c 3
Journal of
Hymenoptera
Research
. .IV
6«**
Volume 15, Number 2
October 2006
ISSN #1070-9428
CONTENTS
BELOKOBYLSKIJ, S. A. and K. MAETO. A new species of the genus Parachremylus Granger
(Hymenoptera: Braconidae), a parasitoid of Conopomorpha lychee pests (Lepidoptera:
Gracillariidae) in Thailand 181
GIBSON, G. A. P., M. W. GATES, and G. D. BUNTIN. Parasitoids (Hymenoptera: Chalcidoidea)
of the cabbage seedpod weevil (Coleoptera: Curculionidae) in Georgia, USA 187
GILES, V. and J. S. ASCHER. A survey of the bees of the Black Rock Forest Preserve, New York
(Hymenoptera: Apoidea) 208
GUMOVSKY, A. V. The biology and morphology of Entedon sylvestris (Hymenoptera:
Eulophidae), a larval endoparasitoid of Ceutorhynchus sisymbrii (Coleoptera:
Curculionidae) 232
KULA, R. R., G. ZOLNEROWICH, and C. J. FERGUSON. Phylogenetic analysis of Chaenusa
sensu lato (Hymenoptera: Braconidae) using mitochondrial NADH 1 dehydrogenase
gene sequences 251
QUINTERO A., D. and R. A. CAMBRA T The genus Allotilla Schuster (Hymenoptera: Mutilli-
dae): phylogenetic analysis of its relationships, first description of the female and
new distribution records 270
RIZZO, M. C. and B. MASSA. Parasitism and sex ratio of the bedeguar gall wasp Diplolqjis
rosae (L.) (Hymenoptera: Cynipidae) in Sicily (Italy) 277
VILHELMSEN, L. and L. KROGMANN. Skeletal anatomy of the mesosoma of Palaeomymar
anomalum (Blood & Kryger, 1922) (Hymenoptera: Mymarommatidae) 290
WHARTON, R. A. The species of Stenmulopius Fischer (Hymenoptera: Braconidae, Opiinae)
and the braconid sternaulus 316
(Continued on back cover)
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J. HYM. RES.
Vol. 15(2), 2006, pp. 181-186
A New Species of the Genus Parachremylns Granger (Hymenoptera:
Braconidae), a Parasitoid of Conopomorpha Lychee Pests (Lepidoptera:
Gracillariidae) in Thailand
Sergey A. Belokobylskij and Kaoru Maeto
(SAB) Zoological Institute Russian Academy of Sciences, St. Petersburg, 199034, Russia,
and Museum and Institute of Zoology Polish Academy of Sciences, Wilcza 64,
Warszawa 00-679, Poland; email: sb@zin.ru
(KM) Laboratory of Insect Science, Faculty of Agriculture, Kobe University, Rokkodai-machi 1-1,
Nada-ku, Kobe 657-8501, Japan; email: maeto@kobe-u.ac.jp
Abstract. — Parachremylns litchii Belokobylskij & Maeto, new species, from Thailand is
described as a parasitoid of larvae of Conopomorpha sinensis Bradley and C. litchiella Bradley, the
major pests of lychee and longan in South-East Asia. The taxonomic position of Parachremylns and
the range of the hosts of related genera of parasitoids are discussed.
Several insect pests are seriously threat- The genus Parachremylns with type spe-
ening lychee (Litchi chinensis Sonn.) and cies P. seyrigi Granger was originally de-
longan (Dimocarpus longan Lour.) (Sapin- scribed from Madagascar (Granger 1949);
daceae) growers. They are the fruit borer this genus occurrs also in continental
{Conopomorpha sinensis Bradley), leaf miner Africa — Nigeria and Niger (Wharton
(Conopomorpha litchiella Bradley), longan 1993). Two additional species of this genus
sucking bug (Tessaratoma papillosa Drury), have already been recorded from the
fruit piercing moth (Othreis fnllonia Oriental region. Parachremylns oblongus
(Clerck)), and twig borer (Zenzera coffeae (Papp) was described from India in the
Nietner) (Menzel 2002). genus Avga Nixon (Papp 1990, 1997), and
Conopomorpma sinensis, the lychee stem- P. temporalis Belokobylskij from Brunei
end borer and the lychee fruit borer in (Belokobylskij 1999). A fourth species of
China, Thailand and India, is the major this genus, similar to P. temporalis, is
pest of lychee and longan in these coun- described below from Thailand. The sys-
tries. Conopomorpha sinensis and the related tematic position of this genus is disputable.
C. litchiella both attack lychee and longan, Parachremylns is included in the subfamily
the latter preferring to mine leaves and Exothecinae (tribe Avgini: Belokobylskij
shoots (Bradley 1986). There have been 1993), or conventionally in subfamily Hor-
only tentative reports on braconid para- miinae (Wharton 1993). In spite of the
sitoids of the pest Conopomorpiha borers: different understanding of the contents of
Phanerotoma sp., Pholetesor (Apanteles) sp., subfamilies, the position of this genus close
and Colastes sp. (Menzel 2002, Anupunt to Avga Nixon is suggested by both
and Sukhvibul 2005), but possibly infor- authors. Belokobylskij (1993) discussed
mation about Colastes is due to misdeter- the relationships of these genera with
mination. Here we report a new braconid Parahormins Nixon, Pseudohormius Tobias
of the genus Parachremylns Granger as & Alexeev and Allobracon Gahan (= Lcur-
a larval parasitoid of C. sinensis and C. inion Muesebeck), which share the loss of
litchiella. the prepectal (epicnemial) carina on the
182
Journal of Hymenoptera Research
mesosoma. Wharton (1993) provisionally
placed Avga near Parahormius and Pseudo-
hormius and showed the possible relation-
ship of Avga and Parachremylus (shared
granulate mesonotal sculpture and the
poorly developed propleural flange). How-
ever, in his opinion, Allobracon does not
appear to be closely related to Parachremy-
lus in spite of it sharing a number of
features with Avga and Parahormius.
The host of Parachremylus has not been
known till now. The new species described
below as P. litchii sp. nov. was reared from
larvae of Conopomorpha sinensis and C.
litchiella (Gracillariidae), both important
pests of lychee and longan trees in South-
East Asia. The members of related genera
of the tribe Avgini {Parahormius, Avga,
Allobracon) are also recorded as parasitoids
of the leaf-rollers or leaf-miners of the
families Tortricidae, Gracillariidae, Lyone-
tiidae, Cosmopterigidae, Coleophoridae,
and Gelechiidae (Belokobylskij 1993, Whar-
ton 1993) as well as rarely (recorded for
Allobracon) of leaf-mining Coleoptera
(Wharton 1993).
The terms of wing venation are used as
defined by Belokobylskij and Tobias (1998).
The following abbreviations are used:
POL — postocellar line; OOL — ocular-ocellar
line; Od — maximum diameter of lateral
ocellus; NIAES— National Institute of
Agro-Environmental Sciences (Tsukuba,
Japan); ZISP— Zoological Institute, Rus-
sian Academy of Sciences (St. Petersburg,
Russia).
Parachremylus litchii Belokobylskij &
Maeto, new species
(Figs 1-11)
Holotype female.— "Horticultural Re-
search Center, Chiang Rai, Thailand, viii,
1996, Supatra Dolsopon", "Host: Conopo-
morpha litchiella larvae on Lychee or Lon-
gan" (NIAES). Paratypes. 2 females, 1
male, with the same labels as holotype
(NIAES, ZISP); 5 females, "Horticultural
Research Center, Chiang Rai, Thailand, 6.
vi, 1997, Supatra Dolsopon", "Host: Con-
opomorpha sinensis larvae" (NIAES, ZISP).
Description. — Female. Body length 2.6-
2.8 mm; fore wing length 2.5-2.6 mm.
Antennae: thickened, almost filiform, 29-
30-segmented, 1.1-1.2 times longer than
body. Scapus 1.7-2.0 times longer than
wide. First flagellar segment 2.5-2.8 times
longer than its apical width, 1.1-1.2 times
longer than second segment. Penultimate
segment 2.0-2.3 times longer than wide,
0.6-0.7 times as long as first flagellar
segment, 0.7-0.75 times as long as apical
segment; the latter with distinct spine
apically. Head: width 1.8-2.0 times its
median length, 1.25-1.4 times width of
mesoscutum. Temple very strongly and
almost linearly narrowed behind eye (dor-
sal view). Transverse diameter of eye
(dorsal view) 5.5-7.0 times longer than
temple length (7.0-7.7 times if measured on
straight line). Ocelli small, in triangle with
base 1.1-1.15 times its sides. POL 0.7-1.0
times Od, 0.3-0.5 times OOL. Vertex with
narrow median longitudinal furrow. Oc-
cipital carina dorsally distinctly curved
towards ocelli, rather widely interrupted
medially; not fused with hypostomal cari-
na ventrally being obliterated for a short
distance. Eye large, sub-round, glabrous,
1.1-1.2 times as high as broad. Malar space
0.25-0.3 times height of eye, 0.8-0.9 times
basal width of mandible. Face width 0.9
times height of eye and 1.2-1.25 times
height of face and clypeus combined.
Malar suture absent. Clypeal suture rather
distinct and complete. Clypeus weakly
convex. Hypoclypeal depression sub-
round, its width 0.8-0.9 times distance
from edge of depression to eye, 0.35 times
width of face. Head below eyes (front
view) strongly and almost linearly nar-
rowed. Mesosoma: length 1.5-1.55 times its
height. Mesoscutum highly and almost
perpendicularly raised above pronotum
(lateral view), with rather fine longitudinal
medioposterior keel (dorsal view). Notauli
rather narrow, shallow anteriorly on verti-
cal surface and very shallow to almost
Volume 15, Number 2, 2006
183
Figs 1-11. Parachremylus litchii sp. nov. 1, Head, frontal view. 2, Head, dorsal view. 3, Propodeum. 4, Six basal
segments of antenna. 5, Mesosoma, lateral view. 6, Hind tibia and two basal segments of hind tarsus. 7,
Metasoma, dorsal view. 8, Hind coxa. 9, Hind femur. 10, Fore wing. 11, Hind wing.
184
Journal of Hymenoptera Research
absent on dorsal surface, finely sculptured.
Prescutellar depression short, shallow,
finely crenulate-granulate, 0.15-0.2 times
as long as scutellum. Scutellum almost flat.
Metanotum medially with small and ob-
tuse tubercle. Subalar depression rather
shallow, wide, densely and curvedly stri-
ate with fine granulation anteriorly. Ster-
nauli shallow, rather wide, weakly curved,
entirely smooth. Wings: Length of fore
wing 2.8-3.0 times its maximum width.
Radial cell not shortened, metacarpus 1.3
times longer than pterostigma. Pterostigma
rather wide, 3.1-3.7 times longer than
wide. Radial vein arising a little or rather
distinctly before middle of pterostigma.
Second radial abscissa 1.5-2.3 times longer
than first abscissa, 0.25-0.3 times as long as
the straight third abscissa, 1.15-1.25 times
longer than the weakly curved first radio-
medial vein. Second radiomedial cell short,
weakly narrowed towards apex, its length
1.5-1.8 times maximum width, 0.9-1.1
times length of brachial cell. First medial
abscissa rather distinctly S-shaped. Recur-
rent vein 0.9-1.0 times as long as second
abscissa of medial vein. Discoidal cell 1.55-
1.65 times longer than wide. Nervulus
strongly postfurcal, distance from nervulus
to basal vein nearly twice nervulus length.
Parallel vein arising a little behind middle
of distal margin of brachial cell. Hind wing
4.5-4.7 times longer than maximum width.
First abscissa of costal vein 0.85-0.9 times
as long as second abscissa. First abscissa of
mediocubital vein 1.15-1.2 times longer
than second abscissa. Recurrent vein short,
unsclerotized, interstitial, curved toward
base of wing. Legs: Hind coxa large, 1.5-1.6
times longer than wide, 0.7-0.75 times as
long as hind femur. Hind femur wide, 3.1-
3.2 times longer than wide. Hind tibia
thickened towards apex. Hind tarsus 1.1
times longer than hind tibia; hind basitar-
sus 0.6-0.65 times combined length of
second-fifth segments (without pretarsus).
Second tarsal segment 0.4-0.45 times as
long as basitarsus, 1,2-1.3 times longer
than fifth segment (without pretarsus).
First-fourth segments of hind tarsus ven-
trally with wide and transparent flanges,
which are pointed on the tops of each
segment. Metnsomn: 1.7-2.0 times longer
than its maximum width, 0.9-1.1 times as
long as head and mesosoma combined.
First tergite strongly, uniformly and line-
arly widened from base to apex; with small
spiracular tubercles before its middle;
laterally with distinct high and rather wide
carinae; fine dorsal carinae fused in basal
0.3 and then extending to apex as a single,
elevated, median carina; dorsope absent.
Apical width of first tergite 2.4-2.7 times its
basal width; its length 0.6-0.65 times apical
width. Second suture rather distinct and
convex. Second and third tergites with
rather distinct and fine longitudinal medi-
an carina. Median length of second tergite
about half its basal width, equal to or 1.1-
1.2 times length of third tergite. Combined
median length of second and third tergites
nearly equal to basal width of second
tergite, 0.7-0.75 times maximum width of
tergites. Ovipositor sheath (visible part in
lateral view) 1.1-1.3 times longer than first
tergite, 1.0-1.2 times longer than hind
basitarsus, 0.25-0.4 times as long as meso-
soma, 0.15-0.17 times as long as fore wing.
Sculpture mid pubescence: Head very densely
and minutely granulate, face additionally
with rather fine and irregular striation.
Mesoscutum very densely and distinctly
granulate, with rather narrow and long
rugulosity in medioposterior half. Scutel-
lum finely and densely granulate. Meso-
pleuron almost smooth in lower half.
Metapleuron coarsely, regularly and curv-
edly striate for the most part, with fine
granulation between striae and anteriorly.
Propodeum almost entirely coarsely and
rather sparsely striate, striae in areola more
or less transverse and partly undulate or
rugulose, with fine granulation partly;
with distinctly delineated basolateral areas;
areola wide, its length 1.0-1.2 times
maximum width; dorsal carina 0.8-1.0
times as long as areola fork. Hind coxa
smooth; hind femur finely punctulate with
Volume 15, Number 2, 2006
185
very fine granulation dorsally, smooth
ventrally. Metasoma entirely densely
granulate, granulation becoming finer to-
wards apex of metasoma. Mesoscutum
entirely shortly and very densely setose.
Hind tibia dorsally with rather short,
dense and semi-erect setae, its length
0.35-0.55 times maximum width of tibia.
Colour: Head and anterior half of meso-
soma (including mesoscutum) yellow,
posterior part of mesosoma and metasoma
pale yellow, metasoma additionally often
with greenish tint. Antenna reddish
brown or brown, scapus mostly yellow.
Palpi pale yellow. Legs yellow, all tarsi
(especially posterior ones) more or less
brown. Ovipositor sheath brown in basal
half and black in apical half. Fore wing
faintly infuscate. Pterostigma brownish
yellow.
Male. — Body length 2.0 mm; fore wing
length 2.4 mm. Head width 2.1 times its
median length. Transverse diameter of eye
(dorsal view) 8.8 times longer than temple
length if measured on straight line. Anten-
na 28-segmented. Otherwise similar to
female.
Diagnosis. — The new species is very
similar to P. temporalis Belokobylskij from
Brunei (Belokobylskij 1999) and differs in
having the recurrent vein as long as second
abscissa of medial vein, the nervulus
strongly postfurcal, the pterostigma rather
wide, the hind femur wide, the second
tergite short, the face rather finely striate,
and the propodeum almost entirely coarse-
ly rugose-striate.
Host. — Conopomorpha sinensis Bradley
and C. litchiella Bradley (Gracillariidae).
Distribution. — Thailand.
Etymology. — This species is named after
the name of the fruit tree — lychee {Litchi
chinensis Sonn.) — on which their hosts
develop.
KEY TO SPECIES OF THE GENUS PARACHREMYLUS GRANGER
1. Temple longer; transverse diameter of eye (dorsal view) 4.0-5.0 times as long as temple
length. Malar space larger than basal width of mandible. Mesopleuron smooth in upper
half, striation partly present in subalar depression only. 1st— 4th segments of hind tarsus
with narrow and partly indistinct flanges 2
- Temple shorter; transverse diameter of eye (dorsal view) 5.5-7.0 times as long as temple
length. Malar space less than basal width of mandible. Mesopleuron distinctly curvedly
striate in upper 0.4-0.5. lst-4* segments of hind tarsus with wide flanges 3
2. Notauli complete, rather deep posteriorly. Metacarpus 1.2-1.3 times as long as pterostigma.
T1 flagellar segment 3.5-3.7 times as long as apical width. Median length of 2nd and 3rd
metasomal tergites combined a little larger than basal width of 2nd tergite. Propodeum
mostly coarsely and sparsely striate. Body length 2.2 mm. — Madagascar P. seyrigi Granger
- Notauli incomplete, almost absent posteriorly. Metacarpus 1.5 times as long as pterostigma.
1st flagellar segment 3.0 times as long as apical width. Median length of 2nd and 3rd
metasomal tergites combined 1.3 times basal width of 2nd tergite. Propodeum mostlv
smooth. Body length 2.0 mm. — India P. oblougus (Papp)
3. Pterostigma narrow, 5.0 times as long as maximum width. Recurrent vein of fore wing about
twice as long as second abscissa of medial vein. Nervulus not strongly postfurcal,
distance from nervulus to basal vein 0.7 times nervulus length. Hind femur 3.5 times as
long as wide. 2nd tergite 0.6 times as long as its basal width. Face almost entirely distinctly
transversely striate. Propodeum within background areolation sparsely striate, mostly
smooth. Body length 2.3 mm. — Brunei P. temporalis Belokobylskij
Journal of Hymenoptera Research
Pterostigma rather wide, 3.1-3.7 times as long as maximum width. Recurrent vein of fore
wing almost as long as second abscissa of medial vein. Nervulus strongly postfurcal,
distance from nervulus to basal vein nearly twice nervulus length. Hind femur 3.1-3.2
times as long as wide. 2nd tergite about 0.5 times as long as its basal width. Face finely and
partly indistinctly transversely striate and with dense fine granulation. Propodeum
within background areolation almost entirely coarsely rugose-striate with fine granula-
tion partly. Body length 2.0-2.8 mm. — Thailand P. litchii sp. nov.
ACKNOWLEDGMENTS
We wish to thank Dr. S. Moriya (Tsukuba, Japan)
and Dr. S. Dolsopon (Bangkok, Thailand) for giving us
a chance to examine the reared material, and two
reviewers for valuable comments of our manuscript.
The present work was supported by the grant of Japan
Society for the Promotion of Science, Invitation
Fellowship Program for Research in Japan
(No. L05564) and grant 2 P04C 001 28 of the Ministry
of Science and Information Society Technologies
(Poland) for the first author, and a Grant-in-Aid for
Scientific Research (A) (No. 15208007) from the Japan
Society for the Promotion of Science for the second
author.
LITERATURE CITED
Anupunt, P. and N. Sukhvibul. 2005. Lychee and
longan production in Thailand. Acta Horticultural'
(ISHS) 665: 53-60.
Beiokobylskij, S. A. 1993. [On the classification and
phylogeny of the braconid wasps of subfamilies
Doryctinae and Exothecinae (Hymenoptera, Bra-
conidae). Part I. On the classification, 2.] Entomo-
logicheskoe Obozrenie 72: 143-164. [In Russian.]
Beiokobylskij, S. A. 1999. [New taxa of the braconid
subfamily Exothecinae (Hymenoptera, Braconi-
dae) from tropical and subtropical regions of the
Old World. I.] Entomologicheskoe Obozrenie 78:
674-693. [In Russian.]
Beiokobylskij, S. A. and V. I. Tobias. 1998. Fam.
Braconidae. Introduction. Pp. 8-26. in: Lehr, P. A.
ed. OpredeliteV nasekomykh Dal'nego Vostoka Rossii
[Keys to the Insects of the Russian Far East]. 4(3).
Dal'nauka, Vladivostok. [In Russian.]
Bradley, J. D. 1986. Identity of the South-East Asian
cocoa moth, Conopomorpha cramerella (Snellen)
(Lepidoptera: Gracillariidae), with descriptions
of three allied new species. Bulletin of Entomolog-
ical Research 76: 41-51.
Granger, C. 1949. Braconides de Madagascar. Mem-
oires de llnstitut Scientifique de Madagascar. Ser. A,
Biologic Animate 2: 1-428.
Menzel, C. 2002. The lychee crop in Asia and the Pacific.
Food and Agriculture Organization of the United
Nations, Bangkok. 120 pp.
Papp, J. 1990. New braconid wasps (Hymenoptera,
Braconidae) in the Hungarian Natural History
Museum, 1. Annates historico-naturales Musei
nationalis hungarici 82: 175-190.
Papp, J. 1997. New braconid wasps (Hymenoptera,
Braconidae) in the Hungarian Natural History
Museum, 5. Annates historico-naturales Musei
nationalis lunigarici 89: 157-175.
Shenefelt, R. D. 1975. Braconidae 8. Exothecinae,
Rogadinae. Hymoiopterorum Catalogus (nova editio)
12: 1115-1262.
Wharton, R. A. 1993. Review of the Hormiini
(Hymenoptera: Braconidae) with description of
new taxa. Journal of Natural History 27: 107-171.
J. HYM. RES.
Vol. 15(2), 2006, pp. 187-207
Parasitoids (Hymenoptera: Chalcidoidea) of the Cabbage Seedpod
Weevil (Coleoptera: Curculionidae) in Georgia, USA
Gary A. P. Gibson, Michael W. Gates and G. David Buntin
(GAP) Agriculture and Agri-Food Canada, Biodiversity and Integrated Pest Management, K. W.
Neatby Bldg., 960 Carling Avenue, Ottawa, ON, Canada, K1A 0C6; email: gibsong@agr.gc.ca
(MWG) Systematic Entomology Laboratory, PSI, Agricultural Research Service,
U.S. Department of Agriculture, c/o National Museum of Natural History,
Smithsonian Institution, Washington, DC 20560-0168, USA
(GDB) Department of Entomology, University of Georgia, Georgia Station, Griffin, GA 30223, USA
Abstract. — Five families and 13 species of Chalcidoidea (Hymenoptera) were obtained from
mass-reared seedpods of Brassica napus L. (Brassicaceae) as putative parasitoids of the cabbage
seedpod weevil, Ceutorhynchus obstrictus (Marsham) (Coleoptera: Curculionidae), in Georgia, USA.
The species are Conura torvina (Cresson) (Chalcididae), Euderus glaucus Yoshimoto and Necremnus
tidius (Walker) (Eulophidae), Brasema allynii (French) n. comb, (from Eupelmus Dalman) and
Eiipclimis cyaniceps Ashmead (Eupelmidae), Eurytoma tylodermatis Ashmead (Eurytomidae), and
Lyrcus incertus (Ashmead), L. maculatus (Gahan), L. perdubius (Girault), Mesopolobus moryoides
Gibson, Neocatolaccus tylodermae (Ashmead), Pteromalus cerealellac (Ashmead) and Pteromalus sp.
(Pteromalidae). An illustrated key is provided to differentiate the taxa. Lyrcus maculatus constituted
about 96% of all reared Pteromalidae and 86% of the total parasitoid fauna. The associations of B.
allynii, E. glaucus, E. cyaniceps, E. tylodermatis, L. incertus, N. tylodermae, Pteromalus sp. and P.
cerealellac with C. obstrictus are new, but some of these species likely are hyperparasitoids or
emerged from insect contaminants of the mass-reared seedpods. The only previous report of
a parasitoid of C. obstrictus in eastern North America, Trichomalus perfectus (Walker) (Pteromalidae),
is a misidentification. The parasitoid fauna of C. obstrictus in Georgia is discussed relative to that
known for western North America.
The cabbage seedpod weevil, Ceuto-
rhynchus obstrictus (Marsham) (Coleoptera:
Curculionidae), was introduced from Eur-
ope to western North America about
70 years ago. Since then it has become the
most important insect pest of canola and
rape, Brassica napus L. and B. rapa L.
(Brassicaceae), in most areas of the conti-
nent where these crops are grown (Car-
camo et al. 2001, Kuhlmann et al. 2002). It
was first reported from eastern North
America in North Carolina, USA (USDA
1960), and is now known to extend from
Georgia to Quebec and Ontario, Canada
(Brodeur et al. 2001, Mason et al. 2004).
There have been several surveys of the
introduced and native chalcid (Hymenop-
tera: Chalcidoidea) parasitoids of the cab-
bage seedpod weevil in western North
America, including Breakey et al. (1944),
Doucette (1944, 1948), Hanson et al. (1948),
Carlson et al. (1951), McLeod (1953), Walz
(1957), and Dosdall et al. (in press).
Murchie and Williams (1998) listed 7
identified and 4 unidentified species in 9
genera and 5 families of Chalcidoidea as
parasitoids of C. obstrictus in North Amer-
ica, but almost all of the species names
either represent misidentifications or are
now recognized as junior synonyms of
older names (Gibson et al. 2005). Dosdall et
al. (in press) reported another six chalcid
species as reared from B. napus and B. rapa
seedpods in Alberta. Consequently, the
188
Journal of Hymenoptera Research
Table 1. Chalcid parasitoids associated with the
cabbage seedpod weevil in North America, including
for Georgia the number of specimens and percentage
(in parenthesis) of total parasitoids reared by Buntin
(1998).
Western North
Taxon
America
Georgia
Chalcididae
Conura albifrons (Walsh)
+
-
? Conura side (Walker)1
+
-
Conura torvina (Cresson)
+
9 (0.8)
Eulophidae
Eudcrus albitarsis
+
-
(Zetterstedt)
Euderus glaucus
—
2 (0.2)
Yoshimoto
Necremnus tidius (Walker)
+
6 (0.5)
Eupelmidae
Brasema allynii (French)
-
5 (0.5)
Eupelmus cyaniceps
—
4 (0.4)
Ashmead
Eupelmus vesicularis
+
-
(Retzius)
Eurytomidae
Eun/toina ti/lodennatis
+
25 (2.2)
Ashmead
Pteromalidae
Chlorocytus sp.
+
-
Lyrcus incertus (Ashmead)
-
6 (0.5)
Lyrcus maculatus (Gahan)
+
967 (86.0)
Lyrcus perdubius (Girault)
+
60 (5.3)
Mesopolobus bruchophagi
+
-
(Gahan)
Mesopolobus mayetiolae
+
-
(Gahan)
Mesopolobus moryoides
+
2 (0.2)
Gibson
Neocatolaccus tylodermae
-
33 (2.9)
(Ashmead)
Pteromalus eerealellae
-
1 (0.1)
(Ashmead)
Pteromalus spp.2
+
4 (0.4)
Trichomalus lucidus
+
-
(Walker)
' Single record, likely a misidentification of C. torvina
(see text).
2 Females in the two regions represent different
species (see text).
chalcid fauna purportedly parasitizing C.
obstrictus in western North America in-
cludes at least 14 species (Table 1). In
contrast, there is only a single published
report of parasitoids of C. obstrictus in
eastern North America. Buntin (1998)
stated that greater than 96% of the para-
sitoids recovered from seedpods of B.
napus in Georgia were Trichomalus perfectus
(Walker) (Pteromalidae). This species is the
most common biological control agent of C.
obstrictus in Europe (Murchie and Williams
1998) and was long thought to have been
introduced to North America along with
the seedpod weevil. However, Gibson et al.
(2005) snowed that all previous reports of
T. perfectus in western North America were
misidentifications of Trichomalus lucidus
(Walker), another European species.
Accurate identification of parasitoid spe-
cies is a prerequisite for successful classical
biological control and integrated pest man-
agement. The senior author examined the
parasitoids reared by Buntin (1998) as part
of a larger study to document the diversity
and identity of the chalcid parasitoids of C.
obstrictus in North America. The primary
purpose of Buntin (1998) had been to
examine the effect of trap cropping on the
number of seedpod weevils and its para-
sitoids in canola crops in Georgia. The
species identities of the parasitoids had
therefore never been thoroughly investi-
gated. The purpose of this paper is to
provide the first comprehensive informa-
tion on the diversity of the chalcid para-
sitoids reared from canola seedpods in
southeastern USA in order to facilitate
future studies of the parasitoid fauna
associated with C. obstrictus throughout
North America.
MATERIALS AND METHODS
The chalcid parasitoids identified in this
study were obtained from mass-reared
seedpods of B. napus collected from the
Bledsoe Research Farm (33 10.635'N
84°24.354'W) located near Griffin, Georgia,
from 1994-1996, as per "Material and
methods" in Buntin (1998). Although not
stated, the pods were screened for insect
contaminants prior to rearing. Contami-
nants mainly included aphids (Hemiptera:
Volume 15, Number 2, 2006 189
Aphididae) and larvae and pupae of the using Adobe Photoshop™ to enhance cla-
diamondback moth, Plutella xylostella (L.) rity.
(Lepidoptera: Plutellidae). The reared
parasitoids had been stored in ethanol, RESULTS
but were critical-point dried, point- A total of 1,127 specimens of Chalcidoi-
mounted, and identified to genus by the dea were sufficiently intact that they could
senior author using the relevant family be identified accurately. Of these, there
keys in Gibson et al. (1997). The senior was a single male Pachyneuron aphidis
author is responsible for all species identi- (Bouche) and a female and male Asnphes
fications except Eurytoma tylodennatis Ash- suspensus (Nees) (Pteromalidae). Members
mead (Eurytomidae), which was identified of Pachyneuron and Asaphes are obligate
by MWG. Information concerning the hyperparasitoids of aphids (Gibson et al.
method of species identification within 1997) and are not dealt with further. The
each genus is provided under the relevant remaining 1,124 specimens included 5
species discussion. Voucher specimens are families, 10 genera, and 13 species of
deposited in the Canadian National Col- Chalcidoidea that are possible parasitoids
lection of Insects and Arachnids (CNC), of C. obstrictus. These taxa are keyed below
Ottawa, Ontario, the University of Georgia and subsequently treated by family in
Museum of Natural History (UGCA), alphabetical order. The key also segregates
Athens, Georgia, and the United States Trichoinalus Thomson and Chlorocytus Gra-
National Museum of Natural History ham, the only two genera reared from C.
(USNM), Washington, District of Colum- obstrictus in western North America (Dos-
bia. Terms used for parasitoid structure dall et al. in press) that were not recovered
follow Gibson (1997). Photographs are by Buntin (1998) (Table 1). The two taxa
composite serial images that were com- are included in the key because species of
bined using Auto-Montage™. These both genera occur in eastern North Amer-
images and the scanning electron micro- ica and may eventually be reared as part of
photographs were digitally retouched the regional C. obstrictus parasitoid fauna.
KEY TO CHALCIDOIDEA PUTATIVELY PARASITIZING C. OBSTRICTUS IN GEORGIA
1 Hind leg with elongate coxa of similar length to conspicuously swollen and ventrally
toothed femur, and with curved tibia (Fig. 1) . . . Conura torvina (Cresson) (Chalcididae)
Hind leg with comparatively short coxa, slender femur, and straight tibia (Figs 3-8) ... 2
2(1) Tarsi 4-segmented; flagellum with 3 or 4 funicular segments, the segments sometimes
branched (Figs 9, 10) (Eulophidae) 3
Tarsi 5-segmented; flagellum with 5 or 6 unbranched funicular segments (Figs 11-18) ... 4
3(2) Meso- and metatarsi with basal 4 segments white; forewing membrane bare dorsally
behind marginal vein, but with clearly visible row of long admarginal setae
(Fig. 42, ams) on ventral surface near marginal vein; flagellum of both sexes with 4
unmodified funicular segments Euderus gJaucus Yoshimoto
Meso- and metatarsi with basal 1 or 2 segments white; forewing membrane uniformly
setose dorsally behind marginal vein, the setae largely obscuring admarginal setae
on ventral surface (Fig. 41); flagellum branched in male (Fig. 10) and with only 3
funicular segments in female (Fig. 9) Necrcmmis tidius (Walker)
4(2) Head and mesosoma with coarse piliferous punctures and non-metallic, dark brown
to black (Figs 3, 4); pronotal collar quadrangular in dorsal view, only slightly
shorter than mesoscutum; male with elongate petiole and flagellar segments having
whorls of conspicuously long setae (Fig. 4)
190 Journal of Hymenoptera Research
Eiin/toma tylodermatis Ashmead (Eurytomidae)
Head and mesosoma with finer mesh-like sculpture and often with metallic green to
bluish luster; pronotum strongly transverse in dorsal view, conspicuously shorter
than mesoscutum; male with short petiole and flagellar segments having short,
inconspicuous setae 5
5(4) Mesopleuron elongate, convex or cushion-like, and uniformly finely sculptured
(Figs 5, 7); middle leg with strong black spines at apex of tibia and on ventral
surface of tarsal segments (Figs 5, 7, sp), the colour of spines contrasting distinctly
with mostly yellowish leg (Eupelmidae: Eupelminae female) 6
Mesopleuron about as high as long, usually with a smooth region dorsally below base
of wings and often with an oblique femoral depression or groove, but at least not
convex or uniformly sculptured (Figs 23-25); middle leg with slender spines at
apex of tibia and on ventral surface of tarsal segments, the colour of spines not
contrasting with leg 7
6(5) Ovipositor sheaths projecting only slightly beyond apex of gaster and uniformly
coloured (Fig. 5); forewing completely setose behind parastigma and marginal vein
(Fig. 5) Brasema allynii (French)
Ovipositor sheaths projecting beyond apex of gaster by about one-third its length and
medially whitish between darker basal and apical bands (Fig. 7); forewing with
slender, oblique bare band (Fig. 7, bb) below parastigma and base of marginal
vein Eupelmus cyaniceps Ashmead
7(5) Flagellum with only basal segment conspicuously differentiated as strongly transverse
ring segment and with 7 or 8 distinct funicular segments; head in frontal view with
inner margin of eyes distinctly divergent over about ventral half; mesotibial spur
much longer and thicker than metatibial spurs, as long as basal tarsal segment and
about one-third length of tarsus (Eupelmidae: Eupelminae male) 8
Flagellum with 2 or 3 basal segments conspicuously differentiated as ring segments
and then with 6 or 5 tubular funicular segments, respectively (Figs 11-18); head in
frontal view with inner margin of eyes subparallel or slightly but uniformly
incurved; mesotibial spur somewhat longer than, but otherwise similar to,
metatibial spurs, the spur obviously shorter than basal tarsal segment and only
about one-quarter length of tarsus (Pteromalidae) 9
8(7) Forewing completely setose behind parastigma and base of marginal vein (Fig. 6);
hind leg with femur yellowish-white and tibia usually more or less distinctly
brown (Fig. 6); flagellum clavate, the segments widening distinctly to clava and
apical funicular segments transverse (Fig. 6) Brasema allynii (French)
Forewing with large, oblique bare region (Fig. 8, bb) behind parastigma and base of
marginal vein; hind leg with femur and tibia dark (Fig. 8); flagellum robust-
filiform, the segments all about the same width and apical funicular segments
quadrate (Fig. 8) Eupelmus cyaniceps Ashmead
9(7) Flagellum with 5 tubular funicular segments and 3 strongly transverse ring segments
(Figs 11, 13, 14) 10
Flagellum with 6 tubular funicular segments and 2 ring segments (Figs 12, 15-18) ... 14
10(9) Female only: head and mesosoma dark with conspicuous, white, lanceolate setae
(Figs 19, 20, 24, 25); costal cell ventrally with setae only within about apical half of
cell (Figs 43, 46) 11
Female or male: head and mesosoma variable in colour, but with inconspicuous hairlike
setae (Figs 21-23); costal cell ventrally with line of setae extending almost entire
length of cell or at least setae present both basally and apically if line more or less
interrupted medially (Figs 44, 45) 13
11(10) Forewing dorsally setose behind marginal vein over about apical half of vein, the setae
partly obscuring at least 3 rows of ventral admarginal setae apically (Fig. 43);
propodeum with transverse ridge or carina within anterior half dividing it into
anterior and posterior sections on either side of median carina (Fig. 30);
Volume 15, Number 2, 2006 191
metapleuron completely sculptured and with anterior margin on same plane as and
abutting mesopleuron (Fig. 25, am) Neocatolaccus tylodertnae (Ashmead)
Forewing dorsally bare behind marginal vein to level at least equal with middle of
stigmal vein, completely exposing 1 or sometimes 2 partial rows of ventral
admarginal setae (Fig. 46, ams); propodeum sometimes with transverse furrow
near middle, but without transverse ridge (Figs 27, 29); metapleuron partly smooth
anteriorly and with anterior margin either curved outward (Figs 24, 29) or
extending anteriorly above and over posterior margin of mesopleuron (Figs 27, 28) 12
12(11) Propodeum with nucha (Fig. 29, nuc) delineated laterally by longitudinal carina
within furrow along posterior margin; metapleuron with anterior margin (Figs 24,
29, am) curved outwards, extending as thin brown flange almost at right angle to
posterior margin of mesopleuron; lower face without evident malar depression,
evenly convex along oral margin between malar sulcus and clypeus
Lyrcus perdubius (Girault)
Propodeum with nucha not delineated laterally by carina, the furrow along posterior
margin of callus continued uninterrupted mesally and anteriorly so as to delineate,
more or less conspicuously, anterior limit of nucha (Fig. 27); metapleuron with
anterior margin (Figs 27, 28, am) raised above and extending over posterior margin
of mesopleuron; lower face with short but distinct, concave malar depression
(Fig. 20, md) between malar sulcus and clypeus Lyrcus incertus (Ashmead)
13(10) Both sexes: mesonotum usually dark with conspicuous pattern of bluish-green spots,
the spots usually most distinct on mesoscutum paramedially behind pronotum and
laterally on lateral lobe adjacent to notaulus, though small specimens sometimes
brown. Female: gaster lanceolate (Fig. 22); forewing dorsally bare behind marginal
vein to level at least equal with middle of stigmal vein (Fig. 45). Male: flagellum
brown with first funicular segment oblong and much longer than combined length
of the 3 ring segments (Fig. 14); marginal vein strong, but only as thick as width of
stigma and with posterior margin straight, parallel with anterior margin
Lyrcus maculatus (Gahan)
Both sexes: mesonotum metallic green. Female: gaster subcircular (Fig. 21); forewing
dorsally bare behind marginal vein, but apically the setae extending to base of
stigmal vein (Fig. 44). Male: flagellum yellowish with first funicular segment
quadrate to slightly wider than long and at most as long as combined length of the
3 ring segments (Gibson et al. 2005, fig. 8); marginal vein conspicuously thickened
relative to slender stigma and with posterior margin slightly convex (Gibson et al.
2005, fig. 31) Mesopiolobus moryoides Gibson
14(9) Male only: forewing with bare band behind marginal vein extending to level about
equal with middle of stigmal vein, and with 1 or at most 2 partial rows of
admarginal setae (Fig. 46, ams) that are obviously longer than setae on dorsal
surface of disc; metapleuron partly smooth and with anterior margin (Figs 24, 28,
am) curved outward or raised above mesopleuron 15
Male or female: forewing with bare region behind marginal vein less extensive, the
discal setae extending to or almost to base of stigmal vein, and with more than 2
rows of admarginal setae of about same length as setae on dorsal surface of disc
(Figs 43, 47-50); metapleuron completely sculptured and with anterior margin
(Fig. 25, am) on same plane as and abutting mesopleuron 16
15(14) Propodeum with nucha (Fig. 29, nuc) delineated laterally by longitudinal carina
within furrow along posterior margin; flagellum with most funicular segments only
slightly longer than wide, the first segment subquadrate and shorter than pedicel
(Fig. 15); metapleuron with anterior margin (Figs 24, 29, am) curved outwards,
extending as thin brown flange almost at right angle to posterior margin of
mesopleuron Lyrcus perdubius (Girault)
Journal of Hymenoptera Research
Propodeum with nucha not delineated laterally by carina, the furrow along posterior
margin of callus extending uninterrupted mesally and anteriorly so as to delineate,
more or less conspicuously, anterior limit of nucha (Fig. 27); flagellum with all
funicular segments clearly oblong, the first segment as long as pedicel (Fig. 16);
metapleuron with anterior margin (Figs 27, 28, am) raised above and extending
anteriorly over posterior margin of mesopleuron Lyrcus incertus (Ashmead)
16(14) Male only: head and body dark with conspicuous, long, slightly lanceolate white setae
(Fig. 25); propodeum with transverse ridge or carina within anterior half dividing it
into anterior and posterior sections on either side of median carina
(Fig. 30) Neocatolaccus tylodermae (Ashmead)
Male or female: head and body metallic green with inconspicuous hairlike setae
(Fig. 26); propodeum with or without median carina but without transverse ridge
(Figs 35-38) 17
17(16) Pronotum anteriorly with collar rounded into neck, the reticulations extending
uninterrupted from dorsal to inclined surface (cf. Figs 23, 24); forewing with
marginal vein comparatively short, less than 1.5 times as long as stigmal vein
(Figs 49, 50); propodeum with convex, reticulate nucha (Figs 31, 32, nuc), reticulate
panels (Figs 31, 32, pnl), and often without distinct median carina 18
Pronotum anteriorly with shiny, transverse carina differentiating collar from neck
(Fig. 26); forewing with marginal vein obviously (at least 1.5 times) longer than
stigmal vein (Figs 47, 48); propodeum with flat or slightly concave, lunate or
triangular adpetiolar strip (Fig. 35, aps) delineated by inverted Y-shaped median
carina anterior to petiolar foramen or, if with reticulate nucha (Fig. 37, nuc), then
with panels (Fig. 37, pnl) partly strigose (having oblique, irregular, fine carinae or
striae) 19
18(17) Female: costal cell with line of setae on ventral surface interrupted medially (Fig. 49);
scutellum anterior to frenum with reticulations distinctly smaller medially than
laterally (Fig. 33); propodeum with plical carina (Fig. 31, pc) directed obliquely
toward inner margin of spiracle Pteromalus cereahilae (Ashmead)
Female: costal cell with entire line of setae ventrally (Fig. 50); scutellum anterior to
frenum with almost uniform meshlike reticulations (Fig. 34); propodeum with
plical carina (Fig. 32, pc) less strongly angled, directed distinctly mesal of inner
margin of spiracle toward outer margin of basal fovea (Fig. 32, bf) ... Pteromalus sp.
19(17) Metacoxa setose dorsally only over about apical half; forewing of female without setae
on basal fold (Fig. 48); propodeum with inverted Y-shaped median carina
delimiting adpetiolar strip (Fig. 35, aps); propodeum in male without complete
plical carina (Fig. 36, pc) and in female largely bare posterior to spiracle, setose only
from callus to postspiracular sulcus (Fig. 35, pss) Chlorocytus Graham3
Metacoxa setose dorsally over at least apical two-thirds and often completely setose to
base; forewing of female with at least a couple of setae on basal fold (Fig. 47, bf)
differentiating basal cell from speculum; propodeum with or without median
carina but with convex, reticulate nucha (Fig. 37, nuc); propodeum in male with
complete plical carina (Fig. 38, pc) and in female extensively setose posterior to
spiracle, from callus to complete, strong plical carina (Fig. 37)
Trichomalus Thomson3
Genus not yet reported parasitizing C. obstrictus in eastern North America.
Volume 15, Number 2, 2006
193
SPECIES NOTES, ARRANGED
BY FAMILY
Chalcididae
One species of Chalcididae was reared —
Conura torvina (Cresson), which comprised
nine specimens (7 99, 2 33) or 0.8% of the
parasitoid fauna. Delvare (1992) keyed the
species-groups of Conura and differentiat-
ed C. torvina as one of eight species of the
side-group in a key to the "common
species" of that group in North America
north of Mexico. Prior to Delvare (1992), C.
torvina was consistently misidentified as
Conura side (Walker). Carlson et al. (1951)
reported that a specimen of C. side was
reared from C. obstrictus in California. We
were unable to locate this specimen to
confirm the identification, but it is possible
that it is conspecific with the Georgia
species because C. torvina is transcontinen-
tal in North America (Delvare 1992, Noyes
2002). Because of the confusion in names
prior to Delvare (1992), the list of published
distribution and host records given for C.
side by Noyes (2002) certainly contains
many records that actually refer to C.
torvina. Based on previous name usage, of
those Curculionidae listed as hosts of C.
side by Noyes (2002), the record of the
cotton boll weevil, Anthonomus grandis
(Boheman), probably does refer to C. side,
whereas the records of Rhynchaenus palli-
cornis (Say) and Hypera spp. likely refer to
C. torvina. Because of variability in the
colour pattern features given by Delvare
(1992), females of C. torvina can be easily
misidentified as Conura albifrons (Walsh),
another transcontinental species that Dos-
dall et al. (in press) reported parasitizing C.
obstrictus in Alberta. Females of both
species have paramedial yellow marks on
the first gastral tergum, but in female C.
torvina the distance between the marks is,
at most, only about equal to the length of
a mark (Fig. 2). In female C. albifrons the
separation between the marks is at least
similar to the width of a mark, if not
conspicuously greater. Males of the two
species are more easily differentiated.
Males of C. torvina have the interantennal
region and lower face yellow, whereas
males of C. albifrons have the clypeus dark
so that they have a conspicuous, angulate
(A-like), yellow band extending dorsally
between the antennal scrobes.
Eulophidae
Two genera and species of Eulophidae
were reared — Euderus glaucus Yoshimoto (2
99; 0.2% of the parasitoid fauna) and
Necremnus tidius (Walker) (1 9, 5 33; 0.5%
of the parasitoid fauna). Although E.
glaucus was known from Florida and Texas
(Noyes 2002), its association with C. ob-
strictus in Georgia represents a new state
distribution record and a possible new host
record. The only other reported host for E.
glaucus is Epiblema obfuscana (Dyar) (Lepi-
doptera: Tortricidae) (Yoshimoto 1971).
Dosdall et al. (in press) reported a second
Euderus species, E. albitarsis (Zetterstedt),
as an incidental parasitoid of C. obstrictus
in Alberta, but this association was also
obtained by mass-rearing seedpods. Eu-
derus glaucus and E. albitarsis are differen-
tiated in Yoshimoto (1971), though prob-
lems remain in species recognition within
the genus.
Necremnus tidius is a comparatively com-
mon parasitoid of C. obstrictus in western
North America, but it was misidentified as
N. duplicatus Gahan prior to Gibson et al.
(2005), who differentiated and illustrated
the species. The specimens from Georgia
represent the first record of the species in
eastern North America.
Eupelmidae
Two genera and species of Eupelmidae
were reared — Eupelmus (Eupelmus) cyani-
ceps Ashmead (2 99, 2 33; 0.4% of the
parasitoid fauna) and Brasema allynii
(French) n. comb, (from Eupelmus Dalman)
(1 9, 4 Jo; 0.5% of the parasitoid fauna).
Brasema Cameron is unrevised for the
region, but there are about 25 described
species in North America north of Mexico.
194
Journal of Hymenoptera Research
Most of the species are currently misclassi-
fied in Eupelmus (Gibson 1995). Gahan
(1933) described and partly illustrated both
sexes of B. allynii as a parasitoid of the
Hessian fly, Mayetiola destructor (Say) (Dip-
tera: Cecidomyiidae). Phillips and Poos
(1921) also provided both a dorsal and
lateral habitus of the female, and for both
sexes illustrated the colour pattern of the
legs, important species-recognition fea-
tures, when they described the immature
stages of B. allynii as a parasitoid of the
wheat jointworm, Tetramesia tritici (Fitch)
(Hymenoptera: Eurytomidae). The sexes of
Eupelminae are strongly dimorphic (Gib-
son 1995), but the more important di-
agnostic features of B. allynii females in-
clude: head and mesosoma variably brown
or dark with metallic green luster, scrobal
depression finely coriaceous and quite
shiny, lower face with relatively sparse
and only inconspicuously lanceolate white
setae, mesonotum finely coriaceous, and
middle legs entirely or largely yellow
beyond coxae (mesofemur and tibia often
with some light brown infusion but meso-
femur not extensively dark). Males of B.
allynii are in part diagnosed within Brasema
by a clavate flagellum with very short and
inconspicuous setae (Fig. 6), head and
mesosoma metallic green, head with only
very slightly lanceolate and comparatively
sparse white setae, and legs with all femora
yellow (Fig. 6). Brasema allynii is trans-
continental in North America and a po-
lyphagous primary or facultative hyper-
parasitoid of hosts in concealed situations.
Noyes (2002) listed 58 host species in 22
families of Coleoptera, Diptera, Hemiptera,
Hymenoptera, and Lepidoptera, though
the putative host record of C. obstrictus is
the first for Curculionidae.
Gibson (1995) recognized three subge-
nera in Eupelmus, including £. (Episolinde-
lia) Girault and E. (Macroneura) Walker in
addition to the nominate subgenus. Noyes
(2002) listed 45 valid species of Eupelmus in
the Nearctic region, but this includes all
three subgenera and several species in-
correctly classified to genus. Eupelmus is
unrevised for the region, but there are
about 15 described species of E. (Eupelmus)
in North America north of Mexico. Eu-
pelmus cyaniceps belongs to the urozonus
species-group sensu Gibson (1995). Hunter
and Pierce (1912, pi. XVIII, f) provided
a dorsal habitus drawing of the female
when they recorded the species as a para-
sitoid of A. grandis. A species revision of E.
(Eupelmus) that includes evaluation of size-
correlated and host-induced variation is
necessary to confidently characterize spe-
cies limits within the subgenus. However,
females of £. cyaniceps are differentiated
from most other regional species of the
subgenus by the following combination of
features: macropterous, the forewing hya-
line and with a linea calva (Fig. 7, bb),
scape dark, mesosoma dark with metallic
green luster, and ovipositor sheaths ex-
tending for a distance at least equal to two-
thirds length of the metatibia and with
a medial white band (Fig. 7). Because of
extreme sexual dimorphism (cf. Figs 7, 8),
species recognition in Eupelmus is based
almost entirely on females; males are not
characterized for most species. The fea-
tures provided in the key for males of E.
cyaniceps are of family and generic level.
Eupelmus cyaniceps is a primary or faculta-
tive hyperparasitoid of hosts in concealed
situations. The rearing from C. obstrictus re-
presents a new putative host record, but
Noyes (2002) listed 17 other species in 11
different genera of Curculionidae as part of
65 host records in 20 families of Coleoptera,
Diptera, Hymenoptera and Lepidoptera.
The only eupelmid previously reported
as a parasitoid of C. obstrictus is Eupelmus
(Macroneura) vesicularis (Retzius) from
Washington state (Hanson et al. 1948) and
British Columbia (McLeod 1953). This
species likely represents one of the earliest
accidental introductions from Europe to
North America; only females are known in
North America and they are brachypterous
(Gibson 1990).
Volume 15, Number 2, 2006 195
Eurytomidae sitoids or as hyperparasitoids. It is beyond
One species of Eurytomidae was reared the scoPe of this study to assess the
—Eurytoma tylodermatis Ashmead, which monophyly of species-groups or species
comprised 25 specimens (8 99, 17 f$) or limits in Eurytoma, but at least the four
2.2% of the parasitoid fauna. The Georgia Eurytoma species listed above have the
rearing represents a new host record for E. propodeum densely setose lateral to the
tylodermatis, but a previously unidentified propodeal foramen, and the petiole
species of Eurytoma was also reared in most (Fi§- 40' Pt}) has one dorsomedial and
surveys of the parasitoid complex of C. two anterolateral processes. Furthermore,
obstrictus in western North America (Douc- the anterior margin of the first gastral
ette 1948, Hanson et al. 1948, McLeod 1953, tergum (Fig. 40, Gtl) is emarginate and
Dosdall et al. in press). Examination of depressed medially, and the tergum is
voucher and additional reared material deeply depressed anterolateral^, to ac-
from Alberta, British Columbia, Idaho, commodate the processes of the petiole
Oregon, and Washington indicates the when the gaster is raised. Bugbee's (1967)
western species is also £. tylodermatis. keY to species differentiates E. tylodermatis,
Noyes (2002) listed 19 other curculionid in part, by stating the sculpturing of the
species in 11 genera as part of 46 reported fourth gastral tergum extends over the
host species in 14 families of Coleoptera, dorsal surface at least narrowly along the
Diptera, Hymenoptera and Lepidoptera. anterior margin (couplet 20). The species
Over 90 nominal species of Eurytoma are description, however, states that the sculp-
known from the Nearctic region (Noyes ture of the fourth tergum is heavy ventro-
2002). Bugbee (1967) revised the North laterally, continues dorsally for about one-
American species, but species recognition half to two-thirds of the surface, and then
remains extremely difficult because of fades out so that the dorsal surface is
variability of the morphological features smooth and shiny. The extent of sculptur-
he used to differentiate species, and be- ing on the fourth gastral tergum appears to
cause sexual dimorphism (cf. Figs 3, 4) be variable in species of Eurytoma, and the
presents difficulties in recognizing conspe- appearance is partly affected by telescop-
cific sexes. Rearing is necessary to make ing of the terga. Features that can be used
the association, and one sex of several in combination to differentiate £. tyloder-
species of Eurytoma remains undescribed. matis from similar species include the
Bugbee (1967) examined over 4,000 speci- malar space lacking an alveolate boss (a
mens originally identified as E. tylodermatis slightly raised area), the ventrolateral
in the USNM and stated that "even this margin of the scrobes (Fig. 39, vis) being
number was not enough to give an produced anteriorly and reflexed poster-
adequate picture of the geographical dis- iorly, and the median channel of the
tribution, or the range of variation of propodeum being distinct and defined
several species in the complex" (Bugbee laterally by carinae formed by longitudi-
1967, p. 492). He keyed E. tylodermatis as nally aligned crenulae (Fig. 40).
one of 48 species of his "tylodermatis A single species of Eurytoma, E. curculio-
complex" and considered the species to num Mayr, has also been reported as reared
be most closely related to £. bolteri Riley, E. from C. obstrictus in Europe (Dmoch 1975).
diastrophi Walsh, and E. pini Bugbee. He Individuals of E. curculionum have a meso-
also stated that the four species were coxal lamella according to Claridge and
probably associated with the larvae of Askew (1960, fig. 2), which is absent from
weevils and small moths that live in stems the North American specimens identified
of various plants, either as primary para- as E. tylodermatis.
196
Pteromalidae
Three species of Lyrcus Walker, one
species of Mesopolobus Westwood, one
species of Neocatolaccus Ashmead, and
what likely are two species of Pteromalus
Swederus comprised about 96% of the
reared parasitoids (Table 1).
Lyrcus is restricted to the New World.
The genus is unrevised for the Nearctic,
but Noyes (2002) listed 16 species from the
region. Species identifications in this study
are based on examination of type material
of the North American species contained in
the USNM, which excludes the four oldest
names assigned to Lyrcus from the Nearc-
tic. Walker (1847) described four species
collected in Florida that are now classified
in Lyrcus (Noyes 2002) and type material of
these species is in The Natural History
Museum, London. Although Burks (1975)
examined the types, the names have yet to
be placed adequately within a species
concept of Lyrcus. Until this is done within
a comprehensive taxonomic revision, it is
possible that one or more of the four
Walker names represents a senior syno-
nym of a name used in this paper.
Lyrcus maculatus (Gahan) was the most
commonly reared species of all the para-
sitoids, comprising 967 specimens (515 99,
452 (J1) or 5.3%
of the parasitoid fauna. Georgia represents
a new state distribution record for the
species. Dosdall et al. (in press) first reared
L. perdubius from canola seedpods in
Alberta, putatively as a parasitoid of C.
obstrictus, and Noyes (2002) listed Antho-
nomus grandis, A. rutilus (Boheman), A.
signatus (Say), Lixus musculus Say, and
Smicronyx tychoides Le Conte (Coleoptera:
Curculionidae) as other hosts.
Six specimens (3 99, 3 SS) 0.5% of the
parasitoid fauna) of Lyrcus incertus (Ash-
mead) were also reared. This species is
widespread throughout southern and east-
ern USA. Although C. obstrictus represents
a new host association, Noyes (2002) listed
several other genera and species of Curcu-
lionidae as hosts, including A. grandis and
a single report of a Ceutorhynchus sp.
(Pierce et al. 1912).
Only one female and male of Mesopolobus
moryoides Gibson were reared, which rep-
Volume 15, Number 2, 2006
197
resent 0.2% of the parasitoid fauna and the
first distribution record for Georgia and
eastern North America. This species is
a common parasitoid of C. obstrictus, its
only known host, in western USA (Gibson
et al. 2005). Two other species of Mesopo-
lobus have also been reported as putative
parasites of C. obstrictus in western North
America, M. mayetiolae (Gahan) in Califor-
nia (Carlson et al. 1951) and M. bruchophagi
(Gahan) in Alberta (Dosdall et al. in press).
Mesopolobus moryoid.es was misidentified as
Mesopolobus morys (Walker) in North
America until Gibson et al. (2005) correctly
identified it and provided features to
differentiate the two species from each
other and from other regional species.
Mesopolobus is yet another unrevised, spe-
ciose genus in North America, with Noyes
(2002) listing 20 valid species for the
region.
A total of 33 Neocatolaccus tylodermae
(Ashmead) (13 99, 20 SS) were reared,
which represent 2.9% of the parasitoid
fauna. Although Georgia is a new state
distribution record, the species was known
from Florida and is transcontinental in the
USA (Noyes 2002). Ceutorhynchus obstrictus
also represents a new putative host associ-
ation, though Noyes (2002) listed 15 other
curculionid species in 10 genera as hosts.
Pierce (1909) reared it along with E.
cyaniceps from Lixus musculus, Wilson and
Andres (1986) reared it along with L.
maculatus from Rhinocyllus conicus, and
there is a single published association with
Anthonomus grandis (Pierce 1909). Boucek
(1993) provided a key to the three de-
scribed North American species of Neoca-
tolaccus. He differentiated N. tylodermae
from N. moneilemae Gahan on the basis of
a rounded rather than medially carinate
pronotum and truncate rather than medi-
ally narrowly emarginate clypeus. Fore-
wing setal differences also help to differ-
entiate the species. In N. tylodermae the
ventral surface of the costal cell has setae
only over about its apical half and dorsally
the forewing is bare behind about the basal
half of the marginal vein so that three or
four rows of ventral admarginal setae are
visible within a distinct speculum (Fig. 43),
whereas individuals of N. moneilemae have
a line of setae along the length of the costal
cell and the forewing is setose behind the
marginal vein more or less to its base so
that a distinct speculum is lacking and the
admarginal setae are covered by dorsal
setae.
Five individuals (2 99, 3 ^ct; 0.5% of
the parasitoid fauna) of Pteromalus were
reared. One female was identified as
Pteromalus cerealellae (Ashmead) based on
examination of type material in the USNM,
but the other specimens remain unidenti-
fied to species (see below). Girault (1917)
provided a key to several species of
Pteromalus (as Habrocytus Thomson) in
North America, but there is no modern
revision and Noyes (2002) listed 46 valid
species names in the Nearctic region. In
western North America, unidentified spe-
cies of Pteromalus have been reported from
surveys in Idaho (Walz 1957), Washington
(Hanson et al. 1948), British Columbia
(McLeod 1953), and Alberta (Dosdall et
al. in press). Examination of voucher speci-
mens from these studies by the senior
author revealed at least one unidentified
species common to the four areas as well as
a single rearing of Pteromalus puparum (L.)
from the surveys reported by McLeod
(1953) in British Columbia (Gibson et al.
2006). The unidentified species from
western North America and P. cerealellae
belong to a comparatively small group of
Nearctic species whose females have the
line of setae on the ventral surface of the
costal cell interrupted medially and the
bare band behind the marginal vein ex-
tending the length of the vein (sometimes
with one or two setae apically within an
otherwise distinct bare band, Fig. 49).
Females of the two species differ from
each other most conspicuously in propo-
deal features. In P. cerealellae the setae on
the callus extend mesal of the postspira-
cular sulcus posteriorly, almost to the
198
Journal of Hymenoptera Research
plical carina (Fig. 31), whereas females of
the species from western North America
have the region between the postspiracular
sulcus and plical carina bare. The propo-
deal structure of P. cerealellae is very similar
to that of the European species Pteromalus
semotus (Walker) (Graham 1969, fig. 385),
which was reported as reared from C.
obstrictus in England and Poland (Murchie
and Williams 1998). Boucek (1977) once
considered the two names conspecific, but
subsequently (Boucek 1988) re-established
P. cerealellae. Among other features, P.
semotus has an entire costal setal line (cf.
Fig. 50). Pteromalus cerealellae was de-
scribed from, and until recently was
thought to be a monophagous parasitoid
of, the Angoumois grain moth, Sitotroga
cerealella (Olivier) (Noble 1932). Flanders
(1932) stated that it would also oviposit
into the tuberworm moth, Phthorimaea
operculella (Zeller) (Lepidoptera: Gelechii-
dae). However, Brower (1991) showed that
it effectively parasitized 12 different spe-
cies in 4 families of Coleoptera (including 3
species of Curculionidae) that are pests of
stored products. He concluded that al-
though the species probably prefers S.
cerealella, it is more habitat specific than
host specific.
The second Pteromalus female reared and
that of P. cerealellae are similar in having
the forewing dorsally bare behind the
entire length of the marginal vein (Figs 49,
50), but differ in those features given in the
key. A comprehensive generic revision is
necessary to establish the correct species
name of the unidentified female. The three
unidentified males may represent the
opposite sex of the unnamed female, based
on the presence of a continuous line of
setae on the costal cell, but species char-
acteristics of male Pteromalus remain large-
ly unknown and the males are not in-
cluded in the key.
DISCUSSION
The parasitoid fauna reared from B.
napus seedpods in Georgia, 1994 through
1996, revealed the same five chalcid fam-
ilies that have been reported as reared from
C. obstrictus in western North America,
including six species apparently shared in
common (Table 1). Of the shared species,
L. maculatus was by far the most commonly
reared parasitoid in Georgia, comprising
about 86% of the fauna. This contrasts to
western North America where it appears to
be only an incidental parasitoid of C.
obstrictus. Furthermore, two common para-
sitoids of C. obstrictus in at least some parts
of western North America, N. tidius and M.
moryoid.es, were reared as only incidental
parasitoids in Georgia. The latter rearings
represent the first distribution records of
the respective species in eastern North
America. If C. obstrictus was introduced to
Georgia from western North America, the
two parasitoid species may have been
introduced accidentally at the same time.
The most common parasitoid of C. obstric-
tus throughout most of western North
America, T. lucidus, was not reared in
Georgia despite the statement of Buntin
(1998) that most of the reared specimens
consisted of T. perfectus (a misidentification
of T. lucidus prior to Gibson et al. 2005). The
second and third most commonly reared
species in Georgia were L. perdubius and N.
tylodermae, respectively. Because of their
relative abundance and because neither
has been reported from hosts other than
Curculionidae (Noyes 2002), both species
very likely are parasitoids of C. obstrictus.
However, it remains to be determined
whether they are primary or hyperparasi-
toids. At least some of the other incidental
species, such as C. torvina, B. allynii and E.
cyaniceps, likely are hyperparasitoids rather
than primary parasitoids. The rearing of £.
glaucus, B. allynii, E. cyaniceps, L. incertus, P.
cerealellae and the unidentified species of
Pteromalus from B. napus seedpods in
Georgia represent new rearing records,
but these are at most incidental parasitoids,
if C. obstrictus was the actual host for all the
species. Buntin (1998) obtained the para-
sitoids from mass-reared seedpods. The
Volume 15, Number 2, 2006
199
very few Asaphes and Pachyneuron that
were reared, along with several Aphidiinae
(Braconidae) also preserved with the ma-
terial, show that some aphid mummies
contaminated the seedpods even though
an attempt was made to remove these prior
to rearing. Likewise, one or more of the
uncommon parasitoid taxa may have
emerged from other undetected insects
within or on the pods. For example, the
only other host record for E. glaucus is
a lepidopteran. Individual rearing of para-
sitoids dissected from seedpods is neces-
sary to definitively prove the host associa-
tions listed in Table 1, which at present are
only inferred.
Both L. incertus and L. perdubius have
been reported previously as parasitoids of
the cotton boll weevil, as has also E.
cyaniceps, E. tylodermatis, and N. tylodermae ,
though not the most commonly reared
parasitoid of C. obstrictus in Georgia, L.
maculatus. These results suggest that the
chalcid parasitoid fauna acquired by C.
obstrictus in any area where it is introduced
is partly influenced by what other curcu-
lionid species occur in the region. If so, the
parasitoid fauna from eastern Canada and
the southeastern USA might be expected to
differ as substantially as between eastern
and western North America.
ACKNOWLEDGMENTS
The senior author gratefully acknowledges Eric
Grissell (USNM) for access to the USNM chalcid
collection and loan of type and other material, without
which most species identifications would not have
been possible. We thank Jennifer Read (CNC) for
preparing the plates of illustrations used to clarify
species differentiation, and John Huber and Peter
Mason (CNC) as well as two anonymous reviewers
for helpful suggestions regarding improving this
manuscript.
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Figs 1-8. 1 and 2, Conura torvina, female: 1, lateral habitus; 2, metasoma, dorsal. 3 and 4, Eurytoma tylodermatis,
lateral habitus: 3, female; 4, male. 5 and 6, Brasema allynii, lateral habitus: 5, female; 6, male. 7 and 8, Eupelmus
cyaniceps, lateral habitus: 7, female (insert: mesotarsus and apex of mesotibia showing spines); 8, male.
(Abbreviations: bb = bare band, sp = spines.)
Volume 15, Number 2, 2006
203
ilus cerealellae 9
Figs 9-18. Antenna, male and female. (Abbreviations: fu = funicular segment.)
204
Journal of Hymenoptera Research
Figs 19-26. 19, Lyrcus perdubius, head. 20, L. incertus, malar space. 21 and 22, dorsal habitus, female: 21,
Mesopolobus moryoides, 22, L. maculatus. 23-25, mesosoma, lateral: 23, L. maculatus; 24, L. perdubius; 25,
Neocatolaccus tylodermae. 26, Trichomalus perfectus, pronotum and mesoscutum. (Abbreviations: am = anterior
margin of metapleuron, md = malar depression.)
Volume 15, Number 2, 2006
205
Figs 27-34. 27 and 28, Lyrcus incertus: 27, propodeum; 28, metapleuron. 29-32, propodeum: 29, L. perdubins; 30,
Neocatolaccus tylodermae; 31, Pteromalus cerealellae; 32, Pteromalus sp. 33 and 34, scutellar-axillar complex: 33, P.
cerealellae; 34, Pteromalus sp. (Abbreviations: am = anterior margin of metapleuron, bf = basal fovea, nuc =
nucha, pc = plical carina, pnl = propodeal panel.)
206
Journal of Hymenoptera Research
WJffMfl'''
"'"f'M/ifibfiiiP'**'
Figs 35^42. 35 and 36, Chlorocytus sp., propodeum: 35, female, posterior; 36 male; posterolateral. 37 and 38,
Trichomalus lucidus, propodeum: 37, female, posterior; 38, male, posterolateral. 39 and 40, Eurytoma tylodermatis
female: 39, head, frontolateral; 40, propodeum and base of metasoma, dorsolateral. 41 and 42, forewing: 41,
Necremnus tidius; 42, Euderus glaucus. (Abbreviations: ams = admarginal setae, aps = adpetiolar strip, Gtl = first
gastral tergite, nuc = nucha, pc = plical carina, pnl = propodeal panel, pss = postspiracular sulcus, ptl =
petiole, vis = ventrolateral margin of scrobes.)
Volume 15, Number 2, 2006
207
Nebcatolaccus tylodermae (43
■Sr\\<
Mesopolobus moryoides v*3
J:
Lvrcus maculatus (40-'
^-r*
bf
^ Trichoma l us lucidus 4J
m&c-'T,
-^ s. »* v
Chlorocytus sp. (48
J? ■-.•v>" . "~ "* ■» - > N
Pteromalus cerealellae (49.
Figs 43-50. Forewing, female. (Abbreviations: ams = admarginal setae, bf = basal fold, cc = costal cell, mv
marginal vein, sv = stigmal vein.)
J. HYM. RES.
Vol. 15(2), 2006, pp. 208-231
A Survey of the Bees of the Black Rock Forest Preserve, New York
(Hymenoptera: Apoidea)
Valerie Giles and John S. Ascher*
Division of Invertebrate Zoology American Museum of Natural History,
Central Park West at 79,h Street, New York, NY 10024, USA
^Address for correspondence: ascher@amnh.org
Abstract— We present the results of a survey of the bee fauna of Black Rock Forest, Orange
County, New York, USA. The survey focused on bees, with more limited data gathered for other
incidentally collected groups such as apoid and vespid wasps. Surveys in 2003 with nets and bowls
recorded 144 bee species (26 genera), 22 vespid species (9 genera) and 23 crabronid species (12
genera). Noteworthy records are detailed. A preliminary checklist of the bee fauna of the BRF is
presented and discussed in relation to that of New York State, selected sites within the state, and of
the northeastern USA as a whole. The cleptoparasitic species Sphecodes fattigi Mitchell, Sphecodes
johnsonii Lovell, and Lasioglossum (Dialictus) michiganense (Mitchell), and the oligolectic species
Osmia (Melanosmia) inermis (Zetterstedt) are newly recorded from New York State. Ecological
patterns pertaining to sociality, nest type, pollen specialization, parasitism, and phenology, are
summarized and discussed, as are the efficacies of different collecting methods. The net collected
sample was richer than the bowl trapped sample in total bee species (117 vs. 113) and in the number
of unique species (29, 20.4% vs. 25, 17.6%).
Key words. — native bees, Bombus, Andrena, Apidae, invertebrate survey, invertebrate biodiversity,
bowl trap, pan trap, trap nest, Black Rock Forest, pollination
Bees (Hymenoptera: Apoidea) are the T. Griswold, unpublished; cf. Hurd 1979).
single most important animal pollinators of Published data on the distribution of bees
both native and cultivated vegetation in in New York State (NY) is limited (e.g.,
most habitats worldwide (Williams et al. Leonard 1928), but Ascher (unpublished
2001, Michener 2000). The mutualist re- information) has compiled a list of 423
lationship between bees and plants forms species known from New York, of which
a key process in the maintenance of both 405 are native to North America,
local biological diversity and agricultural Single-site inventories of poorly known
productivity. As primary pollinators, bees invertebrate groups have the potential to
provide a vital ecosystem service, affecting establish useful quantitative baseline esti-
the integrity of ecological communities as mates of local biodiversity, as well as to
a whole, including the health of humans help illuminate large-scale distributional
(Williams et al. 2001, Nilsson 2000, Cane patterns within those groups. Such esti-
and Tepedino 2001). Despite this ecological mates can prove useful in testing hypoth-
importance, our understanding of some eses arising from practices as diverse as
basic aspects of bee biology, including theoretical biogeography and conservation
species level distributional patterns, re- planning. In addition, geo-referenced spec-
mains incomplete. There are about 20,000 imen data are amenable to re-analysis and
species of bees worldwide (Michener 2000) comparison with related data sets in the
and approximately 3500 described species context of regional studies of biodiversity
occur in America north of Mexico (JSA and across diverse groups. Finally, such in-
Volume 15, Number 2, 2006
209
ventories help to address the need for
natural history information that is crucial
for understanding community-level eco-
logical patterns (e.g. phenological patterns,
host associations, habitat use, etc).
Black Rock Forest (BRF) harbors a variety
of distinct habitat types, many of which are
typical of the larger Hudson Valley Region,
and is managed in part as a long-term
research preserve. Because the landscape
matrix surrounding BRF is under increas-
ing pressure from land conversion and
habitat degradation, a survey of the bee
communities of BRF while surrounding
habitats are still relatively intact should
provide a valuable basis for future com-
parison with a variety of other sites across
a range of spatial and temporal scales. The
effects of environmental change on bee
communities remain insufficiently under-
stood. Many relevant studies have been
published (see, e.g., Matheson et al. 1996),
but few of these are from eastern North
America. Cane (2005) notes that bees
"possess a unique combination of salient
foraging and nesting traits that together set
them apart from other taxa studied in the
context of habitat fragmentation". Many
bees utilize open areas for foraging and
nesting, and may benefit from forest
fragmentation, unlike forest-dwelling
songbirds. However, bees are still poten-
tially vulnerable to habitat change, partic-
ularly the loss of their host plants.
RESEARCH OBJECTIVES
This survey was undertaken with two
primary goals; first, to assemble a faunal
list of the bees (and selected aculeate
wasps) of BRF to serve as a baseline
inventory of use to both ecologists and
conservation biologists, and second, to
compare BRF data with other bee samples
in order to shed light on larger-scale
(regional) patterns of bee distributions
and diversity. Second order objectives in-
cluded enhanced representation of the
regional bee fauna in the collection of the
American Museum of Natural History
(AMNH) and creation of a synoptic (taxo-
nomic reference) collection to be housed at
BRF. Ecological data were also gathered,
such as abundance of bees across the
season and on selected host plants, and
the efficacies of various collecting methods
(net collecting vs. trapping with bowls of
three different colors vs. trap nesting) were
tested.
STUDY SITE AND METHODS
The Black Rock Forest (BRF) is a 1520
hectare preserve and research facility lo-
cated in Orange County, New York (Lat.
41.42267, Long. 74.03039), ca. 50 miles
north of New York City (NYC). The BRF
is situated within the highest portion of the
Hudson Highlands. The terrain comprising
the preserve ranges in elevation from about
135 m to 446 m. A network of closed
canopy dirt roads permits access to within
1 kilometer of any point within the BRF.
The landscape, both within the forest pre-
serve, and across several large adjacent
tracts (including West Point Military Acad-
emy), is mostly forested (upland hardwood
forests dominated by Quercus spp., Barrin-
ger and Clemants 2003). Other local habitat
types include successional hardwood
stands, hemlock coves, chestnut-oak
woods, red maple swamps, ponds, reser-
voirs, and marshes. Important habitats for
bees at BRF include small meadows,
exposed road edges and reservoir edges,
dams, and marshes, where flowering
shrubs (such as Viburnum spp., Ilex verti-
cillata (L.) A. Gray, Kalmia latifolia L.,
Clethra alnifolia L., Spiraea spp., Rhododen-
dron spp.), and herbs (such as Veronica,
Polygonum spp., Gnaphalium, Solidago spp.,
and Viola spp.) provide seasonal sources of
pollen and nectar to bees. In addition,
during early spring (April) prior to leaf-
out, forest habitats hosted bees attracted to
flowering trees such as Acer rubrum L.,
Salix spp. including S. discolor Muhl. and
Prunus spp. In May, other flowering trees
such as Craetaegus macrosperma Ashe, and
other Prunus spp. were important re-
210 Journal of Hymenoptera Research
sources for many Andrena and other bee one of three florescent colors: yellow, white
species. The early spring flower Erythro- or blue. A total of 150 traps were deployed
nium americanum Ker-Gawler flowered on each of 17 survey visits and arrayed in
sparingly during our survey season and ten transects on each visit. Each transect
was visited by relatively few bees. Vacci- consisted of 15 traps (five of each color),
nium species, especially highbush blueber- arrayed in alternating colors. Traps con-
ry Vaccinium corymbosum L. and lowbush tained a solution of Dawn brand blue
blueberry V. august ifloiwn Ait. dominated dishwashing liquid (1 table spoon to 1
large areas of the forest understory at BRF, gallon tap water) and were placed in ten
including dry hillsides, damp forest areas, sites on the ground along transect lines,
open forest gaps created by fires, and wet Traps were deployed over a period of
marsh edges in association with herba- approximately 1 hour beginning at 0730 hr
ceous communities. Beginning in May, and and were in place before 0900 hr on survey
continuing into late June, Vaccinium stands visits during which they were used. In-
composed of several species were visited dividual traps were placed at approximate-
by large concentrations of nectaring and ly one meter apart. Transect sites were
"buzz" pollen-collecting bees. Vaccinum chosen opportunistically and included:
stamineum L. (Deerberry) was moderately open fields, roadsides, reservoir edges,
common in hillside forests. Patches of dams, forest floors and stone outcroppings
Lysimachia were noted. throughout the BRF property. At the close
The survey season during the spring and of each survey visit the traps were re-
summer of 2003 was generally wet and trieved during a two-hour period begin-
cool in southern New York as confirmed ning at approx. 1600 hr. The contents were
by weather data collected at BRF. Above poured through sieves and the recovered
average rainfall and below average tern- specimens were transferred to plastic
peratures would be expected to depress whirl-packs containing 75% ethyl alcohol,
bee numbers and collecting success. Locality data and bowl trap color labels
Sampling schedule. — We conducted bee were recorded,
surveys at BRF during 24 days between 31 Hand netting of bees was conducted
March 2003 and 16 October 2003. Each between 09:00 and 16:00 during 23 survey
survey day began at approximately 0730 hr visits. Collecting by hand-net was under-
and was completed generally between taken opportunistically at sites where bees
1800 hr and 1900 hr. Most fieldwork was were thought to be concentrated. Hand
conducted on days with predominantly netting was pursued most intensively in
sunny skies and warm temperatures. Col- exposed sunny habitats such as fields, road
lection sites visited per sampling day and edges, reservoir and marsh edges, where
the time spent at each site varied. In many shrubs and herbaceous perennials
addition, individual collecting sites were bloom and where bees were most likely to
chosen throughout the BRF property op- occur. When bees were captured they were
portunistically in response to the presence transferred to cyanide killing jars before
of bees or abundance of flowering plants, being stored in vials. Vials were labeled
UTM coordinates were recorded for all and placed in a cooler for transfer to the
sites where bees were collected. laboratory.
Sampling methods.— We collected bees at Twenty wooden 'Binderboard' brand
BRF using 3 principal methods: colored trap-nests were deployed for the duration
plastic pan (or bowl) traps, hand-held of the survey beginning on 27 May 2003.
insect nets, and wooden trap nests. Bowl Ten trap-nests consisted of a wooden block
traps were made from 6 oz. plastic Solo bearing 39 holes, each measuring 5.5 mm
brand bowls that were spray-painted with diameter, and a depth of 10 cm. The
Volume 15, Number 2, 2006
211
remaining 10 trap-nests were similar in
other respects, but each bore 21 holes
measuring 5 mm in diameter, drilled to
a depth of 16 cm. Each hole was lined with
a kraft paper tube to facilitate recovery of
specimens. Trap-nests were mounted in
sets of two, at 10 sites dispersed across the
BRF property. Each nest was hung from
a tree limb approximately 1.5 meters above
the ground with the holes oriented to face
south. Trap nests were checked periodical-
ly to determine if any Hymenoptera had
inhabited the holes and to ensure that they
were intact and undisturbed. Trap nests
were retrieved from BRF on 20 March 2004,
and each trap-nest was examined in the
laboratory for evidence of occupation by
Hymenoptera.
Specimens were sorted, mounted, and
identified to species by the authors (initial-
ly sorted by VG; species determinations
then made or confirmed by JSA) except the
more difficult metallic Lasioglossum (Dialic-
tus) females, determined by S. Droege,
Vespidae, determined by J. M. Carpenter,
and the more difficult Crabronidae, de-
termined by P. Gambino. S. Droege made,
confirmed, and revised identifications for
numerous Nomada, and L. Day made and
confirmed identifications of Bombus sauder-
soni and B. vagnns. Vouchers are deposited
in the collection of the American Museum
of Natural History (AMNH). A synoptic
collection is housed at the BRF research
facility. Duplicate specimens were dis-
persed to various bee specialists.
Comparative data on the North Ameri-
can bee fauna as a whole, and on the fauna
of New York State (NY), and of particular
areas within NY, were compiled by JSA
based on study of relevant taxonomic and
faunistic literature and study of historical
insect collections, especially those housed
at: AMNH; Cornell University (CUIC);
New York State Museum; National Muse-
um of Natural History; University of
Connecticut Insect Collections, Storrs; and
Parker Gambino's personal collection (af-
filiated with the AMNH). Recent collec-
tions from across NY and from nearby
states such as Connecticut were available,
including material collected by the authors,
P. Gambino, S. W. T. Batra, K. N. Mag-
nacca, B. N. Danforth, D. L. Wagner, R. G.
Goelet and their associates. All discussion
of the past and present status and life
histories of bee species found at BRF is
based, in part, on these historical and
recent collections and the literature in
addition to the sample obtained during
the survey of BRF. Totals cited for "south-
ern New York" are for the area encom-
passing New York City (NYC), Long
Island, and all counties adjacent to Orange
County (i.e. the southeast portion of the
state north to Sullivan, Ulster, and Dutch-
ess Counties).
RESULTS AND DISCUSSION
The survey collected and individually
databased 6,542 bee specimens represent-
ing 144 species, of which 138 are native and
six are exotic (Appendix 1), 26 genera, and
five families (Table 1). Of these, several
records detailed below represent notable
range extensions, the only recent known
collection of a species in NY, or otherwise
fill gaps in the known distributions of New
York State bees. Other aculeate specimens
incidentally sampled included 22 vespid
species (9 genera), 24 crabronid species (13
genera), and 2 species of Isodontia (Spheci-
dae sensu stricto) (Appendix 2). Only a sin-
gle bee, an Osmia cornifrons female, em-
erged from our trap nests. These were
occupied primarily by eumenine (Vespi-
dae) and Trxfpoxylon {Trypoxylon) (Crabro-
nidae) wasps and were not examined in
detail due to the lack of bees.
Species totals. — Of the 144 bee species
found at BRF we regard 138 as native to
North America (Appendix 1). These are
discussed first followed by the six species
known or suspected to have been intro-
duced deliberately or accidentally to North
America from Europe or East Asia.
Native bee species. —
212
Journal of Hymenoptera Research
Table 1. Summary of the number of described bee species for each genus known from New York State, with
totals for the Eastern USA (sensu Mitchell 1960, 1962), New York State (NY), southern NY as defined in the text
(SNY), Black Rock Forest (BRF), New York City (NYC; i.e. the five boroughs), and Ithaca (within city limits;
many additional species are known from the Ithaca vicinity in Tompkins Co.). The number of species not native
to North America (i.e. adventive and introduced species) is given in parentheses following the total number of
species. *No recent records.
Superfamily Apoidea: Clade Anthophila (Bees)
EUSA
NYS
SNY
BRF
NYC Ithaca
Family Colletidae:
Subfamily Colletinae
Colletes
Subfamily Hylaeinae: Tribe Hylaeini
Hi/laeits
Family Halictidae
Subfamily Rophitinae: Tribe Rophitini
Dufourea
Subfamily Halictinae
Tribe Augochlorini
Augochlorella
Augochlora
Augochloropsis
Tribe Caenohalictini
Agapostemon
Tribe Halictini
Subtribe Sphecodina
Sphecodes
Subtribe Halictina
Halictus
Lasioglossum
Family Andrenidae
Subfamily Andreninae: Tribe Andrenini
Andrcna
Subfamily Panurginae
Tribe Calliopsini
Calliopsis
Tribe Protandrenini
Pseudopanurgus
Tribe Panurgini
Subtribe Panurginina
Panurginus
Subtribe Perditina
Perdita
Family Melittidae
Subfamily Melittinae
Tribe Macropidini
Mncropis
Tribe Melittini
Melitta
Family Megachilidae
Subfamily Megachilinae
Tribe Anthidiini
Anthidiellum
Anthidium
Paranthidium
35
24(3)
34
5
114(2)
125(1)
4
3
17
14(2)
25
8(2)
15
0
0
8(3) 10(2)
3
1
1
1
1
1
1
1
1
1
1
1
3
1
1
1
1
1
8
3*
0
13
4 3 3 3 3
67(2) 39(1) 29(1) 25(1) 44(2)
87(1) 70(1) 40(1) 58(1) 66(1)
3
1
1
1
1
1
15
4
2
1
1
1
3
1
0
0
0
0
27
7
1
0
3
2
2
1
2
1
1
0
0
1
4(2)
2(2)
2(2)
KD
2(2)
2(2)
1
1
1
0
0
1
Volume 15, Number 2, 2006
213
Table 1. Continued.
Superfamily Apoidea: Clade Anthophila (Bees)
EUSA
NYS
SNY
BRF
NYC
Ithaca
Stelis
15
6
2
2
1
4
Tribe Osmiini
Chelostoma
3(2)
3(2)
2(1)
0
1
3(2)
Heriades
3
3
1
1
2
1
Hoplitis
8(1)
6(1)
3
2
1
4
Osmia
30(4)
20(3)
12(3)
10(2)
5(3)
13(2)
Tribe Megachilini
Megachile
43(5)
22(4)
17(2)
7(2)
16(4)
13(2)
Coelioxys
25
12
7
2
5
9
Family Apidae
Subfamily Xylocopinae
Tribe Xylocopini
Xylocopa
2
1
1
1
1
1
Tribe Ceratinini
Ceratina
4
3
3
2
3
3
Subfamily Nomadinae
Tribe Nomadini
Nomada
76
47
25
18
19
26
Tribe Ammobatoidini
Holcapasites
3
2
1
0
0
2
Tribe Epeolini
Epeolus
21
7
4
0
1
3
Triepeolus
26
7
1
0
1
2
Subfamily Apinae
Tribe Osirini
Epeoloides
1
1*
1*
0
1*
0
Tribe Emphorini
Ptilothrix
1
1
0
0
1
0
Tribe Eucerini
Eucera
7
1
1
0
0
0
Melissodes
27
12
6
2
9
8
Pcpounpis
1
1
1
0
1
1
Svastra
5
1
0
0
1*
0
Tribe Anthophorini
Anthophora
6(1)
4
3
1
3
3
Habropoda
1
1
0
0
0
0
Tribe Bombini
Bombus
21
18
15
8
12
15
Tribe Apini
Apis
1(1)
KD
1(1)
KD
KD
KD
TOTALS:
743(22)
423(18)
269(13)
144(8)
210(15)
274(15)
Colletidae
We collected only two species of Colletes,
C. compactus and C. simulans, neither of
which was numerous. This scarcity of
individuals and species (vs. the nine
known from southern NY; see Table 1)
may reflect the low frequency with which
Colletes is captured in bowls (S. Droege, T.
Griswold, pers. comm.) and perhaps a scar-
city of appropriate sandy nest banks in the
vicinity of the sampling sites. Absence of
Colletes inaequalis Say in net-collected sam-
ples from early spring is surprising, as this
is a conspicuous and locally abundant
species across much of the northeastern
USA and is often encountered as it collects
pollen from maples (Acer spp.)/ which are
numerous at BRF.
214 Journal of Hymenoptera Research
Our sample of Hylaeus, including small below), both widely distributed across NY,
series of only two ubiquitous Hylaeus spp., and by two species, L. acuminatum and L.
H. mesillae and H. modestus, is also im- fuscipenne, restricted to eastern NY (e.g.,
poverished. At least four additional species absent from the Fingerlakes Region; see
are abundant in nearby Putnam County range maps in McGinley 1986).
(JSA and P. Gambino, unpublished) and Two widely distributed, pollen-general-
should occur at BRF. ist species of carinate Lasioglossum (Evy-
laeus) (sensu Michener 2000) were collected
Halictidae (L cinctipes and L. quebecense) in addition to
All three augochlorine species known the more localized L. (Evylaeus) nelumbonis.
from NY are numerous at BRF. The The latter seems to be strongly associated
abundance of Augochlora pura in our with aquatic emergent flowers. In our
samples probably reflects the local avail- study, numerous L. nelumbonis were col-
ability of rotting logs in which this species lected in pan traps placed along a causeway
excavates its nests. The most abundant bee bisecting Jim's Pond, in which grew abun-
in our sample, Augochlorella aurata (1,222 dant Nymphaeaceae (Nymphaea odorata).
individuals collected) is a eusocial, Museum label data suggest that L. nelum-
ground-nesting species that is numerous bonis may be a pollen-specialist of Nym-
across most of eastern North America, phaeaceae and /or Nelumbonaceae, but
Populations of this species from northeast- direct observations of pollen collecting
ern USA and southern Canada were behavior by this species have not yet been
known as A. striata (Provancher) prior to made due to the difficulty of observing and
recent synonymy with A. aurata in Coelho's collecting bees on aquatic vegetation.
(2004) revision of Augochlorella. Lasioglossum (Dialictus) individuals were,
The two most common species of Aga- as expected, particularly abundant in our
postemon in NY (A. sericeus and A. virescens) bowl samples. These were found to belong
were collected, but two species present to 22 identified species (additional, poorly
more locally in southern NY [A. texanus known species may be included among our
Cresson and A. splendens (Lepeletier)] were undetermined metallic Dialictus; most
not found. Absence of A. splendens is not males of this subgenus were not deter-
surprising, as this species seems to be mined) including two socially parasitic
associated with sandy nesting substrates. species (Paralictus sensu Mitchell 1960)
The cleptoparasitic genus Sphecodes was and two black, non-metallic Dialictus spe-
represented by S. galerus, S. levis, S. fattigi, cies ( = noncarinate Evylaeus; see Michener
and S. johnsonii, the last two recorded for 2000). Among the identified species of
the first time in NY (JSA has also collected metallic, pollen-collecting Dialictus collect-
S. johnsonii in Fairfield County, Connecti- ed in BRF (i.e., Dialictus sensu Mitchell
cut, new state record). Two additional 1960) the wood-nesting species L. coeru-
Sphecodes species, S. atlantis Mitchell and leum, L. cressonii, and L. oblongum were each
S. dichrous Smith, not found at BRF were numerous. Other notable metallic, pollen-
collected elsewhere in Orange County in collecting Dialictus species include two
1962 (Tuxedo Park vicinity; AMNH). species typical of northern forests (L.
Three Halictus species ubiquitous in the nigroviride and L. versans), a distinctive
eastern United States were found in good species often found in sand pits (L. hctero-
numbers, but the more precinctive H. gnathum), and an infrequently recorded
parallelus Say was not collected. species (due in part to identification diffi-
Lasioglossum sensu stricto was represent- culties) previously known in NY from a few
ed by L. coriaceum and L. leucozonium specimens collected in or near the lower
(regarded for the first time as exotic, see Hudson River Valley (L. cattellae). Single
Volume 15, Number 2, 2006 215
females of the two socially parasitic Dia- The long malar space of A. rufosignata, in
lictus species were collected, one of which, comparison to its likely sister species
L. michiganense, has previously been re- Andrena mandibularis (LaBerge 1980), may
corded in the literature solely from Mitch- be an adaptation to collecting nectar from
ell's (1960) unique holotype female, col- the bell-shaped corollas of Vaccinium spe-
lected in Wayne County, Michigan, in 1940. cies. Another oligolectic Andrena collected,
Our single female specimen and another A. cornelli, is now thought to be a Rhodo-
female collected 30 June 2004 at the inlet to dendron specialist based on field observa-
Lake Myosotis, Edmund Niles Huyck Pre- tions by JSA in Virginia, label data for
serve, Rensslaerville, Albany County, NY, newly identified material in museum col-
by JSA and C. J. Daley are the first records lections, and the widely spaced scopal
outside of Michigan. Despite a lack of hairs of this species that can be considered
published records, this species is probably an adaptation that holds Rhododendron
widely distributed across the northeastern pollen connected by viscin threads
USA. It was recently found in Maryland (S. (Ascher, unpublished; cf. LaBerge 1980).
Droege, pers. comm.; new state record) and Andrena violae, an oligolege of Viola, pos-
southern Ontario, Canada (L. Packer, pers. sesses elongate maxillary palpi used to
comm; new Canadian record). The male of extract nectar from its host. Viola is other-
this species remains unknown. The female wise most often visited by long-tongued
of L. michiganense possesses a conspicuous, bees such as Osmia that are able to reach its
inner, subapical mandibular tooth, where- concealed nectaries. Andrena violae is nu-
as the mandibles of other parasitic female merous across much of the eastern United
L. (Dialictus) are simple (i.e., lack an inner States, excluding the colder areas of the
tooth) with elongate slender tips. The other northeast, but was previously known in
socially parasitic Dialictns found at BRF, L. NY solely from a single male collected at
cephalotes, has recently been found in NYC Van Natta's Dam, Six Mile Creek, Ithaca,
in Central Park, Prospect Park (JSA, new Tompkins County, 2 May 1936 (specimen
records), and the Bronx (collected by P. examined, CUIC). This species was not
Gambino). represented among collections made on
Viola at this site and elsewhere in the
Andrenidae Fingerlakes Region by JSA during 1997-
Our BRF sample included 40 species of 2002, so evidence of its persistence in NY at
Andrena but is still far from complete, as an a new station of occurrence is welcome,
additional 32 species known from southern Other oligolectic Andrena at BRF include A.
NY were not recorded. Our sample was krigiana, a specialist of Krigia (dwarf dan-
rich in vernal species characteristic of delion), and A. fragilis, a specialist of
northeastern forests such as A. imitatrix Cornus (Svida). Three Andrena specialists
and A. nivalis. Species associated with of Solidago and Aster (tribe Astereae) were
blueberry were particularly well represent- found, A. hirticincta, A. nubecula, and A.
ed including the Vaccininm oligoleges simplex (but not its sister species, A. placata
(pollen specialists) A. hradleyi and A. Mitchell, which has been collected recently
Carolina, and the polylectic A. carlini and in Putnam Co., NY), as was the panurgine
A. rufosignata. The last species is abundant Astereae specialist Pseudopanurgus andre-
(but under-collected; cf. LaBerge 1980) in noides [we recognize genus Pseudopanurgus
northern blueberry bogs, and evidently in the broad sense of Mitchell, 1960,
reaches the southern limits of its range at including Protandrena {Heterosarus) and P.
or near Black Rock Forest, as it is unknown (Pterosarus) of Michener, 2000]. Andrena
from New York City, Long Island, and arabis is a specialist of Brassicaceae that
elsewhere along the mid-Atlantic coast, may actually benefit from spread of in-
216
Journal of Hymenoptera Research
vasive Garlic Mustard Alliaria petiolata
(Bieb.) Cavara & Grande. Many species
that regularly collect pollen from rosaceous
trees and shrubs, and are known or
suspected to be important pollinators of
apples, were collected in good numbers,
including A. miserabilis, A. (Melandrena)
spp., and A. (Trachandrena) species. An-
drena (Trachandrena) nitda was numerous at
BRF, which is near the northern edge of its
range in NY (see map in LaBerge 1973).
Melittidae
Although a deliberate effort was ex-
pended to locate and collect from Vaccinum
stamirieum, the host plant of Mclitta eick-
worti Snelling and Stage (1995), this re-
cently described species was not recorded
during our survey. However, it has been
collected nearby in Putnam County by P.
Gambino, as has O. virga Sandhouse,
another poorly known oligolege of Erica-
ceae (see Cane et al. 1985; they recorded O.
virga, as O. "felti", collecting "surprisingly
pure" loads of Deerberry pollen; this
species also uses other ericaceous hosts,
M. Arduser pers. comm.). No Macropis
were collected in this study although their
host plant Lysimachia was present.
Megachilidae
Native megachilid species collected at
BRF included the cleptoparasites Stelis
(Dolichostelis) louisae (one female) and Stelis
(Stelis) nitida (one female). The former is
a colorfully marked parasite of native resin
bees in subgenus Megachile (Chelosto-
moides), including M. (C.) campamdae (the
likely host in NY and New England),
which reaches its northern distributional
limits in southern New York. Stelis nitida
was described in 1878 from specimens
collected in Canada and NY, but there
have been few subsequent collections from
eastern North America. It is most likely
a northern and montane species that
parasitizes Osmia, or possibly large Hoplitis
species. Our sample of eight native Osmia
species includes series of the forest-associ-
ated O. bucephala and O. pumila. We
collected single specimens of three species
that are scarce or absent in other recent
collections from New York State, O. collin-
siae, O. felti, and O. inermis. The last species,
a probable oligolege of Ericaceae (M. S.
Arduser, pers. comm.) previously unre-
ported from NY, has also been identified
among recent samples of bees from the
Adirondacks (JSA and W. L. Romey, new
record). Another Osmia species, O. dis-
tincta, has been found elsewhere in NY
(e.g., South Hill Swamp, Ithaca, Tompkins
County; and along the Hudson River) and
in Pennsylvania to visit Penstemon, includ-
ing P. digitalis Nutt. The tuft of curved
hairs on the ocellar region of this species
would seem to be an adaptation for
collecting pollen from Penstemon, although
O. distincta is apparently not a strict oligo-
lege of this genus (M. Arduser, pers.
comm.).
Our sample of native Megachile and
associated Coelioxys cleptoparasites is im-
poverished, perhaps reflecting the ineffi-
ciency of bowl traps for capturing these
strong-flying species (although Megachi-
lini can be trapped in numbers in bowls of
appropriate color, S. Droege, pers. comm.).
The species captured are widely distribut-
ed and numerous across New York, ex-
cepting M. montivaga, which is known in
the state from a few collections in southern
NY (e.g., recently collected at Edmund
Niles Huyck Preserve in Albany County).
A report of this species from Ithaca
(Leonard 1928) is based on a misidentified
M. inermis.
Apidae, excluding bumble bees
Large and small carpenter bees were
represented respectively by Xylocopa virgi-
nica (locally very numerous at BRF, but
most uncollected) and by two abundant
sister species of Ceratina (Zadontomenis), C.
calcaratn and C. dupla, that cannot be
distinguished in the females. The related
C. strenna Smith is also common in NY but
was not collected.
Volume 15, Number 2, 2006 217
All 18 identified species belonging to the size and long flight season (JSA has
ruficornis group of Nomada (=Nomada s.str.) observed males flying as late as November
in our samples are known or suspected to 10 in Ithaca, NY, a colder locality than BRF)
be cleptoparasites of Andrena species, characteristic of this species. Other bumble
Commonly encountered species of this bee species encountered include B. per-
group at BRF and other forested areas of plexus and B. vagans, both generally nu-
the northeastern USA and southeastern merous in New York forests and bogs, and
Canada include the large, conspicuous the widely distributed B. bimaculatus and B.
species N. luteolbides (a valid species griseocollis. We found few B. vagans, but the
distinct from N. sulphurata Smith; see extremely similar (and thus infrequently
Schwarz and Gusenleitner 2004) and N. identified) B. Sanderson i was found in
maculata, both cleptoparasites of large surprisingly large numbers, including se-
Andrcna belonging to the subgenus Mclan- ries of queens, males, and workers. Two
drena (Milickzy and Osgood 1995), and N. Bombus ternarius were found. This is
bella, a cleptoparasite of A. imitatrix. A new a species of northern affinities found
probable host association between N. bella commonly south to the Catskills. Leonard
and A. imitatrix was inferred by JSA (new (1928:1031-1032) regarded it as, "Essential-
information) based on repeated co-occur- ly a Canadian and northern transition
rence of these species at several sites species...", and stated that "the species is
across several years. Females of N. bella not found near NYC. (Beq) [indicating J.
have been identified (M. Schwarz, pers. Bequaert as the source]." Long-tongued
comm.) but remain undescribed. Further bumble bee species belonging to subgenera
study of Nomada with bidentate mandibles Fervidobombus [B. fervidus (Fabricius) and B.
( = Gnathias sensu Mitchell 1962) is needed pensylvanicus (Degeer)] and Subterraneo-
to clarify separation of N. bella from N. bombus (B. borealis Kirby) that frequently
ovata, N. lepida, and other similar species, visit clovers (especially Trifolium) were not
We collected a single male Nomada austra- collected. Absence of B. fervidus is surpris-
es, which is one of the three species ing, but B. pensylvanicus has been scarce in
belonging to the erigeronis group ( = Cen- NY in recent years and is no longer, "An
trias) known from NY. These are aestival abundant southern species, common as
cleptoparasites of Agapostemon. far north as central NY..." (Leonard
Anthophora was represented by the 1928:1032). Bombus borealis has always been
wood-nesting species A. (Clisodon) termina- uncommon in New York State (Leonard
lis, which is widely distributed and nu- 1928), and is generally absent from de-
merous in northern and montane forests veloped areas (e.g., it is unknown from the
from Siberia to eastern Canada [Davydova city of Ithaca, NY, but occurs in nearby
and Pesenko 2002; these authors distin- countryside).
guished the Holarctic A. terminalis from the Absence of Bombus (Bombus) affiuis in our
Palearctic A. furcata (Panzer)]. sample of 1261+ bumble bee individuals is
troubling because this species is well
Bumble bees represented in historical collections from
Black Rock Forest is a favorable habitat the northeastern United States, and is
for bumble bees, and certain species were expected to be "...moderately abundant
found in large numbers, especially Bombus in the eastern to southern parts of the [New
(Pyrobombus) impatiens and its social para- York] State..." (Leonard 1928: 1031). How-
site B. (Psithyrus) citrinus (also known to ever, this species has recently disappeared
attack other Bombus species). Large num- from New York (e.g., from Ithaca and the
bers of B. impatiens in our late season NYC area, JSA, unpublished) and else-
samples reflect the unusually large colony where (L. Day, pers. comm.). The regional
218 Journal of Hymenoptera Research
disappearance of B. affinis is coincident NYC) habitats in the eastern United States
with an abrupt decline in B. (Bombus) to the point where it could be classified as
terricola Kirby at Ithaca NY (Ascher, un- invasive. We collected 66 specimens from
published), and elsewhere (L. Day, pers. on or around native vegetation and in
comm.), as well as the extirpation of the bowls, and one female emerged from a trap
closely related B. (Bombus) occidental is nest. Non-specificity to orchards should
Greene from the San Francisco Bay Area not be surprising as Osmia (Osmia) species
and elsewhere in western North America, such as O. cornifrons and the closely related
and the precipitous decline of the endan- native species O. lignaria are polylectic, not
gered B. (Bombus) franklini from its excep- specialists of fruit crops. In areas near
tionally restricted range in southern Ore- where O. cornifrons were deliberately re-
gon and northern California (Thorp 2005). leased (e.g., Patuxent National Wildlife
Populations of B. affinis, and of all North Refuge, see Cane 2003), a very similar
American species of subgenus Bombus, and Asian species, Osmia (Osmia) taurus Smith
their obligate social parasites [e.g., B. has been found to be established. This
(Psitin/rus) ashtoni; a queen of this species species has also been found in Huntingdon
was collected at BRF on June 13 1988, by J. County in south-central Pennsylvania (VG,
G. Rozen], should be carefully monitored, new data), but not yet in NY.
as parasitism by Nosema and other para- We collected 10 Anthidium oblongatum,
sites introduced and spread via the green- a species native to Europe and only re-
house trade in Bombus colonies poses cently detected in North America (Hoebeke
a potentially severe threat to their survival, and Wheeler 1999). This species is now
Introduced bee species. — Our samples in- abundant in the mid-Atlantic States, New
eluded numerous individuals of certain York, and southern New England, usually
exotic bee species that have become estab- in association with favored host plants such
lished and locally invasive in eastern North as Lotus corniculata, a weed generally
America beginning in the 1990's. distributed in waste places such as road-
Megacliile sculpturalis, a giant resin bee sides and abandoned lots, and Sedum.
native to northeastern Asia, was first The halictine species Lasioglossum (L.)
collected in New York State in 1997 leucozonium has long been present in North
(Ascher 2001) and is now widely distrib- America and has therefore been generally
uted and locally abundant in the Finger- regarded as native. However, its North
lakes Region, and in southern NY, in- American range is restricted to northeast-
eluding NYC. Outside of New York, M. ern USA and southeastern Canada and
sculpturalis is now quite widely distributed does not include northwestern Canada or
and has recently been found in additional Alaska (see maps in McGinley 1986). This
northeastern states such as Massachusetts distributional pattern, and association of
(Martha's Vinyard, P. Gambino pers. this species with introduced weeds such as
comm.), Vermont, and New Hampshire Chicorium (Asteraceae), suggests that this
(S. Droege, pers. comm.), as predicted by ground-nesting species is adventive from
Hinojosa-Diaz et al. (2005). Europe, not native as has been assumed.
The horn-faced mason bee Osmia comi- Molecular phylogenetic placement of L.
frons, native to eastern Asia including leucozonium and L. zonulum (Smith) within
Japan, was deliberately introduced by the otherwise exclusively Old World leuco-
USDA scientists as a managed pollinator zonium species group, and lack of signifi-
of apples. After wide distribution and cant genetic differences between Old and
release, this species has recently estab- New World samples (see, e.g., Danforth
lished large populations in natural and and Ji 2001), further support the idea that
urban (e.g., Manhattan and Brooklyn, the occurrence of these species in North
Volume 15, Number 2, 2006 219
America is adventive. It is possible that and 1,179 specimens (18.2%) belong to
these species were introduced in soil parasitic species (Fig. la). Of the 144 bee
carried in ships' ballast as has been species recorded in this study, 116 (80.5%)
hypothesized for another ground-nesting are pollen-collecting species and 28 (19.4%)
bee species native to Europe and found in are parasitic (Fig. lb). The abundance and
our study, Andrena wilkella. Extensive diversity of parasites reflects a rich fauna
sampling of variable molecular markers of vernal Nomada associated with Andrena
such as COI is needed to test hypotheses of hosts. The preponderance of females (4321
native vs. adventive origin for bee species vs. 1977 males vs. 245 of unrecorded sex)
with Holarctic distributions. Mcgachile cen- corresponds with the large number of
tuncularis (L.) may be another early in- workers of eusocial species, including the
traduction from Europe, as this species has two most numerous species at BRF. Of the
not been recorded in Alaska as would be 6,543 bee specimens collected, 1,222
expected for a species with a naturally (18.7%) were Augochlorella aurata and 845
Holarctic range. (12.9%) were Bombus impatiens. The sample
Workers of Apis mellifera (L.) were of 1,113 bumble bees collected was domi-
abundant from mid-June and into October nated by B. impatiens (845, 75.9%) and its
but were generally not collected. social parasite B. (Psithyrus) citrinus (154,
Of the 144 bee species recorded in this 14.1%).
study, six (4.2%) are exotic and 138 (95.8%) Seasonal patterns of occurrence can be
are native. Of the 6,543 specimens collect- obtained from Appendix 1, which gives
ed, 115 (1.7%) belong to exotic species, and extreme dates for BRF (by calendar date),
6,428 (98.2%) belong to native species. NY as a whole (by month), and the entire
Wasps and other non-bees. — Our apoid North American range (by month) for each
wasp samples include 23 crabronid species species. General patterns include an abun-
(12 genera; Appendix 2). Some of these are dance and diversity of Andrena and their
generally numerous in forest edge habitats Nomada parasites at forest floor sites prior
in New York such as Ectemnius continuus, to leaf-out. At more open sites, seasonal
which nests in holes in wood. Other turnover of the bee fauna was apparent,
species collected such as Astata leitthostromi with notable peaks of abundance and
and Bia/rtes quadrifasciata are ground-ne- species diversity corresponding with the
sters that favor more open, often sandy bloom of favored plants such as Vacciniinn
habitats. Our vespid wasp sample includes in late spring (visited by, e.g., Andrena and
long series of the native paper wasp Polistes Osmia spp.) and Solidago in late summer
fuscatus, both sexes of Vespula consobrina, (visited by, e.g., Colletes spp., Andrena
a yellowjacket of northern (Canadian and simplex Pseiidopamirgus andrenoides, and
Transition Zones) affinities, one individual the workers and males of the dominant
of the rather scarce Zethus spinipes, and eusocial species Augochlorella aurata and
a variety of eumenines including cavity- Bombus impatiens). Rather few oligolectic
nesting species found in our trap nests. bee individuals were captured (292, 4.5%
Ecological and behavioral patterns. — Eco- of the total) (Fig. 2a), but these represented
logical information (summarized in Ap- a significant number of species in our
pendix 1) was compiled for each of the 144 sample (19, 13.2%) (Fig. 2b).
bee species from information found in Although soil nesting individuals and
catalogs and revisions, primary literature, species predominated in our samples, hive
and field observations, including those nesters, wood burrowers, and cavity-
made during the BRF survey. nesters were also well represented (Fig. 3a,
Of the 6,543 specimens collected, 5,364 b). Cavity-nesting species were numerous
(82.0%) belong to pollen collecting species, relative to the number of individuals, as
220
Journal of Hymenoptera Research
1A Sociality of Individuals
1 B Sociality of Species
Parasitic
18%
Subsocial
1%
Solitary
27%
Parasitic
21%
Subsocial
Rusocial
21%
Eusocial
54%
Solitary
57%
2A Oligolectic vs. Polyectic Individuals
2B Oligolcctic vs. Polylectic Species
Oligolcctic
5%
Polylectic
Solitary
^ 23%
Other^^
„..--
72%
Oligolectic
13%
| Polylectic
Solitary
44%
3A Nest Substrate by Individuals
3B Nest Substrate by Species
Other
18%
Other
21%
Wood
Burrowers
7%
Cavity
7%
Wood
Burrowers
3%
Cavity
17%
4A Net and Bowl Catch by Individuals
4B Species Catch by Method
Unknown
0%
White Bowl
24%
Net Only
21%
*
Blue Bowl
9%
Yellow Bowl
33%
Both
61%
Bowl Only
18%
Figs 1-4. Summary of ecological properties of Black Rock Forest (BRF) bees. 1, Sociality. The category
"solitary" includes communal species. Some individual nests or local populations of species categorized as
"eusocial" may be solitary: 1 A, percentage of individual bees belonging to each of the four recognized categories
of sociality; IB, percentage of bee species belonging to each of the categories. 2, Pollen specialization; those
classified as oliglolectic are specialists that usually collect pollen from only a single family of plants; the
polylectic category includes polylectic solitary bees only; those in the "other" category include parasites and
social bees, which are necessarily generalists (i.e. polylectic): 2A, percentage of bee individuals that are
Volume 15, Number 2, 2006
221
several cavity-nesters were represented by
singletons or doubletons. By contrast,
wood burrowing species were relatively
few (Fig. 3b), although some of these
species were captured in large numbers
(e.g., Augochlora pura). The large number
of hive-nesting individuals relative to
species likely reflects their eusociality
(see above).
A few species typical of more open and
sandy areas were found at BRF (e.g.,
Lasioglossum heterognathum, Bicyrtes, As-
tata), but sand specialists such as L. vierecki
were not found.
Efficacy and Biases of trapping methods. —
The year 2003 was characterized by long
periods of cold and cloudy weather and
pans may have been particularly useful
under these conditions as these allow catch
during brief windows of sun on days when
net-collecting would be unrewarding.
Nearly twice as many individuals were
bowl trapped than netted (4,322 vs. 2,221)
(Fig. 4a), but the net sample was biased
against certain of the most common and
readily identified taxa (see above). Bowls
were found to be particularly useful in
forest and at the forest edge where flowers
are few, dispersed, or in the case of trees
and shrubs, difficult to reach. Where
flowers are scarce, bowls may be particu-
larly effective due to lack of competition
from real flowers. Using bowls, we found
certain inconspicuous forest-associated spe-
cies rarely taken in nets such as Stelis nitida.
Well known biases of bowl traps reinforced
by our study include low catch rates for
certain groups, especially fast and high-
flying species of, e.g., Colletes, Megachile, and
perhaps Melissodes, and high catch rates for
slow and low-flying species of, e.g., small
Lasioglossum, Andrena, Osmia, and Nomada.
Our results generally support the currently
accepted view that a combination of bowl
trapping using multiple colors and netting
is the best way to efficiently collect a plural-
ity of species (S. Droege et al. protocol).
Only 89 of the 144 bee species collected
(61.8%) were collected by both nets and
bowls, with 30 species (20.8%) unique to
nets and 26 (17.6%) unique to bowls
(Fig. 4b). The net collected sample was
richer than the bowl trapped sample in
total bee species (117 vs. 113) and in number
of unique species (29, 20.4% vs. 25, 17.6%).
Although wood and cavity-nesting bees
were numerous in this survey, only one
individual bee (the introduced Osmia cor-
nifrons) used our trap-nests. The poor
performance of trap-nests might possibly
be explained by an abundance of natural
nesting substrates (standing dead wood) at
BRF. Alternatively, bees may have been
out-competed for the trap-nests by eume-
nines and Trypoxylon, or else the nests may
have been placed in sites that ultimately
proved to be too shady.
Comparison to other bee faunas. — In com-
parison to the bee fauna of NYS as a whole
(423 species) and to the fauna of some well-
sampled localities within the state such as
Ithaca (274 species), the 144 species iden-
tified in our BRF sample is relatively few
(Table 1). However, several of these re-
cords are of considerable biogeographic or
ecological interest (see above). The NY bee
fauna includes many species which are
regionally rare and/or have highly special-
ized ecological requirements, and are
therefore unlikely to be found at BRF.
oligolectic, polylectic and solitary, or other; 2B, percentage of bee species that are oligolectic, polylectic and
solitary, or other. 3, Nest substrates: 3A, percentage of individuals belonging to each nesting category: soil,
cavity, wood burrowers, hive, or other (primarily cleptoparasites that live in the nests of their hosts); 3B,
percentage of bee species known or inferred to use the nest substrate indicated. 4, Collecting method: net vs.
white bowl, vs. blue bowl, vs. yellow bowl: 4A, percentage of bee individuals caught by each method; 4B,
percentage of bee species caught by net only vs. bowl only vs. both net and bowl.
222
Journal of Hymenoptera Research
Nonetheless, it seems highly probable that
at least 250 bee species could be present
at BRF based on totals of 274 species
recorded from Ithaca, Tompkins County,
NY (Ascher, unpublished), in a colder
climate than BRF, and ca. 300 species
recorded from the vicinity of Carlinville
in southern Illinois (Robertson 1929, Marlin
and LaBerge 2001), in seemingly unre-
markable farm country.
The high number and proportion of
singletons (28 spp., 19.4%), of doubletons
(12 spp., 8.3%), of species known from
a single sex (ca. 31 spp., ca. 21.0%)
excluding Lasioglossum, and of rarely col-
lected species (i.e., 3-10 individuals col-
lected: 36 species, 25.3%), indicate that
more prolonged and intensive surveying
using the same methods would reveal
many additional species, likely resulting
in taxonomically and biogeographically
significant specimens.
Another indication of the incomplete-
ness of sampling of the total BRF fauna is
that only 57.8% of the 249 bee species
known from southern New York excluding
NYC and Long Island (i.e., Sullivan, Ulster,
and Dutchess, Orange, Putnam, Rockland,
and Westchester, Counties) were found.
These might be considered to represent
a regional pool of species from an area
relevant to BRF. The total of 249 species
known from an area relevant to BRF is only
59% of the species total for New York State
as a whole (423), which in turn is only 57%
of the 743 bee species known from the
eastern USA. Twenty additional species
recorded in NY only from coastal NYC and
Long Island (e.g., the coastal dune special-
ist Lasioglossum marinum) are less likely to
occur at BRF.
Most bee species at BRF are widely
distributed in NY and have been recorded
from other well-collected sites such as
Ithaca (123 species shared with BRF,
85.4% of the BRF total) and NYC (103
species shared with BRF, 71.5% of the BRF
total). Northern elements of the fauna at
BRF can be defined as those species known
from the northern and montane portions of
NY (e.g., the Adirondack Mountains and in
most cases Ithaca), but absent from NYC,
Long Island, and other warmer and coastal
areas. Examples of northern species occur-
ring at or near their southern limits at BRF
and unknown from NYC include Andrena
rufosignata, A. algida, Stelis nitida, Osmia
felti, O. inermis, Bombus ternarius, and
possibly B. sandersoni (southern distribu-
tional limits of this species remain un-
certain due to identification difficulties
versus B. vagans). Although these northern
species are likely genuinely absent from
NYC, many of the 42 species known from
BRF, but not NYC may be found in the
latter area when more thorough samples
have been made of semi-natural habitats
such as Pelham Bay Park. Southern ele-
ments in the BRF fauna include the
following species that are unknown from
the very well collected Fingerlakes Region
(which includes Ithaca): Lasioglossum bni-
neri, Andrena nuda, A. eonfederata, A. hilaris,
and Melissodes subillata. The apparent ab-
sence of these species from Ithaca and
elsewhere in central and northern New
York is probably genuine and likely reflects
a real faunal difference from BRF. Andrena
violae is another species of southern affin-
ities that is very rare in Ithaca (see above).
The Sorensen index [Cs = 2a /(2a + b + c)
where a is the number of species shared
between two sites, b is the number of
species found at only one site, and c is the
number of species found only at the other
site] was used to quantify similarity be-
tween various sites. The total for BRF vs.
Ithaca is 58.9% whereas the total for BRF
vs. NYC is 58.2%. The similar Sorensen
values for comparisons involving these
two areas (despite BRF's much greater
geographical proximity to NYC) reflect
many shared widespread and northern
species with Ithaca, and significant differ-
ences between BRF and NYC due to the
presence of northern forest elements (e.g.,
blueberry associates) only at BRF and of
coastal /sand associates only in NYC.
Volume 15, Number 2, 2006
223
ACKNOWLEDGMENTS
We are grateful to the staff of Black Rock Forest for
their invaluable advice and assistance with practical
matters, and to the Steifel Foundation for an award
from their Black Rock Forest Consortium Small Grants
for Scientific Research and Education program
awarded to J.G. Rozen and V. Giles. We thank: S.
Droege, L. Day, and T. Griswold for identifying bees,
and M. Arduser for sharing unpublished keys and
other taxonomic information and commenting on the
manuscript. P. Gambino provided unpublished re-
cords of NY bees and identified many of our
crabronid wasps. K. Magnacca provided bee records
and other assistance. The following people kindly
provided access to and information about historical
collections of New York bees: B. N. Danforth, E. R.
Hoebeke, J. K. Liebherr, T. McCabe, T. Schultz, D.
Yanega. S. Kornbluth generously provided assistance
with specimen labeling and data entry and manage-
ment. We thank K. Russell for her helpful discussons
of methods and data base management. C. Encarna-
cion, and E. Burbano prepared our specimens. T.
Nguyen lent his assistance with technical matters. J. G.
Rozen, J. M. Carpenter, A. C. Vallely, and C. Dong
made valuable comments on this manuscript. J. M.
Carpenter identified our vespid wasps. We thank R.
G. Goelet for his support of bee research at the
AMNH. We are especially grateful to J.G. Rozen for
his guidance and support. In addition to his help in
this project, we both owe our interest in bees to his
generosity and encouragement.
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Volume 15, Number 2, 2006
225
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