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Journal of the
New York Entomological Society
VOLUME XC MARCH 1982 NO. 1
CONTENTS
Notes on gall inhabitants of Asphondylia helianthiglobulus Osten Sacken (Dip-
teral Cecidomyiidae) in Western Pennsylvania John D. Plakidas 2-4
Description of the male genitalia of Holcostethus hirtus (Van Duzee) with a revised
key to North American species (Hemiptera: Pentatomidae) F. J. D. McDonald 5-7
Sternal glands in three species of male social wasps of the genus Mischocyttarus
(Hymenoptera: Vespidae) David C. Post and Robert L. Jeanne 8-15
The temporal distribution of Chironomus decorus (chironomidae) in northern New
Jersey, 1979 G. L. Utberg and D. J. Sutherland 16-25
Pit construction by antiion larvae: influences of soil illumination and soil tempera-
ture Bradley G. Klein 26-30
Susceptibility of Kentucky bluegrass cultivars and selections to infestations of and
injury by the bluegrass billbug (Coleoptera: Curculionidae)
Sami Ahmad and C. Reed Funk 31-34
Notes on the natural history of Morpho granadensis polybaptus Butler (Lepidop-
tera; Nymphalidae: Morphinae), and its relation to that of Morpho peleides limpida
Butler Allen M. Young 35-54
Vladimir Nabokov 1899-1977: a note on a late entomologist Michael Juliar 55-57
Book Reviews 58-61
NEW YORK ENTOMOLOGICAL SOCIETY
XC(1), 1982, pp. 2-4
NOTE ON GALL INHABITANTS OF ASPHONDYLIA
HELIANTHIGLOBULUS OSTEN SACKEN
(DIPTERA: CECIDOMYIIDAE) IN
WESTERN PENNSYLVANIA
John D. Plakidas
Abstract. — The insects associated with the gall of Asphondylia helian-
thiglobulus Osten Sacken (Diptera: Cecidomyiidae) were reared during the
summer and winter of 1979-1980. Only those galls formed on the flower disc
of Rudbeckia laciniata L. (Compositae) were investigated. From the gall
cells of A. helianthiglobulus three hymenopterous species were reared: Ri-
leya americana Girault (Eurytomidae), Torymus advenus O.S., and Tory-
mus sp. (Torymidae). Dipterous larvae associated with the gall were rep-
resented by the families Agromyzidae, Cecidomyiidae, Chloropidae, and
Drosophilidae.
Introduction
The gall midge, Asphondylia helianthiglobulus Osten Sacken (Diptera:
Cecidomyiidae) (for synonymy see Rogers et al. 1979) forms an apple like
gall on the flower disc of the coneflower, Rudbeckia laciniata L. (Compos-
itae). The gall is composed of individual cells surrounded by plant tissue.
One host larva develops per cell. Externally there is no evidence of seed
formation, however petals and sepals do develop on the gall.
From field studies it was observed that female midges oviposited into
immature flower discs as early as July 8, with gall formation being visible
by July 15. Adult emergence began the third week in August.
Materials and Methods
In 1979 gall collections began the first week of August and continued on
a weekly basis through the last week of September. In addition, gall collec-
tions were made once monthly from October 1979 through January 1980 to
determine which species utilized the gall as an overwintering reservoir. Two
locations for gall collections were used: stream beds in the Linesville, Penn-
sylvania area, Crawford County; and McCandless Township, Allegheny
County, Pittsburgh, Pennsylvania.
Galls sampled during August and September 1979 were dissected in the
lab. Larvae and pupae recovered from the gall cells of A. helianthiglobulus
were placed on moist paper towels in plastic petri dishes. From both summer
and winter collections dipterous larvae and puparia recovered from the plant
VOLUME XC, NUMBER 1
3
tissue adjacent to the gall cells were reared in the same manner described
above.
Reared adult specimens were sent to the Insect Identification and Bene-
ficial Insects Introduction Institute, Beltsville, Maryland, for identification.
Results
The following is a list of species reared from the gall cells of Asphondylia
helianthiglobulus . None of these species including A. helianthiglobulus
were found to overwinter in the gall.
Eurytomidae (Hymenoptera).
1. Rileya americana Girault. Adults were reared from August 23
through September 11.
Torymidae (Hymenoptera).
2. Torymus advenus (O.S.). Adults emerged from August 21 through
October 15.
3. Torymus sp. Adult emergence was recorded from August 20 through
September 24.
Dipterans associated with the gall were all recovered from the plant tissue
adjacent to the gall cells. Those adults reared are presented below.
Agromyzidae.
1. Melanagromyza matricarioides Spencer. Adult emergence was re-
corded from August 26 through October 3.
Cecidomyiidae.
2. Neolasioptera rudbeckiae (Felt). Adults were reared from both sum-
mer and winter collections. Amblyaspis sp. (Hymenoptera: Platy-
gastridae), a primary endoparasite, was reared from larvae of N.
rudbeckiae.
3. Mycodiplosis sp. A single adult was reared February 15, 1980 from
McCandless.
Chloropidae.
4. Chaetochlorops inquilinus (Coq.). The puparia of this fly are found
overwintering in the gall.
Drosophilidae.
5. Drosophila quinaria Loew. A single adult was reared October 2 from
McCandless.
Unless otherwise noted, all of the species listed above were present in
both the Linesville and McCandless collections.
Acknowledgments
My grateful thanks to the following for identifications: Drs. C. W. Sa-
brosky — Chloropidae, E. E. Grissell — Eurytomidae and Torymidae, G.
4
NEW YORK ENTOMOLOGICAL SOCIETY
Steyskal — Agromyzidae, P. M. Marsh — Platygastridae, R. J. Gagne — Ce-
cidomyiidae, and W. N. Mathis — Drosophilidae.
Literature Cited
Rogers, C. E., T. E. Thompson and R. J. Gagne. 1979. Cecidomyiidae of Helianthus: tax-
onomy, hosts and distribution. Ann. Entomol. Soc. Amer. 72( 1): 109-1 13.
8506 Forest Avenue, Pittsburgh, Pennsylvania 15237.
Received for publication February 12, 1981.
NEW YORK ENTOMOLOGICAL SOCIETY
XC(1), 1982, pp. 5-7
DESCRIPTION OF THE MALE GENITALIA OF
HOLCOSTETHUS HIRTUS (VAN DUZEE)
WITH A REVISED KEY TO NORTH AMERICAN
SPECIES (HEMIPTERA: PENTATOMIDAE)
F. J. D. McDonald
Abstract. — A description of the male genitalia of Holcostethus hirtus is
given and an emended key to the species of the genus is provided.
At the time McDonald (1974) revised the genus Holcostethus in North
America, the male of H. hirtus was unknown. This paper provides a de-
scription of the male genitalia of H. hirtus and an emended key. Specimens
of this species that have lost their long, grey, dorsal setae do not run to the
correct name in the original key.
Holcostethus hirtus (Van Duzee, 1937)
Ventral border of pygophore (Fig. 1) sinuous with very shallow notches.
A pair of small triangular flap-like pseudoclaspers (Fig. 3), one on each side,
within dorsal surface of pygophore. Proctiger box-like (Fig. 2) with vertical
sides, distal margin produced into a distinct lip. Claspers L-shaped (Fig. 4)
with a basal thumb-like tubercle; inner surface of apex finely serrate; a
number of stout setae on outer margin. Theca small, cylindrical with a pair
of small tubercles one on each side at base. Conjunctival appendages mem-
braneous (Fig. 5), bilobed, bluntly rounded, fused basally onto a thecal
shield. Median penial lobes (Fig. 6) plate-like, apically acute, centrally fused
together by a stout cross bar. Ejaculatory duct sinuous, projecting beyond
median penial lobes.
Key to the Species of Holcostethus in N. America
1. Ventral surface of abdomen dark chocolate brown, sometimes mot-
tled, margins may be outlined in yellow 2
- Ventral surface of either yellow, buff or reddish brown with or with-
out black markings 5
2. Anterolateral margins of pronotum straight or only very slightly con-
cave; scutellum with a distinct yellow tip; male theca lacking apical
tubercles piceus (Dallas)
- Anterolateral margins of pronotum distinctly convex; scutellum con-
colorous or at most with a very faint white tip; male theca with
apical tubercles
3
6
NEW YORK ENTOMOLOGICAL SOCIETY
Eig. 1-6. Holcostethus hirtus. 1. pygophore ventral border; 2. pygophore, dorsal; 3. left
pseudoclasper; 4. left clasper, ental; 5. aedeagus, lateral, co-conjunctival appendage, M —
median penial lobe; 6. median penial lobes, E — ejaculatory duct.
Eig. 7-9. Pygophore, ventral border; 7. H. tristis', 8. H. abbreviatus\ 9. H. limbolarius .
3. Connexiva fuscous with continuous narrow yellow border; long grey
setae usually present on dorsum especially pronotum; male ventral
border of pygophore as in Fig. 1 hirtus (Van Duzee)
- Connexiva checked black and yellow; dorsum without long grey
setae; male pygophore with a distinct V-shaped notch centrally (Fig.
7) 4
4. Small species, not more than 7.5 mm long (apex of head to tip of
membrane) and 4.00 mm wide (between lateral angles of pronotum);
VOLUME XC, NUMBER 1
7
male ventral border of pygophore with a small protuberance below
median notch ruckesi McDonald
- Larger species, over 7.5 mm long and 4.00 mm wide; male ventral
border of pygophore without protuberance tristis (Van Duzee)
5. Reddish brown species with distinctive zig-zag black markings on
abdominal sterna; jugae not meeting in front of tylus; restricted to
E. States fulvipes (Ruckes)
- Brown colored species without distinctive markings on abdomen 6
6. Dark punctation on pronotal dorsum continuing irregularly to edge
of anterolateral margins; latter sub marginally impressed; connexiva
with distinct checked pattern of yellow and black; male ventral bor-
der of pygophore as in Fig. 8; dorsal border bearing a large pair of
spines abbreviatus Uhler
- Anterolateral margins of pronotum calloused and without dark punc-
tures, lacking submarginal impressions; connexival margin yellow;
male ventral border of pygophore as in Fig. 9; dorsal border without
spines limbolarius (Stal)
Acknowledgments
I should like to thank Mr. W. R. Dolling of the British Museum for the
loan of material. I am grateful to Professor L. H. Rolston, Entomology
Department, Louisiana Sate University, for pointing out the problems he
encountered with my original key.
Literature Cited
McDonald, F. J. D. 1974. Revision of the Genus Holcostethus in North America (Hemip-
teraiPentatomidae). J. N.Y. Entomol. Soc. 82(4):245-258.
Department of Plant Pathology and Agricultural Entomology, University
of Sydney, N.S.W. Australia.
Received for publication April 3, 1981.
NEW YORK ENTOMOLOGICAL SOCIETY
XC(1), 1982, pp. 8-15
STERNAL GLANDS IN THREE SPECIES OF MALE
SOCIAL WASPS OF THE GENUS MISCHOCYTTARUS
(HYMENOPTERA: VESPIDAE)
David C. Post and Robert L. Jeanne
Abstract. — Males of Mischocyttarus flavitarsis and M. drewseni possess
a large mass of ducted exocrine gland cells on each of the fifth, sixth, and
seventh (terminal) gastral sternites. The epidermal cells in the region of
these glands are elongated, suggesting that they also have a secretory func-
tion. Males of a third species, M. mexicanus, have relatively few ducted
gland cells on the fifth and sixth sternites and none on the seventh sternite.
In M. flavitarsis (but not in M. drewseni or M. mexicanus), the gland cell
ducts of sternites five and six open into a dense brush of long hairs.
Mischocyttarus, even though it is strictly New World in distribution, is
the largest genus of social wasps (Richards 1978). Recent studies of four
species reveal a diversity of male mate-locating behavior within the genus.
Males of Mischocyttarus labiatus (F.) in Colombia, M. mexicanus (de Saus-
sure) in Florida, and M. drewseni de Saussure in Brazil patrol routes in
areas where females converge to feed or to nest (Litte 1981; personal
communication; Jeanne and Castellon Bermudez 1980). The males of M.
flavitarsis (de Saussure) in Arizona pursue two different strategies, depend-
ing on the season: during the summer nesting season they patrol routes in
female foraging areas, but in the fall, during dissolution of the colonies prior
to hibernation, they defend small territories near hibernation sites (Litte
1979). Males of M. labiatus and M. flavitarsis drag and rub the gaster on
perch sites, as though scent-marking (Litte 1981, 1979), while M. mex-
icanus and M. drewseni evidently lack this behavior (Litte, personal com-
munication; Jeanne and Castellon Bermudez 1980). Landolt and Akre (1979)
reported that a M. flavitarsis male from Washington State possessed large
glandular masses on the sixth and seventh (terminal) gastral sternites. The
glands are probably the source of a secretion which the male applies to his
perch (Litte 1979). However, the roles of this behavior and of any phero-
mone that may be deposited remain to be determined.
These behavioral observations suggest differences in the occurrence of
sternal glands among these species. The purpose of this paper is to deter-
mine the presence of these glands and to describe their morphology in males
of three of the species: M. flavitarsis, M. drewseni, and M. mexicanus. This
is the first study of the histology of exocrine glands in male social wasps.
VOLUME XC, NUMBER 1
9
Fig. 1. Location of the class 3 gland cells on the fifth, sixth, and seventh gastral sternites
of the male of M. drewseni.
Methods
Adult males of each species were collected from the localities of the
behavioral studies cited above, as follows. M. flavitarsis navajo Bequaert:
near the Southwestern Research Station, Portal, Arizona (fall of 1959 and
1978); M. mexicanus (de Saussure): Archbold Biological Station, Lake Plac-
id, Florida (early spring of 1980); M. drewseni de Saussure: Fazenda Ta-
perinha, Santarem, Para, Brazil (November, 1978). M. /. flavitarsis (de
Saussure) from Turlock, California, was examined with SEM for comparison
with M. /. navajo.
Specimens were fixed in Kahle’s solution, embedded in Spurr Low- Vis-
cosity embedding media (Poly sciences), and sectioned 2 p. thick for exam-
ination with a light microscope (Post and Jeanne 1980). Specimens were
prepared for scanning electron microscopy as described by Post and Jeanne
(1980) and examined with a JELCO JSM-U3 scanning electron microscope.
The glands of each species differ from one another only in relative size.
For this reason we illustrate the fifth (antepenultimate), sixth (penultimate),
and seventh (terminal) gastral sternites of M. drewseni and discuss the other
species in relation to it.
10
NEW YORK ENTOMOLOGICAL SOCIETY
Figs. 2-4. Longitudinal section (2 /jl thick) through the glandular area of the fifth, sixth, and
seventh gastral sternites of M. drewseni. 2. Fifth sternite. 3. Sixth sternite. 4. Seventh sternite.
Anterior to left. AR = anterior ridge; AT = adipose tissue; E = epidermal cells; GC = glan-
dular cells; Scale = 0.25 mm.
Results
Internal Anatomy
A broad, dense mat of individually ducted gland cells (“class 3 cells” of
Noirot and Quennedey 1974) underlies the anterior margin of each of sternites
5 and 6 in M. drewseni (Figs. 1-4). The duct of each cell passes through the
integument to open onto the surface of the sternite (Fig. 5). The gland cell
mass on the fifth sternite measures 1.9 mm wide by 0.6 mm front to rear
VOLUME XC, NUMBER 1
11
Eig. 5. Longitudinal section (2 /jl thick) through the glandular area of the sixth gastral sternite
of M. flavitarsis navajo. C = cuticle; CD = glandular cell ducts; E = epidermal cells; GC =
glandular cells; H = hairs of the sternal brush. Scale = 0.05 mm.
and is a single cell layer thick (ca. 0.025 mm) (Fig. 2). In contrast the mass
on the sixth sternite is 1-3 cells thick (Fig. 3) and greater in extent (2.0 mm
by 0.7 mm). The seventh sternite has only scattered class 3 cells underlying
its posterior half (Figs. 1, 4). In one specimen there were 115 cells.
The location of class 3 cells on each sternite is the same in M. flavitarsis
navajo as in M. drewseni, but the cells are more numerous. The masses
of cells extend the full width of sternites 5 and 6 (2.8 and 2.6 mm, respec-
tively) and are somewhat bilobed, measuring, respectively 0.8 mm and 0.9
mm along the midline and 1.0 mm and 1.1 mm on the sides. Both masses
of cells are also thicker than in M. drewseni: that on the fifth is cells
thick, while that on the sixth is 2-8 cells thick (both glands are thinner at
the edges). The class 3 cells on the seventh sternite form a dense mat, one
cell thick, comprising about 300 cells in one specimen.
In contrast, M. mexicanus has only a few scattered class 3 cells on the
anterior margins of sternites 5 and 6. In one specimen there were 73 and 48
cells on the two sternites, respectively. The seventh sternite lacks class 3
cells.
In all three species the epidermal layer is thickened in the region of the
class 3 cells, suggesting that these cells are actively secreting ('‘class 1
cells” of Noirot and Quennedey 1974) (Figs. 2^). The layer is much thicker
12
NEW YORK ENTOMOLOGICAL SOCIETY
Fig. 6. SEM photograph of the fifth, sixth, and seventh (terminal) gastral sternites of M.
f. navajo. Sternites are pulled apart to expose their anterior margins. Scale = 2.0 mm.
in M. drewseni (Y = 27.0 fi; SD = 9.5 and x = 33.9 /x; SD = 16.1 on the
fifth and sixth sternites, respectively; 3 specimens) than in M. f. navajo
{x = 20.6 jul; SD = 4.2 and Y = 21.9 jjl; SD = 6.4 on the fifth and sixth
sternites, respectively; 3 specimens). The epidermal cells of M. mexicanus
are not highly modified on these two sternites (6.7 jjl thick; 1 specimen). On
the seventh sternite the elongation is greater in M. /. navajo (Y = 24.9 /x;
SD = 3.4; 3 specimens) than in M. mexicanus (15.5 /x; 1 specimen) and M.
drewseni (Y = 11.1 ^t; SD = 0.3; 2 specimens).
External Morphology
The anterior regions of both the fifth and sixth gastral sternites of M.
flavitarsis navajo (6 specimens) and M. /. flavitarsis (2 specimens) males
are densely covered with long, brush-like hairs, while the seventh sternite
lacks these (Fig. 6). All three sternites of M. drewseni (3 specimens) and
M. mexicanus (3 specimens) lack these hairs, and resemble the seventh
sternite of M. flavitarsis.
Glandular duct openings are located between the long hairs of M. f. na-
vajo (Figs. 5, 7) and between the body hairs of M. drewseni (Fig. 8) and M.
mexicanus.
Discussion
Landolt and Akre (1979) report sternal bushes and ducted, globular gland
cells on the sixth and seventh gastral (seventh and eighth abdominal) ster-
Figs. 7-8. SEM photograph of the glandular cell duct openings (seen as pores) near the
base of the fifth gastral sternite. 7. M. /. navajo. 8. M. drewseni. Scale = 0.05 mm.
VOLUME XC, NUMBER 1
13
14
NEW YORK ENTOMOLOGICAL SOCIETY
nites of a male of M. flavitarsis from Washington State (probably M. f.
idahoensis Bequaert). Our specimens of M.f. navajo from Arizona and M.
f. flavitarsis from California clearly have these structures on gastral sternites
5 and 6. Either Landolt and Akre were in error or the subspecific difference
is real. We were unable to obtain specimens of M. flavitarsis from Wash-
ington State, so we cannot resolve the question.
Since the number of gland cells is fixed in the adult stage, the observed
differences in the number of class 3 gland cells are species differences and
cannot be attributed to developmental differences. Of the three species we
examined, M. flavitarsis navajo has the greatest number of class 3 gland
cells on all three sternites and is the only species with sternal brushes. It
also appears to be the only one of the three whose males rub these sternites
on the substrate (Litte 1979; personal communication; Jeanne and Castellon
Bermudez 1980). A sternal brush appears to be a common structure in
species of aculeate wasps known to rub a secretion onto a substrate, for
example males of Eucerceris spp. and Philanthus spp. (Alcock 1975;
Gwynne 1978; O’Neill 1979), and females of Polistes spp. (Hermann and
Dirks 1974; Turillazzi 1979; Post and Jeanne 1980) and Mischocyttarus spp.
(Jeanne 1970). These facts suggest the possibility that the 5th and 6th sternal
glands serve different functions in M. drewseni and M. mexicanus than in
M. flavitarsis. The location of the class 3 gland cells on the posterior half
of sternite 7 in M. flavitarsis and M. drewseni suggests that their role is
different from that of the cells on the preceding two segments.
The differences in height of the thickened epidermal cells may reflect only
developmental differences. Since we had no information about age or activ-
ity of the specimens we sampled, all we can say is that in all three species
these cells appear to be glandular in function.
This study reveals that species in the genus Mischocyttarus are not uni-
form with respect to the occurrence and size of male sternal glands. It is
likely that further diversity will be found among the remaining 199 species.
Our results, combined with the behavioral studies already completed,
strongly suggest interspecific differences in the importance of chemical sig-
nals in the behavior of males, either on the natal nest, in a reproductive
context, or both.
Acknowledgments
Marcia Litte kindly collected and fixed Mischocyttarus mexicanus and
M. flavitarsis navajo specimens. Roy Snelling provided specimens of M.f.
flavitarsis and M. f. navajo. Martin Garment provided technical assistance
with the SEM. We thank Donas Erica and Violeta Hagmann for their hos-
pitality during our stay in Brazil. Research supported in part by the College
of Agriculture and Life Sciences and The Graduate School, University of
VOLUME XC, NUMBER 1
15
Wisconsin, Madison, and by National Science Foundation grant BNS 77-
04081.
Literature Cited
Alcock, J. 1975. Male mating strategies of some philanthine wasps (Hymenoptera: Sphecidae).
J. Kansas Entomol. Soc. 48:532-545.
Gwynne, D. T. 1978. Male territoriality in the bumblebee wolf, Philanthus bicinctus (Mickel)
(Hymenoptera, Sphecidae): observations on the behavior of individual males. Z. Tier-
psychol. 47:89-103.
Hermann, H. R. and T. F. Dirks. 1974. Sternal glands in polistine wasps: morphology and
associated behavior. J. Georgia Entomol. Soc. 9:1-8.
Jeanne, R. L. 1970. Chemical defense of brood by a social wasp. Science 168:1465-1466.
and E. G. Castellon Bermudez. 1980. Reproductive behavior of a male Neotropical
social wasp, Mischocyttarus drewseni (Hymenoptera: Vespidae). J. Kansas Entomol.
Soc. 53:271-276.
Landolt, P. J. and R. D. Akre. 1979. Occurrence and location of exocrine glands in some
social Vespidae (Hymenoptera). Ann. Entomol. Soc. Am. 72:141-148.
Litte, M. 1981. Social biology of the polistine wasp Mischocyttarus labiatus: survival in
a Colombian rain forest. Smithsonian Contrib. Zool. 327:1-27.
. 1979. Mischocyttarus flavitarsis in Arizona: social and nesting biology of a polistine
wasp. Z. Tierpsychol. 50:282-312.
Noirot, C. and A. Quennedey. 1974. Fine structure of insect epidermal glands. Ann. Rev.
Entomol. 19:61-80.
O’Neill, K. M. 1979. Territorial behavior in males of Philanthus psyche (Hymenoptera,
Sphecidae). Psyche 86:19^3.
Post, D. C. and R. L. Jeanne. 1980. Morphology of the sternal glands of Polistes fuscatus and
P. canadensis (Hymenoptera: Vespidae). Psyche 87:49-58.
Richards, O. W. 1978. The social wasps of the Americas excluding the Vespinae. Brit. Mus.
(Nat. Hist.), London, 580 pp.
Turillazzi, S. 1979. Tegumental glands in the abdomen of some European Polistes (Hyme-
noptera Vespidae). Monitore Zool. Ital. (N.S.) 13:67-70.
Department of Entomology, University of Wisconsin-Madison, Madison,
Wisconsin 53706.
Received for publication April 22, 1981.
NEW YORK ENTOMOLOGICAL SOCIETY
XC(1), 1982, pp. 16-25
THE TEMPORAL DISTRIBUTION OF CHIRONOMUS
DECOR US (CHIRONOMIDAE) IN NORTHERN
NEW JERSEY, 1979^
G. L. Utberg^ and D. J. Sutherland
Abstract. — The temporal distribution of Chironomus decorus, inhabiting
a shallow, brackish pond, was determined over the course of one season.
Data on larval abundance and adult flight activity were collected at regular
intervals to study population trends. Initial population levels, following thaw
of the winter ice cover, were relatively low. As temperatures rose through-
out the spring and early summer, midge abundance gradually increased.
Larval density peaked in mid-summer, averaging 700 third and fourth instar
larvae per 0.02 m^ of substrate. Population numbers decreased rapidly in
late summer, prior to the onset of colder temperatures and winter weather
conditions. Fluctuations in water level, salinity, and dissolved oxygen
levels, are discussed in terms of their limiting affect on the aquatic environ-
ment and its inhabitants. Temperature appears to fill the primary role in
regulating the population of C. decorus in the study area. Peaks in adult
emergence and larval abundance suggest 5-6 generations were completed
during 1979.
Introduction
Residents living adjacent to the Hackensack Meadowlands of New Jersey
are plagued each spring and summer by tremendous swarms of the midge,
Chironomus decorus Johannsen. The aquatic larvae develop in an area that
was once salt marsh meadow, but is now a brackish, muck-bottomed pond.
The habitat is subjected to daily tidal fluctuations, to a depth of one foot
or less at ebb tide, and up to three feet at flood tide. Access roads and gas
pipelines prevent a significant amount of water exchange, and organic ma-
terials are steadily introduced through garbage dump runoff and disposal
plant effluents. The implementation of effective control measures for this
chironomid is hampered by a relative lack of knowledge concerning the
biology and habits of the species in this environment. Field investigations
^ Paper of the Journal Series, New Jersey Agricultural Experiment Station, Rutgers Uni-
versity, the State University of New Jersey, New Brunswick, New Jersey, 08903. This research
was performed under NJAES Project 40401.
^ Pesticide Technology Dept., Penick Corp., Orange, New Jersey 07050.
VOLUME XC, NUMBER 1
17
were undertaken to examine the temporal distribution of C. decorus during
1979, as measured by larval abundance and adult flight activity.
Materials and Methods
A quantitative survey of the larval populations was initiated in March of
1979, immediately following thaw of the winter ice cover. Three aquatic
sampling stations were established along a transect running parallel to the
southern shore of the study area (Fig. 1), and were permanently tagged with
buoys. An airboat was used for transportation through the shallow water,
to avoid disturbance of the upper 5 cm of bottom substrate, where 95% of
the vertical distribution of chironomid larvae has been found to occur (Car-
ter 1976; Mundie 1957). An Ekman dredge was employed as a sampling
device to obtain 0.02 m^ sections of mud, to an approximate depth of 7 cm.
At each of the three stations, five samples were obtained, one from a central
position at the buoy, and one taken 4-5 m from the buoy at each of the four
cardinal directions. The sampling procedure consisted of first raising the
sample from the bottom and immediately releasing the contents of the
dredge into a bucket. The combined mud and water yield of each dredge
sample was approximately four liters. The mud and water were whipped to
a slurry, and a portion of the mixture funneled into a half-liter plastic con-
tainer. The aliquots were transported to the laboratory for subsequent ex-
amination.
Larval sampling was conducted from early March through November.
Collections were made on a semi-monthly basis in the spring and fall, and
at weekly intervals during the summer. Environmental factors were moni-
tored on each sampling date, including benthic mud and surface water tem-
peratures, dissolved oxygen content, pH, and salinity of the water. Tem-
perature measurements were taken with a mercury thermometer, and
dissolved oxygen was determined in the field with a Hach kit.^ A refractom-
eter was used to ascertain salinity levels in ppt, and pH was periodically
determined with commercially prepared indicator papers. Daily sampling
times were selected according to published tide tables, when water levels
accommodated boat usage.
The larvae were separated from the mud slurry in the laboratory by rinsing
each aliquot through a series of U.S. Standard Sieves, with screening rang-
ing from 20 to 50 mesh. Head capsule width measurements were taken to
determine the instar of the larvae (Ford 1959). Preliminary trial separations
indicated that the first and second instars were often lost in the rinse water.
Their recovery by flotation methods was inconsistent and hampered by
^ Model OX-10, Hach Chemical Co., Ames, Iowa.
^ pHydrion Midget, Ward’s, Rochester, New York.
18
NEW YORK ENTOMOLOGICAL SOCIETY
Fig. 1. Study site in the Hackensack Meadowlands, Bergen County, New Jersey, with
locations of larval sampling stations and of adult traps (shaded, land; clear, water).
small detritus. Therefore, only third and fourth instar larval counts were
used to assess population levels.
Light trap data gave an indication of seasonal adult emergence patterns.
One New Jersey light trap, permanently established 600 m from the shore,
was operated once weekly for a 24 hr period, from April to November. Two
CDC portable traps were positioned closer to the shore, at 425 and 275 m
distances. The portable traps were operated one evening a week, between
8:30 pm and 10:30 pm, from May to November. The time selected was based
VOLUME XC, NUMBER 1
19
Table 1. Head capsule width measurements for the four larval instars of Chironomus de-
corus, collected in the Hackensack Meadowlands, Bergen County, New Jersey.
Instar
Range
Head capsule width (mm)
Mean + C.I.
First
0.07-0.13
0.10 ± .0039
Second
0.14-0.20
0.17 ± .0029
Third
0.25-0.33
0.29 ± .0037
Fourth
0.40-0.61
0.50 ± .0108
Upon a study (Ping 1917) reporting that peak flight activity of C. decorus
occurs just after sunset. The weekly trap collections were synchronized
with larval sampling dates, and the numbers of male and female adults were
recorded.
Results
The range of head capsule width measurements for C. decorus was uni-
form for each instar (Table 1). As there was no overlapping of sizes, head
capsule measurements were confirmed as a reliable means of determining
the instar of field-collected larvae. The mean numbers of third and fourth
instar larvae collected per 0.02 m^ are presented on a weekly basis in Figure
2. The samples taken in early March contained only low levels of fourth
instar larvae, and no individuals were detected in April. Third and fourth
instar larvae suddenly appeared in May, and numbers increased as the sum-
mer progressed. A maximum density of 700 larvae per 0.02 m^ was recorded
in mid July. The population then declined in early August, and stabilized at
comparatively low levels for the duration of the fall season.
Mean larval abundance per sample was different at the three stations (Site
1 = 208.6 ± 56.99; Site 2 - 180.5 ± 47.80; Site 3 = 141.4 ± 38.16), while
the numbers of larvae collected from the five locations at any one station
were statistically the same (5% level of significance). Duncan’s multiple
comparison test revealed that adjacent stations (1 and 2, 2 and 3) were the
same, while larval numbers at stations 1 and 3 were significantly different.
This indicates that a density gradient existed, with highest concentrations
of larvae at station 1, and density decreasing toward station 3. The larval
density gradient was accompanied by changes in substrate composition.
Fine silt and ooze predominated at station 1 , and shifted to a coarser mixture
of muck and broken vegetation at station 3. Sublette (1957) and Whitsel et
al. (1963) have also reported that C. decorus prefers to inhabit finer sub-
strates.
Changes in the recorded environmental data reflected the seasonal time
span of the investigation (Figure 3). Water temperatures gradually increased
20
NEW YORK ENTOMOLOGICAL SOCIETY
Mar Apr Moy Jun Jul Aug Sep Oct Nov
I ' ' ' I ' — I r 1
WEEK
Eig. 2. Average number of third and fourth instar larvae of C. decorus, collected in the
Hackensack Meadowlands, Bergen County, New Jersey, 1979.
from 6.5°C in early March, when larval numbers were lowest, to 24°C in
May, when the larval population increased significantly. Throughout the
summer months, temperatures oscillated from week to week, but exhibited
an overall rising trend. A maximum of 33.5°C was attained in early August,
when larval numbers began declining. Temperatures remained above freez-
ing until late October. The dissolved oxygen content of the water was high-
est in early spring, when monitoring was initiated (12 ppm), and fell to
relatively low levels (3-5 ppm) for the summer and fall seasons. The pH of
the water was constant, between 6. 8-7.0, throughout the study. The salinity
of the water, during the spring precipitation period, was zero ppt. Salinity
increased with the onset of dryer, warmer conditions, attaining a maximum
of 13 ppt in August. Salinity readings averaged 9 ppt during the remainder
of the season.
VOLUME XC, NUMBER 1
21
Mar Apr May Jun Jul Aug Sep Oct Nov
1 — 1 n r — ^ 1 — ^ 1 1
WEEK
Fig. 3. Temperature, oxygen, and salinity levels recorded in the aquatic habitat of C.
decorus, Hackensack Meadowlands, Bergen County, New Jersey, 1979.
The warm summer temperatures were accompanied by a proliferation of
submergent vegetation, particularly the green algae Cladophora sp. and
Enteromorpha sp. A vegetative mat had formed over the water surface
by mid July. In late July the plant life rapidly diminished, and had nearly
disappeared by mid August.
The numbers of adults collected from the N.J. light trap and the two
portable traps are plotted on a logarithmic scale in Figure 4. The pattern of
adult flight activity closely resembles the larval population trends. Adults
were less active in the early spring and fall when lower temperatures pre-
vailed, with emergence occurring primarily during June and July.
The use of different trap types prohibited averaging of data from the three
trap locations, and limited statistical analysis. However, the overall flue-
22
NEW YORK ENTOMOLOGICAL SOCIETY
Mar Apr May Jun Jul Auq Sep Oct Nov
I — ^ — I — ^ I ■ 1 1 — I \ 1
WEEK
Eig. 4. Air temperatures and numbers of adult C. decoms collected in one NJ. light trap
and two portable CDC traps, Hackensack Meadowlands, Bergen County, New Jersey, 1979.
tuations in abundance appeared similar among the three sets of data. The
N.J. light trap and portable 2 exhibited the strongest correlation (Table 2),
while the centrally located trap, portable 1, was weakly correlated with the
other two traps. Males dominated the trap counts at portable 2, but the sex
ratio became more equal with increasing distance from the shore.
Discussion
The population trend of C. decoms typifies that of an insect inhabiting a
temperate region, where temperature is the dominant regulator of insect
VOLUME XC, NUMBER 1
23
Table 2. Adult Chironomus decorus collected during a 15 week period from three light
traps located in the Hackensack Meadowlands, Bergen County, New Jersey.
Light trap
Distance
from shore
(meters)
Total adults
Sex ratio
(M:F)
Correlation
coefficient (R)
with N.J. trap
New Jersey
600
8,324
1.8:1
Portable 1
425
1,542
4.4:1
0.48
Portable 2
275
10,339
26.0:1
0.85
activity and population growth (Corbet 1964). A resumption of larval growth
and development following thaw of the winter ice cover is considered
a facultative developmental response to warmer temperatures (Oliver 1971).
The disappearance of third and fourth instar larvae during April, and their
reappearance in May, suggest a generation of individuals developing in syn-
chrony. A more sensitive sampling technique might have detected the pres-
ence of first and second instar larvae during April. This generation was
probably a consequence of an early initial spring emergence of the over-
wintering population (Oliver 1971).
The gradual rise in spring temperatures was accompanied by increases in
the midge population. A sigmoid relationship between temperature and de-
velopment has been demonstrated in some chironomid species (Biever
1967). A faster developmental time and shorter life cycle, in the presence
of higher temperatures, appears likely for C. decorus.
The sudden decline in larval abundance, and accompanying drop in veg-
etative growth, occurred during a period in August when temperatures
reached their highest levels. The temperature fluctuations within the habitat
were probably more extreme on a daily basis than is indicated by the re-
corded data. Local wind and tidal effects reduced the water level at ebb tide
to a few centimeters or less, exposing a great deal of the benthic mud
surface. On a hot day, with ebb tide occurring at midday, the maximum
daily temperatures attained would have been greater than those recorded
during the high tide sampling situation. Temperature may have surpassed
the tolerance levels of C. decorus and the aquatic life (Brauner 1979), and
directly influenced survival.
Dense growths of algae and vegetation, such as were present in the study
site, have been attributed with causing mid summer declines in chironomid
populations (Bay and Anderson 1965). Plant respiration, and decomposition
of an organic aggregate derived from the death of a large algal bloom, can
reduce oxygen tension in the water (Cole 1975). Therefore, vegetation as
well as temperature could have produced an oxygen deficit, detrimental to
the larvae.
The variability in the number of adults collected by the three light traps
was partially due to the use of different types of trap, and the length of the
24
NEW YORK ENTOMOLOGICAL SOCIETY
sampling interval for each type. Undefined differences in light trap efficien-
cy, and local environmental factors, could have caused the weak correlation
observed between the centrally located trap and the other two traps. The
distance between the traps and the source would also be expected to affect
the number of adults trapped. The decreasing male sex ratio in traps located
further from the shore indicates that distance did influence light trap reli-
ability to some extent. The male swarming behavior, in which assemblages
form over stationary points (Downes 1969) may have limited dispersal of
males. A differential flight ability between the sexes, where the female is
the stronger flyer, would also result in fewer males reaching points further
from the source.
The overwintering status of the study population in New Jersey was not
clearly defined by the data collected. However, it was presumed that pop-
ulation growth of C. decorus was minimal after November, when larval
sampling and adult trapping were discontinued. Previous research has in-
dicated that populations of chironomids, located in temperate regions, gen-
erally overwinter as larvae in a state of suspended growth and development
(Oliver 1971). Laboratory reared larvae, of a species closely related to C.
decorus, ceased feeding and became inactive at temperatures of 50°F (10°C)
or lower (Biever 1967). Adults become lethargic at 5°C or lower (Ping 1917).
The number of generations completed by C. decorus can be estimated by
counting the number of apparent peaks in larval abundance and adult flight
activity. Highest levels were attained at points in mid May, June, July,
August, September, and late October. As the season progressed, and gen-
erations began overlapping, the peaks were less distinct and more difficult
to distinguish. Five to six generations may have been completed by C.
decorus in northern New Jersey, during the 1979 reproductive season. This
finding is similar to that of Ping (1917) for a source located in Ithaca, New
York.
Summary and Conclusions
Data obtained from larval sampling and trapping of adults indicated that
the highest levels of C. decorus were attained in midsummer, 1979. Abun-
dance during the spring and fall seasons was relatively low, apparently being
regulated by environmental factors. The initial rise in spring temperatures
resulted in a resumption of larval growth and development, and emergence
of overwintering individuals. Population accrual continued to accompany
rising temperatures throughout the summer, with the highest numbers oc-
curring in July. A sudden drop in larval abundance in late summer may have
been a result of extreme temperatures. In terms of nuisance potential, emer-
gent adults reached peak levels during a limited interval of the reproductive
season. Larval and adult data indicated that C. decorus can complete 5-6
generations in this habitat in a single year.
VOLUME XC, NUMBER 1
25
Acknowledgments
The authors are indebted to the Bergen County Mosquito Commission for
their assistance in the collection of data. We wish to thank Dr. Selwyn S.
Roback for confirming the chironomid species identification, and Drs. Fran-
cesco B. Trama and John Brauner for examining and identifying the aquatic
vegetation.
Literature Cited
Bay, E. C. and L. D. Anderson. 1965. Chironomid control by carps and goldfish. Mosq. News
25:310-316.
Biever, K. D. 1967. Biological studies on chironomids with emphasis on the temperature
relations of the developmental stages. Ph.D. Thesis, Univ. Calif., Riverside. (Libr.
Congr. Card No. Mic. 68-4955). 119 pp. University Microfilms, Ann Arbor, Mich. (Diss.
Abstr.).
Brauner, J. 1979. Personal communication. Rutgers Univ., New Jersey.
Carter, C. E. 1976. A population study of the Chironomidae (Diptera) of Lough Neagh. Oikos
27:346-354.
Cole, G. A. 1975. Textbook of Limnology. C. V. Mosby, Saint Louis, 283 pp.
Corbet, P. S. 1964. Temporal patterns of emergence in aquatic insects. Can. Entomol.
96:264-279.
Downes, J. A. 1969. The swarming and mating flight of Diptera. Ann. Rev. Entomol.
14:271-298.
Ford, J. B. 1959. A study of larval growth, the number of instars and sexual differentiation in
the Chironomidae (Diptera). Proc. Roy. Entomol. Soc. London (A) 34:151-160.
Mundie, J. H. 1957. The ecology of Chironomidae in storage reservoirs. Trans. Roy. Entomol.
Soc. London 109:149-232.
Oliver, D. R. 1971. Life history of the Chironomidae. Ann. Rev. Entomol. 16:211-230.
Ping, C. 1917. Observations on Chironomus decorus (Johannsen). Can. Entomol. 49:418-426.
Sublette, J. E. 1957. The ecology of the macroscopic bottom fauna in Lake Texoma (Denison
Reservoir), Oklahoma and Texas. Amer. Midi. Nat. 57:371-402.
Whitsel, R. H., C. A. Vickery, Jr., C. J. Rogers and C. D. Grant. 1963. Studies on the biology
and control of chironomid midges in the San Francisco Bay Region. Proc. and Papers
Calif. Mosq. Control Assoc. 31:83-94.
Cook College, Rutgers University, New Brunswick, New Jersey.
Received for publication May 14, 1981.
NEW YORK ENTOMOLOGICAL SOCIETY
XC(1), 1982, pp. 26-30
PIT CONSTRUCTION BY ANTLION LARVAE:
INFLUENCES OF SOIL ILLUMINATION AND
SOIL TEMPERATURE
Bradley G. Klein^
Abstract. — Pits constructed by antiion larvae are most frequently found
in areas which are darker and cooler than areas exposed to the sun. Soil
illumination and soil temperature, therefore, are two factors which may play
a role in determining the probability of pit construction. The influence of
these factors upon the probability of pit construction by M. immacidatus
larvae was examined. Soil illumination and soil temperature were indepen-
dently varied in the laboratory. Soil temperature was found to have a sig-
nificant effect upon pit construction, whereas soil illumination was found to
have no such significant effect. This supports the view that the construction
of pits in shaded areas is a response to temperature rather than to light.
Antiions of the genus Mynneleon are best known for the ability of their
larvae to construct inverted conical pits in dry, loose, fine grained soil.
These pits provide the antiion larva with an effective means of capturing
prey (Topoff 1977; Turner 1915). It has been frequently observed that antiion
larvae begin constructing pits at, or immediately following sundown (Haub
1942; Topoff 1977; Wheeler 1930; Youthed and Moran 1969b). It has also
been observed that antiion pits are usually found in shaded areas such as
under ledges of rock, under logs which do not touch the ground at all points
or under man made sources of shade (Green 1955; Haub 1942; Topoff 1977;
Turner 1915). Shaded areas differ from areas exposed to the sun in two
obvious respects; they are darker and they are cooler. These two charac-
teristics also apply to any given area at or after sundown, when it is com-
pared to the same area during the day. It therefore appears that soil illu-
mination and soil temperature are two physical properties of the
environment which may play a role in determining the probability of pit
construction by antiion larvae.
In a series of laboratory experiments using larvae of Mynneleon imma-
cnlatus (De Geer), Haub (1942) demonstrated that soil temperature does
indeed influence the probability of pit construction. A similar result was
demonstrated by Youthed and Moran (1969a) for larvae of M. obscurus
' Present address: West Laboratories, American Museum of Natural History, 79 St. and
Central Park West, New York, New York 10024.
VOLUME XC, NUMBER 1
27
(Rambur). Haub (1942) concluded that, '‘the building of pits after sundown
is due to a temperature factor rather than light” (p. 116). However, he did
not provide evidence to warrant the exclusion of light as a factor controlling
the probability of pit building.
Antiion larvae appear to be well equipped to respond to light. Wheeler
(1930) noted that the larvae possess seven eyes on each side of the head
which appear to be transitional between ocelli and compound eyes. These
eyes point in all directions except backwards. Behavioral responsiveness of
antiion larvae to light has been reported. For example. Turner (1915) has
demonstrated negative phototaxis in M. immaculatus and Comes (1909)
found a similar result for M. formicariiis.
Given the ability of antiion larvae to respond to light, the experiment
discussed below attempts to determine whether the level of soil illumination
influences the probability of pit construction by M. immaculatus larvae. In
short, the study tests the validity of Haub’s (1942) contention regarding the
lack of control of light over pit construction. The study also attempts to
replicate the findings of Haub (1942) and Youthed and Moran (1969b) re-
garding the influence of soil temperature upon probability of pit construc-
tion.
Materials and Methods
Forty-seven M. immaculatus larvae were used. All larvae were collected
from the bank of a creek near the Southwestern Research Station of the
American Museum of Natural History in southeastern Arizona. Only larvae
found at the bottom of pits were used in the study. This insured that all
antiion larvae used were capable of building pits under normal field condi-
tions.
The experiment proper was carried out in an International Radiant Co.
humidity chamber (Model HI 5) in which temperature and relative humidity
could be independently regulated and thermostatically controlled. The
chamber window was covered with cardboard to prevent ambient light from
entering. Within the chamber, larvae were placed in one of two white plastic
containers filled with 6 cm of dry silt. Each container was divided into two
11 X 16 cm compartments by a piece of cardboard and only one larva was
placed in a compartment. Therefore, 4 larvae could be tested in each ex-
perimental session. A 75 W light bulb was used to illuminate the chamber
in the high illuminance condition of the experiment and a 15 W bulb covered
by a red translucent piece of plastic was used in the low illuminance con-
dition. A Gossin Lumasix light meter was used to measure the illuminance
at the silt surfaces within the chamber. Petri dishes containing 0.5 cm of dry
silt were used to store the larvae between the time of collection and the
time of testing.
28
NEW YORK ENTOMOLOGICAL SOCIETY
The 4 larvae and the silt in which they were to be tested were collected
3 h prior to each session. This was the approximate amount of time needed
for the silt to reach its desired temperature after being placed in the envi-
ronmental chamber. During this 3 h period, the larvae were stored in a
shaded area of the laboratory. The experiment was run as a 2 x 2 fac-
torial design with 2 temperature levels (hot and cool) and 2 illumination
levels (light and dark). Therefore, 4 different experimental conditions were
used: (1) hot-light, (2) hot-dark, (3) cool-light and (4) cool-dark. Each ex-
perimental condition was run 3 times, making a total of 12 test sessions, and
each session lasted 3 h. All testing was done between 1200 and 2200 h
(MST). Each time a condition was repeated, it was run at one of 3 different
times during the day: early afternoon, late afternoon or evening. These
periods often overlapped by as much as 1 h. A total of 12 larvae were tested
under each experimental condition, except that due to the death of one
larva, only 1 1 were tested under the hot-light condition. The silt temperature
was checked at the beginning and the end of each test session. Silt temper-
atures in the hot conditions ranged from 38-47°C whereas temperatures in
the cool conditions ranged from 20-24°C. The measured silt surface illu-
minance was 1 ,076 lx in the light conditions and 86 lx in the dark conditions.
At the beginning of each test session, one larva was placed on the surface
of each silt compartment within the environmental chamber. Three hours
later, each compartment was checked for pit construction. A larva was
scored positive for pit construction if: (1) a pit existed in its compartment,
or (2) the larva was in the process of pit construction and had dug out more
than one concentric circle. Larvae which were not scored positive for pit
construction were placed in a box of silt and checked for pit construction
24 h later. All these larvae built pits and this ruled out physical damage
during the experiment as a reason for lack of pit construction.
Results
The mean percentage of larvae constructing pits, per session, was deter-
mined for each condition of the experiment. The greatest amount of pit
construction occurred in the cool-dark condition (58.3%), followed in de-
scending order by the cool-light condition (41.7%), the hot-light condition
(1 1.0%) and the hot-dark condition (8.3%). The mean percentage of larvae
constructing pits was 9.7% per session under the hot temperature level as
compared with 50.0% per session under the cool temperature level. Under
the light illumination level, the mean percentage of larvae constructing pits
was 26.3% per session as compared with 33.3% per session under the dark
illumination level. A two-way ANOVA in conjunction with the arcsine
transformation for percentage data revealed a significant difference between
the two temperature levels (F = 9.16, df = 1,8, P < 0.025). No significant
VOLUME XC, NUMBER 1
29
difference was found between the two illumination levels (F = 0.30, df
1,8, F > 0.05) nor was any significant interaction found between tempera-
ture and illumination (F = 0.48, df = 1,8, F > 0.05).
The overlap of the early afternoon, late afternoon and evening replicates
within each condition prevented the inclusion of time of day as a separate
factor in the design. However, since the early afternoon and evening time
periods never overlapped, differences in pit building activity during these
two time periods were examined. The mean percentage of larvae construct-
ing pits was 12.5% per session during the early afternoon time period (1230
to 1530 h) and 39.5% per session during the evening time period (1835 to
2200 h). A single classification ANOVA in conjunction with the arcsine
transformation for percentage data revealed a significant difference between
the two time periods (F = 6.34, df = 1,6, F < 0.05).
Discussion
The finding that the greatest amount of pit building took place under the
cool-dark condition is consistent with field observations that antiion larvae
frequently build pits in areas that are darker and cooler than nearby areas
exposed to the sun (Green 1955; Haub 1942; Turner 1915). The significant
difference in pit building behavior between the two temperature levels (hot
and cool) supports the finding of Haub (1942) that soil temperature can
influence the probability of pit construction in M. immaculatus larvae. It is
interesting to note that Haub found no pit construction above 28°C and
100% mortality above 38°C. This did not appear to be the case in the present
study since pit construction occurred at temperatures as high as 44°C and
only one out of 24 larvae died within a temperature range of 38^7°C. It
seems that larvae in the present study were better able to withstand and
function at high temperatures than the larvae used by Haub. Although both
studies employ M. immaculatus larvae, those used in the present study
were collected in Arizona whereas those used by Haub were collected in
Ohio. A comparison of summer soil temperatures measured in the field
suggests that this seemingly greater tolerance for high temperatures by Ar-
izona M. immaculatus larvae may represent a regional adaptation to higher
temperature extremes than those encountered by Ohio M. immaculatus
larvae.
The absence of a significant difference in pit building behavior between
the two illumination levels (light and dark) supports the view that the con-
struction of pits after sundown or in shaded areas is a response to temper-
ature rather than to light. In other words, frequent construction of pits in
cool, dark areas is probably a response to coolness as opposed to darkness.
The absence of an influence of illumination level upon pit building proba-
bility was most likely not due to an inability of the larvae to discriminate
30
NEW YORK ENTOMOLOGICAL SOCIETY
between the illumination levels used since Youthed and Moran (1969b)
found that antiion larvae (M. obscurus) can discriminate between illumi-
nation levels similar to those used in the present study.
The significant difference in pit building activity found between the early
afternoon and evening time periods seems to indicate endogenous temporal
variation in pit building since external conditions were the same during both
periods. This would be in agreement with findings of Youthed and Moran
(1969b) who demonstrated an endogenous circadian pit building rhythm in
M. obscurus larvae. Pit building in M. immaculatus larvae therefore appears
to be under the control of internal as well as external factors.
Acknowledgments
I would like to thank Howard Topoff for his thoughtful advice and Rita
Feldman Klein for her technical assistance. This research was supported by
NIMH training grant MH 15341 and the Biopsychology Program, Hunter
College of the City University of New York.
Literature Cited
Comes, S. 1909. Stereotropismo, geotropismo e termotropismo nella larva de Mynneleon
fonnicarius. L. Atti Accad. Gioen. Sc. Nat. Catania 86(4): 1-14.
Green, G. W. 1955. Temperature relations of antiion larvae {Neuroptera: Mynneleontidae).
Can. Ent. 87:441^59.
Haub, J. G. 1942. The pit-building activities of Ohio antiions. Ohio J. Sci. 42:113-116.
Topoff, H. 1977. The pit and the antiion. Nat. Hist. 86:64-71.
Turner, C. H. 1915. Notes on the behavior of the antiion with emphasis on the feeding activities
and letisimulation. Biol. Bull. 29(5): 277-307.
Wheeler, W. M. 1930. Demons of the Dust. W. Norton and Co., New York.
Youthed, G. J. and V. C. Moran. 1969a. Pit construction by myrmeleontid larvae. J. Insect
Physiol. 15:867-875.
. 1969b. The solar day activity rhythm of myrmeleontid larvae. J. Insect Physiol.
15:1103-1116.
Biopsychology Program, Hunter College, CUNY, 695 Park Ave., New
York, New York 10021.
Received for publication July 1, 1981.
NEW YORK ENTOMOLOGICAL SOCIETY
XC(1), 1982, pp. 31-34
SUSCEPTIBILITY OF KENTUCKY BLUEGRASS CULTIVARS AND
SELECTIONS TO INFESTATIONS OF AND INJURY BY THE
BLUEGRASS BILLBUG (COLEOPTERA: CURCULIONIDAE)
Sami Ahmad and C. Reed Funk
Abstract. — Nine cultivars and selections of the Kentucky bluegrass, Poa
pratensis L., were examined for susceptibility to a natural infestation of the
larvae of the bluegrass billbug, Sphenophorus parvulus Gyllenhal. There
was a significant correlation (r = 0.767, df = 1 , P < 0.05) between larval
infestations and percent damage to the bluegrasses. ‘Kenblue’ was among
the least damaged entries and had minimal infestation. This established cul-
tivar is characterized by narrow leaves, an upright (taller) posture, and high
susceptibility to the leaf spot and melting-out disease incited by Drechslera
poae (Baudys) Shoem. The other cultivars and selections had broader
leaves, lower growth, and showed less disease damage. Possibly, these char-
acteristics render these other bluegrasses more apparent than Kenblue and,
thereby, are preferred over Kenblue for oviposition by the adult billbugs.
Introduction
The bluegrass billbug, Sphenophorus parvulus Gyllenhal, often causes
damage to Kentucky bluegrass, Poa pratensis L., which is widely used as
a turf and pasture grass in the northcentral and northeastern U.S. The dam-
age caused by this pest is particularly severe in Nebraska (Kindler and
Kinbacher 1975). In 1978, a natural infestation of bluegrass billbugs in the
turfgrass plots at the N.J. Agric. Exp. Stn. Res. Ctr. at Adelphia, N.J.,
provided an opportunity to assess the relative susceptibility of 9 Kentucky
bluegrass cultivars and selections. The data provided a partial test for the
earlier observations of Kindler and Kinbacher (1975), and 1979 observations
of Lindgren et al. (1981) that suggested differential reaction of Kentucky
bluegrass cultivars to the bluegrass billbug.
Methods
The various cultivars and experimental selections of Kentucky bluegrass
were seeded in a replicated test at Adelphia, during September 1974. The
test site was on a loamy sand. Plot size was 1.22 x 1.83 m with a 0.15 m
unplanted border between each plot. The grass was mowed at 3.8 cm during
the fall of establishment but lowered to 1.9 cm thereafter. Weeds were
controlled, as needed, with DCPA, dicamba, and 2,4-D. Soil was limed to
maintain a pH of 6.0. The test was irrigated as needed for rapid establish-
32
NEW YORK ENTOMOLOGICAL SOCIETY
ment, and later, to prevent severe drought stress. Two nitrogen fertilizer
levels (1.7 and 3.2 kg/92.94 m“) were established on each plot during the
1974-78 period. The fertility level appeared to have little, if any, effect on
bluegrass damage.
During the 2nd week of July 1978, there was evidence of differential
damage to the various cultivar plots, and an initial observation showed that
the damage to the Kentucky bluegrass was primarily due to the larvae of
the bluegrass billbug. Populations of other coleopterous pests such as the
Japanese beetle, Popillia japonica Newman, northern masked chafer Cy-
clocephala borealis Arrow, European chafer Rhizotrogus majalis (Razou-
mowsky), and black turfgrass ataenius Ataenius spretulus (Raid.) that are
also serious pests of turfgrasses in northeastern region (Tashiro 1973), were
not present in the turfgrass plots at levels that would cause significant dam-
age. During the 3rd week of July, the plots were examined for billbug in-
festations and extent of injury.
The infestations were quantified by two separate determinations, each
with two replicates, and each sample covering 0.09 m^ of turfgrass in each
cultivar plot. The larvae were examined and counted by laying back the sod
with sharp knife and removing and examining the sod and soil to a depth of
4 cm. The damage to cultivars was visually assessed twice on the basis of
percent damage (0, 10, 20, ... , 100); the observations were repeated 2
more times, providing 6 determinations for each cultivar and selection. Each
observation of injury was carried out by a different observer to minimize
potential error inherent in this procedure if left entirely to the judgement of
a single individual.
The data on larval counts (4 determinations), and percent injury to cul-
tivars (6 determinations) were pooled to obtain mean values and for statis-
tical analysis to establish significant differences among the sample means.
The data also were analyzed for correlation between larval counts and dam-
age to bluegrass cultivars and selections.
Results and Discussion
Table 1 presents data on damage to Kentucky bluegrass cultivars and
selections and mean numbers (per 0.09 m^) of billbug larvae responsible for
the damage. The counts of billbug larvae ranged from a low of 1.3/0.09 m^
for ‘Kenblue’ to a high of 60.3/0.09 m^, for ‘Nugget.’ Low damage, ca. 25
to 27%, was recorded on Kenblue, ‘E 1757,’ and ‘E 353.’ The entries show-
ing high damage, over 80%, were Nugget, ‘Cheri,’ and ‘K3-182.’
A significant correlation coefficient (r = 0.767, df = 7, < 0.05) was
found between the larval counts and the damage to the bluegrasses. Since
one selection, K3-182, was exceptional in that despite low larval counts
(18.5/0.09 m^) it was most severely damaged (87%), it was of interest to
VOLUME XC, NUMBER 1
33
Table 1. Susceptibility of Kentucky bluegrass cultivars to damage by the bluegrass billbug
in turfgrass trials during September 1978 at Adelphia, NJ.
Bluegrass cultivar
or selection
.V percent turfgrass
damage'
.V number of billbug
larvae per 0.09 m--
Kenblue
27.4 a
1.3 a
E 1757
24.7 a
12.3 b
F 353
26.7 a
13.0 b
Princeton 104
39.1 a
14.8 b
Baron
67.8 b
51.0 c
Merion
76.0 b
57.8 c
Nugget
82.4 b
60.3 c
Cheri
83.5 b
45.3 c
K3-182
87.0 b
18.5 b
^ Assessment based on 3 separate observations, each with 2 replicates; N = 6. Means not
followed by the same letter are significantly different at 5% level by Duncan’s multiple range
test.
^ Based on 2 separate observations, each with 2 replicates; N = 4. Area of each sample
(replicate) = 0.09 m^. Means not followed by the same letter are significantly different at 5%
level by Duncan’s multiple range test.
evaluate the correlation coefficient by deleting this selection from data anal-
ysis. Analyzed in this way the correlation coefficient was even more highly
significant (r = 0.950, df = 6, P < 0.01). Although we have no explanation
to account for the extensive damage of K3-182, inasmuch as the data anal-
ysis with or without inclusion of this cultivar provided a significant corre-
lation coefficient, the observations of Kindler and Kinbacher (1975) and
Lindgren et al. (1981) were reaffirmed. Moreover, in the study of the dif-
ferential reaction of Kentucky bluegrass cultivars to the bluegrass billbug
by Kindler and Kinbacher (1975), the relationship was established with 15
cultivars; we had 9 entries in the present study, with ‘Merion’ and Nugget
being common to both studies. Lindgren et al. (1981) in their 1979 study had
included 38 cultivars, only 3 cultivars, Merion, Nugget, and ‘Baron’ being
common to the present study.
The cause(s) for the differential larval density and susceptibility of the
Kentucky bluegrass cultivars to the billbugs is a matter of speculation at
this time. Of all the entries in the test, Kenblue is an established cultivar
characterized as upright (taller) and narrow-leafed. Moreover, this variety
is highly susceptible to leaf spot and melting-out disease (caused by Drechs-
lera poae). The main purpose of the other entries, was to select Kentucky
bluegrass for resistance to this disease. This selection however generally
resulted in broader leaves and lower growth, relative to Kenblue. It may
well be that these qualities render these selections more apparent and, also,
as more suitable substrates for oviposition by the adult bluegrass billbugs.
34
NEW YORK ENTOMOLOGICAL SOCIETY
More research is clearly necessary to determine the nature of resistance and
the underlying mechanisms. Potentially this kind of information would be
valuable for developing improved billbug resistant cultivars.
Acknowledgments
Paper of the Journal Series, New Jersey Agricultural Experiment Station,
New Brunswick, New Jersey. This work was performed under NJAES Proj-
ect number 08128 and 15265 supported by Hatch Act Eunds.
Literature Cited
Kindler, S. D. and E. J. Kinbacher. 1975. Differential reaction of Kentucky bluegrass cultivars
to the bluegrass billbug, Sphenophonis parvulus Gyllenhal. Crop Science 15:873-874.
Lindgren, D. T., R. C. Shearman, A. H. Bruneau and D. M. Schaaf. 1981. Kentucky bluegrass
cultivar response to bluegrass billbug, Sphenophonis parvulus Gyllenhal. HortScience
16:339.
Tashiro, H. 1973. Bionomics and control of root feeding insect pests: grubs and billbugs. Bull.
Entomol. Soc. Amer. 19:92-94.
Department of Entomology and Economic Zoology, New Jersey Agri-
cultural Experiment Station, Cook College, Rutgers-The State University,
New Brunswick, New Jersey 08903, and Soils and Crops Department,
New Jersey Agricultural Experiment Station, Cook College, Rutgers-The
State University, New Brunswick, New Jersey 08903.
Received for publication August 26, 1981.
NEW YORK ENTOMOLOGICAL SOCIETY
XC(1), 1982, pp. 35-54
NOTES ON THE NATURAL HISTORY OE
MORPHO GRANADENSIS POLYBAPTUS BUTLER
(LEPIDOPTERA: NYMPHALIDAE: MORPHINAE),
AND ITS RELATION TO THAT OF
MORPHO PELEIDES LIMPIDA BUTLER
Allen M. Young
Abstract. — Various aspects of the natural history of the Neotropical but-
terfly Morpho granadensis polybaptus Butler (Lepidoptera: Nymphalidae:
Morphinae) in a zone of geographical overlap with M. peleides limpida
Butler in Costa Rica are reported for the first time. The work reported is
part of a long-range study on the comparative natural history of Central
American Morpho, represented in Costa Rica by at least five valid species.
Of particular interest is the comparison of M. granadensis with M. peleides
since both species are closely related forms in the South American achilles
“superspecies” complex. Morpho granadensis is exceedingly rare in mu-
seum collections and the type specimen is from Costa Rica. The species is
narrowly restricted in Costa Rica to a band of tropical rain forest within
about 100 to 600 meters elevation along the Caribbean watershed of the
Cordillera Central and adjacent highlands. The early stages are strikingly
similar to those of M. peleides, including various aspects of caterpillar be-
havior. Oviposition is single. Although the caterpillars are legume-feeders,
either the scope of food plant suitability is narrower or different between
this species and M. peleides, since the former cannot survive on at least
one common food plant {Mucuna mens) of the latter. Morpho peleides has
a much wider geographical range that M. granadensis, and probably utilizes
a greater range of food plants. The species is also sympatric with M. ama-
thonte Deyrolle; baiting studies indicate that M. granadensis is about half
as abundant as the other two species. A model for further comparative study
of the two species is presented.
Introduction
Some of the seminal papers on the life cycles and natural history of var-
ious species of Morpho butterflies (Lepidoptera: Nymphalidae: Morphinae)
in Central America have appeared recently on the pages of this journal (e.g.,
Young 1972a; Young and Muyshondt 1972a) and elsewhere (Young 1971a,
b, 1972b, 1973, 1974, 1975a, b, 1978b, 1979, 1980a; Young and Muyshondt
1972b, 1973; Young and Thomason 1974). As part of a long-range goal of
studying the biology of all Central American species of this widespread
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NEW YORK ENTOMOLOGICAL SOCIETY
Neotropical genus, field studies were conducted on Morpho granadensis
polybaptus Butler in northeastern Costa Rica, in a region where this species
and another one of the achilles “superspecies,” Morpho peleides limpida
Butler, are sympatric. Of particular interest is the study of M. granadensis
polybaptus in Costa Rica, the site of the type specimen for this species
(Seitz 1924). This paper reports the first description of the early stages and
larval food plant association for this species, and compares these attributes
and others with previously reported data on M. peleides from Costa Rica
(Young and Muyshondt 1973; Young 1972a, b, 1978b; and other papers).
Because adults of both species are very similar (Fig. 1), and one species
{peleides) is far more abundant than the other in zones of geographical
overlap, studies on the comparative natural history of these forms is of
particular interest, in the context of recent discussions of ecological mech-
anisms of environmental exploitation among closely related sympatric
species (e.g., see MacArthur 1972).
The major findings of this paper are the following: (1) the early stages of
M. granadensis are strikingly similar morphologically and behaviorally to
those of M. peleides, (2) one larval food plant of M. granadensis, the woody
vine Machaerium seemannii (Leguminosae) is also one of several legumi-
nous larval food plants of M. peleides in this region (Young and Muyshondt
1973) but caterpillars of M. granadensis die in early instars when reared on
several of these other plant species, suggesting a narrower range of food
plants in M. granadensis, (3) the egg-to-adult developmental times are very
similar in both species, (4) overall average egg production is probably much
lower in M. granadensis as captive females are less prone to oviposit and
lay fewer eggs when they do, and (5) M. granadensis has a relatively re-
stricted distribution within a narrow band of Caribbean watershed tropical
rain forest within about 100 to 600 meters elevation, while M. peleides is
far more catholic in its distribution throughout the whole country. Further-
more, M. granadensis, although very similar in habitat association and nat-
ural history to M. peleides, is probably not as abundant locally as the latter.
Along with M. amathonte Deyrolle, both species readily come to bait (rotten
bananas) although sex ratios are invariably skewed towards males (see also
Young 1974, 1975a; Young and Thomason 1974; Young 1972a). Based on
this study and several previous ones (as cited above), a general model for
ecological segregation in zones of sympatry among the five species of Mor-
pho in Costa Rica is presented as a testable set of hypotheses for further
study.
Methods
Morpho granadensis was studied at “Finca La Tigra,” a locality within
the premontane tropical wet forest zone of northeastern Costa Rica, and
VOLUME XC, NUMBER 1
37
Fig. 1. Dorsal and ventral aspects of Morpho peleides limpida (left column) and M. gra-
nadensis polybaptus from northeastern Costa Rica. Specimens part of reared and wild-caught
series in the collections of the Milwaukee Public Museum.
about 10 km east of La Virgen (10°23'N, 84°07'W; about 220 m elev.),
Heredia Province. This locality is about 15 km west of the site where I
originally found M. granadensis (Young 1972a), the first known collection
of this species since its original discovery in Costa Rica in 1904 (specimen
in the Allyn Museum, a male, collected by A. Hall, at ‘‘Castillo”). This
original locality is probably Castilla (10°43'N, 85°25'W). Through baiting
(with rotten bananas) within the mixed primary-secondary tropical rain for-
est at La Tigra (Fig. 2), it was possible to obtain a new series of specimens
and mated females. Collecting of new material was kept to an absolute
minimum. Periodically between 1977 and 1981, I baited M. granadensis,
along with M. peleides and M. amathonte within an approximately 2,000
square-meter plot of forest, placing three piles of rotting bananas at scat-
38
NEW YORK ENTOMOLOGICAL SOCIETY
Eig. 2. View through the light-gap associated with the plot of mixed primary and secondary
premontane tropical rain forest at “Einca La Tigra” in northeastern Costa Rica where Morpho
butterflies were baited with rotten bananas over several years. Adults of at least three species
(see text) occur in this habitat. It borders on a cacao plantation created largely by “tunnelling”
of advanced secondary forest to allow room for cacao trees.
tered places along two different foot paths within this area. Baits were
always placed at the same places as I was interested in estimating the rel-
ative abundance of the three species in various months at this somewhat
seasonal (although very irregular) locality (Fig. 3). The baiting also provided
fecundated females from which to obtain fertile ova for rearing studies. I
did not attempt to make a thorough study of abundance and temporal dis-
tribution on a daily basis (e.g., Young 1972a), but rather only collected or
observed butterflies at baits whenever possible.
I used the bagging method described earlier (Young 1978b; Young and
Muyshondt 1973) to rear caterpillars. Generally I obtained fertile ova by
confining a freshly-caught female in a large clear plastic bag containing fresh
cuttings of various suspected leguminous food plants (candidates taken from
the list generated in Young and Muyshondt 1973 for M. peleides) and keep-
ing each bag tightly closed. In this manner butterflies were kept for one to
three weeks, during which each was fed daily outside the bag with rotten
bananas. The cuttings were replaced as needed, fresh ones sprayed with
water, and eggs removed at regular intervals. With Morpho, this technique
is very successful. I deliberately tested samples of first instar larvae within
VOLUME XC, NUMBER 1
39
MONTHS
Fig. 3. A typical pattern of rainfall during which some of the studies were made at Finca
La Tigra. Note the general depression of rain during January, February and March, and a
second, less lengthy dip in April. This pattern is fairly typical over a run of several successive
years.
each batch of eggs obtained in this manner on different suspected larval
food plants, with an emphasis on comparing larval performance (survival,
growth rates) on two major food plants of M. peleides, Mucuna mens and
Machaerium seemannii, both stout woody legume vines associated with
forest-edge and secondary habitats (see Young and Muyshondt 1973) in
Costa Rica. I also obtained a third-instar larva of M. granadensis (deter-
mined to be this species from rearing to adult stage) from Philip J. DeVries,
who found it on a small “trailside” seedling of M. seemannii at “Carillo”
(San Jose Province, 600 m elev.). This discovery by Mr. DeVries constituted
the only wild record of the species on a particular food plant, as all of my
information on life cycle was generated by the rearing of larvae from ova
obtained from captive females. Whenever possible, I reared M. peleides
simultaneously with M. granadensis, and using the same food plant mate-
rial. Because both species are closely related in the achilles Linnaeus “su-
perspecies” complex of South America (see Young 1974), I suspected over-
lap in larval food plants when these rearing tests were initiated.
Additional material was reared independently by Philip J. DeVries, using
40
NEW YORK ENTOMOLOGICAL SOCIETY
eggs or larvae collected either at Carillo or ‘'Finca La Selva,” the site of
my original field study on the adults of this species and others (Young
1972a). In addition to the three species mentioned thus far, two others, M.
theseus Deyr, and M. cypris Westw., are also found at the La Tigra locality.
The locality has been the site for studies on the natural history of other
forest-associated day-flying Rhopalocera (e.g.. Young 1977, 1978a, 1980b).
Results
Distribution
Based upon my own observations in Costa Rica (1968-81) and those of
Philip J. DeVries (several pers. comms.), M. granadensis is probably dis-
tributed within a relatively narrow band of tropical rain forest along the
Caribbean watershed of the Cordillera Central, between 100 and 600 meters
above sea level. Because specimens of this form are exceedingly rare in
museum collections throughout the world, there are few data available on
distribution. In Costa Rica, the species is associated with the “Carrillo belt”
(see DeVries 1980), extending to the north slopes of the Poas volcano and
south to the Reventazon River Valley. Populations probably follow the se-
ries of ridges along this imaginary transect. It is probably not found in the
lowland coastal areas of the Caribbean watershed and it is absent from the
Pacific side of the country, even where tropical rain forest is well developed
(e.g., Corcovado National Park on the Osa Peninsula). In contrast, M. pe-
leides is widely distributed in several major climatic zones of Costa Rica
(Young and Muyshondt 1973; Young and Thomason 1974) and it is one of
the most widely distributed species throughout much of Central America,
not to mention the broad distribution of related forms in South America
(Seitz 1924). Unlike M. granadensis, specimens of M. peleides from specific
localities exhibit considerable marked variation in the markings on the un-
dersides of the wings (A. M. Young pers. obs.). Such variation is often
associated with a single batch of eggs (pers. data). The extreme variation
in wing markings and coloration in this species has been emphasized (Seitz
1924) and it exists both within and between populations. Morpho grana-
densis has a much more restricted geographical distribution in southern
Central America and northern South America (Seitz 1924) and specimens
generally lack, in my experience, the marked phenotypic variation so prev-
alent in M. peleides. Further field study in Central America should focus
upon the determination of distribution boundaries of M. gra-
nadensis, suspected to include Nicaragua and Panama (Seitz 1924). The
several specimens I have reared over the past four years match very well
the two male specimens in the Allyn Museum collection (the forementioned
specimen from A. Hall and that of W. J. Kaye, collected at “Pejeballe” in
1926).
VOLUME XC, NUMBER 1
41
Early Stages
The early stages are very similar to those of M. peleides (see Young and
Muyshondt 1973). Detailed taxonomic separation and a key to early stages
in these species and other is awaiting completion of life cycle studies of all
Central American species. A good series of eggs and caterpillars for M.
granadensis and M. peleides is available with the author. Some general
features are described here. The egg is green and within two days after
being deposited, develops a lateral reddish-brown band (Fig. 4). Unlike that
of M. peleides, this band is distinctly broken into small dots (Fig. 4). But
at Corcovado National Park in southwestern Costa Rica, the egg band of
M. peleides is also broken (P. J. DeVries pers. comm.). Egg hatches in 16
days and first instar is yellow and red (Fig. 4). The second instar (Fig. 4) is
similarly patterned but with a marked change in the distribution, coloration,
and sizes of setae. For example, distinct rows of long white setae appear at
the posterior margins of the second and third thoracic segments (Fig. 4).
Setae generally black and evenly distributed on most body segments in the
first instar. The second instar develops a set of dorso-lateral groupings of
long, filamentous red and white setae on the posterior five abdominal seg-
ments, and all body segments have long, whitish lateral setae as well (Fig.
4).
The first instar head capsule of M. granadensis possesses a greater num-
ber of setae than does that of M. peleides (Fig. 5). The medial edge of the
mandible in M. granadensis (first instars) possesses relatively short protu-
berances compared to that of M. peleides (Fig. 5). The cuticle over most of
the head capsule in M. granadensis, and exemplified by the area just above
the ocelli region, is finely studded with many small protuberances while that
of M. peleides is relatively smooth (Fig. 6). These micromorphological dis-
tinctions in the head capsule appear to be very consistent in separating the
first instar caterpillars of these two species.
The third instar is very similar to the second instar, but with further short-
ening of head capsule setae in the former (Fig. 7). Early instars generally
rest on leaves of Machaerium seemanii when not feeding (Fig. 7). As in M.
peleides (Young and Muyshondt 1973), there is a major change in the col-
oration of the caterpillar in the fourth instar (Fig. 7). The caterpillar becomes
shaded in rich hues of brown, pink, and white. There is further reduction
and coalescence of setae into sets of distinct tufts (Fig. 7) and the caterpillar
generally rests on woody stems of M. seemanii when not feeding. The fifth
instar is similar to the fourth, but with colors becoming more bland and
setae reduced in size. Prior to developing into a highly mobile green prepupa
of contracted size, the caterpillar attains a length of about 100 mm and
maximal width (laterally) of about 20 mm. The caterpillar stage, as measured
on a total of 16 individuals reared, lasts 89-96 days. These estimates may
42
NEW YORK ENTOMOLOGICAL SOCIETY
Eig. 4. Early stages of Morplw gmnadensis. Top; egg; note broken “ring” of brownish
pigmentation of the hemispherical green egg (about 1.8 mm dia.). Bottom, left to right; first
and second instar caterpillars respectively.
VOLUME XC, NUMBER 1
43
Fig. 5. Some micromorphological differences between the first instar caterpillars of M.
granadensis and M. peleides. Left column: head capsule (SEM, 20 x mag.) and right mandible
(l,000x mag.) for M. granadensis. Right column, same structures and magnifications, but for
M. peleides. Preparations and SEM work by R. Wolff.
have relatively little biological significance since caterpillars were often
transported around considerably while being reared, and therefore subject
to different environmental conditions.
The pupa (Fig. 7) is uniformly green with a pinkish-brown cremaster. It
is 34-38 mm long and with a maximal dorso-ventral width of 15.5 mm and
lateral width of 16.5 mm. Three prominent spiracles adjacent to the wing
pads are white, while others are much smaller and pinkish. The marked
protuberances of the head region are deep brown or black. The pupa stage
lasts about 24 days. Eclosion is extremely rapid, with the wings fully ex-
panded within one hour. Newly eclosed males less than two days old exude
a rich fragrance similar to that of rancid margarine.
Behavior
Oviposition-proneness in Morpho refers to the readiness of captive fe-
cundated females to deposit eggs on larval food plant cuttings. Females of
44
NEW YORK ENTOMOLOGICAL SOCIETY
Fig. 6. Sculpturing of the cuticle of the head capsules of first instar caterpillars of M.
gmmidensis (top) and M. peleides (below) for medial-lateral sections above region of ocelli
(500x mag., with SEM). Preparations and SEM work by R. Wolff.
VOLUME XC, NUMBER 1
45
Fig. 7. Early stages of Morpho granadensis. Top, left and right: third-instar caterpillars
resting on leaves of a larval food plant, Machaenum seemannii (Leguminosae). Note leaf
damage from feeding in the photograph to the right. Bottom: fourth instar caterpillar (left) and
pupa (right).
46
NEW YORK ENTOMOLOGICAL SOCIETY
M. granadensis generally require two to four days before egg-laying is ini-
tiated under these conditions, while the response is generally less than two
days in M. peleides. Successful mating cannot be achieved by confining a
virgin female with an active male in a clear plastic bag. In one such test,
the two butterflies were kept active for abut two weeks without copulation
taking place. The female was clearly a virgin given the large amounts of
body fluids exuded in the first few days of confinement, indicating a very
recent eclosion. Given tests with both M. granadensis and M. peleides, it
is likely that mating takes place very soon after eclosion in the wild. In
captivity, eggs are placed singly on the food plant, and it is known that
oviposition is single in the wild in M. peleides (Young and Muyshondt 1973).
I suspect single oviposition in M. granadensis in the wild.
Adults of both species regularly show up at fruit baits (see below) and can
sometimes be distinguished by the amount of blue reflectance and size of
the wings. At a closer distance, the distinctive coloration of the ocellus
markings on the ventral sides of both sets of wings between the two species
is very evident (see also Fig. 1). In M. granadensis, both the circulus and
ocellata associated with each of the three eyespot markings on each fore-
wing, and the set of four such markings on each hind wing are colored in
vivid reddish-orange. In M. peleides, these areas are yellowish. In M. gra-
nadensis, the circulus is a very thin reddish-pink or slightly lavender line,
while in M. peleides it is light blue. In M. granadensis the center spot of
each eyespot marking is bluish while it is white in M. peleides. The number
of eyespot markings on the forewings of individual specimens of M. peleides
is often very variable while seldom so in M. granadensis. Relative to M.
peleides, the dorsal marginal black border of both sets of wings is thinner
in M. granadensis. Good descriptions of wing coloration for both species
is given in Seitz (1924). The above comments are made more in the context
of providing a key to an immediate identification of species in the wild. A
detailed study of the evolutionary relationship between these two species
must understandably await for a detailed study of the genus as a whole,
including the examination of early stages and biogeographical patterns.
In laboratory culture, caterpillars of M. granadensis exhibit the same
‘‘dawn-dusk” feeding rhythm as do the caterpillars of M. peleides (see
Young 1972b), with the dusk peak being greater than the dawn peak. Pre-
sumably similar behavior exists in the wild.
Food Plant Preference
When offered various leguminous food plants, known to be acceptable to
the caterpillars of M. peleides from sympatric Costa Rican populations, the
first instar caterpillars of M. granadensis exhibit distinctive patterns of re-
sponse. For example, of a total of 32 first instars offered fresh leaves of
Mac ana are ns, a very common food plant of M. peleides (Young and Muy-
VOLUME XC, NUMBER 1
47
shondt 1973), all died within five days, and following the actual ingestion of
tissues along leaf edges. Similar results were obtained in tests of 17 first
instars with the leaves of Erythhna sp. From studies of M. peleides (e.g..
Young 1978b; Young and Muyshondt 1973) it is known that even first instars
preferentially eat the tissues from older leaves of M. urens and other le-
gumes, and M. granadensis in the present tests were offered both young
and old leaves. Although M. peleides can be succesfully reared to adulthood
on the leaves of peanut, Arachis hypogea (Young 1974), results are more
mixed with M. granadensis: in a total of 15 first instars tested, 10 died in
that instar, two more survived to the next instar, and the others perished
in the third instar (within two days following the molt). At the same time,
all but two of an initial group of 12 M. peleides were reared to adulthood
on the same peanut plants. Although these samples are exceedingly small,
they do provide some tentative evidence of a differential response between
these two species to legumes. The inference is that M. granadensis does
not share all of its food plants with the closely related M. peleides. A total
of 10 caterpillars of M. granadensis have been reared successfully on one
common food plant of M. peleides, Machaerium seemannii. Furthermore,
one partially-grown caterpillar was discovered feeding on this plant in the
wild (Carillo, San Jose Prov., Costa Rica) by Philip J. DeVries, thus con-
firming this woody legume vine as a food plant of M. granadensis in Costa
Rica. This species has also been reared at Finca La Selva by P. J. DeVries.
Both butterflies share the habit of placing eggs on seedlings of M. seemannii
(see also Young 1972a; Young and Muyshondt 1973) since the one cater-
pillar found in the wild was on a small seedling.
Species abundances
The irregular program of baiting Morpho in the same tract of trop-
ical rain forest over several years suggests that M. granadensis is less
abundant than two other sympatric species, M. peleides and M. ama-
thonte (Table 1). While samples are small and widely scattered over
time, they do reflect the low density property of adult populations of these
butterflies, as noted for M. peleides in tropical rain forest elsewhere in Costa
Rica (Young 1973). In tropical wet regions populations of these butterflies
tend to be of low adult densities, except when restricted to a small habitat
(e.g.. Young and Thomason 1974). The patterns of sex ratio observed in the
present study (Table 1) are fairly typical for baited Morpho, in which males
far outnumber females, regardless of the species (see also Young 1972a,
1973; Young and Thomason 1974). All three species were found to be active
at various times of the year, including the drier months of February and
March (Table 1 and Fig. 3). As noted in the footnotes to Table 1, there is
considerable variation in the production of eggs by individual captive fe-
males of M. granadensis, although again samples are very small. Although
hourly samples are not available, there is clearly considerable feeding at
48
NEW YORK ENTOMOLOGICAL SOCIETY
baits during the afternoon hours (Table 1). Based upon a very limited mark-
recapture study, there is some evidence that the same individual butterflies
may appear at a bait on successive days. For example, the male M. gra-
nadensis sighted on 20 February 1980 was captured, marked (with a fast-
drying white enamel paint — see Young and Thomason 1974 for details of
this technique), and released immediately. This male was subsequently re-
sighted at the three baits on 21 and 22 February. Similarly, two males of
M. peleides marked and released on 22 February 1980 were resighted at
the baits, and one of these was seen feeding the following day at noon (Ta-
ble 1).
Discussion
Because both M. granadensis and M. peleides are both members of the
achilles “superspecies” complex (Young 1974), it is not unexpected that
their early stages would be of similar appearance and behavior as reported
in this present study. Yet consistent morphological differences, including
larval characters, adult wing color patterns, and genitalia structure (Young
unpubl. data), indicate the forms to be two valid species. Both undoubtedly
represent the geographical extension of the achilles complex into Central
America, although one, M. granadensis is far more restricted in this eco-
logical range extension than the other. Elsewhere (Young and Muyshondt
1973), it is argued that M. peleides represents the form most successful in
the colonization of secondary habitats in both the non-seasonal and seasonal
tropical forest zones of Central America and Mexico. Morpho granadensis,
in contrast, is a restricted tropical rain forest form associated with inter-
mediate elevations along the Caribbean drainage of southern Central Amer-
ica, and the northern part of South America.
The topic of this paper is not to make an attempt to reconstruct what
might have been the ecology of a progenitor form to both species, and the
selection pressures leading to divergence. Rather, we are at the descriptive
stage of trying to gather basic natural history data that provide some clues
to present-day patterns of differentiation between the two species in a region
of sympatry and ecological overlap. What is of interest in this context is the
considerable overlap in the flying seasons of both species, their co-occur-
rence at baits in tropical rain forest, and a preference for the same habitat.
Yet M. peleides is a far more widely distributed species than M. granaden-
sis, suggesting indirectly that the two forms are not in direct competition
for resources. Rather, I tentatively conclude that the ecological require-
ments of M. granadensis, particularly as related to the exploitation of larval
food plants, represents a subset of those required by M. peleides. The data
and other reports (Young and Muyshondt 1973; Young and Thomason 1974)
suggest that M. peleides has a higher reproductive capacity (defined here
VOLUME XC, NUMBER 1
49
Table 1. Comparative abundance of three species of Morplw butterflies at baits of rotting
bananas in an approximately 2,000 m^ block of tropical rain forest in northeastern Costa Rica
as seen intermittently over several years.
Time
of day
Morpho peleides
Morpho granadensis
Morpho amathonte
Dates
Season
9$
dd
T
99
(?C?
T
99
T
VIII- 14-77
rainy
1300
0
0
0
0
1
1
0
1
1
VIII-15-77
rainy
1300
0
3
3
0
0
0
0
0
0
XI-30-78
rainy
1200
0
1
1
0
0
0
0
0
0
XII- 1-78
rainy
1130
1
0
1
u
0
1
0
2
2
XII-2-78
rainy
1100
0
0
0
0
0
0
0
2
2
XII-3-78
rainy
1330
0
1
1
0
0
0
0
6
6
XII-4-78
rainy
1400
1
0
1
0
0
0
0
2
2
XII-5-78
rainy
1000
0
0
0
0
0
0
0
1
1
III- 11-79
dry
1000
1
0
1
0
0
0
0
0
0
III- 12-79
dry
1000
1
0
1
0
0
0
0
0
0
IX-28-79
rainy
1400
0
0
0
2'’
0
2
0
L
1
X-3-79
rainy
1400
0
1
1
0
1
1
0
1
1
11-12-80
dry
1200
1
2
3
0
0
0
0
3
3
11-13-80
dry
1230
1
1
2
0
0
0
0
1
1
11-20-80
dry
1330
0
0
0
0
0
0
0
1
1
11-21-80
dry
1230
0
0
0
0
0
0
0
1
I
11-22-80
dry
1200
0
2
2
0
0
0
0
1
1
11-23-80
dry
1200
0
1
1
0
0
0
0
0
0
VI- 13-80
rainy
1400
0
2
2
0
5
5
0
0
0
VI- 14-80
rainy
1400
0
0
0
0
2
2
0
1
1
VI-15-80
rainy
1400
0
0
0
0
1
1
0
0
0
VI- 16-80
rainy
1400
0
0
0
0
1
1
0
2
2
II-3-81
dry
1500
u
2
3
0
0
0
0
0
0
II-4-81
dry
1500
0
1
1
0
1
1
0
1
1
II-5-81
dry
1500
0
0
0
0
2
2
0
1
1
VII-24-81
rainy
1300
1
0
1
0
2
2
0
1
1
VII-26-81
rainy
1300
u
0
1
0
1
1
0
0
0
Total butterfly sightings:
% rainy season sightings:
9
17
26
46.1%
3
17
20
75.0%
0
29
29
69.0%
^ In captivity this female produced 53 viable eggs in two weeks.
^ In captivity one female produced 8 viable eggs in 8 days, the other 24 eggs in 2.
This male fed continuously for at least one hour.
** In captivity this female produced 16 viable eggs in 3 days.
® In captivity this female produced 26 viable eggs in 5 days.
as the total number of eggs produced by the average female during her
lifetime) than does M. granadensis, and that this phenotypic trait is asso-
ciated with the ability of the former species to colonize a broad range of
ecological conditions throughout much of the American tropics. Thus the
lower abundance of adult M. granadensis at baits reflects a smaller breeding
population, rather than a necessarily large population dispersed over large
areas. It is interesting to note the similarities in adult abundance between
50
NEW YORK ENTOMOLOGICAL SOCIETY
M. peleides and M. amathonte in this region. Both of these species are
widespread forms throughout the lowland and premontane tropical rain for-
est region of the Caribbean drainage of Costa Rica, and that ranked abun-
dance with M. granadensis is not unexpected (Young 1972a). All three
species have single oviposition and legume-feeding caterpillars, conditions
that promote widespread distribution. Elsewhere (Young and Muyshondt
1972b) it was proposed that M. amathonte, representing the Central Amer-
ican expansion of M. anaxibia from South America, with its dazzling blue
wing colors in the males, is the dominant form of the genus in lowland
tropical rain forests, whereas forms such as M. peleides, with more subdued
wings, are the dominant representation of the genus at higher forested ele-
vations. Morpho granadensis seems to '‘fit in” by being distributed at an
intermediate elevation, and perhaps between the two major faunistic regions
for the other forms mentioned above. Given some preliminary information
available on these and other species of the genus in Central America, I
propose the summary of ecological distributions given in Figure 8. This
scheme is meant to be a starting point for further study, rather than a con-
clusion based on a solid background of field data.
Owing to its relatively narrow geographical distribution in Costa Rica and
presumably elsewhere, M. granadensis is probably a more ecologically-spe-
cialized species of mixed primary-secondary tropical rain forest than the
more cosmopolitan M. peleides. The inability of the caterpillars of M. gra-
nadensis to exploit one of the most abundant larval food plants of M. pe-
leides, Mucuna are ns, and their inability to develop successfully on a cul-
tivar such as peanuts, tentatively suggests ecological specialization in a
direction away from M. peleides. What is lacking is critical data on the
scope of larval food plants of M. granadensis in zones of overlap with M.
peleides. If it is assumed that Machaerium seemannii is the dominant larval
food plant of M. granadensis, much of the distribution of this butterfly may
be explained by the distribution of this woody vine. Although this vine is
abundant at higher elevations in the same region of Costa Rica, where it is
exploited by M. peleides (Young and Muyshondt 1973), the marked absence
of M. granadensis from this region suggests the operation of other factors
generating this distribution. With an absence of critical biogeographical
data, perhaps larval food plant distribution coupled with other factors such
as climate play a joint role in limiting M. granadensis. Whether or not
ecological differentiation within larval food plant populations, in which the
defensive chemistry against herbivorous insects is altered geographically,
plays a role in the absence of M. granadensis from both montane and coastal
tropical rain forest zones cannot be ruled out at this time. As pointed out
by Ehrlich and Raven (1969), there is a need to consider the evolutionary
history of a species in determining patterns of ecological differentiation.
The data on adults suggest that the short, often erratic dry season char-
VOLUME XC, NUMBER 1
51
The species of Moroho butterflies found in premontane-
to- lowland tropical wet forest region of northeastern Costa
Rica, and their general habits.
SPECIES
MAJOR ADULT FLIGHT
HABITAT(S) HEIGHT RANGE
RELATIVE
ABUNDANCE
LARVAL FOOD-
PLANT TAXA
M. Deleides
various secondary;
large forest light gaps
1-3 M
high
several
Leguminosae
M. aranadensis advanced
secondary; large forest
light gaps
1-3M
low
prob. few
Leguminosae
M. amathonte
primary forest
1-6 M
intermediate
prob. few
Leguminosae
M. theseus
primary forest
6-30 M
low
prob. few
Menispermaceae
M. cvpris
primary forest
10-30 M
low
prob. few
Leguminosae
Defined here as 0-200 meters above sea level.
Fig. 8. A schematic summary of the major ecological attributes for the species of Morpho
butterflies found in northeastern Costa Rica. Data on habitats, flight ranges, and relative abun-
dance from direct field observations over several years, and larval food plant data from both
field and laboratory study, and the literature.
acteristic of this region of Costa Rica does not appreciably affect the pop-
ulation structure of the butterflies. In all three species, adults are active at
various times of the year. It is likely that the larval food plants are not
deciduous and therefore remain evergreen during dry periods. Thus breed-
ing can be continuous throughout the year. What may vary is the distribution
of natural adult foods. I have noticed that during dry periods adults arrive
at baits faster than in the rainy season. Increased dryness may result in
increased feeding to maintain water balance and it may also result in a
decrease in density of suitable foods, which include sweet-smelling rotten
fruits and soupy fungal growths (Young 1979, 1980a). Throughout the year,
females are rare at baits, possibly the result of a sexual difference in pref-
52
NEW YORK ENTOMOLOGICAL SOCIETY
Table 2. Summary of the major ecological characteristics of Morpho peleides and M. gra-
nadensis in terms of explaining their distribution.^
Phenotypic and other characters
responding to selection
Determined and/or predicted*’
character states
Morpho peleides
Morpho granadensis
(1)
Geographical range
widespread
restricted
(2)
Local endemism and restriction of gene
flow
low*
high*
(3)
Distribution across marked elevational
gradients
high
low
(4)
Ability to occupy highly seasonal regions
good
poor
(5)
General spatial patchiness of resident
populations over approx. 1,000-meter
sections of a region
low
high
(6)
Dispersal ability of adults
high
moderate-to-low*
(7)
Intrapopulational variation
high*
low*
(8)
Interpopulational variation
low*
moderate-to-high*
(9)
Habitat selection
generalized (low)
grading into
specialized (high)
(10)
Main habitat
grades of secondary
forest
advanced secondary
forest
(11)
Colonizing ability
high
moderate-to-low*
(12)
Local population density
high
low
(13)
Regional breadth of larval foodplants
high
low*
(14)
Acceptability of each other’s larval
foodplants
high
low
(15)
Relative abundance of larval foodplants
per unit area of suitable habitat
high
low*
(16)
Oviposition strategy
single and scattered
single and scattered
(17)
Oviposition proneness in captivity
high
low
(18)
Predation/parasitism on immature stages
high
high‘s
(19)
Overall fecundity (average per female)
high
moderate
(20)
Egg-adult developmental time
very ;
similar*
(21)
Larval dawn-dusk feeding
present
present
(22)
“Bait-ability” of adults
excellent
fair-to-good
(23)
Daily turnover of adults at baits
high
moderate
(24)
Sex ratio of adults at baits
skewed males
skewed males
(25)
(26)
Diurnal rhythmicity of adult feeding
Percent character states in common;
28.0%
present
present
“ Analyses of these characters should include an examination of breeding populations in both
regions of species overlap and non-overlap.
Character states marked with an asterisk are predictions, while unmarked ones are empir-
ical observations from previous studies (Young 1971a, b, 1972a, b, 1973, 1974, 1975a, 1978b,
1979, 1980a; Young and Muyshondt 1973; Young and Thomason 1974) and unpublished data
(Young).
VOLUME XC, NUMBER 1
53
erence for feeding on rotten bananas. Rearing studies (Young and Muy-
shondt 1973) indicate that sex ratio is near unity in egg batches. The com-
plete absence of female M. amathonte from baits may indicate, together
with the scarcity of females in the other two species, that females have
different feeding habits from those of males. Because observations were
made at various times of the day, it is unlikely that females were missed at
the baits. Rather, their food requirementss may be considerably different
from males in all three species, and possibly linked to nutritional demands
associated with egg production.
Table 2 summarizes a proposed model for the overall evolution and eco-
logical differentiation of M. granadensis and M. peleides. This framework
is suggested as a means for developing some testable hypotheses on the
ecological properties of these two species, a topic of considerable interest
since both are members of the same ‘‘superspecies” complex and exhibit
considerable sympatry at the same time. The proposed model can also be
eventually applied to the other species of Morpho as natural history data
are gathered on them. Basically this model makes some testable predictions
about the differences in genetic structure and ecological properties between
a pair of species, one of which is geographically and ecologically more
restricted than the other in the tropics.
Acknowledgments
I thank Dr. J. Robert Hunter for allowing me to conduct these studies at
Finca La Tigra, and Dr. Lee D. Miller of the Allyn Museum of Entomology
for hospitality and access to the Morpho collections there. Mr. Philip J.
DeVries kindly shared with me his independent data on Morpho granaden-
sis. Costs of publication were provided by the Milwaukee Public Museum.
I thank Phil DeVries for reading an earlier draft of this manuscript.
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and soupy fungi in tropical forests: variations on a theme developed by the Muyshondts
and Arthur M. Shapiro. The Entomologist’s Record 90:63-70.
. 1980b. Notes on the behavioral ecology of Perrhyhris lypera (Pieridae) in northeastern
Costa Rica. J. Lepid. Soc. 34:36-47.
and A. Muyshondt. 1972a. Biology of Morpho polyphemus in El Salvador. J. New
York Entomol. Soc. 80:18^2.
and . 1972b. Ecological and geographical expansion in tropical butterflies of the
genus Morpho in evolutionary time. Rev. Biol. Trop. 20:231-263.
and . 1973. Biology of Morpho peleides in Central America. Caribbean Journal
of Science 13: 1-49.
and J. H. Thomason. 1974. Demography of a confined population of the butterfly
Morpho peleides during a tropical dry season. Studies on the Neotropical Fauna 9:1-34.
Section of Invertebrate Zoology, Milwaukee Public Museum, Milwaukee,
Wisconsin 53233.
Received for publication September 10, 1981.
NEW YORK ENTOMOLOGICAL SOCIETY
XC(1), 1982, pp. 55-57
VLADIMIR NABOKOV 1899-1977: A NOTE ON A
LATE ENTOMOLOGIST
“Look, how pretty,” said observant Chateau.
A score of small butterflies, all of one kind, were settled on a damp patch
of sand, their wings erect and closed, showing their pale undersides with
dark dots and tiny orange-rimmed peacock spots along the hindwing mar-
gins; one of Pnin’s shed rubbers disturbed some of them and, revealing the
celestial hue of their upper surface, they fluttered around like blue snow-
flakes before settling again.
“Pity Vladimir Vladimirovich is not here,” remarked Chateau. “He
would have told us all about these enchanting insects.”
“I have always had the impression that his entomology was merely a
pose.”
“Oh no,” said Chateau.
(from Pnin by V. Nabokov, Garden City, N.Y.: Doubleday, 1957)
Not just Timofey Pavlich Pnin, the title character of his 1957 novel, but
many “real” people thought that Vladimir Vladimirovich’s — that is, Vla-
dimir Nabokov’s — entomology was an affectation, a little marginal color-
ation on the hindwings of his prolific writing and busy teaching careers. But
it wasn’t. “My passion for lepidopterological research, in the field, in the
laboratory, in the library, is even more pleasurable than the study and prac-
tice of literature, which is saying a good deal.” (Interview for Wisconsin
Studies in Contemporary Literature in 1966.)
Nabokov was born in old St. Petersburg in 1899. By the age of seven he
was chasing butterflies across Russian meadows and marshes (see Chapter
Six of Speak, Memory: An Autobiography Revisited, New York: Putnam,
1966). In a 1920 entomological article, written during his Western European
emigre period, he described some species he observed in the Crimea. It was
his first published writing in English, 20 years before he was forced to flee
Europe and abandon his “docile” Russian and decided to tame English for
his creative purposes. For seventy years, until he died in Montreux, Swit-
zerland in 1977, Vladimir Nabokov pursued Lycaeides and Plebejinae, dis-
sected and drew their genitals, described their haunts and once said, “I
have often dreamt of a long and exciting career as an obscure curator of
lepidoptera in a great museum.” (Interview for Life Magazine in 1964.)
His published lepidoptera is not minor. From 1941 to 1948 he worked
very hard and very happily over a microscope at the laboratory of the
Museum of Comparative Zoology, Harvard. From 1941 to 1953, at the same
time he was writing and teaching, he published some 10 major articles in
entomological journals (see pp. 314-335 of Strong Opinions, New York:
56
NEW YORK ENTOMOLOGICAL SOCIETY
McGraw-Hill, 1973). '‘I am the author or the revisor of a number of species
and sub-species mainly in the New World . . . Several butterflies and one
moth have been named for me . . . There is also a genus Nabokovia Hem-
ming, in South America.” (Interview for Bayerischer Rundfunk in 1971.) At
one time he had begun work on an illustrated “Butterflies in Art,” from
Egyptian antiquity to the Renaissance.
The playful lepidopterological images that flutter across the pages of his
stories and novels had a very precise purpose. They established contact
between two of the great loves in his life, literature and lepidopterology,
“between the precision of poetry and the excitement of pure science.”
(Interview for the BBC in 1962.)
Michael Juliar,^ 74 Kings Road, Little Silver, New Jersey 07739.
Bibliography
A Eew Notes on Crimean Lepidoptera. London: The Entomologist, Eebruary, 1920, Vol. 53,
No. 681, pp. 29-33.
Notes on the Lepidoptera of the Pyrenees Orientales and the Ariege. London: The Entomol-
ogist, November, 1931, Vol. 64, No. 822, pp. 255-257; and December, 1931, Vol. 64,
No. 823, pp. 268-271.
On Some Asiatic Species of Carterocephalus. New York: Journal of the New York Entomo-
logical Society, September 1941, Vol. 49, No. 3, pp. 221-223.
Lysandra cormion, a New European Butterfly. New York: Journal of the New York Ento-
mological Society, September 1941, Vol. 49, No. 3, pp. 265-267.
Some New or Little-Known Nearctic Neonympha. Cambridge, MA: Psyche, Journal of Ento-
mology, September-December, 1942, Vol. 49, Nos. 3-4, pp. 61-80.
The Eemale of Neonympha maniola Nabokov. Cambridge, MA: Psyche, Journal of Entomol-
ogy, March-June, 1943, Vol. 50, Nos. 1-2, p. 33. (Note to article in Vol. 49, Nos. 3^.)
The Nearctic Eorms of Lycaeides Hub. (Lycaenidae, Lepidoptera). Cambridge, MA: Psyche,
Journal of Entomology, September-December, 1943, Vol. 50, Nos. 3^, pp. 87-99.
Notes on the Morphology of the Genus Lycaeides (Lycaenidae, Lepidoptera). Cambridge,
MA: Psyche, Journal of Entomology, September-December, 1944, Vol. 51, Nos. 3^,
pp. 104-138.
Notes on Neotropical Plebejinae (Lycaenidae, Lepidoptera). Cambridge, MA: Psyche, Journal
of Entomology, March-June, 1945, Vol. 52, Nos. 1-2, pp. 1-61.
A Third Species of Echinargus Nabokov (Lycaenidae, Lepidoptera). Cambridge, MA: Psyche,
Journal of Entomology, September-December, 1945, Vol. 52, Nos. 3-4, p. 193 (Note
to article in Vol. 52, Nos. 1-2).
Southern Pierids in New England, Cambridge, MA: Psyche, Journal of Entomology. Septem-
ber-December, 1946, Vol. 53, Nos. 3-4, p. 42. (Observational note.)
Sphingids over Water. Cambridge, MA: The Lepidopterists’ News, 1947, No. 1, p. 82.
(Observational note.)
A New Species of Cyclargus Nabokov (Lycaenidae, Lepidoptera). London: The Entomologist,
December, 1948, Vol. 81, No. 1027, pp. 273-280.
’ The author, who is working on a bibliography of Nabokov, requests copies or reprints of
the articles listed, or information on where they can be obtained.
VOLUME XC, NUMBER 1
57
The Nearctic Members of the Genus Lycaeides Hubner (Lycaenidae, Lepidoptera). Cambridge,
MA: Bulletin of the Museum of Comparative Zoology, February, 1949, Vol. 101, No. 4,
pp. 479-541. (This article occupies the entire issue.)
Remarks on F. M. Brown’s “Measurements and Lepidoptera.’’ New Haven, CT: The Lepi-
dopterists’ News, 1950, Vol. 4, Nos. 6-7, pp. 75-76.
The Female of Lycaeides argyrognomon sublivens. New Haven, CT: The Lepidopterists’
News, 1952, Vol. 6, Nos. 1-3, pp. 35-36.
On Some Inaccuracies in Klots’ “Field Guide.’’ New Haven, CT: The Lepidopterists’ News,
1952, Vol. 6, Nos. 1-3, p. 41.
Butterfly Collecting in Wyoming. New Haven, CT: The Lepidopterists’ News, 1953, Vol. 7,
No. 2, pp. 49-52.
Comments on Lycaeides argyrognomon in Wisconsin. New Haven, CT: The Lepidopterists’
News, 1953, Vol. 7, No. 2, p. 54. (Quotation from a letter addressed to another
collector.)
NEW YORK ENTOMOLOGICAL SOCIETY
XC(1), 1982, pp. 58-61
BOOK REVIEW
Vectors of Plant Pathogens. Edited by K. F. Harris and K. Maramorosch.
Published by Academic Press, New York, London, Toronto, Sydney and
San Francisco, 1980. ISBN 0-12-326450-2. Price $48.00. Pp. xiv + 467.
This 17-chapter book, authored by 21 contributors from 8 different coun-
tries of the world, is the third in a multi-volume series of treatises on the
general topic of vectors, pathogens and plant diseases. The two preceding
volumes, namely '‘Aphids as Virus Vectors” and ‘‘Leafhopper Vectors and
Plant Disease Agents,” appeared in 1977 and 1979, respectively.
Each one of the first 10 chapters of the book, with a certain exception
concerning Chapter 8 (see below), deals with a different taxonomic group
of insects which is known to contain species that act as vectors of plant
disease agents. Chapters 14-17 are devoted to taxa other than Insecta that
include vectors, to wit: mites, nematodes and fungi.
Two vector-borne plant diseases which in recent years have gained world
notoriety were quite understandably allotted special chapters rather than
being treated in the relevant chapters dealing with their respective vector
group. The diseases are: (a) Dutch elm disease, a barkbeetle-borne fungal
pandemic, prevalent mainly in temperate regions of the world (Chapter 8);
and (b) lethal yellowing of coconut palm, a devastating disease in the tropics
and subtropics, which is presumably caused by a mycoplasma-like organ-
ism, though its vector transmission still requires unequivocal proof (Chapter
11). The singling out of these two plant diseases by apportioning a separate
chapter to each one of them in a book which is chiefly concerned with
animal vectors, seems nevertheless justifiable in view of the worldwide im-
pact of these diseases on both the economy and the environment. Moreover,
thanks to the editors’ apparent prudence there is no duplication of coverage
between Chapter 8 handling the beetle-borne Dutch elm disease and Chapter
7 which comprehensively deals with beetles vectoring plant pathogens, in
spite of the fact that the two chapters were written by different authors.
The same virtually holds true for Chapters 12 and 13 which review the
involvement of insects in the transmission of bacterial and fungal phyto-
pathogens, respectively. By narrowing the scope of the chapters on the
taxonomic groups of insects down to vectors of viral pathogens mainly, the
editors managed to preclude any unnecessary repetitiousness likely to occur
in these two chapters dealing with the transmision by insects of nonviral
plant pathogens viz. bacteria and fungi.
It may perhaps be argued that Chapter 1 (aphids, leafhoppers and plant-
hoppers) is apparently redundant since the two preceding volumes of the
series bear the titles of ‘‘aphid vectors of plant viruses” and “leafhopper
vectors and plant pathogens.” However, in a rapidly developing area of
VOLUME XC, NUMBER 1
59
research like the transmission of plant pathogens by homopterous vectors,
a tri- or even biennial updating of a review is certainly not unwarranted, let
alone the fact that the chapter under discussion (written by one of the edi-
? tors — K. F. Harris) has been condensed into a mere 12-page synoptic re-
! view. At any rate, it would be utterly inconceivable to omit, for whatever
reason, a chapter on aphids and leafhoppers from a textbook entitled “Vec-
I tors of Plant Pathogens.”
|, A marked improvement in the References lists concluding each chapter,
as compared to the two previous volumes of the series, is the providing of
full titles of the cited publications. This will undoubtedly be very much
appreciated by all users of the book.
In a reference book like this, comprising 17 individual contributions of
different authors, a good index, going down to exhaustive detail, is more
than essential. The challenge was indeed well taken up here in the form of
a 29-page index, listing some 2,800 items.
Thus, the careful choice of top expert authors, coupled with a good mea-
; sure of skillful judgment exercised by the editors, and the highly professional
[presentation on the part of the publishers, have all together resulted in a
most welcome, comprehensive and updated compilation of core reference
and background information on its subject. It will doubtlessly be indispen-
sable in providing the most up-to-date handbook on vector transmission of
' plant disease agents currently available. As a university instructor charged
with the teaching of a course on this very subject, I can state with a great
deal of gratitude that the book has definitely made my task very much easier
now. Without any hesitation I thoroughly recommend this book to special-
ists and non-specialists alike who have an interest in vector-related plant
pathology, and/or phytopathogen-related entomology, whether it be from
the research or teaching point of view.
I. Harpaz, Hebrew University of Jerusalem, Rehovot, Israel.
Handbook of Plant Virus Infections: Comparative Diagnosis. E. Kurstak,
ed. Elsevier-North Holland. 944 pp. $192.75.
Plant virus infections can spread in various ways and the most prevalent
is by means of insect vectors. Therefore the subject of this treatise is of
special interest to entomologists. Hundreds of plant virus diseases have
been described all over the world, but the causative viruses were often
inadequately identified and, in many instances, the viral nature of the dis-
eases not properly ascertained. In fact, more than 200 viruses, incompletely
characterized by 1981, remain ungrouped, and only 23 virus groups and 2
60
NEW YORK ENTOMOLOGICAL SOCIETY
families of viruses have been accepted as properly characterized. There
exists a real need for a comprehensive text that would facilitate the proper
diagnosis of plant virus infections, and the present volume is the first attempt
to fill the existing gap.
Many of the basic procedures used for the description and diagnosis of
viruses are described in detail by the contributors of this book. In most
instances the natural means of transmission is highly diagnostic, but there
are certain virus groups with specific vectors, others where vectors are as
yet unknown, and still others transmitted by seed, pollen, or by vegetative
means. The comprehensive contents of this book are divided into 6 parts.
The first deals with plant virus taxonomy, the second with non-enveloped
RNA viruses, the third with enveloped RNA viruses, the fourth with elon-
gate RNA viruses. The fifth part deals with DNA viruses and the last one
with viroids.
Entomologists will be especially interested in the chapters in which insect
and other invertebrate vectors and their interactions with plant-pathogenic
viruses are presented. Chapter 2 describes maize chlorotic dwaft and its only
natural leafhopper vector Graminella nigrifrons. The third chapter deals
with the identification of Tymoviruses, transmitted by beetles. The Tom-
busviruses (Chapter 4) have but a few known natural vectors, such as the
flea beetles of the genera Phyllotreta and Psilliodes for turnip crinkle. Chry-
somelid vectors, Lema melanopa and L. lichensis, as well as Ceratoma
trifurcata transmit southern bean mosaic virus (Chaper 5). Luteoviruses are
transmitted by more than a dozen specific aphid vectors and the persistent
virus-vector interaction accounts for the distribution of infected plants in
the field (Chapter 6). Chapter 8 deals with Comoviruses, transmitted mainly
by chrysomelid beetles. There are very good electron micrographs of the
pea enation mosaic virus (Chapter 10) but none of the virus in its major
aphid vector Acyrthosiphon pisum, in which the virus is known to multiply.
The ubiquitous Cucumoviruses (Chapter 11) are transmitted in nature by
some 75 aphid species. Their biological aspects and vector control have
been described in great detail. Among the Ilarviruses (Chapter 13) one,
tobacco streak, is transmitted by a thrips, Franklinella sp., another (Prunus
ringspot) by a mite, Vasates fockeni. Alfalfa mosaic virus (Chapter 14) is
transmitted by many aphid species in the nonpersistent manner. On the
other hand, Reoviruses (Chapter 15) are transmitted biologically by leaf-
hoppers and planthoppers. This chapter contains detailed tables concerning
vector species as well as outstanding electron micrographs by Prof. E. Shi-
kata of Hokkaido U., Japan, of virions in fatbody cells of Nephotettix cinc-
ticeps infected with rice dwarf virus, and of Unkanodes albifascia with rice
black-streaked dwarf virus. The use of immune electron microscopy for the
identification of plant reoviruses is among the highlights of this chapter. In
the following chapter (No. 16) there is an excellent tabular presentation of
VOLUME XC, NUMBER 1
61
rhabdovirus vectors, and of the world- wide distribution of the diseases.
Electron micrographs of virions in cells of infected plants illustrate this
chapter, prepared by R. I. B. Francki (Australia), E. W. Kitajima (Brazil)
and D. Peters (Netherlands), the three foremost authorities on these ubiq-
uitous viruses. The thrips-borne tomato spotted wilt virus is described in
Chapter 17. The diagnosis of aphid-borne Poty viruses is in Chapter 23. The
plant DNA Caulimoviruses (Chapter 25) are transmitted by Myzus persicae
and Brevicoryne brassicae aphids. Some of the Geminiviruses (Chapter 26)
have vectors belonging to the cicadellid leafhoppers, such as Orosius ar-
gent atus and Nesoclutha pallida, while others are transmitted by whiteflies,
Bemisia tabaci.
The constant growth of the world population increases the need for more
food and fiber, and in light of this it is important to reinforce current
knowledge of plant virus diseases and of plant virus-vector interactions, so
as to control the numerous diseases more efficiently. Plant pathologists,
entomologists, teachers and students will find this large volume to be an
outstanding source of information. Because of its cost, however, it will most
likely be limited to libraries at colleges and universities.
Karl Maramorosch, Waksman Institute of Microbiology, Rutgers-The
State University, New Brunswick, New Jersey.
Insects and Other Invertebrates of the World on Stamps. William E. Stan-
ley, ed. Biology Unit, American Topical Association Handbook No. 98.
140 pp. $10.— ATA, 3306 North 50th St., Milwaukee, WI 53216.
This 140-page handbook is the only complete data source on insects and
other invertebrates, illustrated on stamps of the world. It brings together in
a concise form in three sections (1) butterflies and moths, (2) other insects
and (3) other invertebrates. Taxonomic listings by phyla, classes, subclass-
es, orders and families, with Latin and common names, as well as country
listings follow the style of earlier biological stamp handbooks issued by
ATA. The checklist specifies the country, date of issue, Scott, Minkus or
Stanley Gibbons stamp catalog numbers, face value, Latin name and au-
thority, as well as an indication of whether the reproduction is in natural
colors. This little handbook will be a must for entomologists who collect
insects on stamps.
Karl Maramorosch, Waksman Institute of Microbiology, Rutgers Uni-
versity.
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Journal of the
New York Entomological Society
VOLUME XC JUNE 1982 NO. 2
EDITORIAL BOARD
Editor Associate Editors
Dr. Karl Maramorosch Dr. Lois J. Keller, RSM
Waksman Institute of Microbiology Dr. Herbert T. Streu
Rutgers University
New Brunswick, New Jersey 08903
Publication Committee
Dr. Randall T. Schuh
American Museum of
Natural History
Dr. Louis Trombetta
St. Johns University
CONTENTS
Habitat effect on fungal substrate selection by a leaf-cutting ant Harold G. Fowler 64-69
A new species of Trachymyrmex fungus-growing ant (Hymenoptera: Myrmicinae;
Attini) from Paraguay Harold G. Fowler 70-73
Experimental removal of 17-year cicada nymphs and growth of host apple trees
Richard Karban 74-81
Altered hatch sequence of males and females from unchilled eggs of a “non-diapause”
gypsy moth strain (Lepidoptera: Lymantriidae)
Nancy Fike Knop, Marjorie A. Hoy and Michael E. Montgomery 82-86
Comparative feeding behavior of two aphid species: bean aphid {Aphis fabae
Scopoli) and pea aphid {Acyrthosiphon pisum (Harris)) (Homoptera: Aphididae)
Ronald P. Salyk and Daniel J. Sullivan S.J. 87-93
Seasonal flight of the cranberry girdler determined with pheromone traps
J. A. Kamm and L. M. McDonough 94-98
Notes on the interaction of the skipper butterfly Calpodes ethlius (Lepidoptera:
Hesperiidae) with its larval host plant Canna edulis (Cannaceae) in Mazatlan,
State of Sinaloa, Mexico Allen M. Young 99-1 14
Oviposition by Two Heliconius species: Comments on a paper by Dr. A. Young
L. E. Gilbert 115-116
Errata: Over-exploitation of larval host plants by Heliconius butterflies
Allen M. Young 117-118
Miridae and Coleoptera associated with tulip tree flowers at Ithaca, New York
David Andow 119-124
Susceptibility of gypsy moth larvae to several species of entomogenous fungi
S. S. Wasti and G. C. Hartmann 125-128
Habitat differences in feeding habits and body size of the predatory stinkbug
Perillus circumcinctus (Hemiptera: Pentatomidae) Edward W. Evans 129-133
Book reviews
114, 133, 134
NEW YORK ENTOMOLOGICAL SOCIETY
XC(2), 1982, pp. 64-69
HABITAT EFFECT ON FUNGAL SUBSTRATE SEEECTION
BY A FEAF-CUTTING ANT
Harold G. Fowler
Abstract. — Colonies of the grass-cutting ant, Acromyrmex landolti frac-
ticornis, ranked potential fungal substrate in field preference bioassays as
a function of the dominant grass species in the field in which they were
located. Mahalanobis distances between and among colony preferences be-
tween and among habitats revealed that there was less variability among
colonies within any habitat than between habitats. These results suggest
that fungal substrate preferences are environmentally induced, probably as
a result of early experience of workers.
The extreme Catholicism demonstrated by leaf-cutting ants (genera Atta
and Acromyrmex; Formicidae: Attini) in harvesting vegetation to serve as
fungal substrate is one of the most intriguing and applied problems con-
fronting researchers in Fatin America. Yet, despite the efforts of many
workers, why leaf-cutting ants do what they do is still unknown, although
patterns are beginning to emerge. Cherrett (1968) and Rockwood (1976,
1977) have argued for a conservative management of vegetative resources
by colonies of leaf-cutting ants based upon their experience with tropical
forest species. Recently, Fowler and Stiles (1980) have suggested that a
more parsimonious explanation, based upon the patchiness of the vegetative
resources, might better explain the patterns observed by Cherrett and Rock-
wood.
There is now strong evidence that fungal substrate selection is based, in
part, on plant chemistry (Cherrett and Seaforth 1970; Cherrett 1972; Fittle-
dyke and Cherrett 1975, 1978; Rockwood and Glander 1979; Stradling 1978;
Hubbell and Wiemer 1981). This fact alone, however, cannot explain plant
selection, especially by colonies in different habitats (Rockwood 1976,
1977). I now report on the effect of habitat in determining plant selection in
a leaf-cutting ant. These studies were conducted near Asuncion, Paraguay,
in 1976.
The publication costs of this article were defrayed in part by page charge payment. This article
must therefore be hereby marked “Advertisement" in accordance with 18 U.S.C. §1734 solely
to indicate this fact.
VOLUME XC, NUMBER 2
65
Methods
Based on a previous study (Fowler and Robinson 1977), the ranked pref-
erences for 5 species of introduced forage grasses were determined from
laboratory colonies. These ranked preferences were Digitaria smutsii, Hy-
parrhenia rufa, Digitaria decumbens, Tripsacum laxum, and Pennisetum
ciliare. Thirty field bioassays were conducted on colonies in each of three
essentially mono-specific habitats of D. smutsii, D. decumbens, and P. cil-
iare. Each bioassay consisted in clearing vegetation from nest mounds, en-
closing the mound with a wire and plastic enclosure to prevent foraging,
and then offering 10 uniform pieces of each of the 5 grasses to the colony
in a random array, and recording the numbers taken. Bioassays were ter-
minated when all of one type of grass was entered into the nest.
Data analysis consisted in subjecting the acceptance data to a multivariate
analysis of variance (Morrison 1976) to test for an overall habitat effect, and
the use of discriminant analysis (Morrison 1976) to obtain the Mahalanobis
distances separating the preferences of colonies established in distinct hab-
itats.
Results
The mean number of pieces picked up by the ants and entered into the
nests is given in Table 1. Overall ranking of the grasses taken in each habitat
were the same, with the exception of the ranking by colonies located in P.
ciliare monocultures.
When these data were subjected to a multivariate analysis of variance, it
was found that the vectors characterizing habitat-specific colony prefer-
ences for the 5 forage species (Table 2), were highly significant (Pillai’s
trace = 0.5306, Fao,im = 6.0 , P < 0.0001) indicating an important effect of
habitat on the resulting preferences of the colonies located within them. In
particular, these differences arose from the colony selections relating to P.
Table 1. The effect of habitat on grass-selection by Acromyrmex landolti fracticornis col-
onies located in mono-specific stands of forage in Paraguay.
Grass species taken
Mean (±SD) of 10 grass pieces taken
by colonies located in fields of:
D. smutsii
D. decumbens
p.
ciliare
Digitaria smutsii
7.57 ± 2.49
8.83 ± 2.15
7.83
± 2.03
Hyparrhenia rufa
7.50 ± 2.03
6.23 ± 2.71
6.63
± 2.52
Digitaria decumbens
6.77 ± 2.63
4.86 ± 2.78
6.93
± 2.76
Tripsacum laxum
2.60 ± 1.69
3.20 ± 2.66
2.30
± 1.66
Pennisetum ciliare
1.93 ± 1.62
3.20 ± 2.56
5.43
± 3.02
66
NEW YORK ENTOMOLOGICAL SOCIETY
Table 2. Characteristic roots and vectors of E inverse x H matrices, where E is the error
sum square and cross product matrix, and H is the type IV sum square and cross product
matrix for habitat effect on the preferences of colonies located in the 3 habitats of Table 1.
Characteristic
root
Characteristic vector
D. smutsii
H. rufa
D. decumbens
T. laxum
P. ciliare
0.428
-0.0155
-0.0069
0.0116
-0.0208
0.0396
0.300
-0.0198
0.0200
0.0312
-0.0194
-0.0159
0.000
-0.0083
0.0034
0.0099
0.0456
0.0049
0.000
0.0180
0.0414
-0.0078
0.0044
0.0096
0.000
0.0397
0.0056
0.0211
0.0002
-0.0026
ciliare and D. decumbens (ANOVA F values of 15.41 and 5.32 respectively
for pick-up per habitat).
An analysis of the covariance matrices of grass-selection by colonies in
each habitat (Table 3) revealed a significant deviation from homogeneity
Table 3. Covariance matrices of grass-species preferences exhibited by field colonies of
Acromyrmex landolti fracticornis in habitats where they occur, and a pooled covariance matrix
for overall preferences (intra-habitat). Habitat is indicated in parenthesis.
Grass species:
D. smutsii
H. rufa
D. decumbens
T. laxum
P. ciliare
(D. smutsii)
D. smutsii
6.18
-2.53
0.34
-0.90
0.04
H. rufa
4.12
-0.16
-0.07
0.28
D. decumbens
6.94
1.14
-0.33
T. laxum
2.86
-0.34
P. ciliare
{D. decumbens)
2.62
D. smutsii
4.63
0.73
1.49
-0.21
1.03
H. rufa
7.36
-1.73
-0.53
-0.05
D. decumbens
7.71
1.03
2.23
T. laxum
7.06
3.68
P. ciliare
{P. ciliare)
6.58
D. smutsii
4.14
-2.79
0.47
-0.39
0.49
H. rufa
6.38
-2.58
0.08
0.58
D. decumbens
7.65
1.06
-0.28
T. laxum
2.77
-1.79
P. ciliare
(Pooled — all habitats)
9.15
D. smutsii
4.98
-1.53
0.77
-0.50
0.52
H. rufa
5.95
-1.49
-0.17
0.27
D. decumbens
7.43
1.08
0.54
T. laxum
4.23
0.52
P. ciliare
6.11
VOLUME XC, NUMBER 2
67
Table 4. Mean Mahalanobis distances of substrate preferences from colonies of each habitat
to the habitat mean substrate preferences.
Substrate preferences for
colonies in habitats of
Mahalanobis distance to:
D. smutsii habitat
mean preferences
D. decumbens habitat
mean preferences
P. ciliare habitat
mean preferences
Digitaria smutsii
6.723
10.980
9.457
Digitaria decumbens
8.843
8.753
9.653
Pennisetum ciliare
11.899
11.628
7.783
(x^ = 52.26, P < 0.0033). Thus, the overall pick-up of grass species in each
habitat was determined by the characteristics of the colonies within those
habitats, and a global pattern of substrate ranking and preference cannot be
extracted across habitat boundaries.
Because of the lack of homogeneity in the covariances, a quadratic dis-
criminant function was needed to separate the habitat specific pick-up re-
sponses of colonies. From this, it possible to calculate the generalized
squared distances (Mahalanobis distances) for the habitat specific prefer-
ences (Table 4). Without exception, the preferences of colonies located in
D. smutsii habitats are more dissimilar to those of colonies located in P.
ciliare than they are to the preference means of colonies located in D.
decumbens. Since the overall preferences of these 2 grasses are relatively
high (Table 1), this result is expected. However, colonies located in the low
preference P. ciliare habitats showed a highly variable selection of grasses
(Table 1), because all of the offered grasses were of superior rank. Yet,
these colonies did not harvest significantly more of these than the grass it
was normally foraging, P. ciliare (Table 1).
Discussion
The results presented here suggest that conditioning can greatly affect the
patterns of plant selection in leaf-cutting ants, and may explain, in part, the
intra-colonial differences in substrate selection noted in tropical rain forests
(Rockwood 1976, 1977). In a tropical forest, the surrounding vegetation
pattern is much more complex than that discussed here for grass-cutting
ants in mono-specific pastures. However, the foraging territories of these
species is undoubtedly characterized by its own characteristic vector of
plant species. If worker ants learn, probably through early experience (Jais-
son 1980), the characteristics of its habitat, it is very likely that the colony
specific foraging patterns noted by Rockwood (1976, 1977) could be induced.
The results of this paper also suggest that a more tractable analysis of the
foraging behavior of leaf-cutting ants could probably result from studies in
less complex habitats instead of tropical rain forests. In less complex hab-
itats, many complicating and intertwined parameters are naturally elimini-
68
NEW YORK ENTOMOLOGICAL SOCIETY
nated, which would allow for an analysis of leaf-cutter foraging behavior
from basic to complex elements, instead of complex to basic elements as is
by necessity dictated by studies in tropical rain forests.
Acknowledgments
This research was supported by the following institutions: the New Jersey
Agricultural Experiment Station, the National University of Asuncion, the
Paraguayan Ministry of Agriculture, the U.S. Peace Corps, UNESCO, and
the British Ministry of Overseas Development. I thank S. W. Robinson and
N. Gonzales Romero for their support, and Elena Fowler for her help in data
collection. New Jersey Agricultural Experiment Station Publication No.
D-0800 1-07-82.
Literature Cited
Cherrett, J. M. 1968. The foraging behaviour of Atta cephalotes L. (Hymenoptera; Formici-
dae). I. Foraging patterns and plant species attacked in tropical rain forest. J. Anim.
Ecol. 37:387-403.
. 1972. Chemical aspects of plant attack by leaf-cutting ants. In: J. B. Harboume (ed.).
Phytochemical Ecology. Academic Press, New York.
and J. Seaforth. 1970. Phytochemical arrestants for leaf-cutting ants, Atta cephalotes
(L.) and Acromyrmex octospinosus (Reich), with some notes on the ants’ response.
Bull. Ent. Res. 59:615-625.
Fowler, H. G. and S. W. Robinson. 1977. Foraging and grass-selection by the grass-cutting
ant, Acromyrmex landolti fracticornis (Hymenoptera: Formicidae), in habitats of intro-
duced forage grasses in Paraguay. Bull. Ent. Res. 67:659-666.
and E. W. Stiles. 1980. Conservative resource management by leaf-cutting ants? The
role of foraging territories and trails, and environmental patchiness. Sociobiology 5:25-
41.
Hubbell, S. P. and D. F. Wiemer. 1981. Host-plant selection by an attine ant. In: P. Jaisson
(ed.). Proceedings of the International Symposium on Social Insects in the Tropics (in
press).
Jaisson, P. 1980. Environmental preference induced experimentally in ants (Hymenoptera:
Formicidae). Nature 286:388-389.
Littledyke, M. and J. M. Cherrett. 1975. Variability in the selection of substrate by the leaf-
cutting ants, Atta cephalotes (L.) and Acromyrmex octospinosus (Reich) (Hymenoptera:
Formicidae). Bull. Ent. Res. 65:33-47.
and . 1978. Olfactory responses of the leaf-cutting ants, Atta cephalotes (L.)
and Acromyrmex octospinosus (Reich) (Hymenoptera: Formicidae), in the laboratory.
Bull. Ent. Res. 68:273-282.
Morrison, D. F. 1976. Multivariate Statistical Methods. McGraw-Hill, New York. 2nd Ed.
Rockwood, L. L. 1976. Plant selection and foraging patterns in two species of leaf-cutting ants
{Atta). Ecology 57:48-61.
. 1977. Foraging patterns and plant selection in Costa Rican leaf-cutting ants. J. N.Y.
Entomol. Soc. 85:222-233.
and K. E. Glander. 1979. Howling monkeys and leaf-cutting ants: comparative foraging
in a tropical deciduous forest. Biotropica 11:1-10.
Stradling, D. J. 1978. The influence of size on foraging in the ant Atta cephalotes, and the
effects of some plant defence mechanisms. J. Anim. Ecol. 47:173-188.
VOLUME XC, NUMBER 2
69
Department of Entomology & Economic Zoology, Cook College, New
Jersey Agricultural Experiment Station, Rutgers — The State University,
New Brunswick, New Jersey 08903.
Received for publication January 26, 1981.
NEW YORK ENTOMOLOGICAL SOCIETY
XC(2), 1982, pp. 70-73
A NEW SPECIES OE TRACHYMYRMEX EUNGUS-GROWING ANT
(HYMENOPTERA: MYRMICINAE: ATTINI) EROM PARAGUAY'
Harold G. Eowler
Abstract. — A new species of fungus-growing ant, Trachymyrmex kempfi,
is described from the Chaco Basin of Paraguay. This species necessitates
an expansion of the concept of the generic limits of Trachymyrmex.
Specimens of a new species of Trachymyrmex were collected in north-
western Paraguay (Chaco Basin). Correspondence with the late W. W.
Kempf indicated that this species was common in central Brazil and north-
ern Argentina. At the time of his untimely death, Dr. Kempf was in the
process of revising the species of Trachymyrmex, in which this species was
to be included. Since no generic revision is imminent, I felt it imperative to
provide a description and an available name for this common species.
Trachymyrmex kempfi, n.sp.
Figures 1-3
Holotype. — Worker holotype and 18 worker paratypes, Teniente Enciso,
Departamento Nueva Asuncion, Paraguay, 16 August, 1975 (J. Guerrero).
Holotype and 5 paratypes deposited in the Museum of Comparative Zool-
ogy. Additional paratypes are deposited in the American Museum of Natural
History and the author’s personal collection.
Diagnosis. — Similar to T. iheringi but differing in the following features:
frontal carina vestigial or weakly defined in T. iheringi, prominent in T.
kempfi', lobe of antennal scape 2.5 x longer than broad in T. kempfi, less
than this in T. iheringi ; pronounced tubercle at base of epinotal spine in T.
kempfi, absent or vestigial in T. iheringi (Fig. 2).
Description. — Holotype worker (all measurements in mm): (TL) total
body length, 4.2; (EL) maximum length of compound eye, 0.16; (OMD)
distance between lower margin of compound eye and mandibular base, 0.15;
(SL) scape length, 0.73; (WL) Weber’s length of alitrunk, 1.67; (HW) head
* Publication No. D-0800 1-06-82, New Jersey Agricultural Experiment Station, supported
by state funds.
The publication costs of this article were defrayed in part by page charge payment. This article
must therefore be hereby marked “Advertisement" in accordance with 18 U.S.C. §1734 solely
to indicate this fact.
VOLUME XC, NUMBER 2
71
Figs. 1-3. Trachymyrmex kempfi new species: 1. Paratype worker head, frontal view, 2.
Paratype worker, lateral view without appendages, 3. Paratype worker mesonotum; lateral
view.
72
NEW YORK ENTOMOLOGICAL SOCIETY
width across occiput, 1.02; (HL) head length from anterior clypeal margin
to occipital margin, 1.02; (PW) pronotum width between anterior mesonotal
crests, 0.57; (ML) mandible length, 0.49; (SI) scape index ((HW/SL)(100)),
139; (Cl) cephalic index, ((HW/HL)(100)), 100.
Head: (Fig. 1) In full frontal view, as broad as long, tubercles numerous
on frons and occiput; lobe of frontal carina expanded, covering antennal
condyle (Fig. 1); erect setae covering antennal scape, appressed fine setae
covering flagellomeres; scape fitting tightly into frontal carina; gena with
erect setae; compound eye slightly raised above gena; base of antennal
scape with a pronounced, protruding lobe; gula with appressed setae. Ali-
trunk: (Figs. 2, 3) anterior mesonotal crest 2x as long as basal width, tri-
lobed; posterior mesonotal spine poorly developed; propodeal crest poorly
developed; propodeal spine 2x as long as basal width, sharply pointed, with
large tubercle laterally at base; propodeum with numerous tubercles dorsally
and laterally; setae long and erect on propodeal spine and on dorsum of
alitrunk. Prothoracic femur: expanded. Petiole: (Fig. 2) dorsum of node
evenly convex; setae of dorsum short and erect, longitudinally long and
appressed. Postpetiole: (Fig. 2) dorsum with long, erect setae. Caster: (Fig.
2) first segment with many short tubercles arranged in four longitudinal
rows, from which arise long, semi-erect setae; venter smooth, with short,
thinner appressed hairs. Color: reddish brown.
Holotype worker bears red/handwritten label: Trachymyrmex kempfilHO-
LOTYPE/H. Fowler 1981. Paratype workers bear green/handwritten label:
Trachymyrmex kempfi/PARATYPE/H. Fowler 1981.
Paratypes.—TL, 3.34^.24; EL, 0.15-0.17; OMD, 0.15-0.17; SL, 0.73-
0.78; WL, 1.62-1.67; HW, 0.98-1.07; HL, 0.90-1.03; PW, 0.48-0.57; ML,
0.45-0.53; SI, 134-142, Cl, 97-108.
Etymology . — This species is named in honor of the late W. W. Kempf.
His cheerfulness and prompt identifications benefitted many researchers in
Latin America, and his untimely death has created a void that will not easily
be filled.
Discussion
The similarities of T. kempfi with T. iheringi are numerous, yet these
species are easily distinguished. Emery (1887) has discussed in detail these
morphological traits which distinguish T. iheringi, and which also hold for
T. kempfi.
Kempf (personal communication) indicated that T. kempfi has a broad
geographical range, extending from the Chaco Basin to Rio Grande do Sul,
Brazil, and that it is much more common than T. iheringi which tends to
occur further to the south.
Nests of T. kempfi consist of a small tumulus of excavated soil with one
VOLUME XC, NUMBER 2
73
entrance hole located on the tumulus. The workers described above were
foraging on fresh leguminous vegetation, a habit common in most range-
land Trachymyrmex, although generally more characteristic of the genera
Atta and Acromyrmex (Weber 1972).
This description of T. kempfi necessitates a new interpretation of the
genus Trachymyrmex. Although T. kempfi is weakly polymorphic, as other
species of Trachymyrmex, its large size overlaps with some castes of species
of Acromyrmex. Also T. kempfi lacks pronounced sharp spines or large
tubercles posteriorly on the head, unlike other taxa of Trachymyrmex, al-
though small spines bearing tuberculate setae are present. Also, the thoracic
and propodeal spines of T. kempfi are not well developed, which is generally
characteristic of the genus. However, the alitrunk is much smoother in
lateral view than is usually observed in Trachymyrmex. Nevertheless, T.
kempfi will key out as a Trachymyrmex in the generic key of Weber (1972).
Acknowledgments
The late W. W. Kempf initially identified the species as new, and en-
couraged further collection of the Paraguayan fauna. N. A. Weber, R. B.
Roberts, and M. DuBois all read and offered useful criticisms of the manu-
script, and their interest is gratefully acknowledged.
Literature Cited
Emery, C. 1887. Formiche della provincia di Rio Grande do Sul nel Brasile, raccolte del dott.
Herman von Ihering. Bull. Soc. Entomol. Ital. 19:352-366.
Weber, N. A. 1972. Gardening ants: the attines. Mem. Amer. Philos. Soc. 92:1-146.
Department of Entomology & Economic Zoology, New Jersey Agricul-
tural Experiment Station, Cook College, Rutgers University, New Bruns-
wick, New Jersey 08903.
Received for publication March 2, 1981.
NEW YORK ENTOMOLOGICAL SOCIETY
XC(2), 1982, pp. 74-81
EXPERIMENTAL REMOVAL OF 17- YEAR CICADA NYMPHS
AND GROWTH OF HOST APPLE TREES
Richard Karban
Abstract. — Seventeen-year cicada nymphs feed on xylem sap of most
species of deciduous trees. They attain very great densities in apple orchards
and are known to damage apple trees during oviposition. Newly hatched
nymphs were removed from certain apple trees in 1979 and the growth of
these trees was compared to trees where the nymphs had not been removed.
During 1980, trees without cicada nymphs experienced significantly in-
creased annual wood accumulation compared to control trees with cicadas.
There was no difference in growth between the 2 treatments in 1979 (the
emergence year). Despite the small sample size, these results suggest that
cicada nymphs reduce apple tree wood increment in the years following the
emergence.
Most deciduous woody tree species in the eastern U.S. serve as hosts for
cicada {Magicicada spp.) nymphs (Butler 1886; Dybas and Lloyd 1974;
Lloyd and White 1976; White 1980). Commercial apple, peach, pear, plum
and cherry trees are often heavily infested (Marlatt 1907; Asquith 1954;
Graham and Cochran 1954; Banta 1960; Hamilton and Cleveland 1964).
Adult periodical cicadas, which emerge only every 17 years at any locality
in the northern part of their range, are known to be extremely abundant
(Marlatt 1907; Dybas and Davis 1962). The adults live for two to four weeks
and females oviposit in pencil sized twigs (3-11 mm diam). The damaging
effects of egg laying are well established (Riley 1885; Hopkins 1897; Cory
and Knight 1937; Smock and Neubert 1950; Graham and Cochran 1954;
Hunter and Lund 1960; Lloyd and Dybas 1966; Smith and Linderman 1974;
White 1980).
Periodical cicadas spend their 17 or 13 year nymphal development un-
derground. As nymphs, they feed by sucking relatively great quantities of
xylem fluid, an extremely dilute source of nutrients (Cheung and Marshall
1973; White and Strehl 1978). Few studies have considered the effects of
root sucking nymphs. Assessing the impact of feeding nymphs is very dif-
ficult; trees supporting cicadas are not killed and are still capable of growth
The publication costs of this article were defrayed in part by page charge payment. This article
must therefore be hereby marked '‘Advertisement" in accordance with 18 U.S.C. §1734 solely
to indicate this fact.
VOLUME XC, NUMBER 2
75
and reproduction. Banta (1960), Hamilton (1961) and Hamilton and Cleve-
land (1964) observed that apple trees with cicada nymphs were experiencing
a cessation of growth and a reduction of yield. They were unable to reduce
chemically the cicada population, in a replicated manner, required to test
their hypothesis that cicadas were responsible for the “apple orchard de-
cline.” Karban (1980) compared the growth of parasitized and unparasitized
scrub oak trees {Querciis ilicifolia Wang.). Parasitized and unparasitized
trees did not differ in the amounts that they grew during the years preceding
the cicada emergence, indicating that microenvironmental differences be-
tween parasitized and unparasitized trees were not causing differences in
growth. In the emergence year and the years following the emergence, trees
without cicadas grew significantly more (ca. 30% more radial wood accu-
mulation) than those which supported nymphs.
The purpose of this investigation is to remove cicada nymphs from certain
trees but not others (controls) in order to test the effect of root xylem fluid
feeding on wood increment of apple tree hosts.
Methods and Site Description
A heavy emergence of M. septendecim (L.) and M. cassini (Fisher) oc-
curred in the Jenkins-Leuken orchard near New Paltz, Ulster Co., N.Y.
during the 1st week in June 1979 (Brood II). I selected eleven 'Nothern Spy’
apple trees {Malus pumila Mill.) (Fig. 1). By late June all of these trees
contained many freshly constructed eggnests. Apple trees of other varieties
meters
Eig. 1. Scale diagram of the study area. Circles with letters are ‘Northern Spy’ apple trees
that are included in the study. Circles without numbers are either a variety other than Spy or
were dead, hollow, or rotting Spy trees. The size of the circle represents the approximate
extent of the canopy of each tree. There is no apparent relationship between canopy diam and
tree growth. Trees in which cicadas were removed in 1979 are marked “N”; trees with cicadas
are marked “C.” The location of an overgrown farm road is indicated.
76
NEW YORK ENTOMOLOGICAL SOCIETY
Fig. 2. Bed sheets are set up under one of the experimental trees. As nymphs hatch they
fall from the twigs to the ground to begin their subterranean development. Sheets are used to
remove the nymphs from the experimental trees.
and trees which were dead, had fallen or had rotting or hollowed trunks
were not included. None of the eleven trees in this experiment had been
cropped in 1979 or 1980; previous cropping history is unknown.
Three of the eleven trees were selected at random and served as experi-
mental trees. Cotton bed sheets, used to entrap the newly hatched nymphs,
were spread under the canopies of the experimental trees on July 2, 1979.
The sheets were raised off the ground by wooden stakes at their corners
and rocks were placed in them to keep them from flapping in the wind and
discharging their contents (Fig. 2). The sheets allowed rain water to pass
through but caught the nymphs and unwanted debris. Nymphs were re-
moved from the debris which was then placed on the soil under the sheets.
Eight trees had no sheets placed under their canopies and serve as controls.
Once a female places her eggs in a twig they require 6 to 10 weeks to
hatch. The 1st instars hatch from their twig-borne eggs, fall to the ground
and burrow into the soil. Nymphs from the experimental trees were inter-
cepted in the sheets as they attempted to fall to the ground, leaving the
experimental trees free of cicada nymphs.
VOLUME XC, NUMBER 2
77
The density of 1st instars was estimated for the experimental trees by
placing five aluminum trays (29 x 23 x 11 cm) on the ground under the
canopy in a manner so that they were not covered by bed sheets. Each tray
was filled with a saturated picric acid solution and intercepted an area equal
to one-fifteenth of a square meter (White 1973; Karban in press). The trays
were not allowed to dry out, overflow, or be covered by spider webs. The
trays were set out on July 16 and were removed on Aug. 30 after all the
nymphs had hatched out and fallen.
The 3 experimental and 8 control trees were cored on Dec. 6, 1980 with
an increment borer. The cores were stained with a solution of 1% phloro-
glucinol in 95% ethanol. To prevent bias, an assistant who did not know
which of the cores were from the experimental trees, measured the annual
wood increment with a dissecting microscope and an ocular micrometer.
Annual wood increment was measured for a 10-year period from 1971 to
1980. Although it was possible to measure more years, the probability of
counting a false ring increases as more years are included. One of the control
cores was particularly difficult to measure and is not considered in the anal-
ysis.
Results
The radial growth increment for the 1980 growing season for each tree is
presented in Table 1. There is a 7-fold difference (0.5 to 3.5 mm) in radial
growth for 1980 among the trees, irrespective of treatment. From year to
year, some trees grow consistently more than others; this is undoubtedly
due to habitat, cropping, competitive and predator differences. Comparing
Table 1. 1980 and 1979 annual radial increments (mm) for control trees with feeding cicada
nymphs (with cicadas) and experimental trees without cicadas (no cicadas). The 1979 season,
which was the emergence year, and the 1980 season are compared to the 8-year (1971-1978)
average for each site.
8-year average
increment
(1971-1978)
1980
1980 increment
1979
increment
1979 increment
Treatment
increment
8-year average
8-year average
No cicadas
1.45
1.4
.97
1.8
1.24
No cicadas
1.91
1.8
.94
0.6
.31
No cicadas
2.88
3.5
1.22
2.0
.69
With cicadas
1.50
1.0
.67
1.2
.80
With cicadas
2.09
2.0
.96
2.3
1.10
With cicadas
2.33
1.8
.77
3.1
1.33
With cicadas
1.38
0.7
.51
0.6
.43
With cicadas
2.96
0.9
.30
2.1
.71
With cicadas
2.46
2.8
1.14
3.3
1.34
With cicadas
1.23
0.5
.41
0.7
.57
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NEW YORK ENTOMOLOGICAL SOCIETY
1980
1.5
n No Cicadas
With Cicadas
Eig. 3. The ratio of annual wood increment for 1980 and 1979 compared to the 8-year
average (1971-1978) for each tree. Trees without cicadas are shown as shaded bars, trees with
cicadas are shown as unshaded bars. In 1980, trees without cicadas were ranked 1, 3, and 5.
In 1979 trees without cicadas were ranked 3,7, and 10.
the 1980 annual wood increment with an 8-year average (1971-1978) incre-
ment for each tree provides a more meaningful estimate of the effects of the
removal of cicadas on tree growth than considering 1980 growth alone. By
comparing the 1980 growth to the 8-year average for each tree, the effects
of other factors, which each year create differences in growth between the
trees, are controlled. There is less than a 3-fold difference in average wood
increment over an 8-year period between the trees (1.23 to 2.96 mm). Table
1 and Fig. 3 present the ratio of 1980 increment to the 8-year average for
each tree. The ranks of this ratio are compared using a Mann-Whitney U
test. The null hypothesis is that there is no effect on radial wood increment
of removing the nymphs, i.e. that the samples from the experimental and
control treatments come from populations having the same distribution. This
VOLUME XC, NUMBER 2
79
null hypothesis can be rejected with 94% certainty (U 3, nj = 3, n2 == 7,
P = .058).
This same analysis was applied to the annual radial wood increment for
the 1979 growing season (the emergence year). Table 1 and Fig. 3 present
the ratio of the 1979 increment to the 8-year average for each tree. The
hypothesis that trees without cicadas added relatively more wood than those
with nymphs during 1979 is not supported by the data (U = 7, n^ = 3,
U2 = 7, F = .258).
There is no relationship between canopy diameter and tree growth. Mean
canopy diameter of the three experimental trees was 7.467 ± 1.746 m and
mean canopy diameter of the eight control trees was 6.625 ± 2.271 m (Mann-
Whitney U = 5, n^ = 3, n2 = 7, F = .133). Canopy diameter was not cor-
related with average radial growth over the 8-year period of 1971 to 1978
(Kendall rank correlation N = 2, n = 10, no significance). Canopy diameter
was not correlated with the ratio of radial growth in 1980 to the 8-year
average (Kendall rank correlation, N = 30, n = 10, no significance).
The density of first instars was estimated by catching a sample of those
falling from the tree. The estimated density and SE for each of the three
experimental trees is: 111 ± 56, 1,404 ± 144 and 1,914 ± 252 first instars
per m^. This range of densities corresponds to 20-40 adult cicadas per m^
(Karban in press) and is comparable to nymphal densities found in other
studies in apple orchards (Hamilton 1961; Forsythe 1976; Maier 1980).
Discussion
In a previous study the effects of cicada nymphs on annual wood incre-
ment were most apparent in the years immediately following the emergence
year (Karban 1980). In this study experimental removal of cicadas resulted
in increased radial growth of apple trees in the year following the emergence.
The reduction in wood increment associated with cicadas has now been
shown for apple trees as well as scrub oaks.
Experimental trees, without cicadas, added on average, 61% more radial
wood in 1980 than did control trees, with cicadas. This result was not due
to other differences between experimental and control trees. However, the
experimental trees added 12% more wood over the 8-year period (1971-
1978) than did the control trees. Scrub oaks, without cicadas added 30%
more wood, on average, than trees with cicadas (Karban 1980).
No signficant differences in annual wood accumulation during the emer-
gence year, 1979, were found between apple trees without cicadas and trees
with cicadas. This is in contrast to a previous study in which scrub oaks
with no eggnests in their canopies added significantly more wood during the
emergence year than did those with eggnests (Karban 1980). This suggests
that the negative effect on tree growth observed for the scrub oaks during
the emergence year probably resulted from oviposition damage.
80
NEW YORK ENTOMOLOGICAL SOCIETY
Periodical cicada nymphs feed on root xylem fluid (White and Strehl
1978). Amination, the process of incorporation of inorganic nitrogen into
organic compounds, takes place largely in growing root cells; most of the
nitrogen ascending the stem is already in the form of amino acids (Bollard
1957; Raven et al. 1976; Tromp and Ovaa 1976; Dickson 1979). Wiegert
(1964) points out that xylem feeding insects may be extremely costly to their
host plant because they consume the nitrogen which the plant requires for
growth.
This study has shown that 1st instar cicadas can negatively affect the
radial wood increment of apple trees. However, the small sample size, both
in terms of the number of trees in the study and in terms of the number of
years in which the effect has been found, suggest caution about generalizing
from this result.
Acknowledgments
I wish to thank Mr. Jack Leukin and Mr. Ray Jenkins for permission and
encouragement to work in their orchard. Andrea White and my father helped
in the field. The Mohonk Trust graciously provided a campsite. Leeanne
Omrod measured growth increments. This paper was writen in Chapel Hill
where JoAnn White created a ‘writers retreat,’ second to none. I benefitted
greatly from the comments of Monte Lloyd, Frank Slansky, Alan Smith,
JoAnn White and Truman Young. This work was supported by NSF grant
DEB-7914039.
Literature Cited
Asquith, D. 1954. The periodical cicada in southern Pennsylvania in 1953. J. Econ. Entomol.
47:457^59.
Banta, E. S. 1960. Apple orchard decline. Proc. Ohio Hortic. Soc. 118:88-90.
Bollard, E. G. 1957. Composition of the nitrogen fraction of apple tracheal sap. Aust. J. Biol.
Sci. 10:279-287.
Butler, A. W. 1886. The periodical cicada. Bull. USDA Div. Entomol. 12:24-31.
Cheung, W. W. K. and A. T. Marshall. 1973. Water and ion regulation in cicadas in relation
to xylem feeding. J. Insect Physiol. 19:1801-1816.
Cory, E. N. and P. Knight. 1937. Observations on brood X of the periodical cicada in Mary-
land. J. Econ. Entomol. 30:287-294.
Dickson, R. E. 1979. Xylem translocation of amino acids from roots to shoots in cottonwood
plants. Can. J. Eor. Res. 9:374-378.
Dybas, H. S. and D. D. Davis. 1962. A population census of seventeen year periodical cicadas
(Homoptera:Cicadidae:Mflg/c/cfl<7a). Ecology 43:432-444.
and M. Lloyd. 1974. The habitats of 17-year periodical cicadas. Homoptera:Cicadi-
dae: Magicicada spp. Ecol. Monogr. 44:279-324.
Eorsythe, H. Y. 1976. Estimating nymphal populations of 17-year cicadas in eastern Ohio.
Ohio J. Sci. 76:95-96.
Graham, C. and A. B. Cochran. 1954. The periodical cicada in Maryland in 1953. J. Econ.
Entomol. 47:242-244.
VOLUME XC, NUMBER 2
81
Hamilton, D. W. 1961. Periodical cicadas, Magicicada spp., as pests in apple orchards. Proc.
Indiana Acad. Sci. 71:116-121.
and M. L. Cleveland. 1964. Periodical cicadas in 1963, Brood 23. Ibid. 73:167-170.
Hopkins, A. D. 1897. The periodical cicada. W. Va. Agric. Exp. Stn. Bull. 68. 46 pp.
Hunter, P. E. and H. O. Lund. 1960. Biology of the periodical cicada in Georgia. J. Econ.
Entomol. 53:961-963.
Karban, R. 1980. Periodical cicada nymphs impose periodical oak tree wood accumulation.
Nature (London) 287:326-327.
Karban, R. in press. Increased reproductive success at high densities and predator satiation
for periodical cicadas. Ecology.
Lloyd, M. and H. S. Dybas. 1966. The periodical cicada problem. I. Population ecology.
Evolution 20:133-149.
and J. White. 1976. On the oviposition habits of 13-year versus 17-year periodical
cicadas of the same species. J. N.Y. Entomol. Soc. 84:148-155.
Maier, C. T. 1980. A mole’s eye view of seventeen-year periodical cicada nymphs, Magicicada
septendecim (Hemiptera:Homoptera:Cicadidae). Ann. Ent. Soc. Amer. 73:147-152.
Marlatt, C. L. 1907. The periodical cicada. Bull. USDA Bur. Entomol. 71:1-171.
Raven, P. H., R. E. Evert and H. Curtis. 1976. Biology of Plants. Worth Publ. New York.
685 pp.
Riley, C. V. 1885. The periodical cicada. An account of Cicada septendecim and its tredecim
race, with a chronology of all broods known. Bull. USDA Div. Entomol. 8:1-46.
Smith, E. E. and R. G. Linderman. 1974. Damage to ornamental trees and shrubs resulting
from oviposition by periodical cicadas. Environ. Entomol. 3:725-732.
Smock, R. M. and A. M. Neubert. 1950. Apples and apple products. Interscience. New York.
486 pp.
Tromp, J. and J. C. Ovaa. 1976. Effect of time of nitrogen application on amino-nitrogen
composition of roots and xylem sap of apple. Physiol. Plant. 37:29-34.
White, J. 1980. Resource partitioning by ovipositing cicadas. Am. Nat. 115:1-28.
and C. Strehl. 1978. Xylem feeding by periodical cicada nymphs on tree roots. Ecol.
Entomol. 3:323-327.
Wiegert, R. G. 1964. Population energetics of meadow spittlebugs {Philaenus spumarius) as
affected by migration and habitat. Ecol. Monogr. 34:217-241.
Biology Department, University of Pennsylvania, Philadelphia, Pennsyl-
vania 19104.
Received for publication July 21, 1981.
NEW YORK ENTOMOLOGICAL SOCIETY
XC(2), 1982, pp. 82-86
ALTERED HATCH SEQUENCE OE MALES AND EEMALES EROM
UNCHILLED EGGS OE A ‘ NON-DIAPAUSE ’ GYPSY MOTH
STRAIN (LEPIDOPTERA: LYMANTRIIDAE)
Nancy Eike Knop, Marjorie A. Hoy and Michael E. Montgomery
Abstract. — Larvae hatching from unchilled egg masses of a “non-dia-
pause” gypsy moth strain {Lymantria dispar Linnaeus) were reared to
determine the hatch sequence of males and females. Males tended to hatch
from non-diapause egg masses before females, which is the reverse of the
hatching sequence of chilled wild egg masses. These results were obtained
with an outcrossed non-diapause strain as well. The reversed hatch se-
quence is not due to a skewed sex ratio or to differential mortality.
The gypsy moth {Lymantria dispar Linnaeus) has been intensively stud-
ied in North America even though it has been a difficult laboratory animal.
Development of an artificial diet (Leonard and Doane 1966; ODell and Rol-
linson 1966) and a “non-diapause” strain (Hoy 1977, 1978a, b) have made
laboratory rearing easier. The non-diapause strain facilitates continuous lab-
oratory rearing for parasite or virus production since the gypsy moth eggs
will hatch within 31 days of deposition without exposure to cold. Wild type
gypsy moth eggs require 90-120 days of chill to obtain a good level of hatch.
The non-diapause strain had been selected simultaneously for early hatch
and hatch without chilling (Hoy 1977, 1978a, b). Beginning with generation
5, the first 60% of larvae to hatch from each egg mass were reared and
larvae hatching later were discarded. In generation 6 the first 40% of larvae
to hatch were reared, and in generation 7 only the first 20% to hatch were
reared. At the same time, the sex ratio of the adults reared from these
fractions of egg masses shifted; progressively higher frequencies of males
were produced in each succeeding generation. This study was undertaken
to determine whether the excessive number of males produced was due to
a change in sex ratio in the non-diapause strain or to a change in the time
of hatch of males and females in this strain. Accordingly, all larvae hatching
from twenty-three unchilled non-diapause egg masses were reared.
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VOLUME XC, NUMBER 2
83
Materials and Methods
Intact, unsterilized egg masses were held individually in plastic petri dish-
es (100 X 15 mm). They were supplied with a block of synthetic diet (Media
722A, BioServ, Inc., Frenchtown, N.J.). Larvae began hatching about 30
days after egg deposition. Newly hatched larvae were collected daily from
10 non-diapause (selection generation 8 (Hoy 1977, 1978a)) egg masses.
These larvae were reared in groups of 20 in 475 ml unwaxed cardboard
containers supplied with diet in small cups. Egg masses and larvae were
maintained at 21-22°C and 16 h light. Sex was determined by examining
pupae.
To see if the apparently skewed sex ratio could be modified by outbreed-
ing with a wild gypsy moth stock, an outcross was made of the selected
non-diapause strain to adult moths reared from field-collected wild eggs.
This was followed by a backcross of the progeny to the non-diapause
strain. The resulting strain was selected for “non-diapause” for 3 genera-
tions. Larvae from 13 egg masses of the 4th outcrossed non-diapause gen-
eration were group reared as above except larvae from 4 of the 13 egg
masses were reared individually in 100 x 15 mm petri dishes to reduce
larval mortality due to disease.
Results and Discussion
All three groups of egg masses had the same male-female hatch sequence
(Fig. 1). High frequencies of male pupae were reared from the first collec-
tions and progressively lower proportions were reared from subsequent col-
lections (Fig. 1). This hatch sequence is the reverse of that reported for
chilled wild gypsy moth eggs. Leonard (1968) reported that the first larvae
to hatch from field collected eggs consisted of ca. 30% males while the last
larvae to hatch were ca. 65% males. We observed this wild-type hatching
sequence repeatedly in our own laboratory (Hoy and Knop, unpubl. data).
Mortality data are given in Fig. 2.
Greater susceptibility to mortality factors of females in first collections
and males in later collections might produce a pupal (male-female) sequence
such as we observed. Our data for the group with the lowest mortality
(individually reared outcross non-diapause. Fig. 1, Fig. 2C) show that the
male-female pupal sequence cannot be eliminated even if we include the
dead larvae in the sex ratio calculations. Even if we assume that all the
dead larvae in collections 1-5 were female and all the dead larvae in collec-
tions 10-14 (no. pupae > 10, Fig. 1) were male, collections 1-5 are more
than 65% male and collections 10-14 are less than 40% male. Also, the
number of male pupae in the first collection is significantly greater (x^,
P < .001) than the number of female pupae and dead larvae combined in
84
NEW YORK ENTOMOLOGICAL SOCIETY
Eig. 1. Percentage male gypsy moth pupae reared from collections of larvae hatching from
non-diapause (selected generation 8) and outcrossed non-diapause egg masses, group reared
and individually reared. Numbers of pupae are indicated at each point.
each of the other groups (Fig. 1, Fig. 2A, B). We conclude that a male-
female hatch sequence occurs in these non-diapause strains.
The overall sex ratios of the non-diapause and the outcrossed non-dia-
pause strains are not different from the wild strain. Higher proportions of
males (58-59%) were produced in group rearing but this was the result of
higher mortality among late-hatching (mostly female) larvae (Fig. 2A, B).
Individual rearing eliminated this skewed distribution of mortality (Fig. 2C)
and pupae were 52% male.
Reversal in male-female hatching in the non-diapause gypsy moth strains
is not due to a change in sex ratio or to differential mortality. We do not
know if the reversal is a pleiotropic effect of the non-diapause trait, or is
correlated with it through linkage or through correlated selection. Hoy
(1978b) speculated that the “non-diapause” strain still has a short diapause,
the selection having influenced diapause duration and the requirement for
chilling. Wild egg masses that are chilled for less than the optimal time
produce males first (Montgomery, unpubl. data). The occurrence and du-
ration of chill may be important in determining hatch order in wild and
“non-diapause” strains. (Chilled non-diapause egg masses hatch rapidly
(Hoy 1978a) and any alteration in hatch order is obscured.)
The non-diapause gypsy moth strain was developed to facilitate contin-
VOLUME XC, NUMBER 2
85
90
80
70
60
50
40
30
20
10
0
40
30
20
10
0
40
30
20
10
0
2 4 6 8 10 12 14 16 18 +
HATCH DAY
Fig. 2. Percentage developmental mortality in collections of gypsy moth larvae hatching
from group reared non-diapause (selected generation 8) (A), and outcrossed non-diapause egg
masses either group reared (B) or individually reared (C). The broken line indicates overall
percent mortality.
uous rearing of gypsy moth parasites. The data presented here should fa-
cilitate maintenance of a desired sex ratio in larvae hatching from unchilled
egg masses of this non-diapause strain. A higher proportion of adult females
will result if the first non-diapause larvae to hatch are discarded. This would
be advantageous if the caterpillars were being reared to yield nuclear-poly-
86
NEW YORK ENTOMOLOGICAL SOCIETY
hedrosis virus since a higher yield is obtained from females than from males.
Conversely, if sterile males were being reared for a genetic control program,
it would be most profitable to rear only the first 30% of the larvae to hatch.
Acknowledgments
We wish to thank Karen Ocorr, Marion Winslow and Arthur Bridgeforth
for their assistance during the course of this project.
This work was supported in part by U.S. Department of Agriculture spon-
sored program entitled ‘The Expanded Gypsy Moth Research and Appli-
cations Program.”
Literature Cited
Hoy, M. A. 1977. Rapid response to selection for a non-diapausing gypsy moth. Science
196:1462-1463.
. 1978a. Selection for a non-diapausing gypsy moth: Some biological attributes of a new
laboratory strain. Ann. Entomol. Soc. Amer. 71:75-80.
. 1978b. Variability in diapause attributes of insects and mites: Some evolutionary and
practical implications. In: Evolution of Insect Migration and Diapause, H. Dingle (Ed.)
p. 101-126, Springer- Verlag, New York.
Leonard, D. E. 1968. Sexual differential in time of hatch of eggs of the gypsy moth. J. Econ.
Entomol. 61:698-700.
and C. C. Doane. 1966. An artificial diet for the gypsy moth, Porthetria dispar. Ann.
Entomol. Soc. Amer. 59:462^64.
Masaki, S. 1956. The effect of temperature on the termination of diapause in the eggs of
Lymantria dispar Linne. Jap. J. Appl. Zool. 21:148-157.
ODell, T. M. and W. D. Rollinson. 1966. A technique for rearing the gypsy moth, Porthetria
dispar (L.), on an artificial diet. J. Econ. Entomol. 59:741-742.
(NFK, MAH) Department of Entomological Sciences, University of Cal-
ifornia, Berkeley, California 94720 and (MEM) The Northeast Forest Ex-
periment Station, U.S.F.S., Hamden, Connecticut 06514.
Received for publication August 27, 1981.
NEW YORK ENTOMOLOGICAL SOCIETY
XC(2), 1982, pp. 87-93
COMPARATIVE FEEDING BEHAVIOR OF TWO APHID SPECIES:
BEAN APHID {APHIS FABAE SCOPOLI) AND PEA APHID
{ACYRTHOSIPHON PISUM (HARRIS))
(HOMOPTERA: APHIDIDAE)
Ronald P. Salyk and Daniel J. Sullivan SJ.
Abstract. — A comparative feeding site preference and competition study
was performed in the laboratory on two species of aphids, the bean aphid,
Aphis fabae Scopoli, and the pea aphid, Acyrthosiphon pisum (Harris). The
two species were raised separately and together on broad bean, Vida faba.
When raised separately, A. fabae was found about equally on leaves and
stems, and preferred leaf undersides to leaf uppersides by a 2:1 ratio. A.
pisum, however, strongly preferred leaves to stems, and leaf undersides to
leaf uppersides, both by more than 90%. But, when raised together, A.
fabae developed a noticeable change in preferences, preferring leaves by
83% and leaf undersides by 90%. No major change was seen in preferences
of A. pisum.
Introduction
The preferences in feeding sites of aphids have long been of interest to
a wide variety of entomologists. Feeding site preferences have implications
for the type of damage done to plants, the ease with which chemical and
biological controls can affect the aphids, the accuracy of standard sampling
methods, and the breeding of resistant plant varieties.
Leafage. — Preferences in leaf age were studied by Kennedy et al. (1950)
and Kennedy and Ibbotson (1950). They found that Myzus persicae (Sulzer)
and Aphis fabae Scopoli preferred young or early senescent leaves, while
avoiding mature leaves on the sugar beet and spindle tree. Similar results
were obtained by Wearing (1972) for M. persicae and Brevicoryne brassicae
(Linnaeus) on brussel sprouts. This preference, however, was stronger in
M. persicae. Kennedy and Booth (1951) later found that the reproductive
rate was also similarly affected by leaf age. Preference in M. persicae for
older leaves on potatoes was found by Taylor (1955), but he failed to find
any difference in Aphis nasturtii Kaltenbach. Differences between the two
races of M. persicae were reported by Tanaka (1957), who found the green
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must therefore be hereby marked "Advertisement” in accordance with 18 U.S.C. §1734 solely
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NEW YORK ENTOMOLOGICAL SOCIETY
race preferring younger leaves to a greater extent than the pink race, which
preferred the older.
Position on plant. — Preferences in position on the whole plant have also
been studied. Shands et al. (1954) studied 3 aphid species on potatoes:
Myzns persicae, Macrosiphum solanifolii (Ashmead), and abbreviata
(Patch). M. solanifolii had the highest percentage on the top leaves of the
3 species studied. Even on the same leaf, A. abbreviata and M. persicae
had preferences for different leaflets. Doucette (1961), while studying 3 other
species, found that Macrosiphum scoliopi (Essig) preferred undersides of
lower leaves of Easter lily, while Aphis gossypii Glover preferred edges of
undersides of top leaves. Myzns solani (Kaltenbach) preferred young leaves
and buds near the tip.
Leaf uppersides or undersides. — Whether uppersides or undersides of
leaves are chosen by aphids has also been the subject of some study. With
the aphid, Eucallipterus tiliae (Linnaeus), feeding on lime (Tilia spp.), Kidd
(1976) found that both uppersides and undersides were colonized in the
laboratory, while only undersides were colonized in the field. Dixon and
Logan (1973) reported an underside preference in Drepanosiphum plata-
noides (Schrank). Cruz and Bernardo (1971) found a similar behavior for
Aphis gossypii. Contrasting these studies, Wyatt (1965) noted that a majority
of aphids on the terminal buds of chrysanthemum were on the upper surface,
although this preference was reversed on the lower leaves. Uppersides were
also preferred by Hyalopterus prunii (Geoffroy) on peach and plum in a
study by El-Kady (1971). Four aphids on potatoes prefer undersides: Mac-
rosiphum euphorbiae Thomas, Myzus persicae. Aphis nasturtii, and Aula-
corthum solani (Kaltenbach) (Gibson 1972).
Bean aphid and pea aphid. — In direct relation to this present research.
Aphis fabae Scopoli, the bean aphid, and Acyrthosiphon pisurn (Harris),
the pea aphid, have been the subjects of a number of studies on feeding as
well.
1) Leaves vs. stem: Bond and Lowe (1975) found resistant varieties of V.
faba to be colonized by A. fabae approximately 50% on the leaf and 50%
on the stem, while susceptible varieties were more heavily colonized on the
leaves. Lowe and Taylor (1964) found that both green and red races of A.
pisurn preferred leaves to stems of V. faba, with from 86% to 98% of the
green preferring leaves, and from 91% to 98% of the red preferring leaves.
Lowe (1967), studying only those A. fabae that settled on leaf undersides,
found that they fed only on veins, ignoring lamina and margins. A. pisurn,
on undersides, were found to be on veins 93.3% of the time. On the same
plant at the same time, Tambs-Lyche and Kennedy (1958) found that A.
pisurn was more numerous on the leaves than A. fabae.
2) Leaf uppersides or undersides: In reference to the uppersides or un-
dersides of leaves, Dixon and Wratten (1971) reported that A. fabae feeds
VOLUME XC, NUMBER 2
89
on either surface of V.faba. However, this was not true in the research by
Hull (1964) who found only underside feeding on the sweet pea. A. pisum
was also an underside feeder, ranging from 93% to 98% on leaf undersides.
It should be noted, however, that a different host plant was used in the
study. A. pisum on V. faba was found to be primarily an underside feeder
by Muller et al. (1974).
In this present research, two types of positional preferences were studied
in the laboratory: leaf or stem preference, and upperside or underside pref-
erence, using both the bean aphid. Aphis fabae, and the pea aphid, Acyr-
thosiphon pisum. These preferences were measured in the aphids as they
colonized broad bean, Vida faba, separately, and as they colonized it to-
gether. Preferences and changes in preferences between the two species
raised separately and then raised together were recorded.
3) Aggregation: Aggregation has been found in both species. Kennedy
and Ibbotson (1951) found aggregation in A. fabae on spindle trees and
sugar beets, while Strong (1967) found aggregation in 1st and 2nd instar
nymphs of A. pisum. The results of the present research will be related to
these studies of aggregation.
Materials and Methods
Two species of aphids were used in this study: the bean aphid. Aphis
fabae Scopoli, and the pea aphid, Acyrthosiphon pisum (Harris). The broad
bean, Vida faba Linnaeus (Windsor variety), served as the host plant for
both aphids. Seeds were obtained from the W. Atlee Burpee Co., and ger-
minated in a mixture of Baccto potting soil and vermiculite in a ratio of 2: 1.
Prior to planting, the seeds were dusted with a powdered fungicide, Ortho-
cide, to inhibit the growth of fungi.
Two rows of 4 seeds each were planted in plastic trays and placed in a
dark cabinet for approximately 5 days to ensure germination. When the
seeds had sprouted, they were transferred to an insect cage illuminated with
continuous light by 1 Sylvania Gro-Lux 40 W bulb and 1 Westinghouse 40
W cool white fluorescent bulb. Temperature in the cage varied between 18°
and 25°C.
For the first 8 weeks of the experiment, 2 trays were selected and 6
sprouts were left in each tray through the removal of any stunted or incom-
pletely sprouted seeds. One apterous aphid was placed on each of these 6
sprouts, using a camel’s hair brush. Sprouts selected for the aphids were
approximately 5 cm in height. The gravid females were selected from stock
cultures of aphids reared on broad bean. One tray of broad bean sprouts,
therefore, had 6 gravid bean aphids, one to a plant, and the other tray 6
gravid pea aphids, one to a plant. These were placed in separate insect
cages.
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NEW YORK ENTOMOLOGICAL SOCIETY
Counts were taken on the 2nd, 5th and 7th days. Aphids were counted
and recorded in 3 categories: aphids on stems, on leaf uppersides, and on
leaf undersides. At the end of the 7th day count, the tray was discarded and
a new tray begun.
During the final 4 weeks of the experiment, the procedure was similar
except for the addition of a third tray of 6 sprouts. On each sprout of this
tray, one gravid bean aphid and one gravid pea aphid were placed. This tray
was placed in a third separate continuous light insect cage, and similar
counts were made on the 2nd, 5th and 7th days.
Results and Discussion
1) Reared separately: leaves v^. stem. — a) The bean aphid reared by itself
showed no preference in feeding site, being evenly divided between leaves
(49.4%) and stems (50.6%). These results were very similar to those of Bond
and Lowe (1975) who reported a division of 50% leaf to 50% stem.
b) Pea aphids reared by themselves were much more particular, with
95.0% preferring leaves to stems. This was parallel to the results of Lowe
and Taylor (1964), who found that the green race of the pea aphid preferred
leaves by a percentage varying from 86% to 98%. Tambs-Lyche and Ken-
nedy (1958) also reported that the pea aphid preferred leaves to a greater
extent than did the bean aphid when the two were raised separately.
2) Reared separately: leaf uppersides or undersides . — a) Those bean
aphids found on the leaf preferred the underside of the leaf (64.5%) to the
upperside (35.5%) by a 2:1 ratio. Dixon and Wratten (1971) noted feeding
on both the uppersides and undersides of leaves of Vicia faba by the bean
aphid, but did not quantify these observations. Hull (1964) found only un-
derside leaf feeding in the bean aphid on sweet pea, but these results may
not be directly comparable due to the different host plants.
b) Pea aphids were again more particular, with 93.4% of them preferring
the underside. This is also similar to the results of Hull (1964), who found
the pea aphid preferring leaf undersides on sweet pea from 93% to 98%.
Muller et al. (1974) also found the pea aphid to be primarily a leaf underside
feeder on Vicia faba.
3) Reared together: leaves V5. stem. — No previous studies have measured
differences or changes in feeding site preferences between these two species
when reared together as compared to their behavior when reared separately.
In the present study, the 2 species raised together showed some dramatic
behavioral changes for the bean aphid, but little change for the pea aphid.
a) Bean aphids reared together with pea aphids developed a much stronger
preference for leaves rather than stems: 83.1% compared to the original
49.4% {P < .0001, Chi square test).
b) Pea aphids reared together with bean aphids showed no noticeable
VOLUME XC, NUMBER 2
91
Table 1. Results of experiment on feeding site preferences.
1) Reared separately
Leaf
Stem
No.
%
No.
%
Bean aphid
677
49.4
693
50.6
Pea aphid
3,104
95.0
165
5.0
Leaf upperside
Leaf underside
No.
%
No.
%
Bean aphid
240
35.5
437
64.5
Pea aphid
205
6.6
2,899
93.4
Leaf
Stem
2) Reared together
No.
%
No.
%
Bean aphid
723
83.1
147
16.9
Pea aphid
1,250
94.3
75
5.7
Leaf upperside
Leaf underside
No.
%
No.
%
Bean aphid
70
9.7
653
90.3
Pea aphid
106
8.5
1,144
91.5
change in behavior, preferring leaves by almost the same percentage (94.3%)
as when reared alone.
4) Reared together: leaf uppersides or undersides. — a) When reared to-
gether with the pea aphid, the bean aphids on the leaves also developed a
much stronger preference for leaf undersides (90.3%) {P < .001).
b) The pea aphid preference changed only slightly, with 91.5% preferring
undersides, compared to 93.4% when reared separately. This change, al-
though small, was statistically significant {P < .01).
The results of both parts of the study are presented in Table 1. In sum-
mary, the presence of the pea aphid seemed to cause the bean aphid to
move from stems to leaves and from leaf uppersides to leaf undersides. No
major change was seen in the pea aphid preferences.
Intraspecific aggregation: A possible explanation of these results might
involve the phenomenon of aggregation. Individuals in many aphid species
have been shown to have a strong preference for locations on the host plant
in close proximity to groups of other aphids of the same species. This in-
traspecific aggregation behavior is independent of the quality of the feeding
site. That is to say, given apparently equal feeding sites, aphids often choose
one near another aphid or group of aphids. Kennedy and Ibbotson (1951)
found such aggregation in the bean aphid on spindle trees and sugar beets.
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NEW YORK ENTOMOLOGICAL SOCIETY
and it is said to be well known in other cases (Heathcote 1972). Aggregation
has also been reported in the pea aphid by Strong (1967) who found it in 1st
and 2nd instar nymphs.
Interspecific aggregation: It is conceivable that an interspecific aggrega-
tion has occurred in this research. The pea aphid showed an unchanging but
strong preference for leaves and leaf undersides throughout the study, re-
gardless of the presence or absence of the bean aphid. The bean aphid,
however, which was found to a great degree on both stems and leaf upper-
sides when reared separately, strongly preferred leaf undersides when raised
with the pea aphid. This appears to be a form of interspecific aggregation
or an enhancement of the bean aphid’s tendency to aggregate, and may be
to the bean aphid’s advantage in some way, possibly for protection. This
apparent aggregation was surprising in that an expected result of such a
combination of two species might have been a form of niche splitting. It
would have been thought that increased competition for feeding sites might
have logically led to the bean aphid’s leaving the leaf undersides to the pea
aphid and concentrating on stems and leaf uppersides for feeding sites,
where interspecific competition would be minimal. Surprisingly, the oppo-
site occurred. Aggregation for protection or some other advantage is the
suggested partial answer.
Other possible factors which could affect the feeding site preferences of
these aphids that were not studied here include the relative nutritional qual-
ity of the stems, leaf uppersides and leaf undersides, and the age of the plant
when first colonized by the aphids. These two factors of nutrition and age
will be explored in future work.
Literature Cited
Bond, D. A. and H. J. B. Lowe. 1975. Tests for resistance to Aphis fabae (Horn. Aphididae)
in field beans {Vida faba) (Leguminosae). Ann. Appl. Biol. 81:21-32.
Cruz, Y. P. and E. N. Bernardo. 1971. The biology and feeding behavior of the melon aphid.
Aphis gossypii Glover, on four host plants. Phil. Entomol. 2:155-156.
Dixon, A. F. G. and M. Logan. 1973. Leaf size and availability of space to the sycamore aphid
Drepanosiphum platanoides (Hem., Horn. Aphididae). Oikos 24:58-63.
and S. D. Wratten. 1971. Laboratory studies on aggregation, size and fecundity in the
black bean aphid. Aphis fabae Scop. Bull. Entomol. Res. 61:97-111.
Doucette, C. F. 1961. Systemic insecticides for control of aphids on field grown Easter lily.
J. Econ. Entomol. 54:595-597.
El-Kady, E. A. 1971. Feeding sites of the mealy plum aphid Hyalopterus pranii (Geoffroy)
with respect to its alternative host plants in Egypt. Bull. Soc. Entomol. Egypte 55:35-
40.
Gibson, R. W. 1972. The distribution of aphids in potato leaves in relation to vein size.
Entomol. Exp. Appl. 15:213-223.
Heathcote, G. D. 1972. Evaluating aphid populations on plants. In: Aphid Technology, H. F.
Van Emden (ed.) Academic Press, New York.
VOLUME XC, NUMBER 2
93
Hull, R. 1964. Some observations of Acyrthosiphon pisum (Harris) and Aphis fabae Scop.
colonizing sweet peas. Entomol. Exp. Appl. 7:195-204.
Kennedy, J. S. and C. O. Booth. 1951. Host alternation in Aphis fabae Scop. I. Eeeding
preferences and fecundity in relation to the age and kind of leaves. Ann. Appl. Biol.
38:25-64.
and A. Ibbotson. 1950. The distribution of aphid infestations in relation to leaf age. II.
The progress of Aphis fabae infestations on pot sugar beets. Ann. Appl. Biol. 37:690-
696.
and . 1951. Aggregation in Aphis fabae Scop. I. Aggregation on plants. Ann.
Appl. Biol. 38:65-78.
, C. O. Booth and A. Ibbotson. 1950. The distribution of aphid infestation in relation
to leaf age. I. Myzus persicae Sulz. and Aphis fabae Scop, on spindle trees and sugar
beets. Ann. Appl. Biol. 37:659-679.
Kidd, N. A. C. 1976. Aggregation in the lime aphid {Eucalliptenis tiliae L.). I. Leaf vein
selection and its effect on distribution on the leaf (Horn.). Oecologia 22:299-304.
Lowe, H. J. B. 1967. Interspecific differences in the biology of aphids (Homoptera) (Aphididae)
on leaves of Viciafaba. Entomol. Exp. Appl. 10:347-357.
and L. R. Taylor. 1964. Population parameters, wing production and behavior in red
and green Acyrthosiphon pisum (Harris) (Horn: Aphididae). Entomol. Exp. Appl. 7:287-
295.
Muller, F. P., L. Berger and O. Herrman. 1974. Experiments on the settling behavior of two
races oi Acyrthosiphon pisum (Harris) on Viciafaba L. (Leguminosae) (Horn: Aphididae).
Biol. Zentralbl. 93:675-686.
Shands, W. A., G. W. Simpson and L. B. Reed. 1954. Sub units of sample for estimating
aphid abundance on potatoes. J. Econ. Entomol. 47:1024-1027.
Strong, F. E. 1967. Aggregation behavior of pea aphids, Acyrthosiphon pisum. Entomol. Exp.
Appl. 10:463-475.
Tambs-Lyche, H. and J. S. Kennedy. 1958. Relation between growth pattern and resistance
to Aphis fabae Scop, in three varieties of field bean. Entomol. Exp. Appl. 1:223-239.
Tanaka, T. 1957. Studies on the two ecological forms of Myzus persicae Sulzer. I. Color
variations and distribution of the two colored forms on cabbage in the greenhouse. Jap.
J. Appl. Entomol. Zool. 1:88-94.
Taylor, C. E. 1955. Growth of the potato plant and aphid colonization. Ann. Appl. Biol.
43:151-156.
Wearing, C. H. 1972. Responses of Myzus persicae and Brevicoryne brassicae to leaf age and
water stress in brussel sprout plants. Entomol. Exp. Appl. 15:139-154.
Wyatt, I. J. 1965. The distribution of Myzus persicae on year round chrysanthemum. Ann.
Appl. Biol. 56:439-459.
(RPS) Adjunct Instructor, Department of Biology, School of Education,
City College of C.U.N.Y. and (DJS) Associate Professor, Department of
Biological Sciences, Fordham University, Bronx, New York 10458.
Received for publication October 14, 1981.
NEW YORK ENTOMOLOGICAL SOCIETY
XC(2), 1982, pp. 94-98
SEASONAL FLIGHT OF THE CRANBERRY GIRDLERi
DETERMINED WITH PHEROMONE TRAPS^’"
J. A. Kamm and L. M. McDonough
Abstract. — The seasonal flight of the cranberry girdler, Chrysoteuchia
topiaria (Zeller) was monitored with pheromone traps for 3 years. Daily
trap captures averaged at 5-day intervals were a better indicator of seasonal
flight than single-day catches. Peak flight occurred at different calendar
dates but was close to the mean summation of heat units (857°C degree-
days) for 3 years. More than 90 percent of the flight period occurred within
30-35 calendar days. Even though pheromone traps measured both flight
activity and population density, a combination of pheromone trap data and
cumulative heat units provided a reasonable estimate of the seasonal abun-
dance of adults, excluding data obtained during periods of cool rainy
weather.
The cranberry girdler, Chiysoteuchia topiaria (Zeller) occurs throughout
North America in habitats that vary from coastal cranberry bogs to high
mountain parks. The sex pheromone of this polyphagous feeder has provid-
ed a new way to monitor the seasonal flight of adults (McDonough and
Kamm 1979; Kamm and McDonough 1980). The extent to which trap cap-
tures reflect population density is unknown because various factors influ-
ence the number of insects captured in pheromone traps, e.g. weather,
population density, and various behavioral modes of the insect (Lingren et
al. 1981). Unlike many moths, the cranberry girdler is diurnal and weather
conditions during the day also influence the number responding to phero-
mone traps.
The accumulation of heat units (degree-days) is also useful in monitoring
insect populations (Sevacherian et al. 1977). Since temperature is known to
have a major influence on diapause development of the cranberry girdler
(Kamm 1973), heat unit summations may also have value in monitoring
* Lepidoptera: Pyralidae.
“ Contribution of Agric. Res. Serv., USDA in cooperation with the Agric. Exp. Sta., Oregon
State Univ. Technical Paper No. 5744 of the latter.
Mention of a commercial or proprietary product in the paper does not constitute an en-
dorsement of this product by the USDA.
The publication costs of this article were defrayed in part by page charge payment. This article
must therefore be hereby marked "Advertisement" in accordance with 18 U.S.C. §1734 solely
to indicate this fact.
VOLUME XC, NUMBER 2
95
girdler populations. The present paper describes the seasonal flight period
of the cranberry girdler determined with pheromone traps in relation to daily
weather and heat unit summations.
Materials and Methods
Tests were conducted near Corvallis, Albany, and Woodburn, Oregon,
in commercial seed fields of Kentucky bluegrass or orchardgrass infested
with a natural population of the cranberry girdler. Procedures and methods
for preparing the test baits are described elsewhere (Kamm and McDonough
1980). Pherocon 1C traps were baited with natural rubber septa that con-
tained 1.0 mg (Z)-l 1-hexadecenal and 0.05 mg (Z)-9-hexadecenal.
The flight period of adults was monitored with 6 pheromone traps de-
ployed throughout each field. Traps were serviced daily in the weather
study, and trap catches for 5 consecutive days were averaged each calendar
day of the flight season to obtain a moving average. Otherwise traps were
serviced 2 or 3 times weekly, and trap captures were averaged every 5
calendar days during the flight season. Light intensity was measured at 0900
daily with a General Electric light meter, and 3 classifications of sky con-
ditions were established: (1) Cloudy — total cloud cover, 26-28,000 foot can-
dles (fc); (2) Partially cloudy — 39-50,000 fc; and (3) Sunny — 66-67,500 fc.
Since the calling period of females occurs primarily between 0700-1100
(Kamm 1974), sky conditions during the afternoon were disregarded. No
light intensity values fell between established classifications.
Temperature data were obtained from the Oregon State University weath-
er station. Wintering prepupae of the girdler were capable of some devel-
opment at 10°C, and 5.5°C was considered the developmental threshold
(Kamm 1973). Daily heat units (degree-days) were computed by adding the
maximum and minimum temperature, then dividing by 2 and subtracting
5.5°C.
Results and Discussion
Daily weather and trap capture of males during the flight period is shown
in Fig. 1. Substantial day-to-day variation in trap capture occurred through-
out the flight period that could be attributed to weather. For example, trap
captures during rainy periods were considerably less than captures made
during favorable weather that preceded the rain. Notice also that trap catch
was reduced on cloudy or partially cloudy days when preceded by a sunny
day. Weather clearly exerted a dominant influence on trap catch, and there-
fore trap catch on any given day may not provide a reasonable estimate of
population density. Such variation in trap catches is believed to be the rule
rather than the exception.
MEAN CAPTURE OF MALES PER TRAP
96
NEW YORK ENTOMOLOGICAL SOCIETY
Eig. 1. Weather records and pheromone trap catch of male cranberry girdler in a commer-
cial seed field of bluegrass.
30
June July
Eig. 2. Eive-day moving average of seasonal pheromone trap catch of male cranberry gir-
dler.
VOLUME XC, NUMBER 2
97
Eig. 3. Seasonal flight of the cranberry girdler in 4 fields in different years and locations.
Daily variation in trap data (Fig. 1) can be reduced by computing a 5-day
moving average (Fig. 2). The resulting histogram closely approximates sea-
sonal abundance and duration of the flight period determined with light traps
(Crawford and Harwood 1964). Even though pheromone traps measured
both flight activity and population density, the traps provided a reasonable
estimate of seasonal abundance except when trap catches decreased during
the two periods of cool rainy weather (near June 24 and July 3).
Fig. 3 shows the adult trap captures in 4 fields in different years and
locations when trap captures are averaged every 5 calendar days. Notice
that populations peaked July 1-5 in 1979 and 1981 and July 5-10 in 1980.
The cumulative heat units since Jan. 1 for each year at peak flight were:
Date of peak
Cumulative heat units
Year
flight
°C at peak flight
1979
July 1-5
877
1980
July 5-10
847
1981
July 1-5
849
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NEW YORK ENTOMOLOGICAL SOCIETY
The mean cumulative heat units for the 3 years was 857, and peak flight
in all three years occurred remarkably close to the mean. Peak flight for
each of the three years would be close to the mean summation of 857 heat
units even though peak flight occurred on different calendar dates. Notice
also that 90 percent of the moths were captured in each field over a period
of 30-35 days. The decline of the adult population may be accelerated when
fields are windrowed for harvest by elimination of canopy shelter. However,
none of the bluegrass fields was windrowed for harvest before July 1 1 , and
in 2 of these fields the population had declined drastically before windrow-
ing.
Information about population density in advance of larval damage is dif-
ficult to obtain, and often the stand of grass is damaged before the infestation
becomes noticeable. Since pheromone traps measure both flight activity and
population density, it is nearly impossible to forecast precise population
density with 90-95 percent accuracy. Nevertheless, we feel a combination
of pheromone trap data and cumulative heat units provides a useful estimate
of population density during the flight period. For example, by June 10,
1979, 1980, and 1981, the cumulative heat units were 631, 551, and 577°C,
respectively. Therefore, the 1980 population would be expected at a later
calendar date than the 1979 and probably the 1981 population. With this
information, trap data obtained after June 10 should indicate about when
the population will peak and whether the population is sparse, moderate or
dense, excluding data gotten during cool and rainy weather.
Literature Cited
Crawford, C. S. and R. F. Harwood. 1964. Bionomics and control of insects affecting Wash-
ington grass seed fields. Wash. State Univ. Agric. Exp. Sta. Tech. Bull. 44, 25 p.
Kamm, J. A. 1973. Role of environment during dispause on the phenology of Chrysoteuchia
topiarki (LepidopteraiPyralidae). Ent. Exp. Appl. 16:407^13.
and L. M. McDonough. 1980. Synergism of the sex pheromone of the cranberry girdler.
Environ. Entomol. 9:795-797.
Lingren, P. D., J. R. Raulston, A. N. Sparks and W. W. Wolf. 1981. Insect monitoring
technology for evaluation of suppression via pheromone systems. In: Insect Suppression
with Controlled Release Pheromone Systems. G. Zweig, A. F. Kydonieus and M. Be-
roza, [Eds.], CRC Press, Inc., Boca Raton, Florida (in press).
McDonough, L. M. and J. A. Kamm. 1979. Sex pheromone of the cranberry girdler, Chrys-
oteuchia topiaria (Zeller) (Lepidoptera: Pyralidae). J. Chem. Ecol. 5:211-219.
Sevacherian, V., N. C. Toscano, R. A. VanSteenwyk, R. K. Sharma and R. R. Sanders. 1977.
Forecasting pink bollworm emergence by thermal summation. Environ. Entomol. 2:967-
970.
(JAK) Legume and Grass Seed Prod. Lab., Department of Entomology,
Oregon State University, Corvallis, Oregon 97331 and (LMM) Yakima Ag-
ricultural Research Laboratory, 3706 W. Nob Hill Blvd., Yakima, Wash-
ington 98902.
Received for publication October 23, 1981.
NEW YORK ENTOMOLOGICAL SOCIETY
XC(2), 1982, pp. 99-114
NOTES ON THE INTERACTION OE THE SKIPPER BUTTERFLY
CALPODES ETHLIUS (LEPIDOPTERA: HESPERIIDAE)
WITH ITS LARVAL HOST PLANT CANNA EDULIS
(CANNACEAE) IN MAZATLAN, STATE OF
SINALOA, MEXICO
Allen M. Young
Abstract. — The hesperiid butterfly Calpodes ethlius is very widely dis-
tributed in the subtropical and tropical regions of North, Central, and South
America, where the larval stage is a specialized herbivore of various Can-
naceae, mostly Canna. The larvae are known to be occasional defoliators
of cannas, including ornamental species. Some quantitative aspects of the
interaction between C. ethlius and several patches of an ornamental species
Canna edulis, were examined at Mazatlan, State of Sinaloa, Mexico, at the
beginning of the dry season. Defoliation from C. ethlius was very severe
during the previous wet season as indicated by aging conspicuous leaf dam-
age to most plants in all patches, lack of new vegetative growth, and an
abundance of old fecal pellets from larvae. Only a few late instar larvae and
adults were present at this time. A large number of pupae were found in the
host plant patches but more than 90% were parasitized, apparently by a
chalcid wasp. Although C. edulis blooms profusely here in the early dry
season, very few flowers and fruits were present. Presumably the larval
population of C. ethlius peaks during the latter half of the wet season fol-
lowing a period of steady increase in response to an abundant food supply.
The dry season interrupts this trend in abundance and the population drops
off. Severe attack by chalcid parasites in the late wet season intensifies this
change in population structure. The parasites cause mortality in the pupal
stage thus precluding any regulatory effects on the larval population. Such
interactions lower the ability of the infested larvae to produce flowers and
fruit. Other aspects of these interactions are discussed.
Calpodes ethlius Stoll (Hesperiidae) is a well known medium-sized brown
skipper butterfly with an extensive geographical range comprising predom-
inantly southern North America, Central America, and much of South
America (Klots 1951: MacNeill in Howe 1975). Several early studies re-
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must therefore be hereby marked "Advertisement” in accordance with 18 U.S.C. §1734 solely
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100
NEW YORK ENTOMOLOGICAL SOCIETY
vealed a life history pattern for C. ethlius in which the larvae feed on cannas
{Canna spp., Cannaceae — see Tietz 1972). Although the Cannaceae is a
tropical and subtropical family, native to the West Indies and Central Amer-
ica (Hey wood 1978), the widespread popularity of cannas as ornamentals
has been a probably significant ecological factor in accounting for the ex-
tensive range of this butterfly (MacNeill in Howe 1975). Although the life
cycle and association of C. ethlius larvae with cannas has been examined
(e.g. Cockerell 1892; Scudder 1889; Seitz 1924; Klots 1951, and several other
references) and it has been noted that the larvae are sometimes severe
defoliators (Klots 1951), there are no published studies on the quantitative
aspects of the butterfly’s interaction with the host plant. The purpose of the
present paper is to report one short-term study on the abundance of larvae
and pupae of C. ethlius in different patches of Canna edulis in Mazatlan,
State of Sinaloa, Mexico at the beginning of the dry season, and to make
a preliminary assessment of the possible ecological impact of the butterfly
on the host plant. This is not an attempt to establish broad generalizations
about the population biology of this obviously very successful species, as
to do so would require long-term studies in many regions.
Locality and Methods
The distribution and abundance of larvae and pupae, along with notes on
adults, of C. ethlius were studied in Mazatlan (18°02'N; 96°54'W), State of
Sinaloa, along the Pacific Coast of Mexico, 12-14 November 1979. This
region is best described as coastal tropical dry forest, with mostly open and
scattered scrub forest (Fig. 1) save for large tracts of experimental agricul-
tural lands for Citrus and other crops. A strong and long dry season occurs
here each year, usually from the end of October through June. A large
portion of the natural plant communities of the region go deciduous during
the dry season.
Within the last seven years, Mazatlan experienced a large expansion of
the tourist industry, resulting in the construction of several major hotels
with extensive gardens and recreational grounds containing numerous or-
namental plants, including numerous patches of Canna plants (Fig. 2). Dis-
cussions with three local gardeners employed by hotels revealed that the
patches of Canna were generally less than six years old. In the central (old)
section of the city, however, Canna has been present in scattered small
patches in homes. At the time of this study there was a total of eight
patches of Canna associated with two hotels and one small shopping center
in the tourist area. Other patches of Canna were undoubtedly present at
other hotels but these were not examined.
I counted the number of C. ethlius larvae and pupae (including empty and
parasitized pupae [Fig. 3j) in eight different patches of Canna. This was
VOLUME XC, NUMBER 2
101
Fig. 1. The open scrub forest habitat characteristic of the Mazatlan region.
done by carefully examining each plant within a patch, unfolding rolled
leaves and opening the characteristic tent-like structures made by the larvae.
The relatively large final instar larva (60-65 mm) and pupa (38^2 mm) make
it easy to locate C. ethlius on its host plants. The size distribution of larvae
was noted. With the exception of four larvae taken for rearing to confirm
species, the larvae and living pupae were left undisturbed following the
census. Samples of obviously parasitized (dead) pupae were also taken. The
number of plants in each patch was recorded, along with the heights of the
plants and numbers of flowers and fruits. An estimate of defoliation activity
by C. ethlius was obtained by counting the numbers of leaves with at least
20% surface area missing and bearing the easily-recognized defoliation pat-
tern of the larger larvae of this species. The ground beneath each patch was
also examined for the presence of fresh and old feces of the larvae. The
incidence of successfully-eclosed C. ethlius in each patch was estimated by
counting the number of empty pupal cases (Fig. 3). The pupae and pupal
cases of C. ethlius can generally be counted accurately as they are usually
protected within the individual tent-like structures made by the larvae prior
NEW YORK ENTOMOLOGICAL SOCIETY
102
Fig. 2. (A) Typical hotel garden environment in the tourist area of Mazatlan where patches
of Canna edulis (Cannaceae) are grown as ornamentals. (B) Typical patterns of leaf destruction
from larvae of Calpodes ethlius on its host plant, Canna edulis \ usually only 1-2 per leaf.
VOLUME XC, NUMBER 2
103
Fig. 3. (A) Final instar larvae of C. ethlius. Note internal structures visible through thin
cuticle and the silk netting lining the opened tent-like structure. (B) Parasitized pupa of C.
ethlius showing exit holes of apparently chalcid wasps. (C-D) Intact pupal shells following
adult eclosion; note the long proboscis case (D) characteristic of Calpodes. Healthy, parasit-
ized, and empty pupal shells are found inside the tent-like structures built by the larvae.
104
NEW YORK ENTOMOLOGICAL SOCIETY
to pupation. Most or all individuals pupate on the host plant, or within a
patch of the host plant. The host plant patches selected for study were
widely distributed with most inter-patch distances being greater than 30
meters, but with the closest distance between one particular pair being 500
meters, and between another pair, only one meter. Records were kept on
the coniditions of plants in these patches to determine whether or not there
was a pattern of plant healthiness with the abundance of C. ethlius. Other
ornamentals were examined briefly to check for signs of defoliation. A sam-
ple of healthy larvae was brought to Milwaukee for completion of the life
cycle. Various patches of flowers in the many surrounding gardens were
examined for adult C. ethlius, as this particular skipper is well known to
visit flowers (Klots 1951). Searches for adults were also made at the larval
host plant patches.
Results
The sizes of the eight Canna patches studied as host plant resources for
C. ethlius ranged from 10 to 150 mature plants. These patches (Figs. 4-6)
showed varying but noticeable levels of defoliation by larval C. ethlius (Fig.
7) with the percentage of defoliated plants (Fig. 4) ranging from 90-100%.
The percentages of defoliated leaves (Fig. 7) in patches ranged from 80-
95%. The herbivore damage on most plants in all of the patches was not
fresh. The damaged leaf edges were generally brown and dry. When cut
with a knife, leaf edges of Canna do not become brown for at least three
days and probably considerably longer. A large patch of about 100 plants
contained about 500 leaves whereas small patches, those with 10-15 plants,
contained about 100 leaves. Height range was very similar for all eight
patches, being 0.75 to 1.5 meters. The abundance of flowers (Fig. 7) was
very low: there were 0-3 flowers among the different patches. Fruits were
absent from three of the patches with the following distributions among the
patches: 26 fruits on 13 stalks, 22 fruits on 9 stalks, 8 fruits on 4 stalks. Of
the total of 56 fruits present, 50 were brown and partly dehisced with the
hard black seeds exposed; the remaining few were green. Fruits (Fig. 6)
were scarce relative to the numbers of plants in the patches. Only close to
full-grown larvae and empty pupal shells were present at this time (Fig. 3).
A total of 1 1 larvae were found, with none found in four of the patches. One
patch of about 100 plants had more than 1,000 old fecal pellets beneath
them, while another patch of only 48 plants also had more than 1,000 pellets.
Of 135 pupae found, 132 or 97% were parasitized, all by the same species
of an undetermined chalcid wasp. The parasite exit holes were the same
(Fig. 3) in all of these pupae, suggesting a single species of parasite. Only
two eclosed pupae were found, and one living pupa. No other insects were
found feeding on the cannas at this time. A total of four adult C. ethlius
VOLUME XC, NUMBER 2
105
Fig. 4. One of the patches of Canna edulis examined for defoliation by Calpodes ethlius
in Mazatlan. This patch contains 97 plants; note flower stalks above the leaf canopy.
were seen at two different flower patches on a single morning during a two-
hour observation period split between the patches. The adults exhibited
considerable home-range movements at the flower patches, darting off from
perches on leaves (Fig. 8) only to return a short time later. One other skip-
per, Quinta cannae (Herrich-Schaffer), presumably another Canna-fQQdQV,
was seen at the flower patches. Larvae or pupae of Q. cannae were not
found in the patches studied.
The larva of C. ethlius builds at tent-like structure (Fig. 7) folding over
the edge of a leaf and anchoring it in place with two widely-spaced multi-
stranded bands of silk. Owing to a very thin cuticle (Fig. 3) it is probably
very vulnerable to desiccation, especially in dry areas such as Mazatlan.
Larvae feed primarily after dusk and on the same leaves where their tents
are located (Fig. 7). In the laboratory in Milwaukee, the pupa stage lasted
11 days, with a rapid darkening within 48 hours of eclosion. Full eclosion
is also very rapid, the adult requiring only about four minutes to fully expand
the wings (Fig. 8).
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NEW YORK ENTOMOLOGICAL SOCIETY
Fig. 5. (A-B) Two other patches of cannas in Mazatlan examined for C. ethlius. Note the
considerable ‘‘thinning out” of leaves from larval feeding (B).
VOLUME XC, NUMBER 2
107
Fig. 6. (A-B) In addition to large patches of cannas, there were also two small patches
studied. Both large and small patches exhibited signs of considerable defoliation by C. ethlius
larvae.
Discussion
Peaks in adult populations of C. ethlius in the Mazatlan area, and perhaps
in other very seasonal tropical and subtropical regions, are predicted to be
synchronized with the growing (wet) season, based upon the age-structure
of the population studied here. The low abundance of larvae, pupae, and
adults indicates that the population was beyond its period of greatest abun-
dance. My discovery of predominantly old herbivore damage and old fecal
pellets indicates further that the period of intense larval abundance had
passed. Although the larvae of C. ethlius are well known to be occasional
severe defoliators of Canna (Klots 1951), the impact of the larvae on the
host plant population may be a very seasonal occurrence in regions such as
Mazatlan. The generation time of C. ethlius is probably on the order of 40
days, given the observed duration of the pupa stage and my own experience
studying butterfly life cycles in the tropics for the past 11 years. A wet
season period of about five months permits three, and possibly four, gen-
erations of C. ethlius. For many tropical plants, the wet season is the period
of maximal vegetative growth (Janzen and Schoener 1968), thus providing
an expanded resource base for many herbivorous insects inhabiting seasonal
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NEW YORK ENTOMOLOGICAL SOCIETY
Eig. 7. (A) Generally more than 90% of the leaves within a single patch of cannas were
heavily defoliated by C. ethUiis\ the contrast between a few surviving intact leaves and defo-
liated leaves is shown here. (B) Very few plants possessed the very conspicuous red flowers
even though blooming is generally high early in the dry season. (C) Generally between 20-90%
of the leaf surface area were devoured by C. ethlius larvae in each patch of cannas studied.
(D) In situ tent-like structure housing a C. ethlius larva; the larva feeds on the same leaf.
VOLUME XC, NUMBER 2
109
environments (Wolda 1978). Under such conditions, there will be strong
selection for synchronization of maximal breeding with the wet season in
herbivorous insects such as C. ethlius. Yet the presence of the tail-end of
the immature population during the very early dry season at Mazatlan in-
dicates that such synchronization is somewhat variable, with some overlap
between the seasons. The tent-building behavior of the larvae of C. ethlius
may be adaptive in permitting completion of a generation in the early phase
of the dry season in that the shelter, and after-dusk feeding, reduce the
chances of larval death from severe desiccation in the open exposed habitats
of this region. Such behavior may also preadapt this species to occupy new
environments otherwise impenetrable owing to dry conditions, thus con-
tributing to the widespread geographical distribution of the butterfly. Clearly
more extensive and long-term field studies are needed to confirm these ideas
on the survival of C. ethlius in very seasonal environments. As a basis for
such studies, a preliminary prediction from my observations in Mazatlan in
the early dry season would be that C. ethlius attains peak abundance and
reproductive activity in the latter half of the wet season, and the population
experiences a drastic decline by the beginning of the dry season. Severe
herbivore damage of Canna patches also occurs in the latter half of the wet
season. The age-structure of the C. ethlius population during this period
consists primarily of adults and larvae, particularly the first three instars.
Larvae of two or more generations are abundant at this time. By the begin-
ning of the dry season, the age-structure shifts to a reduced number of adults
and very few larvae if any at all. Whether or not adult C. ethlius breeds
during the long dry season of this region or enters into a reproductive dia-
pause is not known at this time. The observed age-structure of the popu-
lation at the beginning of the dry season suggests a very depressed abun-
dance of adults at this time.
Calpodes ethlius probably has its greatest impact on the Canna edulis
population during the latter half of the wet season, a time in which the larval
population is peaking in abundance. A good example of this impact is seen
in the fact that virtually all plants in the eight patches studied were severely
defoliated by C. ethlius. Very little new growth was present and the plants
were entering into a period of synchronized flowering and fruiting. I learned
from two different gardeners familiar with some of the patches studied that
(1) flowers are most abundant near the end of the wet season, and fruits
during the first half of the dry season, and (2) these patches show signs of
severe defoliation each year. The observed high incidence of parasitized
pupae suggests that most larvae, even those parasitized, complete the larval
stage and pupate, and such mortality therefore does little to regulate her-
bivore pressure on the host plant. It does, however, reduce recruitment of
no
NEW YORK ENTOMOLOGICAL SOCIETY
VOLUME XC, NUMBER 2
111
new adults. Whether or not earlier generations in the wet season experience
intense parasitism needs to be studied.
The point at which larvae are parasitized is another unknown factor is
this system. It is clear, however, that the last major wave of pupae in the
population were heavily parasitized and resulted in very low recruitment of
new adults. Therefore, it is most likely that seasonality molds the population
dynamics of this butterfly. The advancing wet season results in a build up
of the population and a gradual depletion of food supply so that when the
dry season arrives, food supply is even further reduced by lack of new
growth and the breeding population declines. The availability during the late
wet season or early dry season of a reduced larval population consisting of
later instars, provides a suitable resource base for a hymenopterous parasite
which in turn reduces the adult population even further. The early dry sea-
son may also be a period very favorable for the parasites thus increasing
their impact as a regulatory factor on the C. ethlius population. Many par-
asites are generalists on several species of Lepidoptera in a region, and as
some of these host species “drop out” of the herbivore community as the
wet season closes, such generalists concentrate on the few remaining
species.
Given the size and pattern of emergence holes of the parasites seen on
the C. ethlius pupae, the suspected parasite is a chalcid wasp, and perhaps
a species of Spilochalcis. The exit holes and their distribution on pupae
resembles very closely Spilochalcis parasitism of Opsiphanes tamarindi sik-
yon Fruhstorfer in Costa Rica (Young and Muyshondt 1975). Cockerell
(1892) reported high incidence of egg parasitism of C. ethlius by another
hymenopterous parasite, Trichogramma sp., and suggested that such mor-
tality regulates the abundance of this butterfly in Jamaica. With the excep-
tion of this observation and that presented here, little is known about the
parasites of C. ethlius. Whether or not the population in Mazatlan experi-
enced egg parasitism is not known, although perhaps not since larvae were
very abundant in at least the late wet season as indicated by the abundance
of pupae at that time.
When large larval populations build up on Canna patches, they will have
a severe defoliating effect, perhaps to the point of reducing leaf surface area
to the point where differentiation of flowers and fruit is impaired. Very few
Fig. 8. (A) Calpodes ethlius perched on the leaf of an ornamental plant in a garden in
Mazatlan (13 November 1979, 1100 hours). The butterfly is set for flying off as the species
exhibits considerable home-range movements involving sudden darting flights and perching
periods. (B) A fresh C. ethlius within five minutes after eclosion (in Milwaukee, 24 November
1979, 1500 hours).
NEW YORK ENTOMOLOGICAL SOCIETY 1
112
flowers and fruit were present in all eight patches studied and this scarcity
of reproductive structures on mature plants could have been due to the
interplay of severe herbivore pressure from C. ethlius, and the synchroni-
zation of flowering with another time of the year. These observations sug-
gest that herbivore damage by C. ethlius reduces the abundance of flowers
and fruits in infested patches of Canna in Mazatlan. The interaction between |
Canna edulis and C. ethlius is intensified further by the observation that j
this butterfly is a specialized herbivore of Cannaceae, and primarily Canna \
(Scudder 1892; Seitz 1924; Klots 1951; Tietz 1972; MacNeill in Howe (1975)
although the Cannaceae share a very close phylogenetic history with the
bananas (Musaceae), gingers (Zingiberaceae) and arrowroots (Marantaceae)
(Hey wood 1978), checking members (ornamental species) of these groups
in Mazatlan revealed no larvae or pupae of C. ethlius, nor any signs of
extensive herbivore damage indicative of larval C. ethlius.
Based upon the above considerations, I suggest that Klots’s original re-
mark (Klots 1951) about C. ethlius larvae that “sometimes they are abun-
dant enough to be destructive” be expanded to consider the interplay of
seasonality of climate, vegetative growth patterns of the host plants, and
impact of parasites. Another factor to be considered in long-term studies of
this interaction relates to the status of Canna species as ornamentals. A
major portion of the geographical distribution of Canna species results from
these plants being used as ornamentals in regions where wild species do not
occur. Apparently the group had its origin in Central America (Heywood
1978) thriving primarily in moist or wet forest areas. The expansion of these
plants as ornamentals into drier regions such as the Mazatlan area provides
an evolutionary opportunity for the herbivores normally associated with
wild populations to “track the environment” and penetrate new regions
where the plants have been introduced. Whether or not under such condi-
tions C. ethlius leaves behind its natural regulatory agents (predators and/
or parasites) associated with it on wild host plants needs to be determined.
Sometimes when a herbivorous insect enters into a new adaptive zone (host
plant) it leaves behind its natural enemies (Janzen 1973). If wild host plants
are scarce or absent from the region where the ornamentals are being intro-
duced, C. ethlius as a colonizing species will concentrate on infesting
patches of the ornamentals. It would not be surprising to discover that
virtually all patches of Canna edulis in the Mazatlan area are severely dam-
aged each year from such an effect. Given the climatic regime of this area,
it is unlikely that wild patches of host plant exist, and the known distribution
of this butterfly through Baja California and mainland Mexico (MacNeill
1975) is probably largely determined by the presence of ornamental cannas.
Being a very strong flier and having the habit of depositing eggs singly on
the host plant contribute to C. ethlius successfully dispersing itself among
VOLUME XC, NUMBER 2
113
patches of cannas. Depending chiefly upon (1) the intensity of seasonal
conditions affecting phenological patterns of vegetative and reproductive
growth of the host plants at a locality, (2) the relative abundance of wild
and ornamental host plant patches, and (3) the kinds of predators and par-
asites present, C. ethlius adult populations will vary in size at different times
of the year. Thus population structure and dynamics of C. ethlius will vary
considerably both spatially and temporally, to varying degrees. The impact
of this insect as a specialized herbivore on cannas is also expected to vary
considerably, even to the extent that under some conditions it is one of the
determinants influencing the spread of the host plant once the latter is es-
tablished as an ornamental in a new area.
Acknowledgments
This field research was accomplished while I was attending the 27th Trop-
ical Region Meeting of the American Society for Horticultural Sciences held
in Mazatlan, State of Sinaloa, Mexico. I thank Richard I. Smith (ICI de
Mexico) and Miguel Holle (CATIE, Turrialba, Costa Rica) for their assis-
tance with identifying the species of Canna. I thank Lee D. Miller (Allyn
Museum of Enotmology) for confirming the identification of the butterfly,
and also for assistance with some of the early references cited in this paper.
This paper is dedicated to the gardeners at the several hotels along the
northern “tourist strip” in Mazatlan.
Literature Cited
Cockerell, T. D. A. 1892. Notes on the life-history of Calpodes ethlius. Ent. News 2:78-80.
Heywood, V. H. (ed.). 1978. Flowering plants of the world. New York: Mayflower Books,
335 p.
Janzen, D. H. 1973. Comments on host-specificity of tropical herbivores and its relevance to
species richness, pp. 201-211, In Taxonomy and Ecology, V. H. Heywood (ed.), Syst.
Assoc. Spec. Vol. No. 5, New York: Academic Press, 325 p.
and T. W. Schoener. 1968. Differences in insect abundance and diversity between
wetter and drier sites during a tropical dry season. Ecology 49:96-110.
Klots, A. B. 1951. A field guide to the butterflies. Boston: Houghton Mifflin Co., 349 p.
MacNeill, C. D. 1975. Hesperiidae, pp. 423-577, In The Butterflies of North America, W. H.
Howe (ed.). New York: Doubleday, 633 p.
Seitz, A. (ed.). 1924. Macrolepidoptera of the world. Vol. 5. American Rhopalocera. Stuttgart:
A. Kernan Verlag, 1004 p.
Scudder, S. H. 1889. The butterflies of the eastern United States and Canada. Vol. II. Ly-
caenidae, Papilionidae, Hesperidae. Cambridge: S. H. Scudder Pub., 1774 p.
Tietz, H. M. 1972. Index to life histories. North American Macrolepidoptera. Sarasota, Flor-
ida: Allyn Museum & Ent. Reprint Specialists, 1041 p.
Wolda, H. 1978. Seasonal fluctuations in rainfall, food and abundance of tropical insects. J.
Anim. Ecol. 47:369-381.
114
NEW YORK ENTOMOLOGICAL SOCIETY
Young, A. M. and A. Muyshondt. 1975. Studies on the natural history of Central American
butterflies in the family cluster Satyridae-Brassolidae-Morphidae (LepidopteraiNymphal-
oidea). III. Opsiphanes tamarindi and Opsiphanes cassina in Costa Rica and El Salva-
dor. Stud. Neotrop. Eauna 10:19-56.
Department of Invertebrate Zoology, Milwaukee Public Museum, Mil-
waukee, Wisconsin 53233.
Received for publication October 23, 1981.
NEW YORK ENTOMOLOGICAL SOCIETY
XC(2), 1982, p. 114
BOOK REVIEW
Butterflies of the Rocky Mountain States. Clifford D. Ferris and F. Martin
Brown, eds. U. Oklahoma Press. 442 p. 4 color plates. 1981. $35 hard-
bound; $15.95 softbound.
This book is a comprehensive and updated field guide that will be most
welcomed by butterfly collectors who could no longer obtain the out-of-
print Colorado Butterflies by Brown, Eff, and Roger (1957). The prefactory
chapters describe the biogeography and geology of the region from the Ca-
nadian border to northern New Mexico, as well as some butterflies from the
Black Hills and the Pine Ridge Escarpment of South Dakota and Nebraska.
All of the species and most of the butterfly subspecies of North America in
the four major families, Hesperioidea, Papilionoidea, Lycaenoidea, and
Nymphaloidea are illustrated by black-and-white photographs and many are
shown in the 4 color plates. Life histories, flight periods, habitats, as well
as common names are included. There is a complete glossary of terms, a
general bibliography, and distribution maps. The excellent illustrations and
well-organized text add to the usefulness of this book. This very reasonably
priced field guide will serve not only the serious collector, but also a wide
audience, including weekend naturalists and young butterfly collectors.
Karl Maramorosch, Waksman Institute of Microbiology, Rutgers Uni-
versity.
NEW YORK ENTOMOLOGICAL SOCIETY
XC(2), 1982, pp. 115-116
OVIPOSITION BY TWO HELICON lUS SPECIES:
COMMENTS ON A PAPER BY DR. A. YOUNG
L. E. Gilbert
Abstract. — Corrections and comments are made concerning the paper by
A. Young on Heliconius cydno and H. sapho and their host plants.
I disagree with a number of observations and with the taxonomy in
Young’s paper on host-plant relations of Heliconius cydno and sapho (N.Y.
Entomol. Soc. 88, 1980, pp. 217-227). Young places H. sapho in the “mW-
pomene" group with H. cydno. However, the paper cited by Young (Benson
et al. 1976) clearly places H. sapho with H. hewitsoni, H. eleuchia, and H.
congener. These species are more closely allied with the ''erato" group as
they share the trait of pupal mating (Gilbert 1976). The ''sapho group” is
also known to utilize only the Astrophea subgenus of Passiflora (Benson
et al. 1976), one of which, P. pittieri, is widespread in Costa Rican rainfor-
ests, from low to middle elevation.
I have cultured H. cydno, H. sapho and H. hewitsoni in tropical insec-
taries here in Austin and my students and I have observed these and other
species oviposit in the field. H. cydno is a generalist (Smiley 1978); ovipos-
iting on most available Passiflora in forest understory and in light gaps. It
typically lays single eggs but will place two eggs on a shoot when shoots
are limited (see Williams and Gilbert 1981). On the other hand, Atlantic side
H. sapho and its close relative on the Pacific side of Costa Rica, H. hew-
itsoni, are strictly monophagous on P. pittieri. This plant has a tree-like
growth form but can also be a liana. It grows in spurts punctuated by total
absence of new growth. H. sapho and H. hewitsoni always place clusters
on young shoots. In fact, females of these species are very choosy, using
only a restricted stage of shoot development. (Beyond a certain point, young
larvae would be unable to feed on maturing leaves which are extremely
tough in this Passiflora species.)
In Costa Rica H. cydno, a mullerian mimic of H. sapho and its close
relative, H. pachinus, a mullerian mimic of H. hewitsoni, both occasionally
oviposit on P. pittieri but have a lower per capita survivorship on those
The publication costs of this article were defrayed in part by page charge payment. This article
must therefore be hereby marked ''Advertisement" in accordance with 18 U.S.C. §1734 solely
to indicate this fact.
1 16 NEW YORK ENTOMOLOGICAL SOCIETY |
shoots than the clusterlaying specialist (J. Longino, personal communica-
tion).
Young’s Passiflora identifications require corrections. The individual la-
belled ''Granadilla A” (fig. IB) is unmistakably Passiflora pittieri Masters,
an Astrophea. The other '"Granadilla A” (fig. 2A, B) is equally unmistakably
Passiflora ambigua Hemsl., a common liana in Costa Rican rainforests. P.
ambigua is a member of Granadilla and is a recorded host for both H. cydno |
and H. pachinus in Costa Rica (Benson et al. 1976). Both P. pittieri and P. !
ambigua lack tendrils as young understory plants and develop tendrils as |
mature vines. In rearing both of these Passiflora from seedlings, I have
noticed several consistent features which allow identification. The best char-
acter for young plants is the placement of extra floral nectaries. There are
typically Vi to Vs of the way along the petiole toward the leaf base in P.
ambigua as can be clearly seen in Young’s fig. 2A, B. Extra floral nectaries
of P. pittieri are conspicuous at the leaf base in unfolded new leaves
(Young’s fig. 4C) but hidden under the leaf at the apex of the petiole in
older leaves (Young’s fig. IB, fig. 3A).
An additional problem concerns the specific identification of these mi-
metic Heliconius. The black and white butterfly depositing “large clusters”
on P. pittieri is H. sapho, not H. cydno. For example, fig. 4A, p. 223 show
H. sapho, not H. cydno. The shape of the white fore-wing patch, clearly
visible in Young’s fig. 4A, is diagnostic of H. sapho.
Literature Cited
Benson, W. W., K. S. Brown and L. E. Gilbert. 1976. Coevolution of plants and herbivores.
Evolution 29:659-680.
Gilbert, L. E. 1976. Postmating female odor in Heliconius butterflies: A male contributed
antiaphrodisiac? Science 173:419-420.
. 1979. Attempts to build theory in the study of insect plant interactions. In: “Analysis
of ecological systems,” Horn, Mitchell and Stearns, eds.; Ohio State Univ. press, Co-
lumbus.
Smiley, J. T. 1978. Plant chemistry and the evolution of host specificity: New evidence from
Heliconius and Passiflora. Science 201:745-747.
Williams, K. S. and L. E. Gilbert. 1981. Insects as selective agents on plant vegetative mor-
phology: egg mimicry reduces egg-laying by butterflies. Science 212:467-469.
Department of Zoology, University of Texas, Austin, Texas 78712.
Received for publication October 27, 1981.
NEW YORK ENTOMOLOGICAL SOCIETY
XC(2), 1982, pp. 117-118
ERRATA: OVER-EXPLOITATION OF LARVAL HOST
PLANTS BY HELICONIUS BUTTERFLIES
Allen M. Young
I take this opportunity to agree with, and to thank, both Drs. Keith S.
Brown, Jr. and Lawrence E. Gilbert for expressing their concern for the
errors I made in the paper discussed by Gilbert (1982). My errors were
initially pointed out to me by a letter from Dr. Brown, and subsequently I
received a copy of an earlier version of Dr. Gilbert’s manuscript. In this
note I wish to explain the sources of my errors appearing in Young (1980).
My error in confusing Passiflora pit fieri for a species of Granadilla, as
pointed out by both Drs. Brown and Gilbert, arose from a superficial com-
parison of a photograph with the plant shown in figs. 1-3 (Young 1980),
which was determined by Brown (pers. comm., 30 April 1977) to be Gran-
adilla. Gilbert (1982) disagrees with this determination. My error was com-
pounded by not having a voucher specimen for determination. I hesitated
to collect the individual plant in question because I wanted to observe pos-
sible repeated Heliconius oviposition on both plants over a year or longer.
Neither plant was therefore collected. Granadilla and P. pittieri are system-
atically far apart from one another in the evolution of the Passifloraceae
(Benson et al. 1976). Furthermore, Laurifoliae is a series, not a subfamily
(Benson et al. 1976).
My misidentification of Heliconius sapho as H. cydno in fig. 4 (Young
1980) came from missing the clearly diagnostic small red patch at the base
of the hindwing, very visible in the ovipositing butterfly and in the photo-
graph (fig. 4) as a small light area. The butterfly had not been collected to
make a confirmative determination. At the time it did not occur to me that
the shape of the white fore- wing patch (seen in my fig. 4A) is also diagnostic
of H. sapho as pointed out by Dr. Gilbert in the accompanying note. I
simply did not know this. Heliconius sapho is not in the melpomene group
(Benson et al. 1976) as I incorrectly stated in my paper.
Members of the H. sapho group customarily lay large numbers of eggs
on individual host plants (Benson et al. 1976; Brown 1981), thereby explain-
ing the high abundance of eggs I reported (Young 1980). Heliconius sapho
typically lays 10-40 eggs on a rapidly growing meristem (Brown 1981). Dr.
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NEW YORK ENTOMOLOGICAL SOCIETY
Brown (pers. comm.) believes that the reproductive strategy of members of
the H. sapho group includes the possibility of sometimes laying many more
eggs on an individual host plant than can survive, so it is sometimes ob-
served that only one or two larvae make it to pupation. This may be related
to the frequent “explosive” mode of growth of young shoots on the Astro-
phea host plants (Benson et al. 1976). Thus one may assume that the high
density of eggs observed on the second host plant in my study is typical
oviposition behavior of H. sapho, and that the butterfly exhibits the same
behavior on large host plant individuals as well as on small ones which are
expected to grow rapidly.
Gilbert, in the accompanying note, has comprehensively summarized de-
tailed information on the oviposition behavior of both H. cydno and H.
sapho in relation to larval host plants, and has clarified how to distinguish
the larval host plants in my study.
In the study discussed here, I was not able to follow two standard pro-
cedures I use for confirming the identifications of host plant and butterfly
species used in my other published field studies in Costa Rica. I did not, in
Young (1980), collect the butterfly or host plant material, thereby increasing
markedly the potential for my margin of error.
In closing, I am appreciative of the professional concerns expressed by
Drs. Brown and Gilbert in relation to the errors appearing in my paper
(Young 1980). I thank the editor of this journal for giving me the opportunity
to clarify the nature of these unfortunate mistakes. I offer particular thanks
also to Dr. Keith S. Brown, Jr. for his encouragement and assistance with
clarifying these errors, and for reading over an earlier draft of this note.
Literature Cited
Benson, W. W., K. S. Brown, Jr. and L. E. Gilbert. 1976. Coevolution of plants and herbi-
vores; passion flower butterflies. Evolution 29:659-680.
Brown, K. S., Jr. 1981. The biology of Heliconius and related genera. Ann. Rev. Entomol.
26:427-456.
Gilbert, L. E. 1982. Oviposition by two Heliconius species: comments on a paper by Dr. A.
Young. J. New York Entomol. Soc. 90;in press.
Young, A. M. 1980. Over-exploitation of larval host plants by the butterflies Heliconius cydno
and Heliconius sapho (Lepidoptera: NymphaUdae: Heliconiinae: Heliconiini) in Costa
Rica? J. New York Entomol. Soc. 88:217-227.
Department of Invertebrate Zoology, Milwaukee Public Museum, Mil-
waukee, Wisconsin 53233.
Received for publication December 15, 1981.
NEW YORK ENTOMOLOGICAL SOCIETY
XC(2), 1982, pp. 119-124
MIRIDAE AND COLEOPTERA ASSOCIATED WITH
TULIP TREE FLOWERS AT ITHACA, NEW YORK
David Andow
Abstract. — The Miridae and Coleoptera on tulip tree (Liriodendron tu-
lipfera L.) flowers were examined in June, 1979 at Ithaca, New York. The
mirids are mainly Lygocoris caryae and L. omnivagus, and are probably
attracted to tulip tree to feed on the flowers. The mirid-flower association
changes with the age of the flower; young flowers support a larger mirid
fauna than older flowers. The Coleoptera are mainly Cantharidae and Ceph-
aloidae. Notes on the feeding behavior of these insects are recorded. These
are the first feeding records for Cephaloon lepturides, Cantharis scitulus,
C. rectus, and Podabrus brunnicollis, and the first records for Lygocoris
caryae, L. omnivagus, L. tiliae, L. hirticulus, and L. belfragii on tulip tree.
The tulip tree or yellow poplar, Liriodendron tulipifera L. is an important
hardwood in the Appalachian region. It is distributed from central Massa-
chusetts to Ontario and Wisconsin, and southwest to Florida, Mississippi,
and Arkansas (Wiegand and Fames 1925), and is relatively common in rich,
light, loamy woodland soils. Its northern limit passes through upper New
York State.
More than 100 species of insects have been reported to feed on the leaves
and stems of tulip tree (Felt 1905; Felt and Rankin 1932; Herrick 1935;
Doane et al. 1936; Burns and Gibson 1968; Burns 1970; Peigler 1976; Fu-
tuyma and Gould 1979). The pollinators of L. tulipifera are not known but
may be similar to those on Magnolia acuminata L. (Downes 1973; Thien
1974; Thien et al. 1975).
Flowers are an important resource for insects. Current research on pol-
lination ecology has focussed on competition between pollinators for flower
products and between plants for pollinators (Feinsinger 1976; Heinrich 1976;
Rathcke and Poole 1977; Poole and Rathcke 1979). However, parasitoids,
predators, and herbivores also consume flowers and flower products. Evans
and Murdoch (1968) found that 68 percent of the adult insect fauna of an
old field in Michigan were flower feeders. The insects utilizing many species
of flowers have been described (Kerner 1878; Robertson 1928); Wheeler and
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NEW YORK ENTOMOLOGICAL SOCIETY
Henry (1976) present observations on five species of mirids that feed on
honey locust flowers, and Henry (pers. comm.) has observed both nymphs
and adults of Neurocolpus nubilus feeding on and damaging petals of tulip
tree in Pennsylvania. This paper describes the associations of flower feeding
mirids and Coleoptera on Liriodendron tulipifera flowers at Ithaca, New
York.
Materials and Methods
Two individuals of Liriodendron tulipifera on Turkey Hill in Ithaca, New
York were observed on the edge of a weedy lawn, surrounded by poplar,
yellow birch, white pine, white ash, basswood, hickory, and oaks. The trees
are about 50 feet tall, and first started blooming on June 7, 1979, continuing
through June 25.
On June 8 and 9 general collections were made from about 300 flowers.
On June 10, all the insects were removed from the open flowers on four low
branches, and haphazardly twenty of these flowers were designated as “old
flowers.” Eight “young flowers,” opening between the mornings of the 10th
and 11th, were also identified. Insects were collected from these flowers for
the next five consecutive mornings between 0600 and 0700 hours. On June
14, five more newly opened flowers were added to the young flower sample.
On June 10 at 0700 hours, twenty mirids in six flowers were marked with
fluorescent dusts sprayed into the flowers. No insects left these flowers
immediately after marking. At 0800 the following morning, all the mirids in
those six flowers were collected and identified.
Results and Discussion
A total of 55 Miridae and 47 Coleoptera were collected in 150 flower-days
of sampling (Table 1). Most of the Miridae (96 percent) were species of the
genus Lygocoris, and most of the Coleoptera were represented by the fam-
ilies Cantharidae (55 percent) and Cephaloidae (38 percent).
Miridae. — All the mirids were observed feeding in the flowers, mainly on
the lower portion of the carpels. It is not known if they damage the seed.
No nymphs of any of the species were seen on tulip tree during the period
of observation. L. tulipifera is probably only an adult feeding host for these
insects.
Lygocoris caryae breeds on hickory {Carya ovata, and C. alba), and to
a lesser extent on pecan, and walnut (Knight 1917; Blatchley 1926; Smith
1940; Kelton 1971). It has also been found on Rhus typhina, Lilia americana,
Robinia pseudoacacia, Quercus sp., dead Juniperus virginiana (Knight 1941;
Kelton 1971). It can cause economic damage on peaches and pears (Caesar
1920; Ross and Caesar 1921, 1927; Carman 1936; Smith 1940; Knight 1941),
VOLUME XC, NUMBER 2
121
Table 1. Number of insects caught in young and old L. tulipifera flowers. A single flower-
day sample consists of the insects accumulating in one flower during one 24 hour period. L.
belfragii was collected while general collecting on June 8 and 9. All insects were identified by
the author, and are deposited in the Cornell University collection under C.U. Lot #1108.
Young flowers
50 flower-days
Old flowers
100 flower-days
Miridae
37
19
Lygocoris caryae (Knight)
18
9
13.5*
L. omnivagus (Knight)
14
7
10.5*
Other mirids**
5
3
—
Cantharidae
12
14
1.92
Cantharis scitulus Say
0
3
—
C. rectus Melsheimer
9
9
2.25
Podabrus spp.f
3
2
—
Cephaloidae
Cephaloon leptu rides Newman
7
11
0.29
Other Coleopteratt
2
0
—
* P < 0.005, all others n.s.
** L. tiliae (Knight), L. hirticulus (Van Duzee), L. belfragii (Reuter), Taedia sp., and
Monosynamma bohemani (Fallen).
t P. brunnicollis Fab. and P. rugulosus LeConte.
tt Grammoptera haemetites (Newman) [Cerambycidae] and Orsodacne atra (Ahrens)
[Chrysomelidae].
migrating as far as 300 yards into orchards, returning to hickory to oviposit.
Its economic damage is often severe, but highly variable in space and time.
Lygocoris omnivagus has been reported from a large number of hosts,
breeding on oaks {Quercus alba, Q. rubra, and Q. coccinea), and to a lesser
degree on Cornus florida, C. circinata, Castanea sp., and Viburnum acer-
ifolium (Knight 1917; Blatchley 1926; Knight 1941). Kelton (1971) reports
it from numerous other species. It too can cause significant damage to
peaches and pears (Ross and Caesar 1921, 1927; Carman 1928, 1936; Kelton
1971), migrating to orchards to feed, and returning to its host plants to
oviposit.
L. tulipifera is a new feeding record for both of these species, and also
for Lygocoris tiliae, L. hirticulus, and L. belfragii. The individuals I caught
probably migrated to tulip tree from nearby host trees, where they will
return to oviposit. Many of the individuals I observed were covered with
tulip tree pollen, so they may be pollinators. But many bees and wasps were
also observed working the flowers, so the importance of mirids as pollinators
is unknown.
Coleoptera. — Little is known about the biology of the Cantharidae. Can-
tharis spp. are known to feed on aphids, mealybugs, and plant materials
122
NEW YORK ENTOMOLOGICAL SOCIETY
such as pollen and nectar (Motizambert 1908; Essig 1926; Fluke 1929;
Balduf 1935; Claussen 1940). Podabrus spp. also feed on aphids and pollen
(Webster 1876-83; Blatchley 1910; Wilson 1913; Essig 1926; Fluke 1929;
Smith 1936; Claussen 1940; Fender 1973).
The cantharids on L. tulipifera flowers were observed feeding on the
pollen and other litter that collects in the bottoms of the flowers. My ob-
servations are the first feeding records for Cantharis scitulus, C. rectus, and
Podabrus brunnicollis . Although P. rugosuliis is known to eat aphids
(Blatchley 1910; Fluke 1929), this is the first record of it eating plant ma-
terial.
The association of adult Cephaloon lepturides with L. tulipifera is the
first published ecological record for adults of this species. This is an active
beetle, more so than the smaller cantharids, and moves rapidly inside the
flowers. It is not known if these beetles eat plant material. In captivity, I
observed C. lepturides to kill and eat an adult of L. caryae and L. tiliae in
a manner similar to the cantharids (Motizambert 1908; Tilden 1950).
The insect-flower associations change with age, young flowers supporting
a different mirid fauna than old flowers (Table 1). This was also observed
in the general collections on the 8th and 9th. Lygocoris caryae and L.
omnivagus were more abundant in young flowers than old flowers. All the
beetles appeared in flowers older than 3 days. Since these beetles are pollen
feeders, they may be attracted to flowers after anther dehiscence.
The mirids may be displaced from the older flowers by the cantharids and
Cephaloon lepturides. The beetles move around inside the flowers much
more than the mirids, often bumping into them. The mirids stop feeding to
move and avoid the beetles. These interruptions may limit feeding time so
that the mirids leave the older flowers that have beetles to find a better,
more available food resource. Also, since these beetles may be potential
predators, the mirids may be avoiding predation.
Alternatively, flower quality may decline with age and the mirids are
avoiding low quality flowers. Of the marked mirids, only 55 percent stayed
in the same flower for a day. None of these flowers contained beetles, so
this shows that the mirid-flower association is ephemeral and that mirids
move enough to be able to respond rapidly to factors such as changes in
flower quality and availability.
Even in such a temporary resource as flowers there are regular changes
in herbivore presence. Be it a decline in flower quality with time or dis-
placement by beetles, the mirid fauna changes rapidly, apparently exploiting
just a portion of a flower’s life.
Acknowledgments
I would like to thank Dr. A. G. Wheeler, Jr. (Bureau of Plant Industry, Penn-
sylvania Dept, of Agriculture, Harrisburg, Pa.) for his comments on this
VOLUME XC, NUMBER 2
123
paper and for pointing out several of the references used, Mr. Stephen Behr
(Dept, of Nutrition, Cornell University, Ithaca, N.Y.) for the use of the tu-
I lip trees on his property. Dr. Thomas J. Henry (Systematic Entomology
Laboratory, % U.S. National Museum, Washington, D.C.) for his com-
ments and information on the Miridae, and an anonymous reviewer whose
comments strengthened this paper.
Literature Cited
1
Balduf, Walter V. 1935. The Bionomics of Entomophagous Coleoptera (John S. Swift Co.: St.
Louis), 220 p.
Blatchley, W. S. 1910. Coleoptera or Beetles (The Nature Publ. Co.: Indianapolis), 1386 p.
. 1926. Heteroptera or True Bugs of Eastern North America (The Nature Publ. Co.:
Indianapolis), 1116 p.
Burns, Denver P. 1970. Insect Enemies of Yellow-Poplar. USDA Forest Service Research
Paper NE-159.
and Lester P. Gibson. 1968. The leaf-mining weevil of yellow-poplar. Can. Ent.
100:421-429.
i Caesar, L. 1920. Notes on leaf bugs (Miridae) attacking fruit trees in Ontario. 51st Ann. Rpt.
Ent. Soc. Ont. 14-16.
Claussen, C. P. 1940. Entomophagous Insects (McGraw-Hill: New York), 688 p.
Doane, R. W., E. C. Van Dyke, W. J. Chamberlain and H. B. Burke. 1936. Forest Insects
(McGraw-Hill: London), 463 p.
Downes, John A. 1973. Endopterygote insects and the origin of the angiosperm flower. Inter-
national Congress of Systematic and Evolutionary Biology, 1st, Boulder, Colo., 1973.
Abstracts of papers, p. 9.
j Essig, E. O. 1926. Insects of Western North America (Macmillan: New York), 1035 p.
Feinsinger, Peter. 1976. Organization of a tropical guild of nectarivorous birds. Ecol. Monogr.
46:257-291.
Felt, Ephraim P. 1905. Insects Affecting Park and Woodland Trees. New York State Museum
Memoir 8, pt. I and II.
and W. Howard Rankin. 1932. Insects and Diseases of Ornamental Trees and Shrubs
(Macmillan: New York), 507 p.
Fender, Kenneth M. 1973. Ecological notes on Podabrus (Coleoptera: Cantharidae). Coleopt.
Bull. 27:11-17.
Fluke, C. L. 1929. The known predaceous and parasitic enemies of the pea aphid in North
America. U. Wise. Agr. Expt. Sta. Res. Bull. 93.
Futuyma, Douglas J. and Fred Gould. 1979. Associations of plants and insects in a deciduous
forest. Ecol. Monogr. 49:33-50.
Garman, Philip. 1928. Plant bug injury to fruits. Conn. Agr. Expt. Sta. Bull. 305:729-731.
. 1936. Plant bug injury to fruits. Conn. Agr. Expt. Sta. Bull. 383:357-358.
Heinrich, Bernd. 1975. Flowering phenologies: bog, woodland, and disturbed habitats. Ecol-
ogy 57:890-899.
Herrick, Glenn W. 1935. Insect Enemies of Shade Trees (Comstock: Ithaca), 417 p.
Kerner, A. 1878. Flowers and Their Unbidden Guests (C. Kegan Paul: London), 164 p.
Knight, Harry H. 1917. A revision of the genus Lygus as it occurs in America north of Mexico,
with biological data on the species from New York. Cornell Agr. Expt. Sta. Bull. 391.
. 1941. The Plant Bugs, or Miridae, of Illinois. Bull. 111. Nat. Hist. Survey 22(1).
Motizambert, Eric. 1908. Lampyridae and aphids. Can. Ent. 40:36.
Peigler, Richard S. 1976. Observations on host plant relationships and larval nutrition in Cal-
losamia (Saturniidae). J. Lepid. Soc. 30:184-186.
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Poole, Robert W. and Beverly J. Rathcke. 1979. Regularity, randomness, and aggregation in
flowering phenologies. Science 203:470^71.
Rathcke, B. J. and R. W. Poole. 1977. Community patterns of flowering phenologies. Bull.
Ecol. Soc. Amer. 58(2): 15 (Abstract).
Robertson, Charles. 1928. Flowers and Insects: Lists of Visitors of Four Hundred and Fifty-
three Flowers (Science Press: Carlinville, 111.), 221 p.
Ross, W. A. and L. Caesar. 1921. Oak plant bugs. 52nd Ann. Rpt. Ent. Soc. Ont. 44-45.
and . 1927. Oak and hickory plant bugs. 58th Ann. Rpt. Ent. Soc. Ont. 17.
Smith, Edward H. 1940. The biology and control of the hickory capsid, Lygus caryae Knight
(Hemiptera: Miridae), as a peach pest. M.S. Thesis, Cornell University.
Smith, J. W. 1936. Biology of the mealy plum aphid, Hyalopterus pruni (Geoffroy). Hilgardia
10:167-207.
Thien, Leonard B. 1974. Floral biology of magnolia. Amer. J. Bot. 61:1037-1045.
, W. H. Heimermann and R. T. Holman. 1975. Floral odors and quantitative taxonomy
of Magnolia and Liriodendron. Taxon 24:557-568.
Tilden, J. W. 1950. The feeding of Podabms pruinosus LeConte (Cantharidae). Coleopt. Bull.
4:92.
Webster, F. M. 1876-1883. Notes upon the food of predaceous beetles. Bull. 111. St. Lab. Nat.
Hist. 1, 149-152.
Wheeler, A. G., Jr. and Thomas J. Henry. 1976. Biology of the honey locust plant bug,
Diaphnocohs chlorionis, and other mirids associated with ornamental honey locust.
Ann. Amer. Ent. Soc. 69:1095-1104.
Wiegand, Karl M. and Arthur J. Fames. 1925. The Flora of the Cayuga Lake Basin, New
York: Vascular Plants. Cornell Univ. Agr. Expt. Sta. Memoir 92.
Wilson, H. F. 1913. Notes on Podabms pruinosus. J. Econ. Ent. 6:457^58.
Department of Ecology and Systematics, Cornell University, Ithaca, New
York 14853.
Received for publication October 6, 1981.
NEW YORK ENTOMOLOGICAL SOCIETY
XC(2), 1982, pp. 125-128
SUSCEPTIBILITY OF GYPSY MOTH LARVAE TO SEVERAL
SPECIES OF ENTOMOGENOUS FUNGI
S. S. Wasti and G. C. Hartmann
Abstract. — Several species of fungal entomopathogens were tested for
infectivity on larvae of the gypsy moth, Lymantria dispar (L.). Spore sus-
pensions in varying dosages were administered to the larvae via surface
treatment of sterilized oak leaves. High levels of mortality were obtained
for most of the test species of fungi. Conidiobolus thromboides, Fusarium
sp., Hirsutella thompsonii, one strain of Paecilomyces farinosus and Ver-
ticillium sp. were non-infective to gypsy moth larvae under the conditions
of the experiment.
I Introduction
The infectivity of entomogenous fungi on gypsy moth larvae has been
demonstrated on the basis of qualitative and histopathological studies (Hart-
mann and Wasti 1974, 1976; Wasti and Hartmann 1975, 1978; Wasti et al.
1980). Toxicity tests on non-target invertebrate and vertebrate hosts have
established the environmental safety of these candidate biological control
agents (Donovan-Peluso et al. 1980; Hartmann and Wasti 1976, 1980; Hart-
mann et al. 1979; Wasti et al. 1980). Quantitative studies on the effect of
specific dosage levels on mortality of gypsy moth larvae were the primary
objectives of the present study.
Methods
Larvae of the gypsy moth, Lymantria dispar, were reared from eggs sup-
plied by the Forest Insect Disease Laboratory, Hamden, Connecticut. En-
tomogenous fungal species tested for infectivity against these larvae were:
1. Beauveria bassiana (Bals.) Vuill. Russian strain
2. Beauveria bassiana (Bals.) Vuill. {B. tenella) ATCC 26156
3. Cordyceps militaris (Fries) Link ATCC 26848
4. Conidiobolus thromboides Drechsler (Entomophthora virulenta) (Hall
& Dunn) Latge et al. (1980) ATCC 14270
5. Fusarium sp. (local strain)
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NEW YORK ENTOMOLOGICAL SOCIETY
6. Hirsutella thompsonii (Fisher) ATCC 24874
7. Metarhizium anisopliae (Metsch.) Sorokin ATCC 22099
8. Paecilomyces farinosus (Dicks, ex Fr.) Vuill. ATCC 24319
9. Paecilomyces farinosus (Dicks, ex Fr.) Vuill. USDA, Hamden, Ct. strain
10. Paecilomyces fumoso-roscus (Wize) Brown & Smith ATCC 16312
11. Nomuraea rilcyi (Farlow) Samson, Missouri strain
12. Verticillium sp. (local strain)
Stock cultures of the various species of fungi were subcultured on Potato-
dextrose agar and yeast extract (PDAYE) under procedures outlined by
Hartmann and Wasti (1974). Culture isolates of the test species were then
passed in serial succession through larvae of the wax moth. Galleria mel-
lonella L. in order to increase virulence by passage through an easily in- |
oculated host.
Infectivity studies were considered initially using the surface treatment i
technique recommended by Garcia and Ignoffo (1978) for infecting larvae i
with entomogenous fungi. Spore suspensions representing calibrated dos-
ages of the fungal species were administered in sterile Petri dishes (dia. 60
mm) by spreading them over the surface of artificial gypsy moth diet under
conditions recommended by O’Dell and Rollinson (1966). Results with this
technique were unsatisfactory and the method proved unsuitable. There was
difficulty in distributing low spore concentrations uniformly over the surface
and infection rates with early instars were extremely low. Elimination of
the antifungal agents in the diet increased the likelihood of infection by non-
target saprotrophic fungi.
As an alternative, the leaflet technique described by Garcia and Ignoffo
(1978) was used for the infectivity studies and was found to provide satis-
factory results. Eresh oak leaves were surface-sterilized by immersion for
several minutes in a one per cent solution of sodium hypochlorite and thor-
oughly rinsed with sterile distilled water. Appropriate dilutions of fungal
spore suspensions were applied to the surface and each leaf was placed in
a sterile Petri dish.
Fifty second instar larvae were surface-sterilized by immersion in a so-
lution of Zephirine chloride (1:750) followed by a sterile water rinse, placed
in each Petri dish, and allowed to feed on the leaves for 48 hours. Subse-
quently, the larvae were transferred to individual plastic chambers and ob-
served at 24 hr intervals. The following spore dosages of each test organism
were administered: 50, 100, 500, 1,000, 5,000 spores/mm^. Untreated leaves
were set up as controls and fed to a group of 100 2nd instar larvae used as
controls. Each spore dosage was administered to 50 larvae.
Infected larvae were placed on sterile Petri dishes containing Potato Dex-
trose agar plus 0.5% yeast extract (PDAYE). One half gram Streptomycin
127
VOLUME XC, NUMBER 2
Table 1. Percentage mortality of gypsy moth larvae treated with surface applied suspen-
sions of entomogenous fungi.
Dosage spores/mm^
50
100
500
1,000
5,000
72 hr
120 hr
72 hr
120 hr
72 hr
120 hr
72 hr
120 hr
72 hr
120 hr
B. bassiana
56
92
88
88
92
92
90
90
96
96
B. tenella
80
96
90
98
90
90
92
92
98
98
C. militaris
66
90
88
92
92
98
90
98
100
100
M. anisopliae
72
94
94
100
94
100
98
100
98
100
N. rileyi
80
96
90
100
98
100
100
100
92
100
P. farinosus
72
96
88
98
88
98
88
100
100
100
P. fumoso-roseus
84
100
88
100
92
100
96
100
88
100
and 0.5 g Penicillin G were added after the PDA YE was autoclaved for 15
min at 15 pounds pressure. The target organisms were recovered in all cases.
Results
The mortality rates of gypsy moth larvae to various dosages of entomoge-
nous fungal spore suspensions have been summarized under Table 1 . Con-
trol larvae showed 3% mortality after 72 hours and 5% mortality at the end
of 120 hours. Of the 12 test species, 5 were considered non-infective under
the conditions of the experiment. These were, Conidiobolus thromboides,
Fusarium sp., Hirsutella thompsonii, Paecilomyces farinosus (Hamden,
Ct. strain) and V erticillium sp.
Paecilomyces fumoso-roseus, Metarhizium anisopliae and Nomuraea ri-
leyi were considered highly infective as 100% mortality was recorded within
120 hours of treatment with dosage levels as low as 100 spores per square
millimeter of leaf surface. There was a linear dosage-mortality response and
five of the seven infective species produced 100% mortality at the high
dosages of 1,000 and 5,000 spores/mm^. The target fungus was recovered
from dead larvae and survivors did not show any adverse effects as a result
of the treatment.
These tests simulate natural conditions as leaf surfaces were used for
administration of the dosages. Entomogenous fungi exhibit varying levels
of mortality against gypsy moth larvae and 5 of the 12 test species were
non-infective.
Acknowledgments
This research was supported (in part) by the funds provided by the USDA
Forest Service, North Eastern Forest Experiment Station, Broomall, Pa.
128
NEW YORK ENTOMOLOGICAL SOCIETY
Literature Cited
Donovan-Peluso, M., S. S. Wasti and G. C. Hartmann. 1980. Safety of entomogenous fungi
to vertebrate hosts. Appl. Entomol. Zool. 15:498-499.
Garcia, C. and C. M. Ignoffo. 1978. A simplified diet-surface treatment technique for deter-
mining the infectivity of conidia of Nomuraea rileyi. J. Invertebr. Pathol. 32:398-399.
Hartmann, G. C. and S. S. Wasti. 1974. Infection of the gypsy moth, Porthetria dispar (L.)
with the entomogenous fungus, Conidiobolus coronatus (Cost.) Batko. Entomophaga
19:353-360.
and . 1976. Experimental mycoses of the gypsy moth, Porthetria dispar (L.) by
Entomophthora virulenta with notes on its mammalian safety. Entomophaga 21:377-
382.
and . 1980. Avian safety of three species of entomogenous fungi. Comp. Physiol.
Ecol. 5:242-245.
, and D. L. Hendrickson. 1979. Murine safety of two species of entomogenous
fungi, Cordyceps militaris and Paecilomyces fumoso-roseus. Appl. Entomol. 14:217-
220.
Latge, J. P., D. S. King and B. Paperiok. 1980. Synonymie de Entomopthora virulenta Hall
& Dunn et de Conidiobolus thromboides Drechsler. Mycotaxonomy 11:255-268.
O’Dell, T. M. and W. D. Rollinson. 1966. A technique for rearing the gypsy moth, Porthetria
dispar on an artificial diet. J. Econ. Entomol. 59:741-742.
Wasti, S. S. and G. C. Hartmann. 1975. Experimental parasitization of larvae of the gypsy
moth, Porthetria dispar (L.) with the entomogenous fungus, Beauveria bassiana (Bals.)
Vuill. Parasitology 70:341-346.
and . 1978. Host-parasite interactions between larvae of the gypsy moth, Ly-
mantria dispar (L.) and the entomogenous fungus, Nomuraea rileyi (Farlow) Samson.
Appl. Entomol. Zool. 13:23-28.
, and A. J. Rousseau. 1980. Gypsy moth mycoses by two species of entomoge-
nous fungi and assessment of their avian toxicity. Parasitology 80:419^24.
Department of Biology, Rhode Island College, Providence, Rhode Island
02908.
Received for publication November 17, 1981.
NEW YORK ENTOMOLOGICAL SOCIETY
XC(2), 1982, pp. 129-133
HABITAT DIFFERENCES IN FEEDING HABITS AND
BODY SIZE OF THE PREDATORY STINKBUG
PERILLUS CIRCUMCINCTUS
(HEMIPTERA: PENTATOMIDAE)
Edward W. Evans^
Abstract. — Near Ithaca, New York, Perillus circumcinctus reproduces in
stands of Bidens cernua and Solidago spp. (Compositae) in early summer.
Nymphs of this predator consume larvae and adults of the beetle Calligra-
pha calif ornica coreopsivora (Chrysomelidae) in stands of B. cernua, and
primarily larvae of Trirhabda spp. (Chrysomelidae) in stands of Solidago.
Newly molted adults of P. circumcinctus in stands of Solidago are larger
than newly molted adults of P. circumcinctus in stands of B. cernua. It is
suggested that differences in the physical structure of the two habitats result
in nymphs of P. circumcinctus experiencing more difficulty in harvesting
prey in stands of B. cernua. Increased difficulty in harvesting prey results
in smaller adult sizes.
Introduction
The predatory stinkbug Perillus circumcinctus Stal (Hemiptera: Penta-
tomidae) is a specialized predator of beetles of the family Chrysomelidae
(Hart and Gleason 1907; Vestal 1913; Knight 1923; Parshley 1923). In the
Ithaca, New York, area this univoltine predator usually oviposits in stands
of goldenrod {Solidago spp.) and reproduction is timed so that nymphs
mature by feeding almost entirely on beetle larvae of the genus Trirhabda
(Chrysomelidae) (Evans 1982a). While studying the habits of P. circum-
cinctus near Ithaca, I found this species in only one additional habitat,
stands of Bidens cernua (Compositae). A comparison of the seasonal pat-
terns, feeding habits, and adult body sizes of P. circumcinctus in B. cernua
and Solidago provides a perspective on the ability of this predator to exploit
chrysomelid beetles as prey.
Observations
On 3 July 1979, a dense but very local population of P. circumcinctus was
found in stands of B. cernua on the edge of a small pond (Bull Pasture Pond,
^ Present address: Division of Biology, Kansas State University, Manhattan, Kansas 66506.
The publication costs of this article were defrayed in part by page charge payment. This article
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to indicate this fact.
130 NEW YORK ENTOMOLOGICAL SOCIETY ^
Cornell University Golf Course). Both nymphs and adults of the predator n
were feeding on larvae and adults of Calligrapha californica coreopsivora ,i
Brown (Coleoptera: Chrysomelidae). Voucher specimens of both predator ri
and prey have been placed in the Cornell University Insect Collection under n
Lot 1086. The age structure of the P. circumcinctus population onB. cernua i-
was very similar to that of populations of the predator on or about the same n
date in stands of goldenrod (Evans 1982a): fourth- and fifth-instar nymphs >
and adults predominated, although younger nymphs (including a group of Ij
newly hatched first-instar nymphs) were also present. Several clusters of
hatched eggs of P. circumcinctus were found on the vegetation. Relatively :
few late-instars of larvae of the beetle were present but adults were com- j
mon. !
Twenty meters distant grew a small (20 x 10 m) but dense stand of gold- |
enrod where nymphs and adults of P. circumcinctus also occurred. Adults i
of P. circumcinctus were collected in both habitats. Vernier calipers were
used to measure length from the anterior tip of the pronotum to the posterior
tip of the abdomen and width across the widest part of the abdomen. Adults
of P. circumcinctus from goldenrod at Bull Pasture Pond and other localities
were of similar size. However, adults from B. cernua were unusually small
(Tables 1 and 2; P < .001 for length and width of both sexes in t test com-
parisons of adults from B. cernua vs. from goldenrods at all sites).
All adults of P. circumcinctus were marked and released in both habitats,
after measuring them. No marked individuals were recaptured. By mid- July,
individuals of P. circumcinctus were much less numerous in both habitats
and most individuals were adults. No larvae of C. californica coreopsivora
were found and adults were also much less abundant than formerly. Adult
Table 1. The mean (Jc) ± two standard errors (SE), range, and sample size (N), of lengths
and widths (mm) of P. circumcinctus females captured on comparable dates in early and mid-
July in 1979 in stands of B. cernua and Solidago (BPP: Bull Pasture Pond, Brk; Brooktondale;
descriptions of the Brooktondale and Whipple sites appear in Evans 1980).
Vegetation
Site
Date
N
Length
Width
i ± 2 SE
Range
X ± 2 SE
Range
Bidens
BPP
July 3
9
7.9 ± 0.1
7.8-8. 1
5.5 ± 0.1
5.4-5. 6
Solidago
BPP
July 3
2
8.3 —
8. 1-8.5
5.6 —
5. 5-5. 7
Brk
July 3
10
8.4 ± 0.1
8. 2-8. 7
5.8 ± 0.1
5. 5-6.0
Whipple
July 1
22
8.5 ± 0.1
8. 2-8. 8
5.8 ± 0.1
5.6-6. 1
Bidens
BPP
July 18
4
7.9 ± 0.1
7. 6-8. 2
5.4 ± 0.1
5. 3-5.7
Solidago
BPP
July 18
2
8.4 —
8. 3-8. 4
5.6 —
5. 6-5. 7
Brk
July 17
3
8.3 ± 0.1
8. 3-8. 4
5.8 ± 0.2
5. 6-6.0
Whipple
July 19
11
8.3 ± 0.2
7. 5-8. 7
5.6 ± 0.1
5. 2-5. 8
VOLUME XC, NUMBER 2
131
Table 2. The mean (i) ± two standard errors (SE), range, and sample size (N), of lengths
and widths (mm) of P. circumcinctus males captured on comparable dates in early and mid-
July 1979 in stands of B. cernua and Solidago (BPP: Bull Pasture Pond; Brk: Brooktondale).
Vegetation
Site
Date
N
Length
Width
X ± 2 SE
Range >
^ ± 2 SE
Range
Bidens
BPP
July 3
17
7.3 ± 0.1
6. 9-7. 6
5.1 ± 0.1
4. 9-5.4
Solidago
BPP
July 3-4
4
7.7 ± 0.2
7. 6-7. 9
5.5 ± 0.2
5. 3-5. 7
Brk
July 3
13
7.7 ± 0.1
7.^8. 0
5.4 ± 0.1
5. 2-5. 6
Whipple
July 1
11
7.6 ± 0.1
7. 3-8. 2
5.3 ± 0.1
5. 0-5. 6
Bidens
BPP
July 18
4
7.4 ± 0.1
7.3-7.5
5.1 —
5.1
Solidago
BPP
July 18
3
7.6 ± 0.1
7. 5-7.7
5.3 ± 0.1
5. 2-5. 4
Brk
July 17
6
7.3 ± 0.2
6. 8-8. 2
5.1 ± 0.2
5.0-5.6
Whipple
July 19
11
7.4 ± 0.1
7. 2-7. 6
5.1 ± 0.1
4. 8-5.4
females of the predator from B. cernua were still unusually small in com-
parison to females in goldenrod stands (Table \, P < .01 for length, P =
.10 for width in t test comparisons). However, adult males did not differ
significantly in either length or width in the two kinds of vegetation (Ta-
ble 2).
On 5 July 1979 I visited a second stand of B. cernua at the edge of a small
pond several km west of Ithaca. Calligrapha californica coreopsivora was
very abundant (much more so that at Bull Pasture Pond), but only a few
nymphs and two adult males of P. circumcinctus were found in an extensive
search. Sparse stands of goldenrod grew on hillsides surrounding the pond.
In one of these stands, a single P. circumcinctus adult male was collected.
As at Bull Pasture Pond, this individual was larger (length: 7.5 mm, width:
5.4 mm) than the two males collected from B. cernua (7.3 and 7.0 mm in
length, 5.0 and 4.8 mm in width, respectively).
Discussion
The predator-prey relationship of P. circumcinctus and C. californica
coreopsivora in stands of B. cernua bears strong resemblance to the rela-
tionship between the predator and its prey {Trirhabda) in stands of gold-
enrod (Evans 1982a). Both prey have sluggish larvae which briefly occur in
great densities in early summer. Nymphs of P. circumcinctus easily subdue
the sluggish larvae of Trirhabda (Evans 1982b). The weak responses of C.
californica coreopsivora larvae when probed with a pencil suggest that these
larvae are also easily captured by stinkbug nymphs. The similarity in age
structure of P. circumcinctus populations in stands of goldenrod and B.
cernua in July suggests that the predator’s timing of reproduction is similar
132
NEW YORK ENTOMOLOGICAL SOCIETY
in both habitats. Thus the development of P. circumcinctus nymphs appar- ;
ently is well synchronized with availability of C. californica coreopsivora (
larvae as prey in stands of B. cernua in fashion similar to patterns of pred- ■
ator and prey observed in goldenrod stands (Evans 1982a). |j
The major difference between individuals of the predator in the two hab- '
itats is the difference in body size of newly molted adults. The rate that i
nymphs consume prey has a dramatic effect on subsequent adult size in P. :
circumcinctus (Evans 1980). Thus the striking difference in body size of
adults maturing at the same time in adjacent stands of Solidago and B.
cernua suggests that the food supply differed in these two habitats. In par-
ticular P. circumcinctus nymphs may have experienced greater difficulty in
seeking out prey on B. cernua than on Solidago. Stands of B. cernua were
less dense than Solidago stands, and individual plants touched each other
less frequently in B. cernua than in Solidago stands. Many plants of B.
cernua were isolated even more because they grew in shallow water. These
attributes of B. cernua presumably hampered the searching abilities of P.
circumcinctus', hence C. californica coreopsivora was probably protected
from the predators by a “moat effect” (Eickwort 1977). In goldenrod stands,
predators travelled readily from plant to plant and hence larvae of Trirhabda
received little protection (Evans 1982b). Eurther study of the hunting habits
of P. circumcinctus should reveal the degree to which the physical structure
of these two habitats influences this predator’s ability to exploit its prey.
Acknowledgments
These observations were made in partial fulfillment of the Ph.D. degree
at Cornell University. I thank R. B. Root for his support and guidance in
this endeavor. I also thank S. Juliano for bringing the presence of P. cir-
cumcinctus at Bull Pasture Pond to my attention, and E. R. Hoebeke for
confirming the identity of Calligrapha californica coreopsivora. Financial
support was provided in part by N.S.F. Grant DEB 77-25210 to R. B. Root
and by Hatch Project 410, New York State College of Agriculture and Life
Sciences, Cornell University.
Literature Cited
Eickwort, K. R. 1977. Population dynamics of a relatively rare species of milkweed beetle
(Labidomera). Ecology 58:527-538.
Evans, E. W. 1980. Lifeways of predatory stinkbugs: feeding and reproductive patterns of a
generalist and a specialist (Pentatomidae: Podisus macidiventris and Perillus circum-
cinctus). Dissertation. Cornell University, Ithaca, N.Y.
. 1982a. Timing of reproduction by predatory stinkbugs (Hemiptera: Pentatomidae):
patterns and consequences for a generalist and a specialist. Ecology 63: 147-158.
. 1982b. Feeding specialization in predatory insects: hunting and attack behavior of two
stinkbug species (Hemiptera: Pentatomidae). American Midland Naturalist, in press.
VOLUME XC, NUMBER 2
133
Hart, C. A. and H. A. Gleason. 1907. On the biology of the sand dunes of Illinois. Bulletin
of the Illinois State Laboratory of Natural History 7:137-272.
jKnight, H. H. 1923. Studies of the life history and biology of Perillus bioculatus. Report of
the Minnesota State Entomologist 19:50-96.
Parshley, H. M. 1923. Family Pentatomidae. Pages 753-776 in: W. E. Britton, editor. Guide
to the Insects of Connecticut. IV. The Hemiptera or sucking insects of Connecticut.
' Connecticut State Geological and Natural History Survey, Hartford, Connecticut.
Vestal, A. G. 1913. An associational study of Illinois sand prairie. Bulletin of the Illinois State
I Laboratory of Natural History 10: 1-96.
Section of Ecology and Systematics, Cornell University, Ithaca, New
York 14853.
Received for publication July 17, 1981.
NEW YORK ENTOMOLOGICAL SOCIETY
XC(2), 1982, p. 133
BOOK REVIEW
The North American Grasshoppers. Vol. /. Acrididae: Gomphocerinae and
Acridinae. By Daniel Otte. Harvard University Press. 275 p. 1981. $45.
All known species of the slant-faced grasshoppers of North America,
including the Caribbean Islands, are illustrated by detailed drawings and 16
full color plates prepared by the author. Keys for genera and species, dis-
tribution maps, habitat preferences, as well as grasshopper ecology, behav-
ior and life cycles are discussed in detail. Taxonomic changes made in this
volume are included in Appendix I. Genera and species of Gomphocerinae
and Acridinae are in Appendices II and III. A glossary of terms and a
[ taxonomic index complete the volume. The book is carefully written, the
format of the volume is excellent and typographical errors are rare. The
author. Associate Curator of Entomology at the Academy of Natural Sci-
ences in Philadelphia has provided a valuable contribution to the taxonomy
of grasshoppers and a definitive reference that will be especially appreciated
by all who are interested in these grazing herbivores and pests, as well as
by taxonomists, ecologists, and economic entomologists. The forthcoming
volumes II and III will comprise the Oedipodinae, Melanoplinae, Romal-
einae and other smaller groups.
Karl Maramorosch, Waksman Institute of Microbiology, Rutgers Uni-
versity.
NEW YORK ENTOMOLOGICAL SOCIETY i
XC(2), 1982, p. 134 i
BOOK REVIEW
Insects: An Anthology of Arthropods Featuring a Bounty of Beetles. Paint-
ings by Bernard Durin, introduction by Paul Armand Gette, entomological
commentaries by Gerhard Scherer. Translated from German by Georg
Zappler. Hudson Hills. Distributed by Simon & Schuster. 108 p. 34 full
color plates. 1981. $50.
This beautiful, large (11" x 14") format volume can only be described in
superlative. It contains 34 remarkably well reproduced illustrations by Ber-
nard Durin of butterflies, grasshoppers, beetles, wasps, flies, and a scorpion.
The full color plates give the impression of superb color photographs, be-
cause of the faint shadows in the background. The illustrations of arthropods
are unquestionably among the best ever published. Gette’s introduction pro-
vides a link between art and science, with an anthology of poems and writ-
ings about insects by Nabokov, Lewis Carroll, Thoreau, Poe, Heine and
others. The delightful book is a treasury of fact and fancy for entomologists
as well as for wildlife lovers and all those who appreciate art and fine books.
The authors, particularly Bernard Durin, one of the foremost illustrators,
and the publishers are to be congratulated for making available this art book
to entomologists and nature lovers. The volume will constitute a valuable
addition to any university and public library. Its price, justified for an art
book with such fine reproductions of the dazzling paintings, unfortunately
puts it out of the reach of all but the most affluent entomologists.
Karl Maramorosch, Waksman Institute of Microbiology, Rutgers Uni-
versity.
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Journal of the
New York Entomological Society
VOLUME XC SEPTEMBER 1982
NO. 3
EDITORIAL BOARD
Editor Associate Editors
Dr. Karl Maramorosch Dr. Lois J. Keller, RSM
Waksman Institute of Microbiology Dr. Herbert T. Streu
Rutgers University
New Brunswick, New Jersey 08903
Publication Committee
Dr. Randall T. Schuh
American Museum of
Natural History
Dr. Louis Trombetta
St. Johns University
CONTENTS
Onset of phenotaxis and age at first mating in female house crickets, Acheta domesti-
cus (Orthoptera: Gryllidae) Scott K. Sakaluk 136-141
Larval morphology and phylogeny of Trichordestra tacoma (Strecker) (Noctuidae)
Tim L. McCabe and George L. Godfrey 142-146
Descriptions, synonymy and sex associations in the genus Eucerceris (Hymenop-
tera: Philanthidae) George R. Ferguson 147-160
Perching behavior of Canthon viridis (Coleoptera: Scarabaeidae) in Maryland
Orrey P. Young 161-165
Descriptions of the immature stages of Myndus criidus (Homoptera: Fulgoroidea:
Cixiidae) Stephen W. Wilson and James H. Tsai 166-175
Unionicolid mites from central New York R. A. Baker 176-180
Predators, parasites, and associates of Anthophora abrupta Say (Hymenoptera:
Anthophoridae) Beth B. Norden and Aubrey G. Scarbrough 181-185
Entomology in The People’s Republic of China James H. Tsai 186-212
Effects of two insect growth regulators (Hydroprene and R-20458) on the follicular
epithelium and the oocytes of the rice weevil, Sitophilus oryzae (L.) (Coleoptera:
Curculionidae) J. M. Mkhize and A. P. Gupta 213-219
NEW YORK ENTOMOLOGICAL SOCIETY
XC(3), 1982, pp. 136-141
ONSET OF PHONOTAXIS AND AGE AT FIRST MATING IN
FEMALE HOUSE CRICKETS, ACHETA DOMESTICUS
(ORTHOPTERA: GRYLLIDAE)
Scott K. Sakaluk*
Abstract.— ThQ age at which females first become phonotactic to conspe-
cific calling song, and the age at which females first mate were studied in
the house cricket, Acheta domesticus L. (Orthoptera: Gryllidae). The average
age at which virgin females first showed positive phonotaxis to tape recorded
male calling song was 5.4 days after the last molt. In another group of adult
female A. domesticus, the average age at which females first mated was 6.9
days, a significantly later age than that for phonotaxis. Early onset of pho-
notaxis may function to increase mate availability when females become
sexually receptive.
Introduction
Formation of sexual pairs in crickets is facilitated by male calling songs
which attract females (Alexander and Otte 1 967). Species specificity of female
cricket phonotaxis to the calling songs of conspecific males has been dem-
onstrated repeatedly (e.g. Forrest 1980; Paul 1976; Popov and Shuvalov
1977; Shuvalov and Popov 1971, 1973; Ulagaraj and Walker 1973; Walker
1957; Zaretsky 1972). Although the pairing function of male calling songs
is clear, scant evidence exists indicating a developmental link between female
phonotaxis and the readiness of a female to copulate (Stout et al. 1976).
Shuvalov and Popov (1971) found that only sexually mature female Acheta
domesticus (Gryllidae: Gryllinae), 7-10 days after the imaginal molt, exhib-
ited positive phonotaxis. Additionally, Stout et al. (1976) tested responsive
and non-responsive female A. domesticus (determined by whether a female
mounted a male during 30 min of conhnement) for phonotaxis: the speed
of responsive females in phonotactic locomotion was greater than non-
responsive females, although this difference was not significant.
During the course of a study of A. domesticus reproductive behaviour
' Present address: Department of Zoology, Erindale College, University of Toronto, Missis-
sauga, Ontario L5L 1C6.
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VOLUME XC, NUMBER 3
137
(Sakaluk and Cade 1980; Sakaluk 1981), the onset of female phonotaxis and
the age at which females first mate were quantified. I here report experimental
evidence which indicates that females show positive phonotaxis at a signif-
icantly earlier age than that at which they first mate.
Methods of Study
Immature A. domesticus, purchased from Armstrong’s Cricket Farm,
Monroe, Louisiana, were housed in a fibreglass chamber containing ample
food (Purina® Mouse Chow), cotton-plugged test tubes containing water,
and layered egg cartons for cover. Late instar females were removed and
held separately, thus ensuring virginity upon the imaginal molt. The nymphs
were examined every day and adult females were considered 1 day old on
the day they were found to have molted. Crickets were exposed throughout
to a 1 2 h light : 1 2 h dark photoperiod opposite the ambient photoperiod
and at 28 ± 2°C.
To determine the age at which females first mate, 1 day old virgin A.
domesticus were numbered and observed for 1 5 consecutive days in a plastic
mating arena (78 cm X 49 cm X 10 cm). Ajar lid which contained crushed
mouse chow, another which contained moistened vermiculite for oviposi-
tion, and 2 cotton-plugged test tubes containing water were placed in the
arena. The number of females in the arena ranged from 1 5-20 depending
on the molting frequency and the death of some females. The same number
of sexually mature males was placed in the arena each day and observed for
2 h under red lights, 3 h into the dark portion of the photoperiod. Age at
first mating was recorded for each female.
To determine the onset of positive phonotaxis to male calling song in
female A. domesticus, a separate group of females was tested using tape-
recorded A. domesticus song. One day old virgin females obtained from the
same colony were housed in a separate terrarium without males. Trials were
conducted 1 hour into the dark portion of the photoperiod in a circular
arena (105 cm (diam) X 15 cm (ht)) constructed of Styrofoam®; this exper-
iment immediately preceded the test for first mating. The arena contained
2 Phillips AD 0160/T8 loudspeakers embedded in the wall of the arena at
a height of 3 cm and 2 red lights at 90° to the speakers. A Sanyo M22 1 1
cassette tape recorder broadcast previously recorded A. domesticus calling
song through 1 loudspeaker at 70 dB (A scale) as measured with a General
Radio Sound Level Meter -1565B held 10 cm in front of the loudspeaker.
This sound intensity was consistent with previous measurements of A.
domesticus males calling in the laboratory. Recordings were made with a
Uher 240 CR tape recorder and a Uher M640 microphone. In each trial, a
single female was placed under an inverted plastic vial in the centre of the
arena. After a 5 min period in which the female was allowed to become
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NEW YORK ENTOMOLOGICAL SOCIETY
quiescent, the vial was removed and broadcasting of taped song commenced.
A trial lasted until a positive response was recorded, until 5 min had elapsed,
or until the female left the arena. A positive response involved a female
remaining in a marked area ( 1 5 cm X 1 1 cm) under the speaker and/or on
the speaker for a total duration of 1 min of the 5 min trial. All females were
tested once every day until a day of positive response was recorded, and 9
of the 28 females were tested for 1 5 consecutive days regardless of the age
at which they first showed positive phonotaxis.
Results
Data on the age at which females first mated are in Fig. 1. Female A.
domesticus mated for the first time at an average age of 6.9 days (N = 103;
SD = 2.7 days). Data on the age at which females first showed positive
phonotaxis to conspecific calling song are in Fig. 2. Females were first attracted
to a loudspeaker broadcasting A. domesticus song at 5.4 days of adult age
(N = 28; SD = 2.4 days). The mean number of positive phonotactic responses
exhibited by the 9 females tested over 1 5 consecutive days was 7.8 (SD =2.2;
range = 5-11). The mean ages at which females first mated and first became
phonotactic are significantly different (t = 2.6; P < 0.01).
VOLUME XC, NUMBER 3
139
Age at first acoustical receptivity
Fig. 2. The age at which female A. domesticus first showed a positive phonotactic response
to conspecific calling song.
Discussion
The dominant carrier frequency of the male A. domesticus calling song is
4.6 kHz, with the maximum sensitivity of the cricket’s tympanal ear falling
within 4-7 kHz (Counter 1976). However, Shuvalov and Popov (1971)
showed that up to 85% of female A. domesticus, less than 5 days of age after
the imaginal molt, exhibited an escape response when exposed to artihcially
generated square pulse signals emitted at 4 kHz. This conflicts with our
finding that 61% of female A. domesticus exhibited positive phonotaxis to
broadcast male calling song within 5 days of age. The discrepancy in these
results likely reflects the use of different source stimuli.
The close correspondence of onset of female phonotaxis with the age at
which females first mated indicates that female phonotactic responsiveness
and readiness to copulate in A. domesticus are developmentally linked. This
is supported further by the similarity in shape of the frequency distributions.
However, the onset of female phonotaxis occurred at a significantly earlier
age than that at which females first mated. When phonotaxis precedes mat-
ing, selection may favour phonotaxis by non-receptive females since males
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NEW YORK ENTOMOLOGICAL SOCIETY
will thus be present when mating is possible. Pair formation is not the only
plausible result of female phonotaxis. In mole crickets (Forrest 1980; Ula-
garaj and Walker 1973), field crickets (Cade 1979a), and katydids (G. K.
Morris, pers. comm.), females may use the conspecific calling song as a cue
for resource location.
Females tested for phonotaxis over 1 5 consecutive days exhibited, on the
average, 8 phonotactic responses. Cade (1979b) found that when female field
crickets were deprived of males (as was the case here), the frequency of
positive phonotaxis was increased significantly. Nonetheless, female A.
domesticus did not always respond subsequent to the first phonotactic
response. It may be that sexually mature females that do not exhibit pho-
notaxis will not mate.
The age at which females first mate may correspond to when mature eggs
are available for fertilization. No ovariole growth occurs in the last nymphal
stadium, but begins to increase drastically at an adult age of 3 days in A.
domesticus (Woodring et al. 1979). Therefore, adult females that mated
before the availability of mature eggs might encounter reduced fitnesses due
to time and energy expenditures. Also, adult crickets are soft-bodied for
about 3 days after eclosing and females might risk physical damage if they
mated before the cuticle hardened.
A different situation exists for field crickets, Teleogryllus commodus Walker,
as females less than 24 h old will mate. Two females that were 4 and 5 h
old were observed mating even though the cuticle was soft and the ovaries
were immature. Eggs laid by these females were fertilized and yielded viable
offspring (Loher and Edson 1973). Additionally, conspecific male calling
song does not initiate running activity in female T. commodus 3 days of age
after the imaginal molt. This suggests that a female is ready to mate before
she can be attracted to the male calling song (Loher 1979). These differences
clarify the need for further ontogenetical studies, before generalizations con-
cerning the development of female cricket sexual responsiveness can be
made.
Acknowledgments
I thank Dr. William H. Cade for his guidance during this research project
and writing of the manuscript. I also thank Paul Bell, Dr. Glen K. Morris,
Dr. W. Gary Sprules, and an anonymous reviewer for comments on the
manuscript. Financial support was provided by a Natural Sciences and Engi-
neering Research Council grant to W. Cade (A6 174) and an Ontario Graduate
Scholarship to S.K.S. Page charges were met by the Department of Zoology,
Erindale College at the University of Toronto. This study represents a portion
of a thesis submitted to Brock University in partial fulfillment for the M.Sc.
degree.
VOLUME XC, NUMBER 3
141
Literature Cited
Alexander, R. D. and D. Otte. 1 967. The evolution of genitalia and mating behavior in crickets
(Gryllidae) and other Orthoptera. Misc. Publ. Mus. Zool. Univ. Mich. 133:1-62.
Cade, W. H. 1979a. The evolution of alternative male reproductive strategies in field crickets.
In: M. S. Blum and N. A. Blum (eds.). Sexual Selection and Reproductive Competition
in Insects. Academic Press, New York.
. 1979b. Effects of male-deprivation on female phonotaxis in field crickets (Orthoptera:
Gryllidae; Gryllus). Can. Entomol. 111:741-744.
Counter, S. A. Jr. 1976. Fourier and electrophysiological analyses of acoustic communication
in Acheta domesticus. J. Insect Physiol. 22:589-593.
Forrest, T. G. 1980. Phonotaxis in mole crickets: its reproductive significance. Fla. Entomol.
63:45-53.
Loher, W. 1979. Circadian rhythmicity of locomotor behavior and oviposition in female
Teleogryllus commodus. Behav. Ecol. Sociobiol. 5:253-262.
and K. Edson. 1973. The effect of mating on egg production and release in the cricket
Teleogryllus commodus. Entomol. Exp. Appl. 16:483-490.
Paul, R. C. 1976. Species specificity in the phonotaxis of female ground crickets (Orthoptera;
Gryllidae: Nemobiinae). Ann. Ent. Soc. Am. 69:1007-1010.
Popov, A. V. and V. F. Shuvalov. 1977. Phonotactic behavior of crickets. J. Comp. Physiol.
119:111-126.
Sakaluk, S. K. 1981. Sexual behaviour and factors affecting female reproduction in house and
field crickets. M.Sc. thesis. Brock University, St. Catharines. 176 pp.
and W. H. Cade. 1980. Female mating frequency and progeny production in singly
and doubly mated house and field crickets. Can. J. Zool. 58:404-41 1.
Shuvalov, V. F. and A. V. Popov. 1971. Reaction of females of the domestic cricket Acheta
domesticus to sound signals and its changes in ontogenesis. J. Evol. Biochim. Fisiol. 7:
612-616.
and . 1973. Study of the significance of some parameters of calling signals of
male crickets Gryllus bimaculatus for phonotaxis of females. J. Evol. Biochim. Fisiol.
9:177-182.
Stout, J. F., G. Gerard and S. Hasso. 1976. Sexual responsiveness mediated by the corpora
allata and its relationship to phonotaxis in the female cricket, Acheta domesticus L.
J. Comp. Physiol. 108:1-9.
Ulagaraj, S. M. and T. J. Walker. 1973. Phonotaxis of crickets in flight: attraction of male
and female crickets to male calling songs. Science 182:1278-1279.
Walker, T. J. 1957. Specificity in the response of female tree crickets to calling songs of the
males. Ann. Ent. Soc. Am. 50:626-636.
Woodring, J. P., C. W. Clifford and B. R. Beckman. 1979. Food utilization and metabolic
efficiency in larval and adult house crickets. J. Insect Physiol. 25:903-912.
Zaretsky, M. D. 1 972. Specificity of the calling song and short term changes in the phonotactic
response by female crickets, Scapsipedus marginatus (Gryllidae). J. Comp. Physiol. 79:
153-172.
Department of Biological Sciences, Brock University, St. Catharines,
Ontario L2S 3A1.
Received for publication August 25, 1981.
NEW YORK ENTOMOLOGICAL SOCIETY
XC(3), 1982, pp. 142-146
LARVAL MORPHOLOGY AND PHYLOGENY OF
TRICHORDESTRA TACOMA (STRECKER) (NOCTUIDAE)'
Tim L. McCabe and George L. Godfrey
Abstract.— larva of Trichordestra tacoma (Strecker) (Lepidoptera;
Noctuidae) has been misidentified in the past, resulting in a discrepancy in
the phylogenetic placement of this species in separate works based on adults
and on larvae. Both larva and adult prove to be similar to T. rugosa (Morr.)
and T. liquida (Grote). The larva is described and illustrated, and a host list
is presented.
Comparisons of two recent reclassifications of hadenine noctuids based
on larvae (Godfrey 1972) and on a section of the adults (McCabe 1980) has
yielded a discordance in the phylogenetic placement of Mamestra tacoma
Strecker (1900). M. tacoma has been placed in Folia (McDunnough 1938),
but recently the genus Trichordestra was erected for it and its relatives
(McCabe 1980). Trichordestra presently includes eight nearctic species, the
larvae of which are characterized by two inner mandibular teeth and a striped
body pattern in addition to the characters Godfrey (1972) used in his group
seven. The adult moths are best defined by three bandlike signa on the
female bursa copulatrix.
One of us (TLM) cultured the larvae and discovered that specimens pre-
viously determined, by association, as “Po/za” tacoma were actually another,
as yet unidentified noctuid larva. True T. tacoma larvae have the two inner
mandibular teeth so typical of other known Trichordestra larvae (Fig. 1). T.
tacoma larvae were discovered to have two color phases: the typical phase
is green and appears unstriped in life, although faint stripes are traceable in
preserved larvae; a red color phase has prominent stripes and in this manner
is more typical of the genus. Dyar (1904) may have described the green
phase of T. tacoma larvae.
T. tacoma are related to T. liquida (Grote) on the basis of the paired, inner
mandibular teeth with the tooth in the second inner ridge being weakly
developed (broken off or worn down in some specimens of T. tacoma) (Fig.
' Published by permission of the Director, New York State Museum, State Education Depart-
ment, Journal Series No. 324.
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VOLUME XC, NUMBER 3
143
Figs. 1-3. 1. Oral aspect of the left mandibles of the known larvae of Trichordestra. 2.
Hypopharynx of the larva of T. tacoma. 3. Frontal aspect of the head capsule of T. tacoma.
144
NEW YORK ENTOMOLOGICAL SOCIETY
5
Figs. 4, 5. 4. Green phase of living larva of T. tacoma from the Adirondack Mountains. 5.
Red phase larva from the same locality.
1). T. tacoma and T. rugosa (Morr.) adults have an appendix bursa nearly
as large as the bursa copulatrix in the females, asymmetrical valves in the
males, and both lack male coremati. The adult male is also similar to T.
liquida in their similar habitus, similar broad cuculli, and similar asym-
metrical valves.
VOLUME XC, NUMBER 3
145
Confined first instar larvae of T. tacoma fed on Apocynum androsaemi-
folium L. (APOCYNACEAE), Sambucus canadensis L. (CAPRIFOLI-
ACEAE), Betula papyrifera Marsh (CORYLACEAE), Vaccinium myrtil-
loides Michx. (ERICACEAE), and Prunus virginiana L. and Spiraea latifolia
(Ait.) Borkh. (ROSACEAE). First instar larvae rejected Carex spikes
(CYPERACEAE), Abies balsamea L. (PINACEAE), and Viola selkirkii Pursh
(VIOLACEAE). Larvae from an early season female did very well on Prunus
and were reared to maturity on it, however, larvae from a late-season female
had older Prunus leaves and did poorly until switched to a non-woody host
{Apocynum).
First instar larvae have prominent pinaculi, a weakly sclerotized pro-
thoracic shield, and reduced prolegs on the third and fourth abdominal
segments. The mature larva is described below.
General (Figs. 4, 5). — Head 2. 9-3.0 mm wide; total length (fully expanded)
36 ± 6 mm (N = 14). Abdominal prolegs present on 3-6th and 10th seg-
ments.
Coloration (living material). — Green phase (Fig. 4): Soft yellow-green with
pale middorsal line, a pair of slightly darker subdorsal bands on each side
separated by a narrow pale line, then a pale lateral band which includes the
top half of the spiracles followed below by a greenish white subspiracular
band which includes the bottom half of the spiracles. Red phase (Fig. 5):
Similar to preceding, but much darker, with prominent stripes and with the
prevalent color a brownish red.
Head (Fig. 3).— Epicranial suture 0.94 mm long. Height of frons 0.96 mm.
Top half of head with reticulate pattern near epicranial suture and between
setae P-2 and L. Seta P- 1 as long as frons is wide. Ocellar interspaces between
Ocl-Oc2 .5 X diameter of Oc2; Oc2-Oc3 1.5 X Oc2; Oc3-Oc4 .5 X Oc3;
Oc4-Oc5 2.5 X Oc4; Oc5-Oc6 3.0 X Oc5.
— Hypopharyngeal complex (Fig. 2): With spinneret as long
as labial palpi and apical seta combined (Lpg 1&2 and Lp2); distal region
with fine spines; proximolateral region with a row of 1 9-27 spines. Mandible
(Fig. 1): Two shallow inner teeth, 6 triangular outer teeth.
Body segments. — ProXhomx: SD-2 and SD-1 lacking pinaculum; LI and
L2 share weakly developed pinaculum. Meso- and metathorax and abdom-
inal segments without pinaculi. Crochets uniordinal, 16-26 on 3rd, 18-26
on 4th, 20-28 on 5th, 22-30 on 6th, and 28-32 on 10th proleg.
Material examined. — \ A specimens, 10 kilometers east of Indian Lake,
elev. 555 meters, lat. 43.45.30, long. 74.10. 14, Hamilton County, New York,
19 June 1977 (12 specimens) and 23 July 1977 (2 specimens), from ova of
females collected and determined by T. L. McCabe. All larvae, PI 9 ’s, and
reared adults are labelled tlm 77-15 9a (1st 9) and tlm 77-159 (2nd 9).
Larvae are deposited in the New York State Museum, the Illinois Natural
History Survey, and the United States National Museum.
146
NEW YORK ENTOMOLOGICAL SOCIETY
Literature Cited
Dyar, H. G. 1904. Lepidoptera of the Kootenai District. Proc. U.S. National Mus. 27(1376):
779-938.
Godfrey, G. L. 1972. A review and reclassification of larvae of the subfamily Hadeninae
(Lepidoptera, Noctuidae) of America north of Mexico. U.S. Dept. Agric. Tech. Bull.
1450, 265 pp.
McCabe, T. L. 1980. A reclassihcation of the Polia complex for North America (Lepidoptera:
Noctuidae). New York State Mus. Bull. 432, 141 pp.
McDunnough, J. 1 938. Checklist of Lepidoptera of Canada and the United States of America.
Pt. 1, Macrolepidoptera. South. Calif. Acad. Sci. Mem. 1, 275 pp.
Strecker, H. 1 900. Lepidoptera, Rhopaloceres and Heteroceres, Indigenous and Exotic, sup-
plement 3, p. 31.
(TLM) New York State Museum, Albany, New York 12230 and (GLG)
Illinois Natural History Survey, Champaign, Illinois 61820.
Received for publication November 5, 1981.
NEW YORK ENTOMOLOGICAL SOCIETY
XC(3), 1982, pp. 147-160
DESCRIPTIONS, SYNONYMY AND SEX ASSOCIATIONS IN
THE GENUS EUCERCERIS (HYMENOPTERA: PHILANTHIDAE)*
George R. Ferguson
Abstract.— E. geboharti, E. melleoides and E. sculleni, n. spp. are described
from western North America. E. barri Scullen is removed from synonymy,
and E. cavagnaroi Scullen is elevated from subspecies to species status.
Aphilanthops marginipennis Cameron is removed from synonymy with E.
rubripes Cresson and placed in synonymy with E. canaliculata (Say). The
male of E. brunnea Scullen and the female of E. melanovittata Scullen are
characterized. E. menkei Scullen is the female and a junior synonym of E.
melanosa Scullen.
Since Scullen’s (1968) revision of the genus Eucerceris much additional
material has become available. A study of this material has permitted rec-
ognition of three previously undescribed species, some new sex associations,
and new or revised synonymy involving other species. The results and con-
clusions presented here are a continuation of those in Ferguson (1981).
Eucerceris geboharti Ferguson, n. sp.
Figure 1
— Length 12-13 mm; forewing 8-9 mm; mandible with broadly
triangular tooth deflected inward, slightly recurved, weakly bicuspidate; ven-
tral margin of mandible swollen and elbowed slightly distad of middle, apical
margin of clypeal midsection with 2 rounded lobes medially, connected
basally, angled outward from plane of face (Fig. 1); preapical setae arising
from underside of apical lobes; disc of clypeal midsection convex; eyes
diverging below; mesopleural tubercle present; subalar carina strongly pro-
duced laterally into a roundly triangular lobe, projecting beyond tegula when
viewed from above; scutal punctures separated by 1 to 3 puncture diameters;
scutellum and metanotum polished with a few scattered punctures; propo-
deal enclosure polished with a few scattered punctures and/or weak ridges,
median groove linear; mesopleuron ridged between punctures, hypoepi-
' Oregon Agricultural Experiment Station Technical Paper No. 6197.
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NEW YORK ENTOMOLOGICAL SOCIETY
meron with separated punctures; metapleuron densely ridged; propodeum
densely striatopunctate except immediately adjacent to enclosure; second
submarginal cell of forewing petiolate anteriorly; punctures on anterior and
posterior ridges of terga II-IV coarse, well separated, polished between;
transverse depressions of terga II-IV with smaller punctures, closely but
irregularly spaced; hypopygium flat, with transluscent area; pygidium sub-
triangular, narrowly rounded apically.
— Length 10-11 mm; forewing 8-9 mm; mandible edentate with
small membranous lobe on lower side slightly distad of midlength; clypeal
margin with 3 rounded teeth of equal length; eyes slightly divergent below;
clypeus sparsely punctate; lower face moderately punctate, upper face and
vertex more or less contiguously punctate except punctures separated behind
ocelli; subalar carina and punctation of thorax as in female; forewing with
second submarginal cell sessile, marginal cell with a basoposterior pocket;
fimbriae loosely clumped on sterna III and IV, that on III narrower than on
IV; fimbria on sternum V denser, hairs shorter medially, somewhat irregular
in length; pygidium narrowed apically, longer than basal width, distance
between apicolateral teeth about V2 basal width, strongly incised inside each
lateral tooth with middle lobe wider than distance to tooth.
Coloration. — Much like Eucerceris bitruncata Scullen with which it has
been confused; scape, pedicel and basal 5 or 6 flagellar segments fulvous to
red, remainder of flagellum black; ground color of head mostly red in female,
mostly black in male, with several yellow marks; thorax with black back-
ground and numerous yellow markings, more abundant in male; legs variably
yellow, fulvous and red; wing veins yellowish to light brown, membrane
lightly yellow stained, lightly infuscated apically; tergum VI red in female,
pygidium infuscate apically; pygidium of male yellow to ferruginous.
Female, Utah, [Garfield County], Boulder, VI-28-75 (J. Alcock);
University of California, Davis.
Paratypes. — 4 males, 4 females as follows: UTAH: 2 females, same data
as holotype [UCD, OSUj; 2 males, 2 females, Garfield County, Boulder,
7,200 ft., June 5, 1966 (G. E. Bohart) [USU, OSUj; 1 male, Emery County,
July 20, 1921 [OSUj; 1 male, Emery County, Aug. 23, 1921 (Grace O. Wiley)
[OSU].
Other specimen. — \ male, UTAH; Sphinx, June 14, 1950 (G. F. Knowl-
ton), heavily cyanided [USU].
The sternal fimbriae of the males of E. geboharti are very much like those
of Eucerceris provancheri (Dalla Torre). The fimbria on sternum V is nar-
rower, more compact and the hairs are of more uniform length in the latter
species. The two broadly rounded median lobes of the apical clypeal margin
are diagnostic in the female. The strongly expanded subalar carina in both
sexes is unique in the genus. The mesopleural tubercle, unidentate mandible,
diverging eyes, and sparse punctation of the female, and the weak clypeal
VOLUME XC, NUMBER 3
149
teeth of the male place E. geboharti, n. sp. in the same species group with
Eucerceris barri Scullen, E. provancheri, and Eucerceris similis Cresson.
Eucerceris melleoides Ferguson, n. sp.
Figure 2
Eemale.—EQngXh 13 mm; forewing 10 mm; inner margin of mandible
with blunt tooth slightly distad of midlength; clypeus with median domelike
swelling, apicomedial margin with transverse, rectangular lamina beyond
base of preapical setae, lamina as long as pedicel width and 3 times as wide
as long, transverse apically, with poorly developed flanking tooth on each
side (Fig. 2); clypeal punctures irregular, mostly well separated, remainder
of head contiguously punctate; eyes diverging below; interocellar distance
scarcely 2 lateral ocellus diameters and about half ocellocular distance; pro-
podeal enclosure crossridged on anterior half, remainder polished; meta-
pleuron finely ridged; scutellum polished on anterior half, remainder punc-
tate; mesopleuron and propodeum contiguously punctate; mesopleural
tubercle absent; second submarginal cell of forewing petiolate anteriorly;
subalar carina not widened laterally; terga II-IV with dense, uniform, con-
tiguous punctures in transverse depressions well differentiated from polished,
more coarsely punctate anterior and posterior tergal ridges; sterna sparsely
punctate; hypopygium flat, transluscent area distinct; pygidium narrowly
suboval, twice as long as wide.
Afa/e. — Length 12-14 mm; forewing 9-1 1 mm; mandible edentate; clypeal
lip with 3 teeth of equal size; eyes slightly diverging below; ocelli and sculp-
ture of head as in female; forewing with second submarginal cell sessile,
marginal cell with basoposterior pocket; sculpture of thorax as in female;
sterna III-IV with numerous hairs but not differentiated into fimbriae; fim-
bria on sternum V about twice as wide as distance between apices of pygidial
teeth, hairs long, somewhat ragged at the tips, irregularly clumped; pygidium
with narrow median lobe projecting slightly beyond lateral teeth, distance
between median lobe and lateral tooth equal to or greater than width of lobe;
sculpture of abdomen as in female.
Co/ora^/o/t. — Background color black, pale markings yellow; 3 spots on
clypeus, 3 stripes on face, spot behind eye, pronotal ridge, metanotum, spot
on mesopleuron, spot on propodeal side, narrowly interrupted transverse
band on tergum I, narrow transverse band on posterior ridge of terga II-V,
lateral spots on sternum IV, yellow; mandible red, fulvous at base; scape,
pedicel and flagellomeres I-II red, remainder of flagellum black; coxae black,
remainder of legs mostly red; anterior Vs of forewing infuscated, remainder
of forewing and hindwing lightly smoky.
— Female, Mexico, Coahuila, Los Pinos, 19 mi. SE Saltillo, IX-
24-76, 6,800' (J. A. Chemsak, J. Powell, A. and M. Michelbacher). Uni-
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NEW YORK ENTOMOLOGICAL SOCIETY
versity of California, Berkeley. [To be deposited on indefinite loan at the
California Academy of Sciences, San Francisco.]
Paratypes.—4 males, all from Mexico, as follows: 1 male, same data as
holotype [UCBj; 1 male, Hidalgo, Zimapan, VI- 11/14-51, on flowers of
Eysenhardtia polystachya (Ort.) (P. D. Hurd) [UCB]; 2 males, Nueva Leon,
40 mi. W Linares, 5,200 ft.. Sept. 7, 1963 (Scullen and Bolinger) [OSU].
The long middle lobe of the pygidium separates the males of this species
from all others in the genus except E. melanosa Scullen and E. mellea
Scullen. E. melanosa has the fimbria on sternum V reduced to a pair of
pointed tufts medially, whereas the fimbria is distinctly transverse in mellea
and melleoides, n. sp. The female of melleoides, n. sp. is much like mellea,
but the apical margin of the clypeus is trilobed medially in mellea whereas
it is transverse in melleoides, n. sp. I have been unable to separate mellea
and melleoides, n. sp. males except by color; mellea is a red and yellow
species whereas melleoides, n. sp. is essentially a black and white species.
Eucerceris sculleni Ferguson, n. sp.
Figure 3
Eucerceris melanovittata, Scullen 1968:39, figs. 77a, b, d, f, females only,
nec Scullen 1939.
Fcma/c. — Length 14-16 mm; forewing 10-12 mm; mandible large, thick,
with blunt tooth having long axis subparallel to long axis of mandible; disc
of clypeal midsection with large protuberance, bluntly rounded apically with
apex closer to clypeofrontal suture than to apical clypeal margin and above
a line drawn between tentorial pits; lateral clypeal teeth large, bluntly tri-
angular, closer to each other than to eye margin; transverse lamella connected
to underside of lateral teeth, in front of and below preapical setae, with a
bluntly bidentate median elevation on lamella; clypeus and face densely
punctate, punctures essentially contiguous in area below midocellus; inter-
ocellar distance about half ocellocular distance; eyes diverging below; meso-
pleural tubercle indistinct; scutal punctures dense anteriorly, separated by 1
to 2 puncture diameters across middle Vs, surface polished; propodeal enclo-
sure polished, weakly sculptured, a few foveae in midline and a few weak
ridges anteriorly; subalar carina not laterally expanded; forewing with second
submarginal cell petiolate anteriorly; anterior and posterior ridges of middle
terga with large separated punctures, well differentiated from dense, small
punctures in transverse depressions; pygidium twice as long as midwidth,
narrowly rounded apically, slightly narrowed basally; hypopygium with large,
evenly rounded transverse swelling posteriorly (Fig. 3) with transverse or
broadly U-shaped carina at base, transluscent area absent.
Ma/G — Length 13-15 mm; forewing 10-11 mm; clypeal lip with strong
VOLUME XC, NUMBER 3
151
teeth, midtooth slightly longer; eyes slightly diverging below; ocelli and
punctation of head as in female; thorax punctured as in female except scutal
punctures separated by less than 1 puncture diameter; forewing with second
submarginal cell sessile, marginal cell with basoposterior pocket; sterna III-
IV without fimbriae; sternum V with sharply rectangular fimbria, hairs very
dense, uniform in length, fimbria about 3 times as wide as length of hairs;
pygidium transversely swollen at apical Vi with lateral carinae converging
from base to middle, subparallel over apical V2, median lobe not extending
beyond lateral teeth; punctation of abdomen as in female.
Coloration. S\m\\2ir in color pattern to red marked specimens of E. rub-
ripes Cresson; background color black, usually replaced in part by red; pale
markings usually bright yellow; 3 spots on clypeus, 3 lines on face with
middle one more or less reaching midocellus, spot behind eye, sometimes
spots behind ocelli, pronotal collar and lobes, hypoepimeron, scutellum,
metanotum, propodeal sides, spots behind mid and hind coxae, incised or
interrupted band on tergum I, complete transverse band on posterior ridge
of terga II-V, complete or partial transverse band on anterior ridge of terga
II-V, transverse band or lateral spots on one or more of sterna II-IV, yellow;
scape, pedicel and basal 2 to 4 flagellomeres reddish, remainder of flagellum
black; legs red, coxae sometimes infuscated; forewing infuscate along anterior
V3 and distal %. Males colored like females except legs marked with yellow.
Holotype.—FemsiXe, Willcox, [Cochise County], Arizona, July 7, 1956 (A.
D. Telford); University of Arizona, Tucson. [Deposited in the California
Academy of Sciences, San Francisco.]
Paratypes. — 1 6 males, 6 females, as follows: ARIZONA: 1 male, 2 females,
same data as holotype [UAZ, OSU] [Note: one of these females [OSU] carries
a Eucerceris melanovittata Scullen determination label and a yellow Plesio-
type label affixed by Scullen.]; 1 male, Cochise County, 3 mi. SE Willcox,
VIII-29-57, on Cleome sp. (W. F. Barr) [UID]; 1 male, Cochise County,
Willcox, VIII- 18-58 (P. D. Hurd) [UCB]; 2 males, Graham County, 12 mi.
S Stafford, 4,250 ft., IX- 14-62 (H. A. Scullen) [OSU]; 1 male, Navajo County,
1 mi. SW entr. Navajo Nat. Mon. VIII-2-67 (D. C. and K. A. Rentz) [UCB];
1 male, Pima County, Baboquiviri Mts., near Kits Peak, VIII-7/9-1916,
about 3,600 ft., [AMNH]; 2 males, Pima County, Baboquiviri Mts., VIII-
15-1924, VIII- 18- 1924 (O. C. Poling) [CAS]; 1 male, Pima County, Nogales
Springs, Whetstone Mtns., VIII- 17-77, 4,400^,600 ft. (Werner, Olson, Hetz)
[UAZ]; 1 female, Pima County, 3 mi. SE Continental, VIII-26-76 (S. Kuba)
[CDA]; 1 male, Pima County, Greaterville, X-8-80 (B. F. and J. L. Carr)
[ALB]; 1 male, Santa Cruz County, 5 km N Nogales, VIII- 17-79 (J. v. d.
Vecht) [UCD]; 1 male, 1 female, Santa Cruz County, 5 mi. E Nogales, IX-
1-70 (R. M. Bohart) [UCD]; COLORADO: 1 female, Costilla County, San
Luis, VIII- 11-76 (B. F. and J. L. Carr) [ALB]; NEW MEXICO; 1 male.
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NEW YORK ENTOMOLOGICAL SOCIETY
Hidalgo County, Skeleton Cyn., VIII- 12-65 (G. W. Forister) [UCR]; 2 males,
Santa Fe County, Santa Fe, VII- 14-34 (F. E. Lutz), IX-2-34 (P. E. Geier)
[AMNH]; TEXAS: 1 female, Jeff Davis County, Davis Mts., IX-4-44 (F.
Wemer and W. Nutting) [MCZ]. [Note: This specimen carries a Eucerceris
melanovittata Scullen determination label, a yellow Plesiotype label, and an
orange “D” label, the latter indicating that the specimen was used for the
illustration of E. melanovittata in Scullen (1968).]
Other specimen. — \ female, Mexico, Chihuahua, La Campana, IX-8-73
(W. J. Hanson, B. A. Haws) [USU].
The strongly swollen hypopygium separates the female of E. sculleni, n.
sp. from all other species in the genus; otherwise it is very similar to E.
melanovittata except for the larger size and partly red markings of the former.
The sharply rectangular fimbria on sternum V and lack of fimbriae on sterna
III and IV separate the males of E. sculleni, n. sp. from its congeners except
for E. melanovittata and E. sinuata Scullen. E. sinuata has the anterior
transverse ridges of the midterga hnely punctate, similar to the punctures
of the transverse depressions, whereas in the other two species the anterior
transverse ridges are polished between well separated large punctures, the
anterior ridges being similar to the posterior ridges. Males of E. sculleni, n.
sp. are very similar to males of E. melanovittata morphologically. Males of
sculleni, n. sp. are larger (13-15 mm) than males of melanovittata (11-13
mm), and the latter are black and white whereas in sculleni, n. sp. there are
variable amounts of red and the pale markings are yellow. The scape and
pedicel are red in sculleni, n. sp. but black or piceous in melanovittata.
E. sculleni, n. sp. has not been collected in the Portal area of Cochise
County, Arizona, where E. melanovittata is relatively common. The spec-
imen from the state of Chihuahua, Mexico, is black and white, but it has
the strongly swollen hypopygium characteristic of E. sculleni, n. sp.
Eucerceris barri Scullen (Revised Status)
Eucerceris barri Scullen, 1968:19. [Female holotype, Jacob’s Cabin, Hart
Mtn., Lake County, Oregon; U.S. Natl. Mus. Nat. Hist., Washington, D.C.]
Eucerceris similis, Bohart and Grissell 1975:33, nec Cresson, in part; Bohart
and Menke 1976:592, in part; Krombein 1979; 1740, in part.
Bohart and Grissell (1975) synonymized Eucerceris barri Scullen with
Eucerceris similis Cresson presumably because of the similarity of the males
of the two species. Bohart has recognized barri in subsequent determinations.
Females of the two species are quite distinct in clypeal structure. E. similis
has four teeth on the apicomedial margin of the clypeus with all teeth equally
separated from each other. E. barri lacks the two median teeth, and the
apicomedial emargination of the clypeus is bounded by a carina or rim which
connects with the lateral teeth. Each lateral tooth is formed by a blunt.
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153
triangular expansion of the apical rim and the tooth is sometimes bicuspi-
date. In E. similis the preapical setae arise from the apical clypeal margin,
whereas in E. barri they arise from underneath the apical rim.
Males of barri and similis are separable by differences in antennal col-
oration. In similis the flagellum is black or infuscate above and at least the
apical 3 to 5 segments are black below. In barri the flagellum is fulvous to
light brown below for its entire length and partly to entirely pale above. In
about 75% of the male specimens of barri the pale facial maculations are
fused above the antennal sockets and almost fused in the remainder. In
similis the black stripes above the antennal sockets are of more or less
uniform width with no strong tendency toward fusion of the pale stripes.
The pale markings of barri are whitish, whereas they are usually distinctly
yellow in similis. The pale markings are much reduced in barri females, as
the face and head are often completely black, the sterna are black, and the
femora are black with an occasional apical pale spot.
Material examined. — males including 8 paratypes and a male collected
with the holotype female; 29 females including the holotype and 4 paratypes.
The California distribution data are given in detail since these records are
the southern extension of the range of this predominantly northern species.
County records are given for the remaining states with elevations when stated
on the labels. CALIFORNIA: 1 female, Lassen County, 4 mi. S Ravendale,
VIII- 10-59 (J. A. Chemsak), paratype [UCB]; 2 males. Mono County, Crooked
Creek Lab., White Mtns., 10,150 ft., VII- 19-61 (G. L. Stage) [UCB]; 1 male.
Mono County, Blanco’s Corral, White Mtns., 10,150 ft., VIII-25-60 (P. D.
Hurd) [UCB]; 3 males. Mono County, Cottonwood Cr., 9,300 ft., VII- 10-
61 (H. V. Daly) [UCB]. IDAHO: Camas, Cassia, Clark, Custer (5,899 ft.),
Fremont, Jefferson, Jerome, Lemhi, Oneida and Teton Counties [UCD,
UID, OSU, USU]. NEVADA: Lander and White Pine Counties [UCD,
UID]. OREGON: Lake County [OSU, USNM]. UTAH: Rich County [USU].
WYOMING: Albany, Carbon (8,800 ft.), Natrona (7,000 ft.), Sublette,
Sweetwater, Teton, Uinta (7,000 ft.) Counties and Yellowstone National
Park [UCB, UCD, CSU, COR, NEB, OSU, USNM]. In addition, Scullen
(1968) reported a male from Moffat County, Colorado.
Collection dates are July 2 to August 28 with the exception of one male
collected in Lemhi County, Idaho, on June 21.
Eucerceris brunnea Scullen
Eucerceris brunnea Scullen, 1948:159. [Female holotype, Jacala, Hidalgo,
Mexico; Mus. Comp. Zool. Cambridge, Massachusetts]; Scullen 1968:22;
Bohart and Menke 1976:591.
Eucerceris velutina, Scullen 1948:160, males in part; Scullen 1968:69, males
in part.
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NEW YORK ENTOMOLOGICAL SOCIETY
The males of Eucerceris brunnea Scullen have not heretofore been char-
acterized. A series of specimens collected by H. E. Evans at Guadalajara
Mexico, in 1959 and 1965 [COR, OSU, USNM] contained females of both
E. brunnea and Eucerceris velutina Scullen together with a number of super-
ficially similar males which had been determined as E. velutina by Scullen.
It seemed probable that these males were not all conspecific, but that males
of both brunnea and velutina were present. Although females of brunnea
and velutina are easily separated by their markedly different clypeal structure,
the two belong to the same species group on the basis of wing venation and
other characters. A similarity in the males of the two species would therefore
not be surprising.
Since surface sculpture is only weakly sexually dimorphic in the genus
Eucerceris, characters based on sculpture are important clues in associating
the opposite sexes of a species. It was found that there were substantial
differences in surface sculpture between females of brunnea and velutina,
and that the series of males collected at Guadalajara, Mexico, separated
readily into two groups each of which corresponded in sculpture to one of
the two species of females with no intergradation between the two.
Both sexes of velutina have the anterior and posterior transverse ridge of
terga III-IV sharply differentiated in punctation from the transverse furrow
of the same tergum. The transverse furrow is densely, contiguously punctate
with small, mostly hexagonal punctures, whereas the transverse ridges are
polished between large punctures mostly separated by one or more puncture
diameters. Both sexes of brunnea have punctation similar to velutina on the
posterior tergal ridge and in the transverse furrow of the same terga, but the
anterior transverse ridge is punctured almost as densely as the furrow.
Males collected with velutina females in the absence of brunnea females,
and males collected with brunnea females in the absence of velutina females
show an identical pattern of tergal punctation.
The dense, black, “carpet-like,” velvety pile on the scutum of males of
brunnea and velutina is a highly unusual character state. Since Eucerceris
cavagnaroi Scullen, Eucerceris punctifrons (Cameron), and Eucerceris vio-
laceipennis Scullen belong to the same species group on the basis of female
wing venation, I strongly suspect that the males of these latter three species,
when found, will also possess this character.
Specimens examined.— The following records for E. brunnea Scullen are
all from Mexico: CHIHUAHUA: 2 males, 1 female, Cuiteco, VIII-9-69
[UCD]; HIDALGO: 1 male, 28 mi. SW Jacato, 6,300 ft., VI-23-57 [OSU];
JALISCO: 7 males, 4 females, Guadalajara, VII- 14-59 [COR], VII- 17/28-
65 [OSU, USNM]; MICHOACAN: 1 male, 1 female, 6 mi. NW Quiroga,
VII- 1 1-63 [UCD]. I have also seen the male holotype of Eucerceris velutina
Scullen at the U.S. National Museum of Natural History, Washington. It is
not at the California Academy of Sciences, San Francisco, as reported by
Scullen (1968).
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155
Eucerceris canaliculata (Say)
Aphilanthops marginipennis Cameron, 1890:105. [Male holotype, Atoyac,
Veracruz, Mexico; Brit. Mus. (Nat. Hist.), London.] New Synonymy.
Aphilanthops marginipennis Cameron was incorrectly synonymized with
Eucerceris rubripes Cresson by Scullen (1968) and should be added to the
synonymy of Eucerceris canaliculata (Say) as given by Ferguson (1981). The
type locality of marginipennis is outside the known range of rubripes, and
the original description of marginipennis does not fit rubripes.
I have studied the holotype male of marginipennis which consists of a
head and thorax without a gaster. Nevertheless, the available parts of the
holotype taken together with Cameron’s (1890) original description make
identification possible. Cameron described the fimbriae on sterna III and IV
as occupying the middle three-fourths of the sterna and stated that the hairs
were nearly as long as the segments. He described the fimbria on sternum
V as “stiff, comb-like, almost continuous,” and the abdomen as mostly
yellow except for the tergal depressions. This pattern of sternal fimbriae
occurs only in E. canaliculata (Say) and Eucerceris atrata Scullen. E. atrata
is a black and white species known only from elevations of 5,500 to 7,300
ft. in the state of San Luis Potosi, Mexico (Scullen 1968). Eucerceris conata
Scullen has a very narrow, stiff fimbria on sternum V, but the mandible is
rather strongly elbowed medially, quite different from the mandible of the
holotype of marginipennis, and the distribution is similar to that of rubripes.
The holotype of marginipennis has a black scutum and extensive black
markings on the thoracic pleura and propodeum. It is an example of the
dark color form of southern Mexico and Guatemala described by Scullen
(1968) as Eucerceris zimapanensis and previously synonymized with E.
canaliculata by Ferguson (1981).
In my previous paper (Ferguson 1981) I neglected to give the reasons for
deleting Cerceris cameroni Schulz from the synonymy of Eucerceris cana-
liculata (Say) as listed by Scullen (1951, 1968), Bohart and Menke (1976),
and Krombein (1979). Cerceris cameroni was proposed by Schulz (1906) as
a new name for the Indian species Cerceris canaliculata Cameron, nec Say,
nec Perez, and is a synonym of Cerceris bimaculata Cameron according to
Turner (1912). Bohart and Menke (1976) list C. cameroni Schulz in the
synonymy of both Eucerceris canaliculata (Say) and Cerceris bimaculata
Cameron, the latter being correct according to my interpretation.
Eucerceris cavagnaroi Scullen (New Status)
Eucerceris punctifrons cavagnaroi Scullen, 1968:53. [Female holotype,
Volcan de San Salvador, El Salvador; Univ. of California, Davis]; Bohart
and Menke 1976:592.
Scullen (1968) separated this taxon from Eucerceris punctifrons (Cameron)
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NEW YORK ENTOMOLOGICAL SOCIETY
on the basis of color characters. Examination of the types of E. punctifrons
and E. cavagnaroi shows that they are morphologically distinct species. In
E. cavagnaroi the apical clypeal lip is very narrow, not or scarcely wider
than the diameter of an antennal socket, and with a distinct emargination
at the base of the lip on each side. In E. punctifrons the apical clypeal lip is
wider than an antennal socket diameter and forms a shoulder on each side
at its base at which point the width of the lip is about as wide as the width
of the subantennal sclerite. The size of the apical clypeal lip in punctifrons
is intermediate between the very narrow lip of cavagnaroi and the very wide
lip of Eucerceris velutina Scullen, the lip being distinctly wider than the
subantennal sclerite in the latter (Figs. 4-6).
Differences also exist in tergal and propodeal punctation. In cavagnaroi
the anterior transverse tergal ridges of terga III-IV have dense punctures,
almost as dense as those in the transverse tergal depressions, with virtually
no interspaces of more than one puncture diameter. In punctifrons the ante-
rior tergal ridges have numerous interspaces of more than one puncture
diameter being somewhat intermediate between cavagnaroi and velutina.
The transverse tergal depressions are rather broad in cavagnaroi, about 6 to
8 punctures in width, whereas they are quite narrow in punctifrons, being
about 3 to 4 punctures in width. The propodeum of cavagnaroi is densely
punctured with virtually no interspaces of more than one puncture diameter,
whereas the propodeum of punctifrons has numerous interspaces of more
than one puncture diameter.
Material examined. — WoXoXypQ female of Eucerceris punctifrons cavag-
naroi Scullen [UCD]; holotype female of Aphilanthops punctifrons Cameron
[BMNH]; 1 female of E. cavagnaroi Scullen, El Salvador, Santa Tecla, 638
m, 28-VI-1974 (C. G. Dean, B. M. 1974-366) [BMNH].
Eucerceris melanosa Scullen
Eucerceris melanosa Scullen, 1948:163. [Holotype male, Tehuacan, Puebla,
Mexico; U.S. Natl. Mus. Nat. Hist., Washington]; Scullen 1968:38; Bohart
and Menke 1976:591.
Eucerceris menkei Scullen, 1968:42. [Holotype female, 10 mi. NW Tama-
zulapan, Oaxaca, Mexico; Univ. of California, Davis]; Bohart and Menke
1976:591. New Synonymy.
Scullen (1968) suggested that Eucerceris menkei might be the female of
Eucerceris melanosa Scullen. A male and female collected together in the
state of Oaxaca, Mexico, were available to him [OSU]. In addition I have
seen a male collected with two females in the state of Puebla, Mexico [UCB].
The two species are of similar size, color pattern and surface sculpture,
and they are the only two species in south central Mexico for which opposite
sexes have not been identified. These facts and observations taken together
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157
Figs. 1-6. 1, 2. Head of female of E. geboharti, n. sp. (1) and E. melleoides, n. sp. (2); head
width 3.1, 3.5 mm respectively. 3. Side view of terminal gastral segments of E. sculleni, n. sp.
female; length of hypopygium 1.2 mm. 4-6. Clypeus of female E. cavagnaroi Scullen (4), E.
punctifrons (Cameron) (5), and E. velutina Scullen (6); clypeal width 2.9, 2.9 and 3.2 mm
respectively.
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NEW YORK ENTOMOLOGICAL SOCIETY
with the collection records leave no doubt that melanosa and menkei rep-
resent opposite sexes of the same species.
Specimens examined.— ThQ following records are all from Mexico.
HIDALGO: 1 female, Zimapan, VII- 14-68 [UCD]; 1 male, 5 mi. W Pachuca,
7,900 ft. VIII-25-62 [OSU]; MEXICO D.F.: 1 male, 2,300 m, VIII-26-28
[COR]; OAXACA: 1 male, 1 female, 8 mi. SW Oaxaca, Monte Alban ruins,
6,000-6,500 ft., VIII-23-63 [OSU]; PUEBLA: 1 male, 2 females, 2 km E
Tecamachalco, VII-4-74 [UCB]; 1 female, 275 km NE Chapulco, VIII-2-65
[COR]; 1 male paratype, 18 mi. W Tehuacan, 6,200 ft., IX-5-57 [OSU]; 1
male paratype, Cacaloapan, IV-26-62 [OSU]; QUERETARO: 1 male para-
type, 41 mi. N Queretaro, 6,500 ft., IX- 19-63 [OSU].
Eucerceris melanovittata Scullen
Eucerceris melanovittata Scullen, 1948:164. [Male holotype, 25 mi. E El
Paso, Texas; Calif. Acad, of Sci., San Francisco]; Scullen 1968:39, males
only; Bohart and Menke 1976:591; Krombein 1979:1739.
Scullen (1948) described Eucerceris melanovittata from males, and later
(Scullen 1968) described what he thought was the female based on two
specimens of a species described earlier in this paper as Eucerceris sculleni.
The two males mentioned by Scullen (1968) as representing a darker form
of melanovittata belong to the species described earlier in this paper as
Eucerceris melleoides.
The true female of E. melanovittata was found among specimens carrying
Eucerceris arenaria Scullen determination labels. Several females with asso-
ciated males have also been identified in collections submitted for deter-
mination. E. melanovittata and E. arenaria are almost identical in size and
coloration, both being black and white species. In males of arenaria the
fimbria on sternum V is about % as wide as the sternum, and the hairs are
shorter medially; whereas in males of melanovittata this fimbria is narrower,
sharply rectangular and the hairs are of even length. The females differ in
several morphological characters as follows:
E. melanovittata
(1) Clypeal elevation with apex
above the middle
(2) Clypeal teeth with long axis
directed forward
(3) Face below midocellus densely
and regularly punctate
(4) Hypopygium shagreened, with-
out translucent area
E. arenaria
Clypeal elevation with apex below the
middle
Clypeal teeth with long axis directed
slightly outward
Face below midocellus irregularly
punctate with many interspaces
of one puncture diameter or more
Hypopygium with a transluscent area
VOLUME XC, NUMBER 3
159
(5) Propodeal enclosure smooth,
weakly sculptured
(6) Mandible with single blunt tooth
(7) Subapical clypeal process sepa-
rated from lateral teeth by
width of process
(8) Distance from lateral tooth to eye
margin greater than distance
between lateral teeth (ratio
1.5/1. 1)
Propodeal enclosure ridged over most
of surface
Mandible bicuspidate, small tooth
arising from base of larger tooth
Subapical clypeal process separated
from lateral teeth by more than
width of process
Distance from lateral tooth to eye
margin slightly less than distance
between lateral teeth (ratio 1.4/
1.5)
Specimens examined.— \ have studied the holotype male of Eucerceris
melanovittata Scullen [CAS] and the holotype female of Eucerceris arenaria
Scullen [CAS]. In addition I have seen 66 males and 32 females of E.
melanovittata [AMNH, AZS, CAS, CDA, UCB, UCD, COR, UFL, OSU,
PUC, USNM] with the following distribution: ARIZONA: Cochise, Gra-
ham, and Santa Cruz Counties; NEW MEXICO: Hidalgo, San Miguel and
Torrance Counties; TEXAS: Brewster, Culberson, El Paso, Hudspeth, Jeff
Davis and Presidio Counties. I have seen 1 male specimen from each of the
states of Coahuila and Nueva Leon, Mexico.
Eucerceris pimarum Cockerell and Rohwer
Ferguson (1981), following previous authors, incorrectly cited Rohwer as
the author of this species. Menke and Bohart (1979) pointed out that author-
ship of the species is Cockerell, T. D. A. and S. A. Rohwer 1908:326, in
Rohwer, S. A. 1908. New philanthid wasps. Can. Entomol. 40:322-327.
Acknowledgments
In addition to the individuals and institutions previously noted (Ferguson
1981), I wish to thank the following individuals and institutions for their
generous loan of specimens used in the course of the research reported here
[abbreviations in brackets as used in the text]: D. Shpeley, University of
Alberta, Edmonton [ALB]; M. C. Day, C. Vardy, British Museum (Natural
History), London [BMNH]; W. J. Pulawski, California Academy of Sciences,
San Francisco [CAS]; M. W. Wasbauer, California Department of Food and
Agriculture, Sacramento [CDA]; R. M. Bohart, R. O. Schuster, University
of California, Davis [UCD]; M. Hathaway, Museum of Comparative Zool-
ogy, Cambridge, Massachusetts [MCZ]; B. C. Ratcliffe, University of Nebraska
State Museum, Lincoln [NEB]; L. L. Eighme, Pacific Union College, Angwin,
California [PUC]; A. S. Menke, U.S. National Museum of Natural History,
Washington [USNM].
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NEW YORK ENTOMOLOGICAL SOCIETY
I am grateful for the generous advice and assistance given to me by Dr.
R. M. Bohart during a visit to the University of California, Davis, in March
1981.
I thank Bonnie T. Hall for preparing the illustrations.
Literature Cited
Bohart, R. M. and E. E. Grissell. 1975. California wasps of the subfamily Philanthinae. Bull.
Calif Insect Surv. 19:1-92.
and A. S. Menke. 1976. Sphecid Wasps of the World. Univ. of California Press,
Berkeley, ix + 695 pp.
Cameron, P. 1890. Insecta, Hymenoptera v. 2 (Fossores), pp. 65-128. In: F. D. Godman and
D. Salvin. Biologia Centrali-Americana. Taylor and Francis, London.
Ferguson, G. R. 1981. Synonymy and distribution records in the genus Eucerceris. J. N.Y.
Entomol. Soc. 89:172-183.
Krombein, K. V. 1979. Sphecoidea, pp. 1573-1740. In: K. V. Krombein, Paul D. Hurd, Jr.,
David R. Smith, and B. D. Burks. Catalog of Hymenoptera in America North of Mexico.
V. 2:1199-2209. Smithsonian Inst. Press, Washington, D.C.
Menke, A. A. and R. M. Bohart. 1979. Sphecid wasps of the world: errors and omissions.
Proc. Entomol. Soc. Wash. 81:11 1-124.
Schulz, W. A. 1906. Spolia hymenopterologica. A. Pape, Paderbom. 355 pp.
Scullen, H. A. 1948. New species in the genus Eucerceris with notes on recorded species and
a revised key to the genus. Pan-Pac. Entomol. 24:155-180.
. 1968. A revision of the genus Eucerceris Cresson. U.S. Natl. Mus. Bull. 268:1-97.
Turner, R. E. 1912. A monograph of the wasps of the genus Cerceris inhabiting British India.
J. Bombay Nat. Hist. Soc. 21:476-516.
Systematic Entomology Laboratory, Department of Entomology, Oregon
State University, Corvallis, Oregon 97331.
Received for publication January 4, 1982.
NEW YORK ENTOMOLOGICAL SOCIETY
XC(3), 1982, pp. 161-165
PERCHING BEHAVIOR OF CANTHON VIRIDIS
(COLEOPTERA: SCARABAEIDAE) IN MARYLAND ‘
Orrey P. Young
Abstract. — Canthon viridis Beauv. is shown to perch on leaf surfaces near
a food source (dung) before attempting consumption. It is hypothesized that
predator pressure by staphylinids has contributed to the development of this
behavior. Correlated with perching behavior in C. viridis is low population
density, relative absence of diurnal beetle competitors, observed predation
by Staphylinus maculosus Grav., the capability of feeding on old feces aban-
doned by staphylinid predators, and a long survival period without food.
Introduction
It has been known for quite a few years that numerous species of Scara-
baeinae (Coleoptera: Scarabaeidae) can be found perching on vegetation
above the forest floor (Ohaus 1900). Recent investigators have suggested
that perching behavior may function to partition resources among competing
species (Howden and Nealis 1978; Young 1978). All previous reports of this
behavior, however, originate from tropical areas (Halflter and Matthews
1966; Howden and Young 1981). I report herein observations of perching
by a species of Scarabaeinae in a north temperate (Maryland) forest and
present a hypothesis relating to such behavior.
Observations
As part of a two-year study of the arthropods associated with dung in
Maryland, pitfall traps baited with human feces were placed weekly in a 50
hectare oak-hickory forest 4 km NE of Clarksville, Howard County. On 14
May 1979, at 0900 hr, a trap on gently sloping terrain was freshly baited
and then observed for one hour from a distance of three meters upslope.
Five minutes after bait placement, one Canthon viridis Beauv. was observed
flying in a zig-zag manner upslope at a height of approximately 1 5 cm. When
the beetle was within one meter of the bait, it landed on a horizontal leaf
' Scientific Article No. A3 107, Contribution No. 6172 of the Maryland Agricultural Exper-
iment Station, Department of Entomology.
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to indicate this fact.
162
NEW YORK ENTOMOLOGICAL SOCIETY
surface 30 cm above the forest floor. For the next ten minutes it remained
inactive, facing the bait. At the end of this period the beetle flew directly to
the trap and was captured. No other scarabs were attracted to the bait during
the one hour observation period. This same sequence of behaviors was also
observed with single individuals of C. viridis in the same forest near fecal
material on 12 June and 2 July 1979, and on 6 and 9 June 1980. During
the two-year period of this study, no other scarabaeine speeies was observed
perching on leaf surfaces in this forest.
Laboratory Experiments
Starvation. — During the period June-Sept. 1979, individuals of several
dung beetle species were brought into the laboratory and maintained at
ambient temperature (20-30°C) in 473 cc glass mason jars with a screen top
and 4 cm of packed soil in the bottom. Individuals were exposed to cow or
human dung for 48 hr, then removed to a fresh container and monitored
daily for survival without food. Atomized water was routinely added to
maintain suitable moisture conditions. Five individuals of C viridis {x
length = 5 mm) were processed in this manner and lived an average of 32
days (R = 27-36). Similar-sized species of Aphodius did not live as long
under these conditions {A. lividus, x ^ 11 days, R = 1 1-20, n = 50; A. rur-
icola, X = 22 days, R = 12-25, n = 40).
Predation. — During the period June-Sept. 1979, individuals of Staphy-
linus maculosus Grav. {x length = 21 mm, n = 15) were brought into the
laboratory and maintained under the same conditions as in the starvation
experiments previously discussed. Food in this case, however, was small
dung beetles plaeed alive in each cage every second day. On ten separate
occasions, a single C. viridis was placed in a container with one S. maculosus
(six different individuals) and was consumed on the average in 40 minutes
(R = 15-85).
Discussion
In Maryland, Canthon viridis occurs uncommonly in forests at fecal mate-
rial and even more rarely at carrion (pers. obs.). Usually no more than two
individuals will co-occur at a specific food site. Other forest-inhabiting diur-
nal coprophagous scarabs are likewise quite rare, with the vast majority of
the coprophagous scarab guild— such as species of Aphodius, Ateuchus, Can-
thon, Copris, Geotrupes, and Onthophagus— occurring at night.
Areas of the neotropics where perching in scarabaeines has been reported
may contain many species displaying this behavior. At Rio Palenque, Ecua-
dor, of 35 species of scarabaeines captured at human feces, 1 1 species also
perched on leaf surfaces (Howden and Nealis 1978). In a feeding guild of
VOLUME XC, NUMBER 3
163
this size, as the authors suggest, perching behavior may indeed be an impor-
tant component of foraging strategies effecting resource partitioning. In a
Maryland forest, however, where only one scarabaeine species sits on leaves
during the day and other diurnally-active dung scarabs are rare or absent,
resource partitioning among competing species does not appear to be related
to perching behavior.
Daytime competition for dung in Maryland forests appears to occur pri-
marily among the Diptera, with many predaceous Coleoptera usually in
association. Some of the predatory beetles (Staphylinidae) are common, large
(> 15 mm), diumally active, and capable of consuming small and medium-
sized dung beetles. These predators include Creophilus maxillosus (L.), On-
tholestes cingulatus (Grav.), and Staphylinus maculosus. Laboratory exper-
iments show that at least S. maculosus can capture Canthon viridis quite
easily and completely consume one individual in as little as 1 5 minutes.
Perching behavior in C. viridis may be correlated most closely with param-
eters associated with predation. One of those correlations may be between
the age (and odoriferous nature) of dung and the presence of staphylinids
and scarabs. Fresh fecal material in Maryland forests is quickly utilized by
Diptera if produced during the day and by scarabs if produced at night.
Occasionally, dung can be found in the forest that has not been completely
utilized and is a week or more in age. Diptera and staphylinids will not be
present on this old dung, but C. viridis may occasionally occur. In one
experiment during July 1979, human feces left exposed on the forest floor
and observed daily was still intact after 17 days, when the first C. viridis
arrived and was captured. When the feces was collected after 2 1 days, two
more C. viridis were obtained. Several small Aphodius and Onthophagus
were also collected during the same period, but no staphylinids were obtained
after the eighth day. This data is consistent with the hypothesis that C. viridis
is minimizing predation by consuming food that is no longer attractive to
predatory staphylinids.
Species of dung scarabs that are competitively superior to other dung
beetles and have the highest population densities are usually unable to sur-
vive for long periods of time without food (Young, ms.). In a series of
laboratory starvation experiments, C. viridis could survive for 32 days whereas
several Aphodius species of similar size could survive for approximately 20
days without food. Diurnal species of Aphodius do not perch on leaves, can
be among the earliest arrivals at fresh dung, and probably reduce predation
pressure by burrowing immediately into dung and creating tunnels in the
soil underneath. Canthon species, however, may perch on leaves, can also
be among the earliest arrivals at fresh dung but may be among the last, and
are exposed to predation during the formation of a dung-ball and its sub-
sequent rolling away to a burial site. Although the capability of doing without
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NEW YORK ENTOMOLOGICAL SOCIETY
food for long periods of time may be merely a response to low food avail-
ability, it may also allow individuals of C viridis to choose the food site
with the lowest possible level of potential predation.
The strategy of perching near food, before consumption is attempted, may
be a technique for assessing the presence of predators. Although no direct
information is available, research on dung beetles in Panama suggests that
the detection of staphylinid predators may be of an olfactory nature. As
discussed in Howden and Young (1981), at least one species of Canthon can
detect the presence of another species of Canthon (that is competitively
superior) at a food site and will not attempt to obtain food until the superior
species departs. Detection of the allelochemic involved can occur at a dis-
tance of 15 cm or more. Some staphylinids are known to produce phero-
mones (Peschke 1978) and allelochemics (Schildknecht et al. 1976), and
experimental demonstration of olfactory communication between Canthon
species and potential staphylinid predators would not be a great surprise.
A number of criteria may need to be satished before a species such as
Canthon viridis can use perching as part of a predator-assessment strategy.
It should be able to detect the presence of predators, perhaps by olfactory
cues, at a food site. It may employ the complementary strategy of arriving
at food before and/or after potential predators arrive. It should be able to
survive without food for a sufficient period to allow adequate search time
for food sites without predators. And hnally, perching on leaf surfaces should
not increase the likelihood of predation. The hrst three criteria have been
considered, but the final one requires some comment. C viridis, like many
of the leaf-sitting scarabaeines, is brightly colored (green) and shiny. This
probably makes them particularly obvious to leaf-foraging visual predators
such as birds and lizards. It has been suggested (Poulson 1980) that some
species of Canthon in the tropics are distasteful to such predators and/or
belong to a large mimetic complex. This has yet to be determined for C.
viridis, in Maryland, but may function to minimize predation while sitting
on leaves.
Conclusion
Given that the nature of the evidence is largely circumstantial and infer-
ential, it still seems reasonable to suggest that a species such as Canthon
viridis, which is so vulnerable to staphylinid predation, would have evolved
a strategy for minimizing that predation. The hypothesis of predator assess-
ment presented herein has the virtue of being testable. Although the project
that produced these observations has been terminated, it is easy to envisage
a series of laboratory olfactometer experiments with C. viridis that could
demonstrate the presence or absence of staphylinid-detection capabilities.
Also, manipulations in the field at dung involving removal and introduction
VOLUME XC, NUMBER 3
165
procedures with predators and prey could give strong support to the hypoth-
esis. Finally, detailed field observations where diurnal forest dung beetle
populations are more abundant than in Maryland could also be corrobo-
rative.
References
Halffier, G. and E. G. Matthews. 1966. The natural history of dung beetles of the subfamily
Scarabaeinae (Coleoptera, Scarabaeidae). Folia Entomol. Mexicana 12-14:1-312.
Howden, H. F. and V. G. Nealis. 1978. Observations on height of perching in some tropical
dung beetles (Scarabaeidae). Biotropica 10:43^6.
and O. P. Young. 1981. Panamanian Scarabaeinae: taxonomy, distribution, and
habits (Coleoptera, Scarabaeidae). Contrib. Amer. Entomol. Inst. 18:1-204.
Ohaus, F. 1 900. Bericht uber eine entomologische Reise nach Central-brasilien. Stettiner Ent.
Zeitung 60:193-274.
Peschke, K. 1978. The female sex pheromone of the staphylinid beetle, Aleochara curtula. J.
Insect Physiol. 24:197-200.
Poulson, T. L. 1980. Multiple functions for odors of dung-rolling Canthon beetles in a wet
tropical forest. Presented at Entomol. Soc. Amer. National Meetings, Atlanta, GA.
Schildknecht, H., D. Berger, D. Krauss, J. Connert, J. Gehlhaus and H. Essenbreis. 1976.
Defense chemistry of Stenus comma (Coleoptera: Staphylinidae). LXI. J. Chem. Ecol.
2:1-11.
Young, O. P. 1978. Resource partitioning in a neotropical necrophagous scarab guild. Ph.D.
thesis, Univ. of Maryland, College Park. 228 pp.
Department of Entomology, University of Maryland, College Park, Mary-
land 20742.2
Received for publication January 4, 1982.
^ Present address: Southern Grain Insects Research Lab, USDA-SE-ARS, Tifton, Georgia
31794.
NEW YORK ENTOMOLOGICAL SOCIETY
XC(3), 1982, pp. 166-175
DESCRIPTIONS OF THE IMMATURE STAGES OF
MYNDUS CRUDUS (HOMOPTERA: FULGOROIDEA: CIXIIDAE)i
Stephen W. Wilson and James H. Tsai
Abstract.— T\iQ 5 immature stages of Myndus crudus Van Duzee, an appar-
ent vector of lethal yellowing disease of palms, are described and illustrated
and a key for separating nymphal instars is provided. Features useful in
distinguishing nymphal instars include the number of metatarsal segments
(2 in first through third instars, 3 in fourth and fifth instars), the presence
of a tooth on the profemora of fourth and fifth instars, and the increase in
body size, wingpad size, and number of pits during nymphal development.
Myndus crudus Van Duzee has been recorded from Florida south to Ven-
ezuela and west to Panama, central Mexico, and the southern tip of Baja
California (Kramer 1979). This planthopper has been implicated as a vector
of lethal yellowing disease of coconut palms {Cocos nucifera L.) (Howard
and Thomas 1980; Tsai 1980; Tsai and Thomas 1981).
The eggs of M. crudus are laid in moist soil adjacent to grass stolons or
palm roots. The nymphs feed at the stem bases and roots of several species
of grasses, sedges, and palms (Reinert 1977; Tsai et al. 1976; Tsai and Kirsch
1978). Both adults and nymphs are phloem feeders on coconut palm (Fisher
and Tsai 1978). At least nine species of palms as well as several species of
grasses are feeding hosts of M. crudus (Reinert 1977; Tsai 1978). Although
M. crudus has been studied extensively in the laboratory and field (Reinert
1977, 1980; Tsai and Kirsch 1978), including laboratory rearing (Tsai et al.
1976; Tsai and Kirsch 1978), the eggs and nymphs of this species have not
been illustrated or described. Tsai and Kirsch (1978) provided measurements
of the eggs and nymphs but not detailed descriptions. Zenner and Lopez
(1977) published measurements and figures of eggs and nymphs of Haplaxius
pallidus Caldwell, a synonym of M. crudus (Kramer 1979).
There are few available descriptions of the immatures of any Cixiidae and
none of any species of Myndus. Cumber (1952) described the immatures of
Oliarus atkinsoni Myers, and Myers (1929) described the third, fourth, and
fifth instars of Mnemosyne cubana Stal and the fifth instar of Bothriocera
^ Fla. Agric. Exp. Stn. Journal Series No. 3786.
The publication costs of this article were defrayed in part by page charge payment. This article
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to indicate this fact.
VOLUME XC, NUMBER 3
167
signoreti Stal. The fifth instar of Oliarus placitus Van Duzee is being described
by Wilson (unpublished).
This paper includes descriptions of, and keys for separating, the five imma-
ture stages of M. crudus.
Materials and Methods
Specimens to be described were obtained from laboratory stock main-
tained by Tsai, and preserved in 70% ethyl alcohol. The description of each
stage is based on 10 specimens. The first instar is described in detail, but
only major changes from previous instars are described for subsequent instars.
Comparative statements refer to previous instars (e.g., “darker”). Dimen-
sions of eggs and nymphs are expressed in mm as mean ± SE. For nymphs,
length was measured from tip of vertex to tip of abdomen; width was mea-
sured across the widest part of the body, usually the mesothoracic segment.
Thoracic length was measured along the midline from the anterior margin
of the pronotum to the posterior margin of the metanotum; this measurement
was included because total length measurements are affected by differences
in head shape among specimens, and because the abdomen often becomes
distended when preserved in ethyl alcohol. Specimens of each instar were
cleared in 10% KOH in order to observe some structures (e.g., tergite of first
abdominal segment).
Descriptions of Immature Stages
Egg{¥ig. 1).-Length 0.54 ± 0.032; width 0.17 ± 0.030.
Eggs laid singly; elongate, subcylindrical; white; chorion translucent,
smooth; anterior end asymmetrical and pointed, posterior end broadly
rounded.
First Instar {¥ig. 2). — Length 0.64 ± 0.088; thoracic length 0.24 ± 0.041;
width 0.26 ± 0.041.
Form elongate, subcylindrical, slightly flattened dorsoventrally, widest
along mesothorax. Vertex, frons, thoracic nota, and abdominal tergites with
a few, shallow, indistinct pits.
Vertex broadly rounded anteriorly, widest in anterior V2, slightly narrowing
posteriorly. Frons subquadrate, lateral margins slightly convex, dorsal mar-
gin highly convex, juncture with clypeus obscure. Clypeus narrowing distally.
Beak 3 -segmented, extending just beyond metacoxae; segment 1 obscured
by clypeus, segments 2 and 3 subequal. Eyes reduced, barely visible in ventral
view, red. Antennae 3-segmented; scape and pedicel cylindrical and sub-
equal; flagellum bulbous basally, filamentous distally, bulbous portion sub-
equal in size to pedicel.
Thoracic nota divided by a longitudinal mid-dorsal line into 3 pairs of
plates. Pronotum longest medially; each plate subrectangular, anterior mar-
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NEW YORK ENTOMOLOGICAL SOCIETY
Figs. 1, 2. Immature stages of M. crudus. (1) Egg, (2) 1st Instar. Vertical bar = 0.5 mm.
VOLUME XC, NUMBER 3
169
gin almost straight to ca. level of lateral margin of eye then extending pos-
terolaterally, lateral margin slightly convex, posterior margin slightly sinuate.
Mesonotum with median length subequal to that of pronotum; each plate
subrectangular, lateral margin convex, posterior margin broadly curved.
Metanotum with median length ca. % that of mesonotum; each plate sub-
rectangular, lateral margin convex. Pro- and mesocoxae posteromedially
directed; metacoxae smaller, obscured by trochanters. Tarsi 2-segmented,
divisions between segments very obscure; segment 1 somewhat wedge-shaped;
segment 2 subconical, slightly curved, with a pair of slender apical claws.
Abdomen 9-segmented, subcylindrical, widest across segments 2 and 3;
segment 9 elongate vertically, surrounding anus.
Second Instar (Fig. 3). — Length 1.01 ± 0.103; thoracic length 0.40 ±
0.054; width 0.40 ± 0.052.
Antennae with bulbous portion of flagellum ca. % length of pedicel.
Pronotum with each plate bearing 10-15 shallow pits. Each plate of meso-
notum bearing ca. 7 pits with 3 pits extending anterolaterally from postero-
medial corner and 4 pits near lateral margin; posterolateral corners of plate
slightly lobate. Each plate of metanotum bearing ca. 4 pits with 2 pits near
medial border and 2 pits near lateral margin.
Third Instar {Fig. 4). — hQngth 1.29 ± 0.104; thoracic length 0.51 ± 0.048;
width 0.55 ± 0.073.
Vertex somewhat less rounded anteriorly; pits in 2 irregular rows, more
distinct; posterior and lateral margins distinct, slightly carinate, and sinuate.
Frons with 2 irregular rows of pits bordering lateral margins; lateral margins
almost straight, narrowing distally, juncture with clypeus distinct, ventral
margin concave; juncture between anterodorsal postclypeus and postero-
ventral anteclypeus straight and apparent laterally. Antennae with bulbous
portion of flagellum ca. Vi length of pedicel.
Pronotum with each plate bearing ca. 20 large, distinct pits. Each plate of
mesonotum bearing 12-13 large, distinct pits with 1 pit in the anteromedial
comer, an oblique row of 3-4 pits (usually 4) extending anterolaterally from
posteromedial corner, 7-8 pits near lateral margin; posterior margin dis-
tinctly lobate in lateral Vi. Each plate of metanotum bearing 7-8 pits with
1-2 pits near medial border, 2-3 pits ca. midway between medial and lateral
margins and 2-3 pits near lateral margin.
Abdominal tergites of segments 1 and 2 reduced, not extending to lateral
margins. The following number of pits on either side of midline of each
segment: segment 3 with 2 pits on tergite, segments 4-5 each with 5 pits on
tergite, segment 9 with 3 caudal pits. Tergites 6-8 each with a pair of enlarged
subtriangular, dorsoposteriorly oriented waxpads in intermembranous area
posterior to narrow tergite (waxpads probably present but indistinct in pre-
vious instars); each waxpad with a transverse row of 3 very small, obscure
pits near anterodorsal margin.
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NEW YORK ENTOMOLOGICAL SOCIETY
Figs. 3, 4. Immature stages of M. crudus. (3) 2nd Instar, (4) 3rd Instar. Vertical bar = 0.5
mm.
VOLUME XC, NUMBER 3
171
Fourth Instar (Fig. 5).— Length 2.20 ± 0.162; thoracic length 0.81 ±
0.068; width 0.92 ± 0.074.
Vertex, frons, thoracic nota, and abdominal tergite light gray-brown; pits,
intermembranous areas, clypeus, beak, antennae, legs, and sternum white.
Head with pits on vertex and frons more numerous. Antennae with scape
reduced, ca. V3 length of pedicel, bulbous portion of flagellum ca. V3 length
of pedicel.
Pronotum with each plate bearing 25-30 pits. Each plate of mesonotum
bearing 22-24 pits with 1-2 pits in anteromedial corner, an oblique row of
5 pits extending anterolaterally from near posteromedial corner, and 13-15
pits near lateral margin and extending onto wingpad; each wingpad covering
ca. %-% of each metanotal plate laterally. Each plate of metanotum bearing
9-13 pits in the following arrangement: 3 pits near anteromedial border, 3-
4 pits ca. midway between medial and lateral margins and 3-6 pits near
lateral margin. Distal % of profemora with slender tooth on median aspect
of ventral margin. Metatibiae with setae in longitudinal rows on ventral
aspect (present in previous instars but not apparent); distal setae somewhat
stout, almost toothlike or spinelike. Metatarsi 3 -segmented, segments 1 and
2 cylindrical, segment 3 subconical, slightly curved and bearing a pair of
slender apical claws; segment 3 slightly longer than segment 1, segment 2
ca. % length of segment 1.
Abdomen with each segment bearing the following number of pits on
either side of midline: segment 2 with 1 pit on tergite near midline (obscure,
not illustrated), segment 3 with 2 pits on tergite near midline and 1 pit near
lateral margin, segments 4-5 each with a transverse row of 7 pits extending
from near midline to lateral margin, segments 6-8 each with 2 pits on tergite
near lateral margin, segment 9 with 4 caudal pits. Waxpads on segments
6-8 each with a transverse row of 4 very small pits near anterodorsal margin.
Fifth Instar 6). — Length 2.68 ± 0.122; thoracic length 1.17 ± 0.059;
width 1.31 ± 0.077.
Sclerotized portions of body darker.
Head with pits more numerous on vertex and frons.
Each pronotal plate bearing 31-33 pits. Mesonotal plates bearing 25-29
pits with 2 pits in anteromedial corner, an oblique row of 6 pits extending
anterolaterally from near posteromedial corner, 1 7-2 1 pits on wingpad and
near lateral margin of plate; wingpads extending to or beyond apex of meta-
notal wingpad. Each metanotal plate bearing 8 pits with 3 pits in antero-
medial comer, an oblique row of 4 pits extending anterolaterally from near
posteromedial margin, and 1 pit near region overlapped by mesonotal wing-
pad; wingpads extending almost to fourth tergite. Profemora with stouter
ventral tooth.
Each abdominal segment bearing the following number of pits on either
side of midline: segment 2 with 1 pit on tergite near midline; segment 3 with
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NEW YORK ENTOMOLOGICAL SOCIETY
Figs. 5, 6. Immature stages of M. crudus. A. Nymph, B. Distal end of profemur. (5) 4th
Instar, (6) 5th Instar. Vertical bar = 0.5 mm.
VOLUME XC, NUMBER 3
173
Fig. 7. M. crudus nymph with waxy exudate.
2 pits on tergite near midline (lateral pit present in previous instar absent
in this instar); segment 4 with a transverse row of 10-11 pits on tergite;
segment 5 with a transverse row of 1 0 pits on tergite; segment 6 with 4 pits
on tergite laterally, segments 7-8 each with 2 pits on tergite laterally, segment
9 with 4 caudal pits. Waxpads on segments 6-8 each with a transverse row
of 5 very small pits near anterodorsal margin. Waxy exudate present in this
and earlier instars in living specimens (Fig. 7).
Key to the Nymphal Instars of M. crudus
1. Metatarsi 2-segmented; profemora lacking tooth on ventral margin
(Figs. 2-4) 2
- Metatarsi 3 -segmented; profemora bearing tooth on ventral margin
(Figs. 5, 6)
4
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NEW YORK ENTOMOLOGICAL SOCIETY
2. Posterolateral comers of mesonotum distinctly lobate; mesonotum
with more than 10 distinct pits on each side (Fig. 4) Third Instar
- Posterolateral corners of mesonotum not lobate or weakly so; meso-
notum with fewer than 10 pits on each side, pits shallow and often
indistinct (Figs. 2, 3) 3
3. Mesonotum with ca. 7 indistinct pits on each side; basal bulbous
portion of antennal flagellum distinctly smaller than pedicel; thoracic
length greater than 0.35 mm (Fig. 3) Second Instar
- Mesonotum apparently lacking pits; basal bulbous portion of anten-
nal flagellum subequal to pedicel; thoracic length less than 0.30 mm
(Fig. 2) First Instar
4. Mesonotal wingpads extending to apex of metanotal wingpads; meso-
notum with an oblique row of 6 pits on each side extending antero-
laterally from near posteromedial corner (Fig. 6) Fifth Instar
- Mesonotal wingpads not extending to apex of metanotal wingpads;
mesonotum with an oblique row of 5 pits on each side extending
anterolaterally from near posteromedial corner (Fig. 5) Fourth Instar
Acknowledgment
We thank Mr. Willey Durden, Aquatic Plant Laboratory, ARS, Southern
Region, USDA for producing Fig. 7.
Literature Cited
Cumber, R. A. 1952. Studies on Oliarus atkinsoni Myers (Hem. Cixiidae), vector of the
“yellow-leaf’ disease of Phorium tenax Forst. II. The nymphal instars and seasonal
changes in the composition of nymphal populations. New Zealand J. Sci. Technol. 34:
160-165.
Fisher, J. B. and J. H. Tsai. 1978. Feeding sites of leafhoppers and planthoppers on plant
tissues, p. 23. Proc. 3rd Mtg. Int’l. Council Lethal Yellowing. Univ. Florida Publ. FL-
78-2. 43 pp.
Howard, F. W. and D. L. Thomas. 1980. Transmission of palm lethal decline to Veitchia
merrillii by a planthopper Myndus crudus. J. Econ. Entomol. 73:715-717.
Kramer, J. P. 1979. Taxonomic study of the planthopper genus Myndus in the Americas
(Homoptera: Fulgoroidae: Cixiidae). Trans. Amer. Entomol. Soc. 105:301-389.
Myers, J. G. 1929. Observations on the biology of two remarkable cixiid planthoppers
(Homoptera) from Cuba. Psyche 36:283-292.
Reinert, J. A. 1977. Field biology and control of Haplaxius crudus on St. Augustinegrass and
Christmas palm. J. Econ. Entomol. 70:54-56.
. 1980. Phenology and density of Haplaxius crudus (Homoptera: Cixiidae) on three
southern turfgrasses. Environ. Entomol. 9:13-15.
Tsai, J. H. 1978. Vector studies in Florida, p. 24. Proc. 3rd Mtg. Infl. Council Lethal Yel-
lowing. Univ. Florida Publ. FL-78-2. 43 pp.
. 1980. Lethal yellowing of coconut palm: search for a vector, pp. 177-200. In: K. F.
Harris and K. Maramorosch (eds.). Vectors of Plant Pathogens. Academic Press, New
York. 467 pp.
VOLUME XC, NUMBER 3
175
and O. H. Kirsch. 1978. Bionomics of Haplaxius crudus (Homoptera: Cixiidae).
Environ. Entomol. 7:305-308.
and D. L. Thomas. 1981. Transmission of lethal yellowing mycoplasma by Myndus
crudus, pp. 211-229. In: K. Maramorosch and S. P. Raychaudhuri (eds.). Mycoplasma
Diseases of Trees and Shrubs. Academic Press, New York. 362 pp.
, N. L. Woodiel and O. H. Kirsch. 1976. Rearing techniques for Haplaxius crudus
(Homoptera: Cixiidae). Fla. Entomol. 59:41-43.
Zenner de Polonia, I. and A. Lopez. 1977. Apuntes sobre la biologia y habitos del Haplaxius
pallidus, transmisor de la “marchitez sorpresiva” en palma africana. Rev. Columb.
Entomol. 3:49-62.
(SWW) Department of Biology, Central Missouri State University, War-
rensburg, Missouri 64093 and (JHT) Agricultural Research and Education
Center, IFAS, University of Florida, Ft. Lauderdale, Florida 33314.
Received for publication April 12, 1982.
NEW YORK ENTOMOLOGICAL SOCIETY
XC(3), 1982, pp. 176-180
UNIONICOLID MITES FROM CENTRAL NEW YORK
R. A. Baker
Abstract. — yiiXQS of the family Unionicolidae were collected from 110
freshwater bivalves in New York. Five species of adult mite were found. Of
the dominant bivalves found in Conesus Lake, only Lampsilis siliquoidea
(Barnes) contained adult mites. Unionicola fossulata (Koenike) infected 84.8%
of this mussel and 75.3% of the nymphal and adult mites were found between
the gills. There were 2.08 ±0.12 mites per infected L. siliquoidea. At Spencer
creek, Unionicola arcuata (Wolcott) and Najadicola ingens (Koenike) were
found in Alasmidonta undulata (Say). Unionicola formosa-ypsilophora com-
plex (Vidrine 1980) and Unionicola tumida (Wolcott) occurred in Anodonta
cataracta (Say). In the case of N. ingens this is a new host record and the
first published report of its occurrence in New York State. No nymphal or
adult mites were found in 42 Elliptio complanata (Solander) from the two
contrasting sites.
Introduction
Unionicolid mites parasitize freshwater mussels. The family has been
studied by a number of workers in North America including Wolcott (1899),
Mitchell (1955, 1957, 1965a), Cook (1974), and Vidrine (1977, 1979, 1980).
In particular Mitchell (1965b) studied population densities and regulation
of Unionicola fossulata (Koenike) in Lampsilis siliquoidea (Barnes), and
Gordon, Swan and Paterson (1979) and Paterson and Macleod (1979) the
biology of Unionicola formosa (Dana and Whelpley) in Anodonta cataracta
(Say). Najadicola ingens (Koenike) has been studied by Humes and his co-
workers (1950, 1951, 1952).
Materials and Methods
Collections were made using a hand net in shallow water and by SCUBA
diving to reach a depth of 7.5 m the maximum at which bivalves occurred.
The mussel collections were examined as soon as possible on return to
the laboratory. The number and location of the active mites within each
mussel was noted and estimates were made of the numbers per host of eggs,
prelarvae and nymphochrysalids. Prelarvae and nymphochrysalids were dis-
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VOLUME XC, NUMBER 3
177
sected from the mantle and gills respectively. Specimens of all stages were
preserved in Koenike’s fluid.
Results
Five common species of freshwater mussels were collected from two con-
trasting sites.
Conesus Lake is part of the St. Lawrence River system. It is situated south
of Rochester and is the most westerly of the Finger Lakes system. A total
of 88 mussels were collected made up as follows: 46 L. siliquoidea, 38 Elliptio
complanata (Solander) and 4 Anodonta grandis (Say). The only adult mite
found was U. fossulata in L. siliquoidea. E. complanata contained the devel-
opmental stages of a transient species but no nymphal or adult mites.
Spencer Creek the out-flow channel of Spencer Lake is situated in the
town of Spencer, south of Ithaca and forms the northern part of the Sus-
quehanna River system. A total of 22 mussels were collected, in running
water of up to 1 m depth made up as follows: 1 1 Alasmidonta undulata
(Say), 7 A. cataracta and 4 E. complanata. N. ingens and U. arcuata were
found in A. undulata. Of the 7 specimens of A. cataracta 2 contained U.
tumida. The other 5 contained a species of Unionicola closely resembling
U. formosa (Dana and Whelpley) but the males have a discrete posterior
border on coxal plate IV as in Unionicola ypsilophora (Bonz). They have
been referred to as the U. formosa-ypsilophora complex by Vidrine (1980).
U. fossulata in L. siliquoidea. — \n L. siliquoidea 39 out of the 46 specimens
contained nymphal and adult stages of U. fossulata, a 84.8% level of par-
asitization by the active stages. In addition most contained unionicolid eggs
and prelarvae in the mantle and nymphochrysalids in the gills.
A total of 8 1 nymphs and adults of U. fossulata were found in the infected
specimens of L. siliquoidea averaging 2.08 ±0.12 mites per mussel. These
were made up as follows: 35 males, 41 females and 5 nymphs (Table 1). Of
the infected mussels, 89.7% had a male, never more than 1 per mussel and
61.5% had 1 male and at least 1 female. If 1 adult mite was present it was
Table 1. Nymphs and adults of Unionicola fossulata infecting the mussel Lampsilis sili-
quoidea.
Mite
Total numbers
Mean number
per infected host
Mean number
per total number host
Males
35
0.90 ± 0.05
0.76 ± 0.06
Eemales
41
1.05 ± 0.13
0.89 ± 0.13
Nymphs
5
0.13 ± 0.05
0.11 ± 0.05
All stages
81
2.08 ± 0.12
1.76 ± 0.15
46 L. siliquoidea examined, 39 infected with active stages.
178
NEW YORK ENTOMOLOGICAL SOCIETY
most frequently a male, if 2 mites were present they were normally male
and female and when 3 adults occurred in the mussel, 1 male and 2 females
were found.
The nymphal and adult mites of L. siliquoidea showed a preferred location
with respect to the gill surface. Table 2 refers to these results. The site
numbers are those used by Davids (1973). 75.3% of the mites occurred
between the gills, 17.3% between the gills and the foot and only 2.5% were
found between the outer gill surfaces and the mantle.
Measurements of the developmental stages indicated that 1 type of
Unionicola was present in E. complanata and 4 types in L. siliquoidea,
one of which is the same as that found in E. complanata (Jones, personal
communication). Identification to species is not possible since detailed
descriptions of most American unionicolid larvae are not yet available.
N. ingens in A. undulata.—N. ingens is an endobranchial parasitic mite
of freshwater bivalves. It belongs to the family Unionicolidae Oudemans
1909 and is the only genus and species in the subfamily Najadicolinae Viets
1935.
N. ingens was found only in A. undulata. 1 1 specimens of this bivalve
were collected, 8 were infected and a total of 1 5 adult N. ingens recovered.
The presence of N. ingens in A. undulata is a new host record. The present
report is also the first published account of N. ingens being found in New
York State although student project reports from “The International Field
Workshop on Aquatic Invertebrates,” 1978, St. Lawrence University, U.S.A,
listed N. ingens in 7 specimens of Strophitus undulatus from Grannis Brook
(St. Lawrence County, New York) as well as an unstated number of infected
Anodonta sp. (Crowell, personal communication).
N. ingens has previously only been recorded within the gills of its hosts.
In the present collection 7 out of a total of 1 5 mites were found inside the
pericardial region of 6 specimens of A. undulata.
Discussion
The dominant bivalves in Conesus Lake are L. siliquoidea, E. complanata
and A. grandis (Clarke and Berg 1959). The first 2 show an interesting
comparison with regard to parasitism by unionicolid mites. Although both
harbour immature resting stages, only L. siliquoidea had resident nymphal
and adult mites. E. complanata accommodates the transforming develop-
mental stages of a single species of unionicolid. Adult female mites must
visit this mussel in order to lay eggs and then leave. L. siliquoidea appears
to be the natural host for U. fossulata and the preferred host for other species
of immature transient unionicolids in this lake. Mitchell (1955) working in
Michigan, found four species occurred together in L. siliquoidea namely U.
VOLUME XC, NUMBER 3
179
Table 2. Distribution of Unionicola fossulata in infected Lampsilis siliquoidea.
Site in the mussel
Total numbers of active nymphs
and adults at each site
I
1
II
29
III
14
IV
32
V
1
VI
4
Site numbers are those used by Davids (1973).
fossulata, U. abnormipes, U. serrata and U. aculeata of which only the latter
was transient.
Adult mites were most frequently found between the gills and to a lesser
extent between the gills and the foot, which agrees with the work of Mitchell
and Pitchford (1953) for U. ypsilophora, Mitchell (1965b) for U. fossulata
and Davids (1973) but is in contrast to the findings of Gordon, Swan and
Paterson (1979) who worked on U. formosa. The latter workers frequently
found mites on the outer surface of the gills.
Although seasonal collections were not made both Mitchell (1965b) and
Gordon, Swan and Paterson (1979) found there were no significant seasonal
changes in the total number of mites in all stages or in the percentage
infection. The present results agree closely with the findings of Mitchell
(1965b).
A number of authors previously described N. ingens as rare or infrequent
but it has now been recorded from 16 states in North America, from Canada
and recently from Thailand (Vidrine, personal communication). New York
State can now be added to this list.
The new host reported here further supports the view of Vidrine and Bereza
(1977) that N. ingens, unlike the majority of unionicolids, lacks host spec-
ificity and parasitizes a broad spectrum of mussel genera.
Acknowledgments
This work was carried out whilst the author was a visiting faculty member
on the staff of the State University of New York at Brockport, U.S.A. I am
indebted to Professors Starr and Makarewicz for the facilities they provided
in the Department of Biological Sciences. Two Brockport students, Wesley
Byers and Andris Simsons, did the SCUBA diving and I am grateful for
their help and enthusiasm. Professor Berg kindly checked my identification
of the mussels and Dr. M. F. Vidrine confirmed my identification of the
mites.
180
NEW YORK ENTOMOLOGICAL SOCIETY
Literature Cited
Clarke, A. H. and C. O. Berg. 1959. The freshwater mussels of Central New York. Mem.
Cornell Univ. Agric. Exp. Sta., 367.
Cook, D. R. 1974. Water mite genera and subgenera. Mem. Am. Ent. Inst. 21:1-860.
Davids, C. 1973. The relations between mites of the genus Unionicola and the mussels
Anodonta and Unio. Hydrobiologia 41:37-44.
Gordon, M. J., B. K. Swan and C. G. Paterson. 1979. The biology of Unionicola formosa
(Dana and Whelpley): a water mite parasitic in the unionid bivalve, Anodonta cataracta
(Say) in a New Brunswick Lake. Can. J. Zool. 57:1748-1756.
Humes, A. G. and S. K. Harris. 1952. The clam hosts of Najadicola ingens (K.) Acarina in
a Quebec lake. Can. Fid. Nat. 66(3):83-84.
and H. A. Jamnback. 1950. Najadicola ingens (Koenike), a water mite parasitic in
fresh water clams. Psyche 5 7(3): 7 7-8 7.
and H. D. Russell. 1951. Seasonal distribution of Najadicola ingens (K.) (Acarina)
in a New Hampshire pond. Psyche 58(3): 1 1 1-1 19.
Mitchell, R. D. 1955. Anatomy, life history and evolution of the mites parasitizing freshwater
mussels. Misc. Publ. Univ. Mich. Mus. Zool. No. 89:1-28.
. 1957. On the mites parasitizing Anodonta (Unionidea; Mollusca). J. Parasitol. 43:
101-104.
. 1965a. New species of water mites {Unionicola) from Tennessee unionid mussels. J.
Tenn. Acad. Sci. 40:104-106.
. 1965b. Population regulation of a water mite parasitic on unionid mussels. J. Par-
asitol. 51:990-996.
and G. W. Pitchford. 1953. On mites parasitizing Anodonta in England. J. Conch.
London 23:365-370.
Paterson, C. G. and R. K. MacLeod. 1979. Observations on the life history of the water mite,
Unionicola formosa (Acari: Hydrachnellae). Can. J. Zool. 57:2047-2049.
Vidrine, M. F. 1977. New host records for two water mites (Acarina: Unionicolidae) (Abstract).
Association of South Eastern Biologists Bulletin 24(2):92.
. 1979. Unionicola (Pentatax) fossulata (Koenike, 1895) (Arthropoda: Acarina: Union-
icolidae) in Eastern North American fresh- water mussels (Mollusca: Bivalvia: Unionacea:
Unionidae: Lampsilinae: Lampsilini) (Abstract). Proc. La. Acad. Sci. 42:84.
. 1980. Systematics and coevolution of unionicolid water mites and their unionid
mussel hosts in Eastern United States. Ph.D. dissertation, Louisiana State University,
U.S.A.
and D. J. Bereza. 1977. Some considerations and implications of host specificity
studies of unionicolid mite parasites on the systematics of some groups of North Amer-
ican unionacean freshwater mussels. Bull. Am. Mai. Union. Inc., 85-86.
Wolcott, R. H. 1899. On the North American species of the genus Atax (Fabr.) Bruz. Trans.
Amer. Micros. Soc. 20:193-259.
Department of Pure and Applied Zoology, University of Leeds, U.K. and
S.U.N.Y. Brockport, U.S.A.
Received for publication January 21, 1982.
NEW YORK ENTOMOLOGICAL SOCIETY
XC(3), 1982, pp. 181-185
PREDATORS, PARASITES, AND ASSOCIATES OF
ANTHOPHORA ABRUPT A SAY
(HYMENOPTERA: ANTHOPHORIDAE)*
Beth B. Norden and Aubrey G. Scarbrough
— Predators, parasites, and associates of the gregariously nesting
solitary bee, Anthophora abrupta Say (Hymenoptera: Anthophoridae), were
studied in Baltimore County, Maryland during 1976-1979. Adult bees were
preyed on by bam swallows (Hirundo rustica L.) and parasitized by conopid
flies (Physocephala marginata Say). Immature stages were parasitized by
Chrysis sp., Ganperdea apivora (Aldrich), Hornia minutipennis Riley, and
Monodontomerus mandibularis Gahan. Juvenile mortality was high (about
57%), and may have been contributed to by 10 species of fungi (Eumycota)
infesting 50% of the bee cells. Fifteen non-parasitic insect species also nested
at the study site and possibly competed with A. abrupta for nesting space.
Introduction
Anthophora abrupta Say is a solitary bee that nests gregariously in clay
embankments or adobe structures. Each nesting female typically digs a hor-
izontal tunnel into the soil, then constructs and provisions a series of 6-8
cells. Each lipid-lined cell contains one egg laid on the semi-solid provision,
and is sealed with an earthen cap (Norden 1979). In this study, predators,
parasites, and inquiline associates of A. abrupta were recorded.
Materials and Methods
During 1976-1979, bees were observed as they nested in the adobe walls
of a farm shed in Owings Mills, Baltimore County, Maryland. Associates of
A. abrupta were monitored daily during the 1977 and 1978 seasons. Any
dead adult bees found at the site were collected in vials and returned to the
laboratory for examination. At the end of nesting in 1 978, 600 bee cells were
also taken to the lab where they were opened and inspected.
' This paper is derived from research conducted in partial fulfillment of requirements for the
Master of Science degree in the Department of Biological Sciences, Towson State University,
Towson, Maryland.
The publication costs of this article were defrayed in part by page charge payment. This article
must therefore be hereby marked “Advertisement” in accordance with 18 U.S.C. §1734 solely
to indicate this fact.
182
NEW YORK ENTOMOLOGICAL SOCIETY
Results and Discussion
Bam swallows {Hirundo rustica L.) were observed capturing adult female
bees as they flew above the shed on 23-30 May 1977. Swallows were not
observed foraging on bees after 30 May and it was suspected that the emer-
gence of worker bumblebees aflected this behavior. Three bumblebee species
(Bombus bimaculatus Cress., Bombus nevadensis Cress., and Bombus per-
plexus Cress.) that closely resemble A. abrupta also nested at the site. In
contrast to A. abrupta, bumblebees inflict painful stings and Batesian mim-
icry may have discouraged the swallows from preying on bees after 30 May.
It has been noted that gregarious nesters, especially Anthophoridae, are
frequently attacked by parasites (Callan 1977). In this study, five parasitic
insect species were found and are considered briefly below.
1) Chrysis sp. (Hymenoptera: Chrysididae): These metallic green cuckoo
wasps were abundant at the site (1 5 May- 10 Sept.), and frequently were seen
entering bee tunnels. Three sealed cells contained body fragments of adult
Chrysis, but no immature stages were found. Most cuckoo wasps are external
parasites of bee prepupae (Borror and DeLong 1971).
2) Ganperdea apivora (Aldrich) (Diptera: Anthomyiidae): The holotype
and allotype of this parasitic fly were obtained from A. abrupta cells collected
in Missouri in 1877 (Steyskal 1967). Adult flies were very common at the
study site between 3 May and 1 9 July. Their occurrence corresponded closely
to that of adult bees (13 May-15 July). Ganperdea flew close to the wall or
crawled along its surface, frequently entering bee nests. Oviposition probably
occurred while the cell was still being provisioned. One bee cell containing
six fly pupae was found to have been partially provisioned, then filled with
loose clay particles. Five sealed cells opened in the lab contained an average
of 5.6 ± 0.5 fly pupae but no remnants of bee larvae. Bee eggs were probably
killed and the provision consumed by G. apivora larvae.
3) Hornia minutipennis Riley (Coleoptera: Meloidae): Riley described this
beetle from specimens also obtained from A. abrupta cells in Missouri in
1877. During this study, four H. minutipennis were found within sealed A.
abrupta cells (April 1977 and 1978). One specimen was also removed from
a spider web in the shed on 14 June 1977. Rau (1930) reported that female
beetles laid their eggs in the cells in which they were reared. When the eggs
hatched, the mobile triangulins (first instar larvae) crawled out from the bee
cells and attached themselves to passing adult A. abrupta. They were then
carried by adults into cells where they fed on bee eggs and cell provisions.
Only adult meloids were found in this study, and all appeared to have
consumed the provision but not the cell lining (see Norden et al. 1980).
4) Monodontomerus mandibularis Gahan (Hymenoptera: Torymidae):
These minute metallic green wasps were abundant at the wall from May to
October. They mated at the wall following emergence, and then females
VOLUME XC, NUMBER 3
183
entered A. abrupta nests. Bee cells (n = 8) contained 25.5 ± 4.5 wasp larvae
which consumed bee prepupae, leaving only the prepupal skins. Evidence
of more than one generation per season was provided in the laboratory where
adult wasps continued to emerge during September.
5) Physocephala marginata Say (Diptera: Conopidae): One conopid fly
emerged from a dead adult female A. abrupta on 8 July 1977, and another
was collected as it flew in front of the shed on 23 June 1978. Conopids insert
their eggs into bee abdomens during flight (Askew 1971). Bees remained
active as fly larvae fed on their abdominal contents, and pupation of the
conopid did not occur until after the death of the host bee.
Also noteworthy was the emergence of 64 Pediobius williamsoni (Girault)
from the body of a dead female A. abrupta collected on 27 June 1976. These
wasps were probably parasitoids of a conopid parasitizing the bee (E. E.
Grissell, pers. comm.).
Bees may also have succumbed to fungal infections. Examination of cells
(n = 600) in the laboratory revealed that 343 (57.4%) were infested by fungi
(Eumycota). Of these, 50.2% were completely filled with fungal mycelia
which had destroyed brood and provisions, while 7.2% contained living
prepupae with fungal growth limited to cell caps. Fungi were identified as
follows:
Order Eurotiales
Sporothrix sp.
Order Moniliales
Aspergillus niger van Tiegh.
Cladosporium sp.
Fusarium sp.
Paecilomyces sp.
Sporodesmium sp.
Order Mucorales
Mortierella sp.
Mucor sp.
Rhizopus sp.
Order Sphaeriales
Kernia sp.
Fusarium is known to be a faculative parasite of bees (Batra et al. 1973),
however, we could not determine whether this or other fungi were responsible
for larval death. Bees killed by other causes might later have become infested
by saprophytic fungi.
Whatever the causes of death, we found a juvenile mortality rate of about
57%. Anthophora linsleyi Timberlake exhibited a similarly high mortality
rate (about 50%), while maintaining a stable population (Linsley and
184
NEW YORK ENTOMOLOGICAL SOCIETY
MacSwain 1942). Apparently, A. abrupta also produced sufficient cells to
offset this high juvenile loss since the population did not appear to change
significantly during this study.
In addition to those organisms directly effecting A. abrupta, we also found
several non-parasitic insect species in close proximity to active bee nests.
These associates may compete with A. abrupta for nesting space, as they
were found inhabiting old nest cavities. They included:
Anthophoridae
Anthophora bomboides Kirby
Apidae
Bombus bimaculatus Cress.
Bombus nevadensis Cress.
Bombus perplexus Cress.
Eumenidae
Ancistrocerus unifasciatus (Saus.)
Stenodynerus sp.
Symmorphus sp.
Megachilidae
Megachile latimanus Say
Osmia lignaria Say
Prochelostoma philadelphi (Robt.)
Pompilidae
Agenioideus humilis (Cress.)
Sphecidae
Crossocerus sp.
Trypoxylon clavatum Say
Trypoxylon kolazyi Kohl.
Vespidae
Polistes fuscatus F.
Acknowledgments
We thank Dr. and Mrs. Worthley and Mrs. Reese for access to the study
site on their property. We are indebted to Dr. L. R. Batra, Mycology Lab,
Plant Protection Institute, USD A, Belts ville, Maryland, for fungi identifi-
cation, and the following scientists with the Systematic Entomology Lab,
USD A, Beltsville, Maryland, for insect identifications: S. W. T. Batra—
Anthophoridae, Apidae, Megachilidae; R. D. Gordon — Meloidae; E. E. Gris-
sell — Eulophidae, Torymidae; A. S. Menke— Chrysididae, Eumenidae, Pom-
pilidae, Sphecidae, Vespidae; G. Steyskal— Anthomyiidae; and F. G.
Thompson — Conopidae. We also thank Dr. S. Batra and Dr. R. Schroder,
Benificial Insect Introduction Lab, IIBIII, USDA, Beltsville, Maryland, for
critically reading the manuscript.
VOLUME XC, NUMBER 3
185
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Askew, R. 1971. Parasitic Insects. Amer. Elsevier Pub. Co., Inc., N.Y. 316 pp.
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, S. Batra, H. Pales, A. Hefetz and G. Shaw. 1 980. Anthophora bees: unusual glycerides
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Rau, P. 1930. A note on the parasitic beetle, Hornia minutipennis Riley. Psyche 37:1 55-1 56.
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Steyskal, G. 1967. A key to the genera of Anthomyiinae known to occur in North America
north of Mexico, with notes on the genus Ganperdea Aldrich (Diptera: Anthomyiidae).
Proc. Biol. Soc. Wash. 80:1-8.
(BBN) Entomology Department, University of Maryland, College Park,
Maryland 20742 and (AGS) Department of Biological Sciences, Towson
State University, Towson, Maryland 21204.
Received for publication February 10, 1982.
NEW YORK ENTOMOLOGICAL SOCIETY
XC(3), 1982, pp. 186-212
ENTOMOLOGY IN THE PEOPLE’S REPUBLIC OF CHINA^
James H. Tsai
Introduction
L Development of Entomological Studies
A. Historical Overview of China’s Entomology
a. Study of Beneficial Insects
b. Study of Agricultural Pests
c. Insect Control Measures
d. Study of Insect Morphology and Biology
B. The Influence of Western Industrial Revolution in the 19th and
20th Centuries
C TheEraof 1950’s
D. Pre-Cultural Revolution Period (1960-1966)
E. Cultural Revolution Period (1966-1976)
F. Post Cultural Revolution Era
II. Development of Control Methods
III. Current Educational and Research Organizations in PRC
A. Institutes of Higher Education
1. Comprehensive Universities
2. Agricultural Colleges and Universities
3. Other Specialized Institutions of Higher Education
B. Institutes of Research
C. Undergraduate and Graduate Education
Conclusion
Acknowledgments
Literature Cited
Introduction
China is one of the oldest civilizations in the world. The span of Chinese
history is the evolution from half-million-year old Peking-Man to 20th Cen-
^ Ela. Agric. Exp. Stn. Journal Series No. 3918. Supported in part by the grants from the
Ministry of Agriculture, Beijing, The People’s Republic of China, CSCPRC, National Academy
of Sciences, Washington, D.C. and the University of Florida’s Faculty Development Program.
The publication costs of this article were defrayed in part by page charge payment. This article
must therefore be hereby marked “Advertisement” in accordance with 18 U.S.C. §1734 solely
to indicate this fact.
VOLUME XC, NUMBER 3
187
tury technologists. The earliest evidence of Chinese civilization and science
development provided by a series of archaeological findings is set at more
than 5,000 years ago. Under successive dynasties Chinese achievements in
literature, philosophy, art, and certain fields of science are among the highest
in the world. However, the advent of Western technologies at the turn of
1 9th Century had profound consequences for traditional China. In order to
accommodate with the outside world, China has gone through drastic changes
in political, economic, and scientific systems. As with other sciences in China,
entomology has evolved and reached to a level of glorious attainment in the
history. It would be highly inappropriate for anyone to discuss the current
entomology in The People’s Republic of China (PRC) without knowing the
history of Chinese entomology.
I. Development of Entomological Studies
A. Historical Overview of China’s Entomology
a. Study of beneficial insects. — Silkworm {Bombyx mori L.) (Lepidoptera:
Bombycidae): Sericulture and silk technology date back to the period of
Agriculture ca. 4,500-7,000 years ago, as evidenced by the recent discovery
of a Neolithic relic site in Zhejiang Province (Map 1), where the silk material
along with rice seeds unearthed were determined by C^"^ dating method to
be 4,728 ± 100 years old (Anonymous 1980; Chou 1980). The silk tech-
nology advanced further in Yin Dynasty (16th-llth Century B.C.) as the
silk cloth unearthed from the Yin tombs revealed both flat and in relief
patterns. During this period, the tortoiseshells used as paper were found
inscribed with the idiograms
I
denoting the characters silk, silkworm and mulberry tree respectively
(Chou 1980). The mass planting of mulberry and indoor rearing of silk-
worm were recorded in the Xia Dynasty (1711 B.C.) (Anonymous 1980;
Chou 1980). By the 9th Century, as a result of continuous breeding and
rearing of the silkworm, many books on the bionomics of this insect had
been published. The technique for storing the eggs of silkworm in low tem-
peratures was developed in the 4th Century A.D. The domesticated silk-
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worm, B. mori is thought to be evolved from the wild species B. mandarina
(Moore) (Chou 1 980). The silk road was built 138-126 B.C. in Han Dynasty,
but the records showed that the technology of sericulture was not introduced
in Turkey until the 6th Century A.D. It was introduced into Korea as early
as the 12th Century B.C. Because of geographical barriers its introduction
into Japan came as late as the 2nd Century A.D.
Other silk producing insects have also been utilized in China since the
12th Century B.C. The breeding of the other silkworms, Antherea pernyi
Guerin started in the 1st Century A.D.; and Semia cynthia pryeri Butler,
and Eriogyma pyretomm (Westw.) in the 17th Century A.D.
Honeybee {Apis cerana Fab.) (Hymenoptera: Apidae): The history of api-
culture is as old as sericulture in China. The tortoiseshells found in the Yin
tombs bore the inscriptions of the idiograms of bee (Chou 1980).
I
The commercial breeding and the teaching of beekeeping prospered 1 ,800
years ago (Anonymous 1980). The uses of beeswax for candles and pills
were known in the 7th Century A.D. Several monographs dealing with
bee morphology, biology, rearing techniques, social behavior, control of
natural enemies, honey extraction, and apiary management were published
during the period of 1273-1817 A.D.
Wax insect (Ericerus pela Chav.) (Homoptera: Coccidae): The wax of this
scale insect was first utilized in China in 1 300 B.C. In the next two centuries,
a great deal of information on its host ranges, distribution, biology, and
methods of extracting wax was published.
Lac (Laccifer lacca Kem) (Homoptera: Lacciferidae): During the 3rd Cen-
tury A.D., the Chinese were first to describe methods of distributing the
immatures of L. lacca on the tree in order to produce the lac which was
used for dye, lacquer, and medicine (Anonymous 1980; Chou 1980).
Gall insect {Melaphis chinensis Walsh.) (Homoptera: Eriosomatidae): As
early as the 1st Century B.C. the leaf galls produced by M. chinensis were
used for extracting tannin which has been used in dye and medicine (Anon-
ymous 1980; Chou 1980).
Insects as medicine: From 31 B.C. to 1578 A.D. a total of 73 species of
insects had been listed in Chinese medical publication entitled “Compen-
dium Materia Medica.” Several common insects are currently used in Chinese
medicine: the exuviae of cicadas are used against fever. Tannin extracted
from the leaf galls of M. chinensis is an astringent compound. Cantharadin
from the blister beetles, Lytta caraganae Pallas, Myabris phalerata Pallas,
and Epicauta spp., is used as a cure for ulcers and as a abortion agent. The
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189
egg cases of mantid are used for curing impotence. Silkworms infected with
muscardine fungus, Metarrhizium anisopliae Metsch, are used as a cure for
palpitation. Lepidopterous larvae parasitized by Cordyces sp. fungus are
used as a cough medicine, and the bee sting is used for treating arthritis
(Anonymous 1980; Chou 1980).
Edible insects: Historical records showed that men ate the larvae and
pupae of bees and wasps, the nymphs of cicadas, and the immatures of ants
in 1200 B.C. Locusts as food were recorded in 23 A.D. Currently the pupae
of silkworm and giant silkworm and the predaceous diving beetles (Cybister
spp.) are still being used as food by some people in China.
Insects used for other purpose: Insects have become the main theme in
numerous poems and paintings throughout Chinese history. May beetles and
buprestids were used as ornamentals in the 1 1th Century A.D. During 618-
905 A.D., cicadas and crickets were bred for song contests. Since then several
species of crickets {Scapsipedus aspersus (Walker), Homoeogryllus japoniciis
(Haan), Gryllus testaceus Walker, and Brachytrupes portentosus L.) were bred
for fighting contests. As a consequence there were at least five monographs
dealing with cricket identification, food preference, biology, fighting tech-
niques, and artificial rearing techniques published during 1265-1884 A.D.
b. Study of agricultural pests.— China has been an agricultural society for
more than 5,000 years. Undoubtedly, China has a long history of fighting
against various agricultural pests. A book written in the 1 1th Century B.C.,
entitled “Book of Poems” which dealt with a number of agricultural subjects
described the decree issued by the ancient ruler to mobilize peasants in insect
control, and to recommend insect control with fire. A pest control officer
was officially installed in Zhou Dynasty (ca. 240 B.C.). During this period
a series of control measures were employed such as the use of heat, lime,
plant ashes and insecticidal plants (Chou 1980).
Locusts (Locusta migratoria L.): The locust problem has plagued Chinese
agriculture for as long as Chinese history is dated. Probably it has evolved
with the cultivated crops, mainly Gramineae, for about 6,000 years. The
earliest evidence of locust and man association was found in the Yin Dynasty.
The tortoiseshells found in the Yin tombs have a number of idiograms
representing the character “locust.” The first antilocust decree was is-
used in 29 A.D. (Chou 1980). During the period of 707 B.C. to 1911 A.D.,
between 538 and 800 locust outbreaks were recorded (Anonymous 1977a;
Chou 1980). Since the 17th Century A.D. a number of publications have
appeared concerning the breeding sites, host ranges, morphology, biol-
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ogy, ecology, gregarious and migratory behaviors, and control measures of
this insect.
Lepidopterous insects: Stemborers (Diatraea verosata Walker, D. shari-
inensis Eguchi, Chilo simplex Butler) are currently considered to be the
second most important pests of graminaceous crops. Between 7 1 8 B.C. and
1911 A.D. about 50 serious outbreaks were recorded (Chou 1980). Army
worm (Leucania separata Walk.) had 49 outbreaks between 500-1700 A.D.
Other insects such as wild silkworm (Bombyx mandarina Moore), mul-
berry geometrid, {Hemerophila atrilineata Butler), angoumois grain moth
(Sitotroga cerealella Oliv.), black mulberry beetle (Chrysochus chinensis
Baly.), wheat midge (Sitodiplosis mosellana Gehin), and the cricket {Gryllus
spp.) were reported for a number of outbreaks during the period of 26-1839
A.D.
c. Insect control measures.— Physical control: The manual control of insects
was first described 1,800 years ago, the control of silver fish Ctenolepisma
(Lepisma) vilosa (Fabricius) was recorded 2,000 years ago. The use of fire
to control insects was recorded about 240 B.C. (Anonymous 1979).
Cultural control: Weeding and deep plowing were employed to control
insects in 239 B.C. Varying dates of planting and harvesting which played an
important role in reducing the insect pests were reported during 528-549
A.D. Other practices such as crop rotation, irrigation, and the use of resistant
variety were also recorded in the history.
Biological control: This practice has been used in China since 304 A.D.
The classic example is the use of predaceous ants (Oecophylla smaragdina
F.) to control a number of citrus pests such as leaf beetles, curculioes, scar-
abaeus beetles, and stink bugs (Anonymous 1980). The use of ducks to
control pests in paddy field dates back to the period of 16 1 1-1672 A.D. The
decree was issued by the ruler of Late Han Dynasty (948-980 A.D.) to protect
such insect predators as birds and frogs.
Chemical control: The use of plant ashes and lime to control the household
insects has been known for over 3,000 years. Mercury used for flea control
and the treatment of wheat seeds with arsenic for control of underground
insects were reported 2,000 years ago. Other organic compounds such as
aluminum and copper for flea control and sulfur for control of ornamental
insects were reported 1 ,000-1 ,500 years ago (Anonymous 1 980; Chou 1 980).
The use of a variety of insecticidal plants including Zingiber mioga, and
Illicium lanceolatum to control medical and storage insects was recorded
3,000 years ago. Other plants with insecticidal activities were: Aconitum
lycactorum, A. fischeri, Daphne genkwa, Chaenomeles sinensis, Ligusticum
sinensis, Artemisia scoparia, Incarvillea sinensis, Celastrus sp., Croton sp.,
Ruta sp., Stemona sp., Xanthium sp., Spirodela sp., Gleditsia sp. They
were used during 1000 B.C. to 1700 A.D. (Chou 1980).
d. Study of insect morphology and biology.— The earliest record of study-
VOLUME XC, NUMBER 3
191
ing insect morphology came in 250 B.C. when the Chinese described the
exoskeleton and endoskeleton of the insects. The morphological character-
istics of the planthopper, Lycorma delicatula white, and the louse fly, Hyp-
pobosca capensis Olfers were described in detail in 1116 and 1578 A.D.
respectively (Anonymous 1980; Chou 1980). At the same time, the phe-
nomenon of metamorphosis in mantids was reported. Later the metamor-
phosis of Lepidoptera (butterflies and moths) and caddisflies was reported
in 300 and 739 A.D. respectively. The parthenogenesis of insect was recorded
in 600 B.C. Color mimicry was first described in the 1st Century A.D.
The relationship between the prey and predator was first studied in 502
A.D. with sphecid wasps (including the members of Eumenidae, Sphecidae,
Trypoxylidae) carrying the borer larvae to their nests as food for their young
(Anonymous 1977a; Chou 1980). During the 7th Century it was reported
that not only the borer larvae but also spiders were used as food for the
young, and that the eggs of wasps were deposited on the prey. Others reported
that tachinid flies parasitized on the silkworms in the 15th Century A.D.
(Chou 1980).
B. The Influence of Western Industrial Revolution in the
19th and 20th Centuries
Although China has a glorious ancient history of science and civilization,
it was only at the turn of the 1 9th Century that the study of entomology as
part of plant protection was initiated as a modern scientific discipline. China
is a country built on agriculture, and will probably remain an agricultural
nation from some time to come. This is basically due to the fact that the
country covers an area of 9,6000,000 km^ (=3,706,000 mi^), and the current
population is about 1 billion. Two-thirds of the total area is mountainous
or semidesert; only 1 1% of the land is arable. Nearly 90% of the population
is concentrated on the fertile plains and deltas of the east which accounts
for Vg of the land. Geographically speaking, the country is in the Temperate
Zone with the exceptions of the southern portions of the country including
Yunan, Guangdong, and Guangxi Provinces which are within the Tropics.
Therefore, agriculture which provides enough foods and clothes has become
the major theme for every dynasty in Chinese history.
The industrial revolution and Western expansionism of the 19th Century
has brought China a series of military and political humiliations. Realizing
the inability of the old agricultural system in dealing either with internal
difficulties or with foreign encroachments, China started a series of reforms
in a society structured by about 5,000 years of civilization. Since then, China
has been repeatedly subjected to political, economic and intellectual chaos
and revolutions in order to accommodate with the modern world. Like most
other disciplines of science, entomology, in the broader field of plant pro-
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tection, has gone through many phases of changes. A systematic study of
entomological science started in 1911. During the period of 1922-1924, a
Bureau of Entomology was first established in Jiangsu and Zhejiang Prov-
inces. The entomologists in these two organizations engaged in the systematic
study of the major agricultural pests, and medical insects in southern China.
The first “Year Book of the Bureau of Entomology” and “Insects and Plant
Diseases” were both published by the Bureau of Entomology in Zhejiang
Province. Meanwhile they also had trained several scores of entomologists
who were later assigned to other provinces. Immediately after 1924, an
Entomological Institute similar to the Bureau of Entomology was formally
established in Hunan, Jiangxi, Guangdong, and Sichuan Provinces. Several
universities and colleges had established a Department of Plant Pathology
and Entomology, or Division of Entomology. At the same time, formal
teaching and research in entomology were instituted in the colleges and
universities. The prestigious Chinese Academy of Sciences (CAS) was formed
in 1930. Courses in entomology were included in the curriculum of the
Department of Agricultural Zoology in the comprehensive universities and
agricultural colleges.
Plant protection science in general suffered a lot from the impact of World
War I, and internally, China was all but shattered in the era of warlords.
Then came World War II. Most of the research institutes and universities
were forced to evacuate to the interior during the war, and very little progress
was made in the field of entomology during the 1 940’s. In spite of the difficult
situation during the war, a number of institutions maintained or re-estab-
lished a Department of Plant Pathology and Entomology usually in their
Colleges of Agriculture. Students in their junior and senior years would
specialize in either entomology or plant pathology, even though their degree
was awarded by the same department. Only Beijing University and Qinghua
University had established an independent Department of Entomology in
their College of Agriculture during the period 1945-1949.
C. The Era of 1950’s
After the founding of the People’s Republic of China in 1949, the country
enjoyed steady growth for a decade. The leaders of PRC had transformed a
weak and backward China into a strong and modern state. The progress of
Chinese economy and science in the years of rehabilitation following 1949
was very impressive. In 1953, the “First Five-Year Plan” was initiated in
PRC with emphasis on national defense and heavy industry. The economy
was basically still agrarian, with a small modern industry concentrated in
the northeast and east coast. Even though agriculture produced a greater
share of the GNP in this period, it still received little investment from the
state. Progress continued until the experiments of the “Great Leap Forward”
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193
(1958-1960) which featured backyard steel plants plunged China into a
depression in the early 1960’s.
Several outstanding accomplishments can be cited in this golden period.
In 1949, there were 450 million people in PRC with an average of less than
two years of education per capita. In a decade, the school enrollments quad-
rupled reaching 1 00 million and the average number of years of education
rose to 3.5 years per capita. Enrollment in primary school was compulsory,
therefore it became universal, and junior high enrollment was nearly uni-
versal in the cities and surrounding rural areas. The senior high and college
enrollments were rather restricted due to lack of facilities and there was
extreme competition for admission to these two levels. The Western school-
ing in China taking roots in this period was mainly due to the earlier influ-
ences of foreigners such as American, British, German, French and Japanese
in China before the liberation, and the influence of more than 30,000 Chinese
educated abroad. Immediately after the liberation, education was totally
taken out of foreign hands.
In 1954, higher education was reorganized and modelled on the Soviet
system, which emphasized science and technology and the topical special-
ization of institutions. As a consequence, separate Departments of Plant
Pathology and Departments of Entomology were combined into one Depart-
ment of Plant Protection. There were only two major courses in entomology
taught at college: General Entomology and Agricultural Entomology. The
former included morphology, anatomy, physiology, life history, ecology and
taxonomy. Under this system, no elective courses were offered at colleges
and the credit system was also abolished. The comprehensive universities
became the main sources to train college faculty. The graduate program was
also reorganized on the Soviet model. Roughly 16,000 students were trained
by Chinese institutions at the graduate level during 1955-1965. In the same
period, approximately 3,000 other Chinese students were trained at the
graduate level in USSR and Eastern European nations.
The Institute of Entomology was first established in 1 950 under the admin-
istration of CAS followed by the establishment of an Institute of Entomology
in several localities under the administration of different city and provincial
governments. Similarly, the Institute of Plant Protection was also established
in several places under the jurisdiction of the Chinese Academy of Agri-
cultural Sciences (CAAS) as well as city and provincial governments. Besides
the expansion of various research institutions as mentioned above, there
were noted accomplishments in the educational field. Courses in entomology
and plant protection were offered in the Biology Department of the com-
prehensive universities and the College of Forestry respectively. In Central
Government, the Bureau of Plant Protection was formed under the Ministry
of Agriculture. Consequently, the Plant Protection and Inspection Stations
functioned under various city and provincial governments. Plant Quarantine
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NEW YORK ENTOMOLOGICAL SOCIETY
Stations were set up at the major ports by the Foreign Trade Department.
At the same time, medical entomology received a great deal of attention;
several research institutes of medical entomology were brought into the
picture by the Health Department.
During the period 1953-1957, most of the China’s trained entomologists
were brought into the CAS, and its affiliated research institutes. The CAS
served as the focal point for planning and conducting research on a national
scale. The Institute of Entomology served as the center of all entomological
activities throughout the land. By decree of Mao Zedong, all scientihc research
had to point toward practical application. Therefore, the entomological pub-
lications during this era dealt, by and large, with applied research. None-
theless, some basic research of high quality was published by Chinese ento-
mologists trained abroad.
The hrst issue of “Acta Entomologica Sinica” and that of “Bulletin of
Entomological Society of China” were published in 1951 by the Entomo-
logical Society of China. The latter was renamed “Kunchong Zhishi”
(=“Knowledge of Insects”) in 1955. A total of 10 volumes of “Journal of
Economic Entomology” was published in the 1950’s. The Entomological
Society of China grew from 861 members in 1954 to 1,068 in 1958 with 22
branches throughout the country (Yueh 1958). By 1963, the number of
institutions of higher learning had doubled to 400 as compared to 1949,
with the enrollment up from 1 17,000 to 819,000 (Cheng 1963). Both under-
graduate and graduate training in entomology were offered by most of the
50 comprehensive universities and agricultural institutes. A monograph on
the standardization of entomological nomenclature was published in 1956
containing a comprehensive list of the names of insects in Chinese and in
Latin names (Anonymous 1956).
D. Pre-Cultural Revolution Period (1960-1966)
In 1 962, the Plant Protection Society of China was installed and its official
publication “Acta Phytophylacica Sinica” was issued subsequently. The fol-
lowing year a sister journal entitled “Zhiwu Baohu” (=“Plant Protection”)
was published to accommodate the influx of manuscripts submitted to the
society. During the period of 1962-1965, the Beijing Agricultural University
and Nanjing Agricultural College were designated as two “Key” institutions
by the Ministry of Education, specifically to train the college faculty and
researchers in the field of plant protection. The college curriculum was five
years with the additional year devoted to studying such entomological courses
as taxonomy, physiology, toxicology, ecology, and forecasting. In the last
year, the students were allowed to either specialize in plant pathology or
entomology. But this 5 -year college curriculum was reduced to four years a
year before the onset of Cultural Revolution.
VOLUME XC, NUMBER 3
195
The average number of school years reached 5.5 per capita in the pre-
Cultural Revolution period. As a consequence, a flood of graduates entered
the labor markets seeking the city and industrial jobs, and met with dis-
appointment. Meanwhile, the gaps between the better educated and the less
educated, and the cities and the rural areas continued to widen. The lead-
ership in PRC considered that these trends were very unhealthy. Thus the
government instituted the initial program of sending millions of youths down
to the country-side (Xiafang) in order to alleviate the tension and to increase
the manpower in rural areas.
E. Cultural Revolution Period (1966-1976)
In subsequent years, the full scale Cultural Revolution was carried out
which resulted in major changes in Chinese education and virtually all sectors
of Chinese society. The “Xiafang” movement was intensified as millions of
young people who had been enrolled in the school were sent to the rural
areas in order to emphasize learning through practice. Text books and cur-
ricula were designed to emphasize the needs of the area in which each
institution is located, especially if the subject is agriculture production. All
middle school graduates were required to work in the factories or on farms
for at least 2 years before applying for a recommendation to attend colleges.
Higher education was especially a prime target of attack, colleges and uni-
versities were completely closed for a period of about five years. From 1966
to 1978 graduate school training was indeed nonexistent. At the onset of
Cultural Revolution there were 100 research institutes, but that was reduced
to only 40 in 1976. By this time, China had only 1.2 million scientists,
engineers and technicians, less than 1% of the total population. When schools
reopened in 1970, the students were recruited for colleges solely on the basis
of applicant’s political record and family-class background instead of aca-
demic excellence. In the early 1970’s, college curricula were redesigned to
meet needs of production, they were shortened to three or two years, from
five years previously. Similarly the primary and secondary school curricula
were also reduced from six to five years each. No school records were kept
during the cultural revolution period.
It should, however, be noted that a great deal of applied research was
carried out during this period (Guyer 1977), and that some of the students
even though they were not selected on the base of academic excellence, were
indeed intellectually talented. Most of the practical publications that appeared
in this period were prepared by and credited to the editorial committee
instead of individual researchers. The editorial committee consisted of pro-
ducers, scientists, and administrators. Thus information in the publication
met practical needs and was technically sound, and the measures recom-
mended were also administratively feasible (Chiang 1977a).
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NEW YORK ENTOMOLOGICAL SOCIETY
The achievements in agriculture were impressive. The PRC was able to
attain a self-sufficiency in foods even though its population was enormous
and growing. During this period, the policy called for self-reliance at all
levels, from the top of government to small communes. The chief reason
for the success is in the intensive and efficient use of land through multiple
cropping and intercropping along with good water and soil management
practices, and resistant variety development. But multiple cropping and
intercropping systems led to complicated plant protection methods.
F. Post Cultural Revolution Era
Since 1976, educational policy has returned to a pre-Cultural Revolution
era committed to academic excellence. The examination system has been
reintroduced in the schools. High school graduates are no longer required
to work before entering the colleges. Because of a shortage of colleges and
universities, the competition in entrance examination remains very keen.
Thus only those who are well qualified academically as well as politically,
will be admitted to the colleges. To date, college curricula have been length-
ened to four years. In 1977 four modernization programs were initiated,
they include agriculture, industry, defense, and science and technology. It
was since further affirmed that top investment priority is given to the mod-
ernization of agriculture and science and technology (Reardon-Anderson
1978). The training in the secondary schools is heavily oriented toward the
natural sciences and mathematics. Foreign languages begins in the primary
schools and continues through middle school. The research activities for
faculty and student are now evident, and research facilities are being rapidly
procured and upgraded.
The current higher education in PRC can best be characterized as follows:
Since 1949, education is considered to be a major governmental responsi-
bility and a public enterprise. As in many socialist states, a private educa-
tional system is practically non-existent in PRC. The struggle for techno-
logical development and modernization in education is seen as a vital factor
to enable China to establish and maintain a position of importance in the
world. It is clear that science and technology are perceived as the fundamental
features of the modem education. However, the emphasis is placed on applied
science and technology rather than basic sciences.
II. Development of Control Methods
In the 1950’s chemical control played the predominant role in China’s
struggle against such insect pests as migratory locusts, corn borers, sugar
cane borers, rice stem borers, wheat midges, armyworms, cotton bollworms,
pine caterpillars, mites, spiders, and medical insects. During the period of
VOLUME XC, NUMBER 3
197
1953-1957, a total of 170,000 tons of insecticides was used on the cultivated
land, and about 6 million hectares of crops which covered 70% of infested
areas were treated for locust control by aerial dusting. In 1958 alone, an
average of 14.7 kg of indigenous insecticides per hectare was used on various
crops (Chiu 1959). Benzene hexachloride (BHC), DDT, and other organic
phosphorous compounds such as DDVP, malathion, Dipterex and Demeton
were the main pesticides used for control of major pests throughout China.
The use of native plant and mineral products as a supplement to the
imported insecticides was actively encouraged in 1958, about 500 native
products were made into 10 million tons of insecticides and fungicides (Chiu
1959). Two Derris spp. containing 13.5 and 10% rotenone and one nicotine-
bearing species. Anabasis aphylla were found and utilized (Cheng 1963).
The successful examples of using biological control were numerous in the
1950’s. The use of parasitic wasps, Trichogramma spp. for sugarcane borer
control in Guangdong and Guangxi Provinces was highly successful. By
1959, nearly 5,500 hectares of sugarcane plantations were under this control
program as compared to 480 hectares in the previous year. As a result of
biological control, the sugarcane yield increased by about */3 (Pu et al. 1956).
Another noteworthy development in biological control was the rearing tech-
niques for Trichogramma spp. and their biology study. It was found that
the wasps reared on the eggs of pine caterpillar and ricinus silkworm were
larger in size, more active, with a higher fecundity, and higher 9 : <5 ratio
than those raised on angoumois grain moth, Sitotroga cerealella Oliv. The
eggs of the hosts could be stored up to 2-3 months at 4°-0°C respectively
without ill effect on parasite rearing. The adult wasps fed on a honey diet
lived 8.6 times longer and produced 14.7 times more than those reared on
distilled water.
Other examples of biological control were equally successful including the
use of Dibrachys cavus Wilk. against the pink bollworm, the tachinid fly,
Zenillia roseanae B. B. against the rice leaf roller, the rice swarming cater-
pillar and the European com borer (Yang 1958), the lady beetles, Rodolia
cardinalis Mulsant and R. rufopilosa Mulsant against cottony cushion scales
(Chiu 1959). The muscardine fungus, Beauveria bassiana was reported to
be used against the soybean pod borer, the sweet potato weevil, and the pine
caterpillar (Hsu et al. 1959; Lin 1956).
In addition to chemical and biological control measures, other means of
insect control were reportedly employed with limited success. Early or late
sowing and transplanting of rice resulted in less borer damages (Chiu 1959).
Similarly, late sowing of wheat was reported to reduce the damage by wheat
stem maggot, Meromyza saltatrix L. in Shaanxi Province. The removal of
the weed, Leersia hexandra Swartz from the paddy fields was an effective
control of the rice gall-midge, Pachydiplosis oryzae Wood-Mason and the
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planthopper, Nilaparvata lugens Stal. The method of drowning the riee borer,
Tryporyza (=Schoenobius) incertulas (Walker) was practiced over a large
area (Chiu 1959).
In the 1950’s the age-old problem of migratory locust, Locusta migratoria
manilensis was effectively controlled by sophisticated environmental control
techniques. The problems were managed through installation of reservoirs
and drainage systems, transformation of water-logged areas, reclamation of
wastelands along the coast, and elimination of particular breeding habitats
such as lake shore, flooded plain, coastal areas, and river flooded areas (Ma
1958, 1962). An excellent example is that in a decade (1950-1960), a total
of 15,492 km of irrigation systems were built. As a consequence, 39,000
hectares were under flood control, and 370,500 hectares were placed under
irrigation system in Shangdong Province alone (Guyer 1977).
The system of monitoring and forecasting insect population became an
integral part of pest management in China (Anonymous 1977a, b, 1979b):
the program started in 1951 and by 1958 a total of 678 insect monitoring
and forecasting stations was in operation throughout the nation, with 700,000
farmers participating in the program (Su 1959). These stations were mostly
limited to short-range forecasting of insect outbreaks in large rural areas.
However, the information obtained from the study of population dynamics
of migratory locusts made long range forecasting possible.
The pests of major crops and their control measures are cited below.
1 . Rice insects: China is the world’s largest producer of many food crops
including rice, sweet potatoes, sorghum, soybeans, millets, barley, peanuts,
and tea. Of the 124 million hectares of rice land in Asia, about 35 million
hectares of rice were planted in China. The grain production in PRC was
estimated at 255 and 140 million metric tons in 1974 and 1981, respectively
(Kelman and Cook 1977; Anonymous 1982). About 80% of the rice is of
indica varieties; they are developed for certain qualities including high yield,
fast growth under high fertility, early maturity, and short stems. Since rice
is the dominant crop in China, pests are considered as important production-
limiting factors. Pest problems are more serious in the south than central
and northern China. Of the total 1 14 species of rice pests damaging the rice
during either the growing or post harvest seasons, the rice paddy borer,
Tryporyza incertulas (Walker); purplish stem borer, Sesamia inferens Walker;
rice stem borer, Chilo suppressalis Walker; green rice leafhopper, Nephotettix
cincticeps Uhler; brown planthopper, Nilaparvata lugens Stal; white-backed
planthopper, Sogatella furcifera Horvath; rice leaf roller, Cnaphalocrocis
medinalis Guenee; rice skipper, Parnara guttata Bremer and Grey; rice
weevil, Echinocnemus squameus Billb.; the planthopper, Laodelphax stria-
tellus Fallen; rice leafminer, Hydrellia griseola Fallen; rice thrip, Chloethrips
oryzae Williams, Haplothrips aculeotus Fab.; and rice green caterpillar, Na-
rangaaenescens Moore’, are the major ones {Anonymous 1977, 1978a, 1979a).
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199
The use of chemical control was solely based on its effectiveness, safety and
economy. The time to use insecticide was determined through the monitoring
and forecasting system. The monitoring procedures and determining eco-
nomic thresholds for various rice pests were described in recent reports
(Anonymous 1977b; Chiang 1977b).
Cultural controls played an important part in pest control; they included
the use of resistant varieties, removal of alternate hosts, planting trap crops,
alternation of planting dates, and flooding of rice fields. Biological control
agents such as Trichogramma spp.. Bacillus thuringiensis Berliner, ducks,
and frogs were used considerably for pest control. Light traps were used not
only as a monitoring device, but also for insect control. In addition to the in-
dividual control measures, the integrated pest management program was
most frequently used in crop protection (Anonymous 1977b, 1979b).
2. Cotton insects: China is the third largest cotton producing nation in the
world with annual production at 11.1 million 480-lb. bales in 1981 (Anon-
ymous 1982). The cotton producing areas include 19°-45° north latitudes
and 75°-124° east longitudes (Anonymous 1979b) which cover Zhejiang,
Hubei, Sichuan, Anhui, Jiangsu, Shanxi, Shaanxi, Hunan, Yunnan, Guizhou,
Guangdong, Guangxi, Shangdong provinces (Kung 1975). Eight species of
insect and one spider mite were recognized as important pests: cotton aphid,
Aphis gossypii Glover; cutworms, Agrotis ypsilon Rott., A. tokionis Butler;
green plant bugs, Lygus lucorum Meyer-Dur., Adelphocoris suturalis Jak.;
pink bollworm, Pectinophora gossypiella Saunders; cotton bollworm,
Heliothis armigera Hubner; cotton leafhopper, Empoasca biguttula Ishida;
and two spotted spider mite, Tetranychus urticae Koch (Anonymous 1977b,
1979b). Monitoring and forecasting procedures for the above pests were
widely used like those described for rice culture (Anonymous 1977b; Chiang
1977a, b). The measure used to control these pests included integrated pest
control, good cultural practices, biological control, and chemical control
(Anonymous 1977b, 1979a, b). The biological agents used in integrated
control were Chrysopa septempunctata L. and five species of lady beetles
for control of cotton aphids, Dibrachys cavus Walker for control of pink
bollworm, Trichogramma confusum Viggiani, T. dendrolimi Matsumura as
well as B. thuringiensis for control of cotton bollworm (Anonymous 1977b,
1979b).
3. Wheat insects: Wheat is the second largest grain crop in China, the
production in 1981 was estimated at 54.2 million tons (Anonymous 1982).
Nearly 1 20 species of wheat pests consisting of 46 families and 1 1 orders
were reported (Anonymous 1977b). Among these, the wheat aphids, Macro-
siphum avenae Fab.; Rhopalosiphum maidis Fitch; Schizaphis graminum
Rondani; wheat armyworm, Mythimna separata Walker; wheat midges,
Sitodiplosis mosellana Gehin and Contarinia tritici Kirby; wheat stem mag-
gots, Meromyza saltatrix L. and Oscinella pusilla Meigen; wheat shoot mag-
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got, Nanna truncata Fan.; grubs, Hototrichia titanus Reitt, H. diomphalia
Bates, H. serobiculata Brenske, Anomala cuprea Hope; mole crickets, Gryl-
lotalpa africana Palisot de Beauvois and G. unispina Saussure; wireworms,
Agriotes patrualis Frivalsky, Pleonomus canaliculatus Faldermann and
Melanotus caudex Lewis were considered as major pests (Anonymous 1972,
1974, 1 979b). Monitoring and forecasting procedures for wheat aphids, wheat
midges and army worms as well as underground pests were developed (Anon-
ymous 1979b; Chiang 1977b), and the control measures for wheat insects
were similar to those mentioned above.
4. Soybean insects: Soybean is grown throughout China with annual pro-
duction at 7.9 million metric tons in 1981 (Anonymous 1982). The main
production area is in the northeastern part of the country. A dozen species
of insects are considered as major pests: soybean pod borer, Leguminivora
(Grapholitha) glycinivorella Matsumura; soybean borer, Maruca testulalis
Geyer; pea-pod borer, Etiella zinckenella Treitschke; soybean leafroller,
Heaylepta indicata Fah.; bean hawk moth, Clanis bilineata Walker; soybean
tussock moth, Dasychira locuple Walker; bean blister beetle, Epicauta gor-
hami Marseul; scarabaeid beetles, Holotrichia gebleri Faldermann, H. diom-
phala Bates, Anomala corpulent a Motsch, and Maladera orientalis Motsch;
bean shoot aphid. Aphis craccivora Koch; bean leafroller, Matsumuraeses
phaseoli Matsumura; bean weevil, Xylinophorus mongolicus Faust (Anon-
ymous 1977b, 1979b). Control measures for the pod borer are: cultural
control, resistant variety, chemical control, and biological control (Anony-
mous 1979b). For controlling other lepidopterous pests, the most commonly
used methods are light traps, chemical control, cultural control and biological
control (Anonymous 1977b, 1979b).
5. Other crops: Important pests of citrus fruits, deciduous fruits, vegeta-
bles, stored products and structures, man and animals and their control
methods were discussed in detail by earlier reports (Anonymous 1977b,
1979b; Guyer 1977; Williams 1979).
Other achievements in applied entomology during this period are listed
below.
1) Insecticide and its use: Scientists in PRC were very concerned about
the undesirable side effects of various pesticides on human health and the
environment. Therefore the emphasis of pest control was placed on inte-
grated pest control in 1970’s. Insecticides were carefully chosen and used
only if justified. Mostly pesticides were used as a part of integrated pest
management program. In this period, the consideration was given to the
production of highly effective insecticides of low toxicity, nonpersistance,
and low cost. Many organophosphorus insecticides were produced indus-
trially. The most commonly used were trichloiTon, dichlorvos, dimethoate,
phosmet, fenitrothion, phosphamidon, and malathion. Trichlorfon was used
against the cotton pink bollworm, bollworm, rice planthopper and thrips.
VOLUME XC, NUMBER 3
201
and cabbage worms. Dichlorvos was used to control soybean pod borer,
cabbage root maggot, cotton pink bollworm, cotton aphid, and other species
of aphids, housefly, mosquito, and citrus longicorn beetles. Dimethoate was
used for control of the citrus leafminer, leafrollers, mites, rice greenleafhop-
per, rice thrips, rice paddy borer, and cotton aphid. Phosmet was used to
control the tea scale and citrus leafminer. Fenitrothion was used for rice
paddy borer, riee leafhopper and planthopper (Anonymous 1977b, 1979b;
Guyer 1977).
2) Biological control agent and its use: It was the government policy that
all research must be directly applied to the real needs of people. Biological
control fits very well in this concept and has received substantial support
from the government. It has become the main feature of the integrated pest
management program. The most widely researched and used biological con-
trol agents included Trichogramma spp., Anastatus sp., Rodolia spp., Dibra-
chys cavus, Chrysopa spp.. Bacillus thuringiensis and Beauveria bassiana.
By 1974, a total of 12 species of Trichogramma was reported in China
including T. australicum Girault, T. closterae Pang et Chen, T. dendrolimi
Matsumura, T. evanescens Westwood, T. euproctidis Girault, T. ivelae Pang
et Chen, T. japonicum Ashmead (Anonymous 1978b), T. leucaniae Pang et
Chen, T. lingulatum Pang et Chen, T. ostriniae Pang et Chen, T. raoi Naga-
raja, and T. sericini Pang et Chen. Among these, only four species were
widely used in 26 provinces and regions of PRC, they included T. confusum
Riggiani {=T. australicum), T. dendrolimi, T. japonicum and T. ostriniae
for control of such pests as Ostrinia furnacalis Guenee, Chilo sacchariphagas
Bojer, C. infuscatellus Snellen, Argyroploce schistaceana Snellen, Dendro-
limus spp. and Heliothis armigera Hb. with parasitization 70-80% (Anon-
ymous 1978b, 1979b; Guyer 1977). Anastatus sp. was first used to control
litchi stinkbug, Tessaratoma papillosa Drury in 1960’s. It was reported that
the parasitization could reach to 80-90% (Anonymous 1979b; Huang et al.
1974). Their use has been extended to another 1 1 species of lepidopterous
insects and the eggs of several species of Hemiptera and Lepidoptera (Anon-
ymous 1978b). Dibrachys cavus was successfully used for control of the
overwintering population of the pink bollworm, Pectinophora gossypiella
Saunders with parasitization over 80%. Four species of Chrysopa (C sep-
tempunctata Wesmael, C. sinica Tjeder, C. carnea Stephens, C. boninensis
Okamoto) were commonly used for control of Aphis gossypii Glover, Tetran-
ychus urticae Koch, and the egg stage of Heliothis armigera and Ostrinia
furnacalis {Anonymous 1979b).
The establishment of the introduced Rodolia cardinalis Mulsant in south-
ern China had effectively controlled the cottony cushion scale, Icerya pur-
chasi Moskell in the citrus groves. Another species of lady beetles R. rufo-
pilosa Mulsant was also successfully used for control of I. purchasi in south
China (Anonymous 1979b).
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In the decade of 1 966-1976, a total of 1 7 varieties of Bacillus thuringiensis
Berliner including 12 distinct serotypes was isolated and characterized by
the researchers at the Institute of Zoology, Academia Sinica in Beijing (Guyer
1977). The bacteria were mainly used to control the immatures of Lepi-
doptera. There were more than 70 pest species listed with variable results
ranging from 30 to 100% kill. They were effectively used against such insects
as Cnaphalocrocis medinalis Guen, Paranara guttata Br., Tryporyza incer-
tulas Walk., Plutella xylostella (L.), Artogeia rapae (L.), Ostrinia furnacalis
Guenee, Dendrolimus punctatus and Heliothis arrnigera Hubn. (Anonymous
1979b). During the period of 1973-1977, about 1,100 metric tons of micro-
bial materials were used on about 12,800 hectares of cotton (Guyer 1977).
The use of Beauveria bassiana Vuill as a biological control agent was
introduced in 1971, in four years it was expanded to nearly 57,720 hectares
(Guyer 1977). This agent was especially effective against European corn
borer, O. furnacalis in the field with 80-90% kill (Hsiu et al. 1973). The
granular preparations of Beauveria were often used in combination with B.
t., Trichogramma and/or chemicals in the integrated control programs.
3) Integrated pest management: The practice of integrated pest control
(IPC) or integrated pest management (IPM) began in the 1950’s in China.
It became intensihed and popularized during the Cultural Revolution as it
was in line with the teaching of Chairman Mao who advocated that man
conquer nature. Thus IPM received top priority and strong governmental
support. The plant protection systems in PRC were largely based on IPC
techniques. Several integrated approaches were used in this highly successful
plant protection system. First, the scientists in PRC developed an efficient
monitoring and forecasting system which was set up at four levels including
provincial, county, commune and brigade. The provincial forecasting center
operated by the Academy studied the population dynamics, economic dam-
age thresholds of the pests, and the impact on natural enemy. All substations
reported to the provincial center. The county forecasting station determined
the time of occurrence of the pests, and advised the commune and brigade
what and when to carry out control operations. The commune forecasting
station operated at the farm level that consists of 5 to 15 brigade forecasting
stations. The commune forecasting station monitored the start of insect
activities in the spring as determined by light-trap catches, or monitored the
insect populations at the overwintering sites. The brigade forecasting station
monitored the target helds and relayed information to the commune (Chiang
1977b). The system served as a guide to timing chemical applications. The
insecticides were only used if proved economical, effective and safe. Sec-
ondly, cultural control is considered of vital importance in Chinese plant
protection system which emphasized prevention. This practice emphasized
the reduction in pest populations during the period between two crops. The
application of the biological and ecological information of the pest is essential
VOLUME XC, NUMBER 3
203
in this approach. There were several measures commonly used in the cultural
control including the use of light-traps for monitoring and mass trapping,
crop rotation, sanitation, regulation of planting and harvesting dates, the
use of a trap crop and bait trap. Thirdly, biological control was used suc-
cessfully and widely among pests of many crops including forest trees. This
approach is ranked second after cultural control at the production level. The
detailed biological control elements were described in the aforementioned
sections.
Other methods of insect control such as the use of resistant variety, sex
pheromones, hormones and various insect traps were also used to a consid-
erable extent in the integrated management schemes during the Cultural
Revolution period. For details readers should refer to the earlier publications
(Anonymous 1977b, 1979b; Guyer 1977).
III. Current Educational and Research Organizations in PRC
Since 1976, drastic shifts in policy have occurred. The new ideology
emphasized the four modernizations. The development of education and
research in science and technology became the national goal. New policies
are quickly implemented at every level.
A. Institutes of Higher Education
Currently PRC is restoring many higher institutions that were closed down
during the Cultural Revolution period, and establishing new institutions.
The number of institutions of higher learning is about 600 as compared to
430 in 1 966. The secondary schools and colleges and universities are divided
into “key institutions” and “non-key institutions.” The former are desig-
nated as priority institutions for development and receive generous appro-
priation of funds and attention, as well as enjoying higher prestige. They are
run by the Ministry of Education (MOE), the Ministry of Agriculture (MOA)
or the Ministry of Forestry (MOF). Generally speaking, the brightest and
best of the freshmen will go to key institutions. A total of 94 colleges and
universities has been designated as “key” institutions. A list of them which
offer courses in biological sciences or entomology is given in Table 1.
The basic structure of higher education in PRC has not changed much
since the early 1950’s when China patterned herself after the USSR. Most
Chinese institutions of higher education remain highly specialized in certain
fields.
1) Comprehensive universities: About 30 institutions of higher education
are in this category whose curricula include both basic and applied sciences
and the liberal arts. A number of well known institutions is listed in Table 1.
2) Agricultural colleges and universities: As mentioned before, basically
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NEW YORK ENTOMOLOGICAL SOCIETY
Table 1. Key universities and agricultural institutions in PRC.
Name^ Governing^ body Characterisitic
*Anhui Agricultural College
*Beijing Agricultural College
Beijing Agricultural University
Beijing Forestry College
Beijing Normal College
Beijing University
Central China Agricultural College
Central China University
Chinese People’s University
Chonqing University
Fudan University
*Fujian Agricultural College
Gansu Agricultural University
*Guangxi Agricultural College
*Guizhou Agricultural College
*Hebei Agricultural University
*Henan Agricultural College
*Hunan Agricultural College
*Hunan University
* Jiangsu Agricultural College
Jianxi Agricultural University
*Jilin Agricultural College
Jilin University
Jinan University
Lanzhou University
*Liaoning University
Nanjing Agricultural College
Nanjing Forest Industry College
Nanjing University
Nankai University
*Nei Monggol Agricultural College
Nei Monggol University
*Ningxia Agricultural College
Northwest University
Northwestern College of Agriculture
*Qinghai Industrial & Agricultural College
Qinghua University
*Shangdong Agricultural College
Shangdong University
Shanghai Agricultural College
Shanghai Normal University
*Shanxi Agricultural University
*Shenyang Agricultural College
*Sichuan Agricultural College
Sichuan University
South China Agricultural College
MOA
Agricultural
Beijing City
Agricultural
MOA
Comprehensive
MOF
Forestry
MOE
Comprehensive
MOF
Comprehensive
MOA
Agricultural
MOE
Agricultural
MOE
Comprehensive
MOE
Comprehensive
MOE
Comprehensive
PG
Agricultural
PG
Agricultural
PG
Agricultural
PG
Agricultural
PG
Agricultural
PG
Agricultural
PG
Agricultural
MOE
Comprehensive
PG
Agricultural
MOA
Agricultural
MOA
Agricultural
MOE
Comprehensive
MOE
Comprehensive
MOE
Comprehensive
MOE
Comprehensive
MOA
Agricultural
MOF
Forestry
MOE
Comprehensive
MOE
Comprehensive
MOA
Agricultural and
Animal Husbandry
MOE
Comprehensive
MOA
Agricultural
MOE
Comprehensive
MOA
Agricultural
MOA
Agricultural
MOE
Comprehensive
PG
Agricultural
MOE
Comprehensive
Shanghai City
Agricultural
MOE
Comprehensive
PG
Agricultural
PG
Agricultural
PG
Agricultural
MOE
Comprehensive
MOA
Agricultural
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205
Table 1. Continued.
Name^
Governing*> body
Characteristic
*Southwestem Agricultural College
MOA
Agricultural
Southwest University
MOA
Agricultural
Tianjin University
MOE
Comprehensive
Tongji University
MOE
Comprehensive
Wuhan University
MOE
Comprehensive
Xiamen University
MOE
Comprehensive
Xian Jiaotong University
MOE
Comprehensive
Xiangtan University
MOE
Comprehensive
*Xinjiang Bayi Agricultural College
Autonomous
Region
Agricultural
Xinjiang University
MOE
Comprehensive
*Xinjiang Shihetze Agricultural College
MOA
Agricultural
Yenan University
MOE
Comprehensive
Yunnan Forestry College
MOF
Forestry
Yunnan University
MOE
Comprehensive
*Zhejiang Agricultural University
PG
Agricultural
Zhejiang University
CAS
Comprehensive
Zhongshan University
MOE
Comprehensive
“ * = non-key institutions; without * = key institution.
^ MOE = Ministry of Education; MOA = Ministry of Agriculture; MOF = Ministry of
Forestry; CAS = Chinese Academy of Sciences; PG - Provincial Government.
the PRC has an agricultural economy. Virtually all cultivable land is used
for crops, and the intensive cultural techniques have already secured high
yields. To increase the yields any further requires better technology. Thus,
agricultural colleges and universities play an important role in this effort.
There are approximately 90 agricultural schools throughout China, and they
are mostly operated by the Ministry of Agriculture, the Ministry of Agri-
cultural Machinery, or Provincial Bureaus of Agriculture. Nearly all prov-
inces, autonomous regions and special municipalities have one agricultural
college with the exception of Xizang autonomous region (see map). A list
of well known agricultural colleges which deal with teaching and research
in plant protection is given in Table 1.
3) Other specialized institutions of higher education: Other institutions of
higher education can be categorized based on the nature of their curriculum.
They include: (a) polytechnic colleges and universities, (b) normal colleges,
(c) medical colleges, (d) science universities, (e) machine-building colleges,
(f) shipbuilding colleges, (g) aeronautics colleges, (h) electronic and telecom-
munication colleges, (i) light industry colleges, G) construction colleges, (k)
transportation colleges, (1) mining and metallurgy colleges, (m) chemical and
petroleum engineering colleges, (n) electric power college, (o) geology col-
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NEW YORK ENTOMOLOGICAL SOCIETY
leges, (p) meteorology colleges, (q) oceanography colleges, (r) other non-
technical colleges.
B. Institutes of Research
Currently PRC is attempting to upgrade the levels of professional research-
ers as well as training future generations of professionals. Entomological
research in PRC takes place under the auspices of the Chinese Academy of
Sciences, and the Ministry of Agriculture. The former is in charge of for-
mulating science policy, conducting basic research, and running several uni-
versities. Therefore, it is considered as a prestigious organization. There are
over 100 research institutes, five universities, and four libraries under the
jurisdiction of CAS. The primary emphasis of research institutes is in basic
research. However, CAS is also involved in teaching at its five universities
and training researchers at the graduate levels. The research institutes of
CAS dealing with agricultural production or plant protection are listed as
follows: In Beijing: Institute of Chemistry, Institute of Microbiology, Insti-
tute of Genetics, Institute of Zoology, Institute of Botany; In Shanghai:
Institute of Biochemistry, Institute of Cell Biology, Institute of Plant Phys-
iology, Institute of Entomology; In Sichuan Province: Institute of Biology;
VOLUME XC, NUMBER 3
207
In Yunnan Province: Institute of Zoology, Institute of Botany, Institute of
Tropical Plants; In Guangdong Province: Institute of Botany; In Liaoning
Province: Institute of Forestry and Pedology; In Hubei Province: Institute
of Virology, Institute of Botany; In Xinjiang Autonomous Region: Institute
of Chemistry, Institute of Biology, Pedology and Deserts; In Hebei Province:
Luancheng Institute of Agriculture Modernization; In Heilungjiang Prov-
ince: Institute of Agriculture Modernization; In Hunan Province: Taoyuan
Institute of Agriculture Modernization. The universities administered by
CAS are: In Anhui Province: University of Science and Technology of China;
In Beijing: School of Graduate Study, University of Science and Technology
of China; In Zhejiang Province: Zhejiang University; In Sichuan Province:
Chengdu University of Science and Technology; In Heilungjiang Province:
Harbin University of Science and Technology.
The Ministry of Agriculture manages its own institutes numbering several
hundred. The primary emphasis is conducting applied research. In general
the research institutes are under the jurisdiction of the provincial Academy
of Agriculture. The academies coordinate the activities of their subordinate
research institutes. Nearly every province or autonomous region has a pro-
vincial academy of agriculture.
C. Undergraduate and Graduate Education
Currently, the undergraduate curriculum is a four-year curriculum with
the requirement of 1 40 credit hours. However, some universities and colleges
have reinstated a five-year curriculum. The students are selected almost
exclusively on the basis of a nationally standardized entrance examination.
The entrance examination is held simultaneously for a 3-day period at the
various testing centers throughout the nation. As of July 1982, all participants
are required to take three basic subjects including Chinese, politics, and
mathematics. In addition, students majoring in science and technology are
required to take tests in biology, physics and chemistry, and those majoring
in social science and humanities are tested in history and geography. The
third test is the foreign language test, all examinees are given one of the seven
choices: English, French, German, Japanese, Russian, Spanish or Arabic.
The total points that the examinee majoring in science scores out of 600
possible points determines the outcome. However, the score on the foreign
language test is not counted in calculating the overall point total, it is only used
for considering student’s candidacy by the prospective college (Barendsen
1979; Hsu 1979). The passing score varies in different areas. The acceptance
total score for the resident student in Beijing area for 1980 was 370 points
at 500 possible points, where for students in other provinces the cutoff
total score was 348 points. Because of limited capabilities and short
resources, it is estimated that among the 12 million high school gradu-
ates, only about 300,000 students are able to enter the colleges. Cur-
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NEW YORK ENTOMOLOGICAL SOCIETY
rent total undergraduate enrollments in China stand at about 1.2 to 1.5
million. The goal set for 1985 is at 3 million (Abelson 1979). Graduate
enrollments will be increased to 200,000 by 1990. The faculty and student
ratio is at 1:3. This enables close contact between students and professors.
Thus the relationship, as traditional in China, becomes lifelong.
During the four years, all undergraduates majoring in plant protection are
required to take courses in: foreign language, general physics, organic chem-
istry, inorganic chemistry, advanced mathematics, general botany, plant
physiology, agricultural meteorology, philosophy, plant protection, political
economics, plant pathology, agricultural plant pathology, general entomol-
ogy, agricultural entomology, plant immunology, monitoring and forecast-
ing, biostatistics, mycology, plant bacteriology, plant virology, research tech-
niques, plant biochemistry, insect taxonomy, insect ecology, integrated pest
control, insect morphology, plant quarantine, insect physiology and insect
toxicology. The following courses are offered by some colleges: crop culti-
vation, cultivation techniques, plant breeding, soil and fertilizer, fruit tree
cultivation, vegetable production, agricultural machinery, agricultural eco-
nomics. The equipment and facilities in teaching laboratories and the librar-
ies are generally adequate in most institutions. More expansion and improve-
ment are underway. Some newly restored teaching laboratories have been
equal to or superior to comparable institutions in the U.S.
Graduate programs were restored in the fall of 1 978. Similarly, recruitment
into graduate programs is on the basis of both preliminary and subject matter
examinations. The former consists of four basic tests in politics, foreign
language, a basic subject, and one of the specialized subjects. The preliminary
examination is a nationally standardized test. The latter consists of foreign
language, basic subject, specialized subject. The last tests are conducted by
each graduate school. As a result, only one in six college graduates is able
to pass the test. From this test program it is apparent that the leaders in
PRC realize the importance of foreign languages as a tool in learning advanced
science and technology. In 1979 approximately 19,000 graduate students
selected from the 200,000 college graduates were enrolled in over 200 colleges
and universities as well as 150 research institutes of the CAS and CAAS.
The graduate curriculum ranges from 2 to 4 years depending on various
institutions. Because this program is still at its infant stage, it is somewhat
flexible, with more emphasis placed on research, rather than the research
and course work normally required in other countries.
Since the restoration of graduate study programs in 1978, the leaders in
PRC also have realized the need to award the academic degrees which would
help stimulate national interest in scientific research, raise academic stan-
dards, and facilitate international academic exchanges. Subsequently, the
regulations governing the award of three academic degrees (Bachelor, Master
and Doctor) were approved by the National People’s Congress in February
1979. The measures further provide the State Council of PRC with a vehicle
VOLUME XC, NUMBER 3
209
to establish an academic degrees committee. The bachelor degree is awarded
by 458 colleges and universities authorized by the State Council, whereas
master and doctor degrees are awarded by the academic degrees committee.
Candidates enrolled in the institutions of higher learning and research insti-
tutes as well as those who can pass the qualifying examination and satisfy ex-
aminers in an oral examination or a written thesis are eligible for postgraduate
degrees. The same regulations also apply to foreign students studying in
PRC. Provisions are also made to award honorary degrees to outstanding
scholars residing at home and abroad.
In 1980, the State Council approved the designation of the additional
universities as key national institutions of higher education: (Northwest
University, Southwest University, Central China University, South China
University and Shenyang University), serving the needs of all regions. It is
worth noting that these newest key universities have formally separated the
Department of Plant Protection into a Department of Entomology and a
Department of Plant Pathology. They are charged with the responsibility of
training college faculty and researchers for the respective regions, whereas
other colleges of agriculture still retain the Department of Plant Protection
with the responsibility of training plant protection specialists in the fields.
The impact of the Cultural Revolution on China’s education is profound.
China not only has lost a generation of college graduates, but also a generation
of students at other levels as well. Therefore, the tasks of modernization of
a large educational system are enormous and complex. Currently, PRC pro-
duces about 200,000 college graduates per year. The newly initiated edu-
cational exchange programs with the Western countries, and the government
funded overseas study programs are of current priority to the Chinese gov-
ernment. From January 1978 to November 1979, a total of 2,230 students
and scholars was sent overseas including 1 ,600 researchers and scholars, 1 80
graduate students, and 450 undergraduates. Among these, 1,800 students
were majoring in natural sciences, the rest in social study and linguistics.
There were 500 individuals in the U.S., 500 in U.K., 200 in France, 200 in
F.R. Germany and 100 in Japan. China’s aim in sending students and schol-
ars overseas for 1980 was 10-20,000. It was estimated that about 6,000
Chinese students and scholars were studying at the U.S. universities at the
end of 1981.
Conclusions
China has an ancient history of science and civilization. Unfortunately,
China has been repeatedly subjected to socio-economical, political, and intel-
lectual chaos and revolutions in her search for accommodation with the
modem world since the turn of the 1 9th Century. The process of transforming
a traditionally agricultural society into a modern, industrial state has not
been an easy one. After 1948, a decade of remarkable progress had been
210
NEW YORK ENTOMOLOGICAL SOCIETY
made on the unification of the country, self-sufficiency, fostering education
and research. In the last two decades there have been advances, but progress
has been variable. This was due in large part to repeated changes in ideology.
One of the ideological changes with profound effect on education and research
was the Cultural Revolution. As a result, China has lost a generation of
educators and researchers. To implement the four modernizations and to
make up for ground lost during this period are enormous and complex tasks
which would require a vast infusion of foreign technical assistance and the
creation of domestically exceptional capabilities. Hence, there are ample
opportunities for foreign scientists to cooperate with Chinese counterparts.
During the Cultural Revolution, basic research in plant protection was
greatly disrupted, but some applied research flourished. In some aspects of
plant protection, the Chinese are indeed ahead of Western nations and are
probably in a better position than any other nation to provide alternatives
to chemical control of a number of agricultural pests. Many common pests
in the PRC have been effectively managed and controlled by the use of
integrated pest management measures involving cultural control, biological
control, as well as some chemical methods. The success of IPM programs
in China is attributed to the widespread enthusiasm for IPM shared by the
farmers, scientists and administrators, and, to the application of the eco-
logical principles essential to its development. The insect control measures
developed in China are simple, effective and economical. Therefore, they
could be used in other parts of the world where crop production and pro-
tection are carried out mostly by small farmers with little modern technology.
To increase grain production in meeting the need of population growth
in China and to boost its already high per unit area yields pose a great
challenge to Chinese scientists. Basic research and modern technology will
undoubtedly play an important role in developing modern plant protection
techniques in the years ahead. However, Chinese scientists should be aware
of the impact of modem technology on the future plant production and
protection, such example as the impact of the green revolution on plant
protection in tropical and subtropical areas (Smith 1972). Also, they should
be aware of the fact that once truly committed to modernization, they will
face no end of changes, pressures and socio-economic problems. China is
known to be well endowed with many important minerals and energy
resources. Oil reserves are estimated at least three times as large as those of
the U.S. It has tremendous resources of talented, industrious and moral
people. It is to be hoped that their national goals will soon be realized.
Acknowledgments
I am grateful to Professor Kuan Chih-hu, Beijing Agricultural University
and Dr. Chou lo. Northwestern College of Agriculture for providing much
VOLUME XC, NUMBER 3
211
of the information. Appreciation is extended to Drs. S. C. Ma and H. F.
Chu, Institute of Zoology, Academia Sinica, Beijing and Prof. J. S. Lu,
Northwestern College of Agriculture for their suggestions.
Appreciation is extended to the National Science Foundation, Washington,
D.C. and the Entomological Society of America for travel grants for this
invitational paper at the 16th International Congress of Entomology in
Kyoto, Japan. 1980.
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Abelson, P. H. 1979. Education, science and technology in China. Science 203:505-509.
Anonymous. 1956. Insect Names. Science Press, Beijing. 40 pp.
Anonymous. 1972. Pictures of Chinese Crop Diseases and Insect Pests. Publ. No. 13. Vol.
2. Agricultural Publishing House, Beijing. 86 pp.
Anonymous. 1974. Handbook for Plant Protection Workers. Vol. 1-7. Shanghai People’s
Press, Shanghai. 365 pp.
Anonymous. 1977a. Agricultural Entomology. Vol 1. People’s Education Press, Beijing.
325 pp.
Anonymous. 1977b. Agricultural Entomology. Vol. 2. People’s Education Press, Beijing.
274 pp.
Anonymous. 1978a. Colored Illustrations of Rice Pests and Their Natural Enemies. Hubei
People’s Press. 181 pp., 88 plates.
Anonymous. 1978b. Colored Illustrations of the Natural Enemies of Insects. Science Press,
Beijing. 300 pp., 50 plates.
Anonymous. 1979a. An Illustrated Book of Natural Enemies of Rice Pests in Zhejiang Prov-
ince. Zhejiang People’s Press. 210 pp.
Anonymous. 1979b. Integrated Control of Major Pests in China. Science Press, Beijing.
467 pp.
Anonymous. 1980. General Entomology. Vol. 1. Agricultural Publishing House, Beijing.
425 pp.
Anonymous. 1982. Foreign Agriculture Circular. USDA. Foreign Agricultural Service. WCP-
2-82. 15 pp.
Barendsen, R. D. 1979. The 1978 national college entrance examination in the People’s
Republic of China. Office of Education, US Dept. HEW. Washington, D.C.
Cheng, T. H. 1963. Insect control in mainland China. Science 140:269-277.
Chiang, H. C. 1977a. Pest control in the People’s Republic of China. Science 192:675-677.
. 1977b. Pest management in the People’s Republic of China— monitoring and fore-
casting insect population in rice, wheat, cotton and maize. FAO Plant Protection Bull.
25:1-8.
Chiu, S. F. 1959. The accomplishments of controlling agricultural pests in the last decade of
China. Kunchong Zhishi (=Knowledge of Insects) 5:241-243.
Chou, I. 1980. A History of Chinese Entomology. Entomotaxonomia. Wugong, Shaanxi.
213 pp.
Guyer, G. E., Chairman. 1977. Insect Control in the People’s Republic of China: A Trip
Report of the American Insect Control Delegation. CSCPRC Rept. No. 2. Washington,
D.C. NAS. 218 pp.
Huang, M. D., S. H. Mai, W. N. Wu and C. L. Poo. 1974. The bionomics of Anastatus sp.
and its utilization for the control of lichee stinkbug, Tessaratoma papillosa Drury. Acta
Entomol. Sinica 17:362-375.
Hsiu, C. F., Y. Chang, C. M. Kwei, Y. M. Han and H. H. Wang. 1973. Field application with
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Beauveria bassiana (Bals.) Vuill. for European corn borer control. Acta Entomol. Sinica
16:203-206.
Hsu, C. T. 1979. The 1978 examinations. Contemporary China 3:63-68.
Kelman, A. and R. J. Cook. 1977. Plant pathology in the People’s Republic of China. Ann.
Rev. Phytopathol. 17:409-429.
Kung, P. 1975. Farm crops of China. World Crops 123-132.
Lin, P. H. 1956. A preliminary report on the utilization of the muscardin fungus, Beauveria
bassiana (Bals.), to control the sweet-potato weevil, Cylas formicarius Fab. Acta Entomol.
Sinica 6:539-540.
Ma, S. C. 1958. The population dynamics of the oriental migratory locust {Locusta migratoria
manilensis Meyen) in China. Acta Entomol. Sinica 8:1-40.
. 1962. The structure and transformation of the Oriental locust breeding area. Acta
Entomol. Sinica 11:1 7-30.
Poo, C. L., T. H. Tang, C. C. Liu, F. C. Hung and Y. S. Mo. 1956. On the rearing of
Trichogramma evanescens Westw. and its utilization for the control of sugar cane borer.
Acta Entomol. Sinica 6:1-36.
Reardon-Anderson, J. 1978. Science and technology in post-Mao China. Contemporary China
2:37-45.
Smith, R. F. 1972. The impact of the green revolution on plant protection in tropical and
subtropical areas. Bull. Entomol. Soc. Am. 18:7-14.
Su, Y. N. 1959. Accomplishments on the monitoring and forecasting of agricultural pests in
China. Kunchong Zhishi 5:244-245.
Williams, P. H. 1979. Vegetable crop protection in The People’s Republic of China. Ann.
Rev. Phytopathol. 17:311-324.
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three years. Kunchong Zhishi 4:197-198.
Agricultural Research and Education Center, University of Florida, 3205
S. W. 70 Ave., Fort Lauderdale, Florida 33314.
Received for publication May 12, 1982.
NEW YORK ENTOMOLOGICAL SOCIETY
XC(3), 1982, pp. 213-219
EFFECTS OF TWO INSECT GROWTH REGULATORS
(HYDROPRENE AND R-20458) ON THE FOLLICULAR
EPITHELIUM AND THE OOCYTES OF THE RICE WEEVIL,
SITOPHILUS ORYZAE (L.) (COLEOPTERA: CURCULIONIDAE)
J. M. Mkhize* and A. P. Gupta
Abstract.— Effects of two insect growth regulators (IGRs) (hydroprene and
R-20458) on the follicular epithelium and the oocytes of the rice weevil,
Sitophilus oryzae (Coleoptera: Curculionidae) were studied. Comparative
histological studies of the treated and untreated ovarioles revealed that there
were no apparent differences in the development and structure of the oocytes
in the germarium and in the anterior part of the vitellarium. The IGRs,
however, affected the penultimate oocytes and perhaps the basal oocytes,
because the latter ovulated prematurely. Treated penultimate oocytes were
atrophied, because they lacked yolk and karyospheres; in addition, follicular
epithelium was not retracted from the surface of the oolemma. Untreated
penultimate oocytes on the other hand, were larger due to the accumulation
of yolk and they had karyospheres; the follicular epithelium was retracted
from the egg membrane, leaving a space that had materials, which possibly
are blood proteins. A fine brush-like border (microvilli) was observed on
the side of the oolemma facing the follicular epithelium.
Introduction
Laboratory evaluation of IGRs as protectants against pests of stored prod-
ucts have shown considerable promise (Metwally et al. 1972; Bhatnager-
Thomas 1973; Strong and Diekman 1973; Williams and Amos 1974;
McGregor and Kramer 1975; Loschiavo 1976; and Amos and Williams
1977). It has also been demonstrated that treatment of certain species of
insects with IGRs induces permanent ovarial abnormalities that lead to
female sterility (Metwally and Landa 1972; Metwally et al. 1972; Rohdendorf
and Sehnal 1973; Lanzrein 1974; Patterson 1974; Das and Gupta 1977;
Masner et al. 1979; and Deb and Chakra vorty 1981).
The purpose of this work was to hnd out whether hydroprene and
R-20458 will induce ovarial abnormalities that might cause sterility in the
female weevil.
‘ Present address: P.O. Box B75, Maseru 100, Lesotho, South Africa.
The publication costs of this article were defrayed in part by page charge payment. This article
must therefore be hereby marked “Advertisement” in accordance with 18 U.S.C. §1734 solely
to indicate this fact.
214
NEW YORK ENTOMOLOGICAL SOCIETY
Materials and Methods
Several IGRs were screened for their effects, and hydroprene (ethyl(2E,4E)-
3,7,1 1 -trimethyl-2, 4-dodecadienoate) and R-20458 (6,7-epoxy-3,7-dimethyl-
1 -(p-ethylphenoxy)-2-octene) were selected because they were more effective
than others against S. oryzae. Of these two, hydroprene was more effective
(Mkhize and Gupta 1980; Gupta and Mkhize 1982).
Ovaries of rice weevils, which emerged from either IGR-treated wheat
(see Gupta and Mkhize 1 982) or topical applications, were removed in saline
solution and fixed in two successive changes of Bouin’s fluid. The tissues
were then dehydrated in ethanol series, cleared in xylene and embedded in
paraffin wax. Longitudinal sections, each 5 -Mm thick, were stained in hema-
toxylin and eosin and mounted in permount. Histology of treated and
untreated ovaries was compared in order to discover morphological abnor-
malities induced by IGR treatment.
For making ultrathin sections, treated and untreated ovaries were sepa-
rately dissected in physiological saline solution and fixed in 2.5% glutaral-
dehyde in 0.1 M cacodylate buffer (pH 7.2) for four hr. After rinsing the
ovaries in a buffer, they were placed in 1 % solution of osmium tetroxide for
two hr. The ovaries were again rinsed in a buffer and then dehydrated in
acetone as follows: they were placed in 30%, 70%, and 95% acetone for 20
min in each concentration; then in three changes of 100% acetone for 20
min/change. They were embedded in Epon 8 1 2. One /um-thick cross sections
were made by Sorvall MT-2 ultramicrotome. The sections were stained for
one min, and mounted in immersion oil. The sections were then sealed with
permount. Light micrographs of cross sections of both treated and untreated
oocytes were made and compared in order to observe abnormalities owing
to IGR treatment.
Results
Examination of treated and untreated ovarioles revealed that the ger-
marium and the anterior region of the vitellarium showed no apparent
structural or developmental differences between the oocytes in the treated
and untreated ovarioles. Differences, however, were observed in the pen-
ultimate oocytes and the follicular epithelium around them. In the anterior
region of the vitellarium, the cells of the follicular epithelium were columnal
in shape and had elongated nuclei (Fig. 1). The cell boundaries were distinct
and the cytoplasm appeared strongly basophilic, probably indicative of syn-
thetic processes taking place in these cells.
The follicular epithelium around untreated penultimate oocytes had cells
that appeared slightly flattened, with small, elongated nuclei, indicating that
these cells were in a transitional stage from cuboidal to squamous type of
cells (Fig. 2). The epithelium also showed mitotic divisions (Fig. 3) and the
nuclear chromatin material seemed to be evenly dispersed. Figure 4 shows
VOLUME XC, NUMBER 3
215
Figs. 1-6. 1. X-section of treated, young oocyte, showing columnar epithelium. NU ^nucleus
of oocyte; NUC = Nucleolus. X3,500. 2, 3. X-sections of untreated penultimate oocytes, show-
ing squamous follicular epithelium (arrow head) (Fig. 2, X 3,040) and mitotic figure (Fig. 3,
X2,945). 4. X-section of an untreated penultimate oocyte engaged in vitellogenesis. Note space
between oolemma and follicular epithelium and inter-cellular space in the follicular epithelium
(arrow head). X2,945. 5. X-section of a treated penultimate oocyte. Note absence of intercellular
space in the follicular epithelium and between the latter and the oolemma. X 3,040. 6. X-section
of an untreated oocyte engaged in vitellogenesis, showing a germinal vesicle (arrow head).
X3,420.
216
NEW YORK ENTOMOLOGICAL SOCIETY
the follicular epithelium of an untreated penultimatic oocyte actively engaged
in vitellogenesis. This epithelium was retracted from the surface of the egg
membrane (oolemma) creating a space between the two interfaces. In addi-
tion, there were intercellular spaces in the epithelium itself. These spaces
contained material that was less granular than that in the oocyte. Focusing
up and down over the space between the follicular epithelium and the oo-
lemma sometimes revealed brush-like border on the oolemma. Retraction
of the follicular epithelium from the oolemma enables hemolymph proteins
to pass into the oocyte, supposedly by pinocytosis. By contrast, the follicular
epithelium in a treated penultimate oocyte was not retracted from the oo-
lemma (Fig. 5) and no intercellular spaces were present. The follicular cells
were roundish with spherical nuclei. No mitosis was observed.
Untreated young oocytes in the anterior region of the vitellarium in both
treated and untreated oocytes had centrally located nuclei (Fig. 1), with
evenly distributed chromatin material. Mature untreated oocytes engaged
in vitellogenesis showed germinal vesicle (=the enlarged oocyte nucleus due
to accumulation of vesicular fluid) (Fig. 6) and karyosphere (the clumped
chromatin material of the nucleus), both of which were not observed in
treated oocytes.
Discussion
The follicular epithelium plays an important role in vitellogenesis. The
synthesis of enzymes necessary for the uptake of nutrients from the hemo-
lymph (De Loof and Lagasse 1970), proteins (King and Burnett 1959; Zalokar
1960; Beir 1962, 1963b) and of adsorbents that link blood proteins to the
outer surface of the oocyte (Roth and Porter 1964; Anderson and Telfer
1970; Anderson 1971) is reported to occur in the follicular epithelium. The
facts that the follicular epithelium in the IGR-treated ovarioles were not
retracted from the oolemma, showed no mitosis, and the cells were roundish,
suggest abnormalities that prevent successful completion of vitellogenesis.
Retraction of the follicular epithelium from the oolemma enables hemo-
lymph proteins to pass into the oocyte by pinocytosis (Beir 1962, 1963a;
Kessel and Beams 1963; Telfer and Melius 1963; Roth and. Porter 1964;
Stay 1965; Hopkins and King 1966; De Loof and Lagasse 1970). Clearly,
the IGRs in this study blocked retraction of the oolemma and thus vitel-
logenesis.
In most cases, during vitellogenesis the development of the oocyte nucleus
is arrested at meiotic prophase and that just before vitellogensis takes place,
both the nucleus and the cytoplasm become very basophilic and rich in RNA
(Bonhag 1958; Mahowald 1972; De Robertes et al. 1975; Ambrose and Hasty
1978). According to Schlottman and Bonhag (1956) and Gupta and Riley
(1967) at about this time, the nucleus attains maximal size due to accu-
mulation of vesicular fluid and is known as germinal vesicle, whose chro-
matin material is clumped and is called karyosphere (Chandley 1966). The
VOLUME XC, NUMBER 3
217
absence of the germinal vesicle and the karyosphere in the treated oocyte
confirms the disruption of vitellogenesis. As a result of the abnormalities
discussed in the foregoing, the ovarioles in the IGR-treated weevils remain
atrophied, and consequently no Fj progeny is produced.
Acknowledgments
We are grateful to Zoecon Corporation and Stauffer Chemical Co. for gifts
of the IGRs. This report is the New Jersey Agricultural Experiment Station
Publication No. D-081 12-24-82, supported by State Funds and by U.S. Hatch
Act Funds. The secretarial assistance of Evelyn Weinmann is appreciated.
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(JMM) Department of Entomology & Economic Zoology, Cook College,
New Jersey and (APG) Agricultural Experiment Station, Rutgers University,
New Brunswick, New Jersey 08903.
Received for publication May 12, 1982.
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Vol. XC
SISiX'P'V
DECEMBER 1982 ^ (
No. 4
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The Journal of the New York Entomological Society (ISSN 0028-7199) is published quarterly for the Society by Allen Press
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Journal
of the
New York
ENTOMOLOGICAL SOCIETY
Devoted to Entomology in General
VOLUME LXXXVIII
Published by the Society
New York, N.Y.
INDEX OF AUTHORS FOR VOLUME XC
AHMAD, S. and C. R. FUNK. Susceptibility of Kentucky bluegrass cultivars and
selections to infestations of and injury by the bluegrass billbug (Coleoptera; Curculi-
onidae) - 3 1
ANDOW, D. Miridae and Coleoptera associated with tulip tree flowers at Ithaca, New
York 119
BAKER, R. A. Unionicolid mites from central New York.... 176
COTTY, S. and J. H. LASHOMB. Vegetative growth and yield response of eggplant to
varying first generation Colorado potato beetle densities 222
EVANS, E. W. Habitat differences in feeding habits and body size of the predatory
stinkbug Perillus circumcinctus (Hemiptera: Pentatomidae) 129
EVANS, E. W. Influence of weather on predator/prey relations: stinkbugs and tent
caterpillars 243
FERGUSON, G. R. Descriptions, synonymy and sex association in the genus Eucerceris
(Hymenoptera: Philanthidae) 147
FOWLER, H. G. Habitat effect on fungal substrate selection by a leafcutting ant 64
FOWLER, H. G. A new species of Trachymyrmex fungus-growing ant (Hymenoptera:
Myrmicinae: Attini) from Paraguay 70
GAMBINO, P. Phenology of emergence of the spotted tentiform leafminer, Phyllon-
orycter crataegella (Lepidoptera: Gracillariidae) and its parasitoids in New York 231
GILBERT, L. E. Oviposition by two Heliconius species: comments on a paper by Dr.
A. Young 1 1 5
JULIAR, M. Vladimir Nabokov 1899-1977: a note on a late entomologist — 55
KAMM, J. A. and L. M. McDONOUGH. Seasonal flight of the cranberry girdler de-
termined with pheromone traps 94
KARBAN, R. Experimental removal of 17-year cicada nymphs and growth of host
apple trees 74
KLEIN, B. G. Pit construction by antiion larvae: influences of soil illumination and
soil temperature 26
KNOP, N. F., M. A. HOY and M. E. MONTGOMERY. Altered hatch sequence of
males and females from unchilled eggs of a “non-diapause” gypsy moth strain (Lepi-
doptera: Lymantriidae) 82
1
McCABE, T. L. and G. L. GODFREY. Larval morphology and phylogeny of Trichor-
destra tacoma (Strecker) (Noctuidae) 142
McDonald, F. J. D. Description of the male genitalia of Holcostethus hirtus (Van
Duzee) with a revised key to North American species (Hemiptera: Pentatomidae) 5
McKEON, J. P., T. F. BAST and E. M. BOSLER. The Lone Star tick, Amblyomma
americanum (Linnaeus): new distribution record for North America (Acarina: Ixo-
didae) 267
MKHIZE, J. M. and A. P. GUPTA. Effects of two insect growth regulators (Hydroprene
and R-20458) on the follicular epithelium and the oocytes of the rice weevil, Sitophilus
oryzae (L.) (Coleoptera: Curculionidae) 213
NECHOLS, J. R. and P. J. TAUBER. Thermal requirements for post-diapause devel-
opment and survival in the giant silkworm, Hyalophora cercropia (Lepidoptera: Satumi-
idae) 254
NORDEN, B. B. and A. G. SCARBROUGH. Predators, parasites, and associates of
Anthophora abrupta Say (Hymenoptera: Anthophoridae) 181
PLAKIDAS, J. D. Notes on gall inhabitants of Asphondylia helianthiglobulus Osten
Sacken (Diptera: Cecidomyiidae) in Western Pennsylvania 2
POST, D. C. and R. L. JEANNE. Sternal glands in three species of male social wasps
of the genus Mischocyttarus (Hymenoptera: Vespidae) 8
ROITBERG, B. D. and R. J. PROKOPY. Resource assessment by adult and larval
codling moths 260
SAKALUK, S. K. Onset of phenotaxis and age at first mating in female house crickets,
Acheta domesticus (Orthoptera: Gryllidae) 136
SALYK, R. P. and D. J. SULLIVAN S.J. Comparative feeding behavior of two aphid
species: bean aphid {Aphis fabae Scopoli) and pea aphid {Acyrthosiphon pisum (Harris))
(Homoptera: Aphididae) 87
SIDHU, D. S., S. P. KAUR and N. KUMAR. Tissue glycogen of Mylabris pustulata
Thunb. and Periplaneta americana L. (Insecta) 239
SIMON, C. and M. LLOYD. Disjunct synchronic populations of 17-year periodical
cicadas: relicts or evidence of polyphyly? 276
TEPEDINO, V. J. and D. R. FROHLICH. Mortality factors, pollen utilization and sex
ratio in Megachiles pugnata Say (Hymenoptera: Megachilidae), a candidate for com-
mercial sunflower pollination 270
TSAI, J. H. Entomology in The People’s Republic of China 186
UTBERG, G. L. and D. J. SUTHERLAND. The temporal distribution of Chironomus
decorus (Chironomidae) in northern New Jersey, 1979 16
WASTI, S. S. and G. C. HARTMANN. Susceptibility of gypsy moth larvae to several
species of entomogenous fungi 125
WILSON, S. W. and J. H. TSAI. Descriptions of the immature stages of Myndus crudus
(Homoptera: Fulgoroidea: Cixiidae) 166
YOUNG, A. M. Notes on the natural history of Morpho granadensis polybaptus Butler
(Lepidoptera: Nymphalidae: Morphinae), and its relation to that of Morpho peleides
limpida Butler... 35
YOUNG, A. M. Notes on the interaction of the skipper butterfly Calpodes ethlius
(Lepidoptera: Hesperiidae) with its larval host plant Canna edulis (Cannaceae) in
Mazatlan, State of Sinaloa, Mexico 99
YOUNG, A. M. Errata: over-exploitation of larval host plants by Heliconius butter-
flies 1 1 7
YOUNG, O. P. Perching behavior of Canthon viridis (Coleoptera: Scarabaeidae) in
Maryland 161
ii
Book Reviews
HARPAZ, I. Vectors of Plant Pathogens. Edited by K. F. Harris and K. Maramorosch 58
KURTTI, T. J. The Ecology of Pests— Some Australian Case Histories. Edited by R.
L. Kitching and R. E. Jones 304
MARAMOROSCH, K. Handbook of Plant Virus Infections: Comparative Diagnosis.
Edited by E. Kurstak.— 59
MARAMOROSCH, K. Insects and Other Invertebrates of the World on Stamps. Edited
by W. E. Stanley 6 1
MARAMOROSCH, K. The North American Grasshopper. Vol. I. Acrididae: Gom-
phocerinae and Acridinae. D. Otte 133
MARAMOROSCH, K. Insects: An Anthology of Arthropods Featuring a Bounty of
Beetles. Paintings by P. A. Gette, entomological commentaries by G. Scherer. Trans-
lated from German by G. Zappler 134
MARAMOROSCH, K. The American Cockroach. Edited by W. J. Bell and K. G.
Adiyodi. The Laboratory Cockroach. W. J. Bell 303
MARAMOROSCH, K. Plant Protection: An Integrated Interdisciplinary Approach. W.
H. Sill, Jr 304
THOMAS, D. B., JR. The Pentatomoidea (Hemiptera) of Northeastern North America
with Emphasis on the Fauna of Illinois. J. E. McPherson 302
iii
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Journal of the
New York Entomological Society
Vol. XC
DECEMBER 1982
No. 4
EDITORIAL BOARD
Editor Associate Editors
Dr. Karl Maramorosch Dr. Lois J. Keller, RSM
Waksman Institute of Microbiology Dr. Herbert T. Streu
Rutgers University
New Brunswick, New Jersey 08903
Publication Committee
Dr. Randall T. Schuh
American Museum of
Natural History
Dr. Louis Trombetta
St. Johns University
CONTENTS
Vegetative growth and yield response of eggplant to varying first generation Colorado
potato beetle densities Suzanne Cotty and James H. Lashomb 220-228
Phenology of emergence of the spotted tentiform leafminer, Phyllonorycter crataegella
(Lepidoptera: Gracillariidae) and its parasitoids in New York
Parker Gambino and Daniel J. Sullivan, S.J. 229-236
Tissue glycogen of Mylabris pustulata Thunb. and Periplaneta americana L.
(Insecta) Dalbinder Singh Sidhu, Surinder Pal Kaur and Nirmal Kumar 237-240
Influence of weather on predator/prey relations: stinkbugs and tent caterpillars
Edward W. Evans 241-246
Seasonal occurrence of founding queens and the sex ratio of Camponotus pennsyl-
vanicus (Hymenoptera: Formicidae) in New Jersey
Harold G. Fowler and Radclyffe B. Roberts 247-251
Thermal requirements for postdiapause development and survival in the giant silk-
worm, Hyalophora cecropia (Lepidoptera: Satumiidae)
James R. Nechols and Paul J. Tauber 252-257
Resource assessment by adult and larval codling moths
Bernard D. Roitberg and Ronald J. Prokopy 258-265
The Lone Star tick, Amblyomma americanum (Linnaeus): New distribution record
for North America (Acarina: Ixodidae)
John P. McKeon, Thomas F. Bast and Edward M. Bosler 266-268
Mortality factors, pollen utilization and sex ratio in Megachile pugnata Say (Hyme-
noptera: Megachilidae), a candidate for commercial sunflower pollination
V. J. Tepedino and D. R. Frohlich 269-274
Disjunct synchronic populations of 17-year periodical cicadas: Relicts or evidence of
polyphyly? Chris Simon and Monte Lloyd 275-301
Book Reviews 302-305
Acknowledgement 306
Honorary Life and Sustaining Members 306
Index of Scientific Names of Animals and Plants for Volume XC 307-310
Index of Authors for Volume XC i-ii
NEW YORK ENTOMOLOGICAL SOCIETY
XC(4), 1982, pp. 220-228
VEGETATIVE GROWTH AND YIELD RESPONSE OF EGGPLANT
TO VARYING FIRST GENERATION COLORADO
POTATO BEETLE DENSITIES
Suzanne Cotty and James H. Lashomb
Abstract. — Ndirying densities (0, 8, 15, 23 or 30) of newly eclosed first
generation Colorado potato beetle Leptinotarsa decemlineata larvae were
maintained on eggplant to describe their effects on vegetative growth, flow-
ering and yield. Significant depression of height and leaf area was detected
5-6 weeks post-infestation only for 30 larvae/plant. However, all densities
temporarily disrupted flower production. Densities above 8 larvae/plant
temporarily reduced yield. Beetles were excluded in all treatments during
the second generation during which the plants compensated for early yield
losses. Total yield for the season was the same in all treatments.
In New Jersey normal production of the solanaceous crops, eggplant {Sola-
rium melangena L.), potato {S. tuberosum L.) and tomato {Lycopersicon
esculenturn Mill.) is currently complicated by insecticide resistant popula-
tions of the Colorado potato beetle (CPB) {Leptinotarsa decemlineata Say).
In these crops there are no suitable alternatives to insecticides for main-
taining the beetle below an economic threshold. For potato, Logan and
Casagrande (1980) developed a physiological time model to predict yield
losses on cv ‘Superior’ and Hare (1980) concluded that cv ‘Katahdin’ yield
reductions were greatest during emergence and oviposition of summer adults.
For tomato, cv ‘Chico Grande’ yield was negatively correlated with increas-
ing plant damage (Shalk and Stoner 1979). There are no published results
regarding yield losses caused by any insect on eggplant. In Japan, Fugi and
Itagi (1954) showed that mechanical defoliation of eggplant disrupted fruit
set. Saito and Ito (1973) demonstrated that artificial defoliation of eggplant
retarded flower development and the resulting flowers had smaller sepals,
petals, anthers and ovaries with smaller styles. Also, the reduced flowers
were shed at a higher rate than normal ones. Neither Japanese study reported
which cultivar was used or related defoliation intensity to yield.
S. rnelongena has an indeterminant growth habit, for after initial foliar
development, flower production begins and continues until frost. There are
always more flowers produced than fruit (Bailey 1891; Smith 1931; Eguchi
The publication costs of this article were defrayed in part by page charge payment. This article
must therefore be hereby marked “Advertisement” in accordance with 18 U.S.C. §1734 solely
to indicate this fact.
VOLUME XC, NUMBER 4
221
et al. 1958; Prasad and Prakash 1968; Saito and Ito 1973; Free 1975; and
Mohideen et al. 1977). When there is more than one flower in an inflorescence
only the distal one is perfect and sets fruit. The remaining flowers are shed
(Bailey 1891).
The purpose of this study was to determine the effect of varying larval
densities of first generation CPB on vegetative growth, flowering and yield
of eggplant.
Methods
We germinated ‘Harris Special Highbush’ seeds in flats in the greenhouse
8 weeks prior to planting in 1980. The plants were field conditioned for 5
days before transplanting on May 20, 1980 in rows with a 2.7 m spacing.
A 10-10-10 NPK solution was applied at planting and again 1 month later
with 30% ammonium nitrate crystals. Chlorthal (13.5 kg/ha) was applied 1
week after planting for weed control. Thereafter the plants were irrigated
and cultivated regularly.
Twenty-three days post-planting we inoculated the plants with 0, 8, 15,
23, or 30 CPB eggs/plant. After eclosion we maintained larval density by
adding similar aged larvae. The larval densities were selected based on a
preliminary study in 1979 where we inoculated the plants with 0, 15, 30 or
60 larvae/plant. Plants with 60 larvae were soon defoliated, so in 1980 we
used 30 larvae as our maximum density/plant. Each density was replicated
on 12 plants in groups of 4 in 3 different sub-plots using a completely
randomized design. Larvae were maintained on the plants until pupation.
After 2 1 days the few remaining larvae were removed to maintain uniformity
of feeding effects. Adults were not allowed to feed on any plants during the
study. We used fenvalerate (276 g Al per hectare weekly) to maintain the 0
density during the larval feeding period. After larval feeding terminated all
plants were sprayed to prevent adult feeding and oviposition.
Beginning with inoculation of the plants and weekly thereafter the follow-
ing data were collected from each plant: height, leaf area, flower number and
fruit number. To determine leaf area we established 3 leaf size classes: small,
medium and large. We then sampled 20 leaves from each size class and
measured their area with a photoplanimeter (LICOR-3000, LAMBDA
Instruments, Lincoln, Nebraska). The mean values 62 cm^, 250 cm^, and
427 cm^ were used to make cardboard templates for each leaf size class.
Leaves on each plant were then classified as to size using the template and
summed. Fruit were harvested upon maturity which was subjectively deter-
mined to be 454 g and having a shiny purple skin. The data were analyzed
using a completely randomized Analysis of Variance to test the effects due
to differing densities on each variable each week. Testing for differences
between means of each variable was done using the least significant difference
test (Cochran and Cox 1957).
222
NEW YORK ENTOMOLOGICAL SOCIETY
Results
In 1979 the control plants produced only 60% of the normal yield of the
cultivar which may have affected the relative differences between treatments.
Therefore only the 1 980 results are presented here. Each variable is discussed
individually.
Plant height. — shape of the plant height curve was very similar among
the treatment levels, but at the higher larval densities, a slight delay in height
occurred (Fig. 1). Height differences among treatments were not evident
until week 5 (week number refers to time post-CPB inoculation). At this
time, only mean plant height for the 30 larvae/plant treatment was less than
the controls {P < 0.05). By week 6, the mean height for this treatment had
dropped to 23% less than the controls {P < 0.05). Effects due to feeding
could be expected to occur during weeks 5 and 6, since the larvae were in
the fourth and most destructive instar (Tamaki and Butt 1978) and they
tend to feed in the apical portion of the plant. After week 6, mean height of
30 larvae/plant was the same as the controls. Plant heights in the lower 3
larval densities were not different from the controls during the season.
Leaf area. — During weeks 1-4, there were no differences in mean leaf area
between treatments (Fig. 2). By week 5, the effects of feeding became evident
in the 8,15, and 30 larvae/plant treatments, when compared to the controls.
The leaf areas were 22%, 21%, and 25% less than the controls, respectively
(P < 0.05) which might be attributed not only to an actual leaf loss, but
also to a delay in the production of new leaves. This period is followed by
a transition in which no differences in leaf area occurred among treatments.
From week 10 to the end of the season, the mean leaf area of the 30 larvae/
plant treatment ranged from 18 to 34% higher than the leaf area of the
controls {P < 0.05). This difference appears to be a response to injury in
which the heavier defoliated plants compensate for depressed leaf area.
Throughout the season, mean leaf area of the 8, 15, and 23 larvae/plant
treatments were generally intermediate to the controls and the 30 larvae/
plant treatments. Differences in mean leaf area were not significant between
treatments at the end of the season.
Flower production. — Yio^mng began 27 days after planting (4 days post
CPB inoculation) and continued throughout the season (Fig. 3). The first 4
week flower production for all treatments was the same. By week 5, however
all the treated plants (8, 15, 23, and 30 larvae/plant) produced 49, 79, 57,
and 52% fewer flowers than the controls, respectively (P < 0.05). This
depression of flower production continued through week 7, with the treated
plants producing from 14 to 8 1% fewer flowers than the controls {P < 0.05).
A transition then occurred during weeks 8 and 9 when the treated plants
began producing more flowers than the controls and recovered to a level
equal to the controls. From weeks 10 through 12, flower production in the
VOLUME XC, NUMBER 4
223
1 3 5 7 9 11 13 15
WEEKS
Fig. 1 . Weekly mean height increment curves of eggplant inoculated with different densities
of Colorado potato beetle in 1980. Arrows indicate inoculation dates and the starred arrow
indicates termination of larval feeding. The following are Least Significant Difference Tests of
mean height differences due to differing densities of larval feeding (Cochran and Cox 1957).
For a given week treatments with the same letter are not significantly different, P < 0.05. Week
1 is 23 days post-planting.
Larval density/plant
Weeks
0
8
15
23
30
1-4
N.S.
N.S.
N.S.
N.S.
N.S.
5-6
a
ab
ab
ab
b
7-15
N.S.
N.S.
N.S.
N.S.
N.S.
treated plants ranged from 18 to 71% greater than the controls, but not in
any pattern that could be attributed to larval density levels {P < 0.05).
During the last 3 weeks of the season, there were no differences in flower
production among treatments.
224
NEW YORK ENTOMOLOGICAL SOCIETY
1 3 5 7 9 11 13 15
WEEKS
Fig. 2. Weekly mean leaf area of eggplant inoculated with different densities of Colorado
potato beetle larvae in 1980. Arrows indicate inoculation dates and the starred arrow indicates
termination of larval feeding. The following are Least Significant Difference Tests of mean leaf
area differences due to larval feeding (Cochran and Cox 1957). For a given week treatments
with the same letter are not significantly different, P < 0.05. Week 1 is 23 days post-planting.
Week
Larval density/plant
0
8
15
23
30
1-4
N.S.
N.S.
N.S.
N.S.
N.S.
5
a
b
b
ab
b
6
a
ab
ab
ab
b
7
a
a
ab
ab
b
8-9
N.S.
N.S.
N.S.
N.S.
N.S.
10
b
ab
b
ab
a
11-15
b
ab
ab
ab
a
VOLUME XC, NUMBER 4
225
Fruit production.— ThQ first fruits were harvested 62 days after planting.
Mean fruit weight (478 g ± 6.9 g) were similar among density treatments.
From weeks 7 through 10, the controls yielded 56% more fruit than the
treated plants {P < 0.05) (Fig. 3). During weeks 1 1 through 13, the treated
plants produced between 23 and 65% more fruit than the controls {P <
0.05). Then a transition period ensued, when no yield differences occurred
among treatments. After the transition the 8, 23, and 30 larvae/plant treat-
ments yielded an average of 52% more fruit than the controls {P < 0.05).
There were no significant differences in aggregate yield relative to treatment
for the season, for each treatment produced an average of 10.5 fruit, the
expected yield per plant for the cultivar.
Damaged fruit must also be considered in yield evaluations. The outer
skin must be free from any scars or discoloration to be marketable (U.S.D.A.
1955). The controls and the 15 larvae/plant treatments had 14% damaged
fruit, whereas the 8 larvae/plant treatments had 7% damaged fruit. These
figures represent all fruit damage, since insect injury in this study was not
distinguishable from other types of mechanical injury.
Discussion
For the vegetative and fruit growth variables, the same general pattern
emerges. For the first 4 weeks there was an equal growth rate among all
treatments including the controls. Then vegetative growth was disrupted in
response to late larval feeding. Defoliation causes a decrease in photosynthate
production, which decreases the amount of carbohydrate available for stem,
leaf, and flower production (Wang et al. 1977). The differences which occurred
between the partially defoliated plants and the controls early in the season
apparently were caused by reduced carbohydrate supply. In our experiment
the treatments depressed plant height, leaf and flower production during
weeks 5,6, and 7. These weeks correspond to the time when the larvae were
in 4th instar and the week immediately following termination of feeding.
Following this delay, treated plants compensated for the earlier reduced
growth rate and resumed normal growth. A transition phase then occurred,
in which no treatment differences were found. The treated plants actually
surpassed the controls in their production of leaves, flowers, and fruits. The
shift of peak flower production was also due to plant compensation. Above
8 larvae/plant peak flowering was depressed and fruit production was retarded.
Fuji and Itagi (1954) observed that the fruit-setting cycle of eggplant was
disrupted as a result of defoliation. Also, Saito and Ito (1973) stated that
defoliation caused an increase in the production of short-styled flowers.
These inferior flower forms have a high abortive rate, which may account
for the early delay in fruit production experienced in the higher density
treatments.
226
NEW YORK ENTOMOLOGICAL SOCIETY
Fig. 3. Weekly mean flowers and fruit of eggplant inoculated with different densities of
Colorado potato beetle larvae in 1980. Arrows indicated inoculation dates and starred arrows
indicate termination of larval feeding. The following are Least Significant Difference Tests of
mean flowers and fruit difference due to larval feeding (Cochran and Cox 1957). Within a week
treatments with the same letter are not significantly different, P :< 0.05. Week 1 is 23 days
post-planting. Circles indicate flowers and squares indicate fruit.
Week
Mean flowers/plant
Larval density/plant
0
8
15
23
30
1-4
N.S.
N.S.
N.S.
N.S.
N.S.
5
a
b
b
b
b
6
a
ab
be
be
c
7
a
cd
be
ab
d
8
a
b
a
ab
a
9
N.S.
N.S.
N.S.
N.S.
N.S.
10
b
a
ab
ab
a
VOLUME XC, NUMBER 4
227
During 1980 the highest market price for eggplant coincided with the first
two harvests, after which the market value declined during midseason and
recovered slightly in the last weeks (New Jersey Department of Agriculture
1981). In this study, aggregate yield was the same for all treatments because
late in the season the plants compensated for yield lost earlier in the season
when eggplant value was again high. Therefore, no dollar losses were incurred.
This study represents only the response of eggplant to first generation CPB
feeding. Clearly, an interaction exists between first and second generation
CPB feeding. Studies were planned to examine defoliation effects by both
beetle generations in adjacent plots, but they were terminated due to a large
immigration of CPB adults from neighboring fields which completely defo-
liated the plants.
Acknowledgments
New Jersey Agricultural Experiment Station, Publication No. D-08149-
32-82 supported by state fund and by U.S. Hatch Act. We thank Dr. Richard
Trout consulting statistician for his assistance in data analysis.
Literature Cited
Bailey, L. H. 1891. Botany of the eggplant. N.Y. Cornell Agr. Station Bull. 26:21-26.
Cochran, W. G. and G. M. Cox. 1957. Experimental Design, 2nd Edition. John Wiley &
Sons, Inc., New York. 61 1 pp.
11
12
13-15
Week
b
b
N.S.
ab
a
N.S.
ab
a
N.S.
Mean fruit/plant
ab
ab
N.S.
a
ab
N.S.
0
8
15
23
30
6
N.S.
N.S.
N.S.
N.S.
N.S.
7
a
ab
b
a
b
8
a
ab
b
b
b
9
a
ab
ab
b
c
10
a
b
ab
ab
ab
1 1
b
b
a
a
ab
12
b
ab
b
b
a
13
ab
b
ab
ab
a
14
N.S.
N.S.
N.S.
N.S.
N.S.
15
b
a
ab
a
a
228
NEW YORK ENTOMOLOGICAL SOCIETY
Eguchi, T., T. Matsumura and M. Ashizawa. 1959. The effects of nutrition on flower formation
in vegetable crops. Amer. Soc. for Hort. Sci. 72:343-357.
Free, J. B. 1975. Pollination of Capsicum frutescens L., Capsicum annuum L. and Solarium
me/ongena L. (Solanaceae) in Jamaica. Trop. Agric. 52(4):353-357.
Fuji, T. and T. Itagi. 1954. Studies on cyclic setting of fruit in the eggplant. J. Hort. Assoc.
Japan 23:1-8. (Abstract)
Hare, D. J. 1980. Impact of defoliation by the Colorado potato beetle on potato yields. J.
Econ. Entomol. 37:369-373.
Logan, P. A. and R. A. Casagrande. 1980. Predicting Colorado potato beetle (Leptinotarsa
decemlineata) (Say) density and potato yield loss. Environ. Entomol. 9:659-663.
Mohideen, M. K., C. R. Muthukrishan, A. Rajagopal and V. A. Metha. 1977. Studies on the
rate of flowering, flower types and fruitset in relation to yielding potential of certain
eggplant (Solanum melongena L.) varieties with reference to weather conditions. So.
Indian Hort. 24(2):56-61.
New Jersey Department of Agriculture. 1981. Marketing New Jersey fresh vegetables 1980.
New Jersey Crop Reporting Service. 60 pp.
Prasad, D. N. and R. Prakash. 1 968. Floral biology of brinjal {Solanum melongena L.). Indian
J. Agric. Sci. 38(6):1053-1061.
Saito, T. and H. Ito. 1973. Studies on the flowering and fruiting in eggplant. VIII. Effects of
early environmental conditions and cultural treatments on the development and drop
of flowers. J. Japan. Soc. Hort. Sci. 42(2): 155-162.
Schalk, J. M. and A. K. Stoner. 1979. Tomato production in Maryland: effect of different
densities of larvae and adults of the Colorado potato beetle. J. Econ. Entomol. 72:826-
829.
Smith, O. 1931. Characteristics associated with abortion and intersexual flowers in the eggplant.
J. of Agric. Res. 43(l):83-93.
Tamaki, G. and B. A. Butt. 1978. Impact of Perillus bioculatus on the Colorado potato beetle
and plant damage. USDA Tech. Bull. 1581. 11 pp.
U.S.D.A. 1955. Standards for grades of eggplant. Food and Quality Service. Washington,
D.C. 4 pp.
Wang, Y., A. P. Gutierrez, G. Oster and R. Dasl. 1977. A population model for plant growth
and development: coupling cotton-herbivore interaction. Can. Entomol. 1 09: 1 359-1 374.
Department of Entomology and Economic Zoology, Cook College, New
Jersey Agricultural Experiment Station, Rutgers University, New Brunswick,
New Jersey 08903.
Received for publication November 25, 1981.
NEW YORK ENTOMOLOGICAL SOCIETY
XC(4), 1982, pp. 229-236
PHENOLOGY OF EMERGENCE OF THE SPOTTED TENTIFORM
LEAFMINER, PHYLLONORYCTER GRATAE GELLA
(LEPIDOPTERA: GRACILLARIIDAE) AND
ITS PARASITOIDS IN NEW YORK
Parker Gambino and Daniel J. Sullivan, S J.
Abstract.— A phenological emergence study was carried out in Ulster and
Orange counties, New York, using the overwintering generation of the spot-
ted tentiform leafminer, Phyllonorycter crataegella Clemens and its para-
sitoids, Sympiesis marylandensis Girault, Sympiesis conica (Provancher)
(Hymenoptera: Eulophidae) and Apanteles ornigis Weed (Hymenoptera:
Braconidae). S. marylandensis was found to emerge concurrently with P.
crataegella, followed in order by S. conica and A. ornigis. S. conica also
hyperparasitized A. ornigis and thus served a dual role in this system.
Introduction
The spotted tentiform leafminer, Phyllonorycter crataegella Clemens is a
pest of apple trees in the Northeastern United States. It is assumed to be an
induced pest, resulting from the suppression of its natural controlling factors
due to the use of pesticides intended for other orchard pests. Although the
larvae of P. crataegella do not directly affect the fruit, the stress they place
on the tree can result in reduced yield. High levels of infestation have been
implicated in early fruit drop as well. The long term effects of P. crataegella
upon the host tree are unknown. Recent outbreaks of P. crataegella in the
Northeast, as recorded by Prokopy et al. (1980) and Weires et al. (1980),
have caused concern among growers in the mid-Hudson valley and else-
where. The appearance of organophosphate-tolerant strains (Weires 1977)
has intensified this concern.
Integrated pest management programs, which are gaining acceptance among
apple growers, consist of using various pest control techniques while min-
imizing both adverse environmental effects and farming expenditures. The
timing of insecticide applications is a crucial part of this strategy. Since the
insecticides affect benefical insects as well as pests, it is important to know
at what stages these various insects will be most greatly affected by these
treatments and when these vulnerable stages will be present in the orchard
to be sprayed. Studies of basic insect biology and phenology serve to improve
The publication costs of this article were defrayed in part by page charge payment. This article
must therefore be hereby marked “Advertisement” in accordance with 18 U.S.C. §1734 solely
to indicate this fact.
230
NEW YORK ENTOMOLOGICAL SOCIETY
the efficiency of insecticide use by adding to the information which can be
incorporated into an integrated pest management program.
In the mid-Hudson valley, 3 species of wasps commonly parasitize P.
crataegella: Sympiesis marylandensis Girault, Sympiesis conica (Pro-
vancher), and Apanteles ornigis Weed. Since these wasps are potential control
agents for this moth, their preservation would be a significant factor to be
taken into consideration by growers wishing to synchronize or reduce pes-
ticide applications. By knowing the phenological relationships between the
various stages of P. crataegella and its parasitoids, data gained from pro-
cedures used to monitor flights of adult moths could also serve to predict
the emergence of the adult parasitoids. The goal of the present study is to
determine these phenological relationships.
Biology of the host, Phyllonorycter crataegella. — In the mid-Hudson val-
ley, P. crataegella is trivoltine. Pupae overwinter in fallen leaves and adults
emerge in the spring. The eggs are laid on the undersides of newly sprouted
leaves; upon hatching, the larvae enter the leaves and begin feeding on the
spongy mesophyll layer. The first 3 larval instars puncture the plant cells
with their sharp mouthparts and feed on the liquids which drain from the
cells. These stages are known as sap feeders. Fourth and fifth instar larvae
have chewing mouthparts and take bites from the palisade layer of cells,
causing a white spotted appearance of the leaf when viewed from above.
Silk strands attached to the sides of the mine draw the walls of the mine
inward, causing the upper leaf surface to bulge. Pupation occurs within the
mine. Prior to eclosion, the pupa pokes its head through the underside of
the leaf; the adult emerges, leaving behind the extruding pupal exuvium.
The second and third generations follow the same basic pattern. Approxi-
mate periods of adult emergence in the mid-Hudson valley, which vary
according to the weather conditions, are: April, for the overwintering gen-
eration; late June, for the first generation; mid-August, for the second gen-
eration.
Emergence of the overwintering generation is fairly well synchronized, but
later in the season, individual larvae develop at widely differing rates. As a
result, compared to the overwintering generation, the adult flight of the first
generation is more spread out in time. By July, all of the life stages are
present simultaneoulsy in an orchard. The final flight in August is even more
poorly synchronized. Chemical control efforts for this moth have been con-
centrated in the spring, not only to “nip it in the bud,” but also because this
is the time when it is assured that those moths present will be in the same
or similar stages of development.
Biology o/ Sympiesis marylandensis and Sympiesis conicdi. — Sympiesis
marylandensis and Sympiesis conica are quite similar in their life histories
and habits regarding P. crataegella on apple, and details presented here
pertain to both species unless otherwise noted. While a Sympiesis individual
VOLUME XC, NUMBER 4
231
may overwinter as a larva within the mine of the consumed host, the more
common situation is overwintering in the pupal stage. After eclosion within
the mine, the adult chews a small hole in the leaf and exits.
Behavior of the adults after emergence is not well known. According to
Beckham et al. (1950), oviposition by the Sympiesis species occurs on hosts
in the tissue-feeding stages. Evidence presented by Askew (1979) suggests
that in other representatives of the genus Sympiesis, adult females may also
affect mortality of their Phyllonorycter hosts by feeding on the early sap-
feeding stages.
A Sympiesis larva feeds as an ectoparasite and usually consumes the host,
preventing it from reaching the pupal stage. The wasp larva pupates after
voiding a small characteristic meconium and extruding a short thread through
the anus which serves to anchor it within the mine. In the area of study, at
least 3 generations per year occur when P. crataegella serves as host.
Biology of Apanteles omigis.—Apanteles ornigis is an endoparasitoid of
P. crataegella larvae. As in the Sympiesis species, the tissue-feeding stages
of P. crataegella are selected for oviposition by A. ornigis females. Although
Pottinger and LeRoux (1971) report that overwintering by A. ornigis may
occur naked in mines of P. blancardella, my observations are in agreement
with those of Beckham et al. (1950), who found A. ornigis overwintering
only within a cocoon. A. ornigis is the only species of the parasitoid complex
of P. crataegella which constructs a cocoon; it is an elongate white oval
which is attached by a silk thread at each end to the walls of the mine. The
actual overwintering stage was undetermined, since it was concealed within
the cocoon. Upon eclosion, the adult makes a circular incision so that one
end of the cocoon opens as a flap. To emerge from the leaf mine, a small
hole is chewed through the leaf epidermis.
Materials and Methods
During the apple growing season of 1979, several orchards in Ulster County
and Orange County, New York, were surveyed in order to locate infestations
of Phyllonorycter crataegella and to determine which parasitoids were attack-
ing it. The presence of P. crataegella was apparent to the naked eye, the
apple leaves showing the typical spotted mines where moth larvae were
present. To locate populations of parasitoids, mined leaves were dissected.
Pupating parasitoids were placed individually in #3 gelatin capsules, so that
emerging adults could be identified.
The following sites in Ulster County were selected as locations for the
collection of data: Turkey Hill plot, Macintosh Farm plot and Home plot,
VanDuser Orchards, Wallkill; DiStefano Orchards, Modena; E. Wright
Orchards, New Paltz; Haetzler Orchards, Wallkill; and Gerken Orchards,
Wallkill. Data were also collected from Finelli Orchards in Savilton, Orange
232
NEW YORK ENTOMOLOGICAL SOCIETY
County. All collections were made from orchards in commercial production,
receiving regular spray programs.
Only the Finelli Orchards site showed a sizeable population of A. ornigis,
and a separate set of rearings was carried out using material from this site;
description of this rearing experiment will appear near the end of this section.
Since the host moths and all of the parasitoids overwinter within the fallen
apple leaves, collections of live insect material could be made by gathering
leaves from the ground during the winter. Leaves from the various test sites
were collected during one of two field trips, the first in December 1979 and
the second in March 1980. Infested leaves were placed into rearing chambers
made from empty one gallon translucent plastic jugs. The top of each jug
was removed and replaced with a clear plastic specimen jar. Although the
juncture of the jug and the jar was not airtight, the space between the two
was too narrow to allow any of the insects under study to escape.
In an attempt to keep conditions within the rearing chambers as similar
as possible to conditions in the orchards of origin, chambers were held out
of doors on the porch of the Gambino residence, Bronx, New York. There
they received afternoon sun and were sheltered from precipitation. A wooden
enclosure, open on top, protected them from wind disturbance, but allowed
light to reach the plastic jars at the tops of the chambers.
Insects emerging from the leaves in a chamber were attracted to the light
at the top and gathered in the clear jar. All adults were collected on a daily
basis, using an aspirator. After recording the date and chamber number, the
insects were identified and placed individually in #3 gelatin capsules for
future reference.
For the Finelli orchard, in addition to the procedure described, a more
intensive rearing experiment was conducted. Infested leaves were gathered
in March 1980, and leaves bearing P. cmtaegella mines were dissected until
100 healthy occupants were found. Health was judged from the appearance
of the pupa; if it was not crushed or dented, and showed no signs of fungus
growth, then it was selected. In the case of an A. ornigis cocoon, it was
impossible to determine the actual condition of the inhabitant, so the out-
ward appearance of the cocoon was used as a guide.
Insects were placed individually into #3 gelatin capsules and held indoors.
The capsules were kept near a window, and thus were exposed to a natural
photoperiod. Room temperature was approximately 70°F. As insects emerged,
a record was made of the date, species of insect, and whether emergence was
from an Apanteles cocoon or a naked pupa.
Determination of P. crataegella was made by Dr. Don Davis, Smithsonian
Institution, Washington, D.C. Determinations of the 2 Syrnpiesis species
were made by Dr. E. E. Grissell, U.S. National Museum, Washington, D.C.,
and the determination of A. ornigis was made by Dr. Paul Marsh, also of
the U.S. National Museum.
VOLUME XC, NUMBER 4
233
Fig. 1 . Rearing chamber emergence frequencies of overwintering insects.
Results
The data gathered from the rearing chambers, pertaining to the emergence
of P. crataegella, S. marylandensis and S. conica are combined and pre-
sented in Fig. 1, while Table 1 lists significant emergence dates for these 3
species. Due to the low number of individuals recovered, Apanteles ornigis
(3) and other unidentified hymenopterous species (6) were not included in
the presentation of rearing chamber data.
Data from the separate rearing of insects from the Finelli orchard are
presented in Table 2. Of the 100 pupae or cocoons isolated in capsules, 82
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NEW YORK ENTOMOLOGICAL SOCIETY
Table 1 . Rearing chamber emergence dates of P. crataegella, S. rnarylandensis and S. conica.
Dates of emergence
N
First
Last
Median
P. crataegella
41
3 April
27 April
18 April
S. rnarylandensis
230
3 April
2 May
20 April
S. conica
56
18 April
2 July
3 May
yielded adult insects; the 1 8 mortalities did not receive further consideration.
In addition to the 4 species which are the main concern of this study, an
unidentified fifth species, recovered only as an ornigis hyperparasite, was
also present in the Finelli orchard. Data concerning the occurrence of this
unidentified species are included in Table 2, but are not subject to further
analysis.
Discussion
The rearing chamber data (Fig. 1) show that P. crataegella adults appeared
in April, with most (34/41 or 83%) emerging between 12 April and 22 April
inclusive. With a larger sample, it is likely that recordings would have been
made in late March and early May. During this time, the first leaves of the
apple trees were also appearing, so that the behavior of the moths was well
synchronized with phenological events of the host plant; young leaves were
available to serve as oviposition sites.
Like P. crataegella, S. rnarylandensis emerged primarily during April. This
species is recorded by Beckham et al. (1950) as an ectoparasitoid of P.
crataegella fourth and fifth instars. At the very least, there would be 2 weeks
between S. rnarylandensis emergence and appearance in the field of these
suitable stages of P. crataegella. A more typical time interval would be 4
weeks. As has been previously noted, post-emergence behavior of these
wasps is poorly known. Studies currently underway may shed some light on
what S. rnarylandensis does during the weeks prior to oviposition. Despite
its seemingly early emergence, S. rnarylandensis has become the dominant
P. crataegella parasitoid in the area of study.
Sympiesis conica presented an emergence pattern quite different from that
of S. rnarylandensis. Initial S. conica appearance on 1 8 April was more than
2 weeks after the first S. rnarylandensis emergence, and occurred while the
latter was reaching peak emergence. Overwintering S. conica continued to
emerge until 2 July, showing a rather diffuse pattern of emergence. When
the emergence data from P. crataegella and the 2 Sympiesis species are
analyzed using a median contingency table test, the hypothesis of identical
median emergence dates is rejected at the 0.0 1 level. When just P. crataegella
and S. rnarylandensis are considered using the median test, no significant
VOLUME XC, NUMBER 4
235
Table 2. Emergence dates for Finelli orchard material.*
Date
P.c.
S.m.
S.c.
A.o.
Other
4/19
2
4/20
2
4/21
1
4/22
1
15
4/23
8
4/24
4
4/25
6
1
4/26
4
1
4/27
2**
4/28
2
4/29
2
3**
2**
4/30
3**
5/1
1
5/2
2
2
5/3
1
2
5/4
2
5/5
2
5/6
3
5/7
3
2**
5/8
1
5/9
1
5/10
5/11
1
* Key: P.c.—P. crataegella\ S.m.—S. marylandensis\ S.c.—S. conica; A.o.—A. ornigis.
** Indicates hyperparasitism.
difference in the median emergence date is found (P < 0.05), confirming
what Fig. 1 and Table 1 suggest: P. crataegella and S. marylandensis emerge
concurrently, while S. conica appears later in the spring. Indeed, since by 2
July development of the second generation of P. crataegella larvae was
underway, adults of the overwintering generation of S. conica overlapped
with 2 generations of hosts in the field.
Several additional observations support the contention that S. conica
emerges later than S. marylandensis. When later generations of parasitoids
are considered during the growing season, S. conica adults are seen to emerge
consistently later than those of S. marylandensis. Gathering accurate data
on these later generations is difficult, due to the fact that individuals of a
population are not well synchronized as they pass through developmental
stages and generations may overlap. In the case of parasitoids found to be
overwintering as larvae, the majority of those successfully reared turn out
to be S. conica. Occasionally a parasitized mine contains a parasitoid pupa
and the remains of a P. crataegella pupa. These rearings also yield S. conica.
In the case of hyperparasitized A. ornigis, S. marylandensis was not found
to function as an A. ornigis hyperparasitoid on apple, while S. conica was
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NEW YORK ENTOMOLOGICAL SOCIETY
commonly recovered from A. ornigis cocoons (Table 2). This would suggest
that oviposition by S. conica occurred after A. ornigis had reached its cocoon
spinning stage. Thus it appears that the developmental schedule of S. conica
lags behind that of S. rnarylandensis throughout the year.
The limited data on A. ornigis suggest that it is the final species of this
complex to appear in the spring. However, this generalization certainly would
not apply to all individuals. S. conica, with its extreme variability in emer-
gence dates, actually overlapped with all 3 other species, and was the species
with the final recorded emergence from the rearing chambers. Data from
Table 2 indicate that A. ornigis emerges after S. conica. These data are
consistent with those of Johnson et al. (1979), who found that in Ontario,
Canada, the difference in median emergence dates between P. blancardella
(a species similar to P. crataegella) and A. ornigis was approximately 3 weeks.
A. ornigis is heavily hyperparasitized (17 of 35, or 48%), hyperparasitism
being determined when an A. ornigis cocoon yielded an adult of a species
other than A. ornigis. It was necessary to dissect mines in order to assess
the impact of hyperparasitism, and this time-consuming procedure was fol-
lowed only for the additional series of rearings carried out using material
from the Finelli orchard. (To the author’s knowledge, this is the first record
of A. ornigis serving as a host for S. conica.). Data collected from the rearing
chambers cannot give an accurate picture of either the presence or degree
of hyperparasitization of A. ornigis, especially since the predominant A.
ornigis hyperparasitoid, S. conica, also attacks the primary host, P. cratae-
gella.
Literature Cited
Askew, R. R. and M. R. Shaw. 1979. Mortality factors affecting the leafmining stages of
(Lepidoptera: Gracillariidae) on oak and birch. 1 . Analysis of the mortality
factors. 2. Biology of the parasite species. Zool. J. Linn. Soc. 67(l);31-64.
Beckham, C. M., W. S. Hough and C. H. Hill. 1950. Biology and control of the spotted
tentiform leafminer on apple trees. Va. Agric. Exp. Sta. Bull. 1 14:3-12.
Johnson, E. F., R. Trottier and J. E. Laing. 1979. Degree-day relationships to the development
of Lithocolletis blancardella (Lepidoptera: Gracillariidae) and its parasite Apanteles orni-
gis (Hymenoptera: Braconidae). Can. Ent. 1 1 1:1 177-1 184.
Pottinger, R. P. and E. J. LeRoux. 1971. The biology and dynamics of Lithocolletis blancardella
(Lepidoptera: Gracillariidae) on apple in Quebec. Mem. Ent. Soc. Canada 77:41-91.
Prokopy, R. J., R. G. Hislop and W. M. Coli. 1980. Spotted tentiform leafminers: biology,
monitoring, and control. Fruit Notes 45(2):7-12.
Weires, R. W. 1977. Control of Phyllonorycter crataegella in eastern New York. J. Econ. Ent.
70(4):521-523.
, D. R. Davis, J. R. Leeper and W. H. Reissig. 1980. Distribution and parasitism of
gracillariid leafminers on apple in the Northeast. Ann. Ent. Soc. Amer. 73(5):54 1-546.
(PG) Department of Entomological Sciences, University of California,
Berkeley, California and (DJS) Assoeiate Professor, Department of Biological
Sciences, Fordham University, Bronx, New York.
Received for publication March 1, 1982.
NEW YORK ENTOMOLOGICAL SOCIETY
XC(4), 1982, pp. 237-240
TISSUE GLYCOGEN OF MYLABRIS PUSTULATA THUNB.
AND PERIPLANETA AMERICANA L. (INSECT A)
Dalbinder Singh Sidhu, Surinder Pal Kaur and Nirmal Kumar
Abstract.— glycogen contents in the fat bodies, thoracic muscles, fore-
gut, midgut, hindgut, ovaries and testes of Mylabris pustulata and Peripla-
neta americana have been studied. In M. pustulata, the major depot of
glycogen contents (3.42 mg/g) has been noted in the fat bodies, while it is
not so in P. americana. The midgut of both insects under investigation
shows a higher level of glycogen than the rest of the tissues except fat bodies.
In M. pustulata, the glycogen contents of ovaries are at a higher level than
that of testes. In P. americana, the glycogen reserves are comparatively
higher in ovaries and thoracic muscles than M. pustulata. The glycogen
contents. It is known that insects contain digestive amylase in the midgut
ferences {P > 0.05) while the rest of the tissues contain significantly different
contents (P < 0.05).
Introduction
Glycogen, an important carbohydrate reserve in insects, is the first source
of energy to be utilized under stress of starvation (Strauss 1911; Hill and
Goldsworthy 1970). Most of the information with regard to the glycogen
reserves in insects, available until recently, deals with its estimations from
the whole body extracts; which of course does not provide any clue regarding
the capacity of different tissues to retain and synthesize this compound.
Moreover, work on individual tissues permits an insect biochemist to com-
pare his results with the mammalian tissues where experimentation is not
done on the whole body extracts. The distribution of glycogen in the tissues
of insects has been studied in the past only by a few workers like Yeager
and Munson (1941) in insect muscles, Wigglesworth (1949) in abdominal
fat body and Nemec (1977) in the ovaries. The study in hand deals with the
glycogen estimations from the different tissues of Mylabris pustulata and
Periplaneta americana, and clearly, the aim is to elaborate this field so that
in the long run some general and coherent trend in this respect can be
deduced.
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238
NEW YORK ENTOMOLOGICAL SOCIETY
Table 1. Distribution of glycogen in different tissues of Mylabris pustulata and Periplaneta
americana.
Tissues
mg glycogen ±
SD/g wet tissue weight!
M. pustulata
P. americana
Foregut
1.51 ± 0.032**
1.56 ± 0.016**
Midgut
3.17 ±0.110**
2.43 ± 0.123**
Hindgut
1.49 ± 0.036
1.16 ± 0
Ovaries
0.781 ± 0.004*
1.46 ± 0.024
Testes
0.660 ± 0
—
Fat bodies
3.42 ± 0.012
2.12 ± 0.008
Thoracic muscles
1.02 ± 0
1.67 ± 0.018
t Each value mentioned in the table is at least an average of five determinations. SD denotes
standard deviation. P values have been calculated by applying Student’s /-test of significance.
* P < 0.05.
**p < 0.01.
Materials and Methods
Mature adults of Mylabris pustulata were collected from kitchen gardens,
and of Periplaneta americana from places such as kitchens and storerooms
etc. from the Campus of Punjabi University, Patiala, India. Both the insects
were dissected under physiological saline to obtain tissue samples from their
respective foregut, midgut, hindgut, ovaries, testes, fat bodies and thoracic
muscles. The samples taken were weighed and digested independently in
30% KOH for the extraction of glycogen. For extraction, the method of
Heatly (1935) was employed while the estimation of glycogen was made
colorimetrically according to Montgomery (1957).
Results and Discussion
The results of estimations of glycogen from different tissues of M. pustulata
and P. americana are listed in Table 1.
In M. pustulata, the maximum glycogen contents have been recorded in
the fat bodies, which indicates that this tissue is the major storage site for
this energy reserve. This finding is in accordance with the observations of
Kilby (1963), Gilmour (1965) and Wyatt (1967). However, in P. americana,
the fat body glycogen, though quite high is little less than that of its depot
in midgut.
The midgut of both the insects under study is quite rich in glycogen
contents. It is known that insects contain digestive amylase in the midgut
which hydrolyses dietary starch and glycogen and hence the absorption of
digested food material takes place in this region. Treherne (1958) in Peri-
planeta and Schistocerca has observed that the absorption of glucose is largely
confined to the midgut caeca. Thus the presence of a somewhat high con-
VOLUME XC, NUMBER 4
239
centration of glycogen in the midgut of both the insects suggests that a portion
of the glucose after absorption gets converted into glycogen in this region of
the alimentary canal. Wigglesworth (1949) in Drosophila and Mayer and
Candy (1 969) in Locusta have also observed that the midgut cells themselves
are known to contain glycogen stores or it can be said that the glycogen
anabolism takes place in the midgut cells. This is perhaps because the
midgut converts surplus glucose into glycogen which is not readily utilized
by the insect body.
In M. pustulata, the glycogen contents of ovaries are at a higher level than
that of testes (Table 1). This is perhaps because of the necessity of this car-
bohydrate for vitellogenesis and for the formation of glycosaminoglycans
present in the vitelline membrane and the chorion (Rockstein 1978). The
glycogen which also serves as the principal carbohydrate in yolk, is usually
synthesized in the ovary from glucose and trehalose derived from the fat
body and the hemolymph. Similarly, the ovaries of P. americana also contain
high concentration of glycogen contents. Moreover, it is known that the
glycogen is also synthesized in the ovaries during the terminal phase of
vitellogenesis (Rockstein 1978).
A comparison between the glycogen contents of foregut of M. pustulata
and P. americana shows no signihcant difference {P > 0.05). Generally the
foregut of insects is not involved in the absorption of glucose, but in Peri-
planeta, the crop has been found to have little importance in the absorption
of glucose (Eisner 1955). According to above mentioned author, some lipase
activity exists in the crop of Periplaneta as the result of transfer of secretions
from the midgut into the foregut. So the presence of newly formed fatty
acids facilitate a little absorption of glucose through the crop cuticle. Thus,
the presence of more glycogen contents in the foregut of P. americana con-
firms that some absorption also takes place in the crop region of the foregut.
As the foregut of M. pustulata shows no significant difference in this energy
reserve from that of P. americana, it can be suggested that the foregut region
of Mylabris too might be involved in the absorption of glucose like that of
Periplaneta.
The glycogen contents of thoracic muscles of P. americana are significantly
higher than that of M. pustulata (P < 0.05). It has been already reported
that P. americana uses carbohydrate as the major source of energy for flight
(Polacek and Kubista 1960) and the glycogen reserves of thoracic muscu-
lature comprise the major source of the utilized substrate (Downer and
Matthews 1976). So the presence of more glycogen in the thoracic muscles
of P. americana shows that it is more efficient in using this reserve than M.
pustulata.
By comparing the glycogen contents of the two insects, it is evident that
there exists significant variation with respect to midgut, hindgut, ovaries,
fat bodies and thoracic muscles {P < 0.05) revealing a species specific dis-
240
NEW YORK ENTOMOLOGICAL SOCIETY
tinctness in the metabolism of this compound in these tissues. At 1% level
of significance, there is no difference in the glycogen reserves of foregut,
midgut and hindgut. Although at 5% level of significance, foregut is the only
tissue in the two insects where there exists no variation.
Literature Cited
Downer, R. G. H. and J. R. Matthews. 1976. Glycogen depletion of thoracic musculature
during flight in Periplaneta americana L. Comp. Biochem. Physiol. 5 5B: 50 1-502.
Eisner, T. 1955. The digestion and absorption of fats in the foregut of the cockroach. Peri-
planeta americana L. J. Exp. Zool. 130:159-182.
Gilmour, D. 1965. The metabolism of insects. Edinburgh, Oliver and Boyd, xii + 195 + (1),
p. 33.
Heatly, N. G. 1935. The distribution of glycogen in the regions of amphibian gastrula; with
a method for the microdetermination of glycogen. Biochem. J. 29:2568.
Hill, L. and G. J. Goldsworthy. 1970. The utilisation of reserves during starvation of larvae
of the migratory locust. Comp. Biochem. Physiol. 36:61-70.
Kilby, B. A. 1963. The biochemistry of the insect fat body. In: Advances in Insect Physiology
1:11 1-174. Academic Press, London, New York.
Mayer, R. J. and D. J. Candy. 1969. Changes in energy reserves during flight of the desert
locust, Schistocerca gregaria. Comp. Biochem. Physiol. 31:409-418.
Montgomery, R. 1957. Determination of glycogen. Arch. Biochem. Biophys. 67:378-386.
Nemec, V. 1977. Changes in saccharide content in various organs of Pyrrhocoris apterus
during its reproductive cycle. Acta Entomol. Bohemoslov. 74(4):2 19-223.
Polacek, I. and V. Kubista. 1 960. Metabolism of the cockroach, Periplaneta americana during
flight. Physiol. Bohemoslov. 9:228-234.
Rockstein, M. 1978. The functions of carbohydrates in insect life processes. In: M. Rockstein
(ed.). Biochemistry of Insects. Academic Press, New York.
Strauss, J. 1911. Chemical composition during growth and metamorphosis: Apis. Z. Biol. 56:
347-397.
Treheme, J. E. 1958. The absorption of glucose from the alimentary canal of the locust,
Schistocerca gregaria. J. Exp. Biol. 35:297-306.
Wigglesworth, V. B. 1949. Metabolism during the flight and starvation of Drosophila. J. Exp.
Biol. 26:150-163.
Wyatt, G. R. 1967. The biochemistry of sugars and polysaccharides in insects. In: J. W. L.
Beament, J. E. Treheme and V. B. Wigglesworth (eds.). Advances in Insect Physiology
4:287-347. Academic Press, New York.
Yeager, J. F. and S. C. Munson. 1941. Glycogen in Prodenia. J. Agric. Res. 63:257-294.
Department of Zoology, Punjabi University, Patiala- 147002, India.
Received for publication March 3, 1982
NEW YORK ENTOMOLOGICAL SOCIETY
XC(4), 1982, pp. 241-246
INFLUENCE OF WEATHER ON PREDATOR/PREY RELATIONS:
STINKBUGS AND TENT CATERPILLARS
Edward W. Evans*
Abstract. — Coo\ spring temperatures suppressed the activity of predatory
stinkbugs {Podisus spp.) but did not prevent the bugs’ prey, tent caterpillars
(Malacosoma americanum (F.)), from feeding and developing rapidly. Thus
a cold snap in 1977 temporarily reduced the number of stinkbugs at cat-
erpillar tents near Ithaca, New York, and enabled the prey to escape pre-
dation by growing to large sizes before warm weather returned.
Introduction
The physical environment can have an important influence on the effec-
tiveness with which entomophagous insects reduce numbers of their prey.
Studies in biological control reveal that climate often limits the effectiveness
of both introduced and native natural enemies attacking insect pests: climatic
conditions characterizing particular seasons and/or geographic regions often
adversely affect the predators more than their prey (Messenger et al. 1976).
Connell (1970) has suggested that not just the prevailing climate, but also
fluctuations in weather may reduce the effectiveness of natural enemies because
they are often more vulnerable to such fluctuations than are their prey.
Examples include entomophagous insects that experience proportionately
greater mortality than their prey during unusually cold winters (e.g.. Lord
and MacPhee 1953; Clausen 1958). In an intriguing extension of his hypoth-
esis, Connell (1975) suggests that during the growing season, activity of
natural enemies may be reduced temporarily during short periods of unfa-
vorable weather. Prey may escape predation by growing to sizes invulnerable
to predation during these relatively predator-free periods. Here I present an
example illustrating the occurrence of this phenomenon among predatory
insects and their prey.
Each spring in eastern North America, colonies of the eastern tent cat-
erpillar {Malacosoma americanum (F.), Lasiocampidae) are attacked at the
silken tents that they spin by predatory stinkbugs (Hemiptera: Pentatomidae)
of the genus Podisus (Sullivan and Green 1950; Evans 1982). The following
‘ Present address: Division of Biology, Kansas State University, Manhattan, Kansas 66506.
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must therefore be hereby marked “Advertisement” in accordance with 18 U.S.C. §1734 solely
to indicate this fact.
242
NEW YORK ENTOMOLOGICAL SOCIETY
observations document the influence of the vagaries of spring weather on
this predator/prey relationship.
Methods
In 1 977-79 I examined the seasonal abundance of the stinkbugs in relation
to the ontogenetic development of the tent caterpillars in a stand of scattered,
small black cherry {Prunus serotina Ehrh.) and apple (Malus sylvestris Mill.)
trees near Ithaca, New York. The predators were censused on various dates
by counting the number of individuals observed per prey tent in 1977 and
per tent-bearing tree in 1978 and 1979. At individual censuses, generally
made in the afternoon, 50-250 tents were checked in 1977, 50-100 trees in
1978, and 50 trees in 1979. In all years, the entire surface of trees bearing
tents was searched for stinkbugs.
The ontogenetic development of caterpillars was monitored each year by
determining the instar and body length (mm) of representative individuals
on different dates. Caterpillars were sampled by selecting randomly two
caterpillars in 1977 and one in 1979 from the surface of 20-50 tents. A
similar procedure was used to collect 150 caterpillars on 20 May 1978; on
other occasions in 1978, the average length of prey was estimated by mea-
suring representative individuals in the field.
Daily average air temperatures at 60 inches (152 cm) above ground for
Ithaca, New York, were obtained from the Monthly Meteorological Sum-
mary for April and May 1977-79 prepared by the Division of Atmospheric
Sciences, Cornell University. Voucher specimens of the stinkbugs have been
deposited in the Cornell University Insect Collection (Lot No. 1086).
Results and Discussion
Each year stinkbugs first appeared at tents soon after the caterpillars hatched
in April; the bugs peaked in numbers in May when the prey averaged 12-
14 mm in length and were mostly in the fourth instar (Fig. 1). Three species
of stinkbugs (Podisus maculiventris (Say), P. placidus Uhl., and P. modestus
Uhl.) appeared at tents; they are treated together here (see Evans 1982 for
detailed analyses of each species’ exploitation of tent caterpillars). Stinkbug
numbers at tents gradually declined after this peak in 1978 and 1979 as the
caterpillars matured and abandoned tents in search of pupation sites. In
1977, this seasonal pattern was disrupted by inclement weather in early May.
Stinkbug densities at tents during this time were clearly dependent on ambient
temperature (Fig. 2). After peaking on 1 May, stinkbug numbers plummeted
during the cold snap that followed (all three species’ activity was suppressed).
On seven of the next ten days, the average air temperature was less than
1 0°C; the mean temperature during this ten-day period was 9°C (the mean
VOLUME XC, NUMBER 4
243
Fig. 1 . The densities of stinkbugs at tents (solid lines) expressed as the number of bugs per
100 tents in 1977 and per 100 tent-bearing trees in 1978 and 1979, and the mean length (mm)
of tent caterpillars (dashed lines) on various dates each spring.
temperature for 2-1 1 May was calculated as the mean of the ten daily average
temperatures). In contrast, warmer weather prevailed during the same stage
of the caterpillars’ development in 1978 and 1979; the mean temperatures
were 17°C and 14°C during the comparable ten-day periods (i.e., after cat-
erpillars had grown on average to 1 3 mm in length and were fourth instars)
in 1978 and 1979, respectively. Tent caterpillars were observed to feed
actively during the cold snap in 1977; even the several inches of wet snow
244
NEW YORK ENTOMOLOGICAL SOCIETY
APRIL MAY
Z
C
2
DD
m
3D
o
*n
03
c
O
C/)
T3
m
33
O
O
m
Z
H
0)
Fig. 2. The densities of stinkbugs at tents (solid line) and the average air temperature for
various sampling dates, 30 April-16 May 1977 (temperatures were obtained from Monthly
Meteorological Summaries prepared by the Division of Atmospheric Sciences, Cornell Uni-
versity).
that fell on 9 May failed to slow their aetivity for long. When stinkbugs
reappeared at tents in large numbers with the return of warm weather on
12 May (Fig. 2), the caterpillars had achieved large sizes (Fig. 1).
The cold weather limited the effectiveness of the predators in two ways.
First, by preventing predators from being active during a period of vigorous
growth of the prey, the cold weather reduced the exposure of tent caterpillars
to stinkbug attack. Secondly, it also enabled many caterpillars to escape
stinkbug predation after warm weather returned. By the time bugs reappeared
in large numbers, many prey were too large for the predators to subdue. In
held observations of encounters between stinkbugs and healthy caterpillars
>25 mm long, the predators (9-12 mm long) were repulsed by the vigorous
side-to-side thrashings of the prey and soon abandoned the attack. Unlike
earlier in the spring, the prey upon which the predators were found feeding
after the cold snap were on average smaller than individuals in the caterpillar
population at large: the median length of prey taken from stinkbugs on 1 1-
17 May was 23 mm (A^ = 23) vs. a median of 27 mm (A^= 21) in the
VOLUME XC, NUMBER 4
245
caterpillar population at large on 1 1 May {P = 0.06, Wilcoxon two-sample
test).
The quality as well as the relative size of prey attacked by stinkbugs also
changed after the cold snap. Many stinkbugs were found after the cold snap
at “old” tents from which most caterpillars had dispersed. Those few strag-
glers that remained to fall prey to the bugs generally were sluggish, inactive
caterpillars. Probably many of these individuals were diseased and/or par-
asitized; Myers (1981), for example, reports that up to 90% of fifth-instar
larvae of the western tent caterpillar {Malacosoma californicum pluviale
(Dyer)) with delayed maturation are parasitized by tachinid flies. In a similar
predator/prey interaction, Podisus maculiventris probably also acts selec-
tively to remove parasitized and otherwise weakened larvae (with retarded
development and little ability to defend themselves) from older tents of the
fall webworm, Hyphantria cunea (Drury) (Morris 1963). Even the healthy
tent caterpillars among the stragglers at old tents may have had reduced
dispersal and reproductive capacity; such is the case for inactive larvae of
M. californicum pluviale, which emerge from eggs containing few food reserves
(Wellington 1977). Thus it appears that effective predation (i.e., killing of
individuals not already destined for an early death) by stinkbugs was con-
siderably reduced not only during but also after the cold snap in comparison
to before.
In numerous case studies of biological control, natural enemies have proven
more susceptible to pesticides than the insects that they attack (Messenger
et al. 1976). The present example, in which fluctuation in weather adversely
affects predatory insects more than their prey, also emphasizes the relative
sensitivity of entomophagous insects to environmental perturbation. Further
study is needed to test whether predation as a way of life for insects may,
in comparison to herbivory, generally entail greater exposure and vulnera-
bility to vicissitudes of the physical environment.
Acknowledgments
I thank P. Bierzychudek, P. Kareiva, and R. B. Root for their comments
on the manuscript. Support was provided by the Section of Ecology and
Systematics and the Department of Entomology, Cornell University.
Literature Cited
Clausen, C. P. 1958. Biological control of insect pests. Annu. Rev. Entomol. 3:291-310.
Connell, J. H. 1970. On the role of natural enemies in preventing competitive exclusion in
some marine animals and in rain forest trees. Pages 298-312 in: P. J. den Boer and G.
Gradwell (eds.). Dynamics of Populations. Proc. Adv. Study Inst. Dynamics of Numbers
in Populations, Oosterbeek, Netherlands.
. 1975. Some mechanisms producing structure in natural communities: a model and
246
NEW YORK ENTOMOLOGICAL SOCIETY
evidence from field experiments. Pages 460-490 in: M. L. Cody and J. M. Diamond
(eds.). Ecology and Evolution of Communities. Belknap Press, Cambridge, Massachu-
setts.
Evans, E. W. 1982. Niche relations of predatory stinkbugs {Podisus spp., Pentatomidae)
attacking tent caterpillars (Malacosoma americanum, Lasiocampidae). Amer. Midi. Nat.,
in press.
Lord, F. T. and A. W. MacPhee. 1953. The influence of spray programs on the fauna of apple
orchards in Nova Scotia. VI. Low temperatures and the natural control of the oystershell
scale, Lepidosaphes ulmi (L.) (Homoptera: Coccidae). Can. Entomol. 85:282-291.
Messenger, P. S., F. Biliotti and R. van den Bosch. 1976. The importance of natural enemies
in integrated control. Pages 543-563 in: C. B. Huffaker and P. S. Messenger (eds.). Theory
and Practice of Biological Control. Academic Press, New York.
Morris, R. F. 1963. The effect of predator age and prey defense on the functional response
of Podisus maculiventris Say to the density of Hyphantria cunea Drury. Can. Entomol.
95:1009-1020.
Myers, J. H. 1981. Interactions between western tent caterpillars and wild rose: a test of some
general plant herbivore hypotheses. J. Anim. Ecol. 50:1 1-25.
Sullivan, C. R. and G. W. Green. 1 950. Reactions of larvae of the eastern tent caterpillar and
of the spotless fall webworm to pentatomid predators. Canad. Entomol. 82:52.
Wellington, W. G. 1977. Returning the insect to insect ecology: some consequences for pest
management. Environ. Entomol. 6:1-8.
Section of Ecology and Systematics, Cornell University, Ithaca, New York
14853.
Received for publication April 20, 1982.
NEW YORK ENTOMOLOGICAL SOCIETY
XC(4), 1982, pp. 247-251
SEASONAL OCCURRENCE OF FOUNDING QUEENS AND
THE SEX RATIO OF CAMPONOTUS PENNSYLVANICUS
(HYMENOPTERA: FORMICIDAE) IN NEW JERSEY
Harold G. Fowler and Radclyffe B, Roberts
Abstract. — Pounding queens of Camponotus pennsylvanicus were collected
in the field from early March through the middle of July from 1 977 to 1 98 1 .
No synchronization of population nuptial flights was detected. Sex ratios of
colonies are male biased, contrary to previous reports.
The carpenter ant, Camponotus pennsylvanicus (DeGeer), is one of the
most ubiquitous of all urban insects in the Northeast. As early as 1910,
Wheeler reported on its synanthropic adaptations, and, if anything, this
relationship has grown as urbanization has progressed. In spite of this fact,
our knowledge of its biology has not notably improved since Pricer (1908)
published his detailed studies. McCook (1883a, b), Blochmann (1885), and
Pricer (1 908) have described the initiation of an incipient colony by founding
queens. Here, we describe the seasonal distribution of the nuptial flights,
and the sex ratio of field colonies, both of which are important to under-
standing the reproductive strategy of this insect. We also discuss the impli-
cations of our observations on the organization of its population structure.
Methods
The data we report on here is largely field data that we have collected
from 1977 to 1981, on the captures of alate or recently dealated queens of
C. pennsylvanicus in central New Jersey. However, the data from 1979 is
derived from student collections, as we were unable to collect during that
year.
Sex ratio information was taken from colonies collected in their entirety
in early spring, employing a modification of VanderschafFs (1970) method.
In all cases, colonies were collected before the mode of occurrences of nuptial
flights, as inferred from our field data. Dry weights were determined for 30
males and females, and were used for subsequent interpretations.
Statistical tests employed here are of goodness of fit of distributions: the
Kolmogrov-Smimov test statistic, D\ and the Log-Likehood Ratio test sta-
tistic, G (Zar 1974).
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must therefore be hereby marked “Advertisement” in accordance with 18 U.S.C. §1734 solely
to indicate this fact.
248
NEW YORK ENTOMOLOGICAL SOCIETY
Fig. 1 . The distribution of field collections of alate or recently dealate queens of C. penn-
sylvanicus from 1977 through 1981, in central New Jersey.
Results
A plot of the data of the occurrences of field captures of alate or recently
dealate C pennsylvanicus queens (Fig. 1) demonstrates that most nuptial
flights occurred during spring. Surprisingly, founding queens were captured
for more than four months (Fig. 1).
If we compare the data collection on an annual basis (Table 1), we find
that we cannot reject the null hypothesis that there is significant variation
in the numbers of queens captured (Kolmogrov-Smirnov D = 0. 12615, P >
0.05), but we must conclude that there is a significant seasonal, or monthly.
Table 1. The monthly and yearly distributions of collections of alate or recently dealate
queens of C pennsylvanicus given in Fig. 1 .
Month
Year
1977
1978
1979*
1980
1981
Sum
March
0
1
0
1
4
5
April
3
9
0
4
7
23
May
6
18
8
15
6
53
June
5
9
4
8
2
28
July
0
0
0
2
0
2
Sum
14
37
12
29
19
111
All data from student collections.
VOLUME XC, NUMBER 4
249
Table 2. The sex ratios of reproductives from New Jersey field colonies of C. pennsylvanicus,
and a comparison with data given in Pricer ( 1 908).
Colony reference
number
Alates collected
Proportion males
Pricer #1
350
0.4286
Pricer #2
370
0.4708
Pricer #3
323
0.3591
Pricer #4
206
0.4951
Total
1,249
Mean ratio
0.4339
New Jersey colonies collected in spring 1980
Queens present
CP #32
526
0.6026
CP #39
410
0.4805
CP #40
1,009
0.5154
CP #42
1,120
0.6151
CP #45
260
0.4846
Total
3,325
Mean ratio
0.5561
Queens not recovered
CP #37
75
0.7600
CP #38
217
0.4746
CP #41
437
1.0000
Total
729
Mean ratio
0.8189
effect on capture distributions (Kolmogrov-Smirnov D = 0.181982, P<
0.0002).
Field collected colonies of C pennsylvanicus also were variable in sex ratio
(Table 2). The mean ratio of males to females was 1.516, or 60.26% of all
reproductives in the colonies were males. However, if we compare those
colonies from which queens were not recovered with queenright colonies,
we hnd that queenless colonies have a signihcantly higher frequency of males
(0.81893 vs. 0.55188; G = 189.579, P < 0.00001). Also, our colonies col-
lected in New Jersey had a signihcantly higher frequency of males than did
Pricer’s (1908) colonies (0.6026 vs. 0.4339; G = 109.74, P < 0.00001). Mean
dry weight of female alates was 56.8 ± SD 12.3 mg, while that of the males
was 9.3 ± SD 4.3 mg. Assuming equivalence in conversion costs, a female
is thus 6.1 times more costly to produce than a male.
Discussion
Direct observations of synchronous mass mating swarms of Camponotus
spp. have not been recorded. Males produce a mandibular gland pheromone
which stimulates the females to flight (Holldobler and Maschwitz 1965).
However, Veitinghoff-Riesch (1928) and Sanders (1972) have provided indi-
250
NEW YORK ENTOMOLOGICAL SOCIETY
rect evidence that synchronous swarming may occur in C herculeanus (L.).
Typically, alates depart from the colony individually, and ascend until they
are no longer visible, presumably to mate in the air (Eidmann 1929; Sanders
1964).
The data that we have been able to collect does not suggest synchronous
swarming of C pennsylvanicus in New Jersey. Moreover, our data suggest
that nuptial flights occur over a very long period of time, and generally
involve a limited number of queens. Our data do not allow a consideration
of the temporal range of the mating activities of males, but we assume that
it must be similar. The distribution of our capture data does not differ
signihcantly from carpenter ant complaints from the general public in New
Jersey (Fowler and Roberts 1982), suggesting that the general public may
respond to sightings of individual founding queens.
Trivers and Hare (1976) have extended the argument of sex ratio and
incorporated it with the theory of kin-selection to argue that monogynous
ants, such as C. pennsylvanicus, should have a sex ratio controlled by the
workers. If so, the sex ratio should approximate 1 male to 3 females, on a
per weight basis, given the asymmetries of genetic relatedness. A recalcu-
lation of Trivers and Hare’s (1976) data for C pennsylvanicus, based on a
larger sample size for dry weight determinations, gives an inverse ratio of
investment, of the weight ratio of females to males divided by the ratio of
the number of males to females, of 7.92 for Pricer’s (1908) data. For our
data from New Jersey, this ratio is 4.07. If we examine the data from queen-
right and queenless colonies separately, we hnd an inverse ratio of investment
of 5.8 for queenright colonies and 1.35 for queenless colonies. Only when a
queen has died, or is lost, as was probably the case in our queenless colonies,
does this ratio drop, as would be expected, under worker oviposition leading
to increased production of males. As the ratio of investment was highly
variable from colony to colony, it is likely that there is no optimum ratio
of investment, as has been argued by Herbers (1979) and Cannings and Cruz
Drive (1975).
If we consider an elementary model of sex ratio investment (Cannings
and Cruz Drive 1975), in which the more abundant sex is shown to be the
disperser, we may conclude that alate queens do not widely disperse, but
rather males must if outbreeding is to occur. Isozyme studies of localized
populations are needed to test this prediction.
Acknowledgments
We graciously thank the New Jersey Pest Control Association for their
support. New Jersey Agricultural Experiment Station Publication Number
D-081 14-22-82, supported by state funds.
VOLUME XC, NUMBER 4
251
Literature Cited
Blochmann, F. 1885. Ueber die Grundung neuer Nester bei Camponotus ligniperdus Latr.
und anderen einheimischen Ameisen. Zeitschrift fur Wissenschaften Zoologie 41:719-
727.
Cannings, C. and L. M. Cruz Orive. 1975. On the adjustment of the sex ratio and the gregarious
behaviour of animal populations. Journal of Theoretical Biology 55:1 15-136.
Eidmann, H. 1 929. Zur Kenntnis der Biologic der Rossameise {Camponotus herculeanus (L.)).
Zeitschrift fiir Angewandte Entomologie 14:229-253.
Fowler, H. G. and R. B. Roberts. 1982. Activity cycles of carpenter ants {Camponotus)
(Hymenoptera: Formicidae) and subterranean termites {Reticulitermes) Isoptera: Rhi-
notermitidae): inference from synanthropic records.
Herbers, J. M. 1979. The evolution of sex-ratio strategies in hymenopteran societies. American
Naturalist 114:818-834.
Holldobler, B. and U. Maschwitz. 1965. Der Hochzeitsschwarm der Rossameise Camponotus
herculeanus (L.) (Hymenoptera: Formicidae). Zeitschrift fur Vergleisch Physiologic 50:
551-568.
McCook, H. C. 1 883a. How a carpenter ant founds a colony. Annals and Magazine of Natural
History 13:419-423.
. 1 883b. How a carpenter ant founds a colony. Proceedings of the American Academy
of Arts and Sciences of Philadelphia 35:303-307.
Pricer, J. L. 1908. The life history of the carpenter ant. Biological Bulletin 14:177-218.
Sanders, C. J. 1972. Aggregations of alate carpenter ants in Ontario. Proceedings of the
Entomological Society of Ontario 102:13-16.
Trivers, R. L. and H. Hare. 1976. Haplodiploidy and the evolution of social insects. Science
191:249-263.
Vanderschaff, P. 1970. Polymorphism, oviposition by workers and arrested larval develop-
ment in the carpenter ant, Camponotus pennsylvanicus DeGeer (Hymenoptera: For-
micidae). Ph.D. thesis. University of Kansas, Lawrence, Kansas.
Velinghoff-Riesch, A. 1928. Das Verhalten palaarktischer Vogel gegeniiber den wichtigeren
fortschadlichen Insekten (V-IX). Zeitschrift fur Angewandte Entomologie 13:483-512.
Wheeler, W. M. 1910. Ants, Their Structure, Development and Behavior. Columbia Uni-
versity Press, New York, New York.
Zar, J. H. 1974. Biostatistical Analysis. Prentice-Hall, Englewood Cliffs, New Jersey.
Department of Entomology and Economic Zoology, Cook College, New
Jersey Agricultural Experiment Station, Rutgers University, New Brunswick,
New Jersey 08903.
Received for publication April 26, 1982.
NEW YORK ENTOMOLOGICAL SOCIETY
XC(4), 1982, pp. 252-257
THERMAL REQUIREMENTS FOR POSTDIAPAUSE
DEVELOPMENT AND SURVIVAL IN THE GIANT SILKWORM,
HYALOPHORA CECROPIA (LEPIDOPTERA: SATURNIIDAE)
James R. Nechols^ and Paul J. TaubeU
Abstract. — 'Wq investigated the thermal requirements for development and
survival in the giant silkworm, Hyalophora cecropia (L.) (Lepidoptera: Satur-
niidae) at 5 temperatures (L:D 16:8). The developmental times ranged from
an average of 50 days at 15.6°C to 17 days at 26.7°C. Males developed
slightly faster than females at all temperatures except 26.7°C. In females,
the lower thermal threshold, t, and the thermal constant, K, were 10.4°C
and 276 degree days (°d), respectively. In males, the respective t and K values
were 9.2°C and 301 °d. The percentage mortality (pupal to adult) was very
low (0 to 11 %) at all temperatures. The sex ratio was approximately 1:1.
The annual life cycle of a univoltine insect species consists of a period of
activity (reproduction and development), a period of dormancy (including
diapause), and the intervening transitional periods (diapause induction and
postdiapause development). To understand the phenology and population
dynamics of univoltine insects, and to rear these species efficiently in the
laboratory, it is important to understand the primary environmental factors
regulating each phase.
The giant silkworm, Hyalophora cecropia (L.), is the largest and best-
known of the North American saturniids. Much of the research on this
univoltine insect has focused on its dormancy, including physiological and
ecophysiological aspects of its diapause (e.g., Williams 1956, 1969; Mansingh
and Smallman 1966; Sternburg and Waldbauer 1969, 1978; Waldbauer and
Stemburg 1973, 1978; Willis et al. 1974; Waldbauer 1978).
Despite these important studies, a quantitative assessment of the factors
controlling postdiapause development of the cecropia moth has not been
made (see Waldbauer 1978). Because temperature is one of the most impor-
tant abiotic factors influencing development in insects, we investigated the
‘ Present address: College of Agriculture and Life Sciences, University of Guam, Mangilao,
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2 Box 598, Ithaca, New York 14850.
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253
thermal requirements for postdiapause development and survival in H.
cecropia.
Materials and Methods
We obtained ca. 70 viable eggs from a mated female caught in Ithaca,
New York (Tompkins Co.). Following hatching, the larvae had access to a
constant supply of cherry and black willow leaves. Initially, we placed 10
young larvae in each of 7 2-pint jars. As the larvae grew, we separated them
so that there were 1 or 2 larvae per jar. In addition, we reared some of the
large larvae in well-vented 12 X 12 X 6-inch plastic boxes. All of the con-
tainers were cleaned daily.
The eggs and young larvae were kept at room temperature and natural
photoperiod. We maintained the older larvae at 24 ± 1°C, L:D 10:14. After
the cocoons were spun, we held them under a 24 ± 1°C, L:D 10: 14 regimen
for 3V2 to 4 weeks. Because no emergence occurred, we assumed that all
individuals were in diapause.
Subsequently, we transferred the cocoons through a series of decreasing
temperatures: 21, 18.3, 15.6, 10, and 4.4°C (all ±1°C, at L:D 10:14). We
maintained the cocoons for 2 weeks under each condition, except at 4.4°C,
where the cocoons were held for 4 months to allow for the completion of
diapause.
To determine the effect of temperature on postdiapause development, we
divided the cocoons into 6 groups of about 10 each. One group was trans-
ferred to each of the following 5 temperature conditions (15.6, 18.3, 21, 24,
and 26.7°C) under long days (L:D 16:8) (Table 1). The sixth group was
transferred to 24°C and maintained under short days (L:D 10:14); its served
as a control to show that diapause had ended.
We recorded the number of days taken for the moths to emerge in each
condition. The reciprocal of the mean developmental time (1/days) was
plotted against temperature and the resulting curve was tested for linearity
with the ANOVA test (Snedecor and Cochran 1967). The lower theoretical
threshold temperature, t, was obtained by extrapolating the linear regression
of 1/days vs. temperature through the x-axis. We then calculated the thermal
constant, K, by the equation K = y{d — t), where y is the mean developmental
time in days, and d = temperature in °C (Andrewartha and Birch 1954).
Results
Table 1 shows the mean developmental times for each sex of H. cecropia
(pupal to adult stages) at 5 constant temperatures. The number of days to
emergence ranged from 15-20 days at 26.7°C to 38-62 days at 15.6°C. At
all temperatures except 26.7°C, females took slightly longer to emerge than
254
NEW YORK ENTOMOLOGICAL SOCIETY
Fig. 1 . Postdiapause developmental rates for the pupal to adult stages of Hyalophora cecropia
at constant temperatures.
males, but the difference was only significant at 2 1°C (t-test, P < 0.05) (Table
1). The distribution of emergence was continuous for both females and males
at each temperature except 15.6°C, where adults emerged more slowly and
sporadically.
The lower theoretical threshold, t, is 10.4°C for females and 9.2°C for
males (Fig. 1). The thermal constant, K ± SD, is 276 ± 12 and 301 ± 15
degree days (°d) above t for females and males, respectively. Development
varied linearly with temperature between 15.6°C and 26.7°C {P < 0.05).
The overall sex ratio (all temperatures combined) was 57% females and
43% males. This ratio— calculated from a total of 56 including 4 unemerged
animals that were dissected after death— did not differ significantly (chi-
square test, P < 0.05) from a 1:1 ratio.
The percentage mortality of pupae and pharate adults was very low at all
temperatures. However, at 26.7°C, approximately half of the emerged moths
failed to expand their wings fully (Table 1).
VOLUME XC, NUMBER 4
255
Table 1 . Postdiapause developmental rates, mortality, and sex ratio of H. cecropia at various
constant temperatures. Females: t = 10.4°C; K = 276 °d. Males: t = 9.2°C; = 301 °d.
Temp.
(±1°C)
%
mortality
Developmental time (days)
Sex ratio (%)
9 V
3 3
Combined
9
3
26.7
10
17 ± 2.0**
17.8 ± 2.1
17.3 ± 1.9
55
45
(10)’^
(5)
(4)
(9)
24.0
0
19.2 ± 1.3
18.7 ± 0.6
19.0 ± 1.1
62
38
(8)
(5)
(3)
(8)
21.1
0
27.6 ± 1.4
24.7 ± 1.2
26.7 ± 1.9
70
30
(10)
(7)
(3)
(10)
18.3
0
35.2 ± 2.3
33.8 ± 3.1
34.5 ± 2.7
50
50
(10)
(5)
(5)
(10)
15.6
11
52.3 ± 8.4
48.6 ± 7.2
50.0 ± 7.3
44
56
(9)
(3)
(5)
(8)
* (No.) = number tested.
♦♦ Jc ± SD.
Developmental rates of females and males under L:D 10:14 (24°C) were
not significantly different from those under L:D 16:8 (24°C) (t-test, P <
0.05).
Discussion
Development and temperature. —Stemburg and Waldbauer (1969) observed
that H. cecropia males emerge earlier than females under natural conditions.
Our preliminary results (Fig. 1 ) suggest that the earlier emergence of males
can be explained by differences in thermal requirements for the postdiapause
development of the 2 sexes. That is, males begin development at a lower
threshold temperature than females. In addition, males have a slightly faster
rate of development than females at all but the highest temperature tested
(Table 1).
The lower theoretical threshold temperature (t) for development and emer-
gence was ca. 10°C. This is consistent with Williams’ (1956) observation
that some morphological development, but not emergence, occurred in H.
cecropia at 6°C.
Emergence pattern and sex ratio. — Siernburg and Waldbauer (1969) and
Waldbauer and Sternburg (1978) observed a distinct bimodal emergence
pattern in field populations of H. cecropia in Illinois (i.e., emergence in the
2 broods was separated by ca. 1 month). They attributed the bimodality to
variation in diapause termination, not to variation in rates of development
after diapause had ended. In our experiments, emergence did not exhibit
bimodality, but was unimodal at all temperatures. We suggest that the pro-
longed period of chilling experienced by our experimental animals allowed
256
NEW YORK ENTOMOLOGICAL SOCIETY
diapause to terminate in all individuals before transfer to warmer conditions
and thus masked the expression of any variation in diapause termination
that might have been present.
Our sex ratio data and photoperiodic tests are consistent with this hypoth-
esis. Stemburg and Waldbauer ( 1 969) found that 65% of their early-emerging
brood and 43% of their late-emerging brood were females; whereas 57% of
our adults were females (Table 1). This intermediate sex ratio suggests that
if there were two broods in our population, both emerged simultaneously.
Similary, developmental rates were not affected by photoperiod (L:D 16:8
and L:D 10:14), indicating that diapause had terminated in all individuals
before removal from cold conditions. Thus, our results are consistent with
Waldbauer and Stemburg’s conclusion that bimodality in H. cecropia emer-
gence is the result of variation in diapause termination, rather than post-
diapause development.
The unimodal emergence pattern in our experiments may also have resulted
from limited genetic variability. That is, natural H. cecropia populations
from Ithaca, New York, may not have the genetic variability that produces
bimodal termination. It is also possible that our experimental population,
which was derived from limited stock, did not harbor the full range of
variability. Thus, additional studies are needed to establish the relative roles
of environmental and genetic factors in determining emergence patterns in
H. cecropia populations in central New York.
Mortality. — percentage mortality of immatures within coocons was
low ('^4%) and, apparently, unrelated to temperature over the range of
temperatures we used (Table 1). However, at our highest temperature (26.7°C),
almost half of the adults failed to expand their wings fully and died shortly
after emergence. This mortality can be attributed to desiccation during emer-
gence; H. cecropia emerges early in the day (Truman 1972) when daily
temperatures are low and relative humidity is high. Therefore, we conclude
that H. cecropia should be reared at temperatures around 24°C for relatively
fast development and high rates of survival.
Acknowledgments
We thank Maurice J. Tauber and Catherine A. Tauber for their comments
on the manuscript and John J. Obrycki for his cooperation.
Literature Cited
Andrewartha, H. G. and L. C. Birch. 1954. The Distribution and Abundance of Animals.
University of Chicago Press, Chicago. 782 pp.
Mansingh, A. and B. N. Smallman. 1966. Photoperiod control of an ‘obligatory’ pupal dia-
pause. Can. Entomol. 98:613-616.
VOLUME XC, NUMBER 4
257
Snedecor, G. W. and W. G. Cochran. 1967. Statistical Methods, 6th Edition. The Iowa State
University Press, Ames. 593 pp.
Stemburg, J. G. and G. P. Waldbauer. 1969. Bimodal emergence of adult cecropia moths
under natural conditions. Ann. Entomol. Soc. Amer. 62:1422-1429.
and . 1978. Phenological adaptations in diapause termination by cecropia
from different latitudes. Entomol. Exp. Appl. 23:48-54.
Truman, J. W. 1972. Physiology of insect rhythms II. The silkworm brain as the location of
the biological clock controlling eclosion. J. Comp. Physiol. 81:99-1 14.
Waldbauer, G. P. 1978. Phenological adaptation and the polymodal emergence patterns in
insects. Pages 127-144 in: H. Dingle (ed.). Evolution of Insect Migration and Diapause.
Springer- Verlag, New York. 284 pp.
and J. G. Stemburg. 1973. Polymorphic termination of diapause by cecropia: genetic
and geographical aspects. Biol. Bull. 145:627-641.
and . 1978. The bimodal termination of diapause in the laboratory by Hyalo-
phora cecropia. Entomol. Exp. Appl. 23:121-130.
Williams, C. M. 1956. Physiology of insect diapause. X. An endocrine mechanism for the
influence of temperature on the diapausing pupa of the cecropia silkworm. Biol. Bull.
110:201-218.
. 1969. Photoperiodism and the endocrine aspects of insect diapause. Pages 285-300
in: H. W. Woolhouse (ed.). Dormancy and Survival. Academic Press, New York.
Willis, J. H., G. P. Waldbauer, and J. G. Stemburg. 1974. The initiation of development by
the early and late emerging morphs of Hyalophora cecropia. Entomol. Exp. Appl. 17:
219-222.
Department of Entomology, Cornell University, Ithaca, New York 14853.
Received for publication April 26, 1982.
NEW YORK ENTOMOLOGICAL SOCIETY
XC(4), 1982, pp. 258-265
RESOURCE ASSESSMENT BY ADULT AND
LARVAL CODLING MOTHS
Bernard D. Roitberg' and Ronald J. Prokopy
Two-day-old adult and first-instar larval codling moths were
assayed for ability to detect and avoid resource sites (=apples) already occu-
pied by conspecifics (i.e., host discrimination). Under laboratory conditions,
adults oviposited as readily in the presence of conspecific eggs and larvae
as in their absence. Similarly, newly hatched first-instar larvae, when released
in laboratory arenas, located and attempted to mount and bore into apples
regardless of the presence or absence of older conspecific larvae. We contrast
codling moth behavior and ecology with that of two other insect parasites
of apple that are known to host-discriminate, the apple maggot fly and the
European apple sawfly.
Introduction
Accumulating evidence suggests that individuals utilizing restricted
(exhaustible) resources may suffer reduced fitness when living within pop-
ulations below or above some optimal density range (Peters and Barbosa
1977; Prokopy 1981). Thus, selection may favor individuals that are capable
of assessing population density and responding accordingly. Indeed, several
different mechanisms have been demonstrated through which foraging ani-
mals assess resource quality, quantity, and “population load” of conspecifics,
including biochemical, visual, and physical assessment systems (see Prokopy
etal. 1982).
For the past several years we have been investigating resource assessment
behavior of insects that parasitize (sensu Price 1977) apple fruit. Because
individual apples are discrete, exhaustible resource units, we hypothesized
that insects that exploit these hosts may avoid apples already occupied by
a high density of conspecifics. To date, we have demonstrated that two
parasites of apple, the European apple sawfly (Hoplocampa testudinea) (Roit-
berg and Prokopy in MS) and the apple maggot fly (Rhagoletis pomonella)
(Prokopy 1972) are deterred, by an unknown mechanism and marking pher-
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British Columbia, Canada V5A 1S6.
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VOLUME XC, NUMBER 4
259
omone respectively, from ovipositing in developing ovaries and growing
fruit respectively, when such hosts are already parasitized by conspecifics.
Van Lenteren (1976) defines this “refrainment response” as host discrimi-
nation, an ability that has been demonstrated in several entomophagous and
phytophagous parasitic insects (Prokopy et al. 1982). In addition to host
discrimination by the adults, larvae of H. testudinea, when searching for
supplemental fruit, avoid hosts occupied by conspecific larvae (Roitberg and
Prokopy in MS).
Previous research on resource assessment by a third major parasite of
apple, the codling moth, Cydia pomonella, is inconclusive. For example, on
the basis of distribution of codling moth (CM) eggs and larval-infested apples,
Geier (1963) and Jackson (1979) concluded that codling moths probably do
not host-discriminate. By contrast, Maclellan (cited in Wood 1 965) suggested
that freshly laid eggs may deter oviposition by foraging CM females. Van
Lenteren et al. (1978) and Mackay and Singer (1982) demonstrated that,
when used as the sole criterion, egg distribution data is at best weak circum-
stantial evidence on which to base conclusions about host discrimination
ability.
CM females deposit eggs on or near apples. Following hatch, larvae locate,
bore into, and utilize apples for food and shelter. Thus, two very different
life forms must make choices of sites for exploitation. Because Ferro and
Harwood (1973) demonstrated that individual apples support a finite num-
ber of larvae and that competition among larvae may lead to reduced size
and fitness of adults, we were particularly interested in determining whether
either adults or young larvae could detect presence of and avoid competition
with conspecifics.
Materials and Methods
Wild female and male codling moths were collected using an ultraviolet
lamp at Orchard Hill, Amherst, Mass. Moths were placed in plastic (11.5
cm X 8.0 cm) cylinders whose walls were covered with waxed paper. Each
cylinder was provisioned with one small (ca. 3-cm-diam) apple, serving as
an oviposition stimulus (Wearing and Hutchins 1973), and dilute sugar
solution. The resulting eggs were held at 24°C, 40-60% R.H., 16L:8D until
they matured to the black head stage (just prior to hatch). They then received
one of two treatments: (1) transfer to petri dishes for larval behavior assays
(see below), or (2) transfer to ca. 10-cm-diam McIntosh apples. In the latter
case, resulting infested apples were placed in plastic trays that contained
strips of corrugated cardboard which acted as resting and then pupation sites
for mature larvae.
Assay of adult discrimination ability. — ¥o\\ov^ing eclosion, female and
male moths, reared from the 10-cm-diam apples described above, were
260
NEW YORK ENTOMOLOGICAL SOCIETY
placed in waxed-paper-lined cyclinders at a ratio of 1:2, respectively, to
enhance probability of mating and to provide oviposition experience (Roit-
berg and Prokopy 1981). Females that oviposited while within the cylinders
were selected for assays the following day. We performed three different
experiments:
Experiment i: Two, 2-day-old females and three male moths were placed
in a large (60 X 60 X 25 cm) Plexiglas-screen cage 5 hr prior to lights off.
The floor of the cage was covered with roughened. No. 1 grade filter paper
and the front and back walls of the cage were lined with waxed paper to
provide an oviposition substrate. The other walls and ceiling remained as
screen. Using Scotch™ double sticky tape, we afhxed 25 evenly spaced 1 -day-
old CM eggs, collected as described previously, to one of the waxed-paper
walls. To the other waxed-paper wall, we affixed similarly distributed, egg-
free double sticky tape. Moths were left in the cages overnight and numbers
of freshly-laid eggs were counted the following morning. The orientation of
each cage was reversed with each replicate.
Experiment ii: This series of tests was similar to Experiment i except that
no eggs were affixed to either waxed-paper wall. Instead, at the base of one
waxed-paper wall, we placed two, small (ca. 8-cm-diam) McIntosh apples
which were parasitized with two 10-day-old CM larvae. At the base of the
other waxed-paper wall, we placed two unparasitized but similar McIntosh
apples.
Experiment iii: Electrophysiological tests. We employed whole antennal
preparations from 2-day-old CM females. The antennal base was placed on
the indifferent electrode and the end of the flagellum on the recording elec-
trode. Output was recorded on a Hewlett Packard EM tape recorder. We
presented each of the following substrates 3 cm below each antennal prep-
aration:
(1) One unparasitized 10-cm-diam apple.
(2) One 10-cm-old apple parasitized with two 10-day-old CM larvae.
(3) One glass flask, similar in size and shape to the apples and covered
with moist tissue paper (to simulate moisture emanating from apples), to
serve as a control.
(4) One glass microscope slide to which we fastened, with double sticky
tape, 25, 16-hr-old CM eggs.
(5) One glass microscope slide to which we fastened, with double sticky
tape, one 10-day-old CM larva (including some of its frass).
(6) One glass slide to which we fastened double sticky tape, to serve as a
control.
Assays of larval discrimination ability. — ExpmmtnX i: Individual CM eggs,
at the black head stage, were placed in plastic petri dishes (14.5 cm diam)
along with and equidistant from (1) one small (8-cm-diam) unparasitized
McIntosh apple, and (2) a similar apple parasitized by two 10-day-old CM
VOLUME XC, NUMBER 4
261
Table 1. Response of female codling moths to oviposition substrates with or without con-
specifics.
Expt.
Treatment
N
X No. eggs laid/99
p
Substrate harboring 25 con-
i
specific eggs
Substrate harboring no con-
9
10.8 ± 2.1 SE
N.S.
specific eggs
10.7 ± 2.4 SE
Substrate harboring 4 con-
ii
specific larvae/2 apples
Substrate harboring no con-
13
10.3 ± 1.7 SE
N.S. X"
specific larvae/2 apples
11.5 ± 2.5 SE
larvae. Each dish was oriented in a different direction. In addition, each dish
was lined with moist filter paper from which two small circles had been cut.
The apples were placed within the circles so that they did not touch the filter
paper.
After hatch, larvae crawled on the surface of the filter paper and eventually
contacted one of the apples. Due to experimental design, larvae displayed
difficulty mounting apples from the filter paper. Therefore, we terminated
each replicate when a larva attempted to crawl on an apple, recording search
time, and search path of the larva and the parasitization state of the apple.
Larvae were disqualified if (1) they failed to initiate search within 90 min,
or (2) they crawled out of the petri dish.
Experiment ii: Individual eggs, at the black head stage, were placed on
either ( 1 ) a small ( 1 0-cm-diam) unparasitized McIntosh apple, or (2) a similar
apple parasitized by two 10-day-old CM larvae. Larvae were permitted to
search on apples until they either (1) attempted to bore into the apple, (2)
crawled up the stem, or (3) dropped from it. In the latter two cases, larvae
were transferred to apples of the opposite type, and the test was repeated.
In both Experiments i and ii, each apple and larva was used only once.
Results
Females oviposited as readily in the presence of conspecific eggs and larvae
as in their absence (Table 1). In addition, electrophysiological recordings
showed no difference in female antennal response to egg- and larval-treated
glass slides compared with control slides. Also, whereas female antennae
showed a strong and consistent positive response to whole apples versus
none to glass flasks, there were no differences in response to parasitized
versus unparasitized apples.
Results from the larval behavior experiments showed that newly hatched
larvae do not host-discriminate (Table 2). Similar numbers located and
attempted to mount parasitized versus unparasitized apples. In addition.
262
NEW YORK ENTOMOLOGICAL SOCIETY
Table 2. Response of newly hatched codling moth larvae to parasitized and unparasitized
apples.
Host Condition
Treatment Parasitized Unparasitized P
Expt. ii— Larvae released on host
Time to arrival at host
X = 8.5 ± 2.5 SE
X = 9.2 ± 2.7 SE
N.S.
Mann-
(N= 12)
(N= 16)
Whitney U
No. turns >45° while
X = 8.5 ± 2.1 SE
X = 12.2 ± 3.4 SE
N.S.
Mann-
searching for the host
(N= 12)
(N= 16)
Whitney U
Arrivals at host type
12/28
16/28
N.S.
X^
Expt. ii— Larvae released on host
Time until contact with
X = 13.0 ± 4.6 SE
X = 9.3 ± 5.1 SE
N.S.
Mann-
stem base
II
Z
(N= 15)
Whitney U
Time until boring initiated
X= 15.5 ± 4.3 SE
X = 31.6 ± 9.8 SE
N.S.
Mann-
(N = 14)
(N= 15)
Whitney U
Acceptance of host fruit
14/16
15/16
N.S.
X2
there were no statistically significant differences in search speed or turning
rate of larvae that located either parasitized or unparasitized hosts (Table
2). Similarly, when directly placed on apples, newly hatched larvae readily
accepted (i.e., bored into) apples regardless of parasitization state.
Discussion
Resource assessment by most animals is a complex process, shaped in
part by ecological, physiological, and phylogenetic constraints. Thus, differ-
ent animals facing similar foraging problems may, over evolutionary or
contemporary time, employ widely different solutions (cf. Wright’s “adaptive
landscape,” Wright 1931). The present study strongly suggests that the codling
moth, in contrast to at least two other major parasites of apple (R. pomonella
and H. testudinea), does not partition resources through avoidance of occu-
pied resource sites.
Roitberg (1981) analyzed ecological correlates of phytophagous insects
that have been demonstrated to avoid ovipositing at resource sites harboring
high density of conspecifics. Correlates common to most of these species
included: (1) association with host plants that persist over several parasite
generations, (2) comparatively narrow host range, (3) limited mobility of
parents and offspring, and (4) restricted sites of parasitization within indi-
vidual host plants. In addition. Singer and Mandracchia (1982) noted that
most, though not all, host-discriminators lay single eggs. While codling moth
fits the stereotype of a “typical” host-discriminator, two important differ-
ences should be noted. First, adults choose only the proximate area of a host
VOLUME XC, NUMBER 4
263
for subsequent exploitation. It is the larvae that make the final choice as to
which apple will be exploited for food and shelter. Second, larvae are rela-
tively mobile and are not restricted to individual hosts. By contrast, R.
pomonella larvae are unable to emigrate to new hosts should their current
host prove unsuitable (pers. obs.).
Our results provide strong evidence that the codling moth does not dis-
criminate against occupied resources. Still, several alternate hypotheses are
possible. First, adult host-discrimination behavior may have been adversely
affected by experimental conditions, i.e., cage enclosure. We reject this
hypothesis on the basis of evidence from moths enclosed in small (ca. 300-
cc) containers, where eggs were often distributed in small clusters. In contrast,
moths distributed eggs singly in the large assay cages, paralleling the distri-
bution of eggs in nature (Geier 1963). Second, we may have employed such
high concentrations of conspecific eggs or larvae that the host assessment
system was overstimulated, leading moths to oviposit at random. We are
unable to refute or substantiate this hypothesis for codling moth or any other
phytophagous host-discriminator. Third, with regard to CM larvae, the phys-
ical structure of the arena may have disrupted normal host assessment behav-
ior. Our observation of larval behavior argues against this hypothesis, in
that assay larvae moved freely within the assay arenas and appeared to
readily detect the presence of both apples present.
Two features of codling moth behavior, while lacking any overt host-
discrimination, may provide for reduced competition of offspring. First,
adults tend to oviposit in areas of high fruit density (Jackson 1979), thereby
providing greater levels of resource availability. Second, each oviposition is
preceded and followed by a period of flight (Geier 1963). This action reduces
the chance of sib-sib larval competition. Similar behavior has been dem-
onstrated for other insects, including H. testudinea (Roitberg and Prokopy
1980) and R. pomonella (Roitberg et al. 1982).
Finally, we reemphasize the potential danger of unidimensional, deter-
ministic approaches to studies of insect resource assessment behavior and
expectation of universal host-discrimination ability (cf. Gould and Lewontin
1979) . Codling moth adult oviposition and larval host-location behavior is
probably influenced by several factors, all of which may determine future
fitness of individuals. These could include: energetic costs to foraging by
adults and larvae, foraging-associated risks to predation, structure of resource
patches and abiotic factors. As an example, newly hatched larvae may risk
threat to life if resources are not quickly located (Jackson and Harwood
1980) . Therefore, host discrimination may be of lesser importance to these
animals compared to a species such as R. pomonella. The cost of developing
and maintaining a host-discrimination information processing system may
outweigh the benefits (Jackson 1979). In conclusion, while we may analyze
animal behavior from particular theoretical perspectives, we should main-
264
NEW YORK ENTOMOLOGICAL SOCIETY
tain our awareness of the complexity and stochastic nature of behavioral
and ecological events.
Acknowledgments
This research was supported by a grant from the Massachusetts Society
for Promoting Agriculture and Massachusetts Agriculture Experiment Sta-
tion Project 488. We thank R. Crnjar for conducting and analyzing the
electrophysiological experiments and J. Macdougal and D. Eaton for excel-
lent technical assistance.
Literature Cited
Ferro, D. N. and R. F. Harwood. 1973. Intraspecific larval competition by the codling moth,
Laspeyresia pomonella. Environ. Entomol. 2:783-789.
Geier, P. W. 1963. The life history of the codling moth, Cydia pomonella (L.) (Lepidoptera:
Tortricidae), in the Australian Capital Territory. Aust. J. Zool. 1 1:323-367.
Gould, S. J. and R. C. Lewontin. 1979. The spandrels of San Marco and the Panglossian
paradigm: a critique of the adaptationist programme. Proc. Roy. Soc. Lond. B 205:581-
598.
Jackson, D. M. 1979. Codling moth egg distribution on unmanaged apple trees. Ann. Entomol.
Soc. Amer. 72:361-368.
and R. F. Harwood. 1980. Survival potential of first instars of the codling moth in
laboratory experiments. Ann. Entomol. Soc. Amer. 73:160-163.
Mackay, D. A. and M. C. Singer. 1982. The basis of an apparent preference for isolated host
plants by ovipositing Euptychia libye butterflies. Ecol. Entomol. 7:299-303.
Peters, T. M. and P. Barbosa. 1977. Influence of population density on size, fecundity, and
developmental rate of insects in culture. Ann. Rev. Entomol. 22:431-450.
Price, P. W. 1977. General concepts on the evolutionary biology of parasites. Evolution 31:
405-420.
Prokopy , R. J . 1972. Evidence for a pheromone deterring repeated o viposition in apple maggot
flies. Environ. Entomol. 1:326-332.
. 1981. Epideictic pheromones influencing spacing patterns of phytophagous insects.
In: D. A. Nordlund, R. L. Jones and W. J. Lewis (eds.). Semiochemicals: Their Role in
Pest Control. Wiley and Sons, New York.
, B. D. Roitberg and A. Averill. 1982. Chemical mediation of resource partitioning
in insects. In: R. Carde and W. Bell (eds.). Chemical Ecology of Insects. Chapman and
Hall, London.
Roitberg, B. D. 1981. Foraging behaviour of Rhagoletis pomonella, a parasite of hawthorn
{Crataegus). Ph.D. thesis, Univ. Massachusetts.
and R. J. Prokopy. 1980. Oviposition behavior and egg distribution of the European
apple sawfly, Hoplocampa testudinea. J. New York Entomol. Soc. 88:69.
and . 1981. Experience required for pheromone recognition by the apple
maggot fly. Nature 292:540-541.
and . Evidence for epideictic pheromones in larval and adult European apple
sawflies. (in MS)
, J. C. van Lenteren, J. J. M. van Alphen, F. Galis and R. J. Prokopy. 1982. Foraging
behaviour of Rhagoletis pomonella, a parasite of hawthorn {Crataegus), in nature. J.
Anim. Ecol. 51:307-326.
VOLUME XC, NUMBER 4
265
Singer, M. C. and J. Mandracchia. 1982. On the failure of two butterfly species to respond
to the presence of conspecific eggs prior to oviposition. Ecol. Entomol. 7:327-330.
van Lenteren, J. C. 1976. The development of host discrimination and prevention of super-
parasitism in the parasite Pseudeucoila bochei Weld. (Hym., Cynipidae). Neth. J. Zool.
26:1-83.
, K. Bakker and J. J. M. van Alphen. 1978. How to analyze host discrimination. Ecol.
Entomol. 3:71-75.
Wearing, C. H. and R. F. Hutchins. 1973. Alpha famesene, a naturally occurring oviposition
stimulant for the codling moth, Laspeyresia pomonella. J. Ins. Physiol. 19:1251-1256.
Wood, T. G. 1 965. Field observations on flight and oviposition of codling moths (Carpocapsa
pomonella) and mortality of eggs and first-instar larvae in an integrated control orchard.
N. Z. J. Agric. Res. 8:1043-1059.
Wright, S. 1931. Evolution in Mendelian populations. Genetics 16:97-159.
Department of Entomology, University of Massachusetts, Amherst, Mas-
sachusetts 01003.
Received for publication May 3, 1982.
NEW YORK ENTOMOLOGICAL SOCIETY
XC(4), 1982, pp. 266-268
THE LONE STAR TICK, AMBLYOMMA AMERICANUM
(LINNAEUS): NEW DISTRIBUTION RECORD FOR
NORTH AMERICA (ACARINA: IXODIDAE)
John P. McKeon, Thomas F. Bast and Edward M. Bosler
Abstract.— A new distribution record for Amblyomma americanum (L.)
within northeastern New York State is established. Previous reports indicate
this species was limited to extreme southeastern sections of the state. The
finding of three adult ticks in Saratoga County increases the recorded dis-
tribution of this species.
The Lone Star tick, Amblyomma americanum (L.), is known to be well
established throughout the southern section of the United States (Bishopp
and Trembley 1945; Burgdorfer 1969). Previous occurrences of A. ameri-
canum from New York have been limited, reported only from the south-
eastern sections of the state, most notably Suffolk County (Good 1972),
Nassau County (Benach pers. comm. 1 982) and New York City (Rehn 1953;
Benach pers. comm. 1982). One engorged tick was removed from a human
in Catskill, Greene County, N.Y. in 1954 but it was reported that this person
had recently visited the Long Island area previous to the discovery (Jamn-
back 1969). No other reports from upstate New York have occurred during
the previous 50 years (Anastos et al. 1978).
On 8 June 1979, one adult male A. americanum was found preparing to
attach to the upper leg of a human male adult. The tick was encountered
during a walk through an old-field type habitat in the Town of Wilton,
Saratoga County, N.Y. On 1 2 June 1980, one adult female tick was removed
from a human male who had been walking in a field in the Town of Green-
field, Saratoga County. Subsequent field collections failed to recover any
additional ticks. An engorged female tick was removed from a hospital
emergency room female patient on 28 August 1981. The tick was likely
encountered near the patients home in the Town of Moreau, Saratoga County.
The general locations of these three reports are found in Fig. 1 .
These collections indicate that an apparent indigenous, overwintering pop-
ulation of A. americanum now has a limited distribution in Saratoga County.
The medical significance of this new record may be important due to the
Lone Star tick’s capability as a vector of Rocky Mountain spotted fever
The publication costs of this article were defrayed in part by page charge payment. This article
must therefore be hereby marked “Advertisement” in accordance with 18 U.S.C. §1734 solely
to indicate this fact.
VOLUME XC, NUMBER 4
267
SARATOGA COUNTY
NEW YORK
PROVIDENCE ;
A
greenfield
'A
; northum*
.WILTON' bERlXnD
GALWAY
MILTON
SARATOGA
SPRINGS^^
"A
Schuylerville
“iGalway •
Ballstort^/
Spa
L./»
CHARLTON
•MALTA
BALLSTON
I
1-.
I R^und
) Victory
SARATOGA
• ? NLajce
r
! CLIFTON
j
I PARK
STILLWATER
i
/ Stillwater
r*)
V Mechanicville
i
HALF MOON
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/T
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>• Waterford
Fig. 1 . Known distribution of Amblyomma americanum (L.) in Saratoga County, New York.
268
NEW YORK ENTOMOLOGICAL SOCIETY
(Harwood and James 1979) which is known to be endemic in Dermacentor
variabilis (Say) populations near these areas in Saratoga County (McKeon
et al. in preparation for publication).
Acknowledgments
Appreciation is extended to the following individuals for their help in
supplying information; Dr. J. Benach, Dr. C. M. Clifford, Dr. M. L. Margolius
and Mr. B. Fear.
Literature Cited
Anastos, Doss, Furr and Roach, eds. 1978. Index Catalogue of Medical and Veterinary Zool-
ogy, Special Publication No. 3, Ticks and Tick Borne Diseases IV: Geographical Dis-
tribution of Ticks. U.S. Dept, of Agric., Sci. Ed. Admin. 462 pp.
Bishopp, F. C. and H. L. Trembley. 1945. Distribution and hosts of certain North American
ticks. J. Parasit. 3 1(1): 1-54.
Burgdorfer, W. 1969. Ecology of tick vectors of American spotted fever. Bull. Wld. Hlth. Org.
40:375-381.
Good, N. E. 1972. Tick locality and host records from Long Island and southeastern New
York State. Ent. News 83:165-168.
Harwood, R. F. and M. T. James. 1979. Entomology in Human and Animal Health, 7th
Edition. Macmillan Publ. Co., New York. 548 pp.
Jamnback, H. 1969. Bloodsucking flies and other outdoor nuisance arthropods of New York
State. N.Y.S. Ed. Dept. Memoir 19. 90 pp.
Rehn, J. W. H. 1953. The lone star tick in Staten Island, New York (Acarina: Ixodidae). Ent.
News 64(2):46.
(JPM) Bureau of Community Sanitation and Food Protection and (TFB,
EMB) Bureau of Disease Control, New York State Department of Health,
Albany, New York 12237.
Received for publication May 5, 1982.
NEW YORK ENTOMOLOGICAL SOCIETY
XC(4), 1982, pp. 269-274
MORTALITY FACTORS, POLLEN UTILIZATION, AND
SEX RATIO IN MEGACHILE PUGNATA SAY
(HYMENOPTERA: MEGACHILIDAE),
A CANDIDATE FOR COMMERCIAL
SUNFLOWER POLLINATION
V. J. Tepedino and D. R. Frohlich
Abstract. — T)2i\2i on the nesting biology of Megachile pugnata was obtained
from 359 trap-nests at two sites in northern Utah. Bees made an average of
3.8 cells per nest and provisioned cells exclusively with pollen from the
Compositae. Approximately 1 2% of all offspring were killed by either the
clerid predator Trichodes ornatus or the chalcid parasite Melittobia sp. An
additional 30-38% of offspring failed to complete development to the adult
stage. Most of this immature mortality occurred during the overwintering
and incubation stages. The sex ratio of the populations varied between 1.4
to 1.8 ( 6 / 2 ). Nests with with 3 or fewer cells had a lower percentage of
male offspring than all other nests combined. It may be possible to increase
the percentage of female bees by decreasing the depth of the nesting hole.
Introduction
Megachile (Sayapis) pungata Say is a gregariously nesting leafcutting bee
distributed throughout most of the United States and southern Canada (Hurd
1979). The species commonly nests in pre-existing holes in wood and readily
accepts sumac trap-nests (Medler 1964) or artificial wooden domiciles as
nesting sites. The potential importance of M. pugnata as a pollinator in
commercial sunflower fields has recently been recognized (Parker 1981) and
is currently under study (Parker and Frohlich unpubl.).
Relatively little information is available on the biology of this potentially
manageable species. Medler (1964) reported on the nest architecture and
nest associates found in 20 sumac trap nests in Wisconsin. Parker (1981)
and Hurd (1979) noted that M. pugnata visits a variety of flower species,
but Hurd (1979) suggested that composite species are preferred. These rec-
ords are based on flower visitations only; actual pollen usage has not been
examined.
In this paper we report on mortality factors, pollen usage and sex ratio
from 359 nests obtained from artificial domiciles at two sites in northern
The publication costs of this article were defrayed in part by page charge payment. This article
must therefore be hereby marked “Advertisement” in accordance with 18 U.S.C. §1734 solely
to indicate this fact.
270
NEW YORK ENTOMOLOGICAL SOCIETY
Utah. Of especial interest is the relationship between mortality factors, sex
ratio and number of cells per nest. Information on these parameters enable
us to make suggestions regarding the size of nesting materials that might be
offered to populations in commercial sunflower fields so as to minimize
mortality and increase the percentage of females.
Methods
Nests of M. pugnata were obtained from trap-nest blocks placed at two
sites in Cache Co. in northern Utah. The Faust site (42 km south of Logan,
Cache Co., Utah, 1,800 m elevation) is on a hillside with a SW exposure.
It is covered by large stands of mature aspen (Populus tremuloides) which
are surrounded by open, grassland meadows. The Blacksmith Fork site (27
km SE of Logan, 1,500 m elevation) is on a canyon floor surrounded by
maple-box elder with an understory of Hydrophyllum, Cirsium and Bal-
samorrhiza. Trap-blocks were made of pine wood, and contained 50 drilled
holes into which paper soda straws were inserted (7 mm diameter, 15 cm
deep). Blocks were attached to dead trees with holes oriented horizontally.
Blocks were collected during the first week of July and maintained at room
temperature in the laboratory until 1 September when nests were dissected.
During dissection we recorded the contents of each cell, causes of mortality,
and prepared pollen slides (Beattie 1971) from uneaten provisions or from
pollen adhering to the cell walls. Live larvae were put into gelatin capsules
and maintained at 4°C from 30 September until 9 June when they were
incubated at 29°C. Adults were weighed upon emergence and released in a
commercial sunflower field (Parker and Frohlich unpubl.).
Results
We recovered 359 completed and plugged M. pugnata nests from the two
sites (165 Blacksmith Fork, 194 Faust). The sites differed significantly in the
distribution of nests by cells per nest (x^ contingency tables, ^ 1 2.59, df =
6, P = 0.05): Blacksmith Fork nests averaged fewer cells per nest (3.5, SD
1.5) than did Faust nests (4.1, SD 1.6) (Fig. la). Overall, average cells per
nest (3.8, SD 1.6) was similar to Medler’s (1964) findings for holes of the
same depth.
In contrast to Medler’s (1964) report that M. pugnata is univoltine in
Wisconsin, Utah populations had a partial second generation that emerged
during the last half of August. Parker and Frohlich (unpubl.) also found a
partial second generation in Utah populations. The sites differed in the
percent of total offspring that emerged rather than entering diapause: Faust
had significantly fewer emergent bees (3.3%) than did Blacksmith Fork (7.3%)
(x^ = 8.90, P < 0.005). There was rarely more than one emergent individual
per nest.
VOLUME XC, NUMBER 4
271
Fig. 1 . a. Percent frequency distribution of nests by number of cells per nest. b. Percent of
live adults that were males by number of cells per nest. Both sites combined for each figure.
Mortality factors.— yiovXdMXy factors included developmental arrest in all
stages and various predators and parasites (Table 1). Mortality was signifi-
cantly higher at Blacksmith Fork than at Faust (x^ = 8.8, P < 0.005) and
this was primarily due to the larval and sib caused categories. The latter
category requires clarification. Bees that developed directly to the adult stage
without an intervening period of diapause, destroyed any siblings between
themselves and the nest entrance when they emerged. Apparently, egress
could be achieved only at the expense of nest-mates. The higher percentage
of emergent bees at Blacksmith Fork caused a significantly higher incidence
of sib caused mortality at that site than at Faust (x^ = 36.9, P < 0.001).
Most immature mortality occurred during larval and pupal stages: a smaller
proportion of cells contained provisions but no egg, or unhatched eggs (Table
1). Most of the larval and pupal mortality was sustained during the over-
wintering and incubation treatments (Faust, larval 55.1%, pupal 88.3%,
Blaeksmith Fork, larval 64.0%, pupal 77.8%) and may have been due to
maintenance at inappropriate temperatures (see also Medler 1964). The
incubation treatment appeared to be especially detrimental to females; sig-
nificantly more females than males died as pupae or adults during incubation
(Faust, x^ = 12.2, P < 0.001; Blacksmith Fork, x^ = 6.5, P < 0.02).
272
NEW YORK ENTOMOLOGICAL SOCIETY
Table 1. Percent mortality by stages in the life cycle due to developmental arrest or to
parasites and predators (PP). Sib caused mortality was due to emergence of a partial second
generation. Based upon 827 cells (Faust) and 579 cells (Blacksmith Fork).
Provision
Dead egg
Larvae
Pupae
Adult
PP
Sib
caused
2 mor-
tality
Faust
4.1
4.4
8.6
9.3
1.8
12.7
0.7
41.6
Blacksmith Fork
1.9
5.2
14.9
6.2
3.3
11.7
6.4
49.6
Predators and parasites destroyed approximately 1 2% of the cells at each
site. Most of this mortality was caused by clerid beetles (Trichodes ornatus,
47.6% Faust, 71.2% Blacksmith Fork) and the chalcid parasite Melittobia
sp. (39.1% Faust, 18.2% Blacksmith Fork). Small percentages of cells were
destroyed by the chalcid Monodontomerus and the cuckoo bee, Coelioxys
alternata Say. Melittobia and Coelioxys were also recorded by Medler (1964).
We compared the incidence of mortality among nests with different num-
bers of cells to determine if mortality was associated with nest size. Nests
from both sites were combined and then grouped by number of cells. Num-
bers of dead and live individuals were then tabulated for each nest size
category. No association between nest size and mortality was found (x^ =
6.53, df- 5, P > 0.25).
Sex ratio.— Megachile pugnata is a markedly dimorphic bee: using weights
as an estimate of body size gave a 9 / 6 size ratio of 1.82 {x 6 = 62.7 mg,
SD 9.8, N = 65, range = 41.0-84.7; Jc 9 = 114.3 mg, SD 20.3, N - 74,
range = 69.5-165.7). The expected sex ratio for the population was thus,
1.82 5 : 1 2 (Torchio and Tepedino 1980). The sex ratio of live offspring
at Faust (1.76) did not differ from the expected value (x^ = 0.08, P > 0.75)
but the sex ratio at Blacksmith Fork (1.37) was significantly biased towards
females (x^ = 5.55, P < 0.025). These results were unchanged when dead
but sexable individuals were included in the tests.
We examined the occurrence of male and female individuals in nests by
grouping nests with the same number of cells for both sites and then tabu-
lating the number of males and females at each cell position. All nest sizes
(2 to 6 cells/nest; too few 7 and 8 cells available for statistical testing) had
significantly more females in inner cells and more males in outer cells (x^
tests, all P < 0.005 or less). Thus, M. pugnata is similar to the alfalfa leaf-
cutting bee, M. rotundata (Fabr.) (Stephen and Osgood 1965) and other
megachilid bees (Krombein 1967) in the placement of males and females in
the nest.
There was a significant association between the sex ratio and cells per nest
(Fig. lb). Nests with 1-3 cells had lower percentages of males (48.7%) than
did nests with >4 cells (65.5%) (x^ = 17.1, P < 0.001). Thus, as cells are
VOLUME XC, NUMBER 4
273
added to three-celled nests, there is a much greater probability that they will
be males than females.
Pollen utilization.— examined pollen samples from 1,203 cells at both
sites. Few cells contained anything other than composite pollen, although
the presence of more than one species of composite was not uncommon. In
all, only 0.6% of all pollen grains at Faust and 2.5% at Blacksmith Fork were
of non-composite species. It seems clear that the preferred pollen of this
species, at least in northern Utah, is of the Compositae.
Discussion
Megachile pugnata possesses several characteristics in common with the
semi-domesticated pollinator of alfalfa, M. rotundata, which make it an
excellent prospect for the pollination of commercial sunflowers. The species
is widespread in occurrence, accepts man-made nesting materials and pop-
ulations are readily obtainable from appropriate habitats (Parker and Froh-
lich unpubl.). The species can also be manipulated through temperature
treatments to ensure synchronization of emergence with sunflower anthesis.
In addition, Utah populations appear to exclusively collect the pollen of
composites.
Additional work is required to determine the most appropriate temper-
ature at which overwintering bees should be held and subsequently incubated
so as to reduce mortality. This is particularly important because the data
suggest that females, which are more valuable as pollinators than males, are
more susceptible to the temperature treatments employed here. Some of this
mortality may be unavoidable, however, because it is necessary to hold bees
at cold temperatures beyond their “normal” emergence times (June) to syn-
chronize their emergence with sunflower bloom in mid-July.
The depth and diameter of nesting holes influences the number of cells
made per nest and the sex ratio of the alfalfa leafcutting bee, M. rotundata
(Stephen and Osgood 1965; Gerber and Klostermeyer 1972). A similar effect
is likely for M. pugnata. Although we have no data on the influence of
different diameter holes, it is likely that a decrease in the depth of holes
from the 15 cm used here, to 7.5 cm, would result in an increase in the
percentage of female offspring. As Stephen and Osgood (1965) found for M.
rotundata, M. pugnata uses 1 5 cm holes inefficiently: only 3 to 4 cells were
made in most nests and much unutilized space remained. In addition, nests
with 4 or more cells had significantly more males than nests with fewer cells
(Fig. lb). Thus, when space was more completely utilized, it was allocated
to the less valuable (from the viewpoint of pollination) males. It is possible
that a reduction in depth of nesting holes would increase the proportion of
nests with fewer cells and thereby increase the percentage of female offspring.
Whether or not such a procedure would reduce the total number of offspring
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NEW YORK ENTOMOLOGICAL SOCIETY
produced because of increased time spent in plugging shallower nests with
fewer cells, and in searching for new nesting holes (Stephen and Osgood
1965), needs to be studied.
Acknowledgments
We thank P. F. Torchio for donating the nests; K. Ruggeri, G. Trostle,
and J. Higginson for their help with nest dissections and pollen slide prep-
aration and analysis; T. Waldron for help in computerizing the data; and G.
E. Bohart and F. D. Parker for critical comments on the manuscript.
Literature Cited
Beattie, A. J. 1971. A technique for the study of insect-bome pollen. Pan-Pac. Entomol.
47:82.
Gerber, H. S. and E. C. Klostermeyer. 1972. Factors affecting the sex ratio and nesting behavior
of the alfalfa leafcutter bee. Wash. Agric. Exp. Stat. Tech. Bull. 73. 1 1 pp.
Hurd, P. D., Jr. 1979. Superfamily Apoidea. Pages 1741-2209 in: K. V. Krombein, P. D.
Hurd, Jr., D. R. Smith and B. D. Burks (eds.). Catalog of Hymenoptera in America
North of Mexico, Vol. 2. Smithsonian Institution Press, Washington, D.C.
Krombein, K. V. 1967. Trap-nesting wasps and bees: life histories, nests and associates.
Smithsonian Press, Washington, D.C.
Medler, J. T. 1964. A note on Megachile {Sayapis) pugnata Say in trap-nests in Wisconsin
(Hymenoptera: Megachilidae). Can. Entomol. 96:918-921.
Parker, F. D. 1981. Sunflower pollination: abundance, diversity, and seasonality of bees and
their effect on seed yields. J. Apic. Res. 20:49-61.
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. J. Econ. Entomol. 58:965-968.
Torchio, P. F. and V. J. Tepedino. 1980. Sex ratio, body size and seasonality in a solitary
bee, Osmia lignaria propinqua Cresson (Hymenoptera: Megachilidae). Evolution 34:
993-1003.
Bee Biology and Systematics Laboratory, Agricultural Research Service,
USDA, Utah State University, UMC 53, Logan, Utah 84322.
Received for publication May 26, 1982.
NEW YORK ENTOMOLOGICAL SOCIETY
XC(4), 1982, pp. 275-301
DISJUNCT SYNCHRONIC POPULATIONS OF 17-YEAR
PERIODICAL CICADAS: RELICTS OR
EVIDENCE OF POLYPHYLY?
Chris Simon ‘ and Monte Lloyd
Abstract.— T\\q theory and biogeography of brood formation in periodical
cicadas is reviewed in light of the extraordinary number of different “broods”
represented on Long Island and its immediate vicinity. Detailed emergence
records, with dates and exact localities, are given in an appendix. We present
data which suggest that the evolution of Brood I on Long Island may have
been independent of the evolution of Brood I in Virginia. Possible evolu-
tionary scenarios are discussed in light of allozymic and morphometric find-
ings.
Periodical cicadas (Homoptera: Cicadidae: Magicicada) are confined to
the eastern deciduous forest of the United States; they are not found in the
largely coniferous forests of Canada and the northern United States (Marlatt
1907; Dybas and Lloyd 1974). During the most recent (Wisconsin) glacial
advance, ending about 12,000 years ago, most of the periodical cicadas’
present range was covered by spruce forests (Delcourt and Delcourt 1981
and references therein). It is doubtful whether the recent ancestors of Mag-
icicada spp. could have lived in such forests. The species of trees now char-
acteristic of the eastern deciduous forest were confined to small refugia along
river courses in the southern United States. A dry oak/hickory/southern pine
forest covered the southern states outside of the refugia except for peninsular
Florida (sand dune scrub) and the Mississippi basin (cypress/gum).
Today, we find the three morphologically distinct species of periodical
cicadas split up into 12 major dyssynchronous 17 -year broods {M. septen-
decim, M. cassini, M. septendecula) and three 13-year broods {M. tredecim,
M. tredecassini, M. tredecula), each with its characteristic range. These broods,
or year classes, were mapped by Marlatt (1907). (Updated maps can be
found in Lloyd and Dybas 1966; Dybas and Lloyd 1974; and Simon 1979a.)
It is difficult to believe that the broods, as we know them today, could have
existed prior to the Wisconsin glaciation. Periodical cicadas undoubtedly
moved south with the deciduous forest as the ice advanced and must have
' Present address: Department of Zoology, University of Hawaii, Honolulu, Hawaii 96822.
The publication costs of this article were defrayed in part by page charge payment. This article
must therefore be hereby marked “Advertisement” in accordance with 18 U.S.C. §1734 solely
to indicate this fact.
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NEW YORK ENTOMOLOGICAL SOCIETY
migrated back north as the various tree species recolonized (Davis 1976).
It is likely that the past 12,000 years have seen the active proliferation of
new broods from one or a few ancestral broods. The most interesting feature
of periodical cicada evolution is that whatever happened must have involved
all three species because all can now be found coexisting in almost every
brood.
Recently, it has become possible to bring new information to bear on the
questions surrounding periodical cicada evolution, namely electrophoretic
data on allozymes (Simon 1979a, b) and morphometric data on wing vena-
tion (Simon 1982). Thus the alternative evolutionary schemes, which flow
rather easily from biogeography can now graduate from being idle specu-
lation to being testable hypotheses.
We propose to treat this subject in full detail in a series of forthcoming
papers, but our present purpose is more limited and sharply focused. Here,
we will be immediately concerned with only five of the 1 7-year broods: XIV,
X, IX, V, and I. All five of these broods co-occur on Long Island, New York.
They must either have recently immigrated (since Long Island is a terminal
moraine) or very recently have evolved in situ. It is rare to have so many
different broods reported from so small an area. Most of our knowledge
comes from the observations of one man, William T. Davis, who lived on
Staten Island, New York for many years and kept a close watch for periodical
cicadas emerging on Staten Island (Simon 1979c) and on Long Island (see
Appendix I). According to these records. Long Island appears to be either
(1) a microcosm of rampant evolution of cicada broods or (2) an area into
which many broods have recently immigrated, with still more recent extinc-
tions in large mainland areas, leaving behind relict populations of several
broods.
In this paper, we bring together the available evidence from published
records and unpublished correspondence concerning exact localities and
emergence dates for the six periodical cicada broods known for Long Island.
Each of these is compared biogeographically with the nearest known rep-
resentative of the same brood on the mainland (i.e., those populations with
adults emerging in the same year). As we discuss in detail below, all the
Long Island “broods” except XIV, and possibly X, are disjunct, suggesting
either a relict status or a polyphyletic origin, as described earlier (hence the
quotation marks around broods). Further, we have collected adult specimens
of Broods I and XIV from Long Island and from a representative sample
of their geographic ranges and analyzed them phylogenetically using com-
parative wing-morphometric and allozymic data to test the theories of brood
formation.
Although Long Island is an excellent place to study periodical cicadas, we
should acknowledge the possibility that there may, after all, be nothing more
unique about Long Island than the fact that William T. Davis was watching
VOLUME XC, NUMBER 4
277
it. Perhaps equally close observations on other areas would turn up many
more cases of disjunct, relatively small, unrecorded populations which cor-
respond to the emergence years of existing broods.
Theory of Brood Formation
A series of papers (Marlatt 1907; Alexander and Moore 1962; Lloyd and
Dybas 1966; Lloyd and White 1976; White and Lloyd 1975, 1979; Simon
1979a, b, 1982; Simon et al. 1981; Lloyd et al. in prep.) has given us a theory
of brood formation that postulates two kinds of accelerations in the cicadas’
life cycle: (1) those of one year, supposedly brought about by a single his-
torical episode of extraordinary weather, which affects an entire population
within part of the range, and (2) those of four years, supposedly brought on
by nymphal crowding which affects only part of a population and generates
two broods four years out of phase which coexist in the same woodland.
Seventeen-year cicada broods.— \^ we take the centrally-located Brood XIV
as the putative ancestral brood, then the formal scheme shown in Fig. 1 can
be visualized. With both 1-year and 4-year accelerations occurring, it is
possible to derive a given brood in two different ways (or more, if the scheme
is extended). For example. Fig. 1 shows that Brood IX could be derived
from X by a 1-year acceleration, or from XIII by a 4-year acceleration. The
most immediate check we have on the validity of such hypotheses is bio-
geography. Thus the conversion X ^ IX (shown by a solid line in Fig. 1)
was a likely event because Broods IX and X have broadly contiguous ranges
in the Appalachians. Brood XIII, by contrast, is confined to northern Illinois
and eastern Iowa, a less plausible derivation (therefore shown as a dashed
line). In some cases, the biogeography is equivocal, e.g., VI ^ V and IX ^
V both involve broods with closely associated ranges, so each is represented
by a solid line.
Broods III and IV pose a problem: they have nearly contiguous ranges
through east central Kansas, Missouri and Iowa so we can reasonably derive
III from IV, but none of the supposed ancectral broods— V (Ohio), VIII
(Pennsylvania), VII (New York)— have ranges anywhere near those of III
and IV. In this case, we would postulate that Brood IV came from a separate
glacial refugium and III was subsequently derived from it. A less plausible
alternative is that Brood V once existed in the Mid-West (where it was
derived from VI, which is recorded from Wisconsin) but has since become
extinct.
Thirteen-year cicada broods.— lAoyd and Dybas (1966) proposed that
4-year accelerations, by occurring repeatedly, might have been selected for
and the process thereby become genetically assimilated (Waddington 1953,
1956). This scheme has the great advantage that it provides a mechanism
whereby the 13 -year life cycle can be derived from the 17 -year one without
NEW YORK ENTOMOLOGICAL SOCIETY
4-year accelerations — ►
XIV^B-X VI ► 1 1 ►XV — ► (XI)
i I 1 t 1
XIII IX ►V *-l *[xiv]
(xfi) VIII --"►IV -(XVII)
i 1
VII - -►III — -(XVI)
I I
I i
[VI] [„]
Fig. 1 . Formal scheme for deriving all 1 7-year periodical cicada broods from the postulated
ancestor, Brood XIV, by a combination of 4-year and 1 -year accelerations, modified from Lloyd
and Dybas (1966). Solid single arrows indicate that the two broods have contiguous or closely
associated geographic ranges; dashed arrows, that they do not. Triple arrows indicate that the
broods’ ranges are closely associated in three different parts of the country, suggesting the
possibility of polyphyletic origins. Broods in parentheses are of doubtful existence, consisting
of very few records; no arrows lead from them. Broods in square brackets could be derived as
shown, but there is no need to postulate such a derivation, since a simpler way exists to derive
them, starting with Brood XIV.
losing the periodicity in the process (by the generation of intermediates) or
the complete elimination of one life cycle by the other. The problem is that
the biogeography does not seem to fit. The progression XIV ^ X ^ VI
II ^ XV XI does make geological sense, but it ends up with the very
small Brood XI in New England (Brood XI was well known to the Pilgrims
but is now extinct; Lloyd and White 1976) while 13-year cicadas occupy a
very broad range in the southern and Mississippi Valley states.
Biogeography might have us postulate that the 13-year cicadas existed in
northern deciduous forest refugia during Wisconsin times. They may indeed
have evolved from a progression of 4-year accelerations, like we now pos-
tulate for XIV ^ X VI II ^ XV XI, but this may have happened
during a previous interglacial period. This would not be a surprising con-
clusion, since there have been 16 interglacial epochs in the past two million
years (Wright 1976), but we might still need to postulate a separate full-
VOLUME XC, NUMBER 4
279
glacial refuge for the 13-year cicadas. The alternative prediction would have
13-year cicadas evolving in the present interglacial, in situ in part or the
whole of their present range by many 4-year accelerations from one or more
17 -year parental broods. Allozymic data (Simon 1979a) suggest that the
three or four 13 -year broods which are not extinct evolved from a single
1 7-year ancestor. We may never know the history of the other 1 3 -year broods
which were last recorded in the early 1900’s and late 1800’s and were poorly
collected.
The pattern of distribution of 13 -year Brood XIX in relation to 17 -year
Broods III and IV suggests that 13-year cicadas may be advancing into the
range of 17-year cicadas. Lloyd, Kritsky and Simon (in review) propose that
hybridization between 13- and 17 -year cicadas in zones of contact has led
to the exclusion of the 17-year parental brood via nymphal competition
from a second generation 13-year hybrid brood. They present the following
genetic hypothesis: Parental generation— DD = 17 -year and dd = 13 -year;
FI generation— Dd = all 17 -year; F2 generation— DD, Dd = 17 -year and
dd = 13 -year, which could never breed with their 13 -year grandparent’s
descendants, and could only breed with their 1 7-year sib’s descendants every
221 years, but by that time the 17-year brood will have been eliminated
from the area by nymphal competition.
Evidence for four-year accelerations.— ThQ fact that the largest broods of
17 -year cicadas overlap widely and are separated by four years suggested to
Lloyd and Dybas (1966) that they could have been derived from each other
by a 4-year shortening of the life cycle. They suggested that this shortening
could occur through the temporary deletion of a postulated supernumerary
6th nymphal instar. White and Lloyd (1975) excavated 13- and 17-year
cicada nymphs of the same age and found that rather than possessing an
extra instar, the 17-year nymphs differed from the 13-year ones in that the
17-year nymphs grew much more slowly during the first four years of life.
Lloyd and White (1976) postulated that this 4-year inhibition in growth
might be broken by the stimulus of early nymphal crowding, leading part
of the population to emerge four years ahead of schedule. Simon’s (1979a)
phylogenetic analysis of allozymic data for two 17 -year broods and three
13-year broods is compatible with this scheme.
Three years after Lloyd and Dybas (1966) published their hypothesis, a
4-year acceleration was observed in the suburbs of Chicago, where literally
hundreds of thousands of periodical cicadas emerged four years ahead of
schedule. Only one brood of periodical cicadas (Brood XIII) had ever been
recorded from that area, so these cicadas could have had no other origin.
The remainder of the brood emerged, on schedule, in 1973, in enormous
abundance (many millions) with negligibly few appearing during the inter-
vening years. The two species present in Chicago, Magicicada septendecim
and M. cassini, both participated in this event, although the many thousands
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NEW YORK ENTOMOLOGICAL SOCIETY
that emerged in 1969 were not enough to satiate predators, and apparently
left negligibly few descendants (Lloyd and Dybas in prep.). If they had
successfully reproduced, a disjunct population of Brood IX would have been
created, living sympatrically with Brood XIII in Chicago.
Lloyd and White (1976) found Broods X and XIV living sympatrically in
northern Kentucky and documented quantitatively the proportion of cicadas
in each brood by measuring 4-year-old eggnest scars, and those of the current
year (1974) after Brood XIV had emerged. They made the important the-
oretical point that once accelerated individuals establish a “beachhead” that
is large enough to satiate predators, then conversion to the accelerated brood
(Brood X in this case) can be a gradual process extending over many gen-
erations. There is a geographical gradient from mostly Brood XIV in Ten-
nessee to mostly Brood X in Indiana, suggesting that the conversion is taking
place at a faster pace farther north.
An interesting case exists in New Jersey where Brood XV, a very small
brood, is located in counties noted for their abundance of Brood II. A few
thousand individuals of Brood XV emerged in Union, Essex, and Bergen
counties in 1975. This brood was recorded in these localities in the latter
half of the 19th century (Weiss 1916), but no notice had been taken of them
since that time. The cicadas that did emerge disappeared after only a week.
It is unlikely that they left enough progeny to survive 1 7 years, emerge, and
reproduce. Brood XV is probably not (and probably never has been) a self-
reproducing brood, but rather has been regenerated each 17 years from
particularly dense populations of Brood II. Observations of Brood II in 1979
proved that they appeared in abundance in exactly the same suburban yards
where Brood XV had appeared in 1975 (C. Simon unpubl. field notes). In
other words. Brood II may be in a very early stage of the process of converting
to Brood XV, where there is as yet no “beachhead” of XV established.
A similar situation may exist with Brood VI. It is a very wide-ranging but
strongly patchy brood found throughout the eastern United States (Marlatt
1907). Its modern range is poorly documented because of its patchiness.
Brood VI is largely sympatric with Brood X, the next most wide-ranging of
all the periodical cicada broods. If Brood VI was merely repeated 4-year
accelerations of dense populations of X, then the scatter of Vl-populations
would be understandable. However, there are some counties in Georgia and
North Carolina where populations of VI are reported, but none of X (Marlatt
1907). The conversion X VI may therefore be in a stage intermediate
between XIV ^ X and II ^ XV, which is the progression one would expect
if the ancestral brood were XIV (Figs. 1 , 2).
Evidence for one-year accelerations. — examples can be found of
broods that abut each other and are separated by one year (I-II, III-IV,
VIII-IX, IX-X, XXII-XXIII). Broods separated by one year never overlap.
Broods XXII and XXIII are known to occur less than one mile apart near
VOLUME XC, NUMBER 4
281
Fig. 2 . Central Long Island, encompassing all periodical cicada records except possible Brood
II, 1911 localities from Brooklyn (only a few individuals). Locality information is taken from
W. T. Davis’s field notes and publications, from newspapers, and from personal observations.
These sources are detailed in Appendix I. The numbered dots are towns reporting Brood XIV
(dates are given in Appendix I): 1 Hicksville, 2 = Farmingdale, 3 = Massapequa, 4 = Wyan-
danch, 5 = Deer Park, 6 = Half-Hollow Hills, 7 = Dix Hills, 8 = S. Commack, 9 = Ronkon-
koma, 10 = Bohemia, 1 1 = Farmingville, 12 = East Setauket, 13 = Port Jefferson, 14 = Belle
Terre, 15 = Mount Sinai, 16 = Patchogue, 17 = Medford, 18 = Coram, 19 = Miller Place, 20
Sound Beach, 21 = Middle Island, 22 = Rocky Point, 23 = Yaphank, 24 = Ridge, 25 = Brook-
haven National Laboratories, 26 = Shirley, 27 = Mastic, 28 = Manorville, 29 = Center Mor-
iches, 30 = Calverton, 3 1 = Eastport, ? = Riverhead. Crosshatching = Brood XIV from Davis’s
personal records. Inverse cross-hatching (upper left to lower right) = personal observations of
C. Simon in 1974. Broods other than XIV are indicated as follows: light, regular stipple =
Brood X; small, heavy circles or ellipses and hollow “x’s” = Brood V; irregular stipple (near
Lake Panamoka) = Brood IX.
Utica, Mississippi, as do Broods I and II in the George Washington National
Forest near Luray, Virginia (C. Simon unpubl. held notes). Broods IX and
X come close to overlapping, being reported from many of the same counties
in West Virginia (Marlatt 1907), but there are no known cases of their
occurring sympatrically in the same woods. (We predict that none will be
found, for reasons given below.)
The usual geographic pattern is for the accelerated periodical cicada brood
to replace its fellow farther to the north. Alexander and Moore (1962) sug-
gested that, “. . . prolonged or repeated periods of extreme cold caused sum-
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NEW YORK ENTOMOLOGICAL SOCIETY
mation of diapauses . . . and thus caused those populations closer to the
glacial boundary to emerge earlier.” We now understand from more recent
palynological studies (Delcourt and Delcourt 1981) that the deciduous forest
refugia were nowhere near the glacial boundary, but the principle remains
the same: a single historical episode of unusually cold weather could affect
nymphal development.
The physiological machinery by which periodical cicadas seem to “count”
1 7 (or 1 3) years is still a mystery, but it could well be cued to some hormonal
change taking place in their host trees. If unusual weather on some historical
occasion could cause, say, unseasonal defoliation with a second leafing out
later in the summer, the concomitant hormonal change in the tree might
cause the cicada to “count” an extra winter and emerge one year ahead of
schedule (Lloyd and White 1976). The important point is that one would
expect all of the cicadas to be affected by such a change, not just a part of
the population. For this reason, broods which are separated by one year
would not be expected to overlap except by secondary movements which
are known to occur slowly, if at all (Karban 1981; Lloyd, White, and Stanton
1982).
Theoretically, there are three reasons why broods one year out of phase
should never coexist sympatrically in the same woods: (1) The climatic
aberration by which they supposedly arose would be expected to affect the
entire population, as just stated. This reasoning does not apply to 4-year
accelerations if these arise through effects of crowding, which could well
affect only part of a patchily-distributed population. (2) The numerical
response of predators (Murdoch and Oaten 1975) should fall heavily on any
brood lagging one year behind another, probably eliminating the lagging
brood. Escape from predator build-up is theorized to be the whole advantage
of coupling periodicity with the long life cycle. It enables periodical cicadas
to satiate predators on every emergence, and presumably accounts for their
great abundance compared with non-periodical cicada species (Lloyd and
Dybas 1966). Parasitoids such as the tiny wasp Lathromeris cicadae, for
example, prey heavily on periodical cicada eggs during an emergence year
(Marlatt 1907), but must have alternative hosts in other years. After four
years, the numerical response would surely have dissipated itself, so a brood
lagging by four years should not be eliminated by this cause. (3) Newly-
hatched nymphs of the leading brood will settle underground and have the
advantage of being already established by the time young nymphs of the
lagging brood appear. Given that the cicadas are very abundant, the leading
brood may preempt all the suitable feeding sites, leaving nothing for the
lagging brood. This and predator satiation are the two basic assumptions
used by the model of Hoppenstaedt and Keller (1976) to explain the origin
of periodicity. Their model works well, but depends sensitively on the suit-
able choice of parameter values (May 1979).
White and Lloyd (1979) report a case in northern Kentucky where sub-
VOLUME XC, NUMBER 4
283
stantial numbers of cicadas emerged in 1 975, the year a dense emergence
of Brood XIV in 1974. Presumably, this occurred because of malnutrition
associated with extreme crowding— known from previous evidence (White
and Lloyd 1975) to be a real possibility. Based on considerations (2) and (3)
above, they predict that a new “Brood XV” will not become established
sympatrically with Brood XIV.
There is evidence to suggest that considerations (2) and (3) above would
not prevent the coexistence of broods four or more years apart. The same
area in Kentucky has Brood X and XIV living sympatrically. Excavations
of the nymphs (White and Lloyd 1979) show that they are living at different
depths: Brood X below 15 cm, for the most part, and Brood XIV above.
This is reasonable since the larger nymphs probably require larger roots and
the greatest concentration of small roots is near the surface (Rogers 1940).
This means, that 4-year-old nymphs may well have moved down to larger
roots and would then not preempt feeding spaces from another brood lagging
by as much as four years.
Based on these considerations, there is a priori reason to think that the
carrying capacity of the roots of woody plants for cicada nymphs should be
greater if the nymphs belong to two broods four or more years apart than
if all cicadas belong to the same brood. This hypothesis was supported by
the study of Simon, Karban and Lloyd (1981). This argument suggests that
selection should favor 4-year jumps in the life cycle over 1-year jumps
provided that the accelerating individuals are numerous enough to satiate
predators.
Long Island Broods
Fig. 2 summarizes the existing information for Long Island. The patterns
of stippling, cross-hatching, or discrete symbols identify the broods. Arabic
numbers are keyed to place names for Brood XIV only. Locality records,
dates, and sources of information are given in Appendix I. Notice that there
is evidence for the presence of Brood II on Long Island (which is anticipated
from its distribution on the mainland) but it occupied only the extreme
western edge of the island across a narrow straight from Staten Island.
Fig. 3 attempts to place the broods of Long Island into the biogeographical
context of those on the mainland. Ten broods are shown: five that occur on
Long Island (XIV, X, IX, V, I) and five others (XI, VIII, VI, II, XV) that
occur, or formerly occurred, nearby. Two of the broods found on Long Island
(XIV, X) are also found in New Jersey, but the other three (IX, V, I) have
their nearest mainland representatives 350 km or more away, in western
Virginia, West Virginia, and Ohio. These, certainly, must be considered
disjunct distributions.
The relevance of the other broods shown in Fig. 3 (XI, VIII, VI, II, XV)
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NEW YORK ENTOMOLOGICAL SOCIETY
Fig. 3. Biogeography of periodical cicada broods in relation to Long Island, based on county
records from Marlatt (1907). Some of these populations have subsequently become extinct;
recently documented cases of local extinction are plotted as open circles rather than as solid
dots. Left (above): local distributions near Long Island of the six broods on the “main
sequence” of 4-year accelerations postulated by Fig. 1 . All of these broods occur (or did occur)
in the vicinity of Long Island (often with two or three occurring in the same county). The
predominant brood on Staten Island (arrow) is II, but there are well-documented records of
periodical cicadas having emerged at one time or another in synchrony with Broods XIV, X,
VI, and XV (Simon 1979c). Right (facing page): complete distributions for four other broods—
three of these occur on Long Island (I, V, IX) and one on Martha’s Vineyard (VIII), indicated
VOLUME XC, NUMBER 4
285
by the arrrows. These broods, according to Fig. 1 , can all be derived from the “main sequence”
broods by one or two 1 -year accelerations, and in every case the main center of distribution of
the brood is over 500 km from corresponding mainland populations.
can be judged in relation to Fig. 1 . For example, Broods VI and II occur so
nearby on the mainland that they may once have existed on Long Island,
given rise to VI ^ V and II ^ I respectively, then later become locally
extinct. On the other hand. Broods IX and X occur on Long Island (see
Appendix I) so the progression may have been X ^ IX V I. The
question is of unusual interest, since the two kinds of acceleration (1-year
versus 4-year) are postulated to derive from very different causes.
The presence of Brood VIII on Martha’s Vineyard (Fig. 3) is interesting.
In this case, IX ^ VIII is the only reasonable derivation we can offer, since
there is no evidence that a Brood XII ever existed. The records of IX on
Long Island suggest that it may also have occurred on Martha’s Vineyard
at one time. Notice that the two broods on the end of the 4-year acceleration
sequence, XV, and XI, are (or were) located not far from Long Island. Indeed,
it can be said that the 17-year broods in southern New England and Long
Island (with the exception of Brood XIV) are the most derived from the
point of view of Fig. 1 . Furthermore, if we allow the possibility of Brood
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NEW YORK ENTOMOLOGICAL SOCIETY
I- AND 4-YEAR ACCELERATION THEORY
LONG ISLAND
represent broods derived via 4-year accelerations. Broods shown in parentheses in Fig. 1 are
omitted from this figure. Small roman numerals represent broods formed via 1-year acceler-
ations with the exception of the Long Island broods (see text for hypotheses of origin). “A” is
the position of Broods I and II according to a separate Pleistocene refuge theory, while “B”
shows the position of Broods I and II according to Fig. 1 . “C” depicts Brood I plus the Long
Island populations of Brood I (LI-I) as a monophyletic group.
XIV being polyphyletic, it is theoretically possible to derive the northeast-
ernmost XIV populations from Brood XV by a 1 -year acceleration as shown
also in Fig. 1.
Fig. 4 summarizes, phylogenetically, the hypotheses of brood formation
discussed above. Large roman numerals identify “major” broods which are
largest and supposedly derived via 4-year accelerations. Smaller roman
numerals identify broods derived via 1 -year accelerations from larger broods.
Broods I and II are shown in two positions on the tree: position “A” would
result if I and II refuged separately during the Pleistocene (as suggested by
Simon 1979a); position “B” would result if I and II were derived via accel-
erations from Brood VI (as suggested by Lloyd and Dybas, 1966). The Long
Island broods are shown as derived from Brood XIV on Long Island.
Relationships of Brood XIV
Simon (1979a, b) demonstrated that both M. septendecirn (Brood XIV)
and M. tredecim (Brood XXIII) are allozymically homogeneous among pop-
ulations sampled from a broad geographic range, but at the same time,
different from each other and from conspecifics (M. septendecirn Brood XIII,
and M. tredecim Broods XIX and XXII). Data for thirteen populations of
Table 1. Allele Frequencies and Sample Sizes for Brood XIV, Septendecim.
VOLUME XC, NUMBER 4
287
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N. Jet. US 70; KY-A, on KY 70, 1 mi E. Jet. Mammoth Cave; KY-B, on US 127, 4 mi S. Swallowfield.
VOLUME XC, NUMBER 4
289
Brood XIV are given in Table 1 (polymorphic loci only). Twelve of the
enzyme loci studied were monomorphic. Of the seven polymorphic loci,
only one (/3-esterase) showed any among-population heterogeneity. Work-
man and Niswander’s (1970) formula for the heterogeneity chi-square for
allele frequencies was used as a test. As a check, heterogeneity G-statistics
were also calculated (Sokal and Rohlf 1969) and gave identical results. Chi-
square values for each polymorphic locus are given in Table 1.
Although /6-esterase varied among populations, this variation ranged from
only 0.709 to 0.848 (frequency of the most common allele). Note that four
of the 1 3 populations sampled were located on Long Island (Terry ville. Miller
Place, Deer Park, and Mastic; see Fig. 2 for locations). Comparing these four
to the other localities, we see a larger range within a 25 mile radius on Long
Island (0.740-0.848) than is found between North Carolina and Massachu-
setts (0.709-0.753). Furthermore, the Long Island samples were all taken
from very similar pine-oak woodlands; a habitat which differs very little
from that of the Massachusetts samples and which differs a great deal from
the diverse montane forests of the North Carolina samples. In other words,
there appears to be no consistent trend in allele frequency at the /3-esterase
locus with distance, latitude, or habitat type; the small amount of variation
that is present appears to be random. The Long Island populations do not
differ as a group from the other eight populations sampled.
Analysis of periodical cicada morphology seems to show the same picture.
Examination of 4 8 wing- vein characters shows that populations within broods
differ very little in comparison to the differentiation which has occurred
among broods (Simon 1982). Discriminant analysis based on these 48 char-
acters could not distinguish populations, but broods were clearly distinct.
When the results of the discriminant function analysis were graphed, the 95
percent confidence ellipses for all Brood XIV population means overlapped
almost completely. Thus we see a striking degree of morphological and
allozymic homogeneity within Brood XIV and no evidence to suggest poly-
phyly.
Long Island Brood I
As shown in Fig. 3, Brood I occupies a large section of the Shenandoah
Valley of Virginia and near-by valleys in West Virginia. In 1978, we were
interested to find a population of Brood I at Ridge, New York (on Long
Island) which had been described in W. T. Davis’ 1927 correspondence.
Several other Long Island Brood I sites were described in Davis’ publications
and personal notes (see Appendix I) but we were unable to confirm these.
We mapped the Ridge population (Fig. 5) and also documented the presence
of Brood XIV whose eggnests occurred in the exact same trees (Simon et al.
1981). We surveyed the Brood I emergence area by driving and listening
290
NEW YORK ENTOMOLOGICAL SOCIETY
Fig. 5. Map showing the location of the study area near Ridge, Long Island. The William
Floyd Parkway runs along the western edge of the map, Middle Country Road (New York
Route 25) angles east-northeastward near the southern edge, and a power line runs roughly
north-south just to the west of Lake Panamoka. Single lines are sandy fire lanes; double lines,
paved. Areas marked “developed” are in housing; the remainder is sandy oak scrub. The oak
VOLUME XC, NUMBER 4
291
along the sand fire-lanes. Heavy chorusing centers of Brood I are shown in
Fig. 5 by ellipses of heavier stippling among the light stipple. Broods I and
XIV are the only two Long Island broods whose populations have been
documented to occur in exactly the same trees. Our study site at Ridge is
the only case where reproductive success of Long Island Brood I in successive
generations has been verified. Notice from Fig. 2 that Brood XIV occupies
a very large portion of Long Island.
The fact that Brood XIV is much more abundant in the area than is Brood
I would suggest that I is derived from XIV. However, deriving I directly
from XIV would be contrary to the theory (Fig. 1) which makes no provision
for 4-year decelerations.
Given the evidence that Brood XIV on Long Island is not independently
derived, the theory requires that we derive I from XIV by one of three
pathways: XIV ^ X ^ VI - II ^ I, or XIV - X VI -> V ^ I, or XIV
^X^IX^V^I (see Fig. 1). The third hypothesis is appealing in that
there are now populations of Brood V, five or six kilometers northeast of
our study site across Lake Panamoka (Fig. 2) and populations of Brood IX
are known to have emerged very near there in 1918 (Appendix I; Fig. 2).
Furthermore, there are a substantial number of Brood X populations on
Long Island, one of which is directly south of our study site (Fig. 2). Brood
VI has never been reported on Long Island and Brood II is only known from
the westernmost corner of the island. In other words, the third hypothesis
for the origin of Brood I is strengthened not only by the presence of the
intermediary links but also by the complete absence of any other broods.
Allozymic and morphometric data.— The collection of adults from the
Long Island Brood I population (LI-I) allowed us, for the first time, to
examine a disjunct population, in this case separated by 275 km from the
nearest recorded population of this brood and 475 km from the nearest
extant population (Fig. 3). If this disjunct population was morphologically
and/or allozymically distinct, we hypothesized that a phylogeny based on
these data would place LI-I as the sister group of XIV rather than as the
sister group of I.
A comparison of wing morphometric data (characters same as those illus-
scrub is almost a pure stand of Quercus ilicifolia, which bums at erratic intervals and is seldom
more than 1.5 m tall. Interspersed are occasional pitch pines (Pinus rigida) and white oaks
{Quercus alba), 3-4 m high. Blueberries ( Vaccinium sp.) and huckleberries {Gaylussacia sp.)
are also found, as well as sweet fem {Comptonia peregrina), bracken fern {Pteridium aquilinum),
and winged sumac {Rhus copallind). Judging from eggnest scars, Brood XIV emerged throughout
the oak semb area in 1974. Light stippling indicates the area where Brood I also appeared in
1978. The heavier dots within the light stippling represent the loudest chorusing centers of
Brood I. Small squares in the lower left are quadrats sampled by Simon et al. (1981).
292
NEW YORK ENTOMOLOGICAL SOCIETY
Fig. 6. Three-dimensional plot of brood mean scores of wing measurements on three dis-
criminant function axes with 95 percent confidence ellipses about the means. Sample sizes are
given in the text. If we number the octants as shown in the inset cube, then Broods I (ellipses
1-5) and II (ellipses 6-10) fall within octants 2, 3, 5, 6, and 7; LI-XIV (ellipses 13 and 14) falls
within octants 1 and 4; and LI-I (ellipses 1 1 and 12) falls within octant 8. LI-XIV ellipses are
larger due to smaller sample size per population.
trated in Simon 1982) for Broods I, II, XIV, and LI-I via discriminant
function analysis is shown in Fig. 6. Sample size per population/number of
populations were: 20/5, 20/5, 10/2, 20/2, respectively. Note that Broods I
and II are not recognizable as two distinct groups; their populations inter-
mingle even when plotted in three dimensions. Populations of LI-XIV and
LI-I, on the other hand, are quite different from populations of Broods I
and II and from each other (Fig. 6).
A phylogenetic tree was constructed using the wing morphometric data.
Forty-eight wing vein characters were measured for four 17-year broods
(sample size/number of populations): I (100/5), II (100/5), XIV (80/8), XIII
(100/2), and LI-I (40/2); and one 13-year brood: XXIII (80/8). Five popu-
lations of M. tredecassini (90 individuals/5 populations) were measured to
serve as an outgroup to root the tree. Many of these measurement characters
were not significantly different {P > 0.05) among the broods examined. To
VOLUME XC, NUMBER 4
293
CASS LM XIII XIV I II XXIII
SS- STP
CODING
I I 1.5 2.5 3 3.5 4
Fig. 7. An example of the coding procedure used for each of the 48 wing morphometric
characters. The upper half of the figure depicts the bar diagram resulting from a sum of squares
simultaneous test procedure (SS-STP) for a single character. Bars unite statistically homogeneous
groups. The lower half shows the coding of the bars for that character. The coded information
was used to create a phylogenetic tree using the Wagner procedure.
remove differences among populations which were not statistically signih-
cant, all broods which did not differ for a particular character were pooled.
This pooling was accomplished by performing a posteriori grouping tests
(sums of squares simultaneous test procedure, Sokal and Rohlf 1969) for
each character and then coding the resulting bar diagrams (Simon 1983).
CASS LM XIII XIV I II XXIII
Fig. 8. Wagner phylogenetic tree based on wing vein characters. The shortest tree is rep-
resented by solid lines. Dashed lines indicate a phylogenetic hypothesis which is only 1 8 percent
longer.
294
NEW YORK ENTOMOLOGICAL SOCIETY
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VOLUME XC, NUMBER 4
295
An example of the coding procedure is given in Fig. 7. This procedure retains
more information than previously proposed coding procedures (Mickevich
and Johnson 1976). The coded data was used to construct a phylogenetic
tree using the Wagner-78 program written by J. S. Farris (see Farris 1970).
The resulting tree is shown in Fig. 8.
The phylogenetic tree based on wing morphometric data places the LI-I
populations in the most ancestral part of the tree. Because it is so unlikely
that the small population of Brood I on Long Island could be a remnant of
the ancestor of Brood XIV, which covers much of the eastern United States,
alternative phylogenetic hypotheses were examined. Alternative trees which
placed LI-I as the sister group of I and II or XIV were examined but proved
to be less compatible with the data (i.e., produced much longer trees). It
was, however, possible to create a tree in which Broods XIV, XIII, and LI-
I all originated at approximately the same time. This tree (shown by dotted
lines in Fig. 8) was only 1 8 percent longer than the shortest tree. No signif-
icance tests exist for comparison of phylogenetic trees.
The morphometric phylogenetic hypothesis generated above could be
explained by the following evolutionary scenario. Brood XIV was the ances-
tral brood and at one time covered most of the eastern United States. Very
early in its history it gave rise to Broods XIII and LI-I. Broods I and II were
later derivatives followed by the conversion of southern populations into
13-year Brood XXIII. This scenario is compatible with Lloyd and Dybas’
(1966) scheme shown in Fig. 1.
Allozymic analyses of Broods I, II, LI-I, and XIV were less informative.
Table 2 presents the results. Heterogeneity chi-square values indicate that
essentially no differentiation has occurred. The enzymes a-glycerolphosphate
dehydrogenase (a-GPD), phosphoglucomutase (PGM), and nothing dehy-
drogenase (NDHt) are homogeneous in allele frequency across all broods.
Esterase (/3-esterase) shows significant differentiation but Broods I, II, and
LI-I taken as a group are not significantly heterogeneous and LI-I and XIV
taken as a group are not significantly heterogeneous. Mannosephosphate
isomerase (PHI) is significantly heterogeneous across all broods but I, II,
and LI-I and XIV and II are not significantly heterogeneous. Phosphoglucose
isomerase (PGI) is significantly heterogeneous across all broods but I, II and
XIV are not significantly different as a group. In other words there is little
allozymic information on which to base a phylogenetic tree.
Clearly, additional data are needed. Populations of LI-V will be collected
in 1982 in addition to Brood V from Ohio, West Virginia and Virginia.
These cicadas will be analyzed both allozymically and morphometrically.
We can say, however, that LI-I is morphologically different from Broods I,
II and XIV and that a phylogenetic analysis suggests that LI-I evolved
independently of Brood I in Virginia.
296
NEW YORK ENTOMOLOGICAL SOCIETY
Acknowledgments
Tom Kolwalsic, Irving Hulse, Lewins Orchard, and especially Joseph F.
Burke provided historical information on cicada distributions. We are also
grateful to JoAnn White, Richard Karban and James Archie for comments
on the manuscript. JoAnn White, Peter Curtis, Karen Dohrman, Robin Cort,
Richard Karban and James Archie helped with the held work; without their
help this work would have been very difficult. This research was supported
by a grant from the National Science Foundation, DEB 78-107 10 to Monte
Lloyd. This is contribution number 434 from the Department of Ecology
and Evolution, State University of New York, Stony Brook.
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Sokal, R. R. and F. J. Rohlf. 1969. Biometry. W. H. Freeman & Co. San Francisco, Ca.
776 pp.
Waddington, C. H. 1953. The genetic assimilation of an acquired character. Evolution 7:118-
126.
. 1956. Genetic assimilation of the bithorax phenotype. Evolution 10:1-13.
Weiss, H. B. 1916. The distribution of the periodical cicada in New Jersey. Entomol. News
and Proc. of the Entomol. Section of the Acad, of Natur. Sci. Philadelphia 27:337-340.
White, J. and M. Lloyd. 1975. Growth rates of 17- and 13-year periodical cicadas. Amer.
Midi. Natur. 94:127-143.
, , and Stanton. 1982. Dispersal of fungus-infected periodical cicadas to new
habitat. Environmental Entomology 1 1:852-858.
and . 1979. 17-year cicadas emerging after 18 years: a new brood? Evolu-
tion 33:1193-1199.
Workman, P. L. and J. D. Niswander. 1970. Population studies on southeastern Indian tribes.
II. Local genetic differentiation in the Papago. Am. J. Human Genet. 22:24-49.
Wright, H. E., Jr. 1976. The dynamic nature of Holocene vegetation. Quat. Res. 6:581-596.
Appendix I
This appendix summarizes Long Island periodical cicada locality records.
This compilation is useful as many of the records are not conveniently
accessible being contained in newspapers, private notes, and old, not widely
circulated journals. Localities are organized by brood in chronological order.
Brood /. — 1910: . . in the Half Way Hollows near Wyandanch, Long
Island” at the Holmes farm, “they had been very numerous .... The species
occurred in great numbers in the same territory [same trees] in 1906 . . . .
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NEW YORK ENTOMOLOGICAL SOCIETY
In the Long Island locality [in 1910] they were certainly numerous enough
to lay a great many eggs, and may possibly establish a new brood” (Davis
1910). “It was ascertained, however, that the cicadas had occurred in great
numbers in still another locality in 1910, in a place further to the north [of
the Holmes farm] and nearer to the Dix Hills” (Davis 1911). 1927: In a
letter to W. T. Davis from Albert A. Arnold dated July 1 1, 1927, Arnold
reports Magicicada at two localities: 1) Along Middle Island Road (from
Coram) near Camp Upton (now Brookhaven National Laboratory) “cov-
ering all the scrub oaks”; 2) On the road leading from Coram to Riverhead
on the flat portion of land just east of “the road which formerly led into
Camp Upton property and extending for quite some distance on both sides
of this road.” A reference in W. T. Davis’ personal notes explains that
Magicicada collected from Camp Upton on June 15, 1927 were “found in
abundance.” 1944: In a letter to W. T. Davis dated June 22, 1944, Roy
Latham listed the following periodical cicada localities: 1) Dix Hills in between
Commack Road and Deer Park Avenue south of 25; 2) on 25 several miles
east of Ridge, on the north side of the road, “a small colony”; 3) on 25 near
the junction of 25A (east of Ridge) and for a mile farther. “Just east of
there” he reports “a large colony on both sides of the road— greatest abun-
dance on the north”; 4) Along old country road, north of Riverhead and on
25. Scattered individuals were seen “up to two miles east of Riverhead.”
Brood //. — 1911: “The 17-year cicada occurs on Long Island, New York,
in numbers at the most unexpected times .... Brood 2 [1911] was in its
usual abundance on Staten Island and in the valley of the Hudson, but on
Long Island very few of the cicadas were found .... But one or two indi-
viduals were reported from the western end of the island during 1911” (Davis
1915). 1945: We have found no reference to Brood II in 1945 other than
this strange article from Newsday, May 30, 1974 — “Everyone remembers
Brood II, which covered Long Island in 1945 and left newspaper editors,
pesticide manufacturers and other disaster-lovers counting the years until it
was due again in 1962. Only, Brood II did not keep its appointment.” 1962:
On June 14, 1962, The New York Times published a small article entitled,
“Scientists Mystified as 17-year Locust Shuns Two Counties.” In this article
they commented on the extreme abundance of cicadas in the Hudson Valley,
metropolitan New Jersey and Staten Island and added, “but in Westchester
and Nassau counties, the schedule notwithstanding, the shrill cry of the locust
has not been heard.”
Brood K — 1914: Davis (1915) wrote, “Returning to Wading River on July
24 [1914], we noticed from the car window, about a mile to the west of the
railroad station, many oaks and other trees on the north side of the track in
which the cicadas had laid their eggs, causing the death of the smaller branches
... we found that the seventeen-year cicada had been very common about
Deep Pond and on the easterly side of Long Pond [now Lake Panamoka]
VOLUME XC, NUMBER 4
299
. . . . Under the date of June 9, 1914, Dr. Frank Overton . . . wrote me that
the seventeen-year cicadas were spread over several square miles northwest
of Calverton, about two miles from the Deep Pond locality mentioned above.
He said that he had taken particular notice of them for three miles along
the Riverhead, Coram road about one or two miles north of the station.
They reached all the way to the elevated lots and even scattering ones were
found nearly into Riverhead.” Davis (1924) presumably reasoning from
1914 records predicted that, Brood V would appear “in 1931 in the Half
Way Hollow Hills, also in the hills north of Holbrook and at Wading River.”
1965: On June 6, 1965 The New York noted that, “. . . 17 -year locusts
were reported this week in the Lake Panamoka area of Wading River and
a vast peach orchard off North Road in Calverton.” Newsday also carried
the story: “. . . the insects have appeared this year on Long Island in Cal-
verton and around Lake Panamoka near the Riverhead-Brookhaven town
line.” Brewster (1965) reported that “. . . May 23rd the 17-year locust (peri-
odical cicada) emerged in wooded areas at Lake Panamoka and several days
later at Calverton.” A population of cicadas was noted at the Transient
Camp Area of Wildwood State Park, Wading River and also across from
the picnic and parking areas (J. R. Wildt pers. comm.). They were also seen
along River Road in Calverton, in Le win’s Peach Orchard adjoining Wild-
wood State Park (R. H. Brewster pers. comm.), and on 25A approximately
one mile southeast of the junction of Sound Avenue (John Waskewicz pers.
comm.) in that same year (1965). In 1982 they were again seen in Wildwood
State Park (Camping area E and adjoining Lewin’s Orchard). They were very
abundant east of the junction of 25 A and 25. Oviposition was relatively
heavy along the roadside but no flagging was observed.
Brood /3f. — 1918: Davis (1920) found periodical cicadas “in the woods
along the road between Manorville and Wading River.” He collected some
“about one mile north of the Middle Country Road.” These were the only
ones observed in 1918 and none were there in 1919.
Brood X — 1902: Davis (1920) quoted the eighteenth report of the N.Y.
State entomologist (1902, p. 113) as follows, “The insects were observed
. . . at Wantagh, Nassau Co., also between Massapequa and Amity ville,
between Sayville and Oakdale, east of Patchogue to Brookhaven and also
to the north of Medford and Holtsville, and a small brood [sic] northeast of
Riverhead, all in Suffolk Co.” Davis (1907) reported that although a friend
had seen hundreds of exuviae of the 17 -year locust in Prospect Park, Brook-
lyn, he had only obtained three adults and he “attributed their scarcity to
the English Sparrow.” 1919: The New York Times of June 17th 1919 (p. 25:
3) talked with farmers in the vicinity of Farmingdale, Bethpage, and Mas-
sapequa who reported thousands of cicadas doing damage to fruit trees and
other hardwoods. Old residents claimed that 1 7 years before they were not
nearly so numerous. Davis (1919) recorded “ 1 7-year cicadas singing at Mas-
300
NEW YORK ENTOMOLOGICAL SOCIETY
tic, L.I. during the first week of June.” In another publieation (Davis 1920)
he noted them as occurring on South Country Road just east of Carman’s
River, and in the woods just east of Patchogue; also from Wantagh to Far-
mingdale and as far north as Central Park on Long Island; finally, north and
east of Massapequa rail road station. 1936: The New York Times (June 12,
1936, p. 4:7) reported that the cicadas were found “hrst in Carmen Ave.,
Farmingdale . . . since then the swarms have been reported at Massapequa,
and all through Suffolk scrub oak along the Motor Parkway from Medford
westward to Farmingdale.” They were also seen along the Sunrise Highway
in Massapequa Park. 1970: Newsday (June 5, 1970, p. 12) lists two exact
localities Skylark Drive (Holtsville) and Springdale Drive (Ronkonkoma).
They explained that “officials of the State Conservation Department and
County Agricultural Extension Service said . . . that they have reeeived
hundreds of complaints this month about the insects. Most of the calls have
eome from an area including Ronkonkoma, Holtsville, Islip, and Sayville,
where the influx is concentrated.” The same newspaper (June 23, 1970)
reported 17-year cicadas in Bohemia on eighth Street near the South Side
Sportsman’s preserve. They must have been abundant beeause, “50 Bohemia
residents . . . signed petitions appealing for help to fight the alarming problem
of swarming cicada locusts [sic].”
Brood A/K — 1906: From personal observation and from other sourees,
Davis (1923) discovered that the 1906 Brood XIV covered “in a general
way territory along the north shore from Oyster Bay to Wading River,
extending south to Farmingdale and eastward through the eentral portion
of the island to Manorville. There were also colonies at Moriehes and East-
port.” Davis (1 907) additionally noted that cicadas were eollected in Yaphank
in 1906 from the same loeality in whieh they were exceedingly abundant in
1855. 1923: Davis (1924) reported that “during the summer [of 1923] I
received . . . considerable information regarding the distribution of [Brood
XIV] .... The area already given for the appearance of the brood in 1 906
was almost exactly duplicated.” In the same paper, Davis mentions finding
two species (M. septendecim and M. septendecula from the descriptions) at
the Holmes farm near Wyandaneh. “The large form extended southward to
the railroad and beyond.” This loeality is important beeause Davis reported
finding Brood I in the same apple trees in 1910 and 1944. The Brooklyn
Eagle (June 5 and 6, 1923) noted that cicadas were heard in Yaphank and
Camp Upton on the previous Saturday. Among Davis’ personal notes and
letters, we found the following exact locality descriptions: 1) May 30, 1923
F. M. Schott saw many cicada openings in the ground and later heard them
in Pinelawn and Wyandaneh; 2) Arthur H. Helme saw cicadas “in great
numbers 1 mile east of Port Jefferson between Port Jefferson and Crystal
Brook at the place known as Old Place (letter of June 6). On June 14 (in
VOLUME XC, NUMBER 4
301
litt.) they were still emerging at Belle Terre and there was a colony at Middle
Island. Helme also noted great numbers of these insects from a point south-
east of Port Jefferson Station all along the route from there to Rocky Point,
along the wooded stretches south of the new state road, and at some spots
extending north of the railroad; 3) Edward Bell (letter of June 18, 1923) said
that the cicadas were continuous along the Motor Parkway from Farmingdale
to the road running from Smithtown to Islip; 4) J. Nichols (letter of June
20, 1923) reported seeing the cicadas in the woods boarding the Mastic
railroad station; 5) Roy Lantham (letter of August 24, 1923) searched the
island from Orient to Calverton and never found them east of Calverton;
6) Finally, E. S. Miller (letter of September 12, 1923) summarized, “they
seemed to work in a strip across the island from Rocky Point to South
Setauket in width and thence across the island through Ridge, Middle Island
and Coram, thence across to the south side— Moriches to Brookhaven (worst
from Ridge to Coram). 1957: The New York Times (May 28, 1957) notes
“the heaviest infestation is reported in the Huntington-Deerpark area. None
has been reported in Nassau County.” 1974: Newsday (May 30, 1974) listed
two street addresses (52 Livingston St., Deer Park; 1 7 1 Eastwood Ave., Deer
Park) and said that county agents received approximately 350 calls from
“an area ranging from Hicksville in the west to Mastic and Port Jefferson
in the east, with a concentration from the Medford and Dix Hills-Deer Park
area.” Newsday (May 31, 1974) reported cicadas at 19 Deer Lane, East
Setauket. The following list of localities was compiled in 1974 (1 = personal
observation, 2 == record of county agent, 3 = record of exterminators): Baby-
lon (3), Belle Terre (1), Bohemia (2, 3), Brentwood (3), Brookhaven Labs
(1, 2), Calverton (3), Center Moriches (2), Coram (1, 2), South Commack
(1, 3), Deer Park (1,2), Dix Hills (1,2), East Setauket (1,2, 3), Farmingville
(2) , Hicksville (2, 3), Manorville (2), Massapequa (3), Mastic (1,2), Medford
(3) , Middle Island (3), Miller Place (1), Patchogue (2, 3), Port Jefferson Station
(1,2), Ridge (1,3), Riverhead (2), Rocky Point (3), Ronkonkoma (2), Shirley
(3), Sound Beach (2), and Terryville (1, 2).
(CS) Department of Ecology and Evolution, State University of New York,
Stony Brook, New York 1 1794 and Department of Biology, University of
Chicago and (ML) Department of Biology, University of Chicago, Chicago,
Illinois 60637.
Received for publication May 26, 1982.
NEW YORK ENTOMOLOGICAL SOCIETY
XC(4), 1982, pp. 302-305
BOOK REVIEWS
The Pentatomoidea (Hemiptera) of Northeastern North America with
Emphasis on the Fauna of Illinois. J. E. McPherson. Southern Illinois
University Press. 1982. 240 pp. $30.00.
The field of hemipterology, like many others in entomology, suffers not
so much from a lack of research, as from a lack of assimilation. The present
book provides a compendium of biological information on our native stink-
bugs and their allies. In all the book treats 1 20 pentatomoid species, approx-
imately one-third of the Nearctic fauna. The geographical coverage is the
northeastern quadrant of North America; those states and provinces to the
north and east of Missouri.
The book is very much in the tradition of Blatchley’s “Heteroptera of
Eastern North America” and Hart’s “Pentatomoidea of Illinois.” In the years
since the publication of these earlier works virtually every tribe and large
genus in the superfamily has seen some revision. McPherson’s book updates
the nomenclature and provides keys with illustrations for the identification
of species.
The books greatest value, however, is in the presentation of detailed eco-
logical data, gleaned from an extensive but scattered literature on the subject.
This information, including food plants (or prey), behavior, seasonality,
distribution and parasites, is presented with ample documentation in an
objective and scholarly manner. Host plants are not simply cited, but obser-
vations on abundance, whether or not the insect was actually seen feeding,
and if immature stages were present, are also included. This attention to
detail precludes the readers often time-consuming task of tracking down the
original source. Nevertheless, a useful and voluminous bibliography has been
appended.
There are some problems with the key to the genera of Pentatomini, but
these problems will remain until generic distinctions have been clarified by
future taxonomic work. The keys do work better than those presently avail-
able. In this reviewer’s opinion at least some of the space (22 pages) devoted
to Illinois county records could have been put to better use with figures.
There is a complete lack of habitus drawings, and the few body outlines
provided (e.g., Rhytidilomia senilis) only vaguely resemble the insects
intended.
This book is obviously the result of painstaking research, and these minor
problems do not detract from the main value of the book; an informative
and up-to-date reference on the biology of the pentatomoid Hemiptera.
Donald B. Thomas Jr., Lincoln, Nebraska.
VOLUME XC, NUMBER 4
303
Literature Cited
Blatchley, W. S. 1926. Heteroptera or True Bugs of Eastern North America with Especial
Reference to the Faunas of Indiana and Florida. Nature Publ. Co., Indianapolis.
1116 pp.
Hart, C. A. 1919. The Pentatomoidea of Illinois with keys to the Nearctic genera. Illinois
Natur. Hist. Surv. Bull. 13:157-223.
The American Cockroach. William J. Bell and K. G. Adiyodi, eds. Chapman
and Hall, London-New York, and Methuen, New York. 1981. 529 pp.
$65.00.
The Laboratory Cockroach. W. J. Bell. Chapman and Hall-Methuen. 1982.
161 pp. $13.95.
There exist 47 species of cockroaches (Periplaneta) and four occur in the
United States, but to non-experts it will come as a surprise that none of
these are endemic to America. The name of the cosmopolitan American
cockroach, P. americana, is misleading, as this insect is of African origin.
The importance of cockroaches cannot be overemphasized to entomologists.
This large volume, edited by Bell and Adiyodi, provides an integrated account
of the biology of P. americana. Eighteen authors contributed 16 chapters
that are up-to-date and written with authority. The authors describe the
distribution, life cycle and biologic importance, osmoregulation, excretion,
fat body composition and metabolism, nervous system, neurosecretion and
hormones, sense organs, rhythms, muscular activity, reproduction, phero-
mones, behavior, embryonic and post-embryonic development, and regen-
eration. The chapters give an excellent indepth review and they contain a
wealth of information about nearly all aspects of interest to those working
with cockroaches. Each chapter is concise and the information presented in
a critical manner. The more than 1,250 references are combined at the end
of the volume. A short subject index is provided. The volume is well planned
and edited. It will prove quite valuable to entomology students and teachers.
“The Laboratory Cockroach” nicely supplements the authoritative book,
providing lab exercises on the anatomy, nutrition, circulation, metabolism,
reproduction, embryogenesis, regeneration, metamorphosis and behavior. I
missed the inclusion of exercises dealing with cellular and molecular aspects.
The manual can be recommended highly for entomology courses.
Karl Maramorosch, Rutgers— The State University
304
NEW YORK ENTOMOLOGICAL SOCIETY
Plant Protection: An Integrated Interdisciplinary Approach. Webster H. Sill,
Jr. Iowa State University Press. 1982. 297 pp. $23.95.
This unique book brings together aspects of entomology, plant pathology,
nematology and weed science, as they pertain to plant protection. The exten-
sive plant protection experience of the author, who has worked for many
years as consultant in the tropics of India, the Philippines, and Taiwan,
provided the background for this very readable and useful text. The book
is aimed at practicing plant protection workers, at entomologists and plant
pathologists, as well as students at colleges and universities. In the first
chapters political, social, and economic factors of plant protection, quar-
antine, storage and marketing are discussed. Monitoring and forecasting of
outbreaks and the use of computers are stressed. The chapter on chemical
control of insects also deals with pesticides, pheromones, juvenile hormones
and insect sterilants. A separate chapter is devoted to pesticide compatibility
and toxicology and another chapter to biological control of arthropods,
diseases, nematodes, weeds and animal pests. Other chapters cover in detail
biological control by antagonism and synergism, breeding for resistance to
arthropods, and vertebrate pests. Two separate chapters are devoted to inte-
grated and combined controls, using weed and pest, as well as nematode
and pest controls. The volume concludes with a list of cited, and of other,
useful references, and with a subject index. As a whole, this volume is a well
written contribution that brings together for the first time all aspects of plant
protection. In my opinion this book should be of wide interest to entomol-
ogists engaged in biological, chemical, and integrated control and I recom-
mend it with enthusiasm to all who work in the field of crop protection.
Karl Maramorosch, Rutgers— The State University
The Ecology of Pests— Some Australian Case Histories. R. L. Kitching and
R. E. Jones, eds. Published by CSIRO, Melbourne. 1981.
‘‘The Ecology of Pests,” edited by Kitching and Jones, is a soft-cover book
that comprises review articles on pests found within Australia. The editors
have not attempted to include all of the major pests in Australia but instead
have selected pertinent examples of contemporary pest ecology. The pests
included are organisms of rather diverse life history patterns. Most of the
case histories focus on insects (aphids, codling moth, light-brown apple moth,
mosquitoes, Australian bushfly, cabbage butterfly and the Sirex woodwasp)
but there are chapters on the skeleton weed, kangaroos and the “crown of
thorns” starfish. Though these pests are discussed from the Australian per-
spective most are introduced species and will be of interest to ecologists
VOLUME XC, NUMBER 4
305
familiar with these species elsewhere in the world. There are interesting
historical discussions on the introduction of the organism to Australia (if it
is an alien species) and how it came to be regarded as a pest. The chapters
illustrate the type of information needed to evaluate the pest status of an
organism and to develop control measures that are more specific and, in the
long run, more effective than chemical pesticides alone.
The contributors are affiliated with the CSIRO or universities within Aus-
tralia. The authors present a general overview of the current status of the
pest species and have had considerable research experience with the pest
that they discuss. Most have used the life-system analysis approach and
some present formal models of the ecological processes that affect the pop-
ulation dynamics of the pest. Using modern concepts, the contributors dem-
onstrate how data collected in the field and the laboratory can be integrated
to interpret and analyze field populations.
This book will be of interest to ecologists, entomologists and agriculturists.
Most of the data presented result from field studies but results from labo-
ratory experiments are also given. Natural enemies of the pests receive
adequate attention and their potential role as biological control agents is
discussed. The text is carefully edited and well supplemented with figures
and tables. The subject index is good and chapter references are provided.
Thus, this book will be useful to students and serve as additional reading
material for courses on experimental ecology, field ecology and integrated
pest management.
Timothy J. Kurtti, Waksman Institute of Microbiology, Rutgers University
306
NEW YORK ENTOMOLOGICAL SOCIETY
ACKNOWLEDGMENT
The Editors wish to express their appreciation to all those who have helped
in reviewing the manuscripts submitted during 1982 for publication in the
Journal: S. Ahmad, P. Barbosa, D. M. Caron, R. A. Casagrande, H. Fowler,
H. Goulet, E. E. Grissell, A. P. Gupta, K. F. Harris, A. B. Klots, J. P. Kramer,
K. J. Kramer, T. J. Kurtti, J. H. Lashomb, P. E. Marucci, M. L. May, F. G.
Maxwell, H. E. McGregor, R. J. Poole, D. J. Prokopy, E. Rajotte, R. B.
Roberts, J. B. Schmidt, F. C. Swift, T. J. Walker, G. W. Wolfe.
HONORARY, LIFE, AND SUSTAINING MEMBERS
Harvey E. Barton, Raymond Brush, F. S. Chew, Lucy Clausen,
Howard E. Evans, Su Zan Swain Firmage, S. W. Frost, Irving Granek,
Mark Indenbaum, Alexander B. Klots, Kimumaro Okano, Nellie Payne,
L. L. Pechuman, Hubert J. Thelen, George Townes, Asher E. Treat,
Roman Vishniac.
INDEX TO SCIENTIFIC NAMES OF ANIMALS AND PLANTS
VOLUME XC
Generic names begin with capital letters. New genera, species, subspecies and varieties are
printed in italics. The following are not indexed:
1. Key to the species of Holcostethus in North America, pp. 5-7.
2. Fungi identified in bee cells, p. 183.
3. Non-parasitic insect species in close proximity to active bee nests, p. 184.
4. The pests of major crops and their control measures, pp. 198-200.
5. Biological control agent and its use. pp. 201-202.
Abies balsamea, 145
Acheta domesticus, 136
Aconitum fischeri, 190
lycactorum, 190
Acromyrmex, 64, 73
landolti fracticomis, 64
Acyrthosiphon pisum, 60, 87
Alasmidonta undulata, 176
Amblyaspis, 3
Amblyomma americanum, 267
Anabasis aphylla, 197
Anodonta, 178
cataracta, 176
grandis, 177
Antherea pemyi, 188
Anthophora abrupta, 1 8 1
linsleyi, 183
Apanteles omigis, 231
Aphilanthops marginipennis, 147
punctifrons, 156
Aphis abbreviata, 88
fabae, 87
gossypii, 88
nasturtii, 87
Aphodius, 162
lividus, 162
ruricola, 162
Apis cerana, 188
Apocynum, 145
androsaemifolium, 145
Arachis hypogea, 47
Artemisia scoparia, 1 90
Asphondylia helianthiglobulus, 2
Astrophea, 115, 118
Ataenius spretulus, 32
Ateuchus, 162
Atta, 64, 73
Aulacorthum solani, 88
Balsamorhiza, 271
Beauveria bassiana, 125, 197
tenella, 125
Bemisia tabaci, 6 1
Betula papyrifera, 145
Bidens cemua, 129
Bombus bimaculatus, 182
nevadensis, 182
perplexus, 182
Bombyx mandarina, 188
mori, 187
Bothriocera signoreti, 165
Brachytrupes portentosus, 189
Brevicoryne brassicae, 61, 87
Calligrapha califomica coreopsivora, 129
Calpodes ethlius, 99
Camponotus, 251
herculeanus, 252
pennsylvanicus, 249
Canna, 99
edulis, 99
Cantharis, 121
rectus, 1 1 9
scitulus, 1 1 9
Canthon, 162
viridis, 161
Carex, 145
Carya alba, 1 20
ovata, 120
Castanea, 121
Celastrus, 190
Cephaloon lepturides, 1 1 9
Ceratoma trifurcata, 60
Cerceris bimaculata, 1 5 5
cameroni, 155
Chaenomeles sinensis, 1 90
Chaetochlorops inquilinus, 3
Chilo simplex, 190
Chironomus decorus, 1 6
Chrysis, 181
308
NEW YORK ENTOMOLOGICAL SOCIETY
Chrysochus chinensis, 190
Chrysoteuchia topiaria, 94
Cirsium, 271
Citrus, 100
Cladophora, 21
Cleome, 151
Cocos nucifera, 166
Coelioxys altemata, 273
Coleoptera, 121
Comptonia peregrina, 291
Conidiobolus thromboides, 125
Copris, 162
Cordyceps militaris, 125
Cordyces, 189
Comus circinata, 1 2 1
florida, 121
Creophillus maxillosus, 163
Croton, 190
Ctenolepisma (Lepisma) vilosa, 190
Cybister, 189
Cyclocephala borealis, 32
Cydia pomonella, 261
Daphne genkwa, 1 90
Dermacentor variabilis, 269
Derris, 197
Diatraea shariinensis, 190
verosata, 190
Dibrachys cavus, 197
Digitaria decumbens, 65
smutsii, 65
Drechslera poae, 3 1
Drepanosiphum platanoides, 88
Drosophila, 241
quinaria, 3
Elliptio complanata, 176
Enteromorpha, 21
Entomophthora virulenta, 125
Epicauta, 188
Ericerus pela, 188
Eriogyma pyretorum, 188
Erythrina, 47
Eucallipterus tiliae, 88
Eueerceris, 14
arenaria, 158
atrata, 155
barri, 147
bitruncata, 148
brunnea, 147
canaliculata, 147
cavagnaroi, 147
conata, 155
geboharti, 147
melanosa, 147
melanovittata, 147
mellea, 1 50 '
melleoides, 147
menkei, 147
pimarum, 159
provancheri, 148
punctifrons, 154
punctifrons cavagnaroi, 155
rubripes, 147
sculleni, 147
similis, 149
sinuata, 152
velutina, 153
violaceipennis, 154
zimapanensis, 155
Eysenhardtia polystachya, 1 50
Franklinella, 60
Fusarium, 125, 183
Galleria mellonella, 126
Ganperdea apivora, 1 8 1
Gaylussacia, 291
Geotrupes, 162
Gleditsia, 190
Graminella nigrifrons, 60
Granadilla, 1 1 7
A, 116
Gryllus, 190
testaceus, 189
Haplaxius pallidus, 166
Heliconius, 1 1 7
congener, 1 1 5
cydno, 115, 117
eleuchia, 1 1 5
erato, 1 1 5
hewitsoni, 1 1 5
melpomene, 115, 117
pachinus, 1 1 5
sapho, 115, 117
Hemerophila atrilineata, 190
Hirsutella thompsonii, 125
Elirundo rustica, 1 8 1
Holeostethus hirtus, 5
Homoeogryllus japonicus, 189
Hoplocampa testudinea, 260
Homia minutipennis, 181
Hyalophora cercropia, 254
Hyalopterus prunii, 88
Hydrophyllum, 271
Hyparrhenia rufa, 65
Hyphantria cunea, 247
Hyppobosca capensis, 1 9 1
Illicium lanceolatum, 190
Incarvillea sinensis, 1 90
VOLUME XC, NUMBER 4
Juniperus virginiana, 120
Laccifer lacca, 188
Lampsilis siliquoidea, 176
L athromeris cicadae, 283
Leersia hexandra, 197
Lema lichensis, 60
melanopa, 60
Leptinotarsa decemlineata, 222
Leucania separata, 1 90
Ligusticum sinensis, 190
Liriodendron tulipifera, 1 1 9
Locusta, 241
migratoria, 189
migratoria manilensis, 198
Lycopersicon esculentum, 222
Lycorma delicatula, 1 9 1
Lygocoris belfragii, 1 1 9
caryae, 1 1 9
hirticulus, 1 1 9
omnivagus, 1 1 9
tiliae, 1 1 9
Lymantria dispar, 82, 125
Lytta caraganae, 188
Machaerium seemannii, 36
Macrosiphon euphorbiae, 88
scoliopi, 88
solanifolii, 88
Magicicada, 74, 276
cassini, 75, 276
septendecim, 75, 276
septendecula, 276
tredecassini, 276
tredecim, 276
tredecula, 276
Magnolia acuminata, 1 1 9
Malacosoma americanum, 243
califomicum pluviale, 247
Malus pumila, 75
sylvestris, 244
Mamestra tacoma, 142
Megachile pugnata, 270
(Sayapis) pugnata, 270
rotundata, 273
Melanagromyza matricarioides, 3
Melaphis chinensis, 188
Melittobia, 270
Meromyza saltatrix, 197
Metarhizium anisopliae, 126, 189
Miridae, 120
Mischocyttarus, 8
drewseni, 8
flavitarsis, 8
flavitarsis flavitarsis, 9
flavitarsis idahoensis, 1 4
309
flavitarsis navajo, 9
labiatus, 8
mexicanus, 8
Mnemosyne cubana, 166
Monodontomerus, 273
mandibularsis, 181
Morpho, 35
achilles, 35
amathonte, 35
anaxibia, 50
cypris, 40
granadensis, 35
granadensis polybaptus, 35
peleides, 35
peleides limpida, 35
theseus, 40
Mucuna urens, 35
Myabris phalerata, 188
Mycodiplosis, 3
Mylabris pustulata, 239
Myndus crudus, 166
Myrmeleon, 26
formicarius, 27
immaculatus, 26
obscurus, 26
Myzus persicae, 61, 87
solani, 88
Najadicola ingens, 176
Neolasioptera rudbeckiae, 3
Nephotettix cincticeps, 60
Neurocolpus nubilus, 1 20
Nesoclutha pallida, 6 1
Nilaparvata lugens, 198
Nomuraea rileyi, 1 26
Oecophylla smaragdina, 1 90
Oliarus atkinsoni, 1 66
Ontholestes cingulatus, 163
Onthophagus, 162
Opsiphanes tamarindi sikyon, 1 1 1
Orosius argentatus, 6 1
Pachydiplosis oryzae, 197
Paecilomyces farinosus, 125
fumoso-roseus, 126
Passiflora, 1 1 5
ambigua, 1 1 6
pittieri, 115, 117
Pediobius williamsoni, 183
Pennisetum ciliare, 65
Perillus circumcinctus, 1 29
Periplaneta, 240, 303
americana, 239, 303
Philanthus, 14
310
NEW YORK ENTOMOLOGICAL SOCIETY
Phyllonorycter blancardella, 233
crataegella, 23 1
Phyllotreta, 60
Physocephala marginata, 1 8 1
Pinus rigida, 29 1
Poa pratensis, 3 1
Podabrus, 122
brunnicollis, 119
rugosulus, 122
Podisus, 243
maculiventris, 244
modestus, 244
placidus, 244
Polia tacoma, 142
Polistes, 14
Popillia japonica, 32
Populus tremuloides, 27 1
Prunus, 145
serotina, 244
virginiana, 145
Psilliodes, 60
Pteridium aquilinum, 29 1
Quercus, 120
alba, 121, 291
coccinea, 121
ilicifolia, 75, 291
mba, 121
Quinta cannae, 105
Rhagoletis pomonella, 260
Rhizotrogus majalis, 32
Rhus copallina, 29 1
typhina, 120
Rhytidilomia senilis, 302
Rileya americana, 2
Robinia pseudoacacia, 120
Rodolia cardinalis, 197
rufopilosa, 197
Rudbeckia laciniata, 2
Ruta, 190
Sambucus canadensis, 145
Scapsipedus aspersus, 189
Schistocerca, 240
Semia cynthia pryeri, 188
Sitodiplosis mosellana, 190
Sitophilus oryzae, 213
Sitotroga cerealella, 1 90
Solanum melongena, 222
tuberosum, 222
Solidago, 129
Sphenophorus parvulus, 3 1
Spilochalcis, 1 1 1
Spiraea latifolia, 145
Spirodela, 190
Staphylinus maculosus, 1 6 1
Stemona, 190
Strophitus undulatus, 178
Sympiesis conica, 231
marylandensis, 231
Teleogryllus commodus, 140
Tilia, 88
americana, 120
Torymus, 2
advenus, 2
Trachymyrmex iheringi, 70
kempfi, 70
Trichodes omatus, 270
Trichogramma, 111, 197
Trichordestra, 142
liquida, 142
rugosa, 142
tacoma, 142
Tripsacum laxum, 65
Trirhabda, 129
Tryporyza (=Schoenobius) incertulas, 198
Unionicola abnormipes, 179
aculeata, 179
arcuata, 176
formosa, 176
formosa-ypsilophora, 176
fossulata, 176
serrata, 179
tumida, 176
ypsilophora, 177
Unkanodes albifascia, 60
Vaccinium, 291
myrtilloides, 145
Vasates fockeni, 60
Verticillium, 125
Viburnum acerifolium, 121
Vida faba, 87
Viola selkirkii, 145
Xanthium, 190
Zenillia roseanae, 197
Zingiber mioga, 1 90
,r
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Journal of the
New York Entomological Society
published by
The New York Entomological Society
Contents Volume 91, 1983, Numbers 1-4
Number 1
A revision of the genus Linens St^l (Hemiptera: Pentatomidae: Discocephalinae:
Ochlerini) L. H. Ralston 1-47
On the biology and food plants of Lygaeus turcicus (Fabr.) (Hemiptera: Lygaeidae)
James A. Slater 48-56
The small milkweed bug, Lygaeus kalmii (Hemiptera: Lygaeidae): milkweed specialist
or opportunist? A. G. Wheeler, Jr. 57-62
A new species of Cleradini (Hemiptera: Lygaeidae: Rhyparochrominae) from the
Central African Republic and Ghana B. J. Harrington 63-67
Overwintering egg mass adaptations of the eastern tent caterpillar, Malacosoma
americanum (Fab.) (Lepidoptera: Lasiocampidae)
Alejandro Seggara Carmona and Pedro Barbosa 68-74
Melanism in Phigalia titea (Cramer) (Lepidoptera: Geometridae): a fourteen-year
record from central Massachusetts Theodore D. Sargent 75-82
The Diptera breeding on skunk cabbage, Symplocarpus foetidus (Araceae)
David Grimaldi and John Jaenike 83-89
Notes and Comments
Predation of Prosimulium mixtum/fuscum (Diptera: Simuliidae) copulating pairs by
Formica ants (Hymenoptera: Formicidae) Dennis J. White 90-9 1
The first occurrence of Catorhintha mendica Stal in New England (Hemiptera:
Coreidae) James A. Slater 91-92
Book Review
The North American Grasshoppers. Volume I. Acrididae: Gomphocerinae and
Acridinae K. H. L. Key 93-96
Number 2
A revision of the genus Acrosternum Fieber, subgenus Chinavia Orian, in the western
hemisphere (Hemiptera: Pentatomidae) L. H. Ralston 97-176
I Pathogenicity of the fungus Entomophthora culicis for adult mosquitoes: Anopheles
j stephensi and Culex pipiens quinquefasciatus John Paul Kramer 177-182
The genus Paralincus (Hemiptera: Pentatomidae) L. H. Ralston 183-187
Book Review
Advances in cladistics: proceedings of the first meeting of the Willi Hennig Society
James M. Carpenter 188-192
Number 3
Exotic insects reported new to Northeastern United States and Eastern Canada since
1970 E. Richard Hoebeke and A. G. Wheeler, Jr. 193-222
The types of cercerine wasps described by Nathan Banks (Hymenoptera:
Philanthidae) George R. Ferguson 223-234
Two new species and synonymy of three species of North American Cerceris (Hy-
menoptera: Philanthidae) George R. Ferguson 235-241
Gynandromorphic desert fire ant, Solenopsis aurea Wheeler (Hymenoptera:
Formicidae) James C. Cokendolpher and Oscar F. Franke 242-245
A redefinition of Disderia and addition of a new species (Hemiptera: Pentatomidae)
F. H. Ralston 246-251
A study of winged queens of the Colorado honey ant, Myrmecocystus rnexicanus, in
captivity John R. Conway 252-263
Nesting biology of the bee Svastra sabinensis (Hymenoptera: Anthophoridae)
Jerome G. Rozen, Jr. 264-268
Response of a goldenrod beetle to four seldom-encountered goldenrod (Solidago)
species Frank J. Messina 269-272
Precopulatory behavior in the whirligig beetle Dineutes discolor (Coleoptera:
Gyrinidae) Steven A. Kolmes 273-279
SEM study of the antennal sensilla and setae of Solva pallipes (Loew) (Diptera:
Xylomyidae) C. E. Vasey and E. Ritter 280-282
Book Reviews
The Growth of Biological Thought. Diversity, Evolution, and Inheritance
William F. Brown, Jr. 283-284
Vicariance Biogeography: A Critique Stephen W. Nichols 284-288
Number 4
Revision of the Philanthus zebratus group (Hymenoptera: Philanthidae)
George R. Ferguson 289-303
Establishment of Hyles euphorbiae (h.) (Lepidoptera: Sphingidae) in the United States
for control of the weedy spurges Euphorbia esula L. and E. cyparissias L.
S. W. T. Batra 304-311
Patterns of distribution and abundance in small samples of litter-inhabiting orthoptera
in some Costa Rican cacao plantations Allen M. Young 312-327
Coordinated prey capture by Novomessor cockerelli (Hymenoptera: Formicidae)
Harold G. Fowler and Walter G. Whitford
Habitat preferences of carrion beetles in the Great Swamp National Wildlife Refuge,
New Jersey (Coleoptera: Silphidae, Dermestidae, Nitidulidae, Histeridae, Scara-
baeidae) Paul P. Shubeck
Microclimate observations and diel activities of certain carrion arthropods in the
Chihuahuan desert Kenneth Schoenly
Seasonal dynamics of fleas associated with the gray-tailed vole, Microtus canicaudus
Miller, in western Oregon Richard G. Robbins
Histology of the male reproductive systems in the adults and pupae of two doryline
ants, Dorylus (Anomma) wilverthi Emery and D. (A.) nigricans Illiger (Hymenop-
tera: Formicidae) Francis C. Ford and James Forbes
Anatomy and histology of the male reproductive system in the adult and pupa of the
doryline ant, Aenictus gracilis Emery (Hymenoptera: Formicidae)
S. Shyamalanath and James Forbes
Cuticular pigment changes in worker Yellowjackets (Hymenoptera: Vespidae)
Kenneth G. Ross
Ecological and sensory aspects of prey capture by the whirligig beetle Dineutes discolor
(Coleoptera: Gyrinidae) Steven A. Kolmes
Irbisia knighti, a new mirine plant bug (Heteroptera: Miridae) from the Pacific
Northwest Michael D. Schwartz and John D. Lattin
Descriptions of the nymphal instars of Oecleus borealis (Homoptera: Fulgoroidea:
Cixiidae) Stephen W. Wilson, James H. Tsai, and Catherine R. Thompson
On the biology of cave inhabiting Antillocorini with the description of a new species
from New Guinea (Hemiptera: Lygaeidae) James A. Slater
The types of some American Cerceris with lectotype designations (Hymenoptera:
Philanthidae) George R. Ferguson
Type designations and new synonymies for Nearctic species of Phytocoris Fallen
(Hemiptera: Miridae) Thomas J. Henry and Gary M. Stonedahl
An annotated synonymic list of North American and Caribbean wasps of the genus
Cerceris (Hymenoptera: Philanthidae) George R. Ferguson
Recognition of host nest odour by the bumblebee social parasite Psithyrus ashtoni
(Hymenoptera: Apidae) Richard M. Fisher
Notes and Comments
Mimicry, predation and potential pollination by the mantispid, Climaciella brunnea
var. instabilis (Say) (Mantispidae: Neuroptera) Thomas C. Boyden
Book Reviews
The nesting behavior of dung beetles (Scarabaeinae). An ecological and evolutive
approach Brett C. Ratcliffe
Ecologie des insectes forestiers Gordon R. Stairs
The biology of social insects Janice S. Edgerly
Insects of an Amazon forest Harold Fowler
The semiaquatic bugs (Hemiptera, Gerromorpha): Phylogeny, adaptations, biogeog-
raphy, and classification Randall T. Schuh
Reviewers for 1983
328-332
333-341
342-347
348-354
355-376
377-393
394-404
405-412
413-417
418-423
424-430
431-441
442-465
466-502
503-507
508-511
512-516
516
516-521
521
522-523
524
.*ri .
Vi.
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Vol. 91
MARCH 1983
No. 1
Journal
of the
New York
Entomological Society
(ISSN 0028-7199)
Devoted to Entomology in General
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY
Editor: Randall T. Schuh, Department of Entomology, American Museum
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York 10024
Book Review Editor: Quentin D. Wheeler, Department of Entomology,
Cornell University, Ithaca, New York 14853
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The New York Entomological Society
Incorporating The Brooklyn Entomological Society
President: Gerard Iwantsch, Department of Biological Sciences, Eordham
University, Bronx, New York 10458
Vice President: Henry M. Knizeski, Jr., Department of Biology, Mercy
College, Dobbs Ferry, New York 10522
Secretary: Irene E. Matejko, Science Department, The New Lincoln School,
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Treasurer: Louis Sorkin, Department of Entomology, American Museum
of Natural History, New York, New York 10024
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Joseph Cerreta, Columbia University, New York; Durland Fish, Ford-
ham University, Bronx, New York.
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Meetings of the Society are held on the third Tuesday of each month (except June through
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Mailed June 7, 1983
The Journal of the New York Entomological Society (ISSN 0028-7199) is published quarterly (March, June, September,
December) for the Society by Allen Press, Inc., 1041 New Hampshire, Lawrence, Kansas 66044. Second class postage paid
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Known oflicc of publication: American Museum of Natural History, New York, New York 10024.
Journal of the New York Entomological Society, total copies printed 600, paid circulation 443, mail subscription 443, free
distribution by mail 7, total distribution 450, 150 copies left over each quarter.
NEW YORK ENTOMOLOGICAL SOCIETY
91(1), 1983, pp. 1-47
A REVISION OF THE GENUS UNCUS STAl (HEMIPTERA:
PENTATOMIDAE: DISCOCEPHALINAE: OCHLERINI)
L. H. Rolston
Department of Entomology, Louisiana Agricultural Experiment Station,
Louisiana State University, Baton Rouge, Louisiana 70803
Abstract.— Tht genus Linens is redefined and Minilincus Ruckes, 1958, placed in synonymy.
The 30 known species of Linens are described or redescribed and a key provided to assist in
their recognition. New species are: annlatns, breddini, convexns, cronpins,fcitigns, incisns, lami-
natns, leviventris, manchns, modiens, operosns, repizens, singnlaris, sinnosns, snbstyliger, tn-
midifrons, vallis, vandoesbnrgi and varins.
INDEX TO SPECIES
anulatus, new species 22
Breddin 32
breddini, new species 33
convexus, new species 16
croupius, new species 12
dentiger Breddin 39
discessus (Distant) 26
fatigus, new species 10
incisus, new species 9
lamelliger Breddin 35
laminatus, new species 41
leviventris, new species 30
lobuliger Breddin 35
rnanchus, new species 31
modicus, new species 28
operosus, new species 14
parvulus (Ruckes) 1 7
repizeus, new species 24
rufospilotus (Westwood) 36
securiger Breddin 1 7
singularis, new species 18
sinuosus, new species 1 3
The publication costs of this article were defrayed in part by page charge payment. This article
must therefore be hereby marked “Advertisement” m accordance with 18 U.S.C. §1734 solely
to indicate this fact.
2
NEW YORK ENTOMOLOGICAL SOCIETY
sp athuli ger 'QvQddin 10
styliger Breddin 40
substyliger, new species 43
subuliger Breddin 46
tumidifrons, new species 20
vallis, new species 38
vandoesburgi, new species 7
varius, new species 26
The genus Lincus was last defined by Stal ( 1 872) in a key to related genera.
At that time the genus was monotypic, based on Pentatoma rufospUota
Westwood, and characterized largely by an elongated lobe at each antero-
lateral angle of the pronotum. The form of these lobes no longer distinguishes
Lincus because among species subsequently and correctly added to the genus
the pronotal lobes vary much in shape and degree of development. A revision
of Lincus is desirable not only to redefine the genus but also to lessen the
difficulty of identifying the contained species, most of which have been
unrecognized previously.
The genus is meagerly represented in collections, and rare is the series
taken at the same time, unless at a light, that contains both sexes. Conse-
quently, knowledge of geographical distribution and variation is sketchy,
and the correct association of sexes becomes a greater problem than usual,
especially so because most species are poorly endowed with useful taxonomic
characters other than the genitalia. As specimens of the genus accumulate
in collections, so too should information on distribution and variation as
well as evidence to corroborate or refute my conclusions with regard to the
association of sexes.
Lincus Stal
Lincus Stal, 1867, p. 524.
Minilincus Ruckes, 1958, pp. 14-15. New Synonymy.
Type species. Lincus: Pentatoma rufospilotus Westwood, 1837. Minilin-
cus: Minilincus parvulus Ruckes, 1958.
Labium originating behind imaginary plane bisecting head at anterior limit
of eyes when viewed ventrally and perpendicularly to longitudinal axis of
thorax and abdomen; second rostral segment extending a little past meso-
coxae; apex of rostrum reaching penultimate or ultimate abdominal segment.
Antennae 5-segmented, basal segment reaching or surpassing apex of head.
Juga as long as or longer than tylus, sometimes contiguous apically. Length
of head before ocelli 0.6-0. 8 of width across eyes. Anterolateral angles of
VOLUME 91, NUMBER 1
3
pronotum usually developed into prominent lobe, this sometimes expanded
apically. Scutellum longer than wide at base, 0.4-0. 5 as wide at distal end
of frena as at base. Costal angle of coria extending past apex of scutellum.
Metasternum flat or weakly tectiform mesially. Pygophore deeply and broad-
ly emarginate mesially; a hemispherical impression present mesially between
bottom of pygophoral emargination and inferior ridge (Fig. 2).
Comments. Ruckes (1958) separated Minilincus from Lincus on the basis
of smaller size, lighter color and reduced pronotal lobes. These diflerences
are of specific value only and Minilincus is therefore considered to be a
junior synonym of Lincus.
Those few species of Lincus of which sufficient specimens were available
for dissection share several characteristics that may be common to the genus.
These characteristics are excluded from the generic description because of
the meagerness of the evidence.
The proctiger is essentially tubular, usually duck-bill shaped, with a ventral
opening (Figs. 39, 47, 99); it often protrudes well beyond the posterior
pygophoral margin (Figs. 48, 1 12). The parameres are relatively small, not
visible within the genital cup, and doubtfully functional (Figs. 5, 38, 102).
The aedeagus is typically discocephaline, heavily pigmented and sclerotized
excepting the conjunctiva, which is restricted to a sheath around the pro-
jecting portion of the ejaculatory duct; the ejaculatory duct emerges sub-
apically and ventrally (Figs. 4, 42, 103).
The spermathecal bulb is spherical with 3 equidistant appendages that
curve basad and, viewed terminally, clockwise (Figs. 43, 44).
Among ochlerines, the only genera with a flat or weakly tectiform meta-
sternum are Lincus and an unnamed genus. The latter genus difiers from
Lincus in that the costal angles of the coria project little if at all caudad of
the scutellar apex, and the scutellar width at the end of the frena is about
0.6 the basal width rather than 0.5 or less as in Lincus.
The subfamily and tribe to which Lincus belongs have been characterized
recently (Rolston and McDonald, 1979; Rolston, 1981).
SPECIES GROUPS OF CONVENIENCE
As an aid to identification, the 30 known species of Lincus may be arranged
in species groups of convenience.
The “hatchet-lobed” group of 9 species has the anterolateral pronotal
angles enlarged into lobes that are expanded subapically (into a form which
I fancy resembles a hatchet blade) or that are broad throughout their length
(Figs. 1, 12). L. convexus, croupius,fatigus, incisus, operosus, securiger, sinu-
osus, spathuliger and vandoesburgi belong in this group. Most of these species
have relatively large eyes, the width of each eye being equal to or greater
than one-half of the interocular width.
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NEW YORK ENTOMOLOGICAL SOCIETY
The “swollen head” group is readily recognized by the tumescent vertex
of the head. Included here are L. parvulus, singularis and tumidifrons. This
is an artificial assembly if ever there was one.
In the “little-eyes” group the width of each eye is less than one-half of the
interocular width. This group consists of 7 species: L. armiger, discessus,
leviventris, manchus, modicus, repizcus and varius.
In the “big-eyed” group of 10 species, by contrast, the width of each eye
is equal to or greater than one-half of the interocular width. The included
species are L. breddini, dentiger, lamelliger, laminatus, lobuliger, rufospi-
lotus, styliger, substyliger and vallis.
Finally, there is L. anulatus, which stands by itself as the only relatively
gaudy member of a drab genus. The femoral and tibial bands distinguish
this species, which refuses to fall neatly into any of the above species groups.
Too much should not be inferred from the above groupings with respect
to phylogeny. The ancestral form of Lincus presumably resembled L. dis-
cessus and other species of the “little-eyed” group without notably developed
pronotal lobes, but the lines of evolvement from this form are unclear.
Enlargement of the anterolateral pronotal denticles into lobes and widening
of the eyes did not go hand in hand. In both the “little-eyed” and “big-
eyed” groups there is considerable variation in the degree of lobe develop-
ment, and although most species in the “hatched-lobed” group have enlarged
eyes, others do not. The most parsimonious hypothesis regarding intrage-
neric lineage is that either pronotal lobe enlargement or eye widening evolved
more than once, as did the tumescent vertex.
KEY TO Lincus SPECIES
1 . Pronotal lobes expanded subapically, if expansion slight then each lobe wider than
anterior-posterior diameter of an eye (Figs. 1,12) 2
- Pronotal lobes parallel sided or tapering toward apex, if slightly expanded sub-
apically then much narrower than anterior-posterior diameter of an eye (Figs. 30,
34) 10
2(1). Pronotal margins caudad of lobes expanded on each side into obtuse projection
(Figs. 1, 8) 3
- Pronotal margins sinuous or straight (Figs. 10, 12) 4
3(2). Second antennal segments about half as long as first, incision between lobe and
anterolateral margin of pronotum extending mesad little beyond lateral limit of
eye (Fig. 1) .vandoesburgi, new species
Second antennal segments subequal to first in length; incision extending mesad
to middle of eye (Fig. 8) incisus, new species
4(2). Second antennal segments about half as long as first spathuliger Breddin
Length of second antennal segments subequal to or greater than first 5
5(4). Pronotal lobes strongly deflexed 6
- Pronotal lobes horizontal or reflexed 7
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5
6(5).
7(5).
8(7).
9(7).
10(1).
11(10).
12(11).
13(10).
14(13).
15(14).
16(15).
17(16).
18(17).
19(16).
20(15).
Pronotal lobes strongly expanded subapically (Fig. 10); juga contiguous before
tylus fatigus, new species
Pronotal lobes weakly expanded subapically, broad throughout their length (Fig.
12); juga little longer than tylus cwupius, new species
Small species, about 1 0 mm or less in length 8
Larger species, more than 11.5 mm in length 9
Anterolateral pronotal margins strongly sinuous (Fig. 16); margins of mesial py-
gophoral emargination produced anteromesad on each side at dorsal limit of
emargination (Fig. 17) sinuosus, new species
Anterolateral pronotal margins moderately sinuous (Fig. 19); margins of mesial
pygophoral emargination uniformly narrow, without projections (Fig. 20)
operosus, new species
Incision caudad of pronotal lobes reaching mesad to middle of eye; lobes projecting
laterad of eye by 0.2 width of eye or less; lateral margins of juga converging toward
apex (Fig. 22) convexus, new species
Incision caudad of pronotal lobes reaching mesad from lateral limit of eye by
about 0.2 width of eye; lobes projecting laterad of eye by about 0.5 width of eye;
lateral margins of juga subparallel between anteocular concavity and apex (Fig.
25) securiger Breddin
Vertex of head strongly convex, appearing swollen 1 1
Vertex of head normally convex 13
Small species, about 9 mm in length; distance across ocelli greater than interocular
width; second antennal segment at least as long as first parvulus (Ruckes)
Larger species, about 11-14 mm in length; distance across ocelli and interocular
width subequal; second antennal segment shorter than first 1 2
Lateral angle of basal plates produced, projecting over base of 8th paratergites
(Fig. 35); pronotal lobes projecting laterad of eyes about 0.3 width of eye
singularis, new species
Lateral angle of basal plates not produced (Fig. 37); pronotal lobes seldom pro-
jecting as far laterad of eyes as 0.3 width of eye tumidifrons, new species
Femora with broad preapical fuscous ring anulatus, new species
Femora not annulated 1 4
Width of eye less than half of interocular width 1 5
Width of eye at least half of interocular width 21
Pale narrow yellow low ridge on ventral surface of head running from base of
head to antennifer at mesial margin of eye 1 6
Venter of head caudad of eye smooth excepting punctures 20
Lateral margins of abdominal venter with large yellowish brown macule on each
segment 17
Abdominal venter uniformly colored 1 9
Lateral margins of head converging from preocular concavity toward apex
repizcus, new species
Lateral margins of head parallel between preoccular concavity and apex 1 8
Pygophoral emargination U-shaped discessus (Distant)
Pygophoral emargination V-shaped varius, new species
Disk of abdominal venter multitudinously and very shallowly cratered laterally;
mesial margin of 9th paratergites not reflexed modicus, new species
Abdominal venter nearly smooth, impressions faint; mesial margin of 9th para-
tergites narrowly but strongly reflexed (Fig. 65) leviventris, new species
Ocelli large, greatest diameter of each about 0.30-0.35 of distance between ocelli
manchus, new species
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NEW YORK ENTOMOLOGICAL SOCIETY
21(14).
22(21).
23(22).
24(23).
25(24).
26(25).
27(26).
28(25).
29(28).
30(24).
31(23).
32(31).
33(31).
Ocelli smaller, greatest diameter of each about 0.22 of distance between ocelli ...
- — armiger Breddin
Incision caudad of pronotal lobes reaching mesad as far as middle of eye (Fig. 71)
- breddini, new species
Incision caudad of pronotal lobes shallower 22
Length of pronotal lobes from base of incision to apex of lobe subequal to width
of lobes at base (Fig. 75) lamelliger Breddin
Pronotal lobes triangular or if not triangular longer than wide at base (Figs. 90,
95) 23
Males 24
Females 3 1
Apex of proctiger transversely convex, narrowly rounded from dorsal view (Fig.
99) 25
Apex of proctiger flattened and expanded (Figs. 107, 1 12) 30
Mesial emargination of pygophore U-shaped from caudal view (Figs. 80, 85) 26
Mesial emargination of pygophore lyre-shaped or V-shaped from caudal view
(Figs. 91, 97) 28
Posterior margin of pygophore in profile strongly convex (Fig. 81)
lobuliger Breddin
Posterior margin of pygophore in profile sinuously linear (Fig. 86) 27
Elevated rim along lateral margins of pygophoral emargination disappearing be-
neath bottom of emargination (Fig. 85); 2nd and 3rd antennal segments subequal
in length, 5th almost uniformly colored rufospilotus (Westwood)
Elevated rim along lateral margins of pygophoral emargination disappearing on
level with bottom of emargination (Fig. 88); 3rd antennal segment about 1.5 times
length of 2nd, basal 4th of 5th segment paler than remainder of segment
vail is, new species
Pygophoral emargination lyre-shaped from caudal view, narrowed where rim along
emargination first becomes prominent about a 3rd of distance from dorsal margin
of pygophore to bottom of emargination (Fig. 9 1 ) dentiger Breddin
Pygophoral emargination V-shaped from caudal view 29
A single broad lamina present in genital cup on each side of posterior wall (Fig.
99) styliger Breddin
Two smaller laminae present in genital cup on each side of posterior wall (Fig.
101) - laminatus, new species
Apical expansion of proctiger about a 4th wider than proctiger at base of expansion
(Fig. 107); rim of pygophoral emargination continuing into genital cup as lamina
on each side substyliger, new species
Apical expansion of proctiger about three-fourths wider than proctiger at base of
expansion (Fig. 1 1 2); rim of pygophoral emargination continuing into genital cup
as low ridge on each side subuliger Breddin
Lateral margins of head subparallel between anteocular concavity and apex (Fig.
79) 33
Lateral margins of head converging toward apex between anteocular concavity
and apex (Figs. 90, 94) .... 32
Pronotal lobes triangular, projecting laterad of eyes by about 0.2 width of eye ....
dentiger Breddin
Pronotal lobes elongated, projecting laterad of eyes by about half the width of an
eye styliger Breddin
Rugose interstices between punctures on abdominal venter brownish yellow, punc-
tures and area around punctures much darker lobuliger Breddin
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7
- Abdominal venter uniformly dark or nearly so 34
34(33). Smaller species, about 5.7 mm wide at humeri subuliger Breddin
Larger species, about 6.5 mm wide at humeri rufospilotus (Westwood)
Lincus vandoesburgi, new species
(Figs. 1-7)
Description. Appearing dark brown above and below due to thick fuscous
to dark castaneous punctation on yellowish background. Yellowish brown
macule present on humeri, along base of scutellum near angles and in middle,
on disk of each corium and on each connexival segment. Abdominal venter
dark castaneous mesially. Thoracic sterna and much of pleura black. Ap-
pendages predominately yellowish brown.
Antennal segment 2 short; length of segments 0. 8-1.0, 0.4-0. 5, 1.0-1. 2,
1 .6-2.0, 2. 5-2. 6 mm. Head 2. 3-2. 6 mm wide across eyes, 2. 1-2.3 mm long.
Width of eyes about half of interocular width, slightly greater than anterior-
posterior diameter of eyes. Distance across ocelli slightly greater than inter-
ocular width, 1.2- 1.3 and 1. 1-1.2 mm respectively. Juga and tylus equally
long, lateral jugal margins before anteocular concavity converging slightly
toward apex of head (Fig. 1).
Pronotal lobes extending laterad of eye by 0.6-0. 8 width of eye, much
expanded apically and curved dorsad. Anterolateral pronotal margins pro-
duced into obtuse projection laterad of lobe on each side, leaving curved
incision between this margin and lobe. Disk rugosely punctate. Width at
humeri 6.0-6. 7 mm, mesial length 2. 6-2. 8 mm.
Scutellum 3.6-4. 1 mm wide at base, 4. 5-5.0 mm long, with black trian-
gular fovea containing a few punctures at basal angles. Disk rugosely punctate
basally.
Basal plates truncate apically, sometimes lightly impressed along lateral
margins (Fig. 7). Distal portion of spermatheca as in Figure 6; three projec-
tions on bulb curving basad and clockwise from terminal view of bulb.
Large mesial emargination of pygophore slightly concave at dorsal limits
on each side; in dissected pygophore emargination entad of external rim
sinuous (Fig. 2); pygophore in lateral view as in Figure 3. Proctiger pro-
truding, visible from above. Parameres short relative to phallus (Fig. 4),
transversely expanded basally (Fig. 5).
Length 12-13.3 mm.
Distribution. Surinam.
Holotype. 9, labeled “Museum Leiden, Dr. D. C. Geijskes, Paramaribo,
Cultuuruin, 28 VIII 1938” Deposited in the Rijksmuseum van Natuurlijke
Historic.
NEW YORK ENTOMOLOGICAL SOCIETY
Figs. 1-7. Linens vandoesburgi. 1 . Head and pronotum. 2. Pygophore, caudal view, proctiger
removed. 3. Same, lateral view. 4. Relationship of aedeagus and parameres within pygophore,
right lateral view, basal plate omitted. 5. Right paramere, dorsal view. 6. Genital plates, cau-
doventral view; basal plate (BP). 7. Distal parts of sphermatheca.
Paratypes. 599, 1$$. Same data as holotype (299, 3<$<3 RNH; 19 LHR); same
data as holotype except date “24 VIII 1938” (6 RNH), “29 VII 1938” (9
RNH) and “14 IV 1938” without word Cultuuruin (3 LHR); (a) “Surin.
1158-19 P. H. V. Doesburg, Jr.” (b) “611” on red label {S RNH); (a) “on
roots Liberian coffee” (b) “Jaglust plantation” (c) “Paramaribo, Surinam.
D. G. VII-28-33” (d) “Lincus spathuliger Bredd. det. H. G. Barber” (9 AL);
VOLUME 91, NUMBER 1
9
Figs. 8, 9. Lincus incisus. 8. Head and pronotum. 9. Genital plates, caudoventral view.
same data without determiner’s name (5 USNM); (a) Victoria, oliepalm, 15-
6-77, V. Slobbe (b) L. P. 815 (c) Surinam {6 USNM).
Comments. The shape of the pronotum and the short 2nd antennal seg-
ment are together diagnostic. L. spathuliger, which is known only from the
female holotype, appears to differ only in lacking the obtuse projection on
the anterolateral pronotal margins just laterad of the lobes. Since the prove-
nance of L. vandoesburgi is Surinam and that of L. spathuliger is Peru, it is
possible that these two taxa are subspecies.
This species is named in honor of Dr. P. H. van Doesburg, Jr. of the
Rijksmuseum van Natuurlijke Historic.
Lincus incisus, new species
(Figs. 8, 9)
Description. Fuscous to black with rugose interstices between punctures;
discal spot on each corium, last 2 antennal segments, rostrum and tarsi
yellowish brown.
Antennal segments 0.9, 1.2, 1.3, 1.4, 1.9 mm in length; basal segment
extending well beyond apex of head. Rostral segments 2-4 about 2.4, 2.0,
1.8 mm in length. Eyes wider than anterior-posterior diameter, 0.8-0.85
mm wide and 0.6 mm in diameter from dorsal view. Interocular width and
distance across ocelli equal, 1.15 mm; greatest diameter of ocelli 0.3 mm.
Juga nearly contiguous apically, their lateral margins before anteocular con-
cavity tapering slightly toward apex (Fig. 8). Width of head across eyes 2.8
mm, length 2.2 mm. Vertex nearly flat.
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NEW YORK ENTOMOLOGICAL SOCIETY
Pronotal lobes expanded apically, projecting horizontally laterad of eyes
by about 0.3 width of eye; incision caudad of lobes extending mesad from
lateral limit of eye by about 0.4 width of corresponding eye. Width of prono-
tum at humeri 6.5 mm, mesial length 3.0 mm. Disk rugosely punctate,
interstitial rugae yellowish brown. Anterolateral margins produced into ob-
tuse angle caudad of apex of lobes.
Scutellum 4.0 mm wide at base, 5.1 mm long. Triangular black fovea
present in each basal angle. Disk rugose.
Posterior margins of basal plates evenly convex (Fig. 9).
Length about 12.8 mm.
Distribution. Surinam. Known only from holotype.
Holotype. $, labeled “Museum Leiden, Suriname, Mapane area, Blakka
Watra, 26. V. 1960, Ph. van Doesburg, Jr.” Deposited in the Rijksmuseum
van Natuurlijke Historic. The 5th segment of the right antenna is glued on
the label.
Lincus spathuliger Breddin
Uncus spathuliger^xtddm, 1908, pp. 31-33, figs. 15, 16; Gaedike, 1971, p.
99 (lectotype designated).
Diagnosis. Pronotal lobe on each side extending laterad of eye by distance
nearly equal to width of eye, expanding toward apex, leaving deep narrow
incision between posterior margin of lobe and anterolateral margin of prono-
tum; latter margin sinuous, quite convex caudad of lobe. Second antennal
segment about one-half length of first. Juga and tylus subequal in length;
lateral jugal margins subparallel between concavity above antennifers and
apex of head. Posterior margin of basal plates obtusely angled where base
of 9th paratergites meet 8th paratergites.
Length including membranes about 1 1 mm.
Distribution. Known only from lectotype collected at Marcapata, Peru.
Comments. Of those species with apically expanded pronotal lobes, only
this species and vandoesburgi have short second antennal segments, about
half as long as the first. L. spathuliger is distinguished from L. vandoesburgi
by the absence of a subapical projection on the anterolateral pronotal mar-
gins.
The lectotype was examined.
Lincus fatigus, new species
(Figs. 10, 11)
Description. Mostly fuscous; discal spot on each corium, mesial spot at
base of scutellum, last 2 antennal segments, some interstitial rugae on scu-
VOLUME 91, NUMBER 1
Figs. 10, 11. Lincus fatigus. 10. Head and pronotum. 1 1. Genital plates, caudoventral view;
basal plate (BP); 9th paratergite (Pt9).
tellum, tarsi and scattered flecks on venter yellowish brown; some interstitial
rugae on anterior disk of pronotum and humeri reddish.
Antennal segments 1.0, 1.3, 1.4, 1.5, 1.9 mm long; first segment projecting
well beyond apex of head. Head 2.9 mm wide across eyes, 2.4 mm long.
Eyes wider than anterior-posterior length, each eye 0.9 mm wide, 0.6 mm
long. Distance between lateral limits of ocelli greater than interocular dis-
tance, 1.3 mm across ocelli, 1.1 mm between eyes. Juga contiguous before
tylus (Fig. 10).
Pronotal lobe extending laterad of eye on each side about 0.4 width of
eye, much expanded apically and bent ventrad; narrow incision between
posterior margin of pronotal lobe and convex expansion of anterolateral
pronotal margin reaching nearly to middle of eye. Pronotal disk rugosely
punctate. Width at humeri 7.3 mm, mesial length 3.4 mm.
Scutellum 4.6 mm wide at base, 5.3 mm long. Deep fovea present in basal
angles. Basal part of disk comparable to pronotum in rugoseness.
Basal plates broadly rounded at posterior margin, rather smoothly surfaced
with a few large punctures laterally. Paratergite 9 transversely convex (Fig.
11).
Length about 13.5 mm.
Distribution. Surinam. Known only from type.
Holotype. 9, labeled “S. Suriname, Sipalivini, 14 June 1963, P. H. v.
Doesburg Jr.” and on red label “609”. Deposited in the Rijksmuseum van
Natuurlijke Historie. The last 3 rostral segments, most of middle legs and
part of antennal segment 5 on left are missing.
No paratypes.
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NEW YORK ENTOMOLOGICAL SOCIETY
Figs. 12-15. Lincus croupius. 1 2. Head and pronotum. 1 3. Pygophore, caudal view, proctiger
omitted. 14. Same, lateral view. 15. Genital plates, caudo ventral view; basal plates and 9th
paratergites partially opened.
Lincus croupius, new species
(Figs. 12-15)
Description. Mostly fuscous; last 2 segments of antennae, discal spot on
each corium, rostrum and tarsi yellowish brown.
Antennal segments 0.8, 1. 1-1.2, 1.0- 1.1, 1.3, 1.7 mm in length, first pro-
jecting slightly beyond apex of head. Head 2.3 mm wide across eyes, 2.0-
2. 1 mm long. Eyes wider than anterior-posterior diameter, each 0.6-0. 7 mm
wide, 0.45-0.5 mm in diameter. Distance across ocelli greater than inter-
ocular width, 1.1-1.15 mm across ocelli, 1.0-1.05 mm between eyes. Juga
slightly longer than tylus. Rostral segments 2-4 about 2. 6-2. 7, 2.1, 1. 9-2.0
mm in length.
Pronotal lobes broad throughout, little expanded apically, deflexed toward
VOLUME 91, NUMBER 1
13
Figs. 16-18. Lincus sinuosus. 16. Headand pronotum. 17. Pygophore, caudal view, proctiger
omitted. 18. Same, profile.
apex, concave basally, extending laterad of eye on each side about 0.5 width
of eye (Fig. 1 2); emargination caudad of lobe not reaching mesad as far as
lateral limit of eye. Anterolateral margins caudad of lobes nearly straight.
Pronotal disk rugose. Width of pronotum across humeri 5. 8-6. 2 mm, mesial
length 2. 6-2. 8 mm.
Scutellum 3.8 mm wide at base, 4. 3-4. 6 mm long. Basal angles shallowly
foveate. Disk rugose basally, smoother than pronotum.
Genital plates as in Figure 15.
Outline of pygophoral emargination interrupted near dorsal margin dorsad
of setose patch (Fig. 1 3). Posterior margin of pygophore sinuous in profile
(Fig. 14).
Length about 10 mm.
Distribution. Brazil (Amapa), French Guiana.
Holotype. 5, labeled (a) “Rio Pelioca, Terr. Amapa, Brasil, 4-VIII-1957,
J. Lane Leg.” (b) “Colegao J. Lane.” Deposited in the Museum de Zoologia,
Sao Paulo.
Paratype. 9, labeled “Museum Paris, Guyane Frang, Camopi, F. Geay,
1900” (b) “693” (AMNH).
Comment. This species resembles L. fatigus in having deflexed pronotal
lobes. The lobes are much broader basally and the juga much shorter, relative
to the tylus, in L. croupius than in L. fatigus.
Lincus sinuosus, new species
(Figs. 16-18)
Description. Dark brown with yellowish brown vermiform short lines
above and below, most of these marking elevated rugae; basal half of head
fuscous; humeri, 3 basal spots on scutellum, spot on disk of each corium
and in middle of each connexival segment yellowish-brown.
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NEW YORK ENTOMOLOGICAL SOCIETY
Antennal segments 0.7, 0.7, 1.1, 1.1 mm long (5th missing); first segment
reaching apex of head. Head 2.05 mm wide across eyes, 1.9 mm long. Eyes
wider than anterior-posterior diameter, each about 0.5 mm wide, 0.4 mm
in diameter. Interocular width 1.1 mm, greater than distance of 1.0 mm
between lateral margins of ocelli. Juga longer than tylus, convergent apically,
their lateral margins above antennifers subparallel. Rostral segments 2-4
about 2.4, 1.9, 1.7 mm long.
Pronotal lobes expanded apically, their anterolateral margins narrowly
reflexed, each lobe projecting laterad of eye by about 0.7-0. 8 width of eye
(Fig. 16); emargination between pronotal lobe and anterolateral margin of
pronotum on each side extending mesad of lateral limit of eye for 0. 3-0.4
width of eye. Anterolateral margins of pronotum caudad of lobes strongly
sinuous. Disk rugose, impressed adjacent to concavity in anterolateral mar-
gins. Width across humeri 5.1 mm, mesial length 2.2 mm.
Scutellum 3.0 mm wide at base, 3.8 mm long. Disk less strongly rugose
than pronotum. Triangular fovea in basal angles black, containing a few
small punctures.
Mesial emargination of pygophore U-shaped, its margins produced an-
teromesad on each side near dorsal limit of emargination (Fig. 1 7). Dorsal
margin on each side of emargination slightly sinuous from caudal view.
Profile of pygophore as in Figure 18.
Length about 9.9 mm.
Distribution. Peru. Known only from holotype.
Holotype. 6, labeled “Piches & Perene Vs, 2000-3000 ft. Peru, Soc. Geog.
de Lima”. Deposited in U.S. National Museum. Type no. 76687. This
specimen lacks both front legs, the left middle leg, right antenna and last
segment of the left antenna.
No paratypes.
Comments. Within the group of species with apically expanded pronotal
lobes, this species, the preceding species (croupius) and the following species
(operosus) are the smallest. Each differs markedly from the others in the
form of the anterolateral pronotal angles.
Lincus operosus, new species
(Figs. 19-21)
Description. Light brown above and below (perhaps somewhat callow)
with red eyes, dark punctures dorsally and on the thoracic venter; mesial
spot at base of scutellum and on each connexival segment, mottling and
interrupted submarginal lateral band on abdominal venter, all yellowish
brown.
Antennal segments 0.6, 0.5, 0.7, 1.1, 1.5 mm long; first segment slightly
VOLUME 91, NUMBER 1
15
Figs. 1 9-2 1 . Linens operosus. 1 9. Head and pronotum. 20. Pygophore, caudal view, proctiger
omitted. 21. Same, profile.
surpassing apex of head. Head width across eyes and length equal, 2.0 mm.
Eyes wider than anterior-posterior diameter, about 0.48 mm wide, 0.4 mm
in diameter. Interocular width of 1 . 1 5 mm greater than distance of 1 .05 mm
between lateral margins of ocelli. Juga slightly longer than tylus, apically
convergent, their lateral margins subparallel above antennifers. Vertex strongly
convex but not tumescent. Rostral segments 2-4 about 2.1, 1.6, 1.6 mm
long.
Pronotal lobes apically expanded and reflexed, each projecting laterad of
eye by about 0.3 width of eye (Fig. 19); emargination between pronotal lobe
and anterolateral pronotal margin on each side extending mesad of lateral
limit of eye by about 0. 1 width of eye. Anterolateral pronotal margins caudad
of lobes moderately sinuous, narrowly and strongly reflexed. Disk rugosely
punctate; punctures deep but not unusually large. Width of pronotum across
humeri 4.6 mm, mesial length 1.9 mm.
Scutellum 2.9 mm wide basally, 3.3 mm long. Disk rugosely punctate
basally; rugosity and puncture size diminishing toward apex. Fovea in basal
angles triangular, darker than adjacent disk, containing a few fine punctures.
Mesial emargination of pygophore U-shaped, its margins without pro-
ductions, sparcely setose (Fig. 20). Dorsal margin on each side of pygophore
convex. Hemispherical impression at bottom of mesial pygophoral emar-
gination clearly visible, prominent. Posterior surface of pygophore nearly
linear in profile (Fig. 21).
Length about 9.4 mm.
Distribution. Venezuela. Known only from type.
Holotype. S, labeled (a) “Venezuela. On orchid” (b) “Insp. House. D.C.
7.14.39”. Deposited in U.S. National Museum. Type no. 76689.
NEW YORK ENTOMOLOGICAL SOCIETY
16
tiger omitted. 24. Same, lateral view.
No paratypes.
Comments. This specimen was intercepted at a plant quarantine station
in a shipment of orchids which may or may not have been the host.
Lincus convexus, new species
(Figs. 22-24)
Description. Dark castaneous to black with many interstitial rugae between
punctures and a small macule on each humerus, at basal angles and middle
of scutellar base, on disk of each corium and on connexival segments yel-
lowish brown. Dorsum unusually convex, dorsal and ventral surfaces sim-
ilarly arched transversely from lateral margins.
Juga longer than tylus, their lateral margins before anteocular concavity
decidely converging toward apex of head (Fig. 22). Vertex normally convex.
Antennal segments 0.8, 0.9, 1.3, 1.6 mm long (5th missing); basal segment
reaching apex of head. Rostral segments 2-4 about 2.5, 2.1, 2.1 mm in
length. Eyes from dorsal view 0.65-0.7 mm wide, their anterior-posterior
diameter 0.55 mm. Interocular width 1.4 mm; distance across ocelli 1.3
mm. Width of head across eyes 2.7 mm, length 2.3 mm.
Pronotal lobes expanded toward apex, horizontal, extending laterad of
eyes by about 0. 1 width of eye. Incision caudad of pronotal lobes extending
mesad to middle of eye. Width of pronotum at humeri 6.3 mm, mesial
length 2.9 mm. Disk rugose.
Scutellum 4.0 mm wide at base, 5.0 mm long. Fovea in basal angles
triangular, black, containing 1 or 2 deep punctures.
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17
Mesial emargination of pygophore U-shaped; low setose projection within
genital cup partially visible where rim of emargination begins dorsally (Fig.
23). Mesial depression at base of inferior ridge small. Proctiger narrowly
rounded apically. Posterior margin of pygophore in profile sigmoid (Fig. 24).
Distribution. Peru. Known only from holotype.
Holotype. S, labeled (a) “Monson Valley, Tingo Maria, XI-2-1954” (b)
“E. I. Schlinger & E. S. Ross, collectors”. Deposited in the American Museum
of Natural History. Fifth antennal segment on right, 4th and 5th on left,
right tarsi on fore and middle legs and tibiae on left fore leg missing.
Linens securiger Breddin
(Figs. 25-29)
Lincus securiger Breddin, 1904, p. 154; Breddin, 1908, pp. 29-31, figs. 9-
1 1; Gaedike, 1971, p. 99 (lectotype designated).
Diagnosis. Pronotal lobe on each side extending horizontally laterad of
eye by about 0.5 width of eye, expanding toward apex; narrow incision
between lobe and anterolateral pronotal margin extending mesad of lateral
limit of eye by about 0.3 width of eye (Fig. 25). Juga exceeding tylus, their
lateral margins tapering sinuously toward apex. Distance between lateral
limits of ocelli subequal to interocular width. Width of each eye about 0.2
greater than anterior-posterior diameter, 0.6-0. 7 of interocular width. First
and 2nd antennal segments subequal in length, 1 st surpassing slightly apex
of head. Basal plates obtusely angled where base of 9th paratergites and 8th
paratergites meet (Fig. 26). Dorsomesial pygophoral margin on each side of
setose border of mesial emargination with small convexity from caudal view
(Fig. 27), this convexity part of tumescence within genital cup on each side
of mesial emargination (Fig. 29). Rim of mesial emargination convex in
profile (Fig. 28).
Length 12.6-12.8 mm.
Distribution. Bolivia, Brazil (Amazonas), Peru.
Comments. The $ lectotype and 9 paralectotype were examined.
Lincus parvulus (Ruckes), New Combination
(Figs. 30-33)
Minilincus parvulus Ruckes, 1958, pp. 15-17, fig. 2.
Diagnosis. Entire vertex of head tumescent. Second antennal segment
subequal to first and third in length. Distance across ocelli from lateral margin
of each slightly greater than interocular distance. Width and length of eyes
from dorsal view subequal. Juga a little longer than tylus, their lateral margins
sigmoid, briefly parallel between concavity above antennifers and apex (Fig.
30). Pronotal lobes small; emargination caudad of lobe reaching mesad about
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NEW YORK ENTOMOLOGICAL SOCIETY
Figs. 25-29. Lincus securiger. 25. Head and pronotum. 26. Pygophore, caudal view, proc-
tiger omitted. 27. Genital plates, caudoventral view; basal plate (BP); 8th paratergite (Pt8); 9th
paratergite (Pt9). 28. Posterior margin of pygophore, lateral view. 29. Genital cup, dorsal view,
setal tufts (S), proctiger (Pr).
as far as lateral limit of eyes. Anterolateral margins of pronotum weakly
sinuous, notably reflexed. Posterior margins of basal plates convex (Fig. 31);
surface of basal plates smooth. Emargination of pygophore sinuously V-
shaped (Fig. 32).
Length about 9 mm.
Distribution. Peru and Brazil (Acre).
Comment. The much swollen vertex and rather small size together are
diagnostic.
The holotype and allotype were examined.
Lincus singularis, new species
(Figs. 34, 35)
Description. Dark brown to fuscous with humeri, 3 basal spots on scutel-
lum and one on disk of each corium, mesial macule on each connexival
VOLUME 91, NUMBER 1
19
Figs. 30-33. Lincus parvulus. 30. Head and pronotum. 31. Genital plates, caudoventral
view; basal plates (BP). 32. Pygophore, caudoventral view; proctiger (Pr). 33. Same, lateral
view.
segment with enlarged continuation on lateral border of corresponding ster-
nite, and vermiform markings on abdominal venter, all yellowish brown.
Vertex of head tumid (Fig. 34). Juga little longer than tylus, their lateral
margins subparallel above antennifers. Antennal segments 0.8, 0.8, 1.1, 1.5,
2.0 mm in length. Distance across ocelli and interocular width equal, 1.25
mm. Eyes from dorsal view as long as wide, 0.6 mm. Width of head across
eyes 2.4 mm, length 2.2 mm. Rostral segments 2-4 about 2.7, 2.2, 2.0 mm
long.
Anterior and posterior margins of pronotal lobes subparallel, each lobe
extending laterad of corresponding eye by 0. 3-0.4 width of eye; emargination
caudad of lobes extending mesad of lateral limit of eye by about 0.2 width
of eye. Anterolateral pronotal margins moderately sinuous. Disk with mod-
erately rugose band between humeri. Pronotal width across humeri 6.3 mm,
length at meson 2.8 mm.
Scutellum 4.0 mm wide at base, 5.2 mm long. Basal angles scarcely foveate.
Lateral angle of basal plates strongly produced, projecting beneath 8th
paratergites (Fig. 35); 9th paratergites reflexed apically.
Length 12.6 mm.
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NEW YORK ENTOMOLOGICAL SOCIETY
Eigs. 34, 35. Linens singularis. 34. Head and pronotum. 35. Genital plates, caudoventral
view; basal plates (BP); 8th paratergites (Pt8).
Distribution. Peru.
Holotype. 9, labeled (a) “Chauchamayo, Peru” (b) ‘‘From F. FI. Rosen-
burg”. Deposited in U.S. National Museum. Type no. 76690.
No paratypes.
Comment. The basal plates of this species are unique within the genus.
Lincus tumidifrons, new species
(Figs. 36-44)
Description. Fuscous above with some of interstitial rugae yellowish brown;
similarly colored macule on humeri, 3 along base of scutellum, marginal
macule in middle of each connexival segment. Rostrum, legs, basal band on
last antennal segment and sternites yellowish brown, latter with fuscous
punctures; remainder of venter fuscous.
Entire vertex strongly convex (Fig. 36). Antennal segment 2 short, usually
about two-thirds length of 1 and 3, latter 2 segments subequal in length;
length of segments 0. 8-0.9, 0. 5-0.6, 0.8-0. 9, 2.0-2. 3, 2. 6-2. 8. Distance across
ocelli from lateral margin of each subequal to interocular width, 1.25-1.4
mm. Width and length of eye from above subequal, 0.65 mm. Width of
head across eyes 2. 3-2. 7 mm length 2. 3-2.6 mm. Juga little longer than
tylus, their lateral margins sigmoid.
Pronotal lobes extending little if at all laterad of eyes, narrowly rounded
apically. Disk rugose. Anterolateral margins weakly sinuous. Width across
humeri 5. 7-6. 3 mm, mesial length 2. 3-2. 8 mm.
Scutellum shallowly foveate in each basal angle. Disk similar to pronotum
in rugosity. Basal width 3.5-4. 1 mm, length 4. 1-5.0 mm.
VOLUME 91, NUMBER 1
21
Figs. 36-44. Lincus tumidifrons. 36. Head and pronotum. 37. Genital plates, caudoventral
view; basal plates (BP). 38. Paramere. 39. Proctiger, dorsal view. 40. Proctiger, lateral view,
apex to right. 41. Pygophore, caudal view, proctiger omitted. 42. Aedeagus, basal plate omitted.
43. Distal parts of spermatheca (bulb stippled). 44. Spermathecal bulb, terminal view.
Posterior margin of basal plates with pronounced submarginal impression
(Fig. 37).
Mesial emargination of pygophore broad, V-shaped (Fig. 41).
Length about 1 1-13.7 mm.
Distribution. Panama, Trinidad, Venezuela.
Holotype. 9, labeled “Las Cumbres, Panama, 9°06’ N, 79°32’ W, Lt. trap.
2-V-75. Col: Henk Wolda”. Deposited in U.S. National Museum. Type no.
76686.
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NEW YORK ENTOMOLOGICAL SOCIETY
Paratypes. 1099, 6dd. Same data as holotype except date “16-VI-76” (9
HDE), “1 l-VII-76” (9 LHR) and “7-X-75” (9 LHR) “Barro Colorado-C.
Z., Lights, Weir/SM-I, 7-XII-1977, Coll. H. Wolda” (5 HDE); same data
except ‘‘Weir SM-I-III, 3-9-IX-1975” (<5 LHR); “Barro Colorado IsL, Canal
Zone— Lt. trap, 9- VI-76, Col: D. Engleman” (9 HDE); same data except
“23-VI-77, Wolda” (9 HDE); (a) “Panama Canal Zone, Barro Colorado Is.,
9°10’ N, 79°50’ W” (b) “8-VII-1977, H. A. Hespenheide” (9 HDE); “Panama,
dist. Chepo, Altoa de Maje, 17 May 75 at lights, Stockwell-Engleman” (9
HDE); “Panama; Pma Pr., Altos (Isla) de Maje, 9°08’ N, 78°49’ W, 15 May
’76, Stockwell” (S HDE); “Barro Colorado IsL, Canal Zone— Lt. trap 8-IX-
76. Col: D. Engleman” (9 UNAM); “Cabima Pan., May 17-11, August
Busck” (S UNAM); same data except “May 20.1 1” (5 USNM); (a) “Vene-
zuela, on orchid” (b) “Inspect. H., D. C. 6.27.40” (9 USNM); (a) “Venezuela,
on orchid packing” (b) “Inspect. H., Wash. D. C. VII-11-35” (9 USNM);
“Trinida: W.I. Simla, Arima Valley. 11-18-1966. B. Heineman” (S LHR).
Comment. A female from Peru that is probably this species is omitted
from the type series because its identity is uncertain. The juga of this spec-
imen are nearly contiguous apically and the basal plates are less impressed
than in females of the type series.
Lincus anulatus, new species
(Figs. 45-48)
Description. Mostly dark to fuscous, occasional yellowish clay colored with
fuscous punctation; sparsely punctate transverse fascia connecting humeri
irregular, of varying size, contrasting yellowish brown in dark specimens as
is macule on disk of each corium, marginal macule in middle of connexival
segments, mesial spot at base of scutellum and sometimes at basal angles,
small scattered patches on venter, macule on lateral margins of each sternite
corresponding with macule on connexival segment, rostrum and legs ex-
cepting small spots and bands: broad preapical femoral band, one of com-
parable width at apex of tibiae and broader band near base of tibiae all
fuscous; tibial bands occasionally obscure; humeri, connexival macules and
apex of femora sometimes rufous.
Antennal segments 0.5-0. 7, 0.6-0. 8, 0.7-0. 9, 1.0-1. 5, 1.4-1. 9 mm in
length; basal segment nearly reaching to slightly exceeding apex of head.
Rostral segments 2-4 about 1.8-2. 6, 1.6-2. 3, 1.5-2. 2 mm in length; apex
reaching middle of penultimate sternite to apex of abdomen. Eyes as wide
as anterior-posterior diameter from dorsal view, width of each half or some-
what less of interocular distance. Interocular distance slightly greater (usually
0.5 mm) than distance between lateral limits of ocelli. Juga a little longer
than tylus, their lateral margins parallel between concavity above antennifers
VOLUME 91, NUMBER 1
23
Figs. 45-48. Lincus anulatus. 45. Head and pronotum. 46. Genital plates, caudoventral
view; basal plates (BP). 47. Pygophore, caudal view; proctiger (Pr). 48. Same, lateral view;
proctiger (Pr), impression (imp).
and convexity at apices (Fig. 45). Width of head across eyes 2.0-2. 1 mm,
length 1. 7-2.0 mm.
Pronotal lobe small, subacute, directed anterolaterad, usually projecting
laterad of eye by distance equal to 0.2-0. 4 width of an eye, rarely shorter;
emargination caudad of lobe varying in depth, reaching lateral limit of eye
or failing to do so by distance equal to 0.2 width of an eye. Anterolateral
margins of pronotum usually slightly sinuous, sometimes markedly so (5),
occasionally straight or slightly concave. Pronotal width at humeri 4. 6-5. 3
mm, mesial length 1.9-2. 2 mm.
Scutellum 2. 7-3. 3 mm wide at base, 3. 3-4.0 mm long. Fovea in basal
angles inconspicuous.
Posterior margin of basal plates obtusely angulate (Fig. 46). Ninth para-
tergites suboval.
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NEW YORK ENTOMOLOGICAL SOCIETY
Rim of pygophoral emargination produced posteriorly on each side just
dorsad of mesial circular impression at bottom of emargination (Figs. 47,
48).
Length 9.2-10.3 mm.
Distribution. Brazil (Amapa, Amazonas, Mato Grosso, Para) and Panama.
Holotype. 2, labeled “Brazil, Mato Grosso: Lat. 12° 13’ and Long. 55°37’
Sinop, October 1974. M. Alvarenga”. Deposited in the American Museum
of Natural History. The left antennae is aberrant, having 4 segments.
Paratypes. 15, 322. “Panama, Colon Prov. Sta. Rita Ridge, 18-26 VI 76
at lights, Engleman-Thurman” (2 LHR); “Benjamin Constant, Amazonas,
Brasil, 18-28-IX-1962, K. Lenko — col.” (255 MZSP); “Sta. Isabel do Para,
PA Brasil, 30 III 1962, J. Bechyne col.” (2 MZSP), (a) “Jei Amapa, Serra
Novio, X Bicelli col.” (b) “— arape Sucuriji” (Hyphen indicating illegible
letter) (c) “Colegao J. Lane” (5 NZSP).
Comments. The annulated femora and tibiae are diagnostic, although faint
or even obscured in discolored specimens.
There appears to be sexual dimorphism with respect to the anterolateral
margin of the pronotum. In the males this margin is clearly sinuous; in the
females it is at most weakly sinuous. The pronotal lobe is more weakly
developed in males than in females.
Lincus repizcus, new species
(Figs. 49-52)
Description. Dorsum dark brown becoming fuscous on anterior disk of
pronotum, head and connexiva, mottled with yellowish brown rugae and
small macules of same color: one on each humerus, mesially at base of
scutellum, on disk of each corium and on margin in middle of each con-
nexival segment. Antennae dark brown excepting brownish yellow mesial
surface of first segment and basal third of last segment. Rostrum and legs
brownish yellow excepting superior surface of tibia, broad diffuse incomplete
subbasal tibial band and similar subapical femoral band darker. Venter
yellowish brown with punctures and surrounding area brown to fuscous;
head, evaporative areas and plura above coxae fuscous; mesial impunctate
abdominal vitta brown; yellow macule on lateral margins of each sternite
includes lateral trichobothrium.
Head 2. 3-2.4 mm wide across eyes, 2.0-2. 1 mm long. Antennal segments
0.7-0. 8, 0.8, 0.9, 1.4, 1. 6-2.0 mm in length; first segment clearly surpasses
apex of head (Fig. 49). Eyes from dorsal view 0.55 mm wide, anterior-
posterior diameter 0. 5-0.6 mm. Interocular width 1.3 mm; distance across
ocelli 1 .2 mm. Greatest diameter of each ocellus 0.25 mm. Juga a little longer
than tylus, their lateral margins tapering sinuously toward apex. Base of tylus
VOLUME 91, NUMBER 1
25
Figs. 49-52. Lincus repizcus. 49. Head and pronotum. 50. Genital plates, caudoventral
view. 51. Pygophore, caudal view. 52. Same, lateral view.
slightly tumescent. Vertex of head convex, not tumescent, its lateral margins
delineated on each side by densely punctate band of punctures that envelopes
ocellus. Ventrally on each side a narrow but markedly elevated brown ridge
runs from base of head to base of antennifer at mesial limit of eye. Rostral
segments 2-4 about 2. 2-2. 6, 1. 8-2.0, 1.7-1. 9 mm in length.
Pronotal lobes nearly parallel-sided, extending laterad of eye by 0.2-0. 3
width of eye; emargination caudad of lobes reaching mesad of lateral limit
of eye by about 0.1 width of eye. Width of pronotum across humeri 5.5-
5.7 mm, mesial length 2.4-2. 5 mm; anterolateral margins slightly sinuous;
disk rugosely punctate.
Scutellum 3. 4-3. 5 mm wide at base, 4. 3-4. 5 mm long. Fovea in each
basal angle black, triangular, with one large and a few small punctures.
Punctures in basal disk forming transverse vermiform rows separated by
ruga.
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NEW YORK ENTOMOLOGICAL SOCIETY
Greatest width and greatest length of basal plates subequal, about 1 mm
(Fig. 50).
Pygophoral emargination U-shaped, its margins entire (Fig. 51). Posterior
pygophoral margin convex in profile (Fig. 52).
Length about 1 1 mm.
Distribution. Peru.
Holotype. 6, labeled (a) “Valle Chanchamayo (Peru) 800 M, 13-VIII-195 1,
Leg. Weyrauch” (5 1 of the year written over other numbers) (b) “EKW4632”
(c) “ex col. Weyrauch” (d) “coleccion Fundacion M. Lillo, 4000 S. M. Tu-
cuman, Tucuman, Argentina”. Deposited in Fundacion Miguel Lillo, Tu-
cuman, Argentina.
Paratype. 9, labeled (a) “Pucallapa, Peru. 6-6-1956. Jose M. Schunke,
Coll.” (b) “J. C. Lutz Collection, 1961” (USNM).
Lincus discessus Distant
(Figs. 53-56)
Lincus discessus Distant, 1900, pp. 688-689.
Diagnosis. Pronotal lobe on each side extending laterad of eye by less than
0.2 width of eye, curving to rounded apex. Incision between lobes and
anterolateral pronotal margins shallow, scarcely extending mesad as far as
lateral limit of corresponding eye. Antennal segments 2 and 3 subequal in
length. Juga projecting beyond tylus but not markedly convergent (Fig. 53).
Interocular width and distance across ocelli subequal; width of each eye
about 0.42 of interocular width. Posterior margin of each basal plate un-
evenly convex, bending most at posterolateral angle of 9th paratergites; latter
briefly contiguous mesially (Fig. 54). Pygophore without projections ex-
tending into mesial emargination from border, contour of emargination
interrupted only at dorsal corners by sulcus formed by diagonal lamina entad
of posterior pygophoral surface (Fig. 55). Posterior pygophoral surface con-
cave in profile (Fig. 56).
Length 9.3-1 1.5 mm.
Distribution. Costa Rica, Guyana.
Comment. The holotype, a female was examined.
Lincus varius, new species
(Figs. 57-59)
Description. Dorsum dark brown to fuscous with small brownish yellow
macule on each humerus, on disk of each corium, mesially at base of scu-
tellum and marginally in middle of each connexival segment; some rugae
of pronotum and scutellum and all appendages brownish yellow, the 3 basal
antennal segments darker than last 2 segments. Venter fuscous with some
VOLUME 91, NUMBER 1
27
Figs. 53-56. Linens discessus. 53. Head and pronotum, antennifers omitted. 54. Genital
plates, caudo ventral view; basal plates (BP); 9th paratergites (Pt9). 55. Pygophore, caudal view,
proctiger omitted. 56. Same, lateral view.
rugae and broad somewhat discontinuous callus along lateral margins of
abdomen brownish yellow; calli extend mesad to include mesial tricho-
bothrius; punctation inconspicuous.
Head 2.3 mm wide across eyes, 2.2 mm long. Antennal segments 0.7, 0.7,
1.1, 1.4, 2.0 mm in length; first segment reaching apex of head. Eyes from
dorsal view 0.45-0.5 mm wide, anterior-posterior diameter 0.6 mm, their
width 0.35-0.40 of interocular width. Distance across ocelli from lateral
limit of one to lateral limit of other 1 .2 mm; greatest diameter of each ocellus
0.25 mm. Juga slightly longer than tylus, their lateral margins subparallel
between concavity above antennifers and apex of head (Fig. 57). Tylus tu-
mescent at base. Vertex of head convex but not tumescent, its lateral margins
defined on each side by densely punctate band which envelopes ocellus.
Ventrally on each side a slightly elevated narrow ridge runs from base of
head to base of antennifer at mesial limit of eye, this ridge contrastingly
brownish yellow on fuscous surface. Rostral segments 2-4 about 2.8, 2.1,
2.0 mm long.
Pronotal lobes subtriangular, extending laterad of eye by 0.2-0. 3 width of
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NEW YORK ENTOMOLOGICAL SOCIETY
Figs. 57-59. L. varius. 57. Head and pronotum. 58. Pygophore, caudal view; proctiger (Pr).
59. Same, lateral view.
eye; emargination caudad of lobes reaching mesad as far as lateral limit of
eye or nearly so. Width of pronotum across humeri 6.4 mm, mesial length
2.3 mm; anterolateral margins slightly sinuous, disk somewhat rugosely
punctate with weak ruga on meson.
Scutellum 3.4 mm wide at base, 4.4 mm long. Triangular black fovea in
basal angles with few punctures. Most punctures in scutellum arranged in
transverse vermiform rows.
Pygophoral emargination a bowed V from caudal view (Fig. 58). Posterior
margin of pygophore sinuously linear from lateral view (Fig. 59).
Length about 1 1.5 mm.
Distribution. Peru.
Holotype. 6, labeled (a) ‘‘Satipo, Peru. III. 10. 1941. P. Paprzycki” (b) “J.
C. Lutz Collection. 1961” Deposited in U.S. National Museum. Type no.
76691.
No paratypes.
Comments. This is one of 7 species with relatively small eyes. The pale,
calloused band along the lateral margins of the abdominal venter and the
parallel sided juga is a combination that distinguishes the species.
Lincus modicus, new species
(Figs. 60-63)
Description. Fuscous with castaneous or ochraceous macule on each hu-
merus, mesially at base of scutellum and on disk of each corium. Basal band
on last antennal segment and tarsi ochraceous. Rostrum brown.
Antennal segments 0.6-0. 7, 0.8, 0. 9-1.0, 1.2, 1.6 mm in length. Head
little wider than long, 2. 1 mm across eyes, 2.0 mm long. Width and anterior-
posterior diameter of eyes subequal, about 0.5 mm, less than half interocular
width. Interocular width 1.1-1.15 mm, slightly greater than distance across
VOLUME 91, NUMBER 1
29
Figs. 60-63. Lincus modicus. 60. Head and pronotum. 6 1 . Pygophore, caudal view, proctiger
omitted. 62. Pygophore, lateral view, proctiger omitted. 63. Genital plates, caudoventral view;
basal plates (BP); lateral angle (l.a.); mesial angle (m.a.).
ocelli from lateral margin of one to lateral margin of other, 1.05 mm. Juga
longer than tylus, apically convergent (Fig. 60).
Pronotal lobe on each side extending laterad about as far as lateral limit
of eye, with anteromesial and posterolateral margins tapering to narrowly
rounded apex, slightly reflexed apically. Incision between lobe and antero-
lateral margin of pronotum not extending mesad as far as middle of eye.
Pronotal disk strongly punctate, rugosely so laterally. Width across humeri
5. 2-5. 4 mm, mesial length 2.2-2. 3 mm.
Scutellum 3. 2-3. 3 mm wide at base, 3. 9-4.0 mm long. Large black fovea
in each basal angle containing several small punctures. Disk rugosely punc-
tate basally. Connexiva entirely fuscous.
Posterior margin of basal plates broadly rounded mesially, otherwise sub-
truncate (Fig. 63). Greatest width of each plate (1.0 mm) exceeding greatest
length (0.9 mm).
Large projection at dorsal corners of broadly U-shaped emargination in
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pygophore directed cephalad and dorsomesad (Fig. 61). From lateral view
these projections visible above rim of genital cup (Fig. 62).
Length about 10.7-11 mm.
Distribution. Ecuador.
Holotype. 9, labeled “Ecuador, Pastaza: Cuisimi, on Rio Cuisimi, 150 km
SE Puyo, 350 m, May 15-31, 1971. B. Malkin”. Deposited in the American
Museum of Natural History.
Paratypes. Same labeling as holotype {1266, 599 AMNH; 266, 299 LHR).
Lincus leviventris, new species
(Figs. 64, 65)
Description. Dark brown, appendages for most part lighter, with yellowish
brown macule on each humerus, mesially at base of scutellum, on disk of
each corium and marginally at middle of connexival segments.
Head 2.5 mm wide across eyes, 2.4 mm long. Antennal segments 0.8, 0.8,
1.1, 1.6 mm long (last segment missing); first segment reaching apex of head.
Width and anterior-posterior diameter of eyes from dorsal view equal, about
0.55 mm and about 0.45 of interocular width. Interocular width 1.4 mm;
distance across ocelli about 0. 1 less than interocular width. Juga longer than
tylus, their lateral margins subparallel between concavity above antennifers
and apex of head (Fig. 64). Rostral segments 2-4 about 2.9, 2.1, 2.1 mm
long.
Pronotal lobes extending laterad of eye by 0. 1-0.3 width of an eye, tapering
to obtuse apex; emargination caudad of lobe extending about as far mesad
as lateral limit of eye. Width of pronotum across humeri 5.9 mm, mesial
length 2.5 mm; anterolateral margins sinuous; disk rugosely punctate.
Scutellum 3.6 mm wide at base, 4.4 mm long. Triangular black fovea in
basal angles containing a few strong punctures. Basal disk rugosely punctate.
Abdominal venter inconspicuously punctate, smooth excepting some
wrinkles along sutures.
Greatest length of each basal plate 1.1 mm, greatest width 1.0 mm; pos-
terior margin evenly convex; lateral angle anterior to mesial angle formed
by junction of basal plates when viewed caudoventrally (Fig. 65). Ninth
paratergites reflexed as narrow perpendicular ridge where they meet; surface
convex with broad longitudinal ridge nearer mesial than lateral margins (Fig.
65).
Length about 12 mm.
Distribution. Peru (Cuzco). Known only from holotype.
Holotype. 9, labeled (a) “Pilcopata” (2nd and 3rd letters uncertain) “Cuzco.
9-8-64” (b) “trampa de luz” (c) “CUZCO” (d) “1592”. Deposited in Museu
Nacional, Rio de Janeiro, Brazil. No paratypes.
Comments. This species is one of 7 whose interocular width is more than
VOLUME 91, NUMBER 1
31
Figs. 64, 65. Lincus leviventris. 64. Head and pronotum. 65. Genital plates, caudoventral
view; basal plates (BP); 9th paratergites (Pt9).
twice the width of an eye, i.e., it belongs in the “small eyed” group of species.
Within this group it is one of 5 species bearing on each side a low ridge on
the ventral surface of the head behind the eyes, and among these 5 only this
species and L. modicus have the abdominal venter almost uniformly dark.
The genital plates of these 2 species differ.
Lincus manchus, new species
(Figs. 66-68)
Description. Yellowish brown with darker punctures; humeri, narrow dif-
fuse border on anterolateral pronotal margins, basal third of coria, apex of
scutellum, lateral ventral margins of thorax and abdomen, all rufous; large
discal macule on each corium, basal 0.4 of last antennal segment, tarsi and
abdominal venter (excepting last sternite) between spiracular lines yellowish
or brownish yellow, latter with brown punctures on each side of wide im-
punctate mesial area.
Head 2.0 mm wide across eyes, 1.8 mm long. Antennal segments 0.7, 0.6,
1.0, 1.2, 1.7 mm long; first segment reaching apex of head. Width and
anterior-posterior diameter of eyes from dorsal view about 0.45 mm; width
of an eye 0.4-0.45 of interocular width. Interocular width 1.1 mm, distance
across ocelli subequal; ocelli relatively large, greatest diameter of each 0.35
of distance between them. Juga slightly longer than tylus, their lateral margins
subparallel between concavity above antennifers and apex of head (Fig. 66).
Vertex flat, delimited on each side by densely punctate depressed band that
basally flows around ocellus. Rostral segments 2-4 about 2.1, 1.9, 1.8 mm
in length.
Pronotal lobes extending laterad of eye by 0.4-0. 5 width of eye, sides
32
NEW YORK ENTOMOLOGICAL SOCIETY
Figs. 66-68. L. manchus. 66. Head and pronotum. 67. Pygophore caudal view, proctiger
omitted. 68. Same slightly exserted, lateral view.
subparallel, narrowly rounded at apex; emargination caudad of lobes reach-
ing mesad nearly to lateral limit of eyes. Width of pronotum across humeri
4.9 mm, mesial length 2.1 mm. Anterolateral margins sinuous, notably
reflexed. Coarsely punctate disk with narrow irregular callus along meson.
Scutellum 3.0 mm wide at base, 3.7 mm long. Pale mesial line on basal
disk. Punctation similar to that on pronotum.
Mesial emargination of pygophore U-shaped from caudal view; area around
circular impression at base of emargination slightly raised (Fig. 67). Profile
of pygophore nearly linear (Fig. 68).
Distribution. Bolivia. Known only from holotype.
Ho/otype. S, labeled (a) “Tumupasa, Boliv. Dec. W. M. Mann.” (b) “Mul-
ford Biological Exploration. 1921-1922.” Deposited in the U.S. National
Museum. Type no. 76693.
Comment. The small eyes, absence of a pair of ridges on the venter of the
head between the base of the head and base of the antennifers, and large
ocelli are a diagnostic combination.
The holotype may be somewhat callow and as a consequence unusually
light in color.
Linens armiger Breddin
(Figs. 69, 70)
Linens armiger Breddin, 1908, pp. 26-21 , fig. 4.
Diagnosis. Eyes relatively small, width of one eye about 0.4 of interocular
width. Juga as long as tylus, their lateral margins somewhat constricted above
antennifers (Fig. 69). Vertex evenly convex between ocelli; surface of juga
shallowly excavated; base of eyes from ocelli to ommatidia somewhat pol-
ished; eyes not elevated above adjacent part of head; sides of head laterad
of concavity in lateral margin of juga sloping to antennifers and visible from
VOLUME 91, NUMBER 1
33
Figs. 69, 70. Uncus armiger. 69. Head and pronotum. 70. Genital plates, caudoventral
view; basal plates (BP).
above. Anterolateral margins of pronotum sinuous; pronotal lobes extending
laterad of eyes by about one-half width of eye, triangular, narrowly rounded
apically; emargination behind lobes extending no farther mesad than lateral
limit of eyes. Basal plates with depressed area along posterolateral margins
(Fig. 70); second gonocoxae convex. Length 10 mm.
Distribution. Bolivia. Known only from holotype.
Comments. The small eyes together with the form of the head distinguishes
this species.
The female holotype, in the Naturhistorisches Museum Wien, was ex-
amined.
Lincus breddini, new species
(Figs. 71-74)
Description. Mostly fuscous with ochraceous macule on each humeral
angle, mesially at base of scutellum and on disk of each corium. Last 2
segments of antennae, rostrum and tarsi also ochraceous.
Antennal segments 0. 7-1.0, 1.0-1. 4, 1.0-1. 5, 1.4-1. 8, 1.9-2. 3 mm in
length. Rostral segments 2-4 about 2.9, 2.2, 2.1 mm in length. Head 2.6-
2.9 mm wide across eyes 2. 3-2. 6 mm long. Eyes wider by about 0.05 mm
than their anterior-posterior diameter. Distance across ocelli from lateral
margin of one to lateral margin of other usually slightly greater (0. 1 mm or
less) than interocular width. Width of eye (0.75-0.9 mm) more than half of
interocular width (1.1 5-1 .3 mm). Juga longer than tylus, apically convergent
and occasionally contiguous (Fig. 71).
Pronotal lobes extending laterad approximately to lateral limit of eyes.
34
NEW YORK ENTOMOLOGICAL SOCIETY
Eigs. 71-74. Lincus breddini. 71. Head and pronotum. 72. Genital plates, caudoventral
view; basal plates (BP). 73. Pygophore, caudal view, proctiger omitted. 74. Same, lateral view.
longer than wide, usually somewhat reflexed but not expanded apically, their
anteromesial and posterolateral margins subparallel. Incision caudad of lobes
extending mesad at least as far as middle of eye. Pronotal disk rugose. Width
across humeri 6. 2-7. 4 mm, mesial length 2. 3-2.6 mm.
Basal width of scutellum 3. 8-4. 5 mm, length 4. 7-5. 6 mm. Large black
fovea in each basal angle containing several small punctures. Disk rugose
except apically. Apex slightly reflexed on each side.
Posterior margin of basal plates truncately rounded (Fig. 72).
Mesial emargination of pygophore from caudal view with large projection
near dorsal margin directed obliquely dorsomesad and somewhat cephalad
(Fig. 72). Profile of pygophore irregular (Fig. 74).
Length about 12.2-14.3 mm.
Distribution. Surinam.
Holotype. 2, labeled “Museum Leiden. Suriname. Mapane area, Blakka
Watra, 26. V. 1960. Ph. v. Doesburg, Jr.”. Deposited in the Rijksmuseum
van Natuurlijke Historic. The left antenna is deformed, there being only 3
segments, with the basal 2 segments formed normally.
VOLUME 91, NUMBER 1
35
Paratypes. 4$9, 86(3. Same data as holotype (6, 9 LHR; 6, 329 RNH); hand-
written label “Brownsweg, 8 juni ’61”, remainder uncertain (9 RNH); hand-
written label “Suriname 28. VI. 1938”, remainder uncertain (9 RNH); same
labeling except date “16. VIII. 1938” (6 RNH) and “13. X. 1938” (6 RNH);
“Museum Leiden. Suriname exp. 1948-1949” with additional handwritten
data of which only “14. x. ’48” is clearly decipherable (6 RNH); “Suriname,
Faglust op Erthyrina, 28. VI. 1938, Ge[— ]les”.
Comments. The species is named for Gustav Breddin who, among his
numerous contributions to pentatomid taxonomy, described the majority
of species previously recognized in the genus Uncus.
Uncus lamelliger Breddin
(Figs. 75-78)
Uncus lamelliger Brtddm, 1908, pp. 34-35, figs. 19, 20.
Diagnosis. Pronotal lobe on each side extending laterad of eye nearly 0.4
width of eye, about as long as wide at base, broadly rounded apically (Fig.
75); anterolateral pronotal margins somewhat sinuous. Width of each eye
about 0.6 of interocular width. Juga exceeding tylus, their lateral margins
briefly subparallel anterior to antennifers. First antennal segment about 0.6
length of second. Posterior margin of basal plates broadly rounded (Fig. 78).
Dorsal margin of pygophore from caudal view shallowly notched adjacent
to large mesial emargination (Fig. 76); profile of pygophore concave (Fig.
77).
Length about 10.2-10.8 mm.
Distribution. Colombia, French Guiana, Surinam.
Types. Of 2 syntypes one remains in the Naturhistorisches Museum Wien.
This female labeled “Colomb. Sign.”, is designated lectotype.
Comment. The basal width and length of the pronotal lobes are subequal
in this species; this is not true for other species of the “big eyed” group.
Uncus lobuliger Breddin
(Figs. 79-82)
Uncus lobuliger Breddin, 1908, pp. 27-29, figs. 5-8.
Diagnosis. Pronotal lobes variable in length, extending laterad nearly to
lateral limit of corresponding eye, or longer and projecting laterad of eye as
much as 0.4 width of eye (Fig. 79); incision caudad of lobes extending mesad
of lateral limit of corresponding eye by about 0.2 width of eye; anterolateral
pronotal margins sinuous. Width of each eye about 0.6 of interocular width.
Juga exceeding tylus, their lateral margins anterior to anteocular concavity
tapering slightly to apex. First antennal segment about 0.8 as long as second.
Genital plates as in Figure 82. Dorsal margin of pygophore from caudal view
36
NEW YORK ENTOMOLOGICAL SOCIETY
Figs. 75-78. Lincus lamelliger. 75. Head and pronotum. 76. Pygophore, caudal view, proc-
tiger omitted. 77. Same, lateral view. 78. Genital plates, caudo ventral view; basal plates (BP).
bearing on each side a small projection at edge of setose area along mesial
emargination (Fig. 80); posterior pygophoral margin in profile strongly con-
vex subdorsally (Fig. 81).
Length about 9.4-12.0 mm.
Distribution. Brazil (Bahia, Espirto Santo, Rio de Janeiro).
Types. Of the syntype series of 2 females and 1 male, only the male was
located and examined. This specimen, in the Naturhistorisches Museum
Wien, is designated lectotype.
Lincus rufospilotus (Westwood)
(Figs. 83-86)
Pentatoma rufospilota Westwood, 1837, p. 44.
Ochlerus rufospilota: Dallas, 1851, p. 157.
VOLUME 91, NUMBER 1
37
Figs. 79-82. Lincus lobuliger. 79. Head and pronotum. 80. Pygophore, caudal view, proc-
tiger omitted. 81. Same, lateral view. 82. Genital plates, caudoventral view.
Ochlerus villis Walker, 1867, p. 196 (synonymized by Distant, 1899).
Lincus rufospilotus: StM, 1867, p. 254; Distant, 1899, p. 423.
Lincus erythrospilus Stal, 1872, p. 14 (replacement name).
Lincus cultiger Breddin, 1908, pp. 35-36, figs. 21-23. New Synonymy.
Diagnosis. Pronotal lobe on each side extending no further laterad of eye
than 0.2 width of eye, narrowly round and often reflexed apically (Fig. 83).
Incision between posterior margin of lobes and anterolateral pronotal mar-
gins not reaching as far mesad as middle of eyes. Distance across ocelli
slightly greater than interocular width. Width of each eye about 0. 1 greater
than one-half of interocular width, slightly greater than anterior-posterior
diameter of eye from dorsal view. Antennal segments 2 and 3 subequal in
length; segments 4 and 5 lighter in color than basal 3 segments, neither
annulated. Juga projecting beyond tylus, their lateral margins parallel be-
tween anteocular concavity and apex. Posterior margin of basal plates broad-
ly rounded; greatest width of each plate about 0.8 of greatest length (Fig.
84). Large mesial emargination in pygophore from caudal view U-shaped
below tubercles protruding into emargination at its dorsal limit; rim of
emargination continuing for short distance ventrad of emargination (Fig.
38
NEW YORK ENTOMOLOGICAL SOCIETY
Figs. 83-86. Linens rufospilotus. 83. Head and pronotum. 84. Genital plates, caudoventral
view; basal plates (BP); 9th paratergites (Pt9); 2nd gonocoxae (Gx2). 85. Pygophore, caudal
view, proctiger omitted. 86. Same, lateral view.
85); dorsal pygophoral margin on each side of emargination slanting evenly
dorsolaterad to rounded dorsolateral corners. Posterior surface of pygophore
sinuously linear in profile (Fig. 86).
Large species, 1 1.5-15 mm in length.
Distribution. Northern Brazil, Colombia, Guyana, Peru and Surinam.
Comments. The holotypes of Pentatoma rufospilata, Ochlerus vi/is and
Lincus cultiger were examined.
Stal (1872) gave no reason for renaming the species but he presumably
believed that Pentatoma rufospilota was preoccupied.
Lincus vallis, new species
(Figs. 87-89)
Description. Dark castaneous to fuscous with ochraceous macule on each
humeral angle, mesially at base of scutellum and on disk of each corium.
Basal fourth of last antennal segment and mottling on venter also ochraceous.
VOLUME 91, NUMBER 1
39
Antennal segments 0.7, 0.8, 1.3, 2.1, 2.0 mm long. Head 2.4 mm wide
across eyes, 2.1 mm long. Eyes slightly wider (by 0.05-0.1 mm) than an-
terior-posterior diameter, the width of each equal to or slightly greater than
half interocular distance. Distance across ocelli 1.1 mm, 0.1 mm less than
interocular width. Juga longer than tylus, their lateral margins converging
sinuously toward apex (Fig. 87). Rostral segments 2-4 about 2.5, 2.2, 2.0
mm long.
Pronotal lobes extending laterad of eyes by about 0.2 width of eye, about
twice as long from base of incision to apex as wide, parallel sided or slightly
expanded subapically; incision behind lobes reaching mesad well past lateral
limit of corresponding eye but not reaching middle of eye. Disk rugose.
Anterolateral margins sinuous. Width at humeri 5.9 mm, mesial length 2.5
mm.
Scutellum 3.7 mm wide at base, 4.4 mm long. Fovea in each basal angle
triangular, black. Disk rugose except apically.
Fength about 11.7 mm.
Mesial emargination of pygophore from caudal view U-shaped, with setose
projection directed mesad from dorsal limit of emargination (Fig. 88). Dorsal
pygophoral margin on each side of emargination sloping dorsolaterad. Rim
of emargination disappearing at level with bottom of emargination. Posterior
surface of pygophore sinuously linear in prohle (Fig. 89).
Distribution. Peru. Known only from holotype.
Holotype. S, labeled “Tingo Maria, Huan. Peru. Nov. 21, 1946. Alt. 670
m.” Deposited in the U.S. National Museum. Type no. 76695.
Comments. This species is similar to L. rufospilotus but differs in having
antennal segment 3 about 1.5 times as long as segment 2, the basal fourth
of the last antennal segment pale, the distance across the ocelli less than the
interocular width, and the produced rim along the mesial pygophoral emar-
gination disappearing at a level even with the bottom of the emargination.
Lincus dentiger Breddin
(Figs. 90-93)
Lincus dentiger Breddin, 1904, p. 154; Breddin, 1908, pp. 24-26, hgs. 1, 2
(redescription); Gaedike, 1971, p. 84 (type).
Diagnosis. Pronotal lobes on each side extending laterad of eye by less
than 0.2 width of eye, tapering to narrowly rounded apex. Incision between
lobe and anterolateral pronotal margin of pronotum extending mesad about
as far as lateral limit of eye. Antennal segments 2 and 3 subequal in length.
Juga not or scarcely exceeding tylus (Fig. 90). Interocular width and distance
across ocelli subequal, about twice width of one eye. Posterior margin of
basal plates broadly and evenly rounded. Each 9th paratergite subtriangular
(Fig. 93). Emargination in pygophore from caudal view constricted midway
40
NEW YORK ENTOMOLOGICAL SOCIETY
Figs. 87-89. Linens vallis. 87. Head and pronotum. 88. Pygophore, proctiger omitted, caudal
view. 89. Same, lateral view.
in dorsoventral direction, cordiform ventrad of constriction, widening uni-
formly dorsad of constriction (Fig. 91); pygophoral profile as in Figure 92.
Length about 9.5-10 mm.
Distribution. Ecuador, Surinam.
Comments. The type, a female from Ecuador, was examined. Both sexes
were seen from Surinam.
Linens styliger Breddin
(Figs. 94-99)
Linens styliger 1908, p. 31, figs. 12, 13.
Diagnosis. Pronotal lobe on each side extending laterad of eye by nearly
one-half width of eye; length of each lobe from base of emargination almost
twice width of lobe at this point; apex narrowly rounded (Figs. 94, 95);
anterolateral margins of pronotum sinuous. Width of each eye about 0.6 of
interocular width. Juga exceeding tylus slightly, their lateral margins con-
verging slightly toward apex. First antennal segment about 0.8 length of
second. Basal plates subtriangular with uniformly convex posterolateral mar-
gin (Fig. 96). Mesial emargination of pygophore V-shaped; dorsal margin of
pygophore from caudal view notched about midway between mesial emar-
gination and dorsolateral corners (Fig. 97), this notch resulting from inflec-
tion of posterior rim of genital cup on each side into a small dorsolateral
lobe and large plate mesoventrad of this lobe (Fig. 99). Posterior pygophoral
surface convex in profile (Fig. 98).
Length 11.5-12.5 mm.
Distribution. Colombia, Peru.
Types. The 2 syntypes were not located. Of a pair of specimens in the
VOLUME 91, NUMBER 1
41
Figs. 90-93. Lincus dentiger. 90. Head and pronotum. 9 1 . Pygophore, caudal view, proctiger
omitted. 92. Same, lateral view. 93. Genital plates, caudoventral view; basal plates (BP); 9th
paratergites (Pt9).
Akademie der Landwirtschaftswissenschaften der DDR, a male labeled “Peru,
Amer.”, with the abdomen and pygophore both carded, is designated the
voucher specimen.
Lincus laminatus, new species
(Figs. 100-103)
Description. Fuscous to dark brown with following light brown to yellowish
brown: macule on humeri, on base of scutellum mesially and on disk of
each corium; calloused marginal line or macule expanding from middle of
connexival segments to form broken submarginal band on abdominal venter;
many interstitial areas of venter.
Head 2.5 mm wide across eyes, 2.0 mm long. Distance between lateral
limits of ocelli (1.15 mm) slightly greater than interocular width (1.10 mm).
Width of eyes (0.7 mm) greater than anterior-posterior diameter (0.6 mm)
42
NEW YORK ENTOMOLOGICAL SOCIETY
Figs. 94-99. Linens styliger. 94. Head and pronotum. 95. Pronotal lobe. 96. Genital plates,
caudoventral view; basal plate (BP). 97. Dorsal margin and mesial emargination of pygophore,
caudal view, proctiger omitted. 98. Same, lateral view, probable anterior limit of exposure
indicated by broken line. 99. Genital cup, dorsal view; proctiger (Pr).
from dorsal view. Juga exceeding tylus, their lateral margins parallel between
concavity above antennifers and convexity at apices.
Pronotal lobes long, narrow, their anteromesial and posterolateral margins
subparallel, extending laterad of eye by distance equal to 0.2 width of an
eye; emargination caudad of lobes reaching mesad of lateral limit of eye by
distance equal to 0.2 width of eye. Anterolateral margins of pronotum quite
sinuous. Width of pronotum at humeri 7.1 mm, mesial length 3.1 mm.
Scutellum 4.3 mm wide at base, 5.3 mm long; fovea in basal angles with
large deep puncture; apex weakly reflexed on each side.
Pygophoral emargination V-shaped; dorsal margin of pygophore stepped
down toward mesial emargination and riser of step recessed (Fig. 100).
Posterodorsal border of genital cup with 2 laminae on each side of emar-
gination near dorsal margin and dorsad of patch of dense stout setae (Fig.
101); part of dorsal laminae visible above dorsal pygophoral margin from
caudal view, and some of setae visible in mesial emargination. Posterior
surface of pygophore deeply impressed between rim of mesial emargination
and lateral border of pygophore. Parameres as in Figure 102. Aedeagus as
in Figure 103; membranous conjunctiva enveloping much of ejaculatory
VOLUME 91, NUMBER 1
43
Figs. 100-103. Linens laminatus. 100. Dorsal margin and mesial emargination of pygo-
phore, caudal view, proctiger omitted; lamina (1); setae (s). 101. Posterodorsal border of genital
cup; lamina (1); setae (s). 102. Paramere. 103. Aedeagus, lateral view; conjunctiva (c); ejaculatory
duct (ej.d.); median penial lobes (mpl); theca (t).
duct, remainder of aedeagus heavily pigmented and sclerotized; median
penial lobes elongate.
Length about 1 1.5 mm.
Distribution. Peru. Known only from holotype.
Holotype. S, labeled “PERU: Loreto Prov. Amazon Safari Camp. Rio
Manon NNW Iquitos, ca. 3° 42' S 73° 14' W.” Deposited in American
Museum of Natural History. The holotype is completely without appendages.
No paratypes.
Comments. This species is much like L. styliger, differing in the male
genitalia and in the parallel lateral margins of the juga.
Lincus substyliger, new species
(Figs. 104-107)
Description. Dorsum fuscous with humeral angles and macule on disk of
each corium reddish orange; basal fourth of last antennal segment yellowish.
Venter fuscous to dark castaneous.
44
NEW YORK ENTOMOLOGICAL SOCIETY
Figs. 104-107. Lincus substyliger. 104. Head and pronotum. 105. Pygophore, caudal view,
proctiger omitted. 106. Same, profile. 107. Same, dorsal view; proctiger (Pr).
Antennal segments 0.8, 0.8, 1.1, 1.4, 1.9 mm long; basal segment extending
slightly beyond apex of head. Rostral segments 2-4 about 2.5, 2.0, 2.0 mm
long. Anterior-posterior diameter of eyes and width from dorsal view sub-
equal, about 0.6-0.65 mm. Interocular width 1.1 mm. Distance across ocelli
1.2 mm; greatest diameter of each ocellus about 0.25 mm. Juga slightly
longer than tylus, their lateral margins subparallel between anteocular con-
cavity and apex. Width of head across eyes 2.4 mm, length 2.1 mm. Vertex
normally convex.
Pronotal lobes parallel-sided, longer than wide, extending laterad of eyes
by 0. 1-0.2 width of an eye; incision caudad of lobes reaching mesad from
lateral limit of corresponding eye by 0. 3-0.4 width of eye. Width of pronotum
across humeri 5.6 mm, mesial length 2.3 mm. Disk rugosely punctate. An-
terolateral margins weakly sinuous (Fig. 104).
VOLUME 91, NUMBER 1
45
Figs. 108-112. Lincus subuliger. 108. Head and pronotum. 109. Genital plates, caudo ventral
view. 110. Pygophore, caudal view, proctiger omitted. 111. Same, lateral view. 112. Genital
cup; proctiger (Pr).
Scutellum 3.5 mm wide at base, 4.4 mm long. Fovea present in each basal
angle.
Pygophoral emargination narrowly U-shaped from caudal view, notched
at junction with dorsal margin where rim continues into genital cup and
flattens into lamina inclined anterodorsad (Figs. 105, 106). Apex of proctiger
flattened and expanded, attaining width about one-fourth greater than width
of proctiger at base of expansion.
Length about 1 1.5 mm.
Distribution. Colombia. Known only from the holotype.
Holotype. S, labeled (a) “Colombia: Road Bogota-Villavicencio. June 29,
1965. 1500 M.” (b) “P. & B. Wygodzinsky, Collectors.” Deposited in the
American Museum of Natural History.
No paratypes.
Comments. The pygophore of this species is similar to that of L. styliger,
46
NEW YORK ENTOMOLOGICAL SOCIETY
especially in having a lamina within the genital cup on each side of the mesial
emargination near the dorsal margin of the cup. The most conspicuous
difference between these two species with respect to the male genitalia is
that the proctiger of L. styliger is neither flattened nor expanded apically.
This species resembles L. subuliger in having the apex of the proctiger
flattened and expanded, although in L. subuliger the expansion is much
greater, about three-fourths wider than the proctiger at the base of the ex-
pansion. The rim of the mesial pygophoral emargination of L. subuliger
continues into the genital cup as a low ridge on each side of the emargination
and does not enlarge into a lamina as it does in this species.
Lincus subuliger Breddin
(Figs. 108-112)
Lincus subuliger BxQddm, 1908, p. 33, figs. 17, 18.
Diagnosis. Pronotal lobes extending laterad of eyes by 0.2-0. 4 width of
eye, longer than wide at base (Fig. 108). Width of each eye about 0.55 of
interocular width; distance across ocelli subequal to interocular width. Juga
exceeding tylus, their lateral margins subparallel anterior to concavity above
antennifers. First and second antennal segments subequal in length. Basal
plates diagonally truncated apically (Fig. 109). Mesial emargination of py-
gophore narrowly U-shaped; posterolateral corners of pygophore bent pos-
teriorly (Fig. 110); profile of pygophore quite convex (Fig. Ill); proctiger
expanded and flattened apically into small paddle (Fig. 1 12).
Length about 1 1-1 1.5 mm.
Distribution. Colombia, Venezuela.
Comments. Breddin’s type, which was examined, is a female from an
unknown locality.
The proctiger of this species is remarkable for its apical expansion.
DEPOSITION OF PARATYPES
AL Akademie der Landwirtschaftwissenschaften
FIDE H. Dodge Engleman collection
LHR author’s collection
MZSP Museu de Zoologia de Sao Paulo
RNH Rijksmuseum van Natuurlijke Historic
USNM U.S. National Museum
ACKNOWLEDGMENTS
I am particularly grateful to Drs. G. Petersen (Akademie der Landwirtschaftwissenschaften
der DDR), A. Kaltenbach (Naturhistorisches Museum Wien), W. R. Dolling (British Museum
[Natural History]), M. W. R. de V. Graham (Hope Entomological Collections) and Randall T.
VOLUME 91, NUMBER 1
47
Schuh (American Museum of Natural History) for the loan of types and other specimens.
Additionally, specimens were loaned by Drs. H. Brailovsky (Universidad Nacional Autonoma
de Mexico), H. Dodge Engleman, Richard C. Froeschner (U.S. National Museum), J. Grazia
(Universidade Estadual de Campinas) P. van Doesburg (Rijksmuseum van Natuurlijke Historie)
and A. Willink (Universidad Nacional de Tucuman).
LITERATURE CITED
Breddin, G. 1904. Neue Rhynchotenausbeute aus Siid-Amerika. Soc. Entomol. 1 8(20): 153-
154.
Breddin, G. 1908. Beitrage zur Systematik der Pentatomiden Siid-Amerikas. Zweites Stiick.
Sitz.-Ber. Ges. Natf Fr. Berlin (1908):24-36.
Dallas, W. S. 1851. List of the specimens of hemipterous insects in the collection of the British
Museum. London. Part 1.
Distant, W. L. 1899. XLIX. Rhynchotal notes. III. Heteroptera: Discocephalinae and Pen-
tatominae (part). Ann. Mag. Nat. Hist. (7)4:421-445.
Distant, W. L. 1900. Contributions to a knowledge of the Rhynchota II. Rhynchota Central
America. Trans. R. Entomol. Soc. London (1900):687-695.
Gaedike, H. 1971. Katalog der in den Sammlungen des ehemaligen Deutschen Entomolo-
gischen Institutes aufbewahrten Typen-V. Beitr. Entomol. 21(l/2):79-159.
Rolston, L. H. 1981. Ochlerini, a new tribe in Discocephalinae (Hemiptera:Pentatomidae).
J. New York Entomol. Soc. 89(l):40-42.
Rolston, L. H. and F. J. D. McDonald. 1979. Keys and diagnoses for the families of Western
Hemisphere Pentatomoidea, subfamilies of Pentatomidae and tribes of Pentatominae
(Hemiptera). J. New York Entomol. Soc. 87(3): 189-207.
Ruckes, H. 1958. New genera and species of neotropical discocephaline and halyine penta-
tomids (Heteroptera, Pentatomidae). Amer. Mus. Novitates no. 1868, 27, pp.
StM, C. 1867. Bidrag till Hemipterernas systematik. Conspectus generum Pentatomidum
Americae. Ofv. K. Svenska Vet.-Akad. Forhl. 14(7):522-534.
StM, C. 1 872. Enumeratio Hemipterorum. 2. Enumeratio Cimicinorum Americae. K. Svenska
Vet.-Akad. Handl. 10(4):3-65.
Walker, F. 1867. Catalogue of the specimens of Heteropterous-Hemiptera in the collection
of the British Museum. London. Part 1.
Westwood, J. O. in Hope, F. W. 1837. A catalogue of Hemiptera in the collection of the Rev.
F. W. Hope, M. A. with short Latin descriptions of the new species. London. Part 1.
Accepted for publication November 16, 1982.
NEW YORK ENTOMOLOGICAL SOCIETY
91(1), 1983, pp. 48-56
ON THE BIOLOGY AND FOOD PLANTS OF LYGAEUS TURCICUS
(FABR.) (HEMIPTERA: LYGAEIDAE)i
James A. Slater
Section of Systematic and Evolutionary Biology,
University of Connecticut, Storrs, Connecticut 06268
Abstract. — ThQ food plant of Lygaeus turcicus (F.) is shown to be the false sunflower, Heliopsis
helignthoides (L.) (Sweet) (Asteraceae) rather than species of milkweeds (Asclepiadaceae). The
literature is reviewed and the records of L. turcicus on milkweeds are believed to pertain to
Lygaeus kalmii St^l. Laboratory rearing records are summarized for L. turcicus on seeds of
Heliopsis helianthoides, Asclepias syriaca L. and sunflowers. Nymphs of all five instars and the
egg are described. The nymphs are compared and contrasted with those of Lygaeus kalmii.
For many years I have been puzzled by the scarcity of Lygaeus turcicus
(F.) in New England and the upper midwest despite the abundance of its
supposed food plant, the large milkweed, Asclepias syriaca L., and the abun-
dance upon this plant of the closely related milkweed bug, Lygaeus kalmii
Stal. Although on two occasions specimens were taken in late summer on
this milkweed, I have never encountered a breeding population in the north-
east. The scarcity of turcicus could have several causes: 1) it is at the northern
periphery of its range in the northeast; 2) it does not breed in the northern
states, but rather migrates north in the summer, as is the case with Oncopeltus
fasciatus (Dallas) (Dingle, 1965); 3) it is outcompeted by Lygaeus kalmii\
or, 4) it is associated with some plant other than Asclepias syriaca L. With
the last thought in mind, I have spent considerable time in the held observing
other milkweed plants, but without Ending a specimen of L. turcicus.
It has been apparent for sometime (see Slater, 1964) that many records
of L. turcicus actually refer to L. kalmii and others cannot, in the absence
of specimens, be assigned to either species. Indeed some authors, e.g., Hei-
demann (1894), have considered the two to be synonymous; this is certainly
not the case as both adults and nymphs differ in a number of ways.
Uhler (1872) stated that eggs were deposited on Asclepias. In 1878 he
noted that records in the T. W. Harris collection suggested that adults and
nymphs were present on Asclepias syriaca. Provancher (1886) reported tak-
‘ This work was supported in part by a grant from the National Science Foundation.
The publication costs of this article were defrayed in part by page charge payment. This article
must therefore be hereby marked ‘Advertisement” in accordance with 18 U.S.C. §1734 solely
to indicate this fact.
VOLUME 91, NUMBER 1
49
ing it on Asclepias cornuti Dene. {=A. syriaca). Townsend (1887), in a de-
tailed paper treating the life history of what he called turcicus, quoted from
Uhler’s (1878) paper to demonstrate that in Michigan turcicus was taken on
Asclepias syriaca as early as 1832 (Harris collection) and that “larvae” were
present. Townsend also noted however that turcicus was seldom found in
Michigan “on any other plant than A. tuberosa although sometimes on A.
syriaca."^ He does mention an individual specimen taken on ragweed, “a
tall weed,” and a flowering almond, but considered these accidental occur-
rences. In 1891 Townsend again reported turcicus feeding on A. tuberosa L.
and seemed to have little doubt that this was the principal food plant.
There are numerous later records on milkweeds. Robertson (1891) reports
turcicus on flowers of A. cornuti and A. incarnata L., Blatchley (1895) reports
adults and nymphs on A. cornuti, Morrill (1910) states that Asclepias is the
natural food and, as recently as 1944, Froeschner reports it on flowers of
Asclepias tuberosa. Nymphs are recorded as present several times suggesting
that milkweeds are the host plants and that the preferred host may be the
butterfly weed, A. tuberosa. I have attempted to take turcicus from the orange
butterfly weed on a number of occasions without success, although Lygaeus
kalmii occurs there.
It is true that other plants have been associated with turcicus. Morrill
(1910) reports it on cotton (this record apparently repeated by Hargreaves,
1948) and on alfalfa in Texas. Banks (1912) lists it from Ceanothus in
Virginia (record repeated by Barber, 1912, 1923, and Torre-Bueno, 1946,
among others). Blatchley (1926) lists it from flowers o^ Rhus hirta (L.) Sudw.
Robertson’s (1929) compendium of plant associations lists it on flowers of
26 species of plants, only three of which are milkweeds. None of these non-
milkweed records give any indication that immature stages were present and
therefore do not suggest a breeding host relationship.
Several references establishing Lygaeus kalmii Stal as breeding on various
species of milkweeds can be found in the literature (Simanton and Andre,
1936; see Slater, 1 964, for references), and it has been reared in the laboratory
on dry milkweed seeds in a manner similar to Oncopeltus fasciatus (Dallas).
The paucity of recent records of L. turcicus on milkweeds, coincident with
the increase of records of L. kalmii, strongly suggests that the earlier records
of turcicus might in part, or entirely, refer to L. kalmii. (See following article
by Wheeler for kalmii food plants.)
The establishment of the principal food plant of L. turcicus came quite
unexpectedly; the manner in which this question was resolved emphasizes
strongly the importance of, and need for, enthusiastic amateurs in American
entomological study.
In 1979 the Rev. James M. Sullivan of St. Louis, Missouri sent me a
letter expressing his pleasure with the recent publication of my book with
Dr. R. M. Baranowski (Slater and Baranowski, 1978). Rev. Sullivan stated
50
NEW YORK ENTOMOLOGICAL SOCIETY
that for many years he had been collecting host plant records of various
Hemiptera, and he included sample pages for several species, one of which
was Lygaeus turcicus. The Sullivan records indicated that turcicus did not
breed upon milkweeds but rather on the composite Heliopsis helianthoides
(L.) Sweet, the false sunflower. Rev. Sullivan generously made available his
entire file on turcicus, and to him is due the credit for establishing the
definitive host plant.
The Sullivan records list turcicus from 1 8 species of plants in eight families.
Of 49 records 28 are from H. helianthoides, and 1 1 of the remaining 21 are
from other composites (Asteraceae). H. helianthoides was the only plant
upon which copulation was observed and the only plant upon which nymphs
were taken; his records included all collections made later than June 30.
There was only a single record of turcicus occurring on a milkweed— adults
taken on the inflorescence of Asclepias incarnata.
The most compelling of the Sullivan records for L. turcicus on H. helian-
thoides are summarized:
1. September 5, 1976 — nymphs of various instars feeding.
2. September 8 to September 29, 1976 — nymphs present and molting to
later instars on same plants during period.
3. September 15, 1976 — eight nymphs present, one reared to adult.
4. Records of adults copulating June 30, 1972, July 30, 1973, August 28,
1974 and August 9, 1977.
5. More than 50 specimens July 25, 1979.
6. Six records of feeding from June through August of several years.
On August 15, 1982 A. G. Wheeler, Jr. (pers. comm.) took many adults
and first, second and third instar nymphs on H. helianthoides in Randolph
County, West Virginia (Route 2 1 9 midway between Valley Head and Mingo).
In July and August 1979 Rev. Sullivan sent specimens of L. turcicus from
St. Louis, Missouri. They were carried through four generations in this
laboratory entirely upon the dried seed heads of H. helianthoides with almost
no mortality.
As noted below turcicus was reared successfully for more than one gen-
eration on dried sunflower seeds and also upon seeds of Asclepias syriaca.
Specimens were reared both in the open laboratory and in an environmental
chamber. Growth was more rapid on milkweed seeds than upon sunflower
seeds.
Thus it appears that L. turcicus utilizes Heliopsis helianthoides as its
principal and possibly only breeding host. However, it certainly is capable,
in the laboratory at least, of completing its life cycle on other plants, including
milkweeds. This is not really surprising, as in the laboratory Oncopeltus
fasciatus has been successfully maintained upon both sunflower and peanut
seeds, although there is no evidence that it ever breeds upon these plants in
VOLUME 91, NUMBER 1
51
the field. The choice of host plants in the field depends upon many factors,
not merely the ability of the insect to survive and reproduce on a given
plant. Slater and Wilcox (1973) suggest that many essentially host specific
Lygaeidae tend to colonize other plants; occasionally such attempts are suc-
cessful for a generation or two, and in some cases a “better” host may be
selected, which, in time, becomes the primary plant upon which the insect
breeds.
While there is no firm evidence that L. turcicus breeds upon any other
plant than H. helianthoides, the number of Rev. Sullivan’s records from
other composites suggests that under favorable conditions some of these
species may serve as hosts.
What is clearly evident is that L. turcicus is not a milkweed bug. It is
unfortunate that Robertson (1929) apparently did not distinguish turcicus
from kalmii. To my knowledge he is the only previous author to report
turcicus from H. helianthoides. Many of his records are from composites,
suggesting that he did, in part, have turcicus before him. (Robertson worked
at Carlinville, Illinois not far from the St. Louis area.)
The scarcity of L. turcicus in the north, compared with populations in the
middle Mississippi valley area, suggests that its breeding range may be more
southern than previously thought, with a late summer movement northward
in favorable years. Northern records should be carefully checked to attempt
to test this hypothesis.
LABORATORY REARING
Eggs of L. turcicus were obtained from St. Louis County, Missouri, July
25, 1979. The insects were maintained for two generations in plastic con-
tainers with dry seeds of H. helianthoides and a water source. At this time
fresh seed heads were introduced together with dried seeds, and individual
egg masses were isolated in petri dishes. The colonies were first maintained
at room temperatures and later placed in a rearing chamber at 75°F with a
16-hour day cycle for most of their development.
When seed heads of H. helianthoides were introduced into a colony, insects
of all instars sought them actively. Young nymphs moved deep into the
heads and were almost invisible despite the bright red color of the abdomen.
Eggs were laid in clumps or loose masses of 15 to 50, preferentially upon
cotton but sometimes loosely in the litter on the floor of the rearing cages.
Individual nymphs were not isolated, but colonies were examined daily
and notes on egg laying, copulation, molting and death of adults recorded.
While this method is less accurate than isolating individuals, the large num-
ber of observations taken from 35 colonies, some of which were maintained
through several generations, has yielded data that is probably a reasonably
reliable expression of the life cycle. The duration of the first stadium is
appreciably shorter than that of stadia II, III, and IV, the latter three stadia
52
NEW YORK ENTOMOLOGICAL SOCIETY
Table 1. Summary of laboratory rearing of Lygaeus turcicus (F.) reared on Heliopsis he-
lianthoides (L.) Sweet.
N Mean Median Range SD
Copulation to egg laying 1 7
Adult emergence to copulation 1 3
Egg laying to hatching 34
Instar I to instar II 35
Instar II to instar III 45
Instar III to instar IV 51
Instar IV to instar V 52
Instar V to adult 73
Adult longevity 34
5.06
5.5
3-8
1.56
6.92
7
5-10
1.61
7.24
8
6-10
1.13
3.77
4.5
2-7
1.19
6.93
7
4-11
1.88
7
7
3-11
1.93
6.62
7.5
4-12
1.83
9.47
11
4-19
3.15
60.91
59
26-97
19.31
are of equal length, and the duration of the fifth stadium is the longest (Table
1, Fig. 1). This life cycle agrees with those of many other hemipterans. The
longevity of the adults is extremely variable but can be as long as three
months.
In addition to the colonies maintained on Heliopsis helianthoides, similar
colonies were established on commercial sunflower seeds and dried seeds of
Asclepias syriaca. Lygaeus turcicus is capable of completing its life cycle on
both of these food sources. Although these colonies were observed only
sporadically on milkweeds, the length of the life cycle and the mortality
appeared similar to colonies reared on Heliopsis. Where only sunflower seed
was available, mortality was increased, the individual nymphs were smaller,
and the duration of individual stadia appeared more erratic.
In several crosses attempted between Lygaeus kalmii and Lygaeus tur-
cicus, no mating was observed and no fertilized eggs were produced.
IMMATURE STAGES
Nymphs of Lygaeus turcicus are readily distinguishable from those of
Lygaeus kalmii. In the latter species the abdomen is conspicuously longi-
tudinally striped with red and pale yellow. There is a broad, median, red
stripe and an even broader red stripe somewhat laterad of midway between
meson and each lateral margin. The intervening area is pale yellow with a
“sprinkling” of tiny red dots; the lateral margins are broadly white. In turcicus
the abdomen has the appearance of being nearly uniformly red rather than
striped. However, as noted in the descriptions that follow, there is a tendency
for early instars to have obscure stripes. In such cases kalmii nymphs can
readily be distinguished by having a very conspicuous transversely elongate-
elliptical black spot in the center of the sub-lateral red stripe on each ab-
VOLUME 91, NUMBER 1
53
ADULTHOOD TO
COPULATION
, I I , COPULATION TO
' EGG DEPOSITION
,| I I , EGG DEPOSITION
' TO HATCHING
' 1 I ^ ' LENGTH STADIUM I
• 1 I 1 • LENGTH STADIUM II
' 1 I I ' LENGTH STADIUM III
, I I I , LENGTH STADIUM IV
^ j I ^ LENGTH STADIUM V
1 I I I I I I I I I I I 1 I I I I I I
I 2 3 4 5 6 7 8 9 10 II 12 13 14 15 16 17 18 19
Fig. 1. Length of life cycle of Lygaeus turcicus (F.) reared on Heliopsis helianthoides (L.)
Sweet.
dominal tergum from segments two through six. In turcicus this area is
undifferentiated in color. These dark spots, plus the darkened areas around
the abdominal scent glands, give nymphs of kalmii a striking spotted ap-
pearance.
The labium is much longer in turcicus than in kalmii. In kalmii the labium
reaches only the metacoxae in early instars and only between the mesocoxae
in later instars. In turcicus the labium reaches well onto the abdomen as late
as the fourth instar, and even in the fifth instar it attains the posterior end
of the metacoxae.
DESCRIPTION OF Lygacus turcicus nymphs
Fifth instar. Coloration bright orange-red with strongly contrasting choc-
olate brown to black coloration as follows: a broad comma-shaped area that
curves antero-laterad on each pronotal calli area, a small paler spot on
posterior pronotal margin on either side of midline, meso- and metathoracic
wing pads, antero-lateral corners of scutellum, elliptical areas around ab-
dominal scent gland openings between terga 4-5 and 5-6, a mesal patch on
tergum 8 and sterna 7 and 8, all appendages (but distal ends of femora paler).
Dorsal coloration of head chiefly dull reddish brown. Vertex with a speckled,
longitudinal, dark stripe on either side of midline, darkened anteriorly on
tylus along inner margins of juga. Head below orange posterior to antennal
bases, dark brown anteriorly.
General form typical for genus. First antennal segment exceeding apex of
54
NEW YORK ENTOMOLOGICAL SOCIETY
tylus. Labium attaining posterior ends of metacoxae. Length head 1.56^,
width 1 .88, interocular space 1.31. Length pronotum 1.12, width 2.44. Length
mesothoracic wing pads 2.94. Length abdomen 5.50. Length labial segments
I 1.00, II 0.94, III 0.94, IV 0.88. Length antennal segments I 0.44, II 1.12,
III 0.94, IV 1.38. Total length 9.44.
Fourth instar. General form and color as in instar five but dark coloration
reddish brown rather than chocolate brown to black. Labium extending well
onto second abdominal sternum. Length head 0.88, width 1.31, interocular
space 0.81. Length pronotum 0.56, width 1.44. Length wing pads 0.88.
Length abdomen 2.56. Length labial segments I 0.75, II 0.75, III 0.69, IV
0.62. Length antennal segments I 0.38, II 0.75, III 0.62, IV 1.00. Total length
5.50.
Third instar. Similar to instar four. Head markings as noted in instar five
but much more strongly contrasting. Abdomen tending to show a narrow,
darker, longitudinal orange stripe and shading to darker orange laterally.
Lateral abdominal margins with a narrow white stripe present. Length head
0.94, width 1.12, interocular space 0.78. Length pronotum 0.50, width 1.25.
Length wing pads 0.38. Length abdomen 1.56. Length labial segments I 0.62,
II 0.62, III 0.62, IV 0.62. Length antennal segments I 0.31, II 0.56, III 0.62,
IV 0.75. Total length 4.50.
Second instar. Very similar to instar three. Thoracic terga each marked
with an irregular transverse dark “dash.” Abdomen laterally with a broad
pale yellow to translucent white border. Length head 0.66, width 0.76, in-
terocular space 0.51. Length pronotum 0.32; width 0.90. Length abdomen
1 .73. Length labial segments 1 0.42, II 0.42, III 0.42, IV 0.42. Length antennal
segments I 0.20, II 0.37, III 0.37, IV 0.56. Total length 3.05.
First instar. Head and thorax brown, strongly contrasting with bright red
abdomen, each thoracic tergum marked with a nearly black transverse dash,
similar to instar II. Abdomen nearly uniformly red except for pale margins.
(Abdominal coloration variable; some nymphs have abdomen pale yellowish
flecked with red and a darker reddish central longitudinal stripe.) Legs and
antennal segments I-III pale brown, antennal segment IV suffused with
reddish. Labium extending to middle of abdomen. Length head 0.59, width
0.56, interocular space 0.39. Length pronotum 0.20, width 0.56. Length
abdomen 1.02. Length labial segments I 0.34, II 0.34, III 0.34, IV 0.34.
Length antennal segments I 0.15, II 0.29, III 0.27, IV 0.49. Total length
1.81.
Egg. Smooth, glabrous, broadly elliptical with ten short subtruncate cho-
rionic processes around anterior pole. Length 1.34, width 0.76.
^ All measurements are in millimeters.
VOLUME 91, NUMBER 1
55
ACKNOWLEDGMENTS
I should like to express my deep appreciation to the following: Rev. James M. Sullivan of
St. Louis, Missouri for bringing the food plant of Lygaeus turcicus to my attention, allowing
me to utilize his extensive field observations and for sending me live individuals of the species;
Ms. Marianne Hassey (University of Connecticut) for her careful maintenance of the laboratory
colonies and aid in interpretation of the data; Dr. A. G. Wheeler, Jr. (Pennsylvania Department
of Agriculture) for providing me with the West Virginia host record and for his careful review
of the manuscript and to Mrs. Elizabeth Slater (University of Connecticut) for aid in preparation
of the manuscript.
LITERATURE CITED
Banks, N. 1912. At the Ceanothus in Virginia. Entomol. News 23:102-1 10.
Barber, H. G. 1912. Lygaeus turcicus Fab. and its allies. J. New York Entomol. Soc. 20:210-
211.
Barber, H. G. 1923. Family Lygaeidae. In: Britton, Guide to the Insects of Connecticut: Pt.
IV. The Hemiptera or Sucking Insects of Connecticut. Bull. Conn. Geol. Nat. Hist. Surv.
34:708-737.
Blatchley, W. S. 1895. Notes on the winter insect fauna of Vigo County, Indiana II. Psyche
7:267-270, 279-281.
Blatchley, W. S. 1926. Heteroptera or True Bugs of Eastern North America, with Especial
Reference to the Faunas of Indiana and Florida. Nature Publishing Co., Indianapolis,
1116 pp.
Dingle, H. 1965. The relation between age and flight activity in the milkweed bug, Oncopeltus.
J. Exp. Biol. 42:269-283.
Froeschner, R. C. 1944. Contributions to a synopsis of the Hemiptera of Missouri Part III.
Amer. Midland Nat. 31(3):638-683.
Hargreaves, H. 1948. List of recorded cotton insects of the world. Commonwealth Inst.
Entomology, London, pp. 1-50.
Heidemann, O. 1894. In: Proceedings of meetings. Proc. Entomol. Soc. Washington 3:106.
Morrill, A. W. 1910. Plant bugs injurious to cotton bolls. Bull. U.S. Bur. Ent. 86:1-1 10.
Provancher, L. 1 886. Petite faune entomologique du Canada Vol. 3, Les Hemipteres. Quebec,
pp. 1-205.
Robertson, C. 1891. Flowers & insects. Asclepiadaceae to Scrophulariaceae. Trans. St. Louis
Acad. Sci. 1891:569-577.
Robertson, C. 1929. Flowers & Insects. Privately printed, Carlin ville, Illinois, pp. 1-221.
Simanton, W. A. and F. Andre. 1936. A biological study of Lygaeus kalmii Stal (Hemiptera-
Lygaeidae). Bull. Brooklyn Entomol. Soc. 31:99-107.
Slater, J. A. 1964. A Catalogue of the Lygaeidae of the World. Vol. 1. University of Con-
necticut, Storrs, pp. v-xviii + 1-778.
Slater, J. A. and R. M. Baranowski. 1978. How to Know the True Bugs (Hemiptera-Heter-
optera). Wm. C. Brown Co. Publ., Dubuque, Iowa, pp. v-x + 1-256.
Slater, J. A. and D. Wilcox. 1973. The chinch bugs or Blissinae of South Africa (Hemiptera:
Lygaeidae). Mem. Entomol. Soc. Sth. Africa 12:1-135.
Torre-Bueno, J. R. 1946. A synopsis of the Hemiptera-Heteroptera of America north of
Mexico. III. Family XL Lygaeidae. Entomologica Amer. 26:1-141.
Townsend, C. H. T. 1887. On the life history of Lygaeus turcicus Fabr. Entomologica Amer.
3:53-55. (Omitted in bibliography of Slater, 1964.)
Townsend, C. H. T. 1891. Hemiptera collected in southern Michigan. Proc. Ent. Soc. Wash-
ington 2(2):52-56.
56
NEW YORK ENTOMOLOGICAL SOCIETY
Uhler, P. R. 1872. Notices of the Hemiptera of the western territories of the United States
chiefly from the surveys of Dr. F. V. Hayden. In: Prelim. Rep. U.S. Geol. Survey Mont.
1872:392-423.
Uhler, P. R. 1878. Notices of the Hemiptera Heteroptera in the collection of the late T. W.
Harris, M. D. Proc. Boston Soc. Nat. Hist. 19:365-446.
Received July 1, 1982; accepted October 12, 1982.
NEW YORK ENTOMOLOGICAL SOCIETY
91(1), 1983, pp. 57-62
THE SMALL MILKWEED BUG, LYGAEUS KALMII
(HEMIPTERA: LYGAEIDAE):
MILKWEED SPECIALIST OR OPPORTUNIST?
A. G. Wheeler, Jr.
Bureau of Plant Industry, Pennsylvania Department of Agriculture,
Harrisburg, Pennsylvania 17110
Abstract.— Lygaeus kalmii StM, the small milkweed bug, often is considered a milkweed
specialist. A review of recent literature and observations of nymphs and adults on composites
(Asteraceae) and plants of other families in Missouri, Pennsylvania, and West Virginia show
that L. kalmii uses various food sources. The feeding strategies of this lygaeid are briefly
compared with those of Oncopeltus fasciatus (Dallas), a lygaeine restricted to milkweeds and
related plants, and the Palearctic Lygaeus equestris (L.), which prefers asclepiads but feeds on
plants of numerous families.
Relatively few North American insects have been thoroughly studied, and
when a particular species is considered biologically well known, further
investigation may be stifled. Any misconceptions about the habits of that
species often are perpetuated in the literature and are difficult to rectify.
Any large, strikingly colored insect, especially a common one, is apt to
convey the impression “well studied.” Indeed, aposematic species of the
lygaeid genera Oncopeltus and Lygaeus have long been characterized as
milkweed feeders and are now known to sequester and store cardenolides
(cardiac glycosides) from their hosts (Dufley and Scudder, 1972; Scudder
and Dufley, 1972). L. kalmii Stal, known in the United States by the ap-
proved common name small milkweed bug, sometimes has been assumed
representative of the specialized fauna associated with the Asclepiadaceae.
Caldwell (1974) considered its utilization of resources similar to that of O.
fasciatus (Dallas), a definite milkweed specialist, although he acknowledged
the biology of L. kalmii was not as well known. Dailey et al. (1978) referred
to both lygaeine species as “milkweed-specific.”
Several species of Lygaeus, however, seem to furnish examples of hard-
to-correct biological misconceptions. L. turcicus F., a close relative of kalmii,
has been considered a milkweed bug although specific observations were
lacking. Published records from Asclepias spp. are unreliable because many
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58
NEW YORK ENTOMOLOGICAL SOCIETY
observers did not distinguish turcicus from kalrnii. Slater (1983, this issue)
has shown that turcicus feeds mainly on composites, especially ox-eye [He-
liopsis helianthoides (L.) Sweet], rarely, if ever, developing on milkweeds.
Slater’s paper, which I had the opportunity to review, prompted me to
reevaluate my collections of L. kalrnii from various composites, considered
merely fortuitous when first made in 1977. Recent papers by Isman et al.
(1977), Hunt (1979), and Duarte and Calabrese (1982) also led me to reap-
praise the habits of L. kalrnii. Isman et al. found adults of kalrnii collected
from Asclepias spp. in California varied widely in their cardenolide content;
they suggested that some of the adults might have developed on hosts lacking
these compounds. In Michigan, Hunt described kalrnii as an opportunist
that feeds on various food sources, especially when milkweed pods are un-
available. After determining that the chromosome number and sex deter-
mination mechanism of populations in west Texas differed from those pre-
viously recorded, Duarte and Calabrese suggested L. kalrnii might represent
a sibling complex.
Herein I summarize my records of L. kalrnii taken on plants other than
milkweeds. Through the courtesy of the Rev. James M. Sullivan, who has
made extensive collections of Hemiptera-Heteroptera (see Slater, 1983), I
am able to refer to his observations of kalrnii on various plants in Missouri.
My Pennsylvania and West Virginia observations and those made in Mis-
souri refer to the subspecies angustornarginatus Parshley (see Slater and
Knop, 1969). On the basis of these observations tentative conclusions are
drawn regarding the feeding strategies of this lygaeid.
OBSERVATIONS ON COMPOSITES
In late July 1977 I collected an adult and a 5th-instar nymph on the seed
heads of yarrow, Achillea rnillefoliurn L. (Asteraceae = Compositae), in
Adams County, Pennsylvania. Adults and a 5th instar were observed on
heads of yarrow on a return visit to this site in mid-August 1982. At about
this time I found 5 nymphs (instars III-V) and adults on heads of yarrow
growing in an old field on Dolly Sods, a wilderness area at approximately
4,000 ft. (1,219 m) in Tucker County, West Virginia. Milkweeds were not
observed in the vicinity (40-50 m) of either collecting site.
I found a larger population of L. kalrnii associated with Senecio vulgaris
L. growing in isolated colonies in an otherwise weed-free nursery in Somerset
County, Pennsylvania. On 1 4 July 1982 about 1 0 mating pairs were observed
on inflorescences and seed heads (dried capitula); many more adults were
found beneath the plants. A thorough examination of the nursery beds and
nearby hedgerows did not reveal any milkweeds or the association of kalrnii
with other composite or weed species. Two weeks later adults were common
on the ground beneath V. vulgaris; 2 adults, a 3rd, and a 5th instar were
collected on mature heads. On 19 August no nymphs were observed, but 3
VOLUME 91, NUMBER 1
59
adults were taken on fruiting heads and others were observed beneath Se-
necio.
During July and August I collected small numbers of late instars, usually
only 1 or 2 individuals, on fruiting heads or inflorescences of Centaurea
maculosa Lam., Conyza canadensis (L.) Cronq., and Erigeron strigosus Muhl.
Adults only were collected on seed heads of Ambrosia artemesiifolia L.,
Chondrilla juncea L., Daucus carota L., and Eupatorium perfoliatum L.
In Missouri the Rev. Sullivan has observed L. kalmii visiting flowers in
succession, apparently taking nectar from Eupatorium altissimum L. and
Solidago graminifolia (L.) Salisb. In early March an adult was taken on the
fresh receptacle of dandelion. Taraxacum sp., and in April on a dandelion
flower and in gravel beneath the plants. During July-October adults were
observed on inflorescences or old flower heads of Cirsium altissimum (L.)
Spreng., Eupatorium rugosum Houtt., Lactuca saligna L., Solidago altissi-
mum L., and Tragopogon dubius Scop.; apparent feeding took place on
several of the plants. In October a mating pair was found on the dried heads
of Helianthus annuus L. In the literature a mating pair is known from
goldenrod, Solidago sp. (Procter, 1946), and on goldenrod Balduf (1943)
observed an adult feeding on a dead honeybee. Maw (1976) swept L. kalmii
from Cirsium and Taraxacum.
Lygaeus kalmii on plants of other families
Occasionally I found nymphs of L. kalmii on nonmilkweed plants other
than composites. Thirty or more nymphs (instars II-V) and a smaller number
of adults were observed in August on mature inflorescences of buckwheat,
Eagopyrum sagittatum Gilib. (Polygonaceae). The small field, well removed
from milkweed and nearly weed free except for a few plants of common
ragweed. Ambrosia artemesiifolia, was bordered by a pure stand of ragweed.
No nymphs or adults, however, were observed or swept from ragweed.
Smaller numbers of nymphs (1 or 2 individuals of instars III-V) were taken
during August on Hypericum punctatum Lam. (Hypericaceae = Guttiferae)
and on or under Euphorbia spp. (Euphorbiaceae) growing in isolated colonies
in ballast along railroad tracks. In Missouri, the Rev. Sullivan observed 2
adults in early September on fruits of Euphorbia sp. He also observed prob-
able feeding in late August on the calyx of Campanula americana L. (Cam-
panulaceae) and 3 adults beneath these plants; in mid-October he found an
adult with its rostrum inserted in a capsule of Scrophularia marilandica L.
(Scrophulariaceae). In mid-September he collected an adult from an unripe
fruit of Datura stramonium L. (Solanaceae).
DISCUSSION
The observations here summarized, and the studies of Hunt (1979), suggest
that L. kalmii is not as intimately associated with Asclepiadaceae as stated
60
NEW YORK ENTOMOLOGICAL SOCIETY
in much of the literature and thus not a milkweed specialist in the sense of
Oncopeltus fasciatus. Caldwell (1974) eontrasted the migratory strategies of
these lygaeines, noting that kalmii remains in its habitat year round rather
than engaging in migratory flights. According to Caldwell, it flies to new
habitats only when food or mates are searce; he referred to its traeking of
new milkweed stands as “more opportunistic” than in O. fasciatus.
L. kalmii also may be more an opportunist in its utilization of food
resources. Adults obtain nectar from flowers in early spring and later in the
season, sometimes “moving systematically from nectary to nectary” (Hunt,
1979). In late summer and fall L. kalmii may feed on seeds and other
structures found on old flower heads.
L. kalmii also uses milkweeds for reproduction, and various asclepiads
may represent preferred hosts. The similarity in distribution of kalmii and
Aclepias syriaca L. and A. speciosa Torr. (Slater and Knop, 1969) suggests
a dependence on these plants if not an extensive coevolutionary history.
But the finding of nymphs on various composites and on plants of unrelated
families suggests that L. kalmii develops also on nonmilkweed hosts. The
observation of 1 or a few late instars on other plants may be based on nymphs
that have ascended after completing most of their development elsewhere.
It is known that nymphs feed on fallen seeds of their hosts and that 5th
instars may climb plants at metamorphosis (Hunt, 1979). It is likely, how-
ever, that the nymphs found on Senecio vulgaris and buckwheat completed
their development on those plants.
The evidence indicates that L. kalmii reproduees on plants other than
Asclepiadaceae or the related Apocynaceae, perhaps “getting by” before
milkweed pods are formed. Hunt found that this multivoltine lygaeid was
not common on milkweed in southern Michigan until late June and not
abundant until July or August. She characterized kalmii as an opportunistic
forager which preferred milkweeds but fed on a suecession of angiosperms
when asclepiads were scarce or unavailable. It is not clearly stated, however,
whether this foraging involved adults only or if nymphs occurred on non-
milkweed hosts.
Assuming reproduetion takes place on plants other than milkweeds, my
observations and Hunt’s may help explain the relatively few nymphs Dailey
et al. (1978) collected on milkweed in Ohio (152 compared to 1,021 adults).
Nymphs may have fed on fallen milkweed seeds, but it is possible that they
developed on plants of other families. Feeding on nonmilkweed hosts may
also explain the differences in cardenolide content that Isman et al. (1977)
found in field-collected L. kalmii in California.
Observations of L. kalmii on composites and other plants unrelated to
the Asclepiadaceae and Apocynaceae suggest a comparison with the Pale-
arctic lygaeine L. equestris L. This species, although preferring the asclepiad
Cynanchum vincetoxicum (L.) Pers., feeds on a succession of hosts through-
VOLUME 91, NUMBER 1
61
out the season. It has been observed on more than 60 plant species in 37
families; nymphs have been associated with 1 3 species in 1 1 families (Sol-
breck and Kugelberg, 1972). Another milkweed-associated lygaeine that is
known to develop on composites is Spilostethus pandurus var. elegans (Wolff)
in South Africa (Slater and Sperry, 1973). A lygaeine perhaps preferring
seeds of Asteraceae and feeding on some of the same composite genera as
L. kalmii (e.g., Achillea, Senecio) is the widely distributed Neacoryphus
bicrucis (Say) (Solbreck and Pehrson, 1979).
Careful field studies are needed to determine the similarity in feeding
strategies between L. kalmii and L. equestris. Such work will help determine
whether oviposition occurs a considerable distance from potential food plants
(as Kugelberg, 1977, has reported for L. equestris), whether the occurrence
of nymphs on aerial portions of plants reflects only a shortage of seeds on
the ground, the extent to which feeding preferences vary throughout the
geographic range, whether resource utilization changes during development
(as Puchkov, 1956, has discussed for various phytophagous Heteroptera),
and whether feeding on nonmilkweed plants represents a recent shift in the
host spectrum similar to the use of oleander {Neriurn oleander L.— Apocy-
naceae) by Oncopeltus fasciatus in Florida (Klausner et al., 1980).
ACKNOWLEDGMENTS
I thank the Rev. James M. Sullivan (St. Louis, Missouri) for letting me refer to his observations
on Lygaeus kalmii. R. J. Hill (Bureau of Plant Industry, Pennsylvania Department of Agri-
culture, Harrisburg) identified many of the plants from Pennsylvania and West Virginia. For
helpful comments on an early draft of the manuscript I thank E. R. Hoebeke (Department of
Entomology, Cornell University, Ithaca, New York), R. B. Root (Section of Ecology & System-
atics, Cornell University), and J. A. Slater (Biological Sciences Group, University of Connecticut,
Storrs).
LITERATURE CITED
Balduf, W. V. 1943. New food records of entomophagous insects (Hym., Dip., Col., Orth.,
Hemip.). Entomol. News 54:12-15.
Caldwell, R. L. 1974. A comparison of the migratory strategies of two milkweed bugs, On-
copeltus fasciatus and Lygaeus kalmii. Pages 304-316 in: L. Barton-Browne (ed.). Ex-
perimental Analysis of Insect Behaviour. Springer- Verlag, New York.
Dailey, P. J., R. C. Graves and J. L. Herring. 1978. Survey of Hemiptera collected on common
milkweed, Asclepias syriaca, at one site in Ohio. Entomol. News 89:157-162.
Duarte, J. and D. M. Calabrese. 1982. Is the binomen Lygaeus kalmii St;Sl (Hemiptera:
Heteroptera: Lygaeidae) applied to sibling species? Proc. Entomol. Soc. Washington 84:
301-303.
Dulfey, S. S. and G. G. E. Scudder. 1972. Cardiac glycosides in North American Asclepia-
daceae, a basis for unpalatability in brightly coloured Hemiptera and Coleoptera. J.
Insect Physiol. 18:63-78.
Hunt, L.-M. 1979. Observations of the habits of Lygaeus kalmii angustomarginatus (He-
miptera:Lygaeidae) in southern Michigan. Great Lakes Entomol. 12:31-33.
62
NEW YORK ENTOMOLOGICAL SOCIETY
Isman, M. B., S. S. Duffey and G. G. E. Scudder. 1977. Variation in cardenolide content of
the lygaeid bugs, Oncopeltus fasciatus and Lygaeus kalmii kalmii and of their milkweed
hosts {Asclepias spp.) in central California. J. Chem. Ecol. 3:613-624.
Klausner, E., E. R. Miller and H. Dingle. 1980. Nerium oleander as an alternative host plant
for south Florida milkweed bugs, Oncopeltus fasciatus. Ecol. Entomol. 5:137-142.
Kugelberg, O. 1977. Distribution, feeding habits and dispersal of Lygaeus equestris (Heter-
optera) larvae in relation to food supply. Oikos 29:398-406.
Maw, M. G. 1976. An annotated list ofinsects associated with Canada thistle (C/>.s/wm arvense)
in Canada. Can. Entomol. 108:235-244.
Procter, W. 1946. Biological Survey of the Mount Desert Region. Part VII. The Insect Fauna.
Wistar Inst. Anat. Biol., Philadelphia, 566 pp.
Puchkov, V. G. 1956. Basic trophic groups of phytophagous hemipterous insects and changes
in the character of their feeding during the process of development (in Russian). Zool.
Zh. 35:32-44.
Scudder, G. G. E. and S. S. Duffey. 1972. Cardiac glycosides in the Lygaeinae (Hemiptera:
Lygaeidae). Can. J. Zool. 50:35-42.
Slater, J. A. 1983. On the biology and food plants of Lygaeus turcicus (Fabr.) (Hemiptera:
Lygaeidae). J. New York Ent. Soc. 91:48-56.
Slater, J. A. and N. F. Knop. 1969. Geographic variation in the North American milkweed
bugs of the Lygaeus kalmii complex. Ann. Entomol. Soc. Amer. 62:1221-1232.
Slater, J. A. and B. Sperry. 1973. The biology and distribution of the South African Lygaeinae,
with descriptions of new species (Hemiptera:Lygaeidae). Ann. Transvaal Mus. 28:1 17-
201.
Solbreck, C. and O. Kugelberg. 1 972. Field observations on the seasonal occurrence of Lygaeus
equestris (L.) (Het., Lygaeidae) with special reference to food plant phenology. Entomol.
Scand. 3:189-210.
Solbreck, C. and I. Pehrson. 1979. Relations between environment, migration and reproduc-
tion in a seed bug, Neacoryphus bicrucis (Say) (Heteroptera: Lygaeidae). Oecologia 43:
51-62.
Received September 24, 1982; accepted October 13, 1982.
NEW YORK ENTOMOLOGICAL SOCIETY
91(1), 1983, pp. 63-67
A NEW SPECIES OF CLERADINI
(HEMIPTERA: LYGAEIDAE: RHYPAROCHROMINAE) FROM
THE CENTRAL AFRICAN REPUBLIC AND GHANA
B. J. Harrington
Department of Entomology, University of Wisconsin,
Madison, Wisconsin 53706
Abstract.— Navarrus ater, a new Ethiopian species, is described in a formerly monotypic genus
known only from the Oriental Region and New Guinea. Characters are presented to distinguish
N. ater from N. phaeophilus, the type species of the genus. A dorsal view drawing of the holotype
of N. ater is provided.
Members of the little-known hemipteran tribe Cleradini are nest inquilines
found principally in the nests of rodents where they feed on vertebrate blood.
In such a specialized habitat these insects are not commonly collected. In-
deed, some species are known only from the limited material of the original
description. A majority of the genera in this tribe are currently monotypic.
The cleradine genus Navarrus was described by Distant in 1901, with
Rhyparochromus phaeophilus Walker, 1872 as the type species. To date
Navarrus has remained monotypic, with N. phaeophilus known to occur
broadly in the Oriental Region and in New Guinea. The present paper
extends the distribution of Navarrus into the Ethiopian Region by describing
N. ater, a new species from the Central African Republic and Ghana. In the
following description all measurements are in mm and the Villalobos color
chart (Palmer, 1962) has been used as a standard.
Navarrus ater, new species
Virtually entire body surface dark blackish brown; scutellum subtly darker,
almost true black; tarsi, lateral pronotal carina and lateral hemelytral carina
appearing lighter, between blackish brown and sepia; subproximal one-third
antennal segment IV dark tawny, segment IV dark on distal two-thirds and
in very narrow band at proximal end.
Body subshining save for dull granular evaporative area surrounding meta-
thoracic scent gland auricle and adjacent strip along posterior margin me-
sopleuron. Pronotum, scutellum, clavus, and corium evenly covered with
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NEW YORK ENTOMOLOGICAL SOCIETY
Fig. 1 . Metathoracic scent gland auricles and evaporative areas of: (A) Navarrus ater and
(B) N. phaeophilus.
small shallow punctures; punctures larger and deeper on head and latero-
ventral aspects of pro- and mesothorax; abdomen impunctate, marked
lateroventrally on sterna II and III (first two visible) with large finely textured
areas representing a diffuse stridulitrum. Abdomen sparsely clothed with
slender recumbent posteriorly directed hairs; similar distally directed hairs
on legs and antennae; head including eyes with short anteriorly directed
hairs, some stout and bristle-like; each puncture on pronotum, scutellum
and hemelytra containing a very short minute scale-like hair.
Head very short and broad, slightly declivent from flat vertex; tylus nar-
row, appearing markedly so with antenniferous tubercles short to non-
existent; eyes large, seeming to rest on anterior pronotal angles; postocular
distance negligible; ocelli at posterior margin of head; length head 0.56; width
0.90; interocular distance 0.46. Pronotum shield-like, not noticeably bilobed,
declivent from posterior margin; transverse impression obsolete; no collar
apparent on anterior lobe; a broad continuous lateral carina running from
anterior angle to just short of posterior margin; humeral angle elevated and
knob-like, extending posteriorly a short but distinct distance beyond lateral
carina; posterior pronotal margin straight across base of scutellum; length
pronotum 1.00; width posterior margin across humeral angles 1.64; greatest
width including carinae 1.78. Length scutellum 0.90; width 0.94. Hemelytron
vaguely sinuate laterally, with a broad lateral carina; length corium 2.58;
distance apex corium to apex membrane 0.34; length claval commissure
0.50; distance apex clavus to apex corium 1.16. Labium extending between
mesocoxae; length labial segments I 0.30, II 0.26, III 0.82, IV 0.26; bucculae
long extending most of head length, with a broad U-shaped juncture at level
of proximal one-third labial segment II. Antennae with segments relatively
stout; length antennal segments I 0.24, II 0.40, III 0.26, IV 0.68. Fore femur
slightly incrassate, armed on anterior edge of ventral surface with three very
VOLUME 91, NUMBER 1
65
Fig. 2. Navarrus ater, new species. Holotype, dorsal view.
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NEW YORK ENTOMOLOGICAL SOCIETY
minute broad spines each with a stout apical bristle; meso- and metafemora
also relatively swollen; metafemur with several fine denticles on posterior
surface aligned to strike textured areas on abdomen, these two surfaces
probably constituting a stridulatory apparatus. Metathoracic scent gland
auricle large elongate crescent-shaped and arched posteriorly (Fig. la). Total
length 4.44.
Holotype. 9, CENTRAL AFRICAN REPUBLIC: La Maboke 6-9. VI. 1 973
(Linnavouri). In American Museum of Natural History, New York.
Paratypes. CENTRAL AFRICAN REPUBLIC: Same data as holotype,
19. GHANA: Tafo 10.X.1967 (UV trap) (D. Leston), 19. In J. A. Slater and
B. J. Harrington collections.
The paratype from Ghana is somewhat lighter (more toward chestnut)
than the holotype and paratype from the Central African Republic.
N. ater and N. phaeophilus have a very similar general habitus with a
broad ovoid body shape, a distinct shelf-like and reflexed lateral pronotal
Carina, an obsolete pronotal transverse impression, and a broad short head
that is not prolonged at all in the postocular region.
TV. ater can be easily distinguished from N. phaeophilus by its uniform
dark coloration. TV. phaeophilus is lighter with the ground color ranging
between chestnut and tawny. In TV. phaeophilus the hemelytral membrane
is largely fuscous but proximally marked at the midline with a distinct light
colored macula contiguous to the claval commissure. TV. phaeophilus also
has a small rounded scent gland auricle with the evaporative area confined
to the auricle itself (Fig. lb) in contrast to the large crescent-shaped auricle
of TV. ater which is surrounded by a broad evaporative area that extends
onto the posterior margin of the mesopleuron (Fig. la). Additionally, TV. ater
apparently has a stridulatory apparatus while there is no evidence of one in
TV. phaeophilus.
Such striking morphological differences might suggest separate generic
status for TV. ater. However, without male specimens for examination and
pending a comprehensive generic-level reevaluation of the tribe Cleradini
that will shed more light on whether the features shared by N. ater and N.
phaeophilus represent synapomorphies or symplesiomorphies, it seems pref-
erable to be conservative and describe this new species as a member of the
genus Navarrus.
ACKNOWLEDGMENTS
I thank Dr. James A. Slater (University of Connecticut) for the loan of specimens and Mary
Jane Spring (University of Connecticut) for her preparation of the dorsal view illustration of
the holotype. This research was supported by the College of Agricultural and Life Sciences,
University of Wisconsin, Madison and by a grant from the National Institutes of Health (No.
AI17152).
VOLUME 91, NUMBER 1
67
LITERATURE CITED
Distant, W. L. 1901. Rhynchotal notes XL Heteroptera; Family Lygaeidae. Ann. Mag. Nat.
Hist. 7(8):476.
Palmer, R. S. 1962. Handbook of North American Birds. Vol. I. Loons through Flamingos.
Yale Univ. Press, New Haven, col. pi.
Walker, F. 1872. Catalogue of the specimens of Hemiptera Heteroptera in the collection of
the British Museum. London 5:106-107.
Received July 19, 1982; accepted October 14, 1982.
NEW YORK ENTOMOLOGICAL SOCIETY
91(1), 1983, pp. 68-74
OVERWINTERING EGG MASS ADAPTATIONS OF THE EASTERN
TENT CATERPILLAR, MALACOSOMA AMERICANUM
(LEPIDOPTERA: LASIOCAMPIDAE)‘
Alejandro Segarra Carmona and Pedro Barbosa
Department of Entomology, University of Maryland,
College Park, Maryland 20742
ylfc/rac/. — Experiments were designed to determine the role of the spumaline covering eastern
tent caterpillar egg masses as an adaptation for overwintering. The ability of the spumaline to
absorb water from the environment is a direct function of temperature and water concentration
in air. Egg mass temperatures may be significantly higher than ambient temperatures. The
spumaline covering egg masses of this species may act to ameliorate low overwintering tem-
peratures and to prevent desiccation by absorbing as much moisture from surrounding air as
possible.
The survival success of overwintering insects is achieved by many ad-
aptations which overcome or compensate for extremes and fluctuations in
winter conditions. In the temperate regions adaptations like the presence of
cryoprotectants, low supercooling points and reduced metabolic activity
(dormancy) are examples of physiological adaptations. More subtle but equally
important to survival are microenvironmental adaptations, such as location
of hibernacula and external protective structures.
Malacosoma americanum (Fab.), the eastern tent caterpillar, overwinters
as a pharate larva inside its egg shell. Egg masses are laid on branches of
suitable hosts in early summer. Complete embryonation occurs within 3-4
weeks. These mature embryos remain dormant from late July to April of
the following year (Mansing, 1974). Characteristically the egg masses of all
North American species (except M. tigris) are covered with a froth called
spumaline. This material which is produced in accessory glands is deposited
by the ovipositing female on top of the newly laid eggs. Very little is known
about this material. Hodson and Weinman (1945) described important
features of the spumaline on eggs of Malacosoma disstria such as its ability
‘ This research was completed at the Department of Entomology of the University of Mas-
sachusetts, Amherst, Massachusetts.
The publication costs of this article were defrayed in part by page charge payment. This article
must therefore be hereby marked “Advertisement” in accordance with 18 U.S.C. §1734 solely
to indicate this fact.
VOLUME 91, NUMBER 1
69
to absorb atmospheric moisture and elucidated the role of spumaline in the
eclosion process.
This investigation examines, in further detail, the adaptive role of the
spumaline of overwintering Malacosoma americanum egg masses in mod-
ulating environmental temperature and humidity.
MATERIAL AND METHODS
All tent caterpillar egg masses examined were collected from black cherry
trees {Prunus serotina Ehrh.) located in Amherst, Massachusetts (42°23'N,
72°32'W) during the months of January-April 1979.
Hygroscopic properties of spumaline. To test water absorption capabilities
of egg masses, field collected egg masses were exposed to three different
temperatures: 0°C, — 5°C, — 10°C and five different saturation deficits per
temperature for a total of 1 5 treatments, replicated 3 times each with 9 egg
masses per replicate. Before being used in experiments any egg mass sections
not covered by spumaline were covered with wax. All egg masses were placed
in a controlled temperature chamber at 5°C until experiments began.
Our initial attempts to establish the hygroscopicity of the spumaline at
low temperatures and high saturation deficits resulted in water loss to the
surrounding environment. Therefore, subsequent experiments used desic-
cated egg masses, which resulted in better determination of expected water
gain. Water absorption of the spumaline is expressed in terms of percent
weight gain of egg mass (previously desiccated to a constant weight) main-
tained in humidity chambers for 24 hr. The humidity chambers consisted
of closed glass containers (105 X 76 X 76 mm) where egg masses were sus-
pended with zinc wire above a given aqueous sulphuric acid solution (Sol-
omon, 1951). Vapor pressures for corresponding sulphuric acid solutions at
temperatures ranging from 0°C to 235°C can be calculated with the aid of
“International Critical Tables” (Washburn, 1928). Unfortunately, simula-
tion of common mid-winter conditions requires creation of saturation deficits
at temperatures lower than 0°C. To obtain vapor pressures for solutions
below 0°C, extrapolations were made from correlations of loge vapor pressure
vs. temperature. In all correlations performed the Pearson’s correlation coef-
ficient was r = 0.999 {P < 0.001). Saturation deficits (SD) were calculated
from the formula:
SD^ = Cs — e (Rosenberg, 1974)
where e^ is the saturation vapor pressure at temperature t°C and e is the
actual vapor pressure, both in mm Hg.
Insulative properties of the spumaline. To determine the effect of the spu-
maline layer on the temperature of egg masses in the field, temperature
measurements were made of egg masses on trees with a thermocouple probe
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NEW YORK ENTOMOLOGICAL SOCIETY
SATURATION DEFICIT ImmHg).
Fig. I . Loge of percent of weight gain of Malacosoma arnericamun (Fab.) exposed to different
absolute humidities -10°C, -5°C and 0°C for 24 hr (mean and SD).
and a cryothermometer model Bat-5 (Bailey Instrument Co.). For each egg
mass the information recorded included type of day (e.g., cloudy or sunny),
temperature inside the spumaline (but above the eggs themselves) and am-
bient temperature (temperatures were taken to the nearest 0.5°C).
Egg masses stored in the laboratory for two weeks at 1 1°C and 9.8 g/m-
absolute humidity were utilized to study the reflectance spectrum of the
chorion and spumaline. Study of reflectance spectrum is an important tool
in determining how different light wavelengths behave on media and thus
their energy trapping characteristics. The spectrum was measured with a
Shimadzu Spectronic 210 UV Spectrophotometer equipped with an inte-
grating sphere 200 UV attachment for measuring solid materials.
VOLUME 91, NUMBER 1
71
Table 1. Differences between air temperature and egg masses of Malacosoma americanum
on clear days.
Air temperature
range (°C)
Mean temperature
difference: under
the spumaline and
air (°C)
Number of egg
masses examined
9 to 5
+ 1.6
29
4 to 0
+2.7
47
-1 to -5
+3.0
122
-6 to -10
+3.5
73
-11 to -20
+6.0
67
RESULTS
Water absorption decreased logarithmically with increasing saturation def-
icit (Fig. 1), i.e., the less water in the environment the less water the spumaline
absorbs at all the temperatures tested. The regression analyses of water
absorption at (r2 = -0.99, P < 0.01), = -0.98, P < 0.01) and - 10°C
(^2 ^ —0.99, P < 0.001), show that water absorption is a direct function of
temperature and water concentration. Examination of the regression equa-
tions’ slopes reveals an inverse relationship between temperature and the
magnitude of the slope, furthermore, the slope of the - 1 0°C regression line
is significantly larger {t = 3.86, P < 0.0005). These two facts agree with a
fundamental characteristic of hygroscopic materials where decreasing tem-
peratures mark a corresponding increase in any material’s ability to absorb
water (Hodson, 1937).
Although water uptake occurs at the air-spumaline interface, it may also
occur at the chorion-spumaline interface since the chorion also is hygroscopic
(Hodson and Weinman, 1945; and pers. obs.). A differential response to
temperature and/or humidity at each interface could account for the non-
linear nature of the water absorption curves. In addition, it was also observed
that immediately after rainfall, and at temperatures higher than 10°C, field
collected spumaline contains so much water that it can be literally squeezed
out. This illustrates the extent of spumaline’s hygroscopicity at high absolute
humidity (characteristic of higher temperatures). (Also see Hodson and
Weinman, 1945.)
Preliminary field measurements of temperature differences between egg
masses and ambient showed that although marked temperature differences
are observed on clear sunny days these disappeared during cloudy days even
when measurements are taken on the same egg mass. Thus, Table 1 presents
a summary of data for egg masses measured on clear days only. It was
observed that differences varied inversely with environmental temperature.
In some specific comparisons, egg masses were up to 1 2°C warmer than air
72
NEW YORK ENTOMOLOGICAL SOCIETY
LU
O
z
<
H
O
UJ
OC
WAVELENGTH nm
Fig. 2. Percent reflectances of the chorion ( ) and the spumaline ( ) of egg masses
held at 9.8 g/m^ absolute humidity and 1 1°C.
temperature. These differences are in concert with differences of up to 5°C
on an egg mass of M. disstria, during one cold and clear February day found
by Wellington (1950). This observation points at the importance of solar
radiation.
The renectance spectrum of the spumaline and the chorion is presented
in Figure 2. The percent reflected light decreased with decreasing wavelength
with a reflectance minimum at 342 nm, in the near ultraviolet region for
both structures. The fact that the spumaline consistently had high reflectance
values at the infrared (IR) region of the spectrum suggests that: as short high
energy wavelength radiation penetrates the chorion and spumaline some of
its energy is reemitted as IR, which in turn is trapped in the spumaline, due
VOLUME 91, NUMBER 1
73
to its large reflectivity. Thus a greenhouse effect is probably created between
the chorion and the air-spumaline interface. This effect would maintain egg
mass temperatures above those in the air on days when the UV component
of incident light is large (i.e., on sunny clear days).
DISCUSSION
The hibernacula of many insects occur under or within snow cover, bark,
leaf litter or soil. These substrates provide relatively small temperature fluc-
tuations and excellent insulative qualities (Holmquist, 1931). On the other
hand, the overwintering stage of Malacosoma americanum in its arboreal
habitat is exposed to wide environmental fluctuations and low temperatures.
Under these conditions, species like those of Malacosoma tend to have cold-
hardy overwintering stages (MacPhee, 1964). Several physiological mecha-
nisms of cold-hardiness in Malacosoma have been reported namely the
presence of high cryoprotectant levels and low supercooling points (Hanec,
1966; Mansingh, 1974). However, although important these are not the only
ways to compensate for climatic extremes. The egg masses of many over-
wintering arthropods are often protected from cold environments by ma-
terials provided by the maternal parent. The eggs of the gypsy moth can
withstand temperatures of — 50°C if the masses are covered by the hair placed
on them by the female, but die at — 1 9°C if the hairs are removed (Kulagin,
1897, cited in Danks, 1978). Similarly, the egg sac of Floridia bucculenta
protects eggs from desiccation and flooding (Schaefer, 1976). In Malacosoma
the presence of the spumaline appears to serve two purposes: to ameliorate
environmental temperatures and to prevent desiccation. The spumaline in-
sures the capture of the water when the humidity is high and serves as an
interphase for slow evaporation when air moisture is low. Extremely dry
winters seldom occur in the natural range of M. americanum where precip-
itation is usually high. The suggestion of Hodson and Weinman (1945) that
the spumaline of M. disstria was essential for its winter survival appears to
apply equally well to M. americanum.
LITERATURE CITED
Danks, H. V. 1978. Modes of seasonal adaptation in insects. I. Winter survival. Can. Entomol.
110:1167-1205.
Hanec, W. 1966. Cold-hardiness in the forest tent caterpillar, Malacosoma disstria Hubner
(Lasiocampidae:Lepidoptera). J. Insect Physiol. 12:1443-1449.
Harvey, R. B. 1923. Cambial temperatures of trees in winter and their relation to sun scald.
Ecology 4:261-265.
Hodson, A. C. 1937. Some aspects of the role of water in insect hibernation. Ecol. Monogr.
7:271-315.
Hodson, A. C. and C. J. Weinman. 1945. Factors affecting recovery from diapause and
hatching of eggs of the forest tent caterpillar, Malacosoma disstria Hbn. Univ. Minnesota
Agric. Exp. Sta. Tech. Bull. 170:31 pp., illus.
74
NEW YORK ENTOMOLOGICAL SOCIETY
Holmquist, A. M. 1931. Studies of arthropod hibernation. III. Temperatures in the forest
hibemacula. Ecology 12:387-400.
Jensen, R. E., E. F. Savage and R. A. Hayden. 1970. The effect of certain environmental
factors on the cambial temperatures of peach trees. J. Amer. Hort. Sci. 95:286-292.
Kulagin, N. 1897. Zur Biologic Ocneria dispar in Russland. Ulus. Wschr. Ent. 2:418-420.
(Original text not seen; cited in Danks, 1978.)
MacPhee, A. W. 1 964. Cold-hardiness, habitat and winter survival of some orchard arthropods
in Nova Scotia. Can. Entomol. 96:617-625.
Mansingh, A. 1974. Studies on insect dormancy. II. Relationship of cold-hardiness to diapause
and quiescence in the eastern tent caterpillar, Malacosoma americanum (Fab.) (Lasio-
campidae:Lepidoptera). Canad. J. Zool. 53:629-637.
Rosenberg, N. J. 1974. Microclimate: The Biological Environment. John Wiley and Sons,
New York, New York, 315 pp.
Schaefer, M. 1976. An analysis of diapause and resistance in the egg stage of Floridia bucculenta
(Araneida:Linyphiidae). Oecologia (Berlin) 25:155-174.
Solomon, M. E. 1951. Control of humidity with potassium hydroxide, sulphuric acid or other
solutions. Bull. Entomol. Res. 42:543-554.
Washburn, E. W. 1928. International Critical Tables. National Res. Council. McGraw-Hill
Co., New York, Vol. 3, p. 303.
Wellington, W. G. 1950. Effects of radiation on the temperatures of insects and habitats. Sci.
Agric. 30:209-234.
Received July 22, 1982; accepted December 6, 1982.
NEW YORK ENTOMOLOGICAL SOCIETY
91(1), 1983, pp. 75-82
MELANISM IN PHIGALIA TITEA (CRAMER)
(LEPIDOPTERA: GEOMETRIDAE): A FOURTEEN-YEAR
RECORD FROM CENTRAL MASSACHUSETTS
Theodore D. Sargent
Department of Zoology, University of Massachusetts,
Amherst, Massachusetts 01003
Abstract.— melanic morph of Phigalia titea (Cramer), "Aeplorans'" (Franclemont, 1938),
comprised 1 8.7 percent of 3,906 specimens collected at various light sources at a single location
in central Massachusetts between 1968 and 1981. There were no discernible trends in melanic
frequency over that period. Three different light sources (incandescent, fluorescent and mercury
vapor) yielded similar estimates of melanic frequency. There was a decline in melanic frequency
late at night (after 2300 hours), and an increase in melanic frequency during the second half of
the flight season. The implications of these results are briefly discussed.
This report summarizes information on the incidence of melanism in
Phigalia titea (Cramer) (Lepidoptera: Geometridae) as determined by light-
source collecting at a single location in Leverett, Franklin County, Massa-
chusetts from 1 968-198 1 . Previous papers have dealt with part of this period
(1968-1973) (Sargent, 1971, 1974), and the present paper extends and elab-
orates on the overall record.
LOCALITY AND METHODS
Male P. titea were collected at several light sources about my home in
Leverett. (Female P. titea have only rudimentary wings, do not fly, and are
not included in any of the recorded data.)
My home is located in an extensive, mixed deciduous woodland domi-
nated by 40-year-old oaks {Querciis alba L. and Q. velutina Lam.) and
hickories {Carya ovata (Mill.) and C. glabra (Mill.)). Other trees with sub-
stantial representation include birches {Betula papyrifera Marsh, and B. lenta
L.), pine {Pinus strobus L.) and hemlock (Tsuga canadensis L.). Understory
shrubs include mountain laurel {Kalrnia latifolia L.), viburnums (esp. Vi-
burnum acerifolium L.) and blueberries (inc. Vacciniiim corymbosum L. and
V, vacillans Torr.), reflecting an acidic soil. The area shows little evidence
of air-borne pollution, as epiphytic lichens are common on tree trunks that
are themselves not noticeably darkened by soot.
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must therefore be hereby marked “Advertisement” in accordance with 18 U.S.C. §1734 solely
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76
NEW YORK ENTOMOLOGICAL SOCIETY
Fig. 1 . The typical (left) and melanic (right) morphs of P. titea. The upper specimens are
representative of the Leverett population. The lower specimens are an unusually dark typical
(left) and an unusually light melanic (right). Approximately life-size.
Four incandescent spotlights (Westinghouse outdoor projector, 1 50-watt),
a fluorescent blacklight tube (General Electric F15T8 BL, 15-watt), and a
medium-pressure, mercury-vapor discharge lamp (Atlas MB/U, 125-watt)
were all used at various times, and these light sources were carefully distin-
guished in the records. Generally, the lights were checked on several occa-
sions during the night, and the time, as well as the numbers of typical and
melanic P. titea present, were recorded. An effort was made to capture all
of the assembled moths on each visit to a light in order to minimize the
possibility of counting individuals on more than one occasion. The capture
moths were sacrificed, except for some that were color-marked and released
as part of a mark/recapture study during 1969 and 1970 (see Sargent, 1971).
Most collected P. titea were easily assignable to one of two morph cate-
gories: “typical” (light gray ground with strongly contrasting black lines), or
“melanic” {=''deplorans'\ Franclemont, 1938) (blackish overall, with only
a trace of the black lines). Occasional specimens, especially worn individuals,
were somewhat intermediate, but these were assigned to one or the other
category on the basis of whether the scales on the abdomen were light gray
(“typical”) or blackish (“melanic”) (Fig. 1).
(The division of specimens into only two categories on the basis of ap-
pearance is undoubtedly an oversimplification in terms of reflecting the
underlying genetic and/or environmental bases of melanism in this species.
VOLUME 91, NUMBER 1
77
Z
UJ
o
cr
LU
Q.
100
75-
50-
25-
Fig. 2. The total numbers of F. titea taken at light sources in Leverett, Massachusetts for
the years 1968-1981 (below), and the percentage melanic for each of these yearly samples
(above).
However, given our lack of knowledge regarding these underlying factors,
and further uncertainty regarding the effects of wear on phenotypic appear-
ance, it seems pointless to attempt a finer classification at this time.)
RESULTS
A total of 3,906 P. titea was taken at the various light sources over the
14-year period, 1968-1981. Of that total, 730 individuals, or 18.7 percent,
were melanic. The yearly sample sizes varied considerably, but the corre-
sponding percentages of melanics were relatively constant, and there were
no long-term trends of either increasing or decreasing melanic frequency
(Fig. 2).
Light sources. The three different light sources used in this study (incan-
78
NEW YORK ENTOMOLOGICAL SOCIETY
Table 1. The numbers of typical and melanic individuals of P. titea taken at various light
sources during periods of simultaneous operation.
Numbers
Year
Light sources
Typical
Melanic
p*
1971
Spotlights
118
24
>0.50
Blacklight
71
17
1979
Spotlights
652
110
>0.70
Blacklight
574
92
1972
Spotlights
22
5
Blacklight
41
15
>0.40
Mercury vapor
54
12
* Probabilities based on chi-square contingency tests.
descent spotlights, fluorescent tube, and mercury vapor bulb) were appar-
ently sampling identically, as the frequencies of typical and melanic P. titea
taken were not significantly different when compared for any of the sources
during periods of simultaneous operation (Table 1).
Time of night. In a previous paper (Sargent, 1 97 1 ), I suggested that melanic
P. titea might occur at their highest frequency during the early hours of the
evening. That suggestion was not supported by the more extensive data
reported here (Table 2). However, there was a decline in the frequency of
melanics taken late at night (after 2300 hr) (chi-square, 2X3 contingency
test == 5.94; P = 0.05), suggesting that melanics may have a somewhat dif-
ferent daily flight period (shorter, if not earlier) than typicals.
Seasonal effect. No seasonal effects were discernible in the data reported
previously with respect to the proportions of typical and melanic P. titea
taken (Sargent, 1971). Analysis of the current data reveals, however, that
significantly more melanics were taken over the second half of the season
each year (^^%,76s^ 20.9%) than were taken over the first half of the season
(^^%,079. 17.0%) (chi-square 2X2 test, P < 0.002) (Fig. 3B). This effect is
Table 2. The numbers of typical and melanic individuals of P. titea taken at various times
of night (based on data from nights when sampling was repeated no less than hourly) for the
six years when P. titea was most abundant at the Leverett location.
Time of night
Typical
Numbers
Melanic
% melanic
<2100 hr
586
141
19.4
2100-2300 hr
560
148
20.9
>2300 hr
267
45
14.4
VOLUME 91, NUMBER 1
79
2662 1185
2079 1768
DATES HALF SEASONS
Fig. 3. The percentage of melanic individuals of P. titea taken before and after April 13
(left) and during the hrst and second half of the flight season (right), summed for the years
1968-198 1 in Leverett, Massachusetts. The total numbers of individuals taken are given above
the bars.
not apparent when the data are analysed by calendar dates (Fig. 3A), sug-
gesting that the seasonal effect is not based on differential responses of the
two morphs to extrinsic environmental factors (e.g., temperature, daylength),
but rather reflects some intrinsic difference between the morphs with respect
to either their physiology (e.g., development rates, longevity) or their sus-
ceptibility to predation.
DISCUSSION
Aside from a single, very early record from West Roxbury, Massachusetts
(Minot, 1869), melanic P. titea were first reported from most localities in
eastern North America from about 1915-1960 (Owen, 1962). Minot de-
scribed his early specimen as a new species, P. strigataria, but Franclemont
(1938) correctly identified the melanic as a morph of P. titea, naming it
'"deplorans'\ Interestingly, by 1962, ""deplorans"' had not been noted in
certain industrial areas (e.g., Pittsburgh, Chicago) where melanic Piston cog-
nataria (Guenee) had already become quite common, and at that time it
was estimated that "'deplorans'' made up no more than 10 percent of any
P. titea population in North America (Owen, 1962). Few other data are
available until the present records which indicate that the melanic frequency
of P. titea in central Massachusetts has remained stable at close to 20 percent
of the population for the past 14 years.
80
NEW YORK ENTOMOLOGICAL SOCIETY
I have previously pointed out that the area in which these records have
been obtained shows little discernible evidence of industrial air pollution,
and Owen (1962) noted that P. titea was unusual in that the initial reports
of melanism were not from industrial centers. Thus, the bases for melanism
in P. titea remain obscure, although similar problems exist with respect to
explaining melanism in the closely related P. pilosaria (Schilf.) in Britain
(Lees, 1971, 1 98 1 ). It seems certain that factors other than selective predation
based on crypsis are involved in these cases, as they seem to be in Piston
in both the rural central Appalachians in this country (West, 1977) and
southern Britain (e.g.. Steward, 1977).
Evidence for different daily flight periods for the typical and melanic
morphs of a species has not, to my knowledge, been reported previously.
Such a difference would be detected only if sampling were carried out as
here with P. titea, i.e., at intervals throughout the daily flight period of the
species, with all of the moths collected at each sampling time. The signih-
cance of the apparently earlier flying of melanics than of typicals in P. titea
is not immediately apparent, though it may be related in some way to the
differences one would expect to hnd in the warming and cooling character-
istics of dark, as opposed to light, insects (see e.g.. Watt, 1968).
A tendency for melanics to become more common later in the season, as
reported here for P. titea, has been reported for other species as well. Thus,
Bishop, Cook and Muggleton (1978) described an abrupt increase in the
frequency of the melanic ""nigra'' morph of Gonodontis bidentata Clerk
towards the end of the season at two locations in northwest England. These
authors attributed this effect to a later emergence of the ""nigra" morph.
Increasing melanic frequency over the season has also been reported in
Cleora repandata L. in the Scottish Highlands, though in this case the effect
has been attributed to selective predation favoring the melanics (Kettlewell,
1973, p. 86).
There is some evidence to suggest that melanics may have slower devel-
opment rates than typicals in some species (e.g., Spodoptera exigua Hubn.
(Poitout, 1973), and Piston betularia (Kettlewell, 1973, p. 85)). There is no
evidence on this point for P. titea, but Lees (pers. comm.) has detected no
differences between the development rates of typical and melanic P. pilosaria
in his extensive rearing of that species.
Another factor that could lead to higher melanic frequencies later in the
season is an increased viability of melanics relative to typicals in the adult
stage. I am aware of no studies on adult viabilities in an industrial melanic
species, though there is evidence for a greater viability of the larvae of
melanics in certain cases (e.g., Menophra abruptaria Thunb. (Onslow, 1921),
and Cleora repandata {Yord, 1940)). Kettlewell (1973, p. 78), however, found
the evidence for differential viability of larvae of the morphs in industrial
melanic species ‘‘unconvincing”, and the matter needs further investigation.
VOLUME 91, NUMBER 1
The possibility remains that the increased frequency of melanic P. titea
later in the season is due to selective predation that favors the melanics. If
this were so, it seems unlikely that it would be based on crypsis, as the area
in which these studies were conducted is, as previously noted, visibly rural.
Furthermore, experimental tests have shown that melanic P. titea prefer
light backgrounds (Sargent, 1969), a tendency that would seemingly place
them at a cryptic disadvantage.
Other possible bases for differential predation on the morphs of P. titea
should be explored. Whittle et al. (1976) obtained evidence that bats take
more typical than melanic B. betularia, and this possibility should not be
overlooked in P. titea, particularly in light of the evidence that melanics of
this species may fly earlier in the evening or for a shorter period of time
than typicals. Other matters that should be investigated include the relative
palatabilities of typical and melanic P. titea to potential predators, and the
possibility of behavioral differences between the morphs in reacting to pred-
ator attacks.
LITERATURE CITED
Bishop, J. A., L. M. Cook and J. Muggleton. 1978. The response of two species of moths to
industrialization in northwest England. I. Polymorphisms for melanism. Phil. Trans.
Roy. Soc. London B 281:489-515.
Ford, E. B. 1940. Genetic research in the Lepidoptera. Ann. Eugen. London 10:227-252.
Franclemont, J. G. 1938. Description of new melanic forms (Lepidoptera: Geometridae,
Noctuidae and Arctiidae). Entomol. News 49:108-1 14.
Kettlewell, H. B. D. 1973. The Evolution of Melanism. Clarendon Press, Oxford, 424 pp.
Lees, D. R. 1971. The distribution of melanism in the pale brindled beauty moth, Phigalia
pedaria, in Great Britain. Pages 152-174 in: E. R. Creed (ed.). Ecological Genetics and
Evolution. Blackwell Scientific Publications, Oxford.
Lees, D. R. 1981. Industrial melanism: genetic adaptation of animals to air pollution. Pages
129-176 in: J. A. Bishop and L. M. Cook (eds.). Genetic Consequences of Man-Made
Change. Academic Press, New York.
Minot, C. S. 1869. American Lepidoptera 1. Geometridae Latr. Proc. Boston Soc. Nat. Hist.
13:83-85.
Onslow, H. 1921. Inheritance of wing colour in Lepidoptera VII. Melanism in Hemerophila
abruptaria (var. fuscata Tutt). J. Genet. 1 1:293-298.
Owen, D. F. 1961. Industrial melanism in North American moths. Amer. Nat. 95:227-233.
Owen, D. F. 1962. The evolution of melanism in six species of North American geometrid
moths. Ann. Entomol. Soc. Amer. 55:695-703.
Poitout, S. 1973. A study of melanism produced in the adult of Spodoptera exigua (Lepi-
doptera: Noctuidae) reared by strict brother-sister mating. Ann. Soc. Entomol. Fr. 9:
331-344.
Sargent, T. D. 1969. Background selections of the pale and melanic forms of the cryptic moth,
Phigalia titea (Cramer). Nature 222:585-586.
Sargent, T. D. 1971. Melanism in Phigalia titea (Cramer) (Lepidoptera: Geometridae). J. New
York Entomol. Soc. 79:122-129.
Sargent, T. D. 1974. Melanism in moths of central Massachusetts (Noctuidae, Geometridae).
J. Lepid. Soc. 28:145-152.
82
NEW YORK ENTOMOLOGICAL SOCIETY
Steward, R. C. 1977. Industrial and non-industrial melanism in the peppered moth, Biston
betularia (L.). Ecol. Entomol. 2:231-243.
Watt, W. B. 1968. Adaptive significance of pigment polymorphisms in Colias butterflies. I.
Variation of melanin pigment in relation to thermoregulation. Evolution 2:437-458.
West, D. A. 1977. Melanism in Biston (Lepidoptera: Geometridae) in the rural central Ap-
palachians. Heredity 39:75-81.
Whittle, P. D. J., C. Clarke, P. M. Sheppard and J. A. Bishop. 1976. Further studies on the
industrial melanic moth Biston betularia (L.) in the northwest of the British Isles. Proc.
R. Soc. London B 194:467-480.
Received July 28, 1982; accepted September 22, 1982.
NEW YORK ENTOMOLOGICAL SOCIETY
91(1), 1983, pp. 83-89
THE DIPTERA BREEDING ON SKUNK CABBAGE,
SYMPLOCARPUS FOETIDUS (ARACEAE)
David Grimaldi and John Jaenike
Department of Entomology, Cornell University,
Ithaca, New York 14853
Abstract.— larvae of 19 species of flies belonging to 7 families were found to feed in the
rotting portions of leaves, petioles, and flowers of skunk cabbage. Among flies bred from this
plant, drosophilids were by far the most numerous, but there is considerable geographic and
seasonal variation in composition and abundance of the fauna. The sex ratio of one species, a
Bryophaenocladius sp. (Chironomidae), reveals it to be parthenogenetic. The breeding biology
of the flies, with particular reference to other dipteran-aroid relationships, is reviewed.
Symplocarpus foetidus is a common perennial plant of wet lowland areas
in eastern North America. Inflorescences appear in early spring, the leaves
expand about one month later, and seeds are formed in the fall, when the
above-ground portion of the plant dies back. This species forms established
patches that may vary in size from a few square meters to several hectares.
Symplocarpus foetidus would thus seem to be an “apparent” plant to po-
tential consumers, and as such one might expect it to be protected by “quan-
titative” defenses (sensu Feeny, 1976). It is: the tissues contain barbed crys-
tals of calcium oxalate, which may deter a variety of herbivores (B. Meeuse,
pers. comm.). An herbivore that would cope with this defense, however,
would be rewarded with a predictable supply of food. Hence, specialization
of some herbivore species on S. foetidus might be expected to evolve.
Although a few Coleoptera, Psocoptera and Collembola breed in skunk
cabbages, the majority of insects bred from the plant are a variety of dip-
terans. We have never seen visible signs of serpentine or blotch mines,
chewing areas, or petiole girdling which might be indicative of insect activity.
All flies in this study were bred from rotting portions of plants and most
likely fed on bacteria harbored therein. This does not, however, rule out the
possibility that diptera may be the cause of decay in the portions of some
plants. We show here that Symplocarpus foetidus is probably the major
breeding site of the Holarctic species Scaptomyza graminum. Another sig-
nificant finding is that Drosophila recens, previously thought to breed ex-
clusively in mushrooms, also utilizes Symplocarpus as well.
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must therefore be hereby marked “Advertisement” in accordance with 18 U.S.C. §1734 solely
to indicate this fact.
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NEW YORK ENTOMOLOGICAL SOCIETY
METHODS
One collection of 22 flowers (including the spathe and spadix) was made
from two Symplocarpus patches on Deer Isle, Maine on June 28, 1982.
Three collections of 37, 27, and 28 plants were made on May 14, June 25,
and August 6, 1982, respectively, at Chenango Valley State Park, Chenango,
New York. Only flowers were collected in May, since rotting or damaged
leaves were not found. Rotting and damaged petioles and leaves, all from
different plants, comprised the specimens thereafter, as spathes were un-
common. Portions of plants were bagged and transported to the laboratory,
where they were placed on damp sand (except for the Maine collections,
which were placed on Instant Drosophila medium [Carolina Biol. Suppl.
Co.]) in 200 ml plastic containers covered with cardboard tops. To prevent
drying, the specimens were misted every other day with water. Adult insects
were aspirated as they emerged and were preserved in 70 percent ethanol
prior to identification.
Insects were identified using the following references: Wheeler (1960) and
Strickberger (1962) for Drosophila, Wheeler (1952) for Drosophilidae other
than Drosophila, Johannsen (1952) for Chironomidae and Ceratopogonidae,
Saether (1 973) for Bryophaenocladius sp., and Quate (1 960) for Psychodidae.
Scaptomyza ‘sp. A" appears to be a new species and will probably be described
later. Voucher specimens are deposited in the Cornell University collection,
Ithaca, New York. Our identification of Dasyhelea was confirmed by Dr.
Willis W. Wirth and the ephydrid and chloropids were identified by Drs.
W. N. Mathis and C. W. Sabrosky, respectively, all of the Systematic Ento-
mology Laboratory of the U.S. Department of Agriculture.
RESULTS AND DISCUSSION
The most conspicuous aspect of the breeding records (Tables 1 and 2) is
the complete absence of parasitic Hymenoptera: from a total of 1,132 po-
tential dipteran hosts reared (an average of 10 flies per plant part), not one
parasite emerged. This contrasts with the number of parasitic braconids and
cynipids bred from mushrooms containing Diptera also growing at the Che-
nango Valley site (Grimaldi, 1983). Carson et al. (1980) also bred numerous
small wasps from another primary dipteran breeding site, the aroid Calocasia
esculent a in Papua New Guinea.
Not so surprising is the seasonal variation in abundance of some flies,
since insect populations generally fluctuate in abundance seasonally. For
instance, none of the nematocera present during the spring and early summer
breed in Symplocarpus in late summer in New York. In contrast, Scaptomyza
graminum abundance increases almost four-fold towards late summer. Dro-
sophila recens, which is the most common fly in the spring collection, is
virtually nonexistent in the later collections. Of particular interest is the
VOLUME 91, NUMBER 1
85
Table 1. Drosophilidae reared from Symplocarpus foetidus (N = 672).
Numbers
Species
Males
Females
Total
Chenango Valley, New York, May 1982: flowers, 37 plants
Drosophila recens Wheeler
62
84
146
Scaptomyza graminum Fallen
11
15
26
Drosophila affinis-snhgroxxx)
5
15
20
Scaptomyza sp. A
8
4
12
Drosophila putrida Sturtevant
6
4
10
Drosophila falleni Wheeler
7
2
9
Drosophila palustris Spencer
1
4
5
Chymomyza amoena Loew
2
3
5
233
Chenango Valley, New York, June 1982: leaves
and petioles, 27 plants
Scaptomyza graminum
42
36
78
Drosophila palustris
1
6
7
Scaptomyza sp. A
2
1
3_
88
Deer Isle, Maine, June 1982: flowers, 22 plants
Drosophila quinaria Loew
17
20
37
Drosophila busckii Coquillett
2
3
5
Drosophila recens
1
1
2
44
Chenango Valley, New York, August 1982: leaves and petioles, 28 plants
Scaptomyza graminum
137
144
281
Scaptomyza sp. A
7
12
19
Scaptomyza paravittata Wheeler
3
4
7
307
geographic variation of species breeding in skunk cabbages. Drosophila qui-
naria, which is common in S. foetidus in Rochester, New York and Deer
Isle, Maine (Jaenike, 1978) is not present in the Chenango Valley population.
In the latter area, D. recens is the predominant species of Drosophila bred
from this plant.
The Araceae appears to be a pivotal resource in the ecological diversity
of the Drosophilidae. The niche is typical of Scaptomyza, which are leaf
miners. We have bred a large number of S. graminum {=S. borealis [Wheeler,
1981]) from skunk cabbages (this study and Jaenike, 1978). This drosophilid
has, however, been found, occasionally, to use some non-araceous resources,
such as Stellaria aquatica (Caryophyllaceae) in Japan (T. Okada, pers. comm.).
Nasturtium officinalis (Cruciferae) (Wheeler, 1952), Petasites officinalis
(Compositae), Brassica campestris, B. oleracea, B. rapa (Cruciferae), and
86
NEW YORK ENTOMOLOGICAL SOCIETY
Table 2. Non-drosophilid Diptera reared from Symplocarpus foetidus (N = 460).
Family
Species
Males
Numbers
Females
Total
Chenango Valley, New York May 1982: flowers, 37 plants
Psychodidae
Psychoda satchelli Quate
29
52
81
Psychoda alternata Say
2
4
6
Chloropidae
Tricimba lineella (Fallen)
94
Elachiptera costata (Loew)
11
Chenango Valley, New York, June 1982: leaves and petioles, 27 plants
192
Psychodidae
Psychoda satchelli
33
35
68
Ceratopogonidae
Dasyhelea oppressa Thomsen
31
36
67
Chironomidae
Bryophaenocladius sp.
0
29
29
Chloropidae
Tricimba lineella
2
166
Deer Isle, Maine, June 1982: flowers, 22 plants
Psychodidae
Psychoda satchelli
12
11
23
Ceratopogonidae
Dasyhelea opressa
9
3
12
Chloropidae
Elachiptera costata
18
Ephydridae
Athyroglossa granulosa (Cresson)
5
58
Chenango Valley, New York, August 1982: leaves and petioles, 28 plants
Cecidomyiidae
Dyodiplosis sp.
19
22
41
Ephydridae
Athyroglossa granulosa
3
44
Aquilegia vulgaris (Ranunculaceae) (Frost, 1923). Stalker (1945) has bred S.
graminum from Trifolium pratense leaves, but mentions this is not a major
resouree since so few individuals were reared. Stalker also found that S.
graminum in Rochester, New York, like the flies bred in this study, are most
abundant throughout the summer and early fall as found by sweep netting
in open areas. In addition, R. Lacy (pers. comm.) has bred 13 individuals
of S. graminum from a very extensive collection of mushrooms. Our data
show that a major larval resource of Scaptomyza graminum, perhaps the
most abundant species of its genus, is Symplocarpus foetidus.
In the Drosophila quinaria species group, some members, such as D.
quinaria and D. magnaquinaria, specialize on aroids (Jaenike, 1978; Whee-
ler, 1954; this study). Other species in the group (such as D. falleni, D.
phalerata, and D. transversa) breed in a diversity of mushrooms (Jaenike,
1978; Lacy, 1982; Grimaldi, 1983; Shorrocks, 1980). Drosophila recens ?ind
D. limbata are intermediate, breeding in both mushrooms and aroids. D.
limbata has been bred from Arum maculatum fruits (Schatzmann, 1977)
VOLUME 91, NUMBER 1
87
and from Russula mushrooms (Burla and Bachli, 1968) in Switzerland. D.
recens commonly breeds in mushrooms, but utilizes Symplocarpus in the
spring before mushrooms are fruiting.
Although the bulk of dipteran-aroid relationships in the tropics has yet
to be investigated, Heed (1957) gives an indication of the potential impor-
tance of this relationship in structuring Neotropical drosophilid commu-
nities. Approximately one-quarter of the 222 species of Drosophilidae col-
lected in El Salvador can be found in “aroid swamps,” of which Xanthosoma
is a characteristic plant. In Papua New Guinea, Okada and Carson (1979)
and Carson and Okada (1980) have found that several aroids are a “veritable
zoological garden” for drosophilids. Four drosophilid species were reared
from Colocasia esculenta, and 13 different species reared from Alocasia
macrorrhiza. The larvae of most of these species feed on spathes and spad-
ices, and are quite monophagous. Drosophila aproclinata, for example, breeds
only in decaying staminate portions of Alocasia hollrungii spadices. Like
most of the Drosophila breeding in Symplocarpus, Carson and Okada found
the New Guinea Drosophila limited to breeding in the flowers. It appears
that throughout the world the Araceae have repeatedly been invaded by
Drosophilidae.
For some drosophilids, Symplocarpus foetidus is just an incidental re-
source. For example, only 20 individuals of the extremely abundant Dro-
sophila affinis subgroup were bred in this study. Drosophila putrida, D.
falleni, D. busckii, and Chymomyza amoena are also, judging from their
numbers bred from other substrates and caught in banana traps, just inci-
dental rearings.
Another interesting parallel between New Guinea aroid-breeding flies and
the kind we have bred are the Psychodidae. For a family with larvae generally
breeding in detritus (Quate, 1960) such as compost and decaying organic
matter (Johannsen, 1934), aroids are an unusual niche. But, about one-third
of the non-drosophilid Diptera in Symplocarpus foetidus are Psychoda. Car-
son and Okada (1980) also found several psychodids breeding in Calocasia
esculenta. Quite unusual for cecidomyiids are our records for Dyodiplosis
sp. in Symplocarpus. Decaying vegetation may not be an unusual niche for
Dyodiplosis since it occasionally is found in some of our mushroom rearings.
Symplocarpus is not an unusual niche for the other nematocera and the
ephydrid and chloropids bred in this study. Ephydridae and Chloropidae
are commonly associated with wet areas, many members of both these
families boring the stems of plants. Although ceratopogonids and chiron-
omids are almost entirely aquatic groups, Dasyhelea and Bryophaenocladius,
which were bred from skunk cabbages, are among the more derived, ter-
restrial forms. Dasyhelea oppressa is common in bleeding exudates of Elm
( Ulmus) trees in New York (Thomsen, 1937). At least three Dasyhelea species
breed in cacti (Wirth and Hubert, 1960). Other ceratopogonids are quite
88
NEW YORK ENTOMOLOGICAL SOCIETY
terrestrial, larvae of Forcipomyia argenteola group species commonly breed-
ing in decaying portions of banana stems, cacao pods, and Calathea and
Heliconia bracts in the Neotropics (Wirth, 1982). Bryophaenocladius, to-
gether with Smittia and Pseudosmittia, belongs to a related group of Ortho-
cladiinae, some members of which Johannsen (1937) records as breeding in
and among plant roots. Parthenogenesis occurs repeatedly in the Chiron-
omidae, but is most frequent primarily in only the Orthocladiinae among
the eight subfamilies usually recognized. In the Orthocladiinae, species be-
longing to 8 genera, including Bryophaenocladius, are known to be obligately
or facultatively thelytokous (Scholl, 1960). Oliver et al. (1972) used emer-
gence trap data to show female-biased sex ratios occur in three additional
genera of this subfamily.
Most Diptera raised in this study, with the exception of some drosophilids
(and possibly chloropids), probably breed in plant matter other than decaying
Symplocarpus. Nematocera breed in detritus, on the whole, with several of
the species mentioned here adding the bacterial flora of decayed skunk
cabbage to their diet.
ACKNOWLEDGMENTS
We thank Drs. T. Okada and R. Lacy for use of their unpublished data presently being used
for a survey of drosophilid breeding sites. Beth Krause made comments on the manuscript.
This work was supported by a Grant-in-Aid-of-Research to DG by Sigma Xi and NSF Grant
No. DEB80-08574 to JJ.
LITERATURE CITED
Burla, H. and G. Bachli. 1968. Beitrag zur Kenntnis der schweizerischen Dipteren, insbe-
sondere Drosophila- \r\Qn, die sich in Fruchtkorpern von Hutpilzen entwickeln. Viert.
der Natur. Ges. Zurich 1 13:31 1-336.
Carson, H. L. and T. Okada. 1980. Drosophilidae associated with flowers in Papua New
Guinea I. Calocasia esculenta. Kontyu 47:15-29.
Feeny, P. 1975. Biochemical coevolution between plants and their insect herbivores. Pages
3-19 in: L. E. Gilbert and P. H. Raven (eds.), Coevolution of Animals and Plants. Univ.
of Texas Press, Austin.
Frost, S. W. 1923. A study of the leaf mining Diptera of North America. Cornell Agric. Exp.
Sta. Memoir 78.
Grimaldi, D. A. 1983. Ecology and competitive interactions of four coexisting species of
mycophagous Drosophila. M.A. thesis, St. Univ. of New York, Binghamton.
Heed, W. B. 1957. Ecological and distributional notes on the Drosophilidae (Diptera) of El
Salvador. Univ. Texas Publ. 5721:62-78.
Jaenike, J. 1 978. Resource predictability and niche breadth in the Drosophila quinaria species
group. Evolution 32:676-678.
Johannsen, O. A. 1934. Aquatic Diptera I. Nemocera, exclusive of Chironomidae and Cer-
atopogonidae. Cornell Agric. Exp. Sta. Memoir 164:3-70.
Johannsen, O. A. 1937. Aquatic Diptera III. Chironomidae: subfamilies Tanypodinae, Dia-
mesinae, and Orthocladiinae. Cornell Agric. Exp. Sta. Memoir 205:3-84.
VOLUME 91, NUMBER 1
89
Lacy, R. C. In press. Ecological and genetic consequences of mycophagy in Drosophilidae.
In: Q. Wheeler and M. Blackwell (eds.), Fungus-Insect Relationships: Perspectives in
Ecology and Evolution. Columbia U. Press, New York.
Okada, T. and H. L. Carson. 1979. Drosophilidae associated with flowers in Papua New
Guinea II. Alocasia (Araceae). Pac. Insects 22:217-236.
Oliver, D. R. and H. V. Danks. 1972. Sex ratios of some high arctic Chironomidae (Diptera).
Canad. Entomol. 104:1413-1417.
Quate, L. W. 1960. Guide to the insects of Connecticut. VI. Diptera. Seventh fasicle, Psy-
chodidae. Conn. St. Geol. Nat. Hist. Survey 92:1-32.
Saether, O. A. 1973. Four species of Bryophaenocladius Thien., with notes on other Ortho-
cladiinae (Diptera: Chironomidae). Canad. Entomol. 105:51-60.
Schatzmann, E. 1977. Friichte als natiirliche entwicklungs-substrate von Drosophiliden. Mitt.
Schw. Entomol. Gessell. 50:135-148.
Scholl, H. 1960. Die Oogenese einiger parthenogenetischen Orthocladiinen (Diptera). Chro-
mosoma 11:380-401.
Shorrocks, B. 1977. An ecological classification of European Drosophila species. Oecologia
26:335-345.
Stalker, H. D. 1945. On the biology and genetics of Scaptomyza graminum Fallen (Diptera,
Drosophilidae). Genetics 30:266-279.
Strickberger, M. W. 1962. A key to the United States species of Drosophila. Pages 1 1 1-122
in: M. W. Strickberger (ed.). Experiments in Genetics with Drosophila. Wiley, New York.
Thomsen, L. C. 1937. Aquatic Diptera V. Ceratopogonidae. Cornell Agric. Exp. Sta. Memoir
210:57-80.
Wheeler, M. R. 1952. The Drosophilidae of the Nearctic Region, exclusive of the genus
Drosophila. Univ. Texas Publ. 5204:162-218.
Wheeler, M. R. 1954. Taxonomic studies on American Drosophilidae. Univ. Texas Publ.
5422:47-64.
Wheeler, M. R. 1981. The Drosophilidae: a taxonomic overview. Pages 1-84 in: M. Ashburner
et al. (eds.). The Genetics and Biology of Drosophila, Vol. 3a. Academic Press, New
York.
Wirth, W. W. 1982. The cacao-pollinating midges of the Forcipomyia argenteola group (Dip-
tera: Ceratopogonidae). Proc. Entomol. Soc. Wash. 84:568-585.
Wirth, W. W. and A. A. Hubert. 1960. Ceratopogonidae (Diptera) reared from cacti, with a
review of the copiosus group of Culicoides. Ann. Entomol. Soc. Amer. 53:639-658.
Received November 15, 1982; accepted December 23, 1982.
NEW YORK ENTOMOLOGICAL SOCIETY
91(1), 1983, pp. 90-91
NOTES AND COMMENTS
PREDATION OF PROSIMULIUM MIXTUM/FUSCUM
(DIPTERA: SIMULIIDAE) COPULATING PAIRS BY
FORMICA ANTS (HYMENOPTERA: FORMICIDAE)
A comprehensive description of black fly predators, both vertebrate and
invertebrate, was given by Davies (Davies, D. M. 1981. Pages 139-158 in:
M. Laird (ed.), Blackflies. Academic Press, New York). While most predation
of black flies by Hymenoptera occurs by sphecid and vespid wasps, five
species of formicids have been observed preying upon black flies. Peterson
(Peterson, B. V. 1960. Can. Entomol. 92:266-274) lists Formica fusca L.,
F. obscuripes Forel and Myrmica brevinodes Emery. Peterson and Davies
(Peterson, B. V. and D. M. Davies. 1960. Can. J. Zool. 38:9-18) list Lasius
neoniger Emery and Myrmica emeryana Forel. Of these only L. neoniger
and M. emeryana were observed preying upon adult black flies.
On 5 May 1982, I observed large numbers of Formica subnuda Emery
preying on newly emerged Prosimulium males that were unwilling or unable
to fly when disturbed (air temp. = 18°C). The newly emerged males were
presumably drying their wings on the rocks and vegetation adjacent to the
Saranac River, Franklin Co., New York. On 12 and 13 May I observed
predation of mating pairs of Prosimulium mixtum/fuscum by Formica pod-
zolica Fraenkcoeur at a site 0.8 km upstream from the initial site. While
approaching this particular site, I had located a large swarm (estimated at
60 m X 8 m X 10 m high) of male Prosimulium black flies (at 3:00 p.m.,
ambient temperature = 24°C, RH = 37%) swarming in the leeward (3 km/
hr W) shaded sides of 2 20-m hemlocks {Tsuga canadensis) and 1 25 m
quaking aspen (Populus tremuloides) whose leaves had opened about 30-50
percent. Prosimulium mixtum/fuscum adults were dropping onto the road
surface in copula and remained attached for 0-5.75 min. Quite often the
larger female dragged the copulating male behind her as she walked along
the sandy road. A marauding ant randomly traversed the road until it crossed
the trail of a mating pair or came within 3-4 cm of an immobile pair. The
ant then turned abruptly in the same direction that the mating pair had
travelled, increased its pace, followed the same path the flies had taken and,
within 3-4 seconds of locating the pair, immobilized one or both of the flies.
On four separate occasions other ants that had been 20 cm or more from a
copulating pair when it crossed the flies’ trail responded similarly. Shorter
hunting trails were more common.
VOLUME 91, NUMBER 1
91
The F. subnuda that I observed on 5 May were attacking only the male
flies since the females had not yet emerged. The F. podzolica that were
observed on 1 2 and 1 3 May showed no apparent preference for the male or
female black fly. A few ants were observed dragging both male and female
carcasses that were still in copula. More frequently, an ant was successful in
killing only one fly. The other fly would either remain wounded and separate
immobilized from the mate or fly off unscathed.
Once an ant obtained its victim, it quickly turned back toward the colony.
With the captured fly (flies) supported by the ant’s mandibles, the ant com-
pletely ignored the trail of other mating pairs and other flies. On two oc-
casions, ants with flies clasped in their mandibles walked directly over mating
flies, which having been disturbed, separated and flew off. The F. podzolica
colony (ca. 40 cm diam) was in a loose sand area about 1.5 m off the vehicular
access zone on the south side of the road. On 1 3 May, ants from within the
colony were also observed to be discarding Prosimulium carcasses from the
colony interior to the area just outside (3-4 cm) the access holes. Since adult
flies had been emerging since 3 May, the duration of time within the colony
for these discarded Prosimulium is uncertain.
The author is indebted to Dr. John Simeone, SUNY College of Environ-
mental Science and Forestry, Syracuse, New York and to Dr. William Brown,
Department of Entomology, Cornell University, Ithaca, New York for iden-
tification of the ant specimens. I would also like to thank Dr. Charlie Morris,
NYS Department of Health, SUNY College of Environmental Science and
Forestry, Syracuse, New York for reviewing the ms.— Dennis J. White, New
York State Department of Health, Saranac Lake, New York 12983.
NEW YORK ENTOMOLOGICAL SOCIETY
91(1), 1983, pp. 91-92
THE FIRST OCCURRENCE OF CATORHINTHA MENDICA STAl
IN NEW ENGLAND (HEMIPTERA: COREIDAE)
The spread of the coreid bug Catorhintha mendica Stal from the Great
Plains eastward is, as noted by Hoebeke and Wheeler (1982) one of the few
well documented cases of such a range extension known in the Hemiptera.
On August 7, 1982 I collected a series of eight adults of this species from
a small patch of the host plant, Mirabilis nyctaginea (Michx.) MacMill.
92
NEW YORK ENTOMOLOGICAL SOCIETY
growing adjacent to the spillway of the large flood control dam in Mansfield
Center, Connecticut.
The collecting site is several miles from any railroad and the plants oc-
curred within a two square meter area. Hoebeke and Wheeler (1982) support
Baldufs (1957) belief that the spread of the plant has been essentially along
railroad rights of way. This is probably so and it suggests that both plant
and insect may have been present in Connecticut for some time. The col-
lecting site is some distance from any commercial center, several miles from
a railroad and is in the interior hill country of the eastern part of Connecticut.
Dowhan (1979) lists Mirabilis nyctaginea from Connecticut. Meyerhoff (pers.
comm.) informs me that it is not uncommon in suitable habitats in western
Connecticut but that he has not observed it east of the Connecticut River.
The presence of Catorhintha mendica in New England in a sense completes
its sweep eastward to the Atlantic coast. Hoebeke and Wheeler (1982) were
the first to provide an east coast record when they reported it from eastern
Long Island (Yaphank). They also listed several Pennsylvania records. Balduf
(1957) had also reported it from western Pennsylvania and Hoffman (1975)
from inland Virginia.
It gives me particular pleasure to be able to add this insect to the New
England Hemiptera fauna as the species was the subject of my first scientific
paper which was published just forty years ago (Slater, James A.
Slater, Section of Systematic and Evolutionary Biology, University of Con-
necticut, Storrs, Connecticut 06268.
LITERATURE CITED
Balduf, W. V. 1957. The spread of Catorhintha mendica StM (Coreidae, Hemiptera). Proc.
Entomol. Soc. Wash. 59:176-185.
Dowhan, J. J. 1979. Preliminary checklist of the vascular flora of Connecticut. Conn. Geol.
Nat. Hist. Surv. Rept. Investigations no. l:i-x, 1-176.
Hoebeke, E. R. and A. G. Wheeler, Jr. 1982. Catorhintha mendica, a Great Plains coreid now
established on the Atlantic Coast (Hemiptera: Coreidae). Entomol. News 93:
29-31.
Hoffman, R. L. 1975. The Insects of Virginia. No. 9 Squash, broad-headed, and scentless
plant bugs of Virginia (Hemiptera:Coreoidea, Alydidae, Rhopalidae). Virginia Polytech.
State Univ. Res. Div. Bull. 105:1-52.
Slater, J. A. 1 943. Developmental stages of Catorhintha mendica (Coreidae, Hemiptera). Bull.
Brooklyn Entomol. Soc. 38:1-5.
NEW YORK ENTOMOLOGICAL SOCIETY
91(1), 1983, pp. 93-96
BOOK REVIEW
The North American Grasshoppers. Volume I. Acrididae: Gomphocerinae
and Acridinae.— Daniel Otte. 1981. Harvard University Press, Cam-
bridge, Massachusetts. 275 pp. $45.00.
This book is the first volume of a projected three-volume work, the prin-
cipal purpose of which, according to its author, is “to aid in identifying all
described grasshopper species north of the Gulf of Panama, as well as those
in the West Indies”. It deals with two subfamilies of the family Acrididae,
the Gomphocerinae and Acridinae, which together comprise those grass-
hoppers best fitted to bear the name by virtue of the grassy habitats fre-
quented by most of the species. Throughout, the term “grasshopper” is taken
to mean the “short-horned” grasshoppers of the orthopteran suborder Cae-
lifera.
Taxonomic monographs tend to take one of two forms: the monographic
revision and the handbook. The first aims to be a comprehensive, archival
documentation, to be used by, and written in the language of, the specialist
researcher who may wish to carry investigation further in the particular field.
The second is intended as an identification manual and source of interesting
information for a variety of non-specialist readers. Daniel Otte has written
a book which falls between these two extremes, or rather displays some of
the characteristics of both. By employing a variety of novel and ingenious
devices, he has succeeded in meeting many of the requirements of both
groups of readers in a very pleasing format.
He starts with a general introduction in which mating behaviour, sound
production, and problems of identification of grasshoppers in general are
emphasised and structural details illustrated by clear and well labelled draw-
ings. Then follow an illustrated key to the families of North American
grasshoppers, an itemised diagnostic characterisation of the North American
acridid subfamilies that lack a prosternal spine, i.e., the Gomphocerinae,
Acridinae, and Oedipodinae, a general discussion of the Gomphocerinae
and Acridinae, and an illustrated key to their component genera.
The greater part of the book is then devoted to a systematic account of
the 42 genera and 1 24 species of Gomphocerinae and two genera and three
species of Acridinae recognised by Otte. The gomphocerine genera are ar-
ranged in 16 “genus groups”, which in two instances receive also tribal
names. The criteria used for distinguishing the categories genus group and
tribe, and their ranking relationship, are not stated. Under each genus group
containing more than one genus an itemised diagnosis of the component
genera is given, headed “Identification of Genera”. Each genus is then taken
up in turn, with sections on “Recognition”, “Identification of Species” (item-
ised diagnosis), and sometimes a key to the species. There follows individual
94
NEW YORK ENTOMOLOGICAL SOCIETY
treatment of each species, typically under the headings Distribution, Rec-
ognition, Habitat, Life Cycle, and References, often with text figures, and
including an excellent distribution map on a base of state boundaries, omit-
ting confusing topographic detail.
A major feature of the book is the 16 plates illustrating each species in
colour, often including both sexes and sometimes, in variable species, more
than one presentation to show the more distinctive of the alternative patterns.
This takes care of a rather general objection to coloured illustrations— that
they lead the reader to expect close conformity to the published figure,
whereas there may be wide divergence from it amongst individuals. How-
ever, it has been represented to me that the colours in many cases are too
weak. Another interesting feature of both the plates and some of the line
drawings in the text is the presentation in dorsolateral view. This has the
advantage of giving an impression of both the dorsal and lateral character-
istics in a single drawing, although for precise comparisons it cannot replace
separate dorsal and lateral views, especially since it is hardly possible to
maintain exactly the same orientation from drawing to drawing. The tech-
nique is feasible only in the hands of a skilled illustrator, such as Otte
obviously is, and is most appropriate for the less specialised reader.
Four valuable appendices enable the author to separate from the system-
atic treatment in the main part of the book those formal but essential details
concerning synonymy and type species of genera, and synonymy, generic
assignment, and type-specimen identity and location in species, which con-
cern principally the research specialist. They list also taxonomic changes
made for the first time in the book and the divergent subfamily assignments
of certain genera by the principal world monographers of recent years. The
justification for a number of new synonymies is not argued. Appendix 5, on
the pronunciation of generic names is, as the author admits, more contro-
versial, but may assist students. The book is completed by a glossary, a list
of references, and a taxonomic index.
I have some difficulty in interpreting the information given at the generic
level. The genus Ligurotettix will serve as an example chosen at random.
On p. 146 under “Identification of Genera” of the Cibolacris genus group,
four attributes of Ligurotettix are given. On p. 156, under “Recognition” of
the genus, several more are listed, but only one of the earlier four is repre-
sented. Presumably some difference is seen between “identification” and
“recognition”, but what is it? We are not told. There is now virtually no
disagreement that species should be treated as concrete populations whose
members are determined by relations of reproductive compatibility and
incompatibility. In the case of genera there are no such biological criteria,
although some would claim that “evolutionary” criteria are operational.
Important though the type species is in a nomenclatural context, it tells us
VOLUME 91, NUMBER 1
95
what species must not be excluded from the genus, not which species should
be included. The latter is approached nowadays with the aid of some form
of clustering procedure. Having obtained a more or less discrete cluster, we
search for attributes, common to all the species but not to related genera,
which would be jointly or severally diagnostic, and our success in this en-
terprise may determine how far we adhere to the clustering pattern in building
up a practical classification. But not every feature in common need be re-
garded as “necessary” to membership of the genus and is unlikely to be
“sufficient.” This situation has given rise to the distinction between diagnosis
and description of a genus. The diagnosis is a listing of those attributes
which, in the view of the taxonomist, would need to be possessed by any
species being considered for membership, and which at the same time dif-
ferentiate the genus from its near relatives. The description is a statement
of the attributes common to all the species and, if desired, of the range of
interspecific variation in other characters. The diagnosis is a characterisation
of the pigeon-hole, the description a characterisation of the present contents.
The distinction becomes of special importance in monotypic genera, where
the generic diagnosis must be carefully worked out so as not to incorporate
specific attributes that would be unlikely to be considered necessary in some
newly discovered candidate for membership of the genus. I do not believe
that this distinction corresponds to Otte’s “identification” and “recogni-
tion”, both of which seem to combine elements of both diagnosis and de-
scription, though leaning more towards the second.
Similar uncertainties are encountered at the subfamily level. Nearly all
the genera are endemic to the region, but this is not true of the subfamilies.
It must be understood, and was certainly in the mind of the author, that the
characterisations and keys will not necessarily work on a world basis. Otte’s
comments on problems of subfamily classification in the Gomphocerinae
and Acridinae, while familiar outside the subject area, are well supported
by his discussion on p. 17. However, the problem is not simply where to
place individual genera: it extends to the question of whether the two taxa
merit separation at subfamily level. Moreover, the same question arises with
respect to the subfamily distinctness of the Acridinae and Oedipodinae con-
cerning which, along with the ad hoc Truxalinae (extralimital), controversy
and vacillation involving Dirsh and Uvarov continued for years. The case
of Stethophyma, discussed on p. 208 (see also pp. 1 7 and 34) illustrates the
situation very well. This genus not only lacks the principal diagnostic feature
of the Gomphocerinae (the femoral pegs), but it possesses the principal
feature of the Oedipodinae (the intercalary vein); yet Otte tentatively places
it in the Gomphocerinae because of its “behaviour and appearance”.
It is a tribute to the publishers that my (admittedly not completely ex-
haustive) reading of this book has not brought to light a single printing error.
96
NEW YORK ENTOMOLOGICAL SOCIETY
However, attention may be directed to a few minor departures from accepted
terminology and to a few minor factual errors. In Figure 5, on p. 7, the scape
of the antenna and the flagellum are shown and labelled, but not the pedicel,
and in the glossary, p. 253, it is incorrectly stated that the flagellum is the
“main portion of the antennae excluding the basal segment or scape”. In
Figure 8, p. 9, the terminology of the areas and carinae of the hind femur
is unconventional and will not be readily understandable to the specialist;
the conventional terminology can be found in many readily available works,
including those of Rehn, Dirsh, and Uvarov. In Figure 6, p. 8, the structure
labelled “preocular ridge” is conventionally termed the “lateral facial Cari-
na”; the “lateral foveolae” are the “temporal foveolae” (see also p. 253).
On p. 254, the definitions of “sternum” and “sternite” are misleading. A
sternum is the ventral part of a body segment; a sternite is a sclerotised plate
occupying the whole or part of a sternum; since in Acrididae the whole of
each sternum is sclerotised, the two terms are interchangeable (the same
applies to “tergum”, “tergite”). Also on p. 254, under “Subgenital plate”:
The term is applied in both sexes. In the female it is the eighth sternum (or
sternite), in the male the “ninth sternal lobe”, which articulates on the main
part of the sternum. In the “Key to North American Orthopteroid Insects”,
the Acridoidea are shown as having a tympanum, whereas on p. 9 it is
correctly stated that the tympanum may be absent. On p. 253, under “Para-
type”: According to the Glossary of the International Code of Zoological
Nomenclature, a paratype is “every specimen in a type-series other than the
holotype”; according to Article 72(b) of the Code, a type-series of a species
“consists of all the specimens on which its author bases the species, except
any that he refers to as variants, or doubtfully associates with the nominal
species, or expressly excludes from it” — i.e., the author does not have to
specifically “designate” a paratype. On p. 215, under (1), for “tibiae” read
“femora”— a lapsus calami. On p. 208 the question of the type species of
Stethophyma and Mecostethus is presented as though it were open to Otte
to make his own decisions, whereas the valid type species must be deter-
minable under the provisions of the International Code or by ruling of the
International Commission on Zoological Nomenclature.
In summary. Dr. Otte has produced a useful, attractive, and very readable
account of the two subfamilies he deals with. The few minor blemishes
should be avoidable in the second and third volumes and in any new edi-
tion.—K. H. L. Key, Division of Entomology, C.S.I.R.O., Canberra, Austra-
lia.
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Journal of the
New York Entomological Society
VOLUME 91 MARCH 1983 NO. 1
CONTENTS
A revision of the genus Linens StM (Hemiptera: Pentatomidae: Discocephalinae:
Ochlerini) L. H. Ralston 1-47
On the biology and food plants of Lygaeus turcicus (Fabr.) (Hemiptera: Lygaeidae)
James A. Slater 48-56
The small milkweed bug, Lygaeus (Hemiptera: Lygaeidae): milkweed specialist
or opportunist? A. G. Wheeler, Jr. 57-62
A new species of Cleradini (Hemiptera: Lygaeidae: Rhyparochrominae) from the
Central African Republic and Ghana B. J. Harrington 63-67
Overwintering egg mass adaptations of the eastern tent caterpillar, Malacosorna
americanum (Fab.) (Lepidoptera: Lasiocampidae)
Alejandro Seggara Carmona and Pedro Barbosa 68-74
Melanism in Phigalia titea (Cramer) (Lepidoptera: Geometridae): a fourteen-year
record from central Massachusetts Theodore D. Sargent 75-82
The Diptera breeding on skunk cabbage, Symplocarpus foetidus (Araceae)
David Grimaldi and John Jaenike 83-89
Notes and Comments
Predation of Prosimulium mixtum/fuscum (Diptera: Simuliidae) copulating pairs by
Corm/ciz ants (Hymenoptera: Formicidae) Dennis J. White 90-91
The first occurrence of Catorhintha mendica Stal in New England (Hemiptera:
Coreidae) James A. Slater 91-92
Book Review
The North American Grasshoppers. Volume I. Acrididae: Gomphocerinae and
Acridinae A'. H. L. Key 93-96
JUNE 1983
No. 2
Vol. 91
Journal
of the
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Entomological Society
(ISSN 0028-7199)
Devoted to Entomology in General
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY
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Incorporating The Brooklyn Entomological Society
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Joseph Cerreta, Columbia University, New York; Durland Fish, Ford-
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NEW YORK ENTOMOLOGICAL SOCIETY
91(2), 1983, pp. 97-176
A REVISION OF THE GENUS ACROSTERNUM FIEBER,
SUBGENUS CHINA VIA ORIAN, IN THE WESTERN
HEMISPHERE (HEMIPTERA: PENTATOMIDAE)
L. H. Rolston
Department of Entomology, Louisiana Agricultural Experiment Station,
Louisiana State University, Baton Rouge, Louisiana 70803
Abstract.— Tht American species of Acrosternum Fieber, 1860, are placed in the subgenus
Chinavia Orian, 1965, the latter taxon being elevated from synonymy. Chinavia is redefined,
a key provided for the 51 nominal species of the Western Hemisphere and a description or
diagnosis provided for each species. One nominal species is removed from Acrosternum. The
species cataloged by Kirkaldy as Nezara {Acrosternum) hebes Bergroth, 1891, is a junior syn-
onym of Piezodorus guildinii (Westwood, 1837). Additional new synonymy recognized is:
Nezara nigritarsis Stal, 1872, as a junior synonym of Acrosternum {Chinavia) grave (Walker,
1867). Species described in the genus Nezara and here given new combinations as species of
Acrosternum subgenus Chinavia are: fuscopunctatum (Breddin, 1901), longicorialis (Breddin,
1901), nigridorsatum {Breddin, 1901), nigropictum (Breddin, 1906) panamensis {Distant, 1890)
and rogenhoferi (Stfil, 1872). The following 23 species are new: aseadum, australe, bellum,
brasicola, callosum, collis, ecuadorensis, esmeraldum, euri, froeschneri, insulani, istum, rnac-
donaldi, monticola, occasi, occultum, pecosum, pengue, plaumanni, simplicis, teretis, ubicum
and wygodzinskyi.
INDEX TO SPECIES
abnorme {Berg, I S9 2) 113
apicicorne {SpinolsL, 1852) 149
armigerum (Sial, 1859) 108
aseadum Rolston, new species 132
australe Rolston, new species 150
bellum Rolston, new species 1 10
bipunctulum \%11) 166
brasicola Rolston, new species 157
callosum Rolston, new species 1 1 3
collis Rolston, new species 148
dallasi (Distant, 1900) 165
dijfLcile iSxkl, 1860) 123
ecuadorensis Rolston, new species 134
The publication costs of this article were defrayed in part by page charge payment. This article
must therefore be hereby marked “Advertisement” in accordance with 18 U.S.C. §1734 solely
to indicate this fact.
98
NEW YORK ENTOMOLOGICAL SOCIETY
esmeraldum Rolston, new species 120
erythrocnemis (Berg, 1878) 1 6 1
euri Rolston, new species 140
froeschneri Rolston, new species 1 17
fuscopunctatum {Brtddin, 1901) 152
geniculatum 1851) 127
gra ve (Walker, 1867) 1 2 6
herbidum (Sik\, 1859) 169
hi lare {Say, 1831) 155
irnpicticorne {SXa\, \S7 2) 167
insulani Rolston, new species 1 1 1
istum Rolston, new species 121
laetum {SXa\, \S59) 137
longicorialis {Brtddm, 1901) 124
macdonaldi Rolston, new species 142
(Palisot de Beauvois, 1805) 152
monticoia Rolston. new species 144
montivagum (DisXanX, 1890) 147
musivurn (Berg, 1878) 160
napaewn (StM, 1872) 123
nigridorsatum (Breddin, 1901) 133
nigropictum (Breddin, 1906) 128
obstinatum {SXaX, 1860) 1 16
occasi Rolston, new species 145
occiillum Rolston, new species 1 1 5
panamensis (Distant, 1 890) 1 26
pecosurn Rolston. new species i 32
pengue Rolston, new species 170
pennsylmnicum (Palisot de Beauvois, 1 805) 1 30
plaumanni Rolston. new species 1 63
rogenhoferi (Stal, 1872) 1 60
runaspis {DaWas, 1851) 162
scutcHatiim (Distant, 1890) 1 1 7
simplicis Rolston, new species 1 38
sparnium (Dallas, 1851) 1 1 6
teretis Rolston, new species 1 58
ubicum Rolston, new species 1 35
viridans (Stal, 1 859) 1 28
wygodzinskyi Rolston, new species 142
The status and bounds of Acrosternum were especially uncertain for several
decades after Fieber (1860) proposed the genus. The principal difficulty lay
VOLUME 91, NUMBER 2
99
in distinguishing between Acrosternum and Nezara StM, and even so astute
an hemipterist as Stal wavered in his concept of these genera (1865, 1872,
1876). Other eminent hemipterists of the late 19th and early 20th centuries
(Berg, Breddin, Distant, Horvath, Jakovlev, Schouteden, etc.) misunder-
stood Fieber’s genus for more than half a century after it was proposed and
described over a score of Acrosternum species in Nezara.
Sharp (1890) began the process of differentiating more clearly between
Acrosternum and Nezara. After studying the male genitalia of Acrosternum
acutum (Dallas), A. marginatum (Palisot de Beauvois) and Nezara viridula
(L.), he remarked on the similarity of the first two species— even though the
first was African and the second American— and doubted that the three
species should be in the same genus. Nevertheless, he left them all in Nezara.
Bergroth (1914) drew on Sharp’s work and the external form of the meta-
thoracic scent glands to distinguish between these genera. He transferred to
Acrosternum those African species of the genus that StM (1876) had included
in Nezara and all the American species (except viridula) that Kirkaldy ( 1 909)
had cataloged under the “typical subgenus” of Nezara.
The work of Freeman ( 1 940) apparently disposed of any reservations about
the biological validity of Acrosternum or its proper rank. He began his
revision of Nezara by characterizing the genus and stressing the value of the
parameres in differentiating among the genera Nezara, Acrosternum, Pellaea
and Banasa.
Although the validity and status of Acrosternum were now established,
the limits of the genus were not. Orian (1965) proposed the genus Chinavia,
with Rhaphigaster pallidoconspersum Stal as type species, for nine species
from Africa, Madagascar and Mauritius that had been previously included
in Nezara or Acrosternum, and he reserved Acrosternum for A. heegeri
Fieber, the type species, and unspecified species from the Palearctic-Med-
iterranean region. He relied primarily upon characters of the male genitalia
to distinguish Chinavia from Acrosternum. Day (1965) supported Orian’s
proposal but Linnavuori (1972) did not and sank Chinavia in the synonymy
of Acrosternum. Nevertheless, Linnavuori recognized two groups of species
within Acrosternum, the heegeri-gvoup and the pallidoconspersum-group,
which correspond to Orian’s concept o^ Acrosternum and Chinavia, respec-
tively.
In the Eastern Hemisphere, Acrosternum is represented in most if not all
of Africa, in the Mediterranean basin and eastward into Madagascar, the
Mascarine Islands, the Seychelles Islands, Arabia, southern Russia and the
Indian subcontinent. In the Western Hemisphere, representation occurs from
Patagonia in Argentina to southern Canada in the Great Lakes region (On-
tario and Quebec). The northern limit of distribution is about the 45th
parallel in both hemispheres. There are numerous species in both hemi-
spheres, the largest number being American.
The heegeri-gvowp is confined to the Eastern Hemisphere, ranging from
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NEW YORK ENTOMOLOGICAL SOCIETY
the Cape Verde Islands and Canary Islands in the west through the Medi-
terranean basin and northern Africa into Arabia, southern Russia and the
Indian subcontinent. Linnavuori (1972) speculates that the heegeri-gvoup of
species are the products of a radiation from the Sudanese subregion. This
group is apparently adapted to arid and semiarid conditions.
Most of the species studied by Linnavuori belong to the pa/lidoconsper-
surn-group. Their distribution in the Eastern Hemisphere is primarily trop-
ical in Africa, Madagascar, Mauritius and Seychelles. It is to this group that
all American species belong.
From my examination of heegeri, graminea (Fabricius), and millieri (Mul-
sant and Rey) of the heegeri-growp and acutum (Dallas), pallidoconspersiim
(Stal), and punctatorugosum (Stal) of the paUidoconspersiim-group, as well
as all American species, which also belong in the latter group, I conclude
that Chinavia deserves a better fate than synonymy. Pending a more detailed
study of the Eastern Hemisphere species than has so far been undertaken, I
propose to use Chinavia as a subgenus of Acwsternum, placing therein all
American species of Acrosternum as well as the species assigned to Chinavia
by Orian (1965).
The subgenera Acrosternum and Chinavia differ in several respects. In the
females of the 18 American and three African species of Chinavia that were
dissected, the spermathecal bulb and pump are remarkably similar. The bulb
is always spherical with two long, thin, opposed diverticula that usually bend
in opposite directions (Fig. 1 1). The duct at the base of the proximal flange
is moderately expanded. In the nominal subgenus the spermathecal bulb
varies in shape. It is essentially digitiform, without elaboration in heegeri
(Fig. 1) but with rudimentary diverticula in millieri and graminea (Figs. 2,
3, 5). The duct at the base of the proximal flange is greatly expanded in all
3 species, and proximad of this expansion it is convoluted in heegeri and
graminea.
The proctiger of Chinavia is rather flattened dorsally with a median, lon-
gitudinal impression that becomes pronounced before ending subbasally. In
the nominate subgenus the proctiger bears a pair of prominent, conical
projections subapically on the dorsal surface which otherwise is transversely
convex.
The parameres in Chinavia have a basal spur and distally project into the
genital cup (Fig. 10). In the nominate subgenus the basal spur is absent and
the part of the parameres within the genital cup are bent sharply and ap-
pressed to the anterior wall of the genital cup (Fig. 4).
The mesosternal carina in Chinavia is low, conspicuously widened ante-
riorly and evanescent posteriorly. In the nominate subgenus this carina runs
the length of the mesosternum and is about the same width and height
throughout.
VOLUME 91, NUMBER 2
101
In general, the species of Chinavia are larger and darker in color than the
species of the nominate subgenus.
There is no doubt, based on the species studied, that these two groups of
species differ substantially. Whether or not these differences converge when
other species of the Eastern Hemisphere are examined critically remains to
be seen. The classihcatory signihcance of the observed differences is, of
course, a matter of subjective interpretation.
DISPOSITION OF SPECIES REMOVED FROM Acrostemum
One nominal species is removed from Acrostemum. The species cataloged
by Kirkaldy (1 909) as Nezara {Acrostemum) hebes Bergroth, 1 89 1 , is a junior
synonym of Piezodorus guildinii (Westwood, 1837). The type of N. hebes,
which is in the Museum National d’Histoire Naturelle, Paris, was examined.
PARATYPE DEPOSITORIES
Paratype depositories are indicated by the following abbreviations:
AMNH
CAS
DBT
FSCA
HDE
INHS
JG
LHR
MAI
OSU
RNH
TAMU
UCD
USNM
American Museum of Natural History
California Academy of Sciences
Donald B. Thomas collection
Florida State Collection of Arthropods
H. Dodge Engleman collection
Illinois Natural History Survey
Jocelia Grazia collection
author’s collection
M. A. Ivie collection
Ohio State University
Rijksmuseum van Natuurlijke Historie
Texas A&M University
University of California at Davis
U.S. National Museum
Acrostemum Fieber, 1860
Acrostemum Fieber, 1860, p. 79; StM, 1876, p. 90; Bergroth, 1914, p. 25
(diagnosis); Freeman, 1940, pp. 351-352 (cf Nezara); Cachan, 1952, pp.
445-446 (Madagascan spp. keyed); Wagner, 1959, pp. 4 1 3-4 15,417 (Pale-
arctic spp. keyed); Finnavuori, 1972, pp. 416-420 (synonymy, African
spp. keyed).
Nezara: StM, 1865, pp. 196-198 (African spp. keyed section bb); StM, 1876,
pp. 91-92 (African spp. keyed section aa); Sharp, 1890, pp. 406-408, pi.
13, hgs. 11a, lib, 12, 17 (6 genitalia).
Nezara {Acrostemum): Stal, 1872, pp. 41-43 (American spp. keyed).
102
NEW YORK ENTOMOLOGICAL SOCIETY
Figs. 1-6. Fig. 1. A. heegeri. Distal part of spermatheca. Figs. 2-4. A. millieri. 2. Distal part
of spermatheca. 3. Spermathecal bulb and distal flange, end view. 4. Paramere. Fig. 5. A.
graminea. Distal part of spermatheca; spermathecal bulb (SB); distal flange (DF). Fig. 6. A.
montivagum. Length of ostiolar ruga relative to distance from mesial margin of ostiole to lateral
margin of thorax.
Acrosternurn subgenus Chinavia Orian, 1965
Chinavia Orian, 1965, p. 25; Day, 1965, pp. 559, 565 (Madagascan spp.
keyed).
Acrosternurn pallidoconspersum-group: Linnavuori, 1972, pp. 416-418.
Diagnosis. Median projection on sternite 3 (2nd visible) ranging in length
from tubercle not quite attaining metacoxae to spine surpassing mesocoxae
VOLUME 91, NUMBER 2
103
but not reaching procoxae, usually compressed unless rudimentary, rarely
circular in cross-section. First antennal segment not reaching apex of head.
Bucculae percurrent caudad of obtuse anterior tooth, evanescent at base of
head. Rostrum usually terminating between metacoxae, rarely at mesocoxae
or on sternite bearing median projection. Mesosternal carina low, broadened
anteriorly, evanescent posteriorly. Metasternum flat or slightly convex, with-
out marginal rim, not produced ventrad. Femora unarmed. Ostiole on each
side on plane of metasternum; attending sulcus and ruga elongated, latter
reaching about one-half to three-fourths of distance from mesial margin of
ostiole to lateral thoracic margin (Fig. 6), shorter only in A. abnorme.
Spermathecal bulb with two long slender diverticula (Fig. 1 1); expansion
of spermathecal duct at proximal flange with much smaller diameter than
bulb. Spiracles present on 8th paratergite (Fig. 22). Genital cup lacking
tubercles on lateral walls, rarely with carina; posterior wall prominent, emar-
ginated mesially, occasionally reduced to stout projection (Fig. 37); dorsal
rim of posterior wall oriented diagonally or curved, usually denticulate and/
or toothed (Figs. 9, 19), rarely covering genital cup except above proctiger
(Fig. 60).
Large species about 10-18.5 mm long excluding membranes of hemelytra.
Color predominantly green, sometimes blended with yellow; outer margins
of body usually bordered in yellow to crimson; minor black markings usually
present.
CHARACTERS OF INTERSPECIFIC VALUE
The pygophore is one of the taxonomically most useful characters, espe-
cially the structure which here is termed the posterior wall of the genital cup
and which may be homologous with the inferior ridge. The posterior wall
is deeply emarginated or divided mesially and the part on each side variously
oriented when viewed dorsally. Usually the dorsal rim is diagonal, running
anteromesad from near the posterolateral angle of the pygophore (Fig. 1 9)
but sometimes slightly curved (Fig. 9) or reduced to a stout projection (Fig.
37). The dorsal rim of this structure is variously shaped and armed and
usually constant intraspecifically, although notable variation occurs in A.
ubicum (Figs. 95-97). In A. longicorialis the dorsal wall is reflexed cephalad
and covers the genital cup except above the proctiger (Fig. 60). The parameres
follow one general plan, and the slight differences among species are not
often diagnostic.
As usual among pentatomids, the female genital plates are not always
taxonomically useful, and those differences that do occur among species are
usually more subtle than in the male genitalia. However, the outline of the
posterior margin of the basal plates may differ appreciably from the cau-
doventral view, i.e., with the anterior and posterior margins of the plates
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NEW YORK ENTOMOLOGICAL SOCIETY
approximately on the same focal plane. From a caudal view the dorsal edge
of the plates may also differ in form among species. For example, A. mar-
ginatum and A. hilare females are easily separated by the basal plates (Figs.
157, 158, 164, 165). The spermathecae of all species in which this structure
has been examined are similar. Insufficient specimens have been examined
to determine whether or not the differences observed among species are
constant.
The humeri are usually rounded from the dorsal view and little or not at
all produced laterad. However, a few species have the humeri produced
laterad in a right angular to spinose projection. Such species do not form a
phylogenetic group, but the humeral character is nonetheless useful in rec-
ognizing species.
The length of the median projection at the base of the abdominal venter
seems to vary little intraspecifically. Length ranges from a rudimentary tu-
bercle to a spine reaching to or a little past the anterior limit of the mesocoxae.
The spine is usually compressed but in A. froeschneri it is nearly round in
cross-section.
The rather inconspicuous black markings on the cicatrices, basal angles
of the scutellum and connexiva are usually constant intraspecifically, par-
ticularly those on the connexiva. A. laetum is exceptional in the amount of
variation in the connexival markings. Interspecifically the connexiva may
be immaculate, or the black mark confined to the edge of the sternite at the
posterolateral angle (Fig. 34), or spill onto the laterotergite (Fig. 33), or cover
the posterolateral angle of one segment and the anterolateral angle of the
following segment (Fig. 40), or border most or all of the posterior margin
of each segment (Fig. 26) or border the connexival segments on both sides
of the transverse sutures.
The color markings that occur on the femora or both femora and tibiae
of a few species are diagnostically useful.
Many species have each spiracle located on a small, differentially colored
callus. In A. australe the spiracle is contiguous with but not on the callus.
Usually the callus is ivory colored, but in A. collis it is black. Occasionally
the area around each spiracle is more or less ivory colored but not calloused.
The color of the peritreme is diagnostically helpful but unfortunately variable
in some species.
Generally the costal angle of each corium lies over the penultimate con-
nexival segment, terminating intraspecifically from the middle of the segment
to the posterior margin. A few species, however, have longer coria and the
costal angle lies over the last connexival segment. Usually the extremity of
the coria is at or very near the costal angle (Fig. 57), but A. longicorialis is
exceptional in that the corium of each hemelytron intrudes into the mem-
brane and leaves an appreciable amount of membrane along the costal mar-
gin cephalad of the extremity of the corium (Fig. 58).
VOLUME 91, NUMBER 2
105
The rostrum generally terminates between the metacoxae, but in A. mon-
tivagum and A. pennsylvanicum it does not reach the metacoxae and in A.
viridans it extends onto the second visible sternite. This character seems
intraspecifically constant but is, of course, altered in specimens with the head
abnormally positioned.
KEY TO SPECIES OF Acrosteruum
1 . Humeri produced laterad of each corium by more than half the width of an eye,
right angular or spinose 4
Humeri not or less produced, not spinose, usually rounded 2
2(1). Abdominal spine projecting past anterior limit of metacoxae 6
Abdominal spine or tubercle not surpassing metacoxae 3
3(2). Lateral half or more of transverse connexival sutures (entire suture, not just ex-
posed part if partially covered) bordered on one or both sides in black (Fig. 26) ... 47
Connexiva immaculate or with black spot or macule at posterolateral angles of
segments (Figs. 33, 34, 40) 18
4(1). Abdominal spine projecting slightly past anterior limit of mesocoxae; transverse
connexival sutures bordered on both sides with black insulani, new species
Abdominal spine not surpassing metacoxae; connexiva with black dot at pos-
terolateral angles of segments 5
5(4). Lateral margins of head anterior to eyes tapering abruptly to narrow apex of head
(Fig. 12); pygophore moderately emarginate from caudal aspect (Fig. 14)
bellum, new species
Head less tapered, apex moderately rounded (Fig. 7); pygophore deeply emarginate
from caudal view (Fig. 8) armigerum St^l
6(2). Ostiolar ruga not much longer than ostiole abnorme (Berg)
- Ostiolar ruga reaching about halfway or more from mesial margin of ostiole to
lateral margin of thorax 7
7(6). Posterior margin of connexival segments bordered in black 8
- Connexiva with large black macules, each divided by transverse suture, or with
small black mark in posterolateral angle of each segment, or immaculate 10
8(7). Abdominal spine reaching just past metacoxae herbidum (St^l)
Abdominal spine reaching to or beyond posterior margin mesocoxae 9
9(8). Sternites slightly calloused on both sides of transverse sutures and pseudosutures
callosum, new species
Abdominal sternites lacking calli occultum, new species
10(7). Abdominal spine projecting past mesocoxae 1 1
Abdominal spine reaching no farther that anterior margin of mesocoxae 1 2
11(10). Lateral margins of juga black, submarginally yellow banded sparnium (Dallas)
Lateral margins of juga not or thinly edged in black; submarginally green or weakly
suffused with yellow obstinatum (St^l)
12(10). Connexival spots large, each divided by transverse sutures scutellatum (Dallas)
- No more than posterolateral angles of connexival segments black 13
13(12). Abdominal spine conical, round in cross-section fweschneri, new species
Abdominal spine compressed 14
14(13). Distal margin of each corium parabolic, intruding into membrane (Fig. 58); ab-
dominal spine reaching anterior limit of mesocoxae longicorialis (Breddin)
Little or none of membrane laterad of coriaceous part of hemelytra (Fig. 57);
abdominal spine terminating near posterior limit of mesocoxae 1 5
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NEW YORK ENTOMOLOGICAL SOCIETY
15(14). Lateral jugal margins barely concave before eyes (Fig. 43) 16
- Lateral jugal margins clearly concave before eyes (Figs. 50, 52) 17
1 6( 1 5). Black on connexiva confined to edge of stemite at posterolateral angle of segments;
outer margins of dorsum weakly and incompletely bordered in yellow
esmeraldum, new species
- Black spot in posterolateral angle of connexival segments including part of latero-
tergite; outer margins of dorsum clearly bordered in reddish orange
istum, new species
17(15). Broad submarginal depression present dorsally along anterolateral margins of
pronotum napaeum (Stdl)
- Transverse curvature of pronotal disk continuing smoothly to anterolateral mar-
gins dijficile (Stal)
1 8(3). Distal end or more of femora black at least on superior surface; antennae entirely
black 1 9
Distal end of femora green, yellow or red; much or all of antennae usually green
19(18). Large black macule on dorsum comprised of distal part of each corium and
hemelytral membranes panamensis (Distant)
Dorsum not so marked, macules absent or on pronotum and scutellar base 20
20(19). Anterolateral margins of pronotum and lateral margins at base of coria widely
bordered in yellow; often with yellow median stripe on pronotum and/or scutel-
lum, and often lateral margins and/or apex of scutellum yellow grave (Walker)
- Anterolateral margins of pronotum with incomplete and diffuse yellow border or
with none; scutellum and pronotal disk immaculate or marked with black 2 1
21(20). Cicatrices and large subquadrate patch at scutellar base on each side black
nigropictum (Breddin)
Pronotal disk and scutellum immaculate geniculatum (Dallas)
22(18). Dorsal punctation sparse, most punctures separated from nearest puncture by
distance equal to or greater than diameter of second antennal segment
viridans (St^l)
Dorsal punctation much denser 23
23(22). Connexiva immaculate or black mark at posterolateral angles of segments confined
to edge of sternite (Fig. 34) 24
Connexival spots at least on basal segments expanding onto laterotergite (Figs.
33, 40) 37
24(23). Rostrum terminating at mesocoxae; anterolateral margins of pronotum strongly
convex (Fig. 75) pennsylvanicum (P.B.)
Rostrum extending to or beyond metacoxae; anterolateral margins of pronotum
straight or weakly convex 25
25(24). Spiracles black 26
- Spiracles green, yellow, brown, or red 30
26(25). Most of dorsal punctation fuscous; callus around each spiracle weak, not differ-
entiated by color pecosum, new species
- Dorsal punctures concolorous with surrounding area; callus surrounding each
spiracle usually yellowish 27
27(26). Posterolateral angles of sternites with conspicuous black spot 29
- Not more than minute spine at posterolateral angles of sternites dark or black 28
28(27). Basal 3 segments of antennae v/ithout black markings; large tooth present at
anteromesial corner of dorsal rim on each side of posterior wall of pygophore (Fig.
84) aseadurn, new species
- All antennal segments partially black; denticle at anteromesial corner of dorsal
rim little larger than other denticles on rim ecuadorensis, new species
VOLUME 91, NUMBER 2
107
29(27).
30(25).
31(30).
32(31).
33(31).
34(33).
35(33).
36(35).
37(23).
38(37).
39(37).
40(39).
41(40).
42(40).
43(42).
44(39).
45(44).
46(45).
47(3).
Humeri forming obtuse angle (Fig. 87); anterolateral margin of pronotum straight
nigridorsatum (Breddin)
Humeri broadly rounded; anterolateral margin of pronotum slightly convex
ubicum, new species
Black dot present at mesial limit of each cicatrice, sometimes also at lateral limit
laetum (StM)
Cicatrices immaculate 3 1
Ostiolar ruga extending only halfway from mesial margin of ostiole to lateral
margin of thorax 32
Ostiolar ruga extending about % distance from mesial margin of ostiole to lateral
margin of thorax 33
Each spiracle on small ivory callus simplicis, new species
Spiracles not accompanied by callus euri, new species
Each spiracle on ivory callus 35
Spiracles not accompanied by differentially colored callus 34
Costal margin of coria basally and margin of scutellum at apex orange to yellow
macdonaldi, new species
Costal margin of coria and apex of scutellum green wygodzinskyi, new species
Abdominal spine projecting between metacoxae monticola, new species
Abdominal tubercle not or barely attaining metacoxae 36
Posterior margin of basal plates sloping toward meson; antennal segments 4 and
5 black or green distally ubicum, new species
Posterior margin of basal plates nearly transverse mesad of rounded lateral angle;
distal parts of antennal segments 4 and 5 castaneous occasi, new species
Tibiae crimson proximally 38
Proximal end of tibiae green or yellow 39
Costal angle of coria above last connexival segment collis, new species
Costal angle of coria above penultimate connexival segment ... montivagum (Distant)
Spiracles black 44
Spiracles pale 40
Each spiracle on or beside yellow callus 4 1
Spiracles unattended by callus but sometimes in yellow spot 42
Each spiracle in edge of callus 60
Each spiracle contiguous with but not surrounded by callus australe, new species
Costal angle of coria subacute, lying above sixth visible sternite (Fig. 153)
.fuscopunctatum (Breddin)
Costal angle of coria rounded, lying above fifth visible sternite 43
Females; basal plates with posterolateral projection at base of 9th paratergite (Fig.
157); males: posterior margin of pygophore with broad V-shaped emargination
from ventral view (Fig. 159) marginatum (P.B.)
Females: posterior margin of basal plates evenly convex (Fig. 1 64); males: py-
gophoral emargination shallow with small median notch from ventral view (Fig.
166) hilare {S^.y)
Black spot present in basal angles of scutellum brasicola, new species
Basal angles of scutellum immaculate 45
Abdominal tubercle nearly reaching anterior limit of metacoxae .. teretis, new species
Abdominal tubercle barely reaching posterior limit of metacoxae 46
Humeral margin obtusely angular; anterolateral margin of pronotum straight
nigridorsatum (Breddin)
Humeral margin rounded; anterolateral margin of pronotum weakly convex
ubicum, new species
Connexival segments bordered both anteriorly and posteriorly with black 48
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NEW YORK ENTOMOLOGICAL SOCIETY
48(47).
49(48).
50(49).
51(50).
52(47).
53(52).
54(52).
55(54).
56(55).
57(54).
58(57).
59(58).
60(41).
Connexival segments bordered posteriorly only with black 52
Some or all femora with preapical black band musivum (Berg)
Black femoral markings apical or absent 49
Anterolateral margins of pronotum broadly bordered with black near humeri and
yellow anteriorly rogenhoferi (St^l)
Border of anterolateral margins of pronotum entirely red or yellow 50
Sutures and pseudosutures on abdominal venter black erythrocnemis (Berg)
Sutures and pseudosutures of abdominal venter not contrasting strongly in color
with remainder of abdominal venter 5 1
Each spiracle in yellowish callus montivagum (Distant)
Spiracles unaccompanied by callus ninaspis (Dallas)
Base of tibiae with reddish band 53
Tibiae without basal band 54
Basal angles of scutellum with small black macule; spiracles and spiracular calli
black collis, new species
Basal angles of scutellum immaculate, spiracles reddish, spiracular calli yellow ..
montivagum (Distant)
Spiracles black 55
Spiracles pale, only peritreme sometimes thinly ringed in black 57
Abdominal tubercle pronounced; black spot usually present in basal angles of
scutellum and at mesial limit of cicatrices 56
Abdominal tubercle rudimentary; pronotum and scutellum without black mark-
ings plaumanni, new species
Anterior pronotal disk transversely depressed, forming shallow basin; anterolateral
margins of pronotum slightly convex dallasi (Distant)
Pronotal disk weakly convex anteriorly, any depressed area submarginal; antero-
lateral margins of pronotum nearly straight bipunctulum (St^l)
Each spiracle on distinct yellow callus; yellowish macules present along base of
scutellum laetum (Stal)
Callus around each spiracle weakly or not differentiated by color; yellow macules
rarely present at base of scutellum 58
Black spot present in basal angles of scutellum irnpicticorne (St^l)
Basal angles of scutellum without black spot 59
Dorsal borders of body not differentially colored or inconspicuous light yellow;
abdominal spine reaching anterior limit of hind trochanters herbidum (St^l)
Dorsal borders conspicuous, narrow, red; abdominal spine shorter, reaching pos-
terior limit of metacoxae pengue, new species
Black spot present at mesial limit and sometimes also at lateral limit of cicatrices
laetum (StSl)
Cicatrices immaculate apicicorne (Spinola)
Acrosternum (Chinavia) armigenim (StM, 1859)
(Figs. 7-11)
Rhaphigaster armiger Stk], 1859, pp. 229-230.
Nezara {Acrosternum) armigera: Stal, 1872, p. 43 (keyed).
VOLUME 91, NUMBER 2
109
Figs. 7-11. A. annigerum. 7. Humeral angle and head. 8. Pygophore, caudal view. 9. Genital
cup, dorsal view. 10. Paramere. 1 1. Spermathecal bulb and pump.
Nezara armigera: Berg, 1878, p. 28 (descriptive note; reprinted 1879, p. 58);
Berg, 1883, pp. 213-214 (description; reprinted 1884, pp. 29-30).
Acrosternum armigera: Piran, 1948, p. 9 (records); Grazia-Vieira and Casini,
1973, p. 57 (record).
Diagnosis. Humeral angles spinose. Abdominal spine terminating between
metacoxae. Small black spot present at posterolateral angles of connexiva
and sternites; laterotergites immaculate. Spiracles usually black, each in
posterolateral edge of weak small and usually yellow callus. Ostiolar ruga
on each side extending 0.6-0. 7 distance from mesial margin of ostiole to
lateral margin of thorax. Rostrum reaching metacoxae. Femora green. Head
moderately rounded apically (Fig. 7). Cicatrices immaculate. Basal angles
of scutellum lacking black spot. Punctation on dorsum dense, punctures
concolorous with surrounding area. Posterior wall of genital cup subvertical,
deeply emarginate mesially, expanded mesodorsad on each side; this ex-
pansion acute apically, bearing several small black denticles on dorsal rim
(Figs. 8, 9). Parameres and spermatheca as in Figures 10 and 11. Length
about 10-13 mm.
Distribution. Northern Argentina, southern Brazil, Paraguay, Uruguay.
NEW YORK ENTOMOLOGICAL SOCIETY
1 10
Types. Lectotype, here designated, 5, labeled (a) “Buenos Ayres” (b) “Type”
(c) “Typus” (d) “109/79” (e) “Riksmuseum Stockholm”. Paralectotype: 2
labeled (a) “Buenos Ayres” (b) “Westerman” (c) “Allotypus” (d) “1 10/79”
(e) “Riksmuseum Stockholm”.
Acrosternum (Chinavia) bellum, new species
(Figs. 12-16)
Description. Light dull green, becoming yellowish to yellowish green me-
sially below, with narrow yellow to red lateral borders on head, pronotum,
coria basally, connexiva and abdominal venter. A black dot present at pos-
terolateral angles of connexival segments (excluding laterotergites) and ster-
nites. Antennae green. Legs green with base of femora, trochanters and coxae
yellowish. Punctation on dorsum dense, fine; punctures concolorous with
surrounding area; thoracic venter more sparsely and irregularly punctate
than dorsum.
Head 2. 4-2. 6 mm wide across eyes, 1. 9-2.0 mm long, proportionately as
in armigerum but more narrowly rounded at apex (Fig. 12). Antennal seg-
ments 0.4-0. 5, 0. 9-1.0, 1.0-1. 2, 1.2-1. 4, 1.5 mm long. Rostral segments 2
through 4 about 1.2-1. 4, 0. 9-1.0, 0.7-0. 9 mm long; apex reaching meta-
coxae.
Pronotum 7. 0-8. 3 mm wide at humeri, 2. 4-2. 8 mm long at meson. An-
terolateral margins nearly straight. Humeral angles right angular to spinose.
Cicatrices immaculate.
Scutellum 4.0-4. 5 mm wide at base, 4. 3-4. 8 mm long. Four or 5 small
yellow spots spaced along scutellar base.
Abdominal spine reaching only to posterior limit of metacoxae, its anterior
margin subvertical. Each spiracle located in posterolateral edge of small ivory
callus; peritremes black. Ostiolar ruga on each side extending about 0.7
distance from mesial margin of ostiole to lateral margin of thorax.
Female unknown.
Posterior margin of pygophore from ventral view with broad shallow
sinuous concavity (Fig. 1 3). Posterior wall of genital cup diagonal at dorsal
margin on each side, there bearing a large denticle anteriorly (bent antero-
laterad at apex) and elsewhere weak denticles (Figs. 14, 15). Paramere as in
Figure 16.
Length about 1 1.0-1 1.5 mm.
Distribution. Northern Argentina, southern Brazil, Paraguay.
Holotype. S, labeled “Brazil, Embrapa, 28-IX-1976, Coll. Ev. Vogel”.
Deposited in U.S. National Museum, type no. 75560.
Paratypes. 6, labeled “Museum Leiden. Argentina. Pronuntiamente, Prov.
Entre Rios. I, 1965. F. H. Walz.” (RHN); c5, labeled “Museum Leiden. Dr.
VOLUME 91, NUMBER 2
111
Figs. 12-16. A. bellum. 12. Head. 13. Pygophore, ventral view. 14. Pygophore, caudal view.
15. Genital cup. 16. Paramere.
B. Podtiaguine. Assomption; Omgev. Paraguay. 22 Feb. -25 Apr. 1936”
(RNH).
Comments. Superficially resembling A. armigerum, the two species differ
especially in the shape of the head, abdominal tubercle and pygophore.
Acrosternum (Chinavia) insulani, new species
(Figs. 17-20)
Description. Light green above, paler green below, with lateral margins of
head, pronotum, coria basally, abdominal venter as well as apex of scutellum
and connexiva reddish-yellow. Transverse sutures of connexiva broadly bor-
dered on both sides with black, these bands continuing onto abdominal
venter and evanescing before reaching spiracular line. Spot on superior sur-
face of antennifers and antennae excepting base of first segment black. Legs
excepting somewhat darker tarsi concolorous with venter. Punctation on
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NEW YORK ENTOMOLOGICAL SOCIETY
dorsum moderately dense, fine; punetures concolorous with surrounding
area; abdominal venter sparsely punctate and aciculate.
Head 2.85 mm wide across eyes, 2.3 mm long. Antennal segments 0.5,
1.1, 1.5, 1.8, 1.9 mm long. Rostral segments 2 through 4 about 1.5, 1.4, 1.1
mm long; apex reaching metacoxae.
Pronotum 8.0 mm wide at humeri, 5.3 mm long at meson. Humeri pro-
duced laterad, forming right angle. Cicatrices immaculate.
Scutellum 4.6 mm wide at base, 5.3 mm long, without markings other
than pale apex.
Abdominal spine projecting slightly past anterior limit of mesocoxae.
Spiracles yellowish, unattended by callus. Ostiolar ruga on each side ex-
tending 0.7 distance from mesial margin of ostiole to lateral margin of thorax.
Female unknown.
Posterior pygophoral margin shallowly concave from ventral view with
slight protrusion on each side of mesial emargination (Fig. 17), this emar-
gination from caudal aspect deep, U-shaped (Fig. 18). Posterior wall of
genital cup diagonal dorsally on each side with large tubercle at anteromesial
corner of dorsal rim; this tubercle directed anterolaterad, acute apically (Fig.
19). Parameres elongate (Fig. 20).
Length about 14 mm.
Distribution. Hispaniola.
Holotype. 6, labeled “Constanza to Jarabacoa. Aug. ’38. Dom. Rep. 2-
4000 ft. Dari.” Deposited in the American Museum of Natural History.
No paratypes.
VOLUME 91, NUMBER 2
113
Acwsternum (Chinavia) abnonne (Berg, 1892)
(Figs. 21, 22)
Nezara abnormis Berg, 1892, pp. 7-8; Piran, 1966, p. 86, fig. 1 (record, 9
genitalia).
Acwsternum abnorme: Piran, 1968, p. 17 (record).
Diagnosis. Ruga from metathoracic scent gland about as long as ostiole.
Humeri not produced, rounded. Abdominal spine terminating between meta-
and mesocoxae. Each spiracle located in lateral part of large yellowish callus;
peritreme little darker than callus. Rostrum reaching metacoxae. Femora
without black markings. Cicatrices immaculate. Subcalloused orange-yellow
spot present in basal angles of scutellum. Dorsal punctation dense, fine,
concolorous with light green dorsum, in irregular transverse rows with pale
to orange-yellow interstices forming subtle mesial stripe on pronotum and
on scutellum except basally. Pygophore from caudal aspect deeply emargin-
ated mesially, appearing bilobed laterally (Fig. 2 1 ). Genital plates as in Figure
22. Length about 10.5 mm.
Distribution. Northern Argentina, southern Brazil, Uruguay.
Type. Described from a single male preserved in the Universidad Nacional
de La Plata.
Comment. The holotype was examined.
Acrosternum (Chinavia) callosum, new species
(Figs. 23-25)
Description. Light green above with lateral submargins of head and many
interstices between punctures yellow. Venter yellowish green laterally, blend-
ing to yellow mesially. Antennae green with last 2 segments brownish. Coxae
and trochanters yellow, remainder of legs green. Border along posterior
margin of connexival segments, posterolateral angles of sternites, elongated
macule on antennifers and edge of head before eyes, all black. Dorsal punc-
tation moderately dense, fine, in irregular transverse rows on most of prono-
tum, green; abdominal venter lightly and densely aciculate.
Head 2. 6-2. 8 mm wide across eyes, 2. 1-2.2 mm long. Antennal segments
0.4-0. 5, 1.0-1. 1, 1.2-1. 4, 1.6-1. 7, 1.5-1. 6 mm long. Last 3 rostral segments
about 1.3-1. 4, 1.0-1. 1, 0.8-0. 9 mm long; rostrum reaching metacoxae.
Pronotum 6.6-7. 1 mm wide at humeri, 2.4-2. 8 mm long at meson. Hu-
meri not produced laterad, rounded. Anterolateral margins straight. Cica-
trices immaculate.
Scutellum, 4. 2-4. 6 mm wide at base, 4. 5-4.9 mm long; basal angles im-
maculate.
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NEW YORK ENTOMOLOGICAL SOCIETY
Figs. 21, 22. A. abnorme. 21. Pygophore, caudal view. 22. Genital plates, caudoventral
view.
Abdominal spine compressed, projecting slightly beyond mesocoxae. Spi-
racles not on callus, concolorous with surrounding area of sternites. Sternites
slightly calloused on both sides of transverse sutures, latter green except
mesially and at lateral margins. Pseudosuture between spiracle and tricho-
bothria similarly calloused and colored. Ostiolar ruga on each side extending
about 0.7 distance from mesial margin of ostiole to lateral margin of thorax.
Genital plates similar to those of following species (Fig. 27).
Posterior margin of pygophore from ventral view sinuously emarginate,
notched mesially and protruding slightly on each side of notch (Fig. 23);
from caudal view posterior margin sinuously emarginate with denticles vis-
ible on dorsal rim of posterior wall of genital cup (Fig. 24). Posterior wall
of genital cup diagonal dorsally on each side, bearing small black denticles
posteriorly on dorsal rim and large anterolaterally directed curved denticle
anteriorly (Fig. 25).
Length about 1 1.5 mm.
Distribution. Paraguay, Bolivia.
Holotype. 6, labeled (a) “Village #17. Fern. Col. Chaco. Paraguay. III-l-
56.” (b) “cotton” (c) “J. L. Nichel, Collector.” (d) “He-21.” Deposited in
U.S. National Museum, type no. 75561.
Paratypes. 6, 9, labeled “Bolivia, S. C., 1 mi. W. Pto. Pailas. Apr. 21,
1978. C. W. O’Brien & Marshall” (6 HDE; 9 LHR).
Comment. This species is especially distinguished from the following species
by the longer abdominal spine, black margins of the head, and calli along
the transverse sutures and pseudosutures of the abdominal venter. These
two species, among the Acrosternum of the Western Hemisphere whose
abdominal spine projects past the metacoxae, are the only ones having the
posterior margin of the connexival segments completely bordered in black.
VOLUME 91, NUMBER 2
115
view. 25. Genital cup. Figs. 26, 27. A. occultum. 26. Connexival segment. 27. Genital plates,
caudoventral view.
Acrosternum (Chinavia) occultum, new species
(Figs. 26, 27)
Description. Light green above and below including appendages. Venter
blending to yellowish brown mesially; abdominal margins yellow. Posterior
border of connexival segments, posterolateral angles of abdominal sternites
and elongated mark on antennifers black (Fig. 26). Dorsal punctation dense,
fine; punctures concolorous with surrounding area; transverse interstices on
pronotum somewhat rugose.
Head 2.8 mm wide across eyes, 2.2 mm long. Antennal segments 0.5, 1.0,
1.2, 1.5-1. 7, 1.7 mm in length. Last 3 rostral segments about 1.5, 1.1, 0.9-
1.0 mm long; rostrum reaching metacoxae.
Pronotum 7.0 mm wide at humeri, 2.7 mm long. Humeri not produced
laterad, rounded; anterolateral margins slightly convex. Cicatrices immac-
ulate.
Scutellum 4. 6-4. 7 mm wide at base, 4.9 mm long. Obscure dark dot
sometimes present in basal angles.
Abdominal spine compressed, reaching to or between mesocoxae. Spi-
racles not on callus, more or less concolorous with surrounding area of
sternites. Ostiolar ruga on each side extending 0.7 distance from mesial
margin of osiole to lateral margin of thorax.
Genital plates as in Figure 27.
Male unknown.
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Length about 12 mm.
Distribution. Bolivia (Beni).
Holotype. 2, labeled (a) “Bolivia: Dept. Beni, Rio Itenez at mouth of Rio
Baures. X-10-1964”. (b) “J. K. Bouseman, Collector”. Deposited in the
American Museum of Natural History.
Paratype. 2, with same data as holotype except date “IX-X-1964” and
(b) “J. K. Bouseman, J. Lugenhop, Collectors” (LHR).
Comment. Excepting the longer abdominal spine, this species appears
indistinguishable from A. herbidiim. While some intraspecific variation oc-
curs in the length of the abdominal spine, it seems improbable that so much
variability is contained in one species.
Acrosternum (Chinavia) obstinatiim (Stal, 1860)
(Figs. 28-33)
Rhaphigaster obstinatus Stal, 1860, p. 23.
Nezara {Acrosternum) obstinatus: Stal, 1872, p. 42 (keyed, A. difficile com-
pared).
Diagnosis. Humeri obtusely rounded, not produced. Abdominal spine
compressed, reaching anterior limit of mesocoxae. Posterior angles of con-
nexival segments (sometimes including posterolateral angles of laterotergite)
and of sternites with small black spot (Fig. 33). Lateral jugal margins thinly
or not edged in black; distinct submarginal band absent. Each spiracle at
posterolateral edge of weak callus; peritremes brown to narrowly black.
Ostiolar ruga on each side extending 0.6-0. 8 distance from mesial margin
of ostiole to lateral margin of thorax. Femora green. Cicatrices immaculate.
Diffuse yellow spot often present at basal angles of scutellum. Dorsal punc-
tation dense, fine. Pygophore from ventral view with mesial notch in broad
shallow emargination (Fig. 28). Posterior wall of genital cup dorsally diagonal
on each side; dorsal rim with small denticles posteriorly and large denticle
anteriorly, latter bending first anterolaterad then cephalad at acute apex (Fig.
30). Parameres and spermatheca as in Figures 31 and 32.
Length about 11-15 mm.
Distribution. Southern Brazil.
Types. Lectotype, here designated, 3, labeled (a) “Brasil” (b) “F. Sahib”
(c) “Type” (d) “Typus” (e) “122/79” (f) “Riksmuseum Stockholm”; para-
lectotype, 5, labeled (a) “Rio Jan” (b) “Stal” (c) “Type” (d) “Paratypus” (e)
“123/79” (f) “Riksmuseum Stockholm”.
Acrosternum {Chinavia) sparniiim (Dallas, 1851)
(Fig. 34)
Rhaphigaster sparnius Dallas, 1851, p. 280.
Nezara sparnius: Stal, 1872, p. 43 (listed).
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117
Acrosternum sparnium; Rolston, 1976, p. 4 (generic placement).
Diagnosis. Lateral margins of head black with well-defined yellow band
submarginally. Humeral angles rounded, not produced laterad. Abdominal
spine compressed, projecting beyond mesocoxae, ending about midway be-
tween meso- and procoxae. Posterolateral angles of sternites and of con-
nexiva where protruding, black; this mark on connexiva often elongated
anteriorly (Fig. 34). Spiracles unattended by callus; peritremes brown to
rufous. Ostiolar ruga on each side extending 0. 7-0.8 distance from mesial
margin of ostiole to lateral margin of thorax. Rostrum reaching metacoxae.
Femora green to yellowish green. Cicatrices immaculate. A yellowish spot
present at basal angles of scutellum. Dorsal punctation dense, shallow.
Length about 10.5 mm.
Distribution. Jamaica, Cayman Is.
Comment. The female holotype was examined. No male was seen.
Acrosternum (Chinavia) scute/latum (Distant, 1890)
(Figs. 35-40)
Nezara scutellata Distant, 1890, pp. 339-440, pi. 31, fig. 21.
Acrosternum scutellatum: Rolston, 1976, p. 4 (generic placement).
Diagnosis. Connexiva with large black macules, each bisected by trans-
verse connexival suture, continuing onto margin of venter (Fig. 40). Distal
end of femora and proximal end of tibiae usually reddish. Humeri rounded,
not produced laterally. Abdominal spine compressed, ending beyond meta-
coxae, sometimes reaching mesocoxae. Spiracles yellowish, unattended by
callus. Ostiolar ruga on each side extending 0.6-0. 8 distance from mesial
margin of ostiole to lateral margin of thorax. Rostrum extending to meta-
coxae. Cicatrices and basal angles of scutellum immaculate. Dorsal punc-
tation dense, fine. Pygophore broadly emarginate from ventral view; emar-
gination shallow, somewhat sinuous (Fig. 35). Posterior wall of genital cup
represented on each side by large tubercle projecting anterodorsad and ap-
pearing capitate from caudal view (Figs. 36, 37). Paramere and spermatheca
as in Figures 38 and 39.
Length about 14-18 mm.
Distribution. From Mexico (Veracruz) into Panama.
Comment. The female holotype was examined.
Acrosternum (Chinavia) froeschneri, new species
(Figs. 41, 42)
Description. Yellowish green above and below including appendages, tint-
ed reddish orange along lateral margins of head, anterolateral angles of
pronotum and labial groove. Punctation mostly of moderate strength and
density, somewhat rugose on pronotal disk, less dense on scutellum, fine
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Figs. 28-34. Figs. 28-33. A. obstinatum. 28. Pygophore, ventral view. 29. Pygophore, caudal
view. 30. Genital cup. 3 1 . Paramere. 32. Spermathecal bulb and pump. 33. Connexival marking;
laterotergite (L). Fig. 34. A. sparniurn. Connexival marking; edge of sternite (S).
and sparse on abdominal venter; punctures concolorous with surrounding
area.
Head 3.0-3. 1 mm wide across eyes, 2. 5-2.6 mm long. Lateral margin of
juga scarcely concave before eyes, tapering to rather broadly rounded apex
of head (Fig. 42). Length of antennal segments 0. 5-0.6, 1.0-1. 1, 1.3-1. 4,
1.4, 1.4 mm; last 3 segments subequal in length. Rostral segments 2 through
4 about 1.5, 1. 1-1.2, 0. 9-1.0 mm long; rostrum reaching anterior limit of
metacoxae.
Pronotum 8. 2-8. 8 mm wide at humeri, 3. 1-3.5 mm long at meson. Hu-
meri broadly rounded, slightly produced laterad. Cicatrices immaculate.
Scutellum 5. 1-5.5 mm wide at base, 5. 9-6. 5 mm long, without markings.
Posterolateral angles of connexival segments bearing small black mark.
Abdominal spine conical, projecting beneath mesocoxae and there ter-
minating. Spiracles not accompanied by callus; peritremes nearly concol-
orous with surrounding areas of venter. Posterolateral angles of sternites
minutely marked with black. Ostiolar ruga on each side extending 0.7-0. 8
distance from inner margin of ostiole to lateral margin of thorax.
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119
Figs. 35-40. A. scutellatum. 35. Pygophore, ventral view. 36. Pygophore, caudal view. 37.
Genital cup. 38. Paramere. 39. Spermathecal bulb and pump. 40. Connexival markings.
Posterior edge of basal plates produced as small ridge at base of 9th
paratergites (Fig. 41).
Male unknown.
Length about 14.4-15.4 mm.
Figs. 41, 42. A. fweschneri. 41. Genital plates, caudoventral view. 42. Head.
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Distribution. Mexico (Sinaloa).
Holotype. $, labeled (a) ‘‘4 mi. N. San Bias, Sinaloa, Mexico. VIII- 17-65.
H. R. Burke & J. Meyer” (b) ‘‘taken at light”. Deposited in U.S. National
Museum, type no. 75559.
Paratype. 9, labeled as holotype (TAMU).
Comment. This species is dedicated in gratitude to Dr. Richard C. Froesch-
ner of the U.S. National Museum.
Among the species of Acrosternum in the Western Hemisphere whose
abdominal spine projects beyond the metacoxae, only A. scuteUatum, A.
isturn and this species are found in Middle America.
Acrosternum (Chinavia) esmeraldum, new species
(Figs. 43-48)
Description. Dark green dorsally, lighter below and blending irregularly
to yellow mesially; lateral margins of head, pronotum, coria basally and
abdomen very narrowly yellow; antennae and legs dark green, latter often
discolored brown on inferior surface. Punctation on dorsum dense, hne, with
many punctures on pronotum and scutellum forming irregular transverse
lines; punctures dark green.
Head width across eyes 3. 2-3. 3 mm 99, 2.9 mm length 2.4-2. 5 mm
99, 2. 0-2. 2 mm 63. Antennal segments 0.6-0. 7, 1.3, 1.5, 2.0, 2.1 mm long
99, 0.6, 0. 9-1.1, 1.3-1. 6, 1.8, 2.0 mm long 66. Length of rostral segments 2
through 4 about 1. 9-2.0, 1.5, 1. 1-1.2 99, 1.4-1. 5, 1.2-1. 3, 1.0 66; rostrum
reaching metacoxae. Lateral margins of head rather strongly tapered toward
apex (Fig. 43).
Pronotal width at humeri 7. 7-7. 9 mm 99, 6. 6-6. 9 mm 66; mesial length
3. 2-3. 3 mm 99, 2. 7-2. 9 66. Humeri rounded, not produced laterad. Disk
evenly convex transversely between anterolateral margins. Cicatrices with-
out black markings.
Scutellar width at base 5. 1 mm 99, 4. 3-4. 6 mm 66; length 5.9 mm 99, 4.6-
5.2 mm 66. A small ivory callus present at each basal angle.
Abdominal spine compressed, reaching mesocoxae. Each spiracle in pos-
terolateral margin of small ivory callus; peritremes black. Minute spine at
posterolateral angles of connexival segments black, connexiva otherwise
immaculate. Ostiolar ruga on each side reaching 0.6-0. 8 distance from mesial
margin of ostiole to lateral margin of thorax.
Pygophore deeply emarginate from caudal view (Fig. 44). On each side
posterior wall of genital cup expanded mesodorsad; dorsal rim concave,
denticulate, curved from dorsal view; dorsally serrated ridge projecting pos-
teriorly from lateral limit of dorsal rim. Dorsal margin of genital cup bearing
black denticle at angle formed by lateral and anterior concavities, this angle
a little laterad of and cephalad from mesial limit of dorsal rim of posterior
wall (Fig. 45). Ventral surface of pygophore narrowly sulcate submarginally.
VOLUME 91, NUMBER 2
121
Figs. 43-48. A. esmeraldum. 43. Head. 44. Pygophore, caudal view. 45. Genital cup. 46.
Paramere. 47. Right basal plate. 48. Spermathecal bulb and pump.
Posterior surfaces densely setose. Parameres as in Figure 46. Basal plates
and spermatheca as in Figures 47 and 48.
Distribution. Brazil (Minas Gerais).
Holotype. 6, labeled (a) “Brazil. Minas Gerais: Santa Barbara, Sierra do
Caraca” (b) “1450 m. Jan. 1970. F. M. Oliveira.” Deposited in American
Museum of Natural History.
Paratypes. 2SS, 2$9, labeled as holotype (5 LHR; 6, 2$$ AMNH).
Acrosternum (Chinavia) istum, new species
(Fig. 49)
Description. Light green above and below with yellowish orange border
along outer margins. Black markings confined to posterolateral angles of
sternites, connexival spots that include posterolateral angles of laterotergites,
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Figs. 49-51. Fig. 49. A. istum. Basal plates, caudal view (BP). Figs. 50, 51. A. napaeum.
50. Head. 51. Right basal plate.
apex of rostrum, distal one-third of antennal segment 3 and distal one-fourth
of segment 4. Dorsal punctation moderately dense, dark green.
Head 3.0 mm wide across eyes, 2.7 mm long; lateral jugal margins slightly
concave. Antennal segments 0.5, 1 .4, 1.5, 2.0, 2.0 mm long. Rostral segments
2-4 about 1.8, 1.4, 1.1 mm long; rostrum reaching posterior limit of meta-
coxae.
Pronotum 7.8 mm wide at humeri, 3.2 mm long at meson. Humeri nar-
rowly rounded, little produced laterad; anterior pronotal margin straight.
Scutellum 5.0 mm wide at base, 5.9 mm long. Costal angle of coria round-
ed, reaching posterior half of penultimate connexival segment.
Abdominal tubercle compressed, attaining posterior limit of mesocoxae.
Spiracles brown, each in posterolateral margin of bright green subcalloused
spot. Ostiolar ruga on each side extending about three-fourths distance of
mesial margin of ostiole to lateral thoracic margin.
Posterior margin of basal plates protruding beneath 9th paratergite; dorsal
edge of combined plates mesially emarginate from caudal view (Fig. 49).
Male unknown.
Length about 13.7 mm.
Distribution. Panama.
Holotype. 9, labeled “Panama. Pan. Prov. Altos de Maje, Chepo. 14-15
VOLUME 91, NUMBER 2
123
May 1976, at lights. Col: D. Engleman.” Deposited in U.S. National Mu-
seum, type no. 75565. No paratypes.
Comments. This species is similar in appearance to A. marginatum, which
also occurs in Panama, but differs especially in the longer abdominal spine
and less dense dorsal punctation.
Acrosternum (Chinavia) napaeum (Stal, 1872)
(Figs. 50, 51)
Nezara {Acrosternum) napaea Stal, 1872, p. 42.
Diagnosis. Pronotum broadly depressed submarginally along cephalic half
of anterolateral margins. Abdominal spine compressed, terminating between
mesocoxae. Humeral angles rounded, not produced laterad. Connexiva im-
maculate excepting black spine at posterolateral angles of each segment.
Spiracles unattended by callus; peritremes brown. Ostiolar ruga on each side
extending about 0.8 distance from inner margin of ostiole to lateral margin
of thorax. Rostrum reaching metacoxae. Dorsum light green, femora con-
colorous, cicatrices and scutellum immaculate; antennae black excepting
green first segment. Lateral margins of head sigmoid (Fig. 50). Punctation
on dorsum moderate in density and strength, less dense and weaker on
scutellum. Basal plates longitudinally sulcate below 9th paratergite (Fig. 51).
Length about 13.5 mm.
Distribution. Brazil.
Comment. Of this species I have seen only the holotype.
Acrosternum {Chinavia) difficile {Sxk\, 1860)
(Figs. 52-57)
Rhaphigaster difficilis Stal, 1860, p. 23.
Nezara {Acrosternum) difficilis: Stal, 1872, p. 42 (keyed, descriptive note).
Diagnosis. Humeri broadly rounded, not produced laterad. Abdominal
spine compressed, terminating between mesocoxae. Posterolateral connex-
ival angles minutely marked with black. Lateral jugal margins sigmoid (Fig.
52). Weak callus bearing spiracles not strongly differentiated by color; peri-
tremes thinly black. Ostiolar ruga on each side extending about 0.7 distance
from mesial margin of ostiole to lateral margin of thorax. Anterior disk of
pronotum weakly convex without submarginal depression along anterolat-
eral margins. Dorsum dark green; neither cicatrices nor basal angles of scu-
tellum with black spots; femora green. Punctation on dorsum moderately
strong, dense; less dense and weaker on scutellum. Junction between corium
and membrane rounded (Fig. 57). Rostrum reaching metacoxae. Pygophore
from ventral view notched at bottom of broad shallow emargination of
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NEW YORK ENTOMOLOGICAL SOCIETY
Figs. 52-57. A. dijficile. 52. Head. 53. Pygophore, ventral view. 54. Pygophore, caudal view.
55. Genital cup. 56. Paramere. 57. Hemelytron.
posterior margin (Fig. 53); deeply emarginate from caudal view (Fig. 54).
Posterior walls on each side of genital cup curving anteromesad, with large
anterior denticle curving anterolaterad and several small black denticles
posteriorly on dorsal rim (Fig. 55). Paramere as in Figure 56.
Length about 11-15 mm.
Distribution. Southern Brazil.
Comment. The female type was examined.
Acrosternum (Chinavia) longicorialis (Breddin, 1901), New Combination
(Figs. 58-60)
Nezara longicorialis Breddin, 1901, p. 123; Gaedike, 1971, p. 91 (lectotype
designated).
Diagnosis. Posterior margin of corium parabolic, intruding into and re-
ducing membrane of hemelytra (Fig. 58). Humeral angles rounded; antero-
lateral margin of pronotum slightly convex. Abdominal spine compressed.
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125
Figs. 58-65. Figs. 58-60. A. longicorialis. 58. Hemelytron. 59. Genital plates, caudoventral
view. 60. Genital cup. Figs. 61-65. A. grave. 6 1 . Pygophore, ventral view. 62. Pygophore, caudal
view. 63. Genital cup. 64. Paramere. 65. Spermathecal bulb and pump.
reaching anterior limit of mesocoxae. Connexiva immaculate. Spiracles red-
dish, each located laterally in edge of yellow oval spot. Ostiolar ruga on each
side reaching about halfway from inner margin of ostiole to lateral margin
of thorax. Rostrum reddish, terminating between metacoxae. Femora uni-
formly green. Cicatrices and basal angles of scutellum immaculate. Punc-
tation of dorsum fine, dense. Narrow yellowish mesial stripe running length
of pronotum and scutellum. Apex of scutellum subacute. Posterior walls of
genital cup strongly reflexed with low elevation along posterior half of mesial
margin, terminating in projection directed anteromesad which reaches su-
perior ridge of genital cup; only proctiger visible in genital cup, remainder
covered by posterior walls (Fig. 60). Basal plates subtriangular (Fig. 59).
Length about 12 mm.
Distribution. Uruguay (Montevideo) and Argentina (Buenos Aires).
Comment. The lectotype, designated by Gaedike (1971), was examined.
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The male genitalia of this species are distinctive in that the entire genital
cup excepting the proctiger is entirely closed. The shape of the corium,
intruding far into and reducing the membraneous area of the hemelytra, is
also diagnostic.
Acrosternum {Chinavia) panamensis (Distant), New Combination
Nezara panamensis Distant, 1890, p. 339, pi. 32 fig. 1.
Nezara (Pellaea) panamensis: Kirkaldy, 1909, p. 121 (listed).
Pellaea panamensis: Rolston, 1976, p. 7 (systematic position queried).
Description. Large black macule on dorsum comprised of distal part of
each corium and tergites beneath hemelytral membranes; also black: anten-
nae, rostrum excepting parts of first and base of second segments, tarsi,
tibiae, femora distally and on front legs femora proximally and trochanters.
Remainder of insect excepting eyes and ocelli brownish yellow, perhaps green
in life. Dorsal punctation fine, rather dense.
Head 2.9 mm across eyes, 2.4 mm long. Antennae 0.5, 1.0, 1.8, — , — ,
mm long. Rostral segments 2-4 about 1.8, 1.5, 1.2 mm long; apex of rostrum
lying between metacoxae. Juga not convergent apically.
Pronotum 7.8 mm wide at humeri, 3. 1 mm long mesially. Humeri broadly
rounded, not produced laterad. Anterolateral margins slightly convex, with-
out submarginal depression.
Scutellum 5.0 mm wide at base, 5.4 mm long. Coria ending distally near
middle of penultimate connexival segment; boundary between each corium
and membrane slightly convex; membranes heavily fumose excepting nar-
row hyaline border.
Abdominal tubercle reaching a little beyond posterior limits of metacoxae,
compressed apically. Ostiolar ruga on each side reaching about 0.75 distance
from inner margin of ostiole to lateral margin of thorax.
Posterior margins of basal plates forming nearly transverse line between
lateral angles.
Length about 13 mm.
Distribution. Panama. Known only from the female holotype collected at
Bugaba.
Comment. This species meets all of the morphological criteria for the
genus Acrosternum subgenus Chinavia insofar as can be determined from
the single known specimen. Only its coloration seems especially remarkable.
Acrosternum {Chinavia) grave (Walker, 1867)
(Figs. 61-65)
Strachia gravis Walker, 1867, p. 322.
Nezara nigritarsis Stal, 1872, p. 40. New Synonymy.
VOLUME 91, NUMBER 2
127
Nezara gravis: Distant, 1900, p. 393.
Nezara gentilis Breddin, 1903, p. 368; Gaedike, 1971, p. 86 (lectotype des-
ignated). New Synonymy.
Acrosternum grave: Rolston, 1976, p. 4 (generic placement).
Diagnosis. Anterolateral margins of pronotum and coria basally bordered
broadly in yellow; often all or margins of head, median stripe on pronotum
and/or scutellum also yellow. Antennae, most of rostrum and at least apex
of femora black. Humeri rounded, not produced. Abdominal spine just
reaching between metacoxae, sometimes reduced to obtuse tubercle. Con-
nexiva immaculate excepting tiny spine at posterolateral angles of segments
darkened. Membrane of hemelytra dark. Spiracles not accompanied by cal-
lus; peritremes pale. Ostiolar ruga on each side extending about 0.7 distance
from mesial margin of ostiole to lateral margin of thorax. Rostrum termi-
nating between metacoxae. Cicatrices immaculate. Basal angles of scutellum
usually immaculate, rarely with small dark dot. Dorsal punctation moder-
ately dense, dark. Posterior wall on each side of genital cup dorsally diagonal,
with short obtuse projection anteriorly (Figs. 62, 63). Parameres as in Figure
64. Spermatheca as in Figure 65.
Length about 12.5-13.5 mm.
Distribution. Amazon region.
Types. Lectotype of Nezara nigritarsis Stal, here designated, $ with wings
spread, labeled (a) “Amazon” (b) “Stevens” (c) “Type” (d) “Paratype” (e)
“120/79” (f) “Riksmuseum Stockholm”. Paralectotypes, 5, labeled as lec-
totype except (d) “Type” (e) “119/79”; 5, labeled as lectotype except (e)
“121/79”.
Comment. The holotype of Strachia gravis and of Nezara gentilis, both
females, as well as the syntypes of Nezara nigritarsis, were examined,
Acrosternum {Chinavia) geniculatum (Dallas, 1851)
(Figs. 66-68)
Rhaphigaster geniculatus Dallas, 1851, p. 279.
Rhaphigaster obscuricornis Stal, 1860, p. 22 (synonymized by StM, 1872).
Nezara geniculata: StM, 1872, p. 40 (keyed, synonymy).
Acrosternum geniculatum: Rolston, 1976, p. 3 (generic placement).
Diagnosis. Distal end of femora narrowly banded in black excepting in-
ferior surface. Cicatrices and scutellum unicolorous, green. Humeri rounded,
not produced. Abdominal tubercle compressed, terminating between meta-
coxae. Connexiva immaculate excepting spine on posterolateral angles of
segments dark. Spiracles unaccompanied by conspicuous yellow callus; peri-
tremes brown to green. Ostiolar ruga on each side extending about 0.8
distance from mesial margin of ostiole to lateral margin of thorax. Rostrum
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Figs. 66-68. A. geniculatum. 66. Genital plates, caudoventral view. 67. Pygophore, ventral
view. 68. Pygophore, caudal view.
terminating between or slightly caudad of metacoxae. Basal angles of scu-
tellum immaculate. Posterior margin of basal plates nearly transverse except
at convex angles (Fig. 66). Pygophore as in Figures 67 and 68. Length about
14.5-17 mm.
Distribution. Southern Brazil, Bolivia.
Type. Described from one 6 and one 9, the latter mislabeled with France
as locality. This specimen was not located. Lectotype, here designated, 6
bearing labels (a) “Type” (b) “720a” (c) “a” (d) ""Rhaphigaster geniculatus"'.
Acrosternum (Chinavia) nigropictum (Breddin, 1906), New Combination
Nezara nigropicta Breddin, 1906, pp. 193-194; Gaedike, 1971, p. 93 (lec-
totype designated).
Diagnosis. Apparently differing from A. geniculatum only in color. Tylus
excepting narrow mesial wedge at base, cicatrices, large subquadrate patch
on each side of scutellum at base (confluent basally), sutures of abdominal
venter except mesially and at lateral ends, all black. Length about 19 mm.
Distribution. Brazil (Manaus). Known only from lectotype.
Comment. This form may be a color variation of A. geniculatum Dallas.
The lectotype, a 9 designated by Gaedike (1971), was examined.
Acrosternum {Chinavia) viridans {Sxh\, 1859)
(Figs. 69-74)
Rhaphigaster viridans Stal, 1859, p. 228.
Nezara {Acrosternum) viridans: Stal, 1872, p. 41 (keyed).
Nezara viridans: Distant, 1880, p. 79 (synonymy).
Acrosternum viridans: Froeschner, 1981, p. 68 (catalog).
VOLUME 91, NUMBER 2
129
Figs. 69-79. Figs. 69-74. A. viridans. 69. Genital plates, caudoventral view. 70. Pygophore,
ventral view. 71. Pygophore, caudal view. 72. Genital cup. 73. Paramere. 74. Spermathecal
bulb and pump. Figs. 75-79. A pennsylvanicum. 75. Pronotum. 76. Pygophore, ventral view.
77. Pygophore, caudal view. 78. Genital cup. 79. Paramere.
Diagnosis. Dorsum sparsely punctate, most punctures separated from
nearest puncture by distance equal to or greater than diameter of second
antennal segment. Humeri rounded, slightly produced laterad. Abdominal
tubercle reaching posterior limit of metacoxae. Connexiva and sternites with
small black spot at posterolateral angles of each segment, connexival spot
extending onto laterotergite. Spiracles pale, unaccompanied by callus. Os-
tiolar ruga on each side reaching about halfway from mesial margin of ostiole
to lateral margin of thorax. Rostrum extending past metacoxae onto sternite
bearing tubercle. Legs green. Pronotum and scutellum without black mark-
ings. Posterior margin of basal plates at lateral angles bent up against 9th
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paratergite (Fig. 69). Posterior wall on each side of genital cup dorsally
diagonal; dorsal rim straight, denticulate, with large denticle anteriorly (Figs.
71, 72). Spermatheca as in Figure 74. Parameres as in Figure 73. Length
about 10-12.5 mm.
Distribution. Panama, Peru, Galapagos Islands.
Types. Lectotype, here designated, labeled (a) “Callao” (b) “Kinb” (c)
“Typus” (d) “124/79” (e) “Riksmuseum Stockholm”. Paralectotypes: 3,
labeled (a) “Panama” (b) “Kinb” (c) “Paratypus” (d) “125/79” (e) “Riks-
museum Stockholm”; 9, labeled (a) “Panama” (b) “Paratypus” (c) “127/79”
(d) “Riksmuseum Stockholm”; 9, labeled (a) “Ins. Gallop” (b) “Allotypus”
(c) “126/79” (d) “Riksmuseum Stockholm”.
Comments. No other species of the genus is so sparsely punctate dorsally.
Acrostenmm {Chinavia) pennsyh'anicum (Palisot de Beauvois, 1805)
(Figs. 75-79)
Cimex viridis pennsy/vanica DeGeer, 1773, p. 330, pi. 34, fig. 5; Gmelin in
Linnaeus, 1788, p. 2148 (unavailable trinomen).
Pentatorna pennsylvanica Palisot de Beauvois, 1805, p. 186, Hem. pi. 11,
fig. 5.
Pentatorna abrupta Say, 1831, p. 6; Say, 1859, p. 317 (synonymized by
Uhler, 1871).
Rhaphigaster parnisus Dallas, 1851, p. 279 (synonymized by Uhler, 1886).
Rhaphigaster pennsylvanicus: Uhler, 1871, p. 98 (synonymy, records ex-
cluding Panama); Uhler, 1878, p. 380 (identity verified).
Nezara (Acrosternum) pennsylvanica: Stal, 1872, p. 42 (keyed).
Nezara pennsylvanica: Uhler, 1886, p. 8 (synonymy); Osborn, 1892, p. 122
(records); Van Duzee, 1904, pp. 57, 58 (keyed, records); Van Duzee, 1909,
p. 157 (records); Olsen, 1912, p. 55 (records, host); Barber, 1914, p. 523
(records).
Nezara parnisus: Distant, 1900, p. 392 (listed).
Acrosternum pennsylvanicum: Parshley, 1915, p. 175 (keyed); Van Duzee,
1916, p. 7 (listed); Van Duzee, 1917, pp. 59-60 (cataloged); Parshley,
1917, p. 24 (records); Hart, 1919, p. 181 (record); Stoner, 1920, pp. 107-
1 08, pi. 7, fig. 1 (keyed, description); Hussey, 1 922, p. 1 5 (record); Parshley,
1923, p. 767 (record); Blatchley, 1926, pp. 160-161 (keyed, description);
Brimley, 1938, p. 63 (record); Torre Bueno, 1939, p. 236 (keyed); Froesch-
ner, 1941, p. 130 (keyed); McPherson, 1970, pp. 54-55, fig. 52 (records);
Hoffman, 1971, p. 48 (records); Furth, 1974, p. 40, pi. 2, fig. 22, pi. 3, fig.
37 (records).
Diagnosis. Rostrum reaching only to mesocoxae. Anterolateral margins
of pronotum usually strongly convex. Humeri rounded, not produced. Ab-
dominal tubercle compressed, reaching posterior limits of metacoxae. Con-
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131
Figs. 80-86. Figs. 80, 81. A. pecosum. 80. Head. 81. Genital plates, caudoventral view.
Figs. 82-86. A. aseadum. 82. Genital plates, caudoventral view. 83. Pygophore, ventral view.
84. Pygophore, caudal view. 85. Genital cup. 86. Paramere.
nexiva and sternites with small black spot at posterolateral angles, spot not
extending onto laterotergite. Spiracles usually black, each on yellow callus.
Ostiolar ruga on each side extending about 0.6 distance from mesial margin
of ostiole to lateral margin of thorax. Femora green. Pronotum and scutellum
without black markings. Genitalia as in Figures 76 and 77. Length about
10.5-14.5 mm.
Distribution. Eastern United States and southeastern Canada.
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NEW YORK ENTOMOLOGICAL SOCIETY
Acrosternum (Chinavia) pecosum, new species
(Figs. 80, 81)
Description. Emerald green above, paler green below blended with yellow
mesially; lateral margins of head, pronotum and basal part of coria narrowly
bordered in yellow. Connexiva and lateral margins of abdominal venter
yellow excepting spine at posterolateral angles of segments dark. Spiracles
black, surrounding area weakly calloused and not differentially colored. An-
tennae green with apex of segment 2, distal half of segments 3 and 5, distal
three-fourths of segment 4 black; spot on superior surface of antennifers
black. Dorsal punctation on head green, elsewhere distinctly black, mod-
erately dense.
Head 2.7 mm wide across eyes, 2.2 mm long, narrowly parabolic apically
(Fig. 80). Antennal segments 0.5, 1.0, 1 .4, 1 .8, 1 .9 mm long. Rostral segments
2-4 about 1.6, 1.0, 1.0 mm long; rostrum terminating between metacoxae.
Pronotum 6.4 mm wide at humeri, 2.4 mm long at meson. Humeri round-
ed, not produced. Anterolateral margins slightly convex. Cicatrices without
black marking.
Scutellum 4.1 mm wide at base, 4.2 mm long; basal angles immaculate;
very apex yellowish.
Abdominal tubercle compressed, barely attaining metacoxae. Ostiolar ruga
on each side extending 0.6-0. 7 distance from mesial margin of ostiole to
lateral margin of thorax.
Posterior margin of basal plates nearly transverse, thin and hyaline around
mesial angle (Fig. 81).
Male unknown.
Length about 10.9 mm.
Distribution. Peru.
Holotype. 9, labeled “Manu, Peru. X-21-63. Pena, Coll.” Deposited in the
American Museum of Natural History. No paratypes.
Comment. The black dorsal punctation together with the immaculate
connexiva distinguish this species.
Acrosternum (Chinavia) aseadum, new species
(Figs. 82-86)
Description. Light green above and below, becoming yellow mesially on
venter; margins of head, pronotum, coria basally and connexiva usually
bordered thinly in yellow or orange; lateral margins of venter with somewhat
wider border, without black markings on lateral margins of abdomen. Spi-
racles black, each located on ivory callus. Antennae green with last 2 segments
sometimes mostly ferrugineous to fuscous; spot on superior surface of an-
tennifers black. Dorsal punctation dense; punctures concolorous with sur-
rounding surface.
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133
Head 2. 5-2.9 mm wide across eyes, 2. 1-2.3 mm long; jugal margins sin-
uous. Antennal segments 0. 5-0.6, 1. 1-1.4, 1.2-1. 5, 1.6-1. 8, 1.6-1. 7 mm
long. Rostral segments 2-4 about 1 .6-1 .9, 1 .0-1 .2, 0.9-1 . 1 mm long; rostrum
terminating between metacoxae.
Pronotum 6.8-8. 1 mm wide across humeri, 2.5-3. 1 mm long at meson.
Humeri not or little produced laterad, rounded. Cicatrices immaculate.
Scutellum 4.3-5. 1 mm wide at base, 4. 7-5. 8 mm long, without markings.
Abdominal tubercle rudimentary, not projecting past anterior margin of
second visible sternite. Ostiolar ruga on each side extending about 0.8 dis-
tance from mesial margin of ostiole to lateral margin of thorax.
Posterior margin of basal plates sigmoid from caudo ventral view, slightly
concave between 9th paratergite and mesial angle, convex laterally (Fig. 82).
Posterior wall on each side of genital cup dorsally diagonal with anterior
projection ending in laterally curved black tooth; basad of this tooth a small
denticle present on dorsal rim of posterior wall (Figs. 84, 85). Parameres as
in Figure 86.
Length about 11.5-14.3 mm.
Distribution. Southern Brazil and northern Argentina.
Holotype. 6, labeled (a) “Passo Fundo, 1-14-74, RS, E. Heinrichs” (b)
“224”. Deposited in U.S. National Museum, type no. 75562.
Paratypes. 399, “Museum Leiden, Argentina, Cordoba, Almafuerte,
Calamucrita La Cascada, I. 1965, F. H. Walz” (9 RNH); “Museum Leiden,
Argentina, Cordoba, El Quebracho, Calamucrita, I. 1965. F. H. Walz” (9
RNH); (a) as holotype except date “1-15-74” (b) “218” (9 INHS); (a) “E. E.
Gualba, 1-10-74 RS, E. Heinrichs” (b) “223” (6 LHR); (a) “Carazinho, I-l-
74, E. Heinrichs” (b) “220” (<3 INHS).
Acrosternum (Chinavia) nigridorsatum (Breddin, 1901), New Combination
(Figs. 87-89)
Nezara nigridorsata Breddin, 1901, p. 123; Gaedike, 1971, p. 93 (lectotype
designated).
Diagnosis. Humeral angles obtusely angulate (Fig. 87). Abdominal tubercle
compressed, anterior and ventral margins forming slightly acute angle, reach-
ing posterior limit of metacoxae. Black mark at posterolateral angles of
connexival segments confined to edge of sternite, continuing as black spot
in posterolateral angles of sternites; connexiva narrowly red margined. Spi-
racles black, each located on ivory callus. Ostiolar ruga on each side extending
about 0.8 distance from mesial margin of ostiole to lateral margin of thorax.
Rostrum terminating between metacoxae. Femora light green. Cicatrices and
basal angles of scutellum immaculate. Punctation of dorsum dense, green.
Posterior margin of basal plates projecting into concavity of 9th paratergite
(Figs. 88, 89). Length about 12.5 mm.
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NEW YORK ENTOMOLOGICAL SOCIETY
Figs. 87-92. Figs. 87-89. A. nigridorsatum. 87. Humerus. 88. Genital plates, caudoventral
view. 89. Genital plates, caudal view. Figs. 90-92. A. ecuadorensis. 90. Pygophore, ventral
view. 91. Pygophore, caudal view. 92. Genital cup.
Distribution. Southern Brazil (Curitiba). Known only from lectotype.
Comment. Similar to A. aseadum but differing in having angulate humeri,
a black spot in posterolateral angles of the sternites and connexival segments,
shape of the abdominal tubercle and in the form of the basal plates. The
lectotype, a 9 designated by Gaedike (1971), was examined.
Acrosternum (Chinavia) ecuadorensis, new species
(Figs. 90-92)
Description. Light green above and below with narrow yellowish orange
outer margins. Cicatrices and scutellum immaculate. All antennal segments
banded distally with black or fuscous; these bands broadest on last 3 seg-
ments, covering about three-, five-, and five-tenths of last 3 segments, re-
spectively. Black spot present on superior surface of antennifers. Spine at
posterolateral angles of abdominal segments black. Spiracles black, unat-
tended by callus. Legs green. Dorsal punctation moderately dense, fine, green.
Head 2.8 mm wide across eyes, 2. 1 mm long. Outer jugal margins sigmoid,
moderately concave before eyes. Antennal segments 0.5, 1.1, 1.3, 1.9, 1.9
VOLUME 91, NUMBER 2
135
mm long. Rostral segments 2-4 about 1.7, 1.0, 0.9 mm long; apex lying
between metacoxae.
Pronotum 6.7 mm wide at humeri, 2.8 mm long at meson. Humeri scarcely
produced laterad, rounded; anterolateral pronotal margins nearly straight.
Scutellum 4.3 mm wide at base, 4.8 mm long. Coria ending near posterior
margin of penultimate connexival segment, costal angle rounded.
Abdominal spine compressed, reaching posterior limit of metacoxae. Os-
tiolar ruga on each side extending about three-fourths distance from mesial
margin of ostiole to lateral thoracic margin.
Pygophore with somewhat sinuous V-shaped emargination from both ven-
tral and caudal views (Figs. 90, 91). Semicircular mesial projection on ventral
surface of pygophore densely setose. Dorsal rim of posterior wall of genital
cup curved on each side from dorsal view, with several black denticles of
which largest located at mesial corner of rim (Fig. 92).
Female unknown.
Length about 9.5 mm.
Distribution. Ecuador.
Holotype. S, labeled “Ecuador, 25 km. W. Puyo. April 27, 1978. CW &
LB O’Brien & Marshall”. Deposited in U.S. National Museum, type no.
72131. No paratypes.
Acrosternum (Chinavia) ubicum, new species
(Figs. 93-99)
Description. Light green, below often blending to yellow mesially, with
narrow dorsal border of red to yellow along lateral margins of head, prono-
tum, coria basally and abdomen. Connexiva usually with small black spot
at posterolateral angles of segments, this spot sometimes reduced and con-
fined to protruding angle. Antennae green, usually with apex of segment 3
and distal half of segments 4 and 5 dark. Legs green. Spiracles usually black,
each located on conspicuous yellow callus. Dorsal punctation fine, dense,
green.
Head 2. 5-3.0 mm wide across eyes, 2. 0-3. 5 mm long. Antennal segments
0.5, 1.1-1. 4, 1.2-1. 6, 1.5-1. 7, 1.5-1. 7 mm long. Rostral segments 2-4 about
1 .4-1 .8, 1 .0-1 .2, 0.9-1 . 1 mm long; rostrum terminating between metacoxae.
Pronotum 6. 5-8. 2 mm wide at humeri, 2.4-3. 3 mm long at meson. Hu-
meri rounded, not produced. Cicatrices usually immaculate, rarely with
small dark dot at mesial limits.
Scutellum 4. 0-5. 3 mm wide at base, 4. 4-5. 6 mm long, without markings.
Abdominal tubercle compressed, not or barely reaching metacoxae; an-
terior margin subvertical. Ostiolar ruga on each side extending about 0.7-
0.8 distance from inner margin of ostiole to lateral margin of thorax.
Most posterior portion of basal plates at mesial angles (Fig. 93); posterior
136
NEW YORK ENTOMOLOGICAL SOCIETY
Eigs. 93-99. A. ubicum. 93. Genital plates, caudoventral view. 94. Spermathecal bulb and
pump. 95. Pygophore, caudal view. 96. Posterior wall, right side, variation in dorsal rim. 97.
Same, 98. Genital cup. 99. Paramere.
edge from caudal view flattened above 9th paratergite. Spermatheca as in
Figure 94.
Posterior wall of genital cup approximately transverse, expanded dorsad,
mesially curved cephalad on each side of deep mesial emargination (Fig.
95); posterior surface prominently armed with short curved carina on each
side; posteroventral surface with stout median tooth. Dorsal rim of posterior
wall bearing several black denticles; cephalic surface on this wall with several
smaller denticles (Fig. 98); usually one large tooth present on dorsal rim (Fig.
95), sometimes 2 large teeth separated by notch (Fig. 96), or notch without
adjacent large tooth or teeth (Fig. 97). Parameres as in Figure 99.
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137
Length about 10.6-13.7 mm.
Distribution. Hispaniola, Colombia, Guyana, Surinam, Ecuador, Bolivia,
Brazil, Galapagos Islands. Presumably present in other West Indian islands
besides Hispaniola.
Holotype. S, labeled (a) “Bolivia: Dept. Beni, Rio Itez, Pampa de Meio.
IX- 1 1-13-1964” (b) “J. K. Bouseman, J. Lussenhop, Collectors”. Deposited
in American Museum of Natural History.
Paratypes. 699, 10(3(3, (a) “Colombia, Buenaventura XI-6-1950” (b) “Mich-
elbacher and Ross” (9, $ LHR); “Brazil: Ceara State, Barbalha. V-1969. M.
Alvarenga” (299, 2(3(3 AMNH); (a) “British Guiana” (b) “Babcock Coll’n” (9
RNH); “Suriname, Paramaribo, 16-18 VII-75. Coll: D. Engleman” (9 HDE);
“Brazil: Est. Rio Muriqui Mangaratiba. July, 1969. M. Alvarenga” (9 LHR);
(a) “27-III-1925” (b) “Galapagos” (c) “Gift of New York Zoo. Soc. Dept.
Tropical Research, William Beebe, Dir.” (2(3(3 AMNH), “Guaruja Is. Sao
Paulo, Brazil IV- 17- 1966” (b) “Collr. C. A. Triplehorn” ((3 OSU); “Ecuador,
Guayas, Boliche. Oct. 13, 1976 M. E. Irwin, soybean breeding line trial seed
maturation. 1 298” {$ INHS); “Mallares. 23 • 3 • 66. J. Alva N.” (b) “Meunprg.
No. 1160-68” ((3 JG); (a) “La Vega Prov., Rep. Dominicana, ll-X-1967”
(b) “L. H. Rolston, Collector” {26$ LHR).
Acrosternum {Chinavia) laeturn (Stal, 1859)
(Figs. 100-105)
Rhaphigaster laetus Stal, 1859, pp. 228-229.
Nezara {Acrosternum) laeta: Stal, 1872, p. 42 (keyed, descriptive note).
Acrosternum laeturn: Froeschner, 1981, p. 67.
Diagnosis. Humeri rounded, not produced. Abdominal tubercle com-
pressed, reaching posterior margin of metacoxae. Connexiva immaculate or
variously marked with black: minute dot at posterolateral angles of segments,
or narrow border along posterior margin of segments, or macule lying across
transverse sutures. Each spiracle located in posterolateral edge of small yel-
low callus; peritremes red to yellow. Ostiolar ruga on each side extending
0.6-0. 8 distance from mesial margin of ostiole to lateral margin of thorax.
Rostrum ending between or slightly caudad of metacoxae. Femora green.
Small black dot present at mesial limit of cicatrices, another usually present
at lateral limit. Posterior margin of basal plates slanting, most posterior part
at mesial angle (Fig. 100). Spermatheca as in Figure 101. Male genitalia as
in Figures 102-105. Posterior wall on each side of genital cup dorsally
diagonal; large denticle on dorsal rim apposed by acute dorsolateral angle
of pygophore; posterolateral wall of pygophore deeply emarginated (Fig.
103).
Distribution. Northern Chile, Peru, Ecuador.
Types. Lectotype, here designated 6 labeled (a) “Puna” (b) “Kinb” (c)
138
NEW YORK ENTOMOLOGICAL SOCIETY
Figs. 100-105. A. laetum. 100. Genital plates, caudoventral view. 101. Spermathecal bulb
and pump. 102. Posterior margin of exserted pygophore, ventral view. 103. Pygophore, caudal
view. 104. Genital cup. 105. Paramere.
‘‘Type” (d) “Typus” (e) “305/73” (f) “Riksmuseum Stockholm”. Paralec-
totype, 9 labeled as lectotype except (d) “Allotypus” and (e) “304/73”.
Comment. The degree of variability of connexival markings in this species
seems unique within the genus in the hemisphere.
Acrosternum (Chinavia) simplicis, new species
(Figs. 106-110)
Description. Green above with lateral margins of head, pronotum, coria
basally and connexiva narrowly bordered in yellow or red, when red with
very edges of body yellowish in lateral view. Scutellum with 4 small yellowish
spots or transverse macules along base; apex usually bordered narrowly with
yellow. Connexiva with small black mark on edge of sternites at postero-
lateral angles of each segment, this mark sometimes continuing vaguely onto
first laterotergite. Venter paler green than dorsum, becoming yellowish me-
sially. Only peritreme of spiracles dark; each spiracle located in lateral edge
of ivory callus. Black mark at posterolateral angles of sternites small. Femora
VOLUME 91, NUMBER 2
139
Figs. 106-110. A. simplicis. 106. Head. 107. Pygophore, ventral view. 108. Pygophore,
caudal view. 109. Genital cup. 110. Paramere.
green. Small black line or macule on superior surface of antennifer. Antennae
green, sometimes variously blended with fuscous on distal 2 segments. Dor-
sal punctation dense; punctures concolorous with surrounding area.
Apex of head narrowly rounded; lateral jugal margins weakly concave
before eyes (Fig. 106). Width of head across eyes 2.6 mm, length 2.0-2. 1
mm. Antennal segments 0.5, 1.0-1. 1, 1.2, 1.5-1. 6, 1.4-1. 6 mm long. Rostral
segments 2-4 about 1.4-1. 6, 1.2-1. 3, 0.9 mm long; rostrum terminating
between metacoxae.
Pronotum 6. 6-8.0 mm wide at humeri, 2.6-2. 8 mm long at meson. Ci-
catrices immaculate. Humeri slightly produced laterad, rounded.
Scutellum 4.2-4. 5 mm wide at base, 4.4-4. 7 mm long. Basal angles im-
maculate.
Abdominal tubercle barely reaching posterior margin of metacoxae. Os-
tiolar ruga on each side reaching halfway from mesial margin of ostiole to
lateral margin of thorax.
Female unknown.
Posterior wall of genital cup on each side of deep mesial emargination
diagonal dorsally; weakly concave dorsal rim bearing several tiny black
denticles (Figs. 108, 109). Parameres as in Figure 1 10.
Length about 10.0-1 1.3 mm.
Distribution. Paraguay.
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NEW YORK ENTOMOLOGICAL SOCIETY
Holotype. S, labeled “Museum Leiden, Dr. B. Podtiaguine, Assomption
Omgev., Paraguay, 22 Feb. -25 Apr. 1936”. Deposited in Rijksmuseum van
Natuurlijke Historie, Leiden, Netherlands.
Paratypes. 365, labeled as holotype with second label “Cat. No. 9” (6
RNH); labeled as holotype (6 RNH; 6 LHR).
Acrosternum (Chinavia) euri, new species
(Figs. 111-119)
Description. Green above with a liberal am.ount of yellow on interstices
between punctures except on head; lighter green below blending to yellow
mesially at least on thorax. Lateral dorsal and ventral margins of head,
pronotum, coria basally and connexiva crimson; protruding angles of ab-
dominal segments black. Antennae green; black spot present at base of an-
tennifers on superior surface. Very apex of femora, base or all of tibiae, tarsi,
and often rostrum crimson or suffused with crimson. Spiracles narrowly
ringed with black and suffusion of crimson or crimson alone. Neither ci-
catrices nor basal angles of scutellum marked with black. Dorsal punctation
dark green, dense on pronotum and head, less so on scutellum and thorax.
Apex of head moderately rounded; lateral margins ofjuga weakly concave.
Width of head across eyes 2. 7-2. 9 mm, length 2. 2-2. 3 mm. Antennal seg-
ments 0.5, 1.0-1. 2, 1.2-1. 4, 1.6-1. 7, 1.5-1. 6 mm long. Rostral segments 2-
4 about 1 .4-1.6, 1 .2-1.3, 0.9-1 .0 mm long; apex terminating between meta-
coxae.
Pronotum 6. 6-7. 3 mm wide at humeri, 2. 6-2. 9 mm long. Humeral angles
somewhat produced laterad (0.3 mm or less beyond base of coria), narrowly
rounded. Anterolateral margins of pronotum nearly straight.
Scutellum 4. 2-4. 7 mm wide at base, 4. 7-5. 5 mm long.
Abdominal tubercle compressed, reaching between metacoxae. Ostiolar
ruga on each side extending about halfway from mesial margin of ostiole to
lateral margin of thorax.
Posterior margin of basal plates projecting strongly caudad at rounded
mesial angle (Fig. Ill), from caudal view curving ventrad at lateral portion
of 2nd gonacoxae (Fig. 1 12). Spermatheca as in Figure 1 13.
Male genitalia as in Figures 114-119. Posterior wall on each side of genital
cup with stout hook at anteromesial corner of dorsal rim, hook curving
laterad at apex (Figs. 115, 116).
Length about 1 1.1-12.8 mm.
Distribution. Bahama Islands (Mayaguana, Grand Bahama?).
Holotype. 6, labeled “Bahama, VII-28- 1 926”. Deposited in American Mu-
seum of Natural History.
Paratypes. 16, 399, labeled as holotype (6, 9 LHR; 9 AMNH); “Bahamas:
Mayaguana Is. 30-VIII-63, C. Murvosh, black light trap” (9 FSCA).
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141
Figs. 111-119. A. euri. 111. Genital plates, caudoventral view. 112. Same, caudal view.
113. Spermathecal bulb and pump. 1 14. Pygophore, ventral view. 115. Pygophore, caudal view.
116. Genital cup. 117. Paramere. 118. Aedeagus, dorsal view. 119. Aedeagus, lateral view;
conjunctiva (C); median penial lobes (MPL); penisfilum (P); basal plate (BP).
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NEW YORK ENTOMOLOGICAL SOCIETY
Acrosternum (Chinavia) macdonaldi, new species
(Figs. 120-123)
Description. Bright green above, paler below, yellowish on thoracic sterna
and sometimes on abdominal disk. Lateral dorsal and ventral borders of
head, pronotum, coria basally, connexiva and apical borders of scutellum
orange-yellow; connexival borders sometimes suffused over much of latero-
tergites. Cicatrices and basal angles of scutellum immaculate. Antennae green
with apical end of segment 3 and apical half of segments 4 and 5 black;
antennifers rarely with black dot on superior surface near base. Legs green.
Spiracles pale without callus. Punctation dense, fine, green.
Head 2. 8-3.0 mm wide across eyes, 2. 3-2.4 mm long. Antennal segments
0. 5-0.6, 1.1, 1.3-1. 5, 1.9-2. 1, 2. 1-2.2 mm long. Rostral segments 2-4 about
1 .6-1.7, 1 .3-1.4, 1 .0-1 . 1 mm long; rostrum extending a little past metacoxae.
Pronotum 7. 1-7.3 mm wide at humeri, 2. 8-2. 9 mm long. Humeri slightly
produced laterad, rounded.
Scutellum 4. 5-4. 6 mm wide at base, 4. 8-5.0 mm long.
Abdominal tubercle compressed, reaching between metacoxae. Ostiolar
ruga on each side extending about three-fourths of distance from mesial
margin of ostiole to lateral margin of thorax.
Male genitalia as in Figures 1 20-123. Posterior wall on each side of genital
cup dorsally diagonal, dorsal rim concave; denticulate low ridge running
from obtuse mesial corner of dorsal rim onto concave anterolateral surface
of posterior wall (Fig. 122).
Basal plates from caudo ventral view with posterior margin nearly straight
between convex lateral angles; dorsal edge of each slightly sinuous from
caudal view, the two together bowed a little ventrad.
Length about 11.8-12.3 mm.
Distribution. Panama.
Holotype. 3, labeled (a) '‘Canal Zone: Barro Colorado, 3 IV” (b) “W. M.
Wheeler” (c) "light”. Deposited in American Museum of Natural History.
Paratypes. 366, 19, “Panama” (6 LHR); “Coco Solo Hospital, Canal Zone,
27 Apr. 72 L. T. Col: D. Engleman” (6 HDE). “Panama, dist. Chepo, Altos
de Maje. 17 May 75 at lights Stockwell-Engleman” (6 HDE); “Coco Solo
Hosp. C. Z. Panama. Light Trap. 20-V-75. Col: D. Engleman” (9 LHR).
Comment. This species is dedicated to my friend and colleague F. J. D.
McDonald. He has contributed much to our knowledge of pentatomoids.
Acrosternum (Chinavia) wygodzinskyi, new species
(Figs. 124-127)
Description. Dark green above, paler below. Lateral margins of head and
usually anterolateral margins of pronotum thinly bordered in red or yellow.
VOLUME 91, NUMBER 2
143
Figs. 120-127. Figs. 120-123. A. macdonaldi. 120. Pygophore, ventral view. 121. Pygo-
phore, caudal view. 122. Genital cup. 123. Paramere. Figs. 124-127. A. wygodzinskyi. 124.
Pygophore, ventral view. 125. Pygophore, caudal view. 126. Genital cup. 127. Paramere.
Posterolateral angles of connexiva with black mark on edge of sternites;
corresponding angles of sternites more generously blackened ventrally. Ci-
catrices and basal angles of scutellum immaculate. Antennae and legs green
or greenish yellow; black macule present at base of antennifers on superior
surface. Peritremes of spiracles brown, surrounded by yellowish uncalloused
ring. Dorsal punctation dense, green.
Head 2. 9-3. 5 mm wide across eyes, 2.4-2. 9 mm long; lateral jugal margins
weakly concave. Antennal segments 0. 5-0.6, 1.2-1. 5, 1. 5-2.0, 2. 0-2. 3, 1.8-
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NEW YORK ENTOMOLOGICAL SOCIETY
2.1 mm long. Rostral segments 2-4 about 1. 6-2.0, 1.3-1. 6, 1.0-1. 2 mm
long; rostrum terminating between metacoxae.
Pronotum 7. 6-9. 6 mm wide at humeri, 3.0-3. 8 mm long at meson. Hu-
meri produced laterad 0.2-0. 3 mm beyond costal margin of coria; humeral
angle at least slightly obtuse, narrowly rounded; anterolateral margin of
pronotum nearly straight.
Scutellum 4. 7-6.0 mm wide at base, 5. 3-6. 7 mm long.
Abdominal tubercle compressed, projecting between metacoxae. Ostiolar
ruga on each side reaching 0.7-0. 8 distance from mesial margin of ostiole
to lateral margin of thorax.
Posterior margin of basal plates sinuous from caudoventral view, each
slightly concave below second gonocoxae, broadly rounded at lateral angles.
Male genitalia as in Figures 1 24-127. Posterior wall on each side of genital
cup dorsally diagonal, rim shallowly concave; denticulate carina on antero-
lateral face of posterior wall curving posterolaterad from anterior angle of
posterior wall; at base of this carina another thin low carina proceeds an-
terolaterad to lateral margin of pygophore (Fig. 126).
Length about 12.2-17.0 mm.
Distribution. St. Thomas and St. John Islands (Virgin Islands, West Indies).
Holotype. S, labeled “St. Thomas V.I., Est. Lilliendahls El. 1000', 15-X-
78, M. A. Ivies, Coll.” Deposited in U.S. National Museum, type no. 75563.
Paratypes. SSS, 999, same data as holotype (S AMNH); same data as ho-
lotype except date “25-X-78” {2S6 DBT); same data as holotype except date
“24-VIII-78” (S LHR); (a) “Virgin Is: St. Thomas. Est. Lilliendahl. 08 Aug.
1980. At Light” (b) “M. A. Ivie, Coir.” (5 MAI); (a) “Virgin Is.: St. John
V.I.E.R.S. 1 5 Aug. 1 980. At UV Light” (b) “M. A. Ivie, Coir.” (S, 499 UCD);
(a) “Virgin Is. French. Bay Estate. El. 360 ft. St. Thomas. 30 Sep. 1978” (b)
“C. Petrovic, coir.” (S OSU); same data except date “03 Oct. 1978” (3, 9
LHR; 9 UCD); “Virgin Is.: St. Thomas. Estate Lilliendahl. 1000 ft. Oct.
1978, M. A. Ivie, Coir.” (9 MAI); “Virgin Is.: St. Thomas. Frenchman Bay
Estate. 09 July 1978. 750 ft. M. A. Ivie, Coir.” (9 OSU); (a) “Virgin Is. St.
Thomas. Red Hook 14 Oct. 1979” (b) “M. A. Ivie, Coir.”
Comment. This species is similar to the preceding, differing in details of
coloration and the genitalia.
The species is dedicated to Dr. P. Wygodzinsky, whose contributions to
the knowledge of Hemiptera are many.
Acrosternum (Chinavia) monticola, new species
(Figs. 128-130)
Description. Pale green above and below with narrow orange-yellow border
on lateral margins of head, pronotum, base of coria and connexiva. Pos-
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145
terolateral angles of connexiva and sternites with small black spot confined
on connexiva to edge of sternites except on first two segments, there barely
extending onto laterotergites. Connexiva and basal angles of scutellum im-
maculate. Legs and antennae green, antennifers immaculate. Spiracles pale,
surrounded by pale ivory uncalloused area. Dorsal punctation dense, shal-
low, pale green.
Head 2.9 mm wide across eyes, 2.2 mm long. Antennal segments 0.5, 1.1,
1.5, 1.9, 1.9 mm long. Rostral segments 2-4 about 1.7, 1.1, 0.9 mm long;
rostrum terminating between metacoxae.
Pronotum 7.4 mm wide at humeri, 2.9 mm long at meson. Humeri little
produced, obtusely angulate. Anterolateral margins of pronotum straight.
Scutellum 4.6 mm wide at base, 5.0 mm long.
Abdominal tubercle compressed, reaching middle of metacoxae. Ostiolar
ruga on each side extending about 0.7 distance from mesial margin of ostiole
to lateral margin of thorax.
Female unknown.
Male pygophore as in Figures 128-130. Posterior wall on each side of
genital cup dorsally diagonal with laterally bent hook on anterior angle (Figs.
129, 130); concave dorsal rim of posterior walls weakly denticulate.
Distribution. Peru. Known only from holotype.
Holotype. 6, labeled (a) “Peru, 10 Km. S. of Chiclayo. III-21-1961” (b)
“Ross and Michelbacher, Collectors”. Deposited in California Academy of
Sciences. No paratypes.
Acrosternum (Chinavia) occasi, new species
(Fig. 131)
Description. Medium green above, paler below becoming yellowish me-
sially. Narrow yellow lateral border of body sometimes incomplete on head,
entire on pronotum and base of coria, sometimes diffusing over most of
connexiva. Spine at posterolateral angle of connexiva and dot at postero-
lateral angles of sternites black. Cicatrices and basal angles of scutellum
immaculate. Spots (3 or 5) along base of scutellum and narrow apical border
yellow to orange. Antennal segments 4 (except base) and 5, and sometimes
apex of 3 ferrugineous to purple; remainder of antennae green; black macule
present at base of antennifers on superior surface. Each spiracle located in
posterolateral edge of yellow callus; peritremes green. Punctation on dorsum
dense, somewhat rugose on pronotum and scutellum, green; somewhat in-
distinct median longitudinal band on scutellum formed by discontinuous
irregular yellowish calli on interstices between punctures.
Head 2. 7-2.9 mm wide across eyes, 2.4 mm long. Antennal segments 0.5-
0.6, 1.0-1. 2, 1.0-1. 2, 1.3-1. 4, 1.4 mm long. Rostral segments 2-4 about
1.5-1. 6, 1. 1-1.3, 0.9 mm long; rostrum terminating between metacoxae.
146
NEW YORK ENTOMOLOGICAL SOCIETY
Figs. 128-137. Figs. 128-130. A. monticola. 128. Pygophore, ventral view. 129. Pygophore,
caudal view. 130. Genital cup. Fig. 131. /I. occasi. Genital plates, caudoventral view. Figs. 132-
137. A. montivagum. 132. Head. 133. Pygophore, ventral view. 134. Pygophore, caudal view.
135. Genital cup. 136. Posterior wall, left side, dorsomesial view. 137. Paramere.
Pronotum 6. 9-7. 8 mm wide at humeri, 4. 9-5. 7 mm long at meson. Hu-
meri little produced laterad, broadly rounded; anterolateral margins straight.
Scutellum 4. 4-5.0 mm wide at base, 4. 9-5. 7 mm long.
Abdominal tubercle barely or not attaining metacoxae. Ostiolar ruga on
VOLUME 91, NUMBER 2
147
each side extending about 0.7 distance from mesial margin of ostiole to
lateral margin of thorax.
Basal plates forming nearly straight line along posterior margin except at
convex lateral angles (Fig. 131). Paratergite 9 not concave above posterior
margin of basal plates.
Male unknown.
Distribution. Chile (Coquimbo Prov.).
Holotype. 9, labeled (a) “50 Km S. of La Serena, Chile, XII- 1-50” (b)
“Ross and Michelbacher, Collectors”. Right basal plate partially detached,
last antennal segments lacking. Deposited in California Academy of Sciences.
Paratype. “Chile” and “XI”, remainder illegible (9 LHR).
Comment. The abdominal tubercle of this species is nearly as rudimentary
as that of A. aseadum. The basal plates and some markings of the two species
are dissimilar, however.
Acrosternum (Chinavia) montivagum (Distant, 1890)
(Figs. 132-137)
Chlorochroa montivaga Distant, 1890, p. 333, pi. 31, fig. 13.
Nezara majuscula Distant, 1890, p. 339, pi. 31, fig. 20 (synonymized by
Rolston, 1976).
Pentatoma montivaga: Lethierry and Severin, 1893, p. 119.
Acrosternum montivagum: Rolston, 1976, p. 4 (lectotype designated).
Diagnosis. Proximal end of tibiae and broad border along outer margins
of body crimson. Juga converging apically over tylus, rarely contiguous (Fig.
132). Humeri rounded, little produced laterad. Abdominal spine barely at-
taining posterior limit of metacoxae, its anterior margin subvertical. Pos-
terior margins of connexival segments except last black bordered for about
half their width, or black marginal macule mostly on posterolateral angle
but extending onto anterolateral angle of succeeding segment; this macule
continuing onto venter and forming spot at posterolateral angles of sternites.
Distal end of coria ending above penultimate connexival segment. Each
spiracle positioned in posterolateral edge of yellowish callus; peritremes black
or reddish. Ostiolar ruga on each side reaching about 0. 5-0.6 distance from
mesial margin of ostiole to lateral thoracic margin. Rostrum terminating
between meso- and metacoxae. Femora green. Cicatrices and basal angles
of scutellum immaculate. Dorsal punctation dense; punctures concolorous
with surrounding area. Posterior margin of basal plates transverse between
rounded lateral angles from caudoventral view; dorsal margin of each basal
plate slightly convex from caudal view. Male genitalia as in Figures 133-
137. Posterior wall of genital cup diagonal dorsally on each side with shal-
lowly concave rim and line of black denticles on anterolateral face.
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NEW YORK ENTOMOLOGICAL SOCIETY
Length about 13-17 mm.
Distribution. Southern Mexico into Panama, Hispaniola.
Comment. The types of Chlorochroa montivaga and Nezara majiisciila
were examined.
Acrosternum (Chinavia) collis, new species
(Figs. 138-142)
Description. Emerald green above, greenish yellow below with apex of
tibiae and outer margins of body crimson. Posterior margin of connexival
segments bordered with black along lateral half, this border continuing as
black spot at posterolateral angles of sternites. Dot at mesial limit of cica-
trices, basal angles of scutellum, spiracles and wide ring around spiracles,
and antennae, black. Dorsal punctation moderately dense; punctures con-
colorous with surrounding area.
Head 2. 9-3. 3 mm wide across eyes, 2. 2-2. 7 mm long. Antennal segments
0.6-0. 7, 1. 1-1.4, 1.6-2. 1, 1.9-2. 4, 2. 3-2. 5 mm long. Rostral segments 2-4
about 1.6-1. 9, 1. 1-1.4, 0.9-1. 2 mm long; apex of rostrum lying between
metacoxae. Juga not convergent apically.
Pronotum 6. 8-8. 6 mm wide at humeri, 2. 5-3.4 mm long at meson. Hu-
meri little produced laterad, broadly rounded to obtusely angulate. Antero-
lateral margin more or less straight, sometimes weakly concave or sinuous.
Scutellum 4. 2-5. 3 mm wide at base, 4.4-6. 1 mm long. Coria ending dis-
tally above anterior part of last connexival segment.
Abdominal tubercle reaching posterior margin of metacoxae. Ostiolar ruga
on each side extending about 0.7-0. 8 distance from mesial margin of ostiole
to lateral margin of thorax.
Genital plates as in Figure 138. Paratergites 9 flat basally.
Large obtuse tooth projecting posterodorsad from posterolateral angles of
pygophore. Posterior wall of genital cup dorsally diagonal on each side;
posterolateral and anteromesial angles of dorsal rim with smaller tooth pro-
jecting laterad (Figs. 140, 141).
Length about 11.5-15.5 mm.
Distribution. Costa Rica.
Holotype. 3, labeled “Costa Rica, Puntar. - Alajuela: Monteverde For. Res.
1600 m. 17/18 Aug. 1976. E. M. Fisher.” Deposited in U.S. National Mu-
seum, type no. 75564.
Paratypes. 13, 299, labeled as holotype (3, 9 DBT); “Costa Rica: San Jose,
8.3 mi. N. San Isidro del General, 30 June 1972. R. R. & M. E. Murray”
(9 LHR).
Comment. This species is distinguished from A. montivagum, which it
resembles superficially, by the black markings, rostral length and male gen-
italia.
VOLUME 91, NUMBER 2
149
Figs. 138-142. A. collis. 138. Genital plates, caudoventral view. 139. Pygophore, ventral
view. 140. Pygophore, caudal view. 141. Posterior margin of pygophore, dorsal view. 142.
Paramere.
Acrosterniim (Chinavia) apicicorne (Spinola., 1852)
(Figs. 143-147)
Pentatoma apicicorne Spinola in Gay, 1852, pp. 131-132, pi. 1, fig. 1.
Nezara apicicornis: Signoret, 1863, p. 548, pi. 1, fig. 1 (description); Reed,
1898, p. 135 (synonymy); Haglund, 1899, p. 77; Berg, 1900, p. 83 (corrects
Reed’s synonymy).
Rhaphigaster apicicornis: Walker, 1867, p. 360 (listed).
Acrostcrnum apicicornis: Piran, 1948, p. 9 (record).
Diagnosis. Humeral angles rounded, little produced. Abdominal tubercle
scarcely reaching metacoxae. Posterolateral angles of connexiva and sternites
black, macule on connexiva including part of laterotergite and sometimes
anterolateral angle of following connexival segment. Spiracles green to brown,
each located in posterolateral edge of yellow callus. Ostiolar ruga on each
side extending about 0.7-0. 8 distance from mesial margin of ostiole to lateral
thoracic margin. Rostrum terminating between metacoxae. Femora green.
Cicatrices immaculate. Small yellow callus located adjacent to basal angles
of scutellum; 3 additional yellow spots present along scutellar base. Dorsal
punctation dense, concolorous with surrounding area. Posterior margin of
basal plates nearly transverse between rounded lateral angles; spermatheca
as in Figure 147. Male genitalia as in Figures 143-146. Posterior wall of
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NEW YORK ENTOMOLOGICAL SOCIETY
Figs. 143-147. A. apicicorne. 143. Genital cup. 144. Pygophore, ventral view. 145. Pygo-
phore, caudal view. 146. Paramere. 147. Spermathecal bulb and pump.
genital cup diagonal dorsally on each side with small black tooth at antero-
mesial angle of rim curving anterolaterad; from this tooth a subvertical
denticulate black carina descends anterolateral face of wall, with largest
denticles subterminal. Pygophore deeply emarginated subterminally from
lateral view.
Length about 1 1.5-13.0 mm.
Distribution. Chile, Argentina (fide Piran, 1948).
Acrosternum (Chinavia) australe, new species
(Figs. 148-152)
Description. Green above and below with narrow yellow to orange-yellow
border along outer margins, usually excepting head apically; thoracic sterna,
coxae, trochanters, femora and abdominal tubercle yellowish, often with
rufous suffusion. Posterolateral angles of connexiva including corresponding
corner of laterotergites and posterolateral angle of sternites black. Cicatrices
and basal angles of scutellum immaculate. Spot adjacent to each basal angle,
usually 2 additional spots along base, and apex of scutellum yellowish. Spi-
racles green, each contiguous with but not on a small yellowish callus. Dorsal
punctation dense, rugose on pronotum; punctures concolorous with sur-
rounding area.
Head 2. 7-3.0 mm wide across eyes, 2. 2-2. 7 mm long. Antennal segments
0. 5-0.6, 1.0-1. 2, 1.3-1. 7, 1.7-1. 9, 1.3-1. 6 mm long. Rostral segments 2-4
VOLUME 91, NUMBER 2
151
Figs. 148-152. A. australe. 148. Genital cup. 149. Pygophore, ventral view. 150. Pygophore,
caudal view. 151. Paramere. 152. Spermathecal bulb and pump.
about 1.3-1. 6, 1.2-1. 6, 0.8-1. 1 mm long; rostrum terminating between or
projecting a little past metacoxae.
Pronotum 6. 2-8. 2 mm wide at humeri, 2.3-3. 1 mm long at meson. Hu-
meri rounded, little produced laterad. Anterolateral margins straight.
Scutellum 3. 9-5. 2 mm wide at base, 4. 2-5. 6 mm long.
Abdominal tubercle projecting little past posterior limit of metacoxae.
Ostiolar ruga on each side extending about three-fourths distance from me-
sial margin of ostiole to lateral thoracic margin.
Posterior margin of basal plates slightly sinuous, their most posterior part
mesial, there bent slightly dorsad toward 2nd gonocoxae. Spermatheca as
in Figure 152,
Male genitalia as in Figures 148-151. Posterior wall of genital cup diagonal
dorsally on each side; dorsal rim with denticle at anteromesial and postero-
lateral corners, concave between these corners and usually with one or more
small denticles along concavity (Figs. 148, 150).
Length about 10-14 mm.
Distribution. Northern Argentina.
Holotype. S, labeled (a) “5 mi. N. of Dean Fumes, Cordoba, Arg. II-8-51”
(b) “leg. Ross and Michelbacher”. Costal area of left hemelytron swollen
above posterior half of first and anterior half of second connexival segments.
Deposited in American Museum of Natural History.
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NEW YORK ENTOMOLOGICAL SOCIETY
Paratypes. 699, 7(5(5, “Museum Leiden, Argentina, La Rioja, Mascasin, II
1964, F. H. Walz.” (399, 3(5(5 RNH; 9 LHR); “Museum Leidn., Argentina,
Bs. Aires, Fandel, II 1953, F. H. Walz” (9, 3(5(5 RNH; <5 LHR); “Cruz del
Eje. Argentina” (9 HDE).
Comment. The contiguous callus and spiracle arrangement, rather than
the spiracle on the callus, are diagnostic.
Acrosternum (Chinavia) fiiscopunctatum (Breddin, 1901), New Combination
(Figs. 153-156)
Nezara fuscopunctata Breddin, 1901, pp. 123-124; Gaedike, 1971, p. 86
(lectotype designated).
Diagnosis. Pronotum transversely impressed caudad of cicatrices. Coria
nearly reaching posterior margin of last connexival segment; costal angles
of coria subacute (Fig. 153). Humeri rounded, not produced, with short
black line on edge. Abdominal tubercle compressed, reaching middle of
metacoxae. Posterolateral angles of connexiva with triangular black mark
extending onto laterotergite; posterolateral angles of sternites bearing small
black spot. Spiracles light brown, unattended by callus. Ostiolar ruga on
each side extending about 0.65 distance from mesial margin of ostiole to
lateral thoracic margin. Rostrum terminating between metacoxae. Femora
entirely green. Lateral jugal margin moderately concave (Fig. 154). Neither
pronotum nor scutellum with black markings. Dorsal punctation dense,
concolorous with area surrounding punctures. Mesial angle of basal plates
rounded; most posterior part of plates near this angle; posterior margin rather
abruptly reflexed toward 9th paratergites (Figs. 155, 156). Length about 10.7
mm.
Distribution. Brazil (Espirito Santo). Known only from lectotype.
Comment. The lectotype, a 9 designated by Gaedike (1971), was examined.
Acrosternum {Chinavia) marginatum (Palisot de Beauvois, 1805)
(Figs. 157-163)
Pentatoma marginata Palisot de Beauvois, p. 147, Hem. pi. 10, fig. 1.
Nezara marginata: Amyot and Serville, 1843, p. 145; Distant, 1880, p. 79,
pi. 7, fig. 23 (records); Distant, 1890, p. 339 (records); Sharp, 1890, p.
408, pi. 13, fig. 1 7 (5 genitalia); Uhler, 1894a, pp. 1 75-176 (record); Uhler,
1894b, p. 232 (record); Van Duzee, 1904, p. 58 (keyed, records); Van
Duzee, 1907, p. 10 (records).
Rhaphigaster spirans Dallas, 1851, p. 280 (synonymized by Stal, 1872).
Pentatoma {Nezara) marginata: Guerin in Sagra, 1857, p. 374.
Nezara juriosa Stal, 1862, p. 106 (synonymized by Stal, 1872).
Nezara {Acrosternum) marginata: Stal, 1872, p. 42 (keyed, synonymy).
Figs. 153-156. A. fuscopunctatum. 153. Hemelytron. 154. Head. 155. Genital plates, cau-
doventral view. 156. Basal plates, caudal view (BP).
Acrosternum marginatum: Bergroth, 1914, p. 25; Van Duzee, 1917, p. 60
(catalog); Barber, 1923, p. 12 (record); Blatchley, 1926, pp. 160, 163 (keyed,
description); Barber and Bruner, 1932, p. 262 (Cuban record, descriptive
note); Barber, 1939, p. 296 (record); Torre Bueno, 1939, p. 237 (keyed);
Wolcott, 1948, p. 194 (records); Alayo, 1967, pp. 27-28, pi. 1, fig. 3, pi.
9, fig. 5; Gaud and Martorell, 1974, p. 258 (records); Froeschner, 1981,
p. 67 (catalog).
Diagnosis. Humeri little produced laterad, rounded. Abdominal tubercle
compressed, reaching middle of metacoxae. Posterolateral angles of con-
nexival segments including corner of laterotergites and sternites with black
spot. Spiracles unattended by callus; peritremes narrowly and usually in-
completely ringed with black. Ostiolar ruga on each side extending about
three-fourths distance from mesial margin of ostiole to lateral margin of
thorax. Rostrum terminating between metacoxae. Femora green. Cicatrices
and basal angles of scutellum immaculate. Dorsal punctation dense, con-
colorous with area surrounding punctures. Basal plates with projection of
154
NEW YORK ENTOMOLOGICAL SOCIETY
Figs. 1 57-1 63. A. marginatum. 157. Genital plates, caudoventral view. 158. Basal plates,
caudal view (BP). 159. Pygophore, ventral view. 160. Pygophore, caudal view. 161. Genital
cup. 162. Paramere. 163. Spermathecal bulb and pump.
posterior margin reflexed toward 9th paratergites; margin between lateral
angles and projection diagonal, between projection and mesial angles convex
(Figs. 157, 158). Spermatheca as in Figure 163. Posterior margin of pygo-
phore broadly and sinuously emarginate from ventral view (Fig. 159); pos-
terior wall of genital cup dorsally diagonal on each side, rim finely dentic-
ulate, concave, bearing small tooth at anteromesial angle (Figs. 160, 161).
Parameres as in Figure 162. Length about 1 1.5-17.0 mm.
Distribution. Southwestern U.S. into Colombia, Ecuador and Venezuela;
and from Florida as far south as Guadeloupe in the West Indies.
Comment. This is the common species of Acrosternum in Middle America.
Both sexes are readily distinguished from A. hilare, with which it is partially
sympatric and often confused.
VOLUME 91, NUMBER 2
155
Figs. 164-170. A. hilare. 164. Genital plates, caudoventral view. 165. Basal plates, caudal
view (BP). 166. Pygohpore, ventral view. 167. Pygophore, caudal view. 168. Genital cup. 169.
Paramere. 170. Spermathecal bulb and pump.
Acrosternum (Chinavia) hilare {Say, 1831)
(Figs. 164-170)
Pentatoma hilaris Say, 1831, p. 5; Say, 1832, pp. 9-10; Say, 1859, pp. 304-
305, 316-317.
Rhaphigaster sarpinus Dallas, 1851, p. 276 (synonymized by Uhler, 1878).
Nezara {Acrosternum) sarpinus: StM, 1872, p. 43 (keyed).
Nezara hilaris: Uhler, 1878, p. 380 (synonymy); Uhler, 1893, p. 368 (record);
Van Duzee, 1 894, p. 1 72 (record); Van Duzee, 1 904, p. 58 (keyed, records);
Barber, 1906, p. 261 (record); Torre Bueno, 1908, p. 225 (host, records);
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NEW YORK ENTOMOLOGICAL SOCIETY
Van Duzee, 1909, p. 157 (record); Olsen, 1912, p. 55 (hosts, record);
Zimmer, 1912, p. 233 (records); Van Duzee, 1914, p. 5 (record); Barber,
1914, p. 523 (record); Whitmarsh, 1917, pp. 519-552 (biology).
Nezara {Acrosternum) hilaris: Kirkaldy, 1909, p. 119.
Acrosternum hilare: Parshley, 1915, p. 175 (keyed); Parshley, 1917, p. 24
(records); Hart, 1919, p. 181 (keyed, hosts); Stoner, 1920, pp. 107, 108-
109, pi. 7, fig. 2 (keyed, synonymy, description, hosts); Hussey, 1922, p.
15 (records); Parshley, 1923, pp. 767-768 (keyed, records); Blatchley,
1926, pp. 160, 161-162, fig. 33 (keyed, description); Pack and Knowlton,
1930, p. 250 (host, records); Brimley, 1938, p. 63 (record); Froeschner,
1941, p. 130 (keyed); Esselbaugh, 1946, pp. 682-683 (eggs); Esselbaugh,
1948, pp. 41-44 (biology); Russell, 1952, pp. 5, 7-9 (biology); Sailer, 1953,
pp. 70-7 1 (biology); Miner, 1 966, pp. 8-17, figs. 2-5 (biology); McPherson,
1970, p. 54, fig. 51 (keyed, hosts, records); Hoffman, 1971, p. 48, fig. 12
(records); Burks, 1972, pp. 367-368 (parasite); Furth, 1974, pp. 40-41,
pi. 1, fig. 3, pi. 5, figs. 64, 70 (keyed, hosts, records); McPherson and
Mohlenbrock, 1976, pp. 155-156 (hosts, parasites); Nixon, 1976, p. 412
(host).
Acrosternum hilaris: Van Duzee, 1917, p. 60 (synonymy, distribution); Bak-
er, 1931, p. 201, figs. 116, 117 {6 genitalia); Underhill, 1934, pp. 1-26
(biology); Torre Bueno, 1939, p. 236 (keyed).
Diagnosis. Humeri little produced laterad, rounded. Abdominal tubercle
compressed, reaching between metacoxae. Posterolateral angle of connexiva
including corner of laterotergite and of sternites with black spot. Spiracles
green to brown, in posterolateral part of subcalloused and usually yellowish
spot. Ostiolar ruga on each side extending about three-fourths distance from
mesial margin of ostiole to lateral margin of thorax. Rostrum ending between
metacoxae. Femora green. Cicatrices and basal angles of scutellum immac-
ulate. Punctation dense dorsally, green. Posterior margin of basal plates
forming shallow convex arc from caudoventral view, nearly straight from
caudal view (Figs. 164, 165). Spermatheca as in Figure 170. Emargination
of pygophore from ventral view broad, shallow, sinuous (Fig. 166). Posterior
wall dorsally on each side creating triangle with diagonal obtuse ridge pos-
teriorly and transverse ridge anteriorly; rim of latter concave, armed with
several black denticles (Figs. 167, 168). Parameres as in Figure 169. Length
about 1 1-19 mm.
Distribution. Ranging from Ontario and Quebec, Canada, apparently
throughout the United States. A single specimen was seen from Mexico, that
from the state of Jalisco. Uhler’s (1878) record from the West Indies is
dubious, and those from Panama and Brazil are clearly erroneous.
Comment. A. hilare is of some economic importance and there are nu-
VOLUME 91, NUMBER 2
157
Figs. 171-176. A. brasicola. 171. Genital plates, caudoventral view. 172. Basal plates, caudal
view (BP). 173. Pygophore, ventral view. 174. Pygophore, caudal view. 175. Genital cup. 176.
Paramere.
merous references to the species in this connection. Some of these references
have been noted in the synonymy cited, but those of a purely economic
nature have not.
Acrosternum (Chinavia) brasicola, new species
(Figs. 171-176)
Description. Dark green above, paler below, with narrow bright red border
along outer margins. Black spot present at mesial limits of cicatrices and
black fovea in basal angles of scutellum; a small yellow spot usually mesad
of and contiguous with each scutellar fovea. Small black spot in posterolateral
angles of connexiva extending onto laterotergites. Elongate black mark pres-
ent on superior surface at base of antennifers; antennae usually emerald
green with black ring on distal end of segment 3, often much of antennae
apparently discolored, black. Legs green. Spiracles black, each in postero-
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lateral part of callus distinguished in varying degrees by yellowish color.
Dorsal punctation green, fine, dense, less dense on scutellum; many yellowish
small calli scattered on dorsum.
Head 2.9-3. 1 mm wide across eyes, 2. 2-2. 6 mm long. Antennal segments
0. 5-0.6, 1.0-1. 4, 1.3-1. 7, 1. 7-2.1, 2.0-2. 2 mm long. Last 3 rostral segments
about 1.5-1. 7, 1.2-1. 4, 1.0-1. 2 mm long; rostrum reaching metacoxae.
Pronotum 7. 5-9.4 mm wide at base, 4. 8-6. 7 mm long. Humeri rounded,
little or not at all produced laterad. Anterolateral margins straight or weakly
convex.
Scutellum 4. 5-5. 9 mm wide at base, 4. 8-6. 7 mm long.
Abdominal tubercle compressed, projecting little beyond posterior limit
of metacoxae. Ostiolar ruga on each side extending 0.6-0. 8 distance from
mesial margin of ostiole to lateral thoracic margin.
Basal plates subtriangular; posterior margin arched ventrad beneath 9th
paratergites from caudal view (Figs. 171, 172).
Male genitalia as in Figures 173-176. Posterior wall on each side with
tooth near posterior limit; pygophoral margin posterior to this tooth with
field of denticles extending onto dorsal surface of posterolateral projection
on each side (Figs. 174, 175).
Distribution. Southern Brazil.
Holotype. 6, labeled “Brasilien, Nova Teutonia, 27° 1 FB 52° 23'L. Fritz
Plaumann”, on one edge ‘TI 1974”, on other edge “300-500 m”. Deposited
in U.S. National Museum, type no. 75566.
Paratypes. 599, 266, all from same locality: labeled as holotype (299 LHR);
dated “X-3 1-1958” (b) “Ex. Coll. H. Ruckes” {6 AMNH); labeled as pre-
ceding except date “XI-28-195” (c$ LHR); same labeling except date “I. 3.
1962” (9 AMNH); same labeling except date “XI- 12- 1959” (9 AMNH);
labeled as holotype except date “III-3-1960” (9 LHR).
Acrosternum (Chinavia) teretis, new species
(Figs. 177-180)
Description. Light green, below blending to yellow mesially, with reddish-
yellow border along outer margins. Black spot present in posterolateral angles
of connexiva and sternites, those on connexiva extending onto laterotergites,
at mesial and sometimes lateral limits of cicatrices. Small macule on superior
surface at base of antennifers also black; antennae green with black ring at
distal end of segment 3 and short dense setose vestiture giving last 2 segments
brownish appearance. Legs green. Spiracles black, unattended by callus.
Dorsal punctation dense, fine, concolorous with area around punctures.
Head 2. 8-2. 9 mm wide across eyes, 2. 2-2. 3 mm long. Antennal segments
0.6, 1.1-1. 2, 1.5, 1.9, 2.0 mm long. Rostral segments 2-4 about 1.5-1. 6,
1.2-1. 3, 1.1 mm long; apex of rostrum lying between metacoxae.
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159
Figs. 177-185. Figs. 177-180. A. teretis. 177. Pygophore, ventral view. 178. Pygophore,
caudal view. 179. Genital cup. 180. Paramere. Figs. 181-185. A. musivum. 181. Basal plates,
caudal view (BP). 182. Pygophore, ventral view. 183. Pygophore, caudal view. 184. Genital
cup. 185. Paramere.
Pronotum 7. 5-7. 7 mm wide at humeri, 2. 7-2. 9 mm long at meson. Hu-
meri narrowly rounded, somewhat produced laterad. Anterolateral margins
nearly straight or weakly convex.
Scutellum 4. 3-4. 6 mm wide at base, 4.8-5. 1 mm long, without markings.
Abdominal tubercle somewhat compressed, terminating between meta-
coxae. Ostiolar ruga on each side extending 0.6-0. 7 distance from mesial
margin of ostiole to lateral thoracic margin.
Female unknown.
Posterior wall of genital cup extremely produced on each side as flat
protuberance curving dorsolaterad, projecting above tergum; concave dorsal
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rim next to lateral wall denticulate (Figs. 177-179). Parameres projecting
past posterior margin of pygophore, spatulate apically (Fig. 180).
Length about 12-13 mm.
Distribution. Colombia.
Holotype. S, labeled (a) “Colombia; Sierra Nevada de Santa Marta, San
Sebastian de Rabago, 2000 m. April 1-14, 1968” (b) “Borys Malkin, Col-
lector.” Deposited in American Museum of Natural History.
Paratype. 6, labeled as holotype (LHR).
Acrosternum (Chinavia) musivum (Berg, 1878)
(Figs. 181-185)
Nezara musiva Berg, 1878, pp. 26-27; Berg, 1879, pp. 56-57 (reprint).
Acrosternum musiva: Piran, 1948, p. 9 (record); Piran, 1963, p. 337 (record);
Grazia-Vieira and Casini, 1973, p. 58 (record).
Diagnosis. Connexival segments bordered both anteriorly and posteriorly
with black; some or all femora with preapical black band, this often incom-
plete or fragmented; dorsum variously marked with black macules and ver-
miform punctate lines. Apex of scutellum reflexed. Humeral angles rounded,
little produced laterad. Abdominal tubercle not or barely reaching meta-
coxae. Spiracles black, each in posterolateral edge of prominent yellowish
callus. Ostiolar ruga on each side extending about 0.7 distance from mesial
margin of ostiole to lateral thoracic margin. Apex of rostral segment 3 lying
between metacoxae, apex of last segment on sternite 4 (3rd visible). Cicatrices
at least partially black bordered. Large yellowish or green basal callus located
beside each basal angle of scutellum. Many dorsal punctures clustered or in
transverse vermiform lines, these areas of aggregate punctures usually black.
Posterior margin of basal plates emarginated from caudal view at mesial
margin of 9th paratergites (Fig. 181). Male genitalia as in Figures 182-185.
Posterior wall of genital cup on each side reduced to stout hook, this bent
anterolaterad at apex and denticulate along lateral margin (Figs. 183, 184).
Length about 10.5-13 mm.
Distribution. Argentina, Uruguay.
Type. The holotype is preserved at Universidad de La Plata.
Comments. Black borders on both sides of the transverse connexival su-
tures for the entire width of the connexivum coupled with preapical black
femoral bands are diagnostic. The holotype was examined.
Acrosternum (Chinavia) rogenhoferi (Stal, 1872), New Combination
(Figs. 186-190)
Nezara rogenhoferi Stal, 1872, pp. 40-41.
Diagnosis. Both anterior and posterior borders of connexival segments
broadly black bordered; anterior margin of humeri bordered in black, re-
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161
mainder of outer pronotal borders orange-yellow; large yellowish macule at
base of scutellum on each side located about midway between lateral angles
and meson; femora black apically, otherwise green. Humeri rounded, little
or not produced laterad. Abdominal tubercle barely reaching between meta-
coxae. Coria bearing many irregular pale green calli. Spiracles black, unat-
tended by callus. Ostiolar ruga on each side extending from mesial ostiolar
margin 0.65-0.85 distance to lateral thoracic margin. Cicatrices black bor-
dered posteriorly. Basal angles of scutellum immaculate. Dorsal punctation
dense, rather regularly distributed. Posterior margin of basal plates from
caudal view forming shallow V (Fig. 186). Male genitalia as in Figures 187-
1 90. Posterior wall of genital cup dorsally diagonal; dorsal rim slightly convex
and denticulate caudad of moderate sized bent tooth at anterolateral corner
(Figs. 188, 189). Length about 12.5 mm.
Distribution. Southern Brazil.
Comment. This species is easily recognized by the broad, colored border
along the anterolateral pronotal margins. This border is black at the humeri
and otherwise orange-yellow.
Acrosternum (Chinavia) erythrocnemis (Berg, 1878)
(Figs. 191-195)
Nezara erythrocnemis Berg, 1878, pp. 27-28; Berg, 1879, pp. 57-58 (reprint).
Acrosternum erythrocnemis: Piran, 1966, p. 87 (records).
Diagnosis. Connexival segments bordered anteriorly and posteriorly with
black; distal end of femora and at least proximal end of tibiae crimson; outer
border of head crimson, remaining outer border of body yellowish. Humeri
rounded, not produced. Abdominal tubercle compressed, reaching between
metacoxae. Spiracles pale, narrowly ringed with fuscous, not attended by
callus. Ostiolar ruga on each side extending about 0.7 distance from mesial
margin of ostiole to lateral thoracic margin. Rostrum reaching or somewhat
surpassing posterior limit of metacoxae. Cicatrices partially bordered in
black, sometimes entirely black. Basal scutellar angles black. Dorsal punc-
tation dense, punctures concolorous with surrounding area. Basal plates
subtriangular, their posterior margin from caudoventral view forming nearly
straight line between rounded lateral angles. Spermatheca as in Figure 191.
Male genitalia as in Figures 192-195. Posterior wall on each side of genital
cup an obtuse projection with concave anterolateral face and finely dentic-
ulate lateral rim (Figs. 193, 194). Length about 1 1.5-14.5 mm.
Distribution. Southern Brazil, northern Argentina, Uruguay.
Type. The holotype is conserved in the Universidad Nacional de La Plata.
Comments. Of the few species with the transverse connexival suture bor-
dered with black on both sides for at least half the width of the connexivum,
only this species has the femora and tibiae marked with crimson as indicated.
The holotype was examined.
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Figs. 186-190. A. rogenhoferi. 186. Basal plates, caudal view (BP). 187. Pygophore, ventral
view. 188. Pygophore, caudal view. 189. Genital cup. 190. Paramere.
Acrosternum (Chinavia) runaspis (Dallas, 1851)
(Figs. 196-200)
Rhaphigaster runaspis Dallas, 1851, p. 280 (keyed).
Nezara runaspis: StM, 1872, p. 43 (listed).
Acrosternum runaspis: Becker and Grazia- Vieira, 1977, pp. 54-55 (descrip-
tive note, record); Froeschner, 1981, p. 68 (catalog).
Diagnosis. Both anterior and posterior borders of connexival segments
black, dorsum otherwise devoid of black; venter (excluding appendages) with
black only as small macules at base of antennifers and both anterior and
posterior lateral angles of sternites. Legs green. Humeri rounded, not pro-
duced. Abdominal tubercle compressed, reaching between metacoxae. Spi-
racles pale, unaccompanied by callus. Ostiolar ruga reaching about 0.7 dis-
tance from mesial margin of ostiole to lateral thoracic margin. Rostral apex
lying between metacoxae. Dorsal punctation dense, fine, green. Posterior
margin of basal plates from caudoventral view transverse between rounded
lateral angles, rounded dorsoventrally, rather thick from caudal view. Sper-
matheca as in Figure 196. Male genitalia as in Figures 197-200. Posterior
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163
Figs. 191-195. A. erythrocnemis. 191. Spermathecal bulb and pump. 192. Pygophore, ven-
tral view. 193. Pygophore, caudal view. 194. Genital cup. 195. Paramere.
wall of genital cup dorsally diagonal on each side with tooth at anterior angle
bent anterolaterad and small black denticles along concave dorsal rim (Figs.
198, 199). Length about 15-18.5 mm.
Distribution. Argentina (Misiones), Brazil (Bahia, Para, Mato Grosso, San-
ta Catarina, Sao Paulo), Colombia, Ecuador, French Guiana, Paraguay,
Peru (Loreto).
Comment. The type was examined.
Acrosternum (Chinavia) plaumanni, new species
(Figs. 201, 202)
Description. Dark green above; yellowish green below, especially on tho-
racic sterna and abdomen. Outer margins narrowly bordered with orange-
yellow, diffusing mesad along transverse connexival sutures; apex of scutel-
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Figs. 196-202. Figs. 196-200. A. runaspis. 196. Spermathecal bulb and pump. 197. Py-
gophore, ventral view. 198. Pygophore, caudal view. 199. Genital cup. 200. Paramere. Figs.
201, 202. A. plaumanni. 201. Pygophore, ventral view. 202. Pygophore, caudal view.
lum similarly colored. Connexival segments narrowly bordered posteriorly
with black. Spot at base of antennifers on superior surface and spiracles
black, latter in posterolateral edge of paler green subcalloused spot. Cica-
trices, basal angles of scutellum and lateral angles of sternites immaculate.
Antennae green with distal half or somewhat less of last 3 segments fuscous
to black. Legs apparently green (discolored in holotype). Dorsal punctation
dense, hne, green.
Head 2.7 mm wide across eyes, 2.3 mm long. Antennal segments 0.5, 1.1,
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165
1.5, 2.1, 2.0 mm long. Rostral segments 2-4 about 1.6, 1.3, 1.0 mm long;
rostrum reaching posterior limit of metacoxae.
Pronotum 6.7 mm wide at humeri, 2.5 mm long at meson. Humeri round-
ed, slightly produced laterad. Anterolateral margins of pronotum straight.
Scutellum 4.3 mm wide at base, 4.5 mm long.
Abdominal tubercle apically obtuse, scarcely reaching metacoxae. Ostiolar
ruga on each side extending about 0.7 distance from mesial margin of ostiole
to lateral thoracic margin.
Female unknown.
Pygophore incised narrowly between posterior and lateral walls of genital
cup (Figs. 20 1 , 202). Posterior wall of genital cup on each side with diagonal
orientation dorsally; anteromesial corner developed as large tooth bent lat-
erad; dorsal rim concave, roughened but without apparent denticles.
Length about 1 1.6 mm.
Distribution. Ecuador (Pastaza). Known only from holotype.
Holotype. 6, labeled “Ecuador, Pastaza, Ashuara Village on Rio Macuma,
10 km from Rio Morona, 300 m. July 5-16, 1971. B. Malkin.” Deposited
in the American Museum of Natural History.
No paratypes.
Comment. This species is dedicated to Fritz Plaumann, whose industry
in collecting over many years has made the insect fauna of Santa Catarina
the best known in South America.
Acrosternum {Chinavia) dallasi (Distant, 1900)
(Figs. 203-207)
Rhaphigaster marginatus: Dallas, 1851, p. 277 (misdetermination).
Nezara marginata: StM, 1872, p. 43 (listed).
Nezara dallasi Distant, 1900, p. 392.
Nezara {Acrosternum) dallasi: Kirkaldy, 1909, p. 118.
Acrosternum dallasi: Rolston, 1976, p. 3 (generic placement).
Diagnosis. Yellowish green above with head, anterior disk of pronotum,
base of scutellum and median broad line thereon, connexiva excepting border
entirely or predominantly tan; venter and legs mostly tan or entirely green
except as noted below. Posterior margin of connexival segments narrowly
black bordered. Humeral angles rounded, little produced laterad. Abdominal
spine compressed, subacute apically, reaching between metacoxae. Spiracles
black, unaccompanied by callus. Ostiolar ruga on each side extending about
three-fourths of distance from mesial margin of ostiole to lateral thoracic
margin. Rostrum terminating near posterior limit of metacoxae. Cicatrices
with small black macule at mesial and lateral limits. Basal angle of scutellum
marked with black foveate spot. Dorsal punctation dense, color of punctures
similar to surrounding area. Posterior margin of basal plates from caudal
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Figs. 203-207. A. dallasi. 203. Basal plates, caudal view (BP). 204. Pygophore, ventral view.
205. Pygophore, caudal view. 206. Genital cup. 207. Paramere.
view bent dorsad near lateral angle, fitting into concavity of 9th paratergites
(Fig. 203). Posterior walls of genital cup dorsally diagonal on each side;
dorsal rim concave, roughened, anteromesial corner bearing small tooth
directed anterolaterad (Figs. 205, 206). Little of pygophore exposed from
ventral view (Fig. 204). Length about 10.5-14 mm.
Distribution. Brazil (Goiaz, Minas Gerais), Colombia.
Type. Described from 2 specimens, only one of which was located and is
here designated LECTOTYPE: 9, labeled (a) “Type” (b) Rhaphigaster mar-
ginatus'' (c) “b.”
Acrosternum (Chinavia) bipunctulum (StM, 1872)
(Figs. 208-212)
Nezara {Acrosternum) bipunctula StM, 1872, p. 42.
Diagnosis. Humeri rounded, not produced laterad. Abdominal tubercle
compressed, reaching posterior limit of metacoxae. Connexival segments
narrowly bordered posteriorly with black. Spiracles black, on weak some-
times yellow callus. Ostiolar ruga on each side extending about 0.55 to 0.65
distance from mesial margin of ostiole to lateral margin of thorax. Small
black spot present at mesial limit of cicatrices, another often present at lateral
limit of cicatrices and at basal angles of scutellum. Basal plates weakly
convex, posterior margin slightly concave near mesial angle (Fig. 208). Male
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167
Figs. 208-212. A. bipunctulum. 208. Genital plates, caudoventral view. 209. Pygophore,
ventral view. 210. Pygophore, caudal view. 211. Genital cup. 212. Paramere.
genitalia as in Figures 209-212. Posterior wall of genital cup on each side
diagonal dorsally, dorsal rim concave, weakly denticulate, terminating an-
teriorly in stout curved denticle directed anterolaterad (Figs. 210, 211).
Length about 10-12.5 mm.
Distribution. Brazil (Bahia, Ecuador, Peru (Amazonas).
Types. Lectotype, here designated, 2, labeled (a) “Bahia” (b) “Lindg” (c)
“Type” (d) “Allotypus” (e) “1 1 1/79” (f) “Riksmuseum Stockholm”. Para-
lectotype, here designated, 5 with pygophore missing, labeled (a) “Bahia”
(b) “Type” (c) “Typus” (d) “112/79” (e) “Riksmuseum Stockholm”.
Comment. An occasional specimen of^. laetum has connexival markings
like A. bipunctulum and a dark dot at the mesial limits of the cicatrices as
well. Such specimens may be recognized by the well developed callus around
each spiracle, yellowish macules along the base of the scutellum, and the
posterior margin of the basal plates or outline of the pygophore from a caudal
view.
Acrosternum (Chinavia) impicticorne {Sta\, 1872)
(Figs. 213-218)
Nezara {Acrosternum) impicticornis StM, 1872, p. 42.
Nezara vicina Berg, 1892, pp. 8-9; Piran, 1963, p. 337 (record) (synonymized
by Grazia, 1980).
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Figs. 2 1 3-2 1 8. A. impicticorne. 213. Genital plates, caudoventral view. 214. Spermathecal
bulb and pump. 215. Pygophore, caudal view. 216. Genital cup. 217. Posterior wall, left side,
dorsomesial view. 218. Paramere.
Acrosternum impicticorne: Grazia, 1977, p. 163 (records); Grazia, 1980, p.
234, fig. 1 (lectotype designated).
Diagnosis. Humeri rounded, not produced laterad. Abdominal tubercle
reaching posterior limit of metacoxae, its anterior margin subvertical. Each
connexival segment narrowly bordered in black posteriorly; black spot pres-
ent in posterolateral angles of sternites. Area about each spiracle weakly
calloused, not strongly differentiated by color; peritremes usually pale, rarely
thinly circled with black. Ostiolar ruga on each side extending about three-
fourths distance from mesial margin of ostiole to lateral margin of thorax.
Small black spot usually present at basal angles of scutellum, another often
present at mesial limit of cicatrices and occasionally at lateral limits also.
Posterior margin of basal plates nearly transverse mesad of lateral angles
from caudoventral view (Fig. 213); dorsal edge of basal plates transverse
from caudal view. Spermatheca as in Figure 214. Male genitalia as in Figures
215-218. Posterior wall of genital cup diagonal dorsally with dorsal rim
finely and irregularly denticulate, bearing large denticle anteriorly.
Distribution. Argentina, Paraguay, Brazil, Peru, Ecuador, Surinam, Co-
lombia.
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169
Figs. 219-223. A. herbidum. 219. Spermathecal bulb and pump. 220. Pygophore, ventral
view. 221. Pygophore, caudal view. 222. Genital cup. 223. Paramere.
Types. One syntype of N. impicticornis lacks the dark spot in the basal
angles of the scutellum, and this spot is faint in the other syntypes.
Comments. Dr. J. Grazia first called my attention to the synonymy of TV.
impicticornis and TV. vicina.
In all but one of the 6 1 specimens examined, other than the syntypes of
TV. impicticornis, the scutellar spots are quite clear. The occasional specimen
lacking the scutellar spots will run to A. pengue in the key, but it will not
agree with this species in genitalia or markings at the incisures of the ab-
dominal venter.
The type of TV. vicinia as well as the syntypes of TV. impicticornis were
examined.
Acrosternum {Chinavia) herbidum (Stal, 1859)
(Figs. 219-223)
Rhaphigaster herbidus Stal, 1859, p. 229.
Nezara {Acrosternum) herbida: Stal, 1872, p. 42 (keyed).
Nezara herbida: Berg, 1878, p. 28 (reprinted 1879, p. 58); Berg, 1883, pp.
212-213 (description; reprinted 1884, pp. 28-29); Berg, 1892, p. 6 (de-
scription).
Acrosternum herbida: Rufhnelli and Piran, 1959, p. 1 1.
Acrosternum herbidus: Grazia, 1980, p. 236, fig. 2 (lectotype designated).
Diagnosis. Humeri rounded, not produced laterad. Abdominal tubercle
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reaching between metacoxae to anterior limit of trochanters, rarely just past
metacoxae. Connexival segments bordered posteriorly with black, border
sometimes incomplete but covering lateral half of at least some segments.
Spiracles pale, surrounding area weakly or not calloused and concolorous
with adjacent part of sternites. Ostiolar ruga on each side extending about
0.7 distance from mesial margin of ostiole to lateral margin of thorax. Neither
pronotum nor scutellum marked with black. Dorsal border on outer margins
absent or inconspicuous, narrow, pale yellow. Anterolateral margins of
pronotum weakly convex. Posterior margin of basal plates from caudoventral
view nearly straight excepting lateral angles. Spermatheca as in Figure 219.
Male genitalia as in Figures 220-223. Dorsal rim of posterior wall of genital
cup diagonally oriented from dorsal view, anteriorly bearing large denticle
curving anterolaterad and posteriorly small denticles.
Distribution. Argentina, Brazil, Peru, Surinam.
Comments. The syntypes were examined.
Dr. P. H. Van Doesburg called my attention to an apparently consistent
difference in the pygophore of specimens from the north-south extremes of
this species’ range. In the south, the genital cup rim is infolded near the
posterolateral corners. These projections, which vary in size and shape, have
a finely denticulate and usually infuscated edge. In the north, the genital
cups is unelaborated at these corners (Fig. 222). Whether this difference
indicates variation, subspecies or sibling species cannot be determined from
the material available to me. I have taken the conservative course, treating
the difference as interspecific and geographically undefined variation.
A. herbidurn and A. occulturn, the latter known only from females, seem
indistinguishable excepting the shorter abdominal spine in A. herbidurn.
Acrosternum (Chinavia) pengue, new species
(Figs. 224-232)
Description. Light green above and below with narrow crimson border
along outer margins. Posterior margins of connexival segments narrowly
black bordered, this band continuing onto sternites about halfway to spi-
racular line. An elongated black macule present at base of antennifers on
superior surface. Occasionally dark mark present at mesial limit of cicatrices,
less frequently at lateral limit as well. Basal angles of scutellum immaculate.
Antennae dark green. Legs pale green with rufous tint. Spiracles green to
brown; area around spiracles sometimes paler, not or weakly calloused.
Head 2. 6-2. 9 mm wide across eyes, 2.0-2. 3 mm long. Antennal segments
0. 5-0.6, 0.9-1. 1, 1.2-1. 4, 1.3-1. 8, 1.8-2. 1 mm long. Rostral segments 2
through 4 about 1.4-1. 7, 1. 1-1.2, 0.9-1. 1 mm long; rostrum reaching be-
tween metacoxae.
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171
Figs. 224-232. A. pengue. 224. Genital plates, caudoventral view. 225. Same, caudal view.
226. Spermathecal bulb and pump. 227. Abnormal spermathecal bulb. 228. Proximal end of
dilation of spermathecal duct; sclerotized ring (SR). 229. Pygophore, ventral view. 230. Py-
gophore, caudal view. 231. Genital cup. 232. Paramere.
Pronotum 6. 6-7. 6 mm wide at humeri, 2.6-3. 1 mm long at meson. Hu-
meri rounded, not or barely produced laterad; a short line on anterolateral
edge black. Anterolateral margins of pronotum slightly convex.
Scutellum 4. 2-5.0 mm wide at base, 4. 7-5. 6 mm long.
Abdominal tubercle compressed, reaching between metacoxae. Ostiolar
ruga extending about 0.7 distance from mesial margin of ostiole to lateral
margin of thorax. Posterior margin of basal plates slightly concave mesad
of 9th paratergites from caudoventral view (Fig. 224), strongly concave from
caudal view (Fig. 225). Spermatheca as in Figure 226. A sclerotized ring
present at base of spermathecal dilation (Fig. 228).
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Male genitalia as in Figures 229-232. Posterior wall of genital cup dorsally
diagonal on each side; dorsal rim concave, denticulate with large tooth at
anteromesial end and smaller tooth posterolateral end.
Length about 10-13 mm.
Distribution. Argentina (Misiones) and Brazil (Santa Catarina).
Holotype. S, labeled “Brasilien, Nova Teutonia. 27° 1 LB, 52° 23'L, Fritz
Plaumann”, on one edge “10 1 1942”, on other edge “300-500 M”. De-
posited in American Museum of Natural History.
Paratypes. 8SS, 7$9, same data as holotype except dates “I- 1976” (S USNM),
“VI- 1975” (2SS LHR), “XII- 1975” (S, 9 LHR); “Eldorado, Misiones, Ar-
gentina”, ‘‘4-IX-1967” or “16-IX-1967” (3SS AMNH); “Misiones, Argen-
tina” ((5 RNH); “Argentina, Eldorado, Misiones, X- 15- 1964, A. Kovacs” (9
AMNH); as preceding except date “IX- 16- 1964” (9 AMNH); “Museum
Leiden, Argentina, Dos de Mayo, Prov. Misiones, XII- 1964, F. H. Walz”
(9 RNH); “Museum Leiden, Argentina, Misiones, Loreto, IX- 1955, F. H.
Walz” (399 RNH).
Comment. An anomalous condition consisting of a bifurcate spermathecal
diverticulum was found in one specimen (Fig. 227).
An occasional specimen A. impicticorne which lacks the black spot in the
basal angles of the scutellum will key to A. pengue. However, the genitalia
and markings at the incisures of the abdominal venter distinguish such
specimens from A. pengue.
ACKNOWLEDGMENTS
For the opportunity to study type material in their care I am especially grateful to Messrs.
J. Carayon (Museum National d’Histoire Naturelle), W. R. Dolling (British Museum (Natural
History)), Richard C. Froeschner (National Museum of Natural History, Washington, D.C.),
P. Lindskog and P. I. Perrson (Naturhistoriska Riksmuseet), G. Petersen (Akademie der Land-
wirtschaftsivissenschaften der DDR), and Luis de Santis (Universidad Nacional de La Plata).
I also wish to express my appreciation for the specimens loaned or contributed by Paul H.
Arnaud, Jr. (California Academy of Sciences), John K. Bouseman (Illinois Natural History
Survey), Joan B. Chapin (Louisiana State University), H. D. Engleman, M.D., Jocelia Grazia
(Universidade Estadual de Campinas), Frank W. Mead (Florida State Collection of Arthropods),
Randall T. Schuh and P. Wygodzinsky (American Museum of Natural History), Donald B.
Thomas (University of Missouri) and P. van Doesburg, Jr. (Rijksmuseum van Natuurlijke
Historic).
I am particularly grateful to Joseph E. Eger II, H. Dodge Engleman, Richard C. Froeschner,
and P. H. van Doesburg, Jr. for testing the key, noting errors and presenting questions and
comments that were most helpful in revising the manuscript.
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NEW YORK ENTOMOLOGICAL SOCIETY
91(2), 1983, pp. 177-182
PATHOGENICITY OF THE FUNGUS ENTOMOPHTHOR.4 CULICIS
FOR ADUFT MOSQUITOES: ANOPHELES STEPHENSI
AND CULEX PIPIENS QUINQUEEASCIATUS
John Paul Kramer
Department of Entomology, Cornell University,
Ithaca, New York 14853
Abstract. — Tht pathogenicity of Entomophthora culicis for the mosquitoes Anopheles ste-
phensi and Culex pipiens quinquefasciatus was determined by exposing healthy adults to conidial
showers from cadavers of naturally infected Chironomus decorus and from cultures grown in
vitro. While 100 percent of the A. stephensi succumbed to infection, only 20 percent of the C.
pipiens quinquefasciatus did so. Explanations for these differences are suggested. Some mor-
phological characteristics of the E. culicis used in this study are presented.
Adult Aedes aegypti exposed to conidia of the fungus Entomophthora
culicis originating from field-collected cadavers of the midge Chironomus
decorus or from experimentally infected A. aegypti or from yolk cultures can
develop a fatal mycosis (Kramer, 1982). The present study extends our
understanding of this fungus and its disease-causing abilities in two more
medically important mosquito species; namely. Anopheles stephensi and
Culex pipiens quinquefasciatus. The former is an important vector of ma-
larial parasites from the middle East to India while the latter is a principal
developmental host of the filarial worm Wuchereria bancrofti in Southeast
Asia (Harwood and James, 1979).
MATERIALS AND METHODS
Sources of conidia. E. culicis conidia from two sources were used in this
study. These were field-collected cadavers of naturally infected adult Chi-
ronomus decorus found in Ithaca, New York and mycelial mats produced
from conidia discharged from C. decorus cadavers onto the egg-yolk medium
of Miiller-Kogler (1959). Fresh whole cadavers were placed on 1.5 percent
water agar in small plastic dishes to promote the development of the fungus.
The discharge of conidia commenced within 24 hours. Small chunks of
mycelial mats taken from yolk cultures, 7 to 10 days old, were also placed
on water agar. The discharge of conidia from these chunks usually began
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NEW YORK ENTOMOLOGICAL SOCIETY
VOLUME 91, NUMBER 2
179
within 48 hours. Dishes containing cadavers or chunks of mats were held
at 20 ± 2°C in closed chambers containing moist filter paper.
Testing procedures. Adult mosquitoes 24 to 48 hours old from disease-free
insectary cultures were placed in cylindrical transparent plastic cages (height
35 mm, width 30 mm), having two large mesh-covered windows and mesh-
covered tops and bottoms. A small plastic dish containing water agar with
several cadavers or chunks of mycelial mats was inverted over each cage to
allow the conidia being discharged to fall through the mesh-covered top into
the cage of mosquitoes. These cages with dishes were housed in glass battery
jars containing a layer of wet sand. The jars, tightly closed with polyethylene
wrap, were held in cabinets at 20 ± 2°C with an 18:6 photoperiod. After 48
hours the mosquitoes were transferred to glass-covered carton cages pro-
visioned with bottles of a 3 percent sucrose solution. These cages, held at
20 ± 2°C with an 18:6 photoperiod, were checked daily for dead mosquitoes.
The dead ones were placed on water agar and examined for external growth
of E. culicis at irregular intervals up to 48 hours. Only those cadavers that
produced a bloom of E. culicis on their exteriors by the 48th hour, were
scored as infected. Female and male mosquitoes were present in equal num-
bers in each test group. No attempt was made to determine whether one sex
or the other was more susceptible because of the small numbers of individuals
used in each test.
RESULTS AND DISCUSSION
Representative conidia of the E. culicis used in this study are depicted in
Figure 1. The binucleate primary conidia are broadly ellipsoidal with a
papillate apex and a flattened base. Their size depends upon the substrate
from which they developed. Those discharged from cadavers were about 1 4
to 16 by 10 to 11 fxm, while those produced in vitro were consistently larger,
measuring about 17 to 19 by 12 to 15 pm. In both shape and size these
primary conidia are virtually identical to the E. culicis conidia described by
Gustafsson (1965) and others (see MacLeod et al., 1976). The primary co-
nidium produces a binucleate secondary conidium which is ovoid with a
rounded apex; such conidia range in size from 11 to 14 by 8 to 11 pm.
Secondary conidia produced one to four long germ tubes which are some-
times branched. Tertiary conidia were not observed. Conidia discharged
Figs. I, 2. Fig. I. Conidia of Entomophthora culicis. Note primary conidium surrounded
by protoplasmic layer (at left), two secondary conidia with long germ tubes, and two primary
conidia without protoplasmic layer. Bar equals 1 5 )um. Fig. 2. In vitro culture of Entomophthora
culicis showing its typically convoluted growth pattern. Bar equals 0.75 mm.
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NEW YORK ENTOMOLOGICAL SOCIETY
VOLUME 91, NUMBER 2
181
Table 1. Mortality at post-inoculation day 6 among adult mosquitoes experimentally in-
fected with Entomophthora ciilicis.
Sources of conidia
Species of mosquito
Chironomus
deconis
cadavers
Egg-yolk
cultures
Both sources
Anopheles stephensi
100% (18/18)
100% (33/33)
100% (5 1/51)
Cx. p. quinquefasciatus
11% (4/38)
8% (2/26)
9% (6/64)
onto yolk agar produce glassy, greyish-white cerebriform colonies in 10 days
to two weeks at 20 ± 2°C (Fig. 2). Chunks of these colonies placed on water
agar generally developed a velvety hymenium after 24 to 48 hours.
Test mosquitoes and E. ciilicis. The post-mortem appearance of mos-
quitoes that succumbed to an E. culicis infection is shown in Figures 3 and
4. The fungal outgrowths so prominent on these cadavers occur only when
the dead mosquito rests upon a moist surface. In nature midges with fatal
E. culicis infections seek out shaded moist surfaces on which to die and
thereby contribute to the survival and perpetuation of the agent responsible
for their deaths (Kramer, 1 98 1 ). It is conceivable that this pattern of behavior
would also occur among mosquitoes with E. culicis infections in nature.
The results of the infectivity tests indicate that adults of both mosquito
species are susceptible to infection by E. culicis (Table 1). The relative
susceptibilities of these two hosts are, however, quite different. While 100
percent of the A. stephensi succumbed to infection by the end of post-
inoculation day 6, only about 9 percent of the C. p. quinquesfasciatus had
done so in this time period. By post-inoculation day 10 mortality in the
Culex group reached about 20 percent; many adults in this group lived for
more than 10 days but never developed the mycosis. The observed differ-
ences in susceptibility may be attributed in part to the fact that A. stephensi
is the smaller and more delicate of the two species at risk. A. stephensi may
also be less efficient in dislodging conidia during the grooming process.
Whether conidia originated from a cadaver or from an artificial substrate
did not alter their relative ability to fatally infect host mosquitoes (Table 1).
The time of death of infected Anopheles ranged from 3 to 6 days with 50
percent dying on day 3 (Table 2). For the Culex that became infected the
Figs. 3, 4. Cadavers of Anopheles stephensi displaying post-mortem changes caused by
Entomophthora culicis. 1. Male with typical wreath-like growth of fungus surrounding the
thorax. Bar equals 2.5 mm. 2. Female with atypical crescent-like growth around posterior part
of the thorax. Bar equals 3.0 mm.
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NEW YORK ENTOMOLOGICAL SOCIETY
Table 2. The temporal distribution of deaths among adult mosquitoes dying of the mycosis
caused by Entomophthora culicis.
Number
Post-inoculation days of death*
Species of mosquito
infected
1-2
3
4
5
6
7
8
9
10 11-30
Anopheles stephensi
(50)
0
50
8
18
24
0
0
0
0 0
Cx. p. quinquefasciatus
(24)
0
0
0
25
8
42
0
17
8 0
* Daily distribution in percentages.
time of death ranged from 5 to 10 days with 50 percent mortality occurring
by day 7 (Table 2). If these patterns of mortality took place in nature, they
would enhance the pathogen’s chances for survival by keeping a supply of
fresh conidia in the habitat of potential hosts over a period of several days.
In his review of methods for the mass production of E. culicis and other
pathogens, Nolan (1981) suggests that this fungus may find place in the
applied control of black flies. His conjecture can be expanded to include
mosquitoes as well.
ACKNOWLEDGMENTS
This study was funded in part by the UNDP/World Bank/WHO Special Programme for
Research and Training in Tropical Diseases and USDA/SEA Regional Project S- 1 35. The author
thanks Mr. Donald C. Steinkraus for his preparation of the photographs.
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Harwood, R. F. and M. T. James. 1979. Entomology in Human and Animal Health. Mac-
millan, New York, 548 pp.
Kramer, J. P. 1981. On adult midges (Chironomidae) as victims of the fungal pathogen
Entomophthora culicis (Entomophthoraceae). Aquatic Insects 3: 99-102.
Kramer, J. P. 1982. Entomophthora culicis (Zygomycetes, Entomophthorales) as a pathogen
of adult Aedes aegypti (Diptera, Culicidae). Aquatic Insects 4: 73-79.
MacLeod, D. M., E. Miiller-Kogler and N. Wilding. 1976. Entomophthora species with E.
muscae-Mke conidia. Mycologia 68:1-29.
Miiller-Kogler, E. 1959. Zur Isolierung und Kultur insektenpathogener Entomophthoraceen.
Entomophaga 4:261-274.
Nolan, R. A. 1981. Mass production of pathogens. Pages 3 1 9-324 in: M. Laird (ed.), Blackflies.
Academic Press, London.
Received November 5, 1982; accepted December 23, 1982.
NEW YORK ENTOMOLOGICAL SOCIETY
91(2), 1983, pp. 183-187
THE GENUS PARALINCUS (HEMIPTERA: PENTATOMIDAE)
L. H. Rolston
Department of Entomology, Louisiana State University Agricultural Center,
Baton Rouge, Louisiana 70803
Abstract.— T\iQ genus Paralincus is diagnosed, the type species redescribed and the species
keyed. One new species, Paralincus silvae Rolston, is described and the nominal genus Vauriana
Ruckes relegated to the status of a junior synonym of Paralincus Distant.
The genus Paralincus is among those pentatomid genera recently trans-
ferred from Halyini in Pentatominae to Ochlerini in Discocephalinae (Rol-
ston and McDonald, 1979; Rolston, 1981). Until now this genus has been
monotypic, but during work on the ochlerines new generic synonymy and
a new species of Paralincus were discovered, bringing the known number
of species in this genus to three. These findings are reported here, a diagnosis
of the genus is provided and the type species is redescribed.
The location of the eyes, separated from the anterior pronotal margin by
about one half the eye diameter, together with the femoral armature distin-
guish this genus from other ochlerines.
Paralincus Distant
Paralincus Distant, 1911, pp. 246-247.
Vauriana Ruckes, 1958, pp. 10-12. New Synonymy.
Diagnosis. Femora armed distally on inferior surface with small tubercles,
these obscure or reduced in size and number on posterior femora. Eyes
separated from pronotum by about half the diameter of an eye (Fig. 1).
Interocular distance greater than distance across ocelli from lateral limit of
one to lateral limit of other. Juga exceeding tylus, convergent apically, with-
out anteocular process or subapical tooth. Antennae 5 -segmented, basal
segment slightly surpassing apex of head. Labium arising behind imaginary
plane bisecting head at anterior limit of eyes; second segment reaching me-
socoxae; last segment extending onto sternites 4-6. Anterior margin of
pronotum shallowly concave. Coria surpassing apex of scutellum. Proster-
num concave, meso- and metasterna thinly carinate along meson.
The publication costs of this article were defrayed in part by page charge payment. This article
must therefore be hereby marked “Advertisement” in accordance with 18 U.S.C. §1734 solely
to indicate this fact.
184
NEW YORK ENTOMOLOGICAL SOCIETY
Comment. Females have not been associated definitely with males.
Type species. Paralincus: Ochlerus terminalis Walker, 1867, p. 195, by
original designation. Vauriana: V^auriana bimaculata Ruckus, 1958, pp. 12-
14, by original designation.
KEY TO MALES OE PamlinCUS SPECIES
1 . Parameres capitate; antennal segments 2 and 3 subequal in length, segments 4 and 5
each about 'A longer than 2 or 3 and subequal in length terminalis (Walker)
- Parameres laminate; antennal segments 2, 4 and 5 approximately same length, longer
than 3 2
2. Lateral pygophoral emargination arcuately concave (Fig. 5); head of parameres visible
in emargination suboval, curving laterad apically silvae, new species
- Lateral pygophoral emargination angular (Fig. 6); head of paramere visible in emar-
gination subquadrangular bimaculatus (Ruckes)
Paralincus terminalis (Walker)
Ochlerus terminalis Walker, 1867, p. 195.
Paralincus terminalis: Distant, 1911, p. 247.
Redescription. Head 2.9 mm across eyes, 2.2 mm long; interocular width
1.4 mm; distance across ocelli from lateral limit of one to lateral limit of
other 1.3 mm. Antennal segments 0.8, 1.3, 1.2, 1.7, 1.7 mm in length. Rostral
segments 2-4 about 2.5, 2.2, 1.8 mm. Pronotum 6.9 mm wide at humeri,
3.0 mm long at mesom Scutellum 4.2 mm wide at base, 5.0 mm long. Length
about 13 mm.
Vertex of head elevated, with double row of punctures on each side;
U-shaped row of punctures from lateral margin of elevated vertex passing
around ocellus and terminating at eye. Jugal surface impressed submarginally
before eye where lateral margin concave. Apical third of antennal segment
4 and all of 5 pale. Rostrum brownish yellow.
Small tooth at anterolateral angles of pronotum narrowly rounded; an-
terolateral pronotal margin slightly sigmoid; humeri scarcely produced, weakly
emarginated. Disk rugose, deeply punctured between rugae; collar reflexed
as far laterad as ocelli, bordered where reflexed by dense punctation; cica-
trices not well delineated, somewhat elevated.
Fovea in basal angles of scutellum rather large, with one deep impression
and a few strong punctures. Base of scutellum marked mesially with small
brownish yellow macule. Rugae and punctation of disk similar basally to
that on scutellum, becoming less prominent beyond frena.
Costal angle of coria lying above penultimate abdominal segment; junction
with membrane sinuous; inner angle broadly rounded. Endocoria each bear-
ing large rufous macule; punctation moderately strong, weaking apically.
Membrane dark, with about 12 veins. Connexivum partially exposed, fus-
cous, finely and somewhat sparsely punctate.
VOLUME 91, NUMBER 2
185
Punctation of pleura rather uniform, moderately strong, somewhat sparse.
Coxae, trochanters and tarsi brownish yellow. Tubercles on inferior surface
of front and middle femora in 2 irregular rows, obscure on hind femora.
Abdominal venter with broad shallow median sulcus on all but last seg-
ment, elsewhere with large shallow circular depressions centered on short
seta. Brownish yellow band located submarginally on each side of abdominal
venter interrupted broadly at sutures.
Pygophore deeply, arcuately emarginate from caudal view (Fig. 8). Interior
ridge trisinuate, shallowly concave mesially. Head of parameres capitate
with granulated area apically, visible from lateral view in convex emargi-
nation of extracted pygophore (Fig. 7).
Distribution. Known only from holotype, which came from an unspecified
locality in Brazil.
Paralincus silvae, new species
(Figs. 2-5)
Head 3. 0-3. 2 mm wide across eyes, 2. 5-2. 6 mm long; interocular width
1.6- 1. 7 mm; distance across ocelli from lateral limit of one to lateral limit
of other 1.45-1.5 mm. Antennal segments 0. 9-1.0, 1.6-1. 8, 1.3-1. 4, 1.9,
1.6- 1. 8 mm long. Rostral segments 2-4 about 2. 4-2. 8, 2. 2-2. 3, 2.0-2. 1 mm
long. Pronotum 7. 6-7. 9 mm wide at humeri, 3. 4-3. 6 mm long at meson.
Scutellum 4. 8-5.0 mm wide at base, 5. 4-5. 5 mm long. Length about 13.5-
14.5 mm.
Aside from measurements, conforming to description of P. terminalis
except as follows: A row of transversely elongated punctures on each side
of head bordering elevated vertex which is rugosely and irregularly punctate.
Abdominal venter entirely fuscous, lacking paler submarginal band on each
side. Coxae, trochanters and tarsi dark brown to fuscous. All femora clearly
tuberculate on inferior surface, the number of tubercles fewest on hind fem-
ora.
Lateral emargination of pygophore smoothly arcuate, with tiny acute tooth
on cephalic edge of emargination (Fig. 5). Parameres laminate, curving lat-
erad apically, granular on cephalic margin of lateral face (Figs. 2, 3). Theca
without appendages; conjunctiva moderately sclerotized, bearing small dor-
solateral lobe on each side and medioventral lobe; penisfilum nearly straight,
slightly flanged apically (Fig. 4).
Holotype. 5, labeled “Brazil, Para, Belem-Utinga. 07. IV. 1978. R. Neto.”
Deposited in Rijksmuseum van Natuurlijke Historie.
Paratypes. 2SS, “Surin. [illegible] 10 III 58 ([illegible]” (LHR); (a) “Ter.
Ampa, Rio Ampari, J. Lane, Co.” (b) [illegible] “16. VI. 65.” (c) “Colecao
J. Lane” (Museum de Zoologia de Sao Paulo).
Comments. Aside from substantial differences in the form of the pygophore
and parameres, this species seems indistinguishable from P. bimaculatus. It
186
NEW YORK ENTOMOLOGICAL SOCIETY
Fig. 1. P. tenninalis. Head and pronotum in part.
Figs. 2-5. P. silvae. 2. Right paramere, lateral surface. 3. Same, rotated 180°, mesial surface.
4. Aedeagus; theca (t), penisfilum (p), conjunctiva (c), basal plate in part (b). 5. Lateral emar-
gination of pygophore and head of paramere.
Fig. 6. P bimaculatus. Lateral emargination of pygophore and distal part of paramere.
Figs. 7, 8. P. tenninalis. 7. Lateral emargination of pygophore and head of paramere. 8.
Caudal view of pygophore, parameres and proctiger omitted; inferior ridge (ir).
VOLUME 91, NUMBER 2
187
differs notably from P. terminalis in these respects also, and in the relative
lengths of the antennal segments.
Paralincus bimaculatus (Ruckes), New Combination
(Fig. 6)
Vauriana bimaculata Ruckes, 1958, pp. 12-14, fig. 1.
Ruckes (1958) described this species in detail and provided a habitus
drawing. Other than the male genitalia, P. bimaculatus seems indistinguish-
able from the preceding species, P. silvae. The holotype of Vauriana bi-
maculata was examined and the genitalia figured (Fig. 6).
ACKNOWLEDGMENTS
Drs. W. R. Dolling of the British Museum (Natural History), Jocelia Grazia of the Univer-
sidade Federal do Rio Grande do Sul, Randall T. Schuh (American Museum of Natural History)
and P. H. van Doesburg (Rijksmuseum van Natuurlijke Historic) loaned the specimens for this
study. To them I am grateful.
LITERATURE CITED
Distant, W. L. 1911. XXVIII — Rhynchotal notes— LIII. Neotropical Pentatomidae. Ann. &
Mag. Nat. Hist. (8)7:242-258.
Rolston, L. H. 1981. Ochlerini, a new tribe of Discocephalinae (Hemiptera: Pentatomidae).
J. New York Entomol. Soc. 89(l):40-42.
Rolston, L. H. and F. J. D. McDonald. 1979. Keys and diagnoses for the families of Western
Hemisphere Pentatomoidea, subfamilies of Pentatomidae and tribes of Pentatominae
(Hemiptera). J. New York Entomol. Soc. 87(3): 189-207.
Ruckes, H. 1958. New genera and species of neotropical discocephaline and halyine penta-
tomids (Heteroptera, Pentatomidae). Amer. Mus. Nov. no. 1868.
Walker, F. 1867. Catalogue of the specimens of Hemiptera Heteroptera in the collection of
the British Museum. London, 1:1-240.
Accepted for publication January 14, 1983.
NEW YORK ENTOMOLOGICAL SOCIETY
91(2), 1983, pp. 188-192
BOOK REVIEW
Advances in Cladistics: Proceedings of the First Meeting of the Willi Hennig
Society. — V. A. Funk and D. R. Brooks (eds.). 1981. New York Botanical
Garden, New York. 250 pp. $29.50.
The Proceedings of the First Meeting of the Willi Hennig Society are
appropriately titled Advances in Cladistics. Much of the debate concerning
cladistics over the last few years has comprised repetition of arguments (e.g.,
Mayr, 1981), many of which have been repeatedly refuted. As J. S. Farris
indicates in the Founder’s Address, the Hennig Society will serve as a forum
for advancing phylogenetic systematics as a science, by providing an op-
portunity for diverse workers to debate within a cladistic framework. The
1 3 papers contained herein form an interesting cross-section of current re-
search in theory and application, and show that there are many areas for
further development. The meetings from which these papers were drawn
(which occurred in October of 1980) have been reviewed by Schuh (1981).
The papers are grouped in four sections. In the first, “Cladistics and
Molecular Biology”, Farris provides a landmark paper on the use of distance
data in phylogenetic analysis. He discusses techniques for analyzing im-
munological distances, then examines various other types of biochemical
distances, and concludes with a general treatment of distance measures. He
first devotes considerable space to refutation of the contention of Prager and
Wilson (1978) for general superiority of the Fitch-Margoliash method to the
Distance Wagner procedure of Farris. After showing their comparisons to
be improper he alters his algorithm to fit branch lengths so as to allow better
fit in terms of percent standard deviation (i.e., negative branch lengths). This
leads to discussion of the bases for measuring fit, and from that he proceeds
to consider the effect of nonmetricity on distance measures. As he demon-
strates, this vitiates the interpretation of such measures as amounts of evo-
lutionary change between taxa, to the extent of rendering them useless for
constructing evolutionary trees. Therefore the concept of a molecular clock
as drawn from studies using immunological and Nei’s distances is unsup-
ported (since a measure must be ultrametric to show clocklike behavior).
He then shows that Euclidean metrics are not interpretable as change either,
and that whereas the Manhattan metric is, it can be misleading in terms of
treatment of homoplasious characters. Further, sequence differences and the
Manhattan metric applied to frequency data are shown to lead to uninter-
pretable branch lengths. From these conclusions he is led to recommend
analyzing character data directly rather than via distance techniques.
Farris’ paper has already had a pronounced effect on approaches to genetic
distance data. For example, J. Felsenstein at the Evolution meetings in June
of 1982 suggested changing the interpretation of the distances from actual
VOLUME 91, NUMBER 2
189
phenetic distance to expected (in the statistical sense) distance, and thus
permitting branch length fitting. This is in line with Felsenstein’s inveterate
approach to phylogenetic analysis as a statistical problem, but even if his
interpretation is adopted, any molecular clock still needs repair.
D. L. Swofford’s paper describes a modification of Farris’ Distance Wagner
procedure to incorporate all three addition criteria described by Farris in
1972 simultaneously. He then outlines a method for optimizing Distance
branch lengths after the initial construction of the tree which allows proper
comparison in terms of the “F” statistic of Prager and Wilson (1978). His
procedure is quite different from Farris’ in the foregoing paper, but his results
in re Prager and Wilson are the same. Their logic was flawed and their
comparisons when properly done show general superiority for Distance Wag-
ner. Swofford amplifies on certain points made by Farris, and concludes that
a defined algorithm such as Distance Wagner is generally more efficient than
trial-and-error methods. He rationalizes the use of distance data on the
pragmatic grounds that biochemical systematists will undoubtedly continue
to rely on distance measures, which is no doubt true. However, in view of
the enormous sums expended on such studies they ought to be encouraged
to concoct some better justification. Swofford and Selander (1981) describe
a Fortran program that will perform his modified procedure.
M. F. Mickevich and C. Mitter, in the third and final paper in this section,
compare methods for coding characters of terminal taxa when the characters
vary within taxa. Together with Mickevich’s paper on quantitative bio-
geography, this represents the first published exposition of Mickevich’s
“transformation series analysis” (the term is somewhat sententiously capi-
talized everywhere in the book except the index). The method is a further
formalization for assigning polarities to multistate characters so as to obtain
best fit to a given cladogram, ordering states without requiring a pre-defined
polarity. In the application to coding the authors first outline the three
optimality criteria they use: consistency (degree of fit between classification-
implied and data character distributions), boldness (ability to specify a hi-
erarchy) and “Occam’s Probative” (which might be characterized as con-
straining results to the possible) and then compare three coding methods
using electromorph data. These are 1) “Independent alleles”, where the
alleles are the characters and any one can evolve into any other at the same
locus. They consider treating combinations of these as quantitative char-
acters, or as binary characters. 2) “Shared alleles”, in which the locus is the
eharacter and the allelic combinations its states. When more than two alleles
occur they are ordered such that states are linked to those with which they
share the most alleles. 3) The locus is the character and allelic combinations
the states, with order inferred by transformation series analysis. Using a
variety of data sets, they show a better performance for the third coding
method. They conclude by emphasizing the applicability of this method to
190
NEW YORK ENTOMOLOGICAL SOCIETY
Other sorts of polymorphic characters, and in an intriguing idea, suggest
comparing this method with other types of coding methods developed on
the basis of evolutionary assumptions about particular characters as tests of
those assumptions.
The second section, “Theoretical Cladistics”, is monobasic, comprising
D. R. Brooks’ paper on treating classifications as languages. He suggests
various criteria for best classification from an application of an entropy
concept. This approach is here more successful than in Wiley and Brooks
(1982). The criteria converge on parsimony, and he concludes by analogizing
his approach with those of Nelson (1979) and Farris (1969). His treatment
is more general than that of Farris (1979), in that he considers aspects of
the information within the terminal taxa, whereas Farris treated this as a
constant and analyzed transmission of this information, but it is less appli-
cable in that his criteria could be argued (I suppose) and there is ambiguity
in the relationship of the measures for minimum entropy he compares (e.g.,
his treatment of Nelson’s “term information”).
The next section, “Botanical Cladistics”, includes five papers. Three of
these (Sanders, Bolick and Churchill) are cladistic analyses of various plant
groups, and are presumably included in this volume in order to demonstrate
that, yes, phylogenetic systematics can be done with plants. However, at
least the synopsis provided by Churchill (complete with the Latin diagnoses
of new taxa) seems quite out of place. There are scattered fallacies throughout
some of these papers: Sanders lists four assumptions of Hennigian cladistics,
none of which are necessary; he and Funk frequently use “outgroup” for
“sister-group”; Churchill speaks of methods that employ the axioms of evo-
lution; Sanders terms bushes resolved; Funk states that a cladistic analysis
involving non-monophyletic groups is futile; synapomorphy and autapo-
morphy are frequently confused; etc. These statements, recalling as they do
the misunderstandings prevalent in Systematic Zoology debates of ten years
ago, indicate that botanists are not very advanced in their understanding of
the methods. Although there has of late been considerable print expended
on the issue of whether cladistics has somehow been transformed from
Hennig’s methods, the transformation has been in perception. The methods
are just that, a variety of analytical procedures that may be applied to a
given data set. Whether or not that data set is evolutionarily clean is irrel-
evant. Further, the general tenor to the arguments in favor of applying
cladistic methods, implying that zoological studies are easy by comparison,
is occasionally aggravating.
Withal, certain aspects of these papers are instructive. Funk has an ex-
tended discussion of the treatment of putative hybrids in an analysis, con-
cluding, as does Wagner (1980), that they should be removed prior to the
construction of the cladogram, to which they can be added like so many
ornaments afterwards. This problem is more thoroughly examined here than
VOLUME 91, NUMBER 2
191
in theoretical treatments by zoologists. Bremer and Wanntorp give an in-
teresting comparison of cladistic and traditional classifications of major
groups of organisms, arguing for progress in classification.
The final section, “Biogeography and Cladistics”, contains 4 papers. In
the first. Brooks, T. B. Thorson and M. A. Mayes present a study of freshwater
stingray biogeography, testing four narrative hypotheses, each encompassing
several points, such as the monophyly and origin of the group, on the basis
of assertions about the parasite fauna of these fish. They construct cladograms
for helminth parasites, show Pacific relationships and are able to define areas
of endemism that correspond to those shown by other fishes. They further
suggest that an outgroup genus of stingrays may be paraphyletic because of
the pattern of interrelationships shown by the parasites. However, only two
of the parasite groups force this interpretation (figs. 6 and 1 1). All in all, an
excellent example of the “coevolutionary” approach.
C. J. Humphries also presents an interesting study, on the celebrated
Nothofagus. Constructing a cladogram of the species and then a reduced
area cladogram, he compares this with four geological cladograms and is
able to show only a Laurasia/Gondwanaland pattern. Comparison with re-
duced area cladograms of a variety of other organisms shows considerable
incongruence. Thus Nothofagus is relatively uninformative about interre-
lationships of the southern land masses. Throughout his paper there is a
very good emphasis that acceptance of continental drift only provides a
plausible dispersal route for a non-analytic biogeography.
Mickevich contributes an important paper on quantitative biogeography.
In it she shows how transformation series analysis (which is explained by
examples and figures that allow one to perform the procedure) can be of
great use in biogeography. The method can extract the general pattern (a
“biogeographic map”) from a set of area cladograms. It is a logical extension
of parsimony algorithms to analytic biogeography, which may thus be au-
tomated. Readers attempting to follow her example drawn from Rosen (1978)
should note that there are several typos and there are missing values for row
“D” in Table II: the 4 should be a 2. Also her exclusion of area 7 on the
Heterandria cladogram (fig. 7a) from the transformation into area states may
be an oversight, but points out some ambiguity in this part of the process
(cf her assignments of area states with figs. 1 and 2 in Platnick’s paper).
Platnick has the final say in a paper explicating the thoughts of Nelson
and Platnick (1981) on dealing with widespread taxa in biogeography. He
too uses the example from Rosen (1978), analyzing the cladograms under
two different assumptions about the history of the groups. It is interesting
that Mickevich’s method, when area states are defined as she does and area
7 omitted, gives the same pattern under both assumptions. This again shows
ambiguity in the assignment of area states. If the localities are used as area
states (only areas 2 and 10 overlap), the problem of combinability discussed
192
NEW YORK ENTOMOLOGICAL SOCIETY
by Platnick remains. Whether the greater informativeness under Platnick’s
second assumption (amounting here to disjunction, then fusion of areas) is
more than ad hoc can at least be tested as he suggests.
In summary, as published symposia go, the quality of papers in this book
is quite good. In these days of offset printing of this type of publication, it
is refreshing to see superb quality paper and excellent binding for a paper-
back. However, the rather exorbitant price ($29.50) illustrates why offset
printing is used so widely for symposia. Considering that there are no en-
tomological papers in the book, readers of this journal interested in these
proceedings are likely to be systematists interested in some of the theoretical
papers. That being the case, they would probably be better off photocopying
about three or four papers and the unihed bibliography than purchasing the
whole yohxmQ.— James M. Carpenter, Department of Entomology^, Cornell
University, Ithaca, New York 14853.
LITERATURE CITED
Farris, J. S. 1969. A successive approximations approach to character weighting. Syst. Zool.
18:374-385.
Farris, J. S. 1979. The information content of the phylogenetic system. Syst. Zool. 28:483-
519.
Mayr, E. 1981. Biological classification: toward a synthesis of opposing methodologies. Science
214:510-516.
Nelson, G. 1 979. Cladistic analysis and synthesis: principles and definitions, with a historical
note on Adanson’s Families des Plantes (1763-1764). Syst. Zool. 28:1-21.
Nelson, G. and N. I. Platnick. 1981. Systematics and biogeography: cladistics and vicariance.
Columbia Univ. Press, New York.
Prager, E. M. and A. C. Wilson. 1978. Construction of phylogenetic trees for proteins and
nucleic acids: empirical evaluation of alternative matrix methods. J. Mol. Evol. 1 1:129-
142.
Rosen, D. E. 1978. Vicariant patterns and historical explanation in biogeography. Syst. Zool.
27:159-188.
Schuh, R. T. 1981. Willi Hennig Society: report of first annual meeting. Syst. Zool. 30:76-
81.
Swofford, D. L. and R. B. Selander. 1981. BIOSYS-1: a FORTRAN program for the com-
prehensive analysis of electrophoretic data in population genetics and systematics. J.
Heredity 72:281-28^.
Wagner, W. H. 1 980. Origin and philosophy of the groundplan divergence method of cladistics.
Syst. Bot. 5:173-193.
Wiley, E. O. and D. R. Brooks. 1982. Victims of history— a nonequilibrium approach to
evolution. Syst. Zool. 31:1-24.
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Journal of the
New York Entomological Society
VOLUME 91 JUNE 1983 NO. 2
CONTENTS
A revision of the genus Acrosternum Fieber, subgenus Chinavia Orian, in the western
hemisphere (Hemiptera: Pentatomidae) L. H. Ralston 97-176
Pathogenicity of the fungus Entomophthora culicis for adult mosquitoes: Anopheles
Stephens! and Culex pipiens quinquefaseiatus John Paul Kramer 177-182
The genus Paralincus (Hemiptera; Pentatomidae) L. H. Ralston 183-187
Book Review
Advances in cladistics: proceedings of the first meeting of the Willi Hennig Society
James M. Carpenter 188-192
/■.
SEPTEMBER 1983
No. 3
Vol. 91
Journal^
of the
New York
Entomological Society
(ISSN 0028-7199)
^ fli
Devoted to Entomology in General
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY
Editor: Randall T. Schuh, Department of Entomology, American Museum
of Natural History, Central Park West at 79th Street, New York, New
York 10024
Book Review Editor: Quentin D. Wheeler, Department of Entomology,
Cornell University, Ithaca, New York 14853
Publications Committee: Louis Trombetta, St. Johns University, Chair-
man; Alfred G. Wheeler, Jr., Pennsylvania State Department of Ag-
riculture, Harrisburg; Joseph M. Cerreta, Columbia University College
of Physicians and Surgeons, New York.
The New York Entomological Society
Incorporating The Brooklyn Entomological Society
President: Gerard Iwantsch, Department of Biological Sciences, Fordham
University, Bronx, New York 10458
Vice President: Henry M. Knizeski, Jr., Department of Biology, Mercy
College, Dobbs Ferry, New York 10522
Secretary: Irene E. Matejko, Science Department, The New Lincoln School,
New York, New York 10021
Assistant Secretary: Dennis J. Joslyn, Department of Biology, Rutgers
University, Camden, New Jersey 08102
Treasurer: Louis Sorkin, Department of Entomology, American Museum
of Natural History, New York, New York 10024
Trustees: Class of 1 983— FeXcr Chabora, Queens College, New York; Den-
nis J. Joslyn, Rutgers University, Camden, New Jersey; Class of 1984—
Joseph Cerreta, Columbia University, New York; Durland Fish, Ford-
ham University, Bronx, New York.
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Mailed September 27, 1983
The Journal of the New York Entomological Society (ISSN 0028-7199) is published quarterly (March. June, September,
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Known office of publication: American Museum of Natural History, New York, New York 10024.
Journal of the New York Entomological Society, total copies printed 600, paid circulation 443, mail subscription 443, free
distribution by mail 7, total distribution 450, 150 copies left over each quarter.
NEW YORK ENTOMOLOGICAL SOCIETY
91(3), 1983, pp. 193-222
EXOTIC INSECTS REPORTED NEW TO NORTHEASTERN
UNITED STATES AND EASTERN CANADA SINCE 1970'
E. Richard Hoebeke and A. G. Wheeler, Jr.
Department of Entomology, Cornell University,
Ithaca, New York 14853, and
Bureau of Plant Industry, Pennsylvania Department of Agriculture,
Harrisburg, Pennsylvania 17110
Abstract.— To promote interest in the detection of exotic insects, we provide a list of species
accidentally introduced to North America and reported since 1970 as new to the Western
Hemisphere, United States, or Canada. The area of detection is limited to eastern Canada
(Ontario, Quebec, Newfoundland, and the Maritime Provinces) and the U.S. from Maine to
Virginia and west to Pennsylvania and West Virginia. Included for each immigrant species are
the country or area of probable origin, collection data relating to detection, host and habitat
preferences (if known) in the area of endemism, economic importance in North America, and
subsequent references to distribution or habits in North America. The list includes 70 species
new to the Western Hemisphere; 6, to the United States; and 7, to Canada. The 70 insects new
to the Western Hemisphere include 22 Coleoptera, 14 Hemiptera-Heteroptera, 10 Hymenop-
tera, 7 Lepidoptera, 7 Diptera, 6 Hemiptera-Homoptera, and 1 each for Dictyoptera, Psocoptera,
Thysanoptera, and Neuroptera.
The importance of detecting exotic insects in North America, especially
plant pests, has long been recognized. Since colonial times, man’s activities
and commerce have enhanced the passive, worldwide movement of insects
and other invertebrates. In eastern North America many of the most im-
portant pests are immigrants from Europe or Asia, principally because of
the large volume of Eurasian-North American traffic. Sailer (1978) analyzed
the immigrant fauna (by insect order only) in the contiguous United States.
His comprehensive study further stimulated interest in detection and focused
more attention on several questions such as where have introduced insects
originated, what areas of North America are most susceptible to invasion
by exotic species, and where have they been first detected?
' The original version of this paper was presented as a report of the Committee on Insect
Detection, Evaluation and Prediction (IDEP), Eastern Branch, Entomological Society of Amer-
ica, September 29, 1982, Hartford, Connecticut.
The publication costs of this article were defrayed in part by page charge payment. This article
must therefore be hereby marked “Advertisement” in accordance with 18 U.S.C. §1734 solely
to indicate this fact.
194
NEW YORK ENTOMOLOGICAL SOCIETY
Insect detection generally is the responsibility of the U.S. Department of
Agriculture, state departments of agriculture, Agriculture Canada, and other
cooperating agencies and professionals (see Wheeler and Nixon, 1979). Even
with increased vigilance at the international ports of entry, new discoveries
will be made. The prompt detection of invaders, pest species or otherwise,
continues as one of the vital needs for agriculture in the United States and
Canada. Plans now are underway to develop the first computerized database
of essential information on insects and their allies introduced into the West-
ern Hemisphere, with emphasis on North America, especially the U.S. (L.
Knutson, in litt.). A forthcoming book on exotic pests in North American
agriculture by C. L. Wilson and C. L. Graham (U.S. Department of Agri-
culture) also should draw attention to pest detection.
Although detection is acknowledged as a national priority, some would
consider it one of the less glamorous aspects of entomology, sometimes
eliciting disdain from those who pursue more “sophisticated” activities. But
even in integrated pest management, currently a popular area of research,
detection should play a role. In citing criteria for implementing new pest
management strategies, Kim (1979) emphasized the detection of exotic pests.
Too often, however, it is slighted in favor of assessment and forecasting.
The monitoring of pests obviously is crucial to management strategies, but
an overemphasis on “key pests” without considering other members of the
agroecosystem is not conducive to an appropriate level of detection. We
recognize that the disparate objectives of assessment and detection may be
incompatible in some crop monitoring systems, or probably cannot be em-
ployed each time a crop is sampled. Ideally, however, scouts who evaluate
only the most economically important species should also be familiar with
the “typical” or characteristic arthropods associated with the host crop. With
such knowledge, plus some curiosity, a scout is apt to spot some anachro-
nism—to suspect that a particular species is out of place in the crop being
surveyed. Eventually, a few of these apparently unusual insect inhabitants,
that is, ones appearing unfamiliar to a well-trained observer, will prove
interesting— maybe only an undescribed species of interest to taxonomic
specialists of the group, but perhaps a native insect that recently has changed
its host preference, a resurgent pest, or an immigrant that has gone unde-
tected.
From our general collecting of insects in the Northeast since the mid-
1970’s, we have discovered several species new to the Western Hemisphere,
or immigrants known only from a few North American localities. Such
collecting, plus detections made during the “High Hazard Pest Survey”
conducted by USDA-APHIS, has emphasized our inadequate knowledge of
North American insects, or what Turnbull (1979) has called an “. . . igno-
rance of the recent changes in the insect fauna.” As Sailer (1978) noted, “. . .
VOLUME 91, NUMBER 3
195
while we are painfully aware of those species that are pests, and know
something of those that are beneficial, few people, aside from taxonomists,
are aware of the large number having little or no known importance.”
To promote interest in insect detection — among systematists as well as
those involved in pest management and other applied aspects of entomol-
ogy—we have compiled and listed relevant information on insect detection
in the northeastern U.S. and eastern Canada since 1970. Any attempt to
document all known introductions is at best preliminary because of the
scattered literature in which new records are published, thus making omis-
sions inevitable.
In the following list, we include all insect species considered accidentally
introduced into North America (thus species thought to be naturally Hol-
arctic or deliberately introduced are excluded) and reported in the literature
since 1970 as new Western Hemisphere [W. Hem.], United States [U.S.], or
Canadian [Can.] records from states composing the Eastern Branch, Ento-
mological Society of America, or from eastern Canada.^ For each exotic
species we give (1) the country or area of probable origin, (2) information
on first detection including locality, dates, and other collection data, (3) host
preferences or habitat if known in the area of endemism (most often Europe),
(4) economic importance, and (5) subsequent references to distribution or
life history in North America. The arrangement is by insect orders and
families, with taxa listed alphabetically under the family-group names.
DICTYOPTERA
ECTOBIIDAE
Ectobius sylvestris (Poda) [W. Hem.]
Probable origin. Europe.
Detection. Geneva, New York (Ontario County), June 1980, and between
May 25 and July 16, 1981; in a private home (in kitchen) and out-of-doors.
Host preference(s)/habitat. Various habitats; commonly on the ground in
forests, particularly in mountainous areas.
Economic status. Probably noneconomic.
Reference(s). Hoebeke, E. R. and D. A. Nickle. 1981. The forest cockroach,
Ectobius sylvestris (Poda), a European species newly discovered in North
America (Dictyoptera: Blattoidea: Ectobiidae). Proc. Entomol. Soc. Wash-
ington 83:592-595.
^ Eastern Branch States: Connecticut, Delaware, Maine, Maryland, Massachusetts, New
Hampshire, New Jersey, New York, Pennsylvania, Rhode Island, Vermont, Virginia, and West
Virginia; eastern Canada: New Brunswick, Newfoundland, Nova Scotia, Ontario, Prince Edward
Island, and Quebec.
196
NEW YORK ENTOMOLOGICAL SOCIETY
PSOCOPTERA
ELIPSOCIDAE
Cuneopalpus cyanops (Rostock) [W. Hem.]
Probable origin. Western Europe.
Detection. Prospect Park, Brooklyn, New York, summer 1950; from co-
nifers.
Host preference(s) /habit at. Various conifer spp.
Economic status. Probably noneconomic.
Reference(s). New, T. R. and A. M. Nadler. 1970. A North American
record of Cuneopalpus cyanops (Rostock) (Psocoptera). Entomologist
103:44.
THYSANOPTERA
THRIPIDAE
Tmetothrips subapterus (Haliday) [W. Hem.]
Probable origin. Europe.
Detection. Rew, Pennsylvania (McKean County), July 27, 1975; on Stel-
laria graminea L. (little starwort).
Host preference(s)/habitat. Various plants including Plantago maritima L.
(European seaside plantain), Stellaria media (L.) Vill. (chickweed), Galium
palustre L. (marsh bedstraw), Juncus sp. (a rush), Lolium perenne L. (pe-
rennial ryegrass). Clematis vitalba L. (travelersjoy), Lonicera caprifolium L.
(sweet honeysuckle), and Robinia pseudoacacia L. (black locust).
Economic status. Probably noneconomic.
Reference(s). Anonymous. 1976. U.S. Dept. Agric., Coop. Econ. Ins. Rep.
1(43):798.
HEMIPTERA-HETEROPTERA
MICROPHYSIDAE
Loricula pselaphiformis Curtis [W. Hem.]
Probable origin. Europe.
Detection. Halifax, Nova Scotia, July 22, 1976; on trunks of European
beech {Eagus sylvatica var. purpurea Ait.), at Point Pleasant Park.
Host preference(s)/habitat. Among lichens on trunks of trees such as Bet-
ula, Crataegus, Eagus, and Juniperus; also on Acer, Buxus, Eraxiniis, Mains,
Quercus, Salix, Sambuciis, Tilia, Ulmus, Abies, and Pinus. This species is
predaceous on small arthropods.
Economic status. Probably noneconomic.
Reference(s). Kelton, L. A. 1980. First record of a European bug, Loricula
pselaphiformis, in the Nearctic region (Heteroptera: Microphysidae). Canad.
Entomol. 1 1 2: 1 085-1 087.
VOLUME 91, NUMBER 3
197
Myrmedobia exilis (Fallen) [W. Hem.]
Probable origin. Europe.
Detection. St. John’s, Newfoundland, July 20, 1980; “collected by sweep-
ing ground cover on the east side of Gibbet Hill just west of the Information
Centre (Signal Hill National Historic Park) . . . .”
Host preference(s)/habitat. Among mosses and lichens on trees such as
Finns, Abies, Larix, Quercns, and Fagus; also among moss and grass roots
on the ground. This species is predaceous on small arthropods.
Economic status. Probably noneconomic.
Reference(s). Kelton, L. A. 1982 (1981). First record of a European bug,
Myrmedobia exilis (Heteroptera: Microphysidae), in the Nearctic region.
Canad. Entomol. 113:1 125-1 127.
MIRIDAE
Camptozygum aequale (Villers) [W. Hem.]
Probable origin. Europe.
Detection. 11 counties in central and western Pennsylvania in 1971 and
1972; from seedlings and trees of Scotch pine {Finns sylvestris L.) and seed-
lings of Austrian pine {P. nigra Arnold); red pine {P. resinosa Ait.); Swiss
mountain pine {P. mugo Turra); and western yellow pine {P. ponderosa
Douglas).
Host preference(s)/habitat. Conifers, mainly Finns spp.
Economic status. Probably noneconomic.
Reference(s). Wheeler, A. G., Jr. and T. J. Henry. 1973. U.S. Dept. Agric.,
Coop. Econ. Ins. Rep. 23(16):228; Wheeler, A. G., Jr. and T. J. Henry. 1973.
Camptozygum aequale (Villers), a pine feeding mirid new to North America
(Hemiptera: Miridae). Proc. Entomol. Soc. Washington 75:240-246; Whee-
ler, A. G., Jr. 1979. A comparison of the plant-bug fauna of the Ithaca, New
York area in 1910-1919 with that in 1978. Iowa St. J. Res. 54:29-35; Henry,
T. J. and A. G. Wheeler, Jr. 1979. Palearctic Miridae in North America:
records of newly discovered and little-known species (Hemiptera: Heter-
optera). Proc. Entomol. Soc. Washington 81:257-268; Wheeler, A. G., Jr.
1980. Plant bugs at Cornell: a changing fauna. Cornell Plantations 36(1):
3-8.
Dicyphus rhododendri Dolling [W. Hem.]
Probable origin. Great Britain.
Detection. Several localities in Pennsylvania (Centre, Dauphin, Luzerne,
Philadelphia, and Schuylkill counties) during June 1977 to July 1975; also
Cleveland, Ohio (Cuyahoga County), June 24, 1 933 (specimens in the USNM
colln.); on flowers of native and cultivated Rhododendron maximum L. and
mountain laurel {Kalmia latifolia L.). Because the closest relatives apparently
198
NEW YORK ENTOMOLOGICAL SOCIETY
are in the New World, Dolling suggested that rhododendri is a Nearctic
endemic which has been introduced to England.
Host preference(s)/habitat . Rhododendron spp.
Economic status. Probably noneconomic.
Rcference(s). Henry, T. J. and A. G. Wheeler, Jr. 1976. Dicyphus rhodo-
dendri Dolling, first records from North America (Hemiptera: Miridae).
Proc. Entomol. Soc. Washington 78:108-109.
Megalocoleus mollicullus (Fallen) [Can.]
Probable origin. Europe.
Detection. “Southeastern Ontario”; from Canada goldenrod {Solidago
canadensis L.).
Host preference(s)/habitat. Composites (Asteraceae), particularly Achillea
L. and Tanacetum L.
Economic status. Probably noneconomic.
Reference(s). Reid, D. G., C. C. Loan and R. Harmsen. 1976. The mirid
(Hemiptera) fauna of Solidago canadensis (Asteraceae) in south-eastern On-
tario. Canad. Entomol. 108:561-567.
Orthotylus nassatus (F.) [W. Hem.]
Probable origin. Europe.
Detection. Allentown, Pennsylvania (Lehigh County), July 5, 1973; and 3
miles south of Hershey, Pennsylvania (Dauphin County), July 4, 1977; on
pear, Pyrus sp., heavily infested with pear psylla {Psylla pyricola Foerster)
at Allentown, and at blacklight south of Hershey.
Host preference(s)/habitat. On Eraxinus, Quercus, Salix, and Tilia.
Reference(s). Henry, T. J. 1977. Orthotylus nassatus, a European plant
bug new to North America (Heteroptera: Miridae). U.S. Dept. Agric., Coop.
Plant Pest Rep. 2(3 1):605-608; Henry, T. J. and A. G. Wheeler, Jr. 1979.
Palearctic Miridae in North America: records of newly discovered and little-
known species (Hemiptera: Heteroptera). Proc. Entomol. Soc. Washington
81:257-268; Kelton, L. A. 1982. New records of European Pilophorus and
Orthotylus in Canada (Heteroptera: Miridae). Canad. Entomol. 114:283-
287.
Orthotylus viridinervis (Kirschbaum) [W. Hem.]
Probable origin. Europe.
Detection. Niagara Falls, Ontario, June 17, 1978; on American elm, Ulmus
americana L., heavily infested with woolly elm aphid, Eriosoma america-
num (Riley).
Host preference(s)/habitat. On Wych or Scotch elm {Ulmus glabra Huds.),
and on Alnus, Corylus, Quercus, and Salix.
Economic status. Probably noneconomic.
Reference(s). Henry, T. J. and A. G. Wheeler, Jr. 1979. Palearctic Miridae
VOLUME 91, NUMBER 3
199
in North America: records of newly discovered and little-known species
(Hemiptera: Heteroptera). Proc. Entomol. Soc. Washington 81:257-268;
Kelton, L. A. 1982. New records of European Pilophorus and Orthotylus in
Canada (Heteroptera: Miridae). Canad. Entomol. 114:283-287.
Pilophorus confusus (Kirschbaum) [W. Hem.]
Probable origin. Europe.
Detection. Kentville, Nova Scotia (Research Station, Agriculture Canada),
July 10-14, 1976; in orchard on plum trees heavily infested with aphids.
Host preference(s) /habit at. Various deciduous trees and shrubs, including
Alnus, Crataegus, Populus, Ribes, Salix, and Tilia\ feeds mainly on aphids.
Economic status. Probably noneconomic.
Reference(s). Kelton, L. A. 1982. New records of European Pilophorus
and Orthotylus in Canada (Heteroptera: Miridae). Canad. Entomol. 114:
283-287.
Pinalitus rubricatus (Fallen) [W. Hem.]
Probable origin. Europe.
Detection. Wayne County, Pennsylvania, June 26, 1974; on white spruce
(Picea glauca (Moench) Voss).
Host preference(s)/habitat. Various conifers, mainly spruces {Picea spp.)
Economic status. Probably noneconomic.
Reference(s). Henry, T. J. and A. G. Wheeler, Jr. 1974. U.S. Dept. Agric.,
Coop. Econ. Ins. Rep. 24(1 1):103; Henry, T. J. and A. G. Wheeler, Jr. 1974.
Sthenarus dissimilis and Orthops rubricatus: conifer-feeding mirids new to
North America (Hemiptera: Miridae). Proc. Entomol. Soc. Washington 76:
217-224; Wheeler, A. G., Jr. 1979. A comparison of the plant-bug fauna of
the Ithaca, New York area in 1910-1919 with that in 1978. Iowa St. J. Res.
54:29-35; Wheeler, A. G., Jr. 1980. Plant bugs at Cornell: a changing fauna.
Cornell Plantations 36(1): 3-8.
Plagiognathus vitellinus (Scholtz) [W. Hem.]
Probable origin. Europe.
Detection. 17 counties in Pennsylvania during 1972-1973; on Douglas fir
{Pseudotsuga menziesii (Mich.) Franco); European larch {Larix decidua Mill.),
Norway spruce {Picea abies (L.) Karst.), Colorado spruce {P. pungens En-
gelm.), and white spruce {P. glauca).
Host preference(s)/habitat. Spruce {Picea spp.) and larch {Larix spp.).
Economic status. Probably noneconomic.
Reference(s). Henry, T. J. and A. G. Wheeler, Jr. 1973. Plagiognathus
vitellinus (Scholtz), a conifer-feeding mirid new to North America (Hemip-
tera: Miridae). Proc. Entomol. Soc. Washington 75:480-485; Henry, T. J.
and A. G. Wheeler, Jr. 1 974. U.S. Dept. Agric., Coop. Econ. Ins. Rep. 24( 1 1 ):
103; Henry, T. J. and A. G. Wheeler, Jr. 1979. Palearctic Miridae in North
200
NEW YORK ENTOMOLOGICAL SOCIETY
America: records of newly discovered and little-known species (Hemiptera:
Heteroptera). Proc. EntomoL Soc. Washington 81:257-268; Wheeler, A. G.,
Jr. 1979. A comparison of the plant-bug fauna of the Ithaca, New York area
in 1910-1919 with that in 1978. Iowa St. J. Res. 54:29-35; Wheeler, A. G.,
Jr. 1980. Plant bugs at Cornell: a changing fauna. Cornell Plantations 36(1):
3-8.
Psallus variabilis (Fallen) [W. Hem.]
Probable origin. Europe.
Detection. Planting Fields Arboretum, Oyster Bay, Long Island (Nassau
County), New York, June 6, 1979.
Host preference(s)/habitat. Mainly reproductive structures of oaks (Quer-
cus spp.), and recorded from Populus, Salix, and Tilia', also predaceous on
small insects.
Economic status. Probably noneconomic.
Reference(s). Hoebeke, E. R. 1980. U.S. Dept. Agric., Coop. Plant Pest
Rep. 5(33):628; Wheeler, A. G., Jr. and E. R. Hoebeke. 1982. Psallus vari-
abilis and P. albipennis, two European plant bugs established in North Amer-
ica, with notes on taxonomic changes (Hemiptera-Heteroptera: Miridae).
Proc. EntomoL Soc. Washington 84:690-703.
Sthenams dissimilis Reuter [W. Hem.]
Probable origin. Europe.
Detection. 5 counties in southeastern Pennsylvania from early May to late
June 1973; on balsam hr {Abies balsamea (L.) Mill.), concolor hr {A. concolor
(Gord.) Ldl.), and Nordmann hr (A. nordmanniana (Stev.) Spach).
Host preference(s)/habitat. Fir {Abies spp.).
Economic status. Probably noneconomic.
References. Henry, T. J. and A. G. Wheeler, Jr. 1974. U.S. Dept. Agric.,
Coop. Econ. Ins. Rep. 24(1 1):103; Henry, T. J. and A. G. Wheeler, Jr. 1974.
Sthenams dissimilis and Orthops rubricatus: conifer-feeding mirids new to
North America (Hemiptera: Miridae). Proc. EntomoL Soc. Washington 76:
217-224; Wheeler, A. G., Jr. 1979. A comparison of the plant-bug fauna of
the Ithaca, New York area in 1910-1919 with that in 1978. Iowa St. J. Res.
54:29-35; Wheeler, A. G., Jr. 1980. Plant bugs at Cornell: a changing fauna.
Cornell Plantations 36(l):3-8.
Sthenams rotermundi (Scholtz) [W. Hem.]
Probable origin. Europe.
Detection. Monroeville, Pennsylvania (Allegheny County), May 24-26,
1977; near Wilkes-Barre (Luzerne County), May 7, 1978; and Niagara Falls,
Ontario, June 17, 1978; from quaking aspen {Populus tremuloides Michx.),
large-tooth aspen {P. grandidentata Michx.) and from seeds of silver poplar
{P. alba L.).
VOLUME 91, NUMBER 3
201
Host preference(s)/habitat . White or silver poplar {Populus alba) and also
P. canescens Smith and Alnus glutinosa (L.) Gaertn.
Economic status. Probably noneconomic.
Reference(s). Henry, T. J. and A. G. Wheeler, Jr. 1979. Palearctic Miridae
in North America: records of newly discovered and little-known species
(Hemiptera: Heteroptera). Proc. Entomol. Soc. Washington 81:257-268.
RHOPALIDAE
Rhopalus {Brachycarenus) tigrinus (Schilling) [W. Hem.]
Probable origin. Europe.
Detection. North Vineland, New Jersey (Cumberland County), August 18,
1977; from wheat (Triticum aestivum L.) on a farm.
Host preference(s)/habitat. Various crucifers (Brassicaceae).
Economic status. Probably noneconomic.
Reference(s). Hoebeke, E. R. 1977. U.S. Dept. Agric., Coop. Plant Pest
Rep. 2(40):802; Hoebeke, E. R. 1978. U.S. Dept. Agric., Coop. Plant Pest
Rep. 3(40-4 1):579; Hoebeke, E. R. and A. G. Wheeler, Jr. 1982. Rhopalus
{Brachycarenus) tigrinus, recently established in North America, with a key
to the genera and species of Rhopalidae in eastern North America (Hemip-
tera: Heteroptera). Proc. Entomol. Soc. Washington 84:213-224.
CYDNIDAE
Aethus nigritus (F.) [W. Hem.]
Probable origin. Europe.
Detection. Townsend, Delaware (New Castle County), June 8, 1977; from
soybeans on a farm.
Host preference(s)/habitat. Sandy areas, associated with roots of weeds and
grasses, particularly Corynephorus canescens L. (Beauv.) (Poaceae).
Economic status. Probably noneconomic.
Reference(s). Hoebeke, E. R. 1978. U.S. Dept. Agric., Coop. Plant Pest
Rep. 3(29):376; Hoebeke, E. R. 1980. U.S. Dept. Agric., Coop. Plant Pest
Rep. 5(36):691.
PENTATOMIDAE
Picromerus bidens (L.) [Can.]
Probable origin. Europe.
Detection. Lennoxville, Quebec, in 1968, and Ascot Corner, Quebec, in
1969.
Host preference(s)/habitat. A well-known predator of coleopterous and
lepidopterous larvae.
Economic status. Potentially beneficial.
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NEW YORK ENTOMOLOGICAL SOCIETY
Reference(s). Kelton, L. A. 1972. Picromerus bidens in Canada (Heter-
optera: Pentatomidae). Canad. Entomol. 104:1743-1744; Larochelle, A. 1979.
Les punaises a bouclier (Hemiptera: Scutelleroidea) du Quebec. Cordulia,
Suppl. 11:1-84; Larochelle, A. 1980. Picromerus bidens L. (Heteroptera:
Pentatomidae) en Amerique du Nord: repartition geographique, habitat et
biologic. Bull. Invent. Ins. Quebec 2:10-18.
HEMIPTERA-HOMOPTERA
CICADELLIDAE
Anoscopus albifrons (L.) [W. Hem.]
Probable origin, Europe.
Detection. Greenwood, Kings Co., Nova Scotia, August 12, 1973; and also
Marion Co., Oregon, July 23, 1974.
Host preference(s)/habitat. Mostly dry localities, with Agrostis tenuis Sibth.,
Anthoxanthum, Luzula, Holcus mollis L., etc.
Economic status. Probably noneconomic.
Reference(s). Hamilton, K. G. A. 1975. A review of the northern hemi-
sphere Aphrodina (Rhynchota: Homoptera: Cicadellidae), with special ref-
erence to the Nearctic fauna. Canad. Entomol. 107:1009-1027.
Eupteryx atropunctata (Goeze) [U.S.]
Probable origin. Europe.
Detection. Willimantic, Connecticut (Windham County), July 3, 1979; on
beans {Phaseolus vulgaris L.) in garden.
Host preference(s)/habitat. Especially species of Labiatae (mints), but also
on some Asteraceae, Scrophulariaceae, Verbenaceae, Solanaceae, and Che-
nopodiaceae.
Economic status. Potentially economic.
Reference(s). Hoebeke, E. R. 1980. U.S. Dept. Agric., Coop. Plant Pest
Rep. 5(29): 547; Hoebeke, E. R. and A. G. Wheeler, Jr. In press. Eupteryx
atropunctata: North American distribution, seasonal history and host plants,
and description of hfth-instar nymph (Hemiptera-Homoptera: Cicadellidae).
Proc. Entomol. Soc. Washington.
Planaphrodes bifasciata (L.) [W. Hem.]
Probable origin. Europe.
Detection. St. John’s, Newfoundland, July 26, 1967; in a Malaise trap.
Host preference(s)/habitat. Dry meadows.
Economic status. Probably noneconomic.
Reference(s). Hamilton, K. G. A. 1975. A review of the northern hemi-
sphere Aphrodina (Rhynchota: Homoptera: Cicadellidae), with special ref-
erence to the Nearctic fauna. Canad. Entomol. 107:1009-1027.
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PSYLLIDAE
Psylla mali (Schmidberger) [U.S.]
Probable origin. Europe.
Detection. Camden, Maine (Knox County), June 27, 1980; from apple
leaves {Malus sylvestris Mill.) in organically grown orchard.
Host preference(s) /habitat. Malus sylvestris (apple), Prunus sp. (plum), Cy-
donia oblonga Mill, (quince), and Sorbus aucuparia L. (European mountain
ash).
Economic status. Economic.
Reference(s). Anonymous. 1981. Serious European apple pest new in the
U.S. U.S. Dept. Agric., Plant Pest News 1(2):2.
APHIDIDAE
Bmchycolus aspamgi (Mordvilko) [W. Hem.]
Probable origin. Mediterranean area and eastern Europe.
Detection. Orient, Long Island, New York (Suffolk County), July 20, 1 969,
and New Brunswick, New Jersey (Middlesex County), August and November
20, 1969; on red top {Agrostis alba L.) and asparagus {Asparagus officinalis
L.).
Host preference(s)/habitat. Asparagus {Asparagus sp.).
Economic status. Economic.
Reference(s). Leonard, M. D. 1970. U.S. Dept. Agric., Coop. Econ. Ins.
Rep. 20(11):156.
Coloradoa tanacetina (Walker) [W. Hem.]
Probable origin. Europe.
Detection. Veazie, Maine (Penobscot County), August 13, 1977; from
Tanacetum vulgare L. (common tansy).
Host preference(s)/habitat. Tansy {Tanacetum vulgare).
Economic status. Probably noneconomic.
Reference(s). Smith, C. F. 1978. U.S. Dept. Agric., Coop. Plant Pest Rep.
3(11):89.
Hyadaphis tataricae (Ajzenberg) [W. Hem.]
Probable origin. Northern and western Asia.
Detection. Numerous localities in Quebec since 1976; from honeysuckle
{Lonicera spp.).
Host preference(s)/habitat. Lonicera spp., particularly L. tatarica L. and
L. bella Zabel.
Economic status. Economic.
Reference(s). Boisvert, J.-M., C. Cloutier and J. McNeil. 1981. Hyadaphis
tataricae (Homoptera: Aphididae), a pest of honeysuckle new to North
America. Canad. Entomol. 113:415-418; Voegtlin, D. 1981. Notes on a
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NEW YORK ENTOMOLOGICAL SOCIETY
European aphid (Homoptera: Aphididae) new to North America. Proc. Ento-
mol. Soc. Washington 83:361-362; Voegtlin, D. 1982. The distribution and
spread of Hyadaphis tataricae (Homoptera: Aphididae) in the north-central
states with notes on its hosts, biology, and origin. Great Lakes Entomol. 15:
147-152.
PSEUDOCOCCIDAE
Chaetococcus phmgmitis (Marchal) [W. Hem.]
Probable origin. Europe.
Detection. 1 mile east of Tuckerton, New Jersey (Ocean County), July 21,
1975; on Phragmites communis Trin. (common reed).
Host preference(s)/habitat. Phragmites spp.
Economic status. Probably noneconomic.
Reference(s). Nakahara, S. 1975. U.S. Dept. Agric., Coop. Econ. Ins. Rep.
25(45-48):875.
NEUROPTERA
CONIOPTERYGIDAE
Aleuropteryx juniperi Ohm [W. Hem.]
Probable origin. Europe.
Detection. Harrisburg, Pennsylvania (Dauphin County), July 21, 1968 and
6 other Pennsylvania counties; on scale-infested juniper {Juniperus spp.).
Host preference(s)/habitat. Juniperus spp.; larvae prey on juniper scale
{Carulaspis juniperi (Douche)).
Economic status. Beneficial.
Reference(s). Henry, T. J. 1974. U.S. Dept. Agric., Coop. Econ. Ins. Rep.
24(33):659; Anonymous. 1974. U.S. Dept. Agric., Coop. Econ. Ins. Rep.
24(35):703; Henry, T. J. 1976. Aleuropteryx juniperi: a European scale pred-
ator established in North America (Neuroptera: Coniopterygidae). Proc.
Entomol. Soc. Washington 78:195-201; Wheeler, A. G., Jr. 1981. Updated
distribution of Aleuropteryx juniperi (Neuroptera: Coniopterydiae), a pred-
ator of scale insects on ornamental juniper. Proc. Entomol. Soc. Washington
83:173.
COLEOPTERA
CARABIDAE
Harpalus mfipes DeGeer [U.S.]
Probable origin. Europe.
Detection. Orono, Maine (Penobscot County), May 4 and 29, 1970; on
the banks of the Penobscot River.
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205
Host preference(s)/habitat. Open, moderately dry country, especially cul-
tivated fields, waste places, feeding largely upon seeds; a pest of strawberries.
Economic status. Potentially economic.
Reference(s). Larochelle, A. 1976. Premiere mention de VHarpalus nifipes
DeGeer (Coleoptera) pour les Etats-Unis d’Amerique. Cordulia 2:121-122;
Dunn, G. A. 1981. Distribution of Haraplus nifipes DeGeer in Canada and
United States (Coleoptera; Carabidae). Entomol. News 92:186-188; Larson,
D. J. and D. W. Langor. 1982. Carabid beetles of insular Newfoundland
(Coleoptera: Carabidae: Cicindelidae) — 30 years after Lindroth. Canad.
Entomol. 1 14:591-597.
Leistus feirmgineus (L.) [W. Hem.]
Probable origin. Europe.
Detection. Northwest St. John’s, Newfoundland, August-October 1977;
under stones and in litter or humus lying over well-drained, moist, gravelly
soil in mixed Abies-Alnus-Prunus forest.
Host preference(s)/habitat. Open, dry country, often in moss and grass
tufts.
Economic status. Probably noneconomic.
Reference(s). Larson, D. J. 1978. Leistus ferrugineus (L.) (Coleoptera: Ca-
rabidae), new to North America. Coleopt. Bull. 32:307-309; Larson, D. J.
and D. W. Langor. 1982. Carabid beetles of insular Newfoundland (Co-
leoptera: Carabidae: Cicindelidae) — 30 years after Lindroth. Canad. Ento-
mol. 1 14:591-597.
STAPHYLINIDAE
Sepedophilus marshami (Stephens) [W. Hem.]
Probable origin. Europe.
Detection. Cap Rouge, Quebec, June 19, 1959 and subsequently from
Sydney, Nova Scotia, and additional localities in Quebec along the St. Law-
rence and Ottawa river valleys from Quebec City to Hull.
Host preference(s)/habitat. Under logs and loose bark, in rotting wood, in
leaf litter, under stones, at plant roots, etc.
Economic status. Noneconomic.
Reference(s). Campbell, J. M. 1976. A revision of the genus Sepedophilus
Gistel (Coleoptera: Staphylinidae) of America north of Mexico. Mem. Ento-
mol. Soc. Canad., no. 99:89 pp.
Tachinus corticinus Gravenhorst [W. Hem.]
Probable origin. Europe.
Detection. Near Montreal, Quebec, November 21, 1970 and September
2, 1972; and St. Cyrville, Quebec (Drummond County), April 8, 1967.
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NEW YORK ENTOMOLOGICAL SOCIETY
Host preference(s)/habitat. Under rotting matter, including compost, hay
and straw; under moss and decaying leaves.
Economic status. Probably noneconomic.
Reference(s). Campbell, J. M. 1975. New species and records of Tachinus
(Coleoptera: Staphylinidae) from North America. Canad. Entomol. 107:87-
94.
PSELAPHIDAE
Euplectus (Diplectellus) karsteni (Reichenbach) [W. Hem.]
Probable origin. Europe.
Detection. Specimens examined from Montreal, Quebec, June 29, 1898;
Tilbury, Essex Co., Ontario, March 26, June 2 and September 5, 1967; Long
Island, New York; Washington, D.C., September 2, 1964; Latrobe, St. Vin-
cent, Westmoreland Co., Pennsylvania; and from numerous localities in
Ohio, Michigan, Illinois, Iowa, Wisconsin, and Minnesota.
Host preference(s)/habitat. Frequently found in composted or well-rotted
haystacks, corncob piles, horse manure, and occasionally in tree holes.
Economic status. Probably noneconomic.
Reference(s). Wagner, J. A. 1975. Review of the genera Euplectus, Pyc-
noplectus, Leptoplectus, and Acolonia (Coleoptera: Pselaphidae) including
Nearctic species north of Mexico. Entomol. Amer. 49:125-207.
Euplectus (Diplectellus) signatus (Reichenbach) [W. Hem.]
Probable origin. Europe.
Detection. Specimens examined from Laurentide Park, Camp Le Relais,
Quebec, August 29, 1956, from wood shavings and straw compost near
domestic rabbit hatch; Barre, Washington Co., Vermont, July 7, 1961, from
cow manure mixed with sawdust; Tilbury, Essex Co., Ontario, June 2, 1967,
from manure; and Illinois and Wisconsin.
Host preference(s)/habitat. Restricted to rich domestic or farm compost,
especially that with much fungal mycelia.
Economic status. Probably noneconomic.
Reference(s). Wagner, J. A. 1975. Review of the genera Euplectus, Pyc-
noplectus, Leptoplectus, and Acolonia (Coleoptera: Pselaphidae) including
Nearctic species north of Mexico. Entomol. Amer. 49:125-207.
ELATERIDAE
Melanotus dichrous (Erichson) [W. Hem.]
Probable origin. Southern Europe.
Detection. Adjacent to Pier No. 1, Clinton Street, Canton pier area of
Baltimore City, Maryland, late June 1965 to mid-July 1970, in blacklight
trap.
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207
Host preference(s)/habitat. Nocturnal, attracted to lights. Little or nothing
is known about biology.
Economic status. Probably noneconomic.
Reference(s). Ford, E. J., Jr. 1973. U.S. Dept. Agric., Coop. Econ. Ins.
Rep. 23(29):472; Becker, E. C. 1973. A European species of Melanotus now
established at Baltimore, Maryland (Coleoptera: Elateridae). Proc. Entomol.
Soc. Washington 75:454-458.
DERMESTIDAE
Attegenus lobatus Rosenhauer [W. Hem.]
Probable origin. Europe.
Detection. New York, New York, May 15, 1964; in building. Also found
at Detroit, Michigan, February 2, 1960; in building.
Host preference(s)/habitat. In nests of desert owls {Athene noctua bactriana
Hutt.), Old World jumping rats or gerbils (Gerbillinae), long-clawed ground
squirrels {Spermophilopsis leptodactylus Licht.), and various predatory
mammals such as foxes and badgers; a pest of skins, furs, feathers, woolen
goods, grain, and red pepper.
Economic status. Economic.
Reference(s). Beal, R. S., Jr. 1970. A taxonomic and biological study of
species of Attegenini (Coleoptera: Dermestidae) in the United States and
Canada. Entomol. Amer. 45:141-235.
MELYRIDAE
Troglops cephalotes Erichson [W. Hem.]
Probable origin. Central Europe.
Detection. Bedford, Massachusetts (Middlesex County), May 18, 1974 and
May 20, 1975; at window.
Host preference(s)/habitat. Little or nothing is known about biology.
Economic status. Probably noneconomic.
Reference(s). Wittmer, W. 1975. First records of the genus Troglops Er.
(Coleoptera: Malachiidae) in the United States. Coleopt. Bull. 29:250.
EUCNEMIDAE
Dirrhagofarsus lewisi (Fleutiaux) [W. Hem.]
Probable origin. Japan.
Detection. Leakin Park, Baltimore, Maryland, in 1976; larvae and pupae
collected from fallen, dead beech tree (Eagus grandifolia Ehr.) between 4
March and 1 1 August 1976.
Host preference(s)/habitat. Little or nothing is known about biology and
habits in Japan.
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NEW YORK ENTOMOLOGICAL SOCIETY
Economic status. Probably noneconomic.
Reference(s). Ford, E. J., Jr. and T. J. Spilman. 1979. Biology and immature
stages of Dirrhagofarsus lewisi, a species new to the United States (Coleop-
tera, Eucnemidae). Coleopt. Bull. 33:75-83.
COCCINELLIDAE
Coccinella septempunctata (L.) [W. Hem.]
Probable origin. Europe.
Detection. Near Lyndhurst, New Jersey (Bergen County), June 28, 1973;
in Hackensack Meadowlands. Beginning in 1956, the Benehcial Insects Re-
search Laboratory, USDA-ARS, Newark, Delaware, released this species in
the eastern U.S., but no recoveries were made to indicate its establishment.
Host preference(s)/habitat. Predaceous on numerous species of aphids and
coccids, as well as larvae of some lepidopterous and weevil pests of agri-
cultural crops.
Economic status. Benehcial.
Reference(s). Anonymous. 1974. U.S. Dept. Agric., Coop. Econ. Ins. Rep.
24(33):659; Angalet, G. W. and R. L. Jacques. 1975. The establishment of
Coccinella septempunctata L. in the continental United States. U.S. Dept.
Agric., Coop. Econ. Ins. Rep. 25(45-48):883-884; Larochelle, A. and M.-
C. Lariviere. 1979. Coccinella septempunctata L. (Coleoptera; Coccinellidae)
au Quebec: repartition geographique, habitat et biologic. Bull. Invent. Ins.
Quebec l(4):68-76; Hoebeke, E. R. and A. G. Wheeler, Jr. 1980. New
distribution records of Coccinella septempunctata L. in the eastern United
States (Coleoptera: Coccinellidae). Coleopt. Bull. 34:209-212.
Propylea quatuordecimpunctata (L.) [W. Hem.]
Probable origin. Europe.
Detection. Montreal: He de Montreal; Montmorency No. 2: Saint Jean-
d’Orleans; Portneuf: Saint-Augustin; and Quebec: Lorette, Sainte-Foy.
Host preference(s)/habitat. Predaceous on various aphid species.
Economic status. Benehcial.
Reference(s). Landry, L.-P. 1979. Interessantes recoltes dTnsectes a Trois-
Rivieres. Fabreries 5(1):8-10; Larochelle, A. 1979. Repartition de Coleop-
teres Coccinellidae au Quebec. Fabreries 5(2):26-33; Larochelle, A. 1979.
Les Coleopteres Coccinellidae du Quebec. Cordulia, Suppl. 10:1-111; La-
rochelle, A. and M.-C. Lariviere. 1980. Propylea quatuordecimpunctata L.
(Coleoptera: Coccinellidae) en Amerique du Nord: etablissement, habitat
et biologie. Bull. Invent. Ins. Quebec 2(1): 1-9.
Scymnus {Pullus) suturalis Thunberg [W. Hem.]
Probable origin. Europe.
Detection. 1 1 specimens examined from various localities in Pennsylvania
July 1972; on various species of pine (reported as S. coniferarurn Crotch).
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209
Host preference(s)/habitat. Feeds mainly or entirely on aphids infesting
conifers.
Economic status. Potentially beneficial.
Reference(s). Gordon, R. D. 1976. The Scymnini (Coleoptera: Coccinel-
lidae) of the United States and Canada; key to genera and revision of Scym-
nus, Nephus and Diomus. Bull. Buffalo Soc. Nat. Sci. 28:1-362; Gordon, R.
D. 1982. An Old World species of Scymnus {Pullus) established in Penn-
sylvania and New York (Coleoptera: Coccinellidae). Proc. Entomol. Soc.
Washington 84:250-255.
Subcoccinella vigintiquatuorpunctata (L.) [W. Hem.]
Probable origin. Europe.
Detection. Southcentral Pennsylvania in August 1972 and in the Hack-
ensack Meadowlands (Bergen County), New Jersey, in June 1973; on crown-
vetch (Coronilla varia L.).
Host preference(s)/habitat. Approximately 70 host plants, mainly species
of Caryophyllaceae, Chenopodiaceae, and Leguminosae; a pest of alfalfa
{Medicago sativa L.).
Economic status. Potentially economic.
Reference(s). Anonymous. 1974. European alfalfa beetle in the U.S.—
1974. U.S. Dept. Agric., Coop. Econ. Ins. Rep. 24(36):73 1-734; Anonymous.
1975. Distribution of European alfalfa beetle, Subcoccinella vigintiquatuor-
punctata. U.S. Dept. Agric., Coop. Econ. Ins. Rep. 25(1 1):184; Wheeler, A.
G., Jr. and T. J. Henry. 1981. Seasonal history and habits of the European
alfalfa beetle, Subcoccinella vigintiquatuorpunctata (L.) (Coleoptera: Coc-
cinellidae). Coleopt. Bull. 35:197-203.
LATHRIDIIDAE
Adistemia watsoni (Wollaston) [Can.]
Probable origin. Europe?
Detection. Ottawa, Ontario, February 1974; on the wall of an office build-
ing.
Host preference(s)/habitat. Feeds on spores and hyphae of various species
of fungi (molds) which infest seed and herbarium specimens.
Economic status. Potentially economic.
Reference(s). Becker, E. C. 1977. New and noteworthy records of Coleop-
tera in Canada (1). Ann. Entomol. Soc. Quebec 22:14-17.
CHRYSOMELIDAE
Chaetocnema concinna (Marsham) [W. Hem.]
Probable origin. Europe.
Detection. Hingham, Massachusetts (Plymouth County), June 26, 1979;
from sudangrass forage {Sorghum sudanense (Piper) Stapf) on farm.
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NEW YORK ENTOMOLOGICAL SOCIETY
Host preference(s)/habitat. Known from Beta vulgaris L. (beets and man-
gles), Brassica spp. (kale and rutabaga), Chenopodiurn alba L, (common
lambsquarters), Fragaria sp. (strawberry), etc.
Economic status. Potentially economic.
Reference(s). Hoebeke, E. R. 1980. U.S. Dept. Agric., Coop. Plant Pest
Rep. 5(20):374.
Longitarsus luridus (Scopoli) [U.S.]
Probable origin. Europe.
Detection. Amston, Connecticut (Tolland County), July 20, 1979; on beans
in a garden.
Host preference(s)/habitat. Numerous hosts, including plants of Boragi-
naceae, Lamiaceae, Asteraceae, and Plantaginaceae.
Economic status. Probably noneconomic.
Reference(s). Anonymous. 1981. New U.S. detection in high hazard area.
U.S. Dept. Agric., Plant Pest News l(2):2-3; Anonymous. 1981. Longitarsus
luridus extends range. U.S. Dept. Agric., Plant Pest News 1(3):2.
Psylliodes affinis (Paykull) [Can.]
Probable origin. Europe.
Detection. Between Hamilton and Burlington, Ontario (Halton Co.), June
13, 1981; from foliage of bitter nightshade {Solanurn dulcamara L.).
Host preference(s)/habitat. Solanaceous plants, including bitter nightshade
{Solanurn dulcamara), potato (Solanurn tuberosum L.), tobacco (Nicotiana
spp.), tomato (Lycopersicon esculenturn L.), black henbane (Hyoscyarnus
niger L.), Lyciurn sp., and occasionally hops (Hurnulus sp., Cannabinaceae).
Apparently prefers S. dulcamara.
Economic status. Potentially economic.
Reference(s). Wheeler, A. G., Jr. and E. R. Hoebeke. In press. New records
of a Palearctic flea beetle, Psylliodes affinis, in eastern North America. Proc.
Entomol. Soc. Washington.
Psylliodes napi (F.) [W. Hem.]
Probable origin. Europe.
Detection. Ithaca, New York (Tompkins County) area during 1966-1969;
also in Berkshire County, Massachusetts in early June 1968; on collard
(Brassica oleracea L.) and on yellow rocket (Barbarea vulgaris L.).
Host preference(s)/habitat. Various wild crucifers.
Economic status. Potentially economic.
Reference(s). Tahvanainen, J. O. and R. B. Root. 1970. The invasion and
population outbreak of Psylliodes napi (Coleoptera: Chrysomelidae) on yel-
low rocket (Barbarea vulgaris) in New York. Ann. Entomol. Soc. Amer. 63:
1479-1480.
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211
Psylliodes picina (Marsham) [W. Hem.]
Probable origin. Europe.
Detection. Near Ulster, Pennsylvania (Bradford County), July 6, 1978;
from corn {Zea mays L.) on farm.
Host preference(s)/habitat. Known from Cirsium palustre (L.) Scop, (a
thistle), Lysimachia vulgaris L. (garden loosestrife), Lythrum salicaria L.
(purple loosestrife) and Quercus (oaks).
Economic status. Probably noneconomic.
Reference(s). Hoebeke, E. R. 1979. U.S. Dept. Agric., Coop. Plant Pest
Rep. 4(14):202.
Pyrrhalta viburni (Paykull) [W. Hem.]
Probable origin. Eurasia.
Detection. Ottawa, Ontario and Hull, Quebec, summer 1978; from foliage
of Viburnum opulus L., also on ornamental plantings of V. lantana L. and
V. dentatum L. and on native V. rafinesguianum Schultes.
Host preference(s)/habitat. Viburnum spp.
Economic status. Economic.
Reference(s). Becker, E. C. 1979. Pyrrhalta viburni (Coleoptera: Chryso-
melidae), a Eurasian pest of Viburnum recently established in Canada. Ca-
nad. Entomol. 111:417-419.
CURCULIONIDAE
Calomyctems setarius Roelofs [Can.]
Probable origin. Japan.
Detection. Lanark, Ontario, August 1974; in and around a house, and
“fairly abundant along the roadside.”
Host preference(s)/habitat. Variety of herbaceous plants.
Economic status. Probably noneconomic.
Reference(s). Becker, E. C. 1977. New and noteworthy records of Coleop-
tera in Canada (1). Ann. Entomol. Soc. Quebec 22:14-17.
Larinus carlinae (Olivier) [W. Hem.]
Probable origin. Europe.
Detection. Maugansville, Maryland (Washington County), June 15, 1971;
on thistle {Cirsium sp.).
Host preference(s)/habitat. Composites, including Carduus, Cirsium, Car-
lina, and Centaurea.
Economic status. Potentially beneficial.
Reference(s). White, J. C. 1972. A European weevil, Larinus carlinae
Olivier, collected in Maryland. U.S. Dept. Agric., Coop. Econ. Ins. Rep.
22(26):418. [D. R. Whitehead (USDA, Systematic Entomology Laboratory)
considers L. planus (F.) as the correct name.]
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NEW YORK ENTOMOLOGICAL SOCIETY
Otiorhynchus clavipes BonsdorfF [W. Hem.]
Probable origin. Western Europe.
Detection. “Newport County, Rhode Island, July 16, 1942.”
Host preference(s)/habitat. Adults feed on leaves of many fruit trees and
vines, and larvae eat roots of lilac (Syringa), raspberry (Rubus), strawberry
{Fragaria), smooth sumac {Rhus glabra L.), European cranberry bush {Vi-
burnum opulus L.), bigsting nettle {Urtica dioica L.), curly dock {Rumex
crispiis L.), Japanese aucuba {Aucuba), and honeysuckle {Lonicera). A serious
pest of strawberries.
Economic status. Economic.
Reference(s). Warner, R. E. and E. B. Negley. 1976. The genus Otiorhyn-
chus in America north of Mexico (Coleoptera: Curculionidae). Proc. Ento-
mol. Soc. Washington 78:240-262.
Otiorhynchus ligustici (L.) [Can.]
Probable origin. Europe.
Detection. Wolfe Island (in the St. Lawrence River), near Kingston, On-
tario, “about 1965 and later.”
Host preference(s)/babitat. Numerous plants; a serious pest of alfalfa {Med-
icago sativa).
Economic status. Economic.
Rcfcrencc(s). Warner, R. E. and E. B. Negley. 1976. The genus Otiorhyn-
chus in America north of Mexico (Coleoptera: Curculionidae). Proc. Ento-
mol. Soc. Washington 78:240-262; Becker, E. C. 1977. New and noteworthy
records of Coleoptera in Canada (1). Ann. Entomol. Soc. Quebec 22:14-17.
Polydrusus cervinus (L.) [W. Hem.]
Probable origin. Europe.
Detection. Moorestown?, New Jersey (Burlington County), 1969; Hanover,
New Hampshire (Grafton County), August 1963; and environs of Hanover,
late April to mid-October 1963; from the windshield of a car, and probably
from birch {Betula sp.) that “margined the lacrosse playing helds of Dart-
mouth College.”
Host preference(s)/habitat. Larvae live in the roots of orchard grass, Dac-
tylis glomerata L., and the adults feed on leaves of birch, oak, hazel, and
maple.
Economic status. Probably economic.
Reference(s). Warner, R. E. 1971. Polydrusus cervinus (L.), a European
weevil discovered in North America (Coleoptera: Curculionidae). Proc.
Entomol. Soc. Washington 73:57; Hoebeke, E. R. 1980. U.S. Dept. Agric.,
Coop. Plant Pest Rep. 5(29):540.
Campylirhynchus bruchoides (Herbst) [W. Hem.]
Probable origin. Europe.
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213
Detection. Bear, Delaware (New Castle County), July 25, 1979, and Po-
tomac, Maryland (Montgomery County) in 1979; from alfalfa.
Host preference(s)/habitat. Host plants include Polygonum {P. lapathifo-
lium L., P. persicaria L., and P. hydropiper L.), and species of Oenanthe
and Chaerophyllum (Apiaceae).
Economic status. Probably noneconomic.
Reference(s). Hoebeke, E. R. and D. R. Whitehead. 1980. New records of
Rhinoncus bruchoides (Herbst) for the Western Hemisphere and a revised
key to the North American species of the genus Rhinoncus (Coleoptera:
Curculionidae: Ceutorhynchinae). Proc. Entomol. Soc. Washington 82:556-
561.
SCOLYTIDAE
Xyleborus validus Eichhoff [W. Hem.]
Probable origin. Japan.
Detection. Old Westbury, Long Island, New York (Nassau County), May
1976; from infested grove of mature beech trees and possibly a young Norway
maple in arboretum.
Host preference(s) /habit at. Hosts include Abies, Pinus, Tsuga, Fagus, Pru-
nus, and Quercus.
Economic status. Potentially economic.
Reference(s). Willson, H. R. 1976. U.S. Dept. Agric., Coop. Econ. Ins.
Rep. 1(37):610; Wood, S. L. 1977. Introduced and exported American Sco-
lytidae (Coleoptera). Great Basin Nat. 37:67-74; Wood, S. L. 1982. The
bark and ambrosia beetles of North and Central America (Coleoptera: Scol-
ytidae), a taxonomic monograph. Great Basin Nat. Mem., no. 6:1-1359.
LEPIDOPTERA
OCHSENHEIMERIIDAE
Ochsenheimeria vacculella F. Von Roesslerstamm [W. Hem.]
Probable origin. Eurasia.
Detection. Specimens collected from unspecified locality in New York in
1968, and from near Ithaca, New York (Tompkins Co.) in August 1973;
from Slippery Rock, Pennsylvania (Butler Co.) in August 1971; and also
from unspecified locality in Geauga County, Ohio, in 1964, and in Cuyahoga
County, Ohio, in 1967.
Host preference(s) /habitat. Larval host plants include various cereals and
a number of wild grass species.
Economic status. Potentially economic.
Reference(s). Davis, D. R. 1975. A review of Ochsenheimeriidae and the
introduction of the cereal stem moth Ochsenheimeria vacculella into the
United States (Lepidoptera: Tineoidea). Smith. Contrib. ZooL, no. 192:20
214
NEW YORK ENTOMOLOGICAL SOCIETY
pp.; Ellis, C. R. 1978. First record of the cereal stem moth, Ochsenneimeria
vacculella (Lepidoptera: Ochsenheimeriidae) in Canada. Proc. Entomol. Soc.
Ontario 109:82-83.
OECOPHORIDAE (ETHMIIDAE)
Ethmia bipunctella (F.) [W. Hem.]
Probable origin. Europe.
Detection. Laval and Philipsburg, Quebec, May-August 1965-1969, and
also Pincourt, He Perrot, August 1969.
Host preference(s)/habitat . Larval host plants are Echium vulgare L., and
species of Symphytum, Cynoglossum, Anchusa, and Lithospermum (all Bo-
raginaceae).
Economic status. Probably noneconomic.
Reference(s). Sheppard, A. C. 1970. Palearctic Lepidoptera new to the
province of Quebec. Ann. Entomol. Soc. Quebec 15:14-16; Powell, J. A.
1973. A systematic monograph of New World ethmiid moths (Lepidoptera:
Gelechioidea). Smith. Contrib. ZooL, no. 120:302 pp.; Heppner, J. B. and
J. A. Powell. 1974. Ethmia bipunctella in Maryland, Pennsylvania and West
Virginia: the expanding range of an introduced European moth (Gelechioi-
dea). J. Lepid. Soc. 28:302-305.
TORTRICIDAE
Cnephasia stephensiana (Doubleday) [W. Hem.]
Probable origin. Europe.
Detection. 5 specimens collected in Nova Scotia in 1954 (CNC collection),
from Prince Edward Island and Quebec in 1978, and from the vicinity of
Ottawa, Ontario, in 1979.
Host preference(s)/habitat. Larvae polyphagous on herbaceous plants, in-
cluding Chrysanthemum, Inula, Hieracium, Taraxacum, Centaurea, Son-
chus, Heracleum, Vida, Chenopodium, Rumex, Ranunculus, Plantago\ oc-
casionally on cultivated peas and beans.
Economic status. Potentially economic.
Rcfcrencc(s). Mutuura, A. 1982. Cnephasia stephensiana, a species newly
recorded from Canada and compared with the previously recorded C in-
terjectana (Lepidoptera: Tortricidae). Canad. Entomol. 114:667-671.
Gmpholita delineana (Walker) [W. Hem.]
Probable origin. Eurasia.
Detection. Earliest label dates are New York, New York in 1944, and
Madison, Wisconsin and Lexington, Kentucky in 1943; present distribution
records include Minnesota, Iowa, Missouri, Wisconsin, Illinois, Kentucky,
and New York.
VOLUME 91, NUMBER 3
215
Host preference(s)/habitat. Wild hops (Humulus lupulus L.), as well as
hemp (both Cannabinaceae); larvae feed in stems, inflorescences, and de-
veloping seeds.
Economic status. Economic.
Reference(s). Miller, W. E. 1982. Grapholita delineana (Walker), a Eur-
asian hemp moth, discovered in North America. Ann. Entomol. Soc. Amer.
75:184-186.
GEOMETRIDAE
Chlowclystis rectangulata (L.) [W. Hem.]
Probable origin. Europe.
Detection. The following localities in Nova Scotia, summer 1970: Smiley
Brook Provincial Park, near Brooklyn, Hants Co., July 20; Middle River,
Victoria Co., Cape Breton Island, July 27; Baddeck River, near Baddeck
Bridge, Victoria Co., July 29; and the Hants Co. locality, July 23, 1972.
Host preference(s)/habitat. Primary larval hosts include species of Cra-
taegus, Prunus and Pyrus, and blossoms of wild and cultivated apple.
Economic status. Potentially economic.
Reference(s). Ferguson, D. C. 1972. The occurrence of Chloroclystis rec-
tangulata (L.) in North America (Geometridae). J. Lepid. Soc. 26:220-221;
Neil, K. 1980. The occurrence of Chloroclystis rectangulata (Geometridae)
in New Brunswick. J. Lepid. Soc. 34:75.
Eulithis mellinata (F.) (=Lygris associata (Bkh.)) [W. Hem.]
Probable origin. Europe.
Detection. Laval (He Jesus), Quebec, July 10, 1967, and June 24 and July
1, 1973; captured at light in a garden at rear of a home.
Host preference(s)/habitat. Larval host plants are red and black currant
(Ribes rubrum L. and R. nigrum L.).
Economic status. Probably noneconomic.
Reference(s). Sheppard, A. C. 1975. Lygris associata Bork., a new record
for North America (Lepidoptera: Geometridae). Ann. Entomol. Soc. Quebec
20:7; Neil, K. 1978. A second locality for Eulythis mellinata (Geometridae)
in North America. J. Lepid. Soc. 32:224-225.
NOCTUIDAE
Noctua pronuba L. [W. Hem.]
Probable origin. Europe.
Detection. West end Halifax, Nova Scotia, August 8, 1979; under a house
porch light.
Host preference(s)/habitat. Numerous larval hosts include Poa annua L.,
Rumex, Polygonum, Atriplex, Myosotis, Taraxacum spp., and various Bras-
sicaceae.
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NEW YORK ENTOMOLOGICAL SOCIETY
Economic status. Probably noneconomic.
Reference(s). Neil, K. 1981. The occurrence of Noctua pronuba (L.) (Noc-
tuidae) in Nova Scotia: a new North American record. J. Tepid. Soc. 35:
248.
DIPTERA
CHIRONOMIDAE
Orthocladius {Orthocladius) dentifer Brundin [W. Hem.]
Probable origin. Europe.
Detection. Jock River, near Munster Hamlet, Ontario, May 1973; Lake
Ontario, Sta. X, May 1968; Leeds Co., St. Lawrence Nat’l. Pk., center of
Grenadier IsL, May 1975; known also from other northcentral and western
North American localities.
Host preference(s)/habitat . Pupae and adults in and around lakes.
Economic status. Noneconomic.
Reference(s). Soponis, A. R. 1977. A revision of the Nearctic species of
Orthocladius {Orthocladius) Van der Wulp (Diptera: Chironomidae). Mem.
Entomol. Soc. Canad., no. 102:187 pp.
CECIDOMYIIDAE
Coccopsis marginata de Meijere [W. Hem.]
Probable origin. Europe.
Detection. “New York”; in light trap.
Host preference(s)/habitat. Little or nothing is known about biology.
Economic status. Probably noneconomic.
Reference(s). Gagne, R. J. 1976. New Nearctic records and taxonomic
changes in the Cecidomyiidae (Diptera). Ann. Entomol. Soc. Amer. 69:26-
28.
Colomyia hordei Barnes [W. Hem.]
Probable origin. Europe.
Detection. “New Jersey”; in light trap.
Host preference(s)/habitat . Little or nothing is known about biology.
Economic status. Probably noneconomic.
Reference(s). Gagne, R. J. 1976. New Nearctic records and taxonomic
changes in the Cecidomyiidae (Diptera). Ann. Entomol. Soc. Amer. 69:26-
28. [R. J. Gagne (USDA, Systematic Entomology Laboratory) feels that both
cecidomyiids represent “widespread Holarctic species” rather than intro-
ductions.]
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217
RHAGIONIDAE
Rhagio strigosus Meigen [W. Hem.]
Probable origin. Europe.
Detection. Bergen, New York (Genesee County), around August 8, 1981;
from garage window. Additional dead and damaged specimens collected
from garage window sill at same site in summer 1982.
Host preference(s) /habitat. Little or nothing is known about biology.
Economic status. Noneconomic.
Reference(s). Pechuman, L. L. and E. R. Hoebeke. In press. European
Rhagionidae in eastern North America: records of newly discovered species
(Diptera, Brachycera). Entomol. News.
Rhagio tringarius L. [U.S.]
Probable origin. Europe.
Detection. Lenox (Eastover), Massachusetts (Berkshire County), July 7,
1982 and Jam Pond Bog near German, New York (Chenango County),
August 7, 1980.
Host preference(s) /habitat. Among shrubs on damp ground.
Economic status. Noneconomic.
Reference(s). Pechuman, L. L. and E. R. Hoebeke. In press. European
Rhagionidae in eastern North America: records of newly discovered species
(Diptera, Brachycera). Entomol. News.
SPHAEROCERIDAE
Apteromyia claviventris (Strobl) [W. Hem.]
Probable origin. Europe.
Detection. Ontario: Guelph, 4-XI-1981, ex decaying mushroom; Guelph,
IX- 1979; Dundas, Royal Botanical Gardens, lO-IX-1982, ex decaying coral
fungus; and New Brunswick: St. Andrews, 5-V-1978, ex pan trap in seaweed;
St. Andrews, 1 5-VIII-1978, ex carrion trap.
Host preference(s)/habitat. Adults associated with compost piles, also found
in rabbit, mouse, and mole nests; bred from decomposing narcissus bulbs.
Economic status. Probably noneconomic.
Reference(s). Marshall, S. A. and J. R. Rohacek. 1982. Two new species
and a new Nearctic record in genera Apteromyia and Nearcticorpus (Diptera:
Sphaeroceridae). Ann. Entomol. Soc. Amer. 75:642-648.
AGROMYZIDAE
Agromyza fwntella (Rondani) [Can.]
Probable origin. Europe.
Detection. Near St. Armand, Quebec, less than 3 miles from the Vermont
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NEW YORK ENTOMOLOGICAL SOCIETY
border, June 29, 1972; on alfalfa (larvae reared). First reported in U.S.
(Massachusetts) as Agrornyza sp.; now well established in northeastern states.
Host preference(s)/habitat. Host plants are various species of Medicago
{M. falcata L., M. lupuUna L., M. sativa), Melilotus, and Trifoliimp preferred
host is apparently alfalfa, M. sativa.
Economic status. Economic.
Reference(s). Harcourt, D. G. 1973. Agrornyza frontella (Rond.) (Diptera:
Agromyzidae); a pest of alfalfa new to Canada. Ann. Entomol. Soc. Quebec
18:49-51.
ANTHOMYIIDAE
Delia coarctata (Fallen) [W. Hem.]
Probable origin. Europe.
Detection. First documented occurrence at Cap Chat, Quebec, 21-29 July
1954; additional collections in 1954 and 197 1 from Quebec and New Bruns-
wick, and 1980 from Nova Scotia, Prince Edward Island, and Maine.
Host preference(s)/habitat. Primary wild host is couch grass {Agropyron
repens (L.) Beauv.); also important pest of winter wheat, and will infest
barley and rye.
Economic status. Economic.
Reference(s). McAlpine, J. F. and C. Slight. 1981. The wheat bulb fly,
Delia coarctata, in North America (Diptera: Anthomyiidae). Canad. Ento-
mol. 113:615-621.
Emmesomyia soda (Fallen) [W. Hem.]
Probable origin. Europe.
Detection. South of Auburn, New York (Cayuga County), August 6 and
8, 1970; from cells of Oxybelus uniglurnis quadrinotatus Say (a sphecid
wasp).
Host preference(s)/habitat. Adults attracted to animal dung and other de-
caying animal matter; larvae develop in dung.
Ecorjornic status. Probably noneconomic.
Reference(s). Anonymous. 1971. U.S. Dept. Agric., Coop. Econ. Ins. Rep.
2 1(1): 5. [F. C. Thompson (USDA, Systematic Entomology Laboratory) be-
lieves that this record should be referred to the closely related species E.
villica (Meigen).]
HYMENOPTERA
TENTHREDINIDAE
Allantus viennensis (Schrank) [W. Hem.]
Probable origin. Europe.
Detection. Several localities in Tompkins County, New York (vicinity of
VOLUME 91, NUMBER 3
219
the Cornell University campus and Ludlowville) from June 1967 to August
1968; from flowers of Rosa sp. and Rubus sp.
Host preference(s)/habitat. Larval hosts include foliage of Rosa spp., and
also Rubus and Ribes.
Economic status. Potentially economic.
Reference(s). Smith, D. R. 1975. A rose sawfly new to North America
(Hymenoptera: Tenthredinidae). U.S. Dept. Agric., Coop. Econ. Ins. Rep.
25(10):163-165.
BRACONIDAE
Chorebus rondanii (Giard) [U.S.]
Probable origin. Europe.
Detection. South Deerfield, Massachusetts (Franklin County), May 1973;
reared from puparia of asparagus miner (Ophiomyia simplex (Loew)) (Agro-
myzidae).
Host preference(s) /habit at. Known host is Ophiomyia simplex.
Economic status. Potentially beneficial.
Reference(s). Anonymous. 1973. U.S. Dept. Agric., Coop. Econ. Ins. Rep.
23(43):729.
ICHNEUMONIDAE
Homotherus magus (Wesmael) [W. Hem.]
Probable origin. Europe.
Detection. “Newfoundland”; collections made in July and August 1973,
and June-August 1974.
Host preference(s)/habitat. Little or nothing is known about biology.
Economic status. Probably noneconomic.
Reference(s). Heinrich, G. H. 1975. Synopsis of Nearctic Ichneumoninae
Stenopneusticae with particular reference to the northeastern region (Hy-
menoptera). Supplement 5: Ichneumoninae of the island of Newfoundland.
Natur. Canad. 102:753-782.
EULOPHIDAE
Tetrastichus rhosaces (Walker) [W. Hem.]
Probable origin. Europe.
Detection. Winchester, Virginia (Frederick County), July 6, 1973; from
Cassida rubiginosa F. (a chrysomelid beetle) on Carduus nutans L.
Host preference(s)/habitat. Four species of Cassida have been recorded as
hosts.
Economic status. Potentially beneficial.
Reference(s). Anonymous. 1974. U.S. Dept. Agric., Coop. Econ. Ins. Rep.
24(14):203.
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NEW YORK ENTOMOLOGICAL SOCIETY
PTEROMALIDAE
Trlchomalus statutus (Foerster) [W. Hem.]
Probable origin. Europe.
Detection. Blacksburg, Virginia (Montgomery County), August 28, 1970;
emerged from pupae of the frit fly, Oscinella frit (L.) (Chloropidae).
Host preference(s)/habitat. Known host is Oscinella frit.
Economic status. Beneficial.
Reference(s). Allen, W. A. 1971. U.S. Dept. Agric., Coop. Econ. Ins. Rep.
21(23);400.
EUCOILIDAE
Hexacola hexatoma (Hartig) [W. Hem.]
Probable origin. Europe.
Detection. Blacksburg, Virginia (Montgomery County), September 3, 1970;
emerged from pupae of the frit fly, Oscinella frit.
Host preference(s)/habitat. Known host is Oscinella frit.
Economic status. Beneficial.
Reference(s). Allen, W. A. 1971. U.S. Dept. Agric., Coop. Econ. Ins. Rep.
21(23):400.
VESPIDAE
Polistes gallicus (L.) [W. Hem.]
Probable origin. Europe.
Detection. Cambridge, Somerville, Belmont, and Newton (Boston area),
Massachusetts in 1981; also in Cambridge in 1980.
Host preference(s)/habitat. Nests constructed in enclosed places, such as
metal containers and gutter pipes; or in the open, hanging from eaves, branch-
es, or other protective horizontal structures.
Economic status. Probably noneconomic.
Reference(s). Hathaway, M. A. 1981. Polistes gallicus in Massachusetts
(Hymenoptera: Vespidae). Psyche 88:169-173.
Vespula germanica (F.) [W. Hem.]
Probable origin. Europe.
Detection. Earliest known record is Ithaca, New York (Tompkins County),
August 18, 1891; other records include vicinity of Ithaca from 1920-1950;
and Walkersville, Maryland (Frederick County) in 1968. (Previous reports
of germanica in North America have been discounted as they were probably
based on misidentifications of the North American species V. maculifrons
(de Buysson).)
Host preference(s)/habitat . Nests are usually subterranean, but may be
VOLUME 91, NUMBER 3
221
aerial or in roofs, attics, and between walls of houses; workers prey on wide
variety of arthropods and also are opportunistic in exploiting food sources.
Economic status. Economic.
Reference(s). Menke, A. S. and R. Snelling. 1975. Vespula germanica
(Fabricius), an adventive yellow jacket in the northeastern United States
(Hymenoptera: Vespidae). U.S. Dept. Agric., Coop. Econ. Ins. Rep. 25(1 1):
193-200; Akre, R. D., A. Greene, J. F. MacDonald and others. 1981. The
yellowjackets of America north of Mexico. U.S. Dept. Agric., Sci. and Educ.
Admin., Agriculture Handbook, no. 552:102 pp.
MEGACHILIDAE
Chelostoma campanularum (Kirby) [W. Hem.]
Probable origin. Europe.
Detection. Schoharie, Tompkins, Ontario, Schuyler, and Jefferson coun-
ties, New York, since 1973, from July 1-July 27; from inflorescences of
bellflower. Campanula spp. (Campanulaceae).
Host preference(s)/habitat. Strictly oligolectic for pollen of bellflowers.
Campanula spp.; nests constructed in pre-existing cavities, such as holes in
old wood, including stumps, posts, and structural timbers.
Economic status. Probably noneconomic.
Reference(s). Eickwort, G. C. 1980. Two European species of Chelostoma
established in New York State (Hymenoptera: Megachilidae). Psyche 87:
315-323.
Chelostoma fuliginosum (Panzer) [W. Hem.]
Probable origin. Europe.
Detection. Schoharie, Tompkins, and Ontario counties. New York, since
1962, from June 4-July 2; from inflorescences of bellflower. Campanula
spp.
Host preference(s)/habitat. Strictly oligolectic for pollen of bellflowers.
Campanula spp.; nests constructed in pre-existing cavities such as holes in
old wood, including stumps, poles, and structural timbers.
Economic status. Probably noneconomic.
Reference(s). Eickwort, G. C. 1980. Two European species of Chelostoma
established in New York State (Hymenoptera: Megachilidae). Psyche 87:
315-323.
Litharge chrysurus Fonscolombe [W. Hem.]
Probable origin. Western Mediterranean area.
Detection. Phillipsburg, New Jersey (Warren County), June 1977 (the Phil-
lipsburg nest site is only 150 feet from the railroad which serves industries
along the Delaware River); from nests in house siding, consisting of a 4 mm
outer layer of asphalt-impregnated wood fiber.
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NEW YORK ENTOMOLOGICAL SOCIETY
Host preference(s)/habitat. Gathers pollen of spotted knapweed { Centaurea
maculosa (Lamarck)) (Asteraceae), and also from star thistle (C calcitrapa
L.).
Economic status. Probably noneconomic.
Reference(s). Roberts, R. B. 1978. The nesting biology, behavior and im-
mature stages of Lithurge chrysurus, an adventitious wood-boring bee in
New Jersey (Hymenoptera: Megachilidae). J. Kansas Entomol. Soc. 51:735-
745.
ACKNOWLEDGMENTS
We thank members of the 1982 Insect Detection, Evaluation and Prediction Committee
(IDEP) and Executive Committee, Eastern Branch, Entomological Society of America, for their
interest in pest detection. The philosophy developed herein is solely that of the authors. We
are grateful to the research entomologists with the Systematic Entomology Laboratory, U.S.
Department of Agriculture, for verifying names in the manuscript. We also thank Ms. Susan
Pohl for typing the manuscript.
LITERATURE CITED
Kim, K. C. 1979. Cooperative pest detection and monitoring system: a realistic approach.
Bull. Entomol. Soc. Amer. 25:223-230.
Sailer, R. I. 1978. Our immigrant insect fauna. Bull. Entomol. Soc. Amer. 24:3-1 1.
Turnbull, A. L. 1979. Recent changes to the insect fauna of Canada. Pages 180-194 in: H.
V. Danks (ed.), Canada and Its Insect Fauna. Mem. Entomol. Soc. Canad. no. 108.
Wheeler, A. G., Jr. and H. F. Nixon. 1979. Insect survey and detection in state departments
of agriculture. Pennsylvania Department of Agriculture (Harrisburg). Special Publication,
28 pp.
Wilson, C. L. and C. L. Graham. In press. Exotic Pests and North American Agricultural
Plants. Academic Press, New York, New York.
Received October 13, 1982; accepted December 23, 1982.
NEW YORK ENTOMOLOGICAL SOCIETY
91(3), 1983, pp. 223-234
THE TYPES OF CERCERINE WASPS DESCRIBED BY
NATHAN BANKS (HYMENOPTERA: PHILANTHIDAE)*
George R. Ferguson
Department of Entomology, Oregon State University,
Corvallis, Oregon 97331
Abstract. — A. study of the type material of the 52 species-group taxa of Cerceris and one
species of Eucerceris described by Nathan Banks is reported. Lectotypes of 25 species-group
taxa of Cerceris are designated. Banks designated holotypes in his original descriptions for two
species of Cerceris, and the types of the remaining 25 species-group taxa of Cerceris and one
species of Eucerceris are accepted as holotypes since the original descriptions state or imply
that only one specimen was involved and only one specimen was found.
In a series of nine papers from 1912 to 1947 Nathan Banks described 52
species-group taxa in the genus Cerceris and one species of Eucerceris. In
only two cases did he unambiguously designate a holotype in the original
description. He states the number of specimens in the type series in some
cases but not in others. Most, but not all, specimens comprising syntype
series are conspecific. Type material for 49 of Banks’ taxa is in the Museum
of Comparative Zoology, Cambridge, Massachusetts (MCZ), type material
of three species is in the American Museum of Natural History, New York
(AMNH), and one species is represented by type specimens in both insti-
tutions.
Lectotypes are designated below for 25 taxa. Holotypes of two species
were designated by Banks in the original descriptions. The remaining 26
taxa are represented by holotypes since only one type specimen was found
and the original description either states or implies that only one specimen
was involved.
In the MCZ each holotype and each specimen in a syntype series bears a
red MCZ label with the printed word TYPE and the type number handwritten
below it. Each specimen in a syntype series carries the same type number.
In his 1917 and 1947 papers Banks stated the MCZ type number for each
newly described species, but he did not do so in his other papers.
For the taxa described from 1912 to 1917 there is usually a small, red
‘ Oregon Agricultural Experiment Station Technical Paper No. 6484.
The publication costs of this article were defrayed in part by page charge payment. This article
must therefore be hereby marked “Advertisement” in accordance with 18 U.S.C. §1734 solely
to indicate this fact.
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NEW YORK ENTOMOLOGICAL SOCIETY
label with only the handwritten word “type” on one, and only one, specimen
of a syntype series in addition to the red MCZ type label. For the taxa
described from 1912 to 1919 subscript numbers are usually found written
below the letter “E” of the printed word “TYPE” on the red MCZ label.
One specimen of a syntype series is always without a subscript number. The
subscript numbering starts with “2” and proceeds consecutively according
to the number of specimens in the syntype series. Where the small, red
handwritten type label also occurs, it is always on the specimen without a
subscript number. I assume that the presence of the small, red handwritten
type label and/or the absence of a subscript number signifies Banks’ intention
that the specimen so labelled be regarded as the type. To the extent possible
I have been guided by the above labelling in selecting lectotypes. In some
cases there are no such clues. In two cases I have deviated from the above
guideline because of the seriously damaged condition of the specimens which
I would have otherwise selected.
Except as otherwise noted, when a single specimen was found, and nothing
in the original description indicates that more than one specimen was in-
volved, that specimen is accepted as the holotype. With the exception of
Cerceris huachuca, all holotypes and lectotypes are in conformity with the
original descriptions.
Each holotype and one specimen of each syntype series carries a deter-
mination label in Banks’ handwriting. With one exception (C. semiatra) the
word “Type” is written on the label. These “header” labels usually have
several pinholes, and it cannot be assumed that the specimen on which the
header label now occurs is the one on which it was placed by Banks. The
present placement of these header labels on specimens within syntype series
is not always consistent with the occurrence of the small, red handwritten
type label or the presence or absence of the subscript numbers discussed
above. These header labels have largely been ignored in selection of lecto-
types.
Individual specimens of eight species bear a curious blue “Paratype” label
in a handwriting that does not appear to be that of Banks. There is no
apparent pattern as to the specimen in a syntype series on which this label
appears. Hathaway (1981) states “No one here [MCZ] knows anything about
the blue paratype labels”. These labels have been ignored in lectotype se-
lection.
Synonymy and taxonomic status of the 52 species of Cerceris described
by Banks will be treated in a forthcoming synonymic check-list of North
and Central American Cerceris.
Cerceris abbreviata Banks, 1919:84
Described from “Males from Yakima River, Little Spokane and Umatilla,
Washington. June and July (S. Henshaw)”. Number of specimens and MCZ
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225
type number not stated. Five conspecific syntype males are in the MCZ with
the type number 13794 labelled as above. One specimen labelled “Yakima
R., W. T., Nelson’s, July 4,5, ’82” is without a subscript number on the
MCZ type label and is designated lectotype. The other four specimens have
subscript numbers 2, 3, 4, 5 respectively.
Cerceris alaope 1912:22
Described from both sexes “From Falls Church, Va., 5 June on Ceano-
thus'\ Number of specimens and MCZ type number not stated. The male
is described first in the original description, and the female is given rather
cursory treatment. A male and a female in the MCZ with type number 13784
are labelled as above except that the male carries the date “5 July” and the
female the date “26 June”. The date of “5 June” in the original description
is considered to be a lapsus. The male bears a small, red handwritten type
label in addition to the MCZ type label and is designated lectotype. The
female is not conspecific but is a specimen of Cerceris clypeata Dahlbom
with reduced pale markings on the posterior terga.
Cerceris ampla Banks, 1912:16
Described from both sexes from “Fedor, Lee County, Texas (Birkman)”.
Number of specimens and MCZ type number not stated. A female and a
conspecific male are in the MCZ with type number 13769. The female with
a small, red handwritten type label and a MCZ type label without a subscript
number is designated lectotype. The male has the subscript number 2 on
the MCZ type label.
Cerceris ar elate Banks, 1912:18
Described from the female “From Great Falls, Va., 20 June”. Number of
specimens and type number not stated. One female in the MCZ labelled as
above with type number 13779 is accepted as the holotype.
Cerceris (Apiratrix) [sic] arizonella Banks, 1947:32
Described from “One male from Tempe, Arizona, 1 August (Bequaert).
Type M.C.Z. no. 23538”. One male in the MCZ labelled as above is accepted
as the holotype.
Cerceris arno Banks, 1947:19
Described from two conspecific females “From Colton, Calif. (Pilate),
Eddy Collection, and mountains near Claremont (Baker). Type M.C.Z. no.
23542”. The two females labelled as above are in the MCZ. The female
from Colton carries both a red MCZ Paratype label with the number 23542
and a MCZ Holotype label with the “Holo” handwritten before the printed
word TYPE. The anterior four legs and most of the antennae are missing
from this specimen. The second specimen is labelled “Mts. near Claremont,
Cal. Baker” and bears a MCZ type 23542 label as well as a small, white
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NEW YORK ENTOMOLOGICAL SOCIETY
handwritten "\ferruginiof" label. Although the gaster is glued to the second
label, the specimen from “Mts. near Claremont” is in much better condition
and is designated lectotype.
Cerceris astarte Banks, 1913b:424
Described from the female from “Falls Church, Va., 7 and 8 Sept. 1912”.
Number of specimens and type number not stated. One female is in the
MCZ labelled “Falls Church, Va., 7-IX” with the type number 13788 and
a small, red handwritten type label. Since two collection dates were given
in the original description, it is presumed that more than one specimen was
involved. The above labelled specimen is designated lectotype.
Cerceris athene Banks, 1947:20
Described from the female “From Claremont, Calif. (Baker). Type M.C.Z.
no. 23537”. Number of specimens not stated. One female in the MCZ with
the above labelling is accepted as the holotype.
Cerceris atramontensis Banks, 1913b:425
Described from the female “From Valley of Black Mts., N. Car., 23 July,
1906 (Beutenmuller)”. Number of specimens and type number not stated.
One specimen is in the AMNH with the above labelling and bears a label
''Cerceris atramontensis Bks.— Type” in Banks handwriting. It has a red
“Type AMNH” label with no number plus a label reading “Am. Mus. Nat.
Hist., Dept. Invert. Zool. No. 21 180”, and it is accepted as the holotype.
Cerceris belfragei Banks, 1917:114
Described from both sexes from “Type. — M.C.Z. 10029. Texas. G. W.
Belfrage. Coll. Peabody Academy Science, Salem. Four specimens”. Three
females and one male are in the MCZ with the above type number. One
female labelled “Tex.” lacks a subscript number on the MCZ type label and
is designated lectotype. The male has subscript number 2, and two females
have subscript numbers 3 and 4 respectively. The four specimens are con-
specific. The female with subscript number 3 has the blue paratype label
described in the introductory discussion.
Cerceris calodera Banks, 1947:22
Described from “One [male] from Jacumba, Calif., 12 August 1917. Type
M.C.Z. no. 27622”. One male in the MCZ with the above labelling is ac-
cepted as the holotype.
Cerceris jucunda Carolina Banks, 1912:26
Described from specimens “From Southern Pines”. Number of specimens,
sex and MCZ type number not stated. Reference in the original description
to the mesosternal processes indicates that he had males. Three conspecific
males from “Southern Pines, N.C.” are in the MCZ with the type number
13785. A male with the collection date “VI-4-09”, a small, red handwritten
type label and the MCZ type label lacking a subscript number is designated
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227
lectotype. It also carries the blue paratype label described earlier. The other
two males have collection dates of “VI-4- 10” and “VII- 12-09”, and bear
subscript numbers 2 and 3 respectively on the MCZ type labels.
Cerceris carhzonensis Banks, 1 9 1 5:403
Described from the male from Uvalde, Texas (misspelled Walde in the
original description). Number of specimens and MCZ type number not
stated. One male in the MCZ with type number 13772 labelled “Uvalde,
Texas, June 19-20, 930 ft.” is accepted as the holotype.
Cerceris catawba Banks, 1912:25
Described from both sexes “From Southern Pines, N. Car. June, (Manee)”.
Number of specimens and MCZ type number not stated. A female and a
conspecific male with the above labelling in the MCZ have the type number
13787. The female bears a small, red handwritten type label in addition to
the MCZ label, the latter being without a subscript number. The specimen
is labelled “Southern Pines, N.C., VI-5-09” and is designated lectotype. The
male has a collection date of “VI-4-09”, and the MCZ type label has the
subscript number 2.
Cerceris chryssipe Banks, 1912:18
Described from both sexes from “Falls Church, Va. in July on Cicuta'\
Number of specimens and MCZ type number not stated. One female and
six conspecific males are in the MCZ with type number 13791. The female
has a small, red handwritten type label in addition to the MCZ label, the
latter being without a subscript number. It is labelled “Falls Church, Va.,
30-VII” and is designated lectotype. The males have the subscript numbers
2, 3, 4, 5, 6, and 8 on the MCZ type lables. The specimen with subscript
number 7 has been found in the U.S. National Museum, Washington.
Cerceris clymene Banks, 1912:20
Described from both sexes “From Glencarlyn, Va., 23 June, (Ceanothiis),
and Falls Church, Va., 30 July”. Number of specimens and MCZ type
number not stated. One female and three conspecific males are in the MCZ
with type number 13789. All are from “Falls Church, Va., 30-VH”, there
being no specimen from Glencarlyn, Va. The female bears a small, red
handwritten type label, lacks a subscript number on theMCZ type label, and
is here designated lectotype. The males have the subscript numbers 2, 3,
and 4 respectively.
Cerceris cornpleta Banks, 1919:83
Described from “Male from Claremont, California (Baker)”. MCZ type
number not stated. One male labelled as above in the MCZ with type number
13767 is accepted as the holotype.
Cerceris dakotensis Banks, 1915:402
Described from both sexes “From Fargo, N. Dak., July 7 to Sept. 6 (Stevens
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NEW YORK ENTOMOLOGICAL SOCIETY
coll.)”. Number of specimens and MCZ type number not stated. A female
and two conspecific males are in the MCZ with type number 13770. The
female has a small, red handwritten type label and the MCZ label is without
a subscript number. It is labelled ‘‘Fargo, N.D., Jul. 9, 191 1, O. A. Stevens,
Melilotus alba"" and is designated lectotype. Males have subscript numbers
2 and 3 on the MCZ label and have collection dates of “July 7, 1911”, and
“Aug. 2, 1913”, respectively. The collection date of “Sept. 6” is not rep-
resented in the type material.
Cerceris denticularis Banks, 1917:113
Described from both sexes from “Type — M.C.Z. no. 10,028. Oregon:
Umatilla, June, 1 882; Washington: Lone Tree, Yakima River, 30 June, 1 882.
Samuel JJenshaw. Nine specimens.” Four females and two males, all con-
specific, with the above type number are in the MCZ. A female labelled
“Umatilla, Or., June 24, ’82” is without a subscript number on the MCZ
type label and is designated lectotype. Three females have the subscript
numbers 2, 3, and 6, respectively, and two males have the subscript numbers
7 and 8, respectively. The male with subscript number 7 is labelled “Spokane,
W. T., July 27, ’82”, a locality not mentioned in the original description.
The subscript numbers 4 and 5 are not represented.
Cerceris englehardti Banks, 1947:12
Described from “A male from St. John, Arizona, 27 July, 1931 (G. P.
Englehardt). Type M.C.Z. no. 27638”. One male labelled as above is in the
MCZ, but the MCZ label states “MCZ Paratype 27638”. Since the original
description states “a male” and only one specimen was found, I regard the
red paratype label as a lapsus in labelling and accept the specimen as the
holotype.
Cerceris eiirymele Banks, 1947:1 1
Described from “The female holotype from Davis, Calif., 30 June (Bo-
hart), M.C.Z. no. 23546, and males from El Cajon, Calif, May 1 (Van Duzee),
and Santa Barbara, Calif., 18 July (Cockerell), Riverside, Calif., 4 Oct. (Me-
lander)”. The female holotype as designated above is labelled with a red
MCZ holotype label in addition to the MCZ 23546 type label. Three males
each carry a red MCZ paratype label with the type number 23546.
Cerceris floridensis Banks, 1915:403
Described from the male “From Gulfport, Fla. (Reynolds)”. Number of
specimens and MCZ type number not stated. One male labelled as above
in the MCZ with the type number 13765 is accepted as holotype.
Cerceris guarina Banks, 1913a:237
Described from the female “From Vinita, Indian Terr., 7 June (Wickham),
Colorado Springs, Colo., 16 June (Wickham), and Chimney Gulch, Golden,
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229
Colo. (Oslar). Type American Mus. Nat. Hist. Co-type, Author’s Collection”.
Number of specimens and type number not stated. The above record of type
material does not indicate which specimen was placed in the AMNH. A
female in the AMNH with the labelling “Vinita, Ind. T., June 7, 8, ’99” and
''Cercehs gnarina Banks— Type” in Banks’ handwriting is designated lec-
totype. An identically labelled specimen is in the MCZ except that it bears
a red MCZ type label with number 1 3768 and is without a subscript number.
It also bears a label in Banks’ handwriting ""Cerceris gnarina Bks. — Co-type”.
Another female in the MCZ from “Chimney Gulch, Golden, Colorado”,
has the subscript number 2 on the MCZ type label. The Colorado Springs
locality mentioned in the original description is not represented. The three
specimens examined are conspecific.
Cerceris grandis Banks, 1 9 1 3b:423
Described from the female “From Ft. Yuma, Arizona; Amer. Mus. Nat.
Hist.”. Number of specimens not stated. One female in the AMNH with
the above labelling and with ^"Cerceris grandis Bks. — Type” in Banks’ hand-
writing is accepted as the holotype.
Cerceris halone 1912:24
Described from the female “From Falls Church, Va.” Number of speci-
mens and type number not stated. One female in the MCZ from the above
locality with the type number 13777 and bearing a small, red handwritten
type label is accepted as the holotype.
Cerceris hesperina Banks, 1917:115
Described from “Type— M.C.Z. 10,03 1 . Washington: Yakima, July, 1882;
Ainsworth, July, 1882. Samuel Henshaw. Four specimens”. A female and
three males, all conspecihc, are in the MCZ with the above labelling. The
female from “Ainsworth, W. T., July 20, ’82” without a subscript number
on the MCZ type label is designated lectotype. The males have subscript
numbers 2, 3 and 4 respectively. The male with subscript number 3 is labelled
“Yakima City, W. T.” whereas the others are from “Ainsworth, W. T.”
Scullen (1965) incorrectly stated that the female was from Yakima, Wash-
ington.
Cerceris (Apiratrix) [sic] huachuca Banks, 1947:29
Described from the male “From Patagonia, Arizona, 20 August (Be-
quaert). Type M.C.Z. no. 27636”. Number of specimens not stated. One
specimen labelled as above is in the MCZ. The thorax and gaster, in one
piece, are glued to a point with a head glued separately to the same point.
The head is that of a species different from that of the thorax and gaster.
The thorax and gaster agree with Banks’ original description, but the head
does not. The specimen as represented by the thorax and gaster, but not the
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NEW YORK ENTOMOLOGICAL SOCIETY
head, is accepted as the holotype. I have placed a “note” on the specimen
referring to a memorandum on hie at the MCZ detailing my reasons for
concluding that the parts of the specimen are not conspecihc.
Cerceris illota Banks, 1947:23
Described from an unstated number of males “From Tucson, Arizona,
August (Bequaert), also Patagonia, Arizona, 20 August (Bequaert), and Col-
ton, Southern California, 16 August (Pilate); also two from Palmerlee, Ar-
izona (Biedermann). Type M.C.Z. no. 23541”. Eleven conspecihc males are
in the MCZ with the type number 23541, six from Tucson, one from Pat-
agonia, two from Palmerlee and two from Colton. The specimens from
Patagonia, Palmerlee and Colton have MCZ type 23541 labels with the
specimen from Patagonia having two such identical labels. Three specimens
from Tucson have a MCZ type 2354 1 label, and two specimens from Tucson
have a MCZ paratype 23541 label. The sixth specimen labelled “Tucson,
Ariz. VIII” with a MCZ holotype 23541 label is designated lectotype.
Cerceris interjecta Banks, 1919:84
Described from “Male from Lake Point, Utah, 18 July (Titus)”. MCZ
type number not stated. The male in the MCZ labelled as above with the
type number 13766 is accepted as the holotype.
Cerceris irene Bdinks, 1912:26
Described from the female “From Fedor, Lee County, Texas, 25 June
(Birkman)”. Number of specimens and MCZ type number not stated. One
female in the MCZ with the above labelling and MCZ type number 13781
is accepted as the holotype.
Cerceris isolde 1947:24
Described from the male “From Palmerlee, Arizona, Biedermann coll.
Type M.C.Z. no. 23540”. One male in the MCZ labelled as above is accepted
as the holotype.
Cerceris melanthe Banks, 1947:21
Described from the female “From Apache Co., Santa Catalina Mts., Ariz.,
25 July, 5500 ft., J. Bequaert collector. Type M.C.Z. no. 23539”. One female
in the MCZ labelled as above is accepted as the holotype.
Cerceris nigritula Banks, 1915:402
Described from the male “From Colden, N.Y., July 3 (M. C. Van Duzee)”.
Number of specimens and type number not stated. A male in the MCZ with
the type number 13782 and labelled as above is accepted as the holotype.
Cerceris finitima Banks, 1912:27
Described from the male “From Falls Church, Va., in August”. Number
of specimens and MCZ type number not stated. Two males are in the MCZ
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231
labelled as above with the type number 13786. One male, dated “7 Aug.”
with a small, red handwritten type label and no subscript number on the
MCZ type label is designated lectotype. The second male is dated “27-VIII”
and has the subscript number 2 on the MCZ type label.
Cerceris nitida Banks, 1913b:424
Described from an unstated number of both sexes “From Valley of Black
Mts., N. Car., 12 to 22 July, 1906 (Beutenmuller)”. A female labelled as
above, except the date is “June 24, 1906”, bearing a ""Cerceris nitida Bks.
— Type” label in Banks’ handwriting is in the AMNH and is designated
lectotype. The discrepancy in date is considered to be a lapsus. The male
was not found.
Cerceris orestes 1947:13
Described from an unstated number of both sexes “From Patagonia, Ar-
izona, 1 to 14 August (Bequaert). Type M.C.Z. no. 27637”. One female in
the MCZ with the above labelling and type number is designated lectotype.
There is no subscript number on the MCZ type label. The male was not
found.
Cerceris orphne Banks, 1 947: 1 5
Described from “One male from Jemez Springs, N. Mex., 2 July (Engle-
hardt). Type M.C.Z. no. 23536”. A male in the MCZ with the above labelling
is accepted as the holotype.
Cerceris posticata Banks, 1916:64
Described from the male “From Jemez Mountains, New Mexico, 1 1 July
(Woodgate)”. Number of specimens and MCZ type number not stated. One
male in the MCZ with the above labelling bearing the MCZ type number
13771 is accepted as the holotype.
Cerceris prominens Banks, 1912:19
Described from both sexes “From Falls Church, Va., September and Oc-
tober”. Number of specimens and MCZ type number not stated. Three
females and two males, all conspecific, are in the MCZ with the above
labelling and bear the MCZ type number 13790. One female with a small,
red handwritten type label and the MCZ type label without a subscript
number is designated lectotype. The other two females have subscript num-
bers 2 and 3 respectively, and the males have subscript numbers 4 and 5
respectively.
Cerceris psamat he Bsinks, 1912:21
Described from the female “From Fedor, Lee County, Texas (Birkman)”.
Number of specimens and MCZ type number not stated. One female in the
MCZ with type number 13780 bearing the above labelling is accepted as the
holotype.
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NEW YORK ENTOMOLOGICAL SOCIETY
C erceris salorne 1923:21
Described from the female “From Long Island, and Nyack, N.Y., Welles-
ley, Mass. (Morse); and Jones’ Creek, Lee Co., Va.”. Number of specimens
and MCZ type number not stated. Three conspecific females are in the MCZ
with the type number 14705 without subscript numbers on the type labels.
One specimen is simply labelled “N.Y.”. A second specimen has a label
“Coll. A. P. M., Lot 11” and a second label “Coll. A. P. M., Lot 30”. A
third specimen is labelled “Jones’ Cr., Lee Co., Va.” and is designated
lectotype since this is the only specimen in unambiguous agreement with
the original description.
Cerceris sayi Banks, 1923:21
Described from both sexes “From Steele, N. Dakota, July 13 (Stevens)”.
Number of specimens and type number not stated. One female in the MCZ
labelled as above with type number 14706 is designated lectotype. The male
was not found.
Cerceris semiatra Banks, 1947:25
Described from “One male from Patagonia, Arizona, 20 August (Be-
quaert). Type M.C.Z. no. 27620”. A male in the MCZ with the above
labelling is accepted as the holotype. The header label in Banks’ handwriting
spells the name ''semiateC and lacks the word “type”.
Cerceris (Apiratrix) [sic] seminigra Banks, 1947:33
Described from “One male from Patagonia, Arizona, 20 August (Be-
quaert). Type M.C.Z. no. 27621”. A male in the MCZ labelled as above is
accepted as the holotype.
Cerceris sext aides Banks, 1947:10
Described from “Holotype female from Lone Tree, Yakima River, Wash.,
30 June 1882 (S. Henshaw); allotype and paratypes from Nelson’s, Yakima
River, 4-5 July, and Camp Umatilla, 26 June, both Washington and by Mr.
Henshaw; also from Davis, Calif., 1 July (Bohart), and one ‘California’. Type
M.C.Z. no. 23547.” The female holotype labelled as above is in the MCZ
and bears a MCZ holotype label in addition to the MCZ 23547 label. Four
males bear MCZ 23547 paratype labels, and one of these labelled “Yakima
R., W. T. Nelson’s, July 4,5 ’82” bears a MCZ allotype label in addition to
the paratype 23547 label.
Cerceris snowi Banks, 1919:84
Described from an unstated number of males “From Tucson, Arizona
(Snow) and San Diego Co., Cal. (Van Duzee)”. MCZ type number not stated.
Three conspecihc males are in the MCZ with the type number 13764. One
male from “Tucson, Ariz.” is without a subscript number on the MCZ label,
but the specimen consists only of a pinned thorax with a head glued to the
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233
top label. The head is without antennae and the gaster is missing. A male
from “San Diego Co., Cal.”, has the subscript number 2. Another male
labelled “Tucson, Ariz., F. H. Snow” bears the subscript number 3 on the
MCZ label and is designated lectotype due to the badly damaged condition
of the specimen without a subscript number.
Cerceris stevensi Banks, 1923:22
Described “From Steele, N. Dakota, Aug. 10 (Stevens)”. Sex, number of
specimens and MCZ type number not stated. Reference to the clypeal process
indicates a female. One female labelled as above in the MCZ with the type
number 14707 is accepted as the holotype.
Cerceris stigmosalis Banks, 1916:64
Described from the male “From Fargo, North Dakota, September, on
Solidago (Stevens)”. Number of specimens and MCZ type number not stated.
One male in the MCZ labelled as above with type number 13778 is accepted
as the holotype.
Cerceris thione Banks, 1947:18
Described from an unstated number of males “From Colton, Calif (Pilate),
Eddy Collection, 24 August and 4 September, and Claremont, Calif. (Baker).
One specimen from Colton is only 8 mm. long. Type M.C.Z. no. 23593
[sic]”. The type number “23593” of the original description is a typograph-
ical error as the specimens are labelled “23543” and the MCZ log indicates
“23543”. Four conspecific males are in the MCZ with the above labelling.
One specimen from “Colton, Calif”, with the date “8-24” and a red MCZ
holotype label in addition to the MCZ 23543 label is designated lectotype.
The remaining three males have red MCZ paratype 23543 labels.
Cerceris vanduzeei Banks, 1917:1 14
Described from the female from “Type— M.C.Z. 10,030. Calif: San Diego,
June. E. P. Van Duzee. One specimen [sic]”. Despite the statement “one
specimen” there are two females in the MCZ with identical labelling as
above except that one has a small, red handwritten type label and the MCZ
label is without a subscript number. It is designated lectotype. The second
specimen lacks the small, red handwritten type label and has the subscript
number 2 on the MCZ label. I interpret the statement “one specimen” in
the original description as a lapsus.
Cerceris {Apiratrix) [sic] vierecki Banks, 1947:30
Described from the female “From Tempe, Arizona, 1 August (Bequaert).
Type M.C.Z. no. 23544”. Number of specimens not stated. One female in
the MCZ with the above labelling is accepted as the holotype. In addition
there are two males in the MCZ from Tempe, Arizona, with collection dates
“2-VIII” and “31 -VII” bearing red MCZ 23544 paratype labels. Since the
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NEW YORK ENTOMOLOGICAL SOCIETY
male was not mentioned in the original description, these specimens are
without status. They are conspecific with C. arizoneUa Banks described from
a male from the same locality.
Cerceris zelica Banks, 1912:23
Described from both sexes, the male “From Fedor, Lee County, Texas,
7 June (Birkmann)”, and the female “From Lee County, Texas, 7 July
(Birkmann)”. Number of specimens and MCZ type number not stated. A
female and a male in the MCZ with the above labelling and the MCZ type
number 13773 are not conspecific. The female with a small, red handwritten
type label and the MCZ label without a subscript number is designated
lectotype. The male bears the subscript number 2 on the MCZ label, and
the gaster is glued to the top label. It is a male of C. squamulifera Mickel.
Eucerceris apicata Banks, 1915:404
Described from the male “From Yuma, Arizona”. Number of specimens
and MCZ type number not stated. One male in the MCZ labelled as above
with the type number 13742 is accepted as the holotype.
ACKNOWLEDGMENTS
I am greatly indebted to Mary Hathaway and Karen Jepson of the MCZ, and to Marjorie
Favreau of the AMNH for their cooperation in making the type material available for study.
I am especially grateful to Mary Hathaway for taking the time to search the MCZ collection
for all available type specimens of the Banks taxa and for valuable background information.
I thank J. D. Lattin, Curator of the Systematic Entomology Laboratory, Oregon State Uni-
versity, for making facilities available for this study, and my colleague P. W. Oman for time
spent in discussions regarding various technical and procedural aspects of the work reported
here.
LITERATURE CITED
Banks, N. 1912. Notes on the eastern species of Cerceris. Ann. Entomol. Soc. Am. 5:1 1-30.
Banks, N. 1913a. Two new fossorial Hymenoptera. Bull. Am. Mus. Nat. Hist. 32:237-238.
Banks, N. 1913b. New American Philanthidae. Bull. Am. Mus. Nat. Hist. 32:421-425.
Banks, N. 1915. New fossorial Hymenoptera. Canad. Entomol. 47:400-406.
Banks, N. 1916. Two new species of Cerceris. Entomol. News 27:64-65.
Banks, N. 1917. New fossorial Hymenoptera. Bull. Mus. Comp. Zool. 61:97-1 15.
Banks, N. 1919. New Psammocharidae and Philanthidae. Canad. Entomol. 51:81-85.
Banks, N. 1923. Notes and descriptions of some fossorial Hymenoptera. Canad. Entomol.
55:21-22.
Banks, N. 1947. Synopsis of west coast Cerceris. Psyche 54:1-35.
Hathaway, M. 1981. Personal communication.
Scullen, H. A. 1965. Review of the genus Cerceris in America north of Mexico. Proc. U.S.
Natl. Mus. 1 16:333-548.
Received September 1, 1982; accepted November 3, 1982.
NEW YORK ENTOMOLOGICAL SOCIETY
91(3), 1983, pp. 235-241
TWO NEW SPECIES AND SYNONYMY OF THREE
SPECIES OF NORTH AMERICAN CERCERIS
(HYMENOPTERA; PHILANTHIDAE)*
George R. Ferguson
Systematic Entomology Laboratory, Department of Entomology,
Oregon State University, Corvallis, Oregon 97331
Abstract.— \ study of the relevant types in the genus Cerceris has resulted in the following
synonymy (junior subjective synonyms in parentheses); C acanthophila Cockerell (=C chil-
opsidis Viereck and Cockerell), revised synonymy, C. cockerelH Viereck, revised status (=C.
minax Mickel), new synonymy, C. gandarai Rohwer (=C. micheneri Scullen), new synonymy.
Two species are left without names and are described as C grisselli Ferguson n. sp. (=C. minax,
sensu Bohart and Grissell, nec Mickel) and C. rohweri Ferguson n. sp. (=C gandarai, sensu
Scullen, nec Rohwer).
Five species of North American Cerceris are discussed. A study of the
types representing the six names previously applied to these hve species
shows that the six names apply to only three of the species, leaving two
species without names. These two species are described as new to provide
names for a forthcoming synonymic list of North American Cerceris, and
the synonymy for the other three species is presented.
Cerceris acanthophila Cockerell
Cerceris acanthophilus (sic) Cockerell, 1897:135 (Male lectotype, Deming,
New Mexico; ANSP); Viereck and Cockerell, 1904:138; Cresson, 1928:
48; Banks, 1947:30; Scullen, 1951:1004, 1972:17.
Cerceris chilopsidis Viereck and Cockerell, 1 904: 1 36 (Female holotype, Rin-
con, New Mexico; ANSP); Cresson, 1928:48; Scullen, 1951:1005, 1965:
365; Bohart and Grissell, 1975:42; Bohart and Menke, 1976:578; Krom-
bein, 1979:1731. Revised Synonymy. Synonymy first proposed by Banks
(1947).
Banks (1947) first recognized that C. acanthophila and C. chilopsidis were
the male and female, respectively, of the same species. However, Scullen
‘ Oregon Agricultural Experiment Station Technical Paper No. 6661.
The publication costs of this article were defrayed in part by page charge payment. This article
must therefore be hereby marked “Advertisement” in accordance with 18 U.S.C. §1734 solely
to indicate this fact.
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NEW YORK ENTOMOLOGICAL SOCIETY
( 1 960, 1 965) misidentified C. acanthophila and used the name for the species
discussed below as C. cockerelli. Scullen (1972) reversed himself and rec-
ognized C chilopsidis as the female of C acanthophila. Bohart and Grissell
(1975) characterized and keyed both sexes of this species under the name
of C chilopsidis.
Cerceris cockerelli Viereck, Revised Status
Cerceris cockerelli Viereck, 1902:731 (Male lectotype. La Jolla, California;
ANSP); Cresson, 1928:48; Banks, 1947:3 1; Scullen, 1951:1006, 1965:367
(as unrecognized species).
Cerceris minax Mickel, 1918:339 (Female holotype, Sacramento, California;
NEB); Banks, 1947:34; Scullen, 1951:1008, 1972:18. New Synonymy.
Cerceris acanthophila, Scullen, 1960:75, 1965:355, in part; Krombein, 1967:
414; Bohart and Grissell, 1975:38; Bohart and Menke, 1976:576; Krom-
bein, 1979:1729.
Scullen (1960, 1965) incorrectly applied the name C. acanthophila to this
taxon, and this interpretation was followed by subsequent authors. Later he
stated (Scullen, 1972:18) ""Cerceris chilopsidis Viereck and Cockerell is,
therefore, now considered a synonym of C acanthophila Cockerell and C.
minax Mickel is again accepted as a valid species.” However, he ambiguously
retained C. minax Mickel in his list of synonyms of C acanthophila. Bohart
and Grissell (1975) correctly recognized C. cockerelli as belonging to this
taxon, but they used C. acanthophila as the senior name. The type of C.
minax Mickel is conspecific with C. cockerelli Viereck.
Cerceris grisselli, new species
Cerceris acanthophila, Scullen, 1965:355, in part, figs. 108a, b, c.
Cerceris minax, Bohart and Grissell, 1975:49, map 36, figs. 99, 100, 124,
125; Bohart and Menke, 1976:584; Krombein, 1979:1739.
Bohart and Grissell (1975) first recognized this taxon as a species distinct
from C. cockerelli. Although Scullen (1965) had specimens of both species
before him, he did not discriminate between them. Bohart and Grissell (1975)
figured and keyed the dififerences between the two species. Unfortunately,
they used the name C. minax for this species, but the holotype female of C.
minax Mickel is clearly a specimen of C. cockerelli. The minax of Bohart
and Grissell (1975) is thereby left without a name.
The female pygidium in grisselli is abruptly constricted distally with the
sides subparallel over the apical one-third, whereas in cockerelli the sides
are rather evenly tapered toward the apex. In the male the clypeal brush of
VOLUME 91, NUMBER 3
237
cockerelli is dense, rectangular, wider than long, whereas in grisselli the brush
is rather loose and tapered toward the apex.
Description. Jugal lobe of hind wing about V3 as long as submedian cell;
stigmal and prestigmal veins brown, slightly paler than subcosta; apico-
median fossette present on tergum I; sternum II with basal platform; com-
plete longitudinal carina on inner ventral margin of coxa III; tegulae evenly
convex, not pitted; propodeal enclosure usually weakly ridged; scutal punc-
tures separated by 1 to 3 puncture diameters, at least across middle; pro-
podeal punctures dense, separated by ridges of 0.3 to 0.5 puncture diameter;
punctation of preapical terga as on propodeum. FEMALE: Length 8-10 mm;
clypeal midsection with low, transverse lamellate process separated apically
from clypeal margin by scarcely the length of flagellomere IV, apical margin
of process usually ematginate with distinct apicolateral corners and about
as wide as subantennal sclerite; mesopleural tubercle and precoxal carina
strong; pygidium narrow, about 2.5 times as long as greatest width, con-
stricted basally, abruptly narrowed apically with sides subparallel over apical
0.3; pygidial surface smoother on apical 0.3 than on basal portion. MALE:
Length 7-9 mm; clypeal midsection with numerous to dense micropunctures
between macropunctures, with 3 short, blunt teeth apically, and about as
wide as midlength (width measured between tentorial pits); antennocular
distance about equal to length of flagellomere I; clypeal brush scarcely as
wide as least clypeo-ocular distance, tapered toward apex, hairs not agglu-
tinated or waxed; sterna IV-VI without apicolateral teeth; pygidium sub-
rectangular, slightly bowed outward medially. COLOR: Black with yellow
markings as follows: clypeus except lip; subantennal sclerite partly or entirely;
lateral sections of face from above sockets to clypeus; scape below; basal
half of mandible; spot behind eye; tegulae in part; lateral stripes on pronotal
collar; metanotum; spot on hypoepimeron sometimes present in female;
anteriorly emarginate transverse bands on terga I-V (female) and terga I-VI
(male), often reduced on tergum I and penultimate tergum, less strongly
emarginate in male than in female; one or more sterna with lateral spots or
transverse band; trochanter III in part or entirely; apical 0.3 to 0.5 of femora;
tibiae and metatarsi. Apical 0.25 of metatarsus III and tarsomeres II-V
infuscated; some tarsomeres of anterior legs tinged with red.
Holotype. Female, California, Los Angeles County, Tanbark Flat, VI-25-
50 (A. T. McClay); UCD. [The holotype female is the specimen used by
Scullen (1965) for his figures 108a, b, c under the name acanthophila.]
Paratypes. 1 1 males, 26 females all from California, Los Angeles County,
Tanbark Flat as follows: 5 males, 8 females same date as holotype (P. D.
Hurd, A. T. McClay, F. X. Williams) (UCB, UCD, CAS, MIS); 59 males,
10 females, VI-15-50 to VII-25-50 (H. M. Graham, J. C. Hall, P. D. Hurd,
A. T. McClay, W. O. Marshall, K. G. Whitesell, F. X. Williams) (UCB,
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NEW YORK ENTOMOLOGICAL SOCIETY
UCD, CAS, MIS, OSU); 4 males, 5 females, VI-29-52 to VII- 13-52 (R. L.
Anderson, D. E. Barcus, A. A. Grigarick, A. T. McClay, B. Miyagawa, S.
Miyagawa) (UCD); 3 males, 3 females, VI- 1 8-56 to VII- 1 8-56 (R. C. Bechtel,
R. M. Bohart, R. W. Bushing, J. I. Stage) (UCB, UCD).
Other specimens. In addition to the type series I have examined 1 7 1 males
and 52 females.
Distribution. Known only from California. Bohart and Grissell (1975)
provide details of distribution under the name C. minax. A series of spec-
imens (OSU) from California, Modoc County, Adin Pass, 5,000 ft., July 30,
1 955 (G. R. Ferguson, Joe Schuh) extends the range of the species northward.
This species is named for E. E. Grissell who, with R. M. Bohart in Bohart
and Grissell (1975), hrst showed that it was a distinct species.
Cerceris gandarai Rohwer
Cerceris gandarai Rohwer, 1912:470 (Male holotype. Federal District, Mex-
ico; USNM).
Cerceris micheneri Scullen, 1972:86 (Female holotype. Metachic, Chihua-
hua, Mexico; AMNH); Bohart and Menke, 1976:584. New Synonymy.
This species was described, hgured and keyed by Scullen (1972) under the
name of C micheneri. However, a study of the male holotype of C gandarai
shows that Scullen (1972) misidentihed Rohwer’s species. It is in fact con-
specific with and a senior synonym of C micheneri. The type female, allotype
male, and several paratypes of C. micheneri have been studied.
Cerceris rohweri, new species
Cerceris gandari (sic), Scullen, 1972:82, figs. 156a, b, c, d.
Cerceris gandarai, Bohart and Menke, 1976:581; Krombein, 1979:1734.
This species was described, figured and keyed by Scullen (1972) under
the name C gandarai. Since the name C. gandarai must now be applied to
the species previously known as C. micheneri, the species treated by Scullen
(1972) as C. gandarai is left without a name and is described here as C.
rohweri. It and true gandarai belong to an as yet undefined group of small
species, most of which occur in the central plateau area of Mexico and
contiguous portions of the United States. In having a fossette on tergum I
and a variably developed basal platform on sternum II, the group is allied
to the Cerceris fnitima group but separable by the absence of a carina on
coxa III.
In both sexes C. rohweri lacks microridging between punctures on the
scutum, whereas these microridges are conspicuous in C gandarai. Female
C. gandarai has an elevated, truncate process on the clypeal midsection; in
VOLUME 91, NUMBER 3
239
C. rohweri this process is absent, and, instead, there is a pair of somewhat
rounded, subquadrate lobes on the apical margin of the clypeal midsection.
In the male the clypeal brush of C. gandarai is somewhat tapered toward
the apex and not wider than the distance from the tentorial pit to the eye
margin; in C. rohweri the clypeal brush is rectangular and wider than the
distance from the tentorial pit to the eye margin.
Description. Jugal lobe of hind wing 0.3 times length of submedian cell;
fossette present on tergum I; carina absent on coxa III; basal platform of
sternum II a low, rounded swelling in male, essentially absent in female;
propodeal enclosure shiny with numerous micropunctures, median furrow
complete, crenulate; tegulae longer than wide, smooth; deflected posterior
portion of metasternal plate with triangles broadly fused at base, carinate
between. FEMALE: Length 9-1 1 mm; with small, sharp mesopleural tu-
bercle; mandible with large, broadly triangular tooth a little basad of middle;
clypeal midsection evenly convex, without a projecting process, terminating
apically in a pair of broad, subquadrate, slightly outturned lobes, fused at
base and with a transverse depression at base of lobes; eyes slightly divergent
below; macropunctures of clypeus and lower face well separated with nu-
merous micropunctures on lateral clypeal sections and lower face; punctation
denser in front of ocellar triangle than behind; scutal punctures separated
by 0.5-1 puncture diameter, at least across middle; anterior part of meso-
pleuron with a few ridges between punctures, punctures separated by flat
ridges on dise; propodeal punctures separated by many thick ridges of 0.5-
1 puncture diameter on side of and behind enclosure; punctures of black
portions of terga III-IV dense, some contiguous, some separated by flat,
shagreened ridges of 0.25-0.5 puncture diameter, pale portions with well
separated coarse punctures; pygidium narrowly oval, twice as long as mid-
width, uniformly rugose without pits at base, about as wide apically as
basally. MALE: Length 7-8 mm; mandible edentate; clypeal lip edentate,
about as wide as subantennal sclerite; clypeal brush strong, waxed, cupped,
wider than clypeal lip; clypeal midsection 1 .2 times as long as width between
tentorial pits; eyes parallel; punctures of clypeus and lower face separated
mostly by 1 or more puncture diameters; punctures on scutum, mesopleuron,
propodeum and terga as in female except ridges between punctures absent;
pygidium subrectangular, 1.8 times as long as greatest width, not widened
basally, slightly narrowed apically, slightly bowed out on the sides, coarsely
punctate. COLOR: Black with following parts yellow: base of mandible
(sometimes absent in male); clypeus except apically and basally in female
and apically in male; lower face from above sockets; spot on subantennal
selerite in male; small spot behind eye in female; interrupted band on prono-
tum; metanotum; tegulae in part; lateral spot on propodeum of female;
interrupted band or spots on tergum I, sometimes absent in male; posterior
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NEW YORK ENTOMOLOGICAL SOCIETY
bands on preapical terga, shallowly emarginate anteriorly; lateral spots on
sterna III-IV. Legs black and yellow suffused with red; tibiae I-II, except
dark spot behind, and trochanter III pale; femora with narrow, red apical
band in female, basal half pale in male; tarsi I-II reddish, less so in male;
tarsi III infuscate. Forewings with brownish infuscation along anterior one-
third; antennae with scape and pedicel black, flagellum brown below.
Holotype. Female, 17 mi. NE San Luis Potosi, S.L.P., Mexico, 6,200 ft..
Sept. 6, 1963 (Scullen and Bolinger); USNM.
Paratypes. 48 males, 12 females, all from the state of San Luis Potosi,
Mexico, as follows: 40 males, 7 females, same data as holotype (CAS, UCD,
OSU, USNM); 2 males, 19 mi. SW San Luis Potosi, 7,200 ft., IX-4-63
(Scullen and Bolinger) (OSU); 5 males, 1 female, 40 mi. S San Luis Potosi,
5,700 ft., IX-5-63 (Scullen and Bolinger) (OSU); 2 females, 15 mi. E San
Luis Potosi, 6,500 ft., X-3-57 (H. A. Scullen) (OSU); 1 female, 18 mi. SW
San Luis Potosi, 7,300 ft., X-2-57 (H. A. Scullen) (OSU); 1 male, 1 female,
10 mi. NE San Luis Potosi, 6,200 ft., VIII-22-54 (R. R. Dreisbach) (MIS).
Other specimens examined. 1 13 males, 20 females, from several states in
Mexico as follows: OAXACA: 18 males, 2 females, 5,600-6,850 ft., VI-27
to VIII-23 (OSU); PUEBLA: 9 males, 2 females, 5,380-6,900 ft., VI-7 to
VIII-24 (UCB, OSU); MEXICO: 17 males, 9 females, 7,400-8,550 ft., IV-7
to VIII-30 (MIS, OSU); VERACRUZ: 1 male, 5,000 ft., VI-9 (OSU); QUE-
RETARO: 46 males, 2 females, 6,500-7,400 ft., VI- 13 to IX- 19 (OSU);
AGUASCALIENTES: 1 male, 1 female, VIII-3 to VIII-9 (OSU, USNM);
ZACATECAS: 6 males, VII-3 to VIII-20 (MIS, OSU); DURANGO: 9 males,
2 females, 6,000-7,100 ft., VI- 17 to X-23 (MIS, OSU); COAHUILA: 1
male, 4,450 ft., IX-9 (OSU); NUEVO LEON: 5 males, 2 females, 4,600-
7,200 ft., VIII-30 to IX-22 (UCB, OSU, USU).
Scullen (1972) lists additional records (as gandarai) including a female
from Brooks County, Texas.
This species is named for the late S. A. Rohwer, long time student of
aculeate and other Hymenoptera, who described the species with which this
species has been misidentified.
ACKNOWLEDGMENTS
I am indebted to the following individuals and institutions for the loan of specimens used
in the course of this study (abbreviations in parentheses as used in the text): D. Azuma, Academy
of Natural Sciences of Philadelphia (ANSP); J. A. Powell, University of California, Berkeley
(UCB); R. M. Bohart, R. O. Schuster, University of California, Davis (UCD); W. J. Pulawski,
California Academy of Sciences, San Francisco (CAS); M. Favreau, American Museum of
Natural History, New York (AMNH); R. L. Fischer, Michigan State University, East Lansing
(MIS); B. C. Ratcliffe, University of Nebraska State Museum, Lincoln (NEB); J. D. Lattin,
Oregon State University, Corvallis (OSU); A. S. Menke, USDA Systematic Entomology Lab-
oratory, % U.S. National Museum of Natural History, Washington (USNM); F. D. Parker,
Utah State University, Logan (USU).
VOLUME 91, NUMBER 3
241
LITERATURE CITED
Banks, N. 1947. Synopsis of west coast Cerceris. Psyche 54:1-35.
Bohart, R. M. and E. E. Grissell. 1975. California wasps of the subfamily Philanthinae. Bull.
California Insect Surv. 19:1-92.
Bohart, R. M. and A. S. Menke. 1 976. Sphecid Wasps of the World. Univ. of California Press,
Berkeley, ix + 695 pp.
Cockerell, T. D. A. 1897. New Hymenoptera from New Mexico, U. S. A. Entomologist 30:
135-138.
Cresson, E. T. 1928. The types of Hymenoptera in the Academy of Natural Sciences of
Philadelphia other than those of Ezra T. Cresson. Mem. Am. Entomol. Soc. No. 5, pp.
1-90.
Krombein, K. V. 1967. Superfamily Sphecoidea. Pages 386-421 in: K. V. Krombein and B.
D. Burks, Hymenoptera of America North of Mexico, Synoptic Catalog. U.S. Dept, of
Agr. Monogr. 2, 2nd supplement, 584 pp.
Krombein, K. V. 1979. Sphecoidea. Pages 1573-1740 in: K. V. Krombein, Paul D. Hurd,
Jr., David R. Smith and B. D. Burks, Catalog of Hymenoptera in America North of
Mexico. 2:1199-2209. Smithsonian Inst. Press, Washington, D.C.
Mickel, C. E. 1918. New species of Sphecoidea from the central and western states. Nebraska
Univ. Studies (1917) 17:319-341.
Rohwer, S. A. 1912. Descriptions of new species of wasps in the U. S. National Museum.
Proc. U.S. Natl. Mus. 41:447-478.
Scullen, H. A. 1951. Tribe Cercerini. In: C. F. W. Muesebeck et al., Hymenoptera of America
north of Mexico. Synoptic catalog. U.S. Dept. Agric. Monogr. 2:1004-1013.
Scullen, H. A. 1960. Synonymical notes on the genus Cerceris, II. Pan-Pac. Entomol. 36:75-
80.
Scullen, H. A. 1965. Review of the genus Cerceris in America north of Mexico. Proc. U.S.
Natl. Mus. 116:333-548.
Scullen, H. A. 1972. Review of the genus Cerceris Latreille in Mexico and Central America.
Smithsonian Contr. Zool. 110:1-121.
Viereck, H. L. 1902. Hymenoptera from southern California and New Mexico, with descrip-
tions of new species. Proc. Acad. Nat. Sci. Philadelphia 54:728-743.
Viereck, H. L. and T. D. A. Cockerell. 1904. The Philanthidae of New Mexico, II. J. New
York Entomol. Soc. 12:129-146.
Received December 23, 1982; accepted May 13, 1983.
NEW YORK ENTOMOLOGICAL SOCIETY
91(3), 1983, pp. 242-245
GYNANDROMORPHIC DESERT FIRE ANT,
SOLENOPSIS AUREA WHEELER
(HYMENOPTERA: FORMICIDAE)*
James C. Cokendolpher and Oscar F. Francke
Departments of Entomology and Biological Sciences,
Texas Tech University, Lubbock, Texas 79409
Abstract. — \ gynandromorph of Solenopsis aurea Wheeler is described from an ant collected
in western Texas. The specimen is predominantly that of a queen, but the head is noticeably
male on the right half, female on the left half The reproductive system, both internally and
externally, is entirely female.
Gynandromorphs are individuals which have the male and female sexual
characters combined discretely. The character combinations can occur as
right-left halves, dorsoventral halves, antero-posterior halves, or in patches,
resulting in mosaics. Female ant characters can be contributed by any of the
castes present: queen-male (gynandromorph), worker-male (ergatandro-
morph), and soldier-male (dinergatandromorph). Combinations of worker,
queen, and male are also known (Donisthorpe, 1929).
Numerous mechanisms have been proposed to explain the occurrence of
gynandromorphs in animals. Most mechanisms deal either with fertilization-
related phenomena or with cytogenetic complications during early embryo-
genesis. Morgan and Bridges (1919), Rothenbuhler et al. ( 1952), Brust ( 1 966),
and Wigglesworth (1972) provide useful reviews and discussions on the
various mechanisms implicated in formation of insect gynandromorphs.
Gynandromorphs have been reported from 39 species in 22 genera of ants
(Donisthorpe, 1929; Wheeler, 1931, 1937; Buschinger and Stoewesand, 1971;
Hung et al., 1975). Two gynandromorphs have been reported from the genus
Solenopsis. A red imported fire ant, Solenopsis (Solenopsis) invicta Buren,
with male head, mosaic thorax, and female pedicel and gaster was described
by Hunget al. (1975). A thief ant, Solenopsis (Diplorhoptrum) fugax Latreille,
with female head and thorax, and male pedicel and gaster was reported by
‘ Supported by the Texas Department of Agriculture Interagency Agreements lAC (81-82)-
806 and lAC (82-83)- 1651. Contribution No. T-10-146, College of Agricultural Sciences, Texas
Tech University.
The publication costs of this article were defrayed in part by page charge payment. This article
must therefore be hereby marked ‘Advertisement” \n accordance with 18 U.S.C. §1734 solely
to indicate this fact.
VOLUME 9 1 , NUMBER 3 243
Figs. 1-4. Solenopsis aurea. 1. Dorsal view of gynandromorph. 2. Anterior aspect of gyn-
andromorph head. 3. Anterior aspect of male head. 4. Anterior aspect of female head.
Santschi (1910). Another anomaly reported for the red imported fire ant is
that of intercastes, female individuals exhibiting worker and queen characters
(Glancey et ah, 1980).
The present gynandromorph is that of a desert hre ant, Solenopsis {Sole-
nopsis) aurea Wheeler, which is preserved along with normal male and
female siblings (cat. no. 6461) in the Entomological collection. The Museum,
Texas Tech University. Nineteen colonies of S. aurea where collected 2.6
miles ESE of Southland, Garza Co., Texas, on 4 June 1982 and were kept
alive in the laboratory. On 15 June 1982 all the colonies were closely ex-
amined for the presence of external parasites, at which time the malformed
ant was discovered. The gynandromorph and six male and female siblings
were isolated on 2 July for closer observations. The gynandromorph, but
not its siblings, died 6 July 1982.
The sexes of S. aurea are distinctive and easily separated. The gynandro-
morph (Figs. 1, 2) is predominantly female, with male characteristics most
distinctive on the head. A normal male head (Fig. 3) is darkly pigmented
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NEW YORK ENTOMOLOGICAL SOCIETY
and small in size. The ocelli and compound eyes are large, and the mandibles
and antennal scapes are small. The number of segments in the antenna also
differs between the sexes: 12 for males, 10 or 11 for females. The head of
the gynandromorph (Fig. 2) is clearly asymmetrical. The right half is that
of a male: dark, with large ocelli, reduced antennal scape and mandible, and
with a 12-segmented antenna. The left half is distinctly female: light in color,
with small ocellus, large mandible, long antennal scape, and 1 1 -segmented
antenna. The thorax and its appendages, except for minor differences, are
those of a female. The wing bases appear to have been removed by the
gynandromorph’s siblings, or at least dealation did not occur normally. The
pronotum is slightly darker on the right side, although not as dark as on
males. The small ridge on the declivous face of the propodeum, characteristic
of females, is absent on the right half of the postpetiole. The internal anatomy
of the head, thorax, and pedicel was not examined. The abdomen is four
segmented as in females (five segments in males) and shows but a slight
darkening in color on the right half, although still much paler than in males.
Dissection of the abdomen revealed a complete bilaterally symmetrical fe-
male reproductive system. Developing oocytes are present in the ovarioles.
The spermatheca is well developed and as large as that of mated females,
but no spermatozoa were found in a section of spermathecal duct stained
with giemsa. The poison gland and sting are present and of normal size.
ACKNOWLEDGMENTS
We would like to thank Drs. Sherman A. Phillips, Jr., James K. Wangberg, and Jeff Whit-
worth for their comments on the manuscript. Dr. Phillips was also helpful in the examination
of the gynandromorph’s internal anatomy. Thanks are also extended to Ms. Lorie A. Prien for
typing the various drafts of the manuscript.
LITERATURE CITED
Brust, R. H. 1966. Gynandromorphs and intersexes in mosquitoes (Diptera : Culicidae). Can.
J. Zool. 44:911-921.
Buschinger, A. and H. Stoewesand. 1971. Teratologische Untersuchungen an Ameisen (Hy-
menoptera : Formicidae). Beitr. Ent. 21:211-241.
Donisthorpe, H. 1929. Gynandromorphism in ants. Zool. Anz. 52:92-96.
Glancey, B. M., R. K. Vander Meer, A. Glover and C. S. Lofgren. 1980. Observations of
intercastes in Solenopsis invicta Buren. Florida Ent. 63:346-350.
Hung, A. C. F., W. N. Norton and S. B. Vinson. 1975. Gynandromorphism in the Red
Imported Fire Ant, Solenopsis invicta Buren (Hymenoptera : Formicidae). Ent. News
86:45-46.
Morgan, T. H. and C. B. Bridges. 1919. Contributions to the genetics of Drosophila mela-
nogaster. I. The origin of gynandromorphs. Carnegie Inst. Wash. Publ. No. 278, pp. 3-
1 22.
Rothenbuhler, W. C., J. W. Gowen and O. W. Park. 1952. Androgenesis with zygogenesis in
gynandromorphic honeybees {Apis }nellifera L.). Science 1 15:637-638.
VOLUME 91, NUMBER 3
245
Santschi, F. 1910. Contributions a la faune entomologique de la Roumanie, formicides cap-
turees par Mr. A. L. Montandon et determinees par Mr. le Dr. F. Santschi. Bull. Soc.
Sci., Bucarest 19:648-651.
Wheeler, W. M. 1931. Concerning some ant gynandromorphs. Psyche 38:80-85.
Wheeler, W. M. 1 937. Mosaics and Other Anomalies Among Ants. Harvard University Press,
Cambridge, Massachusetts, 95 pp.
Wigglesworth, V. B. 1972. The Principles of Insect Physiology, 7th Edition. Chapman and
Hall, London, 827 pp.
Received January 21, 1983; accepted April 19, 1983.
NEW YORK ENTOMOLOGICAL SOCIETY
91(3), 1983, pp. 246-251
A REDEFINITION OF DISDERIA AND ADDITION OF
A NEW SPECIES (HEMIPTERA: PENTATOMIDAE)
L. H. Rolston
Department of Entomology, Louisiana Agricultural Experiment Station,
Baton Rouge, Louisiana 70803
Abstract.— genus Disderia is redefined and a new species from Oaxaca, Mexico, added.
Parts of the male genitalia of the three known species and the genital plates of the females of
D. decorata and D. inornata are hgured. A key to the species is given.
Bergroth (1910) established the genus Disderia for Phalaecus decoratiis
Distant, and Ruckes (1959) added D. inornata. A third species is described
here and additional details are given concerning the genitalia of the other 2
species. The genus is redefined to broaden and augment the original descrip-
tion.
Among the Pentatomini of the Western Hemisphere, Disderia may be
recognized by the combination of 3-segmented tarsi and a grossly elongated
abdominal spine which reaches the procoxae.
Disderia Bergroth, 1910
Disderia Bergroth, 1910, p. 20; Distant, 1911, p. 252; Ruckes, 1959, p. 27.
Median basal spine of abdomen reaching procoxae, depressed basally,
compressed distally. Ostiolar rugae straight, extending about three-fourths
of distance from inner margin of ostiole to lateral margin of metapleuron.
Mesosternum and metasterum anteriorly with low median carina. Femora
without apical spine or other armament. Tarsi three-segmented.
Length of head about 0.7 width across eyes; juga narrowly contiguous or
nearly so at apex of head (Fig. 14); distal end of first antennal segment
surpassing apex of head; bucculae evanescent at base of head, terminating
near distal end of first rostral segment; apex of rostrum reaching between
mesocoxae. Pronotum about 2.5 times wider than long at meson; anterior
margin contiguous with and projecting a little laterad of eyes; anterolateral
margins entire. Scutellum 0. 1-0.2 longer than wide at base; frena extending
about 0.6 length of scutellum from base toward apex.
Theca of male genitalia without processes on apical margin. Eighth par-
atergites of female and 8th sternite of males bearing spiracles.
lype species. Phalaecus decoratiis Distant by original designation.
VOLUME 91, NUMBER 3
247
Figs. 1-7. D. decorata. 1. Pygophore and last abdominal segment, ventral view. 2. Genital
cup; paramere (PA); proctiger (PR). 3. Paramere, ventral view. 4. Aedeagus, dorsal view; lateral
diverticula (L. DIV.); median diverticulum (M. DIV.). 5. Aedeagus, ventral view; lateral di-
verticula (L. DIV.); ventral diverticula (V. DIV.); median penial plates (M. P. P.). 6. Aedeagus,
lateral view; lateral diverticula (L. DIV.); ventral diverticula (V. DIV.); penisfilum (P.). 7. Genital
plates; triangulum (T.).
Comment. Perhaps it should be emphasized, because of conflicting state-
ments, that Distant (1880) was correct in describing the mesosternum and
metasternum as elevated (p. 83), although Bergroth (1910) may also have
been correct in his belief that Distant mistook the abdominal spine for sterna.
In all 3 species the mesosternum and anterior portion of the metasternum
have a low median carina which is generally rather obscured by the abdom-
inal spine.
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NEW YORK ENTOMOLOGICAL SOCIETY
Figs. 8-13. D. inornata. 8. Pygophore, ventral view. 9. Genital cup; parameres (PA); proc-
tiger (PR). 10. Aedeagus, everted, lateral view; lateral diverticula (L. DIV.); ventral diverticulum
(V. DIV.); penisfilum (P). 1 1. Aedeagus, uneverted; lateral diverticula (L. DIV.). 12. Paramere,
ventrolateral view. 13. Genital plates.
Disderia parda, new species
(Figs. 14-19)
Description. Dorsum dark yellowish brown; punctation black, on coria
especially dense, on pronotum and scutellum disposed in part as irregular
VOLUME 91, NUMBER 3
249
Figs. 14-19. D. parda. 14. Head. 15. Pygophore, ventral view. 16. Genital cup; parameres
(PA). 17. Aedeagus, everted, lateral view, ventral diverticulum (V. DIV.); median penial plates
(M. P. P.); penisfilum (P.). 18. Aedeagus, uneverted, ventral view. 19. Parameres, ventral view.
rows with a general transverse orientation. Venter much paler; thoracic
punctures of moderate size, mostly dark, irregularly distributed; abdominal
punctures for most part progressively larger darker and less numerous from
lateral abdominal margins toward meson. Length of body 7.8 to 8.1 mm.
Head 1.9 mm wide across eyes, 1.4 mm long. Three basal segments of
antennae concolorous with venter, bearing numerous dark dots; ultimate
two segments darker, immaculate; length of segments 0.5; 0.8 to 0.9; 0.9;
1.2; 2.0 mm. Pronotum 5.0 to 5.1 mm wide, 2.0 mm long at meson; an-
terolateral margins slightly convex; humeri rounded, scarcely produced. Scu-
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NEW YORK ENTOMOLOGICAL SOCIETY
tellum 3.1 mm wide at base, 3.5 to 3.6 mm long; basal disk darkened by
diffusion of black about punctures. Coastal angle of coria reaching posterior
margin of penultimate tergite; membrane of hemelytra darkly fumose, veins
simple or furcate. Connexiva little to moderately exposed, black with pale
marginal spot in middle of each segment.
Evaporative area large, each covering much of metapleuron and meso-
pleuron, matte, similar to remainder of pleura in color and punctation. Legs
with fuscous dots, those on femora larger and less numerous than on tibiae;
superior surface of all tibiae sulcate. Spiracular peritremes dark. Abdominal
margins thinly edged in black except in middle of each segment.
Digitiform process on posterolateral angles of pygophore directed oblique-
ly mesad (Figs. 1 5, 1 6); conspicuous impression in lateral pygophoral surface
located outside of genital cup opposite apex of parameres. Parameres irreg-
ularly palmate from dorsal aspect; ventrolateral subapical surface covered
with fine serrate ragae (Fig. 1 9). Median penial plates forming large concave
area, heavily sclerotized and pigmented, with short penishlum emerging
within ventral margin; ventral conjunctival diverticulum increasingly scler-
otized and pigmented toward apex, distally bifurcate, terminating on each
side in hook (Fig. 17); most of conjunctiva and penishlum retractable into
theca, of pigmented parts leaving only distal part of ventral diverticulum
exposed in deep ventral emargination of theca (Fig. 18).
Holotype. 6, labeled (a) “Mexico, Oaxaca, 21.8 mi. n. Juchatengo. 7100'
III, 23, 1966, in bromeliads” (b) “George E. Ball, D. R. Whitehead, collec-
tors”. Deposited in U.S. National Museum, type no. 75558.
Paratype. S, labeled as holotype, in author’s collection.
Discussion. Disderia has no great likeness to any other genus, as both
Bergroth (1910) and Ruckes (1959) remarked, and the intrageneric dissim-
ilarities suggest that the relationship among species is not close.
Although the tibiae of D. decorata are asulcate and those of the other 2
species are sulcate, the external morphology other than the genitalia is con-
sistent with expectations for congenericity. Both D. decorata and D. ornata
have eversible conjunctiva, i.e., the conjunctiva and associated phallic struc-
tures are largely contained within the theca (Figs. 11, 18), from which they
may be extracted manually and from which they are presumably everted
during copulation (Figs. 10, 17). This does not seem to be the case in D.
ornata.
There are notable differences among the 3 species in the conjunctival
structures. D. decorata has a median, conical diverticulum that is posterior
to a dome shaped expanion, a pair of lateral diverticula and a pair of ventral
diverticula (Figs. 4-6). These are entirely membranous with the exception
of the apex of each member of the ventral pair. Small median penial plates
are present at the base of the penishlum. Both D. inornata and D. parda
have an elongated ventral diverticulum that bifurcates distally with each
part terminating in a hook (Figs. 10, 17). The two species diverge sharply
VOLUME 91, NUMBER 3
251
in that D. parda has large median penial plates while D. inornata has none,
and D. inornata has a pair of appendicular diverticula laterally which are
not represented in D. parda.
The parameres of D. ornata and D. parda are distally flattened and es-
sentially horizontal (Figs. 2, 3, 16, 19); those of D. inornata are distally
hooked and essentially vertical (Figs. 9, 12).
Among females, the triangulum is exposed in D. decorata but not in D.
inornata (Figs. 7, 13). The female of D. parda is unknown.
Disderia appears to be an old genus that exhibits a remarkable degree of
divergence in the genitalia of the 3 species. The few species and the large
interspecific differences are consistent with the assumption of generic antiq-
uity. The spiracles on the 8th paratergites and 8th sternite retain the primitive
character state. Eversible conjunctiva, as exhibited by D. inornata and D.
parda, occurs infrequently among Pentatomini, but it is characteristic of
several groups, e.g. the acanthosomatids, that are generally considered to be
less recent phylogenetically than the Pentatomini.
KEY TO SPECIES OE Disderia
1. Tibiae asulcate; large white spot in each basal angle and at apex of scutellum, other
white markings on body; little of pygophore evident from ventral view (Fig. 1 )
D. decorata (Distant)
- Tibiae sulcate; conspicuous pale markings confined to connexiva; pygophore prominent
from ventral view (Figs. 8 and 1 5) 2
2. Dorsum light brown, abdominal punctation nearly concolorous with sternites; length
without membrane about 10 to 12 mm; posterolateral angles of pygophore deeply
incised laterally, without digitiform process, parameres hook-shaped (Figs. 9, 12)
— D. inornata Ruckes
- Dorsum dark brown, abdominal punctation mostly much darker than sternites; length
without membrane about 8 mm, posterolateral angles of pygophore not incised, each
bearing slender digitiform process directed obliquely mesad; parameres irregularly
palmate (Figs. 16, 19) D. parda, new species
ACKNOWLEDGMENT
The description of Disderia parda is based on specimens donated by Dr. George E. Ball of
the University of Alberta.
LITERATURE CITED
Bergroth, E. 1910. Notes on the genus Phalaecus StM. Entomol. News 2 1(1): 18-21.
Distant, W. L. 1880-1893. Insecta. Rhynchota, Hemiptera-Heteroptera. In: F. D. Godman
and O. Salvin, Biologia Centralia-Americana. Vol. 1.
Distant, W. L. 1911. Rhynchotal notes liii. Neotropical Pentatomidae. Ann. Mag. Nat. Hist.
(8)7:242-258.
Ruckes, H. 1959. Disderia inornata, a new species of pentatomid from Mexico (Heteroptera,
Pentatomidae). J. New York Entomol. Soc. 67:27-30.
Received July 1, 1982; accepted January 14, 1983.
NEW YORK ENTOMOLOGICAL SOCIETY
91(3), 1983, pp. 252-263
A STUDY OF WINGED QUEENS OF THE COLORADO HONEY ANT,
MYRMECOCYSTUS MEXICANUS, IN CAPTIVITY
John R. Conway
Department of Biology, Elmhurst College,
Elmhurst, Illinois 60126
Abstract. — queens (24) were captured issuing from a honey ant nest north of Colorado
Springs prior to the nuptial flight and placed in Janet-type nests. Six nests housed one queen
and six housed three queens each. These nests were fed three different diets to determine their
effect on queen longevity and egg production. Maximum longevities occurred in nests given
the most complete diet. All nests produced eggs 7-12 days after swarming, but only one larva
developed, and it lived only 72 days. Data on egg numbers, locations, durations and clump
size are presented. Observations were made on dealation, abdominal distension, mite infes-
tation, and queen behavior and death. Dealation attempts took place over a long period and
were successful in only two queens. Queens developed swollen abdomens in every nest except
one. However, neither dealation nor a swollen abdomen is essential for egg production. There
was no hostility in three-queen nests and they cooperated caring for eggs. Behavior, longevity,
and reproductive potential of virgin queens are compared to fertilized queens in captivity.
Although a number of investigators such as McCook (1882), Wheeler
(1908, 1910, 1913, 1917), Creighton (1950), Cazier and Statham (1962),
Gregg (1963), Cazier and Mortenson (1965), Slocumb (1966), Conway (1975,
1977, 1980a, b, 1981a, b), Holldobler (1976), Snelling (1976), Kay (1978),
Kay and Whitford (1978), Chew (1979), and Holldobler and Lumsden (1980)
have discussed the biology of Myrmecocystus mexicanus, there has been no
systematic study of winged queens in captivity. This study attempts to rem-
edy that deficiency and had the following objectives:
1. To see if winged, presumably virgin queens, issuing from a nest prior
to the nuptial flight, could lay eggs and rear young.
2. To determine longevity of winged queens.
3. To note behavioral differences between winged queens in solitary con-
finement and those kept in groups of three.
4. To investigate the effect of three diets on winged queens.
5. To see what effect natural and artificial dealation would have on queen
behavior and reproduction.
6. To compare behavior and reproductive potential of virgin queens with
fertilized queens (Conway, 1981a).
The publication costs of this article were defrayed in part by page charge payment. This article
must therefore be hereby marked “Advertisement” in accordance with 18 U.S.C. §1734 solely
to indicate this fact.
VOLUME 91, NUMBER 3
253
MATERIALS AND METHODS
Twenty-four winged queens were captured as they left a M. rnexicanus
nest on a mesa north of Colorado Springs on July 28, 1972 prior to the
nuptial flight, and placed in plaster Janet-type nests on July 30.
The plaster nests contain two glass-covered chambers, each 10.8 cm long,
8.9 cm wide, and 0.6 cm high, connected by a passageway 1.3 cm long and
1.6 cm wide. The passageway was blocked with foam to form two com-
partments. Water was added daily to a small sponge or piece of cotton in
each chamber.
Three nests, with six compartments, were set up with one winged queen
in each compartment. Queens in the first nest received the complete diet of
sugar water, protein and fat (see section on sustenance), and water. Queens
in the second nest received sugar water and water, and those in the third
nest received water only.
Three nests, with six compartments, were set up with three winged queens
in each compartment. Queens in the first nest were given the complete diet,
those in the second nest received sugar water and water, and those in the
third nest drank water only.
Sugar water given to colonies was dyed with blue food coloring so its
intake and distribution could be followed.
RESULTS AND DISCUSSION
Queen sustenance— complete-diet nests. Queens in these nests were given
the most complete diet which consisted of a variety of foods, such as sugar
water, honey solutions, milk, sugar-sweetened milk, peanut butter, cereal,
hamburger, live Drosophila, mealworms, and water. Queens fed mainly on
blue sugar water and their abdomens usually became bluish. The only other
foods they ate were honey solutions. Drosophila, and one mealworm. There
were only two observations of a queen drinking water.
Sugar water and water nests. Queens in these nests received blue sucrose
solutions frequently, blue honey water infrequently, and tap water almost
daily. Most queens developed swollen blue abdomens, but there were no
observations of them drinking tap water.
Water only nests. Queens in these nests received tap water almost daily
and periodically were observed drinking it.
Queen longevity. In solitary-queen nests, maximum longevity occurred in
a queen receiving the complete diet, who lived 194 days, and minimum life
span belonged to a queen receiving water only who survived 88 days (Table
1).
Single queens provided with complete diets lived longest (mean 149 days).
Those in water-only nests lived only slightly longer (mean 104.5 days) than
those fed sugar water and water (mean 100.5 days) (Table 1).
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NEW YORK ENTOMOLOGICAL SOCIETY
Table 1. Longevity, wing condition, and method of dealation of solitary and grouped Myr-
mecocystus mexicamis queens supplied with three diets in captivity.
Longevity (days)
Nest
1st queen
2nd queen
3rd queen
Mean
A— solitary queen — sugar
194
water, protein, water
winged
149
B — solitary queen — sugar
104
water, protein, water
winged
A— solitary queen — sugar
107
water and water
B— solitary queen — sugar
winged
94
100.5
water and water
wingless —
natural
A — solitary queen — water
88
only
wingless-
B — solitary queen— water
natural
121
104.5
only
winged
A — 3 queens— sugar water.
275
339
419
protein, water
winged
wingless-
winged
artificial
250
B— 3 queens— sugar water.
5
190
272
protein, water
winged
winged
winged
A-3 queens— sugar water
85
221
249
and water
wingless —
winged
winged
B — 3 queens— sugar water
artificial
128
155
195
172.2
and water
winged
winged
winged
A — 3 queens— water
48
149
237
only
wingless-
winged
winged
B — 3 queens— water
artificial
175
189
194
165.3
only
winged
winged
winged
In three-queen nests, both maximum and minimum life spans, 419 and
5 days respectively, occurred in colonies receiving complete diets. Never-
theless, these nests still exhibited the longest mean life span (250 days). The
queens receiving sugar water and water lived slightly longer (mean 172.2
days) than those given water only (mean 165.3 days) (Table 1).
On the average, solitary queens had shorter life spans (mean 118 days)
than queens in groups of three (mean 195.8 days). Perhaps queen cooperation
and interaction increase longevity. It is interesting to speculate if this situ-
VOLUME 91, NUMBER 3
255
ation is peculiar to virgin queens and whether it would be reversed if fertilized
queens were placed together.
In summary, maximum longevities occurred in one- and three-queen nests
receiving the complete diet. Individuals in these nests received comparable
diets to fertilized queens in another study (Conway, 1981a) who lived even
longer (mean 297.6 days). Thus, as might be expected, fertile queens lived
much longer than their solitary unfertilized counterparts. What is surprising
however, is that when virgin queens were in groups, their mean life span
(250 days) approached that of fertile queens (297.6 days).
Data suggest that wingless virgin queens do not live as long (mean 130.8
days) as winged ones (mean 188.4 days). And, among wingless queens, the
average life of queens who dealated themselves was shorter (9 1 days) than
among those artificially dealated (157.3 days).
First appearance of eggs. Eggs first appeared in virgin-queen colonies 7 to
12 days after swarming (Table 2). They appeared sooner, 4 or 5 days after
the nuptial flight, in fertilized-queen nests (Conway, 1981a).
Eggs were laid in every virgin-queen nest. The number of eggs in the first
clump varied from 1 to 8 (Table 2), but was somewhat larger, 5 to 10, in
fertilized-queen colonies (Conway, 1981a).
Egg survival. The number of days eggs were present in solitary virgin-
queen nests ranged from 66 to 138. There was no correlation with persistence
of eggs and completeness of diet except that eggs survived longer, on the
average, in complete-diet nests (Table 2).
The number of days eggs were present in three-queen nests varied from
92 to 340. The two nests given water only, contained eggs for the shortest
period (mean 94). Both colonies nourished on sugar water had eggs for 145
days, the added carbohydrate seeming to increase egg longevity. Nests pro-
vided with the complete diet had eggs for the longest period (mean 257.5)
(Table 2).
Eggs persisted longest in one- and three-queen nests which received the
complete diet. Eggs also survived much longer in three-queen colonies than
in solitary-queen nests, except those receiving only water. In water-only
nests, egg durations were not significantly diflerent (Table 2).
The only virgin-queen nests comparable to fertilized-queen colonies in
another study (Conway, 1981a) were those fed the complete diet. Fertile
queens maintained eggs a couple weeks longer (mean 132.6 days) than sol-
itary virgin queens in those nests (mean 1 14.5 days), but not nearly as long
as nests with three queens (mean 257.5 days) (Table 2).
Disappearance of eggs. In four of six solitary-queen nests, eggs disappeared
before the queen expired, but in two water-only nests, the queen passed away
while some eggs were still present.
In all three-queen colonies, eggs vanished before the third queen died, and
in one water-only nest, eggs perished before the first queen expired. In nests
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NEW YORK ENTOMOLOGICAL SOCIETY
Table 2. Data on first appearance, duration, maximum number, and disappearance of eggs
in solitary and grouped Mynnecocystus mexicanus queens supplied with three diets in captivity.
Nest
Appearance
of 1st
eggs — days
after
swarming
(7/28/72)
Number of
1st eggs
Disappearance
of eggs
Egg
durations
(days)
Mean
duration
(days)
Egg maxima
and dates
A— solitary queen — sugar
water, protein, water
9
8
10/20/72-
11/29/72
2/1/73
138
114.5
18
(8/13/72)
B— solitary queen— sugar
water, protein, water
11
1
1 1/7/72
91
17
(9/23/72)
A — solitary queen— sugar
water, water
10
4
10/12/72
66
69.5
11
(9/12/72)
B— solitary queen— sugar
water, water
10
8
10/19/72
73
28
(9/10/72)
A— solitary queen— water
only
9
2
10/25/72
81
96
25-30
(9/7/72)
B — solitary queen — water
only
10
5
11/26/72
111
20
(8/22/72)
A— 3 queens— sugar water,
protein, water
8
2
11/30/72-
2/1/73
9/11/73
340
257.5
60
(8/28/72)
B — 3 queens— sugar water,
protein, water
12
4-5
10/4/72-
12/11/72
4/14/73
175
40
(8/28/72)
A— 3 queens— sugar water,
water
8
clump
12/28/72
145
145
60
(8/31/72)
B — 3 queens— sugar water,
water
10
6
1/1/73
145
60
(9/9/72)
A— 3 queens— water
only
8
1
11/5/72
92
94
40
(8/22/72)
B— 3 queens— water
only
7
4
11/8/72
96
44
(9/20/72)
receiving sugar water and water, and the other water-only nest, eggs disap-
peared before the demise of the second queen. Only in the two complete-
diet nests did the third queen continue to lay eggs after the death of the
second.
Fluctuation in egg number. Egg numbers fluctuated in all virgin-queen
nests, and in some, they dropped to zero for a time before increasing again.
This seemed to occur because the queen laid eggs, some deteriorated, and
then she deposited more. Queens were not observed consuming eggs.
VOLUME 91, NUMBER 3
257
In three complete-diet nests, eggs vanished for a period and then reap-
peared (Table 2), but did not reach the maximum number attained originally.
One exception was a complete-diet, solitary-queen nest, which achieved the
original maximum of 18 eggs during the second period of deposition. In
four of seven fertile-queen colonies, eggs also perished before returning again
(Conway, 1981a).
It is hypothesized that queens ceased laying eggs when there was a tem-
porary lack of protein. When protein was added, usually in the form of
Drosophila and mealworms, eggs reappeared, although in one case not until
40-50 days later. In one nest in which eggs did not recur, the solitary queen
probably received very little protein.
In both sugar water and water-only nests, ova never reappeared after
declining to zero for several days, presumably due to a lack of protein.
Maximum number of eggs. The maximum number of eggs in one-queen
nests varied from 1 1 to approximately 30 (mean 20.3). The largest numbers
occurred in a water-only nest (25-30) and sugar water nest (28). The maxima
in the two colonies given the complete diet were 17 and 18 (Table 2). Egg
maxima in fertilized-queen nests given a comparable diet varied from 5 to
34 (mean 13.6) (Conway, 1981a). It is surprising that the mean maximum
(20.3) is higher in virgin, solitary-queen nests than in fertilized-queen nests
(13.6).
In three-queen colonies egg maxima varied from 40 to 60. Greatest num-
bers occurred in nests with the complete diet or receiving sugar water. The
one complete-diet colony in which the total was not 60 may be explained
by the fact that one queen died almost immediately and hence all eggs were
laid by two queens. Two nests receiving water only had egg maxima of 40
and 44 (Table 2).
Evidence is inconsistent with regard to nourishment and egg maxima in
virgin-queen nests. In one-queen nests, greatest numbers appeared in nests
fed either water only or sugar water. But in three-queen colonies, the greatest
numbers were in nests given the complete diet or sugar water.
Egg maxima in three-queen colonies were greater than in solitary-queen
nests, suggesting that more than one queen, and probably all three, laid eggs.
And since the mean total for solitary queens was 20, and the mean maximum
for three-queen colonies was 5 1 , there is the likelihood that each queen lays
a total of about 20 eggs. However, since the mean three-queen total, 5 1 , is
somewhat below the theoretical 60, there may be some inhibition to egg
deposition due to group interaction.
Egg grouping. In five of six solitary-queen nests, eggs were predominantly
or only in one clump. In the sixth nest, eggs were scattered most of the time.
Maximum number of clumps varied from zero to two. In fertilized-queen
colonies eggs were also predominantly in one clump (Conway, 1981a).
Although eggs were mainly in one clump in three-queen colonies, the
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NEW YORK ENTOMOLOGICAL SOCIETY
maximum number of clumps varied from one to four. In one colony there
were almost as many observations of scattered ova as of eggs in a clump.
Eggs were sometimes in two clumps of different sizes. For example, in one
nest there were 40 eggs in one clump and 6 in another.
When total number of eggs is divided by number of clumps, the range is
6V2 to 30 eggs per clump (mean 13.7). It is postulated that the mean of 13.7
reflects an optimum clump size which can best be cared for and transported
by the queen, and that as egg numbers increase, they are divided into several
clumps to preserve optimum size.
Defective eggs. Defective eggs were variously described in my notes as
“small, yellow, shriveled, wrinkled, not firm or turgid, containing internal
globules, caved in, crushed, dried up, decaying, and amorphous” and began
appearing about one week after the first eggs were laid. Actually all ova were
non-viable in that none developed into adults, and only one metamorphosed
into a larva. Defective eggs were sometimes scattered and sometimes in
separate piles from normal eggs. By contrast, there was only one observation
of defective eggs in fertilized-queen colonies (Conway, 1981a).
Egg color. It is probable that yellow and white eggs occur in all nests, but
white eggs (more viable?) seem more abundant in nests given the complete
diet or sugar water whereas yellow eggs seem more numerous in nests re-
ceiving water only. White eggs also seemed to predominate in fertilized-
queen colonies (Conway, 1981a).
Location of eggs. Most observations from virgin-queen (150) and fertile-
queen nests were of eggs close to moisture (damp sponge or piece of cotton).
There were 35 sightings of eggs away from moisture in virgin-queen nests.
Care of eggs. Virgin queens were observed tending eggs in every nest except
one, just as fertile queens do (Conway, 1981a). Tending consists of standing
over, holding, or carrying eggs. Queens seem to tend both healthy and de-
fective eggs.
In three-queen colonies, eggs were most frequently tended by one queen
at a time (72 observations), but often (3 1 observations) all three queens were
around an egg cluster or single egg. There were seven observations of two
queens tending a group of eggs. Sometimes, difierent queens watched over
separate clumps simultaneously (6 observations). There was only one ob-
servation of a queen replacing another over a clump and this occurred when
one queen left the eggs and another came over to carry them away.
Queens are protective towards eggs and often picked them up when nests
were opened. On one occasion, a queen found an egg mired in honey, plucked
it out, and placed it with the others. Despite this attention, queens stepped
on eggs from time to time.
Egg deposition. One of the most significant findings of this study was that
queens which do not participate in the nuptial flight may nevertheless lay
eggs. The question arose as to whether dealation was necessary to trigger egg
VOLUME 91, NUMBER 3
259
laying, but clearly it is not, since eggs appeared in nests with only winged
queens. Nor does a swollen abdomen seem essential, since one queen laid
eggs but was never distended.
Larvae. A larva developed only in one three-queen colony given sugar
water. It emerged 32 days after eggs were first observed and survived 72
days. It emerged later and survived for a shorter duration than larvae in
fertile-queen colonies. In these nests first larvae appeared 17 to 23 days
(mean 20.1) after eggs were laid and were present 171 to 290 days (mean
261.1) (Conway, 1981a).
It seems unusual that the larva developed in a sugar water nest instead of
one given a complete diet.
The larva’s blue midgut attested to its success in soliciting blue sugar
water. Larvae in fertile-queen colonies also fed on eggs, larvae, dead workers
and mealworms (Conway, 1981a) and may explain why they completed
metamorphosis.
All queens seemed to care for and transport the larva. On one occasion,
when it was in water, a queen pulled it out. Despite their solicitude, queens
occasionally stepped on the larva.
Queen cooperation. A primary question of the study was how winged
queens would behave when put together after the nuptial flight. There was
no hostility among queens in the three-queen colonies. In fact, they seemed
to share responsibility caring for eggs.
A possible case of altruism occurred when a queen was accidentally tipped
onto her back. Another queen went over and tapped the first with her an-
tennae. The upside-down queen then seized the helper and righted herself.
Queen trophallaxis. Trophallaxis occurred in every three-queen colony.
The exchange took different forms; either both queens had their mandibles
open, both had them closed, or one had hers open while the other kept hers
closed. One queen regurgitated a droplet between her open jaws and at-
tempted to pass it to others, but if there were no takers, she would swallow
it again. Some queens disgorged fluid on the floor prior to their death.
Movement of gravel. Queens usually scattered the pile of gravel placed in
the nests 2 to 15 days after it was added. They also deposited gravel and
other debris (dried eggs, wings, etc.) on food and water placed in nests.
Queen abdominal distension. Queens developed distended abdomens in
every nest except a solitary-queen one given water only. Queens became
swollen 7 to 43 (usually 7-10) days after the nuptial flight. Fertilized queens
in four of seven nests became inflated somewhat sooner, 4-6 days after the
nuptial flight (Conway, 1981a).
Virgin queens in water-only nests became dilated either by drinking, pro-
ducing eggs, or both. Since the solitary queen in one nest did not enlarge
and yet laid eggs, it follows that a distended abdomen is not an essential
accompaniment of egg production.
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NEW YORK ENTOMOLOGICAL SOCIETY
Queens in sugar water and complete-diet nests developed inflated blue
abdomens, hence, at least part, if not all their swelling, could be attributed
to drinking the blue sucrose solution. In six fertilized-queen nests, the queen
also developed an inflated blue abdomen 1 to 13 days after blue sugar water
was first presented.
All wingless virgin queens, whether artificially or naturally dealated, de-
veloped enlarged abdomens rapidly. Wingless fertilized queens also became
inflated (Conway, 1981a) and thus it was theorized that dealation triggers
distension. That this is not the case was demonstrated by the fact that winged
virgin queens became distended as well.
It was noted that in almost every three-queen colony, one or two queens
were more inflated than others. There were a few observations in both
solitary- and three-queen nests which suggest some queens fluctuated in
abdominal size. This fluctuation also occurred in some fertilized-queen nests
(Conway, 1981a).
Although the majority of queens developed distended gasters, most were
not swollen at death, apparently because they disgorged fluid prior to passing
away. There were four exceptions, two winged and two wingless queens,
which were inflated at death. Some fertilized queens also disgorged fluid and
began to deflate prior to death (Conway, 1981a).
Queen dealation. Fertilized queens dealate themselves quickly. In fact, all
were wingless when collected shortly after the nuptial flight (Conway, 1981a).
By contrast, dealation attempts in virgin-queen colonies were often unsuc-
cessful and took place over a long period of time, beginning as early as
August 5, 1972 and continuing, in one instance, until January 27, 1973.
All queens in three-queen colonies retained their wings, except three de-
alated with forceps. There were a few attempts by queens in two nests to
remove their wings but they were not successful. Queens tried to remove
their wings in all solitary-queen nests except one, and two individuals suc-
ceeded eight days after the nuptial flight.
In conclusion, it appears that solitary virgin queens attempt to dislodge
their wings more often and with more success than grouped queens. If fer-
tilized queens remove their wings in the same fashion as virgin queens
attempt to, they first spread them, lift their hind legs simultaneously over
the tops, and step on them. Sometimes the dealation act was altered or did
not go to completion. A queen might lift her hind legs, but not set them on
the wings. One queen with a wing spread at a right angle, raised her hind
leg over the other wing which was folded straight back. Occasionally, queens
placed only one rear leg on top of the abdomen or wings. Another queen
was observed simply walking, raising her gaster, and spreading the wings
slightly. Infrequently a foreleg was used to dislodge wings, either by hooking
it under the wing junction or moving it back and forth over the point of
articulation.
VOLUME 91, NUMBER 3
261
Fig. 1. Uropidine mite on leg of Mynnecocystus mexicanus worker.
Queen grooming. Although grooming between virgin queens was not ob-
served, both virgin and fertilized queens groomed themselves. They clean
their antennae by pulling them through foreleg spurs and draw their front
legs between mouthparts. One queen bent her abdomen forward between
her legs to clean the tip.
Mite infestation. Although mites were not observed on solitary queens or
fertilized queens, they were in four three-queen colonies. They are probably
the same Uropidine mites (Fig. 1) found on honey ant workers. Mites were
first noted in two nests seven days after queens were placed in them. Six
mites was the maximum number observed in a nest at one time. The greatest
number on a queen was five; two on each middle leg and one on a hind leg.
The most on a single leg was three on a middle leg. The greatest number on
a foreleg or hindleg was two.
Usually mites were on middle legs, less frequently on rear legs, and least
frequently on forelegs. An exception occurred in one nest where the only
mite was usually on the hind leg. Another unusual location was the dorsal
thorax near the meso-metanotal suture.
Mites are mobile and may move about on the queen, wander onto the
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NEW YORK ENTOMOLOGICAL SOCIETY
floor, or transfer to a different queen. On the other hand, they may remain
in the same position for some time. A mite was on a hind leg for nine days,
and another stayed on a middle leg for ten days. One mite was on the dorsal
thorax for seventy days. It could not be determined how long mites live,
but in one nest the single mite was present 138 days and in another nest,
the mite was there 1 1 1 days.
There was no conclusive evidence that any queen perished due to mite
infestation, but it may have been a contributing factor. However, it is known
that some queens were parasitized more heavily and that two passed away
with mites still attached.
Queen death. In many cases there was no forewarning of a queen’s death,
but in other instances there was some portent, such as the queen moving
slowly or disgorging fluid. This behavior was noted 10 to 16 days before
demise in virgin queens and 1 to 7 days before in fertilized queens (Conway,
1981a). Sixteen virgin queens died with their abdomens not swollen, or tele-
scoped somewhat. Two had slightly dilated abdomens and four died with
enlarged abdomens.
ACKNOWLEDGMENTS
I am indebted to R. E. Gregg for consultation on results and guidance during the investigation.
Mites were identified by John Kethley, Division of Insects, Chicago Field Museum of Natural
History. Grants from the Walker Van Riper Fund of the University of Colorado Museum and
from the Kathy Lichty Memorial Fund of the University of Colorado Biology Department
provided financial support for the study.
LITERATURE CITED
Cazier, M. A. and M. A. Mortenson. 1965. Bionomical observations on myrmecophilous
beetles of the genus Cremastocheiliis. J. Kansas Entomol. Soc. 38:19-44.
Cazier, M. A. and M. Statham. 1962. The behavior and habits of the myrmecophilous scarab
Cremastocheilus stathamae Cazier with notes on other species (Coleoptera: Scarabae-
idae). J. New York Entomol. Soc. 70:125-149.
Chew, R. M. 1979. Mammalian predation on honey ants, Mynnecocystus (Formicidae).
Southwestern Nat. 24:677-682.
Conway, J. R. 1975. Selected aspects of the biology of the honey ant, Mynnecocystus mexi-
canus subsp. hortideorum McCook, in the vicinity of Colorado Springs, Colorado. Ph.D.
thesis. University of Colorado, Boulder, 325 pp.
Conway, J. R. 1977. Analysis of clear and dark amber repletes of the honey ant, Mynnecocystus
mexicanus hortideorum. Annals of the Entomological Society of America 70:367-369.
Conway, J. R. 1980a. Foraging behavior of the honey ant, Mynnecocystus mexicanus, in
Colorado. Trans. Illinois State Acad. Sci. 72:81-93.
Conway, J. R. 1980b. The seasonal occurrence of sexual brood and the pre- and post-nuptial
behavior of the honey ant, Mynnecocystus mexicanus Wesmael, in Colorado. J. New
York Entomol. Soc. 88:7-14.
Conway, J. R. 1981a. A study of dealated queens of the honey ant, Mynnecocystus mexicanus
Wesmael, and their progeny in captivity (Hymenoptera: Formicidae). J. Kansas Entomol.
Soc. 54:41-55.
VOLUME 91, NUMBER 3
263
Conway, J. R. 1981b. Honey ants: sweet swell of success. Science Digest 89:56-59, 1 17.
Conway, J. R. In press. Nest architecture and population of the honey ant, Myrmecocystus
mexicanus Wesmael (Formicidae) in Colorado. Southwestern Nat.
Creighton, W. S. 1950. The ants of North America. Bull. Mus. Comp. Zool. 104:1-585.
Gregg, R. E. 1963. The Ants of Colorado. University of Colorado Press, Boulder, 792 pp.
Hdlldobler, B. 1976. Tournaments and slavery in a desert ant. Science 192:912-914.
Hdlldobler, B. and C. J. Lumsden. 1980. Territorial strategies in ants. Science 210:732-739.
Kay, C. A. 1978. Preferred temperatures of desert honey ants (Hymenoptera: Formicidae).
J. Thermal Biol. 3:213-217.
Kay, C. A. and W. G. Whitford. 1978. Critical thermal limits of desert honey ants: possible
ecological implications. Physiol. Zool. 51:206-213.
McCook, H. C. 1882. The Honey Ants of the Garden of the Gods, and the Occident Ants of
the American Plains. J. B. Lippincott & Co., Philadelphia, 188 pp.
Slocumb, L. B. 1966. A report on the honey ants of the plains. M.A.T. thesis, Colorado
College, 31 pp.
Snelling, R. R. 1976. A revision of the honey ants, genus Myrmecocystus (Hymenoptera:
Formicidae). Nat. Hist. Mus. Los Angeles County Sci. Bull. 24:163 pp.
Wheeler, W. M. 1908. Honey ants, with a revision of the American Myrmecocysti. Bull.
Amer. Mus. Nat. Hist. 24:345-397.
Wheeler, W. M. 1910. Ants, Their Structure, Development and Behavior. Columbia Uni-
versity Press, New York, 663 pp.
Wheeler, W. M. 1913. Additions to our knowledge of the ants of the genus Myrmecocystus
Wesmael. Psyche 19:172-181.
Wheeler, W. M. 1917. The pleometrosis of Myrmecocystus. Psyche 24:180-182.
Received September 1, 1982; accepted December 28, 1982.
NEW YORK ENTOMOLOGICAL SOCIETY
91(3), 1983, pp. 264-268
NESTING BIOLOGY OF THE BEE SV ASTRA SABINENSIS
(HYMENOPTERA, ANTHOPHORIDAE)
Jerome G. Rozen, Jr.
Deputy Director for Research and Curator,
American Museum of Natural History,
New York, New York 10024
Abstract. — \ single nest of Svastra sabinensis sabinensis (Cockerell) is described, illustrated,
photographed, and compared with the nests of Svastra obliqua (Say), the only other member
of the genus whose nest has been described. Svastra sabinensis sabinensis is ground nesting,
with a branched, meandering burrow descending to approximately 80 cm, at which level nu-
merous cells occur. Cells are vertical, elongate, and lined with a non-waxlike waterproof lining.
Unlike in any other anthophorid bee, parts of the nest are plugged with sorted pebbles, pre-
sumably by the females associated with this nest, and a single specimen of the cuckoo bee
Triepeolus penicilliferus (Brues) was collected while trying to enter the nest. This is the first
association of T. penicilliferus with any host.
While searching for nests of various bees at Corona de Tucson, a small
development about twenty miles southeast of Tucson, Pima County, Ari-
zona, I discovered and excavated a nest of Svastra sabinensis sabinensis
(Cockerell) on August 27 and 28, 1982. Dr. Wallace E. LaBerge, who kindly
identified two females associated with the nest, suggested that my fragmen-
tary observations would be of interest because so little has been published
on the nests of any species in the genus. Only Rau (1922), Custer (1928,
1929) and Rozen (1964) have contributed nesting information heretofore,
all authors reporting on Svastra obliqua (Say). Like that species, Svastra
sabinensis belongs to the subgenus Epimelissodes, but its range is much more
limited, and the nominate subspecies is known only from Arizona and New
Mexico (LaBerge, 1956).
The single isolated nest penetrated the nearly horizontal ground (Fig. 1)
where there was no shade, although Cercidium, Echinocactus, several species
of Opuntia, and other typical Sonoran desert plants were abundant. The
entrance was open, without turret, and not situated near any prominent
marker such as a stone, and it was not surrounded by a tumulus. The main
shaft (Fig. 3), circular, about 6 mm in diameter, unlined by any secretion.
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VOLUME 91, NUMBER 3
265
Figs. 1,2. 1. Nesting locale of Svastra sabinensis sabinensis at Corona de Tucson, showing
characteristic vegetation. 2. Section of nest of Svastra sabinensis sabinensis showing pebbles
blocking curved branch, side view. Arrow marks point of attachment of main descending branch
with pebble filled one. White material in burrow above is dry, powdered plaster of paris, used
by the author to follow burrow during excavation.
and without masoned walls, descended at about a 30° angle from the surface
for about 8 cm, at which depth a short branch was attached, presumably an
antechamber as described by Custer (1928) and Rozen (1964) for Svastra
obliqua. Below the antechamber the main tunnel turned downward, with
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NEW YORK ENTOMOLOGICAL SOCIETY
Figs. 3, 4. 3. Schematic diagram of nest of Svastra sabinensis sabinensis, side view. 4.
Diagram of cell lumen of same, made from plaster cast using camera lucida to depict exact
shape, side view.
VOLUME 91, NUMBER 3
267
some meandering. At about the 25 cm level, a side tunnel branched, which
was hlled with rather large, sorted pebbles (Fig. 3). The side tunnel was lost
and may have been filled with soil beyond the pebbles. Not having seen
such a plug of pebbles in any anthophorid nest before, I postulated that a
wasp might have constructed the tunnel using the Svastra burrow as an
access tunnel. The open burrow continued to descend to about the 40 cm
level, at which point it branched. One branch curved and was filled for about
4 cm, again with sorted pebbles (Fig. 2); then it was open for about 6 cm
and then again plugged with pebbles for 3 cm more, after which it appeared
to end blindly. The occurrence of pebble plugs at various places in this nest
almost certainly indicated that the female bees occupying the nest had col-
lected and deposited them, but their function, if any, is not clear. The other
branch, completely open, descended, still meandering, to the approximate
depth of 75 cm, where it branched and then branched again at 80 cm. One
of the last branches, extending obliquely downward, ended in a vertical cell
without rising before the cell entrance. In this vicinity four other cells were
found near one another, all presumably arranged singly. Side tunnels leading
to them were filled with soil and not discernible from the nesting substrate.
Although I had hoped to find a cell arrangement similar to that diagrammed
by Custer (1928) for Svastra obliqua or by Rozen (1974) for Thygater analis
(Lepeletier), no such distinctive patterns could be identified.
The cells (Fig. 4) were elongate, with a broadly rounded bottom and a
distinct, deeply concave closure. This closure, a well formed spiral with
approximately five coils to the radius, possessed a rather poorly formed
inner coil. This coil appeared to be less consolidated than that of the other
rows. The plane of two such closures was tilted so that it was not at right
angles to the long axis of the cell (not visible in profile drawn in Fig. 4).
Unlike that of Svastra obliqua (Rozen, 1964), the cell wall did not appear
to be plastered, but was slightly harder than the surrounding soil, suggesting
that it had been either worked by the female or impregnated by her. It was
very smooth and coated with a thin, somewhat shiny, transparent, non-
waxlike material that was tan and translucent when viewed against a plaster
cast made of the cell lumen. This lining extended to the cell closure, but not
beyond, where the wall of the side tunnel was rough, not shiny, and readily
absorbed water droplets.
Cells were 8.0 to 8.2 mm in maximum diameter (3 measurements) and
the cast of one cell from apex of closure to bottom was 19 mm. The cell
diameter at the closure was 6.5 mm (2 measurements).
A number of feeding immature larvae were recovered but none had reached
the last instar. The provisions were orange, homogeneously moist but not
liquid. As in Svastra obliqua, more than one female occupied a single nest.
Two badly worn females were recovered from the nest, and a single Tri-
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NEW YORK ENTOMOLOGICAL SOCIETY
epeolus penicilliferus (Brues)^ was collected while it was trying to enter the
nest, although no eggs or larvae of the cuckoo bee were recovered. This is
the hrst host record for T. penicilliferus.
Samples of cells, and the two female specimens, as well as the Triepeolus
penicilliferus pinned with one of the females, are in the collections of the
American Museum of Natural History.
LITERATURE CITED
Custer, C. P. 1928. On the nesting habits of Melissodes Latr. (Hymenoptera). Canad. Ent. 60:
28-31.
Custer, C. P. 1 929. Notes on cocoons and parasites of Melissodes obliqua and nests of Perdita
opuntiae (Hymenoptera-Apoidea). Psyche 36:293-295.
LaBerge, W. E. 1956. A revision of the bees of the genus Melissodes in North and Central
America. Part I (Hymenoptera, Apidae). Univ. Kans. Sci. Bull. 37(18):91 1-1 194, hgs.
1-129.
Rau, P. 1922. Ecological and behavior notes on Missouri insects. Trans. Acad. Sci. St. Louis
24(7): 1-71, figs. 1-12.
Rozen, J. G., Jr. 1964. The biology of Svastra obliqua obliqua (Say), with a taxonomic
description of its larvae (Apoidea, Anthophoridae). Amer. Mus. Novitates 2170:13 pp.,
figs. 1-15.
Rozen, J. G., Jr. 1974. Nest biology of the Eucerine Bee Thygater analis (Hymenoptera,
Anthophoridae). J. New York Ent. Soc. 82:230-234, figs. 1-2.
Received April 10, 1983; accepted May 10, 1983.
* Compared with specimens from the vicinity of Tucson, Arizona, identified by Dr. Paul D.
Hurd, Jr.
NEW YORK ENTOMOLOGICAL SOCIETY
91(3), 1983, pp. 269-272
RESPONSE OF A GOLDENROD BEETLE TO FOUR
SELDOM-ENCOUNTERED GOLDENROD
(SOLIDAGO) SPECIES
Frank J. Messina*
Section of Ecology & Systematics, Cornell University,
Ithaca, New York 14853
Thegoldenrods Solidago bicolor and S. nemoralis are scarce in the old-field habitat
of a goldenrod leaf beetle, Trirhabda virgata LeConte, in central New York. Two additional
species, S. caesia and S. flexicaulis, are wholly restricted to woodlands. In the laboratory only
S. bicolor supported complete development and reproduction by T. virgata, but this plant was
an inferior host compared to five common meadow goldenrods that had been tested previously.
Nine Solidago spp. can be ranked in four categories of suitability for T. virgata. Host quality
is not well correlated with taxonomic affinities of the plants.
More than 1 5 species of goldenrod (Asteraceae: Solidago) occur in central
New York (Clausen, 1949). These rhizomatous perennials typically grow in
natural openings (e.g., rocky hillsides) and in disturbed areas (e.g., abandoned
fields). In a recent paper I described (Messina, 1982a) how hve common
goldenrods vary in quality for a leaf beetle, Trirhabda virgata LeConte (Co-
leoptera: Chrysomelidae). To provide a broader perspective on the Tri-
rhabda-Solidago association, I report here the performance of T. virgata
when offered four Solidago species that the beetle rarely encounters in nature.
Solidago nernoralis and S. bicolor are least common among seven gold-
enrods in old helds of central New York, perhaps because they prefer dry,
gravelly soils (Wiegand and Fames, 1925; Sinclear, 1979). Although S. bi-
color is rare in open meadows where Trirhabda occurs, it is common along
woodland edges. Two additional goldenrods are restricted to shaded habitats.
Solidago caesia is characteristic of dry woods, whereas S. flexicaulis is com-
mon in moist, well-shaded woods (Wiegand and Fames, 1925). Like most
chrysomelids, Trirhabda beetles fail to disperse into shaded areas. I therefore
tested whether S. caesia, S. flexicaulis, and possibly S. bicolor escaped her-
bivory by T. virgata because of their affinity for woodlands. The life cycle
’ Present address: Boyce Thompson Institute for Plant Research, Tower Road, Ithaca, New
York 14853.
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270
NEW YORK ENTOMOLOGICAL SOCIETY
Table 1. Comparison of the performance of T. virgata larvae and adults on S. canadensis'
and on 4 ‘novel’ goldenrods.
Solidago host
canadensis
bicolor
caesia
flexicaulis
nemoralis
N
Larval
survivorship^
0.45
0.30
0
0
0
40
Developmental
endpoint
adult
adult
instar III
instar III
instar I
Adult longevity^
{x ± SE days)
31 ± 2
20 ± 2
7 ± 1
6 ± 1
5 ± 1
35
‘ Var. scahra = S. a/tissinia (see Messina, 1982a).
2 Fraction surviving from egg hatch to adult emergence.
^ All adults shared S. canadensis as the larval host.
of T. virgata is described elsewhere (Messina, 1982b). Larvae and adults
feed on goldenrod leaves and can reach densities that result in complete
defoliation in open fields.
METHODS
Trirhabda larvae were reared from egg hatch to adult emergence on each
of the four goldenrods in a room-size growth chamber (24 ± 2°C, 16L:8D).
Cohorts of 40 larvae were placed on each host. Every other day I provided
fresh-cut goldenrod stems which were placed in florists’ water-pics to main-
tain plant turgor. I measured variation in plant quality for adults by using
newly emerged adults that had fed as larvae on S. canadensis, an optimal
larval host. Thirty-five adults were used in each host treatment, and fresh-
cut stems of the appropriate host again were provided every other day. I
used plants grown outdoors to ensure that larvae and adults received stems
in the proper phenological stage. Further details are found in Messina (1 982a).
RESULTS AND DISCUSSION
The performance of beetles fed the ‘novel’ goldenrods can be compared
to the performance of beetles reared simultaneously on S. canadensis, the
most common old-field host. Of the four test hosts, only S. /7/co/cr supported
complete larval development; survivorship was similar to that on S. can-
adensis (Table 1). However, larvae fed S. bicolor required on average 38.1
d (N = 1 2) to develop into adults, whereas larvae fed S. canadensis required
only 3 1 .3 d (N = 1 8, F < 0.01, Mann-Whitney test). Solidago bicolor af-
forded slower growth than did any of the five common old-field hosts (Mes-
sina, 1982a).
All first-instar larvae fed S. nemoralis died within 5 d; this mortality is
VOLUME 91, NUMBER 3
271
Table 2. Relationship between the field preference of T. virgata and the suitability of nine
Solidago spp.
Solidago sp.
Host use in the field
Host suitability'
larvae
adults
bicolor
not utilized
+
+
caesia
not utilized
-
-
canadensis
frequent
+ +
+ +
flexicaulis
not utilized
-
-
gigantea
frequent
+ +
+ +
grarninifolia
rare
+
+ +
juncea
rare
+
+ +
nemoralis
not utilized
-
-
rugosa
frequent
+ +
+ +
‘ ++ = optimal host, + = inferior but adequate for survivorship or reproduction, — = un-
suitable.
comparable to that of starved individuals (Messina, 1982b). In contrast,
several larvae fed S. caesia or S. flexicaulis molted to the second stadium,
and three larvae in each treatment progressed to the third (last) stadium
before dying. One larva fed S. caesia lived for 32 d, a period of time that
would be sufficient for complete development on a normal goldenrod host.
The four goldenrod species were ranked similarly in food quality for adults.
Most of the 105 adult beetles fed S. caesia, S. flexicaulis, or S. nemoralis
died within 10 d (Table 1). No females in these treatments became gravid,
even though one female fed S. caesia lived for 42 d. As it was for larvae,
S. bicolor was intermediate in quality for adults. The longevity of beetles
fed S. bicolor was significantly lower than the longevity of beetles fed S.
canadensis (Table 1), but several females survived long enough on S. bicolor
to produce fertile eggs (hatching success = 83%, N = 58).
The rare old-field goldenrods strongly diverged in suitability. Solidago
bicolor was a suboptimal but adequate host, whereas S. nemoralis was the
least suitable goldenrod tested. Solidago nemoralis produces very little leaf
biomass/stem; if the plant were suited to T. virgata it would be rapidly
defoliated. Observations suggest that S. nemoralis leaves may be especially
well defended against many goldenrod herbivores.
The relationship between T. virgata and nine Solidago spp. is presented
in Table 2. The plants constitute four categories of suitability. Solidago
canadensis, S. gigantea, and S. rugosa are frequently utilized, optimal hosts.
Solidago grarninifolia and S. juncea, which afford slower development of
larvae, are rarely used by either larvae or adults, and are colonized only
when they are interspersed among preferred hosts (Messina, 1 982a). Solidago
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NEW YORK ENTOMOLOGICAL SOCIETY
bicolor is an inferior food plant for both stages. Finally, the woodland gold-
enrods and S. nemoralis are unsuitable for larvae and adults.
A plant can escape herbivory by growing in sites that are not invaded by
insects otherwise adapted to feed on it. For example, the woodland crucifer
Dentaria diphylla is suitable for many crucifer-feeding insects that restrict
host search to open areas (Cromartie, 1975). I did not hnd an analogous
relationship between T. virgata and the woodland goldenrods, which are
protected by both extrinsic (habitat) and intrinsic (plant quality) barriers to
colonization.
Only a subset of Solidago species are thus potential hosts for T. virgata.
Plant quality does not seem to follow taxonomic lines within Solidago.
Solidago graminifolia, which is occasionally placed in the separate genus
Euthamia (Cronquist, 1980), is distantly related to all the other goldenrods
but is more suitable than any of the four rarely encountered goldenrods.
Moreover, T. virgata successfully develops and reproduces on certain asters,
such as Aster novae- anglicae (unpubl. data). It remains to be determined
which morphological or chemical characteristics are shared by host gold-
enrods and asters but are lacking in non-host species in each genus.
LITERATURE CITED
Clausen, R. T. 1949. Checklist of the vascular plants of the Cayuga Quadrangle. Cornell
Univ. Agric. Exp. Sta. Mem. 291, 87 pp.
Cromartie, W. J., Jr. 1975. Influence of habitat on colonization of collard plants by Pieris
rapae. Environ. Entomol. 4:783-784.
Cronquist, A. 1980. Vascular Flora of the Southeastern United States. Vol. 1. Asteraceae.
Univ. North Carolina Press, Chapel Hill, 261 pp.
Messina, F. J. 1982a. Food plant choices of two goldenrod beetles: relation to plant quality.
Oecologia 55:342-354.
Messina, F. J. 1982b. Comparative biology of the goldenrod leaf beetles, Trirhabda virgata
and T. borealis (Coleoptera: Chrysomelidae). Coleop. Bull. 36:255-269.
Sinclear, J. C. 1979. Environmental factors that influence the distribution of goldenrods in
central New York. M.S. thesis, Cornell Univ., Ithaca, New York, 42 pp.
Wiegand, K. M. and A. J. Eames. 1925. The flora of the Cayuga Lake Basin, New York.
Cornell Univ. Agric. Exp. Sta. Mem. 92, 491 pp.
Received November 15, 1982; accepted April 19, 1983.
NEW YORK ENTOMOLOGICAL SOCIETY
91(3), 1983, pp. 273-279
PRECOPULATORY BEHAVIOR OF THE WHIRLIGIG BEETLE
DINEUTES DISCOLOR (COLEOPTERA:GYRINIDAE)
Steven A. Kolmes
Department of Zoology, Birge Hall, University of Wisconsin,
Madison, Wisconsin 53706
Abstract. — Tht sexual behavior of whirligig beetles is more complex than has previously been
described. Precopulatory mounting is a behavior pattern performed by males, and proleg-up
is performed by females. These interactions are more in accord with the potential complexities
of mate selection and sexual synchronization than earlier accounts of whirligig courtship be-
havior.
Precopulatory communication is believed to be important in allowing
animals to select mates of the appropriate species and of robust phenotype,
and in synchronization of sexual behavior. It is of special interest therefore,
when the accounts of a species’ reproductive behavior seems to preclude
any interactions complex enough to facilitate these processes. The only de-
scriptions of whirligig beetle precopulatory behavior speak of males merely
leaping on females and inseminating them (Smith, 1926), which in light of
current ideas about sexual communication seems unlikely. I undertook this
study in order to uncover any behavior interactions in gyrinid precopulatory
behavior that might supply a context in which precopulatory communication
of a more substantial sort might occur.
These beetles have four compound eyes, have good vision (Bennett, 1967;
Carthy and Goodman, 1 964) and appear to be very visually oriented (Brown
and Hatch, 1 929). As surface film dwelling predators, whirligigs have evolved
antennal specializations that allow their Johnston’s organs to serve as ex-
tremely sensitive surface vibration detectors (Wilde, 1941; Rudolph, 1967).
Other analogous surface vibration detectors occur in insects found at the
air-water interface (Murphey, 1971, 1973; Murphey and Mendenhall, 1973)
and in some cases precopulatory signals are transmitted via surface film
vibrations (Wilcox, 1972, 1979).
The basic gyrinid body plan consists of streamlined elytra covering the
thorax and abdomen, with flat paddle-like mesothoracic and metathoracic
legs that are efficient swimming organs (Nachtigall, 1961). The long, slender
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NEW YORK ENTOMOLOGICAL SOCIETY
Fig. 1 . A view of Dineutes discolor swimming. The prothoracic leg is shown lowered some-
what so that its form is apparent.
prothoracic legs are used to seize prey and are held under the beetles’ bodies
in recessed grooves when unused (Fig. 1 shows these legs lowered somewhat).
MATERIALS AND METHODS
Dineutes discolor were removed from a swarm located on the Wisconsin
River near Arena (Iowa Co.), Wisconsin in September 1977 and throughout
May and June 1978. The beetles were housed in screen-eovered aquaria
filled with tap water and fed live flightless Drosophila. Approximately 70
beetles individually marked on their elytra with Tester’s PLA enamel paints
were used in my observations.
The observation tank was a 92.5 X 46.5 X 46.5 cm aquarium containing
floating vegetation and live prey. The beetles were therefore free to climb
out of the water or feed, rather than being constrained to interaeting in a
more impoverished environment. Observations were normally for 55 min
sessions, occasionally 44 min.
The rates at whieh every pair of beetles performed different behavior
patterns were recorded and Mann-Whitney U tests (Siegel, 1956) were used
to eompare the behavior patterns performed by beetles whieh mated to the
data collected during observation periods during which the beetles did not
mate. It is important to note that all data from subjects that were never
observed to mate were disearded, so that nonmating data were collected
Fig. 2. A proleg-up performed by a female beetle.
VOLUME 91, NUMBER 3
275
FEMALE PROLEG-UP PERFORMANCE
• = Mating females, n = 9
A= Nonmating females, n=15
COPULATION
1.5
1.2
0.9
0.6
0.3
• • • •
• •
A A A A
• • •
A A A A A A A
• • •
A A A A
A A A-A-^A-^A-^V-A-A-A-nA^-Ar-J-
10 11 12 13 14
MINUTES
Fig. 3. Performance of the proleg-up behavior by female beetles, in observations with one
pair of beetles present. Values for control females are based on the hrst 14 min of each obser-
vation of nonmating pairs (which was the average precopulatory interval in mating pairs).
from animals known to be capable of carrying out successful precopulatory
sequences at other times.
RESULTS
I conducted 56 observation periods, during which 24 copulations were
observed. Twenty-four observation periods had one pair of beetles present
in the experimental apparatus, and 32 of the observations had two pairs
present. Data collected in these two situations were consistent with one
another in form and in statistical significance, and they are reported together.
Data on fifteen behavior patterns were collected. These categories included:
different postures of legs and elytra, mouthpart movements, proximity and
contacts between beetles, diving, swimming, climbing, general activity level,
feeding, and grooming. Of these behavior patterns, only two proved to be
statistically related to successful copulation.
Female beetles perform a behavior pattern called proleg- up more fre-
quently before they copulate (Mann-Whitney U test, P < 0.05). This act
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NEW YORK ENTOMOLOGICAL SOCIETY
MALE MOUNTINGS OF FEMALE BEETLES
COPULATION
Z)
o
2 5
ii_
o
cr
uj 4
OQ
ZD
• = Mating males, n = 9
A= Nonmating males, n=15
• •
A A
• •
A A A A A A A
• A A A A
^^^AAAAAA
J ^ ^ ^ ^ I I I ^ I ^ ^ L
-A-a-^
1 2 3 4 5 6 7 8 9 10 11 12 13 14
MINUTES
Fig. 4. Male mountings of female beetles, in observations with one pair of beetles present.
Values for control males are based on the first 14 min of each observation of nonmating pairs
(which was the average precopulatory interval in mating pairs).
consists of the beetle swinging her leg out toward the lateral margin of her
body, flexing the leg so that its distal region (tibia and tarsus) extends per-
pendicularly up from the water, and then rapidly reversing these movements
(Fig. 2). Male performance of proleg-up is not statistically related to ensuing
copulation.
The data in Figure 3 show the average number of proleg-up performances
plotted against time to eopulation, for female beetles observed with one male
present. The proleg-up behavior has reached almost its final level when
copulation is still 5 min away. It appears therefore to be a signal used early
in precopulatory communication, rather than in the final stages. No specific
distance or direction between male and female during proleg-up performance
was noted, but this may have been an artifact of their relatively enforced
proximity in the observation aquarium.
Male beetles mount female beetles more frequently before they copulate
(Mann-Whitney U test, P < 0.05), with the males elinging to the dorsal
surface of the females for periods ranging from very brief to 10 min. These
mounts do not involve insertion of the male genitalia (those were scored
VOLUME 91, NUMBER 3
111
Fig. 5. Male (above) and female (below) tarsal segments of the prothoracic leg.
separately as copulations). Interestingly although male/male, male/female,
and female/male mounts were all common, only male/female mounts are
especially associated with successful precopulatory sequences.
The data in Figure 4 show the average number of male mountings of
females, plotted as in Figure 3. These mountings continue throughout the
precopulatory exchange, increasing in rate somewhat in the hnal 2 min before
copulation. Beetles are adjacent to one another immediately before a mount-
ing, but no specific direction of approach by the male relative to the female
was apparent.
DISCUSSION
Based on statistical analysis of rates of different precopulatory behavior
patterns, mountings of females by males and proleg-up by females may be
serving in the precopulatory communication of Dineutes discolor.
The proleg-up behavior pattern exposes the distal segment of the pro-
thoracic leg as high above the water line as is possible. This leg region is the
animal’s one strikingly sexually dimorphic characteristic (Fig. 5) aside from
the form of the genitalia themselves. This behavior pattern may therefore
serve to demonstrate sexual identity and indicate willingness to copulate.
The occurrence of males mounting females as a precopulatory display is
known in a wide variety of animal species. These mountings in D. discolor
are best viewed as signals rather than as unsuccessful attempts to copulate.
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NEW YORK ENTOMOLOGICAL SOCIETY
especially in light of the facts that (a) females mount males, (b) males mount
other males and (c) the male genitalia are not extended during any of these
mountings.
The sensory channels that might be involved in the proposed displays of
D. discolor are diverse, but visual transmission seems likely for the proleg-
up and tactile transmission seems likely for male/female mountings. In the
latter, ample tactile cues are certainly present — although contact chemore-
ception cannot be ruled out. In the former, the striking sexual dimorphism
of the distal prothoracic leg segments could be easily apparent via visual
cues— but again, due to the remarkable sensitivity of the gyrinid surface
vibration detector (Rudolph, 1967) a second stimulus channel cannot be
ruled out.
The proleg-up performance by female beetles reaches a plateau early in
precopulatory exchanges, and is more likely to be involved in initially sig-
nalling for female receptivity than in facilitating the final approach of the
male for copulation. Male mountings of females occur throughout the pre-
copulatory period, and increase somewhat in frequency during the last 2
min before copulation. This behavior may be involved both in signalling
sexual readiness, and in facilitating the final synchronization and orientation
of the beetles for copulation.
This type of observational study is inherently limited in its conclusion.
In order to further elucidate the signal value of these behavior patterns,
either their performance or perception would have to be experimentally
manipulated and the results observed. Nonetheless, this study is adequate
to strongly indicate that whirligig precopulatory behavior contains interac-
tions that could supply the reciprocal complexity we have come to expect
in situations where mate selection or sexual synchronization occur.
ACKNOWLEDGMENTS
I greatly appreciate the aid of Professors Jeffrey R. Baylis and Jack P. Hailman in the course
and completion of this study. Among others, Laurence Berg, Clyde Gorsuch, Leigh Hurst,
Katherine C. Noonan, and Scott R. Robinson have given me encouragement and advice. Cheryle
Hughes produced the excellent illustrations. This work was supported in part by Fellowships
from the Graduate School of the University of Wisconsin.
LITERATURE CITED
Bennett, R. R. 1967. Spectral sensitivity studies on the whirligig beetle, Dineutes ciliatus. J.
Insect. Physiol. 13:621-633.
Brown, C. R. and M. H. Hatch. 1929. Orientation and “fright” reactions of whirligig beetles
(Gyrinidae). J. Comp. Psychol. 9:159-189.
Carthy, J. D. and L. J. Goodman. 1964. An electrophysiological investigation of the divided
eye of Gyrinus bicolor. J. Insect Physiol. 10:431-436.
VOLUME 91, NUMBER 3
279
Murphey, R. K. 1971. Sensory aspects of the control of orientation to prey by the waterstrider,
Gerris remigis. Z. Verg. Physiol. 72:168-185.
Murphey, R. K. 1973. Mutual inhibition and the organization of a non- visual orientation in
Notonecta. J. Comp. Physiol. 84:31-40.
Murphey, R. K. and B. Mendenhall. 1973. Localization of receptors controlling orientation
to prey by the Back Swimmer Notonecta undulata. J. Comp. Physiol. 84:19-30.
Nachtigall, W. 1961. Funktionelle Morphologic, Kinematik, und Hydromechanik des Ru-
derapparates von Gyrinus. Z. Vergl. Physiol. 45:193-226.
Rudolph, P. 1967. Zum ortungsverfahren von Gyrinus substriatus (Steph). Z. Vergl. Physiol.
56:341-375.
Siegel, S. 1956. Nonparametric Statistics for the Behavioral Sciences. McGraw Hill Book
Company, New York.
Smith, H. B. 1926. Notes on the behavior of Dineutes americanus. Psyche 33:156-161.
Wilcox, R. S. 1972. Communication by surface waves in the mating behavior of a water
strider (Gerridae). J. Comp. Physiol. 80:255-266.
Wilcox, R. S. 1979. Sex discrimination in Gerris remigis: role of a surface wave signal. Science
206:1325-1327.
Wilde, J. de 1941. Contribution to the physiology of the Johnston organ and its part in the
behavior of the Gyrinus. Arch. Neer. Physiol. Homme Animaux 25:381-400.
Received October 13, 1982; accepted April 19, 1983.
NEW YORK ENTOMOLOGICAL SOCIETY
91(3), 1983, pp. 280-282
SEM STUDY OF THE ANTENNAL SENSILLA AND SETAE
OF SOLVA PALLIPES (LOEW) (DIPTERA: XYLOMYIDAE)
C. E. Vasey and E. Ritter
Department of Biology, State University of New York,
College of Arts and Science, Geneseo, New York 14454
Abstract.— T\\q antennal sensilla and setae of both sexes of Solva palHpes are described using
SEM. Sexual dimorphism is evident on the first flagellomere and its functional significance is
postulated.
Light microscope observations of antenna of xylomyids were made by
Leonard (1930) and Steyskal (1947) with the latter using antennal charac-
teristics to help separate Solva from Xylomya. Steyskal indicated Solva
pallipes (Loew) is widely distributed throughout the Nearctic region. How-
ever, much of its biology is poorly known and electron microscopic inves-
tigations have not been undertaken.
Other investigators have clearly demonstrated the importance of insect
antennal sensilla in perceiving a variety of external stimuli (Acree et al.,
1968; Kellogg, 1970; Mclver, 1974; Chu-Wang et al., 1975; Zacharuk, 1980).
It is anticipated that these SEM observations will be utilized to better un-
derstand the structural components of the antennal sensilla and to provide
some evidence for their functional role in the behavior of this species.
MATERIALS AND METHODS
Dried specimens of both sexes of Solva pallipes were mounted on alu-
minum studs with silver paint and gold coated in Polaron diode sputterer.
Rehydrated specimens were similarly treated and showed less distinctly the
sensillar pattern. Critical point drying was, therefore, not deemed desirable.
Specimens were examined in an ISI Alpha-9 SEM.
RESULTS AND DISCUSSION
The antennae of xylomyid flies consists of a scape attached to the head,
followed by a pedicel and a segmented flagellum. Leonard (1930) indicated
The publication costs of this article were defrayed in part by page charge payment. This article
must therefore be hereby marked ‘Advertisement’' \n accordance with 18 U.S.C. §1734 solely
to indicate this fact.
VOLUME 91, NUMBER 3
281
Figs. 1-6. 1. Antenna of female Solva pallipes showing scape (a) and pedicel Arrows
indicate flagellar annuli. lOOX. 2. Flagellar segments demonstrating typical fluted tricoid sensilla.
Arrow indicates pattern of depressions near flagellar junction. lOOX. 3. Basal segment of male
flagellum with typical coeloconicoid sensilla. 2,000X. 4. Depressions of flagellar segments show-
ing two types of sensilla and cuticular wall surrounding each wall. 1,000X. 5. Scape (a) of
antenna showing antennal socket (arrow) and pattern of tricoid sensilla. 700X. 6. Lateral view
of scape (a) and pedicel (13) showing absence of sensilla on dorsum. 400X.
seven flagellar segments while Steyskal (1947) reported eight for both sexes.
The latter number is supported by our studies (Fig. 1).
The flagellum contains a variety of trichoid sensilla and setae having the
same pattern of distribution on all segments and being the most frequently
encountered type in both sexes. These have evident fluting and are not
socketed (Fig. 2). In males, the basal segment bears a large number of coe-
loconic sensilla which are surrounded by fluted setae (Fig. 3) not found on
the female antennae. The basal region of the female antennae bears a uniform
pattern of basiconic sensilla. Moreover, the basiconic sensilla are found in
both sexes from annulus two through seven, and are always located in “well-
like” depressions surrounded by the fluted setae which appear to be elevated
on a cuticular wall around the well (Figs. 2, 4). Figure 4 shows the distinct
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NEW YORK ENTOMOLOGICAL SOCIETY
arrangement of five to six basiconicoid sensilla characteristic for a given
well, and four to five unfluted basiconicoid sensilla surrounding a shorter
fluted sensillum. The “well-like” areas decrease in number as one proceeds
distally and were not found on the eighth flagellomere.
The scape arises from an antennal socket and bears large, fluted setae
which arise from sockets on the ventral and lateral surfaces. These setae do
not occur on the lower one-third of the scape (Fig. 5). This pattern is seen
on both sexes. Smaller, non-socketed setae cover the entire scape except for
the dorsal surface (Fig. 6). The setal pattern of the upper portion of the scape
continues on to the pedicel.
Although specific evidence is not available from this study, Zacharuk
(1980) has indicated that coeloconic and basiconic sensilla are generally
chemosensory. It is entirely possible, then, that the different sensilla pattern
observed on the first flagellomere of the male might be functional in mating.
ACKNOWLEDGMENT
We thank Dr. F. E. Kurczewski of SUNY College of Environmental Sciences and Forestry
for his advice, suggestions, and reading of the manuscript.
LITERATURE CITED
Acree, F., Jr., R. B. Turner, H. K. Gouck, M. Beroza and N. Smith. 1968. L-lactic acid: a
mosquito attractant isolated from humans. Science 161:1346-1347.
Chu-Wang, I. W., R. C. Axtell and D. L. Kline. 1975. Antennal and palpal sensilla of the
sand fly, Culicoides furens (Poey) (Diptera: Ceratopogonidae). Int. J. Insect Morphol.
Embryol. 4(2): 13 1-149.
Kellogg, F. E. 1970. Water vapor and carbon dioxide receptors in Aedes aegypti (L.). J. Insect
Physiol. 16:99-108.
Leonard, D. M. 1930. A revision of the dipterous family Rhagionidae (Leptidae) in the United
States and Canada. Mem. Am. Entomol. Soc. 7:1-181.
Mclver, S. B. 1974. Fine structure of antennal grooved pegs of the mosquito, Aedes aegypti.
Cell Tiss. Res. 153:327-337.
Steyskal, G. C. 1947. A revision of the Nearctic species of Xylomyia and Solva (Diptera:
Erinnidae). Mich. Acad. Sci. Arts and Letters 31:181-190.
Zacharuk, R. Y. 1980. Ultrastructure and function of insect chemosensilla. Ann. Rev. Ento-
mol. 25:27-47.
Received November 1, 1982; accepted April 19, 1983.
NEW YORK ENTOMOLOGICAL SOCIETY
91(3), 1983, pp. 283-288
BOOK REVIEWS
The Growth of Biological Thought. Diversity, Evolution, and Inheritance.—
Ernst Mayr. 1982. Belknap/Harvard University Press, Cambridge, Mas-
sachusetts, xiii + 974 pp. $30.00.
This is a big book, containing close to a thousand pages of densely packed
facts, interpretations and opinions. It is not light reading, but in general it
is clearly written and unambiguous. After a hrst reading through, I think
many biologists will find it valuable above all as a reference work, albeit
one often as strongly colored by the author’s personal viewpoints as was Dr.
Johnson’s dictionary.
Major themes of the work are the long struggles, now all but completely
won, of biological-evolutionary science against the retarding forces of es-
sentialism, natural theology, and reductionism, and the rise at last of pop-
ulation thinking and natural selection theory. These themes are driven home
relentlessly throughout the book, occasionally through repetition that ex-
ceeds the bounds of didactic advantage. Examples of some other less con-
ventional but welcome lines of thought are Mayr’s espousal of the philosophy
of emergence, and his vigorous correction of the common misconception
that Lamarck was a failed evolutionist.
The range of literature in philosophy and science that is covered (and
referenced) in this history is truly impressive, and the reader learns again
and again of important contributions by scholars whose names have been
barely familiar to most.
Like many histories that reach into present times, this one tends to fall
off into spotty and idiosyncratic, often perfunctory discussion of evolutionary
issues currently in controversy— for example, group selection. I doubt wheth-
er there really is a “general consensus” that most ostensible group selection
cases “can be interpreted in terms of individual selection, except perhaps in
social animals . . . .”
Some other interesting topics, probably even less to Mayr’s taste, are barely
or never mentioned. The revolt against the concept of race (subspecies), so
widely flaring in the fifties, was grudgingly recognized by him in 1963, but
by 1982, the gate has clanged tight against this and other heretical uprisings.
Mayrian peripatric speciation, proposed as a theory of incipient macroevo-
lution in 1954, has become the firm dogma of 1982, although interesting
models exist that offer attractive alternatives to some of its main assertions.
In contrast to his silence on these ideas, Mayr introduces some contributions
of his own, or of his students, that do not seem as cardinally important to
evolutionary theory.
It is difficult indeed to exaggerate the importance of this man in distilling
and in teaching to my generation the systematics and evolutionary theory
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NEW YORK ENTOMOLOGICAL SOCIETY
of the mid-twentieth century, especially through his influential classic of
1942. Yet in a curious way Mayr does himself succeed in producing this
exaggeration because, despite disclaimers to the contrary, this history gives
the impression that the growth of biological thought has reached a sort of
culmination for Mayr in his personal perceptions and opinions (fulsomely
but incompletely indexed on p. 968) of its state as of about 1960. But as the
facts related in this great volume suggest, every contributor and his contri-
bution, no matter how fundamentally correct and triumphant they may
appear contemporaneously, are liable to suffer some revision as the surprises
of time and discovery emerge.
It seems likely to me that the evolutionary understandings of a half century
from now will view many of our current concepts as quaint. But looking
back a full century from then, to 1933 and earlier, they may well find their
agreement with Mayr’s history becoming more substantially complete. Let
us then celebrate and learn from the earlier periods of coverage by this book,
and be cautious about its account of modern times. — William L. Brown, Jr.,
Department of Entomology, Cornell University, Ithaca, New York 14853.
Vicariance Biogeography: A Critique. — Gareth Nelson and Donn E. Rosen
(eds.). 198 1 . Columbia University Press, New York, xvi + 593 pp. $35.00.
Vicariance Biogeography is a historical approach to biogeography which
searches for general patterns of relationship among areas of endemism. These
patterns are discovered through congruence among taxa cladograms— con-
gruence which can presumably be attributed to the vicariance of a widespread
ancestral biota, but not to the combined effect of chance dispersal events.
Vicariance biogeography has also been called the “Platnick, Nelson, and
Rosen method” (Patterson, this volume, p. 466) due to the method’s for-
malization by Platnick and Nelson (1978) and application by Rosen (1978).
A more lengthy explication of the method may be found in Nelson and
Platnick (1981).
Among the more salient factors which have contributed to the formali-
zation of vicariance biogeography are: (1) the growing evidence in support
of continental drift (cf. Darlington, 1957, 1 965; Tarling and Runcorn, 1973);
(2) the introduction of Hennigian phylogenetics into the English language
(Elennig, 1965, 1 966); (3) the union of continental drift theory and Hennigian
phylogenetics (Brundin, 1966); (4) the introduction ofPopperian philosophy
into phylogenetic systematics (Bock, 1973; Ball, 1975; Wiley, 1975); and (5)
the incorporation of various aspects of Croizat’s “Panbiogeography” (Croizat
et ah, 1974; Rosen, 1975). Application of drift theory to biogeography had
already been attempted in Jeannel’s La Genese des Faunes Terrestres ( 1 942).
Unfortunately, this antedated the vindication of continental drift and the
VOLUME 91, NUMBER 3
285
development of an explicit means of inferring the relative recency of common
ancestry among taxa.
MacArthur and Wilson (1967:5) have criticized historical biogeography,
stating that: “The conventional issues relate to specific places and specific
groups of plants and animals” and, therefore, the “major issues are ad hoc
and historically oriented; for example: What was the ultimate origin of the
Antillean vertebrate fauna?”; “Did Central America develop a discrete in-
sular fauna during the Tertiary?”; “How can we account for the phylogenetic
similarities of the biotas of southern South America and New Zealand?”;
“Why is Hawaii rich in species of Nesoprosopis but lacking in other native
bee genera?”. These are exactly the kinds of biogeographic questions which
systematists— neontologists and paleontologists — are most interested in. All
questions concerning the distribution of organisms properly fall under the
heading of biogeography; however, it should be obvious that when one is
asking different questions one might need to employ different methods.
Vicariance biogeography presumably obviates MacArthur and Wilson’s crit-
icism of historical biogeography in that it (1) searches for general patterns
and (2) produces biogeographic hypotheses which are predictive and testable
(Nelson and Platnick, 1981).
Vicariance biogeography has indirectly benefited from a de-emphasis on
speciation via founder events (Mayr, 1942, 1963), a mode of speciation
perfectly amenable to dispersalist biogeography. Templeton (1981), based
upon a review of the population genetics literature, concludes that among
divergence types of speciation (adaptive, clinal, and habitat) adaptive di-
vergence (the erection of an extrinsic isolating barrier followed by indepen-
dent microevolution) “is probably the dominant mode in both plants and
animals” (p. 39). Among transilience modes, Templeton concludes that
hybrid maintenance and hybrid recombination are important, particularly
in plants, and that genetic transilience (speciation via a founders event) can
be important for certain groups and situations. Speciation by chromosomal
transilience is judged to be relatively rare. Among all the speciation modes
discussed by Templeton, adaptive divergence (speciation following vicari-
ance) is painted as the most general. Bush (1975:357) suggested that “the
number of animals that may be speciating sympatrically or parapatrically
(i.e., rodents, parasites, flightless insects, etc.) might exceed or at least equal
the number of those speciating allopatrically.” This is a ridiculous statement
which implies that rodents, parasites, and flightless insects do not speciate
allopatrically.
The present volume, Vicariance Biogeography: A Critique, is the product
of a three day symposium (May 2-4, 1979) organized by the Systematic
Discussion Group of the American Museum of Natural History. The purpose
of the symposium according to Rosen (Introduction, p. 4) was to provide a
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NEW YORK ENTOMOLOGICAL SOCIETY
forum to discuss whether vicariance theory and method as recently discussed
by various authors, was useful, useless, or irrelevant for dealing with prob-
lems of historical biogeography. According to Rosen (Introduction, p. 3):
“It was the decision of the altered committee that the symposium should
include speakers who, except for Croizat, had never before written on vi-
cariance theory and who represented recognizably different points of view
in biogeography.” Although I found the entire text interesting and enjoyable
reading, I question whether the volume as a whole constitutes a thorough
critique of vicariance biogeography. This may be due in part to the choice
of speakers and in part to the speakers’ choice of topics.
Rosen (Introduction, p. 1) refers to Croizat as a vicariance biogeographer,
and yet Croizat ( 1 982) flatly denies being a Hennigian. One can only conclude
from Croizat (1982) that Croizat is not a vicariance biogeographer. Vicar-
iance biogeography is apparently a hybridization between Brundin’s phy-
logenetic biogeography and Croizat’s “Panbiogeography,” and it presumably
incorporates the best aspects of each. “Panbiogeography” offers to vicariance
biogeography the concept of generalized tracts (congruent distribution pat-
terns) against a background of allopatric speciation by vicariance which
sidesteps the Neodarwinistic and largely dispersalistic approaches of Dar-
lington (1957) and Simpson (1965). Croizat’s (1982) falling out with vicar-
iance biogeography may be due to its piece meal incorporation of various
aspects of “Panbiogeography” and the unsolicited editorial notes interjected
within his contribution to the present volume.
Vicariance biogeography, as mentioned previously, searches for congru-
ence between area cladograms (generalized tracks, sensu Platnick, Nelson,
and Rosen). A critique of vicariance biogeography should, therefore, be a
critique of generalized tracks. Other pertinent issues would include phylo-
genetic methods, Popperian philosophy, and models of speciation. Conti-
nental drift is not really an issue.
Six of the twelve invited papers are largely ancillary as critiques of vicar-
iance biogeography. These include; Erwin’s discussion of “taxon pulses”;
Solem’s discussion of land-snail biogeography; Hallam’s review of plate
movements, eustasy, and climate since the early Mesozoic; two papers dis-
cussing evidence for a lost Pacific continent, one by Melville and one by
Nur and Ben-Avraham; and the paper by Haffer on Neotropical bird spe-
ciation. It is interesting, and perhaps no coincidence, that these six papers
are buried centrally and consecutively within the text. Not surprisingly,
discussions of these six papers are equally ancillary as critiques of vicariance
biogeography. This is due to no fault of the discussants.
Of the remaining six formal papers, only the paper by SimberlolT et al.
really constitutes a serious critique of vicariance biogeography by directly
questioning the statistical significance of congruent cladograms. Udvardy’s
VOLUME 91, NUMBER 3
287
paper is a useful interpretation of the possible position of vicariance bio-
geography within biogeography as a whole. The contributions by Brundin
and by Patterson are recommended reading for an understanding of the
difference between “phylogenetic biogeography” and “vicariance biogeog-
raphy,” a division which in some respects parallels the divisions referred to
as “process” and “pattern” cladism (Platnick, 1979). Wolfe’s paper on “Vi-
cariance biogeography of angiosperms in relation to paleobotanical data”
should be noted for the interesting discussion which it elicited. The hnal
invited paper in the volume by Croizat is equally as entertaining as his 1982
paper in Systematic Zoology.
Nelson’s summary of the symposium is clearly partisan and his manip-
ulations of what the participants actually said are unwarranted. His reference
to participants’ reservations and criticisms of vicariance biogeography as
“stumbling blocks” casts an air of naivete upon the participants— a display
of arrogance which will be more of a disservice than a shot in the arm for
vicariance biogeography.
The format chosen for the symposium and this volume — contributed pa-
per followed by discussants’ comments and a hnal response — is excellent. I
detected very few typographical errors in the text. I have reservations about
symposium volumes in general; however, given the excellent format, good
physical production, and the relatively low cost of this volume, I would
recommend it to anyone with more than a passing interest in biogeography. —
Stephen W. Nichols, Department of Entomology, Cornell University, Ithaca,
New York 14853.
LITERATURE CITED
Ball, I. R. 1975 (1976). Nature and formulation of biogeographical hypotheses. Syst. Zool.
24:407-430.
Bock, W. J. 1973 (1974). Philosophical foundations of classical evolutionary taxonomy. Syst.
Zool. 22:375-392.
Brundin, L. 1966. Transantarctic relationships and their significance. K. Sven. Vetenskaps-
akad. Handl., series 4, 11(1): 1-472.
Bush, G. L. 1975. Modes of animal speciation. Ann. Rev. Ecol. Syst. 6:339-364.
Croizat, L., G. Nelson and D. E. Rosen. 1974. Centers of origin and related concepts. Syst.
Zool. 23:265-287.
Croizat, L. 1982. Vicariance/vicariism, panbiogeography, “vicariance biogeography”, etc.: a
clarification. Syst. Zool. 31:291-304.
Darlington, P. J., Jr. 1957. Zoogeography: The Geographical Distribution of Animals. John
Wiley and Sons Inc., New York.
Darlington, P. J., Jr. 1965. Biogeography at the Southern End of the World: Distribution and
History of Far-Southern Life and Land, with an Assessment of Continental Drift. Harvard
University Press, Cambridge, Massachusetts.
Hennig, W. 1965. Phylogenetic systematics. Ann. Rev. Entomol. 10:97-116.
Hennig, W. 1966. Phylogenetic Systematics. University of Illinois Press, Urbana.
Jeannel, R. 1942. La Genese des Faunes Terrestres. Presses Universitaires de France, Paris.
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NEW YORK ENTOMOLOGICAL SOCIETY
MacArthur, R. H. and E. O. Wilson. 1967. The Theory of Island Biogeography. Monographs
in Population Biology, No. 1. Princeton University Press, Princeton, New Jersey.
Mayr, E. 1942. Systematics and the Origin of Species. Columbia University Press, New York.
Mayr, E. 1963. Animal Species and Evolution. Harvard University Press, Cambridge, Mas-
sachusetts.
Nelson, G. and N. I. Platnick. 1981. Systematics and Biogeography. Columbia University
Press, New York.
Platnick, N. I. 1979. Philosophy and the transformation of cladistics. Syst. Zool. 28:537-546.
Platnick, N. I. and G. Nelson. 1978. A method of analysis for historical biogeography. Syst.
Zool. 27:1-16.
Rosen, D. E. 1975 (1976). A vicariance model of Caribbean biogeography. Syst. Zool. 24:
431-464.
Rosen, D. E. 1978. Vicariant patterns and historical explanation in biogeography. Syst. Zool.
27:159-188.
Simpson, G. G. 1965. The Geography of Evolution. Chilton Books, Philadelphia and New
York.
Tarling, D. H. and S. K. Runcorn (eds.). 1973. Implications of Continental Drift to the Earth
Sciences. 2 vols. Academic Press, New York.
Templeton, A. R. 1981. Mechanisms of speciation — a population genetics approach. Ann.
Rev. Ecol. Syst. 12:23-48.
Wiley, E. O. 1975. Karl R. Popper, systematics, and classification: a reply to Walter Bock
and other evolutionary taxonomists. Syst. Zool. 24:233-243.
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Journal of the
New York Entomological Society
VOLUME 91 SEPTEMBER 1983
NO. 3
CONTENTS
Exotic insects reported new to Northeastern United States and Eastern Canada since
1970 E. Richard Hoebeke and A. G. Wheeler, Jr.
The types of cercerine wasps described by Nathan Banks (Hymenoptera:
Philanthidae) George R. Ferguson
Two new species and synonymy of three species of North American Cerceris (Hy-
menoptera: Philanthidae) George R. Ferguson
Gynandromorphic desert fire ant, Solenopsis aurea Wheeler (Hymenoptera:
Formicidae) James C. Cokendolpher and Oscar F. Franke
A redefinition of Disderia and addition of a new species (Hemiptera: Pentatomidae)
F. H. Rolston
A study of winged queens of the Colorado honey ant, Myrmecocystus mexicanus, in
captivity John R. Conway
Nesting biology of the bee Svastra sabinensis (Hymenoptera: Anthophoridae)
Jerome G. Rozen, Jr.
Response of a goldenrod beetle to four seldom-encountered goldenrod (Solidago)
species Frank J. Messina
Precopulatory behavior in the whirligig beetle Dineutes discolor (Coleoptera:
Gyrinidae) Steven A. Kolmes
SEM study of the antennal sensilla and setae of Solva pallipes (Loew) (Diptera:
Xylomyidae) C. E. Vasey and E. Ritter
Book Reviews
The Growth of Biological Thought. Diversity, Evolution, and Inheritance
William F. Brown, Jr.
Vicariance Biogeography: A Critique Stephen W. Nichols
193-222
223-234
235-241
242-245
246-251
252-263
264-268
269-272
273-279
280-282
283- 284
284- 288
ffS,70(>73
Vol. 91 - DECEMBER 1983 No. 4
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MAR 2 7 1984
NW YORK ENTOMOLOGICAL SOCIETY
91(4), 1983, pp. 289-303
ZEBRATUS GROUP
(HYMENOPTERA: PHILANTHIDAE)
George R. Ferguson
Systematic Entomology Laboratory, Department of Entomology,
Oregon State University, Corvallis, Oreogn 97331
Abstract.— Philanthus basilaris Cresson is removed from synonymy with Philanthus zebratus
Cresson. A key to the six species of the Philanthus zebratus group is presented and their
relationships, distribution and intraspecific variation discussed.
The Philanthus zebratus group, as defined by Bohart and Grissell (1975),
is characterized by a narrow vertex and by the inner eye margins diverging
below the eye emarginations. Philanthus basilaris Cresson has been found
to be a good species and is removed from synonymy with Philanthus zebratus
Cresson. The six species in the group are characterized and discussed, and
a key for their separation is presented.
During the past few years I have received for identification a large number
of philanthid wasps collected by Dr. F. D. Parker and colleagues at the
USDA Bee Biology and Systematics Laboratory, Utah State University,
Logan. Most of the collections have been from areas in and around the San
Rafael Desert, Emery County, Utah. Among the 12 species of Philanthus
collected is a long series of P. basilaris Cresson previously synonymized
under P. zebratus Cresson. The studies reported here were undertaken to
clarify the relationships and distribution of P. basilaris and its closely related
congeners. Over 1,500 specimens have been examined in the course of the
work.
The following abbreviations are used in the text and in the key:
LOD— Least distance from lateral ocellus to eye margin measured in lateral
ocellus diameters.
MOD— Least distance from midocellus to eye margin measured in mid-
ocellus diameters.
V/MO— Least width of vertex divided by diameter of midocellus.
H/F Ratio — Maximum width of head divided by width of face between
apices of eye emarginations.
‘ Oregon Agricultural Experiment Station Technical Paper No. 6551.
The publication costs of this article were defrayed in part by page charge payment. This article
must therefore be hereby marked “Advertisement” in accordance with 18 U.S.C. §1734 solely
to indicate this fact.
290
NEW YORK ENTOMOLOGICAL SOCIETY
F/V Ratio— Width of face between apices of eye emarginations divided by
least width of vertex.
C/F Ratio— Maximum width of clypeus divided by width of face at apices
of eye emarginations.
Measurements in the ocellocular area were made at 40-70 x magnification,
and those of the head, face and clypeus at 10-30 x depending on the size of
the specimen. Variation in the vertex and ocellocular ratios is given in the
diagnosis of each species. The H/F and C/F ratios were remarkably constant
within species. Only those ocellocular ratios which show a substantial dif-
ference are used in the key.
Abbreviations of institutional repositories are given under Acknowledg-
ments.
A progressive narrowing of the vertex occurs within the group such that
three subgroups may be recognized. In the gloriosus subgroup, composed of
Philanthus gloriosus Cresson and Philanthus bicinctus (Mickel), the LOD is
O. 7 to 1.5 in males and 2.0 to 2.5 in females; the F/V ratio is 1.8 to 2.3 in
males and about 1.5 in females. In the zebratus subgroup, composed of
Philanthus basilaris Cresson, Philanthus ventralis (Mickel) and Philanthus
zebratus Cresson, the LOD is less than 0.5 in males and about 1 .0 in females;
the F/V ratio is about 3 in males and about 2 in females. In the intermediate
sanbornii subgroup, composed only of Philanthus sanbornii Cresson, the
males have the vertex and ocellar characteristics of the zebratus subgroup,
and the females have those of the gloriosus subgroup.
Strandtmann (1946) recognized four species within the zebratus subgroup,
P. basilaris, P. nitens (Banks), P. ventralis and P. zebratus. Bohart and
Grissell (1975) recognized two species: P. ventralis and P. zebratus, synon-
ymizing P. basilaris and P. nitens under P. zebratus. They suggested that, if
subspecies of zebratus were recognized, the name to be used for the whitish
phase east of California would be P. basilaris with P. nitens as a synonym.
However, both P. basilaris and P. zebratus occur in a whitish phase. As
interpreted here P. basilaris is resurrected as a good species and P. nitens
remains a junior synonym of P. zebratus.
The following key is adapted and modified from that given by Bohart and
Grissell (1975).
KEY TO Philanthus zebratus group
1 . Males 2
- Females 7
2. MOD at least 1.0; LOD at least 0.7 3
- MOD less than 1.0; LOD less than 0.5 4
3. MOD about 1.5; LOD about 1.2; clypeal brush mostly yellowish gloriosus Cresson
- MOD about 1.0; LOD about 0.75; clypeal brush mostly or entirely blackish
bicinctus (Mickel)
VOLUME 91, NUMBER 4
291
4. Scutal punctures dense, mostly separated by about 1 puncture diameter, more or less
uniformly distributed; tergum III with punctures about same size as on scutum; pu-
bescence of head and thorax brown to fuscous; tergum I all dark (black or red) or with
widely separated pale spots sanbornii Cresson
- Scutal punctures, at least medially, unevenly separated by many smooth interspaces
of several puncture diameters; tergum III with at least a few smooth-rimmed punctures
distinctly larger than scutal punctures; pubescence of head and thorax pale yellow to
white; tergum I with a pale band which may be incised medially or narrowly divided
5
5. Antennal flagellum pale below to tip; pale bands of terga I and/or II interrupted
medially ventralis (Mickel)
- Antennal flagellum with apical 1 to 5 flagellomeres black or infuscated below; pale
bands of terga I and II continuous (rarely interrupted in northern Rocky Mountain
forms) 6
6. H/F ratio 1.8; propodeal enclosure usually shagreened; tergum I and/or sternum I
usually marked with red basilaris Cresson
- H/F ratio 1.7; propodeal enclosure polished between punctures; tergum I and sternum
I rarely marked with red zebratus Cresson
7. MOD 2.4 to 3.1; LOD 2.0 to 2.5; V/MO 6 to 7 8
- MOD 1.2 to 1.5; LOD 0.8 to 1.1; V/MO 3.2 to 4.0 10
8. Scutal punctures sparse, unevenly distributed, almost impunctate across middle third;
terga III-V black; about 20 mm long bicinctus (Mickel)
- Scutal punctures dense and more or less evenly distributed, at least on lateral third;
one or more of terga III-V maculated; length 1 8 mm or less 9
9. Punctures of tergum III ravher dense and uniformly distributed; tergum I about twice
as wide as long, distinctly transverse; about % of surface of lateral lobe of clypeus
finely and densely punctate; pale band on tergum II interrupted medially
sanbornii Cresson
- Punctures of tergum III sparse and irregularly spaced; tergum I about 1.5 times as
wide as long, somewhat bell-shaped; about % of surface of lateral clypeal lobe polished,
impunctate; pale band on tergum II complete or anteriorly emarginate, not completely
interrupted gloriosus Cresson
10. Scutal punctures sparse, separated by many large, polished areas; pale facial mark
truncate or crown-shaped above, separated from midocellus by much more than
ocellus diameter zebratus Cresson
- Scutal punctures dense laterally, at least along parapsidal lines; pale facial mark nar-
rowed above, reaching to midocellus or nearly so 1 1
1 1. Pale bands of terga I/II continuous; H/F ratio 1.7; tergum I and/or sternum I usually
marked with red basilaris Cresson
- Pale bands on terga I and II usually interrupted or deeply incised medially; H/F ratio
1.6; terga and sterna never marked with red ventralis (Mickel)
Philanthus gloriosus Cresson
Philanthus gloriosus Cr^s^on, 1865:86. [Female lectotype, Colorado; ANSP];
Strandtmann, 1 946:3 1 and prior literature; Burks, 1 95 1 : 100 1 ; Bohart and
Grissell, 1975:11; Bohart and Menke, 1976:565; Krombein, 1979:1723.
Philanthus insignatus Banks, 1913:421. [Female holotype, Alpine, Texas;
AMNH]; Synonymy by Strandtmann 1946:31.
292
NEW YORK ENTOMOLOGICAL SOCIETY
Diagnosis. Male 14-16 mm; LOD 1.2 (1.0-1. 5); MOD 1.6 (1.3-1. 8); V/
MO 4.2 (3. 8-4.5); H/F 1.6; F/V 1.8; C/F 1.1; clypeal brush pale. Female
16-18 mm; LOD 2.5 (2.4-2. 8); MOD 3.1 (2.8-3. 3); V/MO 7.0 (6. 5-7. 5);
H/F 1.6; F/V 1.4; C/F 1.2; scutal punctures dense except medially. P. glo-
riosus is assumed to be the most generalized member of the group since the
vertex is wider relative to the width of the face than in any other species
within the group.
Variation. Color pattern is quite constant in the populations east of the
Rocky Mountains, eastern Arizona and in Mexico. Females have a deep
tongue-shaped anterior emargination in the pale band of tergum II, and
males have a triangular posterior emargination on tergum II completely
interrupting the pale band. The background color is usually bright red, es-
pecially in females, in the desert areas in and around southeastern Arizona.
Both northward and southward the red gradually changes to black such that
the specimens from Canada and Mexico D.F. have identical coloration with
a black and yellow pattern and few, if any, red markings. A similar double
dine also occurs in Cerceris sexta Say which has a similar distribution.
In northern Arizona (Coconino and Mohave Counties), Utah and Cali-
fornia the emargination on tergum II has been lost in both sexes, the spec-
imens being broadly banded with bright yellow. Dorsally the background
color is largely black. Morphologically this form differs from the eastern
form only in having the terga somewhat more sparsely punctate. An anomaly
is the fact that males collected at Government Holes, 5,000 ft., San Bernadino
County, California, IX- 12-76 [USU] are identical in color pattern to the
populations east of the Rocky Mountains. Specimens of the “yellow” form
are in the collections of ALB, AZS, UCD, CDA, OSU, USNM, and USU.
Flight period. Dates of capture are July-August in the northern plains;
August-September in southern Arizona, New Mexico, Texas and Chihua-
hua; and September-October in the remaining states of Mexico. The species
is apparently univoltine in the above areas.
California records are in June-July and September-October leading Bo-
hart and Grissell (1975) to suggest that there were two broods per year.
Collection dates in northern Arizona and Utah are in May-June and August-
September, indicating that the species is bivoltine in that area as well.
Distribution. West of the 100th meridian in the United States and Canada
from Alberta (Medicine Hat) and Saskatchewan (near Empress, Alberta)
south through the high plains and eastern slope of the Rocky Mountains to
Culberson and Brewster Counties, Texas; Hidalgo County, New Mexico; and
west through eastern and northern Arizona and Utah to southeastern Cal-
ifornia. In Mexico it occurs in the states of Chihuahua, Coahuila, Durango,
San Luis Potosi, Hidalgo, Mexico D.F., and Puebla.
Specimens examined. 134 males, 96 females.
VOLUME 91, NUMBER 4
293
Philanthus bicinctus (Mickel)
Ococletes [sic] bicinctus Mickel, 1916:407. [Female holotype, Ute Creek,
Sage Flats, Colorado; NEB]; 1918:326.
Ococletes [sic] hirticulus Mickel, 1918:326. [Male holotype, Ute Creek, Sage
Flats, Colorado; NEB].
Philanthus bicinctus, Strandtmann, 1946:35; Burks, 1951:1001; Evans, 1964:
275 (larva); Krombein, 1967:413; 1979:1721; Bohart and Grissell, 1975:
5, 7 (in key); Bohart and Menke, 1976:564.
Philanthus hirticulus, Strandtmann, 1946:40; Burks 1951:1001; Synonymy
by Krombein 1967:413.
Diagnosis. Male 17-19 mm; LOD 0.7 (0.6-0.8); MOD 1.1; V/MO 3.2
(3. 1-3.3); H/F 1.6; F/V 2.3; C/F 1.2; clypeal brush blackish, sometimes pale
at tip. Female 20-22 mm; LOD 2.1 (2.0-2.2); MOD 2.7 (2.6-2.7); V/MO
6.4 (6. 1-6.6); H/F 1.6; F/V 1.5; C/F 1.2; scutal punctures sparse, mostly
impunctate over middle third. In males the eyes are somewhat more di-
verging at the clypeus with a C/F ratio of 1.2 compared to 1.1 for the
remaining species of the group.
P. bicinctus is our largest species. The males are yellow banded and re-
semble males of the western form of P. gloriosus, but the larger size and
characters given in the key easily separate them. The female is a strikingly
beautiful insect with a red tergum I, yellow tergum II, and the remaining
terga black.
Biology. Digging, feeding, provisioning, nesting and hunting activities of
females as well as male behavior suggesting territoriality were described by
Armitage (1965). Nests were found in aggregations in a silty clay soil. Prey
consisted of eight species of worker and male Bombus with Megachilidae
{Megachile sp.) and Anthophoridae {Anthophora sp., Melissodes sp.) also
taken. Gwynne (1978) and Gwynne and O’Neill (1980) described and dis-
cussed male territorial behavior. Gwynne (1980, 1981) described female
behavior and nesting activities. Prey records included ten species of Bombus
plus one species of Psithyrus and Apis mellifera in the Apidae; five species
of Megachilidae in two genera; two species of Anthophoridae in two genera;
and one species of Sphecidae {Podalonia communis). Bumblebees consti-
tuted about 99% of the prey in 1976, 1977, but only 39% of the prey in
1978.
Flight period. July to September.
Distribution. Rocky Mountains from Montana to southern Colorado at
recorded elevations of 6,900 to 8,000 ft.
Specimens examined. 6 males, 4 females.
294
NEW YORK ENTOMOLOGICAL SOCIETY
Philanthus sanbornii Cresson
Philanthus sanbornii Cresson, 1865:89. [Female lectotype, Massachusetts;
ANSP]; Strandtmann, 1946:37 and prior literature; Burks, 1951:1002;
Krombein, 1958:197; 1979:1725; Bohart and Grissell, 1975:5, 7 (in key);
Bohart and Menke, 1976:566.
Philanthus scutellaris Cvtsson, 1879:xxxiv. [Male holotype, Kansas; ANSP];
Synonymy by Strandtmann, 1946:37.
Philanthus eurynome Fox, 1890:107. [Female lectotype, Florida; ANSP];
Strandtmann, 1946:34; Burks, 1951:1001; Krombein, 1967:413; Synon-
ymy by Bohart in Bohart and Menke, 1976:566.
Philanthus trumani Dunning, 1897:70. [Male, not female, holotype, Brook-
ings County, South Dakota; location of type unknown]; Synonymy by
Dunning, 1898:152.
Philanthus magdalenae Strandtmann, 1946:39. Name created in synonymy,
authorship (p. 37) incorrectly attributed to Viereck.
Diagnosis. Male 12-14 mm; LOD 0.3 (0.3-0.5); MOD 0.7 (0.6-0.7); V/
MO 2.4 (2.3-2.5); H/F 1.7; F/V 3.1; C/F 1.1; pubescence brown to fuscous
on head and thorax. Female 14-17 mm; LOD 2.0 (1.8-2. 3); MOD 2.4 (2.0-
3.0); V/MO 6.0 (53.-7. 2); H/F 1.6; F/V 1.6; C/F 1.2; scutum, scutellum and
terga rather densely and uniformly punctate. P. sanbornii exhibits an inter-
mediate condition in the evolution of the ocellocular area. The vertex of the
male is narrowed markedly compared to the gloriosus subgroup, whereas
the female vertex is not. The eyes of the male have slightly enlarged to a H/
F ratio of 1.7 compared to 1.6 in both P. gloriosus and P. bicinctus.
Variation. The geographical variation in color pattern is a common one
in philanthid wasps. The Florida specimens are usually heavily marked with
red on the anterior terga, and the western specimens are more highly mac-
ulated with yellow.
Biology. Evans (1955) and Evans and Lin (1959, as P. eurynome) reported
honeybees used as prey with Halictidae also taken. Nesting was in sandy
areas but not in dense aggregations. Megachile inermis Provancher (det. W.
P. Stephen) is also used as prey based on a specimen of P. sanbornii pinned
with this bee from Aweme, Manitoba, VI-28-78 [MTB].
Flight period. Dates of capture are March-April in Florida; June in Geor-
gia; and June 15 to August 30 elsewhere. It is apparently univoltine.
Distribution. United States and southern Canada east of the 100th merid-
ian with occasional captures in the high plains. Peripheral records west of
the 100th meridian are: ALBERTA (Writing-On-Stone) [ALB]; NORTH
DAKOTA (Dunn County) [NDS]; NEBRASKA (Thomas County) [OSU];
and NEW MEXICO (Socorro County), the latter record from Strandtmann
(1946).
Specimens examined. 48 males, 62 females.
VOLUME 91, NUMBER 4
295
Philanthus ventralis (Mickel)
Ococletes [sic] ventralis Mickel, 1918:329. [Female holotype, Sacramento,
California; NEB].
Philanthus ventralis, Strandtmann, 1946:43; Bohart and Grissell, 1975:18;
Bohart and Menke, 1976:566; Krombein, 1979:1726.
Philanthus strandtmanni Burks, 195 1 : 1002. Unnecessary new name for Phi-
lanthus ventralis (Mickel).
Diagnosis. Male 12-14 mm; LOD 0.3 (0. 3-0.4); MOD 0.7 (0.6-0. 8); V/
MO 2.5 (2.4-2.7); H/F 1.7; F/V 2.9 (2.6-3.2); C/F 1.1. Female 13-15 mm;
LOD 1.1 (0.9-1. 2); MOD 1.5 (1.3-1. 7); V/MO 3.9 (3. 5-4.4); H/F 1.6; F/V
2.1 (2. 0-2. 2); C/F 1.2. See Table 1 for additional characters separating this
species from P. basilaris and P. zebratus.
Variation. There is little variation in morphology or color pattern over
most of the range of this species. However, some males from southern
California (San Diego County, Kitchen Creek, Laguna Mountains, IX-31-
78 [LAM], and Kem County, Walker Pass, IX-26/27-57 [UCD]) have the
apical two or three flagellomeres infuscated below. Over the balance of the
range of the species the flagellum is pale below to the apex. This may indicate
some introgression of either P. basilaris or P. zebratus genes.
Flight period. This is a late summer and early fall species with most
captures in August-September. Bohart and Grissell (1975) provide a fre-
quency distribution showing a peak flight period in September. Oregon and
Washington specimens have been collected between August 9 and September
8 with a peak in late August. The earliest collection date is July 6 in Siskiyou
County, California, and the latest date is October 2 1 in Contra Costa County,
California.
Distribution. California and western Nevada to British Columbia. Bohart
and Grissell (1975) presented a distribution map and detailed locality records
for California. In Oregon I have seen specimens from Baker, Harney, Kla-
math, Union and Wheeler Counties, all east of the Cascade range. The Inyo
County, California, specimens listed by Bohart and Grissell (1975) are dis-
cussed under P. basilaris.
Specimens examined. 159 males, 67 females.
Philanthus basilaris Cresson (Revised Status)
Philanthus basilaris Cresson, 1879:xxxiii. [Female lectotype, Colorado;
ANSP]; Strandtmann, 1946:48 and prior literature; Burks, 1951:1001.
Philanthus zebratus, in part, Bohart and Grissell, 1975:1 8; Bohart and Menke,
1976:567; Krombein, 1979:1726.
Diagnosis. Male 12-14 mm; LOD 0.3 (0.2-0.4); MOD 0.8 (0.7-0.9); V/
MO 2.5 (2.3-3.0); H/F 1.8; F/V 2.8 (2.6-3. 2); C/F 1.1. Female 13-15 mm;
296
NEW YORK ENTOMOLOGICAL SOCIETY
LOD 0.8 (0.7-0.9); MOD 1.3 (1. 1-1.6); V/MO 3.4 (3. 1-3.7); H/F 1.7; F/V
2.0 (1.9-2. 1); C/F 1.2. See Table 1 for additional diagnostic characters.
P. basilaris resembles P. ventmlis in having denser scutal punctation com-
pared to P. zebratus, and in its late-season flight period compared to a
midsummer flight period in P. zebratus. The swollen eyes of P. basilaris,
indicated by a H/F ratio of 1.8 in males and 1.7 in females, separate it from
both P. ventralis and P. zebratus. In the latter two species the H/F ratio is
1.7 in males and 1.6 in females. Another distinctive feature is that the tergal
bands of P. basilaris are broad and continuous in both sexes, whereas in P.
ventralis they are interrupted, at least on the anterior terga, in both sexes,
and in P. zebratus the bands are continuous in the males and interrupted in
the females.
Variation. Coloration is rather constant over most of the range of this
species. The broad tergal bands vary from lemon yellow to pale yellowish
white, and tergum I as well as sterna I-II are conspicuously marked with
red. Nevada specimens are whiter, and the red marks are reduced or absent.
The pale markings of a male from Humboldt County, Nevada, are entirely
white. California specimens are more of a bright yellow, and red marks are
absent. Three of four males from Mono County, California [UCD], and one
of eight females from Inyo County, California [UCD] have the pale band of
tergum I narrowly interrupted, indicating a possible introgression of P. ven-
tralis or P. zebratus genes.
Biology. Prey consists of diverse adult Hymenoptera, as O’Neill and Evans
(1981) reported that the contents of an excavated nest of P. basilaris consisted
of two conspecific males and a male of P. bicinctus “intermingled with other
bees and wasps serving as food for the larvae.” Evans and O’Neill (1978)
and Gwynne and O’Neill (1980) described male territoriality in populations
from Weld and Alamosa Counties, Colorado. At that time these populations
were considered to belong to P. zebratus, but later (O’Neill and Evans, 1981)
they were recognized as P. basilaris. These populations were compared to a
population of P. zebratus from Jackson Hole, Wyoming, that did not exhibit
territoriality. Nests were not concentrated into dense aggregations in the
Colorado populations of basilaris as they were in the case of the Wyoming
population of zebratus.
Flight period. P. basilaris, like P. ventralis, is a late-summer and early-fall
species. Dates of capture are mostly in August-September with an early
record of July 17 in Alamosa County, Colorado (Great Sand Dunes), and a
late record of October 12 in Mono County, California. It is apparently
univoltine.
Distribution. From Alberta and Saskatchewan south through the high plains
and eastern slope of the Rocky Mountains to McKinley County, Texas, and
west through northern New Mexico and northern Arizona to Utah, Nevada,
and Mono and Inyo Counties, California. Records and collection dates are
VOLUME 91, NUMBER 4
297
summarized as follows: AL.BERTA (8 km W of Buffalo, VIII-27) [ALB];
SASKATCHEWAN (Tp. 5, Rge 25, W.3 Mer., VIII- 17) [ALB]; NORTH
DAKOTA (Billings and Golden Valley Counties, VIII-30) [AMNH, MCZ,
USNM]; WYOMING (Carbon, Converse, Laramie, Platte and Sweetwater
Counties, VIII-4 to IX-27) [AMNH, UCD, COR, OSU, USNM, USU, UWY];
NEBRASKA (Dawes County, VIII-27) [NEB]; COLORADO (Alamosa,
Grand, Larimer and Weld Counties, VIII-4 to IX-5) [UCD, CSU, MCZ,
OSU, USU]; TEXAS (McKinley County, IX-21) [USU]; NEW MEXICO
(Catron and San Miguel Counties, VIII- 16 to VIII- 19) [ANSP, UCD, OSU];
ARIZONA (Coconino and Navajo Counties, VIII-30 to IX-23) [ALB, UCD,
OSU]; UTAH (Emery, Kane and Washington Counties, VIII- 19 to IX-20)
[UCD, UCR, KAN, OSU, USNM, USU]; NEVADA (Churchill, Douglas
and Humboldt Counties, VIII- 17 to IX-9) [UCD, OSU]; CALIFORNIA
(Inyo and Mono Counties, VIII-23 to X-12) [UCD, MIN, UMO, OSU].
Distribution is essentially sympatric with P. gloriosus over the northern
portion of the range of the latter.
Specimens examined. Female lectotype, 163 males, 76 females.
Philanthus zebratus Cresson
Philanthus zebratus Cresson, 1879:xxxiii. [Male holotype, Nevada; ANSP];
Strandtmann, 1946:46 and prior literature; Burks, 1951:1003; Bohart and
Grissell, 1975:18; Bohart and Menke, 1976:567; Krombein, 1979:1726.
Oclocletes nitens Banks, 1913:423. [Female, not male, holotype, Princeton,
British Columbia; MCZ].
Ococletes [sic] illustris Mickel, 1 9 1 8:326. [Female holotype, American River,
Placer County, California; NEB]; Synonymy by Strandtmann, 1946:46.
Philanthus nitens Strandtmann, 1946:42; Burks, 1951:1002; Synonymy by
Bohart and Grissell, 1975:19.
Diagnosis. Male 12-14 mm; LOD 0.3 (0.2-0. 3); MOD 0.6 (0.5-0. 7); V/
MO 2.3 (2.1-2.4); H/F 1.7; F/V 3.1 (2.8-3. 2); C/F 1.1. Female 14-16 mm;
LOD 0.9 (0.7-1. 1); MOD 1.3 (1. 1-1.6); V/MO 3.8 (3. 3-4.1); H/F 1.6; F/V
2.0 (1.9-2. 2); C/F 1.2. See Table 1 for additional diagnostic characters.
Variation. P. zebratus falls into three distinguishable populations. The
Rocky Mountain population is characterized by the females having red tibiae
and a tendency for the pale maculations to be white in a majority of the
specimens. About one-fourth to one-third of the specimens have yellow
maculations. The tendency for pale markings to change from yellow to white
in a south to north dine is common in western philanthid wasps.
The Siskiyou population is characterized by a reduced amount of black
on the underside of the male flagellum. Commonly only the apical flagello-
mere is infuscated below, which may represent an influence from adjacent
P. ventralis populations in which the flagellum is entirely pale below. The
298
NEW YORK ENTOMOLOGICAL SOCIETY
allochronicity would not be sufficient to prevent early-emerging ventralis
males from mating with late-emerging zebratus females, if such cross-breed-
ing can, in fact, occur. Females of the Siskiyou population usually have the
hind tibiae blotched with red, representing a condition intermediate between
the Rocky Mountain and Sierra Nevada populations— the latter having yel-
low tibiae with only occasional traces of red.
The Sierra Nevada population is characterized by a continued reduction
in density of scutal punctures with the posterior two-thirds of the scutum
being virtually impunctate (see Bohart and Grissell, 1975, Fig. 23). The
transverse band of fine punctures on the posterior margin of the scutum is
4 to 8 punctures wide in the Rocky Mountain and Siskiyou populations,
whereas this band of punctures is only 2 or 3 punctures wide, or virtually
absent, in the Sierra Nevada population.
Occasional males in the Rocky Mountain population have an interrupted
pale band on tergum I and greyish pubescence on the head and thorax. The
band on tergum I is usually continuous and the pubescence usually pale, as
in other populations. The sternal hair brushes of the male are often fuscous
in the Rocky Mountain population and usually amber in the Sierra Nevada
population, but these hair brushes in the Siskiyou population are interme-
diate, being brown to occasionally fuscous. The sterna of males of the Rocky
Mountain population are usually immaculate, but almost always maculated
in other populations.
A series of 9 males and 5 females collected in Wyoming, Sublette County,
3 mi. N Pinedale, VII- 15-71 (R. M. Bohart, E. E. Grissell) [UCD] shows
evidence of hybridization with P. basilaris. The head/face ratio of the males
varies from 1.7 to 1.8. The scutal punctures of the females are very sparse
in one specimen, almost as dense as in basilaris in one specimen, and in-
termediate in three specimens. The outer face of the hind tibia is yellow in
four females and red in one. The pale band of tergum I is interrupted in
three females and continuous in two. The pale band of tergum II is either
interrupted or deeply incised in all five females. The anterior terga and sterna
have an all black background color in four males and two females with the
remainder showing variable amounts of red. The pale facial mark of the
females is crown-shaped above and well separated from the midocellus. The
pale markings are white in females and white on terga III-VI in males.
Typical basilaris is known from Sweetwater County, the adjoining county
on the south, and typical zebratus is found in Teton County, the adjoining
county on the north and west.
The three populations of P. zebratus might be treated as subspecies, but
I have elected not to do so. The ranges of the populations are contiguous
and the differences tend to be clinal in nature. The possibility that Pleistocene
and Recent ash falls and lava flows from volcanic activity have tended to
temporarily isolate these populations cannot be overlooked. The Sierra Ne-
VOLUME 91, NUMBER 4
299
vada and Siskiyou populations of P. zebratus abut in Lassen County, Cal-
ifornia, the site of very recent volcanic activity. The Siskiyou and Rocky
Mountain populations adjoin in northern Klamath County, Oregon, an area
of extensive volcanic activity only a few thousand years ago.
Biology. Males are non-territorial, but rather form aerial swarms over the
densely aggregated nesting sites during the mating period according to Evans
and O’Neill (1978). Evans (1966, 1970) recorded 25 species of wasps and
20 species of bees used as prey by P. zebratus in the vicinity of Jackson
Hole, Teton County, Wyoming. The number of prey specimens was about
equally divided between 75 wasps and 74 bees. Families represented were
Ichneumonidae, Eumenidae, Masaridae, Sphecidae, Colletidae, Andrenidae,
Halictidae, Megachilidae, and Anthophoridae.
A series of 35 female zebratus collected on the same day, VII- 18-48, at
Hope Valley, Alpine County, California, (A. Bartel, J. W. Adams, P. D.
Hurd, J. W. MacSwain) [UCB] is each pinned with one or more prey spec-
imens and labelled “taken as prey.” The bees were determined by Dr. A.
Moldenke and the wasps by the author. One P. zebratus is pinned with two
small bees, i.e., Dufourea sp. (male) and Osmia sp. (female). It is implied
that the single zebratus female was carrying both specimens at the time of
capture. The remaining 34 female zebratus are each pinned with a single
prey specimen as follows:
VESPOIDEA
Ancistrocerus sp. (1 male, 1 female)
SPHECOIDEA
Ammophila stangei Menke (1 male)
Aphilanthops subfrigidus Dunning (3 males)
Podalonia luctuosa (F. Smith) (1 male)
Tachysphex sp. (1 female)
APOIDEA
Andrena (Andrena) sp. (3 females)
Andrena (Euandrena) sp. (1 female)
Andrena (Thysandrena) sp. (1 female)
Andrena (Trachandrena) spp. (2 females of 2 species)
Andrena (subgenus ?) sp. (2 females)
Hoplitis fulgida platyura (Cockerell) (1 female)
Hoplitis albifrons argentifrons (Cresson) (1 male)
Lasioglossum trizonatum (Cresson) (2 females)
Osmia (Chenosmia) sp. (3 females)
Osmia spp. (8 females, 2 males of 4 species)
Total. Eight wasps in 5 genera and 5 species; 28 bees in 5 genera and 16
species.
Since all captures were on the same day, the record probably reflects local
Table 1. Comparison of species of the zebratus subgroup.
300
NEW YORK ENTOMOLOGICAL SOCIETY
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VOLUME 91, NUMBER 4
301
abundance of prey rather than any difference in prey preference or diversity
when compared with Evans’ records above. Insofar as I can ascertain, these
are the first published prey records west of the Rocky Mountains and simply
confirm that P. zebratus uses a diverse assortment of Hymenoptera as prey.
Flight period. Rocky Mountain population: Most collection dates are in
July with an early record of May 27 in Valley County, Idaho, and a late
record of August 25 in Teton County, Wyoming. Siskiyou population: Dates
of capture are in July-August with an early date of June 20 in Trinity County,
California, and a late date of August 14 in Siskiyou County, California.
Sierra Nevada population: Collection dates are mostly in July-August with
an early date of June 12 in Plumas County, California, and a late date of
September 9 in Sierra County, California. Bohart and Grissell (1975) provide
a frequency distribution showing a pronounced peak in July. The species
appears to be univoltine throughout its range.
Distribution. Rocky Mountain population: BRITISH COLUMBIA (Chil-
cotin, Nicola, Okanagan Falls and Osoyoos); MONTANA (Missoula Coun-
ty); WYOMING (Sublette, Teton Counties and Yellowstone National Park);
IDAHO: (Owyhee and Valley Counties); WASHINGTON: (Douglas and
Kittitas Counties); OREGON: (Klamath and Lake Counties). Siskiyou pop-
ulation: CALIFORNIA: (Humboldt, Lassen, Siskiyou and Trinity Counties);
OREGON: (Klamath County). Sierra Nevada population: CALIFORNIA:
(all alpine counties from Tulare County to Lassen County); NEVADA:
(Douglas and Ormsby Counties).
Specimens Examined. Male holotype of Philanthus zebratus Cresson, fe-
male holotype of Oclocletes nitens Banks, and 423 males, 29 1 females. Of
the three populations discussed under Variation: Rocky Mountain popula-
tion 41 males, 50 females; Siskiyou population 94 males, 32 females; Sierra
Nevada population 288 males, 209 females.
SUMMARY
Certain evolutionary trends are exhibited within the zebratus group. The
most obvious is the progressive narrowing of the vertex from an F/V ratio
of 1.8 in the males of P. gloriosus to 2.8 to 3.0 in the sanbornii and zebratus
subgroups, and from 1.4 in the females of P. gloriosus to about 2.0 in the
zebratus subgroup. The size of the eyes in relation to the width of the face
increases from a H/F ratio of 1.6 in males of the gloriosus subgroup to 1.7
in sanbornii, ventralis and zebratus, and to 1.8 in basilaris. In females the
H/F ratio is 1.6 in all species except basilaris in which it is 1.7. There has
been a reduction in the density of scutal punctures with P. bid net us and P.
zebratus showing the sparsest punctation.
Within the three closely related species of the zebratus subgroup the degree
of interspecific variation is less than that between it and the remaining species
of the group; however, the differences shown in Table 1 are sufficient to
302
NEW YORK ENTOMOLOGICAL SOCIETY
justify specific recognition of the three taxa in spite of the evidence of oc-
casional gene flow or hybridization. Noteworthy also is the fact that nesting
is in dense aggregations in the case of P. zebratus but not in P. basilaris,
and males of basilaris exhibit territoriality whereas males of zebratus form
aerial swarms over the nesting site.
ACKNOWLEDGMENTS
I am grateful to the following individuals and institutions for the generous loan of the large
number of specimens used in this investigation [abbreviations as used in the text are in brackets];
D. Azuma, Academy of Natural Sciences of Philadelphia [ANSP]; D. Shpeley, University of
Alberta, Edmonton [ALB]; M. Favreau, American Museum of Natural History, New York
[AMNH]; F. G. Werner, University of Arizona, Tucson [UAZ]; F. F. Hasbrouck, Arizona State
University, Tempe [AZS]; C. Carlton, University of Arkansas, Fayetteville [ARK]; S. C. Can-
nings, University of British Columbia, Vancouver [UBC]; J. A. Powell, University of California,
Berkeley [UCB]; R. M. Bohart, R. O. Schuster, University of California, Davis [UCD]; S. I.
Frommer, University of California, Riverside [UCR]; W. J. Pulawski, California Academy of
Sciences, San Francisco [CAS]; M. S. Wasbauer, California Department of Food and Agriculture,
Sacramento [CDA]; H. E. Evans, Colorado State University, Fort Collins [CSU]; L. L. Pechu-
man, Cornell University, Ithaca, New York [COR]; L. E. Watrous, Field Museum of Natural
History, Chicago [FIELD]; L. A. Stange, Florida State Collection of Arthropods, Gainesville
[FLA]; W. F. Barr, University of Idaho, Moscow [UID]; W. E. Laberge, Illinois Natural History
Survey, Urbana [ILL]; R. E. Lewis, Iowa State University, Ames [IOWA]; G. W. Byers, Uni-
versity of Kansas, Lawrence [KAN]; R. R. Snelling, Los Angeles County Museum of Natural
History, Los Angeles [LAM]; T. D. Galloway, University of Manitoba. Winnipeg [MTB]; R.
L. Fischer, Michigan State University, East Lansing [MIS]; P. J. Clausen, University of Min-
nesota, St. Paul [MIN]; R. Blinn, University of Missouri, Columbia [UMO]; S. Rose, Montana
State University, Bozeman [MSU]; M. Hathaway, Museum of Comparative Zoology, Cam-
bridge, Massachusetts [MCZ]; B. C. Ratcliffe, University of Nebraska State Museum, Lincoln
[NEB]; E. U. Balsbaugh, North Dakota State University, Fargo [NDS]; J. D. Lattin, Oregon
State University, Corvallis [OSU]; L. E. Eighme, Pacific Union College, Angwin, California
[PUC]; R. A. Cannings, Provincial Museum, Victoria, British Columbia [PMV]; A. S. Menke,
Systematic Entomology Laboratory, % U.S. National Museum of Natural History, Washington,
D.C. [USNM]; F. D. Parker, Utah State University, Logan [USU]; R. S. Zack, Washington
State University, Pullman [WSU]; R. J. Lavigne, University of Wyoming, Laramie [UWY];
and the personal collection of A. L. Steiner, Edmonton, Alberta.
LITERATURE CITED
Armitage, K. B. 1965. Notes on the biology of Philanthus bicinctus. J. Kansas Entomol. Soc.
38:89-100.
Banks, N. 1913. New American Philanthidae. Bull. Amer. Mus. Nat. Hist. 32:421-425.
Bohart, R. M. and E. E. Grissell. 1975. California wasps of the subfamily Philanthinae. Bull.
California Insect Surv. 19:1-92.
Bohart, R. M. and A. S. Menke. 1976. Sphecid Wasps of the World. Univ. of California Press,
Berkeley, ix + 695 pp.
Burks, B. D. 1951. Tribe Philanthini. Pages 1000-1004 in: C. F. W. Muesebeck et al., Hy-
menoptera of America North of Mexico, Synoptic catalog. U.S. Dept. Agr. Monograph
No. 2, 1420 pp.
VOLUME 91, NUMBER 4
303
Cresson, E. T. 1865. Monograph of the Philanthidae of North America. Proc. Entomol Soc.
Philadelphia 5:85-132.
Cresson, E. T. 1879. “. . . descriptions of new Hymenoptera.” Trans. Amer. Entomol. Soc.
7:xxxii-xxxiv.
Dunning, S. N. 1897. Notes on Philanthus. Entomol. News 8:68-71.
Dunning, S. N. 1898. Notes on Can. Entomol. 30:152-155.
Evans, H. E. 1955. Philanthus sanbornii Cresson as a predator on honeybees. Bull. Brooklyn
Entomol. Soc. 50:47.
Evans, H. E. 1964. Further studies on the larvae of digger wasps. Trans. Am. Entomol. Soc.
90:235-299.
Evans, H. E. 1966. Nests and prey of two species of Philanthus in Jackson Hole, Wyoming.
Great Basin Naturalist 26:35-40.
Evans, H. E. 1970. Ecological-behavioral studies of the wasps of Jackson Hole, Wyoming.
Bull. Mus. Comp. Zool. 140:451-511.
Evans, H. E. and C. S. Lin. 1959. Biological observations on digger wasps of the genus
Philanthus. Wasmann J. Biol. 17:115-132.
Evans, H. E. and K. M. O’Neill. 1978. Alternative mating strategies in the digger wasp
Philanthus zebratus Cresson. Proc. Natl. Acad. Sci. 74:1901-1903.
Fox, W. J. 1890. Description of three new species of Hymenoptera. Entomol. News 1:106-
108.
Gwynne, D. T. 1978. Male territoriality in the bumblee wolf, Philanthus bicinctus (Mickel):
observations on the behaviour of individual males. Z. Tierpsychol. 47:89-103.
Gwynne, D. T. 1980. Female defence polygyny in the bumblebee wolf, Philanthus bicinctus
Mickel. Behav. Ecol. Sociobiol. 7:213-225.
Gwynne, D. T. 1981. Nesting biology of the bumblebee wolf Philanthus bicinctus Mickel.
Amer. Midland Nat. 105:130-138.
Gwynne, D. T. and K. M. O’Neill. 1980. Territoriality in digger wasps results in sex biased
predation on males. J. Kansas Entomol. Soc. 53:220-224.
Krombein, K. V. 1958. Sphecoidea. Pages 186-204 in: K. V. Krombein, Hymenoptera of
America North of Mexico, Synoptic Catalog. U.S. Dept, of Agr. Monogr. 2, 1st supple-
ment, 305 pp.
Krombein, K. V. 1967. Superfamily Sphecoidea. Pages 386-421 in: K. V. Krombein and B.
D. Burks, Hymenoptera of America North of Mexico, Synoptic Catalog. U.S. Dept, of
Agr. Monogr. 2, 2nd supplement, 584 pp.
Krombein, K. V. 1979. Sphecoidea. Pages 1573-1740 in: K. V. Krombein, Paul D. Hurd,
Jr., David R. Smith and B. D. Burks, Catalog of Hymenoptera in America North of
Mexico. V. 2:1199-2209. Smithsonian Inst. Press, Washington, D.C.
Mickel, C. E. 1916. New species of Hymenoptera of the superfamily Sphecoidea. Trans. Am.
Entomol. Soc. 42:399-434.
Mickel, C. E. 1918. New species of Sphecoidea from the central and western states. Nebraska
Univ. Studies (1917) 17:319-341.
O’Neill, K. M. and H. E. Evans. 1981. Predation on conspecific males by females of the
beewolf Philanthus basilaris Cresson. J. Kansas Entomol. Soc. 54:553-556.
Strandtmann, R. W. 1946. A Review of the North American Species of Philanthus, North of
Mexico. Ohio State Univ. Press, Columbus, 126 pp.
Received October 13, 1982; accepted July 27, 1983.
NEW YORK ENTOMOLOGICAL SOCIETY
91(4), 1983, pp. 304-311
ESTABLISHMENT OE HYLES EUPHORBIAE (L.)
(LEPIDOPTERA: SPHINGIDAE) IN THE UNITED STATES
FOR CONTROL OF THE WEEDY SPURGES
EUPHORBIA ESULA L. AND E. CYPARISSIAS L.
S. W. T. Batra
Beneficial Insect Introduction Laboratory,
Insect Identification and Beneficial Insect Introduction Institute,
Beltsville Agricultural Research Center, USDA, ARS, NER,
Beltsville, Maryland 20705
Abstract. — T\\q European spurge hawkmoth, Hyles euphorbiae (L.) was introduced for control
of leafy spurge {E. esula L.) and cypress spurge {E. cyparissias L.). It was initially established
in the United States at Chestertown, New York, on tetraploid cypress spurge and the hybrid
E. X pseudo-esula Schur. The population increased from 180 to about 1 million insects within
5 years, with total defoliation of spurge in some areas.
Cypress spurge {Euphorbia cyparissias L.), also known as yellowweed or
graveyard weed, was introduced into North America from central and south-
ern Europe by the mid-nineteenth century (Moore and Lindsay, 1953). In
North America and Europe, this plant occurs in two forms— a fertile tetra-
ploid (2n = 40) and a male-sterile diploid (2n = 20); a third form in France
is a fertile diploid (Pritchard, 1961). The seed-producing tetraploid is an
invasive weed that may occupy large areas, but it is relatively rare in North
America; the less weedy sterile diploid is often cultivated as an ornamental
(Moore and Lindsay, 1953). In the United States, stands of the fertile tet-
raploid form in New Hampshire, Massachusetts, and four counties (Franklin,
St. Lawrence, Herkimer, and Orange) in New York were reported by
Muenscher (1936).
Fertile tetraploid cypress spurge has been reported to hybridize with leafy
spurge, E. esula L. {sensu lato, 2n = 60), in Europe and Canada; these mor-
phologically variable hybrids {E. X pseudo-esula Schur, 2n = 50), produce
scant seed (Moore, 1958; Moore and Frankton, 1969).
Leafy spurge {s.l.) is a variable, invasive, fertile weed. It originated in
central and northern Eurasia, and was first recorded in North America in
1827 (Croizat, 1945; Best et al., 1980). Subsequently, it has spread through-
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VOLUME 91, NUMBER 4
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out most of the northern half of the United States and southern Canada,
with greatest impact in the upper Great Plains region (Hanson and Rudd,
1933; SellecketaL, 1962; Dunn, 1979; Best et al., 1980). Leafy spurge infests
1.2 million ha of pasture and rangeland (Sun, 1981). In New York, leafy
spurge has been recorded in 15 counties (Muenscher, 1930; Dunn, 1979),
including Orange, St. Lawrence, and Herkimer. It is possible that leafy spurge
in these latter three counties may hybridize with the tetraploid cypress spurge
known to occur there. These hybrids are known at three sites in Canada
(Moore and Frankton, 1969).
Leafy and cypress spurges possess a milky latex that may cause dermatitis
in humans and animals; the ingested fresh plants may cause illness in live-
stock (Moore and Lindsay, 1953; Best et al., 1980). Sheep graze limited
amounts of leafy spurge without distress (Johnston and Peake, 1959). The
latex contains esters of cocarcinogenic diterpene irritants (Upadhyay et al.,
1978) as well as a related antileukemic diterpenoid diester (Kupchan et al.,
1976). Control of these vigorous perennial weeds is difficult due to shoot
regeneration from deeply buried roots or root fragments, competitiveness
with desirable vegetation, and lack of natural enemies in North America.
Efforts to limit the spread and impact of leafy spurge, and to a lesser extent
cypress spurge, by biological control were initiated in Canada. After several
years of investigation, including host-specihcity testing (as summarized in
Batra, 1982), the Eurasian spurge hawkmoth Hyles {=Celerio) euphorbiae
(L.) was hrst imported in 1965 and released in Canada. This population did
not survive; of numerous subsequent North American releases, only one
established population (released in 1966-1967 on tetraploid cypress spurge
at Braeside, Ontario) is reported (Harris and Alex, 1971). Mortality is at-
tributed to predation by several genera of ants, deer mice {Peromyscus leu-
copus (Raf.)), Xyticus spiders, Polistes wasps, pentatomid bugs (Apateticus
sp.), and four genera of carabid beetles (Harris and Alex, 1971; Forwood
and McCarty, 1980a). Due to the poor survival of released H. euphorbiae,
several other Eurasian host-specihc natural enemies of leafy spurge are being
investigated.
In southern and south-central Europe, Hyles euphorbiae has two genera-
tions annually on Euphorbia spp., with E. cyparissias its preferred larval
host. Adults (Fig. 1) lay about 100 eggs in small clusters (Fig. 2) on terminal
leaves where the growing larvae feed gregariously (Fig. 3). Pupation occurs
2-15 cm below the soil surface, and the crepuscular adults visit Saponaria
officinalis L. or other flowers for nectar (Moscardini, 1947). In laboratory
tests, each larva is capable of consuming 130 linear cm of fresh cypress
spurge stem and foliage (2.7 g dry weight) according to New (1971), or 17.05
g fresh weight (3.37 g dry weight) of leafy spurge (Forwood and McCarty,
1980b).
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NEW YORK ENTOMOLOGICAL SOCIETY
Figs. 1-4. Hyles euphorbiae. 1. Adult; forewings are olive-green and tan tinged with pink,
hind wings are salmon-pink and black. 2. Bright-green eggs. 3. Gregariously feeding young
larvae; first instar larvae are greenish-black, older instars develop green, yellow and black
markings. 4. Mature larva; the aposematic markings of this conspicuous toxic-plant feeder
include orange-red on the head, horn, dorsal stripe, lateral stripes and legs, with numerous pale
yellow to white speckles and spots on a black integument.
VOLUME 91, NUMBER 4
307
HOSTS AND ESTABLISHMENT IN NEW YORK
In July 1977, 180 third and fourth instar larvae collected at Braeside,
Ontario were released within one day at Chestertown (Warren Co.), New
York. The larvae were placed by hand in scattered groups of 3-5 on dense
clusters of E. cyparissias plants, at two release points 1.6 km apart. Most
larvae began feeding immediately. When the 1977 release site near Ches-
tertown was revisited for the first time in July 1982, a thriving population
of H. euphorbiae was encountered.
The stand of fertile tetraploid E. cyparissias (Fig. 5) extends along high-
ways and adjacent nonforested areas for at least 14.4 km east-west and 28.8
km north-south, on fine-textured sand. Growing among the cypress spurge
were grasses, Asclepias syriaca L., Saponaria ojficinalis, Centaurea maculosa
Lam., and Spiraea tomentosa L. as well as roadside patches of spurge that
morphologically resembled the hybrid, E. X pseudo-esula, previously not
reported to occur in the United States (see Moore and Frankton, 1969).
These probable hybrid clones varied considerably in size and fertility. The
male parent, leafy spurge, is not recorded from Warren County, and the
nearest recorded stands are over 160 km to the south (Albany Co.). Its sticky
pollen requires insect transport (Selleck et al., 1962).
A survey of 20 random 1 m^ plots indicated that the population of H.
euphorbiae had expanded from the original 180 released to an estimated 10^
in five years. (Assuming no premature mortality, a 1:1 sex ratio, and 1
generation annually, the population could have increased to 1.13 X 10^ in
this time.) At an average density of 1 larva/m^, the insects had spread from
the two release points to a distance of 0.48 km to the east (at 1 larva/m^),
3.2 km to the west (at 0.5 larva/m^) and 1.6 km to the north (0.1 larva/m-);
the total area occupied was about 4 km^, with some 25 percent of this area
infested by cypress spurge (70-700 stalks/m^) bearing 1 larva/m^. The max-
imum average larval density encountered (12-21 larvae/m^) occurred in a
small area at one of the original release points, where the larvae were effec-
tively defoliating several 50-100 m^ patches (Fig. 6). The larvae migrated
toward, and gregariously fed at, the edges of defoliated areas (Fig. 7), reaching
a maximum density of 32 3-4 instar larvae/m^. According to calculations
based on laboratory feeding studies, 14 larvae/m^ are required to defoliate
and weaken a vigorous stand of cypress spurge (New, 1971).
Larvae of all instars were found at Chestertown between July 13 and 27,
indicating a long oviposition period. In Ontario, there is one generation
annually (New, 1971). Some (3%) of the 1 50 larvae collected at Chestertown
and kept in cages at Beltsville at about 25°C produced second generation
female adults within 4 weeks.
The reasons for the excellent reproduction of H. euphorbiae at Chestertown
and the lack of survival elsewhere in the United States remain to be ascer-
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NEW YORK ENTOMOLOGICAL SOCIETY
Figs. 5-7. Defoliation by //. euphorbiae at Chestertown. 5. Vigorous tetraploid cypress
spurge (flags are 1 m apart). 6. Completely defoliated cypress spurge patch. 7. Detail showing
7 larvae stripping remaining leaves from nearly defoliated stems.
VOLUME 91, NUMBER 4
309
tained. Some factors favoring survival may be the presence of soft, fine, well
drained sand easily entered by larvae for pupation, and the availability of
flowers such as A. syriaca, S. officinalis and Silene cucubalis Wibel. supplying
nectar for adults. Ants are considered to be major predators limiting survival
of H. euphorbiae (Harris and Alex, 1971); however, nests of ants at Ches-
tertown among the spurge were not noticeably less abundant than at other
New York release sites. Myers and Campbell (1976) observed sudden mass
attacks on Tyria jacobaeae L. larvae by Camponotus ants attracted to sap
oozing from feeding sites. Spurge flowers are visited by ants for nectar (Selleck
et al., 1962); and such foraging ants may attack young H. euphorbiae larvae,
but no ant predation was observed. Camponotus pennsylvanicus (DeGeer)
was foraging on spurge flowers and Formica sp. also was present at the
Chestertown site.
Other causes of mortality among larvae brought from Ontario and cultured
at Beltsville between 1977 and 1980 were: (1) 161 pupae buried in a field
plot were all dug out and eaten within 2 days by a raccoon {Procyon lot or
L.); (2) first instar larvae were ingested by large larvae as they ate leaves; (3)
death of caged larvae of all instars, which showed symptoms of polyhedrosis
virus infection; (4) unidentified tachinid flies were reared from larvae; (5)
Formica sp. were found feeding on a mutilated dead larva; however, when
live larvae were placed on the ants’ nest, they dislodged attacking ants by
violently squirming, regurgitating, and quickly crawling away and onto spurge
plants.
OTHER RELEASES OE Hylcs cuphorbiac
IN THE EASTERN UNITED STATES
In July 1976, third and fourth instar larvae of H. euphorbiae on cypress
spurge were collected from Braeside, Ontario. The actively feeding, appar-
ently healthy larvae were released by the author within 1-5 days of collection
as follows:
480 larvae on a large area of cypress spurge in hilltop pasture on shaly
soil near Bland (Bland Co.), Virginia; no recovery through 1982.
1,284 larvae on leafy spurge at two uncultivated brushy lots on loam
near Watertown (Jefferson Co.), New York; no recovery through 1982.
70 larvae on leafy spurge in grass along a roadside near Waddington
(St. Lawrence Co.), New York; no recovery in 1978.
100 larvae on leafy spurge in mixed weeds on loam along a railway
embankment at Calcium (Jefferson Co.), New York; no recovery through
1982.
In 1978, 70 laboratory-reared pupae were buried in sand with diploid
cypress spurge under a cage at Beltsville (Prince George’s Co.), Maryland,
but there was no recovery in 1979.
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NEW YORK ENTOMOLOGICAL SOCIETY
Additional releases of H. euphorbiae in New York were at New Milford
(Orange Co., by R. R. Hahn, 221 larvae, 1978), and by me at Keene Valley
(Essex Co., 20 larvae, 1982), Newburgh (Orange Co., 50 larvae, 1982), and
Pitcairn (St. Lawrence Co., 52 larvae, 1982).
CONCLUSION
Hyles euphorbiae may be a useful agent for the biological control of cypress
and leafy spurge at some locations; however, additional natural enemies of
these weeds are needed. Fifty years ago, this insect was considered to be an
important pest of Euphorbia lathyris L., grown as an oilseed crop in the
USSR (Malyuta, 1934). As a result of recently increased costs of petroleum,
E. lathyris is now being reinvestigated as a commercial source of petroleum
substitutes for fuel and chemical feedstocks, yielding over 1 T of oil per 2.5
ha per 7 months (Nielsen et al., 1977; Buchanan et al., 1978). The potential
benefit of H. euphorbiae for spurge control currently appears to outweigh
possible harm to spurges grown as oil crops; however, this is expected to be
influenced by changing socioeconomic conditions.
ACKNOWLEDGMENTS
I thank J. S. Kelleher, Agriculture Canada, and L. R. Batra, U.S. Department of Agriculture,
for assistance with the field work.
LITERATURE CITED
Batra, S. W. T. 1982. Biological control in agroecosystems. Science 215:134-139.
Best, K. F., G. G. Bowes, A. G. Thomas and M. G. Maw. 1980. The biology of Canadian
weeds. 39. Euphorbia esula L. Can. J. Plant Sci. 60:651-663.
Buchanan, R. A., F. H. Otey, C. R. Russell and I. M. Cull. 1978. Whole-plant oils, potential
new industrial raw materials. J. Am. Oil Chem. Soc. 55:657-662.
Croizat, L. 1945. ""Euphorbia esula'" in North America. Am. Midland Nat. 33:231-243.
Dunn, P. H. 1979. The distribution of leafy spurge {Euphorbia esula) and other weedy Eu-
phorbia spp. in the United States. Weed Sci. 27:509-516.
Forwood, J. R. and M. K. McCarty. 1980a. Control of leafy spurge {Euphorbia esula) in
Nebraska with the spurge hawkmoth {Hyles euphorbiae). Weed Sci. 28:235-240.
Forwood, J. R. and M. K. McCarty. 1 980b. Observations on the life cycle of spurge hawkmoth.
Trans. Nebraska Acad. Sci. 8:31-34.
Hanson, H. C. and V. E. Rudd. 1933. Leafy spurge, life history and habits. N. Dakota Agr.
Exp. Sta. Bull. 266:1-24.
Harris, P. and J. Alex. 1971. Euphorbia esula L., leafy spurge, and E. cyparissias L., cypress
spurge (Euphorbiaceae). Biological control programmes against inseets and weeds in
Canada, 1959-1968. Tech. Commun. No. 4, Commonw. Inst. Biol. Control, Trinidad,
67-76; 83-88.
Johnston, A. and R. W. Peake. 1959. Effect of selective grazing by sheep on the control of
leafy spurge {Euphorbia esula L.). J. Range Manage. 12:192-195.
Kupchan, S. M., I. Uchida, A. R. Branfman, R. G. Dailey, Jr. and B. Y. Fei. 1 976. Antileukemic
principles isolated from Euphorbiaceae plants. Science 191:571-572.
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Malyuta, D. A. 1934. Sphingids as pests of spurge. All Union Inst. Sci. Res. Oil Cult. Kras-
nodar, Publ. No. 7:81-84.
Moore, R. J. 1958. Cytotaxonomy of Euphorbia esu/a in Canada and its hybrid with Euphorbia
cyparissias. Can. J. Bot. 36:547-559.
Moore, R. J. and C. Frankton. 1969. Euphorbia X pseudo-esula {E. cyparissias X E. esula) in
Canada. Can. Field Nat. 83:243-246.
Moore, R. J. and D. R. Lindsay. 1953. Fertility and polyploidy of Euphorbia cyparissias in
Canada. Can. J. Bot. 31:152-163.
Moscardini, C. 1947. Osservazioni morfologico-biologiche su Deilephila euphorbiae L. Atti.
Soc. Nat. Mat. 73:210-213.
Muenscher, W. C. 1930. Euphorbia esula as a weed in New York State. Rhodora 32:100-
102.
Muenscher, W. C. 1936. The production of seed by Euphorbia cyparissias. Rhodora 38:161-
163.
Myers, J. H. and B. J. Campbell. 1976. Predation by carpenter ants: a deterrent to the spread
of cinnabar moth. J. Entomol. Soc. Brit. Columbia 73:7-9.
New, T. R. 1971. The consumption of Euphorbia cyparissias (Euphorbiaceae) by larvae of
Celerio euphorbiae {EQpi(Xop\.QV3. : Sphingidae). Canad. Entomol. 103:59-66.
Nielsen, P. E., H. Nishimura, J. W. Otvos and M. Calvin. 1977. Plant crops as a source of
fuel and hydrocarbon-like materials. Science 198:942-944.
Pritchard, T. 1961. The cytotaxonomy of the weedy species Euphorbia cyparissias L. and
Euphorbia esula L. Recent Advances in Botany (IX Internal. Bot. Congr.), Vol. 1:866-
870.
Selleck, G. W., R. T. Coupland and C. Frankton. 1962. Leafy spurge in Saskatchewan. Ecol.
Monogr. 32:1-29.
Sun, M. 1981. The purge of leafy spurge. Science 214:1 103.
Upadhyay, R. R., F. Bakhtavar, M. Ghaisarzadeh and J. Tilabi. 1978. Cocarcinogenic and
irritant factors of Euphorbia esula L. latex. Tumori 64:99-102.
Received November 17, 1982; accepted April 19, 1983.
NEW YORK ENTOMOLOGICAL SOCIETY
91(4), 1983, pp. 312-327
PATTERNS OF DISTRIBUTION AND ABUNDANCE IN
SMALL SAMPLES OF LITTER-INHABITING ORTHOPTERA
IN SOME COSTA RICAN CACAO PLANTATIONS
Allen M. Young
Invertebrate Zoology Section, Milwaukee Public Museum,
Milwaukee, Wisconsin 53233
.4 fo/racr. — Collections of Orthoptera and Dictyoptera associated with rotting organic litter
in cacao plantations at three localities in Costa Rica are described. These collections, in which
the Blattodea were the most represented, were taken from experimental, replicated treatments
of rotting discs of banana tree trunks placed on the ground, piles of natural leaf litter on the
ground, and arboreal leaf litter contained in large plastic cups suspended from the lower branches
of cacao trees. For two localities, there was one collection each for the lengthy rainy season and
the short, erratic dry season; only one rainy season collection was taken at the third locality.
The most abundant cockroach was Latiblatta sp., occupying arboreal litter samples at all three
localities followed by Eurycotis sp. which occupied both arboreal and ground litter microhabitats
at one locality only. Most of the other taxa were represented by one individual at one locality
only, suggesting very patchy distributions, assuming an adequate sample size. At one locality
(La Lola), the tettigoniid Idiarthwn hamulifemm Beier was found in arboreal litter, as was a
leaf-rolling cricket, Camptonotus sp. prob. affinis. Most orthopterans were found in litter as-
sociated with sunny areas of cacao during the dry season, a probable response to seeking refuge
in the few available moist patches, whereas numbers and densities were always lower in the
well-shaded areas in both rainy and dry periods. The arboreal leaf litter microhabitat in tropical
forests deserves particular attention for understanding the vertical stratification of cockroach
faunas.
This paper summarizes some preliminary field data on the association of
Orthoptera (Tettigoniidae and Gryllacrididae) and Dictyoptera (Blattodea)
with several kinds of rotting organic litter substrates, “microhabitats,” in
cacao plantations at three localities in Costa Rica. The information gathered
in this study was adjunct to an investigation of the occurrence of immature
stages of cacao-pollinating Diptera in these litter substrates (Young, 1982,
1983). Orthoptera and Dictyoptera are commonly associated with the cacao
plantation habitat overall, although seldom are they of any economic im-
portance (e.g., Leston, 1970). The major group discussed in this paper, the
Blattodea or cockroaches, represent only about 10 percent of the total cock-
roach fauna described for Costa Rica (see the review of Fisk, 1971). Never-
theless, the data are new in terms of both describing cacao plantations in
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VOLUME 91, NUMBER 4
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the Neotropical Region as Orthoptera and Dictyoptera habitats, and the
observed patterns of specificity for some of the forms collected.
METHODS
Between 1978 and 1980, replicated series of field “treatments” consisting
of increased abundance of three kinds of natural litter substrates, were dis-
tributed in cacao (Theobroma cacao L.) plantations at three localities or
collecting sites in Costa Rica (see Young, 1982, 1983 for descriptions of
these sites and studies). The localities are: (1) “Tineas La Tigra and El Uno,”
near La Virgen (10°23'N, 84°07^W), Heredia Province, (2) “Tinea Experi-
mental La Lola,” near Siquirres (10°06'N, 83°30'W), Limon Province, and
(3) “Turrialba or CATIE,” Turrialba (9°54'N, 83°4TW), Cartago Province.
These localities are within Lower Montane Tropical Wet Torest (Turrialba),
Premontane Tropical Wet Torest (La Tigra and El Uno), and Lowland Trop-
ical Wet Torest (La Lola) regions (Holdridge, 1 967). Each locality experiences
a short, irregular dry season each year (Tig. 1), during which the ground leaf
litter in sunny areas of cacao plantations (areas with a broken or poor canopy
cover of shade trees) becomes dry and crunchy underfoot.
The La Tigra and El Uno cacao is about 10-15 years old, with the La
Tigra plantation having a broken canopy of natural tree species from regen-
erating trees establishing very shaded conditions. The La Lola cacao plan-
tation area studied consists of the UT-29 variety (La Tigra and El Uno is
mixed varieties) shaded primarily by Erythrina and other Leguminosae, and
with one area (A) with heavy shade (shaded habitat) and another (B) with
less shade (sunny habitat) throughout the year. The La Lola cacao trees are
20-30 years old. The Turrialba plantation area studied consists of “Catongo”
variety trees with a highly variegated canopy establishing both shaded and
sunny areas. Cacao trees here are 20-30 years old.
The three kinds of organic litter substrates studied are: 2 X 2-meter ground
plots of piled leaf litter (mostly cacao leaves), 22-meter ground plots of discs
of banana tree trunks, and 200-cm“-leaf-litter-filled plastic cups suspended
in cacao trees. In addition, observations on Orthoptera, Dictyoptera, and
other insects inhabiting piles of rotting cacao pods were also made.
Tor La Tigra and El Uno cacao plantations each, there are eight replicates
of the “ground leaf litter” treatment, employing sturdy wooden frames to
hold the litter in place beneath the cacao trees. There are also eight replicates
of the rotting discs of banana tree trunks in each of the two areas, as well
as twenty plastic cups, the “arboreal leaf litter” treatment in each area. The
same distribution of treatments is also used for Areas A and B at La Lola
(Tigs. 2-4). The rotting discs of banana tree trunks treatment consists of 20-
30 discs per replicate renewed approximately every 3-4 months. The plastic
cups used for the “arboreal leaf litter” studies (Tig. 3) have perforated bot-
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SEP OCT NOV DEC JAN FEB MAR APR MAY JUN JUL
1980 1981
SUCCESSIVE MONTHS
Fig. 1 . Monthly patterns of rainfall at three cacao-growing localities in Costa Rica during
1980 and 1981. There is a depression in rainfall during the January-February period at these
localities, most evident at the La Lola and Turrialba localities. All three regions, however, are
classified as relatively non-seasonal compared to other tropical regions with a very pronounced
dry season each year.
Fig. 2. The La Lola cacao plantation, near Siquirres, in Limon Province, Costa Rica. Top:
general view of the “sunny habitat” (Area B). Below: ground litter in the shaded habitat (Area
A). Note wooden frame filled with leaf litter in the left foreground.
VOLUME 91, NUMBER 4
Fig. 3. Ground leaf litter treatment used to assess litter-inhabiting Orthoptera. Top: typical
positioning of one replicate beneath a cacao tree. Below: typical distribution of cacao leaf litter
in wooden frame. Photographs taken at La Lola during the “dry” season. Note great abundance
of leaves in the wooden frame at this time.
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NEW YORK ENTOMOLOGICAL SOCIETY
VOLUME 91, NUMBER 4
317
toms to allow drainage of rainwater. The cups are suspended by sturdy wires
from lower branches of individual cacao trees. A pattern of randomization
was used to position treatment replicates in La Lola and Turrialba. For La
Tigra and El Uno, a different system was used, one in which replicates were
placed in distinctive-appearing sub-areas. Whenever the litter substrates
were collected for sampling of insects (see Young, 1982, 1983), fresh litter
was added from the same general areas of the plantations to continue the
treatments. Further details and descriptions of the treatments are given
elsewhere (Young, 1982, 1983).
For the rotting discs of banana tree trunks (Fig. 2), prepared by slicing
freshly fallen banana trees with a machete and allowing the discs to rot for
several months, held notes were taken on the approximate numbers of insects
seen scurrying off when discs were collected in plastic bags for sorting of
insect material in the laboratory. Additional notes were taken only on an
opportunistic basis for orthopterans that scurried off in the laboratory. For
the ground and arboreal leaf litter treatments, however, virtually all of the
orthopterans that were found in the individual substrates were collected. It
is emphasized that the samples were taken during the daylight hours, a period
when many orthopterans were concealed in the litter substrates being studied.
Collected orthopterans were preserved in 70 percent ethanol for subse-
quent determinations. The litter samples were always collected by placing
the contents of a substrate container (wooden frame or cup) into a plastic
bag and tying the bag shut. Orthopterans less than 5 mm long were either
missed or not collected. For La Tigra and El Uno, there was one major
collecting period or sample, August 1980, the mid-rainy season. For La Lola
and Turrialba, however, samples were taken in both the late rainy season
(November 1980) and mid-dry season (February 1981). In addition to these
collections of Orthoptera and Dictyoptera, field observations on the occur-
rence of these insects in these substrates were made at several other times,
although systematic collections were not made.
There is a lapse of at least four months from the time any of the substrates
were initially disturbed after being set up. The substrates were sampled
several times within the time period of the present study, since they were
used to estimate the abundance of cacao-pollinating Diptera (Young, 1982,
1983, and unpubl. data). Interest in the Orthoptera focused upon the com-
parison of ground and arboreal leaf litter as microhabitats, and relative
changes in the estimated abundance of these insects in the rotting discs of
banana tree trunks between rainy and dry periods (for La Lola in particular).
RESULTS
For all three collecting sites combined, more than three times the number
of orthopteran individuals were collected from arboreal leaf litter than from
ground leaf litter in cacao plantations, although number of species in each
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NEW YORK ENTOMOLOGICAL SOCIETY
Fig. 4. Top: arboreal leaf litter replicate, with forceps for scale. Below: rotting disc of banana
tree trunk which forms a suitable microhabitat for roaches throughout the year, particularly
during the dry season. Even though such substrates represent a combined area far less than the
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319
kind of microhabitat was similar (Table 1). By far, the most abundant group,
in terms of species and numbers of individuals in samples, were members
of the Blattaria or cockroaches (Dictyoptera, Blattelidae) (Table 1). Most of
the blattids represented in the limited samples from arboreal and ground
leaf litter were nymphs, and one genus, Latiblatta, dominated the arboreal
samples (Table 1). In some instances, individual roaches jumped from the
cups when the litter samples were being collected, although this error is
estimated at less than 10 percent for all samples.
Of particular interest was the discovery of several individuals of a leaf-
rolling cricket, Camptonotus sp. prob. ajfinis Rehn, in a few of the arboreal
litter samples from Area B (sunny habitat) at La Lola (Fig. 5). This species,
along with the tettigoniid Idiarthron hamuliferum Beier (Fig. 5), were the
only two orthopteran species encountered in the study. For both species,
adults were found hiding beneath the dry leaves in the cups. Camptonotus
specimens formed tubular nests from dead, dry cacao leaves within the cups.
In addition, a single individual of /. hamuliferum was collected from its
hiding place in thick moss in the trunk of a cacao tree at El Uno during the
rainy season (July 1981). Both orthopterans were conspicuously absent, how-
ever, from cups and ground leaf litter collections at both La Tigra and El
Uno throughout the study. Furthermore, both Camptonotus and /. hamu-
liferum were absent from all arboreal and ground leaf litter samples in Area
A (shaded habitat) at La Lola for the entire study period. Thus, these forms
were most abundant, albeit small samples, from the sunny habitat at La
Lola and the mixed shade and sunny habitats in the cacao at Turrialba (the
latter for Camptonotus only).
During both rainy and dry season samples at La Lola, roaches were very
low in abundance in the arboreal leaf litter samples from Area A, the shaded
habitat. Usually only 1-2 cockroaches were seen scurrying out of individual
cups at both times, and with 60-100 percent of the cups without roaches at
all. The leaf litter in these cups, as compared to that of the more exposed
cups in Area B (sunny habitat), was always very damp and water-logged. In
the arboreal samples from the sunny habitat, however, the cups with most
individuals of cockroaches had dense layers of dry cacao leaves. During the
dry season in Area B, between 80-100 percent of the cups had 1-5 roaches
in each, whereas during the rainy season sample, only about 50 percent of
the cups had cockroaches and occupied ones with 1-2 cockroaches each. A
similar pattern was also found at Turrialba between dry and rainy season
samples.
leaf litter treatments used, densities of insects such as roaches are generally far higher in discs
than other treatments, with this effect most noticeable in the dry season in sunny cacao habitats
(La Lola).
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NEW YORK ENTOMOLOGICAL SOCIETY
Table 1. The daytime distribution and abundance of Orthoptera in arboreal ground leaf
litter in Costa Rican cacao plantations.'
Species
No. No. individuals No. individuals
localities in arboreal in ground
represented litter litter Totals
Dictyoptera, Blattellidae
Latiblatta sp.
3
26 (12A, 14N)2
0
26
Eurycotis sp.
1
7(N)
5(N)
12
Ischnoptera panamae
Hebard
1
1 (A)
0
1
Blattella gennanica (L.)
1
0
1 (A)
1
Anaplecta domestica
Saussure & Zehntner
1
0
1 (A)
1
“Undetermined Blattidae”
1
1 (N)
5 (N) (2 spp.)
6
Tettigoniidae, Pseudophyllinae
Idiarthron hamuliferum
Beier
2
2(A)
0
2
Gryllacrididae, Gryllacridinae
Camptonotus sp. prob.
ajfinis Rehn
2
5(A)
1 (A)
6
Total orthopterans by “micro-
habitat”
42
13
57
% nymphal Blattidae in samples
52.4%
76.9%
56.
No. of “species”
7
6
10
‘ Data are combined for three cacao plantation collecting sites or localities and “no. localities
represented” column indicates the localities at which specimens were collected. The three
localities are; “Fincas La Tigra & El Uno” (considered as one site); “Finca Experimental La
Lola”; “Turrialba or CATIE.” See text for descriptions of these sites. For “La Tigra & El Uno”
site, only one rainy season collection was made; for other sites, one rainy season and one dry
season collection was taken for each one.
^ Data are broken down in parentheses for numbers of adults (A) and nymphs (N) in samples.
Most of the orthopterans collected in this study were found exclusively
in the arboreal leaf litter samples, and most of these were found in the rainy
season for all three localities combined (Table 2). The most geographically
widespread form is a species of Latiblatta occurring at all three localities,
even though it is an arboreal leaf litter “specialist.” The terms “specialist”
and “generalist” species are used in this paper to refer to distributional
Fig. 5. Top: the leaf-rolling cricket Captonotus sp. prob. ajfinis Rehn. Below: the tettigoniid
Idiarthron hamuliferon after being dislodged from the leaf litter of an “artificial bromeliad”
(plastic cup) used as a replicate of arboreal leaf litter treatment. Both of these orthopterans were
VOLUME 91, NUMBER 4
321
encountered occupying arboreal leaf litter replicates in the sunny cacao habitat (Area B) at La
Lola.
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NEW YORK ENTOMOLOGICAL SOCIETY
Table 2. Locality, season, and “microhabitat” specialists and generalists' among Orthoptera
found in arboreal and/or ground leaf litter in Costa Rican cacao plantations.
Species
Exclusively
arboreal
Exclusively
ground
Generalist
in both
No. of
localities
Active season(s)
Latiblatta sp.
X
0
0
3
rainy
Eurycotis sp.
0
0
X
1 (La Lola)
rainy & dry
/. panamae
X
0
0
1 (La Lola)
rainy
B. gennanica
0
X
0
1 (La Lola)
dry
A. domestica
0
X
0
1 (Turrialba)
rainy
“Undetermined Blattidae” 0 (?)
0(?)
X
3
rainy
7. hamuliferum
X
0
0
1 (La Lola)
rainy & dry
C. sp. prob. affinis
X
0
0
2 (La Lola
& Turrialba)
rainy & dry
No. arboreal specialist species: 4
No. ground specialist species: 2
No. leaf litter generalist species: 2
No. geographically restricted species^: 5
No. geographically widespread species: 3
No. dry season specialist species: 1
No. rainy season specialist species: 4
No. of tropical season generalist species: 3
' The terms “specialist” and “generalist” are used in this paper to define probable patterns
of ecological distribution within the very limited samples obtained in this study. Similar patterns
may, or may not, exist in whole populations.
^ A geographically (regionally) restricted species is operationally defined in this paper as one
found in samples obtained for only one collecting site or locality, while widespread species are
those found in two or more such localities.
patterns within the very limited samples, and it may be difficult to extrapolate
to whole populations due to severe limitations of the samples. The data
(Table 2), while limited, suggest that there is considerable ecological diver-
sification within the Blattidae of the Neotropical Region in terms of micro-
habitat, geographical distribution, and active season.
The La Lola cacao plantation contained the greatest number of “species”
of litter-associated Orthoptera, with 4-7 species, depending upon the max-
imal number of “undetermined Blattidae” in the samples. In contrast, only
one species was found at La Tigra and El Uno, an obvious underestimate
generated in part by the very limited sampling, while 2-4 species were found
at Turrialba. In all cases, however, the data are taken as pronounced under-
estimates of the actual orthopteran faunas of cacao.
The fact that the cockroaches collected in this study may have different
ecological roles in cacao plantations is reflected in part by the morphological
differences used in noting them under field conditions. Thus Latiblatta sp.
was a “large tan roach” with nymphs either 18-22 mm long (La Tigra and
VOLUME 91, NUMBER 4
323
El Uno, and Turrialba) or 7-22 mm long for La Lola. Ewycotis sp. was
recognized as a “chocolate-brown fat cockroach” with nymphs 15-22 mm
long (La Lola), while I. panamae is a “medium size dark brown cockroach”
with adults about 13 mm long (La Lola). Blattella germanica is a “large
brown cockroach” with body size of 25 mm for the adult (La Lola), and
Anaplecta domestica is a “small dark brown cockroach” of 6-mm-long adults
(Turrialba). There are also three forms of “undetermined Blattidae” rec-
ognized in the held samples: “small dark brown” nymphs (5-8 mm long)
(Turrialba); “small sooty brown” nymphs (6 mm long) (all three localities);
“small rusty brown” nymphs (7 mm long) (Turrialba). The two orthopteran
forms are readily distinguishable in the held (Lig. 5).
Lield counts of cockroaches scurrying from the rotting discs of banana
tree trunks (Lig. 4) indicate some interesting patterns of distribution: (1)
general numbers of cockroach individuals found in discs are very similar
among the three localities or collecting sites; (2) during the dry season (La
Lola and Turrialba samples), 2-3 species are found in individual discs,
mostly nymphs within size range 4-20 mm, with 1-5 individuals per disc
and about 80% of the discs with one or more roaches; (3) during the rainy
season (all three localities), there are 1-2 species present in discs at each
locality, with 1-2 individuals (mostly nymphs) per disc, and about 30-50
percent occupancy rate of discs; (4) during the dry season in a sunny cacao
habitat (Area B at La Lola), there are many more cockroaches (1-3 species,
and 1-10 individuals per disc) in discs, with an occupancy rate of 80-100
percent, than in the nearby shaded habitat (Area A). Cockroaches are least
abundant in all disc samples during the rainy season at all localities.
During the dry season in a sunny cacao habitat (Area B at La Lola), similar
numbers of cockroaches are found in the discs and arboreal leaf litter sam-
ples, while far fewer are found in the ground leaf litter at this time. Such
differences are far less pronounced in the nearby shaded cacao, with an
overall depression of roach numbers during both dry and rainy seasons.
During the lengthy rainy season at all three cacao-growing localities, there
is a profusion of ant colonies (Young, in manuscript) associated with the
three kinds of litter microhabitats, while ant colonies are far less abundant
in these places during the dry season. Orthopterans such as Camptonotiis
and /. hamiilifenim are not found in rotting discs of banana tree trunks and
ground leaf litter throughout most of all of the year. Orthoptera in general
are far less abundant in piles of rotting cacao pod husks than they are in
ground litter throughout the year at the three localities.
The dry season impacts upon the sunny cacao habitat by a marked decline
in all arthropods on a per unit basis of ground leaf litter, while other sub-
strates, such as arboreal leaf litter and rotting discs of banana tree trunks,
may exhibit marked increases in arthropod densities.
324
NEW YORK ENTOMOLOGICAL SOCIETY
DISCUSSION
During the tropical rainy season, optimal conditions for activity in litter-
inhabiting insects may be more evenly distributed over the cacao habitat
than during the dry season, and such an effect is most pronounced in sunny
areas of cacao (see also Young, 1983, for a discussion of these effects). A
partial or broken canopy over cacao during the dry season permits the ground
litter to dry out more thoroughly and faster than in more shaded areas of
cacao. The result is an increased period of environmental thermal or mois-
ture stress to small organisms inhabiting the litter. The data in the present
paper indicate that cockroaches and large-bodied Orthoptera that occupy
leaf litter and other litter substrates (rotting discs of banana tree trunks)
during the daylight hours in cacao plantations may become more abundant
in arboreal litter microhabitats characterized by dense layers of dead leaves.
Rotting discs of banana tree trunks become ecological “refuges” for small-
bodied insects (less than 5 mm long) during the dry season, even in sunny
cacao habitats (Young, 1983). Even though cockroaches in the tropics may
be active nocturnally for feeding and courtship (e.g., Schal, 1982; Schal and
Bell, 1982), the availability of moisture-holding microhabitats in cacao hab-
itats provides a daylight shelter for hiding from thermally stressful conditions
and deleterious biological factors such as predators. Given the large diversity
of the Costa Rican cockroach fauna (Fisk, 1971), the observed patterns of
different distributions in the small number of species studied here indicate
that the group as a whole has undergone considerable ecological diversih-
cation in the Neotropical Region.
Assuming that the sample data are adequate to explain patterns of dis-
tribution in populations, it appears that most of the cacao cockroach fauna
is arboreal in terms of litter microhabitat. Tropical cockroaches are consid-
ered as major converters of decaying litter to other nutrients (e.g., Irmler
and Furch, 1979). Some species, such as Anaplecta domestica Saussure and
Zehntuer, may be ecologically flexible in occupying both arboreal and ground
litter. In the present study, this species was found in ground litter, although
the related A. mexicana Saussure occurs in epiphytes in Costa Rica (Fisk,
1971). Such species, and the very common Lattihlatta sp. may be adapted
to exploit litter and other litter-inhabiting organisms as food in natural
epiphytes such as tank bromeliads as well as in the “artificial bromeliads”
of arboreal leaf litter used in the present study. In a previous study of the
daytime-foraging of insects at experimentally-placed food baits in the lower
understory layer (at heights of 1-2 m) in lowland tropical rain forest (“Finca
La Tirimbina,” adjacent to “Finca La Tigra”) and semi-deciduous wet forest
(“Barranca Site, near Miramar, Puntarenas Prov.”) in Costa Rica during the
1975 dry season (January-February 1975 collections), a total of 5 cock-
roaches (2 species) were collected from three bait-types (pineapple, orange.
VOLUME 91, NUMBER 4
325
and grape jelly) at Barranca at 0900 hours, one on pineapple at 1 300, none
at 1500, and a total of four on three bait-types (pineapple, jelly and ground
beef) at 1 700 hours (A. M. Young, unpubl. data). Only one cockroach foraged
at one bait from 0900 to 1300 hours at La Tirimbina (Young, unpubl. data).
Such data, involving replicated series of food baits along transects in forest
understory, and in which large quantities of other insects were captured at
baits, indicate the low activity level of roaches during these periods and
times of the day.
The layers of dead leaves in the plastic cups most likely provide a suitable
hiding site for Camptonotus and I. hamuliferum during the daytime, and
feeding presumably occurs elsewhere, such as in the cacao canopy. Camp-
tonotus carolinensis (Gerstaecker) is known to feed on scale insects in North
American forests, and this species constructs rolled-leaf nests anchored with
silk, using fresh, attached leaves of the tree (Blatchley, 1920). The use of
dead cacao leaves by Camptonotus in the present study as a rolled-leaf nesting
site represents a departure from the usual habit of using leaves on the host
plant. The absence of Camptonotus sp. from the La Tigra and El Uno arboreal
litter may reflect a difference in food preferences. The occurrence of bro-
meliad-festooned shade trees in both cacao habitats may provide a suitable
dry-season refuge for these orthopterans, whereas at La Lola, at least Camp-
tonotus sp. requires the artificial bromeliads as a refuge. Whether or not
these orthopterans move each evening into the canopy trees for feeding
remains to be studied.
The marked difference in abundance of orthopterans in the small samples
between dry and rainy seasons, with most occurring in the rainy season
samples for all localities, may reflect the complex fluctuations in the abun-
dance of roaches and other foliage-dwelling insects in response to seasonal
conditions in Central America (Wolda, 1978; Wolda and Fisk, 1981).
Nymphal stages of roaches in particular exhibit marked cycles of abundance
(Wolda and Fisk, 1981).
The lower densities of Orthoptera and other litter-inhabiting arthropods
observed for cacao habitats with heavy shade and year-around moist con-
ditions (see also Young, 1983) may be due in part to the more uniformly
moist conditions of such areas and also to a greater abundance of predatory
forms such as ants under these conditions. Shaded cacao habitats tend to
accumulate a high number of ant species (Bigger, 1981). Orthoptera are more
abundant in sunny cacao habitats (Bigger, 1981), as also observed for the
Area B habitat at La Lola in the present study. Rotting discs of banana tree
trunks may exhibit less of a water-loss response to the tropical dry season
than less dense substrates such as dead cacao leaves. Under such conditions,
insects inhabiting discs, both in sunny and shaded cacao habitats, may
exhibit less of a response, in terms of density or shifts in numbers of species.
326
NEW YORK ENTOMOLOGICAL SOCIETY
than insects in leaf litter. Young (1983) found that such discs provided a
suitable refuge for the immature stages of cacao-pollinating Diptera (Cera-
topogonidae) to pass the dry season at La Lola, particularly in the sunny
habitat. Cacao leaf litter may represent a relatively more ephemeral micro-
habitat for many small-bodied insects, including the early instars of cock-
roach nymphs, in sunny cacao habitats, and particularly during the drier
months of the year. Large-bodied insects such as bigger roach nymphs, adult
cockroaches of some species, and other orthopterans (such as Camptonotiis
sp. and I. hamulifenuu), may have little difficulty in occupying the semi-
moist to dry arboreal leaf litter provided by the plastic cups in cacao trees.
But during such periods, small-bodied insects, including ants, are virtually
absent from such microhabitats. For those orthopterans that actually feed
on the leaves of cacao trees, seasonal cycles in the availability of young leaves
may greatly influence the abundance of these insects in cacao plantations
(Majer, 1975). Tettigoniidae, for example, increase in abundance with the
onset of flushes of cacao leaves in Ghana plantations (Majer, 1975). Cock-
roaches, being opportunistic feeders on a variety of food-types, may exhibit
less regular fluctuations in abundance.
The foregoing discussion considers what might be the overall trends in
relative abundance patterns, seasonal distributions, and daytime-resting places
for a relatively small portion of the Costa Rican orthopteran fauna associated
with cacao plantations. Given the type of sampling performed and the num-
ber of collection dates, the data are large underestimates of the probable
fauna resident in such habitats. Had larger samples been obtained for longer
periods of time, it might very well have been shown that individual taxa of
Blattodea exhibit very flexible behavior in terms of daytime-resting sites.
Thus, it would not be surprising to discover that most or all of the cockroach
taxa discussed in this paper are “generalist” forms in terms of hiding in both
arboreal and ground leaf litter. Yet given the tremendous vertical complexity
of tropical rain forests (e.g., Richards, 1964), it would also not be surprising
to discover that some of the taxa are, in fact, canopy or sub-canopy “spe-
cialists,” occupying large epiphytes accumulating organic litter. Such effects
must be taken into account when considering the relatively low vertical
complexity of the cacao habitat, a condition that may alter the distribution
of cockroaches that would otherwise be found in undisturbed tropical rain
forests.
ACKNOWLEDGMENTS
This research was made possible by grants from The American Cocoa Research Institute of
The Chocolate Manufacturers of America. I thank Dr. D. A. Nickle of the Systematic Ento-
mology Laboratory, U.S.D.A., for determinations of the Orthoptera. Special thanks to the
Sweetheart Corporation in Chicago, Illinois for their generous donation of the plastic cups used
as “artificial bromeliads” in this study.
VOLUME 91, NUMBER 4
327
LITERATURE CITED
Bigger, M. 1981. Observations on the insect fauna of shaded and unshaded Amelonado cocoa.
Bull. Ent. Res. 71:107-119.
Blatchley, W. S. 1920. Orthoptera of Northeastern America. Nature Pub. Co., Indianapolis,
783 pp.
Fisk, F. W. 1971. An annotated checklist of Costa Rican cockroaches (Dictyoptera: Blattaria).
Proc. Ent. Soc. Wash. 73:431-444.
Holdridge, L. R. 1967. Life Zone Ecology. Tropical Science Center, San Jose, Costa Rica,
110 pp.
Irmler, U. and K. Furch. 1979. Production, energy, and nutrient turnover of the cockroach
Epilampra innleri Rocha e Silva & Aguiar in a Central-Amazonian indundation forest.
Amazoniana 6:497-520.
Leston, D. 1970. Entomology of the cocoa farm. Ann. Rev. Ent. 15:273-294.
Majer, J. D. 1975. The pattern of leaf production in cocoa and its influence on insect distri-
bution and abundance. Cocoa Grower’s Bulletin, No. 22:18-24.
Richards, P. W. 1964. The Tropical Rain Forest. Cambridge University Press, London, 325
pp.
Schal, C. 1982. Intraspecific vertical stratification as a mate-finding mechanism in tropical
cockroaches. Science 215:1405-1407.
Schal, C. and W. J. Bell. 1982. Ecological correlates of paternal investment of urates in a
tropical cockroach. Science 28:170-173.
Wolda, H. 1978. Seasonal fluctuations in rainfall, food and abundance of tropical insects. J.
Anim. Ecol. 47:369-381.
Wolda, H. and F. W. Fisk. 1981. Seasonality of tropical insects. II. Blattaria in Panama. J.
Anim. Ecol. 50:827-838.
Young, A. M. 1982. Effects of shade cover and availability of midge breeding sites on pol-
linating midge populations and fruit set in two cocoa farms. J. Appl. Ecol. 19:47-63.
Young, A. M. 1983. Seasonal differences in abundance and distribution of cocoa-pollinating
midges in relation to flowering and fruit set between shaded and sunny habitats of the
La Lola Cocoa Farm in Costa Rica. J. Appl. Ecol. 20 (in press).
Received November 8, 1982; accepted April 19, 1983.
NEW YORK ENTOMOLOGICAL SOCIETY
91(4), 1983, pp. 328-332
COORDINATED PREY CAPTURE BY NOVOMESSOR COCKE RE ELI
(HYMENOPTERA: FORMICIDAE)
ELarold G. Fowler and Walter G. Whitford
Department of Biology, New Mexico State University,
Las Cruces, New Mexico 88003
Abstract. —Novomessor cockerelli uses coordinated behavior to subdue large orthopteran prey.
When a partially disabled grasshopper is encountered, the first worker finding it mounts the
dorsum of the grasshopper and clamps its mandibles over the wings, thus preventing escape
by flight. Workers which arrive subsequently systematically remove or subjugate additional
appendages. Behavioral coordination allows N. cockerelli to efficiently exploit large orthopteran
prey which would otherwise escape if workers attempted to capture these individually.
The genus Novomessor comprises three species. Of these, the species N.
cockerelli (Andre) is encountered commonly in the intermountain plains of
the deserts of southern North America (Wheeler and Creighton, 1934). Al-
though the generic name suggests that N. cockerelli is a harvester ant, it is
in fact omnivorous (Creighton, 1950; Chew, 1977; Whitford, 1978), and
insects comprise about one-half of its normal forage (Whitford et al., 1980).
N. cockerelli normally employs an individual foraging strategy (Whitford,
1976; Davidson, 1977) but is also capable of recruiting and cooperatively
carrying large food items (Holldobler et al., 1978). Both chemical (Holldobler
et al., 1978) and vibrational (Markl and Holldobler, 1978) signals are used
during recruitment. Here we report on cooperative and coordinated prey
capture by N. cockerelli under field conditions, paying particular attention
to the adaptive sequences of behaviors by individual foragers.
METHODS AND MATERIALS
Studies were conducted at the Jornada Experimental Range, 60 km NNE
of Las Cruces, New Mexico, from May to July, 1979. The site has been
described in detail elsewhere (Whitford et al., 1980). All studies were per-
formed from 0800-1 100 hr (MDT) during weekly visits to the site to collect
long-term data on related projects.
Two separate observational series were performed. In the first, grasshop-
pers were hand collected and placed in an active entrance to a N. cockerelli
nest. The sequence of attack by workers of N. cockerelli on the body parts
The publication costs of this article were defrayed in part by page charge payment. This article
must therefore be hereby marked “Advertisement” in accordance with 18 U.S.C. §1734 solely
to indicate this fact.
VOLUME 91, NUMBER 4
329
of the grasshopper were recorded, as well as the ability of the workers to
subjugate and capture the grasshopper. The body parts sequenced were the
wings, antennae, pro-, meso-, and metathoracic legs, the cervix of the head,
and the abdomen. All of the latter six body regions were combined and
compared with the sequence of attack on the wings, using the binomial test
(Siegel, 1956). Likewise, the ability of the workers to capture the grasshopper
was dichotomized: capture or escape.
In the second series of observations, grasshoppers were captured and teth-
ered with a fine copper wire which was passed through the body from the
pronotum to the mesosternum. The copper wire was then anchored by a
stone. The length of the copper wire from grasshopper to anchor was ap-
proximately 10 cm, allowing the grasshopper to move but not to take flight.
For each of five distances from the nest entrance (0.5, 1.0, 2.0, 4.0 and 6.0
m), 16 separate grasshoppers were tethered. The number of grasshoppers
found at each distance upwind from the nest entrance per distance class,
and the maximum number of N. cockerelli workers present at the grasshopper
during the 30 min. period were recorded. The sequence of attack by workers
on tethered grasshoppers was recorded as described previously. Only 45 of
the 80 grasshoppers offered were attacked, and only data from these suc-
cessful attacks are presented. The sequences of attack on the body parts were
recorded in inverse order, i.e., 7 = attacked first, 1 = attacked last. The num-
ber of occurrences of attack observed for each sequence was multiplied by
the rank for that sequence. By dividing this weighted sum by 315, or the
expected weighted score if that body part was always attacked first, a relative
preference for attack on each of the 7 body regions was obtained.
RESULTS AND DISCUSSION
A total of 53 grasshoppers were captured and placed at the nest entrances
of active N. cockerelli nests. Of these 53, 39 (74%) escaped. Of the 36
grasshoppers offered by this technique which were attacked, the first worker
attacking the grasshopper mounted the dorsum and clamped its mandibles
over the wings in 28 (78%) of the observations, indicating a highly significant
preference for attacking the wings (binomial test, P < 0.001, Siegel, 1956).
Moreover, all 1 4 grasshoppers which were captured by N. cockerelli workers
were first attacked by this method, as well as 14 of the 36 grasshoppers
attacked which escaped. It should be noted that only about one-quarter ( 1 4/
53) of grasshoppers presented at the entrances of active nests were captured.
The conditional probability of encounter and the maximum number of
workers present at the immobilized grasshoppers declined slightly with in-
creasing distance from the nest (Table 1). However, this reduction was not
dramatic until a distance of 6.0 m was reached (Table 1). Holldobler et al.
(1978) have demonstrated that recruitment is mediated by poison gland
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NEW YORK ENTOMOLOGICAL SOCIETY
Table I. The conditional probability of encounter and the maximum number of workers
arriving at tethered grasshoppers at various distances from the entrance of Novomessor cockerelli
nests.
Distance (m)
from nest
entrance
Conditional
probability
of encounter'
Number of workers
arriving (mean ± 1
standard deviation)^
0.5
0.6875
24.6 ± 16.5
1.0
0.5000
22.6 ± 12.3
2.0
0.6250
19.8 ± 10.6
4.0
0.5625
15.2 ± 8.5
6.0
0.4375
6.0 ± 3.7
‘ The number of grasshoppers encountered within 30 min./ 16, the total number of grass-
hoppers offered at each distance.
^ Calculated only for those grasshoppers which were encountered within 30 minutes.
secretions deposited by individual ants from the food source to the nest.
The decline in the number of workers present at the food source (grasshop-
pers) was similar to the pattern reported by Holldobler et al. (1978) to
artificial food sources.
As in the first series of experiments, the first worker which attacked a
tethered grasshopper mounted the dorsum and clamped its mandibles over
the wings in 34 of the 45 attacks observed. Likewise, a significant preference
for attacking the wings first was found (binomial test, P < 0.0001, Siegel,
1956). The metathoracic legs and the antennae were the body parts of the
grasshopper attacked most frequently sooner than the other body parts after
the wings (Table 2).
The sequence of prey capture employed by N. cockerelli is thus coordi-
nated. Moreover, the sequence of attack is apparently very adaptive. By
pinning the wings of orthopteran prey first, the prey is thus deprived of flight
as an escape mechanism. Subsequent subjugation of the jumping legs and/
or antennae then deprives the grasshopper of saltatorial escape, or flight,
directed by the antennae, if the ant subjugating the wings is dislodged.
Given our results, it is uncertain how frequently N. cockerelli may employ
this adaptive sequence of prey capture. Grasshoppers placed in the entrances
of active nests were only captured about one-quarter of the times tested. It
is unlikely that many grasshoppers would land at an active nest entrance.
More likely, N. cockerelli probably employs the sequence described in cap-
turing injured or weakened grasshoppers. Grasshoppers escaping bird strikes,
or the attacks of lizards, scorpions or other desert predators may be injured
to the extent that rapid escape is not possible. Under such conditions, N.
cockerelli, employing the adaptive sequences of behavior described, may be
able to capture these individuals.
Coordinated prey capture of orthopteran prey, organized around adaptive
VOLUME 91, NUMBER 4
331
Table 2. Observed frequency distribution of the order of attack on the body parts of im-
mobilized grasshoppers by workers of Novomessor cockerelli.
Observed order of attack
body part
1
2
3
4
5
6
7
Totals
Score*
Pin wings
34
5
1
1
4
0
0
45
0.917
Grab/clip
Antenna
2
10
11
5
16
0
1
45
0.679
Prothoracic leg
3
6
9
8
11
7
1
45
0.578
Mesothoracic leg
1
5
14
8
7
8
2
45
0.565
Metathoracic leg
4
14
8
15
3
1
0
45
0.708
Head (cervix)
0
0
2
5
3
16
19
45
0.286
Abdomen
_[
_5
_0
_3
J_
13
45
0.282
Totals
45
45
45
45
45
45
45
315
* Obtained by weighting each frequency (rank 1 = 7, rank 2 = 6, etc.) by multiplying the
frequency by its rank and then dividing by the maximum possible score, 315 (i.e., all 45
observations given rank of 1, or 45 x 7).
sequences of behavior, and the consequential cooperative removal of cap-
tured prey to the nest (Holldobler et aL, 1978), is an efficient strategy of
resource procurement, as well as a means of counteracting interference com-
petition by mass-recruiting ant species. With the exception of army ants,
termite predator specialists of the genus Leptogenys, and some of the slave-
making formicines, we know of no instance of pack-like hunting behavior
being recorded for ants. Indeed, the constancy of the attack sequences re-
ported here is similar to that observed in pack-hunting felines and canines
(Wilson, 1975).
ACKNOWLEDGMENTS
We gratefully acknowledge the initial observations of Dr. Tom Marr which led us to conduct
this series of experiments. These studies were supported by a National Science Foundation
Grant, DEB-77- 16633, to W. G. Whitford.
LITERATURE CITED
Chew, R. M. 1977. Some ecological characteristics of the ants of a desert-shrub community
in southeastern Arizona. Am. Midi. Nat. 98:33-49.
Creighton, W. S. 1950. Ants of North America. Bull. Mus. Comp. Zool. 104:1-585.
Davidson, D. W. 1977. Foraging ecology and community organization in desert seed-eating
ants. Ecology 58:725-737.
Holldobler, B., R. C. Stanton and H. Markl. 1978. Recruitment and food-retrieving behavior
in Novomessor (Formicidae, Hymenoptera). I. Chemical signals. Behav. Ecol. Sociobiol.
4:163-181.
Markl, H. and B. Holldobler. 1978. Recruitment and food-retrieving behavior in Novomessor
(Formicidae, Hymenoptera). II. Vibrational signals. Behav. Ecol. Sociobiol. 4:183-216.
332
NEW YORK ENTOMOLOGICAL SOCIETY
Siegel, S. 1956. Nonparametric Statistics for the Behavioral Sciences. McGraw Hill, New
York.
Wheeler, W. M. and W. S. Creighton. 1934. A study of the ant genera Novomessor and
Veromessor. Proc. Am. Acad. Arts Sci. 69:342-354.
Whitford, W. G. 1976. Foraging behavior in Chihuahuan Desert harvester ants. Am. Midi.
Nat. 95:455-458.
Whitford, W. G. 1978. Structure and seasonal activity of Chihuahuan Desert ant communities.
Ins. Soc. 25:79-88.
Whitford, W. G., E. Depree and P. Johnson. 1980. Foraging ecology of two Chihuahuan
Desert ant species: Novomessor cockerelli and Novomessor albisetosus. Ins. Soc. 27: 148-
156.
Wilson, E. O. 1975. Sociobiology. Belknap Press of Harvard University, Cambridge, Mas-
sachusetts.
Received June 1, 1983; accepted July 27, 1983.
NEW YORK ENTOMOLOGICAL SOCIETY
91(4), 1983, pp. 333-341
HABITAT PREFERENCES OF CARRION BEETLES IN THE
GREAT SWAMP NATIONAL WILDLIFE REFUGE, NEW JERSEY
(COLEOPTERA: SILPHIDAE, DERMESTIDAE, NITIDULIDAE,
HISTERIDAE, SCARABAEIDAE)
Paul P. Shubeck
Biology Department, Montclair State College,
Upper Montclair, New Jersey 07043
Abstract.—^ total of 2,397 individuals representing 8 species of carrion beetles (Silphidae)
and 2,336 individuals representing 4 species of “carrion beetles” from 4 other families were
collected on carrion in the Great Swamp National Wildlife Refuge during June, July and August
in 1980 and during April and May in 1981. Beetles were trapped in 3 habitats— forest, field,
and marsh. Of the 8 silphid species, 5 manifested a strong preference for a given habitat. These
species were Oiceoptoma noveboracense (forest), Necrophila americana (field), Nicrophorus
orbicollis (forest), Nicrophorus pustulatus (forest), and Necrodes surinamensis (forest). Two
silphid species manifested a slight preference for a given habitat: Oiceoptoma inaequale (field),
and Nicrophorus tomentosus (field). One species, Nicrophorus marginatus, manifested a probable
preference for the field. Insofar as other “carrion beetles” are concerned, 1 species, Onthophagus
hecate (Scarabaeidae) showed a strong preference for the field. Three species manifested slight
preferences for given habitats: Omosita colon (Nitidulidae) and Dermestes caninus (Dermesti-
dae) for the forest, and Euspilotus assimilis (Histeridae) for the field.
A search of the literature on carrion beetles indicates that there has been
increased interest and research on this group of beetles during the past quarter
century. Many of the papers that have appeared have dealt with the ecology
and behavior of species of the taxon. Conspicuous, however, has been the
lack of information on the habitat preferences of these beetles. In the 2
papers that have mentioned habitat preferences. Walker (1957) tabulated
arthropod species that were attracted to carrion-baited pitfall traps in 4
habitats— mesic forest, bottom forest, ridge forest and old field and Anderson
(1982) studied Silphidae that were collected in carrion-baited pitfall traps
in 4 very distinct habitats— deciduous forest, coniferous forest, field/mead-
ow, and marsh.
In a reeent study to determine the species composition and seasonal abun-
dance of carrion beetles in an oak-beech forest in the Great Swamp National
Wildlife Refuge (GSNWR), Basking Ridge, New Jersey, 7 species of Sil-
phidae were present (Shubeck et al., 1981). Over 98% of these silphids were
taken from early April through August. Oiceoptoma noveboracense was very
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334
NEW YORK ENTOMOLOGICAL SOCIETY
abundant from April through July with a peak in May. Necrophila americana
was most active from May through August with a pronounced peak in July.
Oiceoptoma inaequale was an early season silphid, being most active from
April through June, with a peak of activity in April. Nicrophorus orbicollis
was active from May through September and peaked in August. Although
Necrodes surinamensis was active from June through September over % of
the individuals were taken in August. Nicrophorus pustulatus was collected
in May and into September with over V2 of the season’s catch taken in June.
Nicrophorus tomentosus was taken from June through October but almost
Vi of these individuals were collected in August. Among the 55 additional
taxa also present were 4 very abundant species from other beetle families
(Nitidulidae, Histeridae, Dermestidae, Scarabaeidae).
For this study it was decided that additional information about the carrion
beetles of GSNWR might be obtained by collecting and comparing the
numbers of carrion beetles in the 3 distinct habitats found in this refuge —
forest (deciduous), field, marsh. The species of carrion beetles included in
this study were: all species of Silphidae, Derrnestes caninus (Dermestidae),
Omosita colon (Nitidulidae), Euspilotus assi mills (Histeridae), and Ontho-
phagus hecate (Scarabaeidae).
MATERIALS AND METHODS
Carrion beetles were trapped in 6 No. 10 food cans (3.78 liter), each of
which was concealed in a wooden box having 1.27 cm wire mesh at the top
and a rain cover 5 cm above the opening. These traps have been described
elsewhere (Shubeck, 1976). Two traps, 10 meters apart, were placed on the
ground in a red oak forest about 2 km northeast of the former refuge head-
quarters building. Two traps, also 10 meters apart, were placed on the ground
in an old field adjacent to the forest. This field collecting station was about
1/2 km northeast of the forest collecting station and about 100 meters from
the edge of the forest. Two traps, 10 meters apart, were individually mounted
on stakes that had been driven into the mud of a marsh adjacent to the field.
The marsh collecting station was about 400 meters east of the field collecting
station but only 5 meters into the marsh. I would have preferred to situate
the traps farther into the marsh but this was not possible because of the very
soft ooze (mud) and about 20 cm of water covering the mud from April
through July. The difference in distance (field traps situated 100 meters
from edge of forest and marsh traps situated 5 meters from the edge of field)
was not considered a problem because a previous study had shown that the
return to baited traps by carrion beetles released at 5 to 75 meters was a
result of random wandering (Shubeck, 1968). The same study showed that
the periphery of odor perception seems to be about 1 meter from carrion.
Given this information it seemed improbable that baited traps would attract
beetles from adjacent habitats.
VOLUME 91, NUMBER 4
335
Each of these 3 habitats is located in the management area which is off-
limits to visitors. The forest is dominated by red oak (Quercus rubrum), but
American beech (Fagus grandifolia) occurs on its moist fringes. This wood-
land stand is about 5 hectares in size. The field is about 2 hectares in size
and is covered by grasses about 1 meter tall. Solidago spp., Daucus carota,
and Aster sp. are also present. The marsh, about 1 00 hectares in size, contains
a variety of hydrophytes including Pontederia cordata, Typha latifolia, Pel-
tandra virginica, and Sagittaria latifolia.
One of the pair of traps situated in each habitat was baited with fish (smelt),
and the second was baited with chicken legs (drumsticks). Carrion bait in
each trap consisted of 3 “fresh” fish (about 90 g total weight) and 3 “stale”
fish (about 90 g), or 1 “fresh” chicken leg (about 90 g) and one “stale”
chicken leg (about 90 g). The fresh and stale components were individually
placed into a styrofoam cup (0.258 liter) so that each trap had a cup of fresh
carrion and one of stale carrion. These traps were initially baited with “fresh”
carrion 1 week before the season’s collecting began and on the Saturday that
collecting was begun “fresh” carrion was added to the “stale” carrion. Each
trap was serviced once per week, throughout the season, at which time the
oldest carrion (and cup) was replaced with fresh carrion (and cup), and all
beetles were collected and preserved in jars containing 70% alcohol. At all
times, therefore, each habitat had one trap baited with fish 1-7 days old
(fresh) and fish 8-14 days old (stale), and a second trap baited with a chicken
leg 1-7 days old (fresh) and a chicken leg 8-14 days old (stale). This technique
(Pirone, 1974) resulted in the presence of fairly uniform “attractive” carrion
continuously.
Weekly collections were made from 7 June to 25 August in 1980 and from
3 April to 30 May in 1981. A previous study in GSNWR had shown that
carrion beetles were most abundant during the months of April through
August (Shubeck et al., 1981).
RESULTS AND DISCUSSION
The catch for both seasons was totaled, by species for each habitat, and
the bar graphs in Figures 1 to 4 show the numbers of individuals, and the
percentage of the total for the habitat. In order to compare these results with
Anderson’s results (1982) I had to revise his percentages [Figs. 15-18] after
removing his data for the coniferous forest habitat. By doing this I was able
to compare results for deciduous forest, field, and marsh habitats in New
Jersey and Canada (Table 1). In preparing Walker’s data (1957) for com-
parison it was necessary to average his figures for the 3 deciduous forest
habitats [Fig. 7] (mesic forest, bottom forest, ridge forest), and to then work
out the percentages for species for deciduous forest versus field for a partial
comparison (Table 1).
NEW YORK ENTOMOLOGICAL SOCIETY
336
Nicroohorus orbicoll ia Nec rcde« aurinamanaia N Ic rophorus toirentoaus
Figs. 1, 2. Habitat association. 1. Oiceoptorna noveboracense {YovsXqv), Necwphila ameri-
cana (L.), Oiceoptorna inaequale (F.) — [Silphidae]. 2. Nicrophorus orbicoUis Say, Necrodes sur-
inamensis (F.), Nicrophorus tornentosus Weber— [Silphidae].
Oiceoptorna noveboracense was the most abundant species collected (1,149
individuals) and it was common in the forest and in the field but the species
preferred the forest habitat (Fig. 1). It was rarely collected in the marsh (2%
of total). Anderson (1982) also found this species most common in the
VOLUME 91, NUMBER 4
337
Table 1. A comparison, by percentage, of carrion beetles and their habitat associations in
Tennessee, Toronto, Canada, and New Jersey. Percentages rounded off to whole numbers.
Tennessee
Canada
New Jersey
For-
est
Field
For-
est
Field
Marsh
For-
est
Field
Marsh
Oiceoptoma noveboracense (Forster)
45
29
26
61
37
2
Necrophila arnericana (L.)
100
0
21
7
72
20
70
10
Oiceoptoma inaequale (F.)
—
94
0
6
42
56
2
Nicrophorus orbicoUis Say
95
5
63
12
25
86
14
0
Necrodes surinamensis (F.)
100
0
0
100
0
69
19
12
Nicrophorus tomentosus Weber
100
0
34
45
21
39
46
15
Nicrophorus pustulatus Herschel
100
0
76
24
0
100
0
0
Nicrophorus marginatus (F.)
—
0
80
20
0
100
0
Dermestes caninus Germ.
9
91
—
44
37
19
Omosita colon (L.)
70
30
57
33
10
Euspilotus assimilis (Payk.)
94
6
43
56
1
Onthophagus hecate Panz.
100
0
9
79
12
deciduous forest and least common in the marsh. Although least common
in the marsh 26% of the individuals taken were, in fact, collected in this
habitat. It should be noted that Anderson’s description of his marsh indi-
cated that it . . underwent seasonal inundation, with water accumulating
in the spring or after heavy rainfall.” It seems clear, therefore, that it was
relatively dry part of the time and may have superhcially resembled an old
field. Walker did not collect this species in his Tennessee study (1957).
Necrophila arnericana, the second most abundant species (927) collected
in GSNWR strongly preferred (70%) the field habitat yet it was somewhat
common (20%) in the forest and less common (10%) in the marsh (Fig. 1).
Anderson’s results were virtually identical for his deciduous forest habitat
but he collected 72% of this species in his marsh and 7% in his field (Table
1). If one compares the New Jersey and Canadian data for this species in
terms of forest versus field and marsh (combined) the results are virtually
identical. The Tennessee data are completely different— all of the 162 in-
dividuals were collected in the 3 forests and none was taken in the field
(Table 1).
Oiceoptoma inaequale, with 165 individuals collected, was the third most
abundant silphid species in Great Swamp. Although it slightly preferred the
field it was, in fact, common in both forest and field but rarely taken in the
marsh (Fig. 1). Anderson found that the bulk (94%) of the individuals col-
lected were taken in the forest, none in the field and 6% in the marsh (Table
1). Walker did not find this species in his study.
The fourth most abundant species in this study was Nicrophorus orbicoUis
and it showed a strong preference for the forest (86%) over the field (14%)
338
NEW YORK ENTOMOLOGICAL SOCIETY
(Fig. 2). Results of the studies in Tennessee and Canada were somewhat
similar since Walker collected 95% of his individuals in the forest and 5%
in the field, and Anderson took 63% of his individuals in the forest, 12% in
the field, and 25% in the marsh (Table 1).
The fifth most abundant silphid species, Necrodes surinamensis, was in
fact not abundant (Fig. 2). Of the 1 6 individuals taken, % were collected in
the forest and the remaining ‘A about equally divided between the field and
marsh. Although Walker did not take a single individual in his field, he
collected 543 individuals in his deciduous forests (Table 1). Anderson took
7 individuals in his field (Table 1) (plus 4 others in his coniferous forest).
In spite of the fact that only 13 individuals of Nicrophorus tomentosus
were taken in New Jersey but 1,488 were collected in Anderson’s study
(1982), a remarkable similarity in the forest : field : marsh percentages was
evident (39:46:15 in New Jersey and 34:45:21 in Canada) (Table 1). On the
other hand, all 5 1 individuals taken by Walker were collected in the 3 forests.
Over 20 years of carrion beetle field studies have made it quite obvious to
me that this is the most active Nicrophorus species of the 4 I have observed.
It is the one that I would expect to have the widest range in its random flight
(Shubeck, 1968) as it searches for carrion. This was supported by the data
from both the New Jersey and Canadian studies.
Nicrophorus pustulatus, the seventh silphid species in order of abundance
numbered but 8 individuals (Fig. 3), all from the forest. Walker’s data were
also limited to a few individuals (13) and they were all taken in the forest
habitats (Table 1). Anderson’s data too, were based on a small sample (17)
and it indicated a preference of 3:1, forest : field (Table 1). In spite of the
small sample in each case the preference of this species for the forest habitat
is consistently clear.
The least abundant silphid in GSNWR was Nicrophorus marginatus which
was limited to 1 individual (Fig. 3). However, it was taken in the open field
like Anderson’s sample of 125 individuals which showed a preference for
the field over the marsh by a 4:1 ratio (Table 1). Although this species was
not taken in the Tenneessee study, 1 individual was taken in a field but in
no other habitat in Maryland (Shubeck, unpublished data, 1981).
The remaining 4 species are not members of the family Silphidae but they
are members of 4 other families which contain species associated with silphid
species on carrion. None of these species was included in Anderson’s study
but they were included in the Tennessee study.
Dermestes caninus (Dermestidae) was the most abundant (77 1 individuals)
non-silphid species collected (Fig. 3). It was common in all 3 habitats but
slightly preferred the forest (44%) over the field (37%). A substantial per-
centage ( 1 9%) of individuals was also taken in the marsh. This species is a
good flyer and apparently ranges widely in search of dry carrion. The majority
of individuals (91%) in Walker’s study was collected in the field (Table 1).
VOLUME 91, NUMBER 4
339
Dwosite cclon Euscilotus essi'"ili8 pntSophe.aus ^ecate
Figs. 3, 4. Habitat association. 3. Nicrophorus pustulatus Herschel, Nicrophorus marginatus
(F.)— [Silphidae], and Dennestes caninus Germ. — [Dermestidae]. 4. Omosita colon (L.)— [Ni-
tidulidae], Euspilotus assimilis (Payk.)— [Histeridae], Onthophagus hecate Panz. — [Scarabaei-
dae].
Omosita colon (Nitidulidae) was also taken in large numbers (723). Al-
though this species was present in the marsh (10%) and common in the field
(33%), it showed a slight preference for the forest (57%) (Fig. 4). Walker also
found in his study that the species preferred the forest to the field in a 7:3
ratio (Table 1).
340
NEW YORK ENTOMOLOGICAL SOCIETY
A total of 661 individuals of the species Euspilotus assimilis (Histeridae)
was collected in GSNWR. The species was rarely taken in the marsh (less
than 1%) and it was common in the forest and field but slightly preferred
the latter (Fig. 4). The overwhelming majority (94%) of Walker’s specimens
in Tennessee was taken in the forest (Table 1).
The last species included in this study is Onthophagus hecate (Scarabaei-
dae). Although it is called a ‘‘dung” beetle, it is, in fact, found on dung and
carrion (Arnett et al., 1980). About Vio of the 181 individuals collected were
taken in the forest and a comparable number in the marsh, but the majority
(79%) was taken in the field (Fig. 4). All of the individuals of this species
collected in Tennessee by Walker were taken in the forest (Table 1).
CONCLUSIONS
Of the 8 silphid species collected in 3 habitats in GSNWR, 5 had a strong
preference for 1 habitat over the other 2 (more than 60% of the species’
representatives were taken in the preferred habitat). These were Oiceoptoma
noveboracense, Necrophila americana, Nicrophorus orbicollis, Nicrophorus
pustulatus, and Necrodes surinamensis.
Two silphid species had a slight preference for 1 habitat over the other 2
(56% or 46% of the species’ representatives were taken in the preferred
habitat). These species were Oiceoptoma inaequale and Nicrophorus tomen-
tosus.
One silphid species manifested a probable preference for 1 habitat. Nicro-
phorus marginatus, although limited to 1 specimen was taken in the same
habitat (field) in this and 2 other studies.
Insofar as the carrion beetles from other families are concerned 1 species,
Onthophagus hecate (Scarabaeidae), showed a strong preference for 1 habitat
(79% of the species’ representatives were taken in the preferred habitat).
Three species showed a slight preference for 1 habitat over the other 2 (44%,
56%, or 57% of the species’ representatives were taken in the preferred
habitat). These species were Dermestes caninus (Dermestidae), Euspilotus
assimilis (Histeridae), and Omosita colon (Nitidulidae).
ACKNOWLEDGMENTS
I would like to thank Mr. John L. Fillio, Refuge Manager of GSNWR for permission to work
in the Swamp. Mr. Theodore W. Gutzke, Assistant Refuge Manager, provided general infor-
mation when called on for assistance.
LITERATURE CITED
Anderson, R. S. 1982. Resource partitioning in the carrion beetle (Coleoptera: Silphidae)
fauna of southern Ontario: ecological and evolutionary considerations. Can. J. Zool. 60:
1314-1325.
VOLUME 91, NUMBER 4
341
Amett, Jr., R. H., N. M. Downie and H. E. Jaques. 1980. How to Know the Beetles, 2nd
edition. Wm. C. Brown Co., Publishers, Dubuque, Iowa, 461 pp.
Pirone, D. 1974. Ecology of necrophilous and carpophilous Coleoptera in a southern New
York woodland (phenology, aspection, trophic and habitat preferences). Ph.D. Thesis,
Fordham University, New York, New York, 769 pp.
Shubeck, P. P. 1968. Orientation of carrion beetles to carrion; random or non-random? J.
New York Entomol. Soc. 76:253-265.
Shubeck, P. P. 1976. An alternative to pitfall traps in carrion beetle studies (Coleoptera). Ent.
News 87:176-178.
Shubeck, P. P., N. M. Downie, R. L. Wenzel and S. B. Peck. 1981. Species composition and
seasonal abundance of carrion beetles (Coleoptera) in an oak-beech forest in Great Swamp
National Wildlife Refuge, N.J. Ent. News 92:7-16.
Walker, T. J. 1957. Ecological studies of the arthropods associated with certain decaying
materials in four habitats. Ecology 38:262-276.
Received February 15, 1983; accepted September 27, 1983.
NEW YORK ENTOMOLOGICAL SOCIETY
91(4), 1983, pp. 342-347
MICROCLIMATE OBSERVATIONS AND DIEL ACTIVITIES
OF CERTAIN CARRION ARTHROPODS IN
THE CHIHUAHUAN DESERT
Kenneth Schoenly
Department of Biology, Angelo State University,
San Angelo, Texas 76909
Abstract. — Die! activity cycles of carrion arthropods and microclimate in and around carcasses
were recorded using a bait trap and electronic telethermometer apparatus, respectively. Ar-
thropod collections and temperature data were gathered for 24 hours each on 2 collection dates
in 12 consecutive 2-hr intervals. Ants, histerid and dermestid beetles were active throughout
the 24 hr period. Temporal segregation was observed between diurnal muscoid flies and noc-
turnal trogid beetles. Small numbers of maggots emerged from carcasses between 1 AM and 9
AM MDT. In the microclimate experiments, highest temperatures were noted on the soil surface
(50°C), whereas the carcass surface (47°C), carcass interior (42°C), and surrounding air tem-
peratures (36°C) were notably lower. Highest mean temperatures were recorded in the carcass
interior and were probably attributable to internal heat generation by bacteria and maggots.
Most previous studies on carrion arthropods have addressed aspects of
insect phenology and succession, trophic relationships and stages of decom-
position. Other studies have examined the influence of environmental factors
on rate of decomposition (Nabaglio, 1973), arthropod succession (Payne,
1965), flight activities of carrion beetles (Shubeck, 1975), and carcass mi-
croclimate (Reed, 1958; Payne, 1965). However, diel activities among car-
rion arthropods relative to microclimates in and around carcasses have
received only casual attention. Diel variations in the carrion fauna (Reed,
1958; Payne, 1965; Shubeck, 1971) indicate that some arthropods (histerids,
ants, silphids, phalangids, and dipterous larvae) that are active on carcasses
by day also are nocturnal; whereas other taxa are almost exclusively diurnal
(adult dipterans) or nocturnal (trogid beetles).
This paper supplements an earlier study (Schoenly and Reid, 1983) on
the community structure of carrion arthropods in the northern Chihuahuan
desert. Here I describe microclimate and diel activity patterns of arthropods
on rabbit carrion during peak periods of arthropod diversity and carcass
decomposition.
materials and METHODS
The study area was located in a desert shrub community adjacent to the
Franklin Mnts in El Paso County, Texas. In this region of the Chihuahuan
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must therefore be hereby marked “Advertisement” in accordance with 18 U.S.C. §1734 solely
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VOLUME 91, NUMBER 4
343
desert, mean annual precipitation is 21 1 mm, temperatures range from -21°
to 43°C, and the growing season averages 238 days (Norquest, 1941).
Observations of diel activity and carcass microclimate were made using
a bait trap and telethermometer apparatus. The trap consisted of a square
wooden box and was designed to collect arthropods immigrating to and
emerging from carcasses (Schoenly, 1981). Arthropods were collected from
the trap with a system of eight killing jars charged with an ethylene glycol
mixture (Morrill, 1975). Adult blacktailed jackrabbits {Lepus californicus
Gray) with an average mass of 2,250 gm were used for bait and placed in
the trap. Sampling was begun when maximum arthropod diversity was
reached; a period of 4-6 days after carcass placement (Schoenly and Reid,
1983). Arthropod collections and temperature data were gathered for a
24 hr period during 12 consecutive 2-hr intervals. Two experiments, using
one rabbit carcass each, were conducted on July 26-27, and August 25-26,
1980. Penetration and contact thermocouples were attached to the carcass
to monitor internal and surface temperatures. Ambient air and soil surface
temperatures were monitored by securing thermocouples to the top and
bottom of a 1 -meter vertical post (Fig. 1). A Yellow Springs Instruments
telethermometer was used to measure temperature. To maximize sampling
efficiency, collecting jars were changed with fresh preserving fluid during
each visit.
Patterns of temporal utilization among carrion arthropod taxa were com-
pared using Levins (1968) measure of niche breadth:
B, = 1/2 p./
j
where py is the importance value (proportion of individuals) of the ith species
found on j resource units. The resource units in this study refer to time of
carcass visitation by arthropods; thus, there were 12 2-hr resource units.
Niche breadth values range from 0 to 1. A value of 1 indicates carrion
utilization on all 1 2 resource units, whereas a value approaching 0 indicates
very restricted use of carrion. Correlation statistics were used to test for
significant differences between variables.
RESULTS AND DISCUSSION
Activity distributions of 5 arthropod taxa and temperature records pooled
from the 2 collection dates are summarized in Figure 2. Although a brief
summer shower temporarily interrupted arthropod activity on July 26, species
composition overlapped considerably in both trials. Arthropods collected
from the July and August experiments included: dipterans (97 and 268),
hister beetles (18 and 109), ants (36 and 30), trogid beetles (4 and 10) and
dermestids (3 and 17). The only other notable difference between the 2
experiments was the absence of maggots in the July trial.
344
NEW YORK ENTOMOLOGICAL SOCIETY
Fig. 1 . Diagrammatic representation of the bait trap and telethermometer apparatus used
for recording microclimate data and diel activity patterns of arthropods frequenting rabbit
carrion. The height of the vertical post is 1 meter. A = ambient air, S = soil surface, C3 = carcass
surface, and C, = carcass interior thermocouples.
Adult dipterans were collected from the trap from 9 AM to 9 PM, and
no activity occurred after 9 PM (Fig. 2A). A relatively high niche breadth
value (0.754) corresponded to a uniform pattern of diurnal activity (1-7
PM) and a large density of individual flies, particularly from 9 to 1 1 AM.
The relationship between diurnal activity of adult flies and air temperature
was significant and negative (r = —0.62, P < 0.05). The calliphorid, Cochlio-
myia macellaria (Fabricius) was the most abundant species observed (N =
329), followed by Musca domestica L. (Muscidae) (24) and the sarcophagid,
Blaesoxipha plinthopyga (Wiedemann) (12). In addition, small numbers of
Fig. 2. (A-E) Number of arthropods collected from jackrabbit carrion during periods of
active decomposition (days 4-6 after placement) in summer, 1 980. Numbers in the right margin
of each graph refer to values of temporal niche breadth (see text). Shaded portions indicate
nocturnal periods. L = fly larvae. (F) Microclimate temperatures taken on the carcass and near
the trap; S = soil surface, A = ambient air, Q = carcass surface and Q = carcass interior tem-
peratures. Partial sun symbols indicate approximate periods of sunset and sunrise (MDT).
TEMP(*C) NUMBER OF INDIVIDUALS
VOLUME 91, NUMBER 4
345
200
0
A) OIPTERANS
N-365
1
L
CocMiomvio mocellorio 0.754
Musco domestfco
BloeSQXiphg Rjinthgpy.qg
Soprinui discoidolis
0.900
20F
C) FORMICIDS
N-66
Conorwvrmo bicolor
(Prfmflypgflftfr
0.870
1— C
ir tloro
f
10 •
0) TR06IOS
N-14
O*
Irgi fuborotut 0.660
' E)
OERMESTIOS
Dtrmtottt mormoratut 0 82i
N-20
f 1
» \
» 1 ■ » ■ ■ I ■ ■ ■ . 1 . . ^ ,
346
NEW YORK ENTOMOLOGICAL SOCIETY
dipteran larvae (N = 22) emerged from earcasses at night between 1 and 7
AM and continued into late morning (Fig. 2A).
The histerid beetle, Saprinus discoidalis LeConte, was second in abun-
dance (N = 127) and had the highest niche value of all the taxa considered
(0.900). This species was active throughout the day and night, but had a
distinct peak of activity 2 hr before sunset (Fig. 2B).
Of the ants observed, Crematogaster clara Mayr and Conomyrma bicolor
(Wheeler) were the most frequent visitors to carrion. Ants were least active
during daylight hours, whereas nocturnal observations showed a rapid in-
crease in numbers beginning at 1 1 PM with a prominent peak 1 hr before
sunrise (Fig. 2C). Ant activity was negatively correlated with increasing air
(r = —0.63, P < 0.02) and soil temperatures (r = —0.56, P < 0.05). A sig-
nificant positive relationship was observed between nocturnal ant activity
and dipteran larvae emergence from carrion (r = 0.93, P < 0.01) suggesting
that ants were preying on the larvae. Predation by ants on larvae in carrion
has been reported previously from the Chihuahuan desert (McKinnerney,
1978; Schoenly and Reid, 1983) and elsewhere.
Trogid beetles {Trox suberosus Fabricius) were collected in small numbers
(N = 14) during nocturnal periods only from 9 PM to 3 AM (Fig. 2D), and
had the lowest value of temporal niche breadth (0.660). During the heat of
the day, trogids remained inside or under carcasses where subdued light
conditions prevailed.
A total of 20 dermestid beetles {Dermestes marmoratus Say) were collected
between 7 AM and 3 AM. Eight or 40% of the individuals collected were
captured in the 2 hr between 7 AM and 9 AM (Fig. 2E).
Smaller numbers of other arthropods (not shown in Fig. 2) also were
collected including 3 solpugids (Eremobates marathoni Muma), 2 arachnids
{Syspira longipes [Simon]), and a staphylinid beetle (Creophilis maxillosus
[Linne]).
Highest temperatures were recorded on the soil (50°C) and carcass surfaces
(47°C) between 1-3 PM, whereas carcass interior and ambient air temper-
atures peaked later at lower temperature (42° and 36°C, respectively). How-
ever, the mean temperature of the interior of carcasses (35°C) was much
higher than the mean temperature of the carcass surface (31°C) and the
surrounding soil surface temperatures (30.6°C) (Fig. IF). Payne (1965) stated
that rising temperatures in the carcass during active and advanced decay
stages are attributable to the actions of bacteria and maggots. My data would
seem to support this hypothesis.
My results indicated that diel variations in the desert carrion fauna are
similar to those reported by Reed (1958) and Payne (1965) in temperate
forest ecosystems. Higher environmental temperatures do not appear to
restrict arthropod activity in or around carrion, at least in those taxa studied
here. Temporal segregation was noted in 2 taxa (flies, trogids) and densities
VOLUME 91, NUMBER 4
347
of all 5 taxa peaked at different times. Analysis of feeding habits revealed
that even among ecologically similar carrion taxa patterns of diel activity
differed. Among the necrophagous taxa, trogid beetles were exclusively noc-
turnal, whereas dipterans and dermestids were both active after sunrise but
displayed distinct peaks of activity at different times (9-11 AM and 7-9
AM, respectively). Of the predaceous arthropods, hister beetles and ants
both are known to prey on maggots (Schoenly and Reid, in press and ref-
erences therein), however, peak densities occurred at dusk for beetles and
dawn for ants. Members of the carrion community in other regions may
show similar patterns in carcass utilization.
ACKNOWLEDGMENTS
I thank R. J. Gagne, A. C. F. Hung, J. M. Kingsolver, Systematic Entomology Laboratory,
USDA; P. Vaurie, L. H. Herman, American Museum of Natural History; V. Roth, R. S. Beal,
Jr., and M. Muma for their determinations.
LITERATURE CITED
Levins, R. 1968. Evolution in Changing Environments. Some Theoretical Explorations.
Princeton Univ. Press.
McKinnemey, M. 1978. Carrion communities in the northern Chihuahuan desert. Southwest.
Nat. 23:563-576.
Morrill, W. L. 1975. Plastic pitfall trap. Environ. Entomol. 4:596.
Nabaglio, L. 1973. Participation of invertebrates in decomposition of rodent carcasses in
forest ecosystems. Ekologia Pol. 21(18):251-270.
Norquest, C. E. 1941. Climate of Texas. Pages 1129-1146 in: Climate and Man, Yearbook
of Agriculture. U.S.D.A., Washington, D.C.
Payne, J. 1965. A summer carrion study of the baby pig Sus scwfa Linnaeus. Ecology 46:
592-602.
Reed, H. B., Jr. 1958. Study of dog carcass communities in Tennessee, with special reference
to the insects. Am. Midi. Nat. 59:213-245.
Schoenly, K. 1981. Demographic bait trap. Environ. Entomol. 10:615-617.
Schoenly, K. and W. Reid. 1983. Community structure of carrion arthropods in the Chihua-
huan desert. J. Arid Environ. 6:253-263.
Shubeck, P. P. 1971. Diel periodicities of certain carrion beetles (Coleoptera: Silphidae).
Coleopt. Bull. 25:41-46.
Shubeck, P. P. 1975. Flight activities of certain carrion beetles: Silpha noveboracensis, Sta-
phylinidae, Histeridae. The William L. Hutcheson Memorial Forest Bull. 3(2):40-43.
Received April 25, 1983; accepted August 17, 1983.
NEW YORK ENTOMOLOGICAL SOCIETY
91(4), 1983, pp. 348-354
SEASONAL DYNAMICS OF FLEAS ASSOCIATED WITH THE
GRAY-TAILED VOLE, MICROTUS CANICAUDUS MILLER,
IN WESTERN OREGON*
Richard G. Robbins
Department of Entomology, Smithsonian Institution,
Washington, D.C. 20560
Abstract. — ^ study of the population dynamics of fleas associated with the gray-tailed vole,
Microtus canicaudus Miller, was conducted on three sites surrounding the city of Corvallis, in
Oregon’s Willamette Valley. Over a period of 12 months, 22,641 adult and larval fleas repre-
senting eight species were recovered from 377 comparable voles and 256 nests. On all sites,
adult and larval flea populations experienced spring and early winter peaks followed by drastic
summer and midwinter declines. These fluctuations are consistent with the thesis that humidity
and temperature are the chief factors influencing flea populations.
This is the third in a series of papers on the fleas that parasitize the gray-
tailed vole, Microtus canicaudus Miller, the principal species of field mouse
inhabiting grassy lowlands between the Cascade and Coast Ranges of western
Oregon and Washington. Earlier, the fleas of this vole served to illustrate
the use of statistics for measuring interspecific associations of vertebrate
ectoparasites (Faulkenberry and Robbins, 1980) and for modeling parasite
populations (Robbins and Faulkenberry, 1982). By contrast, this and suc-
ceeding papers will be confined to an analysis of flea populations specific to
the gray-tailed vole and its immediate mammalian associates.
DESCRIPTION OF THE STUDY SITES
The three sites selected for this study lie north, south, and west of the city
of Corvallis, Oregon, which itself is situated west-centrally in the Willamette
Valley, a rich agricultural region characterized by warm, dry summers and
cool, wet winters. Since the sites are ecologically and physiographically sim-
ilar to one another, data obtained from any one site were compared with
data from the other two which, in effect, served as controls. Additional
criteria used in site selection were accessibility, relative freedom from human
interference, and the presence of significant populations of Norway rats
{Rattus norvegicus (Erxleben)) and house mice (Mus musculus Linnaeus).
‘ Published by permission of the Chairman, Department of Entomology, Smithsonian Insti-
tution, Washington, D.C. 20560.
The publication costs of this article were defrayed in part by page charge payment. This article
must therefore be hereby marked “Advertisement” in accordance with 18 U.S.C. §1734 solely
to indicate this fact.
VOLUME 91, NUMBER 4
349
Each site is bounded by larger areas of similar terrain permitting free move-
ment of animal populations. In addition to the gray-tailed vole, Norway rat
and house mouse, mammals found on all three sites include the vagrant
shrew (Sorex vagrans Baird), Townsend mole {Scapanus townsendii (Bach-
man)), deer mouse {Peromyscus maniculatus (Wagner)), Townsend vole {Mi-
crotus townsendii (Bachman)), and the introduced Eastern cottontail rabbit
{Sylvilagus floridanus (Allen)). While various species of grasses account for
most of the vegetative cover, larger plants common to all sites are sweetbriar
rose (Rosa eglanteria Linnaeus), Himalaya berry (Rubus procerus Mueller),
Northwest nettle (Urtica lyallii Watson), Queen Anne’s lace (Daucus carota
Linnaeus), and Douglas’ hawthorn (Crataegus douglasii Lindley).
The north Corvallis collecting site, hereafter referred to as the North Site,
occupies approximately 1.3 hectares of land within the apex of the inverted
isosceles triangle formed by the tracks of the Southern Pacific Railroad at
the Corvallis Junction. During winter, standing water may persist for long
periods in ditches that parallel the tracks. Tall oat-grass (Arrhenatherum
elatius (Linnaeus) Mertens) is the dominant grass species, covering most of
the site and broken only occasionally by clumps of orchard-grass (Dactylis
glomerata Linnaeus). Because part of this area is used as a public dumping
ground, Norway rats and house mice are most abundant here. Being moist,
the North Site also supports the largest population of Townsend voles.
Extensive real estate development along the southern edge of Corvallis
necessitated several minor shifts in the location of the South Site and effec-
tively curtailed collecting during two months of this study (May-June 1 973).
However, all southern sites were located immediately east or west of U.S.
Route 99W between Millrace Creek and Wake Robin Avenue, and no site
exceeded 0.8 hectares in area. Again, tall oat-grass is the dominant grass
species, but orchard-grass and a variety of weeds are also abundant. The
South Site is only slightly drier than the North Site.
The West Site embraces 0.6 hectares of land along the north side of
Philomath Boulevard 1.3 kilometers west of its junction with 53rd Street.
Both tall oat-grass and orchard-grass are absent here, their place being taken
principally by colonial bent-grass (Agrostis tenuis Sibthorp), followed by
velvet-grass (Holcus lanatus Linnaeus), bristly dog’s-tail grass (Cynosurus
echinatus Linnaeus), meadow fescue (Festuca elatior Linnaeus), and small,
scattered clumps of tufted hair-grass (Deschampsia caespitosa (Linnaeus)
Palisot de Beauvois). On this relatively dry site, the Eastern cottontail and
Townsend mole are conspicuously abundant.
SAMPLING TECHNIQUES
Between December 1972 and January 1974, gray-tailed voles and their
nests were collected at regular monthly intervals on all three sites. Initially,
voles were taken by placing Sherman all-metal live traps (H. B. Sherman,
350
NEW YORK ENTOMOLOGICAL SOCIETY
Table 1. Composition of the adult flea population of the gray-tailed vole.
Flea species
Nests
No.
Percent
No.
Hosts
Percent
Atyphloceras nniltidentatus (C. Fox) 1909
2,550
41
100
20
Catallagia charlottensis (Baker) 1898
3,291
53
252
49
Corrodopsylla curvata (Rothschild) 1915
32
0.5
1 1
2
Hystrichopsylla occidentalis Holland 1949
212
3
25
5
Monopsy/lus wagneh (Baker) 1 904
36
0.5
44
8.5
Nosopsyllus fasciatus (Bose) 1800
71
1
14
3
Peromyscopsylla selesis (Rothschild) 1 906
57
1
62
12
Rhadinopsylla sp.
1
0
3
0.5
Totals
6,250
100
511
100
De Land, Florida) in their runways; however, the winter of 1972-1973 was
so cold that most animals captured in this manner died before they could
be retrieved. Persistent vandalism, which must be expected when working
in a suburban environment, also discouraged this approach. In place of traps,
a variety of old boards and panels were scattered at random over each site.
Voles seeking shelter under these objects were easily caught by hand. Each
vole was then immediately transferred to a large, labeled plastic jar con-
taining fresh grass clippings and pieces of fruit. All jars were fitted with wire
mesh lids. Because some voles died or injured themselves in transit or were
found naturally injured in the field, not all of them could be used in this
study.
Ordinarily, the gray-tailed vole constructs its nest in a chamber located
1 5-30 centimeters below the surface of the ground (Pearson, 1972); however,
should objects be present at the surface the vole will also build under these.
Only surface nests were routinely collected for this study, though during the
summer of 1 973 five subterranean nests were successfully exhumed for com-
parative purposes. As most fleas require about one month to develop, an
eflbrt was made to collect only those nests that had been occupied for at
least this time. Each nest and the loose soil below it was swiftly transferred
to a labeled, half-gallon Freezette-Flat plastic container (Cole-Parmer In-
strument and Equipment Company, Chicago, Illinois) sealed with an airtight
lid. If a litter was found, it was removed and notes were taken on the age
and number of the young.
In the laboratory, all voles were killed by quickly wrapping them in cotton
blankets saturated with chloroform. This technique prevented the escape of
any ectoparasites and preserved them in the positions they occupied on their
host’s body while it was alive. Once dispatched, each vole was placed in a
labeled plastic bag and stored at — 1 6°C. The nest fauna was extracted with
VOLUME 91, NUMBER 4
351
Berlese funnels set for 72 hours and equipped with 60- watt Ken-Rad light
bulbs. All fleas and their larvae were preserved in labeled vials of 75% ethyl
alcohol.
At irregular intervals, groups of 1 2 voles were removed from cold storage,
thawed, and examined for fleas. Though a number of washing and dissolution
techniques have been described for the collection of other ectoparasite groups
(Hopkins, 1949, pp. 395-398; Lipovsky, 1951; Cook, 1954; Henry and
McKee ver, 1971), these are unsatisfactory for fleas, which adhere tenaciously
to the hairs of their host and are easily damaged by caustic chemicals. During
this study, all fleas were recovered by vigorous brushing and careful search-
ing, a process that generally required half an hour per animal. To prevent
ectoparasite loss, these operations were performed against a large, white
enameled tray.
RESULTS AND DISCUSSION
Remarkably severe weather prevailed throughout the collecting period. A
destructive freeze in December 1972 was followed by an unusually dry spring
and a prolonged summer drought. Ten consecutive months of dryness were
finally ended by generous rains in September and October and record-break-
ing rainfall in November. Above-normal precipitation and temperatures
characterized December, but the heavy rains of late January 1974 fell on
ground that had been solidly frozen earlier in the month, producing serious
floods over much of the Willamette Valley. On the whole, it appears that
the normal weather patterns for western Oregon were exaggerated during
the period of this study.
In 1973, the gray- tailed vole experienced a population peak throughout
the Willamette Valley. This was followed in the spring of 1974 by a cata-
strophic decline after which few voles were seen until the end of the year
(R. F. Hoyer, personal communication). Between February 1 973 and January
1974, 428 gray-tailed voles— 377 of them statistically comparable— and 25 1
surface nests were removed from the North, South, and West sites. This
material yielded 22,641 specimens of eight flea species, including 6,761
imagines distributed as shown in Table 1. For most species, identification
of larvae was not possible. Only Atyphloceras multidentatus, Catallagia
charlottensis, Hystrichopsylla occidentalis, and Peromyscopsylla selenis are
regular parasites of voles (Hopkins and Rothschild, 1962, 1971). Corrodop-
sylla curvata is a true shrew flea, Monopsyllus wagneri is a widespread west-
ern deer mouse flea, and Nosopsyllus fasciatus is the introduced northern
rat flea. Owing to the rarity of Rhadinopsylla in collections and the resultant
taxonomic confusion within this large genus, the identity and preferred hosts
of specimens from western Oregon remain unknown.
Table 2 summarizes seasonal changes in the total flea population on all
352
NEW YORK ENTOMOLOGICAL SOCIETY
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VOLUME 91, NUMBER 4
353
three sites. Regardless of species or host preference, fleas were most abundant
during the mild, moist months of spring and early winter and least numerous
during the hot, dry summer. A midwinter decline also was evident. Thus,
in the example of Peromyscopsylla selenis, which despite its generic name
is typically a flea of microtine rodents (Jameson and Brennan, 1957), 85
(71%) of the 1 19 specimens in Table 1 were taken during March-April and
October-December. On the other hand, Corrodopsylla curvata is specific to
insectivores (Hubbard, 1947; Hopkins, 1957), but of 43 accidentals obtained
from nests and pelts of the gray-tailed vole, 27 (63%) were taken during the
peak spring and early winter months. Hystrichopsylla occidentalis is an im-
partial parasite of small mammals within its range (Hopla, 1964), but again
of 237 specimens collected, 170 (72%) were from the same peak months.
These observations are consistent with Holland’s (1 949) thesis that humidity
and temperature are the principal factors influencing flea populations. Sim-
ilar results were obtained by Howell (1955, 1957) in his investigations of
fleas associated with nests of the desert wood rat (Neotoma lepida lepida
Thomas) and by Parker (1958) in a study of fleas— particularly the cerato-
phyllid Thrassis bacchi gladiolis (Jordan)— on the antelope ground squirrel
(Citellus leucurus leucurus (Merriam)). As expected, peaks in larval popu-
lations preceded those of adults; the sole departure from this pattern (North
Site, April) probably was due to sampling error.
The mean number of fleas per vole or per nest is equivalent to a crude
flea index, similar to one proposed early in this century by the British Plague
Commission to monitor flea populations on synanthropic rats in India
(Hirst, 1926, 1927; Muirhead-Thomson, 1968). The sensitivity of this index
can be improved by treating each species separately and by restricting cov-
erage to infested voles and nests (Cole and Koepke, 1947). Both of these
modifications will be demonstrated in forthcoming papers, where the pop-
ulation dynamics of each flea species listed above will be discussed in detail.
ACKNOWLEDGMENTS
Foremost among the recipients of my gratitude is Dr. Gerald W. Krantz, Department of
Entomology, Oregon State University, Corvallis, who enthusiastically supported a research
program largely unrelated to his own profession of acarology. Sincere thanks are also due Mr.
Richard F. Hoyer, long-time Corvallis naturalist, who collected most of the voles and nests
used in this study and offered many important observations drawn from years of experience
in field biology. Dr. Vernon J. Tipton and his colleagues at the Center for Health and Envi-
ronmental Studies, Brigham Young University, Provo, Utah, kindly determined a synoptic
series of the flea species discussed herein. The bulk of the flea collection on which this series
is based has been donated to B.Y.U.
LITERATURE CITED
Cole, L. C. and J. A. Koepke. 1947. Problems of interpretation of the data of rodent-ecto-
parasite surveys. Public Health Reports, Supplement No. 202, pp. 1-24.
354
NEW YORK ENTOMOLOGICAL SOCIETY
Cook, E. F. 1954. A modification of Hopkins’ technique for collecting ectoparasites from
mammalian skins. Entomological News 65:35-37.
Faulkenberry, G. D. and R. G. Robbins. 1 980. Statistical measures of interspecific association
between the fleas of the gray-tailed vole, Microtus canicaudus Miller. Entomological
News 91:93-101.
Henry, L. G. and S. McKeever. 1971. A modification of the washing technique for quantitative
evaluation of the ectoparasite load of small mammals. Journal of Medical Entomology
8:504-505.
Hirst, L. F. 1 926. Researches on the parasitology of plague. Part 1 . Ceylon Journal of Science,
Section D, 1:155-271.
Hirst, L. F. 1927. Researches on the parasitology of plague. Part 2. Ceylon Journal of Science,
Section D, 1:277-455.
Holland, G. P. 1949. The Siphonaptera of Canada. Canada Department of Agriculture Tech-
nical Bulletin No. 70, Ottawa, 306 pp.
Hopkins, G. H. E. 1949. Host-associations of the lice of mammals. Proceedings of the Zoo-
logical Society of London 1 19:387-604.
Hopkins, G. H. E. 1957. Host-associations of Siphonaptera. First symposium on host spec-
ificity among parasites of vertebrates. Switzerland, Universite de Neuchatel, Institut de
Zoologie, pp. 64-87.
Hopkins, G. H. E. and M. Rothschild. 1962, 1971. An illustrated catalogue of the Rothschild
collection of fleas (Siphonaptera) in the British Museum (Natural History). B.M.(N.H.),
London. Vol. Ill, x + 560 pp., Vol. V, viii + 530 pp.
Hopla, C. E. 1964. Alaskan hematophagous insects, their feeding habits and potential as
vectors of pathogenic organisms. Part I: The Siphonaptera of Alaska. Final report between
the Arctic Aeromedical Laboratory, United States Air Force, Fort Wainwright, Alaska,
and the University of Oklahoma Research Institute, Norman, Oklahoma. Contract No.
AF 41(657)-333, x + 346 pp. (Unpublished)
Howell, J. F. 1955. A study of the aspectional variations of Siphonaptera associated with the
nests of the Thomas wood rat Neotoma lepida lepida Thomas. Great Basin Naturalist
15(l-4):35-49.
Howell, J. F. 1957. Fleas associated with nests of the desert wood rat. Journal of Parasitology
43:566-573.
Hubbard, C. A. 1947. Fleas of Western North America. Iowa State College Press, Ames, x +
533 pp.
Jameson, E. W., Jr. and J. M. Brennan. 1957. An environmental analysis of some ectoparasites
of small forest mammals in the Sierra Nevada, California. Ecological Monographs 27:
45-54.
Lipovsky, L. J. 1951. A washing method of ectoparasite recovery with particular reference
to chiggers (Acarina-Trombiculidae). Journal of the Kansas Entomological Society 24:
151-156.
Muirhead-Thomson, R. C. 1 968. Ecology of Insect Vector Populations. Academic Press, New
York and London, viii + 174 pp.
Parker, D. D. 1958. Seasonal occurrence of fleas on antelope ground squirrels in the Great
Salt Lake Desert. Journal of Economic Entomology 51:32-36.
Pearson, J. P. 1972. The influence of behavior and water requirements on the distribution
and habitat selection of the gray-tailed vole {Microtus canicaudus) with notes on Microtus
townsendii. Ph.D. thesis, Oregon State University, Corvallis, 56 numbered pp.
Robbins, R. G. and G. D. Faulkenberry. 1982. A population model for fleas of the gray-tailed
vole, Microtus canicaudus Miller. Entomological News 93:70-74.
Received March 3, 1983; accepted September 30, 1983.
NEW YORK ENTOMOLOGICAL SOCIETY
91(4), 1983, pp. 355-376
HISTOLOGY OF THE MALE REPRODUCTIVE SYSTEMS IN THE
ADULTS AND PUPAE OF TWO DORYLINE ANTS,
DORYLUS (ANOMMA) WILVERTHI EMERY AND
D. {A.) NIGRICANS ILLIGER (HYMENOPTERA: FORMICIDAE)
Francis C. Ford and James Forbes
Department of Biology, Bronx Community College, CUNY,
Bronx, New York 10453, and
Department of Biological Sciences, Fordham University,
Bronx, New York 10458
Abstract.— This paper is the first comprehensive description of the histology of the male
reproductive systems of the adults and pupae of the doryline ants, Dorylus (Anomma) wilverthi,
and D. (A.) nigricans. The reproductive systems consist of the testes and vasa efferentia, the
vasa deferentia, the seminal vesicles, the accessory glands, the bound accessory gland duct, the
ejaculatory duct and wedge, and the aedeagal bladder. The numbers of testicular follicles, the
microanatomy of the organs, the types and amounts of secretions in the lumina of the organs,
as well as the presence or absence of spermatozoa, are compared in the adults and pupae of
these two species. Although similarities exist, significant differences are found in the microanato-
my of the organs, in the types and amounts of secretions produced, and between organs of the
adults and pupae. The histology is also compared with that in the Old World Aenictus gracilis,
the New World Neivamyrmex harrisi, and other studies of Old and New World dorylines.
Important differences exist between the male reproductive systems of the two African Dorylus
species herein studied and the New World N. harrisi. The Dorylus species have a larger number
of testicular follicles, the organs of the system are more complex and produce both acidophilic
and basophilic secretions, the entrance of the ejaculatory duct into the aedeagal bladder, and
the formation of a new dorsal duct at the posterior end of the bladder are different from those
in other dorylines.
This paper describes the histology of the organs of the male reproductive
system in the adults and pupae of Dorylus {Anomma) wilverthi and D. (A.)
nigricans. In an earlier paper, the anatomy of this system in these two ants
has been reported (Ford and Forbes, 1980). The histology described is com-
pared with that in the Old World Aenictus gracilis (Shyamalanath and Forbes,
1983) and in the New World Neivamyrmex harrisi (Forbes and Do-Van-
Quy, 1965) and with the histological comments and illustrations included
in the study of army ant males by Gotwald and Burdette (1981).
Histological studies reveal differences that are not observable by dissection
but that are significant in the microscopic structure of organs, in the types
of secretions produced, and between organs of pupal and adult stages.
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to indicate this fact.
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Fig. 1. Diagram of a lateral dissection of the posterior portion of the gaster of the adult
male ant Dorylus (Anomma) wilverthi. Abbreviations: Ag, accessory gland; Ed, ejaculatory duct;
InV, inner genitalic valve; M, Malpighian tubule; MV, middle genitalic valve; OV, outer genitalic
valve; R, rectum; SV, seminal vesicle; T, testis; V, ventriculus; Vd, vas deferens; VIII-IX,
Roman numerals designate abdominal segments. (From the original of Fig. 1 , F. C. Ford and
J. Forbes, J. New York Entomol. Soc. 88:135.)
MATERIAL AND METHODS
Seven to nine specimens each of the male adults and pupae were furnished
by Albert Raignier, S.J. of Belgium, who collected them in the Republic of
the Congo, Africa, now designated Zaire, in the town of Mayidi. The adults
of nigricans were collected during June of 1956, the pupae of nigricans and
the adults and pupae of wilverthi during November of 1957. The specimens
were preserved in 80% ethyl alcohol. These specimens were sent to the late
T. C. Schnierla of the American Museum of Natural History who gave them
to J. Forbes for anatomical studies.
The pupae of both species were very well developed externally and inter-
nally and were probably close to eclosion. The gastral viscera were removed
entirely and processed by a double infiltration technique (Trombetta and
Forbes, 1977). Sagittal and transverse serial sections were cut at 10 stained
with Harris’ haematoxylin and counterstained with eosin. Although the ma-
terial had been fixed and stored in alcohol for a long period, most of it cut
fairly well. The nigricans adults were in the poorest condition. Their organs
and tissues were brittle when gross dissections were made and were more
resistant to the embedding process, resulting in blocks that shattered during
sectioning. During staining, each group of slides was processed similarly,
but variations in the stained slides were apparent.
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OBSERVATIONS
Throughout this section, the histology of the wilverthi adult system will
be described first. Differences between that system and the pupal system
will follow. Comparisons will be made with the adult and pupal systems of
nigricans.
Testes (Figs. 1, 2, and 3). The testis of wilverthi consists of 35-40 long,
slender, thin-walled follicles. The wall of the follicle is composed of two cell
layers, a basal and an inner one, arranged around a distinct central lumen.
The cells of the basal layer are large, generally cuboidal in shape with in-
distinct lateral margins, and they lie on a distinct basement membrane. The
basally located nuclei of these cells have their long axes parallel to the
basement membrane. The cytoplasm is highly vacuolate throughout but
more granular basally. The inner cell layer consists of flattened nuclei lying
in an irregular layer of cytoplasm above the basal cells. The lumina of the
follicles are filled with clusters of closely packed spermatozoa. The heads of
these sperm clusters lie toward the center, and the tails extend to the pe-
riphery.
The testes of both species are not covered with a capsule. The sections
show a thin covering of many branching tracheae of various sizes. These
tracheae continue into the spaces between the follicles, and they are attached
to the basement membrane of the follicles by very fine, branching, nucleated
fibers. Occasionally, clusters of fat cells are found in the network of tracheae
between the follicles. Embedded in these clusters are a few spheroidal cells,
about the size of adipocytes, that contain irregularly-shaped nuclei and have
the cytoplasm filled with fine, basophilic granules and numerous vacuoles.
These cells resemble urate cells. Acidophilic granular material is found in
spaces between some of the follicles.
In the pupa of wilverthi the testis is similar in its overall histological
arrangement to that of the adult, but the lumina of the follicles are smaller
and contain a network of cytoplasmic strands not connected to the inner
cell layer of the follicle. Clusters of sperm heads are found in the spaces of
the network and are embedded in the cytoplasmic strands, and the tails
extend to the periphery of the follicles.
In nigricans adults no testes were found macroscopically, but the sections
revealed a posterior displaced mass of intertwined large and small tracheae
within which was a large circular mass and a few smaller masses of sper-
matozoa; nothing else was seen. In nigricans pupae each testis consists of
50-55 tubules. The histology of the testicular follicles is similar to the ar-
rangement in the wilverthi pupa.
Vasa efferentia. In wilverthi each testicular follicle ends in a narrow vas
efferens. At the junction of the follicle and the vas efferens there is an abrupt
transition from the larger, basal cells on the wall of the follicle to very low,
columnar cells that form the epithelial lining of the vas efferens. The nuclei
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are centrally located. A basement membrane is not evident, but small muscle
fibers adhere closely to the bases of the epithelial cells and run obliquely
around the vasa efferentia. Clusters of spermatozoa are found only at the
anterior ends of many of these vas efferens. The thin capsule, that extends
between the testicular follicles, continues between the vasa efferentia.
The histology of the vasa efferentia in the wilverthi pupa and in the ni-
gricans pupa is similar to that of the wilverthi adult. No vasa efferentia were
found in the nigricans adult.
Vas deferens. Where the vasa efferentia unite to form the short vas def-
erens, the low columnar cells immediately increase in height to form the
columnar epithelium of the vas deferens. The nuclei of these cells are cen-
trally located. Small clusters of sperm are present in the lumen. The vas
deferens narrows as it proceeds toward the seminal vesicle. Vacuoles are
present in the cytoplasm bordering the lumen. A very faint basement mem-
brane is seen periodically, and the surrounding muscle coat consists of 2 or
3 obliquely arranged fibers. At the junction of the vas deferens with the
seminal vesicle, the columnar cells merge with the tall columnar epithelium
of the seminal vesicle. The muscle coat of the vas deferens continues into
that of the seminal vesicle.
Seminal vesicles (Figs. 1 and 4). The epithelium of the prominent
U-shaped seminal vesicles consists of columnar cells built on a distinct
basement membrane. The cells are arranged in clusters of taller cells inter-
spersed with one or a few shorter, irregularly distributed cells. The nuclei of
the cells are centrally located, and the staining reaction of the cytoplasm is
variable from the basal region to the free surface. The region from the base
to just above the nucleus is packed with granule-filled vacuoles. In the
subnuclear region these granules stain faintly basophilic, and in the region
above the nucleus they stain more strongly. Above this region, there is a
narrow band of strongly acidophilic-staining cytoplasm that extends to the
free surface. Occasionally, in this strongly acidophilic layer, thin columns
of basophilic granules extend toward the surface of the cells. The free surfaces
of most of the cells are dome-shaped, and extensions from the free surfaces
of some cells are pinched-off to form a secretion in the lumen. This secretion
varies in composition and amount along the lumen of these organs. In the
first part, the lumen is filled with a fine-granular, basophilic secretion and
numerous spermatozoa. On the periphery of this secretion are scattered
clusters of a larger, acidophilic granular secretion. Toward the middle region
Figs. 2, 3. Photomicrographs of oblique sections of testicular follicles of Dorylus wilverthi.
x450. 2. Adult. 3. Pupa. B and I, nuclei of basal cell layer and of inner cell layer of testicular
follicles; S, bundles of sperm.
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the secretion is composed of large, acidophilic granules, and some sperm is
present. In this region, also, there is a strongly acidophilic-staining, amor-
phous mass that can vary in amount within the seminal vesicles of the same
ant. In the posterior region, the secretion is granular, stains distinctly ba-
sophilic, and no sperm were seen. The muscle coat is composed chiefly of
obliquely arranged fibers, 4 or 5 fibers in thickness anteriorly and increasing
by 2 or 3 more fibers posteriorly.
In the constricted posterior region of the seminal vesicle, the epithelial
cells are reduced in height. The cytoplasm is basophilic in staining reaction,
and in the upper third of many cells the vacuoles have coalesced to form a
large, clear vacuole. The cytoplasm at the free borders is compact, and there
is a granular, basophilic secretion in the lumen. The muscle coat has again
increased by several fibers in thickness. This constricted end joins the lower,
posterior part of the accessory gland, where it penetrates obliquely through
the muscle coat of the accessory gland.
The organization of the tissues in the seminal vesicles of the pupa of
wilverthi is similar to that of the adult, but no spermatozoa are present. A
granular, basophilic secretion is present in the lumen. The histology of the
seminal vesicles of the nigricans adult and pupa is respectively similar to
that of the wilverthi adult and pupa.
Accessory glands (Figs. 1 and 5). The epithelium of these thick-walled,
S-shaped glands consists of a single layer of cells that varies from cuboidal
to exceptionally tall columnar. No basement membrane is evident. The
epithelium is raised into 2 or 3 oblique folds that extend the length of the
glands. Large, granular, ellipsoidal nuclei are located in the lower half of the
cells. The cytoplasm, variable in appearance from the bases to the free
surfaces, is basophilic in staining throughout the cells. In the basal region it
is highly vacuolate but above the nucleus less so. Within the upper third the
cytoplasm is packed with coarse, spherical, acidophilic-staining granules. In
some regions above this granular, acidophilic layer and at the surface of the
cells there is a thin, hyaline layer which stains lightly basophilic. Some
columnar cells have the cytoplasm above the nucleus filled with acidophilic
granules, and in these cells there is usually a thin separation in the cytoplasm
between the nucleus and the granules. These cells break down to produce
an apocrine secretion that probably contributes to the acidophilic secretion
Figs. 4, 5. Photomicrographs of oblique sections of the seminal vesicle and the accessory
gland in the adult of Dorylus wilverthi. x450. 4. The seminal vesicle, showing the groups of
alternating high and low epithelial cells. 5. The accessory gland, showing the variations in the
staining reactions of the cytoplasm of the epithelial cells. Arrow points to muscle fibers attached
to the bases of epithelial cells. BM, basement membrane; H, hyaline layer; MC, muscle coat;
Se, secretion in lumen; Ss, surface secretion.
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in the center of the lumen of these glands. The surfaces of some cells are
ruptured, and the cell contents produce the hne-granular and the globular,
basophilic secretions found just outside the cells. A well-developed, muscle
coat of 8 to 12 obliquely arranged fibers lies outside the epithelium. Muscle
fibers extend into the epithelial folds, and, in regions around the wall, the
muscle fibers terminate in the basal cytoplasm of the epithelial cells.
Toward the lower ends of the accessory glands, close to where the seminal
vesicles enter, the oblique folds of the epithelium become longitudinal in
direction, and these are located dorsolaterally and ventromedially. Over the
surface of the folds, the cells are exceptionally tall, and, between the folds,
the cells are reduced in height to become cuboidal. In this region, the cy-
toplasm is acidophilic in reaction, less so in the subnuclear region of the
cells. A very fine-granular, acidophilic secretion along the surface of these
cells contributes to a fine-granular, but weakly staining, acidophilic secretion
in the lumen of the ducts.
The median walls of the terminal portions of the accessory glands come
together, the muscle fibers of these median surfaces fuse, and some of the
outer fibers encircle these terminal portions to form a continuous layer. From
here to the end of the system, macroscopically, there appears to be a single
tube. However, histological sections of this single duct in its proximal region
show two lumina; this region is the bound accessory gland duct.
The overall arrangement and appearance of the tissues in the accessory
glands of the nigricans adult and pupa and the wilverthi pupa are similar to
and resemble those of the wilverthi adult. In the pupal stage of these ants,
fewer regions of the epithelium are breaking down, and there is a scattered,
basophilic granular secretion in the lumen. In the nigricans pupa an aci-
dophilic, granular secretion is present, also. In the nigricans adult small
clusters of spermatozoa were seen in the lumen of these glands but only at
the anterior ends.
Bound accessory gland duct. This is a short duct, and in the wilverthi adult
the epithelium, lining the lumina, varies from low to tall columnar cells built
on a basement membrane. The ventral halves of the lumina are lined with
tall columnar cells that are taller on the median and lateral walls. The upper
halves are lined with cells that are lower in height, and there is a middordsal
fold in each lumen. The nuclei are basally located in all the epithelial cells.
The subnuclear cytoplasm is dense and basophilic in staining. In the midre-
gion of the cells, the cytoplasm is condensed into strands, and many large
vacuoles containing basophilic granules lie between these strands. In the
Figs. 6, 7. 6. Photomicrograph of a cross section of the bound accessory gland duct in the
pupa of Dorylus nigricans, x 140. 7. Photomicrograph of a cross section through the anterior
end of the ejaculatory duct in the adult of Dorylus wilverthi. x 140. Se, secretion in lumen.
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ventral halves of the lumina there are large, compact, amorphous, acido-
philic-staining secretions, which are covered dorsally with a basophilic, mu-
cus-like layer. Scattered in the lumina, there is also a small amount of
basophilic granular material. The muscle coat consists of an inner longitu-
dinal and an outer circular or obliquely arranged layer of fibers. The inner
longitudinal fibers are confined to the middorsal, the lateral, and the ventral
walls. Middorsally, this muscle coat is about a dozen fibers in thickness,
whereas on the lateral and ventral walls it is only a few fibers in thickness.
In the median wall the muscle coat is circular and heavier in its midregion.
As the bound accessory gland duct proceeds posteriorly, the median wall
becomes thinner and disappears; first the dorsal half, then the remainder.
The lumina now communicate with each other, and thus begins the ejacu-
latory duct (Fig. 7).
In the nigricans adult the bound accessory gland duct is similar in its
organization to that of the wilverthi adult. In the pupae of wilverthi and
nigricans, the epithelial cells are much taller, and only very small lumina
are present (Fig. 6). A basement membrane is present beneath the epithelium.
The ellipsoidal-shaped nuclei are centrally located in most of these cells.
The subnuclear cytoplasm is compact and acidophilic in staining. The peri-
nuclear cytoplasm in many cells is straplike and separated from that of the
adjoining cells. The cytoplasm toward the surface of the cells is compact
and distinctly acidophilic. The muscle coat is similar to that of the adult.
Ejaculatory duct and wedge (Figs. 1 and 7-14). The epithelium of the
ejaculatory duct continues as simple columnar. The cells on the lateral and
ventrolateral walls are very tall. On the dorsal wall the epithelium on either
side of the midregion is elevated into a lateral fold. The cytoplasm of the
cells on the dorsolateral and lateral walls is vacuolated. The acidophilic
secretion in the lumen does not have the dorsal mucus covering. The un-
derlying muscle coat consists of a number of circular and longitudinally
arranged fibers. Longitudinally arranged fibers are clustered inside the cir-
cular layer under the epithelium of the dorsolateral folds and on the ven-
trolateral walls. The number of longitudinal muscle fibers under the dor-
solateral folds increases, the folds become higher, and their median walls
touch.
A very thin, cuticular layer now covers the lateral and ventrolateral folds
(Fig. 8). This layer, thicker on the median surfaces of the ventrolateral folds,
is the beginning of the lateral arms of the cuticular wedge. The ventrolateral
Figs. 8, 9. Photomicrographs of cross sections of the ejaculatory duct and wedge in Dorylus.
X 140. 8. Section through the anterior arms of the wedge (W) covering the ventrolateral folds
in the pupa of D. nigricans. 9. Section through the lower half of the duct showing the ventral
ridges of the wedge (VR) in the adult of D. wilverthi. Se, secretion in lumen.
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‘Si?
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folds increase in height, and the cuticular covering gets thicker on its un-
derside and projects deeply into the epithelium of these folds (Fig. 9). These
cuticular thickenings on the lateral arms of the wedge are set down in wavy
horizontal layers that are basophilic in staining reaction. A thin surface layer
on the arms of the wedge extends more laterally and is strongly acidophilic.
The clear, cuticular covering of the median surfaces of the ventrolateral folds
joins medially and forms a mid ventral groove (Fig. 9). A little further back
the ventral ridges of the lateral arms of the wedge are directed medially and
the epithelium on the inner sides of these ridges is displaced medially to
form ventral epithelial folds. The midventral groove is elevated, and the
arms of the wedge extend dorsally along the lateral walls of the duct and
reflect medially on themselves. This surface layer is longitudinally ridged.
At this level the epithelial cells on the lateral folds increase in height and
bend downward under the reflected lateral arms of the wedge (Fig. 10). The
epithelium on the dorsal and dorsolateral walls becomes lower. The secretion
is still present in the lumen. The ventral epithelial folds disappear, the ventral
ridges of the lateral arms diminish in height, and the elevated midventral
groove extends upward to become a median ridge of the wedge (Fig. 1 1).
This median ridge is covered with a thick, longitudinally ridged basophilic-
staining layer. The cuticular material within this median ridge is less dense
than the surface layer and does not fill the elevation; its tip is hollow. The
middle portion of the median ridge is reduced to a small ridge. The epithe-
lium beneath the ridge is low columnar and contains basophilic granules.
Under the lateral arms of the wedge the epithelium is flattened. The wedge
at this level is a ridged plate with its lateral arms reflected and tapering
medially (Fig. 12). The remnant of the median ridge elevates slightly again,
but soon disappears. The lateral reflected arms of the ridge are reduced and
disappear. The surface of the wedge becomes smoother, tapers to a U-shaped
trough, which gradually ends on the floor of the ejaculatory duct (Fig. 1 3).
The epithelium on the dorsal and dorsolateral walls decreases in height, and
the distinctly acidophilic cytoplasm along the surface of these cells is con-
densed. The amount of secretion in the lumen increases. The muscle coat
consists of some dorsolateral and ventrolateral longitudinal fibers, which are
surrounded by a heavy band of circular fibers. The ejaculatory duct extends
a short distance beyond the end of the wedge and lies on the roof of the
aedeagal bladder. Here the cells on the dorslateral, lateral, and ventral walls
Figs. 10, 11. Photomicrographs of the lower part of cross sections through the ejaculatory
duct and wedge in the adult of Dorylus wilverthi. x 140. 10. Shows the converging ventral ridges
(VR) of the wedge. 1 1 . Shows the median ridge of the wedge (MR) and the lateral reflected
arms (LRA). Se, secretion in lumen.
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of the ejaculatory duct are reduced to flattened squamous, and the cells on
the ventral wall are covered with a thin cuticular intima. This ventral wall
becomes folded, and the folds deepen when they contact folds on the dorsal
wall of the aedeagal bladder. These deep folds break through, and the ventral
wall of the ejaculatory duct quickly disappears (Fig. 14). The cuticular intima
now covers the remaining epithelium of the ejaculatory duct, and this layer
thickens. The duct decreases in height. Folds on the dorsal wall of the
aedeagal bladder extend upward on either side of the ejaculatory duct and
invade its circular muscle coat. The enclosed lateral walls of the ejaculatory
duct disappear, and the roof of the ejaculatory duct continues for a short
distance as the middorsal wall of the aedeagal bladder. Outside the epithe-
lium, the visceral muscle fibers of the outer muscle coat of the ejaculatory
duct continue for some distance along the dorsolateral walls of the aedeagal
bladder, but the heavy, oblique, body wall muscle fibers of the dorsal wall
of the aedeagal bladder move in from either side.
A cluster of unicellular gland cells is found at the posterior end of the
ejaculatory duct. These cells are situated both outside and within the lon-
gitudinal muscle layer of the roof of the ejaculatory duct. The gland cells
are large, spherical, and ellipsoidal in shape. The nucleus of each gland cell
is vesicular, and the cytoplasm contains a granular, basophilic-staining se-
cretion. These gland cells become smaller in size posteriorly. Minute ductules
emerge from these gland cells and extend tortuously through the thick cuticle
at the end of the ejaculatory duct. A large number of ductules is visible in
some sections; this indicates numerous gland cells (Fig. 15).
In the nigricans adult, the histological organization of the anterior end of
the ejaculatory duct is similar to that of wilverthi. Unfortunately, the tissue
of the remaining portion of this system in the nigricans adult specimens was
too brittle to be sectioned.
At the anterior end of the ejaculatory duct in the pupae of wilverthi and
nigricans, the epithelium is all columnar and arranged into two dorsomedian
folds and two ventrolateral folds. The cells over the dorsolateral folds are
moderate in height, those over the ventrolateral folds are very tall, while
those midventrally are the lowest. The nuclei of these cells are centrally
located, and the cytoplasm along the free borders is condensed and more
acidophilic in staining reaction than that of the remainder of the cell. A
basement membrane is present under the epithelium, but beneath the epi-
Figs. 12, 13. Photomicrographs of cross sections through the ejaculatory duct and wedge
in the adult of Dorylus wilverthi. x 140. 12. The median ridge of the wedge (MR) has decreased
in height. 13. The posterior tip of the wedge (W) is U-shaped. LRA, lateral reflected arms of
the wedge; Se, secretion in lumen.
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Fig. 16. Photomicrograph of a cross section through the dorsal duct in the adult of Dorylus
wilverthi. x 140. This section is a short distance beyond that in the previous figure and is cut
through the anterior ends of the inner genitalic valves. The left side of the photomicrograph
shows the wall of the dorsal duct fusing with the wall of the inner genitalic valve (InV).
thelium of the ventrolateral folds it is a distinct hyaline layer. The lumen is
reduced to thin spaces between the folds of these cells. Throughout the
remainder of the duct and including the wedge, the organization of the tissues
is similar to that of the adult wilverthi, but the cells are taller. There is no
secretion in the lumen of this duct in either pupa.
Aedeagal bladder (Figs. 14-16). The aedeagal bladder has a thick, tightly
folded, cuticular lining, and the nuclei of the flattened epithelial cells lie
between the folds of the cuticle. At the anterior end, the intima on the dorsal
wall is thinner and less folded. In cross section this organ is shallowly
V-shaped with the point of the V directed ventrad. An acidophilic-staining
Figs. 14, 15. 14. Photomicrograph of a cross section through the opening of the ejaculatory
duct into the aedeagal bladder in the adult of Dorylus wilverthi. x 70. DF, dorsal fold of the
aedeagal bladder; S, bundles of sperm; Se, secretion in the bladder. 15. Photomicrograph of a
cross section through the posterior part of the aedeagal bladder in the adult of Dorylus wilverthi
showing the formation of a transverse cuticular fold (Tf) from the dorsolateral wall. This
transverse fold forms a new duct, the dorsal duct, in the upper part of the aedeagal bladder.
x70. BWM, body wall muscles; D, gland ductules from the gland cells, GC; Sd, secretion in
dorsal duct; Se, secretion in aedeagal bladder; VM, visceral muscle.
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secretion, similar to that found in the accessory glands and ejaculatory duct,
and bundles of spermatozoa fill the lumen, but the sperm and the secretion
are not mixed (Fig. 14). In some of the lower portions of the lumen, layers
of a basophilic secretion are interspersed with the acidophilic secretion. The
muscle fibers covering the epithelium of the aedeagal bladder are similar in
diameter to the body wall muscles, and these muscle fibers are generally
oblique in direction. The outermost fibers of the muscle coat lie very close
to the muscles of the external genitalia and insert on the anterior ends of
the genitalic valves. When the valves of the genitalia are removed, the outer
muscle fibers of the aedeagal bladder are disrupted.
Where the ejaculatory duct lies on the roof of the aedeagal bladder, dorsal
folds extend upward on either side of the ejaculatory duct. Shortly thereafter
the ejaculatory duct opens into the aedeagal bladder. The bladder then tapers
posteriorly. At this posterior end of the aedeagal bladder, the cuticle thickens
on the dorsal and dorsolateral walls, and a heavy transverse fold is formed
from the right dorsolateral region (Fig. 1 5). This fold almost completely cuts
off the dorsal region of the aedeagal bladder, but a small channel is still
present on the left side, which connects the dorsal lumen with the remainder
of the aedeagal bladder. More posteriorly a new lateral channel is formed
on the right side of this fold. This results in a block of cuticular material
between the dorsal lumen and the ventral bladder. The cuticular block soon
expands laterally, closes the lateral channels, and completely separates the
dorsal duct from the remainder of the aedeagal bladder. Muscle from the
lateral walls of the aedeagal bladder now covers the ventral wall of the new
dorsal duct. The ventral remaining aedeagal bladder becomes smaller in
diameter and continues for a short distance as a blind pouch, still filled with
a strongly acidophilic-staining secretion and bundles of spermatozoa.
At the posterior end of the aedeagal bladder, numerous unicellular gland
cells, similar to those at the end of the ejaculatory duct, are found on the
dorsal walls. Ductules from these gland cells extend through the thick, dorsal,
cuticular wall of the dorsal duct. These gland cells may contribute to the
secretion that is found in the lumen. The lumen of this dorsal duct is more
or less triangular in shape with the base downward (Fig. 15), but it soon
changes and becomes cruciform (Fig. 16). At this level the gland cells are
on the lateral regions of the duct. This dorsal duct lies immediately under
the spathe of the inner valves. More posteriorly, the heavily sclerotized plates
of the anterior ends of the inner valves are present on either side of the duct,
and the walls of the dorsal duct fuse with the median walls of the inner
valves.
In the pupae of wilverthi and nigricans the histology of the aedeagal bladder
is similar to that of the adult of wilverthi. However, there are neither sper-
matozoa nor secretion in the aedeagal bladder, so that the lumen is not
distended, and the walls are more wrinkled. Also, the entrance of the ejacula-
tory duct into the aedeagal bladder is similar to that of the adult wilverthi.
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DISCUSSION
While the histology of the internal male reproductive organs in the two
African dorylines, Dorylus wilverthi and nigricans, agrees with that of some
of the organs of two other dorylines similarly investigated, the Old World
Aenictus gracilis (Shyamalanath and Forbes, 1983) and the New World
Neivamyrmex harrisi (Forbes and Do-Van-Quy, 1965), significant differ-
ences do occur in other organs.
The thin walls of the testicular follicles in the adults of both D. wilverthi
and A. gracilis are composed of two cell layers, and the arrangement of the
sperm in the lumina is the same. The testes in N. harrisi were apparently
undergoing degenerative changes; the walls of the follicles had only a single
layer of cells, the follicles were filled with granules, and no sperm were
present.
Recently Gotwald and Burdette (1981) investigated the male internal re-
productive system for representative species of army ants. Most of their
specimens were adults, but a few pupae were included in the New and Old
World genera. They found functional testes in the pupae, but in most of the
adults the testes had atrophied so that there was little evidence of their
existence. Prominent testes were found in earlier dissections of adults of
Eciton hamatum (Forbes, 1958). In this study, testes were not found in the
adult of D. nigricans, but the histological sections revealed a few masses of
spermatozoa surrounded by tracheae in the gaster. Sections of the testes in
pupae show that only these organs are packed with sperm. In the pupa of
A. gracilis, the sperm are contained in a large, central vacuole within each
follicular cell. The number of testicular follicles in the pupae examined by
Gotwald and Burdette were not determined. This number is important,
because, from the papers previously cited and one on the male anatomy of
Dorylus wilverthi and nigricans (Ford and Forbes, 1980), New World species
have significantly fewer follicles than do Old World ones. The testes of the
Dorylus species have no visible capsule, whereas a common capsule covers
the testes in A. gracilis and N. harrisi. Each testis has its own capsule in
Eciton hamatum.
The histology of the vasa elferentia and that of the vasa deferentia in the
doryline adults reported is similar, and scattered sperm are present in the
lumina. In the pupae, the epithlium is slightly taller and the lumina are more
constricted in each of these organs, and no sperm are present.
The seminal vesicles have a distinctive histology in both the adult and
the pupal stages that is different from that of the vasa deferentia regardless
of whether sperm is present as it is in the adults or absent as in the pupae.
Our findings, contrary to those of Gotwald and Burdette (1981), show that
no sperm is stored in the seminal vesicles during the pupal stage. The epi-
thelial cells of this organ are larger and taller than those in the vas deferens,
and they are built on a distinct basement membrane. The two Dorylus species
described have the nuclei of the epithelial cells centrally located, the cyto-
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plasm is variable in its staining reaction from base to free surface, and the
secretion in the lumen is both acidophilic and basophilic; this arrangement
seems to be unique. Figure 14 of the Gotwald and Burdette (1981) paper, a
high power photomicrograph of the wall of the seminal vesicle of a species
of the subgenus Anomma, appears similar in many details to Figure 4 of
this paper: The position of the nuclei of the epithelial cells is the same, and
the cytoplasm appears variable in staining reaction but not as granular as it
is in wilverthi and nigricans. In N. harrisi [the organ labelled vas deferens
in Forbes and Do-Van-Quy (1965) has been correctly designed seminal
vesicle by Hung and Vinson (1975)] and in A. gracilis, the nuclei in the
epithelium are basally located and have their long axes parallel to the base-
ment membrane. In the pupa of A. gracilis, the tall columnar cells almost
occlude the lumen of this organ, and the cytoplasm is basophilic.
The accessory glands of the Old World dorylines are shorter than those
of the New World species, and they are not coiled. The epithelium of these
glands in D. wilverthi, nigricans and A. gracilis, which consists of low to tall
columnar cells, is folded the length of the glands and produces both aci-
dophilic and basophilic granular and globular secretions. The staining re-
actions in these cells are more variable in Dorylus than in Aenictus, but the
acidophilic secretion in these glands is produced by an apocrine type of
secretion in the species of both genera. The cells that produce the acidophilic
secretion in the Dorylus species are scattered throughout the epithelium,
whereas in Aenictus these cells are clustered. Within the epithelial folds in
the Dorylus species, muscle fibers join directly to the bases of the epithelial
cells. In the Dorylus specimens of the subgenera Rhogmus and Anomma
examined by Gotwald and Burdette (1981), the lumina of these glands were
empty of secretion, but some spermatozoa were present in limited areas in
the lumen of these glands in Rhogmus. We found in D. nigricans small
bundles of sperm in the lumen of these glands only at the anterior ends. In
N. harrisi, the epithelium of this organ is similar to that of the Old World
dorylines, but it is not folded. The cytoplasm of these cells is filled with fine,
basophilic-staining granules, and the lumen contains a dense, homogeneous,
basophilic secretion that is sometimes coiled and convoluted within the
lumen. In the pupa of A. gracilis, the epithelium is more folded than that
in the adult, these folds almost occlude the lumen, and some neutral-staining
secretion is present in the lumen.
In the accessory gland ducts and in the bound accessory gland duct, epi-
thelial folds are present, but the number and position of these folds differ.
The cells over the folds are usually taller than those between the folds.
Along the lumina of the ejaculatory duct in the various male ants studied,
the epithelial folds undergo changes, and the lateral arms of the wedge are
developed on lateral or ventrolateral folds. The lateral arms of the wedges
in D. wilverthi and A. gracilis arise similarly, but the thickening of the arms
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375
in each is accomplished differently. In both wilverthi and gracilis a prominent
median ridge is formed on the base of the wedge, i.e., where the arms unite.
The formation and appearance of the wedge in N. harrisi is different from
that of the Old World dorylines. The cross-sectional configurations of the
wedges are different in the species of the three genera compared. Throughout
the ejaculatory duct and above the wedge in D. wilverthi there is a consid-
erable amount of secretion, while in A. gracilis and N. harrisi no secretion
is present. In the pupae of D. wilverthi, nigricans, and A. gracilis, the cells
along the ejaculatory ducts are taller, the arrangement of the folds is different
from that in the adults, and no secretion is present in the lumen. The ejacula-
tory duct in N. harrisi and in A. gracilis opens directly through the roof of
the aedeagal bladder at its posterior region, but in the Dorylus species the
end of the ejaculatory duct is accompanied by folds from the roof of the
aedeagal bladder, which envelop it.
Unicellular glands, similar to those found at the posterior end of the
ejaculatory duct and at the posterior end of the aedeagal bladder in the
Dorylus species, were found at the posterior end of the ejaculatory duct in
N. harrisi. No glands were seen in these regions in A. gracilis.
The aedeagal bladder, present in the dorylines, has the same histological
organization in all those studied, and it opens between the inner genitalic
valves. Gotwald and Burdette (1981) did not mention nor figure this organ
in any of the New or Old World army ants they investigated. Of the dorylines
histologically examined so far, it is only in the Dorylus species herein reported
that the posterior part of the aedeagal bladder is divided by a transverse
cuticular fold to form a new dorsal duct, that opens between the inner
genitalic valves. No sperm was present in the aedeagal bladder of N. harrisi
or A. gracilis, but bundles of spermatozoa and an acidophilic secretion filled
this organ in the adult of D. wilverthi, and layers of a basophilic secretion
were interspersed with the acidophilic in the deeper parts. This organ may
be used to store sperm in the adults nearly ready for the nuptial flight.
More doryline male internal reproductive organs will have to be examined
histologically to determine if there are further important generic or subge-
neric differences within this group. This study of the male organs in Dorylus
(Anomma) wilverthi and nigricans and comparisons with the other two dory-
lines similarly reported have shown that the seminal vesicles are distinct
organs in both the pupal and the adult stages. These organs in species of the
subgenus Anomma are more complex than those in Aenictus gracilis and in
the New World Neivamyrmex harrisi in that they produce both acidophilic
and basophilic secretions. The accessory glands of the Old World species
also produce acidophilic and basophilic secretions, while those of N. harrisi
produce only a dense, homogeneous, basophilic secretion. The cuticular
wedges in the ejaculatory ducts arise in similar fashion but are developed
somewhat differently in the three genera. The entrance of the ejaculatory
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NEW YORK ENTOMOLOGICAL SOCIETY
duct into the aedeagal bladder and the formation of a new dorsal duet at
the posterior end of the bladder in D. wilverthi and nigricans are different
from those in other dorylines.
LITERATURE CITED
Forbes, J. 1958. The male reproductive system of the army ant. Eciton hamatum Fabricius.
Proc. Xth Intemat. Congr. Entomol. 1:593-596.
Forbes, J. and D. Do-Van-Quy. 1965. The anatomy and histology of the male reproductive
system of the legionary ant, Neivamyrmex harrisi (Haldeman) (Hymenoptera: Formi-
cidae). J. New York Entomol. Soc. 73:95-1 1 1.
Ford, F. C. and J. Forbes. 1980. Anatomy of the male reproductive systems of the adults and
pupae of two doryline ants, Dorylus (Anomma) wilverthi Emery and D. (A.) nigricans
Illiger. J. New York Entomol. Soc. 88:133-142.
Gotwald, W. H., Jr. and A. W. Burdette. 1981. Morphology of the male internal reproductive
system in army ants: phylogenetic implications (Hymenoptera: Formicidae). Proc. Ento-
mol. Soc. Wash. 83:72-92.
Hung, A. and S. Vinson. 1975. Notes on the male reproductive system in ants (Hymenoptera:
Formicidae). J. New York Entomol. Soc. 83:192-197.
Shyamalanath, S. and J. Forbes. 1983. Anatomy and histology of the male reproductive system
in the adult and pupa of the doryline ant, Aenictus gracilis Emery (Hymenoptera: For-
micidae). J. New York Entomol. Soc. 91:377-393.
Trombetta, L. D. and J. Forbes. 1977. A double infiltration technic for serial sectioning of
heavily sclerotized insects. J. Histotech. 1:25-26.
Received March 7, 1983; accepted September 14, 1983.
NEW YORK ENTOMOLOGICAL SOCIETY
91(4), 1983, pp. 377-393
ANATOMY AND HISTOLOGY OF THE MALE REPRODUCTIVE
SYSTEM IN THE ADULT AND PUPA OF THE DORYLINE ANT,
AEN ICTUS GRACILIS EMERY (HYMENOPTERA: FORMICIDAE)
S. Shyamalanath and James Forbes
Department of Biological Sciences, Fordham University,
Bronx, New York 10458
Abstract.— This paper presents the anatomy and histology of the reproductive system, the
external genitalia, and the terminal gastric sterna in the male adult and advanced-stage pupa
of the Old World doryline ant, Aenictus gracilis. The male reproductive system consists of the
testes, the vasa efferentia, the vasa deferentia, the seminal vesicles, the accessory glands, the
short bound accessory gland duct, the ejaculatory duct and wedge, the aedeagal bladder, and
the external genitalia. The testes are enclosed in a thin common capsule, and each testis is
composed of about 32 follicles. The epithelium of the testicular follicle in the adult consiss of
a basal and an inner layer of cells around a central lumen, which contains scattered or clustered
spermatozoa. In the pupa, the epithelium is composed of irregularly- shaped cells, each with a
large central vacuole filled with spermatozoa. The seminal vesicles are U-shaped in the adult
and pupa. The epithelial cells in the pupa are taller than those in the adult. In the adult,
spermatozoa were present throughout, but in the pupa, sperm were absent throughout. The
accessory glands are tubular and bent around the proximal region of the intestine. The epithelium
is more folded in the pupa than in the adult. In the adult, some regions contain an acidophilic
secretion, some basophilic, and some both. In the ejaculatory duct, a cuticular wedge is present
on the lateral and ventral walls; this wedge is more complex in its structure in the adult. The
duct enters the dorsal, posterior surface of the aedeagal bladder. The epithelium of the bladder
consists of small cells covered by a thick, wrinkled intima. The muscle fibers that surround the
bladder are larger in diameter than the visceral muscles of the ejaculatory duct. The lumen of
the bladder is devoid of secretion. The basal ring of the genitalia, the three pairs of valves, and
the eighth and ninth sterna are described. They are quite different in shape from those of the
New World dorylines, Eciton hamatum and Neivamyrmex harrisi, but in some respects resem-
ble those of the Old World dorylines, Dorylus wilverthi and nigricans. Comparisons made with
previously described Old and New World species have revealed structures and features that
lend support to the concept of the triphyletic origin of the dorylines.
A review of the studies on the male reproductive system in ants was made
by Forbes (1954), and further information has been added by Hung and
Vinson (1975). The study of the male reproductive system in doryline ants
was pioneered by Mukerjee ( 1 926), who described the anatomy of this system
in the Old World Dorylus labiatus. Descriptions of the anatomy and histology
of this system that followed are those of Eciton hamatum (Forbes, 1958),
Neivamyrmex harrisi (Forbes and Do-Van-Quy, 1965), Dorylus wilverthi
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to indicate this fact.
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NEW YORK ENTOMOLOGICAL SOCIETY
and D. nigricans (Ford and Forbes, 1980, 1983). The morphology of the
male internal reproductive system for representative species of Old and New
World army ants has been described by Gotwald and Burdette (1981) to
interpret phylogenetic implications of this morphology. After Clausen’s work
(1938), the genitalic valves have been studied in detail and used as the basis
for the classification of ants even to the subspecies level (Forbes, 1952;
Borgmeier, 1955; Krafchick, 1959).
This is the first anatomical and histological description of the reproductive
system of the male adult and advanced-stage pupa of the Old World doryline
Aenictus gracilis. Comparisons are made with an African Aenictus sp. (Got-
wald and Burdette, 1981) and with other dorylines previously described.
The specimens and methods used in this study were those reported in the
description of the male digestive system of this ant (Shyamalanath and
Forbes, 1980).
OBSERVATIONS AND DISCUSSION
Anatomy of the reproductive system. This system in the male adult and
pupa consists of the testes, the vasa efferentia, the vasa deferentia, the seminal
vesicles, the accessory glands, the short bound accessory gland duct, the
ejaculatory duct and wedge, the aedeagal bladder, and the external genitalia
(Fig. 1).
The testes are enclosed within a very thin, common testicular capsule and
lie in a concavity on the dorsal, posterior half of the ventriculus from the
middle of the 4th to the end of the 5th abdominal segments. Each testis is
composed of about 32 slender follicles, which are longer in the pupa than
in the adult. The tubules of each testis form a compact mass, and their
anterior ends converge medially. The posterior end of each follicle leads into
a short narrow duct, the vas efferens. The vasa efferentia of each testis unite
to form the short, wider vas deferens. The distal end of the vas deferens is
slightly constricted where it leads into a dilated, elongated, U-shaped tube,
the seminal vesicle, situated along the outer margin of the testis. The first
part of the seminal vesicle is convoluted only in the adult and lies beneath
the posterior region of the testis. The proximal arm of the seminal vesicle
lies beneath the distal one, and both arms are covered with the testicular
capsule. The distal arm of the seminal vesicle continues backward, emerges
from the capsule, and opens into the accessory gland at about the middle of
its lateral margin. The seminal vesicle in the pupa is narrower than that in
the adult. The tubular-shaped accessory glands are bent around the proximal
region of the intestine and unite beneath it to form a short bound accessory
gland duct that continues into the ejaculatory duct with the wedge. The
ejaculatory duct opens into the dorsal surface of a thick-walled aedeagal
bladder situated beneath the bound accessory gland duct and the ejaculatory
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379
Fig. 1. Diagram of a lateral dissection of the adult male of A. gracilis. (From the original
of Fig. 1, S. Shyamalanath and J. Forbes, J. New York Entomol. Soc. 88:18.)
duct. The posterior region of the aedeagal bladder lies between the median
walls of the inner genitalic valves and opens to the outside near the ends of
these valves. The aedaegal bladder in the pupa is larger and has a more
spacious lumen than that in the adult.
The anatomy of the reproduction system of A. gracilis bears a close re-
semblance to that of an African Aenictus sp. (Gotwald and Burdette, 1981),
except that the testes were absent in the adult of the African species; these
authors also reported that testes were absent in the adults of some species
of the subfamilies Ecitoninae and Dorylinae. The reduction in the length of
the testicular follicles in the adult of A. gracilis from the length of the follicles
in the pupa could be indicative of the shrinking and eventual total atrophy.
A. gracilis with about 32 testicular follicles resembles the Old World dory-
lines that have comparatively more follicles, 35-40 in the adult and pupa
of Dorylus wilverthi, 50-55 in the pupa of D. nigricans (Ford and Forbes,
1980), and a fair number in D. labiatus (Mukerjee, 1926). The New World
Eciton hamatum has 20 follicles (Forbes, 1958) and Neivamyrmex harrisi
22-25 (Forbes and Do-Vah-Quy, 1965). The common capsule surrounding
the testes of A. gracilis is similar to that in N. harrisi, but no capsule covers
the testes of D. wilverthi, nigricans, or labiatus. In general arrangement the
seminal vesicles resemble those of the other dorylines. However, structures
eomparable to the collecting sac at the anterior end of the seminal duct and
the dilated vesicula seminalis at the posterior end, reported to be present in
D. labiatus, are not present in A. gracilis. The accessory glands of A. gracilis
and the African Aenictus sp. are short, curved glands, which resemble those
of D. labiatus in shape. These glands in D. wilverthi and nigricans are thick
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walled, S-shaped tubes and in E. hamatum and N. harrisi are tightly coiled
tubes situated on either side of the intestine. The bound accessory gland
duct is short and straight in the Old World dorylines, while in the New
World dorylines it is much longer and in E. hamatum it encircles the ven-
triculus 5 or 6 times. The ejaculatory duct in all the dorylines examined
opens into the dorsal, posterior end of the aedaegal bladder. The blind
diverticulum found on the dorsal side of this duct by Mukerjee (1926) in a
male of D. labiatus has not been seen in any other doryline examined or
reported. The aedeagal bladder found in A. gracilis is present in D. wilverthi
and nigricans, in E. hamatum and N. harrisi. It was not reported in D.
labiatus nor in any of the army ants investigated by Gotwald and Burdette
(1981).
EXTERNAL GENITALIA AND TERMINAL GASTRIC STERNA
The external genitalia are composed of a basal ring and 3 pairs of valves,
the outer, the middle, and the inner, all of which are sclerotized. They are
retracted into the genital chamber and only the distal ends of the outer and
inner valves protrude beyond the posterior margin of the gaster (Fig. 1). The
basal ring or lamina annularis is ring-shaped and situated in the posterior
half of the 6th abdominal segment. Its anterior margin is attached to the
reflected intersegmental membranes of the 9th segment. The dorsal surface
of the basal ring is the broadest, its anterior margin is heavily sclerotized,
and there is a distinct median process. Behind this median process there is
a thin, nonsclerotized, V-shaped region that appears as a notch (Fig. 2). The
outer valves or parameres are situated dorsally and laterally behind the basal
ring to which they are attached by a thin membrane. Each has a broad
anterior region, the lamina parameralis, and a narrow, finger-like posterior
region, the paramere. The outer surface of both these regions is convex, and
there is no suture or demarcation between the two regions; both regions are
uniformly sclerotized. Ventrolaterally at the base of the paramere, there is
a distinct indentation (Fig. 3). The laminae paramerales lie close to each
other along the middorsal line, and ventrally their anterior halves are held
together by a thick, nonsclerotized membrane. The parameres are separated
Figs. 2-7. Genitalic valves and terminal gastric sterna of the adult male of A. gracilis. All
figures are drawn to the same scale. 2. Diagram of an anterodorsal view of the basal ring, the
lamina annularis. The dorsal side is downward in this drawing. 3. Diagram of a lateral view
of the right outer genitalic valve, the paramere. 4. Diagram of a lateral view of the right middle
genitalic valve, the volsellaris, attached to an inner part of the lamina parameralis of the outer
valve. 5. Diagram of a lateral view of the right inner genitalic valve, the lamina aedeagalis. 6.
Diagram of a ventral view of the IXth abdominal sternum, the subgenital plate. Dotted lines
indicate the margins of its dorsal plate. 7. Diagram of a ventral view of the Vlllth sternum.
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from each other. Many sensory hairs are present along the upper and lower
margins of the parameres, and some are present on their median surfaces.
The middle valves or volsellares are the smallest of the valves and the most
heavily sclerotized. Each is roughly quadrilateral in shape and is strongly
attached along its anteroventral margin to the ventroposterior margin of the
lamina parameralis (Fig. 4). It lies mediad of the paramere and is inflected
upward. Campaniform sensilla are present on the posterior surfaces, and
sensory hairs are only on the ventroposterior margin. The inner valves or
laminae aedeagales constitute the male intromittent organ. These are mod-
erately sclerotized, narrow, elongated valves lying close to each other. The
posterior half in lateral view is bent downward. When viewed dorsally and
ventrally, they are broader anteriorly and posteriorly and narrower in the
midregion (Fig. 5). The dorsal and ventral inner margins of these valves are
joined by thin membranes enclosing a narrow space that opens to the outside
by a terminal orifice. The dorsal membrane is the spatha. Wavy flaps from
the lateral margins of the posterior region fold ventrally and form a short
ventral trough. The anterior ventral margins of the two valves are attached
to the posterior half of the lateral sides of a triangular plate. The base of this
plate is attached to the posterior margin of the membrane extending between
the ventral anterior halves of the laminae paramerales. This plate is less
sclerotized than the inner valves. Proximally, the aedeagus has a pair of
anterodorsally directed arms, the aedeagal apodemes. These valves are de-
void of sensilla or sensory hairs.
The IXth sternum or subgenital plate is located on the floor of the genital
chamber. It has a broad, shield-shaped body that terminates in two widely
separated processes, and this segment is heavily sclerotized (Fig. 6). The
body consists of a dorsal and ventral plate fused along their lateral margins.
The dorsal plate extends up to half the length of the ventral plate. The forward
margin of the ventral plate has three anteriorly directed apodemes; the mid-
dle one is the shortest. Sensory hairs are found on the posterolateral regions
of the body of this plate and on the terminal processes. The Vlllth sternum
forms the anterior floor of the genital chamber, and it is roughly crescent-
shaped (Fig. 7). The anterior margin has a pair of lateral arms, and the
segment is heavily sclerotized between them. The posterior margin has a
shallow notch in its midregion. Sensory hairs are present on its ventral
posterior half. The external genitalia and terminal gastric sterna of the pupa
are similar in all respects to those of the adult except that the component
parts are less sclerotized.
The external genitalia of A. gracilis conform to the doryline pattern in
general organization and resemble those of the Old World dorylines, D.
wilverthi and nigricans (Ford and Forbes, 1980). The shapes of the com-
ponent parts are strikingly different from those of the New World dorylines,
E. hamaturn (Forbes, 1958) and N. harrisi (Forbes and Do-Van-Quy, 1965).
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While the basal ring of A. gracilis resembles that of the African dorylines,
D. wilverthi and D. nigricans, it has a sclerotized median process on the
anterior margin of the dorsal surface and a posteriorly directed notch behind
this process. The middorsal apodeme present on the ventral part of the basal
ring of D. wilverthi and D. nigricans is absent in A. gracilis. Also, the outer
valves of A. gracilis are not fused to the basal ring along their dorsal, anterior
borders as they are in D. wilverthi and nigricans. The middle valves of A.
gracilis are broad and roughly quadrilateral in shape while those of D. wil-
verthi and nigricans are finger-shaped. These valves of A. gracilis are strongly
attached to the inner surfaces of the outer valves; this is not the arrangement
in D. wilverthi and D. nigricans. The presence of many campaniform sensilla
on the posterior surfaces of the middle valves of A. gracilis has not been
reported in other dorylines. The shapes of the inner valves of all the dorylines
described, the Old World as well as the New, are distinctly different. The
presence of the triangular plate at the anteroventral margins of the inner
valves is characteristic of A. gracilis. The IXth sternum is considered an
integral part of the genitalia because it provides muscle attachment (Kraf-
chick, 1959). In general shape, the IXth sternum of A. gracilis resembles
that of other dorylines (Borgmeier, 1955; Forbes, 1958; Forbes and Do-Van-
Quy, 1965; Ford and Forbes, 1980).
HISTOLOGY OF THE REPRODUCTIVE SYSTEM
The testicular capsule in the adult is a network of interlacing tracheae of
varying diameters, and it continues inward between the testicular follicles.
Fat cells are found on the surface of the capsule. In the pupa, the capsule is
not as well formed; the tracheae are smaller and fewer. The epithelium of
the testicular follicles in the adult consists of a basal layer and an inner layer
of cells arranged around a central lumen (Fig. 8). The larger cuboidal or
pyramidal cells of the basal layer have distinct cell boundaries. The cyto-
plasm of these cells is granular and highly vacuolated, and a large, ellipsoidal
nucleus is located in the basal region. The smaller cells of the inner layer
are without distinct cell boundaries. They have densely granular cytoplasm
and small oval nuclei. The central lumen has clusters of spermatozoa or
scattered spermatozoa and their tails extend into the cytoplasm of the basal
cell layer. This arrangement of the testicular epithlium resembles that in the
adult and pupa of D. wilverthi and in the pupa of D. nigricans (Ford and
Forbes, 1983). In the pupa, large irregularly- shaped cells form the epithelium
of the follicles and there is no central lumen. The cytoplasm is restricted to
the periphery of the cells and encloses a large vacuole that is filled with
spermatozoa (Fig. 9). This condition has not been reported in any of the
other dorylines.
The epithelium of each vas efferens in the adult is a continuation of the
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385
basal epithelial layer of the testicular follicle. These basal cells decrease
abruptly in size and become low and cuboidal in shape (Fig. 8). The nuclei
are centrally located. The lumina of the vasa elferentia contain scattered or
clustered spermatozoa. In the pupa, the cells of these organs are low and
columnar with oval, basally located nuclei and basophilic cytoplasm. The
lumina of the vasa efferentia are narrower than those of the adult and are
devoid of spermatozoa, but they do contain a fine-granular, neutrophilic
secretion that arises from the free surface of the cells.
The epithelial cells of the vas deferens of the adult are larger than those
of the vasa efferentia, and the epithelium of the vas deferens is surrounded
by a thin layer of small, obliquely arranged muscle fibers. The lumen contains
scattered spermatozoa (Fig. 8). In the pupa the single layer of epithelial cells
of this organ is taller than that lining the vasa efferentia, and the cytoplasm
is still basophilic in its staining reaction. The lumen is narrower, is devoid
of spermatozoa and has a secretion similar to that found in the vasa efferentia.
In the seminal vesicles, the epithelium is gradually transformed from the
cuboidal cells of the vas deferens to low columnar. The cytoplasm is granular,
and some vacuoles are present. Each cell has a basally located, ellipsoidal
nucleus; the long axis of the nucleus is parallel to the basement membrane.
A small amount of neutrophilic-staining, granular secretion was seen on the
free surface of some of the cells (Fig. 8). The muscle coat gradually becomes
thicker from the proximal to the distal region of this organ. In the narrow
duct at the posterior end of the seminal vesicle, the epithelium increases
slightly in height and the nuclei change their position to lie at right angles
to the basement membrane. This position of the nuclei continues into the
accessory glands. The lumen of this organ is filled with spermatozoa except
in one adult where the sperm were confined to the testicular follicles. Sper-
matozoa were never found in the distal narrow duct (Fig. 10). In the pupa
the epithelium consists of a single layer of tall columnar cells that almost
occlude the lumen. The cytoplasm is more granular, basophilic and with
fewer vacuoles. The lumen is filled with coarser granules that are less ba-
sophilic than those in the cytoplasm and no spermatozoa are present (Fig.
9).
The histology of the seminal vesicle and the distribution of spermatozoa
are similar in the adults of the dorylines described. Variations in the secre-
tions of this gland have been reported: In N. harrisi (Forbes and Do-Van-
Quy, 1965) the secretion is granular and acidophilic in the constricted ter-
Figs. 8, 9. Photomicrographs of sections through the testes of A. gracilis. 8. Adult testis that
shows testicular follicles, vasa efferentia, vas deferens, and seminal vesicle. x275. 9. Pupal
testis that shows follicles and seminal vesicle, x 600.
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NEW YORK ENTOMOLOGICAL SOCIETY
minal region; in D. wilverthi and nigricans there is mixing of both acidophilic
and basophilic secretions in the anterior region and basophilic in the pos-
terior.
The epithelium of the accessory glands in the adult is folded and the cells
vary in height from low cuboidal to tall columnar. The cytoplasm is granular
and slightly vacuolate. The secretion in the dorsal half of the gland is dis-
tinctly acidophilic and is composed of coarse spherical granules that are
formed in the supranuclear region of the cells (Fig. 10). The cells become
packed with this secretion, the surface membrane of the cells disintegrates,
and the secretion is discharged into the lumen. Only a thin basal layer of
cytoplasm containing the nuclei remains of these epithelial cells that elab-
orated this secretion. In the ventral half of the gland, the secretion is aci-
dophilic toward the median wall, basophilic toward the lateral wall, and a
combination of both in the center of the lumen. The basophilic secretion,
in contrast to the acidophilic, is developed within the cytoplasm of the entire
cell and consists of small, fine granules concentrated in the upper ends of
the cells and is discharged through the surface membrane into the lumen.
The muscle coat is composed of an inner layer of 3 or 4 oblique fibers, 1 or
2 of which extend into the epithelial folds, and an outer layer of 2 or 3
circular fibers (Fig. 10). In the pupa the epithelium consists of a single layer
of columnar cells that vary in height and is more folded than in the adult.
These folds almost occlude the lumen. The cytoplasm is fine-granular
throughout the cells but more dense in the supranuclear regions. A small
amount of neutral staining secretion is present in the lumen.
Spermatozoa are normally absent in these glands, but their unusual pres-
ence has been reported at the anterior ends of these glands in the adults of
D. nigricans (Ford and Forbes, 1983) and in limited areas in these glands
in the subgenus Rhogmus (Gotwald and Burdette, 1981).
The accessory gland ducts are wider proximally, and their epithelial cells
vary in height from cuboidal to columnar. In the adult, where each accessory
gland enters its duct, there are two epithelial folds on the lateral and two on
the medial walls. These folds alternate in position and might serve as a
closing mechanism. The pupa has only two median folds, one dorsal and
one ventral. In the adult the lumen contains a neutral, granular secretion
which is absent in the pupa.
The bound accessory gland duct is formed by the two accessory gland
ducts held together by a common peripheral muscle coat. Anteriorly, the
Figs. 10, 11. Photomicrographs of cross sections of organs in the adult of A. gracilis. 10.
The accessory gland and the seminal vesicle opening into it. x266. 1 1. The bound accessory
gland duct, x 300.
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NEW YORK ENTOMOLOGICAL SOCIETY
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389
epithelium in each duct has two folds, one dorsomedian and one ventro-
median. The cells over these folds are columnar and the rest of the epithelium
is cuboidal. Posteriorly, these folds gradually decrease in height and new
folds are formed on the dorsal, lateral, and ventral walls. The lumina contain
a neutrophilic, granular secretion. The muscle coat is composed of a few
circularly arranged fibers around each accessory gland duct, and a few pe-
ripheral circular and oblique fibers that bind the two ducts together (Fig.
1 1). At the posterior end, the muscle fibers between the two ducts are reduced
in number and disappear. The two lamina join into one and the duct formed
is the ejaculatory duct. Histologically, the pupa is similar to the adult in this
region.
The first part of the ejaculatory duct has two prominent dorsal folds and
smaller lateral and ventral folds. The latter become more pronounced pos-
teriorly. The columnar epithelium in this duct is taller over the folds. In the
pupa, this region has narrow folds; three dorsal, two lateral, and one ventral.
In the adult, the apices of the two dorsal folds become much broader, and,
at tliis level, the formation of the cuticular wedge is first seen as a thin intima
over the epithelium on the ventrolateral folds. The muscle coat consists of
2 or 3 inner oblique fibers and 1 or 2 outer circular fibers. More posteriorly,
a fold is formed between the two dorsal folds, the small lateral folds broaden,
and the ventral one becomes a tall narrow midventral fold. The cuticular
wedge is now broadly W-shaped, thicker ventrally, and the lateral arms are
thinner and wavy in appearance. In the pupa the wedge is short and is less
complex in its structure. The muscle coat on the dorsal and lateral walls in
the adult has increased by several fibers in thickness, and some of these
fibers are longitudinally arranged (Fig. 12). In the pupa, the muscle fibers
are not distinct and are being formed. Further along in the adult, the mid-
ventral portion of the wedge becomes thicker, broader for a short distance,
and then narrows and extends upward into the lumen. The middorsal fold
is reduced in height and disappears, and the midventral fold of the wedge
lowers. The muscle coat becomes thicker laterally, and pushes the lateral
folds inward. Toward the distal region, the lateral arms move mesially and
the wedge is W-shaped (Fig. 13). The posterior tip of the wedge is thinner
and flatter. The distal end of the ejaculatory duct extends through the dorsal
wall of the aedeagal bladder at the level of the anterior region of the inner
Figs. 12-14. Photomicrographs of cross sections through the ejaculatory duct and wedge in
the adult of A. gracilis; all are magnified x215. 12. This section is about at the midregion of
the wedge and shows the extended lateral arms of the wedge. 1 3. This section shows the reduction
of the lateral arms of the wedge, the formation of its base, and the laterally compressed lumen.
14. This section shows the end of the ejaculatory duct opening through the roof of the aedeagal
bladder.
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NEW YORK ENTOMOLOGICAL SOCIETY
15 VC
Fig. 15. Photomicrograph of cross sections through the ejaculatory duct and the aedeagal
bladder in the pupa of A. gracilis. x210.
valves (Fig. 14), and just beyond the end of the wedge the floor of the
ejaculatory duct opens into the aedeagal bladder. The muscles of the dorsal
and lateral walls of the ejaculatory duct are incorporated into the dorsal wall
of the aedeagal bladder. The lumen of the duct throughout, in both adults
and pupae, is devoid of any secretion.
The cuticular wedge on the ventral surface of the ejaculatory duct, referred
to as the chitinous ‘‘penes” in D. labiatus by Mukerjee (1926) and described
in detail by Clausen (1938) in formicine ants, is present in all male ants in
which the histology has been described. The shapes of the wedges and the
histological organization of the ejaculatory ducts exhibit differences in all
male ants. In the origin of the arms of the wedge from the ventrolateral folds
of the duct and in the formation of a ventral median ridge on the base, A.
gracilis resembles the Old World dorylines.
The wall of the aedeagal bladder consits of a thick, wrinkled intima and
an underlying epithelium of small cells with prominent spherical or ellip-
soidal nuclei. The epithelium is surrounded by a thick muscle coat of large
inner circular and outer obliquely arranged fibers; the latter are inserted on
the median walls of the inner valves at their anterior ends. In the adult, the
VOLUME 91, NUMBER 4
' 391
dorsal and lateral walls in the anterior region of the bladder are deeply folded,
and muscle fibers extend into these folds. In the pupa, the walls are not
folded, the lumen is spacious (Fig. 1 5), and the dorsal wall is depressed in
front of the point where the ejaculatory duct opens into the bladder. In the
posterior region, in both the adult and the pupa, the intima is thin and not
wrinkled, and there are small folds on the lateral walls. The lumen of the
bladder opens between the inner valves. There is no secretion anywhere in
the lumen of the aedeagal bladder.
Gotwald and Kupiec (1975), who analyzed the existing information on
the morphology, behavior, and geographical distribution of the doryline
tribes, indicate that the subfamily Dorylinae, as presently constituted, is
triphyletic. The three lineages are the Ecitonini-Cheliomyrmecini, the Dor-
ylini, and the Aenictini. They advocate the retention of the subfamily Dory-
linae to include the tribe Dorylini and make a case for the creation of a
subfamily Ecitoninae, already introduced by Brown (1973) to include the
tribes Ecitonini and Cheliomyrmecini. The status of Aenictini, they con-
clude, remains to be determined by further investigation.
This report of the anatomy and histology of the reproductive system has
brought to light several structures and features of phylogenetic importance.
The larger number of testicular follicles, the histology of the testicular epi-
thelium, the general structure of the accessory glands, the short straight
bound accessory gland duct and ejaculatory duct, the formation of the chi-
tinous wedge in the ejaculatory duct, and the structure of the genitalia are
features in which A. gracilis is different from the New World dorylines. Some
of these structures have been noted and described by Gotwald and Burdette
(1981). It also differs from the Old World genus Dorylus subgenus Anomma
in the presence of a capsule covering the testes, in the histology and secretion
of the seminal vesicles, in the development of the wedge in the ejaculatory
duct, in the entrance of the ejaculatory duct into the aedeagal bladder, in
the absence of unicellular glands at the end of the ejaculatory duct and the
aedeagal bladder, in the absence of a dorsal duct also at the end of the
aedeagal bladder, and in the shapes of the genitalic valves and terminal
sterna. These features exhibited by the male reproductive system in Aenictus
lend support to the elevation of the tribe Aenictini to a subfamily rank, and
to the triphyletic origin of the dorylines.
ABBREVIATIONS
A, anus; AB, aedeagal bladder; AG, accessory gland; AGD, accessory gland
duct; Ao, aorta; Ap, apodeme; ASe, acidophilic secretion; B, basal cell layer;
Br, brain; BR, basal ring; BSe, basophilic secretion; BT, buccal tube; CM,
circular muscle; CS, campaniform sensilla; DIF, dorsolateral fold; DM, di-
lator muscle; DmF, dorsomedian fold; E, epithelium; ED, ejaculatory duct;
392
NEW YORK ENTOMOLOGICAL SOCIETY
EF, epithelial fold; Gl, G2, G3, G4, gastric ganglia; H, heart; I, inner cell
layer; IC, infrabuccal chamber; In, intestine; It, intima; IV, inner genitalic
valve; L, labium; LA, lateral arm; LP, lamina parameralis; Lu, lumen; M,
mandible; MC, muscle coat; MG, mandibular gland; MP, median process;
MT, Malpighian tubule; N, notch; Oc, ocellus; Oe, oesophagus; OL, optic
lobe; OM, oblique muscle; OV, outer genitalic valve; Pa, paramere; PC,
preoral cavity; Pe, petiole; PeG, petiolar ganglion; PG, postpharyngeal gland;
Ph, pharynx; PM, peripheral muscle coat; R, rectum; RP, rectal pad or gland;
RV, rectal valve; S, spermatozoa; SD, salivary duct; Se, secretion; SG, sal-
ivary gland; SI, salivarium; SoG, suboesophageal ganglion; SV, seminal ves-
icle; T, testis; Tl, T2, T3, thoracic ganglia; TC, testicular capsule; TF, tes-
ticular follicle; TP, triangular plate; V, vacuole; VE, vas efferens; VD, vas
deferens; VIF, ventrolateral fold; VmF, ventromedian fold; Vn, ventriculus;
VNC, ventral nerve cord; W, wedge; IX, 9th abdominal sternum; X, 10th
abdominal tergum.
LITERATURE CITED
Borgmeier, T. 1955. Die Wanderameisen der Neotropischen Region (Hymenoptera: Formici-
dae). Studia Entomologia, Nr. 3, Editora Vozes Limitada, Petropolis, R.J. Brasil 3:9-
716.
Brown, W. L., Jr. 1973. A comparison of the Hylean and Congo-West African rain forest ant
faunas. Pages 161-185 in: B. J. Meggers, E. S. Ayensu and W. D. Duckworth (eds.).
Tropical Forest Ecosystems in Africa and South America: A Comparative Review. Smith-
sonian Institution Press, Washington, D.C.
Clausen, R. 1938. Untersuchungen uber den mannlichen Copulatinsapparat der Ameisen,
speciell der Formicinae. Mitteil. Schweiz. Entomol. Gesell. 17:233-346.
Forbes, J. 1952. The genitalia and terminal segments of the male carpenter ant, Camponotus
pennsylvanicus DeGeer, J. New York Entomol. Soc. 60:157-171.
Forbes, J. 1954. The anatomy and histology of the male reproductive system of Camponotus
pennsylvanicus DeGeer. J. Morph. 95:523-556.
Forbes, J. 1958. The male reproductive system of the army ant, Eciton hamatum Fabricius.
Proc. Xth Intemat. Congr. Entomol. 1:593-596.
Forbes, J. and D. Do-Van-Quy. 1965. The anatomy and histology of the male reproductive
system of the legionary ant, Neivamyrmex harrisi (Haldman) (Hymenoptera: Formici-
dae). J. New York Entomol. Soc. 73:95-1 1 1.
Ford, F. C. and J. Forbes. 1980. Anatomy of the male reproductive systems of the adults and
pupae of two doryline ants, Dorylus (Anomma) wilverthi Emery and D. (A.) nigricans
Illiger. J. New York Entomol. Soc. 88:133-142.
Ford, F. C., and J. Forbes. 1983. Histology of the male reproductive systems in the adults
and pupae of two doryline ants, Dorylus (Anomma) wilverthi Emery and D. (A.) nigricans
Illiger (Hymenoptera: Formicidae). J. New York Entomol. Soc. 91:355-376.
Gotwald, W. H., Jr. and A. W. Burdette. 1981. Morphology of the male internal reproductive
system in army ants: phylogenetic implications (Hymenoptera: Formicidae). Proc. Ento-
mol. Soc. Wash. 83:72-92.
Gotwald, W. H., Jr. and B. M. Kupiec. 1975. Taxonomic implications of doryline worker
ant morphology: Cheliomyrmex morosus (Hymenoptera: Formicidae). Ann. Entomol.
Soc. Amer. 68:962-971.
VOLUME 91, NUMBER 4
393
Hung, A. and S. Vinson. 1975. Notes on the male reproductive system in ants (Hymenoptera:
Formicidae). J. New York Entomol. Soc. 83:192-197.
Krafchick, B. 1959. A comparative study of the male genitalia of North American ants
(Formicidae) with emphasis on generic differences. Dissertation, University of Maryland.
Mukerjee, D. 1926. Digestive and reproductive systems of the male ant, Dorylus labiatus
Schuck. J. Proc. Asiatic Soc. Bengal, n.s. 22:87-92.
Shyamalanath, S. and J. Forbes. 1980. Digestive system and associated organs in the adult
and pupal male doryline ant Aenictus gracilis Emery (Hymenoptera: Formicidae). J. New
York Entomol. Soc. 88:15-28.
Received March 7, 1983; accepted September 14, 1983.
NEW YORK ENTOMOLOGICAL SOCIETY
91(4), 1983, pp. 394-404
CUTICULAR PIGMENT CHANGES IN WORKER
YELLOWJACKETS (HYMENOPTERA: VESPIDAE)
Kenneth G. Ross^
Department of Entomology, Cornell University, Ithaca, New York 14853
Abstract.— \5mxs\xdi\ cuticular marks on the yellow regions of the gastral terga of worker
yellowjackets of two species ( Vespula vulgaris and V. maculifrons) were studied. These resem-
bled marks found on the gastral terga of physogastric Vespula foundresses. The marks were
associated with ovarian development among workers of queenless V. vulgaris colonies, and the
change in structure of pigment granules was similar to that occurring in physogastric queens.
Cuticular marks of V. maculifrons workers from queenright colonies were not related to ovarian
development and the constituent pigment of the marks differed from that of foundresses and
laying workers. The evidence presented supports the hypothesis of Ishay and Shimony that
changes in structure of the pigment granules of ovipositing wasps are due to components of the
granules being shunted to the production of nucleic acids.
Many social wasps of the subfamily Vespinae are aposematically colored
with bright yellow or orange pigment on a contrasting dark background. The
development of characteristic dark blotches on the otherwise lightly pig-
mented regions of the cuticle of physogastric queens is well known among
these wasps (Heldmann, 1934; Spradbery, 1973). These marks typically
appear medially on the posterior portions of gastral terga I-IV in late season
foundresses (Edwards, 1980); their cause has variously been attributed to
external wear, precipitation of pigments due to friction between sclerites, or
oxidation of the pigments due to diffusion of substances through the cuticle
(Marchal, 1896; Becker, 1937a; Spradbery, 1973). Ishay and Shimony (1982)
presented evidence that the cuticular marks result from loss and change in
structure of the pigment granules (xanthosomes) comprising the marks. The
pigment granules are thought to contain pteridines (Becker, 1937b) which
may be shunted to the production of nitrogenous bases for DNA synthesis.
In this paper I report the occurrence of anomalous dark marks on the
yellow regions of the gastral tergites of workers of Vespula vulgaris (L.) and
V. maculifrons (Buysson) similar to the marks found on aged queens. Evi-
dence is presented to suggest that the mechanisms giving rise to the marks
in workers are of two distinct types, with one apparently analogous to that
occurring in aged queens.
' Present address: Department of Entomology, University of Georgia, Athens, Georgia 30602.
The publication costs of this article were defrayed in part by page charge payment. This article
must therefore be hereby marked “ Advertisement'’ in accordance with 18 U.S.C. §1734 solely
to indicate this fact.
VOLUME 91, NUMBER 4
395
Table 1. Colony composition and population of five Vespula study colonies.
Date
Population
Found-
ress
Size of
worker
sample
taken
Colony
collected
5
S
9
Total
present
V. vulgaris
VIO
Aug. 27
1,420
—
—
1,420
no
80
V16
Sept. 16
713
96
—
809
no
79
V35
Nov. 1
15
-
-
15
no
15
V. maculifrons
M26
Oct. 8
1,164
454
165
1,783
yes
80
M29
Oct. 15
1,215
552
154
1,921
yes
60
MATERIALS AND METHODS
Three colonies of Vespula vulgaris and two of V. maculifrons were col-
lected during late summer and fall of 1982 in Tompkins County, New York.
These colonies were part of a larger Vespula colony survey in which 35
colonies of five species were sampled; they were chosen for further study on
the basis of observed cuticular anomalies of workers. The colonies were
lightly anesthetized with ethyl ether, excavated, and transported to the lab-
oratory in large plastic bags. The colonies were killed by freezing. Adults
were counted and sorted according to sex and caste; colony composition and
population are presented in Table 1. Workers and queens were held in a
freezer at — 1 0°C until subsequent examination.
Among the study colonies, the V. vulgaris colonies were queenless at the
time of collection, while the V. maculifrons colonies possessed functional
queens (Table 1). Queenlessness was inferred from non-recovery of a found-
ress, presence of workers with well developed ovaries, and presence of su-
pernumerary eggs in the cells (Ross and Visscher, 1983).
The inner and outer surfaces of gastral terga of workers and queens were
studied using light and scanning electron microscopy. The inner fold of the
cuticle of the sclerites was teased away and the specimens were coated with
a thin (250-350 A) layer of gold-palladium in a Balzers® sputter coater. The
external and internal surfaces of the terga were observed with a AMR® Model
1000 SEM at 10 kilovolts.
Random samples of non-teneral workers from within the colonies were
selected for wing wear analysis and dissection of the ovaries. Wing wear was
determined subjectively on a zero (no wear) to three (greatest wear) scale.
Degree of development of gastral cuticular marks was similarly rated on a
zero (no marks) to two (extensive marks) scale. Ovarian development was
assessed using a modification of Cumber’s (1949) ovariole index.
Data were descriptively analyzed with Exploratory Data Analysis (Velle-
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NEW YORK ENTOMOLOGICAL SOCIETY
Table 2. Frequency of occurrence of anomalous cuticular marks on the gasters of workers
of five Vespula colonies. Percentages of total are in parentheses.
Colony
No marks
Number of workers with
Intermediate
development
of marks
Strong
development
of marks
VIO
72 (90%)
6 (7.5%)
2 (2.5%)
V16
52 (65.8%)
1 (1.3%)
26 (32.9%)
V35
1 1 (73.3%)
-
4 (26.7%)
M26
72 (90%)
—
8 (10%)
M29
43 (71.7%)
7 (11.7%)
10(16.7%)
man and Hoaglin, 1981). When significant trends were suggested, data were
further analyzed using conventional statistical tests.
RESULTS
Workers in the five study colonies possessed dark cuticular marks on the
yellow portion of one or more gastral terga (Fig. 1). The frequency and degree
of development of the marks among colony workers are shown in Table 2.
The marks appeared identical to those of physogastric queens when the
isolated terga were viewed under a dissection microscope from internal and
external aspects. The marks occurred on the posterior portions of the terga
and were most prominent medially ; they consisted of areas of reddish-brown
pigment bordered by dark bands. Immediately adjacent were areas of ap-
parently normal yellow pigment. Clear spots throughout the marks represent
the columnae (cuticular pillars between the infolding cuticle, see Becker,
1937b; Shimony and Ishay, 1981). The layer of reddish-brown pigment
comprising the cuticular mark was dry, thin, and brittle and was easily
removed as flakes, while adjacent layers of normal yellow pigment were
considerably thicker and more malleable. Concentric dark rings throughout
the reddish-brown area, conspicuous in physogastric queens (Fig. la), were
absent from the marks of workers. Cuticular marks were not found in males
or fall queens from the study colonies, nor were they found in workers
randomly sampled from 30 other colonies of V. vulgaris, V. maculifrons, V.
germanica (F.), V. flavopilosa Jacobson, or V. vidua (Saussure) collected
during the same season.
For V. maculifrons workers the marks were most commonly found on
gastral tergum I (Fig. 1 c), but occasionally also on T II. Many of these workers
possessed dark discolorations of the lateral and ventral yellow regions of the
gaster as well. For V. vulgaris workers the marks occurred on gastral terga
I-V and were often spread laterally along the posterior margin of the segment
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397
Figs. la-c. la. Cuticular marks on gastral tergites of physogastric queen of V. maculifrons
(arrows). The yellow regions are faded in contrast to fall or spring queens ( x 6.6, bar = 4.0 mm),
lb. Cuticular marks on gastral tergites of laying worker of V. vulgaris (arrow). The yellow regions
of the body are dull and faded in contrast to nestmates without developed ovaries ( x 8.2, bar =
3.0 mm). Ic. Anomalous cuticular mark on gastral tergite I of V. maculifrons worker (arrow)
(x8.2, bar = 3.0 mm).
(Fig. lb). These workers also commonly exhibited extensive lateral discol-
oration. The yellow pigment of V. vulgaris workers with marks was quite
dull and faded in comparison to other nestmates, as is common for phy-
sogastric foundresses. In contrast, the yellow pigment of V. maculifrons
workers with gastral cuticular marks was characteristically bright.
Many workers in the three queenless V. vulgaris colonies had developed
ovaries (Ross, unpublished data) and were acting as functional reproductives.
The presence and degree of development of cuticular marks were positively
associated with ovarian index for workers in these colonies (ANOVA, F =
40.7,23.2, and 103.6; all F < 0.005). Ovarian development was uncommon
among workers in the two queenright V. maculifrons colonies and was not
associated with development of gastral cuticular marks. Presence of the
marks was not associated with wing wear in workers of the five study colonies.
Scanning electron micrographs revealed loss of setae and hairs from the
gastral terga of laying workers of V. vulgaris, similar to the loss of these
structures in physogastric queens (Figs. 2a, b). Loss of the hair and setae
occurred on the same segments as did cuticular marks. However, the areas
of hair loss did not correspond to the exact areas of the cuticular marks.
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NEW YORK ENTOMOLOGICAL SOCIETY
Figs. 2a-d. 2a. Hair and seta loss on gastral tergum of laying worker of V. vulgaris ( x 69,
bar = 0.5 mm). 2b. Hair and seta loss on portion of gastral tergum of physogastric V. maculifrons
queen. Photograph shows the region in which the cuticular mark is found ( x 74, bar = 0.5 mm).
2c. Gastral tergum I of V. maculifrons worker with cuticular mark. Posterior of segment is to
the top (x78, bar = 0.5 mm). 2d. Gastral tergum of virgin V. maculifrons queen collected in
autumn. Posterior of segment is to the left (x 30, bar = 1.0 mm).
Indeed, hair loss of physogastric queens often involved the entire yellow
region of any given tergum. Workers of V. maculifrons with cuticular marks
and virgin queens collected in autumn exhibited an abundance of hair and
setae on the gastral tergites (Figs. 2c, d).
In virgin V. maculifrons queens, pigment granules were deposited in dense
layers throughout the yellow portion of the tergum. The granules were cy-
lindrical and elongate (ca. 0.8 ixm long. Fig. 3a), as shown by Shimony and
Ishay (1981) and Ishay and Shimony (1982) for various vespine species.
Granules from the yellow pigmented areas surrounding the marks of phy-
sogastric queens were more sparsely distributed; many of the granules were
shorter and more barrel-shaped than typical granules from virgin queens
(ca. 0.6 ixm long. Fig. 3b). All granules from the areas of the cuticular marks
VOLUME 91, NUMBER 4
399
Figs. 3a-c. 3a. Pigment granules from yellow region of gastral tergum of virgin V. maculifrons
queen (x 8,300, bar = 3 /urn). 3b. Pigment granules from yellow region of gastral tergum of
physogastric V. maculifrons queen (x 10,300, bar = 3 ixm). 3c. Pigment granules from area of
gastral cuticular mark of physogastric V. maculifrons queen (x 8,600, bar = 3 )um).
of physogastric queens lost their cylindrical structure to an even greater extent
and became barrel-shaped or spherical (Fig. 3c).
Yellow pigment granules from normal V. maculifrons workers exhibited
a short, cylindrical structure (ca. 0.6 ixm long, Fig. 4a). Granules from the
yellow-pigmented regions adjacent to cuticular marks of V. maculifrons
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NEW YORK ENTOMOLOGICAL SOCIETY
Figs. 4a-c. 4a. Pigment granules from yellow region of gastral tergum of normal V. ma-
culifrons worker (x 10,500, bar = 3 ixm). 4b. Pigment granules from yellow region of tergum of
V. maculifrons worker with gastral cuticular mark (x 10,900, bar = 3 yuni). 4c. Pigment mass
from area of gastral cuticular mark of V. maculifrons worker (x 8,500, bar = 3 ixm).
workers resembled granules from normal conspecifics in size and shape (Fig.
4b). Granules from the area of the cuticular mark lost their discrete, cylin-
drical character and formed amorphous masses of pigment (Fig. 4c).
Pigment granules from the yellow regions of gastral terga of normal V.
vulgaris workers were cylindrical (ca. 0.6 ij.m long, Fig. 5a) and occurred in
dense layers. As in physogastric queens, pigment granules from the yellow
VOLUME 91, NUMBER 4
401
Figs. 5a-c. 5a. Pigment granules from yellow region of gastral tergum of normal V. vulgaris
worker (x 10,300, bar = 3 ixm). 5b. Pigment granules from yellow region of gastral tergum of
laying V. vulgaris worker (x 10,150, bar = 3 ixm). 5c. Pigment granules from area of cuticular
mark on gastral tergum of laying V. vulgaris worker (x 10,600, bar = 3
portions of terga of laying workers appeared less dense in their distribution,
and the granules became barrel-shaped or spherical (Fig. 5b). Granules from
the areas of the cuticular marks lost their discrete structure to some extent
and were without exception spherical or irregularly shaped (Fig. 5 c).
402
NEW YORK ENTOMOLOGICAL SOCIETY
DISCUSSION
How do the results of this study bear on hypotheses of the origin of
cuticular marks on the gastral terga of foundress queens? As suggested by
Ishay and Shimony (1982), development of cuticular marks in queens is
associated with changes in the structure of the pigment granules found in
the yellow regions of the wasps, and this change in the granules is associated
with ovarian development. V. vulgaris workers with marks from the present
study resembled physogastric queens in this altered structure of the yellow
pigment granules, particularly in the area of the dark marks. These workers
also exhibited significant development of the ovaries and loss of gastral hair
and setae (probably resulting from friction between the gaster and cell wall
during oviposition). These data suggest that the mechanisms underlying the
development of cuticular marks in queens and in laying workers are identical.
The development of cuticular marks in laying workers was first reported by
Marchal (1896) for Dolichovespula media (Retzius) and is probably common
to all laying vespine workers.
Spradbery (1973) suggested that cuticular marks in queens result from
diffusion of substances through the normally impermeable cuticle via the
sockets of worn-away hairs and setae. This hypothesis cannot be ruled out
by the results of the present study, although two observations diminish the
likelihood of its validity. First, the hair and seta loss in laying workers was
not as dramatic as in physogastric queens, yet cuticular marks in workers
were equally well developed. Secondly, the outline of areas of hair loss did
not approximate the outline of the cuticular marks.
Becker (1937a) suggested that the cuticular marks of queens develop in
response to friction between adjacent margins of gastral tergites due to dis-
tension of the physogastric queen’s gaster. I observed no such friction or
contact between tergal margins of laying workers, so this hypothesis also
seems unlikely.
The development of cuticular marks in vespine queens is postulated by
Ishay and Shimony (1982) to result from the conversion of pteridines in the
yellow pigment granules to purines required for the synthesis of DNA. The
conversion of the pteridine components of the granules results in the ob-
served change in the structure of the granules. Significant amounts of nucleic
acids presumably need to be manufactured for the large number of eggs
produced by late-season foundresses. My findings that structural changes in
the granules accompany ovarian development and oviposition in workers,
as well as in queen yellowjackets, support this hypothesis.
The development of cuticular marks in V. maculifrons workers from the
present study seems unrelated to the development of these marks in ovi-
positing wasps. The V. maculifrons workers with marks exhibited no sig-
nificant ovarian development or loss of hair and setae from the gastral terga,
and these workers did not differ from normal nestmates in level of foraging
VOLUME 91, NUMBER 4
403
activity (deduced from amount of wing wear). Cuticular marks developed
primarily on gastral tergum I in V. maculifrons workers, while these were
most prominent on the more posterior segments of laying workers of V.
vulgaris and of physogastric queens. The change in structure of the pigment
granules from the marks of V. maculifrons workers seems to differ funda-
mentally from that occurring in ovary-developed queens and workers; gran-
ules from V. maculifrons exhibited virtually none of the discrete structure
normally visible. Finally, the marks were confined to a significant proportion
of the workers from only two V. maculifrons colonies collected; these col-
onies did not differ in any obvious respects from 1 3 other conspecific colonies
collected during the same season. From these data I conclude that the cu-
ticular marks of the V. maculifrons workers resulted from an anomaly during
deposition of the yellow pigment layer, perhaps due to a genetic mutation.
Ishay and Shimony (1982) report similar aberrations in patterns of pigmen-
tation and structure of the granules in Vespa orient alis F. workers and males.
If components of the yellow pigment granules are indeed shunted to pro-
duction of DNA in ovipositing wasps, then the pigment may be regarded as
a vital storage substance. Given the variable proportions of yellow coloration
in relation to dark coloration among vespine species, this storage product
may be more limiting in some species than in others. Indeed, we may predict
that the development of cuticular marks in those species with relatively low
proportions of yellow-pigmented cuticle (e.g. V. vulgaris, V. flavopilosd) will
be more extensive than in those species with high proportions of yellow-
pigmented cuticle (e.g. V. germanica). My preliminary observations suggest
that this prediction will hold true.
ACKNOWLEDGMENTS
I thank Mary Ann Moran and James M. Carpenter for reviewing the manuscript. Logistic
and financial support was provided by Roger A. Morse of Cornell University’s Dyce Honey
Bee Laboratory and by the Departments of Entomology at Cornell University and the University
of Georgia. Valuable technical assistance with SEM work was provided by Mary Kay Campenot.
LITERATURE CITED
Becker, E. 1937a. Die rotbraune Zeichnung der Wespennestmiitter, eine durch mechanischen
Reiz ausgeloste Pigmentablagerung in Liesegangschen Ringen. Z. Vergl. Physiol. 24:305-
318.
Becker, E. 1937b. Tiber das Pterinpigment bei Insekten and die Farbung und Zeichnung von
Vespa im besonderen. Z. Morph. Okol. Tiere 32:672-751.
Cumber, R. A. 1949. The biology of humble-bees, with special reference to the production
of the worker caste. Trans. R. Ent. Soc. London 100:1-45.
Edwards, R. 1980. Social Wasps, Their Biology and Control. Rentokil Ltd., East Grinstead,
England, 398 pp.
Heldmann, G. 1934. Einiges fiber Wespenverbreitung and Wespenzeichnung. Zool. Anz. 108:
266-273.
404
NEW YORK ENTOMOLOGICAL SOCIETY
Ishay, J. S. and T. B. Shimony. 1982. Changes in the yellow pigment of the Oriental Hornet,
Vespa orient alls, induced by aging, mutations and allopurinol (Hymenoptera; Vespidae).
Ent. Generalis 7:317-325.
Marchal, P. 1 896. La reproduction et 1’evolution des guepes sociales. Arches. Zool. Exp. Gen.
4:1-100.
Ross, K. G. and P. K. Visscher. 1983. Reproductive plasticity in yellowjacket wasps: a
perennial, polygynous colony of Vespula maculifrons. Psyche. 90:179-191.
Shimony, T. B. and J. S. Ishay. 1981. Pigment granules in the tegumental yellow strips of
social wasps: a scanning electron microscopic study. Z. Mikrosk.-Anat. Forsch. 95:310-
319.
Spradbery, J. P. 1973. Wasps, An Account of the Biology and Natural History of Solitary and
Social Wasps. University of Washington Press, Seattle, Washington, 408 pp.
Velleman, P. F. and D. C. Hoaglin. 1981. Applications, Basics, and Computing of Exploratory
Data Analysis. Duxbury Press, Boston, Massachusetts, 354 pp.
Received April 29, 1983; accepted August 17, 1983.
NEW YORK ENTOMOLOGICAL SOCIETY
91(4), 1983, pp. 405-412
ECOLOGICAL AND SENSORY ASPECTS OF PREY CAPTURE BY
THE WHIRLIGIG BEETLE DINEUTES DISCOLOR
(COLEOPTERA: GYRINIDAE)
Steven A. Kolmes
Department of Zoology, University of Wisconsin,
Madison, Wisconsin 53706
— Whirligig beetles prey on a variety of small soft-bodied invertebrates that enter
their surface film habitat from above or below. They also act as scavengers on dead floating
invertebrates. Deficits in prey capture ability were produced by experimental deprivation of
visual, tactile, or surface vibration cues. Visual deprivation and deprivation of surface vibration
cues produced statistically equivalent deficits in whirligig predatory behavior. When tactile and
surface vibration cues were both unavailable to the beetles, the deficits in their predatory
behavior were more pronounced. These sensory systems are similar to those of Hemipteran
predators that live in the surface film.
The way a predator locates its prey is determined by the predator’s en-
vironment, its endowment of sensory structures, and the nature of its prey.
For whirligig beetles, all three of these factors are unusual. These beetles live
and feed on the surface film of bodies of fresh water, and gyrinid sensory
structures are highly specialized for this environment. Their separate pairs
of compound eyes above and below the water (Fig. 1) possess different
spectral sensitivities suited to their respective surroundings (Bennett, 1967;
Carthy and Goodman, 1964; Pappas, 1974). The Johnston’s organs of their
antennae, running between pedicel and flagellum (Fig. 2), are modified into
extremely sensitive surface vibration detectors (Eggers, 1926; Wilde, 1941)
which can sense vibrations with an amplitude as small as a few microns
(Rudolph, 1967). Gyrinids potentially have available to them prey that enter
the surface film from the air, from underwater or that live in the surface
film itself.
I have examined the predatory activities of the river dwelling species
Dineutes discolor, working in both the field and the laboratory to determine
the natural diet of the beetles and what sensory systems were involved in
their prey capture behavior. I investigated for the use of visual, tactile, and
surface vibration cues in gyrinid predation, all three of which are used by
surface film dwelling water striders or backswimmers (Murphey, 1971a,
1971b, 1973; Murphey and Mendenhall, 1973). Prey animals in the surface
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406
NEW YORK ENTOMOLOGICAL SOCIETY
Fig. 1 . Front and side views of the head region of Dineutes discolor, showing the separate
dorsal and ventral pairs of compound eyes.
film are known to generate distinctive surface vibrations by their movements
(Lang, 1980) which constitute an unusual stimulus available to any predator
in contact with the water’s surface.
MATERIALS AND METHODS
I Studied an aggregation of whirligigs located on the Wisconsin River near
Arena (Iowa Co.). Field observations were carried out with binoculars; for
laboratory study beetles were removed from the river and maintained on a
diet of live flightless Drosophila. Of the approximately 600 beetles examined
for identification during the summers of 1976 and 1977, all but two were
Dineutes discolor. This group composition differs from the multispecies rafts
found in a lake habitat by Heinrich and Vogt (1980).
Beetles observed feeding in the laboratory belonged to one of four treat-
ment groups. The first was normal beetles, which underwent no manipulation
of their sensory structures. The second group was temporarily visually de-
prived, by exposing them to light from a Westinghouse DXC 500-W Pho-
toflood from a distance of 30 cm. The exposure was carried out in a special
reflector-lined aquarium freshly filled with cool water, so that the beetles
were protected from elevated temperatures (they always submerged when
the light was turned on). Once bright-light adapted, this group was observed
feeding under a dim red light for 9 min. The combination of bright-light
adaptation, a short observation period, and the insensitivity of the gyrinid
eye to red light (Bennett, 1967; Carthy and Goodman, 1964) produced the
visual deprivation. The third group was deprived of surface vibration cues
by having their antennal flagellae removed. The Johnston’s organ stretched
between pedicel and flagellum (Fig. 2) depends upon relative motions of
these two parts to detect surface vibrations (Eggers, 1926; Wilde, 1941). The
fourth group was deprived of sensory cues by removal of both their antennal
VOLUME 91, NUMBER 4
407
Fig. 2. The antenna of Dineutes discolor. Labelled parts are flagellum (F), pedicel (P) and
socketed flotation hairs of the pedicel (FH).
pedicels and flagellae. Inspection with a scanning electron microscope re-
vealed that the row of hairs on the pedicel have the typical individually
socketed structure of mechanoreceptors, and so beetles with both pedicels
and flagellae removed were deprived of tactile cues from these hairs as well
as surface vibration cues. (Unfortunately it was impossible to eliminate the
hairs on the pedicel and leave the surface vibration detectors intact, as these
hairs supply the buoyancy that keeps the antennae afloat, and removing
them causes unpredictable changes of the height at which the antennal ped-
icels are floating.)
Beetles were observed in the laboratory one at a time, feeding on live
flightless Drosophila in a 122 x 122 x 30 cm tank that rested on inflated
inner tubes to insulate it from extraneous environmental vibrations. Ap-
proximately 150 prey captures were recorded for each of the four treatment
groups, with ten to twenty beetles comprising each group. Three or four days
of food deprivation before observation periods was adequate to ensure hun-
gry but otherwise healthy beetles.
408
NEW YORK ENTOMOLOGICAL SOCIETY
Table 1. Objects close circled by D. discolor under natural conditions.
Objects close-circled
Consumed
Not consumed
Live invertebrates less than 3 mm long
(gnats and collembolans)
4
3
Live mosquito larvae
3
88
Live flies, 3-5 mm long
2
—
Live flies, 5-10 mm long
1
1
Live coccinellid beetles
—
3
Dead spiders
18
—
Dead unidentified invertebrates
5
12
Duckweed plants
—
33
Unidentified objects
46
1,103
In both laboratory and field, D. discolor performs a stereotypic predatory
sequence I call a close circle. The beetle approaches with its head directed
toward its prey. Keeping its head very close to the prey the beetle circles
around the prey. The raptorial forelegs of the whirligig, normally held in
grooves under its body, reach out toward the prey during this circle. In the
field, one, occasionally two, or rarely more, close circles were performed
about prey animals before they were captured and consumed. For the lab-
oratory data I compared the number of close circles required for prey capture
as a measure of how the various sensory deprivations affected predatory
ability.
RESULTS
A total of 1,322 close circles of objects in the surface film was recorded
during the course of my field observations (Table 1). Identification of objects
being close circled was often difficult due to their small size, as the categories
in the table attest. Prey were only scored as “live” if I saw them move
independently before being close circled by a gyrinid.
All of D. discolor 's prey were soft-bodied invertebrates. They entered the
surface film both from above (gnats, collembolans, flies, etc.) and below
(mosquito larvae). Although small pieces of vegetation were close circled, I
never observed the beetles eating any plant material. Beetles occasionally
dove below the surface, but even when they were surrounded by animals
they would have attacked at the surface film (mosquito larvae), I never saw
a submerged gyrinid attack a prey animal.
In the laboratory, sensory deprivation in visual, surface vibration, and
tactile stimuli all resulted in more close circles being required for prey cap-
ture. Figure 3 shows these data displayed as the cumulative probability of
prey capture vs. the number of close circles performed, which is a form of
VOLUME 91, NUMBER 4
409
LU
cr
3
I-
Q_
<
O
>
LU
CL
CL
o
_l
CD
<
GO
O
CL
CL
LU
>
<
_l
3
3
O
Fig. 3. The cumulative probabilities of prey capture by four experimental groups of Dineutes
discolor fed live flightless Drosophila in the lab. The total number of prey captured is 1 50 ± 5
for each of the groups of beetles.
data organization amenable to analysis using the Kolmogorov-Smirnov test
(Siegel, 1956). Each of the four groups of beetles includes data for 150 ± 5
prey captures.
Both visually deprived beetles and those deprived of their flagellae re-
410
NEW YORK ENTOMOLOGICAL SOCIETY
quired more close circles than normal beetles to capture prey, with signih-
cance levels of 0.025 and 0.001 respectively. The visually deprived beetles
and those deprived of surface vibration cues by flagellar ablation were sta-
tistically indistinguishable from one another in their number of close circles
per prey capture, which can be seen by the virtual overlap of these two lines
in Figure 3.
Beetles deprived of both surface vibration cues and tactile cues by having
their antennae removed required even more close circles to capture prey,
differing from all three previous groups at the 0.001 significance level. Pre-
sumably this deficit reflects an additional sensory deprivation due to the loss
of mechanoreceptive hairs on their antennal pedicels.
DISCUSSION
Whirligig beetles capture and consume prey that enters the surface film
from below and above, thereby taking advantage of aquatic insects that need
to reach the surface for air, as well as flying or terrestrial invertebrates that
may be blown down onto the water or landed in order to lay eggs. My limited
ability to clearly distinguish small invertebrates and their movements from
several meters distance undoubtedly skews Table 1 towards larger prey items
and towards dead food materials more than is the actual case. Nonetheless,
D. discolor clearly acts as both a predator and as a scavenger, and seems to
feed on rather soft-bodied forms.
Virtually all of the live prey of D. discolor listed in Table I are very
ephemeral objects from a whirligig’s viewpoint. When approached by a
predator in the surface film, mosquito larvae submerge, collembolans jump,
gnats take off from the water’s surface, etc. If a gyrinid is to successfully
capture any prey, its predatory sequence must be as rapid as possible. The
fewer number of close circles a gyrinid performs before prey capture, the
more prey it will probably consume. This argument may also obtain for the
dead invertebrates scavenged by the beetles. Not only do D. discolor live in
moving water, but they also feed in aggregations and a dead invertebrate
not captured immediately is apt to be swept downstream or consumed by
a conspecific.
Previous studies have shown that backswimmers and water striders are
capable of utilizing a variety of cues in locating and capturing prey, including
visual, tactile and surface vibration cues (Murphey, 1971a, 1971b, 1973;
Murphey and Mendenhall, 1973). Potential prey insects in the surface film
generate distinctive vibration spectra (Lang, 1980) and the processes of
surface vibration orientation are beginning to be unravelled (Lang, 1980;
Reinig and Uhlemann, 1973; Weise, 1974).
D. discolor appears to use visual, tactile, and surface vibration cues in prey
capture, as indicated by the increased number of close circles required in
my laboratory groups. An alternative hypothesis for these deficits that must
VOLUME 91, NUMBER 4
411
be considered is that the beetles were physically damaged in a more general
sense by their laboratory treatments, but I do not believe this to be the case.
Normal beetles in Figure 3 show probabilities of prey capture with one and
two close circles, respectively, that correspond to 77 and 129 out of 155
total prey captured. The normal laboratory animals were therefore capturing
prey with one or two close circles, much as beetles do in nature. The visually
deprived beetles were exposed only to bright light and my apparatus ensured
that they were not subjected to elevated temperatures, and they show deficits
in capture behavior statistically indistinguishable from those of the beetles
with their antennal flagellae removed. Experimental beetles who were op-
erated on were as vigorous as normal animals, and did not display an in-
creased mortality. Ideally I would have performed sham operations, but
there is no obvious way to do so when the procedure involves an ablation
rather than an incision.
Congeneric whirligigs such as Dineutes hornii are largely quiescent during
the day and forage at night (Heinrich and Vogt, 1 980). This nocturnal feeding
may rely upon different sensory’ modalities than the diurnal predation by D.
discolor. Whether whirligigs forage at night or during the day appears to
depend on the motion of the water in which they live; those that live in a
current must climb out of the water onto emergent vegetation at night in
order to avoid being swept downstream while unable to use visual cues to
maintain their position relative to the shore (Brown and Hatch, 1929; Folk-
erts and Donovan, 1973).
The wave patterns produced by swimming gyrinids have been well de-
scribed by Tucker (1969), and the possibility of these waves reflecting off
prey in a surface film borne echolocation system has seemed a natural pos-
sibility. However, the wavelengths of the waves produced by swimming
gyrinids are very large compared to the portions of the natural prey items
in contact with the surface film (siphons for mosquito larvae, legs for gnats,
etc.) and so efficient wave reflection for echolocation seems unlikely. It is
more probable that any echolocation used by gyrinids involves locating larger
objects in the surface film, like rocks, conspecifics, etc.
ACKNOWLEDGMENTS
I am grateful to Professors Jeffrey R. Baylis, Jack P. Hailman, and Stanley Carlson who have
supplied me with assistance and advice. I am also indebted to Clyde S. Gorsuch, Scott R.
Robinson and Katherine C. Noonan for reading and commenting on various drafts of this
paper. Cheryle Hughes produced the excellent illustrations.
LITERATURE CITED
Bennett, R. 1 967. Spectral sensitivity studies on the whirligig beetle, Dineutes ciliatus. J. Insect
Physiol. 13:621-633.
Brown, C. R. and M. H. Hatch. 1929. Orientation and “fright” reactions of whirligig beetles
(Gyrinidae). J. Comp. Psychol. 9:159-189.
412
NEW YORK ENTOMOLOGICAL SOCIETY
Carthy, J. D. and L. J. Goodman. 1964. An electrophysiological investigation of the divided
eye of Gyrinus bicolor. J. Insect. Physiol. 10:431-436.
Eggers, F. 1926. Die mutmassliche Funktion des Johnstonaschen Sinnesorgans bei Gyrinus.
Zool. Anz. 63:184-192.
Folkerts, G. W. and L. A. Donavan. 1973. Resting sites of stream dwelling Gyrinids (Co-
leoptera). Entomol. News 84:198-201.
Heinrich, B. and F. D. Vogt. 1980. Aggregation and foraging behavior of whirligig beetles
(Gyrinidae). Behav. Ecol. Sociobiol. 7:179-186.
Lang, H. H. 1 980. Surface wave discrimination between prey and nonprey by the backswimmer
Notonecta glauca L. (Hemiptera: Heteroptera). Behav. Ecol. Sociobiol. 6:233-246.
Murphey, R. H. 1971a. Motor control of orientation to prey by the waterstrider Gerris remigis.
Z. vergl. Physiol. 72:159-167.
Murphey, R. H. 1971b. Sensory aspects of the control of orientation to prey by the waterstrider
Gerris remigis. Z. vergl. Physiol. 72:168-185.
Murphey, R. H. 1973. Mutual inhibition and organization of a non-visual orientation in
Notonecta. J. Comp. Physiol. 84:31-40.
Murphey, R. K. and B. Mendenhall. 1973. Localization of receptors controlling orientation
to prey by the backswimmer Notonecta undulata. J. Comp. Physiol. 84:19-30.
Pappas, C. D. 1974. Fine structure of the dorsal and ventral eyes of Dineutes assimilis Kirby
(Coleoptera, Gyrinidae). Unpublished Ph.D. dissertation. University of Illinois at Ur-
bana-Champaign.
Reinig, H. J. and H. Uhlemann. 1973. Uber das ortungsvermogen des Taumelkafers Gyrinus
substriatus Steph. (Coleoptera, Gyrinidae). J. Comp. Physiol. 84:281-298.
Rudolph, P. 1872. Zum ortungsverfahren von Gyrinus substriatus (Steph). Z. vergl. Physiol.
56:341-371.
Siegel, S. 1956. Nonparametric Statistics for the Behavioral Sciences. McGraw Hill Book
Company, New York.
Tucker, V. A. 1969. Wave-making by whirligig beetles (Gyrinidae). Science 166:897-899.
Weise, K. 1974. The mechanoreceptive system of prey localizations in Notonecta. II. The
principle of prey localization. J. Comp. Physiol. 92:317-325.
Wilde, J. de. 1941. Contribution to the physiology of the Johnston organ and its part in the
behaviour of the Gyrinus. Arch. Neer. Physiol. Homme Animaux 25:381-400.
Received May 23, 1983; accepted September 14, 1983.
NEW YORK ENTOMOLOGICAL SOCIETY
91(4), 1983, pp. 413-417
IRBISIA KNIGHTI, A NEW MIRINE PLANT BUG
(HETEROPTERA: MIRIDAE) FROM THE PACIFIC NORTHWEST
Michael D. Schwartz and John D. Lattin
Systematic Entomology Laboratory, Oregon State University,
Corvallis, Oregon 97331
Abstract.— T\iQ new species Irbisia knighti is described and its distribution given.
Irbisia knighti, new species
(Figs. 1-8)
Diagnosis. Similar to I. solani (Heidemann) 1910 in size and surface
features but is consistently recognizable by brachyptery of both sexes and
white vestiture. I. knighti is separated from the brachypterous form of I.
sericans (StM) 1858 by the slightly punctate pronotum and white vestiture
of the former species (Figs. 1 and 2).
Description. Holotype male. Brachypterous. Length 3.90-4.63 (all mea-
surements in millimeters), width 2.18-2.30, shining black, moderately cov-
ered with shining white setae. HEAD: width across eyes 1.25-1.30, vertex
0.59-0.63, dorsal width of eye 0.35-0.36; triangular, smooth; temporal areas
distinct, bordered basally by transverse short sulcus, glabrous median depres-
sion indistinct; basal carina moderately prominent, slightly rounded, decli-
vous to level of vertex, straight, eyes broadly joined to frons, posterolateral
margins arcuate posteriorly in dorsal view; distance from tylus to ventral
margin of eye 0.61-0.63, height of the eye 0.50-0.53, maximum interocular
width (anterior view of frons) 0.90-0.95. ROSTRUM: length 1.70-1.83,
black, surpassing mesocoxae to just attaining apices of metacoxae. ANTEN-
NAE: black; I, length 0.50-0.55, apices fuscous; II, 1.45-1.55; III, 0.78-
0.93; IV, 1.02-1.20. PRONOTUM: length 0.86-0.88, anterior width 0.93-
0.95, posterior width 1.45-1.55; subconical, slightly flattened, distinctly or
confluently punctate, lateral margins straight, broadly rounded at junction
with propleura, anterior angles rounded, indistinct in dorsal view, gently
sulcate in lateral view; calli slightly convex, smooth, narrowly confluent
anteriorly, with transverse depression on inner, and foveate depression of
outer anterior angles reaching antero-lateral margins of pronotum. LEGS:
black; coxae and femora with apices testaceous; tibiae testaceous with bases
of knees black; tarsi black. HEMELYTRA: membrane reduced, extending
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NEW YORK ENTOMOLOGICAL SOCIETY
slightly beyond apex of cuneus, cells not developed; apex of abdomen ex-
posed in dorsal view. VESTITURE: dorsum clothed with suberect white
setae; head and pronotum sparsely covered with thin setae; hemelytra mod-
erately covered with sericeous setae basally, these setae grading into and
being replaced by thin setae apically; venter moderately clothed with white
setae; eyes with sparse minute pubescence between facets. GENITALIA:
Left paramere (Fig. 3): evenly curved; sensory lobe developed but not set
olf by a dorsal suture, with numerous bristles; shaft truncate at apex, with
numerous setae. Right paramere (Fig. 4): subcylindrical; with a single short
apical process and a few short lateral setae. Vesica (Fig. 5): two smooth
membranous lobes, each with a small preapical side lobe; sclerotized process
furcate, primary process strongly serrate; ductus seminis expanded subapi-
cally.
Brachypterous female. More robust than male, but very similar in color,
structure and vestiture. Length 4.40-4.95, width 2.40-2.50 HEAD: width
across eyes 1.35-1.36, vertex 0.66-0.68, dorsal width of eye 0.35-0.38,
distance from tylus to ventral margin of eye 0.66-0.68, height of eye 0.50-
0.54, maximum interocular width 0.96-0.98. ROSTRUM: length 1 .88-1 .93,
barely reaching apices of metacoxae. ANTENNAE: I, length 0.55-0.58; II,
1.53-1.55; III, 0.85-1.00; IV, 1.03-1.15. PRONOTUM: length 0.93-0.96,
anterior width 1.02-1.04, maximum width 1.65-1.70. GENITALIA: Scler-
otized rings (Fig. 6): sclerotized, area within rings narrow, postero-lateral
region of rings with heavily sclerotized partial rings; dorsal wall extending
anterior beyond rings, with a broadly truncate mesal margin. Maximum
length 0.48, maximum width 0.99. Posterior wall (Fig. 7): A— structure
curving, broadly truncate on ventral margin, with slight mesal indentation.
B— structure protruding posteriorly, without indentations on ventral pos-
terior surface; median process gradually expanded dorsad. Maximum length
0.22, maximum width 0.60.
Holotype. Male, Washington, Pacific Co., Nahcotta Oyster Research Sta-
tion, 14 June 1979, ex. Agropyron repens, M. D. Schwartz (CAS type no.
15024). Paratypes: 15 males, 17 females, same data as holotype (AMNH,
CAS, CNC, OSU, USNM).
Other specimens examined. 232 specimens were examined from the fol-
Figs. 1-8. 1. Dorsal view of body, /. knighti. 2. Dorsal view of pronotum, I. sericans. 3.
Dorsal view of left paramere, I. knighti. 4. Dorsal view of right paramere, /. knighti. 5. Ventral
view of vesica, I. knighti. 6. Posterior view of sclerotized rings and dorsal labiate plate, /.
knighti. 7. Posterior view of inter-ramal sclerites, I. knighti. 8. Distribution map, I. knighti.
Specimens illustrated are from Washington, Pacific Co. (OSU).
VOLUME 91, NUMBER 4
415
416
NEW YORK ENTOMOLOGICAL SOCIETY
lowing localities (Fig. 8): CANADA: BRITISH COLUMBIA: Jesse Island—
(Nanaimo), 3 June 1925, 19 (USNM); G. J. Spencer, 16, 399 (UBC); 16
(UCB). Royal Oak, 3 June 1959, L. A. Kelton, 366, 399 (CNC). Victoria, 4
August 1912, J. B. Wallis, 19 (USNM); 25 June 1925, K. F. Auden, 399
(UCB); 19 (USNM); W. Downes, 16 (USNM); 15 July 1923, K. F. Auden,
19 (UCB); 3 June 1933, O. H. Swezey, 299 (CAS); 15 May 1928, W. Downes,
16, 19 (USNM). UNITED STATES: CALIFORNIA: Humboldt Co., 5 mi.
W Loleta— tidal mud flat, 9 June 1963, Ball, Freitag, Lister, McDonald, 19
(UALB). OREGON: Lane Co., 12 mi. N Florence— Big Creek Rd. — Siuslaw
Nat. For., 30 May 1979, ex. Holcus lanatus, M. D. Schwartz, 16, 299 (OSU).
Tillamook Co., 2.3 mi. N Neskowin on rt. 101,21 May 1979, ex. Festuca
rubra, Holcus lanatus, Poa pratensis, M. D. Schwartz, 2766, 899 (OSU); 0.1
mi. N Rockway on rt. 101, 1 June 1979, M. D. Schwartz, 16 (OSU); Sand
Beach Cmpgrd. — 2 mi. W Sand Lake, 13 June 1972, ex. Poa sp. & Carex
sp., J. D. Lattin, 966, 2199 (OSU); Sand Lake, 7 July 1962, G. C. Eickwort,
366, 299 (MSU); Tierra del Mar, 13 June 1972, J. D. Lattin, 16 (OSU).
WASHINGTON: King Co., Seattle, 20 May 1916, 1466, 899 (OSU); 24 May
1928, 16 (OSU); 25 May 1928, 16 (OSU). Pacific Co., Bay Center, 30 June
1931, 19 (OSU); Nahcotta, 14 June 1979, ex. sweeping grasses just above
tide line on bay shore, R. T. Schuh, 1466, 699 (AMNH); G. M. Stonedahl,
16 (OSU); Wallapa Bay— near Nahcotta, 12 June 1971, W. J. Turner, M.
Wiebers, J. A. Novak, D. N. Feno, 2966, 2599 (WSU). Skagit Co., Mt.
Vernon, 25 May 1961, M. C. Lane, 16 (OSU).
Etymology. Named in honor of the late Dr. Harry Hazelton Knight of
Iowa State University, the renowned worker of the North American Miridae.
The junior author, during his undergraduate years at Iowa State, had the
good fortune to work with Dr. Knight who kindled the former’s interest in
the Hemiptera.
Discussion. The genus Irbisia contains 23 species; all are Nearctic (Schwartz,
1981). The junior author found this species with specimens borrowed from
WSU. In addition, the senior author upon examining the Knight collection,
at the United States National Museum at Washington, D.C. found five
specimens of Irbisia with Dr. Knight’s characteristic red labels indicating a
“new species.” These specimens are of I. knighti.
This species is narrowly distributed along the coastal flatlands of the Pacific
Northwest. It occupies a very restricted portion of the Vancouveran Zone
as defined by Van Dyke (1939). Adult specimens were collected from these
hosts: Agropyron repens (L.) Beauv., Carex sp. (a sedge), Festuca rubra L.,
Holcus lanatus L., Poa pratensis L., and Poa sp. None of these grasses {Carex
sp. is a doubtful host) are obligatorily associated with the apparent restricted
habitat of I. knighti. Collection dates are from 15 May to 15 July. All
localities are from sea level to 45 meters (150 feet) elevation.
VOLUME 91, NUMBER 4
417
ACKNOWLEDGMENTS
The authors wish to thank Ms. Bonnie B. Hall, for the dorsal view illustration. The following
institutions and curators kindly loaned specimens: American Museum of Natural History, New
York (AMNH), Dr. Randall T. Schuh; California Academy of Sciences, San Francisco (CAS),
Dr. Paul H. Amaud, Jr.; Canadian National Collection, Ottawa (CNC), Dr. Leonard A. Kelton;
Montana State University, Bozeman (MSU), Ms. Sharon D. Rose; University of Alberta, Ed-
monton, Canada (UALB), Dr. George E. Ball; University of British Columbia, Vancouver,
Canada (UBC), Dr. Geoffrey G. E. Scudder; University of California, Berkeley (UCB), Dr. Jerry
A. Powell; United States National Museum, Washington, D.C. (USNM), Dr. Richard C.
Froeschner and Mr. Thomas J. Henry; James Entomological Collection, Pullman, Washington
(WSU), Dr. William J. Turner.
LITERATURE CITED
Heidemann, Otto. 1910. Description of a new capsid. Proceedings of the Entomological Society
of Washington 12:200-201.
Schwartz, Michael D. 1981. A revision of the black grass bug genus Irbisia Reuter (Heteroptera:
Miridae). Unpublished Master’s thesis, Corvallis, Oregon State University, 222 pp.
St^l, Carl. 1858. Beitrug zur Hemipteren — Fauna Sibiriens und des Russischen N ord- Amerika.
Entomologische Zeitung herausgegben von dem entomologischen Vereine zu Stettin 19:
175-198.
Van Dyke, Edwin C. 1939. The origin and distribution of the coleopterous insect fauna of
North America. Proceedings of the Sixth Pacific Science Congress, Pacific Science As-
sociation 4:255-268.
Received May 19, 1983; accepted September 30, 1983.
NEW YORK ENTOMOLOGICAL SOCIETY
91(4), 1983, pp. 418-423
DESCRIPTIONS OF THE NYMPHAL INSTARS OF
OECLEUS BOREALIS {YiOMOVTEKK: FULGOROIDEA: CIXIIDAE)*
Stephen W. Wilson, James H. Tsai, and Catherine R. Thompson
Department of Biology, Central Missouri State University,
Warrensburg, Missouri 64093 and
Agricultural Research and Education Center, IFAS, University of Florida,
Ft. Lauderdale, Florida 33314
Abstract.— The, 5 nymphal instars of Oecleus borealis Van Duzee are described and illustrated.
Features useful in separating nymphal instars include the size, and form of teeth and spines on
the profemora and tibiae, size of body and wingpads, and the numbers of pits, and metatar-
someres.
Oecleus borealis Van Duzee has been recorded from New York south to
Florida and west to Michigan, Kansas, and Texas (Kramer, 1977). Adults
have been collected on apple (Malus sp.), hickory {Carya sp.). New Jersey
tea {Ceanothus americanus L.) and desert willow (Chilopsis linearis DC.)
(Kramer, 1977). Information on the immatures of any cixiid is very limited
as they are subterranean: Cumber (1952) described the immatures of Oliarus
atkinsoni Myers, Myers (1929) described some immature stages of Mne-
mosyne cubana StM and Bothriocera signoreti Stal. Wilson and Tsai (1982)
recently provided detailed descriptions of the immatures of Myndus crudus
Van Duzee. We describe the five immature stages of Oecleus borealis Van
Duzee in this paper.
MATERIALS AND METHODS
Specimens to be described were collected by Tsai and Thompson at two
localities in peninsular Florida (collecting data given below) and preserved
in 70% ethyl alcohol. The first instar is described in detail, but only major
changes from previous instars are described for subsequent instars. Com-
parative statements refer to previous instars (e.g., more numerous). Mea-
surements are given in mm as mean ± SE. Length was measured from apex
of vertex to apex of abdomen, width across the widest part of the thorax.
' Ra. Agric. Exp. Stn. Journal Series No. 4822.
The publication costs of this article were defrayed in part by page charge payment. This article
must therefore be hereby marked “Advertisement” in accordance with 18 U.S.C. §1734 solely
to indicate this fact.
VOLUME 91, NUMBER 4
419
Figs. 1-3. Nymphs of O. borealis. 1 . First Instar. 2. Second Instar. 3. Third Instar, a. Nymph,
b. Median aspect of apex of profemur. Vertical bars = 0.5 mm.
and thoracic length along the midline from the anterior margin of the prono-
tum to the posterior margin of the metanotum.
Nymphs of O. borealis were first discovered in January 1975 by the junior
author (CRT) in sand pine woods [Pinus clausa (Chapman)], 27-32 km east
of Silver Springs, Marion Co. Florida in Ocala National Forest. The sub-
terranean nymphs were found 30-45 cm deep in white sand areas where
vegetation consisted of saw-palmetto [Serenoa repens (Bartram)] and turkey
oak (Quercus laevis Walter) as well as sand pine. Nymphs were subsequently
found in similar habitats in Orange and Broward counties. An adult reared
from a late instar nymph was identified as O. borealis by J. P. Kramer,
USNM. The economic importance of this insect is unknown.
Collecting data for the specimens used in the descriptions are: FLORIDA:
Broward Co.: Ft. Lauderdale, 28 May 1982 (2 first instars), 23 July 1982 (1
third instar); Orange Co.: Orlando, 18 November 1982 (5 fourth instars, 12
fifth instars); Marion Co.: Ocala National Forest, 15 September 1982 (1
fourth instar), 18 June 1979, (1 first instar, 7 second instars, 3 fourth instars,
3 fifth instars; same locality, no date— 1 third instar, 3 fifth instars).
DESCRIPTIONS OF THE NYMPHAL INSTARS
First instar (Fig. 1). Mean length 0.69; thoracic length 0.27; width 0.26.
Three specimens examined.
Form elongate, subcylindrical, slightly flattened dorsoventrally, widest
across junction of meso- and metathoraxes. Vertex, frons, thoracic nota, and
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NEW YORK ENTOMOLOGICAL SOCIETY
abdominal tergites with a few (less than 10) shallow indistinct pits. Body
white.
Vertex broadly rounded anteriorly, slightly narrowing posteriorly. Frons
with lateral margins convex and forming shelf-like carinae beneath eyes.
Clypeus narrowing distally. Beak apparently 3-segmented [based on com-
parison with later instars and first instars of other cixiids e.g., Myndus; see
Wilson and Tsai (1982)]; extending just beyond metacoxae. Eyes reduced,
barely visible in ventral view, reddish. Antennae 3-segmented; scape and
pedicel subcylindrical and subequal; flagellum bulbous basally, filamentous
distally, bulbous portion ca. Vi length of pedicel.
Thoracic nota divided by a longitudinal mid-dorsal line into 3 pairs of
plates. Pronotum longest laterally; each plate subquadrate, anterior margin
almost straight, posterior margin slightly concave, lateral margin broadly
curved. Mesonotum with median length subequal to that of pronotum; each
plate subquadrate, posterior margin slightly concave, lateral margin slightly
convex. Metanotum with median length ca. % that of mesonotum; each plate
subquadrate, lateral margin sharply angled posteromedially. Pro- and me-
socoxae posteromedially directed; metacoxae smaller, obscured by trochan-
ters. Profemora with a slightly swollen appearance. Pro- and mesotibiae very
short, ca. Vi length of tarsi; metatibiae elongated, slightly longer than meta-
tarsi. Tarsi 2-segmented, divisions between tarsomeres obscure; with a pair
of tiny apical claws.
Abdomen apparently 9-segmented (although specimens were cleared in
10% KOH, segments were very difficult to see; the supposed number of
segments is based on comparison with later instars and first instars of other
cixiids (e.g., Myndus; see Wilson and Tsai, 1982), subcylindrical, widest
basally; posterior-most segment surrounding anus.
Second instar (Fig. 2). Length 1.05 ± 0.05; thoracic length 0.46 ± 0.03;
width 0.46 ± 0.03. Seven specimens examined.
Widest across mesothorax. Thoracic nota with pits generally more nu-
merous. Pro- and mesotibiae with lengths subequal to those of tarsi. Oth-
erwise, similar to first instar.
Third instar. (Fig. 3). Mean length 1.95; thoracic length 0.73; width 0.79.
Two specimens examined.
Frons and clypeus with a continuous row of ca. 1 1 tiny black dotlike pits
paralleling lateral margins. Beak with segment 2 ca. 2 x length of segment
1; segment 3 subequal to 2. Antennae with bulbous portion of fiagellum ca.
V4 length of pedicel.
Thoracic nota with pits generally more numerous. Pronotum with each
plate bearing 4 pits paralleling median line and ca. 13-15 total pits on plate.
Mesonotum with median length ca. 1 Vi x that of pronotum, distinctly lobate
posterolaterally. Metanotum with median length ca. % that of mesonotum;
each plate with 1 pit in anteromedial corner, ca. 4 pits near middle and ca.
VOLUME 91, NUMBER 4
421
Figs. 4, 5. Nymphs of O. borealis. 4. Fourth Instar. 5. Fifth Instar, a. Nymph, b. Median
aspect of apex of profemur, protibia, and protarsus. Vertical bars = 0.5 mm.
3 pits laterally. Profemora with ca. 0.05 mm curved spine in distal Vi. Pro-
and mesotibiae subequal in length to tarsi; protibiae with small spine in
distal Vi, metatibiae with a row of 5 very small spines apically.
Fourth instar (Fig. 4). Length 2.47 ± 0.19; thoracic length 0.98 ± 0.03;
width 1.03 ± 0.05. Nine specimens examined.
Frons and clypeus with a continuous row of ca. 20 tiny black dots par-
alleling lateral margins. Antennae with scape reduced and ring-like. Prono-
tum with each plate bearing 4 pits paralleling median line and ca. 20 pits
on plate. Mesonotum with each plate bearing 1-2 pits near anteromedial
corner, 1 pit near posteromedial corner, an oblique row of 4 pits near middle
and an irregular row of very shallow pits (apparently absent in some spec-
imens) paralleling lateral margin; wingpad lobate and covering ca. of
metanotal plate laterally. Metanotum with each plate bearing ca. 5 pits.
Profemora with ca. 0. 1 5 mm curved spine in distal Vi and a very small tooth
between spine and apex. Protibiae with a large bifid tooth in distal Vi, a small
tooth proximal to it, and 1 large tooth on median aspect (Figs. 4A, B).
Metatibiae with a row of 5 small black-tipped spines apically. Metatarsi
3-segmented; tarsomere 1 cylindrical with a row of 4 very tiny black-tipped
spines apically; tarsomere 2 cylindrical with a small black-tipped spine on
either side at apex; tarsomere 3 subconical, slightly curved and bearing a
pair of slender apical claws.
Abdominal tergites 6-8 each with dorsoposteriorly oriented white wax-
pads in intermembranous areas posterior to tergal plates; waxpads probably
paired and present but indistinct in earlier instars (see Wilson and Tsai,
1982).
Fifth instar (Fig. 5). Length 3.23 ± 0.11; thoracic length 1.34 ± 0.02;
width 1.52 ± 0.04. Eighteen specimens examined.
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NEW YORK ENTOMOLOGICAL SOCIETY
Frons and clypeus with a continuous row of ca. 16-25 tiny black dots
paralleling lateral margins. Antennae with bulbous portion of flagellum ca.
length of pedieel.
Pronotum with each plate bearing 4 pits paralleling median line and ca.
30 or more total pits on plate (many pits very shallow and obscure). Meso-
notum with each plate bearing 1-2 pits near anteromedial corner, an oblique
row of 5 pits near middle of plate and numerous shallow pits paralleling
lateral margins; wingpad extending to or nearly to apex of metanotal wing-
pad. Metanotum with eaeh plate bearing ea. 6-8 shallow, obscure pits. Pro-
femora with ca. 0.2 mm curved spine in distal Vi and a small unifid or bifid
tooth distal to it on median aspect; on lateral aspeet with 2 small teeth in
basal Vi and 1 small tooth in distal Vi. Protibiae with bifid tooth in basal Vi
in median aspeet, bifid tooth near apex and 1 small tooth in basal Vi of
anterior aspeet, and 1 tooth in basal Vi of lateral aspect. Metatarsomere 1
with an apical row of 5 black-tipped teeth.
Abdomen with pits generally more numerous.
KEY TO THE NYMPHAL INSTARS
1. Metatarsi subdivided into 3 tarsomeres; profemur with small tooth distal to curved
spine (Figs. 4, 5) 2
- Metatarsi subdivided into 2 tarsomeres; profemur without small tooth distal to curved
spine or spine lacking (Figs. 1-3) 3
2. Mesothoracic wingpads extending to or nearly to apex of metathoracic wingpads (Fig.
5); metatarsomere 1 with an apical row of 5 spines Fifth Instar
- Mesothoracic wingpads extending at most % length of metathoracic wingpads (Fig. 4);
metatarsomere 1 with an apical row of 4 spines Fourth Instar
3. Profemur with curved spine in distal V2 (Fig. 3b); thoracic length greater than 0.7 mm
Third Instar
- Profemur without curved spine in distal '/2; thoracic length less than 0.6 mm 4
4. Protibia ca. V2 length of protarsus; thoracic length less than 0.35 mm (Fig. 1)
First Instar
- Protibia subequal in length to protarsus; thoracic length greater than 0.35 mm (Fig. 2)
Second Instar
ACKNOWLEDGMENTS
We would like to thank Drs. F. W. Mead, DPI, Florida Department of Agriculture and
Consumer Services, Gainesville, Florida 32602, T. D. Center, Aquatic Plant Laboratory, ARS
Southern Region, USDA, Ft. Lauderdale, Florida 33314, and J. A. Reinert, AREC, University
of Florida, Ft. Lauderdale, Florida 33314 for reviewing the manuscript and Dr. J. P. Kramer,
Department of Entomology, Smithsonian Institution, Washington, D.C. for identifying the adult
male and reviewing the manuscript.
LITERATURE CITED
Cumber, R. A. 1952. Studies on Oliarus atkinsoni Myers (Hem. Cixiidae), vector of the
“yellow-leaf’ disease of Phonnium tenax Forst. II. The nymphal instars and seasonal
changes in the composition of nymphal populations. New Zealand J. Sci. Technol. 34:
160-165.
VOLUME 91, NUMBER 4
423
Kramer, J. P. 1977. Taxonomic study of the planthopper genus Oecleus in the United States
(Homoptera: Fulgoroidea: Cixiidae). Trans. Am. Entomol. Soc. 103:370-449.
Myers, J. G. 1929. Observations on the biology of two remarkable cixiid planthoppers (Ho-
moptera) from Cuba. Psyche 36:283-292.
Wilson, S. W. and J. H. Tsai. 1982. Descriptions of the immature stages of Myndus crudus
(Homoptera: Fulgoroidea: Cixiidae). J. New York Entomol. Soc. 90:166-175.
Received June 20, 1983; accepted September 7, 1983.
NEW YORK ENTOMOLOGICAL SOCIETY
91(4), 1983, pp. 424-430
ON THE BIOLOGY OF CAVE INHABITING ANTILLOCORINI WITH
THE DESCRIPTION OF A NEW SPECIES FROM NEW GUINEA
(HEMIPTERA: LYGAEIDAE)i
James A. Slater
Section of Systematic and Evolutionary Biology,
University of Connecticut,
Storrs, Connecticut 06268
Abstract.— Botocudo cavernicola is described as a new species from bat guano in a cave in
New Guinea. Comments on the cave habitat are included. This is thought to be the first record
of a lygaeid inhabiting caves in the Eastern Hemisphere. The biology of Cligenes subcavicola
Scudder, Darlington and Hill is discussed. It is reported to feed on the seeds of Piper sp. found
in bat guano in caves and abandoned buildings on Trinidad. The species is reported from a
cave in Peru for the first time where it feeds at least in part on the seeds of species of Ficus.
The only species of Lygaeidae previously known to inhabit caves is Cli-
genes subcavicola Scudder, Darlington and Hill. Recently I have had the
opportunity to observe this species in the field in Trinidad.
Through the kindness of Mr. W. R. Dolling and Dr. T. E. Woodward I
have also been able to study a species that inhabits a bat cave in New Guinea
and which is described below as a new species. To my knowledge these are
the only lygaeids thus far known to live in caves where presumably both
feed on mature seeds that have passed through the digestive tracts of fru-
givorous bats.
All measurements are in millimeters.
Botocudo cavernicola, new species
(Fig. 1)
Body broadly elliptical. Head, pronotum, scutellum, antennal segments I,
II, and III, abdomen and thoracic pleura bright red brown. Apex of scutellum
white. Base of head between ocelli darker. Hemelytral ground color white;
area within corial furrow nearly to outer closely set row of punctures trans-
lucent, widening posteriorly. Corium with strongly contrasting red brown
markings as follows: posterior % of lateral corial margin; a large transverse
' This work was supported in part by a grant from the National Science Foundation.
The publication costs of this article were defrayed in part by page charge payment. This article
must therefore be hereby marked “Advertisement” 'm accordance with 18 U.S.C. §1734 solely
to indicate this fact.
VOLUME 91, NUMBER 4
425
vitta at level of claval commissure extending broadly along lateral margin,
extending mesad to corial furrow and continued caudo-mesal along outer
margins of translucent area to reach apical corial margin just laterad of where
apical margin becomes strongly concave; remainder of apical corial margin
red brown terminating in a very large brown terminal macula. (This col-
oration thus creating a large white ovoid spot distally on corium.) Membrane
translucent. Legs and labium uniformly pale yellow. Fourth antennal seg-
ment (except for base) white. Pronotal and scutellar punctures colored as
rest of surface; hemelytral punctures dark red brown. Body lacking upstand-
ing hairs; dorsal surface nearly glabrous but with scattered, very short, se-
riceous hairs present, these more prominent on head, where anteriorly di-
rected.
Head non-declivent, tylus nearly attaining distal end of first antennal
segment. Eyes sessile, in contact with antero-lateral pronotal angles. Length
head 0.50, width 0.68, interocular space 0.40. Pronotum broadly trapezoidal,
lateral margins very strongly narrowing anteriorly, slightly sinuate; no trans-
verse impression; posterior margin evenly and shallowly concave. Length
pronotum 0.68, width 1.40. Scutellum lacking a median elevation. Length
scutellum 0.84, width 0.81. Hemelytra with lateral corial margins nearly
straight, explanate. Apical corial margin deeply concave on inner third.
Membrane slightly exceeding end of abdomen. Length claval commissure
0.20. Midline distance apex clavus-apex corium 0.70. Midline distance apex
corium-apex membrane 0.60. Metathoracic scent gland auricle strongly bent
posteriorly, acute at distal end. Evaporative area small, occupying only a
small area around auricle, outer margin slightly convex. Fore femora slender,
mutic. Labium extending well between mesocoxae. Length labial segments
I 0.40, II 0.46, III 0.36, IV 0.26. Antennae slender, terete. Length antennal
segments I 0.36, II 0.66, III 0.56, IV 0.66. Total body length 3.40.
Holotype 9. PAPUA NEW GUINEA: West Sepik District of Tedi nr.
Tabubil VII-IX. 1978 (N. Plumley) (Woltem Cave: on moist guano). In
British Museum (Natural History). Paratype: 1 female. Same data as holo-
type. In J. A. Slater collection.
Discussion. This strikingly colored species is readily separable from any
Botocudo species yet described from the Eastern Hemisphere. It is larger
than any known species; of those previously described only assimilans Ber-
groth (Philippines) exceeds 3 mm in length. However, assimilans has a
bicolored pronotum in which the posterior pronotal lobe has a broad whitish
fascia and brown rather than pale yellow front legs. Bergroth (1918) relates
assimilans to signandus Distant.
China (1930) in his description of swezeyi from Samoa states that it is
“readily distinguishable from all other species by the uniformly red-brown
pronotum and scutellum.” This is true of cavernicola. However, the two
species are readily distinguishable. B. swezeyi is only 2.3 long (the subspecies
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NEW YORK ENTOMOLOGICAL SOCIETY
major reaches 2.7), has a fuscous brown rather than white fourth antennal
segment, shining meso- and metapleura and dark spots on the pronotal
humeri. The illustration in China’s paper shows a straight apical corial
margin but this is probably an artist’s error.
The most interesting feature of this new species is its habitat. It is the first
species of lygaeid that, to my knowledge, has been taken in a cave in the
Eastern Hemisphere.
Mr. Noel Plumley who collected the type material has kindly sent me
information on the habitat from which the material was taken. Woltem Cave
is located at 5°14'S, 141°13'E in the Victor Emanuel Region of Papua New
Guinea. The lygaeids were taken from bat guano in the entrance series near
where a stream bed enters. The guano in the area was damp, moderately
firm in consistency and probably no more than 5 cm in depth. A large number
of small seeds similar in size to millet seeds were present in this guano. Mr.
Plumley says that a large species of flying fox inhabited part of the entrance
series. It is probable that droppings from these flying foxes provided the
seeds upon which the insects were feeding. A smaller bat species occurred
deeper in the cave but lygaeids were not collected in that area.
Thus we appear to have the independent acquisition of members of the
same lygaeid tribe of the habit of feeding in caves upon seeds that have
passed through the digestive tracts of the fruit eating bats.
Cligenes subcavicola Scudder, Darlington and Hill
Scudder, Darlington and Hill (1967) described Cligenes subcavicola from
the Tamana Caves in Trinidad. This was the first record of a lygaeid living
and breeding in underground caves. The Tamana Caves are limestone caves
inhabited by both insectivorous and frugivorous bats. C. subcavicola was
reported as living on the floor of the cave on and in the surface layers of bat
guano in large numbers, in some instances reaching a density of the order
of 100,000 per square meter. These authors did not establish the feeding
habits but did suggest that they may have been feeding on the many seeds
present in the guano or that they were predatory upon organisms in the
guano. They described the egg and commented upon features of the nymphs.
A year after the publication of Scudder, Darlington and Hill’s paper Drs.
C. W. and L. B. O’Brien collected a series of C. subcavicola from Lechuzas
Cave in Peru (Cueva de Lechuzas, Huanuco Province, near Tingo Maria,
12.VII.1968, C. W. and L. B. O’Brien).
Three years later Dr. R. T. Schuh visited Lechuzas Cave and has been
kind enough to make his field notes available to me. In general his obser-
Fig. 1 . Botocudo cavernicola new species. Holotype, dorsal view.
VOLUME 91, NUMBER 4
427
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NEW YORK ENTOMOLOGICAL SOCIETY
vations agree with the situation at the Tamana Caves on Trinidad and at
Woltem Cave in New Guinea. Dr. Schuh’s notes indicate that the main part
of the cave is divided into two distinct caverns. The first chamber is large
(about 150-200 feet long) and rather cubic in design with considerable light
entering from the outside. The second chamber is somewhat smaller with
very little light present. The lygaeids were concentrated in the outer two-
thirds of the first chamber. The floor of the cave in this area was almost
devoid of large seeds which are primarily those of palm and laurel fruits.
These form the principal dietary items for the oil birds Steatornis, which
occupy the inner portion of the first chamber and all of the second. The
floor of the outer two-thirds of the first chamber consisted of thousands of
tiny seeds mixed with soil and bat and parrot guano. Dr. Schuh found the
Cligenes subcavicola population to be almost exclusively confined to this
outer area of the cave where they occurred in “unbelievably large numbers”
(one bagful of soil collected contained literally (!!!) thousands of lygaeids)
and could be found in the “soil” to a depth of 3-4 inches.
Dr. Schuh feels that a great proportion of the small seeds in the lygaeid
habitat were of one or more species of Ficus.
It is of interest to note that Dr. Schuh also collected litter living lygaeids
immediately outside the cave, in fact inside the outermost overhang of the
cave. He found small seeds in the litter and a species of rhyparochromine
present but not a single specimen of Cligenes subcavicola. This is an addi-
tional indication that the latter may be confined to caves and other closed
habitats where bat guano is present.
Recently Dr. R. M. Baranowski discovered a large breeding colony of
subcavicola in an abandoned building at the Simla Subtropical Station in
the Arima Valley of the North Range on Trinidad. Our collecting party (R.
M. Baranowski, R. Clayton, M. Hassey, J. and E. Slater) visited this site on
August 23, 1 982. The building is constructed of concrete blocks. It apparently
was used for photography and sound control work as there is a single entry
with an interior wall to screen out light. The interior is completely dark
during daylight hours. It is functionally a “cave” and is known to local
investigators as the “bat house” because of the large colony of bats present.
C. subcavicola is abundant on and in the upper layers of bat guano as
described by Scudder et al. (1967). The insects climb upward for a foot or
more on the vertical walls and move about actively, each carrying a small
seed impaled on the end of the rostrum. Most of these seeds are of a species
of Piper, the fruit of which forms a considerable part of the diet of the
frugivorous bats inhabiting the building. Nymphs of all instars were present
and there is no doubt but that this is a thriving population that is feeding
upon seeds that have passed through the digestive systems of the bats.
The occurrence of Cligenes subcavicola in caves as far removed from each
other as Trinidad and Peru raises at least two questions. (1). Is the species
VOLUME 91, NUMBER 4
429
confined to caves? (2). If it is there must still be considerable dispersal or
the populations would presumably show geographic differentiation.
C. subcavicola as noted by Scudder et al. (1967) does not show any of the
adaptations usually associated with true cavernicoles such as loss of body
pigmentation and reduction of the wings, eyes, and ocelli. The insects also
do not occur in the deeper parts of either the Tamana or Lechuzas Caves.
This suggests that they may not be confined to caves. When the first report
appeared I had thought this was primarily an adventitious adaptation to a
plentiful food source. However, the occurrence of the same species in bat
guano in two separate places in Trinidad and in a cave in Peru together with
its complete absence in collections from any other habitat (including light
traps where many other antillocorines are taken in abundance) suggests that
the occurrence is certainly not fortuitous but an adaptive strategy of the
species.
The degree of host specificity in the Antillocorini is also very poorly
understood. The only other species of Cligenes (distinctus) does appear to
be restricted to fallen seeds of various species of Ficus. I have collected long
series of this species on several islands of the West Indies and in southern
Florida, always in this habitat.
Botocudo cavernicola also does not show any obvious adaptations to cave
dwelling. It is in fact one of the more brightly colored species of Botocudo
with large eyes and ocelli. It will probably prove to have similar habits to
those of Cligenes subcavicola.
The carrying of small seeds on the end of the rostrum by Cligenes subca-
vicola is very reminiscent of the actions of species of Stilbocoris in Africa.
Carayon (1964) has given a fascinating account of the activity of members
of this genus. Here the male injects salivary secretion into the mature fallen
seeds of Ficus and offers the softened seed to the female who only then will
allow the male to copulate. Cligenes subcavicola (and probably also Botocudo
cavernicola) can be readily maintained in laboratory culture and it would
be valuable to know if the complex behavior found in species of Stilbocoris
is more widespread in the Rhyparochrominae.
ACKNOWLEDGMENTS
I wish to extend my sincere appreciation to the following: Mr. Noel Plumley (Bayswater,
Western Australia) and Dr. Philip Chapman (University of Bristol) for kindly providing in-
formation on the composition of the New Guinea cave and details of the collecting site of
Botocudo cavernicola, and to Dr. Randall Schuh (American Museum of Natural History) for
details of the collecting site near Tingo Maria, Peru. Mr. W. R. Dolling (British Museum Natural
History) and Dr. T. E. Woodward (University of Queensland) for allowing me to examine
specimens of Botocudo cavernicola. Dr. R. M. Baranowski (University of Florida) for infor-
mation on and aid in the collection of Cligenes subcavicola at Simla, Trinidad. Drs. C. W. and
L. B. O’Brien (Florida A.&M. University) for the gift of specimens. Ms. Mary Jane Spring and
Mrs. Elizabeth Slater (University of Connecticut) for the preparation of the illustration and
assistance with the manuscript respectively.
430
NEW YORK ENTOMOLOGICAL SOCIETY
LITERATURE CITED
Bergroth, E. 1918. Studies in Philippine Heteroptera I. Philippine Jour. Sci. 13(Sec. D):43-
126.
Carayon, J. 1964. Un cas d’offrande nuptiale chez les Heteropteres. Compt. Rendu Hbd.
Seances Acad. Sci. 259:4815-4818.
China, W. E. 1930. Insects of Samoa and other Samoan terrestrial Arthropoda Pt. II. He-
miptera. Ease. 3:81-162. London: British Museum (Natural History).
Scudder, G. G. E., J. P. E. C. Darlington and S. B. Hill. 1967. A new species of Lygaeidae
(Hemiptera) from the Tamana Caves, Trinidad. Ann. Speleologie 22 (Ease. 2):465-469.
Received August 17, 1983; accepted September 30, 1983.
NEW YORK ENTOMOLOGICAL SOCIETY
91(4), 1983, pp. 431-441
THE TYPES OF SOME AMERICAN CERCERIS WITH LECTOTYPE
DESIGNATIONS (HYMENOPTERA: PHILANTHIDAE)^
George R. Ferguson
Systematic Entomology Laboratory, Department of Entomology,
Oregon State University, Corvallis, Oregon 97331
Abstract.— primary types of 37 species-group taxa of American Cerceris are discussed,
and lectotypes are designated for 32 species-group taxa. Four previously designated neotypes
are shown to be invalid, and three holotypes are identified.
As part of a study of North American and Caribbean wasps of the genus
Cerceris the primary types of most of the described taxa have been studied.
Cresson (1916, 1928), Scullen (1965, 1972), and Ferguson (1983) designated
lectotypes for many species, most of which are located in type repositories
in the United States. Most of the lectotypes designated in this paper involve
species described by F. Smith, Saussure and Cameron and are located in
European institutions. In a few cases the specimens comprising a syntype
series are not conspecific; in other cases the specimens of a syntype series
are located in more than one institution. All lectotypes and holotypes dis-
cussed herein agree with the original descriptions unless otherwise noted.
The synonymy and taxonomic status of the species discussed below will be
treated in a forthcoming synonymic list. Abbreviations used in the text for
type repositories are explained in Acknowledgments.
Cerceris acolhua Saussure, 1867:90.
Described from at least two males from “Mexico. In planitie alta (terris
frigidis), . . .” and from “provincia Mechoacan”. A male in Geneva labelled
“Mexico t.f.” is here designated lectotype. A second conspecific syntype
male in Vienna is rather badly damaged.
Cerceris azteca Saussure, 1867:97.
Described from eight females from Cuantla and two females from Orizaba,
Mexico. A female in Vienna bearing a “Cuantla t.c.” label, a printed red
' Oregon Agricultural Experiment Station Technical Paper No. 6698.
The publication costs of this article were defrayed in part by page charge payment. This article
must therefore be hereby marked “Advertisement” in accordance with 18 U.S.C. §1734 solely
to indicate this fact.
432
NEW YORK ENTOMOLOGICAL SOCIETY
“Type” label, a handwritten ""Cerceris azteca Sauss.” label, and an ""azteca
Type det. Sauss.” label is here designated lectotype.
Cerceris bakeri Cameron, 1904:67.
Described from an unstated number of females from “Chinandega, Nic-
aragua”. A female in the BMNH labelled “Chinandega, Nicaragua, Coll.
Baker” is here designated lectotype. It also bears a circular red margined
“Type” label, a printed “Cameron Coll. 1904-313” label, a handwritten
“3055” label, a handwritten '"Cerceris bakeri Cam. Type Nicaragua” label,
and a printed “B. M. Type Hym. 21.1, 371” label. Scullen (1972) saw a
second female at Pomona College, but it has not been found in the USNM
where most of the Baker collection of Hymenoptera is now housed.
Cerceris binodis Spinola, 1841:1 17.
Described from two males from Cayenne, French Guiana. Two conspecific
syntype males are in the Spinola collection in Torino. These have been
standing under the header label "Philanthus binodis m., Didesmus Spinolae
Dlbm., D. Buquet, M. Le Prieur”. The specimens are otherwise unlabelled.
The larger of the two specimens is here designated lectotype.
Cerceris bothriophora Schletterer, 1887:456.
Described from an unstated number of both sexes from “Reg. V. Subreg.
3. (Mexico)”. One female and two males in Vienna bear handwritten "both-
riophora det. Schletterer” labels. These specimens agree with the original
description including the fact noted by Schletterer, that the fossette present
on tergum I of the female is absent in the males. The female also bears the
labels “Withm.” and “Newhaven” (handwritten and almost illegible). It is
a specimen of Cerceris atramontensis Banks, a common species in the area
of New Haven, Connecticut. Scullen (1972) designated and labelled this
specimen neotype, but this designation is rejected as invalid since it was not
shown that the original syntype series was lost or irretrievably damaged.
Although not from Mexico, the female agrees with the original description,
and it seems clear that Schletterer thought it was from a Mexican locality.
One male is labelled “Georg. Am. 1877 II” and is a specimen of Cerceris
compar Cresson. Although the specimen agrees with the rather incomplete
original description, the possibility exists that it is not a syntype because of
the locality label.
The second male has a handwritten “Aug.” label and a printed “Bilimek,
Mexico, 1871” label with an illegible handwritten word below the printing.
This specimen also agrees with the original description and is here designated
lectotype, since the label agrees with the type locality given in the original
description.
Cerceris chinandegaensis Cameron, 1904:66.
Described from an unstated number of males from Chinandega, Nicara-
gua. Scullen (1972) saw four males: one each at BMNH and Cornell, and
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433
two at Pomona College. I have seen three conspecific males: one each at the
BMNH, Cornell, and the Baker collection now at the USNM. The second
male from Pomona College reported by Scullen has not been located. The
three specimens examined are labelled “Chinandega, Nicaragua, Coll. Bak-
er”. The BMNH specimen bearing a circular, red margined “Type” label, a
printed “Cameron Coll. 1904-313” label, a handwritten “3054” label, a
handwritten '"Cerceris chinandegaensis Cam. Type Nicaragua” label, and a
printed “B. M. Type Hym. 21.1, 374” label is here designated lectotype.
The USNM specimen bears a '"Cerceris kennicottii baked Cameron det.
H. A. Scullen” label. The Cornell specimen bears a printed “Cornell U. Lot
546 Sub 207 Baker Coll.” label, and a ""Cerceds kennicottii Cresson det. H.
A. Scullen” label.
Cerceds clypeata Dahlbom, 1844:221.
Described from both sexes from “Carolina meridionali . . . Dr. Zimmer-
man”. Number of specimens not stated. A female and male in Lund are not
conspecific. The female bears a handwritten “105” label, a “E. Sudcarolina
Zimmerman” label, an unmarked red label, and a blue “1982-402” label.
It is here designated lectotype. The male bears a handwritten “104” label,
a “S. Carolina” label, a “C^r. clypeata Kl. Mus. Berol.” label, and a blue
“1982-403” label. It is a male of the homeochromic Cerceds halone Banks.
Cerceris cribrosa Spinola, 1841:119.
Described from two females and one male from Cayenne, French Guiana.
I have studied a syntype female and a conspecific male syntype from the
Spinola collection in Torino. Scullen (1 972) designated the female as neotype,
but this designation is invalid since a syntype cannot become a neotype. I
here designate the same female as lectotype. The male has the gaster missing
beyond the first segment.
Cerceris crotonella Viereck and Cockerell, 1904:139.
Described from one female from Las Cruces, New Mexico. Cresson (1928)
does not list this species. A female in the type collection of the ANSP from
Las Cruces, N. Mex. bears a handwritten ""Cerceris crotonella V. & C.” label
and is accepted as the holotype. Cresson’s ledger in the ANSP has a line
drawn through his entry No. 10039, C. crotonella with the notation “off in
coll. Ckll”. The number 10039 is not on the specimen. Apparently the
specimen was subsequently returned to the ANSP collection.
Cerceris crucis Viereck and Cockerell, 1904:139.
Described as Cerceris rufinoda var. crucis from two females from Las
Cruces, New Mexico. This taxon is not listed by Cresson (1928). The spec-
imen in the type collection at ANSP bears a “Las Cruces, N. M.” label and
a handwritten ""Cerceris rufinoda var. crucis V. & C. Type” label, and it is
here designated lectotype. The second specimen has not been located. Scullen
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NEW YORK ENTOMOLOGICAL SOCIETY
(1965) was incorrect in stating that the type of crucis carried the ANSP No.
10393 as this number applies to quite a different wasp (see Cresson, 1928,
p. 42). The lectotype does not bear an ANSP type number.
Cerceris deserta Say, 1825:343.
Scullen’s (1965, p. 479) statement designating a neotype for this species
is not clear as to whether he or Banks made the designation. Neither the
labels on the specimen nor Banks’ (1912) only published reference to deserta
give any indication that he designated a neotype. The specimen [MCZ] carries
a neotype label affixed by Scullen, although it also bears a Cerceri^ deserta
Say determination label apparently in Banks’ handwriting. The neotype
designation is clearly that of Scullen (1965).
Cerceris dilatata Spinola, 1841:118.
Described from two females and one male from Cayenne, French Guiana.
I have examined a syntype female and a conspecific syntype male from the
Spinola collection in Torino. Scullen (1972) designated the male as neotype,
but this designation is invalid since a syntype cannot become a neotype.
The female is here designated lectotype, since it is the more distinctive of
the two sexes.
Cerceris elegans F. Smith, 1856:467.
Described from an unstated number of males from “Hab. East Florida
(eoll. E. Doubleday, Esq.).” Two conspecific males in the BMNH are each
labelled “E. Doubleday, St. John’s Bluff, E. Florida.” One male bearing a
circular orange margined “Type” label, a handwritten "'elegans Sm. Type”
label, and a printed “B. M. Type Flym. 21.1 365” label is here designated
lectotype. The second male has the gaster missing beyond the first segment.
Cerceris exsecta F. Smith, 1873b:410.
Described from an unstated number of both sexes from “Hab. Mexico.”
I have examined one female syntype and three conspecific males from the
BMNH. The female bearing a small circular “Mex.” label, a handwritten
"Cerceris exsecta Smith” label, a circular orange margined “Type H. T.”
label, and a “B. M. Type Hym. 21.1, 429” label is here designated lectotype.
One male labelled “Mex” is probably a syntype. A second male bears a “F.
Smith Coll.” label, and the third male has a “Smith Coll. pres, by Mrs.
Farren White, 99-303” label. Both carry a “Mexico” locality label, but it is
questionable whether or not they are syntypes.
Cerceris feralis Cameron, 1890:1 13.
Described from an unstated number of males from “Hab. Mexico (coll.
Saussure)”. I have examined six conspecific males from the BMNH of which
two are probably not syntypes. A male bearing a “Cuantla t.c.” label, a
circular orange margined “Type H. T.” label, a handwritten "Cerceris feralis
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435
Cam. Type B.C.A. ii, 113” label, and a printed “B. M. Type Hym. 21.1
372” label is here designated lectotype. Two males labelled “Cuantla t.c.”
and one labelled “Mexiq. Orizaba” each bears a “P. Cameron Coll. 1914-
1 10” label. The italicized type face of the locality labels indicates that they
are from the Saussure collection; these are accepted as syntypes. Two males
each bearing a “N. Yucatan Gaumer” label and a “Cameron Coll. B. M.
1914-110” label are not from the Saussure collection and are probably not
syntypes. The latter two specimens each bears a ^'Cerceris compar geniculata
Cam. det. H. A. Scullen” label.
Cerceris flavida Cameron, 1890:116.
Described from an unstated number of both sexes from “Hab. Mexico,
Cordova (Coll. Saussure).” A female in the BMNH bearing a “Cordova t.c.”
locality label, a circular blue margined “Syntype” label, a handwritten ""Cer-
ceris placida [sic] Cam” label, and a “Syntype Cerceris flavida Cam. det C.
R. Vardy” label is here designated lectotype of Cerceris flavida Cameron.
Despite the ""placida" label the specimen has the aberrant wing venation
described by Cameron in the original description. The second cubital cross
vein is missing on the left front wing but represented by a stub on the right
front wing. The first cubital cross vein is partially missing on the right front
wing.
Cerceris flavomaculata Cameron, 1890:1 15.
Described from an unstated number of both sexes from “Hab. Costa Rica,
Rio Su/^io, Cach^(Rogers).” A female and a male syntype in the BMNH are
not conspecific. The female bearing a circular blue margined “Syntype” label,
a printed “R. Susio, H. Rogers” label, and a printed “B.C.A. Hymen. II
Cerceris flavomaculata Cam.” label is here designated lectotype. The male
bears a circular blue margined “Syntype” label, a circular orange margined
“Type” label, a printed “Cach^ Costa Rica, H. Rogers” label, a printed “B.
C. A. Hymen. II, Cerceris flavomaculata Cam.” label, a handwritten ""Cer-
ceris flavomaculata Cameron, Type, B. C. A. ii, 115” label, and a printed
“B. M. Type Hym. 21.1, 376” label. The specimen is without a gaster and
very dirty. The portions of the specimen that can be clearly seen indicate
that it is a male of Cerceris cooperi Scullen.
Cerceris geniculata Cameron, 1890:1 13.
Described from an unstated number of females from “Hab. Mexico, Cuan-
tla (Coll. Saussure).” A female in the BMNH bearing a “Cuantla, t.c.” locality
label, a “P. Cameron Coll. 1914-110” label, and a handwritten ""Cerceris
geniculata Cam.” label is here designated lectotypes. A second conspecific
female bears a “Mexiq. Orizaba” label, a “P. Cameron Coll. 1914-110”
label, a red margined circular “Type H. T.” label, a “B. M. Type Hym. 21.1,
366” label, and a handwritten ""Cerceris geniculata Cam. Type B. C. A. ii.
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NEW YORK ENTOMOLOGICAL SOCIETY
1 13” label. The Orizaba locality was not mentioned in the original descrip-
tion.
Cerceris hebes Cdivcitr on, 1890:124.
Described from the male from “Hab. Mexico, Chilpancingo in Guerrero
4500 feet (H. H. Smith)”. Number of specimens not stated. A male in the
BMNH bears a “Valladolid, Yucatan, Gaumer” label, a printed “B. C. A.
Hymen. II Cerceris hebes Cam. Type” label, and a handwritten “(? locality
G.C.C.)” label. A head glued to the thorax is that of Eucerceris montana
Cresson {= Cerceris sonorensis Cameron) and is not conspecific with the
remainder of the specimen. The pygidium agrees with the figure accompa-
nying the original description and is sufficient to identify the thorax and
gaster as that of a male of Cerceris graphica F. Smith. The thorax and gaster
is accepted as the holotype despite the discrepancy in the locality label of
the specimen versus the locality stated in the original description. The lo-
cality label on the specimen was “questioned by G. C. C.” Nevertheless the
species is identifiable from the original description and accompanying fig-
ures.
Cerceris intricata F. Smith, 1856:459.
Described from an unstated number of males from “Hab. Brazil (Santa-
rem). (Coll. H. W. Bates)”. A male in the BMNH bearing a blue circular
handwritten “Braz. Santarem” label, a circular orange margined “Type”
label, a handwritten "'intricata Sm. Type” label, a printed “B. M. Type Hym.
21.1, 407” label, and a “Syntype. Another at Oxford prob. conspecific. det.
C. R. Vardy 1976” label is here designated lectotype. A second conspecific
male in the BMNH bears a “Para” label, a blue handwritten "intricata
Smith” label, and a “Smith coll. pres, by Mrs. Farren White 99-303” label,
but it is probably not a syntype.
Cerceris krugi Dewitz, 1881:200.
Described from five specimens of both sexes from Puerto Rico. One female
and two male syntypes, all conspecific, from Berlin have been studied. The
female bearing a green “Portorico Krug” label; a red, printed “TYPE” label;
a white printed “2204 1 ” label; and a green, handwritten "krugii [sic] Dewitz”
label is here designated lectotype.
One male has a green “Portorico Moritz” label; a red, printed “TYPE”
label; a white “4439” label; and a green "krugii [sic] Dewitz” label. The
second male is similarly labelled, and, in addition, has a large, green, hand-
written "krugi Dewitz, Berliner ent. Zeit. XXV” label. The latter specimen
has the head and tergum VII badly damaged. The location of the remaining
two syntypes is not known.
Cerceris laevigata F. Smith, 1856:465.
Described from both sexes from Santo Domingo. Number of specimens
not stated. A syntype female and conspecific syntype male in the BMNH
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437
are labelled “St. Dom. 55.1”. The female bearing a circular orange margined
“Type” label, a handwritten ""laevigata Sm. Type” label, and a printed “B.
M. Type Hym. 21.1, 377” label is here designated lectotype. The left hind
leg is glued to a point. The male bears only the locality label.
Cerceris montezuma Cameron, 1890:108.
Described from an unstated number of females from “Hab. Mexico (coll.
Saussure)”. There are four specimens in the BMNH under this name of
which two are conspecific females and agree with the original description.
The female bearing a handwritten “Mex. Mayo [and an illegible word]”
label, a “P. Cameron Coll. 1914-110” label, and a handwritten ""Cerceris
montezuma Cam.” label is here designated lectotype. A second conspecific
female has a handwritten “Mex.” label, a printed “F. Sm. Coll. 79.22” label,
and a handwritten ""montezuma'’" label. Since the F. Smith collection was
not mentioned in the original description, the lectotype may in fact be a
holotype. However, the F. Smith collection was available to Cameron, and
it seems prudent to make the lectotype designation.
A third female bears a “Omilteme, Guerrero, 8000 ft. Aug. H. H. Smith”
label, a “P. Cameron Coll. 1914-110” label, a “in B. M. 1973 under C.
graphica"" label, and a handwritten ""montezuma"" label. It does not agree
with the original description, but it is a specimen of Cerceris dreisbachi
Scullen.
The fourth specimen consists of a thorax and part of the gaster on a pin
with a head and the remaining part of the gaster glued to the second label.
It is a male of which the head appears to be that of Cerceris californica
Cresson and the remaining parts appear to be Cerceris erythropoda Cameron.
It bears a “Presidio, Mexico, Forrer” label, a printed “B. C. A. Hymen. II,
Cerceris montezuma Cam.” label, a handwritten “probably wrongly named
GCC” label, and a handwritten ""Cerceris montezuma Cam. Type, B. C. A.
iii, 108” label. The composite specimen bears no resemblance to the original
description.
Cerceris nigra Ashmead, 1900:227.
Described from two females from “St. Vincent- Wind ward side” [B. W.
I.]. A female in the USNM bears a printed “Windward side St. Vincent, W.
I., H. H. Smith” label, a red “Type No. 6410 USNM” label, and a hand-
written ""Cerceris nigra Ashm. Type” label. It is here designated lectotype.
A second female in the BMNH also carries a handwritten ""Cerceris nigra
Ashm. Type” label. I have not seen it, but according to comments received
from C. R. Vardy (pers. comm.) it is conspecific.
Cerceris obsoleta Cameron, 1890:1 18.
Described from an unstated number of females from “Hab. Mexico, Teapa
in Tabasco (H. H. Smith)”. Two conspecific females in the BMNH each
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NEW YORK ENTOMOLOGICAL SOCIETY
bears a Teapa, Tabasco, locality label. One bearing a “Teapa, Tabasco,
March H. H. S.” label, and a printed “B. C. A. Hymen. II Cerceris obsoleta
Cam.” label is here designated lectotype. The second specimen is very dirty
and bears a “Teapa, Tabasco, Feb. H. H. S.” label, a circular orange margined
“Type” label, and a handwritten ""Cerceris obsoleta Cam. Type B. C. A. ii
118” label.
Cerceris otomia Saussure, 1867:99.
Described from five males from “. . . urbis Mexico” and “Zinapecuaro in
provincia Mechoacan”, and three females and six males from “. . . Cordillera
oriental! . ...” A female in Vienna bearing a “Mexico, Saussure, Type 1880”
label, a handwritten ""otomia Type det. Sauss.” label, and a printed red
“Type” label is here designated lectotype.
Cerceris pullata F. Smith, 1873a: 105.
Described from two females from “Hab. St. Paulo”. Two conspecific
syntype females in the BMNH each bears a blue circular “St. Paul” label.
One of these bearing an orange margined circular “Type” label, a handwritten
""Cerceris puUatus [sic] Smith” label, and a printed B. M. Type Hym. 21.1,
409” label is here designated lectotype. The second female bears only the
locality label. A third conspecific female bears a handwritten ""Cerceris pul-
latus Smith St. Paulo” label and a printed “Smith coll. pres, by Mrs. Farren
White 99-303” label, but it is probably not a syntype.
Cerceris rufopicta F. Smith, 1856:467.
Described from an unstated number of males from “Hab. E. Florida”.
Two male syntypes in the BMNH are conspecific. One specimen bearing a
circular orange margined “Type H. T.” label, a circular white label with
illegible markings on both sides, a printed “R. Foster, St. John’s Bluff, E.
Florida” label, a handwritten ""rufopicta Sm. Type” label, a circular blue
margined “Syntype” label, and a “B. M. Type Hym. 21.1, 364” label is here
designated lectotype. The second male bears a small handwritten “’71” label,
a printed “E. Doubleday, St. John’s Bluff, E. Florida” label, a handwritten
""rufopicta Smith” label, a printed “Smith Coll. pres, by Mrs. Farren White,
99-303” label, and a circular blue margined “Syntype” label.
Cerceris simplex ¥. Smith, 1856:462.
Described from an unstated number of both sexes from “Hab. Brazil
(Santarem), (Coll. H. W. Bates)”. A female in the BMNH bearing a blue
circular handwritten “Santarem” label, a circular orange margined “Type”
label, a handwritten label with illegible marks, a handwritten ""simplex Sm.
Type” label, and a printed “B. M. Type Hym. 21.1, 438” label is here
designated lectotype. A male syntype has not been located.
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439
Cerceris simulans Saussure, 1867:87.
Described from a male from “Mexico Temperata: in Cordillera oriental!”.
The type has not been located. A headless female in Geneva was designated
neotype by Scullen (1972), but this designation is invalid since the original
description was based on a male and the headless female disagrees with the
original description in several important respects. Furthermore, C simulans
Saussure is readily identifiable from Saussure’s original description and fig-
ures. The headless female is identifiable as Cerceris scapularis Schletterer
which belongs to quite a different species group from C. simulans.
Cerceris smithiana Cameron, 1890:1 19.
Described from an unstated number of females from “Hab. Mexico, Atoy-
ac in Veracruz (H. H. Smith)”. Two conspecific females in the BMNH each
bears an “Atoyac, Veracruz, May. H. H. S.” label and a printed “B. C. A.
Hymen. II Cerceris smithiana Cam.” label. One of these is here designated
lectotype. The second specimen also bears a circular orange margined “Type”
label, a handwritten '^Cerceris smithiana Cam. Type B. C. A. ii 1 19” label,
and a printed “B. M. Type Hym. 21.1, 369” label, but it is very dirty.
Cerceris strigosa Cameron, 1890:1 10.
Described from an unstated number of both sexes from “Hab. Mexico,
Ciudad in Durango 8100 feet (Forrer)”. Two conspecific females in the
BMNH each bears a “Ciudad Mex., 8100 ft., Forrer” label, a printed “B.
C. A. Hymen. II Cerceris strigosa Cam.” label, and a handwritten Cerceris
strigosa Cam.” label. One of these also bears a circular orange margined
“Type” label and a notation on the handwritten label “Type B. C. A. ii
1 10”, and it is here designated lectotype. It is also the cleanest of the two
specimens.
Cerceris subpetiolata Saussure, 1867:95.
Described from “Mexicanus ager. Terrae calidae”, “In Puebla viejo apud
Tampico” two males, and “Cordillera orientalis” one female. A female in
Vienna bearing a “Saussure type 880” label, a “Cordova t.c.” label, a ^"sub-
petiolata type det. Sauss.” label, and a red printed “Type” label is here
designated lectotype. A conspecific syntype male bears a “Cordova t.c.”
label, a “Saussure type 880” label, and a ""subpetiolata det. Sauss.” label.
Cerceris townsendi Viereck and Cockerell, 1904:140.
Described from a male from Las Cruces, New Mexico. The species is not
listed in Cresson (1928). A male in the type collection of the ANSP bears a
“Las Cruces, N. M.” label, a red “Type No. 10389” label, and a handwritten
""Cerceris acanthophiloides V. & C.” label. Cresson’s ledger at the ANSP
contains the entry “C. xanthophiloides [sic] V. & C.” under his number
10389. The specimen agrees in all details of structure and coloration with
the original description of C. townsendi, and I have no doubt that this
440
NEW YORK ENTOMOLOGICAL SOCIETY
specimen is the holotype. Apparently Viereck and Cockerell changed the
name in publication and neglected to change the label on the specimen. I
have affixed a label to the specimen indicating that I consider it to be the
holotype of Cerceris townsendi.
Cerceris truncata Cameron, 1 890: 1 2 1 .
Described from an unstated number of both sexes from “Hab. Mexico,
Temax in North Yucatan (Gaumer)”. I have seen six females and four males
from the BMNH. All are conspecific, and all are apparently syntypes. One
female and two males each bears a “Temax, N. Yucatan, Gaumer” label;
and five females and two males each bears a “N. Yucatan, Gaumer” label.
Each of the ten specimens bears a printed “B. C. A. Hymen. II Cerceris
truncata Cam.” label. The female labelled “Temax, N. Yucatan, Gaumer”
is here designated lectotype since the locality label agrees precisely with the
original description. One female labelled “N. Yucatan, Gaumer” also bears
a circular orange margined “Type” label, a “B. M. Type Hym. 21.1, 433”
label, and a handwritten ""Cerceris truncata Cam. Type” label, but it is in
rather poor condition.
Cerceris vulpina F. Smith, 1856:463.
Described from an unstated number of males from “Hab. Brazil”. A male
in the BMNH bearing a blue circular “Santarem” label, a circular orange
margined “Type” label, a handwritten ""vulpina Sm. Type” label, and a
printed “B. M. Type Hym. 21.1, 418” label is here designated lectotype. A
second conspecific male bears a circular white handwritten “Braz.” label, a
handwritten “C vulpina Sm.” label, and a printed “Smith coll. pres, by Mrs.
Farren White 99-303” label, but it may not be a syntype.
Cerceris zapoteca Saussure, 1867:89.
Described from two males from “prope Tampico”. A male in Vienna
bearing a “Cordova t.c.” label, a handwritten “Saussure Type 1880” label,
a handwritten ""zapotecus [sic] Type det. Sauss.” label, a handwritten ""Cer-
ceris zapotecus [sic] Sss.” label, and a red printed “Type” label is here
designated lectotype. A second conspecific male in the BMNH with the same
locality label is the presumed second syntype.
ACKNOWLEDGMENTS
I am indeed grateful to the following individuals and institutions for their generous loan of
the type material discussed in this paper [abbreviations as used in the text are in brackets]: D.
Azuma, Academy of Natural Sciences of Philadelphia [ANSPj; F. Koch, Zoologisches Museum,
Humboldt Universitat, Berlin, DDR [Berlin]; C. R. Vardy, British Museum (Natural History),
London [BMNH]; Q. D. Wheeler, Cornell University, Ithaca, New York [Cornell]; Cl. Besuchet,
Museum d’Histoire Naturelle, Geneva [Geneva]; R. Danielsson, Museum of Zoology and Ento-
mology, Lund University, Lund, Sweden [Lund]; K. Jepson, Museum of Comparative Zoology,
Harvard University, Cambridge, Massachusetts [MCZ]; P. Passerin d’Entreves, Museo ed Is-
VOLUME 91, NUMBER 4
441
tituto di Zoologia Sistematica, Universita di Torino, Torino, Italy [Torino]; A. S. Menke, USDA
Systematic Entomology Laboratory, % U.S. National Museum of Natural History, Washington
[USNM]; M. Fischer, Naturhistorisches Museum Wien, Vienna [Vienna].
I also thank A. S. Menke, P. W. Oman, Oregon State University, and C. R. Vardy for helpful
comments regarding some of the problems encountered.
LITERATURE CITED
Ashmead, W. H. 1900. Report upon the aculeate Hymenoptera of the islands of St. Vincent
and Grenada .... Trans. Entomol. Soc. London, part II, pp. 207-367.
Banks, N. 1912. Notes on the eastern species of Cerceris. Ann. Entomol. Soc. Am. 5:1 1-30.
Cameron,?. 1888-1891. Insecta, Hymenoptera, v. 2 (Fossores). xi + 413 pp. 7«. F. D. Godman
and D. Salvin, Biologia Central!- Americana. Taylor and Francis, London. (Pp. 1-32 =
1888; pp. 33-64 = 1889; pp. 65-128 = 1890; pp. 129-176 = 1891.)
Cameron, P. 1904. New Hymenoptera mostly from Nicaragua. Invertebrata Pacifica 1:46-
69.
Cresson, E. T. 1916. The Cresson types of Hymenoptera. Mem. Am. Entomol. Soc. No. 1,
pp. 1-141.
Cresson, E. T. 1928. The types of Hymenoptera in the Academy of Natural Sciences of
Philadelphia other than those of Ezra T. Cresson. Mem. Am. Entomol. Soc. No. 5, pp.
1-90. (Posthumously published by E. T. Cresson, Jr.)
Dahlbom, A. G. 1843-1845. Hymenoptera Europaea praecipue borealis . . . . v. 1, xliv -H 528
pp. Lundbergiana, Lund. (Pp. 1-172, fasc. 1 = 1843; pp. 173-352, fasc. 2 = 1844; pp.
353-528, i-xliv, tables = 1845.)
Dewitz, H. 1881. Hymenopteren von Portorico. Berliner Entomol. Z. 25:197-208.
Ferguson, G. R. 1983. The Banks types of cercerine wasps. J. New York Entomol. Soc. 91:
223-234.
Saussure, H. de. 1867. Hymenoptera. Pages 1-138, in Reise der dsterreichischen Frigatte
Novara . . . , Zoologischer Theil, v. 2, 156 pp. Wien.
Say, T. 1825. In Keating, Narrative of an Expedition to the Source of St. Peter’s River . . .
Under the Command of S. H. Long, v. 2, pp. 268-378.
Schletterer, A. 1887. Die Hymenopteren gattung Cerceris Latr. mit Vorzugsweiser Beriick-
sichtigung der Palaarktischen Arten. Zool. Jahrb. 2:349-510.
Scullen, H. A. 1965. Review of the genus Cerceris in America north of Mexico. Proc. U.S.
Natl. Mus. 116:333-548.
Scullen, H. A. 1972. Review of the genus Cerceris Latreille in Mexico and central America.
Smithsonian Contr. Zool. 110:1-121.
Smith, F. 1856. Catalogue of hymenopterous insects in the collection of the British Museum,
part IV, Sphecidae, Larridae, and Crabronidae. London, pp. 207-497.
Smith, F. 1873a. Descriptions of new species of fossorial Hymenoptera in the collection of
the British Museum. Ann. Mag. Nat. Hist. (4) 11:441-451; 12:49-59, 99-108.
Smith, F. 1873b. Descriptions of new species of fossorial Hymenoptera in the British Museum
and of a species of the rare genus Iswara belonging to the family Dorylidae. Ann. Mag.
Nat. Hist. (4) 12:253-260, 291-300, 402-415.
Spinola, M. 1841. Hymenopteres, receuillis a Cayenne en 1839 per M. Leprieur, pharmacien
de la Marine Royale. Ann. Soc. Entomol. France 10:85-149.
Viereck, H. L. and T. D. A. Cockerell. 1904. The Philanthidae of New Mexico, II. J. New
York Entomol. Soc. 12:129-146.
Received May 24, 1983; accepted July 27, 1983.
NEW YORK ENTOMOLOGICAL SOCIETY
91(4), 1983, pp. 442-465
TYPE DESIGNATIONS AND NEW SYNONYMIES FOR NEARCTIC
SPECIES OF PHYTOCORIS FALLEN (HEMIPTERA: MIRIDAE)
Thomas J. Henry and Gary M. Stonedahl
Systematic Entomology Laboratory, IIBIII,
Agriculture Research Service, USD A, % National Museum of
Natural History, Washington, D.C. 20560 and
Department of Entomology, Oregon State University,
Corvallis, Oregon 97331
Abstract.— TYiiny-stvcn lectotype and two neotype designations are made for Nearctic species
of Phytocoris (Hemiptera: Miridae) described by O. M. Reuter, P. R. Uhler, and E. P. Van
Duzee. One holotype is identified, and eight species are considered as new synonyms. Species
are listed alphabetically; and for each, year of publication, original page number, exact label
data, type depository, pertinent body measurements, and discovered paralectotypes are given.
Comments are provided on distributions, hosts, identification keys, and male genitalia.
Prior to this study, 230 species of the plant bug genus Phytocoris were
recognized from Canada and the United States. Most of these were described
by H. H. Knight, O. M. Reuter, P. R. Uhler, and E. P. Van Duzee. Van
Duzee (except for species treated in this paper) and Knight regularly des-
ignated holotypes for their species, thus satisfying Article 73(a) and Rec-
ommendation 73 A of the 1961 International Code of Zoological Nomen-
clature. However, because of the lack of holotype designations and the
frequency of mixed syntype series of species described by Reuter and Uhler
and some by Van Duzee, the remaining species of Phytocoris are often
difficult to identify with certainty. We have found a number of specimens
bearing labels such as “typus,” “lectotype,” “allotype,” and “paratype,”
indicating that someone at the respective museums recognized that these
specimens belonged to the original type series of the species; these desig-
nations, however, as far as we can determine, represent in-house curation
and have never been validated through publication. To insure nomencla-
torial stability in the North American species of Phytocoris, we feel it is
important to make proper type designations, or in a few cases select necessary
neotypes.
We have located most of the original specimens of Reuter and Van Duzee
in the California Academy of Sciences and in several European museums.
The publication costs of this article were defrayed in part by page charge payment. This article
must therefore be hereby marked “Advertisement” in accordance with 18 U.S.C. §1734 solely
to indicate this fact.
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443
The remaining Phytocoris types, including most of Uhler’s, were discovered
in the U.S. National Museum of Natural History. We have not found types
for interspersus Uhler and minutulus Reuter.
In this paper we select 37 lectotypes, identify one holotype, and assign
two neotypes for the North American Phytocoris described by Reuter, Uhler,
and Van Duzee. Eight species names are recognized as new synonyms. When
syntypes were located in the U.S. National Collection and also in European
collections, we have selected the best specimen as the lectotype and have
labeled the remaining syntypes as paralectotypes. If possible, we have chosen
males as lectotypes, or if males are in poor condition, a female was selected.
We recognize a holotype only when the author stated he had a single spec-
imen; in all other cases, even when we could locate only one specimen, we
assumed that there was a series. The lectotypes, neotypes, and holotype are
identified with red rectangular labels having the appropriate type printed at
the top; paralectotypes are identified with blue rectangular labels.
The list of taxa below is arranged alphabetically by species with year of
publication and original page number given after the author. In the same
paragraph, exact label data, set in quotes, are given for each primary type,
followed by type depository, condition of primary type, measurements of
major body structures of primary type, and list of secondary types located.
Synonyms are listed in a new paragraph with the same data as for the valid
name; species listed without this data (e.g., those described by H. H. Knight)
indicate that a holotype was properly designated and accompanied by a
description and measurements. A remarks section under each taxon includes
such information as notes on synonymy, confused identities, sources of keys,
and other pertinent comments. All junior synonyms are listed alphabetically
in the text and cross referenced to the valid senior synonym; e.g., Phytocoris
bipunctatus Van Duzee [See annulicornis Reuter].
The following measurements, in millimeters, are given for each primary
type: Body length = BE (from apex of tylus to apex of wing membrane);
body width = BW (greatest width across hemelytra, usually just above cu-
neus); head width = HW (width across eyes); vertex width = V (greatest
distance between eyes); rostrum length = RL; length of antennal segments =
AI, All, AIII, AIV; pronotal length = PL (mesal length); pronotal width =
PW (basal or posterior width).
The following abbreviations are used for institutions serving as type depos-
itories of the Phytocoris species: CAS (California Academy of Sciences, San
Francisco, California); TMB (Termeszettudmanyi Muzeum, Budapest);
LACM (Los Angeles County Museum, Los Angeles, California); MCZ (Mu-
seum of Comparative Zoology, Harvard University, Cambridge, Massachu-
setts); NRS (Naturhistoriska Riksmuseet, Stockholm); USNM (United States
National Museum of Natural History, Washington, D.C.), and UZMH
(Universitetets Zoologiska Museem, Helsinki).
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NEW YORK ENTOMOLOGICAL SOCIETY
Phytocoris arnericanus Carvalho
Phytocoris angustulus Reuter, 1909:29 [name preoccupied by Phytocoris
angustulus Germar and Berendt, 1856 (fossil mirid)]. Type data (Fig. 1).—
Lectotype male (here designated): Label 1, ‘‘F[or]t. Pendl[e]t[o]n, 10-7,
W[es]t V[irgini]a”; 2, “O. Heidemann Collector”; 3, ""Phytocoris angus-
tulus n. sp. [handwritten], O. M. Reuter det.”; 4 (here added), “Lectotype:
Phytocoris angustulus Reuter, by Henry and Stonedahl, 1983” [USNM
type No. 100402; condition good, except the antennae and one fore- and
one hindleg are missing]. Measurements of lectotype: BL 5.92; BW 1.92;
HW 0.92; V 0.32; RL 2.52; antennae missing; PL 0.92; PW 1.60. Para-
lectotypes: No other syntypes located.
Phytocoris arnericanus Carvalho, 1959:190 [new name for Phytocoris an-
gustulus Reuter].
Remarks. Knight (1923) correctly recognized this species (as angustulus),
figured male genitalia, and gave eastern hemlock, Tsuga canadensis (L.)
Carr., as the host. Knight (1941) recorded arnericanus (as angustulus) from
New York, North Carolina, Nova Scotia, Vermont, and West Virginia;
Wheeler and Henry (1977) added Pennsylvania.
Phytocoris angustulus Reuter
[See arnericanus Carvalho]
Phytocoris annulicornis (Reuter)
Cornpsocerocoris annulicornis Reuter, 1 876:70. Type data. — Lectotype male
(here designated): Label 1, “Texas”; 2, “Belfrage”; 3, “Paratypus”; 4, “307
82”; 5, “Riksmuseum Stockholm”; 6 (here added), “Lectotype: Comp-
socerocoris annulicornis Reuter, by Henry and Stonedahl, 1983” [NRS;
condition good, except most of left wing membrane and the 4th antennal
segments are missing]. Measurements of lectotype: BL 4.58; BW ca. 1.28
(wings spread); HW 0.82; V 0.32; RL 2.08; AI 0.96; All 2.04; AIII ca.
1.40 (slightly curled); AIV missing; PL 0.64; PW 0.72. Paralectotypes: 2
males and 1 female, same data as for lectotype [NRS].
Phytocoris annulicornis: Reuter, 1909:33 (new combination).
Phytocoris bipunctatus Van Duzee, 1910:77. NEW SYNONYMY. Type
data. — Lectotype male (here designated): Label 1, “7-Oaks, Fla., May 1,
’08, Van Duzee”; 2, “Lectotype bipunctatus''' (in red, handwritten); 3, “EP
Van Duzee Collection”; 4 (here added), “Lectotype: Phytocoris bipunctatus
Van Duzee, by Henry and Stonedahl, 1983” [CAS type No. 1995; con-
dition fair, left antenna, right foreleg, and right hindleg missing; antennal
segment III on right side broken, segment IV missing; membrane of right
hemelytron chipped; anterolateral margin of left clavus chipped]. Mea-
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445
surements of lectotype: BL 5.08; BW 1.44; HW 0.87; V 0.33; RL 2.05;
AI 1.04; All 2.21; AIII broken or missing; AIV missing; PL 0.72; PW
1.19. Paralectotypes: 1 male and 1 female, same data as for lectotype; 1
male, Sanford, Fla., Apr. 27, ’08, Van Duzee; 1 female, St. Petersbg., Fla.,
4-28-08, Van Duzee; 1 female, Tampa, Fla., May 2, ’08, Van Duzee; 1
female. Ft. Myers, Fla., May 3-5, ’08, Van Duzee. All paralectotypes are
deposited in the collection of the CAS.
Remarks. Phytocoris annulicornis is recorded from Maine, south to Flor-
ida, west to California, and south to Panama (Carvalho, 1959). We suspect
that the far western records (California, Colorado, and New Mexico) and
the far southern records (Guatemala and Panama) are in error. Although
Carvalho (1959) indicated that Knight (p. 717) in Blatchley (1926) synon-
ymized bipunctatus under annulicornis, Blatchley did not agree with this
opinion, noting that neither Reuter (1876) nor Distant (1883) mention the
prominent pale spots on the corium which are distinctive in bipunctatus.
Knight (1927b) added to Blatchley’s discussion of annulicornis, stating that
he had examined a co-type of annulicornis and that this specimen was the
same as bipunctatus, but felt the final decision on synonymy should come
only after all of Reuter’s specimens were examined. We have examined the
types of both species and can say, without a doubt, that bipunctatus is a
junior synonym of annulicornis.
Blatchley (1926) redescribed and keyed annulicornis. Contrary to his key,
annulicornis (couplet bb) has two white spots at the apex of the corium. His
key should be modified accordingly.
Phytocoris antennalis Reuter
Phytocoris antennalis Reuter, 1909:32. Type data. — Lectotype male (here
designated): Label 1, “Washington, D. C., 24-IX-04”; 2, “O. Heidemann
Collector”; 3, ""Phytocoris antennalis n. sp. [handwritten] O.M. Reuter
det.”; 4, “Mus. Zool. H:fors Spec. typ. No. 9685 Phytocoris antennalis
O.M. Reut.”; 5 (here added), “Lectotype: Phytocoris antennalis Reuter,
by Henry and Stonedahl, 1983” [UZMH; good condition except the 4th
antennal segments are missing and the wings are spread (pinned through
right anterior region of abdomen). Measurements of lectotype: BL 6.10;
BW not measured, wings spread; HW 0.90; V 0.44; RL 2.81; AI 1.60; All
3.28; AIII 2.16; AIV missing; PL 0.79; PW 1.40. Paralectotypes: 4 females,
same locality as for lectotype with dates 15-9, 4-7-1889, 4-7-1901, 19-
VIII- 1901, [3 females, UZMH; 1 female, USNMj.
Remarks. Phytocoris antennalis has been redescribed and keyed by Knight
(1923) and Blatchley (1926). This is a widespread species that occurs from
Massachusetts, south to Florida, and west to Oklahoma (Carvalho, 1959).
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NEW YORK ENTOMOLOGICAL SOCIETY
Phytocoris barbatus Van Duzee
[See roseus (Uhler)]
Phytocoris bakeri Reuter
Phytocoris bakeri Reuter, 1909:28. Type data. — Lectotype male (here des-
ignated): Label 1, “Claremont, Cal., Baker”; 2, “1203”; 3, ""Phytocoris
bakeri n. sp. [handwritten] O. M. Reuter det.”; 4, “Mus. Zool. H:fors,
Spec. typ. No. 9681, Phytocoris bakeri O.M. Reut.”; 5 (here added), “Lec-
totype: Phytocoris bakeri Reuter, by Henry and Stonedahl, 1 983” [UZMH;
good condition, except the 4th antennal segments are missing and the
hemelytra are spread]. Measurements of lectotype: BL 5.25; BW ca. 1.67
(wings spread); HW 0.90; V 0.36; RL 2.02; AI 1.28; All 2.08; AIII 1.28;
AIV missing; PL 0.76; PW 1.40. Paralectotypes: 7 males and 2 females,
same data as for lectotype [5 males and 2 females, CAS; 1 male, UZMH;
2 males, USNM].
Remarks. Phytocoris bakeri is known only from California. Knight (1968)
included it in his key to the western species of Phytocoris.
Phytocoris bipunctatus Van Duzee
[See annulicornis (Reuter)]
Phytocoris breviusculus Reuter
Phytocoris breviusculus Reuter, 1876:68. Type data. — Lectotype male (here
designated): Label 1, “Texas”; 2, “Paratypus”; 3, “417 82”; 4, “Riks-
museum Stockholm”; 5 (here added), “Lectotype: Phytocoris breviusculus
Reuter, by Henry and Stonedahl, 1983” [NRS; good condition, except the
right hindleg and left antenna are missing, and the specimen is pinned
through the right hemelytron]. Measurements of lectotype: BL 4.05; BW
1.57; HW 0.90; V 0.30; RL 1 .84; AI 0.59; All 1 .46; AIII 0.90; AIV missing;
PL 0.79; PW 1.44. Paralectotypes: Same data as for lectotype [1 male and
2 females, NRS; 1 female, UZMH].
Remarks. Phytocoris breviusculus has been keyed and redescribed by Knight
(1941). Froeschner (1949) included it in his key to the Missouri species.
Wheeler and Henry (1977) described and figured the adult and 5th-instar
nymph, studied biology, summarized the literature, and firmly associated
this predatory species with Juniperus spp. and other conifers, as well as with
apple, crabapple, and honeylocust. Phytocoris breviusculus is widespread
from Pennsylvania, south to Alabama, and west to Colorado and Texas
(Carvalho, 1959; Wheeler and Henry, 1977).
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447
Phytocoris canadensis Van Duzee
Phytocoris inops Uhler, 1878:402 [preoccupied by Phytocoris inops Uhler,
1877]. Type data. — Lectotype [sex?] (here designated): Label 1, “53”; 2,
“83”; 3, “M.C.Z. Type 26446”; 4 (here added) “Lectotype: Phytocoris
inops Uhler [1878], by Henry and Stonedahl, 1983” [MCZ; condition very
poor; pinned through scutellum; all legs and antennae, hemelytra (except
for clavi), and abdomen missing]. Measurements of lectotype: HW 1.06;
V 0.40; RL 2.76; PL 0.92; PW 1.76.
Phytocoris canadensis Van Duzee, 1920:346 [Kearney, Ontario, July 29,
1911; CAS type No. 2002].
Remarks. Phytocoris canadensis was described from a unique male. Knight
(1941) redescribed and keyed this species and illustrated male genitalia.
Phytocoris inops Uhler (1878) has remained a mystery since its original
description. Uhler (1877, 1878) used the name Phytocoris inops in two
separate descriptions of new species. It has been assumed that both descrip-
tions referred to the same species (Carvalho, 1959) or that the “eastern”
inops (1878) was a species of Neurocolpus and the “western” inops (1877)
was a true species of Phytocoris (Knight, in Blatchley, 1926:699). We have
examined what we believe to be the original material studied by Uhler (1 878),
and hnd that the first description (1877) is based on a different species (see
Phytocoris inops Uhler, 1877, listed in this paper) than the second (1878).
Two specimens in the W. T. Harris collection (MCZ) bear the labels “53”
just as Uhler (1878) cited. One specimen (also having “N.H.” on the “53”
label) is in relatively good condition, but is a female and pertinent markings
are obscured by body oils. A second specimen (designated as the lectotype
above) is badly broken, but distinct markings on the propleura and pronotum
have enabled us to identify this specimen as what Van Duzee (1920) later
described as canadensis. Phytocoris canadensis, thus, is a junior synonym
of inops Uhler (1878), but inops (1878) is a junior primary homonym of
inops Uhler (1877). Phytocoris canadensis is the next available name, as we
recognize above.
Phytocoris canescens Reuter
Phytocoris canescens Reuter, 1909:30. Type data. — Lectotype male (here
designated): Label 1, “Claremont, Cal., Baker”; 2, Phytocoris canescens
Rent.” (handwritten); 3 (here added), “Lectotype: Phytocoris canescens
Reuter, by Henry and Stonedahl, 1983” [USNM type No. 100387; good
condition, except left 3rd and 4th antennal segments are missing, the right
2nd, 3rd, and 4th segments are curled, and the hemelytra are spread].
Measurements of lectotype: BL 6.42; BW 1.44; HW 0.88; V 0.36; RL
2.29; AI 1.46; All 2.80; AIII 1.26; AIV ca. 0.84; PL 0.66; PW 1.24.
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NEW YORK ENTOMOLOGICAL SOCIETY
Paralectotypes: Same data as for lectotype [1 female, USNM; 1 male,
LACM; 1 male (abdomen missing), UZMH; 3 males and 1 female, CAS].
Remarks. Phytocoris canescens, described and known only from Califor-
nia, can be keyed in Knight (1968).
Phytocoris confluens Reuter
Phytocoris confluens Reuter, 1909:20. Type data (Fig. 5). — Lectotype male
(here designated): Label 1, 2, “Wash[i]ngt[o]n, D.C., 10-7”; 3, “O.
Heidemann Collector”; 4, “Spec, typ.”; 5, '^Phytocoris puella var. con-
fluens n. [handwritten] O. M. Reuter det.”; 6, “Mus. Zool. H:fors. Spec,
typ. No. 9663, Phytocoris puella var. confluens O.M. Reut.”; 7 (here added),
“Lectotype: Phytocoris confluens Reuter, by Henry and Stonedahl, 1983”
[UZMH; condition good, except left front tibia, left 2nd, 3rd, and 4th
antennal segments, and right 4th antennal segment are missing, and the
hemelytra are spread]. Measurements of lectotype: BL 5.17; BW ca. 1.50;
HW 0.84; V 0.24; rostrum broken; AI 1.08; All 2.88; AIII 1.40; AIV
missing; PL 0.72; PW 1.28. Paralectotypes: 3 males, same data as for
lectotype with dates June 23-05, 6-7-90, and 4-7-97 [CAS, UZMH, USNM].
Phytocoris confluens: Knight, 1923:650 (as species).
Remarks. Knight (1923, 1941) redescribed confluens, provided an iden-
tification key, and gave oak, Quercus sp., and red or river birch, Betula nigra
L., as its hosts. Froeschner (1949) included confluens in his key to the
Phytocoris of Missouri. This species is widely distributed in the eastern
United States (Carvalho, 1959).
Phytocoris conspersipes Reuter
Phytocoris conspersipes Reuter, 1909:22. Type data. — Lectotype male (here
designated): Label 1, “Wash[i]ngt[o]n, D. C., 16-7-97”; 2, “O. Heidemann
Collector”; 3, “Mus. Zool. H:fors, Spec. typ. No. 9677, Phytocoris con-
spersipes O.M. Reut.”; 4 (here added), “Lectotype: Phytocoris conspersipes
Reuter, by Henry and Stonedahl, 1983” [UZMH; fair condition, with right
antenna and left 4th antennal segment missing, and the left hindleg glued
to point beside specimen]. Measurements of lectotype: BL 4.58; BW 1.83;
HW 1.00; V 0.40; RL 1.96; AI 0.64; All 1.68; AIII 1.00; AIV missing;
PL 0.76; PW 1.44. Paralectotypes: Same data as for lectotype with dates
10-9, 7-10, 10-7, 26-7-95, 27-7-95, 5-10-96, 16-7-97, 20-7-97; 1-XI[1
male and 3 females, UZMH; 1 male and 2 females, CAS; 2 females,
USNM].
Remarks. Knight (1923) redescribed this pine-inhabiting species, figured
male genitalia (1941), and provided keys to separate it from other eastern
species of Phytocoris.
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449
Phytocoris covilleae Knight
[See ramosus Uhler]
Phytocoris cunealis Van Duzee
Phytocoris cunealis Van Duzee, 1914:16. Type data. — Lectotype male (here
designated): Label 1, “San Diego Co., Cal., 6-5-13, EP Van Duzee”; 2,
“Lectotype cunealis'^ (in red, handwritten); 3, “EP Van Duzee Collection”;
4 (here added), “Lectotype: Phytocoris cunealis Van Duzee, by Henry and
Stonedahl, 1983” [CAS type No. 2005; good condition; right hindleg glued
to point beside specimen]. Measurements of lectotype: BL 6.75; BW 2.29;
HW 1.02; V 0.45; RL 3.33; AI 1.71; All 3.06; AIII 1.67; AIV 1.13; PL
1 .03; PW 1.71. Paralectotypes: 10 males and 9 females, same locality data
as for lectotype with dates 6 May-5 June 1913 and 13 April-6 June 1914
[CAS; 1 male, USNMj.
Remarks. Phytocoris cunealis is known only from southern California (Van
Duzee, 1914). Knight (1968) figured male genitalia and included this species
in his key to the western Phytocoris.
Phytocoris eximius Reuter
Phytocoris eximius Reuter, 1876:67. Type data (Fig. 3). — Lectotype male
(here designated): Label 1, “Texas”; 2, “Belfrage”; 3, "^eximius, Typ.,
Rent.” (handwritten); 4, “Typus” (red label); 5, “310, 82”; 6, “Riksmu-
seum Stockholm”; 7 (here added), “Lectotype: Phytocoris eximius Reuter,
by Henry and Stonedahl, 1983” [NRS; good condition, except the right
3rd and 4th antennal segments are missing, and the specimen is pinned
through the right hemelytronj. Measurements of lectotype: BL 6.25; BW
ca. 1.83 (hemelytra slightly spread); HW 1.00; V 0.28; RL 2.36; AI 1.12;
All 2.72; AIII 1.48; AIV 1.08; PL 0.88; PW 1.64. Paralectotypes: No
other syntypes located.
Phytocoris penepectus Knight, 1920:58. NEW SYNONYMY [described from
East River, Connecticut; USNM type No. 100388].
Phytocoris penepecten: Ycaighy 1923:640; 1941:199 (unjustified emendation
of penepectus— SQQ Steyskal, 1973:208).
Remarks. There is a large amount of literature using the name eximius
(Carvalho, 1959). Most or all of these records, however, are in error. Typical
of many early descriptions of Phytocoris, Reuter’s description probably was
based on a composite of species. Phytocoris eximius belongs to group II of
Knight’s (1941) treatment of Phytocoris. Apparently no one before us ex-
amined the above lectotype that clearly belongs to Reuter’s “type” series.
Knight ( 1 920) redescribed eximius in detail and figured male genitalia, stating
that his redescription was based on a specimen returned by Reuter to Hei-
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NEW YORK ENTOMOLOGICAL SOCIETY
demann at the USNM. We searched the USNM, but could not find the
specimen studied by Knight. Knight apparently reconsidered his definition
of eximius, for later (1941), he recognized canadensis Van Duzee for the
species he called eximius in 1920 (using the same figures and description).
Knight did not attempt to further clarify the identity of eximius after his
reversed decision.
Male genitalia of the “Belfrage” specimen clearly show that eximius is the
senior synonym of penepectus Knight (1920). Most early records of eximius
are confused and should be disregarded. Phytocoris eximius can be recog-
nized using Knight’s (1923, 1941) keys and descriptions of penepectus. This
species is widespread in the eastern U.S. west to Texas (Carvalho, 1959).
Phytocoris fenestratus Reuter
Phytocoris fenestratus Reuter, 1909:24. Type data. — Lectotype female (here
designated): Label 1, “Retreat, NC, 1-6”; 2, “O. Heidemann Collector”;
3, “Spec, typ.”; 4, ""Phytocoris fenestratus n. sp. [handwritten] O. M. Reuter
det.”; 5 (here added), “Lectotype: Phytocoris fenestratus Reuter, by Henry
and Stonedahl, 1983” [USNM type No. 100389; fair condition, with right
middle and hindlegs and left antenna missing]. Measurements of lectotype:
BL 7.75; BW 2.33; HW 1.16; V 0.44; RL 3.20; AI 1.20; All 3.08; AIII
1.64; AIV 0.92; PL 1.04; PW 2.00. Paralectotypes: 2 females. Wash., D.C.,
dates 16-V-02 and 27-6-95, Heidemann coll. [UZMH]; 1 female, same
data as for lectotype [CAS].
Remarks. This pine-inhabiting species, known from Connecticut, Georgia,
North Carolina, and Washington, D.C., has been redescribed and keyed by
Blatchley (1926). Henry (1979) figured male genitalia and separated fenes-
tratus from two closely related species, discoidalis Henry (now a junior
synonym of dreisbachi Knight (Henry, 1982a)) and intermedius Henry.
Phytocoris formosus Van Duzee
Phytocoris reuteri Van Duzee, 1914:18 (name preoccupied by P. reuteri
Saunders, 1875. Type data. — Lectotype male (here designated): Label 1,
“San Diego Co., Cal., 7-4-13, EP Van Duzee”; 2, “Lectotype reuterV' (in
red, handwritten); 3, “EP Van Duzee Collection”; 4 (here added), “Lec-
totype: Phytocoris reuteri Van Duzee, by Henry and Stonedahl, 1983”
[CAS type No. 2000; good condition; antennal segment IV on right side
missing]. Measurements of lectotype: BL 6.00; BW 1.69; HW 0.92; V
0.30; RL ca. 2.88 (apex obscured by point and glue); AI 1.40; All 2.57;
AIII 1.87; AIV 1.24; PL 0.81; PW 1.35. Paralectotypes: 6 males and 10
females, same locality data as for lectotype with dates 4 July 1913 and 5
August 1913 [14, CAS; 2 USNM].
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451
Phytocoris formosus Van Duzee, 1916:37 (new name for P. reuteri Van
Duzee).
Remarks. Phytocoris formosus is known only from Southern California
(Van Duzee, 1914). Knight (1968) recorded Cordylanthus filifolius Nutt, (as
Adenostegia filifolia) as the host and included formosus in his key to the
western species of Phytocoris.
Phytocoris fumatus Reuter
Phytocoris fumatus Reuter, 1909:25. Type data. — Lectotype female (here
designated): Label 1, “Wash[i]ngt[o]n, D. C. 20-6”; 2, “Heidemann Col-
lector”; 3, “7”; 4, “Spec, typ.”; 5, Phytocoris fumatus n. sp. [handwritten]
O. M. Reuter det.”; 6 (here added), “Lectotype: Phytocoris fumatus Reuter,
by Henry and Stonedahl, 1983” [USNM type No. 100391; fair condition,
with antennae and legs on left side missing]. Measurements of lectotype:
BL 7.58; BW 2.58; HW 1. 16; V 0.48; RL 4.83; antennae missing; PL 1.24;
PW 2.00. Paralectotypes: No other syntypes located.
Phytocoris subnitidulusKtuXtr, 1909:26 (synonymized by Knight, 1920:63).
Type data. — Lectotype female (here designated): Label 1, “Plummer’s [sic]
I[sland]., Md. [date obscured by glue]”; 2, “O. Heidemann Collector”; 3,
^"Phytocoris subnitidulus n. sp. [handwritten] O. M. Reuter det.”; 4 (here
added), “Lectotype: Phytocoris subnitidulus Reuter, by Henry and Stone-
dahl, 1983” [USNM type No. 100390; good condition, except the 3rd and
4th antennal segments are missing and a pin has been inserted into and
removed from the right hemelytron; the left hindleg is glued to the locality
label below the specimen]. Measurements of lectotype: BL 7.25; BW ca.
2.50 (hemelytra slightly spread); HW 1.16; V 0.44; RL 4.83; AI 1.92; All
3.75; AIII and AIV missing; PL 1.12; PW 1.92. Paralectotype: 1 male.
Wash., D.C., 10-6, Heidemann [UZMH].
Remarks. This widespread eastern U.S. species (Carvalho, 1959) has been
redescribed and keyed by Knight (1923, 1941). Knight (1920, 1941) figured
male genitalia and correctly placed subnitidulus as a junior synonym of
fumatus. Froeschner (1949) included fumatus in his key to the Phytocoris
of Missouri. We have chosen the female from Plummers Island as the lec-
totype because of the very poor condition of the male from Washington,
D.C.
Phytocoris heidemanni Reuter
Phytocoris heidemanni Reuter, 1909:27. Type data (Fig. 4). — Lectotype fe-
male (here designated): Label 1, “Pecos, N. M., June 23 (C[oc]k[ere]ll)”
(handwritten); 2, “O. Heidemann Collector”; 3, ""Phytocoris heidemanni
n. sp. [handwritten] O. M. Reuter det.”; 4 (here added), “Lectotype: Phy-
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NEW YORK ENTOMOLOGICAL SOCIETY
tocoris heidemanni Reuter, by Henry and Stonedahl, 1983” [USNM type
No. 100401; good condition, except the right middle and right hindlegs
are missing, and a pin has been inserted through and removed from the
scutellum; right foreleg glued to point beside specimen]. Measurements
of lectotype: BL 7.92; BW 2.75; HW 1.32; V 0.52; RL 3.96; AI 1.84; All
3.68; AIII 1.68; AIV 1.24; PL 1.16; PW 2.12. Paralectotypes: No other
syntypes located.
Remarks. This western pine-inhabiting species, recorded from Arizona,
Colorado, New Mexico, and Nevada (Carvalho, 1959), is included in Knight’s
(1968) key to the western species of Phytocoris.
Phytocoris hesperellus Knight
[See inops Uhler]
Phytocoris hesperius Knight
[See inops Uhler]
Phytocoris infuscatus Reuter
Phytocoris puella var. infuscatus Reuter, 1909:20. Type data. — Lectotype
male (here designated): Label 1, “5”; 2, “Wash[i]ngt[o]n, D. C., June 23,
04”; 3, “Heidemann Collector”; 4, “Mus. Zool. H:fors, Spec. typ. No.
9664, Phyt. puella var. infuscata, O. M. Reuter” (handwritten); 5 (here
added), “Lectotype: Phytocoris infuscatus Reuter, by Henry and Stone-
dahl, 1983” [UZMH; good condition, except the right hindleg is missing
and specimen is pinned through left clavus]. Measurements of lectotype:
BL 5.92; BW 2.17; HW 0.96; V 0.48; RL 2.76; AI 1.08; All 2.76; AIII
1.40; AIV 1.00; PL 0.84; PW 1.64. Paralectotypes: 1 female. Wash., D.C.,
27-7-91, Heidemann [UZMH]; 2 males, 2 females. Rock Creek, 17-6-93
and 24-6-93, Heidemann [1 CAS; 1 UZMH; 2 USNM].
Phytocoris infuscatus: Van Duzee, 1914:16 (as species).
Remarks. Phytocoris infuscatus, an eastern U.S. species (Carvalho, 1959),
has been redescribed and keyed (Knight, 1923, 1941; Blatchley, 1926). Knight
(1923, 1941) figured male genitalia. Froeschner (1949) included infuscatus
in his key to the species of Phytocoris from Missouri.
Phytocoris inops Uhler
Phytocoris inops Uhler, 1877:413. Type data. — Lectotype female (here des-
ignated): Label 1, “Clear Cr[eek]. Canon” (handwritten); 2, “PR Uhler
Collection”; 3 (here added), “Lectotype: Phytocoris inops Uhler [1877],
by Henry and Stonedahl, 1983” [USNM type No. 100392; condition fair;
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453
left 3rd and 4th and right 4th antennal segments and middle left and both
hindlegs missing, and specimen is pinned between hemelytra just below
apex of scutellum]. Measurements of lectotype: BL 7.58; BW ca. 2.50;
HW 1.05; V 0.48; RL 3.28; AI 1.80; All 3.48; AIII 1.68; AIV missing;
PL 1.00; PW 1.84. Paralectotypes: No other syntypes located.
Phytocoris vittatus Reuter, 1909:28. NEW SYNONYMY. Type data. — Lec-
totype female (here designated): Label 1, “Lake Placid, NY, 8-12-04”; 2,
“Van Duzee Collector”; 3, ^"Phytocoris vittatus n. sp. [handwritten] O. M.
Reuter det.”; 4 (here added), “Lectotype: Phytocoris vittatus Reuter, by
Henry and Stonedahl, 1983” [USNM type No. 100395; good condition,
except the right middle leg, left antenna, and right 4th antennal segment
are missing; the left hindleg is detached and glued to point]. Measurements
of lectotype: BL 7.42; BW 2.33; HW 1.04; V 0.44; RL 3.24; AI 1.72; All
3.44; AIII 1.72; AIV missing; PL 0.96; PW 1.76. Paralectotypes: No other
syntypes located.
Phytocoris palmeri Reuter, 1909:32. NEW SYNONYMY. Type data. — Lec-
totype male (here designated): Label 1, “Quinze L[a]k[e]., P[rovince].
Q[uebec]., 8-14-07, W.J. Palmer”; 2, “Holotype palmerr (handwritten);
3, “E. P. Van Duzee Collection”; 4 (here added), “Lectotype: Phytocoris
palmeri Reuter, by Henry and Stonedahl, 1983” [CAS type No. 1994;
poor condition, with right 2nd, 3rd, and 4th antennal segments missing,
the right hemelytron broken on the basal Vi of corium, and the wing
membrane folded downward]. Measurements of lectotype: BL ca. 6.83
“(head turned up and wing membrane folded down); BW 1.08; HW 1.05;
V 0.36; RL ca. 3.00 (slightly bent); AI 1.40; All 3.08; AIII 1.52; AIV
1.20; PL 0.92; PW 1.76. Paralectotypes: 1 female, Quinze Lake, PQ, 8-14-
07 [USNM].
Phytocoris hesperius Knight, 1928:44. NEW SYNONYMY [described from
Stonewall, Las Animas County, Colorado; USNM type No. 100393].
Phytocoris hesperellus Knight, 1968:232. NEW SYNONYMY [described
from Salt Lake City, Salt Lake County, Utah; USNM type No. 100394].
Remarks. Phytocoris inops, like eximius Reuter, has been an enigma to
mirid workers since its original description. Uhler (1877) probably had a
composite of species when he considered the range of inops to be Colorado,
Maryland, Massachusetts, New Jersey, Rhode Island, and “lower Canada.”
His remarks that this species resembled Say’s species Phytocoris nubilus
(now in the genus Neurocolpus), that it could be confused in the field with
the genus Psocus (Psocidae), and that some specimens in the “south” had
lead-colored or bluish markings at the apex of the corium further confused
the identity of inops. Additionally, Uhler (1878) published a second de-
scription of inops as a new species based on another species of Phytocoris
(See Phytocoris canadensis listed in this paper).
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NEW YORK ENTOMOLOGICAL SOCIETY
Only one of Uhler’s (1877) localities, “Beaver Brook Gulch, next to Clear
Creek Canon,” can be associated with the species inops. We have found a
single female in with undetermined material at the USNM that bears the
single label “Clear Cr. Canon.” This specimen exactly fits Uhler’s (1877)
description, making us certain that this specimen belongs to his syntype
series.
Because of past confusion, early distribution records of inops cannot be
trusted. Based on the above synonyms, its distribution can be confirmed for
Arizona, Colorado, Nevada, New Mexico, New York, Oregon, Wyoming,
and Quebec. Knight (1923) redescribed (as palmeri) and keyed (1923, 1941)
(as palmeri and vittatus, respectively) inops.
Phytocoris interspersus Uhler
Phytocoris interspersus Uhler, 1895:32. Type data.— Neotype male (here
designated): Label 1, “F[or]t. Garland, Colo., Ute Creek Ranch, Aug. 1 1,
1925, H. H. Knight”; 2, “H. H. Knight Collection”; 3 (here added), “Neo-
type: Phytocoris interspersus Uhler, by Henry and Stonedahl, 1 983” [USNM
type No. 100396; excellent condition]. Measurements of neotype: BL 6.92;
BW 2.25; HW 1.04; V 0.28; RL 2.48; AI 1.12; All 2.64; AIII 1.48; AIV
1.16; PL 0.88; PW 1.60.
Remarks. We could not find the original female described from “Cheyenne
Canon, Colorado Springs, July (Tucker)”; therefore, we have designated the
above specimen from the same general type locality as the neotype to rep-
resent this species. This specimen agrees with Uhler’s description and sub-
sequently identified material of the species.
Knight (1968) recorded Phytocoris interspersus from Arizona, California,
Colorado, Idaho, New Mexico, Utah, and British Columbia, and included
it in his key to the western species of Phytocoris.
Phytocoris jucundus Van Duzee
Phytocoris jucundus Van Duzee, 1914:17. Type data. — Lectotype male (here
designated): Label 1, “San Diego Co., Cal., 10-19-13, EP Van Duzee”; 2,
“Lectotype jucundus'' (in red, handwritten); 3, “EP Van Duzee Collec-
tion”; 4 (here added), “Lectotype: Phytocoris jucundus Van Duzee, by
Henry and Stonedahl, 1983” [CAS type No. 1998; fair condition; left
foreleg and hindleg missing; antennal segments II-IV on right side missing;
hemelytra slightly spread]. Measurements of lectotype: BL 6.59; BW ca.
2.34; HW 1.09; V 0.34; RL ca. 2.41; AI 1.31; All 2.95; AIII 1.48; AIV
0.97; PL 0.95; PW 1.64. Paralectotypes: 8 females, same label data as for
lectotype [7, CAS; 1, USNM].
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455
Remarks. Phytocoris jucundus is known from California, Idaho, Oregon,
and Washington where it occurs commonly on Firms spp. Knight (1968)
included it in his key to the western species of Phytocoris.
Phytocoris laevis (Uhler)
Callodemas laevis Uhler, 1895:33. Type data. — Lectotype male (here des-
ignated): Label 1, “Albuq[uerque], N. M., 9-19-88 [handwritten, printing
unclear for year]”; 2, “PR Uhler Collection”; 3, ^"Callodemas laevis Uhler,
N. M., Osborn” (handwritten); 4, ""Callodemas laevis, N. M., det. Uhler”
(handwritten); 5 (here added), “Lectotype: Callodemas laevis Uhler, by
Henry and Stonedahl, 1983” [USNM type No. 100397; condition poor,
specimen glued to card, abdomen partially eaten by dermestids, and many
appendages broken but glued to card]. Measurements of lectotype: BL
9.00; BW 2.17; HW 1.12; V 0.36; RL ca. 2.48 (obscured under body); AI
1.76; All 3.04; AIII and AIV missing; PL 1.12; PW 2.04. Paralectotypes:
1 female, Colorado [USNM].
Phytocoris laevis: Reuter, 1909:14 (new combination).
Remarks. Phytocoris laevis is known from Arizona and New Mexico, north
to Alberta, Canada (Knight, 1968). Knight (1968) included laevis in his key
to the western species of Phytocoris.
Phytocoris lasiomerus Reuter
Phytocoris lasiomerus Reuter, 1909:34. Lectotype male (here designated):
Label 1, “Long Lake [New York]”; 2, “typus”; 3 (here added), “Lectotype:
Phytocoris lasiomerus Reuter, by Henry and Stonedahl, 1983” [TMB;
good condition, except 3rd and 4th segments on both antennae missing].
Measurements of lectotype: BL 7.67; BW 2.25; HW 1.04; V 0.44; RL
3.40; AI 1.68; All 3.32; AIII and IV missing. Paralectotypes: 1 female.
Cold River [New York]; 1 male. Huckleberry [New York] (both TMB).
Remarks. Phytocoris lasiomerus is known from Quebec south to Massa-
chusetts and New York, west to Colorado, Washington, and Wyoming
(Knight, 1941). Knight (1923) correctly redescribed and keyed this northern
species.
Phytocoris lineatus Reuter
Phytocoris lineatus Reuter, 1909:30. Type data (Fig. 2). — Lectotype male
(here designated): Label 1, “Rifle, Oil, 7-25-00”; 2, “Holotype lineatus""
(in red, handwritten); 3, “EP Van Duzee Collection”; 4 (here added),
“Lectotype: Phytocoris lineatus Reuter, by Henry and Stonedahl, 1983”
[CAS type No. 1993; fair condition, except all of the right legs and antennal
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NEW YORK ENTOMOLOGICAL SOCIETY
segments III and IV (left) and II, III, IV (right) are missing]. Measurements
of lectotype: BL 6.00; BW 1.67; HW 0.92; V 0.44; RL 3.04; AI 1.64; All
3.28; AIII broken at apex; AIV missing; PL 0.80; PW 1 .40. Paralectotypes:
No other syntypes located.
Remarks. Phytocoris lineatus has not been treated since its original de-
scription and is known only from Colorado.
Phytocoris marmoratus (Van Duzee)
[See vanduzeei Reuter]
Phytocoris mi nut ulus Reuter
Phytocoris minutulus Reuter, 1909:24. Type data. — Neotype male (here des-
ignated): Label 1, “Plummers I[sland], 6-7-06, Md.”; 2, “D. H. Clemons
Collector”; 3, ""Phytocoris minutulus Reut., Det. H. H. Knight”; 4 (here
added), “Neotype: Phytocoris minutulus Reuter, by Henry and Stonedahl,
1983” [USNM type No. 100398; excellent condition except the left 4th
antennal segment is missing]. Measurements of neotype: BL 4.48; BW
1.60; HW 0.88; V 0.26; RL 2.22; AI 1.04; All 2.22; AIII 1.44; AIV 0.90;
PL 0.62; PW 1.14.
Remarks. Phytocoris minutulus was described from a single female taken
on Plummers Island, Md., 26 July 1903, by O. Heidemann. We have been
unable to locate this specimen which should be in the USNM; therefore, we
have chosen the above specimen from the same type locality as the neotype.
This species is recorded from Maryland, Massachusetts, New Hampshire,
NewYork, North Carolina, Pennsylvania, and Virginia (Knight, 1941). Knight
(1923) and Blatchley (1926) redescribed and keyed it with other eastern U.S.
species of Phytocoris.
Phytocoris mundus Reuter
Phytocoris mundus Reuter, 1909:18. Type data. — Lectotype male (here des-
ignated): Label 1, “Wash[i]ngt[o]n, D. C., 19-IV-02”; 2, “10”; 3, “O.
Heidemann Collector”; 4, ""Phytocoris sp.^ near breviusculus Reut.” (hand-
written); 5, ""Phytocoris mundus Uhl. [handwritten] O. M. Reuter det.”;
6 (here added), “Lectotype: Phytocoris mundus Reuter, by Henry and
Stonedahl, 1983” [USNM type No. 100399; good condition, except the
right middle leg and segments III and IV are missing on both antennae].
Measurements of lectotype: BL 4.83; BW 1.75; HW 0.92; V 0.40; RL
2.00; AI 0.64; All 1.80; AIII and AIV missing; PL 0.72; PW 1.36. Para-
lectotypes: 1 female. Wash., D.C. 1-7-97, Heidemann [USNM]; 1 female.
Rock Crk., D.C., 6-7-97, Heidemann [CAS]; 1 female, Bladnsbg., Md.,
20-7-92, Heidemann [CAS].
VOLUME 91, NUMBER 4
457
Remarks. Phytocoris mundus, a common eastern, pine-inhabiting species,
has been redescribed and keyed (Knight, 1923, 1941). Knight (1941) figured
male genitalia.
Phytocoris nigripubescens Knight
[See vanduzeei Reuter]
Phytocoris pallidicornis Reuter
Phytocoris pallidicornis Reuter, 1876:69. Type data. — Holotype female: La-
bel 1, “Wisconsin”; 2, “Kumlien.”; 3, “Typus”; 4, ""pallidocornis Rent.”
(folded, handwritten); 5, “311, 82”; 6, “Riksmuseum Stockholm”; 7 (here
added), “Holotype: Phytocoris pallidicornis Reuter, by Henry and Stone-
dahl, 1983” [NRS; condition poor, with the antennae and all but 3 legs
missing, the wing membrane badly folded, and the right clavus forced up
because the specimen is pinned through the middle of the body]. Mea-
surements of holotype: BL ca. 6.67 (wing membrane folded under); BW
2.33; HW 1.00; V 0.44; RL 3.24; antennae missing; PL 0.92; PW 1.64.
Phytocoris pallicornis: Reuter, 1909:33 (misspelling).
Remarks. Reuter (1 876) described this species from a unique female (Reu-
ter, 1909). Phytocoris pallidicornis is a widespread northern species that
occurs over much of northeastern U.S., west to Colorado and British Co-
lumbia. It has been redescribed and keyed by Knight (1923, 1941), Blatchley
(1926), and Kelton (1980); Kelton also figured male genitalia.
Phytocoris palmeri Reuter
[See inops Uhler]
Phytocoris penepectus Knight
[See eximius Reuter]
Phytocoris politus Reuter
Phytocoris politus Reuter, 1909:21. Type data. — Lectotype male (here des-
ignated): Label 1, “Ormsby Co., Nev., July, Baker”; 2, “Mus. Zool. H:
fors. Spec. typ. No. 9672, Phytocoris politus O.M. Reut.”; 3 (here added),
“Lectotype: Phytocoris politus Reuter, by Henry and Stonedahl, 1983”
[UZMH; poor condition with one segment III and both 4th antennal
segments, and all but three legs missing, the hemelytra spread apart, and
the apical parts of the wing membrane broken]. Measurements of lectotype:
BL 6. 17; BW ca. 2.00; HW 1. 12; V 0.36; RLca. 2.28 (broken and imbedded
in glue); AI 1.12; All 2.76; AIII 1.24; AIV missing; PL 0.96; PW 1.80.
458
NEW YORK ENTOMOLOGICAL SOCIETY
Paralectotypes: 3 females, same locality data as for lectotype [1, UZMH;
2, USNM].
Phytocoris rusticus Van Duzee, 1920:348. NEW SYNONYMY [described
from Mt. St. Helena, Napa County, California; CAS type No. 699].
Remarks. Phytocoris politus is known from California, Idaho, Oregon, and
Washington. Knight (1968) included it in his key to the western species of
Phytocoris.
Phytocoris puella Reuter
Phytocoris puella Reuter, 1876:69. Type data. — Lectotype male (here des-
ignated): Label 1, “N[ewj. York”; 2, “Belfrage”; 3, ""puella Rent.” (folded,
handwritten); 4, “Typus”; 5, “312, 82”; 6, “Riksmuseum Stockholm”; 7
(here added), “Lectotype: Phytocoris puella Reuter, by Henry and Stone-
dahl, 1983” [NRS; condition poor (slightly teneral). The specimen is pinned
through the right hemelytron and mounted on a hollow plastic tube that
is attached to the label pin; the left 2nd, 3rd, and 4th antennal segments
are missing; the hemelytra are folded up; and the base of the abdomen
has been eaten by dermestidsj. Measurements of lectotype: BL 4.67; BW
(not measured; body distorted); HW 0.80; V 0.24; RL 1.96; AI 0.96; All
2.04; AIII 1.24; AIV 1.16; PL 0.64; PW 1.20. Paralectotypes: No other
syntypes located.
Remarks. Phytocoris puella is a widespread eastern U.S. species (Carvalho,
1959) found on numerous deciduous trees, including Cary a and Quercus
spp. Knight (1923, 1941) and Blatchley (1926) redescribed and keyed this
species. Froeschner (1949) included puella in his key to the Phytocoris of
Missouri.
Phytocoris ramosus Uhler
Phytocoris ramosus Uhler, 1894:252. Type data. — Lectotype female (here
designated): Label 1, “S[anj. Berna[r]dino Co., Cal.”; 2, “Uhler Type”; 3,
“9”; 4, “Lectotype ramosus'^ (in red, handwritten); 5, ""Phytocoris ramosus
Uhl.” (handwritten); 6 (here added), “Lectotype: Phytocoris ramosus Uhl-
er, by Henry and Stonedahl, 1983” [CAS type No. 555; poor condition,
with right middle and forelegs, left middle and hindlegs, and antennae
missing; wing membrane curled]. Measurements of lectotype: BL 5.56;
BW 2.16; HW 1.01; V 0.40; RL ca. 2.07; antennae missing; PL 1.01; PW
1.87. Paralectotypes: 2 females, same data as for lectotype except with
additional CAS type identification label [CAS].
Phytocoris covilleae Knight, 1925:54 (synonymized by Carvalho, 1959:214).
Remarks. Knight (1968) recorded this species from Arizona, California,
VOLUME 91, NUMBER 4
459
Nevada, and Utah, and included it in his key to the western species of
Phytocoris.
Phytocoris reuteri Van Duzee
[See formosus Van Duzee]
Phytocoris roseus (Uhler)
Compsocerocoris roseus Uhler, 1894:253. Type data. — Lectotype male (here
designated): Label 1, “San Borja, Lower Cal., Mex., Chas D. Haines, May
1889”; 2, “778”; 3, “5”; 4, “Lectotype roseus"' (in red, handwritten); 5
(here added), “Lectotype: Compsocerocoris roseus Uhler, by Henry and
Stonedahl, 1983” [CAS type No. 557; poor condition with hindlegs, left
antenna, and 3rd and 4th segments of right antenna missing; pinned through
scutellum and hemelytra slightly spread; abdomen partially eaten by der-
mestids but leaving genital segment intact]. Measurements of lectotype:
BL 7.34; BW ca. 2.20; HW 1.07; V 0.34; RL 3.19; AI 1.67; All 3.1 1; AIII
and AIV missing; PL 0.95; PW 1.76. Paralectotypes: 2 females, Los An-
geles, no other data [USNM].
Phytocoris roseus: Reuter, 1909:27 (new combination).
Phytocoris barbatus Van Duzee, 1920:353 (synonymized by Carvalho, 1959:
214).
Remarks. Phytocoris roseus is known only from southern California and
Mexico (Carvalho, 1959). Knight (1968) included it in his key to the western
species of Phytocoris.
Phytocoris rufoscriptus Van Duzee
Phytocoris rufoscriptus Van Duzee, 1914:15. Type data. — Lectotype male
(here designated): Label 1, “San Diego Co., Cal., 6-8-13, EP Van Duzee”;
2, “Lectotype rufoscriptus" (in red, handwritten); 3, “EP Van Duzee Col-
lection”; 4 (here added), “Lectotype: Phytocoris rufoscriptus Van Duzee,
by Henry and Stonedahl, 1983” [CAS type No. 2007; good condition
except the left hindleg is missing]. Measurements of lectotype: BL 7.78;
BW 2.45; HW 1.04; V 0.44; RL (not measured; tip obscured by point and
glue); AI 1.94; All 3.53; AIII 1.87; AIV 1.26; PL 1.13; PW 1.85. Para-
lectotypes: 2 males and 1 female, same data as for lectotype except dates
6-6-14 and IV-13-1913 [CAS]; 1 male, same data as for lectotype except
4-13-14 [USNM].
Remarks. Phytocoris rufoscriptus is known only from southern California
(Van Duzee, 1914). Knight (1968) included it in his key to the western species
of Phytocoris.
1
460
NEW YORK ENTOMOLOGICAL SOCIETY
Phytocoris rufus Van Duzee
Phytocoris rufus Van Duzee, 1912:477. Type data. — Lectotype male (here
designated): Label 1, “7-Oaks, Fla., May 1, ’08, Van Duzee”; 2, “Lectotype
rufus’’" (in red, handwritten); 3, “EP Van Duzee Collection”; 4 (here added),
“Lectotype: Phytocoris rufus Van Duzee, by Henry and Stonedahl, 1983”
[CAS type No. 2009; fair condition; the right 3rd and 4th antennal seg-
ments are missing; hemelytra spread, outer margins raised]. Measurements
of lectotype: BL 4.54; BW ca. 1.62; HW 0.77; V 0.25; RL ca. 1.62, tip
obscured by point and glue; AI 0.59; All 1.62; AIII 0.90; AIV 0.79; PL
0.59; PW 1.17. Paralectotypes: 3 females, same data as for lectotype [2,
CAS; 1, USNMj.
Remarks. Phytocoris rufus is known from Florida (Van Duzee, 1912),
Louisiana, and Mississippi (Knight, 1927a). Blatchley (1926) included this
species in his key to the eastern species of Phytocoris. Henry ( 1 982b) recorded
Hypericum spp. as the hosts.
Phytocoris rusticus Van Duzee
[See politus Reuter]
Phytocoris subnitidulus Reuter
[See fumatus Reuter]
Phytocoris tibialis Reuter
Phytocoris tibialis Reuter, 1876:68. Type data. — Lectotype female (here des-
ignated): Label 1, “Wisconsin”; 2, “Kumlien”; 3, “Allotypus”; 4, “414
82”; 5, “Riksmuseum Stockholm”; 6 (here added), “Lectotype: Phytocoris
tibialis Reuter, by Henry and Stonedahl, 1983” [NRS; poor condition with
legs and left antenna missing; abdomen somewhat shriveled]. Measure-
ments of lectotype: BL 5.02; BW 1.78; HW 0.82; V 0.32; RL ca. 2.07; AI
0.92; All 2.03; AIII 1.33; AIV 0.70; PL 0.72; PW 1.40. Paralectotypes:
No other syntypes could be recognized with certainty.
Compsocerocoris vilis Distant, 1883:260 [described from San Deronimo,
Guatemala and Bugaba, Panama; lectotype from Bugaba designated by
Carvalho and Dolling, 1976:809] (synonymized by Reuter, 1909:20.)
Remarks. Two male specimens of this species also were examined from
the collection of the NRS. Both of these were collected in New York by
Belfrage; one bears Reuter’s handwritten determination label reading: ‘7/-
bialis, Typ., Rent.” Although these specimens are not included in the original
description of P. tibialis, their association with the Wisconsin specimen in
the NRS and Reuter’s det. label on the one specimen suggests that they may
VOLUME 91, NUMBER 4
461
be syntypes. In the original description, “Texas (Belfrage)” is given as the
label data for at least part of the syntypes, but it is possible that “Texas”
was recorded in error, or less likely, that the specimens were mislabeled (see
label data for lectotype of puella Reuter).
Phytocoris tibialis is a widespread species occurring from eastern Canada,
west to Minnesota, and south to Panama and Venezuela (Carvalho, 1959).
Knight (1923, 1941) keyed this species, figured genitalia, and indicated that
it could be found among weedy herbaceous plants, including mountain mint
Pycnanthemum sp., in damp situations.
Phytocoris validus Reuter
Phytocoris validus Reuter, 1909:31. Type data (Fig. 6). — Lectotype male
(here designated): Label 1, “F[or]t. Collins, Colo., 7-16-03”; 2, “Van Du-
zee Collector”; 3, “E P Van Duzee Collection”; 4, ""Phytocoris validus
(Uhl.) [handwritten] O. M. Reuter det.”; 5 (here added), “Lectotype: Phy-
tocoris validus Reuter, by Henry and Stonedahl, 1983” [CAS type No.
14257; condition fair; the left antenna, left foreleg, and right hindleg are
missing; abdomen removed for dissection of genitalia and 9th segment
retained in micro vial which is attached below specimen on pin]. Mea-
surements of lectotype: BL ca. 6.83 (wing membrane folded under); BW
2.17; HW 0.96; V 0.42; RL 2.80; AI 1.24; All 2.68; AIII 1.64; AIV 0.88;
PL 1.00; PW 1.68. Paralectotypes: No other syntypes could be recognized
with certainty.
Remarks. Knight (1968) recorded validus from Colorado and North Da-
kota, and included it in his key to the western species of Phytocoris. Kelton
(1980) figured male genitalia, recorded Alberta, Manitoba, and Saskatche-
wan, and keyed it with the Phytocoris of the Prairie Provinces.
Phytocoris vanduzeei Reuter
Lygus vividus Uhler, 1894:260. Type data. — Lectotype male (here designat-
ed): Label 1 , “Comondu”; 2, “Uhler Type”; 3, “3 1 8”; 4, “5”; 5, “Lectotype
vividus'" (in red, handwritten); 6, ""Lygus vividus Uhler” (handwritten); 7
(here added), “Lectotype: Lygus vividus Uhler, by Henry and Stonedahl,
1983” [CAS type No. 561; fair condition, with left hindleg, left antenna,
and right 2nd-4th antennal segments missing; wing membrane curled
upward]. Measurements of lectotype: BL ca. 5.40; BW 1.75; HW 0.97; V
0.28; RL 1.66; AI 0.54; AII-AIV missing; PL 0.90; PW 1.58. Paralecto-
types: No other syntypes located.
Dichrooscytus rnarmoratus Van Duzee, 1910:78. Type data. — Lectotype male
(here designated): Label 1, “Alamogordo, NM, V-7-’02”; 2, “Paratype
rnarmoratus" (in orange, handwritten); 3, “EP Van Duzee Collection”; 4
(here added), “Lectotype: Dichrooscytus rnarmoratus Van Duzee, by Hen-
462
NEW YORK ENTOMOLOGICAL SOCIETY
ry and Stonedahl, 1983” [CAS type No. 2015; good conditioi except left
middle leg, left antennal segments II-IV, and right antennal i gment IV
missing]. Measurements of lectotype: BL 4.86; BW 1.64; 0.98; V
0.30; RL (not measured, obscured by point and glue); AI 0.45; All 1.86;
AIII 0.86; AIV missing; PL 0.77; PW 1.44. Paralectotypes: 1 male and 1
female [USNM]; 8 males and 5 females [CAS]; all having same locality
data as for lectotype with dates ranging from 8 March to 10 May 1902.
Phytocoris vanduzeei Reuter, 1912:30 [new name for Dichrooscytus mar-
moratus Van Duzee (combination Phytocoris marmoratus preoccupied by
Palearctic species of Douglas and Scott, 1869)]; Knight, 1917:640 (syn-
onymy of P. vanduzeei with secondary junior homonym Phytocoris vividus
(Uhler)).
Phytocoris vividus: Knight, 1917: 640 (new combination) (preoccupied by
Lygaeus vividus Fabricius, 1803).
Phytocoris nigripubescens Knight, 1925:55. NEW SYNONYMY [described
from Tucson, Pima County, Arizona; USNM type No. 100400].
Remarks. Reuter (1912) first recognized that Dichrooscytus marmoratus
Van Duzee belonged in the genus Phytocoris and that it was preoccupied by
the Palearctic species Phytocoris marmoratus Douglas and Scott. He re-
named Van Duzee’s species vanduzeei. Knight (1917) considered vanduzeei
a junior synonym of Lygus vividus Uhler, but later (1968), in treating the
western species of Phytocoris he apparently overlooked his earlier decision
and again recognized both vividus and vanduzeei. We agree with Knight’s
earlier work that Dichrooscytus marmoratus is a junior synonym of Lygus
vividus. This situation should allow for the recognition of Phytocoris vividus
(Uhler) as the acceptable name; however, Phytocoris vividus is a secondary
junior homonym of Lygaeus vividus Fabricius. Phytocoris vanduzeei is the
next available name, as we recognize above.
Phytocoris vanduzeei is known from Arizona, California, New Mexico,
and Nevada (Knight, 1968).
Phytocoris vigens (Uhler)
Calocoris vigens Uhler, 1894:255. Type data. — Lectotype male (here des-
ignated): Label 1, “San Jose del Cabo”; 2, “Uhler Type”; 3, “($”; 4, “Lec-
totype vigens"" (in red, handwritten); 5, ""Calocoris vigens Uhl., San Jose
del Cabo” (handwritten); 6 (here added), “Lectotype: Calocoris vigens
Uhler, by Henry and Stonedahl, 1 983” [CAS type No. 558; poor condition
with legs, right 3rd and 4th antennal segments, and left 4th antennal
segment missing; hemelytra spread, membrane tattered; originally pinned
through scutellum and later transferred to a point]. Measurements of lec-
totype: BL ca. 6.59; BW (not measured; hemelytra spread); HW 1.00; V
VOLUME 91, NUMBER 4
463
\'C /WVa
Hehlgmann
Collector ,
I
^.M. Reuter det.
EPVanDuzee
Collection
yhlfvage.
^SkaKfipi;’;
lOHeulemarm
1 Collector
O.M. Reuter det. I
|T^
6
5
1
iVaslyi^tn
I»C
Joiieiitemann
[Tt.Col! J18
Specityp.
P/l
7
O.M.Exniter det.
Mus. Zool. Hdors
Sttfc. typ.^No^<^.^,3
ViinD\ieee
Collector .
EPVanDuzee
Colledtion
^Q$Uc7L^/tLi.f.j
O.M. Reiter det.
Figs. 1-6. Labels for lectotypes of Phytocoris species described by Reuter. 1. angustulus. 2.
lineatus (dark-red 2nd label reads “Holotype lineatus." 3. eximius (dark-red 3rd label reads
“Typus”). 4. heidemanni. 5. confluens. 6. validus (last collection label not in type data; added
by authors).
464
NEW YORK ENTOMOLOGICAL SOCIETY
0.39; RL 2.90; AI 1.57; All 3.1 1; AIII broken or missing; AIV missing;
PL 1.08; PW 2.00. Paralectotypes: 1 male, same locality data as for lec-
totype [CAS].
Phytocoris Vigens: CdiVYdiWio, 1959:221 (new combination).
Remarks. Phytocoris vigens is known only from Baja California, Mexico
(Uhler, 1894).
Phytocoris vilis (Distant)
[See tibialis Reuter]
Phytocoris vittatus Reuter
[See inops Uhler]
Phytocoris vividus (Uhler)
[See vanduzeei Reuter]
ACKNOWLEDGMENTS
We thank P. H. Amaud, Jr. (CAS), C. L. Hogue (LACM), A. Jansson (UZMH), P. Lindskog
(NRS), A. Soos (TMB), T. Vasarhelyi (TMB), and C. Vogt (MCZ) for lending valuable type
specimens examined during this study. R. C. Froeschner (USNM), R. L. Hodges (Systematic
Ent. Lab., IIBIII, ARS, USDA, % USNM), F. C. Thompson (SEL, % USNM), and D. R.
Whitehead (SEL, % USNM) kindly reviewed the manuscript and made valuable comments.
This work was partially supported by a grant to G. M. Stonedahl from the Theodore Roosevelt
Memorial Fund, American Museum of Natural History, New York.
LITERATURE CITED
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Co., Indianapolis, 1116 pp.
Carvalho, J. C. M. 1959. Catalogue of the Miridae of the world. Part IV. Subfamily Mirinae.
Arq. Mus. Nac., Rio de Janeiro 48:1-384.
Carvalho, J. C. M. and W. R. Dolling. 1976. Neotropical Miridae, CCV: type designations
of species described in the “Biologia Centrali Americana” (Hemiptera). Rev. Brasileira
Biol. 36:789-810.
Distant, W. L. 1880-1893. Biologia Centrali-Americana. Insecta. Rhynchota. Hemiptera
Heteroptera. Vol. I, pp. 1-302 (1880-1884); suppl., pp. 304-462 (1884-1893). London.
Froeschner, R. C. 1949. Contribution to a synopsis of the Hemiptera of Missouri. Pt. IV.
Am. Midland Naturalist 42:123-188.
Henry, T. J. 1 979. Descriptions and notes on five new species of Miridae from North America.
Melsheimer Entomol. Ser. 27:1-10.
Henry, T. J. 1 982a. New synonymies and a new combination in the North American Miridae
(Hemiptera). Proc. Entomol. Soc. Wash. 84:337-341.
Henry, T. J. 1982b. Genus Parthenicus in the Eastern United States, with descriptions of new
species (Hemiptera: Miridae). Fla. Entomol. 65:354-366.
Kelton, L. A. 1980. The Plant Bugs of the Prairie Provinces of Canada. Agric. Canada Publ.
1703, Ottawa, 408 pp.
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Knight, H. H. 1917. A revision of the genus Lygus as it occurs in America north of Mexico,
with biological data on the species from New York. Cornell Univ. Agric. Exp. Stn. Bull.
391:555-645.
Knight, H. H. 1920. New and little-known species of Phytocoris from the Eastern United
States (Heteroptera-Miridae). Bull. Brooklyn Entomol. Soc. 15:49-66.
Knight, H. H. 1923. Family Miridae (Capsidae). Pages 422-658 in: W. E. Britton (ed.). The
Hemiptera or Sucking Insects of Connecticut. Conn. St. Geol. Nat. Hist. Surv. Bull. 34.
Knight, H. H. 1 925. Descriptions of thirty new species and two new genera of North American
Miridae (Hemiptera). Bull. Brooklyn Entomol. Soc. 20:33-58.
Knight, H. H. 1927a. Notes on the distribution and host plants of some North American
Miridae (Hemiptera). Can. Entomol. 59:34-44.
Knight, H. H. 1927b. On the Miridae in Blatchley’s “Heteroptera of Eastern North America.”
Bull. Brooklyn Entomol. Soc. 22:98-105.
Knight, H. H. 1928. New species of Phytocoris from North America (Hemiptera, Miridae).
Bull. Brooklyn Entomol. Soc. 23:28-46.
Knight, H. H. 1941. The plant bugs, or Miridae, of Illinois. 111. Nat. Hist. Surv. Bull. 22:1-
234.
Knight, H. H. 1968. Taxonomic review: Miridae of the Nevada Test Site and the Western
United States. Brigham Young Univ. Sci. Bull 9(3): 1-282.
Reuter, O. M. 1876. Capsinae ex America Boreali in Museo Holmiensi asservatae, descriptae.
Ofv. Kong. Vet.-Akad. Forh. 32(9):59-92 (1875).
Reuter, O. M. 1 909. Bemerkungen iiber nearktische Capsiden nebst Beschreibung neuer Arten.
Acta Soc. Sci. Fenn. 36(2): 1-86.
Reuter, O. M. 1912. Hemipterologische miscellen. Ofv. Fins. Vet. -Soc. Forh. 65(7): 1-76.
Steyskal, G. C. 1973. The grammar of names in the catalogue of the Miridae (Heteroptera)
of the world by Carvalho, 1957-1960. Studia Entomol. 16:204-208.
Uhler, P. R. 1877. Report upon the insects collected by P. R. Uhler during the explorations
of 1875, including monographs of the families Cydnidae and Saldae and the Hemiptera
collected by A. S. Pachard, Jr., M.D. Bull. U.S. Geol. Surv. 1877 3:355-475; 765-807.
Uhler, P. R. 1878. Notices of the Hemiptera Heteroptera in the collection of the late T. W.
Harris, M. D. Proc. Boston Soc. Nat. Hist. 19:355-446.
Uhler, P. R. 1 894. Observations upon the heteropterous Hemiptera of Lower California, with
descriptions of new species. Proc. Calif. Acad. Sci. 4:223-295.
Uhler, P. R. 1895. A preliminary list of the Hemiptera of Colorado. In Gillette, C. P. and C.
F. Baker. Colo. Agric. Exp. Stn. Bull. 31:1-137.
Van Duzee, E. P. 1910. Descriptions of some new or unfamiliar North American Hemiptera.
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Van Duzee, E. P. 1914. A preliminary list of the Hemiptera of San Diego County, California.
Trans. San Diego Soc. Nat. Hist. 2:1-57.
Van Duzee, E. P. 1916. Check List of the Hemiptera (Excepting the Aphididae, Aleurodidae
and Coccidae) of America, North of Mexico. New York Entomol. Soc., New York,
1 1 1 pp.
Van Duzee, E. P. 1917. Catalogue of the Hemiptera of America north of Mexico excepting
the Aphididae, Coccidae and Aleurodidae. Univ. Calif. Publ. Tech. Bull. 2:1-902.
Van Duzee, E. P. 1 920. New hemipterous insects of the genera Aradus, Phytocoris and Camp
tobrochys. Proc. Calif. Acad. Sci. 9:331-356.
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Species on arborvitae, false cypress, and juniper. Trans. Am. Entomol. Soc. 103:623-
656.
Received August 1, 1983; accepted September 14, 1983.
NEW YORK ENTOMOLOGICAL SOCIETY
91(4), 1983, pp. 466-502
AN ANNOTATED SYNONYMIC LIST OF NORTH AMERICAN AND
CARIBBEAN WASPS OF THE GENUS CERCERIS
(HYMENOPTERA: PHILANTHIDAE)'
George R. Ferguson
Systematic Entomology Laboratory, Department of Entomology,
Oregon State University, Corvallis, Oregon 97331
/I — Results of a study of primary types representing 258 species-group names in the
genus Cerceris are presented: 43 new synonyms are proposed; 14 names are removed from
synonymy and given species status; 1 0 names previously recognized as subspecies are elevated
to species status; one name is removed from synonymy and given subspecies status; three names
are reduced from species to subspecies status; and eight names are transferred from the syn-
onymy of one species to that of another. Cerceris nitidoides Ferguson is proposed as a New
Name for the preoccupied Cerceris nitida Banks.
As part of a study of North American and Caribbean wasps of the genus
Cerceris I have examined primary types representing 258 species-group
names. The resulting new and revised synonymy is presented as an annotated
list. Included species are those known to occur in North America, south to
and including Panama, and the islands of the Caribbean other than Trinidad.
Exclusive of new names, emendations, and misspellings, some 294 species-
group names have been applied to Cerceris occurring within the geographic
limits indicated. Ferguson and Vardy (1983) removed Cerceris serripes (Fa-
bricius) from the North American faunal list, showing that it was a junior
subjective synonym of the palaearctic Cerceris arenaria (Linnaeus).
Of the remaining 293 species-group names, 137 are recognized as valid
species and four as subspecies. Ten names previously treated as subspecies
' Oregon Agricultural Experiment Station Technical Paper No. 6912.
The publication costs of this article were defrayed in part by page charge payment. This article
must therefore be hereby marked “Advertisement” in accordance with 18 U.S.C. §1734 solely
to indicate this fact.
VOLUME 91, NUMBER 4
467
are elevated to species rank, and 1 5 species-group names are removed from
synonymy and recognized as valid species or subspecies. Three taxa are here
reduced from species to subspecies status. Seven species-group names are
transferred from the synonymy of one species to that of another. Twenty-
seven species-group names are synonymized because of conspecificity of
types (10 of these by sex association); and 1 6 species-group names previously
treated as subspecies are synonymized under the nominate species. One new
name, C. nitidoides Ferguson, is proposed for the preoccupied C. nitida
Banks, here elevated to species status.
Color variation. Sexual dimorphism has led to the description of the op-
posite sexes as different species in many cases. However, the plethora of
names is due largely to extensive color variation in widely distributed species
with many color variants having been described as species or as infraspecific
taxa. Bohart and Grissell (1975) synonymized many of the color variants
which had been recognized as subspecies by Scullen (1965a, 1972). Color
variation in most cases is clinal, and species with similar distributions often
show similar geographic patterns of color variation. I attribute this to con-
vergent responses to extrinsic environmental conditions and do not consider
such variants as worthy of subspecies recognition. These color variants have
been given names by previous authors in some cases but not in others. If
one were to be consistent in naming color forms, a large number of additional
names would have to be created. In their Philanthinae of California, Bohart
and Grissell (1975) consistently ignored these variations in shade and extent
of pale maculations as criteria for establishing taxonomic entities. As a
general rule I have followed the same policy in this paper.
One common pattern of variation is the gradual transition in shade of
pale markings from yellow to white in a south to north dine among several
western species (e.g., C. aequalis, C. calochorti, C. nigrescens, C. vanduzeei).
Another pattern (e.g., C. californica, C. sexta) is the development of broad
pale bands on the terga in the southwestern deserts with a gradually increasing
amount of melanism exhibited in both northerly and southerly directions.
Several species widely distributed in eastern North America exhibit a ten-
dency toward a red instead of black background color in Florida and adjacent
southeastern states (e.g., C. blakei, C. JJavofasciata, C. compar,C. rufopicta
and Philanthus sanbornii). Some widely distributed eastern species have
extended their ranges into the southwestern states where they exhibit a clinal
change from yellowish to whitish pale markings and a gradually increasing
amount of pale color on the body (e.g., C compar, C. compacta, C. insolita).
I do not consider it useful to designate these kinds of variation as subspecies.
Other cases of color convergence are apparently due to mimicry. Saussure
(1867) pointed out the resemblance of color pattern of C. simulans to that
of Polybia emaciata Lucas (misidentified by Saussure as Polybia fasciata
468
NEW YORK ENTOMOLOGICAL SOCIETY
according to Richards, 1978). C. militaris and C scapularis have similar
patterns of pale markings. In the northern part of their ranges C azteca, C.
bakeri, C. dilatata, and C. tolteca, each belonging to a different species group,
exhibit broad pale bands on the posterior terga similar to the color pattern
of some social wasps of the genus Brachygastra. In Costa Rica and adjacent
areas C bakeri, C. kennicottii, and C. mexicana exhibit a color pattern
similar to that of Polybia occidentalis bohemani Holmgren, having an almost
completely yellow propodeum with posterior tergal markings reduced. I
divide C. bakeri, C. kennicottii and C. mexicana into subspecies based on
this pattern of mimicry, which may have biological significance.
Differences in color pattern, as opposed to differences in shade of color
or size of maculations within the same pattern, are important taxonomic
characters, but such differences in pattern are almost always associated with
differences in structural morphology.
Format. All species-group names are listed alphabetically whether origi-
nally described as specific or infraspecific taxa. Synonyms are italicized and
placed in parentheses with the name of the valid species under which each
is cited. A generalized summary of geographic distribution is given after
each name. [Abbreviations used are n., s., e., w. and c. for north, south, east,
west and central respectively]. Under each valid name is placed the biblio-
graphic citation to the original description or emendation for each name
included in the synonymy. The original orthography is used in the literature
citation.
An asterisk (*) following the species-group name in the bibliographic ci-
tation indicates that I have studied the type. A double asterisk (**) indicates
that a type comparison has been made by C. R. Vardy of the British Museum
(Natural History) using specimens supplied by me. Insofar as possible I have
placed in brackets the sex, type locality, and location of the type specimen(s)
following each bibliographic citation. If a neotype or lectotype has been
designated, the reference to such designation is given. Immediately following
the brackets I have attempted to give the first author and citation for the
synonymy.
Type repositories. The following abbreviations for type repositories are
used in the text:
AMNH — American Museum of Natural History, New York
ANSP— Academy of Natural Sciences of Philadelphia
Berlin— Zoologisches Museum, Humboldt Universitat, Berlin (DDR)
BMNH — British Museum (Natural History), London
BPBM — Bishop Museum, Honolulu, Hawaii
CAS — California Academy of Sciences, San Francisco
Geneva — Museum d’Histoire Naturelle, Geneva
VOLUME 91, NUMBER 4
469
Genoa— Museo Civico di Storia Naturale, Genoa, Italy
Halle— Zoologisches Institut, Martin Luther Universitat, Halle am Saale,
DDR
Havana— Academia de Ciencias de Cuba, Havana
KANS — University of Kansas, Lawrence
Lund— Universitets Zoologiska Institut, Lund, Sweden
MACN— Museo Argentino de Ciencias Naturales “Bernardino Rivadavia,”
Buenos Aires
Madrid— Museo Nacional de Ciencias Naturales de Madrid
MCZ— Museum of Comparative Zoology, Cambridge, Massachusetts
NCDA— North Carolina State Department of Agriculture, Raleigh
NEB— University of Nebraska State Museum, Lincoln
Torino — Museo ed Istituto Zoologia Sistematica, Torino, Italy
UCD— University of California, Davis
USNM — United States National Museum of Natural History, Washington
Vienna— Naturhistorisches Museum Wien, Vienna
LIST OF NORTH AMERICAN AND CARIBBEAN Cerceris
{abbreviata Banks = nigrescens F. Smith)
acanthophila Cockerell sw. U.S., n. Mexico
Cerceris acanthophilus* Cockerell, 1897:135. [Male lectotype, Deming,
New Mexico, designated by Cresson (1928); ANSP No. 10038].
Cerceris chilopsidis* Viereck and Cockerell, 1904:136. [Female holotype,
Rincon, New Mexico; ANSP No. 10375]. Synonymy by Banks ( 1 947),
Ferguson (1983b).
Ferguson (1983b) reviewed previous misapplications of the name acan-
thophila.
acolhua Saussure Mexico
Cerceris acolhua* Saussure, 1867:90. [Male lectotype, Mexico, designated
by Ferguson (1984); Geneva]
Scullen (1972) left C. acolhua as an unrecognized species. It is allied to,
but distinct from, C. occipitomaculata Packard. The female is unknown,
aequalis Provancher sw. Canada to nw. Mexico
Cerceris aequalis* Provancher, 1888:417. [Female lectotype, California,
designated by Gahan and Rohwer (1917); USNM].
Cerceris aequalis idahoensis* Scullen, 1965a:454. [Female holotype, Craig’s
Mountain, Idaho; ANSP No. 5040]. New Synonymy.
Scullen (1965a) separated idahoensis from the nominate subspecies by
its creamy white versus yellow pale markings. As in some other western
species, these markings vary from yellow to whitish in a south to north
dine.
470
NEW YORK ENTOMOLOGICAL SOCIETY
(affumata Schletterer = intricata F. Smith)
alacris Mickel (Revised Status) sc. U.S.
Cerceris alacris* Mickel, 191 8:334. [Female holotype, Mitchell, Nebraska;
NEB].
Scullen (1965a) incorrectly synonymized alacris under C. halone Banks.
They are distinct species,
alamos Scullen w. Mexico
Cerceris alamos* Scullen, 1972:18. [Female holotype, 10 mi. se. Alamos,
Sonora, Mexico; CAS],
alaope Banks (Revised Status) e. U.S.
Cerceris alaope* Banks, 1912:22. [Male lectotype, Falls Church, Virginia,
designated by Ferguson (1983a); MCZ No. 13784].
Cerceris banksi* Scullen, 1965a:461. [Female holotype. Falls Church, Vir-
ginia; MCZ No. 30477]. New Synonymy.
Synonymy is by sex association. Scullen (1942) incorrectly synonymized
alaope under C. prominens Banks.
(albida Scullen = insolita Cresson)
(albimana Taschenberg = cribrosa Spinola)
(albinota Scullen = compar Cresson)
{alceste Mickel = sexta Say)
{ampla Banks = intricata F. Smith)
(arbuscula Mickel = atramontensis Banks)
(architis Mickel = halone Banks)
arelate Banks (Revised Status) se. Canada, ne. U.S.
Cerceris arelate* Banks, 1912:18. [Female holotype. Great Falls, Virginia;
MCZ No. 13779].
Cerceris nigritulus* Banks, 1915:402. [Male holotype, Colden, New York;
MCZ No. 13782]. Synonymy by Scullen (1965a).
Cerceris crawfordi* Brimley, 1928:199. [Male holotype, Raleigh, North
Carolina; NCDA]. Synonymy by Scullen (1965a).
Scullen (1965a) treated arelate as a yellow marked subspecies of C.
nigrescens F. Smith, and Bohart and Grissell (1975) synonymized the two
forms. C. arelate is distinct from C. nigrescens in having a triangular
median tooth on the apical margin of the clypeal midsection, whereas in
nigrescens this tooth is rectangular.
(argia Mickel = rufinoda Cresson)
{argyrotricha Rohwer = californica Cresson)
(arizonella Banks = vierecki Banks)
{arno Banks = californica Cresson)
astarte Banks se. Canada, e. U.S.
Cerceris astarte* Banks, 1913b:424. [Female lectotype. Falls Church, Vir-
ginia, designated by Ferguson (1983a); MCZ No. 13788].
{athene Banks = femurrubrum Viereck and Cockerell)
VOLUME 91, NUMBER 4
471
(atlacomulca Scullen = flavida Cameron)
(atrafemori Scullen = insolita Cresson)
atramontensis Banks se. Canada, e. U.S.
Cerceris atramontensis* Banks, 1913b:425. [Female holotype. Valley of
Black Mountains, North Carolina; AMNH].
Cerceris arbuscula* Mickel, 1916:410. [Female holotype, Omaha, Ne-
braska; NEB]. Synonymy by Scullen (1951).
Scullen (1965a) incorrectly stated that the type of atramontensis was at
the MCZ.
{atrata Scullen = echo Mickel)
{aureofacialis Cameron = compacta Cresson)
(austrina Fox = rufopicta F. Smith)
azteca Saussure sw. U.S. to Nicaragua
Cerceris azteca* Saussure, 1867:97. [Female lectotype, Cuantla, Mexico,
designated by Ferguson (1984); Vienna].
Cerceris (Apiratirx) [sic] seminigra* Banks, 1947:33, nec Taschenberg 1875.
[Male holotype, Patagonia, Arizona; MCZ No. 27621]. Synonymy by
Scullen (1961).
bakeri bakeri Cameron (Revised Status) El Salvador to Costa Rica
Cerceris bakeri* Cameron, 1904:67. [Female lectotype, Chinandega, Nic-
aragua, designated by Ferguson (1984); BMNH].
bakeri parkeri Scullen (Revised Status) sw. U.S. to Guatemala
Cerceris parkeri* Scullen, 1972:35. [Female holotype, 48 mi. nw.
Tehuantepec, Oaxaca, Mexico; USNM].
Scullen (1972) misidentified bakeri and used the name as a subspecies
of C kennicottii Cresson. However, the type of C. bakeri is conspecific
with C. parkeri. The nominate subspecies has a yellow tergum I and yellow
propodeum with reduced pale markings on the terga. It is apparently a
mimic of Polybia occidentalis bohemani Holmgren. The subspecies par-
keri has tergum I and the propodeum mostly or entirely black and terga
III-V are broadly banded with yellow apparently mimicking Brachygastra
azteca (Saussure), another common polybiine wasp.
(banksi Scullen == alaope Banks)
(beali Scullen = kennicottii Cresson)
(belfragei Banks = compacta Cresson)
{bicornis Ashmead = bicornuta Guerin)
bicornuta Guerin U.S., n. Mexico
Cerceris bicornuta Guerin, 1844:443. [Female holotype. New Oreleans,
Louisiana; Genoa].
Cerceris dufourii Guerin, 1844:443. [Male holotype. New Orleans, Loui-
siana; Genoa]. Synonymy by Cresson (1875).
Cerceris Venator* Cresson, 1865:1 16. [Male lectotype, Illinois, designated
by Cresson (1916); ANSP No. 1937]. Synonymy by Cresson (1875).
472
NEW YORK ENTOMOLOGICAL SOCIETY
Cerceris curvicornis** Cameron, 1890:124. [Male holotype, Mazatlan,
Mexico; BMNH]. Synonymy by Scullen (1961).
Cerceris bicornis Ashmead, 1899:295. Lapsus.
Cerceris fidelis* Viereck and Cockerell, 1 904: 1 32. [Female holotype, Santa
Fe, New Mexico; ANSP No. 10379]. Synonymy by Bohart and Gris-
sell (1975).
Cerceris venatrix SchuXz, 1906:195. Emendation.
{bifida Scullen = rufopicta F. Smith)
(bilinieata Schletterer = triangulata Cresson)
{bilunata Cresson = triangulata Cresson)
binodis Spinola s. Mexico to Argentina
Cerceris binodis"^ Spinola, 1841:117. [Male lectotype, Cayenne, French
Guiana, designated by Ferguson (1984); Torino].
Diamma spinolae Dahlbom, 1844:225. Described in synonymy.
Cerceris viduata** F. Smith, 1856:463. [Female holotype, Para, Brasil;
BMNH]. Synonymy by Fritz (1971).
Cerceris singularis Brethes, 1910:265. [Female holotype, Cordoba, Ar-
gentina; MACN ]. Synonymy by Fritz (1971).
(biungulata Cresson = sexta Say)
(blackii Schletterer = blakei Cresson)
Make! Cresson e. U.S.
Cerceris elegans* F. Smith, 1856:467, nee Eversmann 1849. [Male lec-
totype, St. John’s Bluff, E. Florida, designated by Ferguson (1984);
BMNH].
Cerceris blakei* Cresson, 1865b: 121. [Female holotype, Georgia; ANSP
No. 1947]. Synonymy by Scullen (1961).
Cerceris blackii Schletterer, 1887:487. Lapsus.
Cerceris elegantissima Schletterer, 1887:490. New name for Cerceris ele-
gans F. Smith.
Scullen (1961) was correct in stating that elegantissima was proposed
as a new name for Cerceris elegans F. Smith, but he was later (Scullen
1965a) incorrect in stating that it was proposed as a new name for Eu-
cerceris elegans Cresson.
boharti Scullen sw. U.S., n. Mexico
Cerceris boharti* Scullen, 1965a:466. [Female holotype. Mount Lemmon
Lodge, Santa Catalina Mountains, Arizona; UCD].
bolingeri Scullen (New Status) nw. U.S.
Cerceris aequalis bolingeri* Scullen, 1965a:453. [Female holotype. Hart
Mountain, Jacob’s Cabin, Lake County, Oregon; USNM].
Although described as a subspecies of aequalis, this taxon is a mor-
phologically distinct species.
{bolingeri Scullen, 1972 = bolingeriana Krombein)
bolingeriana Krombein sw. U.S. to c. Mexico
VOLUME 91, NUMBER 4
473
Cerceris bolingeri* Scullen, 1972:72, nec Scullen 1965a. [Female holotype,
40 mi. nw. Gomez Palacio, Durango, Mexico; USNM].
Cerceris bolingeriana Krombein, 1979:1730. New name for Cerceris bol-
ingeri Scullen, 1972.
bothriophora Schletterer sw. U.S. to El Salvador
Cerceris bothriophora* Schletterer, 1887:456. [Male lectotype, Mexico,
designated by Ferguson (1984); Vienna].
Cerceris geniculata* Cameron, 1890:113. [Female lectotype, Cuantla,
Mexico, designated by Ferguson (1984); BMNHj. New Synonymy.
Cerceris feralis* Cameron, 1890:113. [Male lectotype, Cuantla, Mexico,
designated by Ferguson (1984); BMNHj. Revised Synonymy. Syn-
onym of C. geniculata by Scullen (1962).
Cerceris orestes* Banks, 1947:13. [Female lectotype, Patagonia, Arizona,
designated by Ferguson (1983a); MCZ No. 27673]. New Synonymy.
Scullen (1972) left bothriophora as an unrecognized species, and he
treated geniculata {=feralis) and orestes each as a subspecies of C. compar
Cresson.
bradleyi Scullen s. Mexico to Nicaragua
Cerceris bradleyi* Scullen, 1972:73. [Female holotype. La Calera (?), Nic-
aragua; USNM].
bridwelli Scullen sw. U.S., nw. Mexico
Cerceris bridwelli* Scullen, 1965a:361. [Female holotype. Imperial Coun-
ty, California; USNM].
butleri Scullen sw. U.S., n. Mexico
Cerceris butleri* Scullen, 1965a:363. [Female holotype, 30 mi. s. Stafford,
Arizona; USNM].
cacaloapana Scullen se. Mexico
Cerceris cacaloapana* Scullen, 1972:74. [Female holotype, Cacaloapan,
Puebla, Mexico; UCD].
californica Cresson sw. Canada to c. Mexico
Cerceris californica* Cresson, 1 865b: 1 28. [Male holotype, California; ANSP
No. 1953].
ICerceris texensis Saussure, 1867:89. [Female holotype, Texas; type not
located]. Revised Tentative Synonymy.
Cerceris ferruginior* Viereck and Cockerell, 1904:134. [Male holotype,
Deming, New Mexico; ANSP No. 10378]. Synonymy by Scullen
(1960).
Cerceris gar ciana* Viereck and Cockerell, 1904:135. [Male holotype. Las
Cruces, New Mexico; ANSP No. 10380]. Synonymy by Scullen (1960).
Cerceris populorum* Viereck and Cockerell, 1904:135. [Male lectotype,
Albuquerque, New Mexico, designated by Cresson (1928); ANSP No.
10385]. Synonymy by Scullen (1960).
474
NEW YORK ENTOMOLOGICAL SOCIETY
Cerceris argyrotricha* Rohwer, 1908:324. [Female holotype, Las Cruces,
New Mexico; USNM]. Synonymy by Bohart and Grissell (1975).
Cerceris cognata* Mickel, 1916:408. [Female holotype, Worland, Wyo-
ming; NEB]. Synonymy by Scullen (1960).
Cerceris denticularis* Banks, 1917:1 13. [Female lectotype, Umatilla, Or-
egon, designated by Ferguson (1983a); MCZ No. 10028]. Synonymy
by Scullen (1960).
Cerceris interjecta^ Banks, 1919:84. [Male holotype. Lake Point, Utah;
MCZ No. 13766]. Synonymy by Scullen (1960).
Cerceris arno* Banks, 1947:19. [Female lectotype. Mountains near Clare-
mont, California, designated by Ferguson (1983a); MCZ No. 23542].
Synonymy by Bohart and Grissell (1975), synonym of argyrotricha
by Scullen (1972).
Cerceris calodera* Banks, 1947:22. [Male holotype, Jacumba, California;
MCZ No. 27622]. Synonymy by Scullen (1961).
Cerceris illota* Banks, 1947:23. [Male lectotype, Tucson, Arizona, des-
ignated by Ferguson (1983a); MCZ No. 23541]. Synonymy by Scullen
(1960).
C. texensis Saussure was described from a headless female which has
not been located. Scullen (1961) synonymized it under C. frontata Say,
but the original description does not support this placement. The wing
length of 10 mm, red scutum, and mostly polished propodeal enclosure
make it much more likely that Saussure had a specimen of C californica
Cresson, and I provisionally place it in the above synonymy rather than
under C. frontata.
calochorti Rohwer sw. Canada, w. U.S.
Cerceris calochorti"^ Rohwer, 1908:322. [Female holotype, Boulder, Col-
orado; USNM].
Cerceris varians* Mickel, 1918:336. [Female holotype, Donner Lake, Pla-
cer County, California; NEB]. New Synonymy.
Scullen (1965a) separated calochorti and varians by their whitish and
yellowish pale markings respectively. As in several other western species
the pale markings vary from yellow to white in a south to north dine. The
types of both calochorti and varians are the yellow form. I do not consider
the whitish form worthy of subspecies recognition.
{calodera Banks = californica Cresson)
{caridei Holmberg = dilatata Spinola)
(Carolina Banks = compar Cresson)
carrizonensis Banks sw. U.S.
Cerceris carrizonensis* Banks, 1915:403. [Male holotype, Uvalde, Texas;
MCZ No. 13772].
Cerceris poculum* Scullen, 1965a:391. [Female holotype, 23 mi. ne.
Douglas, Arizona; USNM]. New Synonymy.
VOLUME 91, NUMBER 4
475
Synonymy is by sex association. Scullen (1965a) left carrizonensis as an
unrecognized species.
(catamarcensis Schrottky = intricata F. Smith)
(catawba Banks = compar Cresson)
cavagnaroi Scullen El Salvador to Brasil
Cerceris cavagnaroi* Scullen, 1972:75. [Female holotype, Quezaltepeque,
El Salvador; UCD].
cerverae Giner Mari Cuba
Cerceris cerverae Giner Mari, 1941:333. [Male holotype, Havana, Cuba;
Madrid ?].
(chilopsidis Viereck and Cockerell = acanthophila Cockerell)
{chinandegaensis Cameron = kennicottii Cresson)
chiriquensis Cameron (Revised Status) sw. U.S. to Colombia
Cerceris chiriquensis* Cameron, 1890:114. [Female holotype, Volcan de
Chiriqui, Panama; BMNH].
Scullen (1972) treated this taxon as a subspecies of insolita Cresson, but
it is morphologically distinct.
(chisosensis Scullen = dilatata Spinola)
{chrysogastra Schletterer = simulans Saussure)
(chryssipe Banks = clypeata Dahlbom)
(cincta Dahlbom = fumipennis Say)
(cisandina Brethes = intricata F. Smith
{citrina Scullen =finitima Cresson)
(cleomae Rohwer = crucis Viereck and Cockerell)
(clymene Banks = clypeata Dahlbom)
clypeata Dahlbom se. Canada, e. U.S.
Cerceris clypeata* Dahlbom, 1844:221. [Female lectotype, e. South Car-
olina, designated by Ferguson (1984); Lund].
Cerceris imitator* Cresson, 1865b: 125, nec F. Smith 1856. [Male lecto-
type, Illinois, designated by Cresson (1916); ANSP No. 1951]. Syn-
onymy by Scullen (1960).
Cerceris imitatoria Schletterer, 1887:494. New name for Cerceris imitator
Cresson.
Cerceris chryssipe* Banks, 1912:18. [Female lectotype. Falls Church, Vir-
ginia, designated by Ferguson (1983a); MCZ No. 13791]. Synonymy
by Scullen (1951).
Cerceris clymene* Banks, 1912:20. [Female lectotype. Falls Church, Vir-
ginia, designated by Ferguson (1983a); MCZ No. 13789]. Synonymy
by Scullen (1951).
Cerceris zobeide* Brimley, 1 929: 1 94. [Male holotype, Raleigh, North Car-
olina; NCDA]. Synonymy by Scullen (1960).
Cerceris zosma* Brimley, 1929:195. [Female holotype, Raleigh, North
Carolina; NCDA]. Synonymy by Scullen (1960).
476
NEW YORK ENTOMOLOGICAL SOCIETY
(cochise Scullen = cochisi Scullen)
cochisi Scullen sw. U.S., n. Mexico
Cerceris cochisi"^ Scullen, 1965a:416. [Female holotype, Lordsburg to Sil-
ver City, New Mexico; USNM].
Cerceris cochise Scullen, 1972:42. Lapsus.
cockerelli Viereck sw. Canada, w. U.S.
Cerceris cockerelli* Viereck, 1902:731. [Male lectotype. La Jolla, Califor-
nia, designated by Cresson (1928); ANSP No. 10037].
Cerceris minax* Mickel, 1918:339. [Female holotype, Sacramento, Cal-
ifornia; NEB]. Synonymy by Ferguson (1983b).
(cognata Mickel = californica Cresson)
compacta Cresson e. and sw. U.S. to Costa Rica
Cerceris compacta* Cresson, 1865b: 127. [Female lectotype, Illinois, des-
ignated by Cresson (1916); ANSP No. 1940].
Cerceris huastecae Saussure, 1 867: 102. [Female, male syntypes, Tampico,
Mexico; type specimens not located]. New Synonymy.
Cerceris aureo-facialis** Cameron, 1890:112. [Male holotype, Orizaba,
Mexico; BMNH]. Synonymy by Scullen (1972).
Cerceris solidaginis* Rohwer, 1908:323. [Male holotype. Las Cruces, New
Mexico; USNM]. Synonymy by Scullen (1960).
Cerceris belfragei* Banks, 1917:1 14. [Female lectotype, Texas, designated
by Ferguson (1983a); MCZ No. 10029]. Synonymy by Scullen (1960).
Cerceris Costarica Costarica* Scullen, 1972:44. [Female holotype, 3 mi.
nw. Liberia, Costa Rica; USNM]. New Synonymy.
Cerceris Costarica mitla* Scullen, 1972:46. [Female holotype, Mitla, Oa-
xaca, Mexico; USNM]. New Synonymy.
C huastecae is identifiable from the original description and figures.
The color forms on which Scullen (1972) based his two subspecies of
Costarica are part of a clinal north to south variation, and both huastecae
and aureofacialis are older names for the same color form,
compar Cresson e. and sw. U.S., n. Mexico
Cerceris compar* Cresson, 1865b: 126. [Male lectotype, Illinois, designat-
ed by Cresson (1916); ANSP No. 1949].
Cerceris catawba* Banks, 1912:25. [Female lectotype. Southern Pines,
North Carolina, designated by Ferguson (1983a); MCZ No. 13787].
Synonymy by Scullen (1960).
Cerceris jucunda Carolina* Banks, 1912:26. [Male lectotype. Southern
Pines, North Carolina, designated by Ferguson (1983a); MCZ No.
13785]. Synonymy by Scullen (1951).
Cerceris rufa* Scullen, 1965a:449, nec Taschenberg 1875. [Female holo-
type, South Miami, Florida; USNM]. New Synonymy.
Cerceris compar albinota* Scullen, 1972:60. [Female holotype, 2 mi. ne.
Portal, Cochise County, Arizona; USNM]. New Synonymy.
VOLUME 91, NUMBER 4
477
Cerceris rubrata Bohart and Menke, 1976:586. New name for Cerceris
rufa Scullen.
C compar varies clinally from yellow markings on a partially red back-
ground in Florida to whitish markings on a black background in south-
western U.S. and northern Mexico. The color variation is similar to that
of several other species, and I do not consider these minor color forms
worthy of subspecific names.
{complanata Mickel = vanduzeei Banks)
completa Banks California
Cerceris completa* Banks, 1919:83. [Male holotype, Claremont, Califor-
nia; MCZNo. 13767].
Cerceris grandis percna* Scullen, 1965a:415. [Female holotype, Moke-
lumne Hill, Calaveras County, California; USNMj. Synonymy by
Bohart and Grissell (1975).
conifrons Mickel w. U.S., n. Mexico
Cerceris conifrons* Mickel, 1916:410. [Female holotype, Harrison, Ne-
braska; NEB].
{contractu Taschenberg = dilatata Spinola)
convergens Viereck and Cockerell w. U.S., n. Mexico
Cerceris convergens* Viereck and Cockerell, 1904:136. [Female holotype,
Alamogordo, New Mexico; ANSP No. 10376].
Cerceris rinconis* Viereck and Cockerell, 1904:137. [Female holotype,
Rincon, New Mexico; ANSP No. 10386]. Synonymy by Scullen ( 1 942).
Cerceris hesperina* Banks, 1917:115. [Female lectotype, Ainsworth,
Washington, designated by Ferguson (1983a); MCZNo. 10031]. Syn-
onymy by Scullen (1951).
Cerceris pudorosa* Mickel, 1918:338. [Female holotype. Auburn, Cali-
fornia; NEB]. Synonymy by Scullen (1942).
Cerceris snowi* Banks, 1919:84. [Male lectotype, Tucson, Arizona, des-
ignated by Ferguson (1983a); MCZNo. 13764]. Synonymy by Scullen
(1951).
^ cooperi Scullen s. Mexico to Costa Rica
Cerceris cooperi* Scullen, 1972:43. [Female holotype, 6 mi. w. Turrialba,
Costa Rica; USNM].
The holotype female bears the label “NEOTYPE— cooperi
Scullen,” an obvious lapsus in labelling,
cortezi Scullen (New Status) s. Mexico
Cerceris insolita cortezi* Scullen, 1972:64. [Female holotype, Cuernavaca,
Morelos, Mexico; USNM].
Although described as a subspecies, this taxon is a morphologically
distinct species.
{cosmiocephala Cameron = tolteca Saussure)
{Costarica Scullen = compacta Cresson)
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NEW YORK ENTOMOLOGICAL SOCIETY
crandalli Scullen sw. U.S., nw. Mexico
Cerceris crandalli"^ Scullen, 1965a:372. [Female holotype, Tucson, Ari-
zona; USNM].
{cmwfordi Brimley = arelate Banks)
cribrosa Spinola s. Mexico to Brazil
Cerceris cribrosa"^ Spinola, 1841:1 19. [Female lectotype, Cayenne, French
Guiana, designated by Ferguson (1984); Torino].
Cerceris subpetiolata* Saussure, 1867:95. [Female lectotype, Cordova,
Mexico, designated by Ferguson (1984); Vienna]. Synonymy by Scul-
len (1961).
Cerceris pullatus"^ F. Smith, 1873:105. [Female lectotype, Sao Paulo, Bra-
sil, designated by Ferguson (1984); BMNH]. Synonymy by Scullen
(1962).
Cerceris albimana Taschenberg, 1875:395. [Female lectotype, Venezuela,
designated by Scullen (1962); Halle]. Synonymy by Scullen (1962).
crotonella Viereck and Cockerell sw. U.S., n. Mexico
Cerceris crotonella'^ Viereck and Cockerell 1904:139. [Female holotype.
Las Cruces, New Mexico; ANSP].
crucis Viereck and Cockerell (New Status) s. Canada to n. Mexico
Cerceris rufinoda var. crucis'^ Viereck and Cockerell, 1904:139. [Female
lectotype. Las Cruces, New Mexico, designated by Ferguson (1984);
ANSP].
Cerceris cleomae'^ Rohwer, 1908:325. [Male holotype, Denver, Colorado;
USNM]. New Synonymy.
Cerceris flnitima nigroris* Banks, 1912:27. [Male lectotype. Falls Church,
Virginia, designated by Ferguson (1983a); MCZ No. 13786]. Revised
Synonymy. Synonym of C. flnitima Cresson by Scullen (1965a).
Viereck and Cockerell (1904) and subsequent authors have used the
name C. rufinoda for this species. However, the male holotype of C.
rufinoda Cresson is a specimen of the species which has gone under the
name C argia Mickel. C. crucis is the oldest available name for the
rufinoda of authors. C. cleomae and C. nigroris are both males with a
black tergum I. This color form occurs sporadically over most of the range
of the species. Scullen (1 965a) had left cleomae as an unrecognized species,
cubensis Cresson (Revised Status) Cuba; Jamaica
Cerceris zonata* Cresson, 1865a: 156, nee Say 1823. [Female lectotype,
Cuba, designated by Cresson (1916); ANSP No. 1958].
Cerceris cubensis Cresson, 1 865b: 1 23. Created in synonymy as a new name
for C. zonata Cresson.
Cresson (1865b) proposed C. cubensis as a new name for C. zonata
Cresson because of the similarity of the latter name to C. zonalis (!) F.
Smith 1856. The name cubensis was created in synonymy since zonalis
is not a homonym of zonata. However, when Philanthus zonatus Say was
VOLUME 91, NUMBER 4
479
transferred by Schletterer (1887) from Eucerceris to Cerceris, C. zonata
Cresson became a junior secondary homonym of C zonata (Say). Dalla
Torre (1890, 1897) rejected zonata Cresson and used cubensis Cresson as
the next available name. C. zonata Cresson was thereby permanently
rejected, and cubensis is available even though it was created in synonymy.
Cuernavaca Scullen s. Mexico
Cerceris cuernavaca* Scullen, 1972:46. [Female holotype, Cuernavaca,
Morelos, Mexico; USNM].
{curvicornis Cameron = bicornuta Guerin)
{dakotensis Banks = vicina Cresson)
{denticularis Banks = californica Cresson)
dentifrons Cresson se. Canada, e. U.S.
Cerceris dentifrons"^ Cresson, 1865b: 124. [Female lectotype, Illinois, des-
ignated by Cresson (1916); ANSP No. 1942].
deserta Say se. Canada, ne. U.S.
Cerceris deserta* Say, 1824:343. [Male neotype. South Dakota, designated
by Scullen (1965a); MCZ No. 31113].
Cerceris fulvipes* Cresson, 1865b: 126, nec Eversmann 1849. [Female lec-
totype, Delaware, designated by Cresson (1916); ANSP No. 1941].
Synonymy by Scullen (1942).
Cerceris fuhipediculata Schletterer, 1887:492. New name for Cerceris ful-
vipes Cresson.
dilatata Spinola sw. U.S. to Argentina
Cerceris dilatata* Spinola, 1841:1 18. [Female lectotype, Cayenne, French
Guiana, designated by Ferguson (1984); Torino].
Cerceris atriceps* F. Smith, 1856:458. [Female holotype, Para, Brasil;
BMNH]. New Synonymy.
Cerceris maximiliani* Saussure, 1867:94. [Female lectotype, Mexico, des-
ignated by Scullen (1961); Geneva]. Synonymy by Scullen (1961).
Cerceris contract a Taschenberg, 1875:396. [Female lectotype, Brasil, des-
ignated by Scullen (1962); Halle]. Synonymy by Scullen (1962).
Cerceris caridei Holmberg, 1903:478. [Female holotype, Territorium
Pampa Centralis, Argentina; MACN]. Synonymy by Fritz (1971).
Cerceris vigilii Brethes, 1910:270. [Male lectotype, Alta Gracia, Cordoba,
Argentina, designated by Fritz (1970); MACN]. Synonymy by Fritz
(1971).
Cerceris divisa Brethes, 1910:270. [Female holotype. Las Mercedes, Cha-
co, Argentina; MACN]. Synonymy by Fritz (1971).
Cerceris olymponis* Strand, 1910:140. [Female holotype, Asuncion, Par-
aguay; Berlin]. Synonymy by Scullen (1962).
Cerceris semiatra* Banks, 1947:25. [Male holotype, Patagonia, Arizona;
MCZ No. 27620]. Synonymy by Scullen (1961).
Cerceris dilatata chisosensis* Scullen, 1965a:409. [Female holotype, Chi-
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NEW YORK ENTOMOLOGICAL SOCIETY
SOS Mountains, Big Bend National Park, Texas; USNM]. New Syn-
onymy.
The subspecies chisosensis was based on specimens with a partially red
background color on the propodeum and tergum I. These forms occur
with normally colored specimens in the southwestern U.S. and northern
Mexico.
A dark form in eastern Brasil and Argentina having a pale band on
tergum III with the markings on the posterior terga absent or evanescent
may be worthy of subspecies status. C atriceps F. Smith is the oldest
name applicable to this form, but I have not seen sufficient material to
recognize a subspecies at this time.
{dissita Holmberg = intricat a F. Smith)
{divisa Brethes = dilatata Spinola)
dreisbachi Scullen s. Mexico
Cerceris dreisbachi* Scullen, 1972:77. [Female holotype, Cuernavaca, Mo-
relos, Mexico; USNM].
{dufourii Guerin = bicornuta Guerin)
{duisi Scullen = flavotrochanterica Rohwer)
durango Scullen c. Mexico
Cerceris durango* Scullen, 1972:78. [Female holotype, 15 mi. n. Durango,
Durango, Mexico; USNM].
(eburnea Scullen = vanduzeei Banks)
echo Mickel s. Canada to n. Mexico
Cerceris echo* Mickel, 1916:412. [Female holotype, Monroe Canyon, Sioux
County, Nebraska; NEB].
Cerceris echo atrata* Scullen, 1965a:377. [Female holotype, Camden
County, New Jersey; ANSP No. 5039]. New Synonymy.
The subspecies atrata was based on the color form with a black tergum
I. This color form occurs sporadically throughout the range of the species,
and I do not consider it worthy of subspecies recognition.
{elegans F. Smith = blakei Cresson)
{elegantissima Schletterer = blakei Cresson)
{elephantinops Holmberg = intricata F. Smith)
{emmiltosa Scullen = rufopicta F. Smith)
{englehardti Banks = mimica Cresson)
{eriogoni Viereck and Cockerell = kennicottii Cresson)
erythropoda Cameron s. Mexico
Cerceris erythropoda* Cameron, 1890:126. [Female holotype, Cordova,
Mexico; BMNH].
{esau Schletterer = mimica Cresson)
{eurymele Banks = sextoides Banks)
{eustylicida Williams = nigra Ashmead)
evansi Scullen c. Mexico
VOLUME 91, NUMBER 4
481
Cerceris evansi* Scullen, 1972:80. [Female holotype, 3 mi. nw. Cuerna-
vaca, Morelos, Mexico; USNM].
(exsecta F. Smith = imperialis Saussure)
farri Scullen Jamaica
Cerceris farri* Scullen, 1970:199. [Female holotype, 3.5 mi. sw. Mande-
ville, Manchester, Jamaica; USNM].
{fasciola Cresson = occipitomaculata Packard)
femurrubrum Viereck and Cockerell sw. U.S. to c. Mexico
Cerceris femur- rubrum* Viereck and Cockerell, 1904:135. [Male lecto-
type, Albuquerque, New Mexico, designated by Cresson (1928); ANSP
No. 10040].
Cerceris thione* Banks, 1947:18. [Male lectotype, Colton, California, des-
ignated by Ferguson (1983a); MCZ No. 23543]. Synonymy by Scullen
(1951).
Cerceris athene* Banks, 1947:20. [Female holotype, Claremont, Califor-
nia; MCZ No. 23537]. Synonymy by Bohart and Grissell (1975).
Cerceris femurrubra [sic] rossi* Scullen, 1972:56. [Female holotype. Las
Animas, Sierra Laguna, Baja California Sur, Mexico; CAS]. Synon-
ymy by Bohart in Bohart and Menke (1976).
(feralis Cameron = bothriophora Schletterer)
iferruginior Viereck and Cockerell = calif ornica Cresson).
festiva Cresson Cuba
Cerceris festiva Cresson, 1865a: 156. [Male holotype, Cuba; Havana ?].
Cerceris gratiosa* Schletterer, 1887:460. [Male holotype, Cuba; Vienna].
Synonymy by Scullen in Alayo (1968).
{fidelis Viereck and Cockerell = bicornuta Guerin)
finitima Cresson s. Canada to n. Mexico
Cerceris finitima* Cresson, 1865b: 122. [Female holotype, Illinois; ANSP
No. 1948].
Cerceris finitima citrina* Scullen, 1965a:380. [Female holotype. River-
side, California; CAS]. Synonymy by Bohart and Grissell (1975).
ifirma Cresson = verticalis F. Smith)
flavida Cameron s. Mexico
Cerceris flavida* Cameron, 1890:116. [Female lectotype, Cordova, Mex-
ico, designated by Ferguson (1984); BMNH].
Cerceris atlacomulca* Scullen, 1972:68. [Female holotype, Atlacomulca,
Mexico, Mexico; USNM]. New Synonymy.
Scullen (1972) left C. flavida as an unrecognized species, but the lec-
totype is conspecific with C. atlacomulca.
flavocostalis Cresson Cuba
Cerceris flavocostalis Cresson, 1865a: 153. [Female, male syntypes, Cuba;
Havana ?].
flavofasciata H. Smith e. U.S.
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NEW YORK ENTOMOLOGICAL SOCIETY
Cerceris flavofasciata* H. Smith, 1908:364. [Female holotype, Lincoln,
Nebraska; NEB].
Cerceris floridensis* Banks, 1915:403. [Male holotype, Gulfport, Florida;
MCZ No. 13765]. New Synonymy.
Cerceris natallenus* Brimley, 1927:238. [Female holotype. La Grange,
North Carolina; NCDA]. Synonymy by Scullen (1951).
Scullen (1965a) recognized C. floridensis as a red marked subspecies.
The color variation from north to south is clinal and homeochromic with
C rufopicta F. Smith. The black and yellow northern form passes through
a black, yellow and red phase to a black and red form in southern Florida.
(flavomaculata Cameron = otomia Saussure)
flavotrochanterica Rohwer c. Mexico
Cerceris flavotrochanterica"^ Rohwer, 1912:471. [Male holotype. Federal
District, Mexico; USNM].
Cerceris duisi* Scullen, 1972:47. [Female holotype, 10 mi. sw. Mendoza,
Veracruz, Mexico; USNM]. New Synonymy.
Synonymy is by sex association.
{floridensis Banks = flavofasciata H. Smith)
fortin Scullen s. Mexico
Cerceris fortin* Scullen, 1972:47. [Female holotype, Fortin de las Flores,
Veracruz, Mexico; CAS].
I have not identihed the male of this species,
frontata Say sw. U.S., n. Mexico
Cerceris frontata* Say, 1823:80. [Female neotype, 19 mi. e. Lordsburg,
New Mexico, designated by Scullen (1965a); USNM].
Cerceris occidentalis* Saussure, 1867:100. [Female lectotype, Texas, des-
ignated by Scullen (1961); Geneva]. Synonymy by Scullen (1961).
Cerceris raui* Rohwer, 1920:230. [Female holotype, St. Louis, Missouri;
USNM]. Synonymy by Bohart and Grissell (1975).
{fugatrix Mickel == sexta Say)
(fulvipediculata Schletterer = deserta Say)
(fulvipes Cresson = deserta Say)
fumipennis Say se. Canada, e. U.S.
Cerceris fumipennis* Say, 1837:381. [Male neotype, 3 mi. w. Kaskaskia
River, Boulder, Illinois, designated by Scullen (1965a); USNM].
Cerceris cincta* Dahlbom, 1844:204. [Male holotype (headless). North
America; Lund]. Synonymy by Cresson (1865b).
Cerceris unicincta Taschenberg, 1875:397. [Female holotype, Tennessee;
Halle]. Synonymy by Dalla Torre (1897).
gandarai Rohwer c. Mexico
Cerceris gandarai* Rohwer, 1912:470. [Male holotype. Federal District,
Mexico; USNM].
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483
Cerceris micheneri* Scullen, 1972:86. [Female holotype, Matachic, Chi-
huahua, Mexico; AMNH]. Synonymy by Ferguson (1983b).
(gandari Scullen = rohweri Ferguson)
{ganderi Scullen = rohweri Ferguson)
(garciana Viereck and Cockerell = californica Cresson)
(geniculata Cameron = bothriophora Schletterer)
(gnara Cresson = verticalis F. Smith)
gnarina Banks (Revised Status) sc. Canada to sc. U.S.
Cerceris gnarina* Banks, 1913a:237. [Female lectotype, Vinita, Okla-
homa, designated by Ferguson (1983a); AMNH].
Cerceris posticata* Banks, 1916:64. [Male holotype, Jemez Mountains,
New Mexico; MCZ No. 13771]. New Synonymy.
Synonymy is by sex association. C. gnarina is a valid species and not
a subspecies of C. clypeata as proposed by Scullen (1965a).
grandis Banks sw. U.S., nw. Mexico
Cerceris grandis* Banks, 1913b:423. [Female holotype. Fort Yuma, Ar-
izona; AMNH].
{graphica F. Smith = intricata F. Smith)
{gratiosa Schletterer = festiva Cresson)
grisselli Ferguson California
Cerceris grisselli* Ferguson, 1983b:236. [Female holotype, Tanbark Flat,
Los Angeles County, California; UCD].
See Ferguson (1983b) for a discussion of the synonymy of this species,
halone Banks se. Canada, e. U.S.
Cerceris halone* Banks, 1912:24. [Female holotype. Falls Church, Vir-
ginia; MCZ No. 13777].
Cerceris stigmosalis* Banks, 1916:64. [Male holotype, Fargo, North Da-
kota; MCZ No. 13778]. New Synonymy.
Cerceris architis* Mickel, 1916:409. [Female holotype. South Bend, Ne-
braska; NEB]. Synonymy by Scullen (1960).
Cerceris salome* Banks, 1923:21. [Female lectotype, Jones Creek, Lee
County, Virginia, designated by Ferguson (1983a); MCZ No. 14705].
Synonymy by Scullen (1951).
Cerceris shermani* Brimley, 1928:200. [Female holotype, Raleigh, North
Carolina; NCDA]. Synonymy by Scullen (1951).
The holotype male of C. stigmosalis Banks was misidentified by Scullen
(1965a, 1972) as he applied the name to the whitish form of Cerceris sexta
Say.
hatuey Alayo Cuba
Cerceris hatuey Alayo, 1968:14. [Female, male syntypes, Cuba; Ha-
vana ?].
(hebes Cameron = intricata F. Smith)
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NEW YORK ENTOMOLOGICAL SOCIETY
{hesperina Banks = convergens Viereck and Cockerell)
hidalgo Scullen (New Status) c. Mexico
Cercehs calochorti hidalgo"^ Scullen, 1972:75. [Female holotype, 14 mi.
ne. Durango, Mexico; USNM].
Although described as a subspecies, the holotype female of hidalgo
represents a distinct species. At least three species were included by Scullen
in his determinations under the name C. calochorti hidalgo.
huachuca Banks (Revised Status) sw. U.S. to El Salvador
Cerceris (Apiratrix) [sic] huachuca"^ Banks, 1947:29. [Male holotype (tho-
rax and gaster only), Patagonia, Arizona; MCZ No. 27636].
C. huachuca was incorrectly synonymized under C. acanthophila Cock-
erell by Scullen (1942) and subsequent authors. It is a distinct species
closely allied to C. finitima Cresson. Many of Scullen’s determinations
under the name C. finitima vierecki were this species.
{huastecae Saussure = compact a Cresson)
hurdi Scullen sw. U.S. to Nicaragua
Cerceris hurdi* Scullen, 1972:48. [Female holotype, Ahuacatlan, Nayarit,
Mexico; CAS].
(idahoensis Scullen = aequalis Provancher)
{illota Banks = calif ornica Cresson)
(imitator Cresson = clypeata Dahlbom)
(imitatoria Schletterer = clypeata Dahlbom)
imperialis Saussure c. Mexico to Ecuador
Cerceris imperialis* Saussure, 1867:98. [Female lectotype, Mexico, des-
ignated by Scullen (1961); Geneva].
Cerceris exsecta* F. Smith, 1873:410. [Female lectotype, Mexico, desig-
nated by Ferguson (1984); BMNH]. Synonymy by Scullen (1961).
Cerceris pilosa** Cameron, 1890:128. [Male holotype, Guatemala;
BMNH]. Synonymy by Scullen (1961).
insolita Cresson e. and sw. U.S., e. Mexico
Cerceris insolita* Cresson, 1 865b: 1 29. [Male holotype, Illinois; ANSP No.
1954].
Cerceris intractibilis* Mickel, 1916:41 1. [Female holotype. Child’s Point,
Nebraska; NEB]. Synonymy by Scullen (1962).
Cerceris insolita albida* Scullen, 1965a:447. [Female holotype. Las Cruces,
New Mexico; USNM]. New Synonymy.
Cerceris insolita atrafemori* Scullen, 1965a:448. [Female holotype, Phoe-
nix, Arizona; USNM]. New Synonymy.
Scullen (1965a) based his subspecies albida and atrafemori on rather
minor color differences. The pale markings in this species increase clinally
from east to west as in several other species.
(interject a Banks = calif ornica Cresson)
(intractibilis Mickel = insolita Cresson)
VOLUME 91, NUMBER 4
485
intricata intricata F. Smith Venezuela to Argentina
Cerceris intricata"^ F. Smith, 1856:459. [Male lectotype, Santarem, Brasil,
designated by Ferguson (1984); BMNH].
Cerceris simplex* F. Smith, 1856:462. [Female lectotype, Santarem, Bra-
sil, designated by Ferguson (1984); BMNH]. Synonymy by Fritz
(1971).
Cerceris vulpina* F. Smith, 1856:463. [Male lectotype, Brasil, designated
by Ferguson (1984); BMNH]. Synonymy by Fritz (1971).
Cerceris larvata Taschenberg, 1875:391. [Male holotype, Mendoza, Ar-
gentina; Halle]. Synonymy by Fritz (1971).
Cerceris affumata* Schletterer, 1887:455. [Female holotype, Rio de Ja-
neiro, Brasil; Vienna]. Synonymy by Fritz (1971). Synonym of C.
simplex by Scullen (1965b).
Cerceris melanogaster Holmberg, 1903:481. [Female holotype, Territo-
rium Pampa Centralis, Argentina; MNBA]. Synonymy by Fritz (197 1).
Synonym of C. larvata by Fritz (1962).
Cerceris elephantinops Holmberg, 1903:485. [Male holotype, Territorium
Pampa Centralis, Argentina; location of type unknown]. Synonymy
by Fritz (1971). Synonym of C. larvata by Scullen (1965b).
Cerceris elephantinops var. dissita Holmberg, 1903:485. [Male holotype,
Formosa, Argentina; location of type unknown]. Synonymy by Fritz
(1971). Synonym of C larvata by Fritz (1962).
Cerceris catamarcensis Schrottky, 1909:246. [Female holotype?, Cata-
marca, Argentina; location of type unknown]. Synonymy by Fritz
(1971).
Cerceris cisandina Brethes, 1913:122. [Female holotype, Mendoza, Ar-
gentina; location of type unknown]. Synonymy by Fritz (1971).
intricata graphica F. Smith (Revised Status) sw. U.S. to Peru
Cerceris graphica* F. Smith, 1873:410. [Female holotype, Mexico; BMNH].
Cerceris hebes* Cameron, 1890:124. [Male holotype, (thorax and gaster
only), Chilpancingo, Guerrero, Mexico; BMNH]. Synonymy by Bo-
hart and Grissell (1975).
Cerceris macrosticta* Viereck and Cockerell, 1904:133. [Male holotype
New Mexico; ANSP No. 10381]. Synonymy by Bohart and Grissell
(1975).
Cerceris ampla* Banks, 1912:16. [Female lectotype, designated by Fer-
guson (1984); MCZ No. 13769]. Synonym of C. macrosticta by Scul-
len (1942).
This wide ranging species occurs from the southwestern U.S. to Argen-
tina. Color varies from a highly maculated pale form in the U.S. and
Mexico to an almost completely melanic form in southern South America.
Scullen (1965b) divided it into four subspecies under the specific name
C. simplex. His subspecies were (1) simplex simplex (=affumata) for the
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NEW YORK ENTOMOLOGICAL SOCIETY
most melanic form from Brasil; (2) simplex larvata (=elephantinops) from
Brasil and Argentina; (3) simplex graphica from Mexico to Peru; and (4)
simplex macrosticta {=ampla) from northern Mexico and the southwest-
ern U.S.
Fritz (1971) separated intricat a from macrosticta as distinct species on
the basis of characters in the males, but the characters used are inconsistent
when large number of specimens are examined. The apicolateral corners
of the male pygidium are usually bluntly rounded in the north and sub-
spinose in Argentina. This variation is clinal with specimens from Ven-
ezuela, Colombia and Peru showing an intermediate condition.
I recognize two subspecies. The nominate subspecies occurs from Ven-
ezuela to Argentina and is characterized by having the pale bands on the
terga reduced to narrow bands or absent. The subspecies graphica occurs
from the southwestern U.S. to Colombia and Peru with the pale tergal
bands wide, sometimes completely covering the terga.
irene Banks sc. U.S.
Cerceris irene* Banks, 1912:26. [Female holotype, Fedor, Lee County,
Texas; MCZ No. 13781].
{iresinides Rohwer = kennicottii Cresson)
{irwini Scullen = mexicana Saussure)
Isolde Banks (Revised Status) sw. U.S., nw. Mexico
Cerceris isolde* Banks, 1947:24. [Male holotype, Palmerlee, Arizona; MCZ
No. 23540].
Cerceris sandiegensis* Scullen, 1965a:432. [Female holotype, 2 mi. e.
Anza, Riverside County, California; CAS]. New Synonymy.
Scullen (1960, 1965a) misidentified C. isolde and incorrectly synony-
mized it under C californica Cresson.
ijosei Scullen = mexicana Saussure)
jucunda Cresson (Revised Status) e. U.S.
Cerceris jucunda* Cresson, 1872:231. [Male holotype, Texas; USNM].
Cerceris zelica* Banks, 1912:23. [Female lectotype, Fedor, Texas, desig-
nated by Ferguson (1983a); MCZ No. 13773]. New Synonymy.
Scullen (1951) incorrectly synonymized C jucunda under C. compar
Cresson. Synonymy is by sex association,
kennicottii kennicottii Cresson se. Canada to s. Mexico
Cerceris kennicottii* Cresson, 1 865b: 1 28. [Male holotype, Louisiana; ANSP
No. 1952].
Cerceris montivaga* Cameron, 1890:1 19. [Female holotype, Chilpancin-
go, Guerrero, Mexico: BMNH]. Revised Synonymy. Synonym of C.
kennicottii zapoteca Saussure by Scullen (1972).
Cerceris eriogoni* Viereck and Cockerell, 1 904: 1 39. [Male holotype. Drip-
ping Spring, Organ Mountains, New Mexico; ANSP No. 10377]. Syn-
onymy by Scullen (1965a).
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487
Cerceris kennicottii beali* Scullen, 1965a:386. [Female holotype, Scotts-
dale, Arizona; USNM]. Synonymy by Bohart and Grissell (1975).
kennicottii chinandegaensis Cameron (Revised Status) Guatemala to Pan-
ama
Cerceris chinandegaensis* Cameron, 1904:66. [Male lectotype, Chinan-
dega, Nicaragua, designated by Ferguson (1984); BMNH].
Cerceris iresinides* Rohwer, 1914:522. [Male holotype, Gualan, Guate-
mala; USNM]. Revised Synonymy. Synonym of C. kennicottii bakeri
by Scullen (1972).
This species varies from yellowish pale maculations in the northern and
eastern U.S. to whitish pale markings in most of Mexico and parts of the
southwestern U.S. Scullen (1965a) described beali as a subspecies for these
whitish forms and later (1972) incorrectly used zapoteca Saussure as the
subspecies name with beali as a junior synonym. C zapoteca Saussure is
a distinct species and not a subspecies of kennicottii.
From Guatemala to Costa Rica it has an almost completely yellow
propodeum and tergum I, and apparently mimics Polybia occidentalis
bohemani Holmgren in this area as do some other species of Cerceris.
Scullen (1972) applied the name bakeri to this color form, but bakeri is
a distinct species occurring in the same color form in the same area. Scullen
(1972) applied the name smithiana as a subspecies to an intermediate
color form, but smithiana is the female and a junior synonym of zapoteca
Saussure. I here recognize the form with the mostly or entirely yellow
propodeum as a subspecies, but it must take the name kennicottii chi-
nandegaensis Cameron. See also discussions under the names bakeri and
zapoteca.
krombeini Scullen sw. U.S.; nw. Mexico
Cerceris krombeini* Scullen, 1965a:388. [Female holotype. Continental,
Arizona; USNM].
krugi Dewitz Puerto Rico
Cerceris krugi* Dewitz, 1881:200. [Female lectotype, Puerto Rico, des-
ignated by Ferguson (1984); Berlin].
Cerceris margaratella* Rohwer, 1915:248. [Male holotype, Puerto Rico;
USNM]. New Synonymy.
Synonymy is by sex association. Scullen recognized this synonymy in
determinations.
laevigata F. Smith Dominican Republic
Cerceris laevigata* F. Smith, 1856:465. [Female lectotype, Santo Domin-
go, designated by Ferguson (1984); BMNH].
Cerceris levigata Dalla Torre, 1897:466. Emendation.
(larvata Taschenberg = intricat a F. Smith)
{levigata Dalla Torre = laevigata F. Smith)
lutzi Scullen Panama
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NEW YORK ENTOMOLOGICAL SOCIETY
Cerceris lutzi* Scullen, 1972:85. [Female holotype, El Volcan Chiriqui,
Panama; AMNH].
{macrosticta Viereck and Cockerell = intricat a F. Smith)
macswaini Scullen sw. U.S.
Cerceris macswaini* Scullen, 1965a:485. [Female holotype, Imperial
County, California; USNM].
mandibularis Patton e. U.S.
Cerceris mandibularis* Patton, 1880:403. [Female neotype, Glenside,
Pennsylvania, designated by Scullen (1965a); USNM].
(margaretella Rohwer == krugi Dewitz)
{marginata Cameron = marginula Dalla Torre)
marginula Dalla Torre s. Mexico to Panama
Cerceris marginata** Cameron, 1890:117, nee F. Smith 1856. [Female
holotype, Volcan de Irazu, Costa Rica; BMNH].
Cerceris marginula Dalla Torre, 1897:467. New name for Cerceris mar-
ginata Cameron.
{maximiliani Saussure = dilatata Spinola)
(melanogaster Holmberg = intricat a F. Smith)
melanthe Banks sw. U.S., n. Mexico
Cerceris melanthe* Banks, 1947:21. [Female holotype, Apache Canyon,
Santa Catalina Mountains, Arizona; MCZ No. 23539].
mexicana mexicana Saussure s. Mexico to El Salvador
Cerceris mexicana Saussure, 1867:101. [Female, male syntypes, “Agro
Mexicano calido”; types not located].
Cerceris veracruz veracruz* Scullen, 1972:53. [Female holotype, 1 mi. e.
Cordoba, Veracruz, Mexico; CAS]. New Synonymy.
Cerceris veracruz josei* Scullen, 1972:53. [Female holotype, Quezalte-
peque, El Salvador; UCD]. New Synonymy,
mexicana irwini Scullen (Revised Status) Nicaragua, Costa Rica
Cerceris irwini* Scullen, 1972:49. [Female holotype, 7 mi. s.e. Liberia,
Costa Rica; USNM].
C. mexicana is identifiable from Saussure’s description and figures. C.
irwini is conspecific with mexicana, but represents another mimic of Po-
lybia occidentalis bohemani Holmgren, and I recognize it here as a sub-
species. The subspecies josei is somewhat intermediate between the nom-
inate subspecies and irwini, but I regard it as a minor color variant.
(micheneri Scullen = gandarai Rohwer)
militaris Dahlbom Costa Rica to Brasil
Cerceris militaris Dahlbom, 1844:224. [Female holotype, erroneously re-
ported from ‘‘Cap Bonae Spei”; Lund].
Cerceris rufo-nigra Taschenberg, 1875:399. [Female lectotype, here des-
ignated, Rio de Janeiro, Brasil; Halle]. Synonymy by Empey (1980).
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489
Cerceris rufonigra turrialba* Scullen, 1972:51. [Female holotype, Tur-
rialba, Costa Rica; USNM]. New Synonymy.
The subspecies turrialba proposed by Scullen (1972) has the same pat-
tern of maculations as the nominate form; the individual pale marks are
simply larger. Empey (1980) studied the types of both militaris and ru-
fonigra and established their identity. Empey (personal communication)
states that the type specimen bears a lectotype label affixed by Scullen,
but neither he nor I have found any published reference to the designation
of a lectotype. The type specimen is probably a holotype since Taschenberg
(1875) stated the number of specimens at hand when he had more than
one. In order to avoid any future confusion, the above specimen, labelled
as described by Empey (1980) and bearing the lectotype label affixed by
Scullen, is here designated lectotype despite the fact that such designation
may be redundant.
{miltosa Scullen = rufopicta F. Smith)
mimica Cresson sw. U.S. to c. Mexico
Cerceris mimica* Cresson, 1 872:228. [Female lectotype, Texas, designated
by Cresson (1916); ANSP No. 1943].
Cerceris esau* Schletterer, 1887:458. [Female lectotype, Mexico, desig-
nated by Scullen (1961); Vienna]. Synonymy by Scullen (1961).
Cerceris minima Schletterer, 1887:497. Lapsus.
Cerceris englehardti* Banks, 1947:12. [Male holotype, St. John, Arizona;
MCZ No. 27638]. Synonymy by Scullen (1951).
{minima Schletterer = mimica Cresson)
(minax Mickel = cockerel li Viereck)
(mil la Scullen = compact a Cresson)
montealban Scullen c. Mexico
Cerceris montealban* Scullen, 1972:51. [Female holotype, Oaxaca, Oa-
xaca, Mexico; CAS],
montezuma Cameron c. Mexico
Cerceris montezuma* Cameron, 1890:108. [Female lectotype, Mexico,
designated by Ferguson (1984); BMNH].
(montivaga Cameron = kennicottii Cresson)
{morata Cresson = tepaneca Saussure)
morelos Scullen (New Status) s. Mexico
Cerceris flnitima morelos* Scullen, 1972:25. [Female holotype, 14 mi. s.
Cuernavaca, Mexico; USNM].
Although described as a subspecies, I regard this taxon as a distinct
species closely allied to C. flnitima. It was synonymized under C. flnitima
Cresson by Bohart in Bohart and Menke (1976).
(munda Mickel = nigrescens F. Smith)
(nasica Viereck and Cockerell = tepaneca Saussure)
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NEW YORK ENTOMOLOGICAL SOCIETY
{natallena Brimley = Jlavofasciata H. Smith)
neahminax Scullen sw. U.S.
Cerceris neahminax"^ Scullen, 1965a:390. [Female holotype, Santa Fe,
New Mexico; CAS],
nebrascensis H. Smith nc. U.S.
Cerceris nebrascensis* H. Smith, 1908:368. [Female holotype, Glen, Sioux
County, Nebraska; NEB].
nigra Ashmead St. Vincent, British Guiana, Panama
Cerceris nigra* Ashmead, 1900:227. [Female lectotype, St. Vincent, Brit-
ish West Indies, designated by Ferguson (1984); USNM].
Cerceris eustylicida* Williams, 1928:169. [Female holotype, Blairmont,
Berbice County, British Guiana; BPBM]. New Synonymy.
The types of the above two species and a female from Panama [UCD]
present three slightly different shapes of the longitudinal ridge on the apical
portion of the clypeal midsection. They are inseparable in all other re-
spects, and I conclude that they are all conspecific with minor interpo-
pulation variations.
nigrescens F. Smith Alaska, Canada, n. and w. U.S.
Cerceris nigrescens** F. Smith, 1856:466. [Female holotype. Nova Scotia;
BMNH].
Cerceris munda* Mickel, 1918:337. [Female holotype, Sacramento, Cal-
ifornia; NEB]. Synonymy by Bohart and Grissell (1975).
Cerceris abbreviata* Banks, 1919:84. [Male lectotype. Nelson’s, Yakima
River, Washington, designated by Ferguson (1983a); MCZNo. 13794].
Synonymy by Scullen (1951).
(nigritula Banks = arelate Banks)
(nigroris Banks = crucis Viereck and Cockerell)
(nitida Banks = nitidoides Ferguson)
nitidoides Ferguson (New Name) se. Canada, e. U.S.
Cerceris nitida* Banks, 1913b:424, nec Wesmael 1852. [Female lectotype.
Valley of Black Mountains, North Carolina, designated by Ferguson
(1983a); AMNH].
Scullen (1976) incorrectly synonymized this taxon under C melanthe
Banks as the two are distinct species.
{novomexicana Viereck and Cockerell = occipitomaculata Packard)
oaxaca Scullen s. Mexico
Cerceris oaxaca* Scullen, 1972:90. [Female holotype, 12 mi. se. Oaxaca,
Mexico; USNM].
The allotype male and paratype males are not conspecific, but represent
dark specimens of C truncata Cameron,
obregon Scullen nw. Mexico
Cerceris obregon* Scullen, 1972:34. [Female holotype, 38 mi. nw. Ob-
regon, Sonora, Mexico; USNM].
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491
obsoleta Cameron s. Mexico to Colombia
Cerceris obsoleta* Cameron, 1890:1 18. [Female lectotype, Teapa, Tabas-
co, Mexico, designated by Ferguson (1984); BMNH].
(occidentalis Saussure = frontata Say)
occipitomaculata Packard se. Canada, e. U.S.
Cerceris occipitomaculata* Packard, 1866:62. [Male neotype. Hunt Coun-
ty, Texas, designated by Scullen (1965a); USNM].
Cerceris fasciola* Cresson, 1872:230. [Male holotype, Comal County,
Texas; ANSP No. 1950]. Synonymy by Scullen (1965a).
Cerceris novomexicana* Viereck and Cockerell, 1904:137. [Female ho-
lotype, Johnson Park, New Mexico; ANSP No. 10383]. Synonymy
by Scullen (1965a).
{olymponis Strand = dilatata Spinola)
(orestes Banks = bothriophora Schletterer)
(orphne Banks = sexta Say)
otomia Saussure (Revised Status) s. Mexico to Colombia
Cerceris otomia* Saussure, 1867:99. [Female lectotype, Mexico, desig-
nated by Ferguson (1984); Vienna].
Cerceris flavo-maculata* Cameron, 1890:1 15. [Female lectotype, Rio Su-
sio, Costa Rica, designated by Ferguson (1984); BMNH]. New Syn-
onymy.
Cerceris otomita Dalla Torre, 1897:469. Lapsus.
The apical teeth of the clypeal margin of C.flavomaculata are somewhat
larger than in C. otomia, but I have seen specimens intermediate between
the two and have concluded that they are conspecific.
{otomita Dalla Torre = otomia Saussure)
panama Scullen (New Status) Honduras to Panama
Cerceris insolita panama* Scullen, 1972:67. [Female holotype, 7 mi. se.
Liberia, Costa Rica; USNM].
The female of this species is much like C. bothriophora and the males
are close to C. chiriquensis. It is certainly not C. insolita, and I provi-
sionally elevate the taxon to specific status until this species group can be
more carefully studied.
(parkeri Scullen = bakeri Cameron)
{percna Scullen = completa Banks)
{pilosa Cameron = imperialis Saussure)
{platyrhina Viereck and Cockerell = tepaneca Saussure)
(pleuralis H. Smith = rufopicta F. Smith)
{poculum Scullen = carrizonensis Banks)
{populorum Viereck and Cockerell = californica Cresson)
{posticata Banks = gnarina Banks)
prominens Banks (Revised Status) e. U.S.
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NEW YORK ENTOMOLOGICAL SOCIETY
Cerceris prominens* Banks, 1912:19. [Female lectotype, Falls Church,
Virginia, designated by Ferguson (1983a); MCZ No. 13790].
Scullen (1965a) treated this taxon as a subspecies of C. clypeata, but I
regard it as a distinct species based on the structure of the male clypeus.
psamathe Banks (Revised Status) e. U.S.
Cerceris psamathe* Banks, 1912:21. [Female holotype, Fedor, Lee Coun-
ty, Texas; MCZ No. 13780].
Scullen (1951) incorrectly synonymized this species under C. aequalis,
but they are distinct species.
(pudorosa Mickel = convergens Viereck and Cockerell)
{pullata F. Smith = cribrosa Spinola)
queretaro Scullen sw. U.S. to c. Mexico
Cerceris queretaro* Scullen, 1972:91. [Female holotype, 41 mi. n. Que-
retaro, Mexico; USNM].
{raui Rohwer = frontata Say)
rhois Rohwer sw. U.S.
Cerceris rhois* Rohwer, 1908:325. [Male holotype, Rio Ruidoso, White
Mountains, New Mexico; USNM].
This species is very close to C. halone Banks, and it may prove to be
a senior synonym. C. rhois has broader pale bands on the posterior terga
than C. halone, and the tergal punctures tend to coalesce. In C. halone
the punctures are separated by flat ridges.
(rinconis Viereck and Cockerell = convergens Viereck and Cockerell)
{robertsonii Fox = rufopicta F. Smith)
rohweri Ferguson se. Texas to s. Mexico
Cerceris rohweri* Ferguson, 1983b:238. [Female holotype, 17 mi. ne. San
Luis Potosi, San Luis Potosi, Mexico; USNM].
Cerceris ganderi Scullen, 1968:156. Lapsus.
Cerceris gandari Scullen, 1972:82. Lapsus.
As discussed by Ferguson (1983b) this species was misidentified as C
gandarai Rohwer by Scullen (1968, 1972).
{rossi Scullen = fumurrubrum Viereck and Cockerell)
{rostrata F. Smith = sirnulans Saussure)
rozeni Scullen New Jersey, Florida
Cerceris rozeni* Scullen, 1971:1 30. [Female holotype. Archbold Biological
Station, Highlands County, Florida; AMNH].
{rubrata Bohart and Menke = compar Cresson)
{rufa Scullen = compar Cresson)
rufinoda Cresson c. U.S. to c. Mexico
Cerceris rufinoda* Cresson, 1865b: 121. [Male holotype, Colorado; ANSP
No. 1955].
Cerceris argia* Mickel, 1916:412. [Female holotype, Lincoln, Nebraska;
NEB]. New Synonymy.
This species was discussed under C. crucis.
VOLUME 91, NUMBER 4
493
{rufonigra Taschenberg = militaris Dahlbom)
rufopicta F. Smith se. Canada, e. U.S.
Cerceris rufo-picta"^ F. Smith, 1856:467. [Male lectotype, St. John’s Bluff,
E. Florida, designated by Ferguson (1984); BMNH].
Cerceris robertsonii* Fox, 1893a:55. [Female lectotype, Smithville, South
Dakota, designated by Cresson (1928); ANSP No. 4755]. New Syn-
onymy.
Cerceris austrina* Fox, 1893b:556. [Female lectotype. Southern Florida,
designated by Cresson (1928); ANSP No. 4756]. Revised Synonymy.
Synonym of C. robertsonii by Scullen (1965a).
Cerceris pleuralis* H. Smith, 1908:366. [Female holotype. Rock County,
Nebraska; NEB]. Revised Synonymy. Synonym of C. robertsonii by
Scullen (1965a).
Cerceris robertsonii emmiltosus* Scullen, 1964:144. [Female holotype,
Miami, Florida; USNM]. New Synonymy.
Cerceris robertsonii bifidus* Scullen, 1965a:428. [Female holotype. Kill
Devil Hills, North Carolina; USNM]. New Synonymy.
Cerceris robertsonii miltosus* Scullen, 1965a:429. [Female holotype, Ar-
cadia, Florida; USNM]. New Synonymy.
This species is variable both in color and in the shape of the membrane
appended to the clypeal process of the female. Color varies clinally from
north to south from a black and yellow form in the north to a black, yellow
and red form in the southeastern United States to a black and red form
in southern Florida. The names robertsonii, pleuralis, and bifida apply to
the black and yellow form, and the remaining names apply to forms with
variable amounts of red. The clypeal membrane of the female varies from
the "'robertsoniC shape to the ^^bifida’’" shape within the same population
with all intermediate conditions present. (See Scullen 1965a, figs. 136a
and 137a.)
{salome Banks = halone Banks)
{sandiegensis Scullen = isolde Banks)
{sayi Banks = sexta Say)
scapularis Schletterer (Revised Status) s. Mexico
Cerceris scapularis"^ Schletterer, 1887:457. [Female lectotype, Orizaba,
Mexico, designated by Scullen (1972); Vienna].
Scullen (1972) incorrectly synonymized this species under C. simulans
Saussure. See discussion under C simulans.
(semiatra Banks = dilatata Spinola)
(seminigra Banks = azteca Saussure)
semipetiolata Saussure s. Mexico to Panama
Cerceris semipetiolata Saussure, 1867:88. [Male holotype, Orizaba, Mex-
ico; type not located].
Cerceris williamsi* Scullen, 1972:54. [Female holotype, Fortin de las Flo-
res, Veracruz, Mexico; CAS]. New Synonymy.
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NEW YORK ENTOMOLOGICAL SOCIETY
Synonymy is by sex association. Although Saussure described semipe-
tiolata from a headless male, identification is possible from his description.
The combination of long tergum I, coarsely ridged propodeal enclosure,
and color pattern of the thorax and gaster is distinctive,
sexta Say sc. Canada to c. Mexico
Cerceris sexta* Say, 1837:382. [Male neotype, Colorado, designated by
Scullen (1965a); ANSP No. 5041].
Cerceris biungulata* Cresson, 1865b: 118. [Female holotype, Colorado;
ANSP No. 1956]. Synonymy by Scullen (1960).
Cerceris alceste* Mickel, 1918:333. [Female holotype, Mitchell, Nebraska;
NEB]. New Synonymy.
Cerceris fugatrix* Mickel, 1918:335. [Male holotype, Mitchell, Nebraska;
NEB]. Revised Synonymy. Synonym of C stigmosalis Banks by Scul-
len (1965a).
Cerceris sayi* Banks, 1923:21. [Female lectotype, Steele, North Dakota,
designated by Ferguson (1983a); MCZ No. 14706]. Revised Synon-
ymy. Synonym of C stigmosalis Banks by Scullen (1965a).
Cerceris stevensi* Banks, 1923:22. [Female holotype, Steele, North Da-
kota; MCZ No. 14707]. Revised Synonymy. Synonym of C. stigmo-
salis Banks by Scullen (1965a).
Cerceris orphne* Banks, 1947:15. [Male holotype, Jemez Springs, New
Mexico; MCZ No. 23536]. Synonymy by Scullen (1961).
There is much variation in the size and shape of the female clypeal
process in this species. Pale markings vary from whitish in the northern
plains and Canada to a deep yellow in the southern plains and again whitish
in south central Mexico. Scullen (1965a, 1972) placed the whitish forms
under the name '"stigmosalisC but the type of stigmosalis Banks is a male
of C. halone Banks. The names C. sayi and C stevensi apply to the
northern whitish form,
sextoides Banks sw. Canada to nw. Mexico
Cerceris sextoides* Banks, 1 947: 1 0. [Female holotype. Lone Tree, Yakima
River, Washington; MCZ No. 23547].
Cerceris eurymele* Banks, 1947:1 1. [Female holotype, Davis, California;
MCZ No. 23546]. Synonymy by Scullen (1965a).
(shermani Brimley = halone Banks)
{simplex F. Smith = intricata F. Smith)
simulans Saussure s. Mexico
Cerceris simulans Saussure, 1867:87. [Male holotype, Mexico; type not
located].
Cerceris rostrata** F. Smith, 1873b:409. [Female holotype, Mexico;
BMNH]. New Synonymy.
Cerceris chrysogastra* Schletterer, 1887:458. [Male lectotype, Orizaba,
VOLUME 91, NUMBER 4
495
Mexico, designated by Scullen (1972); Vienna]. Synonymy by Scullen
(1972).
C. simulans is identifiable from Saussure’s original description and fig-
ures. Scullen (1972) identified the male, but he misassociated the female,
as he treated C. rostrata as a separate species and incorrectly placed C.
scapularis in synonymy as the female of simulans. C. rostrata shares the
long tergum I, the basal platform on sternum II, and the fossette on tergum
I with C simulans, whereas these characters are lacking in C scapularis.
(singularis Brethes = binodis Spinola)
{smithiana Cameron = zapoteca Saussure)
(snowi Banks = convergens Viereck and Cockerell)
(solidaginis Rohwer = compacta Cresson)
(spinolae Dahlbom = binodis Spinola)
squamulifera Mickel c. U.S.
Cerceris squamulifera* Mickel, 1916:411. [Female holotype. Imperial,
Nebraska; NEB].
This species is very close to C. rufopicta. However, the males which I
have associated with it lack the sternal fimbriae present in rufopicta.
{stevensi Banks = sexta Say)
(stigmosalis Banks = halone Banks)
strigosa Cameron c. Mexico
Cerceris strigosa* Cameron, 1890:1 10. [Female lectotype, Durango, Mex-
ico, designated by Ferguson (1984); BMNH].
(subpetiolata Saussure = cribrosa Spinola)
tepaneca Saussure sw. U.S. to s. Mexico
Cerceris tepaneca* Saussure, 1867:90. [Male lectotype, Orizaba, Mexico,
designated by Scullen (1972); Vienna].
Cerceris morata* Cresson, 1872:230. [Female lectotype, Texas, designated
by Cresson (1916); ANSP No. 1944]. New Synonymy.
Cerceris thermophila* Schletterer, 1887:463. [Female lectotype, Mexico,
designated by Scullen (1972); Vienna]. New Synonymy. (Listed as
both a synonym of C. clypeata tepaneca and as a good species by
Bohart and Menke, 1976.)
Cerceris nasica* Viereck and Cockerell, 1 904: 1 32. [Female holotype. New
Mexico; ANSP No. 10382]. Revised Synonymy. Synonym of C mor-
ata Cresson by Scullen (1942).
Cerceris platyrhina* Viereck and Cockerell, 1904:133. [Female holotype,
Fillmore Canyon, Organ Mountains, New Mexico; ANSP No. 10384].
Revised Synonymy. Synonym of C. morata Cresson by Scullen (1 942),
synonym of C. vicina Cresson by Scullen (1965a).
Scullen (1942) correctly placed C. platyrhina in synonymy under C.
morata, but he later (1965a) incorrectly placed it in synonymy under C
496
NEW YORK ENTOMOLOGICAL SOCIETY
vicina Cresson. C. tepaneca is very close to C clypeata, and Scullen (1972)
treated it as a subspecies of the latter. However, he regarded C morata
to be a distinct species, and he separated the two by the distinctive red
and yellow coloration of C morata in the northern part of its range. All
intergrades occur in a north to south cline from the red and yellow forms
of the southwestern United States to the black forms with reduced yellow
markings of southern Mexico. I separate C. tepaneca from C. clypeata on
the basis of body sculpture,
texana Scullen Texas
Cerceris texana"^ Scullen, 1965a:511. [Female holotype, Del Rio, Texas;
KANS].
{texensis Saussure = californica Cresson)
(thermophila Schletterer = tepaneca Saussure)
{thione Banks = femurrubrum Viereck and Cockerell)
tolteca Saussure s. U.S. to Panama
Cerceris tolteca* Saussure, 1867:94. [Female lectotype, Cuantla, Mexico,
designated by Scullen (1972); Geneva].
Cerceris cosmiocephala** Cameron, 1904:67. [Male holotype, San Mar-
cos, Nicaragua; BMNH]. Synonymy by Scullen (1961).
townsendi Viereck and Cockerell New Mexico
Cerceris townsendi* Viereck and Cockerell, 1904: 140. [Male holotype. Las
Cruces, New Mexico; ANSP No. 10389].
Ferguson (1984) discussed the identity of the holotype.
triangulata Cresson Cuba
Cerceris triangulata Cresson, 1865a: 154. [Female holotype, Cuba; Ha-
vana?].
Cerceris bilunata* Cresson, 1865a: 155. [Male lectotype, Cuba, designated
by Cresson (1916); ANSP No. 1957]. Synonymy by Alayo (1968).
Cerceris bilinieata Schletterer, 1887:487. Lapsus.
trichiosoma Cameron c. Mexico
Cerceris trichiosoma* Cameron, 1890:127. [Male holotype, Durango,
Mexico; BMNH].
trinitaria Alayo Cuba
Cerceris trinitaria Alayo, 1968:10. [Female, male syntypes, Cuba; Ha-
vana?].
I have not seen this species,
truncata Cameron sw. U.S. to Costa Rica
Cerceris truncata* Cameron, 1890:121. [Female lectotype, Temax, North
Yucatan, Mexico, designated by Ferguson (1984); BMNH].
(turrialba Scullen = militaris Dahlbom)
(unicincta Taschenberg = fumipennis Say)
vanduzeei Banks sw. Canada to nw. Mexico
Cerceris vanduzeei* Banks, 1917:114. [Female lectotype, San Diego Coun-
ty, California, designated by Ferguson (1983a); MCZ No. 10030].
VOLUME 91, NUMBER 4
497
Cerceris complanata* Mickel, 1918:340. [Female holotype, Auburn, Cal-
ifornia; NEB]. Synonymy by Scullen (1960).
Cerceris vanduzeei eburnea* Scullen, 1965a:399. [Female holotype. North
Powder, Union County, Oregon; USNMj. Synonymy by Bohart and
Grissell (1975).
{varians Mickel = calochorti Rohwer)
(Venator Cresson = bicornuta Guerin)
(venatrix Schulz = bicornuta Guerin)
(veracruz Scullen = mexicana Saussure)
verticalis F. Smith se. U.S., ne. Mexico
Cerceris verticalis** F. Smith, 1856:466. [Female holotype, Georgia;
BMNHj.
Cerceris gnara* Cresson, 1872:229. [Male lectotype, Texas, designated by
Cresson (1916); ANSP No. 1938]. Synonymy by Scullen (1961).
Cerceris firma* Cresson, 1872:229. [Female lectotype, Texas, designated
by Cresson (1916); ANSP No. 1945]. Synonymy by Scullen (1961).
vicina Cresson nc. U.S.
Cerceris vicina* Cresson, 1865b: 120. [Female lectotype, Colorado, des-
ignated by Cresson (1916); ANSPNo. 1939].
Cerceris dakotensis* Banks, 1915:402. [Female lectotype, Fargo, North
Dakota, designated by Ferguson (1983a); MCZ No. 13770]. New Syn-
onymy.
Scullen (1965a) treated dakotensis as a subspecies of clypeata, but the
type of dakotensis is clearly conspecific with vicina Cresson. The shape of
the clypeal process is quite distinctive. In his determinations under the
name C clypeata dakotensis Scullen included specimens of clypeata and
prominens as well as vicina. The range of vicina is allopatric with that of
C tepaneca, and I am able to separate males of vicina from males of
tepaneca only on the basis of locality even though the females are quite
distinct.
vicinoides Viereck and Cockerell (Revised Status) sc. U.S.
Cerceris vicinoides* Viereck and Cockerell, 1904:140. [Female holotype,
Pecos, New Mexico; ANSP No. 10387].
Scullen (1951) incorrectly synonymized this species under C aequalis
Provancher, but it is a distinct species.
(viduata F. Smith = binodis Spinola)
vierecki Banks (Revised Status) sw. U.S. to c. Mexico
Cerceris (Apiratrix) [sic] vierecki* Banks, 1947:30. [Female holotype,
Tempe, Arizona; MCZ No. 23544].
Cerceris (Apiratrix) [sic] arizonella* Banks, 1947:32. [Male holotype,
Tempe, Arizona; MCZ No. 23538]. New Synonymy.
Synonymy is by sex association. Scullen (1965a) left C arizonella as an
unrecognized species, and treated vierecki as a whitish subspecies of C
finitima. C. huachuca is a closely related whitish species, and C. finitima
498
NEW YORK ENTOMOLOGICAL SOCIETY
occurs in a whitish phase also. A high percentage of Scullen’s determi-
nations under the name vierecki have been found to be huachuca. Bohart
and Grissell (1975) synonymized vierecki under finitima, but I regard the
two as distinct species.
{vigilii Brethes = dilatata Spinola)
(vulpina F. Smith = intricat a F. Smith)
watlingensis Elliott and Salbert San Salvador, Bahamas
Cerceris watlingensis"^ Elliott and Salbert, 1979:359. [Female holotype,
San Salvador Island, Bahamas; USNM].
{williamsi Scullen = semipetiolata Saussure)
wyomingensis Scullen c. U.S.
Cerceris wyomingensis"^^ Scullen, 1965a:519. [Female holotype, 28 mi. e.
Laramie, Wyoming; USNM].
Zacatecas Scullen c. Mexico
Cerceris zacatecas"^ Scullen, 1972:98. [Female holotype, 9 mi. n. Ojo Cal-
iente, Zacatecas, Mexico; UCD].
zapoteca Saussure (Revised Status) s. Mexico to Costa Rica
Cerceris zapoteca"^ Saussure, 1867:89. [Male lectotype, Cordova, Mexieo,
designated by Ferguson (1984); Vienna].
Cerceris smithiana"^ Cameron, 1890:1 19. [Female lectotype, Atoyac, Ve-
racruz, Mexico, designated by Ferguson (1984); BMNH]. New Syn-
onymy.
Scullen (1972) treated zapoteca and smithiana each as a subspecies of
C. kennicottii Cresson based on color characters, but they represent op-
posite sexes of a distinct species. Most of Scullen’s determinations under
these names are C. kennicottii.
(zelica Banks = jucunda Cresson)
{zobeide Brimley = clypeata Dahlbom)
{zonata Cresson = cubensis Cresson)
{zosma Brimley = clypeata Dahlbom)
zumpango Scullen s. Mexico
Cerceris zumpango"^ Scullen, 1972:38. [Female holotype, Zumpango,
Guerrero, Mexico; UCD].
ACKNOWLEDGMENTS
I am indebted to the following individuals and institutions for the generous loan of type
material and to the individuals named for courtesies and facilities provided during visits to
ANSP, CAS, UCD, and USNM: D. Azuma, ANSP; M. Favreau, AMNH; F. Koch, Berlin; G.
M. Nishida, BPBM; M. C. Day, C. R. Vardy, BMNH; R. M. Bohart, R. O. Schuster, UCD;
W. J. Pulawski, CAS; Cl. Besuchet, Geneva; G. W. Byers, KANS; R. Danielsson, Lund; M.
Hathaway, K. Jepson, C. Vogt, MCZ; K. Ahlstrom, NCDA; B. C. Ratcliffe, NEB; A. S. Menke,
USNM; P. Passerin de’Entreves, Torino; M. Fischer, Vienna.
I am especially grateful to Mary Hathaway, formerly of MCZ, for searching out the Banks
type material and for providing helpful background information; to C. R. Vardy for searching
VOLUME 91, NUMBER 4
499
out type material at the BMNH and for much helpful information as well as making several
type comparisons; to A. S. Menke, USNM, for assistance in obtaining certain literature and
for valuable comments on some nomenclatorial questions; and to R. M. Bohart, UCD, for
providing some literature and for reviewing an early draft of this manuscript.
I thank J. D. Lattin, Curator, Systematic Entomology Laboratory, Oregon State University,
for making available the extensive collection of cercerine wasps built up by Dr. H. A. Scullen
over a period of almost 40 years and for other courtesies and facilities extended. I thank my
colleagues P. W. Oman and W. P. Stephen for helpful comments and discussions concerning
a number of systematic and nomenclatorial questions, and for constructive reviews of the
manuscript.
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Received August 17, 1983; accepted September 30, 1983.
NEW YORK ENTOMOLOGICAL SOCIETY
91(4), 1983, pp. 503-507
RECOGNITION OF HOST NEST ODOUR BY THE
BUMBLEBEE SOCIAL PARASITE PSITHYRUS ASHTONI
(HYMENOPTERA: APIDAE)
Richard M. Fisher
Biology Department, Erindale College,
Mississauga, Ontario, Canada L5L 1C6
Abstract.— T\iQ ability of the social parasite Psithyrus ashtoni to identify and discriminate
between nest odours of different bumblebee species was investigated. Female P. ashtoni were
presented with various combinations of host nest, non-host nest and nest material alone.
Parasites showed preference for the odour of the host nests rather than nest material or nests
of non-host bumblebees. The importance of host nest odour identification to searching parasites
is discussed.
Bees belonging to the genus Psithyrus are obligate brood parasites of bum-
blebees (Hymenoptera: Apidae). Female Psithyrus emerge from hibernation
some time after host bumblebee queens and search for established nests,
most of which are underground in abandoned rodent burrows (Alford, 1975).
How are host nests located? Early investigators of Psithyrus biology (e.g.,
Sladen, 1912; Plath, 1 934) believed that the scent of host nests were detected
by searching parasites. More recently, Cederberg (1979) suggested that Psi-
thyrus rupestris Fab. follows odour trails produced by host Bombus lapi-
darious Linn, workers. These trails are made by workers as they wander in
and out of subterranean burrows leading to the nest (Cederberg, 1977).
Worker extract daubed on filter paper is followed by P. rupestris females.
P. ashtoni Cr. is parasitic in nests of Bombus affinis Cr. and B. terricola
Kby. (Plath, 1922). It searches for nests within one-two weeks of host queen
emergence (Fisher, unpubl.), at a time prior to emergence of the first worker
brood, and therefore cannot use odour trails of workers to recognize nests.
Since bumblebee queens incubating incipient brood clumps leave the nest
at irregular intervals to forage (Alford, 1975), their nests may be difficult to
locate if odour trails of queens are being used. In addition, P. ashtoni is
found only in nests of B. terricola and B. affinis, unlike many other Psithyrus
species which are not host specific. Can P. ashtoni identify host nests by
odour without relying on worker trails, and can it discriminate host from
non-host bumblebee species? Investigation of the ability of P. ashtoni fe-
The publication costs of this article were defrayed in part by page charge payment. This article
must therefore be hereby marked “Advertisement” in accordance with 18 U.S.C. §1734 solely
to indicate this fact.
504
NEW YORK ENTOMOLOGICAL SOCIETY
males to identify and discriminate between odours of different bumblebee
species is the subject of this paper.
METHODS AND MATERIALS
Bumblebees were reared in the laboratory using the techniques of Plow-
right and Jay (1966) and Pomeroy and Plowright (1980) during May 1982.
Female P. ashtoni were caught as they searched for nests and maintained in
small wire cages with access to pollen and sugar-water solution.
Each bumblebee nest starting unit consists of a 17.5 X 12.5 X 10 cm outer
box with glass lid where queens collect sugar water and void faeces. The
outer box is connected by a 1cm diameter hole to an inner box (ID 77 X
77 X 50 mm) with closed glass lid. The inner box contains upholsterer’s
cotton and a moist pollen lump in which captive queens lay eggs. Each inner
box is detachable and easily moved about once the entrance hole is plugged.
I chose representative nests of B. terricola, B. affinis, and a non-host species
{B. bimacidatus Cr.) for this experiment; all were at the same stage of de-
velopment (hrst brood workers in cocoon stage, second brood eggs laid).
This is the approximate stage of nest development when P. ashtoni is nest
searching. I plugged and removed boxes containing nests and queens and
placed them in a darkroom cubicle in another room. The glass lid from each
box was removed and replaced with wire mesh screen stapled in place.
A plexiglass cage which exactly ht the outer dimensions of two nest boxes
placed side by side was constructed. This cage had a removable plexiglass
lid and detachable wire mesh floor. The cage was supported off the nest
boxes by 4 mm strips of plexiglass so that no part of the cage floor came in
contact with the wire mesh tops of the nest boxes.
Boxes containing nests of the three bumblebee species or upholsterer’s
cotton alone were placed side by side in various combinations. A female
Psithyrus was brought into the darkroom and placed in the plexiglass cage.
It was allowed one minute to wander around the cage. A single 40 W red
light illuminated the apparatus from a central location. The midline of the
cage floor had been marked; when the female Psithyrus was in the middle
of the cage it was lowered onto the nest boxes. The proportion of time each
female Psithyrus spent on either side of the midline was then monitored for
a five-minute period. Following the five-minute experimental period the
mesh floor of the plexiglass cage was removed and replaced with another
clean screen floor. Nest boxes were replaced with other combinations and
the procedure repeated.
The following nest box pairings were presented to ten different Psithyrus
females: cotton only-cotton only, cotton only-^. terricola nest, cotton only-
B. qtfinis nest, B. qffinis nest-i?. terricola nest, host nest-R bimacidatus nest.
The order in which these pairings were presented was determined randomly,
except for the host nest-/?, bimacidatus nest pairing, which was always
VOLUME 91, NUMBER 4
505
Table 1. Percent time spent by Psithyrus ashtoni females in either half of plexiglass cage
covering nest pairs (N = 10).
Cotton Cotton
only only
tern-
cola Cotton
nest only
affinis Cotton
nest only
teni-
affinis cola
nest nest
bimacu-
Host latus
nest nest
% time spent
in cage-half
47.20 52.80
78.28 21.72
78.93 21.07
49.78 50.22
73.57 26.43
0.470
47.997
50.228
0.003
37.332
p
NS'
<0.001
<0.001
NS'
<0.001
’ Not significant at 0.05 level.
presented last. Left-right box pairings were also randomized. Each female
Psithyrus was tested five times, one test for each pairing. The host nest
chosen for the host bimaculatus nest pairing was that nest over which
the female Psithyrus spent the least time during the B. affinis-B. terricola
test.
RESULTS
The preference of P. ashtoni females for nest odours of host and non-host
bumblebees is shown in Table 1. Female Psithyrus were able to distinguish
host nest odour from the odour of the nest material alone, and could also
discriminate between the odour of host nests and non-host nests. Collec-
tively, Psithyrus females showed no preference in the cotton-cotton and B.
a ffinis-B. terricola pairings. When over a host nest the movements of female
Psithyrus slowed, and they often spent considerable time chewing at the
screen mesh floor, apparently attempting to get to the nest below. They never
were observed chewing at the wire mesh floor when over nests containing
only cotton, or when over B. bimaculatus nests.
It was not possible to determine individual preference for either B. affinis
or B. terricola nests in one five-minute experiment period. Collectively the
ten females used in this experiment showed no preference. Two Psithyrus
females which spent greater than 70 percent of their time over B. terricola
nests during the B. terricola-B. affinis pairings were given two additional
five-minute periods under the same conditions. In one case the percent time
spent over the B. terricola nest dropped to 48 percent; in the other, the
preference for B. terricola nest odour remained high (78%).
DISCUSSION
The results support the hypothesis that P. ashtoni females can recognize
host nest odour without actual contact with the nest or with worker-laid
trails. The active detection distance of host nest odour is unknown. Plath
(1934) found that when he dug up nests of B. bimaculatus he observed P.
506
NEW YORK ENTOMOLOGICAL SOCIETY
citrinus females flying nearby making a ‘bee-line’ for the nests and alighting
on them. Presumably, female Psithyrus can recognize nest odour while flying.
P. ashtoni females characteristically fly a slow searching pattern close to the
ground, often alighting and searching in detail under leaves and twigs. Since
both B. affinis and B. terricola nest underground (Fisher, unpubl.), often
with extensive tunnels leading to the nest, the female Psithyrus may at first
be guided by sight to appropriate looking nest entrances, and then localize
nests using odours.
That P. ashtoni can distinguish host nests from those of B. bimaculatus
is consistent with their never being found in nests other than those of hosts.
P. ashtoni females do not fight with host queens, but when placed in non-
host nests are attacked by queens (Fisher, unpubl.). Mistakes in host iden-
tification in the field could be costly. It would be of interest to test generalist
Psithyrus species in the same apparatus, using a variety of bumblebee nests
as odour sources. I hypothesize that host odour discrimination is weakly
developed in Psithyrus generalists.
The data presented here suggest that P. ashtoni females either do not
discriminate between their two hosts, or that discrimination is weakly de-
veloped. More detailed analysis of individual responses is needed to satis-
factorily answer this question. In areas where both hosts occur, B. affinis
queens often usurp incipient nests of B. terricola (Plath, 1934; Fisher, un-
publ.). A female Psithyrus invading a nest of B. terricola very early could
find itself in a nest subsequently usurped by a B. affinis queen. It is unlikely
that B. terricola host specificity could occur under these conditions. P. ash-
toni from areas where B. affinis does not occur can be placed in laboratory
nests of this species where they successfully rear offspring (Fisher, unpubl.).
B. terricola and B. affinis are closely related phylogenetically and may have
colony odours which are difficult to distinguish. B. affinis queens which usurp
B. terricola nests are able, at least in small nests, to dominate B. terricola
workers (Fisher, in prep.), indicating that there is some similarity in pher-
omonal communication between these species. The degree of relative nest
parasitism which occurs in field colonies may simply depend on which host
nests are first encountered by searching parasites.
ACKNOWLEDGMENTS
I thank Robin Stuart for reviewing the manuscript, and Trent University for generously
supplying laboratory space. This research was supported by a NSERC post-graduate scholarship
to Richard M. Fisher.
LITERATURE CITED
Alford, D. V. 1975. Bumblebees. Davis-Poynter, London.
Cederberg, B. 1977. Evidence for trail marking in Bombus terrestris workers (Hymenoptera,
Apidae). Zoon 5:143-146.
VOLUME 91, NUMBER4
507
Cederberg, B. 1979. Odour guided host selection in Psithyrus (Hym., Apidae). Ent. Tidscr.
100:128-129.
Fisher, R. M. Invasion success of the social parasite Psithyrus ashtoni in nests of its two
bumblebee hosts. (Submitted.)
Plath, O. E. 1922. Notes on Psithyrus, with records of two new American hosts. Biol. Bull.
Wood’s Hole 43:23-44.
Plath, O. E. 1934. Bumblebees and Their Ways. New York.
Plowright, R. C. and S. C. Jay. 1966. Rearing bumblebee colonies in captivity. J. Apic. Res.
5:155-165.
Pomeroy, N. and R. C. Plowr:ght. 1980. Maintenance of bumble bee colonies in observation
hives (Hymenoptera:Apidae). Can. Ent. 112:321-326.
Sladen, F. W. L. 191 2. The Humblebee, Its Life History and How to Domesticate It. London.
Received January 21, 1983; accepted April 19, 1983.
NEW YORK ENTOMOLOGICAL SOCIETY
91(4), 1983, pp. 508-511
NOTES AND COMMENTS
MIMICRY, PREDATION AND POTENTIAL POLLINATION
BY THE MANTISPID, CLIMACIELLA BRUNNEA VAR.
INSTABILIS (SAY) (MANTISPIDAE: NEUROPTERA)
Mantispids are little-known members of the order Neuroptera. As the
name Mantispidae implies, the adults bear considerable resemblance, at least
superficially, to the more familiar Mantidae, or praying mantids. Indeed,
this similarity may represent an interesting case of convergence in the insect
world (Poivre, 1976). The life cycles and parasitic habits of the larvae of
many species of mantispids have been previously described by a number of
investigators (Batra, 1972; Hungerford, 1936;Kaston, 1938; Killebrew, 1982;
Milliron, 1940; Parfin, 1958; Redborg, 1982; Rehn, 1939; Smith, 1934;
Viets, 1941). However, little is known about adult mantispids. Recently,
Batra (1982) studied courtship and mating in adults of the mantispid, Cli-
rnaciella brunnea, and Opler (1981) studied polymorphism in this species.
Here I report some previously unrecorded observations on mimicry, pre-
dation, and possible pollination by adults of C. brunnea var. instabilis.
Clirnaciella brunnea ranges from the central and western United States to
Central and South America. The five morphs of this species are Batesian
mimics of different species of paper wasps {Polistes spp.) that occur through-
out its range (Opler, 1981). Adults of C. brunnea var. instabilis mimic adults
of the wasp, Polistes instabilis. The similarity in color and pattern between
model and mimic has already been described in detail by Opler (1981).
Observations were made in Meeker Co., Minnesota, from 15-30 July
1982. Although mantispids are uncommon in Minnesota, I discovered an
aggregation of about 20 individuals on flowering plants of milkweed, Ascle-
pias syriaca L. (Asclepiadaceae).
The adult mantispids showed two types of mimetic behavior in response
to a disturbance. The first was usually exhibited when I approached a plant
they were resting on. A disturbed individual would curl its abdomen beneath
it and retain this position for several minutes. In this posture, with its wasp-
like coloration, it strongly resembled a wasp in the stinging position. The
second type of observed behavior was a startling mimetic display performed
by two different individuals (Fig. 1). When threatened by the close proximity
of photographic equipment, the first mantispid spread all four of its wings
and held them erect in a vertical position above its body. The main part of
the abdomen was expanded laterally, so that its conspicuous yellow, wasp-
like stripes were even more prominent than usual. The mantispid then
repeatedly catapulted its abdomen high into the air over its head and thorax
VOLUME 91, NUMBER 4
509
Fig. 1. Adults of the mantispid, Clirnaciella brunnea var. instabilis in (left) normal prey-
capture position (center) mimicry display, first position, where wings are held vertically over
thorax, abdomen inflated (right) mimicry display, second position, where abdomen is flipped
over thorax and head.
and brought it back down again. This was done in rapid sequence, causing
the insect to rock back and forth. Throughout this display, the tip of the
abdomen was held downwards at nearly a right angle to the rest of the
abdomen, simulating a stinging wasp. This performance lasted about 7 sec-
onds. The display is probably designed to startle or frighten a potential
predator and to show off the mantispid’s vivid wasp-like warning colors,
rather than to exactly simulate a wasp’s behavior. The behavior of a second
individual (when threatened by my finger) was similar to that of the first in
all respects. To my knowledge, this is the first time such a display has been
described for any mantispid species.
When hunting prey, adults of C. brunnea generally hung motionless on
the underside of milkweed leaves or flowers. They are sit-and-wait ambush-
predators that depend on the flowers to attract their prey. Their food consists
of small insects, primarily flies. Prey capture occurred infrequently during
the course of this study. Only three mantispids were observed with prey
(small flies), although the study site was visited daily.
Each mantispid normally occupied a separate plant, but on a few occasions,
several individuals were found together on the same milkweed stalk. These
temporary groupings might have been breeding aggregations, attracted by
the produetion of a pheromone by males, which is used during courtship
(Batra, 1972; Eltringham, 1932).
The wasp models {P. instabilis) also occurred in the study area and behaved
quite differently from the mantispids when searching for food. Unlike the
sessile mantispids, the wasps moved quickly from plant to plant in search
of prey. Their movements were jerky and aggressive. Several wasps, also
hunting on plants of A. syriaca, captured large syrphid flies, which they
quickly dismembered and ate on the spot. I observed both the wasps and
510
NEW YORK ENTOMOLOGICAL SOCIETY
Table 1. Pollen loads of honeybees {Apis mellifera and mantispids {Climaciella brunnea
var. instabilis) captured on milkweed plants {Asclepias syriaca).
No. of pollinaria of A. syriaca
insect was carrying
No. of corpusculae of A. syriaca
insect was carrying
Honeybee no. 1
5
0
2
4
0
3
7
0
4
5
0
5
10
0
6
8
1
7
7
1
8
6
0
9
2
0
10
4
0
11
n
2
Total
69
4
Mantispid no. 1
1
0
2
6
0
3
0
0
4
1
0
5
2
0
6
0
0
7
0
1
8
3
1
9
2
0
10
0
1
11
_0
0
Total
15
3
the mantispids drinking nectar from flowers of A. syriaca. As Opler (1981)
suggests, the similarity of habits and habitats of wasp and mantispid may
have led to the evolution of mimetic polymorphism in C brunnea.
Honeybees {Apis mellifera) were the most frequent visitors to the milkweed
flowers in the study area. For purposes of comparison, and to see if man-
tispids might also function as pollinators of A. syriaca, I captured an equal
number of mantispids and honeybees and examined them for pollen.
Asclepiad flowers, like orchids, have compact pollen masses called pollinia
that adhere to and are carried from flower to flower by insects. The complete
set of pollinia with its associated parts (the corpusculum, or viscidium in
orchids, and stipe) is known as the pollinarium. In the case of Asclepias
flowers, the corpusculae (which are grooved, clasp-like structures) remain
behind, attached to the insect’s legs or body after the pollinia are given up
to the flower.
VOLUME 91, NUMBER 4
511
Eight of the 1 1 mantispids examined carried A. syriaca pollinaria or cor-
pusculae, while these were present on all 1 1 bees (Table 1). On both the bees
and the mantispids these were attached to the legs or the mouthparts. Al-
though the honeybees sampled carried many more pollinaria than the man-
tispids, the number of pollinia the two groups had given up was nearly equal.
Thus, the mantispids, though uncommon, may function as incidental or
occasional pollinators by their habit of capturing prey on milkweed flowers. —
Thomas C. Boyden, Department of Botany (KB- 15), University of Washing-
ton, Seattle, Washington 98195.
ACKNOWLEDGMENTS
I thank T. Pietsch, B. Meeuse and S. Fedorko for comments on the manuscript.
LITERATURE CITED
Batra, S. W. T. 1972. Notes on the behavior and ecology of the mantispid, Climaciella brunnea
occidentalis. J. Kansas Ent. Soc. 45:334-340.
Eltringham, H. 1932. On an extrusible glandular structure in the abdomen of M antispa styriaca
Poda (Neuroptera). Trans. Ent. Soc. London 80:103-105.
Hoffman, C. H. 1936. Notes on Climaciella brunnea var. occidentalis Banks (Mantispidae—
Neuroptera). Bull. Brooklyn Ent. Soc. 31:202-203.
Hungerford, H. B. 1936. The Mantispidae of Douglas Lake, Michigan, region, with some
biological observations (Neurop.). Ent. News 47:85-88.
Kaston, B. J. 1938. Mantispidae parasitic on spider egg sacs. J. New York Ent. Soc. 46:147-
151.
Killebrew, D. W. 1982. Mantispa in a Peucetia egg case. J. Arachnology 10:281-282.
Milliron, H. E. 1940. The emergence of a neotropical mantispid from a spider egg sac. Ann.
Ent. Soc. Amer. 33:357-360.
Opler, P. A. 1981. Polymorphic mimicry of polistine wasps by a neotropical neuropteran.
Biotropica 13:165-176.
Parfin, S. 1958. Notes on the bionomics of the Mantispidae (Neuroptera: Planipennia). Ent.
News 69:203-207.
Poivre, C. 1976. Observations sur la biologie, le comportement et le phenomene de conver-
gence chez les Mantispides (Planipennes). L’Entomologiste 32:2-19.
Redborg, K. E. 1982. Interference by the mantispid Mantispa uhleri with the development
of the spider Lycosa rabida. Ecol. Ent. 7:187-196.
Rehn, J. W. H. 1939. Studies in North American Mantispidae (Neuroptera). Trans. Amer.
Ent. Soc. 65:237-263.
Smith, R. C. 1934. Notes on the Neuroptera and Mecoptera of Kansas with keys for the
identification of species. J. Kansas Ent. Soc. 7:120-145.
Viets, D. D. 1941. A biological note on the Mantispidae (Neuroptera). J. Kansas Ent. Soc.
14:70-71.
NEW YORK ENTOMOLOGICAL SOCIETY
91(4), 1983, pp. 512-523
BOOK REVIEWS
The Nesting Behavior of Dung Beetles (Scarabaeinae). An Ecological and
Evolutive Approach. — Gonzalo Halffter and W. D. Edmonds. 1982. Publi-
cation 10, Instituto de Ecologia, Mexico, D.F., 176 pp. $40.00 in the United
States and Canada, $45.00 elsewhere. Available in North America from
Bioquip Products, P.O. Box 61, Santa Monica, CA 90406.
Evolutive? When I first saw the title of this book, I feared that it might
simply be a clone of The Natural History of Dung Beetles of the Subfamily
Scarabaeinae (Coleoptera, Scarabaeidae) by Hallfter and Matthews (1966).
While the lines of descent between these two works are clearly indicated, I
was pleasantly surprised to see that the new book is not a rehash of old
material at the expense of the reader’s time and patience, but a delightful
refinement and reanalysis of both old and new data on the nesting behavior
of scarabaeines. The book is, appropriately enough, dedicated to Jean Henri
Fabre, the French entomologist who first described in any detail the nesting
behavior of dung beetles and who can be rightly considered the founder of
dung beetle ethology. The contents of this current work add substantially to
the cornerstone molded by Fabre in Souvenirs Entomologique. Halffter and
Edmonds are both eminently qualified to address the nesting behavior of
dung beetles because of their long and productive research with these ani-
mals.
The chapters are entitled (1) The Scarabaeinae, (2) The Ecological Evo-
lution of Scarabaeinae, (3) Patterns of Nesting Behavior in Scarabaeinae:
An Overview, (4) Evolution of Nesting Behavior and Sexual Cooperation,
(5) Nest Construction and Architecture in Burrowing Scarabaeinae, (6) Other
Sexual Relationships in Scarabaeinae, and (7) The Ovary and Nesting Be-
havior. The appendices are (1) Outline/Classification of the Subfamily Scar-
abaeinae, (2) Nidification Behavior of Old World Oniticellini by Yves Cam-
befort, (3) Nesting Strategies of Three Species of Coprophagous Scarabaeinae
in the Sahel Region of Niger by Daniel and Christiane Rougon, and (4)
Commentaries on Recent Literature. An extensive bibliography concludes
the volume.
Chapter one is a general introduction to the subfamily Scarabaeinae and
provides a good, concise picture of dung beetles. Also included is a very
informative table comparing characteristics of scarabaeines, geotrupines and
aphodiines. The major adaptive features of scarabaeines are listed as (1)
food relocation behavior, (2) accentuation of body structure to enhance
fossorial capabilities, (3) subterranean nesting behavior, (4) de-emphasis of
courtship behavior, (5) increased male/female cooperation, (6) reduction in
fecundity, (7) adaptation by larvae and pupae for development within an
enclosed space, (8) mouthparts modified for soft food, and (9) adult digestive
VOLUME 91, NUMBER 4
513
tract modified to rapidly process large amounts of food. Halffter and Ed-
monds state that there is little doubt that the Scarabaeinae is a monophyletic
group. I know of no substantive discussion concerning the derivation and/
or presumed phytogeny of this group of scarabs, and such a prima facie
statement seems out of place without further scientific evidence or docu-
mentation. With the vast array of morphological and behavioral data avail-
able to them, Halffter and Edmonds seem to be in an ideal position to now
address such a phylogenetic analysis at the generic level, and we may hope
that they will engage in such a project in the future.
The principal (and only) shortcoming in the first chapter is near the end
of the third paragraph where the authors confuse the concepts of primitive
and derived conditions in their discussion. Firstly, they indicate that “. . .
coprophagy is the primitive (plesiomorphic) condition . . . and that it derived
from the general saprophagous habit of the family; all other feeding behaviors
(necrophagy, mycetophagy, saprophagy, etc.) we consider derived (apo-
morphic)” (italics mine). Their contention that coprophagy is primitive after
having been derived from another state as well as calling saprophagy both
primitive and derived in the same sentence are non sequiturs. Secondly, they
use the terms plesiomorphic and apomorphic (indicating body structure)
when they mean to use the terms plesiotypic or apotypic (referring to char-
acters other than structure, such as behavior). It seems the authors attempted
to use some catch words currently popular in the phylogenetics and evolution
literature but tripped over their application. While initially confusing and
somewhat bothersome, their foray into terra incognita should not detract
from the overall value of the chapter.
Chapter two delves into the ecological evolution of dung beetles. The
major adaptive features of the subfamily listed in chapter one are discussed
in more detail using the framework of r- and K-selection theory to explain
much of the exploitation of the soil/excrement system. The authors are quick
to note that r- and K-selection represent extremes of a continuum, and that
the evolution of any given species may result in a mixture of r- and K-selected
traits; therefore, these concepts are relative. They conclude that the Scara-
baeinae, relative to other scarabs and most other beetles, are essentially
K-strategists. A convincing discussion follows of scarabaeine traits that ex-
emplify the tactics assumed to be promoted by K-selection. These life history
traits characteristic of Scarabaeinae are (1) survivorship influenced primarily
by density dependent factors mostly operating directly on the adults, (2)
ecological specialists, (3) delayed reproduction, slow development and re-
peated reproductive periods, (4) low fecundity and production of few, large
sized offspring, (5) some degree of brood care, (6) low rates of juvenile
mortality relative to adult mortality, and (7) superior competitors relative
to other groups. This group of traits is a correlated set of adaptations which
collectively define scarabaeines as K-strategists and as insects for which
514
NEW YORK ENTOMOLOGICAL SOCIETY
ecological and reproductive efficiency is maximized. Prior to their discussion
of evolutionary trends within the Scarabaeinae, the authors offer an aplogia
for their lack of a statistically rigorous analysis of populations upon which
to base largely quantitative theory. Their approach is that of natural his-
torians and not population ecologists. No such apology is necessary for this
method of establishing a working hypothesis. As the authors themselves
point out, their experience has produced information that shows strong
tendencies of consistency which demand explanation. Moreover (and much
to their credit) Halffter and Edmonds distill all the descriptive data written
on dung beetles and their own extensive observations into a persuasive,
unified theory describing the ecological evolution of these animals, and they
do this well. This chapter is the the most interesting and significant part of
the book to me. It would be of supreme interest to see how the results of a
study on the taxonomic evolution of these beetles would correlate with the
scenario of behavioral evolution so expertly proposed by Halffter and Ed-
monds.
Chapter three explores the different ways dung beetles nest. A classification
of nesting behaviors is included and is an extension and elaboration of that
of Halffter and Matthews (1966) and Halffter (1977). A classification of nest
morphologies, or patterns, is also provided. There is a great deal of descrip-
tive information in this chapter which, although synoptic in nature, is valu-
able to understanding scarabaeine nidification behavior. It is here that we
find much of the supportive evidence for the conclusions about ecological
evolution proposed in the preceding chapter.
The fourth chapter explores the evolution of nesting behavior and sexual
cooperation and, like chapter three, provides data supporting the conclusions
reached in chapter two. Halffter and Edmonds conclude that nesting behav-
iors originated as derivatives of feeding behavior. Detailed discussion is
given to the evolution of nesting behavior in burrowing scarabaeines and
ball rolling scarabaeines respectively.
The next three chapters are largely descriptive. A detailed review of each
nest type, the taxa constructing it, and the behavior associated with these
taxa constitutes chapter five. The sixth chapter deals with sexual relationships
with a summary of encounter and recognition, copulatory, and combat be-
haviors as well as bisexual cooperation independent of nesting. The last
chapter relates the ovary of dung beetles to nesting behavior with the ob-
servation that there is a reduction in the number of maturing egg follicles
as nesting behavior becomes more highly developed, i.e., there is greater
parental care for fewer offspring.
The first appendix is an outline classification of the subfamily at the generic
level. The classification differs in various respects to that given in Halffter
and Matthews ( 1 966). Immediately noticeable in this respect is the increased
use of subgenera, a move that will be variably accepted depending, in large
VOLUME 91, NUMBER 4
515
measure, upon which side of the Atlantic a scarab systematist works. Neo-
canthidium continues as a valid genus even though it was synonymized by
Howden and Young (1981); a difference of opinion, no doubt.
The papers by Cambefort and Rougon and Rougon constitute the next
two appendices and result from a symposium held on evolution and nesting
behavior in beetles. Cambefort details nicely the nesting behavior of Old
World Oniticellini and formulates an evolutionary scenario for nest building
in these insects. His first conclusion equates low fecundity with a primitive
state which seems to be just the opposite conclusion reached by Halffter and
Edmonds. Cambefort notes that additional species of Oniticellini must be
studied before a firm idea of their behavioral and phylogenetic relationships
can emerge.
The following short paper by the Rougons is a well executed descriptive
work describing the nesting strategies of three African species that live in a
very hot, dry climate.
The book is nicely produced on good quality paper. However, I found
many of the photographs to be unclear because too coarse a photographic
screen was used (see especially Fig. 65) and because of lack of suitable
contrast. The line drawings are good, but many should have been reduced
in size to eliminate their coarse look and provide snap to the illustrations.
The size of the book (about 25 x 25 cm) is awkward because it will not
conveniently rest on most book shelves without protruding excessively from
the shelf. Editing is generally good although there are some glaring exceptions
in the two invited papers and on p. 58 where there are six misspellings.
Considering its moderate size, I thought the book was somewhat overpriced
at $40.00, but this seems to be a contagious malaise within the entire pub-
lishing industry.
In summary, then, I can strongly recommend this book. It will have its
greatest appeal to scarab workers, coleopterists in general, ecologists, and
those conducting research on the evolution of taxa, behavior, or ecological
relationships. I know that some paleontologists as well as some applied
entomologists have also been stimulated by this book. It will have broad
appeal and application to anyone whose life is touched by these remarkable
creatures.— C. Ratcliffe, Systernatics Research Collections, W436 Ne-
braska Hall, University of Nebraska, Lincoln, Nebraska 68588-0514.
LITERATURE CITED
Fabre, J. H. 1920-1924. Souvenirs Entomologique. Librairie Delagrave, Paris, 383 pp. (En-
glish translation available.)
Halffter, G. 1977. Evolution of nidification in the Scarabaeinae. Quaest. Ent. 13:231-253.
Halffter, G. and E. G. Matthews. 1 966. The Natural History of Dung Beetles of the Subfamily
Scarabaeinae (Coleoptera, Scarabaeidae). Fol. Ent. Mexicana No. 12:1-312.
Howden, H. F. and O. P. Young. 1981. Panamanian Scarabaeinae: taxonomy, distribution,
and habits (Coleoptera, Scarabaeidae). Contr. American Ent. Instit. 18:1-204.
516
NEW YORK ENTOMOLOGICAL SOCIETY
Ecologie des Insectes Forestiers.— Roger Dajoz. 1980. Gauthier- Villars, Paris,
478 pp. 280 f.f.
This is a hue attempt to present the essential ecologieal information on
the insects associated with forests; particularly, the forests of France and
Europe. The first part of the book (4 chapters) describes the forests of France
and the general associations of insects. In addition, the last two chapters ( 1 7
and 1 8) on the role of insects in the decomposition of wood and the fauna
of the soil constitute an outline of how insects interact with trees. The
remaining chapters concentrate on the biology of the defoliators, scales, gall
makers, seed utilizers, and wood utilizers. In these chapters there is no
attempt to correlate their activities with soil quality, gross stand effects or
stability influences. It would have been useful to relate ecological effects of
defoliators to changes in soil quality, for example.
Because of the limitation of space, much information about the ecology
of forest insects has been omitted. Despite this, the book provides a summary
of the life and activities of some of the more economically important insects
in European forests. The French literature is particularly well covered with
less emphasis on the German and British contributions.
The emphasis on ‘‘pest” species is a natural tendency but there are many
other species in the forest that contribute to the ecology and survival of trees
and associated plants. These species and their interactions should have been
discussed in an ecology textbook. Nevertheless, there is much useful infor-
mation supplemented by numerous graphs, tables and figures which should
prove to be valuable to both the advanced student and professional ento-
mologist. Precise analyses of the population dynamics of several species are
particularly important because attention is often focused on the physiological
adaptations affecting abundance and dispersal. It is this aspect of the book
that makes it most usQiviX. — Gordon R. Stairs, Department of Entomology,
Ohio State University, Columbus, Ohio 43210.
The Biology of Social Insects.— M. D. Breed, C. D. Michener, and H. E.
Evans (eds.). 1982. Westview Press, Boulder, Colorado, 419 pp. $25.00.
During August 9-13, 1982, the International Union for the Study of Social
Insects (lUSSI) held its Ninth Congress in Boulder, Colorado. Upon arrival,
each participant received a copy of the book. The Biology of Social Insects,
which contains papers from 10 symposia as well as abstracts representing
most of what was to be seen and heard during the meeting. Given the usual
hustle and bustle of organizing an international meeting, the production of
a finished book, before the start of the meeting, is laudatory, and it reflects
the current status of knowledge and ideas about many aspects of the study
VOLUME 91, NUMBER 4
517
of social insects. The topics covered were chosen by specialists on social
insects from all over the world who were asked to suggest symposium topics.
The Organizing Committee of the Congress then chose ten from the ap-
proximately 100 suggested subjects. The result was ten very different but
complementary symposia.
The best way to present this book of such varied chapters and authors is
to list each symposium title and to mention what I found to be notable
about each. Before proceeding I should mention that within each symposium
were invited papers and submitted papers, the latter represented by abstracts.
Abstracts of presented papers are useful when one is attending a meeting,
however, the usefulness of abstracts within the book is scant. One imme-
diately knows what kinds of studies are being done, but details of the majority
of the abstracted studies are necessarily lacking. A total of 72 of the total
414 text pages is devoted to the abstracts alone.
Symposium I: Foraging Behavior and Pollination. Four papers and an
introduction by G. D. Waller. Waller states that the direction of future
research on foraging behavior and pollination may be profoundly affected
by what transpired at this symposium. Models are presented which challenge
the theory of optimal foraging and R. Jander discusses a new theory called
‘modal tuning theory.’ The papers vary from empirical to theoretical.
Symposium II: Competition and Population Dynamics in Social Insects.
Three papers and an introduction by J. H. Sudd. Especially interesting is a
paper by N. Koeniger on the interactions among the four species of the genus
Apis. The importation of A. mellifera has occurred worldwide with little
consideration of the unstable condition that occurs when competing species
are brought together. Koeniger examined competition for nesting sites, com-
petition for food, interference during mating, interspecific robbing, predators
and exchange of parasites. The results of this study highlight the importance
of applied, as a complement of basic, studies. The oftentimes neglected but
far-reaching effect of introducing social insects to foreign habitats is also
stressed.
Symposium III: The Roles of Social Insects in Ecosystems. Six papers and
an introduction by W. L. Nutting. This symposium considers the effect of
termites, ants and wasps on ecosystems through modification of habitat,
contribution to energy flow and cycling of nutrients, and interactions with
other organisms. The numerical dominance of social insects in tropical for-
ests is well known, yet the ecological role of these organisms is only vaguely
understood. Abe’s paper on the ecological role of termites in a tropical rain
forest helps to fill this gap in our understanding. Data on density and biomass
of termites as well as food consumption by fungus-growing termites are
presented in valuable summary tables. Collins’ study of the interaction and
impact of cattle and termites in Kenya stresses again the important inter-
action between applied and basic research. Another notable paper is Greene’s
518
NEW YORK ENTOMOLOGICAL SOCIETY
report on comparative early growth and foraging of two naturally established
vespine wasp colonies. Vespula germanica and V. vidua colonies were ob-
served in hopes of determining why V. germanica tends to be the more
successful yellowjacket. In spite of the equal early growth rates of the colonies,
a striking disparity appeared due to longer adult life span and ability to
maintain greater numbers of brood because of more efficient foraging by V.
germanica (1,167 cells for germanica vs. 531 for vidua). In addition, there
was an almost complete lack of overlap in food items with germanica being
omnivorous while vidua is more specialized on arthropod prey. The results
of this study allow insight as to why germanica is one of the world’s most
pestiferous social wasps. Indeed, more comparative studies such as this may
lead to an understanding of the evolution of generalists versus specialists
within the social insects.
Symposium IV: Economically Important Social Insects. Eight papers and
an introduction by S. B. Vinson. By far this symposium is the best written
and most interesting. In a time when insect behavior is a popular field of
study, the potential for applied research exists. Indeed, in a field flooded
with new PhDs, the need for career alternatives increases. The papers pre-
sented in this symposium highlight the opportunity for studies on econom-
ically important social insects. Topics include yellowjackets (according to
Akre, responsible for millions of dollars of damages in the U.S.), leaf-cutting
ants, the use of JH-analogue for control of Monomorium pharaonis (the
control takes advantage of the recruitment behavior of the ants), fire ants,
Polistes and caterpillar suppression, and Africanized bees.
Symposium V: Presocial Behavior. Five papers and an introduction by G.
C. Eickwort. As stated in Eickwort’s introduction, analyses of presocial be-
havior are providing significant insights into the conditions under which
reproductive division of labor and eusocial behavior might evolve. One of
the more interesting findings in recent years, in support of kin-selection
hypotheses, is Aoki’s finding of soldiers in parthenogenic aphid colonies.
These soldiers are morphologically distinct and reproductively sterile. Aoki’s
paper on aphids discusses new findings as well as leads the reader to his
numerous other references on aphid soldiers. Evans and Hook report on
communal nesting in Cerceris digger wasps, which appears to have evolved
in response to natural predators. The comparison between solitary and com-
munal Cerceris species represents the type of study needed to determine the
factors involved in the evolution of social behavior. Another interesting
paper is on subsocial behavior in Coleoptera. In contrast to E. O. Wilson’s
(1975) ideas of the evolution of sexuality as an antisocial phenomenon,
Halffter presents the hypothesis that in Coleoptera there is an evolution
towards subsocial forms of behavior through development of patterns based
on sexuality. Also included in the symposium are a paper on membracid
sociality and one on an experimental induction of multifemale associations
VOLUME 91, NUMBER 4
519
in solitary Ceratina bees. As a whole, this symposium presents four distinct
studies that all emphasize the varied routes from solitary to social behavior.
Symposium VI: The Evolution and Ontogeny of Eusociality. Six papers
and an introduction by M. J. West-Eberhard. The theme of this symposium
is intracolony reproductive competition, with parasitic ovipositions by un-
mated females and polygynous queens. The data reported increase the amount
of evidence of intracolony competition, thus promoting even further the
decline of the once popular image of a social insect colony as an harmonious
supraorganism. The studies stress the need for detailed studies of individual
and colony histories necessary to evaluate evolutionary interpretations. A
notable paper was by Hunt on trophallaxis in Hymenoptera; he proposes
that trophallaxis may be central to the evolution of sociality within the
Hymenoptera. Once proposed as the cornerstone of vespid sociality by Rou-
baud (1916), the idea of trophallaxis as critical to the evolution of eusociality
recently lost favor. Hunt’s ideas may rekindle an interesting controversy.
Also dealing with the evolution of eusociality is a plenary address by R.
H. Crozier (pp. 4-9). Although brief, this overview of the recent controversies
in this important question in evolutionary biology is valuable and quickly
takes the reader to the more important recent papers on this topic.
Symposium VII: Caste and Ergonomics. Four papers and an introduction
by J. M. Herbers. Dominance behavior in primitively eusocial species {Lasi-
oglossum and Polistes) is correlated with reproductive status. In an inter-
esting study. Breed determined that in the highly eusocial honey bee, where
overt dominance interactions are rare, a relationship exists between aggres-
sive behavior towards queens and factors that affect the endocrine status of
the worker honey bee. Another interesting study was by J. M. Herbers on
queen number and colony ergonomics in Leptothorax longispinosus. In an
attempt to determine the significance of multiple queens in a nest, Herbers
constructed an hypothesis which states that increased genotypic variation
in polygynous colonies could result in a broader distribution of worker
phenotypes leading to higher efficiency and higher fitness. No correlation
was found, however, and thus the hypothesis was rejected. Even though the
maintenance of polygynous colonies in nature remains a mystery, Herbers’
study is stimulating and hopefully will lead to more hypothesis-testing.
Symposium VIII: Predation, Social Parasitism and Defense. Six papers
and an introduction by R. W. Matthews. Matthews states that this is a series
of definitive papers focusing on slavery in ants, ant predation and wasp and
bee colony defensive strategies. Most striking were the two papers on army
ants and their effect on social insects. Chadab-Crepet and Rettenmeyer pres-
ent a well-written report on the comparative behavior of social wasps when
attacked by army ants. Intriguing points were that some wasps can discrim-
inate between ant species; Protopolykia exigua, for example, becomes
alarmed by the odor of army ants enabling this species to evacuate before
520
NEW YORK ENTOMOLOGICAL SOCIETY
the ants have plugged the tiny nest entrance. Surprisingly, social wasps have
no effective direct defense against army ants; the only escape is evacuation.
The suggestion that development of polygyny in the Polistinae may be a
result of army ant predation again highlights the importance of natural
enemies in the evolution of eusociality.
Symposium IX: Communication. Six papers and an introduction by R.
Boch. Included were papers on chemical communication in ants and new
exocrine glands, honey bee mandibular glands, swarming in honey bees,
behavior genetics and recruitment in ants. Two papers, one applied and one
theoretical, stood out as exceptional. The first, ‘The adaptive value of prob-
abalistic behavior during food recruitment in ants . . .’ by Pasteels, Verhaeghe
and Deheubourg, via experimental and theoretical evidence, showed that
some level of inaccuracy in communication (recruitment) can be advan-
tageous for the society by increasing the probability of discoveries. The
second, ‘Behavior genetics of honey bee alarm communication’ by A. M.
Collins, was a study of genetic selection as a way to combat the defensiveness
of the Africanized bee.
Symposium X: Neurobiology and Behavior of Social Insects. Twelve pa-
pers and an introduction by R. Menzel. Insects, especially the honey bee,
have long been considered prime subjects for the study of neurobiology.
Within this symposium are technical papers on morphology, a good review
of recent studies of memory, a presentation of the celestial map used by
honey bees in orientation and a study on control of heart rate. Interestingly,
the heart rate in honey bees is not under direct nervous control but rather
the beat frequency is sensitive to the sodium concentration in the hemo-
lymph.
How does this book compare with the newly published four volume set.
Social Insects, edited by H. R. Hermann? In other words, was another book
on social insects needed at this time? Hermann’s book contains lengthy
reviews of many groups of social insects, caste differentiation and evolution
of eusociality. The last two topics were covered in the lUSSI publication as
well and perhaps add little to the available literature. In contrast, the chapters
on the economic importance of social insects, foraging, communication,
competition, neurobiology, and predators are distinct, and the mixture of
applied, empirical and theoretical studies makes the lUSSI book unique.
My impression was that much of the detail would be lost on today’s spe-
cialists, but an overview of studies being conducted on social insects is
worthwhile. The necessarily short papers, although lacking in detail, will
lead the interested reader directly into relevant literature. The book is a
valuable reference for this reason alone.
Most of the contributions to the book were prepared as camera-ready by
the authors and published as received; typographical errors and inaccurate
and sometimes confusing translations are not uncommon distractions. But
VOLUME 91, NUMBER 4
521
on the whole, the papers are of high qn?i\\Xy.— Janice S. Edgerly, Department
of Entomology, Cornell University, Ithaca, New York 14853.
LITERATURE CITED
Roubaud, E. 1916. Recherches biologique sur les guepes solitaires et sociales d’Afrique. La
genese de la vie sociale et revolution de I’instinct matemel chez les vespides. Ann. Sci.
Nat. 1(10):1-160.
Wilson, E. O. 1975. Sociobiology. The New Synthesis. Harvard University Press, 697 pp.
Insects of an Amazon Forest.— Norman D. Penny and Jorge R. Arias. 1982.
Columbia University Press, New York, 269 pp. $30.00.
The authors present the results of a full year of sampling in Brazil. Using
five sampling techniques, aerial (1 and 15 m) light traps, soil emergence
traps, Berlese-Tullgren extractions of leaf litter. Malaise traps, and baited
pitfall traps, a large number of insects were obtained. Specimens were sorted
by order, and, for a few orders, by family. Dry weight estimates were obtained
for those groups collected during the last week of sampling. Undoubtedly
the scope of sampling would preclude the identification of the majority of
specimens unless a large team of taxonomists was involved. Still, the results
of such a study could be highly informative.
The book jacket, and the publisher’s advertisements, proclaim it to be a
“classic study unlike any other for an area of Amazon forest ... of major
interest to tropical ecologists and entomologists as well as biologists con-
cerned with evolution, classification, or the tropics.” Given such a glowing
introduction, readers will undoubtedly be disappointed. Indeed, it is unclear
as to the exact nature of the readership for which the book was written, but
it was not for those previously cited. Excellent drawings of representatives
of many families are given, but their treatment is scarce— averaging about
one paragraph per family. Even here, the treatment is very general, giving
the estimated world-wide number of species for that family, as well as gen-
eralizations of their biology. The book is highly reminiscent of Swan and
Papp’s The Common Insects of North America, although its treatment is
much less clear. If the attempt of the authors was to produce a similar book,
they almost made it, but it is highly doubtful if swarms of amateur ento-
mologists will be able to use their book, unlike The Common Insects of
North America.
However, the large print, and the ample number of illustrations, as well
as the lack of any substantive data, make the book easy reading, and, if one
has any interest in entomology, the entire book can be read in from one to
two hours. This book may make a good gift for a beginning amateur ento-
mologist, but will not serve any useful function in the library of a profes-
sional.—Eowler, University of Elorida, Gainesville, Elorida 32611.
522
NEW YORK ENTOMOLOGICAL SOCIETY
The Semiaquatic Bugs (Hemiptera, Gerromorpha): Phylogeny, Adaptations,
Biogeography, and Classification.— N. Moller Andersen. 1982. Entomono-
graph Vol. 3. Scandinavian Science Press Ltd., Klampenborg, Denmark, 455
pp. $35.00.
Andersen’s book represents something of a landmark in the study of the
Hemiptera, for seldom does such a comprehensive treatment of a group
appear. The last obvious example was “Monograph of Cimicidae” (Usinger,
1966), which dealt with the bedbugs in more detail but with a somewhat
different emphasis. Andersen has singlehandedly attempted a generic level
morphological review and phylogenetic analysis as well as presenting a dis-
cussion of adaptations, historical biogeography, and classification. In the
form of appendices he presents for the Gerromorpha lists of names and keys
for all higher taxa and genera.
The semiaquatic bugs in Andersen’s sense include the families Gerridae,
Hebridae, Hermatobatidae, Hydrometridae, Macroveliidae, Mesoveliidae,
Paraphrynoveliidae, and Veliidae. This usage may be unfamiliar to some
workers, especially in North America, where groups such as the Saldidae,
Gelastocoridae, and Ochteridae are often referred to as semiaquatic He-
miptera. Andersen presents an interesting review of the history of the clas-
sification of the Gerromorpha, documenting its monophyly with what ap-
pears to me to be strong morphological evidence. One must conclude from
the results of his analysis that the more inclusive use of the term semiaquatic
should be abandoned by hemipterists.
Andersen demonstrates his skill as an artist and a technician, presenting
several hundred well executed line drawings based on original light micro-
scopic work as well as many scanning electron micrographs— some published
here for the first time with the remainder gathered from a string of revisionary
works which Andersen began on the Gerromorpha nearly two decades ago.
This volume appears at a time when systematics and biogeography are
being revolutionized by methodological innovation. Nils Andersen has made
a serious attempt to incorporate both the techniques of cladistics and vi-
cariance biogeography into his analysis. I consider his initial explication of
principles basically redundant with what already exists in the literature.
Nonetheless, these principles as outlined by Andersen clearly indicate the
somewhat divergent emphasis in phylogenetics in Andersen’s work— and
probably that of most continental Europeans— with the approach adopted
by many North Americans. Notable is Andersen’s emphasis on reconstruc-
tion of the ground plan.
The minor differences in cladistic methods as espoused by Andersen and
practiced by others are far less important than his desire to be explicit about
his methods. His book presents a theory of relationships for the Gerromor-
pha that contradicts the findings of Cobben (1978) and Popov (1971). None-
theless, it appears to me that Andersen offers a consistent interpretation of
VOLUME 91, NUMBER 4
523
the available evidence and has presented an example of the type of com-
prehensive documentation for which all of us should strive. Whatever the
merits of his conclusions, his hypotheses can be easily understood and readily
tested by all future workers.
The analysis presented by Andersen will be most enthusiastically read by
systematists. Nonetheless, the synthetic nature of Andersen’s work will make
this comparatively inexpensive volume a valuable reference for all biologists
interested in the Gmomor^hsi.— Randall T. Schuh, Department of Ento-
mology, American Museum of Natural History, New York, New York 10024.
LITERATURE CITED
Cobben, R. H. 1978. Evolutionary trends in Heteroptera. Part II. Mouthpart-structures and
feeding strategies. Meded. Landbouwhogeschool, Wageningen, Netherlands, 78-5,
407 pp.
Popov, Y. A. 1971. [Historical development of Hemiptera infraorder Nepomorpha (Heter-
optera).] Trudy Paleontological Institute, Acad. Sci. USSR 129:1-228 (in Russian).
Usinger, R. L. 1966. Monograph of Cimicidae (Hemiptera-Heteroptera). The Thomas Say
Foundation, Entomological Society of America, College Park, Maryland, vol. 7, 585 pp.
REVIEWERS FOR 1983
The Editorial Staff thanks the following persons who reviewed manuscripts
submitted to the Journal for publication: Roger D. Akre, Allen H. Benton,
J. Milton Campbell, Norman T. Davis, Richard C. Froeschner, Charles S.
Henry, Thomas J. Henry, Lee H. Herman, A. C. Hodson, Lloyd Knutson,
James P. Kramer, J. E. McPherson, Arnold S. Menke, David A. Nickle, Lois
O’Brien, Eric Quinter, J. G. Rozen, Jr., Michael Schauff, Michael D. Schwartz,
Alex Slater, James A. Slater, Roy R. Snelling, Frederick W. Stehr, John
Stoffolano, Howard Topoff, Alfred G. Wheeler, Jr., and Pedro Wygodzinsky.
{Continued from back cover)
On the biology of cave inhabiting Antillocorini with the description of a new species
from New Guinea (Hemiptera: Lygaeidae) James A. Slater
The types of some American Cerceris with lectotype designations (Hymenoptera:
Philanthidae) George R. Ferguson
Type designations and new synonymies for Nearctic species of Phytocoris Fallen
(Hemiptera: Miridae) Thomas J. Henry and Gary M. Stonedahl
An annotated synonymic list of North American and Caribbean wasps of the genus
Cerceris (Hymenoptera: Philanthidae) George R. Ferguson
Recognition of host nest odour by the bumblebee social parasite Psithyrus ashtoni
(Hymenoptera: Apidae) Richard M. Fisher
Notes and Comments
Mimicry, predation and potential pollination by the mantispid, Climaciella brunnea
var. instabilis (Say) (Mantispidae: Neuroptera) Thomas C. Boyden
Book Reviews
The nesting behavior of dung beetles (Scarabaeinae). An ecological and evolutive
approach Brett C. Ratclijfe
Ecologie des insectes forestiers Gordon R. Stairs
The biology of social insects Janice S. Edgerly
Insects of an Amazon forest Harold Fowler
The semiaquatic bugs (Hemiptera, Gerromorpha): Phylogeny, adaptations, biogeog-
raphy, and classification Randall T. Schuh
Reviewers for 1983
424-430
431-441
442-465
466-502
503-507
508-511
512-516
516
516-521
521
522-523
524
Journal of the
New York Entomological Society
VOLUME 91 DECEMBER 1983 NO. 4
CONTENTS
Revision of the Philanthus zebratus group (Hymenoptera: Philanthidae)
George R. Ferguson
Establishment of Hyles euphorbiae (L.) (Lepidoptera: Sphingidae) in the United States
for control of the weedy spurges Euphorbia esula L. and E. cyparissias L.
S'. W. T. Batra
Patterns of distribution and abundance in small samples of litter-inhabiting orthoptera
in some Costa Rican cacao plantations Allen M. Young
Coordinated prey capture by Novomessor cockerelli (Hymenoptera; Formicidae)
Harold G. Fowler and Walter G. Whitford
Habitat preferences of carrion beetles in the Great Swamp National Wildlife Refuge,
New Jersey (Coleoptera: Silphidae, Dermestidae, Nitidulidae, Histeridae, Scara-
baeidae) Paul P. Shubeck
Microclimate observations and diel activities of certain carrion arthropods in the
Chihuahuan desert Kenneth Schoenly
Seasonal dynamics of fleas associated with the gray-tailed vole, Microtus canicaudus
Miller, in western Oregon Richard G. Robbins
Histology of the male reproductive systems in the adults and pupae of two doryline
ants, Dorylus {Anomma) wilverthi Emery and D. {A.) nigricans Illiger (Hymenop-
tera; Formicidae) Francis C. Ford and James Forbes
Anatomy and histology of the male reproductive system in the adult and pupa of the
doryline ant, Aenictus gracilis Emery (Hymenoptera; Formicidae)
S. Shyamalanath and James Forbes
Cuticular pigment changes in worker Yellowjackets (Hymenoptera; Vespidae)
Kenneth G. Ross
Ecological and sensory aspects of prey capture by the whirligig beetle Dineutes discolor
(Coleoptera; Gyrinidae) Steven A. Kolmes
Irbisia knighti, a new mirine plant bug (Heteroptera; Miridae) from the Pacific
Northwest Michael D. Schwartz and John D. Lattin
Descriptions of the nymphal instars of Oecleus borealis (Homoptera; Fulgoroidea;
Cixiidae) Stephen W. Wilson, James H. Tsai, and Catherine R. Thompson
289-303
304-311
312-327
328-332
333-341
342-347
348-354
355-376
377-393
394_404
405-412
413-417
418-423
{Continued on inside back cover)
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