The chalcidoid parasites (Hymenoptera, chalcidoidae) of economically important Liriomyza species (Diptera, agromyzidae) in North America

21 Pages·1991·9.6 MB·

by  J Lasalle

PROC. ENTOMOL. SOC. WASH.
93(3), 1991, pp. 571-591
THE CHALCIDOID PARASITES (HYMENOPTERA, CHALCIDOIDAE) OF
ECOMONICALLY IMPORTANT LIRIOMYZA SPECIES
(DIPTERA, AGROMYZIDAE) IN NORTH AMERICA
John LaSalle* and Michael P. Parrella
(JL) Department ofEntomology, University ofCalifornia, Riverside, California 92521:
(MP?) Department ofEntomology, University ofCalifornia, Davis, California 95616.
v4/75/rac/.—Twenty-four species ofchalcidoid parasites ofthe five economically impor-
tant species ofLihomyza in North America are reviewed, plus four species which may
A
proveto parasitizetheseLiriomyzaspecies. key ispresentedtodistinguishthese species,
and each species is discussed, with host and distributional information given. Species
names are corrected to reflect currently applied nomenclature, and a complete list ofall
names that have been used for these parasites is given, indicating currently recognized
names and all other names used for any ofthese species.
Key Words: Agromyzidae, Chalcidoidea, Eulophidae, leafminer, Liriomyza, parasite,
Pteromalidae
Five speciesoiLiriomyza (brassicae, hui- key currently encompasses the entire chal-
dobrensis, sativae, trifoliearum, trifolii) are cid fauna ofLiriomyza parasites. The pres-
considered to be of economic importance entpapertreatsallspeciesofchalcidoidpar-
in North America (Spencer 1973, Parrella asites recorded from of these leafminer
1982). Confusion concerningthe identity of species in North America. An additional
these Liriomyza species has been rectified four species, Diglyphus isaea, D. earlylei,
by several recent papers (Spencer 1981, Chrysocharis liriomyzae, C. ignota, are in-
Spencer and Steyskal 1986, Parrella 1982). cluded in this paper because they are either
The purpose ofthe present paper is to ex- very closely related to included species and
amine the chalcidoid parasite complex of may be confused with them, or they are
these leafminers, present a means for the suspectedtobeparasitesoftheseLiriomyza
identification of the parasites, and to up- species or become so in the future. A key is
grade the nomenclature ofthe parasites in presented to distinguish these 28 species,
light ofseveral recent papers on the system- and each species is discussed in the text,
atics ofthese groups. with diagnostic characters, and host and
Alargeparasitecomplexisassociatedwith distributional information given.
theseleafminers; 24 speciesofChalcidoidea Table 1 gives all names used for the leaf-
are now known from North America. Al- miner parasites in the North American lit-
though keys to identify many of the indi- eratureand indicates: currentlyvalid names;
vidual genera and species are available, no the currently recognized names for species
which are no longer consider valid; and all
*Current mailing address: Dr. John LaSalle, CAB generic combinations (including the valid
International,InstituteofEntomology,56Queen'sGate, one) used for any ofthe specific names.
London, SW7 5JR, UNITED KINGDOM. This paper is not intended to be revi-
572 PROCEEDINGS OFTHE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 1. List ofall names used for Nearctic Lir- sionary in nature, but rather a means of
iomyza parasites. Currently valid names in boldface. identifying known Liriomyza parasites.
Genericnamesinparenthesesareothergenericnames Many ofthe genera treated here are badly
underwhichaspecieshasbeentreatedintheliterature. in need of work, and there are problems
which cannot be resolved without detailed
acantha(Walker), Pediobius
aenea (Walker), Halticoptera [misidentification ofH. revisionsary studies. It is out of the scope
circulus(Walker)] ofthis paperto attempt to solve all ofthese
agromyzae (Crawford), Chrysonotomyia (Derostenus) problems, howeverwetrytopointoutprob-
= C. pimctivenths (Crawford) [although see discus- lem areas when we are aware ofthem. The
sion in text under C. pimctivenths] key, discussions, and generic diagnoses are
ainslieiCrawford, Chrysocharis
basedon femalesonly, andarenot intended
americanumGirault, Zagrammosoma
arizonensis (Crawford), Chrysonotomyia (Derostenus) forgeneral use, only forspecies reared from
= C. punctiventris(Crawford) Liriomyza. Complete revisions ofthe var-
begin!(Ashmead), Diglyphus (Diaulinus, Solenotus) ious genera are referred to in the text and
callichromaCrawford, Diaulinopsis thereadercanconsulttheseifthereisdoubt
cinctipennis Ashmead, Closterocerus
about species identifications.
circulus(Walker), Halticoptera
crassiscapus(Thomson), Chrysocharis Tables which summarize parasite/L/>/o-
diastatae(Howard), Chrysonotmyia (Derostenus) myza interactions have previously been
flavipes(Ashmead), Pnigalio presented (Minkenberg and van Lenteren
ffloarvmoovisraidi(sWeCsrtawwofoodr)d,, CCihrrryosspoinloutsomyia (Achrysochar- 1986,Johnsonand Hara 1987), andwehave
condensedand modifiedthesetablesto em-
ella)
fullawayi (Crawford), Chrysonotomyia (Derostenus, phasize the chalcidoid fauna associated with
Achrysocharella) = C.formosa (Westwood) the five most important Liriomyza species
giraultiYoshimoto, Chrysocharis in North America (Table 2). Records for
ignota Hansson, Chrysocharis parasites not determined to species are
intermedins(Girault), Diglyphus(Solenotus)
omitted. Recent reviews o{Liriomyza con-
isaea(Walker), Diglyphus
lineaticeps (Girault), Zagrammosoma (Mirzagram- tainmorecompletelistsofreferencestopar-
mosoma) asite biology, behavior, and effectiveness
liriomyzaeDelucchi, Chrysocharis (Parrella and Robb 1985, Minkenberg and
mallochi Gahan, Chrysocharis = C. crassiscapus van Lenteren 1986).
(Thomson)
mirum Girault, Zagrammosoma
multilineatum (Ashmead), Zagrammosoma Parasites
oscinidis Ashmead, Chrysocharis
parksiCarwford,Chrysocharis=C.oscinidisAshmead The Chalcidoidea contains 24 species of
patellana(Dalman),Halticoptera [misidentificationof known parasites ofthe 5 economically im-
H. circuhis(Walker)] portant Liriomyza species in the Nearctic
petiolata (Girault), Chrysocharis = C. girauhi Yoshi- region. Thesearepredominantlyinthefam-
moto
ily Eulophidae, with a single Pteromalidae,
pictipes(Crawford), Chrysonotomyia(Derostenus) Halticoptera circulus(Walker). The euloph-
pulchripes(Crawford), Diglyphus (Solenotus)
punctiventris (Crawford), Chrysonotomyia (Derosten- id parasites o{Liriomyzaare in the subfam-
us. Achrysocharella) ilies Entedoninae and Eulophinae (tribes
rugosithorax Crawford, Pediobius (Pleurotropis) = P. Eulophini and Elachertini). Oatman and
acantha(Walker) Kennedy (1976) list a Tetrastichus sp. (Tet-
trifasciatus Westwood, Closterocerus rastichinae), however they do not supply a
utahensisCrawford, Closterocerus
variipes (Crawford), Chrysonotomyia (Derostenus. species name for this parasite.
Achrysocharella) = C.formosa (Westwood) General host associations and distribu-
viridis Provancher, Chrysocharis = C. oscinidis Ash- tion are given for all parasites. This infor-
mead
mation is taken from catalogues of North
websteri(Crawford), Diglyphus (Solenotus)
American Chalcidoidea (Peck 1963), North
m
VOLUME 93, NUMBER 3 573
Table 2. Host associations between Nearctic Liriomyza species and their parasites condensed from Min-
kenberg& van Lentem (1986) and Johnson & Hara (1987).
brassicae hiudobrcnsis tnfoliearuin
Diglyphusbegini + + +
Diglyphus intermedins + +
Diglyphuspulchripes
Diglyphus websteri
Pnigalioflavipes
Chrysocharisainsliei + +
Chrysocharisgiraulti
Chrysochariscrassiscapus +
Chrysocharisoscinidis + +
Chrysonotomyia diastatae
Chrysonotomyiaformosa
Chrysonotomyiapictipes ?*
Chrysonotomyiapunctiventris + + +
Closteroceruscinctipennis
Closterocerustrifasciatus
Closterocerus utahensis
Pediobiusacantha
Zagrammasoma americanu
Zagrammasoma lineaticeps
Zagrammasoma mirum
Zagrammasoma multilineatum
Cirrospilusflavoviridis
Diaulinopsiscallichroma +
Halticoptera circulus + + +
"Oatman&Platner(1969)recordedthespeciesDerostenuspunctipesCrawford.Thereisnosuchspeciesname
and it may have been that they meant to referto this species.
American Hymenoptera (Burks 1979a, b) l'- Tarsi 4-segmented. Antennawith 2^ fumc-
and.r.ev.isions ofthe various genera (which "i^"" segments. (Euiophidae) .... l
arehsted undJereac,hgenus)X,andJ .t.huesework1s 2. Scutellumwith2setae(1 pair)(Figs.4-5,21-
,2). Submargmai vein with 2 dorsal setae
shouldbechecked ifmorecompleterecords (pjg^ 6_8 \i_i2). (Entedoninae) 3
are desired. Complete synonymy lists are 2'. Scuteiiumwithatleast4setae(2pairs)(Figs.
not given for the genera and species ofpar- 42^3). Submarginal vein with more than 2
asites, although all names used in the lit- do^-^^l ^^^^^ (^'g^- ^-^^^ 13-18). (Euioph-
erature on LJi.n.omyza parasi-t^es are given in inae) 15
3 p^^p^^eumwithcompleteparallelplicae,and
thetextandinTable 1. Completesynonymy ^^^mediancarinawhichdivergeposteriorly
lists can be found in the above mentioned (Fig. i). (genus Pediobius)
cataloguesandunderthevariousgenericre- Pediobiusacantha (Walker)
3'. Propodeumwithoutplicae;mediancarina(if
Visions.
present) not posteriorlybifurcate (Figs. 2-3,
25-28) 4
Key TO Species of 4. Forewingwith2or3distinct,althoughsome-
Liriomyza Parasites ^'"^^^ *'^'"^' transverse bands (Fig^ 6). Scape
Based on Females at least slightly flattened and produced ven-
trally (Figs. 29, 30). Funicle usually slightly
Tarsi 5-segmented.Antennawith 6 funicular to distinctly flattened (Fig. 30). (genus Clos-
segments. (Pteromalidae). Notaulicomplete. terocerus) 5
Petioledistinct,althoughmaybesmall. Male 4'. Forewing without dark bands, at most with
withmaxillarypalpienlarged, yellow, (genus a single slightly darkened area beneath stig-
Halticoptera) ....Halticoptera circulus(Walker) mal vein. Scape and funicle not flattened . . 7
574 PROCEEDINGS OFTHE ENTOMOLOGICAL SOCIETY OFWASHINGTON
Scapewidestsubapically, narrowingtoapex; 26). Dorsellum (Fig. 26) slightly wider and
funicle and pedicel not or only very slightly shorter than that seen in following species
flattened (Fig. 29). Mesonotum green (Figs. 27-28) . . ChrysocharisgirauhiYoshimoto
ClosteroceniscinctipennisAshmead 13'. Petiole 1.0-1.5 timeslongerthan wide(Figs.
Scapewidestatapex;funicleandpediceldef- 27-28). Dorsellum somewhat narrower and
initelyflattened(Fig. 30). Entirebodypurple longerthan in C. girauhi 14
to bluish 6 14. Eyes comparatively larger (Fig. 23). Petiole
Propodeumdistinctlysculptured,withatleast 1.0-1.5X longer than wide (Fig. 27). Post-
some raised, reticulate sculpture medial to marginal vein about 2x longerthan stigmal
spiracle (Fig. 2) vein(Fig. 11). Hindfemoraalwayscomplete-
ClosterocerustrifasciatusWestwood ly pale yellow . . ChrysocharisoscinidisAshmead
Propodeum smooth over entire surface, or 14'. Eyescomparativelysmaller(Fig. 24). Petiole
with only very faint sculpturing(Fig. 3) . . . aboutaslongaswide(Fig. 28). Postmarginal
Closterocerus utahensisAshmead vein often less than 2x longer than stigmal
Postmarginal vein shorter than the stigmal vein (Fig. 12). Hind femora usually dark,
vein(Figs. 7, 8). Eyesnotemarginate. Meta- rarely completely pale yellow
somawithoutdistinctpetiole(Figs.4, 5).(ge- ChrysocharisainslieiCrawford
nus Chrysonotomyia) 8 15. Notauliincomplete,iftraceabletohindmar-
Postmarginal vein definitely longer (at least gin ofmesoscutum then they are indistinct
1.5 times) than stigmal vein (Figs. 11, 12). and shallow posteriorly (Fig. 43). (Euloph-
Eyes usually more or less emarginate (Figs. ini) 16
23,24).Petiolepresentanddistinct(Figs.25- 15'. Notaulicompleteanddistinct,clearlyreach-
28). (genus Chrysocharis) 10 ingeitherhindmarginofmesoscutumorax-
Forewingsmoothlyroundedapically;slightly illae (Figs. 40-42). (Elachertini) 21
more than 2x longerthan wide (Fig. 7) . . , 16. Antennawith 2 funicularsegments(Fig. 31).
Chrysonotomyiapictipes(Crawford) (genus Diglyphus) 17
Forewingsomewhattruncateapically;slight- 16'. Antennawith 4 funicularsegments(Fig. 32).
ly less than 2x longerthan wide (Fig. 8). . . 9 (genus Pnigalio) . . .Pnigalioflavipes(Ashmead)
Dorsum ofmesosoma with uniform, reticu- 17. Fore wingdensely setose, the basal cell uni-
late, distinctly raised sculpture(Fig. 4). Scu- formly,denselysetose(Fig. 13). Stigmalvein
tellum uniformly colored and postmarginal vein subequal in length.
Chrysonotomyiaformosa(Westwood) Antennal scape alwaysdark 18
Dorsum ofthorax with at least parts ofthe 17'. Fore wing not so setose, the basal cell not
scutellum smooth, or with engraved sculp- uniformlysetose, withat leastsomeareasof
ture, orwith veryweak raised sculpture, but sparser setation (Figs. 14-16). Stigmal vein
in some way differing from strong, uniform, usually shorter than postmarginal vein. An-
raised reticulations (Fig. 5). Scutellum with tennalscapeeitherentirelydark,orwithbas-
twodistinctcolors, purpletoblueanteriorly, al portion white 19
metallicgreen posteriorly 18. Basal 0.25-0.35 ofhind tibia with dark me-
Chrysonotomyiadiastatae(Howard) talliccoloration, some dusky coloration dis-
Chrysonotomyiapunctiventris(Crawford) tally (Fig. 33) Diglyphusintermedins(Girault)
. .
Occiputwithacompletesharp,raisedcarina, 18'. Atleast0.75 ofhindtibiawithdarkmetallic
which extendsthe entire width ofthe vertex coloration;colorationnotfadingdistally(Fig.
(Fig. 19) .... Chrysocharis liriomyzaeDt\ucc\\\ 34) Diglyphus isaea (Walker)
10' Occiput either without carina, or with a ca- 19. Basalportionofhindtibiawithdark,metallic
rina which is only present medially (Fig. ring less than 0.25 the length oftibia (some
20) II dusky coloration may be present past this
II. Longitudinal median groove present in pos- basalring)(Figs. 35-36).Antennalscapeuni-
teriorpart ofmesoscutum and anterior part formly dark Diglyphus begini(Ashmead)
ofscutellum (Fig. 21) Diglyphuscarlylei(Girault)
Chrysochariscrassiscapus(Thomson) 19'. Basalportionofhindtibiawithdark,metallic
II' Mesoscutum and scutellum without median ringatleast0.50the lengthoftibia(Fig. 37).
longitudinal grooves (Fig. 22) 12 Antennal scape white basally 20
12. Petiole more than 2 times longer than wide 20. Postmarginal vein always distinctly longer
(Fig. 25) Chrysocharis ignota Hansson than stigmal vein (Fig. 15)
12' Petiole less than 2 times longer than wide Diglyphus websteri(Crawford)
(Figs. 26-28) 13 20'. Postmarginalveinatmostslightlylongerthan
13. Petiole 1.5-1.8 times longer than wide (Fig. stigmalvein,usuallyshorterthanstigmalvein;
..
VOLUME 93, NUMBER 3 575
stigmal vein somewhat swollen (Fig. 16) . nusChrysocharisrevisedbyHansson(1987),
Diglyphuspulchripes(Crawford) who provideda keyto species, and hostand
21. Notauli curving to meet axillae, well sepa- distribution information. A separate paper
rated from posterior margin ofmesoscutum
(Fig.40). Headvaultedabovecompoundeyes (Hansson 1985) treats Palaearctic members
(Fig. 38). (genus Zagrammosoma) 22 of this genus. Species of Chrysocharis are,
21'. Notauli straight,extendingtoposteriormar- witha fewexceptions, primary, solitary, en-
gin of mesoscutum (Fig. 41). Head not so doparasites oflarvae and pupae ofleafmin-
vaulted, notextendingdistinctly higherthan ing insects. Most are polyphagous, with an
compound eyes (Fig. 39) 25
22. Metasomaentirely black (Fig. 44) 23 extremely wide host range. For complete
22'. Metasomayellow,althoughwithblackmark- host and distribution lists, see Hansson
ings (Figs. 45^6) 24 (1987), Burks (1979b), Peck (1963).
23. Mesosoma dorsally with wide, median yel-
lowstripe(Fig. 44). Forewing(inaddition to
other markings) without a dark stripe along Chrysocharis ainsliei Crawford
apical margin (Fig. 10) (Figs. 12, 24, 28)
Zagrammosoma mirum Girault
23'. Mesosomadorsally black(may have narrow This species and C. oscinidis (= parksi,
yellow stripe laterally on pronotum). Fore-
wing (in addition to other markings) with a viridis) are distinguished from the other
dark stripe along apical margin in posterior Chrysocharis species because they lack the
half(Fig. 9) distinct, completecarina on the vertex, lack
Zagrammosoma lineaticeps(Girault) thelongitudinal, mediangroovesontheme-
24. Anterior margin of mesoscutum with dark soscutum and scutellum, have the petiole
tfroarnmsivnegrsaespsottri(pseomtehtatimeissceonnlcaeragleeddlbaetneeraaltlhy 1.0-1.5 timeslongerthanwide,andthedor-
posterior margin ofpronotum) (Fig. 45) sellum slightlylongerandnarrowerthanthat
. . C
.... Zagrammosoma multilineatum (Ashmead) seen in girauhi. This species is variable,
24'. Anterior margin of mesoscutum without anddifficulttodistinguishfrom C oscinidis.
transverse stripe (Fig. 46) Chrysocharis oscinidis has completely pale
Zagrammosoma americanumGirault
25. Postmarginal vein much longer (nearly 2x) hind femora, and ainsliei has previously
than stigmal vein (Fig. 17). (genus Diauli- been separated from oscinidis as it has dark
nopsis). Thorax entirely metallicgreen .... hind femora. However, Hansson (1987) has
Diaulinopsis callichromaCrawford shown that this character is variable, and
25'. Postmarginalveinaboutaslongasorshorter specimens o{ainslieiwith pale hind femora
than stigmal vein (Fig. 18). (genus Cirrospi-
lus).Thoraxmetallicgreenwithyellowmark- are known. Apparently the only valid char-
ings CirrospilusflavoviridisCrawford acters for separating these two species are
thosegivenby Hansson(1987)andrepeated
in the key in this paper: the eyes are some-
Family Eulophidae what larger in oscinidis than in ainsliei; the
Subfamily Entedoninae petiole in ainsliei is about as long as wide,
Genus Chrysocharis Forster in oscinidis it ranges from 1.0-1.5 x as long
(Figs. 11-12, 19-28) as wide; the postmarginal vein in ainsliei is
often less than 2x longer than the stigmal
Diagnosis.—Scutellum with 2 setae (1 vein, in oscinidis it is always about 2x lon-
pair); submarginal vein with 2 dorsal setae; ger than the stigmal vein.
propodeum without plicae; wings hyaline; Distribution.—Widely distributed and
postmarginal vein distinctly longer than common throughout the Nearctic region.
stigmal vein; petiole present and distinct, Alsoknown fromCuba, Guam, Hawaii,and
usually at least as longas wide; eyes usually Europe.
—
slightly emarginate. Hosts. Larval-pupal parasite of leaf-
Discussion.—Nearctic species of the ge- mining Agromyzidae.
576 PROCEEDINGS OFTHE ENTOMOLOGICAL SOCIETY OFWASHINGTON
- lU ^1^
VOLUME NUMBER
93, 3 577
Plate 2. (Figs. 6-10). Female forewings. 6. Closterocerus utahensis. 1. Chrysonotomyiapictipes. 8. Chryso-
notomyiadiastatae. 9. Zagranimosoma lineaticeps. 10. Zagrammosoma mirum. Scale bars = 0.3 mm. pmv =
postmarginal vein; smv = submarginal vein; sv = sligmal vein.
578 PROCEEDINGS OFTHE ENTOMOLOGICALSOCIETY OF WASHINGTON
Plate 3. (Figs. 1 1-18). Female forewing ventation. 11. Chrysocharisoscinidis. 12. Chrysochahsainsliei. 13.
Diglyphus intermedius. 14. Diglyphusbegini. 15. Diglyphus websteri. 16. Diglyphyspulchripes. 17. Diaulinopsis
callichroma. 18. Cirrospilusflavoviridis. Scale bars = 0.3 mm. be = basal cell; mv = marginal vein; pmv =
postmarginal vein; smv = submarginal vein; sv = stigmal vein.
Chrysocharis giraulti Yoshimoto the other Chrysocharis species because it
(Fig. 26) lacks the distinct, complete carina on the
vertex,lackthelongitudinal,mediangrooves
C
This species has also been treated as on the mesoscutum and scutellum, the pet-
petiolata Girault. It is distinguished from iole is 1.5-1.8 times longer than wide, and
VOLUME 93, NUMBER 3 579
^H
S80 PROCEEDINGS OFTHE ENTOMOLOGICAL SOCIETY OF WASHINGTON
C
Plate 5. (Figs. 25-28). Chrysocharis spp., 2, petioles. 25. C. ignota. 26. C. giraulti. 27. oscinidis. 28. C.
ainsliei. Scale bars = 0.1 mm. ds = dorsellum; pr = propodeum; pt = petiole.
distinct, completecarinaonthe vertex, lacks from the other Chrysocharis species by the
thelongitudinal, mediangroovesontheme- strong, sharp, complete occipital carina
soscutum and scutellum, and the petiole is which extends from the vertex all the way
at least 2.0 times longer than wide. This onto the gena (Fig. 19). This species was
species was recently described by Hansson introduced into North American as a par-
(1987) who, recorded it from Liriomyza sp. asite o{Agromyzafrontella (Rondani), with
mining herbs. It is included, even though subsequentestablishment (Hendrickson and
the exact host species of Liriomyza is yet Barth 1979, as C. punctifacies). This is the
unknown. only known Nearctic host, although it is
Distribution.—USA, Canada. Also Cen- known to attack Liriomyza species includ-
tral America and northern South America. ingL. trifolii) in Europe(BoucekandAskew
Hosts.—From Liriomyza sp. on herbs. 1968), and it is likely that it will attack Lir-
iomyza in North America.
Chrysocharis liriomvzae Delucchi —
Distribution. Palearctic; introduced into
(Fig. 19) eastern United States, with subsequent re-
This species has also been treated as C. covery (Hendrickson and Barth 1979, as C.
punctifacies Delucchi. It is distinguished punctifacies).