A revision of the Indo-Pacific species of Ooencyrtus (Hymenoptera: Encyrtidae), parasitoids of the immature stages of economically important insect species (mainly Hemiptera and Lepidoptera)

136 Pages·1994·43.3 MB·

by  D-W Huang

1
Bull. nai. Hist. Mus. Lond. (Ent.)63(1):1-136 Issued23June 1994
A
revision of the Indo-Pacific
species of Ooencyrtus
(Hymenoptera: Encyrtidae),
parasitoids of the immature stages
of economically important insect
species (mainly Hemiptera and
Lepidoptera)
D.-W. HUANG & J.S. NOYES*
InstituteofZoology, ChineseAcademy ofSciences, Beijing, P.R. China
*DepartmentofEntomology, The Natural History Museum, Cromwell Road,
London SW75BD
CONTENTS
Synopsis 1
Introduction 2
BiologyofOoencyrtusspp 2
SynopsisofhostsofIndo-Pacificspecies 4
UseofOoencyrtus in biologicalcontrol 7
Depositories 7
Acknowledgements 7
SystematicSection 8
Ooencyrtus Ashmead 8
Commentson genericsynonymy 9
Genericdiagnosis 9
Systematicrelationshipswithin the genus 10
Identificationofspecies 10
Abbreviationsused in text 11
Keyto OrientalspeciesofOoencyrtus(females) 1
Reviewofspecies 16
Excludedspecies 83
References 84
Illustrations 88
Index toscientificnames 134
Synopsis. This revision treats 70 species of the genus Ooencyrtus found in the
Indo-Pacific region, including south China, but excluding Australia and New Zealand.
One new generic and 7 newspecificsynonymiesare proposed, 49species are described
as new 'and 11 lectotypes are designated. Each species is characterized by a diagnosis
and/orfull speciesdescription, itsknown distribution and hostsare reviewed, and a key
isprovidedto the females.
CorrespondencetoDrJ.S. Noyes
©TheNaturalHistoryMuseum. 1994
D.-W.HUANG ANDJ.S. NOYES
INTRODUCTION the solitary eggs of Sphingidae (Battisti et ai,
1988). At least one species is known to be a
gregarious parasitoid of the solitary eggs of but-
The genus Ooencyrtus is of particular interest terflies, e.g. papilionis (see p. 79). Two species
because it is widespread and species-rich in all are known to attack different stages of hosts in
geographic regions and its species parasitise different orders. Ooencyrtus kuvanae, has been
diverse hosts and hoststages. Manyspeciespara- recorded both as a primary solitary parasitoid of
sitise immature stages of pest species and thus theeggsofgypsymoth (Lymantriadispar) andas
have potential value as biological control agents. agregarious hyperparasitoid ofthe prepupal and
Some species also act as minorpests by parasitis- pupal stages of the moth's braconid primary
ing eggs of butterflies being reared for commer- parasitoid. Muesebeck & Dohanian (1927) found
cial purposes in Malaysia and Thailand. that female kuvanae from field-collected gypsy
This interest is reflected by the relatively large moth eggs would oviposit into the immature
amount of material of the genus that is received stages of Apanteles melanoscelus within their
by the identification services of the The Natural cocoons after they had emerged from the gypsy
History Museum and International Institute of moth caterpillar. Up to 17 individuals subse-
Entomology from the Indo-Pacific region. quently issuing from a single braconid host. The
Unfortunately, identification is difficult because same species of Ooencyrtus has also been
the taxonomy of the genus in this part of the recorded as a primary parasitoid of Anastatus
world is inadequate. This revision was under- bifasciatus Fonscolombe in gypsy moth eggs
taken as a first step towards improving this (Howard & Fiske, 1911), although Crossman
situation and it is perhaps not surprising that of (1925) stated that he knew ofonly one such case
the 70 species treated here, more than two-thirds in 14 years ofworking with these species. Ooen-
are described as new. cyrtussubmetallicus (Howard) is best known as a
primary parasitoid ofthe heteropterouseggs, but
it has also been recorded as a parasitoid of the
pupae of a chloropid dipteran (Legner & Bay,
BIOLOGY OF OOENCYRTUS
1965a,b).
Host range Ovary development
The majority of species are parasitoids of the Adult females emerge from the host with unde-
eggs of Hemiptera or Lepidoptera, but species veloped ovaries and the preoviposition period
are known that attack the prepupae of Lepi- mayvary from 1.5 to nearly four daysdepending
doptera, braconid primary parasitoids of cater- on temperature (Tracy & Nechols, 1988).
pillars (Lepidoptera), immature stages of
Dryinidae attacking auchenorrhynchous Mating
Homoptera, nymphal stages of Aphididae
(Homoptera), immature stages of Syrphidae Mating has been described in detail only for
(Diptera) or Coccinellidae (Coleoptera) feeding kuvanae (Brown, 1984). Females may mate sev-
on aphids, or the pupae of Chloropidae eraltimes, but iftheyparticipate in apost-mating
(Diptera). ritual they will mate only once. The post-mating
ritual involves the pair facing each other and the
Host specificity male touching the female with his antennae, fore
legs and head. The female touches the male with
The degree of host specificity is not known with her mouthparts at this time. Newly emerged,
anycertainty. Some species appear to act only as matingfemales may be swarmed overby malesif
solitary parasitoids of lepidopterous eggs laid in populations are large. Mating males appear to
batches, such as pinicolus which parasitises the cleanthe female withtheirmouthparts, they may
eggs of Lymantriidae and Lasiocampidae (see also try to mate with a female before her wings
Trjapitzin, 1989). Other species may be parasi- are fully expanded.
toids of the eggs of Heteroptera or Lepidoptera
whichare laidin batches, e.g.pityocampaewhich Host finding and oviposition
will parasitize the eggs of Lasiocampidae, Not-
odontidae (Lepidoptera) aswell as Coreidae and Evidence suggests that, in at least two species
Pentatomidae (Hemiptera), although the same [fecundusandpityocampae), female adult parasi-
species has also been recorded as a parasitoid of toids are attracted to chemicals or odours pro-
REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS
duced by the adult female host (Laraichi & number of instars is probably four or five since
Voegele, 1975; Battisti, 1989). This may be the these counts are based on the mandibles of the
result of the parasitoid being conditioned by exuviae of previous instars which remain
encountering suitable freshly laid eggs after the attached to the anal shield of the final instar
mass emergence of a suitable host species (Bat- larva.
tisti, 1989). Oviposition may take some time
dependingupon the experience ofthe parasitoid. Development time
In kuvanae experienced females may take only
five minutes to deposit an egg after the initial In both kuvanae and anasae development from
encounterwhilst inexperienced femalesmaytake eggtoadulttakesabout 18-35 daysdependingon
nearly twice as long (Lee & Lee, 1989), whilst in temperature (Muesebeck & Dohanian, 1927;
gregariousspeciesoviposition maytake nearly an Tracy & Nechols, 1987). In manii no develop-
hour per single host, e.g. johnsoni (Maple, ment of immatures occurs if the temperature is
1937). Host feeding, by the adult parasitoid, has over 32.5°C (Rahim et al., 1991). O kuvanae has
been observed in at least two species (kuvanae, 4 or 5 generations per year in more northerly
Lee & Lee, 1989; johnsoni. Maple, 1937) and is parts of the USA (Crossman, 1925) and up to 7
almost certainly common throughout the genus. generations per year in Italy (Prota, 1966) and
overwinters as an adult.
Fecundity
Adult longevity
The species of host egg can influence fecundity
(Laraichi, 1978a). Host density does not affect In manii longevity of both sexes decreases with
fecundity but lower host density increases the an increase in temperature, males will live 4 to
rate of superparasitism in solitary parasitoids 1.2 days with temperatures of 15-36°C whilst
(Laraichi, 1978b). The maximum overall fecun- females live from 10-1.7 days at the same tem-
dity may be as high as 200 (Crossman, 1925). In peratures. Ingeneral, longevityincreaseswith an
species that parasitise egg clusters the number of increase in humidity, with optimum at 50-70%
eggsdeposited bythe parasitoid will be limited to RH (Rahim etal., 1991). Thisis also reflected by
the number of mature eggs present in the ova- longevity and fecundity of the same species dur-
ries, in nezaraethis averages about 17 (Takasu & ing different seasons, the highest fecundity (37
Hirose, 1991). In kuvanaethis normally resultsin eggs/female) being noted in April and the lowest
a lower percentage parasitism of larger egg (0.8 eggs/female) being noted in January and
masses resulting from egg limitation (Weseloh, February (Yadav & Chaudhary, 1984). In some
1972; Williams et al., 1990). Percentage parasit- species day length may effect reproduction and
ism in kuvanae may be as high as 80%, but is longevity. For instance, in kuvanae longer day
generally around 10-40%. The substrate on length results in more progeny but lower longev-
which the host eggs are laid may affect rate of ity (Weseloh, 1986).
parasitism. In kuvanae, the eggsofitshost on red
maple (smooth bark) have a higher rate of para- Sex ratio and sex determination
sitism than those on oak (rough bark) (Bellinger
etal., 1988). The sex ratio varies from about to 1:1 to 4:1 in
favour of females (johnsoni Maple, 1937). In
Immature stages anasae the proportion of females increases with
number of hosts parasitized (Tracy & Nechols,
After oviposition the egg stalk remains protrud- 1987). Sex ratio may also be influenced by tem-
ing through the chorion of the host egg. The perature. For instance, in fecundus (Laraichi,
larva remainsattached to itseggshell forthe first 1978c) all progeny are female if the ovipositing
three instars and is metapneustic (Maple, 1937), female is subjected to temperatures of 30°C but
being able to utilise atmospheric air directly all progeny are male if the ovipositing female is
through the protruding part of the egg (Maple, subjected to a temperature of 35°C (Laraichi,
1937). The number of larval instars recorded 1978c). In O. submetallicus similar temperatures
varies from three (Gerling et al., 1976 in trin- produce similar results with hermaphrodite
idadensis; Matteson, 1981 in utetheisae; Cross- mosaics being produced if the developing prog-
man, 1925 in kuvanae), four (Maple, 1937 in eny are subjected to intermediate temperatures
johnsoni; Laraichi, 1977 infecundus, nigerrimus (Wilson, 1962; Wilson & Woolcock, 1960). This
and telenomicida) to five (Parker, 1933 in kuva- suggests that the sex of the offspring may be
nae;Takasu& Hirose, 1989innezarae). The true under the control of a microorganism similar to
D.-W.HUANG ANDJ.S. NOYES
thatfoundin some speciesof Trichogramma (see resulted in a higher proportion offemales, possi-
Stouthamer et al, 1990; Stouthamer, 1990, bly because of higher male mortality. Seasonal
1991). However, Kamay (in Brown, 1984) fluctuations of temperature influence the sex
reported that if developing kuvanae were ratio oimanii (Yadav & Chaudhary, 1984).
exposed to temperatures of around 35°C this
SYNOPSIS OF HOSTS OF INDO-PACIFIC SPECIES
[] extralimital records
* probably incorrect host association
laboratory reared
!
(H) hyperparasitoid
All reared from eggs except those prefixed by (L) -from larvae, and (P)-from prepupae or pupae.
HOST PARASITOID
(Ooencyrtussp.)
Unknowneggs
COLEOPTERA
Chrysomelidae
PlesispareicheiChapius pindarus
Podontiaquatuordecimpunctata (Linnaeus) corbetti
Podontiaquatuordecimpunctata (Linnaeus) podontiae
Coccinellidae
(P)indet. guamensis
DIPTERA
Syrphidae
(P)Allograptaexotica (Wiedemann) guamensis
(L)Dideopsispura (Curran) guamensis
(L)[Ischiodonscutellaris (Fabricius) guamensis]
(L)[IschiodonaegyptiusWied (Wiedemann) guamensis]
(L)ParagusauritusStuckenberg guamensis
HEMIPTERA
indetfamily ferrierei
Heteroptera
Alydidae
Leptocorisasp. utetheisae
LeptocorisaacutaThunberg utetheisae
[Mirperusjaculus (Thunberg) utetheisae]
Piezodorushybneri(Fabricius) utetheisae
Riptortussp. cybele
Riptortussp. utetheisae
[Riptortusdentipes(Fabricius) utetheisae]
Coreidae
Amblypeltasp. caurus
AmblypeltacocophagaChina ilion
AmblypeltalutescensDistant caurus
AmblypeltapapuensisBrown utetheisae
caurus
[Anoplocnemiscurvipes(Fabricius) utetheisae]
REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS
[ClavigrallaelongataSignoret utetheisae]
[ClavigrallatomentosicollisStal utetheisae]
DasynuskalshoveniBlote icarus
DasynuspiperisChina utetheisae
[GonocerusacutangulatusGoeze \telenomicida]
[GonocerusjuniperiHerich-Schaeffer telenomicida]
Mictisprofana (Fabricius) utetheisae
[Pseudotheraptus wayiBrown utetheisae]
Pentatomidae
indet. lucens
[Brachynemagermarii(Kolenati) telenomicida]
[Dolycorispenicillatus Horvath telenomicida]
Eocantheconafurcellata (Wolff) iulus
Nezara viridula (Linnaeus) utetheisae
Plataspidae
Brachyplatyspacificus (Dallas) pacificus
Cratoplatyssp. ceres
Scutelleridae
[EurygasterintegricepsPuton telenomicida]
[Aeliasp. telenomicida]
Tessaratomidae
Tessaratomapapillosa (Drury) phongi
Tessaratomajavanica (Thunberg) phongi
Pycanumponderosum Stal phongi
Homoptera
indet. family midas
Lophopidae
Pyrillaspp. manii
Pyrillaperpusilla Walker manii
Aphididae
(L)indet. pallidipes *
LEPIDOPTERA
indet. phongi*
indet. javanicus
indet. utetheisae
(P)indet. leafminingfamily ooii
Agonoxenidae
(L)Agonoxenapyrogramma Meyrick shakespearei
Arctiidae
[Utetheisapulchella (Linnaeus) utetheisae
Bombycidae
Rondotiamenciana Moore hercle
Crambidae
ChiloterenellusPagenstecher papilionis
Danaidae
Danauschrysippus (Linnaeus) papilionis
Euploeacore(Cramer) papilionis
Tirumalalimniace(Cramer) papilionis
Epipyropidae
(P)Epiricaniamelanoleuca (Fletcher)
Gracillariidae
(P)AcrocercopsglobuliferaMeyrick oott
(P)Conopomorphacramerella(Snellen) ooii
D.-W.HUANG ANDJ.S. NOYES
Heliconiidae
Heliconiuscharitonius (Linnaeus) papilionis
Hesperiidae
Erionotathrax(Linnaeus) pallidipes
Hasorasp. papilionis
Lasiocampidae
Dendrolimussp. endymion
DendrolimuskikuchiiMatsumura endymion
Dendrolimusspectabilis Butler kuvanae
Malacosomaamericana \kuvanae
Malacosomaneustriatartacea Motschulsky kuvanae
Taragamarepanda Hubner Uelenomicida
Lymantriidae
Aroacometaris Butler papilionis
Euproctischrysorrhoea (Linnaeus) \kuvanae
HemerocampaleucostigmaAbbot & Smith \kuvanae
HemerocampadefinataPackard Ikuvanae
[Lymantriadispar(Linnaeus) kuvanae]
[Lymantriafumida Butler kuvanae]
LymantriaxylinaSwinhoe Ikuvanae
[NygmiaphaeorrhoeaDonovan kuvanae]
[StilpnotiasalicisLinnaeus kuvanae]
Noctuidae
AchaeaJanata (Linnaeus) lucens
(L)Exelastisatomosa (Walsingham) guamensis
Othreisfullonia (Clerck) crassulus
Othreisfullonia (Clerck) papilionis
(L)Mythimnaunipuncta (Haworth) guamensis
Notodontidae
Closteracupreata (Butler) lucina
Stauropuslichenina Butler boreas
Turnacaacuta (Walker) macula
Nymphalidae
Ariadneariadne(Linnaeus) papilionis
Caligomemnon (Felder) pallidipes
Hypolimnasbolina Linnaeus papilionis
Kallimasp. papilionis
Phalantaphalantha (Drury) papilionis
Junonialemonias (Linnaeus) papilionis
TanaeciajuliiBougainville papilionis
Papilionidae
TroidesHelenaLinnaeus papilionis
Papiliosp. papilionis
Papiliosp. plautus
PapilioaegeusDonovan papilionis
Papilioagamemnon (Linnaeus) papilionis
Papiliocitri(?lapsusfordemoleus plautus
Papiliodemoleus Linnaeus papilionis
PapiliohelenusLinnaeus papilionis
Papiliomemnon Linnaeus paplionis
Papiliopolytes Linnaeus papilionis
PapiliorumanzoviaEschscholtz papilionis
Pieridae
(L)Deliassp. larvarum
Euremasp. larvarum
Hebomoiaglaucippe(Linnaeus) hera
Saturniidae
REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS
AttacusatlasFabricius phoebi
CallosamiaprometheaDrury \kuvanae
Criculasp. javanicus
CriculaelaeziaJordan dione
[Eriogynapyretorum Westwood kuvanae]
Hemileucamaia Drury Ikuvanae
HemileucaoliviaeChurchill Ikuvanae
Sphingidae
indet. endymion
AmorphapopuliaustantiStaudinger Uelenomicida
Cephonodeshylas (Linnaeus) papilionis
IGnathothlibaserotuseras (Boisduval) sphingidarum
Hippotioncelerio (Linnaeus) crassulus
NEUROPTERA
Myrmeleontidae
indet.
USE OF OOENCYRTUS IN IZAS Instiute of Zoology, Chinese Academy of
Sciences, Beijing, PR. China
BIOLOGICAL CONTROL MARI Malaysian Agricultural Research and
Development Institute, Kuala Lumpur,
Malaysia
Although species of Ooencyrtus undoubtedly MNHN Museum National d'Histoire Naturelle,
play an important role in the regulation of the
Paris, France
populations of many insect species worldwide, MZB Museum Zoologicum Bogoriense, Bogor,
they have not proven to be ofgreat value in pest Indonesia
control. Their use in classical biological control ORSTOM Office de la Recherche Scientifique et
programmes is summarised in Table 1. Perhaps Technique Outre-Mer, Paris, France
thebest documented example is that ofOoencyr- PPRI Plant Protection Research Institute, Preto-
tus kuvanae for the control of the gypsy moth ria, South Africa
(Lymantria dispar) in North America and OMB Queensland Museum, Brisbane, Australia
Europe. Brown (1984) comments that although TAMU Texas A&M University,Texas, USA
affording some control, the species is unlikely to USNM United States National Museum, Washing-
be ofmuch benefit by itself, although it probably ton, D.C., USA
causes a post outbreak collapse which in turn ZAMU ZoologyDepartment,AligarhMuslimUni-
increases the length of the interval between out- versity, Aligarh, India
breaks. ZISP Zoological Institute, St Petersberg, Russia
A further species, Ooencyrtus fecundus Fer-
riere & Voegele, may be of some benefit ifused
by means of inundative releases against het- ACKNOWLEDGEMENTS
eropterouspestsofwheat inNorth Africa (Larai-
chi & Voegele, 1975).
We thank The Royal Society which provided the
senior author the opportunity to study at The
Natural History Museum in London. Many
DEPOSITORIES thanks to Mr David FosterofThe Royal Society,
who gave the senior author consistent help dur-
BMNH The Natural History Museum, London, ing his stay in UK. The Natural History Museum
England made its collections, facilities and libraries avail-
BPBM Bernice P. BishopMuseum, Hawaii able to us for the project. The senior author's
CNC Canadian National Collection, Ottawa, host institution, Institute of Zoology, Chinese
Canada AcademyofSciencespermittedhimoneyearoff.
IARI IndianAgricultureResearchInstitute,New We also thank Nigel Wyatt, Jeremy Holloway,
Delhi, India Gary Stonedahl, Phil Ackery, Mike Shaffer and
IEE Institito di Entomologfa Espanol, Madrid, Mick Webb for checking host names, and thanks
Spain especially to John LaSalle for his valuable com-
IRSN Institute Royal des Sciences Naturelles de ments concerning the manuscript. Finally we are
Belgique, Brussels, Belgium grateful to Dr M. Schauff (USNM), Dr G.
8 D.-W.HUANG AND J.S. NOYES
Table 1 Asummaryofthe useofOoencyrtusspp. inclassicalbiologicalcontrolprogrammesworldwide
(Abbreviations: NE-notestablished; NR-not released; ?-nosubsequentinformation; SC-successful
control; P-partialcontrol; E-establishedbutnofurtherinformation available;NC-establishedbutno
significantcontrolachieved).
Targetpestspecies Introduced Ooencyrtussp.;country; year; result; source
HEMIPTERA
Amblypeltacocophaga -sp.;Solomon Is; 1937-38; NE;Phillips (1941)
-malayensis(?= utetheisae, misident.); Solomon Is; 1937-38; NE; Phillips(1941)
Amblypeltatheobromae -malayensis(?= utetheisae, misident.); PapuaNewGuinea; 71974; ?; Young
(1982)
Anasatristis -malayensis(?= utetheisae, misident.); USA (Massachusetts); 1981; ?; Coulson et
al. (1988)
Eurygasterintegriceps -fecundus; USSR; ?; NR; Izhevskiy(1988)
-nigerrimus; USSR; ?; NR; Izhevskiy(1988)
-telenomicida; USSR; ?; NR; Izhevskiy(1988)
Murgantiahistrionica -johnsoni; Bermuda; 1953; NE; Bennett& Hughes (1959)
-johnsoni; Hawaii; 1940; E; Clausen in Clausen (1978)
Nezara viridula -johnsoni; Australia; 1953; NR; Wilson (1960)
-malayensis(?= utetheisae, misident.); USA (Massachusetts); 1981; ?; Coulsonet
al. (1988)
-submetallicus; Australia; 1952-57; NE; Wilson (1960)
-submetallicus; Hawaii; 1962; NE; Davis(1964), Waterhouse & Norris(1987)
-submetallicus; USA (Florida); 1973; ?;CIBC(1974)
-trinidadensis; Hawaii; 1962; NE; Davis(1964), Waterhouse & Norris (1987)
Pseudotheraptus wayi -sp. (?= utetheisae); Zanzibar; 1959; ?; Greathead (1971)
-sp. (?= utetheisae); Kenya; 1959; ?; Greathead (1971)
Soybeanstinkbugs -nezarae; Brazil; 1983-1985; ?; Kobayashi & Cosenza (1987)
LEP1DOPTERA
Ascotisselenaria -ennomophagus; Israel; 1977; NR; Wysoki (1979)
Calpodesethlius -sp. (= calpodicus); Bermuda; 1953, 1962-63; NC; Cock (1985)
-sp. (= calpodicus); StVincent; 1950-51; E; Cock (1985)
Conopomorphacramerella -sp. (= ooii); Malaysia (Sabah); 1987; NE; CIBC (1988, 1989, 1990)
Erionotathrax -pallidipes; Hawaii; 1973; SC; Mau, etal. (1980)
-pallidipes; Mauritius: 1971-72; P; Waterhouse & Norris (1989)
Hemerocampaleucostigma -kuvanae; USA; 1917, 1921; NE; Dowden (1962)
Hemileucaoliviae -kuvanae; USA (NewMexico); 1913-16;NE; Clausen (1956)
Lymantriadispar -kuvanae; Algeria; 1925-26, 1931; E; Lepigre (1932), Clausen (1978)
-kuvanae; Canada (Ontario); 1976; E; Brown (1984)
-kuvanae; Czechoslovakia; 1922; E; Clausen (1978)
-kuvanae; Morocco; 1924-26; E; Brown (1984), Clausen (1978)
-kuvanae; Portugal; 1932; E; Brown (1984), Clausen (1978)
-kuvanae; Spain; 1923-27; E; Brown (1984), Clausen (1978)
-kuvanae; USA; 1908-28, 1967-1971, 1981; E; Brown (1984), Peck (1963),
Clausen (1956, 1978), Coulsonetal. (1988)
-kuvanae; CIS; 1987; ?; Volkov& Mirohova (1990)
-kuvanae; Yugoslavia; 71960; ?; Brown (1984)
Othreisfullonia -sp. (?=papilionis); American Samoa; ?; ?; Waterhouse & Norris (1987)
-sp. (?=papilionis); WesternSamoa; ?; ?; Waterhouse & Norris(1987)
Nishida (BPBM), DrJ.-L. Nieves Aldrey (IEE), SYSTEMATIC SECTION
Dr I. Trjapitzin (ZISP), Dr G.L. Prinsloo
(PPRI), Dr G. Gibson (CNC), Dr P. Dessart
(IRSN) andMrE.C. Dahms (QMB) forthe loan OOENCYRTUS Ashmead
orgift ofmaterial.
Ooencyrtus Ashmead, 1900: 381. Type species:
Encyrtus clisiocampae Ashmead, by original
designation.
EchthrodryinusPerkins, 1906: 252. Type species:
REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS
Echthrodryinus destructor Perkins, by mono- from their gypsy moth caterpillar host (see
typy. Synonymy with Ooencyrtus by Noyes & below). Recent work (Noyes, 1985 and Prinsloo,
Hayat, 1984. 1987) has shown that the host range of Ooencyr-
Ectopiognatha Perkins, 1906: 254. Type species: tus spp. is much more diverse than had previ-
Ectopiognatha minor Perkins, by designation ously been thought, many species being noted as
ofGahan & Fagan, 1923: 49. Syn.nov. parasitic on nymphs of Aphididae (Hemiptera)
Schedius Howard, 1910: 2. Type species: Sche- and Syrphidae (Diptera), even to the extent
dius kuvanae Howard, by original designation. where specimens that attack dipterous or
Synonymy with Ooencyrtus by Ferriere, 1931. coleopterous larvae are morphologically indistin-
Tetracnemella Girault, 1915: 170. Type species: guishable from those that attack heteropterous
Tetracnemella australiensis Girault, by original eggs (see Noyes, 1985). In view of this, we are
designation. Synonymy with Ooencyrtus by continuing to treat Echthrodryinus as synony-
Noyes & Hayat, 1984. mous with Ooencyrtus.
Xesmatia Timberlake, 1920: 424. Type species:
Xesmatiaflavipes Timberlake, by original des- Generic diagnosis
ignation. Synonymy with Ooencyrtusby Noyes
& Hayat, 1984. In the most comprehensive classification of the
Pseudolitomastix Risbec, 1954: 1068. Type spe- Encyrtidae currently available, Trjapitzin (1973,
cies: Litomastixcreona Risbec. Synonymywith 1989) includes Ooencyrtus in the Encyrtinae,
Ooencyrtus by Annecke & Mynhardt, 1973. tribe Microteryini, subtribe Ooencyrtina. Unfor-
tunately, this classification does not provide any
Comments on generic synonymy meaningful diagnoses for the tribes or subtribes.
We are therefore hopeful that the following
Species previously combined with the genus diagnosis will enable species to be assigned cor-
Ectopiognathacan be separatedfrom Ooencyrtus rectly to Ooencyrtus:
by the quadridentate mandibles and a broadened
Robust, squat species, never conspicuously
andflattened scape. However, one ofthe species
slender and elongate; thorax with posterior
included in this revision (leander sp.n.) is margin ofmesoscutum weakly to strongly con-
extremely close in general structure to both spe- vex medially and overlying axillae centrally so
cies included in Ectopiognatha except that it has that when thorax is in normal resting position
acylindrical scape. In addition, several described axillae appear to be widely separated;
species of Ooencyrtus have a scape which is mesopleuron posteriorly expanded so that it
distinctly broadened. The biology of Ectopiog- touches base ofgaster, or nearly so, and com-
nathaspp. (parasitoids ofthe eggsofHemiptera) pletely conceals the metapleuron and propo-
falls within the range ofthat found in Ooencyrtus deum from lateral view immediately above the
(see below) and, therefore, we have no hesita- hind coxae; forewing with marginal vein punc-
tiform or not much longer than broad,
tion in treating the two genera as synonymous.
Noyes and Hayat (1984) treated Echthrodryi- Species of Ooencyrtus can be most easily con-
nus as a synonym of Ooencyrtus whilst Gordh & fused with Trichomasthus, Helegonatopus and
Trjapitzin (1978) and laterTrjapitzin (1989) have Psyllaephagus. Both Helegonatopus and Psyl-
treated it as valid. It is not possible to separate laephagus have the mesopleuron normal, not
the two genera on morphological grounds and expanded posteriorly and not touching the base
the only possible basis for continuing to treat ofthe gaster so that in lateral view the metapleu-
them as distinct is that the included species have ron and propodeum are not obscured and touch
different biologies. Ooencyrtus could be the hind coxae. Species of Trichomasthus are
restricted to primary parasitoids (or perhaps generally much larger, usually being about 1.5
mm
hyperparasitoids) of the eggs of various insects, long, the marginal vein of the forewing is
whilst Echthrodryinus could include primary normallyseveral times longerthan broad, andall
parasitoids or hyperparasitoids of the larvae or species are parasitoids of scale insects (Coc-
prepupal stages of certain holometabolous coidea).
insects, e.g. Dryinidae, Braconidae and Gracilla- Superficially, the placement of the genus
riidae. However, one species, Ooencyrtus kuva- within the tribe Microteryini may appear some-
nae, has been recorded both as a primary what questionable, especially on biological
parasitoid of the eggs ofgypsy moth (Lymantria grounds. Morphologically, it ispossibly to see an
dispar) and as a hyperparasitoid of the prepupal evolutionary trend from a Microterys-Mke or
and pupal stages of the braconid Apanteles mel- Trichomasthus-Wke ancestor, some species of
anoscelus in their cocoons after they emerge Microterysand Trichomasthusbeingquitesimilar
10 D.-W.HUANG ANDJ.S. NOYES
structurally and, as stated above, it can be some- the following characters especially helpful in
times quite difficult to separate Ooencyrtus from identifyingspecies:
pTlreiscihoommoarspthhiucs.hoIsftitassisocihaytpiootnhefsoirzetdhethtartibethies Mandibles - several basic types, a) with one
scale insects (Homoptera; Coccoidea) (see tooth and a broad straight, sometimes minutely
Trjapitzin, 1989) then it would be possible to denticulatetruncation (Figs 10, 18, 139, 188, 204,
envisage a switch from parasitising insects with 333, etc.), b) withone tooth and a broad, slightly
hard a scale covering and a host immune system, convex, minutely denticulate truncation (Figs
to parasitising the eggs of other insects which 303), c) one tooth and a distinctly emarginate
haveahardoutershell andnoimmunesystem. A truncation, or with two teeth and a truncation
further switch to hyperparasitism of Braconidae (Figs 75, 143, 150, etc.), d) tridentate (Fig. 69,
or Dryinidae, or parasitism of the immature 91, 268, etc.), all teeth subequal, e) one or two
stages of holometabolous insects (Diptera and teethandanoblique minutelydenticulatetrunca-
Lepidoptera) is much more difficult to envisage, tion, (Fig. 240), f) three acute, unequal teeth
but at least two species are known to parasitise (Fig. 176), or g) four teeth (Fig. 46).
both hemipterous or lepidopterous eggs and the Clava - a) clava with apex rounded and sensory
prepupae or pupae of holometabolous insects, area at extreme apex only (Figs 42, 47, 51, etc.)
i.e. kuvanae(Muesebeck& Dohanian, 1927) and b) clava with sutures transverse and parallel,
submetallicus (see Noyes, 1985). apex obliquely truncate and sensory area
enlarged (Figs 9, 104, 196, etc.), or c) clava with
sutures oblique, apex obliquely truncate and
Systematic relationships within the
sensory area enlarged (Figs 16, 22, 28, 193, etc).
genus
Eyes - a) conspicuously hairy, or b) almost
The relationships within Ooencyrtus are unclear naked.
and we do not attempt to provide any sort of Ocelli - relative distance of ocelli from occipital
formal classificatory framework within the genus margin in terms oftheirown diameters.
by proposing subgeneric categories. However, Antennal toruli- relative distance below eyes or
probable assemblages of closely related species above mouth margin in terms of their own
within the genus are highlighted in the comments lengths.
Interantennalprominence - coloration purple or
sections ofthe appropriate species.
metallic green or blue green, etc.
Scutellum - relative depth and type of sculpture
Identification ofspecies
in relation to mesoscutum.
The following works should be consulted as aids Forewing - relative density and distribution of
to the identification of species from outside the setae on both dorsal and ventral surfacesofbasal
Oriental region: Peck (1963) and Gordh (1979) cell and on ventral surface of costal cell; linea
for North America; Noyes (1985) and De Santis clava closed or open; relative length of marginal
(1988) for South America; Trjapitzin (1989) for and postmarginal veins in relation to stigmal.
the Palaearctic; Prinsloo (1987) for Africa. Legs- the coloration of coxae, femora and tibia
Previous work dealing with the species within seems to be reliable although there may be a
the Indo-Pacific area has not much value, either slight amount ofvariation in the intensity of any
becauseofinadequatecoverage ofthespecies, or brownareason the femora and tibia. The colora-
because it relies on poor characters whilst over- tion of the fore and hind coxae may vary from
looking important diagnostic features. The only yellow to largely brown in some species.
key available (Trjapitzin et ah, 1978) is based Gaster - the relative length may be unreliable
almost entirely on inadequate original descrip- because it may depend on the preservation tech-
tions and is therefore oflittle use. niques used. In air dried specimens the gaster
Although we describe males of each species collapses and may thus be relatively shorter than
when they are available, they are excluded from in Critical Point Dried specimens where the
the key to species. Males are often difficult to gaster remains inflated (see Gordh & Hall,
identify because they do not present many fea- 1976). The shape of the last tergite (apically
tures and are unknown for most of the species rounded, truncate or medially invaginated) may
included here. However, the presence of males be useful as well as the shape ofthe hypopygium
inarearedseriesmayhelptoconfirmthe separa- (rectangular, triangular, with long or short ante-
tion ofsome closely related species, e.g. sphingi- rior lateral projection, relative size of posterior
darum andpapilionis. incision, distribution and relative density of
During the course of this work we have found setae).