Journal of Hymenoptera research

.SOCIETY

Journal of
Hymeno
Research

Volume 7, Number 1 April 1998

ISSN #1070-9428

CONTENTS

ABD-RABOU, S., and M. M. ABOU-SETTA. Parasitism of Siphoninns pbillyreae (Homoptera:

Aleyrodidae) by aphelinid parasitoids at different locations in Egypt 57

ENGEL, M. S. A new species of the Baltic amber bee genus Electrapis (Hymenoptera: Api-

dae) 94

ENGEL, M. S., and R. W. BROOKS. The nocturnal bee genus Megaloptidia (Hymenoptera:

Halictidae) 1

van der ENT, L.-J., and S. R. SHAW. Species richness of Costa Rican Cenocoeliini (Hy-
menoptera: Bracondiae): a latidudinal and altitudinal search for anomalous di-
versity 15

JOHNSON, N. F., and L. MUSETTI. Geographic variation of sex ratio in Pelecinus polyturator

(Drury) (Hymenoptera: Pelecinidae) 48

KIMSEY, L. S., and M. S. WASBAUER. Revision of the American Hphiid genus Quemaya

Pate (Hymenoptera: Tiphiidae: Brachycistidinae) 38

KURCZEWSKI, F. E. Territoriality and mating behavior of Sphex pensylvanicus L. (Hyme-
noptera: Sphecidae) 74

NEWMAN, T. M., and D. L. J. QUICKE. Sperm development in the imaginal testes of

Aleiodes coxalis (Hymenoptera: Braconidae: Rogadinae) 25

RUST, R. W. The effects of cavity diameter and length on the nesting biology of Osmia

lignaria propinqua Cresson (Hymenoptera: Megachilidae) 84

SHAW, S. R., P. M. MARSH, and J. C. FORTIER. Revision of North American Aleiodes
Wesmael (Part 2): the apicalis (Brulle) species-group in the New World (Hymenop-
tera: Braconidae, Rogadinae) 62

(Continued on back cover)

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This issue was mailed 14 May 1998

J. HYM. RES.
Vol. 7(1), 1998, pp. 1-14

The Nocturnal Bee Genus Megaloptidia (Hymenoptera: Halictidae)

Michael S. Engel and Robert W. Brooks

(MSE) Department of Entomology, Comstock Hall, Cornell University, Ithaca, NY 14853 USA;
(RWB) Division of Entomology, Natural History Museum, Uruversity of Kansas,

Lawrence, KS 66045, USA

Abstract. — The nocturnal augochlorine bee genus Megaloptidia is revised and three species rec-
ognized; Mt'galopttidia contradicta (Cockerell), M. nocturna (Friese), and M. saulensis Engel and
Brooks new species. The genus is newly diagnosed, distributional data given, and the male com-
pletely described for the first time. The first flower record for a species of this genus is given as
Dichorisandra iilei (Commelinaceae). A lectotype and two paralectotypes are designated for M.
contradicta. Megalopta angiisticeps Friese is considered a junior synonym of Megaloptidia nocturna
(new synonymy). Megalopta pmllitarsus Friese, previously considered to be a species of Megaloptidia,
is in fact a junior synonym of Megommalion insigne (Smith) (new synonymy).

The neotropical halictine genus Megal-
optidia Cockerell consists of three, rarely
collected, nocturnal bees of the tribe Au-
gochlorini. Individuals of Megaloptidia can
be captured at lights during the night,
however, aside from their nocturnal habit,
the biology of Megaloptidia species remains
unknown. One female of Megaloptidia iwc-
tunia (Friese 1926), however, has been col-
lected at flowers of the monocotyledon Di-
chorisandra iilei (Commelinaceae). Mem-
bers of Dichorisandra are nectarless and it
is therefore believed that pollen is the only
reward for floral visitors. Dichorisandra
species possess poricidal anthers, suggest-
ing "buzzing" as the means of freeing the
pollen (i.e., the bee grasps the anther and
buzzes its wings, thus translating the vi-
bration to the anther and releasing the
pollen through the apical pore). Most au-
gochlorine species which have been stud-
ied are known to be buzz pollinators (e.g.,
Michener 1962, Rego and Albuquerque
1989, Renner 1989) and, in fact, the related
D. hexajidra is recorded to be buzz polli-
nated by the diurnal augochlorines Aitgo-
chloropsis (Paraugochloropsis) cupreola
(Cockerell 1900) and Pseudaugochlora gra-

minea (Fabricius 1804) (Sigrist and Sazima
1991). Within the Commelinaceae, Dichor-
isandra species are among the only ones to
be open during the night (e.g., D. hexandra
opens before dawn in southern Brazil)
(Sigrist and Sazima 1991), making them
suitable targets for nocturnal or crepus-
cular insects. Among related bee genera,
floral associations are only known for Me-
gommation (s. str.) (Moure 1943) which has
been recorded at species of Bauhinia (Fa-
baceae) (Friese 1926) and Ipomoeae (Con-
volvulaceae) (Jorgensen 1912). There is
nothing about the floral morphology of
any of these plants which seems to readily
explain the peculiarly modified mouth-
parts of these genera, which are extremely
narrowed with a distinctly pointed galeal
apex.

Megaloptidia was originally described by
Cockerell (1900) as a subgenus of the com-
mon nocturnal augochlorine genus Megal-
opta Smith (1853) owing to its superficial
similarity to this group. Cockerell distin-
guished his subgenus on the basis of wing
venation. Moure (1958) redescribed the
group, according it generic status for the
first time, and recognized its affinity with

Journal of Hymenoftera Research

his genus Megommation. Eickwort (1969),
in a revision of the genera and subgenera
of Augochlorini, provided a more thor-
ough description of the genus; however,
he did not have the opportunity to ex-
amine the male terminalia thus the distal
male sterna and genitalia remained un-
described. The genus is closely allied to
the genera Ariphaiiarthra (Moure 1951),
Megonmiation (s. lato), and Micromniation
(Moure 1969). All of these genera share
the distinctly narrowed labiomaxillary
complex, pointed galeal apex, orthogonal
epistomal sulcus, absence of the male la-
bral distal process, narrow spiculum on
the male eighth sternum, and the presence
of a parapenial lobe in the male genitalia.
The phylogenetic position of Megaloptidia
will be further elaborated on in a forth-
coming paper concerning the entire tribe
(Engel in prep).

Herein we provide new descriptions for
the genus and all included species. The
male is thoroughly described for the first
time and a key is presented for the iden-
tification of all three species.

MATERIAL AND METHODS

The following abbreviations are used
for institutions where material used in this
study is deposited: American Museum of
Natural History, New York, New York,
J.G. Rozen, Jr. (AMNH); Carnegie Muse-
um of Natural History, Pittsburgh, Penn-
sylvania, R.L. Davidson (CMNH); Field
Museum of Natural History, Chicago, Il-
linois, P.P. Parrillo (FMNH); Cornell Uni-
versity Insect Collection, Ithaca, New
York, J.K. Liebherr and E.R. Hoebeke
(CUIC); Natural History Museum of Los
Angeles County, Los Angeles, California,
R.R. Snelling (LACM); Instituto de Inves-
tigacion de Recursos Biologicos, Alexan-
der von Humboldt, Santa Fe de Bogota,
Colombia, F. Fernandez (UNCB); Instituto
Nacional de Pesquisas da Amazonia, Ma-
naus, Amazonas, Brazil (INPA); Museo de
Invertebrados 'G.B. Fairchild', Universi-
dad de Panama, Estafeta Universitaria,

Panama City, Panama', D. Quintero
(MIUP); Museu Entomologico de Univer-
sidade Federal de Vi^osa, Vigosa, Minas
Gerais, Brazil (UFVB); Museum fiir Natur-
kunde, Humboldt-Universitat, Berlin, Ger-
many, F. Koch, A. Kleine-MoUhoff
(ZMHB); United States National Museum,
Smithsonian Institution, Department of
Entomology, Washington, D.C., R.J.
McGinley (USNM); Division of Entomol-
ogy, Natural History Museum, University
of Kansas, Lawrence, Kansas, R.W. Brooks
(SEMC); G.A.R. Melo collection (GARM);
Philadelphia Academy of Natural Sci-
ences, Philadelphia, Pennsylvania, D.
Azuma (ANSP).

A total of 67 specimens of Megaloptidia
were located and examined in the course
of this study. All measurements were
made using an ocular micrometer on a
WILD-M5a microscope. The abbreviation
"o.d." in the descriptions refers to "ocellar
diameter" and relates the approximate
length of setae to the diameter of the me-
dian ocellus. SI, Tl, and Fl are given as
abbreviations for first metasomal sternum,
first metasomal tergum, and first flagello-
mere respectively.

Genus Megaloptidia Cockerell

Megalopta (Megaloptidia) Cockerell 1900: 373.
Type species: Megalopta {Megaloptidia) coii-
tradicta Cockerell 1900, by monotypy and
original designation. Moure 1958: 180. Eick-
wort 1969: 442.

Diagnosis. — Individuals of Megaloptidia
are robust bees which superficially resem-
ble in general habitus species of the larger
nocturnal genus Megalopta and the sole
species of Megommation (s. str.). Megalop-
tidia differs from the former genus in the
narrowed labiomaxillary complex, the ser-
rate inner hind tibial spur, the irregular
spacing of the distal hamuli, the narrow
gena, the unmodified labral distal keel, the
orthogonal epistomal sulcus, and the ab-
sence of lateral notches on the male S4.
Megommation differs from Megaloptidia in
the acute marginal cell apex, the deeply

Volume 7, Number 1, 1998

concave clypeal apex, the extremely short
and weakly border basitibial plate, and
the medioapical processes on the male
S3-4.

Description. — The following description
is based on all three included species. FE-
MALE: Epistomal sulcus orthogonal.
Clypeus and supraclypeal area strongly
bowed, as in Megommation (Fig. 2). Malar
space short. Mid-region of face gently
sunken around antermal socket. Preoccip-
ital ridge rounded. Inner orbit of com-
pound eye strongly emarginate; eyes
greatly enlarged, much broader than gena
in profile (Figs. 2, 8, 14, 19); eye hairs
short. Ocelli greatly enlarged (Figs. 1, 7,
13, 18), without impressed line running
between lateral ocelli. Vertex extremely
short, barely an ocellar diameter in length,
usually less. Labral basal elevation orbic-
ular, protuberant in profile, distal process
narrowly triangular, labral teeth absent
(Fig. 1). Mandible broad, subapical tooth
well defined (Fig. 1). Hypostomal ridge
carinate, not projecting beyond posterior
margin of head, anterior angle rounded.
Mouthparts greatly narrowed, approxi-
mately 10 times longer than wide (Fig. 4).
Galeal base reaching to base of stipes,
apex pointed, inner strip with setae; galeal
comb absent; maxillary palp not greatly

lengthened. V-shaped brace of salivary
plate absent; combined length of labial
palp segments 2 and 3 shorter than 1.
Pronotal lateral angle obtuse, not pro-
duced; lateral and dorsal ridges rounded.
Mesoscutum broadly rounded anteriorly;
mesoscutal lip low and rounded. Tegula
rounded. Propodeal dorsal ridge rounded;
lateral ridge carinate, ridges slightly di-
vergent; propodeum slightly narrowed
posteriorly; pit of posterior face narrow.
Wing hairs long; apex of marginal cell
truncate (Fig. 3). Anterior basitarsal brush
present. Inner hind tibial spur serrate, ser-
rations sharp (Fig. 5). Basitibial plate nar-
rowly rounded, all borders well defined.
Division of T5 narrow. MALE: As for the
female with the following modifications.
Antenna of moderate length, reaching to
posterior border of mesoscutum; scape
long; F2 shorter than Fl; sensory plate ar-
eas present. Labral basal elevation absent;
distal process absent. Gradulus of T7 ab-
sent. S4 with depression along posterior
margin. S8 with posterior margin pro-
duced into a short, blunt median projec-
tion; spiculum narrow. Gonobasal bridge
narrow; dorsal lobes strong. Dorsal gon-
ostylus a long thin process. Basal process
of gonostylus absent. Parapenial lobe
present. Ventral prong of penis valve pres-
ent.

KEY TO SPECIES OF MEGALOPTIDIA

1. Propodeal lateral surface strongly punctured, punctures almost contiguous; length of pro-
podeal triangle approximately equal to that of metanotum; mesepisternum strongly and
closely punctured, punctures separated by less than a puncture width, integument between
smooth (excluding hypoepimeron); mesoscutum punctured contiguously, integument be-
tween imbricate M. contradicta (Cockerel!)

- Propodeal lateral surface imbricate with punctures separated by 2-3 puncture widths;
length of propodeal triangle little to much greater than that of metanotum; sculpturing of
mesepisternum and mesoscutum variable, but mesoscutum never strongly punctured .... 2

2. Labrum and clypeus amber; mesepisternum minutely punctured, punctures separated by
3^ times puncture width, at least on posterior half, integument between smooth or im-
bricate (excluding hypoepimeron); mesoscutum minutely punctured, punctures separated
by 2-3 times puncture width; lateral ocellus almost touching compound eye, ocellocular
distance 0.25 o.d. in female (Fig. 1), 0.33 o.d. in male (Fig. 7); frontal line ver>' weakly
produced in profile in female (Fig. 2); median ocellus wider than interantennal distance in
female (Fig. 1); scopal hairs of hind femur and tibia amber M. noctuma (Friese)

Journal of Hymenoptera Research

Figs. 1-8. Me^nlopthiin nocturna (Friese), 1-6 female, 7-8 male. 1, 2, head, front and side views respectively.
3, fore wing. 4, mouthparts excluding labium, ventral view, mp = maxillary palps, c = cardo, m = maxilla,
hf = hypostomal fossa. 5, inner hind tibial spur. 6, hind leg and metasoma, side view, circle enlargement
shows scopal hairs on S2 and S3. 7, 8, head, front and side views, respectively.

Volume 7, Number 1, 1998 5

- Labrum and clypeus dark brown; mesepisternum closely and weakly punctured, punctures
separated by less than a puncture width (excluding hypoepimeron); mesoscutum punc-
tured, punctures weak and separated by 2-3 puncture widths centrally, laterally becoming
separated by less than a puncture width; lateral ocellus removed from compound eye 0.5
o.d. (Fig. 18); frontal line protuberant in profile (Fig. 19); median ocellus narrower than
interantennal distance (Fig. 18); scopal hairs of hind tibia black (those of femur amber) . .
M. saulensis n. sp.

Megaloptidia contradicta (Cockerell)

(Figs. 13-17, 20)

Megalopta {Megaloptidia) contradicta Cockerell
1900: 373. Schrottky 1902: 407. Friese 1926:
124.

Megaloptidia contradicta (Cockerell); Moure 1958:
180.

Diagnosis. — Labrum and apical % of
clypeus amber. Mesoscutum closely and
coarsely punctured, punctures separated
by less than a puncture width, except over
central disc punctures becoming v^^eaker
and separated by 1-3 times a puncture
width. Mesepisternum closely punctured,
punctures separated by less than a punc-
ture width, integument between punc-
tures smooth; hypoepimeron smooth with
a few widely spaced minute punctures.
Scopal hairs of hind leg fuscous. Propo-
deal lateral surface strongly punctured,
punctures nearly contiguous; length of
propodeal triangle equal to that of meta-
notum. Only three specimens of this spe-
cies were located, all from Cockerell' s type
series.

Description. — The following description
is based on the lectotype. MALE: Total
body length 11.3 mm; forewing length 8.6
mm. Head wider than long (Fig. 13). Dis-
tal margin of clypeus barely projecting be-
low lower tangent of compound eye (Fig.
14); supraclypeal area wider than long,
0.59 times length of clypeus. Frontal line
carinate from just below antennal sockets
to just above sockets, ending at this point.
Measurements of head of lectotype in mm:
width 3.0; length (to apex of clypeus) 2.5;
clypeal length 0.8; lower interorbital dis-
tance 0.9; upper interorbital distance 1.2;

interantennal distance 0.4; antennocellar
distance (to median ocellus) 0.6; between
lateral ocelli 0.4; median ocellus to lateral
ocellus 0.1; lateral ocellus to compound
eye 0.16; prementum length 1.9, width
0.18. Scape reaching past lateral ocellus,
length 1.1; pedicel slightly longer than
wide, length 0.2, width 0.18; Fl longer
than wide and longer than F2, length 0.26,
width 0.2; F2 as long as wide, length-
width 0.22; F3-9 each longer than wide,
individual lengths 0.24, widths 0.22; FIO
longer than wide and longer than imme-
diately preceding flagellomeres, length
0.26, width 0.22; Fll longer than wide and
longer than all previous flagellomeres,
length 0.42, width 0.22. Median line
strongly impressed; parapsidal lines
weak. Intertegular distance 2.2. Propodeal
triangle about as long as metanotum,
much shorter than scutellum. Basal vein
distad cu-a by 2.5 times vein width; Ir-m
basad Im-cu by vein width; 2r-m distad
2m-cu by 4.5 times vein width, 2r-m
weakly curved. First submarginal cell lon-
ger than second and third combined; sec-
ond narrowed anteriorly; anterior border
of third along Rs almost 2 times as long
as anterior border of second, posterior
border of third 2 times longer than ante-
rior border; marginal cell length 1.2, width
0.3. Distal hamuli arranged 3-1-1-3. S5 api-
cally emarginate and bent ventrally (Fig.
15). S6 apically emarginate and mediolon-
gitudinally concave (Fig. 15). Male termin-
alia as in figures 16 and 17.

Clypeus and supraclypeal area finely
imbricate, with a few weak punctures sep-
arated by 1-3 puncture widths on lateral
borders. Face minutely and closely punc-

Journal of Hymenoptera Research

Figs. 9-17. Megaloptidia noctiirnn (Friese), 9-12 male; Megaloplidia coiitrndicla (Cockerell), 13-17 male. 9, me-
tasoma, apical half, oblique view, numbers refer to sterna. 10, S7 and S8, dorsal view is right half and ventral
view is left. 11, genital capsule, dorsal view is right half and ventral view is left, dl = dorsal gonostylar
process, pi = parapenial lobe, vp = ventral prong of penis valve. 12, S6, ventral view. 13, 14, head, front and
side views, respectively. 16, genital capsule, dorsal view is right half and ventral view is left. 17, S7 and S8,
ventral view is right half and dorsal view is left.

Volume 7, Number 1, 1998

tured, integument between imbricate. Ver-
tex smooth and impunctate. Gena and
postgena impunctate and finely imbricate.
Pronotum finely imbricate. Mesoscutum
closely and coarsely punctured, punctures
separated by a puncture width or less, in-
tegument between punctures imbricate,
medially punctures becoming weaker and
more widely spaced, separated by 1-3
punctured widths. Scutellum with minute
punctures separated by 1-2 puncture
widths, integument between smooth.
Metanotum sparsely and weakly nodu-
late, integument otherwise smooth. Pre-
episternum and mesepisternum closely
punctured, punctures separated by a
puncture width or less, integument be-
tween smooth. Hypoepimeron smooth,
with a few minute punctures separated by
4-5 puncture widths. Metepisternum
smooth. Propodeal triangle imbricate; lat-
eral surface closely punctured, punctures
separated by less than a puncture width;
posterior surface smooth. Terga and ster-
na imbricate.

Head dark brown with a few weak
blue-green highlights; labrum and apical
three-quarters of clypeus amber; mandible
dark brown; antennae brown. Mesosoma
and metasoma dark brown; tegula amber
and translucent along outer margins; legs
dark brown, except amber on protarsus,
apical four mesotarsomeres, and apical 2
metatarsomeres.

Pubescence generally pale or fuscous,
fuscous hairs mostly on mesosoma and
apical terga of metasoma. Scattered simple
hairs (1 o.d.) on face, with suberect
branched hairs (0.5 o.d.) along inner mar-
gins of compound eyes. Similar suberect
and simple hairs on gena. Postgena with
a few simple hairs (1.5-2 o.d.) on lateral
and posterior borders. Hairs of face, gena,
and postgena pale, those of vertex becom-
ing fuscous. Pronotum with scattered sim-
ple hairs (1 o.d.) and laterally with ex-
tremely short, appressed branched hairs
not obscuring the surface. Mesoscutum
with scattered simple hairs (1 o.d.) and

shorter (less than 0.5 o.d.) branched hairs
on lateral and posterior borders. Scutel-
lum and metanotum with sparse simple
hairs (1.5 o.d.). Pleura with scattered sim-
ple pale hairs (1-1.5 o.d.) and shorter (0.5
o.d. or less) suberect branched hairs. Hairs
of forelegs pale or golden, outer margins
with simple hairs (1-1.5 o.d.); mid legs
with pale hairs except fuscous on basitar-
sus and outer margin of tibia, hairs sim-
ple, small tuft of closely packed hairs (1
o.d.) on inner basal margin of femur; hair
of hind legs mostly fuscous, inner surface
of basitarsus with stiff black hairs (1-1.5
o.d.). Terga with scattered simple hair (1-
1.5 o.d.), becoming longer on lateral mar-
gins and more distal terga. Sterna with
sparse simple hair (1-2.5 o.d.); hairs of S4
longest and concentrated on borders of
apical depression and medially towards
basal border; S5 with pair of apical sub-
median hair tufts (Fig. 15); hairs of 56
most numerous, more dense laterally (Fig.
15).

FEMALE: Unknown.

Type material.— BRAZIL: Para: Lecto-
type #345, male, Benevides [1°22'S,
48°15'W], July (CMNH). Paralectotype,
male, Santarem (CMNH). Paralectotype,
male, Santarem, but lacking all metasomal
segments except Tl and SI and both an-
tennae leaving only the scape, pedicel, Fl
and F2 (ANSP).

Remarks. — In 1957 Padre Jesus S. Moure
examined the material listed above and
placed lectotype labels on the specimens,
however, he failed to publish lectotype
designations for this species. We now des-
ignate lectotype and paralectotypes for
Megaloptidia contradicta using the same
specimens. The lectotype specimen now
bears a new label reading "LECTOTYPE:
Megalopta (Megaloptidia) contradicta Cock-
erel!, 1900; desig. M. Engel & R. Brooks",
and the paralectotypes now have similar
labels.

In Eickwort's (1969) study of the Au-
gochlorini he referred to a male of M. con-
tradicta (the paralectotype from ANSP des-

Journal of Hymenoptera Research

ignated above) along with a series of fe-
males he considered as possibly belonging
to M. contradicta. This series of females
from the AMNH are, in actuality, all of M.
nocturna.

Megaloptidia nocturna (Friese)

(Figs. 1-12, 20)

Megalopta nocturnn Friese 1926: 127.

Megalopta angiisticeps Friese 1926: 127. New

synonymy.
Megaloptidia iwctuma (Friese); Moure and Hurd

1987: 242.
Megaloptidia angiisticep's (Friese); Moure and

Hurd 1987: 241.

Diagnosis. — Labrum and clypeus amber.
Mesoscutum minutely punctured, punc-
tures separated by 2-3 times a puncture
width. Mesepisternum minutely punc-
tured, otherwise integument smooth or
imbricate; hypoepimeron as on mesepi-
sternum although punctures more widely
spaced. Scopal hairs of hind femur and
tibia amber. Propodeal lateral surface im-
bricate with punctures separated by 2-3
times a puncture width; length of propo-
deal triangle little to much greater than
that of metanotum. This is the most com-
mon of the three species with 62 speci-
mens examined during the course of this
study.

Description. — The following description
is based on Friese's holotypes. MALE: To-
tal body length 13.3 mm; forewing length
8.3 mm. Head wider than long (Fig. 7).
Distal margin of clypeus barely projecting
below lower tangent of compound eye
(Fig. 8); supraclypeal area wider than
long, 0.68 times length of clypeus. Frontal
line carinate from just below antennal
sockets to just above sockets, ending with
acute projection (Fig. 8). Measurements of
head of holotype in mm: width 2.8; length
(to apex of clypeus) 2.5; clypeal length 0.8;
lower interorbital distance 0.8; upper in-
terorbital distance 1.1; interantennal dis-
tance 0.3; antennocellar distance (to me-
dian ocellus) 0.5; between lateral ocelli 0.3;
median ocellus to lateral ocellus 0.04;

lateral ocellus to compound eye 0.08; pre-
mentum length 2.1, width 0.2. Scape
reaching past lateral ocellus, length 1.0;
pedicel slightly longer than wide, length
0.2, width 0.18; Fl longer than wide and
longer than F2, length 0.24, width 0.18
(Fig. 7); F2 as long as wide, length-width
0.2; F3 and F4 each longer than wide, in-
dividual lengths 0.22, widths 0.2; F5 lon-
ger than wide, length 0.24, width 0.2; F6
longer than wide, length 0.26, width 0.2;
F7 and F8 longer than wide, individual
lengths 0.28, widths 0.2; F9 and FIO longer
than wide, individual lengths 0.3, widths
0.2; Fll longer than wide and longer than
preceding flagellomeres, length 0.44,
width 0.2. Median line strongly im-
pressed; parapsidal lines weak. Intertegu-
lar distance 1.8. Propodeal triangle longer
than metanotum, slightly shorter than scu-
tellum. Basal vein distad cu-a by vein
width; Ir-m distad Im-cu by 5 times vein
width, intersecting second submarginal
cell about half way through cell; 2r-m dis-
tad 2m-cu by 2 times vein width, 2r-m
weakly curved. First submarginal cell lon-
ger than second and third combined; sec-
ond narrowed anteriorly; anterior border
of third along Rs about as long as anterior
border of second, posterior border of third
2 times longer than anterior border; mar-
ginal cell length 2.5, width 0.5. Distal ham-
uli arranged 3-1-3. Male terminalia as in
figures 10 and 11.

Clypeus and supraclypeal area finely
imbricate, with a few weak punctures sep-
arated by 1-2 puncture widths on lateral
borders. Face imbricate. Vertex smooth
and impunctate. Gena sparsely and weak-
ly nodulate, otherwise smooth. Postgena
smooth. Pronotum smooth. Mesoscutum
minutely punctured, punctures separated
by 2-3 times puncture width, integument
between punctures smooth and shining.
Scutellum as on mesoscutum. Metanotum
sparsely and weakly nodulate, integument
otherwise smooth. Pre-episternum and
mesepisternum minutely punctured,
punctures separated by 3-4 times punc-

Volume 7, Number 1, 1998

ture width, integument between smooth;
hypoepimeron as on mesepisternum ex-
cept minute punctures separated by 4
times puncture width. Metepisternum
smooth. Propodeal triangle imbricate; lat-
eral surface imbricate, with punctures sep-
arated by 2-3 puncture widths; posterior
surface smooth. Terga and sterna imbri-
cate, except Tl which is smooth.

Head dark brown with strong metallic
green highlights; labrum and clypeus am-
ber; mandible amber except red at apex;
antennae brown. Mesosoma dark brown
with strong metallic green highlights, ex-
cept pronotum light brown and without
such highlights. Legs pale brown. Meta-
soma dark brown.

Pubescence golden. Scattered simple
hairs (1-1.5 o.d.) on face, with suberect
branched hairs (0.5 o.d.) along inner mar-
gins of compound eyes. Similar suberect
and simple hairs on gena, branched hairs
becoming longer towards postgena. Post-
gena with scattered branched hairs (2
o.d.). Pronotum with scattered simple
hairs (1 o.d.) and laterally with extremely
short, appressed hairs. Mesoscutum with
scattered simple hairs (1 o.d.) and shorter
(less than 0.5 o.d.) branched hairs on lat-
eral and posterior borders. Scutellum and
metanotum with sparse simple hairs (1.5-
2 o.d.) and more numerous shorter simple
hairs (0.5 o.d.). Pleura with scattered sim-
ple pale hairs (1-1.5 o.d.) and shorter (0.5
o.d. or less) suberect branched hairs; me-
tepisternum without longer simple hairs.
Pubescence of legs simple (1-2 o.d.) Terga
with scattered simple hair (1 o.d.), becom-
ing longer on lateral margins and apical
terga. Sterna with sparse simple hair (0.5-
1.5 o.d.); S4 with a few simple hairs (0.5
o.d.) concentrated on borders of apical de-
pression (Fig. 9); hairs of S6 most numer-
ous, laterally more dense than median
area.

FEMALE: As for the male with the
modifications indicated below. Measure-
ments from female type of M. angusticeps.
Total body length 12.2 mm; forewing

length 8.7 mm. Head as in figures 1 and
2. Supraclypeal area 0.8 times length of
clypeus. Head width 3.0; length (to apex
of clypeus) 2.7; clypeal length 0.8; lower
interorbital distance 0.9; upper interorbital
distance 1.1; interantennal distance 0.3; an-
tennocellar distance (to median ocellus)
0.6; between lateral ocelli 0.3; median ocel-
lus to lateral ocellus 0.04; lateral ocellus to
compound eye 0.08; prementum length
2.2, width 0.2. Scape reaching past lateral
ocellus, length 1.4; pedicel slightly longer
than wide, length 0.2, width 0.18; Fl lon-
ger than wide and longer than F2, length
0.24, width 0.22; F2 wider than long,
length 0.2, width 0.22; F3 and F4 each as
long as wide, individual lengths-widths
0.22; F5 and F6 longer than wide, lengths
0.24, widths 0.22; F7 and F8 longer than
wide, lengths 0.26, widths 0.22; F9 longer
than wide, length 0.3, width 0.22; FIO lon-
ger than wide and longer than previous
flagellomeres, length 0.44, width 0.22. In-
tertegular distance 2.1. Wing as in figure
3; basal vein distad cu-a by vein width; Ir-
m confluent with Im-cu; 2r-m distad 2m-
cu by 5 times vein width, 2r-m weakly
curved. First submarginal cell longer than
second and third combined; second nar-
rowed anteriorly; anterior border of third
along Rs slightly longer than anterior bor-
der of second, posterior border of third 2
times longer than anterior border; margin-
al cell length 2.6, width 0.6. Distal hamuli
arranged 3-1-3.

Mesoscutum minutely punctured,
punctures separated by 3^ times punc-
ture width, integument between punc-
tures weakly imbricate and shining. Terga
and sterna weakly imbricate, except Tl
which is smooth.

Head dark brown with strong metallic
green highlights; labrum and clypeus am-
ber; supraclypeal area light brown medi-
ally; mandible amber except black at apex.
Mesoscutum, scutellum, metanotum and
propodeum brown with strong metallic
green highlights; pronotum and pleura
amber with metallic green highlights

10

Journal of Hymenoptera Research

fainter than those of mesoscutum. Legs
amber. Terga amber, except apical mar-
gins brown; sterna amber.

Pubescence golden. Scattered simple
hairs (1-1.5 o.d.) on face. Gena and post-
gena with branched hairs (1.5-2 o.d.). Pu-
bescence of fore- and mid-legs as in male;
hind legs with scopa formed of plumose
hairs (2.5-3.5 o.d.) on trochanter, femur
and imier margin of tibia. Terga with scat-
tered simple hair (1-1.5 o.d.), becoming
longer on lateral margins and apical terga.
Sterna with sparse simple hair (1-1.5 o.d.).

Type material. — BRAZIL: Amazonas:
Holotype, male. Faro, 23 January 1910,
Ducke (ZMHB).

Additional material. — BRAZIL: Amazo-
nas: 1 male, Parintins, 9 October 1919, Par-
ish (FMNH). 1 male, Beruri, Rio Purus, 15
October 1991, G.A.R. Melo, sitting under
leaf (GARM). 6 females, Hwy. BR 174, 86
km N Manaus, ZF6 km 9, 2°16'S, 59°56'W,
3 July 1986, M.V.B. Garcia, attracted to
propane lantern at night (UFVB); 5 fe-
males, same except 4 July 1986 (UFVB); 2
females, same except (SEMC). 3 females,
Reserva Ducke, 26 km NE Manaus, Itacoa-
tiara Hwy., 12-23 May 1972, Munroe
(SEMC). 2 males, Autaz-Mirim, Faz. Sao
Lucas, 25 August 1994, Joao Vidal, malaise
trap (INPA); 2 males, same except
(SEMC); 1 male, same except 28 October
1994 (INPA). 1 female. Campus Universi-
tario, Manaus, 7-24 September 1988, M.
Castilho, J. Elias Bindo, Shannon trap,
baited with feces (INPA). 2 females, Mun.
Itacoatiara, Faz. Aruana, AMOIO km 215,
19-21 September 1990, C Motta, R. An-
dreazze, R. Ferreira (INPA); 2 females,
same except 18-19 September 1990, light
trap (INPA). 1 female, Reserva Ducke, 9-
22 September 1994, J. Rafael, J. Vidal, sus-
pended malaise trap, 20 m (INPA). 1 fe-
male, Manaus, Campus Universitario, 21
June 1982, J. Rafael, malaise trap (INPA).
1 female, 26 km NE Manaus, Reserva
Ducke, 6 October 1988, J. Raphael, sus-
pended malaise trap, 10 m (INPA); 1 fe-
male, same except 9 May 1978, light trap.

J. Arias, N. Penny (INPA). 1 female, Rio
Jau, Meriti, Mun. Novo Airao, 4-10 June
1994, J. Raphael (INPA). Para: 4, females,
Val de Cans. Belem., 20-21 November
1963, Oliveira, Wygodzinsky, at light
(AMNH). 1 male. Baker (LACM). CO-
LOMBIA: Dpto. Guaviare: 1 female, 1
male, Nukak Maku, Cerro Moyano, 200
m, 12 November 1995, malaise trap, F.
Fernandez (UNCB). ECUADOR: Sucum-
bios: 1 female, 0.5°S, 76.5°W, 290 m, Sacha
Lodge, 13-23 June 1994, P. Hibbs, malaise
trap (LACM). FRENCH GUIANA: 1 fe-
male (holotype of Megaloptidia angusti-
cep^s), Nouveau Chantier, Collection le
Moult, June (ZMHB). 1 female, Saint-Lau-
rent, du Maroni (AMNH). 1 female, Saiil,
3°37'N, 53°12'W, 13-18 April 1983, S.
Mori, ex: light trap (AMNH). 2 males,
Roura, 18.4 km SSE, 240 m., 4°36'38"N,
52°13'25"W, 25-29 May 1997, J.S. Ashe and
R.W. Brooks, FG1AB97-081, ex: flight in-
tercept trap (SEMC). 1 male, Roura, 27.4
km SSE, 280 m., 4°44'20"N, 52°13'25"W,
23-24 May 1997, J.S. Ashe and R.W.
Brooks, FG1AB97-022, ex: flight intercept
trap (SEMC). 1 male, Roura, 27.4 km SSE,
280 m., 4°44'20"N, 52°13'25"W, 10 June
1997, J.S. Ashe and R.W. Brooks,
FG1AB97-177, ex: flight intercept trap
(SEMC). 1 male, Matoury, 41.5 km SSW on
Hwy N2, 50 m., 4°37'22"N, 52°22'35"W, 29
May-9 June 1997, J.S. Ashe and R.W.
Brooks, FG1AB97-170, ex: flight intercept
trap (SEMC). GUYANA: Mazaruni-Potaro
District: 1 female, Kartabo Point, 25-27
December 1983, W. Steiner, J. Byrd, J. Hill,
F. Holtzclaw, malaise trap at edge of sec-
ondary forest and farmed field (USNM).
PERU: Cuzco: 1 male. Quince Mil, 750 m,
September 1962, L. Pefia (CUIC). Huanu-
co: 1 female, Tingo Maria, 21 February
1972, J. Schuster, on flower of Dichorisnii-
dra iilei, 5:53am (CUIC). Madre de Dios: 1
male, Estacion Pakitza, Reserva Manu, 18
June-4 July 1993, R. Cambra, T. Amorilla
(MIUP). 1 female, Rio Tambopata Res., 30
air km SW Pto. Maldonaldo, 290 m, 16-20
November 1979, J. Heppner, subtropical

Volume 7, Number 1, 1998

11

19

Figs. 18-19. Mcgnloi'tkiia saulensis Engel c& Brooks n. sp., 18, 19, female head, front and side views respec-
tively.

moist forest (USNM). SURINAME: 3 fe-
males, Raleigh Vallen-Voltzberg Research
Foengoe 4°43'N, 56°12'W, 26 January-15
February 1982, J. Carpenter, D. Trail
(CUIC). VENEZUELA: Amazonas: 2 fe-
males, San Carlos de Rio Negro, 1°56'N,
67°3'W, 6-12 December 1984, R. Brown
(CUIC). 3 females. Camp IV, 0°58'N,
65°57'W, Cerro de la Neblina, 760 m, IS-
IS March 1984, O.S. Flint, Jr., malaise trap
over dry stream channel (USNM).

Flower records. — One female of M. noc-
tunia has been collected on Dichorisandra
iilei (Monocotyledonae: Commelinaceae),
although there was no pollen in her scopa.
This is the first floral association recorded
for any species of Megaloptidia.

Variation. — As typical of nocturnal bees,
Megaloptidia nocturna is lightly melanized
throughout most of its Amazonian range.
However, at the western edge of its dis-
tribution the male specimen from Colom-
bia is light yellow brown and noticeably
lighter than the brown Brazilian male
specimens. This is odd since a female
caught in the same trap on the same day
is darker than the normal color of the Bra-
zilian female specimens. The male speci-
men from Peru, Madre de Dios is almost

black and much darker than the Brazilian
material.

Megaloptidia saulensis Engel and
Brooks, new species

(Figs. 18-20)

Diagnosis. — Labrum and clypeus dark
brown. Mesoscutum weakly punctured,
punctures separated by 2-3 times a punc-
ture width over central disk, laterally
punctures separated by less than a punc-
ture width. Mesepisternum closely and
weakly punctured, punctures separated
by less than a puncture width; hypoepi-
meron with minute punctures separated
by 2-3 times a puncture width. Scopal
hairs of hind tibia black, those of femur
amber. Propodeal lateral surface imbricate
with punctures separated by 2-3 times a
puncture width; length of propodeal tri-
angle little to much greater than that of
metanotum. This species is only known on
the basis of two females.

Description. — The following description
is based on the holotype. FEMALE: Total
body length 12.4 mm; forewing length 8.8
mm. Head wider than long (Fig. 18). Dis-
tal third of clypeus projecting below lower
tangent of compound eye (Figs. 18, 19);

12

Journal of Hymenoptera Research

supraclypeal area wider than long, 0.58
times length of clypeus. Frontal line cari-
nate from below antennal sockets to just
above sockets, ending with an acute pro-
jection and protuberant in profile (Fig. 19).
Measurements of head of holotype in mm:
width 3.4; length (to apex of clypeus) 2.8;
clypeal length 1.0; lower interorbital dis-
tance 1.1; upper interorbital distance 1.4;
interantennal distance 0.4; antennocellar
distance (to median ocellus) 0.6; between
lateral ocelli 0.4; median ocellus to lateral
ocellus 0.08; lateral ocellus to compound
eye 0.2 mm; prementum length 2.0, width
0.2. Scape reaching past lateral ocellus,
length 1.5; pedicel slightly longer than
wide, length 0.24, width 0.18; Fl longer
than wide and longer than F2, length 0.26,
width 0.22 (Fig. 18); F2 as long as wide,
length-width 0.2; F3-6 each longer than
wide, individual lengths 0.22, widths 0.2;
F7 and F8 each longer than wide, individ-
ual lengths 0.24, widths 0.22; F9 longer
than wide, individual lengths 0.26, widths
0.24; FIO longer than wide and longer than
preceding flagellomeres, length 0.4, width
0.24. Median line strongly impressed; par-
apsidal lines weak. Intertegular distance
2.5. Propodeal triangle slightly longer than
metanotum, about half length of scutel-
lum. Basal vein distad of cu-a by vein
width; Ir-m basad Im-cu by vein width;
2r-m distad 2m-cu by 3 times vein width,
2r-m straight. First submarginal cell lon-
ger than second and third combined; sec-
ond narrowed anteriorly; anterior border
of third along Rs as long as anterior bor-
der of second, posterior border of third 2.5
times longer than anterior border; margin-
al cell length 2.4, width 0.6. Distal hamuli
arranged 4-1-1-1-4.

Clypeus and supraclypeal area finely
imbricate, with a few weak punctures sep-
arated by 1-2 puncture widths on lateral
borders. Face imbricate with a few weak
punctures along border with compound
eye. Vertex, gena, and postgena smooth
and impunctate. Pronotum weakly imbri-
cate. Mesoscutum punctured, punctures

separated by 2-3 times puncture width,
integument between punctures imbricate
and shining, except laterally punctures
separated by less than a puncture width.
Scutellum with sparse, minute punctures,
integument smooth. Metanotum sparsely
and weakly nodulate, integument other-
wise weakly imbricate. Pre-episternum
and mesepisternum punctured, punctures
separated by less than a puncture width,
integument between smooth; hypoepime-
ron as on mesepisternum except minute
punctures separated by 2-3 times punc-
ture width. Metepisternum smooth with a
few minute punctures sparsely scattered.
Propodeal triangle imbricate; lateral sur-
face imbricate with punctures separated
by 2-3 puncture widths; posterior surface
smooth. Terga and sterna imbricate.

Head dark brown with metallic blue-
green highlights, except labrum, mandi-
ble, clypeus, supraclypeal area without
such highlights. Mesosoma dark brown
without highlights, except scutellum,
metanotum, and propodeal triangle am-
ber. Legs brown. Metasoma dark brown.

Pubescence pale to golden. Scattered
simple hairs (1-1.5 o.d.) on face, with sub-
erect branched hairs (0.5 o.d.) along inner
margins of compound eyes. Similar sim-
ple hairs on gena, hairs becoming longer
towards postgena; postgena with scat-
tered simple hairs (2 o.d.). Pronotum with
scattered simple hairs (1 o.d.) and laterally
with extremely short, appressed hairs.
Mesoscutum with scattered simple hairs
(1 o.d.) and shorter (less than 0.5 o.d.)
branched hairs on lateral and posterior
borders. Scutellum with sparse simple
hairs (1.5-2 o.d.) and more numerous
shorter simple hairs (0.5 o.d.). Longer
hairs of mesoscutum and scutellum fus-
cous. Metanotum with sparse simple hairs
(1.5-2 o.d.). Pleura with scattered simple
hairs (1-1.5 o.d.) and shorter (0.5 o.d. or
less) suberect branched hairs; metepister-
num with longer simple hairs more dense
on ventral half; pubescence of pre-epister-
num fuscous, remainder pale. Pubescence

Volume 7, Number 1, 1998

13

Fig. 20. Distribution of Mcgnloptidia species. Circles represent locality records for M. iioctunuj (Friese), squares
for M. contradictn (Cockerell), and the single triangle for M. saulensis Engel & Brooks.

of legs simple and amber (1-2 o.d.), except
tarsomeres 1 and 2 (of all legs), apex of
protibia, and entirety of mesotibia and
metatibia, including tibial scopa. Terga
with scattered simple hairs (1 o.d.), be-
coming progressively longer on lateral
margins and more distal terga. Sterna
with sparse simple hairs (1.5-2.5 o.d.),
most with a few branches.

MALE: Unknown.

Type material.— FRENCH GUIANA: Ho-
lotype, female, Saiil, 3°37'N, 53°12'W,
1983, S. Mori, #158 (AMNH). 1 paratype
female, Saul, 3°37'N, 53°12'W, 1983, S.
Mori, #157 (AMNH).

Etymology. — The specific epithet is de-
rived from the type locality of the species.

SPECIES MISTAKENLY PLACED IN
MEGALOPTIDIA

Moure and Hurd (1987), in a catalog of
the Western Hemisphere halictid bee spe-

cies, tentatively placed Megalopta pallitar-
siis Friese as a species of Megaloptidia. In
actuality, examination of the holotype
male for M. pallitarsus (ZMHB) reveals
that this species is a junior synonym of
Megommatioii insigne (Smith 1853). We
therefore propose the following new syn-
onymy:

Megommation insigne (Smith 1853)

Halictus hisigiiif Smith 1853: 65.

Halictus cherazon Vachal 1904: 113.

Megalopta I'irgili Friese 1911: 454.

Megalopta {Megaloptella) iponweae Schrottky
1912: 271.

Megalopta pallitarsus Friese 1926: 125. New syn-
onymy.

Megaloptidia pallitarsus (Friese); Moure and
Hurd 1987: 242.

ACKNOWLEDGMENTS

We gratefully acknowledge the curators listed
above for the sundry loans of material which made

14

Journal of Hymenoptera Research

this study possible. We are thankful to D.B. Baker,
E.E. Grissell, R.J. McGinley, and B.B. Norden for crit-
ically reviewing previous versions of this paper.
Their valuable criticisms served to improve the pre-
sentation of this study. C.R. Hardy graciously shared
with us his knowledge of Commelinaceae biology,
morphology, and systematics. MSE was supported by
a National Science Foundation Predoctoral Fellow-
ship during this work. The generic description pre-
sented herein is taken from a monograph of the gen-
era and subgenera of Augochlorini being completed
by MSE as part of a Ph.D. dissertation at Cornell Uni-
versity. This paper is contribution Nr. 3186, Division
of Entomology, Snow Entomological Collection, Uni-
versity of Kansas Natural History Museum.

LITERATURE CITED

Cockerell, T. D. A. 1900. Descriptions of new bees
collected by Mr. H.H. Smith in Brazil. 1. Proceed-
ings of the Academy of Natural Sciences of Pliiliulel-
phia 52:356-377.

Eickwort, G. C. 1969. A comparative morphological
study and generic revision of the augochlorine
bees (Hymenoptera: Halictidae). University of
Kansas Science Bulletin 48:325-524.

Fabricius, J. C. 1804. Systenia piezatorum secundum or-
dines, genera, species adiectis synonymis, locis, oh-
servationihus, descriptionibus. Reichard, Brunsvi-
gae, xiv + [15]-439, 1-30 pp.

Friese, H. 1911. Neue Bienen aus Siid-Amerika
(Hym.). Deutsche Entomologische Zeitschrift 1911:
453-156.

Friese, H. 1926. Die Nachtbienen-Gattung Megalopta
Sm. Stettiner Entomologische Zeitung 87:111-135.

Jorgensen, P. 1912. Beitrag zur Biologie einiger sii-
damerikanischer Bienen. Zeitschrift fiir loissen-
schaftliche Insektenbiologie 8:268-271.

Michener, C. D. 1962. An interesting method of pol-
len collecting by bees from flowers with tubular
anthers. Revista de Biotogia Tropical 10:167-175.

Moure, ]. S. 1943. Notas sobre abelhas da cole^ao de

Zikan (Hym. Apoidea). Revista de Entoiiiologia 14:
447^84. '

Moure, J. S. 1951. Ariphanarthra, um novo genero de
Halictidae (Hymenopt. — Apoidea). Dusenia 2:
137-140.

Moure, J. S. 1958. On the species of Megalopta de-
scribed by F. Smith (Hymenoptera, Apoidea).
journal of the New York Entomological Society 66:
179-190.

Moure, J. S. 1969. Micrommation, novo genero de Hal-
ictidae do Parana (Hym. Apoidea). Atas Sociedade
de Biologia 12:247-249.

Moure, J. S., and P. D. Hurd, Jr. 1987. An annotated
catalog of the halictid bees of the Western Hemisphere
(Hymenoptera: Halictidae). Smithsonian Institution
Press, Washington, vii + 405 pp.

Rego, M. M. C, and P. M. C. de Albuquerque. 1989.
Comportamento das abelhas visitantes de Muri-
el, Byrsonima crassifolia (L.) Kunth, Malpighi-
aceae. Boletim do Museu Paraense Emilio Goeldi,
Zoologia 5: 179-194.

Renner, S. S. 1989. Floral biological observations in
Heliamphora tatei (Sarraceniaceae) and other
plants from Cerro de la Neblina in Venezuela.
Plant Systematics and Evolution 163:21-30.

Schrottky, C. 1902. Ensaio sobre as abelhas solitarias
do Brasil. Revista do Museu Paulista 5:330-613.

Schrottky, C. 1912. Beschreibung von Megalopta (Me-
galoptella) ipomoeae n. sp. Zeitschrift fur Wissen-
schaftliclien Insektenbiologie 8:271-272.

Sigrist, M. R., and M. Sazima. 1991. Biologia floral e
poliniza^ao por vibra^ao em duas especies sim-
patricas de Dichorisandra (Commelinaceae). XLII
Congresso Nacional da Sociedade Botanica do Brasil,
Sociedade Botanica do Brasil, Universidade Federal de
Golds 1991:484.

Smith, F. 1853. Catalogue ofhymenopterous insects in the
collection of the British Museum, part 1: Andrenidae
and Apidac. British Museum, London, 197 pp., 6
pis.

Vachal, J. 1904. Etude sur les Halictiis d'Amerique
(Hym.). Miscellanea Entomologica 12:9-24, 113-
128, 137-144.

J. HYM. RES.
Vol. 7(1), 1998, pp. 15-24

Species Richness of Costa Rican Cenocoeliini (Hymenoptera:

Braconidae): a Latitudinal and Altitudinal Search for

Anomalous Diversity

Leendert-Jan van DF.R Ent and Scott R. Shaw

Department of Plant, Soil and Insect Sciences, P.O. Box 3354, University of Wyoming,

Laramie, WY 82071, USA

Abstract. — Latitudinal patterns of species diversity of New World Braconidae have been scarcely
surveyed to date. Such patterns may be of biogeographical and ecological interest because some
literature data suggest that some braconid subfamilies do not show an increase in species diversity
towards the equator despite an increase of potential host species (i.e., "anomalous diversity"). In
the present study, species diversity of a "presumptive" anomalous diverse braconid taxa, Ceno-
coeliini, was surveyed in Costa Rica. The results were compared with published distribution data
of North American Cenocoeliini. Also, species richness and abundance of Cenocoeliini from sea-
level to 3400 m altitude in Costa Rica were analyzed to compare latitudinal and altitudinal gra-
dients in species diversity. Costa Rican Cenocoeliini were five times more speciose than those in
Canada and USA combined. The increase in estimated species richness per unit area towards the
equator of North American Cenocoeliini was similar to that of their most common hosts, Cer-
ambycidae and Scolytidae, but exceeded that of the potential hosts in Costa Rica. Diversification
in Costa Rican Cenocoeliini was partly influenced by adaptation to different host families and
host substrates. Most species and individuals of Cenocoeliini were found at low altitudes (<500
m) in Costa Rica. Cenocoeliini were not encountered above 1600 m in Costa Rica, this being in
contrast to their most likely hosts, Cerambycidae and Scolytidae, which also occurred at high
altitudes. Larger-sized Costa Rican Cenocoeliini were often brightly colored suggesting an apo-
sematic function towards visually-oriented predators. New World Cenocoeliini appeared to be
tropical lowland-centered and this is expected to be rather an effect of temperature requirements
than an effect of host-limitation.

Anomalous diversity is defined as a pat- the tropics may provide biased results for

tern in species richness "counter to the large-sized parasitoids (>3 mm) in areas

prevalent trend of increasing species num- where small-sized parasitoids may be

ber in a taxon with decreasing latitude" abundant (Hespenheide 1979; Morrisson

(Rathcke and Price 1976). Owen and et al. 1979). More recently, results of Mal-

Owen (1974) were the first investigators to aise trap sampling revealed that some

show anomalous diversity for parasitic subfamilies of Ichneumonidae displayed

Hymenoptera of the family Ichneumoni- anomalous diversity (Gauld 1986, 1987,

dae, despite an increase of numbers of po- 1995b). Several theories to explain latitu-

tential host species towards the equator, dinal patterns in species diversity of par-

Janzen and Pond (1975) found a similar asitic Hymenoptera have been formulated:

pattern in species richness for parasitic resource fragmentation (Janzen and Pond

Hymenoptera; they were less or equally 1975; Janzen 1981), predation on hosts

speciose in Costa Rica compared to those (Rathcke and Price 1976), predation on

occurring in a meadow in England. Sev- parasitoids (Gauld 1987), and the "nasty"

eral investigators noted that sweep-sam- host hypothesis (Gauld ct al. 1992; Gauld

pie studies of parasitic Hymenoptera in and Gaston 1994). According to these the-

16

Journal of Hymenoptera Research

ories, different ecological groups of para-
sitic Hymenoptera are expected to show
different patterns in latitudinal species di-
versity (Hawkins 1994).

To date, anomalous diversity has not
been fully analyzed for New World Bra-
conidae. Quicke and Kruft (1995) found
some subfamilies of Braconidae (e.g., Aly-
siinae, Aphidiinae, Cheloninae) to be less
speciose from northern (zone 3, 4) towards
southern (zone 5) latitudes in the USA.
This suggests anomalous diversity, how-
ever, these analyses did not include neo-
tropical regions. Species richness of anoth-
er group, Cenocoeliinae (i.e., Cenocoeli-
ini), increased from northern to southern
temperate regions, but did not increase
from middle to lower latitudes in the USA
(Quicke and Kruft 1995). Also, species
richness of Cenocoeliini in southern tem-
perate regions was equal to that of north-
ern and central Mexico combined and to
that of southern Mexico (data from Saffer
1982). To date, not more than two species
of Cenocoeliini have been described from
Costa Rica (Saffer 1977, 1982). Previous
data, therefore, suggest a pattern of anom-
alous diversity in Cenocoeliini. The distri-
bution of Central American Cenocoeliini,
however, is less well documented than
that of North America (Saffer 1982).

Cenocoeliini are diurnally active endo-
parasitic koinobionts of endophytic beetle
larvae (van Achterberg 1994; Saffer 1982).
In North America, Cenocoeliini were
found to parasitize Cerambycidae (68% of
recorded host species), Scolytidae (22%),
and Buprestidae (Saffer 1982). Koino-
bionts are parasitoids which let their hosts
continue to be mobile and defend them-
selves for a while after being parasitized.
Koinobionts are expected to have more
narrow host ranges than idiobionts
(Askew and Shaw 1986; Gauld 1987; Haw-
kins 1994) and are often referred to as
"specialist" parasitoids. Sheehan and
Hawkins (1991), however, noted that com-
parisons of average host ranges between
koinobionts and idiobionts have to be

evaluated carefully. Specialist parasitoids
(i.e., koinobionts) associated to endophytic
hosts may show, as predicted by the the-
ories of resource fragmentation and pre-
dation on parasitoids, a (very) strong de-
crease in species richness from the tem-
perate to tropical zones (Table 4.2, Haw-
kins 1994). At the other hand, these
parasitoids may also show, as predicted
by the theory of predation on hosts and
that of the "nasty" hosts, a weak decrease
to increase in species richness towards the
equator. Thus, based on parasitizing be-
havior, contrasting predictions could be
made for species richness of Costa Rican
Cenocoeliini.

Species richness along altitudinal gra-
dients may demonstrate patterns similar
to those of latitudinal gradients (Brown
1988; Stevens 1992). Noyes (1989) noted
that in general species diversity of para-
sitic Hymenoptera in an Indonesian rain
forest was the highest at low altitudes (<
1000 m). Also the highest diversity of their
hosts, e.g. Lepidoptera, was found at low
altitudes (Holloway 1986). This suggests
that, if patterns in latitudinal and altitu-
dinal species richness are similar, anoma-
lous diversity should not occur among
parasitic Hymenoptera. Results of an alti-
tudinal transect study in the Venezuelan
Andes by Janzen et al. (1976) showed that
species richness of parasitic Hymenoptera
was as high at 200 m as at 1600 m, but
that it was lower at high altitudes (3550
and 3600 m). The decline of species rich-
ness of parasitic Hymenoptera between
1600 and 3550 m was smaller than the de-
cline of species richness of most other
groups of insects such as other Hymenop-
tera (ants, bees, aculeate wasps) and other
insect orders (e.g., Coleoptera and Lepi-
doptera). This suggests that some groups
of parasitic Hymenoptera show anoma-
lous diversity along altitudinal gradients.
At a lower taxonomic level, tropical alti-
tudinal species diversity of Ichneumono-
idea was found to differ among subfami-
lies (Gauld 1985; Gaston and Gauld 1993;

Volume 7, Number 1, 1998

17

Gauld and Hanson, in press). Also,
Ophioninae were found to have different
patterns in altitudinal species diversities
between tropical regions (Gauld and Han-
son, in press).

As noted earlier, anomalous diversity
has not been completely analyzed for New
World Braconidae. Literature data suggest
that the braconid tribe Cenocoeliini may
show a decrease, or at least no increase, in
species diversity towards the equator.
Therefore, the aims of the present study
were: 1) to determine whether species
richness and a-index of diversity of the
Cenocoeliini in temperate regions were
higher than those in tropical regions, 2) to
determine whether potential hosts of Cen-
ocoeliini increased more in species rich-
ness towards the equator than their para-
sitoids, 3) to relate latitudinal species rich-
ness with altitudinal species richness, and
4) to determine whether abundance and
species richness of Cenocoeliini were
higher at intermediate and high than at
low altitudes in Costa Rica. This research
is the first in a series of analyses of differ-
ent groups of Braconidae to be examined
for species diversity in Costa Rica in re-
lation to altitudes.

MATERIALS AND METHODS

The tribe Cenocoeliini is a monophyletic
group in the Helconinae (Shaw 1995), al-
though several authors placed the Ceno-
coeliini as the main tribe in a separate sub-
family, Cenocoeliinae (van Achterberg
1984, 'l993; Shaw and Huddleston 1991;
Wharton 1993). Before the generic revision
by van Achterberg (1994), most Cenocoe-
liini species were considered to be within
the genus Cenocoelius.

Individuals of Cenocoeliini were col-
lected with Malaise traps (85%) and the
rest by hand netting in Costa Rica, mostly
during the last 10 years. Hand collected
specimens for this study were from H.A.
Hespenheide (University of California,
Los Angeles), F. Parker (University of
Utah) and J.A. Ugalde (INBio, Instituto

Nacional de Biodiversidad, Costa Rica).
Malaise traps were located in different
habitat types and at different altitudes
throughout Costa Rica as described by
Gauld (1991, 1995a). Our analyses includ-
ed ca 70 Malaise trap-years of samplings
at ca 60 different sites. Therefore, sample
coverage was expected to be reasonably
representative for the Costa Rican fauna.
From these samples, Braconidae were
sorted and sent to the University of Wy-
oming for identification. Cenocoeliini
were sorted and individuals were deter-
mined to morphospecies using the follow-
ing set of characters: hindwing venation
(relative length vein IM-l-CU to IM, and
IM to Ir-m); color patterns of head, me-
sosoma, metasoma, legs, ovipositor
sheaths and antennae; body size; oviposi-
tor length relative to forewing length;
number of flagellomeres; and shape of
apical flagellomeres. Additional charac-
ters, like sculpture patterns on the pro-
episternum, apex of the propodeum, and
vertex, were included to distinguish
among presumptive sibling species com-
plexes. Data by Saffer (1982) were used to
compare species richness of Braconidae in
North America with that in Costa Rica.
Sample coverage and sample intensity for
Mexico was relatively low and less rep-
resentative for the area than those of Can-
ada, USA and Costa Rica.

Two formulae were used to estimate ex-
pected species richness of the faunae (S')
based on the numbers of individuals per
species in a sample:

1. S^' = S-/(S - S,) (S, = total number of
species with one individual)

2. S"- = S + ((S,)V2S,) (S; = total number
of species with two individuals)

The first formula of expected species rich-
ness (S'') is derived from the formula of
sample coverage (1-(N,/I): Fagen and
Goldmann 1977). In this formula, I is the
total number of behavior types observed
and N| is the total behavior types ob-

18

Journal of Hymenoptera Research

served only once. I was substituted with S
and N, with S,. S""' was calculated as the
inverse of the sample coverage multiplied
by the number of observed species (S).
The second formula of expected species
richness (S'"'), is that of Chao 1 as de-
scribed in Colwell and Coddington (1994).
In addition to species richness, a-index of
diversity of the logarithmic series was cal-
culated because of its good discriminant
ability and its low sensitivity to sample
size (Magurran 1988). To estimate species
richness per unit area the formula S, =
x/a°-^ was used (S, = number of species
per unit area, x = number of observed
species in country or region, a = area of
country or region (10' km-): MacArthur
and Wilson 1967, Gaston et al. 1996). We
also estimated species richness per unit
area for the most important temperate
hosts of Cenocoeliini, the Cerambycidae
(data from Monne and Giesbert 1994) and
the Scolytidae (data from Wood 1982;
Wood et al. 1991).

To examine the effect of altitude on spe-
cies richness of Cenocoeliini, we defined 4
altitude classes: low (0-500 m), low inter-
mediate (500-1500 m), high intermediate
(1500-2500 m) and high (>2500 m) alti-
tudes. These altitude classes reflect the
distribution of different habitat types as
described by Gauld (1995a). We assumed
that there was a linear relationship be-
tween sample effort (i.e.. Malaise trap-
months) and number of individuals
caught in Malaise traps. Because seasonal
variation in abundance of neotropical in-
sects occurs (Owen and Chanter 1970;
Wolda 1988, 1989; Wolda and Wong 1988),
only Malaise traps which operated three
or more consecutive months were includ-
ed in the analysis, and abundances of Cen-
ocoeliini were summed for several year-
round Malaise trap samples. We estimated
expected numbers of Cenocoeliini per al-
titude class by multiplying the total num-
ber of observed Cenocoeliini with the pro-
portion of number of Malaise trap-months
of a particular altitude class to the total

Table 1. Numbers of individuals (N), species rich-
ness (S), two estimates of expected species richness
(S"', S'-; see materials and methods), a-index of di-
versity, numbers of localities (loc) and averaged local
species richness (S,,^) of Cenocoeliini from Canada
and USA combined, Mexico and Costa Rica.

Canada + USA

MfMO.

e i.sla Rica

N

1108

51

301

S

11

13

57

S"

11

24

96

S"=

11

22

123

a-index

1.665

5.291

20.822

loc

233

13

48

s„„

1.1 ± 0.3

1.5 ± 1.0

2.8 ± 4.4

(range)

1-3

1-4

1-25

number of Malaise trap-months. We test-
ed whether observed and expected num-
bers were equal between low (<500 m)
and higher altitude classes combined us-
ing a chi-square test (Sokal and Rohlf
1981).

RESULTS

In total, 290 individuals of Cenocoeliini
were found from 80,000 to 90,000 Bracon-
idae sampled in Costa Rica. The individ-
uals of Cenocoeliini were divided into 55
morphospecies. The two described Costa
Rican Cenocoeliini species by Saffer (1977,
1982) did not match our individuals and
were added to our morphospecies result-
ing in a total of 57 species and 301 indi-
viduals (Table 1). From this set of 57 spe-
cies, 44 species belonged to Capitoiiius, and
7 species to Ceiiocoelius. Of the remaining
species, 4 species may belong to Capitonius
but are rather distinctive and 2 species be-
long to a new genus of Cenocoeliini. The
species varied in size from 3 to 12 mm.
The ovipositors were relatively long, 0.7 to
1.7 times the forewing length. About half
of the individuals and species were rela-
tively large, mostly >5 mm, and bright
yellow-orange to red-orange and often
partly black in color, with partly to com-
pletely darkened wings. Most other indi-
viduals were relatively small, <7 mm, and
blackish, brownish or black and dull red

Volume 7, Number 1, 1998

19

Table 2. Estimated species richness per unit area (S,) of parasitoids (Cenocoeliini) and their most common
temperate hosts (Cerambycidae and Scolytidae) for different geographical regions of North and Central Amer-
ica, using S., = x/a"-' as a standard species-area relationship (see text). Between parentheses are the relative
richnesses when the estimated species richness of Canada was set at 1.0.

Geographical
region

Area
(10' km-')

Cenocoeliini
S.

Cerambycidae
S,"

ScolvHciae

Canada ( + Alaska)

11,49b

0.39 (1.0)

29.4 (1.0)

17.3(1.0)

USA (-Alaska)

7,828

1.17 (3.0)

85.8 (2.9)

50.3 (2.9)

Mexico

1,969

1.95 (5.0)

201.2 (6.8)

90.8 (5.2)

Costa Rica

5\

21.36 (54.8)

260.1 (8.8)

155.3 (9.0)

to reddish brown in color with clear
wings, sometimes with brownish spots on
the forewings.

Highest species richness was found in
Costa Rica and lowest species richness in
Canada and USA combined (Table 1).
Both estimators of expected species rich-
ness (S''' and S'-) predicted that in Mexico
and Costa Rica about half of the total
number of species have been caught to
date. More species were found in 2 Costa
Rican Malaise trap sites (Heredia Prov-
ince, Biological Station of OTS La Selva,
and Puntarenas Province, 24 km west of
Piedras Blancas) than in all trap sites of
Mexico or Canada and USA combined.
The highest diversity (a-index) was found
for Costa Rican Cenocoeliini, the lowest
for those of Canada and USA combined.

The increase of estimated species rich-
ness per unit area for North American
Cenocoeliini was similar to that of their
most common hosts (Table 2). On average,
3 times more species per unit area oc-
curred in the USA than in Canada and
about twice as many species per unit area
occurred in Mexico compared to the USA.
In contrast, Costa Rican Cenocoeliini were
11 times more speciose per unit area than
in Mexico, meanwhile the potential hosts
increased only about 1.5 times in species
richness per unit area for the same areas.

Cenocoeliini were most speciose at low
altitudes in Costa Rica (Table 3). Speci-
mens were not encountered above 1600 m
and Cenocoeliini were, significantly, more

abundant at low (<500 m) than at higher
(>500 m) altitudes.

DISCUSSION

The increase of species richness of Cen-
ocoeliini from Canada to Mexico was
equal to that of their common hosts and
exceeded that of their potential hosts in
Costa Rica. Hence, anomalous diversity
could not be shown for New World Cen-
ocoeliini. This is in contrast to the predic-
tion by resource fragmentation (Janzen
and Pond 1975; Janzen 1981) suggesting
that tropical host trees and their associat-
ed host beetles are too widely distributed
to be exploited by koinobiont parasitoids.
To illustrate this fragmentation, one ha of
tropical lowland rain forest in Costa Rica
contained 3 to 4 times more tree species
than one ha of the most diverse North
American forests (Hartshorn 1983; Whit-
tacker 1965). Also a 50 ha moderate di-
verse lowland rain forest in Panama con-
tained an equal number of tree species as
the whole of western North America,
north of Mexico (Condit et al. 1996; Little
1980). Janzen's (1981) caveat to resource
fragmentation was that tropical koino-
biont parasitoids had developed a broader
host range compared to temperate koino-
bionts, or that they had become very effi-
cient in looking for sparcely distributed
hosts. In the present study, this could not
be analyzed for neotropical Cenocoeliini,
partly due to the lack of host records.
North American Cenocoeliini, on average.

20

Journal of Hymenoptera Research

Table 3. Total species richness (S), number of Malaise trap-months {# tm), observed (N„i,J and expected
(N^.,j,) numbers of individuals of Cenocoeliini per altitude class in Costa Rica. Only Malaise traps which
operated more than three consecutive months were included in the analysis. It was tested if the observed and
expected numbers of Cenocoeliini were equal at low altitudes (<500 m) and at higher altitude classes com-
bined using chi-square test.

Altitude cl

ass

S

# tm

N

N,..„

xMdf = 1)

S500 m

48

546

219

155

78.27

500-1500 m

16

71

8

20

(P < 0.001)

1500-2500 m

5

138

7

15

39

79

>2500 m

0

72

0

20

Totals

57*

827

234

234

* Twelve species occurred at the two lowest altitude classes; one species occurred from 0-1600 m. Individuals
were not found above 1600 m altitude.

parasitize one to 4 different host species
(Saffer 1982). This "narrow" host ranges
would classify the North American Cen-
ocoeliini as specialists, however, we have
to be careful to generalize, because these
host records were not complete (Shaw
1994).

Results of the present study showed
that the increase in species richness per
unit area of Cenocoeliini from Mexico to
Costa Rica was 6 times larger than that of
their potential hosts (Table 2). This sug-
gests that Costa Rican Cenocoeliini para-
sitize a larger proportion of the Ceram-
bycidae and Scolytidae, than those in
North America. At the other hand, some
Costa Rican Cenocoeliini may have adapt-
ed to other beetle families of beetles with
a different biology. The only known host
record from Cenocoeliini in Costa Rica
was that of a species attacking seed-boring
beetle larvae of the family Curculionidae
(Saffer 1977). Long-term research on seed-
boring beetles in Costa Rican dry forest
(Janzen 1980), however, did not reveal ad-
ditional observations of Cenocoeliini on
seed-boring beetles (Janzen, pers. comm.).
Five percent of our surveyed specimens of
Cenocoeliini from Costa Rica were collect-
ed from treefalls by hand (Hespenheide,
unpublished data) and other micro-habi-
tats not were indicated on the collecting
labels. We expect, therefore, that wood-

and bark-boring insects are the most likely
host for Costa Rican Cenocoeliini.

In the present study, we found that few-
er species of Cenocoeliini occur at inter-
mediate than at low altitudes (Table 3).
This is comparable to the reduction in spe-
cies richness towards the equator (Table
1). But the decline in species richness of
Cenocoeliini with altitude in Costa Rica
was higher than expected. As shown in
the present study, Cenocoeliini were not
encountered at 2000 m in Costa Rica. At
latitudes in temperate North America
with a similar mean yearly temperature,
however, 5 to 8 species of Cenocoeliini oc-
curred (Quicke and Kruft 1995). In Costa
Rica, seasonal changes in temperature are
usually smaller than diurnally fluctuation
in temperature (Gauld 1995a). In temper-
ate regions this is often reversed. These
differences in temperature regimes may
affect species richness of New World Cen-
ocoeliini.

Individuals of Cenocoeliini were not en-
countered at altitudes higher than 1600 m,
even though a total of 150 Malaise trap-
months located at 10 different sites were
surveyed. This was an unexpected obser-
vation as potential hosts of Cenocoeliini
have been observed at high altitudes. Cer-
ambycidae are most abundant at low al-
titudes in Costa Rica but occur also at high
altitudes up to timberline (3200-3400 m;

Volume 7, Number 1, 1998

21

Lezama, pers. comm.). Scolytidae are
known to be regularly encountered at
high altitudes in Costa Rica (Wood et al.
1991). It is unlikely, however, that alter-
native hosts for Cenocoeliini, like larvae of
seed-boring beetles do occur at high alti-
tudes. Legume trees, of which the fruits
are among the most frequently attacked
by seed-boring beetles (Janzen 1980), are
scarce at intermediate and absent at high
altitudes (Holdridge et al. 1971). Also, Gas-
ton and Gauld (1993) noted that Pimplinae
(Icneumonidae) were more abundant at
high altitudes than at low altitudes. In
case species of Cenocoeliini would have
been present at high altitudes in Costa
Rica, they likely would have been collect-
ed in the Malaise traps. This suggests that
host presence does not explain absence of
Cenocoeliini at high altitudes in Costa
Rica.

In the present study, it could not be de-
termined if Cenocoeliini were scarce in
Costa Rica or that it is difficult to sample
them by using Malaise traps. On average,
one individual of Cenocoeliini was caught
per 3 to 4 Malaise trap-months. Trap effi-
ciency for Cenocoeliini was twice as high
at low altitude rain forests than at low
middle altitude rain forest or low altitude
dry forest. Also Cenocoeliini were most
frequently caught during the dry season
(Feb. -May; unpublished data). But even in
the optimal habitat type and season, Cen-
ocoeliini was never found to be abundant
suggesting that they occur in low popu-
lation densities.

Another remarkable result was that 77%
of the Mexican and 60% of the Costa Rican
species of Cenocoeliini were represented
by one or 2 individuals. Estimated species
richness of Mexican Cenocoeliini may
have been underestimated as sample cov-
erage by the Malaise traps in Mexico was
low compared to those in Costa Rica and
the USA and Canada combined.

Analyses of geographical distribution of
Costa Rican Cenocoeliini could not be jus-
tified due to low numbers of individuals

in Malaise traps. The observed 24 Ceno-
coeliini species in the Pacific lowland rain
forest around Golfo Dulce in Puntarenas
Province shared 9 species with the 25 Cen-
ocoeliini species in the Atlantic lowland
rain forest of La Selva in Heredia Prov-
ince. This species distribution of Cenocoe-
liini may suggest that many Costa Rican
Cenocoeliini have a restricted geographi-
cal distribution according to Rapoport's
Rule (Stevens 1989, 1992). It may also be
a sample artifact due to the low number
of individuals.

In the present study it was found that
half of the Cenocoeliini and especially
those larger than 5 mm were bright or-
ange and black colored, often with partly
or completely darkened wings. Saffer
(1982) described 2 similar bright colored
Cenocoeliini from southern tropical Mex-
ico, but no such colored species from tem-
perate North America. Bright colors are
common in tropical parasitoids (Quicke
1986a; Shaw 1995). Bright colors occur in
several other neotropical braconid sub-
families such as Agathidinae and Bracon-
inae and they are characteristic for larger
sized (>5 mm) diurnally active Braconi-
dae with long ovipositors, which likely
parasitize concealed hosts (Shaw 1995). In
general, bright colors are characteristic for
lowland insects where it occurs in up to
25% of insects and do not occur at high
altitudes in Costa Rica (Janzen 1973).
Quicke (1986b) noted that in general
bright colors have a warning function to-
wards visually-oriented predators (i.e.,
aposematic coloration). Bright colored
tropical parasitic wasps may mimic sting-
ing aculeates and some larger sized para-
sitoids are capable of stinging by them-
selves (Quicke 1986b). Other authors hy-
pothesized that parasitoids may mimic
unpalatable hosts such as Chrysomelidae
(Gauld, pers. comm.) or Symphyta (anon,
rev., pers. comm.). Gauld and Gaston
(1994), suggested that parasitoids with
bright colors may be unpalatable for pred-
ators after sequestering "nasty" tasting

22

Journal of Hymenoptera Research

secondary plant chemicals from their
hosts. If the latter is true and the "nasty"
host hypothesis has validity for Cenocoe-
liini, bright colored Cenocoeliini may at-
tack seed-eating or phloephageous beetle
larvae, rather than wood-living Scolytidae
or Cerambycidae. Quicke (1986a) found
homeochromatic assemblages for some
large sized Braconidae and their potential
hosts (i.e., Cerambycidae). Hespenheide
(1996) showed that color patterns of
Chrysomelidae may be substrate- related.
More research is needed to elucidate the
underlying defense mechanisms or other
meaning of bright colors in large sized
neotropical wood-boring braconid parasit-
oids.

The smaller sized Costa Rican Cenocoe-
liini from our survey were mostly less
conspicuous colored than the larger sized
ones. Many of these smaller sized Ceno-
coeliini may be ant-mimics, which is ex-
pected to occur frequently in neotropical
Braconidae (Shaw 1995). This color pat-
tern of presumably ant-mimics also oc-
curred among North American Cenocoe-
liini (Saffer 1982). To date, possible behav-
ioral and olfactorial cues involved in ant-
mimicry of Braconidae have not been doc-
umented.

In conclusion, the results of the present
study showed that the species diversity of
the Cenocoeliini increases towards the
equator. This is the normal pattern in lat-
itudinal species richness (Fisher 1960;
Pianka 1966; Stevens 1989; Wilson 1992).
Thus, anomalous diversity was not ob-
served for Cenocoeliini. The increase in
species richness per unit area of Cenocoe-
liini from temperate North America to the
Neotropics exceeded that of their potential
hosts. In Costa Rica, species and individ-
uals of Cenocoeliini were not found above
1600 m altitude, this in contrast to their
potential hosts which also occur at high
altitudes. This suggests that species rich-
ness in Cenocoeliini is not host limited.
The Cenocoeliini apparently is a tropical
lowland-centered group of which a limit-

ed number of species have adapted to
year-round cool conditions at lower mon-
tane rain forests and none to montane for-
ests. Some other species have adapted to
climatological conditions in temperate
regions and evolved overwintering mech-
anisms (Saffer 1982).

ACKNOWLEDGMENTS

We would like to thank Paul Hanson and Ian
Gauld for setting up the Malaise trap network in Cos-
ta Rica and for having the University of Wyoming
involved in identification of Braconidae. We would
like to thank Ian Gauld, David Kazmer, Dave Legg,
Jeff Lockwood, Mark Shaw and an anonymous re-
viewer for their critical comments on earlier versions
of the manuscript.

LITERATURE CITED

Achterberg, C, van. 1984. Essay on the phylogeny of
Braconidae (Hymenoptera: Ichneumonoidea)£ii-
tomologic TuMrift 105: 41-58.

Achterberg, C, van. 1993. Ilustrated key to the sub-
families of Braconidae (Hymenoptera: Ichneu-
monoidae). Zoologische Verhamkiiiigen 283: 1-189.

Achterberg, C, van. 1994. Generic revision of the sub-
family Cenocoeliinae Szepligeti (Hymenoptera:
Braconidae). Zoologische Verhandelingen 292: 1-52.

Askew, R. R. and M. R. Shaw (1986). Parasitoid com-
munities: their size, structure and development.
Pages 225-264, in J. Waage and D. Greathead
(eds) /»str( Parasitoids. 13th Symposium of the
Royal Entomological Society of London. Aca-
demic Press, London.

Brown, J. H. 1988. Species diversity. Pages 57-90, in
A. A. Myers and P. S. Gilles (eds) Analytical Ge-
ography. Chapmann and Hall. London, New
York. "

Colwell, R. K. and ]. A. Coddington. 1994. Estimation
terrestrial biodiversity through extrapolation.
Philosophical Transactions of the Royal Society Lon-
don (series B) 345: 101-118.

Condit, R., S. P. Hubbell, J. V. Lafrankie, R. Sukumar,
N. Manokaran, R. B. Foster and P. S. Ashton.
1996. Species-area and species-individual rela-
tionships for tropical trees: a comparison of three
50-ha plots, journal of Ecology 84: 549-562.

Fagen, R. M. and R. N. Goldmann. 1977. Behavioural
catalogue analysis methods. Animal Behaoioiir 25:
261-271.

Fischer, A. G. 1960. Latitudinal variation in organic
diversity. Evolution 14: 64-81.

Gaston, K. J. and I. D. Gauld. 1993. How many spe-
cies of pimplines (Hymenoptera: Ichneumoni-
dae) are there in Costa Rica? journal of Tropical
Ecology 9: 491^99.

Volume 7, Number 1, 1998

23

Gaston, K. J., I. D. Gauld and P. Hanson. 1996. The
size and composition of the hymenopteran fauna
of Costa Rica. ]ournal cf Bwgeogrnp!iy 23: 105-113.

Gauld, I. D. 1985. A preliminary survey of Ophioni-
nae (Hymenoptera: Ichneumonidae) of Brunei.
Brunei Museum Journal 6: 169-188.

Gauld, I. D. 1986. Latitudinal gradients in ichneu-
monid species-richness in Australia. Ecologicnl
Entomolog}/ 11: 155-161.

Gauld, I. D. 1987. Some factors affecting the compo-
sition of tropical ichneumonid faunas. Biological
loiirnal of the Linnaean Society 30: 299-312.

Gauld, I. D. 1991. The Ichneumonidae of Costa Rica,
1. Memoirs of the American Entomological Institute.
No. 47.

Gauld, I. D. 1995a. Introduction. Pages 1-19, in P. E.
Hanson and I. D. Gauld (eds) The Hymenoptera of
Costa Rica. The Natural History Museum, Lon-
don. Oxford Scientific Publications.

Gauld, I. D. 1995b. Ichneumonidae. Pages 390^31, in
P. E. Hanson and I. D. Gauld (eds) The Hyme-
noptera of Costa Rica. The Natural History Muse-
um, London. Oxford Scientific Publications.

Gauld, I. D., K. J. Gaston and D. H. Janzen. 1992.
Plant allelochemicals, tritrophic interactions and
the anomalous diversity of tropical parasitoids:
the "nasty" host hypothesis. Oikos 65: 353-357.

Gauld, I. D. and K. J. Gaston. 1994. The taste of ene-
my-free space: parasitoids and nasty hosts. Pages
279-299, in B. A. Hawkins and W. Sheehan (eds)
Parasitoid Community Ecology. Oxford Scientific
Publications.

Gauld, 1. D. and P. Hanson. In press. Distribution pat-
terns of cloud forest Hymenoptera in Costa Rica.
In N. Kadkarni and N. Weelright (eds.) Montev-
erde: natural history and conservation. Oxford Uni-
versity Press.

Hartshorn, G. S. 1983. Plants: Introduction. Pages_
118-183, in D. H. Janzen (ed) Costa Rican Natural
History. University of Chicago Press.

Hawkins, B. A. 1994. Patterns & Process in Host-Para-
sitoul Interactions. Cambridge University Press.

Hespenheide, H. A. 1979. Are there fewer parasitoids
in the tropics? The American Naturalist 113: 766-
769.

Hespenheide, H. A. 1996. The role of plants in struc-
turing communities of mimetic insects. Chapter
6. Pages 109-126, in A. C. Gibson (ed) Neotropical
biodiversity and conservation. Occasional Publica-
tion of the Mildred E. Mathias Botanical Garden
1. University of California, Los Angeles.

Holdridge, L. R., W. C. Grenke, W. H. Hatheway, T.
Liang and ]. A. Tosi Jr. 1971. Forest environments
in tropical life zones: a pilot study. Pergamon Press,
Oxford.

Holloway, J. D. 1986. Macrolepidopteran diversity in
the Indo-Australian tropics.: geographic, biotopic

and taxonomic variations. Biological Journal of the
Linnean Society 30: 325-341.

Janzen, D. H. 1973. Sweep samples of tropical foliage
insects: effects of seasons, vegetation types, ele-
vation, times of day, and insularity. Ecology 54:
687-708.

Janzen, D. H. 1980. Specificity of seed-attacking bee-
tles in Costa Rican deciduous forest. Journal of
Ecology 68: 929-952.

Janzen, D. H. 1981. The peak in North American Ich-
neumonid species richness lies between 38° and
42° N. Ecology 62: 532-537.

Janzen, D. H., M. Ataroff, M. Farinas, N. Rincon, A.
Soler, P. Soriano and M. Vera. 1976. Changes in
the arthropod community along an elevational
transect in the Venezuelan Andes. Biotropica 8:
193-203.

Janzen, D. H. and C. Pond. 1975. A comparison, by
sweeping, of the arthropod fauna of secondary
vegetation in Michigan, England and Costa Rica.
Transactions of the Royal Entomological Society Lon-
don {Series B) 127: 33-50.

Little, E. L. 1980. National Audubon Society field guide
to North American trees- western region. Alfred A.-
Knopf, New York.

MacArthur, R. H. and E. O. Wilson. 1967. The theory
of island biogeography . Princeton University Press,
New Jersey.

Magurran, A. E. 1988. Ecological diversity and its mea-
surement. Princeton University Press, New Jersey.

Monne, M. A. and E. F. Giesbert. 1994. Checklist of the
Cerambycidae of the western hemisphere. Wolfgar-
den Books, Burbank California, USA.

Morrison, G., M. Auerbach and E. D. McCoy. 1979.
Anomalous diversity of tropical parasitoids: a
general phenomenon? The American Naturalist
114: 303-307.

Noyes, J. S. 1989. The diversity of Hymenoptera in
the tropics with special reference to Parasitica in
Sulawesi. Ecological Entomology 14: 197-207.

Owen, D. F. and D. O. Chanter. 1970. Species diver-
sity and seasonal abundance in tropical Ichneu-
monidae. Oikos 21: 142-144.

Owen, D. F. and J. Owen. 1974. Species diversity in
temperate and tropical Ichneumonidae. Nature
249: 583-584.

Pianka, E. R. 1966. Latitudinal gradients in species
diversity: a review of concepts. The American Nat-
uralist 100: 33-46.

Quicke, D. L. J. 1986a. Preliminary notes on homeo-
chromatic associations within and between the
Afrotropical Braconinae (Hvmenoptera, Bracon-
idae) and Lamiinae (Coleoptera, Cerambycidae).
Entomologi.it's Monthly Magazine 122: 97-109.

QuickC, D. L. J. 1986b. Warning coloration and mim-
icry, in Betts (ed.) The Hymenopterist's Handbook,
2th edition The amateur Entomologist 7: 74-80.

Quicke, D. L. J. and R. A. Kruft. 1995. Latitudinal

24

Journal of Hymenoptera Research

gradients in North American braconid wasp spe-
cies richness and biology. Journal of Hi/iiieiioftern
Research 4: 194-203.

Rathcke, B. ]. and P. W. Price. 1976. Anomalous di-
versity of tropical ichneumonid parasitoids: a
predation hypothesis. The American Nntiirnlist
110: 889-902.

Saffer, B. 1977. A new species of Cenocoelius from Cos-
ta Rica. Proceedings of the Entomological Society of
Washington 79: 593-596.

Saffer, B. 1982. A systematic revision of the genus
Cenocoelius (Hymenoptera, Braconidae) in North
America including Mexico. Polskie Pismo Ento-
mologiczne 52: 73-167.

Shaw, M. R. 1994. Parasitoid host ranges. Pages 111-
144, in B. A. Hawkins and W. Sheehan (eds) Par-
asitoid Community Ecology. Oxford University
Press.

Shaw, M. R. and T. Huddleston. 1991. Classification
and biology of braconid wasps (Hymenoptera:
Braconidae). in W. R. Dolling and R. R. Askew
(eds) Handbook for identification of British Insects.
Vol. 7, Part iii. Royal Entomological Society of
London.

Shaw, S. R. 1995. Braconidae. Pages 431-463, in P. E.
Hanson and 1. D. Gauld (eds) The Hymenoptera of
Costa Rica. The Natural History Museum, Lon-
don. Oxford University Press.

Sheehan, W. and B. A. Hawkins. 1991. Attack as an
indicator of host range in metopiine and pimp-
line Ichneumonidae (Hymenoptera). Ecological
Entomology 16: 129-131.

Sokal, R. R. and F. J. Rohlf (1981). Biometry. 2nd edi-
tion. W.H. Freeman and Company. San Francis-
co.

Stevens, G. C. 1989. The latitudinal gradient in geo-
graphical range: how so many species coexist in
the tropics. The American Naturalist 133: 240-256.

Stevens, G. C. 1992. The elevational gradient in alti-
tudinal range: an extension of Rapoport's's lati-
tude rule to altitude. The American Naturalist 140:
893-911.

Wharton, R. A. 1993. Bionomics of Braconidae. An-
nual Reviezv of Entomology 38: 121-143.

Whittacker, R. H. 1965. Dominance and diversity in
land plant communities. Science 147: 250-260.

Wilson, E. O. 1992. The diversity of life. Harvard Uni-
versity Press, Cambridge, Massachusetts.

Wolda, H. 1988. Insect seasonality: Why? Annual re-
view of Ecology and Systematics 19: 1-18.

Wolda, H. 1989. Seasonal cues in tropical organisms.
Rainfall? Not necessarily? Oecologia 80: 437^142.

Wolda, H. and M. Wong. 1988. Tropical insect diver-
sity and seasonality. Sweep-samples vs. light-
traps. Proclaniaties Koninklijke Nederlandse Akade-
mie voor Wetenschappen 91: 203-216.

Wood, S. L. 1982. The bark and ambrosia beetles of
North and Central America (Coleoptera: Scoly-
tidae) a taxonomic monograph. The Great Basin
Naturalist Memoirs No.6. Brigham Young Univer-
sity, Provo, Utah.

Wood, S. L., G. C. Stevens and H. J. Lezama. 1991.
Los Scolytidae de Costa Rica: clave de generos y
de subfamilia Hylesinae (Coleoptera). Rei'ista
Biologica Tropica 39: 125-148.

J. HYM. RES.
Vol. 7(1), 1998, pp. 25-37

Sperm Development in the Imaginal Testes of Aleiodes coxalis
(Hymenoptera: Braconidae: Rogadinae)

Terence M. Newman and Donald L. J. Quicke'

Department of Biology, Imperial College of Science, Technology and Medicine, Silwood Park,

Ascot, Berks SL5 7PY, UK

Abstract — Spermatogenesis and spermiogenesis in the imaginal testes of the long-lived braconid
wasp, Aleiodes coxalis (Spinola) has been investigated. The cyclostome group of braconid subfam-
ilies, to which Aleiodes belongs, are considered to have unspecialised sperm, but in Aleiodes several
aspects of spermatogenesis and morphology differ from that reported in other Hymenoptera,
suggesting that reinterpretation of previously published studies may be necessary. In particular,
the centriolar adjunct is found to lie between the nucleus and one of the pair of mitochondrial
derivatives, resulting in the mitochondrial derivatives being offset longitudinally, giving the im-
pression that the mitochondrial derivatives are of different lengths. A projection extends from the
centriolar adjunct to the one mitochondrial derivative which abuts the nucleus. The shape change
undergone by the nucleus during spermatogenesis is associated with an uneven distribution of
peri-nuclear microtubules (microtubular manchette). These are not found over two extra-nuclear
electron-dense regions, ('lateral plates') which appear to add increased rigidity to the nucleus at
these points.

Very little is known about the sperm of
Hymenoptera despite the fact that this is
one of the largest and most diverse of in-
sect orders (Jamieson 1987), and descrip-
tions of spermatogenesis are limited to a
handful of taxa, mostly bees and ants
(Hoage & Kessel 1968, Hogge & King
1975, Cruz-Landim & Beig 1980). A recent
survey of mature sperm morphology and
ultrastructure across the order has re-
vealed considerable variation between
higher taxa that may be important for un-
derstanding phylogenetic relationships
within the order (Quicke et al. 1992), some
of the most extreme modifications occur-
ring in the parasitic wasp family, Bracon-
idae. In order to interpret the structures
and modifications found in some braconid
subfamilies it is first necessary to describe
the 'normal' situation in that family.

The Braconidae is a large family of par-
asitic wasps comprising some 40 subfam-

' To whom correspondence should be addressed.

ilies that can be broadly divided into two
groups, the cyclostomes and the non-cy-
clostomes, which differ in morphology of
the mouth and in their biology (Shaw &
Huddleston 1991). In an initial survey
(Quicke et al. 1992), it was found that in
members of the cyclostome group of brac-
onid subfamilies (e.g. Braconinae, Doryc-
tinae and Rogadinae, the latter including
Aleiodes) the sperm have a normal appear-
ance, being between 80 and 170 |xm long
with the head (nucleus + acrosome) com-
prising 10 to 20% of the sperm's total
length. However, nearly all non-cyclo-
stomes have apomorphic sperm with total
length between 10 and 20 |xm, of which
the head constitutes nearly 50%. Further,
whilst the axonemes of the cyclostome
braconid sperm have a normal 9-1-9-1-2 ar-
rangement of microtubules, as found in
most insects, those of the short apomor-
phic non-cyclostome sperm frequently
have only one or no central microtubules.
These features appear to be highly phy-

26

Journal of Hymenoptera Research

logenetically informative and, in order
better to understand their ontogenies, we
have investigated spermatogenesis in a
range of braconids. Here we report on that
leading to the apparently unmodified
sperm of the rogadine, Aleiodes coxalis
(Spinola), which will form the basis for fu-
ture comparisons. Despite the normal ap-
pearance and ultrastructure of the mature
sperm in Aleiodes, spermatogenesis in this
taxon exhibited several structures that do
not appear to have been reported in other
insects.

It has long been appreciated that the
timing of spermiogenesis and spermato-
genesis in insect testes is a reflection of the
adult life span which may in turn be as-
sociated with the occurrence of sib-mating
(Phillips 1970). Thus imaginal testes of
species with short lived adults, such as
mayflies (Needham et al. 1935) and caddis
flies (Ross 1944), or those in which males
almost invariably mate with their nearby
sisters upon emergence such as many par-
asitic wasps, typically contain spermatids
and spermatozoa, but no gonial and mei-
otic divisions, which would be completed
in the pupal or nymphal stages. In con-
trast, meiotic and even gonial divisions of-
ten continue well into the imaginal stages
in testes of long lived species (e.g. many
beetles and dragonflies) which usually
search for and mate with several unrelat-
ed females. Most rogadine braconid
wasps, such as the Aleiodes species inves-
tigated here, are solitary parasitoids of
Lepidoptera larvae and belong to the sec-
ond category. Males fed on honey water
will often live a month and sometimes
considerably longer, and will mate readily
throughout much of their adult life (M. R.
Shaw pers. comm.). Indeed, some species
are reluctant to mate immediately after
eclosion and only start to mate once they
have fed.

There are no papers on the ultrastruc-
ture of spermatogenesis in any member of
the Ichneumonoidea, and the only para-
sitic Hymenopteran studied to date is the

pteromalid chalcidoid Nasoiiia vitripennis
(Walker) (Hogge & King 1975).

MATERIALS AND METHODS

Adult Aleiodes coxalis (Spinola) were col-
lected at Silwood Park (Imperial College)
in 1995. Testes were dissected under 2%
glutaraldehyde in phosphate buffered sa-
line (pH 7.2), and fixed for two hours. Tis-
sue was transferred to 2% osmium tetrox-
ide in cacodylate buffer (pH 7.2) for 2 hr.
After another buffer wash, tissue pieces
were dehydrated to 50% ethanol and then
further fixed with saturated uranyl acetate
in 50% ethanol prior to complete dehydra-
tion, embedding in Epon resin and poly-
merisation overnight. Large silver sections
were picked-up onto high resolution
grids, stained with uranyl acetate and lead
citrate and examined using a Phillips
EM400 electron microscope.

RESULTS

Adult testes of Aleiodes coxalis contain
follicles (seminal tubules) with multiple
cysts present at various levels of maturity.
Although neighbouring cysts can exhibit
different stages of development, generally
the more posterior the position along the
testes the greater the proportion of later
stages. Although most cysts contain sper-
matids, it is also possible to find in the
adult indications of earlier stages of de-
velopment. Primary spermatocytes still
appear to be present some without any
sign of a normal spindle and with a con-
centric arrangement of the endoplasmic
reticulum around the chromosomes (Fig.
1 long arrow). Centriolar pairs are present
in these cells {arrowheads), as are large
numbers of small mitochondria. This ap-
pearance is considered indicative of an
abortive first meiotic division; the result of
incomplete centriolar polarisation. Centri-
oles are then extruded after meiotic mul-
tiplication by cytoplasmic blebbing (Fig.
2), where centrioles, attached to microtu-
bules (Fig. 2a), are found incorporated
into rounded portions of the plasma mem-

Volume 7, Number 1, 1998

27

brane. This blebbing is also seen in sec-
ondary spermatocytes (which also exhibit
loss of plasma membrane in larger
whorls), although not all blebs exhibit cen-
trioles (Fig. 2b arrow). Large numbers of
polyribosomes are also present at this
stage.

In the secondary spermatocytes it is dif-
ficult to find evidence of synaptonemal-
like structures. At this stage the cells are
considered to form a syncytium because
of incomplete cytogenesis during the ear-
lier mitotic division. The intercellular
bridges (Fig. 3) linking the spermatocytes
are difficult to identify as the cells can be
very closely apposed. It is only during lat-
er spermiogenesis, with cell elongation,
that the connections become apparent
(Fig. 3a) with evidence of organelle conti-
nuity and possibly even intercellular
movement of vesicles and granules (Fig.
3b). This has been suggested as being a
method of transferring organelles from
large to small spermatids where there is
unequal division after the second meiotic
division. No noticeable size difference was
seen in Aleiodes.

Nebenkern formation occurs concomi-
tantly with the above processes (Fig. 4).
Mitochondria begin to accumulate in one
area of the cell (Fig. 4a) where they fuse
to form the beginnings of the nebenkern
(Fig. 4b), a highly convoluted membra-
nous organelle that is the precursor of the
two mitochondrial derivatives. The devel-
opment of the nebenkern into two labyrin-
thian networks has been linked with the
appearance of microtubules in the cyto-
plasm. However, in Aleiodes microtubules
appear to be a consistent component of the
cytoplasm from earlier stages.

Flagellum growth (Fig. 5) is evident pri-
or to nuclear condensation, commencing
with the appearance of a basal body (Fig.
5a arrowhead) in a small depression of the
nucleus (Fig. 5a arrow). The nuclear enve-
lope thickens at this area (Fig 5b) and the
doublet microtubules of the flagellum
grow from the triplet microtubules of the

basal body (Fig 5b arroio). The flagella ini-
tially do not have accessory tubules (Fig.
5c). These appear to develop from sub-fi-
bre b of each doublet as previously re-
ported. The flagella /mitochondria axis is
developed before this takes place (Fig. 5d
arroiv). Although spermiogenesis is ad-
vanced at this stage, separation following
the last meiotic division in many cases is
not complete and the nucleus still appears
spherical. Nuclear condensation begins
with the appearance of polarity in the nu-
clear membrane in the region closely ap-
posed to the spermatid cell membrane
(Figs 5d, 6 arrowheads) where the sperma-
tid is attached to the wall of the cyst (a
single epithelial layer with supportive and
secretory functions). Microtubules appear
around the nucleus but these are not even-
ly located; an arrangement which proba-
bly assists the complicated shape change
that the nucleus undergoes.

The acrosome is detectable in early
spermatid stages as an accumulation of a
large number of small particles termed the
proacrosomal granule (Fig. 7). During
spermiogenesis the proacrosomal granule
becomes located between the plasma
membrane and the nucleus at the region
of nuclear polarity (Fig. 8a). The acrosome
is formed by a flattening of this structure
and an indenting on the side apposed to
the nucleus (Fig. 8b). A small protrusion
is found anteriorly to the acrosome in the
early spermatid (Fig. 8b arroiv). This may
represent an apical expansion of extra-ac-
rosomal material, similar to that found in
Locusta migratoria L. (Szollosi 1974). How-
ever, the structure in Aleiodes appears to
contain electron dense material. Whether
this could be a separate structure, or rep-
resents a later stage of fusion of Golgi de-
rived products, as found in the proacro-
somal granule, is not clear. The situation
is further complicated by the impression
that the is distinct coated membrane
which surrounds the acrosome and ex-
tends posteriorly over the anterior portion
of the nucleus during development, at

28

Journal of Hymenoptera Research

3b

=*■•*?;^■

;4b

J, ;■

Figs. 1-4. 1, Primary spermatocyte from the testes of nn adult male Aleiodes, with anomalous metaphase
plate formation, characterised by the absence of a normal spindle and a concentric arrangement of the en-
doplasmic reticulum around the chromosomes (armii'). Large numbers of small mitochondria and a centriolar
pair (anvu'liciidi), are also present. Scale bar = 0.5 |j.m. 2, Membrane blebbing during spermiogenesis: (a) loss
of centrioles (attached to microtubules) through membrane blebbing; (b) membrane blebbing (iinoiv) without
loss of centrioles. Scale bar a = 0.6 |i,m; b = 0.85 jjim. 3, Syncytium formed by incomplete cvtogenesis during
mitotic division: (a) Intercellular bridges (imou'lwntl) link spermatocytes; (b) linkage provides a method for
organelle movement between spermatocytes. Scale bar = 0.75 jjim. 4, Nebenkern formation in the spermato-

Volume 7, Number 1, 1998

29

least initially does not appear to be contin-
uous with this granular tip. As the acro-
some elongates an acrosomal rod (or per-
foratorium) develops, and extends from a
depression in the nucleus and into the su-
bacrosomal space formed during folding
(Fig. 8c, arrozi'). The acrosomal rod be-
comes hollow as it develops (Fig. 8d), but
does not occupy all of the large sub-acro-
somal space (Fig. Be). At this stage the
complete structure is referred to as the ac-
rosomal complex.

With the formation of the acrosomal
complex and cell elongation, a distinct
centriolar-adjunct appears in the sper-
matocyte (Fig. 9). It develops at the pos-
terior nuclear pole adjacent to the forming
basal body (Fig 9a). Unlike the situation in
some other insect species (Gatenby & Tah-
misian 1959, Breland et al. 1966) it does
not appear to be forming around the cen-
triole that is perpendicular to the axis of
the flagellum. Instead, the basal body is
located between the centriolar adjunct and
the 'perpendicular' centriole; the ultimate
fate of this second centriole is not clear. In
transverse section, the axoneme is associ-
ated with one mitochondrial derivative
and the sectioned centriolar adjunct
(which in the past has been interpreted as
a darkened mitochondrial derivative). In
longitudinal section the centriolar adjunct
can be seen to lie between the posterior
pole of the nucleus and one of the mito-
chondrial derivatives (Fig. 9c), and is
slightly more than 2|xm long. The mito-
chondrial derivatives are thus offset lon-
gitudinally, with the other member of the
pair abutting the nucleus. This may result
in the appearance of the adult mitochon-
drial derivatives, where one member of
the pair often seems to extend further pos-
teriorly, possibly giving the erroneous im-

pression that in fact the mitochondrial de-
rivatives are of different lengths. This is a
different arrangement from that previous-
ly reported for other related species. Fur-
thermore, anteriorly, it can be seen that for
part of the length of the centriolar adjunct,
where the centriolar adjunct and one of
mitochondrial derivatives lie side by side,
there is an extension of the centriolar ad-
junct which surrounds the mitochondrial
derivative on the side facing the centriolar
adjunct (Fig. 9d). Interestingly in a rare ter-
atological sperm-tail with two axonemes
the centriolar adjunct was found to extend
to enclose two mitochondrial derivatives.
This contact is most evident in the region
of the basal body and it is not clear if it
actually extends into the flagellum proper
as defined by the possession of both a ring
of doublet microtubules and a central pair
of microtubules.

The change in shape of the nucleus (Fig.
10) that occurs with the appearance of
peri-nuclear microtubules is also marked
by the formation of two extra-nuclear elec-
tron-dense regions (Fig. 10a, see also Fig.
6). Peri-nuclear microtubules are unevenly
distributed and are not found over the ex-
tra-nuclear electron-dense regions, which
we have called lateral plates, nor do they
occur in the area of the nucleus that will
be convex during the elongation process.
The lateral plates appear to be the locus
for shape changes that occur during the
elongation of the nucleus. Condensation
of the chromatin into coiled fibrillar
threads then follows (Fig. 10b). In trans-
verse section, as the nucleus elongates, the
threads are found associated with that por-
tion of the nuclear membrane that is ad-
jacent to the region lacking microtubules
(Fig. 10c). The side of the nuclear mem-
brane surrounded by the full complement

cyte: (a) Large numbers of small mitochondria accumulate in one area of the spermatocvte; (b) mitochondrial
fusion occurs to form the labyrinthine network of the nebenkern, which will itself then give the two mito-
chondrial derivatives of the mature sperm. Scale bar a = 1.4 (j.m; b = 1.15 \xm.

30

n

Journal of Hymenoptera Research

5b

4^

w

ni

'"^if'/

'':'t^

f

■■:

5c

...^/ r

t

Figs. 5-7. 5, Flagellum formation during spermiogenesis: (a) tlu' Iviscil hcidv (iiirowhciht) forms from the lateral
centriole by insertion into a depression of the nucleus {nnow); (b) doublet microtubules {iinou') grow from the
triplet microtubules of the basal body; (c) accessory tubules arc absent at this stage, although a central pair
of microtubules form in the developing axoneme post-basal body; (d) the flagella/ mitochondria axis is de-
veloped (arrow) before the change in shape of the nucleus is found, although polarity in the nuclear membrane
can be found, n = nucleus. Scale bar a, b = 0.7 jjim; c = 0.8 jj.m; d = 3 (xm. 6, Polarity of the nucleus develops
prior to shape change with increased electron density of the nuclear membrane including the appearance of
two extra-nuclear densities (lateral plates). Microtubules (nrroivlicads) appear around the nucleus. Scale bar =
1 \xm. 7, Proacrosomal granule formed from small Golgi derived vesicles. Scale bar = 1 jim.

Volume 7, Number 1, 1998

8a

:##^:; 8 b

/

31

<w

n

■.•A .Villi. aiWAi

hM0

'•'='

-■■' -

^vvjijigiitti

^,^ ^-;^-: ,

'«ii

'J.

8d

Be

'If:

^#^

#^

I ij;. h. 1 Liiaicition of the acrosomal complex during spermiogoncbis: (a) tlic acrosuni>_ l.t_t_>.,iiii> U.>.atod be-
tween the plasma membrane and the nucleus (urrozc); (b) the acrosome indents to enfold the nucleus, and a
small electron density is found anteriorly; (c) an acrosomal rod ('perforatorium' arrow) occupies the subacro-
somal space formed during folding; (d) the perforatorium (arrpic) is seen in transverse section to be hollow;
(e) the subacrosomal space itself can be seen to be large and is not filled by the perforatorium for its entire
length, n, nucleus. Scale bar a, c, d = 0.5 \j.m; b, e = 0.65 (j.m.

32

Journal of Hymenoptera Research

9d -^ "^

*r\ >.'■-■

«v;

V ' "yV^ ':M^ i?^wiK*r^

p:^^^^

9c 1

K*

^^

^

f

' 1

^

B^

:

^^^

* „ -tf >*'!

1^

/|,

r

^^H_L *

^■

3

■ ^ 1

r

■ '*■»■'««

■^

^-.

Fig. 9. Centriolar adjunct formation during spermiogenesis: (a) arraii^unuiit nl iLiitriolar adjunct {arnnr, 'a'),
basal body (anmu, 'b') and perpendicular centriole (iirroic, 'c'); (b) in transverse section the centriolar adjunct
(nrrou') and a single mitochondrial derivative {arroiclwad) lie in parallel with the axoneme; (c) in longitudinal
section the centriolar adjunct (c) abuts the nucleus anteriorly at the nuclear plate (largo nrrcw) and a mito-
chondrial derivative (curved arrozv) posteriorly. Microtubules (arrcwf) are evident lining the nucleus (n) and
the developing axoneme; (d) at the level of the basal body (arrows) the centriolar adjunct (iinoic/icHife) extends
to contact and enclose the single mitochondrial derivative. Posteriorly to the centriolar adjunct two mitochon-
drial derivatives are found {arrou'). Note also the absence of the central pair of microtubules in the basal
bodies, n, nucleus. Scale bar a = 0.5 jjim; b = 0.3 ixm; c = 0.6 |j.m; d = 0.9 |j.m.

VoLUMK 7, Number 1, 1998

33

10f

10d

r---

10e. *.

^I^B

i

.. ' ■. ■--•

%

- ,,: -;

.^^^%l

^s^fc*^

•-**' ^

Ktm^J^i''* »i wi^^ :'m>^frm^'-

Fig. 10. change in shape of nucleus during spermiogenesis: (a) extra-nuclear electon-dense region (lateral
plate', arnnc) are produced upon nuclear shape-change; (b) condensation of the chromatin into coiled fibrillar
threads (arwxv); (c) the threads divide and become associated with that portion of the nuclear membrane on
either side of the region lacking microtubules (nnmi's); (d) the side of the nucleus with the full complement
of microtubules {lar;^c arrows) flattens, and then infolds with the lateral plates (arrowheads) providing a locus
for shape change; (e) the chromatin becomes a layer on the outer surface of a horse shoe (arrowheads) that
infolds to surround elements of the endoplasmic reticulum; (f ) as the spermatocyte elongates the groove begins
to disappear from the anterior and posterior ends; (g) the dense nuclear material then begins to redistribute
more evenly around the nuclear membrane as the width decreases. Scale bar a = 40 |xm; b = 1 jxm; c, d, e
= 0.6 |j.m; f = 1.2 |j.m; g = 1.7 |j.m.

34

Journal of Hymenoptera Research

of microtubules then flattens (Fig. lOd ar-
row), with the chromatin becoming a layer
on the outer surface of the developing
horse-shoe shape; as viewed in section
(Fig. lOd arrowhead). The lateral plates be-
come the linear portions of the outer sur-
face of the arms of the 'horse-shoe' (Fig.
lOe arrowheads); at the most extreme, the
lateral arms (in transverse section) almost
come into contact with one another. As the
nucleus infolds, so elements of the endo-
plasmic reticulum, which surround the
microtubules, become largely enclosed by
the nucleus (Fig. lOe). It is not clear how
this highly folded nucleus reaches the cy-
lindrical shape of the adult sperm. As the
sperm elongates it appears that the in-
verted side is flattened with the result that
the grove begins to disappear from the an-
terior and posterior ends (Fig. lOf). The
dense nuclear material then begins to re-
distribute more evenly around the nuclear
membrane as the width decreases (Fig.
lOg); however, intermediate stages are dif-
ficult to identify.

The axoneme itself develops into the
9 + 9 + 2 arrangement (Fig. 11a); 9 outer sin-
gle accessory tubules, 9 doublets and 2
central single microtubules. Intratubular
material is abundant (with radial spokes —
Afzelius rays) and indications of the inner
and outer dynein arms. Two deltoid bod-
ies, (also referred to as triangular rods;
Lensky et al. 1979) develop, but their exact
derivation is not clear, although a mem-
brane origin is apparent and a close as-
sociation with the mitochondrial deriva-
tives is probable. This is further supported
by the observation that only one deltoid
body is found at the level of the centriolar
where there is only a single mitochondrial
derivative. At the level of the two deltoid
bodies is a single central rod, as previous-
ly reported in ant spermatozoa (Wheeler
et al. 1990). This arrangement is main-
tained in mature sperm (Fig. lib). By com-
parison with that of the spermatocyte, the
nucleus of the mature sperm is homoge-
neous (Fig. lie). The acrosome develops a

distinctive glycocalyx (Fig. lie arroivheads)
which extends over the anterior portion of
the nucleus. The two mitochondrial deriv-
atives show the offset arrangement that
may be the result of the centriolar adjunct
overlaying one of them anteriorly (Fig. lie
arrows). In addition an end piece with no
mitochondrial derivatives is found (Fig.
lid arrow).

DISCUSSION

In the testes of imaginal male Aleiodes
we have found structures indicative of
early stages of sperm development. In
particular, evidence for an abortive first
meiotic division of the primary spermato-
cyte, a major feature of hymenopterous in-
sects in which males are haploid, has been
found. This appears to involve anomalous
metaphase plate formation, probably
caused by the absence of a normal spindle.
This has been previously described in the
drone of the bee Scaptotrigona postica by
Cruz-Landim & Beig (1980), where it is
characterised by the presence of a concen-
tric arrangement of the endoplasmic retic-
ulum around the chromosomes and the
absence of a normal spindle. In Aleiodes,
as in S. postica, this is connected to an
anomalous movement of spermatocyte
centrioles, which, instead of assuming po-
lar locations, migrate to the cell surface
and are lost by cytoplasmic blebbing
(Hoage & Kessel 1968). The presence of
such early stages of development in the
adult supports our supposition that gonial
development in this species could to be re-
lated to the life style of the adult. The rel-
atively long adult life time of the male
would seem to obviate the need for syn-
chronous maturation of all the individual
sperm cysts.

The manchette of microtubules that as-
sembles around the developing spermatid
nucleus appear to be important for nucle-
ar compression and elongation (Baccetti
1972). The nuclear shape change found
during development in Aleiodes, and the
asymmetric distribution of microtubules.

Volume 7, Number 1, 1998

35

•11a 5i^>•^^•^ ' '■■liiiS^Sife^ %^^H^' lib/- |

■*f

r^

'h

t*

Fig. 11. Axoneme structure and the mature sperm:(a) The arrangement of 9 outer, singlet accessory micro-
tubules (at), 9 doublet (d) and two single central microtubules (t) is evident in the later stages of spcrmiogen-
esis, with intratubular material (i) and prominent radial spikes (s); (b) two deltoid bodies {arroivs) develop in
association with the mitochondrial derivative except at the level of the centriolar adjunct where only one is
found; (c) the mature sperm illustrates the uneven length of the mitochondrial derivatives (arrow) and the
glycocalyx that develops around the mature acrosome {arrowhmds); (d) the tail piece at the posterior of the
sperm (arrow) contains only the 9+2 arrangement of doublet and single microtubules. Scale bar a = 100 |a.m;
b = 0.25 (xm; c = 0.75 (xm; d = 0.4 jjLm; p = perforatorium.

have not previously been described for
parasitic wasps. A gutter-shaped devel-
opmental stage of the nucleus occurs in
some bugs (Lee & Lee 1992) but nuclear

elongation is not accompanied by concav-
ity in many other insects (e.g. SzoUosi
1975, Friedlander 1993, Wolf & Joshi
1995). In some caddis flies, in which the

36

Journal of Hymenoptera Research

spermatid nucleus transiently assumes a
sickle shape, the microtubular manchette
surrounding the nucleus is interrupted
(Wolf & Klein 1995), as it is in the drag-
onfly, Aeshna grandis L., in which the elon-
gating nucleus becomes locally com-
pressed (Kessel 1966). The structures we
have termed lateral plates, which occur
between groups of microtubules of the mi-
crotubular manchette, do not appear to
have been described previously in any in-
sect.

The present study of spermiogenesis in
Aleiodes has implications for published
studies of the centriolar adjunct. The pres-
ence of a centriolar adjunct overlying only
one of the mitochondrial derivatives has
not to our knowledge previously been re-
ported. In particular the connection from
the centriolar adjunct to the mitochondrial
derivative that abuts the nucleus is novel.
This arrangement may be responsible for
observations in related species suggesting
that anteriorly there may be a particularly
electron dense mitochondrial derivative
(Quicke et al. 1992) or a region of overlap
between the axoneme, nucleus and mito-
chondrial derivative (Chauvin et al.
1987) — i.e. some studies may have misin-
terpreted the centriolar adjunct as a par-
ticularly electron dense mitochondrial de-
rivative. Further, the longitudinal offsetting
of the mitochondrial derivatives caused by
the interposition of the centriolar adjunct
between one of them and the nucleus
could be responsible for the appearance of
some sections through the posterior re-
gion of the sperm, just before the tail
piece, in which only one mitochondrial
derivative is found. The presence of a sin-
gle mitochondrial derivative in sections
through this region in other species
prompted previous authors to conclude
that mitochondrial derivatives are of dif-
ferent lengths, which has even been con-
sidered to be a characteristic, not just of
the Hymenoptera, but of all holometabo-
lous insect orders (Wheeler et al. 1990).
However this result could be explained

simply by the arrangement of the centrio-
lar adjunct. In the light of this finding in
Aleiodes, it would be interesting to re-ex-
amine these previously reported groups
for the presence of a similarly located cen-
triolar adjunct. A large centriolar adjunct
has been reported in ant spermatozoa
(Wheeler et al. 1990), and was thought to
distinguish this group from other Hyme-
noptera, such as bees, which appear to
lack a homologous structure (Lensky et al.
1979, Cruz-Landim & Beig 1980). The ar-
rangement of the centriolar adjunct in the
ant spermatozoa, at least in longitudinal
section, is similar to that of Aleiodes. How-
ever, in these the centriolar adjunct can be
seen to overlie both mitochondrial deriv-
atives.

The connection to the mitochondrial de-
rivative is particularly interesting as it oc-
curs at the level of the basal body where
the two central microtubules are absent.
The origin and exact function of the cen-
tral pair is not known. A structural role
for the central pair could imply a similar
role for the centriolar adjunct and the lat-
ter could therefore be compensating for
the absence of the central pair at the re-
gion of the basal body. However, some
studies have provided evidence that the
centriolar adjunct consists partly of RNA
(Baccetti et al. 1969) and how this would
be related to a structural role is uncertain.

The exact relationship between centri-
oles, the basal body, the centriolar adjunct
and the developing axoneme in Aleiodes is
not clear. In another parasitic wasp, the
pteromalid Nasouia vitripeunis (Walker),
the distal centriole is reported to lengthen
to form the basal body of the flagellum,
and then, with the proximal centriole,
form the ring centriole (Hogge & King
1975). This then associates with a spherical
reticulate dense body, which forms adja-
cent to the nuclear membrane opposite a
nuclear pore at the secondary spermato-
cyte stage. As maturation proceeds the
proximal centriole is reported to become
surrounded by granular material and

Volume 7, Numbhr 1, 1998

37

break down, with loss of material from the
spherical, reticulate dense-body. The gran-
ular material, with presumably the re-
mains of the proximal centriole is then
thought to contribute to the centriolar ad-
junct. However, in Aleiodes such a straight-
forward relationship is not clear; in fact it
appears as if the centriolar adjunct may
develop prior to loss of the proximal cen-
triole. Thus this may not contribute to the
formation of the centriolar adjunct at all.
In discussions of the centriolar adjunct in
ant spermatozoa, Wheeler et al. (1990)
suggest caution when applying this term,
as the developmental homology to such
structures in other species, such as mam-
mals, has not been proven.

ACKNOWLEDGEMENTS

The authors gratefully acknowledge the invaluable
assistance of Mr Ian Morris (Electron Microscopy
Unit, Biology Department, Imperial College). This
work was supported by the NERC Initiative in Tax-
onomy.

LITERATURE CITED

Baccetti, B. 1972. Insect sperm cells. Advances in Insect
P/ji/sio/iW/ 9: 315-397.

Baccetti, O. P., R. Dallai, and F. Rosati. 1969. The sper-
matozoon of Arthropoda III. The lowest holomet-
abolic insects. Journal of Microscop}/ 8: 233-248.

Breland, O. P., C. D. Eddleman and J. J. Biesele. 1968.
Studies of Insect Spermatozoa. Entomological
News 79: 197-216.

Chauvin, G., M. El Agoze, C. Hamon and J. Huig-
nard. 1987. Ultrastructure des spermatozoides di-
ploides de Dimlroinus inilchellus Wesmael (Hy-
menoptera: Ichneumonidae). International ]onrnal
of Insect Morphology and Embryology 17: 358-366.

Cruz-Landim, C. and D. Beig. 1980. An electron mi-
croscopical study of spermatogenesis in the
drone of Scaptotrigona postica (Hymenoptera, Ap-
idae). International Journal of Invertebrate Repro-
duction 2: 272-283.

Friedlander, M. 1993. Phylogenetic position of rhy-
acophiloid caddisflies (Insecta, Trichoptera): a
spermatological analysis of Rliyacophilidae and
Glossosomatidae. Zoologica Scripta 22: 299-304.

Gatenby, J. B. and T. N. Tahmisian. 1959. Centriolar
adjunct, centrioles, mitochondria and ergosto-
plasm in orthopteran spermatogenesis. An elec-
tron microscopical study. La Cellule 60: 103-134.

Hoage, T. R. and R. G. Kessel. 1968. An electron mi-

croscopical study of the process of differentiation
during spermatogenesis in the drone honey bee
with special reference to replication and elimi-
nation. Journal of Ultrastructure Research 24: 6-32.
Hogge, M. A. F. and P. E. King. 1975. The ultrastruc-
ture of spermatogenesis in Nasonui vitripennis
(Walker) (Hymenoptera: Pteromalidae). journal of
Suhmicroscopical Ci/tolog}/ 7: 81-996.
Jamieson, B. G. M. 1987. The Ultrastructure and Phy-
logeny of Insect Spermatozoa. Cambridge Univer-
sity Press, Cambridge, 320pp.
Kessel, R. G. 1966. The association between microtu-
bules and nuclei during spermiogenesis in the
dragonflv. Journal of Ultrastructure Research 16:
293-304.
Lee, Y. H. And C. E. Lee. 1992. Ultrastructure of sper-
matozoa and spermatogenesis in Nepomorpha
(Insecta: Heteroptera) with special reference to
phylogeny. Zoological Science 9: 971-981.
Lensky, Y., E.Ben-David and H. Schindler. 1979. Ul-
trastructure of the spermatozoan of the mature
drone honeybee. Journal of Apiculture Research 18:
264-271.
Needham, J. G., ]. R. Traver, and Y. C. Hosu. 1935.
The Biology of Mat/flies. Comstock Publishing As-
sociates, Ithaca, New York.
Phillips, D. M. 1970. Insect sperm: their structure and
morphogenesis. Journal of Cell Biology 44: 243-
277.
Quicke, D. L. J., S. N. Ingram, H. S. Baillie and P. V.
Gaitens. 1992. Sperm structure and ultrastructure
in the Hymenoptera (Insecta). Zoologica Scripta
21: 381-402.
Ross, H. H. 1944. The caddis flies or Trichoptera of
Illinois. Illinois Natural History Survey Bulletin 23:
1-326.
Shaw, M. R. and T. Huddleston. 1991. Classification
and biology of braconid wasps (Hymenoptera:
Braconidae). Handbooks for the Identificatum of Brit-
ish Insects 7(11): 1-126.
Szollosi, A. 1974. Ultrastructural study of the sperma-
todesm of Locusta migratoria migratoroides (R.F): ac-
rosome and cap formation. Acrida 3: 175-192.
Szollosi, A. 1975. Electron microscope study of spermi-
ogenesis in Locusta migratoria (Insecta Orthoptera).
Journal of Ultrastructure Research 50: 322-346.
Wheeler, D. E., E. G. Crichton and P. H. Krutzsch.
1990. Comparative ultrastructure of ant sperma-
tozoa (Formicidae: Hymenoptera). /oMniii/ of Mor-
phology 206: 343-350.
Wolf, K. W. and H. C. Joshi. 1995. Microtubule or-
ganization and the distribution of 7-tubulin in
spermatogenesis of a beetle, Tenehrio nuttitor (Te-
nebrionidae, Coleoptera, Insecta). Journal of Celt
ScH-nce 108: 3855-3865.
Wolf, K. W. and C. Klein. 1995. Development of the
sperm head in the caddis fly Potainophylax rotun-
dipennis (Insecta, Trichoptera). Zoomorphology
115: 109-115.

J. HYM. RES.
Vol. 7(1), 1998, pp. 38-47

Revision of the American Tiphiid Genus Quemaya Pate
(Hymenoptera: Tiphiidae: Brachycistidinae)

Lynn S. Kimsey and Marius S. Wasbauer

Bohart Museum of Entomology, Department of Entomology, University of California,

Davis, CA 95616, USA

Abstract. — The brachycistidine genus Quemaya is reviewed and six new species are described
from southern California, Arizona, northern Mexico and Costa Rica, confusa, costaricensis, eu-
rekaensis, megalops, mexicana, and sonorensis.

The genus Quemaya Pate (1947) is an ob-
scure group of nocturnal tiphiids original-
ly known from five species described from
the southwestern deserts of North Amer-
ica, as far south as Sonora, Mexico. In this
paper we more than double the known
species in this genus, and extend the
known distribution hundreds of miles.
Additionally, intensive collecting in Costa
Rica by the Instituto Nacional de Biodi-
versidad and by Frank Parker has turned
up a new species from the arid north-
western region of Costa Rica. This region
appears to be the southernmost extension
of this arid nearctic desert habitat.

These wasps are easily overlooked be-
cause of their small size, dark coloration
and nocturnal habits. However, males are
strongly attracted to ultraviolet light sources
at night. Females are as yet unknown. The
new Costa Rican species, costaricensis, has
some characteristics unusual for Quemaya,
including the tiny ocelli and sculptured pro-
podeum, but still has the diagnostic features
for the genus discussed below.

Quemaya is characterized by a combina-
tion of primitive and derived features. The
wing venation is reduced (Figs. 10-17). The
forewing has only one, or less commonly
two, submarginal cells, one discoidal and
one subdiscoidal cell, and the marginal cell
is separated from the costal wing margin.
In the hindwing the cubital vein is nearly
straight meeting the transverse cubital vein

at an angle much greater than 135°. Que-
maya species lack the ventrally "tailed" an-
tennal socket carina, forecoxal stridulatory
structure, scrobal sulcus, mandibular cari-
na, basal "ring" carina of the first gastral
tergum, and carinate gastral sterna char-
acteristic of other brachycistidine genera.
Members of Quemaya all have a distinctly
modified epipygium, that has sublateral
carinae and an emarginate, truncate or con-
vex apex (Figs. 18-21).

MATERIALS

Specimens in this study were obtained
from: DAVIS — Bohart Museum of Ento-
mology, University of California, Davis (S.
L. Heydon); ITHACA — Department of En-
tomology, Cornell University, Ithaca, New
York (J. Liebherr); LAWRENCE— Snow
Entomological Museum, University of
Kansas, Lawrence (R. Brooks); LOGAN —
Department of Entomology, Utah State
University, Logan (T. Griswold); OTTA-
WA— Canadian National Insect Collec-
tion, Agriculture Canada, Ottawa (L. Mas-
ner); RIVERSIDE— Department of Ento-
mology, University of California, River-
side (S. Triapitsyn); SAN FRANCISCO—
California Academy of Sciences, San Fran-
cisco (N. Penny); SANTO DOMINGO—
Instituto Nacional de Biodiversidad, Santo
Domingo de Heredia, Costa Rica (C. M.
Rodriguez); TUCSON — University of Ari-
zona, Tucson; WASHINGTON— U.S. Na-

Volume 7, Number 1, 1998 39

tional Museum of Natural History, Smith- Two abbreviations are used below for

sonian Institution, Washington, D. C. (A. the sake of brevity; MOD = midocellus di-

S. Menke, K. V. Krombein). ameter and F = flagellomere.

KEY TO SPECIES OF QUEMAYA (MALES)

1 Forewing with two submarginal cells (Figs. 10, 15, 16) 2

- Forewing with one submarginal cell (Figs. 11-14, 17) 5

2 Mandible with two apical teeth; distance between midocellus and closest eye margin less
than 1.5 MOD (Fig. 1) arenicola Wasbauer

- Mandible with three apical teeth, one may be very small; distance between midocellus
and closest eye margin more than 1.5 MOD 3

3 Mesopleuron densely punctate, with punctures evenly dispersed over entire surface and
1-2 puncture diameters apart; propodeum coarsely sculptured with irregular demarcation
between dorsal and posterior surfaces indicated by rugae confusa, new species

- Mesopleuron sparsely punctate, with punctures generally 4—6 puncture diameters apart,
denser dorsally than posteroventrally, surface above midcoxa nearly impunctate; propo-
deum smooth, without distinct punctation or rugosities, and no demarcation between
dorsal and posterior surfaces 4

4 Midocellus separated from eye margin in frontal view by more than 2 MOD; epipygium
apically bidentate perpunctata (Cockerell)

- Midocellus separated from eye margin in frontal view by more than 1 MOD but less than

2 MOD (Fig. 6); epipygium apically flat and broadly triangular (Fig. 19) ... megalops, new

species

5 Gular carina with basal tooth-like projection near mandible (Fig. 25); clypeus transversely
indented, without medial projection, arcuately raised apicomedially (Fig. 5); epipygium
apex truncate (as in Fig. 21) inermis (Malloch)

- Gular carina simple; clypeus medially bulging or with medial tooth; epipygium various

6

6 F-1 and II subequal in length, about twice as long as broad or longer; epipygium apex
strongly convex and lip-like (Fig. 20) ntarcida (Bradley)

- F-I shorter than II, and between 1.2 and 1.8x as long as broad; epipygium apex medially
emarginate or truncate 7

7 Midocellus larger, separated by 2 MOD or less from nearest eye margin; F-I 1.6X as long

as broad or longer 8

- Midocellus small, separated by 3 or more MOD from nearest eye margin; F-I 1.5x as long

as broad or shorter 9

8 Midocellus separated from nearest eye margin by 1.5 MOD or less; epipygial apex convex;
clypeus with broad, blunt medial projection, bulging and strongly subtriangular in profile;
forewing Rl vein strongly angulate near costal margin, marginal cell broadly parallel-
sided (as in Fig. 15) mexicana, new species

- Midocellus separated from nearest eye margin by 1.8-2.0 MOD; epipygial apex bidentate;
clypeus with narrow noselike or almost digitate medial projection; forewing Rl vein nar-
rowly separated from stigma, curved or indistinguishable near costal margin

paupercula (Bradley)

9 Midocellus five or more MOD from nearest eye margin (Fig. 3); F-I 1.2x as long as broad;
F-II 1.4X as long as broad; epipygium apex truncate (as in Fig. 21); propodeum coarsely
sculptured, with irregular, partial transverse carina (Fig. 24) ... . costaricensis, new species

- Midocellus 3.0-3.5 MOD from nearest eye margin; F-I 1.5x as long as broad; F-II 1.7-
1.9 X as long as broad; epipygium apex truncate or emarginate; propodeum smooth with-
out sculpturing or transverse carina 10

40

Journal of Hymenoptera Research

10 Clypeus with small, sharp medial projection, apical truncation 1.5 MOD wide (Fig. 4);

epipygial apex medially emarginate, with sublateral carina eurekaensis, new species

- Clypeus with broad, transverse medial projection subtended by discrete declivity, apical

truncation 2 MOD wide (Fig. 9); epipygial apex truncate sonorensis, new species

Quetnaya arenicola Wasbauer

(Figs. 1, 10, 22)

Quemaya arenicola Wasbauer 1967:169. Holotype
male; California: Inyo Co., 6 mi w Glamis
(SAN FRANCISCO, type No. 9306).

Male. — Body length 4-6 mm; clypeus
with narrow apical truncation 0.6 MOD
wide (Fig. 1); mandible with two apical

teeth; gular carina simple; F-I and II length
twice breadth; interantennal distance 0.3
MOD wide; midocellus separated from
nearest eye margin by 1.1-1.2 MOD; fore-
wing with two submarginal cells, second
cell triangular or subquadrate, completely
underlying the first (Fig. 10); mesopleural
punctures 0.2-0.5 puncture diameter
apart; epipygium with sublateral carina

4. eurekensis

5 inermis

7. marcida ^ mexicana

Figs. 1-9. Front view of m,ik' fan', k'ft iintcnna removod.

6 megalops

9. sonorensis

Volume 7, Number 1, 1998

41

10. arenicola

12. eurekaensis

14. marcida

16. perpunctata

22. arenicola 23. marcida

1 1 . costaricensis

13. inermis

15. megalops

17. sonorensis

24. costaricensis

18. confusa

19. megalops

20. marcida

21. eurekaensis

25. inermis

Figs. 10-25. Figs. 10-17. Forewing venation. Figs. 18-21. Apex of epipygium (last gastral tergum). Figs. 22-
23. Lateral view of male genital capsule. Fig. 24. Posterior view of male propodeum. Fig. 25. Oblique posterior
view of male head, showing genal tooth.

42

Journal of Hymenoptera Research

ending in an apical lobe, apical margin
medially emarginate; paramere subapi-
cally expanded, apical margin linear ter-
minating in acute ventral angle (Fig. 22).
Body color reddish brown; antennae pale
red; wing veins clear and at most faintly
tinted with dark brown stigma.

Material examined. — 84 specimens from
California: Imperial, San Bernardino, Riv-
erside and San Diego Co.; Arizona: Yuma
Co.; Mexico: Baja California Norte and So-
nora.

Discussion.— This is one of the few Que-
maya species with two submarginal cells.
It can be immediately distinguished from
the others with similar venation by having
only two mandibular teeth and large ocelli
narrowly separated from the eye margin.

Quemaya confiisa Kimsey and
Wasbauer, new species

(Figs. 2, 18)

Male. — Body length 4-6 mm long; face
(Fig. 2): clypeus medially projecting and
nose-like in profile, apical truncation me-
dially emarginate or slightly trilobate and
1.3 MOD wide; mandible with three apical
teeth, two subsidiary teeth considerably
smaller than apical one; gular carina sim-
ple; F-I and F-II 1.7x as long as wide; dis-
tance between midocellus and closest eye
margin 2 MOD; interantennal space 0.5
MOD wide; mesopleural punctures 0.5-
1.0 puncture diameter apart; forewing
with one submarginal cell; marginal cell
rhomboid, apically broad and Rl angu-
late; epipygium apicomedially emargin-
ate, with two sublateral carinae forming
an ovoid, slightly sunken, medial area and
terminating in apical lobes (Fig. 18); par-
amere nearly parallel-sided, only slightly
widened subapically before acute ventral
angle. Body color black to dark brown; leg
brown; antenna bicolored, paler ventrally
than dorsally; wing veins pale brown-tint-
ed, stigma dark brown; wing membrane
untinted.

Etymology. — This species is named
"confusa" because of the strong similarity

between it and other species of Quemaya
found in the same localities.

Typw material. — Holotype male: MEXI-
CO: Sonora, 6 km nnw San Carlos, 11-15
July 1983, E. Fisher, malaise trap (DAVIS).
Paratypes, 55 males (DAVIS, SAN FRAN-
CISCO, RIVERSIDE, LAWRENCE, OT-
TAWA, WASHINGTON, TUCSON):
two— USA: CALIFORNIA: Riverside Co.,
Blythe, 21 June 1963, F. D. Parker and L.
A. Stange; one — Deep Canyon, 24 June
1964, E. I. Schlinger; one— Millard Cyn., 20
June 1963, E. 1. Schlinger; two — Winches-
ter, 10 July and 14 Aug. 1967, W. Icenogle;
four — Imperial Co., 3 mi n Glamis, 15—16
Sept. 1977, M. Wasbauer & A. Hardy;
one— HoltviUe, 8 July 1965, R. A. Flock;
three — Chocolate Mts., Ogilby Rd., 3 mi s
Jet. Hwy. 78, 16-21 Oct. 1977, M. Was-
bauer; five — San Bernardino Co., 10 mi w
29 Palms, 27 May 1966, M. Wasbauer;
two— NEW MEXICO: Dona Ana Co., Las
Cruces, 2 June 1965, R. M. Bohart; one — 2
mi. e Mesilla; ten— ARIZONA: Pinal Co.,
Marana, 6 July 1955, Butler & Werner;
one — Maricopa Co., Gila Bend, Bohart &
Butler, 12 Aug. 1954; one — Wickenburg,
10 Aug. 1950, H. K Lloyd; one— Pinal Co.,
10 mi. s Toltec, 21 June 1953, T. R. Haig;
nine — 4 mi. se Casa Grande, 18 June 1964,
Smith & Baker; one — Pima Co., Continen-
tal, 17 July 1966, H. K. Court; one— TEX-
AS: Brewster Co., Chisos Mts. 10-12 June
1908, Mitchell & Cushman; one— MEXI-
CO: Sonora, Cocorit, 23 May 1968, Parker
& Stange; one — 11 June, F. D. Parker;
two— 18 mi. e El Puerto, 7 Aug. 1960, Ar-
naud, Ross & Rentz; four — 40 mi. n Her-
mosillo, 8 Aug. 1960, Amaud, Ross &
Rentz; one — 6 km nnw San Carlos, 11-15
July 1983; one — 4.9 mi n Magdalena, Rt.
15, 25 Aug. 1964, M. E. Irwin.

Discussion. — Quemaya confusa is another
species with two submarginal cells. It can
be distinguished from other species with
this kind of wing venation by the densely
and evenly punctate mesopleuron, coarse-
ly sculptured propodeum, and mandibles
with the normal three apical teeth. This

Volume 7, Number 1, 1998

43

species is probably most similar to perpiiii-
cata and megalops.

Quemaya costaricensis Kimsey and
Wasbauer, new species

(Figs. 3, 11, 24)

Male. — Body length 4.0-5.5 mm; face
(Fig. 3): clypeus flat in profile, with apical
truncation 2.5 MOD wide; mandible with
three apical teeth, preapical teeth subequal
in length; gular carina simple; F-I length
1.2X breadth; F-II 1.4x as long as broad;
interantennal distance 0.9 MOD wide; dis-
tance from midocellus to closest eye mar-
gin 5.7 MOD; mesopleuron with punc-
tures 1-2 puncture diameters apart; fore-
wing with one submarginal cell; marginal
cell small, more than 3x as long as broad,
widely separated from costal margin (Fig.
11); propodeum coarsely rugose dorsally,
with broken irregular transverse ridge
separating dorsal from posterior surfaces
(Fig. 24) epipygium apicomedially trun-
cate, with two sublateral carinae above
apex forming an ovoid, slightly sunken
medial area; paramere gently tapering,
apicoventral angle narrowly rounded.
Body color black; antenna bicolored, paler
ventrally than dorsally; wing veins pale
brown, stigma dark brown; wing mem-
brane slightly brown tinted.

Etymologi/. — This species is named after
the country of collection, which is the
southernmost record for the tiphiid sub-
family Brachycistidinae.

Ti/pe material. — Holotype male: Costa
Rica: Guanacaste Prov., 14 km s Caiias, F.
D Parker, 17 Feb. 1989 (LOGAN). Para-
types, 52 males (DAVIS, LOGAN, SANTO
DOMINGO): eight— same data as holo-
type; one: 4-5 Mar. 1989; two— 24 Feb.
1989; one— 28-29 Jan. 1989; three— 18 Feb.
1989; three— 28 Feb. 1989; one— 9 Mar.
1989; one— 11-13 Jan. 1990; one— 16 Feb.
1989; one— 15-24 Feb. 1990; one— 5 Mar.
1989; two— 1-11 Feb. 1990; one— south of
Cafias, 9-14 Feb. 1989, F. D. Parker; one—
25-28 Feb. 1989; one— 25 Feb.-8 Mar.

1989; twenty-one — Santa Rosa Natl. Pk.,
21 Feb.-ll-14 Mar. 1981.

Discussion. — There are many distinctive
features of this species, which will imme-
diately separate it from all other Quemaya,
including: the tiny ocelli and small eyes,
coarsely sculptured propodeum and short,
broad flagellomeres. It is closest to sono-
reiisis and eurekaensis based on the small
ocelli, single submarginal cell and short
broad flagellomeres.

Quemaya eurekaensis Kimsey and
Wasbauer, new species

(Figs. 4, 12, 21)

Male. — Body length 3.5^.0 mm; face
(Fig. 4): clypeus with small, sharp medial
projection, apical truncation 1.5 MOD
wide; mandible with three apical teeth;
gular carina simple, without tooth-like
projection; F-1 length 1.5x breadth; F-II
length 1.9 X breadth; interantennal dis-
tance 0.5 MOD wide; midocellus separat-
ed from nearest eye margin by 3.2 MOD;
mesopleural punctures 3-6 puncture di-
ameters apart; forewing with one submar-
ginal cell (Fig. 12); epipygium with sub-
lateral carinae each ending in an apical
lobe, epipygial apex medially emarginate
(Fig. 21); gonostylar shape apically nar-
rowed into single acute apical angle. Body
dark reddish brown, antennal and leg col-
or pale reddish brown; wing veins untint-
ed, except stigma dark reddish brown.

Etymology. — This species is known pri-
marily from Eureka Valley, thus the name.

Type material. — Holotype male: Califor-
nia: Inyo Co., Eureka Valley, 13 July 1975,
F. Andrews & A. Hardy (DAVIS). Para-
types, eight males (DAVIS, SAN FRAN-
CISCO); six — same data as holotype;
one — 19 June 1972 Derham & Guiliani;
one — Saline Valley dunes, 30 km e Inde-
pendence, 26 May 1993, D. E. Russell,
malaise trap.

Discussion.— The diagnostic features for
eurekaensis are the small ocelli, separated
from the eye margin by more than 3
MOD, medially emarginate epipygium

44

Journal of Hymenoptera Research

apex, and short F-I and II. This species is
closest to sonorensis and less so costaricen-
sis, but can be immediately distinguished
from sonorensis by the sharp clypeal pro-
jection and emarginate epipygium. The
larger ocelli and smooth propodeum will
separate eurekae)isis from costaricensis.

Quemaya inermis (Malloch)

(Figs. 5, 13, 25)

Brachycistis inermis Malloch 1924:23. Holotype
male; Arizona: Higley (WASHINGTON).

Male. — Body length 2>-^ mm; face (Fig.
5): clypeus transversely indented, without
medial projection, apical truncation 0.9
MOD wide; mandible with three apical
teeth, preapical teeth subequal; gular ca-
rina with tooth-like projection near base of
mandible (Fig. 25); forewing with one sub-
marginal cell (Fig. 13); F-I and II length
1.7X breadth; interantennal distance 0.8
MOD wide; midocellus separated from
eye margin by 2 MOD; mesopleuron an-
terior part with punctures 1-3 puncture
diameters apart, posteriorly nearly im-
punctate and polished; epipygium with-
out sublateral carinae, only slightly in-
dented apicomedially; paramere gently ta-
pering, apicoventral angle narrowly
rounded. Body color reddish brown; an-
tenna paler than body; wing veins and
stigma nearly colorless.

Material examined. — 239 specimens from:
California: Riverside, San Bernardino, and
Imperial Cos.; Arizona: Yuma, Santa Cruz
and Maricopa Cos. Nevada: Lincoln Co.;
Mexico: Sonora and Baja California Norte.

Discussion. — This is the only Quemaya
species with a tooth-like projection on the
gular carina; a feature typically found in
members of the genus Brachycistis. Que-
maya inermis can be immediately recog-
nized by the transversely medially indent-
ed clypeus, and the apically truncate epi-
pygium.

Quemaya marcida (Bradley)

(Figs. 7, 14, 20, 23)

Bracln/cisttfi marcida (Bradley) 1917:283. Holo-
type male; USA: California, Imperial Co.
(ITHACA).

Male. — Body length 3-5 mm; face (Fig.
7): clypeus with sharp medial projection,
appearing nasiform in profile, apical trun-
cation 1.7 MOD wide; mandible with three
apical teeth; gular carina simple; F-I and
F-II twice as long as broad; interantennal
distance 0.6 MOD wide; midocellus sepa-
rated from eye margin by 2.5 MOD; meso-
pleuron sparsely punctate, punctures 4-6
puncture diameters apart or more; fore-
wing with one submarginal cell; marginal
cell nearly parallel-sided, Rl vein strongly
angulate and joining stigma at or below
costal margin (Fig. 14); epipygium apico-
medially rounded and lip-like, with sub-
lateral carina beginning above lip (Fig. 20);
paramere apically truncate with apical
margin concave (Fig. 23). Body reddish
brown to brown; antenna and legs paler
reddish brown than body color, forewing
veins pale brown tinted; stigma brown.

Material examined. — 873 specimens from
CALIFORNIA: San Bernardino, Riverside
and Imperial Cos.; Arizona: Yuma and Co-
chise Cos.; Nevada: Nye; Mexico: Sonora.

Discussion. — As with the majority of
Quemaya species, marcida has one submar-
ginal cell in the forewing and a simple gu-
lar carina. However, of these species only
marcida, arenicola and perpuncata have the
first two flagellomeres twice as long as
broad. Quemaya marcida is the only one
that has the epipygial apex strongly con-
vex and lip-like. Additionally, the ocelli
are large and narrowly separated from the
nearest eye margin.

Quemaya megalops Kimsey and
Wasbauer, new species

(Figs. 6, 15, 19)

Male. — Body length 4-5 mm; face (Fig.
6): clypeus with large tuberculate medial
projection, strongly nasiform in profile,
apical truncation 0.9 MOD wide; mandible
with three apical teeth, subsidiary ones
considerably shorter than primary tooth;
gular carina simple; F-1 and 11 length 1.9-
2.0 X breadth; interantennal distance 0.5
MOD wide; midocellus separated from eye

Volume 7, Number 1, 1998

45

margin by 1.6 MOD; mesopleuron nearly
impunctate; forewing with two submargin-
al cells, the second large, nearly rectangular
and Rs aligned with 2rs-m; marginal cell
large and parallel-sided with Rl vein
strongly angulate before joining stigma at
costal margin (Fig. 15); epipygium apically
truncate, with distinct sublateral carinae,
flat medially (Fig. 19); paramere lanceolate,
narrowly tapering apically. Body reddish
brown to brown with yellow legs and an-
tenna; wing veins and stigma transparent
and lightly yellow tinted.

Ehpiiologi/. — mega = large, ops = eyes; i.
(Gr.). The name refers to the greatly en-
larged eyes and ocelli.

Ti/pe material. — Holotype male: Califor-
nia: Riverside Co., 5 mi nw Desert Center,
22 Oct. 1955, M. Wasbauer (DAVIS). Para-
types: 7 males (DAVIS, RIVERSIDE, SAN
FRANCISCO): one — same data as holo-
type; two: Indio, Aug. 1977, Allen and
Duffy (DAVIS); one — Magnesia Cyn. 2
July 1952; one — Inyo Co., 7 mi nne Pana-
mint Spr., 15 May 1969, P.Rude; one — Ar-
izona: Yuma Co., 4 mi w Salome, 8 June
1958, D & G. MacNeill; one — Mohawk,
Apr. 1963, Timberlake.

Discussion. — This species most closely
resembles perpuncata as both have two
submarginal cells, a sparsely punctate
mesopleuron and smooth propodeum.
Quemaya megalops can be distinguished
from perpuncata by the much larger ocelli,
separated from the nearest eye margin by
less than two midocellus diameters, pale
stigma, and the very distinctive epipy-
gium, which appears broadly triangular
with the apex narrowly truncate. The
wing venation is also diagnostic, with the
second submarginal cell much larger than
in other Quemaya species.

Quemaya mexicana Kimsey and
Wasbauer, new species

(Fig. 8)

Male. — Body length 2.5-4.5 mm; face
(Fig. 8): clypeus with broad, blunt medial
projection, bulging and strongly subtrian-

gular in profile, apical truncation 1 MOD
wide; mandible with three apical teeth;
gular carina simple; F-I-II 2.0-2.4 X as
long as broad; interantennal distance 0.4
MOD wide; midocellus separated from
eye margin by 1.5-2.0 MOD; mesopleuron
sparsely punctate, punctures 4-6 puncture
diameters apart or more; forewing with
one submarginal cell; marginal cell nearly
parallel-sided, Rl vein strongly angulate
and joining stigma at or below costal mar-
gin; epipygium apicomedially convex,
otherwise similar to sonorensis; gonostylar
apex truncate, with apical margin linear.
Body brown to dark brown; antenna and
legs paler reddish brown than body color,
forewing veins pale brown tinted; stigma
brown.

Etymology. — The name refers to the pre-
ponderance of specimens having been col-
lected in northern Mexico.

Material examined. — Holotype male:
MEXICO: Baja California Norte, 1 km s El
Rosario, 24-25 July 1992, D. E. Russell, MT
(DAVIS). Paratypes: 46 males (DAVIS):
37— Diablo Cyn, Dry Lake, 16 July 1979,
D. Giuliani; four — same data as holotype;
three — eastern base of Sierra de Juarez be-
low Rumorosa, 11 Sept. 1961, I. L. Wig-
gins; one — USA: California, Andrade, 4
Aug. 1966, M. Wasbauer; one — Calexico,
19 June 1969.

Discussion. — This species closely resem-
bles marcida as discussed under that spe-
cies. However, mexicana can be readily
distinguished from marcida by the shorter
flagellomeres, broad blunt clypeal projec-
tion, and distinctively convex epipygial
apex.

Quemaya pauperciila (Bradley)

Brachycistis pauperciila Bradley 1917:282. Holo-
type male; California: Calexico (ITHACA,
type No. 129.1)

Male. — Body length 3-4.5 mm; clypeus
with medial nose-like projection in profile,
apical truncation 1.1 MOD wide; mandible
with three apical teeth; gular carina sim-

46

Journal of Hymenoptera Research

pie; F-I and II length 1.6X breadth; inter-
antennal distance 0.3 MOD wide; mido-
cellus separated from eye margin by 1.9
MOD; mesopleural punctures almost ab-
sent; forewing with one submarginal cell;
marginal cell narrow, closed on costal
margin; epipygial apex truncate, subapi-
cally concave; paramere nearly parallel-
sided, apex abruptly truncate, with acute
ventral and dorsal angle and apical mar-
gin somewhat concave between. Body col-
or dark brown; antenna paler than body;
wing veins nearly colorless, except stigma
dark brown.

Material examined. — 109 specimens from
California: Riverside, Imperial, Kern, San
Bernardino and San Diego Co.; Arizona:
Coconino, Yuma Co.; Nevada: Lincoln
Co.; Texas: Brewster and and Presidio Co.;
New Mexico: Dona Ana Co.; Mexico: Baja
California Norte and Sonora.

Discussion. — Queniaya paupercula is one
of the most commonly collected species of
the genus. It is most readily confused with
eurekaensis, which has a very restricted
distribution. Both species have a medially
emarginate epipygial apex, short flagel-
lomeres, two submarginal cells and a sim-
ple gular carina. However, the larger size
of the ocelli will readily separate pauper-
cula from eurekaensis.

Quemaya perpunctata (Cockerell)

(Fig. 16)

Brachycistis perpunctata Cockerell 1896:291. Ho-
lotype male; New Mexico: Las Cruces (PHIL-
ADELPHIA).

Male. — Body length 4.0-5.5 mm; clypeus
with low medial projection, apical margin
1 MOD wide; mandible with three apical
teeth; gular carina simple; F-I length twice
breadth; F-II 2.2X as long as broad; inter-
antennal distance 0.5 MOD wide; mido-
cellus separated from eye margin by 2.8
MOD; forewing with two submarginal
cells, second cell triangular or subquad-
rate, completely underlying the first (Fig.
16); mesopleural punctures 1-2 puncture

diameters apart, nearly impunctate above
midcoxa; epipygium with sublateral cari-
nae each ending in an apical lobe, epipy-
gial apex medially emarginate; paramere
expanded subapically, apex truncate, api-
cal margin linear, between two acute an-
gles. Head and body dark brown to black,
except antennae, clypeal apex, palpi and
tegula reddish brown, mandibles and tarsi
yellow, femora and tibiae brown; stigma
dark brown, nearly black, veins brown.

Material examined. — 249 specimens from
California: San Bernardino, Riverside, Im-
perial and San Diego Cos.; Nevada: Lin-
coln Co.; Texas: Brewster Co.; Arizona:
Yuma Co.; Mexico: Baja California Sur.

Discussion. — The most striking feature
of this species is the wide range of varia-
tion in coloration. This variation appears
to be geographically correlated. In Califor-
nia specimens some red is always present
on at least the head. The prothorax and
often the entire thorax may also be red,
giving the specimen a strongly bicolored
appearance. Otherwise, perpuiwtata can be
recognized by having two submarginal
cells, three mandibular teeth, propodeum
without distinct sculpturing, and small
ocelli.

Quemaya sonorensis Kimsey and
Wasbauer, new species

(Figs. 9, 17)

Male. — Body length 3.5^.0 mm; face
(Fig. 9): clypeus with broad transverse me-
dial projection subtended by discrete de-
clivity, apical truncation 2 MOD wide;
mandible with three apical teeth; gular ca-
rina simple; F-I length 1.5x breadth; F-II
length 1.7X breadth; interantennal dis-
tance 0.7 MOD wide; midocellus separat-
ed from eye margin by 3 MOD; mesopleu-
ron smooth, punctures 1-4 puncture di-
ameters apart becoming sparser above
midcoxa; forewing with one submarginal
cell, venation as in paupercula (Fig. 17);
epipygium apex truncate; paramere as in
costaricensis. Body and antennal color pale

Volume 7, Number 1, 1998

47

reddish brown; wing veins untinted, ex-
cept stigma reddish brown.

Etymologi/. — The type series of this spe-
cies is from Sonora, Mexico; thus the
name.

Type material. — Holotype male: MEXI-
CO: Sonora, 6 km nnw San Carlos, 11-15
July 1983, E. Fisher, MT (DAVIS). Para-
types: 11 males (DAVIS), same data as ho-
lotype.

Discussion. — The small ocelli, short fla-
gellomeres, single submarginal cell, trun-
cate epipygial apex and lack of a gular
tooth distinguish sonorensis from most oth-
er Qiiemaya species, except eurekaensis and
costaricensis. Quemaya sonorensis can be
distinguished from these two species by
the larger ocelli and flagellomeres I and II
1.5 X as long as broad or longer, not short-
er as in costaricensis, narrow clypeal trun-
cation and broad transverse clypeal pro-
jection.

ACKNOWLEDGMENTS

This study was made possible by the assistance of
all of the collections and curators who provided spec-
imens. Thanks also to Maria Garcia who did the final
preparation of the ink drawings, and to the editors
for their suffering.

LITERATURE CITED

Bradley, J. C. 1917. Contributions toward a mono-
graph of the Mutillidae and their allies of Amer-
ica north of Mexico. Transactions of the American
Entomological Society 43:247-290.

Cockerell, T. D. A. 1896. Descriptions of new Hyme-
noptera. Transactions of the American Entomologi-
cal Society 22:289-297.

Malloch, ]. R. 1924. A new species of the genus Bra-
chycistis. Bulletin of the Brooklyn Entomological So-
ciety 19:23.

Pate, V. S. L. 1947. A conspectus of the Tiphiidae,
with particular reference to the nearctic forms.
journal of the New York Entomological Society 54:
115-145.

Wasbauer, M. S. 1967. A new species of Quemaya
from the Colorado desert of California. Proceed-
ings of the Biological Society of Washington 80:169.

J. HYM. RES.
Vol. 7(1), 1998, pp. 48-56

Geographic Variation of Sex Ratio in Pelecinus polyturator (Drury)

(Hymenoptera: Pelecinidae)

Norman F. Johnson and Luciana Musetti

Department of Entomology, Museum of Biological Diversity, 1315 Kinnear Road,
The Ohio State University, Columbus, OH 43212-1192, USA

Abstract. — The relative abundance of males and females of Piieciiius poh/turafor (Drury) (Hy-
menoptera: Pelecinidae) was examined on the basis of specimens held in natural history collec-
tions. The species may be divided into two groups of populations. Those in the United States and
Canada (between 28°N and SFN) are primarily thelytokous: males form only 4% of the total
number of individual specimens. Populations from localities 23°N and southward have a sub-
stantially higher frequency of males (36%). Within each group of populations, there is no demon-
strable change in sex ratio with latitude. Male emergence dates generally precede those of females,
but there seems to be no significant difference in the time period in which the two sexes are flying.
Within the U.S. and Canada, the uncommon males are not randomly distributed.

The parasitoid wasp Pelecinus polytura-
tor (Drury) (Hymenoptera: Pelecinidae) is
a large and familiar inhabitant of moist
deciduous forests in the Nearctic. Its range
extends well beyond this, generally from
southeastern Canada to central Argentina
(Muesebeck 1979, Masner 1993). Surpris-
ingly little is known of Pelecinus biology
despite its relative abundance. The few
published host records indicate that this
wasp is an internal larval parasitoid of
soil-dwelling Scarabaeidae (Coleoptera)
(summarized in Lim et al., 1980). One as-
pect of its biology that has received wide
comment, however, is that males are ex-
tremely uncommon in the northern por-
tion of the range.

Brues (1928) cited Pelecinus polyturator
as an example of the phenomenon of geo-
graphic parthenogenesis. Arrhenotokous
parthenogenesis is the most common
mode of reproduction in the Hymenop-
tera, in which males are usually produced
from unfertilized eggs. Thelytoky is not at
all rare, being known from at least twenty
families (Stouthamer et al. 1990). Pelecinus
appears to demonstrate both modes: the-

lytoky in the north temperate region and
arrhenotoky elsewhere.

At the time that he wrote his paper
Brues admitted that he had seen very few
male specimens of P. polyturator from the
United States or Canada. His analysis of
the sex ratio of the species was anecdotal.
His sampling of Pelecinus was never de-
scribed and must have been quite limited
(see below). Neither did he actually quan-
tify observed sex ratios in tropical and
temperate regions. Our purpose here is to
quantify the issue of sex ratio variation in
this wasp throughout its geographical
range.

MATERIALS AND METHODS

Specimen data. — Our information on the
distribution and relative abundance of
males and females is based upon an ex-
tensive survey of the holdings of Pelecinus
in natural history collections. Material for
this study was borrowed or data acquired
from 91 collections around the world (see
Appendix). The information associated
with specimens varies extensively in ac-
curacy and completeness, especially given

Volume 7, Number 1, 1998

49

the small size of the typical label attached
to a specimen.

The data were transcribed and stored in
a specimen-level relational database. The
table structure is slightly modified from
the information model developed by the
Association of Systematics Collections
(1993) and is implemented in the Oracle?®
environment on a Silicon Graphics (UNIX)
workstation. This combination of hard-
ware and software was chosen to deal
with the large numbers of specimens in
insect collections and for its ability to in-
terface with other software (geographic in-
formation systems, mapping software,
and World Wide Web servers). The data-
base stores all of the information on spec-
imen labels (place, time, method of collec-
tion, etc.), characteristics of the specimens
(e.g., sex, color pattern), source of materi-
al, and literature references. The relational
structure allows the development of ad
hoc queries unconstrained by the format
of the original data. As such, the system
is not only suited to the questions we ask
here, but is also applicable to collection
management, diversity assessments, taxo-
nomic studies, host-parasitoid biology,
etc. The database (Johnson & Musetti
1996), intended to represent the sum of
documented geographic and temporal in-
formation available, contains data from
7,188 specimens of the genus Pelecinus.

Latitude and longitude of collecting lo-
calities were added where these could be
determined with relative confidence.
These are stored in two separate tables, for
points and polygons, reflecting the level of
accuracy of the cited collecting locality
and our ability to locate the sites in atlases
and gazeteers. Only those classed as
"points" were used in the analyses below.
In practice, this means that the collecting
records for points consist of localities
identified with specific populated places,
recreational areas, manmade features (e.g.,
monuments), or geographic features such
as mountains and lakes.

Brues (1928) pointed out that Neotrop-

Table 1. Numbers of specimens of males and fe-
males of Pelecinus polyturator used in analyses of sex
ratio.

latitude Ul

localities

Xumber of
6£ [%1

Number o\

S 9 (%]

T,it.il
mini her

25°N-5rN

38°S-25°N

Total

119 (4.2]
616 |38.4]
735 116.5)

2723 [95.8]
990 161.6]

3713 183.5]

2842
1606
4448

ical specimens of Pelecinus exhibit notable
variation in color patterns. Many of these
were described as distinct species in the
early 19th Century, but the present taxo-
nomic consensus (dating from Schletterer
1890) is that only a single species, P. po-
lyturator, is recognized as valid. To avoid
confounding data from possible distinct
species, we chose to include in our anal-
yses only those specimens conforming to
the color pattern of specimens from the
U.S. and Canada: the head, mesosoma,
and metasoma are uniformly black or very
dark brown, and the fore wings are clear
or gradually infuscate toward the apex. A
summary of the numbers of specimens
used is presented in Table 1.

Analyses. — Coordinates of latitude and
longitude of collecting localities were ex-
tracted from the database by sex. Brues
(1928) asserted (as does conventional wis-
dom) that males and females are not iden-
tically distributed. This was tested by
comparing the cumulative relative fre-
quency distribution of specimens by 1° of
latitude using the Kolmogorov-Smirnov
test (Sokal & Rohlf 1995).

If there is variation in sex ratio among
sites, especially over the vast area occu-
pied by this species, one reason may be
that northern females are substantially
longer-lived as adults than males, thus
leading to an overabundance of females in
collection records. To examine this possi-
bility, we sorted the collecting date re-
cords for specimens by sex and combined
them into groups for each 10 degrees of
latitude (from 50°N to 40"S). Collecting
dates were expressed in terms of polar co-

50

Journal of Hymenoptera Research

Fig. 1. World distribution of Pticciiius polyturnlcr.

ordinates by Julian date (1-365) and mean
and standard deviation of the dates were
calculated for each sex in every 10° band.
Sex ratio data are expressed as the pro-
portion of males in the total population of
specimens. Specimens for every 5° band of
latitude were pooled and the relationship
between sex ratio and latitude of collec-
tion examined using regression. For rea-
sons developed below, these data were
partitioned into two groups north and
south of 25°N. The two partitions then
were separately analyzed for relationship
between latitude and sex ratio. Finally, on
the basis of this data partition, we exam-

ined the hypothesis that males in the U.S.
and Canada are randomly distributed.
Specimens were pooled from blocks of 5°
of latitude and longitude and the ob-
served number of specimens was com-
pared using a x' test with that expected
using the observed sex ratio of all speci-
mens north of Mexico.

RESULTS

The documented range of Pelecinii^ poh/-
turator extends from a maximum of 51°N
in Quebec and Ontario south to 38°S in
Argentina (Fig. 1 ). The species does not
occur in Chile or the West Indies (includ-

Volume 7, Number 1, 1998

51

100

>■
o

s

K
IL

III

>

m

80

60

20

max diff = 0 367
P< 0001

f j^fflO-*'""*'**

-40 -20 0 20 40

LATITUDE, DEGREES

60

Fig. 2. Cumulative relative frequency distribution
(%) of male and female specimens of Pelkiinis poly-
luratcr by latitude. Localities grouped for every 1° of
latitude. Maximum difference and probability of
identity of distributions using Kolmogorov-Smirnov
test.

ing Trinidad). Specimens are otherwise
found throughout South America, al-
though material from the Amazon Basin
is very scarce. In the United States, the
range of the species extends west to 106°
in Colorado and New Mexico. The species
is apparently absent from peninsular Flor-
ida south of the Gainesville area.

The cumulative relative frequency dis-
tributions of the two sexes, pooled into 1°
increments of latitude, is presented in Fig.
2. The comparison of these two distribu-
tions clearly leads to rejection of the null
hypothesis that the two are identically dis-
tributed throughout America. Specimens
of males are clearly more abundant out-
side of temperate North America.

Two possible sources of sampling error
that could lead to the observation of high-
ly skewed sex ratios are (1) the samples
were taken in particular years in which
one sex is either very rare or extraordinar-
ily abundant, and (2) if populations are
strongly protandrous or adult females sur-
vive much longer than males, then the ap-
parent rarity of males in some areas may
be a collecting artifact. The frequency dis-

300

S 200

O
ui

a.
w

III

a

s

3

100

I ,l.li

11

187S 1900 192S 1950 1975 2000

YEAR

Fig. 3. Frequency distribution of collecting years of
PelecinKS polyturator. Three specimens collected be-
tween 1804 and 1875 are not illustrated.

tribution of collecting dates by year is il-
lustrated in Fig. 3. The distribution of col-
lecting dates for each sex were pooled into
groups by 10° of latitude (Fig. 4); statistics
for each sex in the latitudinal bands is pre-
sented in Table 2. The average collecting
dates for males generally precedes those
of females, but do not differ significantly.
Further, it seems to be possible to find
males through most of the flight time of
females.

Figure 5 demonstrates a significant
negative relationship between latitude
and sex ratio (with southern latitude ex-
pressed as negative numbers). The ob-
served sex ratio varies from 0.0-0.60, with
an average of 0.19. Even cursory exami-
nation of the data reveals that this is not
a continuous decrease in the frequency of
males, but that the change to spanan-
drous populations (with <10% males) oc-
curs rather abruptly. We have no knowl-
edge of any specimens of Pelecinus col-
lected between 23° and 28°N latitude, and
the sex ratios on either side of this gap
differ strongly. Therefore the data were
partitioned into two components at 25°N
(Fig. 6). Separate regression analyses re-
sult in a change in slope from negative to

52

N

0°-10°183

10°- 20°

o or>o

20°- 30

30°- 40'

40°- 50°

Fig. 4. Collecting dates for males and females of Piiecinus polyturator. Localities grouped for every 10° of
latitude. Julian day 1 = 1 January; 92 = 2 April; 183 = 2 July; 274 = 1 October; N: north latitudes; S: south
latitudes.

Journal of Hymenoptera Research

s

cf 9

positive, but neither significantly differs
from a slope of 0, i.e., there is no demon-
strable relationship between latitude and
sex ratio in the two groups. The average
sex ratio for the southern populations is

0.36 (0.20-0.60), and that for the northern
populations is 0.04 (0.00-0.06). Figure 7
maps the abundance of males and fe-
males in 5° blocks of latitude and longi-
tude. Pooling specimens by latitude or

Volume 7, Number 1, 1998

53

Table 2. Dates of collection of Pelecinin pclyturalor specimens, localities within 10' bands of latitude pooled
together, v: mean Julian date of collection (calendar date); sd: standard deviation; N: number of specimens.

Degrees
latitude

\orth

South

f''

0°-10°

X

144.7 (25 May)

153.4 (2 Jun)

22.7 (23 Jan)

344.8 (11 Dec)

sd

7.1

8.5

20.7

26.3

N

137

222

16

27

10°-20°

.V

181.0 (30 Jun)

185.7(5 Jul)

344.0 (10 Dec)

24.3 (24 Jan)

sd

4.8

5.4

0.5

17.9

N

133

390

2

11

20°-30°

X

209.3 (28 Jul)

195.9 (15 Jul)

39.0 (8 Feb)

37.5 (7 Feb)

sd

3.4

6.9

12.2

18.5

N

26

30

107

189

30°-t0°

.Y

213.2 (1 Aug)

222.4 (10 Aug)

48.3 (17 Feb)

25.7 (26 Jan)

sd

2.3

3.4

0.8

22.0

N

18

897

3

8

40°-50°

X

228.1 (16 Aug)

230.7 (19 Aug)

sd

2.4

2.8

N

88

1277

70^

y = -0 45x + 27 25
r = 0.334

LATITUDE, DEGREES

Fig. 5. Sex ratio for all specimens of Pdcciiins ;'o/i/-
lurntcr (proportion of males in total) as a function of
latitude, with regression and 95% confidence limits.
Individual localities pooled for every 5° of latitude;
south latitudes expressed as negative numbers.

longitude (Table 3) reveals that males do
not appear to be randomly distributed
through the U.S. and Canada.

DISCUSSION

Collecting records are not random sam-
ples and we recognize a number of pos-
sible biases in the data. Female Pelecinus
are large and "unusual," easily identifia-
ble, relatively slow fliers, and are often
found resting on vegetation at heights ac-
cessible to collectors. Thus, females are
commonly found in the holdings of even
small collections and may be overrepre-
sented. On the other hand, their numerical
abundance and the fact that only a single
species is recognized may cause experi-
enced collectors to ignore them. Males
may be relatively scarce in collections be-
cause their abundance in some areas may
be truly low, or they may be overrepre-
sented precisely because of their rarity, at
least in the eyes of North American col-

54

Journal of Hymenoptera Research

80

70 I- y = 0406x-21 17
r = 0.088

-40 -20 0 20 40

LATITUDE, DEGREES

Fig. 6. Sex ratio for Peleciuus polyturator (proportion
of males in total) as a function of latitude, with re-
gression and 95% confidence limits. Data partitioned
into two groups: localities north of 25°N, and locali-
ties south of 25°N. Individual localities pooled for ev-
ery 5^ of latitude; south latitudes expressed as nega-
tive numbers.

lectors. The magnitude and net effect of
these biases are impossible to quantify.
However these collections represent the
material foundation upon which every-
thing we know about this species is based

Table 3. Test of null hypothesis that male speci-
mens in Fig. 7 are distributed randomly among
blocks in the U.S. and Canada. When pooling by lon-
gitude, the specimens from the two westernmost col-
umns of cells and the three easternmost columns of
cells were summed to obtain expected numbers great-
er than five. Expected numbers of males based on
overall sex ratio in America north of Mexico: 4.0%
males. ": probability < 0.01; *: probability < 0.05.

specimens in 5" blocks pooled by lati-
tude " 38.6" 2
specimens in 5° blocks pooled by longi-
tude 10.2* 4

and is the only sample available from
which to estimate the sex ratio. We believe
that we can fairly judge the hypothesis
presented by Brues with cautious use of
the specimen data from collections.

Our survey of collections produced only
83 male specimens of Pelecinus that we are
certain would have been available to
Brues (i.e., collected in 1928 or earlier; a
further 75 males have no year of collection
on the label). Even on the basis of such
limited data, it appears that Brues gener-

-

2d'
94?
2.1%

8^
143?
5.3%

Od-
26?
0.0%

Od-
4?
0.0%

27 d'
89?
23.3%

Od'
36?
0.0%

Od*
4?
0.0%

Oo-

3 ?

0.0%

37.^
399?*'-^
8.5%

1 cf
15?
6.3%

1 rf
55?
1.8%

2d'
91 ?
2.2%

17 0-
292 ?
5.7%

90"
80?

iai%

15 d"
343?
4.2%

lOd'
482?
2.0%

80"
19 ?
29.6%

Oo-
5 ?
0.0%

63 o'
1382 $
4.4%

30*
30?

12?

5^
70?
6.7%

Od-
110?
0.0%

1 d'

203?
0.5%

Sd"
344?.
1.4%

Oo-
7 ?
0.0%

So*
23 ?
25.8%

Oo-
8 9
0.0%

14 d'
776?
1.8%

9.1%

0.0%

Orf
2?
0.0%

Od-
12?
0.0%

Oo"
38?
0.0%

Od-
110?
0.0%

Oo"
3?
0.0%

lOd-
525 ?
1.9%

Od-

165 ?
0.0%

4^
45?
8.2%

163?
1.8%

15d"
316 ?
4.5%

170*
466 ?
3.5%

lOd*
397 ?
2.5%

47 0"
779?
5.7%

114o'

2722?

4.0%

Fig. 7. Numbers of males, females, and sex ratio (proportion of males In total) of thclytoknus populations
of Pclcciiius polyturator in blocks of 5° latitude and longitude in the U.S. and Canada.

Volume 7, Number 1, 1998

55

ally described the true situation: males are
very scarce in temperate North America,
and elsewhere they occur in numbers con-
sistent with a 0.50 sex ratio. There is an
abrupt transition between the two popu-
lations that corresponds with a geograph-
ical disjunction in southern Texas and
northern Mexico. Further focussed collec-
tions are needed to determine whether
this disjunction is real and, if not, what
happens to the males in that area.

We cannot yet identify any diagnostic
morphological differences between the
northern thelytokous populations and the
bisexual populations to the south. Speci-
mens from southern Mexico (Chiapas) and
Central America are often distinguishable,
but typical black specimens are found
from Mexico to Argentina. There is pre-
cious little information on the biology of
Pelecinus, but Aguiar (1997) has recently
described the copulatory behavior of in-
dividuals in Brazil, consistent with the
idea that males in the tropics and south
temperate regions are functional, i.e., that
females do indeed mate.

Brues (1928) additionally speculated on
the genetics of Pelecinus. He asserted that
the largest specimens were found in the
north and suggested that these may be tet-
raploid. The largest specimens we have
seen, however, are from Argentina and a
great range of sizes may be found even in
single populations in the U.S. The size
variation could be better explained by
variation in host size than by invoking
unexplored genetic mechanisms.

The discovery of the nonrandom distri-
bution of males in the United States and
Canada was particularly surprising. This
could be the result of a statistical artifact
or something unusual may be occurring in
some populations. The most notable of
these is a population near Ottawa, Ontar-
io: male specimens have been consistently
collected in this single site over a period
of ten years. Young (1990) described a po-
tentially similar situation in southern Wis-
consin in which he suggested that a bisex-

ual population may have replaced the the-
lytokous strain. The Ottawa population
could be a promising one upon which to
focus in order to better understand the
role of males in the northern temperate
populations of Pelecinus.

ACKNOWLEDGMENTS

Thanks to the many curators who made their ma-
terial and data available to us; to A. P. Aguiar, A.
Sharkov, and J. W. Wenzel for constructive comments
on the manuscript. This material is based in part
upon work supported bv the National Science Foun-
dation under Grant No. DEB-9521648.

LITERATURE CITED

Aguiar, A. P. 1997. Mating behavior of Peteaiuis po-
lyturatcr (Hymenoptera: Pelecinidae). Entomolog-
ical News 108: 117-121.

Association of Systematics Collections. 1993. An in-
formation model for biological collections. Re-
port of the Biological Collections Data Standards
Workshop, August 18-24, 1992. URL: gopher://
kau\keil.ukans.edu:70/ll/staniiards/asc.

Brues, C. T. 1928. A note on the genus Pelecinus. Psy-
che 35: 205-209.

Johnson, N. F. and L. Musetti. 1996. The Pelecinus pro-
ject. URL: http://iris.hiosci.ohio-state.edu/Pelecinus.

Lim, K. P., W. N. Yule, and R. K. Stewart. 1980. A
note on Pelecinus polyturator (Hymenoptera: Pe-
lecinidae), a parasite of Phyllophaga anxin (Cole-
optera: Scarabaeidae). The Canadian Entomologist
112: 219-220.

Masner, L. 1993. Chapter 13. Superfamily Proctotru-
poidea. Pages 537-557, in H. Goulet & ]. Huber,
eds., Hymenoptera of the world: an identification
guide to families. Research Branch, Agriculture
Canada, Publication 1894/ E. 668 pp.

Muesebeck, C. F. W. 1979. Superfamily Pelecinoidea.
Pages II 19-1 120, m K. V. Krombein, P. D. Hurd,
Jr., D. R. Smith, and B. D. Burks, Catalog of Hy-
menoptera in America north of Mexico. Smith-
sonian Institution Press, Washington, DC. 3 vol.,
2735 pp.

Schletterer, A. 1890. Die Hymenopteren-Gattungen
Stenophasmus Smith, Monomachus Westw., Peleci-
nus Latr. und Megalyra Westw. Berliner Entomo-
togische Zeitschrift 33: 197-250.

Sokal, R. R. and F. J. Rohlf. 1995. Biometry. The prin-
ciples and practice of statistics in biological re-
search. Third edition. W. H. Freeman and Com-
pany, New York. 887 pp.

Stouthamer, R., J. D. Pinto, G. R. Platner, and R. F.
Luck. 1990. Taxonomic status of thelytokous
forms of Trichogramma (Hymenoptera: Tricho-

56

Journal of H'imenoptera Research

grammatidae). Annals of the Entomological Societi/
of America 83: 475-481.
Young, D. K. 1990. Distribution of Pciecinus polytura-
tor in Wisconsin (Hymenoptera: Pelccinidae),
with speculations regarding geographical par-
thenogenesis. Tlie Great Lakes Entomologist 23:
1-4.

APPENDIX

Sources of material. American Entomological Insti-
tute, Gainesville, FL; American Museum of Natural
History, New York, NY; Academy of Natural Sci-
ences, Philadelphia, PA; Buffalo Museum of Science,
Buffalo, NY; California Academy of Sciences, San
Francisco, CA; Albertson College of Idaho, Caldwell,
ID; Carnegie Museum of Natural History, Pittsburgh,
PA; Canadian National Collection of Insects, Ottawa,
ON; Colorado State University, Fort Collins, CO; Cor-
nell University, Ithaca, NY; Cambridge University
Museum of Zoology, Cambridge, UK; Deutsches En-
tomologisches Institut, Eberswalde, Germany; Uni-
versity of New Hampshire, Durham, NH; College of
Environmental Science & Forestry, Syracuse, NY;
Denver Museum of Natural History, Denver, CO; Es-
cuela Agricola Panamericana, Zamorano, Honduras;
Estacion de Biologia "Chamela", UNAM, San Patri-
cio, Mexico; North Carolina Department of Agricul-
ture; University of California, Berkeley, CA; Utah
State University, Logan, UT; University of Wyoming,
Laramie, WY; Funda^ao Instituto Oswaldo Cruz, Rio
de Janeiro, RJ, Brazil; Field Museum of Natural His-
tory, Chicago, IL; Florida State Collection of Arthro-
pods, Gainesville, FL; Instituto Miguel Lillo, San Mi-
guel de Tucuman, Tucuman, Argentina; Instituto Na-
cional de Biodiversidad, Santo Domingo, Costa Rica;
Illinois Natural History Survey, Urbana, IL; Instituto
Nacional de Pesquisas da Amazonia, Manaus, AM,
Brazil; University of Wisconsin, Madison, WI; Uni-
versidad Central de Venezuela, Maracay, Venezuela;
Kansas State University, Manhattan, KS; Natural His-
tory Museum, Los Angeles, CA; Loyola University,
Chicago, IL; M.A. Ivie private collection; Milwaukee
Public Museum, Milwaukee, WI; Museum of Com-
parative Zoology, Cambridge, MA; Mississippi State
University, Mississippi State, MS; Museum d'Histoire
Naturelle, Geneva, Switzerland; Universidad Nacio-
nal de La Plata, La Plata, Argentina; Museum Na-

tional d'Histoire Naturelle de Paris, France; Michigan
State University, East Lansing, MI; Montana State
University, Bozeman, MT; Museu de Zoologia da
Universidade de Sao Paulo, Sao Paulo, SP, Brazil;
North Carolina State University, Raleigh, NC; North
Dakota State University, Fargo, ND; Naturhistorisch-
es Museum, Vienna, Austria; Naturhistoriska Riks-
museet, Stockliolm, Sweden; New York State Muse-
um, Albany, NY; Oklahoma State University, Nor-
man, OK; Ohio State University, Columbus, OH; P.K.
Lago private collection; Peabody Museum of Natural
History, Yale University, New Haven, CT; Pennsyl-
vania State University, State College, PA; Purdue
University, West Lafayette, IN; Museu Nacional, Rio
de Janeiro, RJ, Brazil; Pontificia Universidad Catolica
del Ecuador, Quito, Ecuador; Nationaal Natuurhis-
torisch Museum, Leiden, The Netherlands; Royal On-
tario Museum, Toronto, Ontario, Canada; R.S. Miller
private collection; Rutgers State University, New
Brunswick, NJ; R. Willis Flowers collection; South
Dakota State University, Brookings, SD; Servicio En-
tomologico Autonomo, Nicaragua; University of
Kansas, Lawrence, KS; Southern Illinois University,
Carbondale, IL; Smithsonian Tropical Research Insti-
tute, Panama; Texas A&M University, College Sta-
tion, TX; T.K. Philips private collection; University of
Arkansas, Fayetteville, AK; University of Arizona,
Tucson, AZ; Universidad de Concepcion, Concep-
cion, Chile; University of California, Davis, CA; Uni-
versity of Colorado, Boulder, CO; University of Con-
necticut, Storrs, CT; University of California, River-
side, CA; University of Delaware, Newark, DE; Uni-
versity of Georgia, Athens, GA; University of
Louisville, Louisville, KY; University of Massachu-
setts, Amherst, MA; University of Mississippi, Ox-
ford, MS; Museum of Zoology, University of Michi-
gan, Ann Arbor, MI; University of Missouri, Colum-
bia, MO; University of Minnesota, St. Paul, MN;
Universidad Nacional Autonoma de Mexico, Mexico
City, Mexico; University of Nebraska State Museum,
Lincoln, NE; National Museum of Natural History,
Washington, DC; University of Vermont, Burlington,
VT; Virginia Tech University, Blacksburg, VA; Uni-
versity of Idaho, Moscow, ID; James Entomological
Collection, Washington State University, Pullman,
WA; West Virginia University, Morgantown, WV;
Humboldt Universitiit, Berlin, Germany; Zoologische
Staatssammlung, Munich, Germany.

J. HYM. RES.
Vol. 7(1), 1998, pp. 57-61

Parasitism of Siphotiinus phillyreae (Homoptera: Aleyrodidae) by
Aphelinid Parasitoids at Different Locations in Egypt

S. Abd-Rabou and M. M. Abou-Setta
Plant Protection Research Institute, Nadi El-Said Street, Dokki, Giza, 12618, Egypt

Abstract. — Seven species of aphelinid parasitoids (Hymenoptera: Aphelinidae) were reared from
second and third larval stages and pupae of pomegranate whiteflies, Siphoninus phillyreae (Hali-
day) (Homoptera; Aleyrodidae) from three locations in Egypt, during a one year survey (June 94-
June 95). Three species, Eretmocerus nniiuius Mercet, Encarsia davuii Viggiani and Mazzone, and £.
galilea Rivnay, were reported from Arish (Northeast Egypt). Parasitism was greatest (45.7%) in
Arish during September 1994, out of which 38% was caused by E. mundiis. Eiicarsia inaron (Walker)
was the dominant parasitoid of S. phillyreae in Giza (Central Egypt) and Assiut (Upper Egypt),
with average parasitism rates of 38 and 46.5% over the year, respectively. In Giza, total parasitism
reached a maximum of 80% during August 1994, with Encarsia inaron being responsible for 66.1%.
In Assiut, parasitism peaked at 93.1% in August 1994 with E. inaron accounting for 78% of the
total. Eretmocerus diversicilatus Silvestri and Encarsia lutea (Masi) were reported only from Giza.
Eretmocerus corni Haldeman was reported only from Assiut. These species had much less impact
than E. inaron in both locations. The presence of different parasitoids at different locations was
attributed to geographical factors as well as tolerance of the parasitoids to weather factors and
probably availability of alternative hosts. A key for the reported parasitoids is presented.

The pomegranate v^'hitefly, Siphoninus
piiillyreae (Haliday) (Homoptera: Aleyrod-
idae), is the most important pest of pome-
granate in Egypt (Priesner and Hosny
1932). This insect's host range is restricted
to deciduous fruit crops (Byrne et al.
1990). In Egypt, the host range of S. phil-
lyreae includes apple, Pyrus mains L.; pear,
Pyrus comnntnis L.; quince, Pyrus cydonia
L., and pomegranate, Puuica granatum L.
(Abd-Rabou 1990). Pomegranate orchards
extend from the north coast to southern
Egypt. Pomegranate leaves heavily infest-
ed with S. phillyreae have the demand for
fluid transport substantially increased be-
yond the tree's normal capacity to re-
spond. The loss of phloem fluids certainly
represents a loss of potential productivity
and probably contributes to the reported
reduction in fruit size (Costacos 1963). El-
wan (1982) studied the biology of S. phil-
lyreae and showed that its developmental
period varied according to temperature

and relative humidity. Various aphelinid
parasitoids (Hymenoptera: Aphelinidae)
have been reported from S. phillyreae, in-
cluding Encarsia inaron (Walker), £. siplion-
ini Silvestri, £. galilea Rivnay, £. hispida De
Santis, £. pseudopartenopea Viggiani and
Mazzone and Eretmocerus corni Haldeman
(Priesner and Hosny 1940; Mentzeloz 1967;
Viggiani and Mazzone 1980a,b; Viggiani
and Battaglia 1983; Rivnay and Gerling
1987; Polaszek et al. 1992). Puuica grana-
tum is a deciduous fruit crop which defo-
liates in winter.

The purpose of this study was to mon-
itor the rate of aphelinid parasitism on S.
phillyreae infested pomegranate leaves in
three distinctive regions in Egypt, to as-
sess the impact of biological control on the
whitefly, and to determine in what ways
it can be improved.

MATERIALS AND METHODS
Second and third larval stages and pu-
pae of Siphoninus phillyreae were sampled

58

Journal of Hymenoptera Research

Table 1. Percent parasitism of Siphoiiiinis pyhillyreae
by different aphelinid parasitoids in Arish, Egypt.

JJASONDJFMAMJ
1994 1995

Month

Fig. 1. Monthly mean maximum and minimum
temperatures and percent relative humidity at the
three locations over the survey period.

on pomegranate leaves collected monthly
(30 infested leaves per sample) from one
site representing each of three distinctive
regions in Egypt. The number of trees var-
ied by location. Arish is located in North-
east Sinai (i.e., coastal area), Giza is locat-
ed south of Nile Delta and Assiut in
southern Egypt. Environmental data were
obtained from local weather stations and
mean monthly values were plotted in Fig.
1. Pomegranate trees in the three locations
did not receive any chemical treatments
except at the Assiut area, which was
sprayed for stem borers during defoliation
(i.e., January-February 1995). Defoliation
time was longer in the Arish area than at
Giza or Assiut. No data for the whitefly is
presented during the defoliation periods.
Pomegranate leaves were transferred to
the laboratory in well-ventilated boxes. S.
phillyreae eggs and first larval stages were

Whitefly -
individuals/
leaf

Percent parasitism

Er.
miindiis

En.

En.

June 1994

July

August

September

October

November

December

January 1995

February

March

April

May

June

41.6
66.6
110.0
100.0
70.4
35.0
16.6

17.0
23.0
31
38

27

10

5

1.5

4.5

8

6.5

3

0.5

0

0

0.3

0.8

1.2

2.0

0.5

0

32.6

13

1.1

0.8

— Data was not available because of defoliation.

eliminated as well as other insects. Total
number of S. phillyreae individuals in each
stage were recorded per leaf. Each leaf
was stored in well-ventilated glass emer-
gence tube and monitored daily for para-
sitoid emergence. Parasitoid adults were
slide mounted in Hoyer's medium and
identified to species, and a diagnostic key
was constructed for their identification.

RESULTS

Parasitoid species emerging from sam-
ples of S. phillyreae on pomegranate varied
according to the area from which they
were collected in Egypt. In the Northeast-
ern Sinai, where Arish is located, Erctmo-
cerus miimius Mercet, Encarsia davidi Vig-
giani, and E. galilea parasitized an average
of 25% of the total S. phillyreae population.
Eretmocerus miindus was responsible for
20.5% of the total parasitism and the other
4.4% by the other two parasitoids (Table
1). In the Arish area, parasitism peaked at
45.7% in September (Table 1). The parasit-
oid species reported from Arish area (i.e.
£. mundus, E. davidi and E. galilea) were not
recovered from samples in Giza or Assiut.

In Giza (Central Egypt), S. phillyreae was
parasitized by £. iiiaroii, Eretinocents di-
versicilatus Silvestri, and £. liitea (Masi) at

Volume 7, Number 1, 1998

59

Table 2. Percent parasitism of Siiilwmnnf phiUyrecic
by different aphelinid parasitoids in Giza, Egypt.

Table 3. Percent parasitism of Siyhouiuui phillyrcae
by different aphelinid parasitoids in Assiut, Eg\'pt.

Whileflv

individu^ils/

leal

Percent parasitisir

1

Date

Whitefly
individuals/ —
leaf

Percent parasitism

En.
iiiiiron

Er
iluvn-

£.1

Fn imlrim

Fr conn

D.iti-

June 1994
July

70.1

50 0

9.0
12.1

June 1994

27.4

24.1

1.5

0.5

113.8

73.0

July

33.7

39.0

4.5

2.0

August

189.0

78.0

15.1

August

56.3

66.1

8.1

6.0

September

177.8

66.0

17.1

September

68.9

69.0

6.5

3.5

October

140.4

45.1

19.1

October

66.4

53.1

3.0

1.5

November

100.0

20.0

14.0

November

30.6

9.0

0.5

!.0

December

20.7

18.0

10.0

December

10.6

3.1

0.0

0.0

January 1995

—

—

—

January 1995

—

—

—

—

February

—

—

—

February

—

—

—

—

March

—

—

—

March

—

—

—

—

April

10.3

0.0

0.0

April

—

—

—

—

May

40.3

32.2

4.1

May

17.3

0

0

0

June

21

37.1

5.1

June

22.1

25.0

1.1

1.0

— Data was not

available beca

use of defoliation.

Data was not available because of defoliation.

average rates of 38, 4.5, and 2.2%, respec-
tively (Table 2). The maximum rate of par-
asitism reached 80% during August 1994,
of which £. itiaron was responsible for
66.1% of the total.

In Assiut, parasitism averaged 46.5% by
£. inaroti and 12% by Eretmocerus corui (Ta-
ble 3). Parasitism peaked to 93.1% during
August 1994, where £. inaron was respon-
sible for 78% of the total.

All reported parasitoids were primary
parasitoids except for Encarsia males that
are known to be hyperparsites on females
of their own species or other parasitoid
species (Viggiani 1981). No hyperparasi-
tism was observed.

DISCUSSION

The three areas of Egypt surveyed were
distinctive in their locations as well as
their weather (Fig. 1). The Arish area is
located in Northeast Sinai and can be
characterized by colder and longer win-
ters, and higher relative humidity than in
Giza or Assiut. Also, Arish is isolated
from the other two locations by the vast
desert area of the Sinai, which may ac-
count for difference in the S. pliillyreae par-
asitoid complex. The parasitoid species

collected from Arish area were not report-
ed in the other two locations. £. inaron was
the dominant parasitoid of S. phillyreae in
both Giza and Assiut. Both of these areas
are in the Nile River Valley, south of Nile
Delta with Assiut about 300 km south of
Giza. Encarsia lutea and £. diversicilatus
were obtained only in samples from Giza,
while £. corni was recovered only from
Assiut. Higher temperature in Assiut may
correlate to both higher whitefly popula-
tions and higher rates of parasitism.

Encarsia davidi was recorded by Abd-Ra-
bou (1994) as a parasitoid of Aleurolobus
niloticus Priesner and Hosny on Ziziphus
spinachristi in Egypt. Encarsia galilea was
recorded by Abd-Rabou (1994) as a para-
sitoid of S. phillyreae on P. granatum. Pries-
ner and Hosny (1932) recorded Encarsia in-
aron as a parasitoid of S. phillyreae on P.
granatum and indicated a rate of parasit-
ism as high as 80%. In our survey, the
peak of parasitism by £. inaron occurred
in September (69%) and in August (78%)
in Giza and Assiut, respectively.

Encarsia lutea (Masi) was recorded by
Abdel-Fattah et al. (1984) as a parasitoid
of Bettiisia tabaci (Genn.) on tomato plants,
whereas Eretmocerus corni was recorded
by Priesner and Hosny (1940) as a para-

60

Journal of Hymenoptera Research

sitoid of B. tahaci on Lantana camara. Eret-
mocerus diversicilatus was recorded for the
first time from Egypt associated with B.
tahaci by BChalifa and El-Khidir (1965), and
Eretmocerus mundus has been recorded
from Egypt on B. tahaci by El-Helay et al.
(1971). In the present study, £. mundus
was the dominant species in the Arish lo-
cation, with the peak parasitism occurring
in September (38%).

Parasitoid tolerance to different ranges

of temperature and relative humidity is
not clearly known. The colder and longer
winters of Arish, as well as lower humid-
ity in Assiut, may be limiting factors con-
trolling the presence and absence of each
parasitoid species of S. phillyreac. In addi-
tion, the availability of alternate hosts for
the parasitoids in the same area is proba-
bly another factor. Terminology used in
the key follows that of Polaszek et al.
(1992).

KEY TO APHELINID PARASITOIDS OF SIPHONINLIS PHILLYREAE FROM EGYPT

Antennal flagellum 3-segmented in female, club one elongate segment, tarsi 4- segmented,

male flagellum 1-segmented Genus Eretmocerus 2

Antennal flagellum 6-segmented in female, club 2-3 segments, tarsi S-segmented, male

flagellum 5 or 6-segmented Genus Encarsia 4

Mesoscutum with 2 pairs of setae, male pedicel dark brown 3

Mesoscutum with 3 pairs of setae, first funicle segment triangular, club 6-7 times as long

as wide, male pedicel yellow £. comi Haldeman

First funicle segment quadrate, second funicle segment longer than wide, submarginal

vein with 3 seta E. mundus Mercet

First funicle segment triangular, second funicle segment transverse, submarginal vein

with 2 setae E. diversicilatus Silvestri

Head and mesosoma dark brown to black with 6-10 pairs of setae, metasoma yellow,

antennal club 2-segmented E. inaron (Walker)

Entire body yellow or orange, or nearly so, metasoma tergite I often infuscate, antennal

club 3 segmented 5

Valvular III dark brown, first funicle segment quadrate or wider than long, male funicle

segments F1-F3 expanded 6

Valvular III yellow, first funicle cylindrical, 1.6-2.3 as long as wide, male unknown ....

£. galilea Rivnay

Valvular III short, as long as width at base and 0.3 times as long as ovipositor, tibia II

1.0-1.1 times as long as ovipositor E. lutea (Masi)

Valvular III elongate, 2 times as long as width at base and 0.4 times as long as ovipositor,
tibia II 0.8 times as long as ovipositor E. davidi Viggiani and Mazzone

LITERATURE CITED

Abd-Rabou, S. 1990. Taxonomic studies of whiteflies
of Egypt (Homoptera: Aleyrodidae). M.Sc. Tlwsis,
Fac. of Sciena; Aiii SImins Univ., 193 pp.

Abd-Rabou, S. 1994. Taxonomic and biological stud-
ies on the parasites of whiteflies (Hemiptera: AI-
eyrodidae) in Egypt. Ph.D. Thesis, Fac. of Science,
Cairo Univ., 83 pp.

Abdel-Fattah, M. I., A. Hendi, M. O. Koliab and A.
El-Said. 1984. Studies on Prosfmllelln lulca Masi
(Hymenoptera: Aphelinidae), a primary parasite
of the cotton whitetly, Bcniisin tabnci (Genn.) in

Egypt (Hemiptera: Aleyrodidae). Bulletin ile la So-
ciete Entoinolo\;upie ti'Egypte 65: 119-129.
Byrne, D. N., T. S. Bellows and M. P. Parrella. 1990.
Whiteflies in Agricultural Systems, 227-261, in:
Gerling, D. (ed.) "Whiteflies: their Bionomics,
Pest Status and Management." Intercept Ltd., 348

PP-

Costacos, T. A. 1963. On a severe attack bv Siphomiiiis
phiUyreae (Hal.) (Hemiptera: Aleyrodidae) sub-
species inneqiinlis Gautier on fruit trees and its
control. Geoponika 105; 3-7.

El-Hclay, M. S., A. Y. El-Shazli and F. H. El-Gayar.
1971. Biological studies on Beniisia tnlnici (Genn.)

Volume 7, Number 1, 1998

61

(Hemiptera: Aleyrodidae) in Egypt. Zeitschrifl
feur Angewnndtc Entomologie 69(1): 48-55.

Elwan, E. A. 1982. Biological and ecological studies
on the pomegranate whitefly, Siphoiiiniis phiUy-
rene (Hal.) (Hemiptera: Aleyrodidae). Mij.';/<t cf
Science Thesis, Faculty of Agriculture, Cniro Uni-
versity, 1-97.

Khalifa, A. and E. El-Khidir. 1965. Biological study
on Trialeurodes lubia El-Khidir and Khalifa and
Bemisia tabaci (Germ.) (Hemiptera: Aleyrodidae).
Bulletin lie In Societe Entonwiogique d'Egypte 18:
120-155.

Mentzeloz, I. A. 1967. Contribution to the study of
the entomophagous insects on Siphoninus pliilly-
reae (Hal.) (Hemiptera: Aleyrodidae) on pear
trees in central Macedonia. Report of the Plant
Protection Agriculture Research Station. Thessa-
loniki 3: 92-102.

Polaszek, A., G. A. Evans and F. D. Bennett. 1992.
Encarsia parasitoids of Bemisia tabaci (Hymenop-
tera: Aphelinidae, Homoptera: Aleyrodidae): a
preliminary guide to identification. Bulletin ofEn-
tmnological Research 82: 375-392.

Priesner, H. and M. Hosny. 1932. Contributions to a
knowledge of whiteflies of Egypt. Bidletin Min-
istry of Agriculture, Egypt, 121, 8 pp.

Priesner, H. and M. Hosny. 1940. Notes on parasites
and predators of Coccidae and Aleyrodidae in
Egypt. Bulletin de la Societe Ento>nologiqiie d'Egi/pte
24: 58-70.

Rivnay, T. and D. Gerling. 1987. Aphelinidae para-
sitoids (Hymenoptera: Chalcidoidae) of white-
flies (Hemiptera: Aleyrodidae) in Israel with de-
scription of 3 new species. Entonwp'haga 32: 463-
475.

Viggiani, G. 1981. The role of hyperparasitism on bi-
ological control: A symposium, pp. 19-26. Uni-
versity of California, Division of Agricultural Sci-
ences.

Viggiani, G. and Battaglia, D. 1983. Le specie italian
del genera Eretnwcerus Haldeman (Hymenop-
tera: Aphelinidae). Bollettino del Laboratorio di En-
tcnwlogia Agraria Fdippo Sdvestri 40: 97-101.

Viggiani, G. and P. Mazzone. 1980a. Encarsia ptseudo-
parlenopea n. sp., parasita di Siphoninus phillyreae
(Haliday) (Homoptera: Aleyrodidae). Bollettnw
del Laboratorio di Entomologia Agraria Fdippio Sil-
vestri 37: 9-12.

Viggiani, G. and P. Mazzone. 1980b. Le specie Pa-
leartiche di Encarsia del gruppe Lutea (Masi) (Hy-
menoptera: Aphelinidae), con descrizione de due
nuove specie. Bollettino del Laboratotio di Entomo-
logia Agraria Fdippo Silvestrii 37: 51-57.

J. HYM. RES.
Vol. 7(1), 1998, pp. 62-73

Revision of North American Aleiodes Wesmael (Part 2):

the apicalis (Brulle) Species-group in the New World

(Hymenoptera: Braconidae, Rogadinae)

Scott R. Shaw, Paul M. Marsh, and Joseph C. Fortier

(SRS, JCF) Department of Plant, Soil, and Insect Science, P.O. Box 3354, University of Wyoming,

Laramie, Wyoming 82071, USA; (PMM) Cooperating Scientist, USDA Systematic Entomology

Laboratory, c/o U.S. National Museum of Natural History, NHB-168, Washington, D.C. 20560, USA

(correspondence address; P.O. Box 384, North Newton, Kansas 67117, USA)

Abstract. — The Aleiodes apicalis (Brulle) species-group is defined to include the following previ-
ously described species: apicalis (Brulle, 1832), grandis Giraud, 1857 { = Rhogas malaisei Shestakov,
1940 NEW SYNONYMY), parasiticus Norton, 1869, atriceps Cresson, 1869, abdorninalis Cresson,
1869, rileyi Cresson, 1869, molestus (Cresson, 1872) NEW COMBINATION, schirjajewi Kokujev,
1898, convexiis van Achterberg, 1991 ( = Chelonorhogas rufithorax Enderlein, 1912), and brethesi
Shenefelt, 1975 NEW COMBINATION. One newly described species, flavitarsiis Marsh and
Shaw, is also included. The apicalis species-group is regarded as monophyletic based on the pres-
ence of dense setal mats on the apical metasomal terga of males. The genus Dimorphomastax
Shenefelt, 1967 is synonymized under Aleiodes, and the species Dimorphomastax peculiaris Shenefelt,
1979 is regarded as a junior synonym of Aleiodes atriceps Cresson. Rogas rufocoxalis Gahan, 1917 is
newly synonymized as a junior synonym of Aleiodes molestus (Cresson). A key to the New World
species of the apicalis species-group is provided, and species treatments are given for Nearctic
species, including diagnostic characters, distribution, and biological information.

The rogadine braconid genus Aleiodes
Wesmael is worldwide in distribution, but
is particularly species-rich in the Holarctic
region. Aleiodes is diverse in North Amer-
ica, with at least 90 species in the United
States and Canada (Shaw et al. 1997). This
paper is the second contribution in a series
of planned papers on Aleiodes species-
groups, intended to provide a complete
revision of the genus for North America.
In this paper we treat a distinctive mono-
phyletic assemblage, the species of the Al-
eiodes apicalis species-group, as it occurs in
the New World region. All members of
this group have dense setal mats on male
terga 4-7 (Fig. 1). Our definition of the
species-group includes all species known
to us, worldwide. However, because our
main intent is to provide a revision for
North American species, species treat-
ments are limited to the Nearctic fauna.

Since only one other New World species
is known to us, this is included in the key
for convenience. The European species are
currently being revised by Kees van Ach-
terberg and Mark Shaw.

Aleiodes species are koinobiont endopar-
asitoids of lepidopteran larvae, especially
macrolepidoptera of the superfamilies
Noctuoidea and Geometroidea, and to a
lesser extent, Arctioidea, Sphingoidea, and
Papilionoidea (Shaw et al. 1997). Members
of the apicalis group, as far as known, are
mostly parasitoids of Noctuidae. The
method of parasitism, unique to the tribe
Rogadini, is noteworthy: the Aleiodes larva
completes its feeding and pupates within
the shrunken and mummified remains of
the host caterpillar. The form of the mum-
my caused by a particular Aleiodes species
is usually characteristic for that host and
parasitoid, so the mummified remains are

Volume 7, Number 1, 1998

63

of considerable diagnostic value and
should be retained with the parasitoid,
when reared. For more complete discus-
sions of Aleiodes biology, readers may re-
fer to Shaw (1983, 1994), Shaw and Hud-
dleston (1991), Shaw (1995), and Shaw et
al. (1997).

METHODS

Species covered in this paper can be
identified as members of the subfamily
Rogadinae using the keys of Shaw and
Huddleston (1991), van Achterberg (1993),
or Shaw (1995). Our definition of Aleiodes
follows that of van Achterberg (1991),
Shaw (1993), and Shaw et al. (1997). Spec-
imens can be determined as Aleiodes using
the keys of Marsh et al. (1987), van Ach-
terberg (1991), or Shaw (1997). Specimens
keyed through Marsh et al. (1987) will key
to couplet 185, at which point they can be
separated from Rogas by the presence of a
discrete median carina on the propodeum,
the lack of a foveate sternaulus on the
mesopleuron, and the lack of a blunt basal
tooth on the tarsal claw. In practice, more
than 99% of U.S. and Canadian specimens
encountered will be Aleiodes, as Rogas s.s.
is only infrequently encountered north of
Mexico (but increases in species richness
in the neotropics). The species-groups of
North American Aleiodes can be keyed us-
ing the key provided in Shaw et al. (1997).
The species treated in this paper were for-
merly assigned to the diictor Thunberg
species-group by Shaw et al. (1997) follow-
ing a recent interpretation of that species
by Papp (1985). However, Kees van Ach-
terberg (pers. comm.) has indicated to us
that previous interpretations of ductor are
not correct, and that the species treated
here are better called the apicalis species-
group.

Terminology follows that used for Aleio-
des by Shaw et al. (1997), Shaw (1993) and
Marsh (1989). Microsculpture terminology
follows that of Harris (1979). Wing vena-
tion terminology (see Fig. 16) follows that
of Shaw (1997) and Shaw et al. (1997).

Abbreviations for museums are as fol-
lows: ANSP, Academy of Natural Sci-
ences, Philadelphia; AEI, American Ento-
mological Institute, Gainesville; AMNH,
American Museum of Natural History,
New York; CAS, California Academy of
Sciences, San Francisco; CNC, Canadian
National Collection, Ottawa; CUI, Cornell
University, Ithaca; FSCA, Florida State
Collection of Arthropods, Gainesville;
HNHM, Hungarian Natural History Mu-
seum, Budapest; INHS, Illinois Natural
History Survey, Urbana; MISU, Michigan
State University, East Lansing; MSSU,
Mississippi State University, Mississippi
State; OKSU, Oklahoma State University;
RMNH, Nationaal Natuurhistorisch Mu-
seum, Leiden; TAMU, Texas A. & M. Uni-
versity, College Station; UCD, University
of California, Davis; UMSP, University of
Minnesota, St. Paul; RMSEL, Rocky
Mountain Systematic Entomology Labo-
ratory, University of Wyoming, Laramie;
USNM, U.S. National Museum of Natural
History, Washington, D.C.

Authorship of species is attributed to
the senior authors (PMM and SRS) in the
order indicated.

ALEIODES APICALIS SPECIES-GROUP

Included species: apicalis (BruUe, 1832),
grandis Giraud, 1857 ( = »;fl/rt(St'/ Shestakov,
1940 new synonymy), parasiticus Norton,
1869, atriceps Cresson, 1869 revised com-
bination { = Diiuorphoi)iastax peculiaris
Shenefelt, 1979 new synonymy), abdomi-
nalis Cresson, 1869 ( = lectus Cresson, 1869),
rilei/i Cresson, 1869 revised combination,
niolestus (Cresson, 1872) new combination
( = rufocoxalis (Gahan, 1917) new synony-
my), schirjajezvi Kokujev, 1898, coiwexus
van Achterberg, 1991 ( = Chelonorliogas ruf-
ithorax Enderlein, 1912), hrethesi Shenefelt,
1975 new combination (replacement name
for nigriceps Brethes, 1909, preoccupied by
iiigriceps Wesmael, 1838), and flavitarsus
Marsh and Shaw, new species.

Diagnostic characters. — Ocellar diameter
small, ratio of ocellar diameter to distance

64

Journal of Hymenoptera Research

Figs. 1-6. Fig. 1. Densely setose metasomal tergum 4. A. alniomiiiaUs, male. Figs. 2-3. Mandible and oral
space. 2. A. atriceps, male. 3. A. abdominalis, male. Figs. 4-6. Mesonotal sculpture. 4. A. abdominalis. 5. A.
parasiticus. 6. A. hrcthesi.

between lateral ocellus and compound eye
less than 1; occipital carina meeting hy-
postomal carina laterally; dense setal mat
present on male terga 4-7 and subdivided
medially (Fig. 1); medial ridge extending
down frons 0.55 or more of distance from
line between base of scape to clypeus;
mesonotal disc sculpture finely granulate
to smooth (Figs. 4-6), sparsely or not se-
tose; tarsal claws strongly pectinate (Figs.
12-14).

Remarks. — A moderate-sized, circumpo-
lar and neotropical monophyletic group,
associated mostly with noctuids (see Fig.
15). There are some recorded associations
with geometrids, lymantriids, pyralids,
and even sawflies, but these need confir-
mation and the latter seems unlikely.

The dense setal mats on male terga 4-7
are undoubtedly synapomorphic. The
function of the dense setal mats on male
terga 4—7 is unknown, but perhaps they

Volume 7, Number 1, 1998 65

may serve to disperse pheromones during {Aleiodes, Cheloiwrhogas, Diniorphojnastax,

courtship and mating. and Ro;^as). Chiionorliogas was synonymized

Even fairly recently (Shenefelt, 1975; with Aleiodes by van Achterberg (1991), but

Marsh, 1979), species belonging in this retained Cheloiwrhogas as a valid subgenus,

group have been classified in several genera to which the apicalis-group is assigned.

KEY TO NEW WORLD SPECIES OF THE ALEIODES APICALIS SPECIES-GROUP

1 Mandible of male with a large, curved, tusk-like accessory tooth situated near the pos-
terior condyle (Fig. 2), or female with a small accessory tooth situated near the posterior
condyle; oral space unusually large; southwestern U.S. and Mexico . , A. atriceps Cresson

- Mandible normal and unmodified, lacking a large, tusk-like tooth (males) or small tooth
(females) near the posterior condyle (Fig. 3); oral space smaller (Fig. 3) 2

2(1) Mesosoma (excluding legs) entirely black 3

- Mesosoma color varying from entirely orange, to orange with extensive black markings
posteriorly and laterally, but pronotum, mesonotum, and scutellar disc always colored
bright orange 4

3(1) Hind tarsomeres orange to brown, similar to color of tibia; body length 6.0-8.0 mm;
fore wing vein Icu-a beyond vein IM by less than 3 times its length; hindwing vein m-
cu present (Fig. 16) A. abdominalis Cresson

- Hind tarsomeres 1-4 yellow, much lighter than color of tibia; body length 4.5-5.5 mm;
fore wing vein Icu-a beyond vein IM by 3 times its length; hindwing vein m-cu absent
(Fig. 17) A. flavitarsiis Marsh & Shaw, new species

4(3) Body entirely orange to orangish brown; mesonotum granulate and dull

A. rileyi Cresson

- Body only partly orange, head entirely black, legs, mesosoma, and apex of metasoma
with variable black markings; mesosoma sculpture variable, but always somewhat shin-
ing and polished 5

5(4) Mesosoma entirely orange, legs entirely black; mesonotal disc entirely smooth and high-
ly polished, virtually devoid of setae (Fig. 6); South American species

A. brethesi (Shenefelt)

- Mesosoma usually orange and black (rarely entirely orange), leg color orange and black,
coxae at least always orange; mesonotal disc not so completely smooth, with numerous
setal pits (Fig. 5); North American species 6

6(5) Hind femur and tibia banded, orange on basal half, black on apical half; northern species

associated with boreal forests A. parasiticus Norton

- Hind femur and tibia entirely black; southern species frequently associated with agroe-
cosytems A. molestus (Cresson)

Aleiodes atriceps Cresson, revised palpi yellow, mesosoma varying from en-
combination tirely black to entirely orange, tegula yel-
(Fig. 2) low, metasoma orange, legs orange, wings

hyaline, veins brown except vein C + Sc + R

Aleiodes atriceps Cresson, 1869, Amer. Ent. Soc. ^^^ ^^-^^^ yellow; body length, 6.0-7.0

Trans. 2:380. .,.„-,,.„ mm; 46-53 antennomeres; malar space

Dn«orptom«stop.c«/mns Shenefelt, 1979, Proc. ^^ ^^^^, ^^^^^^ ^^ mandible; oral

Ent. Soc. Wash. 81:133. New synonymy " . .... ^ . i i

, , .,. c\. < ^ openme circular, diameter equal to malar

based on exammation of holotype. ^ P , , , , ., .

space m female and greater than malar

Diagnosis. — Body bicolored, head in- space in male; ocelli small, ocellocular dis-

cluding antenna black, mandibles and tance about twice diameter of lateral ocel-

66

Journal of Hymenoptera Research

lus; mandible in female with short trian-
gular tooth at condyle, in male with large
sickle-shaped tooth at condyle; occipital
carina meeting hypostomal carina; head
entirely coriaceous, sometimes smoother
in male; pronotum rugose laterally; meso-
notum and scutellum coriaceous, notauli
weakly scrobiculate; mesopleuron smooth,
subalar sulcus rugose, sternaulus absent;
propodeum rugose, median carina com-
plete; metasomal terga 1 and 2 costate-ru-
gose, median carina complete; third meta-
somal tergum longitudinally costate over
basal 0.25 to 0.5, smooth posteriorly; terga
4—7 in male with dense row of short yel-
low setae at base, and dense patches of
yellow setae on each side of mid-line; fore
wing with vein Icu-a beyond vein IM by
distance nearly twice length of Icu-a, hind
wing with marginal cell gradually broad-
ening to apex, vein RS straight on basal
half and slightly bent downward on apical
half, vein m-cu present; tarsal claws
strongly pectinate with 6-7 stout teeth,
apical 3 teeth of pectin about 0.75 as long
as apical claw, remaining teeth gradually
shorter towards base of claw; apical tibial
spurs of male blunt.

Type material examined. — Aleiodes atriceps
Cresson, holotype female, Mexico (ANSP).
Diniorphomastax peculiaris Shenefelt, para-
type female. Portal, Arizona (AEI).

Distribution. — Mexico, Arizona, and
Texas.

Biology. — Unknown. Adults have been
collected from July through December.

Conwients. — Shenefelt (1975) classified
atriceps under Rogas, but we are moving it
here back to its original combination with
Aleiodes. Although the unusual and dis-
tinctive shape of the mandible (Fig. 2)
prompted Shenefelt (1979) to create a new
genus for this species, we consider it to be
only a specialized species of Aleiodes with
peculiar mandibles. The species is, in most
other characters, a typical Aleiodes and we
consider the mandible shape, as well as
the blunt tibial spurs of the male, to be
autapomorphies. Dimorphomastax Shene-

felt, 1969 is therefore considered a junior
synonym of Aleiodes, new synonymy.

The unusual accessory mandibular
tooth of this species is strongly sexually
dimorphic (small triangular tooth in the
female, large sickle-shaped projection in
the male) suggesting a possible role in
courtship and mating. The sexual dimor-
phism raises doubts about whether it may
serve any primary cutting function, such
as assisting in escape from the host m.um-
my.

Aleiodes abdominalis Cresson

(Figs. 1, 3, 4, 7, 12, 16)

Aleiodes abdominalis Cresson, 1869, Amer. Ent.
Soc. Trans. 2:379.

Aleiodes lectus Cresson, 1869, Amer. Ent. Soc.
Trans. 2:379. Possible synonymy with Aleio-
des abdominalis Cresson indicated by Cresson,
1869. Synonymized by Muesebeck & Walk-
ley, 1951.

Diagnosis. — Body bicolored, head and
mesosoma black, metasomal terga 1-3 or-
ange, remainder of terga orange to black,
antenna and legs orange, wings lightly
dusky, veins brown, tegula yellow; body
length, 6.0-8.0 mm; 54-63 antemiomeres;
malar space longer than basal width of
mandible; face costate with distinct raised
ridge between antennae, frons, vertex and
temple coriaceous; oral opening circular,
diameter about equal to basal width of
mandible; propleuron rugose; mesonotum
and scutellum finely coriaceous, notauli
weakly scrobiculate and meeting in small
rugose are before scutellum; mesopleuron
smooth, subalar sulcus rugose, sternaulus
absent; propodeum rugose-coriaceous,
median carina on basal half only; first and
second metasomal terga costate, median
carina complete, third tergum costate at
base, remainder smooth; apical terga of
males, especially terga 4-7 densely cov-
ered with silvery setae, except along me-
dian line; fore wing with vein Icu-a wing
beyond vein IM by distance greater than
length of Icu-a; hind wing with marginal
cell gradually widening, vein RS slightly

Volume 7, Numbkk 1, 1998

67

s

^i

y^Bfli^^B^ "^

Figs. 7-12. Figs. 7-8. Propodeal sculpture. 7. A. ahdominalis. 8. A. parasitictis. Figs. 9-10. Metasomal sculpture.
9. A. brethesi, first tergum. 10. A. brethesi, border of second and third metasomal terga. Figs. 11-12. Tarsal
claws. \\. A. parasiticus. 12. A. abdominalis.

curved downward, vein m-cu present; tar-
sal claws strongly pectinate with 6-7 stout
teeth, apical 3 teeth of pectin about 0.75 as
long as apical claw, remaining teeth grad-
ually shorter towards base of claw.

Type material examined. — Aleiodes abdom-
inalis Cresson, holotype female, Pennsyl-
vania (ANSP). Aleiodes lectiis Cresson, ho-
lotype male, Illinois (ANSP).

Distribution. — Widely distributed in

eastern North America from Quebec and
Ontario south to North Carolina, west to
South Dakota and Arizona; more com-
monly encountered in the eastern parts of
its range.

Biologi/. — Unknown. One specimen
from Maryland is associated with an un-
determined noctuid. The mummy is dark
brown, smooth, and about 1 cm long.
Adults appear in early June in the north-

68

Journal of Hymenoptera Research

m

Figs. 13-15. Figs. 13-14. Compound microscope photographs of slide-mounted tarsal claws. 13. A. hkyi. 14.
A. niclcstiis. Fig. 15. Mummified plusiine noctuid host larva parasitized by A. tnolestus.

ern parts of its range; the specimens from
Arizona were collected in late September.
Comments. — This is the most common
member of the species-group with a most-
ly black body. The only other North
American species in the group with an all-
black mesosoma is flavitarsus, from which
abdominalis can be distinguished by its
larger body size (6.0-8.0 mm), fore wing
vein Icu-a beyond vein IM by less than 3
times its length, and hindwing vein m-cu
present (Fig. 16). The European species, A.
apicalis, is also similar in body color and
general appearance. However, abdominalis
can be distinguished by its finely coria-
ceous to granulate mesonotum (Fig. 4),
while the disc of the mesonotum is
smooth-punctate in apicalis. Superficially,
abdominalis is similar in color pattern
(black and orange) to the very common
species A. terminalis Cresson, but terminalis
is a member of a different species-group,
and can be easily separated by the species-
group key provided in Shaw et al. (1997).
Males of terminalis do not have densely se-
tose metasomal terga 4-7.

Aleiodes flavitarsus Marsh and Shaw,
new species

(Fig. 17)
Female. — Body color: head black, man-
dibles and mouthparts yellow, antenna

light brown; mesosoma black, propleuron
sometimes brown or orange; tegula yel-
low; legs yellow except apical tarsal seg-
ments, apical 0.25 of hind femur and api-
cal 0.5 of hind tibia which are black; meta-
soma with segments 1-3 yellow-orange
except tergum 1 black medially and ter-
gum 3 black apically, segments 4-8 black;
wings hyaline, veins light brown, tegula
yellow. Body length, 4.5-5.5mm. Head:
malar space short, equal to basal width of
mandible and about eye height; temple
narrow, about 0.5 eye width; occipital ca-
rina not quite meeting hypostomal carina;
oral space small and oval, width equal to
malar space and about 0.5 face height; 46-
49 antennomeres, all flagellomeres slightly
longer than wide, first slightly longer than
second; ocelli small, ocellocular distance
equal to or slightly greater than diameter
of lateral ocellus; face rugulose-coriaceous
with median carina between antennae;
frons coriaceous; vertex and temples cori-
aceous; occiput smooth and shining; max-
illary palpus not swollen; mandibles
small, tips not overlapping when closed.
Mesosoma: propleuron rugose, often
smooth medially; mesonotum and scutel-
lum coriaceous; notauli scrobiculate, meet-
ing in small rugose area before scutellum;
mesopleuron smooth and shining, rugose

Volume 7, Number 1, 1998

69

Figs. 16-17. Wings showing venation terminology. 16. A. abdominaU^. 17. A. flnvitarfus.

dorsally and in subalar sulcus; sternaulus
absent; propodeum rugose-coriaceous
dorsally, coriaceous laterally, median ca-
rina obscured apically. Legs: tarsal claws
strongly pectinate with 4-5 stout teeth,

apical 3 teeth of pectin about 0.75 as long
as apical claw, remaining teeth gradually
shorter towards base of claw; inner spur
of hind tibia slightly less than 0.5 length
of hind basitarsus; hind coxa smooth dor-

70

Journal of Hymenoptera Research

sally. Wings: hyaline; fore wing with vein
r 0.5 length of 3RSa, vein Icu-a beyond IM
by nearly 3 times length of 1 cu-a, vein
ICUa slightly longer than ICUb; hind
wing with vein RS straight, cell IRl grad-
ually widening to wing apex, vein r-m
slightly shorter than IM, vein M + Cu
slightly longer than IM, vein m-cu absent.
Metasoma: first tergum rugulostriate, me-
dian carina complete, length equal to api-
cal width; second tergum rugulostriate,
median carina complete; third tergum ru-
gulostriate basally, smooth apically, me-
dian carina absent; fourth and following
terga smooth; ovipositor short, about 0.5
length of hind basitarsus.

Male. — Essentially as in female.

Holotype.— Female: MICHIGAN: Mar-
quette County, August 14, 1959, R. And K.
Dreisbach. Deposited in USNM.

Parnti/pes.— CANADA, BRITISH CO-
LUMBIA: 2 females, Gagnon Rd., 6 mi W
Terrace, June 20, 1960, J. C. Chillcott, W.
W. Moss; 4 females, Hixon, July 11, 1965-
July 7, 1966, E. D. A. Dyer; 1 female. Ter-
race, July 9, 1960, W. R. Richards; 1 fe-
male, 10 km S Fernie, July 29, 1980, D.
Williams. MANITOBA: 2 females. Big
Eddy, em March 13, 1953, ex Aiitographn
sp. NEW BRUNSWICK: 2 males, Kouchi-
bougnac N.P., July 21, 1977, S. J. Miller.
NEWFOUNDLAND: 1 male, Agr. Exp.
Sta., St. John's, July 16, 1967, J. F. Mc-
Alpine. ONTARIO: 1 female. One Sided
Lake, July 12, 1960, S. M. Clark; 1 male.
North Branch, July 23, 1960, S. M. Clark;
1 male, Cumberland, June 13, 1975, L.
Ling. QUEBEC: 1 female, 1 male, Parke
Reserve, Kam. Co., July 5-12, 1957, G. E.
Shewell. UNITED STATES, MAINE: 1 fe-
male, Oxford Co., Bryand Pond, July 23,
1976, leg. Heinrich. MICHIGAN: 1 female.
Delta Co., August 6, 1959, R. & K. Dreis-
bach; 1 male, Houghton Co., August 20,
1959, R. & K. Dreisbach; 1 male, School-
kraft Co., August 5, 1959, R. & K. Dreis-
bach. MINNESOTA: 1 female, Itasca St.
Park, September 1927, S. Carthside.
WASHINGTON: 1 female. Lake Cush-

man, July 22, 1917, A.L. Melander. Depos-
ited in CNC, HNHM, MCZ, RMNH,
RMSEL, USNM.

Distribution. — Widely distributed across
Canada and the northern United States.

Biology. — Two specimens from Manito-
ba were reared from an unknown species
of the genus Autographa (Noctuidae).
Adults are active from late June through
September. The specimens from Manitoba
were labeled as emerging in March, but
were probably collected during the pre-
vious summer and emergence may have
been under laboratory conditions.

Comments. — This species is similar to ah-
dominalis, but differs by its smaller body
size (4.5-5.5 mm), vein Icu-a of the fore
wing being beyond vein IM by three
times its length (Fig. 17), by having hind
tarsomeres 1^ yellow, and by the absence
of vein m-cu in the hind wing (Fig. 17).

Etymology. — The specific name is from
the Latin flaviis meaning yellow, in refer-
ence to the yellow hind tarsus.

Aleiodes molestus (Cresson), new
combination

(Figs. 14, 15)

Rogas molestus Cresson, 1872, Trans. Amer. Ent.
Soc. 4:188.

Rogas rufocoxalis Gahan, 1917, Proc. U.S. Natl.
Mus. 53:207. New synonymy based on ex-
amination of holotype.

Diagnosis. — Body bicolored, mesonotum
orange, mesopleuron and propodeum
varying from entirely black to entirely or-
ange, with various intermediate forms oc-
curring, metasomal terga 1-3 always or-
ange, rest of terga varying from orange to
black, legs beyond coxae dark brown or
black, wings slightly dusky, veins brown,
tegula orange; body length, 5.0-7.0 mm;
45-47 antennomeres; malar space long,
slightly greater than basal width of man-
dible; face, frons and vertex rugulose, tem-
ple smooth; mesonotum and mesopleuron
smooth; propodeum rugose, median cari-
na complete; first and second metasomal
terga strigate-rugose to costate, median

Volume 7, Number 1, 1998

71

carina complete; fore wing with vein
Icu-a beyond IM by distance nearly twice
length of Icu-a; marginal cell of hind wing
narrowest basally, gradually widening to-
ward wing apex; tarsal claws strongly
pectinate with 6-8 stout teeth, apical 3
teeth of pectin about 0.75 as long as apical
claw, remaining teeth gradually shorter
towards base of claw.

Type material examined. — Rogas molestus
Cresson, holotype female, Texas, G.W.
Belfrage collection, [USNM]. Rogas rufo-
coxalis Gahan, holotype female, Colorado,
Rocky Ford [USNM].

Distribution. — South Dakota south to
Arkansas, Louisiana, Texas, and Mexico,
west to Wyoming, Utah, Arizona, and
southern California.

Biology. — Although A. molestus is com-
monly collected by Malaise trap, sweep
net, or at lights, verified rearing records
are less common. In the USNM collection
there are single specimens labelled as
reared from several plusiine and noctuine
species including Autoplusia egena (Gn.),
the soybean looper, Pseudoplusia includens
(Wlkr.), the cabbage looper, Trichoplusia ni
(Hbn.), and the variegated cutworm, Per-
idroma saucia (Hbn.). The known hosts are
all generalist feeders on a variety of low
vegetation including numerous crop spe-
cies. Adults have been collected from late
April through mid-September in the
southern parts of its range.

Comments. — A. molestus is a fairly com-
mon midwestern and southern species fa-
voring open fields, low vegetation, and
agroecosystems. It belongs to the parasiti-
cus assemblage, comprising parasiticus
Norton, molestus (Cresson) and brethesi
(Shenefelt), all of which have an orange
mesonotum that is smooth and shining (as
in Figs. 5-6) and well-developed longitu-
dinal sculpture on tergum 2 -(-3 (as in Fig.
10). A. molestus differs from parasiticus by
having the legs entirely dark brown or
black beyond the coxae (the femorae and
tibiae are banded in parasiticus). It differs
from brethesi by having orange coxae.

while the legs are entirely black in that
South American species.

There is considerable variation in the
extent of dark coloration on the mesopleu-
ron and propodeum in molestus, conse-
quently rufocoxalis, which differs only by
having an entirely orange mesosoma, can-
not be held as a valid species. In his de-
scription of rufocoxalis, Gahan (1917) stat-
ed that "it would not be surprising if it
{rufocoxalis) would ultimately turn out to
be merely a color variety of Cresson's spe-
cies (molestus)." Indeed, although some-
times the mesopleuron and propodeum
are entirely black, intermediates with less
dark color and orange patches showing
through are fairly common, so it would
appear that Gahan's prediction is correct.

Aleiodes parasiticus Norton

(Figs. 5, 8, 12)

Aleiodes parasiticus Norton, 1869, Trans. Amer.
Ent. See. 2:327.

Diagnosis. — Body bicolored black and
orange, head black, antenna orange ba-
sally to black apically, mesosoma orange
except mesopleuron below subalar sulcus
and propodeum black, first and second
metasomal terga orange, third tergum or-
ange on basal 0.5, black on apical 0.5, re-
mainder of terga black, fore and middle
legs orange except apical tarsomeres
black, hind coxa and trochanters orange,
hind femur orange on basal 0.66, black on
apical 0.33, hind femur yellow on basal
0.5, black on apical 0.5, hind tarsus or-
ange except apical tarsomere black,
wings lightly dusky, veins brown, tegula
yellow; body length, 5.0-7.0 mm; face
costate, frons and vertex rugose, temple
punctate, occipital carina scrobiculate;
malar space longer than basal width of
mandible and about 0.5 eye height; ocel-
locular distance longer than diameter of
lateral ocellus; pronotum rugose; meso-
notum smooth, mesopleuron smooth me-
dially, subalar sulcus and sternaulus ru-
gose; propodeum rugose dorsally, punc-

72

Journal of Hymenoptera Research

tate laterally, median carina complete;
first and second metasomal terga costate-
rugose, median carina complete, third
tergum costate on basal 0.5, smooth on
apical 0.5, median carina absent, remain-
der of terga smooth, fourth-seventh terga
of male with dense patches of setae on
apical 0.5 on each side of mid-line; fore
wing with vein Icu-a beyond IM by dis-
tance twice length of Icu-a, marginal cell
of hind wing gradually widening, vein
RS straight, vein m-cu absent; tarsal
claws strongly pectinate with 5-6 stout
teeth, apical 3 teeth of pectin about 0.75
as long as apical claw, remaining teeth
gradually shorter towards base of claw.

Type material examined. — Aleiodes parasi-
ticus Norton, holotype female, Connecticut
(MCZ).

Distribution. — Eastern Canada and Unit-
ed States south to Maryland, west to
North Dakota, Wyoming, and Colorado.

Biology. — We have examined speci-
mens from the CNC and USNM collec-
tions labelled as reared from the plusiine
noctuids Anagrapha falcifera (Kby.) and
Syngrapha epigaea (Grt.). The former is a
generalist on low plants including blue-
berries and clover, while the later is a
generalist feeding on conifers including
pines, spruces, and firs. The mummy
formed is typically cream-colored and
fairly smooth. Norton (1869) and Shene-
felt (1975) listed this species as having
been reared from the diprionid sawfly
Neodiprion abietis (Harris) on Abies, but
this seems very unlikely. Adults of A. par-
asiticus are active from late May to early
September.

Comments. — A. parasiticus is similar to
molestus, but parasiticus differs in having
the posterior femur and tibia banded (or-
ange on basal 0.5, black on apical 0.5). A.
parasiticus is a distinctly northern species
associated with boreal forests, while mo-
lestus is a midwestern and southern spe-
cies favoring drier and more open habi-
tats, including agroecosytems.

Aleiodes rileyi Cresson, revised
combination

(Fig. 13)

Aleiodes rileyi Cresson, 1869, Trans. Amer. Ent.
Soc. 2:382.

Diagnosis. — Body unicolored orange or
honey yellow, flagellum black, wings hy-
aline, veins brown, stigma yellow to light
brown; body length, 5.5-8.0 mm; 53-55
antennomeres; malar space short, about
equal to basal width of mandible and 0.25
eye height; ocelli large, ocellocular dis-
tance equal to or slightly less (about 0.75)
diameter of lateral ocellus; oral operung
small and circular, diameter about equal
to basal width of mandible; face rugose,
frons smooth, vertex and temple coria-
ceous; occipital carina not meeting hypos-
tomal carina; pronotum procate; mesono-
tum and scutellum coriaceous; mesopleu-
ron smooth, subalar sulcus rugose, ster-
naulus absent; propodeum rugose
dorsally, coriaceous laterally, median ca-
rina complete; first and second metasomal
terga rugose costate, median carinae com-
plete; third tergum costate on basal 0.33,
smooth or weakly coriaceous on apical
0.66, sometimes entirely smooth or weakly
coriaceous, median carina absent; remain-
der of terga weakly coriaceous; terga 4—6
in male with lateral patches of dense gold
hair; fore wing with vein Icu-a beyond IM
by distance slightly greater than length of
Icu-a; hind wing with vein RS arched in
middle, marginal cell narrowest in mid-
dle; tarsal claws strongly pectinate with 7-
8 stout teeth, apical 3-4 teeth of pectin
about 0.75 as long as apical claw, remain-
ing teeth gradually shorter towards base
of claw.

Type material examined. — Aleiodes rileyi
Cresson, holotype female, Missouri
(ANSP).

Distribution. — Connecticut south to Flor-
ida, west to Michigan, Kansas, and Sas-
katchewan. It probably occurs throughout
the eastern half of North America.

Biology. — We have examined specimens

Volume 7, Number 1, 1998

73

from the CNC, INHS, and USNM collec-
tions labelled as reared from the noctuids
Acrom/cta oblinaia (J.E. Sm.), Melanclirn pic-
ta (Harr.), and Nep^heloiies minians Gn., the
lymantriid Dasychira vagans (B. & McD.),
and possibly the pyralids Ostriiiia obiim-
bratalis (Led.), and O. penitalis (Grt.). The
known hosts are all generalist feeders on
a variety of low vegetation, shrubs, and
low trees such as willows. Adults of Alcio-
des rileyi are active in Florida as early as
January, but in northern parts of its range
(Illinois) adult activity is in late summer
(August).

Comments. — Marsh (1979) classified rile-
yi under Rogas, but we are moving it here
back to its original combination with Al-
eiodes. A. rileyi is quite distinctive by being
the only member of the species-group that
is entirely orange; all other North Ameri-
can species in the apicalis-group have at
least some black coloration on the body.

ACKNOWLEDGMENTS

This research was supported by grant DEB-930-
6314 from the National Science Foundation. Addi-
tional support was provided by supplemental Re-
search Experience for Undergraduates (REU) grants
in 1994, 1995, and 1996. Support was also provided
by a Faculty Grant-in-Aid from the University of Wy-
oming Research Office, U.W. Experiment Station Pro-
ject WYO-256-90, and a CANACOL Foundation grant
to the junior author (JCF). Additional thanks are due
to Ms. Teresa Williams, of the Western Research In-
stitute, for assistance with the Environmental Scan-
ning Electron Microscope and photography. We also
thank the curators of the museums mentioned in the
Methods section for the loan of specimens used in
this study.

LITERATURE CITED

Achterberg, C. van. 1991. Revision of the genera of
the Afrotropical and W. Palaearctical Rogadinae
Foerster (Hymenoptera: Braconidae). Zoolcgische
Verhandelingen 273: 1-102.

Achterberg, C. van. 1993. Illustrated key to the sub-
families of Braconidae (Hymenoptera: Braconi-
dae). ZiX'logisclic Vcrhamlchngcn 283: 1-189.

Cresson, E. T. 1869. List of the North American spe-
cies of the genus Aleiocia Wesmael. Transactions
of the American Entwnological Societi/ 2: 377-382.

Gahan, A. B. 1917. Descriptions of some new parasitic

Hymenoptera. Proceedings of the United States Na-
tional Museum 53: 195-217.

Harris, R. A. 1979. A glossary of surface sculpturing.
Occasional Papers in Entomology 28: 1-31.

Marsh, P. M. 1979. Family Braconidae. Pp. 144-313,
In K. V. Krombein, P. D. Hurd Jr., D. R. Smith,
and B. D. Burks (eds.). Catalog of Hymenoptera in
America North of Mexico, Smithsonian Institution
Press, Washington, D.C.

Marsh, P. M. 1989. Notes on Braconidae (Hymenop-
tera) associated with jojoba (Snnmomisia chmen-
sis) and descriptions of new species. Pan-Pacific
Entomologist 65: 58-67.

Marsh, P. M., S. R. Shaw and R. A. Wharton. 1987.
An identification manual for the North American
genera of the Family Braconidae (Hymenoptera).
Memoirs of the Entomological Society of Washington
13: 1-98.

Norton, E. 1869. American Hymenoptera. Catalog of
the described Tenthredinidae and Uroceridae of
North America. Transactions of the American En-
tomological Society 2: 321-368.

Papp, J. 1985. Contributions to the braconid fauna of
Hungary, VII. Rogadinae (Hymenoptera: Bracon-
idae). Folia Entomologica Hungarica 46: 143-164.

Shaw, M. R. 1983. On[e] evolution of endoparasitism:
the biology of some genera of Rogadinae (Bra-
conidae). Contributions of the American Entomolog-
ical Institute 20: 307-328.

Shaw, M. R. 1994. Parasitoid host ranges. Chapter 7,
pp. 112-144, In B.A. Hawkins and W. Sheehan
(eds.), Parasitoid Community Ecology, Oxford Uni-
versity Press, Oxford.

Shaw, M. R. and T. Huddleston. 1991. Classifcation
and biology of braconid wasps. Handbooks for the
Identification of British Insects 7: 1-126.

Shaw, S. R. 1993. Systematic status of Eucystomastax
Brues and characterization of the Neotropical
species. Journal of Hymenoptera Research 2: 1-11.

Shaw, S. R. 1995. Braconidae. Chapter 12.2, pp. 431-
463, In P.E. Hanson and l.D. Cauld [eds.j. The
Hymenoptera of Costa Rica, Oxford University
Press, Oxford.

Shaw, S. R. 1997. Subfamily Rogadinae s.s. Journal of
Hymenoptera Research, Special Publication 1997:
4()2-412.

Shaw, S. R., P. M. Marsh and J. C. Fortier. 1997. Re-
vision of North American Aleiodes Wesmael (Part
1): the pulchripes Wesmael species-group in the
New World (Hymenoptera: Braconidae, Roga-
dinae). Journal of Hymenoptera Research 6: 10-35.

Shenefelt, R. D. 1975. Braconidae 8: Exothecinae, Ro-
gadinae. Pp. 1115-1262, In van der Vccht and
R.D. Shenefelt, (eds.), Hymenopterorum Catologiis
(novo cditio), W. Junk B.V., The Hague.

Shenefelt, R. D. 1979. Some unusual Braconidae (Hy-
menoptera). Proceedings of the Entomological Soci-
ety of Washington 81: 125-134.

J. HYM. RES.
Vol. 7(1), 1998, pp. 74-83

Territoriality and Mating Behavior of Sphex pensylvanicus L.
(Hymenoptera: Sphecidae)

Frank E. Kurczewski

Environmental and Forest Biology, State University of New York College of Environmental
Science and Forestry, Syracuse, New York 13210-2778, USA

Abstract. — Daily observations were made on nine individually marked males of Sphex pensyl-
vanicus in upstate New York during 25 July-8 August 1982. Males occupied territories on or near
a grate atop a storm sewer drain in which 12 females nested. They obtained honeydew at a maple
tree and slept and fed on white sweet clover growing on a nearby hillside. The following main-
tenance and reproductive activities of males were defined: (1) perching at stations, (2) swivelling,
(3) cleaning, (4) wing raising, (5) defecating, (6) spontaneous flights, (7) feeding flights, (8) pounc-
ing on conspecific males, (9) pursuit flights, (10) grappling, (11) trailing conspecific females, (12)
clasping conspecific females, and (13) copulation. Aggressive interactions between territorial con-
specific males occupied more time than all other maintenance and reproductive activities com-
bined.

Almost nothing was known about the
behavior of male solitary wasps until Lin's
(1963) study of male territoriality in Splie-
cius speciosus (Drury), the cicada killer. Re-
cent interest in male behavior surfaced fol-
lowing the revival of Darwin's (1859) sex-
ual selection theory, especially as cham-
pioned by Trivers (1972) and his
contemporaries. The activities of male sol-
itary wasps are primarily aimed at obtain-
ing matings. Males feed on nectar, rest on
plants or in burrows, or bask in the sun
when not in pursuit of females (Evans and
O'Neill 1988). Males of most species of
Sphecidae are free from parental duties
and their reproductive success is solely
determined by the number of eggs they
fertilize. In other words, males of most
species contribute little more than genes to
their offspring.

Male solitary wasps tend to emerge be-
fore the females, a phenomenon known as
protandry (Evans 1966). Circumstantial
evidence suggests that females of most
digger wasps mate only once during their
lifetime (Alcock et al. 1978). Although

nesting may extend for several weeks in
certain species of Sphecidae (Hager and
Kurczewski 1986, Kurczewski 1997), cop-
ulations in most species take place only
during the first week or two. The majority
of contacts between males and females do
not end in successful copulation (O'Neill
1979). Once females are actively nesting,
they rebuff all attempted matings by
males (Evans and O'Neill 1988). The rela-
tive low fecundity of the females may
mean that they actually gain little from
additional matings (O'Neill 1985). Alcock
et al. (1978) address the costs and benefits
associated with multiple matings in spe-
cies of aculeate Hymenoptera.

Territoriality, as defined by spacing,
maintenance of stations, and aggressive
encounters between conspecific males,
does not occur in all sphecid wasps (Hag-
er and Kurczewski 1985). But, it can be ex-
aggerated in some species (Minkiewicz
1934, Lin 1963, Evans and O'Neill 1988).
Males may establish stations or territories
near prominent landmarks on the ground
(Astata, Minkiewicz 1934; Tachysphex, Kur-

Volume 7, Number 1, 1998

75

czewski 1966) or emergence holes {Splie-
ciiis, Lin 1963), or they may scent mark
plants to attract females to a site (Philan-
thus, Evans and O'Neill 1988). The estab-
lishment and maintenance of stations or
territories clearly facilitates mating be-
tween resident males and females in the
vicinity (Alcock et al. 1978).

Studies on the behavior of male solitary
wasps lag far behind those on female nest-
ing behavior (Evans 1966, Alcock et al.
1978, Evans and O'Neill 1988). Male soli-
tary wasps are often smaller, shorter lived,
and do not maintain a nest making them
more inconspicuous to the average ob-
server (Kurczewski 1966, Hager and Kur-
czewski 1985). Among the species of Sphe-
cinae (R. Bohart and Menke 1963, 1976),
many of the studies on male behavior in-
volve the genus Ammophila (Turner 1912,
Baerends 1941, G. Bohart and Knowlton
1953, Olberg 1959, Powell 1964, Hager and
Kurczewski 1985). Little is known about
territoriality and male behavior in the ge-
nus Sphex including the nearctic S. pensyl-
vauicus Linnaeus, the Great Black Wasp of
John Bartram (Rau 1944). Rigley and Hays
(1977) noted dominance, grappling, and
attempted copulation in a few males of S.
peiisi/lvanicus. Gillaspy (1962) described
mating behavior in the nearctic S. tepane-
cus Saussure. Janvier (1928) noted perch-
ing and grappling in the neotropical S. la-
treillei Lepeletier. Surprisingly, nothing is
known of male behavior in a common
nearctic species, S. ichneutiioneus (Linnae-
us)(Brockmann 1980).

My paper presents new and interesting
information on the behavior of males of S.
pensi/lvanicus and provides a verbal out-
line of the mating system employed by
this species. The literature on reproductive
behavior in sphecids is depauperate and
observations such as those reported herein
are needed to rectify this situation. Terri-
toriality and mating behavior in this spe-
cies were observed concurrent with an in-
vestigation on the sequential daily activi-
ties of the females (Kurczewski 1997). 1

chronologically sequenced and described
individual male behaviors, especially
those connected with spacing, aggression,
and copulation, and attributed possible
significance to them, thereby adding to the
overall knowledge of male behavior in the
family Sphecidae.

METHODS

Males of S. pensylvanicus were observed
on a daily basis, weather permitting, from
25 July to 8 August 1982 from 0730 to 2100
hrs (EDT). The study area was examined
before and after these dates and there was
no sign of male activity. One must as-
sume, therefore, that the males had not
emerged before 25 July and either had
died or left the area after 8 August. Nine
males each were color-coded by placing a
drop of Tester's model paint on their me-
soscutum with a tiny paint brush from
which had been removed most of the
hairs. The activities of these males were
observed and recorded at or near a storm
sewer grate for a total of 62.5 hrs during
10 of the 15 days that they were in evi-
dence. The individual life spans of the
nine males was only 10-14 days, with
three males living an entire two-weeks-
long period. I similarly marked all 12 fe-
males from this aggregation thus yielding
a secondary sex ratio of 3:2 in favor of fe-
males. One male and one female each
were collected before individually mark-
ing them and placed as voucher speci-
mens in the insect museum of the State
University of New York College of Envi-
ronmental Science and Forestry, Syracuse,
New York.

Territoriality in this species facilitated
simultaneous observation of all nine
males; however, some time was allocated
to observe certain focal males more exten-
sively during fixed observation periods.
Individual male behaviors were described
and chronologically sequenced. Particular
emphasis was placed on reproductive be-
havior as it occurred near a storm sewer
drain in which the females nested. Less

76

Journal of Hymenoptera Research

emphasis was placed on male mainte-
nance behaviors such as feeding, sleeping,
and basking in the sun that took place
away from this site.

EMERGENCE AND LOCATION OF
STUDY

Both males and females emerged from
soil near a broken tile at the bottom of the
sewer drain situated in an asphalt drive-
way beside the Marcellus Senior High
School, Marcellus, Onondaga County,
New York. Three males emerged on 25
July 1982, two days before the first female
appeared. The emergences of the other six
males shortly thereafter (27-31 July 1982)
were interspersed among those of early
emerging females. Behavior that repre-
sented aspects of territoriality such as
spacing, maintaining stations, and aggres-
sion between conspecific males was fo-
cused on a 70 X 70 cm grate atop the 70
cm-deep drain in which the females nest-
ed (Fig. 1, Kurczewski 1997). Males also
visited a maple tree 13 m south of the sew-
er drain in order to obtain honeydew, and
slept and fed at a stand of Melilotus alba
(white sweet clover) 55 m away (Fig. 2,
Kurczewski 1997). Both males and females
fed on the flowers of other plant species
nearby but not as frequently as at the flo-
rets of white sweet clover, probably be-
cause this species was more abundant in
the vicinity (Kurczewski 1997).

The nine males left their sleeping roosts
on white sweet clover daily on warm sun-
ny days, occasionally fed on nectar or
basked in the sun, and then made low cir-
cling or figure-8 flights over the sewer
grate as early as 0828-0837 hrs (EDT) at
an air temperature as low as 16°C. They
stationed themselves on and /or near the
grate from 0845 to 1643 hrs, and once as
late as 1818 hrs, at air temperatures of
16.5-26°C. Males interrupted this territo-
rial activity to feed on the flowers of Mel-
ilotus alba from 1125 to 1800 hrs at air tem-
peratures of 23-26°C. They circled the
drain in flight between 1819 and 2019 hrs.

alit briefly on or entered through openings
in the sewer grate, but then quickly exited
and flew to sleeping roosts. Males did not
maintain stations at or near the grate dur-
ing this time. They did not make extensive
and intricate flights for the purpose of ori-
entation to the immediate environs as did
the provisioning females (Kurczewski
1997).

MALE ACTIVITY

The following activities of male S. pen-
sylvanicus were delineated: (1) perching at
stations, (2) swivelling, (3) cleaning, (4)
wing raising, (5) defecating, (6) spontane-
ous flights, (7) feeding flights, (8) pounc-
ing on conspecific males, (9) pursuit
flights, (10) grappling, (11) trailing conspe-
cific females, (12) clasping conspecific fe-
males, and (13) copulation. Definition and
significance of these activities are as fol-
lows:

(1) Perching at stations. — Males perched
on or near the sewer grate with mid- and
hindlegs outstretched yet raised and
wings folded flat on the dorsum. They pe-
riodically moved their head or anterior
body from side to side. While perching,
males moved their antennae or held them
still in an upward and outward attitude.
The forelegs were either "fishhooked" me-
dially or held backward beneath the body,
raised above the substrate, and moved
back and forth slowly or in short rapid
bursts. Positioning by certain males close
to the grate openings gave them a decided
advantage over males stationed farther
away with respect to access to entering or
exiting females. These males subsequently
obtained more claspings and copulations
than males stationed farther from the sew-
er grate (see below).

Following emergence and through the
first week of nesting, males maintained
stations near the female nesting site. Males
spent much time at a preferred site or sta-
tion; however, most males maintained
several stations in close proximity moving
from one to another throughout the day.

Volume 7, Number 1, 1998

77

On 27 July 1982, five males perched at sta-
tions on or near the sewer grate. Two of
the males occupied opposite ends of the
grate, being only 50-70 cm apart. The oth-
er three males were located near the grate
at cardinal points of the compass, each
less than 1 m from the two males. The
males near the grate were not allowed to
approach the males on the grate any closer
than this without being chased away. On
the following day, four additional, newly
emerged males took up stations near the
grate less than 1 m from the older males.
Nine males now maintained one or more
stations within a diameter of 2 m from the
grate. The two males stationed on the
grate and a third male nearest the grate
perched at their stations for periods of 20-
150 min (x = 37.9 min, n = 37 observa-
tions), except for chasing away conspecific
males, females, and other insects, and
making periodic flights with no discern-
ible function. The six most peripherally
situated males, on the other hand, perched
for an average of only 3.8 (<l-7) min (n
= 41) and then flew away for 1-71 (x =
32.4, n = 41) min before returning. In oth-
er words, the three males nearest the grate
spent most of their time on or near it
whereas the six males positioned periph-
eral to the grate occupied most of their
time away from it.

The two males perching on the grate of-
ten approached to within 30 cm of one an-
other without any apparent sign of inter-
ference between them. When one of the
males flew away to feed, the other male
encroached on his territory often tempo-
rarily appropriating some of the stations.
Such replacement implies that males com-
pete for stations, the defense of which
clearly represents a form of territoriality.
The spacing distance of about 1 m be-
tween most territorial conspecific males
probably diminished the amount and in-
tensity of aggressive interaction between
them and thereby increased their chance
for successful copulation but 1 have no
concrete evidence to substantiate this.

(2) Swivelling. — Males often moved or
turned their entire body to face in a slight-
ly different direction. This movement was
made so smoothly that the wasps ap-
peared to be on a swivel. Swivelling was,
therefore, a ramification of perching.
Change in vantage ground by certain
males may have ensured that they would
not miss viewing females approaching or
exiting the drain.

(3) Cleaning. — Males frequently cleaned
their antennae, compound eyes, and
mouthparts with the forelegs, and their
wings and abdomen with the hindlegs.
Wing and abdominal cleaning followed 63
flights during which a male pursued an-
other male, female, or other insect. Only
five times did a male land from such a
flight without cleaning. Cleaning of the
antennae, eyes, and mouthparts occurred
seemingly spontaneously with or without
the presence of another insect.

(4) Wing raising. — Males perching at sta-
tions often held the wings outward at an
angle or raised and flicked the wings re-
peatedly. The former behavior was not ob-
served to be associated with any specific
incident. Sixty-one times wing flicking
was associated with the appearance of a
conspecific male, female, or other insect.
After the intruder passed by, the perched
male either lowered the wings onto the
dorsum or cleaned the wings with the hin-
dlegs and then lowered them. Raising and
flicking the wings followed by wing clean-
ing invariably followed the pursuit in
flight of another male and landing (see
above). The two males stationed only 50-
70 cm apart on the grate often wing
flicked upon visible movement of the oth-
er. Raising and flicking the wings may
have represented an intention movement
to fly as this behavior was performed usu-
ally at the appearance of another male, fe-
male, or other insect.

(5) Defecati)ig. — The two males stationed
on the sewer grate for long periods of time
periodically discharged a viscous liquid
from the anus to a distance of 10-20 cm.

78

Journal of Hymenoptera Research

Often, this fluid was squirted in two or
three directions by the wasp moving his
abdomen to a shghtly different angle.

(6) Spontaneous flights. — Every 1-8 (x =
2.1, n = 78) min, a male made a short brief
flight of no discernible function directly
away from his station, turned 180°, and re-
turned along the same route to resume
perching. Such flights were often less than
a meter in length and of only 1-2 sec du-
ration, but occasionally they were as long
as 3-6 m and several seconds in duration.
They were usually made in different,
sometimes opposite, directions from a sta-
tion. Possible functions of these flights in-
clude moving into cooler strata of air to
reduce body temperature or surveillance
of the immediate area and its occupants.

Other flights made by males were
slower, more deliberate, and longer in dis-
tance and duration. They were more or
less straight in line, often repeatedly cov-
ered the same ground, were 1-2 m in
length or longer, and invariably lasted for
more than a second or two. Some flights
took the form of repetitive short distance
figure 8's. The longer flights possibly func-
tion in helping locate receptive females.

(7) Feeding flights. — Periodically, males
temporarily left their perches and flew to
a nearby maple tree where they obtained
honeydew or to a stand of white sweet
clover from which they gathered nectar.
As many as six males were simultaneous-
ly seen in the maple tree or feeding on
Melilotus alba from late morning to early
evening. Flowers of other plant species
nearby such as Queen Anne's lace or wild
carrot, goldenrod, yarrow, and white clo-
ver were visited less frequently for nectar.

(8) Pouncing on conspecific males. — When
males at adjacent stations moved closer
than 30 cm apart, one of them often made
a short pouncing flight at the other wasp.
The latter frequently flew out of reach of
the former and then sometimes was pur-
sued in a low, sinuous flight for a short
distance. Less commonly the two individ-
uals, after making contact, grappled with

one another. This behavior may have re-
inforced the spacing of individual males.

(9) Pursuit flights. — Males maintaining
stations near the female nesting site fre-
quently flew at other males flying close to
the sewer grate or at neighboring males if
they approached too closely. Two wasps
stationed on the grate periodically chased
more peripherally located males in sinu-
ous or figure-8 flights, 1-2 m above the
ground, to distances of 3-5 m. Some of
these flights transformed into ascending
spiral flights, 3-A m high. Continuous pur-
suit flights sometimes lasted for as long as
1.5 min. Regardless of the form of the
flight, as many as four or five males par-
ticipated in a single chase with distances
of 20-30 cm separating individuals in tan-
dem. After several seconds, such flights
broke up into pairs of males or three in-
dividuals involved in chases. Flights ter-
minated when the participants returned to
their respective stations on or near the
grate. Some chases involving two or three
males continued into and out of the sewer
drain. Physical contact occurred only rare-
ly between males in pursuit of one anoth-
er. In such cases one wasp grasped anoth-
er, both fell to the ground, they grappled
for several seconds, separated, and then
flew to respective stations.

In addition to chasing conspecific males
and females, territorial males also pursued
dragonflies, moths, butterflies, horseflies,
and other wasps and bees. The initiation
of chasing depended entirely upon the
visible presence of another insect, es-
pecially a conspecific. When other males
were in the vicinity, males left their sta-
tions, took flight, and were air-borne con-
tinuously until the visitors(s) left the area.
The sight of a non-provisioning female en-
tering or exiting through the sewer grate
often incited males stationed nearby to act
aggressively toward one another and this
frequently led to pursuit flights. By chas-
ing other males out of the immediate area,
males stationed on or near the grate
would have more mating opportunities

Volume 7, Number 1, 1998

79

available to them. Such males, in fact, did
obtain more copulations with females
than males situated on the periphery of
the aggregation. Where all males were
viewed simultaneously, the three males
perching on or near the grate obtained
eight (67%) of the 12 observed matings
while the six males positioned farther
away gained only four (33%). The three
wasps stationed on or near the grate drove
away all other males and permitted fe-
males to freely fly into the nesting area.
However, copulations were successful
only with females exiting through open-
ings in the grate. Copulations with fe-
males entering the sewer drain were in-
variably unsuccessful.

(10) Grappling. — When two males ap-
proached to within 30 cm of one another
they leapt at each other, held one another
in a venter to venter position with the
legs, and bit with the mandibles. Such
grappling occurred in both anterior-ante-
rior and anterior-posterior body positions.
Grappling invariably ensued when a res-
ident male pounced upon an intruding
male flying low near the grate. Such
wasps either separated in flight and re-
turned to their respective stations or fell
to the ground, making a buzzing noise,
and continued grappling. Males stationed
next to one another on or near the grate
grappled for 2-20 (x = 11.8, n = 13) sec,
separated, and flew to respective stations.
One male stationed on the grate twice
grappled with and drove away a larger in-
truder from a peripheral station. On three
occasions two or three grappling males
fell through openings in the sewer grate,
disappeared from view, and resurfaced in
tandem flight 3-5 sec later. After exiting,
the three grappling males resumed grasp-
ing one another in flight above the grate
with the legs and continued biting with
the mandibles. Bouts of grappling even
took place within ascending spiral flights
up to 3^ m in height. Much grappling oc-
curred when non-provisioning females
flew into the area by-passing the stations

of adjacent males. One to several males
pursued such an incoming female, imme-
diately bringing them into close contact
with one another. They briefly grappled
with each other, one chased another away,
and the victor continued to pursue the fe-
male in flight. Spacing, an expression of
territoriality in males maintaining stations
near female nests, was clearly established
through bouts of grappling. Such premier
positions often facilitated successful cop-
ulation (see above).

(11) Trailing conspecific females. — During
the height of male activity the trailing
and / or pursuit of conspecific females was
secondary to chasing other males. In fact,
many females were allowed into the nest-
ing area and permitted to land without be-
ing chased or contacted by males. None-
theless, some non-provisioning females
flying near but not entering the sewer
grate were pursued by territorial males for
distances of 1^ m after which the male
returned to his station. In addition, males
stationed on the grate sometimes followed
females in flight into and out of the sewer
drain and often interfered with females
making orientation flights. Orienting fe-
males were pounced upon, bumped, and
jostled in mid-air before they flew away.
However, males did not pursue females
whose orientation flights took them away
from the drain. Attempts by males to con-
tact females in flight probably served as a
prelude to mating, but other males in the
vicinity constantly disrupted copulation
efforts.

(12) Clasping conspecific females. — Males
clasped females by flying onto their dor-
sum and holding them with the legs. Fe-
males so grasped were deprived of the use
of their wings and either plummeted
downward to the ground from heights of
30 cm to 2 m or glided onto a nearby
lawn, 2^ m from the grate. Such clasped
wasps rolled over several times while
making a buzzing sound as the female at-
tempted to escape. Sometimes the male
was dorsum down, still retaining his

80

Journal of Hymenoptera Research

grasp of the female which was also dor-
sum down with her legs dangling in the
air. One pair remained clasped together
for 3 min as they continually rolled over
and twisted on the ground while making
a constant buzzing sound. They eventu-
ally separated but the male clasped the fe-
male again and repeated his attempted
copulation for an additional 1.5 min. Un-
successful copulation in the form of clasp-
ing, however, usually lasted only 5-45 (x
= 18.7, N = 14) sec. Following attempted
copulation, males cleaned the sides of
their abdomen alternately with the hin-
dlegs and their eyes, antennae, and
mouthparts alternately with the forelegs.
Clasping of females by males preceded
mating but most such engagements never
went to completion.

(13) Copulation.— Twelve of 29 (41%)
pairings that began as clasping culminat-
ed in several minutes-long matings. Gen-
italic union was not achieved in 17 pair-
ings that dissolved after 5 sec-4 min (x =
47.8 sec) of clasping. The twelve appar-
ently successful matings were observed
between 1021 and 1231 hrs during 27-31
July 1982. This period, when males were
2-6 and females only 1-4 days old, rep-
resented the pinnacle of reproductive ac-
tivity insofar as frequency of claspings
and copulations. Attempted copulations
were sparse before and after this time.
Matings were not seen during the second
week of a male's life.

Successful copulation proceeded
through a sequence of behaviors. Males al-
ways mounted females dorsal side up in
a head above head position. Usually, the
male remained on top with the female on
the bottom. However, sometimes the male
was on his side or, rarely, on his back
holding the female ventral side up with
her legs dangling in the air. Regardless of
orientation, the male retained his grasp of
the female by using his legs to grip her
wings, body, and legs.

In the beginning stages of copulation, a
male stroked a female's antennae with his

antennae. This behavior seemingly aided
in keeping her acquiesced. When the fe-
male became restless, she sporadically
produced a buzzing sound. In order to
contact her genitalia, the male rubbed the
sides of his abdomen against the under-
side of the female's abdomen. The male's
abdomen had to be telescoped and the fe-
male's abdomen mildly contracted to ex-
ecute coupling. The last half of his abdo-
men had to be twisted nearly 45° and the
female's abdomen turned somewhat.
Once coupling was achieved, periodic
sound production ceased. There was some
rhythmic abdominal movement from both
participants during copulation.

Males and females remained coupled
for an average of 5.1 (R = 2.5-14.0, N =
12) min. Following an apparently success-
ful copulation, the female feebly attempt-
ed to release herself from the male by
slowly moving her legs and antennae and
twisting her head in an arc. In some cases,
the female tried to break loose by walking
on the substrate dragging the attached
male behind. Coupling, separation, and
recoupling of male and female occurred
up to three times in certain pairs. The ar-
rival of other males or, rarely, females of-
ten led to disruption of mating and, al-
most invariably, to dissolution of the pair-
ing. Twice, intruding females entering the
mix led to genitalic separation and cessa-
tion of mating. Seven times an intruding
male mounted the first male while he was
mating, dislodged him, and disjoined the
coupling. Recoupling by the original pair
was sometimes achieved after such a dis-
ruption.

Once, as many as three males attempted
to copulate with a single female, the par-
ticipants being stacked in a pile atop one
another. The pile of bodies kept falling to
one side, but the pedestal male retained
his grasp of the female and the other
males their grasp to one another. Whether
the pile remained upright or fell onto its
side made no difference in the respective
positioning of the males, probably because

Volume 7, Number 1, 1998

81

the pedestal male was holding the bases
of the female's wings with his forelegs and
ad infinitum. After 35 sec, the uppermost
male flew away for a few seconds only to
return and rejoin the pile. In his absence,
the remaining two males each tried to
make genitalic contact with the female by
fencing for prime position with the ends
of their abdomens. Sound production oc-
curred on and off during the entire at-
tempted copulation. After 4 min of re-
maining together, the participants sepa-
rated and flew away without successfully
mating.

Male activity summary. — The activities of
four focal males stationed on or near the
grate were observed and recorded for 30
min-long periods at air temperatures of
23-24°C between 1015 and 1553 hrs on 29-
31 July 1982 for the purpose of summariz-
ing and ranking them. Of the combined
120 min spent by these males, 67 (56%)
min were utilized for flying, chasing, and
grappling, 43 (36%) min for perching,
swivelling, cleaning, wing raising, and
defecating, 6 (5%) min for clasping and
copulating, and 4 (3%) min for feeding on
honeydew and the flowers of white sweet
clover. The large proportion of time spent
air-borne by these males coincided with
the presence of conspecific males and fe-
males in the area. Ninety percent of the
time (60/67 min) used for flying, chasing,
and grappling by males stationed on or
near the sewer grate included the intru-
sion of conspecific males stationed nearby.

DISCUSSION

Male solitary wasps often emerge one
or a few days before the females (Evans
1966). This trend, known as protandry,
and an overall shorter flight season syn-
chronize male activity with female emer-
gence (Evans and O'Neill 1988). Early
emergence of males gives them an oppor-
tunity to mate with unmated, recently
emerged females (Bulmer 1983). One-third
of the males of Sphex pensylvatiicus that I
studied emerged two days before the re-

mainder of the males and the first females.
The early emerging males were more suc-
cessful in establishing stations near the fe-
male nesting site than later emergents and
this eventually resulted in a higher pro-
portion of matings for these individuals.

Rigley and Hays (1977) noted a "domi-
nance order" among males of Sphex pcn-
sylvanicus for about a week during the
"latter part" of July before the females
started provisioning. One male positioned
himself nearer the female burrows than
two other males, continually chased them
from the area, occasionally caught and
grappled with them, and temporarily
drove them away. The "dominant" male
"solicited" females at their burrow en-
trances by repeatedly flicking his wings,
flew after females as they exited their en-
trances, clasped them in flight, disap-
peared from view, and then returned 2-5
min later to resume perching near the
nests.

Behavior of male sphecid wasps is pre-
dominantly directed toward obtaining
matings (Hager and Kurczewski 1985,
Evans and O'Neill 1988). Males of many
digger wasp species maintain territories or
perches near female nests in order to gain
a reproductive advantage (Lin 1963,
O'Neill 1979, Evans and O'Neill 1988).
Copulatory attempts near nesting sites un-
derline the importance of territories or
perches to facilitate mating (Alcock et al.
1978). In Sphex pcnsylvanicus, females often
nest aggregatorily in one area (Reinhard
1929, Frisch 1938, Rigley and Hays 1977,
pers. obs.). Males establish stations near
the female nests, the two sexes are contin-
ually brought into contact with one anoth-
er, and mating is expedited.

Male territoriality, as defined by spac-
ing, maintenance of stations or perches,
and aggression between conspecifics, has
been demonstrated for a number of sphe-
cids (Lin 1963, Evans 1966, Kurczewski
1966, Alcock et al. 1978, O'Neill 1979,
Evans and O'Neill 1988, Hastings 1989).
Applying this definition to Sphex pcnsyl-

82

JuuRNAi, or- Hymenoptera Research

imiiicus, territoriality clearly is operational
among the males (Rigley and Hays 1977,
pers. obs.)- In this species spacing and sta-
tion maintenance and defense may func-
tion in retiucing conflict between conspe-
cific males and promoting successful cop-
ulation.

Although mating is the ultimate goal of
male digger wasps, aggressive interac-
tions between conspecifics are clearly the
most conspicuous activities of territorial
males (Evans and O'Neill 1988). More
than 90% of the time spent by males of
Sphcx pcnsi/lvniiiciis during a three day-
long observation period involved aggres-
sive activities connected with territoriality.
Males used most of this time in making
flights, grappling with and pursuing other
males, and vigorously defending stations
against conspecifics. Males of Sphcx pcn-
si/lvanicus utilized only about 5% of their
available time for clasping and mating.
Some males, in fact, consistently ignored
females flying into the area in order to
pursue other males. However, if a male
spends too much time on aggressive in-
teractions with conspecific males, he
wastes time and energy that could be used
for locating and contacting females
(O'Neill 1979).

Copulation in Sphcx pcnsylvanicuf is
similar in configuration and relative posi-
tions of male and female to that of other
sphecines, especially members of the ge-
nus Aiiiiuophila (Turner 1912, Baerends
1941, Olberg 1959, Powell 1964, Hager and
Kurczewski 1985). In the former species,
duration of coupling is apparentlv briefer
than in species of AiinuopliiLj and the male
and female abdomens are held outward
rather than raised upward. Uncoupling
and recoupling are characteristic features
of copulation in btith Sphcx and Aintiiophila
(Baerends 1941, Olberg 1959, Hager and
Kurczewski 1985, pers. obs.). Coupling,
separation, and recoupling in certain pairs
of Sphcx pctitii/lvanicii:^ occurred up to three
times during one copulation event. Dis-
ruption of matings by conspecifics occurs

frequently in AmmophUa and Sphex. Two
or more males attempting to mate with a
single female and, in the process, disen-
gaging the initial coupling appears to be a
common strategy in both genera. In such
a case an intruding male niay benefit im-
mediately or later via successful copula-
tion with the disjoined female (Hager and
Kurczewski 1985).

ACKNOWLEDGMENTS

1 am grateful to M. G. Spofford for assistance in the
field. 1 thank B. ]. Hager, K. M. O'Neill, L. S. Vlietstra,
and two anonymous reviewers for improving the
manuscript.

LITERATURE CITED

Alcock, J., E. M. Barrows, G. Gordh, L. J. Hubbard,
L. Kirkendall, D. W. Pyle, T. L. Ponder, and F.
G. Zalom. 1978. The ecology and evolution of
male reproductive behaviour in the bees and
wasps. ZooU^\;icnl lounuil iV'/u' Uiuuiciiii Si'cicty 64:
293-326.

Baerends, G. P. 1941. Fortptlanzungsverhalten und
drientierung der Grabwespe Ainiiti'fhihi aimpes-
trif. Jur. Tijiticlirift iwr Eiitcinokigic 84: 68-275.

Bohart, G. E. and G. F. Knowlton. 1953. Notes on mat-
ing, prey provisioning and nesting in Sphcx pro-
cerus (Dahlbom). Prcccctiiiigs of the Entomological
Society of WiKhington 55: loVlOl.

Bohart, R. M. and A. S. Menke. 1963. A reclassifica-
tion of the Sphecinae with a revision of the nearc-
tic species of the tribes Sceliphronini and Sphe-
cini. University of Californm Publications m Ento-
mology 30: 91-182.

Bohart, R. M. and A. S. Menke. 1976. Sphcciti Wasps
of the World: A Generic Revision. University of Cal-
ifornia Press, Berkeley. 695 pp.

Brockmann, H. J. 1980. The control of nest depth in a
digger wasp (Splwx ichneiimoneiis L.). Animal Be-
haviour 28: 425-445.

Bulmer, M. G. 1983. The significance ot prot.indrv in
social Hvmenoptera. American Naturalist 121:
540-551.

Darwin, C. 1859. On the Origin of Species. (Reprint of
first edition, 1964]. Harvard University Press,
Cambridge, Massachusetts. 502 pp.

Evans, H. E. 1966. The behavior patterns of solitary
wasps. Annual Revierv of Entomology 11: 123-154.

Evans, H. E. and K. M. O'Neill. 1988. Tlie Natural His-
tory ami Behavior of North American Beewolves.
Comstock Publishing Associates, Ithaca, N. Y.
27S pp.

Frisch, J. A. 1938. The life-history and habits of the
digger-wasp Ammobia pennsylvanica (Linn.).
American Miilland Naturalist 19: 673-677.

Vol iiMi: 7, Number 1, 1998

83

Gillaspy, J. E. 1962. Nesting behavior of Sphex tq>a-
nccus Saussure. Bulletin of the Brooklyn Entomolog-
ical Society 57: 15-17.

Hager, B. J. and F. E. Kurczewski. 1985. Reproductive
behavior of male Ammophila harli (Fer-
nald)(Hymenoptera: Sphecidae). Proceedings of
the Lntomological Society of Washington 87: 597-
605.

Hager, B. J. and F. E. Kurczewsl<i. 1986. Nesting be-
havior of Ammophila harti (Fernald) (Hymcnop-
tera: Sphecidae). The American Micilanil Naturalist
116: 7-24.

Hastings, J. M. 1989. The influence of size, age, and
residency status on territory defense in male
western cicada killer wasps (Sphecius grandis, Hy-
menoptera: Sphecidae). journal of the Kansas En-
tomological Society 62: 363-373.

Janvier, H. 1928. Recherches biologiques sur les pre-
dateurs du Chili. Annates des Sciences Nalurelles
Zoologie et Biologic Animate 11 (10): 67-207.

Kurczewski, F. E. 1966. Comparative behavior of
male digger wasps of the genus Tachysphex (Hy-
menoptera: Sphecidae, Larrinae). journal of the
Kansas Entomological Society 39: 436-453.

Kurczewski, F. E. 1997. Activity patterns in a nesting
aggregation of Sphex pensylvanicus L. (Hymenop-
tera: Sphecidae). journal of Hymenoptera Research
6: 231-242.

Lin, N. 1963. Territorial behavior in the cicada killer
wasp, Sphecius speciosus (Drury). I. Behaviour 20:
115-133.

Minkiewicz, R. 1934. Les types de comportement des
males des sphegiens. Polskie Pismo Entomologiczne
13: 1-20.

Olberg, G. 1959. Das Verhalten der solitaren Wespen
Mitteleuropas. Deutscher Verlag Wisscnschaften,
Berlin. 401 pp.

O'Neill, K. M. 1979. Territorial behavior in males of
Philanthus psyche (Hymenoptera, Sphecidae). Psy-
che 86: 19^3.

O'Neill, K. M. 1985. Egg size, prey size, and sexual
size dimorphism in digger wasps (Hymenoptera:
Sphecidae). Canadian journal of Zoology 63: 2187-
2193.

Powell, J. A. 1964. Additions to the knowledge of the
nesting behavior of North American Ammophila.
journal of the Kansas Entomological Society 37: 240-
258.

Rau, P. 1944. The nesting habits of the wasp, Chlorion
(Ammobia) pennsylvanicum L. Annals of the Ento-
mological Society of America 37: 439^140.

Reinhard, E. G. 1929. The Witchery of Wasps. Century
Company, New York. 291 pp.

Rigley, L. and H. Hays. 1977. Field observations in-
cluding acoustic behavior of the Black-Digger
Wasp, Sphex pennsylvanicus (Linn.). Proceedings of
the Pennsylvania Academy of Science 51: 32-34.

Trivers, R. L. 1972. Parental investment and sexual
selection, pp. 136-179. In: B. Campbell (ed). Sex-
ual Selection and the Descent of Man. Aldine, Chi-
cago.

Turner, C. H. 1912. The copulation of Ammophila ab-
breviata Fabr. Psyche 19: 137.

J. HYM. RES.
Vol. 7(1), 1998, pp. 84-93

The Effects of Cavity Diameter and Length on the Nesting Biology
of Osmia lignaria propinqua Cresson (Hymenoptera: Megachilidae)

Richard W. Rust
Biology Department, University of Nevada, Reno, Nevada 89557, USA

Abstract. — When offered equally available trap-nests of 6, 7, or 8 mm diameter and 80, 140, or
230 mm length, Osmia lignaria propinqua Cresson females chose significantly more of the deepest
cavities for nesting and produced 69% of their offspring in them. Proportionately more females
were produced in both years in cavities of greater length and diameter. Male and female weight
varied between years, cavity lengths, and cavity diameters. Mortality was not significantly related
to either nest diameter or length. Nesting parameters of ten species of megachilid bees showed a
positive significant relationship between both female body width and nest cavity diameter and
female body length and cavity length. However, there was no relationship between female body
width or length and number of cells per nest.

The cavity or wood-nesting wasps and
bees (Malyshev 1935; Krombein 1967),
with species found in several families
(Krombein 1967; Evans and West-Eber-
hard 1970; Stephen et al. 1969; Gauld and
Bolton 1988), represent one of the biolog-
ical groupings within the Hymenoptera.
These provisioning wasps and bees use
natural cavities for the placement of food
for rearing their offspring (Krombein
1967). Two quite different life histories ex-
ist in the group (Evans and West-Eberhard
1970). The primitive members never de-
veloped the ability to make a nest, where-
as the advanced members typically make
a series of cells, separated by partitions.
The latter group has received much atten-
tion because of the development of the
technique of "trap-nesting" (Krombein
1967) and their economic value as man-
aged pollinators (Torchio, 1987, 1990;
Bosch et al. 1992).

Evidence supporting Fisher's sex ratio
theory (Fisher 1958) has come from both
natural history observations on cavity
nesting bees and wasps (Rau 1928, 1937;
Malyshev 1935; Krombein 1967; Danks
1970, 1971; Maeta 1978) and experimental

studies (Stephen and Osgood 1965; Gerber
and Klostermeyer 1972; Phillips and Klos-
termeyer 1978; Torchio and Tepedino
1980; Cowan 1981; Freeman 1980; Tepe-
dino and Torchio 1982a, 1982b, 1989; Te-
pedino and Parker 1983, 1984; Frohlich
and Tepedino 1986; Johnson 1988, 1990;
Sugiura and Maeta 1989; Bosch 1994). The
resource quality model of cavity size
(Charnov 1982) appears to be the domi-
nant factor in the facultative sex ratios ob-
served. Female parents of these sexually
dimorphic species produce a greater pro-
portion of progeny of the larger sex (usu-
ally females) in wider diameter cavities,
and /or additional progeny of the smaller
sex (usually males) in narrowed cavities.
Size distribution of nest cavities available
to the parent generation will shift sex ra-
tios; niore males are produced in narrow
cavities, and more females are produced
in wide cavities (Charnov et al. 1981).

The genus Osmia contains several spe-
cies that have been developed as commer-
cial pollinators of fruit crops (Maeta 1978,
Torchio 1987, 1989, 1990, Bosch 1994).
Much of the biological information ob-
tained concerns the procurement and uti-

Volume 7, Number 1, 1998

85

lization of these species as pollinators. It
has already been well established that
both the sex ratio and the weight of indi-
vidual bees of Osmia lignaria pwpinqua
Cresson increased with cavity diameter
(Torchio and Tepedino 1980; Tepedino
and Torchio 1982a, 1982b, 1989). Bosch
(1994) found that when Osmia coniiita
Latr. was given a choice of cavity lengths
for nesting they preferred the longest cav-
ity. Maeta (1978) reported the use of an
average cavity length and diameter for
five Japanese Osmia species when given an
assortment of diameter and length reeds
for nesting.

Osmia I. pwpinqua has not been offered
trap-nests of varying length (Torchio 1976,
1984a, 1984b, 1985). This paper examines
the combined effects of cavity diameter
and length on cavity acceptance, cavity
use, offspring weight and sex, mud use for
cell partitions and plugs, and mortality in
Osmia lignaria pwpinqua. Also, the relation-
ship of average female body size to aver-
age cavity selection in other cavity nesting
megachilid bees is examined.

MATERIALS AND METHODS

Osmia 1. pwpinqua individuals were ob-
tained from trap-nests placed out during
the spring and summer of 1991 and 1992
in Lake City, Modoc Co., California. Trap-
nest were pine blocks drilled with 6, 7, or
8 mm holes to a depth of 80, 140, or 230
mm. Trap-nests were bundled together in
groups of nine, one each by diameter and
length, and 30 bundles or 270 nesting cav-
ities were placed in each of four wooden
boxes (30 X 30 X 60 cm) supported by
metal fence posts 1.5 m off the ground.
Trap-nest openings faced southeast. Boxes
were placed out in mid March near the
edge of two mixed fruit (apricots, apples,
pears, peaches and plums) orchards and
were from 100 to 200 m apart. With the
onset of apricot bloom, trap-nests were
checked daily for nesting. Completed,
plugged nests were marked and dated.
Trap-nests were removed in September, x-

rayed, and held out-doors in Reno, Wash-
oe Co., Nevada. In March, nests were
opened and nesting information was re-
corded and adult bees were weighted.
Temperature and precipitation were re-
corded daily from a station adjacent to one
orchard. The duration of seasonal nesting
activity was based on daylight tempera-
tures above 15°C on precipitation free
days (Torchio 1976).

Mean body lengths and widths for fe-
male Osmia were from Sandhouse (1939),
Yasumatsu and Hirashima (1950), Rust
(1974), and Peters (1977); for Hoplitis from
Michener (1947); and for Megachile from
Mitchell (1962).

Analysis of variance (GLM in SAS 1990)
was used for all comparative analyses be-
tween years, cavity diameters, and cavity
lengths and the cells per nest, mud per cell
per nest, placement of first and last cell in
a nest, and percentage (arcsine transfor-
mation) of mortality per nest using Type
III sums of squares due to unequal obser-
vations (Cody and Smith 1991). Duncan's
multiple-range test was used for multiple
comparisons when analysis of variance in-
dicated a significant difference. Female
body part measurements were compared
to nest dimensions using linear regression.
Chi-square test and G-test (Sokal and
Rohlf 1969) were used to compare trap-
nest usage patterns.

RESULTS

In 1991, nesting began on 4 May: on 22
May the first nest was completed. Forty-
three days (4 May to 15 June) passed to
accumulate 200 hours of 15°C or greater
temperatures. During that period, rain or
snow (57 mm precipitation) fell on 11 days
and an additional three days were below
15°C. In 1992, nesting began on 14 April:
on 29 April the first nest was completed.
Thirty-four days (14 April to 17 May)
passed to accumulate 200 hours of 15°C or
greater temperatures. During that period,
rain (20 mm) fell on two days and another
six days were below 15°C.

86

Journal of Hymenoptera Research

Table 1. Nests and cells produced by Osniia lignaria propinqua in cavities with 6, 7, and 8 mm diameters
and 80, 140 and 230 mm lengths in 1992 and 1993 at Lake City, California.

Nest.s

Cell:

Cell

Diameter

1991

1992

Total

1991

iw;

Total

Mean ± SD

80 mm length

2.8 ± 1.2

6.0

4

3

7

14

7

21

7.0

0

2

2

—

6

6

8.0

1

2

3

3
140 mm length

10

13

5.4 ± 2.2

6.0

7

9

16

43

38

81

7.0

6

5

11

39

22

61

8.0

7

3

10

47
230 mm length

13

60

8.1 ± 4.0

6.0

12

9

21

101

55

156

7.0

7

10

17

70

62

132

8.0

12

9

21

117
Totals

78

195

56

52

108

434

291

725

Total Nest Mean ± SD

Total Cell Mean ± SD

7.7 ± 3.4

5.4 ±

3.6

A total of 108 nests containing 725 cells
were produced (Table 1). Most nests (59)
and cells (482) were approximately evenly
divided among the 6 mm, 7 mm, and 8
mm diameter long cavities (230 mm) (Ta-
ble 1). The least nests (12) and cells (40)
were in the 6 to 8 mm diameter by 80 mm
short cavities. The distribution of nests
was significantly different from the avail-
ability of cavities for nesting (x* = 35.23,
df = 8, P < 0.001). Partitioning the nests
by cavity length and cavity diameter
showed no significant pattern (G = 2.66,
df = 4, 0.75 > P > 0.50).

The mean number of cells per nest was
significantly different between the years
with more cells in 1991 than in 1992 (F =
4.51, P = 0.03). Cell distribution paralleled
nest distribution with significantly more
cells in longer cavities (F = 15.42, P <
0.001) but a similar number of cells in all
cavity diameters (F = 0.4, P = 0.66). Nests
in 230 mm length cavities contained sig-
nificantly more cells than did nests in 140
mm and they contained more than did
nests in 80 m cavities (Table 1).

The distribution of adults by cavity

length was 69% in 230 mm cavities, 28%
in 140 mm cavities, and 3% in 80 mm cav-
ities and by cavity diameter was 32% in 6
mm cavities, 30% in 7 mm cavities, and
38% in 8 mm cavities. Five hundred and
forty-nine adults (317 males and 232 fe-
males) were produced (Table 2). In 1991,
312 adults (166 males and 146 females)
were produced, and in 1992, 248 adults
(151 males and 86 females) were pro-
duced. The distribution of males or fe-
males by cavity length was similar to the
distribution of all individuals. However,
the distribution of males or females by
cavity diameter showed different patterns
with the number of males decreasing with
increasing cavity diameter (6 mm — 44%, 7
mm— 30%, and 8 mm— 26%) and the
number of females increasing with in-
creasing cavity diameter (6 mm — 17%, 7
mm— 30%, and 8 mm— 53%).

Both male and female weights showed
a significant year by cavity length by cav-
ity diameter interaction (males F = 3.81, P
= 0.02; and females F = 3.88, P = 0.02)
and were thus separated by year to ex-

Volume 7, Number 1, 1998

87

Table 2. Male and female production by Osmia //sf-
nnrm propmqun in cavities with 6, 7, and 8 mm di-
ameters and 80, 140, and 230 mm lengths in 1991 and
1992 at Lake City, California.

Males

Females

Didnieter

IMUI

1W2

Ti.t.il

I9»l

loo:

Ti.tal

80

mm length

6.0

0

5

5

0

1

1

7.0

0

1

1

0

0

0

8.0

1

2

3

2

5

7

Total

8

8

140

mm length

6.0

27

26

53

7

3

10

7.0

18

13

31

16

5

21

8.0

14

4

18

13

8

21

Total

102

52

230

mm length

6.0

46

35

81

20

8

28

7.0

34

28

62

24

24

48

8.0

26

37

63

64

32

96

Total

206
Totals

172

166

151

317

146

86

232

amine the effects of cavity length and di-
ameter on individual vv^eights (Table 3).

In 1991, male weights differed signifi-
cantly among cavity lengths (F = 23.5, P
= 0.0001), cavity diameters (F = 7.79, P =
0.001), and the cavity length by diameter
interaction (F = 130.5, P = 0.01) (Table 3).

Males in 80 mm length cavities were
heavier than males in 140 mm or 230 mm
cavities. Males in 8 mm diameter cavities
v^ere heavier than males in 7 mm or 6 mm
cavities. However, in 1992 male weights
were not significantly different in cavity
lengths, cavity diameters, or cavity length
by diameter interaction.

In 1991, female weights in were signifi-
cantly different for cavity lengths (F =
6.75, P = 0.002), cavity diameters (F =
2.72, P = 0.07), and the cavity length by
diameter interaction (F = 2.94, P = 0.06)
(Table 3). Females in 80 mm cavities were
significantly heavier than females in 140
mm or 230 mm. Females in 6 mm cavities
were significantly lighter than females in
7 mm or 8 mm cavities. However, in 1992
female weights were not significantly dif-
ferent among cavity diameters or in cavity
lengths and or their interaction.

The sex ratios based on mean weight of
all males and females ranged from 1.2:1 to
1.9:1 (males: females) and were slightly
different between years (Table 4). Sex ra-
tios increased in both years with increas-
ing cavity lengths (1.2:1 to 1.8:1) but not
diameters (1992—1.6:1 to 1.9:1 and 1991—
1.6:1 to 1.5:1).

The amount of mud used for each cell
per nest was significantly different be-

Table 3. Mean weights and standard deviations (mg) of male and female Ounia lignnrui prop'inqua produced
in cavities with 6, 7, and 8 mm diameters and 80, 140, and 230 mm lengths in 1991 and 1992 at Lake City,
California. * equals one individual.

M.

ales

Females

Parameter

HMI

IW2

IWl

igq:

Cavity diameter

6.0

35.8 ± 5.7

34.2 ± 4.8

56.8 ±11.2

54.8 ± 7.6

7.0

36.0 ± 7.3

35.1 ± 5.8

63.3 ± 9.1

59.7 ± 11.6

8.0

44.9 i 9.5

32.6 ± 6.9

Cavity length

68.9 ± 12.7

61.7 ± 12.6

80

70.9'

32.6 ± 5.6

83,7 ± 5.5

53.2 ± 4.1

140

40.6 ± 6.5

35.3 ± 5.1

67.9 -t 10.2

62.2 ± 12.1

230

33.7 ± 5,2

33.4 ± 6.1

Total

61.8 ± 11.7

60.6 ± 12.3

Total

37.7 ± 5.1

33.9 ± 5.8

64.1 ± 10.1

60.3 ±11.4

88

Journal of Hymenoptera Research

Table 4. Sex ratio of Osiiiin Ui;iuuiii prcpiiiqun pro-
duced in cavities with 6, 7, and 8 mm diameters and
80, 140, and 230 mm lengths in 1991 and 1992 at Lake
City, California.

Table 5. Percent mortality in Osiiiin Ugiuirin pircpin-
qua cells produced in cavities with 6, 7, and 8 mm
diameters and 80, 140, and 230 mm lengths in 1991
and 1992 at Lake City, California.

P.ir.inicttT

mm

um:

DianifltT

|W1

IMu;

IoCjI

Cavity diameter

80 mm length

6.0

1.6:1

1.6:1

6.0

100.0

14.3

71.4

7.0

1.7:1

1.7:1

7.0

—

83.3

83.3

8.0

1.5:1

1.9:1

8.0

0.0

30.0

23.1

Cavity length

140 mm length

80

1.2:1

1.6:1

6.0

20.9

23.7

22.2

140

1.7:1

1.8:1

7.0

12.8

18.1

14.7

230

1.8:1

1.8:1

8.0

42.5

7.6

35.0

Totals

230 mm length

Total

1.7:1

1.8:1

6.0

34.6

21.8

30.1

7.0

17.1

16.1

16.6

8.0

23.1

11.5

18.5

tween years (F = 7.32, P = 0.008) with less
mud used in 1991 (mean 92.2 ± 33.2 mg

Total for:
Total for:
TOTAL

80
6

mm 60.0,

mm 31.0,

28.1

140 mm 23.7,
7 mm 18.1,
18.5

230
8

mm 21.7
mm 22.4
24.2

Tjor-mic IQQO-

mc^r\ 1 C;c; T -1- Ql^ '

I t-v-\rr\ \

A^^A

investment per cell by cavity length and
diameter was also significantly different
(cavity length F = 6.46, P = 0.002, cavity
diameter F = 16.4, P = 0.001). There was
more mud per cell in the 80 mm cavities
(163.3 ± 85.0 mg) when compared to the
140 mm (116.9 ± 77.8 mg) and 230 mm
(117.7 ± 73/2 mg). The 6 mm diameter
cavity nests contained less mud per cell
(88.7 ± 36.9 mg) than either the 7 mm
(146.0 ± 68.9 mg) or 8 mm (145.6 ± 102.9
mg).

Bees using 230 mm length cavities
placed the first cell not at the bottom of
the cavity but at an average of 21.4 ± 35.2
mm from the bottom of the cavity. Bees
nesting in 80 or 140 mm length cavities
place the first cell at the bottom of the cav-
ity. The last cell in a nest was significantly
closer to the entrance in 80 mm cavities
(mean 33.7 ± 15.8 mm) than in either 140
mm (mean 50.7 ± 27.5 mm) or 230 mm
(mean 75.1 ± 54.4 mm) cavities (F = 5.17,
P = 0.007). For both the placement of the
first and last cell in a nest, neither year,
cavity diameter, or any of the interactions
were significant.

The percent mortality averaged 24.2%
for all cells produced and was not signif-

icantly different between years or among
cavity lengths, cavity diameters, or any of
the interactions (Table 5). Chalk brood
{Ascosphaera torchioi Youssef and Mc-
Manus) caused the greatest loss in 1991
(15.6%) whereas egg death or failure to
hatch caused the greatest loss in 1992
(7.5%) (Table 6). Chalk brood was the
overall highest mortality factor (10.9%).

Nesting parameters of ten species of
megachilid bees have been reported on
where they were provided with a choice
of cavity lengths (Table 6). There was a
positive relationship between female
mean body width and mean preferred
cavity diameter (Y (cavity diameter) =
1.10 -H 1.50X (body width); F = 2.62, P =
0.144). The relationship became significant
when Osmia marginata was removed from
analysis (Y (cavity diameter) = -0.48 +
1.84X (body width); F = 16.6, P = 0.005).
Osmia marginata does not place cells in a
linear series in large diameter cavities, but
fits cells to the cavity dimension that al-
lows a maximum use of the space (Tepe-
dino and Parker 1983). Female mean body
length and mean cavity length also
showed a significant positive relationship

Volume 7, Number 1, 1998

89

Table 6. Female size and nest parameters of megachilid bees from studies where different length cavities
were available for nesting.

Female size

Nest characteristics

Mean wn

dth

Mean length

Mean diameter Mean lenj^th

Mean »

Spi-vK's

(mm)

(mm)

(mm)

(mm)

lit cells

Ketereiue

Osiriin

iinaii

3.2

8.7

5.3

9.9

8.9

Maeta 78

tiiunii

4.1

11.0

6.8

14.9

9.0

Maeta 78

cornifrons

3.5

9.7

5.8

14.8

8.0

Maeta 78

pctiicornis

4.3

12.5

6.7

14.6

6.8

Maeta 78

L'xcazmta

3.8

10.7

6.5

15.0

7.2

Maeta 78

corntita

4.0

12.0

8.0

17.1

4.0

Bosch 94

iiuu\;iniit{i

3.5

9.0

9.0

9.0

8.2

Tepedino & Parker 83

lt};iuirm

3.8

11.5

6.9

18.2
Hoptlitis

6.6

present

fulgida

3.5

9.7

6.0

8.5
Mff;iuhik

4.9

Tepedino & Parker 84

rotundata

3.0

8.5

4.9

7.8

6.3

Gerber & Klost. 72

(Y (cavity length) = -9.95 + 2.22X (body
length); F = 17.18, P = 0.003). There was
no relationship between female body
width or length and the mean number of
cells per nest.

DISCUSSION

Females of Osmia I. propinqua chose sig-
nificantly deeper drilled holes in trap-
nests when presented with an equal dis-
tribution of cavity lengths for nesting.
These deeper holes were filled with more
cells and offspring. However, females
showed no preference for a particular di-
ameter trap-nest from within the range
available. This nest selection pattern was
observed in both years. In similar studies,
Bosch (1994) found a significant prefer-
ence in Osmia cornuta for longer cavities
for nesting (12, 15, or 21 cm, cavity di-
ameter was 8 mm) and produced more
cells in them. Osmia marginata Michener
females preferred nest-traps in drilled el-
derberry {Snmbuciis spp.) stems with the
longest (90 mm) and widest (9 mm) cavi-
ties (Tepedino and Parker 1983). They pro-
duce significantly more cells in them. Te-
pedino and Parker (1984) observed the
same selection pattern in Hoplitis fulgida

(Cresson) nesting in drilled elderberry
stems. The opposite cavity usage pattern
was observed in the completed nests of
Megachile rotundata (F.) which decreased
from 100% in 1.25 and 2.5 cm length cav-
ities to 16% in 15 cm cavities (Stephen and
Osgood 1965). However, nest utilization
increased as cavity diameter increased
from 4.0 to 6.0 mm. Gerber and Kloster-
meyer (1972) also found Megachile rotun-
data to use more short (4 cm compared to
8, 12 or 16 cm cavity lengths) trap-nests
for nesting in a three years study. How-
ever, unlike the Stephen and Osgood
(1965) results, Gerber and Klostermeyer
(1972) found more cells were produced in
8 to 16 cm length cavities.

Maeta (1978) provided Osmia imaii Hir-
ashima, O. taurus Smith, O. cornifrons (Ra-
doszkowski), O. pedicornis Cockerell, and
O. excavata Alfken with a broad selection
of reed cavities for nesting (4 to 11.9 mm
in diameter and 3 to 33 cm in length). His
presentation of nesting materials attempts
to represent what is most likely available
to the species in nature and consequently
the usage preference observed best repre-
sents the species natural usage patterns
(Table 6).

90

Journal of Hymenoptera Research

The general pattern of cavity choice
suggests that female size dictates her
choice in both the diameter and length of
cavity. This assumes that vk'ithin the nat-
ural habitat there exists a variety of cavi-
ties for nesting and that females visit sev-
eral cavities before making a selection. Se-
lection of a short cavity will require find-
ing a second or even third cavity to
continue the nesting process. Selection of
a long cavity simply requires the female
to initiate cell construction at the "aver-
age" depth to maximize the cells or off-
spring produced. The bee is leaving un-
used cavity behind the first cell. The pres-
ence of an empty space behind the last cell
was evident in only in the 230 mm long
cavities. Maeta (1978) found an increasing
percentage of empty spaces with increas-
ing cavity length in all five Osmia species
studied.

Cavity selection for nesting affects the
general population structure of the spe-
cies. All studies show that selection of
long wide cavities allows females to pro-
duce more cells with larger offspring, and
more females (Stephen and Osgood 1965,
Gerber and Klostermeyer 1972, Maeta
1978, Tepedino and Parker, 1983, 1984,
Frohlich and Tepedino 1986, Tepedino
and Torchio 1989, Sugiura and Maeta
1989, Bosch 1994). In bee species that ex-
cavate a tunnel in a twig or stem and then
construct and provision linear series of
cells there is no relationship between tun-
nel diameter or tunnel length and the sex
ratio of the offspring (Garofalo et al. 1981
for Lithiirgiis, Johnson 1988 for Ceratina,
and Watmough 1983 for Xylocopa).

Why should short narrow cavities be se-
lected by any individual females? The an-
swer is variation both in body size and
cavity size. Females are selecting from an
large assortment of potential nest cavities
the cavity that best matches their body
width configuration. Selection of short
cavities is difficult to interpret. Why
should a female invest more time in for-
aging for and constructing cell partitions

and nest plugs than in offspring produc-
tion? Jayasingh and Taffe (1982) and Rust
(1993) have reported on the greater cost to
produce offspring in short cavities. Rust
(1993) has also shown that nest plugs cost
more to produce than cell partitions in O.
/. pwpinqiia and Osmia rihifloris Cresson.
He suggested that a nest should contain
four or more cells to equalize the extra
cost of nest plug. Individuals nesting in
the 80 mm length cavities were producing
on average only 2.8 cells and used signif-
icantly more mud per cell than in the oth-
er cavity length nests.

The selection of a short cavity also im-
plies that the female must spend addition-
al time searching for a second and perhaps
third cavity for nesting. Naturally occur-
ring nest sites must be considered as
clumped; beetle borings in dead trees and
logs, shrubs with hollow stems, etc. This
clumped distribution suggests that new
site searching may be minimal. Tepedino
and Torchio (1989) showed no pattern or
preference for a given diameter nest when
O. I. pwpinqiia searched for and initiated a
second or third nest.

Parasite or predator load may be a
strong selective factor favoring females
that select several different nest sites. Both
parasite and predator build-up can be-
come a serious problem with high mortal-
ity in commercial populations of cavity
nesting bees (Torchio 1970, 1972; Stephen
and Undurraga 1978; Eves et al. 1980). Fe-
males selecting one long cavity will be at
a disadvantage in a high density parasite
or predator site.

The overall immature mortality in O. I.
propinqua was low, less than 30%, and is
similar to other reports on cavity-nesting,
non-social bees and wasps (Krombein
1967, Danks 1971, Raw 1972, Cross et al.
1975, Freeman 1977, Maeta 1978, Taffee

1979, Smith 1979, Jayasingh and Freeman

1980, Tepedino and Frohlich 1982, Tepe-
dino and Parker 1983, 1984). The various
mortality agents or factors were unrelated
to either nest diameter of length in the

Volume 7, Number 1, 1998

91

present study. Chalk brood was the only
agent to showed a substantial yearly
change. Rust and Torchio (1991) also re-
ported extreme year to year variations in
chalk brood mortality within populations
of O. I. propinqua. Tepedino and Parker
(1983) reported a significantly greater
mortality due to developmental failure in
large diameter, long nests of O. marginata.
They suggest the reason to be a departure
in cell construction from a linear array of
cells to an array of cells perpendicular to
the long axis of the cavity. There was no
difference in parasite or predator attacks
in the various nests. In Hoplitis fulgida, Te-
pedino and Parker (1984) found signifi-
cantly less mortality in the short, least
used nests.

Since several species of Osmia and
Megachile rotuiidata have been developed
for commercial pollination (Torchio 1987,
1990), the choice of the appropriate cavity
size is paramount to maximize pollinator
production in a management strategy. The
economics of producing effective commer-
cial nest cavities requires the availability
of materials and tools to manufacture the
"average" cavity for a commercial popu-
lation. This cavity may not be the opti-
mum for the species. The choice of the
standard length (15 to 17 cm) drill bits and
the difficulties of obtaining wood with
grain pattern suitable for the manufacture
of "bee boards" with many straight, close,
deep holes (greater than 17 cm) resulted
in the production the commercial nest cav-
ity for O. /. propinqua of a 7 X 170 mm
paper soda straw inserted into a 8 X 170
mm hole in redwood (see Torchio 1982a,
1982b for details). This nest cavity allows
for the production of sustainable popula-
tions of O. /. propinqua for both apple and
almond pollination. My study suggests
that holes deeper than 170 mm should be
provided for O. I. propinqua for maximize
its offspring production even in a com-
mercial situation.

ACKNOWLEDGMENTS

I thank Phil Torchio and Jordi Bosch for discus-
sions that lead to this research and the development
of the manuscript. Two anonymous reviewer inputs
were most helpful and appreciated. The research
would not have been possible without the use of the
orchards and kind assistance of the Kent and Libby
Upchurch and Lynn Nardella and Jason and Sophia
Sheppard, all of Lake City, California.

LITERATURED CITED

Bosch, J. 1994. Osniii! coniutn Latr. (Hym., Mcgachili-
dae) as a potential pollinator in almond orchard.
Journal of Applied Entomology 117:151-157.

Bosch, J., M. Bias and A. Lacasa. 1992. Osmia cornuta
(Hymenoptera; Megachilidae), un nuevo polini-
zador para los almendros. Fniticiilturn Profession-
al 44:65-71.

Charnov, E. L. 1982. The theory of sex allocation. Prince-
ton University Press, Princeton, New Jersey,
USA.

Charnov, E. L., R. L. Los-den Hartogh, W. T. Jones
and J. van den Assem. 1981. Sex ratio evolution
in a variable environment. Nature 289:27-33.

Cody, R. P. and J. K. Smith. 1991. Applied Statistics
and the SAS programming language. Prentice-Hall,
Englewood Cliffs, New Jersey, USA.

Cowan, D. P. 1981. Parental investment in two soli-
tary wasps Ancistrocerus adiahatus and Euodyne-
rus foraminatus (Eumenidae: Hymenoptera). Be-
hai'orial Ecotog}/ and Sociobiology 9:95-102.

Cross, E. A., M. G. Smith and T. R. Bauman. 1975.
Bionomics of the organpipe mud-dauber, Trypox-
ylon politum (Hymenoptera: Sphecoidea). Annals
of the Entomological Society of America 68:901-916.

Danks, H. V. 1970. Biology of some stem-nesting acu-
leate Hymenoptera. Transactions of the Ro\/at Eii-
tomologi/ Society of London 122:323-399.

Danks, H. V. 1971. Populations and nesting-sites of
some aculeate Hymenoptera nesting in Rulnis.
Journal of Animal Ecology 40:63-77.

Evans, H. E. and M. J. West-Eberhard. 1970. The
waspjs. University of Michigan Press, Ann Arbor,
Michigan, USA.

Eves, J. D., D. P. Mayer and C. A. Johansen. 1980.
Parasites, predators, and nest destroyers of the
alfalfa leafcutting bee, Megachile rotundata. West-
ern Regional Extension Publication 32:1-15.

Fisher, R. A. 1958. The genetical theory of natural selec-
tion. 2nd. Dover, New York, New York, USA.

Freeman, B. E. 1977. Aspects of the regulation of size
of the Jamaican population of Sceliphron assimile
Dahlbom (Hymenoptera: Sphecidae). journal of
Animal Ecology 46:231-247.

Freeman, B. E. 1980. Parental investment, maternal
size and population dynamics of a solitar)' wasp.
American Naturalist 117:357-362.

92

Journal of Hymenoptera Research

Frohlich, D. R. and V. J. Tepedino. 1986, Sex ratio,
parental investment, and interparent variability
in nesting success in a solitary bee. Ei'clntioii 40:
142-151.

Garofalo, C. A., E. Camillo, M. J. O. Campos, R. Zuc-
chi and J. C. Serrano. 1981. Bionomical aspects of
Lithurgus conimbae (Hymenoptera, Megachili-
dae), including evolutionary considerations on
the nesting behavior of the genus. Brazilian Jour-
nal of Genetics IV 2:165-182.

Gauld, 1. and B. Bolton. 1988. Tlw Hymcuoptcra. Ox-
ford University Press, Oxford, Great Britain.

Gerber, H. S. and E. C. Klostermeyer. 1972. Factors
affecting the sex ratio and nesting behavior of the
alfalfa leaf-cutter bee. Washington Agricultural Ex-
periment Station Tcchanical Bulletin 73:1-11.

Jayasingh, D. B. and B. E. Freeman. 1980. The com-
parative population dynamics of eight solitary
bees and wasps (Aculeata; Apocrita; Hymenop-
tera) trapnested in Jamaica. Biotropica 12:214-219.

Jayasingh, D. B. and C. A. Taffe. 1982. The biology of
the eumenid mud-wasp Pachodynerus nasidens in
trapnests. Ecological Entomology 7:283-289.

Johnson, M. D. 1988. The relationship of provision
weight to adult weight and sex ratio in the soli-
tary bee, Ceratina calcarata. Ecological Entomology
13:165-170.

Johnson, M. D. 1990. Female size and fecundity in the
small carpenter bee, Ceratina calcarata (Robertson)
(Hymenoptera: Anthophoridae). Journal of the
Kansas Entomological Society 63:414-419.

Krombein, K. V. 1967. Trap-nesting wasps and bees: life
histories, nests, and associates. Smithsonian Insti-
tution, Washington, D.C., USA.

Maeta, Y. 1978. Comparative studies on the biology
of the bees of the genus Osmia of Japan, with
special reference to thier managements for pol-
linations of crops (Hymenoptera: Megachilidae).
Tohoku National Agricultural Expvriment Station
Bulletin 57:1-221.

Malyshev, S. I. 1935. The nesting habits of solitary
bees. Eos (Spain) 11:201-309.

Michener, C. D. 1947. A revision of the American spe-
cies of Hoplitis (Hymenoptera, Megachilidae).
Bulletin of the American Museum of Natural History
89:259-317.

Mitchell, T. B. 1962. Bees of the eastern United States.
Volume II. North Carolina Agricultural Experinwnt
Station Techincal Bulletin 152:1-557.

Peters, D. S. 1977. Systematik und Zoogeographie der
west-palaarktischen Arten von Osmia Panzer
1806 s. str., Monosmia Tkalcu 1974 und Orientos-
mia n. subgen. (Insecta: Hymenoptycra: Mega-
chilidae). Senckenbaergutna Bwlogia 58:287-346.

Phillips, J. K. and E. C. Klostermeyer. 1978. Nesting
behavior of Osmia lignaria propinqua Cresson
(Hymenoptera: Megachilidae). journal of the Kan-
sas Entomological Society 51:91-108.

Rau, P. 1928. Field studies in the behavior of the non-
social wasps. Transactions of the Academy of Sci-
ence of St. Louis 25:325-489.

Rau, P. 1937. The life history of Osmia lignaria and
Osmia cordata, with notes on Osmia conjuncta. An-
nals of the Entomological Society of America 39:324-
342.

Raw, A. 1972. The biology of the solitary bee Osmia
rufa (L.) (Megachilidae). Transactions of the Royal
Eutoniological Society of London 124:213-229.

Rust, R. W. 1974. The systematics and biology of the
genus Osmia, subgenera Osmia, Chalcosmia, and
Cephalosmia (Hymenoptera: Megachilidae). Wass-
inann Journal of Biology 32:1-93.

Rust, R. W. 1993. Cell and nest construction costs in
two cavity-nesting bees {Osmia lignaria propinc]ua
and Osmia ribifloris bicdermannii) (Hymenoptera:
Megachilidae). Annals of the Entomological Society
of America 86:327-332.

Rust, R. W. and P. F. Torchio. 1991. Induction of As-
cosphaera (Ascomycetes: Ascosphaerales) infec-
tions in field populations of Osmia lignaria pro-
pinqua Cresson (Hymenoptera: Megachilidae).
Pan-Pacific Entomologist 67:251-257.

Sandhouse, G. A. 1939. The North American bees of
the genus Osmia. Memoirs of the Entomological
Society of Washington 1:1-167.

SAS. 1990. SAS/stat users guuie. Volumes 1 and 2. SAS
Institute, Cary, North Carolina, USA.

Smith, A. 1979. Life strategy and mortality factors of
Sceliphron laetum (Smith) (Hymenoptera: Spheci-
dae) in Australia. Australian Journal of Ecology 4:
181-186.

Sokal, R. R. and F. J. Rohlf. 1969. Biometry. Freeman,
San Francisco, California, USA.

Stephen, W. P., G. E. Bohart and P. F. Torchio. 1969.
The biology and external morphology of bees. Oregon
Agricultural Experiment Station, Corvallis, Ore-
gon, USA.

Stephen, W. P. and C. E. Osgood. 1965. Influence of
tunnel size and nesting medium on sex ratios in
a leaf-cutter bee, Megachile rotundata. Journal of
Economic Entomology 58:965-968.

Stephen, W. P. and J. M. Undurraga. 1978. Chalk
brood disease in the leafcutting bee. Oregon State
Universiti/ Agricultural Experinwnt Station Bulletin
630.

Sugiura, N. and Y. Maeta. 1989. Parental investment
and offspring sex ratio in a solitary mason bee,
Osmia corniforns (Radoszkowski) (Hymenoptera:
Megachilidae). Japaiwse Journal of Entomology 57:
861-875.

Taffe, C. A. 1979. The ecology of two West Indian
species of mud-wasps (Euenidae: Hymenoptera).
Biological Journal of the Linnean Society 11:1-17.

Tepedino, V. J. and D. R. Frohlich. 1982. Mortality
factors, pollen utilization, and sex rat'O in Mega-
chile pugnala Say (H:ymenoptcra: Megachilidae),

Volume 7, Number 1, 1998

93

a candidate for commercial sunflower pollina-
tion, journal of the New York Entomological Society
90:269-274.

Tepedino, V. J. and F. D. Parker. 1983. Nest size, mor-
tality and sex ratio in Osinia marginata Michener.
Southwest Entomologist 8:154-167

Tepedino, V. ]. and F. D. Parker. 1984. Nest selection,
mortality and sex ratio in Hoplitis flugiila (Cres-
son) (Hymenoptera: Megachilidae). Journal of tlie
Kansas Entomological Socu-t\/ 57:181-189.

Tepedino, V. J. and P. F. Torchio. 1982a. Temporal
variability in the sex ratio of a non-social bee,
Osmia lignaria propinqua Cresson: Extrinsic deter-
mination or the tracking of an optimum? Oikos
38:177-182.

Tepedino, V. ]. and P. F. Torchio. 1982b. Phenotypic
variability in nesting success among Osmia lig-
naria propinqua females in a glasshouse environ-
ment (Hymenoptera: Megachilidae). Ecologial En-
tomology 7:453^62.

Tepedino, V. J. and P. F. Torchio. 1989. The influence
of nest-hole selection on sex ratio and progeny
size in Osmia lignaria propinqua (Hymenoptera:
Megachilidae). Annals of the Entomological Society
of America 82:355-360.

Torchio, P. F. 1970. The biology of Sapi/ga pumila Cres-
son and its importance as a parasite of the alfalfa
leafcutter bee, Megachile rotundata (Fabricius). Ar-
kansas Agricultural Experiment Station Miscella-
neous Publication 127:84-90.

Torchio, P. F. 1972. Sapyga pumila Cresson, a parasite
of Megachile rotundata (F.) (Hymenoptera: Sapy-
gidae; Megachilidae). II Methods for control. Me-
landeria 10:23-30.

Torchio, P. F. 1976. Use of Osmia lignaria Say (Hy-
menoptera: Megachilidae) as a pollinator in an
apple and prune orchard. Journal of the Kansas
Entomological Society 49:475-t82.

Torchio, P. F. 1982a. Field experiments with Osmia
lignaria propinqua Cresson as a pollinator in al-

mond orchards: II, 1976 studies, journal of the
Kansas Entomological Society 54:824-836.

Torchio, P. F. 1982b. Field experiments with the pol-
linator species, Osmia lignaria propinqua Cresson
as a pollinator in almond orchards: III, 1977 (Hy-
menoptera: Megachilidae). journal of the Kansas
Entomological Society 55:136-144.

Torchio, P. F. 1984a. Field experiments with the pol-
linator species, Osmia lignaria propinqua Cresson,
in apple orchards: III, 1977 studies, journal of the
Kansas Entomological Societ}/ 57:517-521.

Torchio, P. F. 1984b. Field experiments with the pol-
linator species, Osmia lignaria propinqua Cresson,
in apple orchards: IV, 1978 studies, journal of the
Kansas Entomological Society 57:689-694.

Torchio, P. F. 1985. Field experiments with the polli-
nator species, Osmia lignaria propinqua Cresson,
in apple orchards: V (1979-1980), methods of in-
troducing bees, nesting success, seed counts, fruit
yields (Hymenoptera: Megachilidae). journal of
the Kansas Entomological Society 58:448-464.

Torchio, P. F. 1987. Use of non-honey bee species as
pollinators of crops. Proceedings of the Entomolog-
ical Society of Ontario 18:1-32.

Torchio, P. F. 1989. In-nest biologies and develop-
ment of immature stages of three Osmia species.
Annals of the Entomological Society of America 82:
599-615.

Torchio, P. F. 1990. Diversification of pollination
strategies for U.S. crops. Environmental Entomol-
ogy 19:1649-1656.

Torchio, P. F. and V. J. Tepedino. 1980. Sex ratio,
body size and seasonality in a solitary bee, Osmia
lignaria propinqua Cresson (Hymenoptera: Mega-
chilidae). Evolution 34:993-1003.

Watmough, R. H. 1983. Mortality, sex ratio and fe-
cundity in natural populations of large carpenter
bees {Xylocopa spp.). journal of Animal Eai/n^n/ 52:
111-125.

Yasumatsu, K. and Y. Hirashima. 1950. Revision of
the genus Osmia of Japan and Korea. Mushi 21:
1-21.

J. HYM. RES.
Vol. 7(1), 1998, pp. 94-101

A New Species of the Baltic Amber Bee Genus Electrapis
(Hymenoptera: Apidae)

Michael S. Engel
Department of Entomology, Comstock Hall, Cornell University, Ithaca, NY 14853, USA

Abstract. — Electrapis stilbonota, a new species of fossil bee is described and figured from two
female specimens preserved in a single piece of Eocene Baltic amber. The species is assigned to
a new subgenus, Melikertes n. subg., characterized by the sparse hairs of the scutellum, outwardly
curved scape, few distal hamuli, absence of hind tibial spurs, tear-drop shaped tegula, and absence
of setal bands on the apices of the metasomal terga. The specimens of E. stilbonota are morpho-
logically workers and were presumably from a highly eusocial colony. The classification of Elec-
trapis among apines is briefly discussed, and the subtribe Electrapina proposed to accommodate
the genus. The proposal that Electrap>is and its presumed sister. Apis, coexisted in time is briefly
examined and found to be unsupported.

The Eocene Baltic amber contains a fas-
cinating, although uncommon, bee fauna.
Those few specimens that are known pres-
ent the picture of an assemblage of groups
unlike anything seen today. Of the species
represented in the Baltic amber only one
is currently assigned to a modern genus,
this being Andrena zvrisleyi Salt (1931), al-
though the generic assignment of this spe-
cies is of considerable question and it is
possibly a melittid (Michener and Poinar
1996). The remainder, however, are as-
signed to extinct genera whose affinities
are difficult to ascertain and in some cases
cannot confidently be placed to tribe. By
comparison, bees of the Dominican amber,
which is Oligo-Miocene in age (Grimaldi
1995), are referable to modern day genera
or extinct groups closely allied to extant
genera (Engel 1995, 1996, 1997, Michener
and Poinar 1996, Rozen 1996).

In 1909 Prof. Theodore D.A. Cockerell
described a number of Baltic amber Hy-
menoptera among which was the genus
Electrapis (1909a). The genus is a member
of the corbiculate apine tribe Apini which
contains only one other genus, the familiar
honey bees (Apis L.). Electrapis was erected

to accommodate the type species Apis me-
liponoides Buttel-Reepen (1906) which, as
the specific epithet suggests, possessed
characters both Buttel-Reepen and Cock-
erell took to be intermediate between the
Apini and their sister tribe, the Meliponini
(the stingless bees). Since the time of its
description, Electrapis has acquired a total
of nine species segregated into three sub-
genera. Table 1 summarizes the current
classification of the known species.

Herein I describe a tenth species of Elec-
trapis and assign it to a new subgenus, Me-
likertes. In the descriptions the following
abbreviations are used for morphological
terms: F, flagellomere; S, sternum; T, ter-
gum. All measurements were made using
an ocular micrometer on a WILD-M5a mi-
croscope and are in millimeters. All mea-
sures are approximate since the best po-
sition for viewing a specific structure was
not always achievable owing to the cur-
vature of the amber surface. Measure-
ments which were not possible to make
for a given specimen are indicated by an
asterisk (*). Values given in the specific de-
scription are for the holotype with the cor-
responding measure of the paratype indi-
cated in brackets.

Volume 7, Number 1, 1998

95

Table 1. Current classification of £/t'cfra;ns species.
ElL'ctnipis fnoarii (Menge) is tentatively included in the
subgenus Mclikertes.

Subgenus

Species

KfU'rtTUf

Eleclnipis s.str.

nielipoiioidcs

Buttel-Reepen 1906

Electrapis s.str.

tornquisti

Cockerell 1909b

Electrapis s.str.

apciites

Manning 1960

Electrapis s.str.

ijiiinitn

Kelner-Pillault 1970

Electrapis s.str.

hoinhusohlcs

Kelner-Pillault 1974

Protohvnbiis

inciecisiis

Cockerell 1909a

Protobomhus

tristcllus

Cockerell 1909a

Roiiss}/ana

pmlmiiickeuensis

Roussy 1937

Melikcrtes

stilhonota

present study

7

proava

Menge 1856

Genus Electrapis Cockerell
Melikertes Engel, new subgenus

Diagnosis. — Roussyana-like species with-
out dense pubescence covering the scutel-
lum; hairs generally sparse, never obscur-
ing integument. Clypeus flat. Labrum U-
shaped, broader than long, with apical
fringe of simple hairs, hairs laterally short,
becoming longer by middle. Labial palpus
four segmented, basal segment longest, al-
most as long as following three segments
combined. Minute flabellum at apex of
glossa. Antennae set well below mid-line
of face; scape slightly curved outwards at
apex, inner concave surface without pu-
bescence; pedicel longer than Fl; Fl as
long as F2 and F3 combined. Compound
eyes bare. Face relatively flat. Vertex
scarcely elevated above ocelli; preoccipital
ridge with a weak carina behind vertex,
becoming rounded by gena (Fig. 3). Meso-
scutal anterior border weakly rounded,
nearly straight; median line and notauli
not apparent, parapsidal lines faintly evi-
dent. Tegula tear-drop shaped, with blunt
apex pointing posteriorly. Scutellum not
bulging, surface flat, not reaching back to
obscure portions of the metanotum; ante-
rior border nearly straight medially, pos-
terior border broadly rounded. Strigular
concavity set on a slightly protuberant
shelf; velum and malus simple, velum not
divided. Mesocoxae separated medially.

Hind tibia without inner apical spurs; cor-
bicula comprising apical three-quarters of
hind tibia, posterior apical border round-
ed; corbicular surface glabrous and not
strongly depressed, with a few sparsely
scattered long simple hairs, otherwise
hairs restricted to tibial lateral borders,
such hairs long and branched; inner sur-
face of tibia with a dense medial field of
extremely short, simple hairs; strong ras-
tellum on inner apical surface of hind tib-
ia, extending the full width of tibial apex;
penicillum absent; hind basitarsus rough-
ly quadrangular, about as broad at base as
at apex; attachment to tarsomere 2 set
forth on short distal process on anterior
border; auricle present; inner surface of
hind basitarsus with series of stiff setal
rows (as in Apis); claw with minute tooth
on lower third of irmer margin. Pterostig-
ma small; distal wing venation strong;
posterior border of second submarginal
cell extended posteriorly (Fig. 4), greatly
narrowed anteriorly; basal vein basad cu-
a, vein relatively straight; cu-a curved out-
wards, not orthogonal with Cu or A; less
than 10 hamuli on anterior margin of hind
wing; jugal lobe present, just over one half
length of vannal lobe; jugal and vannal in-
cisions shallow. Metasoma without distal
rows of hair on terga (Fig. 3).

Ti/pe species. — Electrapis (Melikertes) stilhon-
ota Engel, new species, present designation.

Etymology. — ^The subgeneric name is taken
from the mythology of ancient Crete. Meli-
kertes (meaning "honey-cutter") was associ-
ated with Corinth and Melissa, priestess to
the mother-goddess Demeter.

Remarks. — Electrapis proava is possibly mis-
placed in Roussyana and should be included
in Melikertes. Official transfer of this species
will have to wait, however, until £. proava
can be studied in more detail.

Electrapis (Melikertes) stilhonota
Engel, new species

(Figs. 1-5)

Description. — FEMALE (male un-
known): Body form Trigona-\\ke. Total

96

Journal of Hymenoftera Research

^0S--

Figs. 1-2. Eleclrapis (McUkcrtes) stilbouota new species. 1, holotype, dorsal view. 2, paratype, lateral view.

body length 3.76 [3.72]. Head wider than
long (length 1.10 [1.12], width * [1.32]). In-
ner margins of compound eyes straight,
nearly parallel; upper interorbital distance
0.84 [0.86]; lower interorbital distance *
[0.82]; eye length 0.74 [0.76], width 0.32
[0.30]. Gena width 0.28 [0.24]. Mandible
with two blunt denticles on upper half of
apical margin, length 0.50 [0.44]; malar
space length 0.04 [0.06]. Labrum length
(median) 0.20 [0.24], width (basal) 0.44
[0.44]; clypeus length * [0.18], width *
[0.64]; clypeoantennal distance 0.08 [0.08].
Scape length 0.36 [0.36]; pedicel length
0.10 [0.06]; flagellum length 0.88 [0.86]; Fl
length 0.10 [0.10]; F2 length 0.04 [0.04]; F3
length 0.04 [0.04]; FIO length 0.18 [0.18];
F6-10 with dense sensillar plates on inner
surfaces; interantennal distance * [0.20];
antennal-ocellar distance 0.58 [0.56]. Me-
dian ocellus diameter 0.12 [0.12]; distance
between lateral ocelli 0.26 [0.26]; distance
between median ocellus and lateral ocel-

lus 0.08 [0.08]; distance from lateral ocel-
lus to eye 0.28 [0.28]; distance from lateral
ocellus to occiput 0.16 [0.16]. Mesosoma
length 1.34 [1.32]; intertegular distance
0.82 [0.80]; mesoscutum length 0.64 [0.60];
scutellum length 0.26 [0.28], width 0.48
[0.44]; metanotum length 0.04 [0.04]; pro-
podeal triangle over four times longer
than metanotum, length 0.18 [0.18]. Me-
socoxae separated by more than mesocox-
al width; hind tibia length (median) 1.02
[1.02], width (basal) 0.12 [0.12], width (api-
cal) 0.28 [0.30]; hind basitarsus length 0.36
[0.32], width 0.28 [0.28]. Wings hyaline, all
veins brown and strong; basal vein basad
cu-a by 2 times vein width; pterostigma
small; Im-cu bisecting second submargin-
al cell; 2r-m distad 2m-cu by 1.5 times vein
width; marginal cell length 0.84 [0.86],
width 0.24 [0.22]; first submarginal cell
shorter than second and third combined;
length of anterior border of second sub-
marginal cell one-tenth that of posterior

Volume 7, Number 1, 1998

97

border; length of anterior border of third
submarginal cell half of that of posterior
border, just over 3 times length of anterior
border of second submarginal cell; fore-
wing length 3.00 [3.16]; venation of fore-
wing depicted in figure 4; six distal ham-
uli on outer margin of hind wing; cu-a of
hind wing orthogonal to M + Cu; hind
wing length 2.08 [2.16]; venation of hind
wing depicted in figure 5. Metasoma
length 1.32 [1.28].

Integument over entire bee smooth and
glabrous, except on metanotum where the
integument is apparently rugulose. S3-6
apparently weakly nodulate, nodules scat-
tered over the surface, integument be-
tween nodules smooth and shining as on
previous sterna. Color not well preserved,
apparently dark brown to black, metallic
and shining, without any maculations.

Pubescence generally pale. Hairs of face
widely scattered, simple, and short. Such
hairs becoming longer by vertex. Gena with
simple, short, suberect hairs. Postgena with
long, simple hairs sparsely scattered over
integument. Pronotal collar without pubes-
cence; pronotal border with mesoscutum
with short, simple hairs; lateral surface with
similar minute hairs, such hairs appressed
to surface, not obscuring integument. Me-
soscutum with scattered simple hairs, more
sparsely scattered and shorter over central
disc, those hairs on anterolateral borders
with a few short branches. Scutellum like
that of mesoscutum except pubescence lon-
ger and restricted to posterior border. Meta-
notum with dense, minute, simple hairs, not
obscuring the surface. Hypoepimeron with-
out pubescence; mesepistemum with scat-
tered simple hairs, becoming longer ven-
trally, central disc, however, without hairs.
Propodeal triangle and posterior surface
without pubescence; lateral surface with
scattered long simple hairs and shorter, ap-
pressed hairs, partially obscuring the sur-
face. Pubescence of fore- and midlegs gen-
erally simple and scattered, except inner
surfaces of midtrochanter and femur with-
out pubescence, and outer surface of mid-

tibia with dense, branched hairs. Inner sur-
face of hind femur and trochanter without
pubescence, except apical quarter of femur
with dense field of minute hairs similar to
those on inner surface of hind tibia (see ge-
neric diagnosis). Eight comb rows on inner
surface of hind basitarsus; outer surface
with scattered, long, simple hairs. Tl with-
out hairs over central disc, a few simple
hairs on lateral borders. T2 as on Tl, except
a few simple hairs on posterolateral bor-
ders. T3 with simple hairs, longer than
those of Tl-2, sparsely scattered over cen-
tral disc, more concentrated on lateral mar-
gins. T4-6 similar to T3. Sterna with sparse-
ly scattered simple hairs.

Material examined. — Holotype: female
(Figs. 1 & 3), worker, Samland, Eocene
Baltic amber, specimen In. 17778, Depart-
ment of Palaeontology, the Natural His-
tory Museum (British Museum, London).
Paratype: female (Figs. 2, 4-5), worker,
same piece of amber and same accession
information as holotype.

Preservation. — The bees presented herein
are exceptionally well preserved. The only
hindrance to their examination is the un-
even surface of the amber and the small
block of storage media they are preserved
in. A few small fracture planes arising
from the wings do not obscure any im-
portant structures, although one small
fracture near the face of the holotype spec-
imen obscures some features of the clyp-
eus and lower face. A bit of mold on the
legs along with the remains of what might
have been collected pollen in the corbicu-
la, while slightly demoting them from per-
fect specimens, does not detract in any sig-
nificant way from examining their mor-
phology. The inner teeth of the claws are
minute and difficult to see. The best view
of these structures can be achieved by
back-lighting the specimens and examin-
ing the extended hind legs.

Etymology. — The specific epithet is de-
rived from stilbo (Gr. shine) and noton (Gr.
back), and is a reference to the glabrous

98

Journal of Hymenoptera Research

Figs. 3-5. Electrapis (Melikerks) stilbonota new species. 3, close-up of holotype, dorsal view, showing integ-
ument of metasoma and propodeal triangle as well as vertex and preoccipital ridge. 4, left forewing of para-
type. 5, left hind wing of paratype.

integument of the dorsum, in particular
that of the propodeal triangle.

DISCUSSION

Bees of the tribes Apini and Meliponini
are all advanced eusocial, except for a few
parasitic forms among the stingless bees
(Michener 1974), and, based on their sis-
ter-group relationship (Chavarria and
Carpenter 1994), presumably inherited
this aspect of their biology from a com-
mon ancestor that was similarly eusocial.
The phylogenetic position of the genus
Electrapis within the tribe Apini suggests
that species of Electrapis were also highly
eusocial with a well developed caste sys-
tem. Also suggestive of eusociality among
Electrapis species is the fact that the spec-
imens described herein are morphologi-
cally workers. As in many eusocial species
the loss of ovarian development in the
worker caste results in a greatly reduced
metasoma, a feature seen in both speci-

mens. Similar lines of evidence were used
to make the inference that the oldest
known fossil bee, Trigoua prisca, was a
worker of a similarly advanced eusocial
society (Michener and Grimaldi 1988a, b).
While Electrapis runs to the tribe Apini
in Michener's (1990) key to the corbiculate
bee tribes (treated as subfamilies of Api-
dae in that work), there are significant
enough differences between Electrapis and
its sister Apis which would more than jus-
tify placing Electrapis in a tribe of its own.
Recognition of a separate tribe for these
bees, however, would obscure the rela-
tionship of Electrapis with members of the
Apini as both possess a jugal lobe, bifid
claws, complete distal wing venation, and
a marginal cell apex gently pulled away
from the anterior wing margin. A more
practical approach to the problem is the
recognition of subtribes, retaining a
broadly defined Apini. The separation of

VoLUMK 7, Number 1, 1998

99

Table 2. Brief summary of the subtribal classification of Apini Latreille (based on worker caste). An elab-
oration of each character is given in the text.

Apin.1

Eye hairs:
Labral apex:
Mandible:
Vertex:
Scutellum:
Propodeum:
Mesocoxae:
Marginal cell:
Marginal cell:
Basal vein:

present

concave

without dentition

short

bulging

short, dcclivious

nearly meeting

reaching wing apex

not tapering

distad cu-a

HIeclrj

Lipinj

absent

convex

with or without dentition

long

weakly convex or flat

long, not declivious

well separated

not reaching wing apex

gently tapering

basad to just distad cu-a

these subgroups is as follows (a summary
of the differences is given in Table 2):

Electrapina (new subtribe containing
only the typical genus Elect rapis): Com-
pound eyes without hairs; labral apex con-
vex; mandible with or without dentition;
malar space extremely short, much less
than basal width of mandible; vertex as
long as ocellar diameter, or more; scutel-
lum not bulging, surface weakly convex to
flat; propodeal triangle with defined sur-
face, not declivious; mesocoxae separated
by at least their width; marginal cell not
reaching to wing apex, gently tapering
over its length; basal vein basad to just
distad cu-a, never strongly distad (7 times
vein width or more).

Apina: Compound eyes covered with
long hairs; labral apex gently concave;
mandible lacking dentition; malar space as
long as, or longer than basal width of
mandible; vertex extremely short, much
less than ocellar diameter; scutellum
strongly convex and bulging, obscuring
metanotum and propodeal triangle; pro-
podeal triangle extremely short and decli-
vious; mesocoxae nearly meeting medial-
ly; marginal cell long, nearly reaching
wing apex, not gently tapering over its
length; basal vein confluent (in some fossil
Apns) to strongly distad cu-a (over 7 times
vein width), never basad cu-a.

Arillo et al. (1996) have recently sug-
gested that Apis and Electrapis overlapped
in geologic time. Specimens of Electrapis

are only known from the Baltic amber
which is Eocene in age (Kosmowska-Cer-
anowicz 1987, Kosmowska-Ceranowicz
and Miiller 1985) while Apis species are
unknown until the middle Oligocene (Cul-
liney 1983, Engel in press, Michener 1990,
Ruttner 1988, Zeuner and Manning 1976).
Thus, the little available evidence in no
way suggests that these taxa were coinci-
dent in time. Arillo et al. (1996) are correct,
however, in their assertion that there is no
reason to believe Electrapis is the direct an-
cestor of the true honey bees. Apis, as has
been done by some earlier authors (e.g.,
Statz 1931, Zeuner and Manning 1976).
Lastly, these authors have peculiarly used
invalid family-group names for bees. For
example, they refer to Michener (1986) for
the recognition of Rophitidae in place of
Halictidae. In fact, Michener (1986) clearly
advises the use of Halictidae (even in his
fairly short abstract), a proposal which
was later supported by Michener (1991)
and validated by the International Com-
mission on Zoological Nomenclature
(1993). Therefore, these authors should not
be followed in their use of family-group
names for bees.

ACKNOWLEDGMENTS

I am thankful to A. J. Ross for hosting me during
my visit to the Department of Palaeontologv, the Nat-
ural History Museum (British Museum, London), and
for bringing these specimens to my attention. J. K.
Liebherr kindly allowed use of his camera-micro-
scope mount for the production of the plates. I am

100

Journal of Hymenoptera Research

grateful to G. Chavarria, E. E. Grissell, and J. G. Roz-
en, Jr., for kindly reading the manuscript prior to
publication. Their constructive criticisms helped im-
prove the final draft. Support for this work was pro-
vided by an Ernst Mayr Award from Harvard Uni-
versity's Museum of Comparative Zoology and by a
National Science Foundation Predoctoral Fellowship.

LITERATURE CITED

Arillo, A., A. Nel, and V. M. Ortuiio. 1996. Two fossil
bees from the Oligocene of Izarra (Alava, Spain)
(Hymenoptera, Apoidea). Bulletin de la Societe en-
tomohgique de France 101:59-64.
Buttel-Reepen, H. v. 1906. Apistica. Beitrage zur Sys-
tematik. Biologic, sowie zur geschichtlichen und
geographischen Verbreitung der Honigbiene
(Apis mellifica L.), ihrer Varietaten und der iiber-
igen Apis-Arten. Mitteilungen aus dem Zoologisch-
en Museum in Berlin 3:117-201.
Chavarria, G., and J. M. Carpenter. 1994. "Total evi-
dence" and the evolution of highly social bees.
Oadistks 10:229-258.
Cockerell, T. D. A. 1909a. Descriptions of Hymenop-
tera from Baltic amber. Sclmften der PJn/sikalisch-
okonomischen Ceiellschaft 50:1-20.
Cockerell, T. D. A. 1909b. Some additional bees from
Prussian amber. Schriften der Physikalisch-dkon-
oinisclien Gesellschaft 50:21-25.
Culliney, T. W. 1983. Origin and evolutionary history

of the honeybees Apis. Bee World 64:29-38.
Engel, M. S. 1995. Neocorynura electra, a new fossil bee
species from Dominican amber (Hymenoptera:
Halictidae). journal of the Neu< York Entonwlogical
Society 103:317-323.
Engel, M. S. 1996. New augochlorine bees (Hyme-
noptera: Halictidae) in Dominican amber, with a
brief review of fossil Halictidae. journal of the
Kansas Entomological Society, Supplement 69:334-
345.
Engel, M. S. 1997. A new fossil bee from the Oligo-
Miocene Dominican amber (Hymenoptera: Hal-
ictidae). Apuiologie 28:97-102.
Engel, M. S. In press. Fossil honey bees and evolution
in the genus Apis (Hymenoptera: Apidae). Api-
dologie.
Grimaldi, D. A. 1995. The age of Dominican amber,
pp. 203-217. In Anderson, K.B., and J.C. Crelling,
eds.. Amber, Resinite, and Fossil Resins. American
Chemical Society Symposium Volume, Washing-
ton, D.C.
I.C.Z.N. 1993. Opinion 1713: Some bee family-group
names (Insecta, Hymenoptera): names based on
Colletes Latreille, 1802, on Paracolletes Smith,
1853, on Halictus Latreille, 1804, on Anthuiium Fa-
bricius, 1804 and on Anthoptiora Latreille, 1803
given precedence over some senior names. Bul-
letin of Zoological Nomenclature 50:85-89.
Kelner-Pillault, S. 1970. L'ambre Balte et sa faune en-

tomologique avec description de deux apoides
nouveaux. Annates de la Societe entomologique de
France 6:3-24.
Kelner-Pillault, S. 1974. Etat d'evolution des apides
de l'ambre balte. Annates de la Societe entomolo-
gique de France 10:623-634.
Kosmowska-Ceranowicz, B. 1987. Mineralogical-pet-
rographic characteristics of the Eocene amber-
bearing sediments in the area of Chlapowo, and
the Palaeogene sediments of Northern Poland.
Biuletyn Instytut Geologii 356:29-50.
Kosmowska-Ceranowicz, B., and C. Miiller. 1985. Li-
thology and calcareous nannoplankton in amber
bearing Tertiary sediments from boreholes, Chla-
powo (Northern Poland). Bulletin of the Polish
Academy of Sciences 33:119-129.
Manning, F. J. 1960. A new fossil bee from Baltic am-
ber. Proceedings of the 11"' International Congress of
Entomology, Vienna 1:306-308.
Menge, A. 1856. Lehenszeichen vortu'eltUcher im Bern-
stein eingeschlossener Thiere. Programm Petrischu-
ler Danzig.
Michener, C. D. 1974. The social behavior of the bees: a
comparative study. Harvard University Press,
Cambridge.
Michener, C. D. 1986. Family-group names among
bees. Journal of the Kansas Entomological Society 59:
219-234.
Michener, C. D. 1990. Classification of the Apidae
(Hymenoptera). University of Kansas Science Bul-
letin 54:75-164.
Michener, C. D. 1991. Case 2535. Proposed prece-
dence of some bee family-group names (Insecta,
Hymenoptera): names based on Colletes Latreille,
1802, on Paracolletes Smith, 1853, on Halictus La-
treille, 1804, on Anthidium Fabricius, 1804 and on
Anthophora Latreille, 1803 to have precedence
over some senior names. Bulletin of Zoological No-
menclature 48:227-235.
Michener, C. D., and D. A. Grimaldi. 1988a. A new
Trigona from Late Cretaceous amber of New Jer-
sey (Hymenoptera: Apidae: Meliponinae). Amer-
ican Museum Novitates 2917:1-10.
Michener, C. D., and D. A. Grimaldi. 1988b. The old-
est fossil bee: apoid history, evolutionary stasis,
and antiquity of social behavior. Proceedings of the
National Academy of Sciences. USA 85:6424-6426.
Michener, C. D., and G. O. Poinar, Jr. 1996. The
known bee fauna of the Dominican amber, four-
nal of the Kansas Entomological Society, Supplement
69:353-361.
Roussy, L. 1937. Contribution a I'etude de I'abeille
tertiaire de ses parasites et de ses ennemis. Ga-
zette apicole 388:49-72.
Rozen, J. G., Jr. 1996. A new species of the bee Hct-
erosariis from Dominican amber (Hymenoptera:
Andrenidae; Panurginae). journal of the Kansas
Entomological Society, Supplement 69:346-352.

Volume 7, Number 1, 1998

101

Ruttner, F. 1988. Biogeography ami taxonomy of honey-
bees. Springer- Verlag, Berlin.

Salt, G. 1931. Three bees from Baltic amber. Bernstein-
Forscluingen 2:136-147.

Statz, G. 1931. Eine neue Bienenart aus Rott am Sie-
bengebirge: ein Beitrag zur Kermtnis der fossilen

Honigbienen. Wissenschaftlichen Milleihingen ties
Vereinsfiir Natiir- und Heimatkunde, Koln 1:39-60.
Zeuner, F. E., and F. ]. Manning. 1976. A monograph
on fossil bees (Hymenoptera: Apoidea). Bulletin
of the British Museum of Natural History (Geology)
27:149-268.

NOTE ADDED IN PROOF

A paper has recently reached me concerning a Middle-Eocene bee from Germany which is
attributable to Electrapis [H. Lutz. 1993. Eekfeldapis eleetrnpotdes nov. gen. n. sp., eine "Honigbiene"
aus dem Mittle-Eozan des "Eckfelder Maares" bei Manderscheid/Eifel, Deutschland (Hymenop-
tera: Apidae, Apinae). Maimer naturwissensclniftliches Arehiv 31:177-199]. This bee is clearly a
species of Electrapis s. str. and, based on the wing venation, appears most similar to £. apoides.
I, therefore, here synonymize Eekfeldapis (new synonymy) with Electrapis, and place its only
included species as a species of the latter: Electrapis (Electrapis) electrapoides (Lutz), new combi-
nation. It must also be noted that Lutz's figure 3h, labeled as the wing venation of Apis inellifcra,
should be disregarded as it resembles very little the venation of this species (particularly in the
position of the basal vein and cu-a).

J. HYM. RES.
Vol. 7(1), 1998, pp. 102-115

Neotropical Eucoilidae (Cynipoidea) Associated with Fruit-infesting

Tephritidae, with New Records from Argentina,

Bolivia and Costa Rica

R. A. Wharton, S. M. Ovruski, and F. E. Gilstrap

(RAW, FEG) Texas A&M University, Department of Entomology, College Station, TX 77843-
2475, USA; (SMO) CIRPON, C. C. C. 90 4000, San Miguel de Tucuman, Argentina

Abstract. — Host and distribution records are presented for five species of Neotropical Eucoilidae
(Hymenoptera: Cynipoidea) reared in association with fruit-infesting Tephritidae. All previously
recorded tephritid host associations for New World species are critically reviewed with the con-
clusion that several of these records are doubtful. Members of the genera Aganaspis and Odonto-
sema are confirmed as parasitoids of Anastrepha and Ceratitis. Based on isolated puparia, Dicera-
taspis and Lopheucoila are recorded as parasitoids of Drosophilidae and Lonchaeidae, respectively.
It is suggested that Dicerataspis is unlikely to attack Tephritidae, and records of Lopheucoila from
Tephritidae require confirmation. One new species reared from Tephritidae, Aganaspis nordlan-
deri Wharton, is described.

The Eucoilidae are solitary endoparasi-
toids that oviposit in the larval stage of
cyclorrhaphous Diptera and emerge as
adults from the host puparium. Several
eucoilid species have been implicated as
important natural enemies of different
phytophagous dipteran species (Wishart
and Monteith 1954, Harding 1965, Valla-
dares et al. 1982, Johnson 1987), and two
species have been used for the biological
control of fruit fly pests (Clausen 1978).
The Asian species Aganaspis daci (Weld)
has been introduced to the New World,
and the Neotropical endemic Aganaspis
pelleranoi (Brethes) was reared and re-
leased from 1941 to 1945 in several areas
of Tucuman, Argentina (Nasca 1973). At
present, A. pelleranoi is being mass-pro-
duced in Metapa de Dominguez, Chiapas,
Mexico (Ruiz et al. 1996).

Members of the genus Aganaspis are the
only eucoilids thus far utilized in biologi-
cal control efforts against Tephritidae. The
two species involved, A. daci and A. pel-
leranoi, are also the only eucoilids attack-
ing tephritids for which biological infor-

mation other than host records has been
published. Different aspects of the basic
biology of A. daci were studied in the lab-
oratory associated with programs directed
against Bactrocera dorsalis (Hendel) in Ha-
waii (Clausen et al. 1965) and Anastrepha
suspensa (Loew) in Florida (Nunez-Bueno
1982). This species has also been intro-
duced to Mexico (Jimenez-Jimenez 1956)
and Costa Rica (Wharton et al. 1981, Jiron
and Mexzon 1989). Establishment in Mex-
ico and Costa Rica is doubtful, but in Flor-
ida it is established on Anastrepha suspensa
(Loew), though in low numbers (Bara-
nowski et al. 1993). A detailed biology of
A. pelleranoi was given by Ovruski (1994a,
1994b).

The aim of this note is to provide pre-
liminary information on the diversity of
eucoilid species associated with tephritid
fruit flies in the Neotropics, and clarify the
status of species previously recorded as te-
phritid parasitoids. While several species
have been associated with tephritids, few
of these have been reared from puparia
that were sufficiently isolated to enable

Volume 7, Number 1, 1998

103

verification of the host. Data presented
here are based on surveys of tephritid par-
asitoids and on literature records.

MATERIALS AND METHODS

Fruits damaged by tephritid larvae
were collected from 1991 to 1994 in Tu-
cuman, Catamarca and La Rioja provinces
in northvv'estern Argentina, and from Au-
gust, 1979 through November, 1982 in the
provinces of Alajuela, Cartago, Guana-
caste, Heredia, Limon, Puntarenas, and
San Jose in Costa Rica. Additional eucoilid
specimens were also received from Boliv-
ia, and all reared material housed in the
U. S. National Museum of Natural Histo-
ry, Washington, D. C. (USNM) was ex-
amined. Samples collected in Argentina
and Costa Rica consisted of fallen fruit
and fruit still on the tree. In Argentina,
fruit samples were placed in styrofoam
boxes with damp sand in the bottom as a
pupation substrate. Fruit fly puparia were
recovered weekly and transferred to a
closed wooden box for holding until emer-
gence of flies or parasitoids. The proce-
dure differed slightly for the samples from
Costa Rica (Wharton et al. 1981), where
81,279 puparia were isolated (most of
them in individual vials) for verification of
host records. Tephritids of the genera
Anastrepha Schiner and Ceratitis MacLeay
(or their parasitoids) accounted for 69,012
of these puparia, with C. capitata repre-
senting 64.8% of the total tephritids. Lon-
chaeidae (also discussed below) were rep-
resented by 4583 puparia.

Specimens reported on here are housed
at Museo de La Plata, Argentina (MLP),
Museo de Ciencias Naturales Bernardino
Rivadavia, Buenos Aires, Argentina
(MBR), Instituto Fundacion Miguel Lillo,
Tucuman, Argentina, Texas A&M Univer-
sity, College Station (TAMU), and USNM.
Measurements for the description of the
new species are as described by Nordlan-
der (1978, 1982).

RESULTS AND DISCUSSION

Several species of Eucoilidae, represent-
ing at least five genera, have been reared
in association with fruit-infesting tephri-
tids. Most of the species are almost cer-
tainly attacking other Diptera associated
with ripe and decomposing fruit (e.g. Dro-
sophilidae, Lonchaeidae, Phoridae, Neri-
idae). We have verifiable host records
from Tephritidae for Agauaspis and Odon-
tosetna. The published records for Dicera-
taspis, Lopheucoila, Rhoptoiiicris, and Tri/-
bliographa attacking tephritids in the New
World need confirmation. Species of the
genus Leptopilina, well-known parasitoids
of Drosophilidae, may also be reared com-
monly from rotting fruit. Though Droso-
philidae tend to colonize fruit after te-
phritids, fallen, broken fruit may have
more rapidly decaying portions inhabited
by Drosophilidae at the same time as more
sound portions still inhabited by tephritid
larvae. Since fruit is often collected and
reared in bulk, it is easy to obtain parasit-
oids of both Drosophilidae and Tephriti-
dae, for example, from the same sample.
Unless puparia are isolated individually,
correct host associations cannot be made.

These seven genera may be separated
by the characters in Table 1. Additionally,
Lopheucoila is unique within this group of
genera in having a small spine on the dor-
sal plate of the scutellum and longitudinal
ridges on the mesoscutum. See also papers
by Weld (1952), the updated classification
by Nordlander (1978, 1980, 1981) and the
description of Aganaspis by Lin (1987).

Agartaspis Lin

The genus Aganaspis was relatively re-
cently described (Lin 1987) to accomodate
four species from southeast Asia (Taiwan
and Malaysia). One of these species, A.
daci (Weld), was originally described in
Tn/hliographa (Weld 1951b) but its generic
placement had always been problematic
(Kerrich and Quinlan 1960, Nordlander
1981). Aganaspis daci is the only one of the

104

Journal of Hymenoptera Research

Table 1. Genera of Eucoilidae reported from fruit-infesting Tephritidae in the Neotropics compared with
Lqjtopili}ia, parasitoids of Drosophilidae commonh' reared from the same fruits.

Scuteilar disc
posteriorly in
dorsal view

Fore wing

Posterior-dorsal

margin of
pronota! plate

4th antennal segment

{2nd flagellomere)

of male

Anterior and posterior
parts ot pronotal

plate fused or
separate laterall\'

Agciimspis

Dicerataspis

Lopheucoila

Odontoscma

Rhoptrcineris

TrybUographa

LL'ptopilinn

rounded or
truncate or
weakly bi-
lobed

bifurcate, with 2
tooth-like
lobes directed
posteriorly

weakly dentate,
with 4 lobes;
somewhat
truncate

distinctly setose

distinctly setose

bare or nearly
so

bifurcate, with 2 bare or nearly
tooth-like so

lobes directed
posteriorly

rounded or
truncate

distinctly setose

rounded or
truncate

distinctly setose

rounded or
truncate

distinctly setose

protruding
above anteri-
or margin of
mesoscutum;
deeply con-
cave medially

weakly protrud-
ing above an-
terior margin
of mesoscu-
tum; undu-
lant, with 4
rounded
lobes

protruding
above anteri-
or margin of
mesoscutum;
deeply con-
cave medially

protruding
above anteri-
or margin of
mesoscutum;
deeply con-
cave medially

not protruding
above anteri-
or margin of
mesoscutum;
evenly round-
ed or very
weakly con-
cave medially

not protruding
above anteri-
or margin of
mesoscutum;
evenly round-
ed or very
weakly con-
cave medially

not protruding
above anteri-
or margin of
mesoscutum;
evenly round-
ed or very
weakly con-
cave medially

3 not bent

4 < 3 not bent

4 < 3 not bent

4 < 3 not bent

4 > 3 bent out-
wardly

4 > 3 usually
bent

widely to nar-
rowly sepa-
rated

contiguous or
narrowly sep-
arated

fused or contig-
uous

fused or contig-
uous

fused

4 £ 3 not bent fused

widely separat-
ed '

Volume 7, Number 1, 1998

105

four originally included species for which
hosts have been recorded. Nordlander (in
lift.) suggested that the New World spe-
cies pellemiwi should also be placed in
Aganaspis. This transfer was made by
Ovruski (1994a), bringing the total num-
ber of species in Aganaspis to five. All four
of the Old World species have distinctly
setose eyes (more noticeable in the fe-
male), while those from the Neotropics do
not. This is one of the easiest ways to rec-
ognize A. daci in those areas of the New
World where it has been introduced.

The placement of pelleranoi and nonilan-
deri, n. sp. (described below) in Aganaspis
broadens the limits of this genus, and
opens up the possibility that several of the
Neotropical species formerly placed in ei-
ther Ttybliographa or Pseudeiicoila may ac-
tually belong here. As indicated by Nor-
dlander (1981, Table 2), several generic
names are available for these species, and
placement of most of the previously de-
scribed species will not be possible with-
out a revision of the entire group. Collec-
tion records (e.g.. Weld 1932) and label
data on specimens in the USNM suggest
that several of these species have been
reared from tephritids, but most records
are not sufficiently precise to preclude the
possibility that the actual hosts may be
other fruit-inhabiting flies. See further dis-
cussion below under Tryhliographa.

Aganaspis pelleranoi (Brethes)

De Santis (1965) placed Ganaspis carvalhoi
Dettmer, 1929 as a junior subjective syn-
onym of Eucoila pelleranoi Brethes, 1924. The
senior author has confirmed this synonymy
through comparison of one of Dettmer's
syntypes in USNM with one of the syntypes
of pelleranoi from the Brethes collection
(MBR). Both types match the specimens we
reared from Argentina and Costa Rica, con-
firming their identity as pelleranoi.

Known hosts and distribution records
of A. pelleranoi are as follows:

Hosts. — A. ludens (Loew), A. obliqua
(Macquart) (Aluja et al. 1990), A. serpentina

(Wiedemann) (Costa Lima 1940), A. striata
Schiner (Clausen 1978), A. distincta Greene
(Katiyar et al. 1995), A. fraterculus (Wie-
demann) (Brethes 1924), Ceratitis capitata
(Wiedemann) (De Santis 1965), and Rhag-
oletis turpiniae Hernandez-Ortiz (Hernan-
dez-Ortiz 1993). The records from "Lon-
chaea sp." and "L. pendula Bezzi" (Borg-
meier 1935 and Costa Lima 1948, respec-
tively) are suspect, and require
verification. As noted by McAlpine and
Steyskal (1982), the name pendula has been
misapplied on numerous occasions to
Neotropical lonchaeids of the genus Neo-
silba McAlpine that have been reared from
fruit. Our own records suggest that A. pel-
leranoi may only rarely attack lonchaeids.
Of 295 specimens of A. pelleranoi that we
reared from isolated puparia in Costa
Rica, 191 (64.7%) were from C. capitata
(mostly in coffee), 81 from Anastrepha
(nearly all A. striata), and only two were
from a lonchaeid (Neosilba hatesi (Curran),
new record). Twenty-one other specimens
of Aganaspis were also reared from Neosil-
ba in Costa Rica, but these are only tenta-
tively assigned to pelleranoi because of
slight but consistent color differences rel-
ative to the other specimens of A. pelleranoi
reared during these studies.

We reared A. pelleranoi from infested
fruits of the following species: Casimiroa
ediilis Llave and Lex., Citrus aurantiifolia
(Christm.) Swingle, Coffea arabica L., Ficus
carica L., Juglans australis Grisebach, Pru-
nus domestica L., Prunus persica (L.) Batsch,
Psidium guajaim L., P. friedrichsthalianum
(O. Berg) Niedenzu, P. littorale Raddi
( = cattleianum), Syzygium jambos (L.) Al-
ston, and Terminalia catappa L. These data,
together with previously published re-
cords, suggest that A. pelleranoi, like sev-
eral of the other commonly encountered
tephritid parasitoids, has little or no host
plant preferences. See additional com-
ments below under discussion of Odonto-

sema.

Distribution. — Argentina: Buenos Aires,
Misiones, Salta, Tucuman, Jujuy, Corrien-

106

Journal of Hymenoptera Research

tes (De Santis 1967, Diaz 1986), La Rioja
and Catamarca (new records); Bolivia:
Santa Cruz de La Sierra (new record);
Peru (Clausen 1978); Brazil (Dettmer
1929); Venezuela (Katiyar et al. 1995); Co-
lombia (Yepes and Velez 1989); Costa Rica
(Wharton et al. 1981); El Salvador (Ovru-
ski et al. 1996); and Mexico (Aluja et al.
1990). There are also specimens in the
USNM from Panama, Guatemala, and Be-
lize (all new records).

During the survey for tephritid parasit-
oids in Costa Rica (Wharton et al. 1981),
several species resembling A. pelleraiwi
were reared. Only one of these was re-
peatedly reared from tephritids, and it is
described next.

Aganaspis nordlanderi Wharton, new
species

(Figs. 1, 3, 4, 6-8, 10, 12)

Quantitative measurements, based on 5
females and 2 males, are presented either
as ranges or means to the nearest 0.05.

Female (Fig. 1). — Body length 2.3-3.5
mm; fore wing length 2.3-3.1 mm.

Head in dorsal view 1.05-1.15 times
wider than mesoscutum; 1.75-1.90 times
wider than long, when length measured in
profile; temples strongly receding in dor-
sal view. Frons, vertex, and occiput bare;
eyes without visible setae at 50 x. Face
shining, unsculptured; malar sulcus a
weak, narrow groove, without additional
striae; antennal base elevated, especially
laterally, forming shallow depression be-
tween antemia and eye. Posterior ocelli
widely separated: distance between them
1.2-1.3 times distance between posterior
ocellus and eye. Antenna (Fig. 4) short,
about 2.5 times height of head; without
distinct club, the segments gradually
broadening distally, first 4-5 flagellomeres
weakly clavate: each slightly broader sub-
apically than medially; first flagellomere
slightly shorter than second, relative
lengths, first 8 flagellomeres: 1.0:1.1:1.1:
1.05:1.0:1.0:0.95:0.95; ratio of length to
maximum width, first 8 flagellomeres: 2.4:

2.3:2.2:2.0:1.9:1.8:1.65:1.55; flagellomeres
each with 3-5 whorls of setae.

Mesosoma 1.25 times longer than high;
1.6 times longer than wide; 1.3 times high-
er than wide. Pronotal collar (Figs. 1, 6)
distinctly protruding above anterior mar-
gin of mesoscutum in lateral view, deeply
bilobed; median bridge wider than ante-
rior ocellus; lateral arms of anterior and
posterior parts narrowly but distinctly
separated; posterior part of pronotal plate
with a few, completely decumbent setae,
otherwise bare and polished, 2.3-2.8 times
wider than median bridge. Mesoscutum
divided into three parts of approximately
equal width by two longitudinal rows of
5-6 decumbent setae per row; margin
with scattered setae separated from each
other by their own length; short, shallow,
crescentic grooves present on posterior
half directly anteriad lateral bars of scu-
tellum. Scutellar disc (Figs. 7, 8, 10) dis-
tinctly reticulate, with dorsoposteriorly-
directed setae somewhat longer than in
pelleranoi; in dorsal view (Fig. 8) disc
slightly excavated medially, and thus
weakly bilobed, the lobes protruding
slightly beyond cup; posterior margin of
disc sinuate in profile; height of posterior
margin of scutellum about 1.5 times
length (in lateral view) of flat portion of
cup; scutellar cup (Fig. 8) large, broadly
tear-drop shaped, with posterior margin
weakly rounded, nearly truncate, anterior
margin short, not extending through scu-
tellar fovea, surface nearly flat, not droop-
ing posteriorly, shallowly excavated me-
dially, with only one or two punctures
and 2^ short, erect setae laterally; width
of cup 0.5-0.6 times width of disc. Fore
wing (Fig. 12) 2.6-2.9 times longer than
wide; marginal setae short, longest sub-
apical seta 0.08-0.1 times maximum width
of wing; radial cell deep, completely open
along wing margin, second radial abscissa
2.7-3.1 times longer than first; costal cell
densely setose, ventral surface with 3-4
longitudinal rows of setae. Mid and hind
coxae (Fig. 1) with extensive patches of

Volume 7, Number 1, 1998

107

Figs. 1-5. Agimaspna spp. 1, A. nonilaiideri, habitus. 2, A. pcUcranoi, mid and hind coxae, lateral view, showing
small patches of dense setae dorsoposteriorly. 3, A. nordlanderi, male antenna, setae not shown except for 8th
flagellomere. 4, A. nordlanderi, female antenna, setae not shown except along margins of 7th and 8th flagel-
lomeres. 5, A. pelleranoi, basal 5 antennal segments of female.

dense setae, patch on mid coxa extending
about half length of coxa from base, patch
on hind coxa extending more than half
length of coxa.

Second metasomal tergum 1.0-1.1 times
length of mesosoma.

Male. — As in female except as follows:
head in dorsal view 1.15-1.2 times wider
than mesosoma; antenna (Fig. 3) long, 3.2-
3.3 times height of head; relative lengths,
first 5 flagellomeres: 1.0:0.8:0.9:0.95:0.95,
ratio of length to maximum width, flagel-
lomeres 1-5 and 10: 2.85:2.05:2.25:2.25:2.3:
2.6; mesosoma 1.2 times longer than high;

fore wing 3.6-3.75 times longer than wide;
second tergum 0.85-0.9 times length of
mesosoma. Color. Head and mesosoma
black except pedicel and basal flagellom-
eres usually dark reddish brown, flagel-
lomeres gradually darkening apically,
more rarely with antenna entirely dark
brown to black; gaster reddish-orange, ex-
cept apical terga black posteriorly; legs
red-brown (slightly browner than gaster).
Hosts. — The specimens forming the type
series were reared from individually iso-
lated puparia of Ceratitis cnpitntn and Anas-
trepha striata, and the puparia from which

Figs. 6-11. Aganaspis spp, mesosoma. 6, A. nordlandcn, pronotum in dorsal view. 7, A. nordlanderi, propodeum
in posterior view, showing posterior portion of scutellum, setae not shown except portion of basal ring on
metasoma. 8, A. nordlanderi, scutellum in dorsal view. 9, A. pelleranoi. scutellum in dorsal view. 10, A. nor-
dlanderi, .scutellum in lateral view. 11, A. pelleranoi. scutellum in lateral view.

108

Journal of Hymenoptera Research

Volume 7, Number 1, 1998

109

Figs. 12, 13. Agaiuispis, fore wings. 12, A. nordlandcn,
showing setal pattern in costal cell, marginal setae,
and shape of radial cell, other setae not shown. 13, A.
fieUeranoi, showing relatively reduced pattern of setae
in costal cell and closed radial cell.

they emerged are pinned with the parasit-
oids.

Holoiype female. — "Costa Rica: Cartago
Turrialba, Catie 10.vii.l980 ex: guava R.
Chavez." Reared from Anastrepha striata in
guava. Deposited in USNM. Paratypes
(TAMU), all reared from Ceratitis capitata
in Costa Rica: 3 females, 2 males, same lo-
cality, 8-vi-1980, 26-vi-1980, 30-ix-1981,
and 8-vi-1982, J. Duran and R. Chavez,
collectors, from coffee and naranja agria; 1
female, Puntarenas, San Vito, 23-ix-1980,
R. Chavez, from guava.

Diagnosis. — This species differs from pel-
leraiwi in the more densely setose costal
cell of the fore wing (with only a single
longitudinal row of setae on ventral sur-
face in pelleraiwi: compare Figs. 12 and 13),
the completely open radial cell, the more
extensively furry hind coxa (Fig. 1 vs. Fig.
2), the smaller scutellar cup (width of
cup/ width of disc = 0.55 in nordlamieri vs.
0.75 in pelleraiwi; scutellar cup extending
nearly to anterior margin of scutellar fo-
vea in pelleranoi: compare Figs. 8 and 10
with Figs. 9 and 11), and the relative pro-
portions of the flagellomeres (Fig. 4 vs.
Fig. 5) and first and second radial abscis-
sae.

Discussion. — Aganaspis nordlanderi has a
distinctly smaller scutellar cup than either
pelleranoi or daci, and the surface is not

quite as flat as in these two other species.
The first flagellomere of the female is also
slightly shorter than the second in nordlan-
deri but slightly longer than the second in
pelleranoi and daci. Otherwise, nordlanderi
and pelleranoi share several features which
suggest that they are more closely related
to each other than either is to daci and the
other Old World species of Aganaspis de-
scribed by Lin (1987). This relationship be-
tween the New World species is based on
the absence of distinct setae on the eyes,
the shape of the scutellar disc which pro-
trudes posteriorly beyond the cup, the
lack of a deep, median depression in the
posterior margin of the metapleuron, and
the more elongate basal flagellomeres and
less distinctive club of the female antenna.
Until the genera occurring in the Neotrop-
ical Region become better known, we pre-
fer to retain pelleranoi and nordlanderi in
Aganaspis, with the full realization that
this placement may need to be reconsid-
ered at some later date.

Dicerataspis Ashmead

There are two described species, and
Weld (1952) recorded an additional, ap-
parently undescribed species from Flori-
da. The genus is known from Mexico and
Florida south through the Caribbean to
Brazil and Argenhna (Weld 1921, 1952,
Diaz 1974). The species have not been re-
vised, and most of the reared material in
the USNM has been tentatively associated
with the name D. grenadensis Ashmead,
1896. At least two species are represented
in our material from Costa Rica. The spe-
cies differ in the sculpture of the scutel-
lum, infumation of the wing, and shape of
the radial cell. A revision of the genus is
needed before species names can be as-
signed with any degree of confidence.

Based on records in the USNM, largely
from the rearings by Zetek in the Panama
Canal Zone, members of this genus have
been reared from a variety of fruits (Ficiis,
Labatia, Psidiiitn, Carica, and Anacardium)
in association with several different spe-

110

Journal of Hymenoptera Research

cies of the tephritid genus Anastrepha. Di-
cerataspis has also been associated with
one species of Rhagoletis from Mexico
(Hernandez-Ortiz 1993). However, precise
associations have rarely if ever been made.
Our own material was reared in associa-
tion with Anastrepha fraterculus and Anas-
trq}ha sp. from Psidium guajaim (guava) in
Tucuman, Argentina, and from Droso-
philidae in peaches and guava in Costa
Rica. The single specimen from guava in
Costa Rica was reared from a carefully
isolated drosophilid puparium and repre-
sents the first specific, verified host asso-
ciation to our knowledge. All of our other
material was batch-reared, and could not
be directly associated with the individual
puparia from which they emerged. The
specimens from guava in both Costa Rica
and Argentina agree most closely with the
description of D. grenadensis.

Given the small size of the species of
Dicerataspis and their development as sol-
itary parasitoids, normal hosts are likely
to be drosophilids or other small Diptera
in fruit, rather than tephritids. Parasitism
of Anastrepha or other tephritids needs
verification.

Lopheucoila Weld

Weld (1951a) described the genus Lo-
pheucoila for three New World species,
only one of which, L. anastrephae (Roh-
wer), has ever been associated with spe-
cific hosts. Lopheucoila anastrephae was
originally described from specimens sup-
posedly reared from Anastrepha sp. in
Trinidad (Rohwer 1919). Weld (1951a)
subsequently reported this species from A.
fraterculus in Panama and Lonchaea sp. in
Brazil, as well as from Mexico and Peru
(unassociated with hosts). As noted above
under the discussion of hosts of A. peller-
anoi, the Brazilian record from Lonchaea
undoubtedly refers to a species of Neosilba.
All of the specimens of Lopheucoila reared
from Argentina and Costa Rica during the
present study were L. anastrephae, based
on comparison with type material of the

three described species housed in the
USNM.

In Argentina, L. anastrephae was ob-
tained from guava fruits in association
with A. fraterculus and Anastrepha spp. In
Costa Rica, it was reared from coffee, gua-
va, papaya, sour orange (Citrus aurantium
L.), and tangerine (Citrus reticulata Blan-
co), and only from Lonchaeidae. All 74
specimens collected in Costa Rica were
reared from isolated puparia of Neosilba
batesi (Curran), the most abundant of the
two species of Lonchaeidae in the Costa
Rican samples. Although 69,000 puparia
of Anastrepha and Ceratitis were collected
from 1979-1982 in Costa Rica, L. anastre-
phae was never reared from a tephritid
during this period.

Odontosema Kieffer

Kieffer (1909) based the genus Odonto-
sema on a single species from Brazil. Borg-
meier (1935) later described a second bra-
zilian species, O. anastrephae Borgmeier,
collected in association with A. fraterculus
from guava. One undescribed species has
been recorded from A. striata and A. fra-
terculus in guava in Veracruz, Mexico
(Hernandez-Ortiz et al. 1994), and an un-
identified Odontosema species was reared
from A. fraterculus in fruit of four species
of Myrtaceae in southern Brazil (Salles
1996). In Costa Rica, O. anastrephae has
been reported from C. capitata and Anas-
trepha spp. inhabiting several different
fruits (coffee, orange and guava) (Wharton
et al. 1981). Odontosema is probably widely
distributed throughout the Neotropical
Region, but at present it has only been re-
corded from Brazil (Borgmeier 1935, Costa
Lima 1948, Salles 1996), Costa Rica (Whar-
ton et al. 1981) and Mexico (Hernandez-
Ortiz et al. 1994, Lopez et al. 1996).

Odontosema anastrephae was collected
only in Costa Rica, and was not found in
Argentina during the present study. This
species showed distinct host preferences,
with 74% of the 193 reared individuals
coming from Anastrepha in guavas. An ad-

Volume 7, Number 1, 1998

111

ditional 15.5% of the reared individuals
were from C. capitata in citrus. Coffee, the
most heavily sampled fruit in Costa Rica,
yielded only four individuals. Only 2% of
the individuals were reared from Lon-
chaeidae (all on citrus), with the remain-
ing 98% on either Anastrepha or Ceratitis.
These results are in direct contrast with
those for A. pelleraiioi, which was collected
predominantly from C. capitata in coffee in
Costa Rica. Slightly less than half as many
A. pelleranoi were reared from Anastrqjha
in guava as from Ceratitis in coffee. The
pattern of host utilization by A. pelleranoi
can thus be directly correlated with sam-
pling frequency, with the mostly com-
monly sampled fruits and tephritids yield-
ing the greatest numbers of pelleranoi.

Rhoptromeris Forster

Only one species of this genus has been
associated with tephritid fruit flies. This
species, R. hai/zoardi (Blanchard 1947), was
originally included in Eucoila and later
transferred to Rhoptromeris (De Santis
1967). R. hayzvardi was described from Ar-
gentina and Uruguay (Blanchard 1947)
and has been reared in association with A.
fraterculus and C. capitata (De Santis 1967),
as well as Anastrepha spp., from Carica
quercifolia Hill, Ficus carica, Phoebe porphyr-
ia Gris, Psidium j^uajava, and Primus persica
in Tucuman (Turica and Mallo 1961). De-
spite intensive surveys of fruit fly parasit-
oids that were conducted in several areas
of the province of Tucuman for this and
related studies (Ovruski 1995) the host fly
and host plant associations recorded by
Turica and Mallo have not been verified.

Nasca et al. (1980) obtained exception-
ally large numbers of R. hai/wardi and the
diapriid Trichopria anastrephae Costa Lima
(nearly 3,500 and 1,000 individuals respec-
tively) by using a modified model of Hay-
ward's (1940) parasitoid fly-trap. This col-
lecting method consisted of a pit in the
soil beneath the host plant into which fall-
en host-fruit were placed, the pit was then
covered with a thin sieve permitting only

the capture of parasitoids. It is very likely
this method facilitated the production of
drosophilids and their parasitoids. For ex-
ample T. anastrephae is also known to at-
tack drosophilids (Turica and Mallo 1961).
This, together with our failure to rear hai/-
wardi from tephritid puparia in the prov-
ince of Tucuman, lead us to question the
recorded host associations of this species.
Parasitism of Tephritidae needs to be ver-
ified.

The generic placement of this species
also needs verification following Nordlan-
der's (1978) revision of Rhoptromeris. As
restricted by Nordlander (1978), Rlioptrom-
eris consists of parasitoids of Chloropidae
and other small dipterous larvae such as
those that commonly breed in the base of
grass stems (Poaceae), and it is unlikely
that haifwardi belongs to this largely Hol-
arctic genus.

Trybliographa Forster

Several eucoilids associated with fruit-
infesting Diptera in the Neotropical Re-
gion have been placed in the genus Try-
bliographa at one time or another. These in-
clude species placed in Pseudeucoila, a ju-
nior synonym of Trybliographa (Hellen
1960, Nordlander 1980), as well as various
subgenera of either Trybliographa or Pseu-
deucoila (Weld 1952). Three of these, in-
cluding the species originally described as
Trybliographa daci by Weld (1951b), are
treated above under the genus Aganaspis.
Most of the others, however, cannot be
readily assigned to genus at the present
time because the Neotropical genera are
badly in need of revision. These species
appear to belong to what Nordlander
(1982) referred to as the Ganaspis group of
genera. Nordlander (1981, 1982) discussed
some of the problems associated with the
Trybliographa and Ganaspis groups of gen-
era, presented a list of generic names that
he considered valid, and provided useful
information on the location and status of
the type material. According to Nordlan-
der (1981), Trybliographa is predominantly

112

Journal of Hymenoptera Research

Holarctic. It is therefore unlikely that any
of the species previously associated with
fruit-infesting Diptera in the neotropics
belong to this genus. We have examined
several such specimens in the USNM col-
lection of Tri/bliograplia and Pseudeucoila
labelled as reared from tephritids or other
fruit-infesting flies, and were unable to
discover any that belong to Trybliographa
s. s. as defined by Nordlander (1981).
Aside from the species now transferred to
Aganaspis, we are aware of only two other
species that have been described, and for
which tephritid hosts have been reported
(additional records available to us are
largely in the form of unpublished label
data). These are bmsiliensis von Ihering,
1905 and hookeri Crawford, 1913.

Von Ihering's species was first de-
scribed as Hexamerocera brasiliensis and lat-
er as Eiicoela (Hexamerocera) eobrasiliensis
(von Ihering 1914). It was transferred,
along with the subgenus Hexamerocera, to
Pseudeucoila by Weld (1932). Nordlander
(1978), however, treated Hexamerocera as a
synonym of Rhoptromeris. Pseudeucoila, as
noted above, is now a synonym of Try-
bliographa. Von Ihering's brasiliensis does
not fit the current definition of either
Rhoptromeris or Trybliographa, and thus,
like haywardi, remains unplaced in the Eu-
coilidae.

Von Ihering's brasiliensis is known from
Brazil and Panama (Borgmeier 1935) and
it was introduced to Puerto Rico during
1935-37 along with other unidentified eu-
coilid species to aid in the control of A.
obliqua and A. suspensa (Bartlett 1941). The
recorded hosts were A. fraterculus, Anas-
trepha sp., C. capitafa and Drosophilidae
(Borgmeier 1935, Costa Lima 1948). The
type material was collected from peaches,
where it was thought to be a parasitoid of
A. fraterculus (von Ihering 1905). Though
von Ihering (1912) provides evidence to
support his view of brasiliensis as a para-
sitoid of Anastrepha, the actual host was
not identified and the possibility that this
is a drosophilid parasitoid (because of its

small size) cannot be discounted. Hosts
for this species thus need verification.

Crawford's hookeri was originally de-
scribed in Ganaspis, but was placed under
Hexamerocera by Weld in his arrangement
of the USNM collection (though formal
transfer to Pseudeucoila (Hexamerocera) was
apparently never published). This is a
similarly diminutive species, and there-
fore unlikely to be a parasitoid of Anastre-
pha, as originally recorded. Both hookeri
(from Puerto Rico) and brasiliensis have a
complete hairy ring at the base of the sec-
ond tergum, unlike similarly-sized species
of Leptopilina, in which the second tergum
is bare dorso-medially. The scutellar cup
is much larger in brasiliensis than it is in
hookeri.

CONCLUSIONS

Information on host specificity in eu-
coilids is largely lacking other than for the
work by van Alphen, Vet and colleagues
on species of Leptopilina attacking Droso-
philidae (e.g. van Alphen et al. 1991, Pool-
man Simons et al. 1992). There are often
many species of Diptera, representing sev-
eral families, present in fruit attractive to
eucoilids. Thus, in order to assess host
specificity accurately, it is essential to ver-
ify all records by isolating individual pu-
paria or exposing known hosts to ovipos-
iting females. For many of the older re-
cords (including label data from unpub-
lished studies), host associations were
based on eucoilids reared from bulk sam-
ples of fruits containing pest tephritids.
These must be viewed with caution be-
cause of the inevitable inclusion of other
flies, such as drosophilids and lonchaeids,
in these samples.

ACKNOWLEDGMENTS

We are grateful to the following people for the loan
of types and other specimens: Arnold Menke and Da-
vid Smith (U.S.D.A., Systematic Entomology Labo-
ratory, Washington, D.C.) and A. Roig-Alsina (MBR).
We are also indebted to Goran Nordlander for con-
siderable assistance in helping us to understand the
classification of Eucoilidae. S. Ovruski is deeply

Volume 7, Number 1, 1998

113

grateful to N. B. Diaz (Facultad de Ciencias Naturales
y Museo de La Plata, Argentina: MLP) for identifying
the Dicercitnfpns species from Argentina, C. Pruett
(Universidad Autonoma Gabriel Rene Moreno, Santa
Cruz, Bolivia) for sending the specimens from Boliv-
ia, P. Fidalgo (Instituto Fundacion Miguel Lillo), N.B.
Diaz and L. De Santis (MLP) for supplying important
bibliographic material. Finally, S. Ovruski wishes to
thank Consejo Nacional de Investigaciones Cientifi-
cas y Tecnicas de la Republica Argentina for its con-
stant support. R. Wharton and F. Gilstrap are most
grateful to P. Krauter for organizing the database for
the Costa Rican collections and providing printouts
for host associations, and to M. Fischel, J. Duran, and
R. Chavez for their crucial role in conducting the field
work in Costa Rica and isolating individual puparia.
R. Wharton is also most appreciative of the assistance
provided by C. Yoshimoto in making available notes
and other information from L. Weld. Illustrations
were provided by B. Flahey.

LITERATURE CITED

Aluja, M., J. Guillen, P. Liedo, M. Cabrera, E. Rios, G.
de la Rosa, H. Celedonio, and D. Mota. 1990.
Fruit infesting tephritids (Diptera: Tephritidae)
and associated parasitoids in Chiapas, Mexico.
E)itoinofJhagn 35: 39-48.

Baranowski, R., H. Glenn, and ]. Sivinski. 1993. Bio-
logical control of the Caribbean fruit fly (Diptera:
Tephritidae). Florida Entomologist 76: 245-251.

Bartlett, K. A. 1941. The introduction and coloniza-
tion in Puerto Rico of beneficial insects parasitic
on west indian fruit flies. The journal of Agricul-
ture of the Unit'ersily of Puerto Rico 25: 25-31.

Blanchard, E. 1947. Insectos del Uruguay. Comunica-
cionei Zoologicas, Museo tie Historia Natural de
Montevideo 2(42): 11.

Borgmeier, T. 1935. Sobre alguns cynipidos parasiti-
cos e cecidogenos do Brasil (Hymenoptera: Cy-
nipidae). Archivos Instituto Biologico y Vegetal, Rio
de Janeiro 2: 97-124.

Brethes, J. 1924. Varios himenopteros de la America
del Sur. Nunqunm Otiosus 2: 2-16.

Clausen, C. P. 1978. Tephritidae. Pp. 320-325. In: C.
P. Clausen (Ed.) Introduced parasites and predators
of arthropod pests and weeds: a world review. United
States Department of Agriculture, Agriculture
Handbook 480.

Clausen, C. P., D. W. Clancy and Q. C. Chock. 1965.
Biological control of the oriental fruit fly {Dacus
dorsalis Hendel). United States Department of Ag-
riculture Technical Bulletin 1322: 102 pp.

Costa Lima, A. da. 1940. Alguns parasites de moscas
de frutas. Analcs da Academia Brasdeira de Scwncias
12; 17.

Costa Lima, A. da. 1948. Entomofagos sul amcricanos
(parasites e predadores) de insetos nocivos a

agricultura. Boletim de Sociedade Brasiliera de
Agronomia, Rio de Janeiro 11: 1-82.

De Santis, L. 1965. Nota sobre un parasito de la mosca
sudamericana de la fruta (Hymenoptera: Cyni-
pidae). Rei'ista de la Sociedad Entomolgica Argen-
tina 27: 73-74.

De Santis, L. 1967. Catdlogo de los himenopteros argen-
tinos de la serie Parasitica, incluyendo Bethyloidea.
Publicacion de la Comision de Investigacion
Cientifica de la Provincia de Buenos Aires 337

PP-

Dettmer, H. 1929. Ganaspis carvalhoi n. sp. (Hymenop-
tera: Cynipidae), um novo parasita da mosca das
frutas {Anastrepha fraterculus Wied.). Boletim
Biologica, Sao Paulo 16: 70-74.

Diaz, N. B. 1974. Anotaciones sobre cinipoideos ar-
gentinos. I. Neotropica 20: 17-20.

Diaz, N. B. 1986. Ampliacion de la distribucion geo-
grafica de cinipoideos parasitoides en la Repiib-
lica Argentina y Brasil. Revista de la Sociedad En-
tomoldgica Argentina 44: 32.

Harding, J. A. 1965. Parasitism of the leaf miner Lir-
iomi/za munda in the winter garden area of Texas.
journal of Economic Entomology 58: 442^143.

Hayward, K. ]. 1940. Lucha biologica contra las mos-
cas de las frutas. Dispositivo que permite la sal-
ida de los parasitos beneficos del pozo donde se
arroja la fruta atacada. Revista Industrial Agri'cola
de Tucumdn 10-12: 230-233.

Hellen, W. 1960. Die Eucoilinen Finnlands (Hym.
Cyn.). Fauna Fennica 9: 1-30.

Hernandez-Ortiz, V. 1993 Description of a new Rhag-
oletis species from tropical Mexico (Diptera: Te-
phritidae). Proceedings of the Entomological Society
of Washington 95: 418-424.

Hernandez-Ortiz, V., R. Perez-Alonso, and R. A.
Wharton. 1994. Native parasitoids associated
with the genus Anastrepiha (Dipt.: Tephritidae) in
Los Tuxtlas, Veracruz, Mexico. Entomophaga 39:
171-178.

Jimenez-Jimenez, E. 1956. Las moscas de la fruta y
sus enemigos naturales. Fitdfito 16: 4-11.

Jiron, L. F. and R. G. Mexzon. 1989. Parasitoid hv-
menopterans of Costa Rica: geographical distri-
bution of the species associated with fruit flies
(Diptera: Tephritidae). Entomophaga 34: 53-60.

Johnson, M. 1987. Parasitization of Lirumiyza spp.
(Diptera: Agromyzidae) infesting commercial
watermelon plantings in Hawaii, journal of Eco-
nomic Entomology 80: 56-61.

Katiyar, K. P., J. Camacho, F. Geraud, and R. Ma-
theus. 1995. Parasitoides hvmenopteros de mos-
cas de las frutas (Diptera: Tephritidae) en la re-
gion occidental de Venezuela. Revista de la Facul-
tad de Agronoma (LUZ) 12: 303-312.

Kerrich, G. J. and J. Quinlan. 1960. Studies on eucoi-
line Cynipoidea (Hym.). Opuscuta Entonwlogica
25:179-196.

114

Journal of Hymenoptera Research

Kiefter, ]. J. 1909. Description de nouveaux cynipides
zoophages. BuUctiu dc In Societe d'Histoire Natii-
rclle, Metz (3)2: 8-96.

Lin, K. S. 1987. Aganaspis, a new genus of Eucoilidae
(Hymenoptera: Cynipoidea). Taiwan Agricultural
Rt'scarf/i Institute, Special Puhlicaticn 22: 67-79.

Lopez, M., ]. Sivinski, L. Guillen, C. Ruiz, J. Pinero,
and M. Aluja. 1996. Reservorios de parasitoides
de moscas de la fruta (Diptera: Tephritidae) en
el estado de Veracruz, Mexico. In: 2nd Meeting
of the working group on fruit flies of the western
hemisphere. Via del Mar, Chile, 1996: 70.

McAlpine, J. F. and G. C. Steyskal. 1982. A revision
of Neosilba McAlpine with a key to the World
genera of Lonchaeidae (Diptera). The Canaiiian
Entomologist 114: 105-137.

Nasca, A. J. 1973. Parasites de "moscas de los frutos"
establecidos en algunas zonas de Tucuman. Re-
vista Agn'cola del Noroeste Argentino 10: 31—13.

Nasca, A. J., R. V. Fernandez, D. R. Sosa, D. Fernan-
dez, and A. De Guerrero. 1980. Eficiencia del
pozo trampa en el manejo de enemigos naturales
de moscas de los frutos. Adas del 2 ° Congreso Na-
clonal de Citricultura, Entre Rios 2: 141-162.

Nordlander, G. 1978. Revision of the genus Rhoptrom-
eris Forster, 1869 with reference to north-western
European species. Studies on Eucoilidae (Hym.:
Cynipoidea) II. Entonwiogica Scandinnvica 9: 47-
62.

Nordlander, G. 1980. Revision of the genus Lepioptliua
Forster, 1869, with notes on the status of some
other genera (Hymenoptera, Cynipoidea: Eucoil-
idae). Entomologica Scandinavica 11: 428^53.

Nordlander, G. 1981. A review of the genus Tryblio-
grapha Forster, 1869 (Hymenoptera, Cynipoidea:
Eucoilidae). Entomologica Scandinavica 12: 381-
402.

Nordlander, G. 1982. Identities and relationships of
the previously confused genera Odontcucoila, Co-
neucoila and Trichoplasta (Hym., Cynipoidea: Eu-
coilidae). Entomologica Scandinavica 13: 269-292.

Nunez-Bueno, L. 1982. Trybliographa daci Weld (Hy-
menoptera: Cynipidae): biology and aspects of
the relationship with its host Anastreplm suspensa
(Loew) (Diptera; Tephritidae). Ph. D. Disserta-
tion, University of Florida: 152 pp.

Ovruski, S. M. 1994a. Comportamiento en la detec-
cion del huesped de Agaiuispis pclkranoi (Hyme-
noptera: Eucoilidae) parasitoide de larvas de Cer-
atitis capitata (Diptera: Tephritidae). Revista de la
Sociedad Entomologica Argentina 53: 121-127.

Ovruski, S. M. 1994b. immature stages of Aganaspis
pclleranoi (Brethes) (Hymenoptera: Cynipoidea:
Eucoilidae), a parasitoid of Ceratitis capitata
(Wied.) and Anastrepha spp. (Diptera: Tephriti-
dae). Journal of H\/menoptera Research 3: 233-239.

Ovruski, S. M. 1995. Pupal and larval-pupal parasit-
oids (Hymenoptera) obtained from Anastrepha

spp. and Ceratitis capitata (Dipt.: Tephritidae) pu-
pae collected in four localities of Tucuman prov-
ince, Argentina. Entoniopliaga 40: 367-370.

Ovruski, S. M., S. Fuentes, F. Nunez, and J. G. Gra-
nados Zuniga. 1996. Himenopteros parasitoides
de moscas de la fruta (Diptera: Tephritidae) pre-
sentes en la Republica de El Salvador. Revista de
Ingenieros Agronomos de El Salvador 14: 8-14.

Poolman Simons, M. T. T., B. P. Suverkropp, L. E. M.
Vet, and G. de Moes. 1992. Comparison of learn-
ing in related generalist and specialist eucoilid
parasitoids. Entomologia Experimeutalis et Appli-
cata 64: 117-124.

Rohwer, S. A. 1919. Description of a new cynipoid
from Trinidad. Proceedings of the Entomological So-
ciety of Washington 21: 156.

Ruiz, S. L., J. L. Cancino D., and M. Aluja. 1996. Co-
lonizacion de parasitoides nativos para el control
biologico de moscas de la fruta. In: 2nd Meeting
of the working group on fruit flies of the western
hemisphere, Via del Mar, Chile, 1996: 70.

Salles, L. A. 1996. Parasitismo de Anastrepha fratercu-
lus (Wied.) (Diptera: Tephritidae) por Hymenop-
tera na regiao de Pelotas, R.S. Pesquisa Agrope-
cuaria Brasileira, Brasilia 31: 769-774.

Turica, A. and R. G. Mallo. Observaciones sobre la
poblacion de las "Tephritidae" y sus endopar-
asitos en algunas regiones citricolas argentinas.
IDIA 6: 145-161.

Valladares, G., N. Diaz, and L. De Santis. 1982. Tres
notas sobre dipteros agronomicidos de la Repub-
lica Argentina y sus himenopteros parasitoides.
Revista de la Sociedad Entomolgia Argentina 42:319-
330.

Van Alphen, J. J. M., G. Nordlander, and I. Eijs. 1991.
Host habitat finding and host selection of the
Drosophila parasitoid Lcptopilina australis (Hyme-
noptera, Eucoilidae), with a comparison of the
niches of Eurcipean Leptopilina species. Oecologia
87:324-329.

Von Ihering, R. 1905. As moscas das frutas e sua des-
trui^ao. 1° edi^ao. Secretaria de Agricultura, Sao
Paulo. Pp. 12-13.

Von Ihering, R. 1912. As moscas das frutas e sua des-
trui^ao. 2° edii^ao. Typographia Brazil, Sao Paulo.
48 pp.

Von Ihering, R. 1914. Diagnose de uma Eucoela, par-
asita das moscas. Revista Museo Paulista 9: 224-
225.

Weld, L. H. 1921. Notes on certain genera of parasitic
Cynipidae proposed by Ashmead with descrip-
tions of genotypes. Proceedings of the United States
National Museum 59: 433-451.

Weld, L. H. 1932. Synonymical and descriptive note
on Pseudeucoila hrasiUensis (R. v. Ihering, 1905)
(Hym., Cynipidae), Revista de Entomologia 2: 24-
27.

Weld, L. H. 1951a. New Eucollinae (I hnienoptera,

Volume 7, Number 1, 1998

115

Cynipoidea). Pwceeditigs of the Entomological So-
ciety of Wmtnngton 53: 223-226.

Weld, L. H. 1951b. A new species of Tryhliografilui
(Hymenoptera: Cynipidae). Proceedings of the Ha-
umiinn Entomological Society 14: 331-332.

Weld, L. H. 1952. Cynipoidea (Hym.) 1905-1950. Ann
Arbor, Privately Printed.

Wharton, R. A., F. E. Gilstrap, R. H. Rhode, M. Fis-
chel-M., and W. G. Hart. 1981. Fiymenopterous
egg-pupal and larval-pupal parasitoids of Cera-
titis capitata and Anastrepha spp. (Diptera: Te-

phritidae) in Costa Rica. Entomophaga 26: 285-
290.

Wishart, G. and E. Monteith. 1954. Tryhliographa rapae
(Westw.) (Hymenoptera: Cynipidae), a parasite
of Hylemya spp. (Diptera: Anthomiidae). The Ca-
nadian Entomologist 86: 145-154.

Yepes, R. and R. Velez. 1989. Contribucion al cono-
cimiento de las moscas de las frutas (Tephritidae)
y sus parasitoides en el departamento de Antio-
quia. Rei'ista de la Facultad Nacional de Agronoini'a
de Medellin 42: 73-98.

J. HYM. RES.
Vol. 7(1), 1998, pp. 116-117

NOTE

First Chromosome Record for the Family Dryinidae: The Karyotype
of Anteon brevicome Dalman (Hymenoptera: Chrysidoidea)

Vladimir E. Gokhman and Kirill A. Kolesnichenko
Botanical Garden, Moscow State University, Moscow 119899, Russia

Though karyotypes of some members of
the superfamily Chrysidoidea have been
examined during the last few years (Ho-
shiba and Imai 1993, Gokhman and
Quicke 1995, Quicke and Gokhman 1996),
the family Dryinidae remains totally un-
touched by chromosomal investigation.
We have studied for the first time chro-
mosome number and karyotype of the
dryinid, Anteon brevicome Dalman. Chro-
mosome preparation was obtained from
an adult wasp collected from the wild at
the Botanical Garden, Moscow State Uni-
versity, Moscow, Russia, in May 1997.
Preparation was made according to the
previously described protocol (Gokhman
and Quicke 1995). Chromosomes were
subdivided into four groups — metacen-
trics, submetacentrics, subtelocentrics and
acrocentrics following Levan et al. (1964)
and Imai et al. (1977). The voucher speci-
men is deposited in the Zoological Muse-
um, Moscow State University, Moscow,
Russia.

RESULTS

Eleven well-spread metaphase plates
were obtained from the individual stud-
ied, all of them having the same diploid
chromosome number, 2n = 10 (Fig. 1). All
chromosomes are obviously two-armed
and thus arm number (NF) in this species
is 20. The karyotype comprises three pairs
of submetacentric chromosomes and two
pairs of subtelocentric ones. However, all
chromosomes differ notably in size, each
chromosome pair being at least about 1.5

times longer or shorter than the others.
Submetacentrics of the first two pairs are
the longest in the set (3^ jxm), those of
the third pair are the shortest (0.5 ixm),
and subtelocentrics are of intermediate
length (1-2 ixm).

DISCUSSION

The above results, together with accu-
mulated data on chromosomes of the oth-
er Chrysidoidea, provide qualitatively
new karyotypic information for the super-
family. First, chromosome number of A.
brevicome is the lowest in the Chrysidoi-
dea and one of the lowest in all aculeate
Hymenoptera. Except for a few ant spe-
cies, only four predominantly unrelated
members of the Aculeata (although two of
them belong to the bee genus, Andrena)
were reported to have n values of 5 or
fewer (Goodpasture 1974, Hoshiba and
Imai 1993). Second, the chromosome set of
A. brevicome is highly asymmetric (White
1973), apart from karyotypes of the other
Chrysidoidea, where chromosomes show
a continuous gradation in length (see for
example Hoshiba and Imai 1993, Fig. 2c
and 8c, and Quicke and Gokhman 1996,
Fig. Ic). Finally, ranges of variation in
chromosome number in all studied fami-
lies of the Chrysidoidea do not overlap.
Specifically, n values of 10-14, 19-21 and
5 were found in the Bethylidae, Chrysidi-
dae and Dryinidae respectively. Though it
is difficult at present to determine path-
ways of karyotype evolution in the Chry-
sidoidea, low chromosome number and

Volume 7, Number 1, 1998

117

n t

I) it

Fig. 1. Karyotj'pe of Anteoii brevicorne. Scale bar in-
dicates 1 (im.

karyotype structure found in the Dryini-
dae and perhaps resulted from multiple
chromosome fusions suggest that those
features may be apomorphic together with
the other apomorphies of this specialized
group (Brothers and Carpenter 1993).
However, further chromosomal investi-
gation of the Dryinidae and other Chry-
sidoidea will be necessary to confirm this
assumption.

LITERATURE CITED

Brothers, D. J. and J. M. Carpenter. 1993. Phylogeny
of Aculeata: Chrysidoidea and Vespoidca (Hy-
menoptera). Journal of Hyniciwplern Rcfcanh 2:
227-302.

Gokhman, V. E. and D. L. J. Quicke. 1995. The last
twenty years of parasitic Hymenoptera karyolo-
gy: An update and phylogenetic implications.
Journal of Hymenoptera Research 4: 41-63.

Goodpasture, C. 1974. Cytological data and classifi-
cation of the Hymenoptera. Unpublished Ph.D.
thesis. University of California, Davis. 178 pp.

Quicke, D. L. J. and V. E. Gokhman. 1996. First chro-
mosome records for the superfamily Ceraphron-
oidea and new data for some genera and species
of Evanioidea and Chrysididae (Hymenoptera:
Chrysidoidea). Journal of Hymenoptera Research 5:
203-205.

Hoshiba, H. and H. T. Imai. 1993. Chromosome evo-
lution of bees and wasps (Hymenoptera, Apo-
crita) on the basis of C-banding pattern analyses.
Japanese Journal of Entomology 61: 465-492.

Imai, H. T., R. H. Crozier and R. W. Taylor. 1977.
Karyotype evolution in Australian ants. Chro-
mosoma 59: 341-393.

Levan, A., K. Fredga and A. A. Sandberg. 1964. No-
menclature for centromeric position on chromo-
somes. Hereditas 52: 201-220.

White, M. J. D. 1973. Animal Cytology and Evolution.
Cambridge University Press, Cambridge. 961 pp.

]. HYM. RES.
Vol. 7(1), 1998, pp. 118-121

NOTE

A Putative Pheromone-gland Associated Modification of the Hind
Tibia in Vipio moneilemae (Hymenoptera: Braconidae: Braconinae)

Donald L. J. Quicke and Jose Vincent Falco

(DLJQ) Department of Biology, Imperial College of Science, Technology and Medicine, Silwood

Park, Ascot, Berks SL5 7PY, UK; (JVF) Departamento de Ciencias Ambientales y Recursos

Naturales, Universidad de Alicante, Ap. Correos 99, E-03080 Alicante, Spain

More is known about the pheromone
and other exocrine glands of braconid
wasps than any other group of parasitic
Hymenoptera, although behavioural evi-
dence suggests that they are present in
many if not all groups and that their prod-
ucts collectively have a variety of roles in-
cluding mate location, host marking, de-
fence and spacing (Quicke, 1997). The ma-
jority of pheromone glands identified to
date are located in the metasoma (Wese-
loh 1980; Tagawa 1983; Buckingham and
Sharkey 1988; Quicke 1991; Field and Kel-
ler 1994; Quicke et al. 1996, 1997) although
recent studies have indicated that the an-
tennae also have a wide range of intrinsic
glands that are involved in courtship as
well as host assessment (Bin et al. 1986,
1989; Isidoro and Bin 1995; Isidoro et al.
1996). Here we describe and illustrate a
modification of the hind tibia of a bracon-
ine wasp, Vipio moneilefiiae Gahan, that
was noticed during revision of the North
American species of Vipio (Inayatullah et
al. 1998). The morphology of the structure
suggests that it is yet another exocrine
gland, and its presence only in males fur-
ther suggests that, if the newly described
structure is associated with a gland, then
this may be a release site for sex phero-
mones. It is worth noting that the hind tib-
ia of an unrelated braconid, the chelonine
Ascogaster reticulatus Watanabe, has been

identified as the source of a sex phero-
mone, but in this case the glands are pres-
ent only in females (Kainoh and Oishi
1993). Males of this species apparently fol-
low female pheromone trails as do the
males of the aphelinid Aphelinus asychis
Walker (Fauvergue et al. 1995).

The male hind tibia of V. moneilemae is
especially thickened for most of its length
(Figs. 1, 3) whereas that of the female (Fig.
2, 4) is essentially unmodified. Along al-
most the whole of the dorsal surface of the
male hind tibia, there is a deep groove
bordered laterally by a distinct ridge on
both the inner and outer sides. Within this
groove, there is a dense arrangement of
setae that point somewhat posteriorly and
whose tips converge towards the mid-line
of the groove (Figs. 5, 6). In some uncle-
aned specimens, the setae are covered by
a deposit, perhaps the dried secretion of
an associated gland, though this has not
been confirmed. We were not able to see
any cuticular pores but these may be small
and the setae obscured the view of most
of the floor of the groove. The arrange-
ment and close-spacing of the setae in the
tibial groove suggest that they could act
as a wick and provide an increased evap-
orative area for the release of pheromones;
similar evaporative setal structures have
been found associated with the metasomal
glands of agathidine braconids (Bucking-

Volume 7, Number 1, 1998

119

Figs. 1-6. Features of the hind leg of Vipio inonciUinac. 1, 3, 5, 6, male leg showing sivollen tibia with dorsal
groove and associated setal arrangement; 2, 4, female showing unmodified tibia. Scale bar: I = 860 ji.m; 2, 3
= 380 ^.m; 4 = 136 (j.m; 5 = 100 |j.m; 6 = 50 M-m.

120

Journal of Hymenoptera Research

ham and Sharkey 1988). No similar struc-
tures have been found on the legs of any
other species of Vipio, nor on those of oth-
er braconines.

Vipio species are idiobiont larval ecto-
parasitoids of concealed hosts living in
wood. Unfortunately, very little is knov^n
about V. moneilemae apart from the fact
that it has been reared from a species of
Moneilema (Cerambycidae) in Mexico (Ga-
han 1930), and it is known from only a
handful of specimens from Mexico and
the southern USA (California, Texas, Utah:
most of the known specimens are in the
United States National Museum of Natu-
ral History). The majority of parasitic
wasps are protandrous, and the males of
many that attack wood-boring hosts con-
gregate at sites from which females are
about to emerge, and where a variety of
competitive tactics may ensue. If the struc-
ture seen on the hind tibia of male V. mo-
neilemae is indeed associated with a sex
pheromone gland, it may suggest a radi-
cally different mating strategy.

Large putative sex pheromone glands
are found in the metasomata of males of
many genera of braconines (Quicke 1991)
as well as in a range of other braconids
such as many opiines, alysiines and agath-
idines (Buckingham and Sharkey 1988).
Most work, including the chemical analy-
sis of glandular products, has been carried
out on those of some Opiinae (Williams t'f
al 1988) and it has been proposed that
these have a role in courtship although a
defensive role cannot be excluded. With-
out doubt, more work needs to be done
on the roles of male exocrine glands in the
Braconidae and other parasitic wasps.

ACKNOWLEDGEMENTS

We would like to thank Rachel Kruft for assistance
with electron microscopy and Dave Smith for the
'oan of specimens of V. moneikmnc. This work was
supported by the NERC (Natural Environment Re-
search Council) Initiative in Taxonomy.

LITERATURE CITED

Bin, F., S. Colazza, N. Isidoro, M. Solinas and S. B.
Vinson. 1989. Antennal chemosensilla and
glands, and their possible meaning in the repro-
ductive behavior of Trissolcus basalis (Woll.)
(Hym.: Scelionidae). Entomcilogicn 24; 33-97.
Bin, F., M. R. Strand and S. B. Vinson. 1986. Antenna
structures and mating behavior in Trissolciif htis-
nlis (Woll.) (Hym.: Scelionidae), egg parasitoid of
the green stink bug. In INRA (ed.) Trichogramnia
and Other Egg Parasites. 2nd International Sympo-
sium, Guangzhou (China), Nov. 10-15 1986. Les
Colloques de ITNRA No. 43, Paris, pp. 144-151.

Buckingham, G. R. and M. J. Sharkey. 1988. Abdom-
inal exocrine glands in Braconidae (Hymenop-
tera). pp. 199-242. In V. K. Gupta (ed.) Advances
in Parasitic Hymenoptera Research. E. J. Brill Pub-
lishing Co., Leiden. 546 pp.

Fauvergue, X., K. R. Hopper and M. F. Antolin. 1995.
Mate finding via a trail sex-pheromone by a par-
asitoid wasp. Proceedings of the National Acadenn/
of Sciences of the U.S.A. 92:' 900-904.

Field, S. A. and M. A. Keller. 1994. Localization of the
female sex pheromone gland in Cotesia rubecula
Marshall (Hymenoptera: Braconidae). journal of
Hymenoptera Researcli 3: 151-156.

Gahan, A. B. 1930. Synoptical and descriptive notes
on parasitic Hymenoptera. Proceedings of the
United States National Museum 77: 1-12.

Inayatullah, M., S. R. Shaw and D. L. J. Quicke. 1998.
The genus Vipio LatreiUe (Hymenoptera: Bracon-
idae) of America North of Mexico. Journal of Nat-
ural History 32: 117-148.

Isidoro, N. and F. Bin. 1995. Male antennal gland of
Amitus spinifcrus (Br.thes) (Hymenoptera: Platy-
gastridae), likely involved in courtship behav-
iour, htternational Journal of Insect Morphology and
Embryology 24: 365-373.

Isidoro, N., F. Bin, S. Colazza and S. B. Vmson. 1996.
Morphology of antennal gustatory sensilla and
glands in some parasitoid Hymenoptera with hy-
pothesis on their role in sex and host recognition.
Journal of Hymenoptera Research 5: 206-239.

Kainoh, Y. and Y. Oishi. 1993. Source of sex phero-
mone of the egg-larval parasitoid, /l.';cc\\;i!sfi'r re-
twulatus Watanabe (Hymenoptera, Braconidae).
Journal of Chemical Ecology 19: 963-969.

Quicke, D. L. J. 1991. Tergal and inter-tergal glands
of male Braconinae. Zoologica Scripta 19: 413—423.

Quicke, D. L. J. 1997. Parasitic Wasps. Chapman &
Hall, London, 470 pp.

Quicke, D. L. J., R. A. Wharton and H. Sittertz-Bhat-
kar. 1996. Recto-tergal fusion in the Braconinae
(Hymenoptera, Braconidae): Distribution and
structure, journal of Hymenoptera Research 5: 73-
79.

Quicke, D. L. ]., K.A. Wharton and H. Sittertz-Bhat-
kar. 1997. The antero-lateral metasomal scent

Volume 7, Number 1, 1998

121

glands of the Braconinae (Hymenoptera, Bracon-

idae): Structure & function, journal of Hymcnoy-

tera Research 6: 219-230.
Tagawa, ]. 1983. Female sex pheromone glands in the

parasitic wasps, genus Apanteles. Ap^'Iicd Ento-

mologi/ and Zoology 18: 416-427.
Weseloh, R. M. 1980. Sex pheromone gland of the

gypsy moth parasitoid, Apanteles metanoscelus:

Revaluation and ultrastructural survey. Annals
of the Entomological Societ}/ of America 73: 576-
580.
Williams, H., M. A. Wong, R. A. Wharton and S. B.
Vinson. 1988. Hagen's gland morphology and
chemical content analysis for three species of
parasitic wasps (Hymenoptera: Braconidae).
Journal of Chemical Ecology 14; 1727-1736.

J. HYM. RES.
Vol. 7(1), 1998, pp. 122-123

NOTE

A Note on the Mating Behavior of Anoplitis amethystinus Fabricius

(Hymenoptera: Pompilidae)

Karen R. Sime' and David B. Wahl

(KRS) Department of Entomology and Section of Ecology and Systematics, Cornell University,

Ithaca, New York 14853, USA; (DBW) American Entomological Institute, 3005 S.W. 56th

Avenue, Gainesville, Florida 32608, USA

To date, nothing has been reported on
the biology of Anopliiis (Notiochares) ame-
thystinus Fabricius, a widespread New
World pompihd ranging from Argentina
to parts of the southern United States (Cal-
ifornia, Arizona, New Mexico, Texas,
Georgia, Florida) (Wasbauer & Kimsey
1985). A handful of remarks may be found
on the biology of related species; to this
growing knowledge we add the following
observations on A. amethystinus.

The encounter occurred in the Riverside
Pine Island (longleaf pine (Pinus palustris
Miller) woods) of the Ocala National For-
est, Florida (29°25'N, 8r47'W). On 11
April 1997, around noon on a lightly over-
cast day, we noticed a group of male pom-
pilids in the tire rut of a seldom-used,
sandy dirt road. Our presence scattered
them, but as soon as we stepped back 1-
2 meters, they returned to an oblong area
of sand (about 10 by 30 cm) in which it
appeared that the surface debris of pine
needles, dead leaves, and small twigs had
been displaced to the periphery by the ac-
tivities of the male wasps. We counted 12-
15 males in this space at any one time,
some flying off occasionally to circle the
area or to visit nearby shrubs. They
walked around in small circles, flicking
their wings and drumming their antennae

' Corresponding author

on the ground. They did not fight or oth-
erwise respond to each others' presence,
although in their tight quarters they often
touched each other.

After watching for a few minutes and
capturing some of the males that flew out
from the group, we swept aside the loose
sand (about 5 mm deep) in the clear area,
and then started excavating the compact-
ed, cement-like sandy substrate. A few
millimeters down we found the opening
of a burrow (6 mm diameter) with a pom-
pilid female near the top, scraping at the
sand in an apparent effort to dig herself
out: her wings were open and dry, and
she flew directly into the waiting net.
When we stepped aside, 4 or 5 males re-
turned; several popped headfirst into the
burrow, then backed out after a few sec-
onds. Further excavation revealed an
empty nest similar to those reported for
Anoplius (Pompilinus) tenebwsus Cresson,
A. (P.) viaticus L. (Aim & Kurczewski
1984), and A. (Arachnophroctotius) apicula-
tus pretiosis Banks (Kurczewski & Spofford
1986): exit tunnel straight, ca. 5 cm long,
angled 50-60° from surface, with terminus
unicellular and widened only very slight-

Similar behavior in male Anoplius has
been reported in a few other species. Rau
(1922) found four males of A. (Notiochares)
atramentarius Dahlbom gathered around a

Volume 7, Number 1, 1998

123

newly emerged female, and males of A.
tenebrosus (Aim & Kurczewski 1984) and
A. apiculatus auttimnalis Banks (Evans et al.
1953) have been observed to fly low over
sandy areas and occasionally alight. These
observations and ours are consistent with
the suggestion of Evans et al. (1953) that
mating may only be successful with vir-
gins in some species (but do not rule out
other strategies). Such behaviors also im-
ply the use of pheromones in combination
with habitat cues in locating female emer-
gence areas, and our observations in par-
ticular suggest that antennation of the
ground may be involved in location of fe-
males at short range.

Voucher specimens (three males and the
female) are deposited at the American En-
tomological Institute. We are grateful for
the help of Mike McDonald, Rex Rowan,
Bill Wcislo, John Wenzel, and several

anonymous reviewers, and we thank the
staff of the Lake George Ranger District
for granting us permission to work in the
Ocala National Forest.

LITERATURE CITED

Aim, S. R., and F. E. Kurczewski. 1984. Ethology of
AiiopHiui tenchrosus (Cresson) (Hymenoptera:
Pompilidae). Proceedings of the Entomological So-
ciet}/ of Washington 86: 110-119.

Evans, H. E., C.-S. Lin, and C. M. Yoshimoto. 1953.
A biological study of Anopilius apiculatus nutum-
nalis (Banks) and its parasite, Evageles mohai'e
(Banks) (Hymenoptera, Pompilidae). Journal of
the New York Entomological Society 61: 61-77.

Kurczewski, F. E., and M. G. Spofford. 1986. Obser-
vations on the behaviors of some Scoliidae and
Pompilidae (Hymenoptera) in Florida. Florida
Entomologist 69: 636-644.

Rau, P. 1922. Ecological and behavior notes on Mis-
souri insects. Transactions of the Academy of Sci-
ence of St. Loins 24(7): 1-71.

Wasbauer, M., and L. Kimsey. 1985. California spider
wasps of the subfamily Pompilinae. Bulletin of the
California insect Sunvy 26: 1-130.

Vol. 7(1)

J. HYM. RES.
1998, pp. 124-126

BOOK REVIEW

Annotated Keys to the Genera of Nearctic
Chalcidoidea (Hi/nienoptern). Gary A. P. Gib-
son, John T. Huber and James B. Woolley
(Editors.). NRC Research Press, Ottawa,
1997. xi + 794 pp. Price: Canada
CAN$64.95, other countries the equivalent
of US$64.95 (hardcover). ISBN 066016669-0.

It has been said that it is dangerous to
study the Parasitic Hymenoptera, for
those who do are inclined to become al-
coholics or end up in mental institutions.
Notions of such despair undoubtedly stem
from the daunting task that confronts
those who venture to unravel the identity
of this vast and taxonomically complex as-
semblage of enigmatic wasps.

The superfamily Chalcidoidea alone
comprises close to 19,000 valid species in
2,000 genera, with estimates of the actual
number of species ranging from between
60,000 to 100,000 world wide. Compound-
ing the difficulties in coming to grips with
the identity of this bewildering array of
mostly poorly known taxa is the plethora
of scattered and often inaccessible descrip-
tive literature spanning some 200 years of
taxonomic endeavour on the group. The
task is made even more arduous by a
shortage of essential identification tools
such as modern catalogues, revisions,
keys and adequately documented refer-
ence collections. In Britain, arguably the
cradle of Chalcidology, there are for in-
stance no modern identification keys for
more than half of the 5,000 or so species
of Hymenoptera (Weeks et al. 1997, Bull.
Ent. Res. 87: 203), many of which belong
to chalcidoid groups.

What is clearly needed, is to take stock
of what is known, collate and synthesize
the vast pool of scattered knowledge,
much of which is buried in a few inquir-
ing minds, and make it readily accessible
in such a way that it is both palatable and

of practical value to all. Indeed, this is not
only imperative in giving credence to the
ideals of the Convention on Biological Di-
versity, but also in proving the worth of
insect systematists in a world of changing
attitudes to science.

The 17 authors of Annotated Keys to the
Genera of Nearctic Chalcidoidea receive
full credit for having done exactly that. In
harnessing their collective skills to the full
they have produced an outstanding syn-
opsis of the 19 families and 706 chalcidoid
genera known to occur in the region. But
the true value of this book lies far beyond
that of a manual merely for identifying
specimens. It its also an impressive refer-
ence work, crammed with invaluable no-
menclatural, bibliographical and host in-
formation on the Nearctic chalcidoid fau-
na.

The book comprises 22 chapters. The in-
troduction, which contains interesting
background information on the history of
Chalcidology in North America, numbers
of taxa and the rate at which genera have
been described from the region through
the years, is followed by a chapter on mor-
phology. Morphological terms pertaining
to the keys and various family diagnoses
are conveniently highlighted in bold,
clearly explained and depicted in a series
of uncluttered line drawings and good
quality scamiing electron micrographs.
For quick reference, terms are listed al-
phabetically in a separate appendix with
their abbreviations and reference to fig-
ures. A second appendix cross-references
the abbreviations with terms. The disad-
vantage of having this information sepa-
rate, and not in figure legends below the
plates, is that some paging is required to
determine the meanings of the abbrevia-
tions depicted in the illustrations.

Chapter three provides, besides a key to

Volume 7, Number 1, 1998

125

families, an overview of the Chalcidoidea
and covers topics such as recognition of
the group, mode of parasitism (feeding
types), host ranges and relationships, be-
haviour, collecting methods and voucher
specimens. Written in clear language, this
is essential reading matter which should
appeal in particular to the non-specialist
seeking general background information
on the superfamily at a glance.

The family key is primarily a derived
combination of those found in A Hand-
book of the families of Nearctic Chalci-
doidea and Hymenoptera of the World. In
recognizing 19 families, the authors have
opted for the traditional system of classi-
fying chalcidoids at this level. Considering
the intended purpose of the book and the
instability of the higher classification of
these wasps, in which anything from nine
to 24 families have been recognized, this
is a sensible choice.

Preceding the key is a flow chart which
gives a pleasing visual impression of how
the 19 families are keyed out in the 40 cou-
plets. The key itself is visually less pleas-
ing. Placing the figures associated with
each alternate of a couplet directly above
it allows for easy comparison between fig-
ures and text, but often results in insuffi-
cient space for both alternates on the same
page. Consequently, many pages are left
with unsightly blank spaces, while the al-
ternates of some couplets appear on dif-
ferent pages, making comparisons some-
what cumbersome. More important
though, is the fact that the key works well.
Evidently, much careful thought has gone
into making it both accurate and workable
for the novice and specialist alike. Chosen
for diagnostic reliability rather than con-
venience, the characters are not necessar-
ily "easy" to use, but being appropriately
and clearly illustrated there should be no
problem in keying specimens out.

The bulk of the remaining text is devot-
ed to the family treatments. Each of the 19
chapters reviews one family and includes,
apart from a generic key, sections on its

recognition, systematics, biology, fauna
and literature. For larger families a useful
index to genera based on couplet numbers
is also included. These topics are all well
covered, serving as a concise and infor-
mative introduction to the 19 families as
represented in the Nearctic region.

In essence, this book is all about generic
keys to Nearctic chalcidoids, and its real
value will ultimately be judged by how
well these keys perform. So I decided to
put some of them to the test, selecting
specimens not only from the Nearctic re-
gion, but also from further afield. The re-
sults were excellent, bearing testimony to
the skillful and competent way in which
these keys have been constructed. The
contents of the couplets are unambiguous
and the taxonomic language easy to com-
prehend. I was pleased to see that the au-
thors have, in all but one of the tested
keys, managed to avoid using the handy
but dreadful "either/or" method of sepa-
rating taxa. All of this adds much to the
relative ease with which the user is guided
through the keys, even in the case of gen-
era which are difficult to define. Although
these keys have been designed specifically
for the Nearctic region, they will, if used
with the necessary insight and care, also
serve as a very useful tool in the study of
extra-limital forms, as was evident from
the large selection of Afrotropical speci-
mens which keyed out perfectly well.
Greatly enhancing the value of these keys
are brief annotations to each genus com-
prising references to existing species keys,
estimated number of species, known dis-
tribution and host range in the region.

The use of the keys is facilitated by
1,880 line drawings and scanning electron
n\icrographs which, despite the inputs
from a variety of illustrators, are of a con-
sistently high aesthetic and technical stan-
dard. With the intended readership of the
book in mind, a few more eye-catching
habitus drawings, which are completely
lacking for some families, would have
added a special touch.

126

Journal of Hymenoptera Research

The book concludes with four very use-
ful indices. The first one comprises about
130 generic and subgeneric chalcidoid
names (with their nomenclatural fate)
which have become invalid since publi-
cation of the most recent (1979) edition of
Catalog of Hymenoptera in America
North of Mexico. Following this index are
separate lists of plant and animal hosts,
and supraspecific chalcidoid names.

In summary, I applaud the contents of
this book, hence the lack of any serious
criticism. Naturally, this is not the last
word on the supraspecific identity of Ne-

arctic chalcidoids, for our knowledge and
understanding of the fauna is far from
complete. But the authors have succeeded
admirably in synthesizing what is pres-
ently known while providing an excellent
framework for further revisionary work
on the group. 1 recommend this book as
an indispensable reference and identifica-
tion tool for anyone, specialists and gen-
eralists alike, involved in the study of par-
asitic wasps.

Gerhard Prinsloo, Biosystematics Divi-
sion ARC-Plant Protection Research Institute
Pretoria, South Africa.

ANNOUNCING THE PUBLICATION OF

MANUAL OF THE NEW WORLD

GENERA OF THE FAMILY

BRACONIDAE

Edited by
Robert A. Wharton, Paul M. Marsh and Michael J. Sharkey

Contributing Authors

Paul C. Dangerfield, Paul M. Marsh, Donald L. J. Quicke,

Michael J. Sharkey, Scott R. Shaw, Cornelis van Achterberg,

Robert A. Wharton, and James B. Whitfield

Special Publication Number 1
International Society of Hymenopterists

This 439 page, multiauthored identification manual presents illustrated
keys to the 34 subfamilies and 404 genera of the family Braconidae from
the New World. Information about all New World genera described up
to 1994 is included. The manual consists of 37 chapters by world au-
thorities plus an index to generic names including subgenera and syn-
onyms, and an appendix listing all the New World genera alphabetically
by subfamily. The first chapter provides an introduction including a re-
view of braconid biology, literature, classification, biogeography, collec-
tion and curation, and a discussion of the manual format. The second
chapter is an extensive illustrated discussion of braconid morphology
and terminology used in the keys. The third chapter is a fully illustrated
key to the New World subfamilies of the Braconidae. The New World
subfamilies are then presented separately in the remaining 34 chapters.
For each, a key to the New World genera is included. Each of these keys
is annotated to include estimated number of species, distribution, hosts,
and critical references for each genus. Each subfamily chapter is exten-
sively illustrated and the entire manual contains over 750 line drawings
and scanning electron micrographs.

A valuable identification tool for biological control workers and museum curators

Price: US$20.00

In the United States, prepaid orders are sent postage free. Outside the U.S., postage will
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Send orders (check made out to "Department of Entomology") to:

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CONTENTS
(Continued from front cover)

WHARTON, R. A., S. M. OVRUSKI, and F. E. GILSTRAP. Neotropical Eucoilidae (Cyni-
poidea) associated with fruit-infesting Tephritidae, with new records from Argen-
tina, Bolivia and Costa Rica 102

NOTES

GOKHMAN, V. E., and K. A. KOLESNICHENKO. First chromosome record for the
family Dryinidae: The karyotype of Anteon brevicorne Dalman (Hymenoptera:
Chrysidoidea) 116

QUICKE, D. L. J., and J. V. FALCO. A putative pheromone-gland associated modifi-
cation of the hind tibia in Vipio moneilemae (Hymenoptera: Braconidae: Bracon-
inae) 118

SIME, K. R., and D. B. WAHL. A note on the mating behavior oi Anoplius amethysiinus

Fabricius (Hymenoptera: Pompilidae) 122

BOOK REVIEW

PRINSLOO, G.— Gary A. P. Gibson, John T. Huber and James B. Woolley (Editors).

1997. Annotated Keys to the Genera of Nearctic Chalcidoidea (Hymenoptera) .... 124

ANNOUNCEMENT

International Society of Hymenopterists, Special Publication Number 1: Manual of the

New World Genera of the Family Braconidae 127

c,oClETy

Journal of

Hymenoptera
Research

^3

'^M/olume 7, Number 2

October 1998

ISSN #1070-9428

CONTENTS

ARCHER, M. E. Worker versus sexual, and sex ratio investments in the social u^asp Vespula

vulgaris (L.) (Hymenoptera: Vespinae) in England 257

GESS, F. W. Priscomasaris namihiensis Gess, a new genus and species of Masarinae (Hyme-
noptera: Vespidae) from Namibia, southern Africa, with a discussion of its position
within the subfamily 296

GIBSON, G. A. P., and V. VIKBERG. The species of Asaphes Walker from America north of
Mexico, with remarks on extralimital distributions and taxa (Hymenoptera: Chal-
cidoidea, Pteromalidae) 209

LIU, ZHIWEI. A new species of Ihalia from Borneo, with a revised phylogeny and historical

biogeography of Ibaliidae (Hymenoptera: Cynipoidea) 149

MACEDO, M. V. DE, M. C. P. PIMENTEL, and R. C. VIEIRA. Response of Pithecellobium
tortum Martius (Leguminosae) seeds to the attack of the phytophagous braconid
AUorhogas dyspistus Marsh (Hymenoptera: Braconidae) 274

MARTINS, R. P., L. A. SOARES, and D. YANEGA. The nesHng behavior and dynamics of
Bicyrtes angiilata (F. Smith) with a comparison to other species in the genus (Hy-
menoptera: Sphecidae) 165

NAUMANN, I. D., and H. GROTH. A revision of the Philomastigine sawflies of the world

(Hymenoptera: Pergidae) 127

OLIVEIRA, L., R. MELO, and ]. TAVARES. Response of Glypiapanteks militaris (Walsh)
(Hymenoptera: Braconidae), a larval parasitoid of the armyworm, Mythimna uni-
puncta (Haworth) (Lepidoptera: Noctuidae), to different temperatures 268

(Continued on back cover)

INTERNATIONAL SOCIETY OF HYMENOPTERISTS

Organized 1982; Incorporated 1991

OFFICERS FOR 1998

James M. Carpenter, President

Andrew D. Austin, President-Elect

James B. WooUey, Secretary

John T. Huber, Treasurer

E. Eric Grissell, Editor

Subject Editors

Symphyta and Parasitica Aculeata

Biology: Mark Shaw Biology: Sydney Cameron

Systematics: Donald Quicke Systematics: Wojciech Pulawski

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Title of Publication: Journal of Hymenoptera Research.

Frequency of Issue: Twice a year.

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of Hymenopterists, % Department of Entomology, NHB 168, Smithsonian Institution, Wash-
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ural History, Washington, DC 20560-0168.

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This issue was mailed 20 October 1998

J. HYM. RES.
Vol. 7(2), 1998, pp. 127-148

A Revision of the Philomastigine Sawflies of the World
(Hymenoptera: Pergidae)

I. D. Naumann and H. Groth

(IDN) CSIRO Division of Entomology, P. O. Box 1700, Canberra, Australian Capital Territory
2601, Australia; (HG) "Timbarrah", M/S 1073, Crows Nest, Queensland 4355, Australia

Abstract. — The world genera and species of Philomastiginae (Cerospastus Konow, Ecopatus Smith
and Philomastix Froggatt) are revised and an illustrated, identification key is provided. Cewspastus
volupis Konow and Ecopatus penai Smith from South America and Philomastix nancarrowi Froggatt,
P. macleaii (Westwood) and P. xanthophylax sp. n. from eastern Australia are described or re-
described. Larvae of C. volupis feed on foliage of species of Nothofagus (Fagaceae); those of Phi-
lonmstix on foliage of Rulms (Rosaceae) oi Alphitonin (Rhamnaceae). All species of Philomastix
exhibit maternal guarding of eggs and larvae. The trans-Antarctic distribution of the Philomasti-
ginae suggests a Gondwanan origin for the subfamily.

Distinctive Gondwanan elements are
novi' known within numerous Australian
families of Hymenoptera (Naumann
1991). Over a dozen families or subfami-
lies and even a few genera are shared by
Australia and South America or by Aus-
tralia, South America, New Zealand and
southern Africa. Examples of such trans-
Antarctic distribution patterns can be
found among the Symphyta, Megalyro-
idea, Evanioidea, Ichneumonoidea, Proc-
totrupoidea, Platygasteroidea, Chalcidoi-
dea, Vespoidea and Apoidea and include
phytophages, parasitoids and predators.
Generally these Gondwanan elements are
readily recognisable as archaic clades
within their respective superfamilies or
families. Some, such as the parasitoid ge-
nus Monomachus Klug (family Monomach-
idae) are morphologically conservative
and comprise in total only a few dozen
species (Johnson 1992). Some, such as
Monomachus, are very rarely collected or
are restricted to temperate, forest habitats.
In contrast such Gondwanan groups as
the thynnine Tiphiidae are spectacularly
varied and speciose; the many hundreds
of species (Naumann 1991) occur over a

wide range of habitats and some of these
species are extremely common.

Seven subfamilies or tribes of Symphy-
ta, the Dereycyrtinae (Xiphydriidae), the
Guiglini and Leptorussini (Orussidae), the
Scobini (Argidae), and the Pergulinae,
Perreyinae and Philomastiginae (Pergi-
dae) all exhibit trans-Antarctic distribu-
tion patterns. The Dereycyrtinae compris-
es the monotypic Austrocyrta Riek in Aus-
tralia and three genera totalling sixteen
species in South America (Smith 1988;
Smith 1995). Larvae of this subfamily are
unknown but probably develop within
wood. The Guiglini comprise six Austra-
lian and two New World species, and the
Leptorussini consist just a South American
species, five Australian species and a spe-
cies in South Africa (Smith 1988). All orus-
sid larvae are believed to be ectoparasitic
on the larvae of wood-boring beetles and
siricid wood wasps. The orussid tribe
Ophrynopini, with representatives in
south-east Asia and South America might
also be considered to be a "trans-Antarc-
tic" taxon (N. Springate, pers.comm.). The
Scobinini comprise Scobinn Lepeletier and
Serville, a common New World genus of

128

Journal of Hymenoptera Research

about 50 species, and Antargidium Morice,
an Australian genus of six species (Nau-
mann 1991). Larvae of two species of Scob-
ina feed on Sida Linnaeus (Malvaceae) but
nothing is known of the biology of Antar-
gidium. The subfamily Pergulinae (Pergi-
dae) comprises the genus Haplostegus Ko-
now with fifteen species in South America
and the monotypic Pergula Morice in
south-western, mainland Australia (Smith
1990). The biology of the single, rare spe-
cies of Pergula is unknown but larvae of
the South American pergulines have been
recorded feeding on Myrtaceae, especially
Psidium guajava L. (guava). Perreyinae are
represented on the western side of the Pa-
cific ocean by two species with flightless
females (described respectively from Su-
lawesi and New Guinea) and east of the
Pacific by about 80 species in South Amer-
ica (Smith 1990). Larvae of South Ameri-
can species feed variously on Asteraceae
and Malvaceae. The Philomastiginae con-
sists of two monotypic South American
genera, Cerospastus Konow and Ecopatus
Smith, and three eastern Australian spe-
cies of Philomastix Froggatt (Fig. 75). The
two previously described species of Phi-
lomastix are well known species of tropical
and subtropical rainforests. Their larvae
feed on the foliage of species of Rubus L.
(Rosaceae) (Leask 1944) or Alphitonia Reis-
sek ex Endl. (Rhamnaceae) (Jackson 1993)
and females are commonly encountered
standing guard over egg batches and
young larvae (Fig. 72). Members of the
subfamily can be recognised using the
keys of Smith (1990) and Naumann (1991).
The present paper revises the generic
diagnoses and key of Smith's (1990) treat-
ment of the South American Philomasti-
ginae, redescribes all previously named
species, describes a new species of Philo-
mastix from south-eastern Queensland
(Fig. 1), and collates all known biological
and distributional data on the subfamily.
Morphological and biological studies of
the larvae of Philomastix are continuing
(Groth and Naumann unpublished).

Fig. 1. Philomastix xanthophylax. Scale line = 2mm.

TERMINOLOGY

The morphological terminology used
here follows Goulet and Huber (1993).

DEPOSITORIES

AMSA Australian Museum, Sydney,
Australia

ANIC Australian National Insect Col-
lection, CSIRO Division of En-
tomology, Canberra, Australia

BPBM Bernice P. Bishop Museum, Ho-
nolulu, Hawaii, U.S.A.

CNC Canadian National Collections

of Insects, Arachnids and Nem-
atodes, Ottawa, Canada

BCRI Biological and Chemical Re-
search Institute, N. S. W. De-
partment of Agriculture, Rydal-
mere, Australia

BMNH The Natural History Museum,
London, U. K.

DEIC Institut fur Pflanzenschutzfor-
schung der Akademie der
Land wirtschaftswissenschaf ten
(formerly:Deutsches Entomolo-
gisches Institut, Eberswalde,
Germany)

FCNI State Forests Research Institute,
Beecroft, Australia

Volume 7, Number 2, 1998

129

HGCN H. Groth Collection, "Timbar-
rah/' via. Crows Nest, Australia

MVMA Museum of Victoria, Mel-
bourne, Australia

UMO Hope Department of Entomol-
ogy, University Museum, Ox-
ford, U. K.

QMBA Queensland Museum, Brisbane,
Australia

QDPl Entomology Collection, Queens-
land Department of Primary In-
dustries, Brisbane, Australia

SAMA South Australian Museum,
Adelaide, Australia

UQIC Department of Entomology,
Uruversity of Queensland, Bris-
bane, Australia

USNM National Museum of Natural
History, Smithsonian Institu-
tion, Washington, D. C, U. S. A.

PHILOMASTIGINAE Rohwer

Pterygophorinae (part): Froggatt 1890b: 696;

Ashmead 1898: 231.
Ptergophorides (part): Konow 1898: 248; Ko-

now 1905b: 37.

Philomastiginae Rohwer 1911a: 220; Benson
1935: 224; Benson 1938: 379; Riek 1970a: 891;
Riek 1970b: 218; Smith 1978: 159; Naumann
1984: 345, 347; Smith 1990: 13, 21-23; Nau-
mann 1991: 934-935; Macdonald and Ohmart
1993:493-^95.

Diagnosis. — Antenna 14-21-segmented,
filiform in female, serrate in male. Head
capsule open (neither postgenae nor hy-
postomae forming continuous bridge be-
tween occipital and foramen and oral fos-
sa). Maxillary palp 4- or 6-segmented; la-
bial palp 3- or 4-segmented. Labium api-
cally single or tri-lobed. Ventral arms of
cervical sclerites pointed, neither meeting
nor forming precoxal bridge with proster-
num; notauli complete; mesothoracic ster-
nopleural suture present; distance be-
tween cenchri less than half width of cen-
chrus. Mid- and hind tibia each with pre-
apical spine; foretibia with 2 apical spines.
Costa of forewing narrower than intercos-
tal area; forewing without anal cell. Ab-
dominal tergum 1 sclerotised.

KEY TO GENERA OF PHILOMASTIGINAE

1. Maxillary palp 4-segmented; labial palp 3-segmented, apical segment with large sensory
cup occupying more than half length of segment (Fig. 70), maxillary palp with smaller
sensory cup; labium single lobed; female without cercus Philomastix Froggatt

- Maxillary palp 6-segmented; labial palp 4-segmented, apical segments without sensory
cup; labium tri-lobed; female with cercus 2

2. Forewing with 4 cubital cells (IRl, IRs, 2Rs, 3Rs) (Fig. 4); forewing radial cell closed; fourth
maxillary palp segment slender (Fig. 9); antenna of female weakly clavate (Fig. 3); meso-
scutum conspicuously setose Cerospastiis Konow

- Forewing with 2 or 3 cubital cells; fore-wing radial cell open (Fig. 5); fourth maxillary palp
segment apically distinctly wider than other segments (Fig. 8); antenna of female not cla-
vate (Fig. 2); mesoscutum almost devoid of setae Ecopatiis Smith

PHILOMASTIX Froggatt

Philomastix Froggatt 1890a: 487^88; Froggatt
1890b:696; Froggatt 1901: 1070; Dalla Torre
1894: 308; Ashmead 1898: 231; Konow 1898;
248, 249; Konow 1905: 36-37; Schulz 1906:
81-82; Rohwer 1911: 87; Morice 1919: 248,

255, 287; Tillyard 1926: 265; Benson 1935: 224;
Benson 1938: 379-380; Riek 1970a: 880, 889,
891; Smith 1978: 159; Smith 1990: 21-23; Pag-
liano and Scaramozzino 1990: 157; Abe and
Smith 1991: 65; Naumann 1991: 926, 933, 935;
Macdonald and Ohmart 1993: 493-496; Nau-
mann 1994: 414-415. Type species: Perga nan-

130

Journal of Hymenoitera Research

4,5

2,6,7

8,9

Figs. 2-9. Ecopatus penae, Cerospastus volnpis: 2, £. pcnae, female antenna; 3, C.volupis, female antenna; 4, C.
vohipis, forewing (part); 5, £. penac. forewing (part); 6, £. pcnac, lateral panel of first abdominal tergum; 7, C.
I'olupis, apex of male abdomen, dorsal view; 8, £. penae, maxillary palp palp; 9, C. mhipia, maxillary palp.
Scale lines = 1.0 mm.

carrowi Froggatt (designated by Rohwer

1911).
Perga Leach: Westwood 1880: 372; Kirby 1882:

26. Dalla Torre 1894: 351. (part)
Heptncola Konow: Konow 1905a: 167 (part)

Female. — Vertex conspicuously setose.
Face with some fine microsculpture. Malar
space narrower than diameter of anterior
ocellus. Antenna 14-19-segmented, weak-
ly serrate, not clavate. Right mandible
simple (Fig. 69). Maxillary palp 4-seg-
mented, filiform (Fig. 70); apical segment
with conspicuous sensory cup, this less
than half as long as segment. Labial palp
3-segmented, apical segment with con-
spicuous sensory cup, this more than half
as long as segment. Labium single lobed.
Thorax dorsally conspicuously setose. No-
tauli, median mesoscutal line deeply im-
pressed. Mesoscutellum swollen so that

posterior margin concealed from above.
Mesepisternum with tubercle. Metascutel-
lum in form of transverse band. Forewing
with closed radial cell and 4 cubital cells
(e.g. Fig. 1). Median, second cubital and
third cubital cells each with nygma (small,
corneous spot). Abdominal terga dorsally
densely setose. Second tergum predomi-
nantly smooth. Cercus absent. Ovipositor
sheath posteriorly strongly expanded, in
posteroventral view with prominent, flat-
tened, semicircular surface (e.g. Fig. 21).

Head, mesoscutellum, metascutellum
orange-yellow. Legs brown to dark
brown. Wings with background smokey
brown appearance, especially in female.
Costal cell of fore wing distinctly more
darkened than remainder of wing.

Male. — Antenna 15-21-segmented, stron-
glyserrate (e.g. Fig. 11). Eighth tergum pos-

Volume 7, Number 2, 1998

131

teriorly with moderate or very weak emar-
gination (e.g. Fig. 20).

Discussion. — Larvae of Philomastix have
long been known to feed externally on
leaves of various shrubby and scandent
species of Riibus (known as blackberries or
native raspberries) (Leask 1944). More re-
cently (present paper and Jackson 1993)
Philomastix larvae have been discovered
defoliating trees of the genus Alphitonia
(Figs. 72-74). Two species of Alphitonia are
attacked: Alphitonia petriei Braid & C. T.
White, the pink ash, which is widespread
in Queensland and the northern part of
the Northern Territory, and Alphitonia ex-
celsa (Fenzl) Benth., the red ash, which oc-
curs in rainforests and eucalypt wood-
lands of Queensland, New South Wales,
and the northern parts of the Northern
Territory and Western Australia (Francis
and Chippendale 1970; Lazarides and
Hince 1993). The hosts of Philomastix are
thus a shrub and a tree from different
plant families. Rubus and Alphitonia do
have at least one, ecological characteristic
in common — both are pioneer species.
Various Rtthiis are well known species of
paths, clearings and margins of closed for-
ests and A. petriei is the most prominent
recolonising tree species in upland rain-
forest clearings in north Queensland (Jack-
son 1993).

A species of "Philomastix" has been re-
corded feeding on Eucalyptus sp. at Laun-
ceston, Tasmania (Anonymous 1980). This
is almost certainly a misidentification: it is
the only record of a eucalypt as host and
we have seen no authentic specimens of
Philomastix from Tasmania in the course of
this revision.

Philomastix does not occur over the en-
tire range of its host plants. The sawfly ge-
nus is restricted to the eastern Australian
states of Queensland and New South
Wales between the latitudes 16°S and 35°S.
However Alphitonia occurs beyond this
range in the Northern Territory and West-
ern Australia and suitable species of Rubus
are common in Victoria (Bruzzese 1980).

P. nancarwwi and P. macleaii are strictly
allopatric (Fig. 76). P. nancarwwi is con-
fined to the rainforests and closely adja-
cent eucalypt woodlands of the Queens-
land Wet Tropics between the northern
end of the Atherton Tableland and the
Mount Spec-Paluma area. Almost all spec-
imens have been collected at altitudes
above 500m. P. macleaii is very widely dis-
tributed from Eungella in central Queens-
land to the lllawarra district in south-east-
ern New South Wales. It occurs in tropi-
cal, subtropical and temperate rainforest
and nearby moist woodlands. In the
northern parts of its range (Eungella,
Kroombit tops, Bunya Mountains) P. ma-
cleaii is not known below 1000m but south
of about Brisbane it occurs near sea level.
The region of eastern Queensland between
Paluma and Eungella which separates P.
nancarrowi and P. macleaii is relatively dry;
the vegetaHon is dominated by eucalypt
woodland and there are no significant
patches of rainforest. North from Paluma
the so-called "base-of-peninsula" rainfor-
est system stretches with some interrup-
tions to Cooktown. On the other hand
Eungella stands as the northern extremity
of a rainforest system that extends, also
with interruptions, all the way to southern
New South Wales. There are taxonomic
discontinuities in several orders of insects
(especially mesothermic Odonata, Plecop-
tera and Megaloptera) at the gap between
Paluma and Eungella (Kikkawa et al.
1981; Watson and Theischinger 1984) and
these are thought to reflect Pleistocene
fluctuations in climate. During Pleistocene
glacial periods, increased aridity resulted
in a contraction of the rainforests and a
pronounced inhospitable gap between
moist refugia north of Paluma and south
of Eungella. Presumably vicariance speci-
ation occured on either side of this barrier
within Philomastix and various odonate,
stonefly and alderfly genera.

P. xanthophylax has a much more cir-
cumscribed distribution pattern (Fig. 76).
It has been collected in moist woodlands

132 Journal of Hymenoptera Research

rather than rainforest but in the Brisbane nal care. Females stand over or near their

area it is sympatric with P. macleaii. There egg masses and young larvae (Fig. 72).

is at present no model to explain the ori- When disturbed they shake from side to

gin of this species. side and rapidly vibrate their wings to

The oviposition behaviour of Philomas- produce a loud buzzing sound. Females
tix is unusual for a pergid sawfly. The will guard their offspring, usually from a
adult female pierces the leaf of the host position on a petiole or stem, and have
plant, pushes each elongate egg through been observed to guide larvae to new
the perforation, and attaches one end of leaves (Leask 1944). Females eventually
the egg to the underside of the leaf (Mac- die at their post, and sometimes dozens of
donald and Ohmart 1993). The other end dried bodies remain hanging on defoliat-
of the egg is pushed free of the lower sur- ed host plants. Parental defence of off-
face of the leaf. The more typical behav- spring is widespread within the Symphy-
iour for pergids is to cut a shallow slit in ta. It has been recorded in three other sub-
the host plant and insert the egg into this families of Pergidae (Naumann 1984; Mac-
slit. The lancet of most pergids is saw-like donald and Ohmart 1993) and in the
in appearance but the lancet of Philomastix Pamphiliidae (Kudo et al. 1992). In those
(Figs. 22, 39, 58, 71) is auger-shaped, pre- species which have been studied quanti-
sumably to facilitate the "pierce-and- tively it has been shown that female
push" oviposition habit. guarding reduces predation on eggs by

All species of Philomastix exhibit mater- other arthropods.

KEY TO SPECIES OF PHILOMASTIX

Female; abdomen with saw-like ovipositor often concealed within sheath (Figs. 1, 21, 46,

55) 2

Male; ovipositor and sheath absent 4

Abdomen orange-yellow (Fig. 1); antennal flagellum black; ovipositor sheath posterome-

dially dentate (Figs. 54, 55) xanthophylax sp.n.

Abdomen with extensive dark brown, black and cream markings (Figs. 62,64); antennal
flagellum orange-yellow; ovipositor sheath not posteromedially dentate (Figs. 21, 46) .... 3
Mesoscutellum usually with posterolateral tubercles (Figs. 30-34); mesepisternal tubercle
strong (Fig. 38); abdominal terga 3-5 usually with cream, lateral maculae visible from
above (Fig. 62); posterolateral margin of first abdominal tergum weakly curved or straight
(Fig. 37), spiracle separated from margin of tergum by a distance 0.9-1.1 times maximum

diameter of spiracle macleaii (Westwood)

Mesoscutellum without posterolateral tubercles (Fig. 16) although sometimes posteriorly
emarginate; mesepisternal tubercle usually weak (Fig. 26); abdominal terga 3-5 without
lateral, cream maculae visible from above (Fig. 64); posterolateral margin of first abdominal
tergum strongly curved almost angulate (Fig. 19), spiracle separated from margin of ter-
gum by a distance 1.6-1.7 times maximum diameter of spiracle nancarroivi Froggatt

Mid lobe of mesoscutum with brown to black coloration reaching anterior margin (Fig. 63);
diameter of anterior ocellus 0.9-1.0 times minimum distance between antennal sockets;
sixth abdominal tergum with lateral, cream or orange-yellow macula visible from above

(Fig. 63); mesoscutellum usually with posterolateral tubercles mflc/ffliiXWestwood)

Mid lobe of mesoscutum reddish orange anteriorly and brown to black posteriorly or lobe
entirely reddish orange; diameter of anterior ocellus approximately 0.7 times minimum
distance between antennal sockets; sixth abdominal tergum usually without orange or
yellow maculae visible from above; mesoscutellum without tubercles, although posterior

margin sometimes distinctly concave 5

Seventh abdominal tergum with widely separated, yellow or orange maculae; posterolat-

Volume 7, Number 2, 1998

133

eral margin of first abdominal tergum strongly curved, almost angulate (as in Fig. 19);

mesoscutellum posteriorly weakly emarginate (Fig. 17) or straight mmcarrowi Froggatt

Seventh abdominal tergum with yellow or orange band across posterior margin, band
sometimes narrowly interrupted medially; posterolateral margin of first abdominal tergum
weakly curved (as in Fig. 53) or straight; mesoscutellum posteriorly straight or weakly
convex xanthophylax sp.n.

Philomastix nancarroivi Froggatt
(Figs. 10-26, 64-66, 76)

Philomastix nancarrowi Froggatt 1890a: 488-489;
Froggatt 1890b: 696; Konow 1898: 250; Schulz
1906: 82-83; Rohwer 191 IB: 87; Morice 1919:
287-288, Plate XI, fig. 13, Plate XII, figs 5,6,
Plate XIII, fig. 15; Leask 1943: 2; Leask 1944:
1-3; Smith 1978: 159; Bruzzese 1980: 4; Abe
and Smith 1991: 65; Macdonald and Ohmart
1993: 493-494; Naumann et al. 1994: 71.

Philomastix nancarrowi: Dalla Torre 1894: 308;
Konow 1905: 37; Rohwer 1911: 87. (Miss-
spelling)

nee Philomastix nancarrowi : Forsiusl927: 283.
(Misidentification)

Female. — Body length 9.5-11.0 mm.
Forewing length 10.6-12.3 mm. Distance
between antennal sockets 2.8-3.9 times di-
ameter of anterior ocellus. Antenna (Fig.
10) 15-18-segmented; apical 2-A segments
sometimes fused so that antenna appar-
ently 13-16-segmented (traces of interseg-
mental sutures sometimes present). First
flagellar segment 2.8-3.8 times longer than
wide. Second flagellar segment 1.7-1.9
times wider apically than basally. Meso-
scutellum without posterolateral tuber-
cles, posteriorly rounded or weakly emar-
ginate (Fig. 16); anterodorsal surface flat
(Fig. 15) to weakly concave. Mesepisternal
tubercle weak (Fig. 26). Ovipositor sheath
not posteromedially dentate (Fig. 21). Pos-
terolateral margin of first abdominal ter-
gum strongly curved (Fig. 19), almost an-
gulate, spiracle separated from margin by
a distance 1.6-1.7 times maximum diam-
eter of spiracle. Lancet as in Figs. 22-25,
about 8 ctenidia terminating in dorsal
tooth, anterior teeth weak

Flagellum orange-yellow. Mid lobe of
mesoscutum orange-yellow (Fig. 64). Each

lateral lobe of mesoscutum with large,
dark brown to black macula occupying
most of length of lobe. Mesepisternum
with broad, brown band adjacent to ster-
nopleural suture (Fig. 66). Mesosternum
brown with narrow, orange-yellow band
along anterior and lateral margins. Meta-
scutum orange-yellow to pale brown, usu-
ally not much darker than cenchri. Fore-
wing with broad, transverse, brown band;
cell IM usually completely brown (Fig. 12)
or forewing entirely brown (Fig. 13). Ab-
domen predominantly dark coloured with
some metallic reflections. Abdominal ter-
gum 1 orange-yellow to pale brown. Terga
3-5 dark brown, black or metallic blue.
Terga 6 and 7 each with cream, lateral
macula; maculae narrowly separated mid-
dorsally by dark brown or black, or mac-
ulae contiguous. Tergum 8 cream. Tergum
9 orange-yellow. Abdominal sterna 2 and
3 at least in part orange-yellow or cream.
Sterna 4—7 dark brown to black. Oviposi-
tor sheath orange-yellow.

Male. — Body length 7.1-9.4 mm. Fore-
wing length 7.1-8.1 mm. Distance be-
tween antennal sockets 1.7-2.0 times di-
ameter of anterior ocellus. Antenna (Fig.
11) 15-17-segmented, apical 2 segments
sometimes fused. First flagellar segment
1.6-2.0 times wider than long. Second fla-
gellar segment 1.8-2.2 times wider apical-
ly than basally. Mesoscutellum sometimes
distinctly emarginate posteriorly (Fig. 17);
anterodorsal surface usually more con-
cave (Fig. 14). Mesepisternal tubercle as in
female. Tergum 8 posteriorly with deep
emargination (Fig. 20). Genitalia as in Fig.
18, paramere moderately broad, gonolaci-
nia not strongly hooked, penis valve trun-
cate.

134

Journal of Hymenoptera Research

■12,

-10. II

13 1«, 22 23-25

14-17,19,20,21,26

Figs. 10-26. Pliiloinastix luiucanoun: 10, female antenna; 11, male antenna; 12, 13, forewings; 14, male meso-
scutellum, surface contour as seen from rear of insect; 15, female mesoscutellum, surface contour as seen from
rear of insect, 16, female mesoscutellum, dorsal view; 17, male mesoscutellum, dorsal view; 18, male genitalia,
dorsal view; 19, lateral panel of first abdominal tergum; 20, apex of male abdomen, dorsal view; 21, apex of
ovipositor sheath, posteroventral view; 22-25, lancet; 26, mesepisternum, profile. Scale lines = 1.0 mm for 10-
17, 19-21, 26; = 0.1 mm for 18, 22-25.

VoLUMK 7, Number 2, 1998

135

Clypeus and labrum creamy white. Mid
lobe of mesoscutum anteriorly orange-yel-
low, posteriorly brown (Fig. 65). Lateral
lobe of mesoscutum entirely brown to
dark brown. Abdominal terga 5 and 6
without orange-yellow, lateral maculae.
Tergum 7 with lemon-yellow, lateral mac-
ula.

Type. — Holotype female. Cairns (ANIC,
on permanent loan from Macleay Muse-
um, University of Sydney, examined).

Material examined. — Queensland: 1 fe-
male, Myola, 1909 (ANIC); 10 females, 22
males, Kuranda, IV. 1902, R. E. Turner
(BMNH, USNM, QMBA); 1 female, same
locality, 3. V.-20. VI. 1913, R. E. Turner
(BMNH);1 male, same locality, 15. IV.
1931, A. N. Burns (MVMA); 1 female,
same locality, H. J. Carter (AMSA); 1 fe-
male, 2 males, 1.5 km SE Kuranda, 16-17.
V. 1980, I. D. Naumann, J. C. Cardale
(ANIC); 2 females, Barron Falls, via Ku-
randa, 16. VI. 1971, E. F. Riek (ANIC); 1
female. Cairns, 1905, H. Elgner (ANIC); 1
male. Cairns district, F. P. Dodd (SAMA);
1 female, Danbulla (forestry Reserve), 14.

V. 1959, G. W. S. (BMNH);1 female, Mt
Baldy,approximately 8 km W of Atherton,
13. 1. 1977, M. S. and B. J. Moulds (AMSA);
1 male, 12 miles (19.3 km) from Ivanhoe
(?Mine), 5. III. 1961, R. Straatman (ANIC);
1 female, Herberton, 17. 111. 1922 (QMBA);
1 female, same locality, 30. V. 1943, M. F.
Leask (QMBA); 4 females, same locality, 4.

VI. 1944, M. F. Leask (AMSA); 3 females,
Eubenangee, 14. V. 1950, G. Brooks
(MVMA); 1 female, Bartle Frere, 5. V. 1928
(QDPI); 1 female, Laceys Creek, Mission
Beach, 22. IV. 1970, S. R. Curtis (ANIC); 1
female, Tully, 24. IV. 1931, A. N. Burns
(MVMA); 1 female, 4 miles (6.4 km) W
Paluma, 13. IV. 1969, I. F. B. Common, M.
S. Upton (ANIC); 2 females, Paluma, II.-
IV. 1992, R. Jackson (AN1C);18 females.
Little Crystal Creek, Mt Spec, 16. V. 1971,
E. F. Riek (ANIC). Unlocalised: 2 females,
"North Queensland," V. 1944, M. F. Leask
(QMBA); 1 female, "Atherton Tableland,"
14. III. 1934 (QDPI); 1 female, "Atherton

Tableland," 19. 111. 1958, N. H. L. Krauss
(USNM); 1 female (QMBA).

Distribution. — See Fig. 76. The series
split between the BMNH, USNM and
QMBA is labelled "Cairns Kur 4. 02." 1
have interpreted this as meaning that the
specimens were collected at Kuranda, a
well known entomological collecting lo-
cality approximately 15 km north west of
Cairns. Locality labels for specimens col-
lected at Kuranda during the early 1900s
commonly bear the additional information
"Cairns" or "Cairns district." The collec-
tor's name is omitted from the USNM and
QMBA specimens. Forsius (1927) recorded
P. nancarroivi (and P. macleaii) from speci-
mens collected at Dorrigo, in north eastern
New South Wales by W. Heron and de-
posited in the SAMA. 1 have re-examined
these specimens and all are clearly iden-
tifiable as macleaii.

Discussion. — One male from Mt Lewis,
north Queensland listed below under
"Other material examined", differs signif-
icantly from the description of P. nancar-
roivi males given above. In the Mt Lewis
male the mesoscutellum is rather truncate
posteriorly and thus approaches the tu-
berculate condition of P. macleaii; tergum
8 is only weakly emarginate; the dark
brown coloration of the vertex does not
extend to the posterior surface of the head;
and the the clypeus, labrum and meso-
scutum are all orange-yellow. The speci-
men may represent an extreme of varia-
tion in P. nancarroivi or an additional, un-
described species. It is perhaps not
surprising that an anomalous Pbilomastix
occurs on Mt Lewis. The mountain is the
most northerly, known locality for the ge-
nus. It is part of the Carbine Tableland, a
discrete, rainforest-covered upland re-
markable for the uniqueness of its fauna.
The Carbine Tableland supports the high-
est number of endemic vertebrate species
in the Queensland Wet Tropics (Nix 1991)
and Mt Lewis itself harbours several en-
demic species of insects: a stag beetle of
the genus Sphaenognathus Buquet which is

136

Journal of Hymenoptera Research

known elsewhere only from the Black-
down Tableland in central Queensland
and Andean South America (Moore 1978;
Monteith 1996), a primitive leafhopper
representing a tribe otherwise unknown
from Australia but recorded from Mada-
gascar, New Zealand, Chile and Juan Fer-
nandez and a species of flightless dung
beetle (Kikkawa et al. 1981).

Larval host plants. — Rosaceae: Rubus ros-
ifolius Sm. (Leask 1943, 1944); Riilms hillii
F. Muell.(Leask 1944; Bruzzese 1980); Ru-
biis moluccamis L. (Bruzzese 1980). Rham-
naceae: Alphitonia petriei (Jackson 1993).

Other material examined. — QUEENS-
LAND: 1 male, Mt Lewis, via Julatten, 4.
V. 1970, S. R. Curtis (ANIC).

Philomastix tnacleaii (Westwood)

(Figs. 27^7, 62, 63 76)

Perga madeaii Westwood 1880: 372-373, Plate
XXXV, fig. 2.; Kirby 1882: 26.

Pergn madeayi : Dalla Torre 1894: 351 (unjusti-
fied emendation).

Heptacola madeayi : Konow 1905a: 167.

Philomastix madeaii : Froggatt 1918: 671; Smith
1978: 159; Bruzzese 1980: 4; Smith 1980: 342;
Macdonald and Ohmart 1993: 493-494; Nau-
mann 1993: 8, 115, 183.

Philomastix madeayi : Morice 1919: 248, 265, 287-
288; Tillyard 1926: 265, Plate 21, fig. 2; For-
sius 1927:283 (part).

Philomastix nancarroivi : Forsius 1927: 283 ( mis-
identification).

Philomastix glabra Froggatt 1890a: 489-490;
Froggatt 1892; 201; Konow 1898: 248; Frog-
gatt 1901; 1070; Schulz 1906: 83-84. (Synon-
ymised, as glaber, with madeaii by Froggatt
1918: 671.)

Philomastix glaber : Froggatt 1890b: 696; Froggatt
1893: 201; Froggatt 1901: 1070, Plate; Froggatt
1907: 73, Plate XI; Froggatt 1918: 671.(Mis-
spelling of glabra).

Female. — Body length 12.0-14.3 mm.
Forewing length 12.2-13.7 mm. Distance
between antennal sockets 2.2-2.6 times di-
ameter of anterior. Antenna (Fig. 27) 17-
19-segmented; apical 2-3 segments some-
times fused so that antenna apparently
15-18-segmented. First flagellar segment

2.5-2.9 times longer than wide. Second fla-
gellar segment 1.6-1.9 times wider apical-
ly than basally. Mesoscutellum usually
with posterolateral tubercles (Figs. 30-34);
anterodorsal surface distinctly (Fig. 36) or
weakly (Fig. 35) concave. Mesepisternal
tubercle strong (Fig. 38). Posterolateral
margin of first abdominal tergum weakly
curved (Fig. 37) or straight, spiracle sepa-
rated from margin by a distance 0.9-1.1
times maximum diameter of spiracle. Ovi-
positor sheath not posteromedially den-
tate (Fig. 46). Lancet as in Figs. 39^2,
about 5 ctenidia terminating in dorsal
tooth.

Flagellum orange-yellow. Mid lobe of
mesoscutum usually with large, brown
macula occupying more than half of
length of lobe (Fig. 62); mid-lobe some-
times uniformly orange-yellow. Lateral
lobe of mesoscutum with large, dark
brown to black macula occupying most of
length of lobe. Mesepisternum usually
with broad, brown band adjacent to ster-
nopleural suture. Mesosternum usually
brown to anterior margin and sternopleur-
al sulcus, sometimes entirely orange-yel-
low. Metascutum brown to dark brown,
usually distinctly darker than cenchri.
Forewing (Fig. 47) with narrow, trans-
verse, brown band; cell IM rarely com-
pletely brown. Abdomen predominantly
dark coloured with some metallic reflec-
tions. Abdominal tergum 1 dark brown to
black. Tergum 2 brown to black or metal-
lic blue, on each side with large, cream,
macula. Terga 3 and 4 dark brown, black
or metallic blue, with smaller, cream lat-
eral maculae. Terga 5, 6 and 7 each with
large, cream, lateral macula; maculae nar-
rowly separated by dark brown or black,
or contiguous. Tergum 8 anteriorly dark
brown to metallic blue (usually concealed
by preceding tergum), posteriorly cream.
Tergum 9 dark brown to black. Abdomi-
nal sterna 1 and 2 at least in part yellow
or cream. Sterna 3-6 dark brown to black.
Ovipositor sheath orange-yellow, with or
without dark brown markings.

Volume 7, Number 2, 1998

137

,27-29

30-38

Figs. 27-38. P/ii/oHMs/i.v rimcleair. 27, female antenna; 28, male antenna, basal antennomeres, frontal view; 29,
male antenna, lateral view; 30-34, mesoscutellum, dorsal view, variation in shape; 35, 36, mesoscutellum,
variation in surface contour as seen from rear of insect; 37, lateral panel of first abdominal tergum; 38, mes-
episternum, profile. Scale lines =1.0 mm.

Male.— Body length 11.0-12.8 mm. Fore-
wing length 7.4-11.4 mm. Distance be-
tween antennal sockets 1.0-1.3 times di-
ameter of anterior ocellus. Antenna (Figs.

28, 29) 18-21-segmented; apical 2 seg-
ments sometimes fused (occasionally
without trace of sutures) so that antenna
apparently 17-21-segmented. First flagel-

138

Journal of Hymenoptera Research

Figs. 39^7. Plntiimastix madeaii: 39^2, lancet; 43, 44, apex of male abdomen, dorsal view; 45, male genitalia,
dorsal view; 46, apex of ovipositor sheath, posteroventral view; 47, forewing. Scale lines = 0.1 mm for 39-42,
45; = 1.0 mm for 43, 44, 46, 47.

lar segment 1.2-1.6 times longer than
wide. Second flagellar segment 1.9-2.4
times wider apically than basally. Meso-
scutum and mesepisternal tubercle as in
female. Tergum 8 posteriorly with shallow
emargination (Fig. 43, 44). Genitalia as in
Fig. 45, paramere broad, gonolacinia not
strongly hooked, penis valve apically
rounded.

Clypeus and labrum orange-yellow.

Mesoscutum entirely brown to dark
brown (Fig. 63). Abdominal terga 5 and 6
with orange-yellow, lateral macula. Ter-
gum 7 with lemon yellow, lateral macula.
Types. — P. madeaii: holotype female,
"Australia" (UMO; examined). P. glabra: 4
syntypes, "Australasia," Dunoon, Rich-
mond River (ANIC, on permanent loan
from Macleay Museum, University of
Sydney, examined).

Volume 7, Number 2, 1998

139

Other material examined. — Queensland: 1
male, Mt William, Eungella National Park,
1100-1200 m,19. IV. 1979, G. B. Monteith
(QMBA); 1 male, Olmara Hills, via Dal-
rymple Heights, 1000m, 5. IV. 1975, D. K.
McAlpine (AMSA); 1 female, Eungella, 25.
IV. 1931, W. A. McDougall (QDPI); 1 fe-
male. Three Moon Creek, Kroombit Tops,
3-4 II. 1984, G. B. Monteith, C. Hagen, D.
Yeates (QMBA); 1 female, Kroombit Tops,
1000-1100 m, 22-26. II. 1982, G. B. Mon-
teith, G. Thompson, D. Yeates (QMBA); 1
female. Forest Station, Bulburin State For-
est, 600 m, 12-13. IV. 1974, I. D. Naumann
(UQIC); 2 females, Imbil, 13. V. 1937, 30.
III. 1938, A. R. Brimblecombe (QDPI); 1 fe-
male, Palmwoods, 15. IV. 1911, Miss Ede
(QMBA); 1 female, Montville, C. Deane
(UQIC); 1 female, 6 males, Mt Kiangarow,
Bunya Mountains, 27. 1.1993, K. J. and C.
L. Lambkin (QMBA, ANIC); 1 female,
Bunya Mountains National Park, 11-13.
XII. 1979, M. Schneider (UQIC); 1 female,
Mt Glorious, 4. IV. 1959, K. H. L. Key
(ANIC); 1 female, same locality, II. 1960, J.
Bryan (UQIC); 2 females, same locality,
853 m, 13. III. 1960, R. Straatman (ANIC);
1 female, 1 male, same locality, 5-8. II.
1961, 10. IV. 1962, J. L. and M. Gressitt,
Malaise trap (BPBM, USNM); 2 females,
same locality, 17. I. 1963, T. Brooks
(BPBM, USNM); 1 female, same locality, 1.
II. 1968, H. McDougall (UQIC); 1 male, Mt
Tenison-Woods, 4. II. 1983, G. Daniels
(UQIC); 1 female, Highvale, 12. III. 1960,
A. Cameron (UQIC); 1 female, Brisbane,
24. III. 1929, L. F. (MVMA); 1 male, same
locality. III. 1953, L. W. Rule (UQIC); 5 fe-
males, Brookfield, 30. III. 1990, 2. IV. 1990,
10. IV. 1993, 20. IV. 1994, J. Grigg (UQIC,
ANIC); 1 male, Toowoomba, 30. XII. 1917,
J. A. Bock (UQIC); 1 female, Mt Tambori-
ne, 1893, C. Wild (QMBA); 1 female, same
locality, 20. II. 1911, W. W. Froggatt
(ANIC); 1 female, same locality, 1923, W.
H. Davidson (QMBA); 1 female, same lo-
cality, 21. II. 1927, H. Hacker (QMBA); 1
female, same locality, C. Deane (UQIC); 1
female, same locality (QDPI); 1 female.

Boonah, 5. FV. 1948 (QDPI); 1 female, Can-
ungra, 7. IV. 1928 (QDPI); 1 female, Cun-
ningham's Gap, 800m, R. Eastwood
(UQIC); 1 female, same locality. III. 1972,
R. Baldwin (QMBA); 1 female, 1 male,
Beechmont, 1. II. 1972, A. and G. Daniels
(AMSA); 1 male, Mt Huntley, 1250 m, 29-
30 I. 1993, G. B. Monteith (QMBA); 1 male.
Bald Mountain, 1219 m, 28-31 I. 1972, 1.
D. Naumann (UQIC); 3 females. Upper
Nerang, III. 1891, H. Tryon (QMBA,
QMBA); 1 female. Upper Currumbin, 27.
IV. 1932, L. Franzen (MVMA); 3 females,
1 male, McPherson Range, XI. 1928, A. J.
Turner (MVMA); 1 male, same locality, H.
Tryon (QDPI); 6 females, Lamington Na-
tional Park, (some labelled 300 feet = 914
m), 2-3. I. 1921, 26-27. II. 1921, 1-11. III.
1921 (QMBA); 7 females, same locality,
XII. 1921, H. Hacker (QMBA, ANIC); 2
males, same locality, 900-lOOOm, 16-18. II.
1964, J. Sedlacek (BPBM); 1 female, same
locality, 930 m, 3 II. 1983, W. C. Paine
(ANIC); 2 females, same locality, 914 m,
6-7. III. 1980. H. E. Evans, A. Hook
(UQIC); 1 female, Binna Burra, I. 1943
(QDPI); 3 females, 2 males, Springbrook,
1930, R. Blackwood (MVMA, ANIC); 1 fe-
male, same locality, 2. XI. 1961, 1. C. Cun-
ningham (UQlC).New South Wales: 1 fe-
male, 2 males, Mt Clunie, 16. XII. 1972, I.
D. Naumann (UQIC); 4 females, Tweed
River, 1897 and no date (BRI, USNM,
BMNH); 1 female, same locality, 1920, W.
W. Froggatt (BMNH); 1 female, Wiangaree
State Forest, 12. II. 1978, K. Walker
(UQIC); 2 males, Richmond River (BRI); 1
male, Huonbrook, 2. III. 1964, D. K.
McAlpine (AMSA); 6 females, 1 male.
Glen Innes, 17 II. 1974, M. S. Moulds
(AMSA, ANIC); 1 female, same locality,
19. I. 1975, R. Gallagher (BRI); 2 females.
Platypus Flat camp. Wild Cattle Creek
State Forest, 7 IV. 1993, C. Reid (ANIC); 1
female, 1 male, Guyra, II. 1949, A. Dyce
(ANIC); 2 females, Ulong, III. 1920, W.
Heron (AMSA); 11 females, Brooklana,
1929, W. Heron (BMNH, AMSA, ANIC); 1
female, Coffs Harbour, 11. I. 1950, F. D.

140

Journal of Hymenoptera Research

(MVMA); 11 females, Dorrigo, W. Heron
(BMNH, MVMA, SAMAA, AMSA); 1 fe-
male, same locality, 914 m, G. Heron
(USNM); 2 females, same locality (QMBA,
BMNH); 1 male, same locality, 914 m, 17.
II. 1932, P. J. Darlington (USNM); 1 male,
same locality, 12 II. 1968, D. H. Colless
(ANIC); 1 female, same locality, 14. II.
1981, D. A. Doolan (AMSA); 1 female,
same locality, 13. II. 1984, L. Masner
(CNC); 4 females, 1 male. Deer Vale, 12-
13. I. 1931, 13. I. 1933, A. N. Burns
(MVMA); 2 females, same locality, 30. I.
1972, G. Daniels (AMSA); 1 female, 4
males, Ebor, I. 1934, F. E. Wilson (MVMA,
ANIC); 3 females, same locality, 12. XII.
1962, T. V. Bourke (BRI); 2 females, 3
males, Armidale, 5. II. 1915 (QMBA); 3 fe-
males, Bellangry, 2. V. 1894, W. W. Frog-
gatt (ANIC, BRI, MVMA, CNC); 4 fe-
males. Hanging Rock, 7. I. 1955, K. M.
Moore (FCNI); 1 female. Elands, 30. 1. 1928
(BRI); 1 female, Comboyne Scarp, near
Upper Lansdowne, 6. IV. 1987, D. K.
Mc Alpine, S. Day, R. de Keyzer (AMSA);
1 female. Dingo State Forest, 26-27. II.
1981, G. and T. Williams (AMSA); 1 fe-
male. Bay's Hill, Taree, 28. III. 1992, G.
Williams (AMSA); 9 females, 2 males,
Tuncurry, 21 III. 1931, 15-25. III. 1932, no
date, J. Parkes (AMSA, ANIC); 1 female.
Upper Allyn River, 6. IV. 1958, R. Mackay
(AMSA); 1 female. Upper Allyn, near Ec-
cleston, 10 III. 1970, D. K. McAlpine, G.
Holloway (AMSA); 1 female, 1 male, same
locality, 26. II. 1970, D. K. McAlpine
(AMSA); 1 female, same locality, 16. II.
1967, D. K. McAlpine (AMSA); 6 females,
Maitland, 1892, W. W. Froggatt (ANIC,
MVMA, USNM, BMNH); 1 female, Olney
State Forest, 15. III. 1986, J. Grigg (UQIC);
5 females, Ourimbah, IV. 1904, S. W. Jack-
son (AMSA); 1 female, Narara, 23 I. 1911
(AMSA); 1 female, same locality, 2. III.
1950, P. C. Hely (BRI); 2 males. Upper
Colo, 10. III. 1990, G. R. Brown, M. A. Ter-
ras (BRI); 4 females, 8 males, Mt Wilson,
4. I. 1931, A. N. Burns (MVMA); 1 female,
same locality, 7 II. 1959, D. K. McAlpine

(AMSA); 1 female, same locality, II. 1921
(AMSA); 1 male, same locality, 1067 m, I.
1932, P. J. Darlington (USNM); 1 female.
Hartley Vale, 29. III. 1975, G. Daniels
(AMSA); 3 females, Mt York, 21. III. 1964,
D. K. McAlpine (AMSA, ANIC); 2 males,
Katoomba, 26. 1. 1955, K. M. Moore
(FCNI); 2 females, 1 male, same locality,
21. II. 1969, G. Hardy (AMSA); 2 females
(with eggs), Woodford, 28. II. 1984, M. Hill
(BRI); 1 female, 2 males, Beecroft, 18. II.
1968, O. M. Williams (BRI, ANIC); 2 fe-
males, same locality, 18. III. 1967, C. E.
Chadwick (BRI, ANIC); 5, females. Glen-
brook, III. 1994, L. Turton (BRI, ANIC); 1
female, 2 males. Lane Cove, 27. IV. 1946,
30. III. 1947, 3. IV. 1948 (AMSA); 1 male,
Sydney, 13. III. 1932, G. A. Waterhouse
(MVMA); 1 female, same locality. III. 1949,
A. Dyce (ANIC); 1 female, same locality,
III. 1977, D. Clyne (ANIC). 1 male. Heath-
cote, 20. III. 1952, L. Cascysand (BRI); 1
female, Heathcote National Park, 24. FV.
1994, Cowdrey (AMSA); 2 males, Mt Kei-
ra, 23. II. 1983, G. A. Holloway (AMSA); 1
male, Jamberoo Mtn, 16. II. 1963, C. E.
Chadwick (BRI); 1 male, Jamberoo, 11. 1.
1950 (AMSA). Unrealised: 1 female, 1
male, W. Heron (BMNH); 2 females, 1
male (QDPI); 1 female (MVMA); 3 fe-
males, 1 male (UQIC); 1 female (QMBA);
1 female (ANIC).

Distribution. — See Fig. 76. Several fe-
males in the ANIC and the BMNH are la-
belled "Brooklana Sydney." Since Brook-
lana and Sydney are approximately 400
km apart, clearly mislabelling has oc-
curred. Reliably labelled specimens indi-
cate that P. macleaii occurs at both Brook-
lana and Sydney. P. macleaii was not re-
corded from Victoria by Bruzzese (1980) in
a survey of insects on Rubus.

Larval host plants.— Rosaceae: Rubus mol-
uccanus L. (Froggatt 1893); Rubus rosifolius
Smith.

Discussion. — There is some variation in
wing venation. In the forewing, Rl mav
continue beyond the junction with Rs. In
the hindwing cross-vein m-cu may be

Volume 7, Number 2, 1998

141

48,49

■57, 5S
5*)-61

. 50-56

Figs. 48-61. Philomastix xn]itlioplnihi\: 48, female antenna; 49, male antenna; 50, mesoscutellum, dorsal view;
51, mesoscutellum, surface contour as seen from rear of insect; 52, mebepisternum, profile; 5.3, lateral panel
of first abdominal tergum; 54, apex of ovipositor sheath, dorsal view; 55, same, posteroventral view; 56, apex
of male abdomen, dorsal view; 57, male genitalia, dorsal view; 58-61, lancet. Scale lines = 1.0 mm for 48-56;
= 0.1 mm for 57-61.

142

Journal of Hymenoptera Research

Figs. 62-66. Philomastix spp.: 62, P. madeaii, female, dorsal view; 63, P. madcaii, male, dorsal view; 64, P.
nancarrowi, female, dorsal view; 65, P. nancarroun, male thorax (part), dorsal view; 66, P. nancarrowi, female
mesepisternum, lateral view. Scale lines = 1.0 mm.

present or absent. The mesoscutellar tu-
bercles are usually strong (Figs. 30, 31) but
may be weak (Fig. 32), absent on one side
(Fig. 33) or absent altogether (Fig. 34). Cell
IM is usually at least in part hyaline prox-
imally, but in the female from Eungella
cell IM is entirely dark and the infuscation
extends to cell R.

Philomastix xanthophylax Naumann
and Groth, sp. n.

(Figs. 1, 48-61, 67-74, 76)
Female. — Body length 9.0-12.0 mm.
Forewing length 9.4—11.5 mm. Distance
between antennal sockets 2.2-2.6 times
greater than diameter of anterior ocellus.
Antenna (Figs. 48, 67) 14-16 segmented;

Figs. 67-71. Philoinastix xautlwpln/liix: 67, female, apical antcnnomcres; b8, left mandible; 69, right mandible;
70, labium, maxilla; 71, lancets, ventral view. Scale line = 1.0 mm, for 68-70; see 48, 58 for scale to 67, 71.

Volume 7, Number 2, 1998

Figs. 72-74. Pluliiiiiii-.li\ \iintlioplti/liix on Alphitonia excelsa; 72, two adult females, one guaiduii; fgg mass,
other guarding batch ot first instar larvae; 73, batch of first instar larvae; 74, third instar larva. Adult sawflies
in 72 each approximately 9.5 mm long; larvae in 73, 74 approximately 5 and 17 mm long respectively.

apical 2-5 segments sometimes fused so
that antenna apparently 12-13-segmented.
First flagellar segment 2.9-3.0 times longer
than wide. Second flagellar segment 1.8-
1.9 times wider apically than basally. Me-
soscutellum without posterolateral tuber-
cles (Fig. 50); anterodorsal surface more or
less flat (Fig. 51). Mesepisternal tubercle
weak (Fig. 52). Posterolateral margin of
first abdominal tergum weakly curved
(Fig. 53) or straight, spiracle separated
from margin by a distance 1.2-1.3 times
maximum diameter of spiracle. Ovipositor
sheath posteromedially dentate (Figs. 54,
55). Lancet as in Figs. 58-61, 71, about 8

ctenidia terminating in dorsal tooth, antior
teeth strong.

Flagellum black (Fig. 1). Mid lobe of
mesoscutum orange-yellow. Lateral lobe
of mesoscutum orange-yellow, sometimes
with small, brown macula less than half
as long as lobe. Mesepisternum without
brown band adjacent to sternopleural su-
ture. Mesosternum brown, with orange-
yellow band along anterior and lateral
margins. Metascutum and cenchri orange-
yellow. Forewing with narrow, transverse,
brown band; cell IM not completely
brown. Abdomen entirely orange-yellow.

Male. — Body length 7.3-8.9 mm. Fore-

0 750 150O

Kilometres

^^

Fig. 75. Distribution of Philomastiginae: woM distribution of Philomastix, Cerospastus and Ecopatus.

144

Journal of Hymenoptera Research

0

400 800

:

Kilometres

. 1

1
\

\

i\

Xj

'--^_/ 0 Philomastix macleaii

.:

y^ —

h 0

400 800

Kiiometres

■/ \

t

1

A

'^^,

. *i

/

"■-■J"

'^v^ ^a*

■---. i 9 Philomastix nancarrowl

y^ — '■ ^ Philomastix xanthophylax

-'-g'

Fig. 76. Australian distribution of Philomastix nancarroun, P. macleaii and P. xanthophylax.

wing length 6.5-8.8 mm. Distance be-
tween antennal sockets 1.3-1.7 times
greater than diameter of anterior ocellus.
Antenna (Fig. 49) 15-segmented. First fla-
gellar segment 1.7-1.8 times longer than
wide. Second flagellar segment 2.0-2.1
times wider apically than basally. Mesos-
cutellum without posterolateral tubercles;
dorsally slightly concave. Mesepisternal
tubercle weak. Abdominal tergum 8 pos-
teriorly with deep emargination (Fig. 56).
Genitalia as in Fig. 57, paramere slender,
gonolacinia strongly hooked, penis valve
apically rounded.

Clypeus and labrum lemon-yellow.
Mid-lobe of mesoscutum orange-yellow.
Lateral lobe of mesoscutum entirely
brown to dark brown. Abdominal terga 5
and 6 without orange-yellow, lateral mac-
ulae. Tergum 7 with lemon-yellow, lateral
macula.

Material examined. — Holotype female.

27.28S 151.56E, 10 km N Toowoomba,
Queensland, 12. IV. 1992, H. Groth
(ANIC). Paratypes: Queensland: 2 fe-
males, same data as holotype (ANIC,
HGCN); 8 females, same locality and col-
lector as holotype, 3. IV. 1991, 1. III. 1992,
29. III. 1992, 16. III. 1992 (ANIC, BMNH,
USNM, HGCN); 4 males, same locality
and collector as holotype, but labelled
"Highfields," reared from eggs collected
V. 1992, emerged as adults 30. III. 1993, 22.
IV. 1993, 24. IV. 1993, 26. IV. 1993 (ANIC,
HGCN); 3 females, 4 males, Brookfield, 22.
III. 1994, 29. III. 1994, 3. IV. 1994, J. Grigg
(UQIC, ANIC); 2 females, Bellbird Park,
Brisbane, IV. 1994, R. Nattrass (QMBA).

Etymology. — The species name is de-
rived from the Greek words xanthos, yel-
low or golden, and phylax, a guard, with
reference to the maternal guarding behav-
iour common to all species of Philomastix.

Distribution. — See Fig. 76.

Volume 7, Number 2, 1998

145

Larval food plants. — Alpliitonia excelsa
(Fenzl) Benth. (Rhamnaceae).

Discussion. — There is conspicuous vari-
ation in wing venation within the type se-
ries. In the forewing: (1) Rl sometimes
continues as a short spur beyond the junc-
tion of Rl and Rs; (2) there may be one,
two or no cross-veins between C and R;
(3) there may be an incomplete cross-vein
distal to 3r-m; and (4) a diagonal vein
sometimes defines a small, triangular cell
in the anterobasal comer of cell 3M. In the
hindwing cross-vein m-cu is rather vari-
able: (1) it may be present or absent; (2) it
may curve smoothly into CuA, in which
case there is no distal abscissa of CuA; (3)
it may join M before or after the junction
of cross-vein 2r-m and M.

CEROSPASTUS Konow

Cerospastus Konow 1899: 404--i05; Konow 1905:
36-37; Rohwer 1911: 76. Benson 1935: 224.
Benson 1938: 379; Pagliano and Scaramozzi-
no 1990: 58; Smith 1990: 21-23; Abe and
Smith 1991: 18. Type species: Cerospastus vol-
upis Konow (by monotypy).

Ceratospastus: Schuiz 1906: 84 (unjustified emen-
dation).

Female. — Vertex conspicuously setose.
Face with some fine microsculpture. Malar
space narrower than diameter of anterior
ocellus. Antenna (Fig. 3) 14-(-20, Smith
1990) segmented, weakly serrate, weakly
clavate. Right mandible simple (Smith
1990). Maxillary palp (Fig. 9) 6-segmented,
filiform, without sensory cup. Labial palp
4-segmented, without sensory cup. Labi-
um tri-lobed. Thorax dorsally consci-
cuously setose. Notauli, median mesoscu-
tal line deeply impressed. Mesoscutellum
not swollen, posterior margin visible from
above. Mesepisternum without tubercle.
Metascutellum not band like. Forewing
(Fig. 4) with closed radial cell and 4 cu-
bital cells; median and second cubital cells
each with nygma. Abdominal terga not
conspicuously setose. Second tergum pre-
dominantly fine transversely striate. Cer-

cus present. Ovipositor sheath not strong-
ly expanded posteriorly.

Male. — Antenna 20-segmented (19-21
according to Smith 1990), strongly serrate.
Median cell of forewing without nygma.
Eighth tergum posteriorly with very deep,
broad emargination (Fig. 7).

Cerospastus volupis Konow

(Figs. 3, 4, 7, 9, 76)

Cerospastus volupis Konow 1899: 404; Konow
1905: 37; Rohwer 1911: 76; Smith 1978: 160;
Oehike and Wudowenz 1984: 419; Pagliano
and Scaramozzino 1990: 58; Smith 1990: 22-
23; Smith 1993: 11.

Female. — Body length 10.0-11.5 mm.
Forewing length 11.0-12.0 mm. Distance
between antennal sockets 1.2-1.7 times
greater than diameter of anterior ocellus.
First flagellar segment 2.5 times longer
than wide. Second flagellar segment 1.7-
2.0 times wider apically than basally. Me-
soscutellum without posterolateral tuber-
cles, posterior margin convex; dorsal sur-
face weakly convex. Mesepisternum with-
out tubercles. Posterolateral margin of
first abdominal tergum smoothly curved
or slightly angulate, spiracle separated
from margin by a distance 0.5-1.3 times
maximum diameter of spiracle. Ovipositor
sheath not posteromedially dentate. Lan-
cet as in Smith (1990: Fig. 34).

Flagellum very pale brown, scape and
pedicel orange-yellow. Either, head pre-
dominantly orange-yellow; upper frons
with transverse brown joining upper ex-
tremities of compound eyes and encom-
passing ocellar triangle; or most of frons,
lowermost gena, vertex medially brown.
Mandibles brown, remaining mouthparts
orange-yellow. Thorax and legs predomi-
nantly orange-yellow to cream. Mesono-
tum orange-yellow, with brown maculae
occupying either most of length of mid
and lateral lobes or only anterior half of
lateral lobe. Cenchri, metascutum, most of
mid and hind tarsi, first abdominal ter-
gum and sometimes also ventral surfaces
of thorax brown to pale brown. Remain-

146

Journal of Hymenoptera Research

der of abdomen orange to yellow. Wings
hyaline with faint yellow Hnge.

Male. — Body length 7.5 mm. Forewing
length 7.3 mm. distance between antennal
sockets 0.7-0.8 times greater than diame-
ter of anterior ocellus. First flagellar seg-
ment 1.3 times longer than wide. Second
flagellar segment 2.4 times wider apically
than basally. Mesoscutellum, mesepister-
num as in female. Genitalia as in Smith
(1990: Fig. 31).

Head, thorax predominantly black to
dark brown. Antenna pale brown. Clype-
us orange-brown. Mandibles red-brown.
Maxilla, labium orange-yellow. Spiracular
lobe of pronotum orange-yellow. Legs or-
ange to yellow. Abdomen dorsally dark
brown to brown, ventrally orange to yel-
low. Wings hyaline with faint brown
tinge.

Type. — Lectotype female (designated by
Smith 1990), Valdivia, Chile, 1897, Loss-
berg (DEIC; examined).

Other material examined. — 1 female, 1
male, Parque Nac. Conguillio. Province
Temuco, Chile, 31. XII. 1976, O. Puentes
(USNM).

Distribution. — Known only from a few
specimens from central western Argentina
(Neuquen Province) and central Chile
(Cautin, Malleco, Maule and Valdivia).

Lariml liost plant. —Nothofagus sp. (Faga-
ceae) (Smith 1990).

ECOPATUS Smith

Ecopatiis Smith 1990: 23-24. Type species: Eco-
patus penai Smith (by original designation
and monotypy).

Female. — Vertex almost bare. Face al-
most completely smooth. Malar space
very slightly wider than diameter of an-
terior ocellus. Antenna (Fig. 2) 18-seg-
mented (20-21 according to Smith 1990),
filiform, not clavate. Right mandible with
2 inner teeth. Maxillary palp (Fig. 8) 6-seg-
mented, 4th segment apically distinctly
wider than other segments; apical seg-

ment without sensory cup. Labial palp 4-
segmented, apical segment without sen-
sory cup. Labium tri-lobed. Thorax dor-
sally almost bare of setae. Notauli, median
mesoscutal line shallow. Mesoscutellum
not swollen, posterior margin visible from
above. Mesopleuron without tubercle. Me-
tascutellum not band like. Forewing (Fig.
5) with open radial cell and 2-3 cubital
cells; median and second cubital cells each
with nygma. Abdominal terga dorsally
not conspicuously setose. Second tergum
with faint, reticulate microsculpture. Cer-
cus present. Ovipositor sheath not strong-
ly expanded posteriorly.
Male. — Unknown.

Ecopatiis penai Smith

(Figs. 2, 5, 6, 8)

Ecopatiis penai Smith 1990: 24-25.

Female. — Body length 6.0-7.0 mm. fore-
wing length 7.7 mm. distance between an-
tennal sockets 2.3-2.4 times greater than
diameter of anterior ocellus, first flagellar
segment 3.6-3.7 times longer than wide.
Second flagellar segment 1.1-1.2 times
wider apically than basally. Mesoscutel-
lum without tubercles; anterodorrsal sur-
face convex. Mesepisternum without tu-
bercle. Posterolateral margin of first ab-
dominal tergum sinuate (Fig. 6), spiracle
separated from margin by distance 4.6
times maximum diameter of spiracle. Ovi-
positor sheath not medially dentate. Lan-
cet as in Smith (1990: Fig. 39).

Head, thorax, abdomen predominantly
dark brown to black. Following orange,
orange-yellow to cream: scape or scape
and pedicel, frons just above and below
antennal sockets, clypeus, mandible (ex-
cluding red-brown teeth), maxilla, labium,
posterior margin of pronotum, tegula, ax-
illar sclerites, large macula anterodorsal to
extremity of mesepisternum, legs (except
pale brown extremities of tarsi), abdomen
ventrally. Wings uniformly strongly
brown tinged.

Ti/f^es. — Holotype female, Caramavida,

Volume 7, Number 2, 1998

147

Province Arauco, 5-10. II. 1953, L. Pena
(USNM; not seen). Paratypes: 2 females,
same data as holotype (not seen); 1 female,
Curacautin, Rio Blanco, 27-31. I. 1950, L.
Pefia (USNM) (examined).

Distribution. — Known only from a few
specimens from central Chile (Araucan
and Curacautin Provinces).

Larval host plants. — Unknown.

ACKNOWLEDGEMENTS

I thank Mr M. Moulds (AMSA), Mr P. Gillespie and
Mr J.Macdonald (BCRl), Mr D. G. Notfon (BMNH),
Dr H. Goulet (CNC), Drs A. Taeger and S. M. Blank
(DEIC), Ms C. Urquhart (FCNI), Dr K. Walker
(MVMA), Dr C. O'Toole (UMO), Dr C. Burwell
(QMBA), Mr J. Donaldson and Dr M. Elson-Harris
(QDPI), Ms J. Forrest (SAMAA), Mr G. Daniels
(UQIC) and Dr D. Smith (USNM) for the loan of ma-
terial. Mr N. Springate (c/ -Natural History Museum,
London, U.K.), Dr Goulet and Mr Macdonald kindly
commented on an earlier draft of this paper. Botanical
names were checked by Dr L.S. Springate (Royal Bo-
tanic Garden, Edinburgh, U.K.).

LITERATURE CITED

Anonymous. 1980. Insect pest occurrences in Tas-
mania, 1978/79. Tasmanian Department of Agri-
culture. Insect Pest Suroei/ 12: 1-39.

Abe, M. and D. R. Smith. 1991. The genus-group
names of Symphyta (Hymenoptera) and their
type species. Esakia 31: 1-115.

Ashmead. W. H. 1898. Classification of the horntails
and sawflies, or the sub-order Phytophaga. Ca-
nadian Entonwhgist 30: 225-232.

Benson, R. B. 1935. New Australian sawflies. Memoirs
of the Queensland Museum 10: 211-229.

Benson, R. B. 1938. On the classification of sawflies
(Hymenoptera Symphyta). Transactions of the
Royal Entomological Society of London 87: 353-384.

Bruzzese, E. 1980. The phytophagous insect fauna of
Ruhus spp. (Rosaceae) in Victoria, a study on the
biological control of blackberry ( Ruhus fructico-
sus L. agg.). Journal of the Australian Entoniotogwal
Societ]/ 19: 1-6.

Dalla Torre, C. G. 1894. Tenthredinidae incl. Urocer-
idae (Phyllophaga cSc Xylophaga). Catalogiis Hy-
menoftterorum 1:1-459.

Forsius, R. 1927. On some sawflies from the Austra-
lian region (Hymenoptera Tenthredinidae). Rec-
ords of the South Australian Museum 3: 283-308.

Francis, W. D. and G. M. Chippendale 1970. Austra-
tian Raiu'Forest Trees. Australian Government
Publishing Service: Canberra, xvi, 468 pp.

Froggatt, W. W. 1890a. Descriptions of a new genus
and two new species of Tenthredinidae. Proceed-

ings of the Lmnean Society of Neiv South Wales 5:
487^90.

Froggatt, W. W. 1890b. Catalogue of the described
Hymenoptera of Australia. Proceedijigs of the Lm-
nean Society of Nt-ic South Wales 5: 689-762.

Froggatt, W. W. 1892. (Notes and exhibits]. Proceed-
ings of the Lmnean Society of New South Wales 7:
201.

Froggatt, W. W. 1901. The pear and cherry slug (Er-
iocampa limacina, Retz), generally known as Selan-
dria cerasi, with notes on Australian sawflies. Ag-
ricultural Gazette of Neiv South Wales 12: 1063-
1073.

Froggatt, W. W. 1907. Australian Insects. Brooks: Syd-
ney, i-xiv, 1^149.

Froggatt, W. W. 1918. Notes on Australian sawflies
(Tenthredinidae). Proceedings of the Linnean Soci-
ety of New South Wales 43: 668-672.

Goulet, H. and J. T. Huber (1993). Hymenoptera of the
World. Agriculture Canada: Ottawa, viii, 668 pp.

lackson, R. 1993. Insect herbivory. Program and Ab-
stracts Australian Entomological Society Conference
and 24th A. G. M.: 46.

Johnson, N. F. 1992. Catalogue of world species of
Proctotrupoidea, exclusive of Platygastridae (Hy-
menoptera). Memoirs of the American Entomologi-
cal Institute 51: 1-825.

Kikkawa, J., G. B. Monteith and G. Ingram. 1981.
Cape York Peninsula: major region of faunal in-
terchange, pp. 1695-1742. In A. Keast (ed.) Eco-
logical Biogeography of Australia. W. Junk: The
Hague.

Kirby, W. F. 1882. List of Hymenoptera loith Descriptions
and Figures of the typical Specimens in the British
Museum. Vol.1. British Museum: London, xxviiii,
450 pp., 16 plates.

Konow, F. W. 1898. Uber die Tenthrediniden-tribus
Lophyrini. Entomologische Nachrichten. Berlin 30:
248-249.

Konow, F. W. 1899. Neue Siidamerikanische Tenthre-
dinidae. Anales del Museo Nacional de Buenos Aires
6: 397-417.

Konow, F. W. 1905a. Drei neue Syzygoniides aus
Australien. Zeilschrift fiir Systematische Hymenop-
terologie und Dipterologie 5: 166-169.

Konow, F. W. 1905b. Hymenoptera Fam. Tenthredi-
nidae. Genera Insectorum 29: 1-176.

Kudo, S., K. Maeto and K. Ozaki 1992. Maternal care
in the red-headed spruce web-spinning sawtly,
Cephalcia isshikii (Hymenoptera: Pamphiliidae).
Journal of Insect Behavior 5: 783-795.

Lazarides, M. and B. Hince 1993. CSIRO Handbook of
Economic Plants of Australia. CSIRO: Melbourne,
xiv, 330 pp.

Leask, M. F. 1943. Records of two sawflies. The North
Queensland Naturalist 11 (69): 2.

Leask, M. F. 1944. Habits of sawflies (Philomastix

148

Journal of Hymenoptera Research

nancarrowi). Hymenoptera, Tenthredinidae. The
North Queensland Nahmilist 11(72): 1-3.

Macdonald, J. and C. P. Ohmart. 1993.Life history
strategies of Australian pergid sawflies and their
interactions with host plants, pp. 485-502. In M.
R. Wagner and K. F. Raffa (eds) Snvflij Life His-
tory Adaptations to Woody Plants. Academic Press:
San Diego, xx, 581 pp.

Monteith, G. B. 1996. Recent exciting insect discov-
eries. Queensland Museum Association Neiosletter
11(2): 3.

Moore, B. P. 1978. A new Australian stag beetle (Co-
leoptera: Lucanidae) with Neotropical affinities.
Journal of the Australian Entomological Society 17:
99-103.

Morice, F. D. 1919. Notes on Australian sawflies, es-
pecially the "Authors' Types" and other speci-
mens in the British Museum of Natural History
and the Hope Collections of the Oxford Univer-
sity Museum; with diagnostic synopses of the
genera and species, and photographs illustrating
their structural characters. Transactions of the En-
tomological Society of London 66: 247-333.

Naumann, I. D. 1984. An apterous female sawfly (Hy-
menoptera: Symphyta) from Papua New Guinea.
Systematic Entomology 9: 339-349.

Naumann, I. D. 1991. Hymenoptera. pp.916-1000. /;;
CSIRO (ed.) The Insects of Australia, 2nd edition.
Melbourne University Press: Carlton.

Naumann, I. D. 1993. CSIRO Handbook of Australian
Insect Names. CSIRO: East Melbourne. 200 pp.

Naumann, 1. D. 1994. Systematic and Applied Entomol-
ogy. Melbourne University Press: Carlton. 484

PP-

Naumann, 1., J. C. Cardale, R. W. Taylor and J. Mac-
donald. 1994. Type specimens of Australian Hy-
menoptera (Insecta) transferred from the Ma-
cleay Museum, University of Sydney, to the Aus-
tralian National Insect Collection, Canberra. Pro-
ceedings of the Linnean society of New South Wales
UA: 69-72.

Nix, H. 1991. Biogeography: pattern and process. Ko-
warri 1: 11-39.

Oehlke, J. and J. Wudowenz. 1984. Katalog der in den
Sammlungen der Abteilung Taxonomie der In-
sekten des Institutes fur Pflanzenschutzfor-
schung, Bereich Eberswalde (ehemals Deutsches
Entomologisches Institut), aufbewahrten typen-
XXll (Hymenoptera: Symphyta). Beitriige zur En-
tomologic 34: 363-420.

Pagliano, G. and P. Scaramozzino. 1990. Elenco dei
Generi di Hymenoptera del niondo. Memorie del-
la Societa Entomologica Italiana 122: 3-210.

Riek, E. F. 1970a. Hymenoptera. pp.867-959.//i CSIRO
(ed.) The Insects of Australia, 1st edition. Mel-
bourne University Press: Carlton.

Riek, E. F. 1970b. A redefinition of the subfamily Phy-
lacteophaginae with description of a new genus
and species (Hymenoptera: Symphyta: Pergi-
dae). Journal of the Australian entomological Society
9: 215-218.

Rohwer, S. A. 1911a. A classification of the suborder
Chalastrogastra of the Hymenoptera. Proceedings
of the Entomological Society of Washington 13: 215—
226.

Rohwer, S. A. 1911b. II. The genotypes of the sawflies
and wood wasps, or the superfamily Tenthredi-
noidea. Technical Series. Bureau of Entomology,
United States Department of Agruidture 20:69-109.

Schuiz, W. A. 1906. Spolia Hymenopterologica. Pader-
born. 355pp.

Smith, D. R. 1978. Suborder Symphyta (Xyelidae, Par-
archexyelidae, Paramphiliidae, Xyelydidae, Kar-
ativitidae, Gigasiricidae, Sepulcidae, Pseudosiri-
cidae, Anaxyelidae, Siricidae, Xiphydriidae, Pa-
roryssidae, Xyelotomidae, Blasticotomidae, Per-
gidae). Hymenopterorum Catalogus 14: i-iii, 1-193.

Smith, D. R. 1980. Pergidae (Hymenoptera) from
New Guinea and Australia in the Bishop Muse-
um. Pacific Insects 22: 329-346.

Smith, D. R. 1988. A synopsis of the sawflies (Hy-
menoptera: Symphyta) of America south of the
United States: introduction, Xyelidae, Pamphili-
idae, Cimbicidae, Diprionidae, Xiphydriidae, Sir-
icidae, Orussidae, Cephidae. Systematic Entomol-
ogy 13: 205-261.

Smith, D. R. 1990. A synopsis of the sawflies (Hy-
menoptera, Symphyta) of America south of the
United States: Pergidae. Rei'ista Brasdeira de En-
tomologia 34: 7-200.

Smith, D. R. 1992. A synopsis of the sawflies (Hy-
menoptera: Symphyta) of America south of the
United States: Argidae. Memoirs of the American
Entomological Society 39: 1-201.

Smith, D. R. 1993. Systematics, life history, and dis-
tribution of sawflies.pp. 3-32. In M. R. Wagner
and K. F. Raffa (eds) Sawfly Life History Adaptions
to Woody Plants. Academic Press: San Diego, xx,
581 pp.

Smith, D. R. 1995. A new species of Xiphydriidae
(Hymenoptera) from Chile. Revista Chilena de En-
tomologui 22: 21-24.

Tillyard, R. J. 1926. The Insects of Australia and Nezo
Zealand. Angus and Robertson: Sydney, xvi, 560
pp.

Watson, J. A. L. and G. Theischinger. 1984. Regions
of taxonomie disjunction in Australian Odonata
and other freshwater insects. Odonatologua 13:
147-157.

Westwood, J. O. 1880. A monograph of the sawflies
composing the Australian genus Perga of Leach.
Proceedings of the Zoological Society of London :359-
379.

J. HYM. RES.
Vol. 7(2), 1998, pp. 149-156

A New Species of Ibalia from Borneo, with a Revised Phylogeny and
Historical Biogeography of Ibaliidae (Hymenoptera: Cynipoidea)^

Zhiwei Liu

Department of Entomology, Swedish University of Agricultural Sciences,
P.O. Box 7044, S-75007, Uppsala, Sweden

Abstract. — Ibalia kalimantanica Liu from eastern Kalimantan, Borneo, is described here as a
new species. Reanalysis of a previously published character matrix of the family Ibaliidae with
the new species included shows that /. kalimantanica belongs to the subgenus Tremibalia and is the
sister species to the clade of ((I.mirabilis, I. japonica), I. hunanica). Biogeographical analysis of the
expanded data set strengthened support for an earlier hypothesis concerning the historical bio-
geography of the Ibaliidae postulating early radiation of the family in the eastern Palaearctic —
Oriental region. The separation of the 7. kalimatanica clade is suggested to have been caused by
changes of land area configuration in Southeast Asia as a result of global sea level changes during
late Oligocene to early Miocene.

The Ibaliidae constitute a small family
of parasitic cynipoids comprising nineteen
known species, of which all but one are
restricted in the Northern Hemisphere.
The species belong to the three genera Ei-
leenella Fergusson, Heteribalia Sakagami
and Ibalia Latreille. They parasitize siricid
woodwasps, both in conifers and hard-
woods, and some species of Ibalia have
been used in the biological control of sir-
icid pests in conifer plantations. Ibaliidae
is of interest owing to its near-basal phy-
logenetic position within the superfamily
Cynipoidea (Ronquist 1995). Recently, Liu
& Nordlander (1992, 1994) studied the
North-American species of the Ibaliidae
and presented a review of the world spe-
cies of the family, and Nordlander et al.
(1996) studied their phylogeny and histor-
ical biogeography. In this study, a new
species of Ibalia Latreille is described from

' This paper appeared in Acta Uinzrrsitntis Suieciae
Sihestria 62(1998) but was not intended for perma-
nent scientific record as stated in the Disclaimer pub-
lished on page 6 of that publication. This Disclaimer
satisfies Chapter III, Article 8, section b, of the ICZN.
(Editor)

the tropical rain forests of eastern Kali-
mantan, Borneo, Indonesia. The character
matrix of Nordlander et al. (1996) has been
reanalyzed in order to determine the phy-
logenetic position of the new species in re-
lation to other Ibalia species, and to inves-
tigate whether the topology of the phylo-
genetic tree of the genus would be thus
affected.

Species of Ibalia have previously been
described only from the Northern Hemi-
sphere; the new species represents the first
tropical species of the genus. The only oth-
er tropical ibaliid species, Eileenella cather-
inae, is from New Guinea, and is the sister
species to all other ibaliids (Fig. 2). There-
fore, the phylogenetic position of the new
Ibalia species will provide new evidence
for testing the previous biogeographical
scenario of Nordlander et al. (1996).

MATERIALS AND METHODS

Terminology used in this article follows
that of Ronquist and Nordlander (1989)
and Liu and Nordlander (1994).

Only a single female of the new species
was available. The character-state coding
of the new species was made in compari-

150

Journal of Hymenoptera Research

Table 1. Character states tor Ibcjliii knlimatitiDiicn. Characters and character states are coded as in Nordlander
et al. (1996), with the following addition of character state: Character 55. Sculpture of mesopleural speculum:
(2) distinctly vertically costate with secondary irregular foveolate sculpture. The table is supplementary to the
previously published matrix including all the other species.

character

/. kalimantanka

11???

11100

10100

01-00

0-01 1

00-10

00120

010-1

son with representatives of the genus Het-
erihalia Sakagami and each of the two sub-
genera of Ibalia, as well as of the out-
groups Liopteridae and Eileenella Fergus-
son.

The characters and character coding
were the same as in the previous cladistic
analysis of the Ibaliidae (Nordlander et al.
1996). For characters 31, 34, 35, and 69, a
polymorphic condition was coded as a
separate, intermediate state and each step
was given the weight 0.5, so that a change
between non-polymorphic states would
count as one step instead of two. Of the
multi-state characters, characters 6, 45, 52,
63, 66, 67, and 80 were unordered; the oth-
ers were ordered in the sequence 012. The
only change in relation to the previous
study was that an autapomorphy for /. kal-
immitauica required additional state for
character 55 (see Table 1).

Methods used for phylogenetic and bio-
geographical analyses were in general as
described in Nordlander et al. (1996), with
some minor modifications. As in Nordlan-
der et al. (1996), PAUP 3.1 were used for

Fig. 1. Ilnilin kalimantaiucii, new species.

phylogenetic analysis, but less extensive
options were adopted when calculating
support indexes for the branches. Options
for bootstrapping included heuristic
search, random addition sequence, 1000
replications, and for each replication tree
search options are simple addition se-
quence and tree bisection reconnection
(TBR) swapping. The decay index (or Bre-
mer support), the number of extra steps
needed to break up the group, were ob-
tained using branch and bound search
and simple addition sequence. For biogeo-
graphical analysis, the previously defined
distribution area Eastern Palaearctic -I-
North-east Oriental was extended to in-
clude oceanic Southeast Asia, and defined
as Eastern Palaearctic -I- Eastern Oriental.
DIVA 1.1 was used in the present study for
historical reconstruction (Ronquist 1996,
1997). It is basically the same as the earlier
version (DIVA 1.0) used by Nordlander et
al. (1996), but with some performance im-
provements (Ronquist 1996) that should
not affect comparison of the results.

Ibalia (Tremibalia) kalimantanica Liu,
new species

(Fig. 1)

Female.— Body length 10.0 mm. Colorn-
tion: Head yellow except eyes, upper face,
and a narrow longitudinal median strip
through lower face, which are black. Fla-
gellomeres 4—10 of female antenna white
to pale, contrasting with the remaining
darker antennal segments. Pronotum pale
yellow with anterior plate of pronotum
medially, dorsal pronotal area entirely,
and lateral pronotal area posteriorly black.

Volume 7, Number 2, 1998
Table 1. Extended.

151

41

4h

Tl

Sh

hi

66

71

7h

SI

0-012

01111

00102

10100

0-011

11001

?100?

--

Mesothorax mainly black, a small area yellow with dorsal parts of coxa, trochan-
posteriorly on mesoscutum, mesopleural ter, and femur yellow to brown. Metaso-
triangle entirely, and a broad transverse ma pale yellow with three narrow, trans-
band across middle of scutellum yellow, verse to oblique, dark brown strips (Fig 1).
Metathorax and propodeum black. Legs Head. — Vertex rather weakly longitudi-

X

100/10

95/2

0

2-5

1

X

99/10

10-23
97/5

86/2

1

2-6

73/1

1

1-2

1

2-2.5

5-9.5
64/2

93/4.5

4-4.5

5-6

76/2

3

2-4.5

81/3

1-2

0-/

78/2

3-5

2-4

10-25

2

7-12

55/2

2-3.5

5-6

52/2

3.5

1

2-13.5

2-7,5

55/1 .5

1-2

1.5-4

48/1

1-2

^-o

58/1

2

1-3

Liopteridae

catherinae

Eileenella

confluens t

nishijimai

divergens

Heteribalia

subtilis

aureopilosa ,

jakowlewi

anceps

ornata

Ibalia

mirabilis

(Tremibalia)

hunanica

japonica ^

aprilina

rufipes

montana

kirki

Ibalia
(ibalia)

anzonica

ruficollis

leucospoides

Fig. 2. Shortest tree of interspecific relationships in the Ibaliidae according to the previous analysis by Nord-
lander et al. (1996). Figures above and below branches as in Fig. 3.

152

Journal of Hymenoptera Research

nally carinate and with rather dense, ad-
pressed pubescence. Upper face complete-
ly rugose; antennal scrobes indistinctly de-
limited by a lateral carina and not distinct-
ly depressed. Gena largely glabrate with
shallow foveae, postero-ventrally distinct-
ly costate. Eye length about 3.1 times
length of malar space.

Antenna. — Female with 11 flagellomeres;
2nd flagellomere distinctly longer than 1st
(F2/F1 = 4/3).

Mesoso7na. — Pronotal crest without me-
dial incision. Pronotum costate almost en-
tirely, covered with dense, adpressed pu-
bescence. Propleuron protruding strongly
ventrally. Scutellar foveae rugose, separat-
ed by median carina. Posterior processes
of scutellum raised weakly posteriorly.
Distance between outer sides of posterior
scutellar processes about 0.7 times maxi-
mum width of scutellum. Femoral groove
of mesopleuron almost smooth with faint
longitudinal carination ventrally; specu-
lum vertically striate with secondary ir-
regular foveolate sculpture. Metepister-
num with vertical costulae. Anterolateral
propodeal process distinct; posterior pro-
podeal process low; lateral propodeal ca-
rina not elevated medially.

Wings. — Forewing subhyaline with
wide dark strip along outer margin and
distinct, narrow, dark band between

Rs-I-M and Cul (behind submarginal cell)
along outer side of M. Areolet present and
very small. Hindwing faintly fuscous
along outer margin; with three hamuli.

Legs. — Anterior lateral crest of metacoxa
rounded and low. Anterior mesotibial
spur present. Anterior apical process of
2nd metatarsomere reaching to middle of
4th tarsomere.

Metasotna. — Metasoma as long as head
and mesosoma combined. Tergum 8 with
sparse hairs.

Male. — Unknown.

B iology. — Unknown .

Distribution. — Indonesia: E. Kalimantan,
Borneo.

Type material. — Holotype, 9, INDONE-
SIA: Eastern Kalimantan, Kayan-Menta-
rang Natural Reserve, WWF Station, Low-
land Dipterocarpus Forest (2°52'N, 115°
49'E), Malaise Trap head, iii.1993 (D.C.
Darling and U. Rosichon) (Museum Zool-
ogi Bogor, Indonesia).

For a comprehensive comparison with
other species of the family Ibaliidae, the
character matrix of Nordlander et al.
(1996) should be consulted. For identifi-
cation, /. kali7nnnta)iicn may be keyed out
by introducing an additional couplet 3a
following the second item of couplet 2 in
Liu & Nordlander's (1994) key to the
world species of Ibalia:

3a. Female antenna contrastingly colored with flagellomeres 4-10 white and the remaining
segments dark. Upper face lacks regular sculpture. Speculum vertically striate. Posterior
processes of scutellum posteriorly only weakly raised . . . /. kalimatttanica Liu, new species

- Female antenna evenly light yellow to dark brown throughout or becoming darker to-
ward apex, but never with contrasting colors. Upper face longitudinally or transversely
carinate at least in antennal scrobes. Speculum finely striolate longitudinally. Posterior
processes of scutellum raised distinctly posteriorly 3

REVISED PHYLOGENY AND
HISTORICAL BIOGEOGRAPHY OF

IBALIIDAE
The sum of minimum and maximum
possible lengths over all characters in the
data matrix of Nordlander et al. (1996, Ta-

ble 1) with the data of /. kalimantanica add-
ed (Table 1) was 99 and 414 respectively.
Parsimony analysis using the branch-and-
bound algorithm of PAUP resulted in one
optimal tree of length 149 (CI - 0.67, RI
= 0.84). Compared with Nordlander et

Volume 7, Number 2, 1998

153

X

95/2

G

2-5

1 00/1 0

1

X

10-20

86/2

73/1

1

.-0|

1

1-2

1

94/5

90/4

4

99/10

5-7

4-5

4-6

57/2

2-3

75/4

3-5

53/2

6

2-4

57/2

39/1

2-3

10-22

4-b

1-9

1-3

3-4

8-10

54/2

2-4

5-6

61/3

4

11-13

3-6

57/1,5

1-2

2-4

47/1

55/1

1-2

O Q

2

1-3

Liopteridae

catherinae

confluens

nishijimai

divergens

subtilis

aureopilosa ^

jakowlewi

ornata

anceps

kalimantanica

hunanica

japonica

mirabilis

aprilina

rufipes

montana

kirki

arizonica

ruficollis

leucospoides

Eileenella

Heteribalia

Ibalia
(Tremibalia)

Ibalia
(Ibalia)

Fig. 3. Shortest tree of interspecific relationships in the Ibaliidae, obtained with the hranch-and-bound al-
gorithm of PAUP (length = 149, CI = 0.67, RI = 0.84). Shown above each branch in the tree is the support
for the corresponding clade, measured as the percentage with which the clade appeared among the shortest
trees in 1000 bootstrap replications of the analysis, followed by the decay index (or Bremer support). Below
each branch are the minimum and maximum number of character changes along that branch. Clades marked
'x' were constrained to be monophyletic according to results of Ronquist (1995).

al.'s previous optimal tree (length = 141,
CI = 0.67, RI = 0.85), the new tree is al-
most identical in terms of fitness. Com-
pared with the earlier phylogeny of Ibali-
idae (Fig. 2) as presented by Nordlander
et al. (1996), the topology of the phyloge-
netic tree remained unchanged with the
addition of /. kalimantanica, except for two
local changes within the Tremibalia clade.

The first change concerns the relative re-
lationship within the clade (/. mirabilis
Yasumatsu, /. japonica Matsumura, /. hu-
nanica Liu & Nordlander), and the other
concerns the relative positions of /. anceps
Say and /. ornata Belizin (Figs. 3, 4). The
monophyly of Ibalia (Ibalia) became better
supported after the inclusion of /. kaliman-
tanica in the analysis, the bootstrap value

154

Journal of Hymenoptera Research

I I I I I

1 3 8 12 71

Legend:

— 0^1

ESS 2^1

^ l-»0
^ 0-»2

I I H — jakowlewi

20 35 45 52

1+1-1-

49 64 65 66

I I I I — ornata

57 60 63 73

+++

25 53 72

35 69

4H-

24 70

anceps

1 [] Q I I g — kalimantanica

2 12 32 40 47 55

++++

30 4143 57

-S — hunanica

31 66

20

mirabilis

19 25 72

japonica

27

Fig. 4. The clade of Ibalia (Treniibnlia) of the shortest tree with all character changes that could be mapped
unambiguously.

increased from 64% to 72% and Bremer
support from 2 to 4.

As a result of the changes in tree topol-
ogy, the previously suggested Eocene-Oli-
gocene separation of 7. anceps (ca 33-34
MY A) in the subgenus Tremibalia is now
one node further from the base of the tree
(Nordlander et al. 1996). The origin of the
Ibaliidae, based on node /branch distance
calculation, is now estimated to be about
160 MY A, a negligible difference from the
previous estimate of 150 MYA with regard
to potential calculation errors. The esti-
mated time for the origin of the Ibaliidae
is still the Late Jurassic.

An exact search of the updated distri-
bution matrix using DIVA 1.1 resulted in
one single reconstruction of the distribu-
tion history requiring nine dispersals (Fig.
5). The present reconstruction postulates a
center of origin for Heteribalia and Ibalia in
the Eastern Palaearctic — North Oriental
region, and is in complete accordance with
that preferred by Nordlander et al. (1996,
Fig. 7).

The separation of the /. kaliniaiitaiiica
clade probably resulted from a dispersal
within the Oriental as early as in the end
of Oligocene (29-24 MYA) and its subse-
quent isolation from its sister clade. The

Volume 7, Number 2, 1998

155

Legend:

A = West Nearctic
B = East Nearctic
C ~ West Polaearctic
D — East Polaearctic

and East Oriental
E = New Guinea

E. catherinae
H, subtilis

H. aureopilosa
H, divergens
H, nishijimai
H. confluens

T.) mirabilis
T.) japonica
T.) hunanica
T.) kalimantanica
J.) anceps
T.) ornata
[T.) jakowlewi

.) aprilina

.) rufipes

.) montana

.) Wrki

.) arizonica

.) ruficollis

,) leucospoides

Fig. 5. Reconstruction of ancestral distribution of Ibaliidae using DIVA 1.1 resulted in one exact solution
that requires nine dispersals. Dispersal events are indicated on the branches, and implied between-area vi-
cariance events are indicated by hyphens in the ancestral distributions.

land area configuration of the Southeast mained high from Palaeocene through

Asia has varied greatly since late Oligo- Oligocene (65-30 MYA). By late Oligocene

cene as a result of global sea level changes (29 MYA) there occurred a spectacular fall

(Heaney 1991). The global sea levels re- in sea level to about 250 m below the pres-

156

Journal of Hymenoptera Research

ent level, and it then recovered to present
level by end of Oligocene (24 MYA). From
then onwards the sea levels progressively
rose, with minor drops, to about 220 m
above the present level in the middle Mio-
cene (13 MYA). This was followed by sev-
eral cycles of fluctuating sea levels (Hutch-
ison 1989). During times of low sea levels,
Sumatra, Java and Borneo were part of a
peninsula projecting south from continen-
tal Asia (often referred to as Sundaland)
(Morley & Flenley 1987, Heaney 1991), fa-
cilitating the dispersal of the stem species
of /. kalimatanica and its sister species from
the continental Asia to Borneo, and the
continuous ancestral distribution was sub-
sequently split when sea level rose again.
Although, any of these sea level cycles
could have been responsible for the spe-
ciation of 7. kalimantanica, the many auta-
pomorphies of /. kalimantanica and the
rather many synapomorphies for its sister
group indicates that the event probably
occurred rather early. Using the same dat-
ing method as in Nordlander et al. (1996),
the vicariant event separating I. kaliman-
tanica from its sister species was estimated
as having occurred at about 21 MYA. This
is in general accordance with the sea level
recovery since late Oligocene from the late
Oligocene dramatic drop, which could
have facilitated the dispersal of the ances-
tral species.

ACKNOWLEDGMENTS

I thank Goran Nordlander and Fredrik Ronquist
for valuable comments, Chris Darling for presenting
me with this pleasant specimen, and Rune Axelsson
for photographic assistance. The study was support-

ed by a scholarship from the Oscar and Lili Lamm's
Memorial Foundation and by a grant from the Swed-
ish Natural Science Research Council to F. Ronquist
and G. Nordlander.

LITERATURE CITED

Heaney, L. 1991. A synopsis of climatic and vegeta-
tional change in Southeast Asia. In Schneider, S.
H (ed.) Cliniiitic dw)i\;c 19(2-2), special i^iuc: Tropj-
iail foresti ami climate. Kluwer Academic Publish-
ers, Dordrecht, pp 53-61.

Hutchison, C. S. 1989. Geological evolution of South-
East Asia. Oxford Monographs on Geology and Geo-
pliisics, No.l3. Clarendon Press, Oxford.

Liu, Z. and G. Nordlander. 1992. Ibaliid parasitoids
of siricid woodwasps in North America: two
new Ibalia species and a key to species (Hyme-
noptera: Cynipoidea). Proceedings of the entomo-
logical Societi/ of Washington 94: 500^507.

Liu, Z. and G. Nordlander. 1994. Review of the family
Ibaliidae (Hymenoptera: Cynipoidea) with keys
to genera and species of the World. Entomologica
Seandinavica 25: 377-392.

Morley, R. J. and J. R. Flenley. 1987. Late Cainozoic
vegetafional and environmental changes in the
Malay Archipelago. In Whitmore, T. C. (ed.) Bio-
geographical evolution of the Malay Archipichigo. Ox-
ford Monographs on Geology and Geophisics,
No. 4. Clarendon Press, Oxford, pp 50-59.

Nordlander, G., Z. Liu, and F. Ronquist. 1996. Phy-
logeny and historical biogeography of the cyni-
poid wasp family Ibaliidae (Hymenoptera). Sys-
tematic Entomology 21: 151-166.

Ronquist, F. 1995. Phylogeny and early evolution of
the Cynipoidea (Hymenoptera). Systematic Ento-
mology 20: 309-335.

Ronquist, F. 1996. DIVA version 1.1 computer program
and manual available by anonymous FTP from
Uppsala University (ftp.uu.se or ftp.systbot.uu.se).

Ronquist, F. 1997. Dispersal — vicariance analysis: a
new approach to the quantification of the histor-
ical biogeography. Systematic Biology 46: 195-203.

Ronquist, F. and G. Nordlander. 1989. Skeletal mor-
phology of an archaic cvnipoid, Ibalia riifipes (Hy-
menoptera: Ibaliidae). Entomologica Scandinaz'ica
Supplements 33: 1-60.

J. HYM. RES.
Vol. 7(2), 1998, pp. 157-164

Taxonomy, Mature Larva, and Observations on the Biology of
Gnamptopelta obsidianator (Brulle) (Hymenoptera: Ichneumonidae,

Ichneumoninae)

Karen R. Sime and David B. Wahl

(KRS) Department of Entomology and Section of Ecology and Systematics, Cornell University,

Ithaca, New York 14853, USA, (DBW) American Entomological Institute, 3005 S.W. 56th

Avenue, Gainesville, Florida 32608, USA

Ahstrnct. — The two nominal species in Gnamptopelta, C. obsidiatiator (Brulle) and G. austrina
(Cresson), do not merit separate species or subspecies status, and the latter is synonymized under
the former; the genus is hence monotypic. Specimens reared from Atnphion floridensis (B.P. Clark)
(Lepidoptera: Sphingidae), probably attacked in captivity, represent the first verified host record
for Gnamptopelta. A description of the mature larval exuviae is provided. Observations of the
wasps in the field and in captivity suggest that G. obsidianator directs its host-searching to grape-
vines (Vitis spp.), and that its host range may not include all grapeleaf-feeding sphingids.

Gnamptopelta obsidianator (Brulle) is one
of the largest ichneumonids in eastern
North America and one of the most strik-
ing, with its black body, yellow antennae,
and, in some southern populations, exten-
sive reddish coloring of the head and an-
terior mesosoma. While it is commonly
collected on the wing and is well repre-
sented in collections, no credible hosts
have previously been reported. The genus
is currently placed in the Trogini, subtribe
Callajoppina; nearly all recorded callajop-
pine hosts are Sphingidae (Heinrich 1962).
Heinrich (1977) noted that G. obsidianator
frequents grapevines (Vitaceae: Vitis spp.),
which are the food plants for at least 10
sphingids in North America (Forbes 1948;
Hodges 1971). Thus, guided by taxonomic
inference and a few incidental observa-
tions, we set out to identify the hosts of G.
obsidianator by rearing Vifis-feeding sphin-
gids.

We also decided to use this opportunity
to clarify the status of species within
Gnamptopelta Hopper, in which two spe-
cies, obsidianator (Brulle) and austrina
(Cresson) were originally included. Sub-

sequent authors (Townes and Townes
1951; Heinrich 1962, 1977; Carlson 1979)
have considered austrina to be a subspe-
cies of obsidianator, although Heinrich
(1977) hedged that the subspecies could be
"ecologically differentiated species para-
sitizing 2 different hosts living on the
same plant."

The Gnamptopelta specimens examined
in this study are in the American Ento-
mological Institute (Gainesville, Florida:
AEIC) and the Florida State Collection of
Arthropods (Gainesville, FL: FSCA).

TAXONOMY

Heinrich (1962) placed Gnamptopelta in
the tribe Trogini, subtribe Callajoppina,
which also contains the genera Afrotrogus,
Callajoppa, Catadelphus, Conocalama, Dimae-
tha, Holojoppa, Pepsijoppa, Stirojoppa, Tme-
togaster, Tricyphus, and Yeppoona (Ward
and Gauld 1987; Yu and Horstman 1997).
All reliable host records indicate that these
genera parasitize only Sphingidae.

Hopper (1939) erected Gnamptopelta for
two North American species, Trogus obsi-
dianator Brulle and Trogus austrinus Cres-

158

Journal of Hymenoptera Research

son. These were distinguished on the basis
of color: G. austrina, found in South Car-
ohna, Georgia, and Florida, had the head
and anterior mesosoma reddish while G.
obsidianator, found in the remaining part of
eastern North America west to the Rocky
Mountains, had the entire body black ex-
cept for the yellowish-white paraocular
area. With the exception of Townes (1944),
later authors (Townes and Townes 1951;
Heinrich 1962, 1977; Carlson 1979) have
treated austrina as a subspecies of obsidi-
anator. Heinrich (1962) could not decide
whether the taxa represented two species
or a single species with two color forms.
After studying the Florida ichneumonine
fauna (Heinrich 1977), he concluded that
for Gnamptopelta: 1) "austrina . . . occupies
the peninsula of Florida except its most
northern part, and that the uniformly
black obsidianator occupies the entire east-
ern territory of continental North Ameri-
ca, including the base of Florida"; 2)
"Very sporadically, however, obsidianator
also occurs in the territory of austrina, and
likewise, sporadically, austrina is found in
the southern parts of the territory of obsi-
dianator"; 3) ". . .it also must be mentioned
that "intergrades" between the 2 forms
have never been found" {op. cit., p. 285).

Our examination of over 550 Gnampto-
pelta specimens reveals that, contrary to
Heinrich's assertions, intermediates do oc-
cur and all are found in Florida. Most of
these individuals are found north of
Ocala, in the northern third of the state
(above 29°N). Black and intermediate
specimens are found throughout the flight
period (mid-February through mid-No-
vember); there is no sex bias in the color
forms. An interesting series of eight males
was collected at the American Entomolog-
ical Institute in March and April of 1986
[AEIC]. Every variation is present, from
uniformly black with no reddish markings
to the extreme of an entirely reddish head,
reddish anterior mesosoma, and reddish
postpetiole. This series alone refutes the
notion of two separate entities. We there-

fore place austrina as a junior synonym of
obsidianator (NEW SYNONYM), noting a
particularly apt remark that summarizes
the past Gnamptopelta literature: ". . .too
much time has been wasted on trying to
assemble data, much of it illusory, to
maintain already proposed names for sub-
species" (Franclemont 1973).

All records to date indicate that G. ob-
sidianator does not occur west of the front
range of the Rocky Mountains. We have,
however, seen one specimen from Califor-
nia ("Merced Co.; Los Banos National
Wildlife Refuge, 4 mi. N. Los Banos; 12-
vi-1981; D. Jamieson"; FSCA). It is uni-
formly black and identical to specimens
from the northeastern part of the range.
Queries of the major California collections
have not revealed any more specimens
from the West Coast. At this time, we con-
sider the specimen to be mislabeled or
perhaps the result of an import to the area.

MATURE LARVA OF GNAMPTOPELTA
OBSIDIANATOR

Materials and Terminology. — The termi-
nology of the cephalic sclerites of the ma-
ture larva is that of Gillespie and Finlay-
son (1983) and Short (1978), with modifi-
cations by Wahl (1990). The epistomal su-
ture is the lateral groove between the
anterior tentorial pits (Matsuda 1965). It is
present as a distinct depression in mature
ichneumonid larvae. Although the area is
usually unsclerotized, a continuous scler-
otized band extends across it in the An-
omaloninae, Ichneumoninae, Metopiinae,
and Pimplini (and isolated taxa in other
subfamilies). Previous ichneumonid work-
ers have referred to this area as the "ep-
istoma," including in it the portion of the
pleurostoma above the superior pleuro-
stomal process. This use of "epistoma"
should be eschewed. We suggest the fol-
lowing descriptive terms: a) epistomal su-
ture unsclerotized; b) epistomal suture
partially unsclerotized, medially incom-
plete (as in Cryptus albitarsis (Cresson);
Short 1978: fig. 238); c) epistomal suture

Volume 7, Number 2, 1998

159

Fig. 1. Mature larva of Cnainptopclta ohsiduviator: (a) cephalic sclerites, (b) prothoracic spiracle. The scale lines
represent 0.1 mm.

completely sclerotized forming an episto-
mal hand.

Methods of larval preparation are those
of Wahl (1989). Wahl's notaHon for larval
preparations follows the museum acro-
nym. It consists of his initials, the day,
month, year, and a letter designating the
individual preparation.

Description. — Epistomal band present,
with 14 pores. Labral sclerite, hypostomal
spur, stipital sclerite, labial sclerite, and
prelabial sclerite, absent. Clypeolabral
plates present and triangular in outline,
not connected to epistomal band, and not
joined or touching medially. Pleurostoma
and hypostoma strongly sclerotized. Ac-
cessory pleurostomal area moderately
sclerotized. Hypostoma straight, posterior
end not elongate or upcurved. Accessory
reticulate sclerotization prominent. Max-
illary palpus with four sensilla and an ad-
jacent sclerotized seta; labial palpus with

five sensilla and four adjacent setae. Man-
dible strongly and uniformly sclerotized;
blade about 0.4 X as long as mandible,
without denticles. Antenna disc-like, rim
moderately sclerotized; central papillus
absent. Parietal band present and weakly
sclerotized. Prothoracic spiracle as in fig.
lb: atrium with width about 0.7x as long
as closing apparatus and interior with nu-
merous long heavy spines; atrial opening
with conspicuous rim; closing apparatus
adjacent to atrium. Skin smooth, with
widely scattered setae and without spines.
Fig. 1 illustrates the cephalic sclerites
and prothoracic spiracles of the mature
larva; it is a composite of two individuals
collected in the Ocala National Forest
(USA: Florida; Marion Co.; Lake Delancy
campground area, 5.5 miles N-NW Salt
Springs, 29° 25'N, 81° 47' W; K.R. Sime &
D.B. Wahl; emerged 31 May 1997 [AEIC:
DBW l.vi.l997a] and emerged 19 June

160

Journal of Hymenoptera Research

1997 [AEIC: DBW 20.vi.l997a], both from
Aviphioii floridensis). The characteristic ab-
sence of the labial sclerite in ichneumoni-
nes resulted in distortion of the labial re-
gion in both preparations. Consequently,
placement of the salivary orifice, palpi,
and associated setae is only approximate.
Discussion. — Short (1978) and Gillespie
and Finlayson (1983) provide keys to final-
instar larvae of ichneumonine tribes and
genera. Townes and Heinrich differed re-
garding placement of the Callajoppina.
Short follows Townes (Townes et al. 1961)
in placing them in the Heresiarchini ("Ich-
neumonini" of Short); Finlayson accepts
Heirurich's placement of these genera as
the subtribe Callajoppina in the Trogini
(her "Trogusini"). Both sets of keys will
take Gnamptopelta to the correct group of
genera. Further use of Short will run
Gnamptopelta to Callajoppa; Gillespie and
Finlayson will key it to Conocalama. Mod-
ification of these keys will not, in our
opinion, be useful. Both are inadequate
samples of the world ichneumonine fau-
na: of approximately 370 genera world-
wide (Wahl 1993), Gillespie and Finlayson
treat 25 and Short 53. Any attempted cor-
rections of the generic keys will risk defeat
when the next previously undescribed lar-
va is found.

BIOLOGY OF GNAMPTOPELTA
OBSIDIANATOR

Observations. — Field work was conduct-
ed in the Ocala National Forest, in north-
central Florida (29° 25'N, 81° 47' W),
where both grape and G. obsidianator are
abundant in the spring (March to May).
The study site is part of an extensive stand
of longleaf pine (Pinaceae: Pinus paiustris
Miller), with an open understory domi-
nated by wiregrass (Gramineae: Aristida
striata Micheaux) and scattered low
shrubs, chiefly scrubby oaks (Fagaceae:
Quercus marilandica Muenchhausen and Q.
laevis Walter), and pawpaw (Annonaceae:
Asimina speciosa Nash and A. obovata
Willdenow). Two species of grape are

Table 1. Summary of field observations of female
obsiiiianator. Identification of plants landed upon with
searching times, recorded when possible, in paren-
theses. Before 8 April we had not confirmed the iden-
tities of Viti:^ species.

Wasp No,

Searching Sequence

1 (23 March) Vitif (2-3s); V;f/s (2-3s); Vitis (2-3s)

2 (23 March) Quercus (<ls); Quercus (<ls); Vitis

(25s); Vitis (lost)

3 (23 March) Vitis (2-3s); Vitis (2-3s); Vitis (2-3s)

4 (23 March) Vitis (2-3s): Vitis (2-3s); Quercus (2-3s)

5 (23 March) Vitis (2-3); Vitis (2-3s); Quercus (2-3s)

6 (8 April) V. rotundifolia (>20s); V. aestivalis (10s)

7 (8 April) V. aestivalis (captured after 2-3s)

8 (17 April) V. rotundifolia (>60s)

9 (24 April) V. aestivalis (captured after 2-3s)

found here, both extremely common and
often covering large expanses: Vitis aesti-
valis Micheaux and V. rotundifolia Mi-
cheaux. In 1997, most grape plants of both
species were fully leafed out by the mid-
dle of March, and both continued to put
out new leaves throughout the spring. A
few G. obsidianator were seen as early as
10 March; the peak of abundance ap-
peared to be during the last week of the
month, when for a few days 12-15 wasps
were seen each day. From then until the
middle of May, when observations ceased,
we usually saw 1-3 individuals in the
course of 4-5 hours spent almost every
day in the field.

Males fly rapidly, 2-3 meters above the
ground, land rarely, and follow wide cir-
cuits in patches of forest abounding in
grapevines. They were perhaps patrolling
areas in which females were likely to
eclose or to enter in the course of search-
ing for hosts, although we witnessed no
encounters between the sexes.

The females as well as the males are as-
sociated with grape at this field site. The
females also fly quite fast, but land often.
Table 1 summarizes our observations of
searching behavior. Females appear to fly
deliberately from one grape patch to the
next, less often landing on other common
low shrubs, which were usually surround-

Volume 7, Number 2, 1998

161

ed by if not overlaid v^ith grapevines. The
wasps land on the upper surface of the
leaf and tap it several times with the ven-
tral surfaces of the antennae; usually, the
inspection lasted just a few seconds, and
the wasp then proceeded to the next plant,
but in some cases the wasps stayed longer
and conducted a thorough search of near-
by leaves and stems. We suspect that the
length of the search might be related to the
presence of feeding damage or other trac-
es left by host larvae (Vinson 1984). How-
ever, we observed no encounters with
hosts in the field: the leaves searched, in-
cluding those examined for just a few sec-
onds, often bore herbivore damage, but
we never found larvae in their vicinity.

Between 31 March and 2 May 1997 we
collected 66 sphingid larvae on Vitis. We
reared them in small groups (3-10 larvae)
in plastic boxes which we supplied with
fresh grape leaves every 1-2 days. Using
Forbes's (1948) key, we were able to iden-
tify the caterpillars to species when they
reached the 4th instar (and confirmed the
identifications when adults emerged). All
were macroglossine sphingids: Darapsa
myron Cramer (33 reared to pupation),
Amphiou floridensis B.P. Clark (= nessus
Cramer) (5 reared), and Euinorpha achemon
Drury (2 reared). Larval mortality was
due to braconid parasitism (Aleiodes iex-
ensis) killing 4th instars (10 victims, all D.
myron) and unknown causes among 2nd-
instar larvae (too small to identify). Of the
D. myron and A. floridensis found during
the first two weeks (about 45 larvae), all
but one were small second and third in-
stars; later collections included a few 4th
and 5th instars. The two £. achemon larvae
were found side-by-side on April 26 as
very nearly mature 5th instars.

All larvae were found on V. aestivalis.
Although we often found compelling
feeding damage on V. rotundifolia, and this
plant was inspected both by us and by
Gnamptopelta females, we found no sphin-
gid larvae on it: if not coincidence, this
could be evidence for an oviposition pref-

erence in the adult sphingids, or for higher
larval survivorship on V. aestivalis. Fur-
thermore, in rearing the larvae, we found
that they preferred to feed on V. aestiimlis
when both species of grape were in their
boxes. This could reflect the habituation of
the larvae to the plant on which they ini-
tially fed (Jermy 1987), or it could repre-
sent an inherently greater palatability of
V. aestivalis to the larvae. As most of the
larvae were D. myron, these observations
may apply only to that species; the num-
bers of the other two species collected
were too small for useful speculation. In
addition, we noted that all larvae fed on
full-sized leaves, never on the youngest
leaves, which might represent an aversion
to the denser pubescence of the youngest
leaves (Southwood 1986) or perhaps to
higher levels of allelochemicals (Feeny
1976).

The larvae of D. myron and A. floridensis
have many habits that make them incon-
spicuous in the field. Whether feeding or
resting, all instars are invariably on the
undersides of the leaves, and they usually
rest along the leaf veins, which the light-
green young instars match particularly
well in color and shape. Often the larvae
rest on undamaged foliage one, two, or
three leaves removed from the feeding
site. The younger larvae feed in a distinc-
tive fashion, by eating symmetrical holes
on either side of the leaf, near the base, a
pattern that blends extremely well with
the deep rounded lobes and hollows of the
leaves of V. aestivalis. Older larvae tend to
feed along the edges of the leaf, eating off
large pieces. Heinrich (1979) noted similar
behaviors in the sphingid Sphecodina ab-
botti Swainson feeding on V. vulpina L. (in
Minnesota) and suggested that this, along
with the habit of staying under the leaves,
conceals the caterpillars from predators
that hunt by eye, particularly birds. We
did not happen to observe any birds in-
specting grapevines, but speculate that
these behaviors may be somewhat effec-
tive against G. obsidianator. As the wasps

162

Journal of Hymenoptera Research

land initially on the top surface of the leaf,
a caterpillar resting underneath may gain
some extra time to fall off without being
noticed, and its removal from the feeding
site while resting might help frustrate a
wasp that was initially attracted (by sight
or by odor) to feeding damage (Vet and
Dicke 1992).

Our observations of G. obsidianator and
sphingid larvae in captivity suggest that
letting go from the leaves is an effective
means by which caterpillars elude wasps.
We placed three field-caught females in a
small glass aquarium (23-cm cube) with 4
larvae (at a time) feeding on grape sprigs.
Some larvae (both D. myron and A. flori-
densis) dropped off their leaves as soon as
wasps set foot on the leaves; others
dropped only after the wasps' antennae or
tarsi touched them. If the wasps tried to
oviposit, the time it took them to bring
their ovipositors into position, after mak-
ing antennal contact, was long enough for
the larvae to drop down and thus con-
found the oviposition attempt. Dropping
appeared to be the main defense of small-
er larvae; fourth- and fifth-instar larvae,
when touched by a wasp, stayed put and
swung their heads about with such vio-
lence as to parry any attempt at oviposi-
tion quite effectively, sometimes hurling
the wasp against the side of the cage. The
wasps, if not thrown, in many instances
had so much difficulty inserting their ovi-
positors into the writhing caterpillars that
they simply gave up.

When exploring the aquarium, the
wasps tapped steadily with the extended
antennae, lightly touching the apical quar-
ter (flattened ventral surface) to the sub-
strate. The wasps showed little interest in
D. myron larvae, never attempting to sting
them even after finding and examining
them. In contrast, A. floridensis elicited
great excitement: upon encountering a
chewed leaf margin or a particle of frass,
the wasps slowed their walking pace, vi-
brating the antennae much more rapidly
and pressing them more firmly against the

surface, with more of the antennal area,
approximately the apical third, making
contact. Upon finding A. floridensis larvae,
the wasps tapped them for 1-2 seconds
with the antennae and then turned around
to sting. We witnessed six apparent ovi-
positions, by two females; it appears that
the eggs may be inserted anywhere into
the host's body apart perhaps from the
head capsule. Attempts at oviposition oc-
curred with third-, fourth-, and fifth-instar
larvae (we did not expose younger lar-
vae).

The G. obsidianator that we reared came
from A. floridensis and resulted from those
larvae exposed as 3rd and 5th instars. In
total, we had six A. floridensis larvae that
grew large enough to be identified: two
went to term and emerged as moths, and
four others were exposed, of which one
died just before and one some time after
pupation and two produced the wasps.
Six of the D. myron larvae (also 3rd, 4th,
and 5th instars) were exposed to the
wasps and continuously observed, but no
ovipositions were attempted with these
larvae and all developed into adult moths.

The sphingids, parasitized and not, pu-
pated by forming loose cocoons amid
leaves and paper towels. The adult wasps
emerged by cutting off the anterior 0.1 of
the host pupa. Such an emergence hole is
apparently characteristic of the Callajop-
pina (Mell and Heinrich 1931), and ap-
pears to be the primitive condition for the
Ichneumoninae (Gillespie and Finlayson
1983). The ichneumonid's rudimentary co-
coon, located in the detached anterior sec-
tion, consists of a cap of dark brown silk
just inside the cut margin.

Conclusions. — Although it was the most
common grape-feeding sphingid at our
field site in spring 1997, our observations
suggest that D. myron is not a host of G.
obsidianator. Of 33 larvae collected on
plants exposed to a considerable popula-
tion of G. obsidianator, no wasps emerged;
the wasps showed no interest in this spe-
cies in captivity (and much interest in A.

Volume 7, Number 2, 1998

163

floridensis under the same conditions). It is
possible that the larvae we collected were
too small for successful oviposition or
were not old enough to have spent much
time exposed to wasps, thus lowering the
probability of parasitism, or that parasit-
ism rates are generally very low; however,
the combined field and captivity observa-
tions argue that D. myrou is at least not a
preferred host.

That A. floridensis is a natural host is
strongly supported by our successful rear-
ing in captivity: a koinobiont larva-pupal
endoparasitoid, G. obsidianaior is not likely
to develop successfully in randomly en-
countered moth larvae because it must
survive the immunological defenses of the
host and also coordinate its development
with the onset of pupation in the host
(Askew and Shaw 1986; Omata 1984). Am-
phion floridensis occurs throughout the
geographical range of G. obsidianator and
is thus potentially the sole host: its range
is described as extending "from Florida,
north to Nova Scotia, west to Calgary, Al-
berta, and south to Kansas" (Hodges
1971), and our queries of various collec-
tions expand this to central Colorado and
southern Texas as well. In Florida, at least,
the flight period of G. obsidianator coin-
cides with the larval stage of A. floridensis
(Kimball 1965). However, we can say
nothing about £. achemon, because we
found only two specimens and did not ex-
pose them to wasps; some 10 other sphin-
gids feeding on grape in eastern North
America (Hodges 1971) also demand in-
vestigation.

Hopper (1939) lists as a host Papilio po-
lyxenes Fabricius (Lepidoptera: Papilioni-
dae), referring to a New York list for
which voucher specimens are unobtaina-
ble. As may be the case for many such iso-
lated records (Shaw 1994), this one is al-
most certainly specious. This swallowtail
is often reared by amateurs and profes-
sionals alike, as the caterpillar is quite
striking and very often found on garden
herbs and common roadside weeds (Um-

belliferae). We have seen scores of speci-
mens of Trogus pennator Fabricius (Ichneu-
moninae) reared from P. polyxenes in col-
lections, but no reared G. obsidianator; a
life-history study of P. polyxenes in central
New York State, where G. obsidianator is
common, found that about 10% of 128 pu-
pae were parasitized by T. pennator, with
a few other parasitoids occurring infre-
quently, but no G. obsidianator (Feeny et al.
1985).

We conclude that G. obsidianator para-
sitizes at least one species of Viffs-feeding
sphingid. The degree of specialization
within this group is unknown, but our ob-
servations of captive wasps suggest that
G. obsidianator does not uniformly attack
available sphingid species and that it
could be restricted, at least locally in Flor-
ida, to A. floridensis. Our results support
Heinrich's (1962) contention that the gen-
era of the Callajoppa group are strictly par-
asitoids of Sphingidae.

ACKNOWLEDGMENTS

We thank Tim Carr, Dan Janzen, Mark Shaw, Bob
Wharton, and two anonymous reviewers for advice
and assistance. Steve Heydon (U.C. Davis), Wojciech
Pulawski (California Academy of Science), Roy Snell-
ing (Los Angeles County Museum), Serguei Triapit-
syn (U.C. Riverside), and Robert Zuparko (U.C.
Berkeley) searched collections for western North
American records of Guainptopelln; Robert Brooks
(University of Kansas), Richard Brown (Mississippi
State University), John Heppner (Florida State Collec-
tion of Arthropods), Eric Quinter (American Museum
of Natural History), Edward Riley (Texas A&M Uni-
versity), and James Whitfield (University of Arkan-
sas) checked A. floridensis distribution records. We are
indebted to James Wiley for the FSCA Gnamptopelta
loan. We thank the staff of the Lake George Ranger
District, Florida, for granting permission to work in
the Ocala National Forest. This material is based
upon work supported under a National Science
Foundation Graduate Research Fellowship (KRS) and
funded by NSF grant IBN-9600094 to KRS and Dr. P.
P. Feeny.

LITERATURE CITED

Askew, R. R. and Shaw, M. R. 1986. Parasitoid com-
munities: their size, structure, and development. In-
sect Parasitoids (eds J. Waage & D. J. Greathead), pp.
225-264. Academic Press, London.

164

Journal of Hymenoptera Research

Carlson, R. W. 1979. Family Ichneumonidae. Catalog of
Hymenoptera in America north of Mexico, vol. 1 (eds
K. V. Krombein, P. D. Hurd, D. R. Smith, & B. D.
Burks). 1198 pp. Smithsonian Institution Press,
Washington, D.C.

Feeny, P. P. 1976. Plant apparency and chemical de-
fense. Recent Aiizninces in Phytochemistry , Vol. 10 (eds
J. W. Wallace & R. L. Mansell), pp. l^W. Plenum
Press, New York.

Feeny, P. P., W. Blau, and P. Kareiva. 1985. Larval
growth and survivorship of the black swallowtail
butterfly in central New York. Ecological Monograplis
55: 167-187.

Forbes, W. T. M. 1948. Lepiiioptera ofNeiv York and neigh-
boring states. Part //. Cornell University Agricultural
Experiment Station, Ithaca, New York.

Frandemont, J. 1973. Tlie moths of America north of Mex-
ico: Fascicle 20.1. Mimallonoidea, Mimallonidae and
Bombycoidea, Apatelodidae, Bombycidae, Lasiocantpidae.
E.W. Classey Ltd. & R.B.D. Publications, Inc., Lon-
don.

Gillespie, D. R. and T. Finlayson. 1983. Classification of
final-instar larvae of the Ichneumoninae (Hyme-
noptera: Ichneumonidae). Memoirs ofttie Entomolog-
ical Society of Canada 124: 1-81.

Heinrich, B. 1979. Foraging strategies of caterpillars: leaf
damage and possible predator avoidance strategies.
Oecologia 42: 325-337.

Heinrich, G. H. 1962. Synopsis of the Nearctic Ichneu-
monidae Stenopneusticae with particular reference
to the northeastern region (Hymenoptera): Part VII:
Synopsis of the Trogini. Canadian Entomologist sup'-
plcment 29: 807-860.

Heinrich, G. H. 1977. Ichneumoninae of Florida and
neighboring states. Arthropods of Florida and neigh-
boring land areas 9: 1-350.

Hodges, R. W. 1971. Tlie moths of America north of Mexico,
Fascicle 11. Sphhigoidea (Hawkmotlis). E.W. Classey
Ltd. & R.B.D. Publications, Inc., London.

Hopper, H. P. 1939. A synophcal revision of the tribe
Trogini Ashmead of the United States and Canada
(Hymenoptera: Ichneumonidae). Transactions of the
American Entomological Society LXV: 307-347.

Jermy, T. 1987. The role of experience in the host selec-
tion of phytophagous insects. Persp'ectiivs in chc-
moreception and behavior (eds R. F. Chapman, E. A.
Bemays, & J. G. Stoffolano, Jr.), pp. 143-157. Spring-
er-Verlag, New York.

Kimball, C. P. 1965. The Lepidoptera of Honda: an an-
notated checklist. Arthropods of Florida and nciglibor-
ing land areas 1:1-363.

Matsuda, R. 1965. Morphology and evolution of the in-
sect head. Memoirs of tlie American Entomological In-
stitute 4:1-334.

Mell, R. and G. Heinrich. 1931. Zur Biologie and Sys-
tematik der siidchinesischen Ichneumoninae Ashm.

(Fam. Ichneumonidae Hym.). Zeitschrift fiir ange-
wandte Entomologie 18: 371^03.

Omata, K. 1984. Influence of the physiological condition
of the host, Papilio xuthus L. (Lepidoptera: Papili-
onidae) on the development of the parasite, Trogus
mactator Tosquinet (Hymenoptera: Ichneumoni-
dae). Applied Entomology and Zoology 19: 430-^35.

Shaw, M. R. 1994. Parasitoid host ranges. Parasitoid com-
munity ecologi/ (eds B. A. Hawkins and W. Shee-
han), pp. 111-144. Oxford University Press, Oxford.

Short, J. R. T. 1978. The final larval instars of the Ich-
neumonidae. Memoirs of the American Entomological
Institute 25: 1-508.

Southwood, T. R. E. 1986. Plant surfaces and insects —
an overview. Insects and the plant surface (eds B. Ju-
niper & T. R. E. Southwood), pp. 1-22. Edward Ar-
nold Ltd., London.

Townes, H. 1944. A catalogue and reclassification of the
Nearctic Ichneumonidae (Hymenoptera). Part I.
The subfamilies Ichneumoninae, Tryphoninae,
Cryptinae, Paeogeninae and Lissonotinae. Memoirs
of the American Entomological Society 11(1): 1^77.

Townes, H., and M. Townes. 1951. Family Ichneumon-
idae. Hymenoptera of America north of Mexico— syn-
optic catalog (eds C. F. W. Muesebeck, K. V. Krom-
bein, & H. K. Townes). Agriculture Monograph 2: 1-
1420.

Townes, H., M. Townes, and V. K. Gupta. 1961. A Cat-
alog and Reclassification of the Indo-Australian Ich-
neumonidae: Tribe Trogini. Memoirs of tlie American
Entomological Institute 1: 1-522.

Vet, L. E. M., and M. Dicke. 1992. Ecology of infodi-
emical use by natural enemies in a tritrophic con-
text. Annual Rei'ieios of Entomolog}/ 37: 141-172.

Vinson, S. B. 1984. How parasitoids locate their hosts: a
case of insect espionage. Insect Communication (ed.
T. Lewis), pp. 325-348. Royal Entomological Sod-
ety, London.

Wahl, D. B. 1989. Further notes on preparation of exu-
viae of parasitic Hymenoptera. Entomological Neics
100: 181-182.

Wahl, D. B. 1990. A review of the mature larvae of Di-
plazontinae, with notes on larvae of Acaenitinae
and Orthocentrinae and proposal of two new sub-
faniilies (Insects: Hymenoptera, Ichneumonidae).
Journal of Natural History 24:27-52.

Wahl, D. B. 1993. Family Ichneumonidae. Hymenoptera
of the World: an identification guide to families (eds H.
Goulet & J. T. Huber), pp. 395-442. Canada Com-
munications Group, Ottawa.

Ward, S. and Gauld, 1. 1987. The callajoppine parasitoids
of sphingids in Central America (Hvmenoptera:
Ichneumonidae). Systematic Entoniologif 12: 503-508.

Yu, D. S. and Horstmann, K. 1997. A catalogue of world
Ichneumonidae (Hymenoptera). Memoirs of tin-
American Entomological Institute 58 (1-2): 1-1558.

J. HYM. RES.
Vol. 7(2), 1998, pp. 165-177

The Nesting Behavior and Dynamics of Bicyrtes angulata (F. Smith)

with a Comparison to other Species in the Genus

(Hymenoptera: Sphecidae)

RoGERio Parentoni Martins*, Lourdes Aragao Scares, and Douglas Yanega

Laboratorio de Ecologia e Comportamento de Insetos, Departamento de Biologia Geral,

ICB-UFMG, Cx. Postal 486, 30.161-970-Belo Horizonte-MG-Brazil;

*E-mail: Wasp@dedalus.lcc.ufmg.br

Abstract. — Data are presented on the nesting behavior and dynamics of a population of Bicyrtes
angulata (F. Smith) found on the campus of the Universidade Federal de Minas Gerais, Belo
Horizonte, Minas Gerais, Brazil. Many aspects of the biology of this population of B. angulata are
similar to all (or the majority) of the 8 other species in the genus that have been studied in some
detail. Other features were atypical in comparison, such as: (1) nesting in sandy soil far from
water (2) constructing relatively short burrows (less than 10 cm on average) (3) taking more than
two hours to dig a nest. New information is presented on the level of nest failure (roughly a third
of all nests initiated are not completed), mortality (roughly 90% of all completed nests fail to
produce adult wasps), and variability in egg to adult developmental time (44 to 375 days) due to
bimodal prepupal dormancy. Factors affecting mortality are discussed, the most important being
termites and ants. The number of nests made per female varied slightly over the year, this ratio
being lowest in the dry season (winter and spring), and was not correlated with the total number
of nesting females, thus suggesting that there is little or no competition for nesting space in the
area. Possible alternative explanations for these patterns are offered, in relation to prey abundance
and female mobility and longevity, and we suggest that the availability of suitable patches for
nesting may potentially be of importance in the nesting dynamics of this species.

Of the 23 described species of Bicyrtes, al species should help address questions

12 are found exclusively in South Ameri- related to patterns of behavioral evolution

ca, 8 in North America and 3 occur in both within the genus.

continents (see Bohart and Menke 1976 for Bicyrtes angulata (F. Smith) occurs in

distributions; Willink, 1947, revised the French Guiana, Paraguay, Argentina, and

South American species). Their biology is Brazil (Bohart and Menke 1976). Records

largely either unknown or poorly known, for B. angulata in Brazil include some

with the exception of the North American states in the North and Northeast (Nasci-

B. quadrifasciata (Say) (see Evans 1966 for mento and Overall 1980) and Sao Paulo

a review). Some addiHonal information is (Martins 1991), but its biology has been

available for B. discisa (Taschenberg), B. unknown until the present, and is com-

simillima (Smith) and B. variegata (Olivier) P^''^'^ ^^^^ ^° ^^^^ has been recorded for

(Genise 1979, 1982), B. cingulata Burmeis- o*^"" ^P^^^^^ '" 'he genus. This includes

ter (Evans and Matthews 1974), and also "°^«^' information on prepupal dormancy,

for the Cuban B. spuwsa (Fabricius) (San- ^"'^ ""f ^' ^"'^ ^^"^^^ °^ "^^* ^^''^''^ ^"'^

chez and Genaro 1992). Few of the details "^o^t^lity.

of nesting and provisioning behavior have STUDY SITE AND METHODS

proven to be uniform across all species We observed an aggregation of B. an-

(see Discussion). Information on addition- gulata (147 marked nests in 1993 and 80 in

166

o Neslsin1993
• Nests in 1994
^■y Bushes
Ji* Grasses
>!|r WaltherLa

Journal of Hymenoptera Research

<?-

)b M. «> ^»>

^.iff ^

4fc W *. ^ dk' U, 4t" ^

*#''=«fi»«^^*'*8.„^^i^bo •r-v^f

Fig. 1. Diagram of the nesting site of Bicyrtes angulata showing nest distributions in 1993 and 1994.

1994) on the Campus of the Federal Uni-
versity of Minas Gerais (UFMG), Belo Ho-
rizonte, Minas Gerais, Brazil, from Janu-
ary 1993 to June 1994. We spent a total of
556 hours making ad libitum and focal in-
dividual observations {sensu Martin and
Bateson 1986), mostly between 0900 and
1200 h when the majority of nesting activ-
ity occurred, and opportunistically at the
beginning and end of some days. The
study site, Prefeitura, is a secondary
growth, relatively undisturbed area, most-
ly covered with grasses, scattered bushes,
and trees (including a few exotic and or-

chard species; see photo in Gaimari and
Martins 1996). Nearby fields of corn, bean
and manihot support a substantial popu-
lahon of Waltheria americana (Sterculi-
aceae), a weed whose flowers are com-
monly visited by B. angulata (Macedo and
Martins 1998).

Most nests of B. angulata were concen-
trated in a 35 m long and 0.65 m wide
strip of a dirt road (approx. 700 m- total)
on a superficially compacted well-drained
sandy soil. Nesting soil was completely
free of vegetation and exposed directly to
the sun (Figs. 1 and 2). We built a tem-

■r'

^

L

.y

Fig. 2. Nesting site of Bicyrtes nii^uhUn showing the plastic-cup emergence traps.

Volume 7, Number 2, 1998

167

porary fence around the nesting area to
prevent vehicular and pedestrian traffic.
In 1993 the area also contained numerous
nests of other sphecid wasps: 193 of Riih-
rica uasuta (Christ) (Pimenta and Martins
unpublished data), 54 of Ammophila graci-
lis Lepeletier (Gaimari and Martins 1996),
4 of Bicyrtes discisa, 4 of Tracln/pus sp., and
1 of Prionyx fervens (Linnaeus). There were
also a few bee nests at the same time: 7 of
Centris aenea Lepeletier, and 1 of Megnchile
iieoxanthoptera Cockerell.

We marked and released 180 females
(127 in 1993 and 53 in 1994) and 31 males
(19 in 1993 and 12 in 1994) that were cap-
tured in or near the neshng site while ei-
ther searching for a place to begin exca-
vation, in the process of excavating a nest,
flying in search of prey or in transit to the
nest, or visiting the flowers of Waltheria
americana (see Macedo and Martins 1998,
for armual records of B. angulata visits to
this plant). Numerous additional wasps
were marked in areas outside the nesting
area, but none of these wasps were sub-
sequently encountered there. Wasps were
restrained and marked individually with
fast-drying non-toxic acrylic paint using
combinations of 4 colored dots on the cor-
ners of the scutum, making it possible to
observe individual variation in daily phe-
nology, nesting (note that many marked
females made no nests in the study site),
provisioning, male behavior, longevity, in-
teractions with other insects and plants,
and movements in the aggregation and its
surroundings. Adult longevity was calcu-
lated as the length of time between when
a wasp was marked and the last time it
was seen, and is thus likely to be a very
conservative estimate, though many
wasps were never seen again after mark-
ing (64 females and 21 males), and we did
not include these individuals in the aver-
age. Differences between male and female
longevity were tested using Student's t-
test.

Nests were marked during interrup-
tions in the nest excavation process, using

4 cm-long and 5 mm-wide aluminum ar-
rows fixed to the ground with nails. Ar-
rows were distinguished by colors corre-
sponding to those of the resident female
and /or by numbers. After observing a de-
finitive nest closure, we nailed an
emergence trap (a marked plastic cup) to
the ground at the nest entrance (Fig. 2), to
record egg-adult development time and
the emergence of nest parasites. This also
helped us to estimate the number of an-
nual generations. Wasps from the first
generation that emerged in the field or in
the laboratory were marked and released
in the study site, so as not to extirpate the
population.

We excavated all 227 marked nests to
determine their architecture and contents,
assessing whether they were deserted,
parasitized, destroyed by ants or termites,
or completed. The status of the nests dur-
ing the study was categorized as follows:
(1) Deserted nests were those in which we
observed a female working but which
proved to be empty upon excavation (this
class of nests thus includes both abandon-
ment and female mortality events) (2) Par-
asitized nests were those in which para-
sites appeared in the emergence traps (3)
Nests destroyed by ants or termites were
those in which we found pieces of de-
stroyed cocoons and prey remains (4)
Completed nests were those that we ob-
served being provisioned and in which we
found all remains of cocoons and /or prey.
Only nests which were not deserted were
included in the analysis of nesting phe-
nology and counts of nests per females.

We counted 307 prey that were con-
sumed by larvae in 26 of the nests; it was
possible to count the exact number of con-
sumed prey because their heads and scuta
remained intact in the cell. Eight days af-
ter provisioning ceased, seven nests were
excavated to measure the time of devel-
opment from egg to prepupae. The num-
ber of pores of 14 cocoons were counted
and the larger and smaller diameter mea-
sured for 8 of them. We made casts of 2

168

Journal of Hymenoptera Research

Fig. 3. Architecture of the nest of Bicyrtes angulata.

completed nests with plaster of Paris to
record their shape (Fig. 3), and casts of
two incomplete nests. Nesting activity
was measured as the number of nests ini-
tiated per month. We plotted the number
of nesting females per month versus the
average number of nests made per female
to assess evidence for density-dependent
effects. For quantitative measurements we
calculated averages with standard devia-
tions.

Voucher specimens of wasp and prey
were deposited at the Laboratorio de Eco-
logia e Comportamento de Insetos of the
Departamento de Biologia Geral, ICB-
UFMG, Belo Horizonte, MG, Brazil.

RESULTS

Nest excavation. — Over the entire study,
a total of 180 females excavated 227 nests,
and 80 of these females excavated at least
one nest within a week of being marked.
In 1993, the final nest density was approx-
imately 16.5 nests/m-. We also observed
the origin of a new nest site in 1994, when
29 out of the 80 nests of B. angulata found
were in an area close to, but separate
from, the original site (Fig. 1).

Females walk in a slow and meandering
path over the nesting site, occasionally an-
tennating the ground, until they find a
suitable spot (the selection criteria are un-
known, but are likely to be tactile) and be-
gin to dig. Females dig primarily with
their mandibles, using the forelegs to
scoop loosened soil backwards, throwing
the soil out from under the body. This
movement is accompanied by a rapid
rocking motion of the body, lifting the ab-

domen each time the soil is thrown, and
simultaneously tilhng the head down over
the soil. The excavated soil accumulates,
forming a mound (tumulus) close to the
nest entrance. Over the entire period of ex-
cavating a single nest, females spend an
average of 2.26 ± 0.13 h (n = 4) actually
digging the nest, but also 2.39 ± 0.24 h (n
= 4) flying to and from the nests, or sim-
ply resting in the tunnel. Completed nests
(n = 5 with all measurements taken) have
a straight tunnel, 9.6 ± 1.1 cm long and
0.8 ± 0.1 cm in diameter, angled from 20°
to 35° relative to the soil surface (Fig. 3),
and ending in a single ellipsoid cell (only
two nests had two cells) measuring 2.6 cm
long by 1.2 cm in diameter (all 5 measured
cells identical), at 5.6 ± 0.54 cm depth. Fe-
males typically complete nest excavation
in the afternoon. Afterwards, they close
the nest entrance and fly away to feed on
nectar, and, from March 16 to April 20, to
sleep in mixed male /female aggregations
on inflorescences of Panicum maximum
(Poaceae), in proximity to adults of Rub-
rica nasuta (described below) (Fig. 4). No
form of orientation flight was ever ob-
served, either after nest construction, af-
ternoon closure, or in between provision-
ing trips.

Nest provisioning. — Out of 227 nests, 147
were succesfully provisioned by 103 dif-
ferent females. Provisioning can occasion-
ally start the same afternoon that nest ex-
cavation is finished (n = 2) but is gener-
ally begun the following day. In the latter
case, by 0830 to 0900, the females return
and open the nest entrances with their
forelegs, using motions as described
above. Hunting trips occur between 0845
and 1545 h. As they leave the nest for the
first and all subsequent hunting trips, they
emerge headfirst, and temporarily close
the entrance by scooping the tumulus
backwards into the entrance. However,
sometimes nest entrances stay partially
open due to careless and hasty closure. If
heavy rains level off the mound, females
still have little or no difficulty finding

Volume 7, Number 2, 1998

169

Fig. 4. Mixed male-female sleeping site of Bkyrtes angulata.

their nest. Sometimes they seem confused
initially, but soon find the nest entrance
and reopen it.

Prey are immatures (sometimes adults)
of Megnlotomus sp. and Apidaurus sp. (He-
miptera: Coreidae: Alydini [=Alydidae]),
accounting for 302 prey items; only one
anomalous cell was found, containing 5
pentatomid nymphs. Each prey is stung
and partially paralyzed, then the wasp
flies back to the nest holding it venter up
with her middle legs. She then removes
the nest closure with her front legs, keep-
ing the prey in the same position, and in
this way she enters the nest headfirst.
Once, after removing the nest closure, a
female dropped the prey at the nest en-
trance, entered, came back out headfirst,
grabbed the prey in her mandibles, and
dragged it down to the cell by moving
backwards. Hunting trip lengths are quite
variable (45.6 ± 35.9 min., n = 13; range
1-95). After capturing the first prey and
placing it in the nest, one egg is attached

erect on the metastemum between the
middle or hind coxae, exactly as shown by
Evans (1966:158). Since the time between
the first and second provisioning trips can
vary considerably, it appears that the egg
may hatch either before or after the second
prey is supplied, but in most cases it is
probably after two or three prey are al-
ready in the nest.

Provisioning is progressive and some-
what lengthy, lasting approximately 6
consecutive days per nest (6.5 ± 0.53, n =
7), with roughly 12 prey per completely
stocked cell (11.88 ± 4.46, n = 26; range
7-24). The final closure of completed nests
begins with females scraping the tumulus
into the nest entrance, as when making a
temporary closure, but is somewhat more
thorough, and this is followed by a bout
of "hammering," during which a female
delivers vigorous blows with the tip of the
abdomen until the nest entrance is com-
pacted and levelled. The egg-prepupa de-
velopmental period is rapid, as we found

170

Journal of Hymenoptera Research

in

ro

E

E

3

■ Number marked
D Number nesting

F M A M

1994

Fig. 5. Number of marked and nesting females of Bicyrtes angulata in 1993 and 1994. Numbers above the
bars indicate the total number of nests constructed in that time interval.

prepupae already inside their cocoons (n
= 7) when excavated 8 or 9 days after pro-
visioning had ceased. Cocoons had 4 to 6
pores (5.5 ± 1.95, n = 14) and averaged
17 ± 0.9 mm long by 5.3 ± 0.06 mm (n =
8) in diameter. The entire cycle for a single
nest, from first excavation to final closure,
typically takes 8 days (8.5 ± 0.5, n = 6).

Seasonal phenology and demographic pat-
terns.— Adults are found during the whole
year, mainly visiting flowers, but nests are
built mainly from February to May (mid-
summer and autumn; 85 of 97 provisioned
nests in 1993, and all 50 in 1994; Fig. 5).
The ratio of nests provisioned per nesting
female was generally higher than one ex-
cept in the winter and spring (June
through November, Fig. 6; in October the
ratio was higher because two females pro-

visioned five nests in total). Overall, there
is no relationship between the number of
nesting females per month and the aver-
age number of nests per female per
month. If we consider the number of nests
relative to the total number of marked fe-
males, it is obvious that many females
were present in the area that never made
nests (in 7 of 14 months in which nesting
was observed, there were more marked fe-
males alive than nests provisioned). De-
spite the fact that the range of number of
nests per nesting female was one to seven,
the average number of nests per marked
female only varied from 1.0 to 1.5 in dif-
ferent months.

Females seem to feed on flowers in ar-
eas other than those in proximity to their
nests; only three out of 77 females marked

Volume 7, Number 2, 1998

4n

171

■ Nests per marked female
D Nests per nesting female

FMAMJJASONDJFMAM

1993 1994

Fig. 6. Monthly ratio of number of nests to females of Bicyrtes aji^iilata in 1993 and 1994.

on W. americana, close to the study site,
were seen nesting in the aggregation.
However, 77 out of 103 females that were
marked while visibly searching for a place
to begin excavation were later seen nest-
ing there. Females stayed in the aggrega-
tion from 1 to 84 days (24.1 ± 17.5, n =
116). One female was observed nesting
until the age of 65 days, with no apparent
mandibular or wing wear over this time.
The estimate of male longevity (31.4 ±
20.4 days, n = 10; range 3-56) was not sig-
nificantly different from that of females
(Student's t = -1.25, p > 0.2).

There are at least 2 annual generations,
though these are staggered and overlap-

ping rather than synchronized and dis-
crete (see Table 1 ). The range of time from
oviposition to adult emergence was 44 to
375 days (200.1 ± 137.1, n = 13; Table 1).
The origin of such variability is in the pre-
pupal stage, which may remain dormant
for widely varying intervals of time. This
variability in dormancy can occur not only
among but within the broods of individ-
ual females; two eggs from one female
produced adult males after 55 and 375
days of development. The operational sex
ratio at this site was apparently female bi-
ased; 15 wasps from the first generation
emerged (in the field or from cocoons
brought into the laboratory), of which 9

172

Journal of Hymenoptera Research

Table 1. Developmental intervals of Bin/rte angulata (arranged by starting date).

Nest

Nesl starting date

Nest closing date

Adult emergence date

« dav^
egg-adiill

Sex

1

3

12 Feb 1993

13 Feb 1993

18 Feb 1993

19 Feb 1993

16 Apr 1993
9 Mar 1994

57

375

F
M

2
12

16 Feb 1993
19 Feb 1993

3 Mar 1993
26 Feb 1993

27 Apr 1993
3 Feb 1994'

55
342

M
F

4

3 Mar 1993

9 Mar 1993

19 Aug 1993

163

M

10

8 Mar 1993

7

7 Feb 1994

ca. 310

F

13
11

17 Mar 1993
30 Mar 1993

1 Apr 1993
13 Apr 1993

30 Jan 1994
3 Feb 1994*

335
296

F

M

15

5
7
6
9

1 Apr 1993

24 Apr 1993

1 May 1993

6 May 1993

13 Oct 1993

13 Apr 1993
6 May 1993
6 May 1993

7

28 Oct 1993

11 Apr 1994*
29 Jan 1994
4 Sep 1993
31 Jan 1994
24 Jan 1994

356

328

88

ca. 310
88

F

F

M

F

F

8

28 Oct 1993

5 Nov 1993

24 Jan 1994

80

F

14

11 Feb 1994

22 Feb 1994

8 Apr 1994

44

M

* These individuals emerged in thie laboratory.

were females (60%). We found 5.7 females
per male (85% females) over the period of
the study, though males were rarely seen
on the aggregation and thus their abun-
dance was likely underestimated.

Movement pat tents and diel phenology. —
Adults of B. angulata spend nights on
plants. From March 16 to April 20, marked
B. angulata and R. nasuta were seen sleep-
ing on inflorescences of Panicum maximum
(Fig. 4). The highest number of wasps seen
in a single group was 17 B. angulata and 9
R. nasuta (J. F. Macedo, pers. comm.). Fe-
males leave the sleeping site between 0800
and 0830 h to take nectar prior to the start
of nesting activities (other phenological
data above). Usually after 1600 h, or when
it is very cloudy, they return to the sleep-
ing site. One day they left the sleeping site
early, returned around 0900, but then re-
sumed nesting activities between 0900 to
1000 h, when the sun began shining again.
B. angulata comprised 34% of the individ-
uals of the 29 bee and 51 wasp species vis-
iting Waltheria americana between 1000 and
1200 h in April 1993 and April 1994 (Ma-
cedo 1995). The highest number of B. an-
gulata seen on W. americana during one
month was 35 females and 7 males in June
1993 at a site some distance from the nest-

ing site (despite the availability of flowers,
no visits were observed on the same
plants in June 1994, reflecting the large
variation in the abundance of wasps be-
tween years).

Only 8 males were captured and
marked (of 31 total) while they were pa-
trolling the aggregation by flying close to
the soil, continuously going back and
forth. Occasionally they clashed briefly in
mid-air with other males, females, or other
insects flying in the patrolling area. The
highest levels of patrolling activity were
observed in April between 0845 to 1145 h,
and 1300 to 1600 h. Despite the patrolling
activity, copulation was never seen in the
aggregation, nor on flowers of W. ameri-
cana or in the sleeping site.

Four new adults were observed emerg-
ing from their cells between 1045 and 1145
h, and new adults were generally found
in the emergence traps in the mornings,
suggesting that there is a preferred daily
emergence time.

Interactions with natural enemies and other
insects. — The female actively defends the
nest entrance against insects. Once, when
an army ant trail crossed the entrance of
one nest, the female tried to drive them
out by touching them with its abdomen or

Volume 7, Number 2, 1998

Table 2. Survivorship and success of Bici/rtcs angulata nests.

173

Nutnber ot nests per year

Nest status

1003

IW4

Total

Marked

147

80

227

Marked

Abandoned

46

30

76

Marked

Cast in plaster

4

0

4

Marked

Completed

97

50

147

Marked

Completed

Failed

83

49

132

Marked

Completed

Survived

14

1

13

hovering above them. However, we never
observed females carrying ants into the air
and dropping them a few cm away, as
Ammophila gracilis did in our study site
(Gaimari and Martins 1996). A butterfly
that was flying repeatedly over a nest was
also driven away by the resident female;
in this case the wasp hovered above the
butterfly, approaching and touching it
several times until it was driven off. Aerial
clashes were common between males and
females of B. angulata and R. nasiita. Once,
a female of B. angulata entered a nest of R.
nasuta while the latter was discarding a
prey item not consumed by her larva, and
when the Rubrica returned, she found the
Bicyrtes and drove her away. A few min-
utes afterwards, this female started to ex-
cavate her own nest close to that of the
Rubrica.

Table 2 summarizes the fate of the nests
iniHated in 1993 and 1994. Out of 227
nests, 147 were completed; 76 were aban-
doned and four were cast in plaster; ex-
cluding the latter, then, some 34% of nests
were abandoned. Apparent causes of nest
abandonment were: excavation in hard
soil (n = 8); proximity to ant nests (2);
heavy rains (2); female disturbed by a fe-
male of R. nasuta (1); and the remainder
(63) were due to unknown causes.

Of 147 provisioned nests, only 15 pro-
duced adult wasps; 100 were destroyed by
ants (principally Solenopsis sp.) and ter-
mites; 16 experienced immature mortality
of unknown nature; 7 pupae died due to
flooding of the nesting site; 6 nests were

parasitized by Ligyra morio (Diptera: Bom-
byliidae; 5 adults emerged in 5 traps); one
nest was apparently parasitized by Meto-
pia n. sp. nr. sinipalpis (Diptera: Sarco-
phagidae: Miltogramminae; emerged in
one trap), a major parasite of Ammophila
gracilis in the study site (Gaimari and Mar-
tins 1996); and two nests were excavated
prematurely to document egg placement
on prey. Excluding the latter two nests,
mortality of completed nests was approx-
imately 90%. One entire aggregation of 25
marked nests was extirpated in 1991 by
ants and termites at the Ecological Station
of the UFMG.

DISCUSSION AND CONCLUSIONS

The data from this study are only ex-
ceeded by those available for B. quadrifas-
ciata, obtained by several workers in nu-
merous localities in the United States (see
Evans 1966 for a review). As Evans (1966)
has claimed, a detailed comparison of the
behavior of the species of Bicyrtes requires
that other species like B. angulata are stud-
ied in detail. Following Evans' (1966) sum-
mary of features of the ethology of the
species of Bicyrtes, we have thus con-
firmed some general patterns within the
few members of the genus that have been
studied, as well as added new information
(Tables 2, 3 and 4), though, as mentioned,
only a few of the details of nesting and
provisioning behavior are uniform across
all species.

Adults of B. angulata are like all the oth-
er studied species in the genus in some

174

Journal of Hymenoptera Research

Table 3. Comparison of qualitative features of Bicyrtes nesting biology.

aiigtiliifa

W)St/Sd

I'liric^ala

^imillinui

qiiaiirifii^cttila

Provisioning

progressive

mass

mass

mass

delaved

Prey paralyzation (partial)

yes

yes

yes

yes

7

Prey types*

N a 4 (+A)

N >4

N5

N5

N

Prey families**

C

P

P

P

CPRLSCy

Foraging during morning-afternoon

yes

?

7

yes

yes

Mound (tumulus) near nest entrance

yes

yes

no

yes

no

Nests aggregated***

yes

no

yes

no

no

Accessory burrow

no

no

yes

yes

7

Sleep in accessory burrow

no

no

?

yes

7

Sleep in mixed association

yes

7

7

no

7

Citation****

present work

C1982

G1982

G1982

El 966

* N = nymphs (followed by instar numbers if specified), A = adults, ( + A) indicates adults rarely taken.

** C = Coreidae (s.l.), P = Pentatomidae, R = Reduviidae, L = Lygaeidae, S = Scutelleridae, Cy = Cydnidae,

Rh = Rhopalidae, Py = Pyrrhocoridae.

*** Statistics not available; category represents subjective evaluation of authors as to dispersion of nests relative

to apparent available space.

****G1982 = Genise 1982, E1966 = Evans 1966, E&M1974 = Evans & Matthews 1974, S&G1992 = Sanchez &

Genaro 1992.

respects: (1) they are frequent visitors at
flowers for nectar (they do not feed upon
the prey they capture for their larvae) (2)
they are solitary ground-nesting predators
(3) prey are immature and /or adult het-
eropterans (4) nest closure is maintained
while outside the nest (5) foraging is pri-
marily in the morning and afternoon. Oth-
er features which appear similar to the
majority of species studied include: (1)
nesting in multispecies aggregations (2)
sleeping on vegetation (3) partial paralysis
of prey (4) nests with one or two cells (5)
low nest angles (generally less than 45°).

Other features were atypical, such as: (1)
nesting in sandy soil far from water (other
species except for B. spiiiosa nest along wa-
ter courses) (2) relatively short burrows
(less than 10 cm on average) (3) more than
two hours to dig a nest. The presence of a
tumulus can neither be considered excep-
tional nor general, and for other features
comparative information is lacking.

A few of these points merit further dis-
cussion. Despite the general tendency of B.
angulata, like other species, to intersperse
its nests with those of other digger wasp
and bees, it tends to aggregate intraspecif-

Table 4. Comparison of quantitative features of Bicyrtcs nesting biologv.

iitii^iiiiiiii

ilisani

t'llrif^tllii

^ittnlhttiil

i^iiitiinfii^iiitlii

Number of cells per burrow

1-2

3

2-5

1

2-3

Number of prey per cell

7-24

16

3-6

11

8-11

Time to dig cell /nest (min)

146 ± 14

30

60

>60

60-120

Angle of entrance burrow

20°-35°

30°-35°

30°-35°

30°

30°-60°

Nest depth (cm)

5.6 ± 0.54

5

8-10

20

7

Nest length, burrow plus cell (cm)

9.6 ± 1.1

12

12

24

8-43

Lengt of cell (mm)

25-26

18-20

25-30

17

20-35

Width of cell (mm)

12-13

10-12

12-15

12

5-27.5

Number of pores in cocoon

5.5 ± 1.95

7

5

?

7

Egg-adult developmental time (days)

44-375

7

7

?

49

Number of generations per year

2

7

7

7

7

Citations as for Table 3.

Volume 7, Number 2, 1998

175

Table 3. Extended.

nlr„h~

lo<lu->!<

'iVn/.if.;

delayed ? progressive progressive

? ? ? no

N N + A N + A N( + A)

CPRLSCy PS Rh CPLSPy

? ? yes yes

no no no yes

variable ? ? variable

? ? ?

? ? ?

? 7 7

El 966

E1966 E&M1974

no
no

no
S&G1992

ic nests within these areas. As in its con-
geners, B. angulata maintains an outer clo-
sure at all times when the female is not in
the nest, but no inner closure is ever
made. Prey are Coreidae (sensu lato), in-
dicating in our case a possibly high level
of local specialization, compared for ex-
ample with the Cuban generalist B. spinosa
that preys upon several species of five het-
eropteran families (Sanchez and Genaro
1992). Additionally, the prey is partially
paralyzed and carried in flight with the
middle legs, clasped tightly beneath the
base of the wasp's abdomen, and is not
usually deposited on the ground at any
time. The egg is laid in a semierect posi-
tion on the mid-ventral line of the first
prey item. Our observations established
that B. angulata is a progressive provision-

Table 4.

Extended.

:y',ti.lh~

h;lu-il<

.,.,„;,„„

,.,„„„,

1-2

3-11

60-120

2-5
10-23

7

1
7

7

1

7

7

45°

4-8

20-30

20-30

45°-60°
8-13.5
12.5-18.5

7

20°-45°
8.5-10.5
12-14

7

45°

4.5-8.5

9.9 ± 2.6

7

8-12

7

10-12

7

?

7

7

40-42

7
7

7.6

± 1.54

7

'

-)

7

7

er like B. spinosa and B. cingulata, though
mass provisioning and delayed provision-
ing are found in other species.

Among the novel data for B. angulata is
the huge variability in the time of prepu-
pal dormancy within the same brood. This
variability (sometimes called "parsivoltin-
ism"; Torchio and Tepedino 1982) is scarce-
ly documented for Nearctic and Neotrop-
ical species of solitary wasps and bees (see
Evans 1966; Stephen et al. 1969; Evans and
West-Eberhard 1970; Torchio and Tepedi-
no 1982; Roubik 1989; Wcislo and Cane
1996). A comparable variability was also
found for other species of solitary wasps
and bees in the same study site or in other
sites in the Ecological Station of the UFMG
(Martins et al. 1996; Almeida et al. 1997;
Pimenta and Martins unpublished data).
One of the possible interpretations for
such a pattern is given by theories of risk
spreading of reproductive effort (or "bet-
hedging") in unpredictable environments
(Danks 1987; Tauber et al. 1986). The pos-
sibility that parsivoltinism may help avoid
predation or parasitism in some way
seems reasonable, but we cannot confi-
dently identify the precise mechanism of
selection for such an adaptation in the
present case. We do believe, however, that
we can exclude variability in the occur-
rence of rains (as in deserts where similar
insect life cycles occur, e.g. Hanski 1988),
because our study site is strikingly season-
al and predictable in the occurrence of
rains (see Martins and Antonini 1994) and
dormancy seems to not correlate with
rains (R. P. Martins unpublished data).

There is an apparent paradox in the
data on number of marked females versus
number of nests made (see Fig. 5). Many
marked females visiting flowers at the
study site did not nest there, so the total
number of females exceeded the number
of nests in 7 of the 14 months when fe-
males were present. Why are there so
many active females that are apparently
not nesting? This same phenomenon oc-
curs in R. nasuta (Pimenta and Martins un-

176

Journal of Hymenoptera Research

published data), and for that species, prey
are seasonally distributed within the year,
and the hypothesis is that females should
wait for a burst of prey availability to
make nests. It is likely that prey abun-
dance for B. angulata also exhibits seasonal
variation, because many plant-feeding in-
sects in this environment have strongly
seasonal life cycles, and a similar expla-
nation may apply here. It is alternatively
possible, though questionable, that these
excess females were nesting elsewhere but
all foraging in a limited area together; i.e.,
the females sampled at the flowers repre-
sented the combined population of several
nesting areas. However, this was not the
only patch of suitable flowers in the area,
and there is no obvious reason why the
wasps would be concentrated in this par-
ticular patch.

Because we found no relationship be-
tween the number of nesting females and
the average number of nests per female,
we suggest that crowding has no effect on
the number of nests made in the aggre-
gation, despite the preference to nest in a
somewhat limited area. It is also possible
that females made only one or two nests
in the aggregation and then dispersed to
other areas, given that a female can live
for up to 84 days, yet few were in resi-
dence this long. That is, rather than inter-
preting the small number of nests per fe-
male as a result of high mortality, there
may be some undetected nesting activity
outside of the known aggregation area;
further work with marked females might
help resolve this. In either case, the avail-
ability of adequate patches (whether ag-
gregations form in them or not) may be of
primary importance in the nesting dynam-
ics of this species. In addition to other
forces that could potentially influence
nesting dynamics is the frequent extirpa-
tion of nests or entire aggregations by ter-
mites and ants, and the high frequency of
nest desertion, due to several causes.
Therefore, the patchiness of the environ-
ment and the dynamics within the aggre-

gations are likely to be crucial in the pop-
ulation dynamics of this species.

ACKNOWLEDGMENTS

Servio Tiilio P. do Amarante identified Bicyrtes an-
gulata, B. discisa, B. tricolorata, Rubrica nasuta, and Pri-
onyx fervens. Padre J. S. Moure identified Megachile
ncoxantJwfJtera and Centrii aenea. Arnold S. Menke
identified Amincphila gracilis. ]. A. M. Fernandes iden-
tified McgalotcniKS sp. and Apidaunis sp. The late Her-
mogenes F. Leitao Filho identified Walthetia amcncana
and ]. F. Macedo Panicum maximum. We would also
like to thank William T. Wcislo for useful comments
and criticisms. The Brazilian Conselho Nacional de
Desenvolvimento Cientifico (CNPq) and Fundai^ao de
Amparo a Pesquisa de Minas Gerais (FAPEMIG) con-
ceded grants, the CNPq also provided research schol-
arships to the senior authors (R.P.M. and L.A.S.), a
Visifing Researcher Fellowship (301019/96-7 RN) to
the junior author (D.Y.), and the U. S. Fish and Wild-
life Service provided some logistical support. This
study is a contribution of the Program in Ecology,
Conservation and Wildlife Management (ECMVS) of
the Universidade Federal de Minas Gerais, Belo Ho-
rizonte, MG, Brazil.

LITERATURE CITED

Almeida, D. A. O., R. P. Marfins and M. L. T. Bus-
chini. 1997. Behavior and nesting dynamics of
the Neotropical cavity-nesting specialist bee
MegachUe assumptinnis Schrottky, with compari-
sons to the Nearctic Megadule brcvis Say (Hy-
menoptera: Megachilidae). ]ouriud of Hymenoptera
Research 6: 344-352.

Bohart, R. M and A. S. Menke. 1976. Sphecid Wasps of
the World, University of California Press, Berke-
ley, CA. 695 pp.

Danks, H. V. 1987. Insect Dormancy: an ecological
perspective. Biological Survey of Canada Mono-
graph Series, No. 1. Ottawa, 439 pp.

Evans, H. E. 1966. The comparative ethology and evolu-
tion of the sand wasps. Harvard University Press,
Cambridge, MA, 526 pp.

Evans, H. E. and M. J. West-Eberhard. 1970. The
u'asps. University of Michigan Press, Ann Arbor,
Ml. 265 pp.

Evans, H. E. and R. W. Matthews. 1974. Observations
on the nest behavior of South American sand
wasps (Hymenoptera). Biotropica 6 (2): 130-134.

Gaimari, S. D. and R. P. Martins. 1996. Nesting be-
havior and nest distributions of Ammophila grac-
dis Lepelefier (Hymenoptera; Sphecidae) in Bra-
zil. Journal of Hymenoptera Research 5: 240-248.

Genise, J. F. 1979. Comportamiento de nidificacion de
Bic\/rtes variegata (Ol.) e Bicyrtes discisa (Tasch.)
(Hymenoptera: Sphecidae). Rei'ista da Sociedad
Entomologica Argentina 38 (1-4): 123-126.

Volume 7, Number 2, 1998

177

Genise, J. F. 1982. Estudios sobre el comportamiento
de Bembicini neotropicales. I. Bici/rtes simUlima
(Smith) y Bkyrtes discisa (Tasch) (Hymenoptera:
Sphecidae) con una revision de los tipos de
aprovisionamiento em Bembicini. Physis (Buenos
Aires) Secc. C, 40 (99): 93-99.

Hanski, I. 1988. Four kinds of extra long diapause in
insects: a review of theory and observations. An-
nales Zootogici Fennici 25: 37-53.

Macedo, J. F. 1995. A importancia de Walthcria indica
(Sterculiaceae) para abelhas e vespas. M. Sc. Dis-
sertation, Universidade Federal de Minas Gerais,
Belo Horizonte, M.G., Brazil. 74 pp.

Macedo, J. F. and R. P. Martins. 1998. Potencial da
erva daninha Walthcria ainericana (Sterculiaceae)
no manejo integrado de pragas e polinizadores:
visitas de abelhas e vespas. Anais da Soctedade En-
tomologica do Brasil 27: 29-40.

Martin, P. and Bateson, P. 1986. Measuring Behavior.
Cambridge University Press, Cambridge, U.K.
200 pp.

Martins, R. P. 1991. Biologia e Comportamento de
Comunidades de Vespas Escavadoras (Hyme-
noptera: Aculeata). Ph.D. Dissertation, Universi-
dade Estadual de Campinas, Campinas, S.P., Bra-
zil. 116 pp.

Martins, R. P. and D. A. O. Almeida 1994. Is the bee
Megaehile assumptionis a cavity-nesting specialist?
lournal of Insect Behavior 7 (5): 754-765.

Martins, R. P. and Y. Antonini. 1994. The biology of
Diadasina distincta Holmberg 1803, (Hymenop-
tera: Anthophoridae). Proceedings of the Entomo-
logical Society of Washington 96 (3): 553-560.

Martins, R. P., F. G. Guimaraes and C. M. Dias. 1996.
Nesting biology of Ptilothnx plumata Smith, with
a comparison to the other species in the genus
(Hymenoptera: Anthophoridae). Journal of the
Kansas Entomological Society 69 (1): 9-16.

Nascimento, P. T. R. and W. L. Overall. 1980. Catal-
ogo da Cole^ao Entomologica do Museu Goeldi.
Hymenoptera: Sphecidae. Boletim do Museu Par-
tiensc Eniilio Goeldi, Zoologia 99: 1-14.

Roubik, D. W. 1989. The Ecology and Natural History
of Tropical Bees. Cambridge Tropical Biological
Series. Cambridge University Press, Cambridge,
U.K. 514 pp.

Sanchez, C. S. and J. A. Genaro. 1992. Observaciones
sobre la conduta de nidificacion en esfecidos de
Cuba (Hymenoptera). Bicyrtes spinosa (Fabr.) Poy-
eana 411: 1-8.

Stephen, W. P., G. E. Bohart and P. F. Torchio. 1969.
The hiology and external morphology of bees. Agri-
cultural Experimental Station, Oregon State Uni-
versity, Corvallis, OR. 140 pp.

Tauber, M. J., C. A. Tauber and S. Masaki. 1986. Sea-
sonal Adaptations of Insects. Oxford Uruversity
Press, NY. 411 pp.

Torchio, P. F., and V. J. Tepedino. 1982. Parsivoltin-
ism in three species of Osmia bees. Psyche 89:
221-238.

Wcislo, W. T., and J. H. Cane. 1996. Floral resource
utilization by solitary bees (Hymenoptera: Apo-
idea), and exploitation of their stored food by
natural enemies. Annual Review of Entonwlo<fy 41:
257-286.

Willink, A. 1947. Las especies Argentinas de Bembi-
cini. Acta Zoologica Lilloana 4: 509-651.

J, HYM. RES.
Vol. 7(2), 1998, pp. 178-181

Sexual Dimorphism Of Wasp Antennal Structure in Relation to
Parasitic and Non-parasitic Behavior (Hymenoptera: Sphecidae)

William T. Wcislo

Smithsonian Tropical Research Institute, Balboa, Republic of Panama
(Address for correspondence; Smithsonian Tropical Research Institute,
Unit 0948, APO AA 34002-0948, USA; E-mail, WcisloW@tivoli.si.edu)

Abstract. — To assess the relationship between sensory ecology and behavior of non-parasitic
and parasitic spheciforme wasps (Sphecidae), I measured the lengths of scapes, flagella, and body
size (intertegular distance) of males and females of 29 species, representing 7 subfamilies and 26
tribes. Unlike a previous study with bees (Wcislo, 1995), spheciforme wasps show no consistent
sexual dimorphism in relative antennal size for free-living versus parasitic species.

Brood parasitism (cuckoo behavior) and
social parasitism have evolved repeatedly
among bees, aculeate wasps and ants (e.g.,
Wcislo 1987; Wcislo and Cane 1996; H611-
dobler and Wilson 1990; Cervo and Dani
1996). Parasites utilize host-derived re-
sources (a nest, stored food, or worker la-
bor) to rear their own offspring. Maternal
behavior of parasitic and non-parasitic
species differs (Wcislo 1987), while respec-
tive males do not differ essentially in mat-
ing behavior, although data are scant (e.g.,
compare Cederberg et al. 1984 with Al-
cock and Alcock 1983). Few studies have
investigated the sensory ecology (sensu
Dusenbury 1992) of parasitic and non-par-
asitic Aculeata to ascertain if differences in
sensory structures co-occur with behavior-
al differences. Non-parasitic bees (Apo-
idea) usually are strongly sexually dimor-
phic for antennal structures; at a given
body size, males tend to have shorter
scapes and longer flagella (Wcislo 1995;
Miiller 1872). Parasitic bees, in contrast,
usually are not sexually dimorphic for rel-
ative size of antennal structures. Among
ants, a fusion of antennal flagellomeres is
part of a syndrome of structural characters
associated with parasitic behavior (Holl-
dobler and Wilson 1990).

Some clades of spheciforme wasps ( =
"Sphecidae" of Bohart and Menke 1976)
together with the bees form a monophy-
letic group, Apoidea (e.g., Alexander 1992;
Brothers and Carpenter 1993). Parasitism
has evolved repeatedly in bees (e.g., Wcis-
lo and Cane 1996), and has probably
evolved twice among spheciforme wasps,
once in the common ancestor of the genus
Stizoides and once in the common ancestor
of Nyssonini (see Bohart and Menke 1976).
Thus, the evolution of parasitism among
sphecid wasps provides additional exam-
ples to assess whether female parasites are
similar to males in their sensory ecology
and relevant structures. This note presents
information on antennal size for parasitic
and non-parasitic wasps (Sphecidae), as
part of an on-going comparative study of
the relationship between morphological
and behavioral evolution within aculeate
Hymenoptera (cf. Wcislo 1989).

MATERIALS AND METHODS

In an effort to minimize phylogenetic
bias I used 29 species, representing 7 sub-
families and 26 tribes of the 10 subfamilies
and 30 tribes that Menke (1997) lists for
Sphecidae (see Appendix). Intertegular
distances, scape length, pedicel length.

Volume 7, Number 2, 1998

179

and total flagellar length was measured on
5 individuals of each sex using methods
in VVcislo (1995). Values are reported as
means with standard errors. Data were
analyzed using SYSTAT (Wilkinson 1988)
on a personal computer, unless otherwise
indicated. Phylogenetic bias probably ex-
ists within these data due to noninde-
pendence of the taxa (e.g., Harvey and Pa-
gel 1991). There is, however, no widely-
accepted phylogenetic hypothesis avail-
able for the taxa included here, and tax-
onomy may be a poor indicator of phylog-
eny (see discussion in Alexander 1992).

RESULTS

Parasitism is relatively rare among
spheciforme wasps. Among non-parasitic
species males and females, on average,
were not significantly different in body
size (mean intertegular distance in mm: fe-
males, 1.95 ± 0.17; males, 1.66 ± 0.15;
Mann-Whitney U = 402.5, P > 0.2). The
sexes did not differ in mean length of the
flagella (in mm, females: 3.44 ± 0.46;
males: 3.28 ± 0.39; Mann-Whitney U =
348, P > 0.8 ) (Figure 1, bottom).' Males
had significantly smaller mean scape
length than females (in mm, females: 0.55
± 0.053; males: 0.43 ± 0.04; Mann-Whit-
ney U = 461, P = 0.02; Figure 1, top).
Within species, non-parasitic females
more frequently had a larger body size
than males (two-tailed sign test, P < 0.05,
T = 22, N = 26), and had longer scapes
(two-tailed sign test, P < 0.05, T = 23, N
= 23), but females did not have longer fla-
gella (two-tailed sign test, 0.1 > P > 0.05,
T = 17, N = 25) (sample sizes differ be-
cause ties were eliminated; Conover 1971).
Regression equations for non-parasitic
males versus females were not significant-
ly different for scape or flagellum length
as a function of body size (P > 0.05, com-
paring y-intercepts or slopes). Male and
female parasites did not differ in body
size, nor in lengths of scapes and flagella.

2,0

z

LU

LU

<

1 2 3

BODY SIZE, mm

12 3

BODY SIZE, mm

Fig. 1. Scape (top) and flagellum (bottom) length of
spheciforme wasps as a function of body size (inter-
tegular distance). Open circle = non-parasitic female;
open triangle = non-parasitic male; closed circle =
parasitic female; closed triangle = parasitic male.

DISCUSSION

Among non-parasitic spheciforme wasps
females often are larger than males, and
have longer scapes. Small samples pre-
clude statistical analyses, but parasitic fe-
males are not conspicuously larger than
conspecific males, and have similarly sized
scapes and flagella. In contrast, a study of
114 bee species showed that 1) female non-
parasitic bees of a given body size have

180

Journal of Hymenoptera Research

longer scapes but shorter flagella than con-
specific males, and 2) for parasitic bees,
scapes and flagella are siniilar in size, on
average, between males and females of a
species (Wcislo 1995; also Miiller 1872).
Since bees are closely related to, and prob-
ably arose from within spheciforme wasps
(e.g., Alexander 1992), the sexual differ-
ences are probably derived among bees.
Relative to non-parasitic females, female
parasites may be more "male-like" in their
search behavior, but pertinent data are
scarce (refs. in Wcislo 1995). If substanti-
ated, these behavioral differences can help
explain similarities in sensory structures
among males of parasitic and non-parasitic
species and females of parasitic species,
which differ from non-parasitic females.

Antennae have olfactory, gustatory, and
tactile sensory receptors; they are used
like calipers during nest construction; and
they can be used to drum, tap, or stroke
parts of the female's body (refs. in Wcislo
1995). The scape of some male spheciforme
wasps is broadly expanded (e.g., Dinetus),
like males of a parasitic bee, Doeringiella
(Roig-Alsina 1989); these expansions may
house glands for use during courtship and
mating, as known for other Hymenoptera
(Isidoro et al. 1996). The functional mor-
phology of antennae has been studied for
relatively few species, and typically only
for one or two sensory modalities. Anten-
nae have multiple functions, highlighting
the need for more detailed studies relating
behavior to an animal's sensory world
("utmvelt"), as pointed out long ago by
von Uexkiill (1934).

ACKNOWLEDGMENTS

I am grateful to Richard Hoebeke for unrestricted
access to the Cornell University Insect Collection; and
to Arnold Menke for arranging the loan of specimens
of two important species from the National Museum
of Natural History of the Smithsonian Institution.
Hermogenes Fernandez helped with some measure-
ments. An anonymous reviewer provided very help-
ful comments. General research funds from the
Smithsonian Tropical Research Institute are gratefully
acknowledged.

LITERATURE CITED

Alcock, J. and J. P. Alcock. 1983. Male behaviour in
two bumblebees, Bonihus nevadeniiis aurkimms
and B. grisciccllis (Hymenoptera: Apidae). jcurnal
cf Zoology. London 200: 561-570.

Alexander, B. A. 1992. An exploratory analysis of cla-
distic relationships within the superfamily Apo-
idea, with special reference to sphecid wasps
(Hymenoptera). lournal of Hymenoptera Research
1: 25-61.

Bohart, R. M. and A. S. Menke. 1976. Spheaci Wasps
of the Worhl, A Generic Revision. Berkely: Univer-
sity of California Press, i-ix + 1-695 pp.

Brothers, D. J. and ]. M. Carpenter. 1993. Phylogeny
of Aculeata: Chrysidoidea and Vespoidea (Hy-
menoptera). ]ourunI of Hymenoptera Research 2:
227-304.

Cederberg, B., B. G. Svensson, G. Bergstrom, M. Ap-
pelgren, and I. Groth. 1984. Male marking pher-
omones in north European cuckoo bumble bees,
Psitliyrus (Hymenoptera, Apidae). Nova Acta Re-
giae Societatis Scientiarum Upsaliensis, Serie V:C 3:
161-166.

Cervo, R. and F. R. Dani. 1996. Social parasitism and
its evolution in Polistes, pp. 98-1 12. In: Turillazzi,
S. and M. ]. West-Eberhard (eds.). Natural History
and Evolution of Paper Wasps New York: Oxford
University Press, i-xiv + 400 pp.

Conover, W. J. 1971. Practical Nonparametric Statistics.
New York: John Wiley & Sons, Inc., i-x + 462
pp.

Dusenbury, D. B. 1992. Sensory Ecology. New York:
W.H. Freeman & Co. i-xx + 558 pp.

Harvey, P. H. and M. D. Pagel. 1991. The Comparative
Method in Evolutionary Biology. New York: Oxford
University Press, i-iv + 239 pp.

Holldobler, B. and E. O. Wilson. 1990. The Ants. Cam-
bridge: Harvard University Press, pp. i-xii + 732
pp.

Isidoro, N. F., F. Bin, S. Colazza, and S. B. Vinson.
1996. Morphology of antennal gustatory sensilla
and glands in some parasitoid Hymenoptera
with hypothesis on their role in sex and host rec-
ognition. Journal of Hymenoptera Research 5: 206-
239.

Menke, A. S. 1997. Family-group names in Sphecidae
(Hymenoptera: Apoidea). Journal of Hymenoptera
Research 6: 243-255.

Miiller, H. 1872. Anwendung der Darwinschen Lehre
auf Bienen. Verhandlungen des naturhistorischen
Vereines der preiissischen Rheinlande uiul Westphal-
ens 29: 1-96.

Roig-Alsina, A. 1989. A revision of the bee genus
Doeringiella (Hymenoptera, Anthophoridae, No-
madinae). University of Kansas Science Bulletin 53:
576-621.

Uexkiill, von J. 1934. A stroll through the worlds of
animals and men, pp. 5-80 [reprinted and trans-

Volume 7, Number 2, 1998

181

lated, 1957]. In: Schiller, C. H. (ed.) Instinctive Be-
havior: The Development of a Modern Concept. Lon-
don: Methuen & Co. Ltd.

Wcislo, W. T. 1987. The roles of seasonality, host syn-
chrony, and behaviour in the evolutions and dis-
tributions of nest parasites in Hymenoptera (In-
secta), with special reference to bees (Apoidea).
Biological Reviezva of the Cambridge Philosophical So-
ciety 62: 415-443.

Wcislo, W. T. 1989. Behavioral environments and
evolutionary change. Annual of Ecology & System-
atics 20: 137-169.

Wcislo, W. T. 1995. Sensilla numbers and antennal
morphology of parasitic and non-parasitic bees
(Hymenoptera: Apoidea). International journal of
Insect Morphology & Embryology 24: 63-81.

Wcislo, W. T. and J. H. Cane. 1996. Resource utiliza-
tion by solitary bees (Hymenoptera: Apoidea),
and exploitation by their natural enemies. Annual
Revierv of Entomology 41: 257-286.

Wilkinson, L. 1988. SYSTAT. Evanston IL: SYSTAT
Inc.

APPENDIX

List of spheciforme wasp species from which an-
tennal measurements were taken. * = parasitic taxon;
? = taxon is probably parasitic, but behavioral data
are unavailable. Nomenclature follows Bohart and
Menke (1976), as modified by Menke (1997).

AMPULICINAE
AMPULICINI

Ampulex compressa (Fabricius)
DOLICHURINI

Dolichurus cornicuhis (Spinola)

ASTATINAE
DINETINI

Dinetus pictus (Fabricius)
ASTATINI

Astata mexicana Cresson

PHILANTHINAE
EREMIASPHECIINI

Eremiasphecium schmiedeknechtii Kohl
APHILANTHOPSINl

Aphilanthops friguius (Smith)
PHILANTHINI

Philanthus solivagus Say
CERCERINI

Cerceris frontata Say

PEMPHREDONINAE

PSENINI

Psenulus pallipes (Panzer)
PEMPHREDONINI

Stigmus americanus Packard

SPHECINAE
AMMOPHILINI

Ammoplula polita Cresson
SCELIPHRONINI

Podium nifipes Fabricius
SPHECINl

Sphex dorsalis ( = singularis} Smith

BEMBICINAE
HELIOCAUSINI

Heliocausus larroides (Spinola)
MELLININI

Mellinus arvensts (Linnaeus)
STIZINI

Bembicinus U'heeleri Krombein & Willink
*Stizoides unicinctus { = renicinctus) (Say)

GORYTINI

Gorytes simillimus Smith
*NYSSONINI

'Nysson (Epinysson) mellipes (Cresson)
*7Synnevrus aequalis (Patton)
'Nyssoii simplicicornis Fox

BEMBICINI

Bembix texana Cresson

CRABRONINAE
LARRINI

Larra bicolor Fabricius
MISCOPHINl

Solierella plenoculoides (Fox)
PALARINI

Palarus latifrons Kohl
TRYPOXYLONINl

Trypoxylon lactitarse Saussure
SCAPHEUTINI

Scapheutes brasilianus Handlirsch
CRABRONINI

Crabro cribrellifer (Packard)
OXYBELINI

Oxybclus emarginatus Say

]. HYM. RES.

Vol. 7(2), 1998, pp. 182-208

Systematics of Costa Rican Meteonis (Hymenoptera: Braconidae:
Meteorinae) Species Lacking a Dorsope

Nina M. Zitani, Scott R. Shaw, and Daniel H. Janzen

(NMZ, SRS) Department of Renewable Resources, University of Wyoming, Laramie, Wyoming,

82071-3354, USA, ninaz@uwyo.edu, braconid@uwyo.edu; (DHJ) Department of Biology,

University of Pennsylvania, Philadelphia, Pennsylvania 19104, USA, djanzen@sas.upenn.edu

Abstract. — This study of the genus Meteonis (Hymenoptera: Braconidae: Meteorinae) treats the
19 known Costa Rican species that lack a dorsope (a pit on the dorsal surface of the first metasomal
tergite). Fourteen new species all attributed to Zitani are described and illustrated: M. alejandro-
masisi, M. camilocamargoi, M. coffeatus, M. corniculatus, M. desmiae, M. dos, M. mariamartae,
M. megalops, M. micrommatus, M. pseudodimidiatus, M. rogerblancoi, M. sterictae, M. uno,
and M. yamijuanum. An identification key to 19 species is provided, including the previously
described species: M. cougregatus Muesebeck, M. dimidiatus (Cresson), M. lapln/gnme Viereck, M.
papiliovorus Zitani, and M. rubens (Nees). Biological information, host associations, and cocoon-
forming behavior are included for M. cougregatus, M. rubens and the new species M. alejandromasisi,
M. camUocamargoi, M. desmiae and M. sterictae. This work provides the first record of Meteorus
attacking Megalopygidae and Hesperiidae including Chiomara asi/chis (StoU), Pyrgus sp., Stnph\/lus
azteca (Scudder), and Staphylus sp. Other new host records include the pyralids Desmia tages
(Cram.), Omiodes stigmosalis (Warr.), and Stericta albifasciata (Druce), and the sphingids Manducn
sexta (L.), and Unzela japix (Cram.). Meteorus cougregatus, M. dimidiatus, and M. rubens are recorded
for the first time in Costa Rica.

The Meteorinae is a moderately large,
cosmopolitan subfamily with at least 183
described species worldwide. The subfam-
ily consists of solitary or gregarious koino-
biont endoparasitoids of larval Coleoptera
and Lepidoptera (Muesebeck 1923; van
Achterberg 1979; Huddleston 1980). The
genus Meteorus Haliday is the most di-
verse and frequently encountered genus
of the Meteorinae {seiisii Shaw 1985, 1995;
Maeto 1990b; Shaw and Huddleston 1991),
and is the sister-group of the subfamily
Euphorinae Foerster s.s. (Shaw 1985,
1988). For a detailed review of the biology
of the Meteorinae see Shaw (1997), Zitani
et al. (1997), Shaw (1995), and Shaw and
Huddleston (1991).

There has been some revisionary work
on the Meteorus species that occur in
North America (Muesebeck 1923), Africa
(Nixon 1943), the western Palearctic re-

gion (Huddleston 1980), and Australia
and New Guinea (Huddleston 1983), but
none for the Neotropical region. Although
six species of Meteorus have been de-
scribed from various localities in the neo-
tropics (Shenefelt 1969), and the North
American species M. laphygmae Viereck
has been recorded from Costa Rica (Shene-
felt 1969), the first new species from Costa
Rica was described only recently (Zitani et
al. 1997).

In order to keep the present work at a
reasonable length only those species lack-
ing a dorsope are treated here. A dorsope
is a pit lying anterior to the spiracles on
the dorsal surface of the first metasomal
tergite; these pits occur in a pair (Fig. 1).
Species lacking a dorsope (Figs. 3, 5) rep-
resent the majority of the material from
the examined Costa Rican collections. This
character has been used to separate spe-

Volume 7, Number 2, 1998

183

cies of Meteorus by several previous au-
thors (Muesebeck 1923; Huddleston 1980;
Maeto 1989, 1990a). This group of species
was also chosen because considerable bi-
ological information was available for 6
included species.

METHODS

Species treated in this work can be iden-
tified as members of the subfamily Meteor-
inae using the keys of Shaw (1995) or
Shaw and Huddleston (1991). Specimens
can be determined as Meteorus using the
key of Shaw (1997) or Marsh et al. (1987).

Approximately 1,200 specimens of Me-
teorus were examined. The following in-
stitutions provided specimens for this
study:

ANS: The Academy of Natural Sci-
ences, Philadelphia, PA, USA.

INBio: Instituto Nacional de Biodiver-
sidad, Santo Domingo de Here-
dia, Costa Rica.

PAS: The Polish Academy of Sciences,

Warszawa, Poland.

RMSEL: Rocky Mountain Systematic
Laboratory, University of Wyo-
ming, Laramie, WY, USA.

USNM: United States National Museum,
Smithsonian Institution, Wash-
ington, DC.

UCR: Museo de Insectos, Universidad
de Costa Rica, San Pedro, San
Jose, Costa Rica.

Specimens reared by D.H. Janzen and
W. Hallwachs at the Area de Conserva-
cion Guanacaste (ACG), Guanacaste Prov-
ince, Costa Rica, are identified by a vouch-
er number with "SRNP" in the middle.
Each Janzen and Hallwachs rearing
voucher number is for a single host larva.
Information about the rearings can also be
found at http:/ /janzen.sas.upenn.edu/ in-
dex.html.

The majority of the specimens were pre-
pared and mounted at RMSEL using the
technique of Sharkey (1988). Specimen la-
bel data were recorded exactly as they ap-

pear on the collection label to avoid mis-
interpreting data. However, when avail-
able, collection site information is added
in brackets []. Authorship of all new spe-
cies is attributed to the senior author
(NMZ).

TAXONOMIC CHARACTERS AND
TERMINOLOGY

The taxonomic characters used here fol-
low the work of Huddleston (1980, 1983),
Maeto (1989, 1990a), Shaw (1985), and Zi-
tani et al. (1997). Microsculpture terminol-
ogy follows that of Harris (1979). Morpho-
logical terminology follows Wharton et al.
(1997).

As noted by Huddleston (1980), many
characters of the head are important for
distinguishing one species from another,
especially the following: the size and de-
gree of convergence of the eyes, width of
the face in relation to the height, and the
amount of twist of the mandibles. In this
study, eye size was determined by the
length of the eye, in anterior view, relative
to the length of the head in anterior view.
Eyes were designated as 'small' if the
head height was equal to or greater than
1.6X the eye height in anterior view, and
'large' if the head height was less than
1.6X the eye height in anterior view. Eye
convergence was designated as 'strongly
convergent', 'convergent', or 'nearly par-
allel'. Eyes were called 'strongly conver-
gent' if the maximum /minimum face
width ratio was 1.7 or greater, 'conver-
gent' if the maximum /minimum face
width ratio was 1.4-1.7, and 'nearly par-
allel' if the maximum/minimum face
width ratio was less than 1.4.

The amount of twist of the mandibles
can sometimes be difficult to determine.
An untwisted mandible, in anterior view,
is flat and has two visible teeth. In a
strongly twisted mandible it appears as
though the ventral tooth has rotated to a
position posterior to the dorsal tooth.
Thus the mandible, in anterior view, has
only one apparent tooth, and the apical

184

Journal of Hymenoptera Research

portion appears twisted (Fig. 9). Also,
mandibles that are untwisted are usually
larger (longer and broader in anterior
view) than strongly twisted mandibles.
Moderately twisted mandibles, where the
ventral tooth has only partially rotated,
are the most difficult to determine but are
present in only two species, Meteorus cof-
featiis and Meteorus micrommatus (Fig. 10).
Moderately twisted mandibles are also
longer, and broader at the base, compared
to strongly twisted mandibles.

The number of flagellomeres and size of
the ocelli usually vary only slightly within
a species, and these characters, therefore,
are also useful (Huddleston 1980). In this
text, the diameter of a lateral ocellus is ex-
pressed relative to the length of the ocell-
ocular distance (distance from edge of lat-
eral ocellus to border of compound eye).
The acronym OCD is used for ocellar di-
ameter, and OCOD for ocell-ocular dis-
tance. The term 'small ocelli' is used when
the OCOD is greater than or equal to 1.5X
OCD, and the term 'large ocelli' when the
OCOD is less than 1.5X OCD. The width
of the apical flagellomere was measured
as the width of the base of the flagello-
mere.

Wing venation characters generally are
not stable within species and therefore
should not be used as diagnostic charac-
ters (Huddleston 1980). For example, the
position of the forewing vein m-cu
(whether it is antefurcal, interstitial, or
postfurcal) often exists in two of these
states within a species. One exception may
be the shape of the forewing second sub-
marginal cell, which appears to be stable
in some species (e.g., M. congregatus, M.
papiliovorus).

Muesebeck (1923) and Huddleston
(1980) have pointed out the importance of:
1) the presence or absence of a dorsope

and 2) whether or not the ventral borders
of the tergite are joined. Huddleston
(1980) noted that when a dorsope is pres-
ent it is distinct (Fig. 1), even if it is small,
and that the ventral borders of the first ter-
gite are never joined beneath, and usually
widely separated (Fig. 2). In specimens
where the dorsope is absent (Figs. 3, 5),
the ventral borders are always touching or
nearly touching (Figs. 4, 6), partially fused
(Fig. 7) or completely fused (Fig. 8).

The length of the ovipositor was mea-
sured from the base of the hypopygium to
the tip of the ovipositor and was ex-
pressed as relative to the length of the first
tergite. The term 'long ovipositor' was
used when the length of the ovipositor
was equal to or greater than 2.0 X the
length of the first tergite, and the term
'short ovipositor' was used when the
length of the ovipositor was less than 2.0
X the length of the first tergite.

As noted by Huddleston (1980), color
and body size are among the least stable
characters and often vary within a species.
The color of a specimen can vary depend-
ing on how it was preserved, how long it
remained in alcohol before it was pre-
pared, and exposure to light. Although the
colors 'white' and 'yellow' are used in this
text, often the specimens described as hav-
ing these colors actually have a clear in-
tegument, and it is the internal tissues that
provide the color. Overall, color is some-
what generalized in the species descrip-
tions and should not be used without oth-
er diagnostic characters.

Specimens can have a variety of meta-
somal positions, therefore the body length
is a combined measurement of the length
from the head to the end of propodeum,
added to the length from the base of the
first tergite to the end of the metasoma
(not including the ovipositor).

KEY TO COSTA RICAN METEORUS FEMALES WITHOUT A DORSOPE

1. Mandible strongly twisted (Fig. 9); propodeum rugose (Figs. 15-16) or areolate-rugose
(Fig. 17), but never carinate, with a distinct median and transverse carina creating
very large, defined areolae (Fig. 18) 2

Volume 7, Number 2, 1998

185

Figs. 1-6. 1, 3, 5. First metasonial Ifrgite, dorsal view. 1, Metcorus sp., dorsope present; 3, Mcteorus rogerblancoi,
dorsope absent, surface smooth and rounded; 5, Meteorus pseudodimidiatus, dorsope absent, surface not com-
pletely smooth and tergite flattened laterally. 2, 4, 6. First metasomal tergite, ventral view. 2, Meteorus sp.,
(dorsope present) ventral borders widely separated; 4, Meteorus rogerblancoi, (dorsope absent) ventral borders
of first tergite joined completely along basal Vi of segment and suture apparent; 6, Meteorus diwidmlus. (dorsope
absent) ventral borders touching for only short distance apically.

2(1).

Mandible moderately twisted (Fig. 10) or not twisted, broad and flat at base; propo-
deum carinate, with a distinct median and transverse carina creating very large, de-
fined areolae (Fig. 18), or rugose (Figs. 15-16) 15

Wings very dark, infused with blackish pigment, and second submarginal cell of forewing
strongly narrowed anteriorly; forewing vein 3RSa usually much shorter than r, or some-

186

Journal of Hymenoptera Research

Figs. 7-12. 7-8. Metcorus con^rc^atiiff, first metasomal tergite, ventral view (dorsope absent). 7, ventral borders
partially fused (separated basally, fused apically); 8, ventral borders fused, no indication of suture. 9-10.
Anterior view of head. 9, Mctecruf rogcrbhiiicoi, mandibles strongly twisted, with one visible tooth; 10, Mcfctinis
coffciitiis, mandibles moderately twisted, with ventral tooth partially visible. 11-12. Mesoscutum, dorsal view,
showing notauli. 11, Mctcorus ivgcrblancoi, notauli not distinct, broad, rugose, and converge posteriorly in a
rugose area; mesonotal lobes not well-defined; 12, Meteorus megalops, notauli distinct, linear, foveolate ante-
riorly, and converge posteriorly in a rugose area; mesonotal lobes well-defined.

times nearly equal; eyes small and nearly parallel; notauli distinct and linear, mesonotal

lobes well-defined (Fig. 12) 3

Wings not dark, at most with brownish pigment; second submarginal cell of forewing
not strongly narrowed anteriorly; forewing vein r usually much shorter than, or some-
times equal to, 3RSa; eyes small or large, convergent or nearly parallel; notauli distinct,
linear and mesonotal lobes well-defined (Fig. 12), or not (Fig. 11) 4

Volume 7, Number 2, 1998 187

3(2). Head height 2.0x eye height; occipital carina incomplete, widely separated medially
(Fig. 23); first tergite with costae parallel, usually yellowish in color and only slightly
darker apically, ventral borders fused (Figs. 7-8); highly gregarious (about 25-250

wasps per larva), attacking Sphingidae (Figs. 25-27) M. congregatus Muesebeck

Eyes somewhat larger, head height 1.7x eye height; occipital carina not complete but
with only a small separation medially (Fig. 21); first tergite with costae convergent
posteriorly, usually yellow basally, nearly black apically, ventral borders joined com-
pletely but not fused (Fig. 4); metasoma nearly black dorsally; solitary or gregarious,
attacking Papilionidae (1-5 wasps per larva) (Fig. 28) M. papiliovonis Zitani

4(2). Ventral borders of first tergite not joined completely along basal Vi of segment (Fig.
6); notauli distinct, linear, and mesonotal lobes well-defined (Fig. 12), or not (Fig. 11)

5

Ventral borders of first tergite joined completely along basal Vi of segment (Fig. 4)
and notauli not distinct, broad, mesonotal lobes not well-defined (Fig. 11) 7

5(4). Notauli not distinct, broad, and mesonotal lobes not well-defined (Fig. 11)

M. rubens (Nees)

Notauli distinct, linear, and mesonotal lobes well-defined (Fig. 12) 6

6(5). Ovipositor long, equal to, or greater than 2.0 X length of first tergite; malar space

about equal to basal width of mandible M. dimidiatus (Cresson)

Ovipositor short, less than 2.0x length of first tergite; malar space longer than basal
width of mandible M. pseudodimidiatus Zitani, new species

7(4). Ocelli small (OCOD equal to, or greater than 1.5X OCD) 8

Ocelli large (OCOD less than 1.5x OCD) 9

8(7). Occipital carina complete (Fig. 19), hind coxa rugulose . . . M. rogerblancoi Zitani, new

species
Occipital carina not complete, poorly defined medially (Fig. 21), hind coxa polished,
punctate M. uno Zitani, new species

9(7). Ovipositor short, less than 2.0 X length of first tergite; tarsal claws with a well-defined

lobe (Fig. 22) or without (Figs. 20, 24) 10

Ovipositor long, equal to, or greater than 2.0 x length of first tergite, and tarsal claws

with a well-defined lobe (Fig. 22) 12

10(9). Occipital carina not complete, poorly defined medially (Fig. 21); longitudinal costae

of first tergite parallel M. alejandromasisi Zitani, new species

Occipital carina complete (Fig. 19); longitudinal costae of first tergite convergent pos-
teriorly 11

11(10). Malar space length 0.5x length of mandible basally; frons with a median tubercle
anterior to median ocellus; sternaulus foveolate, not broad (Fig. 14); hind coxa ru-
gulose, with distinct but very fine carinae M. dos Zitani, new species

Malar space greater than 0.5x mandible width basally; sternaulus rugose, broad, and

long (Fig. 13); hind coxa rugulose-punctate M. laphygmae Viereck

12(9). First tergite laterally flattened (Fig. 5), and dorsal surface rugulose medially (costae

not distinctly linear); eyes not so large and protuberant laterally . . . M. sterictae Zitani,

new species

First tergite not laterally flattened (Fig. 3), with longitudinal costae distinct, parallel,

but sometimes convergent posteriorly and, eyes very large, strongly protuberant ... 13

13(12). Eyes convergent; clypeus somewhat flattened; occipital carina complete (Fig. 19); first

tergite with costae somewhat convergent posteriorly; vertex not concave behind eyes

M. desmiae Zitani, new species

Eyes nearly parallel; clypeus not flattened; occipital carina not complete, poorly de-
fined medially (Fig. 21); first tergite with costae parallel; vertex strongly concave be-
hind eyes M. camilocamargoi Zitani, new species

14(1). Mandibles moderately twisted (Fig. 10) 15

Mandibles not twisted 16

188

Journal of Hymenoptera Research

15(14). Malar space very short, 0.3 x rnandible width basally, eyes nearly touching base of

mandible; eyes strongly convergent M. coffeatus Zitani, new species

- Malar space very long, 1.1 x mandible width basally; eyes parallel . . . M. micrommattis

Zitani, new species
16(14). Propodeum rugose, scutellar disc coarsely rugose and raised into a sharp point; eyes

protuberant M. cortticulattts Zitani, new species

- Propodeum carinate, with a distinct median and transverse carina creating very large,
defined areolae (Fig. 18); scutellar disc rounded and smooth; eyes protuberant or not

17

17(16). Eyes huge, bulging, and strongly convergent; antennae with 25 flagellomeres; large

species, body length greater than 5.0 mm M. megalops Zitani, new species

- Eyes large but not huge and bulging; 19-20 flagellomeres, very small species, body
length less than 4.0 mm 18

18(17). Notauli linear and converging posteriorly in a distinct V-shape; antennae with two
white bands on F1-F3, and F9-F16; first tergite with very faint costae or none at all

M. mariamartae Zitani, new species

Notauli distinctly linear anteriorly but converging posteriorly in a rugose area (Fig.
12); antennae without white bands; first tergite with distinct lateral costae

M. yamijuanum Zitani, new species

Genus Meteorus Haliday

Meteorus Haliday 1835: 24.

Diagnosis for species lacking a dorsope:
Body color: variations of w^hite, yellow,
orange, reddish-brown, brown, dark
brown, or nearly black. Body length: 2.5-
6.0mm. Head: maxillary palpi 6-segment-
ed; labial palpi 3-segmented; antenna with
19-34 flagellomeres, antennae filiform,
apical flagellomere pointed; head 1.1-1.4X
wider than high, head height 1.3-2. OX eye
height; eyes nearly parallel, convergent, or
strongly convergent in anterior view;
mandible strongly twisted, moderately
twisted, or not twisted; occipital carina
complete or incomplete. Mesosoma: no-
tauli and sternaulus present; epicnemial
carina present. Legs: tarsal claw lobed or
simple. Wings: vein m-cu antefurcal, in-
terstitial, or postfurcal. Metasoma: ventral
borders of first tergite fused completely or
partially, joined completely, but not fused,
along basal Vi of segment, or, meeting for
only a short distance; dorsal surface of
first tergite usually with costae posterior-
ly; metasoma polished dorsally, smooth
and shining; ovipositor 1. 3-3.6 X longer

than first tergite, usually straight but
sometimes curved.

Meteorus alejandromasisi Zitani, new
species

(Fig. 21)

Holotype female. — Body color: Yellow ex-
cept antennae light brown; metanotum
brown; propodeum brown dorsally and
posteriorly; hind leg with coxa and femur
brown apically, tibia and tarsus brown;
first tergite brown apically; T 2-1-3 brown
laterally, T 4 brown. Body length: 3.2 mm.
Head: antenna with 28 flagellomeres; fla-
gellar length /width ratios as follows: Fl
= 2.5, F2 = 2.5, F3 = 2.5, F24 = 3.0, F25
= 3.0, F26 = 2.0, F27 = 2.0, F28 = 3.0;
head 1.2X wider than high, head height
1.6X eye height, eyes small but protuber-
ant, slightly convergent in anterior view;
maximum face width 1.3X minimum face
width; minimum face width 1.4x clypeus
width; malar space length 1.3X mandible
width basally; ocelli large, OCOD 0.9 X
OCD; mandible strongly twisted; occipital
carina not complete, poorly defined me-
dially (Fig. 21). Mesosoma: notauli not
distinct, broad, rugulose, and mesonotal
lobes not well-defined (as in Fig. 11); scu-

Volume 7, Number 2, 1998

189

Figs. 13-18. 13-14. Mesopleuron, showing sternaulus. 13, Meteoruti layih\igmae, stemaulus broad and rugose;
14, Meteorus yninifuanum, sternaulus foveolate but not broad. 15-18. Propodeum, dorso-posterior view. 15,
Mftforus psfiidmiiiiudintiis, showing finely rugose sculpturing; 16, Mctconis rogcrblancoi, showing rugose sculp-
turing; 17, M('f('iir».< ct^iigrcgiitiis, showing areolate-rugose sculpturing; 18, Mfffonis niegalofs, showing carinate
sculpturing. Notice a distinct median and transverse carina creating very large defined areolae.

tellar furrow with 3 carinae; sternaulus ru-
gose, long but not broad; mesopleuron
polished, shining around sternaulus; pro-
podeum areolate-rugose, median depres-
sion weakly present. Legs: hind coxa ru-
gulose; tarsal claw with a small lobe,
strongly curved. Wings: forewing length

3.0 mm; vein m-cu interstitial; vein 3RSa
1.8X length of r. Metasoma: (as in Fig. 3);
ventral borders of first tergite joined com-
pletely along basal Vi of segment (as in Fig.
4); first tergite dorsally longitudinally cos-
tate with costae parallel; ovipositor short,
1.8X longer than first tergite.

190

Journal of Hymenoptera Research

Figs. 19-24. 19, 21, 23. Posterior view of head showing occipital carina. 19, Mclcoimr )0\;crMtiiici.n, occipital
carina complete; 21, MetccrKS alejuniinviuisisi, occipital carina not complete, poorly defined medially; 23, Mc-
tcorus congrcgatus, occipital carina widely separated medially. 20, 22, 24. Tarsal claw. 20, Mctcorni. nicgdlcfv,
simple tarsal claw; 22, Mclconis cnmilocnmaigci, tarsal claw with well-defined lobe and strongly curved; 24,
Mctcorm pscKtlodiniidintm, tarsal claw with small lobe or basal tooth.

Variation of paratype females. — Brown ar-
eas vary from nearly black to nearly yel-
low; 26 flagellomeres; OCOD 1-1. 3x
OCD; forewing vein 3RSa 0.8 X length of
r, second submarginal cell slighty nar-
rowed anteriorly; median depression of
propodeum strongly present; dorsal cos-

tae of first tergite somewhat convergent
posteriorly.

Variation of paratype males. — Brown areas
vary from nearly black to nearly yellow;
head height 1.8X eye height.

Cocoon. — Pale brown to brown, some-
what wooly, ovoid; length 3.9-^.1 mm.

Volume 7, Number 2, 1998

191

1.6-1.8 mm wide medially; usually termi-
nating anteriorly with a rounded nipple-
like projection; not stuck together but of-
ten in a cluster and pointing outward
from interior of caterpillar leaf shelter and
next to dead or nearly-dead host; usually
with anterior end attached to leaf sub-
strate by a thread 0.7-2.0 mm in length
which originates just posterior to adult
emergence hole (note that thread origi-
nates from anterior end of cocoon and not
the posterior end as is usual for Meteorus).

Material examined. — Holotype female:
COSTA RICA: Guanacaste Province:
ACG, Sector Horizontes, Vado La Esper-
anza, reared from last instar larva of Sta-
phylus prob. vulgata (Moschler), 1993, Jan-
zen & Hallwachs voucher 93-SRNP-5688.
Deposited in RMSEL. Paratypes: Guana-
caste Province: 8 females, 1 male, same
data as holotype; 4 females, ACG, Sector
Santa Rosa, Area Administrativa, reared
fr. Chiomara asychis (Stoll), larva coll. 27
July 1993, wasps pupated 31 July, wasps
eclosed 9 Aug., Janzen & Hallwachs
voucher 93-SRNP-4212; 5 females, 1 male,
ACG, Sector Santa Rosa, Cerco de Piedra,
reared fr. Pyrgus syi., larva coll. 25 Sept.
1993, wasps pupated 28 Sept., wasps
eclosed 5 Oct., Janzen & Hallwachs vouch-
er 93-SRNP-6387; 11 females, 1 male,
ACG, Sector Cacao, Gongora, reared fr.
Staphyliis sp., larva coll. 21 Aug. 1994,
wasps eclosed 4 Sept., Janzen & Hall-
wachs voucher 94-SRNP-6574; 4 females,
ACG, Sector Cacao, Gongora, reared fr.
Staphyliis sp., larva coll. 21 Aug. 1994,
wasps eclosed 9 Sept., Janzen & Hall-
wachs voucher 94-SRNP-6577. Deposited
in INBio, UCR, RMSEL.

Distribution. — Known only from the ho-
lotype and paratype localities in the ACG
(Sector Santa Rosa, dry forest, and Gon-
gora, intergrade between wet and dry for-
est), Guanacaste Province.

Biology. — The rearing of this gregarious
species (4-13 wasps per larva) constitutes
the first record of Meteorus attacking Hes-
periidae. All of the parasitized larvae were

feeding on low plants in isolated dry for-
est old field succession. The type speci-
mens were all reared from last instar lar-
vae of Hesperiidae as follows: Staphylus
prob. vulgata and Staphylus sp. feeding on
Amaranthaceae, Pyrgus sp. feeding on Sida
rhombifolia L. (Malvaceae), and Chiomara
asychis, feeding on Baiiisteriopsis muricata
(Cav.) Cuatr. (Malpighiaceae). Wasps pu-
pated 3-4 days after collection of hosts
and eclosed 7-9 days later.

Comments. — Meteorus alejandromasisi
shares with M. dos and M. laphygmae the
following combination of characters:
strongly twisted mandibles, completely
joined ventral borders of the first tergite,
large ocelli, and a short ovipositor. Meteo-
rus alejandromasisi can be distinguished
from these species by an incomplete occip-
ital carina. Intraspecific variation is high in
this species.

Etymology. — This species is named in
honor of Sr. Alejandro Masis in recogni-
tion of his outstanding interest in facilitat-
ing the parataxonomists in the Research
Program of the ACG, and for being a ma-
jor supporter of the Research Program ac-
tivities and a Hesperiidae fanatic.

Meteorus camilocamargoi Zitani, new
species

(Fig. 22)

Holotype female. — Body color: orange.
Body length: 6.1 mm. Head: antenna with
35 flagellomeres; flagellar length /width
ratios as follows: Fl = 2.6, F2 = 2.6, F3 =
2.0, F31 = 1.5, F32 = 1.5, F33 = 1.5, F34 =
1.5, F35 = 2.0; head 1.3x wider than high,
head height 1.5x eye height, eyes large,
nearly parallel in anterior view; maximum
face width 1.3x minimum face width;
minimum face width = clypeus width;
malar space length = mandible width ba-
sally; ocelli large, OCOD 0.6 X OCD; man-
dible strongly twisted (as in Fig. 9); occip-
ital carina not complete, poorly defined
medially (as in Fig. 21); vertex concave be-
hind eyes. Mesosoma: notauli uncertain
(see comments section); scutellar furrow

192

Journal of Hymenoptera Research

with 3 carinae; sternaulus rugose, not
deep or broad but long; propodeum are-
olate-rugose (as in Fig. 17), median de-
pression present. Legs: hind coxa punc-
tate; tarsal claws lobed, strongly curved
(Fig. 22). Wings: forewing length 4.6 mm;
vein m-cu antefurcal; vein 3RSa 1.3x
length of r. Metasoma: (as in Fig. 3); ven-
tral borders of first tergite joined com-
pletely along basal V2 of segment (as in Fig.
4); first tergite with longitudinal costae
parallel; ovipositor long, 3.5 x longer than
first tergite.

Variation of parati/pe female. — Body
length 5.5 mm.

Variation of paratype male. — Unknown.

Cocoon. — Pale brown, ovoid; length 7.9
mm, 2.6 mm wide medially; does not ter-
minate in a nipple-like projection; formed
within the host cocoon; attached to the in-
side of the host cocoon by a thread, 0.7
mm in length, which originates from the
wasp cocoon just posterior to the adult
emergence hole (not from the posterior
apex of the cocoon as is usual for Meteor-
us).

Material examined. — Holotype female:
COSTA RICA: Guanacaste Province:
ACG, Sector Santa Rosa, Casona, reared
from Omiodes stigmosalis (Warr.) prepupa,
1978, Janzen & Hallwachs voucher 78-
SRNP-143. Deposited in RMSEL. Para-
type: Guanacaste Province: 1 female,
same data as holotype except Janzen &
Hallwachs voucher 78-SRNP-143.1. De-
posited in RMSEL. It is not known to
which specimen the cocoon belongs, de-
posited in RMSEL.

Distribution. — Known only from the
type-locality, the dry forest of the ACG,
Guanacaste Province.

Biology. — All type specimens were
reared from larvae of the pyralid, Omiodes
stigmosalis, feeding on Ficus oimlis Miq.
(Moraceae). The host larva webs and
mines through nearly ripe to fully ripe figs
and turns them brown. The host larvae
were collected 25 December 1978 and be-
came prepupae 3 days later. The wasp lar-

vae emerged from the prepupae and
formed cocoons within the host cocoon.
The wasps eclosed 18 January 1979.

Comments. — Meteorus camilocamargoi, M.
desmiae, and M. sterictae can be recognized
by the following combination of charac-
ters: strongly twisted mandibles, first ter-
gite with completely joined ventral bor-
ders, large ocelli, long ovipositor, and
lobed tarsal claws. Meteorus camilocamargoi
can be separated from these species by the
following characters: first tergite not lat-
erally flattened, parallel eyes, and a con-
cave vertex. The type specimens were
pinned through the mesonotum, therefore
most of the notauli are destroyed.

Eti/mology. — This species is named in
honor of Sr. Camilo Camargo in recogni-
tion of his extreme enthusiasm for facili-
tating the full computerization of the Re-
search Program and other programs of the
ACG, and being a major supporter of the
Research Program activities.

Meteorus coffeatus Zitani, new species

(Fig. 10)

Holotype female. — Body color: brown ex-
cept face light brown; mesosoma dark
brown except fore and mid coxae yellow;
legs brown; metasoma brown except first
tergite dark brown; basal % of T 2-1-3 yel-
low. Body length: 4.0 mm. Head: antenna
with 29 flagellomeres; flagellar length/
width ratios as follows: Fl = 3.5, F2 = 3.5,
F3 = 3.0, F25 = 1.5, F26 = 1.5, F27 = 1.5,
F28 = 1.0, F29 = 2.0; head 1.2X wider than
high, head height 1.6X eye height; eye
small, strongly convergent in anterior
view; maximum face width 1.7x mini-
mum face width; minimum face width =
clypeus width; malar space length 0.3 x
mandible width basally; ocelli small,
OCOD 2.5 X OCD; face polished, punctate;
clypeus somewhat more convex than usu-
al; antennal bases low on face; occipital ca-
rina complete; mandibles moderately
twisted, broad and flat at base (Fig. 10).
Mesosoma: notauli distinct, linear, and
mesonotal lobes well-defined (as in Fig.

Volume 7, Number 2, 1998

193

12); scutellar furrow with one median ca-
rina; sternaulus not broad or long, foveo-
late (as in Fig. 14); propodeum rugose,
median depression absent. Legs: hind
coxa pohshed, punctate; tarsal claws sim-
ple, swollen at base. Wings: fore wing
length 3.8 mm; vein m-cu interstitial; vein
3RSa 1.7x length of r. Metasoma: ventral
borders of first tergite separated basally,
joined apically; first tergite dorsally lon-
gitudinally costate; ovipositor long, 2.6 X
longer than first tergite.

Variation of paratype female. — Forewing
vein m-cu postfurcal.

Variation of parati/pe males. — T 2-1-3 al-
most completely yellow; eyes smaller,
head height 1.8X eye height.

Cocoon. — Unknown.

Material examined. — Holotype female:
COSTA RICA: San Jose Province: Zurqui
de Moravia, 1600 m, vii. 1990, Paul Han-
son. Deposited in RMSEL. Paratypes: San
Jose Province: 1 female, Zurqui de Mora-
via, 1600 m, vii 1992, P. Hanson; 3 males,
Zurqui de Moravia, 1600 m, iii 1991, P.
Hanson. Deposited in INBio, UCR,
RMSEL.

Distribution. — Known only from the ho-
lotype and paratype localities in San Jose
Province.

Biology. — Unknown.

Comments. — Meteorus coffeatus and M.
micrommatus both have moderately twist-
ed mandibles but they can easily be sep-
arated by the size and degree of conver-
gence of the eyes: Meteorus coffeatus has
large convergent eyes whereas M. microm-
matus has small parallel eyes. Meteorus cof-
featus can easily be distinguished from the
species with untwisted mandibles by the
sculpturing on the propodeum.

Etymology. — From Latin meaning "the
color of roasted coffee beans," for the
brown body color of this species.

Meteorus congregatiis Muesebeck

(Figs. 7-8, 17, 23, 25-27)

Meteorus congregatus Muesebeck 1939: 86.
Diagnosis of females. — Body color: yel-

lowish-white except antennae black; head
orange dorsally; pronotum orange dorsal-
ly; mesonotum orange; fore and mid legs
darker with tarsi brown apically; hind leg
with tibia dark brown apically, tarsus
dark brown; first tergite light brown dor-
sally; metasoma light brown dorso-medi-
ally; wings very dark, infused with black-
ish pigment. Body length: 3.8-4.1 mm.
Head: antenna with 27 flagellomeres; fla-
gellar length /width ratios as follows: Fl
= 2.5, F2 = 2.5, F3 = 3.0, F23 = 3.0, F24
= 3.0, F25 = 3.0, F26 = 2.0, F27 3.0x; head
1.1 X wider than high, head height 2.0x
eye height; eyes small, nearly parallel in
anterior view; maximum face width 1.1 X
minimum face width; minimum face
width 1.3X clypeus width; malar space
length 2.0 X mandible width basally; face,
clypeus polished, minutely punctate; ocel-
li small, OCOD 1. 8-2.0 X OCD; mandible
strongly twisted; occipital carina incom-
plete, usually, but not always widely sep-
arated medially (Fig. 23). Mesosoma: no-
tauli distinct, deep, foveolate, and meso-
notal lobes well-defined; mesonotum pol-
ished, punctate, with short carina
dorso-medially; scutellar furrow with one
median carina; mesopleuron polished,
punctate; sternaulus rugose; propodeum
areolate-rugose, somewhat coarse (Fig.
17); median depression present. Legs:
hind coxa smooth; tarsal claws simple,
swollen basally. Wings: forewing length
3.3 mm; vein m-cu interstitial, antefurcal;
vein 3RSa 0.6 x length of r; second sub-
marginal cell strongly narrowed anterior-
ly. Metasoma: (as in Fig. 3); ventral bor-
ders of first tergite partially fused (Fig. 7)
or completely fused (Fig. 8); first tergite
dorsally longitudinally costate, costae par-
allel; ovipositor short, thick at base, some-
times curved, 1.1-1.5X longer than first
tergite.

Variation of males. — 32 flagellomeres;
head height l.Sx eye height.

Cocoon.— (Figs. 25, 27) A cluster of 25-
250 pinkish-beige, wooly cocoons, all
massed together next to host caterpillar

194

Journal of Hymenoptera Research

Figs. 25-28. 25-27. Meteorus coiigregatus and last instar host, Maudtiai Acxtii, at the ACG. Janzon & Hallvvachs
voucher 95-SRNP-7538. 25, Large cocoon cluster next to host cadaver; 26, Close-up of host cadaver showing
exit holes of M. congrcgatus larvae; 27, Close-up of cocoons. Notice cocoons terminate with a distinct nipple-
like projection. 28. Meteorus papiiliovorus cocoons next to host caterpillars, Papilio nnehisiihies. at Santa Rosa dry
forest, ACG. Notice cocoons terminate with a distinct nipple-like projection. Janzen & Hallwachs voucher 90-
SRNP-2177.

and stuck to leaf substrate; each terminat-
ing with a distinct nipple-like projection.
Material examined.— COSTA RICA:
Guanacaste Province: 51 females, 34
males, ACG, Sector Puente, Est. San Cris-
tobal, 2 km W. of Dos Rios, reared from
Mandiica sexta (L.) larva on Solamim tor-
vum Sw., larva collected 4 Aug. 1995,
wasps eclosed 11-12 Aug. 1995, Janzen &
Hallwachs voucher 95-7538; 92 females,
162 males, ACG, Sector Puente, Est. San

Cristobal, 2 km W. of Dos Rios, reared
from Manduca sexta larva on Solatium tor-
viim, larva collected 4 Aug. 1995, wasps
eclosed 11-12 Aug. 1995, Janzen & Hall-
wachs voucher 95-SRNP-7539; 19 females,
7 males, ACG, [Sector El Hacha] Los Al-
mendros, reared from Uiizela japix (Cram.)
larva, Janzen & Hallwachs voucher 96-
SRNP-11208, 1996. Deposited in INBio,
UCR, RMSEL.

Distributicvi. — Recorded only from local-

Volume 7, Number 2, 1998

195

ities in wet forest of the eastern end of the
ACG, Guanacaste Province, and in Pana-
ma (Canal Zone).

Biology. — M. congregatus is a gregarious
species attacking sphingid larvae. A large
number of wasp larvae (about 25-250)
emerge from a single host larva. In Costa
Rica M. congregatus was reared from Man-
diica sexta feeding on Solamtm tonniiu (So-
lanaceae), and Unzela japix feeding on Te-
tracern vohihilis L. (Dilleniaceae). Wasp lar-
vae emerged from the last instar larva of
the host (Figs. 25-27).

Comments. — Meteorus congregatus was
described by Muesebeck (1939) from spec-
imens reared from the larva of a sphingid,
Eri)ini/is ello (L.), feeding on papaya, from
Panama (Canal Zone). The Costa Rican
specimens of M. congregatus show some
morphological variation not discussed by
Muesebeck in the original description: oc-
cipital carina with a wide separation me-
dially (Fig. 23); ventral borders of first ter-
gite partially or completely fused (Figs. 7-
8); and ovipositor sometimes curved.

Meteorus congregatus is very similar
morphologically to M. papiliovorus, and
shares with M. papiliovorus and one other
Neotropical species, M. townsendi Muese-
beck, a Brazilian species, which has not
been found to occur in Costa Rica, a nar-
rowed second submarginal cell of the fore-
wing. M. congregatus can be separated
from these species by its widely separated
occipital carina, fused ventral borders, and
parallel costae of the first tergite. M. con-
gregatus also has a cocoon that is very dif-
ferent from that of M. papiliovorus (Figs.
25, 28), and feeds on another family of
Lepidoptera.

Meteorus comiculatus Zitani, new
species

Holotype female. — Body color: body red-
dish-orange except antenna yellow basal-
ly, brown apically; legs yellow except me-
sothoracic leg with femur brown apically;
metathoracic leg with femur, tibia, and
tarsus brown apically; metasoma dark

brown except first tergite orange. Body
length: 4.5 mm. Head: antenna with 26
flagellomeres; flagellar length /width ra-
tios as follows: Fl = 5.0, F2 = 5.0, F3 =
4.0, F22 = 2.3, F23 = 2.3, F24 = 2.0, F25 =
1.5, F26 = 2.7; head 1.3X wider than high,
head height 1 .5 X eye height; eye large and
protuberant, convergent in anterior view;
maximum face width 1.4X minimum face
width; minimum face width equal to clyp-
eus width; malar space short, malar space
length 0.5 X mandible width basally; ocelli
small, OCOD 2.3 x OCD; face, clypeus
polished; mandible not twisted. Mesoso-
ma: notauli distinct, linear, foveolate, and
mesonotal lobes well-defined; mesonotum
polished except scutellar disc coarsely ru-
gose and raised in a sharp point; scutellar
furrow with a distinct median carina, two
lateral carinae weakly present; mesopleu-
ron rugose; stemaulus rugose, long and
broad; propodeum coarsely areolate-ru-
gose, median depression absent. Legs:
hind coxa polished, punctate; tarsal claws
simple (as in Fig.20). Wings: forewing
length 3.6 mm; vein m-cu antefurcal; r
0.6 X length of 3RSa; second submarginal
cell quadrate. Metasoma: (as in Fig. 3);
ventral borders of first tergite joined com-
pletely along basal Vi of segment; first ter-
gite polished with faint costae posteriorly;
ovipositor long, 2.2 x longer than first ter-
gite.

Variation of paratype females. — Body (ex-
cept legs) almost entirely dark reddish-
brown.

Variation of paratype male. — Antennae
broken; head 1.2x wider than high; head
height 1.8X eye height.

Cocoon. — Unknown.

Material examined. — Holotype female:
COSTA RICA: Puntarenas Province: San
Vito, Las Cruces, 1200 msnm, VIII-IX
1988, Coll. P. Hanson. Deposited in
RMSEL. Paratypes: Guanacaste Province:
2 females, ACG, [Sector] Santa Rosa,
Bosque Humedo-12-C, Malaise, Janzen &
Gauld, 3.viii-24.viii.l985; 1 female, ACG,
[Sector El Hacha] Los Almendros, E. Lo-

196

Journal of Hymenoptera Research

pez, 1 a 22 July 1992, L-N 334800, 369800,
INBIO CRIOOO 735444; Puntarenas Prov-
ince: 1 male, Rancho Quemado, 200 m.
Peninsula de Osa, Oct 1992, M. Segura, L-
S 292500, 511000 INBIO CRIOOO 969047; 2
females, San Vito, Estac. Biol. Las Alturas,
1500 m, vi. 1992, traps #1 & #2, Malaise,
Paul Hanson; 1 female, R.F. Golfo Dulce,
3 km SW Rincon, 10 m, vi. 1991, Paul Han-
son. San Jose Province: Zurqui de Mora-
via, 1600 m, viii. 1995, Malaise, Paul Han-
son. Deposited in INBio, UCR, RMSEL.

Distribution. — Known only from the ho-
lotype and paratype localities in Guana-
caste, Puntarenas, and San Jose Provinces.

B iology . — Unknown .

Comments. — This species has an unusual
character found in no other Meteorus spe-
cies: the scutellar disc is coarsely rugose
and raised into a sharp point. In all other
species the scutellar disc is rounded and
smooth. The combination of the form of
the scutellar disc, untwisted mandibles,
and a coarsely rugose propodeum makes
this species very easy to recognize.

Etymology. — Named for the form of the
scutellar disc, from Latin meaning "hav-
ing a horn-shaped appendage."

Meteorus desmiae Zitani, new species

Holotype female. — Body color: yellow-or-
ange except head with temples and vertex
black, antennae brown; pronotum black
dorsally; propleura mostly black; meso-
notum black; mesopleuron black anteri-
orly; propodeum dark brown dorsally;
first tergite with lateral brown spots. Body
length: 6.0 mm. Head: antenna with 31
flagellomeres; flagellar length /width ra-
tios as follows: Fl = 3.5, F2 = 3.5, F3 =
3.0, F27 = 3.0, F28 = 2.0, F29 = 2.0, F30 =
2.0, F31 = 3.0; head 1.2x wider than high,
head height 1.4x eye height; eye large and
protuberant, ventral margin of eye nearly
touching base of mandible; eyes slightly
convergent in anterior view; maximum
face width 1.6X minimum face width;
minimum face width 0.8 X clypeus width;
malar space length 0.5 X mandible width

basally; ocelli large, OCOD 0.5 X OCD;
face, clypeus punctate; clypeus somewhat
flattened; mandible strongly twisted (as in
Fig. 9); occipital carina complete (as in Fig.
19). Mesosoma: notauli not distinct, ru-
gulose, and mesonotal lobes not well-de-
fined (as in Fig. 11); mesoscutum polished,
punctate, with median carina anteriorly;
scutellar furrow with 3 distinct carinae;
mesopleuron polished, punctate; sternau-
lus rugose, very long and extending dor-
sally; propodeum rugose, with two dis-
tinct carinae posterio-ventrad, median de-
pression weakly present. Legs: hind coxa
polished, punctate; tarsal claws lobed,
strongly curved (as in Fig. 22). Wings:
forewing length 4.6 mm; vein m-cu ante-
furcal; vein 3RSa 1.8X length of r. Meta-
soma: ventral borders of first tergite joined
completely along basal V2 of segment (as
in Fig. 4); first tergite dorsally longitudi-
nally costate, costae parallel; ovipositor
long, 2.5 X longer than first tergite.

Variation of females. — Unknown.

Variation of paratype male. — Body length
5.0 mm; eyes nearly parallel, maximum
face width 1.3x minimum face width;
minimum face width equal to clypeus
width; fore wing length 3.6 mm; vein 3RSa
1.3x length of r; hind coxa rugulose.

Cocoon. — Ovoid, brown; length 5.9 mm,
2.4 mm wide medially; formed within the
remains of the larval leaf roll.

Material examined. — Holotype female:
COSTA RICA: Guanacaste Province:
[ACG, Sector Pitilla] Est. Pitilla, 700 m, 9
km S Sta Cecilia, P. Rios & C. Moraga, Oct
1990, L-N-330200, 380200, INBIO CRIOOO
398209. Deposited in RMSEL. Paratype:
COSTA RICA: Guanacaste Province: 1
male with cocoon, ACG, [Sector] Santa
Rosa, Casetilla Entrada, reared from Dt's-
mia tages (Cram.), 1983, Janzen & Hall-
wachs voucher 83-SRNP-574A. Deposited
in RMSEL.

Distribution. — Known only from the ho-
lotype and paratype localities in the ACG,
Guanacaste Province.

Biology. — The male paratype was reared

Volume 7, Number 2, 1998

197

from a pyralid, Desmia tages, feeding on
Hatnelia patens Jacq. (Rubiaceae). This py-
ralid is a leaf roller /webber. The parasit-
oid emerged from the last instar larva.

Comments. — Meteonis desmiae shares
with M. cnmilocamargoi and M. sterictae the
following characters: strongly twisted
mandibles, completely joined ventral bor-
ders of the first tergite, large ocelli, long
ovipositor, and lobed tarsal claws. Meteo-
nis desmiae can be separated from these
species by its first tergite being not later-
ally flattened, convergent eyes, and com-
plete occipital carina.

Etymology. — This species is named for
the genus of the pyralid caterpillar it at-
tacks, Destnia.

Meteonis dimidiatus (Cresson)

(Fig. 6)

Perilitus dimidiatus Cresson 1872:83.

Diagnosis of females. — Body color: dark
brown except antennae brown, orange
around eyes, face yellow, pronotum yel-
low ventrally, legs yellow except hind leg
with tibia and tarsus light brown, meta-
soma brown. Body length: 3.6 mm. Head:
antenna with 27 flagellomeres; flagellar
length /width ratios as follows: Fl = 4.0,
F2 = 3.5, F3 = 3.0, F23 = 1.5, F24 = 1.5,
F25 = 1.5, F26 = 2.0, F27 = 3.0; head 1.1 X
wider than high, head height 1.7x eye
height, eye small, convergent in anterior
view; maximum face width 1.4X mini-
mum face width; minimum face width
1.1 X clypeus width; malar space short,
malar space length = mandible width ba-
sally; ocelli small, OCOD 2.0 X OCD; man-
dible strongly twisted (as in Fig. 9); occip-
ital carina not complete. Mesosoma: no-
tauli distinct, finely areolate, and meso-
notal lobes well-defined; scutellar furrow
with one distinct median carina, 4 carinae
weakly present; mesopleuron polished,
punctate; sternaulus not deep or broad but
long; propodeum finely areolate-rugose,
with a weak carina medially (as in Fig. 15);
median depression weakly present. Legs:

hind coxa polished, punctate; tarsal claws
simple but swollen at base. Wings: fore-
wing length 3.3 mm; vein m-cu post fur-
cal; vein 3RSa 1.3 X length of r. Metasoma:
(as in Fig. 5); ventral borders of first tergite
joined for only a short distance apically
(Fig. 6); first tergite with lateral longitu-
dinal carinae, rugulose medially; oviposi-
tor long, 2.6 X longer than first tergite.

Variation of males. — Unknown.

Cocoon. — Unknown.

Material examined. — COSTA RICA:
Cuanacaste Province: 1 female, [ACG,
Sector Cacao, Estacion Cacao =] Estac.
Mengo, S.W. Volcan Cacao, 1100 m IX-X
1989; 1 female, [ACG, Sector Cacao, Esta-
cion Cacao =] Estac. Mengo, S.W. Volcan
Cacao, 1100 m, 1988-1989; Puntarenas
Province: 5 females, San Vito, Estac. Biol.
Las Alturas, 1500 m, xii, 1991, Paul Han-
son; 1 female, Estac. Biol. Las Alturas, xi
1991; 1 female, Estac. Biol. Las Alturas, i,
1992; 1 female Estac. Biol. Las Alturas, ii,
1992; San Jose Province: 1 male, Zurqui
de Moravia, vi. 1990, 1600 m, P. Hanson;
1 female, Zurqui de Moravia, 1600 m, x-
xii, 1990; 1 female, Zurqui de Moravia, iii,
1991; 2 females, Zurqui de Moravia, iii,
1992, P. Hanson. Deposited in INBio,
UCR, RMSEL.

Distribution. — In Costa Rica Known
from Guanacaste, Puntarenas, and San
Jose Provinces. It is also widely distribut-
ed in North America and recorded from
Patagonia (Shenefelt 1969).

Biology. — Unknown.

Comments. — M. dimidiatus and M. pseu-
dodimidiatus are very similar morphologi-
cally, but can be separated by the length
of the ovipositor and size of the eyes.
These are the first records of M. dimidiatus
in Costa Rica.

Meteonis dos Zitani, new species

Holotype female. — Body color: Head yel-
low, antenna black; mesosoma white ex-
cept pronotum dark brown dorsally,
mesonotum dark brown anteriorly and
laterally, mesopleuron dark brown ven-

198

Journal of Hymenoptera Research

trally, propodeum dark brown dorsally
and posteriorly, legs brown apically, hind
coxa with dark brown patches laterally;
metasoma white except first tergite dark
brown apical %, T 2+3 dark brown later-
ally. Body length: 4.6 mm. Head: antenna
with 28 flagellomeres; flagellar length/
width ratios as follows: Fl = 3.5, F2 = 3.0,
F3 = 2.5, F24 = 2.5, F25 = 2.0, F26 = 2.0,
F27 = 2.0, F28 = 2.5; head 1.2x wider than
high, head height 1.3x eye height, eyes
large and protuberant, ventral margins
nearly touching base of mandible; slightly
convergent in anterior view; maximum
face width 1.3X minimum face width;
minimum face width = clypeus width;
malar space length 0.5 X mandible width
basally; ocelli large, ocell-ocular distance
0.8 X greatest diameter of a lateral ocellus
mandible strongly twisted (as in Fig. 9)
occipital carina complete (as in Fig. 19)
frons with a median tubercle anterior to
median ocellus. Mesosoma: notauli not
distinct, broad, rugulose, and mesonotal
lobes not well-defined (as in Fig. 11); scu-
tellar furrow with 4 carinae; mesopleuron
polished, minutely punctate; stemaulus
foveolate, not broad or long; propodeum
finely areolate-rugose, median depression
weakly present. Legs; hind coxa rugulose,
with distinct but fine carinae; tarsal claw
with a very small lobe or basal tooth (as
in Fig. 24). Wings: forewing length 4.4
mm; vein m-cu post furcal; vein 3RSa 1.7x
length of r. Metasoma: (as in Fig. 3); ven-
tral borders of first tergite joined com-
pletely along basal 1/2 of segment (as in
Fig. 4); first tergite dorsally longitudinally
costate, costae convergent posteriorly; ovi-
positor short, 1.5x longer than first tergite.

Variation of females. — Unknown.

Variation of males. — Unknown.

Cocoon. — Unknown.

Material examined. — Holotype female:
COSTA RICA: San Jose Province: Zurqui
de Moravia, 1600 m. I-II 1989, P. Hanson
& 1. Gauld. Deposited in RMSEL.

Distribution. — Known only from the ho-
lotype locality in San Jose Province.

Biology. — Unknown.

Comments. — This species has an unusual
color pattern of dark brown and white
patches, similar to the color pattern of M.
uno, giving these two species a superficial
similarity. Morphologically M. dos resem-
bles M. laphygmae, as they share strongly
twisted mandibles, completely joined ven-
tral borders of the first tergite, large ocelli,
short ovipositor, and a complete occipital
carina. Meteorus dos can be further sepa-
rated from this species by its very large
eyes and median tubercle on the frons.

Etymology. — The specific epithet is an
arbitrary combination of letters.

Meteorus laphygmae Viereck

(Fig' 13)

Meteorus laphygmae Viereck, 1913, Proc. U.S.
Nat. Mus. 44560.

Diagnosis of females. — Body color: yel-
low. Body length: 3.6-5.4 mm. Head: an-
tenna with 31-34 flagellomeres; flagellar
length /width ratios as follows: Fl = 5.0,
F2 = 5.0, F3 = 2.7, F27 = 1.5, F28 = 1.5,
F29 = 1.0, F30 = 1.3, F31 = 2.0; head 1.2X
wider than high, head height 1.5X eye
height, eyes large and protuberant; nearly
parallel in anterior view; maximum face
width 1.1 X minimum face width; mini-
mum face width = clypeus width; malar
space length 0.8-1 .2 X mandible width ba-
sally; ocelli large, OCOD 1.0-1.4X OCD;
mandible strongly twisted (as in Fig. 9);
occipital carina complete (as in Fig. 19).
Mesosoma: notauli not distinct, broad, ru-
gulose, and mesonotal lobes not well-de-
fined (as in Fig. 11); scutellar furrow with
3 carinae; mesopleuron polished, minutely
punctate; stemaulus rugulose to rugose,
broad and long (as in Fig. 13); propodeum
rugose (as in Fig. 16) to areolate-rugose,
median depression present, sometimes
weak. Legs: hind coxa rugulose; tarsal
claw with a small lobe, strongly curved.
Wings: forewing length 3.2-4.7 mm; vein
m-cu antefurcal or interstitial; vein 3RSa
1. 0-0.4 X length of r. Metasoma: (as in Fig.

Volume 7, Number 2, 1998

199

3); ventral borders of first tergite joined
completely along basal Vz of segment (as
in Fig. 4); first tergite dorsally longitudi-
nally costate, costae convergent posteri-
orly; ovipositor short, sometimes thick at
base, 1.6-1.8X longer than first tergite.

Variation of males. — 34 flagellomeres;
eyes smaller, head height 1.9x eye height;
maximum face width = minimum face
width; propleura darker, propodeum
brown dorsally, hind leg with tibia and
tarsus darker, first tergite of metasoma
brown apically..

Cocoon. — No reared specimens were ex-
amined from Costa Rica.

Material examined.— COSTA RICA:
Guanacaste Province: 5 females. Hacien-
da El Vieja, Filadelfia, sugarcane, 110 m,
V 1989, M. Garcia; 3 females, 1 male,
[ACG, Sector Cacao] Volcan Cacao, Cerro
Pedregal, 1000 m, Il-IV 1989, 1. Gauld &
D. Janzen; 4 females, 3 males, Sotobosque,
W side Volcan Cacao, 1100 m, II 1989, 1.
Gauld. Deposited in INBio, UCR, RMSEL.
Limon Province: 1 female, Los Diamantes,
Guapiles, 200m, 20/ V/ 1988, col. Paul
Hanson. Deposited in INBio, UCR,
RMSEL.

Distribution. — In Costa Rica known
from Guanacaste and Limon Provinces.
Previously recorded from Venezuela, Co-
lombia, and North America, and intro-
duced into Hawai'i (Shenefelt 1969).

Biology. — Unknown.

Comments. — Meteoriis laphi/gmae shares
with M. dos and M. alejandromasisi the fol-
lowing combination of characters: strong-
ly twisted mandibles, completely joined
ventral borders of the first tergite, large
ocelli, and a short ovipositor. Meteorus
laphygmae can be separated from these
species by a complete occipital carina and
a longer malar space.

Meteonis mariamartae Zitani, new
species

Holotype female. — Body color: orange-
brown except head dark brown, antenna
brown except F1-F3 white and with white

annulus F9-F16; legs yellow; metasoma
slightly paler than mesosoma. Body
length: 2.5 mm. Head: antenna with 20
flagellomeres; flagellar length /width ra-
tios as follows: Fl = 5.0, F2 = 5.0, F3 =
4.0, F16 = 2.0, F17 = 2.0, F18 = 2.0, F19 =
2.0, F20 = 3.0; antenna slightly tapered ba-
sally; head 1.4X wider than high, head
height 1.5X eye height; eye large, conver-
gent in anterior view; maximum face
width 1.6x minimum face width; mini-
mum face width 0.7x clypeus width; ma-
lar space short, malar space length 0.6 x
mandible width basally; ocelli small,
OCOD 3.0 X OCD; mandible not twisted;
occipital carina complete (as in Fig. 19).
Mesosoma: notauli finely areolate, dis-
tinctly linear, converging posteriorly in a
distinct v-shape, and mesonotal lobes
well-defined; scutellar furrow with one
median carina; mesopleuron polished,
smooth and shining; stemaulus foveolate,
not broad (as in Fig. 14); propodeum car-
inate, with a distinct median and trans-
verse carina creating very large, defined
areolae (as in Fig. 18); median depression
absent. Legs: hind coxa rugulose; tarsal
claws simple (as in Fig. 20). Wings: fore
wing length 1.9 mm; vein m-cu antefurcal;
vein 3RSa 4.0 X length of r. Metasoma:
ventral borders of first tergite joined al-
most completely beneath, open basally for
a short distance; first tergite dorsal surface
rugulose and without any indication of
costae; ovipositor long, 3.2 x longer than
first tergite.

Variation of paratype females. — Body color
dark brown; dorsal surface of first tergite
sometimes with very faint costae laterally.

Variation of males. — Unknown.

Cocoon. — Unknown.

Material examined. — Holotype female:
COSTA RICA: Puntarenas Province:
P[arque].N[acional]. Corcovado, Est. Sir-
ena, 50 m, x-xii 1990. Deposited in
RMSEL. Paratypes: Guanacaste Province:
1 female, [ACG, Sector Cacao, Estacion
Cacao = ] Estac. Mengo, SW Volcan Cacao,
1100 m, 1988-1989. Puntarenas Province:

200

Journal of Hymenoptera Research

2 females, same data as holotype; 1 fe-
male, Rancho Quemado, 200 m Peninsula
de Osa, Die 1992, M. Segura, L-S 292500,
511000. Deposited in INBio, UCR, RMSEL.

Distribution. — Known only from the ho-
lotype and paratype localities in Guana-
caste and Puntarenas Provinces.

Biology. — Unknown.

Comments. — Meteonts mariamartae close-
ly resembles M. i/amijumium, and M. me-
galops. These species have untwisted man-
dibles and a carinate propodeum (with
large, defined areolae). Meteorus mariamar-
tae can be distinguished from these other
two species by its smaller eyes, unique no-
tauli (converging posteriorly in a distinct
V-shape), and small body size.

Etymology. — This species is named in
honor of Srta. Maria Marta Chavarria Diaz
in recognition of her unflagging support
for the concept of biodiversity conserva-
tion throughout the history of the ACG
and INBio.

Meteorus megalops Zitani, new species

(Figs. 12, 18, 20)

Holotype female. — Body color: Dark
brown except; antenna brown with white
annulus F13-F19; mesosoma with fore and
mid coxae pale yellow; metasoma with
first tergite pale yellow basally and api-
cally, tergite 2 + 3 pale yellow for basal %.
Body length: 5.4 mm. Head: antenna fili-
form with 25 flagellomeres; flagellar
length /width ratios as follows: Fl = 5.0,
F2 = 4.0, F3 = 3.0, F21 = 1.5, F22 = 1.5,
F23 = 1.5, F24 = 1.0, F25 2.0x; head 1.3x
wider than high, head height 1.4x eye
height; eye large, strongly convergent in
anterior view, bulging anteriorly and lat-
erally; maximum face width 1.7X mini-
mum face width; minimum face width
0.7x clypeus width; malar space short,
malar space length 0.5 x mandible width
basally; ocelli small, OCOD 2.0 x OCD;
mandible not twisted; occipital carina
complete (as in Fig. 19). Mesosoma: no-
tauli distinct, foveolate, and mesonotal
lobes well-defined (Fig. 12); scutellar fur-

row with a distinct median carina, 2 lat-
eral carinae weakly present; mesopleuron
smooth, polished; sternaulus foveolate (as
in Fig. 14); propodeum carinate, with a
distinct median and transverse carina cre-
ating very large, defined areolae (Fig. 18);
median depression absent. Legs: hind
coxa polished, punctate; tarsal claws sim-
ple (Fig. 20). Wings: forewing length 4.8
mm; vein m-cu antefurcal; r 0.6 X length
of 3RSa; second submarginal cell slightly
narrowed anteriorly. Metasoma: ventral
borders of first tergite separated basally,
joined apically; first tergite dorsally lon-
gitudinally costate; ovipositor long, thick
at base, 3.6 X longer than first tergite.

Variation of paratype females. — Mesopleu-
ron reddish-orange ventrally; 24 flagel-
lomeres, annulus on F9-F14;

Variation of paratype males. — 29 flagel-
lomeres; antennae without annulus, com-
pletely brown; eyes not so large and
bulging.

Cocoon. — Unknown.

Material examined. — Holotype female:
COSTA RICA: San Jose Province: Zurqui
de Moravia, 1600 m, X-Xll, 1989, col. Paul
Hanson. Deposited in RMSEL. Paratypes:
COSTA RICA: Guanacaste Province: 1 fe-
male, [ACG, Sector Pitilla] Est. Pitilla, 700
m, 9 km S. Sta. Cecilia, P.N. Guanacaste,
C. Moraga, 31 mar-15 abr 1992, L-N
330200.380200. Puntarenas Province: 3 fe-
males, 8 males, San Vito, Estac. Biol. Las
Alturas, 1500 m, vi. 1992, traps #1 & #2,
Malaise, Paul Hanson; 1 female, San Vito,
Estac. Biol. Las Alturas, 1500 m, v. 1992,
in the forest. Malaise, Paul Hanson; 1 fe-
male, San Vito, Estac. Biol. Las Alturas,
1500 m, V. 1992, forest border. Malaise,
Paul Hanson; 1 female, San Vito, Estac.
Biol. Las Alturas, 1500 m, vi. 1992, Mal-
aise, Paul Hanson; 1 female. Est. Biol. Las
Alturas, 1500 m, Coto Brus, M.A. Zum-
bado, Nov 1991, L-S-322500,591300; 1 fe-
male. Est. Biol. Las Alturas, 1500 m, Coto
Brus, F. Arayo, 23 mar a 2 may 1992, L-S-
322500,591300; 2 females. Est. La Casona,
1520 m. Res. Biol. Monteverde, Ago 1992,

Volume 7, Number 2, 1998

201

N. Obando L-N 253250,449700; San Jose
Province: 1 female, Zurqui de Moravia,
1600 m lV/1989, col. Paul Hanson. Depos-
ited in INBio, UCR, RMSEL.

Distribution. — Known only from the ho-
lotype and paratype localities in Guana-
caste, Puntarenas, and San Jose Provinces
in Costa Rica.

B iologi/. — Unknown.

Comments. — Meteorus megalops very
closely resembles M. mariamartae and M.
yamijiianum. These 3 species share in com-
mon untwisted mandibles and a carinate
propodeum (with large, defined areolae.
Fig. 18). Meteorus megalops is distinctive
because of its huge, bulging, convergent
eyes, longer antennae, and large body
size.

Etymologif. — From Greek meaning
"large eyes."

Meteorus micrommatiis Zitani, new
species

Holotype female. — Body color: head
black, antennae dark brown; mesosoma
orange-brown except propleura dark
brown; fore leg with coxa yellow, femur,
tibia and tarsus brown; middle leg with
coxa yellow, femur, tibia, tarsus brown;
hind leg brown; wings dark; metasoma
dark brown except first tergite orange ba-
sally, brown apically; tergite 2 + 3 yellow
basally. Body length: 4.6 mm. Head: an-
tennae broken, at least 22 flagellomeres;
flagellar length /width ratios as follows:
Fl = 3.0, F2 = 3.0, F3 = 4.0; head 1.2x
wider than high, head height 1.9x eye
height, eye small, nearly parallel in ante-
rior view; maximum face width 1.1 x min-
imum face width; minimum face width
1.2x clypeus width; malar space length
long, 1.1 x mandible width basally; ocelli
small, OCOD 3.3 x OCD; mandible mod-
erately twisted, broad at base (as in Fig.
10); occipital carina complete (as in Fig.
19). Mesosoma: notauli distinct, foveolate,
converging posteriorly in a depressed ru-
gose area with 4 distinct carinae, and
mesonotal lobes well-defined; scutellar

furrow with 3 carinae; sternaulus distinct
but not broad (as in Fig. 14); mesopleuron
polished; propodeum areolate-rugose, me-
dian depression absent. Legs: hind coxa
polished, punctate; tarsal claws simple.
Wings: fore wing length 3.5 mm; vein m-
cu antefurcal; vein 3RSa = length of r. Me-
tasoma: ventral borders of first tergite sep-
arated but nearly touching apically; dorsal
surface of first tergite with lateral costae,
rugulose medially; ovipositor short, 1.4X
longer than first tergite.

Variation of paratype females. — Mesosoma
brown; body entirely dark brown; 29 fla-
gellomeres.

Variation of males. — Unknown.

Cocoon. — Unknown.

Material examined. — Holotype female:
COSTA RICA: Puntarenas Province: Fca.
Cafrosa, 1300 m. Est. Las Mellizas, P. In-
ternac. La Amistad, M. Ramirez, Jul 1991,
L-S-316100, 596100, INBIO CRIOOO 624079.
Deposited in RMSEL. Paratypes: Guana-
caste Province: 1 female, [ACG, Sector Ca-
cao] Est. Cacao, 1000-1400 m, Lado SO
Vol. Cacao, C. Chaves, Ago 1991, L-N-
32300, 375700, INBIO CRIOOO 571335;
Puntarenas Province: 1 female. Est. La Ca-
sona, 1520 m. Res. Biol. Monteverde, J. A.
Ugalde, Jul 1991, L-N-253250, 449700, IN-
BIO CRIOOO 551283; 1 female. Est. Biol.
Las Alturas, 1500m, Goto Brus, F. Araya,
23 mar a 2 may 1992, L-S 322500, 591300,
INBIO CRIOOO 792767. Deposited in IN-
Bio, UCR, RMSEL.

Distribution. — Known only from the ho-
lotype and paratype localities in Guana-
caste and Puntarenas Provinces.

Biology. — Unknown.

Comments. — Meteorus micrommatus is
unusual in that although it lacks dorsopes,
the ventral borders of the first tergite are
very slightly separated. It shares with M.
coffeatus moderately twisted mandibles
but can easily be distinguished by its very
small (head height 1.9 X eye height) par-
allel eyes.

Etymology. — From Greek meaning
"small eyes."

202

Journal of Hymenoftera Research

Meteorus papiliovorus Zitani

(Fig. 28)

Meteorus papiliovorus Zitani 1997:181. See Zitani
et al. (1997) for a species diagnosis. No new
material has been examined.

Meteorus pseudodimidiatus Zitani, new
species

(Figs. 5, 15, 24)

Holotype female. — Body color: Dark
brown except yellow around ventral bor-
ders of eyes, face yellow; pronotum with
yellow spot ventrally, mesopleuron yel-
low on ventral Vz; legs light brown; meta-
soma with T 2-1-3 white on basal Vi, brown
apically; T 4, 5 brown, white posteriorly
and ventrally. Body length: 4.3 mm.
Head: antenna with 29 flagellomeres; fla-
gellar length /width ratios as follows: Fl
= 4.5, F2 = 4.0, F3 = 3.5, F25 = 2.0, F26
= 1.5, F27 = 1.5, F28 = 1.5, F29 = 2.0;
head 1.1 X wider than high, head height
1.8X eye height; eye small, convergent in
anterior view; maximum face width 1.5x
minimum face width; minimum face
width = clypeus width; malar space
length 1.5x mandible width basally; ocelli
small, OCOD 2.0 X OCD; mandible strong-
ly twisted. Mesosoma: notauli distinct
and mesonotal lobes well-defined (as in
Fig. 12); scutellar furrow with a median
carina; stemaulus foveolate, not broad or
long (as in Fig. 14); propodeum finely ru-
gose, with a weak median carina (Fig. 15),
median depression absent. Legs: hind
coxa rugulose; tarsal claw with a small
lobe or basal tooth (Fig. 24). Wings: fore-
wing length 4.1 mm; vein m-cu post fur-
cal; r 0.4X length of 3RSa. Metasoma: (Fig.
5); ventral borders of first tergite joined for
only a short distance (as in Fig. 6); first
tergite dorsally longitudinally costate; ovi-
positor short, 1.3x longer than first tergite.

Variation of paratype females. — 30 flagel-
lomeres.

Variation of males. — Unknown.

Cocoon. — Unknown.

Material examined. — Holotype female:
COSTA RICA: Heredia Province: Vara

Blanca, Finca Georgina, 2100 m, i-ii 1990,
P. Hanson. Deposited in RMSEL. Para-
types: Heredia Province: 1 female. Vara
Blanca, Finca Georgina, 2100 m, iii-iv
1990, P. Hanson; 1 female. Vara Blanca,
Finca Georgina, 2100 m, vi-viii 1990, P.
Hanson. San Jose Province: 1 female,
16km S. Empalme, 2600 m, III-IV 1989, P.
Hanson & I. Gauld; 3 females, Cerro
Muerte, 20 km S. Empalme, 2800 m, xi 88-
i 1989, P. Hanson. Deposited in INBio,
UCR, RMSEL.

Distribution. — Known only from the ho-
lotype and paratype localities in Heredia
and San Jose Provinces.

Biologif. — Based on the holotype and
paratype localities M. pseudodimidiatus oc-
curs at high elevations. The paratype rec-
ord on Cerro de la Muerte, at 2800 m, is
the highest elevation record for the known
Costa Rican Meteorus.

Comments. — M. pseudodimidiatus is
based on 7 female specimens and there is
almost no morphological variation in the
type specimens. This species is very simi-
lar to M. dimidiatus, and can be separated
by the shorter length of the ovipositor and
small size of the eyes.

Etymologif. — Derived from Greek mean-
ing "false dimidiatus" in reference to the
many morphological similarities between
this species and M. dimidiatus.

Meteorus rogerblancoi Zitani, new
species

(Figs. 3-4, 9, 11, 16, 19)

Holotype female. — Body color: Very dark
brown except antennae with white annu-
lus F23-28; face light brown, orange
around eyes; mesonotum yellow medially,
scutellar disc yellow; pronotum white
ventrally; propleura white; mesopleuron
white medially; metapleura white; fore
coxa white, leg light brown; mid coxa
white, leg light brown; hind coxa white
anteriorly, leg brown; first tergite white
basally; T 2 + 3 white medially; metasoma
white apically and ventrally. Body length:
4.3 mm. Head: antenna with 28 tlagello-

Volume 7, Number 2, 1998

203

meres; flagellar length/ width ratios as fol-
lows: Fl = 3.5, F2 = 3.5, F3 = 2.5, F24 =
1.0, F25 = 1.0, F26 = 1.0, F27 = 1.0, F28 =
1.5; head 1.2X wider than high, head
height 1.6X eye height, eye small, nearly
parallel in anterior view; maximum face
width 1.2x minimum face width; mini-
mum face width = clypeus width; malar
space length 1.5x mandible width basally;
ocelli small, OCOD 1.50 x OCD; occipital
carina complete (fig. 19); mandible strong-
ly twisted, and frons with faint horizontal
carinae, clypeus rugulose (Fig. 9). Meso-
soma: notauli not distinct, broad, rugu-
lose, and mesonotal lobes not well-defined
(Fig. 11); scutellar furrow with one distinct
median carina, 4 lateral carinae weakly
present; mesopleuron polished, punctate;
sternaulus shallow but not broad or long;
propodeum rugose (Fig. 16); median de-
pression weakly present. Legs: hind coxa
rugulose; tarsal claw with a small lobe,
strongly curved. Wings: fore wing length
4.0 mm; vein m-cu interstitial; vein 3RSa
2.0 X length of r. Metasoma: (Fig. 3); ven-
tral borders of first tergite joined com-
pletely along basal Vi of segment (Fig. 4);
first tergite with longitudinal costae con-
vergent ventrally; ovipositor short, 1.9x
longer than first tergite.

Variation of paratype females. — Head
completely yellowish; pronotum and
mesopleuron completely white; body
length 4.1-4.3 mm; 28-32 flagellomeres;
maximum face width = minimum face
width; fore wing vein m-cu postfurcal;
ovipositor 2.0 X length of first tergite.

Variation of paratype males. — Overall
body color darker; 31 flagellomeres, an-
nulus F27-F31, yellowish; head height
1.7X eye height.

Cocoon. — Unknown.

Material examined. — Holotype female:
COSTA RICA: Puntarenas Province: San
Vito, Estac. Biol. Las Alturas, 1750 m, IX-
XI 1992, P. Hanson. Deposited in RMSEL.
Paratypes: Puntarenas Province: San Vito,
Estac. Biol. Las Alturas, 1500 m, P. Han-
son; 9 females, 30 males, xi. 1991; 5 fe-

males, 16 males, xii 1991; 3 females, 20
males, i. 1992; 4 males, ii. 1992; 9 females,
21 males, forest border, v. 1992; 19 males,
in the forest v. 1992; 6 females, 35 males,
traps #1 & #2, Malaise, vi. 1992; 1 male,
vi-vii 1992; 2 female, 8 males, same data
as holotype; 5 females, 1 male, 1700 m, ii-
iv, 1993. San Jose Province: 1 female, Zur-
qui de Moravia, 1600 m, vii. 1992, P. Han-
son. Deposited in INBio, UCR, RMSEL.

Distribution. — Known only from the ho-
lotype and paratype localities at Biological
Station Las Alturas, Puntarenas Province,
and Zurqui de Moravia, San Jose Prov-
ince.

Biology. — Unknown.

Comments. — Meteorus rogerblancoi shares
with M. imo strongly twisted mandibles,
completely joined borders of the first ter-
gite, and small ocelli but can easily be sep-
arated from this species by its complete
occipital carina. This is the most numer-
ous of the species collected from Malaise
traps, with nearly 200 individuals trapped
at Biological Station Las Alturas during
1991-1993, and one individual from Zur-
qui de Moravia.

Etymologi/. — This species is named in
honor of Sr. Roger Blanco in recognition
of his tenacious and omnipresent guardi-
anship of the coordination of the ACG Re-
search Program, from its inception in 1990
to the present.

Meteorus rubens (Nees von Esenbeck)

For most of this century New World
specimens of M. rubens were treated under
the name Meteorus imlgaris (Cresson)
(Muesebeck 1923) and Meteorus leviventris
(Wesmael) (Shenefelt 1969; Marsh 1979).
The correct taxonomic status of M. rubens
was clarified by Huddleston (1980). This
species was redescribed by Huddleston
(1980) and Maeto (1990a), and the Costa
Rican specimens do not differ from these
descriptions.

Material examined.— COST A RICA:
Guanacaste Province: 5 females, 3 males,
ACG, Sector Orosi, Est. Maritza, reared

204

Journal of Hymenoptera Research

from Megalopygidae, larva coll. 10 March,
wasps eclosed 23 March 1992, Janzen &
Hallwachs 92-SRNP-549. Deposited in IN-
Bio, UCR, RMSEL.

Cocoon. — Brown, wooly, ovoid; length
3.9^.7 mm, 1.7-1.9 mm wide medially; in
mass of 30 (not all adults eclosed) cocoons,
most terminating in a nipple-like projec-
tion.

Distribution. — In Costa Rica known only
from the ACG, Guanacaste Province.

Biology. — 30 wasp larvae emerged from
a last instar megalopygid larva feeding on
Ardisia revoluta Kunth (Myrsinaceae). The
host larva was collected on 10 March 1992,
the wasps pupated on 19 March, and the
wasps eclosed 23 March 1992 (only 8
adults eclosed).

Comments. — Meteorus rubens is a wide-
spread species, Holarctic in distribution,
and also recorded from Mexico and Brazil
(Shenefelt 1969).

Meteorus rubens shares with M. dimidia-
tus and M. pseudodimidiatus strongly twist-
ed mandibles and ventral borders of first
tergite not joined completely. Meteorus
rubens can easily be separated from these
species by its indistinct and broad notauli,
and poorly defined mesonotal lobes. This
is the first record of any species of Meteor-
us being reared from a megalopygid.

Meteorus sterictae Zitani, new species

Holotype female. — Body color: yellow ex-
cept antennae brown; propodeum brown
dorso-medially; hind leg with femur
brown apically, tibia and tarsus brown;
first tergite dark brown. Body length: 4.6
mm. Head: antenna with 28 flagellomeres;
flagellar length /width ratios as follows:
Fl = 3.6, F2 = 3.2, F3 = 2.3, F24 = 3.0,
F25 = 2.0, F26 = 3.0, F27 = 2.0, F28 3.0;
head 1.1 X wider than high, head height
1.5X eye height; eyes large, nearly parallel
in anterior view; maximum face width
1.3X minimum face width; minimum face
width 1.1 X clypeus width; malar space
length 0.7X mandible width basally; ocelli
large, OCOD = OCD; face punctate, with

faint transverse carinae near base of anten-
nae; clypeus rugulose; mandible strongly
twisted (as in Fig. 9); occipital carina com-
plete (as in Fig. 19). Mesosoma: notauli
not distinct, broad, rugulose, and meso-
notal lobes not well-defined (as in Fig. 11),
mesoscutum polished, punctate laterally
scutellar furrow with 3 distinct carinae,
mesopleuron polished, punctate; sternau-
lus rugose, broad and long (as in Fig. 13);
propodeum areolate-rugose, median de-
pression present. Legs: hind coxa rugu-
lose; tarsal claw lobed, strongly curved (as
in Fig. 22). Wings: forewing length 3.4
mm; vein m-cu antefurcal; 3RSa 1.6x
length of r; Metasoma: (as in Fig. 3); ven-
tral borders joined completely along basal
Vi of segment (as in Fig. 4); first tergite
with dorsal, lateral, longitudinal costae
which converge posteriorly, medially ru-
gulose; ovipositor long, 3.0 x longer than
first tergite.

Variation of paratype female. — yellow ar-
eas may be orange; body length 3.9 mm;
forewing length 3.3 mm; forewing vein
3RSa 2.0 X length of vein r; ovipositor 3.4 X
longer than first tergite

Variation of paratype males. — body color
as in females and propodeum almost com-
pletely brown; 29 flagellomeres; body
length 3.8 mm; head height 1.8 X eye
height; malar space length 1.5x basal
width of mandible.

Cocoon. — Brown, ovoid; length 4.6 mm,
1.9 mm wide medially; apparently not at-
tached to substrate.

Material examined. — Holotype female:
COSTA RICA: Guanacaste Province:
ACG, [Sector] Santa Rosa, dry forest,
reared from Stericta albifasciata (Druce) lar-
va, 1996, Janzen & Hallwachs voucher 96-
SRNP-9074. Deposited in RMSEL. Para-
types: Guanacaste Province: 1 male, same
data as holotype; 1 female, same data as
holotype except Janzen & Hallwachs
voucher 96-SRNP-9103; 1 male with co-
coon, same data as holotype except Janzen
& Hallwachs voucher 96-SRNP-9102. De-
posited in RMSEL.

Volume 7, Number 2, 1998

205

Distribution. — Known only from the ho-
lotype and paratype localities in the dry
forest of the ACG, Guanacaste Province,
Costa Rica.

Biolog}/. — Meteorus sterictae attacks a py-
ralid, Sterictn albifnsciata, which is a gre-
garious webber on Ocotea veraguensis
(Meisn.) Mez in the Santa Rosa dry forest.
Meteorus sterictae is solitary; each wasp
emerged from a single last instar larva.

Comments. — Meteorus sterictae resembles
M. camilocamargoi and M. desmiae, by shar-
ing the following characters: strongly
twisted mandibles, completely joined ven-
tral borders of the first tergite, large ocelli,
long ovipositor, and lobed tarsal claws.
However, M. sterictae can be distinguished
from these two species by its laterally flat-
tened first tergite, rugulose sculpturing on
the dorsal surface of the first tergite, and
smaller eyes. The rugulose sculpturing
may give the false appearance of dor-
sopes.

Etymology. — This species is named after
the genus of the pyralid caterpillar it at-
tacks, Stericta.

Meteorus uno Zitani, new species

Holoti/pe female. — Body color: dark
brown except head orange, area between
ocelli black; pronotum orange dorsally,
white ventrally; mesonotum orange; pro-
podeum white laterally and posteriorly;
fore and mid legs pale yellow; first tergite
white basally; metasoma white laterally
and ventrally; wings dark, infused with
brown pigment. Body length: 4.3 mm.
Head: antenna with 28 flagellomeres; fla-
gellar length /width ratios as follows: Fl
= 2.7, F2 = 2.3, F3 = 2.0, F24 = 1.5, F25
= 1.5, F26 = 3.0, F27 = 2.0, F28 = 3.0;
head 1.2x wider than high, head height
1.6X eye height; eye small but protuber-
ant, nearly parallel in anterior view; max-
imum face width 1.2x minimum face
width; minimum face width 1.5X clypeus
width; malar space length ].7x mandible
width basally; ocelli small, OCOD 1.5X
OCD; mandible strongly twisted (as in

Fig. 9); occipital carina not complete, poor-
ly defined medially (as in Fig. 21); vertex,
in dorsal view, descending vertically be-
hind lateral ocelli. Mesosoma: notauli ru-
gulose, not distinct, and mesonotal lobes
not well-defined; scutellar furrow with
one median carina; mesopleuron polished,
punctate; sternaulus rugulose, broad but
not long; propodeum areolate-rugose, me-
dian depression absent. Legs: hind coxa
polished, punctate; tarsal claw with a
small lobe or basal tooth, strongly curved.
Wings: forewing length 4.0 mm; vein m-
cu post furcal; r 0.6x length of 3RSa. Me-
tasoma: (as in Fig. 3); ventral borders of
first tergite joined completely along basal
Vi of segment (as in Fig. 4); first tergite
dorsally longitudinally costate, costae
slightly convergent posteriorly; ovipositor
short, thick at base, 1.7x longer than first
tergite.

Variation of females and males. — Un-
known.

Cocoon. — Unknown.

Material examined. — Holotype female:
COSTA RICA: Puntarenas Province: San
Vito, Las Cruces, 1200msnm, VIII-IX 1988,
Coll. P. Hanson. Deposited in RMSEL.

Distribution. — Known only from the ho-
lotype locality in Puntarenas Province.

Biolog}/. — Unknown.

Comments. — Meteorus uno shares with
M. rogerhlancoi strongly twisted mandi-
bles, completely joined borders of the first
tergite, and small ocelli, but can easily be
separated from this species by its incom-
plete occipital carina.

Etymology. — The specific epithet is an
arbitrary combination of letters.

Meteorus yamijuanum Zitani, new
species

(Fig. 14)

Holotype female. — Body color: body dark
brown except antenna light brown; pro-
notum yellow anteriorly; propleuron yel-
low; legs yellow except prothoracic tarsus
light brown; mesothoracic tibia and tarsus
light brown; metathoracic tibia and tarsus

206

Journal of Hymenoptera Research

light brown; metasoma brown except first
tergite yellow basally and apically, tergite
2 + 3 yellow basally. Body length: 3.5 mm.
Head: antenna with 19 flagellomeres; fla-
gellar length /width rahos as follows: Fl
= 4.5, F2 = 3.5, F3 = 3.0, F15 = 2.0, F16
= 1.5, F17 = 1.5, F18 = 1.5, F19 = 3.0;
head 1.1 X wider than high, head height
1.7x eye height; eye small, not protuber-
ant, strongly convergent in anterior view;
maximum face width 1.8X minimum face
width; minimum face width 0.6 X clypeus
width; malar space short, malar space
length 0.5 X mandible width basally; ocelli
small, OCOD 2.3 x OCD; face, clypeus
polished, punctate; mandible not twisted;
occipital carina complete (as in Fig. 19).
Mesosoma notauli distinct, foveolate, and
mesonotal lobes well-defined (as in Fig.
12); mesoscutum polished; scutellar fur-
row with a median carina; mesopleuron
polished; sternaulus foveolate, not broad
(Fig. 14); propodeum carinate, with a dis-
tinct median and transverse carina creat-
ing very large defined areolae (as in Fig.
18), median depression absent. Legs: hind
coxa polished; tarsal claws simple. Wings:
forewing length 3.1 mm; vein m-cu ante-
furcal; r 0.8 X length of 3RSa; second sub-
marginal cell quadrate. Metasoma: ventral
borders of first tergite separated basally,
joined apically; first tergite polished with
costae posteriorly; ovipositor 2.6 X longer
than first tergite.

Variation of paratype females. — Body
length 2.9-3.5 mm; 18 flagellomeres; ovi-
positor 2.6-3.0 X length of first tergite.

Variation of males. — Unknown.

Cocoon. — Unknown.

Material examined. — Holotype female:
COSTA RICA: San Jose Province: Zurqui
de Moravia, 1600 m, vii 1990, Col Paul
Hanson. Deposited in RMSEL. Paratypes:
Puntarenas Province: 10 females, San
Vito, Estac. Biol. Las Alturas, 1500 m, vi
1992, traps #1 & #2, Malaise, P. Hanson; 1
female, San Vito, Estac. Biol. Las Alturas,
1500 m, V 1992, forest border. Malaise, col.
P. Hanson; 5 females, San Vito, Estac. Biol.

Las Alturas, 1500 m, v 1992, in the forest,
Malaise, col. P. Hanson; San Jose Prov-
ince: 1 female, Zurqui de Moravia, 1600
m, vii 1990, col. Paul Hanson. Deposited
in INBio, UCR, RMSEL.

Distribution. — Known only from the ho-
lotype and paratype localities in Puntar-
enas and San Jose Provinces.

Biology. — Unknown.

Comments. — Meteorus yamijuanum close-
ly resembles M. mariamartae, and M. me-
galops. These species have untwisted man-
dibles and a carinate propodeum (with
large, defined areolae). Meteorus yamijuan-
um can be distinguished from these other
two species by its smaller eyes and the
condition of the notauli.

Etymology. — This species is named in
honor of Yamilet Acosa and Juan Acosa,
in recognition of their hospitality as care-
takers of Pitilla Biological Station, ACG,
Guanacaste Province in 1995 and 1996.

DISCUSSION

A close examination of M. congregatus
revises our concept of Meteorus. ESEM im-
ages show that the ventral borders of the
first metasomal tergite are partially fused
(Fig. 7) or completely fused (Fig. 8). Pre-
viously the ventral borders of the first ter-
gite in Meteorus have been known to be
completely joined with a suture clearly
visible (Muesebeck 1923; Huddleston
1980) (as in Fig. 4). This is the first docu-
mentation of the total fusion of the ventral
borders of the first metasomal tergite in
Meteorus. Also, M. congregatus is unusual
in that this character varies within the spe-
cies (Figs. 7-8).

The biological information for M. con-
gregatus, M. alejandromasisi, and M. rubens
is also of interest. Most meteorines are sol-
itary (Shaw 1995). Meteorus congregatus is
unusual in that it is highly gregarious (25-
250 individuals per host larva). Meteorus
alejandromasisi is also gregarious (4-13 in-
dividuals per host larva). These rearings
provide the first records of Meteorus at-
tacking the sphingid genera Manduca and

Volume 7, Number 2, 1998

207

Unzela, and the family Hesperiidae. Along
with M. papiliovorus, M. cotigregatus and
M. alejandromasisi suspend their cocoons
from very short threads.

The new biological information present-
ed here, combined with what is known
about M. papiliovorus, provides a basis for
speculation about a possible phylogeny of
Meteorus. Species of Meteonis attacking
concealed coleopteran larvae form unsus-
pended cocoons in the protected environ-
ment of the host, and these species have
been hypothesized to be relatively primi-
tive (Maeto 1990b). Species that suspend
their cocoons also attack exposed hosts,
and given that this suspended cocoon
makes the parasitoid less accessible to po-
tential enemies (Shaw and Huddleston
1991) it can be regarded as an adaptation
to attacking exposed hosts. It follows that
species that form a suspended cocoon are
derived with respect to the more primitive
species that form unsuspended cocoons in
the host's pupation chamber.

Multiple egg-laying gregarious parasit-
ism probably evolved from solitary para-
sitism in the Hymenoptera (Hanson and
Gauld 1995). Gregariousness in Meteorus is
presumed to be derived from solitary par-
asitism. Gregarious species such as M.
papiliovorus, M. congregatus, and M. alejan-
dromasisi may have secondarily lost their
longer cocoon threads. In addition to the
short cocoon threads, these three species
have cocoons that terminate with a dis-
tinct nipple-like projection. Based on ob-
servations of photographs of a highly gre-
garious African species that forms a very
organized cocoon mass, that species also
has cocoons that terminate in a nipple-like
projection. These gregarious species with
this unique cocoon shape are hypothe-
sized to be derived, and probably form a
monophyletic group.

Of the 4 new species with host associa-
tions (M. desmiae, M. camilocamargoi, M.
alejandromasisi, and M. sterictae), only one
individual (a female of M. desmiae) was
found in the examined Costa Rican Mal-

aise trap material. This suggests that there
may be quite a few more species of Meteo-
rus in Costa Rica, and that host rearing, or
some other method of trapping, will be
necessary to find them.

ACKNOWLEDGMENTS

We would like to thank the following individuals
and institutions: The curators of the institutions who
lent specimens; Teresa Williams, Western Research
Institute, for assistance with ESEM images; Jeffrey A.
Lockwood, Nancy L. Stanton and R. Greg Thorn for
helpful comments on the manuscript; The Area de
Conservacion Guanacaste for support of much of the
rearing and Malaise trapping; INBio and the National
System of Conservation Areas (SINAC) for the u.se of
their biological resources; the Norwegian Agency for
International Development (NORAD) for financing
the publication. Special thanks to Paul Hanson for so
many years of maintaining Malaise traps and sorting
specimens in Costa Rica.

The following provided funding for this research:
University of Wyoming Office of Research; UW Col-
lege of Agriculture and Office of Research; UW De-
partment of Plant, Soil, and Insect Sciences; UW In-
sect Museum; UW Graduate School; UW McNair
Scholars Program; UW Program for the Environment
and Nahjral Resources; NSF-BSR 90-24770 and DEB
94-00829 grants to D.H. Janzen.

Figure credits: 1-24, N. Zitani, S. Shaw, and T. Wil-
liams; 25-28, D.H. Janzen.

LITERATURE CITED

Achterberg, C. van. 1979. A revision of the subfamily
Zelinae auct. (Hymenoptera: Braconidae).
Tijdschrift voor Entomologie 122:241—479.

Cresson, E. T. 1872. Descriptions of North American
Hymenoptera, No. 3. Canadian Entonicilcf;ist. 4:81-
84.

Haliday, A. H. 1835. Essay on parasitic Hymenoptera
of the Icltncumoncf Adsciti. Entomological Magazine
3:20-45.

Hanson, P. E., and Gauld, I. D., eds. 1995. The Hy-
inenoptera of Costa Rica. Oxford University Press,
New York. 893 pp.

Harris, R. A. 1979. A glossary of surface sculpturing.
Occasional Papers in Entomolog]/ 28:1-31.

Huddleston, T. 1980. A revision of the western Pale-
arctic species of the genus Meteonis (Hymenop-
tera: Braconidae). Bulletin of the British Museum
(Natural History): Entomohgi/ 41:1-58.

Huddleston, T. 1983. Meteorus (Hymenoptera: Bra-
conidae) of Australia and New Guinea. Si/sfi'm-
atic Entomohgi/ 8: 393^20.

Maeto, K. 1989. Systematic studies on the tribe Me-
teorini (Hymenoptera, Braconidae) from Japan.

208

Journal of Hymenqptera Research

V. The pulchricoriiK group of the genus Meteorus
(1). Japanese journal of Entcmwiogy 57:581-595.

Maeto, K. 1990a. Systematic studies on the tribe Me-
teorini (Hymenoptera, Braconidae) from Japan.
VII. The groups of Meteorus letericus and M. rub-
ens. Japanese Journal of Entomology 58:81-94.

Maeto, K. 1990b. Phylogenetic relationships and host
associations of the subfamily Meteorinae Cresson
(Hymenoptera, Braconidae). Japmnese Journal of
Entomology 58:383-396.

Marsh, P. M. 1979. Family Braconidae. Pp. 144-313,
In: Krombein, K.V., P. D. Hurd Jr., D. R. Smith,
and B. D. Burks |eds.]. Catalog of Hymenoptera in
America North of Mexico, Smithsonian Institution
Press, Washington, D.C.

Marsh, P. M., S. R. Shaw, and R. A. Wharton. 1987.
An identification manual for the North American
genera of the Family Braconidae (Hymenoptera).
Memoirs of the Entonu^logical Society of Washington
13:1-98.

Muesebeck, C. F. W. 1923. A revision of the North
American species of ichneumon-flies belonging
to the genus Meteorus Haliday. Proceedings of the
United States National Museum 63: 1^44.

Muesebeck, C. F. W. 1939. Five new species of Mefco-
rus (Hymenoptera: Braconidae). Proceedings of the
Entomological Society of Washington 41:83-87.

Nixon. G. E. J. 1943. A synopsis of the African species
of Meteorus (Hymenoptera: Braconidae). Bulletin
of Entomological Rescarcli 34:53-64.

Sharkey, M. J. 1988. A taxonomic revision of Alaba-
grus (Hymenoptera: Braconidae). Bulletin of the
British Museum (Natural History): Entomology 57:
311-437.

Shaw, M. R., and Huddleston, T. 1991 Classification
and biology of braconid wasps (Hymenoptera:

Braconidae). Handbooks for the Identification of Brit-
ish Insects 7:1-126.

Shaw, S. R. 1985. A phylogenetic studv of the subfam-
ilies Meteorinae and Euphorinae (Hymenoptera:
Braconidae). Entomography 3:277-370.

Shaw, S. R. 1988. Euphorine phylogeny: the evolution
of diversity in host-utilization by parasitoid
wasps (Hymenoptera: Braconidae). Ecological En-
tomology 13:323-335.

Shaw, S. R. 1995. The Braconidae, pp. 431-163. In:
Hanson, P. and Gauld, I. (eds.). The Hymenoptera
of Costa Rica. Oxford University Press, New York.
893 pp.

Shaw, S. R. 1997. Subfamily Meteorinae, pp. 326-330.
In: Wharton, R. A., P. M. Marsh and M. J. Shar-
key, eds. Manual of the New World Genera of the
Family Braconidae (Hymenoptera). Special Publi-
cation of the International Society of Hymenop-
terists. No. 1, 439 pp.

Shenefelt R. D. 1969. Hymenopterorum Catalogus,
Braconidae 1 Hybrizoninae, Euphorinae, Cos-
mophorinae, Neoneurinae, Macrocentrinae, W.
Junk, The Hague. 306 pp.

Viereck, H. L. 1913. Descriptions of ten new genera
and twenty-three new species of ichneumon-
tlies. Proc. U.S. Natn. Mus. 44:555-568.

Wharton, R. A., P. M. Marsh and M. J. Sharkey, eds.
1997. Manual of the Nezv World Genera of the Family
Braconidae (Hymenoptera). Special Publication of
the International Society of Hymenopterists, No.
1, 439 pp.

Zitani, N. M., S. R. Shaw, and D. H. Janzen. 1997.
Description and biology of a new species of Me-
teorus Haliday (Hymenoptera: Braconidae, Me-
teorinae) from Costa Rica, parasitizing larvae of
Papilio and Paruies (Lepidoptera: Papilionidae).
Journal of Hynwnoplera Research 6:178-185.

J. HYM. RES.
Vol. 7(2), 1998, pp. 209-256

The Species of Asaphes Walker from America North of Mexico, with

Remarks on Extralimital Distributions and Taxa (Hymenoptera:

Chalcidoidea, Pteromalidae)

Gary A. P. Gibson and Veli Vikberg

(GAPG) Agriculture and Agri-Food Canada, Eastern Cereal and Oilseed Research Centre,

Biological Resources Program, K. W. Neatby Building, Ottawa, Ontario, Canada KIA OC6;

(W) Liinalammintie 11 as. 6, FIN-14200 Turenki, Finland

Abstract. — The species of Asaphes Walker (Pteromalidae: Asaphinae) are reviewed for the world
and revised for America north of Mexico. Six species are recognized as valid in the region: Asaphes
brevipetiolattis n. sp. (Canada, USA, Finland), A. californkus Girault, A. hirsutus n. sp. (Canada,
USA, Mexico, Austria, Czech Republic, Finland, Greenland, Norway, Russia, Sweden), A. petiolatus
Zetterstedt, A. suspensus (Nees), and A. indgaris Walker. Asaphes petiolatus (revised status) is
removed from synonymy under A. vulgaris. Asaphes indicus (Bhatnagar), described from India,
is removed from synonymy under A. vulgaris and along with A. nifipes Brues, A. lucens (Pro-
vancher), A. fletcheri (Crawford), and A. americanus Girault is synonymized under A. suspensus
(new synonymies). A lectotype is designated for A. americanus. Asaphes hiiebrichi (Brethes) and
A. bonariensis (Brethes), described from Argentina and previously synonymized under A. fletcheri
and A. lucens, respectively, are treated as nomina inqiiirenda. A key is given to distinguish males
and females of the species in the region; diagnostic features are illustrated using scanning electron
micrographs. Distribution and host data are summarized for each species in the Nearctic region,
including records of A. brevipetiolatus parasitizing Choristoneura fumiferana (Clemens) (spruce bud-
worm) and Neodipriou abietis (Harris) (balsam fir sawfly), likely as a hyperparasite. Remarks are
also given on extralimital distributions of world species and generic composition of the subfamily,
including the existence of an undescribed genus of Asaphinae in the Neotropical region, the
likelihood of at least three undescribed species of Asaphes in regions other than the Nearctic, and
the likelihood that most or all records of A. suspensus and A. vulgaris from the Neotropical region,
and of A. vulgaris from at least the Afrotropical region of Africa, are misidentifications.

Species of Asaphes Walker are known might prove to be a synonym of A. s!<s-
from all continents except Antarctica and p^ensus and Peck (1951, 1963), Graham
usually are considered to be exclusively (1969), and Burks (1979) all questioned the
hyperparasites of aphids. Graham (1969) presence of A. viil;^aris in North America,
revised the western European species and The only other currently recognized valid
Kamijo and Takada (1973) revised the Jap- name of Asaphes is A. aphidi (Risbec),
anese species. Two species were recog- which was described from Madagascar by
nized from each area, A. suspensus (Nees) Risbec (1959). Though Boucek (1976) clar-
and A. vulgaris Walker from western Eu- ified the generic placement of this species
rope, and A. suspensus and A. pubescens he stated that further study was required
Kamijo & Takada from Japan. Burks to determine whether it was a good spe-
(1979) catalogued four species from Amer- cies or a synonym of one of what was then
ica north of Mexico, A. californicus Girault, thought to be two cosmopolitan species,
A. lucens (Provancher), A. rufipes Brues, A. suspensus and A. vulgaris. All other
and A. vulgaris, though Graham (1969) names described from other regions, in-
had previously suggested that A. lucens eluding India and South America, were

210

Journal of Hymenoptera Research

synonymized under A. suspensus or A. vul-
garis. The purpose of this study was to
clarify the species composition and host
relationships of Asaphes in America north
of Mexico. To do so, other genera of Asa-
phinae and Asapihes from other regions
were examined and this has resulted in re-
marks being included on extralimital dis-
tributions and taxa. The discovery that A.
petiolatus was incorrectly synonymized
under A. vulgaris and that other species in
Europe, North America, and elsewhere ex-
ist that are similar to A. vulgaris or A. sus-
pensus suggest that all currently accepted
synonyms of these names should be re-
examined.

Unlike in most pteromalids, males of
most species of Asaphes are more readily
identified to species than are females, par-
ticularly by differences in the antennal
scape. Also important for differentiating
species, in either sex, are setal pattern of
the forewing (setation of basal cell and
structure of speculum), presence or ab-
sence of setae on the metapleuron, sculp-
ture of the frenum, and color pattern of
the legs. Although females of A. brevipe-
tiolatus n. sp. are distinguished by head
structure, females of all the other species
can only be identified correctly by using
combinations of the above features be-
cause of overlap in character states. Fur-
ther, different species, or even opposite
sexes of different species, can be collected
or reared at the same place, time, and even
from the same host. Consequently, it is
easy to misidentify some specimens, par-
ticularly females or poorly preserved and
mounted individuals.

MATERIALS AND METHODS

Terms and abreviations used for struc-
tures follow Gibson (1997), with the fol-
lowing clarifications: 'scutellum' is used
for that part of the scutellum anterior to
the frenum (Fig. 50); the 'disc' of the fore-
wing is the entire membranous region be-
yond the basal setal line (Fig. 68); and the
'funicle' is composed of all flagellar seg-

ments basal to the club (Fig. 19), including
the basal-most ring-like segment, which is
designated as fu, (Figs. 19, 20). Head
width is maximum width measured in
dorsal view; head height is maximum
height excluding the mandibles measured
in lateral view; measurements of eye
length and malar space are maximum
lengths in lateral view with both end-
points equally in focus; petiole width is
maximum width behind the anterior
flange and petiole length is maximum
length from the anterior flange to the
abruptly declined posterior edge. In order
to obtain the accurate measurements and
observations of structure and sculpture
necessary to correctly identify specimens,
glare from incandescent light sources
must be reduced by using flourescent light
or by placing some light-diffusing mate-
rial, such as a piece of translucent tracing
acetate, close to the specimen (see Goulet
and Mason, 1993: 60). Color of the coxae
is not included as part of the leg color pat-
tern because the coxae can be yellowish or
brownish in teneral specimens. However,
color of the trochanter is important and
sometimes differs from the trochantellus
so that it is important that these two struc-
tures be differentiated.

Diagnoses of females and males are
based on Nearctic specimens, with addi-
tional variability based on specimens from
other regions included in brackets or dis-
cussed under 'Remarks'. The distribution
listed and mapped for each species is
based only on specimens examined, not
literature records. Exact locality and other
label and museum data are given only for
the two newly described species and the
two relatively uncommon species in North
America, A. petiolatus and A. vulgaris.
However, in order to facilitate future stud-
ies in other regions, all countries from
which extralimital specimens were seen
are listed under 'Distribution', along with
acronyms of collections containing the
specimens. Extralimital distributions list-
ed for A. petiolatus, A. suspe)isHS, and A.

Volume 7, Number 2, 1998

211

vulgaris are not exhaustive because no at-
tempt was made to examine collections
extensively in other than the Nearctic re-
gion. Unless stated otherv^ise, listed spec-
imens, including paratypes of the two
new species, are in the CNCI. Acronyms
for the museums from which material for
this study was examined are listed in ac-
knowledgments. Label collection data has
been standardized to a single format, with
any additional explanatory data given in
brackets. Some records from Canada have
an F.I.S. number, which refers to the For-
est Insect Survey of the Canadian Forest
Service; localities from Finland often have
two sets of numbers separated by a colon,
which is a uniform grid reference (Grid
27°E) explained in Heikinheimo and Raa-
tikainen (1981). LocaHty records from
Canada are listed by province from west
to east and alphabetically by state for the
USA. Locality records for Finland, Nor-
way and Sweden are ordered by biologi-
cal province listed south to north and west
to east; abbreviations used on the labels
for each province are included in brackets.
The numerical codes included as part of
the label data of specimens reared in Swe-
den by Gardenfors are partly explained in
Gardenfors (1986: 21).

Aphidae (Homoptera) nomenclature is
based on Eastop and Hille Ris Lambers
(1976). Invalid combinations as given on
labels of paratypic material are not itali-
cized when given in association with the
valid combination. Mackauer (1968) is
used for nomenclature of Aphidiinae (Hy-
menoptera: Braconidae). The publications
of A.A. Girault are numbered according to
the bibliography of Dahms (1978).

Specimens for scanning electron micros-
copy (SEM) were prepared following Bolte
(1996). The SEM micrograph negatives
were scanned into a computer with a
35mm scanner, digitized, enhanced, and
the final plates compiled and labelled us-
ing Adobe Photoshop®. Distribution
maps were generated using Quickmap ®.
Only those localities whose position could

be determined unequivocally were
mapped so that the maps generally are
less comprehensive than the listed re-
cords.

ASAPHINAE

Pteromalidae, Miscogasterinae, Asaphini Ash-
mead 1904: 328; Peck 1951: 536; Peck 1963:
601; Peck et al, 1964: 36.

Pteromalidae, Asaphinae; Graham 1969: 77;
Burks 1979: 785; De Santis 1979: 125; Dzhan-
okmen 1987: 112; Boucek 1988: 343.

Diagnosis. — Head with occipital carina
(Figs. 13-17) and genal carina (Figs. 8, 10,
12); left mandible bidentate and right
mandible tridentate (Fig. 18) (except Aii-
sasaphes). Antenna inserted conspicuously
below middle of face, dorsal margin of to-
rulus slightly above to distinctly below
lower orbit (Figs. 1, 3, 5, 7, 9, 11) (except
Ausasaphes), 13-segmented with 1-3 basal
flagellar segments ring-like and /or with-
out longitudinal sensilla (Figs. 19-30). Pro-
notum in dorsal view transverse-rectan-
gular and convexly rounded into neck
(shape not evident in Figs. 43-48 because
of view). Mesoscutum (Figs. 43-48) with
complete, sulcate notauli; axillae not ad-
vanced anterior to base of scutellum. Fore-
wing of fully winged individuals (some
Ausasaphes brachypterous) with marginal
and submarginal veins separated by very
short hyaline region, with relatively long
sHgmal and postmarginal veins, and with
marginal vein at most 2.4 times as long as
stigmal vein. Metasoma with transverse to
elongate, tubular, sculptured petiole (Figs.
43--18).

Asaphes Walker

Asaphcs Walker 1834: 151. Type species: Asaphes
vulgaris Walker; by monotypy.

Isocratus Forster 1856: 53, 58. Unjustified re-
placement name according to Gahan and Pa-
gan 1923: 18; incorrectly considered as pre-
occupied by Asaphus Brongniart.

Notopodwn Dahlbom 1857: 295. Type species:
Asaphes vulgaris Walker; subsequently des-
ignated by Graham 1990: 200. Synonymy by
Graham 1990: 200.

212

Journal of Hymenoptera Research

Parectroma Brethes 1913: 91. Type species: Pa-
redroma huebrichi Brethes; by original desig-
nation. Synonymy by De Santis 1960: 113.

Diagnosis. — Body black or, more com-
monly, dark with obscure to bright metal-
lic green luster on head and mesosoma.
Head with broad, shallow scrobal depres-
sion occupying most of region between
eyes and anterior ocellus, and smoothly
rounded into parascrobal regions (Figs. 1-
12). Clypeus distinctly delineated by su-
tures and at least slightly convex (Figs. 4,
6). Eye superficially bare or sparsely se-
tose (Figs. 1-16). Flagellum (Figs. 19^2)
with fu, ring-like; fu, ring-like to longer
than wide but without longitudinal sen-
silla; fuj with longitudinal sensilla. Pro-
notum about half as long as mesoscutum
(length not evident in Figs. 43^8 because
of view), evenly setose and with very fine
to distinct net-like engraved sculpture ex-
cept along posterior margin. Mesoscutum
(Figs. 43-48) setose and sculptured like
pronotum except lateral lobes usually
broadly bare and sometimes smooth me-
dially; axillae widely separated; scutoscu-
tellar suture deeply sulcate along anterior
margin of scutellum and for short distance
along anteromedial margins of axillae
(Figs. 43-48); scutellum bare at least pos-
teromedially, with bare frenum delineated
by variedly distinct transverse sulcus
(Figs. 50, 52, 54); prepectus bare (Figs. 61,
63); mesopleuron with shiny, characteris-
tically shaped femoral depression that in-
cludes abrupt or carinate anterior margin,
deep transepistemal pit posteromedially,
and arcuate transepistemal sulcus be-
tween pit and anterobasal edge of meso-
coxa (Figs. 61, 63). Metanotum with three
or more long, curved, paramedial setae di-
rected toward median (Figs. 43-48); meta-
pleuron with supracoxal flange and with
two or more carinae at least on flange
(Figs. 64-66); metacoxa conspicuously se-
tose dorsobasally (Figs. 55-66). Propo-
deum coarsely, irregularly sculptured,
without complete median carina though

often with variedly distinct, inverted Y-
shaped carina, and with long white setae
lateral to each spiracle (Figs. 43-48). Fore-
wing with marginal vein relatively short,
at most 0.3 length of submarginal vein,
subequal in length or shorter than stigmal
vein and shorter than postmarginal vein.
Petiole at least two-thirds as long as pro-
podeum, tubular but divided into dorsal
and ventral parts by lateral sulcus (Figs.
63-66), dorsally reticulate and usually
with irregular longitudinal carinae or
stronger parallel keels (Figs. 43-48, 55-60),
ventrally concave (Figs. 75, 76), and with
anterior margin carinate (Figs. 43-48, 55-
60). Gaster (Fig. 74) with terga low convex,
non-collapsing, and mostly smooth and
shiny, at most with very fine micropunc-
tulate or coriaceous sculpture on Gt,-Gt-;
Gt, and Gt, occupying at least half length
of metasoma; Gt, basally with at most 4
setae dorsolaterally near petiole (Figs. 43-
48, 55-60); GSj concave basolaterally for
reception of posterolateral margin of pet-
iole and with A-or n -shaped, often lon-
gitudinally carinate, basomedian projec-
tion (Figs. 75, 77, 78).

Discussion. — Asaphinae was first estab-
lished as the tribe Asaphini in the pter-
omalid subfamily Miscogasterinae by
Ashmead (1904). The group has been rec-
ognized as a subfamily of Pteromalidae
since Graham (1969). Graham (1969: 78)
stated that "the group appears to me dis-
tinct enough to be regarded, at least pro-
visionally, as a subfamily," but he did not
list any features that he considered differ-
ential and simply keyed out assigned gen-
era in three places in his key to pteromalid
subfamilies. He classified three genera in
Asaphinae, Asaphes, Hi/pehmerus Girault,
and Bainvnlia Waterston, and tentatively
proposed that Bairamlia Waterston (1929)
was a junior synonym of Parasiiphodes
Schulz (1906) (Graham 1969: 84). Howev-
er, Boucek (1988) not only treated Parasa-
phodes as a valid genus, he established the
new subfamily Parasaphodinae for it.
Boucek (1988) also removed Bairamlia

Volume 7, Number 2, 1998

213

from Asaphinae, stating (p. 343) that it
"seems to have closer relation with the
current Miscogasterinae," and assigned
two additional genera to the subfamily,
Ausasaphes Boucek and Enoggera Girault.
In addition to keying out assigned genera
in two places in his key to genera Boucek
(1988: 343) listed several features that
"seem to be of major importance" for the
subfamily. In the course of this study we
saw specimens (CNCI) of an undeter-
mined number of species from Central
and South America that appear to belong
to a fifth, undescribed genus that should
be classified in Asaphinae.

It is beyond the scope of this study to
determine the validity or limits of the sub-
family in Pteromalidae or to resolve the
relationships of the genera currently as-
signed to Asaphinae. The diagnoses given
above are based on examination of the
four currently assigned genera and the un-
described genus from the Neotropical re-
gion. Asaphes is distinguished from the
other four genera by one apparent autapo-
morphy, the presence of long, paramedial,
medially-directed setae on the metanotum
(Figs. 43-48, 55-60). Other Asaphinae
have the metanotum bare.

Both Ausasaphes and Enoggera are re-
stricted to Australia and can be identified
using the key provided by Boucek (1988).
Hyperimerus was previously thought to be
Holarctic, but during the course of this
study we saw two females of a new spe-
cies from Guatemala (CNCI) that is simi-
lar to H. corvus Girault and a single female
(CNCI) from Thailand that represents an-
other new species. Asaphes is the most
widely distributed genus, though it likely
was introduced into Australia by man
(Boucek 1988). The keys of Graham (1969),
Boucek and Rasplus (1991), or Boucek and
Heydon (1997) can be used to differentiate
Asaphes from Hi/perimerus and from other
pteromalid genera, though features used
in all three keys require slight clarification.
Graham (1969: 78) was inaccurate in stat-
ing that the gena of Hyperimerus is not bor-

dered by a sharp carina. Also, the petiole
of Asaphes is transverse in some species
though certainly it is much longer and
more conspicuous than that of Hyperime-
rus (see Graham 1969: 78 and Boucek and
Rasplus 1991: 30). The differentiating cou-
plet for Asaphes in Boucek and Heydon
(1997: 567) has to be modified with the
discovery that some species have a dis-
tinctly sculptured frenum (Figs. 50, 52).
Asaphes is further distinguished from Hy-
perimerus by having a setose metanotum
(Figs. 43-60), the basal gastral tergum
sparsely (Figs. 43-48, 55-60) rather than
extensively setose near the petiole (Gra-
ham 1969), the prepectus bare (Figs. 61,
63) rather than setose, and the anterior
margin of the femoral depression abruptly
margined (Figs. 61, 63).

Species of what we consider to be the
new Neotropical genus more closely re-
semble species of Asaphes than Hyperime-
rus because individuals have a long peti-
ole and sparsely setose gaster. However,
they are readily distinguished from spe-
cies of Asaphes by a distinctly reticulate
femoral depression, mostly smooth and
shiny propodeum with a straight median
carina, uniformly cylindrical and longitu-
dinally carinate petiole, evenly convex
and entirely or anteriorly longitudinally
carinate first gastral sternum, and conspic-
uously setose eyes.

Biology. — Species of Asaphes have usu-
ally been considered to be hyperparasites
of aphids (Homoptera: Aphididae),
through the following primary parasites:
Aphidiinae (Ichneumonoidea: Braconi-
dae), Encyrtidae, and Aphelinidae (Chal-
cidoidea) (Hagen and van den Bosch
1968). However, there are a few published
records of other Homoptera or predators
of aphids as hosts. Lai (1934) reared a spe-
cies identified as A. vulgaris from nymphs
of the pear psylla, Psylla pyricola Forster
(Homoptera: Psyllidae), in Scotland, and
McMullen (1966) reared A. suspensus (mis-
identified as A. vulgaris in McMullen 1996,
1971, and in Philogene and Chang 1978)

214

Journal of Hymenoptera Research

from P. pyrkola in British Columbia, Can-
ada. In both instances the Asaphes likely
was a hyperparasite because also reared
were Prionomitus mitratus Dalman (Encyr-
tidae) and Trechnites insidiosus (Crawford)
(Encyrtidae), respectively. Brown and
Clark (1960) reared three females of A.
brevipetiolatiis (as Asaphes sp.) from a pu-
parium of Neocnemodon coxalis (Curran)
(Diptera: Syrphidae), a predator of the bal-
sam woolly aphid, Adelges piceae (Ratze-
burg), likely as a hyperparasite because
four specimens of Syrphophagus quadrima-
culatus (Ashmead) (Encyrtidae) were
reared from the same puparium. We have
seen additional specimens of A. brevipe-
tiolatus whose labels and host remains in-
dicate they were reared from syrphid lar-
vae, as well as rearings from the spruce
budworm, Choristoneura fumiferana (Clem-
ens) (Lepidoptera: Tortricidae), and the
balsam fir sawfly, Neodiprion abietis (Har-
ris) (Hymenoptera: Diprionidae). One
specimen reared from C. fumiferana had an
unidentified braconid cocoon associated,
indicating that A. brevipetiolatus is a hy-
perparasite of the spruce budworm. Mul-
tiple specimens of A. brevipetiolatus asso-
ciated with some rearings indicate that it
is or can be a gregarious parasite.

Species of Asaphes are ectoparasites and
except for A. brevipetiolatus apparently al-
most always solitary hyperparasites. Usu-
ally they are secondary hyperparasites,
ovipositing into mature larvae, prepupae
or pupae of the aphidiine, encyrtid, or
aphelinid primary parasite within the
aphid mummy. The primary parasite is al-
most immediately paralysed by a venom
as part of oviposition and further devel-

opment is prevented regardless of the
stage attacked (Sullivan 1972, Keller and
Sullivan 1976, Bocchino and Sullivan
1981). Studies have also shown that spe-
cies can be tertiary hyperparasites if the
primary parasite has already been parasit-
ized by some other secondary parasite,
such as by species of Megaspilidae (Hy-
menoptera: Ceraphronoidea) or Charipi-
dae (Hymenoptera: Cynipoidea) (Gris-
wold 1929, Sullivan 1972, Carew and Sul-
livan 1993), or by another Asaphes (Levine
and Sullivan 1983). Sullivan (1972) also
showed for A. californicus, and Keller and
Sullivan (1976) for A. suspensus (as A. lu-
cens), that females will make multiple at-
tacks and drillings of the same aphid
mummy and frequently deposit more
than one egg, though in all cases only a
single adult emerged. However, superpar-
asitism and multiparasitism are possible
though probably very rare because Carew
and Sullivan (1993) reared two dwarf fe-
male A. suspensus (as A. luceus) from one
aphid mummy, and a dwarf male together
or with another secondary parasite from
another mummy.

Host feeding was demonstrated for A.
californicus by Sullivan (1972), for A. sus-
pensus by Keller and Sullivan (1976), and
for A. vulgaris by Le Ralec (1995). A feed-
ing tube is constructed prior to oviposi-
tion. Once feeding is completed the ovi-
positor is reinserted, the feeding tube bro-
ken by the ovipositor, and an egg depos-
ited (Keller and Sullivan 1976). Griswold
(1929) described and illustrated the im-
mature stages. Mating habits are de-
scribed by Griswold (1929) and Sekhar
(1958).

KEY TO SPECIES OF ASAPHES WALKER FROM AMERICA NORTH OF MEXICO

1 Female 2

Male 7

2(1) Head in frontal view subquadrate with dorsal margin of torulus distinctly below lower
orbit (Figs. 1, 2); malar space as long as width of eye or longer and at least 0.7 eye
length (Figs. 1, 2); frenum with distinct net-like sculpture similar to that on scutellum
(Figs. 49, 50, 55); forewing with very narrow speculum, the bare area closed basally

Volume 7, Number 2, 1998 215

by setae and with line or lines of setae immediately behind submarginal vein on disc
(Fig. 73) Asaphes brevipetiolatus n. sp.

- Head in frontal view transverse-subtriangular with dorsal margin of torulus about in
line with lower orbit (Figs. 5-12); malar space at most about 0.8 width of eye and less
than 0.65 eye length; frenum usually smooth and shiny except for longitudinal carinae
near lateral margin (Figs. 56, 57, 59, 60); forewing sometimes with broad speculum
and without setae near submarginal vein on disc (Figs. 68, 70) 3

3(2) Frenum with distinct, uniform, engraved net-like sculpture similar to scutellum (Figs.
51, 52)'; metapleuron bare (Fig. 65); petiole at m.ost as long as wide and usually slightly
though obviously transverse (Fig. 51); legs with at least trochanters and all but extreme
apices of femora dark (trochantelli rarely yellowish); forewing with speculum basally
closed by setae and with line or lines of setae immediately behind submarginal vein

on disc (Fig. 72) Asaphes petiolatus Zetterstedt

Frenum usually broadly smooth and shiny except for longitudinal carinae near lateral
margins (Figs. 56, 59, 60), but if with obscure net-like sculpture paramedially and along
posterior margin (Fig. 54) then metapleuron with at least some setae anteroventrally
and often conspicuously setose (Figs. 61, 62); petiole usually at least slightly longer
than wide except sometimes if metapleuron also setose (Figs. 44-48); legs usually en-
tirely yellowish or at least with trochanters yellowish; forewing sometimes with broad,
open specidum and without setae near submarginal vein on disc (Figs. 68, 70) 4

4(3) Metapleuron with at least several setae anteroventrally and usually more extensively
setose (Figs. 61, 62); legs with trochanters dark, concolorous with femora; forewing
with speculum basally closed by setae and with line or lines of setae immediately

behind submarginal vein on disc (Fig. 71) Asaphes hirsntus n. sp.

Metapleuron bare (Figs. 63-66) or at most with one or two short seta anteroventrally;
legs usually with at least trochanters yellowish and often uniformly light colored, but
if trochanters dark then speculum broad, open, and disc without setae near submar-
ginal vein (Figs. 68, 70) 5

5(4) Speculum distinct, broad basally and narrowed toward stigmal vein, without setae
near submarginal vein and immediately beyond basal setal line at least as wide as
distance between first and third setal lines on disc (Fig. 70); head in dorsal view
dishnctly concave between inner orbits (Fig. 15); hind leg with trochanter and femur
(except often trochantellus and apex) infuscate to black [east of Manitoba in Canada
and northeastern seaboard states in USA] Asaphes vulgaris Walker

- Speculum variedly distinct, most often with line of setae behind submarginal vein
separated from vein by distance at most equal to distance to next setal line (Fig. 67),
but sometimes with relatively broad speculum basally closed by one or more setae
(Fig. 68); head in dorsal view shallowly concave between irmer orbits (Fig. 16); hind
leg uniformly yellowish or with trochanter and trochantellus usually yellowish (very
rarely black) in contrast to infuscate or black femur [transcontinental] 6

6(5) Legs more or less uniformly light-colored, yellow; forewing always with several setae
close to submarginal vein on disc, hence speculum very poorly developed (Fig. 69). .

Asaphes suspensus Walker

Legs with at least metafemur in part darker than light-colored metatrochanter unless
metacoxa also light-colored, and then forewing with broad speculum except for one
or two setae interrupting bare area (Fig. 68) Asaphes califomicus Girault

7(1) Antenna with scape and pedicel yellowish, contrasting distinctly with dark flagellum;
scape with ventral margin sinuate, protuberant ventrobasally and tapered apically
(Figs. 31, 32); head with dorsal margin of torulus distinctly below lower orbit (Figs. 3,

4) Asaphes brevipetiolatus n. sp.

Antenna more or less uniformly dark or with flagellum lighter in color; scape usually
almost cylindrical or spindle-shaped (Figs. 34, 36, 38, 40, 42); head with dorsal margin
of torulus almost in line with lower orbit (Figs. 5-12) 8

216 Journal of Hymenoptera Research

8(7) Outer surface of scape with variously elongate-oval or lanceolate microsetose sensory
region (Figs. 33, 34a); legs usually u^ith at least metafemur distinctly darker than meta-
trochanter unless metacoxa also yellov/ish-brown Asaphes califortiicus Girault

- Outer surface of scape without distinct setal patch, at most with linear setal band along
ventral margin (Fig. 40a); legs varied in color, but often uniformly yellowish or with
trochanters similarly dark as femora 9

9(8) Metapleuron with at least 3 setae in extreme anteroventral angle, and often much more
extensively setose (Figs. 61, 62); forewing without distinct speculum, disc setose behind
submarginal vein, the distance between vein and first setal line only slightly greater
than distance between first and second setal lines; metafemur and metatrochanter
usually dark; scape usually with slightly concave to flat ventral surface over at least

apical two-thirds (Fig. 36a) Asaphes hirsuhis n. sp.

Metapleuron bare (Figs. 63-66); other features varied but usually either forewing with
distinct speculum (Figs. 68, 70) or legs yellowish beyond coxae and scape without

distinctly differentiated ventral surface 10

10(9) Forewing disc with broad speculum narrowed toward stigmal vein, but over most of
length width of bare band about equal to distance from first to third setal lines on
dorsal surface of disc (Fig. 70); basal cell with single line of setae behind submarginal
vein except apically near basal setal line (Fig. 70); legs often with at least metafemur
dark; frenum smooth and shiny (Fig. 60); petiole at least 1.25 times as long as wide
(Figs. 59, 60) Asaphes vulgaris (Walker)

- Forewing disc with at least 3 setae on dorsal surface separated from submarginal vein
by distance about equal to width of vein or by distance between first and second setal
lines (Figs. 69, 72); basal cell with more than one line of setae behind submarginal vein
over most of length and often more or less uniformly setose (Figs. 69, 72); legs usually
entirely yellowish; frenum sometimes with fine sculpture (Figs. 52, 58); petiole some-
times less than 1.2 times as long as wide (Fig. 58) 11

11(10) Legs entirely yellowish beyond coxae; frenum smooth and shiny (Fig. 59); petiole

usually at least 1.2 times as long as wide (Fig. 59) Asaphes suspensus (Nees)

- Legs with trochanters, trochantelli and most of femora dark-; frenum with fine sculp-
ture at least paramedially (Fig. 58); petiole at most 1.15 times as long as wide (Fig. 58)
Asaphes petiolatus Zetterstedt

' Females from western Europe sometimes with frenum quite broadly smooth with only very obscure sculp-
ture.

^ Some western European males with legs almost entirely yellow beyond coxae, see 'Recognition' for A. pe-
tiolatus.

Asaphes brevivetiolatllS Gibson and 45'W, 2000 m, 9.VII.86, H. Goulet, subalpine meadow

Vikbere new species '^ ^ '■ ^'''^^'^^ '^'^""' 12.VIII.53, F.I.S. No. A803D, ex.

I ia 1? 10 on •Ti To ^^ 1Q svrphid, hostprob.aphidO 9,NFRC). FortMacKay,

(Figs. 1^, 13, 14, 19, 20, 31, 32, 43, 49, 2.8 km N bridge, 2.IX.79, G.J. Hilchie & J. Ryan, mix-

50, 55, 64, 73) conifer forest (2 9 ). Jasper Gate, 2.111,51, F.I.S. No

A3125B, ex. syrphid, host prob. aphid (2 9, NFRC).

T>/fK- matcnnl.-Holoti/pc. female (CNCI, Type No. j^^p^^ National Park, Maligne Road, 7.VI11.50, F.I.S.

22267): CANADA, New Brunswick, F'ton [Frederic- ^o. A615A, ex. Metasyrphus lapponicus (3 5, NFRC).

ton), em. June 23, 1966, R.C. Clark, AP.66-10-2, ex. obed, 21.VIII.50, F.I.sl No. A20H9C, ex. Metas\/rp^hus

Syrphidae. Allotype, male (CNCI): same data as ho- lapponicus (4 9, NFRC). Rocky Mountain House, 23

lotype. Paratypes: CANADA. Yukon Territory: Ross mi. NW, 17.VIII.53, F.I.S. No. A781D, ex. syrphid.

River, 16.IV-31.VIII.84, S.&J. Peck, aspen willow river host prob. aphid (1 9, 1 S, NFRC). Seebe, 13.VI.68,

terrace (1 9, 2 S). British Columbia: Manning Pro- ex. Croiiuirtiuin coinivulnu- (iun^us). Pinus contort:! wyr.

vincial Park, 2 km N Blackwall Peak, 49°07'N 121' latifolia, 688 1190 03. Manitoba: Picnic Bog, 6.VI.61,

Volume 7, Number 2, 1998

217

F.I.S. (1 9, NFRC). Warkworth near Churchill,
29.VI.52, J.G. Chillcott (1 9 ). Quebec: Ct. Jette, RIF
'40, 5103B, em. 18.VI1I, ex. Syrphidae sp.? (2 9).
Forbes, 26.VI.52, L. Daviault, associated with Choris-
toneiira fiimiferana, rearing no. 20 (5 9 ). Laniel,
9.VI1I.40, C.E. Atwood, experiment no. 12131-69, Cn-
coecia fumiferana (2 9). Montcalm, em. 19.V1.11 from
sp. B [?] worm, parasite of Tortris fumiferana (1 3 ).
New Brunswick: same data as holotype (2 9, 3 d; 1
9 and 6 used for SEM). Fredericton, em. 8.VII.47,
N.R. Brown, 18839-4d5, ex. Neocnenwiim coxalis (8 9,
1 u ). Newfoundland and Labrador: Gallants,
9.VI1.57, 57-0016(01) BIO (4 9, AFRC), 14.VI1.59, 59-
6009(01) A1-A9 {19,6 6 ), F.I.S., host: Neodiprion abie-
tis. Uncertain locality (likely Newfoundland or New
Brunswick): 23 [mi. ?) NW Rocky, 17.VIII.53,
53A781D, ex. syrphid (6 9, 8 d). USA. Alaska: Fair-
banks, 23.V1.45, 45-19058, par. aphids on wild rasp-
berry (1 9, USNM). Colorado: Green Mountain Falls,
Canyon, 10.VIII.41, 10,000', H.H. Ross (1 9, INHS).
Idaho: Bounds Creek, Fairfield, 5.1X.63, H. Flake & K.
Lister, Hopk US no. 50-718a, host: attached pupae
(Syrphidae) (8 9, USNM). Maine: Liberty, em.
8.VL48, parasite 48-Cll syrphid fly, beaten ex. Fir (2
9, USNM). Megalloway, 23.VL49, parasite 49-255,
Neodiprion abietis, beaten ex. Fir (6 6, USNM). Wash-
ington: Yakima Co., Green Lake Road, 31.V1II.94, K.S.
Pike, ex. Bhicus sp. or ichneumonid, from Cinara chi-
nookiana or unknown on Abies tasiocarpa (subalpine
fir) (1 3, WSUC).

EXTRALIMITAL— FINLAND (FENNIA, SUOMI).
Savonia australis [= Sa, ESJ: Ristiina, 6826:502,
29.V11.83, M. Koponen (1 6, DAZH). Ostrobottnia
borealis, N part [= ObN]: Pello, Pentik, 7417:368,
28.VL97, V. Vikberg (1 9, VVPC). Kuusamo [= Ks]:
Kuusamo, 7358:596, 27.V1.82, M. Koponen (1 9,
DAZH).

Etymology. — From the Latin brevis, short,
and petiolus, stalk, in reference to the
transverse petiole of this species.

Female. — Head and mesosoma black
with obscure metallic green luster under
some angles of light; legs entirely black or,
more often, black or dark brown with ex-
treme apex of femora, extreme base and
apex of tibiae to entire tibiae, and tarsi
lighter in color, rufous to yellowish. Head
subquadrate in frontal view (Fig. 1), width
at most 1.2 times height, and in lateral
view lower face abruptly to almost right-
angled relative to upper face (Fig. 2); in-
terorbital region in dorsal view deeply,
broadly concave (Fig. 13); gena as long as
or slightly longer than eye width and at

least 0.72 eye length; dorsal margin of to-
rulus distinctly below lower orbit (Figs. 1,
2). Antenna (Fig. 19) with pedicel length
about 3 times greatest width; funicle with
fu, ring-like, fu.-fuj subquadrate (Fig. 20),
and fus-fu„ increasingly transverse. Me-
soscutum (Fig. 43) with mesoscutal lateral
lobes broadly bare medially, and with fine
engraved net-like sculpture over bare
area. Scutellum mostly bare except along
extreme anterior and lateral margins (Fig.
43); frenum with distinct, uniform, en-
graved net-like sculpture similar to scutel-
lum (Fig. 50). Metapleuron with at most 5
setae anteroventrally, these setae mostly
in line along base of metapleural flange.
Forewing with basal cell evenly setose
(Fig. 73); disc with narrow speculum, the
speculum closed basally by setae and with
one or more lines of setae immediately be-
hind submarginal vein (Fig. 73). Petiole in
dorsal view distinctly transverse, length
1.4-1.7 times width, reticulate with irreg-
ular longitudinal carinae or stronger keels
(Fig. 49).

Male. — Antenna with scape, pedicel and
fu, yellow to yellowish-orange, contrast-
ing distinctly in color with rest of dark fla-
gellum; color pattern otherwise similar to
female except head and mesosoma usually
with more distinct metallic green luster;
legs usually more extensively light-col-
ored, but at least hind leg brownish ba-
sally, including trochanter. Scape (Figs.
31-32) with dorsal margin slightly convex,
ventral margin sinuate so as to be broad-
est subbasally and tapered to apex; inner
and outer surfaces uniformly setose and
sculptured. Pedicel (Figs. 31-32) length
about 3 times width and about 0.40-0.45
scape length. Flagellum length subequal
to head width; funicle (Fig. 31) with all
segments at least slightly transverse or
with one or more of fu,_, quadrate to very
slightly longer than wide. Structure oth-
erwise similar to female except head in
dorsal view with interorbital region even
more deeply, broadly concave (Fig. 14);
malar space only about 0.77-0.87 eye

218

Journal of Hymenoptera Research

width and 0.5-0.6 eye length; petiole usu-
ally only slightly transverse (Fig. 55); and
sculpture of frenum often distinctly finer
than on scutellum (Fig. 55). Setal pattern
similar to female except metapleuron with
at most 2 short setae.

Distribution. — Holarctic; in North Amer-
ica transcontinental within the Boreal re-
gion and extending south along the Cas-
cade and Rocky Mountains into Colorado
(Fig. 79).

Biology. — Asaphes brei'ipetiolatus is a par-
asite of Neocnemodou coxalis (Curran), Meta-
syrphus lapponicus (Zetterstedt) and possi-
bly other Syrphidae (Diptera) based on as-
sociated host pupal remains and label
data. The balsam fir sawfly is also indi-
cated as a host because two cocoons of
Neodiprion abietis (Harris), one with two
emergence holes and the other torn in
half, are preserved with specimens. Labels
from three different rearings in Quebec in-
dicate that A. brevipetiolatus is also a par-
asite of the spruce budworm, Choristoneu-
ra fumiferana (Clemens). Although there
are no associated host pupae, one of the
two Laniel specimens has an unidentified
braconid cocoon preserved with it, indi-
cating that A. brevipetiolatus could be a hy-
perparasite of the spruce budworm
through braconid primary parasites. There
is also a single record, based on label data,
of the species parasitizing an unidentified
aphid on wild raspberry, plus a rearing
from either Cinara chinookiana Hottes or an
unknown aphid via either Blacus sp. (Bra-
conidae) or an unidentified ichneumonid.
More than one emergence hole in some
syrphid puparia and one balsam fir sawfly
cocoon, plus multiple-mounted speci-
mens, labelling, or both, for these two
hosts and two of the three spruce bud-
worm records indicate that A. brevipetiola-
tus is often gregarious when parasitizing
larger non-aphid hosts. It remains to be
demonstrated more conclusively that the
balsam fir sawfly and spruce budworm
are more than just accidental hosts, and
whether A. brevipetiolatus is a primary or

hyperparasite of these two species. How-
ever, based on current evidence it seems
likely that host acceptance for A. brevipet-
iolatus includes not only syrphid larvae
but other relatively large, oblong, brown-
ish pupae or cocoons on coniferous trees.
This suggests an evolutionary progression
in Asaphes from parasitism of hymenop-
terous primary parasites of aphids, to syr-
phid-larvae predators of aphids and their
parasites, to other hosts that resemble syr-
phid larvae.

Remarks. — Males are easily distin-
guished by structure and color of their
scape and pedicel, both are yellowish in
distinct contrast to the flagellum beyond
fu,. Also, the pedicel is conspicuously
long, almost half the length of the scape,
and the scape is widened subbasally so
that its ventral margin is sinuate (Figs. 31,
32).

Females are distinguished by a combi-
nation of features, including a sculptured
frenum (Fig. 50), dark trochanters and tro-
chantelli, closed speculum (Fig. 73), trans-
verse petiole (Fig. 49), and unique head
structure (Figs. 1, 2). Because the legs are
almost uniformly dark basally, females
most closely resemble those of A. vulgaris,
A. petiolatus and A. hirsutus, but are distin-
guished from all three species by de-
scribed head structure. Absence of a broad
speculum further differentiates females
from those of A. vulgaris. Those A. brevi-
pietiolatus females with setae on the meta-
pleuron are more likely to be mistaken for
females of A hirsutus, particularly because
some A. hirsutus females have a slightly
transverse petiole (Fig. 53) and some have
the frenum extensively, though finely
sculptured. However, in A. hirsutus the
frenum is always more or less broadly
smooth immediately behind the frenal sul-
cus even though often sculptured poste-
riorly (Fig. 54). Only head structure readi-
ly differentiates females of A. brevipetiola-
tus and A. petiolatus.

Volume 7, Number 2, 1998

219

Asaphes califomicus Girault

(Figs. 7, 8, 21, 22, 33, 34, 44, 56, 63, 67,
68, 76)

Asn/'/ies califomicus Girault 1917[330]: 1. Type
data: USA: California [Spreckels], [25] Sep-
tember [1916]; reared from [parasitized]
Aphis nunicis; [Chittenden No. 1671]; C.F.
Stahl [collector]. Sex described: female. Ho-
lotype by monotypy; USNM, type no. 21411.

Female. — Head and mesosoma dark
with varying intensity of olive green me-
tallic luster under some angles of light;
legs with trochanters and trochantelli al-
most always uniformly yellowish to yel-
lowish-brown, at least middle and hind
femora black except often apically, and
tibiae and tarsi usually yellowish to red-
dish-brown. Head transverse-triangular in
frontal view (Fig. 7), width at least 1.25
times height, and in lateral view lower
face evenly curved into upper face (Fig. 8);
interorbital region in dorsal view relative-
ly shallowly concave; gena length about
0.62-0.80 eye width and 0.50-0.65 eye
length; dorsal margin of torulus approxi-
mately in line with lower orbit (Figs. 7, 8).
Antenna (Figs. 21, 22) with pedicel length
at most about 2.5 times width; funicle with
fu, ring-like, fu_,-fu4 quadrate to trans-
verse, and fu5-fu8 increasingly transverse.
Mesoscutum (Fig. 44) with lateral lobes
broadly bare medially, and either smooth
and shiny or with fine, engraved net-like
sculpture over bare area. Scutellum (Fig.
44) mostly bare except along extreme an-
terior and lateral margins; frenum smooth
and shiny except finely carinate laterally.
Metapleuron bare. Forewing with basal
cell evenly setose to bare except for single
row of setae on dorsal surface (Fig. 68);
disc often without distinct speculum, dis-
tance between either basal setal line or
submarginal vein and first setal line on
disc at most about equal to distance be-
tween first and third setal lines on disc
(Fig. 67) or, if with large and conspicuous
speculum, then bare region with at least 2
dorsal setae within bare region, the setae

usually separated from basal vein and / or
submarginal vein by distance about equal
to length of setae or less (Fig. 68). Petiole
at least quadrate and usually slightly lon-
ger than wide, but less than 1.3 times as
long as wide (Fig. 44), reticulate with ir-
regular longitudinal carinae (Fig. 44) or
stronger keels.

Male. — Color pattern similar to female
except legs sometimes entirely yellow
(more commonly with at least metafemur
infuscate to black); antenna usually uni-
formly brown to black except sometimes
apex of pedicel and fu, lighter in color.
Scape (Figs. 33, 34) length about 3.5-5.0
times width, ovoid to spindle-shaped,
with dorsal and ventral margins convex to
subparallel over most of length; outer sur-
face with subbasal, ovoid to elongate-lan-
ceolate, microsetose sensory region (Figs.
33, 34a); inner surface with line of ven-
trally directed setae, often from along fine
ridge, parallelling ventral margin of scape
below midline and mediolongitudinal
bare, smooth band (Fig. 34b). Combined
length of pedicel and flagellum less than
2.5 times scape length (Fig. 33) and at
most subequal in length to head width; fu-
nicle with all segments at least slightly
transverse or with one or more of fu,_s
quadrate to very slightly longer than
wide. Structure and setal pattern other-
wise similar to female except petiole al-
ways distinctly (about 1.25-1.80 times)
longer tlian wide (Fig. 56) and sometimes
entirely reticulate without longitudinal ca-
rinae.

Distribution. — Restricted to western
North America within the Nearctic region,
except for one anomalous record from
Georgia (see further under 'Remarks')
(Fig. 82). CANADA. Yukon Territory,
British Columbia, Alberta. USA. Alaska,
Arizona, California, Colorado, ? Georgia,
Idaho, Kansas, New Mexico, Nevada, Or-
egon, Washington, Utah. EXTRALIMI-
TAL. MEXICO (BMNH: 9, (5;CNCI: 9,3;
EMEC: ?), ARGENTINA (MLPA: 9, S;
TAMU: 9, 6); BOLIVIA (USNM: 9),

220

Journal of Hymenoptera Research

Table 1. Host information for Asaphcf cnlifoniiciis based on observed specimens; a question mark follows
rearings or identifications indicated as questionable on the labels. Unless otherwise footnoted, all 'aphid or
other hosts' are Homoptera (Aphidoidea: Aphididae) and all 'associated primary hosts' are Hymenoptera
(Braconidae: Aphidiinae).

Aphid or other Hosts

Associated Primarv Hosts

Museum
Acronvms

Acyrthosiphon lactucae (Pas-
serin)
A. pisuin (Harris)

Amphoroplwrii rubi (Kalten-

bach)
Aphis sp.
A. ccanoDii Clarke
A. fabnc Scopoli
A. goss\/pii Glover

A. hederae pseiniohetierae

Theobald
A. helianthi Monell

A. holociisci Robinson

A. rumicis L.

A. sambuci L.

A. spiraecola Patch

A. I'arians Patch
Aphthargelia symphoricarpi

(Thomas)
Brachycauiius caniui (L.)
6. helkhrysi (Kaltenbach)
6. tragopogonis (Kaltenbach)
Brachycorynelln aspaiagi

(Mordvilko)
Braggia sp.

B. eriogoni (Cowen)
Brevicoryne hrassicae (L.)

Capitophorus claeagni (del

Guercio)
Ciwariella acgopodii (Scopoli)

C. paslinacac (L.)

Acuiitliociuidus sp. ?

Ap^liidius sp.

A. alius Muesebeck

A. nigripes Ashmead

A. smithi Sharma & Subba Rao

Diacretus sp.

Ephedrus sp.

Lysiphlebus {Adialytus) salicaphis

(Fitch)
L. (Phlebus) testaceipes (Cresson)
Pauesia cnlifornicus (Ashmead)
Praon sp.

T. (Trioxys) ccmplniwtiis Quilis
T. (Trioxys) curvicaudtis Mackauer
Praon sp.

Apliidius sp., A. eroi Haliday, A. smi-

tlii Sharma & Subba Rao
Praon pcquodorum Viereck

Lysiphlebus sp.

Aphidius sp.; Lysiphlebus (Phlebus)
testaceipes (Cresson)

Lysiphlebus (Phlebus) testaceipes
(Cresson)

Ephedrus californicus Baker; Lysiphle-
bus (Phlebus) testaceipes (Cresson);
Praon sp.

Alhxysta sp.

Lysiphlebus (Phlebus) testaceipes
(Cresson); Praon sp.

Diaeretiella rapae (M'Intosh)

Lysiphlebus (Phlebus) testaceipes
(Cresson)

Diaeretiella rapae (M'Intosh)

VVA

WSUC

CA

USNM, UCRC

CA

UCRC

CA

UCRC

CA

USNM, EMEC

CA

UCRC

CA

EMEC

CA

EMEC

CA

UCRC

CA

USNM, EMEC

CA

UCRC

CA

EMEC

CA

EMEC

WA

WSUC

CA, OR, WA

UCRC, EMEC,

USNM, WSUC

WA

WSUC

CA

EMEC, UCRC

WA

WSUC

WA

WSUC

CA, NM

EMEC, UCRC,

USNM

CA

UCRC

WA

WSUC

WA

WSUC

CA

USNM

WA

WSUC

WA

WSUC

WA

WSUC

WA

WSUC

WA

WSUC

CA

EMEC

WA

WSUC

WA

WSUC

Aphidius salicis Haliday
Aphidius salicis Haliday

WA

WA

CA, OR, WA

WA

CA, WA

VVA

WSUC

WSUC
EMEC, OSUC,

WSUC
WSUC

CASC, WSUC
WSUC

Volume 7, Number 2, 1998
Table 1. Continued.

221

Aphid or other Hosts

Associated IVimarv Hosts

Museum
Acronvms

Chaeto$ip!wn {Peiilnlriclwpuf)

fragarnefolii (Cockerell)
Chaitophorus pcpulifotii

(Essig)
Chaitophorus salkkola Essig
Cinara ponderosae (Williams)
Diuraphis iioxia (Mordvilko)
Dysapliis (Pomapliii) plaii-

tagiuca (Passerini)?
Elatobium aWietiiium

(Walker)
Ericaphis gentneri (Mason)
Essigella sp.
E. californica (Essig)
E. pini Wilson
Eucalliptcni!. tiliae (L.)
Euceraphis punctipcnnis (Zet-

terstedt)
Flabellomicrosiplnmi knOivl-

toni Smith ?
Hayhurstia atriplkis (L.)
Hyiihpterus pruni (Geoffroy)
Hi/peromyzus {Neoiiasonovia)

nigrkornis (Knowlton)
lllinoia sp.
/. nzalene (Mason)
/. tirwikndri (Monell)
/. niorrisL'iti (Swain)
/. sinipsoui (MacGillivrav)
/. subviridis (MacDougall)
MacrosiphpiikUa ludcivkianac

(Oestlund)
Macrosip'hum sp.

M. dydcsmithi Robinson

M. creelii Davis

M. euphorhiac (Thomas)

M. parvifclii Richards
M. rosae (L.)

Metopu'lcphiiiKm dirfwdum

(Walker)
Mkrolophium cariwsuni

(Buckton)
Myzocallh sp.
M. coryli (Goeze)
Myzus [Nectarcfiphcn) persi-

cae (Sulzer)

Nasoucvui nquilegiae (Essig)

Aphidius sp.
Ephcdnca sp.

Pauesia sp.

Diaretklla rapae (M'lntosh)

Praon uiikuni Smith

Aphidius sp.

Alloxysta sp.; Prnon uiuciim Smith

Diaeretus sp.

Aphidius sp.; Diaeretus sp.

Priwu sp.

Diaretielln rapae (M'lntosh)
Praon sp.
Praon sp.

Alloxysta sp.; Aphidius sp.

Praon sp.
Aphidius sp.

Aphidius sp.; Praon sp.

Praon sp.

Alloxysta sp.; Aphelinus sp.'; Af/ii-
rfius nigripes Ashmead

Aphelinus sp.'; Aphidius sp., /4. a/iks
Muesebeck; Praon sp.

Praon sp.

T. (Trioxys) pallidas (Haliday)
T. (Trioxys) pallidas (Haliday)
Aphidius sp., A. matricarme Haliday;
Diaeretus rapae (M'lntosh); Lysiphlc
bus (Phlebus) testaceipes (Cresson);
Praon sp., P. unicum Smith

CA

WA

WA

WA

WA
WA

AB, CA,
WA

WA

OR,

UCRC, EMEC

WSUC

CA

EMEC, USNM

MT

WSUC

WA

WSUC

WA

WSUC

EC

UCRC, NFRC

WA

WSUC

CA

EMEC

CA

UCRC

CA

UCRC, USNM

CA

EMEC

CA

USNM

WSUC

WA

WSUC

CA

EMEC

WA

WSUC

WA

WSUC

WA

WSUC

CA

CNCl, EMEC

CA

EMEC

BC

EMEC

WA

WSUC

CA

UCRC

BC, CA, WA

CNCI, EMEC,

WSUC

WA

WSUC

WA

WSUC

CA

EMEC, UCRC

BC

CNCI

CA, WA

CASC, EMEC,

UCRC USNM,

WSUC

WA

WSUC

WSUC

WSUC
WSUC
CNCI, EMEC,

UCRC, USNM,

WSUC

WSUC

222

Table 1. Continued.

Journal of Hymenoptera Research

AphKl or i.ther Hostb

Associated Priniarv Hosts

Museum
Acron\'ms

Obtuskauiia sp.

O. artemisiphita (Knowlton
& Allen) ?

O. coweni (Hunter)

O. filifcliac (Gillette & Palm-
er)?

Oi'atus cratacgarius (Walk-
er)?

Phytomyza ilicis Curtis-

Phowdon humidi (Schrank)

Pseudoepameihaphis tridcnta-
tae (Wilson)

Rhodobiuni poroiiis (Sander-
son)

Rhopnloniyzitf (Jiidciikon) /on-
iccrac (Siebold)

R)ioptiloiiphuiii iiiscrtiini
(Walker)

R. viaidis (Fitch)

R. padi (L.)
Schizolachnus piiurndiatnc

(Davidson)
Sitobium <n'cnae (Fab.)
S. pteridis (Wilson)
Spdoccccus iiiipliciitus Ferris'
Therioaphis riclinu (Borner)
T. trifolii (Monell)

TIjripsnphis sp.

Urok'Kcoii (Lnndiersius) kn-
lonkac (Hottes)

U. russellac (Hille Ris Lam-
bers)

Wahlgreitiella ncrvnta (Gil-
lette)

WA

wsuc

WA

wsuc

WA

wsuc

WA

wsuc

WA

T. (Trioxys) boniicvdleiisis Smith

Praon sp.

Prnoii unkum Smith

Lysiphlcbus (Phkbiis) tcstaccipes
(Cresson)

Apliidius sp., A. avet\ap]ns (Fitch)

Praon amerkanum (Ashmead)
Praon exoktum palitans Muesebeck;
Trioxys (Trioxys) complanatus Quil-
ls
Praon sp.

Praon sp.

WSUC

BC

CNCI

WA

WSUC

WA

WSUC

CA

EMEC

WA

WSUC

WA

WSUC

WA

WSUC

WA

wsuc

CA

USNM

OR, WA

osuc, wsuc

BC, CA, OR

EMEC, UCRC

CA

EMEC

CA

UCRC

CA

UCRC, USNM

WA

WSUC

CA

EMEC

WA

WSUC

WA

WSUC

' Chalcidoidea (Aphelinidae).

' Diptera (Agromyzidae).

' Coccoidea (Pseudococcidae).

CHILE (ANIC:?; BMNH: 9,6; CNCI: ?;
EMEC: 9; TAMU: 9, S; UCDC: 9,6),
COLOMBIA (CNCI: 9), COSTA RICA
(CNCI: 9,6), DOMINICAN REPUBLIC
(CNCI: 9, 6), ECUADOR (CNCI: 9),
GUATEMALA (CNCI: 9,6), URUGUAY
(USNM: 9 ), VENEZUELA (CNCI: 9 ).

Biology. — Nearctic specimens with host
data indicate that A. californicus is strictly
a hyperparasite of aphids through aphi-

diine and aphelinid primary parasites,
though there are two anomalous records
from Phytomyza ilicis Curtis and Spilococ-
cus implicntus Ferris (Table 1).

Ti/;'e material examined. — The holotype
consists of a point-mounted specimen
and, in the general collection, a slide with
one hind leg and both antennae. The ho-
lotype otherwise has the head, right pair
of wings, and right front leg missing.

Volume 7, Number 2, 1998

223

Remarks. — We are uncertain both as to
the limits of morphological variation and
the true range of A. califoniiciis because
there may well be more than one species
represented within a species complex in
the New World. Our concept of A. califor-
nicus is influenced largely by the males,
which have a more or less well developed,
but usually elongate-lanceolate or oval,
sensory region on the outer surface of the
scape (Fig. 34a). Apparently the region is
always setose but the setae are not always
distinct because of what appears to be an
exudate covering the area in some speci-
mens. Also, the more elongate spindle-
shaped is the scape, the less obvious is the
line of setae on its inner surface (Fig. 34b)
because the line is closer to the ventral
margin. At least within the Nearctic re-
gion, females seem to have quite a stable
leg color pattern, with at least the hind leg
having yellowish trochanters in contrast to
darker femora. The size of the speculum
varies considerably, some individuals
have a broad bare band like A. vulgaris
(Fig. 68), others have the disc almost uni-
formly setose (Fig. 67), and others have in-
termediate states. Males and females from
the Nearctic region, like other species rec-
ognized from the Northern Hemisphere,
also consistently have a distinctly sulcate,
crenulate frenal groove (Figs. 44, 56). Be-
cause males sometimes have the femora
entirely yellow or only inconspicuously
infuscate they can easily be mistaken for
males of A. suspeusus if structure of the
scape is not examined carefully. A differ-
ence in antennal color often suffices to dif-
ferentiate unmounted males in ethanol.
Males of A. californicus have uniformly
brown antennae, whereas males of A. sus-
pensiis have the flagellum yellowish or at
least distinctly lighter than the scape.

We have seen one anomalous female
from Georgia (Mcintosh Co., Sapelo Is-
land, 28.IV-9.V.97, scrub sand dunes)
(CNCI). Not only is this locality substan-
tially outside the apparent range of A. cal-
ifornicus, the specimen has legs with the

trochanters to femora almost uniformly
dark (trochantelli yellowish-brown, simi-
lar in color to tibiae), which is an aberrant
color pattern for the species. A basally
closed and relatively narrow speculum in-
dicates that this specimen does not belong
to A. vulgaris, and it does not possess the
distinguishing features of other Nearctic
species whose females are characterized
by dark legs. The only species of Asaphes
otherwise known to occur in the south-
eastern USA is Asapihes suspiensus, which
has entirely yellow legs. Because it was
collected from coastal Georgia it is possi-
ble that it represents an accidentally intro-
duced species that is morphologically sim-
ilar to A. californicus. However, until ad-
ditional females with associated males can
be collected and examined, we tentatively
include the female in A. californicus. The
outlier from Kansas (Wellington, USNM:
2 9,3 6) is typical of the species and de-
finitively identified.

We have seen mostly females of Asaphes
from the Neotropical region, but included
are specimens from the countries listed
above that either greatly increase limits of
variation for A. californicus, A. suspensus
and A. vulgaris, or that represent addition-
al species that are very similar to these
species. Some females look superficially
like A. suspensus females because they
have entirely yellow legs (sometimes also
with the coxae brownish rather than dark
with metallic luster). Others are similar to
A. vulgaris females because they have the
trochanters, trochantelli and femora, and
sometimes also the tibiae and tarsi simi-
larly dark. These, and other females hav-
ing a color pattern more typical of A. cal-
ifornicus, can also have different combi-
nations of the following features: specu-
lum usually similarly broad as for A.
vulgaris, though sometimes with 1-3 setae
somewhere within the bare band; frenal
groove often poorly developed, indicated
only by a faint transverse line or if dis-
tinctly sulcate then not distinctly crenu-
late; mesonotum usually shiny with dis-

224

Journal of Hymenoptera Research

tinct metallic sheen, very finely net-like
coriaceous with bare region of mesoscutal
lateral lobe smooth; petiole often obvious-
ly elongate, 1.25-1.60 times as long as
wide; petiole with strong longitudinal ca-
rinae and irregular surface sculpture to
evenly, finely, longitudinally striate-retic-
ulate without longitudinal carinae. Fe-
males with entirely yellow legs tend to
have a shorter petiole with distinct cari-
nae, and always a broad speculum that
distinguishes them from A. suspensus fe-
males. Females with dark legs tend to
have a more elongate petiole and /or one
that is evenly sculptured without or with
only obscure carinae, and often with three
or more setae within or basally closing an
otherwise broad speculum. Both color
forms usually also have a non-crenulate
frenal groove. Finally, all males seen from
South America, including all those asso-
ciated with any of the different female col-
or forms, have a scape similar in structure
to A. californicus, including a varied sen-
sory region on the outer surface. Because
of what currently appears to be a contin-
uum of states in females and the presence
of some sort of sensory region on the
scape of all males we are uncertain of spe-
cies limits in the Neotropical region. How-
ever, we have not seen any Asaphes from
America south of Mexico that we believe
belong to A. suspensus or A. vulgaris (see
respective sections for these two species).
In addition to the New World we have
seen males with an elongate sensory re-
gion on the outer surface of the scape from
the Oriental region [India (CNCI), Taiwan
(UCRC)] and the Palaearctic region [Iran
(TAMU), Morocco (UCRC), Turkey
(UCRC)]. The males from India and Tai-
wan appear to belong to an undescribed
species based on a much longer flagel-
lum — the combined length of the flagel-
lum and pedicel in these specimens is
more than 2.5 times the length of the scape
and distinctly longer (more than 1.2 times)
than the width of the head, and all but the
apical one or two segments are monili-

form to slightly longer than wide. Males,
and associated females, have yellowish or
at least lighter-colored trochanters con-
trasting with variedly infuscate or dark
femora, a coloration similar to that in A.
californicus. The females could be mistaken
for those of A. vulgaris but both females
and males have the speculum closed ba-
sally by setae or have a few setae within
the speculum. Those males from Iran, Mo-
rocco and Turkey with an elongate sen-
sory region on the outer surface of the
scape are otherwise very similar to males
of A. suspensus, including having a similar
forewing setal pattern, entirely yellowish
legs, and short antenna with the pedicel
and flagellum, just the pedicel, or only the
apex of the pedicel yellowish. Further,
some males from Iran were reared along
with typical males of A. suspensus. We
could not find differences in associated fe-
males. It is unknown whether the sensory
region on the scape of some males in the
middle east and northern Africa indicates
a separate species or whether scape struc-
ture is more variable for A. suspensus in
these regions.

Asaphes hirsuttis Gibson and Vikberg,
new species

(Figs. 5, 6, 17, 18, 23, 24, 35, 36, 45, 53,
54, 57, 61, 62, 71, 74)

Type material. — Holcti/pe, female (CNCI, Type No.
22266): CANADA, British Columbia, Cassiar High-
way, Boyar Lake to Stikene River, 6.VI11.1988, S.&J.
Peck, day car netting boreal forest. Allotype, male
(CNCI): same data as holotype. Paratypes: CANADA.
Yukon Territory: Dempster Hwy, 28.VI-2.VIII.82,
D.M. Wood (2 9, 1 6). Heynes ^nction 10, Deza-
deash Lake, 700-900 m, 2.VIII.89, S.&J. Peck, car net-
ting boreal dry forest (1 9). Herschel Island, 16-24,
26.V1I.71, W.R.M. Mason (1 9, 1 S). Ross River,
16.VI-31.VIII.84, S.&J. Peck (3 9, 1 UCDC). North-
west Territories: Banks Island, Masik River,
10.VII.68, W.M. Mason (1 9). Salmita Mines, 64"05'N
liri3'W, 18.V1.33, J.G. Chillcott (1 9). British Co-
lumbia: same data as holotype (11 9, 5 6). Anahim
Lake to Redstone, 1000-1500 m, 17.V1I.88, S.&J. Peck,
car netting pine sand land (2 9, 3 S). Charlie Lake,
25.V1II.80, R.J. Cannings (1 9, SMDV). Manning Pro-
vincial Park, 2 km N Blackwall Peak, 49°07'N
12r45'W, 2000 m, 9.VI1,96, H. Goulet, subalpine

Volume 7, Number 2, 1998

225

meadow (1 6). McLeod Lake, 12.VII.66, #P66-7-12a,
P. A. Rauch, Castilleia (1 6, EMEC); #F66-7-12e, E.G.
Andrews, Castilleia (1 S, EMEC). Manning Provincial
Park, 1400m, 12-14.VII.88, H. Goulet (2 9 ). Mt. Re-
velstoke, 51°02'N 118° 05'W, 1800m, 30.VII.86, H.
Goulet (1 9 ). Mt. Seymour, N Vancouver, 790 m,
17.V.73, J.R. Vockeroth (1 6). Ocean Falls, 13.VII.60,
E.I. Schlinger, 60-7-28P, secondary parasite reared
from Aphiciius sp., host aphid Neomyzaphis iilnetina (4
9, 4 c?, UCRC). Squamish, Diamond Head Trail,
4600', 10, 11, 29, 30.VIII.53, W.R.M. Mason (8 9, 2 cj).
Summerland, 5.VI.59, R.E. Leech (1 6). Terrace,
18.VI.60, J.G. Chillcott (1 9). Vancouver Island, For-
bidden Plateau, W of Courtenay, 1.VII.66, #F66-6-29b,
on Olopanaz homdiim (1 9, 1 6, EMEC), #F66-7-lb,
ex. aphids on Achly^ tnphylla (1 9, 1 c5, EMEC), #F66-
7-le, ex. Macrosiphini on Vaccinium ovalifolium (1 S,
EMEC), #F66-7-lf, 2 mi. NW of Courtenay, on man-
zita-like Ericaceae (2 6 , EMEC), E.G. Andrews. Van-
couver Island, 12 mi. E Port Alberni, 30.VI.66, P.
Raunch, P66-6-30A, Woodumniia sp., Macrosiphum
ptcriiits (1 9, 1 (J, EMEC). Winfield, 20.III.54, F.I.S. no.
541, PiHHS pondemsa cone (1 6). Alberta: Aspen
Beach, 25.VIII. 44, O. Peck (1 6). Edmonton, 11, 16,
22, 23, 24.V.46, 27.V.46, 2, 6, 17.VI.46, W.R.M. Mason
(10 9, 8 6); 24.IX.86, Jorgensen & Anderson (5 3).
Fortress Mountain, 6 km S, 51°52'N 115°10', 1700 m,
21.VI.86, H. Goulet (2 9). Fort McMurray, 30.VI.53,
G.E. Ball (1 9). Seebe, reared from duff layer under
Pinus contorta var latifclia, em. 11.1.71, L.S. Skaley (2
6, NFRC). W Waterton Lake National Park, 49°05'N
113°52'W, 1300 m, 6.V1I.91 (1 9), 3 km E Cameron
Lake, 49U3'N 114°0rW, 1650 m, 13-14.VII.91 (1 6),
H. Goulet. Saskatchewan: Prince Albert National
Park, 23.VII.54, ex. Cimra hottesi (G. & P.), Picea mar-
lana (1 9). Snowden, 26.VII.44, O. Peck (1 9). Mani-
toba: Winnipeg, coll. 31, 31-114, em. 28.VIII.86, H.G.
Wylie (1 9, EDUM). Quebec: Chimo, 17-18.VIII.59,
W.R.M. Mason (2 9, 4 cJ). Great Whale River, 11. VII,
31.VIII.49, J.R. Vockeroth (2 6 ); 8.V1II.59, sand dunes,
W.R.M. Mason (1 6). Louvicourt, 16 km S, km 473.5,
17.VI.85, H. Goulet & D.R. Smith (6 9, 5 <J). Pare de
la Gaspesie, Mont Jacques-Cartier, 29.IX.91, C. Isabel,
zone subalpine, sphaigne humide et krumholz (1 9).
Mont Saint Marie, Low, 1800', 20.IX.65, J.R. Vocker-
oth (1 6). Quebec City area, 29.VIII.88, reared ex.
mummy Aphuinis nigripcs in potato field (1 9). St.
Gedeon, 26. VIII. 76, Remaudiere, ex. Calaphi» on Betula
(3 6, BMNH). Ste-Catherine (de Portneuf), 5.VIII.87
(2 9), 18.VIII.87 (1 9), J. Brodeur. New Brunswick:
Fredericton, 9.Vin.50, Myzus persicae on potato {\ 3),
24.VIII,50, Macroiiphum stilanifolii on potato (1 9 ), B.P.
Spicer; 4.X.50, J.B. Adams, Euladuuis agitis on Scotch
pine (1 6). Kouchibouguac National Park, 17.VII.77,
I. Smith (1 9); 20.IX.77, S.J. Miller (1 9). St. John Co.,
Mary Pitcher Lake, IV.97, R'rd D. O'Shea, aphid on
Picea glauea needle (1 9 ). Prince Edward Island: Har-
rington, 23-29.VIII.86, M.E. Smith, in potato field (1

9). Nova Scotia: Coldbrook, 2, 7, 16, 30.X.63, H.B.
Specht, host Aeyrtliosiplium pisum (Harris) on alfalfa
(1 9, 5 d). Mantua, 23.X.63, H.B. Specht, host Acyr-
thoiiphum pisiini (Harris) on alfalfa (1 9 ) Newfound-
land and Labrador: Junction Pond, Notre Dame Cp.,
19.VII.61, CP. Alexander (1 9, USNM). Labrador,
Goose Bay, 7.V1I.52 (19).

USA. Alaska: Bering Sea, St. Paul Island, T. Kin-
caid (1 9, 1 6, USNM), 16.VIII.15, G.D. Hanna (1 9,
USNM). Cape Thompson, 25.VII.61, R. Madge (1 9 ).
Cartwell, Denali Highway Route #8, mi. 85-130,
24.VII.84, S.&J. Peck (9 9,7 6). Cold Bay, 163° W,
26.VII.52, W.R. Mason, on tundra (1 9). Colorado
Creek, 60"40'N 149°30'W, 9.VII.94, ex. aphid mum-
mies on Alnuf sp., D. Collet (1 9, DCPC). Dave's
Creek, 60°30'N 149°45'W, 3.1X.94, swept from Salix
baniayi, D. Collet (1 6, DCPC). Deering, 8-19.VIII.68,
J. Matthews (1 9, 1 S). Isabel Pass, mi. 206, Richard-
son Hwy, 2900', 17.VII.62, P.J. Skitsko (1 9). Kasilof,
60°15'N 151°15'W, 27.VI.94 (2 9, 2 6, DCPC),
30.VII.94 (2 9, DCPC), ex. aphid mummies on Salix
barclayi, D. Collet. Kenai Peninsula, trail to Bryan gla-
cier, vicinity Portage Glacier, 22.VI1.78, P. H. Arnaud,
Jr. (1 9, CASC). King Salmon, Naknek River,
15.VII.52, W.R. Mason (1 9). Kotzebue., 420,
14.VIII.58, Lindroth (1 9). Matanuska, 6.X.45, J.G.
Chamberlin (2 9, USNM). Steese Highway, mi. 96.4,
4.IX.48, G. Jefferson (1 9, USNM). Soldotna, 60°30'N
151°00'W, 16.VI1.95, ex. aphid mummies on Betula, D.
Collet (2 9, DCPC). Sterling, in front of Collet house,
60°30'N 150°45'W, 23.VII.94 (1 9, DCPC), 31.VII.94 (1
6 , DCPC), ex. aphid mummies on Salix barclayi, D.
Collet; Gann site, 60°30'N 150°45'W, 4.V.93, swept
from Salix barclayi (1 9, DCPC), 3.V.93, swept from
Betula nana (1 9, DCPC), D. Collet. Unalakeet,
27.VI.61, B.S. Hemming (1 9); 15.VII.61, R. Madge (1
9). Unknown locality, 15.VIII.94, D. Collet (1 9,
DCPC). Arizona: Cochise Co., Chiricahua Mountains,
Rustler Park, 8000', 15.VIII.82, G.A.P. Gibson (2 9 );
12 km S Sierra Vista, Ramsey Canyon, 1700 m, 1986,
B.V. Brown (5 i). Tucson Co., Catalina Mountains,
Mt. Lemmon, Crystal Springs Canyon, 8000',
11.VII.90, L. Masner (1 9). California: Glenn Co.,
Mendocino National Park, Plaskett Meadows, 7000',
28.VI.81, J.B. Whitfield (1 9, EMEC). Lake Tahoe,
Pope Beach, 26.IX.75, E.G. Toftner & R.O. Schuster (1
9). Nevada Co., Sagehen Creek, 1.VII.70, Salix, E.E.
Grissell (1 6, UCDC). Yosemite National Park, near
Dog Lake, em IX.59, ex. Essigella mummy, Pinus con-
torta var. murrayana Englm. (1 d, UCRC); near Gaylor
Lakes, X.59, Pinus contorta var. murrayana Englm. (1
6, UCRC). Colorado: Boulder Co., Chataqua Pk. W
Boulder, 4.VI.90, S.L. Heydon (1 9, UCDC). Echo
Lake, Mt. Evans, 10600', 8.VII.61, S.M. Clark (2 6).
Estes Park, 11.VII1.53, R.R. Dreisbach (19, USNM).
Fort Collins, 86 km W, 4.VIII.72, R.B. Penfield, Hop-
kins US no. 36751-V-198, Arceuthobium cyanocarpum (1
6, USNM). New Castle, Hopkins US no. 34211-K, lot

226

Journal of Hymenoptera Research

no. 47-946, Picea engehmnni (1 2, USNM). Pitkin Co.,
10,5 km SE Aspen, 2900m, 9.VII.92, S.L. Heydon (2
6. UCDC). Idaho: Clarkia, 24.V.60, R.E. Denton,
Hopkins US no. 20366, Pimis moiiticohi foliage (1 6,
USNM). Shoshone Co., Thompson Pass Summit,
29.VI1.95, K.S. Pike, ex. AiJhidius ptoli/gcnapiliis or Praon
Immutaphidis from Ulinoia sp. on Vacciniuin sp. (1 9,
WSUC). Maine: Aroostook Co., 9.VI, 10, 21, 31. VII, 4,
10.IX.58, ex. Macrosiphum solanifolii (3 9, 4 3, USNM).
New Hampshire: Mount Washington, 4.V1II.50, S.
Ristich (1 9); 1676m, 20.V1I1.81, H. Goulet (1 9); Al-
pine Garden, 5200-5600', 7.VI1I.54, Becker, Monroe &
Mason (1 9, 1 d); Cow Pasture, 5700', 2.VI1I.54, Beck-
er, Monroe & Mason (1 9); Lakes of the Clouds,
5000', 3, 9.V11I.54, Becker, Monroe & Mason (2 9,1
6); Tuckerman's Ravine, 31.V11.54, Becker, Monroe &
Mason (1 d). New Mexico: Lincoln National Forest,
Karr Canyon, 28.V11.77, L. Masner (1 9, 1 S). Otero
Co., 2 mi. N Cloudcroft, 5.V1.80, S.L. Heydon (1 9,
UCDC). Valencia Co., 20 mi. W Los Lunas, Corritzo
Arroyo, 1-23.VII1.77, S.&J. Peck (1 9 ). Oregon: Klam-
ath Co., Sprague River, Hwy 87, Collier, 2.V1I.85,
D.G. Denning (1 i ). Washington: Benton Co., WSU-
Prosser Roza Unit, 27.V1.94, K.S. Pike, ex. Aphiiiius
ervi from Sitobion avenae on Tritkum aestivum (wheat)
(1 6, WSUC). Clallam Co., 10 mi. E Pysht, 26.VI.66,
#C66-6-26a, D. Calvert (1 9,1 6, EMEC); Lake Oz-
ette, 24.V11.90, J.D. Pinto (2 9,16). King Co., Stam-
pede Pass lookout point, 8.V1I1.96, K.S. Pike, un-
known on Rhmlodcndrcii albiflonim (white-flowered
azalea) (1 9, WSUC). Kittikas Co., Lost Lake,
6.V1II.97, K.S. Pike, ex. Aphidius sp. from Ulinoia sp.
on Lonicera involucrata (black twinberry) (2 9,
WSUC); Manashtash Cr., mile 8, 16.V111.96, K.S. Pike,
ex. Alloxysta sp. or Aphidius ohiocnsis or Dendroccnis
sp. or Ephednis cnlifoniicus from Mncwsiphutn creelii on
Vicia sp. (vetch) (2 9, WSUC); Manashtash Cr.,
milel2, 25.VI.97, K.S. Pike, ex. Alloxysta or Aphelinus
on Lonicera involucrata (1 9, WSUC); 2 mi E. of Quartz
Mtn, 25.V11.96, K.S. Pike, ex. Aphidius sp. or Praon sp.
from unknown aphid on Rubus lasiococcus (dwarf
bramble) (19). Lewis Co., Rainier National Park, Tip-
soo Lake, 11.IX.96, K.S. Pike, from Macrosiplnim sp.
on Rhododendron albiflonim (white-flowered azalea) (1
9, WSUC). Mount Baker, 2.V111.86, 1700 m, H. Goulet
(2 9 ). Mount Rainier National Park, Van Trump Park,
1500-1800 m, 29.V1I.85, L. Masner (8 9, 17 6). Pierce
Co., Mount Rainier National Park, Chinook Pass,
8.1X.95, K.S. Pike, unknown on Spiraea densiflora (1 9,
1 6, WSUC). Wenatchee, 22.V1I.83, D. Carroll, host
Schizapltis graminum via Praon sp. on Agropyron repens
(1 9, Carroll); near bridge, 17.V1.82, D. Carroll, host
Rhopalosiphuni insertuni via Praon uincum on Crataegus
douglasii (1 9, Carroll). Yakima Co., 15 mi. E of Chi-
nook Pass, 11.1X.96, K.S. Pike, free flying parasitoids
on Achlys triphylla (vanilla-leaf) (19, 1 S, WSCU);
Green Lake, 20.VII.94, K.S. Pike, ex. Alloxysta sp. or
Monoctonus sp. from Ulinoia sp. on Rhododendron al-

bifolorum (white-flowered azalea) (2 9, WSUC); Green
Lake Road, 31.V11I.94, K.S. Pike, ex. Blacus sp. or ich-
neumonid, from Cinara cliinookiana or unknown on
Abies laswearpa (subalpine fir) (5 9, WSUC); trail, 1.5
mi. from RdlOlO, 14.1X.95, K.S. Pike, ex. Alloxysta sp.
or Lysiphtebus testaceipjes from Aphis fabae on Cirsium
vulgare (bull thistle) (13 9, 10 (J, WSCU); Yakima In-
dian Reservation, Howard Lake, 19.1X.95, K.S. Pike,
ex. Aphidius pwlygonaphis or Praon sp. from Sitobion sp.
on Pteridium aquilinum (2 9, WSUC). Wyoming: Big-
horn Co., Northern Bighorn Mountains, Sheep Moun-
tain, 2800 m, 22.Vn.88, H. Goulet (1 S).

EXTRALIMIJ AL~Neotropical: MEXICO. Rio Frio,
3150 m, 8.V.79, G. Remaudiere, ex. Macrosipjhum on
Eupihorbia peplus (5 9, 4 c5 glued on 6 cards along with
specimens of A, californicus, BMNH). Palaearctic:
AUSTRIA. Dolomiten, S.-Tirol, Platzwiese, 200 m,
Diirrnstein Geb., 1968, Schimitschek, ex. Pauesia sim-
ilts Sta. in Cinara ceinbrae (1 9, BMNH). CZECH RE-
PUBLIC. Bohemia: Studnice, nr Jablonec nad Nisou,
860m, 28.V1.63, V. Martinek (19). Teplice, Nove Mes-
te, 850m, 2.V11.63 (1 6), 12.V11.63 (I 6), 30.V1I.63 (2
(J), V. Martinek. FINLAND (FENNIA, SUOMl). Ka-
relia borealis [= Kb]: Eno, Ahveninen, 698:65,
22.VI11.68, V. Vikberg (1 9, WPC). Osfrobottnia me-
dia [= Om, KPl: Kestila, 7135:461, 29.VI.78, M. Ko-
ponen (1 9, DAZH). Osfrobottnia borealis, N part
[= ObN, PP]: Ranua, 7312:479, 20.V1I.80, M. Koponen
(1 6. DAZH). Rovaniemi, 737:44, 16.VI11.80, J. Halme
(5 9, 1 6, DAZH). Kuusamo [= Ks|: Kuusamo, 735:
61, 2.V1I.79, V. Vikberg (3 6, WPC); 7366:603,
26.VI.79 (1 9), 30.V1.79 (4 9, 3 cJ), 26.V1.82 (1 9), M.
Koponen (DAZH). Lapponia kemensis, E part [ =
LkE, KemL|: Sodankyla, 7481:467, 11.V111.83 (1 6),
7582:516, 6.V11.89 (1 9), M. Koponen (DAZH). Lap-
ponia kemensis, W part [= LkW]: Kolari, 7501:382,
6.V11.97, M. Koponen (2 9, 1 6, DAZH), 7504:379,
4.V1I.97, K. Silvonen (1 9, DAZH); Kolari, Yllas, 7502:
380, 29.VI.97, V. Vikberg (1 9, WPC); Kolari, Yllas,
Varkaankuru, 7502:382, V. Vikberg (19, WPC). Mu-
onio, Olostunturi, 7541:366, 28.VI.97, V. Vikberg (1 9,
WPC). Lapponia enontekiensis [= Le, EnL]: Hetta,
759:36, 3, 4.V11.82, V. Vikberg (2 9, WPC). Karesu-
vanto, 760:31, 29.V1.82, V. Vikberg (1 9, WPC); 7605:
313, 29.V1.82 M. Koponen (2 9, DAZH). Kilpisjarvi,
Pien Malta, 2.V11.50, W. Hellen (1 9, UZMH); region
subalpina, 767:25, 17.V1.71, V. Vikberg (1 9, WPC);
Siilastupa, 1.V11.50 (1 S, UZMH), 10-19.V1I.50 (1 9,
UZMH), W. Hellen. Ropinsalmi, 2.V1I.82, Y. Zhongqi
(1 9, DAZH). Lapponia inarensis [= Li, InL]: Inari,
7591:478, 1.V1I.89 (1 9), 7617:521, 6.V1I.89 (2 9, 2 6),
7664:504, 12.VII1.83 (2 9), 7615:517, 4.VI1.89 (2 9), M.
Koponen (DAZH); Inari, kk [= kirkonkyla = 'church
village'], 24.V1.60 (1 9), 28.V1.60 (1 9), 29.VI.60 (2 9),
V. Vikberg (WPC); Inari, Kaunispaii, 154/80, 759:50,
12.VI11.80 (1 9, 1 6), 159:51, 11.VI1I.80 (1 9), 159:51,
15.VI11.80 (2 S ), J. Halme (DAZH); Inari, Opukasjarvi,
772:55, 4.V1I.60, V. Vikberg (1 9, WPC). Utsjoki,

Volume 7, Number 2, 1998

227

7741:500, 13.VIII.83, M. Koponen (1 9, DAZH); Uts-
joki, kk [= kirkonkyla = 'church village'], 775:50,
4.VI.60 (1 V), 6.VI.60 (2 9, 1 6), 11.VI.60 (1 9),
16.VI.60 (4 9, 4 (J), 18.VI.60 (1 9, 3 d), 19.VI.60 (2
9 ), 22.VI.60 (2 9 ), 29.V1.60 (2 9 ), V. Vikberg (VVPC);
Utsjoki, Karigasniemi, 770:46, 30.V1.60, V. Vikberg (1
9, VVPC). NORWAY. Oppland [= O]: Dovre, Foks-
tua, 13.VII.53, W. Hellen (1 9, UZMH). RUSSIA.
Khabarovsk Kray: Ochotsk [= Okhotsk], 1841, F.
Sahib. 1= Sahlberg] (1 9, UZMH). Kamchatka
Oblast: N. Kuril Islands, 5.VIII.64, aphid on Sulix (3
6, ZMAS). Murmansk Oblast: Ylaluostari, VI1.30, W.
Hellen (1 9, UZMH). SWEDEN (SUECIA). Norr-
botten (= Nb.]: Boden, Trehorningen, RN 1783/7326
810722-22 [Equisetum siliuiticiim; Sitchioii cqiiiscti;
Aphidiuf picipes], U. Gardenfors (1 9, MZLU). Lulea,
Lulviken, RN 1793/7286 810721-30, [Ephedni^ mum-
my on needle of Piiun a/h'cstris on ground] (1 9,
MZLU), RN 1793/7286 810721-11, [Wahlgrenicllii vac-
cinii; Praon inyzophnguiii, plus Ephedrus plngiator, plus
Aphidiui sp.; Aphelinui sp.j (2 9, 1 (J, MZLU), U. Gar-
denfors. Lulea, Svartostaden, RN 1794/7288 800807-
08, [Epilohium aiigustifoliuin; Mncrosiphuiii rosae; Aplii-
dius sp., plus Ephedrus sp.], U. Gardenfors (1 9,
MZLU). overkalix, RN 1815/7376 810722-08, [Rosa
sp.; Macwsiplnim rosae; Praon sp.] (1 S , MZLU), RN
1815/7376 810722-02, [Lomcera sp. culture; Rhopnlo-
niyziis lonicerac; Ephedrus sp.] (3 9, 8 6, MZLU), U.
Gardenfors. Tome Lappmark ]= Lpl.]: Abisko,
15.VIII.51, J.R, Vockeroth (1 9).

Etymology. — From the Latin hirsuhis,
hairy, in reference to the partially setose
metapleuron.

Female. — Head and mesosoma dark,
usually with obscure metallic green luster
under some angles of light; legs entirely
dark or with at least coxae, trochanters
and most of femora dark, the trochantelli
sometimes, femora apically, and tibiae
and tarsi often yellowish or distinctly
lighter in color than rest of legs. Head
transverse-subtriangular in frontal view
(Fig. 5), width at least 1.25 times height,
and in lateral view lower face evenly
curved into upper face (Fig. 6); interorbital
region in dorsal view relatively shallowly
concave (Fig. 17); gena length about 0.64-
0.82 eye width and 0.53-0.66 eye length;
dorsal margin of torulus approximately in
line with lower orbit (Figs. 5, 6). Antenna
(Figs. 23-24) with pedicel length about
2.5-3.0 times greatest width; funicle with
fu, transverse, fu,-fu„ transverse to very

slightly longer than wide {iu^-hi^ or fu^
usually more or less moniliform), fuy
quadrate to slightly transverse, and fUg
transverse. Mesoscutum (Fig. 45) with lat-
eral lobes broadly bare medially, and with
fine engraved net-like sculpture over at
least posterior half of bare area. Scutellum
(Fig. 45) mostly bare except along extreme
anterior and lateral margins; frenum
broadly smooth and shiny or with lateral
longitudinal rugae continued narrowly
along posterior margin and with fine, en-
graved, net-like sculpture except for an-
teromedial smooth region (Figs. 53, 54).
Metapleuron with at least a few setae in
anteroventral angle and often extensively
setose (Figs. 61, 62). Forewing with basal
cell often evenly setose (Fig. 71) but at
least setose apically and with one or more
lines of setae along length; disc either
without evident speculum (Fig. 71) or
speculum narrow, closed basally by setae
and with line or lines of setae close to sub-
marginal vein. Petiole in dorsal view
slightly transverse to longer than wide,
length 0.8-1.2 times width, reticulate with
irregular longitudinal carina or stronger
keels (Figs. 45, 53).

Male. — Color pattern similar to female
except legs sometimes more extensively
light-colored, rarely almost uniformly yel-
low beyond coxae, the femora only slight-
ly infuscate; antenna uniformly dark.
Scape (Figs. 35, 36) length about 3.7-4.5
times greatest width, often widest subbas-
ally but at least slightly tapered to apex,
and in lateral view inner surface evenly
setose but with flat to slightly concave
ventrally or externally angled, sparsely se-
tose or bare and shiny, surface over at
least apical two-thirds (Fig. 36a). Pedicel
(Figs. 35, 36) length about 2-2.25 times
width and about 0.36-0.47 scape length.
Combined length of pedicel and flagellum
less than 2.5 times scape length, and fla-
gellum at most as long as width of head;
funicle (Fig. 35) with all segments slightly
to distinctly transverse or with one or
more of fu,^ slightly longer than wide.

228

Journal of Hymenoptera Research

quadrate, or moniliform (segments in-
creasingly more transverse apically).
Structure otherwise similar to female ex-
cept petiole always at least slightly longer
than wide, about 1.1-1.66 times as long as
wide (Fig. 57). Setal patterns similar to fe-
male except more commonly with only a
few setae in anteroventral angle of meta-
pleuron.

Distribution. — Holarctic; in North Amer-
ica transcontinental across the Boreal re-
gion and extending south within and west
of the Rocky Mountains in the USA (Fig.
80) and into Mexico along the Sierra Ma-
dre Occidental.

Biology. — All host records indicate that
A. hirsutus is a hyperparasite of aphids, in-
cluding: Acyrthosiphon pisum (Harris),
Aphis fabae Scopoli, Calaphis sp., Cinara
cembrae (Seitner), Cinara chinookiana Hottes ?,
Cinara hottesi (Gillette & Palmer), Elato-
bium abietinum (Walker), Essigella sp., Eu-
lachnus agilis (Kaltenbach), Illinoia sp., Mac-
rosiphum creelii Davis, Macrosiphum eiiphor-
biae (Thomas), Myzus persicae (Sulzer),
Rhopalosiphtim insertum (Walker), Sitobion
avenae (Fabricius), Sitobion pteridis (Wil-
son), and Schizaphis graminum (Rondani).
Label data also indicate species of Praon
and Aphidius (Braconidae), and possibly
Alloxysta, Blacus, Lysiphlebus, Ephedriis,
Monoctonus (Braconidae) and Dendrocerus
(Megaspilidae), as host primary parasites.

Remarks. — In addition to the paratypic
material listed above, in 1984 the junior
author examined two females and two
males of A. hirsutus from the Greenland
Hymenoptera Collection of the Zoological
Museum, University of Copenhagen, Den-
mark. Lundbeck (1897) collected these
specimens in 1889 from Arsuk and Ser-
miligarssuk fjord (= Sermiliarsuk) in
southwestern Greenland. The four speci-
mens had 1955 determination labels by O.
Bakkendorf, as A. vulgaris. At the time of
preparation of this manuscript these spec-
imens could not be located in the museum
(N. Kristensen, pers. comm.). However,
they formed the basis for the literature re-

cords of A. vulgaris in GreerJand cited in
Bakkendorf (1955). It is possible that all rec-
ords of A. vulgaris from Greenland are
based on misidentifications of A. hirsutus.
The setose metapleuron (Figs. 61, 62)
readily distinguishes most males and fe-
males of A. hirsutus from most other spe-
cies of Asaphes in the Nearctic region.
Those specimens with only a few setae
within the anteroventral angle of the me-
tapleuron are very similar to individuals
of A. petiolatus but females of A. petiolatus,
at least in the Nearctic region, have a dis-
tinctly sculptured frenum (Fig. 52). In ad-
dition to the absence of any setae on the
metapleuron, males of A. petiolatus also
have a slightly different structure of the
scape. In lateral view the scape is more
elongate-slender, without a distinct ven-
trally or externally angled surface but
with a longitudinal, bare, shiny band on
the inner surface, at least in larger speci-
mens {cf. Figs. 36, 38). Some A. brevipetiol-
atus females also have setae on the meta-
pleuron but are distinguished by head
structure from A. hirsutus females. The
combination of almost uniformly setose
forewing and dark trochanters will differ-
entiate females from those of A. vulgaris
and A. californicus, respectively. Because of
its setose metapleuron, setose forewings,
and relatively dark legs, A. hirsutus is also
very similar to the Japanese species A. pu-
bescens Kamijo and Takada, but is distin-
guished by having the mesoscutal lateral
lobes broadly bare medially (Fig. 45) rath-
er than evenly setose. We saw three fe-
males from Nepal (CNCI) with the meta-
pleuron entirely setose, which closely re-
semble A. pubescens because they have the
mesonotal lateral lobes evenly setose.
These specimens likely belong to an un-
described species because the flagellar
segments, including fu,, are all at least
slightly transverse.

Asaphes petiolatus Zetterstedt,
revised status

(Figs. 25, 26, 37, 38, 46, 51, 52, 58, 65, 72, 75)

Asaphes petiolatus Zetterstedt 1838; 423. Type
data: Lapponia [Swedish Lapland]: Wittangi.

Volume 7, Number 2, 1998

229

Sex described: female. Holotype by mono-
typy; MZLU.

Female. — Head and mesosoma black
with obscure metallic green luster under
some angles of light [some western Euro-
pean females with distinct metallic green
luster]; legs entirely black or, more often,
black to dark brown with extreme apex of
femora, extreme base and apex of tibiae to
entire tibiae, and tarsi lighter in color, ru-
fous to yellowish. Head transverse-sub-
triangular in frontal view, width at least
1.25 times height, and in lateral view low-
er face evenly curved into upper face; in-
terorbital region in dorsal view relatively
shallowly concave; gena length about
0.72-0.85 eye width and 0.57-0.65 eye
length; dorsal margin of torulus approxi-
mately in line with lower orbit. Antenna
(Figs. 25, 26) with pedicel length about
2.0-2.5 times greatest width; funicle with
fu, ring-like, fu, quadrate to slightly lon-
ger than wide, fu^-fu^ or fuj-fu^ subquad-
rate, but at least fu^ and fu„ transverse.
Mesoscutum (Fig. 46) with lateral lobes
broadly bare medially, and with fine en-
graved net-like sculpture over bare area.
Scutellum mostly bare except along ex-
treme anterior and lateral margins (Fig.
46); frenum with distinct, uniform, en-
graved net-like sculpture similar to scutel-
lum (Figs. 51, 52) [some specimens from
western Europe with frenum medially or
mostly shiny and smooth except for very
fine and obscure net-like sculpture]. Me-
tapleuron bare (Fig. 65). Forewing with
basal cell evenly setose (Fig. 72); disc with
speculum closed basally by setae and with
line or lines of setae close to submarginal
vein (Fig. 72). Petiole in dorsal view var-
iedly distinctly, almost always definitely
transverse (Fig. 51), width 1.15-1.3 times
length, reticulate with irregular longitu-
dinal carinae or stronger keels.

Male. — Color pattern similar to female
[western European specimens sometimes
with head and mesosoma having distinct
metallic green luster; legs usually more

extensively light-colored, sometimes al-
most entirely yellow except metafemur
partly infuscate]; antenna uniformly dark.
Scape (Figs. 37, 38) length about 4.5-5.5
times greatest width, spindle-shaped with
ventral and dorsal margins symmetrically
tapered to apex; in lateral view with flat
ventral surface differentiated only near
pedicel, and at least larger individuals
with elongate, bare, smooth band over
most of inner surface (Fig. 38b). Pedicel
length about 2.0-2.5 times width and
about 0.36-0.42 scape length (Figs. 37, 38).
Combined length of pedicel and flagellum
less than 2.5 Hmes scape length, and fla-
gellum length slightly less than head
width; funicle (Fig. 37) with at least fu^
and fus subquadrate to moniliform, and
sometimes with fu, longer than wide and
fu, s subquadrate to moniliform. Structure
otherwise similar to female except malar
space about 0.62-0.74 eye width and about
0.51-0.54 eye length; petiole subquadrate
to definitely longer than wide, but length
less than 1.25 times width (Fig. 58). Setal
patterns similar to female.

Distribution. — Holarctic; in North Amer-
ica restricted to the Boreal region (Fig. 79)
and not commonly collected. CANADA.
Northwest Territories: Kovaluk River
[69°11'N 131°W], 2-6.VlII.71, W.R.M. Ma-
son (1 9 ). British Columbia: Upper Car-
manah Valley, UTM lOU CJ 801991,
28.V11.92, N. Winchester (2 6, PFRC). Al-
berta: Edmonton, 10.IX.86, A.T. Finna-
more (1 9); 24.IX.86, Jorgensen & An-
drews (1 6, used for SEM). Saskatche-
wan: Prince Albert National Park,
23.VII.54, ex. Cinara twttesi (G. & P.) (2 9,
1 used for SEM). Hudson Bay, 15.IX.59,
J.R. Vockeroth (2 6). Manitoba: Wark-
worth Creek near Churchill, 21. VI. 52, J.G.
Chillcott (1 9). Ontario: Wawa, 2.VII.57,
Forest Insect Survey record no. 710 (1 9 ).
Quebec: Lac Brule, 7.VIII.45, O. Peck (1
9 ). USA. Alaska: Sterling, in front of Col-
let house, 60°30'N 150°45'W, 3.VIII.94, ex.
aphid mummies on Picea glauca, D. Collet
(1 9, DCPC). EXTRAl'iMITAL. FIN-

230

Journal of Hymenoptera Research

LAND (DAZH, WPC), ITALY (BMNH),
SWEDEN (CNCI, MZLU), SWITZER-
LAND (BMNH).

Biology. — Apparently a hyperparasite of
aphids in North America, including Cinara
hottest (Gillette & Palmer).

Type material examined. — The holotype
female of A. petiolatus is glued by its me-
tasoma to the side of a pin that also bears
the holotype of Pteromalus violaceus Zetter-
stedt (see Graham 1969: 81). It is entire,
except for the right antenna beyond the
pedicel, and has the following features:
frenum almost entirely coriaceous except
for narrow median smooth band that does
not quite extend to the posterior margin
of the frenum; speculum closed basally by
setae and with four, almost evenly spaced,
setae behind the submarginal vein that are
separated from the vein by a distance less
than the length of a seta; petiole about 1.2
times as wide as long; and head and me-
sosoma with only relatively obscure me-
tallic green luster under some angles of
light.

Remarks. — We have seen too few speci-
mens of this species to estimate limits of
variation reliably, both within the Nearctic
region and across its known range. All
Nearctic females seen had the frenum dis-
tinctly sculptured whereas some females
from western Europe have a shiny, only
very finely and obscurely sculptured fre-
num. Males from both regions have the
frenum quite shiny, smooth medially
though finely sculptured paramedially.
Such males from the Nearctic had only
very obscure metallic green luster and
dark legs, whereas those from western Eu-
rope were distinctly metallic green and
had the legs almost entirely yellow except
for a partly infuscate metatrochanter and/
or metafemur. These males could easily be
mistaken for males of A. suspeiisus because
of their indistinct speculum and shallowly
concave interorbital region, but the anten-
na is uniformly dark and the scape and
basal flagellar segments are more elongate
than for males of A. suspeusus {cf. Figs. 37,

39). Most males and females of A. petiola-
tus, particularly those with the frenum
more or less smooth and shiny medially,
are more likely to be mistaken for those of
A. vulgaris. However, in addition to hav-
ing a narrow, closed speculum, A. petiola-
tus also has the interorbital region more
shallowly concave {cf. Fig. 17) and usually
has an obviously shorter petiole than A.
vulgaris (cf. Figs. 58, 60). In both species
the petiole of the female is shorter relative
to the male so that females of A. petiolatus
usually have an obviously transverse pet-
iole (Fig. 46) whereas A. vulgaris females
have the petiole at least quadrate and al-
most always slightly to distinctly longer
than wide (Fig. 48). Males of A. petiolatus
usually have a subquadrate to only slight-
ly elongate petiole (Fig. 58), whereas
males of A. vulgaris have a more obviously
elongate petiole (Fig. 60). However, in
both cases the most elongate petioles of
specimens assigned to A. petiolatus ap-
proach the least elongate petioles of spec-
imens assigned to A. vulgaris based on
forewing setal pattern. Structures of the
scape are also similar. Males of A. vulgaris
do not have a distinct bare band on the
inner surface of the scape (cf Figs. 38b,
42b), but because of poor preservation of
specimens we are uncertain whether this
feature is characteristic of and distinct for
all A. petiolatus males. Except for the ab-
sence of any metapleural setae, males of
A. petiolatus are also quite similar to males
of A. hirsutus. The slight differences in de-
scribed scape structure of the two species
help to differentiate those males of A. hir-
sutus that have only a few inconspicuous
setae (see 'Remarks' for A. hirsutus). Only
head structure reliably distinguishes fe-
males of A. petiolatus and A. brevipetiolatus.

Asaphes stispenstis (Nees)
(Figs. 9, 10, 16, 27, 28, 39, 40, 47, 59, 66,

69, 77, 78)

Chr[/solampus fuspeiiaus Nees 1834: 127. Type
data: Germany: Sickershusi province, 2 July
1813; reared from Aphidii rosaruni. Female de-

Volume 7, Number 2, 1998

231

scribed. Lectotype designated by Graham
1969: 82; Hope Entomological Collection, Ox-
ford, England.

Chn/solampiis altiveiitris Nees 1834: 127. Type
data: Germany: Sickershausen [female: 21
April 1811; male: 17 September]. Both sexes
described. Syntypes, lost. Synonymy by Gra-
ham, 1969: 82.

Pteromnlus petioliventris Zetterstedt 1838; 429.
Type data: Lapponia [Swedish Lapland] be-
tween Karesuando and Kengis in August.
Described questionably as male [Graham,
1969: 82 erroneously stated that it was de-
scribed as a female]. Holotype male by
monotypy; MZLU. Synonymy by Graham,
1969: 82.

? Colnx nphidii Curtis 1842: 60. Unknown type
status, lost. Tentative synonymy by Graham
1969: 82, based on original description.

CIm/soIampus nphhiiphagus Ratzeburg 1844: 181.
Holotype, lost. Synonymy by Graham 1969:
82, based on original description; incorrectly
synonymized with A. vulgaris by Kurdjumov
1913: 24.

Chn/solnmpus nphidicoln Rondani 1848: 19-21.
Type data: reared from an aphid [Apihis ro-
sae]. Female described. Lectotype designated
by Boucek 1974: 244; Museo Zoologico 'La
Specola', Florence, Italy. Synonymy by Bou-
cek 1974: 244, 275; incorrectly synonymized
with A. vulgaris by Delucchi 1955: 174.

Euplectrus lucens Provancher 1887: 207. Type
data: Canada: Quebec, Cap Rouge; Ontario,
Ottawa. Female described. Lectotype desig-
nated by Gahan and Rohwer 1917: 399; Univ-
ersite Laval Insect Collection, Quebec City,
Canada, type no. 1369. New synonymy.

Asaphes rufipes Brues 1908: 160. Type data: USA:
Massachusetts, Forest Hills, 30.X.1908, P.
Hayhurst; reared from Aphis, probably A.
atriplicis L., on Chenopodium album. Female
described. Holotype by original designation;
MCPM. New synonymy.

Mcgorismus Fletchen Crawford 1909: 98. Type
data: Ottawa [15 Aug.] Canada [emerged];
bred from [ex.] Nectarophora pisi; Arthur Gib-
son collector. Both sexes described. Holotype
female by original designation; USNM, type
no. 12197. Previous synonymy with A. lucens
by Burks, 1964: 1258. New synonymy.

Asaphes americana Girault 1914[219]: 114. Type
data: USA: Iowa, Hampton, June 1912, R.L.
Webster, exp. 101. Both sexes described. Lec-

totype female hereby designated; USNM,
type no. 15655. Previous synonymy with A.
flefcheri by Burks, 1958: 74 and with A. lucens
by Burks, 1964: 1258. New synonymy.

Pachycrcpoidcs indicus Bhatnagar 1951: 160-163.
Type data: India: Chaubattia (U.P.),
15.V.1946, Z.A. Siddiqi; reared from Aphis
helichrysi Kalt. Female described. Holotype
by monotypy; type depository unknown.
Tentative synonymy with A. vulgaris by Bou-
cek et al. 1978: 437, based on original descrip-
tion and illustrations. New synonymy.

? Asaphes sawraji Sharma & Subba Rao 1958:
181-183. Type data: India: Kalka, Punjab;
reared from Acyrthosiphon (Macrosiphon) pisi
on the garden pea, Lathyrus odoratus L. Both
sexes described. Unknown type status; lARI.
Synonymy by Boucek et al. 1978: 436-437.

Pachyneuron uniarticulata Mani & Saraswat
1974: 96-98. Type data: India: Northwest
Himalayas, DaUiousie (Ahla catchment area),
M.K. Kamath, 25.V.1971. Female described.
Holotype by original designation; USNM.
Synonymy by Boucek et al. 1978: 436-437.

Asaphes vulgaris; McMuUen 1966: 236, 239;
McMullen 1971: 34; Philogene and Chang
1978: 54; Batulla and Robinson 1985: 36. Mis-
identifications of A. suspensus.

Female. — Head and mesosoma with me-
tallic green luster under some angles of
light, and usually bright metallic; legs
usually uniformly light-colored, yellowish
to yellowish-orange, but femora some-
times darker, yellowish-brown, medially.
Head transverse-subtriangular in frontal
view (Fig. 9), width at least 1.25 times
height, and in lateral view lower face
evenly curved into upper face; interorbital
region in dorsal view shallowly concave
(Fig. 16); gena length about 0.6-0.72 eye
width and 0.5-0.65 eye length; dorsal mar-
gin of torulus approximately in line with
lower orbit (Figs. 9, 10). Antenna (Figs. 27,
28) with pedicel length at most about
twice width; funicle with fu, strongly
transverse, fu, ring-like, and usually all
segments at least slightly transverse, but
fui-fus sometimes moniliform. Mesoscu-
tum (Fig. 47) with lateral lobes broadly
bare medially, and with finely engraved

232

Journal of Hymenoptera Research

Table 2. Host information for Asaphes suspensus based on observed specimens; a question mark follows
rearings or identifications indicated as questionable on the labels. Unless otherwise footnoted, all 'aphid or
other hosts' are Homoptera (Aphidoidea: Aphididae) and all 'associated primary hosts' are Hymenoptera
(Braconidae: Aphidiinae).

Aphid or other Hosts

Associated Primary Hosts

Museum
Acronyms

Acauiius convclvuli

Nevsky ?
Atyrtlwsiphon lactucae (Pas-

serini)
A. pisum (Harris)

Amphorophcrn nibi (Kalten-

bach) ?
Aphis atripilicis (L.)
A. briisskcie L.
A. craccivora Koch

A. fabae Scopoli
A. gossypii Glover
A. heliantbi Monell
A. hohdisci Robinson
A. ilUnoisensii Shimer
A. nnstiirtii Kaltenbach
A. rtimicis L.
A. spiraecola Patch ?

A. intis Scopoli
Aphtlmrgelia symphoncarpi

(Thomas)
Brachycaudus luiicbri/si (Kal-
tenbach)

B. (Appelia) tragopogoiiis
(Kaltenbach)

Brachycornclla asparagi

(Modviko)
Braggia sp.
Brevicoryne brassicac (L.)

Calaphis betulaecolens (Fitch)
Cavariella aegopodii (Scopoli)
Chaitophorus salicicola Essig
Chactosiphcii {Pcntatricliopua)

fragacfoUi (Cockerell )
Chromaphis jughmdkola

(Kaltenbach) ?

Aphclinui semiflaviis' Howard

Aphelinus sp.'

Aphidius sp.

A. stnithi Sharma & Subba Rao

Ephednis incompletiis Provancher

Praoii aguti Smith

P. exoletum palitans Muesebeck

T. (Trioxys) complanatus Quilis

T. (Trioxys) pallidits (Haliday)

Prnoii sp.

Aphidius sp., A. ervi Haliday; Praon
sp.

OH

USNM

OH

USNM

CA, NY

CUIC, USNM

CA

EMEC

VA

USNM

CA

EMEC

CA

EMEC, USNM

CA

EMEC

CA

EMEC

NB

CNCI

WA

WSUC

CA, IL, MB, CNCI, EDUM,

NB, NS, OH, EMEC, INHS,

ON, OR, WA OSUC, USNM,
WSUC

WA WSUC

MA

CUIC

CA,

lA, 1

KS

USNM

Lysiphkbus (Phkbus) testnceipcs

WA

WSUC

(Cresson)

Lysip'hk'btis sp.

WA

WSUC

CA,

DC

UCRC, USNM

Pracn sp., P. unicum Smith

WA
WA
DE
ON

WSUC
WSUC
USNM
CNCI

AB,

CA,

NB

CNCI, USNM

AUoxystn sp.

WA,
KS

WV

USNM, WSUC
USNM

Lysiphkbus (Phkbus) kstiiceipes

WA

WSUC

(Cresson)

Praon sp.

WA
WA

WSUC

wsuc

DiaercticUa rapae (M'Intosh)

WA
WA

WSUC

wsuc

Aphidius ervi Haliday; DuicrcticJIa ra-

CA,

MD,

WA,

EMEC, USNM,

pae (M'Intosh)

Wl

WSUC

NB

CNCI

Aphidius salicis Haliday

NB,
CA
CA

WA

CNCI, WSUC

USNM

UCRC

T. (Trioxys) pnllidus (Haliday)

WA

WSUC

Volume 7, Number 2, 1998
Table 2. Continued.

233

Aphid or (tlher Hosts

Associated Pnnl<ir\' Hti'^t'i

Museum
Acronvms

Diurapliis iioxiii (Mord\'ilko)

Dyiaplus {Pomai'luf) p'lanta-

ginea (Passerini)
Elaiohuim abtctiuiim (Walk-
er)
Ericaphis gentneri (Mason)
Eriosonm amerkanum (Riley)
£. laiiHgtiiosmn (Hartig)
Eucnlliptcnis tilinc (L.)
Eitccrnplui piuiictipeiuiis (Zet-

lerstedt)
Hayhurstia atripUcis (L.)
Hyaloptcrus priini (Geoffroy)
Hyperomyzus lactucae (L.)
Hypewmyzui {Ncimasonovm)

lugriconiif (Knowlton)
lUimiiii sp.

/. Urioiientiri (Monell)
/. spirneae (MacGillivary)
Lipapliis crysimi (Kalten-

bach)
Liofcmaphis hcrbcriilis (Kal-

tenbach)
Mncw^iphoncilln ludoviciajiac

(Oestlund)
Miicrofiplmni sp.

M. cnvlii Davis

M. mphcrihiK (Thomas)

M. rosac (L.)
Malacosoma sp.-
Mctopolopliiiiin dirlwtltiin

(Walker)
Moncllia caryae (Monell)
Myzaptm rosarum (Kalten-

bach)
Mi/2»i(S [Nectarosiphoii) per-

sicae (Sulzer)

Nasonovia (Kakimia) sp.
Nearctaphis hakeri Cowen
Ohtusicauda coweni (Palmer)
OiHitiif cratacgarius

(Walker) ?
Paraphif jiiglaiuiis (Goeze)
Periphylhis h/rcpictui^ (Kes-

sler)
P. iwgundiuh (Thomas)
Phorodcu humuli (Schrank)
PIcctrichophous sp. ?
Psylla pyricola Forster'

Apludiuf ervi Halliday; Diitcictielln

rapae (MTntosh)
Praon sp., P. unicum Smith

Prtion unicum Smith
Aphelinuf mail (Haldeman)

Praon sp.
Allcxysta sp.

Prcwn sp.

Aphidius sp.; Prnon sp.

Pnion sp.
Aphidius sp.
Priwn sp.
Trwxys sp.

Apliidius sp.; Dmeretielhi rapac
(MTntosh); Praon sp., P. unicun
Smith

Praon unicum Smith

T. (Trioxys) pallidus (Haliday)
Eunphidius setiger Mackauer

Praon unicum Smith

TX, '

WA

TAMU, WSUC

WA

WSUC

BC

NFRC

WA

WSUC

SK

CNCI

NY

USNM

CA

CNCI

CA

USNM

WA

WSUC

CA

EMEC

NB,

WA

CNCI, WSUC

WA

WSUC

WA

WSUC

CA,

DC

EMEC, USNM

ME,

WA

USNM, WSUC

CA,

OK

CNCI, EMEC

NB

CNCI

WA

WSUC

NB,

OH,

ON,

CNCI, USNM

WA

WA

WSUC

DC,

ME,

NB,

CNCI, USNM,

NJ

, WA

„ WI

UWEM, WSUC

CA

CASC, UCRC

BC

CNCI

WA

WSUC

WA

WSUC

DC

USNM

CA,

MD,

MN,

CNCI, EMEC,

NB, OH, WA

UCRC, USNM,

WSUC

WA

WSUC

WA

WSUC

WA

WSUC

WA

WSUC

WA

WSUC

WA

WSUC

NB

CNCI

WA

WSUC

WA

WSUC

BC, (

ON

CNCI

234

Journal of

Hymenoptera Research

Table 2. Continued.

Museuin

Aphid or other Hosts

Associated Primary' Hosts

LocaHties

Acronvms

Rhadobium porosus (Sander-

NB

CNCl

son)

Rhopalomyzus (Judenkoa) hn-

Praon sp.

WA

WSUC

icerae (Siebold)

Rhopahsiplioiiiiiiis (M\/zosi-

NB

CNCI

phcn) folani (Thomas)

Rtwpahisiplium sp.

ON

CNCI

R. ccrasifoliiic (Fitch)

Lysiphlebus (Phlebus) testaceipes
(Cresson)

WA

WSUC

R. insertion (Walker)

Lysiphlebus (Phlebus) testaceipes
(Cresson); Pracin unieum Smith

WA

WSUC

R. maidis Fitch

Alloxxista sp.

MN, WA

USNM, WSUC

R. padi (L.)

Dineretiella rapae (M'lntosh); Lysi-
phlebus (Phlebus) testaceipes (Cres-
son)

WA

WSUC

Schizaphis graminum (Ron-

IL, SK

CNCI, INHS

dani)

Sipilia {Rungfia) nwydis Pas-

ON

CNCI

serini

Siphocoryne sp.

DC

USNM

Sitobion avenue (Fab.)

Aphuiius sp.; Diaeretiella rapae
(MTntosh); Praon sp.

OR, WA

OSUC, WSUC

Therioaphis trifoUi (Monell)

Aphelinus semiflavus Howard'

CA, WA

EMEC, WSUC

Thriptsaphis sp.

Praon sp.

WA

WSUC

Uwleiicon sp.

Alloxysla sp.; Praon sp.

WA

WSUC

U. nmbrosine (Thomas)

Aphidius pclygonaphis Fitch ?

MD, ON

CNCI, USNM

U. (Lambersiu) madia Swain

NB, N], ON

CNCI, USNM

U. sonchi (L.) ?

WA

WSUC

Wahlgreniella nervata (Gil-

Praon sp.

WA

WSUC

lette)

' Chalcidoidea (Aphelinidae).

- From "egg mass", but likely some aphid with the egg mass.

' Homoptera (Psyllidae).

or subeffaced net-like sculpture over bare
area. Scutellum mostly bare except along
extreme anterior and lateral margins (Fig.
47); frenum smooth and shiny except fine-
ly carinate laterally. Metapleuron bare
(Fig. 65). Forewing with at least two rows
of setae in basal cell (Fig. 69); disc with
very narrow speculum, its dorsal surface
with at least three setae close to submar-
ginal vein (within distance equal to or less
than length of setae) (Fig. 69). Petiole (Fig.
47) length at least slightly greater than
width (up to about 1.25 times), and
strongly carinate with reticulate or subef-
faced sculpture between carinae.

Male. — Color pattern similar to female;

antenna rarely uniformly dark brown,
much more commonly with flagellum and
usually pedicel yellowish or at least dis-
tinctly lighter brown than dark scape.
Scape (Figs. 39, 40) robust-subcylindrical,
length about 4-5 times width, with dorsal
and ventral margins subparallel, and usu-
ally with obscure linear or very narrow
ventrally or externally angled, flat, micro-
setose sensory strip (Fig. 40a); inner sur-
face low convex to flat, smooth, shiny and
sparsely setose (Fig. 40b); outer surface
(Fig. 40a) more distinctly convex. Com-
bined length of pedicel and flagellum less
than 2.5 times scape length and shorter
than head width; funicle (Fig. 39) with all

Volume 7, Number 2, 1998

235

segments at least slightly transverse. Setal
pattern similar to female except metapleu-
ron rarely with one short setae directed
ventrally toward metacoxa. Structure sim-
ilar to female, with petiole always at least
slightly longer than wide (up to about 1.35
Hmes) (Fig. 59).

Distribution. — A naturally occurring Hol-
arctic species (see also under 'Remarks'). In
North America distributed throughout the
Nearctic region (Fig. 81) and extending
south into Mexico. The absence of observed
specimens from the states and provinces
listed below undoubtedly reflects artifacts
of collection (except possibly for Newfound-
land and Labrador). CANADA: all territo-
ries and provinces except Yukon, and New-
foundland and Labrador. USA: all states ex-
cept Alabama, Louisiana, New Jersey,
North Dakota, Rhode Island, and Vermont.
EXTRALIMITAL. Neotropical: MEXICO
(CNCl). Palaearctic: AZORES (USNM), CA-
NARY ISLANDS (CNCl), CZECH REPUB-
LIC (CNCl), DENMARK (CNCl), FIN-
LAND (CNCl, DAZH, WPC), FRANCE
(BMNH, CNCl EMEC, UCRC, USNM),
GERMANY (CNCl), GREAT BRITAIN
(BMNH, EMEC), HUNGARY (CNCl),
ICELAND (GNME, LUND, ZMCU),
IRAN (ANIC, BMNH, CNCl, UCRC), IS-
RAEL (EMEC), ITALY (BMNH, CNCL
UCRC), JAPAN (CNCL EMEC, UCRC,
USNM), SOUTH KOREA (UCRC), MA-
DEIRA ISLANDS (BMNH), ? MOROC-
CO (UCRC), NETHERLANDS (TAMU),
PEOPLE'S REPUBLIC OF CHINA (CNCl,
TAMU), POLAND (TAMU), PORTUGAL
(BMNH), SPAIN (CNCL UCRC), SWE-
DEN (BMNH, CNCL MZLU), ? TURKEY
(UCRC), UKRAINE (USNM). Oriental: IN-
DIA (CNCL UCRC), NEPAL (CNCl), PA-
KISTAN (UCRC).

Biolog}/. — Specimens with host data
from the Nearctic region indicate that A.
suspensiis is usually a hyperparasite of
aphids through aphidiine and aphelinid
primary parasites, and rarely also a para-
site of Psi/lla (Homoptera: Psyllidae) (Ta-

ble 2). One record from a lepidopteran egg
mass is undoubtedly erroneous.

Synonymy and type material examined. —
Asaphes suspensus was generally consid-
ered to be a synonym of A. vulgaris until
Graham (1969) reestablished the name as
valid. Our synonymy of A. rufipes with A.
suspensus is based on Graham's (1969)
concept and examination of the holotype
of A. rufipes. It is glued to a point and is
entire, though the right wings are glued
over the metasoma and the body is oth-
erwise covered by a film of glue except for
one antenna, about the dorsal half of the
head and the dorsal surface of the meso-
soma. The specimen is labelled as from
"Boston Mass." rather than "Forest Hills,"
which was given in the original descrip-
tion, and has the additional labels "3671,"
"27315," "TYPE," "Asaphes rufipes
Brues," "Holotype Asaphes rufipes Brues,
V. Vikberg 1986," "Asaphes suspensus
(Nees) det. V. Vikberg 1986."

The holotype female of A. fletcheri is
point-mounted and entire. It has a hand
written label with "Megorismus fletcheri
Cwfd. $ type" and a red "Holotype" la-
bel. There are also 5 9 and 1 6 labelled
as paratypes in the USNM as well as 5 9
labelled as paratypes in the CNCL All
specimens are labelled identically except
for type labels. Because Crawford explic-
itly referred to 'paratypes' in the original
description we consider the female la-
belled originally as 'Type' to be the holo-
type by original designation. The holotype
is point-mounted and entire.

The type series of A. americanus consists
of 4 9 and 1 6 syntypes on five slides.
The Hampton female from experiment 102
is labelled as 'Type', the male from exper-
iment 101 as 'Allotype', and the other
three females as 'Paratypes'. However, in
the original description Girault referred to
all specimens as 'Types'. We hereby des-
ignate the Hampton female from experi-
ment 101 as lectotype and the other four
specimens as paralectotypes. The female
originally labelled as type is not selected

236

Journal of Hymenoptera Research

as lectotype because the cover slip over
the specimen is crushed.

Boucek et al. (1978) transferred Pachi/-
crepoides indicus Bhatnager to Asaphes
based on the original description and ac-
companying illustrations, and tentatively
synonymized the name under A. vulgaris
with the statement "judging from some
points as e.g. head from above rectangu-
lar." We agree with the generic placement,
but the original description states "Legs
except the three coxae which are black,
rest of legs are light yellowish-brown."
This statement leads us to believe the
name is a junior synonym of A. suspensus.

Remarks. — Females of A. suspensus are
recognized by the combination of entirely
yellowish legs and relatively narrow spec-
ulum with at least three setae very close
to the submarginal vein (Fig. 69). Because
of color variation, without associated
males it is sometimes very difficult to dis-
tinguish females of A. suspensus and A. cal-
ifornicus in western North America. Males
of A. californicus and A. vulgaris can have
entirely yellowish legs, but structure of
the scape distinguishes males of A. califor-
nicus and a broad speculum distinguishes
males of A. vulgaris. Males of these last
two species usually also have uniformly
brown antennae (see under A. californicus).

Asaphes suspensus is undoubtedly wide-
spread throughout the Palaearctic region
from western Europe to Japan, much more
so than is indicated by the relatively few
countries listed above or in Graham
(1969). Bakkendorf (1955), and probably
based on him Graham (1969), reported A.
vulgaris from Iceland. The junior author
examined 3 females identified as A. vul-
garis by Bakkendorf in 1955 (GNME),
which were specimens of A. suspensus.
Asaphes suspensus was also reported from
southern Iceland and the Westman Islands
by Lindroth et al. (1973). Because all spec-
imens of Asaphes that we have seen from
Iceland are A. suspensus it seems likely
that this is the only species occurring in
Iceland.

Farooqi and Subba Rao (1986) list A.
suspensus from India and Pakistan, which
we confirm. De Santis (1967, 1979) and De
Sands and Fidalgo (1994) also recorded A.
suspensus (as A. lucens and A. rufipes) with-
in the Neotropical region as far south as
Chile and Argentina. We saw specimens
from La Plata and Jujuy, Argentina
(MLPA) that were identified as A. fletcheri
and A. rufipes by De Santis, but which fit
within our concept of the californicus-com-
plex. As discussed in the respective sec-
tion for A. californicus, we suspect that
most if not all records of A. suspensus
south of Mexico result from misidentifi-
cation of the californicus-complex.

Asaphes vulgaris Walker

(Figs. 11, 12, 15, 29, 30, 41, 41, 48, 60, 70)

Asaphes vulgaris Walker 1834: 152. Both sexes
described. Lectotype female designated by
Graham 1969: 80-81; BMNH.

Eurytoma aenea Nees 1834: 42. Type data: Ger-
many: Sickershausen, 11 June 1813. Female
described. Holotype by monotypy, lost. Syn-
onymy by Walker 1846: 23.

CIm/solampus aeneus Ratzeburg 1848: 185. Fe-
male described. Holotype by monotypy, ?
lost (see Graham 1969: 81). Synonymy by
Reinhard 1857: 76.

Chrysolampus aphidophila Rondani 1848: 21-22.
Female described. Lectotype designated by
Boucek 1974: 244; Museo Zoologico "La Spe-
cola," Florence, Itatly. Synonymy by Boucek
1974: 244, 275.

Female. — Head and mesosoma dark
with variedly distinct olive green metallic
luster under some angles of light; legs
mostly dark, at least middle and hind legs
with trochanters infuscate to black [light-
colored in some regions of western Eu-
rope] and femora black except apically,
trochantelli often much lighter to yellow-
ish, and tibia and tarsus usually yellowish.
Head transverse-subtriangular in frontal
view (Fig. 11), width at least 1.25 times
height, and in lateral view lower face
evenly curved into upper face (Fig. 12); in-
terorbital region in dorsal view relatively

Volume 7, Number 2, 1998

237

deeply concave (Fig. 15); gena length
about 0.57-0.67 eye width and 0.5-0.6 eye
length; dorsal margin of torulus approxi-
mately in line with lower orbit (Figs. 11,
12). Antenna (Figs. 29, 30) with pedicel
length 1.65-2.3 times greatest width; fu-
nicle (Fig. 29) with fu, ring-like, fuj-fuj
quadrate to transverse (Fig. 30), and fur-
fus increasingly transverse. Mesoscutum
(Fig. 48) with lateral lobes broadly bare
medially, and with fine but distinct, en-
graved net-like sculpture over bare area.
Scutellum mostly bare except along ex-
treme anterior and lateral margins (Fig.
48); frenum smooth and shiny except fine-
ly carinate laterally. Metapleuron at most
with one or two short setae directed ven-
trally toward metacoxa. Forewing with
basal cell usually almost bare or with only
single row of setae on dorsal surface (Fig.
70) except apically; disc with broad spec-
ulum, dorsal surface without setae near
submarginal vein and with distance be-
tween basal setal line and first setal line
on disc about equal to distance between
first and fourth setal lines on disc (Fig. 70)
(note: ventral surface of wing can have a
few setae on bare band closer to submar-
ginal vein). Petiole at least as long as and
usually slightly longer than wide (up to
1.2 times), entirely or mostly reticulate
with at most few irregular carinae (Fig. 48)
to strongly carinate with subeffaced sculp-
ture between carinae.

Male. — Color pattern similar to female
except legs usually more extensively yel-
lowish, with trochanters and trochantelli
usually yellowish and often legs mostly or
entirely yellow; antenna uniformly brown
to black except possibly apex of pedicel
and fu, lighter in color. Scape (Figs. 41, 42)
elongate-subcylindrical or spindle-shaped,
length about 5-6 times width, with dorsal
and ventral margins subparallel; outer
surface with shorter setae along line ven-
trally but without well defined microse-
tose area or smooth band (Fig. 42a); inner
surface with sparse setae (Fig. 42b). Com-
bined length of pedicel and flagellum less

than 2.5 times scape length and subequal
in length to head width; funicle (Fig. 35)
usually with all or most segments slightly
to distinctly transverse, but at least fu, dis-
tinctly transverse, fu, transverse to quad-
rate, fu, and fuj transverse to slightly lon-
ger than wide, fu=, and fu^ transverse to
quadrate, and iu-_^ distinctly transverse.
Setal pattern and structure otherwise sim-
ilar to female except petiole longer, length
1.25-1.60 times width (Fig. 60).

Distribution. — True world distribution is
uncertain because of remaining taxonomic
problems in differentiating the species (see
under 'Remarks'). In North America A.
vulgaris is restricted to eastern Canada and
northeastern USA (Fig. 82). Based on this
distribution and no collection records seen
prior to 1953 it is probable that the species
it is not naturally Holarctic but was intro-
duced accidentally from Europe. Litera-
ture records from California and other
western North American localities are
based on misidentifications, probably
mostly of A. californicus. At least some lit-
erature records from eastern North Amer-
ican localities are probably also based on
misidentifications. CANADA. Ontario:
Rondeau Provincial Park, 9-26.VI.80, H.
Goulet (1 9). Quebec: Quebec city,
27. VII. 86, J. Brodeur, ex. Aphidius nigripes
from Macrosiphum euphorhiae (1 9, 2 d).
Nova Scotia: Aylesford, 24.VII.63, H.
Specht, host Acyrthosiphum pisum (1 9 ).
Centerville, 15. VIII, 9.IX.63, R. Foley, (4 9,
1 i); Coldbrook, 30.IX., 7, 10, 16, 21.X.63
(6 9, 3 6), Coldwell near Canard, 8,
23.VIII.63 (1 9, 1 i), Garland, 6.VIII.63 (1
i), Picketts Wharf near Canard, 2.VII.63
(2 9,1 (J)— all H. B. Specht, host Acyrtho-
sipluim pisum (Harris) on alfalfa or clover.
Kentville, greenhouse, 28.11.68, H.B.
Specht, host pea aphid (1 S). Lockport,
20.VII.58, J.R. Vockeroth (1 9). Prince Ed-
ward Island: N Tryon, 26.VI.91, M.E.M.
Smith, in potato field (1 9, 1 S). USA.
Maine: Aroostook Co., 1953, ex. Macrosi-
phum solaiufolii (4 9, 3 6). Massachusetts:
Amherst, Nadel, clover (1 9, USNM). EX-

238

Journal of Hymenoptera Research

TRALIMITAL. Palaearctic: AUSTRIA
(CNCI), CZECH REPUBLIC (CNCI,
UCRC), DENMARK (CNCI), FINLAND
(DAZH, WPC), FRANCE (CNCI, EMEC,
UCRC, USNM), GERMANY (BMNH,
CNCI, USNM), GREAT BRITAIN (ANIC,
BMNH, CNCI, EMEC, USNM), GREECE
(CNCI, TAMU, UCRC), IRELAND
(CNCI), ISRAEL (UCRC, USNM), ITALY
(CNCI, UCRC, USNM), LEBANON
(UCRC), MOROCCO (UCRC), SPAIN
(CNCI, UCDC, UCRC), SWEDEN (BMNH,
CNCI, MZLU), TURKEY (UCRC, USNM).
Afrotwpical: ? ERITREA (UCRC), ? ETHI-
OPIA (CNCI), ? SOUTH AFRICA (CNCI,
USNM), ? ZIMBABWE (CNCI). Austral-
asian: AUSTRALIA (Tasmania) (ANIC,
BMNH), NEW ZEALAND (CNCI, EMEC).

Biology. — Host records based on exam-
ined specimens indicate that in North
America A. vulgaris is a hyperparasite of
aphids, including Acyrthosiplwn pisum
(Harris) and Macrosiphum euphorbiae (Tho-
mas) through aphidiines, including Aphi-
dius nigripes (Ashmead). Most of the un-
substantiated aphid host records from
western North American localities given
in Peck (1963) likely refer to A. californicus,
whereas non-aphid hosts other than Syr-
phidae likely are erroneous.

Remarks. — Females of A. vulgaris are dis-
tinguished by a combination of features,
but primarily by their broad, distinct spec-
ulum (Fig. 70), dark trochanters (at least
in the Nearctic region), and relatively
deeply concave interorbital region (Fig.
15). Females of A. californicus with a broad
speculum could be confused with A. vul-
garis females, but ranges of the two spe-
cies apparently do not overlap in the New
World (Fig. 82). Further, Nearctic A. cali-
fornicus females have more or less uni-
formly light-colored trochanters and tro-
chantelli, whereas the trochanters and tro-
chantelli are dark in A. vulgaris or at least
the trochanters are distinctly darker than
the trochantelli, more similar in color to
the femora. Some females of A. vulgaris
from western Europe have both the tro-

chanters and trochantelli yellowish in dis-
tinct contrast to dark femora, but their
broad speculum readily differentiates
these from other known species in western
Europe. All specimens tentatively identi-
fied as A. vulgaris from the Afrotropical
region also have yellow trochanters and
trochantelli plus a broad speculum. Only
one male was seen from Zimbabwe but a
series of both sexes were seen from Ethi-
opia, Eritrea and South Africa. The Afri-
can males have scapes with a variedly dis-
tinct, flat, densely setose, sometimes lat-
erally margined ventral surface. The
scapes of most specimens are also distinct-
ly shorter and more robust (more similar
to A. suspensus, cf Fig. 39) than the rela-
tively elongate-slender, shinier, and much
more sparsely setose scape of A. vulgaris
males from North America or Europe (Fig.
41). Further, both sexes of the African
specimens appear to have the interorbital
region more shallowly concave (cf. Figs.
16, 17 with Fig. 15), though this feature is
variable and difficult to quantify. Only fe-
males were seen from Lebanon and Mo-
rocco. The Afrotropical specimens are ten-
tatively identified as A. vulgaris for the
purposes of this study, but a comprehen-
sive study of Asaphes is required from
throughout the Palaearctic region and Af-
rica to determine character variation and
species limits. It is very possible that spec-
imens with dark femora and yellowish
trochanters and trochantelli from at least
the Aftrotropical region of Africa belong
to an undescribed species different from
A. vulgaris.

Males of A. vulgaris, particularly from
regions outside of the Nearctic, can have
the legs entirely yellow but are differen-
tiated from males of A. suspensus by their
broad speculum and longer scape (cf Figs.
39, 40), though the range of variation is
greater than stated by Graham (1969) and
the shortest scape of A. vulgaris males is
similar in relative length to the longest A.
suspensus scape. Separation from males of

Volume 7, Number 2, 1998

239

other species is discussed under the re-
spective section for these species.

De Santis (1967, 1979, 1980) hsted A.
vulgaris from San Vincente, Juan Fernan-
dez, Argentina, Brazil, and Chile. We saw
specimens from La Plata (9, cJ) and jujuy
(?), Argentina (MLPA) that were identi-
fied by De Santis as A. vulgaris, and these
fit within our concept of the califoniicus-
complex from South America. We have
not seen any specimen from the Neotrop-
ical region that we identify as A. vulgaris
(see 'Remarks' for A. californicus) and it is
likely that most or all records of A. znilgaris
from South America are based on misi-
dentifications (see also 'Remarks' for A.
suspensus). Records of A. vulgaris from
Greenland may also be based on misiden-
tifications of A. hirsutus (see 'Remarks' for
A. hirsutus).

Nomina inquirenda

Parectroma hiibrichi Brethes 1913: 91-92. Type
data: Argentina: Rosario (J. Hiibrich) and
Buenos Aires (A. Zotta). Female described.
Syntypes [Rosario female examined]; MLPA.
Synonymized with A. fletcheri by De Santis
1960: 114.

Pachycrefoideus bonariensis Brethes 1916: 427.
Type data: Argentina: Buenos Aires, 5. XI.
1915. Female described. Holotype by mono-
typy. Synonymized with A. liicens by De San-
tis 1967: 189.

Remarks.— De Santis (1960, 1967) syn-
onymized, respectively, A. huebrichi (Bre-
thes, 1913) under A. fletcheri (Crawford,
1909) and A. bonariensis (Brethes, 1916) un-
der A. lucens (Provancher, 1887). Burks
(1964) synonymized A. fletcheri under A.
lucens, which in this paper we synonymize
under A. suspensus (Nees, 1834). We ex-
amined the 'Rosario' female syntype of A.
huebrichi. It is mounted laterally on a slide,
is crushed, and otherwise is poorly pre-
served. The petiole is quadrate, although
this appearance may partly be because it
is flattened somewhat by the cover slip.
The specimen also has uniformly yellow-
ish legs, which led De Santis to synony-

mize the name under A. fletcheri. Howev-
er, one forewing remains attached and al-
though it lies over the hind wing a broad
speculum is visible. Because of the ab-
sence of a row of setae immediately be-
hind the submarginal vein on the disc we
do not consider that A. huebrichi is syn-
onymous with A. suspensus. The specimen
has brownish coxae, a feature that we
have observed only in some females from
South America with uniformly yellowish
legs and a broad speculum, which we cur-
rently include in the californicus-complex
from South America (see 'Remarks' for A.
californicus). At this time we do not for-
mally synonymize the names A. huebrichi,
A. bonariensis, and A. californicus. Formal
synonymy of either A. huebrichi (1913) or
A. bonariensis (1916) with A. californicus
(1917) would result in A. californicus be-
coming the junior synonym. Our study of
Neotropical material has been insufficient
to confidently resolve the limits of varia-
tion and number of species of Asaphes oc-
curring in the Neotropical region. For rea-
sons of stability we do not consider it ap-
propriate to make formal nomenclatural
changes until concepts of species are bet-
ter resolved and topotypic material of A.
huebrichi and A. bonariensis can be studied.
For this reason we treat these names as
nomina inquirenda.

ACKNOWLEDGMENTS

We thank Drs. J. Huber and H. Goulet, Eastern Ce-
real and Oilseed Research Centre (ECORC) for criti-
cally reviewing this manuscript; Mr. K. Bolte (Cana-
dian Forest Service) prepared the specimens for SEM
and produced the micrographs, Ms. Inna Nei pre-
pared the distribution maps; and Ms. J. Read
(ECORC) produced the plates of illustrations. We
thank Dr. K. Kamijo, Hokkaido Forest Experiment
Station, |apan, for the gift of representatives of both
lapanese species. The following collections and cu-
rators also provided specimens for this study:

AEIC American Entomological Institute, Gaines-
ville, FL (D. Wahl).

ANIC Australian National Insect Collection, CSI-
RO, Canberra Cit>', Australia (I. Naumann).

AFRC Atlantic Forestry Research Centre, Canadi-

240

Journal of Hymenoptera Research

■■i^M

^B||H

^^F ' ' /jM^^^SSf^SS^St^^Ss^^^^^^Ss/^t

^^^

^S^^^^^^^^^^Kk ^ ^1

H^^9

^SH^^^Pl^^^H

^^P^^JhR

^^^^^^^^^bN^ ^^ ~ ^^%'I^HhK^

^^^■^^^H

^^^^^^^^^^^^^^^^^^H^^l

I^^^O

Figs. 1-6. Head: 1 and 2, /4sflp/ics brevipetiolatus (9); 3 and 4, /I. bivvipctiolntu^ (6); 5 and 6, /^. /!irs»f»s (9).
(abbreviation: cly = clypeus; scale bars = microns)

Volume 7, Number 1, 1998

241

Figs. 7-12. Head (9): 7 and 8, Aia\>hes catifornicKs; 9 and 10, A. susjvmsks; 1 1 and 12, A. vulgaris, (abbreviation:
gc = genal carina; scale bars = microns)

242

Journal of Hymenoptera Research

■

1

■

i

1

■

m

^^^

1

1

I

■

I

t^7J

Figs. 13-18. 13-17. Head, dorsal; 13, Asnpluv hrcvifetiotatiif. (9); 14, A. hirvii't'tu^lntiis (d); 15, A. vulgaris (9);
16, A. suspensus (9); 17, A. hirsutus (9). 18. Clypeus and mandibles, A. Iiir^iifii^ (9). (abbreviation: occ =
occipital carina; scale bars = microns)

Volume 7, Number 2, 1998

243

Figs. 19-24. 19 and 20. Asa^'hef brevipetiolatus (9): 19, antenna; 20, basal 4 flagellar segments. 21 and 22. A.
californicus (9): 21, antenna; 22, basal 4 flagellar segments. 23 and 24. A. hir^utuf. (9): 23, antenna; 24, basal 4
flagellar segments, (abbreviation: fu, = first funicular segment; scale bars = microns)

244

Journal of Hymenoptera Research

Figs. 25-30. 25 and 26. Asnphcf pftiohitm (V); 25, antenn.i; 26, b.isal 4 fldgoUar segments. 27 and 28. A.
suspoi.sKs (9): 27, antenna; 28, basal 4 flagellar .segments. 29 and 30. A. vulgaris {9): 29, antenna; 30, basal 4
flagellar segments, (scale bars = microns)

Volume 7, Number 2, 1998

245

Figs. 31-36. 31 and 32. Asaphes hreinpetiolatus (cJ): 31, antenna; 32, scape — a, outer view, b, inner view. 33
and 34. A. californicus (S): 33, antenna; 34, scape — a, outer view, b, inner view. 35 and 36. A. hirsutus (d): 35,
antenna; 36, scape — a, outer view (arrow points to flat surface), b, inner view, (scale bars = microns)

246

Journal of Hymenoitera Research

Figs. 37-42. 37 and 38. Asaplws petiolatus (cj): 37, antenna; 38, scape — a, outer view, b, inner view. 34 and
40. A. suspensus (6): 39, antenna; 40, scape — a, outer view, b, inner view. 41 and 42. A. vulgaris (cj): 41, antenna;
42, scape — a, outer view, b, inner view, (scale bars = microns)

Volume 7, Number 2, 1998

247

Figs. 43-48. Mesosoma, dorsal (9). 43, Asaphes brevipetiolatus; 44, A. califormcuf, 45, A. Imsuhis; 46, A. pct-
wlatiis; 47, A. smpciisus; 48, A. vulgaris, (abbreviation: sss = scutoscutellar suture; scale bars = microns)

248

Journal of Hymenoptera Research

Figs. 49-54. 49 and 50. Aaiiphes hmnpetiolcitiis (9): 49, frenum to petiole; 50, sculpture of scutellum and
frenum. 51 and 52. A. petiolatua (9): 51, frenum to petiole; 52, sculpture of scutellum and frenum. 5.1 and ,54.
A. /iirsKfus (9): 53, frenum to petiole; 54, sculpture of scutellum and frenum. (abbreviations: fre = frenum,
set = scutellum; scale bars = microns)

Volume 7, Number 2, 1998

249

Figs. 55-60. Frenum to petiole (d): 55, A. breinyctiolatus; 56, A. catifonitcus; 57, A. hirsiitus; 58, A. pefiolatus;
59, A. suspensus; 60, A. vulgaris, (scale bars = microns)

250

Journal of Hymenoptera Research

Figs. 61-66. 61 and 62. Asaphci liir^iitiis: 61, lateral mcsosoma (9); 62, niL'tapleuron (c!). 63. A. cnUtcniicii-i:
lateral mesosoma (d). 64-66. Metathorax to petiole, lateral (9) (arrow points to lateral sulcus ot petiole): 64,
A. brevipeliolatu'^; 65, A. petwlnttis; 66, A. s»s;v«s»s. (scale bars = microns)

Volume 7, Number 2, 1998

251

Figs. 67-72. Forewing (9): 67 and 68, Aaaphcs cahfonuctis; 69, A. ^uspensus; 70, A. vulgaris; 71, A. htnittus; 72,
A. petiolatiis. (abbreviations: be = basal cell, bsl = basal setal line, cc = costal cell, smv = submarginal veini
spc = speculum; scale bars = microns)

252

Journal of Hymenoptera Research

Figs. 73-78. 73. Forewing: Asaphcs brcvifetiolahis ( 9 ). 74. Gaster: A. himiitii^ ( 9 ). 75. Petiole and base of gaster,
ventral view: A. pctiolatus (9). 76. Petiole, ventral view: A. cnlifoniiciis (9). 77 and 78. A. stisff.ifui, base of
gaster (9): 77, ventral view; 78, ventrolateral view, (scale bars = microns)

Volume 7, Number 2, 1998

253

Figs. 79-80. Nearctic distribution: 79, Asaphes brevi-
petiolatus (•) and Asaphes petiolatuf (A). 80. Asaphes
hirsutus.

Figs. 81-82. Nearctic distribution: 81, Asaphes sus-
pensus. 82, Asaphes califcrnicus (•) and A. vulgaris (A).

an Forest Service, Insect Collection, St.

John's, NF (G. Smith). DCPC

BMNH The Natural History Museum, London, En-
gland (J. Noyes, Z. Boucek). EDUM

CASC California Academy of Sciences, Depart-
ment of Entomology, San Francisco, CA (D. EMEC
Ubick, W. Pulawski).

CISC California Insect Survey, Division of Ento- GNME
mology and Parasitology, University of
California, Berkeley, CA (R. Zuparko). HAPC

CNCI Canadian National Collection of Insects

and Arachnids, Ottawa, ON. INHS

CUIC Cornell University Insect Collection, De-
partment of Entomology, Cornell Univer- MLPA
sity, Ithaca, NY (E. R. Hoebeke).

DAZH Department of Applied Zoology, Universi-

ty of Helsinki, Helsinki, Finland (M. Ko-
ponen).

Dominique Collet private collection. Ster-
ling, AK (D. Collet).

Department of Entomology, University of
Manitoba, Winnipeg, MB (T. Gallowav).
Essig Museum of Entomology, University
of California, Berkeley, CA (R. Zuparko).
Department of Entomology, Naturhisoriska
Museet, Goeborg, Sweden (G. Andersson).
Harry Anderson Private Collection, Hun-
tington Beach, CA (H. Anderson).
Illinois Natural History Survey Insect Col-
lection, Champaign, IL (K.R. Zeiders).
Museo de la Plata, Universidad Nacional
de La Plata, Division Entomologia, La Pla-
ta, Argentma (L. De Santis).

254

Journal of Hymenoptera Research

MZLU Museum of Zoology, Lund University,
Lund, Sweden (R. Danielsson, U. Garden-
fors).
NFRC Northern Forest Research Centre, Canadian
Forest Service, Edmonton, AB (D. Wil-
liams).
PFRC Pacific Forestry Research Centre, Canadian

Forest Service, Victoria, BC (L. Humble).
OSUC Collection of Insects and Spiders, Ohio
State University, Columbus, OH (A. Shar-
kov).
OSUO Department of Entomology Collection, Or-
egon State University, Corvallis, OR (D.C.
Darling).
SMDV Spencer Museum, Department of Zoology,
University of British Columbia, Vancouver,
BC (K. Needham).
TAMU Department of Entomology, Texas A&M
University, College Station, TX (J. Woolley,
E. Riley).
UCDC The Bohart Museum, University of Califor-
nia, Davis, CA (S. Heydon).
UCRC UCR Entomological Training and Research
Collection, University of California, River-
side, CA (S. Triapitsyn).
USNM United States National Entomological Col-
lection, U.S. National Museum of Natural
History, Washington, DC (E.E. Grissell).
UZMH Zoological Museum, University of Helsin-
ki, Helsinki, Finland (A. Albrecht, M. Ko-
ponen).
VVPC Veli Vikberg personal collection, Turenki,

Finland (V. Vikberg).
WSUC James Entomological Collection, Depart-
ment of Entomology Collection, Washing-
ton State University, Pullman, WA (K.S.
Pike), and Tree Fruit Research Center,
Washington State University, Wenatchee,
WA (D. Carroll).
ZMAS Zoological Museum, Academy of Sciences,

Saint Petersburg, Russia (V.A. Trjapitzin).
ZMUC Zoological Museum, University of Copen-
hagen, Copenhagen, Denmark (B. Petersen).

LITERATURE CITED

Ashmead, W. H. 1904. Classification of the chalcid
flies of the superfamily Chalcidoidea, with de-
scriptions of new species in the Carnegie Muse-
um, collected in South America by Herbert H.
Smith. Memoirs of tlw Carncf^ie Miisi'iiin 1: i-ix +
225-551.

Bakkendorf, O. 1955. Notes on Icelandic and Green-
landic Chalcidoideous Hymenoptera. Eutomolo-
giske Mcddek'lser 27: 135-162.

Batulla, B. A. and A. G. Robinson. 1985(1984). Hy-
menopterous parasitoids of aphids (Homoptera:
Aphididae) in Manitoba. Proccciimgs of the Ento-
mological Socicti/ of Manitoba 40: 30-38.

Bhatnager, S. P. 1951. Descriptions of new and re-
cords of known Chalcidoidea (Parasitic Hyme-
noptera) from India. Indian journal of Agricultural
Science 21: 155-178.
Bocchino, F. J. and D. J. Sullivan, S.J. 1981. Effects of
venoms from two aphid hyperparasitoids, Asa-
phes tucens and Dendrocerus carpenteri (Hymenop-
tera: Pteromalidae and Megaspilidae), on larvae
of Aphidius smithi (Hymenoptera: Aphidiidae).
Canadian Entomologist 113: 887-889.
Bolte, K. B. 1996. Techniques for obtaining scanning
electron micrographs of minute arthropods. Pro-
ceedings of the Entomological Societi/ of Ontario 127:
67-87.
Boucek, Z. 1974. On the Chalcidoidea (Hymenoptera)

described by C. Rondani. Redia 55: 241-285.
Boucek, Z. 1976. African Pteromalidae (Hymenop-
tera): new taxa, synonymies and combinations.
journal of the Entomological Societi/ of Southern Af-
rica 39: 9-31.
Boucek, Z. 1988. Australasian Chalcidoidea (Hymenop-
tera). A biosystematic revision of genera of fourteen
families, loith a reclassification of species. CAB In-
ternational, Wallingford. 832 pp.
Boucek, Z. and S. L. Heydon. 1997. Chapter 17. Pter-
omalidae. Pages 541-692 in Gibson, G. A. P., J.
T. Huber, and J. B. Woolley (eds). Annotated Keys
to the Genera of Ncarctic Chalcidoidea (Hymenop-
tera). National Research Council Canada, Re-
search Press, Ottawa. 794 pp.
Boucek, Z. and J.-Y. Rasplus. 1991. Illustrated key to
West-Palearctic genera of Pteromalidae (Hymenop-
tera: Chalcidoidea). Institut National de la Recher-
che Agronomique, Paris. 140 pp.
Boucek, Z., B. R. Subba Rao, and S. I. Farooqi. 1978.
A preliminary review of Pteromalidae (Hyme-
noptera) of India and adjacent countries. Oriental
Insects 12: 433-468.
Brethes, J. 1913. Himenopteros de la America Merid-
ional. Anales del Museo Nacional de Historia Nat-
ural de Buenos Aires 24: 35-166.
Brethes, J. 1916. Hymenopteres parasites de I'Ame-
rique Meridionale. Anales del Museo Nacional de
Historia Natural de Buenos Aires 27: 401-130.
Brown, N. R. and R. C. Clark. 1960. Studies of pred-
ators of the balsam woolly aphid, Adelges picea
(Ratz.) (Homoptera: Adelgidae). VIII. Syrphidae
(Diptera). Canadian Entomologist 92: 801^11.
Brues, C. T. 1908. Notes and descriptions of North
American parasitic Hvmenoptera. VII. Bulletin of
the Wisconsin Natural History Society 6: 160-163.
Burks, B. D. 1958. Superfamily Chalcidoidea. Pages
62-84 (11 Krombein, K. V. (ed.). Hymenoptera of
America North of Mexico. Synoptic Catalog. United
States Department of Agriculture, Agricultural
Monograph No. 2, First Supplement. Washington,
D.C, U.S. Government Printing Office. 305 pp.
Burks, B. D. 1964 (1963). The Provancher species of

Volume 7, Number 2, 1998

255

Chalcidoidea (Hymenoptera). Camuiinn Entomol-
ogist 95: 1254-1263.

Burks, B. D. 1979. Family Pteromalidae. Pages 768-
835 in Krombein, k" V., P. D. Hurd, Jr., D. R.
Smith, and B. D. Burks (eds). Catalog of Hymenop-
tera in America North of Mexico. Vol. 1. Symphyta
and Apocrita (Parasitica). Smithsonian Institution
Press, Washington, D.C. i-xvi, 1-1198.

Carew, W. P. and D. J. Sullivan. 1993. Interspecific
parasitism between two aphid hyperparasitoids,
Dendrocerus carpciiteri (Hymenoptera: Megaspili-
dae) and Asaphes luccns (Hymenoptera: Pterom-
alidae). Annals of the Entomological Society of
America 86: 794-798.

Crawford, J. C. 1909. Notes on some Chalcidoidea.
Canadian Entomologist 41: 98-99.

Curtis, J. 1842. Observations on the natural history
and economy of various insects affecting the tur-
nip-crops; including the plant-lice, maggots of
flies, caterpillers of moths, etc. journal of the Royal
Agricultural Society of England 3: 49-78.

Dahlbom, A. G. 1857. Svenska Sma-Ichneumonernas
familjer och slagten. Kongliga Si'cnska Vetcnskap-
sakademiens Handlingnr 14: 289-298.

Dahms, E. C. 1978. A checklist of the types of Aus-
tralian Hymenoptera described by Alexandre Ar-
sene Girault: 1. Introduction, acknowledgments,
biography, bibliography and localities. Memoirs
of the Queensland Museum 19: 127-190.

Delucchi, V. 1955. Notes sur les Pteromalides (Hym.,
Chalcidoidea). Memoircs dc la Societe royale beige
d'Entomologie 27: 171-175.

De Santis, L. 1960. Anotaciones sobre Calcidoideos
Argentinos. II. (Hymenoptera). Revista de la Fa-
cultad dc Agronomia (3a epoca) 36: 109-119.

De Santis, L. 1967. Catdlogo dc los Himenoptcros Argen-
tinos de la Serie Parasitica, Incluyendo Bethyloidea.
Provincia de Buenos Aires Gobernacion, Comi-
sion de Investigacion Cientifica, La Plata. 337 pp.

De Santis, L. 1979. Catdlogo de los Himenoptcros Chal-
cidoideos de America al sur de los Estados Unidos.
Comision de Investigaciones Cientificas de la
Provincia de Buenos Aires, Publicacion Especial,
La Plata. 488 pp.

De Santis, L. 1980. Catdlogo de los Himenoptcros Brasi-
lei'ws de la Serie Parasitica Incluyendo Bethyloidea.
Editora da Universidade Federal do Parana, Cur-
itiba. 395 pp.

De Santis, L. and P. Fidalgo. 1994. Catalogo de Hi-
menopteros Calcidoideos. Serie de la Academin na-
cional de Agronomia i/ Vetcrinaria 13: 1-154.

Dzhanokmen, K. A. 1987. 5. Familv Pteromalidae
(pteromalids). Pages 88-411 in Medvedev, G.S.
(editor-in-chief). Keys to the insects of the European
part of the USSR. Vol. III. Hymenoptera. Part //. Am-
erind Publishing. Pvt. Ltd., New Delhi, i-xii +
1341 pp. lEnglish translation of Medvedev, G. S.
(editor-in-chief). 1978. Handbook of the Insects of the

European Part of the U.S.S.R. Vol. III. Hi/nwnoptera,
Second part. Leningrad, Nauka (in Russian)).

Eastop, V. F. and D. Hille Ris Lambers. 1976. Sun'cy
of the zoorld's ap'hids. Dr. W. Junk b.v.. The Hague.
573 pp.

Farooqi, S. I. and B. R. Subba Rao. 1986. Family Pter-
omalidae. Pages 279-306 in Subba Rao, B. R. and
M. Hayat (eds). The Chalcidoidea (Insecta: Hy-
menoptera) of India and the adjacent countries.
Oriental Insects 20: 1^30.

Forster, A. 1856. Hymenopterologisclie Studien. II. Heft.
Chalcidiae und Proctotrupii. Aachen. 152 pp.

Gahan, A. B. and M. M. Fagan. 1923. The type species
of the genera of Chalcidoidea or chalcid-flies.
Bulletin of the United States National Museum 124:
1-173.

Gahan, A. B. and S. A. Rowher. 1917. Lectotypes of
the species of Hymenoptera (except Apoidea) de-
scribed by Abbe Provancher. Canadian Entomolo-
gist 49: 391^00.

Gardenfors, U. 1986. Taxonomic and biological revi-
sion of Palearctic Ephedrus Haliday (Hymenop-
tera: Braconidae, Aphidiinae). Entomologica Scan-
dinavica Supplement 27: 1-95.

Gibson, G. A. P. 1997. Chapter 2. Morphology and
Terminology. Pages 16-44 in Gibson, G. A. P., J.
T. Huber, and J. B. Woolley (eds). Annotated Keys
to the Genera of Nenrctic Chalcuioidea (Hymenop-
tera). National Research Council Canada, Re-
search Press, Ottawa. 794 pp.

Girault, A. A. 1914(219]. Descriptions of new chalcid-
flies. Proceedings of the Entomological Society of
Washington 16: 109-119.

Girault, A. A. 1917[330]. Descnptumes Hymcnoptero-
rum Chalcidoidicarum variorum cum ohservationi-
mus V. Private publication, Glendale. 16 pp.

Goulet, H. and W. R. M. Mason. 1993. Chapter 4. Use
of keys. Pages 60-64 in Goulet, H. and J. T. Hu-
ber (eds). Hymenoptera of the world: An identifica-
tion guide to famUies. Research Branch, Agricul-
ture Canada Publication 1894/ E, Ottawa. 668 pp.

Graham, M. W. R. de V. 1969. The Pteromalidae of
North-Western Europe (Hymenoptera: Chalci-
doidea). Bulletin of the British Museum (Natural
History). Entomology 16: 1-908.

Graham, M. W. R. de V. 1990. The identity of certain
problematic Dahlbom genera of Chalcidoidea
(Hym.), some represented by original material in
Zoologiska Institutionen, Lund. Entomologist's
Monthly Magazine 126(1516-1519): 197-200.

Griswold, G. H. 1929. On the bionomics of a primary
parasite and of two hyperparasites of the gera-
nium aphid. Annals of the Entomological Society of
America 22: 438-457.

Hagen, K. S. and R. van den Bosch. 1968. Impact of
pathogens, parasites, and predators on aphids.
Annual Review of Entomology 13: 325-384.

Heikinlieimo, O. and M. Raatikainen. 1981. Grid refer-

256

Journal of Hymenoptera Research

ences and names of localities in the recording of
biological finds in Finland. Notulae Entomohgkae 61:
133-154. [In Finnish].

Kamijo, K. and H. Takada. 1973. Studies on aphid hy-
perparasites of Japan, II. Aphid hyperparasites of
the Pteromalidae occurring in Japan (Hymenop-
tera). Insects Matsurnnrmm, n.s. 2: 39-76.

Keller, L. J. and D. J. Sullivan, S. J. 1976. Oviposition
behavior and host feeding of Asaphes lucens an
aphid hyperparasitoid. Journal of tlw Niic York En-
tomological Societi/ 84:206-211.

Kurdjumov, N. V. 1913. Notes on Pteromalidae (Hy-
menoptera, Chalcidoidea). Reinie Riisse d'Ento-
tiwlogic 13: 1-24.

Lai, K. B. 1934. Insect parasites of Psyllidae. Parasi-
tology 26: 325-334.

Le Ralec, A. 1995. Egg contents in relation to host-
feeding in some parasitic Hymenoptera. Ento-
niofhaga 40: 87-93.

Levine, L. and D. J. Sullivan, S.J. 1983. Intraspecific
tertiary parasitoidism in Asaphes lucens (Hyme-
noptera: Pteromalidae), an aphid hyperparasi-
toid. Canadian Entomologist 115; 1653-1658.

Lindroth, C. H., H. Andersson, H. Bodvarsson, and
S. H. Richter. 1973. Surtsey, Iceland. The devel-
opment of a new fauna, 1963-1970. Terrestrial In-
vertebrates. Entomologica Scandinavica, Suppl. 5:
1-280.

Lundbeck, W. 1897(1896). Hymenoptera Groenlandi-
ca. Videnskabelige Meddclelser fra Dansk naturhis-
torisk Forening i Kjebenhavn 1896: 220-251.

Mackauer, M. 1968. Aphidiidae. Hymenopterorum
Catalogus Pars 3: 1-103.

Mani, M. S. and G. G. Saraswat. 1974. Part III. Pages
85-107 in Mani, M. S., O. P. Dubey, B. K. Kaul,
and G. G. Saraswat. Descriptions of some new
and new records of some known Chalcidoidea
(Hymenoptera) from India. Memoirs of the School
of Entomology, St. John's College, no. 3. 377 pp.

McMuUen, R. D. 1966. New records of chalcidoid par-
asites and hyperparasites of Psylla p'yricola Fors-
ter in British Columbia. Canadian Entomologist 98:
236-239.

McMullen, R. D. 1971. Psylla pi/ricola Forster, pear
psylla (Hemiptera: Psyllidae). Pages 33-38 in Bi-
ological Control Programmes Against Insects and
Weeds in Canada 1959-1968. Commonwealth In-
stitute of Biological Control Technical Commu-
nication 4. 266 pp.

Nees ab Esenbeck, C. G. 1834. Hymenopterorum Ich-
neumonibus affiniuni monographiae, genera Enro-
peaea et species ilhistrantes. Vol. 2. ]. G. Cottae,
Stuttgart and Tubingen. 448 pp.

Peck, O. 1951. Family Pteromalidae. Pages 534-568 in
Muesebeck, C. F. W., K. V. Krombein, and H. K.
Townes (eds). Hymenoptera of America North of
Mexico — synoptic catalog. United States Depart
ment of Agriculture Monograph No. 2. 1420 pp.

Peck, O. 1963. A catalogue of the Nearctic Chalcidoi-
dea. Canadian Entomologist, Supplement 30: 1-
1092.

Peck, O., Z. Boucek, and G. Hoffer. 1964. Keys to the
Chalcidoidea of Czechoslovakia (Insecta: Hyme-
noptera). Memoirs of the Entomological Society of
Canada 34. 120 pp.

Philogene, B. J. R. and J. F. Chang. 1978. New records
of parasitic chalcidoids of pear psylla (Homop-
tera: Psyllidae) in Ontario, with observations on
the current world status of its parasitoids and
predators. Proceedings of the Entomological Society
of Ontario 109: 53-60.

Provancher, L. 1887. Additions et corrections a la Faune
Hymenopterologicpie de la province de Quebec. C.
Darveau, Quebec. 477 pp.

Ratzeburg, J. T. C. 1844. Die Ichneumonen der Forstin-
secten in forstlicher und entoinologischer Beziehung,
vol. 1. Berlin. 224 pp.

Ratzeburg, J. T. C. 1848. Die Ichneumonen der Forstin-
secten hi forstlicher und entomologischer Beziehung,
vol. 2. Berlin, vi 4- 238 pp.

Reinhard, H. 1857. Beitrageb zur Geschichte und Syn-
onymie der Pteromalinen. Berliner entomologische
Zeitschrift 1; 70-80.

Risbec, J. 1959. Pteromalidae de Madagascar (Hyme-
noptera: Chalcidoidea). Memoires de I'lnstitut
Scientifique de Madagascar, serie E II: 129-171.

Rondani, C. 1848. Osservazioni sopra parecchie spe-
cies di esapodi afidicidi e sui loro nemici. Nuovi
Annali delle Scienze Naurali e Rendiconto dei Lai'ori
dell Accademia delle Scienze dell'lstituto e delta So-
cieta Agraria di Bologna 9: 5-37.

Schulz, W. A. 1906. Spolia Hymenopterologica. Pader-
bom. 355 pp.

Sekhar, P. S. 1958. Studies on Asaphes fletcheri (Craw-
ford), a hyperparasite of Aphidius testaceipes
(Cresson) and Praon aguti (Smith), primary par-
asites of aphids. Annals of the Entomological Soci-
ety of America 51: 1-7.

Sharma, A. K. and B. R. Subba Rao. 1958. Description
of two new parasites of an aphid from North In-
dia (Aphidiidae: Ichneumonoidea and Pterom-
alidae: Chalcidoidea). Indian Journal of Entomolo-
gy 20: 181-188.

Sullivan, D. J. 1972. Comparative behavior and com-
petition between two aphid hyperparasites: Al-
loxysta victrix and Asaphes californicus (Hymenop-
tera: Cynipidae; Pteromalidae). Environmental
Entomology 1: 234-244.

Walker, F. 1834. Monographia Chalciditum. The En-
tomological Magazine 2: 148-179.

Walker, F. 1846. 1846. List of the specimens of liymenop-
terous insects in the collection of the British Museum.
Part I. — Chalcidites. E. Newman, London. 100 pp.

Waterston, J. 1929. On a chalcidoid bred from a flea
larva. Parasitology 21: 103-106.

Zetterstedt, J. W. 1838. Sectio secunda. Hymenoptera.
Pages [4171-476 in Zetterstedt, J. W.', 1838-1840,
Insecta lapponica descripta. Lipsiae. 1139 pp.

J. HYM. RES.
Vol. 7(2), 1998, pp. 257-267

Worker Versus Sexual, and Sex Ratio Investments in the Social Wasp
Vespula vulgaris (L.) (Hymenoptera: Vespinae) in England

Michael E. Archer

University College of Ripon and York St. John, Lord Mayor's Walk,
York Y031 TEX, England, U.K.

Abstract. — An attempt has been made to quantify the relative investment in workers versus
sexuals, and queens versus males in Vespula luilgaris (L.). In particular I have investigated
MacNair's hypothesis that the queen invests equally in queen and male eggs, and that her in-
vestment stops after she has laid her eggs. The dry weights and calorific values of the workers,
autumn queens, and small-cell, and large-cell, reared males were determined. The lipid content
of the autumn queens was 39.9%, and the crop-solid of males was 34.5%, of their dry weight. The
seasonal change in worker dry weight was related to the varying work loads of the workers. A
simulation model using the compartmental system approach was used to estimate the number of
workers, queens and males produced by a successful colony. Investment in workers, either as dry
weight or calorific equivalent units, represented nearly 50% of the total colony investment. This
is considered a relatively high investment and can be related to the delay of sexual production.
Males and queens are produced in about equal numbers but, in terms of dry weight or calorific
equivalent units, there is a bias toward queen production. Workers were found to be selectively
destroying male brood probably derived from queen eggs. MacNair's hypothesis could not be
supported.

In England the annual life-history of
Vespula vulgaris (Linn.) starts with the
emergence of the males and new queens
from the mature colonies in the autumn.
The sexuals remain in the colonies for a
few days before leaving on their mating
flights. After a short time the fertilised
queens enter over-wintering sites while
the males die. When the queens emerge in
the spring they each search for a nest site,
build a queen-nest and, by early June, rear
the first workers. The workers take over
the jobs of building the nest and looking
after the brood from the queen which con-
tinues to lay eggs. At first, the workers
build small cells (small-cell colony) in
which more workers, and later, males are
reared. From the beginning of August, the
workers build large cells (large-cell colo-
ny) in which the queens and sometimes
males are reared. The colony with its
queen and workers usually dies by the

end of October or early November (Archer
1981a, 1984).

With the emergence of the new queens,
brood in the small cells is neglected and
not well fed (Montagner 1966). The ne-
glected larvae and sealed brood are fre-
quently pulled out of their cells by the
workers and dropped some distance from
the nest. This destruction of brood has
been frequently recorded (Duncan 1939,
Spradbery 1973, Archer 1981a, Greene
1991). The neglected brood also are eaten
by dipterous larvae, e.g. Volucella sp.
(Archer 1981a).

The above life-history in which the re-
productive females fly away from their
natal colonies and found new colonies in-
dependently is called 'Alate Dispersing'
(Nonacs 1993). All alate dispersing colo-
nies must make two investment decisions.
The first is concerned with the proportion
of resources devoted either to the produc-

258

Journal of Hymenoptera Research

tion of the workers or to the sexuals. The
second is concerned with the proportion
of resources devoted to either male or
queen production (Nonacs 1993).

The first decision is a process of max-
imising sexual production. Sexual produc-
tion could be delayed by the production
of more workers for colony maintenance
with the trade-off consequence of ensur-
ing a greater future sexual production.
Brian (1965) produced a general model of
colonial growth which showed that queen
production increases with increased in-
vestment in workers. Pamilo (1991) devel-
oped models for perennial colonies, find-
ing that greater investment in worker pro-
duction was related to a smaller chance of
queens founding new colonies and the old
colonies surviving. With the annual colo-
nies of V. vulgaris, the old colonies do not
survive and the queens have a very small
chance of founding new colonies (Archer
1984) so worker production relative to
sexual production should be higher. The
timing of the switch from worker to sexual
production varies between the species of
vespine social wasps (Archer 1980, 1981a).

The second decision concerning propor-
tional investments in males and queens fo-
cuses on the queen-worker conflict due to
asymmetrical genetical relationships
(Trivers and Hare 1976, Benford 1978,
MacNair 1978, Nonacs 1986, Boomsma
1989, Ratnieks and Reeve 1992). A wide
range of variation in sex investment ratios
has been found (Crozier and Pamilo 1996),
often due to multiple mating by queens
and worker reproduction in queenright
colonies (Trivers and Hare 1976, Benford
1978).

Usually sex ratio investment is mea-
sured by determining the dry weight pro-
duction of queens and males (Trivers and
Hare 1976, Crozier and Pamilo 1996). In a
vespine colony with a singly-mated queen
and no worker reproduction, it is expected
that queens will adjust the sex ratio in-
vestment to give equal dry weights of
queens and males. Workers, because of

asymmetrical genetical relationships, will
increase the investments in queens to
three times that of males. If the queen has
mated with more than one male and /or
there is worker reproduction, the sex ratio
investment of the queen is unchanged, but
the workers will increase their investment
in the males, although investment in
queens will still be greater.

MacNair (1978) argued that queen in-
vestment stops after the eggs are laid: fur-
ther investment then is carried out by the
workers. Thus the queen should distribute
her parental investment equally between
queen and male eggs to produce a pri-
mary sex ratio of 1:1. For workers to skew
the investment towards queens, they
should try to prevent the queen from lay-
ing male eggs or destroy the brood de-
rived from the male eggs of the queen.

The queens have larger bodies in which
to carry sufficient fat bodies to enable
them to over-winter. The males do not
over-winter but die soon after mating, or
attempting to mate, for which purpose a
smaller body seems adequate. Thus the
departure from equal investment in queen
and male eggs by the queen to a relatively
greater queen investment does not neces-
sarily imply the workers have succeeded
in altering the sex ratio investment of the
queen. To demonstrate that workers have
succeeded in producing a relatively great-
er investment in queens, it is necessary to
show that male brood derived from queen
eggs have been destroyed.

Ideally, the determination of worker and
sexual production should be carried out di-
rectly by visual observations on colonies un-
der natural conditions. This is difficult to
achieve since colonies of V. vulgaris are sur-
rounded by envelopes, are often under-
ground, and the workers are aggressive
when disturbed during investigations. Vi-
sual observations on healthy colonies main-
tained in an observation box have so far
been restricted to one comb, the lowest
comb of the nest (Potter 1964).

However, indirect methods can be used

Volume 7, Number 2, 1998

259

to estimate the production of workers and
sexuals. Colonies can be collected
throughout the year and counts made of
the brood and adults present. At the end
of the last larval stage the gut contents are
evacuated to form the black meconium at
the bottom of the cell. Counts of these me-
conia can be used as an estimate of the
number of adults reared. Since a meconi-
um is only evidence that a larva has pu-
pated, the result could be an over-estimate
if incipient adults died during the pupal
stage. In addition, the meconial remains
do not reveal the sex or caste of the former
occupants of the cells. Thus precise counts
of worker and sexual production cannot
be made from collected colonies, although
counts of meconia and queen and male
sealed brood can give a first approxima-
tion (Archer 1993).

To improve the accuracy of estimating
worker and sexual production a simula-
tion model of a successful colony, i.e. one
that rears many queens, has been devel-
oped (Archer 1981a, 1981b). The simula-
tion model not only used the data from
198 collected colonies of V. vulgaris but
also laboratory observations on the length
of life of the brood stages and adult work-
ers. The model incorporates meconial in-
formation for adult production, and rates
of cell building, brood stage addition and
adult appearance estimated from the col-
lected data. Brood neglect and mortality
were incorporated into the model to come
into effect when adult and brood stages
became too numerous in comparison with
the data from collected colonies. The mod-
el achieves greater realism at the expense
of complexity, with the use of 253 param-
eters and variables. To handle such a com-
plex model the compartmental system ap-
proach (Odum 1971) was used.

In this paper, I will try to determine the
extent of any destruction of queen-derived
male brood to test the proposal of Mac-
Nair (1978), and derive estimates of the
production of workers, males and queens
to determine the relative importance of

worker production, and the relative in-
vestment in males and queens.

METHODS

Sources of, and treatment of workers. —
Workers were bait-trapped during 1970 at
Averhams Plantation, between Flaxton
and Claxton, about 12 kilometres to the
north-east of York, England. Averhams
Plantation was an open site with a dense
herb layer and recently planted conifers.
The trapping station consisted of 16 sub-
stations arranged in a square (4 X 4) with
two traps at each sub-station. The traps
were attached to canes about 70cm above
the ground. Each trap consisted of a white
polythene container (75mm deep, 75mm
diameter) with a 10mm diameter hole in
the lid, and contained a jam solution with
added yeast. The fermenting jam solution
was changed once every two weeks and
the catch of workers collected once a
week. Trapping was continuous from July
until October. The workers were pre-
served in 70% ethanol.

Samples of 50 workers from each week-
ly catch were dried to constant weight in
an air oven at 60°C. During July and Oc-
tober when the number of workers
trapped were smaller all workers were
dried. Each weekly sample of dried work-
ers was weighed to O.lmg. The weekly dry
weight was divided by the number of
workers in the sample to give the mean
worker dry weight.

Workers from seven colonies were col-
lected during late June and preserved by
deep-freezing. Later the dry weights of
these workers were obtained as previous-
ly described except that workers were
weighed individually so a standard devi-
ation could be calculated.

Correction for dry weight of workers due to
70% ethanol preservation. — To determine if
ethanol-preserved workers lost dry
weight, samples of 50 or 55 workers from
three colonies were preserved in 70% eth-
anol and by deep-freezing. After about six
months of such preservation dry weights

260

Journal of Hymenoptera Research

were obtained as previously described.
Ethanol preserved workers showed a loss
of dry weight compared with deep-frozen
preserved workers from all three colonies.
The percentage dry weight loss was
20.5%.

Sources of, and treatment of queens. — Au-
tumn queens with fully developed fat
bodies were collected from four colonies.
From three colonies 172 queens were pre-
served by deep-freezing and from one col-
ony 47 queens were preserved in 70% eth-
anol. Dry weights were obtained as pre-
viously described for workers except that
queens were weighed individually. Dry
weight loss from ethanol preservation was
found to be 11.4% based on queens col-
lected from one colony when 55 queens
were preserved in 70% ethanol and 63 by
deep-freezing.

Sources of, and treatment of males. — Males
with full crops were collected from a col-
ony which was rearing males both in the
small and large cells. All 102 males were
preserved in 70% ethanol. Dry weights
were obtained as previously described for
workers except that males were weighed
individually. The dry weights of the males
showed a bimodal distribution indicating
a weight difference between males reared
in the small and large cells. A method giv-
en by Lewis and Taylor (1967) was used
to separate the 56 small-cell and 46 large-
cell reared males.

Males with full crops were collected
from three colonies which had reared
males only in the small cells. From the
three colonies 146 males were preserved
by deep-freezing. Dry weights were ob-
tained as previously described for workers
except that males were weighed individ-
ually.

Dry weight loss from ethanol preserva-
tion was found to be 37.7% based on
males collected from three colonies when
201 males were preserved in 70% ethanol
and 146 by deep-freezing.

Lipid determinations. — Samples of work-
ers, autumn queens and males were pre-

served by deep freezing and extracted in
the Soxhlet apparatus using trichlorethyl-
ene as a solvent. Extraction was continued
until constant weight was obtained. Nine
queens were extracted individually, 27
males in batches of threes, and 74 workers
from three colonies in colony batches.

Male crop content determinations. — When
males left their natal colonies in the au-
tumn their crops were found to contain a
clear viscous fluid. From one colony 50
males were collected and preserved by
deep freezing. Their crop fluid was col-
lected by cutting the gaster away from the
rest of the body and gently squeezing the
gaster so that the crop fluid could be ab-
sorbed by a known dry weight of filter pa-
per. The filter paper with its absorbed
crop fluids were dried in an air oven at
60°C to constant weight. All weightings
were made to O.lmg. Some general bio-
chemical tests were performed on the crop
fluid.

Ash content of workers, autumn queens and
males. — A sample of 22 autumn queens
was collected, preserved by deep-freezing,
and dried by freeze-drying. Samples of 74
workers and 27 males were obtained from
the lipid-extracted individuals. The sam-
ples were heated in a furnace at 500°C for
three hours and the residue ash weighed
to O.lmg.

Calorific determinations. — Samples of
workers, autumn queens with fat bodies
and males with full crops were preserved
by deep freezing, dried by freeze-drying
and their calorific values determined with
the aid of a Phillipson Oxygen Microbomb
Calorimeter (Phillipson 1964).

Due to the high lipid content of the au-
tumn queens oil was lost when the animal
tissue was pelleted in preparation for
bomb calorimetry. Attempts to bomb non-
pelleted material as suggested by Wood-
land et al. (1968) and Howell and Fisher
(1977) failed because oil was lost from the
sample and found at the bottom of the
bomb. The lipids were extracted, as pre-

Volume 7, Number 2, 1998
25

124

23

Q

^22

o

5 21

c
n)

(D

5 20

19

July

August ! September

Fig. 1. Mean worker dry weight (mg) versus date
for Vt'sp'ula viilgnris.

viously described, and the calorific deter-
minations made on the remaining tissues.

Difficulties sometimes were experienced
when freeze-drying males in that the vis-
cous contents of the crops would not
freeze-dry but rather exploded out of the
male into the freeze-drying equipment.
Calorific determinations were made on
crop-extracted males, as previously de-
scribed, and on males with crop fluids
where freeze-drying was satisfactorily
completed.

Simulation model of colonial determina-
tion.— The development of a simulation
model is given in Archer (1981a). The
model was developed on the DEC system-
10 computer at the University of York, En-
gland. Recently the model has been trans-
ferred to a Personal Computer.

RESULTS

Seasonal variation of worker weights. — The
seasonal change in the dry weights of
workers from the bait traps is shown in
Fig. 1. These dry weights have been cor-
rected for dry weight loss during ethanol
preservation. From a high mean dry
weight during early July there is a de-
crease to a low value during August, fol-
lowed by an increase during the first half
of September, after which there is a slight
decrease during late September and Oc-
tober.

261

Table 1. The mean dry weights of workers of Ves-
puln viil^nrif from late July colonies.

Dale

No,
workers

Mean dry
weight (m's)

standard
deviation

17

13

25.1

3.43

20

14

22.0

4.11

29

33

21.2

5.24

30

15

24.6

5.59

30

34

21.9

5.09

30

35

24.4

5.66

The mean dry weights of the workers
from the late June colonies are given in
Table 1. The mean dry weight from the
seven colonies is 23.3mg which is slightly
higher than the value for early July (Fig

Lipid content of workers, autumn queens
and males. — The lipid content of the work-
ers is given in Table 2. The higher lipid
content corresponds with the higher mean
dry body weight found during September.

The mean hpid content of a male was
7.4mg (range 4.1-9.1) representing 10.3%
of the mean dry body weight.

The mean lipid content of a queen was
65.0mg (s.d. 8.2) representing 39.9% of the
mean dry body weight.

Male crop fluid. — The mean dry weight
of the crop fluid was 32.3mg (s.d. 9.98)
which represented 60.5% of the wet
weight of the crop fluid. The mean dry
weight of the crop fluid represented 34.5%
of the mean dry body weight. The dry
weight of the crop fluid of each male was
positively correlated with its dry body
weight (n = 50, r = 0.57, p < 0.001).

The crop fluid gave a positive result
with two general carbohydrate tests: (a)

Table 2. The lipid content of adult workers of Ves-
puta vulgaris.

.\o
workers

evtracted ".. lipid ot

(nig) dry bodv weight

20 Jul.
26 Aug.
23 Sep.

20

25

23.0

16.7

43, S

5.7

3.1

10.1

262

Journal of Hymenoptera Research

Table 3. The mean dry weight of autumn queens
of Vespula vulgaris.

Table 4. The mean dry weight of small-cell reared
males of Vespuhi inilgnrif.

No,
queens

Mean dry
weight

Standard
deviation

Dale

No, males

Mean drv
weight (mg)

Standard
de\iation

Date

12 Sep.
16 Sep.
23 Sep.
31 Oct.

50
56
50
46

81.1
87.3»
96.3
74.0

12 Sep.

20 Sep.

9 Oct.

31 Oct.

22
47
59
91

163.7
167.1*
167.6
155.3

11.18

11.75

10.43

8.43

14.3

14.7
23.0
13.9

' Corrected for 70% ethanol preservation.

alcoholic thymol and cone, hydrochloric
acid and (b) alcoholic alpha-naphol and
concentrated sulphuric acid. A positive re-
action also was obtained with Benedict's
reagent indicating the presence of reduc-
ing sugar.

Queen dry weight. — The mean dry
weights of autumn queens with fat bodies
are given in Table 3. The closeness of the
deep-freeze and corrected ethanol-pre-
served queens gives confidence in the use
of the correction factor. The mean of the
four samples is 163.4mg. Since these au-
tumn queens consisted of 39.9% lipid con-
tent the mean dry weight would be made
up 65.2mg lipid and 98.2mg non-lipid
substances. The dry weight of lipid of
each queen was positively correlated with
its dry body weight (n = 19, r = 0.79, p
< 0.0001).

Male dry weight. — The mean dry weights
of males reared in the small cells are given
in Table 4. The closeness of the deep-
freeze and corrected ethanol-preserved
males gives confidence in the use of the
correction factor. The dry weight of the
four samples is 84.7mg. Since the crop sol-
id consisted of 34.5% of the body dry
weight, the mean dry weight would be
made up of 29.2mg crop solid and 55.5mg
for the rest of the body.

The mean dry weight of males reared in
large cells was 130.7mg. This dry weight
has been corrected for dry weight loss
during ethanol preservation. Assuming
34.5% of this dry weight was crop solid,
the mean dry weight would be made up

of 45.1mg crop solid and 85.6mg for the
rest of the body.

Calorific determinations. — The ash con-
tent of the adults was so low that it was
unlikely to complicate the calorific deter-
mination. The ash percentage of mean dry
weight for autumn queens was 2.8%, for
males 2.9% and for workers 2.5%.

The mean calorific values of lipid-ex-
tracted queens, males and workers are
given in Table 5. Assuming the calorific
equivalent of queen lipid to be 9.2 cal/mg
(Sawicka-Kapusta 1975) and knowing the
lipid percentage of mean dry body weight
then the calorific value of the total dry
body weight of the queen would be 6.627
cal/mg dry weight.

Knowing the mean calorific values of
crop-extracted males and males with
crops (Table 5), it can be calculated that
the crop solid has a calorific value of 4.232
cal/mg dry weight. Such a calorific value
indicates that the crop solid was a protein
or carbohydrate rather than a lipid sub-
stance. Earlier results showed that the

Table 5. Calorific values of autumn queens, males
and workers of Vespula vulgaris.

Sample

Cal/mg

Standard

size

dry weight

deMJtion

Queen (lipid extracted)

10

4.820

0.085

Male (crop extracted)

10

5.201

0.149

Male (with crop)

10

4.872

0.142

Worker— 17Jun.

10

5.092

0.122

Worker — 28 Jul.

10

5.273

0.147

Worker — 24 Sep.

10

5.220

0.109

Worker — combined

30

5.195

0.145

Volume 7, Number 2, 1998

263

Table 6. The number, biomass (dry weight) and
calorific equivalent of queens, queen lipid, males,
male crop content and workers produced in a simu-
lation colony of Vespula vulgaris.

Table 7. The dry weights of workers of Vespula
vulgaris during the seasonal development of a colony
derived from Fig. 1 and Spradbery (1972).

CalorilR

^■L]ui\alenl

\ umber

Biomass (mj;)

(cal)

Workers

10,248

217,072.3

1,127,690.6

Males

Small cells

763

64,626.1

314,858.4

Large cells

261

34,112.7

166,197.1

Total

1,024

98,738.8

481,055.5

Crop solids

34,064.9

144,162.6

Queens

969

158,334.6

1,049,283.4

Lipid

62,985.0

579,462.0

Total

12,241

474,145.7

2,658,029.5

crop solid is a carbohydrate, probably re-
ducing sugar.

Since the calorific values of workers var-
ies little from samples collected during
June, July and September a combined val-
ue has been calculated (Table 5).

The mean calorific values for workers
and crop-extracted males are similar, but
less than the mean calorific value of au-
tumn queens because of the high lipid
content of queens. The low mean calorific
value of males is due to the high carbo-
hydrate content of the crop. The lipid-ex-
tracted queens have a lower mean calorific
value than the workers and crop-extracted
males, as these workers and males have
some lipid in their bodies.

Production estimates of a successful colo-
ny.— The number, biomass and calorific
equivalents of the workers, males and
queens produced in the simulation model
are given in Table 6. The biomass and cal-
orific equivalents of the males and queens
were calculated by multiplying the num-
ber of males and queens produced by the
model by the appropriate mean dry
weight and calorific values. The crop solid
of the males represents 30.0% of their cal-
orific equivalent and the lipid of the
queens 55.2% of their calorific equivalent.

Since the dry weight of workers varies
during the development of the colony

Dale

Dr\' weight (mg)

6 Jun.

11.75 (Spradbery)

16 Jun.

11.75 (Spradbery)

6 Jul.

22.4

11 Aug.

19.5

14 Sep.

24.9

5 Oct.

24.3

(Fig. 1), it is necessary to know the dry
weight of the workers on the days that
they emerged as adults. These emergence
dry weights were assumed to be those of
the collected workers, half the length of
worker life before the date on which the
workers were collected (Archer 1981a).
The model calculated the number of adult
workers produced each day which was
multiplied by the appropriate mean dry
weight derived from Fig. 1 and given in
Table 7. Increases and decreases in mean
dry weight were assumed to be linear. The
total worker dry weight produced could
be multiplied by worker mean calorific
value to give the total calorific equivalent.

The relative investment in workers,
males and queens of a successful colony
is given in Table 8. Just over five workers
are needed to rear each sexual, but be-
cause sexuals are heavier than workers a
greater biomass or calorific equivalent of
sexuals is produced than for workers.
Workers represent 45.8% by dry weight
and 42.4% by calorific equivalent of the to-
tal production of the colony.

Slightly more males than queens are

Table 8. The relative investment in workers, males
and queens in a simulation model of a successful col-
ony of Vespula vulgaris.

Biomass cainnlic

Number (dry weight) equivalent

Sexuals: Workers
Queens: Workers
Queens: Males

1:5.14

1:10.58

1:1.06

1:0.84
1:1.37

1:0.62

1:0.74
1:1.07

1:0.46

264

Journal of Hymenoptera Research

reared, but, because queens are heavier
and of higher calorific value, the sex ratio
investment changes markedly in favour of
the queens. Because the males have higher
metabolic rates than queens, the energetic
cost ratio (Boomsma 1989, Bourke and
Franks 1995) can be used. The sex ratio
investment then becomes one queen to
0.72 male, still indicating an investment in
favour of queens.

DISCUSSION

Worker dry weight. — A similar seasonal
change in the dry weight of workers of V.
vulgaris from early July until October (Fig.
1) was found by Spradbery (1972) in En-
gland and by Malham (1996) in New Zea-
land for the equivalent season. Spradbery
(1972) also found a very low worker dry
weight during June (Table 7): these were
queen-reared workers. Brian and Brian
(1952) also found that queen-reared work-
ers of Dolichovespula sylvestris (Scopoli)
had low weights.

Malham (1996) found that in areas
where insecticide had been used to dras-
tically reduce the number of workers, the
dry weight of workers during March
(equivalent to September in England, Fig.
1) was markedly higher than in untreated
areas. This difference was less pronounced
earlier in the season and had disappeared
by the end of the season. Malham (1996)
attributed the difference to food availabil-
ity. In treated areas, relatively more food
would be available per forager, so larvae
would receive more food and produce
workers of a heavier weight.

The low worker dry weight during Au-
gust (equivalent to February in New Zea-
land) varied from 12-13mg (Malham 1996)
to 17.4mg (Spradbery 1972) and 19.5mg in
the present study. The high worker dry
weight during September (equivalent to
March in New Zealand) varied from 17-
20mg (Malham 1996) to 20.8mg (Sprad-
bery 1972) and 24.9mg in the present
study. Following Malham (1996) these
variabilities in dry weights could be due

to shortages in food supply as forager
density increases.

The variation in worker dry weight also
can be related to variation of the work
load on workers. The work load will de-
pend on the number of larvae to feed (lar-
va/worker ratio), the number of cells to
build, and the amount of soil to be exca-
vated to make the cavity for the nest. Time
spent in excavation and building could re-
duce the time available to feed the larvae.

The low weights of queen-reared work-
ers could be a consequence of very high
larvae per queen ratio, up to 30 larvae per
queen, and high cell building rates, in ex-
cess of two cells per day (Archer, unpub-
lished).

The relatively rapid increase in worker
dry weight by late June until early July
could be a consequence of workers aiding
the queen in brood rearing. At this time
the larva /worker ratio rapidly decreases
to about 3-4 and the cell building rate to
about one cell per worker per day (Archer
1981a).

The decline in worker dry weight from
early July until August coincides with the
development of the small-cell colony dur-
ing which a large worker population is
reared (Archer 1981a). Workers sampled
during August also had the lowest lipid
content (Table 2). Despite the exponential
growth of the small-cell colony, the work
load on workers continues to decrease
with larva /worker ratio decreasing to
about one, and cell building rate per day
per worker approaching zero. However
the amount of excavation greatly increases
and over 90% of the outgoing workers
from a colony may be carrying earth par-
ticles (Archer, unpublished).

The increase in dry weight from August
until September coincides with the devel-
opment of the large-cell colony when the
future sexuals are reared. Workers sam-
pled during September also had the high-
est lipid content (Table 2). During this
time the worker load remains low. Larva/
worker ratio remains at about one, and

Volume 7, Number 2, 1998

265

large cell building rate per worker per day
is very low at about 0.04 (Archer 1981a).
Soil excavation continues but usually less
than 20% of outgoing foragers are carry-
ing earth particles (Archer, unpublished).

Queen dry weight. — Spradbery (1973)
and Harris and Beggs (1995) found that
nearly 40% of the dry weight of autumn
queens was lipid, which is similar to the
value given earlier in this paper. The same
authors found that the lipid was used as
a food source of which about three-quar-
ters was used during the over-wintering
period. By dissecting queens of Vespa affin-
is (Linn.) from southern Japan, Martin
(1993) found that the contents of the fat
bodies were used up during the over-win-
tering period of four to five months.

Harris and Beggs (1995) found the mean
dry weight of autumn queens from New
Zealand was 121.7mg (range 108.0-154.5),
which is about 26% lower than the mean
weight reported in the current study. They
suggested the low weight of autumn
queens was because the fat bodies of the
queens had not reached their maximum
level of lipid storage. The low weight of
queens also could be a consequence of the
very high colony densities: up to 33 colo-
nies per hectare found in New Zealand,
compared with up to about two colonies
per hectare in England (Edwards 1980). At
lower colony densities relatively more
food resources might be available for
queen rearing.

Male dry weight. — By measuring the
wing length of males of Vesya crabro Linn.,
Potter (1964) also found a bimodal distri-
bution of male size. Measurements were
carried out on callow adults found in their
cells. He also recorded that males of some
colonies of V. vulgaris showed a bimodal
size distribution but gave no numerical
details. Potter's observations support the
interpretation of the bimodal size distri-
bution of males presented in this paper.

The carbohydrate food reserve found in
the crops of the males seems to be a new
observation. The function of this food re-

serve would be to provide a readily avail-
able source of energy needed by the males
when flying around their mating circuits
(Edwards 1980).

Worker-sexual ratio investment. — Invest-
ment in workers represents nearly 50% of
the total biomass investment of a colony
(Table 8). By reference to Pamilo (1991, Ta-
ble 2) the investment in the workers can
be considered to be relatively high. This
large investment in workers can be related
to the delay of sexual production until
September in V. vulgaris, with the conse-
quence of a larger output of queens (Brian
1983). Sexual production in Dolichoz>espnila
sylvestris (Scopoli) occurs earlier, during
July, so this species has a relatively small-
er investment in workers, and also a
smaller output of queens (Edwards 1980).

Sex ratio investment. — The more-or-less
equal production of queens and males in
V. vulgaris (Table 8) seems rather surpris-
ing since at the sealed brood stage, the
number of males is usually twice the num-
ber of queens (Archer 1981a, Greene 1991).
However, Archer (1981b) found that 45%
of large-cell male sealed brood was de-
stroyed, and observed large-cell male
sealed brood and mature larvae were car-
ried away from the colonies by the work-
ers. If these large-cell sealed brood are in-
cluded in the calculation of the primary
sex ratio, the ratio becomes one queen to
1.28 males.

The simulation model allows for this
destruction of large-cell larvae and sealed
brood, as well as for the neglect and de-
struction of small-cell larvae and sealed
brood. The output of the model indicates
that 18.4% small-cell larvae and sealed
brood will be neglected and destroyed.
Since this destruction occurs in the later
part of colonial development, most of the
destroyed small-cell brood will be males
(Archer 1981a). Thus the primary sex ratio
would be even more biased towards the
males.

The interpretation of the above obser-
vations and calculations would indicate

266

Journal of Hymenoptera Research

that the queen is not laying an equal num-
ber of male and queen eggs as MacNair
(1978) suggested. Since males are smaller
than queens, the queen would seem to be
laying relatively more male eggs so that
the investment in adult queens and males
eventually becomes equal.

It is possible that the extra male eggs are
derived from the workers and the destruc-
tion of the male brood an example of
worker policing against male production
by other workers (Ratnieks 1988), which is
predicted when the queen mates with
more than two males. Multiple matings by
queens of V. vulgaris are highly likely
(Page 1986). However, Ross 1986 and
Bourke 1988, failed to find evidence of
Vespula workers laying eggs in queenright
colonies.

The workers would appear to be de-
stroying male brood so as to bias invest-
ment towards the queens. However the
sex-ratio investnient does not reach three
queens for every male (Table 8) probably
because of multiple mating by the queen
(Page 1986).

The destruction of males by the workers
does not take place until the male brood
has reached the mature larval and sealed
brood stages. Thus, the queen would seem
to be able to disguise the sex of her off-
spring during the egg and early larval
stages. MacNair (1978) proposed that
there would be an evolutionary race be-
tween the queens and workers, with genes
selected which favour the disguise of the
sex of the brood, followed by genes which
enable the disguise to be penetrated. The
evolutionary race seems to have reached
an equilibrium with detection occurring at
the late larval and sealed brood stages af-
ter the workers have made a considerable
investment in rearing males.

Brood destruction. — Brood neglect and
destruction has been linked to the death
or physiological breakdown of the queen,
leading to the disintegration of the social
life of the colony (Spradbery 1973). How-
ever, the queen brood are not neglected

but well fed (Montagner 1963), and selec-
tive destruction of male brood occurs in
the large cells (Archer 1981b).

One consequence of the loss of queen
influence is the appearance of a domi-
nance struggle among the workers (Mon-
tagner 1966), whose ovaries start devel-
oping (Greene 1991). Workers with devel-
oped ovaries lay eggs destined to become
males, although due to the lateness of the
season it is unlikely these males will be
reared, or if reared, will successfully mate
(Ross 1985). Probably the capacity of
workers to rear males from worker eggs
is an adaptive response to the premature
death or physiological breakdown of the
queen (Ross 1985). About 28% of the col-
onies surviving until at least September
(Archer, unpublished) are unsuccessful in
producing many queens (Archer 1981b),
but nevertheless rear males in the small
cells. Many of these males could be de-
rived from worker eggs.

In conclusion the outcomes of the two
investment decisions of successful colo-
nies of V. vulgaris have been found as fol-
lows. The first decision of the trade-off be-
tween worker production for colony
maintenance and sexual production is to
delay sexual production in order to pro-
duce more workers and hence to produce
relatively more sexuals. In the second de-
cision, there is a bias towards queen pro-
duction at the expense of male production.
This bias depends on worker action in the
destruction of male brood probably de-
rived from queen eggs.

ACKNOWLEDGMENTS

Robin Edwards made many helpful comments on

this manuscript.

LITERATURE CITED

Archer, M. E. 1980. Population dynamics: 172-207, in
Edwards, R. Social Wasp-,. The Rentokil Library,
East Grinstead.

Archer, M. E. 1981a, A simulation model tor the colonial
development of Pamvcfpuhi vitl^nri^ (Linnaeus) and
Dolichovct^puk sy'i'csfn'.s (Scopoli) (Hymenoptera:
Vespidae). Mclimdcrm 36: 1-59.

Volume 7, Number 2, 1998

267

Archer, M. E. 1981b. Successful and unsuccessful devel-
opment of colonies of Vesj.nda vulgaris (Linn.) (Hy-
menoptera; Vespidae). Ecohgicnl Eiilciniclogy 6: 1-10.

Archer, M. E. 1984. Life and fertility tables for the wasp
species Vespnda vulgaris and Dolichoi'espula syhvstris
(Hymenoptera: Vespidae) in England. Entomologia
Generally 9: 181-188.

Archer, M. E. 1993. The life-history and colonial char-
acteristics of the Hornet, Vespa crabro L. (Hym., Ves-
pinae). Entomologist's monthly Magazine 129: 151-
163.

Benford, F. A. 1978. Fisher's theory of the sex ratio ap-
plied to the social Hymenoptera. journal of theoreti-
cal Biology 72: 701-727.

Boomsma, J. J. 1989. Sex-investment ratios in ants: has
female bias been systematically overestimated? The
American Naturalist 133: 517-532.

Bourke, A. F. G. 1988. Worker reproduction in tlie high-
er eusocial Hymenoptera. Quarterly Review of Biol-
ogy 63: 291-312.

Bourke, A. F. G. and Franks, N. R. 1995. Social ei'olution
in ants. Princeton University Press, U.S.A.

Brian, M. V. 1965. Social Insect Populations. Academic
Press, London.

Brian, M. V. 1983. Social bisects. Ecology/ and Beltaz'ioural
Biology. Chapman & Hall, London.

Brian, M. V. and Brian, A. D. 1952. The wasp, Vespula
sylvestris Scopoli: feeding, foraging and colony de-
velopment. Transactions of the Royal entomological So-
cieh/ of London 103: 1-26.

Crozier, R. H. and Pamilo, P. 1996. Evolution of social
insect colonies. Sex allocation and kin Selection. Oxford
University Press, Oxford.

Duncan, C. D. 1939. A contribution to the biology of
North American Vespine wasps. Stanford Unii'ersity
Publications, Biological Science 8: 1-272.

Edwards, R. 1980. Social wasps. Their biology and control.
The Rentokil Libran', East Grinstead, England.

Greene, A. 1991. Dolichovesputa and Vespula: 263-305, in
Ross, K. G. and Matthews, R. W. (eds) The Social
Biology of Wasps, Cornell Universit)' Press: 263-305.

Harris, R. J. and Beggs, J. R. 1995. Variation in the qual-
ity of Vespula vulgaris (L.) queens (Hymenoptera:
Vespidae) and its significance in wasp population
dynamics. Netv Zealand loumal of Zoology 22: 131-
142,

Howell, ]. and Fisher, R. C. 1977. Food conversion effi-
ciency of a parasitic wasp, Nemeritis connexens. Eco-
logical Entomology 2: 143-151.

Lewis, T. and Taylor, L. R. 1967. Introduction to experi-
mental ecology/. Academic Press, London.

MacNair, M. R. 1978. An ESS for the sex ratio in ani-
mals, with particular reference to the social Hy-
menoptera. Journal of theoretical Biology 70:449-459.

Malham, J. 1996. How much does that wasp weigh?
Wasp Times 24:2.

Martin, S. J. 1993. Weight changes in adult hornets, Ves-
pa affinis (Hymenoptera: Vespidae). Insectes sociaux
40: 363-368.

Montagner, H. 1963. Relations entre les adultes et le cou-
vain chez les guepes sociales du genre Vespa, au
moyen d'un radio-isotOf>e. Insectes scoiaux 10: 153-
165.

Montagner, H. 1966. Sur le determinisme du couvain
abortif dans les nids de guepes du genre Vespa.
Compte rendu hebdomadaire des seances de I'Acadeniie
des sciences, Paris 263: 826-829.

Nonacs, P. 1993. The effects of polygyny and colony life
history on optimal sex investment: 110-131, in Kel-
ler, L. (ed.) Queen Nundter and Socialihi in Insects.
Oxford Scientific Publications, Oxford.

Odum, E. P. 1971. Fundamentals of ecology. Saunders,
London.

Page, R. E. 1986. Sperm utilization in social insects. An-
nual Review of Entomology 31: 297-320.

Pamilo, P. 1991. Evolution of colony characteristics in
social insects. 1. Sex allocation. Tlie American Natu-
ralist 137: 83-107.

Phillipson, J. 1964. A miniature bomb calorimeter for
small biological samples. Oikos 15: 130-139.

Potter, N. B. 1964. A study of the biology of the Common
Wasp, Vespula vulgaris L., with spvcial reference to the
foraging beliaviour. Unpublished Ph.D. thesis. Uni-
versity of Bristol, England.

Ratnieks, F. L. W. 1988. Reproductive harmony via mu-
tual policing by workers in eusocial Hymenoptera.
American Naturalist 132: 217-236.

Ratueks, F. L. W. and Reeve, H. K. 1992. Conflict in
single-queen Hymenoptera societies: the structure
of conflict and processes that reduce conflict in ad-
vanced eusodal species, journal of theoretical Biology
158:33-65.

Ross, K. G. 1985. Aspects of worker reproduction in four
social wasp species (Insecta: Hymenoptera: Vespi-
dae). Journal of Zoology, London (A) 205: 411424.

Ross, K. G. 1986. Kin selection and the problem of sperm
utilization in social insects. Nature 323: 798-800.

Sawicka-Kapusta, K. 1975. Fat extraction in the Soxhlet
apparatiis: 288-292, in Klekowski, G. IBP Handbook
No 24, Methods for Ecological Bio-energetics. Black-
well, Oxford.

Spradbery, J. P. 1972. A biometric study of seasonal vari-
ation in worker wasps (Hymenoptera: Vespidae).
Journal of Entomology (Series A) 47:61-69.

Spradber)', ]. P. 1973. Wasps. An account of the biology and
natural history of social and solitary waspis. Sidgwick
and Jackson, London.

Trivers, R. L. and Hare, H. 1976. Haplodiploidy and the
evolution of the social insects. Science 191: 249-263.

Woodland, D. J., HaU, B. K. and Calder, J. 1968. Gross
bioenergetics of Blaltella germanica. Physiological Zo-
ology 41:424-431.

J. HYM. RES.
Vol. 7(2), 1998, pp. 268-273

Response of Glyptapanteles militaris (Walsh) (Hymenoptera:

Braconidae), a Larval Parasitoid of the Armyworm,

Mythimna unipuncta (Haworth) (Lepidoptera: Noctuidae), to

Different Temperatures

L. Oliveira, R. Melo and J. Tavares

Universidade dos Azores, Departamento de Biologia, P-9502 Ponta Delgada Codex,

Azores, Portugal

Abstract. — The effect of four different temperatures (15, 20 25 and 30°C), on biological param-
eters of the Azorean population of Gh/ptapantelcs militaris (Walsh) was studied, using Mythimna
unipuncta (Haworth) as the host. Thirteen biological parameters of the host-parasitoid interaction
were analysed: percentage of hosts that died without producing parasitoids; percentage of larvae
parasitized from which parasitoids emerged; percentage of hosts surviving to pupate after the
parasitoid's sting; egg-larval development time; pupal period; total developmental time; adult
longevity; total number of larvae per host; number of larval parasitoids that fail to emerge from
each host; mean number of parasitoids that emerged from host larva but failed to spin a cocoon;
mean number of cocoons per host; parasitoid sex-ratio; and finally emergence rate of adult par-
asitoid progeny. The percentage of hosts that died without producing parasitoids increased with
increasing temperature. Developmental times significantly decreased with increasing temperature.
The mean number of cocoons per host; mean number of parasitoids that emerged from each host
larva but failed to spin a cocoon; and total number of larvae per host were higher when the
temperature was lower. Parasitoid sex ratio and emergence rate of adult progeny were not affected
by the temperatures tested.

Mythimna unipuncta (Havk^orth) is the
most important pest in Azorean pastures.
Serious population explosions that require
the use of pesticides often occur (Tavares
1992). Glyptapanteles militaris (Walsh) is a
larval parasitoid of the armyworm in all
islands of the Archipelago (Oliveira 1996).
It is desirable to increase the natural pop-
ulation of G. militaris by field releases of
wasps produced in the laboratory, during
the first generation of M. unipuncta.

One of the most important abiotic fac-
tors that affect insects is temperature. In
parasitoids this can influence develop-
ment, fecundity, mortality, sex ratio, col-
oration and other characteristics in vari-
ous species (Kaya and Tanada 1969; Yu
and Luck 1988; Klein 1988; Lysyk and
Nealis 1988; Spivac et al. 1992).

Temperature increases, within a favour-

able range, will speed up insect metabo-
lism and consequently increase the rate of
development. Each species and each stage
in the life history may develop at its own
rate (Sedlacek et al 1990, Spivac et al. 1992,
Gullan and Cranston 1994).

A previous study of the effect of two
different temperatures on the biological
parameters of G. militaris was performed
by Oliveira (1991, 1992). In the present
study, we analyse the effect of two ex-
treme temperatures (15 and 30°C) and two
intermediate temperatures (20 and 25°C),
on some biological parameters of the Azo-
rean population of G. militaris using M.
unipuncta as host.

MATERIAL AND METHODS

Glyptapanteles militaris used in this ex-
periment emerged from naturally parasit-

Volume 7, Number 2, 1998

269

ized M. unipuncta larvae, collected in pas-
tures of Sao Miguel island. Groups of one
hundred cocoons were placed in 400 ml
glass vials until adult females were re-
moved for experiments. Adults were sup-
plied with honey solution (10%). We used
as hosts M. unipuncta larvae from labora-
tory cultures, established from eggs laid
by field-collected females.

On the third day after adult parasitoid
emergence, one isolated female wasp was
allowed to parasitize one isolated third in-
star larva of M. unipuncta. After the first
sting, the host was removed from the par-
asitoid and individually kept, until emer-
gence of the parasitoids, in a plastic con-
tainer (4.5 X 3cm). Each host larva was
supplied with a small piece (Icm^) of ar-
tificial diet every two days as described by
Poitout and Bues (1970) and modified by
Oliveira (1991). After parasitization, each
group of fifty parasitized host larvae were
kept at a different temperature (15±0.5°C,
20±0.5°C, 25±0.5°C and 30±0.5°C), under
75±0.5% R.H. and 16:8 [L:D] photoperiod.

After larval parasitoid emergence and
construction of the cocoons, each group
was maintained in a plastic container (4.5
X 3cm) with a hole covered by nylon tis-
sue. Emerged adults were kept in the
same conditions and were supplied with
honey solution (10%) until their death.

Thirteen biological parameters of the
host-parasitoid interaction were analysed:
percentage of larvae parasitized from
which parasitoids emerged; percentage of
dead hosts; percentage of hosts surviving
to pupate after the parasitoid's sting; egg-
larval development time; pupal period; to-
tal developmental time; adult longevity;
total number of larvae per host; number
of larval parasitoids that fail to emerge
from each host; mean number of parasit-
oids that emerged from host larva but
failed to spin a cocoon; mean number of
cocoons per host; parasitoid sex-ratio (per-
centage of females); and finally emergence
rate of adult progeny.

The first three parameters were ana-

100

I parasitized hosts
B dead hosts
[]] pupated hosts

2U 2.";

Tempei^ture (°C)

Fig. 1. Percentages of pupated, dead and parasitized
larvae of Mf/thiinna tiinpuncta, at four different tem-
peratures (±standard error). A test for multiple com-
parison of proportions was used. Each column that
is followed by a ciifferent letter is significantly differ-
ent (p < 0.05).

lysed by a test for multiple comparisons
of proportions (Zar 1996), and the others
parameters were analysed by non-para-
metric "Kruskal-Wallis" and Multiple
Comparison tests (p < 0.05) (Scherrer
1984) to compare the results obtained
from different temperatures. To compare
the effect of temperature on all studied pa-
rameters, a discriminant factorial analysis
(Thioulouse 1989), was performed.

RESULTS AND DISCUSSION

Parasitized larvae of M. unipuncta were
able to develop normally at the four dif-
ferent temperatures, with high percent-
ages of hosts that were successfully para-
sitized and producing parasite progeny
(Fig. 1). However the percentage of para-
sitized larvae from which parasitoids
emerged differed significantly between 15
and 30°C (t = 1.983, p < 0.05), achieving
maximum values at 15°C.

The percentage of hosts that died before
emergence of the parasitoids increased
with temperature. It was least at 15°C, in-
termediete and similar at 20 and 25°C, and
most at 30°C fFig. 1). A significant differ-
ence was observed for the percentage of
dead hosts at 15 and 30°C (t = 2.280, p <
0.05). The percentage of surviving hosts
after they had been stung by the parasit-

270

Journal of Hymenoptera Research

Table 1. Mean {±standard deviation) days of egg-larval development time in Mytlniiiini innpuucla (Dl),
pupal period (D2), and the total developmental time (Dl + D2) of Glyp^tapantelcs luilitaris, at four different
temperatures.

Temperature
°C

D2
X ± sd

15

40

37.50 ± 4.90a

40

18.25 ± 1.89a

40

55.75 ± 5.87a

20

34

19.00 ± 2.89b

33

07.27 ± 1.15b

33

26.18 ± 3.07b

25

34

13.56 ± 2.14c

31

05.84 ± 0.69c

31

19.36 ± 2.32c

30

31

11.90 ± 2.01c

20

05.45 ± 1.47c

20

17.15 ± 2.01c

H value

118.495

96.684

108.677

P value

<0.0001

O.OOOl

<0.0001

Kruskal-Wallis H and P values and Multiple Comparisons test. Means in each column that are followed by
a different letter are significantly different (p < 0.05).

oid was very low (2^ %) and therefore
they were not statistically analysed.

The relation between temperature and
rate of development of poikilotherms is an
important aspect of ecological studies and
basic to the development of pest manage-
ment strategies (Lysyk and Nealis 1988).
According to several authors, the period
between parasitization and parasitoid
emergence decreases with increasing tem-
perature (Nealis and Fraser 1988; Gould
and Elkinton 1990; Allen and Keller 1991;
Tillman and Powell 1991). A similar result
was obtained in this study with G. militaris
and M. unipuncta. We divided the total de-
velopment period in two parts: egg-larval
development time, and pupal develop-
ment (strictly, duration of the cocooned
stages). We found similar results for both
periods (Table 1), and significant differ-
ences were observed between the different
temperatures with only one exception (25
and 30°C).

Temperature significantly affected the
longevity of adult G. militaris obtained in
this study except between 20 and 25°C. A
similar result was obtained by Allen and
Keller (1991) in a study of Cotesia urabne
Austin and Allen reared from Uraba lugens
Walker. The maximum longevity of G.
militaris was obtained at 15°C (8.1 days)
and the minimum was at 30°C (2.3 days).
At the intermediate two temperatures we
obtained 2.9 days. Comparing these val-
ues with the 16 days of maximum longev-

ity previously observed by Oliveira (1996),
we conclude that our results are very low
and they may be due to abiotic conditions,
such as an insufficient level of ventilation
in climatic chambers.

The temperature established during the
development time of G. militaris can affect
the mean number of cocoons found per
host, the mean number of parasitoids that
emerged from the host larva but failed to
spin a cocoon, and the total number of dis-
cernable parasitoid larvae. The mean
number of cocoons per host decreased
with increasing temperature. A significant
difference was observed between 30°C
and 15 and 20°C (Table 2). Similar results
were reported by Oliveira Filho and
Foerster (1986) with "Apanteles" muesebecki
Blanchard parasitising Pseiidaletia sequax
Franclemont. The mean number of para-
sitoid larvae that emerged from the host
but then failed to spin a cocoon was rela-
tively small in each case; the differences
found (Table 2), are, though statistically
significant, hard to explain in view of the
non-linearity of the result and they may
not really be informative. The number of
parasitoid larvae that failed to leave the
host larva was similar at the four temper-
atures tested (Table 2). The total number
of larvae per host decreased with increas-
ing temperature (Table 2). A significant
difference was found between the result
obtained at 30°C and the other three tem-
peratures.

Volume 7, Number 2, 1998

271

Table 2. Mean (±standard deviation) number of Gli/ptnfuintcles ruilitiiris cocoons, the number of larvae that
failed to spin a cocoon, and that failed to emerge from the host of Mythimna unipuncia, at four different
temperatures.

lfmptT.itiirf

.\

N^ coccons / host

Par lar\'ae

not pupated

X ± sd

I'ar, larvae

not emerged

X ± sd

total larvae
V ± sd

15

40

55.40 ± 38.24ab

8.50 ± 13.06ab

10.00 ± 16.46a

73.95 ± 31.43a

20

34

50.41 ± 28.71b

2.50 ± 04.86a

8.91 ± 14.83a

61.82 ± 29.25a

25

34

35.12 ± 27.81bc

8.94 ± 13.55ab

12.24 ± 21.84a

56.29 ± 29.21a

30

31

19.42 ± 16.29c

5.55 ± 04.25b

9.19 ± 12.33a

34.16 ± 21.49b

H value

27.36

12.84

3.20

30.19

P value

<0.0001

0.0050

0.3616

<0.0001

Kruskal-Wallis H and P values and Multiple Comparisons test. Means in each column that are followed by
a different letter are significantly different (p < 0.05).

The sex ratios (percentage of females)
were lower than 19%, in all cases (Table
3). No significant differences as a function
of temperature were found. Similar results
were obtained by Kolodny-Hirsch (1988)
in his study of Cotesia melanoscela (Ratz-
burg) and the host Lymantria dispar (L.). In
all laboratory cultures of the Azorean G.
militaris population we systematically ob-
tained low sex ratios (Oliveira 1991, 1992,
1996), when compared with the observed
sex ratio of natural populations, usually
between 48 and 78% (Oliveira 1991, 1996).
This is due to a high percentage of the fe-
males that oviposit failing to produce fe-
male progeny, indicating a high level of
unmated females under laboratory condi-
tions. This is an important aspect that will
require to be overcome, if we want to do

Table 3. Clyptafmntele^ militaris sex-ratio (percent-
age of females) and adult emergence rates (±stan-
dard deviation), at four different temperatures.

Tempera*
ture °C

Se\ ratio

"u Emergence

N

X ± sd

N

X ± sd

15

40

0.19 ± 0.29

40

70.90 ± 21.80

20

33

0.16 ± 0.23

34

63.00 ± 25.60

25

31

0.18 ± 0.25

34

70.30 ± 25.80

30

21

0.13 ± 0.22

31

49.80 ± 40.70

H value

0.595

5.012

P value

0.8976

0.1709

Kruskal-Wallis H and P values (p > 0.05).

mass releases of G. militaris to control M.
unipuncta.

The adult emergence rates were not
very high, always less than 71%, but no
significant differences were observed be-
tween temperatures (Table 3). However,
an interesting result was observed at 30°C
with a high number of adults dying before
complete emergence from the cocoons.

To compare the effect of the different
temperatures on all parameters a discrim-
inant factorial analysis was performed.
This analysis demonstrated: 1) a super-
position of the values obtained at 25 and
30°C; 2) a light separation at 20°C; 3) a
complete separation of the results at 15°C
(Fig. 2). The parameters that had most in-
fluence on this separation were the egg-
larval development time, the pupal peri-
od, adult longevity, and the number of co-
coons per host.

Finally, these results confirm that G. mil-
itaris is very well adapted to the climatic
conditions of the Azores, since the para-
sitoid develops better between 15 and
20"C than at 25 and 30°C, and the temper-
atures found on the Azores pastures range
between 13 and 23°C during the period of
highest activity of the parasitoid.

ACKNOWLEDGMENTS

This research u'as performed on CIRN (Centro
para a Investiga^ao dos Recursos Naturais) and fi-
nanciallv supported bv the Universidade dos Azores,

272

Journal of Hymenoptera Research

; ^30^ ■■..

\ \25| \

15° '•,

Axis 1 (% of inertia=98.5)

Fig 2. Discriminant factorial analysis performed
with nine biological parameters of Glyptapanteles mil-
itaris: egg-larval development time, pupal period,
adult longevity, mean number of cocoons per host,
mean number of parasitoids that emerged from host
larva but failed to spin a cocoon, mean number of
larval parasitoids that failed to emerge from each
host, total number of larvae per host, parasitoid sex
ratio, and emergence rate of adult progeny.

by the Secretaria Regional da Agricultura e Pescas,
and by the Program STRIDE (# ST RDB/C/AGR/
194/92).

LITERATURE CITED

Allen, G. R. and M. A. Keller 1991. Urabn lumens (Lep-
idoptera: Noctuidae) and its parasitoids (Hyme-
noptera: Braconidae): temperature, host size, and
development. Environmental Entomology, 20 (2),
458-469.

Gould, ]. R. and J. S. Elkinton 1990. Temperature-de-
pendent growth of Cotesia melanoscela (Hyme-
noptera: Braconidae), a parasitoid of the Gypsy
moth (Lepidoptera: Lymantriidae). Environmental
Entomology. 19 (4), 859-865.

Gullan, P. J. and P. S. Cranston 1994. The Insects: An
Outline of Entonwlogi/. Chapman & Hall. London.
1-491.

Kaya, H. and Y. Tanada 1969. Responses to high tem-
perature of the parasite Apmnteles niilitaris and of
its host, the armyworm, Pseudaletia unipimcta.
Annals of the Entomological Society of America, 62
(6), 1303-1306.

Klein, M. 1988. Colour morphs induced under con-
trolled environmental conditions in adult Earias
insulana (Lepidoptera: Noctuidae). Environmental
Entomology , 17: 162-165.

Kolodny-Hirsch, D. M. 1988. Influence of some en-
vironmental factors on the laboratory production
of Cotesia melanoscela (Braconidae: Hymenop-
tera): a larval parasitoid of Lymantria dispar. En-
vironmental Entomology , 17 (1), 127-131.

Lysyk, T. J. and V. G. Nealis 1988. Temperature re-
quirements for development of the jack pine
budworm (Lepidoptera: Tortricidae) and two of
its parasitoids (Hymenoptera). Journal of Econom-
ic Entomology, 81 (4): 1045-1051

Nealis, V. G. and S. Fraser 1988. Rate of development,
reproduction, and mass-rearing of Apanteles fii-
miferanae Vier. (Hymenoptera: Braconidae) under
controlled conditions. Cauadiau Entomologist, 120
(3), 197-204.

Oliveira, L. 1991. Bioecologia de Apanteles mUitaris
(Walsh, 1861) (Hymenoptera, Braconidae). Protias
de acesso « categoria de Assistente de bwestiga(;do,
Universidade dos Agores, 1-72.

Oliveira, L. 1992. Influencia da temperatura no cicio
biologico de Apanteles militaris (Walsh) (Hyme-
noptera, Braconidae). Actas do V Congresso Ib-
erico de Entomologia. Boletim da Sociedade porlii-
guesa de Entomologia, 2 (3), 357-362.

Oliveira, L. 1996. Apanteles mditaris (Walsh) (Hyme-
noptera, Braconidae) parasitoide das larvas de
Myihimna unipuncta (Haworth) (Lepidoptera,
Noctuidae). Tese de Doutoramento, Universidade
dos Azores, 1-196.

Oliveira Filho, J. and L. A. Foerster 1986. Ciclo evo-
lutivo e preferencia para oviposi^ao de Apanteles
muesehecki Blanchard, 1947 (Hymenoptera: Bra-
conidae), parasitoide de Pseudaletia sequax Fran-
clemont, 1951 (Lepidoptera: Noctuidae). Anais da
Sociedade de Entomologia do Brasd, 15 (2), 371-377.

Poitout, S. and R. Bues 1970. elevage de plusieurs es-
peces de Lepidopteres Noctuidae sur milieu ar-
tificial riche et sur milieu artificiel simplifie. Ann
Zool Ecol Anim., 2: 79-91.

Scherrer, B. 1984 Biostatistique. Gaetan Morin editeur.
Quebec, 1-850.

Sedlacek, J. D., K. V. Yeargan and P. H. Freytag 1990.
Effect of temperature on the development of the
blackfaced leafhopper (Homoptera: Cicadelli-
dae). Environmental Entomology, 19: 209-214.

Spivac, M., A. Zeltzer, G. Degrandi-Hoffman and J.
H. Martin 1992. Influence of temperature on the
rate of development and colour patterns of
queen honey bees (Hymenoptera: Apidae). En-
vironmental Entomology, 21 (2): 364-370.

Tavares, J. 1992. A importancia econtimica da lagarta
das pastagens Mythimna unipuncta (Haworth)
(Lep., Noctuidae). Aforeana, 7 (3), 407^14.

Tillman, P. G. and J. E. Powell 1991. Developmental
time in relation to temperature for Microplitis cro-
ceipes, M. demolitor, Cotesia kazak (Hymenoptera:
Braconidae) and tlyposoter didymator (Hymenop-
tera: Ichneumonidae), endoparasitoids of the to-

Volume 7, Number 2, 1998

273

bacco budworm (Lepidoptera: Noctuidae). Etwi-

ronmcntal Entoitwlcgy, 20 (1), 61-64.
Thioulouse, J. 1989. Statistical analysis and graphical

display of multivariate data on the Macintosh.

Cominiter Apptlicntions in the Biosciences, 5, 287-

292.
Yu, D. S. and R. F. Luck 1988. Temperature-depen-

dent size and development of California red
scale (Homoptera: Diaspididae) and its effect on
host availability for the ectoparasitoid, Aphitis
melinus DeBach (Hymenoptera: Aphelinidae).
Environmental Entomologii, 17 (2), 154-161.
Zar, J. H. 1996. Biostattstical analysis. Prentice-Hall In-
ternational Editions, London, 1-662.

]. HYM. RES.
Vol. 7(2), 1998, pp. 274-279

Response of Pithecellobium tortum Martius (Leguminosae) Seeds to

the Attack of the Phytophagous Braconid AUorhogas dyspistus

Marsh (Hymenoptera: Braconidae)

Margarete Valverde de Macedo, Marina C. P. Pimentel, and
RiCARDO Cardoso Vieira

(MVM, MCPP) Laboratorio de Ecologia de Insetos, Departamento de Ecologia, CP 68020, IB,

Universidade Federal do Rio de Janeiro, ILha do Fundao, Rio de Janeiro, Brasil,

CEP 21941-590; (MVM) Departamento de Zoologia, CP 6109, Universidade Estadual de Campii\as,

Campinas, Sao Paulo, Brasil, CEP 13083-970; (RCV) Departamento de Botanica,

Universidade Federal do Rio de Janeiro, Ilha do Fundao, Rio de Janeiro,

Brasil, CEP 21941-590

Abstract. — This work describes morphological changes on the seeds of Pithecellobium tortum
caused by the braconid AUorhogas di/spistus, at Restinga (Costal scrub) of Barra de Marica, Rio de
Janeiro State, Brazil. This species was shown to be a gall maker on P. Tortum seeds whose galls
result from the proliferation of parenchymatous cells near the tegument but not of the seed coat
cells. Its impact on the host plant consists of decreases in plant reproductive potential not only
by directly reducing seed viability, but also by contributing to seed mortality via the adult emer-
gence hole which allows invasion by pathogenic micro-organisms. The braconid's way of eating
the seed, keeping itself in a chamber apart from the seed embryo, which remains alive and there-
fore demanding nutrients, accords it the profile of a "manipulative parasite" in the sense of Weis
& Abrahamson (1986).

Signs of insect herbivory on plants vary
greatly. Some are simple feeding marks
left on the host plant which normally do
not involve any apparent morphological
response. Other signs, however, are very
complex, resulting from a noticeable mor-
phological and /or physiological response
of the plant. This response may be defen-
sive, pathological or one which benefits
the herbivore (Price 1980, Weis & Abra-
hamson 1986). Herbivores that are capable
of manipulating the response of their host
plant for their own benefit have been
called "manipulative parasites" (Weis &
Abrahamson 1986). Gall makers induce
the development of localised growing
structures resulting from the abnormal in-
crease in number and /or size of plant cells
(Darlington 1975). Normally, the galls are
induced in undifferentiated tissues, which
have their development manipulated

(Weis et ah 1988). The gall phenotype is
the result of two genotypes: the one of the
gall maker, responsible for the stimulus,
and the other of the plant, which produces
the reaction (Abrahamson & Weis 1987).
From an evolutionary perspective, gall
morphology is the product of natural se-
lection on the insect stimulating the de-
velopment of a structure for protection
and nutrition and on the plant resisting or
trying to avoid the insect stimulus (Weis
et al. 1988).

Most of the known entomogenous galls
are induced by Diptera (especially Ceci-
domyiidae), Hymenoptera, Homoptera
and Thysanoptera (Meyer 1987; Short-
house and Rohfritsch 1992). Within the
Hymenoptera the Cynipidae is the most
important family, but there are also re-
cords for Tenthredinidae, Eurytomidae,
Eulophidae, Pteromalidae and Tanaostig-

Volume 7, Number 2, 1998

275

matidae (Fernandes 1987). Guimaraes
(1957) reared Allorhogas muesebecki Gui-
maraes from Aneniopaegnin miraiidutn
Alph. DC. (Bignoniaceae) galls. Because
this braconid was the only species to
emerge the author concluded that this was
the gall inducer. Since the first record of a
phytophagous braconid (Macedo & Mon-
teiro 1989) and its specific description by
Marsh (1991) as Allorhogas dyspisttis, Infan-
te et nl. (1995) redescribed Monitoriella elon-
gata Hedqvist and recorded it as a new
case of phytophagy within the Braconidae.
This species, like A. dyspistiis, belongs to
the Doryctinae, a group which, according
to Wharton (1993), should be searched for
more cases of phytophagy. Ramirez &
Marsh (1996) described two new Psenobol-
us species (Braconidae: Doryctinae) which
appear to develop as inquilines on plant
tissue in fig flowers after their pollination
by Agaonidae wasps. More recently, Aus-
tin & Dangerfield (1998) recorded the bi-
ology of Mesostoa kerri Austin and Whar-
ton, a member of the endemic Australian
subfamily Mesostoinae, as a new case of
galling Braconidae.

Pithecellobium tortum Martius (Legumi-
nosae) seeds are enclosed in fruits contain-
ing about 30 seeds side by side. They are
attacked by the braconid Allorhogas dyspis-
tiis Marsh, which oviposits directly into
immature seeds, when abundant endo-
sperm and a small embryo are still pres-
ent. After oviposition by the braconid the
seed divides internally and externally, re-
sulting in an intact region, joined to the
funicle, where the seed embryo is usually
found (Macedo & Monteiro 1989). In
many cases this region of the seed contin-
ues growing even after the adult insect
has emerged. More than one A. dyspistus
can be found in a single seed. In these
cases, more than one division occurs and
still a single intact region within the seed
embryo is found. The main purpose of this
study is to describe and discuss the mor-
phological seed changes caused by Allor-
hogas dyspistus in Pithecellobium tortum and

to evaluate this impact of the insect on the
host plant.

MATERIALS AND METHODS

All seeds were collected at Restinga
(Coastal scrub) de Barra de Marica
(22°57'S and 12° 52'W), Marica county, Rio
de Janeiro state, Brazil.

To investigate seed tissue morphology,
intact and attacked seeds were collected
during June 1994, and fixed in 70% alco-
hol. Seeds were then laid in 10% ethyle-
nediamine for one week, as suggested by
Carlquist (1982) to soften them. After
washing in distilled water three times for
a 2 hours period each time, the seeds were
dehydrated in ethanolic series and then
embedded in paraffin wax following Jo-
hansen (1940). Longitudinal serial 15 |xm
sections were obtained with a rotary mi-
crotome and stained with basic blue astra-
fucsin (Roeser 1962).

In 1994, a further sample of 2990 seeds
from 150 fruits obtained from five P. tor-
tum individuals were collected and dis-
sected in order to evaluate the rate of A.
dyspistus attack and to check if the at-
tacked seeds died or continued develop-
ment until complete maturation. In 1995,
327 fruits from 12 plant individuals were
also collected and dissected to evaluated
A. dyspistus seed attack rate.

At the end of the 1995 fruiting season in
June, mature fruits of P. tortum were col-
lected from their parent plant and from
the ground. Attacked and non-attacked
seeds were then obtained to perform the
tetrazolium viability test (Delouche et al.
1962). This test was performed immediate-
ly after the collection of seeds from the
plant and from the ground, and also after
three and eight months of laboratory stor-
age at room temperature for seeds collect-
ed from the ground. All tested seeds were
cut lengthwise and one half of each seed
was completely immersed in 0.5% chlo-
ride of 2,3,5-triphenyl tetrazolium solution
and the other half boiled before being sub-
mitted to this tetrazolium test of viability.

276

Journal of Hymenoptera Research

Fig. 1. Longitudinal sections of PitheceUohium tortiini
seeds, a) Non-attacked seed with its embryo (em); b)
seed attacked by two AUcrlwgaf lii/fpistus individuals,
showing two attacked regions (ar); a larva (la) can be
observed in one of them. The embryo (em) is present
at the center in the preserved region (pr); c) an at-
tacked region showing the seed coat palisade tissue
(pt) which does not cover all of the region involving
the larva. Note the thicker parenchymatous layer (pi).

This procedure was necessary in order to
eliminate the possibility that the red col-
oration of the embryo was due to reduced
ions and not to hydrogenases produced
during the respiration process of the living

embryo. If the dead boiled embryo col-
oured, the test would be invalid. All beak-
ers were kept in darkness at room tem-
perature and the result checked after 12
hours.

RESULTS

Comparison between non-attacked (Fig.
la) and attacked (Fig. lb) seeds shows that
the tissue associated with A. dyspistus lar-
vae probably results from the proliferation
of the fundamental parenchymatous cells
near the inside tegument. The gall, the re-
gion resulting from tissue proliferation,
keeps the insect larva apart from the seed
embryo through a clear division of the
seed. The seed coat does not cover the en-
tire proliferated region where the braconid
larva is found (Fig. Ic). This can also be
seen by the naked eye because the texture
of this attacked region is clearly different
from that where the embr^'o is found.

Allorbogas di/spistus attacked 55.85% of
the dissected seeds. In all, only 6.1%, out
of 1670 attacked seeds appeared healthy
after insect emergence or death. All other
attacked seeds died mainly because of
contamination, probably fungus, which
probably entered the seed through the A.
dyspistus or its parasitoid exit holes.

All mature attacked and non-attacked
seeds collected from the plant itself were
viable according to the tetrazolium test, as
well as all mature non-attacked seeds col-
lected from the ground (Table 1). A small-
er proportion of attacked seeds were via-
ble in the three-months stored group and
an even smaller proportion of the attacked
seeds were viable in the eight-months
stored group. It is worth noting that in all
groups the embryos from the intact seeds
exhibited a clearly darker pink coloration
in comparison with the viable embryos
from the attacked seeds. No boiled seeds
were coloured thus validating the above
results.

Allorfwgas dyspistus seed attack reached
more than 70% in five out of the 12 plants

Volume 7, Number 2, 1998

277

Table 1. Percentage of viable seeds according to
tetrazolium test of attacked and non-attacked seeds
under four situations of collection and storage. The
number of tested seeds is in parentheses.

Collection / situation of
the tested seeds

Non-attacked Attacked
seeds seeds

From plant /immediately after

collection 100(50) 100(22)

Ground /immediately after col-
lection 100(25) 100(18)

Ground /after three months of

storage 100 (20) 73 (15)

Ground /after eight months of

storage 100(6) 18(11)

Studied and in two of these the rates were
very near 100% (Table 2).

DISCUSSION

Clear cell proliferation, characteristic of
the process of gall formation, indicates
that A. dysTpistus induces galls in P. tortum
immature seeds. This kind of gall is, ac-
cording to Gagne (1994), a simple gall,
which does not differ from the normal
plant tissue except for cell proliferation.
This is very different from most of the
complex galls {sensu Gagne 1994) on plant
vegetative parts {e.g. Redfern & Askew
1992). Monitoriella elongata (Infante et al.
1995) and Mesostoa kerri (Austin and Dan-
gerfield 1998), are two Braconidae leaf
gallers whose galls are apparently as com-
plex as any Cecidomyiidae leaf gall and
quite different from the simple galls of A.
di/spistiis. As seeds are attacked when im-
mature and still in the process of devel-
opment, meristematic tissue is available,
and this is where galls are primarily pro-
duced (Mani 1964, Bronner 1977). Quicke
(1997) points out that there is growing ev-
idence that at least seed predation actually
involves the stimulation of proliferation of
some plant cells, that is, a kind of incipient
gall formation or simple gall (sensK Gagne
1994).

Other insect groups which develop in-
side seeds, such as Bruchidae (Coleop-
tera), do not promote any similar response

Table 2. Numbers of seeds collected (N), attacked
seeds and percentage of seed attack by AUorhogai dys-
f)is/i(s on each of the 12 plant individuals sampled in
1995.

All.ii.ked
seeds

"n attack
.4, i/i/s;'i5M<s

1

359

41

11.42

2

288

33

11.46

3

386

130

33.68

4

339

83

24.48

5

252

26

10.32

6

365

290

79.45

7

394

274

69.54

8

558

39

6.99

9

385

307

79.74

10

468

436

93.16

11

436

435

99.77

12

338

337

99.70

and most of them feed upon embryo and
cotyledons (Southgate 1979). Even Riiw-
chenus stigma (L.) (Coleoptera; Curculion-
idae), which eats the cotyledons but not
the embryo of Hymenaea (Leguminosae)
seeds, does not induce any kind of seed
division (T.M. Lewinsohn pers. com.).

The way of attack by the braconid leav-
ing the seed embryo region intact may be
considered a way of maintaining nutrient
demand by the living seed and therefore
for the insect, which could not survive
otherwise. The aggregated pattern of A.
dyspistus occurrence in the fruits (M.V.
Macedo, pers. obs.) could, if the seeds
died, cause selective abortion of most at-
tacked fruits, as in Cassia grandis L. (Le-
guminosae) attacked by bruchids (Janzen
1971). This negative effect for the insect is
more evident in younger fruits (Stephen-
son 1981). Thus, A. dyspistus may be con-
sidered a manipulative parasite of P. tor-
tum immature seeds (sensu Weis & Abra-
hamson 1986).

It is clear that the potential negative ef-
fect is very high where reproductive tissue
is attacked (Abrahamson & Weis 1987).
Reduction of P. tortum seed viability over
time is a clear effect of A. dyspistus attack
(Table 1). Furthermore, the capability of
germination of attacked living seeds still

278

Journal of Hymenoptera Research

remains to be tested. The tetrazolium test
only proved that attacked seeds are alive;
these seeds, however, could have lower
chances of germination and establishment,
as may be suggested by attacked seeds
having a lower degree of respiratory activ-
ity which was shown by their lighter col-
oration in the test. Moreover, A. dyspistus
exit holes seem to serve as a communicat-
ing channel between the seed and the ex-
ternal environment, which makes it pos-
sible for micro-organisms to enter and
eventually kill most of the attacked seeds.
This work shows that, directly or indi-
rectly, A. dyspnstus considerably reduces
plant fitness, and, because the species may
attack up to 100% of the seeds in a plant,
it can be considered a good example of a
galler that imposes high negative effects
on its host plant. We suggest that the gall-
er's behaviour of maintaining the seed
embryo alive while feeding upon seed tis-
sue may avoid abortion of immature seeds
increasing the galler's chances of survival.

ACKNOWLEDGMENTS

The authors are indebted to Paul Marsh, Gislene
Ganade, Mark Shaw, Carlos Roberto Brandao, Tho-
mas Lewinsohn, Rogerio Parentoni Martins, Ricardo
Monteiro, Eric Grissell and an anonymous reviewer
for their suggestions on the manuscript, and to Andy
Austin for his special help in sending his manuscript.
Conselho Nacional de Desenvolvimento Cientifico e
Tecnologico (CNPq), Fundagao Coordenaqao de
Aperfeigoamento de Pessoal de Ni'vel Superior
(CAPES), Funda^ao de Amparo a Pesquisa do Estado
do Rio de Janeiro (FAPERJ), Conselho de Ensino para
Graduados (CEPG/UFRJ) and Funda^ao Jose Boni-
facio (FUJB/UFRJ) provided the financial support.

LITERATURE CITED

Abrahamson, W. G, & Weis, A. E. 1987. Nutritional
ecology of arthropod gall makers, pp. 235-258.
In Slansky Jr, F. & Rodriguez, J. G. (eds.) Nutri-
tional ecology of insects, mites and sptitiers. John Wi-
ley & Sons. New York.

Austin, A. D. & Dangerfield, P. C. 1998. Biology of
the Mesostoa kerri Austin and Wharton (Insecta:
Hymenoptera: Braconidae: Mesostoinae), an en-
demic Australian wasp that causes stem galls on
Banksia marginala Cav. Australian journal of Bota-
ny, in press.

Bronner, R. 1977. Contribution a I'etude histoqui-

mique des tissus nourriciers des zoocecidies.
Marcellia 40: 1-34

Carlquist, S. 1982. The use of Ethylenediamine in soft-
ening hard plant structures for paraffin section-
ing. Stain Technology 57: 311-317.

Darlington, A. 1975. The pocket encyclopaedia of plant
galls. Blandford Press.

Delouche, J. C, Still, T. W., Raspet, M & Lienhard,
M. 1962. The tetrazolio test for seed viability.
Miss. Agric. Exp. Stn. Tech. Bull. 51: 1-63.

Fernandes, G. W. 1987. Gall forming insects: their
economic importance and control. Rei'ista brasi-
leira de Entomologia 31: 379-398.

Gagne, R. J. 1994. The gall nudges of the neotropical re-
gion. Cornell University Press. Ithaca.

Guimaraes, J. A. 1957. Contrihui(;ao ao estudo da Ceci-
dologia Brasiliana. Tese Escola Nacional Agron-
omia. (UFRRJ), Rio de Janeiro.

Infante, F.; Hanson, P. & Wharton, R. 1995. Phyto-
phagy in the genus Monitoriella (Hymenoptera:
Braconidae) with description of new species. An-
nals of the Entomological Society of Anu^rica 88: 406-
415.

Janzen, D. H. 1971. Escape of Cassia grandis L. beans
from predators in time and space. Ecology 52:
964-979.

Johansen, D. A. 1940. Plant nucrotechiuque. McGraw
Hill, New York.

Macedo, M. V. & Monteiro, R. F. 1989. Seed predation
by a braconid wasp, Allcrhogas sp. (Hymenop-
tera). journal of the Nezv York Entomological Society
97: 358-362.

Mani, M. S. 1964. The ecology of plant galls. The Hague,
Junk.

Marsh, P. M. 1991. Description of a phytophagous
Doryctine Braconid from Brasil (Hymenoptera:
Braconidae). Proceedings of the Entomological So-
ciety of Washington 93: 92-95.

Meyer, J. 1987. Plant galls and gall inducers. Borntrae-
ger, Berlin.

Price, P. W. 1980. Evolutionan/ biology of parasites.
Princeton University Press. Princeton.

Quicke, D. L. J. 1997. Parasitic wasps. Chapman &.
Hall, London.

Ramirez, W. B. & Marsh, P. M. 1996. A review of the
genus Psenobolus (Hymenoptera: Braconidae)
from Costa Rica, an inquiline fig wasp with bra-
chypterous males, with description of two new
species. Journal of Hymenoptera Research 5: 64-72.

Redfern, M. & Askew, R. R. 1992. Plant galls. Rich-
mond Publishing Co. Ltd.

Roeser, K. R. 1962. Die Nadel der Schwarzkiefer-Mas-
sen Produkt und Kunstwert der Natur. Mikrokos-
mos 61: 33-36.

Shorthouse, J. D. & Rohfritsch, O. 1992. Biology of in-
sect induced galls. Oxford University Press, Ox-
ford.

Volume 7, Number 2, 1998

279

Southgate, B. J. 1979. Biology of the Bruchidae. An-
nual Review cf Entomology 24: 449^73.

Stephenson, A. G. 1981. Flower and fruit abortion:
proximate causes and ultimate functions. Annual
Revieiv of Ecologi/ and Si/stematics 12: 253-279.

Weis, A. E. & Abrahamson, W. G. 1986. Evolution of
host-plant manipulation by gall makers: ecolog-

ical and genetic factors in the Soliiiago-Euro^ta
system. American Naturalist 127: 681-695.

Weis, A. E., Walton, R. & Crego, C. L. 1988. Reactive
plant tissue sites and the population biology of gall
makers. Annual Rn'ieiv of Entomology 33: 467-486.

Wharton, R. A. 1993. Bionomics of the Braconidae.
Annual Rei'iezv of Entomology 38: 121-143.

J. HYM. RES.
Vol. 7(2), 1998, pp. 280-295

Morphological Caste Differences in the Neotropical Swarm-founding

Polistine Wasps IV: Pseudopolybia vespiceps, with Preliminary

Considerations on the Role of Intermediate Females in the

Social Organization of the Epiponini

(Hymenoptera, Vespidae)

SULENE NORIKO ShIMA, FERNANDO BaRBOSA NOLL, RONALDO ZUCCHl AND

SoiCHi Yamane

(SNS) Departamento de Zoologia e Centro de Estudos de Insetos Socials. Institute Biociencias,

Universidade Estadual Paulista-UNESP, 13506-900 Rio Claro (SP), Brazil; (FBN, RZ)

Departamento de Biologia, Faculdade de Filosofia, Ciencias e Letras de Ribeirao Preto,

Universidade de Sao Paulo— USP,14049-901— Ribeirao Preto (SP), Brazil; (SY) Biological

Laboratory, Faculty of Education, Ibaraki University Mito, 310-8512 Japan

Abstract. — The measurements of 22 body parts and counts of hamulus number for 300 wasps
from a colony of Pseudopoh/hia vespiceps evidenced the occurrence of three types of females, i.e.,
queens (inseminated egg-layers), workers (uninseminated, non egg-layers), and intermediates
(with well developed ovaries but uninseminated). Caste-linked aspects in this species are, thus,
complex. Queens (Q) and workers (W) differed in only two characters (basal width of the second
gastral tergite and hamulus number) (Bonferroni-test, p < 0.05), so, these castes can be considered
as morphometrically similar. On the other hand, intermediates (I) were significantly larger than
queens and workers in some body parts (p < 0.05). Canonical discriminant analysis revealed that
Mahalanobis distances (D-) between Q/W, Q/I and I/W were very small, 0.99, 1.51 and 1.12,
respectively. In spite of this, color patterns of head and gastral parts separate queens and workers
very well. In such aspects intermediates combine queen-worker similarities.

In social insects, approaches related to than workers in most of the morphologi-

caste differences constitute a form of evi- cal characters (Protonectarina sylveirae: Shi-

dencing queen-worker specializations. To ma et al., 1996b). Moreover, the taxa under

study the diversity of such aspects, the such opposite trends showed also differ-

swarm-founding polistine wasps are very ent color patterns allied to conspicuous

helpful because the castal differences so external morphological peculiarities,

far known can be arranged along a spec- However, the most pronounced dimor-

trum ranging from taxa in which queens phism among polistine wasps has been

and workers are externally similar, to oth- found in some Agelaia (Stelopoh/bia auct.)

ers with fairly distinct caste attributes. De- species, such as, A. flavipeniiis (Evans &

spite occurence of transitional steps (e.g. West-Eberhard 1970); A. areata (Jeanne

Protopolybia exigiia cf. Simones 1977; Noll and Fagen 1974; Jeanne 1980 1991), A. pal-

et al. 1996), our previous serial papers lipes and A. multipicta (Noll et al. 1997) and

showed two contrasting cases: queen A. vidua (Sakagami et al. 1996; Baio et al.

dwarfism, as promoted by the prevalence in press).

of smaller body parts in queens relatively The genus Pseudopolybia is poorly known,

to workers {Afioica flavissima and Poh/bia Indeed, only some details on nest architec-

dimidiata: Shima et al. 1994 and Shima et ture and fragmentary morphometric fea-

al. 1996a, respectively), and queens larger tures had so far been reported by Richards

Volume 7, Number 2, 1998

281

(1978, and references therein), before the re-
cent paper on Ps. difficilis (Jeanne 1996).
Contrary to those taxa previously studied in
our serial papers, Ps. vespkeps shows no
clear-cut morphological caste dimorphism.
In addition, Ps. vespiceps evidenced the oc-
currence of intermediate females which are
characterized by the combination of well
developed ovaries and the absence of in-
semination (Richards & Richards 1951). Al-
though the bionomic importance of such fe-
males remains largely speculative, their sig-
nificance in the colonial sodo-economics is
apparently relevant because Naumann
(1970), Simoes (1977) and M. V. Baio (un-
publ.) have found out that intermediates in
Protopolybia acutiscutis (cited as P. pumila)
and P. exigim exigiia, frequently laid eggs
which were primarily eaten by the layer
herself. Based on the morphometric analysis
adopted in our serial work, this paper deals
with caste differences and related aspects in
Pseudopoh/bia vespicqis testacea Ducke.

MATERIAL AND METHODS

A mature colony (A) of Pseudopolyhia
vespiceps testacea, from which a small num-
ber of individuals escaped during collec-
tion, was taken on 20 January 1975 in Ri-
beirao Preto, Sao Paulo State, southeastern
Brazil. The 603 collected wasps, all of
which were females, were fixed in Die-
trich's solution and then kept in 70% eth-
anol until dissection. From this sample,
300 wasps were randomly chosen for ex-
amination. In order to detect caste differ-
ences the following 22 external body parts
were measured and the number of hamuli
was counted for each specimen under a
binocular microscope: (1) head width
(HW), (2) head length (HL), (3) and (4)
maximum and minimum interorbital dis-
tances (IDx, IDm), respectively, (5) gena
width (GW), (6) eye width (EW), (7) pron-
otal width (PW), (8), (9) length and width
of mesoscutum (MSL and MSW), respec-
tively, (10) mesoscutellar length (MTL),
(11) metanotal length (MNL), (12) meso-
somal height (MSH), (13) alitrunk length

(AL), (14) propodeum length (PL), (15)
length of gastral tergite I (T,L), (16), (17)
basal and apical heights of T, (T,BH,
T,AH), respectively, (18), (19) basal and
apical widths of gastral tergite II (T,BW,
T,AW), respectively, (20), (21) length and
height of T, (T,L, T^H), respectively, and
(22) partial length of the forewing (WL)
(See Fig. 2 in Shima et al. 1994). In addi-
tion, color patterns and other morpholog-
ical peculiarities were examined. The nu-
merical data were statistically analyzed in
relation to ovarian and spermathecal con-
ditions.

The statistical analyses including the ca-
nonical discriminant analysis (CDA: Rao
1973) and Mahalanobis's distances (An-
derson 1968) through CDA were per-
formed with the SAS Program Package at
the Computer Center of Kyoto University.
Afterwards, a second colony (B) was col-
lected on February 3, 1995 in Pedregulho,
Sao Paulo State. The illustrations (Fig. lA,
B, C, and D) pertain to this colony.

RESULTS

Nest architecture and colony composi-
tion.— Both nests (A,B) were found in
shady places at about 3 m high above-
ground and both were hung from termi-
nal small branches, of which ramifications
pierced the outer envelope at upper parts,
but not the combs. In shape (ellipsoidal),
size (17.0 cm high, 12 cm 0) and number
of combs (8 and 7, respectively) both nests
were very similar and so these features
may represent the size reached by the ma-
ture colonies of Ps. vespiceps (Fig. 1). The
multilayered greyish envelope covered
combs entirely. It had several, sometimes
indistinct, vertical rows formed by several
pulp additions during construction (Fig.
lA-D). The nest top, which incorporated
some twigs, was somewhat spongy, but
latterally the envelope was formed by two
to four layers of delicate and loose sheets
(Fig. IB). The nest entrance, which was
large and devoid of special structures (Fig.
IC), was located at the lowermost part of

282

Journal of Hymenoptera Research

Fig. 1. Nest of Pseudopol\^bia vespiceps: A — held in tlie position as in nature; B — with the outer envelope
removed partially; C — oriented as to exhibit an enlarged entrance and a part of the lower comb; D — seen
from the top; E — lowermost combs and pedicel. A-D, nest 2; E, nest 1.

Volume 7, Number 2, 1998

283

the nest. Each comb was hung below the
other by a single centrally located paper
pillar (1.0 cm long, 1.0 cm wide; Fig. IE).
The only exception was the uppermost
comb which was attached to the substrate,
a twig, by a row of 4 or 5 thin pillars. The
first comb was solely supported by these
pillars and, therefore, all combs hung
from the upper one independently of the
envelope. As recorded by Richards (1978)
the general plan of Pseudopolybia nests is
similar to Vespuln nonvegica of subfamily
Vespinae. As stated above, colony A used
in this study had 603 adult females (118
queens, 396 workers, 89 intermediates)
and no males. The queen ratio (total queen
number /total female number ) was 0.196.
Eight combs had 729 eggs, 584 larvae, 586
cocoons and 287 empty cells.

Ovarian development and insemination. —
In the analyzed sample four types of ova-
ry development were recognized (Fig. 2):
type A (short ovarioles): A„ thread-like
with no sign of development (n = 60), A.,
slightly developed (n = 74) and A, (n =
10), more developed than A^; type B (in
developmental stages similar to type A,
but with longer ovarioles); B„ B,, B, (n =
13, 33 and 7, respectively); type C (n = 44),
moderately developed with a few (1 to 3)
mature oocytes (mean 1.9, n = 85) and
type D (n = 59), well developed with an
average of 3.7 mature oocytes (range 1 to
7), n = 176. Since insemination was de-
tected exclusively in females with type D
ovaries, only these females are regarded
as queens. All the others females were un-
inseminated, and those with type C (de-
veloped) ovaries were regarded as inter-
mediates (sensu Richards and Richards
1951; Richards 1971). Conversely, females
with A and B ovary types constitute the
cohort of workers. Although measure-
ments were not made, mature oocytes in
the intermediates were clearly smaller
than those in queens.

Ovariole length, wing-wear patterns and
relative amount of fat tissue. — Mean ovariole
length (MOL) was longer in queens

(8.52±0.46 mm, n = 46), followed by B
typed workers (7.63±0.38 mm), interme-
diates (6.95±0.57 mm), and smallest in A
typed workers (6.69 ±0.39 mm). These dif-
ferences were statistically significant
(ANOVA, p < 0.01, F = 0.242). Another
interesting tendency was found connect-
ing ovary development and amount of fat
tissue. Indeed, individuals showing pro-
gressive kinds of ovary development pre-
sented also larger amounts of fat tissue.
The culmination of such tendency was
found in D typed females (queens) which
showed higher amounts of fat reserve
(grade IV, cf. captions in Fig. 3). Most un-
inseminated females (type A ovaries) had
shorter ovarioles relative to the B typed
ones, and both had little (I) or moderate
(II) amounts of fat tissue. But, some un-
inseminated callow females, which were
recognized by their pale and soft cuticule,
had larger amounts of fat tissues also
(grade IV). In this case, it seems that the
mere presence of larger amounts of fat tis-
sue is not conclusively connected to fur-
ther queenhood. Indeed, extensive dissec-
tions in representatives of taxa with clear-
cut caste differences suggest that callow
stages of both castes invariably present
conspicuous fat amounts (Noll unpubl.).

Figure 3 shows the relations between
ovary development and wing-wear in the
different kinds of Ps. vespiceps females.
Grades of wing-wear showed relationship
to the relative amount of fat tissues. Most
analyzed females, including all queens,
had fresh wings (pattern 0), while some
workers and intermediates showed consis-
tent wing-wear (patterns 1-3, Fig. 3).
Wing-wear may indicate relative age pro-
gression and flight activities. The fact that
all queens had fresh wings suggest their
absence in extranidal tasks.

Morphometries and hamulus number. — -
Differences of mean values of 24 charac-
ters measured or counted were tested in
workers, queens and intermediates by
Bonferroni-test (Table 1). Between workers
and queens, significant differences were

284

Journal of Hymenoptera Research

1mm

Fig. 2. Grades of ovarian development in Piiciidopolybia vespiceps. Type A: short ovariole type. A, -thread-like;
A, and Aj-showing initial development of the oocytes); Type B: long ovaries type. B.-thread-like as in A,; B2
and B3-comparable to A, and A^ respectively; Type C: moderately developed with a few mature oocytes; and
Type D: well developed and bearing many mature oocytes. Other explanations, in the text.

Volume 7, Number 2, 1998

285

WING PATTERN

CLASSES OF OVARIOLE LENGTHS ( mm j

WORKERS
6.69 t 0.39 7.63 t 0.38

INTERMEDIATES
6,95 i 0,57

QUE ENS
8,52 1 0,46

16 5

2

TYPES OF OVARY
DEVELOPMENT

Ad

Fig. 3. Relationships between wing patterns (0 = fresh; 1-3 = worn in ascending order) and relative amounts
of fat tissue (I = few; II = moderate; III = abundant; IV = much abundant). Number of individuals for each
class of fat tissue is expressed by the size of circle, in which numbers of individuals of different ovarian types
are shown.

detected only in T,BW, being larger in
workers, and NH, larger in queens (p <
0.05). Between queens and intermediates,
means of five characters, i.e., HW and HL,
Idx, TX, T,BW and NH differed (p <
0.05). Values of all but one character (NH)
were larger in the intermediates. Between
intermediates and workers, differences
were detected in four characters, HW, IDx,
MSW and T,H (p < 0.05), all of which
were larger in intermediates. No signifi-
cant difference was detected in AL, re-
garded as a reference length for compar-
ing sizes, among the three classes of fe-
males. This implies that size differences
are so small that virtually no size poly-
morphism exists in this species. But, it is
noteworthy that the intermediates are
larger in some characters than workers
and even queens.

The results of the canonical discrimi-
nant analysis (CD A) based on 23 charac-
ters also show the absence of clear dimor-
phism between inseminated egg-layers
(queens) and uninseminated non-egg-lay-
ers (workers) (Fig. 4). Queens and workers
were plotted against scales of the CAN,
and CAN; variables between -3.0 and 3.0
and —2.5 and 2.5, respectively. The distri-
bution pattern of the intermediates was
very peculiar because their plots showed
a vertical distribution at the right side in
the scatter diagram. Only one queen and
one worker presented higher CAN, values
than intermediates.

To calculate the CAN, and CAN, values
the following equations were used (see ta-
ble 1): CAN, = -0.47 (HW - 3.79) + 5.24
(HL - 3.04)- 0.36 (Idx - 3.17) - 5.13
(IDm - 1.81) + 0.53 (GW - 0.94) - 1.98

286

Journal of Hymenoptera Research

Table 1. Means and CAN, and CAN, values for 23 characters examined.

Means (mm) ± SD

CAN, ■

values

CAN,

values

Characters

Queens

Workers

Intermediates

Standardized

Ra\^

Standardized

Raw

HEAD

HW»»-

3.78 ±

0.07

3.82 ±

0.06

3.78

±

0.08

-0.04

-0.47

0.67

8.86

HL"

3.02 ±

0.09

3.07 ±

0.08

3.04

-t-

0.08

0.45

5.24

-0.41

-4.81

IDx*!

3.17 ±

0.06

3.20 ±

0.08

3.17

±

0.08

0.03

-0.36

-0.11

-1.51

IDm

1.81 ±

0.04

1.83 ±

0.05

1.81

±

0.05

-0.24

-5.13

-0.24

-5.19

GW

0.94 ±

0.06

0.95 ±

0.05

0.94

±

0.06

-0.03

0.53

0.09

1.53

EW

0.88 ±

0.06

0.88 ±

0.04

0.88

■+-

0.06

-0.11

-1.98

-0.12

-2.30

MESOSOMA

MSL

2.48 ±

0.06

2.49 ±

0.07

2.48

-+-

0.07

-0.01

-0.12

0.22

3.36

MSW

2.50 ±

0.06

2.53 ±

0.06

2.50

■+■

0.07

0.13

1.86

0.16

2.30

PW

3.20 ±

0.08

3.24 ±

0.07

3.21

-+■

0.09

-0.09

1.02

0.15

1.75

MTL

1.14 ±

0.04

1.15 ±

0.04

1.13

±

0.04

-0.10

2.35

0.25

6.10

MNL

0.66 ±

0.05

0.66 ±

0.05

0.65

■+■

0.05

-0.30

-5.88

0.29

5.67

MSH

3.55 ±

0.13

3.57 ±

0.13

3.53

■+-

0.14

-0.02

-0.14

0.19

1.41

AL

4.93 ±

0.12

4.95 ±

0.11

4.92

-

0.13

-0.18

-1.47

0.05

0.40

METASOMA

PL

1.44 ±

0.08

1.47 ±

0.08

1,44

±

0.08

0.34

4.14

0.07

0.82

T,L

2.10 ±

0.08

2.14 ±

0.08

2.12

±

0.10

0.05

0.49

-0.07

-0.80

T,BH

0.87 ±

0.09

0.86 ±

0.07

0.86

±

0.08

-0.34

-4.12

-0.13

-1.58

T,AH

1.53 ±

0.09

1.56 ±

0.10

1.55

±

0.10

0.17

1.71

-0.15

-1.53

T,L"

2.90 ±

0.13

2.97 ±

0.10

2.92

-t-

0.13

0.32

2.44

-0.23

-1.78

T,BW**'

2.33 ±

0.11

2.42 ±

0.13

2.38

+

0.14

0.56

4.21

-0.22

-1.62

TjAW

3.68 ±

0.12

3.70 ±

0.15

3.67

-*-

0.15

-0.31

-2.17

0.04

0.29

T,H'

3.19 ±

0.16

3.26 ±

0.14

3.14

±

0.16

0.34

2.11

0.74

4.54

WING

WL

5.09 ±

0.10

5.13 ±

0.10

5.10

-•-

0.13

0.02

0.20

-0.42

-3.43

NH***

9.59 ±

0.89

9.07 ±

0.73

9.26

-*-

0.84

-0.57

-0.57

0,04

0.04

Full names of characters are explained in the text (see Material and Methods),

*,**,' Mean significant differences between queens and workers, queens and intermediates and intermediates

and workers, respectively (Bonferroni t-test, p < 0,05).

(EW -
(MSW

(MTL - 1.14) -
(MSH - 3.54)
(PL - 1.45) +
(TIBH - 0.86)
(T,L - 2.92) 4
(T,AW- 3.68)
(WL - 5.10) ~

0.88) - 0.12 (MSL -

- 2.51) + 1.02 (PW -

5.88 (MNL -

- 1.47 (AL -
0.49 (TIL -

- 1.71 (TIAH
4.21 (T,BW

- 2.11 (T,H -

2.48) + 1.86
3.21) - 2.35
0.66) - 0.14
4.92) + 4.14
2.12) - 4.12
- 1.55) + 2.44
-2.37) - 2.17
3.17) + 0.20

0.67 (NH - 9.30)

CAN2 = 8.86 (HW - 3.79) - 4.81 (HL -
3.04) - 1.51 (IDx - 3.17) - 5.19 (IDm - 1.81)
+ 1.53 (GW - 0.94) - 2.30 (EW - 0.88) +
3.36 (MSL - 2.48) + 2.30 (MSW - 2.51) +
1.75 (PW - 3.21) + 6.10 (MTL - 1.14) + 5.67
(MNL - 0.66) + 1.41 (MSH - 3.54) + 0.40

(AL - 4.92) + 0.82 (PL - 1.45) - 0.80 (TIL
- 2.12) - 1.58 (TIBH - 0.86) - 1.53
(TIAH - 1.55) - 1.78 (T2L - 2.92) - 1.62
(T2BW - 2.37) + 0.29 (T2AW - 3.68) +
4.54 (T2H - 3.17) - 3.43 (WL - 5.10) +
0.04 (NH - 9.30)

The most important variable to deter-
mine CANl was MNL. Some other vari-
ables, such as HL, IDm, T,BW and T.BH
were also important. The most important
variable to determine CAN, was HW. Ma-
halanobis's distances (D-) between queens
and workers, queens and intermediates
and intermediates and workers were 0.99,
1.51 and 1.12, respectively, showing that, as

Volume 7, Number 2, 1998

287

3,0-

2 5-

2,0-

K5-

1.0-

0,5-

<

o

00-

-05-

-1.0-

-1.5-

-2,0-

-25-

-30-

A = QUEEN

O = VVORKER

D = INTERMEDIATE

-4

-1

CAN

1

Fig. 4. Discrimination among queens, workers and intermediates of Pseudopolybia vespiceps by the canonical
discriminant analysis based on 23 characters. Other explanations in the text.

expected from the result of basic statistics,
queens are morphologically more similar
to workers than to intermediates.

Aspects related to color patterns and exter-
nal morphology. — Differences that were
more conspicuous than morphometric
characters were found in head, mesosoma
and metasoma.

Head (Fig. 5): Workers heads are of various
shapes. Despite sequentially continuous,
these characteristics can be grouped into
three major types, i.e., flat, ovate and
pointed types (types I-III), and four color
patterns (A, to Aj). A tendency was de-
tected and it shows that the coloration of
the flat head type is lighter than in the

288

Journal of Hymenoptera Research

EZ

^ 1

1 mm

YETLLOW ■.

I I LIGHTER I I

LIGHT

DARK

chestnut:

L.IGHT

DARK

DARKER

DARK ■■

Fig. 5. Color patterns and shapes of head in queens, workers and intermediates of Pseudopolybia vespiceps.
Color: A, to A,; Shape: 1. flattened; II. ovate; III. pointed; IV. ovate but more enlarged apically.

Volume 7, Number 2, 1998

289

ovate and pointed head types. On the oth-
er hand, ovate head (type IV) character-
izes the queens in which two color pat-
terns (A3, A4) appear, and these are very
similar to the workers darkest forms.

Mesosoma (Fig. 6): Workers present six
mesosomal color forms ranging from light
(B,) to dark (B„). Mesoscutellum of each
type presents (or not, see type a, B,) a pair
of dark spots which vary in size and form
(separated circles, ellipsoids partly fused)
(e.g., types b-e, B,). Except for type B„
there are no double transverse bands on
the mesoscutellum. On the other hand,
queens present only one color form (B;.),
but as workers, they present various spot
types (a-e) on the mesoscutellum. In ad-
dition, a pair of arched transverse bands
occur near the basal and apical margins of
the mesoscutellum (apical bands are often
lighter in color and more vestigial than ba-
sal ones). Such bands are seen through the
transparent cuticule (Fig. 6, B^, b-d, B, a-
e). Despite color patterns overlap among
the diverse types of females, mesoscutellar
bands easily discriminated queens from
workers. In intermediates, the patterns of
the head and thorax were similar to work-
ers including a tendency for the flat type
of head (n = 30) and lighter colors (A,, n
= 9; A,, n = 7; Aj, n = 14). Indeed, only
one intermediate presented A4 II pattern
which is similar to queens. From 16 color
patterns of the worker's thorax, 12 were
similar in intermediates with higher fre-
quency of B, b (12); B, e (7), B, e (8). Con-
versely, only B, a,c; B, b and B4 e were not
observed in such females.

Metasoma (Fig. 7): Three major color
forms (G,-G,) occur, and each form has
three to five color variations on T2 (C,-
C,,). Queens and intermediates present
lighter gastral patterns (G, and C,-Cj),
among which C, is the most frequent (n =
28). Workers have darker gasters (G, and
G„ and C,, C5-C,,), among which C^ and
Ch are the most frequent (n = 80 and 77,
respectively). In intermediates, six forms
occur (G], C -C4, and G,, C^ with the high-

est frequency in C, (n = 18) and C, (n =
14). Although form C, is shared by all
kinds of females, queens generally present
lighter and enlarged gasters than workers.

DISCUSSION

Although caste differentiation in social
wasps is most conspicuous in the Vespi-
nae, especially Vespula (Blackith 1958;
Spradbery 1972), certain swarm-founding
polistine wasps have also evolved distinct
size dimorphism. Such a fact was clearly
stated long ago by von Ihering (1903), who
described remarkable morphological caste
differences in Agelaia vicina, but his results
have been overlooked by modern special-
ists in spite of its citation by Richards and
Richards (1951). Indeed, additional records
on quite conspicuous caste differentiation
in the Polistinae, analyzed either qualita-
tively and /or quantitatively by statistical
methods, appeared only recently. Genera
and species examined are: New World:
Agelaia flavipennis (Evans and West-Eber-
hard 1970); A. areata (Jeanne and Fagen
1974); Polybia emaciata (Hebling and Letizio
1973); Polybia dimidiata (Maule-Rodrigues
and Santos 1974; Shima et al. 1996a); Apoi-
ca flavissima (Shima et al. 1994); A. pallens
(Jeanne et al. 1995); Protonectarina sylveirae
(Shima et al. 1996b), Pseudopolybia dijficilis
(Jeanne 1996), and Epipona guerini (Hunt et
al. 1996), and Old World genera: Ropalidia
montana (Yamane et al. 1983); R. bambusae
and R. leopardi (Kojima and Kojima 1994);
Polybioides tabidus (Richards 1969; Turillazi
et al. 1994), etc. In the Polistinae, accord-
ing to Richards (1978) at least three pat-
terns of caste differentiation are found: 1)
Conspicuous size and allometric differ-
ences present, with queens larger than
workers in the absence of intermediates
(Agelaia spp: A. areata, Jeanne and Fagen
1974; A. pallipes and A. multipicta, Noll et
al. 1997; A. vicina, Sakagami et al. 1996;
Protonectarina sylveirae, Shima et al. 1996b;
Epipona guerini. Hunt et al. 1996); 2) Con-
spicuous dimorphism present, with
queens smaller than workers in most char-

290

Journal of Hymenoptera Research

(A

.2
E

9)

i

C
0)

3

0

bed

(TD (H3 (^

Fig. 6. Color patterns (B, to B^) and mesoscutellar spots (a-e) in queens, workers and intermediates of Pscii-
dopolybia vespiceps.

Volume 7, Number 2, 1998

291

Queen + Intermediates ( large frequency )

Worker + Intermediates ( only Gj C )

CHESTNUT

OWN/

BLACK

Fig. 7. Color patterns (G.-G,) in the gastral parts (Ci-C,,) of queens, workers and intermediates of Pseudo-
polybia vespiceps.

acters and intermediates absent (Apoica
flavissima, Shima et al. 1994; Polybia dimi-
diata, Shima et al. 1996a); 3) Morphological
differences slight or indistinct, and inter-
mediates present (Parachartergiis smithii,
Mateus et al. 1997; Protopolybia exigua, Noll
et al. 1996; and Pseudopolybia vespiceps,
present work). However, in some groups,
queens are significantly smaller than

workers in some characters and larger in
others (Apoica pallens, Jeanne et al. 1995;
Pseudopohfbia difficilis, Jeanne, 1996), and
according to Jeanne et al. (1995) this is
considered as non-size-based caste dimor-
phism probably due to a reprogramma-
tion in growth parameters (Wheeler,
1991).

Differently from Pseudopolybia difficilis

292

Journal of Hymenoptera Research

(Jeanne 1996), P. vespiceps presented slight
caste dimorphism between queens and
workers. Most analyzed characters
showed no differences between queens
and workers. Such slight distinction is
comparable to Parachartergus smithii (Ma-
teus et al. 1997) and Protopolybia exigua ex-
igua (Noll et al. 1996). Using log-log plots
of the most discriminant characters (Fig. 8)
and considering three distinct groups
(queens, workers and intermediates),
queens presented a type of allometric
growth (Fig. 8A) comparable to Epipona
guerini (Hunt et al. 1996) while workers
and intermediates presented non-allome-
tric growth. However, disregarding inter-
mediates (as done by Jeanne 1996 in Ps.
dijficilis) and considering only two groups,
i.e. individuals with or without ovary de-
velopment (Fig. SB), Pseudopoli/bia vespi-
ceps presented only non-allometric growth
and such pattern is quite similar to that
found in Ps. dijficilis (Jeanne 1996). Such a
fact is very important because origin and
role of intermediates relies primarily on
their careful detection according to taxa
and colony cycle.

In addition to diverse degrees of caste
differentiation in this subfamily, the oc-
currence of intermediates, brought about
complexity in the caste problems. Rich-
ards (1971) assumed that their role is the
production of either trophic eggs or males
but, Forsyth (1978), West-Eberhard (1978)
and Gastreich et al. (1993) considered
them as young uninseminated queens.
Richards (1971) and West-Eberhard (1978)
considered that, in a general way, inter-
mediates have ovary development related
to queen number. That is, in the presence
of a few queens they present larger ova-
ries and vice-versa. Intermediates are
present in species with low caste dimor-
phism (Richards, 1978; Noll et al., 1996;
Mateus et al., 1997; present paper). On the
other hand, species with pronounced caste
dimorphism as Agelaia vicina (Sakagami et
al. 1996), A. pallipes and A. multipicta (Noll
et al. 1997) and Apoica flavissima (Shima et

3.7 -

♦

♦

*

3.5 -

D

* D

□

Q

* ♦

A «

3.3 -

A

□

□ □

es * •

A

A

-o — o-

-a --IT --a---'

3.1 ■

□

a

□ □

ASA

D

A

D

Q

□ □

ES A

2.9 -

A

A
A

ES

A

A A
A A

A A

2.7 -

A

A

D

2.5 -

'

'

' 1

1.8 2

2.2

2.4

2.6 2.8

T2BW

♦

Intermediates

D Queens

worKers

Log tintermeaiales)

■■Log (Queens)

Log (WorkefS)

3 5 -
33 -

♦ ♦ ♦

♦ • ♦

• * * ♦ A ♦

A * • * • • «

-•..A.A.A.A-A. A...

A A » • ' ' ' ,

I 3 1 ■
2.9 ■
2 7 -
25 -

* * « * A * A
♦ A *** »»A

A A 4 A A A A
A A A A
A

A

♦

22 24

T2BW

2.6

2.8

Intermediates Queens

Workers
"Log (Intermediales Queens)
=Log (Workers)

B

Fig. 8. Discrimination between queens, intermedi-
ates and workers of Psctidopolyhia vcspiccpf using the
log-log plots of basal width of tergite II (T,BW) and
head length (HL). A — Queens, workers and inter-
mediates were separatedly plotted, B — Queens and
intermediates were put together representing ovary-
developed individuals and workers undeveloped-
ovary individuals.

al. 1994) present sterile workers. However,
uninseminated queens were found in A.
vidua (Sakagami et al. 1996, Baio et al. in
press.), Polyhia dimidiata (Maule-Rodrigues
& Santos 1974, Shima et al. 1996) and Ro-

Volume 7, Number 2, 1998

293

& Santos 1974, Shima et al. 1996) and Ro-
palidia margiiiata (Chandrashekara & Gad-
agkar 1991), and so their contribution to
male production seems likely. In a general
sense, intermediate females have been re-
corded in several other taxa, e.g., Poh/bia
chrysothorax, P. jurinei, Parachartergiis fra-
terniis and Angiopolybia spp. (Richards and
Richards 1951); Brachygastra scutellaris
(Carpenter and Ross 1984), B. lecheguana
(Machado et al. 1988). But, most of these
species must be reanalyzed with a stan-
dardized statistical method. Richards
(1971, 1978) mentioned that the interme-
diates characterize taxa in which structur-
al caste differences are absent. This is,
however, only partly valid, because P.
dimidiata (Shima et al. 1996a) albeit char-
acterized by clear cut caste dimorphism
with queens smaller than workers, have
intermediates also. In addition, Naumann
(1970) and Simoes (1977) and M. V. Baio
(unpubl.) discovered their occurrence in
two taxa, Pwtopolybia acutiscutis (cited as
P. pumila) and P. exigua, which bear quite
clear intercaste morphological differences.
The latter observations are very important
because in both cases the eggs laid by the
intermediates were invariably eaten by
nearby mates, which suggests likely im-
portance in colony socio-economics, and
remarkable similarities to the system of
worker's trophic eggs widespread in the
stingless bees (Zucchi 1993 and ref. there-
in). By using basic and multivariate statis-
tics, the present study revealed another
important facet about intermediates. It is
noteworthy that conspicuous morphomet-
ric differences between queens and work-
ers were not detected in Pseitdopolybia ves-
piceps whereas intermediates tended to be
larger (significantly larger in some char-
acters, such as HW and PW) than queens
and workers. Such trend was also sug-
gested by distribution of plots on CAN,
and CAN, axes (Fig. 4) and Mahalanobis
distances "(1.51 for Q/I vs. 0.99 for Q/W).
In contrast, intermediates were more sim-
ilar to queens in gaster color than to work-

ers (Fig. 5-7), while similar to workers in
head color and mesoscutellar spots of the
mesoscutellum.

Another interesting fact refers to head
form since most intermediates had flat-
tened head or queen-unlike head (Fig. 5),
as also suggested by the significant differ-
ences in HW. Intermediates were also dif-
ferent in ovaries which were shorter and
had eggs smaller than those in queens
(Figs. 2, 3). As relative age estimated by
wing-wear was apparently not related to
ovarian development (Fig. 3), it may sug-
gest that long ovaries did not develop
from shorter ones. Indeed, older females
(Fig. 3) presented both ovary lengths. So,
intermediates are apparently uninseminat-
ed specialized females that have invari-
ably shorter but developed ovaries. There
is a possibility that their eggs are trophic,
and invariably eaten as in the afore men-
tioned Pwtopolybia species. In addition,
they generally bear more abundant fat tis-
sue than typical workers, and in some in-
stances they may present as much fat tis-
sue as true queens (Fig. 3, IV).

Colonies with intermediate females
show curious combination of differences
and similarities among the three types of
females. These are probably related to the
degree of caste differentiation: (1) species
with intermediates more similar to work-
ers usualy have clear-cut caste dimor-
phism {Pwtopolybia exigua, Simoes 1977; P.
pumila (= acutiscutis), Richards 1978; Nau-
mann 1970); P. sedula, Agelaia lobipleura
melanogaster, Richards 1978 and Polybia
emaciata, Hebling and Letizio 1973); (2)
species with intermediates more similar to
queens have castes distinct in some char-
acteristics (Brachygastra bilineolata, Pseudo-
polybia compressa morph laticincta, Rich-
ards 1978) or present clear-cut size dimor-
phism (P. dimidiata: Shima et al. 1996a)
and (3) species with intermediates distinct
in some characters can bear slight or no
caste differences (Pseudopolybia z'espiceps,
present results, and Belonogaster junceus,
Richards 1969) in which intermediates dif-

294

Journal of Hymenoptera Research

fered significantly from both castes in
hamulus number (similar to P. vespiceps)
and wing length, suggesting that these fe-
males are not merely queens or even ova-
ry-developed workers (Richards, 1969).

In the studied case it is possible to con-
sider intermediates by two ways: 1 ) Inter-
mediates as young or uninseminated
queens as pointed out by Forsyth (1978),
West-Eberhard (1978) and Gastreich et al.
(1993). In this case the pattern (Fig. 8B) is
similar to that found in Ps. difficilis
(Jeanne, 1996). 2) Specialized workers hav-
ing a combination of queen and worker
characters. In this case, intermediates have
non-allometric growth but, queens pre-
sented allometric growth (Fig. 8A).

The present example shows how the
caste system in neotropical swarm-found-
ing polistine is complex and diverse.
Moreover, the occurrence of intermediates
in several polistine taxa turn the under-
standing of their social systems fascinat-
ing. The results at our hands suggest that
the intermediates represent a specialized
state. However, in the lack of substantial
data a final conclusion has to be post-
poned.

ACKNOWLEDGMENTS

This study was supported in part by FAPESP (Fun-
daf ao de Amparo a Pesquisa do Estado de Sao Paulo)
and CNPQ (Conselho Nacional de Desenvolvimento
Cientffico e Tecnologico). Special thanks are also due
to: to the late Prof. T. Inoue (Kyoto University) for
his invaluable help in statistics and informatics, and
Sidnei Mateus for photographs in Fig. 1. Mr. J.R. So-
mera prepared the drawings, and C.M.S. Rovai allied
to S.R. Bonatti prepared the preliminary version of
the manuscript.

LITERATURE CITED

Anderson, T. W. 1968. bitwiiucUcvi to iiuiltivnrintc stn-
tifticnl rt)ii;/i/s/s. John Willey & Sons. Inc., New
York, 374 pp.

Baio, M. v., F. B. Noll, R. Zucchi, and S. Simoes. In
press. Non-allometric caste differences in Agelaui
vidua (Hymenoptera, Vespidae, Epiponini). So-
ciobiology 34

Blackith, R.E. 1958. An analysis of polymorphism in
social wasps. /Msccfcs Socianx 5: 263-272.

Carpenter, J. M. and K. G. Ross. 1984. Colony com-

position in four species of Polistinae from Suri-
name, with a description of the larva of Bracliy-
j;fls(ra sciitellaris (Hymenoptera, Vespidae). Psi/-
Wif 91: 237-250.

Chandrashekara, K. and R. Gadagkar. 1991. Unmated
queen in the primitively eusocial wasp Ropalidia
marginata Lep. (Hymenoptera, Vespidae). Instxtes
Sociaux 38: 213-216.

Evans, H. E. and M. J. West-Eberhard. 1970. The
waspi. University Michigan Press, Ann. Arbor,
Michigan, 265 pp.

Forsyth, A. 1978. Studies on the behavioral ecology
of polygynous social wasps. PhD thesis. Harvard
University, Cambridge.

Gastreich, K. R., J. E. Strassmann and D. C. Queller.
1993. Determinants of high genetic relatedness in
the swarm-founding wasp, ProtcfKilylna exigua.
Ethclcgy, Ecolog}/ and Evolution 5: 529-539.

Hebling, N. J. and V. L. G. Letizio. 1973. Polimorfismo
de las castas femininas de Polybia emaciata. Bole-
tini dc la Sociedad Entomclcgica 7: 23-24.

Hunt, J. H., D. K. Schmidt, S.S. Mulkey and M. A.
Williams. 1996. Caste dimorphism in Epipona
guerini (Hymenoptera: Vespidae): Further evi-
dence for larval determination. Journal of the Kan-
sas Entomological Society 69: 362-369.

Ihering, R. von, 1903. Contributions a I'etude des ves-
pides de I'Amerique du Sud. Annalles de la Socicte
Entomologique de France 72: 144—155.

Jeanne, R. L. 1980. Evolution of social behavior in the
Vespidae. Annual Review of Entomology 25: 371-
395.

Jeanne, R. L. 1991. The swarm-founding Polistinae.
In: K. G. Ross and R. M. Matthews (eds.). The
social biology ofunisps: 191-231. Cornell University
Press, Ithaca.

Jeanne, R. L. 1996. Non-allometric queen-worker di-
morphism in Pseudopolybia difficilis (Hymenop-
tera: Vespidae). Journal of the Kansas Entomological
Society 69: 370-374.

Jeanne, R. L. and R. Fagen, 1974. Polymorphism in
Stelopolybia areata (Hymenoptera, Vespidae). Psy-
che SI : 155-166.

Jeanne, R. L., C. A. Graf and B. S. Yandell. 1995. Non-
size-based morphological castes in a social insect.
Naturwissenschaften 82: 296-298.

Kojima, J. and K. Kojima. 1994. Biometric comparison
of body size in queens and workers in New
Guinean swarm-founding Ropalidia. Tropics 4:
49-55.

Machado, V. L. L., S. Gravena and E. Giannotti. 1988.
Analise populacional e morfometrica em uma co-
lonia de Bracln/gastra leehegnana (Latreille, 1824),
na fase reprodutiva. Anais da Sociedade Entonu^-
logica do Brasil 2: 491-506.

Mateus, S. A., F. B. Noll and R. Zucchi. 1997. Mor-
phological caste differences in neotropical
swarm-founding polistinae wasps: Parachartergus

Volume 7, Number 2, 1998

295

smithii (Hymenoptera: Vespidae). lournal of the
New York Entomological Society 105: 129-139.

Maule-Rodrigues, V. and B. B. Santos. 1974. Vespi'-
deos sociais: Estudo de uma colonia de Poh/hia
dimiiiiata (Olivier, 1791) (Hymenoptera, Polisti-
nae). Rcvista Bnisilcira de Entoniologia 18: 37^2.

Naumann, M. G. 1970. The nesting behavior of Pro-
topoli/bicj pumila in Panama (Hymenoptera, Ves-
pidae). Ph.D. Thesis. Univ. of Kansas, Lawrence.
182 pp.

Noll, F. B., S. Mateus and R. Zucchi. 1996. Morpho-
logical caste differences in neotropical swarm-
founding polistinae wasps. \-Protopolybia exigua
exigiin (Hymenoptera: Vespidae). ]ournal of the
New York Entomological Society 104: 61-68.

Noll, F. B., D. Simoes and R. Zucchi. 1997. Morpho-
logical caste differences in neotropical swarm-
founding polistinae wasps. Agelnia m. muttipicta
and A. p. pallipes (Hymenoptera: Vespidae).
Ethology Ecology and Evolution 9: 361-372.

Rao, C. R. 1973. Linear statistical inference. John VVilley
c& Sons, New York.

Richards, O. W. 1969. The biology of some W. African
social wasps (Hymenoptera, Vespidae, Polisti-
nae). Mcmorins de la Societd Entomological Italiana
48: 79-93.

Richards, O. W. 1971. The biology of the social wasps
(Hymenoptera, Vespidae). Biological Reviezvs 46:
483-528.

Richards, O. W. 1978. The social wasps of the Americas
excluding the Vespimie. British Museum of Natural
History, London. 580 pp.

Richards, O. W. and M. J. Richards. 1951. Observa-
tions on the social wasps of South America (Hy-
menoptera, Vespidae). Transactions of the Royal
Entomological Society of London 102: 1-170.

Sakagami, S. F., R. Zucchi, So. Yamane, F. B. Noll and
J. M. F. Camargo. 1996. Morphological caste dif-
ferences in Agelaia vicina, the neotropical swarm-
founding polistine wasp with the largest colony
size among social wasps (Hymenoptera: Vespi-
dae). Sociobiolog)! 28: 1-17.

Shima, S. N., So. Yamane and R. Zucchi. 1994. Mor-
phological caste differences in some Neotropical

swarm-founding polistine wasps. 1. Apoica flav-
issinm (Hymenoptera, Vespidae). Japanese Journal
of Entomology 62: 811-822.

Shima, S. N., So. Yamane and R. Zucchi. 1996 a. Mor-
phological caste differences in some Neotropical
swarm — founding polistine wasps. II. Pol\/l'ia
dimidiata (Hymenoptera, Vespidae). Japanese jour-
nal of Entomology 64: 131-144.

Shima, S.N., So. Yamane and R. Zucchi. 1996 b. Mor-
phological caste differentiation in some Neotrop-
ical swarm — founding polistine wasps III. Proto-
ncctarina sylveirae (Hymenoptera, Vespidae). Bul-
letin of the Faculty of Education, Ibaraki University
42: 57-67.

Simoes, D. 1977. Etologia e diferencia^ao de castas em
algumas vespas sociais (Hymenoptera, Vespi-
dae). PhD thesis. Universidade de Sao Paulo, Ri-
beirao Preto. 182 pp.

Spradbery, J. P. 1972. A biometric study of seasonal
variation in worker wasps (Hymenoptera, Ves-
pidae). lournal of Entomolog}/ 47: 61-69.

Turillazzi, S., E. Francescato, A. Baldini Tosi and J. M.
Carpenter, 1994. A distinct caste difference in
Polybioides tahidus (Fabricius)(Hymenoptera: Ves-
pidae). Insectes Sociaux 41: 327-330.

West-Eberhard, M. J. 1978. Temporary queens in
Metapoiybia wasps: non-reproductive helpers
without altruism? Science 200: 441-^43.

Yamane, So., ]. Kojima and S. K. Yamane. 1983.
Queen/worker size dimorphism in an Oriental
polistine wasp, Ropalidia montana Carl (Hyme-
noptera, Vespidae). Insectes Sociaux 30: 416-422.

Wheeler, D. E. 1991. The developmental basis of
worker castes polymorphism in ants. AnuTican
Naturalist 138: 1218-1238.

Zucchi, R. 1993. Ritualized dominance, evolution of
queen-worker interactions and related aspects in
stingless bees (Hymenoptera: Apidae). In: T. In-
oue and So. Yamane (eds.). Evolution of Insect So-
cieties: 207-249. Hakuhinsha, Tokyo.

J. HYM. RES.
Vol, 7(2), 1998, pp. 296-304

Priscomasaris namibiensis Gess, a New Genus and Species of

Masarinae (Hymenoptera: Vespidae) from Namibia, Southern Africa,

with a Discussion of its Position Within the Subfamily

Friedrich W. Gess
Albany Museum, Grahamstown, 6140 South Africa

Abstract. — A new genus and species of Masarinae (Vespidae), Priscomasaris namibiensis Gess,
is described from Namibia. Cladistic analysis of the Masarinae shows that Priscomasaris belongs
within the tribe Masarini, in which it is the most plesiomorphic member, and that it represents a
new subtribe, here named Priscomasarina, sister to the subtribes Paragiina and Masarina com-
bined. The justification by Carpenter (1997) for recognising subtribes, that is to have a group name
for the Australian masarines, remains untouched. P. namibiensis visits flowers of Gisekia and Li-
meiim (Moluginaceae) and at pools lands on the water surface to drink.

The subject of this paper, an unde-
scribed species of Masarinae, was discov-
ered by the author and S.K.Gess while
they were engaged in field studies in Na-
mibia in March and April 1997. This wasp
was widespread and abundant together
with a species of Jiigurtia Saussure of sim-
ilar size and colouring. Its habit of alight-
ing on water immediately attracted atten-
tion and distinguished it from the ]ugurtia
which, like all members of that genus,
drinks at the water's edge.

Priscomasaris Gess, genus novum

Type species: Priscomasaris namibiensis
Gess, sp. n.

Recognition. — Priscomasaris namibiensis,
the only included species, is immediately
recognisable by its very short tongue
(glossa), the unique development of the
scutellar (= axillary) processes, and the
very shiny metasomal integument with a
unique pattern of large pale spots situated
posterolaterally on terga 1-IV and poster-
omedially on terga II-V.

Description. — Head wider than long in
frontal view. Clypeal dorsal margin
straight, ventral margin broadly truncate.
Eye with inner margin sinuous but

smooth (like that of Paragia Shuckard lack-
ing an interior emargination). Postocular
and preoccipital carinae fused. Tempora at
midheight slightly narrower than eye in
lateral view at same level. Antenna in both
sexes with ten flagellomeres and no club.
Scape (with radicle) narrow and short, in
male 2.3 X and in female 3x as long as
greatest width. Labrum width about half
interantennal distance, ventral margin
broadly rounded. Female mandible tri-
dentate, that of male quadridentate. Glos-
sa short, neither the section basal to the
bifurcation nor the glossal lobes elongate,
section basal to bifurcation slightly shorter
than glossal lobe; paraglossa extending
slightly beyond bifurcation. Length of ex-
tended tongue (measured from anterior
edge of labrum to end of glossal lobes)
equal to 0.07x body length. Acroglossal
buttons present. Prementum whole, hy-
postomal bridge level, glossal sac absent.
In both sexes maxillary palpus six-seg-
mented, labial palpus four-segmented.

Propleuron neither grooved nor de-
pressed, nor medially diverging dorso-
medially to form a fossa. Pronotum ven-
tro-laterally with an inconspicuous
groove, pretegular area non-carinate. Teg-
ula subtriangular, about as wide as long.

Volume 7, Number 2, 1998

297

Scutellum triangular, roundly pointed
posteriorly; antero-lateral angle conspicu-
ously produced backward into an out-
wardly convex and apically pointed pro-
cess, superficially reminiscent of the simi-
lar looking process (parategula) arising
from the postero-lateral mesoscutal angle
of some Eumeninae. Propodeum dorso-
laterally produced into short, posteriorly
directed spine.

Forewing not longitudinally folded
when at rest. Marginal cell truncate basal-
ly, slightly wider basally than apically, 2r-
rs straight basal to insertion of RS, with
proximally tubular but distally merely
pigmented appendix. Two submarginal
cells present. CuA2 and A meeting at a
near right angle, a stub of free apical sec-
tion of A present. First discal cell longer
than subbasal cell. Junction of RS and M
slightly thickened. Prestigma short, about
one quarter length of pterostigma. Cu-a
situated shortly distad of fork of M and
CuA.

Hindwing CuA diverging from
M + CuA basad of insertion of cu-a; cu-a
inserting on CuA and aligned with A; free
apical section of A absent. Jugal lobe pres-
ent but only about one quarter length of
subbasal cell. Pre-axillary excision evanes-
cent.

Male front trochanter without process;
female front tarsal setae straight; middle
coxa narrow; middle tibia with two spurs;
hind coxa non-carinate; claws of all legs
toothed.

Metasomal tergum I and sternum I sep-
arate. Metasomal segments after II non-re-
tractile. Male genitalia with sharply point-
ed parameral spines, volsella separate
from paramere.

Relationships. — The determination of the
phylogenetic position of Priscomasaris
would have been greatly facilitated had it
been possible to add its character states to
the data matrix used by Carpenter to gen-
erate his cladogram of the masarine gen-
era (Carpenter 1993: Fig. 7.4). This was not
possible to do as the characters and data

matrix used in generating the cladogram
have not been published.

As Priscomasaris has many plesiomorph-
ic characters, the present analysis confines
comparison to the basal taxa of Carpen-
ter's cladogram. [See also cladogram of
Australian Masarinae (Carpenter 1997:
Fig. 17).] These taxa are the Gayellini (only
characters common to Gayella Spinola and
Paramasaris Cameron are used), Paragia
Shuckard, the basal member of the Aus-
tralian Masarini (= subtribe Paragiina
(Carpenter 1997)) and Ceramius Latreille,
the basal representative of the remaining
Masarini (sensu Carpenter) (= subtribe
Masarina, the sister-group of the Paragiina
(Carpenter 1997)). The use of Paragia and
Ceramius is adequately justified as these
two genera exhibit all the groundplan con-
ditions for each of their respective sub-
tribes for all of the characters considered
in the analysis. Euparagia Cresson is in-
cluded as the outgroup.

The characters considered, mostly
drawn from Carpenter (1982, 1993 and
1997), some with modification, are:

Forewing

1. Marginal cell: not narrower basally
than apically (0); 2r-rs curving basal to
insertion of RS so that it is narrower (1).

The basally sinuously narrowed mar-
ginal cell is a synapomorphy of Paragiina
(Carpenter 1997).

2. Submarginal cell number: three (0); two

Two submarginal cells is a synapomor-
phy of Masarini (Carpenter 1982).

3. CuA2 and A: angled where meeting (0);
rounded together (1).

The apically smoothly rounded subdis-
cal cell is a synapomorphy of Paragiina
(Carpenter 1997).

4. First discal cell: shorter than subbasal
cell (0); as long or longer than subbasal
cell (1). The short discal cell in Gayellini
is considered a reversion from the state

298

Journal of Hymenoitera Research

of an elongate discal cell in other Ves-
pidae and is thus an autapomorphy of
Gayellini (Carpenter 1989).

Hindwing

5. CuA: diverging from M+CuA slightly
distad of the insertion of cu-a or at the
insertion of cu-a (0); distal to the inser-
tion of cu-a at a distance much greater
than the length of cu-a (1); basad to the
insertion of cu-a (2). Nonaddihve.

Divergence slightly distad of the inser-
tion of cu-a or at the insertion of cu-a, as
in Euparngia, is considered to be the ple-
siomorphic condition for Vespidae; diver-
gence distal to the insertion of cu-a as in
Gayellini appears to be a reversion to the
plesiomorphic condition for aculeates and
is an autapomorphy of Gayellini; and di-
vergence basad to the insertion of cu-a is
a synapomorphy of Masarini (Carpenter
1982).

6. Cu-a: transverse (0); inserted on CuA
and aligned with A (1).

In the primitive state for aculeates, cu-a
is transverse as in Euparagin and Gayellini.
Derived states are to have cu-a inserted on
CuA and either aligned with A (as in Ma-
sarini) or strongly angled with A (as for
example in Eumeninae) (Carpenter 1982).

7. Free apical section of A: present (0); ab-
sent (1).

Loss of the apical section of A is syna-
pomorphic for Masarini (Carpenter 1982).

Head

8. Clypeal dorsal margin: straight (0); bis-
inuate (1).

The dorsally bisinuate clypeus is an aut-
apomorphy of Gayellini (Carpenter 1982).

9. Ocular emargination: present (0); ab-
sent (1).

Though relatively uncommon in aculea-
tes, emarginate eyes are characteristic of
Vespidae and absence of emargination
within the Masarinae is interpreted as de-

rived (Carpenter 1997). Loss of eye emar-
gination, stated by Carpenter to be a fun-
damental autapomorphy of Paragia, is
shared with Priscomasaris.

10. Number of male antennal articles: 13
(0); 12 (1).

Primitively, aculeate male antennae are
composed of 13 articles. Reduction to 12
articles, as in Masarini, is apomorphic
(Carpenter 1982).

Mouthparts

11. Labrum: narrow (0); broad (1).

A narrow labrum represents the
ground-plan condition of Vespidae, while
a broad labrum, resembling the plesiom-
orphic aculeate condition, is derived (Car-
penter 1982).

12. Female mandibles: quadridentate (0);
tridentate (1); bidentate (2). Nonaddi-
tive.

13. Distal section of glossal lobes: without
processes (0); each with two rows of
processes but these not together form-
ing a tube (1); each with two rows of
flattened overlapping (imbricate) pro-
cesses curved such that their tips come
together forming a tube (2). Nonad-
ditive.

14. Unbranched basal section of glossa:
shorter than paraglossae (0); longer
than paraglossae (1).

Elongation of the unbranched basal sec-
tion of the glossa beyond the level of the
tips of the paraglossae is a synapomorphy
for Masarina (Carpenter 1997).

Mesosoma

15. Pretegular carina: present (0); absent

Polarity as in Carpenter (1997, character
17).

16. Propodeal spiracle: lateral (0); more or
less dorsal (1).

Polarity as in Carpenter (1997, character
24).

Volume 7, Number 2, 1998

299

Priscomasans

Paragia

Fig. 1. Cladogram showing the position of Prisco-
»i(7S(?ns relative to Gayellini, and to Paragia and Cer-
aniiui, the basal members respectively of Carpenter's
subtribes Paragiina and Masarina of the Masarini.

17. Male foretrochanter: without process
(0); with process (1).
Polarity as in Carpenter (1997, character
27).

The distribution of the 17 polarized
characters is given below:

Euparagia 00010 00000 02000 00

Gayellini 00001 00100 00001 00

Priscomasaris 01012 11011 11101 00

Paragia 11112 11011 11200 11

Ceramius 01012 11001 11210 01

A cladistic analysis using Hennig86
computer program (Ferris 1988) produces
one cladogram with a length of 22 steps,
consistency index of 0.90 and retention in-
dex of 0.81 (Fig. 1).

This analysis demonstrates that Prisco-
masaris belongs to the Masarini as it has
the characters identified by Carpenter
(1982, 1993) as supporting the monophyly
of this tribe: forewing with two submar-
ginal cells; hindwing with cu-a inserting
on CuA and aligned with A; CuA diverg-
ing from M + CuA basad of insertion of cu-
a; loss of free abscissa of A; labrum broad;
male antennae 12-segmented. It, however,
shows that Priscomasaris does not belong
within either Paragiina, as represented by
Paragia, or Masarina, as represented by
Ceramius, being more primitive than ei-
ther. The glossa is very short and exhibits
no elongation of either the glossal lobes or
the section basal to their bifurcation. These
two means of elongating the tongue have

been shown by Carpenter (1996: 390) to
define major lineages, of Paragia (and Me-
taparagia Meade-Waldo), and of Ceramius
(and the remaining Masarini). Whereas
the difference between Priscomasaris and
Paragia with regard to glossal length may
not be very obvious, differences in the dis-
tal section of the glossal lobes are very ap-
parent. In Priscomasaris each glossal lobe
has two rows of flattened processes form-
ing a sponge-like extension; in Paragia and
Ceramius the flattened processes are over-
lapping (imbricate) and are curved such
that their tips come together to form a
tube.

Priscomasaris shares with Carpenter's
subtribe Masarina a plesiomorphic fore-
wing venation: a basally truncate margin-
al cell with 2r-rs straight basal to insertion
of RS, and CuA2 and A angled where
meeting, clearly distinguishing it from the
genera of Carpenter's subtribe Paragiina
which show a derived, synapomorphic
condiHon (see Carpenter 1996: 393).

The absence of a pretegular carina, an
apomorphy according to Carpenter, dis-
tinguishes Priscomasaris from both Paragia
and Ceramius.

The unique scutellar (= axillary) pro-
cesses constitute an autapomorphy of Pris-
comasaris.

Clearly, if subtribes are to be main-
tained, the discovery of Priscomasaris ne-
cessitates the recognition of a third sub-
tribe, the sister-group of Paragiina and
Masarina combined. This subtribe can ap-
propriately be named Priscomasarina. The
justification by Carpenter (1997) for recog-
nising subtribes, that is to have a group
name for the Australian masarines, re-
mains untouched.

Eti/molog}/. — The name Priscomasaris is
derived from the Latin adjective priscus
meaning ancient, antique, belonging to
old times, and Masaris, the type genus of
the subfamily Masarinae. It is intended to
indicate its primitive nature.

300

Journal of Hymenoptera Research

Figs. 2-i. Prisconmsaris namibieitsis, female habitus.
2, dorsal; 3, dorso-lateral; and 4, lateral views (X 7.5).

Priscomasaris namibiensis
Gess, species nova

(Figs. 2-9)

Female. — (Figs. 2-8). Black. The follow-
ing are white: rarely small spot on each
side of frons near top of inner orbit; rarely
small spot dorsally behind eye; usually
short transverse marking medially on pro-
notum; usually small spot on humeral an-
gle; large oval spot on prepectus; rarely

small streak medially on mesoscutum be-
tween notauli; usually part or whole of
scutellar processes; propodeal spines and
area at their base; large suboval postero-
lateral spots on terga I-IV; postero-medial
spot (of varying size and present in only
ca. 40% of individuals) on tergum I: large
postero-medial spots on terga II-V. The
following are reddish: mandible (except
extreme base and apical teeth); clypeus
along ventral margin; labrum; labial and
maxillary palpi; most of pronotum (except
for black ventral margin and, if present,
white markings); variously developed dif-
fuse markings on mesoscutum — small
streak (if not yellow) medially between
notauli and two pairs of spots flanking the
notauli antero-laterally and postero-later-
ally; tegula; scutellar processes (if not yel-
low); scutellar disk; rarely diffuse area
next to yellow markings at base of pro-
podeal spines; most of terga I and II (other
than white spots); rarely diffuse transverse
posterior bands (between white spots) on
terga III-V; tergum VI; most of sternum II
(other than anterior transverse groove);
diffuse transverse posterior bands on ster-
na III-V; apex of sternum VI; diffuse areas
on all coxae and trochanters, distal half to
three-quarters of femora, and entire tibiae
and tarsi. Wings lightly infuscated, ante-
rior half of marginal cell somewhat
darker.

Pilosity pale; moderately dense, fine
and semidecumbent on clypeus and lower
half of frons; sparse, coarse and erect on
upper half of frons, vertex, dorsal surface
of pronotum, mesoscutum and scutellum;
sparse, fine and erect on metasoma (es-
pecially on terga I and II).

Punctures of central part of clypeal disk,
most of frons and all of vertex coarse, sep-
arated by shiny, unsculptured interstices;
widest interstices, subequal to puncture
diameter, on upper frons and vertex;
punctures at base and sides of clypeus and
between and around antennal sockets fin-
er and closer. Punctures on dorsal surface
of pronotum, mesoscutum, mesopleuron

Volume 7, Number 2, 1998

301

[6j

^^^^^^^^^■1

W^^^^^^^^^t^^t

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^^^PP^^^^^^^^^^.;T^^^^^^^^H

^^^^^^.^vJCMHjHHM^^^VBiPjgar^b. •' •. ^^^^^^^^^1

fflH^^^^^H

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Figs. 5-8. Priscoitmsurib luiiiiihiciisii. 5, Frontal view of head ot female ( ■
(X 25). 7, Glossa of female (x 150). 8, Front leg of female (x 25).

25). 6, Dorsal view of head of female

and scutellum similar to those on vertex
or coarser (particularly on mesoscutum),
separated by shiny unsculptured interstic-
es subequal to puncture diameter or wider
(on mesopleuron). Punctures of metasoma
smaller and more widely separated (par-
ticularly on tergum II), interspaces un-
sculptured, strikingly shiny.

Length 7.8-8.2 mm (average of 5: 7.9
mm); length of front wing 5.6-5.9 mm (av-
erage of 5: 5.7 mm); hamuli 9-12.

Head (Figs. 5 and 6) noticeably wider
(1.25X) than long in frontal view. Eye in
frontal view markedly convex; inner mar-
gin smoothly sinuous, non-emarginate.
Clypeus evenly convex, wider (1.4x) than

long; ventral margin broad, subtruncate,
minimally curved; junction of ventral and
lateral margins rounded. Antennal sockets
separated by 3X their diameter; interocu-
lar distance at level of sockets 2.25 X
length of scape (with radicle); length of
scape (with radicle) 3x greatest width,
equal to 1.5x of combined length of ped-
icel and flagellomere I; flagellomeres I-X,
respectively, with the following relative
lengths (and breadths) [length of flagello-
mere I = 1.0]: 1.0 (0.50), 0.64 (0.54), 0.50
(0.57), 0.50 (0.71), 0.50 (0.79), 0.50 (0.86),
0.54 (0.86), 0.50 (0.86), 0.54 (0.86), 0.86 (ta-
pering and end of segment narrowly
rounded). POL: OOL = 1:1.1. Vertex be-

302

Journal of Hymenoptera Research

hind ocelli shallowly depressed in front of
preoccipital carina. Glossa (Fig. 7).

Pronotum with carina running posteri-
orly from humeral angle and clearly sep-
arating dorso-lateral and ventro-lateral
faces; with shallow groove ventro-lateral-
ly; pretegular area with no carina or
groove, lobe slightly depressed. Mesoscu-
tum with median line in anterior half and
notauli deeply and widely depressed; no-
tauli subtransversely carinate, especially
posteriorly where concurrent; parapsidal
lines distinct. Tegula subtriangular, about
as wide as long. Mesopleuron anteriorly
divided by sinuous carina into anteriorly
and laterally facing surfaces; carinae
prominent antero-ventrally, joining each
other in front of coxae in wide, subtrans-
verse curve; episternal scrobe unusually
conspicuous. Scutellum triangular, poste-
riorly roundly pointed; antero-lateral an-
gle conspicuously backwardly produced
into outwardly convex and apically point-
ed process; pair of processes bracketing
the basal three-eighths of the scutellum;
basal quarter of scutellum a wide trans-
verse furrow crossed by about twelve lon-
gitudinal carinae; posterior three-quarters
of scutellum raised, markedly convex.
Metanotum fully exposed over entire
width, medially subvertical and slightly
overhung by scutellum, laterally widened
and excavated. Propodeum dorso-laterally
produced into short, posteriorly directed
spine.

Metasoma slightly constricted between
terga I and II, widest across middle of ter-
gum II and progressively narrowing pos-
teriorly to rounded tergum VI. Tergum I
2.4 X as wide as long; tergum II 1.6X as
long as tergum I and 1.7x as wide as long;
tergum I in posterior half with fine, im-
pressed median line; terga I and II with
fine but well defined longitudinal line
above their lateral margins. Sternum II
transversely grooved basally.

Front leg (Fig. 8); hind coxa non-cari-
nate; tibiae of middle and hind legs on the
exterior surface with scattered, small red-

1 mm

Fig. 9. Prifconittfan^ uainibteniis. Male genitalia, ven-
tral view on left, dorsal view on right.

dish peg-like setae (difficult to see
amongst the longer pale hairs); middle tib-
ia with two spurs; claws of all legs
toothed.

Male. — (Fig. 9). Black. The following are
yellowish-white: clypeal disk; large mark-
ing on mandible; small spot dorsally be-
hind eye; broad transverse band on ante-
rior aspect of pronotum; large suboval
spot on prepectus; small streak medially
on mesoscutum between notauli; rarely
greater part of tegula; entire scutellar pro-
cesses; in some individuals posterior half
or less of scutellar disk; in some indiduals
a small medial spot on metanotum; pro-
podeal spines and area at their base; large
suboval postero-lateral spots on terga I-IV
and usually also on V and postero-medial
spots on tergum 1 (usually), on terga Il-V
(always) and on terga VI and VII (usual-
ly); generally a complete or partially com-
plete dorsal streak on front tibia and rare-
ly diffuse dorsal markings on middle and
hind tibiae. The following are reddish:
mandible subapically; narrow streak
along ventral margin of clypeus; labial

Volume 7, Number 2, 1998

303

and maxillary palpi; side of pronotum;
tegula; anterior half to entire scutellar
disk; usually diffuse area next to yellow
markings at base of propodeal spines;
most of terga I and II (other than for yel-
lowish-white spots); usually diffuse trans-
verse posterior bands (between yellowish-
white spots) on terga III or IV- VI; most if
not all of tergum VII; most of sternum II
(other than for anterior transverse
groove); diffuse posterior bands on sterna
III-VII; entire sternum VIII; legs as in fe-
male.

Length 6.9-7.8 mm (average of 5: 7.3
mm); length of front wing 5.3-5.9 mm (av-
erage of 5: 5.6 mm); hamuli 10-11.

In general fades similar to the female,
the chief differences being as follow. Head
width relative to head length even greater
(1.33X). Mandible quadridentate. Anten-
nal sockets separated by 1.9x their diam-
eter; interocular distance at level of sock-
ets 1.85X length of scape (with radicle).
Antenna longer; scape (with radicle) 2.3 x
as long as greatest width and 1.4x as long
as combined length of pedicel and flagel-
lomere I; flagellomeres I-X, respectively,
with the following relative lengths (and
breadths) [length of flagellomere I = 1.0]:
1.0 (0.5), 0.69 (0.59), 0.66 (0.66), 0.63 (0.75),
0.63 (0.81), 0.56 (0.81), 0.56 (0.81), 0.56
(0.81), 0.56 (0.78), 0.86 (tapering and end
of segment narrowly rounded). Tergum
VII subtruncate with hind margin round-
ed laterally. Apical sternum with hind
margin truncate, deeply, narrowly emar-
ginate medially.

Front trochanter and metasomal sterna
without any processes.

Genitalia (Fig. 9).

Material examined. — Holotype: female,
Namibia: between Palm and Khorixas
(20.17S 14.05E), 31.iii.l997 (F. W. and S. K.
Gess) (on pink flowers of Gisekia africaiia
(Lour.) Kuntze, Moluginaceae) [Albany
Museum, Grahamstown]. Paratypes (216
females, 7 males): Namibia: between Palm
and Khorixas (20.17S 14.05E), 31.iii.l997
(F. W. and S. K. Gess), 3 females, 5 males

(2 males on pink flowers of Gisekia africana
(Lour.) Kuntze, Moluginaceae, 3 females
and 3 males on white flowers of Limeum
argute-carinatum Wawra & Peyr., Molugi-
naceae); 15.5 km W Khorixas (20.26S
14.54E), l.iv.l997 (F. W. and S. K. Gess), 8
females (on water); between Khorixas and
Uis (20.31S 14.56E), l.iv.l997 (F. W. and S.
K. Gess), 2 males (on small white flowers
of Limeum mi/osotis H. Walter, Molugina-
ceae); 43 km"s Mariental (24.58S 17.55E),
4.iv.l997 (F. W. and S. K. Gess), 154 fe-
males (on water and flying about above
water of pool); 97 km S Mariental (25.24S
17.54E), 4.iv.l997 (F. W. and S. K. Gess),
47 females (on water and flying about
above water of pool); 161 km S Mariental,
Tses (25.53S 18.07E), 4.iv.l997 (F. W. and
S. K. Gess), 3 females (on water); same lo-
cality, 17.iv.l998 (F. W. and S. K. Gess), 1
female (on water) [Albany Museum,
Grahamstown; Namibian National Insect
Collection, Windhoek; South African Mu-
seum, Cape Town; American Museum of
Natural History, New York; California
Academy of Sciences, San Francisco; and
Natural History Museum, London].

Distribution. — The species appears to be
widely distributed in Namibia, the present
records covering six degrees of latitude
(20-26S).

Behaviour. — Females visit pools of water
in drainage channels and river beds in or-
der to obtain water, presumably for use in
nest construction. When filling their crops
they alight on the water surface in the
manner of some species of Ceramius, Par-
agia, Metaparagia confluens (Snelling), and
M. nocatunga (Richards) (see Gess 1996:
67-76). When observed at pools, this be-
haviour, together with the habit of hold-
ing the wings erect whilst imbibing water,
immediately distinguishes P. namihiensis
from similarly sized and coloured species
of jugurtia Saussure, which alight on the
saturated soil at the water's edge and low-
er their wings. Where common, females
"swarm" in the air above water.

Both sexes forage on the small shallow

304

Journal of Hymenoptera Research

flowers of species of Gisekia and Limeum
(Moluginaceae), for which their short
tongues are adequate.

Nesting has not been observed.

Etyjnology. — The name namihiensis, an
adjective, is derived from Namibia and re-
fers to the provenance of the species.

ACKNOWLEDGMENTS

Thanks are expressed to the following for much ap-
preciated assistance as specified: Sarah Gess of the Al-
bany Museum, Grahamstown, co-collector of the ma-
terial, for valuable discussion and encouragement; Fer-
dy de Moor of the Albany Museum, Grahamstown
who ran the computer program; Robin Cross and Shir-
ley Pinchuck of the Electron Microscopy Unit, Rhodes
University, Grahamstown, for producing the scanning
electron micrographs; The Namibian Ministry of En-
vironment and Tourism for granting a permit to con-
duct research and collect biological specimens in that
country; Coleen Mannheimer of the National Botanical
Research Institute, National Herbarium of Namibia,
Windhoek, for idenhfying the plant specimens; Wo-
jciech Pulawski, Michael Prentice, Roy Snelling and
James Carpenter for their comments on earlier versions

of the manuscript; The South African Foundation for
Research Development for a running expenses grant
for field work during the course of which the present
material was collected.

LITERATURE CITED

Carpenter, J. M. 1982. The phylogenetic relationships
and natural classification of the Vespoidea (Hy-
menoptera). Systematic Entomology 7: 11-38.

Carpenter, J. M. 1989. The phylogenetic system of the
Gayellini (Hymenoptera: Vespidae; Masarinae).
Psyt'/it'95: 211-241.

Carpenter, J. M. 1993. Biogeographic patterns in the
Vespidae (Hymenoptera): Two views of Africa
and South America. In Goldblatt, P. ed., Biological
relationships between Africa and South America.
Yale University Press, New Haven and London,
pp. 139-155.

Carpenter, J. M. 1997. Generic classification of the
Australian pollen wasps (Hymenoptera: Vespi-
dae; Masarinae). Journal of the Kansas Entomolog-
ical Society 69: 384-400.

Farris, J. S. 1988. Hennig86, version 1.5. Port Jefferson
Station, New York.

Gess, S. K. 1996. The Pollen Wasps: Ecology and Natural
History of the Masarinae. Harvard University
Press, Cambridge, Massachusetts, 340 pp.

J. HYM. RES.
Vol. 7(2), 1998, p. 305

ANNOUNCEMENTS

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J. HYM. RES.
Vol. 7(2), 1998, p. 306

EDITOR'S NOTE

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change in my address, so please use the
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% National Museum of Natural History
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CONTENTS
(Continued from front cover)

SHIMA, S. N., F. B. NOLL, R. ZUCCHI, and S. YAMANE. Morphological caste differences
in the neotropical swarm-founding Polistine wasps IV: Pseudopolybia vespiceps, with
preliminary considerations on the role of intermediate females in the social orga-
nization of the Epiponini (Hymenoptera, Vespidae) 280

SIME, K. R., and D. B. WAHL. Taxonomy, mature larva, and observations on the biology
of Gnamptopelta obsidianator (Brulle) (Hymenoptera: Ichneumonidae, Ichneumoni-
nae) 157

WCISLO, W. T. Sexual dimorphism of wasp antennal structure in relation to parasitic and

non-parasitic behavior (Hymenoptera: Sphecidae) 178

ZITANI, N. M., S. R. SHAW, and D. H, JANZEN. Systematics of Costa Rican Meteorus

(Hymenoptera: Braconidae: Meteorinae) species lacking a dorsope 182

ANNOUNCEMENTS

Membership Search Service Now Available 305

Call for Bids for the 5th Conference of the International Society of Hymenopterists . . 305

EDITOR'S NOTE 306

5OCIETV

Journal of

Hymenoptera
Research

JUN 1 0 1999

Volume 8, Number 1 --^.^^BrariES,^ April 1999

ISSN #1070-9428

CONTENTS

AZEVEDO, C. O. A key to world species of Scolebythidae (Hymenoptera: Chrysidoidea),

with description of a new species of Dominihyihus from Brazil 1

BELOKOBYLSKIJ, S. A. and D. L. ]. QUICKE. A new genus and two new species of bra-

chypterous Lysiterminae (Braconidae) 120

COELHO, J. R. and K. WIEDMAN. Functional morphology of the hind tibial spurs of the

cicada killer (Sfhecius speciosus Drury) (Hymenoptera; Sphecidae) 6

KIMSEY, L. S. and M. S. WASBAUER. New genera and species of Brachycistidine wasps

from Southwestern North America (Hymenoptera: Tiphiidae: Brachycistidinae) . . 65

LaSALLE, J. A new species group and two new species of Euderomphale Girault (Hyme-
noptera: Eulophidae) from North America 116

MARSH, P. M. and G. A. R. MELO. Biology and systematics of New World Heterospilus
(Hymenoptera: Braconidae) attacking Pemphredoninae (Hymenoptera: Spheci-
dae) 13

MARSH, P. M. and S. R. SHAW. Revision of North American Aleiodes Wesmael (Part
5): The melanopterus (Erichson) species-group (Hymenoptera: Braconidae: Ro-
gadinae) 98

NEWMAN, T. M. and D. L. J. QUICKE. Ultrastructure of imaginal spermatozoa of sawflies

(Hymenoptera: Symphyta) 35

RASNITSYN, A. P., W. J. PULAWSKI, and X. MARTINEZ-DELCLOS. Cretaceous digger
wasps of the new genus Bestiola Pulawski and Rasnitsyn (Hymenoptera: Sphecidae:
Angarosphecinae) 23

ROMANl, R., N. ISIDORO, and F. BIN. Further evidence of male antennal glands in Aphel-

inidae: The case of Aphytis melinus DeBach (Hymenoptera: Aphelinidae) 109

(Continued on back cover)

INTERNATIONAL SOCIETY OF HYMENOPTERISTS

Organized 1982; Incorporated 1991

OFFICERS FOR 1999

Andrew D. Austin, President

John LaSalle, Presideiit-Elect

James B. Woolley, Secretary

John T. Huber, Treasurer

E. Eric Grissell, Editor

Subject Editors
Symphyta and Parasitica Aculeata

Biology: Mark Shaw Biology: Sydney Cameron

Systematics: Donald Quicke Systematics: Wojciech Pulawski

All correspondence concerning Society business should be mailed to the appropriate officer at the
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Membership. Members shall be persons who have demonstrated interest in the science of ento-
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Journal. The Journal of Hymenoptera Research is published twice a year by the International Society
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Statement of Ownership

Title of Publication: Journal of Hymenoptera Research.

Frequency of Issue: Twice a year.

Location of Office of Publication, Business Office of Publisher and Owner: International Society
of Hymenopterists, %, Department of Entomology, NHB 168, Smithsonian Institution, Wash-
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Editor: E. Eric Grissell, Systematic Entomology Laboratory, USDA, % National Museum of Nat-
ural History, Washington, DC 20560-0168.

Managing Editor and Known Bondholders or other Security Holders: none.

This issue was mailed 3 May 1999

J. HYM. RES.
Vol. 8(1), 1999, pp. 1-5

A Key to World Species of Scolebythidae

(Hymenoptera: Chrysidoidea), with Description of

a New Species of Dominihythiis from Brazil

Celso Oliveira Azevedo

Universidade Federal do Espi'rito Santo, Departamento de Biologia, Av. Marechal Campos 1468,

Marui'pe, 29040-090 Vitoria, ES, Brazil

Abstract. — Dominihythiis strictus sp. n. from Brazil, the first extant species of the genus, is
described and illustrated. Additional specimens of the rarely collected Ch/stopsenelta longivcutris
Kieffer 1911 are examined and the species variation analyzed. A world key to the fossil and extant
species of Scolebythidae is given.

Scolebythidae is a family of Aculeata
with five monotypic genera. Evans (1963)
proposed this family based on Clystopse-
itelln Kieffer 1911, from Brazil, a genus
transferred from the Bethylidae, and on
Scolehi/thus Evans 1963, from Madagascar.
Nagy (1975) added the third genus, Yca-
ploca, from South Africa and Australia de-
scribing the first male of Scolebythidae.
Day (1977) described and illustrated the
male genitalia of Clystopseiielln. Evans et
al. (1979) reanalyzed Scolebythus with de-
scription of the male and sting apparatus.

All extant species are exclusive austral,
but Prentice et al. (1996) described two fos-
sil genera, Libanohythtis and Dominibytlnis,
from Lebanese and Dominican amber.

Biological data on the species suggest
that the species of Scolebythidae are pos-
sibly gregarious ectoparasitoids of wood-
boring beetle larvae (Evans 1963, Nagy
1975, Day 1977, Evans et al. 1979), but
Gauld (1995) pointed out that all published
information is questionable.

In this paper, Domiiiibi/thus strictus sp. n.
from Brazil is described and illustrated.

and new taxonomic data are provided for
Ch/stopseuella longiventris Kieffer 1911. A
world key to the fossil and extant species
of this family is given.

The examined material used in the pres-
ent paper was provided by Canadian Na-
tional Collection of Insects, Canada (CNCl,
J. T. Huber); Universidade Federal do Pa-
rana, Brazil (DZPR, K. Zanol); and Instituto
Brasileiro de Geografia e Estatistica, Brasi-
lia, Brazil (IBCE, B. Dias).

Abbreviations for the main measure-
ments used in this study are as follow: LH,
length of head; WH, width of head; WF,
width of frons; HE, height of eye; OOL,
ocello-ocular line; WOT, width of the ocel-
lar triangle, including the ocelli; DAO, di-
ameter of anterior ocellus; VOL, vertex-oc-
ular line; LEW, length of forewing.

The nomenclature of the integument
follows Eady (1968) for the term coria-
ceous and Harris (1979) for the other tex-
tures. Terminology generally follows
Evans (1963), and the terminology of wing
cells and veins follows Gauld and Bolton
(1988).

KEY TO FOSSIL AND EXTANT SPECIES OF WORLD SCOLEBYTHIDAE

1. Mesoscutum with notaulus absent or at least incomplete; prostemum large, its width at
least 2.5 x length of propleuron; forewing with three closed cells, Rs vein shorter than
stigma (Domitiibythiis) 2

2 Journal of Hymenoptera Research

- Mesoscutum with complete notaulus; prosternum smaller than above, its width at most 2
X length of propleuron; forewing with five or six closed cells, Rs vein much longer than
stigma 3

2. Frons with an arched prominence; eye forming the widest part of head; notaulus present
anteriorly; occipital carina present dorsally; width of prosternum 2.5 x length of propleu-
ron; Cu vein nebulous Dominibythus inopinatus Prentice and Poinar

- Frons without prominence; gena forming the widest part of head; notaulus absent; occipital
carina absent dorsally; width of prosternum 2.8 X length of propleuron; Cu vein as a short
stub Dominibythus strictns Azevedo, new species

3. Pronotal disc enlarged, 1.57 x longer than mesoscutum; parapsidal furrows absent; fore-
wing without metacarpus, with five closed cells, marginal cell opened; tibial spur formula
1,1,1 Lybanobythus milkii Prentice & Poinar

- Pronotal disc shorter than mesoscutum, about 0.6-0.7 x longer than mesoscutum; parap-
sidal furrows present; forewing with metacarpus, with six closed cells, marginal cell closed;
tibial spur formula 1,2,2 4

4. Frons with a median prominence between antermal sockets; malar space virtually nonex-
istent; forewing with submarginal cell longer than marginal, extending beyond the basal
half of marginal cell Ycaploca evansi Nagy

- Frons without median prominence between antennal sockets; malar space short; forewing
with submarginal cell slightly shorter than marginal, not extending beyond the basal half

of marginal cell 5

5. Occipital carina absent; malar space well over half as long as basal width of mandible;
apex of marginal cell arched away from anterior margin of forewing; posterior area of
metasomal sternite V with two groups of appressed and dense setae

Clystopsenella longiventris Kieffer

- Occipital carina present; malar space short, less than half as long as basal width of man-
dible; apex of marginal cell on anterior margin of forewing; posterior area of metasomal
sternite V without special groups of setae Scolebythtts madecassus Evans

Dominibythus strictns Azevedo, apical teeth (Fig. 3), and with a shallow

new species suture below upper tooth. Clypeus ex-

(Figs. 1-7) tremely short, median lobe angulate, with-
out median carina. Anterma short, nearly

Description of female holotype: length reaching the pronotum; first four antennal

of body 3.93 mm; LFW 2.25 mm. segments in a ratio of about 7:4:1:2, fla-

Color: body castaneous, head and me- geHomere IX 1.2 X as long as thick, sen-
sosoma slightly darker; vertex with two g^ji^g circular. Head slightly narrower be-
lighter streaks starting on the crest and ex- j^^ Antennal sockets separated by less
tending between the ocellar triangle and ^y^^^ t^gj^ o^vn diameters. Frons weakly
eye; malar space, antenna, midtibia and coriaceous, with a very small number of
tarsi slightly lighter than head; wings hy- gmall and shallow punctures; frontal ca-
aline, veins castaneous. rina Iqw and short. Malar space broad, 1.0

Pubescence: very sparse and long over- x the basal width of mandible. LH 1.27 X

all; setae short and dense on antenna, WH; WF 0.5 x WH; WF 0.85 X HE; ocellar

some erect; setae noticeably longer on triangle very compact, OOL 1.45 x WOT

hindtibia; nearly absent at dorsum of gas- and situated on an imaginary line between

tral tergites, and concentrated at posterior eye tops, its frontal angle obtuse; posterior

half in gastral sternites. ocelli distant from vertex crest by 2.94 x

Head (Figs. 1-2): mandible wholly di- DAO. Eye subtriangular, with rounded

rected downward slightly, with two sharp corners; gena forming widest part of head.

Volume 8, Number 1, 1999

^:^

M+Cu

Figs. 1-7. Dominiln/thiis sfr;Vf»s. 1. Bodv, lateral. 2. Body, dorsal. 3. Mandible, frontal. 4. Mesosoma, ventral.
5. Forewing. 6. Foreleg, lateral. 7. Hindleg, lateral. Scale bars = 0.5 mm.

Journal of Hymenoptera Research

Vertex rounded, VOL 0.73 X HE; sides of
head straight and subparallel and sHghtly
convergent below. Occipital carina weak,
present only ventrally. Palpal formula 6:4,
segment I-III of labial palpi and segments
I-IV of maxillary palpi flattened.

Mesosoma (Figs. 1-2, 4): thorax weakly
coriaceous. Pronotal disc very short, 0.4 X
length of mesoscutum; anterior margin of
pronotum evenly convex in dorsal view.
Mesoscutum without notauli; parapsidal
furrows well-impressed, absent on the an-
terior half of mesoscutum, paralleled by
an outer carina. Scutellum long, only
slightly shorter than mesoscutum, prescu-
tellar sulcus narrow posterad laterally,
crossing entire anterior margin. Metano-
tum very narrow medially. Propodeal disc
0.56 X as long as wide; anterior margin of
propodeum with a transverse carina, pos-
terior half of median propodeal line with
a shallow groove; spiracle laterad, decliv-
ity without carinae. Propleuron very large,
produced strongly forw^ard. Prosternum
very long, its width 2.8 X the length of
propleuron, and 3 X longer than forefe-
mur. Mesopleuron with a scrobal pit
above midheight. Mesosterna separated
by a wide longitudinal groove. Metaster-
num narrow, separating the base of mid-
coxa. Legs without spines. Forewing (Fig.
5) with only costal, basal and first discal
cells, stigma wide, 0.63 X as long as wide;
Rs vein very short, 0.37 X the length of
stigma; Icu-a vein nearly vertical; Rs-(-M
vein reaching Sc-I-R vein far from stigma;
Cu vein as a very short stub. Hindwing
with anal lobe distinct. Forefemur 3 X as
long as thick (Fig. 6). Hindtibia longer
than hindfemur. Hindcoxae closely set.
Basitarsus of hindleg very long, longer
than half length of hindtibia and longer
than remaining tarsomeres together (Fig.
7). Tibial spur formula 1:1:1. Claws simple
and arched, dilated basally.

Metasoma (Figs. 1-2): coriaceous, wider
than both mesosoma and head. Tergite VII
with anterior margin deeply concavous.
Gonostylus 0.28 x the length of sting.

Sting and lancets without teeth; bulbous
little dilated; sting 0.63 mm long.

Material examined: 1 female holotype
BRAZIL, Distrito Federal, Brasilia, Ronca-
dor Ecological Station, shrub savanna;
12.ix.l979-21.x.l982; window trap; J. Dal-
maceo col. (IBGE). Paratypes, 17 females
BRAZIL, 16 females Distrito Federal, same
data as holotype except places savanna
grassland, shrub savanna, and swamp,
and data 12.ix.l979-21.x.l982 (IBGE); 1 fe-
male Rio de Janeiro, Rio de Janeiro, Cor-
covado; i.l960; Seabra & Alvarenga col.
(DZPR).

Variation: body wholly dark casta-
neous, clypeus distinctly lighter than
head; specimen from Rio de Janeiro with
clypeus projecting with a distinctly wider
median lobe; WH 1.17-1.20 X LH; WF
0.45-0.55 X WH; WF 0.85-0.96 X HE;
OOL 1.2-1.44 X WOT; posterior ocelli dis-
tant from the vertex crest 2.78-3.52 X
DAO; VOL 0.68-0.8 x HE; parapsidal fur-
rows not paralleled by outer carina.

Remarks: Dominihythus was first de-
scribed from a fossil of Late Eocene to Late
Oligocene Dominican amber (Prentice et
al. 1990). Dominihythus strictiis sp. n. is the
first living species of the genus. It may be
identified as Domiiiibytlms by the 3 closed
cells of the forewing. Dominibytlnis strictus
differs from D. inopiuatiis by the absence
of a frontal prominence, absence of occip-
ital carina dorsally and notauli, and pres-
ence of a tubular apical abscissa of the Cu
vein, although extremely short, as a stub,
while Cu vein in D. inopinatus is entire
nebulous. The color pattern of the malar
space and the lighter streaks on the vertex
of D. strictus resembles that of Clystopse-
nella longiventris.

Etymology: The name refers to the short
Rs vein of the forewing.

Clystopsenella longiventris Kieffer

This species is first recorded for Bahia
and Minas Gerais. The specimens are about
8 mm long and the light color streaks on
the vertex are very weak or nearly absent;

Volume 8, Number 1, 1999

mandible and malar space distinctly lighter
than head, legs lighter than mesosoma;
WH 1.06-1.10 X LH; WF 0.61-0.65 x WH;
WF 1.10-1.29 X HE; OOL 0.85-0.86 X
WOT; posterior ocelli distant from the ver-
tex crest 4.0-5.4 x DAO; VOL 0.54-0.61 x
HE; pronotal disc about 0.51 X the meso-
scutum length; notauli and parapsidal fur-
rows complete or nearly so; propodeal disc
about 0.5 X as long as wide.

New material examined: 8 females
BRAZIL, 1 female, Bahia, Encruzilhada;
xi.l974; M. Alvarenga col. (CNCI); 4 fe-
males Minas Gerais, Pedra Azul; xi.l974;
M. Alvarenga col. (CNCI); 2 females Dis-
trito Federal, Brasilia, Roncador Ecological
Station, shrub savanna; 23.1.1982 and
7.iv.l983; window trap; J. Dalmaceo col.
(IBGE); 1 female Sao Paulo, Mogi-gua^u,
Campiminas Farm; 3.i.l970; J. M. & G. R.
Campbell col. (CNCI).

Distribution: Brazil (Bahia, Minas Ger-
ais, Distrito Federal, Sao Paulo, Mato
Grosso de Sul, Santa Catarina). Gauld
(1995) commented that there are a few un-
described species of Chjstopsenella in mu-
seum collections from Neotropics and
Australia, and one or, possibly two species
in Costa Rica.

ACKNOWLEDGMENTS

I wish to thank E. R. Bortolini (UFES) and C. R. F.
Brandao (MZSP) for the loan of the camara lucida.

and to curators cited in the text for the loan of the
material studied here.

LITERATURE CITED

Day, M. C. 1977. A new genus of Plumariidae from
southern Africa, with notes on Scolebythidae
(Hymenoptera, Chrysidoidea). Cimbcbasia, Series
A, 4:171-177.

F.ady, R. D. 1968. Some Illustrations of Microsculp-
ture in the Hymenoptera. Proceedings of the Royal
Entomological Society of London (A) 43(3-6): 66-72.

Evans, H. E. 1963. A new family of wasps. Psyche 70:
7-16.

Evans, H. E., C. Kugler, and W. L. Brown Jr. 1979.
Rediscovery of Scolebythus madecassus, with a
description of the male and of the female sting
apparatus (Hymenoptera, Scolebvthidae). Psyche
86:45-51.

Gauld, 1. D. 1995. Scolebythidae. p. 468-470. In: Han-
son, P. E. and 1. D. Gauld (eds). Hymenoptera of
Costa Rica. Oxford, Oxford University Press. 859

PP-

Gauld, I. D. and B. Bolton. 1988. The Hymenoptera.
Oxford, Oxford University Press. 332 pp.

Harris, R. A. 1979. A glossary of surface sculpturing.
Occasional Papers m Entomology, Department of
Food and Agriculture, California 28:1-31.

Nagy, C. G. 1975. A new genus of Scolebythidae (Hy-
menoptera) from South Africa and Australia.
Journal of t lie Entomological Society of South Africa
38(l):75-78.

Prentice, M. A., G. O. Poinar Jr., and R. Milki. 1996.
Fossil scolebythids (Hymenoptera, Scolebythi-
dae) from Lebanese and Dominican amber. Pro-
ceedings of the Entomological Societ\f of Washington
98(4):802-8n.

]. HYM. RES.
Vol. 8(1), 1999, pp. 6-12

Functional Morphology of the Hind Tibial Spurs of the Cicada Killer
{Sphecins speciosus Drury) (Hymenoptera: Sphecidae)

Joseph R. Coelho' and Kimberly Wiedman

Department of Biological Sciences, Western Illinois University, 1 University Circle, Macomb,
Illinois 61455, USA, (E-mail: JR-Coelho@wiu.edu)

Abstrnct. — The functional morphology of the hind tibial spurs in female cicada killers is exam-
ined in relation to digging. The spurs are controlled by a passive mechanical system using the
corium; when the tarsus is flexed, the spurs are extended. The spurs resist extension, but not
flexion. Videography demonstrated that the hind legs are used in burrow excavation to clear soil
from the path of the wasp and that the position of the legs makes use of the spurs' resistance to
extension. Spur function is not consistent with use during prey carriage, which should cause the
spurs to lie flat. Enlarged spurs may have evolved from smaller structures used in grooming, and
may reduce the energetic and temporal costs of burrowing.

The bodies of winged insects function in
both aerial and terrestrial locomotion,
which may present conflicting selection
pressures on morphology. For example,
flight performance may be improved by
increasing the ratio of flight muscle mass
to body mass (Marden 1987) at the ex-
pense of legs or associated muscles, but
terrestrial locomotion would be adversely
affected. Many insects also dig or burrow,
which increases selection pressure for en-
hancement of the structures associated
with the terrestrial mode. Although fos-
sorial adaptation is relatively well under-
stood in vertebrates (Hildebrand 1985), lit-
tle of the function of specific adaptations
of insects to fossorial life has been dem-
onstrated. The Hymenoptera are generally
excellent flyers, but many also dig bur-
rows for nests. Therefore, they can be used
to show how animals well adapted to
flight handle the requirements of burrow-
ing.

Various morphological modifications
for fossorial life, such as foretarsal rakes.

' To whom all correspondence stiould be ad-
dressed.

have been described in wasps (Bohart and
Menke 1976), but their function has not
been ascertained. An exception is provid-
ed by Gorb (1996), who examined pretar-
sal anatomy and function in a variety of
insects. Fossorial species, such as the sphe-
cid Benibix rostraia L., have a large ungui-
tractor with well-developed microtrichia
for the production of strong frictional forc-
es with the substrate.

Cicada killers {Sphecius speciosus Drury)
are the largest (by mass) North American
sphecid (Horn 1976), and they are superb
flyers. Relative to other Hymenoptera,
they have a high ratio of flight muscle to
body mass, resulting in a high degree of
maneuverability. Females weigh nearly
one gram in body mass, which allows
them to carry cicadas {Tibiceu spp.) heavi-
er than themselves (Coelho 1997). Meta-
bolic rate during hovering is high (Joos
and Casey 1992), and body temperature
during flight is elevated and nearly con-
stant (J.R. Coelho, pers. obs.).

Female cicada killers also dig extensive
burrows. A cicada killer may move up to
1000 times her body mass in dry soil while
excavating her burrow (J.R. Coelho and

Volume 8, Number 1, 1999

A.J. Ross, pers. obs.). The opening of the
burrow is elevated in comparison to the
rest of the burrow, and a large tumulus
accumulates outside the entrance. The en-
trance tends to run at a 35-45° angle
downward and then becomes more level
(Evans 1966). The burrow branches into an
average of 15.8 cells (Dambach and Good
1943), which are used to store paralyzed
cicadas. The female lays one egg per cell.
On average, cells may vary in diameter
from 2 to 3 cm. Burrows vary in length
from 30 cm to over 100 cm (Evans 1966).
After provisioning the cell, the female ci-
cada killer places a wall of dirt across the
entrance. A variety of burrow architec-
tures, as determined by excavation, are
depicted by Riley (1892) and Evans (1966).
After the main burrow is dug, the dirt
from new cells is apparently used to plug
old ones, although soil may be added to
the tumulus as new cells are excavated.

The digging method uses a variety of
body parts and behaviors. The cicada kill-
er uses her mouthparts to loosen compact
soil to begin excavating. She uses her fore
legs to rapidly rake the dirt under and be-
hind her body. Finally, she uses her hind
legs to push the soil behind her as she
backs out of the burrow (Frost 1942; Dam-
bach and Good 1943; Evans and West
Eberhard 1970). The latter behavior results
in the formation of a prominent trough
through the tumulus leading away from
the burrow entrance (Dambach and Good
1943; Evans 1966).

Cicada killer females have a pair of en-
larged spurs on the distal end of the hind
tibia (see Evans [1966] for line drawings).
Males have similar spurs which, by com-
parison, are much reduced (Dambach and
Good 1943). While the presence of such
spurs in bees and wasps has been a useful
taxonomic tool (Eickwort and Fischer
1963), their functional significance has
rarely been addressed (except by Cane
[1979]). Some suggest that the spurs of fe-
male S. •^peciosiif are used to support the
cicada during prey carriage (Howes 1919;

Evans 1962), but our preliminary obser-
vations suggest that they are used to move
soil during burrow excavation.

Hildebrand (1985) lists five require-
ments of a fossorial animal to loosen and
move resistant material: a digging tool,
the capacity to produce and transmit large
forces, a transport mechanism for soil,
passive resistance to various loads, and
the ability to sustain activity. The cicada
killer hind leg functions as part of the
transport mechanism for soil, and we hy-
pothesize that the spurs are morphological
adaptations for that mechanism.

In this study we investigate the func-
tional morphology of the hind tibial spurs
in female cicada killers. Their mechanism
of action is demonstrated, and their poten-
tial function relative to digging and prey
carriage is examined.

MATERIALS AND METHODS

Origin of specimens. — Live cicada killers
were obtained from local nesting aggre-
gations in McDonough County, Illinois;
and the legs of freshly killed individuals
were used in all experiments. Unless oth-
erwise mentioned, all data are reported as
mean ± SEM (N).

Dissection. — We first investigated the
mechanism for movement of the spurs.
Under a dissecting microscope, a section
of the exoskeleton was removed from the
tibia near the tibiotarsal joint. Muscle tis-
sue was examined for any connection to
the spurs or to the corium, the unsclero-
tized conjunctival membrane which forms
the distal end of the tibia and from which
the spurs arise.

Mechanical linkage. — Five hind legs were
obtained. When the tarsus of each was
manually flexed, the spurs extended. Sim-
ilarly, when the tarsus was extended, the
spurs became flexed. The corium was sev-
ered by making an incision perpendicular
to a line between the spurs and tarsus us-
ing a 30-ga syringe needle. The tarsus was
flexed and extended again and the results
were recorded.

Journal of Hymenoptera Research

sensor arm

/femur/ spurs

corium

steel bar

Fig. 1. Apparatus used to measure the force pro-
duced by the spurs. The spurs were extended by low-
ering the steel bar, to which the tibia was glued. The
stationary force transducer measured the force exert-
ed by the spurs during extension. To measure the
force exerted during flexion, a similar set-up (not
shown) was used, except that the sensor arm was
placed above the spurs, and the steel bar was raised.

Having found that tarsal flexion causes
spur extension, we measured the force
generated by the spurs when they were
extended in this manner. The tibia of an
intact hind leg was glued to a steel bar.
The tarsus was manually extended at dif-
ferent angles to the tibia, causing the spurs
to extend. The spurs were oriented just be-
low and perpendicular to the sensor arm
of an optical force transducer (Narcosys-
tems type A), which was calibrated with
weights of known mass and connected to
a physiograph (Narcosystems MK-111).
The force exerted by the spurs was mea-
sured at tibia-to-tarsus angles of 180, 135,
and 90° and recorded by the physiograph.

Force-displacement curves. — To determine
the direction in which the spurs were ca-
pable of resisting force, the hind leg of a
female wasp was removed and the tibia
was glued to a steel bar with the spurs
facing upward. To measure the effect of
extension, spurs were placed on top of
and perpendicular to the sensor arm of a
force transducer connected to a physio-
graph (Fig. 1). An adjustable ringstand
was calibrated so that fractions of a rota-

tion of the knurled adjustment ring could
be used to move the bar known vertical
distances. The angle between the spurs
and the tibia was gradually increased by
moving the bar downward in 0.07-mm in-
crements. Measurements were taken until
the ends of the spurs slipped off the force
transducer. At this point the force was
maximal, but the spurs had not yet bro-
ken. In this way, the resistant force exerted
by the spurs when they were extended to
various degrees was recorded by the phy-
siograph.

To measure the effects of flexion, the
spurs were extended manually, then
placed just below and perpendicular to
the axis of the force transducer. The bar
was moved upward in 0.07-mm incre-
ments until the spurs laid flat against the
tarsus. Hence, the force exerted by the
spurs in resistance to varying degrees of
flexion was recorded by the physiograph.

Videograpin/. — Cicada killers were re-
corded with a Sony VHS-C videotape re-
corder while digging. Burrow entrances
were plugged with a small amount of soil
to induce digging in most cases, but spon-
taneous digging was also recorded. The
tapes were replayed using slow motion to
examine and describe aspects of digging
behavior in detail.

RESULTS

Dissections. — Of the 10 hind legs dis-
sected, no muscle or tendon attachment to
the spurs or corium was found. The tarsus
was traversed by a single tendon arising
from the pretarsal muscles in the femur
and tibia, an arrangement essentially iden-
tical to that described for honey bees
(Snodgrass, 1956).

Mechanical linkage. — When the tarsus
was manipulated and the corium was in-
tact, tarsal flexion caused spur extension,
and tarsal extension caused spur flexion in
every case. The spurs always moved in
one plane and resisted lateral movements.
After the corium was severed, the spurs

Volume 8, Number 1, 1999

1 15

Displacement (mm)

Fig. 2. Representative force-displacement curves for
extension and flexion of the female cicada killer hind-
tibial spurs.

no longer moved when the tarsus was ma-
nipulated.

When the tarsus was held at increasing
angles relative to the tibia, the force pro-
duced by the extension of the spurs in-
creased. When the tarsus was held at 180°
to the tibia, no measurable force was de-
tected. When the tarsus was held at 135°,
an average force of 0.315 ± 0.041(7) mN
was detected. Finally, when the angle was
decreased to 90°, an average force of 0.615
± 0.055(7) mN was produced.

Force-displacement curves. — Fig. 2 dem-
onstrates that as the spurs are displaced,
the force resisting extension increases lin-
early at first, then levels off near the max-
imum, while the force resisting flexion re-
mains constant at zero until the spurs lie
flat and touch the tarsus. Extension pro-
duced a maximum force of 21.68 ± 2.13(8)
mN, which is eight times greater than the
maximum force resulting from flexion,
2.64 ± 1.04(8) mN.

Video records. — Review of videotaped
digging behavior in slow motion demon-
strated how the female cicada killer uses
the hind legs to move soil. A wasp broke
off bits of soil from the burrow wall using
the mandibles and threw them posteriorly
under the body using rapid motions of the
fore legs. She then backed out of the bur-
row, alternately thrusting her hind legs in
arcs that began in a posterior direction
and ended in a lateral direction, removing
dirt from her path. Both front and rear

pairs of legs operated simultaneously,
with fore leg raking being repeated during
the recovery stroke of each hind leg. The
tibiotarsal angle was near 90° at the begin-
ning of the thrusting motion when the leg
was cocked, then the angle increased as
the leg was extended.

DISCUSSION

In dissections of the cicada killer hind
tibia, no muscles or tendons were found
leading to the spurs or corium. The spurs
were extended with increasing force as the
tarsus was flexed at increasing angles,
suggesting that the spurs are not under di-
rect muscular control, but that their de-
gree of extension depends instead upon
the angle of the tibiotarsal joint. When the
corium was severed, the spurs no longer
moved when the tarsus was manipulated.
The corium, a flexible, unsclerotized con-
junctival membrane, connects the tarsus
and spurs. When the tarsus is flexed, it
pulls on the lateral margin of the corium,
which, in turn, pulls on the medial bases
of the spurs, causing them to extend.
When the corium is severed, this mechan-
ical linkage is broken. Hence, the spurs are
controlled by a passive mechanical system
using the corium, as suggested by Cane
(1979).

The spur extension system generates a
very small force, less than 1 mN, which is
sufficient to extend the spurs, but likely to
be of little use otherwise. Hence, resistant
forces determine what tasks can be accom-
plished by the spurs. The spurs produced
a substantial resistant force to being ex-
tended outward, but not to being flexed
inward (whereupon they simply fold flat
against the tarsus). Only behaviors which
flex the tibiotarsal joint, using the spurs'
resistance to extension, are likely to effec-
tively use the action of the spurs.

Hymenopteran hind tibial spurs are
commonly used in grooming the contra-
lateral hind leg (Farish 1972, Cane 1979),
and we observed this behavior in captive
cicada killers. The tibiotarsal joint is flexed

10

Journal of Hymenoptera Research

and brought under the body to the contra-
lateral leg, where vertical motions up and
down the second leg, held between the
two spurs, occur (J.R. Coelho, pers. obs.).
Known as L,-L, grooming, this behavior
occurs in all 15 superfamilies of bees and
wasps tested by Parish (1972). Our data
suggest that the downstroke is the most
useful in Lj-L, grooming, as it is the power
stroke, where the spurs' resistance to ex-
tension would be used. The upstroke is
simply a return stroke. Although the spurs
may be used to groom the ipsilateral wing
(Parish 1972), we did not observe this be-
havior in cicada killers.

The utility of hind tibial spurs for Li-L,
grooming suggests that the spurs evolved
from smaller setae, but it does not ade-
quately explain their exaggerated size in
female Sphechis. It seems unlikely that the
grooming requirements of the female ci-
cada killer would be so much greater than
that of a male, or of other Hymenoptera,
that such elaborate structures would
evolve.

Howes (1919) suggests that the spurs of
female S. speciosus are used to support the
cicada during flight as "she squeezes [the
spurs] against the cicada's sides and thus
secures her burden during the overland
journey to the burrow." When the spurs
were removed from one wasp, the next ci-
cada was carried in a more vertical posi-
tion (Howes 1919). While Howes' (1919)
single anecdotal observation weakly sup-
ports his suggestion that the spurs have a
role in prey carriage, we do not believe the
spurs can function in the manner suggest-
ed. The hind legs wrap around the cicada
at high tibiotarsal angles (J.R. Coelho pers.
obs.); therefore, the spurs would not be ex-
tended. If squeezed against the sides of
the cicada, the spurs will lie flat against
the tarsus, as they have almost no capacity
to resist flexion.

In reviewing videotaped bouts of dig-
ging, we noted that although cicada killers
are capable of buzzing to produce a pow-
erful vibration (Coelho 1998), and many

Hymenoptera use vibration to loosen soil
(Spangler 1973), cicada killers did not
buzz while digging. The compact soil was
chewed from the walls of the burrow us-
ing the mandibles, and the fore legs were
used in rapid motions to rake the loose
particles below and behind the wasp.
Hence, cicada killers are "rakers" in that
they use the fore legs as rakes to move soil
(Evans and West Eberhard 1970). Spines
on the tarsus of the fore leg of S. speciosus
form a pecten (rake), which is believed to
augment the efficiency of raking (Evans
1966, Evans and West Eberhard 1970). Al-
though raking moves the soil beyond the
posterior margin of the animal, it does not
necessarily clear it from the burrow or its
entrance. As a unique type of "pusher, "
the female cicada killer uses the hind legs,
as opposed to the abdomen, to move soil
out of the burrow and beyond (Evans and
West Eberhard 1970). She backs out of the
burrow, thrusting each hind leg first pos-
teriorly then laterally, removing soil from
her path. The tibiotarsal angle appears to
be low (near 90°) at the beginning of the
motion when the leg is cocked, which ex-
tended the spurs. The path of the hind leg
during the power stroke caused the spurs
to be pushed against the soil in the direc-
tion of extension.

Hence, the high resistive force of the
spurs is used to move additional dirt, thus
increasing the efficacy of digging. Since
little force is required for flexion, the spurs
would not hinder the recovery stroke to
complete the motion. This effect is similar
to that of the "oars" of aquatic insects as
they swim through the water (Gullan and
Cranston 1994). When the spurs are fully
extended, the effective surface area of the
hind leg is substantially increased. Nu-
merous setae on the tibia and tarsus prob-
ably also contribute to the effective surface
area, which presumably allows the female
to move more soil per stroke than she
could otherwise, increasing the energetic
efficiency of digging.

The resulting energy savings could be

Volume 8, Number 1, 1999

11

used to dig more extensive burrows, to
forage for additional cicadas, or for other
fitness-enhancing activities. However, per-
haps time is more limiting than energy.
With an average lifespan of 13 days (Has-
tings 1989), they have little time to waste
on burrow construction. Observations
suggest that they carry out much of their
digging at night (Dambach and Good
1943), which would avoid temporal con-
flicts with their strictly diurnal foraging
bouts. Preliminary data from our labora-
tory (J.R. Coelho and A.J. Ross, pers. obs.)
suggest that digging rates of cicada killers
are consistent with completing a burrow
in one night, as suggested by Dambach
and Good (1943).

Hymenoptera tend to have long, thin,
and even delicate legs, while a transport
mechanism for soil should be large, broad,
firm, and thick (Hildebrand 1985) as it is
in mole crickets {Gryllotalpa), cicada
nymphs (Magicicnda) and certain scarab
beetles (Canthon) (Gullan and Cranston,
1994). Evans (1966) states that the cicada
killers' use of hind legs to move soil is
"unusual among digger wasps," and Pate
(1936) mentions the spurs as taxonomical-
ly important structural features of the ge-
nus Spheciiis. The large hind tibial spurs of
cicada killers appear to have been en-
hanced to function in digging while re-
taining their original function of groom-
ing. Enlarged spurs appear to be an im-
perfect solution to the problem of fossorial
adaptation in comparison to the highly
derived morphology of fully fossorial in-
sects. Nonetheless, cicada killers clearly
are accomplished burrowers in addition to
being excellent flyers. The spurs are light-
weight and collapsible, which may make
them a suitable compromise between the
demands of different locomotory habits.

ACKNOWLEDGMENTS

This study was supported by a Western Illinois
University Research Council Grant to ]RC and by
minigrants from the Honors Councils of the Illinois
Region and Western Illinois University to KW. We

thank Allan J. Ross for field and laboratory assistance,
and Jacqueline A. Runestad and two anonymous re-
viewers for comments on the manuscript.

LITERATURE CITED

Bohart, R.M. and A.S. Menke. 1976. Sphcciii waspjs of
the world: a generic reinsion. University of Califor-
nia Press, Berkeley, 695 pp.

Cane, J.H. 1979. The hind tibiotarsal and tibial spur
articulations in bees (Hymenoptera: Apoidea).
Journal of the Kansas Entomological Society 52:123-
137.

Coelho, J.R. 1997. Sexual size dimorphism and flight
behavior in cicada killers (Sphecius speciosus). Oi-
kos 79:371-375.

Coelho, J.R, 1998. An acoustical and physiological
analysis of buzzing in cicada killer wasps (Splie-
cius speciosus). Journal of Comparative Physiology A
183:745-751.

Dambach, C.A. and E. Good. 1943. Life history and
habits of the cicada killer in Ohio. The Ohio jour-
nal of Science 43:32-41.

Eickwort, G.C. and R.L. Fischer. 1963. A morpholog-
ical study of the tibial spurs of bees of the sub-
family Halictinae (Hymenoptera: Halictidae) of
eastern North America. Annals of the Entomologi-
cal Society of America 56:348-354.

Evans, H.E. 1962. The evolution of prey-carrying
mechanisms in wasps. Evolution 16:468-483.

Evans, H.E. 1966. The comparative ethology and evolu-
tion of the sand wasps. Harvard University Press,
Cambridge. 526 pp.

Evans, H.E. and M.J. West Eberhard 1970. The wasps.
University of Michigan Press, Ann Arbor. 265

PP-

Parish, D.J. 1972. The evolutionary implications of
qualitative variation in the grooming behaviour
of the Hymenoptera (Insecta). Animal Behaviour
20:662-676.

Gorb, S.V. 1996. Design of insect unguitractor appa-
ratus. Journal of Morphology 230:219-220.

Hastings, J. 1989. Protandry in western cicada killer
wasps, (Sphecius grandis, Hymenoptera: Spheci-
dae): an empirical study of emergence time and
mating opportunity-. Behavioral Ecologx/ and Socio-
hiology 25:255-260.

Hildebrand, M. 1985. Digging of quadrupeds, pp. 89-
109 in Functional vertebrate morphology. Hilde-
brand, M., D.M. Bramble, K.F. Liem, and D.B.
Wake (eds.). Harvard University Press, Cam-
bridge. 430 pp.

Horn, D.J. 1976. Biology of insects. W.B. Saunders
Company, Philadelphia. 439 pp.

Howes, P.G. 1919. Insect behavior. Gorhani Press, Bos-
ton. 176 pp.

Joos, B. and T.M. Casey 1992. Flight energetics of the
cicada killer wasp. American Zoologist 32:54A [ab-
stract].

12 Journal of Hymenoptera Research

Neukirch, A. 1982. Dependence of the life span of the cidae: Gorytini). Bulletin of tlic Bwokhin Eiitoiiio-

honevbee {Apif iiwUifica) upon flight performance logical Societi/ 31:198-200.

and energy consumption. Journal of Comparative Riley, C.V. 1892. The larger digger wasp. Insect life 4:

Physiology B 146:35-10. 248-252.

Pate, V.S.L. 1936. Studies in the nyssonine wasps. II. Snodgrass, R.E. 1956. Anatomy of the Iwney bee. Com-

The subgenera of Spliecius (Hymenoptera: Sphe- stock Publishing Associates, Ithaca. 334 pp.

]. HYM. RES.
Vol. 8(1), 1999, pp. 13-22

Biology and Systematics of New World Heterospilus

(Hymenoptera: Braconidae) Attacking Pemphredoninae

(Hymenoptera: Sphecidae)

Paul M. Marsh and Gabriel A. R. Melo

(PMM) Cooperating Scientist, USD A Systematic Entomology Laboratory, c/o National Museum

of Natural History, Washington, DC 20560 (correspondence address: P. O. Box 384, North

Newton, Kansas 67117); (GARM) Snow Entomological Museum, University of Kansas,

Lawrence, Kansas 66045 (present address: Departamento de Biologia, FFCLRP-USP, Av.

Bandeirantes 3900, 14040-901, Ribeirao Preto, S.P., Brazil)

Abstract. — Four new species of the braconid genus Heterospilus are described from Brazil and
Costa Rica and a key to the five species known to attack crabronid wasps of the genera Micros-
tigmus and Spihvnenn is presented. Biological notes are given for the hosts and the parasitoids and
comments on their relationships are also offered.

The genus Heterospilus Haliday is one of
the largest of the family Braconidae with
an estimated 500 or more species in the
New World, most of which are unde-
scribed. The genus belongs to the subfam-
ily Doryctinae characterized by a circular
opening present between the clypeus and
mandibles (oral cavity) and a row of stout
spines along the anterior edge of the fore
tibia. It can be distinguished from most
other braconid genera by the reduction or
absence of fore wing vein 2RS and can be
identified by the key to genera in Marsh
(1997).

We have very little information about
the biology of species in the genus Heter-
ospilus, which are, as far as known, idio-
biont ectoparasitoids (Shaw and Huddles-
ton 1991). Most records suggest parasitism
of wood boring Coleoptera, especially
Scolytidae, but a few species have been
reared from stem boring Lepidoptera and
one species from stem boring Symphyta.
However, in an unusual host association,
one species has been described from nests
of the crabronid wasp genus Microstigmus
Ducke (Richards 1935; Matthews 196'8).

Recently, one of us (GARM) reared sev-
eral species of Heterospilus from nests of
species of Microstigmus and Spilomeua
Shuckard that were being studied in Bra-
zil and Costa Rica. The purpose of this pa-
per is to provide descriptions of four new
species of Heterospilus and present a brief
discussion of the biologies of the hosts and
parasitoids.

BIOLOGICAL NOTES

The genus Microstigmus constitutes a
distinctive group of crabronid wasps, in
particular because of the social behavior
and the elaborate suspended nests found
in several of its species (Matthews 1968;
Richards 1972; West-Eberhard 1977; Mat-
thews 1991; Melo in press). Microstigmus
together with Arpnctophilus Smith, Spilo-
meua and Xi/sma Pate form the subtribe
Spilomenina of the Pemphredonini (Men-
ke 1989). Melo (1994) presented evidence
that Microstigmus is closely related to a
group of species within Spilomena contain-
ing S. nlini Antropov, S. kimseyi Antropov
and other related undescribed species. The
use also of S. atini as host bv Heterospilus,

14

Journal of Hymenoptera Research

reported here for the first time, therefore
should not be considered unexpected. Het-
erospilus has not been found parasitizing
other species of Neotropical Spilomena
whose nests have been studied (Melo un-
publ.).

Very little is known about the biology
of Heterospilus attacking Microstignnis
wasps. The only available information is
provided by Matthews (1968, 1991), Rich-
ards (1972) and Melo and Campos (1993).
Matthews (1968, 1991) reported on some
aspects of the biology of H. microstigmi
Richards parasitizing M. comes Krombein,
and Melo and Campos (1993) reported M.
myersi Turner being parasitized by a then
undescribed species of Heterospilus, here
named H. mattheivsi, new species.

Parasitoid females have never been
found inside the host nests and apparently
always lay eggs directly from the outside.
The positive correlation between the
length of the parasitoid ovipositor and the
thickness of the host nest wall found
among the different species of Heterospilus
(see below) provides additional evidence
for this behavior. Matthews (1968) ob-
served females of H. microstigmi crawling
over nests of M. comes and inserting their
ovipositors repeatedly into the nest. He
was not able to ascertain if these insertions
represented only probing or repeated ovi-
positions. However, considering that only
one egg or small parasitoid larva has been
found on each of the attacked host im-
matures and that in most Microstigmus
nests no more than one immature in the
appropriate stage is likely to be found,
these insertions may be only for probing
or for stinging and paralysis of the host
larva.

Only host pre-pupae seem to be at-
tacked, since eggs and small larvae of Het-
erospilus have been found only on imma-
tures at this stage. The pre-pupal stage in
Microstigmus is relatively short because
these wasps are not known to enter dia-
pause, although data on the duration of
this stage is lacking. The egg is laid di-

rectly on the surface of the integument of
the host larva. Parasitized larvae seem to
have their development interrupted which
as probably caused by pre-oviposition
stinging. In their Table 1, Melo and Mat-
thews (1997) gave a record of a small par-
asitic larva, attributed by them to Heter-
ospilus, on a male pupa of Microstig77ius fla-
vus Melo and Matthews (this record was
erroneously printed as associated with
nest 308 instead of nest 303). Since no
adult Heterospilus has been reared from
nests of this species of Microstigmus, it is
possible that this larva represents another
species of parasitoid (the only record of an
unidentified Chalcididae attacking Micros-
tigmus was obtained from this species),
and not the result of exceptional behavior
by a female Heterospilus.

The length of the female ovipositor is
clearly correlated with the thickness of the
host nest wall or the distance of the brood
cells from the nest outer surface. The lon-
gest ovipositors are found in H. richardsi,
new species, a species attacking Spilomena
alini and an undescribed species of Mi-
crostigmus. In both host species, the brood
cells are usually situated deep inside the
nests (see account under H. richardsi). The
second longest ovipositors are found in fe-
males of H. matthewsi, new species, a spe-
cies associated with M. myersi. The thick-
ness of the nest walls in M. myersi is quite
variable (Melo and Campos 1993). This
variation seems to be related to the nest
age, with young nests having thinner
walls and older nests tending to have an
extra covering layer of dirt particles. The
females of the other Heterospilus species
have shorter ovipositors and attack Mi-
crostigmus species whose nest walls are
relatively thin.

Matthews (1991) found a strong corre-
lation between the number of brood cells
in nests of M. comes and frequency of par-
asitism by H. microstigmi; almost all para-
sitized nests had eight or more cells. He
explained this pattern also in terms of the
relation between the length of the parasit-

Volume 8, Number 1, 1999

15

oid ovipositor and the thickness of the
host nest walls. Cells in nests with few
cells tend to be centrally placed and to
have thick walls, and therefore are out of
reach for the short ovipositor of females of
H. microstigmi. New cells are sequentially
added toward the periphery of the nest
and as their number increases, the outer
wall of the nest becomes thinner, making
the cells more accessible to the parasitoid's
ovipositor. Except for silk secreted by fe-
males, addition of new construction ma-
terial to the nest as it grows, a behavior
observed for example in M. meyersi (Melo
and Campos 1993), does not occur in M.
comes (Matthews and Starr 1984), the new
cells being added by stretching the nest
walls and holding the newly created pock-
ets (future cells) with silk.

One would expect females of Heterospi-
lus to have an ovipositor long enough to
cope with this kind of variation in wall
thickness of the host's nests, since a long
ovipositor could reach deep cells as well
as those closer to the nest surface. How-
ever, considering the situation found in H.
niicwstigmi this does not seem to be the
case. Apparently these parasitic wasps are
under strong selective pressure to have
the length of their ovipositor matching
very closely the average depth of the ma-
jority of their host's cells. As more mate-
rial becomes available, it would be inter-
esting to investigate the amount of intra-
specific variation in ovipositor length.

Cocoon spinning by larvae of Heteros-
pilus also varies among the species de-
scribed here. Matthews (1968) mentioned
that H. microstigmi spins an opaque white
cocoon, usually near the bottom of the
host cell. Melo and Campos (1993) ob-
served that larvae of H. mntthezvsi spin
very rigid cocoons, which adults of M.
myersi are apparently unable to open.
Cells of M. myersi successfully parasitized
by H. matthezosi become useless after
emergence of the adult parasitoid. The lar-
vae of H. richardsi spin cocoons somewhat
thicker than those of H. microstigmi, but

much thinner than in H. matthewsi. On the
other hand, mature larvae and pupae of
H. brasilophagous, new species, and H. ar-
leiophagous, new species, were not encased
by any sort of conspicuous cocoons. This
apparent absence of cocoon spinning be-
havior needs confirmation, however, be-
cause cells of Microstigmus are lined with
silk and it would have been easy to over-
look a very loose cocoon adhered to the
cell walls. Newly emerged adults of Het-
erospilus leave their host's nests directly to
the outside by chewing a hole in the nest
wall, without passing through the nest en-
trance. In active nests, these emergence
holes are sealed later on with silk and par-
ticles by the Microstigmus females.

Body color also varies in an interesting
way among the present species of Heter-
ospilus. Species of Heterospihis attacking
Microstigmus with a predominately black
integument (including S. alini) also have a
dark body color, whereas species associ-
ated with light colored Microstigmus (pre-
dominately or entirely yellow) have a light
body color. The significance of this varia-
tion in body color is unknown.

The species of Heterospilus show a high
degree of specificity regarding their hosts.
Four of the present species are associated
with only one species of Microstigmus (H.
matthewsi, H. arleiophagus, H. hrnsilophagus)
or at most with a group of closely related
species (H. microstigmi). Only H. richardsi
is known to attack hosts in different
groups, but even in this case the two hosts
nest in the same type of habitat. Most of
the material used in the present study was
collected by the second author while
studying the biology of Microstigmus and
Spilomena wasps in the region of Vi(josa
(Minas Gerais, southeastern Brazil). Ex-
cept for H. microstigmi, the remaining four
species coexist sympatrically in this local-
ity. The hosts of H. microstigmi, Microstig-
mus species in the group theridii, are
known only from the Amazon basin and
Central America. Judging from this degree
of specificity, additional collecting will

16

Journal of Hymenoptera Research

probably reveal twice as many species of
Heterospiliis attacking these crabronid
wasps.

Additional biological notes are included
with the descriptions of the Heterospiliis
species presented below.

TAXONOMY OF HETEROSPILUS

Although the genus Heterospiliis is large
and badly in need of study for the entire
Western Hemisphere, preliminary study
indicates there will be many distinct spe-

cies groups. The species in this study fall
in a group with the following characters:
vertex, mesonotum and mesopleuron co-
riaceous (Figs. 9-11); flagellum unicolored
without white band or tip; metasoma ter-
gum 24-3 with two transverse scrobiculate
grooves which enclose a slightly raised
median area (Figs. 6-8, see arrow on Fig.
6). The following key is based heavily on
characters found in the female although it
will work with most males also. The iden-
tification of males is best done by associ-
ated rearings.

KEY TO HETEROSPILUS SPECIES PARASITIC ON MICROSTIGMUS AND SPILOMENA

1. Body of both sexes entirely brown 2

Body of both sexes honey yellow, metanotum and metasomal terga occasionally marked
with brown 3

2(1). Ovipositor of female equal to or longer than metasoma; fore wing with vein 3RSa longer

than r-m (Fig. 1); hind femur yellow or at most yellowish-brown

richardsi Marsh and Melo, new species

- Ovipositor Vi to Vi length of metasoma; vein 3RSa equal to r-m (Fig. 2); hind femur
brown on apical %, yellow on basal Va matthewsi Marsh and Melo, new species

3(1). Antennal flagellum yellow, first flagellomere longer than second; forewing vein 3RSa
longer than r-m (Fig. 5); ovipositor of female usually equal to length of second and third
metasomal terga combined microstigmi Richards

- Antennal flagellum black, first flagellomere usually equal to second; forewing vein 3RSa
equal to or shorter than r-m; ovipositor usually equal to length of first metasomal tergum 4

4(3). Fore wing vein 3RSa equal to vein r-m (Fig. 4); median transverse area of metasomal
terga 2 + 3 (between transverse scrobiculate grooves) coriaceous medially, striate laterally

brasilophagits Marsh and Melo, new species

Fore wing vein 3RSa shorter than vein r-m (Fig. 3); median transverse area of metasomal
terga 2+3 entirely coriaceous arleiophagits Marsh and Melo, new species

Heterospilus arleiophagus Marsh and
Melo, new species

(Fig. 3)

Female. — Body color: head yellow, palpi
light yellow, antennal flagellum black,
first flagellomere yellow at base, pedicel
black, scape yellow with black longitudi-
nal stripe on outer edge; mesosoma yel-
low, metanotum, sides of scutellum and
propodeum occasionally black; legs yel-
low, wings slightly dusky, veins brown;
metasoma yellow, first tergum occasion-
ally marked with brown laterally, terga 2-

4 marked with brown medially. Body
length: 2.5 mm. Head: vertex and frons
weakly coriaceous, face and temple
smooth; 26-27 antennomeres, first flagel-
lomere equal in length to second; malar
space about Vi eye height. Mesosoma:
pronotum coriaceous dorsally with medi-
an scrobiculate groove; mesonotal lobes
coriaceous, median lobe with median lon-
gitudinal depression, notauli scrobiculate
and meeting in a wide rugulose area near-
ly as wide as scutellum; scutellum coria-
ceous; mesopleuron coriaceous, sternaulus
short and weakly scrobiculate; propo-

Volume 8, Number 1, 1999

17

Figs. 1-5. Wings of Heterospilus species.: 1, rlchardsi, new species; 2, matthcwsi, new species; 3, arkwplmgus,
new species; 4, brasihpljagus, new species; 5, micwsfiginl Richards.

deum rugose with small basal lateral co-
riaceous spots. Legs: fore tibia with row
of 4-6 short spines. Wings: fore wing vein
3RSa shorter than vein r-m (Fig. 3). Me-
tasoma: first tergum slightly longer than
apical width, carinate rugulose, raised me-
dian area not set off by carinae; second
tergum carinate, transverse median area
between two transverse scrobiculate
grooves entirely coriaceous; third tergum
coriaceous; remainder of terga weakly co-
riaceous; ovipositor as long as first meta-
somal tergum.

Male. — Agrees with description of fe-
male except as follows: scape usually en-
tirely yellow; hind wing with oblong stig-
ma at base (as in Fig. 1).

Holoti/pe. — Female: BRAZIL, Viqosa,
MG, January 28, 1990, G. A. R. Melo, col-
lected in nest of Microstigmus arlei, nest
330. Deposited in the Museu de Zoologia,
Universidade de Sao Paulo, Brazil.

Paratypes. — BRAZIL: 2 females, 1 male.

same data as holotype with additional
dates of August 7, 1990 and February 2,
1992, nests 330, 367, 368, 603. Deposited in
the Museu de Zoologia, Universidade de
Sao Paulo, Brazil.

Biology. — Reared from nests of Micros-
tigmus arlei Richards. This Microstigmus
species makes suspended petiolated nests.
The nests have been found attached only
to surfaces other than plant leaves, like in-
clined tree trunks or branches, earth bank
walls, and structural timber of exposed
roofs of huts; the nests reported by Rich-
ards (1972) were hanging from the face of
a rock. Cocoons have not been observed
for this species.

Etymology. — The species name is based
on the host wasp.

Heterospilus brasilophagus Marsh and
Melo, new species

(Fig. 4)
Female. — Body color: head yellow, palpi
light yellow, antennal flagellum black.

18

Journal or Hymenoitera Research

first flagellomere yellow at base, pedicel
black, scape yellow with black longitudi-
nal stripe on outer edge; mesosoma yel-
low, metanotum, sides of scutellum and
scutellar sulcus black; legs yellow; wings
slightly dusky, veins brown; metasoma
yellow, terga 2-3 marked with brown.
Body length: 2.5 mm. Head: vertex and
frons weakly coriaceous, face and temple
smooth; 26-27 antennomeres, first flagel-
lomere equal in length to second; malar
space about Vi eye height. Mesosoma:
pronotum coriaceous dorsally with medi-
an scrobiculate groove; mesonotal lobes
coriaceous, median lobe with median lon-
gitudinal depression, notauli scrobiculate
and meeting in a wide longitudinal cari-
nate area nearly as wide as scutellum; scu-
tellum coriaceous; mesopleuron coria-
ceous, sternaulus short and weakly scro-
biculate; propodeum rugose with small
basal lateral coriaceous spots. Legs: fore
tibia with row of 4-6 short spines. Wings:
fore wing vein 3RSa usually as long as r-
m (Fig. 4). Metasoma: first tergum slightly
longer than apical width, carinate rugu-
lose, raised median area not set off by ca-
rinae; second tergum carinate, transverse
median area between two transverse scro-
biculate grooves coriaceous, striate later-
ally; third tergum coriaceous; remainder
of terga weakly coriaceous; ovipositor as
long as first metasomal tergum.

Male. — Essentially as in female; hind
wing with oblong stigma at base; with 25-
27 antennomeres.

Holoti/pe. — Female: BRAZIL, Vi^osa,
MG, February 15, 1992, G.A.R. Melo, col-
lected in nest of Micwstigmus brasiliettsis,
nest 582. Deposited in the Museu de Zoo-
logia, Universidade de Sao Paulo, Brazil.

Pamti/pes. — BRAZIL: 5 males, same data
as holotype with additional date of July
25, 1992, nests 579, 580, 619. Deposited in
the Museu de Zoologia, Universidade de
Sao Paulo, Brazil.

Biology. — Reared from nests of Micw-
stigmus brasiliensis Melo. This Microstigmus
species builds the type of pendulous nests

considered typical for this genus (see
Richards 1972, West-Eberhard 1977); its
nests were described and illustrated in
Melo (1992).

Etymology. — The species name is based
on the host wasp.

Heterospiliis matthezvsi Marsh and
Melo, new species

(Figs. 2, 6)

Female. — Body color: head brown, palpi
yellow; scape yellow with brown longi-
tudinal strip laterally, pedicel brown, fla-
gellum dark brown; mesosoma and me-
tasoma dark brown; legs yellow, hind fe-
mur brown on apical %; wings hyaline,
veins light brown, tegula yellow. Body
length: 3 mm. Head: vertex and frons fine-
ly coriaceous, temple smooth, face smooth
with fine striations laterally; 26-28 anten-
nomeres; malar space about V3 eye height;
maxillary palpus longer than fore tarsus.
Mesosoma: pronotum coriaceous and
shining dorsally with median scrobiculate
groove; mesonotal lobes finely coriaceous,
median lobe with median longitudinal de-
pression, scutellum finely coriaceous and
shining; mesopleuron coriaceous, sternau-
lus short and weakly scrobiculate; propo-
deum rugose, median rugae more distinct
indicating vague areola, small basal lateral
coriaceous spots. Legs: fore tibia with row
of 4-5 short spines on anterior edge.
Wings: fore wing with vein 3RSa equal in
length to or slightly longer than vein r-m
(Fig. 2). Metasoma (Fig.6): first tergum
slightly longer than apical width, carinate
rugulose, median raised area set off by
complete distinct longitudinal carinae;
second tergum weakly carinate coria-
ceous, ending in distinct transverse scro-
biculate groove; third tergum with second
transverse scrobiculate groove which
meets first groove at sides, tergum cari-
nate coriaceous before this groove and be-
yond to end of tergum; remainder of terga
finely coriaceous and shining; ovipositor
V3 to V2 length of metasoma.

Volume 8, Number 1, 1999

19

Male. — Essentially as in female; hind
wing with oblong stigma at base.

Holotype. — Female: BRAZIL, Viqosa,
MG, February 10, 1992, G. A. R. Melo, col-
lected in nest of Microstigmus myersi, nest
586. Deposited in the Museu de Zoologia,
Universidade de Sao Paulo, Brazil.

Paratypes. — BRAZIL: 3 females, 2 males,
same data as holotype with additional
dates of March 16, 1992, July 31, 1992, Jan-
uary 28, 1990, nests 585, 586, 610, 620, 637;
1 female, Vargem Alta, ES, September 4,
1992, J. N. C. Louzada, collected in nest of
MicrosHginus myersi, nest 637; 1 male, Belo
Horizonte, MG,' July 18, 1992, J. N. C. Lou-
zada, collected in nest of Microstigmus
myersi. Deposited in the Museu de Zoolo-
gia, Universidade de Sao Paulo, Brazil.

Biology. — Reared from the nests of Mi-
crostigmus myersi Turner. Some aspects of
the biology of H. matthewsi were presented
in Melo and Campos (1993); this species
was referred to as Heterospilus sp. in their
paper. The rigid cocoons spun by its lar-
vae set H. matthewsi apart from other Het-
erospilus attacking Microstigmus, whose
larvae spin only a thin cocoon or no co-
coon at all. It would be interesting to in-
vestigate the significance of these rigid co-
coons.

Distribution. — Known only from Brazil.

Etymology. — Named for R. W. Matthews
who gave the first complete description of
the biology of Heterospilus microstigmi (see
Matthews 1968).

Heterospilus microstigmi Richards

(Figs. 5, 7, 9-11)

Heterospilus microstignti Richards 1935:131. Ho-
lotype female, deposited in The Natural His-
tory Museum, London.

Female. — Body color: head, mesosoma
and metasoma yellow or honey-yellow,
mesonotal lobes, metanotum, propodeum
dorsally, and metasomal terga 1-4 often
marked with brown; antenna varying
from entirely brown to scape, pedicel and
basal flagellomeres yellow, apical flagel-

lomeres brown; legs yellow; wings hyaline
or slightly dusky, veins light brown, te-
gula yellow. Body length: 2.5-3.0 mm.
Head: vertex and frons finely coriaceous
(Fig. 11), face and temple smooth; 24-28
antennomeres; malar space about Vs eye
height; maxillary palpus longer than fore
tarsus. Mesosoma (Figs. 9, 10): pronotum
coriaceous and shining dorsally with me-
dian scrobiculate groove; mesonotal lobes
finely coriaceous, median lobe with me-
dian longitudinal depression, scutellum
finely coriaceous; mesopleuron coria-
ceous, sternaulus short and weakly scro-
biculate; propodeum rugose, median ru-
gae more distinct indicating vague areola,
small basal lateral coriaceous spots. Legs:
fore tibia with row of 4-5 short spines on
anterior edge. Wings: fore wing with vein
3RSa about equal in length to vein r-m
(Fig. 5). Metasoma (Fig. 7): first tergum
slightly longer than apical width, carinate
rugulose, median raised area set off by
short indistinct basal carinae; second ter-
gum carinate rugulose, ending in distinct
transverse scrobiculate groove; third ter-
gum with second transverse scrobiculate
groove which meets first groove at sides,
tergum carinate rugulose before this
groove, strongly coriaceous beyond to end
of tergum; remainder of terga coriaceous;
ovipositor Vi to % length of metasoma.

Male. — Essentially as in female; hind
wing with oblong stigma at base.

Distribution. — Trinidad, Costa Rica, Bra-
zil. For this study we have seen 23 speci-
mens from the following localities in Bra-
zil: Manaus, AM; Beruri, AM; Vi^osa, MG;
Ma to G rosso.

Biology. — Previously reared from nests
of Microstigmus theridii Ducke and M.
comes Krombein (Richards 1935; Matthews
1968, 1991). Specimens from this study
have been reared from nests of species of
the M. theridii group which includes M.
comes (nests 442, 443, 447, 531). This spe-
cies group is restricted to northern South
America and Central America (Melo un-
publ.)

20

Journal of Hymenoptera Research

i-iia^i

IP'JQ

^^^^^^T''^'?^^* ^

1B^^H| ■ J^ ■ ■ r-.

^ J

H '^

^^^Bn/^^^^^^^^Ha|

HW ''' '^j^M

^^Sl

^^^^^^^^^^H

^^^pHjk

^^3

Figs. 6-11. Body parts of Hcterospilus species: 6, metasoma, iiuittlicwsi, new species (arrow points to raised
median area mentioned in the descriptions); 7, metasoma, iiuciosti^iiii Richards; 8, nietasoma, liclmnisi, new
species; 9, mesosoma, lateral view, inicroiitignn; 10, mesosoma, dorsal view, inicwstigDii; 11, vertex, iincrostigiin.

Heterospilus richardsi Marsh and Melo,
new species

(Figs. 1, 8)

Female. — Body color: head brown, face
light brown, palpi yellow; scape yellow
with brown longitudinal strip laterally,
pedicel brown, flagellum dark brown; me-
sosoma and metasoma dark brown; legs
yellow; wings hyaline, veins light brown,
tegula yellow. Body length: 3 mm. Head:
vertex and frons finely coriaceous, temple
smooth, face smooth with fine striations
laterally; 24-26 antennomeres; malar
space about % eye height; maxillary pal-
pus longer than fore tarsus. Mesosoma:
pronotum coriaceous and shining dorsally
with median scrobiculate groove; mesono-
tal lobes finely coriaceous, median lobe
with median longitudinal depression, scu-

tellum smooth and shining; mesopleuron
coriaceous, sternaulus short and weakly
scrobiculate; propodeum rugose, median
rugae more distinct indicating vague are-
ola, small basal lateral coriaceous spots.
Legs: fore tibia with row of 4-5 short
spines on anterior edge. Wings: fore wing
with vein 3RSa longer than vein r-m (Fig.
1). Metasoma (Fig. 8): first tergum slightly
longer than apical width, carinate rugu-
lose, median raised area set off by short
indistinct basal carinae; second tergum
carinate rugulose, ending in distinct trans-
verse scrobiculate groove; third tergum
with second transverse scrobiculate
groove which meets first groove at sides,
tergum carinate rugulose before this
groove, smooth beyond to end of tergum;
remainder of terga smooth and shining;

Volume 8, Number 1, 1999

21

ovipositor as long as or longer than me-
tasoma.

Male. — Essentially as in female except as
follows; scape usually entirely yellow;
hind wing with oblong stigma at base.

Holotype. — Female: BRAZIL, Araponga,
MG, March 22, 1992, G. A. R. Melo, col-
lected in Spilomena aliui nest. Deposited in
the Museu de Zoologia, Universidade de
Sao Paulo, Brazil.

Parah/pes. — BRAZIL: 1 female, 1 male,
same data as holotype; 1 male, Vi^osa,
MG, April 26, 1989, collected in nest of
Spilomena alini. COSTA RICA: 3 females, 2
males, Heredia, Estacion Biol. La Selva,
10°25'N, 84°0'W, 80m, June 14, 1996,
reared from nests of Micwstigmns sp.,
nests 677, 680, GAR Melo. Deposited in
the Museu de Zoologia, Universidade de
Sao Paulo, Brazil and the national Muse-
um of Natural History, Washington, DC.

Biology. — Reared from the nests of Spi-
lomena alini Antropov in Brazil and an un-
described species of Microstigmus from
Costa Rica closely related to M. xanthos-
celes Melo and Matthews. Spilomena alini
excavates its nests in earth bank walls and
in small soil clumps hanging from rootlets
in banks (Carvalho and Zucchi 1989; Melo
unpubl.; see Fig. 1 in Melo and Campos
(1993) for an illustration of this kind of
habitat). Some nests have also been found
inside abandoned mud cells of eumenine
wasps hanging from roots in banks (in this
case, the nest tunnels and cells were dug
in the loose detritus filling up the mud
cells). Hetewspilus richardsi was reared
only from nests of S. alini built in soil
clumps. In these nests, some of the host
cells are close to the surface, especially in
the smaller clumps, and therefore within
reach of the parasitoid ovipositor. Nests
built within bank walls are apparently
protected from parasitism by Heterospilus.

The nests of the second host of H. ri-
chardsi, Microstigmus sp., are small to me-
dium bags (3-12 mm long) built on hang-
ing rootlets in earth banks or in tree
trunks. The external walls of the nest are

made of soil particles (for nests in banks)
or particles of dead wood (nests in tree
trunks) aggregated with silk from the fe-
male's silk glands. The central part of the
nest has a sponge-like appearance and is
formed by anastomosing pillars and chan-
nels; except in the upper part of the nest,
this central portion is not in contact with
the external walls. The brood cells are lo-
cated in the central portion. It seems that
only small nests, in which the cells are rel-
atively close to the nest surface, are subject
to parasitism by Heterospilus. One female
and one male of H. richardsi were reared
from a nest with six Microstigmus females
(nest 677), while two females and one
male were reared from a nest with only
one Microstigmus female (nest 680). Four
additional nests, containing four, five, 11
and 27 adult Microstigmus respectively,
produced no Heterospilus.

Distribution. — Brazil, Costa Rica.

Etymology. — Named for O. W. Richards
who described the first species of Heter-
ospilus reared from Microstigmus wasps.

Heterospilus species

We have seen one female (nest 598)
reared from the nests of an apparently un-
described species of Microstigmus species
from the bicolor group in Vi^osa. As in sev-
eral other species of the group (West-Eber-
hard 1977), this Microstigmus species feeds
its larvae progressively with Cicadellidae
nymphs. This single female Heterospilus is
similar to microstigmi but differs in having
the body somewhat more coarsely coria-
ceous or punctate and darker wings. More
specimens are needed to determine if it is
a variation of microstigmi or another spe-
cies.

Also, we have seen one badly damaged
female specimen and one male specimen
(nest 403) reared from Microstigmus leuder-
ivaldti species group in Manaus, northern
Brazil. These specimens are similar to mi-
crostigmi but are darker than identified
members of that species. Exact placement

22

Journal of Hymenoptera Research

of these must wait until more undamaged
specimens are collected.

ACKNOWLEDGMENTS

We thank Tom Huddleston, The Natural History
Museum, London, for comparison of specimens of
Hetcrospilus micnistigini with the holotype. Additional
specimens of H. microstigmi were also provided by
Robert Matthews, University of Georgia, Athens. The
assistance of Bruce Cutler, EM Laboratory, University
of Kansas, Lawrence, is greatly appreciated. Partial
funding for the first author's (PMM) studies of the
Doryctinae of Costa Rica was provided by National
Science Foundation grant DEB-972614L

LITERATURE CITED

Carvalho, L. M. and R. Zucchi. 1989. Aspectos fen-
ologicos de Spilonicnn sp. (Hymenoptera, Sphe-
cidae, Peniphredoninae). Revista Brasileira dc Bio-
logia 49:799-807.

Marsh, P. M. 1997. Subfamily Doryctinae, pp. 206-
233. In: K. A. Wharton, P. M. Marsh and M. J.
Sharkey (eds.). Manual of the New World Gen-
era of the Family Braconidae (Hymenoptera).
Special Publication of the International Societi/ ofHy-
menopterists No. 1, 439 pp.

Matthews, R. W. 1968. Nesting biology of the social
wasp Microstigmuf conies (Hymenoptera: Sphe-
cidae, Peniphredoninae). Psyche 75:23-45.

Matthews, R. W. 1991. Iivolution of social behavior
in sphecid wasps, pp. 570-602. ///: K. G. Ross and
R. W. Matthews (eds.). The Social Biology of
Wasps. Comstock, Ithaca.

Matthews, R. W. and K. C. Starr. 1984. Micwstigmiis
comes wasps have a method of nest construction
unique among social insects. Biotiopica 16:55-58.

Melo, G. A. R. 1994. Origin and diversification of the
Mierostignnis wasps (Hymenoptera, Sphecidae),
p. 357. In: A. Lenoir, G. Arnold and M. Lepage
(eds.), Lcs Insectes Sociaiix (12o Congresso Inter-
nacional da lUSSI). Univ. Paris Nord, Paris.

Melo, G. A. R. In press. Comportamento social em
vespas da familia Sphecidae (Hymenoptera,
Apoidea). Oecologica Brasiliensis.

Melo, G. A. R. and L. A. O. Campos. 1993. Nesting
biology of Microstigmus myersi Turner, a wasp
with long-haired larvae (Hymenoptera: Spheci-
dae, Pemphredoninae). Journal of Hymenoptera
Research 2:183-188.

Melo, G. A. R. and R. W. Matthews. 1997. Six new
species of Microstigmus wasps (Hymenoptera:
Sphecidae), with notes on their biology. Journal
of Natural History 31:421-437.

Menke, A. S. 1989. Arpactofihilus reassessed, with
three bizarre new species from New Guinea (Hy-
menoptera: Sphecidae: Pemphredoninae). Inver-
tebrate Taxonomy 2:737-747.

Richards, O. W. 1935. Two new parasites of aculeate
Hymenoptera from Trinidad. Stylops 4(6):131-
133.

Richards, O. W. 1972. The species of the South Amer-
ican wasps of the genus Microstigmus Ducke (Hy-
menoptera: Sphecidae, Pemphredoninae). Trans-
actions of the Royal Entomological Society of London
124:123-148.

Shaw, M. R. and T. Huddleston. 1991. Classification
and Biology of Braconid Wasps (Hymenoptera:
Braconidae). Handbooks for the Identification of Brit-
ish Insects 7(11):1-126.

West-Eberhard, M. J. 1977. Morphology and behavior
in the taxonomy of Microstigmus wasps, pp. 123-
125. Proceedings of the 8th International Congress of
the lUSSI. Wageningen, Holanda.

J. HYM. RES.
Vol. 8(1), 1999, pp. 23-34

Cretaceous Digger Wasps of the New Genus Bestiola Pulawski and
Rasnitsyn (Hymenoptera: Sphecidae: Angarosphecinae)

Alexandr p. Rasnitsyn, Wojciech J. Pulawski, and Xavier Martinez-Delclos

(APR) Paleontological Institute, Russian Academy of Sciences, Moscow 117647, Russia; E-mail;

rasna@glasnet.ru; (WJP) Department of Entomology, California Academy of Sciences, Golden

Gate Park, San Francisco, California 94118, USA; E-mail: wpulawski@calacademy.org; (XMD)

Departament de Geologia Dinamica, Geofisica y Paleontologia, Facultad de Geologia,

Universidad de Barcelona, 08071 Barcelona, Spain; E-mail: delclos@natura.geo.ub.es

Absfrncf. — The new genus Bestiola Pulawski and Rasnitsyn, characterized by a unique wing
venation, is described for the following four new species from the Lower Cretaceous: hispanica
Martinez Delclos and Rasnitsyn (type species) from northeastern Spain, communis Pulawski and
Rasnitsyn from central Mongolia, subpetiolata Pulawski and Rasnitsyn from central Mongolia,
and temiipes Pulawski and Rasnitsyn from Eastern Siberia. The genus is assigned to Angaro-
sphecinae Rasnitsyn, 1975, new status by Rasnitsyn, which is treated as a paraphylehc Lower
Cretaceous subfamily of Sphecidae. The family name Baissodidae Rasnitsyn, 1975, is synonymized
with Angarosphecidae Rasnitsyn, 1975, by Rasnitsyn.

Sphecid wasps of the archaic subfamily
Angarosphecinae (= Baissodinae) are the
most abundant taxa among the Early Cre-
taceous Hymenoptera, particularly in the
middle interval of that epoch, probably af-
ter Berriasian and before Albian, 140-113
myr before present (Rasnitsyn et al. 1998).
Only a fraction of the material accumulat-
ed in the collections has been described
(Evans 1969; Rasnitsyn 1975, 1986, 1990;
Hong 1984; Zhang 1985, 1992; Darling and
Sharkey 1990; Jarzembowski 1991; Ansor-
ge 1993; Ren et al. 1995; Rasnitsyn et al.
1998). The Early Cretaceous fossils de-
scribed here originate from three distant
areas of Eurasia (central Mongolia, East-
ern Siberia, and Spain), but have a unique
wing venation and are all approximately
the same age.

Most of the specimens examined were
collected in Bon Tsagan, Central Mongo-
lia, a rich fossil site 5-8 km north of Bon
Tsagan Nuur (= Bon Tsagan Lake), in
Bayanhongor Aymag (= Region). Speci-
mens were impressed in marl of the Khur-
ilt rock unit, Bon Tsagan Series (Sinitza

1993), possibly of Aptian age (Ponomar-
enko 1990). This hymenopteran assem-
blage is related to the Wealden Super-
group of the Southern England (Valangi-
nian to Barremian: Rasnitsyn et al. 1998).

Siberian material was collected in two
localities east of Lake Baikal. One is Bais-
sa, a riverside outcrop on the left bank of
the Vitim River, 3 km downstream of the
former lodge Baissa and 45 air km up-
stream of the Romanovka Village, Buryat
Republic. The fossils, impressed in marl of
the Zaza Formation, are related to the Pur-
beck of South England (Berriassian), based
on their hymenopteran assemblages (Ras-
nitsyn et al. 1998). The other Siberian lo-
cality, Semyon, is at Semyon Creek, 3.5
km SW of Elizavetino Village, west of Chi-
ta, Chita Oblast'. The age of insectiferous
mudstones, disputable within the Early
Cretaceous, is correlated either with Bais-
sa (Zherikhin 1978) or with Bon-Tsagan
(Dmitriev and Zherikhin 1988).

The Spanish specimen originates from
the Montsec Range, central Lerida Prov-
ince, and comes from lithographic lime-

24

Journal of Hymenoptera Research

Stones named La Pedrera de Meia, located

5 km W of Santa Maria de Meia in La No-
guera comarca, possibly of Berriassian-Va-
langinian age (Martinez-Delclos 1995).

The following morphological structures,
variously termed in the literature, are here
defined or redefined as follows for clarity
and convenience sake:

- mesosoma: thorax and propodeum
combined;

- metasoma: abdomen excluding the pro-
podeum (= gaster of Bohart and Menke
1976);

- metapostnotum: propodeal enclosure of
Bohart and Menke 1976;

- spiracular lobe (as in Rasnitsyn 1988):
pronotal lobe of Bohart and Menke
1976;

- adlateral lines: parapsidal line of Bohart
and Menke (1976); we prefer this term
to avoid confusion, as parapsidal lines
of other entomologists correspond to
notauli of most hymenopterists;

- cell l+2r: submarginal cell I of Bohart
and Menke (1976);

- 3r: marginal cell of Bohart and Menke
(1976);

- 2rm and 3rm: submarginal cells II and
111 of Bohart and Menke (1976);

- Imcu and 2 mcu: discoidal cells 1 and II
of Bohart and Menke (1976);

- crossveins 2r-m and 3r-m: distal mar-
gins of 2rm and 3rm, respectively (as in
Richards, 1956, and Gauld and Bolton,
1988), and corresponding to Ir-m and
2r-m of Bohart and Menke, 1976); unlike
the latter two authors, we call Ir-m the
vein that extends from cell l+2r to cell
Imcu (i.e., the vein that separates the
basal cell from cell 2rm) in the xyelid
genus Pleroiieurn;

- vein IRS: a veinlet between basal cell ( =
medial cell of Bohart and Menke, 1976)
and cell l-(-2r;

- vein 2RS: a veinlet between cells l-l-2r
and 2rm;

- vein 2r-rs: a veinlet between cells l-l-2r
and 3r, called 2r by Bohart and Menke,
1976.

The abbreviation PIN stands for the Pa-
leontological Institute, Russian Academy
of Sciences, Moscow, Russia.

TAXONOMY

Family Sphecidae Latreille

Subfamily Angarosphecinae Rasnitsyn,
new status

Angarosphecidae Rasnitsyn 1975:109. Type ge-
nus: Angnrosphcx Rasnitsyn 1975:110.

Baissodidae Rasnitsyn:1975:122. Type genus:
Bnissodes Rasnitsyn 1975:123. New synonym
by Rasnitsyn.

Angarosphecinae are archaic Mesozoic
wasps that lack the synapomorphies of
any extant sphecid subfamily. They may
be paraphyletic with respect to other Apo-
idea because they are not defined by any
synapomorphy. They are treated here as a
subfamily of Sphecidae because there is
evidence in some specimens of two
unique synapomorphies of Apoidea (they
also lack any synapomorphy that would
ally them with any other Aculeata). In
particular, Pompilopterus corpus Rasnitsyn
and Jarzembowski has an elongate spirac-
ular lobe, pronotum thickened preapical-
ly, and an enlarged metapostnotum (Ras-
nitsyn, Jarzembowski, Ross 1998, Fig. 36),
and Angawsphex myrmicopterus Rasnitsyn
has an enlarged metapostnotum (Rasnit-
syn 1980, Fig. 172). A large metapostno-
tum is also found in Bestiola tenuipes (Fig.
7). None of the Angarosphecinae has plu-
mose setae or enlarged hindbasitarsi typ-
ical of bees. The form of the adlateral line
of Bestiola and other Angarosphecidae dif-
fers from that of all extant Apoidea in that
it extends to the posterior margin of the
mesoscutum, as pointed out to us by M.
A. Prentice (oral communication). This in-
dicates that Angarosphecinae very proba-
bly represent the most basal lineage of
known Apoidea.

Rasnitsyn (1975) recognized Baissodi-
dae based on the presence of a unique me-
dian scutal sulcus believed to be lacking
in all other non-bethyloid Aculeata (in-

Volume 8, Number 1, 1999

25

eluding Angarosphecidae). Subsequently,
he (Rasnitsyn 1980) included Angarosphex
in the Sphecidae and hypothesized that
the Baissodidae were sphecid ancestors.
However, lAngarospihex pallidus Rasnitsyn,
1986, from the lowermost Lower Creta-
ceous of Mongolia, combines the wing ve-
nation of A)igarosphex with the presence of
a median scutal sulcus, although the latter
is only slightly indicated. Because of this
combination, Baissodidae are here synon-
ymized with Angarosphecidae.

Bestiola Pulawski et Rasnitsyn, new
genus

Name derivation. — Bestiola, Latin for
small beast. Gender feminine.

Ti/pe species. — Bestiola hispanica Martinez
Delclos et Rasnitsyn, new species. Lower
Cretaceous of Spain.

Recognition. — Bestiola is easily recognized
by its unique wing venation (Figs 1-8). It
has three radiomedian (= submarginal)
cells, and cell 2rm receives veins Im-cu and
2m-cu (= both recurrent veins). The com-
bination of three unusual features differ-
entiates it from all other sphecid genera
with these characteristics, both extinct and
extant: 1. cell 3rm broader on the costal
side than on the anal side (as in the North
American genus Xenosphex and some Pa-
lanis); 2. crossvein 3r-m joining RS near the
distal end of the latter (as in the North
American genus Eucerceris and some Pala-
nis); and 3. crossvein 2r-m equidistant from
2m-cu and 3r-m or closer to the latter (as
in the Old World Tacln/sphex brevipennis
Mercet, and several other Larrini and some
Diploplectron approach this condition).

Description. — Size medium to large,
length of forewing 5-15 mm. Antenna
with no conspicuous modifications, at
least basal flagellomeres longer than wide
(all flagellomeres in most species). Occip-
ital carina almost circular, reaching hypos-
tomal carina. Ocelli not modified, distant
from eye. Pronotum short, wide, thick-
ened preapically, separated from meson-
otum by a groove (Fig. 8). Mesoscutum

without median scutal sulcus, with long
notauli and adlateral lines; mesopleuron
with episternal and scrobal sulci, hyper-
sternaulus, and possibly omaulus. Meta-
notum short, metapostnotum long, trun-
cated, with median line. Propodeal spira-
cle elongate, slit-like. Forewing: pterostig-
ma well defined; basal vein distant from
pterostigma, evenly arched; cell 3r acu-
minate at wing foremargin; crossvein 2r-
rs longer than width of pterostigma; 2r-m
sinuate or arching outwardly, closer to 2r-
rs than to 3r-m on RS, equidistant from
2m-cu and 3r-m on M or closer to the lat-
ter; 3r-m straight or arching outwardly;
Im-cu received near base of cell 2rm; M
sharply angled at 2m-cu; cu-a interstitial
with M or narrowly postfurcal. Hindwing
venation complete, cu-a meeting Cu well
beyond M-I-Cu fork. Fore and mid femora
with well-defined, narrow trochantellus
(hindleg condition unknown), but no ob-
vious specializations. Metasoma rounded
basally (neither petiolate nor peduncu-
late).

Composition. — Four species from the
Lower Cretaceous of Spain, Eastern Sibe-
ria, and Mongolia, as described hereafter.

Taxonomic position. — Bestiola is a mem-
ber of Aculeata s. s. (= Vespoidea -I- Apo-
idea) as evidenced by its sexually dimor-
phic antennal flagellum of 10 (female) and
11 articles (male). Unlike all Chrysidoidea,
it possesses a complete set of forewing
and hindwing cells. It belongs to Apoidea
because it shares two unique synapomor-
phies of the superfamily: an elongate me-
tapostnotum and a pronotum thickened
preapically. The genus belongs to Spheci-
dae because it lacks the enlarged hindbas-
itarsus and plumose body setae of Apidae
s. 1. The genus shares with most other An-
garosphecinae the position of Im-cu
which inserts near the RS+M fork. This
feature occurs elsewhere only in some oth-
er Apoidea. The genus also lacks any syn-
apomorphy that would place it in any ex-
tant sphecid subfamily and so we attribute
it to Angarosphecinae.

26

Journal of Hymenoptera Research

KEY TO SPECIES OF BESTIOLA

1. Forewing cell 2rm conspicuously narrowing anterad, its costal margin markedly shorter than
crossvein 2r-rs; the latter emerging shortly after pterostigma's midlength (Fig. 8)

Bestiola subpetiolata Pulawski and Rasnitsyn, new species

- Forewing cell 2rm moderately narrowing anterad, its costal margin about as long as cross-
vein 2r-rs; the latter emerging markedly beyond pterostigma's midlength 2

2. Forewing length 5 mm; crossvein 2r-rs markedly shorter than 2RS (Fig. 1); hindfemur and
gastral terga with pale spots (Fig. 1); metasoma conspicuously setose (Fig. 1)

Bestiola hispanica Martinez Delclos and Rasnitsyn, new species

- Forewing length 11.0-14.5 mm; crossvein 2r-rs longer to minimally shorter than 2RS (Figs.
2-7); hindfemur and gastral terga without pale spots (Figs. 2-7); metasoma not setose or
less conspicuously setose (Fig. 5) 3

3. Head, wing veins, and legs except coxae light; forefemur slightly more elongate (Fig. 7)

Bestiola tentiipes Pulawski and Rasnitsyn, new species

- Head, wing veins, and legs dark; forefemur slightly stouter (Figs. 2, 3)

Bestiola communis Pulawski and Rasnitsyn, new species

Bestiola hispanica Martinez Delclos and
Rasnitsyn, sp. n.

(Fig. 1)

Name derivation. — Hispanica, Latin for
Spanish.

Recognition. — Bestiola hispanica differs
from its congeners by its small size (fore-
wing length 5 mm rather than 11.0-14.5
mn\), crossvein 2r-rs markedly shorter
than 2RS (longer to minimally shorter in
the other species), the presence of pale
spots on the hindfemur and gastral terga,
and a conspicuously setose metasoma.

Description. — Female unknown, male:
Fig. 1. Body dark (including wing veins),
but hindfemur with pale spot posteroapi-
cally and terga 1-lV each with a pair of
preapical spots. Head and metasoma con-
spicuously setose. Flagellomeres nearly
equal in width, longer than wide, becom-
ing shorter toward apex. Head relatively
small, narrowing toward mouthparts;
gena somewhat inflated; malar space half
length of eye; clypeal free margin project-
ing mesally; hindocellus separated from
eye by about its own diameter. Forewing:
posterior pterostigmal margin straight; 2r-
rs joining pterostigma near the latter's
apex, about as long as costal margin of cell
2rm and half as long as 2RS; 3r-m straight;

costal margin of cell 2rm about twice as
long as anal margin; cu-a almost intersti-
tial with M+Cu fork. Hindwing with long
row of hamuli (10 preserved) and cu-a an-
gling at Cu. Legs not elongate; hindfemur
as long as head width, moderately thick
subbasally, not attenuated apically, its
dorsal margin convex except subapically,
ventral margin almost straight (ventral
margin slightly convex on right femur,
probably due to fossil compression).
Hind tibia 1.25 times as long as femur,
with no spines but with one spur. Hind-
tarsus slightly longer than tibia, basitarsus
longer than tarsomeres II-IV combined.
Apical gastral segments and genitalia not
preserved. Body length ca 9 mm, distance
from forewing base to apex of cell 3rm 5.1
mm.

Material examined. — Holotype (LP92/
SC/3662): male, Spain: Lerida Province:
La Pedrera de Meia 5 km W Santa Maria
de Meia (Institut d'Estudis Ilerdencs, Ler-
ida, Spain).

Bestiola commtmis Pulawski et
Rasnitsyn, new species

(Figs. 2-6)

Name derivation. — Communis, Latin for
common.

Volume 8, Number 1, 1999

27

Fig. 1. Bestwla lufpauicn Martinez-Delclos and Rasnitsyn, new species, holotype: cly — clypeus; cocc — occipital
carina; f — femur; to — occipital foramen; md — mandible; ppl — propleuron; shy — hypostomal suture; sipg —
interpostgenal suture.

Recognition. — The following combina-
tion of characters is unique to Bestiola cotn-
miinis: forewing length 11.0-14.5 mm (5
mm in hispanica); costal margin of cell 2rm
about as long as crossvein 2r-rs (markedly
shorter in suhpetiolata); 2r-rs longer to min-
imally shorter than 2RS (markedly shorter
in hispanica), emerging markedly beyond
pterostigma's midlength (near pterostig-
ma's midlength in suhpetiolata); hindfemur
and gastral terga without pale spots (with
pale spots in hispanica); and forefemur not
elongate (slightly elongate in teniiipcs,

compare Figs. 2 and 3 and 7). Unlike ten-
iiipcs, the body of communis is all dark, in-
cluding the wing veins.

Description. — Female (Fig. 2), sex un-
known in remaining specimens (Figs. 3-
6). Body and appendages uniformly dark,
metasoma inconspicuously setose (Fig. 5).
Scape about as long as midflagellar arti-
cles; pedicel transverse; flagellomeres
more than twice as long as wide, flagel-
lomere 1 almost as long as II and 111 com-
bined, following ones gradually shorter
and thinner toward antennal apex, apical

28

Journal of Hymenoptera Research

Fig. 2. Bcstiola coniiiniiuf Pulawski and Rasnitsyn, new species, hoiotype: d — discrimen (interpleural suture);
liy — hypostoma; hys — hypersternaulus or signum?; Nl — pronotum; oa — omaulus?; ss — scrobal suture; tl —
trochantellus; tr — trochanter; other abbreviations as in Fig. 1.

flagellomere almost 3 times as long as
wide. Head nearly circular in front view;
eyes elongate, widest below midheight,
with inner margin concave; malar space
present; anterior clypeal margin protrud-
ing into wide medial lobe that is shallowly
emarginate apically. Notauli and adlateral
lines complete or nearly so. Mesopleuron
with long, almost straight scrobal sulcus
and hypersternaulus, possibly also with
omaulus. Wing venation as in hispaiiica ex-
cept 2r-rs subequal in length to 2RS and
3r-m arched. Fore- and midfemora thick-
est subbasally, narrow apically, with dor-
sal margin straight and ventral convex;
forefemur as long as head width, midfe-
mur slightly longer; fore- and possibly
midtibiae shorter than respective femora.
Metasoma missing in type series but pres-
ent in specimen PIN 3559/4526 (that is ex-
cluded from type series). Forewing length
13.0-14.5 mm (11.0 mm in the specimen

PIN, 3559/4526 not included into the type
series. Fig. 5).

Material examined. — Hoiotype: Central
Mongolia, Bon-Tsagan, bed 87/8 (PIN.
3559/4525, incomplete female specimen
with propodeum, hindlegs and metasoma
missing).

Paratypes: same locality and bed (PIN,
3559/4528, incomplete specimen with
most of antennae and legs, part of thorax
and all metasoma missing; PIN, 3559/
4530, two damaged wings mixed with
other insect remains in vertebrate drop-
ping).

Excluded from type series but possibly
conspecific: same locality and bed (PIN,
3559/4526, a somewhat damaged speci-
men: Fig. 5); Eastern Siberia, Semyon
(PIN, 2385/2392, an isolated wing with
venation nearly identical to those of coiii-
niiinis and tenuipe^, and attributed to the

Volume 8, Number 1, 1999

29

Fig. 3. Bestwhi ccmmiiiuf Pulawski and Rasnitsyn, new species, paratype PIN, 3559/4528: a — antennal fora-
men; al — adlateral line; h — hypostome; n — notaulus; otherwise as in Figs. 1, 2.

former because of its dark wing veins; Fig.
6).

Bestiola tenttipes Pulawski et Rasnitsyn,
new species

(Fig. 7)

Name derivation. — Tenuipes, from the
Latin words tenuis, thin, and pes, leg;
with reference to the elongate forefemur.

Recognition. — The wing venation of B.
tenuipes is as in communis, but the head,
wing veins, and legs are light rather than

Fig. 4. Bestiola communis Pulawski and Rasnitsyn,
new species, paratype, PIN, 3559/4530.

dark. Also, the forefemur is slightly longer
(compare Figs. 7 with 2 and 3), although
this difference is difficult to quantify. See
Recognition of cotnmunis for differences
with hispanica and subpetiolata.

Description. — Male (Fig. 7). Female un-
known. Antenna and mesosoma (possibly
in part) dark, metasoma infuscated toward
apex, otherwise coloration light (including
wing venation). Thorax with well-defined
although shallow punctures that are about
1 diameter apart, and with moderately
short and moderately dense setae. Basal
flagellomeres unrecognizable, remaining
flagellomeres 2.0-2.5 times as long as wide,
gradually becoming shorter and thinner to-
ward apex. Adlateral line of mesoscutum
complete; scutellum wide, elongate; metas-
cutellum contrastingly short; metapostno-
tum trapezoid, with basal impression, me-
dian longitudinal line, and rounded pos-
terior angles. Propodeal spiracle elongate,
narrow, slightly bent S-like. Wing venation

30

Journal of Hymenoptera Research

5mm

Fig. 5. Bcsticla ? communis Pulawski and Rasnitsyn, new species, PIN, 3559/4526; N3 — metanotum; pp-
propodeum; scl — scutelluni; other abbreviations as in Figs. 1-3.

as in communis. Legs relatively long, fore-
femur slightly longer than head width,
with dorsal margin straight and ventral
margin convex; hindfemur elongate, al-
most symmetrical, attenuated apically,
both dorsal and ventral margins straight
(except basally); midtarsus markedly lon-
ger than midfemur; midbasitarsus shorter

Fig. 6. Bcstiola ? connminis Pulawski and Rasnitsyn,
new species, PIN, 2385/2392.

than three following tarsomeres combined.
Genitalia elongate, ovoid, with smooth
contour, with gonostyle apex narrow
rounded. Body length 21 mm as preserved,
forewing length 11.5 mm.

Material examined. — Holotype: Russia:
Eastern Siberia: Baissa, bed 31 (PIN, 3064/
2055, incompletely preserved male).

Bestiola subpetiolata Pulawski et
Rasnitsyn, new species

(Fig. 8)

Name derivation. — From the Latin petiol-
us, little foot, stalk, stem; and the prefix
sub-, Latin for under, somewhat, less than;
with reference to the shape of forewing
cell 2rm.

Recognition. — The following details of

Volume 8, Number 1, 1999

31

5 mm

Fig. 7. Bestioln teiiuifvs Pulawski and Rasnitsyn, new species, holotype: pN3— metapostnoUim; sp— propodeal
spiracle; other synibols as in Figs. 1-5.

32

Journal of Hymenoitera Research

5 mm

Fig. 8. B('sf/o/(7 fiihpctiolttta Pulawski and Rasnitsyn, new species, luilotype: es — episternal suture; nipl — me-
tapleura; other abbreviations as in Figs. 1-7.

the forewing venation distinguish suhpe-
tiolata from all of its congeners: vein 2r-rs
meeting pterostigma near the latter's mid-
length, costal margin of cell 2rm markedly
shorter than 2r-rs, and anterior end of vein
cu-a closer to wing base than M + Cu fork.
In addition, subapical flagellomeres ap-

pear to be only slightly longer than wide,
thus markedly shorter than in other Bes-
tiola.

Description. — Sex unknown (Fig. 8).
Body moderately dark (including wing
veins) but antennal apex, tarsi (except
hindbasitarsus basally), and metasoma

Volume 8, Number 1, 1999

33

light. Metasoma at least partly setose (se-
tae preserved only along hind margin of
last preserved tergum). Length of basal
flagellomeres about 3 times width, sub-
apical ones probably subquadrate. Eye
large, elongate, almost symmetrical. Malar
space probably short. Pronotum short, spi-
racular lobe not elongate. Mesoscutum:
notauli and adlateral lines complete or
nearly so. Mesopleuron with complete,
crenulate episternal sulcus; and with an-
teriorly crenulate hypersternaulus. Meta-
pleuron wide, crossed by subhorizontal
sulcus. Other thoracic structures unrecog-
nizable due to deformation. Forewing
vein 2r-rs meeting pterostigma near the
latter's midlength, meeting RS near 2rm
(longer than costal margin of cell 2rm),
2rm weakly arching, 3r-m straight, ante-
rior end of cu-a slightly closer to wing
base than M-l-Cu fork. Hindwing vein cu-
a meeting M relatively close to M-I-Cu
fork. Midfemur: dorsal margin straight,
ventral margin convex. Hindfemur about
as long as head height, widest subbasally,
attenuated apically, with dorsal margin
concave in apical half and ventral margin
straight except basally. Midtibia slightly
shorter, hindtibia slightly longer, than re-
spective femur. Mid- and hindtarsi longer
than respective tibiae, respective basitarsi
slightly shorter than following 3 tarso-
meres combined. Metasoma somewhat at-
tenuated basally, with apex missing, but
probably shorter than head and thorax
combined. Body length, as preserved, 10
mm, forewing length 7.0 mm

Material examined. — Holotype: Central
Mongolia, Bon-Tsagan, bed 87/8 (PIN,
3559/4529, incomplete, rather poorly pre-
served specimen with somewhat crum-
pled thorax).

ACKNOWLEDGMENTS

We sincerely thank Michael A. Prentice for his con-
structive criticism of the manuscript, as well as Eric
Crissell and Arnold S. Menke. We are grateful to
Robert L. Zuparko for reviewing an earlier draft of
the manuscript. The project was partly supported by

a grant to A. P. Rasrutsyn from the Royal Society Joint
Project with the Former Soviet Union.

LITERATURE CITED

Ansorge, J. 1993. Bemerkenswerte Lebenspuren und
'fCrftosphcx catnliiniciis n. sp. (Insecta; Hymenop-
tera) aus den unterkretazischen Plattenkalken del
Sierra del Montsec (Provinz Lerida, NE-Spanien).
Neues ]ahrhiicli fiir Ceologie und Paldontologie Moii-
(itshefte 190: 19-35.

Bohart, R. M., and A. S. Menke 1976. Sphecid wasps of
the world. A generic revision. University of Cali-
fornia Press, Berkeley, Los Angeles, London, 1
color pi., ix + 695 pp.

Darling, D. Ch., and M. J. Sharkey. 1990. Order Hy-
menoptera in Grimaldi D. A. (editor). Insects
from the Santana Formation, Lower Cretaceous,
of Brasil. Bulletin of the Ameriain Museum of Nnt-
ural Histori/ 95: 124-129.

Dmitriev, V. Yu, and V. V. Zherikhin. 1988. Changes
in diversity of insect families as revealed by the
method of accumulated appearances, p. 208-215
in A. G. Ponomarenko (editor). Cretaceous hiocen-
otic crisis and evolutum of the insects. Nauka Press,
Moscow. 230 pp. (in Russian).

Evans, H. E. 1969. Three new Cretaceous wasps (Hy-
menoptera). Psi/che 7b: 251-261.

Gauld, I., and B. Bolton. 1988. The Hymenoptera. Brit-
ish Museum (Natural History) and Oxford Uni-
versity Press, Oxford, 332 pp.

Hong, Y.-c. 1984 New fossil insects of Liayang Group
from Laiyang Basin, Shandong Province. Profes-
sional Papers of Stratigraplni and Palaeontology No.
11 1984: 31-41.

Jarzembowski, E. A. 1991. New insects from the
Weald Clay of the Weald. Proceedings of the Ge-
ologists' Association 102: 93-108.

Martinez-Delclos, X. (editor). 1995. Montsec and Mont-
Rat-Alcover, two Konsen'at-Lagerstdtten. Institut
d'Estudis Illerdencs, Catalonia, Spain. 97 pp.

Ponomarenko, A. G. 1990. Insects and the Lower Cre-
taceous stratigraphy of Mongolia, p. 103-108 in
Krassilov V. A. (editor). Non-marine Cretaceous of
the USSR. The Submission from the Conference of
the Soviet Working Group of the IGCP Project 245,
Vladivostok, 19SS. Far Eastern Branch of the USSR
Academy of Sciences, Vladivostok. 226 pp. [In
Russian].

Rasnitsyn, A. P. 1975. Hymenoptera Apocrita of Me-
sozoic. Academy of Sciences of the USSR. Transac-
tions of the Paleontological Institute. 147: 134 pp. [In
Russian].

Rasnitsyn, A. P. 1980. Origin and ei'otution ofhymenop-
terous insect.:. Nauka, Moskva, 189 + two un-
numbered pp. (in Russian[

Rasnitsyn, A. P. 1986. Vespida (= Hymenoptera) in
Rasnitsyn A. P. (editor). In.sects in the Early Cre-
taceous Ecosystems of the West Mongolia. Trans-

34

Journal of Hymenoptera Research

actions of tlw loint Soviet-Mongolian Paleontolcgical
Expedition. No. 28: 154-164 (in Russian).

Rasnitsyn, A. P. 1988. An outline of evolution of the
hymenopterous insects (order Vespida). Oriental
Insects 22: 115-145.

Rasnitsyn, A. P. 1990. Hymenoptera in Ponomarenko
A. G. (editor). Late Mesozoic insects of Eastern
Transbaikalian. Academy of Sciences of the USSR.
Transactions of the Paleontolcgical Institute. 239.
Nauka Press, Moscow: 177-205. [In Russian].

Rasnitsyn, A. P., E. A. Jarzembowski, and A. J. Ross.
1998. Wasps (Insecta: Vespida = Hymenoptera)
from the Purbeck and Wealden Supergroups
(Lower Cretaceous) of Southern England and
their hiostratigraphical and paleoenvironmental
significance. Cretaceous Research 19: 329-391.

Ren, D., L. Lu, Z. Guo, and S. Ji. 1995. Faunae and
stratigraphy of Jurassic-Cretaceous in Beijing and the
adjacent areas. Seismic Publishing House, Beijing.
222 pp.

Richards, O. W. 1956. Handbooks to the identificatwn of
British Insects. H[inienoptera. Introduction and keys
to fanuties. Royal Entomological Society of Lon-
don, London. 94 pp.

Sinitza, S. M. 1993. Jurassic and Lower Cretaceous of the
Central Mongolia. Transactions of the Joint Sox'iet-
Mongoltau Palcontological E.\pcdition. No. 42: 1-239
pp (in Russian).

Zhang, Jun-feng 1985. New data on the Mesozoic fos-
sil insects from Laiyang in Shandong. Geology of
Shandong 1: 23-39.

Zhang, Jun-feng 1992. Descriptions of two new gen-
era and two new species of Baissodidae from
China (Sphecidae, Hymenoptera). Acta Entonio-
logica Sinica 35: 483-489.

Zherikhin, V. V. 1978. Development and changes of
the Cretaceous and Cenozoic faunistic assem-
blages (Tracheata and Chelicerata). Academy of
Sciences of the USSR. Transactions of the Palconto-
logical Institute 165: 1-200.

J. HYM. RES.
Vol. 8(1), 1999, pp. 35-47

Ultrastructure of Imaginal Spermatozoa of Sawflies
(Hymenoptera: Symphyta)

Terence M. Newman and Donald L. J. Quicke

Department of Biology, Imperial College of Science, Technology and Medicine, Silwood Park,

Ascot, Berkshire SL5 7PY U.K.

Abstract. — We present the first ultrastructural study of sperm from representatives of three
superfamilies of sawflies (Hymenoptera: Symphyta): Xyela julii (Xyeloidea), Cephalcia arvensis
(Pamphiloidea) and Tremex sp. (Siricoidea), with particular attention being paid to characters that
may be phylogenetically informative. Differences in the location of the centriolar adjunct, partic-
ularly in relation to the mitochondrial derivatives, would suggest Cephakia has a better claim than
Xyela as having sperm that may be representative of a common ancestral form. The centriolar
adjunct of Cephalcia overlies both mitochondrial derivatives symmetrically, as found in ants and
bees, whereas in Tremex, which its sperm otherwise closely resemble, the centriolar adjunct is
located asymmetrically, abutting a single mitochondrial derivative and thus offsetting the pair of
mitochondrial derivatives longitudinally. Xyela has radically different sperm in terms of size and
both the arrangement and appearance of the organelles, especially acrosomal substructure and
lack of an acrosomal rod.

Very little is known about sperm ultra-
structure among the Hymenoptera com-
pared with most other insect orders (Phil-
lips 1970; Jamieson 1987; Quicke 1997),
and most of the studies that do exist deal
largely with common aculeates such as
bees and ants (e.g. Dallai and Afzelius.
1990; Wheeler and Krutzsch. 1992). How-
ever, an initial study of the spermatozoa
of some species (Quicke et al. 1992) re-
vealed a considerable number of ultra-
structural features that differ between
taxa, raising the possibility that such var-
iation might provide new phylogenetic in-
dicators, as has been possible in many oth-
er groups of insects (Jamieson 1987). The
phytophagous sawflies (Symphyta) consti-
tute a relatively underived basal grade
within the order of Hymenoptera. As such
they are important for our understanding
of the relationships and development of
both the social species of the Aculeata
(ants, wasps and bees) and members of
the paraphyletic group of the ten or eleven
currently recognised, extant superfamilies

generally referred to as the 'Parasitica'.
This is especially so, since the sister group
for the Hymenoptera is not at all certain
at present (Whiting ei al. 1997), and so it
is not possible to make use of outgroup
comparison to determine the ancestral
sperm morphology of the order (Watrous
and Wheeler 1981). Groundplan sperm ul-
trastructure may therefore be determined
best by considering the sperm of those ex-
tant taxa (i.e. the sawflies) which represent
the most basal hymenopteran lineages
(Gibson 1993; Yeates 1995). The only pre-
vious work on sawfly sperm ultrastruc-
ture (Quicke et al. 1992) presented data for
only two of the six symphytan superfam-
ilies, the Tenthredinoidea and the Cephoi-
dea. We have therefore examined sperm
ultrastructure, and in particular that of cell
organelles, in detail in representatives of
three further superfamilies, the Xyeloidea,
Pamphiloidea and Siricoidea, leaving only
the rare, through interesting, Orussoidea
unstudied. Two of the superfamilies ex-
amined here, the Xyeloidea represented

36

Journal of Hymenoptera Research

by Xyela julii (Brebisson) and the Pamphi-
loidea, represented by the pamphiliid, Ce-
phalcia arvensis Panzer have usually been
considered to be among the most primi-
tive of sawflies. In contrast, the Siricoidea
represented by Tremex sp., are close to the
origin of the Apocrita (Rasnitsyn 1980,
1988; Heraty et al. 1994; Vilhelmsen 1997).
The results are discussed in terms of the
likely plesiomorphic states for various
subcellular features in the Hymenoptera.

MATERIALS AND METHODS

Testes were obtained from adult males
of Xyela julii, Ceyhalcia arvensis and Tremex
sp., which had been maintained on dilute
honey solution for a maximum of 3 days.
Xi/ela were collected as adults in Silwood
Park, Berkshire, U.K., the Cephalcia were
reared from larvae collected in Italy and
the Tremex were collected as adults in Cal-
ifornia and couriered to the U.K. for prep-
aration.

Light microscopy. — Vas deferenda and
testes were dissected from living sawflies
in insect saline and teased apart on a clean
microscope slide. After a few minutes to
allow the sperm /spermatodesmata to
swim free of the disrupted tissue, the
slides were dried on an hot plate at c.
80°C. The smear was then flooded with
double-filtered, 0.1% w/w toluidine blue
in 1% w/w aqueous sodium borate and
stained at 80°C until crystallisation of the
stain had started. Following washing in
distilled water they were permanently
stored dry.

Transmission electron microscopy. — Geni-
talia were dissected out under 2% glutar-
aldehyde in phosphate buffered saline
(pH 7.2), and fixed for two hours. Tissue
was transferred to 2% osmium tetroxide in
cacodylate buffer (pH 7.2) for 2 hr. After
another buffer wash, tissue pieces were
dehydrated to 50% ethanol and then fur-
ther fixed with saturated uranyl acetate in
50% ethanol prior to complete dehydra-
tion, embedding in Epon resin and poly-
merisation overnight. Silver sections were

picked-up on to high resolution grids,
stained with uranyl acetate and lead.

RESULTS

Woodwasps of the superfamily Siricoi-
dea are considered to be amongst the most
advanced of the sawflies, sharing a num-
ber of derived morphological features
with the Apocrita (Vilhelmsen 1997). As
has been reported previously for other
sawflies (Quicke et al. 1992), the mature
sperm of siricid, Tremex, stored within the
vas deferens and seminal vesicles are
present in spermatodesmata bundles (Fig.
1 ), though by the time they reach the sper-
matheca of the females they have broken
up completely and only isolated sperm are
present (Naito, personal communication).
In our preparation of Tremex from male
seminal vesicles, a small proportion of iso-
lated sperm were also present but it is not
clear whether they were the result of sper-
matodesmata fragmentation upon fixation
or whether they indicate a normal pre-
transfer phenomenon.

The sperm heads are inserted through-
out the fairly electron-dense and elongate
cap of the spermatodesmata, with those
sperm located more centrally being insert-
ed more anteriorly (Figs 1, 2). As a result,
many different levels of sperm are evident
in a single transverse section of each sper-
matodesmata (Fig. 2). It is therefore pos-
sible, in the same transverse section, to lo-
cate adjacent sperm sectioned through ac-
rosome and acrosomal rod (perforatori-
um), through the nucleus, the basal body
with centriolar adjunct, and through the
axoneme with mitochondrial derivatives.
Also, in transverse section, the acrosome
is clearly seen to have a membrane around
both the outside and around the invagi-
nated portion of the structure (Fig. 4c). Be-
tween the acrosomal membrane and the
plasma menibrane is an electron dense re-
gion extending from the acrosomal mem-
brane (Fig. 4c, arrou'lieads). This may be
comparable to the material reported to
surround the acrosome in other sperma-

Volume 8, Number 1, 1999

37

Fig. 1. Longitudinal section of a spermatodesmatum in the imaginal testes of the sawfly, Tremex (Siricoidea).
A, acrosome; C, centriolar adjunct; M, mitochondrial derivative; N, nucleus; R, acrosomal rod; X, axoneme.
Scale bar = 1.0 nm.

38

Journal of Hymenoptera Research

Figs. 2-3. Features of spermatodesmata and sperm in the imaginal testes of the sawtlv, Tivincx (Siricoidea).
2, transverse sections through several spermatodesmata at different levels showing that the more centrallv
located spermatozoa have their heads inserted more anteriorly; 3, nuclear-associated organelles showing in
3a, the insertion of the acrosomal rod (R) into the anterior of the nucleus (note also the small anterior sac at
the head of the acrcisome), and 3b, the position of the centriolar adjunct (arrowed) in relation to the nucleus
and striated mitochondrial tierivative. Scale bars: 2 = 2.0 p.m; 3 = 0.5 [im.

Volume 8, Number 1, 1999

39

Fig. 4. Organelles of Tremex sperm (Siricoidea) seen in tranverse and oblique section: 4a, showing the cen-
triolar adjuncts (small arrows) of a number of spermatozoa, and in the section indicated by the large arrow
the centriolar adjunct can be seen to make contact with both mitochondrial derivatives; 4b, section at the level
of the centriolar adjunct (arrowed) where it occupies all the extra-axonemal area; 4c, sections through anterior
of nucleus and acrosome showing the acrosomal rod (r) fitting tightly into the nucleus but loosely in the sub-
acrosomal space (note: the clear membranes surrounding acrosome and nucleus but not the rod; granular
material between acrosomal and plasma membranes (arrowheads); putative nuclear 'pore' (large arrow)); 4d,
axoneme with small deltoid bodies (small arrows) and central rod (large arrows) (note also mitochondrial
derivatives with distinct membrane and internal structure). Scale bar: 4a = 0.66 \Lm; 4b,c = 0.25 ^m; 4d =
0.2 (Jim.

40

Journal of Hymenoitera Research

Volume 8, Number 1, 1999

41

tozoa (Quicke et al. 1992), although the lat-
ter structures are larger and have a sub-
layered appearance in at least some taxa.
In longitudinal section (Fig. 3), the ac-
rosome of Tremex can be seen to have a
large sub-acrosomal space (Fig. 3a) which
is partly occupied by the acrosomal rod.
This rod extends into the nucleus for al-
most the same length again as it does into
the acrosome. Unlike the nucleus and ac-
rosome, the rod is not membrane bound,
but where the rod is inserted into the nu-
cleus there is no surrounding space, giv-
ing the impression that the rod is being
held by the nucleus. The plasma mem-
brane surrounding the acrosome extends
slightly anteriorly to produce a small ex-
tra-acrosomal space. A membrane bound
area found within the nucleus (Fig. 4c, ar-
row) may represent a longitudinally run-
ning pore. The nucleus (Fig. 3b) is abutted
posteriorly by the axoneme at the level of
the latter's basal body, where the axoneme
lacks the central pair of microtubules. A
large centriolar adjunct is present, and this
in turn contacts the mitochondrial deriv-
atives which have very clearly defined
membrane bound cristae (Fig. 3b, arrow).
There is at least one membrane separating
the centriolar adjunct from the nuclear
membrane. The exact arrangement of the
centriolar adjunct and nucleus, and in par-
ticular, how the centriolar adjunct contacts
the nucleus, is not always obvious. In
many insect spermatozoa this has given
rise to confused interpretations of the

structure, even to suggestions that the cen-
triolar adjunct is absent. From the present
study, the relationship becomes clear at
higher magnification where several sper-
matozoa lie in close proximity (Fig. 4). The
centriolar adjunct (Fig. 4a, arrows) contacts
two equally sized mitochondrial deriva-
tives, and can be seen to extend some way
in between the two, forming what appear
in transverse sections, tail-like structures
(Fig. 4a, large arrow). The mitochondrial
derivatives are thus intimately connected
to the centriolar adjunct. In transverse sec-
tion, certainly at the level of the basal
body, this produces an arrangement
where the centriolar adjunct and basal
body occupy most of the area of section;
an appearance which could be mistakenly
interpreted as there being an extension of
the nucleus overlying the axoneme (Fig.
4b).

Although the acrosome is smaller in di-
ameter than the nucleus, there is not a
great discrepancy. The axoneme itself has
the 9-1-9+2 arrangement (Fig. 4d) common
to Hymenoptera; 9 outer single accessory
tubules, 9 doublets and 2 central single
microtubules. Intertubular material is
abundant with radial spokes (Afzelius
rays) and indications of the inner and out-
er dynein arms. Two deltoid bodies, (also
referred to as triangular rods, (Lensky et
al. 1979) are present, but they are not
large. Between the two bodies, and the
two mitochondrial derivatives (Fig. 4d) is
a single central rod, as previously report-

Fig. 5. Cell ultrastructure of Cqilmlcia sperm (Pamphiloidea): 5a, sperniatadesmata with surrounding cap
material (arrowed); 5b, centriolar adjunct (small arrow) abuts nucleus at membranous complex, extending
beyond the level of the basal body (arrowhead) to abut a mitochondrial derivative (large arrow); 5c, showing
that the centriolar adjunct does not overly both mitochondrial derivatives with one of the two derivatives
(arrowed) abutting the nucleus at the region of the membranous complex; 5d, transverse section at level of
basal body showing that the centriolar adjunct extends to partially enclose the parallel mitochondrial deriv-
ative (arrowed); 5e, transverse section through axoneme showing only a single mitochondrial derivative pos-
teriorly near the tail piece; 5f, transverse section through midregion of axoneme with two mitochondrial
derivatives (note distinct sub-structure at periphery of each mitochondrial derivative); 5g, transverse sections
showing acrosomal rod (r) insertions into the nucleus; 5h, transverse sections showing insertion of acrosomal
rod (r) into the acrosome. Scale bar: a = 1.3 (xm; b = 0.5 |xm; c, d = 0.42 (xm; e = 0.3 (xm; f = 0.27 (xm; g =
0.7 |xm; h = 0.57 (xm.

42

Journal of Hymenoptera Research

Fig. 6. bpLTiiijliitodcsmaUi ol Xi/cUi (Xyeloidea): 6a, showing the extended length o( Uic spcinialodL'.siiuU.il
cap; 6b, showing the distinctive arrangement of the acrosome at the region of insertion into the cap with
particulate material (small arrows), distinct periodicity in the core material (arrowheads; appears like longi-
tudinal striations), and multilayered membrane coat to the acrosome (large arrows). Scale bar: a = 2.5 jjim; b
= 0.66 |xm.

Volume 8, Number 1, 1999

43

ed in ant spermatozoa (Wheeler et al.
1990). A tail piece is found where the ax-
oneme has no associated mitochondrial
derivatives.

Of the more basally derived sawfly su-
perfamilies investigated, the Pamphilo-
idea (Cephalcia) represents a slightly more
advanced evolutionary lineage than the
Xyeloidea although it was once included
in the same family. The spermatozoa of
Cephalcia are arranged in spermatodes-
mata (Fig. 5a) and have heads (nucleus
plus acrosome) approximately 28fJLm long,
and tail, 75|xm long. Ultrastructurally,
they are very similar to those of Tremex,
especially in terms of the size of the ac-
rosomal rod, its position within the sub-
acrosomal space (Fig. 5h), and its insertion
into the nucleus (Fig. 5g). The most no-
ticeable difference between the two is in
the position of the centriolar adjunct. In
Cephalcia the centriolar adjianct can be seen
to run parallel to one to the pair of mito-
chondrial derivatives (Fig. 5b), rather than
overlying both as in Tremex, as is evident
in the region where one of the mitochon-
drial derivatives is found to abut the nu-
cleus (Fig. 5c, arroic). For part of its length
(at the level of the basal body) the centrio-
lar adjunct contacts and even partially en-
closes the single mitochondrial derivative
that lies parallel to it (Fig. 5d, large arrow).
Possibly as a result of this arrangement, a
region occurs at the posterior part of the
sperm, where there is only a single mito-
chondrial derivative lying next to the ax-
oneme (Fig 5e); here there is also only a
single deltoid body, as opposed to the two
found in normal section (Fig. 5f). Similarly
at the level of the centriolar adjunct the
single mitochondrial derivative has only a
single deltoid body. The axoneme is again
similar to that of Tremex in the arrange-
ment of elements. The ray material is par-
ticularly evident, with Afzelius rays hav-
ing distinct spoke heads, and with distinct
electron-opaque granules between the pe-
ripheral singlets (Bairati and Baccetti
1965).

Xyela spermatozoa, although present in
spermatodesmata (Fig. 6), differ in a num-
ber of ways from those of both other saw-
flies described here as well as from the
tenthredinoids described by Quicke et al.
(1992). The spermatozoa are extremely
long with the head ( = nucleus plus acro-
some) being approximately 60^,m long,
and the tail 150 [xm long (Fig. 6a). This
elongation compared with sperm of other
sawflies, at all levels, viz. the acrosome,
nucleus and tail, is also apparent in lon-
gitudinal section. At the anterior end the
acrosomes can be seen to be asymmetrical
and pointed, containing two types of ma-
terial: an irregularly granular material and
a core material that has an almost crystal-
line periodicity, aligned parallel to the
long axis of the spermatozoa. In trans-
verse section (Fig. 7) the most prominent
of the features is an enlargement of one of
the mitochondrial derivatives to a diame-
ter greater than that of the axoneme (Fig.
7a), with a concomitant enlargement of
that mitochondrial derivative's deltoid
body (Fig. 7a, right arrowhead). This dis-
places the other mitochondrial derivative
which, together with its deltoid body, now
occupies an area approximately equiva-
lent to the other, larger, deltoid body
alone.

In Xyela, it is not immediately apparent
if there is a centriolar adjunct. In some
transverse sections, at the position of the
smaller mitochondrial derivative /deltoid
body a darker structure is present (Fig. 7b,
arrows). This does not seem to be simply
a denser mitochondrial derivative because
it lacks a deltoid body and generally the
two occur together (in shape it is actually
closer to a deltoid body). In longitudinal
section (Fig. 7c) a structure abutting on to
the smaller mitochondrial derivative can
be found. This closely resembles the situ-
ation in Cephalcia. The structure does not
however extend to the nucleus like the
other centriolar adjuncts found. Instead, at
the region of the basal body, identifiable
by the absence of the central pair of mi-

44

Journal of Hymenoptera Research

Fig. 7. Organelles of Xycla spermatozoa (Xyeloidea): 7a, asymmetric mitochondrial derivatives (arrows) and
well-devloped deltoid bodies (arrovi'heads); 7b, in one region the smaller mitochondrial derivative is replaced
by a darker structure similar to a centriolar adjunct (arrows) (note also the numerous, smaller nuclear-like
cross sections, smaller than other readily identified nuclei); 7c, longitudinal section showing a centriolar ad-
junct-like body (arrow) that abuts the smaller mitochondrial derivative; 7d, transverse section at the level of
the basal body (arrows) showing that the centriolar adjunct-like organelle and the smaller of the two mito-
chondrial derivatives are both absent, and that the larger mitochondrial derivative (m) partiallv co\ers the
basal body; 7e, showing asymmetric insertion of a cone of acrosomal (a) material into ihe nucleus isolating a

Volume 8, Number 1, 1999

45

crotubules, the larger mitochondrial deriv-
ative wraps round to partially enclose the
basal body and the smaller mitochondrial
derivative /deltoid body is absent (Fig.
7d). If there is a centriolar adjunct, then
how it terminates anteriorly and its rela-
tionship with the nucleus remains unclear.
The nucleus itself appears similar to those
of the other sawflies in density and mem-
brane organisation at the level of the sper-
matodesmata. However, perhaps in ac-
cord with its greater length, there appears
to be an area, posterior to its insertion into
the cap of the spermatodesmata, that has
a relatively smaller diameter and where it
is significantly smaller than the tail region
with its enlarged mitochondrial derivative
(Fig. 7b, d).

In Xyela the interface of the acrosome
with the nucleus also appears different.
There is no discernible rod. Instead acro-
somes, which have a distinct, paracrysta-
line substructure, contact the nucleus and
may even be partially enclosed by it (Fig.
7e). This insertion is displaced to one side,
and this asymmetry is also present in the
acrosome itself (Fig. 7f). A ridge runs
down one side of the acrosome (Fig. 7f,
iirwzv). Spermatozoa are orientated within
the spermatodesmata so that the ridges all
point in the same direction. Interestingly,
this is also the same side of the sperma-
tozoon that the acrosome inserts into the
anterior of the nucleus, although at this
point the acrosomal material appears to
have lost the ridge, and the acrosome at
this level only shows the core of 'periodic
material'. The ridge itself contains the par-
ticulate matter. In some areas the granules
surround membranes resembling the mul-
tilayered coated complex that surrounds
the acrosome itself. There is also an exten-
sion of the outer layers of this coat to form

a small ridge to one side of the acrosome.
This position of this smaller ridge is again
consistent amongst the spermatozoa.

DISCUSSION

At least with regard to the ultrastruc-
ture of the spermatozoa, Cephalcia appears
to have a better claim than Xyela as having
sperm that may be representative of a
common ancestral form. Cephalcia sperm
are very similar to those of Tremex. It is
mainly in the positioning of the centriolar
adjunct that Cephalcia varies from Tremex,
having an asymmetric location overlying
only one mitochondrial derivative. Tremex
by comparison has a centriolar adjunct
overlying both mitochondrial derivatives.
The arrangement of the centriolar adjunct
might seem to have possible usage as a
phylogenetic indicator. Unfortunately the
arrangement of this organelle has often
been poorly understood (e.g. Wilkes and
Lee 1965), and so it is difficult to draw any
conclusion from all previously reported
works. From studies of bee sperm Jamie-
son (1987) concluded that in bees the cen-
triolar adjunct also lies between the nucle-
us and one of the mitochondrial deriva-
tives. Recently we have described the ul-
trastructure of the parasitic braconid
wasp, Aleiodes, which appears to have rel-
atively underived sperm (Quicke et al.
1992; Newman and Quicke 1998). The ul-
trastructure of individual Aleiodes sper-
matozoa closely resembles both Tremex
and Cephalcia. The mitochondrial deriva-
tives are similarly sized and the acrosomal
rod is similarly positioned. The centriolar
adjunct is, however, asymmetric and
hence similar to Cephalcia. Given the prim-
itive status proposed for Cephalcia this
might be considered to be the archetype
arrangement retained through evolution.

small amount of membrane bound nuclear material (arrowed); 7(, showing the periodic appearance of the
acrosome within the multilayered coat and membrane material (arrowed). Scale bar: a = 0.27 jjim; b, c = 0.5
Jim; d = 0.6 jjim; e = 0.5 |j.m; f = 0.25 p.m.

46

Journal of Hymenoptera Research

The case of Tremex, with symmetric cen-
triolar adjunct overlying both mitochon-
drial derivatives, would have to be con-
sidered as an apomorphic development.
However, it is not clear at present which
of the arrangements of the centriolar ad-
junct represents the groundplan for either
the Hymenoptera as a whole or for any of
the major lineages within it, and more
careful study is necessary.

As was made clear in the results, the po-
sitioning of the centriolar adjunct is not
clear in Xyela. It appears to be asymmetric,
but unlike the arrangement in other saw-
flies it does not appear to abut the nucle-
us. Xyela has sperm with a structure that
is extremely divergent in a number of oth-
er ways; the shape and arrangement of the
acrosome, the apparent absence of the ac-
rosomal rod and the size difference in the
two mitochondrial derivatives. It seems
likely that these must represent a response
to selective pressures subsequent to the di-
vergence of both the other sawflies and
the main body of the order of Hymenop-
tera.

The presence of an asymmetric centrio-
lar adjunct in Cqjhalcia appears to cause
the mitochondrial derivatives to be offset
longitudinally, and this may explain why
some sections through the posterior part
of the spermatozoa have only a single mi-
tochondrial derivative (e.g. Fig. 5e). Where
there is only one mitochondrial derivative
the deltoid body is also absent suggesting
they may be a good marker for mitochon-
drial derivative identification.

From the present observations, it seems
probable that the identification of the coat
material surrounding the acrosome in
many taxa may be incorrect. In the Hy-
menoptera, this material has been referred
to as extracellular matrix (Quicke et al.
1992) and in some insect orders (e.g. Or-
thoptera) it has been reported that extra-
cellular matrix granules accumulate
around the plasma membrane of the ac-
rosome to form an extracellular cap (Szol-
losi 1974). However, these structures are

often highly complex, with layered or re-
peated substructure (see for example. Fig.
6b in Quicke et al. 1992), and it is not im-
mediately clear how such a structure
could be secreted if extracellular; the pos-
sibility that they are produced by the ep-
ithelia of a deferent duct cannot be ex-
cluded. Many plasma membranes possess
a glycocalyx which comprises the carbo-
hydrate portion of integral membrane gly-
coproteins and glycolipids together with
associated glycosamminoglycans and pro-
teoglycans, and these carbohydrates ex-
tend from the plasma membrane into the
extracellular space. Where organelle mem-
branes become glycosilated, as in the case
of secretory granules that will eventually
fuse with a plasma membrane, the coated
face of the membrane that opposes the in-
terior of the organelle is the one that will,
upon fusion, face the extracellular space.
In the sawflies investigated here, it is clear
that the coat lies between two membranes,
and is not extracellular as previously re-
ported. An intracellular origin for this
structure would at least allow a more con-
ventional, although as yet, completely un-
recognised, mechanism for its production.

ACKNOWLEDGMENTS

The authors gratefully acknowledge the cheerful
and patient assistance of Ian Morris (EM Unit, De-
partment of Biology). A. Battisti kindly sent us the
live Ctyhalcin, and Paul Johnson, the Tremex. This re-
search was supported by the NERC (Natural Envi-
ronment Research Council) Initiative in Taxonomy.

LITERATURE CITED

Bairati, A. Jr. and B. Baccetti. 1963. Indagini compar-
ative sull'ultrastruttura delle cellule germinali
maschili in Dacuf oleae Gmel. ed in Dm^cpliihi me-
laiwgn^ter Meig. (Ins. Diptera). 11. Nuovi reperti
ultrastrutturali sul filamento assile degli sper-
matozoi. Redia 49: 81-85.

Dallai, R. and B. A. Afzelius. 1990. Microtubule di-
versity in insect spermatozoa: results obtained
with a new fixative. Icurmil of StructKni! Biolcgti
103: 164-179.

Gibson, G. A. P. 1993. Groundplan structure and ho-
mology of the pleuron in H\nienoptera based on
a comparison of the skeletonuisculaturo of \ve-
lidae (Hymenoptera) and raphiliidae (Neurop-

Volume 8, Number 1, 1999

47

tera). Memoirs of the Entomological Society of Can-
ada 165: 165-187.

Heraty, J. M., J. B. Woolley and D. C. Darling. 1994.
Phylogenetic implications of the mesofurca and
mesopostnotum in Hymenoptera. Journal of Hy-
menoptcra Research 3: 241-277.

Jamieson, B. G. M. 1987. The Ultrastructure ami Pliy-
logeny of Insect Spermatozoa. Cambridge Univer-
sity Press, Cambridge, 320pp.

Lensky, Y., E. Ben-David and H. Schindler. 1979. Ul-
trastructure of the spermatozoan of the mature
drone honeybee, journal of Apiculture Research 18:
264-271,

Newman, T. N. and D. L. J. Quicke. 1998. Sperm de-
velopment in the imaginal testes of Aleiodes cox-
alis (Hymenoptera: Braconidae: Rogadinae). Jour-
nal of Hymenoptera Research

Phillips, D. M. 1970. hisect sperm: their structure and
morphogenesis. Journal of Cell Biology 44: 243-
277.

Quicke, D. L. J. 1997. Parasitic Wasps. Chapman &
Hall, London, 470pp.

Quicke, D. L. J., S. N. Ingram, H. S. Baillie and P. V.
Gaitens. 1992. Sperm structure and ultrastructure
in the Hymenoptera (Insecta). Zoologica Scripta
21: 381-102.

Rasnitsyn, A. P. 1980. Origin and evolution of Hy-
menoptera. Trudy Paleontolagicheskogo Instituta
174: 1-190.

Rasnitsyn, A. P. 1988. An outline of evolution of the
hymenopterous insects (Order Vespida). Oriental
Insects 22: 115-145.

Szollosi, A. 1974. Ultrastructural study of the sper-
matodesm of Locusta migratoria migratorouies
(R.F): acrosome and cap formation. Acrida 3: 175-
192.

Vilhelmsen, L. 1997. The phytogeny of lower Hyme-
noptera (Insecta), with a summary of the early
evolutionary history of the order, lournat of Zoo-
logical Systematics and Evolutionary Research 35:
49-70.

Watrous L. E. and Q. D. Wheeler. 1981. The outgroup
comparison method of character analysis. Sys-
tematic Zoology 30: 1-11.

Wheeler, D. E., E. G. Crichton and P. H. Krutzsch.
1990. Comparative ultrastructure of ant sperma-
tozoa (Formicidae: Hymenoptera). Journal of Mor-
pholog}/ 206: 343-350.

Wheeler, D. E. and P. H. Krutzsch. 1992. Internal Re-
productive system in adult males of the genus
CampHvu^tus (hymenoptera: Formicidae: Formici-
nae). Journal of Morphology 211: 307-317.

Whiting, M. F., J. M., Carpenter, Q. D. Wheeler and
W. C. Wheeler. 1997. The Strepsiptera problem:
Phylogeny of the holometabolous insect orders
inferred from 185 and 285 ribosomal DNA se-
quences and morpholog\'. Systenuitic Biologi/ 46:
1-68.

Wilkes, A. and P. E. Lee. 1965. The ultrastructure of
dimorphic spermatozoa in the hymenopteron
DahWomnius fuscipennts (Zett.) (Eulophidae). Gi-
nadian Journal of Genetics and Cytology 7: 609-619.

Yeates, D. K. 1995. Groundplans and exemplars:
Paths to the tree of life. Cladistics 11: 343-357.

J. HYM. RES.
Vol. 8(1), 1999, pp. 48-64

A Review of the Old World Genus Fopius Wharton (Hymenoptera:

Braconidae: Opiinae), with Description of Two New Species Reared

from Fruit-infesting Tephritidae (Diptera)

R. A. Wharton
Department of Entomology, Texas A&M University, College Station, Texas 77843

Abstract. — Two new species of the Old World genus Fopius are described: ceratitivonis from
Kenya and schlingeri from Queensland, Australia. Both species were reared from fruit-infesting
Tephritidae; ceratitivorus from Ceratitis and schlingeri from Bactrocera. Details are provided on dif-
ferentiation of the known species of Fopius, with discussion of their hosts, host specificity, and
distribution. The parasitoids of fruit-infesting tephritids from Kenya are closely related to those
from Madagascar.

The most recent comprehensive classi-
fication of the Opiinae is the three-volume
monograph published by Fischer (1972,
1977, 1987). This collective work estab-
lished a basis for more intense scrutiny of
the Opiinae, resulting in several subse-
quent modifications and additions to the
classification, including the description of
the Old World genus Fopius (Wharton
1987, van Achterberg and Maeto 1990).
The numerous name changes affecting
opiine parasitoids of fruit-infesting Te-
phritidae were recently reviewed by
Wharton (1997b), who also provided sug-
gestions for delineation of species groups
within the genus Fopius. Additional
changes in nomenclature, some of these
affecting opiine parasitoids of tephritids,
were published by van Achterberg and
Salvo (1997) and Quicke et al. (1997). Keys
to most of the species of Fopius can be
found in Wharton and Gilstrap (1983) and
Fischer (1987), with both works treating
the species under the generic name Bios-
teres. Palacio et al. (1992) provide addi-
tional information on separation of males
and immatures of two sympatric species.

All opiine braconids reared to date are
koinobiont endoparasitoids of cyclorrha-

phous Diptera, and all emerge from the
puparium of their hosts. Hosts are known
for one-third of the approximately 1500
described species, with most of these rec-
ords pertaining to Agromyzidae and Te-
phritidae. All reared species of Fopius are
parasitoids of Tephritidae. Summaries of
the literature on hosts and biology of the
opiine parasitoids of Tephritidae can be
found in Fischer (1972, 1977, 1987), Clau-
sen (1978), Wharton and Marsh (1978),
Wharton and Gilstrap (1983), Gilstrap and
Hart (1987), Wharton (1989, 1997a, b).
Messing (1996), Sivinski (1996), and Siv-
inski et al. (1997).

The primary purpose of the work pre-
sented here is to facilitate on-going studies
in biological control by providing names
for two recently discovered, undescribed
species. Both species are of interest with
respect to tephritid biological control be-
cause of their potential for attacking eggs
or early instars, and one of these is a na-
tive parasitoid of the Mediterranean fruit
fly, Ceratitis capitata (Wiedemann). The use
of parasitic Hymenoptera for the biologi-
cal control of tephritid pests has received
considerable attention in recent years
(Knipling 1992, Waterhouse 1993, Head-

Volume 8, Number 1, 1999

49

rick and Goeden 1996, Purcell, 1998), and
there are active programs currently un-
derway in several countries.

MATERIALS AND METHODS

With the exception of C. Granger's type
material from the Paris Museum and a
single, swept specimen of Fopiiis schliiigeri,
n. sp., all material used in the descriptions
of the new species was reared from fruit
in association with various fruit-infesting
Tephritidae. In some cases, parasitoids
were reared from bulk fruit samples, with
unconfirmed host associations. Most of
the material, however, was reared from
isolated puparia. In several of the rearings
for Fopius cerntitivorus, n. sp., puparia
were individually isolated prior to emer-
gence. Though this procedure decreases
the percent emergence (due primarily to
desiccation and /or physical damage), it
enables correct association of the wasp
with the host from which it was reared.

Specimens of the newly described spe-
cies have been deposited in the following
institutions: University of Queensland,
Brisbane (UQBA), Australian National In-
sect Collection, Canberra (ANIC), Texas
A&M University, College Station
(TAMU), Bernice P. Bishop Museum,
Honolulu (BPBM), Hawaii Department of
Agriculture, Honolulu (HDA), Queens-
land Department of Primary Industries,
Indooroopilly (QDPI), Nationaal Natu-
urhistorisch Museum, Leiden (RMNH),
The Natural History Museum, London
(BMNH), National Museum of Kenya,
Nairobi, International Centre of Insect
Physiology and Ecology, Nairobi (ICIPE),
and U.S. National Museum of Natural
History, Washington, D. C. (USNM).

Descriptive terminology follows Whar-
ton (1987, 1988, 1997b) and Sharkey and
Wharton (1997), and is based largely on
the works of Fischer (1972). A tabular
summary is presented rather than a di-
chotomous key to facilitate assessment of
relationships and point out gaps in our
knowledge.

IDENTIFICATION, RELATIONSHIPS,

HOSTS, AND DISTRIBUTION

PATTERNS

The tephritid parasitoids in the genus
Fopius are readily distinguished from oth-
er opiines by the possession of crenulate
notauli extending posteriorly to the me-
sonotal midpit, an oblique ridge ventral-
laterally on the propleuron, a short second
submarginal cell (3RSa < 2RS), and a long
ovipositor (1.3-4.5 X longer than meso-
soma). Character states useful for identi-
fying species with known tephritid host
records are provided in Table 1, and the
characters themselves are discussed be-
low. Eight closely related species for
which host records are lacking have also
been included in the table. The table is
deemed more informative than a dichot-
omous key because it provides prelimi-
nary data for phylogenetic analysis, which
is beyond the scope and purpose of the
present work, as well as providing sup-
plemental characters for assistance in
identification. More than half of the spe-
cies in Table 1 (including several of the
tephritid parasitoids) are known only
from the type or the type plus a few other
specimens. Additional collecting is essen-
tial before progress can be made in our
understanding of these species. Individual
species and species groups are treated fur-
ther following discussion of the charac-
ters.

Cliaracter 1. — Striate sculpture on the sec-
ond and third metasomal terga. 0 = stri-
ae absent; 1 = striae present on tergum
2 only; 2 = striae present on tergum 2
and at least base of tergum 3. For most
species, assignment of either character
state 0 or character state 1 is unambig-
uous. Fopius deeralensis (Fullaway), how-
ever, has weak striae on tergum 2, and
the sculpture is not always readily vis-
ible (see diagnosis following description
of F. schliugeri, n. sp.). Fopius skiniwri
(Fullaway), from the Philippines, is the
only species in which striae are usually

50

Journal of Hymenoptera Research

Table 1. Matrix of coded character states for species in the genus Fopntis (see text for character definition).

1

:

T

4

-

h

-

s

-

111

11

i:

p.

14

dcnticidifcr (van Achterberg & Maeto)

0

0

0

2

5

1

0

0

3

1

0

4

4.1-4.6

0

nwraiigensis (Fischer)

0

0

0

2

5

1

0

0

2b

1

7

4

1.7

7

taiwanicus (Fischer)

0

0

0

2

5

1

0

0

2b

1

7

4

-3.0

0

ruficornis (Granger)

0

1

0

1,2

3

1

0

0

1

2

0

2

1.5

0

ruhrithorax (Granger)

0

1

0

1

2

0

1

1

0

1

0

1.3

1

bei'isi (Brues)

0

0

1

1

0

1

0

0

1

0

2.2-2.3

1

dcsideratui (Bridwell)

0

0

1

1

0

1

0

0,1

1

0

3.0-3.2

1

uigcr (Szepligeti)

0

0

1

1

0

1

0

1

0,1

0

2.6-2.7

1

otfohvnoaiius (Fullaway)

0

0

1

I

0

1

0

1

1

0

2.8

1

rufotcftnccuf (Granger)

0

0

1

1

0

i

1

0

0,1

0

2.0-2.3

1

altcrnatae (Tobias)

0

0

0

0

0

7

0

3

1

7

2

-1.25

7

arisiinus (Sonan)

0

0

0

0

0

0

0

2

0,1

0

3

2.5-2.8

0

caiyomyiae (Silvestri)

0

0

0

0

0

0

0

2a

0

1

2

1.5-1.7

0

myolejae (Tobias)

0

0

0

0

0

7

0

3

1

7

2

-2.0

7

pcrsulcatus (Silvestri)

0

0

0

0

7

0

0

2a

0

7

3

7

7

skinneri (Fullaway)

0

0

0

0

0

0

0

2b

1

0

3?

-2.65

0

vniuienhoschi (Fullaway)

0

0

0

0

0

0

0

2b

0,1

0

2

2.5-2.7

0

ceraiitivorus n. sp.

0

0

0

0

5

0

0

0

0

0

0,1

2

1.7-1.9

0

longicnuda (Granger)

0

0

0

0

0

0

0

0

1

0

0

2

2.6

1

pykucthcrax (Fischer)

0

0

0?

7

7

■?

7

0,1

0

1

7

7

7

7

silivftrii (Wharton)

0

0

0

0

0

0

0

0

0

1

0

2

2.5-2.6

0

decralensis (Fullaway)

1

0

0

0,2

5

0

0

0

4

0

0

1

3

1

schlingeri n. sp.

0

0

0

0

5

0

0

0

4

0

0

3

2.3-2.5

1

caudatus (Szepligeti)

0

0

0

1

4

0

0

0,1

0

1

1

3

7

1

present on tergum 3, though the sculp-
turing on tergum 3 is usually not exten-
sive.

Character 2. — Occipital carina. 0 = present
laterally; 1 = completely absent. The oc-
cipital carina is absent mid-dorsally in
Fopius, but present laterally in nearly all
species. There is also some variation
among species in the height of the oc-
cipital carina, as exemplified by the two
species described below, and this vari-
ation may eventually prove useful in
demonstrating character state transfor-
mations leading to complete loss of the
occipital carina.

Character 3. — Setal pattern on the oviposi-
tor sheath. 0 = two or more rows of
densely spaced setae; 1 = setae sparse,
at most with a row of long, moderately
sparse setae basally and short, widely
spaced setae apically. This coding is
useful for segregating groups of species,
but oversimplifies the complexity of the
character states that may eventually be

useful for delineation of additional spe-
cies. Setal rows are difficult to count,
however, and many of the specimens
examined were in such poor condition
that it could not be determined if setae
were sparsely arranged or merely bro-
ken off. Of the species coded 0 in Table
1, setal density was greatest in F. detiti-
culifer (van Achterberg and Maeto) and
least in F. schlingeri, n. sp. and F. ruhri-
thorax (Granger).
Character 4. — Ventral margin of clypeus. 0
= thin, sharp, and evenly convex, with-
out median projection; 1 = somewhat
thickened medially, and slightly pro-
truding, with labrum sometimes par-
tially exposed; 2 = with median, ven-
trally-directed, tooth-like (i. e. pointed)
projection (clypeus completely occludes
labrum). Differences between states 0
and 1 may not be apparent without dis-
section to reveal the thickened margin.
The tooth-like projection is very small in
deeralensis, and the margin thinner than

Volume 8, Number 1, 1999

51

in members of the F. marangensis (Fi-
scher) species group, as reflected by its
coding in Table 1.

Character 5. — Pattern of sculpture and se-
tae on frons. 0 = densely setose and
punctate, the punctures tending to coa-
lesce to some degree, giving the appear-
ance of transversely rugulose lines (Fig.
4), midline longitudinally rugose; 1 =
transversely striate and impunctate over
middle half of frons, with deep, widely
spaced punctures laterally; 2 = laterally
as in state 1, but largely unsculptured
medially (at most with a few, irregular,
very weak wrinkles), midline with
sharp carina basally; 3 = smooth, im-
punctate, depressed along midline; 4 =
broad, transverse band of deep punc-
tures extending from ocelli to eye, oth-
erwise smooth; 5 = densely setose and
punctate, the punctures discrete, with
no indication of rugosities as in state 0;
for state 5, the punctures are very
densely spaced in marangensis, F. taiivan-
icus (Fischer) and denticulifer, less so in
deeralensis and schlingeri, and least in
ceratitivorus (where they are virtually
absent basal-laterally).

Character 6. — Postpectal carina. 0 = well
developed; 1 = weak to absent.

Character 7. — Relative length of first two
flagellomeres. 0 = first flagellomere
about same length as second (ratio vary-
ing from 0.9-1.1); 1 = first flagellomere
distinctly shorter than second (0.8 X
length or less).

Character 8. — Shape of petiole. 0 = petiole
length equal to or shorter than apical
width, strongly widening apically; 1 =
petiole appearing more parallel-sided,
with length distinctly greater (at least
1.3 times) than apical width. The petiole
of f. caiidatus (Szepligeti) is somewhat
intermediate, as reflected by its coding
in Table 1. The petiole is not necessarily
more parallel-sided in state 1 than in
state 0 (width at apex may be twice
width at base in both), but appears to

be so because the petiole is longer in
state 1.

Character 9. — Geographic distribution. 0 =
continental Africa; 1 = Madagascar; 2 =
southern Asia (2a = India; 2b = south-
east Asia, including Indonesia, Philip-
pines, and Taiwan); 3 = Japan, eastern
Russia; 4 = northeastern Australia. Dis-
tribution patterns given here do not re-
flect the successful introductions of F.
arisaniis (Sonan) and F. vaiuienboschi
(Fullaway) to Hawaii and arisanus to
Central America.

Character 10. — Color of mesosoma. 0 =
largely pale (red, orange, yellow, or
brownish-white); 1 = dark black to
brown; 2 = pale with large black spots
on mesoscutum and mesopleuron. As-
sessment of coloration is somewhat
problematic due to postmortem chang-
es, especially in shades of red, yellow,
and orange. Also, there is almost a com-
plete continuum in shades of red from
pale through nearly black (skinneri is
dark reddish-brown). Two of the spe-
cies for which there is abundant mate-
rial (e. g. arisanus and vandenboschi) are
color-variable.

Character 11. — Dorsal carinae of petiole. 0
= dorsal carinae extending posteriorly
beyond spiracle for at least a short dis-
tance as a distinctly elevated ridge; 1 =
dorsal carinae not extending past spi-
racle as a distinctly elevated ridge. At
least three species are variable in this
feature, as reflected by the coding in Ta-
ble 1.

Character 12. — Configuration of ovipositor
tip. 0 = distinct double node dorsally; 1
= weak node or swelling dorsally; 2 =
parallel-sided at apex, with little or no
node; 3 = strongly tapered apically to a
fine, smooth point, narrowest subapi-
cally; 4 = strongly tapered apically as in
state 3, but with tip flattened dorsal-
ventrally. States 1 and 2 merely repre-
sent different degrees of development of
a transverse ridge near the tip of the
ovipositor; and states 3 and 4 represent

52

Journal of Hymenoptera Research

conditions for which it is hypothesized
here that nodes and /or transverse ridg-
es have been lost. Detailed SEM work is
still needed to elucidate these character
states for many of the species.

Character 13. — Approximate ovipositor
length. Values given are total ovipositor
length divided by length of mesosoma.
Accurate measurement of ovipositor
length often requires dissection, which
was not possible for some of the species.

Character 14. — Mesopleural setae. 0 = at
least some setae present on mesopleu-
ron dorsal to the speculum (the dorsal-
posterior section of the mesopleuron); 1
= setae completely absent above spec-
ulum.

Table 1 has been arranged to facilitate
identification of both species and species
groups. Several of the species groups are
quite distinctive and thus readily recog-
nizable (Wharton 1997b), and these will be
treated first in the following discussion.
Since the focus of this paper is on parasit-
oids of fruit-infesting Tephritidae, F. riifi-
cornis (Granger) and the marangensis spe-
cies group are not further discussed be-
cause there are no host records and the
species are readily identified using Ta-
ble 1.

The desideratus species group of Fopius
consists of bevisi (Brues), desideratus (Brid-
well), niger (Szepligeti), ottotomoamis (Ful-
laway), and riifotestaceiis (Granger). As
noted by Wharton and Gilstrap (1983), the
species of the desideratus group are very
similar to one another. For example, riifo-
testaceus is virtually identical to bevisi, but
has the mesosoma red rather than yellow
or yellow-orange. Unfortunately, few
specimens have been available for study
of intraspecific variation in the color pat-
terns currently used to differentiate the
species of this group. There are published
host records (summarized by Wharton
and Gilstrap 1983) for all but riifotestaceiis.
Most of the specimens examined were
reared from Dacus infesting Cucurbita-

ceae. Both desideratus and ottototiioauus
have been recorded from undetermined
species of Dacus in cucurbits (Bridwell
1919, Fullaway 1957), and uiger was
reared from D. humeralis Bezzi (Wharton
and Gilstrap 1983). The few remaining
published records (Bridwell 1919, Clausen
et al. 1965) are from Ceratitis aiiouae Gra-
ham on Myriauthus arboreus (specimens of
desideratus) and Trirhithrum queritum Mun-
ro on Stri/chiios usanibarensis (a specimen
tentatively identified as bevisi). All mem-
bers of this species group have large, sub-
apical nodes on the ovipositor. Based on
comparisons of ovipositor morphology
with species of known biology in the re-
lated genus Diachasmimorpha Ashmead, it
is suggested here that members of the de-
sideratus group attack late instar larvae of
their hosts. Members of this group are
known from Cameroon, Kenya, Nigeria,
Tanzania, South Africa, and Madagascar,
and undoubtedly occur throughout sub-
saharan Africa. Fopius rubrithorax (Grang-
er) is very similar to the other species
mentioned here, despite the reduced
sculpture on the frons and a slightly more
setose ovipositor sheath; and I therefore
place it as a basal member of this group.
This placement assumes that both the re-
duced setal pattern on the ovipositor
sheath and the pattern of sculpture on the
frons of the five other species of the desi-
deratus group are derived relative to the
conditions in rubrithorax; this remains to
be tested in a more rigorous fashion.

The persulcatus species group of Fopius,
characterized largely by striate sculpture
on the second metasomal tergum, consists
of altcniatae (Tobias), arisanus (Sonan), car-
ponn/iae (Silvestri), myolejae (Tobias), per-
sulcatus (Silvestri), skinneri (Fullaway), and
vaiulenboschi (Fullaway). The species are
very similar to one another, but differ pri-
marily in coloration, length of ovipositor,
and configuration of the ovipositor tip.
Following their successful introduction to
Hawaii during the biological control pro-
gram against oriental fruit fly (Clausen et

Volume 8, Number 1, 1999

53

al. 1965), arisaniis and vandenboschi were
intensively studied, and much is now
known about their biology (with most of
the early literature on arisaiius published
under the name Opiiis oophihis Fullaway).
In their native range, centered around Ma-
laysia and Indonesia, both are parasitoids
of tephritids in the dacine genus Bactwcera
Macquart. Unlike skinneri, neither is at-
tracted to cucurbit-infesting flies. Other
than the original host records little is
known about the other species in this spe-
cies group, including skinneri. Data on per-
sukaius (type material reared from B. car-
i/eae Kapoor) are particularly problematic
because of widespread confusion regard-
ing the identity of this species during the
Hawaiian oriental fruit fly program, and
the subsequent description of several sub-
species (Fischer 1965). The other species
(viz. carpomi/iae, tm/olejae, and alternatae)
have been reared, respectively, from try-
petine tephritids in the genera Carpomya
Costa, Myoleja Rondani, and Rhagoletis
Loew. Based on the similarities in the
shape of the ovipositor tip, all species in
this species group preferentially attack ei-
ther the egg or early instar larva of their
host (though all eventually emerge from
the puparium). This biology, however, has
only been confirmed for arisanus (attack-
ing eggs) and vandenboschi (attacking pri-
marily first instars). The species of this
group are known from Pakistan and India
east through Indonesia and north through
Taiwan, Japan, and far eastern Russia.

Wharton (1997b) delimited a silvestrii
species group containing longicauda
(Granger), pyknothorax (Fischer), and sil-
vestrii (Wharton). One of the species de-
scribed below, ceratitivorus, also belongs
here. This group is currently defined
largely by the absence of features that de-
fine the three species groups already men-
tioned: the clypeus lacks a median tooth
on the ventral margin, the setae on the
ovipositor sheath are not reduced, and the
second metasomal tergum is unsculp-
tured. Reduction of features on the dorsal

valve of the ovipositor suggests either a
sister group relationship to the persulcatus
species group or a parallel loss relative to
the desideratus species group, but this hy-
pothesis needs to be tested more rigorous-
ly. Both sili>estrii and ceratitivorus have
been reared from ceratitine tephritids in-
festing coffee, and silvestrii has also been
reared from Dacus bivittatus (Bigot) infest-
ing squash (Steck et al. 1986, Wharton
1987). Other members of this group have
not been reared. Members of the silvestrii
species group have much the same distri-
bution pattern as those of the desideratus
species group, and are differentiated from
one another largely by color (silvestrii and
pyknothrorax are dark, longicauda and cer-
atitivorus are pale) and ovipositor length.
The remaining three species, caudatus,
deeralensis, and schlingeri, do not readily
cluster into distinctive species groups.
Identification of deeralensis and schlingeri is
discussed below under the diagnosis fol-
lowing the description of schlingeri; cau-
datus is readily separated from all other
species of Fopius by the distinctive band of
setae and punctures on the frons. Both
deeralensis and schlingeri are from Queens-
land, where (as noted below under the de-
scription of schlingeri and in Clausen et al.
1965) they have been reared from various
species of Bactrocera in a variety of host
fruits. Fopius caudatus has thus far been
reared exclusively from ceratitines (Steck
et al. 1986). It is known from tropical re-
gions of both eastern and western Africa,
where it has been reared from coffee ber-
ries containing the ceratitines Trirhithruin
coffeae Bezzi, C. anonae and C. capitata as
well as from other fruits containing anon-
ae. Specific host records for caudatus need
confirmation, in part because of earlier
confusion regarding its identity (Wharton
1987). This species resembles members of
the desideratus species group in the mor-
phology of the clypeus and petiole, but
has a distinctly different ovipositor
(strongly narrowed, suggesting oviposi-
tion in the host egg) as well as several fea-

54

Journal of Hymenoptera Research

tures unusual for members of the genus
Fopms (Wharton 1997b).

A few generalizations can be made
about hosts and distribution patterns,
even though our current knowledge is
somewhat limited. Rearing records from
within their native ranges (Clausen et al.
1965, Steck et al. 1986) suggest that the 16
species for which we have host records are
restricted to fruit-infesting tephritids, but
that there are different levels of specificity.
Some are currently known only from a
single host, others (e. g. caudatus on cera-
titines) have only been reared from a nar-
row group of hosts, and several have been
reared from hosts in two or three different
tribes. Most of the known hosts belong to
the tribes Ceratitini and Dacini, both in the
tephritid subfamily Dacinae (White and
Elson-Harris 1992). Except where they
have been introduced for biological con-
trol, members of the persukatus group oc-
cur outside the range of fruit-infesting
Ceratitini, and several of them have been
reared from Trypetini in the tephritid sub-
family Trypetinae. Where introduced out-
side their native range for biological con-
trol, arisanus and vandenboschi have been
able to attack other fruit-infesting tephri-
tids (Clausen et al. 1965, Wharton et al.
1981). Yet, while some of the species of
Diachasmimorpha Viereck introduced to
Hawaii to control fruit-infesting Tephriti-
dae occasionally attack gall-making (but
not flower-infesting) Tephritidae, arisanus
and vandenboschi do not (Duan et al. 1996).
The genus Fopius provides evidence for
a close relationship between the fauna of
Madagascar and that of adjacent regions
of continental Africa (as do its host te-
phritids). Although a few of the Madagas-
car elements (notably rubrithorax and es-
pecially ruficornis) are unique in several
respects, both rufotestaceus and longicauda
have their closest known relatives (bevisi
and ceratitivorus respectively) on the adja-
cent mainland.

DESCRIPTIONS

Fopius ceratitivorus Wharton, new
species

(Figs. 1, 2, 7, 8, 10, 11, 13-15, 21)

Female.— Head: 1.55-1.75 (m = 1.65 ±.07)
times broader than long; 1.25-1.35 Hmes
broader than mesoscutum; face distinctly
punctate throughout, pattern variable but
spacing between most punctures about
equal to diameter of punctures; setae short,
somewhat decumbent; midridge low, pol-
ished, more prominent dorsally, extending
between antennal bases (toruli) as a low,
flat ridge; distance between toruli greater
than distance from torulus to eye; frons
longitudinally rugulose along midline,
highly polished and weakly depressed ba-
sally on either side of rugulose midline,
deeply punctate elsewhere, the patch of
punctures on each side anteriorad ocelli
usually more densely spaced, occasionally
with punctures coalescent, ocellar triangle
almost completely margined by a crenulate
sulcus. Occipital carina in lateral view ex-
tending dorsally from base of mandible to
a point just below top of eye. Clypeus in
profile slightly bulging dorsomedially;
ventral margin of clypeus thin and evenly
convex, not thickened medially; setae on
clypeus very sparse, at least twice length of
those in middle of face, weakly directed
ventrally; clypeus completely concealing
labrum when mandibles closed. Eye (at
50x) apparently bare, large, 2.85-3.8
(m=3.2±0.3) times longer temple; temples
very weakly receding in dorsal view; width
of head at temples slightly less than width
at eyes. Antenna 31-37 segmented; roughly
3.0-3.1 times longer than mesosoma; first
flagellomere 0.9-0.95 times length of sec-
ond. Maxillary palps longer than height of
head. Mesosoma: 1.2.5-1.35 (m = 1.3±0.05)
times longer than high, 1.55-1.7
(m = 1.65±0.05) times longer than broad.
Median lobe of mesoscutum with 2 paral-
lel, rugosopunctate, longitudinal grooves
extending more than half length of median
lobe, median lobe otherwise setose, with

Volume 8, Number 1, 1999

55

Figs. 1-6. Heads of Fopniii spp.: 1 and 2, ccratitivonis frons and face; 3, sc/i/diycn face and cljpeus; 4, aiiiHiiHi
frons; 5, i^chlingeri face and clypeus; 6, liccrulciifis face and clypeus (most setae broken), arrow = median
projection on ventral margin of clypeus.

scattered, deep punctures; lateral lobes
bare and impunctate medially, with nu-
merous, relatively long, inwardly directed
setae around margins; notauli broadening
posteriorly, distinctly crenulate through-
out, with pits usually becoming elongate
posteromedially where the ridges between
the pits form a small strigose area; space

between strigose area and scutellar sulcus
with scattered, deep punctures, postero-
median area either broadly and very shal-
lowly depressed or with a shallow, more
discrete midpit. Scutellar sulcus broader
medially than laterally, the posterior mar-
gin with a distinct median excavation;
number of longitudinal ridges in sulcus

56

Journal of Hymenoptera Research

variable. Metanotum with relatively low
median ridge. Propodeum finely, densely
rugose, the sculpture without obvious pat-
tern; elevated median longitudinal carina
usually distinct only on anterior 0.25-0.35;
propodeum laterally not separated from
metapleuron by a well-defined pleural ca-
rina, the demarcation represented only by
the transition to the weakly sculptured
dorsal portion of the metapleuron. Stemau-
lus broad, deep, crenulate throughout, ex-
tending posteriorly roughly 0.7 times dis-
tance from anterior margin of mesopleuron
to mid coxa; crenulate sculpture extending
dorsally along anterior margin of meso-
pleuron through subalar depression; pos-
terior margin crenulate ventrad speculum,
but with unsculptured sulcus dorsally; me-
sopleural disc setose throughout; postpec-
tal carina present, but variously developed.
Wing: SHgma 2.7-2.9 (m=2.85±0.1) times
longer than wide, with r arising slightly
distad its midpoint; 2RS weakly sinuate,
1.2-1.45 (m=1.3±0.05) times longer than
3RSa; 3RSa 1.55-2.5 (m=2.1±0.3) times lon-
ger than r; 3RSb ending slightly but dis-
tinctly anteriorad wing tip; (RS-l-M)a sinu-
ate; (RS-l-]VI)b present, m-cu nearly always
arising distinctly basad 2RS; Icu-a incli-
vous, usually postfurcal relative to IM but
varying from interstitial to postfurcal by
0.4 times its length. Hind wing m-cu recli-
vous, straight or very weakly recurved
near wing margin, extending to wing mar-
gin or nearly so as well-developed, deeply
impressed crease, usually weakly pigment-
ed anteriorly. Metasoma: Petiole 0.95-1.05
(m = 1.0 ±0.05) times longer than apical
width, apex 1.80-2.15 (m=1.95±0.1) Hmes
wider than base; densely and finely striate;
dorsal carinae well-developed over basal
two-thirds, weaker posteriorly but distinct
to posterior margin, carinae very weakly
converging, with distance between carinae
at posterior margin roughly equal to dis-
tance to lateral margin; dorsope present
but not extending basally as a deep pit.
Metasoma unsculptured beyond petiole.
Hypopygium strongly narrowed and

pointed posteriorly but short, not greatly
attenuate. Ovipositor tip weakly narrowed
apically, without distinct dorsal node or ca-
rina but with weak ventral serrations; 1.65-
1.95 (m = 1.8 ±0.1) times longer than meso-
soma; ovipositor sheath densely setose
with multiple rows of at least 30 setae each,
the number of rows difficult to distinguish
because of density of setae; sheath 1.35-
1.55 (m = 1.45 ±0.05) times longer than me-
sosoma. Color: Pale, yellow to orange, the
exact hue dependent largely on manner of
preservation; ovipositor sheath, veins, and
stigma brown; antenna brown, with scape,
pedicel, and basal flagellomeres usually
yellow to orange medially. Wings hyaline.

Male. — As in female except eye 2.9±0.25
times longer than temple; antenna 30-35
segmented; petiole narrowed at apex, 1.1-
1.25 (m = 1.15±0.05) times longer than api-
cal width, apex 1.6-1.95 (m=1.75±0.1)
times wider than base; dorsope less dis-
tinct. Length: 2.0-3.4 (9) and 1.85-2.9 (6)
mm.

Hosts. — This species has been reared
from isolated puparia of fruit-infesting Te-
phritidae attacking coffee in central Ken-
ya. It has also been reared from bulk sam-
ples of coffee. The tephritids from these
samples, in order of abundance, were Cer-
atitis capitata, C. rosa Karsch, and Trirliith-
ruin coffeae. All are members of the tribe
Ceratitini, subtribe Ceratitina.

Material examined. — Holotype female,
"Kenya: Ruiru C. R. F. 17.IX.1996 ex: te-
phritid on coffee berries Ref. No. CB03"
Deposited in Kenya National Museum,
Nairobi. Paratypes (BMNH, BPBM,
RMNH, HDA, ICIPE, TAMU, USNM): 5 9,
76, "Kenya: Nairobi 20.V.1997 ex: Ceratitis
capitata, coffee M. Ramadan & R. Mess-
ing"; 339, 25 9, "Kenya E. Province,
Mbeere Distr Mbeti south Rurima 30.iv.97
ICIPE Fruitfly Project ex Fruittly on Coffee
berries"; 19, 16, "Kenya: Ruiru 15mi
NNE Nairobi 10.iv.l995 ex: coffee No.
CB05 ICIPE Collections"; 19, "Kenya:
Western Prov. Koru iv.l995 ex. Coffea ca-
neophora CAB Collections" and, 1 6

Volume 8, Number 1, 1999

57

Figs. 7-10. fi);iii(s spp.: 7, ccratitiz\^n(^ head, arrows = top of occipital carina ancJ mid-dorsal elevation of
clypeus; 8 and 10, cernlitivonis dorsal view of mesosoma; 9, schliiigeri dorsal view of mesosoma.

58

Journal of Hymenoptera Research

"Kenya: E. Province Mbeere District Rur-
ima Farm 0°38'29"S, 37°29'49"E 3.X.1997 ex
tephritid in coffee Wharton, Kimani, Ov-
erholt." This species is known only from
central Kenya.

Diagnosis. — This species closely resem-
bles longicauda, known only from Mada-
gascar. The two species are similarly col-
ored, have a densely punctate frons,
densely setose ovipositor sheath, and
identical configuration of the clypeus. Fop-
itts ceratitivorus differs from longicauda pri-
marily in the possession of a shorter ovi-
positor (sheaths at least 2 times longer
than mesosoma in longicauda). The median
lobe of the mesoscutum and the junction
of the notauli (Fig. 10) are also more
heavily sculptured in ceratitivorus than in
longicauda, and hind wing m-cu is straight-
er. Both ceratitivorus and longicauda differ
from other Old World species of Fopius ei-
ther in coloration, sculpture of the frons,
length of ovipositor, and /or shape of the
clypeus and its relative degree of conceal-
ment of the labrum. From other orange
opiines reared from tephritids in coffee in
Kenya, ceratitivorus may be readily distin-
guished by the short second submarginal
cell with fore wing m-cu distinctly sepa-
rated from 2RS (Fig. 21) and by the com-
pletely sculptured notauli.

Discussion. — I place ceratitivorus in the
silvestrii species group of Fopius (Wharton
1997b). The ovipositor tip of ceratitivorus,
though narrowed, does not have exactly
the same morphology as found in arisanus
and schlingeri n. sp. Thus, females proba-
bly do not oviposit in host eggs, but based
on the shape of the ovipositor tip, they
may attack early instars. The same may be
true of longicauda, the holotype of which
appears to have a similar ovipositor.

A weak negative correlation was ob-
served between body length and relative
length of the ovipositor, but the sample
size (N = 10) was too small to confirm this
apparent trend.

Fopius schlingeri Wharton, new species

(Figs. 3, 5, 9, 12, 16, 18-20)

Female.— Head: 1.55-1.65 (m = 1.6±.05)
times broader than long; 1.3-1.4
(m = 1.35 ±.05) times broader than meso-
scutum; face distinctly punctate through-
out, pattern variable but spacing between
most punctures distinctly greater than di-
ameter of punctures; setae short, some-
what decumbent; midridge low, polished,
narrower dorsally, extending between to-
ruli; distance between toruli varying from
slightly to distinctly greater than distance
from torulus to eye; frons with polished,
weakly elevated, crenulately margined,
triangular projection extending from me-
dian ocellus at least half distance to toru-
lus; frons otherwise punctate, the punc-
tures anteriorad ocellar field dense, with
spacing between punctures about equal to
diameter of punctures; ocellar triangle
margined at least in part by an impressed
line. Occipital carina in lateral view ex-
tending dorsally from base of mandible to
about middle of eye. Clypeus in profile
weakly to distinctly bulging dorsomedi-
ally; ventral margin of clypeus thin and
weakly but evenly convex, not thickened
medially nor with median projection; se-
tae on clypeus sparse, about twice length
of those on face, weakly directed ventral-
ly; ventral margin of clypeus not suffi-
ciently convex to completely conceal la-
brum when mandibles closed. Eye usually
with 1-4 minute setae visible in dorsal
view, very large, 5.3-7.5 (m = 6.45 ±0.75)
times longer than temple; temples weakly
receding in dorsal view; width of head at
temples about 0.9 times width at eyes. An-
tenna 41-47 segmented; roughly 3.5 times
longer than mesosoma; first flagellomere
equal in length to second. Length of max-
illary palps equal to height of head. Me-
sosoma: 1.25-1.35 (m = 1.3) times longer
than high, 1.75-1.80 times longer than
broad. Median lobe of mesoscutum with 2
parallel, unsculptured, longitudinal
grooves extending more than half length

Volume 8, Number 1, 1999

59

Figs. 11-16. /-iij'/ii-. '-pp.: 11, ccratitworus propodeum; 12, schliiigeri propodeum; 13 and 14, ccrnlili,\'iii> pro-
pleuron, armw = oblique carina; 15, ccriititivoriis petiole; 16, .•ic/i//);,sji'r( petiole.

60

Journal of Hymenoitera Research

Figs. 17 cind 18. Ovipositors of Fopius spp; 17, deernlensis; 18, sir,

rlilingcri.

of median lobe, median lobe otherwise se-
tose with numerous, very fine, widely
spaced punctures; lateral lobes with nu-
merous, relatively long setae around mar-
gins and more sparsely scattered setae me-
dially; notauli distinctly crenulate though-
out, meeting posteriorly in a clearly de-
fined midpit that often extends narrowly
to posterior margin. Scutellar sulcus par-
allel sided or nearly so, usually with 3
well-developed longitudinal carinae plus
several additional weaker ones. Metano-
tum with distinctly elevated median
flange posteriorly. Propodeum densely ru-
gose, the sculpture largely without obvi-
ous pattern though elevated median lon-
gitudinal carina distinct on anterior 0.25,
and posterior 0.25 often with remnants of
the parallel ridges from a median areola;
propodeum laterally not separated from
metapleuron by a well-defined pleural ca-

rina, the demarcation represented only by
the transition to the weakly sculptured
dorsal portion of the metapleuron. Ster-
naulus broad, deep, crenulate throughout,
extending posteriorly roughly 0.7 times
the distance from anterior margin of me-
sopleuron to mid coxa; crenulate sculpture
extending dorsally along anterior margin
of mesopleuron throughout subalar de-
pression; posterior margin crenulate ven-
trad speculum, with unsculptured sulcus
dorsally; mesopleural disc setose; postpec-
tal carina well developed medially. Wing:
Stigma 2.6-3.0 (m = 2.8±0.15) times longer
than wide, with r arising slightly distad its
midpoint; 2RS nearly as sinuate as
(RS-HM)a, 1.15-1.3 (m = 1.25±.05) times
longer than 3RSa; 3RSa 1.7-2.25
(m = 1.9±0.2) times longer than r; 3RSb
ending nearly at wing tip; (RS-l-M)a sin-
uate; (RS-(-M)b present and fairly long,

Volume 8, Number 1, 1999

61

roughly 0.25 times length of m-cu; Icu-a
inclivous, usually slightly postfurcal rela-
tive to IM but varying from nearly inter-
stitial to postfurcal by 0.5 times its length.
Hind wing m-cu strongly reclivous, dis-
tinctly recurved near wing margin, ex-
tending to wing margin or nearly so as a
deeply impressed, completely pigmented
crease. Metasoma: Length of petiole 0.85-
0.95 (m=0.9±0.05) times width at apex;
apex 2.3-2.5 (m=2.4±0.1) times wider
than base; moderately and somewhat ir-
regularly striate posteriorly; dorsal carinae
well-developed over basal two-thirds,
weaker posteriorly, often indistinct at pos-
terior margin, parallel to very weakly con-
verging posteriorly; dorsope weakly de-
veloped. Metasoma unsculptured beyond
petiole. Hypopygium strongly narrowed
and distinctly pointed at extreme posterior
end, but short, not greatly attenuate,
length along midline about 0.55 times
width at base. Ovipositor tip strongly nar-
rowed subapically, without dorsal node or
carina, ventral serrations indistinct to ab-
sent; 2.3-2.55 (m = 2.4±0.1) times longer
than mesosoma; ovipositor sheath mod-
erately setose with 3 rows of setae, two of
which have 30-35 setae per row with the
third row more sparsely setose, distinct
tuft of longer setae at apex, sheath 2.0-2.25
(m = 2.09±0.1) times longer than mesoso-
ma. Color: Orange; propleuron and pro-
podeum often paler, at least in part, some-
times nearly white; ovipositor sheath,
hind tarsi, flagellum, and sometimes ped-
icel dorsally brown to light brown; base of
arolium dark brown. Wings weakly to dis-
tinctly infumate: more noticeably infu-
mate in larger specimens.

Male. — As in female except eye distinct-
ly smaller, 4.4-5.85 (m = 5.15±0.5) times
longer than temple; petiole narrower at
apex with length equal to apical width
and apex 1.95-2.25 (m=2.15) times wider
than at base. Length: 3.05-4.9 mm.

Hosts. — This species has been reared
from guava (Myrtaceae) infested with Bnc-
twccra tri/oni (Froggatt), Rauwenhoffia lei-

chardtii (Armonaceae) infested with B. hal-
fordiae (Tryon) and B. neohumeralis (Har-
dy), Sijzygium hamagense (Myrtaceae) in-
fested with B. rufofusciila (Drew and
Hancock), and Fagraea cnmbngei (Logani-
aceae) infested with B. peninsularis (Drew
and Hardy). It thus appears to be able to
attack several species of Bactrocera (Tribe
Dacini) developing in the fruit of at least
three plant families.

Material examined. — Holotype female,
"Australia: QLD Mt. Glorious 26.1.1994 G.
Quimio MG9413 ex: Rauwenhoffia lei-
chardtii fruit containing Bactrocera halfor-
diae and B. neohumeralis" Deposited in
ANIC. Paratypes (ANIC, BMNH, QDPI,
TAMU, UQBA, USNM): 179, \76, same
data as holotype; 29, 26, "Nambour Qld
24.iv.95 Guava G. Quimio"; 2 9, 13, same
data except 29.V.95; 19, "Australia: QLD
Wongabel 6 km S Atherton 1-28-1990 R.
Wharton; 19, "Malanda NQ 22.xi.1987 M.
Elson-Harris Ex Dacus rufofusculus 249";
2 9, "Sydney, N.S.W., 28-6.1954 G. J.
Snowball" one of these with an additional
label "37/54" and the other "51/54". Ad-
ditional material (not paratypes): 11 9, 3(5,
Australia, North Queensland, Balinda,
10. vi. 1993 from fruits of Fagraea canibagei
infested with Bactrocera peninsularis. This
species is known only from the eastern
coast of Australia.

Diagnosis. — As with ceratitivorus, schlin-
geri also closely resembles the Madagascar
species longicauda. The latter has a smaller
eye (slightly less than 4 times longer than
temple), somewhat more densely setose
ovipositor sheath, a smaller gap between
clypeus and mandibles when mandibles
closed, and the distal portion of the ovi-
positor is parallel-sided rather than suba-
pically narrowed relative to schlingeri. Of
the species known from Queensland,
schlingeri most closely resembles deeralen-
sis. The latter has the ventral margin of the
clypeus distinctly pointed midventrally
(Fig. 6, with mid-ventral projection more
distinct when head rotated forward), a
distinct subapical ridge on the ovipositor

62

Journal of Hymenoptera Research

Figs. 19-21. Wings of Fopnus spp.: 19 and 20, f.cJiliiigcri tore and hind wing; 21, cfintitivcnif fore wing.

(Fig. 17), and the second metasomal ter-
gum is usually weakly striate, at least ba-
sally.

Discussion. — Wharton (1997b) observed
that the hypopygium is strongly attenuate
in many of the parasitoids of fruit-infest-
ing Tephritidae. He also noted in his re-
description of Fopius that the hypopygium
varies from weakly to strongly produced
posteromedially. Though distinctly nar-
rowed and projecting posteriorly in both
ceratitivorus and schlingeri, the hypopy-
gium is much less strongly produced than
it is in members of the Fopius nmraiigensis
species group or in other fruit-infesting te-
phritid parasitoids such as the members of
the Diachasmimorpha longicaudata (Ash-
mead) species group or the species of Psi/t-

talia. Fischer (1987) includes species with
a partially visible labrum in Diachasma
Foerster. Wharton (1997b) briefly dis-
cussed variation in this character in Fopius
and other genera, noting that some of the
"variation" can be attributed to angle of
view or the degree to which the mandibles
are closed on any given specimen (com-
pare Figs. 3 and 5). Nevertheless, there are
slight differences among species of Fopius
in the exposure of the labrum, and schlin-
geri provides a good example of a species
with a partially exposed labrum (in con-
trast to the completely concealed labrum
of ceratitivorus). The ovipositor in scliliugeri
(Fig. 18) is virtually identical in form to
that of arisanus, strongly suggesting a bi-
ology similar to the latter in which the

Volume 8, Number 1, 1999

63

parasitoid oviposits into the egg of its
host. Field observations kindly supplied
by Greg Quimio of the University of
Queensland support this.

ACKNOWLEDGMENTS

I am especially grateful to the following collabo-
rators for providing reared material for this study: M.
Elson-Harris, F. Gilstrap, S. Kimani, C. Lopez-Vaa-
monde, S. Lux, R. Messing, W. Overholt, G. Quimio,
M. Ramadan, G. Steck, and T. Wong. 1 also thank M.
Trostle and S. Kimani for assistance in isolating pu-
paria from which FopiKs ccrntitivorus were reared. Ad-
ditionally, the following people kindly made material
available for examination, without which the com-
parative aspects of this work would have been im-
possible: Greg Daniels and Gimme Walter (UQBA,
Brisbane), Ian Naumann (ANIC, Canberra), David
Wahl (American Entomological Institute, Gaines-
ville), Valter Raineri (Museo Civico di Storia Naturale
"Giacomo Doria," Genoa), Gordon Nishida, Keith
Arakawa, and David Preston (BPBM, Honolulu),
Dick Drew and all the members of his research group
(formerly at QDPl, Indooroopilly), Kaoru Maeto (Shi-
koku Research Centre, Kochi), Tom Huddleston (for-
merly at BMNH, London), Ermenegildo Tremblay
(Universita de Napoli, Naples), Claire Villemant (Mu-
seum National d'Histoire Naturelle, Paris), Sergey
Belokobylskij (Zoological Institute, St. Petersburg),
Zafar Qureshi, Atomic Energy Research Centre, Tan-
do Jam, and Paul Marsh and David Smith (Systematic
Entomology Laboratory, USD A, Washington, D. C).
This work was funded in part by USDA-CSREES Spe-
cial Grant No. 96-34135, Tropical and Subtropical Ag-
riculture Research (to R. Messing), USDA/NRl, and
the National Science Foundation (DEB9712543), with
additional travel support in Australia provided bv R.
A. 1. Drew and A. Austin. Support of the Texas Ag-
riculture Experiment Station, the International Centre
of Insect Physiology and Ecology, and The Electron
Microscopy Center of Texas A&M University is also
gratefully acknowledged. Finally, 1 am indebted to
Nando Bin for assistance in obtaining specimens, and
Imelda Mercado, Ken Wilks, and Randy Scott for il-
lustrations.

LITERATURE CITED

Bridwell, J. C. 1919. Descriptions of new species of
hymenopterous parasites of muscoid Diptera
with notes on their habits. Pwceeiiiii^f of the Ha-
umiiim EntonwIo^iciU Sccict\/ 4: 166-179.

Clausen, C. P. 1978. Tephritidae (Tr\'petidae, Trupa-
neidae), pp. 320-335. In: C. P. Clausen (ed.). In-
troduced parasites and predators of arthropod
pests and weeds: a world review. Uiiitcil Stak?
Department of Agriculture Agricultural Handbook
No. 480.

Clausen, C. P., D. W. Clancy and Q. C. Chock. 1965.
Biological control of the Oriental fruit fly {Drtc»s
ilor>aUf Hendel) and other fruit flies in Hawaii.
Llnited Sffl/cs Department of Agriculture Technical
Bulletin 1322: 1-103.

Duan, J. J., M. F. Purcell and R. H. Messing. 1996.
Parasitoids of non-target tephritid flies in Ha-
waii; implications for biological control of fruit
fly pests. Eiitonwphaga 41: 245-256.

Fischer, M. (1963) 1965. Die orientalischen und aus-
tralischen Arten der Gattung Opius Wesm. Acta
Entoniologica Mu^ei Nationalif Pragae 35: 197-242.

Fischer, M. 1972. Hymenoptera: Braconidae (Opiinae
I). Das Tierreich 91; 1-620.

Fischer, M. 1977. Hymenoptera: Braconidae (Opiinae
Il-Amerika). Das Tierreich 97: 1-1001.

Fischer, M. 1987. Hymenoptera: Opiinae III — athiop-
ische, orientalische, australische und ozeanische
Region. Das Tierreich 105; 1-734.

Fullaway, D. T. 1957. A new reared Opius from Africa
(Hymenoptera, Braconidae). Proceedings of the En-
tomological Society of Washington 59: 98-99.

Gilstrap, F. E. and W. G. Hart. 1987. Biological control
of the Mediterranean fruit fly in the United States
and Central America. United States Department of
Agriculture ARS-56: 1-64.

Headrick, D. H. and R. D. Goeden. 1996. Issues con-
cerning the eradication or establishment and bi-
ological control of the Mediterranean fruit tly,
Ceratitis capitata (Wiedemann) (Diptera, Tephri-
tidae), in California. Biological Control 6: 412—121.

Knipling, E. F. 1992. Principles of insect parasitism
analyzed from new perspectives; practical impli-
cations for regulating insect populations by bio-
logical means. United States Department of Agri-
culture Handbook 693: 1-337.

Messing, R. M. 1996. Status and needs of biological
control research for tephritid flies, pp. 365-367.
In; McPheron, B. A. and G. J. Steck (eds.). Fruit
Fly Pests — A world assessment of their biology and
management. St. Lucie Press, Delray Beach, Flor-
ida.

Palacio, I. P., R. Ibrahim and A. G. Ibrahim. 1992.
Identification of immatures and male adults of
the opiine parasitoids (Biostercs spp.) of the Ori-
ental fruit fly, Bactrocera dorsalis (Hendel). The
Philippines Entomologist 8; 1124-1146.

Purcell, M. 1998. Contributions of biological control
to integrated pest management of tephritid fruit
flies in the tropics and subtropics. Integrated Pest
Management Rei'iews 3: 1-21.

Quicke, D. L. J., K. van Achterberg and H. C. J. God-
fray. 1997. Comparative morphology of the ven-
om gland and reservoir in opiine and alysiine
braconid wasps (Insecta, Hymenoptera, Bracon-
idae). Zoologica Scripta 26: 23-50.

Sharkey, M. J. and R. A. Wharton. 1997. Morphology
and Terminology, pp. 19-37. In: Wharton, R. A.,

64

Journal of Hymenoptera Research

P. M. Marsh, and M. J. Sharkey (eds.). Manual of
the New World Genern of the Ftvnihj Braconidae (Hy-
menoptera). Special Publication of the Internation-
al Society of Hymenopterists 1.

Sivinski, J. 1996. The past and present of biological
control of fruit flies, pp. 369-375. In: McPheron,
B. A. and G. J. Steck (eds.) Fruit Fly Pests-A
worhi assessment of tlieir bioh^i/ and management.
St. Lucie Press, Delray Beach, Florida.

Sivinski, J., M. Aluja and M. Lopez. 1997. Spatial and
temporal distributions of parasitoids of Mexican
Annstrepha species (Diptera: Tephritidae) within
the canopies of fruit trees. Annals of tlie Entomo-
logical Society of America 90: 604-618.

Steck, G. J., F. E. Gilstrap, R. A. Wharton and W. G.
Hart. 1986. Braconid parasitoids of Tephritidae
(Diptera) infesting coffee and other fruits in
west-central Africa. Entomopliaga 31: 59-67.

Tobias, V. 1. (1977) 1978. The genus Opius Wesm. (Hy-
menoptera, Braconidae) as parasites of fruit flies
(Diptera, Tephritidae). Entomological Review 56:
132-139.

Van Achterberg, C. and K. Maeto. 1990. Two new
aberrant species of Braconidae (Hymenoptera)
from Japan. Zoologische Mededelingen Eeiden 64:
59-70.

Van Achterberg, C. and A. Salvo. 1997. Reared Opi-
inae (Hymenoptera: Braconidae) from Argentina.
Zoologische Mededelingen Leiden 71: 189-214.

Waterhouse, D. F. 1993. Bwlogical Control: Pacific Pros-
pects—Supplement 2. Australian Centre for Inter-
national Agricultural Research, Canberra. 138
pp.

Wharton, R. A. 1987. Changes in nomenclature and
classification of some opiine Braconidae (Hyme-
noptera). Proceedings oftlie Entomological Society of
Washington 89: 61-73.

Wharton, R. A. 1988. Classification of the braconid
subfamily Opiinae (Hymenoptera). The Canadian
Entomologist 121: 333-360.
Wharton, R. A. 1989. Classical Biological Control of
Fruit-infesting Tephritidae, pp. 303-313. In: Rob-
inson, A. S. and G. Hooper (eds.). Fruit Flies,
Their Biology, Natural Enemies and Control. Elsev-
ier Science Publishers B.V., Amsterdam.
Wharton, R. A. 1997a. Subfamily Opiinae, pp. 378-
395. In: Wharton, R. A., P. M. Marsh, and M. J.
Sharkey (eds.). Manual of the New World Genera of
the Family Braconidae (Hymenoptera). Special Pub-
lication of the International Society of Hymen-
opterists 1.
Wharton, R. A. 1997b. Generic relationships of opiine
Braconidae (Hymenoptera) parasitic on fruit-in-
festing Tephritidae (Diptera). Contributions of the
American Entomological Institute 30: 1-53.
Wharton, R. A. and F. E. Gilstrap. 1983. Key to and
status of opiine braconid (Hymenoptera) parasit-
oids used in biological control of Ceratitis and
Dacus s. 1. (Diptera: Tephritidae). Annals of the
Entomological Society of America 76: 721-742.
Wharton, R. A., F. E. Gilstrap, R. H. Rhode, M. Fis-
chel-M. and W. G. Hart. 1981. Hymenopterous
egg-pupal and larval-pupal parasitoids of Cera-
titis capitala and Anastrepha spp. in Costa Rica.
Entomopliaga 26: 285-290.
Wharton, R. A. and P. M. Marsh. 1978. New World
Opiinae (Hymenoptera: Braconidae) parasitic on
Tephritidae (Diptera). Journal of the Washington
Academy of Sciences 68: 147-167.
White, 1. M. and M. M. Elson-Harris. 1992. Fruit Flies
of Economic Significance: Their Identification and Bi-
onomics. CAB International, London and ACIAR,
Canberra. 601 pp.

J. HYM. RES.
Vol. 8(1), 1999, pp. 65-73

New Genera and Species of Brachycistidine Wasps from

Southwestern North America

(Hymenoptera: Tiphiidae: Brachycistidinae)

Lynn S. Kimsey and Marius S. Wasbauer

Bohart Museum of Entomology, Department of Entomology, University of California,

Davis 95616

Abstract. — The new genera Brachymaya, Paraquemaya and Sedomaya are described from
southwestern Arizona, Baja California and Sonora Mexico, and Imperial Co., California. Brachy-
maya is based on the new species mexicana. Paraquemaya is based on the new species pallida, and
Sedomaya is based on the new species glamisensis. Three other new species of Paraquemaya are
also described: bitincta, maxima, and bajaensis.

Until recently (Kimsey and Wasbauer
1998) the subfamily Brachycistidinae had
not received taxonomic attention in nearly
three decades. The last major revisions of
any kind of members of this group were
published by Mickel and Krombein (1942)
and later by Wasbauer (1958, 1966 and
1968). Studies of unidentified male bra-
chycistidines, which have accumulated in
collections in the past 30 years, have re-
vealed a variety of new taxa.

Six of these undescribed species had a
combination of features, which did not fit
any of the current generic groupings. Pre-
liminary cladistic analyses of the Brachy-
cistidinae indicated the need to describe
three new genera for these species. Each
of these genera is characterized by a
unique combination of apomorphic fea-
tures, which occur in other brachycistidine
genera in various combinations. They in-
clude a tailed antennal socket, external
mandibular carina, stridulatory structure
on the forecoxa, an elongate digitus,
changes in the position of the carina en-
closing the oral fossa, loss of the hindcoxal
carina, loss of the basolateral carinae on
the first gastral tergum, and highly re-
duced wing venation. The first of these
genera, Paraquemaya, has a large number

of primitive features, although it is still
more highly derived than Quemaya. Para-
quemaya has the following apomorphic
characteristics: a tailed antennal socket,
presence of a mandibular carina, an elon-
gate digitus and hindwing venation simi-
lar to that seen in Brachycistellus and Had-
rocistis. The second new genus, Brachy-
maya, appears to be most closely related to
Bracliycistina and Hadrocistis, based on the
slender, ecarinate mandible and lack of
the dorsal carina on the hindcoxa. Finally,
the third genus, Sedomaya, belongs to a
group of genera with the stridulatory
structure on the forecoxa and the first gas-
tral sternum with a short medial carina.
This group also includes Brachycistis, Bra-
chycistellus and Colocistis.

As with many of the brachycistidine
genera the undoubtedly wingless, and
probably nocturnal, females are unknown.
Although a number of genera have been
described based only on females (Mickel
and Krombein 1942) the taxonomy of the
Brachycistidinae is essentially based on
the males. None of the female-based gen-
era described by Mickel and Krombein
(1942) are known to occur in the same re-
gion as the taxa described below (Fig. 1),
although this is no guarantee that there is

66

Journal of Hymenoptera Research

Fig. 1. Distribution map showing collection locali-
ties of the brachycistidine genera Brachymaya, Para-
queniayn and Scdomnya.

no geographic overlap. In addition, the
genera described by Mickel and Krombein
are relatively large, ranging in length from
5-12 mm. The females of the new genera
described below should be small, about 2-
3 mm in length, roughly the size of female
Quemaya, which are also as yet unde-
scribed. Female brachycistidines described
thus far are considerably larger than the
females of Erachymaya, Paraqiiemaya and
Sedomaya should be based on the male to
female body size ratio seen in species
where both sexes have been described.
Additionally, no hosts are known for any
of the new species described below. De-
scription of these new genera is essential
to enable us to finalize phylogenetic ana-
lyses of the subfamily Brachycistidinae.

MATERIALS

Specimens used in this study came from
the Bohart Museum of Entomology, Uni-
versity of California, Davis, S. L. Heydon

(DAVIS); California Academy of Sciences,
San Francisco, N. Penny (SAN FRANCIS-
CO); Los Angeles County Museum, Cali-
fornia, R. R. Snelling (LOS ANGELES);
University of California, Riverside, S.
Frommer, S. Triapitsyn (RIVERSIDE), and
the personal collection of M. S. Wasbauer.
Type repositories are indicated by the cap-
italized name given in parentheses.

Brachymaya Kimsey and Wasbauer,
new genus

(Figs. 2, 6, 18, 19)

Description of male. — Face (Fig. 6): Man-
dible with two apical teeth and without
longitudinal carina on external surface;
palpi long, extending well outside of oral
fossa, maxillary palpus 6-segmented; la-
bial palpus 4-segmented; inner eye margin
converging medially; flagellomeres some-
what arcuate, particularly apical articles;
antennal carina greatly thickened into
ventral subtriangular process, with tail-
like carinule; gular carina with large sub-
truncate swelling near mandible; clypeus
medially evenly convex, apical margin
without ventral bevel; forecoxa without
stridulatory patch; scrobal pit small and
circular; propodeum without longitudinal
groove on dorsal surface and without
transverse carina; metasternum without
medial ridge terminating in two lobes
near hindcoxal base; hindcoxa without
ventral or dorsal longitudinal carinae;
wings (Fig. 2): forewing with one large
rhomboid submarginal cell, and smaller
second submarginal cell, marginal cell
nearly parallel-sided, Rl barely visible
and bending away from the costal margin
toward the stigma, one discoidal cell, one
subdiscoidal cell; hindwing M vein di-
verging from Cu + M after cross vein cu-
a; gastral segment I, tergum with well-de-
veloped lateral carina at base, sternum
without longitudinal carina extending
from base; epipygium delimited by small
sublateral welt, apex truncate or some-
what short and apically rounded; genital
capsule (Figs. 18, 19): paramere slender

VoLUMi; 8, NUMBEK 1, 1999

67

and tapering dorsally; volsella with small
submedial lobe externally, inner surface
with several rows of denticles; digitus
elongate terminating in acutely pointed
lobe.

Distribution. — This genus is known only
from the vicinity of San Augustine, BCN,
Mexico.

Eti/iuolo^i/. — The name is intended as a
nonsense combination of letters, taken
from Brachycistis and Quemaya, and is as-
sumed to be feminine.

Type species. — Bracbymaya mexicana n.
sp.

Discussion. — Bracbymaya is somewhat
similar to Bracbycistiim and Hadrocistis. All
three genera are characterized by having
a slender mandible, with subsidiary den-
tition reduced to one or two very small
teeth, and the hindcoxa lacking a dorso-
basal carina. As with most brachycistidine
genera Brachymaya has the digitus elon-
gate and pointed. However, Brachymaya
has the antennal socket subtended by a
large triangular enlargement of the carina
and the gular carina is dilated, forming a
truncate projection near the mandibular
base. In addition, the wing venation is re-
duced, with one discoidal and one sub-
marginal cell.

Brachymaifa mexicana Kimsey and

Wasbauer, new species

(Figs. 2, 6, 8, 19)

Description of male. — Body length 5-6
mm; face (Fig. 6); clypeus evenly convex
medially, apical truncation 0.9-1.0 mid-
ocellus diameters wide; interantennal dis-
tance 0.7-0.8 midocellus diameters wide;
distance between midocellus and nearest
eye margin 1.4-1.5 midocellus diameters;
flagellomeres I-II 2.2-2.4 times as long as
broad; facial and thoracic punctures small,
shallow and widely spaced, 4—6 puncture
diameters apart; forewing with two sub-
marginal cells (Fig. 2); abdominal seg-
ments appearing impunctate, integument
finely shagreened; genital capsule (Figs.
18, 19). Head dark brown; thorax and legs

pale yellowish brown; abdomen darker
brown; forewing venation yellow, except
stigma brown; hindwing venation yellow;
wing membrane faintly yellow tinted.

Type material. — Holotype male: Mexico:
BCN, San Augustine, 20 Oct. 1956, R. Mat-
toni (LOS ANGELES). Paratypes: three-
same data as holotype (LOS ANGELES,
DAVIS); one — 3 mi. s. San Augustine, 14
June 1973, J. Doyen (DAVIS).

Etymology. — This species is named after
the country of collection.

Discussion. — Although there are cur-
rently no other species placed in this ge-
nus, species distinctions probably include
proportions of the flagellomeres, ocello-
cular distances, punctation and coloration
of the wing veins.

Paraqiiemaya Kimsey and Wasbauer,
new genus

(Figs. 3, 4, 7-9, 11, 12, 14-17)

Description of male. — Face (Figs. 7-9):
Mandible with three apical teeth and lon-
gitudinal carina on external surface; palpi
long, extending well outside of oral fossa,
maxillary palpus 6-segmented; labial pal-
pus 4-segmented; flagellomeres somewhat
arcuate; antennal carina slightly thickened
ventrally, but without tail-like carinule;
gular carina with tooth-like projection
near mandible (Fig. 12); forecoxa without
stridulatory structure; scrobal pit small
and circular; propodeum with longitudi-
nal groove on dorsal surface and no trans-
verse carina; metasternum with medial
ridge terminating in two small lobes near
hindcoxal base; hindcoxa without ventral
or dorsal longitudinal carinae; wings
(Figs. 3, 4): forewing with one large rhom-
boid submarginal cell, and smaller second
submarginal cell in several species, mar-
ginal cell nearly parallel-sided, Rl barely
visible and bending away from the costal
margin toward the stigma, one discoidal
cell, one subdiscoidal cell; hindwing M
vein diverging from Cu -I- M after cross
vein cu-a; gastral segment I, tergum with
well-developed lateral carina at base, vis-

68

Journal of Hymenoptera Research

5 Sednmaya
glamisensis

13 Scdomaya
glamisensis

Figs. 2-13. Figs. 2-5. Fore and hindwing of males. Figs. 6-11. Front view of male face, with right antenna
removed. Figs. 12, 13. Side view of male head.

ible in most specimens, sternum with
short longitudinal carina extending from
base; epipygium delimited by small sub-
lateral welt, apex truncate or somewhat
rounded; genital capsule (Figs. 14-17):
parameres slender and tapering apically;
volsella with small rounded submedial

lobe, inner margin with small denticles;
digitus elongate and awl-shaped.

Distribution. — This genus occurs in
southern Arizona and Baja California,
Mexico.

Etytnolo^y. — The generic name refers to
the superficial similarity, because of their

Volume 8, Number 1, 1999

69

small size, to species of Qufiiun/a; femi

nine.

Discussion. — Although Paraqucniai/a re-
sembles Quemaya in terms of its reduced
wing venation and small size, it shares the
majority of characteristics with more de-
rived brachycistidine genera including
Brachycistis and Brncliycistelhis. Features
shared with these genera include the elon-
gate digitus, carinate mandible, unmodi-
fied palpi, basal gastral sternum with a

medial carina, and basal tergum with a
well-developed lateral carina adjacent to
the petiolar insertion. The presence of a
gular tooth is a characteristic shared with
some species of Brachycistis and one Que-
maya species. Additional diagnostic fea-
tures include the lack of hindcoxal carinae,
no forecoxal stridulatory structure and the
configuration of the hindwing venation,
which is very similar to that of Hadrocistis
and Brachycistellus.

KEY TO THE SPECIES OF PARAQUEMAYA (MALES)

1. Forewing with two submarginal cells (Fig. 3); interantennal distance 0.5 midocellus di-
ameter wide or less; body usually appearing bicolored with thorax paler than head and
abdomen 2

- Forewing with one submarginal cell (Fig. 4); interantennal distance more than 0.5 mid-
ocellus diameter; body concolorous 3

2. Shortest distance between midocellus and eye margin 1-1.2 midocellus diameters wide
(Fig. 11); flagellomeres I and II subequal, 2.3-2.6 x as long as broad . . maxima new species

- Shortest distance between midocellus and eye margin 1.4-1.6 midocellus diameters wide
(Fig. 8); flagellomere I shorter than II, 2.2-2.3 times as long as broad and II 2.3-2.5 x as
long as broad bitincta new species

3. Flagellomere I 2.0-2.2 times as long as broad (Fig. 9); flagellomere II 2.2-2.4 x as long as
broad; interantennal distance 0.6-0.8 midocellus diameters wide; shortest distance between
midocellus and eye margin 1.8-2.0 midocellus diameters wide pallida new species

- Flagellomere I 1.7-1.9X as long as broad (Fig. 7), flagellomere II length 2.0-2.2X breadth;
interantennal distance 0.8-0.9 midocellus diameters wide; shortest distance between mid-
ocellus and eye margin 2.1-2.3 midocellus diameters wide bajaensis new species

Paraquemaya bajaensis Kimsey and
Wasbauer, new species

(Fig. 7)

Description of male. — Body length 4 mm;
face (Fig. 7); clypeus with narrowly acute
medial projection, apical truncation 0.9
midocellus diameter wide; interantennal
distance 0.8 midocellus diameter; distance
between midocellus and nearest eye mar-
gin 2.3 midocellus diameters; flagellomere
I twice as long as broad; flagellomere II
length 2.4 X breadth; facial and thoracic
punctures small and widely spaced, 4-6
puncture diameters apart; forewing with
one submarginal cell; abdominal segments
appearing impunctate, integument finely
shagreened. Body color dark brown; fore-

wing venation light brown-tinted, except
stigma darker brown; hindwing venation
faintly brown-tinted; wing membrane un-
tinted.

Type material. — Holotype male; Mexico,
BCS, 14 mi s El Arco, 2 Nov. 1965, W.
Ewart & R. Dickson (RIVERSIDE). Para-
types: 1 male same data as holotype, ex-
cept 5 Nov. 1965; one male: BCN, El Con-
suelo, 0-50 ft. elev., 11 Sept. 1983, R. R.
Snelling (DAVIS).

Etymologi/. — The name is derived from
the region of collection, Baja California,
Mexico.

Discussion. — This species most closely
resembles pallida, as both have a single
submarginal cell and the body color is

70

Journal of Hymenoptera Research

16. ---^ 17.

Paraquemaya bitincta

Paraquemaya maxima

Brachymaya mexicana

21.
Sedomaya glamisensis

Figs. 14-21. Male genital capsule. Figs. 14, 16, 18, 20. Ventral view. Figs. 15, 19, 21. Dorsal view. Fig. 17.
Lateral view.

concolorous. These two also have the void-
est ocellocular distance and widest inter-
antennal distance. P. bajaensis has the
smallest ocelli in the genus, with the mid-
ocellus separated from the eye margin by
2.3 midocellar diameters. This species also
has the longest basal flagellomeres of any
species of Paraqueniai/a.

Paraquemaya bitincta Kimsey and
Wasbauer, new species

(Figs. 3, 8, 16, 17)
Description of male. — Body length 5 mm;
face (Fig. 8); clypeus with narrowly acute
medial projection, apical truncation 1 mid-
ocellus diameter wide; interantennal dis-
tance 0.3 midocellus diameter; distance

Volume 8, Number 1, 1999

71

between midocellus and nearest eye mar-
gin 1.6 midocellus diameters; flagello-
meres I and II subequal, length 2.3-2.4 X
breadth; facial and thoracic punctures
small and widely spaced, 4-8 puncture di-
ameters apart; forewing with two submar-
ginal cells (Fig. 3); abdominal segments es-
sentially impunctate, integument finely
shagreened; genital capsule (Figs. 16, 17).
Body color reddish brown, head and ab-
domen darker brown than thorax; fore-
wing venation untinted, except stigma
brown; hindwing venation untinted; wing
membrane untinted.

Ti/pe mnterinl. — Holotype male: Arizona,
Pima Co., Organ Pipe, 12 April 1947, A. L.
Melander (RIVERSIDE). Three paratypes,
same data as holotype (RIVERSIDE, DA-
VIS).

Eti/mologi/. — The name, bitincta, is de-
rived from the two-tone brown color of
the body.

Discussion. — This species can be distin-
guished from pallida and bajaejisis by the
presence of two submarginal cells. It can
be separated from maxima by the smaller
body size, shorter flagellomeres shorter,
and greater distance between the eye and
the midocellus.

Paraqiiemaya maxima Kimsey and
Wasbauer, new species

(Figs. 11, 14, 15)

Description of male. — Body length 7 mm;
face (Fig. 11); clypeus with acute, narrow-
ly hooked medial projection, apical trun-
cation 1 midocellus diameter wide; inter-
antennal distance 0.5-0.6 midocellus di-
ameters; distance between midocellus and
nearest eye margin 0.1-1.1 midocellus di-
ameters; flagellomeres I and II length 2.6 x
breadth; facial and thoracic punctures
small and widely spaced, 4-6 puncture di-
ameters apart; forewing with two submar-
ginal cells; abdominal segment I with
broad, shallow irregular punctures and
shagreening; segments II-V with sparse
shallow punctures, 4-6 puncture diame-
ters apart; genital capsule (Figs. 14, 15).

Body color: head and abdomen brown,
thorax lighter reddish brown; forewing
venation pale brown-tinted, except stigma
reddish brown; hindwing venation nearly
colorless; wing membrane untinted.

Ti/pe material. — Holotype male: Mexico,
Sonora, 22 km se Quitovac, 14 Nov. 1965,
W. Edward & R. Dickson (RIVERSIDE).
Two paratypes, same data as holotype
(DAVIS, RIVERSIDE).

Eti/mologi/. — The name is based on the
unusually large body size for this genus.

Discussion. — Other than the large body
size of this species, the two submarginal
cells and long flagellomeres will distin-
guish maxima from other species placed in
Paraquemaya.

Paraqiiemaya pallida Kimsey and
Wasbauer, new species

(Figs. 4, 9)

Description of male. — Body length 3.5-6.0
mm; face (Fig. 9); clypeus with small acute
medial projection, apical truncation 1.0-
1.3 midocellus diameters wide; interanten-
nal distance 0.6-0.8 midocellus diameters;
distance between midocellus and nearest
eye margin 2.0-2.2 midocellus diameters;
flagellomere I 2.0-2.2 times as long as
broad; flagellomere II length 2.2-2.4 X
breadth; facial and thoracic punctures
small and widely spaced, 3-5 puncture di-
ameters apart; forewing with one submar-
ginal cell (Fig. 4); abdominal segments
with sparse shallow punctures, 2-4 punc-
ture diameters apart. Body color pale red-
dish brown, rarely darker brown; fore-
wing venation slightly brown-tinted, ex-
cept stigma darker; hindwing venation
untinted.

Ti/pe material. — Holotype male: MEXI-
CO: Baja California Sur, 4 mi. wsw Mira-
flores, 23-24 April 1979, M. Wasbauer
(DAVIS). Paratypes, 68 males (DAVIS,
SAN FRANCISCO): 25— same data as ho-
lotype; four — El Salto, 8 mi ne Todos San-
tos, 9 Oct. 1983, D. Faulkner & F. An-
drews; twenty-six — Los Barriles, 24 Mar.-
5 April 1984, J. H. Lynch; two — Puerto Es-

72

Journal of Hymenoptera Research

condito, 10-14 July 1989, R. Shaver; one—
BCN, Sierra Calamajue, 11 km e Chapala,
29°31'N 115°42'E, 23 Aug. 1994, S. Hey-
don; one — BCS: El Pescadero, Playa Los
Cerritos, 16-17 April 1979, M. Wasbauer;
one— BCS, 13 mi nw La Paz, 10 Nov. 1965,
W. Ewart and R. Dicksen; one — Sonora, 14
mi w Guaymas, 13 Nov. 1965, Ewart and
R. Dicksen.

Etymology. — The name, pallida, refers to
the pale coloration of the male.

Discussion. — Paraquemaya pallida is char-
acterized by having one submarginal cell,
flagellomeres I and II subequal in length
and about 2.2 X as long as broad, and the
midocellus separated from the eye by 2
midocellus diameters or slightly less.

Sedornaya Kimsey and Wasbauer,
new genus

(Figs. 5, 10, 13, 20, 21)

Description of male. — Face (Fig. 10): man-
dible with three apical teeth and longitu-
dinal carina on external surface; palpi
long, extending well outside of oral fossa,
maxillary palpus 6-segmented; labial pal-
pus 4-segmented; eye strongly converging
medially, inner eye margin slightly in-
dented; flagellomeres somewhat arcuate,
particularly apical articles; antennal carina
slightly thickened ventrally, but without
tail-like carinule; gular carina with large
subtruncate swelling near mandible (Fig.
13); clypeus medially concave and slightly
trilobate apically, apical margin with ven-
tral bevel; forecoxa with small stridulatory
patch; scrobal pit small and circular; pro-
podeum with obsolescent longitudinal
groove on dorsal surface and no trans-
verse carina; metasternum with medial
ridge terminating in two small lobes near
hindcoxal base; hindcoxa with dorsal lon-
gitudinal carinae, without ventral one;
wings (Fig. 5): forewing with one large
rhomboid submarginal cell, and smaller
second submarginal cell, marginal cell
nearly parallel-sided, Rl barely visible
and bending away from the costal margin
toward the stigma, one discoidal cell, one

subdiscoidal cell; hindwing M vein di-
verging from M + Cu after cross vein cu-
a; gastral segment I, tergum with well-de-
veloped lateral carina at base, visible in
most specimens, sternum with short lon-
gitudinal carina extending from base; api-
cal tergum evenly convex, apex truncate
or somewhat rounded; short and apically
rounded; genital capsule (Figs. 20, 21):
paramere slender and tapering apically;
volsella with rounded medial lobe, inner
margin with numerous small denticles;
digitus elongate and apically acute.

Distribution. — This genus is known only
from the vicinity of Glamis, in the Algo-
dones Dunes, Imperial Co., California.

Etymology. — The name is a nonsense
combination of letters, and is assumed to
be feminine.

Type species. — Sedornaya glamisensis n. sp.

Discussion. — No one genus appears to
be the sister group of Sedomaya. It belongs
to the group of genera characterized by
the presence of a stridulatory structure on
the forecoxae, elongate digitus and the
first gastral sternum with a short longitu-
dinal carina extending posteromedially
from the base. Sedomaya, Dolichetropis,
Acanthetropis and Colocistis all have a ven-
tral clypeal bevel below the apex. How-
ever, unlike the latter three genera Sedo-
maya has highly reduced wing venation,
having only two small forewing submar-
ginal cells and one discoidal cell.

Sedornaya glamisensis Kimsey and
Wasbauer, new species

(Figs. 5, 10, 13, 20, 21)

Description of male. — Body length 5-7
mm; face (Figs. 10, 13); clypeus medially
convex, apex slightly trilobate with medial
lobe projecting anteriorly in profile; apical
truncation 0.9-1.0 midocellus diameters
wide; interantennal distance 0.4-0.5 mido-
cellus diameters; flagellomere I-II 2.9-3.1
times as long as broad; facial and thoracic
punctures small, shallow and widely
spaced, 4-6 puncture diameters apart;
forewing with one submarginal cell (Fig.

Volume 8, Number 1, 1999

73

5); abdominal segments appearing im-
punctate, integument finely shagreened;
genital capsule (Figs. 20, 21). Body color
pale reddish brown; fore and hindwing
venation yellow; wing membrane slightly
yellow tinted.

Type material. — Holotype male: Califor-
nia, Imperial Co., Glamis, 23 April 1972,
M. Wasbauer, blacklight (DAVIS). Para-
types: twenty-five males (DAVIS); two —
same data as holotype; twenty-two — 3 mi
n Glamis, 15-16 Sept. 1972, M. Wasbauer
and A. Hardy; four— 10 Sept 1974, M.
Wasbauer and R. McMaster.

Eti/mologi/. — This species is named after
the vicinity of the collection sites in and
around Glamis, on the edge of the Algo-
dones Dunes.

Discussion. — As with Brachymaya mexi-
cana, species distinctions within this genus
will probably be based on flagellar dimen-
sions, the size of the midocellus and its
distance to the ocular margin, other facial

dimensions and perhaps the configuration
of the clypeus.

ACKNOWLEDGMENTS

Our thanks to Karl Krombein for his help with the
manuscript, and to the fine collections support and
helpful collection managers who made this study
possible.

LITERATURE CITED

Kimsey, L. S. & M. S. Wasbauer. 1998. Revision of the
American tiphiid genus Qiicniaya Pate, journal of
Hymeiioptcra Rc^cnrcli 71:38-47.

Mickel, C. E. and K. V. Krombein. 1942. Glyptometopa
Ashmead and related genera in the Brachycisti-
dinae with descriptions of new genera and spe-
cies. Aincriam Midland Naturalist 28:648-679.

Wasbauer, M. S. 1958. A new genus of brachycisti-
dine wasps. Pan-Pacific Entoniologtst 34:139-142.

Wasbauer, M. S. 1966. Revision of the male wasps of
the genus Bracliyctstis in America north of Mex-
ico. University of California Publicatioiif in Ento-
mology 43:1-96.

Wasbauer, M. S. 1968. New genera of male Brachy-
cistidinae with a redescription of Brachycistelhis
Baker and a key to North American genera. Pan-
Pacific Entomologist 44:184-197.

J. HYM. RES.
Vol. 8(1), 1999, pp. 74-97

Deceptive Similarity in Army Ants of the Genus Neivamyrmex

(Hymenoptera: Formicidae): Taxonomy, Distribution and Biology of

N. califomicus (Mayr) and N. nigrescens (Cresson)

Philip S. Ward
Department of Entomology, University of California, Davis, California 95616, USA

Abstract. — The army ant Neivamyrmex califomicus (Mayr) is demonstrated to be a distinct species,
endemic to California and adjacent Baja California, whose range overlaps that of N. nigrescens
(Cresson), with which it has been previously confused. N. nigrescens is widespread throughout
the southern Nearctic region, and shows extensive morphological variation in shape, size and
sculpture. In the northwestern extremities of its range (i.e., north-central California, Nevada, Utah,
and southwestern Colorado) N. nigrescens is convergently similar to N. califomicus in certain as-
pects of worker morphology: workers lack the densely punctulate head sculpture typical of this
species, and in their more shiny appearance they are superficially similar to workers of N. cali-
fomicus. Many records of "N. califomicus" (and the description of its supposed queen) actually
refer to this shiny morph of N. nigrescens. That the shiny form is conspecific with, although
partially differentiated from, other populations of N. nigrescens is indicated by the occurrence of
intermediate populations in a relatively narrow transition zone in the San Gabriel Mountains of
southern California. No intermediates have been observed between N. nigrescens and N. califor-
nicus. The latter species appears to be more closely related to N. opacitborax Emery than to N.
nigrescens. In California N. nigrescens and N. califomicus raid the nests of other ants, including
Messor andrei, Pheidole califomica, P. hyatti, Solenopsis molesta, and Formica moki. Field observations,
combined with indirect evidence from the contents of ant nest middens, suggest considerable
overlap in prey choice and habitat use. Both N. nigrescens and N. califomicus are commonly sym-
patric in southern California (where N. nigrescens retains its distinctive granular-punctulate body
sculpture), but less frequently so in northern California where they are more similar in appearance,
and where N. califomicus tends to be confined to more mesic, coastal areas than N. nigrescens.

Army ants in the genus Neivamyrmex given that the reproductive females
are a frequent component of ant commu- (queens) of these and other army ants are
nities in tropical and warm temperate re- entirely wingless and have quite limited
gions of the New World. The group has powers of dispersal (Gotwald 1995), a fac-
received considerable attention from tax- tor that favors population differentiation,
onomists (e.g., Smith 1942; Borgmeier The present contribution is concerned
1953, 1955, 1958; Watkins 1976, 1982, with clarifying the taxonomy of Neivmnyr-
1985), with the resulting recognition of mex califomicus (Mayr) and several closely
about 120 species (Bolton 1995). Taxonom- related species. Earlier treatments of these
ic problems persist, however, partly be- species are shown to be misleading. Sim-
cause many names are based on either pie morphometric analyses help to resolve
workers or males only — and the names for and diagnose N. califomicus and N. iii^res-
the two castes are not yet cross-referenced cens (Cresson), two species whose taxo-
and synonymized — but also because there nomic distinctness was previously called
is substantial and often confusing intra- into question (Watkins 1985). Both these
specific variation in morphology (Smith species — but especially N. nigrescens —
1942). Such variation is not unexpected show considerable variation in integu-

Volume 8, Number 1, 1999

75

ment sculpture, and this phenomenon
misled earlier investigators about species
limits.

The range of Neivamyrmex californicus is
more restricted than previously thought,
the species being confined to the Califor-
nian floristic province (California and ad-
jacent northern Baja California), where it
is broadly sympatric with N. uigrescens.
The latter is shown to be a highly poly-
typic species with an extensive transcon-
tinental distribution.

MATERIALS AND METHODS

Specimens were examined in the fol-
lowing collections:

CASC California Academy of Sciences,
San Francisco, California, U. S. A.

CDAE California Department of Food &
Agriculture, Sacramento, Califor-
nia, U. S. A.

JTLC John T. Longino Collection,
Olympia, Washington, U. S. A.

LACM Natural History Museum of Los
Angeles County, Los Angeles,
California, U. S. A.

MCZC Museum of Comparative Zoolo-
gy, Harvard University, Cam-
bridge, Massachusetts, U. S. A.

MHNG Museum d'Histoire Naturelle,
Geneva, Switzerland

NHMV Naturhistorisches Museum, Vien-
na, Austria

RAJC Robert A. Johnson Collection,
Tempe, Arizona, U. S. A.

UCDC Bohart Museum of Entomology,
University of California, Davis,
California, U. S. A.

USNM National Museum of Natural His-
tory, Smithsonian Institution,
Washington, D. C, U. S. A.

Other collections cited are:

ANSP Academy of Natural Sciences, Phil-
adelphia, U. S. A.

MCSN Museo Civico di Storia Naturale,
Genoa, Italy

The following measurements and indi-

ces were used for workers and queens. All
measurements were taken at 50 X magni-
fication with a Wild MSA microscope and
a pair of Nikon stage micrometers wired
to a digital readout. Measurements were
recorded to the nearest thousandth of a
millimeter.

HW Maximum width of head, mea-
sured in full-face (frontal) view

HL Maximum length of head in fron-
tal view, from the midpoint of a
line drawn across the posterior
margin to the anteriormost point
on the clypeal margin. This ex-
cludes the thin, lamelliform clypeal
apron, which projects forward
from the clypeal margin and may
be partly hidden by the mandibles.

MFC Minimum distance between the
frontal carinae

SL Length of the scape, excluding the
basal neck

WL Weber's length: length of the me-
sosoma, as seen in lateral view,
from the anterior pronotal margin
(excluding the collar) to the pos-
terior extremity of the metapleu-
ron

PL Length of the petiole, measured in
lateral view along the long axis of
the petiole, from the anterior mar-
gin (excluding the short peduncle)
to the posterior extremity

PH Petiole height, measured at right
angles to PL, from the summit of
the petiole to the petiolar venter,
excluding the anteroventral tooth
(if present)

DPW Maximum dorsal width of the pet-
iole, measured in dorsal view

PPW Maximum dorsal width of the
postpetiole, measured in dorsal
view

MTL Length of the metatibia, excluding
the basal condyle (Fig. 3)

QGL Length of the gaster (queen caste
only), measured in lateral view
from the posterior end of the gas-

76

Journal of Hymenoptera Research

ter to the anterior extremity of ab-
dominal segment 3, excluding the
helcium. This measurement was
taken only on non-physogastric
queens.
CI Cephalic index: HW/HL

SI Scape index: SL/HW

FCI Frontal carinal index: MFC/ HW
PLI Petiole length index: PH/ PL
PWI Petiole width index: DPW/ PL
MTI Meta tibial index: MTL/HW

Neivamyrmex califomictis and
allied species.

Among the North American army ants
of the genus Neivamyrmex three species —
N. californicus, N. nigrescens and N. texan-
us — can be diagnosed in the worker caste
by having a mandible whose basal margin
rounds gradually into masticatory margin,
without an angular junction (Figs. \, 4, 7).
The species have many other traits in com-
mon, including: moderately large size
(HW 0.60-1.42); well developed clypeal
apron; relatively prominent ocellus-like
eye with convex surface; weakly devel-
oped occipital lobes; lamellate and ven-
trally directed lower pronotal flange; pet-
iole longer than high (PLI 0.68-0.96) and
markedly longer than wide (PWI 0.52-
0.74); and dense punctulate sculpture cov-
ering part of the body but without con-
spicuous overlying coarse impressions or
rugae.

These taxa are part of a larger group of
species, including N. chamelensis Watkins,
N. cornutus Watkins, N. manni (Wheeler),
N. opacithorax (Emery) and N. stimidirnsti
(Norton), that can be placed together, on
the basis of similarities in worker mor-
phology and male genitalia, in an assem-
blage of Neivamyrmex species termed

"Gruppe VI" by Borgmeier (1955: 490).
Although the three species treated in this
paper are evidently closely related, their
exact phylogenetic relationship to one an-
other and to these other taxa remains to
be resolved.

List of species considered here, with
known castes (w = worker, m = male, q
= queen) and distribution:

californicus (Mayr 1870) (w)

U. S. A.: Cahfornia

Mexico: Baja California
nigrescens (Cresson 1872) (w, q, m)

U. S. A.: California, Nevada, Utah, Ari-
zona, New Mexico, Colorado, and
east-central United States

Mexico: Baja California, Baja California
Sur, Jalisco, Nayarit, Oaxaca, San
Luis Potosi, Sonora, Tamaulipas
texanus Watkins 1972 (w, q, m)

U. S. A.: Arizona, New Mexico, Colo-
rado, and east-central United States

Mexico: Chihuahua, Durango, Hidalgo,
Jalisco, Nuevo Leon, San Luis Po-
tosi, Sonora

SPECIES ACCOUNTS

Neivamyrmex californicus (Mayr 1870)

(Figs. 1-3, 10)

Eciton cnlifornicum Mayr 1870: 969. Nineteen
syntype workers, San Francisco (Schaufuss)
(NHMV) [examined]. One syntype worker
here designated lectotype.

Ecitoii {Acainatus) cnlifornicum Mayr; Emery
1894; 182.

Eciton {Acainntiis) cnlifornicum var. obscurn Forel
1914: 265. Two syntype workers, Vista, Cal-
ifornia (E. Hindle) (MHNG) [examined]. One
syntype worker here designated lectotype.
Synonymy by Borgmeier 1955: 517; here con-
firmed.

Eciton {Ncivnnn/rmcx) cnlifornicum Mayr; Smith

Figs. 1-12. Neivamyrmex workers, full-face view of head (1, 4, 7), lateral view of mesosoma, petiole and
postpetiole (2, 5, 8), lateral view of metatibia (3, 6, 9), and dorsal view of propodeum, petiole and postpetiole
(10-12). 1-3, 10: N. californicus, San Francisco, California, lectotype worker; 4-6, 11: N. nigrescens. shiny form,
2km SB Mt. Vaca, California; 7-9, 12: N. nigrescens, typical form with opaque head, Sevilleta NWR, New
Mexico. Fig. 3 indicates measurement of MTL.

Volume 8, Number 1, 1999

r?

3(fl)T

V

MIL

y

w

78

Journal of Hymenoptera Research

15

1.0 mm

Figs. 13-16. Neivcinn/riucx males, dorsal view of head (13, 14) and lateral view of left paramere (15, 16). 13,
15: N. nigrescens, Ash Mountain Kaweah Power Stn.#3, Tulare Co., California; 14, 16: Neivamyrmex species,
probably opncithornx, same locality.

1942: 560. First combination in subgenus Nei-
vamyrmex, but the material examined and de-
scribed by M. R. Smith was N. opncithornx not
N. californicus.

Neivami/rmex californicus v. obscurus (Forel);
Borgmeier 1953: 8.

Neivamyrmex californicus (Mayr); Borgmeier
1953:' 11.

Neivamyrmex californicus (Mayr); Borgmeier
1955: 517.

Neivamyrmex californicus (Mayr); Watkins 1972:
363 (part). Description of queen (p. 364) is
that of N. nigrescens, not N. californicus.

Neivamyrmex californicus (Mayr); Watkins 1985:
482 (part). Key (p. 482) and distribution map
(figure 4, p. 500) refer partly to N. californicus
and partly to N. nigrescens.

Worker measurements. — (n = 28). HW
0.63-1.10, HL 0.69-1.11, WL 1.03-1.67,
MTL 0.68-1.18, CI 0.86-1.00, FCI 0.033-
0.061, SI 0.68-0.80, MTI 1.05-1.14, PLI
0.78-0.88, PWI 0.62-0.73.

Worker description. — Body of moderate
size (see HW, HL and MTL measure-
ments) and somewhat compact (see plot of

WL on HW; Figs. 19, 23); head broad, CI
approaching 1.00 in largest workers, i.e.,
those in which HW and MTL > 1.00 mm;
mandible with basal margin rounding
gradually into masticatory margin (Fig. 1);
masticatory margin with a small tooth at
the terminus of this rounding, followed by
1^ denticles (tending to increase in size),
then a more prominent tooth midway
along the margin; distal portion of masti-
catory margin generally edentate (a small
denticle or two may follow the mid-point
tooth) except for the acute apical tooth; an-
terior margin of torulus separated from
anterior margin of clypeus (ignoring the
thin diaphanous clypeal apron) by about
0.2 X the diameter of the torulus; frontal
carinae moderately well separated (MFC
0.022-0.061), diverging anteriorly, and
protruding very slightly (largest workers)
or not at all (most workers) beyond the
anterior clypeal margin, when the head is
seen in frontal view; anterior clypeal mar-
gin more or less straight (weakly convex

Volume 8, Number 1, 1999

79

in smallest workers, and slightly concave
in largest workers); clypeal apron relative-
ly well developed, its anterior margin gen-
erally slightly convex or subangulate;
clypeal apron extending anteromedially
beyond the clypeal margin proper by a
distance equal to 0.4-O.5X the torulus di-
ameter; each compound eye consisting of
a single convex ommatidium, breaking the
surface of the head, its maximum diame-
ter approximately 0.06-0.08 mm (6-10% of
head width); scapes of moderate length,
exceeding the eye when held back against
the head (SI 0.68-0.80; SI2 0.67-0.75) (see
also plot of SL on HW and SI on MTL;
Figs. 18, 22, 26); posterior margin of head
usually concave, in frontal view; occipital
lobes weakly developed, not protruding
conspicuously when the head is seen in
frontal view; anterior pronotum descend-
ing gradually towards the collar, trans-
verse ridge weakly developed; lower
pronotal flange thin, lamellate, directed
more or less ventrally; dorsal profile of
mesosoma rather flat, and dorsal face of
propodeum only slightly depressed below
the level of the mesonotum (Fig. 2); dorsal
face of propodeum rounding into, and
subequal in length to, the declivitous face;
latter flat to weakly concave, in profile;
legs relatively short, MTL/HW ( = MTI) <
1.15; petiole short, high, and with short
vertical anterior face, followed by a more
or less evenly convex dorsal surface (as
seen in profile; Fig. 2), or with a slightly
steeper posterodorsal than anterodorsal
slope; in dorsal view, petiole subrectan-
gular, but with somewhat convex sides,
about 1.5 times longer than wide; antero-
ventral process of petiole a thickened
transverse shelf, in lateral view appearing
as a relatively small, blunt tooth directed
anteroventrally; a similar, less protrusive
structure at the anteroventral extremity of
the postpetiole; in dorsal view postpetiole
subtrapezoidal (Fig. 10), with straight, di-
verging sides, broadest posteriorly, and
slightly broader than long. Upper surface
of mandible finely and densely striate

with scattered punctures, subopaque, lat-
eral surface of mandible and area imme-
diately preceding the masticatory margin
smooth and shiny with scattered punc-
tures. Head largely smooth and shining,
with numerous piligerous punctures sep-
arated by several to many diameters; in
larger workers (and in more southern
populations) the punctures may be coarser
and denser, and parts of the intervening
shiny integument dulled very slightly by
weak reticulation. Mesosoma densely
punctulate, opaque, but in smaller work-
ers the sculpture weakened laterally on
the pronotum (which becomes finely retic-
ulate and sublucid) and replaced by
smooth shiny areas on the dorsum of the
promesonotum. In workers of all sizes, in-
cluding heavily sculptured large individ-
uals, the center of the mesonotum is near-
ly always smooth and shiny, with a few
larger piligerous punctures, and contrasts
with the predominantly punctulate and
opaque dorsal face of the propodeum (ex-
ceptions include some heavily sculptured
workers from southern California in
which the mesonotum center is weakly re-
ticulate and sublucid, but still contrasts
with the opaque dorsal surface of the pro-
podeum; and small, shiny workers from
northern California in which even the pro-
podeum dorsum loses its punctulate
sculpture centrally, so that contrast be-
tween the mesonotum and the propo-
deum is lessened). Petiole laterally reticu-
lo-punctate and subopaque to sublucid,
petiolar dorsum mostly shining, but with
variable traces of punctulate reticulation;
postpetiole and gaster smooth and shiny,
with scattered piligerous punctures. Long,
fine, golden pilosity conspicuous on body
and appendages (scape, funiculus, legs),
mostly suberect to subdecumbent. Body
light castaneous brown to deep reddish-
brown, tending towards a lighter yellow-
brown on the postpetiole and gaster. Man-
dible medium to dark brown, usually con-
trasting somewhat with the lighter head
color.

80

Journal of Hymenoptera Research

Queen. — Unknown. The "N. califoniicus"
queen from Davis, California, described
by Watkins (1972), is that of the shiny
form of N. nigrescens. A single queen, in
rather poor condition, from Monterey,
California (31. v. 1963; leg. Roy Johnson;
USNM), could be either N. opacithorax or
N. californiciis. It is relatively small in size
(HW 1.65, WL 2.68, MTL 1.33), with short
scapes (SI 0.46), rounded occipital lobes,
short legs (MTI 0.81), and a long gaster
(QGL/WL = 2.34). The short appendages
and elongate gaster are features that dis-
tinguish N. opacithorax queens from those
of N. nigrescens, but they might also be
characteristic of N, californiciis. There are
apparently no worker specimens associ-
ated with the Monterey queen, leaving its
specific identity uncertain.

Male. — Unknown. In LACM and CASC
there are a series of Neivamyrmex males,
collected in California but unassociated
with workers, that appear to be N. opaci-
thorax on the basis of head shape and (es-
pecially) male genitalia (Figs. 14, 16). The
proximal flange on the ventral margin of
the paramere would seem to be particu-
larly characteristic of that species. Never-
theless, until worker-associated males of
N. californiciis are discovered, one cannot
exclude the possibility that some of these
males (from Contra Costa and Tulare
Counties) represent N. californiciis.

Comments. — Workers of N. californiciis
can be distinguished from those of N. ni-
grescens by their more compact body,
shorter legs and scapes, and broader head
and petiole (Figs. 1-3). The simplest quan-
titative diagnostic is the metatibial index
(MTI = MTL/HW) which ranges from
1.05 to 1.14 in N. californiciis (n = 28) com-
pared to 1.16 to 1.52 in N. nigrescens (n =
89). A bivariate plot of the relevant mea-
surements (MTL and HW) demonstrates
the distinction (Fig. 17). Other differences
between N. californiciis and N. nigrescens
are captured by plots of scape length (SL)
on head width (HW) and Weber's length
(WL) on head width (Figs. 18, 19, 21, 22).

When samples from all populations of N.
nigrescens are considered (i.e., including
workers from the Southwest and from
eastern United States) there is a slight
overlap in the distribution of points (Figs.
18, 19); but when confining the compari-
sons to populations of N. nigrescens that
are sympatric with N. californiciis (i.e.,
populations from California) these and
other bivariate plots produce non-overlap-
ping clouds of points (see Figs. 21-28). Be-
cause of allometry, neither the cephalic in-
dex (CI = HW/HL) nor the scape index
(SI = SL/HW) are diagnostic in them-
selves, but when plotted against the me-
tatibia length (MTL) a sharp distinction is
seen between the two species (Figs. 25-
26). Thus, within a given size class (as
measured by MTL) there is no overlap in
CI or SI; but small workers of N. califor-
niciis have scape and cephalic indices that
overlap with large workers of N. nigres-
cens.

Workers of N. californicus also have the
dorsal face of the propodeum less de-
pressed below the level of the mesonotum
than in N. nigrescens (compare Figs. 2 and
5), and this difference is diagnostic. Less
tangible are differences in body sculpture:
the dense punctulate sculpture that is so
prominent in most populations of N. ni-
grescens and that imparts a granular ap-
pearance to the head and mesosoma is
much less developed in N. californicus,
such that the head, mesonotum, and post-
petiole are largely smooth and shiny (cov-
ered only with scattered piligerous punc-
tures). But a reliance on sculptural differ-
ences led to the past confusion between
these two species: populations of N. ni-
grescens that are broadly sympatric with
N. californicus in central and northern Cal-
ifornia also have weakened body sculp-
ture and are superficially similar to those
of N. californicus. On average, N. californi-
cus is still the shinier of the two (see dis-
tinctions in key couplet 9A below) but the
differences are subtle. The contrasts in
body shape and leg length documented

Volume 8, Number 1, 1999

81

17

■e t.2

E,

18

■ califomlcus
A opacrthorax
O nigrescens
o texanus

«gp

05 06 07 08 09 1 11 12 13 14 15
HW(mm)

04 05 06 07 08 09 1 11 12
HW(mm)

13 14 15

19

IS

t 16
E,
_J
5 14

12

o

■ califomicus

A opacrthorax

□ 0

;

D nigrescens
0 texanus

o o

D

:

^^■'^

: #

■

.t

A

20"

■

■ califomicus

^ opacrthorax

■ ■■

'•■"■.

A

0:^5

<" • A

^^ a'^

■

■

02

'

zA

I

A

^A

■

0 15

A

04 05 06 07 08

09 1
HW(mm)

II 12 13 14 15

04 05 06

07 0 8 0 9 1 11 12
HW(mni)

21'

1 L>

■ califomicus

1 b

- o nigrescens (CA)

D

- a

- a

P a a

a

D °

a ° f ■

■

09

08

07

■

22'

0 5 0 6 0 7 0 8 0 9
HW(mm)

■ calrfomicus
n nigrescens (CA)

0

;

OdoP

D *

■

D

■

1 1 12

05 06 07 08 09 1 11 1.2
HW(lTim)

Figs. 17-22. Bivariate plots of various metric measurements in workers ot tour species of Neivamyniifx.

82

Journal of Hymenoptera Research

l

3„

■ califomicus
D nigrescens (CA)

P
P

q, a
T] a a

5 14

: o

;

B

A '

; B.

24'

■ calffomicus
D nigrescens (CA)

Q

" G° °

G ■
G G

: °G s

G P

G a

G CP^^GJ"-

a ■

B ° ■

06 07 08 09 1 11 12

HW(mm)

05 06 07 08 09 1 11 1,2

HW(mm)

25

5 09

■

■ califoniicus
D nigrescens (CA)

;

■

■
■ €

a ; S ^

G G
G

D

■;■ ' G

a G

^ p°

D G
P P
n ° P
ft" ^C?° GP

P P

26

G ^

G
0

P °
D

p p °

D nigrescens (CA)

095

0

°D

n

G

09

4d

D

f^"*; °

P r, °

085

°G

0° G °

08

t a

° P

"...

° P

075

■\

1 ■

cb

a

■ ^

Of

s

a

a

0,7 0,8 0.9 1 11 12 1.3 1.4 1.5 16

MTL (mm)

0,6 07 08 09 1 11 12 13 14 1.5 1.6

MTL (mm)

27'

■ califomicus
D nigrescens (CA)

a

-

■

■
■ I

■•''1

CD

■ .' ° □ □

a a □
f *

D D

28

(^ -

: p Q
- p

^,«

■ calrfomicus

: G

: G G

D nigrescens (CA)

14

1 35

: pP „a °

i GO ""f^
r on G dWaG,

P 125

S 1 2

: 0

1 15
1 1
105

■

■«V;.-

06 07 08 09 1 11 12 13 14 15 16
MTL (mm)

0 16 Olfl 0 2 0 22 0 24 0 26 0 28 0 3 0 32

OPW/MTL

Figs. 23-28. Bivariate plots of metric measurements and indices in workers of Nctzuiw\fnm'X culifonucui^ and
N. nigrescens (California populations only).

Volume 8, Number 1, 1999

29

83

; O nigr»scens (CA, punctate form)
■ T nigrescens (CA, shiny form)

0

D

D

▼

30

0 5 0 6 0 7 0 8 0 9

W 07

.

: D nigrescens (CA. punctate form)
■ T nigrescens (CA. shiny form)

a

C
D

T

: o

D »»

\

T

a o

T

0.5 06 07 Oe 09

HW{mm)

31

u ^

A texanus

D nigrescens {SW. East)

A

015

A

A

A

01

A
^ A

1 0 D

0.05

AD Do D
i^ □ D

0 D

D

32'

A texanus
. D nigrescens (SW, East)

A

AA

* A

A

A \ A O
A A* tU

A ^O °'°

0.4 0.5 0 6 0 7 0 8 0 9 1 11 12 13 1.4 1.5
HW(mm)

0.6 0-8

12 1,4 1,6 1.8

o

J>

1

1 A texanus

D nigrescens (SW. East)

A

0 15

lo,

005

A

A

A

A

^ A ^ O^

i a rp °

° DO

3

34

UD

A lexanus

0 55

D nigrescens (SW, East)

A
A

'.

OS

:

A
A *°

0 45

"

04

A

to

■S"

ACA°

y

n*

/.

3

025

□

05 06 07 08 09 1 11 12 13 14 15

HL (mm)

05 06 07 08 09 1 11 12 13 14 15

HL (mm)

Figs. 29-34. Bivariate plots of metric measurements and indices in workers of Ncivntmfrmex tcxauus and
various populations oi N. fii<^resce}is.

84

Journal of Hymenoptera Research

above are niore reliable for distinguishing
these two species.

N. californkus differs more obviously
from N. texanus. Like N. nigrescens, work-
ers of N. texanus have longer legs (MTI
1.18-1.48), longer scapes (SI2 0.75-0.89),
and a more gracile body than those of N.
californiciis. They also average larger in
size, have more widely separated frontal
carinae (FCI 0.066-0.125 compared to
0.033-0.061 in N. californiciis), and have a
consistently opaque, densely punctulate
head and mesosoma such that they are
unlikely to be confused with N. californicus
workers. As far as known, the ranges of
the two species do not overlap (Fig. 36).

The bivariate plots of measurements
(Figs. 17-19) show that in many respects
N. californicus is more similar to N. opaci-
thorax than to N. nigrescens or N. texanus.
N. opacithorax can be distinguished from
N. californicus by the angular basal margin
of the mandible; shorter clypeal apron;
narrower petiole (Fig. 20); and different
pattern of body sculpture (side of prono-
tum usually smooth and shiny, and con-
trasting with the rugulose-punctulate me-
sosoma dorsum).

Material examined. — (CASC, CDAE,
JTLQ LACM, MCZC, MHNG, NHMV,
RAJC, UCDC, USNM)

MEXICO Baja California: 28km E Ensenada, 750m
(P. S. Ward).

UNITED STATES California Cpntrn Costa Co.: 9km
ENE Danville, 490m (P. S. Ward); El Dorado Co.: 14km
NW Shingle Springs, 340m (P. S. Ward; G. C. Snell-
ing); Los Angeles Co.: Arcadia (c.u.); La Verne (A. C.
Oberle); Los Angeles (A. Mallis & ]. Schwartz); Mon-
terey Co.: Salinas (B. Oliver c& J. Bunch); Orange Co.:
Irvine Park (A. Mintzer); Laguna Hills (R. J. Hamton);
Limestone Canyon, El Toro Rd., I.8mi E Cooks Cor-
ner (A. Suarez); Tonner Canyon (W. P. Mackay); Riv-
erside Co.: Riverside (K. Cooper; K. W. Cooper; E. I.
Schlinger); Temecula (A. Suarez); San Bernardino Co.:
Chino Hills State Park (G. C. Snelling et al.); Chino
Hills State Park, 700 ft. (R. A. Johnson); San Diego Co.:
5mi. NE Poway, 600m (M. S. Trepanier); Camp Pen-
dleton (J. H. Hunt); Chula Vista, E end, 160m (P. S.
Ward); Chula Vista, 70m (A. Suarez); Elliott Reserve,
150m (A. Suarez); La Jolla (c.u.); Mt. Laguna, MSP
site, 6050 ft. (J. H. Hunt); National City (K. Ross); San
Diego (c.u.); Vista (E. Hindle); San Francisco Co.: San

Francisco (Schaufuss); San Mateo Co.: Jasper Ridge,
150m (K. G. Human; N. J. Sanders); Santa Barbara Co.:
9km N Goleta, 490m (P. S. Ward); Cachuma Saddle,
1100m (J. Longino); Santa Clara Co.: 11km S Palo Alto,
490m (P. S. Ward); Santa Cruz Co.: Santa Cruz (K.
Brown).

Neivamyrtnex nigrescens (Cresson 1972)

(Figs. 4-9, 11-12)

Lahidiis nigrescens Cresson 1872: 194. Holotype

male, Bosque Co., Texas (Belfrage) (ANSP)

[not examined].
Eciton nigrescens (Cresson); Dalla Torre 1893: 5.
Eciton (Acamatus) schmitti Emery 1894: 183. Syn-

type workers, Doniphan, Missouri (Pergan-

de) (MSCN) [not examined], MCZC, USNM

[examined]. Synonymy by M. R. Smith 1938:

160.
Eciton (Labidus) nigrescens (Cresson); Emery

1895: 258.
Eciton (Acnrnntus) nigrescens (Cresson); Emery

1900: 187.
Eciton sitmichrnsti; Wheeler (nee Norton) 1900:

564. Description of queen (as "£. sumichras-

ti").
Eciton (Neivnmi/rmex) nigrescens (Cresson);

Smith 1942: 550 (part). Description of worker

(part), queen and male.
Eciton (Neivamyrmex) californicum; Creighton

(nee Mayr) 1950: 70 (part).
Neivamyrmex nigrescens (Cresson); Borgmeier

1953: 6.
Neivamyrmex nigrescens (Cresson); Borgmeier

1955: 494 (part). Description of worker (part)

and queen. Male (p. 496) is that of N. texanus

(Watkins 1972).
Neivamyrmex sp. e; Borgmeier 1955: 531. De-
scription of male.
Neivamyrmex nigrescens (Cresson); Watkins

1972:' 358.
Neivamyrmex californicus; Watkins (nee Mayr)

1972: 363 (part). Description of queen (as "N.

californicus").
Neivamyrmex nigrescens (Cresson); Watkins

1985:482.
Neivamyrmex californicus; Watkins (nee Mayr)

1985: 482 (part).

Worker tneasiirenients. — (n = 89). HW
0.60-1.31, HL 0.71-1.31, WL 1.10-2.12,
MTL 0.76-1.76, CI 0.80-1.00, FCI 0.030-
0.106, SI 0.75-1.01, MTI 1.16-1.52, PLI
0.68-O.91, PWI 0.52-0.71.

Volume 8, Number 1, 1999

85

Worker diagnosis. — Moderately large
body size (see HW, HL and MTL mea-
surements); mandible with basal margin
rounding gradually into masticatory mar-
gin (Figs. 4, 7); frontal carinae moderately
well separated (MFC 0.021-0.138); clypeal
apron well developed and, in all but the
largest workers, produced anteromedially
by an amount subequal to, or greater than,
the minimum distance between the frontal
carinae (MFC); scapes relatively long, SI >
0.69 (see also plot of SL on HW and SI on
MTL; Figs. 18, 22, 26); occipital lobes
weakly to moderately developed; anterior
margin of pronotum with transverse ridge
generally well developed; dorsal (= basal)
face of propodeum conspicuously de-
pressed below the level of the promesono-
tum (Figs. 5, 8), and rounding into the de-
clivitous face, the latter more or less flat
(or weakly concave) in profile; legs rela-
tively long, MTL/HW ( = MTI) > 1.15; pet-
iole subrectangular, somewhat variable in
shape (see PLI and PWI values), but al-
ways longer than high or wide (Figs. 5, 8,
11, 12). Head and mesosoma typically
densely punctulate, and having an
opaque, granular appearance; populations
from northwestern portions of the species'
range, however, have the sculpture much
weakened such that the head is partly
smooth and shining, with scattered pili-
gerous punctures and variable amounts of
finer reticulate sculpture that partly dulls
the sheen; and in this "shiny morph" the
mesosoma is partly sublucid, although
with at least weak reticulate-punctulate
sculpture on most surfaces. In all popula-
tions sculpture weakened on the surface
of metapleural gland bulla, such that the
lower half or more is smooth (or weakly
reticulate) and conspicuously shiny. Peti-
ole densely punctulate, subopaque, post-
petiole tending to be more lightly sculp-
tured. Body varying from light castaneous
brown to dark reddish-brown, the post-
petiole and gaster usually lighter than the
rest of body.

Queen diagnosis. — Eye distinct, consist-

ing of single convex ommatidium. Head
as broad as, or slightly broader than, long
(CI 0.96-1.02, n = 6). Occipital corners
generally angular and projecting, but may
be weakly angulate or rounded. Pronotum
without posterior dorsolateral projection.
Propodeum (and sometimes also meso-
notum) with a median longitudinal im-
pression. Metatibial index (MTI) 0.89-1.07
(n = 6). Petiole subquadrate, slightly
broader than long (PWI approximately
1.1-1.2), with a vertical anterior face that
rounds into a flat dorsal face, as seen in
lateral view. Petiole width much less than
the length of the metatibia, DPW/MTL
0.58-0.72 (n = 6). Length of gaster (non-
physogastric) less than twice WL (QGL/
WL = 1.70-1.90, n = 5).

Male diagnosis. — Mandibles broad, spat-
ulate (not sickle-shaped). Ocelli moderate
in size and separated from the upper mar-
gin of the compound eye by a distance
greater than twice the diameter of the me-
dian ocellus. Prominent transverse swell-
ing above antennal fossa, discerned most
clearly in dorsal view (Fig. 13). Setae on
venter of petiole typically short, whitish,
and slanted posteroventrally, but in some
western populations the setae are longer,
golden, and suberect. Gaster typically
black or blackish-brown, often reddish-
brown in western populations. In profile,
paramere (= stipes) linear subrectangular,
with an oblique (anterodorsal to postero-
ventral) posterior margin, and a straight
ventral margin, not produced anteroven-
trally (Fig. 15); posterodorsal extremity
with a low, triangular projection (Watkins,
1985, Plate 9, fig. 4), tending to become ob-
solete in western populations (Fig. 15);
volsella unforked, with long, slender up-
turned apex; aedeagus ( = sagitta) with an
apically upturned posteroventral process,
just exceeding the posterodorsal process
(in posterior reach).

Comments. — The characteristics that dis-
tinguish workers of N. nigrescens from
those of N. californicus and N. texanus are
discussed under those respective species.

86

Journal of Hymenoptera Research

N. nigrescens differs from N. opacithorax by
the shape of the worker mandibles, by the
heavier body sculpture, especially on the
side of the pronotum, by the longer
scapes, mesosoma and legs (Figs. 17-19),
and by the more strongly convex profile
of the promesonotum.

A remaining question concerns the sta-
tus of the nigrescens-Mke populations with
shiny worker heads, that are superficially
similar to those of N. californicus. This
"shiny morph" of N. nigrescens is found in
north-central California, parts of the Great
Basin, and in upper sections of the Colo-
rado River drainage, while the more "typ-
ical" form, with densely punctulate and
opaque worker head, is distributed widely
from southern California, Arizona and ad-
jacent regions of Mexico to southeastern
United States (Fig. 37). What happens in
zones of contact between the two forms?
Records are too sparse to answer this
question for the Colorado River basin, but
collections from southern California reveal
a zone of intergradation centered on the
north side of the San Gabriel Mountains.
In fact patterns of sculpturation are inter-
mediate in samples taken from this region,
so that the distinction between the two
"forms" becomes quite arbitrary. For this
reason it seems clear that they must be
treated as conspecific, although the pat-
terns of distribution are suggestive of a
previous period of isolation followed by
secondary contact and introgression.

Material examined. — (1) Typical form
with opaque, densely punctulate head
(CASC, CDAE, LACM, MCZC, RAJC,
UCDC, USNM)

MEXICO Baja California: 28km E Ensenada, 750m
(P. S, Ward); 31.7mi WNW Bahia de los Angeles
(Hardy, Andrews & Giuliani); 6mi SE Laguna Cha-
pala (A. E. Lewis); Baja California Sur: 12mi S Santa
Rosalia (Michelbacher & Ross); 15mi S San Domingo
(Ross & Bohart); 20mi W La Paz (E. L. Sleeper); 26km
NW Santa Rosalia (R. A. Johnson); Coyote Cove, Con-
ception Bay (Michelbacher & Ross); Isla San Jose, Imi
S Punta Colorada [as "Punta Colorado"] (J. T. Doy-
en); San Hilario (E. L. Sleeper); San Ignacio, 140m (M.
Bennett); Jalisco: 3mi SE Plan de Barrancas (F. D.

Parker &. L. A. Stange); Nayarit: hlas Tres Marias: Isla
Cleofas (R. R. Snelling); Isla Magdalena (R. R. Snell-
ing); San Luis Potosi: Cd. Valles, El Bariito (J. F. Wat-
kins); Sonora: 2km N Bahia de la Cruz, Isla Tiburon,
10m (P. S. Ward); 2km SW Punta Narragansett, Isla
Tiburon, 5m (P. S. Ward); 37mi N Hermosillo, 1700
ft. (R. R. Snelling); 5mi S Cananea (V. D. Roth); 8km
N Desemboque de los Seris (R. A. Johnson); Los Hor-
cones, Rte. 16, 4km E La Colorada (B. Bestelmayer);
Tamaulipas: Cd. Victoria (J. F. Watkins).

UNITED STATES Alabama: Dallas Co.: Selma (W.
H. Patton); Jefferson Co.: Birmingham (R. D. Jordan);
Lauttcrdale Co.: Florence (W. Cloyd; F. Moore);
Latvrence Co.: King Cove, Bankhead Natl. Forest (E.
O. Wilson); Mobile Co.: Kushla (A. C. Sturtevant);
Spring Hill [as "Springhill"] (W. S. Creighton); Mor-
gan Co.: Decatur (Murphree); Arizona: Cochise Co.:
2mi NE Portal (G. D. Alpert); Chiricahua Mts. (W. &
E. MacKay; J. F. Watkins); Chiricahua Mts., 5000 ft.
(J. F. Watkins); Chiricahua Mtns., 13mi NW jet. Rte.
80 on FSR 74, 5850 ft. (S. P. Cover); Chiricahua Mtns.,
Cave Creek Canyon, SW Res. Stn., 5400 ft. (S. P. Cov-
er); Copper Canyon, 8. Imi SE Sunnyside, 5900-6000
ft. (R. R. Snelling); Copper Canyon, Huachuca Mtns.,
22km SSW Sierra Vista, 1770m (S. G. Brady); Douglas
(W. W. Jones); Huachuca Mts., Miller Canyon (A. E.
Lewis); Miller Canyon, Huachuca Mtns., 6000 ft. (W.
M. Wheeler); Palmerlee, Huachuca Mtns., 5300 ft. (W.
M. Wheeler); Paradise Rd., 1.3mi W Portal Rd. Chir-
icahua Mts. (G. C. Snelling); Portal (Gotwald); Ram-
sey Canyon, Huachuca Mts. (W. S. Creighton); Ram-
sey Canyon, Huachuca Mts., 5800 ft. (W. M. Mann);
SWRS, Portal, 5600 ft. (W. S. Creighton); Texas Pass,
Dragoon (as "Dragon"] Mtns. (W. M. Wheeler); Gila
Co.: Pinal Mts., 8000 ft. (R. A. Flock); Sierra Anchas,
Hwy. 288 at Exp. Res. Stn., 4800 ft. (R. A. Johnson);
Maricopa Co.: Four Peaks Wilderness, nr. Pidgeon
Springs, 5600 ft. (R. A. Johnson); Mazatzal Mtns., on
Four Peaks Rd., 10.3mi E Hwy. 87, 4000 ft. (R. A.
Johnson); South Phoenix Park (P. S. Ward); Mojave
Co.: Hualapai Mts., s. of Kingman, 1450m (E. 1. Schlin-
ger); Pima Co.: Baboquivari Mtns, Forestry Cabin,
3500 ft. (W. S. Creighton); Buehman Canyon, Santa
Catalina Mts., 2900-3000 ft. (R. R. Snelling & G. C.
Snelling); Santa Catalina Mts. (M. Chrisman); Tucson
(R. H. Crandall); Piiml Co.: Oracle, 4500 ft. (W. M.
Wheeler); Sniita Cruz Co.: 6.7mi W 1-19 on Ruby Rd.
(G. C. Snelling); Bog Springs Cpgrd., Madera Canyon,
Santa Rita Mts. (G. C. Snelling); Madera Canvon (R.
H. Crandall); Madera Canyon, Santa Rita Mts. (R. H.
Crandall); Nogales (Ehringer); Pajarito Mtns., Ruby
Rd., 6.7mi W I-IO, 4000 ft. (R. A. Johnson); Pena Blan-
ca Lake (B. V. Brown & D. H. Feener); Ruby Rd., 7mi
W Pefia Blanca (G. C. Snelling); Yavapai Co.: 7.2mi E
Chino Valley, 4600 ft. (R. A. Johnson); Yuma Co.: Bur-
ro Cyn., 2mi SE jet. 24, Kofa Game Refuge (P. Melhop
& R. R. Snelling); California: li'.-^ .Angeles Co.: Clare-
mont (B. Crow); E fork, San Gabriel R., Angeles N. F.

Volume 8, Number 1, 1999

87

(C. Ishida); Eaton Canyon (Sutton?); Eaton Canyon
I'k. (R. H. Crandall); Jet. Angeles Crest & Angeles
Forest Hwys. (G. C. Snelling); Millard Canyon, San
Gabriel Mts. (R. H. Crandall); Placerita Canyon Park
(F. T. Hovore); Orange Co.: Tonner Canyon (W. P.
MacKay); Rivcr^ulc Co.: Camino Resales {A. Suarez);
Margarita Summit (A. Suarez); near Perris (Mallis,
Zschokke & Schwartz); Pinyon Flat, 1220m (P. S.
Ward); Riverside (K. W. Cooper; M. E. Irwin); San
Timoteo Cyn. (M. Wasbauer & A. Hardy); Temecula
Cyn., Sta. Margarita R. (E. I. Schlinger); San Bernar-
dino Co.: 2mi E Mentone (W. S. Creighton); Aliso Cr.,
Chino Hills State Park (M. Bennett et al); Chino Hills
State Park, 700 ft. (R. A. Johnson); San Diego Co.: 18km
E Mt. Laguna, 300m (P. S. Ward); 5mi N Descanso,
MSP primary site, 3000 ft. (J. H. Hunt); 5mi. E La Jolla
(M. S. Trepanier); Chula Vista (E end), 160m (P. S.
Ward); La Jolla (M. S. Trepanier); La Mesa (F. X. Wil-
liams); Nate Harrison Rd. nr. Mt. Palomar, 4800 ft. (E.
I. Schlinger); nf. La Mesa (F. X. Williams); Point Loma
(P. Leonard; A. Suarez); Ramona, 450m (M. S. Tre-
panier); San Diego (c.u.); Colorado: Chaffee Co.: Salida
(C.U.); Salida, 7050 ft. (W. M. Wheeler); Georgia: Jnek-
son Co.: Commerce (Vanderford); Illinois: Adama Co.:
Quincy (T. E. Musselman); lohnson Co.: Feme Clyffe
St. Pk. (W. S. Creighton); Iowa: Harrison Co.: Little
Sioux (W. F. Buren); Woodbiny Co.: Sioux City (C. N.
Ainslie; W. F. Buren); Kansas: Douglas Co.: Lawrence
(F. X. Williams); Nat. Hist. Reser., Lawrence (C. W.
Rettenmeyer); Harvey Co.: Sedgwick [as "Sedwick")
(A. J. McCurray); Jefferson Co.: Valley Falls (A. Mattis);
McPherson Co.: McPherson (W. Knaus); Pottawatomie
Co.: Onaga (F. F. Crevecoeur); Rdei/ Co.: Jardine Terr.
0- F. Watkms); Manhattan (A. J. Mattis; R. C. Smith);
no specific locality (F. Marlatt; J. B. Norton; J. F. Wat-
kins); Sedgwick Co.: Wichita (J. R. Horton); Kentucky:
Marshall Co.: Kentucky Dam (W. L. Brown); Louisi-
ana: Acadia Co.: Crowley (C. E. Hood); Beauregard Co.:
DeRidder (W. F. Buren); £i!s( Baton Rouge Co.: Baton
Rouge (T. H. Jones); Lafai/ette Co.: Lafayette (E. S.
Tucker); Madison Co.: Tallulah (McGehee); Plaquemi-
nes Co.: Buras y. R. Horton); Naomi [as "Naomie")
(c.u.); Rapides Co.: Alexandria (E. S. Tucker); Missis-
sippi: Adams Co.: Natchez (G. W. Alexander); Clarke
Co.: Quitman (Murphree); Clay Co.: Cedar Bluff (c.u.);
Cedar Bluff, Trimcane (G. W. Haug); Harrison Co.:
Landon (Murphree); Humphreys Co.: Belzoni (Mur-
phree); lones Co.: Laurel (M. R. Smith); Lozvndes Co.:
Columbus (Murphree); Monroe Co.: Aberdeen (Mur-
phree); Oktibbeha Co.: Maben (L. C. Murphree); Stark-
ville (W. W. Love; M. R. Smith); State College [as
"Agr. Col. Miss."] (M. R. Smith); Stone Co.: Bond
(Murphree); Wiggins (Murphree); Washington Co.:
Greenville (G. L. Snodgrass); Missouri: Boone Co.: Co-
lumbia (A. C. Cole; L. Haseman; M. Talbot); Butler
Co.: Poplar Bluff (D. E. Read); Cape Girardeau Co.:
Cape Girardeau (D. E. Read); Cole Co.: Jefferson City
(A. C. Burrill); lohnson Co.: Knob Noster State Park

(M. B. & J. R. DuBois); Ripley Co.: Doniphan (Pergan-
de); Sf. Charles Co.: St. Charles (M. Talbot); St. Louis
Co.: Webster Groves [as "Webster Grove") (G. Loef-
fel); New Mexico: Colfax Co.: Cimarron Canyon, Ci-
marron (A. C. Cole); Dona Ann Co.: 45km NE Las Cru-
ces (W. MacKay); University Ranch (C. A. Kay); Grant
Co.: 5km NW Silver City, 1900m (P. S. Ward); 1-10,
3mi E Separ (R. A. Johnson); Hidalgo Co.: 4km N Ro-
deo, 1250m (P. S. Ward); San Simon Valley, 0.25mi W
jet. Rte. 80 on Portal Rd. (NM533), 4250 ft. (K. Helms);
San Simon Valley, 0.3mi NE jet. State Line Rd. & Rte.
533, 4250 ft. (S. P. Cover); San Simon Valley, 0.5mi W
jet. Rte. 80 on Portal Rd. (NM533), 4250 ft. (D. Gor-
don); San Simon Valley, jet. State Line Rd. & Portal
Rd. (NM533), 4250 ft. (S. P. Cover); Santa Fe Co.: Santa
Fe (W. M. Mann); Santa Fe, 0.5mi N 1-25 on Cerillos
Rd., 6200 ft. (S. P. Cover); Sierra Co.: Hillsboro, 1600m
(P. S. Ward); Socorro Co.: Sevilleta NWR (M. Kaspari);
Torrance Co.: lOmi S Mountainair, 6650 ft. (A. C.
Cole); Union Co.: Clayton (W. M. Wheeler); North
Carolina: Swain Co.: Great Smoky Mts. Natl. Park,
5500 ft. (E. S. Ross); Oklahoma: Cimarron Co.: Kenton
(T. H. Hubbell); Kay Co.: Ponea City (A. C. Burrill);
Tennessee: Chester Co.: Henderson (Murphree); Da-
vidson Co.: Nashville (W. S. Creighton; A. R. Laskey);
near Nashville (L. Wesson); Hazokms Co.: Rogersville
(W. S. Creighton); Knox Co.: Knoxville (A. C. Cole);
McMinn Co.: Athens (Murphree); Monroe Co.: no spe-
cific locality (Jones); Shelby Co.: Memphis (Murphree);
Wayne Co.: Clifton (Murphree); Texas: Bell Co.: Bow-
mer Ranch (J. F. Watkins); Be.xar Co.: 20mi S San An-
tonio (E. S. Ross); San Antonio (E. S. Ross); Calhoun
Co.: Port Lavaca (McGehee); Dallas Co.: Dallas (F. C.
Bishop; E. W. Laake; W. D. Pierce; Vanderford);
Grimes Co.: Shiro (W. Buren); Hall Co.: 6mi SE Turkey
(C. W. O'Brien); Harris Co.: Houston (H. C. Milleri-
der); Jeff Davis Co.: Fort Davis State Park (J. F. Wat-
kins); McLennan Co.: Waco (R. S. Baldridge; J. F. Wat-
kins); Montgomery Co.: Willis (J. C. Bridwell); Travis
Co.: Austin (W. M. Wheeler); Val Verde Co.: Del Rio
(c.u.).; West Virginia: Mason Co.: West Columbia
(Murphree).

(2) Form with shiny head (CASC,
CDAE, JTLC, LACM, MCZC, UCDC,
USNM)

UNITED STATES California: Amador Co.: 9km
WNW Plymouth, 200m (P. S. Ward); Colusa Co.: 1km
W Fouts Springs, 600m (P. S. Ward); CoHfra Costa Co.:
Danville [as "Dannville"] (F. X. Williams); El Dorado
Co.: 14km NW Shingle Springs, 340m (P. S. Ward);
9km SW Pilot Hill, 340m (P. S. Ward); Kern Co.: Lone-
tree Cyn., 6.8mi S jet. Randsburg [as "Randsbury")
Rd. & Hwy 14 (F Andrews & M. Wasbauer); Lake
Co.: 14km ENE Lower Lake, 290ni (P. S. Ward); 19km
ESE Lower Lake, 700m (P. S. Ward); 20km ESE Lower
Lake (B. L. Fisher); 6km NW Middletown, 490m (P.
S. Ward); Nice-Bartlett Sprgs (A. Andrasfalvy); Los

88

Journal of Hymenoptera Research

Angtiesi Co.: Los Angeles (A. Mallis); Pearblossom
Hwy & Barrel Spr. Rd. (G. C. Snelling); Mcndocnw Co.:
Hopland Field Stn., 240m (P. S. Ward); Monterey Co.:
10km SSW Jolon, Fort Hunter Liggett MR, 340m (P.
S. Ward); 14km SW Jolon, Fort Hunter Liggett MR,
640m (P. S. Ward); 15km SW Jolon, Fort Hunter Lig-
gett MR, 490m (P. S. Ward); Paraiso Springs (c.u.);
Napa Co.: 5km ENE Rutherford, 120m (P. S. Ward);
5km W Oakville, 560m (P. S. Ward); N. side Howell
Mtn., 3km NNE Angwin, 3%m (H. B. Leech); Placer
Co.: 2km E Colfax, 490m (P. S. Ward); RiversiLte Co.:
Red Cloud Mine, Chuckwalla Mts., 2700 ft. (G. C.
Snelling); San Benito Co.: 16.8mi N New Idria (A. J.
Gilbert & N. Smith); San Bernardino Co.: Cima (c.u.);
San Luis Obispo Co.: 19km SSE California Valley, Car-
rizo Plain Natural Area, 800m (P. S. Ward); 2.5mi. S
Arroyo Grande (G. I. Stage); 20km ESE California
Valley, Carrizo Plain Natural Area, 800m {P. S.
Ward); Santa Barbara Co.: Arroyo Burro, 800m (J. Lon-
gino); Cachuma Saddle, 1100m (J. Longino); Cachu-
ma Saddle, Los Padres N. F., 930m (P. S. Ward); N
end Sedgwick Ranch, 610m (P. S. Ward); N end Sedg-
wick Ranch, 730m (P. S. Ward); near top of Las Cru-
ces Mts (Heath); Solano Co.: 2km SE Mt. Vaca, 680m
(P. S. Ward); Cold Canyon, 19km NNW Vacaville,
120m (D. M. Olson; P. S. Ward); Cold Canyon, 19km
NNW Vacaville, 300m (P. S. Ward); Cold Canyon,
19km NNW Vacaville, 360m (P. S. Ward); Cold Can-
yon, 19km NNW Vacaville, 420m (P. S. Ward); Cold
Canyon, 19kni NNW Vacaville, 600m (P. S. Ward);
Pleasants Ridge, 530m (P. S. Ward); Sonoma Co.: 1km
NNE Sonoma, 170m (P. S. Ward); 3km N Sonoma,
290m (P. S. Ward); 8km NNW Cazadero, 300m (P. S.
Ward); Pepperwood Ranch, 15km N Santa Rosa,
360m (P. S. Ward); Stanislaus Co.: Del Puerto Canyon,
18km WSW Patterson, 300m (P. S. Ward); Tulare Co.:
Ash Mtn. Kwh Pwr Stn. #3 (J. A. Halstead); Tuolumne
Co.: 2mi SE jet. Hwy. 49 & 120, 1840 ft. (G. C. Snell-
ing); Yolo Co.: 6km SW Winters, 45m (D. A. Holway);
8km SE Davis, lOm (P. S. Ward); Davis (J. J. DuBois;
A. Mallis); Woodland (E. I. Schlinger); Colorado:
Montezuma Co.: Mesa Verde N. P., 6300 ft. (E. V.
Gregg); Nevada: U/on Co.: East Walker River, 26km
SSE Yerington, 1460m (P. S. Ward); Weeks, 1280m (P.
S. Ward); Utah: Millard Co.: White Sage Valley [as
"White Valley"! (R- W. Fautin); Washington Co.:
Springdale, 0.25mi S Zion Natl. Park (G. C. Snelling).
Note. Workers from sites in Los Angeles County,
California are intermediate in sculpture between the
two forms.

Neivamyrmex texanus Watkins 1972

Ncivainynncx texanus Watkins 1972: 353. Holo-
type male, Austin, Texas (W. M. Wheeler)
(MCZC) [examined].

Neivamyrmex itigrcscens; Watkins 1972: 358
(part).

Worker tneasiiretnents. — (n = 22). HW
0.71-1.42, HL 0.81-1.38, WL 1.27-2.26,
MTL 0.87-1.68, CI 0.83-1.06, FCI 0.066-
0.125, SI 0.70-0.93, MTI 1.18-1.48, PLI
0.78-0.96, PWI 0.58-0.74.

Worker diagnosis. — Rather large body
size (see HW, HL and MTL measure-
ments); mandible with blunt basal tooth in
largest workers, such a tooth becoming in-
distinct to absent in smallest workers; bas-
al margin of mandible rounding into mas-
ticatory margin; frontal carinae well sep-
arated (MFC 0.047-0.168); clypeal apron
conspicuous but less well developed than
in nigrescetis; scapes relatively long, SI >
0.69 (see also plot of SL on HW; Fig. 18);
occipital lobes generally well developed;
anterior margin of pronotum with trans-
verse ridge well developed; dorsal face of
propodeum conspicuously depressed be-
low the level of the promesonotum, and
forming a subangulate juncture with the
declivitous face, the latter more or less
concave in profile; legs relatively long,
MTL/HW ( = MTI) > 1.15; petiole subrect-
angular, variable in shape (see PLI and
PWI values), always longer than high or
wide, although generally shorter and
higher than in iiigrescens. Head, mesoso-
ma, petiole and postpetiole densely punc-
tulate, and having an opaque, granular
appearance. Most of the surface of meta-
pleural gland bulla densely punctulate
and opaque, although sometimes with a
very small, thin shiny strip immediately
above the flange of the metapleural gland
orifice. Body varying from dark reddish-
brown to blackish-brown, the gaster and
legs lighter.

Queen diagnosis. — Eye distinct, consist-
ing of single convex ommatidium. Head
slightly broader than long (CI 1.05-1.12, n
= 5). Occipital corners rounded. Prono-
tum with a distinct posterior dorsolateral
projection (Watkins, 1972, fig. 11). Meso-
notum and propodeum without a median
longitudinal impression, but propodeum
with a shallow longitudinal concavity.
Metatibial index (MTI) 0.81-0.90 (n = 5).

Volume 8, Number 1, 1999

Petiole transverse, markedly broader than
long (PWI approximately 1.2-1.7) and, in
lateral view, with a single convex antero-
dorsal face. Petiole width only slightly less
than the length of the metatibia, DPW/
MIL = 0.75-0.91 (n = 4). Length of gaster
(non-physogastric) approximately twice
WL (QGL/WL = 1.88-2.25, n = 3).

Male diagnosis. — Mandibles broad, spat-
ulate (not sickle-shaped). Ocelli moderate
in size and separated from the upper mar-
gin of the compound eye by a distance
greater than twice the diameter of the me-
dian oecllus. No prominent transverse
swelling above antennal fossa. Setae on
venter of petiole long, golden, erect or
suberect. Gaster reddish-brown. In profile,
paramere (= stipes) slender, with a trun-
cate posterior margin, and a tall, angular
posterodorsal projection (Watkins 1985,
Plate 9, fig. 5); volsella unforked, with
long, slender upturned apex; aedeagus
( = sagitta) with straight posteroventral
process, just exceeding the posterodorsal
process (in posterior reach).

Comments. — Despite the distinctive
males and queens of this species, workers
of N. texaniis are difficult to distinguish
from those of N. nigrescens. As Watkins
(1972, 1985) noted, workers of N. texanus
have the declivitous face of the propo-
deum slightly concave in profile (more or
less straight in N. nigrescens), and forming
a more distinctive angle with the dorsal
face of the propodeum, but the difference
is a subtle one. Workers of N. texajius also
have the frontal carinae more widely sep-
arated than those of N. nigrescens, and
plots of MFC (the minimum distance be-
tween the frontal carinae) against various
measures of body size (e.g., HW, HL,
MTL) reveal strong and almost diagnostic
differences (Figs. 31-33), when consider-
ing those populations of N. nigrescens in
the Southwest and eastern United States
that are sympatric with N. texanus. Work-
ers of N. texanus also tend to have a short-
er, higher petiole than those of N. nigres-
cens (Fig. 34). Finally, the swelling of the

metapleural gland (bulla) is densely punc-
tulate and more or less opaque over near-
ly all its surface in workers of N. texanus,
while in N. nigrescens it presents a more
shiny appearance. Differences between N.
texanus and N. californicus have been con-
sidered under the latter species.

Material examined.— {CASC, LACM,
MCZC, RAJC, UCDC, USNM)

MEXICO Chihuahua: Mpio. Riva Palacio, Bella-
vista (W. & E. MacKay); Mpio Chihuahua, 45km S
Sueco (Carnada): Durango: 37mi W Durango (W. S.
Creighton); Hidalgo: 5mi S Tizacuya (W. S. Ross);
Guerrero Mill (W. M. Mann); Pachuca (W. M. Mann);
San Miguel (W. M. Mann); Jalisco: Guadalajara
(McClendon; W. M. Mann; Cadwallader); Japopan
(Cadwallader); Nuevo Leon: Vallecillo (W. F. Buren);
Sonora: lOmi S Agua Prieta (V. Roth); 26mi NW Ba-
hia Kino (E. M. Fisher).

UNITED STATES Arizona: Cochise Cc: 5.8mi SE
Sunnyside, 5700 ft. (R. R. Snelling); S.lmi SE Sunny-
side, 5950 ft. (R. R. Snelling); Chiricahua Mts., Rucker
Camp, T19S R29E Sect.22 (W. MacKay); Douglas (W.
W. Jones); Huachuca Mtns., 3mi SW Rte. 92 on Coro-
nado Natl. Mon. Rd., 5100 ft. (S. P. Cover); Huachuca
Mtns., 3nii S jet. Rt. 92 on rd. to Coronado Natl. Mon.,
5100 ft. (S. P. Cover); Huachuca Mtns., Bear Creek,
19km SW Sierra Vista, 1640m (S. G. Brady); Palmerlee,
Huachuca Mtns., 5300 ft. (W. M. Wheeler); San Ber-
nardino Valley, Imi NW jet. Rt. 80 on FSR 74 (Rucker
Canyon Road), 4700 ft. (S. P. Cover); Piiim Co.: 16mi W
Tucson (S. Prchal; G. C. Snelling); Baboquivari Mtns,
Brown Canyon (A. C. Cole); Baboquivari Mtns, For-
estry Cabin, 3500 ft. (W. S. Creighton); Brown Canyon,
Baboquivari Mtns. (Menke & Stange); Sabino Canyon
(R, H. Crandall); Tucson Mtns. (F. R. Gehlbach); Santa
Cruz Co.: Nogales (Buren); Colorado: EI Paso Co.: Col-
orado Springs (J. G. Jack); Colorado Springs and vicin-
ity (W. M. Wheeler); ]efferson Co.: Prospect Park (c.u.);
Florida: Alachua Co.: 29°34.5'N, 82°29'W (R. W. Lund-
gren); Austin Carey Forest, Gainesville (G. B. Fair-
child); Gainesville (T. H. Hubbell; A. Van Pelt); Pierce
Homestead, Gainesville (W. R. Pierce); Escaiiil'ia Co.:
Pensacola (R. M. Lhamon); hiilum River Co.: Sebastian
(Nelson); Leon Co.: Woodville (D. E. Read); Volusia Co.:
Daytona [Beach] (W. F. Buren); New Mexico: Dona
Ana Co.: 45km NE Las Cruces (W. Mackay); EMi/ Co.:
Los Medanos, T22S R31E, Sect.l5 (c.u.); San Miguel Co.:
Las Vegas (W. M. Wheeler); North Carolina: Nczo Han-
over Co.: Wilmington (Vanderford); Yancey Co.{7): Black
Mts. (c.u.); South Carolina: Oconee Co.: Clemson Col-
lege (J. Berly & M. Smith); Texas: Bexar Co.: San An-
tonio (E. S. Ross); Croeketl Co.: Ozona (A. C. Cole); Jeff
Davis Co.: Fort Davis (T. W. Taylor); Kimhie Co.: Junc-
tion 0- F- Watkins); Llano Co.: Llano (A. W. Morrill);
McLennan Co.: Baylor Camp Q. F. Watkins); Waco (J.

90 Journal of Hymenoptera Research

Briga; S. Davis; S. Eldridge; F. R. Gehlbach; O. L. Nich- Chesterfield Co.: Warwick (Bond); Essex Co.: Imi SE

olson; R. W. Plsek; C. A. Rhines; J. F. Watkins); Taylor Durmsville (D. R. Smith); Henry Co.: Chatmoss Plant,

Co.: Abilene rest stop (W. S. Creighton); Travis Co.: Martinsville (S. Schaeffer); Loin'sn Co.: 4mi S Cuckoo (J.

Austin (W. M. Wheeler); Austin, Brackenridge Field Kloke & D. R. Smith); Norfolk city: Norfolk (Vander-

Lab (S. D. Porter); Victoria Co.: Victoria (c.u.).; Virginia: ford).

MODIFICATION OF EXISTING IDENTIFICATION KEYS

Couplets 5 and 9 of Watkins' (1985) worker key to the United States species of hleivamyrmex
need modification to take into account the much greater intraspecific variability in body sculpture
in N. nigresceiis. In the updated portions of the key (below) I have also documented addihonal
features of N. tcxanus and N. opncithornx which will facilitate their identification.

5 In lateral view declivitous face of propodeum slightly concave and forming a somewhat
angular corner with the dorsal (basal) surface; frontal carinae well separated (MFC 0.05-
0.17), as revealed in bivariate plots of MFC on various measures of body size (Figs. 31-
33); petiole relatively short and high (Fig. 34); surface of most of the metapleural gland
bulla densely punctulate and opaque, although sometimes with a very small, thin shiny
strip immediately above the flange of the metapleural gland orifice ... N. texanus Watkins

- Declivitous face of propodeum more or less straight in lateral view and forming a round-
ed angle with the dorsal (basal) surface; frontal carinae usually more closely contiguous
(Figs. 31-33) and petiole height tending to be lower (Fig. 34); sculpture on surface of
metapleural gland bulla becoming obsolete, such that the lower half or more is smooth
(or weakly reticulate) and conspicuously shiny N. nigrescens (Cresson) (part)

9 Inner basal margin of mandible with a straight edge which forms an angular corner or
small tooth at its juncture with the masticatory margin; side of pronotum usually smooth
and shiny, and contrasting with the rugulose-punctulate sculpture that covers at least

part of the mesosoma dorsum N. opacithorax (Emery)

Inner basal margin of mandible with a convex edge which curves into the masticatory
margin without forming an angular corner (Fig. 1); pronotal sculpture variable, but side
of pronotum usually at least weakly reticulate-punctulate rather than exhibiting a smooth,
shiny surface that contrasts with the sculptured mesosoma dorsum 9A

9A Body, scapes and legs shorter, head broader; MTI 1.05-1.14; see also plots of MTL on
HW, SL on HW, WL on HW, CI on MTL and SI on MTL (Figs. 17-19, 21-23, 25, 26);
dorsal surface of propodeum only slightly depressed below the level of the mesonotum
(Fig. 2); dorsum of postpehole and (typically) center of mesonotum smooth and shining,
with scattered piligerous punctures but little or no trace of reticulation or punctulae . . .

N. califomiais (Mayr)

Body, scapes and legs longer; MTI 1.16-1.52; see also Figs. 17-19, 21-23, 25, 26; dorsal
surface of propodeum conspicuously depressed below the level of the mesonotum (Fig.
5); postpetiole and mesonotum varying from opaque to sublucid, with traces of reticu-
lation or punctulae usually evident N. nigrescens (Cresson) (part)

In the key to United States Neivamyrmcx based on queens (Watkins 1972: 350-351) "cnliforniciis"
in couplet 7 should be replaced with "nigrescens (part)". The queen of N. californiciis is not yet
definitively known (see discussion above, under that species).

Couplet 8 of Watkins' (1982) key to Mexican species of Ncivnmyrmux can be modified as follows
to incorporate N. califoniicus and the more recently described N. cluimetcnsis Watkins (1986). The
treatment of N. nigrescens requires no change since the shiny form of N. nigrescens is not known
to occur in Mexico.

8 Head and gaster of smaller workers blackish brown or reddish brown with a black over-
cast, mesosoma reddish brown without a blackish overcast; dorsum of promesonotum

Volume 8, Number 1, 1999

91

slightly convex, and dorsum of propodeum as short or shorter than the node of petiole

in lateral view N. manni (Wheeler)

Head and mesosoma same color (reddish brown), gaster slightly lighter; posterior one-
half of promesonotum flattened; dorsum of propodeum variable in length 8A

8A Dorsal face of propodeum shorter than the declivitous face, and conspicuously depressed
below the level of the promesonotum, at least in larger workers (Watkins 1986, fig. 2);

postpetiole as long as petiole N. chamelensis Watkins

Dorsal face of propodeum as long as, or longer than, the declivitous face, and not con-
spicuously depressed below the level of the promesonotum (Fig. 2); postpetiole shorter
than petiole 8B

8B Inner basal margin of mandible with a straight edge which forms an angular comer or
small tooth at its juncture with the masticatory margin; clypeal apron short, extending
anteriorly by a distance equal to about 0.2-0.4 times the diameter of the torulus; petiole
relatively narrow (Fig. 20); side of pronotum usually smooth and shiny, and contrasting
with the rugulose-punchilate sculpture that covers at least part of the mesosoma dorsum

N. opacithorax (Emery)

Inner basal margin of mandible with a convex edge which curves into the masticatory
margin without forming an angular corner (Fig. 1); clypeal apron longer, extending an-
teriorly by a distance equal to about 0.4-0.5 times the diameter of the torulus; petiole
broader (Fig. 20); pronotal sculpture variable, but side of pronotum usually at least weak-
ly reticulate-punctulate rather than exhibiting a smooth, shiny surface that contrasts with
the sculptured mesosoma dorsum N. califomicus (Mayr)

BIOLOGICAL OBSERVATIONS

Habitat Preferences

Although N. nigrescens and N. califomi-
cus are probably not sister taxa (see below)
they appear to have similar ecological
preferences in California. In southern Cal-
ifornia and northern Baja California,
where morphological differences between
the two species are greatest, their geo-
graphical distributions overlap broadly
(Figs. 35, 36) and they occur together in
such habitats as chaparral, coastal sage
scrub and oak woodland. In northern Cal-
ifornia workers of N. nigrescens are more
difficult to distinguish from those of N.
califomicus, due to convergence in body
sculpture. In this region the two species
are less frequently sympatric. N. califomi-
cus tends to be confined to more mesic,
coastal locations than N. nigrescens, al-
though the habitats occupied are similar:
oak woodland, riparian woodland, chap-
arral and grassland. An exception to this
is the occurrence of N. califomicus at a xe-

ric inland site in the Sierra Nevada foot-
hills (14km NW Shingle Springs, El Do-
rado County) where it co-occurs with N.
nigrescens and N. opacithorax in a distinc-
tive chaparral vegetation on gabbro soil. It
was the sympatric association at this lo-
cality that alerted me to the distinction be-
tween true N. califomicus and the "shiny
morph" of N. nigrescens. The gabbro site
has a number of rare plant species (Hunter
and Horenstein 1992) and N. califomicus
likewise gives the impression of being an
isolated, relic population here (the north-
ern-most point in Fig. 36).

Throughout California both N. califor-
nicus and N. nigrescens are limited to low
elevations, essentially below the conifer-
ous forest zones. Most recorded popula-
tions come from locations below 1500m
(N. califomicus: sea level to 1840m; N. ni-
grescens: sea level to 1460m).

Outside the range of N. califomicus, pop-
ulations of N. nigrescens show a wide lat-
itude of habitat choice, being found in
scrubland, grassland and canyons of the

92

Journal of Hymenoptera Research

35

36

■ nigrescens (s.s.)
^nigrescens (shiny form)

Figs. 35-38. Known distributions in southwestern United States and Mexico of Neivamyrmex nigrescens (25),
N. californicus and N. tcxanus (26), shiny and punctate forms of N. nigrescens (27), and N. opntcitliorax (28). N.
nigrescens, N. opncitlwrnx and N. tcxanus also occur in eastern United States (see Watkins 1985: 499-500).

Sonoran and Chihuahuan deserts, pine-
oak-juniper woodland, prairie grassland,
and eastern deciduous forest (Gregg 1963;
Mirenda et al. 1980; Schneirla 1958; Smith
1942). It is perhaps unsurprising that over
this broad range of ecological conditions
the species displays considerable intraspe-
cific variation in worker morphology.

Associations with Messor atidrei
In California both Neivamyrmex nigres-
cens and N. californicus are often associated
with nests of the common seed-harvesting
ant, Messor andrei (Mayr). Indeed, one of
the most efficient ways to determine the
presence of these species at a locality is to
examine the chaff piles of Messcr tvuirci

Volume 8, Number 1, 1999

93

nests. Such middens frequently contain
the corpses of Neivamyrmex workers.
There are several possible explanations for
this.

First, Messor amirei workers appear to be
efficient scavengers that collect dead and
dying ants (of all kinds) and later discard
the dried corpses in their middens. This is
indicated by the fact that the remains of
other ants, including species of Campono-
tus, Creinntogaster, Don/mynnex, Forelius,
Formica, Leptothorax, Monomorium, Pheidole,
Prenolepis, Solenopsis and Stenamma, are of-
ten encountered in M. audrei middens
(Ward, pers. obs.).

Second, both N. californicus and N. ni-
grescens have been observed attacking
nests of Messor mnirei. In Amador County,
California I observed nocturnal foraging
columns of N. nigrescens workers entering
two adjacent M. ivuirei nests — one contain-
ing a large Messor colony, the other a
small incipient colony. The raid on the
small nest was partially successfully, with
N. nigrescens workers carrying off para-
lyzed Messor worker minims, although
some Messor workers (and a dealate
queen) escaped capture by climbing short
grass stalks. No prey were observed to be
taken from the larger nest, which con-
tained normal-sized Messor workers. Mark
Brown (1999) recorded N. californicus
workers attacking Messor andrei nests at
Jasper Ridge Preserve, near Stanford Uni-
versity. These raids on Messor andrei nests
by N. californicus and N. nigrescens do not
appear to be especially effective, yet in the
observed cases the army ants persisted in
their assaults.

Third, the high frequency of Neivamyr-
mex worker corpses in Messor middens
may reflect a tendency of Neivamyrmex
colonies to temporarily occupy part of the
underground chambers of the Messor
nests, leading to an increased frequency of
interactions between the two species.
Some evidence for this comes from an ob-
servation that I made in early April 1984
near Rutherford, Napa County, northern
California: a colony of N. nigrescens

("shiny form"), containing larvae, was lo-
cated in wet but well-aerated soil imme-
diately adjacent to an active Messor andrei
nest. The N. nigrescens workers were
emerging from the soil and slowly milling
about on the ground surface during day-
light hours, an unusual behavior but one
which occurs in spring (March-May) in
northern California before N. nigrescens
begins its period of summer-active — and
predominantly nocturnal — surface raid-
ing. Thus, it appeared that this Rutherford
colony of N. nigrescens had over-wintered
in the soil in abandoned sections of the
Messor andrei nest. Schneirla (1963) report-
ed the use of pre-empted ant nests (species
not specified) by over-wintering colonies
of N. nigrescens in Arizona.

Interactions with Other California
Ant Species

Other ants besides Messor andrei that are
subject to raids by Neivamyrmex nigrescens
in California include Pheidole californica
Mayr, P. hyatti, Soloiopsis molesta (Say) and
Formica moki Wheeler (Ward, pers. obs.).
Mallis (1938, 1941) reported N. nigrescens
(misidentified as N. californicus) foraging
nocturnally for insects attracted to street
lamps on the Davis campus of the Uni-
versity of California, and attacking nests
of the introduced ant, Tetramorium caespi-
tum (Linnaeus). At the same location Mal-
lis (1938) also recorded an altercation be-
tween Neivamyrmex and Argentine ants,
Linepithema humile (Mayr), in which Line-
pithema emerged the victor. L. humile is
now very abundant on the UC Davis cam-
pus and, during 17 years of observation
here, 1 have seen no evidence that popu-
lations of Neivamyrmex nigrescens survive
on campus, although the species occurs
8km southeast of Davis at a site not yet
overrun by Linepithema humile. Work by
Suarez et al. (1998) in southern California
shows that Neivamyrmex species, includ-
ing N. nigrescens, are among the first ant
species to disappear from patches of rem-
nant coastal sage scrub when Linepithema
humile invades from adjacent urban habitat.

94

Journal of Hymenoptera Research

Table 1. Tests of nest evacuation response in potential prey of Neivnmyrmex. Each test in\'olved placement
of one to several live Neivamyrmex workers at the nest entrance of an active test ant colony. All locations are
in California.

Ncn'ajJii/rTiu'x species

Mass evacuation

Test species

Location

(and source population)

response?

Pheidole desertonim

Pinyon Flats

nigrescens (Pinyon Flats)

Yes

Pheidok hyatti

Carrizo Plain

nigrescens (Carrizo Plain)

Yes

Pheidole hyatti

Del Puerto Canyon

nigrescens (Cold Canyon)

Yes

Pheidole californ ica

Cold Canyon

cnlifornicus (El Dorado Co.)

Yes

Pheidole enlifoni icn

Cold Canyon

nigrescens (Cold Canyon)

Yes

Pheidole califoniicn

Davis

nigrescens (Cold Canyon)

Variable-

Pheidole enlifornicn

Davis

nigrescens (Carrizo Plain)

No

Pheidole enlifornicn

Del Puerto Canyon

nigrescens (Cold Canyon)

No

Pheidole califoniicn

Ventura Co.

nigrescens (Carrizo Plain)

No

Pheidole califoniicn

El Dorado Co.

opacithorax (El Dorado Co.)

No

Messor andrei

Cold Canyon

nigrescens (Cold Canyon)

No

Doryniynnex bicolor

Pinyon Flats

nigrescens (Pinyon Flats)

No

Dorymyrniex insnnus

Davis

nigrescens (Cold Canyon)

No

■' Evacuation response seen in one of two trials.

Elsewhere, across its broad transconti-
nental distribution, N. nigrescens is report-
ed to preferentially raid ant nests, espe-
cially those of Pheidole species (Mirenda et
al. 1980), but also including colonies of
Aphaenogaster, Camponotus, Dorymyrniex,
Formica, Leptothorax and Solenopsis (LaMon
and Topoff 1981; Mirenda et al. 1980;
Schneirla 1958, 1963; Smith, 1927). Both
ant brood, adult workers, and (when
available) sexual alates are taken as prey,
as are occasionally termites and non-social
insects.

Less is known about the prey preferenc-
es of Neivamyrmex cnlifornicus but they ap-
pear to be similar to those of N. nigrescens.
Mark Brown (1999) observed N. califortii-
cus attacking both Messor andrei and Phei-
dole californica colonies at Jasper Ridge.
There is indirect evidence that Solenopsis
molesta and Pheidole hyatti are also preyed
upon. In a collection of dead N. californicus
workers from a Messor andrei chaff pile at
Jasper Ridge (collected by Nate Sanders)
several individuals had dead workers of
Solenopsis molesta attached (with closed
mandibles) to legs and /or antennae. Dead
workers of N. californicus (as well as those
of N. nigrescens) have been found in the
nest middens of Pheidole hyatti (Ward,

pers. obs.). This and certain other Pheidole
species in California show an enemy-spe-
cific nest evacuation response to the pres-
ence of N. californicus and N. nigrescens
workers.

Observations on this nest absconding be-
havior in California Pheidole are summa-
rized in Table 1. A single Neivatnyrmex
worker, held with a pair of forceps at the
nest entrance, can be sufficient to cause
mass evacuation of workers and brood in
Pheidole californica, P. desertorum, and P.
hyatti. In Pheidole californica the response is
not invariably observed, however (Table 1),
and seems to vary with location, ambient
conditions, and possibly as a function of
previous experience (as documented for
another prey species, Aphaenogaster cocker-
elli Andre (McDonald and Topoff 1986)).
Similar nest evacuation behavior has been
studied in Arizona populations of Pheidole
desertorum and P. hyatti attacked by N. ni-
grescens (Droual 1983, 1984).

POPULATION DIFFERENTIATION
AND BIOGEOGRAPHY

The taxonomic confusion surrounding
N. californicus and N. nigrescens can be
traced to undue reliance on superficial fea-
tures of head sculpture, lack of attention

Volume 8, Number 1, 1999

95

to other structural differences, and insuf-
ficient appreciation of the degree of vari-
ability in N. nigrescens. In Neivamyrmex
and in all other genera of New World
army ants (which together comprise the
subfamily Ecitoninae) the queens are en-
tirely wingless and have limited capacities
for dispersal (Gotwald 1995). Because of
the population viscosity associated with
limited female movement — only partly
mitigated by gene flow via dispersing,
winged males — and perhaps also because
of the lower effective population sizes
achieved by these highly predacious (and
hence higher trophic level) organisms, one
expects conspecific allopatric populations
of army ants to develop substantial differ-
ences. The prediction is borne out, at least
among the more widespread species of
ecitonine army ants, in which there exists
a large amount of geographical variation
(Borgmeier 1955, 1956). It seems desirable
for taxonomists to be cautious in establish-
ing new species of army ants especially
when dealing with closely related allopat-
ric populations. In the present paper I
have refrained from giving a new name to
the shiny form of N. nigrescens, because it
is essentially allopatric to the more typical
granulate-punctate N. nigrescens. The two
are not known to both occur sympatricly
and retain their distinctness; rather, in geo-
graphically intermediate localities (i.e., in
the San Gabriel Mountains of southern
California) we find morphologically tran-
sitional populations. A genetic analysis of
the transition zone would be interesting,
especially in view of its relative narrow-
ness.

Thus, the distribution of the two forms
of N. nigrescens (Figure 37) suggests a pre-
vious fragmentation of the range of N. ni-
grescens, and the consequent differentia-
tion of populations but not to a degree
sufficient to cause reproductive isolation.
The Transverse Ranges of California and
the upper Colorado River basin emerge as
the probable sites of geographic barriers.

The divergence between the common

ancestor of N. nigrescens (sensu lato) and
N. cnlifornicus must have occurred consid-
erably earlier. On the basis of structural
similarities (Figs. 17-19) N. californicus ac-
tually appears to be more closely related
to N. opacithorax than to N. nigrescens, with
the latter species being more closely relat-
ed to N. texanus. This hypothesis could be
tested with the study of additional char-
acters and (crucially) the inclusion of ad-
ditional taxa from Mexico, especially N.
chamelensis, N. cornutus, N. manni, and N.
sumichrasti. The possibility that the wide-
spread N. nigrescens is paraphyletic should
also be considered.

A final point of biogeographic interest
concerns the distribution limits of Neiva-
myrmex californicus (Fig. 36) and N. opaci-
thorax (Fig. 38) in northern California.
Both species appear to be restricted to ar-
eas east and south of the Sacramento River
and the San Francisco /San Pablo Bays.
This drainage system can be expected to
be a significant barrier to dispersal in hy-
menopterans such as army ants whose re-
productive females are wingless.

ACKNOWLEDGMENTS

I am grateful to the following persons for access to
material in collections: Wojciech Pulawski (CASC),
John Sorensen (CDAE), Jack Longino OTLC), Roy
Snelling (LACM), Stefan Cover (MCZC), Ivan Lobl
(MHNG), Stefan Schodl (NHMV), and Ted Schultz
(USNM). Additional useful material was received
from Melissa Bennett, Sean Brady, Don Feener, Rob-
ert Johnson, Mike Kaspari, Nate Sanders, Gordon
Snelling, Andy Suarez and Mark Trepanier. I thank
Sean Brady, Brian Fisher, Jack Longino and two
anonymous reviewers for comments on the manu-
script. Research support was received from the Uni-
versity of California and the National Science Foun-
dation.

LITERATURE CITED

Bolton, B. 1995. A nciv general catalogue of the ants of
Ike zocrhi. Cambridge, Massachusetts: Harvard
University Press, 504 pp.

Borgmeier, T. 1953. Vorarbeiten zu einer Revision der
neotropischen Wanderameisen. Slmlia Entomolo-
gica 2: 1-51.

Borgmeier, T. 1955. Die Wanderameisen der neotro-
pischen Region. Studia Eiitoinologica 3: 1-720.

96

Journal of Hymenoitera Research

Borgmeier, T. 1956. Ueher Rassen bei Eciton (Hym.
Formicidae). Revista Brasileira tie Entomologia 4;
209-212.

Borgmeier, T. 1958. Nachtraege zu meiner Monogra-
phie der neotropischen Wanderameisen (Hym.
Formicidae). Stiidia Entoinolcgica (n.s.)l: 197-208.

Brown, M. J. F. 1999. Nest relocation and encounters
between colonies of the seed-harvesting ant Mt'S-
sor andrei. Insectes Sociaux, in press.

Creighton, W. S. 1950. The ants of North America.
Bulletin of the Miiicum of Comparative Zoology 104:
1-585.

Cresson, E. T. 1872. Hymenoptera Texana. Transae-
tions of the Ainerieau Eiitontological Soeiety 4: 153-
292.

Dalla Torre, K. W. v. 1893. Catalogus Hymcnopterorum
hucusque descriptorum systemaiicus et synonymicus.
Vol. 7. Formicidae Heterogyna. Leipzig: W. En-
gelmann, 289 pp.

Droual, R. 1983. The organization of nest evacuation
in PheidoU- de^ertorKin Wheeler and P. hyatti Em-
ery (Hymenoptera: Formicidae). Beluwioral Ecol-
ogy and Sociobiology 12: 203-208.

Droual, R. 1984. Anti-predator behaviour in the ant
Pheidole desertonim: the importance of multiple
nests. Animal Behaviour 32: 1054-1058.

Emery, C. 1894. Studi sulle formiche delta fauna neo-
tropica. Vl-XVI. Bullettino della Societii Entomolo-
gica Italiana 26: 137-241.

Emery, C. 1895. Beitrage zur Kenntniss der nord-
amerikanischen Ameisenfauna. Schluss. Zoolo-
gisches Jahrhiicher. Abteilung fiir Systematik, Geo-
grapbie und Bioh^gie der Ticre 8: 257-360.

Emery, C. 1900. Nuovi studi sul genere Eciton. ^4e-
morie della Reah' Accadcn na delle Scicnzc dell'Istituto
di Bologna 58: 173-188 [pagination of separate:
511-526].

Forel, A. 1914. Formicides d'Afrique et d'Amerique
nouveaux ou peu connus. Bulletin de la Societe
Vnudoife des Sciences Naturelles 50: 211-288.

Gotwald, W. H., Jr. 1995. Army ants: the biology of so-
cial predation, Ithaca, New York: Cornell Univer-
sity Press, xviii + 302 pp.

Gregg, R. E. 1963. The ants of Colorado, with reference
to their ecology, taxonomy, and geographic distribu-
tion. Boulder: University of Colorado Press, xvi
+ 792 pp.

Hunter, J. C, Horenstein, J. E. 1992. The vegetation
of the Pine Hill area (California) and its relation
to substratum. Pp. 197-206 in: Baker, A. J. M.,
Proctor, J., Reeves, R. D. (eds.) The vegetation of
ultramafic (serpentine) soils. Andover, U. K.: In-
tercept Ltd., XX + 509 pp.

LaMon, B., Topoff, H. 1981. Avoiding predation by
army ants: defensive behaviours of three ant spe-
cies of the genus Campninotus. Aniniai Behaviour
29: 1070-1081.

Mallis, A. 1938. Army ants in California. Scientific
Monthly 47: 220-226.

Mallis, A. 1941. A list of the ants of California with
notes on their habits and distribution. Bulletin of
the Southern California Academy of Sciences 40: 61-
100.

Mayr, G. 1870. Formicidae novogranadenses. Sit-
zungsberichte der Kaiserlichen Akademie der Wissen-
schaften in Wien. Mathematisch-Naturwissenschaft-
liche Classe. Abteilung I 61: 370-^17.

Mcdonald, P., Topoff, H. 1986. The development of
defensive behavior against predation by army
ants. Developmental Psychobiology 19: 351-368.

Mirenda, J. T., Eakins, D. G., Gravelle, K., Topoff, H.
1980. Predatory behavior and prey selection by
army ants in a desert-grassland habitat. Behavior-
al Ecology and Sociobiology 7: 119-127.

Sclineirla, T. C. 1958. The behavior and biology of
certain Nearctic army ants. Last part of the func-
tional season, southeastern Arizona. Insectes So-
ciaux 5: 215-255.

Schneirla, T. C. 1963. The behaviour and biology of
certain Nearctic army ants; springtime resur-
gence of cyclic function — southeastern Arizona.
Animal Behavwur 11: 583-595.

Smith, M. R. 1927. A contribution to the biology and
distribution of one of the legionarv ants, Eciton
schmitti Emery. Annals of the Entomological Society
of America 20: 401-404.

Smith, M. R. 1938. Notes on the legionary ants Eciton,
subgenus Acamatus with a record of new specific
synonymy. Proceedings of the Entomological Society
of Washington 40: 157-160.

Smith, M. R. 1942. The legionary ants of the United
States belonging to Eciton subgenus Neivamyrmex
Borgmeier. American Midland Naturalist 27: 537-
590.

Snelling, R. R., George, C. D. 1979. The taxonomy, dis-
tribution and ecology of California desert ants. Re-
port to California Desert Plan Program, Bureau
of Land Management, U.S. Dept. Interior. 335 +
89 pp.

Suarez, A. V., Bolger, D. T., Case, T. J. 1998. Effects
of fragmentation and invasion on native ant
communities in coastal southern California. Ecol-
ogy 79: 2041-2056.

Watkins, J. F., II. 1972. The taxonomv of Neiiwnyrmex
texanus, n. sp., N. nigrescens and N. californicus
(Formicidae: Dorylinae), with distribution map
and keys to the species of Neivamyrmex of the
United States. Journal of the Kansas Entomological
Society 45: 347-372.

Watkins, j. F., II. 1976. The identification and distribu-
tion of Nejv World army ants (Dorylinae: Tormici-
dae). Waco, Texas: Bavlor Universitv Press, 102

PP-
Watkins, J. F., II. 1982. The armv anls of Mexico (Hv-

Volume 8, Number 1, 1999

97

menoptera: Formicidae: Ecitoninae). loiirnnl oftlic
Kansas Eiitoiiwlcgical Society 55: 197-247.

Watkins, J. F., II. 1985. The identification and distribu-
tion of the army ants of the United States of Amer-
ica (Hymenoptera, Formicidae, Ecitoninae). journal
of the Kansas Entonwlo\;ical Soach/ 58: 479-502.

Watkins, J. F., II. 1986. Nnvann/rmcx chamelcnsis. n. sp.
(Hymenoptera: Formicidae: Ecitoninae) from Jal-

isco, Mexico. Journal of the Kamas Entomological
Society 59: 361-366.

Wheeler, W. M. 1900. The female of Eciton sumichrasti
Norton, with some notes on the habits of Texan
Ecitons. American Naturalist 34: 563-574.

Wheeler, W. M. 1908. The ants of Texas, New Mexico
and Arizona. Part I. Bulletin of the American Mu-
seum of Natural Histon/ 24: 399^85.

J. HYM. RES.
Vol. 8(1), 1999, pp. 98-108

Revision of North American Aleiodes Wesmael (Part 5):

The melanopterus (Erichson) Species-Group

(Hymenoptera: Braconidae, Rogadinae)

Paul M. Marsh and Scott R. Shaw

(PMM) Cooperating Scientist, USDA Systematic Entomology Laboratory, c/o National Museum

of Natural History, Washington, DC, USA (correspondence address: P. O. Box 384, North

Newton, Kansas 67117 USA); (SRS) Entomology Section, Department of Renewable Resources,

P. O. Box 3354, University of Wyoming, Laramie, Wyoming 82071 USA

Abstract. — The Aleiodes melanopterus (Erichson) species-group is regarded as monophyletic based
on the large oral opening and narrow clypeus. It is defined to include 13 Palaearctic and Neo-
tropical species plus the following North American species: inandibtilaris (Cresson) new com-
bination, megastomus new species, melanopodus new species, mexkanus Cresson, miani new
species, and politiceps (Gahan) new combination. A key to the North American species of the
melanopterus species-group is presented.

The rogadine braconid genus Aleiodes
Wesmael is worldwide in distribution, but
is particularly species-rich in the Holarctic
Region. Aleiodes is well diversified in
North America, with at least 90 species in
the United States and Canada (S. Shaw et
al., 1997). This study is the fifth in a series
of planned papers on Aleiodes species-
groups, intended to provide a complete
revision of the genus for North America
(see S. Shaw et al, 1997, 1998a, 1998b;
Marsh and Shaw, 1998). The melanopterus
(Erichson) group is moderate sized with
species occurring in the Palaearctic, Ne-
arctic and Neotropical Regions. This is a
distinctive monophyletic group with all
species having a large oval oral opening
and narrow clypeus. Our definition of this
species-group includes all species known
to us worldwide. However, because our
main intent is to provide a revision of the
North American species, species treat-
ments are limited to the Nearctic fauna.

Aleiodes species are koinobiont endopar-
asitoids of lepidopterous larvae, especially
macrolepidoptera of the superfamilies
Noctuoidea and Geometroidea, and to a

lesser extent, Sphingoidea and Papiliono-
idea (S. Shaw et al, 1997). Very little is
known about the biology of the melanop-
terus species-group but the few records in-
dicate parasitism of noctuid larvae. The
method of parasitism, unique to the tribe
Rogadini, is noteworthy: the Aleiodes larva
completes its feeding and pupates within
the shrunken and mummified remains of
the host caterpillar. In all known cases, the
form of the mummy caused by a particu-
lar Aleiodes species is characteristic for that
host and parasitoid, so mummified re-
mains are of considerable diagnostic value
and should be retained with the parasitoid
when reared. For a more complete discus-
sion of Aleiodes biology, readers may refer
to M. Shaw (1983, 1994), M. Shaw and
Huddleston (1991), S. Shaw (1995) and S.
Shaw et al. (1997). We have host informa-
tion for only one North American species,
politiceps (Gahan), which has been reared
from at least two species of noctuid larvae.

METHODS

Species covered in this paper can be
identified as members of the subfamily

Volume 8, Number 1, 1999

99

Rogadinae using the keys of S. Shaw
(1995), M. Shaw and Huddleston (1991) or
Wharton et al. (1997). Our definition of Al-
eiodes follows that of S. Shaw (1993), S.
Shaw et al. (1997) and van Achterberg
(1991). Specimens can be determined as
Aleiodes using the keys of Wharton et al.
(1997). The species-groups of North Amer-
ican Aleiodes can be identified using the
key provided in S. Shaw et al. (1997).

Terminology follows that used for Aleio-
des by S. Shaw et al. (1997), S. Shaw (1993)
and Marsh (1989). Microsculpture termi-
nology follows that of Harris (1979). Wing
vein terminology agrees with the system
adopted by Wharton et al. (1997) and
agrees closely with that of Goulet and
Huber (1993). A labeled diagram of wing
veins was provided by S. Shaw et al.
(1997).

Acronyms for collections where type
material is deposited are as follows: AEI
(American Entomological Institute,
Gainesville, FL), AMNH (American Mu-
seum of Natural History, New York, NY),
ANSP (Academy of Natural Sciences,
Philadelphia, PA), CNC (Canadian Na-
tional Collection, Ottawa, Canada), CAS
(California Academy of Sciences, San
Francisco, CA), FSCA (Florida State Col-
lection of Arthropods, Gainesville, FL),
MCZ (Museum of Comparative Zoology,
Harvard University, Cambridge, MA),
MSU (Montana State University, Boze-
man, MT), NCSU (North Carolina State
University, Raleigh, NC), NNML (Natio-
naal Natuurhistorisch Museum, Leiden,
The Netherlands), RMSEL (Rocky Moun-
tain Systematic Entomology Laboratory,
University of Wyoming, Laramie, WY),
TAMU (Texas A&M University, College
Station, TX), UCD (University of Califor-
nia, Davis, CA), USNM (National Muse-
um of Natural History, Smithsonian Insti-
tution, Washington, DC).

ALEIODES MELANOPTERUS
SPECIES-GROUP

Included species. — miiuatiis (Herrich-
Schaeffer) 1838, new combination, Europe,

North Africa; melanopterus (Erichson) 1848,
new combination, South America; aestiio-
siis (Reinhard) 1863, new combination,
eastern Europe, Middle East; mexicanus
Cresson 1869, Mexico, southern United
States; maiidibularis (Cresson) 1872, new
combination, central United States; kruli-
koivskii (Kokoujev) 1898, new combination,
eastern Europe, Mongolia; veimstulus (Ko-
koujev) 1905, new combination, eastern
Europe; lucidus (Szepligeti) 1906, new
combination, Bolivia; politiceps (Gahan)
1917, new combination, eastern United
States, Central America; wadai (Watanabe)
1937, new combination, Japan; agilis (Te-
lenga) 1941, new combination, eastern Eu-
rope; desertus (Telenga) 1941, new combi-
nation, eastern Europe; /(7/jri»^^fn (Telenga)
1941, new combination, Mongolia; glaber
(Telenga) 1941, new combination, eastern
Europe, Mongolia; riificeps (Telenga) 1941,
new combination, eastern Europe; flavis-
tignia Shaw 1993, Brazil; megastomus, new
species; melaiwpodus, new species; miani,
new species.

Diagnostic characters. — Oral opening
(Figs. 5-9) large and oval, width equal to
or greater than height of face, clypeus very
narrow; eyes and ocelli large, the ocellar
diameter equal to or slighter greater than
diameter of lateral ocellus; mesonotum
and mesopleuron usually smooth and pol-
ished; hind wing vein RS straight, margin-
al cell gradually widening to wing apex
(Figs. 1^). A discussion of the phyloge-
netic relations of the species-groups of Al-
eiodes can be found in Fortier (1997).

Comments. — This is a moderately sized
species-group associated with noctuids
and distinguished by the large oval oral
opening. The Neotropical species have
been reviewed by Shaw (1993) where he
placed them in the subgenus Eucystomas-
tax. Although the name melanopterus used
for this species-group is not the oldest
name, it has been used previously by
Shaw (1993) in his study of the Neotropi-
cal species and Fortier (1997) in his study
of Aleiodes phylogeny. For this reason and

100

Journal of Hymenoptera Research

because the ICZN does not provide for the
naming of species groups, we have decid-

ed to continue the usage of melanopterus in
this paper.

KEY TO THE NORTH AMERICAN SPECIES OF THE MELANOPTERUS SPECIES-GROUP

1. Width of oral opening about equal to height of face (from clypeus to antennal sockets);

malar space about equal to basal width of mandible (Figs. 8, 9) 2

Width of oral opening greater than height of face; malar space much less than basal

width of mandible (Figs. 5-7) 3

2(1). First and second metasomal terga strongly porcate (Fig. 11); fore wing vein ICUa longer

than Icu-a, wings strongly infumated (Fig. 3) politiceps (Gahan)

First and second metasomal terga weakly costate rugulose (Fig. 10); fore wing vein ICUa

equal in length to Icu-a, wings hyaline or weakly infumated

miani Marsh and Shaw, new species

3(1). Body bicolored, at least legs black, differently colored than body 4

- Body unicolored, legs concolorus with rest of body 5

4(3). Head and legs entirely black mexicanus Cresson

Head orange, femora, tibiae and tarsi only black

melanopodus Marsh and Shaw, new species

5(3). Fore wing vein ICUa as long as or shorter than vein Icu-a (Fig. 4); clypeus flat, narrow,

without distinct apical border megastomus Marsh and Shaw, new species

Fore wing vein ICUa longer than Icu-a (Fig. 1); clypeus protruding, with distinct apical
carinate border mandibttlaris (Cresson)

Aleiodes mandibularis (Cresson),
new combination

(Figs. 1, 6)

Rogas mandibularis Cresson 1872:188.

Diagnosis. — Body unicolored orange, an-
terma brown, wings hyaline or slightly
dusky, veins brown; body length, 8.5-10.0
mm; 55-65 antennomeres; oral opening
width greater than height of face (Fig. 6);
malar space short, equal to or less than
basal width of mandible; face costate,
frons, vertex and temple smooth; ocelli
small, diameter of lateral ocellus less than
ocell-ocular distance; pronotum rugose
medially; mesonotum, scutellum and me-
sopleuron punctate and shining, subalar
sulcus and sternaulus weakly rugose; pro-
podeum rugose, median carina complete;
first, second and basal Vi of third metaso-
mal terga costate-rugose, median carina
complete to middle of third terga; fore
wing vein Icu-a beyond IM by distance
greater than length of Icu-a, vein ICUa

longer than Icu-a (Fig. 1); hind wing mar-
ginal cell narrowest basally, gradually
widening apically, vein RS straight, vein
m-cu present (Fig. 1); tarsal claws strongly
pectinate.

Type material examined. — Holotype male
of Rogas mandibularis Cresson is not in the
ANSP and apparently lost. A neotype has
been selected as follows: male, TEXAS,
Victoria, September 18, 1904. Deposited in
USNM.

Distribution. — Scattered collections
throughout central United States from
Ohio south to Georgia, west to Nebraska
and Texas.

Biology. — Host unknown.

Comments. — Although mandibularis is a
large and distinctive species, it is not com-
monly collected. Superficially it resembles
politiceps because of its large orange body,
but the more greatly enlarged oral open-
ing, robust mandibles, and hyaline wings
will easily distinguish mandibularis from
that species.

Volume 8, Number 1, 1999

101

Figs. 1^. Wings of Aleiodes species: 1, iiiaiidibulans (Cresson); 2, nichuwpwdiif n. sp.; 3, politicei's (Gahan); 4,
iiifgiKtonutf n. sp.

Aleiodes megastomus Marsh and Shaw,
new species

(Figs. 4, 5)

Female. — Body color: varying from dark
honey yellow to dark brown or black, legs
and mandibles always brown; wings hy-
aline, veins including stigma dark brown,
stigma occasionally lighter brown or yel-
low. Body length: 7-9 mm; fore wing
length, 7-9 mm. Head (Fig. 5): 57-61 an-
tennomeres, flagellomeres beyond first
slightly longer than wide, first flagello-
mere nearly twice as long as second; malar
space short, !/, eye height and % basal
width of mandible; temple % eye width;
occipital carina not meeting hypostomal
carina; oral opening very wide and oval,
width three times malar space and slightly
greater than height of face; clypeus flat.

without distinct apical carinate border;
ocelli large, greatest diameter of lateral
ocellus slightly greater ('4) than ocell-ocu-
lar distance; face rugulose, frequently with
median longitudinal ridge below anten-
nae; frons smooth, occasionally rugulose
behind antennae; vertex punctate; temples
punctate, rugulose near base of mandi-
bles; maxillary palpus not swollen; man-
dibles large, when closed Hps going well
beyond middle of oral opening. Mesoso-
ma: pronotum rugulose; mesonotum and
scutellum smooth but often with conspic-
uous hair pits making it appear punctate;
notauli weakly scrobiculate, meeting pos-
teriorly in triangular rugose area; scutellar
furrow wide, scrobiculate posteriorly,
separated from mesonotum by transcutal
ridge; mesopleuron smooth, subalar groove

102

Journal of Hymenoptera Research

scrobiculate; stemaulus absent; propodeum
weakly rugulose dorsally, smooth laterally,
propodeal carina present but often weak.
Legs: tarsal daw weakly pectinate, with 3-
4 short teeth near base; inner spur of hind
tibia slightly less than half length of hind
basitarsus; hind coxa rugulose dorsally.
Wings (Fig. 4): hyaline; fore wing vein r
Vi length of 3RSa and of m-cu, vein Icu-a
beyond IM by distance less than length of
Icu-a, vein ICUa about '4 length of ICUb,
vein ICUa about equal to Icu-a; hind
wing marginal cell gradually widening
from about middle to wing margin, vein
r-m equal to or slightly shorter than IM,
vein M+CU only slightly longer than IM,
vein m-cu present. Metasoma: first ter-
gum weakly strigate with raised smooth
triangular area at base, as long as apical
width, basal width Vi apical width, medi-
an carina usually present, often weak and
occasionally absent; second tergum weak-
ly strigate, often smooth at apex, median
carina weak and often absent; third and
following terga smooth, third occasionally
punctate at base, median carina absent on
third tergum; ovipositor sheaths % length
of hind basitarsus.

Male. — Essentially as in female.

Ho/ofi/;'f.— Female: CALIFORNIA, Cor-
coran, Kings County, March 10, 1965, F. H.
Surber, light trap. Deposited in USNM.

Paratypes.— ARIZONA: 2 females. No-
gales, January 30, 1954, March 9, 1961, A.
C. Valcarce; 1 female, Prescott, April 22,
1936, Cby. and Bish.; 3 females, Tempe,
February 5, 1923, February 8, 1926, E. V.
Walter; 10 females, 5, males, Tuscon, Jan-
uary 27, 1935, February 4, 1935, December
10-20, 1939, January 8, 1940, January 26,
1953, R. H. Crandall, Hubbard, Samual
Green, E. C. Narschall, Donald Foote, G.
C. Butler, Bryant collectors, some at light;
1 female, Cameron, Coconimo Co., Feb-
ruary 28, 1978, R. C. Miller; 1 female, Bea-
ver Dam, Mojave Co., April 2, 1969, P. M.
Marsh; 1 fenale, 1 male, Baboquivari
Mtns., August 1, 1924, O. C. Poling. CAL-
IFORNIA: 1 female, Manzanita Lake, Las-

sen Nat. Park, May 23, 1941, P. D. Hurd;

1 female, 1 male. May Lake, Yosemite Pk.
July 26, 1948, H. K. Townes; 6 females, 5
males, 3 mi. SW Somerset, El Dorado Co.,
dates ranging from April 20, 1974 to May
20, 1978, R. Wharton; 2 females, 2 males.
Thousand Palms, February 14-22, 1955,
W.R.M. Mason, J.E.H. Martin; 1 female.
Desert Hot Springs, February 14, 1965,
J.E.H. Martin; 1 female, Wrightwood, San
Bernardino Co., June 16, 1964, J. S. Buck-
ett; 2 females, Quincy, Plumas Co., June 5,

1963, G. Jeskey; 1 female, McClure Vly.,
Kings Co., March 6, 1953, J. C. Hall; 1 fe-
male, Huntington Lake, June 26, 1961, A.
T. McClay; 5 females, 1 male, Calaveras
Co., 4.8 km S West Point, May 1-9, 1981,
Stanley C. Williams; 1 female, Napa Co.,
N. side Howell Mtn., 2 mi NNE Angwin,
1300 ft., April 11, 1978, H. B. Leech; 1 fe-
male, Napa Co., 1 mi N. Angwin, May 9,

1964, H. B. Leech; 1 female, Nevada City,
May 16, 1930, E. P. Van Duzee; 1 female,
Fresno Co., Coalinga, Los Gatos Cn.,
March 17, 1931, E. P. Van Duzee; 1 female,
Mt.. Diablo, April 23, 1939. IDAHO: 1
male, Murtaugh, May 27, 1931, D. E. Fox;

2 females, Lewiston, April 27, 1936, alt.
550 ft., R. E. Miller. KANSAS: 1, female,
Manhattan, April 24, 1926, R. T. Cotton, at
light. MONTANA: 1 female, Gallatin Co.,
May 8, 1932. NEVADA: 1 female, Reno,
May 14, 1915; 1 female, Kyle Cyn., Chlstn,
Mt., May 25, 1940, G. E. Bohart. UTAH: 1
female. Blue Spruce Camp, 18 mi. N Es-
calante, Garfield Co., 8000 ft., July 30,

1965, F., P. & M. Rindge; 6 females, Rich-
field, May 22, 1929, May 28, 1930, light
trap; 2 females, 1 male, Wellsville, May 17,
1961, G. E. Bohart; 1 female. Baker,
S14,T13N,R69E, May 4, 1939, T. O. Thatch-
er. WASHINGTON: 3 females, 1 male,
Yakima, June-September, March 30, 1932,
at light, A. R. Rolfs; 1 male, Orville WY-
OMING: 2 females, Stratton Expt. Water-
shed, nr. Saratoga, May 16-17, 1972, June
4-8, 1972, J. M. Schmid; 26 females, 2
males, Albany Co., T15N R73W, 7500',
May 15, 1978, May 20, 1980, C. D. Ferris;

Volume 8, Number 1, 1999

103

1 female, Carbon Co., 3 mi. N of WY
130W, between mi. 54 and 55, June 25-
July 4, 1991, Malaise trap, Mian; 1 female,
Fremont Co., Sinks Cyn., 12.1 mi. SW
Lander, June 23, 1980, LT, Mike Pogue,
Robert Lavigne; 2 females, 1 male, Albany
Co., 1 mi. E Laramie, May 10-18, 1994,
June 8, 1996, BLT 2217, Skyview Lane,
7465 ft., J. S. Nordin. SASKATCHEWAN:

1 female, Saskatoon, May 12, 1933, A. R.
Brooks. Deposited in USNM, RMSEL,
AMNH, TAMU, MCZ, MSU, CNC, CAS,
UCD, NNML.

Distribution. — This species is common in
western North America from Saskatche-
wan to New Mexico and west. The single
specimen from Kansas is this species and
with more collecting it probably will occur
throughout the western half of North
America.

Biology. — No host records are available.
Many of the type series were collected at
light or in light traps. The general habitus
and color of this species is similar to spe-
cies of Homolobus which frequent light
traps.

Comments. — This species is similar to
mandibularis because of the wide oral
space. It differs in several characters: the
ocellus of megastomus is larger than in
mandibularis, the clypeus of megastomus is
flat rather than protruding as in mandibu-
laris, and vein Icu-a in the fore wing of
megastomus is closer to IM than in mandi-
bularis.

Eti/mologi/. — The specific name is from
the Greek megas meaning large or great
and stoma meaning mouth referring to the
large wide oral space.

Aleiodes tnelanopodus Marsh and Shaw,
new species

(Figs. 2, 7)

Female. — Body color: head, mesosoma,
metasoma, coxae and trochanters orange;
antenna, palpi, ovipositor sheaths, and
legs beyond trochanters black, trochanter

2 sometimes dark; wings dusky. Body
length: 8-9 mm, fore wing length, 7-8

mm. Head (Fig. 7): 53-58 antermomeres,
first flagellomere twice as long as wide,
flagellomeres 10-30 as wide as long; malar
space short, Va eye height and % basal
width of mandible; temple bulging, about
as wide as eye; occipital carina nearly
meeting hypostomal carina; oral opening
wide and oval, width 3.5 times malar
space and about twice height of face; clyp-
eus protruding; ocelli small, greatest di-
ameter of lateral ocellus about % ocello-
cular distance; face costate, with median
longitudinal ridge below antennae; frons
smooth, occasionally weakly rugulose be-
hind antennae; vertex and temple smooth;
maxillary palpus not swollen; mandibles
large, when closed tips going beyond mid-
dle of oral space. Mesosoma: pronotum
rugulose laterally; mesonotum and scutel-
lum smooth and shining; notauli weakly
scrobiculate, meeting posteriorly in trian-
gular rugulose area with central longitu-
dinal carina; mesopleuron smooth and
shining, subalar sulcus rugulose, sternau-
lus short and wide, weakly scrobiculate;
propodeum strongly rugose dorsally,
smooth laterally with rugosities near hind
coxa, median carina complete. Legs: tarsal
claws strongly pectinate, with 6-8 teeth on
entire inner surface of claw; hind coxa
smooth dorsally. Wings (Fig. 2): dusky;
fore wing vein r K length of 3RSa and %
length of m-cu, vein Icu-a beyond IM by
distance about twice length of Icu-a, vein
ICUa about X length of ICUb, vein ICUa
longer than Icu-a; hind wing marginal cell
gradually widening from about middle to
wing margin, vein r-m about % length of
IM, vein M + CU about 1.4 times longer
than IM, vein m-cu present. Metasoma:
first tergum costate-rugose, length slightly
less than apical width, median carina com-
plete; second tergum costate-rugose, me-
dian carina strong on basal half; third ter-
gum costate on basal %, median carina ab-
sent; fourth and following terga smooth,
fourth weakly coriaceous on basal half;
ovipositor sheaths about 1.5 times longer
than hind basitarsus.

104

Journal of Hymenoitera Research

Figs. 5-8. Faces of Aleiodes species: 5, mcgnstomiis n. sp.; 6, numdibulnnf (Cresson); 7, nninuopiMiuf n. sp.; 8,
iniaiii n. sp.

Male. — Essentially as in female.

Holoh/pe.— Female: MARYLAND, Cabin
John, (label actually states Washington,
DC which is in error), September 24, 1922,
H. S. Barber collector. Deposited in
USNM.

Parntypes.— KENTUCKY: 1 female, 1
male. Golden Pond, October 1-8, 1964,
September-October, 1965, S. G. Breeland.
MARYLAND: 1 male. Cabin John, Octo-
ber 1, 1916, R. M. Fouts collector; 1 male.

Bethesda, September 23, 1914, R. C. Shan-
non collector; 1 female. Prince George's
Co., Beltsville, September 19, 1964, Paul H.
Arnaud, Jr.; 1 female, Riverdale, Prince
George's Co., September 26, 1979, T. Wa-
ters; 1 female, Takoma Park, September
29, 1945, H. and M. Townes. LOUISIANA:
1 male. Sunshine, October 27, 1972, V. A.
Brou. NORTH CAROLINA: 1 male. Wake
Co., October 16, 1959; 1 female. Wake Co.,
October 3, 1959, O. F. Vargas; 1 male.

Volume 8, Number 1, 1999

105

Clayton, Cent. Crops. Res. Sta., 1959, A.
Saldarriaga; 1 male, Raleigh, October 3,
1959, L. Self; 1 female, Pender Co., October
9, 1954, D. A. Becker; 1 male. Wake Co.,
Raleigh, Centienial Campus, October 13,
1992, E. D. Karoly; 1 male. Wake Co., Ra-
leigh, November 5, 1988, D. L. Moncol.
VIRGINIA: 1 female, Richmond, collection
Ashmead; 1 female. Falls Church, Septem-
ber 26 (no year). Banks collector. Depos-
ited in USNM, RMSEL, MCZ, CAS,
NCSU, FSCA, AEI.

Distribution. — Known from Maryland,
Virginia, North Carolina, Kentucky and
Louisiana.

Biology. — Unknown.

Comments. — This species is very similar
to mandibiilaris by the shape of the mouth
and the body color. It differs in having
black legs, smaller ocelli, and longer ovi-
positor.

Etymology. — The specific name is from
the Greek melanos meaning dark or black
and the Greek podos meaning foot refer-
ring to the distinctive black legs.

Aleiodes mexicanus Cresson

Aleiodes mexicamis Cresson 1869:378.

Diagnosis. — Body bicolored, head, an-
tenna, pronotum, propleuron and legs
black, mesosoma and metasoma orange,
wings dusky, veins brown, tegula orange;
body length, 9 mm; 64 antennomeres; ma-
lar space very short, about V2 basal width
of mandible; oral opening large, diameter
5 times malar space length and about
equal to face height; ocelli large, ocello-
cular distance less than diameter of lateral
ocellus; frons, vertex and temple smooth,
face rugulose; mesonotum and mesopleu-
ron smooth; propodeum with dorsal api-
cal corners sharp, median carina complete;
first, second and basal % of third metaso-
mal terga strigate, median carina complete
of first and second terga which also have
lateral carinate edges; fore wing vein Icu-
a beyond vein IM by distance slightly
greater than length of Icu-a, vein ICUa

slightly longer than Icu-a; hind wing mar-
ginal cell narrowest at base, gradually
widening to apex, vein RS straight; tarsal
claws pectinate, with 4-6 slender spines.

Type matericd examined. — Aleiodes mexi-
canus Cresson, holotype female (not male
as in original description), Mexico
[ANSP].

Distribution. — In addition to Mexico, we
have seen one specimen from Mississippi.

Biology. — Unknown.

Comments. — This species is distinctive
by its color, short malar space and wide
oral opening. The one specimen from Mis-
sissippi may be an accidental introduction
and the establishment of this species in the
U.S. needs to be confirmed by further col-
lecting in the southern United States.
Shaw (1993) provides a description and
key to this species and other related Neo-
tropical species.

Aleiodes miani Marsh and Shaw,
new species

(Figs. 8, 10)

Female. — Body color: honey yellow ex-
cept antenna, ocellar triangle, palpi, pro-
pleuron, mesosternum, metanotum, pro-
podeum, basal half of first metasomal ter-
gum, apex of all femora, fore and middle
tibiae, apical half of hind tibia and all tarsi
black, wings hyaline, veins dark brown,
tegula yellow. Body length: 7.0-9.0 mm;
fore wing length, 5.0-7.0 mm. Head (Fig.
8): eyes and ocelli not unusually large; 59-
60 antennomeres, all flagellomeres beyond
the first about as wide as long; malar
space shorter than basal width of mandi-
ble and about Va eye height; temple broad,
slightly less than eye width; occipital ca-
rina complete on vertex, not reaching hy-
postomal carina; oral opening large, cir-
cular, width greater than basal width of
mandible and face height; clypeus narrow,
not swollen; ocelli small, diameter of lat-
eral ocellus slightly less than ocellocular
distance; face, frons and malar space ru-
gose, temple and vertex smooth except for
hair pits; maxillary palpus not swollen;

106

Journal of Hymenoptera Research

Figs. 9-11. Morphological features of Alciccics species: 4, face ol poltluc^'^ (Cjahaii); 10, metasomal terga of
miani n. sp.; 11, metasomal terga oi poUticqps (Gahan).

Volume 8, Number 1, 1999

107

mandibles large, tips crossing when
closed. Mesosoma: pronotum costate ru-
gose laterally, costate ventrally; mesono-
tum and scutellum smooth except for hair
pits, notauli weakly scrobiculate, meeting
in shallow rugose area before scutellum;
mesopleuron smooth, subalar sulcus ru-
gose, sternaulus carinate; propodeum ru-
gose dorsally, smooth laterally, median
carina weak but complete. Legs: tarsal
claws weakly pectinate; hind coxa punc-
tate dorsally. Wings: fore wing vein r
about Vs length of 3RSa and m-cu, vein
Icu-a beyond IM by distance equal to
length of Icu-a, vein ICUa about Va length
of ICUb; hind wing vein RS straight, mar-
ginal cell gradually broadening to apex,
vein r-m shorter than IM, vein M + CU
slightly longer than IM, vein m-cu weak
and leaving IM before junction of r-m.
Metasoma (Fig. 10): first tergum costate
rugose, apical width about equal to
length, median carina complete; second
tergum costate rugose, median carina not
quite complete; third tergum costate ru-
gose on basal half, coriaceous on apical
half, median carina absent; remainder of
terga coriaceous; ovipositor less than Vi
length of hind basitarsus.

Male. — Essentially as in female except
mesopleuron and pronotum nearly and
first metasomal tergum entirely black.

Ho/ofi/p.— Female: WYOMING, Carbon
Co., 3/4 mi. N of WY 130 W between mi. 56
& 55, mixed forest near water. Malaise,
Mian, July 14-27, 1991. Deposited in
RMSEL.

ParaU/pes.— WYOMING, 1 female, 4
males, same data as holotype with dates
of June 25 through August 6, 1991. De-
posited in USNM, RMSEL.

Distribution. — Known only from Wyo-
ming.

Biologi/. — Unknown.

Comments. — This species is similar to
mandihularis but can be distinguished by
its different coloration with the black pro-
podeum and first tergum and its smaller
oral opening.

Etymology. — Named for Mian Inayatul-
lah who collected the type series.

Aleiodes politiceps (Gahan),
new combination

(Figs. 3, 9, 11)

Rogns politiceps Gahan 1917:206.

Diagnosis. — Body unicolored orange, an-
tenna black, legs orange except apex of
hind tibia and all tarsi black, wings black;
body length, 7.0-9.0 mm; 60-65 anten-
nomeres; malar space shorter than basal
width of mandible; oral opening large
(Fig. 9), diameter greater than malar
space; ocell-ocular distance greater than
diameter of lateral ocellus; head smooth
and shining, face weakly costate; mesono-
tum, scutellum and mesopleuron smooth
and shining, propodeum areolate-rugose,
median carina complete; first, second and
basal half of third metasomal terga strong-
ly porcate (Fig. 11), median carina com-
plete on first and second terga; fore wing
vein Icu-a beyond IM by distance greater
than length of Icu-a (Fig. 3); hind wing
marginal cell gradually widening, vein RS
straight (Fig. 3); tarsal claws pectinate.

Type material examined. — Rogas politiceps
Gahan, holotype female, Nashville, Ten-
nessee (USNM).

Distribution. — Virginia south to Florida,
west to Arkansas and Texas. Also record-
ed from San Jose, Costa Rica (new record),
at an elevation of 1300m. Not yet recorded
from other parts of Central America.

Biology. — The usual hosts appear to be
moderately large noctuid larvae, includ-
ing Anicla infecta (Ochs.) and Pseudaletia
unipimcta (Haw.).

Comments. — This is one of the common-
est large Aleiodes species occurring in the
southern states. It is frequently taken in
Malaise traps, and in large numbers at
black lights. The entirely dark wings,
bright orange body, and strongly porcate
metasomal terga make this species quite
distinctive. It is one of the few Aleiodes
species that can be reliably identified even

108

Journal of Hymenoptera Research

without a microscope. The very coarse
sculpture of metasomal terga is unique
among the species covered in this study.

ACKNOWLEDGMENTS

We thank the curators of all the collections listed
in Methods for the loan of specimens for this study.
Kent Hampton, Kansas State University, prepared the
scanning electron micrographs and Linda Lawrence,
USDA Systematic Entomology Laboratory, prepared
the wing drawings. This research was supported, in
part, by grant DEB-930-6314 from the National Sci-
ence Foundation to S. R. Shaw. Additional support
was provided by supplemental REU grants in 1994,
1995 and 1996 (Research Experience for Undergrad-
uates).

LITERATURE CITED

Cresson, E. T. 1869. List of the North American spe-
cies of the genus Aleiodes Wesmael. Tnuisactioiis
of the Ainericnii Eittcmologicnl Sccieti/ 2:377-382.

Cresson, E. T. 1872. Hymenoptera Texana. Transac-
tions of the American Entomological Society 4:153-
292.

Fortier, J. C. 1997. Cladistics of the Aleioiies lineage of
the subfamily Rogadinae (Hymenoptera: Bracon-
idae). Unpublished Ph.D. thesis submitted to the
University of Wyoming.

Gahan, A. B. 1917. Description of some new parasitic
Hymenoptera. Proceedings of the United States Na-
tional Museum 53:195-217.

Goulet, H. and J. T. Huber. 1993. Hytneiioptera of ttie
World: an identification guide to families. Agricul-
ture Canada Publication 1894 /E, Ottawa.

Harris, R. A. 1979. A glossary of surface sculpturing.
Occasional Papers in Entomology of the California
Department of Food and Agriculture no. 28, pp. 1-
31.

Marsh, P. M. 1989. Notes on Braconidae (Hymenop-
tera) associated with jojoba {Simmondsia cliinen-
sis) and descriptions of new species. Pan-Pacific
Entomologist 65:58-67.

Marsh, P. M. and S. R. Shaw. 1998. Revision of North
American Aleiodes Wesmael (Part 3): the seriatus
(Herrich-Schaeffer) species-group (Hymenop-
tera: Braconidae, Rogadinae). Proceedings of the
Entomological Society of Washington 100(3):395-
408.

Shaw, M. R. 1983. On[e] evolution of endoparasitism:
the biology of some genera of Rogadinae (Bra-
conidae). Contributions of the American Entomolog-
ical Institute 20:307-328.

Shaw, M. R. 1994. Chapter 7, Parasitoid host ranges.
Pp. 112-144, In: Hawkins, B. A. and W. Sheehan
[eds.], Parasitoid Community Ecology. Oxford Uni-
versity Press, Oxford.

Shaw, M. R. and T. Huddleston. 1991. Classification
and biology of braconid wasps. Handbooks for the
Identification of British Insects 7:1-126.

Shaw, S. R. 1993. Systematic status of Eucystomastax
Brues and Characterization of the Neotropical
species (Hymenoptera: Braconidae: Rogadinae).
Journal of Hymenoptera Research 2:1-11.

Shaw, S. R. 1995. Chapter 12.2, Braconidae. Pp. 431-
463, In: Hanson, P. E. and I. D. Gauld |eds.|. The
Hymenoptera of Costa Rica. Oxford University
Press, Oxford.

Shaw, S. R., P. M. Marsh and J. C. Fortier. 1997. Re-
vision of North American Aleiodes Wesmael (Part
1): the pulchripes Wesmael species-group in the
New World (Hymenoptera: Braconidae, Roga-
dinae). Journal of Hymenoptera Research 6(l):10-35.

Shaw, S. R., P. M. Marsh and J. C. Fortier. 1998a. Re-
vision of North American Aleiodes Wesmael (Part
2): the ductcr Thunberg species-group in the New
World (Hymenoptera: Braconidae, Rogadinae).
Journal of Hymenoptera Research 7(l):62-73.

Shaw, S. R., P. M. Marsh and J. C. Fortier. 1998b. Re-
vision of North American Aleiodes Wesmael (Part
4): the albitibia (Herrich-Schaeffer) and praetor
(Reinhard) species-groups (Hvmenoptera: Bra-
conidae, Rogadinae). Proceedings of the Entomolog-
ical Society of Washington 100(3):553-565.

Wharton, R. A., P. M. Marsh and M. j. Sharkey, eds.
1997. Manual of New World genera of the family
Braconidae. Special Publication of the International
Society of Hynienopterists. No.l, 438 pp.

J. HYM. RES.

Vol. 8(1), 1999, pp. 109-115

Further Evidence of Male Antennal Glands in Aphelinidae: The Case
of Aphytis melimis DeBach (Hymenoptera: Aphelinidae)

R. ROMANI, N. ISIDORO, AND F. BiN

Agricultural Entomology Institute, University of Perugia, 06121 Perugia, Italy,

E-mail: Fbin@unipg.it

Abstract. — Ultrastructural investigations of the elongated male club, antennomere 6, in Aphytis
melimis DeBach have shown that a small oval ventral area, bearing numerous minute setae, is not
a sensory complex but rather the release site of a glandular complex with unicellular unit type 1.
This finding, combined with behavioral observations reported in the literature, strongly indicates
that the secretion induces sex recognition during pre-coital phase. This morpho-functional inter-
pretation is discussed in other aphelinids exhibiting similar structures on different antennomeres.

In the last few years, some peculiar an-
tennal structures of parasitoid hymenop-
terans have received increasing interest,
and have revealed new aspects for func-
tional morphology, biosystematics, and
phylogeny (Isidoro et al. 1996). In fact, the
male antennae, which were previously be-
lieved to be only sensory appendages,
have instead been demonstrated as also
having a secretory function through epi-
dermal glands involved in courtship be-
havior (Bin et al. 1997). The secretory func-
tion has been reported in families Eulo-
phidae (Dahms 1984), Scelionidae (Bin
and Vinson 1986), Platygastridae (Isidoro
and Bin 1995), Ichneumonidae (Isidoro et
al. 1997), Diapriidae (Romani et al. 1997;
Sacchetti et al. 1997) and Eucoilidae (Isi-
doro et al. submitted).

Male antennal glands were reported for
the first time in Encarsia asterobemisiae Vig-
giani et Mazzone (Pedata et al. 1995), a
species supplied with two peculiar ventral
features, respectively on antennomeres
four and five, which likely appear to be
used during pre- and post-coital phases
(Viggiani and Laudonia 1989). In another
aphelinid, Aphytis iiielinus DeBach, an im-
portant worldwide biocontrol agent of cit-
rus scales, the courtship behavior was also

described stressing the importance of an-
tennation during the precoital phase
(Gordh and DeBach 1978). The occurrence
of this behavior, along with the presence
of a "specialized sensory area" on the ven-
tral side of the male elongated club, has
prompted an ultrastructural study of this
"small oval area or plate bearing numer-
ous minute setae" (Rosen and DeBach
1979).

This study proves that in A. melimis
such an area, present on A6 in the form of
an elongated club, is the release site of ad-
jacent integumentary glands. In addition,
for the first time in hymenopterans, glands
are reported on the apical antennomere.

MATERIALS AND METHODS

A colony of A. meliuus, laboratory
reared on Aonidiella aiirantii (Maskell), was
provided by the Entomology Department
of the University of California (Riverside).

For scanning electron microscopy (SEM)
observations, 10 males, newly emerged
and anaesthetized in CO,, were beheaded
and immediately immersed in 50% etha-
nol water solution and kept overnight at
4°C. After dehydration in a graded ethanol
series, the heads with antennae were crit-
ical point dried in a Balzers Union CPD

no

Journal of Hymenoptera Research

020 unit, gold coated in a Balzers Union
SCD 040 unit, and finally examined with
a Philips XL 30.

For transmission electron microscopy
(TEM) observations, 10 males were anaes-
thetized in C02 and immediately im-
mersed in 2.5% glutaraldehyde in 0.1 M
cacodylate buffer + 5% sucrose, pH 7.2-
7.3. The apical antennomeres were de-
tached to aid fixative penetration, and left
at 4°C for 2h. After rinsing overnight in a
cacodylate buffer, the specimens were
postfixed in 1% osmium tetroxide at 4°C
for Ih and rinsed in the same buffer. De-
hydration in a graded ethanol series was
followed by embedding in Epon-Araldite
with propylene oxide as a bridging sol-
vent. Thin sections were taken with a di-
amond knife on a L.K.B. "Nova" ultra-
microtome, and mounted on collodium-
coated 50 mesh grids. Finally, the sections
were investigated with a Philips EM 400T,
after staining with uranyl acetate (20 min,
room temperature) and lead citrate (5 min,
room temperature).

RESULTS

The geniculate antennae of male A. mel-
inus consist of six antennomeres. The api-
cal antennomere, A6 or club, is elongated
and bears an oval area on the proximal
ventral side which is the release site struc-
ture (RSS) of the integumentary glands
(Fig la). This area is covered by minute,
non-socketed microtrichia and bordered
by one row of socketed trichoid sensilla
(Fig lb); while the former are not inner-
vated the latter are provided of one mech-
anosensory neuron. SEM observations of
the ventral side of A6 show the oval area
slightly depressed and partially cut off
from the surrounding club portion by in-
distinct grooves.

Serial longitudinal and cross sections of
the apical antennomere reveal a well de-
veloped glandular epithelium adhering to
the internal wall of the oval area (Fig 2a).
This glandular complex consists of nu-
merous, unicellular secretory units vary-

ing in size and shape. Each glandular cell
has a large, round and regularly shaped
nucleus which is often located in the basal
part of the cell (Fig 2b). Chromatin is not
abundant and most of it is apposed to the
nuclear membrane. The perinuclear region
of the cytoplasm contains mitochondria
with conventional cristae and abundant
free ribosomes while few signs of granular
or smooth endoplasmic reticulum were
observed. The basal plasma membrane of
the cell has deep, irregular invaginations
forming a lacunar system. The apical cell
membrane is surrounded by densely
packed microvilli, delimiting a narrow ex-
tracellular space. The cuticle associated
with the glandular epithelium is pierced
by numerous tiny pores randomly distrib-
uted. The secretory apparatus underneath
each pore is formed by a spherical cham-
ber, from which numerous cuticular fila-
ments radiate (Fig 2c). These filaments,
apparently a specialization of the external
epicuticle, have a tubular structure and
extend deeply in the extracellular space
between the microvilli of the apical cell
membrane.

DISCUSSION

The "small oval area or plate bearing
numerous minute setae" on the ventral
side of male club in A. melinus (Rosen and
DeBach 1979) is not a "specialized sensory
area" but the release site of epidermal
glands. These glands are unicellular secre-
tory units belonging to the type 1 gland
cell (Noirot and Quennedey 1974, 1991;
Quennedey 1998). The cytological features
of the secretory cells do not allow us to
attempt an interpretation on the nature of
the secretion which in other hymenopter-
ans acts on contact (Isidoro et al. 1996) or
is volatile (Felicioli et al. 1998). The pecu-
liar releasing apparatus consists of numer-
ous pores so tiny that neither the external
openings nor the material secreted can be
seen with SEM, contrary to what has been
reported for other parasitoids (Bin and
Vinson 1986; Isidoro and Bin 1995). The

Volume 8, Number 1, 1999

111

Fig. 1. Aplii/tis "iWdiks male: a) \'cntr(.)-latLTal \ievv of antcnnomcre fa (Afa) showing the relcabo bile structure
(RSS); b) detail of the RSS covered by numerous microtrichia (MI) and bordered by a single row of tactile
setae (TS).

conceivable function of no-socketed mi- Encarsia asterobemisiae has two glandular

crotrichia is that of increasing the release complexes, respectively on A3 and A4, be-

surface while that of socketed trichoid longing to the same type 1 but different in

sensilla is that of perceiving tactile stimuli, cytological characteristics and release site

112

Journal of Hymenoptera Research

5 tjm

ilmf-^-ar'

Fig. 2. Apln/ti-i niiiniu^ male: j) cross section ot antennomere 6 at about half level showing the extension of
the glandular area (GA) on the ventral side; b) perinuclear detail of a secretorv cell; c) apical detail of a
secretory cell showing the tiny cuticular pores (P). N, nucleus; EF, epicuticular filaments; H, haemocele; M,
mitochondrion; MV, microvilli; R, ribosomes.

Volume 8, Number 1, 1999

113

features. This could indicate a difference
in composition and role of the relative se-
cretions (Pedata et al. 1995). From these
two ascertained cases it appears that the
so called sensorial complexes described in
other aphelinids need to be investigated to
define their real nature. Morphological
and behavioral observations strongly sug-
gest in fact that glands are present on dif-
ferent antennomeres involving the scape
(Al), from one to 3 intermediate anten-
nomeres (A3 to A5) or the apical anten-
nomere (A6). Some examples are as fol-
lows:

On Al of Physcus testaceus Masi there
are special structures which could be re-
lated to a gland (Viggiani et al. 1986) and
something similar occurs in Pteroptryx chi-
nensis (Howard) (Viggiaru and Ren 1993).
Numerous species of Encarsia Foerster
have one up to three antennomeres, A3-
A5, which may or may not be noticeably
enlarged, and ventrally provided of one
specialized structure or two different ones.
When two or more antennomeres are
glanded the release sites are of two types;
furthermore, one enlarged antennomere
does not necessarily bear a specialized
structure, as A3 in the case of £. asterobem-
isiae (Pedata et al. 1995). Therefore, some
of the following examples having two or
three enlarged antennomeres may need to
be confirmed: A3 in E)icar^ia albiscutellum
(Girault) (Viggiani 1985), A4 in Encarsia
aleurotubae Viggiani, £. hemdoui (Girault)
(Viggiani 1987), A3-A4 in Encarsia olivina
(Masi) (Viggiani and Mazzone 1982), £. g(-
gas (Tchumakova), £. optileiita Silvestri
and £. perniciosi (Tower) (Viggiani and
Laudonia 1989), A3-A5 in Encarsia antiopa
(Girault) (Viggiani 1985).

As regards the apical antennomere the
A6 of several species of Aphytis Howard,
as illustrated by Rosen and DeBach (1979),
have external features similar to those de-
scribed for melinus, and therefore could
have glands: A. cochereaui DeBach and Ro-
sen, A. fabresi DeBach and Rosen, A. chi-
lensis Howard, A. coluinbi (Girault), A. par-

amaculicornis DeBach and Rosen, A. dias-
pidis (Howard), A. lingnanensis Compere,
A. roseni DeBach and Gordh, A. pilosus
DeBach and Rosen.

Behavioral observations indicate that
the antennation mechanism during mat-
ing is different in species having one or
two glanded antennomeres. While the sin-
gle release site seems to be logically relat-
ed to the pre-coital phase, e.g. triggering
sex-recognition, female stimulation or fe-
male sedation (Gordh and DeBach 1978;
Viggiani t'f al. 1986), the presence of a sec-
ond and different release site has been ob-
served in species performing also a post-
coital phase, e.g. E. asterobemisiae (Pedata
et al. 1995).

These chemo-physical mechanisms can
only partially explain the sexual isolation
since there also are other chemicals in-
volved in mating behavior. In some spe-
cies of Aphytis a female sex pheromone
(Rao and DeBach 1969) and a male aph-
rodisiac pheromone (Khasimuddin and
DeBach 1975), whose sources are as yet
unknown (Gordh and DeBach 1978), have
in fact been reported. In another aphelin-
id, Aphelinus asychis Walker, the mate
finding is mediated by a female trail sex
pheromone deposited in a yet unknown
way on the substrate while walking (Fau-
vergue et al. 1995).

As it is being shown in other parasitoid
groups (Bin et al. 1997; Isidoro et al. sub-
mitted) the glandular nature of the anten-
nal structures provides a new perspective
to define the sex selection strategies of
aphelinids as well as additional characters
for taxonomy and phylogeny.

ACKNOWLEDGMENTS

We are verv grateful to I'rof. R. F. Luck, who kind-
ly supplied insects. We also thank C. Dentini for tech-
nical assistance with fixation and embedding of the
specimens and A. Mommi for film processing and
photographic printmg. SEM and TEM pictures were
made using the Electron Microscopy Center of Pe-
rugia University (CUME). This research was finan-
cially supported by the Ministry for University and
Scientific Technological Research (M.U.R.S.T. 40%).

114

Journal of Hymenoptera Research

The authors have contributed equally to different as-
pects of this paper.

LITERATURE CITED

Bin, F. and S. B. Vinson. 1986. Morphology of the
anteruial sex-gland in male Trissolcus basalis
(Woll.) (Hymenoptera: Scelionidae), an egg par-
asitoid of the green stink bug, Nczara viriiiula
(Hemiptera: Pentatomidae). bilcrnationnl journal
of Insect Morphologfi and Enilm/ology 15: 129-138.

Bin, F., N. Isidoro, R. Romani, and S. B. Vinson. 1997.
Antennal functional areas for sex recognition in
some parasitoid hymenopterans. Boletin de la
Asociacion cspaiiola de Entomologia, Suplemento al
Volumen n°2V. 68-69.

Dahms, E. C. 1984. An interpretation of the structure
and function of the antennal sense organs of Mel-
tittohia australica (Hymenoptera: Eulophidae)
with the discovery of a large dermal gland in the
male scape. Memoirs of the Queensland Museum
21(2): 361-377.

Fauvergue, X., K. R. Hopper, and M. F. Antolin. 1995.
Mate finding via a trail sex pheromone by a par-
asitoid wasp. Proceedings of the National Academy
of Siences of the U. S. A. 92: 900-904.

Felicioli, A., N. Isidoro, R. Romani, F. Bin, and M.
Pinzauti. 1998. Ethological and morphological
analysis of mating behaviour in Osniia coriiuta
Latr. (Hymenoptera, Megachilidae). Insect Social
Life 2: 137-144.

Gordh, G. and P. DeBach. 1978. Courtship behaviour
in the Aphytis lingnanensis group, its potential
usefulness in taxonomy, and a review of sexual
behavior in the parasitic Hymenoptera (Chalci-
doidea: Aphelinidae). Hilgardia 46: 37-75.

Isidoro, N. and F. Bin. 1995. Male antennal gland of
Atnitus spiniferus (Brethes) (Hymenoptera: Platy-
gastridae), likely involved in courtship behavior.
International journal of Insect Morpliolog\/ and Em-
bryology 24: 365-373.

Isidoro, N., F. Bin, S. Colazza, and S. B. Vinson. 1996.
Morphology of antennal gustatory sensilla and
glands in some parasitoid Hymenoptera with hy-
pothesis on their role in sex and host recognition.
journal of Hymenoptera Research 5: 206-239.

Isidoro, N., F. L. Wackers, R. Romani, and F. Bin.
1997. Antennal tyloids are release structures of
sex recognition pheromone in Pimpla turionellae
(L.) (Hymenoptera, Ichneumonidae). Boletin de la
Asociacion espanola de Entonwlogia. Suplemento al
Volumen «°21: 70-71.

Khasimuddin, S. and P. DeBach. 1975. Mating behav-
iour and evidence of a male sex pheromone in
species of the genus Aphytis. Annals of the Ento-
mological Society of America 68(5): 893-896.

Noirot, C. and A. Quennedey. 1974. Fine structure of
insect epidermal glands. Annual Review of Ento-
mology 19: 61-80.

Noirot, C. and A. Quennedey. 1991. Glands, gland
cells, glandular units: some comments on ternvi-
nology and classification. Annates de la Societe en-
tomologique de France 27: 123-128.

Pedata, P. A., N. Isidoro, and G. Viggiani. 1995. Evi-
dence of male sex glands of the antennae of En-
carsia asterohemisiae Viggiani et Mazzone (Hy-
menoptera; Aphelinidae). Bollettino del Laboratorio
di Entonuilogia Agraria "Fdippo Silvestri" 50 (1993):
271-280.

Quennedey, A. 1998. Insect epidermal gland cells: ul-
trastructure and morphogenesis. Microscopic
Anatomy of bwertebrates 11 A: 177-207.

Rao, S. V. and P. DeBach. 1969. Experimental studies
on hybridization and sexual isolation between
some Apthytis species (Hymenoptera: Aphelini-
dae). I. Experimental hybridization and an inter-
pretation of evolutionary relationships among
the species. HUgardia 39: 515-553.

Romani, R., N. Isidoro, and F. Bin. 1997. Antennal
structures and sex recognition in Trichopria dro-
sophilae (Hymenoptera, Diapriidae). Boletin de la
Asociacion espafwla de Entomologia, Suplemento al
Volumen n'21: 142.

Rosen, D. and P. DeBach. 1979. Species of Aphytis of
the world (Hymenoptera: Aphelinidae). W. Junk, The
Hague, pp. 1-801.

Sacchetti, P., A. Belcari, F. Fagnani, N. Isidoro, and F.
Bin. 1997. Antennal structures of CopHera occiden-
talis mues. (Hymenoptera, Diapriidae) involved
in mating behaviour and host recognition. Boletin
de la Asociacion espafwla de Entonwlogia. Supleme}i-
to al Volumen n° 21: 72.

Viggiani, G. 1985. Additional notes and illustrations
on some species of aphelinids described by A. A.
Girault and A. P. Dodd in the genera Coccophagus
Westw., Encarsia Foerst. and Prospaltella Ashm.
(Hym.: Chalcidoidea). Bollettino del Laboratorio di
Entomologia Agraria "Filippo Silvestri" 42: 233-
255.

Viggiani, G. 1987. Le specie italiane del genere En-
carsia Foerst. (Hymenoptera: Aphelinidae). Bol-
lettino del Laboratorio di Entomologia Agraria "Filip-
po Silvestri" 44: 121-179.

Viggiani, G. and P. Mazzone. 1982. Antennal sensilla
of some Encarsia Foerster (Hymenoptera: Aphel-
inidae), with particular reference to sensorial
complexes of the male. Bollettino del Laboratorio di
Entomologia Agrarui "Filippo Silvestri" 39: 19-26.

Viggiani, G., D. Battaglia, and R. Jesu. 1986.
L'accoppiamento di Physcus testaceus Masi (Hym.
Aphelinidae), con notizie preliminari sulla strut-
tura dello scapo antennale maschile. Bollettino del
Laboratorio di Entonwlogia Agraria "Filippo Silves-
tri" 43: 3-6.

Viggiani, G. and S. Laudonia. 1989. Su alcuni com-
plessi sensoriali delle antenne maschili di tre spe-

Volume 8, Number 1, 1999 115

cie del genere Eiicarsia Foerster (Hymenoptera: Viggiani, G. and H. Ren. 1993. New species and rec-

Aphelinidae) e il loro rapporto con le fasi ords of Aphelinidae (Hymenoptera: Chalcidoi-

dell'accoppiamento. Bcllettino iU-I Laboratorio di dea) from China. Bollettiiw del Lahoratorio di En-

Entcmologm Agiaim "Filippo Silvestri" 45 (1988): tomologia Agraria "FUippo Silvestri" 48 (1991):

67-75. 219-239.

J. HYM. RES.
Vol. 8(1), 1999, pp. 116-119

A New Species Group and Two New Species of Enderomphale
Girault (Hymenoptera: Eulophidae) from North America

John LaSalle

Unit for Parasitoid Systematics, CABI Bioscience UK Centre (Ascot), Department of Biology,
Imperial College at Silwood Park, Ascot, Berks., SL5 7PY, UK

Abstract. — Two new species of Euderomphale, E. sinuata and E. suzannae, are described from
North America. These represent a distinct species group within the genus Euderomphale, which is
here referred to as the sinuata group. £. sinuata is brachypterous, and this represents the first
record of brachyptery in the Euderomphalini.

Euderomphale is a cosmopolitan genus
containing parasitoids of whiteflies. Thir-
teen described species in this genus were
listed by LaSalle and Schauff (1994). No
comprehensive keys to species have been
published, but there are keys to limited
numbers of regional species (Erdos 1966,
for Hungary; Hulden 1986, for Finland).
Euderomphale belongs in the tribe Euder-
omphalini. This tribe, containing parasit-
oids of whiteflies, was treated at the world
level by LaSalle and Schauff (1994), who
presented keys to, diagnoses for, and sup-
port for the monophyly of seven included
genera in two genus groups. The present
paper describes two new species which
represent a new species group within the
genus Euderomphale, the sinuata group. Eu-
deromphale is thus separated into two spe-

cies groups: the flavimedia group (based on
the valid name for the type species) con-
tains 13 species (LaSalle and Schauff 1994),
and the sinuata group contains the two
species described in this paper. Biology
for the two new species in the sinuata
group is unknown, but it is presumed
they are parasitoids of whiteflies like all
other members of the Euderomphalini. £.
sinuata is brachypterous, and this repre-
sents the first record of brachyptery in the
Euderomphalini. It is also the first record
of brachyptery in females of any Entedon-
inae, although brachyptery is known in
males of Microdonophagus ivoodlei/i Schauff
(Schauff 1986). Kerrich (1973) reported the
males of Pediobius arnconae Kerrich to be
micropterous, although in this species the
wing is only slightly reduced and does not
have a highly modified shape.

KEY TO SPECIES GROUPS WITHIN EUDEROMPHALE AND SPECIES OF THE
SINUATA GROUP (FEMALES)

1. Axilla smoothly rounded anteriorly (Fig. 4). Vertex smoothly rounded posteriorly (Fig. 4),
with the lateral ocellus placed on the top of the head. Prepecti.is partially fused to meso-
pleuron anteriorly flavimedia group (discussed by LaSalle and Schauff 1994)

- Axilla sinuate and concave anteriorly (Figs. 1, 3). Vertex with a distinct transverse carina;
lateral ocellus behind this carina (Figs. 1-3). Prepectus free, not fused to mesopleuron
sinuata group 2

2. Brachypterous; wings short, not extending past first gastral tergite (Fig. 1), infumated.
Axilla with anterior margin deepl concave, anteromedial lobe narrower and more distinct

(cf. Figs. 1 and 3). Gaster uniformly dark brown £. simiata LaSalle

Volume 8, Number 1, 1999

117

Figs. 1-4. Eudcromphale species, head and mesosoma. 1-2. Eudewmphale sinuata. 3. Eiideromphale suzannae. 4.
Eudcroniphiilc sp. (flaviiiicdin group).

Not brachypterous; wings normal, extending to or slightly past apex of abdomen, hyaline
or very lightly infumate. Axilla with anterior margin shallowly concave, anteromedial lobe
wider and less distinct (cf. Figs. 1 and 3). Gaster yellow or pale brown, with several dark
transverse stripes E. suzamiae LaSalle

Characters for the Eiidero»ip}ialc genus
group (within the Euderomphalini): Head
usually smooth to lightly sculptured.
Frontal suture, when present, placed just
ventral to median ocellus. Vertexal suture
present, either placed between the ocelli.

or behind the ocelli. Malar sulcus usually
present, although sometimes incomplete.
Funicle with 1 or 2 segments. Dorsum of
mesosoma more or less flattened and usu-
ally smooth to lightly sculptured, gener-
ally when distinct sculpture is present it is

118

Journal of Hymenoptera Research

incised. Midlobe of mesoscutum usually
with only 1 or 2 pairs of setae. Scutellum
distinctly wider than long, with 1 or 2
pairs of setae. Axilla either completely
separated from mesoscutum by a com-
plete suture, or partially fused to meso-
scutum. Submarginal vein with one or
two setae on dorsal surface. See LaSalle
and Schauff (1994) for further discussion
and illustration of these characters, and
characters to define the Euderomphalini.

Characters for Euderomphale: Scrobal su-
ture and frontal suture absent. Vertexal
suture extending from the eye margin be-
tween the median and lateral ocelli. Malar
sulcus present, although incomplete and
extending away from the mouth margin
(postero-laterally from the eye). Antenna
with two funicular segments, although the
first is reduced to not much larger than
the size of an anellus (the small first fu-
nicular segment can generally be distin-
guished from an anellus by the presence
of setae). Mesosoma always black, non-
metallic, more or less flattened in lateral
view, sculpture on dorsal surface usually
either finely incised or smooth. Midlobe of
mesoscutum generally with 2 pairs of se-
tae near the anterior margin. Scutellum
distinctly wider than long, and with two
pairs of setae: a larger one near the middle
of the scutellum and a smaller one at the
posterior margin. Axilla large (as wide as
long), and completely separated from me-
soscutum by a sulcus. Dorsal surface of
submarginal vein with 2 setae (a single
seta in the brachypterous sinuata). See
LaSalle and Schauff (1994) for further dis-
cussion and illustration of these charac-
ters.

Characters for the flainmedia species
group in Eiidewniphale: Vertex smoothly
rounded posteriorly. Axilla smoothly
rounded anteriorly. Prepectus reduced
and partially fused to the mesopleuron.

Characters for the sinuata species group
in Euderomphale: Vertex with a distinct
transverse carina; lateral ocelli placed pos-
terior to this carina. Axilla sinuate and

concave anteriorly. Prepectus reduced
(strongly so in sinuain) but separate, not
fused to mesopleuron.

Euderomphale sinuata LaSalle, sp.n.

(Figs. 1-2)

Female. — Length 0.75 mm. Head and
mesosoma black, gaster dark brown.
Scape yellow, pedicel and flagellum
brown. All coxae black; fore and middle
femora and tibiae dark brown to black;
hind femur brown to light brown, hind
tibia brown basally, light brown apically;
tarsi brown, with apical segment dark
brown. Wings infumated. Head with del-
icate, small, regular incised sculpture. Ver-
texal suture present near the eye margin;
joining or running just anterior to vertexal
carina. Mesosoma with delicate, small,
regular incised sculpture. Axilla with an-
terior margin distinctly concave, antero-
medial lobe narrower and more distinct
than in suzannae. Prepectus quite reduced,
separate from the mesopleuron. Wings
greatly reduced, not reaching apex of first
gastral tergite. Forewing with a single
strong seta on submarginal vein and an-
other on marginal vein. Gaster with tergite
5 extending over the telescoped tergites 6
and 7 and short ovipositor.

Holotype female, USA, Florida, Levy
Co., 5 km SW Archer, 2-15.ix.l987, FIT,
turkey oak shrubs, BRC HYM. TEAM (Ca-
nadian National Collection, Ottawa).

Euderomphale suzannae LaSalle, sp.n.

(Fig. 3)

Fe?n(7/e.— Length 0.90-0.95 mm. Head
and mesosoma black, gaster yellow with
transverse brown stripe on tergites 2-5,
and a median longitudinal brown stripe
on tergite 1. Scape yellow, pedicel and fla-
gellum brown. Fore and middle coxae
dark brown to black, hind coxa brown to
light brown; fore femur dark brown; mid-
dle femur light brown, dark brown dor-
soapically, hind femur yellow; fore tibia
yellow to light brown, dark brown dor-
sally, middle tibia dark brown basally.

Volume 8, Number 1, 1999

119

yellow to light brown apically, hind femur
yellow to light brown; all tarsi yellow to
light brown. Wings hyaline or very lightly
infumated. Head with small, regular
slightly raised sculpture (slightly stronger
than in sinuata). Vertexal suture present
near the eye margin; joining or running
just anterior to vertexal carina. Mesosoma
with small, regular incised sculpture
(slightly stronger than in sinuata). Axilla
with anterior margin moderately concave,
anteromedial lobe broader and not as dis-
tinct as in sinuata. Prepectus moderately
reduced, separate from the mesopleuron.
Wings normal. Forewing with two strong
setae on the submarginal vein. Gaster with
last tergite not extending over the ovipos-
itor.

Holotype female, USA, Missouri,
Wayne Co., Williamsville, 10-26.vi.l987,
MT, J. Becker (Canadian National Collec-
tion, Ottawa). Paratype female, USA, Mis-
souri, Wayne Co., Williamsville, vii.1987,
MT, J. Becker (Canadian National Collec-
tion, Ottawa).

Et^mologi/. — Named for Suzanne Lewis.

ACKNOWLEDGMENTS

Space and facilities during this study were kindly
provided by the Department of Entomology, The
Natural History Museum, London; technical assis-
tance from the SEM and photography units of the
BMNH is also gratefully acknowledged. Special
thanks to Suzanne Lewis (BMNH) and Georgina
Godwin (CABI) for assistance with the taking of the
photomicrographs, and Nick Hayes (BMNH) for the
printing of the photomicrographs.

LITERATURE CITED

Erdos, J. 1966. Nonnulae Eulophidae novae Hungar-
icae (Hymenoptera: Chalcidoidea). Annales His-
torico-Ntituralci Musci Natiouiilii^ Hungarici, pars
Zoologica 58: 395-420.

Hulden, L. 1986. The whiteflies (Homoptera: Aley-
rododea) and their parasites in Finland. Nctiilae
Entoniologtccic 66: 1—10.

Kerrich, G.J. 1973. A revision of the tropical and sub-
tropical species of the eulophid genus Pediobius
Walker (Hymenoptera: Chalcidoidea). Bulletin of
the British Museum (Natural Histon/I. Entomology
29: 115-200.

LaSalle, J. and Schauff, M.E. 1994. Systematics of the
tribe Euderomphalini (Hymenoptera: Eulophi-
dae): parasitoids of whiteflies (Homoptera: Al-
eyrodidae). Systematie Entomology 19: 235-258.

Schauff, M.E. 1986. Microdonophagus, a new entedon-
ine genus (Hymenoptera: Eulophidae) from Pan-
ama. Proceedings of the Entomological Society of
Washington 88: 167-173.

J. HYM. RES.
Vol. 8(1), 1999, pp. 120-125

A New Genus and Two New Species of Brachypterous Lysiterminae

(Braconidae)

Sergey A. Belokobylskij and Donald L. J. Quicke

(SAB) Zoological Institute, Russian Academy of Sciences, St. Petersburg 199034, Russia;

(DLJQ) Unit of Parasitoid Systematics, CABI Bioscience UK Centre (Ascot), Department of

Biology, Imperial College at Silwood Park, Ascot, Berks SL5 7PY, UK and Department of

Entomology, The Natural History Museum, London SW7 5BD, UK

Abstract. — A new genus Neolysitemtus gen. n. and two new species N. tumeri sp. n. and N.
spinator sp. n. (tribe Lysitermini), both from South Africa, are described and illustrated. These
are the first brachypterous species belonging to the subfamily Lysiterminae.

Aptery and brachyptery are known in
only a few subfamilies of Braconidae, in-
cluding the Alysiinae, Aphidiinae, Blaci-
nae, Doryctinae, Hormiinae, Masoninae,
Pambolinae and Orgilinae. In the first of
these, aptery is probably a result of their
hosts dwelling within subterranean or
filthy habitats and the associated difficulty
in negotiating soil particles or costs of
wing fouling, however, in the others, no
obvious host habitat association is appar-
ent, though many other brachypterous
parasitoids are associated with stored
products or with tree trunks /tall shrubs.
The brachypterous and apterous braco-
nids are more or less equally split between
ecto- and endoparasitoids and between
idiobiont and koinobiont taxa, suggesting
that at least in this family when consid-
ered at subfamily level, brachyptery is not
dependent upon major life history fea-
tures. However, formal comparative anal-
ysis at species level will be required before
firm conclusions can be reached.

The Lysiterminae have previously often
been regarded as a tribe within either the
Rogadinae, Exothecinae or the Hormiinae
(van Achterberg 1976, 1982; Quicke & van
Achterberg 1990; Wharton 1993; Beloko-

bylskij 1993) but was afforded subfamily
status by van Achterberg (1993, 1995) and
van Achterberg & Steiner (1996) because it
shares no obvious synapomorphies with
either Rogadinae s.s. or Hormiinae. This
arrangement seems best at present since
there is considerable doubt about the
monophyly of the Hormiinae as treated
(conservatively) by some workers (see for
example, Whitfield & Wharton 1997). New
molecular data, as well as the investiga-
tion of more character systems, will be re-
quired to resolve this (see Quicke et al.
1992; Belshaw et al. 1998).

No apterous or brachypterous species
have previously been described in the
Lysiterminae. Although little is known bi-
ologically about the Lysiterminae, they
appear to have diverse host associations
including being ectoparasitoids, or pre-
sumed ectoparasitoids, of bagworms (Psy-
chidae) and web-feeders (Xyloryctidae =
Stenomidae), and also apparently endo-
parasitoids of Orthoptera (Hedqvist 1963;
Wharton 1993; van Achterberg & Steiner
1996). Interestingly, all of these hosts live
in retreats involving silk.

The Lysiterminae Tobias, 1968 includes
only seven genera in the Old World (van

Volume 8, Number 1, 1999

121

Achterberg 1995; Belokobylskij 1995; van
Achterberg & Steiner 1996). Unfortunate-
ly, differences between lysitermine genera
mostly concern fore wing venation,
though members of the subtribe Triter-
mina, with 2 genera {Triiermus van Ach-
terberg and Afroiritermus Belokobylskij)
differ from the other genera of Lysitermi-
nae in having fused, and therefore im-
mobile, 1st and 2nd metasomal tergites
(Belokobylskij 1993). Because of the major
reliance of lysitermine systematics on
wing venation, interpreting the relation-
ships of Neolysitermus gen. n., with its re-
duced wings, is not so straightforward.
Apart from the reduced wings, Neolysiter-
mus gen. n. has a distinct median emar-
gination of the posterior margin of 3rd
metasomal tergite — a character not known
in any other species of Lysiterminae.

TERMINOLOGY AND COLLECTIONS

The wing venation terms used largely
follow Tobias (1986). The following abbre-
viation are used: POL — postocellar line;
OOL — ocular-ocellar line; Od — maximum
diameter of lateral ocellus. Collections are
abbreviated as follows: The Natural His-
tory Museum, London (BMNH); Zoologi-
cal Institute, Russian Academy of Scienc-
es, St. Petersburg, Russia (ZIP).

Neolysitenmts Belokobylskij and
Quicke, new genus

Type species: Neolysitermus turucri new species

Etymology. — From "neo" (new) and
" Lysitermus" the genus of the subfamily
Lysiterminae.

Diagnosis. — The position of this new ge-
nus is not clear. Differences between gen-
era of the tribe Lysitermini are connected
mostly with the venation of the fore wing,
but species of Neolysitermus gen. n. have
very short wings with reduced venation.
This new genus differs from other genera

of Lysitermini (in addition to the very
short wings) by the presence of distinct
median emargination on the posterior
margin of 3rd tergite, a character that is
absent in all described species of this tribe.

Description. — Head weakly transverse
(Figs 2, 11). Scapus (Fig. 4, 13) thick, with
distinct round cut in outer apical side. Fla-
gellum filiform. Apical flagellomere with-
out apical spine. Maxillary palpi 6-seg-
mented, labial palpi 4-segmented; 3rd la-
bial segment distinctly shortened. Malar
suture absent. Clypeus strongly convex
(Figs 3, 12). Eyes glabrous. Occipital carina
fused with hypostomal one higher man-
dibles. Hypostomal keel distinct. Ocelli
small, forming an almost equilateral tri-
angle. Mesosoma reduced, but with all su-
tures and depressions (Figs 7, 8, 16, 17).
Propleura with longitudinal median cari-
na on basal half. Pronotum with distinct
lateral median corners (see Figs 8, 17). No-
tauli deep and wide. Lateral lobes of me-
soscutum with oval depressions postero-
laterally. Scutellum with high lateral cari-
nae. Sternauli rather deep, wide, oblique,
coarsely crenulate. Prepectal carina very
strong. Postpectal carina absent. Meta-
pleural flange long and narrow. Propodeal
areola incomplete or absent. Fore wing
very short and narrow, stylet-shaped or
oval. Hind tibia and tarsus slender. Hind
tibial spurs very short. Hind basitarsus
0.9-0.95 X as long as 2nd-5th segments
combined. First and second metasomal
tergites not fused, mobile (Figs 9, 10, 18,
19). Dorsope of first tergite small. Second
suture deep. Dorsal carina of first tergite
semicircularly united basally. Third tergite
without spines and carina posteriorly,
with deep and rather narrow median
emargination, with small single tooth ven-
tro-laterally (Figs 9, 18).

Distribution. — Afrotropical Region (South
Africa).

122

Journal of Hymenoptera Research

KEY TO SPECIES OF NEOIYSITERMUS

Antenna 17-segmented; apical segment dark. Length of mesosoma 1.7-1.8 x height. Scu-
tellum flat. Propodeum without lateral spines. Metasoma narrow, its length 2.3-2.4 x max-
imum width. First metasomal tergite longer, apically 1.2-1.3 x wider than long

N. tumeri sp. n.

Antenna 14-segmented; apical segment whitish. Length of mesosoma 1.3-1.4 X height.
Scutellum strongly pointedly convex. Propodeum with distinct pointed lateral spines. Me-
tasoma wide, its length twice maximum width. First metasomal tergite shorter, apically
1.7-1.8 X wider than long N. spinator sp. n.

Neolysitermus tumeri Belokobylskij and
Quicke, new species

(Figs 1-10)

Holotype female. — "S. Africa. R.E. Turner.
Brit. Mus. 1924-136", "Port St. John, Pon-
doland. 6-25. Feb. 1924" (BMNH).

Paratypes. — 1 female, same data as ho-
lotype (BMNH); 1 female, "S. Africa. R.E.
Turner. Brit. Mus. 1924-97", "Port St.
John, Pondoland. Jan. 1924" (ZIP); 1 fe-
male, "S. Africa. R.E. Turner. Brit. Mus.
1924-109", "Port St. John, Pondoland. 29.1
- 5.II.1924" (BMNH).

Description. — Female. Body length 1.8-
2.3 mm. Head: 1.5-1.6 X wider than me-
dially long; 1.7-1.8 X width of mesoscu-
tum. Antennae 17-segmented. Scapus 1.5-
1.6 X as long as wide. First flagellar seg-
ment 3.7-4.2 X longer than apically wide,
slightly longer than 2nd segment. Penul-
timate segment 3 X as long as wide, 0.7 X
as long as 1st flagellar segment, 0.9-1 X
as long as apical segment. Temple roundly
narrowed behind eyes. Transverse diam-
eter of eye 1.4-1.6 X length of temple (dor-
sal view). POL 1.2-1.5 X Od, 0.3-0.4 X
OOL. Antennal socket diameter 0.8-1 X
distance between sockets, almost twice
distance between socket and eye. Eye 1.2-
1.3 X taller than broad. Cheek height 0.7-
0.8 X height of eye, 1.6-1.7 X basal width
of mandible. Face 1.2-1.4 X wider than
eye height and 1.2 (wider than height of
face and clypeus combined. Clypeus with
distinct narrow flange along lower mar-
gin. Hypoclypeal depression oval, 0.7-0.8

X wider than distance from depression to
eye. Head distinctly and roundly nar-
rowed below eyes. Mesosoma: 1.7-1.8 X
longer than high, almost twice longer than
wide. Median lobe of mesoscutum with
antero-lateral teeth. Prescutellar depres-
sion long, with median carina, granulose-
crenulate, 0.5-0.6 X as long as scutellum.
Scutellum rather flat. Subalar depression
deep, narrow, crenulate. Mesopleura with-
out median furrow. Propodeum without
lateral spines. Fore wing stylet-shaped,
3.2^.5 X longer than wide, 0.35-0.4 X
length of mesosoma. Hind femur 4.5-5 X
longer than wide. Hind tarsus 0.8-0.85 X
hind tibia. Second tarsal segment 0.3-0.35
X length of 1st segment, almost as long as
5th segment (excluding pretarsus). Meta-
soma: Convex and narrow, its length 2.3-
2.4 X maximum width (on the level of
middle of 2nd tergite), 1.4-1.5 x as long
as mesosoma. First tergite distinctly and
roundly narrowed towards base, rather
long, its apical width 1.2-1.3 X length,
2.0-2.2 X its minimum width. Second ter-
gite almost as long as basally wide, 0.8-
0.9 X maximum width, 1.3-1.4 X length
of 1st tergite, 0.8-0.9 X length of 3rd ter-
gite. Third tergite regularly and almost
linearly narrowed toward apex. Oviposi-
tor sheath 0.5-0.6 X as long as metasoma,
0.8-0.9 X as long as mesosoma. Sculpture
and pubescence: Head densely granulate,
vertex strongly granulate and usually
with fine rugae; face finely granulate. Me-
sothorax densely and entirely granulate.
Propodeum with median carina, which is

Volume 8, Number 1, 1999

123

Figs. 1-10. Neoli/sitenuKs tunwri gen. et sp. n.: 1 — head, frontal view; 2 — head, dorsal view; 3 — head, lateral
view; 4 — basal and apical segments of antenna; 5 — hind femur; 6 — hind tibia; 7 — thorax, lateral view; 8 —
thorax, dorsal view; 9 — metasoma, dorsal view; 10 — metasoma, lateral view.

1-1.4 X as long as furca; basolateral areas
densely granulate; posterior half of pro-
podeum transversely striate with dense
granulation and 2 short submedian lon-
gitudinal carinae. Hind legs finely granu-
late. First to 3rd metasomal tergites
strongly striate, with fine and dense trans-
verse rugae between striae; striae at 2nd
and 3rd tergites weakly convexly curved.
Setae on dorsal side of hind tibia sparse,
short and not erect. Colour: Body light
reddish brown, sometimes dorsally dark-
er. Head yellow. Antenna yellowish
brown, submedially slightly darkened, 5-
6 subapical segments whitish, apical seg-
ment dark. Palps pale yellow. Legs yel-
low.

Male. — Ur^known. .-•

Neolysitermus spinator Belokobylskij
and Quicke, new species

(Figs 11-19)

Holotype female. — "S. Africa. R.E. Turner.
Brit. Mus. 1924-136", "Port St. John, Pon-
doland. 6-25. Feb. 1924" (BMNH).

Paratypes. — 1 female, same data as ho-
lotype (BMNH).

Description. — Female. Body length 1.8-
2.1 mm. Head: 1.4-1.5 X wider than me-
dially long; 1.6-1.9 X width of mesoscu-
tum. Temple roundly narrowed behind
eyes. Transverse diameter of eye 1.7-2 X
length of temple (dorsal view). POL 1-1.3
X Od, 0.3-0.4 X OOL. Antennal socket di-
ameter 1.5-2 X distance between sockets,

124

Journal of Hymenoptera Research

<23 ' '

Figs. 11-19. Neolysitermus spinator gen. et sp. n.: 11 — head, dorsal view; 12 — head, lateral view; 13 — basal
and apical segments of antenna; 14 — hind tibia; 15 — hind femur; 16 — thorax, lateral view; 17 — thorax, dorsal
view; 18 — metasoma, dorsal view; 19 — metasoma, lateral view.

almost 1.5 X distance between socket and
eye. Eye 1.3-1.4 x as taller than broad.
Cheek height 0.8 X height of eye, nearly
twice basal width of mandible. Face 1.3 x
wider than eye height and 1.25 x wider
than height of face and clypeus combined.
Clypeus with distinct narrow flange along
lower margin. Head distinctly and round-
ly narrowed below eyes. Antennae 14-seg-
mented. Scapus 1.5-1.6 x as long as wide.
First flagellar segment 3.7-4.0 X longer
than apically wide, 1-1.2 X as long as 2nd
segment. Penultimate segment 2.5-2.7 X
as long as wide, 0.8 X as long as 1st fla-
gellar segment, as long as apical segment.
Mesosoma: 1.3-1.4 X longer than high,
1.5-1.8 X longer than wide. Median lobe
of mesoscutum with antero-lateral teeth.
Prescutellar depression long, with median
carina, granulose-crenulate, 0.6-0.7 X as
long as scutellum. Scutellum strongly

pointedly convex. Subalar depression
deep, narrow, granulose-crenulate. Meso-
pleura with fine oblique median furrow.
Propodeum with distinct pointed lateral
spines and mediobasal flat small lobe.
Fore wing oval and short. Hind femur 4.5-
5 X longer than wide. Hind tarsus 0.8 x
length of hind tibia. Second tarsal segment
0.3 X as long as 1st segment, nearly as
long as 5th segment (excluding pretarsus).
Metasoma: Convex and wide, nearly
twice longer than maximally wide (at lev-
el of middle of 2nd tergite), 1.7-1.9 X
longer than mesosoma. First tergite dis-
tinctly and roundly narrowed towards
base, short, its apical width 1.7-1.8 x
length, 2.4 X its minimum width. Second
tergite 0.9-1 x longer than basally wide,
0.8 X its maximum width, 1.7 x length of
1st tergite, as long as 3rd tergite. Third ter-
gite regularly and roundly narrowed to-

Volume 8, Number 1, 1999

125

ward apex. Ovipositor sheath 0.3-0.4 X as
long as metasoma, 0.6-0.8 x as long as
mesosoma. Sculpture and pubescence:
Head largely densely granulate, vertex
sometimes (paratype) with rugae, frons
striate with granulation, face finely gran-
ulate. Mesothorax densely and almost en-
tirely granulate. Propodeum with trans-
verse median carina between spines; ba-
solateral areas densely granulate; posteri-
or half rugulose-striate, with fine
granulation. Hind legs finely granulate.
First to 3rd metasomal tergites strongly
longitudinally striate, with fine and dense
transverse rugae between striae; striae of
3rd tergite distinctly converging posteri-
orly. Setae on dorsal side of hind tibia
sparse, short and not erect. Colour: Body
light reddish brown, metasoma partly
darker. Head yellow. Antenna yellowish
brown, submedially slightly darkened, 5
distal segments whitish, apical segment
dark. Palps pale yellow. Legs yellow.
Male. — Unknown.

ACKNOWLEDGEMENTS

This work was supported by the Natural Environ-
ment Research Council, Initiative in Taxonomy.

LITERATURE CITED

Achterberg, C. van. 1976. A preliminary key to the
subfamilies of the Braconidae (Hymenoptera).
TijiUchrift voor Eittcimologic, LchIcii 119: 33-78.

Achterberg, C. van. 1982. A new genus of the Roga-
dinae-Lysitermini from Kazakhstan (Hymenop-
tera, Braconidae). Entonuilogische Beriditcn, Am-
sterdam 42: 125-128.

Achterberg, C. van. 1993. Illustrated key to the sub-
families of the Braconidae (Hymenoptera: Ich-
neumonoidea). Zcv/o^'/st'/it' Vcrlininli'liiigcn. Lciiicn
283: 1-189.

Achterberg, C. van. 1995. Generic revision of the sub-
familv Betvlobraconinae (Hymenoptera: Bracon-
idae) and other groups with modified fore tarsus.
Zoologische Verhandtiingen, Leiden 298: 1-242.

Achterberg, C. van and H. Steiner. 1996. A new genus
of Tetratermini (Hymenoptera: Braconidae: Lys-

iterminae) parasitic on grasshoppers (Gryllacri-
didae). Zoologische Mededelingen, Leiden 70: 249-
259,

Belokobylskij S. A. 1992. On the classification and
phylogeny of the braconid wasp subfamilies Do-
ryctinae and Exothecinae (Hymenoptera, Bracon-
idae). Part 1. On the classification, 1. Entonwlogi-
cheskoye Obozreniye 71: 900-928. (In Russian.)

Belokobylskij S. A. 1993. On the classification and
phylogeny of the braconid wasp subfamilies Do-
ryctinae and Exothecinae (Hymenoptera, Bracon-
idae). Part 1. On the classification, 2. Entomotogi-
chcskoye Obozreniye 72: 143-164. (In Russian.)

Belokobylskij S. A. 1995. Two new genera and two
new subgenera of the subfamilies Exothecinae
and doryctinae from the Old world (Hymenop-
tera: Braconidae). Zoologische Mededelingen, Leiden
69: 37-52.

Belshaw, R., E. Herniou, C. Gimeno, M. G. Fitton and
D. L. ]. Quicke. 1998. Molecular phylogeny of the
Ichneumonoidea (Hymenoptera) based on D2 ex-
pansion region of 28S rDNA. Systematic Entomol-
ogy 23: 109-123.

Hedqvist, K.-J. 1963. Notes on Hormiinae with de-
scription of new genera and species (Hym., Ich-
neumonoidea, Braconidae). Entomokigisk Tidskrift
84: 30-61.

Quicke, D. L. ]. and C. van Achterberg. 1990. The type
specimens of Enderlein's Braconinae (Hymenop-
tera: Braconidae) housed in Warsaw. Tijdschrift
voor Entomologie 133: 251-264.

Quicke, D. L. J., j. Tunstead, J. V. Falco, and P. M.
Marsh. 1992. Venom gland apparatus in cyclo-
stome braconid wasps with special reference to
the subfamily Doryctinae (Insecta, Hymenoptera,
Braconidae). Zoologica Scrip'ta 21: 403^16.

Tobias, V. 1. 1968. Voprosv klassifikatsii 1 filogenii
sem. Braconidae (Hymenoptera). Cliteniya Pamy-
ati N. A. Kholodkovskoga (Moscow-Leningrad)
[1967]: 3^3. (In Russian.)

Tobias V.I. 1986. Order Hymenoptera. Family Bracon-
idae. Introduction. In: Medvedev, G.S. (ed.).
Opjredelitel' nasekoniykh evropeyskoy chasti SSSR.
Pereponchatokn/lye [Keys to the insects of the Eu-
ropean part of the USSR. Hymenoptera). Lenin-
grad: Nauka. Vol 3, pt. 4. P. 7-15. (In Russian.)

Wharton, R. A. 1993. Review of the Hormiini (Hy-
menoptera: Braconidae) with a description of
new taxa. Journal of Natural History 27: 107-171.

Whitfield, J. B. and R. A. Wharton. 1997. Subfamily
Hormiinae. In R. A. Wharton, P. M. Marsh & M.
]. Sharkey (eds) Identification manual to the Neiv
World genera of Braconidae. Special Publication of the
International Society of Hymenopterists 1: 285-301.

J. HYM. RES.
Vol. 8(1), 1999, pp. 126

NOTE

New Host Record of a Ceraphronid (Hymenoptera) in
Trichoptera Pupae

J. C. LUHMAN, R. W. HOLZENTHAL, AND J. K. KjAERANDSEN

(JCL) Minnesota Department of Agriculture, Biological Control Lab,
90 W. Plato Blvd., St. Paul, MN 55107-2094; (RWH) Department of Entomology, Uiiiversity of
Minnesota, St. Paul, MN 55108; (JKK) Museum of Zoology, University of Bergen, Museplass,

3 Bergen, N-5007, Norway

Pupae of the microcaddisfly Ochrotrichia
moselyi Flint (Trichoptera: Hydroptilidae)
were collected in Costa Rica that con-
tained pupae of Aphanogmiis sp. (Hyme-
noptera: Ceraphronidae). The caddisflies
were collected in Puntarenas Province,
Bellavista River, ca. 1.5 km NW of Las Al-
turas, at 1400 m elevation (8.95rN,
82.846°W). Collections were made June
15-17, 1986. All material is housed in the
University of Minnesota Insect Collection,
St. Paul, Minnesota. Three collections in
alcohol yielded one Apbnnogmus pupa
each. There were a total of 12 Ochrotrichia
moselyi cocoons of which 3 contained
Aphanogmus pupae, 6 contained eaten cad-
disfly pupae, and the remainder, devel-
oped caddisflies. The cocoons with the
parasitoid pupae contained only the
wings and cast larval skin of the caddisfly.
There was one Aphanogmus per parasit-
ized cocoon. Dr Paul Dessart, a ceraphron-
id specialist in Belgium, confirmed Luh-
man's identifications of the Aphanogmus in
the cocoons and stated this to be a new
ordinal and family host record for Cera-
phronidae (pers. comm., 1996). Heretofore
recorded hosts of Ceraphronidae included

Diptera, Homoptera, Hymenoptera, Neu-
roptera, and Thysanoptera (Muesebeck
1979; Hanson and Cauld 1995).

The Aphanogmus were discovered inside
the cocoons of Ochrotichia moselyi, but out-
side of the caddisfly pupa. Only the de-
veloped wings remained uneaten. The
Ochrotichia cocoons may have been para-
sitized when exposed to the surface near
the water substrate interface. Ochrotichia
pupate within their larval cases in small
clusters of individuals on the sides of
rocks and boulders. During the dry sea-
son, pupae are often exposed or closer to
the surface.

ACKNOWLEDGMENTS

Trichoptera specimens were collected under NSF
grants BSR 8512368 and DEB-9400632 to R. W. Hol-
zenthal.

LITERATURE CITED

Hanson, V. E. and I. D. Gauld (eds.). 1995. The Hy-
menoptera of Costa Rica. Oxford University Press,
NY.

Muesebeck, C. F. W. 1979. Superfamily Ceraphrono-
idea, pp.1187-1195. In K. V. Krombein, P. D.
Hurd, Jr., D. R. Smith, and B. D. Burks eds., Cat-
alog of ll\iiucnof'tcra of America Nortli of Mexico,
\'ol. 2. Si/mpln/tn and Apocnta, Washington, D.C.:
Smithsonian Institution Press

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CONTENTS
(Continued from front cover)

t:

WARD, P. S. Deceptive similarity in army ants of the genus Neivamyrmex (Hymenoptera:
Formicidae): taxonomy, distribution and biology of N. californicus (Mayr) and N.
nigrescens (Cresson) 74

WHARTON, R. A. A review of the Old World genus Fopius Wharton (Hymenoptera: Bra-
conidae: Opiinae), with description of two new species reared from fruit-infesting
Tephritidae (Diptera) 48

NOTE

LUHMAN, J. C, R. W. HOLZENTHAL, and J, K. KJAERANDSEN. New host record

of a ceraphronid (Hymenoptera) in Trichoptera pupae 126

ANNOUNCEMENT

Special opportuniHes for new and current members to purchase back issues of JHR . . 127

^^OOET,^^

Journal of

Hymenoptera
Research

Volume 8, Number 2 October 1999

ISSN #1070-9428
CONTENTS

BARBALHO, S. M., A. M. PENTEADO-DIAS, and P. M. MARSH. Descriptions of new
genera from Brazil in the tribes HeterospilLni and Spathiini with similar wing
venation (Hymenoptera: Braconidae, Doryctinae) 139

BROOKS, R. W. and C. D. MICHENER. The Chilicola megalostigma species group and notes

on two lost types of Chilicola (Hymenoptera: CoUetidae, Xeromelissinae) 132

COOPERBAND, M. R, R. A. WHARTON, G. W. FRANKIE, and S. B. VINSON. New host
and distribution records for Leucospis (Hymenoptera: Leucospidae) associated
primarily with nests of Centris (Hymenoptera: Anthophoridae) in the dry forests
of Costa Rica 154

ENGEL, M. S. The taxonomy of recent and fossil honey bees (Hymenoptera: Apidae;

Apis) 165

FORTIER, J. C. and S. R. SHAW. Cladistics of the Aleiodes lineage of the subfamily

Rogadinae (Hymenoptera: Braconidae) 204

LEBLANC, L. The NearcHc species of Protarchus Foerster (Hymenoptera: Ichneumonidae:

Ctenopelmatinae) 251

LOCH, A. D. and G. H. WALTER. Does the mating system of Trissolcus basalts

(Wollaston) (Hymenoptera: Scelionidae) allow outbreeding? 238

NEWMAN, T M. and D. L. J. QUICKE. Ultrashxicture of spermatozoa in Leptopilina

(Hymenoptera: Cynipoidea: Eucoilidae) 197

SHAW, M. R. and R. R. ASKEW. Reproductive variability in Pediobius alcaeus (Walker)
(Hymenoptera: Eulophidae), a parasitoid of Phyllonorycter (Lepidoptera:
Gracillariidae) 127

BOOK REVIEW

MUELLER, U. B. — Detlef Mader. Geologische und biologische Entomookologie der rezenten

Seidenbiene CoUetes. Volume 1 268

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J. HYM. RES.
Vol. 8(2), 1999, pp. 127-131

Reproductive Variability in Pediobius alcaeiis (Walker)

(Hymenoptera: Eulophidae), a Parasitoid of Phyllonorycter

(Lepidoptera: Gracillariidae)

M. R. Shaw and R. R. Askew

(MRS) National Museums of Scotland, Chambers Street, Edinburgh EHl IJF, UK;
(RRA) 5 Beeston Hall Mews, Beeston, Tarporley, Cheshire CW6 9TZ, UK

Abstract. — Separate, but sympatric, sexual and asexual populations of Pediobius akaeus (Walker)
(Hymenoptera: Eulophidae), a parasitoid of leaf-mining Phyllonorycter species (Lepidoptera: Gra-
cillariidae), are shown to exist and to be associated with hosts on different tree species in Britain.

Pediobius akaeus (Walker) is a solitary
koinobiont endoparasitoid of many spe-
cies of Phi/lloncrycter mining the leaves of
deciduous trees and shrubs (Askew &
Shaw 1974, 1979). Eggs are laid inside
third instar or older host larvae which
continue to develop as far as the pupal
stage before being killed, and the adult P.
alcaeus emerge from the host pupae
(Askew & Shaw 1979). In Britain, Phyllon-
orycter feeding on Almis gliitinosn, Corylus
avellnna and Quercus (both petraea and rob-
ur) seem to be particularly liable to attack
from P. alcaeus, but the chalcidoid is also
frequently associated with PIn/llonon/cter
hosts on Fagus sylvatica, Malus spp. and
Carpinus betulus (Askew 1994).

Our long-standing interest in chalcidoid
parasitoids of leaf mining and other in-
sects is, in part, manifested by a records
database maintained by one of us (RRA),
into which are entered all rearing records
from our own field-work, from host col-
lections made by others who submit par-
asitoids to us for identification, and from
reared specimens we have otherwise seen
(i.e. all records entering the database are
of parasitoids identified by us). As they
currently stand, our total records continue
to indicate that Pediobius alcaeus is exclu-
sively associated with Phyllonorycter spe-
cies mining the leaves of deciduous trees

and shrubs. Our accumulated rearing re-
cords of P. alcaeus for which sex was
scored are given in Table 1: an earlier, con-
densed compilation giving fewer data was
published by Askew (1994), but the rear-
ing records of P. alcaeus previously ana-
lysed by Askew & Shaw (1974) are not all
included in Table 1 as not all of those sam-
ples were sexed. For most host species the
rearing records of P. alcaeus presented in
Table 1 come from numerous collections
made over a wide geographical spread of
localities in Britain and sometimes also
continental Europe.

As was first noted by Askew (1975),
samples of P. alcaeus originating from
hosts on different tree genera have one of
two distinct categories of sex ratio: either
males are well-represented (35-48 per
cent), or the sample is virtually entirely
unisexual with males comprising at most
only about three per cent (Table 1, see also
Askew 1994). Samples from Aluus, Carpi-
nus and Corylus are sufficiently large to
permit their categorization as host tree
genera which support unisexual (female)
populations of P. alcaeus, whilst bisexual
populations are associated with Betula, Fa-
gus, Quercus, Malus, Sorbus and probably
Acer. Different Phyllonorycter species feed-
ing on the same genus of host plants are
attacked by P. alcaeus having the same

128

Journal of Hymenoitera Research

Table 1. Accumulated total rearing records of Pediobitis akaeus (Walker) of known sex (specimens all seen
by the authors).

PhyllomrycU-r

Country

/'

alcaeii^

Host plant

.• V

i-i

Acer campestre

sylvelia (Haworth)

Britain

1

0

A. platanoides

plataiwidella (Joannis)

Britain

0

1

A. pfeiidof'latnnus

genkuklla (Ragonot)

Britain

1

0

Alnus glutinosa

froelichklla {ZeWer) / kkemanella (Fabricius)

Britain

1

62

Belgium

0

1

rajella (Linnaeus)

Britain

0

21

Netherlands

0

1

Germany

1

780

stettincnsis (Nicelli)

Britain

0

4

Germany

0

43

A. inama

strigulatellii (Zeller)

Britain

0

20

Germany

3

135

A. cordata

rajella

Britain

0

4

A. I'iridis

alpina (Frey)

Switzerland

0

1

Betula spp.

ulmifotklla (Hiibner)

Britain

13

26

Netherlands

3

4

Carpinus hetulus

qidnnata (Geoffroy)

Britain

6

219

Netherlands

0

1

tenerella (Joannis)

Britain

1

4

Belgium

0

1

Corylus avellana

coryli (Nicelli)

Britain

0

58

Belgium

0

9

Netherlands

0

2

nketlii (Stainton)

Britain

0

47

Crataegus monogyna

oxyacanthae (Frey)

Britain

0

1

Fagiis si/lvatka

maestingella (Miiller)

Britain

83

107

Netherlands

2

0

Mains domestka

corylifoliella (Hiibner)

Britain

0

1

M. sylvestris

hlancardelln (Fabricius)

Britain

0

3

cydoinella (Denis & Schiffermiiller)

Britain

7

5

Quercus robur / petraea

quercifolklla (ZeWer) / harrisella (Linnaeus)

Britain

110

155

Netherlands

0

1

lautella (Zeller)

Britain

10

21

France

2

2

Netherlands

2

0

saportella (Duponchel)

Britain

0

1

Salix caprea

salkkolella (Sircom )

Britain

0

1

species indet.

Britain

0

2

S. aurita

viminklla (Sircom)

Britain

0

1

Sorhus aucKparia

sorhi (Frey)

Britain

3

2

Netherlands

1

0

type of sex ratio; further, our limited data
indicate that the sex ratio type /host plant
relationship probably remains constant
across Europe. Careful comparison failed
to reveal any consistent morphological
differences between P. alcaeus from the
unisexual and bisexual populations.
In this paper we report the results of

sleeving experiments in which virgin fe-
male P. alcaeus from either the unisexual
or bisexual populations were presented
with developing leaf-mines on the para-
sitoids' own or on an alien host plant. The
aims of this investigation were first, to es-
tablish the type of parthenogenetic repro-
duction employed by each of the two pop-

Volume 8, Number 2, 1999

129

Table 2. Sleeves on Quercus robur and Cori/hi$ avellana into which Pln/llonon/cter species, and later virgin
Pediobiu^ alcaeus females from varying sources, were introduced. The number of moths and the number and
sex of P. alcaeuf reared in each sleeve are recorded.

Sleeve
No.

Host
tree

rili/llLilwry^lfr
species

I'clmbJUi \
no, & source

\o ot
molhs
reared

reared

1

2
3

Q. robur
Q. robur
Q. robur

quercifoliella
quercifoliella
quercifoliella

1, ex P. quercifoliella

1, ex P. coryli

1, ex Phyllonorycter sp. underside

36

40

1

46i

34 9 9

6 99

4
5
6

C. avellana
C. avellana
C. avellana

coryli
coryli
nicellii

miner on Abuts glutinosa
2, ex P. coryli
2, ex P. coryli
2, ex Phyllonorycter sp. underside

miner on Alnus glulinosa

4

14
14

29 9 9
40 9 9
23 9 9

7

C. avellana

nicellii

2, ex P. con//;

36

14 9 9

ulations, and second, to determine wheth-
er the host plant range of the unisexual
population could be expanded under ar-
tificial rearing conditions.

METHODS

Seven muslin sleeves were fastened to
branches of oak (Quercus robur) (sleeves 1-
3) and hazel {Corylus avellana) (sleeves 4—
7) in the period 2i-25.v.l979. At the same
time, or shortly afterwards, reared adult
Phi/llonorycter were introduced. The three
Quercus sleeves each received two male
and two female P. quercifoliella (an under-
surface miner), whilst six male and three
female P. coryli (an uppersurface miner)
were put into sleeves 4 and 5, and four
male and two female P. nicellii (an under-
surface miner) were introduced to sleeves
6 and 7. Variation in the numbers of
moths introduced was partly due to short-
age of livestock, but was also experimen-
tal against the risk of overstocking sleeves
(which might cause leaf abscission as the
mines developed).

Virgin female Pediobius alcaeus were put
in the sleeves during the period 22-
25. vi. 1979, at which time developing third
and fourth instar mines could be seen in
most sleeves. The Pediobius had been in-
dividually reared in tubes and came from
three sources: P. quercifoliella mines on
Quercus, P. coryli mines on Corylus and
from undersurface leaf-mines, probably of

P. froelichiella, on Alnus glutinosa. One or
two virgin Pediobius were introduced into
each sleeve; their host origins are given in
Table 2. Variation in stocking levels was
entirely due to shortage of livestock. All
host and parasitoid material came from
sites in the vicinity of Reading, Berkshire
where the experiments were performed.

The sleeves were brought indoors on
10.vii.l979 (sleeves 1-3), 13.vii.l979
(sleeves 4,5) or l.viii.l979 (sleeves 6,7). All
mature and most immature mines were
removed and stored separately in plastic
boxes for rearing adult insects. When
emergence had finished, the numbers and
sex of Pediobius and numbers of moths
produced in each sleeve were recorded.

RESULTS

Despite sometimes heavy leaf abscission
due to population explosions of enclosed
aphids, mines matured in all sleeves,
mostly in good numbers but rather sparse-
ly in sleeve 3. The first P. quercifoliella, a
leaf undersurface miner, and P. coryli,
an uppersurface miner, emerged 12-
14.vii.l979, whilst P. nicellii, another un-
dersurface miner, emerged somewhat
later about 2.viii.l979. Pediobius also com-
menced emergence about this latter date.
Total emergences of Pediobius and moths
are shown in Table 2.

130

Journal of Hymenoptera Research

DISCUSSION

Sleeve 1, the only sleeve holding P. al-
caeus from a bisexual population, yielded
only male Pediobius. This unequivocal re-
sult demonstrates that the bisexual popu-
lation of P. nicaetis on Qiiercus produces
males parthenogenetically by arrhenoto-
ky, which is usual in haplodiploid Hy-
menoptera. In contrast the other sleeves,
in which Pediobius from unisexual popu-
lations were introduced to their own or
alien hosts, all produced only female P. al-
caeus. This confirms that the populations
which appear to be unisexual from rearing
records (Table 1) are indeed thelytokous,
and not the result of inseminated females
exercising control over fertilisation of their
eggs.

The successful parasitization of P. quer-
cifoliella on Quercus, naturally a host of bi-
sexual populations of the parasitoid, by
Pediobius originating from unisexual pop-
ulations on Con/lus and Ahius (sleeves 2
and 3), shows that hosts on the parasit-
oid's 'wrong' host food-plant can support
it under artificial rearing conditions. Also,
less surprisingly, Pediobius from Alnus un-
dersurface mines was reared through un-
dersurface mines on Con/lus (sleeve 6),
and Pediobius from uppersurface leaf-
mines on Corylus was reared through un-
dersurface mines on the same tree (sleeve
7).

Populations of P. alcaeus therefore differ
in their reproductive biology. The bisexual
populations reproduce by haplodiploidy
and only males result from unfertilised
eggs (arrhenotokous parthenogenesis). Re-
production in the unisexual populations is
thelytokous with females developing from
unfertilised eggs. It is not clear whether
the occasional males appearing in rearings
of normally unisexual populations, as on
Alnus and Con/lus (Table 1), are the prog-
eny of 'stray' bisexual females, in which
case they would presumably be potential-
ly reproductively functional in relation to
the bisexual race, or whether they are

progeny of normally thelytokous females,
in which case they may be entirely repro-
ductively non-functional.

The situation in Pediobius alcaeus is sim-
ilar in some respects to that pertaining in
Diplolepis spinosissimae (Giraud) (Hym.,
Cynipidae) (Plantard et al. 1998). This rose
gallwasp is thelytokous (up to 4 per cent
males) in populations on the Atlantic coast
of France, but in two inland populations
males comprise 21 and 29 per cent of the
populations. The thelytokous populations
were found to be infected by the endo-
symbiotic bacterium Wolhachia which pre-
vents the formation of males. Bisexual
populations of D. spiiwsissitnae were free
of Wolbacbia. Whether or not a micro-or-
ganism is implicated in the thelytoky of
Pediobius alcaeus remains to be determined,
but even if this were to be the case, the
apparent segregation of sexual and thely-
tokous populations onto different tree
genera, which in Pediobius alcaeus occurs
sympatrically and (certainly in Britain)
consistently, would still require explana-
tion.

ACKNOWLEDGMENTS

The experimental part of this work was carried out
while MRS was supported by a Reading University
Research Fellowship. Eric Gissell made helpful com-
ment on an earlier version of the manuscript.

LITERATURE CITED

Askew, R. R. 1975. The organisation of chalcid-dom-
inated parasitoid communities centred upon en-
dophytic hosts, pp. 130-153. In: Price, P.W. (ed.),
Evolnlwmm/ strntcgief^ ofpnrnsitic iiifcctf ami iiiiks.
Plenum Press.

Askew, R. R. 1994. Parasitoids of leaf-mining Lepi-
doptera: what determines their host ranges? pp.
177-202. In: Hawkins, B.A. and W. Sheehan
(eds), Pimmtoid cotnmunily ecology. Oxford Uni-
versity Press.

Askew, R. R. and M. R. Shaw. 1974. An account of
the Chalcidoidea (Hymenoptera) parasitising
leaf-mining insects of deciduous trees in Britain.
Biologicnl journal of the Linncan Socich/ 6: 289-335.

Askew, R. R. and M. R. Shaw. 1979. Mortality factors
affecting the leaf-mining stages of Plii/llonon/ctfr
(Lepidoptera: Gracillariidae) on oak and hirch. 2.

Volume 8, Number 2, 1999 131

Biology of the parasite species. Zoological loiininl oky in the rose gallwasp Diphtepis s;ii»i)siss/»ii7i'

()/ the Unnean Society 67: 51-64. (Giraud) (Hymenoptera: Cynipidae), and its con-

Plantard, O., J.-Y. Rasplus, G. Mondor, I. Le Clainche, sequences on the genetic structure of its host.

and M. Solignac. 1998. Wi);;irtc/;((7-induced thelyt- Proceedings of the Royal Society B 265: 1075-1080.

J. HYM. RES.
Vol. 8(2), 1999, pp. 132-138

The Chilicola megalostigma Species Group and Notes on Two Lost
Types of Chilicola (Hymenoptera: Colletidae, Xeromelissinae)

Robert W. Brooks and Charles D. Michener

Entomology Division, University of Kansas Natural History Museum,
Lawrence, Kansas 66045, U.S.A.

Abstract, — The three species of the megalostigma group of Chilicola (Hylaeosoma) are reviewed
and a key for their separation is provided. Chilicola (H.) stenocephala Brooks and Michener,
new species, is described from Amazonian Colombia. Chilicola (H.) polita Michener is recorded for
the first time from Costa Rica. Lectotypes are designated for two Brazilian Chilicola species orig-
inally described as Oediscelis huberi Ducke and O. minima Ducke.

The xeromelissine genus Chilicola Spi-
nola has been characterized by Toro and
Moldenke (1979) and Michener (1994,
1995). It consists of small, slender, usually
black bees superficially similar to species
of Hylaeiis in the subfamily Hylaeinae, but
differing notably in the presence of a weak
scopa on the posterior femora and tibiae,
and on the first three metasomal sterna,
the hairs on the second sternum being the
best developed part of the scopa. Chilicola
is particularly abundant and diverse in
temperate South America (Chile, Argenti-
na) but ranges north to central Mexico and
to St. Vincent in the Lesser Antilles.

The subgenus Hylaeosoma, characterized
by Michener (1994, 1995), occurs from Bra-
zil to central Mexico. It consists of unusu-
ally slender species with a depression for
the reception of the antennal scape above
each antennal alveolus.

THE CHILICOLA MEGALOSTIGMA
SPECIES GROUP

Within the subgenus Hylaeosoma there is
a species-group consisting of very smooth,
shiny species with a strong, flaring pre-
occipital carina. This group, here known
as the megalostigma group, which was also
characterized as Group B by Michener
(1994:83), consists of C. megalostigma

Ducke from northeastern Brazil, C. polita
Michener from Mexico to Panama, and the
new species described below from Colom-
bia.

A hitherto unrecognized characteristic
of the megalostigma group is the modified
front tarsus of females, perhaps serving to
pull pollen from minute, deep flowers or
tubular anthers. The front tarsus of an or-
dinary species of Hylaeosoma, Chilicola
mexicana Toro and Michener, is illustrated
in Figure 1. Figures 2 to 4 show the front
tarsi of the three species of the megalostig-
ma group. All (including C. mexicana) are
bristly, with strong curved bristles as well
as straight ones, as shown in Figure 4.
This vestiture is omitted in Figures 1 to 3.
In females of the megalostigma group, the
front basitarsus is shortened if one ignores
the apical process, whereas the second
segment is relatively large, compared to
that of other species groups. More details
are indicated in the figures and in the key
to species, below.

Abbreviations used are the following: S,
sternum; T, tergum. KSEM, Entomology
Division, Snow Collections, University of
Kansas Natural History Museum,
Lawrence, Kansas, U.S.A. MPEG, Museu
Paraense Emflio Goeldi, Belem, Para, Bra-
zil.

Volume 8, Number 2, 1999

133

KEY TO THE SPECIES OF THE MECALOSTICMA GROUP OF CHILICOLA

Minimum distance between eyes about three-fourths of width of eye on same horizontal
line seen from front (Fig. 8); malar area distinct, over half as long as wide (Fig. 9). First
and second front tarsal segments of female each ending in long, down-curved, horn-like

process with blunt apex (Fig. 4) stenocephala Brooks & Michener

Minimum distance between eyes about equal to width of eye on same horizontal line seen
from front; malar area short, less than half as long as wide. First and second front tarsal

segments of female each ending in long process terminated by curved bristle 2

Frons with pit on frontal line above level of antennae, forming equilateral triangle with
the two antennal sockets; first and second front tarsal segments of female each ending in

long process terminated by curved bristle coarser than other tarsal bristles (Fig. 3)

megalostigma (Ducke)

Frons without pit on frontal line; first and second tarsal segments of female each with

apical process ending with curved bristle similar to bristles elsewhere on tarsus

polita Michener

^==^:=^^^^^S^

Figs. 1^. Outer views of anterior tarsi of females of
Chilicola (Hi/taeoscma). All are bristly with both curved
and relatively straight bristles and hairs, as shown in
Figure 4. 1, C. mexicaiui Tore and Michener; 2, C. politn
Michener; 3, C. megahsligmcj (Ducke); 4, C. itenocephii-
la n. sp. In Figure 3 the bases of only two large setae
are indicated arising from the apices of the processes
of the first two tarsal segments of C. megalostigma.
Comparable but seemingly more slender setae exist
in C. polita (Fig. 2).

Chilicola (Hylaeosoma) megalostigma
(Ducke)

Oeldiscelis] megalostigma Ducke 1908:62; Ducke

1912:83.
Oediscelis megalostigma, Nascimento 1979:7.
Clhilicola] (Hlylaeosoma]) megalostigma, Michener

1994:87.

This species is known from the Serra de
Baturite, 4°15'S, 39°05'W, altitude 700 m,
in the state of Ceara, Brazil. Although on
a mountain, this location in the xeric
northeast of Brazil is a very different hab-
itat from the moist forest areas where oth-
er species of this group are found. The
specimens were taken by Ducke on the
flowers of Borreria verticillata Mey. (Rubi-
aceae).

The lectotype, selected but not pub-
lished by J. S. Moure and C. D. Michener,
was designated (i.e., published) by Nas-
cimento (1979); it is in MPEG. Lectopara-
types are in MPEG, KSEM, and presum-
ably in the collection of Padre J. S. Moure,
Universidade Federal do Parana, Curitiba.
Specimens are also in the Departamento
de Zoologia, Universidade de Sao Paulo,
Sao Paulo, Brazil.

A front tarsus from two different fe-
males was removed and examined in glyc-
erin (and preserved in microvials on the
pins). The apical processes of the first two

134

Journal of Hymenoptera Research

tarsal segments each ends in a curved
bristle coarser than the other tarsal bris-
tles. Bases of these bristles are shown in
Figure 3. A bristle that is nearly as coarse,
however, arises from the lower distal end
of the third tarsal segment. These bristles
are brittle, easily broken off so that they
look like pegs even at a magnification of
SOX.

Chilicola (Hylaeosoma) polita Michener

Chilicoln (Hi/laeosomn) polita Michener 1994:87.

Except for the characters indicated in
the key to species, C. polita appears not to
differ from C. megalostigma. When the
front tarsus of the female is in its usual
slightly down-curved position, the process
of the basitarsus is appressed against the
under side of the second segment and is
unrecognizable, its hairs seemingly arising
from the second segment. We recognized
the process only when the tarsus was re-
laxed and bent under a dissecting micro-
scope (Fig. 2). We have not been able to
examine the front tarsi of females of C. pol-
ita at high magnification in a liquid such
as glycerin; there may be no consistent dif-
ference between C. polita and C. megalostig-
ma in this structure (see key to species).

Chilicola polita is known from Veracruz
and Oaxaca, Mexico, and Chiriqui Prov-
ince, Panama. A new record is as follows:
Costa Rica: San Jose Province, San Anto-
nio de Escazu (Eberhardt's house) [9°55'N,
84°08'W], May 27, 1996, on Ciiphea (G.
Melo, collector). [See Addendum.]

The specimen from Costa Rica as well
as the holotype from Veracruz are in
KSEM.

Chilicola {Hylaeosoma) stenocephala
Brooks and Michener, new species

In its elongate body form (Figs. 5, 6) this
species resembles other species of Hylaeo-
soma, although the length is exaggerated
by the long head which in all specimens
available is directed forward in a prog-
nathous position (Fig. 7), rather than being

hypognathous as in most bees. Other spe-
cies of the subgenus have a tendency to
the same condition, especially in species
with long heads. This is likely to be relat-
ed to pushing down into small deep flow-
ers or flower clusters. The key to species
indicates some of the principal differences
between C. stenocephala and its closest rel-
atives, C. megalocephala and polita.

Male: Body length 6 mm. Coloration:
Black, the following parts testaceous: lon-
gitudinal band on clypeus (yellowish), la-
brum, mandible, malar area (dark), ante-
rior part of hypostomal area, antennal
scape and pedicel, under side of flagellar
segments 2 to 4 (sometimes dark brown),
legs (femur, tibia and tarsus of posterior
leg infuscated). The following transparent
amber: posterior pronotal lobe, tegula, ax-
illary sclerites of wings, apical margins of
metasomal terga. Bases of T2 to T4 and
metasomal sterna except SI brown. Wings
transparent with veins and stigma black.
Body surface: Highly polished and shin-
ing, with widely separated, not sharply
defined punctures, except propodeum;
dorsal surface of propodeum with fine
longitudinal, radiating striae; sides of pro-
podeum lineolate with small punctures,
rather dull; metasomal sterna lineolate.
Hairs: Sparse, simple, erect, dark testa-
ceous to dull whitish, notably long on me-
tasoma, especially posteriorly; frons with
V-shaped pattern of hairs, lower end be-
tween antennae, arms extending to upper
part of frons (Fig. 8); pronotal lobe mar-
gined with short, appressed white hairs;
propodeum with short, appressed white
hairs that do not hide surface, except for
hairless striate dorsal surface. Structure:
Head elongate (Figs. 7, 8), proportions as
shown in illustrations, paraocular area,
with well developed depression for anten-
nal scape extending upward toward ocel-
locular area from antennal base (as in oth-
er Hylaeosoma); medial to depression a lon-
gitudinal, impunctate welt (white on left
of Fig. 8) extending from antennal base to
upper ocular tangent; first flagellar seg-

Volume 8, Numbek 2, 1999

135

Figs, 5-7. Clnluohi {Hiiliicosoiiin) stciioccpluihi n. sp. 5, male holotype; 6, temale paratype; 7, dorsal view to
show thorax, male.

ment shorter than pedicel, about 1.5 times
as long as broad, segments 2 and 3 broad-
er than long, remaining segments longer
than broad, 11 over three times as long as
broad; malar area about as long as wide
(Fig. 9); ocelloccipital distance about two
ocellar diameters measured to apex of
high preoccipital carina; genal area above
about as wide as eye seen from side, be-
low tapering to vanishing point. Maxillary
palpus about as long as prementum, near-
ly as long as head, first two segments
short, remaining four segments long,
straight, progressively slightly shortened
toward apex, so that segment 3 is longest
(also broadest). Pronotum with large dor-
sal surface (Figs. 5, 7) about on level with
scutum which is over three times as long
as mid-dorsal part of pronotum. Legs
slender (Fig. 5). Wings as shown in Figure
5; apex of marginal cell minutely truncate.
Tl longer than broad, in profile a slight

constriction between Tl and T2 (Fig. 5); T7
rather narrowly rounded at apex; poste-
rior half of S6 with strongly elevated lon-
gitudinal ridge tapering posteriorly, end-
ing in narrow, sharp, bristled apex (Figs.
10, 11). T7, T8 and genitalia as in Figures
12 to 14; genitalia with apices of gonofor-
ceps much elongated and curved mesad,
thus differing from those of the other spe-
cies of this group (see Michener 1994, Fig.
10) as well as from all other Chilicola (see
Toro and Moldenke 1979).

Female: As described for male except as
follows: Clypeus black, without testaceous
band; flagellum blackish; pronotal lobe
dark brown; legs infuscated dark brown
except trochanters testaceous; metasoma
black, dark brown ventrally. Scopa of
nearly erect whitish plumose hairs on SI
to S3, longest on S2. Scopal hairs of hind
femur plumose, shorter than femoral di-
ameter; hairs of hind tibia long and sim-

136

Journal of Hymenoptera Research

Figs. 8-14. Male of Chilicola {Hi/laeoacmii) steiwccphain n. sp. 8, face, sculpturing on left, hairs on right; 9,
side view of head; 10, S5 and S6; 11, side view of S6, ventral side to the left; 12, genitalia, dorsal on left; 13,
S7, right hand half, ventral view; 14, S8, dotted line across extreme base represents actual condition of spec-
imen; but the base has been drawn in what is presumably its undamaged shape.

pie. Flagellum short, segments broader
than long except 8 and 9 which are slight-
ly longer than broad and 10 which is
about twice as long as broad; last two seg-
ments of maxillary palpus missing, prob-
ably broken off in the single specimen; an-
terior tarsus bristly, segments 1 and 2 with
large, apical, down-curved processes lack-

ing apical hairs or pegs (Fig. 4); Tl about
as long as broad; apex of metasoma un-
modified.

Holotype male and one male and one
female paratype: Colombia: Amazonas: La
Chorrera [0°44'S, 73°0rW], 24-31 August,
1976 (M Cooper). One additional male
paratype, same data but 3 September,

VoLUMb 8, Number 2, 1999

137

1976. All bear the Natural History Muse-
um (London) number BM 1976-727, and
the specimens are in that Museum except
one male paratype in KSEM.

NOTES ON TYPES

While on the subject of tropical species
of Chilicola, it is appropriate to publish in-
formation on the types of two species that
had been lost. In 1955 Padre J. S. Moure
and C.D.M. remounted many specimens
in the Ducke collection in the Museu Par-
aense Emilio Goeldi in Belem, Para, Brazil
(MPEG). Remounting was necessary be-
cause of corroded pins that broke at the
levels of labels and inside of specimens,
sometimes bursting them. Many species
were represented by series of syntypes.
We selected and labeled lectotypes and
lectoparatypes. These were published, and
thus formally designated, by Nascimento
(1979) in his catalog of hymenopteran
types in MPEG.

That catalog makes no mention of Oed-
iscelis Iniheri Ducke and O. minima Ducke,
although types of O. megalostigma Ducke
and other species described by Ducke in
the same paper were designated. The rea-
son has only now become evident, since
the lectotypes of O. hiiberi and minima
were found by G. Melo in KSEM, where
they must have been left by C.D.M. or by
Padre Moure. They will be returned to
MPEG. The following notations and lec-
totype designations are in the style of Nas-
cimento's (1979) catalog of types.

Oediscelis huberi Ducke 1908:63.

Lectotipo. Brasil: Ceara, Serra de Batur-
ite [4°15'S, 39°05'W], 600 m, 3- VIII-
08 (Ducke). (Des. Moure e Miche-
ner, 1955).

The headless female is badly mounted
on a paper point. To judge by the wing
venation this is a member of the subgenus
Hi/laeofoma, where it was placed by Mich-
ener (1995). The abundant punctures and

unmodified front tarsus exclude it from
the megalofitigma group.

Oediscelis minima Ducke 1908:63.

Lectotipo: Brasil: Ceara, Quixada'
[04°58'S, 39°01'W], 4-VII-08 (Ducke).
(Des. Moure e Michener 1955).

The specimen consists of a female head
and anterior half of thorax, with one badly
damaged forewing, but no other wings,
badly mounted on a paper point. This spe-
cies is a member of the subgenus Proso-
poides, where it was placed by Michener
(1995).

ACKNOWLEDGMENTS

For the loan of the specimens of Chilicola stenoce-
phntn we thank Suzanne Lewis and Christine Taylor
of the Natural History Museum [London). This is
contribution number 3231 from the Division of En-
tomology, University of Kansas Natural History Mu-
seum, Lawrence, Kansas 66045, U.S.A.

LITERATURE CITED

Ducke, A. 1908. Contribution a la connaissance de la
faune hymenolopterologic]ue du nord-est du Bre-
sil, II Hymenopteres recoltes dans I'etat de Ceara
en 1908. Revue d'Entomologie 27: 57-87.

Ducke, A. 1912. Die natiirlichen Bienengenera Siida-
merikas. Zoologische lahrbucher, Abteilung fiir Sys-
tcinatik, Ceograpliie unci Biolngie lier Ticre 34: 51-
116.

Michener, C. D. 1994. Mexican and Central American
species of Chilicola. Folia Entoiuologica Mexwaiui
no. 85 (1992): 77-93.

Michener, C. D. 1995. A classification of the bees of
the subfamily Xeromelissinae. Journal of the Kan-
sas Entomological Society 68: 332-345.

Nascimento, P. T. R. 1979. Catalogo de tipos ento-
mologicos do Museu Goeldi, Hymenoptera. Bol-
etim do Museu Paraense Emilio Goeldi, Zoologia no.
96: 1-18.

Toro, H. and A. Moldenke. 1979, Revision de los Xe-
romelissinae Chilenos. Anales del Museo de His-
toria Natural Valparaiso 12: 95-182.

ADDENDUM

We have just received four specimens of Chilicola (Hy-
laeosoma) polila Midiener that add substantially to its dis-
tribution as indicated in the bod\' of this pap>er. Tlie first
two, listed below, are from the Centre for Land and
Biological Resources, Ottawa, Ontario, Canada. The oth-
er two were received through Laurence Packer, York

138

Journal of Hymenoptera Research

University, North York, Ontario, Canada. The data are
as follows (one specimen from each locality):

Guatemala: Zacapa, 3.5 km southeast of La Union,
1500 m altitude, 20-27 June 1993, in flight intercept
trap (J. Ashe, R. Brooks).

Mexico: Chiapas: Pk. Lago Belgica, 14 June 1989, in
flight intercept trap (H. Howden); Puebla: 22 km

north of Xicotepec de Juarez, 1070 m altitude, 17 June
1983 (M. Kaulbars); Tamaulipas: Rancho del Cielo
near Gomez Farias, 1000 m altitude, 7 Aug. 1983 (M.
Kaulbars).

The last locality is the northernmost record for the
genus Cliilicoln.

J. HYM. RES.
Vol. 8(2), 1999, pp. 139-153

Descriptions of New Genera from Brazil in the Tribes Heterospilini

and Spathiini With Similar Wing Venation

(Hymenoptera: Braconidae, Doryctinae)

Sandra M. Barbalho, Angelica M. Penteado-Dias and Paul M. Marsh

(SMB, AMP-D) Departamento de Ecologia e Biologia Evolutiva,

Universidade Federal de Sao Carlos, Via Washington Luiz km 235, Caixa Postal 676,

CEP 13-565-905, Sao Carlos, SP, Brazil; (PMM) P. O. Box 384, North Newton, KS 67117 USA

(Cooperating Scientist, Systematic Entomology Laboratory, USD A, Washington, DC)

Abstract. — Six new genera and 11 new species are described from Brazil. These new taxa all
have wing venation similar to that found in the genus Heterospilus, with fore wing vein 2RS absent
or weakly indicated. The new genera are placed in the tribes Heterospilini and Spathiini and a
key to the New World genera of these tribes is given. The new taxa described are as follows:
Amazondonictes Barbalho and Penteado-Dias, n.gen., A. bicolor Barbalho and Penteado-Dias,
n. sp., A. ater Barbalho and Penteado-Dias, n. sp.; Canchim Barbalho and Penteado-Dias, n.gen.,
C. carinatus Barbalho and Penteado-Dias, n. sp., C. enigosus Barbalho and Penteado-Dias, n.
sp.; Heterospathius Barbalho and Penteado-Dias, n.gen., H. belokohylskiji Barbalho and Pen-
teado-Dias, n. sp., H. petiolatus Barbalho and Penteado-Dias, n. sp., H. silvaticiis Barbalho and
Penteado-Dias, n. sp.; Jataiella pilosa Barbalho and Penteado-Dias, n.gen., n.sp.; Leptodonjctes
luizi Barbalho and Penteado-Dias, n.gen., n.sp.; Pioscelus austrimis Marsh, n. sp.; Spathiospilus
brasiliensis Marsh, n. gen., n. sp.

The subfamily Doryctinae is one of the
most diverse groups in the Braconidae, es-
pecially in the Old and Ne^N World trop-
ics. Recent studies by Marsh for the Neo-
tropics (see for instance Marsh 1993) and
Belokobyl'skij for the Old World tropics
(see for instance Belokobyl'skij 1994a, b,
1995) have shown the incredible diversity
at the generic level and have lead to re-
vised concepts of previously described
genera.

The genus Heterospilus was described by
Haliday in 1836 and characterized by the
absence of fore wing vein 2RS, thus the
first and second submarginal cells are
more or less confluent. In some cases, vein
2RS may be sclerotized but only appears
as an infuscate line with no distinct vein
edges such as a true tubular vein (see Ma-
son 1986 for definition of wing vein types
in braconids). This character was unique

for the Bracorudae and for many years
Heterospilus was the only genus known
with such wing venation. Several more
genera that were subsequently described
with this type of wing venation have been
synonymized with Heterospilus (Shenefelt
and Marsh 1976; Belokobyl'skij 1992).
Muesebeck and Walkley (1951) proposed
the genus Pioscelus for two species which
differed from the typical Heterospilus in
other morphological characters. Hedqvist
(1963) described the genus Labania which
had this same wing venation; he placed it
in the subfamily Hormiinae but stated that
it had more affinities to the Doryctinae.
The genus Heterospilus is a very large ge-
nus with about 500 species in the New
World, most of which are undescribed.
Thus, most doryctines with the fore wing
vein 2RS absent have been placed in this
genus.

140

Journal of Hymenoptera Research

Recent studies of the Neotropical fauna
of the Doryctinae have revealed several
forms with this wing venation but which
differ in many distinct characters from
typical Hetewspiliis. Until recently, these
would have all been placed near Hetewspi-
liis in the tribe Heterospilini. However,
some of these are more closely related to
the genus Spathius Nees with its petiolate
metasoma. These genera are properly
placed in the tribe Spathiini as defined by
Belokobyl'skij (1992). The genus Labania is
placed in its own tribe, Labaniini, and is
more closely related to the tribe Ecphylini.

The purpose of this paper is to provide
descriptions of several new genera from
Brazil in the tribes Heterospilini and Spa-
thiini which have this characteristic wing
venation. Keys to the New World genera
are also provided. This is part of a long
term program by the first author (SMB) to
study the Doryctinae of Brazil.

MATERIALS AND METHODS

The subfamily Doryctinae can be iden-
tified using the keys provided by Wharton
et al. 1997. The New World genera can be
identified using the key provided by
Marsh (1997) and a key to the tribes of
Doryctinae is provided by Belokobyl'skij

(1992). Morphological and wing venation
characters are based on Wharton et al.
(1997). Taxonomic authority for the new
genera and species described here is as in-
dicated for each taxon. Abbreviations for
institutions that provided specimens for
this study and where type specimens are
deposited are: DCBU, Departamento de
Ecologia e Biologia Evolutiva da Univer-
sidade Federal de Sao Carlos, Sao Carlos,
SP, Brazil; INPA, Instituto Nacional de
Pesquisas da Amazonia, Manaus, Brazil;
UFPR, Universidade Federal do Parana,
Brazil; USNM, National Museum of Nat-
ural History, Smithsonian Institution,
Washington, DC.

TRIBE HETEROSPILINI FISCHER

This tribe is distinguished from the Spa-
thiini by the following characters: meta-
soma not petiolate, the first metasomal ter-
gum being usually broad and short with
length not much longer than apical width;
the acrosternite of the first metasomal seg-
ment short, less than K length of the ter-
gum and not fusing with the lateral mar-
gins (Fig. 13); dorsope on first metasomal
tergum usually distinct and deep (see Be-
lokobyl'skij 1992).

KEY TO THE NEW WORLD GENERA OF THE TRIBE HETEROSPILINI

L Fore wing vein r-m absent (Figs. 4, 5) 2

Fore wing vein r-m present (Figs. 1-3) 3

2(1). Hind wing veins cu-a and m-cu present (Fig. 5)

Canchim Barbalho and Penleado-Dias, new genus

- Hind wing veins cu-a and m-cu absent (Fig. 4)

Leptodoryctes Barbalho and Penteado-Dias, new genus

3(1). Hind coxa with a more or less distinct antero-ventral basal tubercle or tooth 4

- Hind coxa round at base, without antero-ventral basal tubercle or tooth 5

4(3). Body densely covered with long white hair (Figs. 7-9)

Jataiella Barbalho and Penteado-Dias, new genus

- Body usually sparsely covered with short hair Heterospilus Haliday

5(3). Second metasomal tergum with two posteriorly converging grooves (Fig. 15)

Pioscelus Muesebeck and Walkley

- First and second metasomal terga with two median parallel carinae (Fig. 29)

Amazondoryctes Barbalho and Penteado-Dias, new genus

Volume 8, Number 2, 1999

141

Figs. 1-9. Figs. 1-6, wings: 1, Jataiella pihsa n. sp.; 2, Amazondon/ctes ater n. sp.; 3, Heterospathiiis pelichitus n.
sp.; 4, Leptodoryctes liiizi n. sp.; 5, Canchim cariimtus n. sp.; 6, Spathiospihis brasiliensis n. sp. Figs. 7-9, /. pdofa:
7, mesonotum; 8, propodeum; 9, metasoma.

142

Journal of Hymenoptera Research

Amazondoryctes Barbalho and
Penteado-Dias, new genus

Type species. — Amazondoryctes bicolor
Barbalho and Penteado-Dias, new species.

Diagnosis. — Face striate; vertex, malar
space and temples smooth and shining,
frons smooth or slightly striate; propleu-
ron with transverse carinae; pronotum
weakly granulate with longitudinal scro-
biculate groove; mesopleuron smooth; no-
tauli distinctly scrobiculate; sternaulus
complete; propodeum strongly areolate-
rugose; hind coxa round at base, without
basal tubercle; fore wing vein 2RS absent,
first and second submarginal cells conflu-
ent, first subdiscal cell open at apex (Fig.
2); hind wing vein M-I-CU shorter than

IM; first, second and base of third meta-
somal terga striate, remaining terga slight-
ly granulate, first and second terga with
strong complete parallel carinae (Fig. 29).

Distribution. — Amazonas State of Brazil.
One of us (PMM) has also seen an appar-
ently undescribed species from Costa
Rica.

Comments. — This genus is similar to
Pioscelus and will run to that genus in the
key provided by Marsh (1997) but is dis-
tinguished by the parallel, rather than di-
verging, carinae on the second metasomal
carina.

Etymology. — The generic name is in ref-
erence to the locality of the two known
species from Amazonia.

KEY TO SPECIES OF THE GENUS AMAZONDORYCTES

1. First and secorid metasomal terga striate, remaining terga smooth and shining; frons slight-
ly excavated A. bicolor Barbalho and Pentado-Dias, new species

- First, second and basal half of third metasomal terga striate; frons not excavated

A. ater Barbalho and Penteado-Dias, new species

Amazondoryctes bicolor Barbalho and
Penteado-Dias, new species

(Fig. 30)

Female holotype. — Head: circular; 28 an-
tennomeres, first flagellomere shorter than
scape plus pedicel; occipital carina meet-
ing hypostomal carina; oral cavity small,
diameter equal to malar space and Vs eye
height; face striate, 1.2 times longer than
eye height; vertex, frons, malar space and
temples smooth and shining; frons slightly
excavated. Mesosoma (Fig. 30): propleu-
ron with transverse carinae; pronotum
weakly granulate, with longitudinal scro-
biculate groove; mesopleuron smooth;
mesonotum angled and declivous anteri-
orly; notauli scrobiculate and meeting be-
fore scutellum in costate-rugose area; mid-
dle mesonotal lobe granulate basally, al-
veolate apically, lateral lobes granulate;
sternaulus complete, weakly scrobiculate;

metapleuron areolate; propodeum areo-
late with two longitudinal carinae lateral-
ly. Legs: fore tibia with single row of 8
spines on anterior edge; hind coxa without
basal tubercle. Wings (as in Fig. 2): fore
wing vein 2RS absent, first subdiscal cell
open at apex, vein 2cu-a absent or repre-
sented by a fuscous spot; hind wing vein
M-hCU shorter than IM, Ir-m more than
half length of IM, m-cu weak or absent.
Metasoma: first and second terga striate,
remaining terga weakly granulate; parallel
median carinae on first and second terga
complete and strong; ovipositor shorter
than metasoma. Color: body and antenna
dark brown, face yellowish, vertex dark
brown, first metasomal tergum black, re-
maining terga brown, fore and middle
legs yellow, hind coxa and femur brown,
tibia yellow basally, brown apically, ovi-
positor sheaths yellow with black at apex,

Volume 8, Number 2, 1999

143

wings slightly dusky. Body length: 3.1
mm.

Male. — Unknown.

Holotype female. — BRAZIL: Amazonas,
Manaus, ZF3, Km 23, Fazenda Esteio Res.
1112, November 26, 1986, B. Klein col. De-
posited in INPA.

Eti/mologi/. — The specific name is in ref-
erence to the bicolored body with the face
yellow and mesosoma and metasoma
dark brown or black.

Amazondoryctes ater Barbalho and
Penteado-Dias, new species

(Figs. 2, 29)

Female holotype. — Agrees with the de-
scription of A. bicolor except as follows: 38
antennomeres; face with converging
sculpture; frons not excavated, slightly
striate and with rugose sculpture between
toruli and eyes; eye height 2.4 times lon-
ger than diameter of oral cavity; first, sec-
ond and basal half of third metasomal ter-
ga striate (Fig. 29); fore tibia with 3 rows
of 22 strong spines; head and mesosoma
entirely black, first and second metasomal
terga black, remaining terga black on bas-
al half, light brown on apical half; body
length 4.6 mm.

Male. — Unknown.

Holotype female. — BRAZIL: Amazonas,
Manaus, ZF3, Km23, Fazenda Esteio Res.
1208, February 9, 1985, B. Klein col. De-
posited in INPA.

Paratypes. — BRAZIL: 1 female, Amazon-
as, Manaus, ZF3, Fazenda Esteio Res.
1112, February 9, 1985. Deposited in
DCBU.

Etymology. — The specific name is in ref-
erence to the black color.

Canchim Barbalho and Penteado-Dias,
new genus

Type species. — Canchim cnrinatus Barbal-
ho and Penteado-Dias, new species.

Diagnosis. — Vertex and frons striate or
smooth; face only slightly striate; temple
and malar space smooth; mesonotum de-
clivous anteriorly; notauli and sternaulus
complete and scrobiculate; propodeum
areolate-rugose; hind coxa rugose and
with basal tubercle; fore tibia with row of
8 spines on anterior edge; fore wing veins
r-m and 2RS both absent; hind wing vein
M-f-CU slightly longer than IM; first and
at least basal half of second metasomal
terga striate, terga beyond third smooth
and shining.

Distribution. — Sao Paulo State of Brazil.

Comments. — This genus will run to Het-
erospilus in the key provided by Marsh
(1997) but is distinguished by the absence
of fore wing vein r-m, thus all submargin-
al cells are confluent.

Etymology. — The generic name is in ref-
erence to the localities for the known spe-
cies, Fazenda Canchim, Sao Carlos, SP,
Brazil.

KEY TO SPECIES OF THE GENUS CANCHIM

1. First, second and basal 'A of third metasomal terga striate; vertex and frons striate

C. carinatus Barbalho and Penteado-Dias, new species

- First and basal Vi of second metasomal terga striate; vertex and frons often smooth

C. enigostis Barbalho and Penteado-Dias, new species

Canchim carinatus Barbalho and
Penteado-Dias, new species

(Figs. 5, 26-28)
Female. — Head (Fig. 28): occipital pre-
sent and meeting hypostomal carina; ver-

tex and frons striate; face slightly striate;
temple and malar space smooth; face
height 1.4 times longer than eye height;
eye width 2.2 times temple width; oral
cavity about equal to malar space; 17 an-

144

Journal of Hymenoptera Research

tennomeres; first flagellomere equal to
length of scape and pedicel. Mesosoma
(Fig. 26): pronotum, mesopleuron and me-
sonotum granulate; mesonotum declivous
anteriorly; notauli complete and scrobic-
ulate; sternaulus scrobiculate, not com-
plete; propodeum areolate-rugoae. Legs:
hind coxa rugose, with basal tubercle; fore
tibia with row of 8 short spines on anterior
edge. Wings (Fig. 5): fore wing veins r-m
and 2RS both absent, first subdiscal cell
open at apex, vein 2cu-a absent; hind wing
vein M + CU slightly shorter than IM. Me-
tasoma (Fig. 27): length of first metasomal
tergum equal to apical width; first, second
and basal third of third terga striate, re-
maining terga smooth and shining; ovi-
positor V4 length of metasoma. Color: head
light brown; mesosoma and metasoma
dark brown; second tergum sometimes
with triangular yellow spot at base; legs
yellow; wings hyaline, veins brown. Body
length 2 mm.

Male. — Unknown.

Holotype female. — BRAZIL: Fazenda
Canchim, Sao Carlos, Sao Paulo State, Oc-
tober 9, 1996. Deposited in DCBU.

Parah/pes. — BRAZIL: 2 females, same
data as holotype except dates of June 27,
1985 and July 26, 1989. Deposited in
DCBU.

Etymology. — The specific name is in ref-
erence to the sculpture of the head.

Canchim erugosus Barbalho and
Penteado-Dias, new species

(Fig. 25)

Female. — Agrees with the description of
C. carinatiis except as follows: head
smooth and shining, vertex slightly striate
(Fig. 25); mesopleuron slightly granulate;
first and basal % of second metasomal ter-
ga striate; ovipositor about Vi length of
metasoma; body length 1.8 mm.

Male. — Unknown.

Holotype female. — BRAZIL: Fazenda
Canchim, Sao Carlos, Sao Paulo State,
April 25, 1985, A. S. Soares col. Deposited
in DCBU.

Paratypes. — BRAZIL: 1 female, Telemaco
Borba, PR, September 1, 1986, PROFAU-
PAR (Levantamento da Fauna Entomolo-
gica do Estado do Parana). Deposited in
UFPR.

Etymology. — The specific name is in ref-
erence to the smooth head.

Jataiella Barbalho and Penteado-Dias,
new genus

Type s/'fcr'fs. — Jataiella pilosa Barbalho
and Penteado-Dias, new species.

Diagnosis. — Body densely covered with
long white hair; face, temples and vertex
smooth and shining; frons striate; prono-
tum striate laterally; propleuron and me-
sopleuron smooth; notauli deep, smooth
and meeting in a triangular area sculp-
tured (Fig. 7); sternaulus absent; hind coxa
with a small rounded tubercle at base; fore
wing vein 2RS absent except for short
stub, first subdiscal cell open (Fig. 1); hind
wing vein M + CU shorter than IM, male
with stigma in hind wing; first metasomal
tergum (Fig. 9) slightly longer than apical
width; metasomal terga 1-3 and base of 4
striate, remaining terga striate at base,
granular at apex.

Distribution. — Sao Paulo State of Brazil.

Comments. — This genus is distinguished
from other genera of Heterospilini by the
very densely hairy body, which will sep-
arate it from Heterospilus in the key pro-
vided by Marsh (1997).

Etymology. — The generic name is in ref-
erence to the locality of the type species.

Jataiella pilosa Barbalho and
Penteado-Dias, new species

(Figs. 1, 7-9)

Female holotype. — Head: densely hairy;
occipital carina present and meeting hy-
postomal carina; face and vertex smooth
and shining, frons striate; face height 1.7
times eye height; eye width 1.8 times tem-
ple width; malar space Vj oral cavity; first
flagellomere equal to length of scape and
pedicel combined. Mesosoma: densely
hairy; pronotum striate laterally; propleu-

Volume 8, Number 2, 1999

145

ron and mesopleuron smooth and shining;
middle mesonotal lobe more elevated than
lateral lobes, middle lobe with complete
median longitudinal groove (Fig. 7); no-
tauli deep, smooth, meeting before scutel-
lum in area with longitudinal carinae and
few cross carinae (Fig. 7); sternaulus ab-
sent; propodeum rugose-areolate with
two basal smooth areas (Fig. 8). Legs: hind
coxa with small tubercle at base; fore tibia
with row of 9 spines on anterior edge.
Wings (Fig. 1): fore wing vein 2RS absent
except for short stub, vein r-m present,
first subdiscal cell open at apex; hind wing
vein M+CU shorter than IM, r-m less
than half length of IM. Metasoma (Fig. 9):
first metasonal tergum 1.2 times longer
than apical width; terga 1-3 and base of 4
striate, remaining terga striate at base,
granular at apex; ovipositor about %
length of metasoma. Color: body black,
mesopleuron dark brown; wings infuscat-
ed. Body length: 5.6 mm.

Male. — Essentially as in female except as
follows: propodeum smooth; all metaso-
mal terga except apical one striate; stigma
in hind wing (Fig. 1); only lower part of
mesopleuron dark brown.

Holotype female. — BRAZIL: Rio Mogi-
Guagii, Luiz Antonio, Sao Paulo, Novem-
ber 26, 1993, L. A. Joaquim col. Deposited
in DCBU.

Paratypes. — BRAZIL: 1 male, same data
as holotype. Deposited in DCBU.

Leptodoryctes Barbalho and
Penteado-Dias, new genus

Type species. — Leptodoryctes luizi Barbal-
ho and Penteado-Dias, new species.

Diagnosis. — Head, pronotum, propleu-
ron, mesopleuron and mesonotum smooth
and shining; notauli meeting well before
scutellum, sometimes incomplete; sternau-
lus complete and scrobiculate; propodeum
rugulose; hind coxa round at base; fore
wing veins 2RS, r-m, 2M and 3M absent,
vein 2-lA also absent and, thus, first sub-
discal cell absent; hind wing veins cu-a
and m-cu absent; first metasomal tergum

weakly striate, remaining terga smooth
and shining.

Distribution. — Sao Paulo, Amazonas and
Rio de Janeiro States of Brazil.

Comments. — This genus is distinguished
from most other braconids by the wing ve-
nation with the absence of several veins in
both fore and hind wings. It can be defi-
nitely placed in the Doryctinae by the
presence of a row of stout spines along the
anterior edge of the fore tibia, presence of
a flange at the apico-lateral corner of the
propleuron, circular opening between
clypeus and mandibles and presence of
the occipital carina. The genus will not fit
well in the key to genera provided by
Marsh (1997) but would possible run to
Heterospilus because of the absence of fore
wing vein 2RS.

Etymology. — The generic name is from
the Greek leptos, meaning slender, weak,
in reference to the fragile appearance of
this genus.

Leptodoryctes luizi Barbalho and
Penteado-Dias, new species

(Figs. 4, 10-12, 31)

Female holotype. — Head (Fig. 12): com-
pletely smooth and shining; occipital ca-
rina present and meeting hypostomal ca-
rina; face height 2.3 times eye height; eye
width equal to temple width; oral cavity
diameter 1.2 times malar space. Mesosoma
(Fig. 11): pronotum, propleuron, meso-
pleuron and mesonotum smooth and
shining; notauli not complete, meeting
well before scutellum; sternaulus com-
plete and scrobiculate; propodeum ru-
gose. Legs: hind coxa round at base; fore
tibia with row of 8 spines on anterior
edge. Wings (Fig. 4): fore wing veins 2RS,
r-m, 2M and 3M absent, first subdiscal cell
absent, vein 2-1 A absent; hind wing veins
cu-a and m-cu absent. Metasoma (Fig. 10):
First metasomal tergum 3.2 times longer
than apical width, weakly striate; remain-
ing terga smooth and shining; ovipositor
as long as entire body. Color: head, me-
sosoma, legs and first metasomal terga

146

Journal of Hymenoptera Research

Figs. 10-19. Figs. 10-12, Leptodoryctes hiizi: 10, meldhoma; 11, mesosoma; 12, head. Fig. 13, Hctcnvpilui sp.,
ventral view, first metasomal segment. Fig. 14, Notiospathiiis sp., ventral view, first metasomal segment. Fig.
15, Pioscclus aiistrinus n. sp., metasoma, dorsal view. Figs. 16-19, SpMithiospihis hnisiUcnsis: 16, habitus; 17, face;
18, first metasomal segment, ventral view; 19, mesosoma and metasoma, dorsal view.

yellow, remaining terga and hind femur
light brown; wings lightly infuscated.
Body length: 1.6 mm.

Male. — Essentially as in female except as
follows: head and mesosoma dark brown,
metasoma light brown, legs yellow; fore
wing vein lA weak.

Holotyye female. — BRAZIL: Amazonas,

Manaus, Reserva Ducke, September 6,
1993, M. T. Tavares, col. Deposited in
DCBU.

Parati/pes. — BRAZIL: 2 males, Ilha
Grande, Rio de Janeiro, RJ, May 6, 1997, L.
A. Joaquim, col.; 1 male, Esta^ao Experi-
mental de Ubatuba, SP, November 15, 1990,
L. A. Joaquim, col. Deposited in DCBU.

Volume 8, Number 1, 1999

147

Etymology. — This species is named af-
ter the collector and our friend Luiz Joa-
quim.

Pioscelus Muesebeck and Walkley

Piosccliis Muesebeck and Walkley, 1951: 180.

Pioscelus austrimis Marsh, new species

(Fig. 15)

Female. — Body color: head honey yel-
low; scape, pedicel and basal 3-6 flagel-
lomeres honey yellow, remainder of fla-
gellomeres brown; mesonotum brown,
mesonotum and mesopleuron often light
brown; fore and middle legs yellow, hind
coxa except at apex and hind femur
brown, apex of hind coxa, trochanters, tib-
ia and tarsus yellow; metasoma brown,
grooves on second metasomal tergum
sometimes lighter; wings hyaline, veins
including stigma light brown. Body size:
4.0 mm. Head: antenna with at least 19 an-
tennomeres (broken in all females of type
series), all flagellomeres at least 5 times
longer than wide, first flagellomere very
slightly shorter than second; vertex and
temple smooth and shining; frons exca-
vated, mostly smooth and shining but
with a few striations medially; face rugu-
lose and dull; eyes large, malar space
about Vi eye height; ocelli small, ocell-oc-
ular distance about twice diameter of lat-
eral ocellus; occipital carina complete,
reaching hypostomal carina. Mesosoma:
pronotum coriaceous with median scro-
biculate groove; mesonotum coriaceous,
median lobe sharply declivous anteriorly
with lateral corners broadly produced, no-
tauli scrobiculate, meeting before scutel-
lum in narrow longitudinally rugose-car-
inate area; mesopleuron smooth medially,
subalar area broadly scrobiculate, sternau-
lus smooth, as long as mesopleuron; pro-
podeum slightly longer than first metaso-
mal tergum, not declivous apically, ru-
gose-areolate laterally, rugose dorsally
with basal lateral areas indistinct, coria-
ceous. Wings: fore wing vein 2RS weak or
absent, at most indicated by weak infus-

cated line apically, vein 3RSa twice as long
a vein r, first subdiscal cell open at apex,
vein 2cu-a absent; hind wing vein cu-a
present, vein M + CU about Vi length of
IM. Legs: hind coxa angled at base but
without a distinct antero-ventral basal tu-
bercle; hind femur short and swollen,
about three times longer than width. Me-
tasoma (Fig. 15): first tergum with length
about twice as long as apical width,
strongly longitudinally carinate; second
tergum with two converging grooves en-
closing a basal semicircular carinate area,
tergum carinate laterally; third tergum
separated from second by transverse ar-
cuate groove, carinate on basal %, coria-
ceous apically; remainder of terga coria-
ceous basally, smooth apically; ovipositor
about Vi length of metasoma.

Male. — Essentially as in female; hind fe-
mur more greatly swollen, about twice as
long as wide; hind wing without stigma.

Holotype female. — BRAZIL: Manaus,
ZF3, Km23, Faz. Esteio, Res., 1208, B. Klein
col., November 5, 1985. Deposited in
INPA.

Paratypes. — BRAZIL: 2 females, same
data as holotype, February 27, 1985, No-
vember 18, 1987; 1 female, Cerrado, Can-
chim, Sao Carlos, S.P., December 4, 1989,
L. A. Joaquim, col.; 1 male, Mata Canchim,
Sao Carlos, S.P., April 26, 1996, L. A. Joa-
quim, col. Desposited in DCBU.

Etymology. — The specific name is from
the Latin anstriiuis meaning southern in
reference to this species being the first one
recorded from South America.

TRIBE SPATHIINI FOERSTER

This tribe is distinguished from the
Heterospilini by the following charac-
ters: metasoma petiolate, first tergum
usually long and narrow, acrosternite
very long, at least Vi length but usually
nearly as long as the tergum, fused with
lateral margins (Figs. 14, 18); dorsope on
first tergum weak or absent (see Belo-
kobyl'skij 1992).

148 Journal of Hymenoptera Research

KEY TO THE NEW WORLD GENERA OF THE TRIBE SPATHIINI

1. Fore wing vein 2RS absent or not sclerotized (Figs. 3, 6) 2

Fore wing vein 2RS present and sclerotized 3

2(1). Eyes large, malar space very short or absent (Fig. 17); fore wing vein RS+M arched (Fig.

6); hind coxa with small but distinct antero-ventral basal tubercle

Spathiospilus Marsh, new genus

- Eyes smaller, malar space at least % eye height; fore wing vein RS + M not arched (Fig.

3); hind coxa without basal tubercle (Fig. 21)

Heterospathius Barbalho and Penteado-Dias, new genus

3(1). Hind coxa round at base, without basal tubercle 4

- Hind coxa with distinct basal tubercle or tooth at base 5

4(3). Hind wing vein m-cu curved toward wing apex, hind wing vein M+CU % length of

vein IM Psetwbolus Reinhard

- Hind wing vein m-cu curved toward wing base, hind wing vein M + CU Va length of
vein IM Notiospathins Matthews and Marsh

5(3). First metasomal tergum without triangle area at base Spathiiis Nees

- First metasomal tergum. with distinct triang.ular area .at. base 6

6(5). Fore wing vein m-cu asising distad of 2RS Ptesimogaster Marsh

- Fore wing vein m-cu arising basad or directly in line with 2RS

Trigonophasmus Enderlein

Heterospathius Barbalho and long and slender, length at least 4 time

Penteado-Dias, new genus apical width.

rr ,, , ... .■ , . Distribution. — Amazonas, Para, Ron-

Tuve spectes. — Heterosvatnius vetwlatus ,,. ,^ „,^ ^t, ■, .<

D u iu J r> i. J A- ■ donia and Sao Paulo States of Brazil. Also,

Barbalho and Penteado-Uias, new species. ^ , , ,

D.agnosis.-Diameter of oral cavity °"^ °^ "^ (^^M) ^^^ ^'^^^ ^^^^""^l ""'^^^-

about equal to malar space; occipital ca- ^"^^^ ^P^"^^ ^'■°'" *=°^t^ ^'^^' ^° *^ '^•^-

rina meeting hypostomal carina; 25-35 an- tnbuHon of this genus is probably over

tennomeres; mesonotum declivous anteri- Central and South America,

orly; notauli complete, scrobiculate; ster- Comments.— This genus is similar to No-

naulus complete, scrobiculate; propodeum tiospathius but is disHnguished by the ab-

horizontal for basal Va, usually with two sence of vein 2RS in the fore wing; it will

lateral longitudinal carinae; hind coxa ru" ^o Heterospihis in the key provided by

without basal hjbercle; fore wing vein 2RS Marsh (1997) but is separated by the shape

absent, first subdiscal cell open at apex, of the petiolate metasomal segment,

vein 2cu-a absent, hind wing vein M + CU Eti/mology.—The generic name is in ref-

shorter than IM, m-cu absent, r-m less erence to the fore wing similarity to Het-

than V2 length of IM, no stigma in hind erospilus and the first metasomal segment

wing of male; first metasomal segment similarity to Notiospnthiiis.

KEY TO SPECIES OF THE GENUS HETEROSPATHIUS

1. Vertex and face strongly striate-rugose; ovipositor about 1.5 times longer than body

(Fig. 24) H. belokobylskiji Barbalho and Penteado-Dias, new species

Vertex (Fig. 22) and face finely to weakly striate; ovipositor equal to or shorter than
body 2

2(1). Ovipositor about V2 length of metasoma

H. petiolatus Barbalho and Penteado-Dias, new species

Volume 8, Number 2, 1999

149

Ovipositor about equal to body length

H. silvaticiis Barbalho and Penteado-Dias, new species

Heterospathiiis petiolatus Barbalho and
Penteado-Dias, new species

(Figs. 3, 20-23)

Female holotype. — Head (Fig. 22): occipi-
tal carina meeting hypostomal carina; face
and vertex striate; frons smooth; temple
smooth and shining; face height 2.1 times
eye height; face width 1.6-2.1 times eye
width; diameter oral cavity about Vj eye
height; eye width 1.4-1.7 times temple
width; malar space about equal to diam-
eter of oral cavity; 25 antennomeres, first
flagellomere longer than scape and pedi-
cel combined. Mesosoma (Fig. 20): prono-
tum rugulose-granulate; mesonotum de-
clivous anteriorly; mesonotum rugulose-
granulate; notauli scrobiculate, meeting at
scutellum in triangular rugose area; me-
sopleuron granulate; sternaulus complete,
scrobiculate; propodeum areolate-rugose,
with distinct longitudinal carinae. Legs:
fore tibia with single row of 7 spines on
anterior edge; hind coxa (Fig. 21) rugose,
without basal tubercle; first tarsomere of
hind tarsus 2.3 times longer than second,
second equal to length of third and fourth
combined, fifth twice as long as fourth.
Wings (Fig. 3): fore wing vein 2RS absent
or represented by short stub, first subdis-
cal cell open at apex, vein 2cu-a absent;
hind wing vein M-l-CU much shorter than
IM, r-m less than half length of IM. Me-
tasoma (Fig. 23): first tergum rugose, slen-
der, length 5.7 times apical width, apical
width equal to basal width; remaining ter-
ga smooth and shining; ovipositor about
Vz length of metasoma. Color: head, me-
sosoma and first metasomal tergum dark
yellow, remaining terga dark to light yel-
low; ovipositor light yellow with apex
black; apical 6-12 flagellomeres white, re-
mainder brown; wings slightly dusly,
veins brown, stigma brown with white at
basal third. Body length: 3.3 mm.

Variation in female. — Face occasionally
weakly striate or granular, width 1.6-2.1
times eye width; eye width 1.4-1.7 times
temple width; 20-32 antennomeres; fore
tibia with row of 7-13 spines; metasoma
with apical terga occasionally black; body
length 1.5-3.7 mm.

Male. — Essentially as in female except as
follows: 20-25 antennomeres, apical 2-6
flagellomeres white; diameter of oral cav-
ity slightly greater than malar space; face
width 1.5-1.9 times eye width; frons
smooth or granulate; fore tibia with row
of 7-13 spines; metasoma completely light
yellow or with apical Va dark brown; no
stigma in hind wing.

Holotype female. — BRAZIL: Amazonas,
Manaus, ZF3, Fazenda Esteio, November,
1984. Deposited in INPA.

Parati/pes. — BRAZIL: 2 females, 1 male,
Rondonia, Ariquemes, Rio Ji, October 28,
1986, R. A. Rafael, col; 1 female, Fazenda
Canchim, Sao Carlos, Sao Paulo state,
April 30, 1987, L. A. Joaquim, col; 1 fe-
male, Rio Tocantins, Tucurui, Para state,
November, 22, 1989, N. Degullier, col.; 1
female, Santarem-Cucuruna, Para state,
February, 1996, A. R. Lisboa col.; 2 fe-
males, Amazonas, Manaus, ZF3, Faz. Es-
teio, Res. 1301, January 22, 1986; January
29, 1986, B. Klein col.; 1 female, 1 male
Amazonas, Manaus, ZF3, Faz. Esteio, Res.
1208, October 22, 1986, B. Klein col.; 1 fe-
male, Amazonas, Manaus, km 60, PDBFF/
WWF, Res. 1210, November 8, 1984, B.
Klein col; 1 female, Rio Tocantins, Tucu-
rui, Para state; 2 females, Rio Branco, Acre
state, October 25-November 8, 1991, F.
Ramos, A. Henriques, I. Gorayeb & N. Bit-
tencourt cols.; 1 male, Amazonas, Manaus,
ZF3, Faz. Esteio, Res. 1113, January 30,
1986, B. Klein col.; 1 male, Manaus, ZF3,
Faz. Esteio, Res. 1113, January 23, 1986, B.
Klein col.; 2 males, Manaus, ZF3, Faz. Es-

150

Journal of Hymenoptera Research

Figs. 20-31. Fig. 20-23, Hctcrospntliius pctiolntus n. sp.: 20, propodeum; 21, hind coxae; 22, vertex of head; 23,
metasoma. Fig. 24, Hcterospatlui{~^ Ivtckoln/hklji n. sp., metasoma. Fig. 25, Cancltiiii criigcaui^ n. sp., vertex of
head. Figs. 26-28, C. cnriiititus n. sp.: 26, mesopleuron; 27, metasoma; 28, vertex of head. Fig. 29, Ainazomti'ri/ctcf
ntcr n. sp., metasoma. Fig. 30, A. bicvhr n. sp., mesopleuron. Fig. 31, Lcptodorydcti liiizt. habitus.

Volume 8, Number 2, 1999

151

teio, Res. 1208, November 20, 1984; Octo-
ber 17, 1984, B. Klein col.; Concei^ao do
Araguaia, Para state, January 19-31, 1983,
R. Nonato col. Deposited in DCBU, INPA,
USNM.

Etymology. — The specific name is in ref-
erence to the petiolate metasoma.

Heterospathitis belokobylskiji Barbalho
and Penteado-Dias, new species

(Fig. 24)

Female. — Agrees with the description of
petiolatiis except as follows: 35-38 anten-
nomeres, apical 12-13 flagellomeres white;
face and vertex strongly striate-rugose,
vertex occasionally strongly striate; frons
striate; temples smooth; face height 1.6
times eye height; fore and mid granulate;
propodeum without strong longitudinal
carina; ovipositor 1.5 times longer than
body (Fig. 24); body entirely brown; body
length 5.5 mm.

Male. — Unknown.

Holoti/pe female. — BRAZIL: Amazonas,
Manaus, ZF3, Km32, Fazenda Esteio, Jan-
uary, 1986, B. Klein col. Deposited in
INPA.

Paratypes. — BRAZIL: 1 female, same
data as holotype; 1 female. Para, Santa-
rem-Cucuruna, February, 1996, A. Pentea-
do-Dias, col. Deposited in INPA, DCBU.

Etymology. — This species is named in
honor of our colleague and fellow re-
searcher on the Doryctinae, Sergey Belo-
kobyl'skij. Zoological Institute, Russian
Academy of Sciences, St. Petersburg, Rus-
sia.

Heterospathius silvaticus Barbalho and
Penteado-Dias, new species

Female holotype. — Agrees with descrip-
tion of petiolatiis except as follows: 27 an-
tennomeres, apical 8 flagellomeres white;
face striate; vertex only slightly striate;
frons, temple and malar space smooth and
shining; face width 2.2 times eye width;
face height 1.7 times eye height; diameter
of oral cavity slightly greater than malar
space; no longitudinal carinae on propo-

deum; fore tibia with row of 7 spines on
anterior adge; fore and middle coxa weak-
ly striate; ovipositor about equal to body
length; body length 2.8 mm.

Male. — Agrees with female except as
follows: entire body light brown, propo-
deum dark brown.

Holotype female. — BRAZIL: Amazonas,
Manaus, ZF3, Km23, Fazenda Esteio, No-
vember 8, 1984, B. Klein col. Deposited in
INPA.

Paratypes. — BRAZIL: 1 male, Amazonas,
Sao Gabriel da Cachoeira, April 29, 1982,
J. A. Arias, col. Deposited in DCBU.

Etymology. — The specific name is from
the Latin silvaticus meaning belonging to
woods or trees in reference to the collec-
tion of this species in the jungle.

Spathiospilus Marsh, new genus

Type-species. — Spathiospilus brasilieiisis
Marsh, new species.

Diagnosis. — Cyclostome braconid, oral
cavity circular, labrum concave; eyes
large, malar space very small or absent;
flagellomeres with double row of placodes
separated by ridge around middle of fla-
gellomere; mesonotum strongly declivous
anteriorly; fore wing vein 2RS absent or
weakly present apically near vein r, vein
RS + M strongly arched, hind wing of male
with stigma; for tibia with row of short
stout spines along outer edge, hind coxa
with small but distinct antero-ventral bas-
al tubercle; metasoma petiolate, first ter-
gum slender, parallel sided, rest of meta-
soma suddenly widened, acrosternum
nearly as long as tergum and fused with
tergum.

Distribution. — Sao Paulo State of Brazil.
We have also seen one undescribed spe-
cies from Panama.

Comments. — This genus is similar to Het-
erospathius in the Spathini by the absence
of fore wing vein 2RS but distinguished
by the strongly arched fore wing vein
RS + M and the large eyes. In the key pro-
vided by Marsh (1997) it will run to Het-
erospnlus but is distinguished by the long

152

Journal of Hymenoptera Research

and fused acrostemum of the first meta-
somal segment.

Etymolog}/. — The generic name refers to
the similarity to Hetewspihts by the wing
venation and to Spathius by the petiolate
metasoma.

Spathiospilus brasiliensis Marsh,
new species

(Figs. 6, 16-19)

Female. — Body color: head, mesosoma
and metasoma reddish-brown, metasomal
terga 2-5 sometimes infused with black;
scape and pedicle yellow, flagellum yel-
low basally turning brown to apex; legs
yellow; wings hyaline, veins yellow at
base and apex, stigma and veins across
middle of wing brown; ovipositor sheaths
yellow, black at tip. Body size: 3-4 mm.
Head (Fig. 17): 29-31 antennomeres, fla-
gellomeres with two rows of placodes sep-
arated by ridge around middle of each fla-
gellomere; eyes large, covering most of
head; malar space extremely short or ab-
sent, lower margin of eye nearly touching
base of mandible; face narrow, width less
than length from clypeus to antennal sock-
ets; oral cavity circular, diameter slightly
greater than basal width of mandible; tem-
ple very narrow, about K, eye width; ocell-
ocular distance shorter than diameter of
lateral ocellus; face, frons and vertex ru-
gulose-coriaceous, temple coriaceous; oc-
cipital carina scrobiculate along vertex
and temple side. Mesosoma (Fig. 19):
pronotum with scrobiculate grove across
dorsal surface extending laterally on each
side, bordered laterally by strong carina,
dorsally coriaceous; mesonotum strongly
declivous anteriorly, mesonotal lobes co-
riaceous, notauli scrobiculate, median lobe
with short and wide carinate-rugulose
area before scutellum and with median
raised line extending to pronotum; scutel-
lum coriaceous, bordered laterally by ca-
rina, scutellar sulcus deep with 5 cross ca-
rinae; mesopleuron coriaceous, subalar
area rugose, sternaulus scrobiculate; pro-
podeum strongly areolate-rugose apically

and laterally, with semicular coriaceous
areas basolaterally bordered by distinct
carinae. Wings (Fig. 6): fore wing vein r V2
length of 3RSa, vein RS-(-M strongly
arched; hind wing vein RS absent, vein m-
cu curved toward wing apex. Metasoma
(Fig. 19): first tergum petiolate, parallel
sided with apical and basal widths equal,
longitudinally costate, weakly rugulose
between costae; second tergum longitudi-
nally costate, weakly rugulose between
costae, triangular shaped with basal width
abiut V2 apical width, weak line between
second and third tergum; third tergum
longitudinally costate on basal %, coira-
ceous on apical Vs; remainder of terga co-
riaceous; ovipositor about X; length of me-
tasoma.

Male. — Essentially as in female; 26 an-
tennomeres; stigma present in hind wing.

Holotype Female. — BRAZIL: Rio Mogi
Gua^u, Luis Antonio, S.P., February 18,
1988, L. A. Joaquim collector. Deposited in
DCBU.

Paratypes. — BRAZIL: 1 female, 3 males,
same data as holotype with additional
dates of March 20-27, 1987; 1 female, Luis
Antonio, S.P., Reserva Ecol. do Jatai, Feb-
ruary 8, 1994, A. S. Scares collector; 1 fe-
male, Faz. Canchim, Sao Carlos, S.P., June
20, 1985, A. S. Soares collector. Deposited
in USNM, DCBU.

Etymology. — The specific names is in ref-
erence to the localities of the types series
from Brazil.

ACKNOWLEDGMENTS

We thank the following for the loan of specimens
for this study: Dra. Keti M. Rocha Zanol and Dr. Ren-
ato Contin Marinoni of the Universidade Federal do
Parana (UFPR), the Institute Nacional de Pesquisas
da Amazonia (INPA), and the National Museum of
Natural History, Washington, DC (USNM). Financial
support was provided by the Conselho Nacional de
Desenvolvimento Cientifico e Tecnologico (CNPq)
and the Fundaijao de Amparo a Pesquisa do Estado
de Sao Paulo (FAPESP). Permits to collect specimens
in the area of the Canchim Farm were provided by
Centro de Pesquisa de Pecuaria do Sudeste (EMBRA-
PA, CPPSE, Sao Carlos, SP).

Volume 8, Number 2, 1999

153

LITERATURE CITED

Belokobyl'skij, S. A. 1992. On the classification and
phylogeny of the braconid wasp subfamilies Do-
ryctinae and Exothecinae (Hymenoptera, Bracon-
idae). Part 1. On the classification, 1. Eiitoinolo-
gkiwskof Obi-izrenie 71:900-928. In Russian. (En-
glish version published in 1993, Entomological Rc-
vieu' 77:109-137).

Belokobyl'skij, S. A. 1994a. A review of parasitic
wasps of the subfamilies Doryctinae and Exothe-
cinae (Hymenoptera, Braconidae) of the Far East.
Hymenoptera Insects of Siheria and Far East. No. 3,
pp. 5-77.

Belokobyl'skij, S. A. 1994b. A new tribe of the sub-
family Doryctinae from Papua New Guinea (Hy-
menoptera: Braconidae). Zoosi/stematica Rossica
3(1):141-145.

Belokobyl'skij, S. A. 1995. Two new genera and two
new subgenera of the subfamilies Exothecinae
and Doryctinae from the Old World (Hvmenop-
tera: Braconidae). Zoologische MededcUngen 69(3):
37-52.

Haliday, A. H. 1836. Essav on parasitic Hymenoptera.
Entomologists Magazine 4:38-59.

Hedqvist, K.-J. 1963. Notes on Hormiinae with de-
scriptions of new genera and species (Hym., Ich-
neumonoidea, Braconidae). Entomologisk Tidskrift
84:30-61.

Marsh, P. M. 1993. Descriptions of new Western
Hemisphere genera of the subfamily Doryctinae
(Hymenoptera: Braconidae). Contributions of the
American Entomological Institute 28(l):l-58.

Marsh, P. M. 1997. Subfamily Doryctinae, pp. 206-
233. In: R. A. Wharton, P. M. Marsh and M. J.
Sharkey (eds.), Manual of the New World Gen-
era of the Family Braconidae (Hymenoptera).
Special Publication of the International Society ofHy-
menopterists No. 1, 439 pp.

Mason, W. R. M. 1986. Standard drawing conventions
and definitions for venational and other features
of wings of Hymenoptera. Proceedings of the En-
tomological Society of Washington 88:1-7.

Muesebeck, C. F. W. and L. M. Walkley. 1951. Family
Braconidae (pp. 90-184). In: C. F. W. Muesebeck
et al., Hymenoptera of America North of Mexico.
Synoptic Catalog. U. S. Department of Agriculture,
Agricultural monograph 2, 1420 pp.

Shenefelt, R. D. and P. M. Marsh. 1976. Braconidae 9,
Doryctinae. In: J. van der Vecht and R. D. She-
nefelt, eds., Hymenopteroruni Catalogus (new edi-
tion), part 13, pp. 1263-1424. Dr. W. Junk, The
Hague.

Wharton, R. A., P. M. Marsh and M. J. Sharkey. 1997.
Manual of the New World genera of the family
Braconidae (Hymenoptera). Special Publication of
tlie International Society of Hymenopterists Number
1, 439 pp.

J. HYM. RES.
Vol. 8(2), 1999, pp. 154-164

New Host and Distribution Records for Leucospis (Hymenoptera:

Leucospidae) Associated Primarily with Nests of Centris
(Hymenoptera: Anthophoridae) in the Dry Forests of Costa Rica

Miriam F. Cooperband, Robert A. Wharton, Gordon W. Frankie, and

S. Bradleigh Vinson

(MFC, RAW, SBV) Department of Entomology, Texas A&M University,

College Station, TX 77843-2475;

(GWF) Department of Environmental Science, Policy, and Management,

University of California, Berkeley, CA 94720

Abstract. — A study conducted in the dry forests of Lomas Barbudal Biological Reserve, Guan-
acaste Province, Costa Rica, revealed new host records and elevation data for five species of
Leucospis: nztecn, hulbiventris, cnyennensis, egaia, and latifrons. Four species of trap nesting Centris
bees (Hymenoptera: Anthophoridae) were attacked, as well as at least 4 species of trap nesting
bees in the family Megachilidae. Of 295 Leucospis wasps reared from these trap nesting bees, it
was possible to associate 236 with hosts, 189 of which were from nests of Centris bicornuta. Ele-
vational data and host species data are summarized for each Leucospis species reared, as well as
sex ratios of reared material. Additional notes on biology and behavior are provided, along with
a key to the five species reared from trap-nesting Centris bees in Costa Rica. The taxonomic status
of L. hulbiventris, a sexually dimorphic species, is discussed relative to L. manaica.

Members of the family Leucospidae are
among the largest species of Chalcidoidea,
and all are parasitoids of solitary and sub-
social aculeate bees and wasps. They drill
through the hardened cell walls of their
hosts' nests with their unusual oviposi-
tors, and develop as ectoparasitoids. Ex-
cellent summaries of leucospid biology
are provided by Clausen (1940), Habu
(1962), and Boucek (1974). The family was
revised on a world basis by Boucek (1974),
who also summarized the known host
data. Hosts were recorded for 32 of the
130 species of Leucospidae recognized by
Boucek (1974). Prior to Boucek's (1974) re-
vision, no species of leucospids had been
recorded as parasitoids of Centris bees,
though other anthophorids (notably spe-
cies of Xylocopa) were known as hosts of
at least three species of Leucospis. Subse-

'MFC has relocated to the Department of Entomol-
ogy, University of California, Riverside, CA 92521.

quently. Chandler et al. (1985) reared two
individuals of L. cayennensis Westwood
from Centris in Minas Gerais, Brazil. Leu-
cospids frequently parasitize megachilid
bees in California (GWF, personal obser-
vations), and there are several published
records of species of Leucospis attacking
various megachilids (Boucek 1974, Burgis
1995).

Centris is a large genus of bees in the
hymenopteran family Anthophoridae,
with at least 32 species occurring in Costa
Rica (Snelling, 1984). Their nesting habits
are diverse. Some species make their nests
in the ground while others utilize holes
left in wood by other insects; some pro-
vision multiple cells per nest, others pro-
vision only one cell per nest. Aside from
collecting pollen, all of them collect oil
from plants that have oil producing flow-
ers, and this oil is used in their nesting
biology (Vinson et al. 1996).

Centris bees, because of their large size.

Volume 8, Number 2, 1999

155

are a readily observed component of the
Mesoamerican dry forest habitat during
the dry season. Trap-nests are very effec-
tive means of studying the activity of
wood-cavity nesting species (Frankie et nl.
1988, 1993, Vinson et al. 1996). In the area
of this study, six species of Ceniris have
been found to nest in tree holes, however
only four of these are commonly found in
the trap nests (Frankie et al. 1988). Centris
biconiuta Mocsary is one of the most abun-
dant of the trap nesting species in this area
(Frankie et al. 1988, 1993, 1997). A number
of inquilines and parasitoids can be reared
from trap-nests, and in the Guanacaste
Province of Costa Rica, leucospids were
one of the dominant parasitoids of some
of the species of Centris being studied
there. Below we record the first specific
host associations for Leucospidae on Cen-
tris in Costa Rica.

MATERIALS AND METHODS

Stud}/ sites. — The study was conducted
at the Lomas Barbudal Biological Reserve
in the dry forest in Guanacaste Province,
Costa Rica. Two bundles, each containing
six block monitoring units (BMUs) (Fran-
kie et al. 1993), were placed in dry forest
sites at 100 m, 300 m, 600 m, and 800 m
elevations in a transect extending from
Hacienda Monteverde (at 100 m, 8 km
NW Bagaces) northwards towards Volcan
Rincon de la Vieja. All BMUs were hung
at eye level on tree trunks in shaded lo-
cations, and were monitored at 14 day in-
tervals throughout the dry season (late
December to May). The 100 m site had a
mixture of oak forest and riparian ever-
green forest (Frankie et al. 1988), and had
been disturbed by agricultural develop-
ment. The 300-800 m sites were largely in-
tact oak forest with several other scattered
tree species. The oak species, Quercus oleo-
ides Schlecht. & Cham., was the same at
all four sites.

Trap nests. — The nesting activities of
several species of Centris were monitored
using BMUs. These BMUs consisted of 12

small wooden sticks (11.5 X 2.3 X 2 cm)
with holes drilled in one end, bundled to-
gether to form a block. The wooden sticks
(consisting of pine or two local hard wood
species) were drilled lengthwise to a
depth of between 7.0 to 11.0 cm depend-
ing on the hole sizes (diameters of 4.5, 6.5,
8, 9.5 and 11 mm were used). Two sticks
representing each of the hole diameters,
along with two additional sticks with a
hole diameter of 8 mm, were bundled to-
gether using fine wire or twine. The 8 mm
hole size was doubled as it is the hole size
most commonly used by Centris bees
(Frankie et al. 1988, 1993). SHcks were lay-
ered within the block so that a drilled end
of a particular hole size always alternated
with a non-drilled end, and small to large
hole sizes descended from the top to bot-
tom of a block.

Emergence. — Each BMU was numbered
and identified as to location, altitude, and
time and date of placement. Every 14 days
the sticks with completed bee nests were
replaced with a new stick of the same hole
diameter and additional information was
recorded on the removed stick, including
date removed. Sticks with completed nests
were placed in large wire baskets (—20 x
20 X 40 cm) made of 2.5 cm open mesh
hardware cloth that prevented access by
mammals, but not parasitoids. These bas-
kets were hung from nails on nearby trees.
During the wet season, glass scintillation
vials were taped to the nest entrances to
intercept any emergences. Emergences
were monitored on a two week basis and
parasitoid and host associations recorded.
Data presented here were collected during
four consecutive years (December 1993 to
December 1997).

Behavior and development. — All observa-
tions on female wasp behavior were made
at 100 m sites where 2-4 BMUs were con-
tinuously being monitored. Completed
nests, 1 to 14 days old, were removed
from BMUs, placed in baskets at these
sites, and observed daily during daylight
hours for 3 weeks. Each basket contained

156

Journal of Hymenoptera Research

12

0)

ro

o
ro

O

X

5

Chalicodoma

Unknown spp

^^g

D L- azteca
B L- bulbivenths
?ZL. cayennensis
S/.. ega/a
■ L. latifrons

sv\\v.\\\\\<.v?^?a

^^3

'//////A

^

0% 25% 50% 75% 100%

Percent of Leucospis species

Fig. 1. Percent of each Leucospis species found on
different hosts. Total for each Leucospis species across
all hosts adds to 100%.

approximately 100 sticks arranged in three
rows of two sticks deep so that each stick
had at least one surface exposed. The
number of leucospids searching or at-
tempting to oviposit on nests was record-
ed until the experiment was terminated.
One stick from each basket was removed
daily during week 3 of the experiment,
and opened to monitor bee development,
and that of any present parasites, parasit-
oids, and diseases. This experiment was
replicated 4 times.

Additional observations were made on
16 ovipositing female leucospids where,
following completion of oviposition and
departure of the leucospid, the exact dril-
ling location was marked. The marked
stick was then removed and dissected to
determine the stage of the host attacked
and the placement of the leucospid egg.
Observations on oviposition behavior
were also made on nests removed from
BMUs and placed in baskets at the 100 m
site.

Specimen repositories. — Voucher speci-
mens for the Ceritris and megachilid bees
are at University of California, Berkeley,
and those for the Leucospis species are at

UC Berkeley and Texas A&M University
(TAMU). Material for comparison, includ-
ing primary types, was borrowed from the
Philadelphia Academy of Natural Sciences
(ANSP), the Natural History Museum,
London (BMNH), and TAMU.

RESULTS AND DISCUSSION

Emergence data. — We reared 295 leucos-
pids, representing five species, from at
least seven species of wood-hole nesting
bees in Guanacaste Province. Of 236 host
bees, 189 were Centris bicornuta which was
attacked by all five species of Leucospis: L.
azteca Cresson, L. bulbiventris Cresson, L.
cayennensis Westwood, L. egaia Walker,
and L. latifrons Schletterer (Fig. 1). Centris
nitida F. Smith and C. vittata Lepeletier
each hosted four different species of Leu-
cospis (Fig. 1). The most abundant leucos-
pid in our samples was L. bulbiventris (56%
of reared individuals) (Fig. 2). Discounting
L. egaia, represented only by three reared
individuals in our samples, all leucospids
were more abundant at lower elevations
(Fig. 3). Leucospis azteca and L. cayeniwnsis
were most abundant at the 100 m site; L.
bulbiventris and L. latifrons were most
abundant at the 300 m site (Fig. 3). In ad-
dition to C. bicornuta, C. nitida, and C. vit-
tata, we also reared leucospids from C. an-
alis F., undetermined species of Chalicodo-
ma and Anthidium (Hymenoptera: Me-
gachilidae), and at least two other
undetermined species of megachilid bees,
all from the trap nests described above.

The experimental environment em-
ployed in this study was artificial owing
to the fact that the Ct'nfns-infested trap
nests were highly accessible to leucospids
by being clustered in baskets. The nests
were thus at much higher densities than
would be encountered in nature. In Costa
Rican dry forests, Centris nests tend to be
more widely dispersed, and the bees are
not limited to nesting in holes in small
sticks, which are maximally exposed for
Leucospis oviposition. Therefore, the large
number of leucospids recorded here is at

Volume 8, Number 2, 1999

157

Frequency of Each Leucospis Species Reared

azteca

21%

bulbiventris

56%

latifrons
11% cayennensis

11%

Fig. 2. Relative proportions of Leucospis species reared from trap nesting bees.

least in part a reflection of the experimen-
tal manipulations.

Observations on oviposition. — No female
leucospids were observed around the wire
baskets or on the sticks until nests were at
least 3 weeks old (Fig. 4). Peak activity oc-
curred towards the end of the third week,
but since the experiment was terminated
before all activity ceased, we lack data on
how long nests remain attractive. Female
leucospids fly up wind to the wire enclo-
sures (100% N = 19), and ultimately land
on one of the sticks containing bee nests.
Generally, they walk the length of the
stick slowly while alternately drumming
the surface with their antennae. Prior to
drilling, they stop to antennate the wood
surface with both antennae held close to-
gether, then move forward half a body
length to drill the antennated spot with
the ovipositor.

Based on dissections of the 16 marked

cells into which leucospids were ob-
served ovipositing, female leucospids
only attacked cells in which the mature
larvae had at least begun to spin a co-
coon. Leucospid eggs were always locat-
ed inside the cocoons, either on the sur-
face of a host larvae still finishing its co-
coon (N = 1), a prepupa (N = 3), or a
pupa (N = 12). Of the 18 other leucospid
larvae or pupae recorded from randomly
dissected nests, all were within the co-
coon of a bee, indicating that either the
bee is allowed to develop to a prepupa
and spin a cocoon, or they are only par-
asitized following cocoon formation. The
failure to find leucospid larvae on youn-
ger stages of bee larvae, and the delay in
leucospid response to newly provisioned
bee nests, suggests that these leucospids,
at least, do not attack earlier stages of
their hosts. Parasitized host larvae did
not move, but it was not clear if they

158

Journal of Hymenoptera Research

100%

75%

50%

25%

Proportions of Four Species of
Leucospis Found at Different Elevations

n L azteca
UL. bulbiventris
0/.. cayennensis
■ L. latifrons

_^^|_,

100

300

600

800

Elevation (m)

Fig. 3. Proportions of four species of Leucofpiii found at different elevations. Total for each Leucospis species
across all elevations adds to 100"Xi.

Average Number of Leucospis Attacks on Bee Nests

7
6
5

4 -
3 -
2
1
0

3

day 1

day 2

day 3

WEEK 1 WEEK 2 1-4-

day 4

WEEKS

day 0

dJV '-■'

-♦I

day 1
WEEK 4

Fig. 4. Be
at the begi
of week 4

havioral observations. Number of Leucosfus attacks on provisioned sticks observed over time. Nests
nning of week 1 were from 1 to 14 days old. Observations were not extended beyond tlie first day

Volume 8, Number 2, 1999

159

were paralyzed since larvae at this stage
of development are lethargic.

There have been few prior studies in
which more than one species of leucos-
pid has been reared from a single host
species. In addition to the five specific
examples listed by Boucek (1974), RAW
(unpublished) has collected both L. his-
trio Maindron and L. niolei/rei Maindron
from nests of a single species of Xi/locopm
in Papua New Guinea. The data pre-
sented here represent the first record, to
our knowledge, of five leucospid species
reared from a single host species. Fur-
ther, we know of only one species of
Leucospidae previously associated with
Centris bees (Fritz and Genise 1980, De
Santis 1983, Chandler et al. 1985).

Parasitoid identifications. — Preliminary
identifications of the Leucospis species
were greatly facilitated by the excellent
detail provided in the monograph by
Boucek (1974). In our attempt to confirm
the identifications of these species, how-
ever, we uncovered several problems as-

sociated with the primary types. The ho-
lotypes of bulbiveutris and dubiosa Cres-
son and lectotype of azteca should be in
ANSP. They were all examined by Bou-
cek, and the types returned as indicated
by correspondence at ANSP. Despite
considerable effort by D. Azuma, how-
ever, no leucospid primary types could
be found, though several others should
also be at ANSP.

The key presented here to the leucos-
pids attacking Centris bees in dry forests
of Costa Rica is adapted largely from this
work, as is the terminology. The key has
been greatly simplified for ease in identi-
fication of leucospids attacking Centris in
dry forests of Costa Rica, and should only
be used in this context (or for comparison
with Centris parasitoids from dry forests
elsewhere). There are several other species
of Leucospis known from Mesoamerica,
and Boucek's work should therefore be
consulted for any species not reared from
Centris, and for rigorous confirmation of
suspect individuals.

KEY TO SPECIES OF LEUCOSPIS ASSOCIATED WITH CENTRIS BEES IN COSTA RICAN DRY

FOREST HABITATS

I- Pronotum with a transverse, premarginal cross carina and /or narrow yellow stripe near
posterior margin. Mandible with triang^ular indentation on mesal chewing edge (Fig. 5) . . 2

- Pronotum without transverse, premarginal cross carina or narrow yellow stripe. Mandible
with semicircular indentation on mesal, chewing edge (Fig. 6) . . . L. cayennensis Westwood

2. Scutellum at least partly yellow 3

- Scutellum completely without yellow coloration L. bulbiveutris Cresson

3. Hind femur with fewer than 10 small teeth on ventral margin. Hind coxa with a slender
tooth (spine) on its dorsal posterior edge (as in Figs. 8, 9) 4

- Hind femur with more than 10 small teeth on ventral margin. Hind coxa with a thin,
partially translucent lobe on dorsal-posterior edge (as in Fig. 7), never with a spine-like
tooth L. egaia Walker

4. Setae on hind coxa converge towards center of depression (Fig. 8). Ovipositorial furrow
on first metasomal tergum in the form of a simple, smooth, convex ridge down the midline
(Fig. 11). Yellow coloration on scutellum restricted to posterior half, at least anterior half

of scutellum black L. latifrons Schletterer

- Setae on hind coxa nearly all pointing in the same direction, not converging towards the
center (Fig. 9). Ovipositorial furrow on first metasomal tergum with a smooth, shiny, con-
vex ridge down the midline, and concave slopes on either side of the ridge (Fig. 12). Nearly
entire scutellum yellow or yellowish, with only anterior edge black L. azteca Cresson

160

Journal of Hymenoptera Research

Figs. 5-9. Mandibles and hind legs of Leucospis species. 5. Mandible of L. latifrons, female. 6. Mandible of L.
cayennensis, female. 7. Hind leg of L. bulbiventris, female. 8. Hind leg of L. latifrons, female. 9. Hind leg of L
azteca, female.

Leucospis azteca Cresson

(Figs. 9, 12)

Leucospis azteca, previously known only
from the three specimens of the original
type series, lacks obvious diagnostic fea-
tures. It can be identified by the following
combination of characters: mandible with
triangular indentation (Fig. 5); pronotum
with premarginal carina (a weak, trans-
verse ridge within a yellow band that runs
parallel and slightly anterior to the pos-
terior margin of the pronotum), median
lobe of metanotum (= dorsellum) coarsely
sculptured, but without distinctly carinate
lateral margins; hind tibia apically trun-
cate; hind coxa regularly punctate and se-
tose (as in Fig. 9); and ovipositorial furrow

as in Fig. 12. Boucek (1974) noted that L.
azteca shared several features in common
with both L. latifrons and L. affinis Say, and
that the species had been variously con-
fused in previous studies. Leucospis latif-
rons is readily identified by the pattern of
dense pubescence on the hind coxa (com-
pare Figs. 8 and 9), but one of the subspe-
cies of L. affinis treated by Boucek, L. a.
dubiosa Cresson, is particularly problem-
atic as it is very similar to L. azteca. The
apparent loss of the holotype of L. dubiosa
and lectotype of L. azteca makes it even
more difficult to separate L. affinis from L.
azteca. Fortunately, the two other members
of the original type series of L. azteca were
located in ANSP. Structurally, both of

Volume 8, Number 2, 1999

161

Figs. 10-12. Ovipositorial furrows on the first nietasomal terga of Leucospis species. 10. L. I'ulbhvntiis, female.
11. L, latifrons, female. 12. L. azteca, female.

these agree very closely with our material,
but there are some differences in color.
This is particularly noticeable on the scu-
tellum. In our material, the yellow color-
ation covers at least the posterior half of
the scutellum, providing a readily observ-
able field characteristic for separating this
species from the otherwise similar L. latif-
rons. In both of the L. azteca paratypes,
however, the scutellum is much less ex-
tensively yellow, and similar to our latif-
rons in this regard. Given this variation in
color, we concur with Boucek (1974) that
the differences in the ovipositorial furrow
on the first metasomal tergum are impor-
tant for distinguishing L. affinis from L. az-
teca, and it is on this basis that we have
determined our material as L. azteca. In L.
azteca, the furrow is generally not as deep
as in affinis (a feature that is difficult to
assess without side by side comparison),
and scattered setae occur on the polished
median ridge. In the specimens of L. affinis
available to us from California and south-
em Texas, the median, polished ridge
lacks setae (though these are abundant
along the edge of the furrow). We have
found that density of punctation on the

hind leg (a feature used by Boucek 1974)
is too variable to be used for separation of
L. affinis, L. azteca, and L. latifrons unless
side by side comparison is possible with a
good series of specimens representing all
three species (which we were fortunate to
have at our disposal).

The biology of this species was previ-
ously unknown. Our data suggest that it
is a generalist, capable of attacking several
different species. Of the 58 individuals for
which we had host data, 41 came from C.
bicornuta, 1 from C. nitida, 3 from C. vittata,
4 from Anthidium, 5 from Chalicodoma, and
4 from other undetermined species of Me-
gachilidae. Unlike L. latifrons, L. azteca was
confined to lower elevations. Of the 53
specimens for which we had altitudinal
data, 98%, were reared from nests at 100
m, and only 2% from 300 m (Fig. 3). Of
the 63 individuals we reared, only 17%
were male, showing a strong female bias
(Fig. 13). Leucospis affinis has been reared
from a wide variety of megachilid bees,
and could conceivably be found on the
same hosts as L. azteca where their ranges
overlap (e.g. in Mexico). As these species

162

Journal of Hymenoptera Research

are very difficult to separate, caution must
be exercised when identifying them.

Leucospis bulbwentris Cresson

(Figs. 7, 10)

Leucospis bulbiveiitris is readily identifi-
able by the complete absence of yellow
coloration on the scutellum, the shape of
the ovipositorial furrow, which is excep-
tionally broad anteriorly (Fig. 10), the
deep, triangular incision of the mandible
(much deeper than in Fig. 5), and the
shape of the hind coxa, which has a thin,
partially translucent lobe on its dorsal
posterior edge (Fig. 7). It is a large but
slender species with a distinctly petiolate
abdomen.

Leucospis bulbiveiitris was previously
known only from the male holotype col-
lected in Mexico. A second nominal spe-
cies, L. maimica Roman, described from
Brazil, has heretofore been known only
from five females that are similar in many
respects to the holotype of L. bulbiventris
(Boucek 1974). We reared a good series of
males and females, in several cases from
the same host nest, enabling us to associ-
ate the sexes with certainty. Boucek (1974)
was the first to suggest that perhaps L. bul-
biveiitris and L. maimica represent different
sexes of the same species. We confirm that
the differences in setation and overall
shape between L. maiiaica and L. bulbiveii-
tris noted by Boucek represent sexual di-
morphism. Based on our rearings, we
therefore strongly suspect that L. iiiaiiaica
and L. bulbiventris are the same; but be-
cause we do not have males from South
America, and, more importantly, cannot
locate the holotype of L. bulbiventris, we
must unfortunately leave this problem un-
resolved. The name bulbiventris has prior-
ity over manaica, and since males from our
material fit the description of L. bulbiven-
tris provided by Boucek (1974), we have
therefore used this name for our species.

No biological information has previous-
ly been published for either L. bulbiventris
or L. manaica, nor have either of these been

recorded before from Costa Rica. Of the

116 individuals for which we have host
data, the majority (108) came from nests
of C. bicornuta. The others were reared
from C. aiialis (1), C. nitida (1), and C. vit-
tata (6). Of 150 reared individuals for
which we have elevation data, 21% were
from 100 m, 73% from 300 m, and only 7%
from nests at 600 m. None were found at
800 m (Fig. 3). Of 165 individuals reared,
only 19% were males, showing a strong
female bias (Fig. 13).

Leucospis cayennensis Westwood

(Fig. 6)

This is a widespread Neotropical spe-
cies recorded from Mexico to Argentina as
well as the Caribbean (Fidalgo 1980, De
Santis 1983). It is readily identified by the
semicircular indentation of the mandible
(Fig. 6), the complete lack of a transverse
premarginal carina on the pronotum, and
the relatively smooth hind coxa (with dor-
sal two-thirds of the depression smooth,
shiny, bare and impunctate).

Fritz and Genise (1980) were the first to
record Centris tarsata Smith as a host of L.
cayennensis, and this is the only specific
host recorded to date (Chandler et al. 1985,
De Santis 1983). In Fritz and Genise's
(1980) study, L. cayennensis was reared
from 14% of the C. tarsata cells in old,
abandoned Sceliphron asiaticum (L.) nests.
Our data show that L. cayennensis attacks
at least four other species of Centris bees.
Of the 30 individuals for which we have
host data, 22 came from nests of C. bicor-
nuta, 4 from C. nitida, 3 from C. analis, and
1 from C. vittata. Leucospis cayennensis was
found at all four elevations sampled dur-
ing this study, but seemed to have a pref-
erence for lower elevations. Approximate-
ly 61% of the individuals for which we
have elevational data were reared from
nests at 100 m, 21% from 300 m, 14% from
600 m, and 4% from 800 m (Fig. 3). Of 33
individuals collected, one third were
males, showing a female sex bias (Fig. 13).

Volume 8, Number 2, 1999

163

100%

75%

50%

25% -

0%

Sex Ratios for Reared Leucospis Species (N)

83%

81%

Dmale 90%

D female

17%

19%

67%

33%

10%

azteca (63) bulbiventris (165) cayennensis (33)

Fig. 13. Sex ratios of Lcucospiis species reared from trap-nesting hosts.

latifrons (31)

Leucospis egaia Walker

This species is similar in general ap-
pearance and coloration to L. aztecn and L.
latifrons, but has a carinately margined
dorsellum, a more rounded, translucent
lobe rather than a spinose tooth on the
hind coxa, and an oblique rather than
truncate margin to the hind tibia. No host
records existed previously for L. egaia
(Boucek 1974). We reared a male from C.
biconinta at 100 m, a male from a megachi-
lid bee at 600 m, and a female from a me-
gachilid bee at 600 m. Since only 3 indi-
viduals of this species were found, we
cannot make general inferences about its
biology.

Leucospis latifrons Schletterer

(Figs. 5, 8, 11)

This is another widespread Neotropical
species, occurring from Mexico to Bolivia.
It is readily identified by the arrangement
of the dense patch of setae in the central
depression of the hind coxa (Fig. 8). The
color pattern on the scutellum was suffi-
ciently stable in our material to use for

separation of L. azteca from L. latifrons in
the field. As noted above, however, these
color patterns may vary from one locality
to the next, and should be used cautiously
for identification purposes. Hosts were
previously unknown for L. latifrons. Of the
29 individuals for which we have rearing
data, 17 came from C. hicornitta, 6 from
megachilid bees, 4 from C. nitida, I from
C. vittata, and 1 from Chalicodoma. Of the
32 individuals with altitudinal data, 6%
came from nests at 100 m, 52% came from
nests at 300 m, 19% from nests at 600 m,
and 23% from nests at 800 m (Fig. 3). This
species seems to be more of a generalist,
able to adapt to a variety of hosts and el-
evations, with an apparent preference for
habitats at 300 m. Although L. latifrojis and
L. azteca are extremely similar morpholog-
ically, the elevation data suggest a biolog-
ical difference supporting Boucek's (1974)
finding that they are two distinct species.
Of 31 individuals reared, only 10% were
male, showing a strong female bias, as in
all the other species in our samples (Fig.
13).

164

Journal of Hymenoptera Research

ACKNOWLEDGMENTS

We are most grateful to D. Azuma, Z. Boucek, and
E. Grissell for their assistance in our attempts to lo-
cate the type material. The Friends of Lomas Barbu-
dal offered logistic support for this research. The Cal-
ifornia Agricultural Experiment Station provided fi-
nancial support for most of the field work. We also
thank J. Oswald who offered the use of his scanner
and computer during the final phase of preparing the
illustrations, and two reviewers (J. Noyes and J.
LaSalle) for suggesting improvements in the text.

LITERATURE CITED

Boucek, Z. 1974. A revision of the Leucospidae (Hy-
menoptera: Chalcidoidea) of the world. BiiUctm
of the British Miisctini (Natural History) Entomolo-
gy, Supplement 23:1-241.

Burgis, H. 1995. Leucospis gigas (Chalcidoidea: Leu-
cospidae) as parasite of the mortal bee Megachilc
sicula (Apoidea: Megachilidae). Bembix 5:27-32.

Chandler, L., J. A. F. Barrigossi, and E. B. S. Diaz.
1985. The first definitive host record for Leucospis
cai/ennensis Westwood (Hymenoptera: Leucospi-
dae). Revistn Ceres 32:170-174.

Clausen, C. P. 1940. Entomophagous Insects. McGraw-
Hill, New York. 688 pp.

De Santis, L. 1981. Catalogo de los Himenopteros Cal-
cidoideos de America al Sur de los Estados Un-
idos — Primer Suplemento. Revista Peruana ilc En-
tomologia 24:1-38.

Fidalgo, A. P. 1980. Nuevas citas de calcidoideos para

Argentina, Bolivia y Peru (Hymenoptera). Neo-
tropica 26:193-196.

Frankie, G. W., L. E. Newstrom, and S. B. Vinson.
1993. Nesting-habitat preferences of selected Cen-
tris bee species in Costa Rican dry forest. Biotro-
pica 25:322-333.

Frankie, G. W., S. B. Vinson, L. E. Newstrom, and J.
F. Barthell. 1988. Nest site and habitat preferenc-
es of Centris bees in the Costa Rican dry forest.
Biotropica 20:301-310.

Frankie, G. W., S. V. Vinson, M. A. Rizzardi, T. L.
Griswold, 5. O'Keefe, and R. R. Snelling. 1997.
Diversity and abundance of bees visiting a mass
flowering tree species in disturbed seasonal dry
forest, Costa Rica, journal of the Kansas Entomo-
logical Societ}/ 70:281-296.

Fritz, M. A. and J. A. Genise. 1980. Notas sobre nido
de barro de Sphecidae (Hymenoptera) construc-
tores, inquilinos, parasitoides, cleptoparasitos y
detritivoros. Revista de la Sociedad Entomohigica
Argentina 39:67-81.

Habu, A. 1962. Chalcididae and Leucospidae and Po-
dagrionidae (Insecta: Hymenoptera). Fauna ]a-
ponica 1-232.

Snelling, R. R. 1984. Studies on the taxonomy and
distribution of American centridine bees (Hy-
menoptera: Anfhophoridae). Contributions in Sci-
ence (Los Angeles). 347:1-69.

Vinson, S. B., G. W. Frankie, and H. J. Williams. 1996.
Chemical ecology of bees of the genus Ccutris
(Hymenoptera: Apidae). Florida Entonu^logist 79:
109-129.

]. HYM. RES.
Vol. 8(2), 1999, pp. 165-196

The Taxonomy of Recent and Fossil Honey Bees
(Hymenoptera: Apidae; Apis)

Michael S. Engel

Department of Entomology, American Museum of Natural History,
Central Park West at 79th Street, New York, New York 10024-5192, USA

Abstract. — An attempt is made to clarify the complicated and error fraught taxonomic history
of the honey bees (genus Apis Linnaeus) by cataloguing the numerous names proposed for Apis.
The 178 species- and 10 genus-group names (including those proposed herein) are brought for
the first time into accord with the modern classification of the honey bees. The results of this
synthesis reveal a number of names in current usage to be taxonomically confused. The authorship
of the species Apis koschevnikovi is established as Enderlein and not Buttel-Reepen. The correct
names for four subspecies of the Western honey bee, A. mtilifera, are established as A. mellifcra
remipes Gerstacker (not A. mellifera anuenincn Skorikov), A. mellifera sicilianci Grassi (not A. mellifera
Simla Montagano), A. mellifera jemenitica Ruttner (not A. mellifera yemenitica Ruttner), and A. mel-
lifera Caucasia Pollmann (not A. mellifera caiicasica Gorbachev). The correct authorship of the sub-
species A. mellifcra iberica is established as Ruttner and not Goetze, of A. mellifera syriaca as Sko-
rikov and not Buttel-Reepen, and of A. mellifera intermissa as Maa and not Buttel-Reepen. Three
replacement names are proposed for preoccupied subspecific taxa: A. mellifera artemisia Engel
(new name for A. mellifera acervorum Skorikov, nee Linnaeus), A. mellifera iberiensis Engel (new
name for the aforementioned A. mellifera iberica Ruttner, nee Skorikov), and A. mellifera sossimai
Engel (new name for A. cerifera Gerstacker, nee Scopoli). The previously confused Himalayan
honey bee (infrequently associated with A. cerana skorikovi Maa, nomen nudum or A. cerana "Him-
alaya race") is proposed and validated as A. cerana skorikovi Engel (new subspecies). Similarly
the large black race of China (Aba race) is proposed as A. cerana heimifeng Engel (new subspecies).
A lectotype and 11 paralectotypes are designated for A. mellifera ruttneri (by Sheppard et «/.), a
lectotype is designated for A. koschevnikovi, and a neotype is designated for A. mellifera iberica
Ruttner. Apis cuenoti Theobald is newly synonymized under A. henshaun Cockerell, A. mellifera
rodopica Petrov is synonymized with A. mellifera macedonica Ruttner, while Hauffapis scheeri Arm-
bruster, H. scharmanni Armbruster, H. scheeri gallauni Armbruster, H. scheeri rahdei Armbruster, H.
scheuthlei seemanni Armbruster, and H. scheuthlei zeuneri Armbruster are all synonymized under
A. armbrusteri Zeuner (new synonymies). The name A. vetustus Engel for a fossil from the Oli-
gocene of Europe is emended to A. vetiista Engel. All genus-group taxa are characterized on the
basis of adult and larval morphology and ethology. Two new subgenera are proposed to accom-
modate fossil honey bee species: Cascapis Engel new subgenus (type: A. armbrusteri Zeuner) and
Priorapis Engel new subgenus (type: A. vetusta Engel).

The honey bees (genus Apis Linnaeus) pict early humans collecting honey combs,

are the most famous of all insects owing Similarly, the honey bees figure promi-

to their import for the pollination of crops, nently in human cultures, mythologies,

their social organization, and the honey and religions (e.g., see Ransome 1937).

they produce. There has been a long as- Numerous world revisions have been at-

sociation between Apis and man as is at- tempted for the genus (Gerstacker 1862,

tested by the Paleolithic Arafia Cave 1863, Smith 1865, Ashmead 1904, Buttel-

paintings near Valencia, Spain which de- Reepen 1906, Enderlein 1906, Skorikov

166

Journal of Hymenoptera Research

Table 1. Classificatory structure of major honey bee revisions since Gerstacker (1862); living species only.
Subspecies and varieties omitted so as to save space. Species are in alphabetical order and do not necessarily
correspond in rows from one column to the next. 1 have replaced uses of wcWficn by some of these authors
with mellifera. Although Maa (1953) observes that Skorikov (1929b) recognized 14 species, the later author left
one unnamed (the "Egyptian bee") and 1 have therefore listed only 13 here. G = genus, sg = subgenus.

Gerstacker
1862

Smith
1865

Ashmt'.id

1')II4

Butlcl-Reepen

Endt'rlfin

SkonkiH'
W24b

\Ki,i

I'resent author

G. Aph

G. Apis

G. Apis

G. Apis

G. Apis

G. Apis

G. Apis

G. Apis

dorsata

adansoiiii

cerana

dorsata

dorsata

sg. Apis

sg. Apis

sg. Apis

florea

dorsata

iitdica

florea

florea

adansoiiii

adansonii

cerana

indica

florea

mellifera

mellifera

indica

cerana

intennissa

koschevnikovi

mellifera

indiea

nigritarum

mellifera

cypria

lamarckii

mellifera

mellifera

nigrocincta

indica

meda

nigrocincta

nigrocincta

unicolor

japonica

mellifera

sg. Megapis

sinensis

G. Megapis

johni

remipes

dorsata

zonata

dorsata

meda

unicolor

sg. Micrapis

zonata

mellifera

sg. Sigtnatapis

andreniformis

G. Micrapis

remipes

cerana

florea

florea

syriaca

indica

1871

unicolor

javana

added:

sg. Megapis

johni

laboriosa

dorsata

sg. Micrapis

florea

koschcviiikovi

licftiiicki

nigrocincta

peroni

philippina

samarensis

vecliti

G. Megapis

bingliami

breviligula

dorsata

laboriosa

G. Micrapis

andreniformis

florea

1929b, Maa 1953) as well as an unpub-
lished faunal revision from Thailand (Ma-
laipan 1972). Since the first detailed study
by Gerstacker (1862), anywhere from 3 to
24 species have been recognized while the
four subgenera have at times been treated
as distinct genera (Table 1). Despite the
consistent and concentrated effort on Apis
taxonomy for well over a century, a clear
picture of the species and the numerous
names associated with each has yet to be
achieved. It is hoped that this paper (al-
though mostly an exercise in "bookkeep-
ing") will make a small step towards
achieving these goals.

The primary objective of this paper has

been to pull together the extensive litera-
ture pertaining to the taxonomy of honey
bees and to associate the long lists of ju-
nior synonyms with their valid counter-
parts. The last catalogue of Apis names
was undertaken by Maa (1953); however,
his list of names contains a number of er-
rors and his rather extreme classification
is not easily reconcilable with current us-
age. Moreover, Maa did not treat the nu-
merous fossil honey bees and at least 60
names have either been discovered or
were proposed since his study (approxi-
mately one-third of the total number of
names). Below I have compiled a listing of
all names applied to honey bees, both liv-

Volume 8, Number 2, 1999

167

ing and extinct. This compilation contains
178 species- and 10 genus-group names.
Although Maa (op. cit.) and Ruttner (1988)
noted that over 600 species-group names
have been proposed for Apis, this does not
mean that 600 names have been proposed
for honey bees. Many of these names were
proposed under a Linnean concept of the
genus which included all bees. Thus, most
of the 600-1- names pertain to species now
placed in other genera and families (e.g.,
Anthopliora, Bonihiis, Megachile, Trigoiin,
&c.). Interestingly most of the names ap-
plicable to honey bees have been pro-
posed during this century. A breakdown
of the list reveals eight names appearing
between 1758 and 1799, 17 from 1800
through 1849, 26 between 1850 and 1899,
66 in the period 1900 to 1949, and 61
names between the years 1950 and 1999.
Of these names 146 are applicable to the
living species (90 associated with Apis itiel-
lifera alone!), 25 for the fossil species, and
seven names are of dubious taxonomic
status.

Included with this catalogue I have pro-
vided descriptions of all genus-group taxa
now included in Apis, thereby adding to
this work a subgeneric revision of the ge-
nus. Table 2 summarizes the hierarchical
classification of Apis as it is proposed
herein.

Phylogenetic studies on the genus have
recently been undertaken by Alexander
(1991 a,b), Engel (1998a), and Engel and
Schultz (1997: see also works cited there-
in). The current hypothesis of relation-
ships among the subgenera and species is
depicted in Figure 1 (modified from Engel
1998a). General works on honey bee anat-
omy, biology, behavioral ecology, and di-
versity have been presented by Snodgrass
(1956), Winston (1987), Seeley (1985, 1995),
and Smith (1991a) respectively. The distri-
bution of the various species and subspe-
cies has been thoroughly treated by Rutt-
ner (1988, 1992), Otis (1996), and Hepburn
and Radloff (1998). The position of the Ap-
ini among the other tribes of corbiculate

Table 2. Current hierarchical classification of the
honey bees; tribe Apini Latreille (excluding infraspe-
cific taxa). Daggers (t) indicate extinct taxa.

Genus APIS Linnaeus
subgenus Apiii Linnaeus

cerana Fabricius

koschevnikovi Enderlein

meltifera Linnaeus

nigrocincta Smith
t subgenus Cnscapiis Engel

t armhrusteri Zeuner
subgenus Megnpis Ashmoad

dorsata Fabricius
subgenus Micrapns Ashmoad

andreniformis Smith

florea Fabricius
+ subgenus Priorapis Engel

t vetusta Engel
t subgenus Symipis Cockerell

t ht'nslui-d'i Cockerell

t longtibia Zhang

t mioccnkci Hong

+ pctrcfactn (Rilia)

bees has been considered most recently by
Chavarria and Carpenter (1994), Engel
(1998b, unpubl. data), Noll (1998, in
prep.), and Schultz et al. (1999).

FORMAT

Standard formats for taxonomic histo-
ries are used. Comments provided for
each taxon give information on the assign-
ment of authorship to certain names, sub-
specific classifications, studies of the rec-
ognition of difficult taxa, and occasionally
historical information on the biogeogra-
phy or biology of the species. Several hon-
ey bee names were used in publications as
iiomina iiudn and in those cases where no
description was later provided they are in-
dicated as such. However, whenever a
name was later made available by an as-
sociated description I have only listed the
publication making the name available
under the rules of the I.C.Z.N. (1985) and
not the original paper in which it ap-
peared as a nomen nudum.

For those species in which subspecies
are currently recognized (A. cerana, A. dor-

168

Journal of Hymenoptera Research

£

vetusta
henshawi
petrefacta
longtibia
miocenica
armbrusteri
florea

andreniformis
dorsata
- mellifera
nigrocincta
cerana
koschevnikovi

I Priorapis

Synapis

Cascapis
Micrapis
Megapis

Apis

Fig. 1. Phylogeny of the honey bees (genus A\m Linnaeus) with subgenera indicated (modified from Engel
1998a).

sata, and A. ineUifera) I have associated
each junior synonym with its correspond-
ing subspecies. Names in brackets at the
end of each entry indicate which subspe-
cies the name corresponds to. In some cas-
es (e.g., nomina iiiida) this decision cannot
always be made with certainty and so are
accordingly noted with an interrogative
mark.

Descriptions are provided for genus-
group taxa and are based on information
from all adult castes, the mature larva (of
workers), and ethological information.
Too little information exists at this time on
Apis pupae to know whether additional
character data can be gleaned from this
life-stage. Characters given for workers
also apply to drones and queens (except
when indicated), although characters giv-
en for drones and queens do not necessar-
ily occur in the worker caste. The abbre-
viations Tl, T2, . . . , T8 are used for the

corresponding metasomal terga (S for ster-
num).

Since the literature on honey bees is
vast, some names may have been inadver-
tently overlooked. Researchers discover-
ing names (nomina niida, nova, &c.) omit-
ted here or of earlier usage than the dates
cited are urged to contact me and, if pos-
sible, to send a copy of the relevant pa-
per(s). All in all, however, this list should
prove a reasonable starting point for fu-
ture research.

RECENT HONEY BEES

Genus Apis Linnaeus

Diagnosis. — Modified and updated from
Michener (1990: as Apinae). WORKER:
Labral apex gently and often weakly con-
cave; labrum three to four times broader
than long (median width). Malar space as
long as, or longer than basal width of
mandible. Mandible without dentition

Volume 8, Number 2, 1999

169

Figs. 2-3. Honey bee structural characteristics rep-
resented by Apis {Apis) cerana Fabricius. 2, Worker
head (scale bar = 500 M-m). 3, Inner surface of meta-
tibia-metabasitarsus junction showing pollen press
(auricle), rastellum, and absence of tibial spurs (scale
bar = 200 (jim).

(Fig. 2) (not so for queens or males). Hairs
of compound eye long and erect (Fig. 2).
Vertex extremely short, much less than
ocellar diameter. Scutellum strongly con-
vex and bulging, obscuring metanotum
and basal area of propodeum. Basal area
of propodeum extremely short and decli-
vious. Mesocoxae nearly meeting medial-
ly. Strigilis with prong on anterior margin.
Corbicula, rastellum, and auricle present

(workers only). Metatibial spurs absent
(Fig. 3); metatibia without penicillum; in-
ner surface of metabasitarsus with setal
comb rows (worker only: referred to as
brush combs in Michener 1990). Claws cleft;
arolia present. Distal wing venation strong
and complete; Ir-m and 2r-m strongly an-
gled respective to M (Fig. 3). Marginal cell
long (Fig. 3) and bluntly rounded at apex,
not gently tapering over its length. Jugal
lobe present. Sting straight (worker only).
DRONE: Mandible usually bidentate, fre-
quently only weakly. Holoptic (compound
eyes meeting at top of head). S8 reduced
to transverse bar, without spiculum. Spa-
tha and volsella absent. QUEEN: Mandi-
ble bidentate. Inner surface of metabasi-
tarsus unmodified. Sting curved. MA-
TURE LARVA: Without small dorsal tu-
bercles on segments 1^. Mandible weakly
sclerotized, bluntly pointed, without con-
cavity on inner surface. ETHOLOGY: Nest
a vertical comb of cells made of wax.
Dance language communication system
present. Highly eusocial (with morpholog-
ically distinct castes). New colonies found-
ed by fission and include the old queen.
Brood and storage cells similar. Imma-
tures progressively provisioned.

Biogeography. — Honey bees are predom-
inantly a tropical group and arose in the
Indo-Malayan region during the early Ol-
igocene (Engel 1998a). The genus is native
to Europe, Africa, Madagascar, Arabia, the
Near East, as well as Central and Southern
Asia inclusive of most Southeast Asian Is-
lands. Honey bees are not native to the
Australian or American continents having
been moved to these regions by humans.
Species radiated from southern Asia per-
haps being limited only by temperature
extremes. The clade of living species (sub-
genera Apis, Megapis, and Micrapis: Fig. 1)
arose sometime in the latter half of the
Miocene (Engel 1998a). The development
of cavity-nesting behavior in the earliest
Pliocene (perhaps in the latest Miocene)
enabled at least two of the species, A. cer-
ana and A. melUfera, to extend their ranges

170

Journal of Hymenoptera Research

into more temperate areas. In the case of
A. mellifera this was into Europe and
northwestern Asia while for A. cerana this
was into northeastern China, and the east-
ern regions of the former Soviet Union.
The fossil species were presumably open-
nesting (refer to Engel 1998a for a cladistic
reconstruction of this behavior) and the
slightly warmer temperatures in the Mio-
cene would have partly allowed species
such as A. armbusteri to exist in Europe
even in the absence of well developed
thermoregulatory capabilities. As the tem-
peratures continued to cool (e.g., by the
Pliocene) open-nesting species would
have been forced from these areas and the
cavity-nesting species would be free to
colonize.

Subgenus Apis Linnaeus

Apis Linnaeus 1758: 343, 574. Type species: Apis
mcUifica Linnaeus 1761 (= Apis mellifera Lin-
naeus 1758), designation of Latreille (1810).
Michener 1944: 292. Maa 1953: 557. Michener
1990; 140.

Apicnla Rafinesque 1814: 29. Unjustified re-
placement name for Apis Linnaeus 1758.

Apiarus Rafinesque 1815: 123. Unjustified re-
placement name for Apis Linnaeus 1758.

Apis (Siginntapis) Maa 1953: 556. Type species:
Apis cerana Fabricius 1793, original designa-
tion.

Diagnosis. — WORKER: Forewing length
7-10 mm. Basal vein frequently gently
curved, strongly distad cu-a (Fig. 4). An-
gle of posteroapical margin of first sub-
marginal cell less than 45°. Distal abscissa
of vein M in hind wing variable (present
[Fig. 5] or absent). DRONE: Hind basitar-
sus without thumb-like process. Vertical
arm of T8 longer than horizontal arm; S7
and SB fused mesally. Gonobase absent.
Ventral gonocoxa membranous; dorsal
gonocoxa reduced. Ventral cornua of en-
dophallus recurved ventrally. MATURE
LARVA: Labrum with peg-like setae re-
stricted to apical surfaces and tubercles.
Galea smaller than maxillary palpus. La-
bial palpus not spiculate. Epipharynx with

or without setae. Atrial inner walls
smooth. ETHOLOGY: Nest constructed in
a cavity. Dance language performed on
vertical surface; wagging metasoma posi-
tioned parallel to dance surface; recruits
positioned next to dancer's metasoma
(within a near field sound range). Drone
cell cap variable (present or absent).

Apis (Apis) cerana Fabricius
The Eastern or Asian Honey Bee

Apis cerana Fabricius 1793: 327. [cerana Fabri-
cius]

Apis indica Fabricius 1798: 274. [indica Fabricius]

Apis socialis Latreille 1804a: 390. [indica Fabri-
cius]

Apis peroni Latreille 1804b: 173. [? indica Fabri-
cius: see treatment of javana subspecies]

Apis gronovii Guillou 1841: 323. [? indica Fabri-
cius: see treatment of javana subspecies]

Apis perrolletii Guerin-Meneville 1844: 461.
[indica Fabricius]

Apis delessertii Guerin-Meneville 1844: 461.
[indica Fabricius]

Aprs sinensis Smith 1865: 380. [cerana Fabricius]

Apis mellifica variety japonica Radoszkowski
1887: 436. [japonica Radoszkowski]

Apis delesserti Buttel-Reepen 1906: 168. Unjusti-
fied emendation, [indica Fabricius]

Apis indica variety javana Enderlein 1906: 337.
[javana Enderlein]

Apis indrca Baldensperger 1928: 173. Lapsus cal-
ami, [indica Fabricius]

Apis johni Skorikov 1929b: 251. [johni Skorikov]

Apis indica philippina Skorikov 1929b: 252.
[indica Fabricius]

Apis indica skorikovi Maa 1944: 4. Nomen nudum,
[skorikovi Engel]

Apis mellifera gandhiann Muttoo 1951: 153. No-
men nudum, [indica Fabricius]

Apis {Sigmatnpis) lieftincki Maa 1953: 572. [johni
Skorikov]

Apis (Sigmatapis) samarensis Maa 1953: 580.
[indica Fabricius]

Apis indica sinensis ussuriensis Goetze 1964: 26.
Nomen nudum. Refer to comments under A.
mellifera. [cerana Fabricius]

Apis cerana himalaya Smith 1991b: 154. Nomen
nudum (see below), [skorikovi Engel]

Apis ccrcna Willis, Winston, and Honda 1992:
169. Lapsus calami, [cerana Fabricius]

Volume 8, Number 2, 1999

171

cu-a

Figs. 4-6. Honey bee wing venation. 4, Apii (/l;'/s) cerana Fabricius, worker, forewing. 5, A. {A.) ceraiia,
worker, hind wing. 6, A. {Synapns) hcnshawi Cockerel], worker, forewing reconstruction. In the forewing M
indicates an abscissa of vein M that is called the basal vein, while in the hind wing a distal abscissa is indicated
which is referred to as the indica vein by some authors; cu-a is the cubital-anal crossvein, also called cu-v by
some authors.

Apis nulucnsis Tingek, Koeniger, and Koeniger
1996 [1997]: 116. [mduensis Tingek et ai]

Apis ccrnim hcimifcng Engel 1999: see below.
[heimifeng Engel]

Apis cerana skorikovi Engel 1999: see below, [sko-
rikovi Engel]

Comments. — The name gmnihiaim Mut-
too (1951) was also used by Muttoo (1956)
for an invalid variety of A. cerana. The
name uuluensis was synonymized by my-
self in Schultz et al. (1999) and therefore
appears here for the first time as a sub-

specific entity of A. cerana (see treatment
of subspecies below).

Distribution. — Refer to treatment of in-
dividual subspecies below.

Apis (Apis) koschevnikovi Enderlein,
corrected authorship

The Sundaland Honey Bee

Apis tncllifica imiica variety koschcv)\ikovi Buttel-
Reepen 1906: 192. Unavailable (I.C.Z.N. 1985:
Arts. 10c, 23j, and 50c).

172

Journal of Hymenoitera Research

Apis indica variety koscheimikovi Enderlein 1906:

335. First available usage.
Apis (Sigmatnpis) vechti Maa 1953: 572.
Apis (Sigmatapis) vechti linda Maa 1953: 574.
Apis melUfica adansonii koschevnikowi Goetze

1964: 25. Unjustified emendation.

Comments. — The authorship of this
name has almost universally been given to
Buttel-Reepen who first proposed it in
1906. However, Buttel-Reepen used the
name infrasubspecifically and thus ac-
cording to the I.C.Z.N. (1985: Art. 10c)
does not become available until it is first
used as a species or subspecies, and then
the authorship is assigned to the author
who used the name in such a sense (Arts.
23j and 50c). Enderlein (1906) was the first
to use the name koscheimikovi subspecifi-
cally and he must therefore be considered
as the author.

This species was only recently reinstat-
ed although under the junior synonym A.
vechti (Koeniger et al. 1988, Tingek et al.
1988). Ruttner et al. (1989) recognized that
the correct name for this species was A.
koschevnikovi. There are presently no sub-
specific forms recognized.

Distribution. — Apis koschevnikovi occurs
on the Malay Peninsula, Borneo, Brunei,
Java, Sabah, Sarawak, and Sumatra.

Lectotype.—V^ orkev; BORNEO: Kinaba-
lugebirge, John Worterstradt, L. [leg.];
now with an additional label reading,
"Lectotype, Apis koschevnikovi Enderlein,
desig. M. S. Engel" (deposited in the Mu-
seum fiir Naturkunde, Berlin). The origi-
nal series of A. koschevnikovi used by But-
tel-Reepen is located in the Museum fiir
Naturkunde, Berlin. The type series con-
sists of one worker from northern Borneo
and eight labeled as from Cameroon. No
cerana-MVe bees are natively known from
the African continent, yet the specimens
from "Cameroon" are clearly conspecific
with the one from Borneo, and all corre-
spond to the species widely recognized as
A. koschevnikovi today. The specimens
from Camaroon are very likely misla-
beled; something even Buttel-Reepen

(1906) suspected. Although Maa (1953)
chose Cameroon as the type locality, this
cannot be considered a type fixation since
there are eight specimens from this local-
ity and none are labeled as the type (in
fact, Maa admits having never seen any
specimens of A. koschevnikovi). I therefore
have selected the single specimen from
Borneo to serve as the lectotype.

Apis (Apis) mellifera Linnaeus
The Western Honey Bee

Apis mellifera Linnaeus 1758: 576. [mellifera Lin-
naeus]

Apis mellifica Linnaeus 1761: 421. Unjustified
emendation, [mellifera Linnaeus]

^;);s gregaria Geoffroy 1762: 407. [mellifera Lin-
naeus]

Apis cerifera Scopoli 1770: 16. [mellifera Linnae-
us]

Apis imicolor Latreille 1804b: 168. [iiuicolor La-
treille]

Apis fasciata Latreille 1804b: 171. Preoccupied
{nee Linnaeus 1767, Scopoli 1770). [lamarckii
Cockerell]

Apis adansonii Latreille 1804b: 172. [adansonii La-
treille]

Apis ligustica Spinola 1806: 35. [ligustica Spinola]

Apis capensis Eschscholtz 1822: 97. [capensis
Eschscholtz]

Apis caffra Lepeletier de Saint Fargeau 1836:
402. Preoccupied {nee Linnaeus 1767). [scutel-
lata Lepeletier de Saint Fargeau]

Apis scutellata Lepeletier de Saint Fargeau 1836:
404. [scutellata Lepeletier de Saint Fargeau]

Apis nigritarum Lepeletier de Saint Fargeau
1836: 406. [adansonii Latreille]

Apis daurica Fischer von Waldheim 1843: 1.
[mellifera Linnaeus]

Apis mellifica variety cecropia Kiesenwetter 1860:
315. [cecropia Kiesenwetter]

Apis australis Kiesenwetter 1860: 317. Unjusti-
fied replacement name for Apis ligustica Spi-
nola 1806. [ligustica Spmola]

Apis cerifera Gerstacker 1862: 60. Preoccupied
(nee Scopoli 1770). [sossimai Engel]

Apis mellifica variety remipes Gerstacker 1862:
61. [remipes Gerstacker]

Apis liguna Smith 1861b: 14. Nomoi nudum. [?
ligustica Spinola]

Volume 8, Number 2, 1999

173

Apis mellifica germanica Pollmann 1879: 1. [mel-
lifera Linnaeus]

Apis mellifica caniica Pollmann 1879: 45. [carfiicn
Pollmann]

Apis mellifica hymettea Pollmann 1879: 50. Un-
justified replacement name for Apis mellifica
carnica Pollmann 1879. [carnica Pollmann]

Apis mellifica cypria Pollmann 1879: 52. [cypria
Pollmann]

Apis siciliana Grassi 1881: 1. [siciliana Grassi]

Apis mellifica variety nigrita Lucas 1882: 62. [mel-
lifera Linnaeus]

Apis mcllifida Pollmann 1889: 90. Lapsus calami,
[mellifera Linnaeus]

Apns mellifida [sic] variety caiicasia Pollmann
1889: 90. [caiicasia Pollmann]

Apis ligiirica Dalla Torre 1896: 602. Nomen nu-
dum (uec Tegetmeier 1859, 1860: see below).
Corrected authorship, [ligustica Spinola]

Apis mellifera carniolica Koschevnikov 1900: 1.
Unjustified emendation, [carnica Pollmann]

Apis mellifera cypriaca Koschevnikov 1900: 1.
Unjustified emendation, [cypria Pollmann]

Apis mellifica mellifica variety siziliana Buttel-
Reepen 1906: 168. Unjustified emendation.
[siciliana Grassi]

Apis mellifica unicolor variety syriaca Buttel-Ree-
pen 1906: 175. Unavailable (I.C.Z.N. 1985:
Arts. 10c, 23j, 50c). [syriaca Skorikov]

Apis mellifica mellifica variety lelizeni Buttel-Ree-
pen 1906: 184. Unavailable (I.C.Z.N. 1985:
Arts. 10c, 23j, 50c). [mellifera Linnaeus]

Apis mellifica unicolor variety intermissa Buttel-
Reepen 1906: 187. Unavailable (I.C.Z.N. 1985:
Arts. 10c, 23j, 50c). [intermissa Maa]

Apis mellifica unicolor v ariety friesei Buttel-Ree-
pen 1906: 188. Unavailable (I.C.Z.N. 1985:
Arts. 10c, 23j, 50c). [adansonii Latreille]

Apis mellifera lamarckii Cockerell 1906: 166. Re-
placement name for Apis fasciata Latreille
1804b. [lamarckii Cockerell]

Apis mcllefica Enderlein 1906: 331. Lapsus calami,
[mellifera Linnaeus]

Apis mellifica unicolor variety frisei Enderlein
1906: 335. Lapsus calami, [adansonii Latreille]

Apis mellifera sicula Montagano 1911: 26. [sicili-
ana Grassi]

Apis ndamsoni Meunier 1915: 210. Lapsus calami,
[adansonii Latreille]

Apis fiiscata Meunier 1915: 210. Lapsus calami,
[lamarckii Cockerell]

Apis mellifica kaffra Jack 1916: 397. Lapsus calami,
[scutellata Lepeletier de Saint Fargeau]

Apis mellifera caucasica Gorbachev 1916: 39. Un-
justified emendation, [caucasia Pollmann]

Apis mellifica variety banatica Grozdanic 1926:
57. [carnica Pollmann]

Apis fascrata Baldensperger 1928: 173. Lapsus
calami, [lamarckii Cockerell]

Apis eurasiatica Skorikov 1929a: 14. Unjustified
replacement name for Apis mellifica variety
remipes Gerstacker 1862. [remipcs Gerstacker]

Apis mellifera mellifera natio tesquorum Skorikov
1929a; 29. Unavailable (I.C.Z.N. 1985: Arts.
10c, 23j, 50c). [artemisia Engel]

Apis mellifera remipes natio absuatna Skorikov
1929a: 32. Unavailable (I.C.Z.N. 1985: Arts.
10c, 23j, 50c). [remipes Gerstacker]

Apis mellifera remipes natio siganica Skorikov
1929a: 32. Unavailable (I.C.Z.N. 1985: Arts.
10c, 23j, 50c). [remipes Gerstacker]

Apis mellifera remipes natio georgica Skorikov
1929a: 32. Proposed as new again in Skorikov
(1929b). Unavailable (I.C.Z.N. 1985: Arts. 10c,
23j, 50c). [remipes Gerstacker]

Apis meda Skorikov 1929b: 253. [meda Skorikov]

Apis mellifera natio acervoriim Skorikov 1929b:
253. Preoccupied {nee Linnaeus 1758). [arte-
misia Engel]

Apis remipes transcaucasica Skorikov 1929b: 254.
[remipes Gerstacker]

Apis remipes transcaucasica natio absuana Skori-
kov 1929b: 254. Lapsus calami, [remipes Ger-
stacker]

Apis remipes transcaucasica natio iberica Skorikov
1929b: 254. Unavailable (I.C.Z.N. 1985: Arts.
10c, 23j, 50c). [remipes Gerstacker]

Apis remipes armeniaca Skorikov 1929b: 254. [re-
mipes Gerstacker]

Apis mellifera banata Skorikov 1929b: 263. Un-
justified emendation, [carnica Pollmann]

Apis (Apis) syriaca Skorikov 1929b: pi. 1. First
available usage, [syriaca Skorikov]

Apis mellifica variety sahariensis Baldensperger
1932: 829. [sahariensis Baldensperger]

Apis niger Baldensperger 1932: 830. Nomen nu-
dum. [? mellifera Linnaeus]

Apis mellifera mingrelica Lavrezhin 1935: 656.
Nomen nudum, [remipes Gerstacker]

Apis mellifera taurica Alpatov 1935: 665. Pro-
posed as new again in Alpatov (1938). [taur-
ica Alpatov]

Apis (Apis) intermissa Maa 1953: 591. First avail-
able usage [intermissa Maa].

Apis (Apis) mellifera anatoliaca Maa 1953: 599.
[anatoliaca Maa]

174

Journal of Hymenoptera Research

Apis meUifera siriaca Kerr and Amaral 1960: 12.
Lapsus ailnnii. [si/rincn Skorikov]

Apis mellifcra monticola Smith 1961a: 258. Pro-
posed as new again in Smith {1961b). [imvi-
ticola Smith]

Apis mellifera litorea Smith 1961a: 259. Proposed
as new again in Smith (1961b). [litorea Smith]

Apis mellifera lamarchii Smith 1961b: 148. Lapsus
calami, [lamarckii Cockerell]

Apis millifica Goetze 1964; 9. Lapsus calami, [mel-
lifera Linnaeus]

Apis mellifica intermissa tellica Goetze 1964: 25.
Nomen nudum. See also below, [intermissa
Maa]

Apis mellifica mellifica silvarum Goetze 1964: 26.
Nomen nudum. See also below, [mellifera Lin-
naeus]

Apis mellifica lamarcki Ruttner 1968; 41. Unjus-
tified emendation, [lamarckii Cockerell]

Apis mellifica adami Ruttner 1975: 271. Proposed
as new again in Ruttner (1980) with the spe-
cies called mellifera. [adami Ruttner]

Apis mellifera major Ruttner 1976a; 354. [inter-
missa Maa]

Apis mellifera nubica Ruttner 1976a: 359. [jcmen-
itica Ruttner]

Apis mellifera littorea Ruttner 1976a; 361. Lapsus
calami, [litorea Smith]

Apis mellifcra jemenitica Ruttner 1976a: 366. [je-
menitica Ruttner]

Aptis mellifera carpatica Barac 1977: 270. [cnrnica
PoUmann]

Apis mellifera anatolia Winston 1987; 12. Lapsus
calami, [anatoliaca Maa]

Apis mellifera yemenitica Ruttner 1988: 212. Un-
justified emendation, [jemenitica Ruttner]

Apis mellifera sudanensis Ruttner 1988; 214. No-
men nudum (see below), [jemenitica Ruttner]

Apis mellifera iberica Ruttner 1988: 236. Preoc-
cupied {nee Skorikov 1929b, Goetze 1964: see
treatment of A. mellifera subspecies below);
corrected authorship, [iberiensis Engel]

Apis mellifera macedonica Ruttner 1988: 249. [ma-
cedonica Ruttner]

Apis mellifera logustica Kugler 1988: 268. Lapsus
calami, [ligustica Spinola]

Apis mellifica rodopica Petrov 1991: 17. Proposed
as new again in Petrov (1996). [macedonica
Ruttner]

Apis millifern Willis, Winston, and Honda 1992;
169. Lapsus calami, [mellifera Linnaeus]

Apis mellitera Petrov 1993: 36. Lapsus calami.
[mellifera Linnaeus]

Apis mellifica capratica Petrov 1996; 58. Lapsus
calami, [carnica Pollmann]

Apis mellifera bandasii Radloff and Hepburn
1997; 57. Nomen nudum (see below), [jemeni-
tica Ruttner]

Apis mellifera ruttneri Sheppard, Arias, Grech,
and Meixner 1997 [1998]; 290. [ruttneri Shep-
pard et (?/.]

Apis mellifera artemisia Engel 1999: see below.
Replacement name for Apis mellifera acervo-
runi Skorikov 1929b {nee Linnaeus 1758). [ar-
temisia Engel]

Apis mellifera iberiensis Engel 1999; see below.
Replacement name for Apis mellifera iberica
Ruttner 1988 {nee Skorikov 1929b). [iberiensis
Engel]

Apis mellifera sossimai Engel 1999: see below. Re-
placement name for Apis cerifera Gerstacker
1862 (nee Scopoli 1770). [sossimai Engel]

Comments. — The term "natio" as used
by Skorikov (1929a, b) appears to be syn-
onymous witli "variety" as used by other
authors.

Both Maa (1953) and Goetze (1964) in-
clude the name A. domestica Ray (1710) in
their taxonomic treatment of the v^estern
honey bee. This is a pre-Linnean name
and it is therefore excluded from zoolog-
ical nomenclature (LC.Z.N. 1985: Art. 3)
and my treatment above.

It might be interpreted from my treat-
ment of Pollmann's names above that A.
mellifida as used by this author was not a
lapsus calami but in actuality an unjustified
emendation due to the fact that it appears
to have been used twice (under mellifida
and mellifida caucasia). This is not the case.
The name tnellifida was only used once in
Pollmann's (1889) work in the proposal of
the variety caucasia. It appears twice in my
list since two taxonomic entities are in-
volved in this one action by Pollmann; one
as the lapsus calami mellifida and the second
for the proposal of the variety (now sub-
species) caucasia.

Maa (1953) and Dalla Torre (1896) both
cite the name A. li;^urica with one author
attributing it to Tegetmeier (1859) and the
other to that paper as well as to Teget-
meier (1860). Interestingly enough neither

Volume 8, Number 2, 1999

175

of Tegetmeier's contributions use this
name. Tegetmeier's 1859 paper, the pur-
ported first usage of the epithet, is merely
an untitled note about an exhibition of
Apis specimens at a meeting of the Ento-
mological Society of London and simply
reads, "Mr. Tegetmeier exhibited speci-
mens of Apis ligustica, of which he had
lately received living examples of the
queen and workers from the Continent,
where it is considered a more profitable
species to the owner than the common
honey bee; he hoped, during the next sea-
son, to test the correctness of this opin-
ion." Tegetmeier (1860), a similarly unti-
tled note from a societal meeting, is mere-
ly a follow up to his 1859 notice and uses
the name A. ligustica as well (not A. lig-
urica). A perusal of other issues of the Pro-
ceedings of the Entomological Society of Lon-
don failed to find any usage of A. ligurica,
and it would therefore appear that the first
appearance of such a name was in Dalla
Torre's catalog (1896).

The name mingrelica was used by La-
vrezhin (1935) and attributed to Skorikov
(1929a); however, Skorikov did not pro-
pose such a specific epithet. Skorikov
(1929a: 32, 41, 44) used the name

MHHipejT&cKaa

only in Russian and as part of a vernacu-
lar name (it even has a Russian adjectival
ending which changes in the text as the
case of the noun it modifies changes), nev-
er once proposing it as a latinized taxo-
nomic entity with a description. The same
can be said for several of the nomiua niida
listed by Goetze (1964); these being silva-
rum attributed to Alpatov (1935), tellica at-
tributed to Buttel-Reepen (1906), and us-
suriensis attributed to Lawrjochin (1960).
These names, as used by Goetze (op. cit.),
are also excluded from nomenclature ow-
ing to the fact that they are infrasubspe-
cific names proposed after 1960 (I.C.Z.N.
1985: Art. lb and 45e-g).

Radloff and Hepburn (1997) employ the
subspecific names bandasii (attributed to

Mogga) and sudanensis (attributed to Ras-
had) both of which are nomina nuda. I
failed to find in publications of Rashad
available to me any official proposal of a
subspecies with the latinized name suda-
nensis that would satisfy the rules of no-
menclature, although many of his papers
mention the Sudanese bees as variable
and distinctive (e.g., Rashad and El-Sarrag
1978). Ruttner (1988) also employs the
name sudanensis (attributing it to an un-
published manuscript by Rashad) and
thus it is a nomen nudum with Ruttner ap-
parently being the correct author. The
name baiuiasii, however, was proposed by
Mogga (1988) and would meet the appro-
priate criteria except that this name was
only treated in his unpublished Master's
thesis, is therefore excluded from zoolog-
ical nomenclature [I.C.Z.N., 1985: Art.
9(11)], and means that its use by Radloff
and Hepburn (op. cit.) is a nomen nudum.
Both of these names were once again used
by Hepburn and Radloff (1998) although
they were only mentioned as synonyms of
jemenitica in their consideration of African
races, a synonymy considered amply jus-
tified by both Ruttner (1988) and myself
(above).

This seems an appropriate venue to
comment on the apparent confusion of the
valid name for this species. The persistent
use of the unjustified replacement name A.
mellifica by some modern authors is dis-
tressing and either demonstrates a general
ignorance of taxonomic rules or a flagrant
dismissal of the International Code of
Zoological Nomenclature. Ignorance of
the Code is understandable as not all bi-
ologists are intimately familiar, or even
have an express need to be so familiar,
with its rules and nuances. Dismissal of
the Code is, however, a more serious sit-
uation. The criticism that the name melli-
fica is more descriptive than mellifera is be-
side the point and, if the argument of Ben-
ton (1904) is followed, vastly incorrect
anyway. Suffice to say, the use of A. mel-
lifera versus A. mellifica is not a matter of

176

Journal of Hymenoptera Research

personal choice. Apis melUfera is the only
valid name for this species and authors
should abide by this.

Distribution. —Ihe western honey bee
presumably arose in the Near East or Pon-
tocaspian regions at some point during
the early Pliocene. This species subse-
quently dispersed throughout Africa and
Madagascar (entering via the Arabian
Peninsula), into Europe, and into North-
western Asia as far as the Russian Steppes.
Localized adaptation gave rise to the nu-
merous subspecies which are recognized
today. Spread of this species elsewhere in
the world has been through the action of
human interference. Refer to treatment of
individual subspecies for further details.

Apis mellifera was introduced into the
New World by European settlers during
their colonization of the western hemi-
sphere. Although some early authors felt
that that North American honey bees were
specifically distinct from the European
honey bees (e.g., Belknap 1792) it was
quickly recognized by naturalists of the
time that Apis was not native to the New
World (e.g.. Barton 1793), a belief con-
firmed by their discussions with the Na-
tive American populace. In fact. President
Thomas Jefferson commented that the Na-
tive Americans referred to the honey bees
as "the white man's fly" (Jefferson 1787)
and signaled to them the advance of col-
onists. Early reports that honey bees oc-
curred in the New World (Hernandez
1648) were based on misidentifications
with species of Melipona (Gerstacker 1863).
Most early authors correctly referred to
the transport of honey bee colonies from
Europe by colonists (e.g., Josselyn 1674).
Despite the arguments raging in the late
1700's over the "distinctiveness" and na-
tivity of A. mellifera in the Americas, no
names were proposed for them.

Apis (Apis) nigrocittcta Smith

The Sulawesian Honey Bee

Apis nigrocincta Smith 1861a: 93.

Apis mdlifica indica variety picea Buttel-Reepen

1906: 193. Unavailable (I.C.Z.N. 1985: Arts.
10c, 23j, 50c).
Apis {Signmtapis} nigrocincta marginella Maa
1953: 575.

Co»;?«e»fs.— Although 1 have previously
been hesitant to recognize this species as
distinct from A. cerana (e.g., Engel 1998a),
I here correct this following the excellent
work of Hadisoesilo et al. (1995), Hadisoe-
silo and Otis (1996, 1998), and Hadisoesilo
(1997). The absence of a pore in the drone
cell cap is a plesiomorphic feature of Apis
in general and for the cerana-c\ade in par-
ticular (Fig. 1). Apis nigrocincta is the only
member of the cerana-dade to lack this fea-
ture. The apomorphic presence of such a
pore unites A. cerana and A. koschevnikovi.
No subspecies are recognized in A. nigro-
cincta.

Distribution.— Ihis species only occurs
on the island of Sulawesi except in the ex-
treme southwest corner in a lower eleva-
hon band from a bit northwest of Ujung
Padang around to Bulukumba and be-
yond (G. W. Otis pers. comm.). Previous
records of this species on Mindanao (Otis
1996, Damns and Otis 1997) are probably
A. cerana as revealed by the development
of the drone cell which had a hard cap
with a pore (G. W. Otis pers. comm.), this
being a feature of A. cerana and not A. ni-
grocincta (Hadisoesilo and Ohs 1998). Pop-
ulations of Mindanao are certainly distinc-
tive from typical A. cerana and need to be
investigated further.

Subgenus Megapis Ashmead

Megapis Ashmead 1904: 120. Type species: Apis
dorsata Fabricius 1793, original designaHon.
Maa 1953: 552.

Diflgttosis.— WORKER: Forewing length
12-15' mm. Basal vein frequently gently
curved, strongly distad cu-a (similar to
Fig. 4). Angle of posteroapical margin of
first submarginal cell less than 45°. Distal
abscissa of vein M in hind wing present
(similar to Fig. 4). DRONE: Hind basitar-
sus without thumb-like process. Vertical

Volume 8, Numbek 2, 1999

177

arm of T8 as long as horizontal arm; S7
and S8 not fused mesally. Gonobase ab-
sent. Ventral gonocoxa membranous; dor-
sal gonocoxa reduced. Ventral cornua of
endophallus not recurved ventrally. MA-
TURE LARVA: Labrum with peg-like se-
tae scattered over surface. Galea larger
than maxillary palpus. Labial palpus spic-
ulate. Epipharynx without setae. Atrial in-
ner walls ridged. ETHOLOGY: Nest con-
structed in the open. Dance language per-
formed on vertical surface; wagging me-
tasoma held above dance surface; recruits
potentially far from dancer's metasoma
(potentially outside a near field sound
range). Drone cell cap without pore.

Apis (Megapis) dorsata Fabricius
The Giant Honey Bee

Apis dorsata Fabricius 1793: 328. [dorsata Fabri-
cius]

Apis nigripennis Latreille 1804b: 170. [dorsata Fa-
bricius]

Apis bicolor Klug 1807: 264. Preoccupied (nee Fa-
bricius 1781, Villers 1789). [dorsata Fabricius]

Apis testaeea Smith 1858: 49. [dorsata Fabricius]

Apis zoiiata Smith 1859: 8. Preoccupied (jit'c Lin-
naeus 1758). [binghajiii Cockerell]

Apis laborwsa Smith m Moore et al. 1871; 249.
[laboriosa Smith]

Apis testaeea Smith 1871: 396. Lapsis ealami. [dor-
sata Fabricius]

Megapis zonata (Smith); Ashmead 1904: 121.
[binghami Cockerell]

Megapis dorsata (Smith); Ashmead 1904: 121.
[dorsata Fabricius]

Apis dorsata binghaiiii Cockerell 1906: 166. Re-
placement name for Apis zonata Smith 1859.
[binghami Cockerell]

Apis binghami sladcni Cockerell 1914: 13. [labor-
iosa Smith]

Apis darsata Baldensperger 1928: 173. Lapsus eal-
ami. [dorsata Fabricius]

Apis himnlai/ana Maa 1944: 4. Nomen nudum, [la-
boriosa Smith]

Megapis breviligula Maa 1953: 563. [brevdigula
(Maa)]

Megapis binghann (Cockerell); Maa 1953: 564.
[binghami Cockerell]

Megapis laboriosa (Smith); Maa 1953: 570. [labor-
iosa Smith]

Apis dorsatao Ruttner 1988: 118. Lapsus calami.

[dorsata Fabricius]
Apis labortiosa Willis, Winston, and Honda 1992:

169. Lapsus calami, [laboriosa Smith]

Distribution. — Refer to treatment of in-
dividual subspecies below.

Subgenus Micrapis Ashmead

Micrapis Ashmead 1904: 122. Type species: Apis
florea Fabricius 1787, monobasic and original
designation. Maa 1953: 557.

Diagnosis. — WORKER: Forewing length
6-7 mm. Basal vein frequently gently
curved, strongly distad cu-a (similar to
Fig. 4). Angle of posteroapical margin of
first submarginal cell less than 45°. Distal
abscissa of vein M in hind wing absent.
DRONE: Hind basitarsus with thumb-like
process. Vertical arm of T8 as long as hor-
izontal arm; S7 and S8 not fused mesally.
Gonobase represented by isolated frag-
ments. Ventral gonocoxa sclerotized as
transverse bar; dorsal gonocoxa not re-
duced. Ventral cornua of endophallus not
recurved ventrally. MATURE LARVA: La-
brum with peg-like setae scattered over
surface. Galea larger than maxillary pal-
pus. Labial palpus spiculate. Epipharynx
with setae. Atrial inner walls ridged.
ETHOLOGY: Nest constructed in the
open. Dance language performed on hor-
izontal surface; wagging metasoma held
above dance surface; recruits potentially
far from dancer's metasoma (potentially
outside a near field sound range). Drone
cell cap without pore.

Apis (Micrapis) andreniformis Smith
The Black Dwarf Honey Bee

Apis andreniformis Smith 1858: 49.

Apis florea andreniformis variety sumatrana En-

derlein 1906: 339. Unavailable (I.C.Z.N. 1985:

Arts. 10c, 23j, 50c).
Micrapis andreniformis (Smith); Maa 1953: 601.

Comments. — This species was reinstated
just over a decade ago by Wu and Kuang
(1986, 1987) with additional evicience for
this decision provided by Wongsiri et al.

178

Journal of Hymenoptera Research

(1989). It's common name comes from the
nearly black Tl-2 while in A. florea these
structures are reddish brown. No subspe-
cies are recognized.

Distribution. — Apis andreiiifonnis occurs
as far north as southern-most Yunnan
Province in China, south into Malaysia,
eastward on the mainland to the coast of
Vietnam, and westward as far as north-
western India along the eastern border of
Nepal. Although specimens are not re-
corded from Bhutan, Cambodia, Myan-
mar, or Nepal the records of A. andreiiifor-
iiiis in the Bengal, Sikkim, and Assam re-
gions of India suggests that it will even-
tually be found in these countries (Otis
1996) unless it has since become locally ex-
tinct.

Apis (Micrapis) florea Fabricius
The Red Dwarf Honey Bee

Apis florea Fabricius 1787: 305.

Ayis seminifa Hoffmannsegg 1818: 60.

Apis lobatn Smith 1854: 416.

Apis floralis Home and Smith 1870: 181. Lapsus

calami.
Apis testacea Bingham 1898: 129. Preoccupied

(nee Smith 1858).
Micrapis florea (Fabricius); Ashmead 1904: 122.

[see Comments below]
Apis florea variety rufiventris Friese in Buttel-

Reepen 1906: 167, 170.
Apis florea florea variety fnscata Enderlein 1906:

338. Unavailable (I.C.Z.N. 1985: Arts. 10c, 23j,

50c).
Apis nursci Cockerel! 1911a: 319. Replacement

name for Apis testacea Bingham 1898.
Apis florea nasicana Cockerell 1911b: 241.

Comments. — Although when proposing
the genus Micrapis Ashmead (1904) did
not explicitly transfer the type species, A.
florea, into this genus (he shll referred to
the species as A. florea and not Micrapis
florea) he is to be consider as having made
the combination as the proposal of Mi-
crapis was done prior to 1961 [I.C.Z.N.
1985: Art. 51c(ii)]. There are presently no
subspecies recognized in A. florea.

Distribution. — This species is distributed

from the eastern regions of the Arabian
Peninsula, through southern Iran and Iraq
to Pakistan and India as far north as the
Himalayan uplift. The species extends
eastward as far as Vietnam, north to Yun-
nan Province, China, and south into In-
donesia. Refer to Otis (1996) for more pre-
cise locality records. Apis florea was intro-
duced into Sudan a little over a decade
ago from Pakistan and has become estab-
lished (Lord and Nagi 1987, Mogga and
Ruttner 1988).

SUBSPECIES

Owing to both the high visibility and
dramatic geographic variation of the hon-
ey bees, the more common species have
been heavily divided infraspecifically. Be-
low I have attempted to outline the pres-
ently recognized subspecies. There are no
valid subspecies of A. andreniformis, A. flo-
rea, A. koschevnikovi, or A. >iigrocincta and
1 have not recognized any subspecies
among the fossil forms. A few of the
names treated below are used incorrectly
under the rules of zoological nomencla-
ture (I.C.Z.N. 1985) and are therefore
treated in further detail, otherwise I have
not given detailed taxonomic histories for
each subspecies. I have tried to provide
common names for these morphs as well
as historical, geographical, or important
biological comments for some taxa.

As far as I have been able to ascertain,
holotypes or neotypes have not been des-
ignated for certain subspecies. This will be
necessary to assure stability in nomencla-
ture. The general problems with the rec-
ognition of subspecies have been dis-
cussed by several authors, the best cri-
tique being that by Wilson and Brown
(1953). My recommendation is a complete
cessation of proposals of new subspecies
in Apis (primarily A. mellifera) until the
taxonomy has been more thoroughly ex-
amined (something which 1 hope this pa-
per will facilitate).

Volume 8, Number 2, 1999

179

Apis {Apis) cerana Fabricius

The populations of A. cerana in the Phil-
ippines are somewhat distinctive (e.g.,
Ruttner 1988, Smith 1991b, Smith and Ha-
gen 1996, Damus and Otis 1997) and could
possibly be elevated to subspecific status.
If this is done the name of this race would
become A. cerana philippina Skorikov with
A. samareusis Maa as a junior synonym.

1. Apis cerana cerana Fabricius: The

Eastern Honey Bee

Distribution. — This subspecies occurs as
far east as Afghanistan and south along
the central deserts and mountain ranges.
Along the Himalayan uplift in Tibet it is
replaced by A. cerana skorikovi and further
south in India it is replaced by A. cerana
indica. The subspecies then ranges across
most of central and southern China al-
though not in the high regions of central
China where the black bee (A. cerana heiin-
ifeng) occurs. Apis cerana cerana then rang-
es along the eastern edge of Asia up to
Korea and the southern-most areas of the
former Soviet Union (Ussuria) and south
to the northern reaches of Vietnam.

2. Apis cerana heimifeng Engel, new
subspecies: The Black Chinese Honey
Bee

Apis cerana, Aba race, Peng et al. 1989: 15.

Diagnosis. — Distinguishable from typi-
cal A. cerana cerana in China by the dark
brown to black scutellum and T3-4 (these
are yellow in A. cerana cerana). In this re-
spect it resembles the Tibetan Honey Bee,
A. cerana skorikovi, from which it differs in
the larger body size (body length 12.5-13.7
mm; forewing length 8.8-9.3 mm) and low
tendency to swarm (very strong in A. cer-
ana skorikoz'i). This is the largest A. cerana
race in China and is a relatively dark bee
(hence its common name) with a dark me-
tasoma, scutellum, and legs. This dark col-
oration is partly owing to the very narrow
to completely absent bands of orange on
T3-5. Peng et al. (1989) provide several

morphometric measures for this bee (as
"Aba race").

Distribution. — This is a distinctive race
in central China and occurs in relatively
high elevation areas (although certainly
not at the highest points for this region)
mostly near river valleys. Its general dis-
tribution is in northern Sichuan Province,
southwestern Gansu Province, and eastern
Qinghai Province.

Holoti/pe.— Worker; CHINA: [Northern
Sichuan Province], Nanping, 21 April 1920
(deposited in the American Museum of
Natural History, New York).

Etyniologi/. — The specific epithet is de-
rived from the Mandarin words hei (mean-
ing "dark") and mifeng (meaning "honey
bee") and refers to the dark coloration of
this race. The name is a noun in apposi-
tion.

3. Apis cerana indica Fabricius: The

Indian Honey Bee

Distribution. — This race occurs through
most of India except in the higher regions
towards the north where it is replaced by
A. cerana cerana or in Tibet by A. cerana
skorikovi.

4. Apis cerana japonica Radoszkowski:

The Japanese Honey Bee

Distribution. — This race occurs on the is-
lands of Japan.

5. Apis cerana javana Enderlein: The

Javanese Honey Bee

Comments. — The taxa described as A.
peroni by Latreille (1804b) and A. gronovii
by Guillou (1841) may be the same as En-
derlein's A. cerana javana. Owing to this
uncertainty 1 have for now retained A. cer-
aim javana as the name of this taxon and
considered A. peroni and A. gronovii to be
A. cerana indica. Researchers, however,
should be aware of this uncertainty and if
the consubspecific nature of these taxa is
confirmed in the future, then the name of
the subspecies would have to be changed
to A. cerana peroni with gronovii and javana

180

Journal of Hymenoptera Research

as junior synonyms. This situation was
also noted by Maa (1953: 580). If this un-
fortunately turns out to be the case, then
it would probably be advisable to petition
the I.C.Z.N. to suppress A. peroni and A.
gronovii in favor of A. cerana javana al-
though such a case would not be a very
strong one owing to the fact that A. peroni
has been in use within the last 50 years
(e.g., Maa 1953).

Distribution. — This subspecies is pres-
ently known from Java and as far east as
Timor.

6. Apis cerana johni Skorikov: The

Sumatran Honey Bee

Distribution. — This subspecies only oc-
curs on the island of Sumatra.

7. Apis cerana mihiensis Tingek,
Koeniger, and Koeniger: The Malaysian
Mountain Honey Bee

Distribution. — This morph occurs in the
mountains of Sabah, Malaysia. Otis (1996)
provides details of various collection lo-
cales in Malaysia.

8. Apis cerana skorikovi Engel, new
subspecies: The Himalayan Honey Bee

Apis cerana skorikovi Maa 1944: 4. Nonien nudum.
Apis cerana, Himalaya race, Ruttner 1988: 121,

158.
Apis cerana, Xizang race, Peng et al. 1989: 15.
Apis cerana himalaya Smith 1991b: 154. Nomen

nudum.

Diagnosis. — This race is fairly gray in
overall body coloration, some areas being
slightly more yellow while others are clos-
er to black (e.g., T4 is entirely black) and
with paired yellow spots on S3. There are
distinctive white hairs on the terga which
contribute to the overall grayish appear-
ance. Apis cerana skorikovi, like A. cera)m
heimifeng, also differs from A. cerana cerana
(as well as A. cerana indica and other sub-
species) in the dark scutellum and slightly
larger size (body length around 12.2 mm;
forewing length around 8.75 mm). Addi-

tional morphometric values are provided
by Peng et al. (1989).

Comments. — The Himalayan race of A.
cerana (sometimes called the Tibetan race)
was referred to as A. cerana skorikovi by
Maa (1944); however he never provided a
description for this subspecies. Many au-
thors have subsequently recognized this
race as distinctive but never associated a
name with the numerous descriptions
simply referring to this taxon as the Ti-
betan honey bee (among other common
names). Peng et al. (1989) have done the
best to characterize and describe this tax-
on referring to it as the Xizang race (Xi-
zang is Chinese for Tibet, sometimes
spelled Sitsang). I here validate the name
A. cerana skorikovi for this taxon, referring
readers to the diagnosis provided above
as well as the descriptive comments given
by Peng et al. (1989: as "Xizang race") and
Ruttner (1988: as A. cerana "Himalaya
race"). Smith (1991b) referred to this race
as A. cerana hinialai/a making a Latin name
out of Ruttner's "A. cerana Himalaya
race". Ruttner's name was not used as,
nor intended as, a scientific name and A.
cerana himalaya as used by Smith (1991b)
is a nomen nudum.

Distribution. — This subspecies occurs
along the Himalayan uplift at elevations
of 1900-4000 meters.

Ho/ofype.— Worker; NEPAL: Ko Pasi
near Panauti, 19 March 1982, B. A. Under-
wood (deposited in the Cornell University
Insect Collection, Ithaca).

Paratypes. — Three workers, same collec-
tion data as holotype, deposited in the
same collection as the holotype; two work-
ers, same collection data as holotype, de-
posited in the American Museum of Nat-
ural History, New York.

Etymology. — I have retained the specific
epithet used by Maa (1944) for this race.
The epithet is a patronymic honoring Dr.
A. S. Skorikov who did much work on the
classification of bees; honey bees and
bumble bees in particular.

Volume 8, Number 2, 1999

181

Apis (Apis) mellifera Linnaeus

The western honey bee, A. mellifera, is
remarkable for its geographic variation
and, for this reason, has been broken into
a plethora of taxonomic entities over the
past two centuries. In earlier studies these
forms were often treated as distinct spe-
cies; however, today they are recognized
as subspecies of a single variable species.

1. Apis mellifera adami Ruttner: The

Cretan Honey Bee

Comments. — No type has been designat-
ed for this race. Ruttner's specimens are
among the collection of the Institut fiir Bi-
enenkunde in Oberursel, Germany. A sin-
gle worker from this material should be
selected as the lectotype and the others
designated as paralectotypes.

Distribution. — This race occurs on the is-
land of Crete in the Mediterranean Sea.

2. Apis mellifera adansonii Latreille: The

West African Honey Bee

Distribution. — This subspecies has a
wide distribution in western Africa rang-
ing from Niger in the north, east to Sene-
gal, and as far south as Zaire.

3. Apis mellifera anatoliaca Maa: The

Anatolian Honey Bee

Distribution. — This race occurs through-
out Turkey, bordering with A. mellifera ma-
cedonica in the European portion of that
country. In the east, A. mellifera anatoliaca
borders with A. tnellifera caucasia and A.
mellifera remipes in the area of Azerbaijan,
Armenia, and Georgia and with A. melli-
fera meda in the southeastern-most regions
of Turkey.

4. Apis mellifera artemisia Engel, new
name: The Russian Steppe Honey Bee

Apis mellifera incUifera natio tesquorum Skorikov
1929a: 29. Unavailable (I.C.Z.N. 1985: Arts.
10c, 23), 50c).

Apis mellifera acervorum Skorikov 1929b: 253.
Preoccupied (nee Linnaeus 1758).

Etymology. — Named for Artemis (Greek;
Diana in Roman mythology), the goddess
for whom the honey bee was a symbol
and whose temple at Ephesus, the Artemi-
sium, was listed by Callimachus of Cyrene
and Philo of Byzantium as one of the sev-
en wonders of the world.

Distribution. — This race occurs in the
central Russian Steppes.

5. Apis mellifera capensis Eschscholtz:

The Cape Honey Bee

Comments. — This subspecies is biologi-
cally distinctive for its ability to undergo
parthenogenetic reproduction when de-
prived of a queen (Jack 1916, Anderson
1963, Ruttner 1977).

Distribution. — As is indicated by the
name of this race, it occurs in the Cape
region of South Africa.

6. Apis mellifera caniica Pollmann: The
Carnolian Honey Bee

Distribution. — This European race oc-
curs south of the Alps, east into northern
Italy, and west into Yugoslavia and Ro-
mania.

7. Apis mellifera caucasia Pollmann,
reinstated name: The Caucasian Honey
Bee

Apis mellifida [sic] variety caucasia Pollmann
1889: 90.

Apis mellifera caucasica Gorbachev 1916: 39. Un-
justified emendation.

Distribution. — This race occurs in the
Caucasus Mountains. It is sometimes re-
ferred to as the "grey" Caucasian honey
bee.

8. Apis mellifera cecropia Kiesenwetter:
The Greek Honey Bee

Distribution. — This race occurs through-
out most of Greece and surrounding Ae-
gean islands. It borders A. mellifera mace-
donica in the northeast regions of Greece
and A. mellifera carnica in the northwest
corner of the country.

182

Journal of Hymenoptera Research

9. Apis mellifera cypria PoUmann: The
Cyprian Honey Bee

Distribution. — This race is presently
known only from the island of Cyprus.

10. Apis mellifera iberiensis Engel, new
name: The Iberian Honey Bee

Apis nicllifica intermissa ibericn Goetze 1964: 25.
Infrasubspecific name proposed after 1960
and therefore unavailable (I.C.Z.N. 1985: Art.
lb); preoccupied (nee Skorikov 1929b).

Apis mellifera iberica Ruttner 1988: 236. Correct-
ed authorship (first available usage); preoc-
cupied {nee Skorikov 1929b).

Comments. — Goetze (1964) proposed
this form as a variety of the subspecies in-
termissa. Since this is a varietal name (i.e.,
infrasubspecific) proposed after 1960 it is
unavailable and does not enter into zoo-
logical nomenclature (I.C.Z.N. 1985: Arts,
lb and 45e-g). The first correct application
of this name as a subspecies was by Rutt-
ner (1988) which thereby validates the
name to that author. Unfortunately, a fur-
ther complication arises as the name is a
primary junior homonym of an available
name proposed by Skorikov (1929b) for
another type of honey bee from Azerbai-
jan. Since no type exists for Ruttner's sub-
species I here designate a neotype for A.
mellifera iberica, the name of which will be
replaced by A. mellifera iberiensis.

Neotype. — Worker; SPAIN: Cordoba,
Andalusia, September 1986, D. Smith and
R. Hagen; now with additional labels
reading "NEOTYPE, Apis mellifera iberica
Ruttner, desig. M. S. Engel, 1999" and
"Apis mellifera iberiensis Engel" (deposited
in the Snow Entomological Collection,
Natural History Museum, University of
Kansas, Lawrence). Three additional
workers, identical to the neotype, have
also been deposited with the University of
Kansas and the American Museum of
Natural History, New York.

Etymology. — The new subspecific epi-
thet refers to the distribution of this sub-
species on the Iberian Peninsula.

Distribution. — This race natively occurs
on the Iberian Peninsula in Europe. Colo-
nies were transferred to the South Amer-
ican tropics several decades ago, prior to
the introduction of A. mellifera scutellata,
but populations never became established.

Two ecotypes of A. mellifera iberiensis oc-
cur in Spain as shown by Santiago et al.
(1986) as well as two mitochondrial types
as demonstrated by Smith et al. (1991) and
Smith and Glenn (1995). The neotype se-
lected above comes from the Cordoba
populations sampled by Smith and Glenn
(1995: their colony 1-1).

11. Apis mellifera intermissa Maa,
corrected authorship: The Tellian Honey
Bee

Comments. — As was noted in the taxo-
nomic history of A. mellifera, the name in-
termissa as used by Buttel-Reepen (1906) is
unavailable for the same reasons as pre-
sented for Buttel-Reepen's name koschev-
nikovi (see Comments for A. koschevnikovi).
The name of this subspecies was first
made available by Maa (1953).

Distribution. — This race has a tight dis-
tribution along the northern coast of Af-
rica as far west as Morocco, into Tunisia
in the east, but bordered by the Atlas
range in the south.

12. Apis mellifera jemenitica Ruttner,
reinstated name: The Arabian or Nubian
Honey Bee

Apis mellifern nnbien Ruttner 1976a: 359.

Apis mellifera jemenitica Ruttner 1976a: 366.

Apis mellifera yemenitica Ruttner 1988: 212. Un-
justified emendation.

Apis mellifera sudanensis Ruttner 1988: 214. No-
men nudum. Corrected authorship.

Apis mellifera bandasii Radkiff and Hepburn
1997: 57. Nomeii nudum. Corrected author-
ship.

Comments. — Ruttner (1988) synony-
mized A. mellifera jemenitica with A. melli-
fera nubica and acting as first reviser
(I.C.Z.N. 1985: Art. 24) gave jemenitica pri-
ority over nubica. In addition, the name/V-

Volume 8, Number 2, 1999

183

menitica was unnecessarily emended from
its original spelling. The correct name of
this taxon should be A. mellifera jemenitka,
not yemenitica.

No type has been designated for this
race. See comments under A. mellifera ada-
mi for location of Ruttner's original mate-
rial.

Distribution. — This subspecies of small
bees occurs in hot arid zones of eastern
Africa and the Arabian Peninsula. Coun-
tries in which it occurs include Chad,
Oman, Saudi Arabia, Somalia, Sudan, and
Yemen.

13. Apis mellifera lamarckii Cockerel!:
The Egyptian Honey Bee

Distribution. — This honey bee race oc-
curs in a narrow range along the Egyptian
Nile Valley.

14. Apis mellifera ligustica Spinola: The
Italian Honey Bee

Distribution. — This subspecies occurs
along the Italian Peninsula although it has
been commercially transported through-
out the world. The bees currently distrib-
uted in Italy have hybridized much with
A. mellifera mellifera and A. mellifera carnica
in the north. In fact, untainted populations
of A. mellifera ligustica appear to be con-
fined to Kangaroo Island, Australia where
they are being maintained as an unhybri-
dized strain (Ruttner 1976b). I recently
(January 1999) had the opportunity to visit
Kangaroo Island and to see one of these
colonies.

15. Apis mellifera litorea Smith: The

East African Honey Bee

Comments. — This name was originally
proposed by Smith (1961a) but was also
proposed as new by Smith (1961b). No
type appears to have been designated for
this subspecies.

Distribution. — This subspecies is distrib-
uted along the eastern coast of tropical Af-
rica occurring from Kenya (perhaps even

the southern-most portions of Somalia)
south to Mozambique.

16. Apis mellifera macedonica Ruttner:
The Macedonian Honey Bee

A}ns mellifera macedonica Ruttner 1988: 249.
Apis mellifica rodopica Petrov 1991: 17. New syn-
onymy.

Comments. — No type has been designat-
ed for this race. See comments under A.
mellifera adami for location of Ruttner's
original material. The name rodopica was
proposed again as new by the same au-
thor five years later (Petrov 1996). There
appears to have been no type designated
for Petrov's race either.

Distribution. — This subspecies occurs as
far north as southern Romania, east to Yu-
goslavia, and south to northern Greece
where it borders A. mellifera cecropia. In the
Carpathian Mountains and in Yugoslavia
it borders A. mellifera carnica.

17. Apis mellifera meda Skorikov: The
Median Honey Bee

Distribution. — This race is most common
in Iran and Iraq but does range into south-
eastern Turkey and northern Syria.

18. Apis mellifera mellifera Linnaeus:
The Western or European Honey Bee

Distribution. — This subspecies originally
ranged throughout central Europe north
of the Alps, as far south as southern
France in the west, southern Sweden in
the north, central Russia in the east, and
on the British Isles. In the Ukraine there is
a transition over the steppe region to A.
mellifera sossimai.

19. Apis mellifera monticola Smith: The
East African Mountain Honey Bee

Comments. — The name was first pro-
posed by Smith (1961a) but was designat-
ed as new a second time in Smith (1961b).
No type appears to have been designated
for this subspecies.

Distribution. — This race occurs within
the mountains of eastern Africa (e.g., in

184

Journal of Hymenoptera Research

Kenya and Tanzania). The occurrence of

A. mellifera motiticola in the mountains of
Cameroon should be checked carefully.

20. Apis mellifera remipes Gerstacker,
reinstated name: The Yellow Armenian
Honey Bee

Apis remipes Gerstacker 1862: 61.

Apis mellifera armeniaca Skorikov 1929b: 254.

Comments. — The name armeniaca was
subjectively chosen by Ruttner (1988: 192)
over remipes. This decision is invalid be-
cause remipes has priority and the senior
synonym must be recognized as the name
for the subspecies.

Distribution. — This race occurs in Ar-
menia and may be the same as A. mellifera
anatoliaca, in which case Gerstacker's
name has priority for the subspecies.

21. Apis mellifera ruttneri Sheppard,
Arias, Grech, and Meixner: The Maltese
Honey Bee

Comments. — No type was originally des-
ignated for the subspecies but this is here
corrected by the original authors thereby
stabilizing the name of this taxon.

Lectotype. — Worker; MALTA: St. Julians,
10 August 1995, W. S. Sheppard; desig-
nation of W. S. Sheppard, M. C. Arias, A.
Grech, and M. D. Meixner. The lectotype
now bears a label indicating it as such and
is deposited in the M. T. James Entomo-
logical Museum, Washington State Uni-
versity.

Paralectotypes. — Nine workers; same col-
lection data as lectotype; deposited in the
same collection as the lectotype. A further
two paralectotype workers are deposited
in the American Museum of Natural His-
tory, New York. All paralectotypes desig-
nated by W. S. Sheppard, M. C. Arias, A.
Grech, and M. D. Meixner.

Distribution. — This race is only distrib-
uted on the island of Malta in the Medi-
terranean Sea.

22. Apis mellifera sahariensis
Baldensperger: The Saharan Honey Bee

Comments. — Ruttner (1988) attributed
this name to Baldensperger (1923). He has,
however, unfortunately confused recog-
nition of the subspecific entity with taxo-
nomic proposal of the subspecies. Al-
though Baldensperger (1923) notes the
distinctive character of the Saharan honey
bees he does not provide a latinized name
for them (in fact, nowhere in his work
does he even mention the name Apis, let
alone sahariensis). Instead Baldensperger
(1923) only uses the vernacular name
'T'abeille saharienne". Thus, as far as I
have been able to determine, the name is
first made taxonomically available by Bal-
densperger in his later paper (1932) where
he uses the latinized name and provides a
diagnosis.

This is one of the races for which I know
that no type was ever designated; how-
ever I have not been able to confirm
whether any material survives of Balden-
sperger's original colony which he
brought back to France. I have therefore
hesitated to designate a neotype. This race
is markedly lighter in coloration, particu-
larly on Tl-3, than the more common A.
mellifera intermissa (which has a noticeably
shiny, dark brown to black integument
with more sparse pubescence). See Rutt-
ner (1988) for further descriptive details. It
is possible that this race is synonymous
with A. mellifera lamarckii in which case the
name lamarckii has priority for the subspe-
cies. This possibility should be closely ex-
amined in the future.

Distribution. — This race, like A. mellifera
intermissa, has a tight range in northwest-
ern Africa. It occurs along the southern
side of the Atlas range.

23. Apis mellifera scutellata Lepeletier
de Saint Fargeau: The African Honey
Bee

Comments. — This is the race introduced
into Brazil in the 1950's which has capti-

Vol UME 8, Number 2, 1999

185

vated the apicultural world as well the
imagination of the public; popularly
known as the "Africanized Honey Bee" or
"Killer Bee". An account of the transport
of African bees to Brazil and their acciden-
tal release is given by Kerr (1957, 1967: see
also Michener 1975, Spivak et al. 1991,
Taylor 1977, 1985 for information on the
introduction, spread, and effect of this
race in the western hemisphere and for di-
rection to other literature sources).

Distribution. — This subspecies also has a
large distribution in Africa and ranges
from South Africa northward along the
eastern half of the continent to about So-
malia, it apparently does not occur along
the eastern coastal plain where A. nielli fern
litorea is found.

24. Apis meUifera siciliana Grassi,
reinstated name: The Sicilian Honey Bee

Apis siciliana Grassi 1881: 1.

Apis mellifica mellificn variety siziliana Buttel-

Reepen 1906: 168. Unjustified emendation.
Apis sicula Montagano 1911: 26.

Comments. — Ruttner's (1988) arbitrary
choice of the epithet sicula over siciliana for
the Sicilian race of honey bees does not
meet the criterion of priority and thus the
name siciliana must be reinstated for this
morph.

Distribution. — This subspecies occurs on
the island of Sicily in the Mediterranean
Sea.

25. Apis mellifera sossimai Engel, new
name: The Ukrainian Honey Bee

Apis cerifera Gerstacker 1862; 60. Preoccupied
{nee Scopoli 1770).

Etymologi/. — The new specific epithet is
derived from St. Sossima, patron saint of
beekeeping in the Ukraine. St. Sossima
may be a Christian version of the bee-god
Zosim of some early pagan tribes of Rus-
sia.

Distribution. — This race occurs along the
eastern borders of A. nwllifera carnica's
range. It occurs mostly in the Ukraine and

easterly over to the northern regions of the
Caucasus Mountains where it borders A.
mellifera caucasia. South in the Crimea it is
replaced by A. mellifera taurica.

26. Apis mellifera syriaca Skorikov,
corrected authorship: The Syrian Honey
Bee

Comments. — As was noted in the taxo-
nomic history of A. mellifera, the name si/r-
iaca as used by Buttel-Reepen (1906) is un-
available for the same reasons as present-
ed for Buttel-Reepen's name koschevnikovi
(see Comments for A. koschevnikovi). The
name of this subspecies was first made
available by Skorikov (1929b).

Distribution. — Apis mellifera syriaca oc-
curs along the eastern shores of the Med-
iterranean Sea; north from Syria to the Ne-
gev Desert in the south. It is sometimes
known as the Palestine honey bee.

27. Apis mellifera taurica Alpatov: The
Crimean Honey Bee

Distribution. — This race occurs along the
north-central shores of the Black Sea; in
the Crimea.

28. Apis mellifera tmicolor Latreille: The

Malagasy Honey Bee

Distribution. — The distribution of this
race in Madagascar is discussed by Brooks
and Michener (1988).

Apis (Megapis) dorsata Fabricius

Some authors prefer to recognize one or
more of the A. dorsata races as distinct spe-
cies. This is most often done with A. dor-
sata laboriosa followed by A. dorsata brevi-
li^ula and to a much lesser degree with A.
dorsata biiighami.

1. Apis dorsata binghami Cockerell: The

Giant Sulavv'esi Honey Bee

Distribution. — This race of giant honey
bees occurs on the island of Sulawesi.

186

Journal of Hymenoptera Research

2. Apis dorsata breviligula (Maa): The
Giant Philippine Honey Bee

Comments. — Since Maa (1953) originally
proposed the name breviligula in the genus
Megapis his name should be written in pa-
rentheses (I.C.Z.N. 1985: Art. 51c).

Distribution. — This morph of giant hon-
ey bees occurs in the Philippines. The spe-
cies status of this race is oft argued based
on nest site characteristics as reported by
Morse and Laigo (1969: as A. dorsata) and
Starr et al. (1987).

3. Apis dorsata dorsata Fabricius: The
Common Giant Honey Bee

Distribution. — This subspecies has the
largest distribution of the group; ranging
from India, east to the coast of Vietnam,
and into the southeast Asian islands
where it is sometimes replaced by other
subspecies (see accounts for binghami and
breviligula).

4. Apis dorsata laboriosa Smith: The
Giant Himalayan Honey Bee

Comments. — I currently do not recognize
the subspecific form laboriosa as a separate
species (see also Engel 1998a; but see Sak-
agami et al. 1980, Roubik et al. 1985,
McEvoy and Underwood 1988, Under-
wood 1990a, b). Growing evidence, how-
ever, suggests that the distinction may be
valid and my decision to place laboriosa
within dorsata should be examined more
closely. Sakagami et al. (1980) provided a
detailed account of the morphology of la-
boriosa versus typical dorsata. Many of the
characters they use to justify specific sta-
tus are, however, quite variable (e.g., pro-
tuberance of the ocelli, general pubescence
patterns, &c.) and across a large range of
specimens blend naturally into one anoth-
er except some measures of body size
which I presently feel are more indicative
of a subspecies rather than of a separate
species. One character which appears to
me to be somewhat reliable is the minute,
broad medioapical extension of the grad-

ulus on S3 in dorsata while this is com-
pletely absent in laboriosa.

Distribution. — This subspecies is distrib-
uted at high aWtudes (1,200 to 4,000 m)
on the slopes of the mountains from
northernmost India to the northernmost
boundary of Laos. Further details of its
distribution are provided by Otis (1996).

FOSSIL HONEY BEES

More paleontological work has focused
on the honey bees than on any other
group of bees. Recently I have treated
some of these extinct species and attempt-
ed to place them into a phylogenetic
framework with the living species (Engel
1998a: Fig. 1). Petrov (1992, 1997) also dis-
cussed Apis evolution with reference to
the fossil record, in particular focusing on
the origin of the Bulgarian honey bee (A.
mellifera rodopica = A. mellifera macedonica
in my system). Hong (1984) described a
compression fossil from the Early Creta-
ceous of China as the oldest fossil bee and
as a relative of Apis. Hong's Palaeapis bei-
boziensis is in actuality a sphecid wasp
(Michener 1997, Engel 1998a) and has no
bearing on apine evolution. Lastly, Nel et
al. (1999) presented a small review of fossil
Apis specimens but overlooked the species
presented by myself, recent treatments of
Apini (e.g., Michener 1990), as well as the
numerous synonymies within the group,
instead simply regurgitating the classifi-
cation of Zeuner and Manning (1976)
which is fraught with errors (e.g., refer to
Engel 1998a). Nel et al. (op. cit.) have also
followed Zeuner and Manning (1976) and
other authors in over splitting "species" of
honey bees based on minor morphometric
differences in wing venation and size (e.g.,
their unnamed species A-J: Nel et al. 1999);
characters which distinguish mere subspe-
cies in Apis at best. These authors conclud-
ed (p. 31) that the fossil honey bees could
not be studied from a phylogenetic per-
spective; however, such an attempt had al-
ready been successfully undertaken at
least three times previously (e.g., Buttel-

VdiUME 8, Number 2, 1999

187

Reepen 1906, Statz 1931, Engel 1998a). I
am presently involved in a monographic
study of the fossil bees, particularly those
from Baltic amber, and cladistic analyses
have been successfully completed for the
living and fossil corbiculate bees, includ-
ing Apini (Engel 1998b, in prep.).

In order to maintain the monophyly of
the generally recognized subgenera (re-
cent and fossil) it seems appropriate to
propose two new subgenera thereby
breaking up the paraphyletic subgenus
Si/iiapis. One of these new groups was
called Hnuffnpis by Armbruster (1938), but
he failed to designate a type species for
the genus, thereby making this name un-
available according to the I. C.Z.N. (1985:
Art. 13c) (see also discussions in Michener
1990, 1997). None of the fossil species be-
longs to the living subgenera Apis, Megnp-
is, or Micrapis.

Subgenus Cascapis Engel, new
subgenus

Hauffapis Armbruster 1938: 37. Unavailable as
no type species was originally designated
(I.C.Z.N. 1985: Art. 13c). See also Michener
(1990, 1997).

Ti/pe species. — Apis arnibrusteri Zeuner
1931.

Diagnosis. — WORKER: Basal vein gently
curved, strongly distad cu-a (similar to
Fig. 4). Angle of posteroapical margin of
first submarginal cell greater than 45°. Dis-
tal abscissa of vein M in hind wing present
(similar to Fig. 5). Drone, mature larva,
queen, and ethology all unknown.

Etymology. — The new genus-group
name is a combination of cascus (L. old)
and apis (L. bee).

Comments. — The subsequent designa-
tion of Hauffapis scheiithlei by Zeuner and
Manning (1976) as the type species was in
the synonymy of Hauffapis with Apis and
is therefore invalid (I.C.Z.N. 1985: Art.
lie) (see also Michener 1990).

Apis (Cascapis) armbnisteri Zeuner

Armbruster's Honey Bee

Ayis armbnisteri Zeuner 1931: 292.
Hnuffnpis sclieiithlci Armbruster 1938: 43.
Hnuffapis scheeri Armbruster 1938: 43. New syn-
onymy.
Hnuffnpis schnnuniiiii Armbruster 1938: 44. New

synonymy.
Hnuffnpis scheeri variety gaUnuni Armbruster

1938: 45. New synonymy.
Hnuffnpis scheeri variety rnhdei Armbruster 1938:

45. New synonymy.
Hnuffnpis scheuthlei variety seeiiminii Armbruster

1938: 45. New synonymy.
Hnuffnpis scheuthlei variety zeuiieri Armbruster

1938: 45. New synonymy.
Apis nnnbruiteri chcuthlei Hong and Miao 1992:

2. Lnpsus cnlnmi.

Comments. — The species and subspecies
of Hauffapis proposed by Armbruster (op.
cit.) were all described from the Miocene
of Randecker Maar and were based on mi-
nor wing variations. None of these varia-
tions justifies subspecific status even un-
der the extreme criteria used for splitting
morphs of extant species (e.g., A. mellifera).
The species is presently known from the
Miocene of Germany.

Subgenus Synapis Cockerell

Apis (Synnpis) Cockerell 1907: 229. Type species:
Apis {Syunpis) henshmvi Cockerell 1907,
monobasic. Zeuner and Manning 1976: 240.

Diagnosis. — WORKER: Basal vein gently
curved, only slightly distad cu-a (Fig. 6).
Angle of posteroapical margin of first sub-
marginal cell greater than 45°. Distal ab-
scissa of vein M in hind wing present (as
in Fig. 5). Drone, mature larva, queen, and
ethology all unknown.

Apis (Synapis) henshaivi Cockerell

Henshaw's Honey Bee

Apis (Synnpis) henshnwi Cockerell 1907: 229.
Apis donnitnus Cockerell 1907: 228. Preoccupied

(uec Heyden 1862).
Apis oligoceuicn Meunier 1915: 210.
Syuapis dorinitniis (Cockerell); Statz 1931: 45.
Synapis henshnzei (Cockerell); Statz 1931: 45.

188

Journal of Hymenoitera Research

Synapis kaschkei Statz 1931: 50.

Apis cuenoti Theobald 1937: 401. New synony-
my.

Apis oligocaenica Goetze 1964: 9. Lapsus calami.

Apis heiischnuii Goetze 1964: 9. Lnpsiis calami.

Apis knschki Goetze 1964: 9. Lapsus calami.

Apis henshawi dormiens Zeuner and Manning
1976: 241. Replacement name for Apis dor-
mitans Cockerell 1907.

Apis henshawi kaschkei (Statz); Zeuner and Man-
ning 1976: 243.

Apis aquisextana Nel, Martinez-Delclos, Arillo,
and Pefialver in Arillo et al. 1996: 60. Pro-
posed as new again in Nel et al. (1999).

Comments. — This species is perhaps the
most famous of the fossil honey bees. The
type was redescribed and new synony-
mies presented by Engel (1998a). This spe-
cies is from the Oligocene of Europe (in
Germany, France, and Spain).

Apis (.Synapis) lotigtibia Zhang

The Long-legged Honey Bee

Apis longtibia Zhang 1990: 85.

Comments. — Engel (1998a) gave some
characters to support the recognition of
this species, although far much work re-
mains to be done on the fossil honey bees
of Asia. This fossil is from the Miocene of
Shandong Province, China.

Apis (Synapis) miocenica Hong

The Chinese Miocene Honey Bee

Apis miocenica Hong 1983: 10.

Apisfota Zhang 1989: 323.

Apis shandongica Zhang 1989: 325.

Comments. — This species was briefly
treated by Engel (1998a) and reasons for
the synonymy of A.fota and A. shandongica
were given. The species is from the Mio-
cene of Shandong Province, China.

Apis (Synapis) petrefacta (Riha)

The Petrified Honey Bee

Sipiapis petrefacta Riha 1973: 217.

Apis petrefacta (Riha); Engel 1998a: 275.

strata of the Ceske Stredhori Mountains of
the Czech Republic.

Subgenus Priorapis Engel, new
subgenus

Type species. — Apis vetiista Engel 1998a.

Diagnosis.— WORKER: Basal vein
straight, confluent with cu-a. Angle of
posteroapical margin of first submarginal
cell greater than 45°. Distal abscissa of
vein M in hind wing present. Drone, ma-
ture larva, queen, and ethology all un-
known.

Etymologij. — The new subgeneric name
is derived from prior (L. earlier) and apis
(L. bee).

Apis (Priorapis) vetusta Engel,
emended name

The Aged Honey Bee

Apis vetustus Engel 1998a: 271.

Comments. — Through a lapse on my
part I failed to adjust the gender of the
specific epithet to match that of the genus
(which is feminine). I here correct this er-
ror. The species occurred in the Oligocene
of Germany.

NOMINA DUBIA AND
UNASSOCIATED NOMINA NUDA

Apis aenigmatica Rayment, nomen
dubiiim

Apis aenigmaticus Rayment 1925: 67.

Comments. — As pointed out by Cardale
(1993), this name was based on the draw-
ing of a comb and anecdotal reports of the
insect in the absence of any specimens.
Rayment (1935) gives another lively ac-
count of this elusive "species" which still
had not been found at that time (nor has
it been found since). See also discussion in
Michener (1965: 232).

Apis catanensis Roussy, nomen dubiutn

Apis catanensis Roussy 1960: 8.
Apis catanensis avolii Roussy 1960: 8.

Comments. — A fossil form from Miocene Comments. — ^Judging from the original

Volume 8, Number 2, 1999

189

description, which is exceedingly incom-
plete, this may not be a honey bee at all.
The subspecific name avolii is unnecessary
and invalid since no subspecific forms
were recognized; besides, any subspecies
based on the type specimen would have
to take the nominate specific epithet and
would therefore become A. catanensis ca-
tanensis, with A. cataiiensis avolii being an
invalid synonym. This is reportedly a fos-
sil species from Miocene Sicilian amber.

Apis melistiga Zeuner and Manning,
nomen dtibittm

"Apidae" melisuga Handlirsch 1907: 893. Un-
available (I.C.Z.N. 1985: Art. llh[iii]).
Apis melisuga Zeuner and Manning 1976: 248.

Comments. — The original proposal of
this epithet by the great paleoentomolo-
gist Anton Handlirsch (1907) was not in
accord with the principles of zoological
nomenclature and is therefore unavailable
under Article llh (iii) (I.C.Z.N. 1985). Zeu-
ner and Manning (1976) made the name
available through their monograph by
publishing this name is combination with
a genus-group name and a diagnosis; al-
though these authors attributed the name
to Handlirsch. Neither Prof. Zeuner nor
Dr. Manning had seen the type and their
diagnosis of the species was a mere re-
gurgitation of Handlirsch's original de-
scriptive comments that the specimen re-
sembled A. mellifera. In the absence of the
type (which is presumably lost) and any
real character information it is impossible
to confidently place this as a species of
Apis. The specimen was reportedly from
the Miocene of Italy. Refer to Engel
(1998a) for a more thorough treatment of
this name.

Apis postadamitica Buttel-Reepen,
nomen nudum

Apis postadamitica Buttel-Reepen 1906: 163.

Comments. — Buttel-Reepen (op. cit.) con-
sidered this as the hypothetical direct an-
cestor of A. mellifera and postulated its oc-

currence in the Pliocene. No specimen for
this species exists, and it was a mere hy-
pothesis of Buttel-Reepen's that this spe-
cies existed and would someday be dis-
covered. The fact that this name is for a
hypothetical taxon means that it is also ex-
cluded from zoological nomenclature
(I.C.Z.N. 1985: Art. lb).

Apis styriaca Pongracz, nomen nudum

Apis styriaca Pongracz 1931: 105.

Comments. — A supposed fossil species
from the Miocene of Germany represented
by a wing fragment. No description or fig-
ure was provided and the whereabouts of
this material is unknown.

Apis trigona Rayment, nomen nudum

Apis trigona Rayment 1925: 69. Preoccupied (nee
Schrank 1798).

Comments. — The decision to place this
as a nomen nudum was made by Cardale
(1993) and is followed here.

HONEY BEE VERSUS HONEYBEE

In closing I should like to make a brief
comment on common names for this
group of bees. The great arthropod mor-
phologist Robert E. Snodgrass presented a
short discussion on the common name for
honey bees in the preface to his work con-
cerning the anatomy of A. mellifera (Snod-
grass 1956). His brief discussion advocat-
ing the use of the two word common
name {honey bee) over a single word (bo)2-
eybee) does not appear to have been wide-
ly accepted since apiculturists still com-
monly use "honeybee." Although there
are no absolute rules for the use of this
and other common names, I agree with
Snodgrass' preference for a two word
name and the logic by which he justified
this position. Since I cannot hope to word
his position more eloquently, I quote here
his short argument: "Regardless of dictio-
naries, we have in entomology a rule for
insect common names that can be fol-
lowed. It says: If the insect is what the

190

Journal of Hymenoptera Research

name implies, write the two words sepa-
rately; otherwise run them together. Thus
we have such names as house fli/, blow fly,
and robber fly contrasted with dragonfly,
caddicefly, and butterfly, because the later
are not flies, just as an aphislion is not a
lion and a silverfish is not a fish. The honey
bee is an insect and is preeminently a bee;
'honeybee' is equivalent to 'Johnsmith.'"

ACKNOWLEDGMENTS

I am sincerely grateful to the librarians of Cornell
University's Comstock Memorial Library and the
Kroch Rare and Manuscript Collection and of the li-
brary of the American Museum of Natural History
for their help in locating references examined in the
course of this study. Kumar and Valerie Krishna
helped locate several older references and provided
delightful company during these searches. Numerous
bee biologists and paleontologists throughout the
world graciously supplied copies of their work and
shared discussions on Apis classification; to each of
them I extend my thanks. David A. Grimaldi, E. Eric
Grissell, Charles D. Michener, Card W. Otis, Wojciech
Pulawski, Molly G. Rightmyer, and Jerome G. Rozen,
Jr., kindly read versions of the manuscript and made
valuable corrections and criticisms. Their assistance
greatly improved the presentation of this material.
Not all agreed with my classification or conclusions
and any errors or idiosyncrasies which remain are, of
course, my own. 1 am particularly thankful to Mich-
ener for discussions on this material. I owe additional
thanks to Jeffrey G. Engel for assistance during the
preparation of the manuscript and to Molly G. Right-
myer and Zhiwei Liu for sharing with me their
knowledge of Mandarin in the construction of the ep-
ithet heimifeng. I am indebted to numerous institu-
tions and individuals for hosting me during my trav-
els and /or for the loan or donation of material; in
this regard 1 should particularlv recognize Robert W.
Brooks, Gabriela Chavarria, E. Richard Hoebeke,
Card W. Otis, W. Steve Sheppard, and Deborah R.
Smith. W. Steve Sheppard and his coauthors kindly
allowed me to include their lectotype designation for
A. mellifera ruttneri herein and I am grateful to them
for this honor. Donald B. Baker was the first to rec-
ognize the correct authorship of A. koschcviiikm'i (pers.
comm. to C. D. Michener); I am grateful for his advice
and assistance.

This work is dedicated in memory of Prof. Dr.
Friedrich Ruttner (1914-1998), leading authoritv on
Apis classification who did much to clarify honey bee
systematics (as well as contributing enormously to
many other fields of apiculture). A lovely account cel-
ebrating Prof. Ruttner's life was given by Koeniger
(1998).

LITERATURE CITED

Alexander, B. A. 1991a. A cladistic analysis of the ge-
nus Apis. Pages 1-28, in D. R. Smith (ed.) Diivr-
siti/ in the Genus Apis. Westview Press, Boulder,
Colorado, United States, xiv-l-265 pp.

Alexander, B. A. 1991b. A phylogenetic analysis of
the genus Apis (Hymenoptera: Apidae). Annals of
the Entomological Society of America 84: 137-149.

Alpatov, V. V. 1935. Contribution to the study of var-
iation in the honey bee. 3. The cubital cell on the
wings of different forms of the genus Apis and
its taxonomical and evolutionary significance.
Zoohgischeskiy Zhnrnal 14: 664-673. [In Russian
with English summary]

Alpatov, V. V. 1938. Contribution to the study of var-
iation in the honey bee. 4. Carnolian and Cri-
mean bees and their place among other forms of
Apis mellifera L. Zootogisctieskiy Zhurnal 17: 473-
481. [In Russian with English summary]

Anderson, R. H. 1963. The laying worker in the Cape
honeybee. Apis mellifera capensis. lounial of Api-
cultiiral Research 2: 85-92.

Arillo, A., A. Nel, and V. M. Ortuno. 1996. Two fossil
bees from the Oligocene of Izarra (Alava, Spain)
(Hymenoptera, Apoidea). Bulletin tie la Societe en-
tomologique de France 101: 59-64.

Armbruster, L. 1938. Versteinerte Horugbienen aus
dem obermiocanen Randecker Maar. Archiv fiir
Bienenkumte 19: l-t8, 97-133.

Ashmead, W. H. 1904. Remarks on honey bees. Pro-
ceciiings of the Entomological Society of Washington
6: 120-122.

Baldensperger, P. J. 1923. Sur I'abeille saharienne. 6"'
Congrcs International d' Apiculture, Marseille 1922:
61-64.

Baldensperger, P. J. 1928. Bees in their natural state
and with beekeepers. Bee World 9: 173-174.

Baldensperger, P. J. 1932. Varietes d'abeilles en Af-
rique du Nord. 5"' Congres International
d'Entomologie, Pans 1932: 829-839.

Barac, I. 1977. Preserving the genetic pool of Apis mel-
lifera carpatica. Proceedings of the International Bee-
keeping Congress 26: 270-274.

Barton, B. S. 1793. An inquiry into the question
whether the Apis nwUifica, or true Honey-bee, is
a native of America. Transactions of the Anurican
Philosophical Society, Philadelphia 3: 251-261.

Belknap, J. 1792. A Discourse Intended to Commemorate
the Discovery of America by Christopher Columbus;
Delivered at the Request of the Historical Society in
Massachusetts, on the 23d Day of October, 1792, be-
ing the Completion of the Third Century siiue that
Me\norable Event. To zvhich are added Four Disser-
tations, connected with various Parts of the Dis-
course. Apollo, Belknap & Hall, Boston, Massa-
chusetts, United States, 132 pp.

Benton, F. 1904. The specific name of the common

Volume 8, Number 2, 1999

191

honey-bee. Proceedings of Ihe Entoniolcigical Societi/
of Wasliinghm 6: 71-73.

Bingham, C. T. 1898. On some new species of Indian
Hymenoptera. journal of the Bombay Natural His-
tory Society 12: 115-130.

Brool<s, R. W., and C. D. Michener. 1988. The Apidae
of Madagascar and nests of Liotrigona (Hyme-
noptera). Sociobiology 14: 299-323.

Buttel-Reepen, H., von. 1906. Apistica. Beitrage zur
Systematik, Biologie, sowie zur geschichtlichen
und geographischen Verbreitung der Honigbiene
{Apiis mellificn L.), ihrer Varietaten und der iiber-
igen /Ijii.s-Arten. Mittcilungen aus licni Zoologiscli-
eii Museum in Berlin 3: 117-201.

Cardale, J. C. 1993. Hymenoptera: Apoidea. Pages 1-
405, in W. W. K. Houston and G. V. Maynard
(eds.) Zoological Catalogue of Australia, vol. 10.
Australian Government Publishing Service, Can-
berra, Australia, ix + 406 pp.

Chavarria, G., and J. M Carpenter. 1994. "Total evi-
dence" and the evolution of highly social bees.
Chulistics 10: 229-258.

Cockerel!, T. D. A. 1906. New Rocky Mountain bees,
and other notes. Canadian Entomologist 38: 160-
166.

Cockerell, T. D. A. 1907. A fossil honey-bee. Entomol-
ogist 40: 227-229.

Cockerell, T. D. A. 1911a. Descriptions and records of
bees, XXXV. Annals and Magazine of Natural His-
tory, series S, 7: 310-319.

Cockerell, T. D. A. 1911b. New and little-known bees.
Transactions of the American Entomological Society
37: 217-241.

Cockerell, T. D. A. 1914. Descriptions and records of
bees, LX. Annals and Magazine of Natural History,
series 8, 14: 1-13.

Dalla Torre, C. G., de [K. W., von). 1896. Catalogus
Hymenopterorum hucusque Descriptorum Systema-
ticus et Synonymicus. Vol. 10. Apidae (Anthophila).
Engelmann, Lipsiae [Leipzig], Germany,
viii-)-643 pp.

Damus, M. S., and G. W. Otis. 1997. A morphometric
analysis of Apis cerana F and Apis nigrocincta
Smith populations from Southeast Asia. Apidol-
ogie 28: 309-323.

Enderlein, G. 1906. Neue Honigbienen und Beitrage
zur Kenntnis der Verbreitung der Gattung Apis.
Stettiner Entonwlogische Zeitung 67: 331-344.

Engel, M. S. 1998a. Fossil honey bees and evolution
in the genus Apis (Hymenoptera: Apidae). Api-
dologie 29: 265-281.

Engel, M. S. 1998b. Comparative morphology and the
phylogenv of the corbiculate bees (Hymenoptera:
Apidae; Apinae). Proceedings of the 13"' Interna-
twnal Congress of ILISSI, Adelaide 1998: 151.

Engel, M. S., and T. R. Schultz. 1997. Phylogeny and
behavior in honev bees (Hymenoptera: Apidae).

Annals of the Entomological Society of America 90:
43-53.

Eschscholtz, J. F. 1822. Entomographien, vol. 1. Reimer,
Berlin, Germany, 59-186 pp.

Fabricius, ]. C. 1781. Species Insectorum Exhibentes cor-
am Differentias Specificas, Synonyma Auctorum,
Loca Natala, Metamorphosin adiectis Observationi-
bus, Descriptionibus, vol. 1. Bohn, Hamburgi et
Kilonii [Hamburg and Cologne], Germany,
yiii-l-552 pp.

Fabricius, J. C. 1787. Mantissa Insectorum Sistems cor-
am Species nuper Detectas adiectis Characteribus Ge-
nericus, Differentiis Specificis, Emendationihus, Ob-
scrvationibus, vol. 1. Proft, Hafniae [Copenhagen],
Denmark, xx + 348 pp.

Fabricius, J. C. 1793. Entomologia Systematica Emendata
et Aucta. Secundum Classes, Ordines, Genera, Spe-
cies adiectis Synonymis, Locis, Observationibus, Des-
criptionibus, vol. 2. Proft, Hafniae [Copenhagen],
Denmark, yiii-l-519 pp.

Fabricius, J. C. 1798. Supplcmcntum Entomologiae Sys-
tematicae. Proft, Hafniae [Copenhagen], Den-
mark, [2] + 572 pp.

Fischer von Waldheim, G. 1843. Observata quaedam
de Hymenopteris Rossicis. Magasin de Zoologie,
d' Anatomic comparee et de Paleontologie 5: 1-122.

Geoffroy, E. L. 1762. Histoire abregee des Insectes qui se
trouvent aux environs de Paris, 2 vols. Durand, Par-
is, France, vol. 1, 690 pp., 11 pis.; vol. 2, 744 pp,
12 pis.

Gerstacker, C. E. A. 1862. Uber die geographische Ver-
breitung und die Abiinderungen der Honigbiene
nebst Bcmerkungen itber die ausldndischen Honigbi-
enen der alien Welt. Festschrift XI Wander- Versa-
mmlung deutscher Bienenwirthe, Potsdam, Ger-
many, 75 pp.

Gerstacker, C. E. A. 1863. On the geographical distri-
bution and varieties of the honey-bee, with re-
marks upon the exotic honey-bees of the old
world. Annals and Magazine of Natural History, se-
ries 3, 11: 270-283, 333-347. [English translation
of Gerstacker 1862[

Goetze, G. K. L. 1964. Die Honigbiene in natiirlicher
und kiinstlicher Zuchtauslese, Teil 1: Systematik,
Zeugung und Vererbung. Monographien zur an-
geivandte Entomologie 19: 1-120.

Gorbachev, A. N. 1916. The gray mountain Caucasian
bee (Apis mellifera caucasica) and its place among
other bees. Tiflis 1916: 1^0. [In Russian with En-
glish summar^'l

Grassi, B. 1880. Saggio di una Monografia delle Api
d'ltalia. Milano [Milan[, Italy.

Grozdanic, S. S. 1926. Die "gelbe" banater Biene. Acta
Socictatis Entomologicae Serbo-Croato-Slovenae 1:
45-60.

Giierin-Meneville, F. E. 1844. Iconograp'hie du Regne
Animal de C. Cuvier, ou Representation d'apres Na-
ture de I'une des Especes les plus Renuirquables et

192

Journal of Hymenoptera Research

Scuvaiit noil Encore Figurees, de Chaque Genre
il'Anhnnux, vol. 3. Texte Explicatif, Insectes. J. B.
Bailliere, Paris, France, 576 pp., 106 pis.
Guillou, E. J. F., le. 1841. Des Insectes Hymenopteres
recueillis dans le Voyage de circumnavigation
des Corvettes 1' Astrolabe et la Zelee. Annales tie
ta Societe entonwlogique de France 10: 311-324.
Hadisoesilo, S. 1997. A Comparative Study of Two Spe-
cies of Cavity-Nesting Honey Bees of Sulazoesi, In-
donesia. Ph.D. Dissertation, University of Guelph,
Guelph, Ontario, Canada, 199 pp.
Hadisoesilo, S., and G. W. Otis. 1996. Drone flight
times confirm the species status of Apis nigro-
cincta Smith, 1861 to be a species distinct from
Apis cerana F, 1793, in Sulawesi, Indonesia. Api-
dologie 27: 361-369.
Hadisoesilo, S., and G. W. Otis. 1998. Differences in
drone cappings of Api^ cerana and Apis nigrocinc-
ta. journal of Apicultural Research 37: 11-15.
Hadisoesilo, S., G. W. Otis, and M. Meixner. 1995.
Two distinct populations of cavity-nesting honey
bees (Hymenoptera: Apidae) in South Sulawesi,
Indonesia, journal of the Kansas Entomological So-
ciety 68: 399-407.
Handlirsch, A. 1907. Die fossilen liisel<ten iind die Phy-
logenie der rezenten Forinen. Ein Handhiich fur Pa-
Idontologen uiid Zoologen. Engelmann, Leipzig,
Germany, 641-1120 pp., 37-51 pis.
Hepburn, H. R., and S. E. Radloff. 1998. Honex/hees of
Africa. Springer Verlag, Berlin, Germany, xv + 370
pp.
Hernandez, F. 1648. Rerum Medicaruin Novae Hispan-
iae Thesaurus: Nova Plantarnw, Animalium et Mi-
neralium Historia. Vitalis Mascardi, Romae
[Rome], Italy, 950+[10] + [90] + [6] pp.
Heyden, C, von. 1862. Gliedertiere aus der Braun-
kohle des Niederrheins, der Wetterau und der
Rhon. Palaeontogiaphica 10: 62-82.
Hoffmannsegg, J. C. G., von. 1818. Entomologische
Bemerkungen bei Gelegenlieit der Abhandlun-
gen uber amerikanische Insekten. Zoologischcs
Magazin, von Wiedemann 1(2): 49-110.
Hong, Y. C. 1983. Fossil insects in the diatoms of
Shanwang. Bulletin of the Tianjin Institute of Ge-
ology and Mineral Resources 8: 1-11. [In Chinese
with English summary]
Hong, Y. C. 1984. New fossil insects of Laiyang
Group from Laiyang Basin, Shandong Province.
Professional Papers of Stratigraphy and Paleontology
11: 31-41. [In Chinese with English summary)
Hong, Y. C, and S. ]. Miao. 1992. Fossil bee [sic] and
its origin with discussion on the origin of the an-
giosperm. Memoirs of the Beijing Natural History
Museum 51: 1-19. [In Chinese with English sum-
mary]
Home, C, and F. Smith. 1870. Notes on the habits of
some hymenopterous insects of the NW provinc-

es of India. Transactions of the Zoological Society of
London 7: 161-196.
I.e. Z.N. 1985. International Code of Zoological Nomen-
clature. Charlesworth & Co. Ltd., Huddersfield,
United Kingdom, xx + 338 pp.
Jack, R. W. 1916. Parthenogenesis amongst the work-
ers of the Cape honey-bee: Mr. G. W. Onions'
experiments. Transactions of the Entomological So-
ciety of London 1916: 396--103.
Jefferson, T. 1787. Notes on the State of Virginia. Stock-
dale, London, United Kingdom, 382 pp.
Josselyn, J. 1674. An Account of Two Voyages to New-
England Made During the Years 163S, 1663. Wid-
dows, London, United Kingdom, vii + 211 pp.
Kerr, W. E. 1957. lntrodu(;ao de abelhas africanas no

Brasil. Brasil Apicola 3: 211-213.
Kerr, W. E. 1967. The history of the introduction of
Africanized bees to Brazil. South African Bee Jour-
nal 39: 3-5.
Kerr, W. E., and E. Amaral. 1960. Apicultura: Cientifica
e Prdctica. Diretoria de Publicidade Agricola da
Secretaria da Agricultura, Sao Paulo, Brazil, 148
pp.
Kiesenwetter, E. A. H., von. 1860. Ueber die Bienen
des Hymettus. Berliner Entomologische Zeitschrift
4: 315-317.
Klug, J. C. F. 1807. Species Apiariarum familie novas
descripsit; generumque characteres adjecit. Ma-
gazin der Cesellschaft Naturforschender Frcunde zu
Berlin 1: 263-265.
Koeniger, N. 1998. Friedrich Ruttner, 15 May 1914-3

February 1998. Bee World 79: 110-111.
Koeniger, N., G. Koeniger, S. Tingek, M. Mardan, and
T. E. Rinderer. 1988. Reproductive isolation by
different time of drone flight between Apis cerana
Fabricius, 1793 and Apis vechti (Maa, 1953). Api-
dologie 19: 103-106.
Koschevnikov, G. A. 1900. Materials for the natural
history of the honey bee. Izviestiia Iinperatorskago
ohshchestva liubitelei estest-ooznaniia Antropologiy i
Etnografiy 99: 1-144. [In Russian]
Kugler, ]. 1988. The zoogeography of social insects of
Israel and Sinai. Pages 251-275, in Y. Yom-Tov
and E. Tchemov (eds.) The Zoogeography of Israel:
The Distribution and Abundance at a Zoogeograph-
ical Crossroad (Monographiae Biologicae, vol. 62).
Dr. W. Junk Publishers, Dordrecht, the Nether-
lands, ix + 600 pp.
Latreille, P. A. 1804a. Memoire sur un Gateau de
Ruche d'une Abeille des Grandes-Indos, et sur
les Differences des Abeilles proprement dites, vi-
vant en grande Societe, de I'ancien Continent et
du nouveau. Annales du Musee d'Histoire Naturellc
4: 383-394.
Latreille, P. A. 1804b. Notice des Especes d'Abeilles
vivant en grande Societe, ou Abeilles propre-
ment dites, et Description d'Especes nou\elles.
Annales du Musee d'Histoire Naturellc 5: 161-178.

Volume 8, Number 2, 1999

193

Latreille, P. A. 1810. Considerations Generates siir
I'Ordre Naturel des Animaux Composant les Classes
des Crustaces, des Arachnides, et des Insecies; avec
un Tableau Me'thodique de leurs Genres, disposes en
Families. F. Schoell, Paris, France, 444 pp.

Lavrezhin, F. A. 1935. Contribution to the study of
variation of the honey bee. 2. A comparative bio-
metric characteristic of the sexual appendages of
the drones belonging to different forms of the
genus Apis. Zoologicheskiy Zhurnal 14: 655-663.
|In Russian with English summary]

Lawrjochin, F. A. 1960. Uber die ersten Versuche zur
Einfiihrung der wildlebenden ussurischen Bi-
enen {Apis indica F.) in den europaischen Teil der
Sowjetunion. XVII Congresso Internazioimlc degli
Apicoltori. Roma. vol. 2 1950; 237-238.

Lepeletier de Saint Fargeau, A. L. M. 1836. Histoire
Nalurelle des Inscctes. Suites a Bujfon. Hymenopter-
es, vol. 1. Roref, Paris, France, 547 pp.

Linnaeus, C. 1758. Systema Naturae per Regna Tria Na-
turae, Secundum Classes, Ordines, Genera, Species,
cum Characteribus, Differentiis, Synonymis, Locis,
ed. 10, vol. 1 Refonnata. Salviae, Holmiae [Stock-
holm], Sweden, 824 pp.

Linnaeus, C. 1761. Fauna S'vecica, Sistens Animalia Sve-
ciae Regni: Mammalia, Aves, Amphibia, Pisces, In-
secta. Vermes, Distributa per Classes & Ordines,
Genera & Species, cum Differentiis Specierum, Syn-
onymis Auctorum, Nominibus Incolarum, Locis Na-
talium, Dcscriptionihus Insectorum. Salvii, Stock-
holmiae [Stockholm], Sweden, \lvii + 578 pp., 2
pis.

Linnaeus, C. 1767. Systema Naturae per Regna Tria Na-
turae, Secundum Classes, Ordines, Genera, Species,
cum Characteribus, Differentiis, Synonymis, Locis,
ed. 12, Reformata, vol. 1, pt. 2. Thomas, Vindobon-
ae [Vienna], Austria, 533-1328+ [xxxvi] pp.

Lord, W. G., and S. K. Nagi. 1987. Apis florea discov-
ered in Africa. Bee World 68: 39^0.

Lucas, H. 1882. Communique une note relative a un
Hymenoptere social. Bulletin des Seances de la So-
ciete entomologique de France 1882: 42.

Maa, T. C. 1944. On the classification and phylogeny
of the Chinese honeybees. Entomology SIxaowuana
1: 4-5. [In Chinese: this paper was not seen since
no copies of the journal could be traced. 1 follow
Maa's (1953) restatement of its contents.]

Maa, T. C. 1953. An inquiry into the svstematics of
the tribus Apidini or honeybees (Hym). Treubia
21: 525-640.

Malaipan, S. 1972. A Revision of the Genus o/Apis of
Thailand (Hymenoptera: Apidae). M.S. Thesis, Ka-
setsart University, Bangkok, Thailand, 139 pp.

McEvoy, M. V., and B. A. Underwood. 1988. The
drone and species status of the Himalavan honey
bee. Apis laboriosa (Hymenoptera: Apidae). jour-
nal of the Kansas Entomological Society 61: 246-249.

Meunier, M. F. 1915. Uber einige fossil Insekfen aus

den Braunkohlenschichten (Aquitanien) von Rott
(Siebengebirge). Zeitschrift der deutschen geolo-
gischen Gesellschaft 67: 205-217, 219-230.

Michener, C. D. 1944. Comparative external mor-
phology, phylogeny, and a classification of the
bees (Hymenoptera). Bulletin of the Anurican Mu-
seum of Natural History 82: 151-326.

Michener, C. D. 1965. A classification of the bees of
the Australian and South Pacific regions. Bulletin
of the American Museum of Natural History 130: 1-
362.

Michener, C. D. 1975. The Brazilian bee problem. An-
nual Review of Entoinology 20: 399^16.

Michener, C. D. 1990. Classification of the Apidae
(Hymenoptera). University of Kansas Science Bul-
letin 54: 75-153.

Michener, C. D. 1997. Genus-group names of bees
and supplemental family-group names. Scientific
Papers, Natural History Museum, University of Kan-
sas 1: 1-81.

Mogga, J. B. 1988. The Taxonomy and Geographical Var-
iability of the Honeybee Apis mellifera L. ni Sudan.
M.S. Thesis, University of Khartoum, Khartoum,
Sudan.

Mogga, J. B., and F. Ruttner. 1988. Apis florea in Af-
rica: Source of the founder population. Bee World
69: 100-103.

Montagano, J. 1911. Relation sur I'Apis sicida. Proceed-
ings of the International Beekeeping Congress 5: 26—
29.

Moore, F., F. Walker, and F. Smith. 1871. Descriptions
of some new insects collected by Dr. Anderson
during the expedition to Yunan. Proceedings of the
Zoological Society of London 1871: 244-249.

Morse, R. A., and F. M. Laigo. 1969. Apis dorsata in
the Philippines (including an annotated bibliog-
raphy). Monograph of the Plulippine Association of
Entomologists 1: 1-96.

Muttoo, R. N. 1951. The correct scientific nomencla-
ture for our Indian hive bees. Indian Bee Journal
13: 150-153.

Muttoo, R. N. 1956. Facts about beekeeping in India.
Bee World 37: 125-133, 154-157.

Nel, A., X. Martinez-Delclos, A. Arillo, and E. Pefi-
alver. 1999. A review of the Eurasian fossil spe-
cies of the bee Apis. Paleontology 42: 243-285.

Noll, F. B. 1998. Are behavioral characters informa-
tive of phylogenetic relationships among the cor-
biculate bees (Hymenoptera, Apidae, Apinae).
Proceedings of the 13"' International Congress oflUS-
SL Adelaide 1998: 347.

Otis, G. W. 1996. Distribution of recently recognized
species of honey bees (Hymenoptera: Apidae;
Apis). Journal of the Kansas Entomological Society,
supplement 69: 311-333.

Peng, Y. S., M. E. Nasr, and S. J. Locke. 1989. Geo-
graphical races of Apis cerana Fabricius in China
and their distribution. Review of recent Chinese

194

Journal of Hymenoptera Research

publications and a preliminary statistical analy-
sis. Apidologie 20: 9-20.
Petrov, P. 1991. Systematics of tlie Bulgarian bees.

Pchelnrstvo 9: 15-17. [In Russian]
Petrov, P. 1992. Distribution and phylogenesis of the
bee genus Apns (Hymenoptera, Apidae). Uspcdii
sovrememwi Bioh^gii/, Moscow 112: 359-372. [In
Russian]
Petrov, P. 1993. Man and the honey bee. Pchelovodstvo

11/12: 36-37. [In Bulgarian]
Petrov, P. 1996. Contribution to the taxonomy of the
Bulgarian honey bee. lounml of Animal Sdeiicc,
Agrkiiltiiral Acndcwy 33: 58-61. [In Bulgarian
with Russian and English summaries]
Petrov, P. 1997. On the origin of the Bulgarian honey
bees (Apis mellifica wdopka). journal of Animal Sci-
ence, Agricidtural Academy 34: 88-93.
PoUmann, A. 1879. Werth der verschiedenen Bienenraccn
und deren Varietdten, bestimmt durch Urtheile nam-
hafter Biencnziichtcr. Voigt, Leipzig, Germany, 69
pp.
PoUmann, A. 1889. Wcrtli der verschiedenen Bienenraccn
und deren Varietdten, bestimmt durch Urtheile nam-
hafter Bienenziichter. Voigt, Leipzig, Germany,
viii + 100 pp.
Pongracz, A. 1931. Bemerkungen uber die Insekten-
fauna von Oeningen nebst Revision der
Heer'schen Typen. Verhandlungen des Naturhis-
torisch-medizinisdwn Vereins zu Heidelberg 17: 104-
125.
Radloff, S. E., and H. R. Hepburn. 1997. Multivariate
analysis of honeybees. Apis mellifera Linnaeus
(Hymenoptera: Apidae), of the Horn of Africa.
African Entomology 5: 57-64.
Radoszkowski, O. 1887. Hymenopteres de Koree. Ho-
me Societatis Enfomologicae Rossicae 21: 428-436.
Rafinesque, C. S. 1814. Principes Fondamentaux de So-
miologie. Impr. de F. Abate aux depens de
I'auteur, Palerme [Palermo], Italy, 50-1- [2] pp.
Rafinesque, C. S. 1815. Analyse de la Nature: ou Tableau
de I'Univers et des corfs Organises. Impr. de F.
Abate aux depens de I'auteur, Palerme [Paler-
mo], Italy, 224 pp.
Ransome, H. M. 1937. The Sacred Bee in Ancient Times
and Folklore. Hougton Mifflin Company, New
York, United States, 308 pp.
Rashad, S. E., and M. S. A. El-Sarrag. 1978. Beekeep-
ing in Sudan. Bee World 59: 105-111.
Ray, J. 1710. Historia Insectorum. A. and J. Churchill,

London, United Kingdom, xv-l-400 pp.
Rayment, T. 1925. A new species of honey-bee. Aus-
tralasian Beekeeper 27: 67-69.
Rayment, T. 1935. A Cluster of Bees. Sixty Essays on the
Life-Histories of Australian Bees, with Specific De-
scriptions of over 100 New Species, and an Introduc-
tion by Professor £. F. Phillips. Endeavour Press,
Sydney, Australia, 752 pp.
Riha, P. 1973. Synapis petrefacta sp. n., cine neue Bi-

enenart aus dem Tertiar des B5hmischen Mittel-
gebirges. Vi'stnik Usti'ednilio listavu geologickeho
48: 217-220.
Roubik, D. W., S. F. Sakagami, and I. Kudo. 1985. A
note on distribution and nesting of the Himala-
yan honey bee Apis laboriosa Smith (Hymenop-
tera: Apidae). journal of the Kansas Entomological
Society 58: 746-749.
Roussy, L. 1960. Insectes et abeilles fossiles de I'ambre
de Sicile. Migrations, localisations, peuplement
du Nouveau Monde, de I'Australie, de le Nou-
velle Zelande. La Gazette Apicole, Montfavet 635:
5-8.
Ruttner, F. 1968. Les races d'abeilles. Pages 27-14, in
R. Chauvin (ed.) Traite' de Biologic de I'Abeille. vol.
1. Masson et Cie, Paris, France, xii^-547 pp.
Ruttner, F. 1975. Die kretische Biene, Apis mellifica
adami. Allgemeine deutsche Imkerzcitimg 9: 271-
272.
Ruttner, F. 1976a. Les races d'abeilles de I'afrique.
XXV Ccngri'S International d'Apicnlture, Grenoble
1976: 347-367.
Ruttner, F. 1976b. Isolated populations of honeybees
in Australia. Journal of Apicultural Research 15: 97-
104.
Ruttner, F. 1977. The problem of the Cape bee {Apis
mellifera capensis Escholtz ]sic]): parthenogenesis,
size of population, evolution. Apidologie 8: 281-
294.
Ruttner, F. 1980. Apis mellifera adami (n. ssp.), die Kre-
tische Biene. Apidologie 11: 385-400.
Ruttner, F. 1988. Biogeography and Ta.xonomy of Hon-
tM/('f«. Springer Verlag, Berlin, Germany,
xxii + 284 pp.
Ruttner, F. 1992. Naturgeschichte der Honigbienen. Eh-

renwirth, Munich, Germany, 357 pp.
Ruttner, F., D. Kauhausen, and N. Koeniger. 1989. Po-
sition of the red honey bee. Apis koschevnikovi
(Buttel-Reepen 1906), within the genus Apis. Ap-
idologie 20: 395-404.
Sakagami S. F., T. Matsumura, and K. Ito. 1980. .4;'/.-;
laboriosa in Himalaya, the little known world
largest honeybee (Hymenoptera: Apidae). Insecta
Matsnmurana 19: 47-77.
Santiago, E., J. Albornoz, A. Dominguez, and J. I. In-
quierdo. 1986. Etude biometrique des popula-
tions d'abeilles (Apis mellifera) du nord-ouest de
I'Espagne. Apidologie 16: 71-92.
Schrank, F., von Paula. 1798. Fauna Boica: Durehge-
dachte Geschichte der in Baiern einlieimischen und
zahmen Thiere. Stein'schen Buchhandlung, Nurn-
berg, Germany.
Schultz, T. R., M. S. Engel, and M. Prentice. 1999. Re-
solving conflict between morphological and mo-
lecular evidence for the origin of eusociality in
the "corbiculate" bees (Hymenoptera: Apidae):
A hypothesis-testing approach. The University of

Volume 8, Number 2, 1999

195

Kansas Natural Histori/ Museum Special Publication
24: 110-123.

Scopoli, J. A. 1770. Annus Historicc Naturalis. Annus
IV. Hilscher, Lipsiae [Leipzig], Germany, 150 pp.,
2 pis.

Seeley, T. D. 1985. Honeybee Ecology: A Study of Ad-
aptation in Social Life. Princeton University Press,
Princeton, New Jersey, United States, x + 201 pp.

Seeley, T. D. 1995. The Wisdom of the Hive: The Social
Physiology of Honey Bee Colonies. Harvard Univer-
sity Press, Cambridge, Massachusetts, United
States, xiv+295 pp.

Sheppard, W. S., M. C. Arias, A. Grech, and M. D.
Meixner. 1997 [1998]. Apis mellifera ruttneri, a
new honey bee subspecies from Malta. Apidologie
28: 287-293.

Skorikov, A. S. 1929a. Beitrage zur Kenntnis der kau-
kasischen Honigbienenrassen, 1-V. Rcp'orts on Ap-
plied Entomology, Leningrad 4: 1-60. [In Russian
with German summary]

Skorikov, A. S. 1929b. Eine neue Basis fiir eine Revi-
sion der Gattung Apis L. Reports on Applied En-
tomology, Leningrad 4: 249-270. [In Russian with
German summary]

Smith, D. R. 1991a. Diversity in the Genus Apis. West-
view Press, Boulder, Colorado, United States,
xiv-(-265 pp.

Smith, D. R. 1991b. Mitochondrial DNA and honev
bee biogeography. Pages 131-176, in D. R. Smith
(ed.) Diversity in the Genus Apis. Westview Press,
Boulder, Colorado, United States, xiv-l-265 pp.

Smith, D. R., and T. C. Glenn. 1995. Allozyme poly-
morphisms in Spanish honeybees {Apis mellifera
iberica). journal of Heredity 86: 12-16.

Smith, D. R., and R. H. Hagen. 1996. The biogeogra-
phy of Apis cerana as revealed by mitochondrial
DNA sequence data. Journal of the Kansas Ento-
mological Society, supplement 69: 294-310.

Smith, D. R., M. F. Palopoli, B. R. Taylor, J. M. Cor-
nuet, L. Garnery, M. Solignac, and W. M. Brown.
1991. Geographical overlap of two mitochondrial
genomes in the Spanish honey bees. Apis mellifera
iberica. Journal of Heredity 82: 96-100.

Smith, F. 1854. Catalogue of Hymeiiopterous Insects in
the Collection of the British Museum, Part 2, Apidae.
British Museum, London, United Kingdom, 199-
465 pp., vii-xii pis.

Smith, F. 1858. Catalogue of the hymenopterous in-
sects collected at Sarawak, Borneo; Mount Ophir,
Malacca; and at Singapore, by A. R. Wallace. Pro-
ceedings of the Linnean Society. London 2: 42-130.

Smith, F. 1859. Catalogue of hymenopterous insects
collected at Celebes by Mr. A. R. Wallace. Pro-
ceedings of the Linnean Societi/, London 3: 4-27.

Smith, F. 1861a. Descriptions of new species of hy-
menopterous insects collected by Mr. A. R. Wal-
lace at Celebes. Proceedings of the Linnean Society,
London 5: 57-93.

Smith, F. 1861b. [Untitled]. Proceedings of the Entomo-
logical Society of London 7: 14.

Smith, F. 1865. On the species and varieties of the
honey-bees belonging to the genus Apis. Annals
and Magazine of Natural History, series 3, 15: 372-
380.

Smith, F. 1871. A catalogue of the aculeate Hvmenop-
tcra and Ichneumonidae of India and the eastern
Archipelago. Journal of the Linnean Society 11:
285-415.

Smith, F. G. 1961a. The races of honeybees in Africa.
Bee World 42: 255-260.

Smith, F. G. 1961b. Races of honeybees in Africa. 18"'
Congreso Internacional de Apicnitura, Madrid 1961:
148.

Snodgrass, R. E. 1956. Anatomy of the Honey Bee. Cor-
nell University Press, Ithaca, New York, United
States, xiv+334 pp.

Spinola, M. 1806. Insectorum Liguriae Species Novae aut
Rariores, Quas in Agro Ligustico nuper Detexit, Des-
cripsit. el Iconibus Ulustravit Maxiinilianus Spinola,
adjecto Catalogo Specierum Auctoribus jam Enumer-
atarum, Quae in Eadem Regione Passim Occurrunt,
vol. 1. Gravier, Genuae [Genoa], Italy,
xvii + 159 + [l] pp., 2 pis.

Spivak, M., D. J. C. Fletcher, and M. D. Breed. 1991.
The "African" Honey Bee. Westview Press, Boul-
der, Colorado, United States, [ii]+435 pp.

Starr, C. K., P. J. Schmidt, and J. O. Schmidt. 1987.
Nest-site preferences of the giant honev bee. Apis
dorsata (Hymenoptera: Apidae), in Borneo. Pan-
Pacific Entomologist 63: 37-42.

Statz, G. 1931. Eine neue Bienenart aus Rott am Sie-
bengebirge. Ein Beitrag zur Kenntnis der fossilen
Honigbienen. Wissenschaftlichen Mitteduiigen des
Vereins fiir Natur- iind Hetmatkunde, Koln 1: 39-
60.

Taylor, O. R., Jr. 1977. The past and possible future
spread of Africanized honeybees in the Ameri-
cas. Bee World 58: 19-30.

Taylor, O. R., Jr. 1985. African bees: Potential impact
in the United States. Bulletin of the Entomological
Society of America 31: 14-24.

Tegetmeier, W. B. 1859. [Untitled|. Proceedings of the
Entomological Society of London 5: 88.

Tegetmeier, W. B. 1860. [Untitled]. Proceedings of the
Entomological Society of London 6: 126.

Theobald, N. 1937. Les Insectes Fossiles des Terrains Ol-
igocenes de France. George Thomas, Nancy,
France, 473 pp., 29 pis.

Tingek, S., M. Mardan, T. E. Rinderer, N. Koeniger,
and G. Koeniger. 1988. Rediscovery o( Apis vechti
(Maa, 1953): The Saban honey bee. Apidologie 19:
97-102.

Tingek, S., G. Koeniger, and N. Koeniger. 1996 []997|.
Description of a new cavity nesting species of
Apis (Apis nulueiisis n. sp.) from Sabah, Borneo,

196

Journal of HYMENOrrERA Research

with notes on its occurrence and reproductive bi-
ology. Senckenbergiana biologicn 7b: 115-119.

Underwood, B. A. 1990a. Seasonal nesting cycle and
migration patterns of the Himalayan honey bee
Apis lahoriosn. Nnliounl GcogrnpJiic Reseiirch 6: 276-
290.

Underwood, B. A. 1990b. Time ot drone flight of Aph
lahoriosa Smith in Nepal. Apiciolcgie 21: 501-504.

Villers, C, de. 1789. Cawli Linnaei Entomologia, Faunae
Suecicae Descriptionibus Aucta, D.D. Scopoli, Geof-
frey, de Geer, Fabricii, Schrank, &c., Speciebus vel in
Systemate non Enumeratis, vel Nuperrimc Detectis,
vel Speciebuf^ Galliae Australis Locupletata, Generii
Specieriinupie Rariorum Iconibm Ornata; Curante et
Augeiite Carolci de Villers, vol. 3. Piestre et Dela-
molliere, Lugduni [Lyon], France, 657 pp., 4 pis.

Willis, L. G., M. L. Winston, and B. M. Honda. 1992.
Phylogenetic relationships in the honeybee (ge-
nus Apis) as determined by the sequence of the
cytochrome oxidase II region of mitochondrial
DNA. Molecular Phylogenetics and Evolution 1:
169-178.

Wilson, E. O., and W. L. Brown, Jr. 1953. The sub-
species concept and its taxonomic application.
Systematic Zoology 2: 97-111.

Winston, M. L. 1987. The Biology of the Honey Bee. Har-
vard University Press, Cambridge, Massachu-
setts, United States, viii+281 pp.

Wongsiri, S., K. Limbipichai, P. Tanganasing, M.
Mardan, T. E. Rinderer, H. A. Sylvester, G. Ko-
eniger, and G. W. Otis. 1989. Evidence of repro-
ductive isolation confirms that Apis andreniforniis
(Smith, 1858) is a separate species from sympat-
ric Apis florea (Fabricius, 1787). Apidologie 21: 47-
52.

Wu, Y., and B. Kuang. 1986. A study of the genus
Micrapis (Apidae). Zoological Researcli 7: 99-102.
[In Chinese]

Wu, Y., and B. Kuang. 1987. Two species of small
honeybee — a study of the genus Micrapis. Bee
World 68: 153-155.

Zeuner, F. E. 1931. Die hisektenfauna des Bottinger
Marmors. Fortschritte der Geologic und Palaeonto-
logie 9: 247--106.

Zeuner, F. E., and F. J. Manning. 1976. A monograph
on fossil bees (Hymenoptera: Apoidea). Bulletin
of the British Museum (Natural History). Geology
27: 149-268.

Zhang, F. J. 1989. FossiV Insects from Shamoang, Shan-
dong, China. Science and Technology Publishing
House, Jinan, China, 459 pp., 92 pis. [In Chinese
with English summary]

Zhang, F. J. 1990. New fossil species of Apoidea (In-
secta: Hymenoptera). Act.i Zootaxonomica Sinica
15: 83-91. [In Chinese with English summarv]

J. HYM. RES.
Vol. 8(2), 1999, pp. 197-203

Ultrastructure of Spermatozoa in Leptopilina
(Hymenoptera: Cynipoidea: Eucoilidae)

Terence M. Newman and Donald L. J. Quicke

(TMM) Unit of Parasitoid Systematics, CABI Bioscience UK Centre (Ascot), Department of

Biology, Imperial College at Silwood Park, Ascot, Berkshire SL5 7PY, UK;

(DLJQ) Unit of Parasitoid Systematics, CABI Bioscience UK Centre (Ascot),

Department of Biology, Imperial College at Silwood Park, Ascot, Berkshire SL5 7PY, UK and

Department of Entomology, The Natural History Museum, London SW7 5BD, UK

Abstract. — The mature spermatozoa of the eucoilid wasp, Leptopilina heterotoma (Thomson), is
characterised by a solid corkscrew-shaped nucleus with a posteriorly directed flange that is
unique, not only among other reported Hymenoptera, but among all insect sperm previously
investigated. Leptopilina spermatozoa are further characterised by possession a complex acrosomal
structure, asymmetric mitochondrial derivatives and a centriolar adjunct interposed between the
smaller mitochondrial derivative and the nucleus. Because of their uniqueness, Leptopilina sperm
offer little insight into the relationship between the eucoilids and other members of the Cynipo-
idea, or the relationships of the Cyrupoidea to the rest of the Hymenoptera.

The Hymenoptera is one of the largest
orders of insects, but ciespite this there
have been remarkably ie-w studies of hy-
menopteran sperm ultrastructure, and
particularly among the parasitic taxa con-
sidering their diversity and species rich-
ness (Baccetti 1972; Jamieson 1987; New-
man and Quicke, in press). The sperm of
some social taxa such as ants and bees
(Hoage and Kessel 1968; Lensky et al.
1979; Thompson and Blum 1967; Wheeler
et al. 1990; Dallai and Afzelius 1990), and
a few Chalcidoidea among the 'Parasitica'
(Wilkes and Lee 1965; Lingmei and Dunsu
1987) are relatively well described, but
there are still a number of superfamilies
and many families for which there is very
little spermatological information. Sperm
structure in the superfamily Cynipoidea,
for example, is known only through one
investigation of a member of the family
Figitidae (Quicke et al. 1992). The relation-
ship of the Cynipoidea to the other apo-
critan (wasp-waisted) Hymenoptera is not
well understood. They have been consid-
ered as being related to the Chalcidoidea

(Konigsmann 1977), but also to Diapriidae
(Proctotrupoidea sensu lata) (Rasnitsyn
1988; Ronquist 1994, 1995) and to Ropron-
iidae + Vanhorniidae (Proctotrupoidea sen-
su stricto) (Dowton and Austin 1994).
Greater knowledge of the sperm structure
of other cynipoids may therefore shed
new light on the relationships of this su-
per-family to the other Hymenoptera.

The present paper deals with the sperm
of another member of the superfamily Cy-
nipoidea, Leptopilina heterotoma (Thomson)
(Quicke 1997), a koinobiont endoparasi-
toid of Drosophila larvae. This is the first
member of its family, the Eucoilidae,
whose sperm morphology has been inves-
tigated. It has a solid-corkscrew nucleus,
that is not only different to that of the only
other cynipoid so far described, but is un-
like any other Hymenopteran sperm pre-
viously reported.

MATERIAL AND METHODS

Testes and vas deferens from freshly
eclosed adult males of Leptopilina heteroto-
ma, kept in culture on their host {Drosophi-

198

Journal of Hymenoptera Research

t vi^^

Fig. 1. Low power view of sectioned vas defferens of Leptci\)Hina hetcrotoma with sections through acrosome
(A), nucleus (N) and tail piece (T) including some with only one mitochondrial derivative (arrowed) (scale
bar = 1 jjLm).

la melariogaster) were studied. Preparation
was based on the protocol used by New-
man and Quicke (1998). The tissues were
dissected under 2% glutaraldehyde in
phosphate buffered saline (0.1 M; pH 7.2),
fixed for two hours, then transferred to 2%
osmium tetroxide in cacodylate buffer (0.1
M; pH 7.2) for a further 2 hr fixation. After
two buffer washes, tissue was dehydrated
to 50% ethanol and then contrasted with a
saturated solution of uranyl acetate in 50%
ethanol prior to complete dehydration,
embedding in Epon resin and polymeri-
sation overnight. Large silver sections
were picked up on high resolution grids
and contrasted with uranyl acetate and
lead citrate.

RESULTS

The mature sperm of Lqjtopiliua hetcro-
toma exhibit many of the structures de-
scribed for other parasitic wasps, e.g. cris-
tate mitochondrial derivatives, deltoid
bodies and an axoneme with 9-1-9-1-2 ar-
rangement of microtubules (Fig. 1). How-
ever, they also have several features dif-
ferent from previously reported examples.
The acrosome is a much more complicated
structure than any yet described, forming
part of an acrosomal complex (Fig. 2 ar-
roic). Posteriorly the acrosomal rod (per-
foratorium) lies to one side of the nucleus
and is not held tightly there. The anterior
portion of the acrosomal rod protrudes

Figs. 2-5. Features of Leptopilina heterotoma sperm: 2, longitudinal section with acrosomal complex surround-
ed by extracellular cap (arrowed) and showing alternating protuberances from the nucleus which are the
result of sectioning through the spiral ridge (scale bar = 1 (xm); 3, high power oblique section through

Volume 8, Number 2, 1999

199

acrosomal complex illustrating the extracellular cap, acrosomal rod, inner sheath (I), granular mass (G), gran-
ular extracellular cap surrounding anterior part of nucleus (curved arrow, left hand section) [note also more
posterior section to right of middle which still has cytoplasm indicative of immature sperm] (scale bar = 100
(im); 4, transverse sections through nucleus (N) showing (lower right section) comma-shaped profile charac-
teristic of anterior and medial part, and (left section) round profile towards posterior; acrosomal com-
plex(middle two sections), showing asymmetric electron dense inner sheath of acrosome (I) and acrosomal
rod (A) (scale bar = 100 jim); 5, transverse section through acrosomal complex at level of acrosomal rod,
illustrating that the inner sheath does not completely enclose the granular material of the sub-acrosomal space,
and showing denser structure of extracellular cap (C) adjacent to inner sheath (scale bar = 100 ^.m).

200

Journal of Hymenoitera Research

into a mass of fine fibrous material which
partially fills the sub-acrosomal space
(Figs. 2-5). The true acrosome, that is the
portion derived from the acrosomal vesi-
cle, has a conical shape. This forms a
membrane-bound inner sheath below the
true cell plasma membrane (Fig. 3). The
acrosomal contents are asymmetrically
distributed around the fibrous material.
Thus, posteriorly, in section, (Fig. 4, up'per
profile) there is almost a complete ring of
material, whereas more anteriorly (Fig. 4,
middle profile), on one side of the acroso-
mal complex the acrosomal contents are
absent and only the membrane is found.
Surrounding this acrosomal complex is an
extracellular cap (Figs. 2 and 5). In longi-
tudinal section this cap covers, and com-
pletely encloses, the anterior of the nucle-
us (Figs. 2, 3 curved arrozv).

It is in the structure of the nucleus that
the main difference between the sperm of
Leptopiliua and those of other hymenop-
terans studied to date is found. In most
transverse nuclear sections, there is a pro-
tuberance, giving the nucleus a comma-
shaped profile (Figs. 1, 3, 4). In longitudi-
nal section, these protuberances alternate
along either side of the nucleus for most
of its length (Fig. 2). Posteriorly the nucle-
us abuts the axoneme at the area of the
basal body (Fig. 6), where the nucleus
overlaps the larger of two mitochondrial
derivatives for a short distance (Fig. 6 open
arrow). In transverse section (Fig. 7, arroiv)
an electron dense line is found that may
be a continuation of a similar structure
found at the interface of the axoneme and
the nucleus. Next to the axoneme, and
also abutting the nucleus, is a large elec-
tron-dense centriolar adjunct. (Fig. 8, open
arrow). Posteriorly, this centriolar adjunct
contacts the smaller of the two mitochon-

drial derivatives (Fig. 9). The positiorung
of these structures is such that at the re-
gion of overlap it is possible to obtain
oblique sections with centriolar adjunct,
mitochondrial derivative and nucleus all
present (Fig. 10). Closely apposed to the
mitochondrial derivatives are the two del-
toid bodies (also called triangular rods;
Lensky et al. 1979). Although membra-
nous, their derivation is not clear, but they
are probably associated with the mito-
chondrial derivatives because they are
also of different sizes (Fig. 11, arrows). This
offsetting of the mitochondrial derivatives
probably results in the tail piece of the
wasp spermatozoon only containing a sin-
gle (small) mitochondrial derivative.

Observations of living sperm expressed
from the vas deferens into saline showed
(X 400; phase contrast) that the nucleus is
a straight, rigid-appearing structure which
appears to rotate around its long axis as
because waves that we believe correspond
to the nuclear ridges described here can be
seen passing posteriorly along them.

DISCUSSION

The most probable interpretation of the
nuclear structure, as derived from longi-
tudinal and transverse sections, is a cyl-
inder with a posteriorly-directed ridge
spiralling down along its surface. This
therefore resembles a solid-cored cork-
screw. Observations of living sperm indi-
cate that the profile of the sperm head
does follow the nuclear cork-screw, and
that this shape may serve some fluid dy-
namic function, perhaps inducing the ob-
served rotatory motion. The spiral ridge
may therefore have significance for the
fertilisation process.

Spirally formed sperm are known in
two other groups of Hymenoptera, an un-

Figs. 6-11. Features of of Leptofnlina hetewioma sperm: 6, large mitochondrial derivative (open arrow) over-
laps the nucleus (N) vi'hich abuts the axoneme at the level of the basal body (B) (scale bar = 100 fim); 7,
transverse section showing overlap of large mitochondrial derivative and nucleus (N), with a electron dense

Volume 8, Number 2, 1999

201

area separating the two (scale bar = 100 (j.m); 8, centriolar adjunct (open arrow) abutting nucleus (N) anteriorly
(scale bar = 100 jjim); 9, posterior portion of the centriolar adjunct (CA) abuts the smaller mitochondrial
derivative (scale bar = 100 ^.m); 10, showing close proximity of the nucleus (N) mitochondrial derivative (M)
and centriolar adjunct (CA) which means that in oblique sections it is possible to have all three in the same
plane (scale bar = 100 ji-m); 11, tail piece showing that the larger mitochondrial derivative has an associated
large deltoid body (large arrow), whereas the smaller mitochondrial derivative has a correspondingly smaller
deltoid bodv (small arrow) (scale bar = 100 \xm).

202

Journal of Hymenoptera Research

identified member of the Diapriidae
which belongs to the Proctotrupoidea sen-
su lato (Quicke et al. 1992) and some but
not all chalcidoids (Lee and Wilkes 1965;
Hogge and King 1975; Quicke 1997). How-
ever, the sperm of the only other cynipoid
studied to date, Figites sp. (Cynipoidea:
Figitidae), shows no sign of spiralling
(Quicke et al. 1992). In all the other hy-
menopterans with spiral sperm structure,
the spiralling includes the axoneme and
mitochondrial derivatives as well as the
nucleus, and there is no protuberance
from the nucleus itself which has a normal
circular profile in cross section. The spi-
ralling in wasps other than Leptopilina can
be better likened to that of an open cork-
screw. This phylogenetic distribution of
spiralling suggests that this feature is
quite homoplastic, but until a robust phy-
logeny for the Chalcidoidea is available,
and more taxa have been investigated
spermatologically, it is not possible to say
how many times the open corkscrew type
spiralling has evolved or been lost. The
solid corkscrew form of nucleus with a
protruding ridge or keel has not been ob-
served in any other insect, but a quite sim-
ilar arrangement is found in all Chilopo-
da, and in the Onychophora and some ol-
igochaetes (Jamieson 1987). In these, the
keel is not so well developed as in Lepto-
pilina and in some the nucleus itself ad-
ditionally forms a more or less open cork-
screw.

Most of the spermatozoa observed in
the sections through the vas deferens were
mature, but a few still had a membrane-
bound layer of granular cytoplasm sur-
rounding them (Fig. 3) probably indicat-
ing that these were still at a late stage of
spermiogenesis, implying that not all
sperm are mature in the seminal vesicle
and vas deferens at least of recently
eclosed imaginal wasps.

ACKNOWLEDGMENTS

The authors gratefully acknowledge the cheerful
assistance of Mr Ian Morris (EM Unit, Department of

Biology, Imperial College). We thank Lex Kraaijeveld
for supplying the wasps and Robert Belshaw for com-
ments on an earlier draft. This research was support-
ed by the NERC (Natural Environment Research
Council), Initiative in Taxonomy and a grant from the
Leverhulme Trust to DLJQ and Mike Fitton (The Nat-
ural History Museum).

LITERATURE CITED

Baccetti, B. 1972. Insect sperm cells. Aclviiiiccs in Insect
Pln/siolog}/ 9:315-397.

Dallai, R. and B. A. Afzelius. 1990. Microtubular di-
versity in insect spermatozoa: Results obtained
with a new fixative, journal of Structural Biolog}/
103:164-169.

Dowton, M. and A. D. Austin. 1994. Molecular phy-
logeny of the insect order Hymenoptera: Apocri-
tan relationships. Proccetliugs of the NatiounI Acaci-
cmy of Sciences. USA, 91:9911-9915.

Hoage, T. R. and R. G. Kessel. 1968. An electron mi-
croscopical study of the process of differentiation
during spermatogenesis in the drone honey bee
with special reference to replication and elimi-
nation, journal of Ultrastructural Research 24:6-32.

Hogge, M. A. F. and P. E. King. 1975. The ultrastruc-
ture of spermatogenesis in Nasonia vitripennis
(Walker) (Hymenoptera: Pteromalidae). journal of
Subniicroscopncal Cytology 7:81-996.

Jamieson, B. G. M. 1987. The Ultrastructure and Phy-
logeni/ of Insect Spermatozoa. Cambridge Univer-
sity Press, Cambridge, 320 pp.

Konigsmann, E. 1977. Das phylogenetische System
der Hymenoptera: Teil 2: Svmphyta). Dcntsches
Entoniologische Zeitschrift 24:1-40.

Lee, P. E. and A. Wilkes. 1965. Polymorphic sper-
matozoa in the hvmenopterous wasp Dahlbom-
inus. Science, 147:1445-1446.

Lensky, Y., E. Ben-David, and H. Schindler. 1979. Ul-
trastructure of the spermatozoan of the mature
drone honeybee. Journal of Apicultural Research
18:264-271. '

Lingmei, D. and W. Dunsu. 1987. Ultrastructural ob-
servations on the spermatozoa of Trichogramma.
Acta Zoologica Siuica 33:262-266.

Newman, T. M. and D. L. J. Quicke. 1998. Sperm de-
velopment in the imaginal testes of Aleknies cox-
alis (Hymenoptera: Braconidae: Rogadinae). jour-
nal of Hymenoptera Research 7:25-37.

Newman, T. M. and D. L. J. Quicke. 1999. Ultrastruc-
ture of imaginal spermatozoa of sawflies (Insec-
ta: Hymenoptera: Svmphyta). journal of Hyme-
noptera Researcli 8:35-47.

Quicke, D. L. J. 1997. Parasitic Wasps. Chapm.m and
Hall, London, 470 pp.

Quicke, D. L. J., S. N. Ingram, H. S. Baillie and P. V.
Gaitens. 1992. Sperm structure and ultrastructure
in the Hymenoptera (Insecta). Zoologica Scripta
21:381-402.

Volume 8, Number 2, 1999

203

Rasnitsyn, A. P. 1988. An outline of evolution of the
hymenopterous insects (Order Vespida). Orifiitiil
Insects 22:115-145.

Ronquist, F. 1994. Morphology, phylogeny and evo-
lution of cynipoid wasps. Acta Uiiiversitath Upp-
snhieiisis 38:1-29.

Ronquist, F. 1995. Phylogeny and early evolution of
the Cvnipoidea (Hymenoptera). Systcincitic Ento-
mology 20:309-335.

Thompson, T. E. and M. S. Blum. 1967. Structure and
behaviour of spermatozoa of the fire ant Solcnop-

sis saeinssinia (Hvmenoptera: Formicidae). Annuls
of the Entomological Societi/ of America 60:632-642.

Wheeler, D. E., E. G. Crichton and P. H. Krutzsch.
1990. Comparative ultrastructure of ant sperma-
tozoa (Formicidae: Hymenoptera). ]ournal of Mor-
yhology 206:343-350.

Wilkes, A. and P. E. Lee. 1965. The ultrastructure of
dimorphic spermatozoa in the hymenopteron
Dalilbominiis fiiscipennis (Zett.) (Eulophidae). Ca-
nmiian journal of Genetetics and Cytology 7:609-
619.

J. HYM. RES.
Vol. 8(2), 1999, pp. 204-237

Cladistics of the Aleiodes Lineage of the Subfamily Rogadinae
(Hymenoptera: Braconidae)

Joseph C. Fortier and Scott R. Shaw

(JCF) Department of Biology,

Wheeling Jesuit University, P.O. Box 6295, Wheeling, West Virginia 26003 USA;

(SRS) Department of Natural Resources, P.O. Box 3354, University of Wyoming, Laramie,

Wyoming 82071 USA

Abstract. — A ciadisHc analysis was made for 208 Aleiodes species. Seventy three characters were
examined. Host association data from literature and museum specimens were mapped onto the
cladogram. The genus consists of 18 species-groups within three sections: basal, intermediate, and
derived. Basal and intermediate species tend to be generalist feeders while derived species tend
to attack noctuids. The monophyly of seven subgenera was discussed, which were subsumed
under Aleiodes.

The purpose of this research was to pro-
vide a cladistic analysis of world species
of Aleiodes Wesmael 1838, which could be
used to interpret patterns of host-use by
these wasps. Particular attention was giv-
en to defining species-groups by way of
cladistic analysis based on morphological
characters. Host association data was
mapped onto the cladogram to determine
any associations between parasitoid phy-
logeny and host-utilization. Such associa-
tions might indicate coevolutionary pat-
terns within host-parasitoid relationships
that developed over evolutionary time.

Efforts have also elucidated intergeneric
relationships within subfamily Rogadinae,
as well as interspecific relationships with-
in Aleiodes (Achterberg 1991, 1993b; Shaw
1983, Shaw 1993, Shaw 1994, Shaw et al.
1997, Whitfield 1992). For many years
there has been confusion about the defi-
nitions of the genera Aleiodes and Rogas
Nees von Esenbeck. This problem has
been clarified by Achterberg (1982, 1991),
Achterberg and Penteado-Dias (1995),
Shaw (1993) and Shenefelt (1969).

Morphological synapomorphies which
define the subfamily: Rogadinae are: 1)

propodeum either without distinct area
superomedia, or if present, small; 2) dorsal
carinae of metasomal tergite I converging
posteriad; 3) metasomal tergites II and III
well sclerotized and with extensive, dis-
tinct surface sculpturing (Shaw 1995).

The following features define Aleiodes as
monophyletic: 1) basal portion of tarsal
claw rounded with a pectin of spines
(Shaw 1995); 2) tergite II carinate and an-
teromedial area with a polished triangular
region that continues posteriad as a me-
dian carina (Shaw 1995); 3) median carina
of propodeum never forking in anterior
half of the segment and never diverging
posteromedially into an areola (Shaw
1995); 4) fore wing RS + MB vein long
(Shaw 1995); 5) emergence hole from host
mummy even and circular (Shaw 1995); 6)
ovipositor sheath widened and flattened
(Achterberg 1991).

Most Rogadini appear to be solitary
parasitoids, but two gregarious species are
known {A. stigriiator (Say), A. pallesceus
Hellen). As far as known, most Rogas spe-
cies attack Limacodidae, Lycaenidae,
Riodinidae, and Zygaenidae (Shaw, 1995).
Aleiodes species utilize a wide variety of

Volume 8, Number 2, 1999

205

hosts (Shenefelt 1975; Shaw and Huddles-
ton 1991, Shaw 1995).

Aleiodes usually glue their host mum-
mies to a substrate by way of a hole
chewed in the ventral area of the host pro-
thorax. Emerging Aleiodes adults cut an
emergence hole at the posterior end of the
mummified caterpillar (Shaw 1995).

Aleiodes is a useful group of parasitoids
for comparative studies for three reasons:
1) the physiological character of host
mummification preserves the host for tax-
onomic study; 2) the genus is relatively
species-rich, and thus offers a large sam-
ple of species with which to carry out a
phylogenetic analysis; 3) in general, Aleio-
des species tend to have narrow host rang-
es (Shaw 1994), yet across the genus, a
wide variety of lepidopteran groups are
attacked.

METHODS AND MATERIALS

Characters and character states. — Two
hundred eight Aleiodes species were ana-
lyzed. A list of 62 evolutionary transfor-
mation series (Hennig 1966, Mickevich
1982, Mickevich and Weller 1990, and Wi-
ley 1981) including 73 characters (num-
bered 0-72) and component character
states, was derived from Aleiodes speci-
mens examined (see Character Summary).
Terms used to describe sclerite sculptur-
ing features follow those of Harris (1979),
while those used for sclerites and append-
ages follow Huber and Sharkey (1993).
Wing venation terminology follows Shaw
et al. (1997).

The characters used fell under two gen-
eral categories: qualitative and morpho-
metric. Qualitative categories included 1)
sculpturing on specific sclerites, 2) shape
of specific sclerites, 3) presence or absence
of setation on specific body parts, 4) pres-
ence or absence of tarsal claw features, 5)
presence or absence of specific wing veins,
and 6) coloration. Morphometric catego-
ries included 1) number of flagellomeres,
and 2) morphometric ratios of sizes of
parts, or distances between parts.

Species included in the study. — Descrip-
tions of each character are included below
(Character Summary). Appendix I lists the
Aleiodes species analyzed, including host
information. The following species of Ro-
l^as, the putative sister-group of Aleiodes
(Whitfield 1992) were used for outgroup
comparison to best estimate polarization
of transformation series for each character:
R. discoideus (Cresson), R. fusciceps (Cres-
son), and R. bucculatricis (Ashmead). Four
unnamed morphospecies of Clinocentrus
Haliday were also used, as well as the fol-
lowing Stiropius (Whitfield) species: S. buc-
culatricis (Ashmead) and S. californicus
Whitfield.

Character polarization. — Aleiodes charac-
ter states which most closely resembled
those of Rogas, Stiropius, and /or Clinocen-
trus (Whitfield, 1992) were hypothesized
to resemble most closely the ancestral
state (Hennig, 1981). For those characters
for which there was significant, consistent
departure in the Rogas state in comparison
to the Aleiodes state(s), the latter two gen-
era were examined to determine whether
the Aleiodes or Rogas condition should be
considered ancestral. If a given character
state for Rogas species departed from that
of Stiropius species, Clinocentrus species,
and Aleiodes species, it was hypothesized
to be derived. The state the latter three
genera expressed in common was then hy-
pothesized to be the ancestral state. If a
given state expressed by Stiropius species
departed from one that Aleiodes and Cli-
nocentrus held in common, but not Rogas,
and the Rogas state differed from the Sti-
ropius state, the state that Aleiodes and Cli-
nocentrus shared was hypothesized to be
ancestral to Aleiodes, and the other two
were hypothesized to be independently
derived autapomorphies with respect to
the four genera taken together. For cases
in which a Rogas state was hypothesized
to be derived, Stiropius was hypothesized
to be more closely related to the ancestor
of Rogas + Aleiodes than Clinocentrus
(Whitfield 1992).

206

Journal of Hymenoptera Research

Ordering transformation series. — The or-
dering of character states with respect to
one another was determined by polariza-
tion based on the hypothetical ancestor, as
described above, and parsimony, as fol-
lows: For those transformation series with
more than one state, once the ancestral
state was hypothesized, various combina-
tions of states for a given transformation
series were tested with the Hennig86 phy-
logenetic analysis program (Farris 1988) as
described in the next section. The ordering
of character states for the character in
question on the most parsimonious Nel-
son consensus tree among those derived
from respective Hennig86 runs was hy-
pothesized to represent the best estima-
tion of phylogeny for that transformation
series. All characters in this study consist
of ordered character states. The Nelson
consensus provides a strict consensus tree
from equally parsimonious trees derived
from a Hennig86 run. Exceptions to as-
signing '0' to the ancestral state were the
following classes of hypotheses: 1) bipolar
linear evolutionary transformation series,
in which character states evolved in two
directions from the ancestor, diagrammed
thus:

a <- b ^ c

and reticulate transformation series (Mick-
evich 1982, Mickevich and Weller 1990), in
which either /) character states were hy-
pothesized to have evolved from more
than two directions from the ancestor,
and /or ii) the ancestral character state for
a given Aleiodes lineage or closely related
lineages was an Aleiodes character state
other than the ancestral Aleiodes character
state for that entire transformation series.
We adopted the method of TSA for
treating complex multi-state characters
such as vertex sculpturing (Shaw et al.
1997), tarsal claw form, clypeal carina
shape, propodeum sculpturing (Shaw et
al. 1997), mesopleuron sculpturing, meta-
somal tergite sculpturing, and hind wing
RS vein shape (Achterberg 1993a). This

method has been described by Lipscomb
1992, Mickevich and Lipscomb 1991, and
Mickevich and Weller 1990, and is com-
monly used (Lipscomb 1992, Mitter and
Silverfine 1988, and Schuh 1991). It has
been critiqued by Lipscomb (1992).

Alternate arrangements of character
states of transformation series, in which
more than one arrangement was possible,
and it was not obvious from morphology
alone how these character states were
most logically ordered, were tested using
Hennig86 program 1.5 (Farris, 1988) on a
Dell Omniplex 466 system, to find the
most parsimonious scheme for each re-
spective reticulate transformation series.
The most parsimonious scheme was then
incorporated into the matrix.

The matrix and execution of the phyloge-
netic analysis. — An integer matrix (Table 1)
based on integers representing coding for
the states of the characters (see Character
Summary), arrayed with the hypothetical
ancestor and Aleiodes species in rows and
characters in columns was analyzed by
computerized algorithms (Farris 1970), us-
ing the Hennig86 program Version 1.5
(Farris 1988) on a Dell Omniplex 466 sys-
tem. The results were expressed as Nelson
consensuses of multiple equally parsimo-
nious, minimum length Wagner trees, ob-
tained by using the mhennig* option with
extended branch swapping (bb*) with and
without successive approximations to
character weighting (xs w; cc;) until stable
solutions were reached (Farris 1988), in
tandem with the CLADOS program (Nix-
on 1991). The CLADOS program was use-
ful for 1) obtaining definitions of nodes
based on transformation series changes for
characters at each given node, 2) carrying
out manual branch swapping to further
minimize Nelson consensus tree lengths,
and 3) illustration. The trees obtained
were the most parsimonious explanations
of the distribution of the observed data set
that could be found, barring use of the
Hennig86 ie* option (Farris 1988), which
was found impractical (see below). After

Volume 8, Number 2, 1999

207

a-«— b— *>c— •►d-
Characters 7-8 g

a— •-b— ►c— ►d
Characters 17-18

g '

t t

a— ► b— •► c— *- d— »► e

f J

Characters 21-24

f

t

a— *-b— ►c— •-d

Characters 31-32

t

a— ►b— »-c

\

Characters 34-36

f

i

.i

t

a^4-b-

-►c-

\

\

e

h

Characters 42-44

a— *>b— #>c

\

d

Characters 48-49

Fig. 1. Diagrams of reticulate transformation series.

-b— ►►c

Characters 68-69

208 Journal of Hymenoptera Research

Table ]. Matrix of Alciodcs species and character states. See Appendix for complete species names and text
for explanation of characters.

01234567

ANCESTOR 0000000111000000000000100010000010010000001110000000010000001000000100000

ABDOMINA 021102031101000013 0002100011101132110001002110000021121000101000221100000

ACICULAT 0111010311000000010001000111000121010000001210020021011000001000210100000

AESTUOSU 021112031101000213 0012100010001212110001002110001121023000001000211100100

AGILIS 0111120311000002130013100110001211110001002110001121123000001000110000100

ALBITIBI 0111011311000000110002110011000111110000001110000011011010001000210100000

ALBOANNU 011112030101000003 0002100111701131110111002110001021121000101000210101000

ALUTACEU 011112031100000003 0001200111000111110010101110000021010000000000110100000

ALIGARHE 0111001211000000010001100011000111010000001110100011011000001000220100000

ALTERNAT 0111010311000000010001000111000121110000001210010021011000001000221100000

ANGUSTAT 0121011211100000010001000111000121110000001210010071010000001000210101000

ANNULATU 0101010211000?00030001100111000111010000001110000021010000000100?10100000

ANTENNAT 0111120311000010130002100010001111110111003110000021101000001000210100000

APICALIS 0211020311010000130012100011101132110001002110000021121000101000221100000

ARCTICUS 0111010311000000010001100011000111100000001310030021011010002000210100000

ARIZONIE 0211120311100010030002100010011111210010003110000021111000002000221100000

ARMATUS 0111011211000000010001100011001111110000001210000021010000001000210100000

ARNOLDI 0111120311010000130012100111701131110001002110000021022000001000210101000

ARSENJEV 0111120311010010030002100010011111110111103110000020001000001000710100000

ATRICEPS 0211020311010000130002100011101132110001002110000021122000001000221100000

ATRICORN 0111120311010000030012100111001141110001002110002021112000001000210101000

AUTOGRAP 0111011211000000010001100011000111110000001110000011011000011000210100000

BAKERI 1111011211010000020001100111000111110000001110000071011000001000210100000

BICOLOR 0111010311000000010001000111000121100000001310010021011010001000210100000

BOREALIS 0111011211000000010001100011000111110000001110000011011000011000210100000

BRETHESI 0211020311010000130002100011101132110001002110000021131000001000221100000

BREVIPEN 01110103110017000301013 00111000110100000111110000020000000000100110100000

BREVIRAD 0111020311010000030001000111001121100000001210000021001700001000710100000

BUOCULUS 0111011211000010010001100011000111110000001110000011011000001001210100000

BURRUS 0111120311010000130013100111001131100001002110001021122000001000210101000

CAMERONI 0211120301010010030012100010011111210000003110000021011000002000221100000

CANTHARI 0111011211000000010001100011000111110000001110000011017000011001710100000

CARINATU 0111120311010000130013100111001131110001002110001021112000001000221101000

CARINIVE 0211120311010000130013100011001131110001002120000021122010001000221100000

CAUCASIC 01111213110000001300121000110111311000010021100000210010000010002 7 7100000

CAZIERI 0211120311110010030002100010001131210000002110000021111000002000221100000

CHLOROTI 0111111301000010010002100110070111210000003110000021110000001001110100000

CIRCUMSC 0111011211000000010001100011000111110000001110000011011000011000210100000

COMPRESS 0221001211100000010001100011000111110000001110100011011000001000220100010

CONFORMI 0111120311010000130013100111001131110001002110001021122000001000210101000

CONVEXUS 0211020311010000130002100011101212110001002110000021121010001000221100001

COXALIS 0111010311000000010001000111000121100000001310010021011010001000210100000

COXATOR 0211010311010000010001000111000121100000001210010021011000001000221100000

CRASSIPE 0101011311170000010001100111000111110000001110010071011000001000710100000

CRUENTIS 021112031101000013 0013100110001311120001002120001021122000001000221101100

CURTISPI 0111010311000000010001200111000111110000001110000021011100000000 710100000

DENDROLI 0111011211000010010001100010000111110000001110000011011010001000210100000

DESERTUS 0111120311000002130012100110001211110001002110001171723000001001121000100

DISPAR 011102031100100003 01013 00111001111110010111110000021000000000100110100000

DISSECTO 021112031101000013 0012100010001311120001002120001021122010001000221101100

DIVERSIC 0111010311071000030001200111000121010010101110000021101000000100110100000

DIVERSUS 0211120311010000130012100010 701211110001002120001021022011001000221101100

DUCTOR 0211020311010000130002100011101132110001002110000021121000101000221100000

EARINOS 0111121311000010030012100110011131210000003110000021101000002000220100000

ECUADORI 0111120311010000030014100111001131110011002110000021102000001000210101000

EURINUS 0111120311010000130013100111001131110001002110001021122000101000210101000

EXCAVATU 0111010311001000030101300111001121110000111211030020000000001100110100000

FAHRINGE 0111020311010002130012100010001212110001002110002021123000001000210001100

FEMORATU 1111010211000000030001100111000111010000001110000011010000001000210100000

FERRUGIL 0111020311000000130012100111001141110001002110000021121000001000210001000

FLAVIDUS 0111121301170010030002100020001131210000003110000021111000001000211100000

FLAVISTI 0211020321010001130012100000001101130001002110002021132110001000221101100

Volume 8, Number 2, 1999 209

Table 1. Continuod.

01234567
FLAVITAR 011102031101000003 0002100011101112110001002100000021111000001000221100000
FORTIPES 0111121311070000030012100011011131110000003110000021011000002000210100000
FORTIS 0111120311010000130013100111001131110001002110001021122000001000210101000
FUSCIPEN 01110213110000000 10002110011000111 1100000011100000 71021000001000710100000
GASTERAT 0211120311010000130013100111001131110001002110001021122000101000211101000
GASTRITO 0111011211000100010001100011000111110000001110000011011000001001110100000
GEOMETRA 0211121301000010030012100010011111210000003110000021011000002000221100000
GLABER 0121720311010012130012100110001211120001002100001121023000002000110000100
GOSSYPII 0111011211000000010001100011000111110000001110000011011000001000210100000
GRANDIS 021112031101000003 0011000011101121100001002110000021111010001000221100000
GRANULAT 0111010311001000010001100111000111010000101210020011010000001100210100000
GRAPHICU 0111110311000010111002100001000101110000101110000021021000001000210100000
GRASSATO 011112031101000013 0013100111001131110001002110000021121000001070210101000
GRESSITT 0111001011000000010011100011000111111000001110000011012000001000210100000
HARRIMAN 0111020311010000130012100110001131110001002100001021121000101000210101000
HELLENIC 0111011311000000010001000111001121110000001217020021010000000000710100000
HIRTUS 0211020311010000130012100111001131110001002110001021121000101000210101000
INCERTOI 0111010311000000010001000111000121100000001210020071011000001000210100000
INCERTUS 0111010311000000010001000111000121110000001210010021111000001000210000000
INDESCRE 0211011311000010010001100011000111110000001110000011011000001000221100000
JAKOWLEW 011101031110000001000100011100012110001000131003 0021111000011000710100000
JAROSLAW 021112031101000013 0012100011700131110001002110000020121000101000221101000
KRULIKOW 0111020311010002130013100110001131120001002120001021103000101000210101100
KUZTLIZK 1111011211000000010001100111000112110000001210000011010000011000210100000
LAPHYGMA 0111011211000000010001100011000111110000001110000011011000001001210100000
LONGICOR 7111010311170000030101200111000111010010001110000021110000000100770100000
LONGIPEN 0111010311101000030101300111000120100000101211000020000000000100110100000
LUCIDUS 0211020321010001130012100000001101140001002100002031232110001000221201100
LYMANTRI 0211011211000010010001100010001111210000001110000011011000011000221100000
MALACOSO 0111011211000000010001100011000111110000001110000011011000001001110100000
MANDIBUL 0211120311010002130012100110001211110001002110002021123000001000221101100
MEDIANUS 0221020311010000030002100111001111110000002110000071021000001000221101000
MELANOPT 0211020321010001130012100000001101140001002100002021232110001000221201100
MEXICANU 021102031101000113 001210000000110113 0001002110002021132000001000221101100
MICROCUL 011112031101000003 00031001117 70310120001102110000021101010001000221101100
MINIATUS 012112031101000113 0013101111001311120000002120002021123000001000210001100
MODESTUS 0111010311000000010012100110000111120000002120001021022000001000210101000
MOLDAVIC 0101010311000000010001000111001121100000001310020021011000001000710100000
MOLESTUS 021102031101000013 0002100111101132110001002110000021121000001000221100000
MORIO 0111120311010000130012100111001311110001002110000021121010001000221101000
NARANGAE 0111010211001700010001000011000111110000001210020011000000000100110100000
NEGATIVU 0211010311000000010001000110001121110001101210000021110000001000221101000
NEOTROPI 7111011311170000010001100011000111110000001110000021011000001070710100000
NIGRIBAS 1111011211100010020001100111000111110000001110000071110000001000210100000
NIGRICOR 0111011211000000010001100011000111110000001110000011011000011000110100000
NIGRISTE 1111010211000000020001100111000111010000001110000011010000001000210100000
NOCTURNU 0121001211000010010011100010000111110000001110000071011000001001210100000
NOLOPHAN 0111011211000000010001100011000111110000001110000011011000001001110100000
NOTOZOPH 0211121301000010030002100010011131210000003110000021011000002000221100000
^aMBERGI 0111010311000000010001100111000121100000101310010021011010001000210100000
PALLESCE 0111011211000100010001100011000111110000001110000011011000001001110100000
PALLIDAT 0211011211000000010001100111000111100000001110000011011000001000211100000
PALLIDIS 0111120311010000130003100111001131110001002110001021112000000000210101000
PALMATUS 0121001211100000010001100011000111110000001110100011011000001000220100010
PARASITI 0211020311010000130002100111101132110001002110000021121000001000221100000
PEDALIS 0111121301100010030002100020071131210000003110000021111000001000211100000
PELLUCEN 0111001211000000010001100010000111110000001110000011011000001000210100000
PERCURRE 1111011211000 700010001100011000111110000001110000011010000001001210100000
PERINETE 0111010311001000030001200111000121010000001110000021000000000000110100000
PERISCEL 0211020311000000130012100011001311120000002120001021122010001000211101000
PERPLEXU 0111011211000000010001100011000111110000001110000011011000001001210100000
PICTUS 0111011211000000010001100011000111110000001110000011011000011000210100000
PLATYPTE 0111011211000000010001100011000111110000001110000011011000001001210100000

210 Journal of Hymenoitera Research

Table 1. Continued.

01234567
PLEURILI 0111010311010000030101000111000121010000001210000021001000000000110100000
POLITICE 021112031101000113 0012100000001101130001002110002021122000001000221101100
PRAETOR 0111110311000010111012100011000111110000101110000021021000001000210100000
PROCERUS 0111010311101000010?011000110011211100000011100100?1011000001000210100000
PULCHRIP 021112031101001003 0012110010001131110000002110000021011000002000221100000
PUNCTIPE 0111010311000700010001100111000111110000001110000021011000001000110100000
QUADRUM 02 111203111100001300121001110012111100010021107010211 22 000001070221101000
QUEBECEN 0211120301010010010012100010001131210110003110000021011000002000221100000
RILEYI 0211120311010000010012100011101131110000002110000021111000001000221100000
ROGEZENS 0111010311100000030101201111000111010010001110000020110000000100710100000
ROSSI 0211121301000010030012100010011111210000003110000021011000002000221100000
ROSSICUS 0101001211000000010001100011000111110000001110000011011000001000210100000
RUFICEPS 0111121311010001130012100101001211110001002110000021122000001000210101000
RUFICORN 0111120311010000130013100111001131110001002110000021122000101000210101000
RUFIPES 0111120311010000130002100111001131110001002110001021122000001000211101000
RUGOSICO 0111010311001000030101300111001110110000111111000020000000000100110100000
RUGULOSU 0211120311010000130012100111001311110001002120001021121010001000221100100
SANCTIHY 0111010312000000010001000111000121100000101310010021011001001000210100000
SANCTIVE 1111000211100000010001100111001111110000001110000010000000000000210100000
SATANAS 0111010311000000010001000111001121110000001210010071011011001000210100000
SCHIRJAE 0111020311070000130002100111001131110001002110000021121000001000210101000
SCRIPTUS 1111010211000000010001100111001111010000001110000021010000001000710100000
SCRUTATO 0111011211000000010001100011000111110000001110000011011000011000210100000
SERIATUS 1111010211000000030001100111000111110000001110000011010000001000210100000
SHESTAKO 0211120301010000130002100110001312120001102120000021122110001000221101100
SIBIRICU 0211020311010000130002100111001131110001002110000021121000001000221100000
SIGNATUS 0111010311000000010001000111000121100000001110010021011001101000211100000
SIMILIS 0111011211000000010001100011000111110000001110000011011000011000210100000
SIMILLIM 0111011211000010010001100010000111110000001110000011010000001001210100000
STIGMATO 0211011211000100010001100011000111110000001110000011011000001001111110000
SUDATORI 0111010311000000010001100111000111110000001110000021011000001000 710100000
TATIANAE 0111020311101000030101300111000121110010111317030020000010001100110100000
TESTACEU 0111001211000100010001100011001111110000001110000011011000001001210100000
TERMINAL 0111020311010000130012100111001131110001002110000021121000101000210101000
TETRASPH 0111010311000000010001000011000121100000001310040021011000001000210100000
TEXANUS 0111110411000010111002100001000101110000101110000021021000001000210100000
TRANSVER 0111020311111000030001200111000121010000001110000021111000000100770100000
TRISTIS 0211010311000000010001000111000121110000001210010071011010001000211100000
TURKESTA 0111120311070000130013100111001131110001002100000021122000101000210101000
UFEI 0111120311110000130012100111001211110001002110101071122000101000210101010
UNGULARI 0211010311010000010002100011000121100000002110000021021000001000211101000
UNIPUNCT 0111020311010000130012100110001131110001002100000021122000101000210101000
VAUGHANI 02110203 01010010030012100010001131210000003110000021011000002000221100000
VENUSTUL 0111020311070001130012100011001211120001002120000021122000001000210101100
WADAI 01111203 01010002130002100111001211120001002120001021122000001000210101100
XANTHUS 0111071211000010010001100011001111110000001110000011011000001001710100000
0111011211101000010001300111000111010000001210020011010000000100210100000
0111010312000000010001000111000121100000101310010011011001001000210000000
0111011211000000010001100011000111110000001110000021011000001001110100000
0111120311010000130013100111001131110001002110001021122000001000221101000
0111010311010000010001000111000111110000001210000011011000001000210100000
0111120311010000130013100111001131110001002110001021022000001000221101000
NEW SP.7 0111011211010000010001000111000111110000001110000011011000001000210100000
NEW SP.8 0111110311000010111001100011000111110000101110000021011000001000210100000
NEW SP.9 0111011211100000010001100011001111110000001110000011010000001000210100000
NEW SPIO 0111010311000000010001000111000121110000001210010021011000001000210100000
NEW SPll 0111010311000000010001000111000121100000001310010021011010001000210100000
NEW SP12 0111011211101010010001100110000111110000001210010011011000001000210100000
NEW SP13 0211010311000000010001000111000121100000001310010021011010001000210100000
NEW SP14 011101031100000001000110001100011110000010131003 0021011010001000210100000
NEW SP15 0111001211000100010001100011000111110000001210000011011000001011210100000
NEW SP16 0221001011000000010001100011000111111000001010000011011000001000111110000
NEW SP17 0111010311000000010001100011000111100000101310030021011010001000210100000

NEW

SP.

.1

NEW

SP,

,2

NEW

SP.

.3

NEW

SP.

.4

NEW

SP.

.5

NEW

SP.

.6

Volume 8, Number 2, 1999
Table 1. Continued.

211

NEW
NEW
NEW
NEW
NEW
NEW
NEW
NEW
NEW
NEW
NEW
NEW
NEW
NEW
NEW
NEW
NEW
NEW
NEW
NEW
NEW

SP18
SP19
SP20
SP21
SP22
SP23
SP24
SP25
SP26
SP27
SP28
SP29
SP30
SP31
SP32
SP33
SP34
SP35
SP36
SP37
SP38

02110112
01110012
01111203
01110203
02111203
02111203
01110112
01110203
02110103
01110103
01111203
01111104
11110112
01111203
01110112
11110112
02110103
01110012
01010103
02110103
01110112

1

11000
11000
11010
11000
11010
11010
11100
11010
12000
11001
11010
11000
11000
11010
11000
11000
11000
11000
11000
11000
10100

01001
00001
00013
00213
00213
00013
00001
00013
00001
00001
00003
00011
00003
00013
00001
00002
00001
00001
00001
01001
01001

00011
00011
00131
00121
00121
00121
00011
00121
00010
00010
00141
10021
00011
00121
00011
00011
00010
00011
00010
00011
00011

00010
00011
00111
00010
00100
00110
00011
00110
00111
00111
00111
00001
00111
00111
00011
00111
00111
00011
00111
00111
00011

00011
00011
00113
00121
00131
00121
00011
00113
00012
00012
00013
00010
00011
00113
00011
00011
00012
00011
00012
00111
00111

1110000
1110000
1110001
1110001
1110001
1110001
1010000
1110001
1110000
1100000
1110011
1110000
1010000
1110001
1110000
1010000
1100000
1110000
1100000
1110000
1110000

001110
001110
002110
002100
002110
002110
000110
002110
101210
001310
002110
101110
001110
002110
001110
001110
001210
001110
001210
001110
001110

00001
00001
00102
00112
00102
00102
11001
00102
01002
01002
00002
00002
00001
00102
00101
0000?
01002
00001
00002
0000?
00001

10110
10110
11220
10230
11330
11230
10110
11210
10110
10110
11120
10210
10100
11220
10110
10100
11110
10110
10110
10110
10110

000020
000010
000010
000010
000010
000010
000010
001010
000010
000010
000010
000010
000010
000010
000010
000010
000010
000010
000010
000010
000110

00221
11210
00210
00210
00221
00221
00220
00210
00221
00210
00210
00210
00210
00210
01210
00210
00221
11210
00210
00221
00210

100000
100000
101000
000100
101100
001100
100010
101000
100000
100000
101000
100000
100000
101000
101000
100000
100000
100000
100000
100000
100000

the analysis exceeded 128 taxa, a compre-
hensive matrix (Table 1) was maintained
from which 'sample matrices' were drawn
for analysis of newly analyzed species
within the context of a representative sam-
pling of species analyzed to date.

'Species-group' definition refinement. — It
was found that "species-groups" consis-
tently clustered on cladograms derived
from Nelson consensuses throughout the
course of the study (see Results). Refine-
ment of these species-group definitions
was aided by use of sample matrices as
follows. For a given species-group, all spe-
cies previously found to be in the group
were included in a sample matrix as de-
scribed above, as well as most or all spe-
cies in contiguous species-groups. Manual
branch swapping was applied to the Nel-
son consensus tree after running the ma-
trix on Hennig86 (m*; bb*; xs w; cc) in or-
der to further refine the definition of the
species-group by further reducing the
length of the tree.

After definitions of all characters, char-
acter states, transformation series, and
species-groups had been finalized, a final
matrix with an approximately equal per-
centage of species from each species-

group (about 61.5%) was analyzed using
the mhennig*; option of the Hennig86 pro-
gram, in order to find the most parsimo-
nious tree (Farris 1988). This percentage
was found to meet the requirement of ob-
taining the tree with 128 taxa, with taxa
most evenly distributed across it with re-
spect to species-groups.

Host association analysis. — After a clado-
gram had been generated as described
above, host associations were mapped
onto the cladogram. Based on available lit-
erature (Shenefelt 1975, Achterberg 1985,
Marsh 1979, Shaw 1983, 1994; Shaw 1995,
Shaw et al. 1997), and museum specimens
with host information attached were ex-
amined (Rocky Mountain Systematic En-
tomology Museum, Laramie, WY and Ca-
nadian National Collection, Ottawa, ON),
host associations were assigned.

CHARACTER SUMMARY

Following is a list describing characters
and evolutionary transformation series of
those characters in this analysis, along
with explanations of how polarities were
hypothesized for each transformation se-
ries. The number assigned to each char-
acter identifies it on the matrix and on the

212

Journal of Hymenoitera Research

cladogram illustrated by CLADOS. The
numbers in brackets indicate character
states within the transformation series for
a given character, and also appear under-
neath the numbers assigned to the respec-
tive characters in the matrix, and on the
CLADOS-generated cladograms for each
species-group. Squares on cladogram
branches below designated character
numbers represent character state trans-
formations which help define the node or
species following the squares. If the
square is darkened, the square represents
an increase in character state number; if
white, a reversal is indicated. Unless oth-
erwise stated, the [0] state was that hy-
pothesized to be ancestral based on Cli-
noceutrus, Rogas and Stiropius specimens
examined.

I. Head Characters

4. Shape of 15th flagellomere from base, fe-
males: [0] width/length less than 0.83; [1]
compact: width /length greater than 0.83.
State [1] was found only in some Aleiodes
species. Thus, state [0] was hypothesized
to be the ancestral state.

6. Occipital cari)ia: [0] strong, complete
medially; [1] weak or effaced medially.
State [1] was found in most Stiropius and
no Cliuoceutriis or Rogas examined, and
was hypothesized to be ancestral. This hy-
pothesis was tested by comparing relative
parsimony with respect to character po-
larization. The same matrix was run on
Hennig86 as described above for two runs
except that polarization was switched for
each. Since the alternate polarity was most
parsimonious (strong, complete occipital
carina ancestral), this alternate hypothesis
was accepted; the RogaslClinocentrus con-
dition was hypothesized to be ancestral.

13. Flagellum, female: [0] > 33 flagello-
meres; [1] s 33 flagellomeres. All Stiropius
species examined expressed state [1], and
no Clinocentrus or Rogas species did. The
more parsimonious character polarization
was found by running each of the two
possibilities on Hennig86. Based on par-

simony, the reduced number of flagello-
meres was hypothesized to be derived
where it occurred in Aleiodes.

14. Ocellar diameter/distance between lat-
eral ocellus and compound eye: [0] < 2.5; [1]
> 2.5 (Shaw et al. 1997, Figs. 1-5). State [1]
was observed in all Rogas species exam-
ined, and no Stiropius. A comparison of
parsimony in which the polarity of this
character was switched between two oth-
erwise identical matrices showed that the
polarization used here was most parsi-
monious. Thus, state [0] was hypothesized
to be the ancestral state.

15. Ratio of clypeo-antennal space to width
of oral space: [0] greater than 0.82; [1] 0.69-
0.82; [2] less than 0.69. Since states [1] and
[2] were not found in non-Aleiodes species
examined, state [0] was hypothesized to
be ancestral.

20. Occipital carina at hypostomal carina:
[0] complete or nearly so; [1] incomplete,
effaced well before reaching hypostomal
carina. Determining polarity was prob-
lematic, since of four Rogas species exam-
ined, two expressed state [0], and two ex-
pressed [1]. The Clinocentrus species ex-
amined (four unnamed morphospecies)
consistently exhibited state [0]. Two Sti-
ropius species examined consistently pos-
sessed state [1]. Both codings were tried
for the ancestor, with no change in the
consistency index (c.i.). The above polari-
zation represents the most parsimonious
solution. Thus, state [0] was hypothesized
to be the ancestral state.

27. Ratio of malar space to mandibular bas-
al width: [0] less than 1; [1] greater than or
equal to 1. State [0] was found in all Rogas
and Cli)iocentrus species examined. State
[1] was observed in all Stiropius examined.
A comparison of parsimony between al-
ternate polarizations revealed that the po-
larization presented here is the simplest
explanation.

31. Vertex sculpturing (path a-d): [0]
coarsely granulate; [1] smooth, granulate,
or faintly rugulose (Shaw ef al. 1997, Fig.
6); [2] smooth with shallow, dense punc-

Volume 8, Number 2, 1999

213

tation; [3] shiny, coarsely and densely
punctate.

32. Vertex sculpturing {path e-h): [0] nit-
id; [1] one of conditions in #31; [2] with
strong laterally running ridges, or rugose;
[3] with fine laterally running ridges (gen-
eralized advanced condition; Shaw et al.
1997, Fig. 5); [4] finely rugulose. The retic-
ulate character described by the above 2
paths is hypothesized as follows: a =
31[0]; b = 31[1]; c = 31[2]; d = 31[3]; e =
32[0]; f = 32[2]; g = 32[3]; h = 32[04].

a

b

c

d

path

n

0

1

2

1

a-c (#48)

n + 1

0

0

0

1

b-d (#49)

a

b

c

d

e

f

g

h

path

n

0

1

2

3

1

1

1

1

a-d (#31)

n

+ 1

1

1

1

1

0

2

3

4

e-h (#32)

The above phylogenetic hypothesis is il-
lustrated in Fig. Id. Stiropius species ob-
served all expressed state (a), and Rogns
species all expressed state (e). A compar-
ison of parsimony between alternate po-
larizations revealed that the polarization
presented here is the simplest explanation.

33. Medial carina extending down frons:
[0] absent; [1] present, extending less than
0.55 of distance from line between bases
of scapes to clypeus; [2] present, extending
0.55 or more of distance from line between
bases of scapes to clypeus. States [1] and
[2] were observed in no non-Aleiodes spe-
cies examined. Thus, state [0] was hypoth-
esized to be the ancestral state.

38. Lightnesf of midsection of antennae: [0]
not lighter than both basal and apical sec-
tions; [1] lighter than both basal and apical
sections (Shaw et al. 1997, Fig. 23). State
[1] was not found in non-Aleiodes species
examined. Thus, state [0] was hypothe-
sized to be the ancestral state.

48. Clypeal carina (path a~-c): [0] absent;
[1] present, not shelflike; [2] shelflike.

49. Clypeal carina (path b-d): [0] clypeus
either not elongate, or with a carina; [1]
elongate clypeus without carina. The retic-
ulate character described by the above 2
paths is hypothesized as follows: a =
49[0]; b = 49[1]; c = 49[2]; d = 50[1]

The above phylogenetic hypothesis is il-
lustrated in Fig. Ig. States [1] and [2] were
observed in no non-Aleiodes examined.
Thus, state [0] was hypothesized to be the
ancestral state.

51. Length of first flagellomere: [0] s
scape length; [1] < scape length. State [1]
was not observed in non-Aleiodes species
examined. Thus, state [0] was hypothe-
sized to be the ancestral state.

54. Cli/peus height divided by icidth: [0] >
0.65; [1] < 0.65 > 0.42; [2] <\42 > .30; [3]
^ 0.30. Height was defined as distance be-
tween clypeo-labral suture and dorsal
edge of clypeus. Width was defined as
distance between inner edges of tentorial
pits. All Stiropius and Rogas examined ex-
pressed state [0]. Clinocentrus expressed
state [1]; However, given that Stiropius
and Rogas are phylogenetically closer to
Aleiodes than Clinocentrus (Whitfield 1992),
the elongated clypeal condition in Clino-
centrus was hypothesized to be indepen-
dently derived with respect to the Aleiodes
condition, and state [0] was hypothesized
to be ancestral.

55. Third maxillary palpomere: [0] not
swollen; [1] swollen. State [1] was not ob-
served in non-Aleiodes species examined.
Thus, state [0] was hypothesized to be the
ancestral state.

69. Clypeus shape: [0] not abruptly
edged, not flat ventrad (ancestor); [1]
abruptly edged, flat ventrad. State [1] was
not observed in non-Aleiodes species ex-
amined. Thus, state [0] was hypothesized
to be the ancestral state.

70. Wide, flat flange on lower side of man-
dible: [0] not strongly present; [1] strongly
present. State [1] was not observed in non-
Aleiodes species examined. Thus, state [0]
was hypothesized to be the ancestral state.

214

Jouy<NAL OF Hymenoptera Research

II. Mesosomal Characters

3. Median carina of propodeum: [0] short
and forking before halfway point of pro-
podeum (Achterberg 1991, Fig. 121); [1]
not forking before halfway point of pro-
podeum. Rogas and Clinoceiitrus species
examined expressed the [0] state. Stiropins
and Aleiodes independently expressed dif-
ferent states which were not [0]. Thus,
state [1] was hypothesized to be the an-
cestral state for Aleiodes.

7. Sculpture of propodeiiw antero-laterally
(path a-e): [0] finely, smoothly granulate;
[1] coarsely granulate, or areolate; [2] ru-
gulocostate over granulate surface, not
shiny; [3] rugose over granulate surface or
no underlying granulation; [4] smooth-
punctate.

8. Sculpture of propodeum antero-laterally
(path f-g): [0] granulate with faint rugulos-
ity or granulate (Shaw et al. 1997, Figs. 7-
10); [1] one of states in #7; [2] nitid. The
above two paths constitute a reticulate
transformation series, which is hypothe-
sized as follows: a = 7[0]; b = 7[1]; c =
7[2]; d = 7[3]; e = 7[4]; f = 8[0]; g = 8[2]

a

b

c

d

e

f

g

path

n 0

1

2

3

4

3

3

a-e (#7)

n -H 1 1

1

1

1

1

2

0

f-g (#8)

The above phylogenetic hypothesis is il-
lustrated in Fig. la. The states represented
by (b) were not found in Aleiodes. The pro-
podeums of Clinocentrus and Rogas were
areolate; that of Stiropins was coarsely
granulate. Thus, one of the states repre-
sented by (b) was hypothesized to be the
ancestral state.

12. Pronotum angle: [0] not shelflike
(Achterberg 1991, Figs. 283, 294, 380); [1]
shelflike (Achterberg and Penteado-Dias
1995, Figs. 6, 11, 16, 21, 26) State [1] was
not observed in any non-Aleiodes species
examined. Thus, state [0] was hypothe-
sized to be the ancestral state.

16. Large setal pits on scutellum: [0] ab-
sent; [1] present. Since state [1] was not

found in any non-Aleiodes species exam-
ined, state [0] was hypothesized to be an-
cestral.

19. Prescutellar carina: [0] without an an-
terior invagination; [1] with an anterior in-
vagination. State [1] was observed in no
non-Aleiodes species. Thus, state [0] was
hypothesized to be the ancestral state.

21. Mesopleuron central disc sculpturing
(path a-e): [0] shiny coarse granulation; [1]
granulate; [2] smooth-punctate; [3] heavily
punctate.

22. Mesopleuron central disc sculpturing
(path f-h): [0] coarsely rugulose; [1] gran-
ulate, or other states in #21; [2] costate
from granulate or partly costate, partly
granulate; [3] partly smooth, partly rugu-
lose.

23. Mesopleuron central disc sculpturing
(c-i): [0] not nitid; [1] nitid.

24. Mesopleuron central disc sculpturing
(d-j): [0] not costate from punctate; [1] cos-
tate from punctate. The reticulate charac-
ter described by the above 4 paths is hy-
pothesized as follows: a = 21[0]; b = 21[1];
c = 21[2]; d = 21[2]; e = 21[4]; f = 22[0];
g = 22[2]; h = 22[3]; i = 23[0]; j = 24[1].

a

b

c

d

e

f

g h i

i

path

n

0

1

2

3

4

1

1 1 2

3

a-e (#21)

n-hl

1

1

1

1

1

0

2 3 1

1

f-h (#22)

n-F2

1

1

1

1

1

1

1 1 0

1

c-i (#23)

n-^3

0

0

0

0

0

0

0 0 0

1

d-j (#24)

The above phylogenetic hypothesis is il-
lustrated in Fig. Ic. In this case, a reticu-
late transformation series was considered
the best way to interpret the data because
of the high degree of variability. No Aleio-
des species were coded with (a). Clinocen-
trus and Rogas species examined ex-
pressed the nitid condition, and Stiropins
expressed shiny, coarse granulation. Giv-
en the plasticity of mesopleural sculptur-
ing within genus Aleiodes, as can be seen
by the above complex reticulation of char-
acter states, the Aleiodes nitid condition
was hypothesized to have evolved inde-

Volume 8, Number 2, 1999

215

pendently from the Rogas nitid condition.
The shiny coarse (a) condition was hy-
pothesized to have been the ancestral con-
dition for Aleiodes.

25. Stenumliis: [0] not sculptured; [1] ru-
gose or rugulose. Stiropius expressed state
[0]. Rogas expressed a uniquely foveate
condition not found anywhere else, and
Clinocentrus expressed rugation less regu-
lar than the Rogas condition, but more so
than the condition observed in Aleiodes.
Since only Aleiodes species expressed the
[0] condition in common with Stiropius,
the [0] condition was hypothesized to be
ancestral.

39. Mesopleiiral pit posteriad to central
disc of mesopletiroiv. [0] absent; [1] present.
State [1] was not found in non-Aleiodes
species examined. Thus, state [0] was hy-
pothesized to be the ancestral state.

40. Protuberances or a raised ridge imme-
diately anteriad to propodeal apex: [0] absent;
[1] present. State [1] was not found in non-
Aleiodes species examined. Thus, state [0]
was hypothesized to be the ancestral state.

41. Pronotal anterior flange: [0] < 0.28 of
pronotal length; [1] > 0.28 of pronotal
length. State [1] was not found in non-Al-
eiodes species examined. Thus, state [0]
was hypothesized to be the ancestral state.

50. Postero-dorsal surface of scutum: [0]
mostly nitid, sometimes weakly rugulose;
[1] weakly rugulose, not shiny; [2] at least
some strong costae; costae smooth; [3]
smooth, not nitid. State [0] was observed
in Clinocentrus, Rogas, and Stiropius species
examined, but in no Aleiodes, and was hy-
pothesized to be the ancestral condition.

52. Notauli at mid-dorsal surface of scu-
tum: [0] without carinae or foveae or finely
foveate, non-carinate; [1] coarsely foveate,
and /or usually carinate posteriad; [2]
smooth, not nitid. States [1] and [2] were
not found in any Rogas or Stiropius exam-
ined. However, the coarsely foveate con-
dition was ubiquitous in Clinocentrus. Giv-
en the above, it was hypothesized that
state [0] was ancestral, and that the Cli-

nocentrus condition was derived in parallel
with the state's appearance in Aleiodes.

53. Scutal sculpturing excluding postero-
dorsal surface and notauli: [0] rugulose
sculpturing over a nitid surface; [1] gran-
ulate; [2] punctate; [3] nitid or smooth, not
nitid. States [1], [2], and [3] were not ex-
pressed by non-Aleiodes species examined.
Stiropius expressed the coarsely granulate
condition, Rogas expressed rugulose
sculpturing over a nitid surface, and Cli-
nocoitrus expressed a nitid-punctate scutal
surface. Since Rogas is considered the sis-
ter group of Aleiodes (Whitfield 1992),
state [0] was hypothesized to be ancestral.

61. Propodeum dorsal profile: [0] rounded;
[1] flattened. State [1] was not observed in
non-i4/t';o(ft's species examined. State [0]
was hypothesized to be ancestral.

63. Ratio of pronotal medial length to head
length: [0] >'0.30; [1] < 0.30. Head length
was defined as distance from the occipital
carina at the vertex to the line between
posterior edges of antennal sutures. State
[1] was observed in Stiropius and some Al-
eiodes, but not in Rogas or Clinocentrus.
Given the very small ocelli of Stiropius,
such that the ocellocular distance /ocellar
diameter ratio was consistently much
greater than one, the vertex broad in com-
parison with the three other genera, and
the antennal flagellomeres over twice as
long as wide in contrast with the three
other genera, it was hypothesized that the
Stiropius condition was independently de-
rived. Thus, it was hypothesized that state
[0] was the ancestral condition.

III. Metasomal Characters

2. Median carina of tergite II: [0] undif-
ferentiated from other coarse carinae
(Achterberg 1991, Fig. 124); [1] differenti-
ated from other carinae (Achterberg 1991,
Fig. 377); [2] undifferentiated from other
fine carinae. State [0] is the Stiropius/Rogas
condition. Clinocentrus shared [1] with
some Aleiodes. Since Clinocentrus is consid-
ered most distantly related to Aleiodes of
the three outgroup genera (Whitfield

216

Journal of Hymenoptera Research

1992), it was hypothesized that state [1]
was independently derived in Clinocentrus
and Aleiodes.

5. Median triangle of tergite U: [0] absent;
[1] small and without well developed an-
terior carinae (Achterberg 1991, Fig. 290);
[2] large and associated with anterior ca-
rinae which run laterally nearly to mar-
gins of tergite (Achterberg 1991, Fig. 300).
The presence of a median triangle differ-
entiates most Aleiodes with respect to Ro-
gas and Stiropiiis. Thus, state [0] was hy-
pothesized to be the ancestral state.

10. Length of metasomal tergite I, female:
[0] tergite apical width /tergite length
greater than or equal to 0.87 (not elon-
gate); [1] tergite apical width/tergite
length less than 0.87 (elongate). An elon-
gate first metasomal tergite was not pre-
sent in Stiropius, and only occasionally
present in Clinocentrus and Rogas. Thus, its
occasional presence in Aleiodes, Clinocen-
trus, and Rogas was hypothesized to be in-
dependently derived.

28. Setal mat on metasomal tergites 4-6 of
males: [0] absent; [1] present (Shaw et al.
1998, Fig. 1). State [1] was expressed by no
non-Aleiodes species examined. Thus, state
[0] was hypothesized to be the ancestral
state.

29. Medial pits on tergites 4-7, males: [0]
absent; [1] present (Shaw et al. 1997, Figs.
13, 15, 17) State [1] was expressed by no
non-Aleiodes species examined. Thus, state
[0] was hypothesized to be the ancestral
state.

34. Sculpturing of metasomal tergite I
(path a-c): [0] weakly or strongly costate,
costae relatively widely spaced, often over
weak granulation; [1] rugulocostate or
rugocostate; or costate, costae narrowly
spaced, no underlying sculpturing; [2]
finely rugulose to finely rugulocostate
(Shaw et al. 1997, Figs. 11, 12).

35. Sculpturing of metasomal tergite I
(path e-fl: [0] rugose or rugulose; [1] one
of conditions of #34; [2] smoothly rugose;
[3] strongly costate, costae narrowly
spaced, no underlying sculpturing; [4]

weakly costate, costae narrowly spaced,
no underlying sculpturing, or nitid.

36. Scidpturing of metasomal tergite I
(path b-h): [0] not smoothly, finely granu-
late; [1] smoothly, finely granulate. The re-
ticulate character described by the above
two paths is hypothesized as follows: a =
34[0]; b = 34[1]; c = 34[2]; d = 35[3]; e =
35[0]; f = 35[4]; g = 35[2]; h = 36[1].

abcde fgh path

n 0 12 1

1

1

1 1 a-c (#34)

n -f- 1 1 1 1 3

0

4

2 1 e-f(#35)

n+2 0 0 0 0

0

0

0 1 b-h (#36)

The above phylogenetic hypothesis is il-
lustrated in Fig. le. State (a) was observed
in all Clinocentrus and Rogas examined,
and in some Aleiodes species. The coarsely
granulate Stiropius condition was thus hy-
pothesized to be a derived synapomorphy
for that group, and (a) was hypothesized
to be ancestral.

42. Metasomal tergite III sculpturing (path
a-d): [0] completely, finely smooth-acicu-
late; [1] weakly to strongly rugulocostate
or weakly costate over granulate back-
ground; costae not smooth, tergite not
shiny; [2] smooth-aciculate or smoothly
rugulose anteriad, nitid or smooth-punc-
tate posteriad, or entirely nitid or smooth
punctate; [3] shallowly rugulose or rugu-
lopunctate anteriad, finely punctate pos-
teriad or completely finely punctate.

43. Metasomal tergite III sculpturing (path
^^S^' [0] smooth, faintly granulate, shiny;
[1] one of states of #42; [2] entirely cari-
nate; [3] strongly rugose.

44. Metasomal tergite III sculpturing (path
h-i): [0] completely smooth, shiny, no
trace of granulation; [1] one of conditions
of #42; [2] densely punctate or rugulo-
punctate anteriad; shiny posteriad, or en-
tirely densely punctate. The reticulate
character described by the above 3 paths
is hypothesized as follows: a = 42[0]; b =
42[1]; c = 42[2] d = 42[3]; e = 43[0] f =
43[2]; g = 43[3]; h = 44[0]; i = 44[2].

Volume 8, Number 2, 1999

217

a b c d e f

g h i path

n 0 12 3 11

1 2 2 a-d(#42)

n+ 1 1 1 1 1 0 2

3 1 1 e-g(#43)

n + 2 1 1 1 1 1 1

1 0 2 h-i (#44)

The ancestral state is (b). The above
phylogenetic hypothesis is illustrated in
Fig. If. Since state (a) was ubiquitous in
all Rogns species examined and in many
Aleiodes, it was hypothesized to be ances-
tral.

45. Excavated medial areas on metasomal
tergites II and III: [0] absent; [1] present
(Achterberg 1985, Fig. 12). State [1] was
not found in non-Aleiodes species exam-
ined. Thus, state [0] was hypothesized to
be the ancestral state.

46. Apex of abdomen, females: [0] not
compressed; [1] compressed. State [1] was
not found in non- Aleiodes species exam-
ined. Thus, state [0] was hypothesized to
be the ancestral state.

47. Fourth metasomal tergite (MT IV): [0]
MT IV unsculptured; [1]'mT IV slightly
sculptured; [2] MT IV mostly sculptured,
not heavily rugulose; [3] MT IV heavily
rugulose, but not a complete carapace
over apicad tergites; [4] MT IV a heavily
rugulose, complete carapace over apicad
tergites. States [I] through [4] were found
in no non-Aleiodes species examined. Thus,
state [0] was hypothesized to be the an-
cestral state.

56. Metasomal dorsum color: [0] not en-
tirely black; [1] entirely black. State [1]
was observed in some unicolored Stiropius
and Clinocentrus species, and in some Al-
eiodes species. Since all Rogas species ob-
served were uniformly yellow, and many
Aleiodes expressing state [1] had heads
and /or mesosoma that were not black, it
was hypothesized that the Stiropius and
Clinocentrus conditions were independent-
ly derived from the condition in Aleiodes,
and that state [0] was ancestral.

57. Metapleural pit: [0] not slightly larger
than propodeal spiracle, deep and round;

[1] slightly larger than propodeal spiracle,
deep and round. State [1] was not ob-
served in non-Aleiodes species examined.
Thus, state [0] was hypothesized to be the
ancestral state.

58. Head/mesosoma/apical metasomal col-
oration (metasomal tergite III and posteriad;
females): [0] not all black with bicolored
metasoma; [1] all black with bicolored me-
tasoma. State [1] was not observed in non-
Aleiodes species examined. Thus, state [0]
was hypothesized to be the ancestral state.

59. Metasomal tergite II coloration: [0] not
consistently black laterally, yellow medi-
ally; [1] consistently black laterally, yellow
medially. State [1] was not observed in
non-Aleiodes species examined. Thus, state
[0] was hypothesized to be the ancestral
state.

62. Ovipositor length: [0] less than Vi
length of metafemur; [1] greater than or
equal to V2 length of metafemur. All Sti-
ropius species examined had short ovipos-
itors, and most Rogas species and all Cli-
nocentrus species examined had long ovi-
positors. Since most Aleiodes species ex-
amined had short ovipositors, short
ovipositor length was hypothesized to be
the ancestral condition.

71 . Lateral edges of metasomal tergite I: [0]
not parallel; [1] parallel. State [1] was not
observed in non-Aleiodes species exam-
ined. Thus, state [0] was hypothesized to
be the ancestral state.

72. Metasomal tergite III: [0] not cara-
pace-like; [1] carapace-like, extending over
apicad tergites. State [1] was not observed
in non-Aleiodes species examined. Thus,
state [0] was hypothesized to be the an-
cestral state.

IV. Wing Characters

9. Second suhmarginal cell: [0] long, nar-
row, vein 2RS/vein 2-M less than 0.4; 2RS
parallel or nearly so with r-m; [1] trape-
zoidal or rectangular, defining veins often
arched, 2RS/2-M greater than 0.45 and
less than 0.71; [2] quadrate; 2RS parallel or
nearly so with r-m, and 2RS/2-M greater

218

Journal of Hymenoptera Research

than 0.71. Since the second submarginal
cells of Cliiwcentru:^, Rogas, and Stiropius
species examined all expressed state [1],
and since the ordering described above is
most logical, it was hypothesized that this
is a bipolar character

17. RS vein of hind wing (path a-ci): [0]
marginal cell widest basally, RS vein not
recurved apically (Achterberg 1991, Fig.
54); [1] marginal cell narrowest in middle;
RS vein moderately to slightly recurved
apically, never nearly touching wing mar-
gin (Achterberg 1991, Fig. 278); [2] RS vein
parallel with costal wing margin in basal
1/6-%, abruptly angling posteriad apically;
[3] marginal cell not narrowest in middle;
RS vein not recurved or angling; straight
or nearly (Achterberg 1991, Fig. 291).

18. RS vein of hind wing (path b-e): [0]
not extremely narrow in middle; vein not
nearly touching wing margin; [1] extreme-
ly narrow in middle, vein nearly touching
wing margin (Achterberg 1993a, plate 31).
The reticulate character described by the
above two paths is hypothesized as fol-
lows: a = 17[0]; b = 17[1]; c = 17[2]; d =
17[3]; e = 18[1].

a b c d e path

n 0

1

2

3

1 a-d (#17)

n -H 1 0

0

0

0

1 b-e (#18)

State (a) was observed in Clinocentnis,
Rogas, and Stiropius species examined, but
not in any Aleiodes. Thus, it was hypothe-
sized to be the ancestral state. State (b)
was hypothesized to be the most basal
state for this character within Aleiodes,
since it represents only one modification
from the ancestral state: the recurvation of
the apical section of the RS vein. States (c)
and (d) were hypothesized to represent a
progressive straightening of the RS vein
from the original curved condition. State
(e) was hypothesized to represent a bifur-
cation in the evolution of this character,
such that the (b) condition evolved in
some Aleiodes lineages into the straight-
ened (c) and (d) conditions, and on the

other hand into the extremely recurved (e)
condition in another. The above phyloge-
netic hypothesis is illustrated in Fig. lb.

26. Wing coloration: [0] fumate; [1] hya-
line; [2] patterned. State [1] was universal
for all Clinocentnis, Rogas, and Stiropius
species examined, and most Aleiodes spe-
cies. States [0] and [2] were not observed
in any non-Aleiodes species examined. Giv-
en the ubiquity of state [1], were non-hy-
aline wings to occur in non-Aleiodes spe-
cies, they would almost certainly represent
a parallel derived condition. Thus, state
[1] was hypothesized to be the ancestral
state.

30. Hindwing vein lA: [0] not extending
past vein cu-a; [1] extending past vein cu-
a. State [1] was found in no non-Aleiodes
species examined. Thus, state [0] was hy-
pothesized to be the ancestral state.

60. Fore wing width divided by length: [0]
less than 0.29 (long, narrow); [i] 0.29-0.34;
[2] greater than or equal to 0.35 (broad).
States [0] and [2] were not observed in
non-Aleiodes species examined. State [1]
was hypothesized to be ancestral; thus, the
character was hypothesized to be bipolar.

V. Leg Characters

0. Inner apex of hind tibia: [0] with setae
normal and unmodified; [1] with setae
flattened coming to a point apically, form-
ing an apical fringe (Achterberg 1991, Fig.
122). State [1] is the norm for Rogas, but
does not occur in any Stiropius or Clino-
centrus species so far as known (Shaw
1993). It was considered to be derived in
Rogas and in any Aleiodes in which it oc-
curred. Thus, state [0] was hypothesized
to be the ancestral state.

1. Tarsal claiv: [0] either no tarsal teeth
or if present, preapical lobe present (Ach-
terberg 1991, Figs. 123, 322); [1] no preap-
ical lobe, with basal tarsal teeth only; [2]
no preapical lobe, tarsal teeth extending
almost to apical tooth, or tarsi completely
pectinate (Fig. 2c; Shaw et al. 1997, Figs.
20-22). Since the presence of tarsal teeth
with no preapical lobe is an autapomor-

Volume 8, Number 2, 1999

219

^c S.A. melanoplenis ^^^^^Kf^mnMSJ^ [<' 1^' ^''Smator |

Fig. 2. Tarsal claws, a, "new species 21;" b, A. nlbitihui; c, ,4. iiichiin'plcnis; d, A. ^tignmtor.

phy for genus Aleiodes distinguishing it
from Clinoccntnis, Rogas, and Stiropins,
state [0] in part defines the hypothetical
ancestor. Thus, state [0] was hypothesized
to be the ancestral state for these four gen-
era, and state [1] was hypothesized to be
the ancestral state for Aleiodes.

11. Tarsal segment 4, female: [0] elongate:
length > 1.6 times apical width; [1] com-
pact: length < 1.6 times width. State [1]
was not observed in non-Aleiodes species
examined. Thus, state [0] was hypothe-
sized to be the ancestral state.

37. Lightness of hind tarsus: [0] not con-
siderably lighter than tibia; [1] consider-
ably lighter than tibia (Shaw et al. 1997,
Fig. 24). State [1] was not found in non-
Aleiodes species examined. Thus, state [0]
was hypothesized to be the ancestral state.

64. Number of tarsal ehnv teeth: [0] none;

[1] 5 or less; [2] more than 5. State [0] was
shared by some Rogas, all Clinocentrus and
Stiropius, so far as known, and was hy-
pothesized to be the ancestral state.

65. Shape of apical tarsal tooth: [0] not
present; [1] incompletely toothed; [2] com-
pletely toothed (Fig. 2c; Shaw et al. 1997,
Figs. 19-22). The incomplete form was
reminiscent of a thickened seta, with a
hair-like, flexible apex. State [0] was hy-
pothesized to be ancestral, for reasons
stated for character 64.

66. Setae between apical tarsal tooth and
clazo: [0] present (Fig. 2a, b); [1] absent
(Fig. 2c, d). Several Aleiodes species which
expressed incompletely toothed apical tar-
sal teeth, also expressed basal tarsal teeth
only. In acidition, they expressed a gra-
dation of thickened, prominent setae
which became progressively thinner with

220

Journal of Hymenoptera Research

increasing apical position apicad of the
apical tooth. Thus, it was hypothesized
that state [1] represented the completion
of a transformation of apical setae into
teeth, and thus that state [0] was ancestral.

67. Shape of apical tarsal claw (path a-c):
[0] nearly without a hook (Fig. 2a); [1] not
abruptly hooked (Fig. 2b); [2] abruptly
hooked with straight shank which angles
at nearly 90 degrees (Fig. 2c).

68. Shape of apical tarsal claw (path b-d):
[0] one of above states; [1] abruptly
hooked with straight shank which angles
at less than 90 degrees (Fig. 2d). The retic-
ulate character described by the above two
paths is hypothesized as follows: a =
67[0]; b = 67[1]; c = 67[2]; d = 68[1].

abed path

n

0

1

2

1

a-c (#67)

n + 1

0

0

0

1

b-d (#68)

State (b) was universal in all non-Aleio-
des species examined, and most Aleiodes
species. Thus, it was hypothesized to be
the ancestral state. The above phylogenet-
ic hypothesis is illustrated in Fig. Ih.

RESULTS

Throughout the development of this
analysis, species-groups emerged and de-
fined themselves by consistently cluster-
ing on the most parsimonious consensus
tree. Once all species had been analyzed,
18 well-defined species-groups had
emerged. Figs. 6 to 8 show the major spe-
cies-groups within genus Aleiodes. Fig. 5
shows the overall phylogenetic pattern of
the genus, with species-groups represent-
ed by exemplar species.

Initially assigning equal weight to all
characters resulted in one most parsimo-
nious tree using the mhennig* option. This
tree had a length of 475 steps, a consisten-
cy index (c.i.) of .25, and a retention index
(r.i.) of .78. Extended branch swapping
(bb* option) resulted in 755 equally par-
simonious trees all with lengths of 475
steps.

The final solution for successive itera-
tions to character weighting applied to the
trees before the Nelson consensus was run
resulted in 755 most parsimonious trees,
each with length of 624 steps, a c.i. of .56,
and an r.i. of .91. The modified Nelson
consensus tree of the final solution for the
overall unweighted analysis, after manual
branch swapping, is shown in Fig. 5. The
Nelson consensus tree length of the un-
weighted tree loaded into CLADOS was
reduced by 140 steps by manual branch
swapping.

The genus can be divided into three ma-
jor sections: basal, intermediate, and de-
rived species. Basal species are defined as
those in which 1) the occipital carina is
usually weak or effaced medially, 2) hind-
wing RS vein is recurved, 3) hind-wing
vein lA usually does not extend past vein
cu-a, 4) propodeal sculpturing is coarsely
granular, or rugulocostate over a granu-
late surface, 5) mesopleuron is granulate,
6) mesopleural pit posteriad to mesopleu-
ron central disk is absent, and 7) median
triangle of tergite II is small or absent and
without well developed anterior carinae.
In the overall cladogram (Fig. 3), groups
1-6 are basal groups.

Intermediate species are defined as
those with the above characteristics except
1) the occipital carina is complete medi-
ally, 2) propodeal sculpturing is rugose, 3)
the hind-wing RS vein may be straight,
and 4) mesopleuron may be rugulose
(groups 7-9).

Derived species are defined as those
with the characteristics for intermediate
species except 1) mesopleuron sculpturing
is smooth punctate or smooth, 2) the hind-
wing RS vein is almost always straight, 3)
third metasomal tergite is either smooth-
aciculate or smoothly rugulose or rugulo-
costate anteriad and either smooth-shiny
or smoothly rugulose posteriad, or entire-
ly smooth or entirely punctate 4) hind-
wing vein lA extends past vein cu-a, and
5) the antero-medial triangle of metasomal
tergite II is large and associated with an-

Volume 8, Number 2, 1999

221

terior carinae which run laterally. There is
an increasing tendency with derived po-
sition on the cladogram for the mesopleur-
al pit posteriad to mesopleuron central
disk to be present (groups 10-18).

Species-group definitions

Some species in unresolved clusters, or
which did not clearly fall within any clus-
ter, were not placed within any species-
group. These species either were in the
basal section, or fell at the base of a well-
defined species-group. Such species were
A. armatus, A. arnoldi, A. cariniz'entris, A.
ferrugileti, A. jawslawensis, A. medianus, A.
modestus, A. pellucens, A. quadrum, A. ros-
sicus, A. ruficeps, A. sibiricus, A. iingularis,
and new species 7 and 9.

Species included below which will be
published later by Shaw et al. are desig-
nated as new species (number). This text
does not constitute a publication of any
new species.

1. The gressitti species-group (Fig. 4a)
consists of A. gressitti (Muesebeck) and
new species 16. This is the basal-most
group in the genus, and is defined by
propodeum and metasomal tergite I
finely, smooth-shiny granulate.

2. The compressor species-group (Fig.
4a) consists of known species A. ali-
garhensi (Quadri), A. compressor (Her-
rich-Schaffer), A. palmatus (Walley),
and new species 24. The group is
monophyletic, and defined by the fol-
lowing synapomorphies: 1) apex of fe-
male abdomen compressed, and 2)
apical tarsal tooth complete.

3. The gastritor species-group (Fig. 4b) is
a large monophyletic basal group de-
fined by pronotal medial length :£
.30% of head length. It consists of A.
biioculus Marsh, A. cantharius (Lyle), A.
dendrolimi (Matsumura), A. gastritor
(Thunberg), A. laphygmae (Viereck), A.
malacosomatos (Mason), A. noctuniiis
Telenga, A. iiolophatiae (Ashmead), A.
pallescens Hellen, A. perplexus (Gahan),
A. platypterygis (Ashmead), A. similli-

miis (Ashmead), A. stigmator (Say), A.
testaceus (Telenga), A. xanthus (Mar-
shall), and new species 3, 15, 32, and
35.

4. The circumscriptus species-group (Fig.
6c) is a basal, monophyletic group de-
fined by metasomal tergite II consis-
tently yellow medially and black lat-
erally. Species within the group are: A.
aiitographae (Viereck), A. borealis
Thomson, A. circumscriptus (Nees von
Esenbeck), A. nigricornis Wesmael, A.
pictiis (Herrich-Schaffer), A. similis
(Curtis), A. scrutator (Say), and new
species 38.

5. The pallidator species-group (Fig. 4d),
also basal and monophyletic, is de-
fined by eye diameter greater than
2.5X ocellocular space, and tarsal
claws pectinate. The species in this
group are A. indiscretus (Reardon), A.
lymantriae (Watanabe), A. martini
Marsh and Shaw, A. pallidator (Thun-
berg) and new species 37.

6. The seriatus species-group (Marsh et
al. 1998; Fig. 4e) is basal and mono-
phyletic and is defined by hind tibial
apex with flattened setae forming a
fringe. The group consists of A. bakeri
(Brues), A. femoratus Cresson, A. kus-
litzkyi Tobias, A. nigribasis (Enderlein),
A. nigristemmaticum (Enderlein), A.
sanctivincentensis (Shenefelt), A. percur-
rens (Lyle), A. scriptus (Enderlein), A.
seriatus (Herrich-Schaffer), and new
species 30 and 33.

7. The procerus species-group (Fig. 5a),
which is monophyletic, consists of A.
angustatus (Papp), A. crassipes Telenga,
A. granulatus (De Gant), A. narangae
(Rohwer), and A. procerus Wesmael,
and new species 1. It is closely related
to the A. dispar species-group. The
group shares the following synapo-
morphies: 1) metasomal tergite 4
slightly sculptured, and 2) metasomal
tergite 1 elongate. All except A. cras-
sipes and A. angustatus share a long,
shelflike pronotum.

222

Journal of Hymenoptera Research

ANCESTOR

^^^^f '6 A. gressilli species-eroiin

PALMATUS N

-ALIGARHE Ce H No

K,

— PELLUCEN

— ROSSICUS D G L Ly N Py
NEWSP 35
NEWSP 19

r- TESTACEU
">— XANTHUS

— DENDROLI L
CANTHARI G

I_Hmalacoso l

NEWSP 3 N

STIGMATO G L Ly N
GASTRITO C d G I No
PALLESCE N No
I— NEWSP 7

KUZLITZK

PERCURRE N Nc

Ur— SERIATUS A Uo N
I L FEMORATU
NEWSP 30

— SCRIPTUS

— NIGRIBAS
NIGRICOR G N PY
NEWSP 36 G

A. compix.ssor
spetics-groiip

"— 1-GAST
LpALL

A. };a.\thior
species-tjroup

_ A. seriatus

spccics-giuiip

-AUTOGRAP N

-CIRCUMSC B G Ly N No Ps Py t

-SIMILIS

PALLIDA! G GE LY N PS T
NEWSP 18
LYMANTRI Ly

NEWSP 9

- ARMATUS G N

PROCERUS
CRASSIPE

NEWSP 12

NEWSP 1
ANGUSTAT
NEOTROPI
PUNCTIPE

Host Legend

Arctiidae

Bombycidae

Catocallnae

Choreutidae

Drepanidae

Gelechlidae

Geometrldae

Incurvarildae

Lasiocampidae

Limacodldae

Lycaenidae

Lymantrildae

Noctuldae

Notodontidae

Nymphalidae

Psychidae

Pterophoridae

Pyralidae

Sphingidae

Tortrlcidae

Zygaenldae

■ A. circuniscrijiiiii
spccics-gioup

- A. iHillidaior
spccics-gioui)

L- A. procerus
species-gioup

CURTISPI

- PLEURILI

- ANNUUTU

- TRANSVER
-ROGEZENS

-A. dispor
species-tfoup

I DISPAR

'l-TATIANAE

r— RUGOSICO

•— BREVIPEN

A

B

Nc

C

D

Ge

G

I

L

Li

Lc

Ly

N

No

Ny

Ps

P

Py

S

T

Z

Fig. 3. Cladogram showing overall phylogenetic pattern of Akwdes. Species-groups are represented by ex-
emplar species. Hosts and respective codes are listed in legend. Host associations for Aleioiies species are
represented by codes next to species.

The coxalis species-group (Fig. 5b) is
an intermediate monophyletic group
defined by: 1) rugulose to rugose me-
sopleuron, and 2) vertex with widely
separated laterally running ridges.
The following species fall in this
group: A. aciculatus Cresson, A. alter-

nator (Nees von Esenbeck), A. arcticus
Thomson, A. bicolor (Spinola), A. cox-
alis (Spinola), A. coxator Telenga, A.
helleniciis Papp, A. incertoides Telenga,
A. incertus Kokoujev, A. jakolezvi Ko-
koujev, A. mohiavicus Tobias, A. iwga-
tivus Tobias, A. nunbergi Noskiewicz,

Volume 8, Number 2, 1999

223

-{l

NEWSP 5
r— NEWSP 36

NEGATIVU N

L Ly

{'

HELLENIC

1 TRISTIS

L_ SATANAS
"NEWSP 10 Lc
■ ALTERNAT A
r- INCERTOI

-NEWSP 2 Ly
- NEWSP 13
~\- BICOLOR O Lc Ly N Ny P Z
COXALIS
■ MOLDAVIC

— JAKOWLEW
- TETRASPH

- NEWSP U G
-ARCTICUS

- FUSCIPEN

-PRAETOR S
TEXANUS S
GRAPHICU S

UNGULARI

MEDIANUS

CAUCASIC
p PULCHRIP L Ly N No
I — CA2IERI

■ PEDALIS

- ANTENNAT

-VAUGHANI N

CAMERONI No

GEOMETRA G
ROSSI
RILEY! Ly N Py

GRANDIS N

FLAVITAR N
P MOLESTUS N
-ABDOMINA N

' — ATRICEPS
CONVEXUS

pPR>

M

L

U

I— SIBIRICU

FERRUGIL
r TERMINAL N
I ^ HARRIMAN
_ L UNIPUNCT

GRASSATO
r-TURKESTA

EURINUS
NEWSP 31
— PALLIDIS

-I| ATRICORN N

I NEWSP 28 N

BURRUS N
_ NEWSP 4

-c

A . coxalls
species-group

,) albillhia
species-group

.'). praetor
species-group

A. pulchripes
species-group

A. ductor
species-group

A schirjajewi
species-group

A . gaslerator
species-group

RUFIPES

Fig. 3. Continued.

224

Journal of Hymenoitera Research

UFEI N
r- MORIO N

r-RUGULOSU N

■DIVERSUS N
DISSECTO N

-■ A. ufei species-group

''\^ A. rugulo.sus

species-group

^

■ MICROCUL ^

SHESTAKO

NEWSP 23

h:

NEWSP 22

-FLAVISTI

MELANOPT

LUCIDUS
MINIATUS

I FAHRINGE

c

^

— DESERTUS
AGILIS

AESTUOSU N
NEWSP 21
GLABER

Fig. 3. Continued.

— A. mekmopterus
species-group

A. sanctihyacinthi (Provancher), A. sa-
tauus Telenga, A. signatus (Nees von
Esenbeck), subgenus Tetrasphaeropyx
Ashmead; A. tristis Wesmael, and new
species 2, 5, 10, 11, 13, 14, 17, 26, 27,
34, and 36.
9. The dispar species-group (Fig. 5c), a
monophyletic group, consists of A.
aleutacens Granger, A. annulatus
Granger, A. hrevipeiidulatits Achter-
berg, A. breviradialis Granger, A. cur-
tispina Granger, A. dispar (Curtis), A.
diversicornis Granger, A. excavatus (Te-
lenga), A. longicornis Granger, A. lon-
gipendulatus Achterberg, A. neotropical-
is (Shenefelt), A. perinetensis Shenefelt,
A. phmlineatus (Cameron), A. punctipes
Thomson, A. rogezensis Granger, A. rii-
gosicostalis Achterberg, A. iatianae (Te-
lenga), and A. transversestriatus Grang-
er. It is defined by the narrow fore-
wing, which has a length /widest
width ratio < 0.29.
10. The albitibia species-group (Shaw et
al. 1998b; Fig. 6b), consisting of A. al-
bitibia (Herrich-Schaffer) and A. fiisci-
pennis (Szepligeti), is defined by nitid

mesopleuron. This character state also
occurs in A. pulchripes Wesmael,
which falls well outside of this group.

11. The praetor species-group (Shaw ei al.
1998b; Fig. 6b) is monophyletic and
consists of A. graphicus (Cresson), A.
praetor (Reinhard), A. texauus Cresson,
and new species 8 and 29. The group
is defined by: 1) the strongly recurved
hindwing radiellen vein which nearly
touches the costal margin; 2) lateral
ocellar diameter greater than 2.5 times
ocellocular distance; and 3) short 15th
flagellomere.

12. The apicalis species-group (Shaw et al.
1998a; Fig 6a) is a derived monophy-
letic group.

13. The pulchripes species-group (Shaw et
al. 1997; Fig. 6c) is a derived mono-
phyletic group.

14. The gasterator species-group (Fig. 7a)
is paraphyletic, and it is the basal spe-
cies-group within the derived species-
group cluster of the derived section.
This species-group cluster is defined
by clypeus abruptly edged, flat vcn-
trad (69[1]), and is composed of the A.

Volume 8, Number 2, 1999

225

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those underneath represent character states of taxa distal of the boxes.

226

Journal of Hymenoptera Research

Fig. 5. Species-groups within the interniedi.ite section of AIcimIcs.

Volume 8, Number 2, 1999

227

4 20SJ
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228

Journal of Hymenoptera Research

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Fig. 7. The base of the apical species-group cluster of the derived section.

Volume 8, Number 2, 1999

229

r-»-»-lH)-RUFICEPS

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230

Journal of Hymenoptera Research

gasterator (Jurine), A. rugulosus (Nees
von Esenbeck), and A. melanoptenis
(Erichson) species-subgroups.
The A. gasterator species-group is com-
posed of A. arnoldi Tobias and new species
31 basally, and three species subgroups:
the A. atricornis (Cresson), A. gasterator
(Jurine), and A. unipunctator (Thunberg)
species-subgroups. The group does not
easily cluster based on any good synapo-
morphy. The retention of tarsal claw teeth
basal only in the A. gasterator species-
group distinguishes it from species-
groups apicad to it. Although three spe-
cies clusters exist within the A. gasterator
species-group, they were considered sub-
groups, since they 1) clustered based on
characters which reversed within the spe-
cies-group or subgroup {A. atricornis spe-
cies-subgroup); 2) were based on a color
character, which are notoriously plastic
across the genus {A. unipunctator species-
subgroup), or 3) resulted in a paraphyletic
group, such as the A. unipunctator species-
subgroup, in which A. eurinus shares char-
acter states which define both the A. uni-
punctator and A. gasterator species-groups.

15. The ufei species-group (Fig. 8) is com-
posed of one species, so far as known:
A. ufei (Walley). The bicolored body is
similar to the A. unipunctator species-
subgroup, but the species differs in
that the sides of the first metasomal
tergite are parallel, and the female ab-
dominal apex is compressed. Al-
though this group is monospecific so
far as known, it was given species-
group status based on the unique me-
tasomal features described above.

16. The rugulosus species-group (Fig. 8a)
is monophyletic, and is defined by the
uniquely shiny, coarsely and densely
rugulopunctate vertex sculpturing
and all black body. It consists of A.
cruentus (Nees von Esenbeck), A. dis-
sector (Nees von Esenbeck), A. diversus
(Szepligeti), A. microculatus Watanabe,
A. morio (Reinhard), A. periscelis (Rein-

hard), A. rugulosus (Nees von Esen-
beck), and A. shestakovi (Shenefelt).

17. The melanoptenis species-group (Fig.
8b) is monophyletic and is defined by
the clypeo-antennal space /width of
oral space ratio < 0.69, scutal sculp-
turing excluding postero-medial dor-
sal surface smooth or finely punctate,
and clypeus height/ width less than
0.42. Members of the group are A. aes-
tuosus (Reinhard), A. agilis (Telenga),
A. desertus (Telenga), A. fahringeri (Te-
lenga), A. flavistigma Shaw, A. glaber
(Telenga), A. krulikoivskii Kokoujev, A.
lucidus (Szepligeti), A. melanopterus
(Erichson), A. mexicanus Cresson, A.
tnandihularis (Cresson), A. niiniatus
(Herrich-Schaffer), A. politiceps (Ga-
han), A. ruficeps (Telenga), A. venustu-
lus (Kokoujev), A. wadai (Watanabe),
and new species 21, 22, and 23.

17a. The melanopterus species-subgroup
(Shaw 1993, Fig 8b), a monophyletic
subset of the above, consists of A. flavi-
stigma Shaw, A. lucidus (Szepligeti), A.
mandihularis (Cresson),. A. melanopte-
rus (Erichson), A. mexicanus Cresson,
A. politiceps (Gahan) and new species
23. The subgroup is defined by pecti-
nate tarsal claws (Fig. 2c) and strongly
protruding clypeal carina.

DISCUSSION OF HOST ASSOCIATION

The following data and that in Appen-
dix 1 were drawn from Shenefelt (1975),
Shaw (1983, 1994), and S. R. Shaw and P.
M. Marsh (unpublished data). Of 208 Al-
eiodes species analyzed, host records were
available for 40.4% of Aleiodes species.
Host records were available for 65.4% of
basal species, 27.3% of intermediate spe-
cies, and 39.4% of derived species (Appen-
dix 1).

Basal species for which host associations
were known either attacked host species
from more than one family, or attacked
hosts other than non-catocaline Noctui-
dae, Notodontidae, or Sphingidae. Excep-
tions were A. gossi/pii, A. laphygmae, A. noc-

Volume 8, Number 2, 1999

231

turniis, and A. nolophanae. Two basal spe-
cies in the A. gastritor species group (A.
pallesceus and A. stigmator) are gregarious.

Within intermediate species, the A. cox-
alis species-group showed a tendency to
attack setose hosts within Arctiidae, Lasio-
campidae, and Lymantriidae. Aleiodes cox-
alis species-group species comprise 50.0%
of Aleiodes species known to attack arctiids
and lymantriids. Of the 15 intermediate
species with host records, 40% attack ex-
clusively non-catocaline noctuids so far as
known.

Within the derived section apicad of the
A. pulchripes and A. apicalis species groups,
only trifine Noctuidae are attacked so far
as known. The A. praetor species-group at-
tacks only sphingids. The A. albitihin spe-
cies-group attacks only notodontids. With-
in the A. pulchripes species-group, geome-
trids, noctuids, and notodontids are at-
tacked, so far as known.

Of the seven of 11 A. apicalis species-
group species with known hosts, four at-
tack plusiine noctuids (A. brethesi, A. api-
calis, A. molestus, and A. parasiticus). One
is associated with an unidentified noctuid
mummy {A. abdominalis), and three attack
non-noctuids.

Of all derived Aleiodes with host re-
cords, 69.7% attack trifine noctuids. Of the
above except the A. albitibia and A. praetor
species groups, 84.6% attack trifine noc-
tuids.

Basal and intermediate species tend to
have broad host ranges, and derived spe-
cies tend to specialize on Noctuidae, No-
todontidae, or Sphingidae. The only
monophyletic groups which specialize on
Notodontidae and Sphingidae lie at the
base of the apical section. In the basal sec-
tion, the A. pallidator species-group tends
to specialize on Lymantriidae. The only
Aleiodes species with host records for Ca-
tocalinae are in the A. seriatus species-
group in the basal section. Among inter-
mediate species, the A. coxalis species-
group tends to specialize on setose hosts,
especially arctiids and lymantriids.

According to the classification scheme
of Nielsen and Common (1991), Noctuidae
is the most derived lepidopteran family.
The lepidopteran families Notodontidae,
Lymantriidae, and Arctiidae are placed in
superfamily Noctuoidea with Noctuidae.
Notodontidae is the most basal noctuoid
family (Nielsen and Common, 1991). Su-
perfamily Sphingoidea, consisting of one
family (Sphingidae), is immediately basal
to Noctuoidea (Nielsen and Common,
1991).

Based on order of placement in a list of
19 subfamilies of Noctuidae, subfamily
Catocalinae is placed within the four bas-
al-most subfamilies (Nielsen and Com-
mon, 1991). Catocaline noctuids are uti-
lized only by one Aleiodes species-group in
the basal section, so far as known.

Family Arctiidae is placed just basad to
the noctuid subfamilies referred to in the
study (Nielsen and Common 1991). The
only basal Aleiodes species known to attack
arctiids are within the A. seriatus species-
group.

Mitchell et al. have recently investigated
the phylogeny of superfamily Noctuoidea
by using characters derived from the nu-
clear gene EF-la, which encodes elonga-
tion factor- la protein (Mitchell et al. 1997).
Based on their most parsimonious tree de-
rived from all sites of EF-la, the authors
concluded that 1) Notodontidae comprises
a monophyletic group at the base of Noc-
tuoidea; 2) "quadrifine" Noctuidae com-
prise a paraphyletic group which consists
of a monophyletic subgroup (Eutellinae,
Nolinae, Sarrothripinae), and the basal-
most subfamilies (Catocalinae, Hermini-
inae) of a more derived monophyletic
group. Arctiidae and Lymantriidae are sis-
ter-groups at the apex of this latter mono-
phyletic group; 3) the "trifine" noctuids,
which comprise the remainder of Noctui-
dae, form a monophyletic group at the
apex of the cladogram (Mitchell et al.
1997).

The findings of Mitchell et al. corrobo-
rate Nielsen and Common with respect to

232

Journal of Hymenoptera Research

Noctuoidea, except that Mitchell et al. find
that Noctuidae is a paraphyletic group,
with the quadrifine noctuids (including
Catocalinae) more closely related to Arc-
tiidae and Lymantriidae than to trifine
noctuids (Mitchell et al. 1997). The latter
finding is corroborated by Weller et al.
(1994).

Since older koinobiont parasitoid spe-
cies are thought to have broader host
ranges while newer species are thought to
have narrower ranges (Shaw 1994), and
since this study provides evidence that
basal and intermediate Aleiodes species
have broader host ranges while derived
species have narrower host ranges, the
overall pattern of phylogenetic inferences
made from morphological data in this
study is corroborated by host association
data in light of the above.

This study suggests that genus Aleiodes
overall has co-evolved with its Lepidop-
teran hosts. Basal host families appear
more likely to be attacked by basal Aleio-
des species while the most derived host
family, trifine Noctuidae, is more likely to
be attacked by derived Aleiodes species.
Non-derived groups with well defined
synapomorphies such as the basal A. ser-
iatus and A. pallidator species groups, and
the intermediate A. coxalis species-group,
appear to have co-evolved with noctuoid
groups basal to trifine Noctuidae such as
catocalines, lymantriids and arctiids. The
exclusive associations of Aleiodes species
within the derived section with specific
host groups, such as that of the A. praetor
species-group with sphingids, of the A. al-
bitibia species-group with notodontids,
and of Aleiodes species apicad of the A. ap-
icalis and A. pulchripes species-groups with
Noctuidae add weight to this co-evolu-
tionary hypothesis.

ACKNOWLEDGMENTS

The following collections provided specimens for
this study. The North American collections cited in
Shaw et al., 1997, The Koninklijk Museum voor Mid-
den-Afrika, Belgium, The Museum of Natural His-

tory, Warsaw, Poland, the insect collection of Lund
University, Lund, Sweden, The Natural History Mu-
seum, London, England, The Natural History Muse-
um of Paris, France, The Natural History Museum of
Vienna, Austria, The Royal Institute of Natural Sci-
ences of Belgium, Brussels, The Natural History Mu-
seum of Hungary, The Museum of Natural History,
St. Petersburg, Russia. This research was supported
by grant DEB-930-6314 from the National Science
Foundation. Additional support was provided by a
Canadian National Collection grant and by Univer-
sity of Wyoming summer stipends.

LITERATURE CITED

Achterberg, C. van. 1982. Notes on some type-species
described by Fabricius of the subfamilies Bracon-
inae, Rogadinae, Microgastrinae, and Agathidi-
nae (Hymenoptera: Braconidae). Eutonwliigishdw
Benchtcn, 42: 133-139.

Achterberg, C. van. 1985. Notes on Braconidae I-IV.
Zoolcgische Mededclingen 59: 163-187.

Achterberg, C. van. 1991. Revision of the genera of
the Afrotropical and W. Palearctic Rogadinae
Foerster (Hymenoptera: Braconidae). Zcologische
Vfrhnmielingen 273: 3-102.

Achterberg, C. van. 1993. Illustrated key to the sub-
families of the Braconidae (Hymenoptera: Ich-
neumonoidea). Zoohgische Verhaudclingen 283: 3-
189.

Achterberg, C. van. 1993 Generic revision of the sub-
family Betylobraconinae (Hymenoptera: Bracon-
idae) and other groups with modified fore tarsus.
Zoclogischc Verlmndelingen 298: 3-242.

Achterberg, C. van and A. M. Penteado-Dias. 1995.
Six new species of the Alcwda di-ipar group (Hy-
menoptera: Braconidae: Rogadinae). Zoologhclie
Mededeiingen Leiden 69: 1-18.

Farris, J. S. 1970. Methods for computing Wagner
trees. Systematic Zoology 19: 83-92.

Farris, J. S. 1988. Hcnnig86 reference, version 1.5. Port
Jefferson, New York.

Harris, R. A. 1979. A glossary of surface sculpturing.
Occasional Papers in Entomology 28: 1-33.

Hennig, W. 1966. Phylogenetic Systematics. John Wiley
and Sons. New York.

Hennig, W. 1981. Insect Phytogeny. John Wiley and
Sons. New York.

Huber, J. and M. Sharkey. 1993. Chapter 3. Structure.
In: Hymenoptera of the World: An Identification
Guide to Families (ed. by H. Goulet and J. T.
Huber). Agriculture Canada Publication 1894/ E.
Ottawa.

Lipscomb, D. L. 1992. Parsimony, Homology and the
Analysis of Multistate Characters. Cladistics 8:
45-65.

Marsh, P. 1979. The braconid (Hymenoptera) para-
sites of the gypsy moth, Lymantria dispar (Lepi-

Volume 8, Number 2, 1999

233

dopfera: Lymantriidae). Annals of the Entomolog-
ical Society of America 72: 794-810.

Marsh, P. M. and S. R. Shaw 1998. Revision of North
American Aleiodes Wesmael (Part 3): the seriatus
species-group in the New World (Hymenoptera;
Braconidae, Rogadinae). Proceeiiings of the Ento-
mological Societ}/ of Washington 100: 395—408.

Mickevich, M. F. 1982. Transformation series analysis.
Systematic Zoology 31: 461—178

Mickevich, M. F. and D. Lipscomb. 1991. Parsimony
and the choice between different transformations
for the same character set. Claiiistics 7: 111-139.

Mickevich, M. F. and S. J. Weller. 1990. Evolutionary
character analysis: tracing character change on a
cladogram. Cladistics 6: 137-170.

Mitchell, A., S. Cho, J. C. Regier, C. Mitter, R. W.
Poole, and M. Matthews. 1997. Phylogenetic util-
ity of elongation factor-la in Noctuoidea (Insec-
ta: Lepidoptera): The limits of synonymnous sub-
stitution. Molecular Biology and Evolution 14(4);
381-390.

Mitter, C. and E. Silverfine. 1988. On the systematic
position of Catocala Schrank (Lepidoptera: Noc-
tuidae). Systematic Entomology 13: 67-84.

Nielsen, E. S. and 1. F. B. Common. 1991. Chapter 41.
Lepidoptera. In: The Insects of Australia. Cornell
University Press, Ithaca, New York.

Nixon, K. C. 1991. CLADOS Version 1.0. Cornell Uni-
versity, Ithaca, New York.

Shaw, M. 1983. On(e) the evolution of endoparasit-
ism: the biology of some genera of Rogadinae
(Braconidae). Contributions of the American Ento-
mological Institute 20: 307-328.

Shaw, M. 1994. Chapter 7. Parasitoid host ranges, in
Parasitoid Community Ecology, (ed. by B.A. Haw-
kins and W. Sheehan). Oxford University Press,
Oxford, Great Britain.

Shaw, M. and T. Huddleston. 1991. Classification and
biology of braconid wasps. Handbooks for the Iden-
tification of British Insects Vol. 7, Part II 7: 1-126.

Shaw, S. R. 1993. Systematic status of Eucystomastax

Brues and characterization of the Neotropical
species, journal of Hymenoptera Research 2: 1-11.

Shaw, S. R. 1995. Chapter 12.2. Braconidae. In: The
Hymenoptera of Costa Rica (ed. by P. Hanson and
I. D. Gauld). Oxford University Press, Oxford,
Great Britain.

Shaw, S. R., P. M. Marsh, and J. C. Fortier. 1997. Re-
vision of North American Aleiodes Wesmael (Part
1): the flai'idus Cresson species-group in the New
World (Hymenoptera: Braconidae, Rogadinae).
journal of Hymenoptera Research 6: 10-35.

Shaw, S. R., P.M. Marsh, and J. C. Fortier. 1998a.
Revision of North American Aleiodes Wesmael
(Part 2): the apicalis (Brulle) species-group in the
New World (Hymenoptera: Braconidae, Roga-
dinae). Journal of Hymenoptera Research 7: 62-73.

Shaw, S. R. and P. M. Marsh. 1998b. Revision of
North American Aleiodes Wesmael (Part 4): the
praetor and atbitibia groups in the New World
(Hymenoptera:Braconidae, Rogadinae). Proceed-
ings of the Entomological Society of Washington 100:
553-565.

Shenefelt, R. D. 1969. Notes on some rogadine genera
(Hymenoptera: Braconidae). Proceedings of the En-
tomological Society of Washington 71: 428-444.

Shenefelt, R. D. 1975. Braconidae 8. Exothecinae, Ro-
gadinae. Hymenopterorum Catalogiis (nova editio)
12: 1163-1262.

Schuh, R. T. 1991. Phylogenetic, host and biogeo-
graphic analyses of the Pilophorini (Heteroptera:
Miridae: Phylinae). Cladistics 7: 157-189.

Weller, S. J., D. P. Pashley, J. A. Martin, and J. L.
Constable. 1994. Phylogeny of noctuoid moths
and the utility of combining independent nuclear
and mitochondrial genes Systematic Biology 43:
194-211.

Whitfield, J. B. 1992. The polyphyletic origin of en-
doparasitism in the cyclostome lineages of Bra-
conidae (Hymenoptera). Systematic Entomology
17: 273-286.

Wiley, E. O. 1981. Phylogenetics; the Theory and Practice
of Phylogenetic Systematics. John Wiley and Sons,
Inc. New York.

234

Journal of Hymenoptera Research

Appendix I. List of Akiotlcf species examined.

I losl t.imil\(ifs)

abdominalis Cresson 1869
aciculatus Cresson 1869
aestuosiis (Reinhard) 1863
agilif (Telenga) 1941
albitWm (Herrich-Schaffer) 1838
alboannulalus Belokobylskyj
alutaccus Granger 1949
aligharensi (Quadri) 1933
alternator (Nees von Esenbeck) 1934
angustatus (Papp) 1969
annulatiis Granger 1949
antennatus Belokobylskyj 1996
apicalis (BruUe) 1832

arctkus Thomson 1891
arizoniensis Marsh and Shaw 1997
armatus Wesmael 1838
arnoldii Tobias 1976
atricqjs Cresson 1869
atricornis (Cresson) 1872
autograpliae (Viereck) 1910
bakeri (Brues) 1912
bicolor (Spinola) 1808

borealis Thomson 1891

brethesi (Shenefelt) 1909

breinpendulatKS Achterberg and Penteado-Dias, 1995

brei'iradialis Granger 1949

buoculus Marsh 1989

burrus Cresson 1869

cameronii (Dalla Torre) 1898

cantherius (Lyle) 1919

carinatus (Ashmead) 1888

cariniventris (Enderlein) 1912

caucasicus Tobias 1976

cazieri Marsh and Shaw 1997

chloroticus Shestakov 1940

circuniscriptus (Nees von Esenbeck) 1834

compressor (Herrich-Schaffer) 1838

confonnis (Muesebeck) 1960

convexus Achterberg 1991

coxalis (Spinola) 1808

coxator Telenga 1941

crassipes Thomson 1891

cruentis (Nees von Esenbeck) 1834

curtispina Granger 1949

dendroliwi (Matsumura) 1925

desertiis (Telenga) 1941

dispar (Curtis) 1834

dissector (Nees von Esenbeck) 1834

diversicornis Granger 1949

dhvrsus (Szepligeti) 1903

Noctuidao

Noctuidae

Noctuidae (Noctuinae, Ophiderinae)

unknown

Notodontidae

unknown

unknown

Gelechiidae, Noctuidae, Notodontidae

Arctiidae, Lasiocampidae, Lymantriidae

unknown

unknown

unknown

Arctiidae, Geometridae, Lasiocampidae, Noctuidae

(Hadeninae, Plusiinae)
unknown
unknown

Geometridae, Noctuidae
unknown
unknown

Noctuidae (Hadeninae)
Noctuidae
unknown
Geometridae, Lycaenidae, Lymantriidae, Noctuidae,

Nymphalidae, Pterophoridae, Zygaenidae
unknown

Noctuidae (Plusiinae)
unknown
unknown
unknown

Noctuidae (Acronictinae, Noctuinae)
Notodontidae
Geometridae
unknown
unknown
unknown
unknown
unknown
Bombycidae, Geometridae, Lymantriidae, Noctuidae,

Notodontidae, Psychidae, Pyralidae, Tortricidae
Gelechiidae, Geometridae, Limacodidae, Noctuidae,

Notodontidae
unknown
unknown

Hesperiidae, Satyridae
unknown
unknown

Noctuidae (Hadeninae), Lymantriidae
unknown
Lasiocampidae
unknown

Noctuidae, Nvmphalidac
Noctuidae (Acronictinae, Hadeninae)
unknown
Noctuidae (Acronictinae, Hadeninae)

Volume 8, Number 2, 1999

235

Appendix 1. Continued.

,\/i'iit(irs specifs

Host tamiK'des)

eariiiof Marsh and Shaw 1997
ecuadi'ricima (Brues) 1926
eurinus (Telenga) 1941
excavatus (Telenga) 1941
fahringeri (Telenga) 1941
femoratui Cresson 1869
ferrugilcti (Shenefelt) 1975
flai'idiis (Cresson) 1865
flavisligma Shaw 1993
flaviUnsus Marsh & Shaw 1998
fortipcf. (Reinhard) 1863
forth (Muesebeck) 1960
fuscipennis (Szepligeti) 1904
gasterator (Jurine) 1807
gastritor (Thunberg) 1822

geomctrac (Ashmead) 1888

glabcr (Telenga) 1941

gossypii (Muesebeck) 1960

grmuiif Giraud 1857

graiiK/rtfus (De Gant) 1930

graphkus Cresson 1872

grassaior (Thunberg) 1822

gressitti (Muesebeck) 1964

harrimaiii (Ashmead) 1902

helleniai!. Papp 1985

hirtiia (Thomson) 1891

iiicertoides Telenga 1941

incerlus Kokoujev 1898

(>irf;sfri'fi(S (Reardon)

jakolewi Kokoujev 1898

krulikou'skii Kokoujev 1898

kiisUtzk\/i Tobias 1976

lapln/gnme (Viereck) 1912

longiccrnis Granger 1949

longipemiiilatiif Achterberg and Penteado-Dias 1995

hichlus (Szepligeti) 1906

lymantrmc (Watanabe) 1937

mnhico^omcitos (Mason) 1979

mandibularis (Cresson) 1872

medianus (Thomson) 1896

mclanoptcrus (Erichson) 1848

mexicaniis Cresson 1869

microculntiii (Watanabe) 1937

}niiiiatiis (Herrich-Schaffer) 1838

nnidcftus (Reinhard) 1863

H;oMi!i'k»s Tobias 1986

»i()/i'sfi(s (Cresson) 1872

inoriti (Reinhard) 1863

narangac (Rohwer) 1934

negativus Tobias 1960

neotropiailis (Shenefelt) 1975

nigrilmsif (Enderlein) 1918

nigricorni^ Wesmael 1838

unknown

unknown

unknown

unknown

unknown

unknown

unknown

unknown

Unknown

Noctuidae

unknown

unknown

unknown

Noctuidae (Plusiinae)

Choreutidae, Drepanidae, Geometridae, Incurvari-

idae, Notodontidae
Geometridae
unknown
Noctuidae

Noctuidae (Acronictinae)
Noctuidae
Sphingidae
unknown
unknown
unknown
unknown
unknown
unknown
unknown
Lvmantriidae
unknown
unknown
unknown
Noctuidae
unknown
unknown
unknown
Lvmantriidae
Lasiocampidae
unknown
unknown
unknown
unknown
unknown
unknown

Geometridae, Lasiocampidae
unknown

Noctuidae (Plusiinae, Noctuinae)
Noctuidae
unknown
Noctuidae
unknown
unknown
Noctuidae, Pvralidae, Geometridae

236

Journal of Hymenoptera Research

Appendix I. Continued.

,Ut'(oi(fS species

Host tamiiv(ies)

nigristemmaticiim (Enderlein) 1918
nocturnus Tobias 1960
nohphanae (Ashmead) 1888
notozophus Marsh and Shaw 1997
luinbergi Noskiewicz 1956
pallcscens Hellen 1927
pallidator (Thunberg) 1822

pnllidistiginus (Telenga) 1941
palmatus (Walley) 1941
parasiticus Norton 1869
pedalis Cresson 1869
pellucens (Telenga) 1941
pcrcurrens (Lyle) 1921
perinetensis (Shenet'elt) 1975
perisCL'lis (Reinliard) 1863
perplexiis (Gahan) 1917
pictiis (Herrich-Schatfer) 1838
platypterygis (Ashmead) 1888
plurilineatus (Cameron) 1911
politicqjs (Gahan) 1917
praetor (Reinhard) 1863
procerus Wesmael 1838
pulchripcs Wesmael 1838

punctipes Thomson 1891
quadrum Tobias 1976
quebecensis (Provancher) 1880
rileyi Cresson 1869

rogezensis Granger 1949
rossi Marsh and Shaw 1997
rossicus Kokoiijev 1898

riificeps (Telenga) 1941
rutficoniis (Herrich-Schaffer) 1838

rufipes (Thomson) 1891

rugosicostalis Achterberg and Penteado-Dias, 1995

rugulosui (Nees von Esenbeck) 1811

sanctihyacinthi (Provancher) 1880

sanctivincentcusis (Shenefelt), 1975

satanas Telenga 1941

schirjajnoi (Kokoujev) 1899

scriptus (Enderlein) 1918

scrutator (Say) 1836

seriatus (Herrich-Schaffer) 1838

shestakovi (Shenefelt) 1975

sibiricus (Kokoujev) 1903

sigimtus (Nees von Esenbeck) 1911

similis (Curtis) 1834

simillinius (Ashmead) 1889

stigmator (Say) 1824

Noctuidae (Catocalinae)

Noctuidae

Noctuidae

unknown

Geometridae

Noctuidae, Notodontidae (gregarious)

Gelechiidae, Geometridae, Lymantriidae, Noctuidae,

Psychidae, Tortricidae
unknown
Noctuidae
Noctuidae
unknown
unknown

Noctuidae (Catocalinae, Acronictinae)
unknown
unknown

Geometridae, Noctuidae
unknown
Drepanidae
unknown

Noctuidae (Hadeninae, Noctuinae)
Sphingidae
unknown

Lasiocampidae, Lymantriidae, Noctuidae (Acronicti-
nae), Notodontidae

unknown

unknown

Noctuidae (Acronictinae)

Lymantriidae, Noctuidae (Acronictinae, Hadeninae),
Pyralidae

unknown

unknown

Drepanidae, Geometridae, Lasiocampidae, Lymantri-
idae, Noctuidae, Pyralidae

unknown

Lasiocampidae, Lymantriidae, Noctuidae (Hadeninae,
Noctuinae, Plusiinae)

unknown

unknown

Noctuidae (Acronictinae)

Arctiidae

unknown

unknown

unknown

unknown

Choreutidae, Incurvariidae, Notodontidae

Arctiidae, Lyonitiidae, Noctuidae

unknown

unknown

unknown

unknown

Geometridae

Geometridae, Lasiocampidae, Lymantriidae, Noctui-
dae (gregarious)

Volume 8, Number 2, 1999

237

Appendix I. Continued.

Host tamiiv(ies)

sudatorium Papp 1985

tatianae (Telenga) 1941

testaceus (Telenga) 1941

terminalh (Cresson) 1869

Tetrafphaeropyx Ashmead 1888 spp.

texanui (Cresson) 1869

transversestriatuf Granger 1949

tristis Wesmael 1838

turkcitanicus (Telenga) 1941

ufei (Walley) 1941

ungulnris Thomson 1891

unicolor Wesmael 1838

unipundator (Thunberg) 1822

vaughani (Muesebeck) 1960

venustulus (Kokoujev) 1905

wadai (Watanabe) 1937

xanthus (Marshall) 1892

new species 1

new species 2

new species 3

new species 4

new species 5

new species 6

new species 7

new species 8

new species 9

new species 10

new species 11

new species 12

new species 13

new species 14

new species 15

new species 16

new species 27

new species 18

new species 19

new species 20

new species 21

new species 22

new species 23

new species 24

new species 25

new species 26

new species 27

new species 28

new species 29

new species 30

new species 31

new species 32

new species 33

new species 34

new species 35

new species 36

new species 37

new species 38

unknown

unknown

unknown

Noctuidae (Acronictinae, Hadeninae, Noctuinae)

Geometridae

Sphingidae

unknown

unknown

unknown

Noctuidae (Noctuinae)

unknown

unknown

Noctuidae (Acronictinae, Hadeninae)

Noctuidae (Acronictinae)

unknown

unknown

unknown

imknown

Lvmantriidae

Noctuidae

unknown

unknown

unknown

unknown

Sphingidae

unknown

Lycaenidae

unknown

unknown

unknown

Notodontidae

unknown

unknown

unknown

unknown

unknown

unknown

unknown

unknown

unknown

unknown

Noctuidae (Cuculliinae)

Arctiidae

unknown

Noctuidae (Hypeninae)

Sphingidae

unknown

unknown

Lycaenidae

unknown

unknown

unknown

unknown

L\mantriidae

Geometridae

I. HYM. RES.
Vol. 8(2), 1999, pp. 238-250

Does the Mating System of Trissolcus basalis (Wollaston)
(Hymenoptera: Scelionidae) Allow Outbreeding?

A. D. Loch and G. H. Walter

(ADL and GHW) Department of Zoology and Entomology, The University of Queensland,

Brisbane, Queensland, 4072, Australia; (ADL Present address: CSIRO Entomology,

c/- Department of Conservation and Land Management, Brain St, Manjimup, Western

Australia, 6258, Australia)

Abstract. — The quasi-gregarious egg parasitoid Trissolcus basalis (Wollaston) is generally consid-
ered to be an entirely inbreeding species because it is a sib-mating species that has female-biased
sex ratios. Whether the species also outbreeds has not been previously investigated although
several aspects of its mating behaviour suggest this might be possible. This question was inves-
tigated indirectly in two ways by quantifying: (1) the inseminative capacity of T. basalis males in
relation to the rate of female emergence, and (2) the effects of age and mating status on sexual
receptivity of T. basalis. Trissolcus basalis females emerged over a period of several days, concen-
trating their emergence in the morning hours. Males were able to inseminate many females (>
50) in rapid succession, apparently without sperm depletion. However, approximately 20% of
females did not produce female offspring, probably because they did not mate. Although the
mated females produced proportionately more male offspring with time, this outcome is not
readily explained by sperm depletion of their mating partiiers and remains an unresolved issue.
Male sexual receptivity appears to be unaffected by age and would be expected to be unaffected
by mating status because males typically are polygynous. Although a previously successful mating
encounter did not preclude females from mating again, female sexual receptivity decreased sig-
nificantly after mating. Female sexual receptivity also decreased significantly with age. These
results suggest that both T. basalis males and females have the ability to mate away from the natal
site and that outbreeding is possible in this species. Whether males and females can locate one
another away from their own natal site therefore warrants further investigation.

Arrhenotokous parthenogenesis is the al 1993; Godfray 1994; but see Walter and

usual means of reproduction in Hyme- Clarke 1992; Ode et al. 1997). Because fe-

noptera. Female-biased sex ratios and sib- male Hymenoptera can control the fertil-

mating are characteristic of many arrhen- isation of each egg they deposit, LMC the-

otokous species, especially those v^hose ory predicts that single foundress broods

males develop in the vicinity of their fe- of gregarious and quasi-gregarious spe-

male siblings and emerge before them cies will contain only enough male off-

(protandry). This occurs most frequently spring to mate all of their sisters in the

in gregarious parasitoids, which deposit immediate vicinity (Hardy et al. 1998).
many eggs per host, and quasi-gregarious Several recent studies have shown that

ones (van den Assem et al. 1980), which strict local mating does not occur in some

lay one egg per host into hosts that are species with female-biased sex ratios and

invariably aggregated. Hamilton's (1967) which therefore should be inbreeding spe-

local mate competition (LMC) theory is cies (e.g. Myint and Walter 1990; Nadel

generally seen as the best explanation of and Luck 1992; Molbo and Parker 1996;

female-biased sex ratios (e.g. Waage and Hardy et al. 1999). The term partial local

Lane 1984; Waage and Ng 1984; Hardy et mate competition has been often used for

Volume 8, Number 2, 1999

239

such cases. Another species whose mating
behaviour appears not to be strictly local
is Trissolciis ba:^alis (Wollaston) (Hymenop-
tera: Scelionidae), a quasi-gregarious egg
parasitoid of the green vegetable bug, Ne-
zara viridula (L.) (Hemiptera: Pentatomi-
dae). Trissolcus basalis is regarded as an in-
breeding species that manifests local mate
competition (LMC) (Hamilton 1967) be-
cause it has female-biased sex ratios and
sib-mating (Noble 1937; Anon. 1939; Smith
1945; Wilson 1961; Thomas 1972). How-
ever, several aspects of its mating system
are inconsistent with LMC and indicate
that a proportion of each brood may out-
breed if they are to transmit genes beyond
the next generation. Field observations in-
dicate that nearly 20% of newly-emerged
females depart the egg mass unmated, ap-
proximately 25% of mated females were
mated more than once and often by mul-
tiple males, virgin and mated females re-
mained nearby the egg mass for up to sev-
eral hours after emergence, and males dis-
persed from the natal site (A. D. Loch and
G. H. Walter unpublished data). Given
that both males and females leave the na-
tal site in the field, it is certainly possible
that unrelated males and females meet
and mate away from their natal site.

The female mating pattern in T. basalis
may be a consequence of the males mating
many females in quick succession, and be-
coming sperm depleted. Females mated
by sperm depleted males may receive in-
sufficient sperm to produce the usual pro-
portion of daughters in a brood. They may
therefore mate again, with outbreeding
being more likely in such circumstances.
The inseminative capacity of T. basalis
males was therefore investigated in rela-
tion to the rate and sequence of female
emergence from the host patch. The ex-
periment was designed to emulate typical
rates of sibling emergence and mating in
the field, rather than the unnaturally high
rates used in some studies (e.g. Nadel and
Luck 1985). Specifically, the number (and
proportion) of females emerging from a

single egg mass and inseminated by one
male was quantified. The consequence of
female emergence position for the amount
of sperm received from the male was
quantified by recording the number and
sex ratio of progeny from every tenth fe-
male to emerge. Whether T. basalis males
become sperm depleted at mating rates
typical in the field could thus be deter-
mined.

The potential for outbreeding in T. ba-
salis was also evaluated by determining
whether males and females are sexually
receptive after mating and /or leaving the
natal site. We therefore investigated the
effects of age and mating status (virgin or
mated) on T. basalis sexual receptivity in
the laboratory, by exposing different aged
virgin and once-mated females to newly-
emerged virgin males. For completeness,
the effect of age on male sexual receptivity
was also studied by mating different aged
virgin males with newly-emerged virgin
females.

MATERIALS AND METHODS

Laboratory cultures. — Green vegetable
bugs were reared at 28 ± TC, 65 ± 10%
R.H., 16L:8D in mesh cages (0.45 m sides)
on a diet of green bean pods (Phaseolus
inilgaris L.), shelled peanuts (Arachis hy-
pogea L.) and water. Cultures were aug-
mented regularly with field-collected
bugs. Green vegetable bug egg masses
were collected daily from cages and were
used to maintain cultures of green vege-
table bug or T. basalis (see below).

Laboratory cultures of T. basalis were es-
tablished from parasitoids that emerged
from green vegetable bug egg masses col-
lected from mungbean, Vigna radiata (L.)
Wilezek, and soybean. Glycine max (L.)
Merr., during March-April 1997 and Jan-
uary-April 1998 at Pittsworth (27" 43'S,
151° 38'E), Bongeen (27° 34'S, 151° 27'E)
and Cecil Plains (27° 32'S, 151° 12'E) in
south-eastern Queensland, Australia. All
T. basalis individuals that emerged from a
single egg mass were held together in a

240

Journal of Hymenoptera Research

ventilated vial streaked with honey. Cul-
tures of T. basalis were kept at 15 ± 1°C
65 ± 10% R.H. and 16L:8D. The identifi-
cation of r. basalis was confirmed by Dr
Norman Johnson (Ohio State University).
Voucher specimens from the T. basalis cul-
ture are deposited in The University of
Queensland Insect Collection.

In all experiments, virgin wasps of the
F ,- Ft generation were used. Wasp virgin-
ity was ensured by holding single wasp
pupae in ventilated vials with honey, after
breaking the host egg mass into individual
eggs soon after parasitoid pupation. For
each experimental replicate, wasps were
derived from different field-collected egg
masses to ensure siblings were not includ-
ed as replicates.

Inseminative capacity. — To determine
male inseminative capacity, all of the T.
basalis females that emerged from each of
10 parasitised green vegetable bug egg
masses were tested for insemination (see
below). These original egg masses con-
tained 85 ± 5 eggs, the mean size for
green vegetable bug egg masses in south-
eastern Queensland. Each egg mass had
been parasitised by a single, once-mated
female T. basalis over two days in a 50 x
25 mm ventilated vial. Self-superparasit-
ism is unlikely to arise under such condi-
tions because females use a chemical
marker to mark parasitised eggs (Wilson
1961; Ganesalingam 1966; Field et al.
1998). After 9-10 days, when the first
males began to emerge, vials containing
the parasitised egg masses were moni-
tored frequently (every 5-10 minutes) dur-
ing the 10 hours of artificial laboratory
light each day. Before females began to
emerge, all males were removed except
for the dominant male occupying the egg
mass. The dominant male was lightly
marked on the thorax with fluorescent
dust to distinguish him from males that
emerged subsequently. These latter males
were removed immediately they ap-
peared.

At each morutoring period any females

that had emerged were removed and each
was placed alone in a ventilated vial and
provided with honey. Females were typi-
cally found at the top of the vial. All fe-
males, except those used to assess fecun-
dity (see below), were provided 5-10 fresh
green vegetable bug eggs to establish
whether they produced female offspring,
a certain indication they had been insem-
inated (Wilson 1961). Females were al-
lowed 24 hours to parasitise eggs before
being removed.

Lifetime fecundity was assessed for the
first emerging female and for every tenth
female that emerged from each egg mass.
Each was provided with a frozen ( — 70°C)
green vegetable bug egg mass each day
for the first 12 days. The frozen eggs were
< 1 month old and still viable for T. basalis
(Powell and Shepard 1982, Kelly 1987).
Earlier trials (n = 6) indicated that daily
fecundity decreased rapidly and females
were unlikely to produce offspring after
12 days. Large egg masses (85 ± 5 eggs
each) were supplied on each of the first
two days, half masses (40 ± 5 eggs) for
each of the next four days, and small
masses (20 ± 5 eggs) for each of the last
six days, so that females had an excess of
hosts at all times (see Results). Parasitised
egg masses were placed singly in ventilat-
ed vials and incubated until all offspring
had emerged. Eggs that were obviously
parasitised, but from which parasitoids
failed to emerge, were dissected and the
parasitoid removed for sexing. Counts of
the numbers of male and female offspring
produced per female per day were then
made.

Adult size. — Adults were measured to
assess whether their size was affected by
emergence sequence and whether fecun-
dity was influenced by size. Two mea-
surements were taken from all males and
females to emerge from each egg mass:
head width and right hind tibial length.
Head width was measured as the distance
between the outermost points of the eyes.

Volume 8, Number 2, 1999

241

Measurements were made under a dis-
secting microscope, accurate to 0.01 mm.

Sexual receptivity. — Two experiments
were conducted to investigate the effect of
adult age and mating status (virgin or
mated) on sexual receptivity. The first ex-
amined whether age affected the male's
readiness to mate. A single virgin male
aged \, 5, 10, 15 or 20 days old was intro-
duced into one end of a 50 X 12 mm ven-
tilated vial containing a virgin female less
than 24 hours old at the other end. The
male and female were observed until mat-
ing occurred or for 20 minutes, as virgin
males and females would usually mate
within 10 minutes with an average pre-
mating time of ca 3 minutes.

The number of contacts between the
male and female before mating was re-
corded, as were the pre-mating and mat-
ing times. In addition, pre-mating and
mating behaviours were observed for any
differences between treatments. Once mat-
ing had taken place, the male was re-
moved and the female provided with ca
10 green vegetable bug eggs. The eggs
were removed one day later and incubat-
ed at 28 ± 1°C until offspring emerged.
Because T. basalis is arrhenotokous (Wil-
son 1961), a female was regarded as suc-
cessfully inseminated if any female off-
spring were produced.

The other experiment examined wheth-
er female age and mating status affected
her readiness to mate. Virgin females were
assigned to two groups. Those in one
group were not mated, whereas the others
were mated within 24 hours of emergence
by a virgin male. All females were held,
until needed, in a ventilated vial streaked
with honey. Subgroups of females (virgin
or once-mated) were exposed to virgin
males less than 24 hours old, at ages 1, 5,
10, 15 or 20 days, one pair per 50 X 12
mm ventilated vial. Procedures and con-
ditions were the same as for the first ex-
periment. For both experiments 15 repli-
cates of each treatment were conducted,
all at 25 ± 2°C and 65 ± 10% R.H.

Statistical analysis. — Logistic analyses
were conducted to test whether emer-
gence position influences the probability
of a female being inseminated. A logistic
regression was conducted for each of the
10 experimental replicates, in which the
binary response variable, whether a fe-
male was inseminated (assigned 1) or un-
inseminated (assigned 0), was regressed
against her position in the emergence se-
quence.

The fecundity of females in different
emergence positions and their offspring's
sex ratio were analysed by 1-way ANOVA
after log(x + 0.5) and arcsine(Vp) trans-
formations, respectively. The significance
of any differences was assessed by Fish-
er's protected least significant difference
test. Linear regression was employed to
assess the relationship between fecundity
and female head width or hind tibial
length.

The effect of age of males, virgin fe-
males and mated females on the number
of pre-mating contacts, pre-mating time
and mating time was tested by 1-way AN-
OVA after data were log(x + 0.5) trans-
Table 1. Summary statistics from logistic analyses
testing whether emergence position influences the
probability of a female being inseminated. A logistic
regression was conducted for each of the 10 experi-
mental replicates, in which the binary response var-
iable, whether a female was inseminated (assigned 1)
or uninseminated (assigned 0), was regressed against
her position in the emergence sequence. Relationship
refers to whether females later in the emergence se-
quence tended to be uninseminated (negative) or in-
seminated (positive).

\..

KL-plll.UC-

k'm.iles

\

p.^,,lu.

Ki-I.itionship

1

78

0.58

0.45

negative

2

40

1.52

0.22

positive

3

30

0.56

0.45

negative

4

73

0.30

0.58

positive

5

59

5.74

0.02

negative

6

55

0.80

0.37

positive

7

60

11.41

<0.01

negative

8

57

0.15

0.70

negative

9

61

2.78

0.10

positive

10

78

2.56

0.11

positive

242

Journal of Hymenoptera Research

Day

Fig. 1. Pattern in which Tnsso/cHS basnlis siblings
emerged from parasitised green vegetable bug egg
masses. Number of males and females that emerged
each day from each of 10 egg masses (parasitised on
day 0) in the laboratory at 28 ± rC, 65 ± 107o R.H.
and 16L:8D. Error bars represent standard errors for
the mean number of siblings (males and females) that
emerged each day.

formed. G-tests were employed to test the
effect of age of males, virgin females and
mated females on the number of females
mated within 20 minutes. G-tests were
also employed to test if the number of fe-
males inseminated was related to age of
males and virgin females. The effect of
mated female age on the probability of be-
ing inseminated was not analysed statis-
tically because mated females were pre-
sumed to have been successfully insemi-
nated at their first mating.

The effects of female age and mating
status (virgin or once-mated) on the num-
ber of pre-mating contacts, pre-mating
time and mating time were tested by two-
way ANOVA after data were log(x + 0.5)
transformed. A log-linear analysis was
employed to test the effects of female age
and mating status on the number of fe-
males mated within 20 minutes.

RESULTS

Emergence patterns. — Most wasps (>
95%) emerged under lighted conditions,
with the majority emerging during the
first 3-4 hours of morning light. A mean
± s.e. of 67.9 ± 4.6 wasps emerged from

Table 2. Overall number of offspring and off-
spring sex ratio (mean ± s.e.) produced by Trissokus
basnlis females in different positions in the entire
emergence sequence. Fecundity and sex ratio values
derive only from those females that had been insem-
inated (as indicated by their production of daugh-
ters). See Fig. 2 and Table 1 for details.

Emergence No. No nol
position nitiled mated

(t< m.i!e)

1

8

2

139 ± 13

0.23 ± 0.03

10

8

1

125 ± 17

0.26 ± 0.04

20

9

1

129 ± 16

0.29 ± 0.05

30

9

1

116 ± 11

0.36 ± 0.06

40

8

0

100 ± 12

0.30 ± 0.08

50

7

1

153 ± 23

0.30 ± 0.04

60

5

1

95 ± 15

0.22 ± 0.08

70

1

2

116

0.72

' Column means for fecundity (F-^„ = 1. 12, p = 0.37)
and sex ratio (F-„ = 1.61, p = 0.16) were not signif-
icantly different.

each of the 10 original egg masses, com-
prising 59.1 ± 5.0 females and 8.8 ± 2.3
males. Males began emerging on day nine
with emergence peaking on days 10 and
11 (Fig. 1). Few males, if any, emerged
from egg masses later than day 14. Fe-
males emerged on days 10-19 with emer-
gence peaking on days 11-13. The largest
number of females that emerged from any
one egg mass in one day was 57 females
on day 12.

Inseminative capacity and fecundity. — The
dominant males that were left alone on
egg masses to mate their sisters insemi-
nated a mean ± s.e. of 48.3 ± 3.9 females,
with 68 females being the maximum num-
ber inseminated by one male. The propor-
tion (mean ± s.e.) of emerging females
that was inseminated by the dominant
males was 0.82 ± 0.02 across egg masses.
Of the 10 replicates conducted, five
showed a positive relationship between
the probability of a female being insemi-
nated and her emergence position and five
showed a negative relationship (Table 1).
Two of the negative relationships were
significant at < 5% and two of the positive
relationships were significant at < 11%
(Table 1).

Volume 8, Number 2, 1999

243

Table 3. Number and sex ratio (mean ± s.e.) of
offspring produced by females that emerged as
adults during a single day, but in different positions
in the emergence sequence. Only females that
emerged on the first day of female offspring emer-
gence were included in the analysis because at this
stage males would presumably have had a full sperm
supply, and short term rates of sperm depletion could
be assessed most accurately. Fecundity and sex ratio
values are calculated only from mated females in
each position.

Emerge me
position

No.
mated

No. nol
mated

f-ecunditv

Sev r.ilio
(., male)

1

8

2

139 ± 13

0.23 ± 0.03

10

6

0

112 ± 19

0.27 ± 0.04

20

3

1

150 ± 4

0.27 ± 0.05

30

2

0

108 ± 4

0.46 ± 0.17

40

2

0

99 ± 45

0.21 ± 0.00

50

1

0

257

0.35

Column means for fecundity (F,,^ = 2.31, p = 0.09)
and sex ratio (F^,„ = 1.47, p = 0.25) were not signif-
icantly different.

Fecundity was highly variable, and
ranged from 42 to 257 (mean ± s.e. = 121
± 5, n = 65) offspring per female. Progeny
production peaked during the first 24
hours after emergence with about 40-50
offspring on average, and then decreased
rapidly with time (Fig. 2). The number of
offspring produced per day was always
less than the number of hosts provided.
The sex ratio (proportion male) of off-
spring increased with time such that fe-
males produced few or no female off-

spring after 10 days, although by then few
offspring were being produced (Fig. 2).
Fecundity and brood sex ratio were not
significantly affected by the position of
parent females in the overall emergence
sequence (Table 2), nor by the position of
females in the emergence sequence on the
first day of female emergence (Table 3).

Adult size. — Head widths and hind tibial
lengths for male and female T. basalts
showed little variation within and across
replicates. Females were significantly larg-
er than males: mean ± s.e. head widths
were 0.61 ± 0.001 mm and 0.58 ± 0.002
mm for females and males respectively
(F,,„ = 410.3, p < 0.0001), and their re-
spective hind tibial lengths were 0.41 ±
0.001 mm and 0.39 ± 0.001 mm (F,,,,, =
52.3, p < 0.0001). No trend between emer-
gence position and head width or hind
tibial length was apparent except that the
last 1-5 wasps to emerge from an egg
mass tended to have head widths and
hind tibial lengths up to 0.05 mm smaller
than previously emerged wasps.

Fecundity increased significantly with
increases in female head width and hind
tibial length (Fig. 3). However, regressions
of fecundity against each of the two size
measurements fitted poorly (r- £ 0.10).

Sexual receptivity — The age of males had
no significant effect on the number of pre-
mating contacts, mating time or the num-

Table 4. Effect of virgin male age on their propensity to mate within 20 minutes of exposure to a virgin
female (expressed as number of females mated). Also given is the number of females inseminated, number
of pre-mating contacts, pre-mating time and mating time (last three values are mean ± s.e.). The number of
males used to calculate each mean and s.e. is the number of males mated in 20 minutes (first row) from the
15 replicates.

Male aj;e (days)

1

5

111

1^

20

No. mated

lS,i

ISa

1 5a

15a

15a

No. inseminated

12a

12a

15a

12a

12a

No. contacts

2.6 ± 0.3a

1.9 ± 0.3a

1.8 ± 0.3a

2.7 ± 0.4a

2.2 ± 0.2a

Pre-maling time

(s)

173 ± 37a

92 ± 17b

88 ± 21b

181 ± 33a

176 i 24a

Mating time (s)

12.0 ± 1.3a

10.8 ± 1.0a

12.6 ± 0.6a

12.4 ± 0.8a

14.5 ± 2.1a

Row means tolknved h\ the same letter are not significantly ditterent (t;-test tor first 2 rows, l-way ANOVA
for last 3 rows, P > 0.05).

244

Journal of Hymenoptera Research

Table 5. Effect of virgin female age on their propensity to mate within 20 minutes of exposure to a virgin
male (expressed as number of females mated). Also given is the number of mated females that was successfully
inseminated, number of pre-mating contacts, pre-mating time and mating time (last three values are mean ±
s.e.). The number of females used to calculate each mean and s.e. is the number of females that mated within
20 minutes (first row) from the 15 replicates.

Virgin female age (

;days)

1

5

10

15

20

No. mated

15a

15a

14ab

lib

10b

No. inseminated

12a

11a

10a

8ab

3b

No. contacts

2.6 ± 0.3a

5.3 ± 0.8b

8.6 ± 1.9b

5.0 ± 1.1b

5.8 ± 0.8b

Pre-mating time

(s)

173 ± 37a

303 ± 43b

419 ± 113b

275 ± 93ab

370 ± 73b

Mating time (s)

12.0 ± 1.3a

11.5 ± 0.7a

8.1 ± 0.8b

10.9 ± 0.7a

9.3 ± 1.5ab

Row means followed by the same letter are not significantlv different (G-test for first 2 rows, 1-way ANOVA
for last 3 rows, P > 0.05).

ber of females mated or inseminated (Ta-
ble 4). Pre-mating time was affected by
male age with males aged 5 and 10 days
old having a significantly shorter pre-mat-
ing time than males aged 1, 15 and 20
days old. No differences in male pre-mat-
ing or mating behaviour were observed
between males of different ages.

In contrast, virgin female pre-mating
and mating behaviours were affected by
their age (Table 5). Females aged 5-20
days old tended to resist the males' mat-
ing attempts by moving away from them,
aggressively chasing males away and /or
refusing to allow males to copulate after
mounting. The numbers of pre-mating
contacts and pre-mating times were great-
er for females aged 5-20 days than for 1
day old females (Table 5). Females aged 1,
5 and 15 days old mated for significantly

longer than 10 day old females. The num-
ber of females mated within 20 minutes
and the number successfully inseminated
decreased significantly with female age
(Table 5).

The age of mated females also affected
their pre-mating and mating behaviours,
with mated females aged 5-20 days gen-
erally resisting mating attempts in the
way described above for virgin females of
different age. The mating propensity of
mated 5-20 day old females was signifi-
cantly less than that of one day old mated
females (Table 6). The numbers of pre-
mating contacts and pre-mating times in-
creased significantly with female age, but
mating time was not significantly affected
by their age (Table 6).

Two-way ANOVA examining the ef-
fects of female age and sexual status (vir-

Table 6. Effect of age of previously-mated females on their propensity to mate within 20 minutes of ex-
posure to a virgin male (expressed as number of females mated). Also given is the number of pre-mating
contacts, pre-mating time and mating time (values are mean ± s.e.). The number of females used to calculate
each mean and s.e. is the number of females that mated within 20 minutes (first row) from the 15 replicates.

Age

of pre\'iouslv-mafed female (

da\s)

1

5

10

IS

:ii

No. mated

15a

6b

8b

9b

6b

No. contacts

4.7 ± 1.0a

9.7 ± 2.9b

7.0 ± l.Oab

8.1 i 1.1b

9.3 ± 3.4b

Pre-mating time (s)

235 ± 48a

549 ± 143b

373 ± 74ab

514 ± 97b

547 ± 205ab

Mating time (s)

8.6 ± 0.7a

8.4 ± 1.1a

6.1 ± 1.1a

7.4 ± 1.5a

9.7 ± 2.2a

Row means followed by the same letter are not significantly different (G-test for first row, 1-wav ANOVA for
last 3 rows, P > 0.05). '

Volume 8, Number 2, 1999

245

Table 7, Summary of results from 2-way ANOVA (log (x + 0.5) transformed) testing whether the individual
and interactional effects of the factors, female age and mating status (virgin or once-mated), affected the
number of pre-mating contacts, pre-mating time and mating time.

.ilin^ tinu-

Milling linu'

Age

Mating status

Age X mating status

F^„ = 7.06, P < 0.0001
F,„ = 8.37, P = 0.005
F,« = 0.89, P = 0.47

F^^ = 4.50, P = 0.002
F,^ = 8.65, P = 0.004
F,« = 0.83, P = 0.51

3.31, p = 0.01

F,„ = 10.01, P
F^'^ = 0.96, P =

= 0.002
0.44

gin or once-mated) on mating propensity
indicated that the interaction between fe-
male age and mating status was not sig-
nificant for all three measures of mating
propensity (Table 7). As single factors, fe-
male age and mating status significantly
affected the number of pre-mating con-
tacts, pre-mating time and mating time
(Table 7).

A log-linear analysis on the effect of fe-
male age on the propensity of virgin and
mated females to mate within 20 minutes
revealed that the model could be de-
scribed best by two interactions: female
mating status and the number of females
mated, and female age and the number of
females mated (Maximum likelihood x" =
8.38, df = 8, p = 0.40). The propensity of
females to mate within 20 minutes was
significantly greater for virgin females and
younger females.

DISCUSSION

The results from this study suggest that
strict local mating does not occur in T. ba-
salts and that outbreeding away from the
natal site may commonly occur. Results
that are inconsistent with LMC theory in-
clude: (1) males do not achieve the maxi-
mum rate of insemination expected de-
spite showing no apparent signs of sperm
depletion, (2) males remain sexually re-
ceptive probably throughout their lifetime
(Table 4), and (3) females can be mated
multiple times, despite becoming decreas-
ingly sexually receptive after mating and
with age (Tables 6, 7). We discuss the im-
plications that these results have on the
mating system of T. hasalis and ask wheth-
er this species is likely to outbreed.

Emergence of T. basalis females is con-
centrated during the early morning hours
over several days (Wilson 1961; Fig. 1). In
the field a dominant male usually guards
the parasitised egg mass from which fe-
males are emerging. These females are his
sisters unless the egg mass has been su-
perparasitised. In the laboratory, the sin-
gle male left on the mass successfully mat-
ed many females in succession, with
sometimes up to 50 or more females
emerging over several hours (Table 3). Fe-
males in all positions in the emergence se-
quence apparently received similar quan-
tities of sperm because their offspring sex
ratios were not affected by emergence po-
sition (Table 2; Fig. 2), even if those fe-
males all emerged and were mated on the
same day by a single male (Table 3).

Although the above results suggest that
a single T. basalis male can fully insemi-
nate each of his female siblings from the
same egg mass, two observations indicate
that the dominant male does not achieve
the maximum rate of insemination that is
possible. First, only ca 80% of females
were inseminated by dominant males (Ta-
ble 1, see also Wilson 1961). Wilson (1961)
proposed that temporary sperm depletion
in the dominant male may be the cause.
But even when large numbers of females
emerged in a day, the offspring sex ratio
produced by inseminated females did not
vary with their position in the mating se-
quence (Table 3). In addition, uninsemi-
nated females appeared throughout the
emergence sequence (Tables 1-3). Field
observations have shown that a similar
percentage of emerging females is not

246

Journal of Hymenoptera Research

mated by the dominant male guarding the
egg mass (A.D. Loch and G.H. Walter un-
published data). Also, ca 18% of mahngs
between virgin males and females in the
laboratory do not lead to successful in-
semination (A.D. Loch and G.H. Walter
unpublished data). In our experiments, we
did not observe females to confirm they
mated or to ascertain why they may not
have mated. However, the high rate of un-
inseminated females is likely to be partly
the consequence of simultaneous female
emergences, during which males become
occupied with some emerging females,
while others move unmated to the top of
the vial, a behaviour that has parallels in
the field (A.D. Loch and G.H. Walter un-
published data).

The second observation suggesting that
maximum insemination rates are not
achieved by the dominant male, is that
even those females that were inseminated
produced proportionately more male off-
spring with age (Fig. 2). This trend has
also been reported in other studies of T.
basalis fecundity (Powell and Shepard
1982; Correa-Ferreira and Zamataro 1989;
Awan et al. 1990) and in work on the con-
familial Telenomus busseolae (Gahan) (Cha-
bi Olaye et al. 1997). Females apparently
do not receive sufficient sperm to fertilise
all their eggs. However, temporary sperm
depletion in males or insufficient sperm
transfer by males are unlikely explana-
tions. First, this trend was uniform for fe-
males in all emergence positions (Fig. 2),
indicating that sperm depletion in the
male was not the cause. Second, females
held with males throughout their lifetime,
and therefore assumed to be mated mul-
tiple times, also produce proportionately
more male offspring with time (Powell

and Shepard 1982; Awan et al. 1990), sug-
gesting that this trend occurs irrespective
of the number of times a female is mated.
The reason for this trend is not clear, but
a decrease in sperm viability over time is
possible, or it could have a behavioural or
physiological basis.

Results from this study suggest that T.
basalis males do not become sperm de-
pleted at rates of mating that are typical
for this species in nature. In this study,
green vegetable bug egg masses of 85 ± 5
eggs were used, and represent the largest
known host masses for T. basalis, in terms
of the number of eggs. The test males
were, therefore, exposed to a high number
and frequency of matings. In any case,
dominant males in control of egg masses
in the field are unlikely to become sperm
depleted because changeovers in male
dominance occur frequently (A.D. Loch
and G.H. Walter unpublished data), and
female emergence continues over several
days (Fig. 1).

Sexual receptivity of T. basalis males ap-
pears unaffected by age (Table 4) and mat-
ing status, thus enabling males to mate
probably throughout their lifetime. The
only aspect of male sexual receptivity that
was affected by age was pre-mating time,
which was significantly shorter for 5-10
day old males than for 1, 15 and 20 day
old males. They may be more receptive at
5-10 days because they emerge up to sev-
eral days before females (Anon. 1939; No-
ble 1937; Smith 1945; Wilson 1961; Thom-
as 1972) and would therefore not normally
need to mate immediately upon emer-
gence.

In contrast, female sexual receptivity
decreased rapidly after mating and with
age (Tables 5-7). Such decreases are con-

Fig. 2. Number (mean ± s.e.) of progeny produced each day after emergence by inseminated Tnssc/CKS
basalis females. Data are presented separately for each group of parent females according to their position in
the emergence sequence (i.e. (a) 1st, (b) 10th, (c) 20th, (d) .3()th, (e) 40th, (f) 50th, (g) 60th and (h) 70th), and
thus the sequence in which they were inseminated by the dominant male on their host egg mass. Numbers
above error bars indicate the number of females still alive at that time.

Volume 8, Number 2, 1999

(a) Female 1
70

247

■ Females
Q Males

12 3 4 5 6

r^r^-

9 10 11 12

(e) Female 40

70 -
60 -

(f) Female 50

9 10 11 12

(g) Female 60

E

z

r^r— ir^

9 10 11 12

1 2 3 4 5 6 7 6 aiU1112

70 •
60 ■
50

I.

HmnH^

1 1 1

Female age (days)

123456789 10 1112

Female age (days)

248

Journal of H-imenoptera Research

sistent with LMC theory because mating
is assumed to occur only at the natal site
among siblings (Hamilton 1967). Howev-
er, females can be mated multiple times,
which is inconsistent with LMC theory.
This inconsistency coupled with others
such as male dispersal from the natal site
(A. D. Loch and G. H. Walter unpublished
data), and males remaining sexually re-
ceptive probably throughout their lifetime
(Table 4) suggest that T. basalis of both
sexes may mate away from the natal site
and therefore outbreed.

Trissolciis basalis may outbreed if males
and females can locate and / or attract each
other once they have left the natal site.
Males may be able to locate unrelated,
newly-emerged (and thus sexually recep-
tive) females directly, or they could do so
indirectly by locating hosts parasitised by
T. basalis and then competing with emerg-
ing males for mating access to females.
Similarly, newly-emerged females may be
able to locate males directly, or indirectly
by searching for parasitised hosts with
males in occupation. Currently, no evi-
dence is available on whether males or fe-
males can locate potential mating partners
away from the natal site. LMC models as-
suming strict local mating (Hamilton
1967) suggest that T. basalis males and fe-
males will not be able to locate each other
in the field. However, in other hymenop-
terous species with female-biased sex ra-
tios, such as Spalangia cameroui (Perkins)
(Myint and Walter 1990) and Pacln/crepo-
ideus vindemiae (Rondani) (Nadel and Luck
1992), males are able to locate hosts and
thus potential mating partners, a feature
likely to be found in other species (Hardy
1994).

Males may not only be able to outbreed
with newly-emerged virgin females but
also with newly-emerged mated females
because a previous mating encounter did
not preclude females from mating again
(Table 6). This result is not likely to be an
artefact of laboratory conditions or pro-
cedures because females have been ob-

Head width (mm)

Fig. 3. Fecundity of Trissolcus basalis females in re-
lation to head width (y = 1711x - 928, r^ = 0.10, n
= 65, p = 0.01). The trend for fecundity versus hind
tibial length (y = 1294x - 407, r- = 0.07, n = 65, p
= 0.04) is not shown because it was similar to the
displayed trend. Both trends were determined irre-
spective of female position in the emergence se-
quence (see Table 2).

served to be mated multiple times and by
multiple males in the field (A. D. Loch and
G. H. Walter unpublished data). Whether
T. basalis females are truly polyandrous
has yet to be established, for matings after
the first successful mating may not lead to
successful insemination. For instance,
mating plugs may be used by males to en-
sure additional matings do not result in
insemination.

This study also made a number of find-
ings pertaining to the fecundity of T. ba-
salis. The mean fecundity recorded in this
study is higher than fecundities recorded
by Noble (1937), Ganesalingam (1966) and
Thomas (1972), but similar to values re-
corded by Powell and Shepard (1982) and
Correa-Ferreira and Moscardi (1994), and
lower than fecundities recorded by Cor-
rea-Ferreira and Zamataro (1989) and
Awan et al. (1990) for the same species.
These differences are likely to be the result
of differences in laboratory procedures
and conditions, although differences in
adult female size may have contributed
because fecundity is greater for larger fe-
males (Fig. 3). The trend whereby fecun-
dity peaked on the first day after female
emergence and decreased rapidly over

Volume 8, Number 2, 1999

249

time, differs somewhat from the results of
Ganesalingam (1966) and Powell and
Shepard (1982), who showed that fecun-
dity peaked on day 2. These differences
are less readily attributable to different
laboratory procedures and conditions, and
their significance is unclear. The claim by
Field et al. (1998) that T. basalis is a syno-
vigenic species was supported by our re-
sults because females laid eggs for 10-12
days with progressively fewer eggs each
day (Fig. 2) despite sufficient hosts being
available during the first few days for
them to have deposited their lifetime com-
plement of eggs then.

In conclusion, although uninseminated
females leaving the egg mass may well be
mated by the other males (also likely to be
their siblings) that wait around the egg
mass, the possibility that these females
could mate unrelated males near or away
from the natal site may not be low. Al-
though we have no direct evidence of T.
basalis outbreeding in nature, the results
from this study and other related studies
(A.D. Loch and G.H. Walter unpublished
data), suggest that it may be more fre-
quent than anticipated by LMC theory. In
addition, outbreeding is likely to occur
when > 1 female oviposits in an egg mass.
Further research investigating the mating
system of T. basalis is required before the
question of the species' outbreeding can
be resolved. Specific issues that need to be
addressed include the questions of wheth-
er T. basalis haa a means of mate-attrac-
tion, and whether T. basalis females are
truly polyandrous.

ACKNOWLEDGMENTS

We thank Tony Clarke for discussing aspects of
this manuscript, and Norman Johnson (Ohio State
University) for confirming the identification of Tris-
solcus basalis. We also thank Ian Hardy and Peter
Mayhew for their comments and suggestions on ear-
lier versions of the manuscript. The senior author was
supported bv a Grains Research and Development
Corporation lunior Research Fellowship.

LITERATURE CITED

Anon. 1939. The egg parasite of the green vegetable
bug. The Agricultural Gazette of Neu' Simf/i Wales
50; 277-278.
Awan, M. S., Wilson, L. T. and Hoffman, M. P. 1990.
Comparative biology of three geographic popu-
lations of Trissolcus basalis (Hymenoptera; Sce-
lionidae). Environmental Entomology 19; 387-392.
Chabi Olaye, A., Schulthess, F., Shanower, T. G. and
Bosque-Perez, N. A. 1997. Factors influencing the
developmental rates and reproductive potentials
of Teteiumius busseolae (Gahan) (Hym.: Scelioni-
dae), an egg parasitoid of Sesamia calamistis
Hampson (Lep.: Noctuidae). Biological Control 8;
15-21.
Correa-Ferreira, B. S. and Moscardi, F. 1994. Temper-
ature effect on the biology and reproductive per-
formance of the egg parasitoid Trissolcus basalis
(WoU.). Anais da Sociedade Entomologica do Brasil
23: 399-108.
Correa-Ferreira, B. S. and Zamataro, C. E. O. 1989.
Reproductive capability and longevity of the egg
parasitoids Trissolcus basalis (WoUaston) and
Trissolcus mitsukurii Ashmead (Hymenoptera;
Scelionidae). Revista Brasileira de Biologui 49; 621-
626.

Field, S. A., Keller, M. A. and Austin, A. D. 1998.
Field ecology and behaviour of the egg parasit-
oid Trissolcus basalis (WoUaston) (Hymenoptera;
Scelionidae). Transactions of the Royal Society of
South Australia 122: 65-71.

Ganesalingam, V. K. 1966. Some environmental fac-
tors influencing parasitisation of the eggs of Ne-
zara viridula L. (Pentatomidae) by Telenornus ba-
salis WoUaston (Hymenoptera: Scelionidae). Cey-
lon Journal of Science (Biological Sciences) 6: 1-14.

Godfray, H. C. ]. 1994. Parasitoids: behavioural and evo-
lutionary ecolog}/. Princeton University Press,
Princeton, 473 pp.

Hamilton, W. D. 1967. Extraordinary sex ratios. Sci-
ence 156; 477-188.

Hardy, I. C. W. 1994. Sex ratio and mating structure
in the parasitoid Hymenoptera. Oikos 69: 3-20.

Hardy, I. C. W., Dijkstra, L. J., Gillis, J. E. M. and Luft,
P. A. 1998. Patterns of sex ratio, virginity and
developmental mortality in gregarious parasit-
oids. Biological journal of the Linnean Society 64:
239-270.

Hardy, 1. C. W., Ode, P. J. and Strand, M. R. 1993.
Factors influencing brood sex ratios in polyem-
bryonic Hymenoptera. Oecologia 93: 343-348.

Hardy, I. C. W., Pedersen, J. B., Sejr, M. K. and Lin-
deroth, U. H. 1999. Local mating, dispersal and
sex ratio in a gregarious parasitoid wasp. Ethol-
ogy 105: 57-72.

Kelly, G. L. 1987. Factors affecting the success of Tns-
solcus basalis (Hvmenoptera: Scelionidae) as a bi-
ological control agent of the green vegetable bug.

250

Journal of Hymenoptera Research

Nezara viriciula (Hemiptera: Pentatomidae). Un-
published PhD thesis. University of Sydney, Syd-
ney, 329 pp.

Molbo, D. and Parker Jr, E. D. 19%. Mating structure
and sex ratio variation in a natural population of
Nnsi'ii/n I'itripennis. Proceedings of the Roi/nl Societt/
cf Lciidoii. Series B. Biological Sciences 263: 1703-
1709.

Myint, W. W. and Walter, G. H. 1990. Behaviour of
Si'alangia cameroni males and sex ratio theory. Oi-
kos 59;' 163-174.

Nadel, H. and Luck, R. F. 1985. Span of female emer-
gence and male sperm depletion in the female-
biased, quasi-gregarious parasitoid, Piicln/crepo-
ideus vindcmiae (Hymenoptera: Pteromalidae).
Annals of the Entomological Society of America 78:
410-414.

Nadel, H. and Luck, R. F. 1992. Dispersal and mating
structure of a parasitoid with a female-biased sex
ratio: implications for theory. Evolutionary Ecolo-
gy 6: 270-278.

Noble, N. S. 1937. An egg parasite of the green veg-
etable bug. Agricultural Gazette of Neiu South
Wn/cs48: 337-341.

Ode, P. J., Antolin, M. F. and Strand, M. R. 1997. Con-
strained oviposition and female-biased sex allo-
cation in a parasitic wasp. Oecologia 109: 547-555.

Powell, J. E. and Shepard, M. 1982. Biology of Aus-
tralian and United States strains of Trissolcus ba-

salis, a parasitoid of the green vegetable bug, Ne-
zara viridula. Australian Journal of Ecologi/ 7: 181-
186.

Smith, J. H. 1945. Useful parasitic insects. Queensland
Agricultural journal 60: 340-351.

Thomas, J. W. Jr. 1972. Evaluation of Trissolcus basalts
(Wollaston) as an egg parasite of Nezara viridula
(Linnaeus). Unpublished MSc thesis, Louisiana
State University, Louisiana, 99 pp.

van den Assem, J., Gijswijt, M. J. and Niibel, B. K.
1980. Observations on courtship - and mating
strategies in a few species of parasitic wasps
(Chalcidoidea). Netherlands Journal of Zoology 30:
208-227.

Waage, J. K. and Lane, J. A. 1984. The reproductive
strategy of a parasitic wasp. II. Sex allocation and
local mate competition in Trichogramma evanes-
cens. journal of Animal Ecolog]/ 53: 417^26.

Waage, J. K. and Ng, S. M. 1984. The reproductive
strategy of a parasitic wasp. I. Optimal progeny
and sex allocation in Trichogramma evanescens.
journal of Animal Ecology 53: 401^15.

Walter, G. H. and Clarke, A. R. 1992. Unisexual
broods and sex ratios in a polyembr\'onic encyr-
tid parasitoid (Copndosonia sp.: Hymenoptera).
Oecologia 89: 147-149.

Wilson, F. 1961. Adult reproductive behaviour in
Asolcus basalts (Hymenoptera: Scelionidae). Aus-
tralian journal of Zoology 9: 739-751.

J. HYM. RES.
Vol. 8(2), 1999, pp. 251-267

The Nearctic Species of Protarchiis Foerster
(Hymenoptera: Ichneumonidae: Ctenopelmatinae)

Luc Leblanc

Eastern Cereal and Oilseed Research Centre, Agriculture and Agri-Food Canada, Ottawa,

Ontario, KIA 0C6, Canada

Abstract. — The Nearctic species of the Holarctic genus Protnrchus Foerster (Ichneumonidae, Cten-
opelmatinae, Mesoleiini) are reviewed. A key to the Nearctic species is provided. Seven species
are recognized. P. testntorius (Thunberg) and P. sarin (Ratzeburg) are Holarctic, P. bolbogaster
Leblanc, n. sp. and P. mellipes (Provancher) are transcontinental Nearctic, P. magnus (Davis) and
P. pallidiconiis (Walley) are eastern Nearctic, and P. atrofacies Leblanc, n. sp. is found in Alaska.
The name P. longipes (Cushman) is synonymized with P. sorbi, new synonymy.

The Holarctic genus Protarchus Foerster
belongs to the ichneumonid subfamily
Ctenopelmatinae (Scolobatinae sensu
Townes 1970) and tribe Mesoleiini. The
tribe is the most speciose and taxonomi-
cally difficult in the subfamily. Only a few
genera have been studied in detail (Kaur
1989, Leblanc 1989, Viitasaari 1979). The
Palearctic species of Protarchus were re-
vised by Viitasaari (1979), who recognized
four species and discussed the biology of
the species that parasitize sawflies of the
genus Trichiosoina (Cimbicidae) in bogs,
based on extensive collecting and rearing.
The Nearctic Protarchus were studied by
Walley (1938), who recognized four spe-
cies based on specimens in the Canadian
National Collection.

Ichneumonids of the genus Protarchus
are large sized, the nervellus in the hind
wing is intercepted above its middle, the
areolet is present, the clypeus is small and
the medial dorsal carinae on the first me-
tasomal tergite extend beyond the spira-
cles. Both Nearctic and Palearctic species
parasitize Trichiosoma spp.

Study of the Nearctic and Palearctic
specimens shows that the Nearctic P. lon-
gipes (Cushman) and the Palearctic P. sorbi
(Ratzeburg) represent a single Holarctic

species. In addition, P. testatorius (Thun-
berg) is Holarctic but had not been re-
ported for the Nearctic region by previous
authors. The discovery of Holarctic distri-
bution patterns and of two undescribed
Nearctic species encouraged me to revise
the Nearctic species of Protarchus.

MATERIALS, METHODS AND
TERMINOLOGY

Material studied. — A total of 87 Nearctic
specimens were examined from 5 collec-
tions as follows (collections acronyms are
from Amett and Samuelson (1986)): AEIC:
American Entomological Institute, Gaines-
ville, Florida, D.B. Wahl, 31 specimens;
ANSP: Academy of Natural Sciences, Phil-
adelphia, Pennsylvania, D. Azuma, 2 spec-
imens; CNCI: Canadian National Collec-
tion of Insects, Ottawa, Ontario, J.R. Bar-
ron, 43 specimens; LLIC: Luc Leblanc pri-
vate collection, Montreal, Canada, 1
specimen; USNM: National Museum of
Natural History, Washington, D.C., B.
Danforth, 10 specimens.

Morphological terms. — The morphologi-
cal terms used in the descriptions are from
Gauld (1984) except that mesosoma is
used instead of thorax, and metasoma is
used instead of gaster. Terms used to

252

Journal of Hymenoptera Research

characterize microsculpture are from Al-
len and Ball (1980). In characterizing me-
tasomal punctures and sculpture, only the
second tergum was utilized as sculpture
is best defined on that segment.

Color descriptions. — Areas used to de-
scribe color patterns are as follows. Anten-
nae are divided into dorsal and ventral
surfaces and legs are divided into anterior,
posterior, dorsal and ventral surfaces if
imagined as stretched out horizontally at
right angles from the body. Areas in bilat-
erally symmetrical parts (face, clypeus,
epicnemium, dorsal and ventral surfaces
of mesosoma and metasomal terga) are
identified as median /lateral and basal/
apical for face, clypeus and epicnemium,
or median /apical and anterior /posterior
for other parts. Areas in asymmetrical
parts are identified as anterior /posterior
and upper /lower.

Measurements. — A total of 37 females
and 20 males were measured with an oc-
ular micrometer. Length of the forewing
was measured from the junction of costal
vein with humeral plate to the most dis-
tant tip between the ends of veins Rs and
M. Height of face is the distance between
a line connecting the lower margins of an-
tennal sockets and a line connecting the
tentorial pits. Width of face is the distance
between the inner margins of the com-
pound eyes at the level of the middle of
face height. Length of the first metasomal
tergum was measured in lateral view from
the base to the tip of its dorsal surface.
Width of the first metasomal tergum was
measured in dorsal view at its widest part
near the apex. The length of the hind fe-
mur was measured on the anterior surface
from the base to the apex. The width of
the hind femur was measured at its mid
length. The widths of the hind tibia and
hind tarsomere 1 were measured at their
widest part in lateral view.

Genus PROTARCHUS Foerster

Protnrchus Foerster 1869:201. Type species: Try-
pluvi rufus Gravenhorst (= testatorius Thun-

berg). Designated by Woldstedt 1877:460.
Lectotype not examined.

Zacalles Foerster 1869:204. Type species: Zacalles
mngnus Davis. Designated by Davis 1898:283.
Synonymized by Cushman 1924:8. Holotype
examined.

Protarchoidcs Cushman 1922:25. Type species:
Protarchoidcs loiigipes Cushman. Original des-
ignation. Synonymized by Townes 1945:505.
Holotype examined.

Diagnostic characters. — Large (forewing
9.8 to 18.3 mm long). Clypeus (Fig. 3)
small, transversly convex near base (near
middle in P. sorbi), apically almost flat,
and with apical margin truncate. Mandi-
ble short and broad, its upper tooth a little
wider and longer than lower tooth (Fig. 4)
or much wider than lower tooth (Fig. 5).
Forewing with areolet usually present,
large (Fig. 28) or very small (Fig. 29). Ves-
tige of vein 1/Rs + M (= ramulus) often
present (Fig. 29) (always absent in other
genera of Mesoleiini). Vein cu-a separated
from vein Rs-I-M by 0.2 to 0.5 of its length.
Hind wing with vein 1/Cu longer than
vein cu-a ("nervellus intercepted above
middle"). Tibial spurs of middle and hind
legs unequal, the longest spur about 0.3 to
0.4 as long as first tarsomere. First tergum
moderately stout, with median dorsal ca-
rinae well defined and strong (Figs. 17,19),
or reduced to a median furrow (Fig. 21)
but always extending beyond spiracle.
Terga 2 to 4 in some species each with two
large sublateral swellings where punc-
tures are sparser (Fig. 23). Hairs on female
hypopygium directed backward.

Biology and biogeography. — Hosts of Pro-
tarchus are almost invariably larvae of Tri-
chiosoma (Hymenoptera: Cimbicidae).
Published host records for the Palearctic
region are: T. nanae Vikberg & Viitasaari,
parasitized by P. testatorius in Finland (Vi-
itasaari 1979, Vikberg & Viitasaari 1991);
T. ?lucoruni L., parasitized by P. testatorius
and P. sorbi in Finland (Viitasaari 1976,
1979). Records of Palaeociiubex femorata,
parasitized by P. testatorius (Townes et al.
1965) and Cindu'x parasitized by P. Iieros

Volume 8, Number 2, 1999

253

(Holmgren 1876) would require confir-
mation. Label data included with Nearctic
specimens indicate that Trichiosoma trian-
guliim is attacked by P. testatorius, P. sorbi
and P. mellipes. Bogs are the usual habitat
of Protarchus spp. (Viitasaari 1979). P. tes-
tatorius, P. sorbi and P. bolbogaster release
a strong odor when picked up, as indicat-

ed by label data. The species P. sorbi and
P. testatorius are present in the boreal zone
of North America (Figs. 31-32) as well as
across the Palearctic region. The new spe-
cies P. atrofascies, from Alaska, may also
be Holarctic. The remaining three species,
on the other hand, are apparently restrict-
ed to north-eastern North America.

KEY TO NEARCTIC SPECIES OF PROTARCHUS

1. Hind tibiae entirely light colored, brown or yellowish brown; ocelli enlarged and hind
ocelli separated by about their diameter (Fig. 1) 2

- Hind tibiae apically to entirely dark, reddish black to black; ocelli smaller and hind ocelli
separated by more than their diameter (Fig. 2) 4

2. Epomia clearly defined and prominent (Fig. 12); metasoma black or brownish black beyond
tergum 2; size forewing 12.3-16.0 mm long 3

- Epomia indistinct; metasoma generally mostly brown, but in some specimens black beyond
tergum 2; forewing 16.5-18.3 mm long magniis (Davis)

3. Face black, or sometimes black with a yellowish brown median spot; mesosoma entirely
black except light tegula; metasoma entirely black mellipes (Provancher)

- Face brown or yellowish brown; mesosoma brownish black with extensive yellowish or-
ange and brown markings; metasoma brownish black with tergum 1 and part of tergum

2 brown pallidicontis (Walley)

4. Metasomal terga black, with extensive orange on terga 2 to 4; areolet usually present and
very small (Fig. 29) testatorius (Thunberg)

- Metasomal terga entirely black; areolet large (Fig. 28) or absent 5

5. Metasomal terga 2 to 4 each with two large sublateral swellings (Fig. 22); upper mandibular
tooth much wider than lower tooth (Fig. 5); hind tibia entirely black

bolbogaster Leblanc, n. sp.

- Metasomal terga 2 to 4 without sublateral swellings (Fig. 23); upper mandibular tooth
subequal to lower tooth (Fig. 4); hind tibia basally yellow to orange and apically black ... 6

6. Notaulus weak and reduced to shallow impressions (Figs. 6,7); hind tibia of male black in
apical 0.6; face of male yellow sorbi (Ratzeburg)

- Notaulus anteriorly strong and sharply defined (as Figs. 8,9); hind tibia of male black m
apical 0.5; face of male black except faint brown median spot (female unknown)

atrofacies Leblanc, n. sp.

Protarchus atrofacies Leblanc,
new species

(Fig. 32)

Diagnostic combination. — Face 1.2 X as
wide as high (wider in other species), ar-
eolet absent, face of male black except
faint light spot (female unknown), meso-
soma and metasoma predominantly black.

Description. — Structure: Antenna with
>20 flagellomeres (broken). Ocelli en-
larged and sitting on a swelling, hind ocel-

li separated by about their diameter. An-
tennal sockets in lateral view forming a
moderately strong angle with vertical axis
of compound eye. Frons not strongly de-
pressed. Face 1.2 X as wide as high. Me-
dian swelling of face moderate. Upper
mandibular tooth subequal in size and
shape to lower tooth. Notaulus strong and
sharply defined only at anterior end of
mesoscutum. Epomia weak. Mesopleuron
separated by 0.5-2.0 of their diameter and

254

Journal of Hymenoptera Research

3

■^^

^ v^^^i

Figs. 1-12. 1-2, head in dorsal view: 1, P. mngniif, 2, I', tcftatoniis. 3, face ot i'. -n/i'i 4-S, mandibular teeth;
4, P. testiitoriiis; 5, P. bolbof:af,ter. 6-11, niesoscutum and notauli, dorsal and lateral: 6-7, P. scrbi; 8-9, /'. ningniif,
10-11, P. (I'sfnfpriKS. 12, pronotum of P. iiwUipcf, with epomia (EPM).

microsculpture with meshes well outlined
and sculpticells convex; well defined on
posterior 0.3 of mesopleuron below spec-
ulum and gradually fading towards other
parts of mesepisternum. Carinae of pro-
podeum strongly defined. Forewing 12.3
mm long, areolet absent. Hind femur 3.7
mm long and 6.2 X as long as wide. Hind
tarsomere 1 weakly compressed laterally

and 0.55 X as wide as hind tibia near
apex. First metasomal tergum 2.1 X as
long as wide, with median dorsal carinae
reduced to a median furrow; in lateral
view weakly and regularly curved at mid-
length. Sublateral swellings on terga ab-
sent. Metasomal tergum 2 with punctures
almost coalescent with outlines almost po-
lygonal when close, and microsculpture

Volume 8, Number 2, 1999

255

Figs. 13-24. 13-14, mesopleuron: 13, P. sor/'i; 14, P. l'olbo\;ii^U-i. 15-16, propiidt-um: 15, P. forbi; 16, P. Ic~^l(itnriiis.
17-21, metasomal tergum 1, dorsal and lateral: 17-18, P. holhogasfer. 19-20, P. forbi. 21, P. tefnfdmis. 22-24,
metasomal tergum 2, dorsal: 22, P. maginis; 23, P. bolbogaster; 24, P. sorbi.

well defined with sculpticells convex
along posterior and lateral borders of ter-
gum, at most suggested at centre of ter-
gum and almost entirely faded anteriorly.
Coloration: Antenna with scape, pedicel
and flagellum black except traces of yel-
lowish brown on flagellomeres 9 to 13.
Head black except faint brown median
spot on face below tubercle, brown clyp-

eus and brown anterior surfaces and api-
cal third of lateral surfaces of mandibles.
Mesosoma black. Wings with light yellow
infuscation. Legs with all coxae and tro-
chanters black. Femur, tibia and tarsus of
all legs orange except apical half of hind
tibia and whole hind tarsus black. Meta-
soma black.

Etymology. — From the Latin aier (black)

256

Journal of Hymenoptera Research

and fades (face), referring to the dark col-
ored face.

Distribution. — Known only from the
type locality (Fig. 32).

Specimen examined. — Holotype male:
"Unalakleet, Alaska 8.viii.l961 R. Madge".
Condition of type: missing left antenna be-
yond flagellomere 17 and right antenna
beyond flagellomere 20. [CNCI].

Protarchtts bolbogaster Leblanc,
new species

(Figs. 5,14,17,18,23,30)

Diagnostic combination. — Upper mandib-
ular tooth much wider than lower tooth
(Fig. 5) (subequal in other species), subla-
teral swellings on metasomal tergites 2 to
4 (Fig. 22) (absent in other species), me-
sosoma and metasoma predominantly
black.

Description. — Structure: Antenna with
48-53 flagellomeres. Ocelli of moderate
size and not sitting on a swelling, hind
ocelli separated by more than their diam-
eter. Antennal sockets in lateral view
forming a moderately strong angle with
vertical axis of compound eye. Frons not
strongly depressed. Face 1.7-1.8 (female)
and 1.5-1.7 (male) X as wide as high. Me-
dian swelling of face moderate. Upper
mandibular tooth much wider than lower
tooth (Fig. 5). Notaulus strong and sharply
defined to middle of mesoscutum. Epomia
indistinct. Mesopleuron (Fig. 14) with
punctures separated by 0.5-2.0 of their di-
ameter and microsculpture with meshes
lightly impressed and sculpticells slightly
convex, but more convex anterior to spec-
ulum. Carinae of propodeum strongly de-
fined. Forewing 13.6-15.2 (female) and
13.9-14.8 (male) mm long, areolet present
and large. Hind femur 4.2-4.6 mm long
and 5.2-6.2 X as long as wide. Hind tar-
somere 1 weakly compressed laterally and
0.50 X as wide as hind tibia near apex.
First metasomal tergum 1.8-2.1 X as long
as wide, with median dorsal carinae well
defined and strong (Fig. 17); in lateral
view decurved with a strong angle before

midlength (Fig. 18). Terga 2 to 4 each with
two large sublateral swellings with spars-
er punctures (Fig. 23). Metasomal tergum
2 (Fig. 23) with punctures almost coales-
cent with outlines polygonal, but sparser
and faded on sublateral swellings, and mi-
crosculpture well defined along posterior
and lateral borders, but with meshes
much smaller and flat along anterior bor-
der. Dorsal notch on ovipositor with an-
terior margin without a strong angle,
gradually sloping. Coloration: Female. —
Antenna with scape and pedicel reddish
black, and flagellum with flagellomeres
dorsally reddish brown and ventrally
brownish orange gradually turning red-
dish brown near apex. Head black except
anterior surfaces and apical half of lateral
surfaces of mandibles brown. Mesosoma
black except tegulae yellowish brown.
Wings with yellow infuscation. Legs or-
ange except hind tibia and tarsus reddish
black to black. Metasoma black. Male. —
Antenna with scape and pedicel brown,
and flagellum with flagellomeres dorsally
reddish brown and ventrally brown. Head
black except usually brown median lon-
gitudinal spot on face below tubercle,
brown to reddish black clypeus and
brown anterior surfaces and apical half of
lateral surfaces of mandibles. Mesosoma
black except tegulae yellowish brown.
Wings with yellow infuscation. Legs or-
ange except hind tibia and tarsus reddish
black to black. Metasoma black.

Etymology. — From the Greek bolbos
(swelling) and gaster (belly), referring to
the characteristic sublateral swellings on
metasomal terga 2 to 4.

Distribution. — Transcontinental in cold
temperate and boreal regions (Fig. 30).

Specimens examined. — 3 females and 6
males. Holotype male, "Estes Pk.
Colo.[rado] 7500 ft 7 mi. E. 8-18-48
Evans"; "Protarchus Det. W.R.M. Mason
'48". Condition of type: intact. [CNCI].
Paratypes: CANADA. BRITISH COLUM-
BIA:' Jesmond, 14. ix. 1938, J.K. Jacob
(1F,CNCI); Racing River, 2400',

Volume 8, Number 2, 1999

257

24.viii.1973, H.&M. Townes (1F,AEIC).
ONTARIO: Orrville, 21.vii.l958, L.L. Pe-
chuman (1M,AEIC). QUEBEC: Lac Rol-
land, R.I.F. 41, specimen no. 12150-B
(1M,CNCI). YUKON TERRITORY: 14
mi.E.Dawson, 1300', 30.vi.l962, R.E. Leech
(1F,CNCI) (used for SEM). UNITED
STATES. MAINE: Dryden, 30.viii.l959
(1M,AEIC); Roque Bluff, 10.viii.l907, J.A.
Cushman, "Pwtarclwides maiidibulnris" Al-
lotype # 25975, USNM (1M,USNM).
MICHIGAN: Huron Mts, 25.viii.1959, H.
Townes, "strong Pimpla odor" (1M,AEIC).
Remarks. — This species is closely related
to the Palearctic P. heros (Holmgren), both
species sharing as synapomorphies the
unique mandible shape and the large sub-
lateral swellings on terga 2 to 4. P. holbo-
gaster differs from P. hews by its predom-
inantly orange legs. Cushman's original
allotype of P. mamiibularis belongs to P.
bolbogaster.

Protarchus magnus (Davis)

(Figs. 1,8,9,22,26,30)

Znccnles mngnus Davis 1898 (1897):283.

Diagnostic combination. — Whole body
predominantly light colored (brown), hind
tibiae entirely light colored, ocelli en-
larged and hind ocelli separated by about
their diameter, dorsal notch on ovipositor
with anterior margin with strong angle,
abruptly sloping (Fig. 26).

Description. — Structure: Antenna with
47^8 flagellomeres. Ocelli enlarged and
sitting on a swelling, hind ocelli separated
by about their diameter (Fig. 1). Antennal
sockets in lateral view forming a moder-
ately strong angle with vertical axis of
compound eye. Frons not strongly de-
pressed (Fig. 1). Face 1.5-1.7 X as wide as
high. Median swelling of face very weak.
Upper mandibular tooth subequal in size
and shape to lower tooth. Notaulus strong
and sharply defined only at anterior end
of mesoscutum (Fig. 8,9). Epomia weak.
Mesopleuron with punctures separated by
0.5-2.0 of their diameter and microsculp-

ture with sculpticells very convex, making
surface mat; meshes well outlined on pos-
terior half below speculum, but gradually
fading anteriorly. Carinae of propodeum
strongly defined. Forewing 16.5-18.3 mm
long, areolet present and large. Hind fe-
mur 4.6-5.2 mm long and 6.8-8.0 X as
long as wide. Hind tarsomere 1 weakly
compressed laterally and 0.50-0.55 X as
wide as hind tibia near apex. First meta-
somal tergum 2.1-2.6 x as long as wide,
with median dorsal carinae reduced to a
median furrow; in lateral view weakly
and regularly curved at midlength. Sub-
lateral swellings on terga absent. Metaso-
mal tergum 2 (Fig. 22) with punctures
round and not coalescent, and microsculp-
ture uniform over all tergum, with sculp-
ticells convex. Dorsal notch on ovipositor
with anterior margin with a strong angle,
apruptly sloping (Fig. 26). Coloration: Fe-
male.— Body uniformly brown except the
following. Yellowish brown parts are an-
tennae, clypeus, mandibles, hind corner
lobe of pronotum, tegula, subalar promi-
nence, mesepimeron and legs beyond cox-
ae. Black parts are base of first antennal
flagellomere and narrow bands at apices
of metasomal terga 3 to 8. Wings with yel-
low infuscation. Color variation: Some fe-
males have extensive black markings:
frons, vertex and genae, propleuron, all
pronotum except upper and posterior
margins and hind corner lobe, dorsal sur-
face of forecoxa, anteromedian band on
mesoscutum to half of mesoscutum length
and two bands on mesoscutum lateral to
notauli, upper fourth of epicnemium, up-
per fourth to third of mesopleuron except
anterior and posterior margins, subtegular
ridge, speculum, mesosternum except an-
terolateral corners, large central spot on
metasomal tergum 2 and whole metasoma
beyond tergum 2.

Distribution. — Northeastern United
States and Southeastern Canada (Fig. 30).

Specimens examined. — 10 females. Holo-
type female, 4 labels, "N!N"; "HoloTYPE
4340"; "Zaccales magnus Davis"; "Collec-

258

Journal of Hymenoptera Research

Figs. 25-27. 25, P. testatoriiis, metasonial tergum 2, dorsal. 26-27, ovipositor; 26, P. magnum; 27, P. sorb/.

tion of THE ACADEMY OF NATURAL
SCIENCES of Philadelphia. ANSP". Con-
dition of type: missing left flagellum be-
yond flagellomere 11, right antenna, left
fore wing; right fore and hind wings
pinned below specimen. [ANSP]. Other
specimens: CANADA. QUEBEC: Joliette
Co., Ste-Beatrix, 24.VIII.1978 (1F,LLIC)
(used for SEM). UNITED STATES. MICH-
IGAN: Midland, vii.1950 (1F,AEIC). NEW
YORK: Allegany St.Pk., 30.vii.l938
(1F,USNM), 31.vii.l938 (1F,USNM), A.R.
Shaddle; Essex Co., Keene Valley,
17.ix.l917, H. Nortman (1F,ANSP); Six
Miles Creek, Ithaca, 17.vii.l947, J.G. Fran-
clemont (3F,AEIC). PENNSYLVANIA:
Glenside, "10.12.1929", G.G. Sleesman
(1F,USNM).

Protarchus mellipes (Provancher)

(Figs. 12,31)

Coelocentrus mellipes Provancher 1886:113.
Protarchoides mellipes: Walley 1938:231.
Protarchoides pnllipes Cushman 1927:15. Synon-
ymy by Walley 1938:231.

Diagnostic combination. — Epomia clearly
defined and prominent (Fig. 12), ocelli en-
larged and hind ocelli separated by about
their diameter, face entirely black, or at
most with yellowish brown median spot,

mesosoma and metasoma almost entirely
black, hind tibiae entirely light colored.

Description. — Structure: Antennal flagel-
lum with 43^6 (female) and 41^3 (male)
flagellomeres. Ocelli enlarged and sitting
on a swelling, hind ocelli separated by
about their diameter. Antennal sockets in
lateral view forming a moderately strong
angle with vertical axis of compound eye.
Frons not strongly depressed. Face 1.5-1.7
X as wide as high. Median swelling of
face moderate. Upper mandibular tooth
subequal in size and shape to lower tooth.
Notaulus strong and sharply defined only
at anterior end of mesoscutum. Epomia
clearly defined and prominent (Fig. 12).
Mesopleuron with punctures separateed
by 0.5-2.0 of their diameter and micro-
sculpture with sculpticells convex, making
surface slightly mat; meshes well outlined
on posterior half below speculum, but
gradually fading anteriorly. Carinae of
propodeum well defined and strong. Fore-
wing 13.8-16.0 (female) and 12.3-13.2
(male) mm long. Areolet present and
large. Hind femur 3.9-4.9 mm long and
6.0-7.0 X as long as wide. Hind tarsomere
1 weakly compressed laterally and 0.50-
0.55 X as wide as hind tibia near apex.
First metasomal tergum 2.3 x as long as

Volume 8, Number 2, 1999

259

wide with median dorsal carinae reduced
to a median furrow and; in lateral view
weakly and regularly curved at mid-
length. Sublateral swellings on terga ab-
sent. Metasomal tergum 2 with punctures
round and not coalescent, and microsculp-
ture uniform over all tergum with sculp-
ticells convex. Dorsal notch on ovipositor
with anterior margin without a strong an-
gle, gradually sloping. Coloration: Fe-
male.— Antenna with scape and pedicel
black or black with ventral surfaces yel-
lowish orange, and flagellum with basal
section yellowish orange, except basal half
of first flagellomere and dorsal surfaces of
first few flagellomeres generally black,
and apical section black to brown. Head
black except clypeus reddish brown to
yellowish brown, lower margin of face
along clypeus sometimes yellowish brown
and mandibles black to yellowish brown.
Mesosoma black except yellowish brown
to reddish black tegula. Wings with yel-
low infuscation. Legs yellowish brown ex-
cept fore, middle and hind coxae and usu-
ally fore, middle and hind trochanters 1
black, usually a narrow brownish orange
longitudinal line on black dorsal surface
of hind coxa and brownish orange fore
and middle tarsomeres 5. Metasoma
black. Male. — Antenna with scape and
pedicel black, and flagellum with basal
section yellowish orange, except dorsal
surfaces of first few flagellomeres black
and with apical section dark brown. Head
black except clypeus yellowish brown
and, in one specimen, faint traces of yel-
lowish brown on face. Mesosoma black
except tegula yellowish brown. Wings
with yellow infuscation. Legs yellowish
brown except fore, middle and hind coxae
and trochanters 1 reddish black. Metaso-
ma black.

Distribution. — Transcontinental in bore-
al region (Fig. 31).

Specimens examined. — 5 females and 2
males. Type material: Lectotype of Coelocen-
trus mellipes Provancher, designated by
Barron 1975:508; male, 6 labels: "Holotype

male Coleocentrus mellipes Provancher No.
4235"; "Coleocentrus n. spec!"; "G 484";
"This must be the type of Coleocentrus mel-
lipes Prov. which g.[ahan] & R.[ohwer]
could not locate. It was evidently returned
to geddes by Prov. & the label [# 2] in red
ink is in Geddes hand (note by G.S. Wal-
ley april/20/37)"; "Lectotype Coleocentrus
mellipes Provancher Comeau '40"; "LEC-
TOTYPE Coleocentrus mellipes Provancher
G 484 Barron '71". Specimen from Rocky
Mountains according to Walley, 1938.
Condition of type: missing right fore and
middle legs beyond coxae, left middle leg
beyond tibia and both hind tarsi but tar-
someres 1-2 of one hind leg glued on first
label. [CNCI]. Holotype of Protarchoides
pallipes Cushman. Female. 3 labels: "Ed-
monton, Alberta 23.viii.1916 G. Salt";
"Type No. 40444 U.S.N.M."; Protarchoides
pallipes Type. Cush.". Condition of type:
missing entire left antenna and right hind
tarsus; left hind tarsus broken and re-
glued between tarsomeres 2 and 3.
[USNM]. Other specimens. CANADA. AL-
BERTA: 15 mi.E.Morley, 14.viii.l962, K.C.
Herrmann (1F,CNCI) (used for SEM); Ed-
monton, 23.Vm.1926, G. Salt, paratype #
40444, USNM (1F,USNM). BRITISH CO-
LUMBIA: Houston, 17.vi.l959, Forest In-
sect Survey specimen no. 58-1792-0119, ex.
Trichiosoma triangulum (1M,CNCI); Rob-
son, 13.ix.l949, H.R. Foxlee (1F,CNCI).
ONTARIO: Smoky Falls, near Kapuska-
sing, 9.viii.l937, R.V. Whelan (1F,CNCI).

Remarks. — The specimen used by Pro-
vancher, collected by G. Geddes in the
Rocky Mountains (Provancher 1886),
could not be located by Rohwer in his
1915 visit to the Provancher collection
(Gahan and Rohwer 1917, Cushman and
Rohwer 1920). Walley (1938) discovered a
specimen in the Geddes collection, donat-
ed to CNCI, labelled "Coleocentrus n sp"
and agreeing with Provancher's descrip-
tion. He accepted this specimen as the one
originally used by Provancher. Barron
(1975) designated the specimen as lecto-
type.

260

Journal of Hymenoptera Research

Figs. 28-29. Forewing, showing areolet: 28, P. sorhi; 29, P. testatcriKS.

Protarchus pallidicomis (Walley)

(Fig. 30)

Protarchoides pnllidicoruis Walley, 1938:231.

Diagnostic combination. — Epomia clearly
defined and prominent (Fig. 12), ocelli en-
larged and hind ocelli separated by about
their diameter, face entirely brown or yel-
lowish brown, mesosoma and metasoma
predominantly brownish black with ex-
tensive yellowish orange and brown
markings, hind tibiae entirely light col-
ored.

Description. — Structure: Antenna with
45 (female) flagellomeres. Ocelli enlarged
and sitting on a swelling, hind ocelli sep-
arated by about their diameter. Antennal
sockets in lateral view forming a moder-
ately strong angle with vertical axis of
compound eye. Frons not strongly de-
pressed. Face 1.5 X as wide as high. Me-
dian swelling of face moderate. Upper
mandibular tooth subequal in size and
shape to lower tooth. Notaulus strong and
sharply defined only at anterior end of
mesoscutum. Epomia clearly defined and
prominent. Mesopleuron with punctures
separated by 0.5-2.0 of their diameter
apart and microsculpture with sculpticells
convex, making surface slightly mat and
meshes well outlined on posterior half be-
low speculum, but gradually fading ante-
riorly. Carinae of propodeum strongly de-
fined. Forewing 14.5 (female) and 12.8
(male) mm long, areolet present and large.
Hind femur 4.3 (female) and 4.7 (male)

mm long and 6.9 (female) and 7.8 (male)
X as long as wide. Hind tarsomere 1
weakly compressed laterally and 0.50 x as
wide as hind tibia near apex. First meta-
somal tergum 2.4 x as long as wide, with
median dorsal carinae reduced to a me-
dian furrow; in lateral view weakly and
regularly curved at midlength. Sublateral
swellings on terga absent. Metasomal ter-
gum 2 with punctures round and not co-
alescent, and microsculpture uniform over
all tergum with sculpticells convex. Dorsal
notch on ovipositor with anterior margin
without a strong angle, gradually sloping.
Coloration: Female. — Antenna brownish
orange except dorsal half of scape brown-
ish black. Head brown except frons, area
around ocelli and vertex behind ocelli
brownish black. Mesosoma brownish
black except the following. Yellowish or-
ange parts are hind corner lobe of prono-
tum, tegula, subalar prominence, anterior
margin of mesopleuron, mesepimeron and
upper division of metapleuron. Brown
parts are sublateral longitudinal bands on
mesoscutum along notauli from anterior
margin to three quarters of scutum length
and lateral bands along mesoscutum mar-
gin from wing tegula to end of mesoscu-
tum, lower half of epicnemium, anterolat-
eral corner of mesosternum, lower margin
of mesopleuron, speculum, scutellum in-
cluding axillae, postscutellum, upper an-
terior corner of metapleuron, anterior half
of area lateralis of pronotum and along
median furrow of propodeum. Wings

Volume 8, Number 2, 1999

261

• P
A P
■ P

bolbogai^ter

magnus

pallidicoirnis

Fig. 30. Distribution of P. I'ltlbogaster (circles), P. magnus (triangles) and P. pallidicornis (squares).

with yellow infuscation. Legs yellowish
brown except faint traces of brownish
black on dorsal surface of forecoxa and
apical half of dorsal surfaces of mid and
hind coxae. Metasoma brownish black ex-
cept first tergum brown and faint traces of
brown on second tergum. Male. — Antenna
with scape and pedicel dorsally brownish
black and ventrally yellowish brown, and
flagellomere 1 yellowish brown (rest of
antennae missing). Head brownish black
except face, clypeus, malar space and
mandibles yellowish brown. Mesosoma
brownish black except the following. Yel-
lowish orange parts are hind corner lobe
of pronotum, tegula, subalar prominence
and mesepimeron. Brown parts are two
sublateral longitudinal bands on mesoscu-
tum along notauli from anterior margin to
three quarters of scutum length and two
lateral bands along margins of scutum
from base of sublateral band to three quar-
ters of scutum length, lower two thirds of
epicnemium, anterolateral corner of me-
sosternum, lower margin of mesopleuron,
speculum, scutellum including axillae,
postscutellum and entire propodeum ex-

cept faint brownish black on metapleuron.
Wings with yellow infuscation. Legs yel-
lowish brown. Metasoma brownish black
except first tergum and basal half of sec-
ond tergum brown and faint brown traces
on anterolateral corners of third tergum.

Distribution. — Ontario, Maine (Fig. 30).

Specimens examined. — 1 female and 1
male. Holotype, examined, female, 2 labels:
"Holotype female Protarchoides pallidicor-
nis Walley No. 4410"; "Smokey Falls,
Ont[ario] (near Kapuskasing) Aug 4, 1937
R.V. Whelan". Condition of type: missing
hind tarsomeres 4-5. [CNCI]. Other speci-
men. UNITED STATES. MAINE: Glen-
burn, 5.vii.l928, Gypsy moth Lab, 10088
NIG, ex. Tenthredinid (!) (1M,USNM).

Remarks. — This species is closely related
to P. mellipes. The prominent epomia (Fig.
12) is a synapomorphy.

Protarchtis sorbi (Ratzeburg)

(Figs. 3,6,7, 1 3, 1 5, 19,20,24,27,28,31 )

Trypho)! sorbi Ratzeburg 1844:126. Holotype

lost.
PsUosargc (!) longipes Ashmead, in Slosson 1902:

321 {noincn nmiuw).

262

Journal of Hymenoptera Research

Protarchoides longipes Cushman 1922:26. Protar-
chus longipes: Townes, 1945:505. NEW SYN-
ONYM.

Protnrchoides mainiibidnris Cushman 1924:9. Des-
ignated synonym to P. longipes by Townes
1945:505. NEW SYNONYM.

Diagnostic combination. — Hind tarsomere
1 strongly compressed laterally and 0.55-
0.70 as wide as hind tibia near apex, me-
tasomal tergum 1 decurved with a strong
angle (Fig. 20), face of male yellow, me-
sosoma and metasoma predominantly
black.

Description. — Structure: Antenna with
36-45 flagellomeres. Ocelli of moderate
size and not sitting on a swelling, hind
ocelli separated by more than their diam-
eter. Antennal sockets in lateral view
forming a moderately strong angle with
vertical axis of compound eye. Frons not
strongly depressed. Face (Fig. 3) 1.6-1.9 X
as wide as high. Median swelling of face
almost absent. Upper mandibular tooth
subequal in size and shape to lower tooth.
Notaulus weak and reduced to shallow
impressions (Figs. 6,7). Epomia indistinct
or very weak. Mesopleuron (Fig. 13) with
punctures separated by less than 0.1 of
their diameter and microsculpture vari-
able in different specimens from absent
over all mesopleuron to almost absent
with meshes absent to lightly convex be-
low speculum to irregularly striated
through fusion of sculpticells on posterior
0.5 of mesopleuron below speculum. Ca-
rinae of propodeum strongly defined (Fig.
15). Forewing 9.9-13.9 (female) and 9.8-
12.5 (male) mm long, areolet present and
large (Fig. 28). Hind femur 2.8-3.9 mm
long and 5.2-6.5 X as long as wide. Hind
tarsomere 1 strongly compressed laterally
and 0.55-0.70 x as wide as hind tibia near
apex. First metasomal tergum 1.4-1.9 X as
long as wide, with median dorsal carinae
well defined and strong (Fig. 19) or re-
duced to a median furrow; in lateral view
decurved with a strong angle before mid-
length (Fig. 20). Sublateral swellings on
terga absent. Metasomal tergum 2 (Fig. 24)

with punctures very dense, polygonal,
crater-like but not coalescent, and micro-
sculpture with sculpticells convex along
posterior border of tergum, otherwise
completely faded on the rest of tergum.
Dorsal notch on ovipositor with anterior
margin without a strong angle, gradually
sloping (Fig. 27). Coloration: Female. — An-
tenna with scape and pedicel black, and
flagellum with flagellomeres dorsally
brownish black and ventrally brownish
orange and gradually turning brownish
black near apex, or sometimes entire fla-
gellum brownish black. Head black except
anterior surface of mandible to entire
mandible yellowish orange, and clypeus
brownish orange to reddish black or
black. Mesosoma black except hind corner
lobe of pronotum and tegula reddish
brown. Wings hyaline. Legs orange except
sometimes traces of reddish black anteri-
orly at base of each coxae, and apical two
thirds to nine tenth of hind tibia and
whole hind tarsus reddish black or black.
Metasoma black. Male. — Antenna with
scape and pedicel reddish black except
usually yellow ventral surface of scape
and sometimes yellow ventral surface of
pedicel, and flagellum with flagellomeres
dorsally brownish black and ventrally
brownish black to brownish orange. Head
black except orbits between compound
eyes and antennal sockets yellow, and
face, clypeus and mandibles yellow. Me-
sosoma black except tegula yellow. Wings
hyaline. Legs orange except apical six
tenths of hind tibia and whole hind tarsus
reddish black or black and sometimes yel-
low fore and middle coxae, fore, middle
and hind trochanters, foretibia, foretarsus
and basal half of middle tibia. Metasoma
black.

Distribution. — Palearctic (Sweden, Fin-
land, Russia) and transcontinental in
North America (Fig. 31).

Specimens examined. — 17 females and 14
males. Type material. Holotype of Protar-
choides longipes Cushman. Female. 5 labels:
"MT.WASH'[ingto]N."; "39"; "Type No.

Volume 8, Number 2, 1999

263

• P. sorbi (
A P. mellipesi

Fig. 31. Distribution of P. sinbi (circles) and P. melliftef (triangles).

<^^t^t:>

/"

p. testaioriu^
P. atrofacies

Fig. 32. Distribution of P. tc^tatoriin (circles) and P. u/ra/flot's (triangles).

264

Journal of Hymenoptera Research

25030 U.S.N.M."; "Psilosage longipes .Type
Ashm[ead]"; "Protarchoides longipes Type.
Cush.". [New Hampshire, A.T. Slosson,
1902]. Condition of type: missing left hind
tarsus; both antennae broken and glued
on first label. [USNM]. Holotype of Protar-
choides mandibiilaris Cushman. Female. 4
labels: "Wellington B.C. 28.V1I.[19]04";
"1260"; "Type No 25975 U.S.N.M."; "Pro-
tarchoides mandihularis Type. Cush.". Con-
dition of type: missing: left antenna be-
yond flagellomere 7, right antenna beyond
flagellomere 37, right foreleg tarsus be-
yond tarsomere 1 and right hind leg tarsus
beyond tarsomere 1; right antenna section
containing flagellomeres 4 to 37 and right
hind leg beyond coxa broken and glued
on locality label. [USNM]. Other specimens:
CANADA. ALBERTA: Edmonton,
28.vii.1926, E.H. Strickland (1M,AEIC);
Entranse, emerged 21.iii.l950, Forest In-
sect Survey specimen no. A113313, ex. Tri-
chiosoma sp. (1M,CNCI). BRITISH CO-
LUMBIA: Summit Lake, Mile 392, Alaska
Hwy, 4600', 16.vii.l959 (1F,CNCI), 4700',
15.vii.l959 (1F,CNC1), 5000', 6.vii.l959
(1M,CNCI); Stone Mtn Nat. Pk., "3800',
12.vii.l975, 13.vii.l973, 18.vii.l973,
20.vii.l973 (7M,AEIC). MANITOBA:
Christopher Lake, emerged 4.iii.l963, For-
est Insect Survey specimen no. 1914(02),
ex. Trichiosorna triangulum (1F,CNCI); Tur-
tle Mtn., 21.vii.l953, Brooks-Kelton
(1F,CNCI); Wanless, 29.vi.1961, H.E. Mil-
liron (1F,CNCI). PRINCE EDWARD IS-
LAND: Alberton, 15.vii.l940, G.S. Walley
(1F,CNCI). QUEBEC: Hemmingford,
l.vii.l928, G.H. Hammond (1F,CNCI);
Aylmer, 20.V.1920, G.H. Hammond, ex.
Trichiosorna sp. (1M,CNCI). SASKATCH-
EWAN. Buffalo Narrov^s, emerged
4.iii.l964, Forest Insect Survey specimen
no. 63W-2241(03), ex. Trichiosorna triangu-
laris (!) (1F,CNCI); Candle Lake, emerged
10. ii. 1951, Forest Insect Survey specimen
no. W50-42786, ex. Trichiosorna triangulum
(1M,CNC1); Parr Hill, emerged 17.ii.l961,
Forest Insect Survey specimen no. W61-
2392(04), ex. Trichiosorna triangulum

(1F,CNCI); Waskesiu, 21. vi. 1938, J.G.
Rempel (1F,AEIC). UNKNOWN PROV-
INCE: Crimson Lake, emerged before
12.vii.l961, Forest Insect Survey specimen
no. 60A1409-03, ex. Trichiosoma sp.
(1F,CNCI) (used for SEM). UNITED
STATES. ALASKA: Seaward, <300',
25.vii.1951, W.J. Brown (1F,CNCI).
OREGON: Mt. Hood, 3500', 19.vii.l978,
24.vii.1978, H.&M. Townes (2F,AEIC).
WASHINGTON: Mt. Rainier, 4200',
15.vii.l940, H.&M. Townes, "odor like
Pirnpla" (1M,AEIC). UNKNOWN LOCAL-
ITY. Reared specimen labelled "99 88281"
(1M,CNCI).

Protarchus testatorius (Thunberg)

(Figs. 2,4,10,11,16,21,25,29,32)

Ichneiinion testatorius Thunberg 1822:276. Lec-
totype (designated by Roman, 1912) not ex-
amined.

Tryphon ruftis Gravenhorst 1829:200. Holotype
lost.

Mesoleius (Protarchus) welanurus Thomson 1895:
2020. Leptotype designation and synonymy
by Viitasaari 1979:34. Lectotype not exam-
ined.

Diagnostic combination. — Areolet very
small or sometimes absent (Fig. 29), me-
tasoma black with extensive orange mark-
ings on tergites 2 to 4.

Description. — Structure: Antenna with
40-44 flagellomeres. Ocelli of moderate
size and not sitting on a swelling, hind
ocelli separated by more than their diam-
eter (Fig. 2). Antennal sockets in lateral
view forming a very strong angle with
vertical axis of compound eye. Frons
strongly depressed (Fig. 2). Face 1.3-1.4 X
as wide as high. Median swelling of face
moderate. Upper mandibular tooth sub-
equal in size and shape to lower tooth
(Fig. 4). Notauli strong and sharply de-
fined to middle of mesoscutum length
(Figs. 10,11). Epomia indistinct. Mesopleu-
ron with punctures separated by 0.5-2.0 of
their diameter and microsculpture gener-
ally absent, at most expressed as slightly
convex sculpticells below speculum. Ca-

Volume 8, Number 2, 1999

265

rinae of propodeum weakly defined to al-
most absent (Fig. 16). Forewing 11.2-13.3
mm long. Areolet in forewing very small
(Fig. 29) or sometimes absent. Hind femur
3.6-4.4 mm long and 6.0-6.8 x as long as
wide. Hind tarsomere 1 weakly com-
pressed laterally and 0.45-0.50 X as wide
as hind tibia near apex. First metasomal
tergum 1.8-2.2 X as long as wide, with
median dorsal carinae reduced to a me-
dian furrow (Fig. 21); in lateral view
weakly and regularly curved at mid-
length. Sublateral swellings on terga ab-
sent. Metasomal tergum 2 (Fig. 25) with
punctures very dense, slightly polygonal
but not coalescent, but density decreasing
towards posterior margin, and with mi-
crosculpture well defined on apical 0.3,
flat in middle 0.3 and completely faded
basally. Dorsal notch on ovipositor with
anterior margin without a strong angle,
gradually sloping. Coloration: Female. —
Antenna with scape and pedicel reddish
black, and flagellum brownish orange ex-
cept basal portion of flagellomere 1 red-
dish black and dorsal surfaces of first few
flagellomeres reddish black gradually
turning brownish orange to yellowish or-
ange near apex. Head black except the fol-
lowing. Yellow parts are orbits between
eyes and antennal sockets, clypeus, man-
dibles and face except narrow median red-
dish black line below tubercle. Brownish
orange parts are genal orbits from poste-
rior ocelli down to two thirds to a quarter
of eye height. Mesosoma black except the
following. Yellow parts are tegula, scutel-
lum, postscutellum and often four small
spots on anterior margin of mesoscutum
on the sides of notauli. Wings with light
yellow infuscation. Legs with coxae black
except anteromedian line on or whole an-
terior surface of forecoxa yellow and an-
teroapical spot on middle coxa yellow.
Trochanters 1 black except anterior sur-
face of foretrochanter 1 and apex of dorsal
surface of middle trochanter 1 yellow.
Fore and middle legs beyond trochanters
1 yellowish orange except ventral basal

quarter to half of middle femur reddish
black. Hind trochanter 2 yellowish orange.
Hind femur black except yellowish orange
at base and apex. Hind tibia with basal 0.6
yellowish orange and apical 0.4 reddish
black. Hind tarsi yellowish orange to
brownish orange. Metasoma black with
the following brownish orange: apical 0.25
of tergum 1, apical 0.33 to 0.5 of tergum 2
to entire tergum 2 except lateral black
markings on basal 0.5, entire tergum 3 ex-
cept sometimes a large median black spot,
basal 0.15 of tergum 4 with a small sub-
apical spot to entire tergum 4 except api-
cal 0.1. Color variatio?is: Two females have
brownish orange markings on propodeum
anterior to postpectal carina and lateral to
median furrow and extending laterally be-
low lateral carinae. These markings are
present in most Palearctic specimens of P.
testatorius.

Distribution. — Palearctic (Sweden, Fin-
land, Germany, Russia) and transconti-
nental in North America (Fig. 32).

Remarks. — The lectotypes from Europe
have not been examined because a good
series of Palearctic material (AEIC,
USNM) was studied and found to be con-
specific with the Nearctic material.

Specimens examined. — 28 females. AL-
BERTA: Banff, 18.viii.l946, E.H. Strickland
(1F,AEIC); Banff, Black's Camp Ground,
emerged 28.vi.1958, Forest Insect Survey
specimen no. 58A108-03, ex. Trichiosoma
sp. on alder (1F,CNCI); Cameron Lake,
9.vii.l949, C.P. Alexander (1F,AE1C); Can-
yon Creek, emerged 9.iii.l953, Forest In-
sect Survey specimen no. A2182A, ex. Tri-
chiosoma sp. (1F,CNCI); Eisenhower Junc-
tion, Banff National Park, 9.viii.l962, K.C.
Hermann (1F,CNCI); Fawcett, emerged
9.vii.l956, Forest Insect Survey specimen
no. 55A1438-12, ex. Trichiosoma sp.
(1F,CNCI); Kanamaskis, emerged
23.ii.1949, Forest Insect Survey specimen
no. A-455-K, ex. Trichiosoma sp. (1F,CNCI);
Lake Louise, 5600', 26.vii.1938, G.S. VVal-
ley (1F,CNCI); Nordegg, 21.vii.l926, E.H.
Strickland (1F,CNCI); 18 mi.W. Strachan,

266

Journal of Hymenoptera Research

emerged 24.iii.1959, Forest Insect Survey
specimen no. 58A1821-02, ex. Trichiosoma
sp. (1F,CNCI). BRITISH COLUMBIA:
Hope Mts, 20.viii.l931, A.N. Gartrell
(1F,CNCI) (used for SEM); Saanich,
emerged 23.V.1958, Forest Insect Survey
specimen no. 58-3-01-A, ex. Trichiosoma
triaugulum (1F,CNCI). ONTARIO: Sud-
bury, 1891 (1F,CNCI). QUEBEC: Lac St-
Jean, 20.viii.l939, P.L. Mercier (1F,AEIC);
Kazabazua, 28.viii.1928, G.H. Fisk
(1F,CNCI); Ste-Agathe des Monts,
7.viii.l937, G.S. Walley (1F,CNCI). YU-
KON TERRITORY: Whitehorse,
7.viii.l948, W.R. Mason (1F,CNCI). UN-
KNOWN PROVINCE: Hot Springs Road,
2.viii.l902, N.B. Sanson (1F,CNCI).
FRANCE. Miquelon Island, 15.viii.l990,
D. Abraham, collected in bog (1F,CNCI).
UNITED STATES. ALASKA: Thompson
Pass, 14.viii.l973, H.&M. Townes
(1F,AEIC); Tsaina River, 17.viii.l973,
18.viii.l973, H.&M. Townes (2F,AEIC).
COLORADO: Gould, 5.viii.l974,
6.viii.l974, H.&M. Townes (3F,AEIC).
NEW HAMPSHIRE: Franconia
(1F,USNM). WASHINGTON: Mt. Rainier,
22.vii.1940, H.&M. Townes, "strong Pim-
pla odor" (2F,AEIC).

ACKNOWLEDGMENTS

The museum curators listed in the paper are sin-
cerely acknowledged for their cooperation in the loan
of specimens. For reviewing the manuscript, 1 thank
Henri Goulet, from the Biological Resources Program,
ECORC, Agriculture and Agri-Food Canada, Ottawa,
as well as the three external reviewers. Dr. Goulet's
contribution in accurately describing microsculpture
is highly appreciated. Scanning Electron Micrographs
were taken by K. Bolte, from ECORC. I want to ded-
icate this study to Dr. John Barron (1932-1997), who
has greatly helped me to improve this paper and who
has always encouraged me to study Ctenopelmati-
nae.

LITTERATURE CITED

Allen, R. T. and G. E. Ball. 1980. Synopsis of Mexican
taxa of the Lo.\nth1ruf series (Coleoptera: Carabi-
dae: Pterostichini). Transactions of tlw American
Entomological Socicti/. 105:481-576.

Arnett, R. H. and G. A. Samuelson. 1986. The insect
and spider collections of llie world. E.J. Brill/ Flora

and Fauna Publications. Gainesville, Florida. 220

PP-

Barron, J. R. 1975. Provancher's collections of insects,
particularely those of Hymenoptera, and a study
of the types of his species of Ichneumonidae. Na-
tiiraliste Camidien. 102:387-591.

Cushman, R. A. 1922. On the Ashmead manuscript
species of Ichneumonidae of Mrs. Slosson's
Mount Washington lists. Proceedings of the United
States National Museum. 61(8);l-30.

Cushman, R. A. 1924. On the genera of the Ichneu-
mon-flies of the tribe Paniscini, Ashmead, with
discussions and descriptions and discussion of
related genera and species. Proceedings of the
United States National Museum. 64(20):l-48.

Cushman, R. A. 1927. Miscellaneous notes and de-
scriptions of Ichneumon-flies. Proceedings of the
United States National Museum. 72(13):l-22.

Cushman, R. A. and S. A. Rohwer. 1920. The North
American Ichneumon-flies of the tribe Acoeniti-
ni. Proceedings of the United States National Muse-
um. 57:503-523.

Davis, G. C. 1898(1897). A review of the Ichneumonid
subfamily Tryphoninae. Transactions of the Amer-
ican Entomological Society. 24:193-348.

Foerster, A. 1869. Synopsis fiir Familien und Gattun-
gen der Ichneumonen. Verli. Naturli. Ver. Rliein-
lande. 25:135-227.

Gahan, A. B. and S. A. Rohwer. 1917. Lectotypes of
the species of Hymenoptera (except Apoidea) de-
scribed by Abbe Provancher. Canadian Entomolo-
gist. 49:391^00.

Gauld, I. D. 1984. An introduction to the Ichneumonidae
of Australia. British Museum (Natural History).
London. 413 pp.

Gravenhorst, J. L. C. 1829. Ichneumonologia Europaea,
Vratislaviae, 1. 989 pp.

Holmgren, A. E. 1876. Dispositio synoptica meso-
leiorum scandinaviae. Koiigl. Svenska Vetenskap-
sakad. Handl. (N.F.) 13:3.

Kaur, R. 1989. A revision of the mesoleiinae genus
Dentimachus Heinrich (Hymenoptera: Ichneu-
monidae). Oriental Insects. 23:291-305.

Leblanc, L. 1989. A taxonomic revision of the Nearctic
species of Himerta (Hymenoptera: Ichneumoni-
dae). Contributions of the American Entomological
Institute. 25(3):l-76.

Provancher, L. 1886. Additions et corrections nu volume
11 de la faiine entomologique du Canada. C. Darveau.
Quebec. 475 pp. Fam IV. Ichneumonidae pp. 29-
121.

Ratzeburg, J. T. C. 1844. Die Ichneumonen der Forstin-
sekten in forstliclier und entomologisclwr Beziehung.
Ein Anhang zur Abbildung und Beschreibung
der Forstinsekten. 1, VlII:I-224.

Roman, A. 1912. Die Ichneumonidentvpen C.P. Thun-
bergs. Zool. Bidrag Upp'^ala 1:229-293.

Slosson, A. T. 1902. Addilional list of insects taken in

Volume 8, Number 1, 1999

267

Alpine Region of Mt. Washington. Eiiloniological
News. 13:319-321.

Thomson, C. G. 1895. Opiaai EntoiHotci\;ica. Fascicle
19: pp. 1971-2137.

Thunberg, C. P. 1822. Ichneumonidea, Insecta Hy-
menoptera illustrata. Memoires de I'Acadernie
Iniperiale des Sciences de St.Petershourg. 8:249-
281.

Townes, H. K. 1945. A catalogue and reclassification
of the Nearctic Ichneumonidae (Hvmenoptera),
Memoirs of the American Eiitoinolo^;iail Institute.
No 11 (2):480-925.

Townes, H. K. 1970. The genera of Ichneumonidae,
part 3. Memoirs of the American Entomological In-
stitute. 13:1-307.

Townes, H. K., S. Momoi and M. Townes. 1965. A
catalogue and reclassification of the Eastern Pa-

learctic Ichneumonidae. Memoirs of the American

Entomological Institute. 5:1-661.
Viitasaari, M. 1976. Protarchus sorbi (Ratz.) new to the

Finish fauna. Annates Entomologici Fennici 42:146.
Viitasaari, M. 1979. A studv of the Palearctic species

of the genus Protarchus Forster (Hvmenoptera,

Ichneumonidae). Notiilae Ent. 59:33-39.
Vikberg, V. and M. Viitasaari. 1991. Tncluosoma naiuie

sp. n., a monophag on Betula nana from Finland

(Hvmenoptera, Cimbicidae). Entomologica Fcnni-

ca.' 2:67-77.
VValley, G. S. 1938. Notes on the genus Protarchoides

(Hymen., Ichneumonidae. Canadian Entomologist.

70:230-232.
Woldstedt, F. W. 1877. Beitrag zur Kenntnis den um

St. Petersbourg vorkommenden Ichneumoniden.

Bulletin de I'Acadernie Imperiale des Sciences de

St. Petersbourg. 23:432-460.

J. HYM. RES.
Vol. 8(2), 1999, pp. 268-269

BOOK REVIEW

Geologische und biologische Entomookologie
der rezenten Seidenbiene CoUetes. Volume I.
Detlef Mader. Logabook, Koln, 1999.
xliii + 807. Price: Germany DM 98.00 or
50.00 Euro (hardcover). ISBN 3-87361-263-1.

In our concept-driven time dominated
by molecular biology, few authors dare to
publish lengthy accounts on pure natural
history, summarizing raw facts of organ-
ismal life or the idiosynchrasies of a par-
ticular species. I happen to enjoy pulling
those books off library shelves and im-
mersing myself in organismal trivia. I fre-
quently find a library note slipped into
those books saying that they have not
been circulating for a decade and that, un-
less soon used, they will be permanently
impounded in compact storage. So I reg-
ularly check out these books from the li-
brary, often to return them immediately. I
like to think that this fools the librarians,
such that they will grant these books an-
other decade of accessible existence on a
prime shelf. And I imagine that some fu-
ture scientist will stumble, like me, over
these books and keep alive some natural
history minutia that otherwise may be
buried forever.

Detlef Mader's book on the Geological
and Biological Entomo-Ecology of CoUetes
Bees is one of those natural history ac-
counts. It is thick in detail, particularly the
nesting idiosynchrasies of the species Col-
letes daviesanus. It is a specialist's book,
therefore, and its audience may be rather
limited. But it has enourmous depth in
natural history, inspired by a Humbold-
tian appetite for careful and complete doc-
umentation. Leafing through pages and
pages of flower records, or the geological
details of the substrate used by C. davie-
snniis for nest construction, I came to ad-
mire Mader for taking the time to sum-
marize his decades of work on the nesting

habits of this species. Few of us ever make
the time to do that for the species that are
close to our heart.

Bees in the basal bee genus CoUetes are
best known for their unique nest architec-
ture. The typical coUetid bee is solitary,
constructs an underground nest (some
species use twigs or rotting wood), and
lines the nest tunnel with a cellophanelike
tapestry derived from glandular secre-
tions. A series of cells are constructed in-
side this tunnel, separated by partitions of
additional tapestry. The tapestry linings
are tranlucent, giving the appearance that
these bees keep their brood in a series of
plastic bags. Hence also the vernacular
name of the plastic-bag bee.

Most colletids prefer to nest in level
ground, but C. daviesnnus, the most com-
mon species in central Europe, is an ex-
ception. It prefers to nest in vertical em-
bankments or cliffs, and there only in sub-
strate of particular kinds, such as loose
sandstone. Much of Maders' book is de-
voted to documenting these substrate
preferences for populations in central Ger-
many, with comparative references to oth-
er populations throughout Europe. This
treatment has a rather geological flavor,
and I admit that the geological aspects of
this work were rather lost on me, as I as-
sume they will be lost on anyone unfa-
miliar with the geology and stratigraphy
of central Europe. Other parts of Mader's
book are more accessible, summarizing for
example the literature on flowers visited
by CoUetes, or profiling the communities of
other soil-dwelling species that secondar-
ily use tunnels of abandoned C. daviesaniis
nests.

Mader's book actually comes in two
volumes, only the first of which is pub-
lished at this point. The second volume is
scheduled for publication next year, but a

Volume 8, Number 2, 1999

269

table of contents is already available.
While both volumes are organized around
aspects of the nesting biology of C. dav-
ieaniis and other Colletes species, the first
volume seems to focus more on biological
aspects (e.g., presenting information on
Colletes parasites, flower visitation, etc.),
while the second volume seems more spe-
cialized and appears to concentrate more
on geological aspects.

Who would benefit from these vol-
umes? Clearly the readership is rather lim-
ited, yet it certainly should not be missing
from libraries specializing on hymenop-
teran literature or general natural history.
But apart from Colletes afficionados inter-
ested in knowing everything about Colle-
tes, including the geological trivia of nest

substrate choice of a single species in cen-
tral Europe, I do not think that these vol-
umes will find much use among bee bi-
ologists. This is not a reflection of the
quality of the work, but more a reflection
of the currently perceived value of natural
history accounts. So I imagine that the true
value of these volumes will not be appar-
ent until C. dai'iesaniis, though abundant
at this point, may become extinct some
time in the future. The same is certainly
true for many other species for which time
did not permit a comprehensive study of
all life-history aspects, as for example the
nesting habits of the passenger pigeon.

Ulrich G. Mueller, Integrative Biology,
Patterson Labs, The University of Texas at
Austin, Austin, TX, 78712, USA.

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