteNP,PlayaNaranja, 11.iii.1990,J.S.Noyes, segment which are subquadrate or distinctly longer
J. Cox). This species will be dealtwithelsewhere. than broad; clava apically rounded, 3-segmented and
about 1/3 as long as funicle; longitudinal sensilla on
allflagellarsegments;longestsetaeabouthalfaslong
CALDENCYRTUS gen. nov. aswidthofcorrespondingsegment.Frontovertexwith
shallow piliferous punctures, weakly metallic, the
Type species: Caldencyrtusmitchelli sp. nov. Gender sculpturemoderateandmoreorlesstransverselyelon-
masculine. gate; setae on frontovertex inconspicuous and each
shorter than diameter of anterior ocellus; eye appar-
Diagnostic characters. About 2-2.5 mm in ently naked, but with sparse extremely short setae.
length; body generally blackish with ametallic green Mandibleswithtwoteethandatruncation(Fig. 115);
lustre; mandibles with two teeth and a truncation; maxillarypalpi4-segmented,labialpalpi3-segmented.
occipital margin sharp, carinate; mouth opening at Thorax in side view moderately robust with meso-
least about as wide as frontovertex (Fig. 114); wings scutumandscutellummoderatelyconvex;mesopleuron
hyaline to slightly infuscate; forewing venation with slightlyenlargedposteriorlyandnearlytouchingbase
marginal veinthreeorfourtimesaslongasbroadand ofgaster so that propodeum appears tobe well-sepa-
subequal in lengthto stigmal andpostmarginal veins, rated from hind coxae. In dorsal view the posterior
the stigmal vein forming and angle ofabout45° with margin ofpronotum moderately concave; visiblepart
anteriorwingmargin(Fig.31);mesopleuronexpanded of mesoscutum slightly less than twice as broad as
posteriorly and more or less touching base ofgaster long, with notaular lines absent, its posterior margin
andthuspropodeumnotclearlyvisiblebetweenitand slightly to moderately produced; axillae meeting me-
base ofgasterin side view. Female: antennal toruli at dially or nearly so; scutellum with apex rounded,
level with lowest eye margins (Fig. 114); scape aboutas long asbroad; propodeum medially about 1/
subcylindrical; funicle 6-segmented; clava 3-seg- 7aslongasscutellumandwithsomeshallow,irregular
mented; gaster with hypopygium generally reaching sculpture and with a conspicuous group of silvery