Palearctic Sympiesis acalle and Sympiesis gordius (Hymenoptera: Eulophidae) in north America : Taxonomic changes and a review of nearctic host records

2006·6.6 MB·

by  Chris T Maier

PROC. ENTOMOL. SOC. WASH.
108(1), 2006, pp. 14-23
PALEARCTIC SYMPIESIS ACALLE AND SYMPIESIS GORDIUS
(HYMENOPTERA: EULOPHIDAE) IN NORTH AMERICA: TAXONOMIC
CHANGES AND A REVIEW OF NEARCTIC HOST RECORDS
CHRIS T. MAIER AND CHRISTER HANSSON
(CTM) Department of Entomology, Connecticut Agricultural Experiment Station, P.O.
Box 1106, New Haven, CT 06504, U.S.A. (e-mail: [email protected]); (CH)
Department of COB, Zoology, Helgonavagen 3, SE-223 62 Lund, Sweden (e-mail:
Christer. Hansson @cob.lu.se)
Abstract.4The Palearctic eulophids, Sympiesis acalle (Walker) and S. gordius (Walker),
are widespread in North America where they have been known as S. bimaculatipennis
Girault and S. marylandensis Girault, respectively. To update their taxonomy, we have
established seven new synonyms: Astichus bimaculatipennis Girault, Sympiesis bimacu-
lata Crawford, and S. meteori Girault = S. acalle (Walker); and, S. marylandensis Girault,
S. rex Girault, S. miltoni Girault, and S. lexingtonensis Girault = S. gordius (Walker). We
also have designated six lectotypes: Eulophus acalle Walker, E. gordius Walker, E. ala-
parus Walker, E. pisenor Walker, Sympiesis rex Girault, and S. lexingtonensis Girault.
Finally, we summarize the North American host records of Sympiesis acalle and S. gor-
dius, which frequently are cited as biocontrol agents of pestiferous leafminers.
Key Words: Eulophidae, hosts, leafminer, Phyllonorycter, Sympiesis bimaculatipennis,
Sympiesis marylandensis
Many eulophid wasps (Chalcidoidea: Eu- idoptera: Gracillariidae), of cultivated ap-
lophidae) have a large distribution, with ple, Malus domestica Borkhausen. Among
some species occurring on two or more their images were two eulophid species,
continents (Noyes 2002). Numerous species Sympiesis acalle (Walker) and S. gordius
occur in both Europe and North America (Walker) that appeared remarkably similar
(e.g., Hansson 1987), a fact often over- to the North American species, S. bimacu-
looked because certain species have a dif- latipennis (Girault) and S. marylandensis
ferent name in Europe and North America. Girault, respectively. In North America,
Such nomenclatural mistakes make it diffi- Miller (1970), Maier (1988a, b), and others
cult to understand fully the natural history have recorded hosts of these two parasitoids
of a species and even might interfere with to be species of Phyllonorycter and other
biocontrol efforts. Here we reveal two ad- Lepidoptera whose larvae live in mines and
ditional cases of mistaken identity, both in other concealed situations. In Canada and
the genus Sympiesis Forster. In the process, the United States, S. marylandensis is an
we establish seven new synonyms and des- important natural enemy of leafminers, es-
ignate six lectotypes. pecially the apple pests, Phyllonorycter
Bellostas et al. (1998) have published blancardella (Fabricius), P. crataegella
color photographs of parasitoids of Euro- (Clemens), and P. mespilella (Hiibner)
pean leafminers, Phyllonorcyter spp. (Lep- (e.g., Maier 1994, 2002). In this paper, we
VOLUME 108, NUMBER 1 15
note the synonymy and summarize the single remaining specimen agrees with the
North American records of hosts of the two original description. Walker (1848) did not
Sympiesis spp. This information hopefully indicate on how many specimens his de-
will assist economic entomologists in rec- scription was based; therefore, to keep the
ognizing that the biocontrol previously at- nomenclature stable, the name hereby is
tributed to two <8native=99 species is, in fact, fixed to the male specimen which is here
caused by two European species, or possi- designated the lectotype.
bly holarctic species, that have long been The female holotypes of A. bimaculati-
associated with apple leafminers in Euro- pennis, S. bimaculata, and S. meteori agree
pean orchards. well with our interpretation of S. acalle
(i1.e., with females linked to males that are
LECTOTYPE DESIGNATIONS AND conspecific with the lectotype male of S.
NEw SYNONYMS acalle); and, S. acalle, S$. bimaculatipennis,
Type specimens discussed below are de- S. bimaculata, and S. meteori undoubtedly
posited in the following museums: BMNH, belong to the same species. The holotype of
The Natural History Museum, London, A. bimaculatipennis has the mesosoma, in-
England; INHS, Illinois Natural History cluding only front left leg, glued to a point
Survey, Champaign, IL, USA; USNM, (type no. 45.012); the head (squashed) and
Smithsonian Institution, National Museum the antennae are on a slide (type no.
of Natural History, Washington, DC, USA. 45.013); wings, most of the legs, and the
gaster are missing. The holotype of S. bi-
Sympiesis acalle (Walker) maculata is glued to a point (type no.
15094); the entire left antenna, right anten-
Eulophus acalle Walker 1848: 234. Lecto-
na beyond the scape, and right wing pair
type male (new designation) in BMNH,
are missing. The holotype of S. meteori is
examined.
glued to a point (type no. 19995) and is
Entedon nubeculatus Ratzeburg 1848: 158.
broken in two parts, gaster and mesosoma;
Synonymized by Bouéek (1959).
the head, pair of right wings and left hind
Eulophus bifasciatus Thomson 1878: 230.
wing, right fore tarsus, right mid tibia and
Synonymized by Bouéek (1959).
tarsus, and left and right hind tibiae and tar-
Eulophus punctifrons Thomson 1878: 231.
Si are missing.
Synonymized by Bouéek (1959).
Astichus bimaculatipennis Girault 1912: 8.
Symptesis gordius (Walker)
Holotype female in INHS, examined.
Eulophus gordius Walker 1839: 129. Lec-
New synonymy.
totype male (new designation) in
Sympiesis bimaculata Crawford 1913: 259.
BMNH, examined.
Holotype female in USNM, examined.
Eulophus alaparus Walker 1839: 163. Lec-
New synonymy.
totype female (new designation) in
Sympiesis meteori Girault 1916: 37. Holo-
BMNH, examined. Synonymized by
type female in USNM, examined. New
Bouéek (1959).
synonymy.
Eulophus pisenor Walker 1839: 153. Lec-
Sympiesis acalle: Graham 1959: 182.
totype male (new designation) in
A single male with the name 8<Eulophus BMNH, examined. Synonymized by
acalle=9 is in the collection at BMNH (with Bouéek and Askew (1968).
the label <Eulophus acalle Walker9? and Eulophus cervicornis Forster 1841: 43.
our lectotype label). The gaster has been Synonymized by Bouéek (1959).
gnawed on left side, but otherwise the spec- Entedon padellae Ratzeburg 1844: 166.
imen is intact. The original description was Synonymized by Bouéek (1959).
based on one or more male specimens. The Eulophus bulmerincqii Ratzeburg 1848:
16 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
155. Synonymized by Boucek and USNM. Girault (1917d) did not indicate on
Askew (1968). how many specimens he based his descrip-
Eulophus laevissimus Ratzeburg 1848: 157. tion. Therefore, to stabilize the nomencla-
Synonymized by Bouéek (1959). ture, we chose this specimen as the lecto-
Eulophus stramineipes Thomson 1878: type for S. lexingtonensis. The specimen
232. Synonymized by Bouéek (1959). carries the labels <1866,= <20196,= <8Sym-
Sympiesis rex Girault 1917a: 2-3. Lecto- piesis lexingtonensis Gir. female type,= and
type female (new designation) in our lectotype label.
USNM, examined. New synonymy. The female holotype of S$. marylandensis,
Sympiesis miltoni Girault 1917b: 7. Lecto- and the lectotypes of S. rex, S. miltoni, and
type female, designated by Miller (1970), S. lexingtonensis agree well with our inter-
in USNM, examined. New synonymy. pretation of S. gordius (1.e., with females
Sympiesis marylandensis Girault 1917c: 37. linked to males that are conspecific with the
Holotype female in USNM, examined. lectotype male of S. gordius); and, S. gor-
New synonymy. dius, S. rex, S. miltoni, S. marylandensis,
Sympiesis lexingtonensis Girault 1917d: 6. and S. /exingtonensis undoubtedly belong to
Lectotype female (new designation) in the same species. The lectotype of S. rex is
USNM, examined. New synonymy. glued to a point (type no. 20218); the head
Eulophus albiscapus Erdos 1954: 327. Syn- and right wing pair are missing. The lec-
onymized by Bouéek (1959). totype of S. miltoni is glued to a point (type
Sympiesis gordius: Graham 1959: 183. no. 21399), and the type is intact. The ho-
lotype of S. marylandensis is glued to a
Single specimens under the names <Eu- point (type no. 20137); the head is missing
lophus gordius= (a male), <<Eulophus ala- (according to the original description the
parus= (a female), and <8Eulophus pisenor= head is on a slide, not examined by us). The
(a male) are in the collection at BMNH. lectotype of S. lexingtonensis is glued to a
Walker9s (1839) descriptions of these three point (type no. 20196); the head, right legs
species were based on an unknown number (all three), and pair of right wings are miss-
of specimens. Therefore, to stabilize the no- ing.
menclature, we chose these three specimens
as the lectotypes for each species. The spec- BIOLOGY
imens are labelled 8<8Eulophus gordius Hosts of both S. acalle and S. gordius
Walker,= <<Eulophus alaparus Walker,= and that were discovered during original North
**Eulophus pisenor Walker9; each also has American studies (Table 1) are repeated in
our lectotype label. the catalogs of Peck (1951, 1963) and
A single specimen under the name Sym- Burks (1979) and, to some extent, in the
piesis rex is in the collection at USNM. Gi- revision of Miller (1970). Green (1979) and
rault (1917a) did not indicate on how many Maier (1988b) have since reported many
specimens he based his description. There- new North American hosts. Sympiesis acal-
fore, to stabilize the nomenclature, we le, previously known as S. bimaculatipennis
chose this specimen as the lectotype for S. in North America, is both a primary and
rex. The specimen carries the labels secondary ectoparasitoid of lepidopteran
8<8Quaintance No. 11468,= <Salem, Ore, larvae and pupae (Table 1) that usually live
vi.15,= <<Reared Phyllonorycter crataegel- in concealed places, such as blotch mines
la,= <*E. J. Newcomer collector,9 <20218,= or rolled leaves. Sympiesis acalle has at
<Sympiesis rex Girault female type,9 and least 15 hosts in North America. It is a pri-
our lectotype label. mary parasitoid of Tischeriidae (1 species),
A single specimen stands under the name Gracillariidae (5), Oecophoridae (1), and
Sympiesis lexingtonensis in the collection at Tortricidae (6). In addition, it parasitizes the
VOLUME 108, NUMBER 1 7
Table 1. Hosts of Sympiesis acalle and S. gordius that were recorded as S. bimaculatipennis and S. mary-
landensis, respectively, in the North American literature. Due to taxonomic revisions or changes in generic use
and species concepts, some of the hosts cannot be placed in a genus and are excluded here. Hosts given only
to genus are followed by the genus of their host plant in brackets; if these hosts could be one of the fully
identified hosts, they are not included in the total number of hosts given in the text. Generic records without an
associated host plant are excluded from the main part of the table. To the best of our knowledge, all of the
scientific names are the ones currently in use.
Order, Family, and Species of Host S. acalle 4 S. gordius Literature Sources
LEPIDOPTERA
Nepticulidae
Nepticula variella (Braun) x Green (1979)
Tischeriidae
Tischeria malifoliella Clemens* x xX Dunnam (1924), Peck (1951, 1963), Miller
(1970), Burks (1979), Oatman (1985)
Tischeria sp. [Solidago] x Maier (1994, unpublished data)
Heliozelidae
Coptodisca powellella Opler x Green (1979)
Lyonetidae
Bacculatrix albertiella Busck x Green (1979)
Gracillariidae
Caloptilia agrifoliella Opler xX Green (1979)
Caloptilia sp. [Betula] x Miller (1970), Burks (1979)
Cameraria agrifoliella (Braun) x Green (1979)
C. caryaefoliella (Clemens) xX Heyerdahl and Dutcher (1985)
C. wislizeniella Opler xX Green (1979)
Cameraria spp. [Ostrya] xX Maier (1988b)
Cremastobombycia solidaginis x Maier (1988b)
(Frey and Boll)
Neurobathra bohartiella (Opler) x Green (1979)
Parectopa robiniella Clemens XxX Weaver and Dorsey (1965)
Parornix geminatella (Packard)* x XxX Haseman (1916), Girault (1916), Copenhafer
and Parker (1938), Peck (1951, 1963), Mill-
er (1970), Burks (1979), Oatman (1985),
Maier (1988b)
Parornix sp. [Betula] xX Maier (1988b)
Parornix sp. [Lyonia] x Maier (1988b)
Phyllocnistis populiella Chambers Xx Miller (1970), Burks (1979)
Phyllonorycter aeriferella (Clemens) xX Maier (1988b)
P. albanotella (Chambers) x Maier (1988b)
P. argentifrimbriella (Clemens) x Maier (1988b)
P. argentinotella (Clemens) xX Maier (1988b)
P. antiochella (Opler) xX Green (1979)
P. auronitens (Frey and Boll) x Maier (1988b)
P. basistrigella (Clemens) xX Miller (1970), Burks (1979), Maier (1988b)
P. blancardella (Fabricius) x x Miller (1970), Pottinger and LeRoux (1971),
Johnson et al. (1976), Dutcher and Howitt
(1978), Burks (1979), Weires et al. (1980),
Van Driesche and Taub (1983), Maier
(1984, 1988b, 1992, 1993, 1994, 2002),
Hagley (1985), Ridgway and Mahr (1985,
1989, 1990), Biggs and Hagley (1988),
Hagley and Barber (1991), Gagné and Bar-
rett (1994), Bishop et al. (2001)
P. caryaealbella (Chambers) x Heyerdahl and Dutcher (1985)
18 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 1. Continued.
Order, Family, and Species of Host S. acalle S. gordius Literature Sources
P. celtisella (Chambers) x Miller (1970), Burks (1979)
P. crataegella (Clemens) xX x Beckham et al. (1950), Peck (1951, 1963), Par-
ent (1962), Miller (1970), Burks (1979), Wei-
res et al. (1980), Maier (1982, 1988b, 1992,
1993, 1994), Gambino and Sullivan (1982),
Van Driesche (1983), Van Driesche and Taub
(1983), Drummond et al. (1985), Oatman
(1985), Van Driesche et al. (1985), Gagné and
Barrett (1994), Maier and Weseloh (1995)
P. emberizaepenella (Bouché) x Maier (1988b)
P. felinelle Heinrich Xx Miller (1970), Burks (1979)
P. fitchella (Clemens) x Maier (1988b)
P. inusitatella (Braun) xX Green (1979)
P. kearfottella (Braun) xX Maier (1988b)
P. lucetiella (Clemens) Xx Maier (1988b)
P. lucidicostella (Clemens)* x Miller (1970), Burks (1979), Maier (1988b)
P. mespilella (Hiibner) x x Doganlar and Beirne (1980), Barrett and Jor-
gensen (1986), Varela dn Welter (1992),
Cossentine and Jensen (1994), McGregor
(1996)
P. morrisella (Fitch) xX Maier (1988b)
P. obscuricostella (Clemens) Xx Maier (1988b)
P. propinquinella (Braun) x Miller (1970), Burks (1979), Maier (1988a, b,
1994)
P. restrictella (Braun) Xx Maier (1988b)
P. robiniella (Clemens) xX Weaver and Dorsey (1965), Miller (1970),
Burks (1979), Maier (1988b)
P. salicifoliella (Chambers)! xX Miller (1970), Burks (1979), Maier (1988b)
P. sandraella (Opler) x Green (1979)
P. scudderella (Frey and Boll) xX Maier (1988b)
P. tremuloidiella (Braun) Xx Maier (1988b)
P. trinotella (Braun)! xX Maier (1988b)
P. tritaenianella (Chambers) Xx Maier (1988b)
Phyllonorycter sp. near crataegella x xX Maier (1988a, b, 1994)
(Clemens)
Phyllonorycter sp. [Alnus] Xx Miller (1970)
Phyllonorycter sp. [Amphicarpa| x Miller (1970)
Phyllonorycter sp. [Fagus]' xX Miller (1970)
Phyllonorycter sp. |Lonicera] xX Miller (1970)
Phyllonorycter sp. [Malus] x Girault (1917a), Peck (1951, 1963), Newcom-
ber (1958), Miller (1970)
Phyllonorycter sp. [Populus] xX Miller (1970)
Phyllonorycter sp. [Prunus] xX Miller (1970)
Phyllonorycter sp. [Tilia] XxX Miller (1970)
Oecophoridae
Agonopteryx posticella (Wal- xX Miller (1970), Burks (1979)
singham)
Cosmopterygidae
Stilbosis dulcedo (Hodges) Xx Green (1979)
Gelechiidae
Coleotechnites thujaella (Kearfott) xX Miller (1970), Burks (1979)
Tortricidae
Acleris chalybeana (Fernald) xX Miller (1970), Burks (1979)
VOLUME 108, NUMBER 1 IS)
Table 1. Continued.
Order, Family, and Species of Host S. acalle S. gordius Literature Sources
Ancylis comptana (Frolich) x Miller (1970), Burks (1979)
Choristoneura rosaceana (Harris) x Schuh and Mote (1948), Peck (1963)
Epinotia laracana (Kearfott) xX Miller (1970)
Rhopobota naevana (Hibner) x Fitzpatrick et al. (1994)
Spilonota laricana (Heinemann) xX Burks (1979)
Spilonota ocellana (Denis and xX Sadava and Miller (1967), Miller (1970),
Schiffermiiller) Burks (1979)
HYMENOPTERA
Tenthredinidae
Heterarthrus nemoratus (Fallén) Miller (1970), Burks (1970)
Braconidae"
Pholetesor ornigis (Weed)' x Pottinger and LeRoux (1971), Maier (1982,
1984), Bishop et al. (2001)
Chalcididae
Spilochalcis sp.' x Bishop et al. (2001)
*Dunnam9s (1924) record apparently is repeated by Peck (1951, 1963), Miller (1970), and Burks (1979).
> Both Miller (1970) and Burks (1979) gave the Caloptila host as C. elongella (Linnaeus), but this species is
not known to occur in North America.
© Haseman9s (1916) study provided the record of S. acalle given in Girault (1916) and Peck (1951, 1963), and
Copenhagen and Parker9s (1938) record apparently also was repeated for one of the junior synonyms of S. acalle
in Peck (1963); the record in Burks (1979) probably was repeated from an earlier catalog.
4 Girault (1917d) listed a host as a blotch miner on maple, which may have been a Phyllonorycter sp. or
possibly a Cameraria sp.
© This species on Populus sp. mentioned in Miller (1970) and probably repeated by Burks (1979) is more
likely to be Phyllonorycter tremuloidiella (Braun) or another Phyllonorycter sp. on Populus; P. salicifoliella is
a willow-feeder.
8Procter (1938, 1946) mentioned S. gordius as a parasite of blotch miners on Fagus (probably Phyllonorycter
restrictella) and Salix (probably Phyllonorycter scudderella or P. salicifoliella), and the record of the blotch
miner on Fagus is repeated in Peck (1951, 1963); the record of the blotch miner on Fagus is duplicated in Peck
(1951, 1963) and Miller (1970).
= Newcomer (1958) did not report his record from an apple-feeding Phyllonorycter sp. (misidentified as P.
crataegella) until 1958; this record is the one mentioned in Girault (1917a).
" Webster (1909) retrieved S. acalle from an insectary and reported that is parasitized an unidentified internal
primary parasitoid of the tortricid moth, Acleris minuta (Robinson), a record that was repeated, sometimes in
altered form, by Girault (1912), Frison (1927), Peck (1951, 1963), and Burks (1979).
8Primary hosts of S. acalle on Pholetesor ornigis were Phyllonorycter blancardella and P. crataegella; that
from Spilochalcis sp. was P. blancardella.
braconid Pholetesor ornigis (Weed) and the Under the name of S$. marylandensis in
chalcid Spilochalcis sp., which both attack the North American literature, S. gordius is
apple leafminers, Phyllonorycter spp. The known to be a primary ectoparasitoid of lar-
casual rearing methods used in some stud- vae and sometimes pupae of miners, skel-
ies (e.g., Oatman 1985) may have resulted etonizers, and rollers of leaves, and of other
in overlooking hyperparasitic associations insects (Table 1) that live in sheltered situ-
or in identifying the wrong primary host. In ations. Most of the original host records are
northeastern North America, S. acalle is a from Miller (1970), Green (1979), and
facultative hyperparasitoid of the gracillari- Maier (1988b). In North America, S. gor-
id leafminers Phyllonorycter blancardella dius parasitizes at least 53 species in the
and P. crataegella, attacking especially families Nepticulidae (1), Tischeriidae (2),
their braconid parasitoid, Pholetesor ornigis Heliozelidae (1), Lyonetidae (1), Gracillar-
(e.g., Maier 1982, 1984). lidae (44), Cosmopterygidae (1), Gelechi-
20 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
idae (1), Tortricidae (1), and Tenthredinidae entry into North America. Although it is
(1). Maier (1988b) has concluded that in the less likely, these Sympiesis spp., known so
northeastern United States S. gordius pre- long by their junior synonyms in North
fers gracillariid larvae inhabiting mines on America, actually could be Holarctic or
the underside of leaves. Some of the west- could have spread from North America to
ern host records may refer to parasitization Europe. Comparative studies of genetic di-
by S. yuekseli Doganlar (1979), which is versity of populations in Europe and North
similar to S. gordius and was described af- America might clarify the continent(s) of
ter some of the records in Table 1 were pub- origin of these Sympiesis spp.
lished. Based on host records given by Bou-
cek and Askew (1968), Storozheva (1982), ACKNOWLEDGMENTS
and others, both S. acalle and S. gordius We are grateful to C. Favret (INHS), M.
have similar hosts in the Palearctic and Ne- W. Gates (USNM), and S. Lewis (BMNH)
arctic regions. Delucchi (1958), Boucek and for the loan of specimens.
Askew (1968), Storozheva (1982), and oth-
ers have reported that S. gordius sometimes LITERATURE CITED
is a secondary parasitoid of braconids and
Baggiolini, M. 1960. Observations sur la biologie de
eulophids, but this type of association is not
deux mineuses du genre Lithocolletis: L. coryli-
yet known in North America. foliella et L. blancardella (Lep. Gracilariidae), nu-
Both S. acalle and S. gordius are men- isibles aux arbres fruitiers en Suisse romande.
tioned in the biocontrol literature about gra- Mitteilungen der Schweizerischen Entomologisch-
en Gesellschaft 32: 385-397.
cillariids in apple orchards in Europe (e.g.,
Barrett, B. A. and C. D. Jorgensen. 1986. Parasitoids
Baggiolini 1960, Casas and Baumgartner
of the western tentiform leafminer, Phyllonorycter
1990, Bellostas et al. 1998) and in North elmaella (Lepidoptera: Gracillariidae), in Utah ap-
America (Maier 1994, 2002, and references ple orchards. Environmental Entomology 15:
therein). Especially S$. gordius has been 635-641.
Beckham, C. M., W. S. Hough, and C. H. Hill. 1950.
touted as an important natural enemy of the
Biology and control of the spotted tentiform leaf
pestiferous apple leafminers, Phyllonoryc-
miner on apple trees. Virginia (Blacksburg) Ag-
ter blancardella, P. crataegella, and P. ricultural Experiment Station Technical Bulletin
mespilella (see references in Table 1). 114: 1-19.
Maier (1984, 1992) and Gagné and Barrett Bellostas, J. J., E. Olivella, M. J. Verdu, M. J. Sarastia,
and J. Avilla. 1998. Fauna parasitoide de Phyllon-
(1994) have examined the seasonal flight
orycter spp. en plantaciones de manzano de Llei-
activity of S. gordius; Maier (1992) sug-
da. Boletin de Sanidad Vegetal 24: 313-320.
gested that it could have six or seven gen- Biggs, A. R. and E. A. C. Hagley. 1988. Effects of
erations per year while its host P. cratae- two sterol-inhibiting fungicides on populations of
gella has only three. Also, Maier and We- pest and beneficial arthropods on apple. Agricul-
ture, Ecosystems & Environment 20: 235-244.
seloh (1995) have shown that S. gordius
Bishop, S. D., R. EF Smith, C. Vincent, H. Goulet, J.
tended to have higher abundance near the
Huber, G. Gibson, M. Sharkey, and J. H. Borden.
edge than the center of apple orchards. 2001. Hymenopterous parasites associated with
We consider it to be highly likely that the Phyllonorycter blancardella (Lepidoptera: Gracil-
two European eulophids were introduced lariidae] in Nova Scotia and Quebec. Phytopro-
tection 82: 65-71.
accidentally into North America with one
Boucek, Z. 1959. A study of Central European Eulo-
or both of their European hosts, P. blan-
phidae. I: Eulophinae (Hymenoptera). Acta Ento-
cardella and P. mespilella, which are leaf- mologica Musei Nationalis Pragae 33: 1174170.
miners of non-native apple. The European Bouéek, Z. and R. R. Askew. 1968. Index of Ento-
facultative hyperparasitoid, S$. sericeicornis mophagous Insects. Palearctic Eulophidae (excl.
Tetrastichinae) (Hym. Chalcidoidea). Le Frangois,
(Nees), which for many years masqueraded
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