SOCIETy,
Journal o
Hymenoptera
Research
Volume 3
August 1994
ISSN 1070-9428
CONTENTS
ANTROPOV, A. V. A review of the agile species group of Pison (Hymenoptera: Sphecidae: Trypoxylini) 119
AZEVEDO, C. O. Descriptions of two new species and notes on the genus Bakeriella Kieffer from Brazil and Ecuador
(Hymenoptera, Bethylidae) 145
BELSHAW, R. and B. BOLTON. A survey of the leaf litter ant fauna in Gahana, West Africa (Hymenoptera: Formicidae) ... 5
BITONDI, M. M. G., Z. L. P. SIMOES, A. M. do NASCIMENTO and S. L. GARCIA. Variation in the haemolymph protein
composition of confined Apis mellifera and potential restoration of vitellogenin titre by juvenile hormone analogue
treatment 107
BOHART, R. M. A review of North American Belomicrus (Hymenoptera, Sphecidae, Crabroninae) 207
DINIZ, I. R. and K. KITAYAMA. Colony densities and preferences for nest habitats of some social wasps in Mato Grosso State,
Brazil (Hymenoptera, Vespidae) 133
FIELD, S. A. and M. A. KELLER. Localization of the female sex pheromone gland in Cotesia rubecula Marshall (Hymenoptera:
Braconidae) 151
GUPTA, V. K. A review of the world species of Orthomiscus Mason (Hymenoptera: Ichneumonidae: Tryphoninae) 157
HERATY, ]. M., ]. B. WOOLLEY and D. C. DARLING. Phylogenetic implications of the mesofurca and mesopostnotum in
Hymenoptera 241
KERR, W. E and E. B. SOARES REZENDE. Genetic characters of African bees that have high adaptive value in the tropics . 1
KUGLER, C. Revision of the ant genus Rogeria with descriptions of the sting apparatus (Hymenoptera: Formicidae) 17
NIEVES-ALDREY, J. L. Revision of West-European genera of the tribe Aylacini Ashmead (Hymenoptera, Cynipidae) 175
OVRUSKI, S. M. Immature stages of Aganaspis pelleranoi (Brethes) (Hymenoptera: Cynipoidea: Eucoilidae), a parasitoid of
Ceratitis capitata (Wied.) and Anastreplm spp. (Diptera: Tephritidae) 233
POLASZEK, A. and K. V. KROMBEIN. The genera of Bethylinae (Hymenoptera: Bethylidae) 91
QUICKE, D. L. J. Myosomatoides gen. nov. (Hymenoptera: Braconidae), a Neotropical larval parasitoid of stem-borer pests,
Diatraea (Lepidoptera: Pyralidae) 227
QUINTERO A., D. and R. A. CAMBRA T. Systematics of Pseudomethoca areta (Cameron): sex association, description of the male
and a gynandromorph, and a new synonymy (Hymenoptera: Mutillidae) 303
TANG, Y. and P. M. MARSH. A taxonomic study of the genus Ascogaster in China (Hymenoptera: Braconidae: Cheloninae) .
279
Additions and corrections to Volume 2, Number 1, 1993 309
INTERNATIONAL SOCIETY OF HYMENOPTERISTS
Organized 1982; Incorporated 1991
OFFICERS FOR 1994
George C. Eikwort, President
Donald L. J. Quicke, President-Elect
Michael E. Schauff, Secretary
Gary A. P. Gibson, Treasurer
Paul M. Marsh, Editor
Subject Editors
John Huber, Arnold Menke, David Rosen, Mark Shaw, Robert Matthews
All correspondence concerning Society business should be mailed to the appropriate officer at the following
addresses: President, Department of Entomology, Cornell University, Ithaca, New York 14853; President-
Elect, Department of Biology, Imperial College at Silwood Park, Ascot, Berks SL5 7PY, England; Secretary,
c/o Department of Entomology, NHB 168, Smithsonian Institution, Washington, DC 20560; Treasurer,
Biological Resources Division, CLBRR, Agriculture Canada, K.W. Neatby Building, Ottawa, Ontario,
Canada K1A 0C6; Editor, P. O. Box 384, North Newton, Kansas 67117.
Membership. Members shall be persons who have demonstrated interest in the science of entomology.
Annual dues for members are $25.00 (U.S. currency) per year, payable to The International Society of
Hymenopterists. Requests for membership should be sent to the Treasurer (address above).
Journal. The journal is published once a year by the International Sociey of Hymenopterists, c/o Department
of Entomology, NHB 168, Smithsonian Institution, Washington, DC 20560, U.S.A. Members in good
standing receive the Journal of Hymenoptera Research. Nonmember subscriptions are $50.00 (U.S. currency)
per year.
The Society does not exchange its publications for those of other societies
Please see inside back cover of this issue for information regarding
preparation of manuscripts.
Statement of Ownership
Title of Publication: Journal of Hymenoptera Research.
Frequence of Issue: Once a year (currently).
Location of Office of Publication, Business Office of Publisher and Owner: International Society of
Hymenopterists, c/o Department of Entomology, NHB 168, Smithsonian Institution, Washington, DC
20560, U.S.A.
Editor: Paul M. Marsh, P. O. Box 384, North Newton, Kansas 67117.
Managing Editor and Known Bondholders or other Security Holders: none
This issue was mailed 15 October 1994
George Campbell Eickwort
1940-1994
We were saddened to hear of the untimely death of George
Eickwort, President, International Society of Hymenopterists,
on July 11, 1994. George died from injuries suffered in an
automobile accident while vacationing in Jamaica. This issue
of the Journal of Hymenoptera Research is dedicated to his
memory and to honor his contributions to the study of
Hymenoptera and to this Society.
First Announcement
Third International
Hymenoptera Conference
August 12-17, 1995
University of California
Davis, California, USA
Talks, poster presentations and symposia on all aspects of Hymenoptera,
including:
•Biological control
•Behavior
•Morphology
•Genetics
•Systematics
Also, possiPle tours of the Bohart Museum, California Academy of Sciences,
San Francisco and the northern California wine country
To be put on mailing list tor second announcement with registrationand call
for papers, send name, address, phone, fax and/or e-mail numbers to:
Dr. Lynn S. Kimsey
Department of Entomology
University of California
Davis, California 95616 USA
Phone: (916) 753-5373 FAX (916) 752-1 537
A few grants to cover travel costs may Pe availaPle to participants from
countries with foreign currency or other financial proPlems. Mention your
need when requesting the second accouncement.
J. HYM. RES.
Vol 3, 1994, pp. 1-4
Genetic Characters of African Bees That Have High
Adaptive Value in the Tropics
Warwick Estevam Kerr and Eduardo Badue Soares Rezende
Universidade Federal de Uberlandia, Departamento de Biociencias, 38400-902, Uberlandia, MG, Brazil
Abstract- The Ac gene present in Africanized Apis mellifera populations is male limited and confers bronze color to the
abdomen; females are non-affected. The gene ac confers yellow abdomen equally to Italian female and male bees. Afncan-mt-
DNA is found in African and most Africanized populations, and European-mt-DNA occurs in European populations, and
descendents. The original frequency in Camaqua in 1957 of ac was 0.939 and for Ac 0.061. In 1991, after about 68 generations,
these frequencies changed to 0.038 toacand 0.962 to Ac, what gives an adaptive value to ac in the tropics of 0.952 (considering
1.00 to Ac). The same mav happen tothemt-DNAs, what mav cause the mates European-mt-DNA queens X Afncan-mt-DNA
males to be less fit than the Afncan-mt-DNA queens X European-mt-DNA males.
INTRODUCTION
There is a constant search in tropical research
for characters that give to the bearer species a
higher adaptive value when compared to Euro-
pean ones. This happened with corn, wheat, cab-
bage, apples, cattle, goats, etc. Many morphologi-
cal, behavioral and physiological characters are
being studied in European and African bees, and
in their hybrids under different ecological condi-
tions. Among them, two of the African characters
are particularly interesting because within a few
generations they became predominant in a tropi-
cal environment, but not in a temperate one. These
characters are the gene Ac and the African-mt-
DNA (that acts as a single gene). The population of
this area is being well studied as far as their
enzymes are concerned and show that the
Africanized honeybee is an admixture of Apis
mellifera mellifera (19.5%), Apis mellifera ligustica
(3.8%) and Apis mellifera scutellata (76.7%) (Lobo et
al. 1989; Del Lama et al. 1990).
The African-mt-DNA has been studied by
Hall and Muralidharan (1989), Sheppard et al.
(1991), Sheppard et al. (1991), Soares (1992) and
others. It shows indications of high adaptive value
in the tropics and low in southern South America
(that has European-like climate), where it is domi-
nated by European mt-DNA.
The Ac gene has been observed by Brazilian
bee biologists since 1957. In 1969 the information
published that it is male sex limited: it confers
bronze color to the abdomen, especially to the
tergites, while workers are yellow (Kerr 1969). It
was present in 100% of the 145 queens collected in
Africa and brought to Brazil in 1956. There was
some information (Prof.V. Portugal Araujo, p.c.)
that yellow males were occasionally present in
Angolan populations. However, they were never
seen by W. E. Kerr in his 1956 trip to Africa. It is an
allele of b(=black) (Woyke and Kerr 1989), segre-
gates lAc: lac in different genetic background,
and is not linked to five components of agressive
behavior (Stort 1978).
MATERIAL AND METHODS
Between 19 July 1990 and 17 September 1991,
14 swarms of Africanized bees (Apis mellifera Linne)
entered emptv stingless bee hive boxes located in
the Apiary of the Universidade Federal de
Uberlandia, MG, Brazil. Hive boxes varied in size,
from 15 to 45 litres in volume, located 420 km from
Camaqua, the point of introduction of African
bees, in 1957. All swarms contained many males.
A sample of males was taken from each swarm
and the numbers expressing the Ac and ac alleles
were determined.
Journal of Hymenoptera Research
Males from an additional 17 colonies of
Africanized bees were sampled from commercial
honeybee hives that were occupied by Africanized
swarms. Frequencies of the ac and Ac genes were
calculated and compared with estimates of the
frequencies of the same alleles in Camaqua in
1957.
RESULTS
Six hundred and ninety (690) Ac drones
(0.9623) and 27 ac drones (0.0377) were sampled
from the 31 colonies. No data for 1992 and 1993
were collected because all swarms (22 and 9, re-
spectively) had only Ac drones. Drones in a swarm
come from several colonies.
Some additional observations made are im-
portant to note: 1. Of the 14 swarms, only one had
two queens (the same proportion found in Kerr et
al. 1970), 2. in three swarms many bees with wax
scales were seen, which indicates that bees of
different ages were in the swarms, 3. in 1992 two
and in 1993 two swarms arrived without males.
The lack of ac drones in 1992 and 1993 swarms
is assumed to be a consequence of the population
reaching fixation (100% Ac genes) or near fixation.
DISCUSSION
Since the gene Ac segregates independent of
the xo gene, is independent of genes for defensive
behavior and is an allele of black (b), the hypoth-
esis that it was linked and continues to be linked
with genes for high fitness after 68 generations of
meiosis was discarded.
Several traits of Africanized bees have been
and are being selected for Brazilian conditions
(Page and Kerr 1991; Kerr 1992). The bees are
becoming less aggressive; they are being selected
for greater resistance to the Varroa jacobsoni mite
(Moretto et al. 1991); they do not any more reject
Italian foundation and they use fewer armadillo
holes (Kerr 1992). The high frequencies of Ac and
of African-mt-DNA are also a result of natural
selection, since the degree of natural crosses be-
tween Africanized x Italian and Italian x
Africanized are about equal when queens and
drones of both races use the same mating ground
(Kerr and Bueno 1970).
In 1956, there were 400 Italian hives located in
the same Eucalyptus forest into which 26 swarms
of Africanbees escaped. All Italian colonies had ac
and all African colonies had Ac genes. Therefore,
assuming equal contributions of all escaped colo-
nies to the breeding population, a frequency of
0.939 for the gene ac and 0.061 for Ac in 1957 was
obtained for that original population. According
to Winston (1992 pg. 40-42) an individual
Africanized colony swarms about 16 times per
year. Of course, this can only happen in the expan-
sion phase of the population, before the popula-
tion approaches the carrying capacity of the envi-
ronment. According to Nascimento (1981 page
166), however, this figure, obtained in 1980, is 1.5
swarms per year. For our estimate a conservative
"intermediate" figure of two successful swarms
per colony per year will be used.
The original frequency F1"57 of the gene ac
(0.939) in 1957, multiplied, per generation, by its
relative adaptive value (w) will give us the fre-
quency F1*"1 for ac in 1991, that is 0.0377, 34 years
later, or after 68 generations. Therefore: F1991 = wn.
F11^7. In this formula n, the number of generations,
is 68, F1957 is 0.939, F,MM1 is 0.0377 and our estimation
for w, the fitness of ac, is 0.952 (considering 1.00 to
Ac). This relatively low adaptive value w of ac is
the reason for the almost universal presence of Ac
in feral populations of Apis meUifern in tropical
South America. It may be a similar reason for the
the high frequency of African-mt-DNA found in
Africanized populations, that is colonies of Euro-
pean-mt-DNA queens x African-mt-DNA drones
should be less fit than colonies of African-mt-
DNA queens x European-mt-DNA drones.
The bronze color of the Ac drones may not be
the cause of its fitness, since the workers are yel-
low and heat preservation by dark color would be
better in temperate climate; the physiological rea-
son are being studied.
ACKNOWLEDGMENTS
We thank the FAPEMIG (Stateof Minas Gerais Research
Foundation) and CNPq (Brazilian National Research Coun-
cil) for financial help and Dr. Robert E. Page, Jr. forcorrecting
our English and for sound suggestions.
Volume 3, 1994
LITERATURE CITED
Del Lama, M. A., J. A. Lobo, A. E. E. Soares, S. N. Del Lama,
1990. Genetic differentiation estimated by isozymic
analysis of Africanized honeybee populations from Bra-
zil and from Central America. Apidologie 21: 271-280.
Hall, H. G. and K Muralidharan. 1989. Evidence from the
mitochondrial DNA that African honey bees spread as
continuous maternal lineages. Nature 339: 211-213.
Kerr, W. E. 1969. Genetica e melhoramento de abelhas. In:
Melhoramento e Genetica. Organized by W.E.Kerr, in
homage to Prof. F.G. Bneger. EDUSP, Melhoramentos,
USP, Sao Paulo. Cap. XIV pg. 263-295.
Kerr, W. E. 1992. Abejas africanas: su introduccion y expan-
sion en el continente americano. Subespecies y ecotipos
africanos. Industria Apicola N" 13: 12-21.
Kerr, W. E. and D. Bueno. 1970. Natural crossing between
Apis mellifera adansonii and Apis mellifera ligustica. Evolu-
tion 24(1): 145-148.
Kerr, W. E., L. S. Goncalves, L. F. Blotta and H. B. Maciel. 1970.
Biologia comparada entre abelhas italianas (Apis mellifera
ligustica), africana (Apis mellifera adansonii) e suas
hibridas. Anais do V Congresso Brasileiro de Apicultura
(Flonanopolis, SC) pg. 151-185.
Lobo, J. A., M. A. Del Lama, and M. A. Mestnner. 1989.
Population differentiation and racial admisture in the
Africanized honeybee (Apis mellifera L.). Evolution 43(4):
794-802.
Moretto, G., L. S. Goncalves and D. De Jong. 1991 . Africanized
bees are more efficient at removing Varroa jacobsoni.
Preliminary data. American Bee Journal. 131: 434.
Nascimento, A. F., Jr. 1981. Estudo da mfluencia de fatores
ambientais no comportamento enxameatono, migratono
e no desenvolvimento de colmeias africanizadas.
M.Sc. Thesis presented to the University of Sao Paulo at
Ribeirao Preto, Brazil.
Page, R. E, Jr., and W. E. Kerr. 1991 . Honey bee genetics and
breeding. 8th article of The "African" Honey Bee, pp 157
- 186. Ed. Maria Spivak, David J. C. Fletcher and Michael
D. Breed. Westview Studies in Insect Biology. Westview
Press, Boulder, USA.
Sheppard, W. S., A. E. E. Soares, D. De Jong and H. Shimanuki
1991. Hybrid status of honey bee populations near the
historic origin of afncanization in Brazil. Apidologia 22:
643-652.
Sheppard, W.S., T.E. Rinderer, J. A. Mazolli, J. A. Steiner and
H. Shimanuki. 1991. Gene flow between African and
European derived honey bee population in Argentina.
Nature 349: 782-784.
Soares, A.E.E. 1992. A utilizacao da Genetica molecular e da
morfometna na caractenzacao de populacoes de abelhas
africanizadas. Naturalia (Edic;ao Especial, 14 a 18 de
setembro de 1992) pg. 117-125.
Stort, A. C. 1978. Genetic study of the aggressiveness of two
subspecies of Apis mellifera in Brazil. VII. Correlation of
the various aggressiveness characters among each other
and with the genes for abdominal color. Ciencia e Cultura
30(4): 492-496.
Winston, Mark L. 1992. Killer Bee - The Africanized Honey Bee
in the Americas. Harvard University Press, London, En-
gland
Woyke, I. and W.E. Kerr. 1989. Linkage test between a sex
limited color gene and sex alleles in the honey bee.
Brazilian Journal of Genetics 12(1): 9-15.
Journal of Hymenoptera Research
Table 1. Data on swarms and hives at Uberlandia, Minas Gerais, Brazil, with reference to frequency of
Ac and ac alleles.
N°
Day
Arrival
Hour
Hive Volume
(Liters)
Frequency
Of Genes
Place
01
19.07.90
15:00
27
25 Ac
Kerr's House
02
05.08.90
11:00
27
16 Ac
Kerr's House
03
12.08.90
08:45
(1)(3)
10 Ac
Kerr's House
04
19.08.90
10:45
27
16 Ac
Kerr's House
05
11.09.90
12:30
27
20 Ac
Kerr's House
06
21.07.91
13:15
27
24 Ac
Kerr's House
07
14.08.91
14:20
27
15 Ac
Kerr's House
08
04.08.91
15:15
27
08 Ac
Kerr's House
09
05.08.91
14:28
(li
05 Ac
Kerr's House
10
08.08.91
15:00
27
10 Ac
Kerr's House
11
09.08.91
12:00
45
23 Ac
Kerr's House
12
14.08.91
14:20
27 (3)
14 Ac
Kerr's House
13
27.08.91
12:29
27
47 Ac
Kerr's House
14
24.11.91
(1)
15 (3)
34 Ac
Kerr's House
15
12.05.91
(2)
—
43 Ac
Alvorada Farm
16
11.08.91
(2)
—
23 Ac
Rio Pedras Farm
17
11.08.91
(2)
—
23 Ac
Rio Pedras Farm
18
11.08.91
(2)
—
23 Ac
Rio Pedras Farm
19
11.08.91
(2)
—
23 Ac
Alvorada Farm
20
11.08.91
(2)
—
23 Ac
Alvorada Farm
21
11.08.91
(2)
—
23 Ac
Sta. Rita Farm
22
06.08.91
(2)
—
24 Ac
Sta. Rita Farm
23
06.08.91
(2)
—
24 Ac
St\ Rita Farm
24
04.09.91
(2)
—
24 Ac
Alvorada Farm
25
04.09.91
(2)
—
23 Ac
Alvorada Farm
26
04.09.91
(2)
—
24 Ac
Rio Pedras Farm
27
06.08.91
(2)
—
24 Ac
Rio Pedras Farm
28
17.09.91
(2)
—
24 Ac
Alvorada Farm
29
17.09.91
(2)
—
23 Ac
Alvorada Farm
30
17.09.91
(2)
—
23 Ac
Alvorada Farm
31
17.09.91
(2)
34 Ac
690 Ac (96.23%)
27 ac (3.77%)
Rio Pedras Farm
(l)Hour
not noticed.
(2) Langstroth hive.
(3) Enormous swarm.
J. HYM. RES.
Vol. 3, 1994, pp. 5-16
A Survey of the Leaf Litter Ant Fauna in Ghana, West Africa
(Hymenoptera: Formicidae)
Robert Belshaw and Barry Bolton
Biodiversity Division, Department of Entomology, Natural Historv Museum, Cromwell Road, London SW7 5BD, U.K.
Abstract. — Leaf litter samples were taken from 34 sites scattered across the moist tropical forest zone in Ghana. They
included areas of primary forest, secondary forest and cocoa. Over 40,000 individual ants were extracted using Winkler bags
and identified. The species found are listed together with their abundance and a summary of their distribution. A total of 176
species was found (excluding stray workers of arboreal and surface-foraging species), almost two-thirds of which were
Myrmicinae. The composition of the fauna is discussed and compared with that found in other tropical forests The species
composition at the different sites showed little variation either between the different forest types or with geographic distance.
INTRODUCTION
In the West African forest belt there has been
little quantitative sampling of the ant fauna; stud-
ies have been carried out in the Tai Forest Reserve,
Cote d'lvoire (see Levieux 1982 and included ref-
erences) and the Reserve de Campo, Cameroun
(Halle and Pacal 1992: 65-109). In Ghana there has
been no quantitative sampling except in the main
tree-crop, cocoa. This has been studied in detail
(e.g. Majer 1976 and included references) but the
extent to which it resembles the original forest
fauna is not known. Nevertheless, this research
has led to the taxonomy of West African ants being
more advanced than that of most tropical ant
faunas (e.g. Bolton 1987 and included references).
In consequence we can survey elements of this
fauna with the hope of accurately identifying much
of it to species.
Ghana has two main terrestrial biomes, sa-
vannah and forest, and these have distinct ant
faunas. In turn, the forest zone is readily divisible
into a canopy and a ground fauna. In this paper we
survey the leaf litter element of the forest ground
fauna by sampling at different localities across
Ghana. In addition to identifying the species
present, we discuss the composition of the fauna
and compare it with that from other tropical for-
ests. We also examine how the species composi-
tion at the sites varies geographically and between
the different forest types.
Voucher specimens of all taxa are deposited in
the Natural History Museum, London.
METHODS
Sites. — The locations of the sampling sites in
Ghana are shown in Figure 1, with brief descrip-
tions and sampling dates given in Table 1. Sites
designated by the same letter but with different
numbers (e.g. jl and j2) are within 3km of each
other. With one exception, the sites are within the
moist semi-deciduous forest zone of Hall and
Swaine (1976).
We sampled in a wide range of the forest
habitats found in Ghana, including 14 areas of
primary forest (in the broad sense of forest with a
closed high canopy), 10 areas of secondary forest
(of varying age, mostly on agricultural land) and
10 cocoa farms. Sampling was carried out between
December 1991 and November 1992.
Sampling. — At each site an area of approxi-
mately 1000m: was measured out. Within this area
ten lm2 quadrats were placed at random. All the
leaf litter inside a quadrat was collected, shaken
through a 1cm sieve, and then left for three days in
a Winkler bag. The extracted ants were combined
to form a single total for each site, each site being
sampled on only one occasion. All sampling was
done between 9.30 a.m. and 3.00 p.m.
At three sites (h, ql and cj2) an additional soil
Journal of Hymenoptera Research
sample was taken from each quadrat. This was
done by collecting the soil from a 25cm by 25cm
quadrat (= 0.0625m2) to a depth of approximately
5cm. This soil was then sieved and left for three
days in Winkler bags in the same manner as the
overlying leaf litter.
The Winkler bag (Besuchet el al. 1987) oper-
ates in a similar fashion to a Berlese Funnel except
that the material is left hanging in a mesh bag to
dry in air rather than exposed to a heat source.
Winkler bags are much cheaper and easier to use
and to transport than Berlese Funnels. Litter-sift-
ing followed by extraction in Winkler bags records
many species which do not turn up in pitfall traps
(Olson 1991). Our extraction period of three days
was chosen on the basis of a trial extraction, with
daily sorting of a sample over a two week period.
We found that within the first three days 86% of
the individuals and 88% of the species had
emerged.
We ignored winged reproductives and wing-
less queens found without workers, except in spe-
cies where the queen is known to forage during
nest foundation.
Analyses. — In order to assess the complete-
ness of our survey for the region sampled, i.e. the
moist semi-deciduous zone plus disturbed habi-
tats within, we plotted a species accumulation
curve. We first arranged the sites in five random
sequences. In each sequence we calculated the
number of species found at the first site, the first
two sites combined, the first three sites combined,
and so on. Finally, the mean of the five sequences
was calculated. The extent to which the resulting
curve flattens out indicates the proportion of the
actual fauna which has been recorded; a failure to
flatten out indicates that additional species would
have been found if the sampling had been contin-
ued. Palmer (1990, 1991 ) compares and tests meth-
ods for estimating the species richness of a region
from samples taken within it. He concludes that
the first-order jacknife is the most precise method,
i.e. the one whose estimates are closest to the true
value, and we therefore also apply this analysis to
our data.
We converted the body lengths of each species
to biomass (= dry weight) using the following
equation, taken from Gowing and Recher (1984).
Logn weight(mg) = -4.0 + 2.5(logn length(mm))
Table 1. Sampling sites with dates and habitat description.
Sacred groves are small pieces of forest left in agricultural
areas for religious reasons. (Note that Ghanaian place name
spellings are often variable.)
a Sui River Forest Reserve, 1.x. 1992, primary forest,
b Mabang, 18.xii.1991, secondary forest,
c Tinte Bepo Forest Reserve, 31.iii.1992, primary forest,
d Mankrang Forest Reserve, 11. in. 1992, primary forest,
e Poano, 9.1x1992, cocoa,
f near Ofinso, 2.X1.1992, cocoa.
g Jachie, 20. iv. 1992, sacred grove (28 acres), primary for-
est,
h Effiduase, 17.xi.1992, cocoa.
i Bobiri. 1 - 6.1V.1992, Forest Reserve (primary forest); 2
8. iv. 1992, Forest Reserve (primary forest but all mature
trees killed with sodium arsenide in 1947); 3 - 13. iv. 1992,
secondary forest (farmland left in 1982 and burnt in
1983).
I Juaso. 1 - 21. ix. 1992, secondary forest (area of Dome
River Forest Reserve burnt in 1983); 2 - 23.ix.1992, sec-
ondary forest (farmland left for c.20 years); 3 - 8.ix.l992,
cocoa
k Southern Scarp Forest Reserve (North-West of Mpraeso
near Osubeng), 23.x. 1992, secondary forest (burnt in
1983).
1 Kade. 1 - 6.x. 1992, primary forest (in Aiyeola Forest
Reserve); 2 - 12.x. 1992, secondary forest (farmland left
in 1957); 3- 12.x. 1992, cocoa,
m Esukawkaw Forest Reserve, 27.x. 1992, primary forest,
n Nkawanda (near Nkawkaw), 12.xii.1991, roadside sec-
ondary forest,
o Atewa Forest Reserve, primary forest. 1 - 2.iii.l992, near
Kibi; 2 - 24.ih.1992, near Potrase; 3 - 26.h.l992, near
Sagymasi (logged in 1970's); 4 - 27. m. 1992, nearSagymasi
(logged in 1970's).
p Asiakwa, 1. v. 1992, cocoa.
q Bunso. 1 -6.xi.1992, Crops Research Institute arboretum
(15 acres), primary forest; 2 - 17.iv.1992, secondary for-
est (primary forest partially cleared ca 20 years previ-
ously, left undisturbed for ca 12 years); 3 - secondary
forest (cocoa left in 1981);4 -24.ii.1992, cocoa;5-6.iii. 1992,
cocoa,
r Old Tafo sacred grove (ca 3 acres), 31. i. 1992, primary
forest.
s New Tafo (Cocoa Research Institute of Ghana). 1 -
23.xii.1991, secondary forest (farmland left for en 40-50
years); 2 - ll.xii.1991, cocoa,
t Nankasi, 17.ix.1992, cocoa.
Body lengths (= the outstretched length of a
point-mounted worker including mandibles) were
taken from the literature or from an average of five
specimens. In species with a dimorphic worker
caste we did not count major and minor workers
separately. Instead we used an estimate of 30:1 for
the ratio of minor to major workers in all cases.
To assess the effect of the distance between
Volume 3. 1 994
1 1 1
IVORY r
-8° y
1 "\ \n
TOGO
coastt
\\ \ V ( »" V
/ DS •
!''• 6''
cl> 0 f; /
1 * - , ^ Sunyani
-7°/ b C
/ MS
Pvf ' /
(9 S- - - /
r < \ ' /
9^ > \ ' - \
\ N - - -a, , .
\ v Wiawso
-6° W
Kumasi ***. ^^
h ]
e Lake
Bosumtwi a
1
rs
^\ C /''Ho\
Koforidua ~~~X /
\ \ ME
Dunkwa % *'
Accra
WE \
\ i
,' ■' _ *Lape Coast
t J^ Takoradi C*J
O^
100 km
3° 2°
i°
0° 1°
1 "
Fig. 1 . Map of the forest zone of Ghana showing sampling sites (see table 1 for details). Forest and forest-type boundaries are
shown by dotted lines ( — ). Forest-type abbreviations: DS = Dry Semi-deciduous; MS = Moist Semi-deciduous; ME = Moist
Evergreen; WE = Wet Evergreen. From Hall and Swaine (1981).
sites on their species composition we first mea-
sured the similarity between all possible pairs of
sites using the Morisita Index (see Wolda 1981).
Their values were then plotted against the loga-
rithm of the distance between the sites, and a
regression line fitted. As the data points are not
independent, this procedure is of questionable
statistical validity; it does, however, allow us to
see the general pattern.
RESULTS AND DISCUSSION
We extracted 43,824 ants in 197 species and 47
genera. At least 17 of these species and two of the
genera were undescribed (one of these genera is
now described in Belshaw and Bolton (1994) and
the other is currently being described by Prof.
W.L. Brown under the name Loboponera). A further
57 species are in groups which lack keys or
synthesising taxonomic studies, and we can there-
fore neither identify them nor recognise them as
being undescribed. The species are listed in Table
2, along with the total number of individuals
found in each species and a summary of their
distribution among the sites.
Of the 197 species recorded, three were only
found in the soil samples and a further 18 are
known not to forage in the leaf litter and are
classed here as tourists. The latter are all arboreal
species except for Camponotus, which forage on the
surface of the ground. Although these tourists
s
Journal of Hymenoptera Research
comprised 9% of the species found, they com-
prised only 2% of the total number of individuals.
In the leaf litter the average density of indi-
viduals (excluding tourists) was 117m2. Using
Berlese funnels, other studies have found similar
densities of ants in leaf litter: in tropical moist
forest on Barro Colorado Island, Panama, in the
wet season it was ca 200m: (Levings 1983); in
temperate deciduous forest in Maryland, U.S.A.
between May and September the mean monthly
density was 194m 2 (Lynch and Johnson 1988).
Composition of the fauna. — The relative impor-
tance of the different subfamilies in the combined
soil and leaf litter samples is shown in Fig. 2.
The fauna is dominated by Myrmicinae. Where
their biology is known, the species we found are
all generalist predators / scavengers except for most
Dacetonini, the workers of which forage singly for
Collembola and other soft-bodied arthropods (us-
inghighly specialised mandibles), and Decamorium
decern (Forel), which has been observed attacking
termites (Bolton pers. obs.). Paedalgus distinctus
Bolton and Belshaw has also been found in asso-
ciation with termites (Bolton and Belshaw 1993,
see below).
The Ponerinae is the second most important
subfamily in our survey. Many ponerines are large
ants, so although the subfamily comprises only 8%
of the total number of individuals, it comprises
27% of the total biomass. A good example is
Paltothyreus tarsatus (Fabricius): only 25 individu-
als of this species were found during the survey,
but its contribution to the total ant biomass was
exceeded by only two other species. This subfam-
ily contains a high proportion of genera which are
specialised predators. In our survey we recorded
Ambh/opone (preys on geophilomorph centipedes),
Plectroctena macgeei Bolton (other members of the
genus prey on millipedes), Paltothi/reus tarsatus
(preys on termites), and Discothyrea (preys on
arthropod eggs) (all sources in Holldobler and
Wilson 1990: 559).
Driver ants (Aenictinae and Dorylinae) have
highly aggregated distributions - their colonies
are nomadic and very large (with between 60,000
and 20,000,000 workers in other species (Holldobler
and Wilson 1990: 581)). They are represented in
our survey by single workers found at three sites
and by almost 1500 workers from one soil quadrat
that hit part of a Dorylus nest. This group has
clearly not been adequately sampled in our sur-
vey, and it is probably a much more important
component of African forests than our results
indicate (see Holldobler and Wilson 1990: 588).
We found one worker of Apomyrma stygia
Brown, Gotwald and Levieux in damp leaf litter a
mile inside the Esukawkaw Forest Reserve. This is
the sole described species in the Apomyrminae
and was recorded previously only from one local-
ity in Cote D'lvoire (Brown etal. 1970), where four
nests were found in soil under gallery forest and
one under adjacent unburnt savannah. The spe-
cies is clearly subterranean (the workers are blind ),
and the remains of a geophilomorph centipede
were found in one of the Cote D'lvoire nests,
raising the possiblity that the species specialises
on this prey item.
Habitat and geograpliic variation. — In a separate
paper we have investigated in detail the differ-
ences in the ant assemblages between the primary
forest, secondary forest and cocoa sites in this
study (Belshaw and Bolton 1993). We found that
they did not differ significantly either in species
composition or in species richness. Only two com-
mon species show an association with a particular
forest type (see Table 2): Serrastruma hijae (Forel)
was with the exception of a single individual
found only in the Esukawkaw and Atewa Forest
Reserves, and one Oligomyrmex species (sp.indet.4)
mostly occurred in cocoa. Given the large number
of species present, one would predict that, even if
all species were randomly distributed among the
three forest types, a handful of such apparent
associations would occur purely from chance.
The slope of the regression line in Fig. 4 is very
shallow, and the (logarithmically-transformed)
distance between sites only accounts for 1.9% of
the variation in similarity. The distance between
sites therefore had at the most only a slight effect
on species composition.
Comparison with other faunas. — Several other
studies of tropical forest ant faunas contain lists of
leaf litter species: dry forest at Kimberley, N. Aus-
tralia (Andersen and Majer 1991); moist forest on
Barro Colorado Island, Panama (Levings 1983);
and dry forest in Madagascar (Olson and Ward, in
press). Following elimination of obvious arboreal
species, these studies all show the Myrmicinae to
be the largest subfamily, followed by the Ponerinae
Volume 3, 1994
100
80
o
c
o
60
40
20
0
[1 Biomass
■ Individuals
D Species
ii
L
nl ^1 nL, 11 J I 1 In ^1
'V*
Subfamily
Fig. 2. Relative importance of the different ant subfamilies in the total fauna (leaf litter and soil samples combined, tourists
excluded). Names are abbreviated (see table 2).
(in our study comprising 63% and 22% of the total
number of species respectively). They comprised
52% and 30% at Kimberley (33 species in total),
65% and 27% on Barro Colorado Island (93 species
in total), and 55% and 20% in Madagascar (44
species in total). The figures in the last study
would have been higher but for the unusually
large number of Cerapachys species. The absence of
doryline and aenictine ants from the island has
possibly permitted diversification in this genus of
specialised predators of other ants (Wilson 1971:
68). Unfortunately, it is not possible to compare
the species richness of the four areas owing to the
differences in sampling area, effort and method
(Berlese funnel, Winkler bag and pitfall trapping).
10
Journal of Hymenoptera Research
■M
e
u
200
o
o, 150
Cm
O
,0
a
100
50
0
0
10
20
30
40
Number of sites
Fig. 3. Species accumulation curve for the leaf Utter samples. Line fitted by DWLS smoothing option of computer program
SYSTAT (Wilkinson 1990).
Efficiency of survey. — The species accumula-
tion curve is shown in Fig. 3. If the fitted line is
extrapolated beyond the data it does not continue
to rise. Estimating the true regional species rich-
ness using the first-order jacknife method, we find
our total of 176 species represents 81% of the actual
species present. We infer from this that, within the
area of Ghana sampled, we recorded a large ma-
jority of the species foraging within leaf litter.
In tropical forests the majority of ground ant
species nest in the leaf litter, either in small pieces
of rotting wood or between compressed leaves
(Wilson 1959; Bolton pers. obs.). In addition, some
other species nest in the top l-2cm of the soil but
forage in the leaf litter. However, there are other
smaller elements of the ground fauna which, be-
cause of their nesting or foraging habits, are likely
to have been missed by our sampling method.
1) Completely subterranean species (= ones
which nest and forage only in the soil). In Ghana
only a few such species are known, e.g. Plectroctena
anops Bolton and P. hastifera (Santschi) (the work-
ers of which are either blind {anops) or with very
small eyes {hastifera)). However, this microhabitat
Volume 3. 1994
11
s
75
1.0
X
0.8
<D
•d
c
1— 1
«
06
l/l
•*H
«H
0
>> 0.4
£ 0.2
0.0
0
1
Log10 (distance (km) + 1)
Fig. 4. Plot of similarity in species composition against distance for all possible pairwise comparisons between sites. Line fitted
by linear regression (y = 0.392 - 0.060x), r2 = 0.019.
has to date hardly been sampled at all, and its
fauna is not understood.
2) Species which nest in rotting logs. Such
species usually also restrict their foraging to this
microhabitat (Wilson 1959). For example, we failed
to find Plectroctena minor Emery, which is not
uncommon in eastern Ghana where it is usually
found in or under rotten logs (Bolton 1974). Many
Tetramoriini also nest in rotting logs (see Bolton
1980, and below).
3) Obligatory termitolestic species (= species
which nest and forage only within termitaria).
This is a small group: in Ghana the habit is con-
firmed only in species of Carebara and
Centromyrmex. We found Paedalgus distinctus and
P. saritus Bolton and Belshaw foraging in leaf litter
which indicates that, despite observations on other
members of the genus (see Bolton and Belshaw
1993), they do not belong in this group.
4) Species which only forage nocturnally are
likely to be under- rep resented. In Ghana, among
the ground fauna, this habit appears to be re-
stricted to species of Leptogenys and Camponotus.
As expected, we found only one of the 12 species
12
Journal of Hymenoptera Research
of Leptogem/s previously recorded from Ghana
(Bolton 1975), plus one undescribed species. The
genus Camponotus is poorly known taxonomically,
but there are more than 15 species from Ghana in
the collection of the Natural History Museum
which we did not find. Non-arboreal members of
this genus, although nesting in the soil, are fast-
moving surface foragers, which may also have
contributed to their poor representation in our
survey.
The three largest myrmicine tribes are suffi-
ciently well known taxonomically for us to com-
pare the list of species recorded in our survey with
that of species already known from Ghana. In the
Dacetonini, 25 species were previously known
(Bolton 1983), including two known to be arbo-
real. We found 18 of the remaining 23 species, plus
14 previously unrecorded in Ghana. In the
Solenopsidrni, 15 species were known (Bolton
1987), including three savannah species. We found
10 of the remaining 12 species, plus six previously
unrecorded . These results show that we have found
many small cryptic species which had previously
been overlooked. The world-wide household pest
Monomorium pharaonis (L.) (Pharaoh's Ant) was
recorded from nine sites (including primary for-
est). Its range is clearly not restricted to disturbed
habitats in Ghana.
In contrast, we found a smaller proportion of
the Tetramoriini species known from Ghana. Pre-
viously 53 species were known (Bolton 1976, 1980),
including 10 savannah and three arboreal species.
We found only 21 of the remaining 40 species, plus
six previously unrecorded in Ghana. Members of
this tribe, however, often nest in exposed sites or in
rotten logs, which may explain its relatively poor
representation in our survey.
We cannot compare our results for the
Pheidologetonini, Pheidolini or Crematogastrini
as no synthesising taxonomic studies have been
undertaken. The remaining myrmicine tribes con-
tain a total of 40 species previously recorded from
Ghana (Bolton 1981a, 1981b, 1982) including 20
arboreal (chiefly Cataidacus), four savannah, and
two species which tunnel in living wood
(Melissotarsus). We found 8 of the remaining 14
species: one of the three Cardiocondyla, one of the
three Leptothorax, two of the four Calyptomyrmex,
both Pristomyrmex, the sole Meranoplus and the
sole Baracidris. In most cases nothing is known of
the ecology of the species not found in our survey
except for earlier collection localities and dates.
However, their absence from our survey shows
that they either do not forage in the leaf litter or are
rare.
The soil samples. — The density of individuals
was eight times higher in the soil samples than in
the leaf litter (mean of the three soil samples = 982
m 2 (S.E.= 62). Lynch and Johnson (1988) also found
that in a temperate forest the density of ants was
higher in the soil than the leaf litter. These data
may, however, be misleading as in both studies
the soil samples probably included species which,
although nesting in the top few cm of the soil,
forage primarily in the leaf litter.
Although similar in overall species composi-
tion to the leaf litter samples (also found by Lynch
and Johnson 1988), the three soil samples con-
tained a small distinctive subterranean element.
We extracted 30 workers of Leptanilla boltoni Baroni
Urbani from a single soil quadrat in a small patch
of primary forest at Bunso. The sole Afrotropical
representative of the genus, it is known previously
only from 6 specimens extracted by Berlese funnel
from leaf litter in a cocoa plantation at Mampong,
Ghana (Baroni Urbani 1977). The genus contains a
total of 33 described species, all of which appear to
be very rare (Holldobler and Wilson 1990: 590).
The biology of only one species (L. japonica Baroni
Urbani) is known: it is strictly subterranean and
appears to have true legionary behaviour, em-
ploying both group predation (of geophilomorph
centipedes) and colony migration (Masuko, 1990).
Of the 17 individuals of Acropyga sp., 10 were
found in soil samples. This species resembles oth-
ers in the genus which tend coccids in subterra-
nean nests for their sugary secretions (Holldobler
and Wilson 1990: 527). It is the only species found
in our survey which is known not to be primarily
predacious, although the habit probably occurs in
species of several other formicine genera
(Pseudolasius, Paratrechiiia and Lepisiota). One of
the species of the undescribed Ponerini genus also
appeared to be soil-dwelling.
The results of our few soil samples point
tantalisingly to a rich and poorly known subterra-
nean fauna that would repay further collecting.
Volume 3, 1994
13
Table 2. Species recorded during the survey (see table 1 for details of sites, and figure 1 for their location). "Widespread"
denotes the species was recorded from more than five differently-lettered sites (the number is given in parentheses). An
asterisk (*) denotes species classed as tourists (see text).
Species
Total number
() = from
soil samples
Sites
Species
Total number
() = from
soil samples
Sites
AENICTINAE
OECOPHYLLINI
Aenictus sp. (indet) (1)
1
si
"Oecophylla longinoda
15
d,i3,j3,p
A. sp. (indet) (2)
1
o3
PLAGIOLEPIDINI
APOMYRMINAE
Acropyga sp. (indet.)
17(10)
ai.h.ql
Apomyrma stygia
1
m
Lepisiota sp. (indet.) (1)
4
f,sl
L. sp. (indet.) (2)
2
o3
L. sp. (indet.) (3)
1
o4
CERAPACHYINAE
L. sp. (indet.) (4)
10
f,q5
Cerapachvs foreh
55
widespread (1 1)
L. sp. (indet.) (5)
1
qs
C. nitidulus
299(134)
widespread (12)
L. sp. (indet.) (6)
1
q5
C. n.sp.
3
m,o2,sl
Plagiolepis sp. (indet.)
LEPTANILLINAE
19
d
DOLICHODERINAE
Leptanilla boltoni
30(30)
qi
Tapinoma sp. (indet.)
2
r
Technomyrmex sp. (indet.) (1) 821(2)
widespread (14)
MYRMICINAE
T. sp. (indet.) (2)
163(8)
widespread (11)
CATAULACINI
T. sp (indet.) (3)
3
si
'Cataulacus guineensis
1
d
T. sp (indet.) (4)
3
d,p
T. sp. (indet.) (5)
530
widespread (10)
CREMATOGASTR1NI
*Crematogaster depressa
499
widespread (6)
*C. bequaerti
9
o2,p,q2
DORYLINAE
*C. clariventris
24
p,q4
Dorvlus sp. (indet.)
1477(1468)
12,ql
C. stnatula
*C. sp. (indet. ) (1)
2210
1
widespread (13)
d
C. sp. (indet.) (2)
1
s2
FORMICINAE
C. sp. (indet.) (3)
1
o3
CAMPONOT1NI
C. sp. (indet.) (4)
9
o3
"Camponotus acvapimensis
2
q4-5
*C. sp. (indet.) (5)
103
i3
*C. flavomarginatus
15
q4-5,s2
*C. maculatus
2
qs-t
DACETON1NI
*C. vividus
2
qs
Epitntus laticeps
27
k,ll-3,m,q4
"Polyrhachis decemdentata
2
q4,r
E. roomi
53(16)
widespread (12)
*P. militans
24
p,q2
E. tiglath
9
d,f,k,Il-2
*P. rufipalpis
2
a,o4
Glamyromyrmex crvpturus
57(5)
f,il,o3,q2-3,sl
*P. weissi
1
r
G. sistrurus
110
widespread (6)
G tetragnathus
31
widespread (6)
LASIINI
G tukultus
20
e.ql
Paratrechina sp. (indet.) (1)
794
widespread (12)
Microdaceton tibialis
53
widespread (7)
P. sp. (indet.) (2)
48
d,i3,jl,m,q4-5
Quadnstruma emmae
1
n
P. sp. (indet.) (3)
148(48)
h,i2-3,q2,t
Serrastruma concolor
238
widespread (8)
P. sp. (indet.) (4)
1
P
S. ludovici
20
g,12,o2,t
Pseudolasius sp. (indet.)
270(4)
t,il,o3,ql-3
S lujae
802
j3,m,ol-4
S. serrula
823(4)
widespread (16)
Smithistruma cavinasis
3
11
S. enkara
7
a,q2
14
Journal of Hymenoptera Research
Species
Total number
() = from
soil samples
Sites
Species
Total number
() = from
soil samples
Sites
S. fulda
2
a
P. sp. (indet.) (3)
180(74)
d,g,i2,p,q2
S. hensekta
58
widespread (9)
P. sp. (indet.) (4)
67
d,q5,s2
S. malaplax
69
sl-2
P. sp. (indet.) (5)
213(1)
widespread (9)
S. marginata
1
d
P. sp. (indet.) (6)
10
a,o4,p,t
S. minkara
55
f,12,o3,q3
P. sp. (indet.) (7)
98
o4
S. ninda
55
f,h,n,ql+5
P. sp. (indet.) (8)
8
ol,h
S. sharra
25
d,f,ll,o2-3
P. sp. (indet.) (9)
496
widespread (7)
S. facta
56
widespread (8)
P. sp .(indet.) (10)
_1
e,i3,q4
S. tignlla
9
d,ll,o2
P. sp (indet.) (11)
30
a
S. n.sp.
4
h,q5
Strumigenys korahvla
1
d
SOLENOPSIDINI
S- nimbrata
70(4)
widespread (7)
Monomonum bicolor
1
d
S. petiolata
1439(31)
widespread (20)
M. cryptobium
1475(242)
widespread (16)
S. rogen
177
widespread (6)
M. draxocum
5
d,g
S. rufobrunea
1059
widespread (15)
M. exiguum
605
widespread (16)
S. spathoda
KD
^
M. floncola
13
13
S. vazerka
22(1)
a,13,o2,ql+3,s2
M. egens
8
widespread (7)
M. gabnelense
42
a,b,g,o2-3,t
FORMICOXENINI
M guineense
55
g.'2
Cardiocondyla neferka
1
d
M. invidium
3432(62)
widespread (19)
Leptothorax angulatus
1
qs
M. pharaonis
17
widespread (6)
"Terataner piceus
1
n
M. rosae
7
f,q4-5
M. tanysum
2
d
MERANOPLINI
M. trake
10
d,i2
Meranoplus inermis
1
si
M. n.sp. (1)
22
d
M. n.sp. (2) (exiguum group) 13
p,q5
MYRMECININI
M n.sp. (3)
75
i2,jl+3,p
Pristomyrmex afncanus
79(3)
widespread (9)
P. orbiceps
505(3)
widespread (13)
STENAMMINI
Baracidns meketra
8(3)
j3,12,m,n,ql-2
PHEIDOLOGETONINI
Calyptomyrmex kaurus
18
k,ll-2,o3
Afroxyidris cngensis
2
e,f
C. nummuliticus
1 1
12,o3,p,q3
Oligomyrmex sp. (indet.) (1) 440(7)
widespread (10)
O. sp. (indet.) (2)
1957(333)
widespread (15)
TETRAMORIINI
O. sp- (indet.) (3)
1532(175)
widespread (12)
Decamonum decern
119
widespread (7)
O. sp. (indet.) (4)
123(10)
e.j3,13,q2+5,s2,t
"Tetramonum aculeatum
3
ol+4,i3
O. sp. (indet.) (5)
245
widespread (6)
T. amentete
106
widespread (8)
O. sp. (indet.) (6)
967(81)
widespread (13)
T. antrema
162
widespread (10)
O. sp. (indet.) (7)
98(92)
ql-2+5
T ataxium
140(4)
widespread (11)
O. sp. (indet ) (S)
602
widespread (6)
T brevispinosum
118
widespread (6)
O. sp. (indet.) (9)
156(34)
ql,o3
T. camerunense
2
ol
O. sp. (indet.) (10)
197
g,jl+3,m
T. distincrum
552(16)
widespread (16)
O. sp. (indet.) (11)
39(27)
h
T flavithorax
350
widespread (17)
O. sp. (indet.) (12)
104
ll-3,q3
T. furtivum
689(33)
widespread (8)
Paedalgus distinctus
228(22)
widespread (13)
I guineense
417
widespread (10)
P. saritus
42
|3,n,o2
T ictidum
1
o2
T. invictum
65
widespread (6)
PHEIDOLINI
T. |ugatum
4
d,12,t
Pheidolesp. (indet.) (1)
778
widespread (10)
T. lucayanum
3
e,13
P. sp. (indet ) (2)
5474(94)
widespread (20)
T. menkaura
21
d,o2-4,sl,t
Volume 3, 1994
15
Species
Total number
() = from
soil samples
Sites
T. minimum
368(25)
widespread (15)
T. muralti
189
widespread (10)
T. muscorum
18
widespread (6)
T. peutli
28
widespread (6)
T. quadndentatum
1
q4
T. rhetidum
971(6)
widespread (9)
T. youngi
7
o2-3
T. zambesium
2293(13)
widespread (17)
T. zapyrum
33
d,r,t
T. n.sp. (weitzeckeri
group;
30
H,o4,q2
T. n.sp. (convexum grou
P)
5
i3,m
T. n.sp. (dumezi gro
up)
2
q3
Species
Total number
() = from
soil samples
Sites
PONERINAE
AMBLYOPONINI
Amblyopone mutica
3
i2,t
A. santschii
6
f,g,jl+3,ql
ECTATOMMINI
Discothyrea oculata
1
q2
D. mixta
3
f,ll,o4
D. n.sp (1)
5(1)
f,ll,o4,ql
D. n.sp (2)
3
a
PLATYTHYREINI
•Platythyrea conradti
1
r
Probolomyrmex guineensis
46
f,k,ll-3,sl
PONERINI
Anochetus afncanus
170
g,ll-3,ol+3-4,p,ql-3
A. bequaerti
43(2)
widespread (9)
A. katonae
568(70)
widespread (17)
A. maynei
1
si
A. siphneus
6
f,h,]3,t
A. n.sp.
4
o3-4
Asphinctopone silvestni
5(1)
f,ql-3
Cryptopone n.sp.
2
jl
Hypoponera sp. (indet.) (1)
244(29)
widespread (13)
H. sp. (indet.) (2)
1828(35)
widespread (19)
H. sp. (indet.) (3)
11
d,g,k
H. sp. (indet.) (4)
98(16)
widespread (6)
H. sp. (indet.) (5)
223(2)
widespread (9)
H. sp. (indet.) (6)
6(1)
i3,ol,q2
Leptogenys nuserra
15(1)
13,m,ol-2,q2,t
L. n.sp.
1
o2
Odontomachus assiniensis
8
si
O. troglodytes
22
widespread (7)
Pachycondyla ambigua
24
a,d,f,12,ol-2+4
P. brunoi
24(9)
widespread (8)
P. caffraria
86
widespread (8)
P. fugax
12
h,il,12,o3,t
P. pachyderma 16 widespread (7)
P. soror 2 ]3,q2
P. sp. (indet.) 18(7) p,ql+3,sl
P. n.sp. 1 i3
Paltothyreus tarsatus 25 widespread (7)
Phrynoponera bequaerti 13 11+3
P. gabonensis 24(1) widespread (6)
Plectroctena macgeei 3 jl,o3
n. gen., n.sp. (1) 3(3) q2,h
n. gen., n.sp. (2) 1 13
ACKNOWLEDGMENTS
This project was funded by the Leverhulme Trust (Grant
F.696A). The field work was based at the Cocoa Research
Institute of Ghana, New Tafo, and we are very grateful for
their hospitality and assistance. We would also like to thank
the Ghanaian Forestry Department; Cocoa Services Division;
Crops Research Institute; the Ghana Association for the Con-
servation of Nature; and the Tafohene, Old Tafo.
In addition, Paul Eggleton and two reviewers made
valuable comments on the manuscript.
LITERATURE CITED
Andersen, A. N. and J. D. Ma]er. 1991. The structure and
biogeography of rainforest ant communities in the
Kimberley region of north-western Australia, pp. 333-
346. In McKenzie N.L..R.B. Johnston and P.G. Kendrick,
eds. Kimberley Rainforests of Australia. Surrey Beatty and
Sons, Chipping Norton, Australia. 490 pp.
Baroni Urbani, C. 1977. Materiah per una revisione della
sottofamiglia Leptanillinae Emery (Hymenoptera:
Formicidae). Entomologica Brasiliensia 2: 427-488.
Belshaw, R. and B. Bolton. 1993. The effect of forest distur-
bance on the leaf litter ant fauna in Ghana. Biodiversity
and Conservation 2: 656-666.
Belshaw, R. and B. Bolton. 1994. A new myrmicine ant genus
from cocoa leaf litter in Ghana (Hymenoptera:
Formicidae). Journal of Natural History 28: 631-634.
Besuchet, C, D. H. Burckhardt and I. Lobl. 1987. The
"Winkler/Moczarski" eclector as an efficient extractor
for fungus and litter Coleoptera. The Coleopterists Bulle-
tin 41: 392-394.
Bolton, B. 1974. A revision of the ponenne ant genus
Plectroctena F. Smith (Hymenoptera: Formicidae). Bulle-
tin of the British Museum (Natural History) (Entomology)
30: 309-338.
Bolton, B. 1975. A revision of the ant genus Leptogenys Roger
(Hymenoptera: Formicidae) in the Ethiopian region:
with a review of the Malagasy species. Bulletin of the
British Museum (Natural History) (Entomology) 31: 235-
305.
L6
Journal of Hymenoptera Research
Bolton, B. 1976. The ant tribe Tetramornni (Hymenoptera:
Formicidae): constituent genera, review of smaller gen-
era and revision of Triglyphothrix Forel. Bulletin of the
British Museum (Natural History) (Entomology) 34: 283-
379.
Bolton, B. 1980. The ant tribe Tetramornni (Hymenoptera:
Formicidae): the genus Tetramorium Mayr in the Ethio-
pian zoogeographical region. Bulletin of the British Mu-
seum (Natural History) (Entomology) 40: 193-384
Bolton, B. 1981a. A revision of the ant genera Meranoplus F.
Smith, Dieroaspts Emery and Calyptomyrmex Emery (Hy-
menoptera: Formicidae) in the Ethiopian zoogeographi-
cal region. Bulletin of the British Museum (Natural His-
tory) (Entomology) 42: 43-81.
Bolton, B. 1981b. A revision of six minor genera of My rmicmae
(Hymenoptera: Formicidae) in the Ethiopian region.
Bulletin of the British Museum (Natural History) (Entomol-
ogy) 43: 245-307.
Bolton, B. 1982. Afrotropical species of the myrmicine ant
genera Cardiocondyla, Leptothorax, Melissotarsus, Messor
and Cataulaeus (Formicidae). Bulletin of the British Mu-
seum (Natural History) (Entomology) 45: 307-370.
Bolton, B. 1983. The Afrotropical dacetine ants (Formicidae)
Bulletin of the British Museum (Natural History) (Entomol-
ogy) 46: 267-416.
Bolton, B. 1987. A review of the Solenopsis genus-group and
revision of Afrotropical Monornorium Mayr (Hy-
menoptera: Formicidae). Bulletin of the British Museum
(Natural History) (Entomology) 54: 263-452.
Bolton, B. and R. Belshaw. 1993. Taxonomy and biology of the
supposedly lestobiotic ant genus Paedalgus (Hym.:
Formicidae). Systematic Entomology 18: 181-189.
Brown, W. L., W. H. Gotwald Jr. and J. Levieux. 1970. A new
genus of ponerine ants from West Africa (Hvmenoptera:
Formicidae) with ecological notes. Psyche 77: 259-275.
Gowing, G. and H. F. Recher. 1984. Length-weight relation-
ships for invertebrates from forests in south-eastern
New South Wales. Australian Journal of Ecology 9: 5-8
Hall, J. B. and M. D. Swaine. 1976. Classification and ecologv
of closed-canopy forest in Ghana. Journal of Ecology h4
913-951.
Hall, J. B. and M. D. Swaine. 1981. Distribution and ecology
of vascular plants in a tropical rain forest: forest vegeta-
tion in Ghana. Geobotany 1: 1-383.
Halle, H and O. Pacal, eds. 1992. Biologie d'une canopee deforet
equatoriale: volume 2. Unpublished report.
Holldobler, B. and E.O. Wilson 1990 The ants Springer-
Verlag, Berlin. 732 pp.
Levieux, J. 1982. A comparison of the ground dwelling ant
populations between a guinea savanna and an ever-
green rain forest of the Ivory Coast, pp. 48-53. In Breed,
M.D., CD. Michener, and HE. Evans, eds. The biology of
social insects. Westview Press, Boulder, Colorado. 419
PP-
Levings, S. C. 1983. Seasonal, annual, and among-site varia-
tion in the ground ant community of a deciduous tropi-
cal forest: some causes of patchy species distributions.
Ecological Monographs 53: 435-455.
Lynch, J. F. and A. K. Johnson. 1988. Spatial and temporal
variation in the abundance and diversity of ants (Hy-
menoptera: Formicidae) in the soil and litter layers of a
Maryland forest. The American Midland Naturalist 119:
31-44.
Majer, J. D. 1976. The influence of ant and ant manipulation
on the cocoa farm fauna. Journal of Applied Ecology 13:
157-175.
Masuko, K. 1990, Behavior and ecology of the enigmatic ant
Leptamlla japonica Baroni Urbani. hisectes Socwux 37: 31-
57.
Olson, D. M. 1991. A comparison of the efficacy of litter
sifting and pitfall traps tor sampling leaf litter ants
(Hvmenoptera, Formicidae) in a tropical wet forest,
Costa Rica. Biotropica 23: 166-172.
Olson, D. M. and P. S. Ward, in press. The ant fauna (Hy-
menoptera: Formicidae) of Kinndy Forest (tropical dry
forest) in Western Madagascar. In Ganzhorn, J., ed. The
ecology and economy of a tropical dry forest. Springer Verlag,
Berlin
Palmer, M. W. 1990. The estimation of species richness by
extrapolation. Ecology 71: 1195-1198.
Palmer, M. W. 1991. Estimating species richness: the second
order jacknife reconsidered. Ecology 72: 1512-1513.
Wilkinson, L. 1990. SYSTAT: the system for statistics- Evanston,
IL: SYSTAT, Inc.
Wilson, E. O. 1959. Some ecological characteristics of ants in
New Guinea rain forests. Ecology 40: 437-447.
Wilson, E. O. 1971. The insect societies Belknap Press, Cam-
bridge, Massachusetts.
Wolda, H. 1981 . Similarity indices, sample size and diversity.
Oecologia 50: 296-302.
J. HYM. RES.
Vol. 3, 1994, pp. 17-89
Revision of the Ant Genus Rogeria (Hymenoptera: Formicidae)
with Descriptions of the Sting Apparatus
Charles Kugler
Biology Department, Radford University, Radford, VA 24142 USA
Abstract. — This is the first complete revision of the century-old ant genus Rogeria. The revision recognizes 39 species, of
which 19 are new species: alzatei n. sp. (Panama, South America, Dominican Republic), besucheti n. sp. (Paraguay, Colombia,
Peru), carinata n. sp. (Caribbean), ciliosa n. sp. (Venezuela, Ecuador), cornuta n. sp. (Belize, southern Mexico), cuneola n. sp.
(Mexico, Central America), gibba n. sp. (Colombia, Ecuador), innotabilis n. sp. (Mexico, Central America), leptonana n. sp.
(Mexico, Central America, Dominican Republic) hrata n. sp. (Trinidad, South America), megastigmatica n. sp. (Solomon
Islands), merenbergiana n. sp. (Colombia, Ecuador), neilyensis n sp. (Costa Rica), nevadensis n. sp. (Colombia), prommula n. sp.
(Brazil), scobinata n. sp. (Trinidad, South America), terescandens n. sp. (Costa Rica), tribrocca n. sp. (Colombia), and unguispina
n. sp. (Venezuela). Previously recognized species are belli Mann 1922, blanda Fr. Smith 1858, bruclu Santschi 1922, brunnea
Santschi 1930, creightoni Snelling 1973, curvipubens Emery 1894, exsulans Wilson and Tavlor 1967, foreli Emery 1894, germaim
Emery 1894, inermis Mann 1922, lacertosa Kempf 1963, micromma Kempf 1961, minima Kusnezov 1958, pellecta Kempf 1963,
procera Emery 1896, scandens Mann 1922, sicaria Kempf 1962b, stigmatica Emery 1897, subarmata Kempf 1962a, and tonduzi Forel
1899. Nine new synonyms are proposed: caraiba Santschi 1936, cubensis Santschi 1936, habanica Santschi 1936, and scabra Weber
1934 = brunnea Santschi 1930; gaigei Forel 1914, and huachucana Snelling 1973 = foreh Emery 1894, minensis Santschi 1923 =
germaini Emery 1894; sublei'inodis Emery 1914, and manni Santschi 1922 = stigmatica Emery 1897. Seven new combinations are
proposed: epmotalis Mann 1919, rugosa Mann 1921, striatella Mann 1921, tortuosa Mann 1921, tortuosa levifrons Mann 1921,
tortuosa pohta Mann 1921, and tortuosa stonert Mann 1925 to Lordomyrma Three species occur in Polynesia and Melanesia; the
rest are New World: southwestern United States to southern Paraguay A key to the workers is provided All species are briefly
described and illustrated to show variation. Available biological information is summarized. Six informal species groups are
defined for 29 species; possible affiliations of the remaining 10 species are indicated. A new character system, the sting
apparatus, is used to help define the species and species-groups. Pilositv also provides an important new set of characters for
species identification in this genus.
INTRODUCTION AND DISCUSSION
This revision will redefine the genus, establish
new synonymies, create informal species groups,
redescribe species, provide a key to the workers,
and summarize ecological information. Keys to
the reproducti ves are not included, because queens
are known or proposed for less than half of the
Rogeria species and males for only four species.
The genus Rogeria was erected by Emery ( 1 894 )
for a few neotropical ants then in Myrmica and
Tetramorium. He then extended the distribution of
the genus to New Guinea with the addition of
stigmatica (Emery 1897). Discovery of other
neotropical species led Emery (1915) to create the
subgenus Irogera for those with a clavate petiole.
This subdivision was attractive, and for a brief
period Irogera even enjoyed generic status (Brown
1953), but with a better understanding of the varia-
tion in petiole shape within species, the splitting of
Rogeria along these lines became untenable ( Kempf
1965). Meanwhile, Melanesian and Polynesian
species were being added to the genus with no
discussion of the disjunct distribution of the genus
or comparisons of New World and Old World
species (Mann 1919, 1921, 1925; Santschi 1922,
194 1 ). Brown (1953) recognized that many of these
were actually Lordomyrma species, but still, the
Pacific species including stigmatica, epinotalis, and
later, exsulans remained in Rogeria. Wilson and
Taylor (1967) called this "...one of the most anoma-
lous discontinuous distributions found in ants."
18
Journal of Hymenoptera Research
The disjunct distribution of Rogeria was ques-
tioned by Kusnezov (1958) on the basis of palpal
formula differences between stigmatica and a South
American species, and by Kugler (1978b) on the
basis of the sting apparatus. In contrast to all other
myrmicine genera, there were two distinctly dif-
ferent sting apparatuses within the genus, and
that difference coincided with the disjunct distri-
bution of the genus. The sting of inermis from
Central America was strong, acute and with an
unusual low dorsal flange; the sting of stigmatica
from the Pacific had a weak, narrowly spatulate
sting shaft and no dorsal flange (among other
differences). I believed a revision of Rogeria using
sting apparatus characters would separate the
Pacific and New World species into distinct gen-
era.
The potential of the sting apparatus for defin-
ing ant genera and inferring phylogeny has been
demonstrated by a number of comparative mor-
phological studies (Kugler 1978b, 1980, 1986, 1992).
In addition, Bolton (1973, 1982, 1987) has often
used externally visible parts of the sting to help
define some myrmicine taxa. But this is the first
time that characters from the whole apparatus
have been used as an integral part of a taxonomic
revision. This work, then, also tests the usefulness
of the sting apparatus in the alpha-taxonomy of
ants.
I examined sting apparatuses of 79 workers
and queens in 27 Rogeria species and seven
Lordomyrma species. In the five species in which
both queens and workers were dissected, there
were only the usual individual differences, such as
numbers of sensilla. Contrary to expectations, the
Western Samoan species exsulans is unmistakably
related to Central American Rogeria species in
sting apparatus, pygidium, and external anatomy.
Moreover, some South American species (ciliosa,
gibba, besucheti) had sting apparatuses and pygidia
with the same distinctive features of stigmatica. It
seems the distribution of Rogeria really is disjunct.
Sting and pygidial characters provided sup-
port for numerous other taxonomic decisions as
well. For example, they provided strong support
for creating the stigmatica-group of species, and
could conceivably be used to distinguish that group
as a separate genus. Sting apparatus anatomy also
supported Brown's (1953) belief that levifrons,
striatella, and tortuosa are Lordomyrma species.
Autapomorphic shape of the sting shaft confirmed
that germaini and lirata are sister species. The un-
usual shape of the spiracular plate ultimately con-
vinced me to propose innotabilis as a new species
after long deliberation over the welter of variation
within the creightoni-gioup.
This character system must be used with caution
however. The apparatus is prone to reduction and
convergence of form. The sting of Lordomyrma
epinotalis is more like those of the Rogeria stigmatica-
group than those of Lordomyrma. In this case, I
based my decision to transfer epinotalis to
Lordomyrma on external characters, since the com-
mon sting features could well be due to reduction
convergence.
Hair has not been used previously in Rogeria
systematics, but in this revision I came to rely more
and more on characteristics of pilosity for distin-
guishing species. Consequently it has become nec-
essary for species descriptions to identify pre-
cisely the several kinds of hairs and their distribu-
tions on the ants, sometimes including the num-
bers of pairs of erect hairs on the head, mesosoma,
and nodes. This means, of course, that care must
be taken to not disturb the hair while cleaning and
mounting the ants.
I have attempted to take a fairly conservative
approach to synonymy and description of new
species, choosing in ambiguous situations to err
on the side of not changing the number of species
in the genus. Thus, for lack of sufficient evidence,
some species may actually contain several sibling
species: alzatei, belli, creightoni, foreli, leptonana, and
scandens. On the other hand, I have retained some
names that may fall when collections improve:
brnchi,micromma. Treatment of the creightoni-group
has perhaps been somewhat less conservative (See
the Species Group section).
All told, this study recognizes 39 species (in-
cluding 19 new species), establishes nine new
synonymies and transfers seven species to
Lordomyrma. It fails to resurrect the subgenus
Irogera. It retains three Polynesian /Melanesian
species (stigmatica, megastigmatica, exsulans) with
the Neotropical bulk of the genus.
As presently constituted, members of the ge-
nus Rogeria are distributed from Buenos Aires to
southern Texas and Arizona, and in the Pacific
between ION and 25°S from Tahiti to the western
end of the island of New Guinea. So far it is
unknown in Australia or southeastern United
States. Table 1 shows the distribution of species in
Volume 3, 1994
19
Table 1. Geographic distribution of Rogena species. The North American region is from southwestern United
States through Panama. The northern region of South America extends from the north and west coasts
through the Amazonian basin. The southern region of South America includes Paraguay, ad)acent Brazil, and
Argentina south to Buenos Aires. Trinidad is the only known Caribbean locality of three species (blandajirata,
scobinata). A question mark indicates that presumed queens, but no workers, were collected in that region. See
text for further discussion.
Speices
North Caribbean
America
South America
Pacific
Northern
Southern
alzatai
+
belti
+
besucheti
blanda
+
bruchi
brunnea
carinata
ciliosa
+
cornuta
creightoni
+
cuneola
+
cunnpubens
+
exsulans
foreli
+
germaini
gibba
uierrms
+
mnotabilis
+
lacertosa
leptonana
+
lirata
metastigmatica
merenbergiana
micromma
minima
neilyensis
+
nevadensis
pellecta
procera
prominula
scandens
+
scobinata
+
sicarta
stigtnatica
subarmata
terescandens
+
tonduzi
+
tnbrocca
unguispina
+?
+
+
+
20
Journal of Hymenoptera Research
more detail. In the North American region, most
species occur below the Isthmus of Tehuantepec,
but two species (creightoni, cuneola ) extend north-
ward through the eastern lowlands of Mexico.
Only foreli and creightoni have been collected in the
United States. The North American region con-
tains 8-10 endemic species (depending on the un-
certain distributions of innotabilis and leptonana);
the Caribbean, two endemic species; South America
(including Trinidad), 19 endemic species and the
Pacific, three endemic species. Only 5-7 species
(depending on innotabilis and leptonana) are found
in both Central America and northern South
America.
Little is known of the biology of these cryptic
ants. Collection records usually range from sea
level to 1000m, but five species extend higher and
two (ungiiispina and merenbergiana) can be found
at 2000m. Rogeria species are generally collected in
moist forests (primary or secondary forests, coffee
or cacao plantations), but at higher elevations can
be found in pastures {leptonana, merenbergiana).
Several species (creightoni, cuneola, foreli) have been
found in both moist and dry climates. Rogeria foreli
is the most unusual, with some members dwelling
at over 1800m in the temperate mountains of south-
ern Arizona.
Most Rogeria species have only been collected
as strays or by Berlese or Winkler sampling, usu-
ally in leaf litter and rotten wood, but occasionally
among epiphytes and moss (belti, creightoni,
exsulans). Nests of several species (belti, blanda,
merenbergiana) have been found under loose bark
of rotten logs. Nests of blanda and tondnzi have
been taken from the trunks of cacao trees. A nest of
leptonana was found at 1750m under a rock in a
pasture.
Because nests are so rarely found, males are
known for only four species (belti, blanda, leptonana
and stigmatica), and queens associated through
nest series for only nine species (See Synonymic
List of Species).
METHODS AND TERMINOLOGY
Dissections
Mouthparts, sting apparatus, pygidium,
hy popy gium, and male genitalia preparations were
obtained by rehydrating ants in 70% ethanol, dis-
secting these structures from the ants, clearing in
55-60°C lactophenol for 5 minutes (or longer if
necessary), rinsing twice in 70% ethanol, and rins-
ing twice in 95% ethanol. After clearing, sting
apparatuses and genitalia were usually cut in half,
except for the aedeagus and sting, which were
separated from the other sclerites. Stings, pygidia,
and hypopvgia were mounted in glycerin jelly for
ease of repositioning. Mouthparts, genitalia, and
other sting apparatus sclerites were soaked in
xylene, then mounted in Canada balsam. Occa-
sionally, whole ants were similarly cleared and
mounted in balsam. Preparations were examined
with a Zeiss KF-2 phase contrast microscope.
The "Materials Examined" sections of each
species description identifies which specimens
were dissected. Following the locality of the dis-
sected ant, brackets identify the structures slide-
mounted ("sting" means sting apparatus, py-
gidium and hypopygium). Dissected specimens
are workers unless otherwise identified.
Pinned vouchers are identified with the label
"Kugler 1991 Dissection Voucher." All are depos-
ited in the collection of the MCZ, except for vouch-
ers of lacertosa, pellecta, subarmata, the Agudos,
Brazil voucher of alzatei, and the Surinam vouch-
ers of curvipubens, which are all in the MZSP.
Illustrations
Drawings were made using a grid eyepieces
and grid-backed tracing paper. Drawings of sting
apparatus preparations were made at 400X with
estimated accuracy of <0. 001mm. Scanning elec-
tron micrographs were taken on an AMR-900 and
a JEOL JSM-35C.
Dorsal views of heads are in the same full-face
view used to measure HL. Lateral views of
mesosomas are in the same view used to measure
WL, SpL, and MHI.
Drawings attempt to show all hairs on the left
side of the body that project well above the body
outline. Care was taken to present typical pilosity,
rather than matted or damaged hairs. Hair is not
included in line drawings of: 1) dorsal views of
heads, 2) ventral portions of head profiles, 3) most
dorsal views of waists, and 3) queens and males.
All line drawings of sting apparatuses are
from workers.
Male genitalia drawings show the lateral view
of the aedeagus and medial view of the right
paramere and volsella (with associated part of the
Volume 3, 1994
21
phallobase).
Nontype specimens are identified in figure
legends by affixing their collection localities in
parentheses.
Measurements and Indices
Measurements and drawings of whole ants,
except for the very largest, were made using a
Zeiss SR stereomicroscope at 125X magnification
and a fiber optic ring lamp. Estimated accuracy: ±
0.01mm.
All specimens were measured for Weber's
Length (WL) then at least the largest and smallest
from each locality were measured completely. The
maximum and minimum of each measurement
and index were double-checked, as were all mea-
surements of holotypes. The number of specimens
(N) that follows the list of metric characters at the
beginning of a species description is the number of
ants that were measured completely. The numbers
at the ends of "Material Examined" sections are
total specimens studied.
In listing metric characters for new species,
the ranges found in all type material are followed
by the holotype measurements in parentheses.
If no queens or males are described for a
species, none were available for study.
CL Cephalic index: HW/HL
EL Eye length: Maximum diameter of compound eye
viewed straight on.
FLW Frontal lobe width: In full face dorsal view, maxi-
mum transverse distance across frontal lobes.
GL Gaster length: In lateral view, from anterior edge of
first tergum to posteriormost point usually T2 or
T3, but sometimes to end of pvgidium)
GW Gasterwidth: Maximum transversedistanceacross
gaster.
HL Head length: Maximum longitudinal distance from
the antenormost portion of the clypeus (usually
the clypeal apron, but sometimes the more promi-
nent body of the clypeus) to the midpoint of a line
across the back of the head.
HW Head width: Maximum width in same view as for
HL, excluding eves. For males, HW includes com-
pound eves.
ML Mandible length: In same view as for HL, from
antenormost portion of head to apex of closed
mandibles.
OI Ocular index: EL/HW.
PetL Petiole length: In lateral view and with petiole not
strongly flexed up or down, the axial distance from
the dorsal corner of the posterior peduncle to the
nearest edge of the metapleural lobe.
PetW Petiole width: Maximum transversedistanceacross
the node.
PpetL Postpetiole length: In lateral view, the axial dis-
tance from base of node in front to tip of posterior
peduncle.
PpetW Postpetiole width: Maximum transverse distance
across the postpetiole.
PSI Petiolar spine index: SpL/WL.
PW Pronotal width: Maximum width across pronotum
in dorsal view.
SI Scape index: SL/HW.
SL Scape length: Maximum length excluding basal
condyle and neck.
SpL Propodeal spine length (Fig. 1): In same view as for
WL, from tip of propodeal spine to nearest edge of
propodeal spiracle pentreme.
MH Mesosoma height (Fig 1): In perfect lateral view,
the vertical distance from a line tangent to the
ventralmost points of the of meso- and metasterna
to the highest point on the mesosoma profile (for a
broadlv arching pronotum, estimated as the mid-
point of the arc).
MHI Mesosoma height index (Fig. 1): Mesosoma height
(MH) divided by the horizontal distance from the
base of the MH line to the posteroventral corner of
the propodeum.
TL Total length: Sum of ML + HL + WL + PetL + PpetL
+ GL to the nearest 0.1mm.
WL Weber's length (Fig. 1): In perfect lateral view of
the mesosoma, diagonally from posteroventral
corner of mesosoma to farthest point on anterior
face of pronotum, excluding the neck.
Additional Terminology
Terms such as vertex, occiput, malar and genal
areas are of dubious accuracy when applied to
ants. Consequently, there is disagreement over
how to name the regions of the head (Trager 1989;
Snelling 1989; Wheeler 1989). Moreover, these
terms are confusing jargon that make the use of
keys and descriptions difficult for the uninitiated.
Instead, I prefer to rely, as much as possible, on the
generally understood directional terms, anterior,
lateral, etc., assuming that the mandibles are ante-
rior. Five regions of the head usually contain dis-
tinctive sculpture in Rogeria (Fig. 1). 1) The
middorsum of the head is the median portion of
the dorsal surface between the maximum retrac-
tion of the scapes and extending from the frontal
area to a change in sculpture near the rear of the
head. 2) The posterior (region of the) head in-
cludes the actual posterior surface of the head and
the posterodorsum, the dorsal surface between
the middorsum and the posterior outline of the
head. The "posterior head" seems equivalent to
22
Journal of Hymenoptera Research
the "vertex" of some authors who consider the
mandibles to be ventral and the back of the head to
be the top, rather than posterior. 3) The laterodorsa
are the portions of the head seen in dorsal view
that lie beneath the sweeps of the scapes. 4) The
venter is the ventral surface of the head not seen in
lateral view. 5) The sides of the head lie between
the venter and the laterodorsa (between the lateral
outline of the head in dorsal view and the ventral
outline of the head in lateral view).
In triangular mandibles the basal angle is
distinct and nearly a right angle (e.g. Figs. 43, 49,
63); in subtriangular mandibles the basal angle is
reduced and more obtuse (e.g. Figs. 14, 58, 61).
Sometimes tiny denticles are intercalated between
or replace teeth (e.g. Figs. 35, 55); these are not
included in tooth counts. The palpal formula is
the number of maxillary palpomeres followed by
number of labial palpomeres. Palpal formula is
usually determined by dissection, thus the sample
size is very small and may not show the variation
within the species. The body of the clypeus is the
medial portion of the clypeus anterior to the fron-
tal lobes and dorsal to the clypeal apron, which is
the anterior rim just above the mandibles (Fig. 1).
Clypeal apron shape is described as seen from
dorsal view. In describing eye shape, elliptical or
ellipsoid means similarly rounded at both ends,
whereas oval or ovoid means the anterior end is
more narrowly rounded. The shape of the poste-
rior outline of the head is as seen in the full dorsal
view used for measuring HL since shape some-
times varies with the pitch of the head. The nuchal
grooves are longitudinal impressions in the
posteroventral corners of the head, into which the
lower corners of the pronotum fit when the head is
retracted.
A rough index of the degree of inclination of
the propodeal spines is expressed by imagining a
bisecting line that divides the lateral aspect of the
spine in half and extends forward over the
mesosoma side, and noting where it intersects the
edge of the mesopleuron or pronotum (Fig. 1). It is
important that the mesosoma be in perfect lateral
view. The diameter of a propodeal spiracle in-
cludes the peritreme and is as seen in full lateral
view of the mesosoma. The spiracle's position is
measured from the outer edge of the peritreme to
the nearest edge of the propodeum, which is the
free edge of the infradental lamella.
The most common shapes of postpetioles as
seen from dorsal view are subrectangular, which
have evenly convex sides that are widest midlength
(e.g. Figs. 53, 58), and subtrapezoidal, which have
convex sides that diverge caudad (e.g. Figs. 49, 50,
51).
The terga and sterna of the gaster are referred
to by abbreviations Tl, SI, etc. Terminal seg-
ments of gaster rotated ventrad means that T2
and T3 are distinctly longer then their correspond-
ing sterna, causing the pygidium and hypopygium
(T4 and S4) to shift anteroventrad and making T2
or T3 the distalmost sclerite of the gaster. Figs. 28,
40, 43, and 63 show rotated terminal segments;
Figs. 2, 5, and 8 show unrotated segments.
Pygidial gland sculpture refers to a pair of
microareolate patches on the anterior edge of the
pygidium. The presence of this characteristic sculp-
ture strongly suggests the presence of pygidial
glands, but its absence does not necessarily mean
the glands are absent (Holldobler, et al. 1976;
Kugler 1978a; Holldobler and Engel 1978). In most
species I was able to see the gland reservoirs
attached to these patches, but failure to see the
reservoirs does not mean they are absent; they
could easily be lost during dissection.
Sting apparatus terms are identified in Fig-
ures 3 and 42.
Sculpture descriptions follow illustrations and
definitions of Harris (1979). It seems to me that the
term "areolate" in Harris' nomenclature best de-
scribes much of the sculpture in Rogeria, but I
suspect many would prefer the term "reticulate."
I have called the ridges on the head and mesosoma
of many species "rugae," because they are irregu-
lar, and wrinkled to varying degrees, but others
may prefer to call them "carinae." Also following
Harris, in compound descriptors, such as rugose-
areolate, the predominant type of sculpture is
first.
The following are used to describe approxi-
mate hair inclination as seen in side view (Fig. 1):
erect (> 70 ), suberect (70-50), subdecumbent
(50-25), decumbent (25-5), appressed (< 5 ) (in-
ference from Wilson 1955). To arrive at the num-
bers of erect hairs on the mesosoma dorsum, I
used both dark and light backgrounds and a vari-
ety of positions to count all erect hairs on one half
of the dorsum from the anterior edge of the pronotal
disc to the bases of the propodeal spines. Since the
hairs are distributed symmetrically, that count
gives the number of pairs of erect hairs. Frequently,
Volume 3. 1994
23
both sides were counted in order to check for the
presence of broken hairs. Erect hair on head dor-
sum means on frontal lobes, middorsum, and
posterior head, but not under the sweep of the
scapes. Erect hair on scapes means on the dorsal
or anterior surfaces. Usually no attempt is made to
describe hair on ventral and lateral aspects of
body, nor on other than extensor surfaces of femora
and tibiae.
DEPOSITORIES
ANIC
BMNH
CKC
CUIC
DMOC
FML
JTLC
LACM
MCSN
MCZ
MHN
MIZA
MZSP
NMB
PSWC
USNM
Australian National Insect Collection, CSIRO,
Canberra, Australia.
British Museum (Natural History), London, U.K.
Charles Kugler Collection, Radford University,
Radford, VA, U.S.A.
Cornell University Insect Collection, Ithaca, NY,
U.S.A.
David M. Olson Collection, University of Califor-
nia, Davis, CA, U.S.A.
Fundacion Miguel Lillo, Institute de Zoologia,
Miguel Lillo 251, Tucuman, Argentina
John T. Longino Collection
Los Angeles County Museum, Los Angeles, CA,
U.S.A.
Museo Civico di Storia Naturale "Giacomo Doha,"
Genoa, Italy.
Museum of Comparative Zoology, Cambridge, MA,
U.S.A.
Museum d'Histoire Naturelle, Geneva, Switzer-
land.
Museo Instituto de Zoologia Agricola, Universidad
Central de Venezuela, Maracay, Venezuela.
Museu de Zoologia da Universidade de Sao Paulo,
Sao Paulo, Brazil.
Naturhistorisches Museum, Basel, Switzerland.
Philip S. Ward Collection, University of California,
Davis, CA, U.S.A.
United States National Museum, Washington, DC,
U.S.A.
Genus ROGERIA Emery
Myrmica (in part); Fr. Smith 1858:131. Type: Formica rubra
Linneaus, 1758, designation of Wheeler, 1911
Tetramorium (in part); Mayr 1886:359. Type: Formica caespitum
Linneaus, 1758.
Rogeria Emery 1894:188. Type: Rogeria curvipubens Emery,
1894, by designation of Wheeler, 1911.
Rogeria subgenus Irogera Emery 1915:191. Type: Rogeria procera
Emery, by original designation.
Macromedia (in part); Mann 1922:30. Type: Macromischa
purpurata Roger 1863, by designation of Wheeler, 1911.
Irogera; Brown 1953:4.
Irogera; Kempf 1962a:435. [Redescnbed].
Rogeria; Kempf 1963:195. [Key to South American species],
Irogera; Kempf 1964:66. [Key to species]
Rogeria; Kempf 1965:185.
Rogeria; Kempf 1972:227. [Catalog of New World species].
Worker Diagnosis. — Monomorphic myrmi-
cines. Antenna 12-segmented; scape not reaching
posterior margin of head; distinct 3-segmented
club longer than rest of funiculus; apical
antennomere longer than combined lengths of
other two club segments. No antennal scrobes or
fossae. Clypeus projects narrowly between frontal
lobes at least to posterior edge of antennal inser-
tions. Body of clypeus with one or more pair of
longitudinal carinulae. Lateral clypeus not raised
into a ridge in front of antennal insertions. Nuchal
grooves present on posteroventral corners of head .
Anteroventral corners of pronotum angular to
dentate and fitting into nuchal grooves. Propodeal
spiracle 3 diameters or less from the edge of the
propodeum below the propodeal spines.
Metapleural lobes not sharply pointed.
Worker Description.— Mandibles usually trian-
gular. Except in some ciliosa and foreli, mandibles
with 5-7 teeth (3 apical teeth decreasing in size
basad, followed by 2-3 smaller basal teeth). Addi-
tional denticles may occur among basal teeth or
any basal tooth may be replaced by a pair of
denticles. Palpal formula usually 3,2 or 2,2, but 3,3
in some stigmatica-group, and 2,1 in the very tiny
minima. Scape neither elbowed nor ridged at the
base, nor with an apron around the peduncle.
Clypeus in profile usually with a very narrow
anterior apron. Body of clypeus rises near verti-
cally in most species, but occasionally projecting
beyond the clypeal apron. Frontal lobes narrow,
but covering antennal insertions; at most feebly
notched behind. Frontal triangle small, depressed
slightly. Eyes with 1-100 facets; located on sides, in
the anterior half of the head (excluding mandibles).
Sides of head widest just behind the eyes, forming
rounded corners with the posterior head, which is
weakly concave to weakly convex in full face
view.
Mesosoma generally compact, broad shoul-
dered. Anterior face of pronotum rises nearly ver-
tically from the neck and usually forms a distinct,
rounded angle with the dorsal surface. Mesosoma
24
Journal of Hymenoptera Research
dorsum without sutures; no mesonotal groove;
metanotal groove absent to distinct. Anterior edge
of propodeum often marked by a transverse ca-
rina. Propodeal spines absent to long. Metapleural
lobes low carinae to rounded triangular. Legs not
incrassate. No tibial spurs on middle or hind legs.
Tarsal claws simple.
Petiolar peduncle with or without a ventral
keel; inferior process dentate except in stigmatica
group. Node unarmed; poorly to well differenti-
ated from peduncle. Postpetiole with short pe-
duncles and a low node that is broader than long.
Terminal segments of gaster rotated ventrad in all
but the stigmatica group.
All pygidia dissected have a pair of small
pygidial gland reservoirs and/or paired
microareolate patches present on anterior edge,
except in the stigmatica group. Common features
of the sting apparatus are: 1) medial connection of
spiracular plate incompletely sclerotized, 2)
gonostylus single-segmented, 3) dorsoterminal
chaeta present, 4) at least one companion seta
(except gibba), 5) each lancet with a single moder-
ate to large valve, 6) sting bulb large, with arched
sting base.
Mandibles usually predominantly smooth,
with piligerous punctures and vestigial carinulae
at insertions, but carinulae stronger and more
extensive in some members of the stigmatica-group.
In all but ciliosa, the body of the clypeus is smooth
with a pair of carinulae extending from the frontal
lobes and stopping short of the clypeal apron;
these are sometimes accompanied laterally by 1-2
shorter, weaker carinulae. Lateral extremities of
clypeus and adjacent cheeks with longitudinal
carinulae. Frontal triangle smooth, except in ciliosa.
If macrosculpture present, frontal lobes and
middorsum with diverging longitudinal rugae;
rest of body areolate, rugose, or occasionally cari-
nate. Microsculpture when present usually
microareolate, appearing granular or punctured
at lower magnifications. Posterior face of
propodeum smooth, except in gibba. Legs smooth
and shiny. First segment of gaster smooth and
shiny; less so mprominula, and minima. Pygidium
and sometimes other terminal terga with micro-
scopic areolate sculpture on exposed posterior
surfaces; pygidium may also possess minute
piligerous tubercles.
Short appressed or decumbent pilosity com-
mon, most dependably on legs (except blanda and
procera ) and antennae. Terminal segments of gaster
with erect hair. No erect hair on laterodorsa of
head (under sweep of scapes). Body of clypeus just
above apron with pairs of erect hairs; members of
stigmatica group each with an additional median
hair.
Color from brownish yellow to black, with man-
dibles, antennae, and legs lighter. Most species
also with a lighter triangle on clypeus, cheeks, and
frontal area.
Malpighian tubules in the one species exam-
ined (belti) 5 in the worker (N=2); 5 to 6 in the male
(N=3) (Brown 1988).
Wings. — Notation (Fig. 30) follows Brown and
Nutting (1949) and Holldobler and Wilson (1990:9).
First and second radial crossveins of primitive
formicid wings are lost, creating a single large
submarginal cell. The distal portion of the radial
sector vein (Rsf4+5) vein is straight or curves
forward, but never reaches the edge of the wing.
The r-m crossvein present (Fig. 30) or absent (Fig.
37). Median and cubital veins of variable length,
but not reaching wing margins. Medio-cubital
crossvein always present in females, but may be
lost in males.
Larvae. — Similarities among the larvae of belti,
scandens, procera and blanda found by Wheeler and
Wheeler (1973, 1976, 1986, 1988) are summarized
here. Form pheidoloid, i.e. "abdomen short, stout
and straight; head ventral near anterior end,
mounted on short stout neck, which is the protho-
rax; ends rounded, one end more so than the
other" (Wheeler and Wheeler 1976:8). Thorax and
abdomen with at least bifid and anchor-tipped
hairs; bifid hair on head (sometimes others). Cra-
nium subtrapezoidal; clypeus bulging; antennae
with 3 sensilla; labrum bilobed and narrowed
dorsally. Mandibles ectatommoid (stout, gradu-
ally tapered and curved, with an apical tooth and
additional teeth and denticles in some species).
Maxilla shape paraboloid or lobose, with cylindri-
cal galea and palps of varying length. Labium
narrow; palps only slight elevations. Maxillae and
labial surfaces coarsely spinose.
COMPARISONS WITH SIMILAR GENERA
Some members of Leptotlwrax (placed with
Rogeria in the Leptothoracini) have antennae like
Rogeria and some have a narrow posterior lobe of
the clypeus, but these have rounded anteroventral
Volume 3, 1994
25
corners of the pronotum and no nuchal grooves.
Of 65 species of Leptothorax examined at the MCZ,
only one had an angular inferior corner of the
pronotum, but in that species the scapes extend
beyond the head, the posterior lobe of the clypeus
is wider, and nuchal grooves are absent.
Some species of Lordomymm (Myrmecinini), a
genus sometimes confused with Rogeria in pacific
islands, have similar antennal and clypeal fea-
tures, but have a rounded anteroventral corner of
the pronotum and lack nuchal grooves. The stings
of Rogeria species are dramatically different from
those of five Lordomyrma species I have examined
(caledonica Ern Andre, levifrons Mann, punctiventris
Wheeler, stnatella Mann, tortuosa Mann) in shape
of quadrate plate, oblong plate, gonostylus, trian-
gular plate and sting. Another species, L. epinotalis
is very different from the other Lordomyrma I ex-
amined and has a sting apparatus somewhat like
that of Rogeria besucheti or R. ciliosa, but lacks the
pronotal corner or nuchal grooves of Rogeria. Wings
of three Lordomyrma species I have been able to
examine (epinotalis, striatella, leae Wheeler) are dis-
tinct from the known Rogeria wings in having the
Rsf4+5 vein curve anteriorly to the wing margin.
In L. caledonica this vein is like Rogeria in failing to
reach the margin, but the m-cu vein is absent. The
larvae of an unidentified Lordomyrma species dif-
fers from the known larvae of Rogeria as follows:
form pogonomyrmecoid, cranium subelliptical,
hairs fewer, none anchor-tipped (Wheeler and
Wheeler 1977).
Stenamma (Pheidolini) workers are similar in
form of clypeus, including narrow posterior por-
tion between frontal lobes, and some have 3-seg-
mented antennal clubs, but in that case the apical
segment is shorter than the combined length of the
other two segments. Also, Stenamma has no nuchal
grooves, the anteroventral corner of the pronotum
is rounded, and the metanotal groove is generally
more distinct than in Rogeria species. Larvae of
Stenamma differ from those of Rogeria as follows:
form aphaenogastroid; cranium subhexagonal;
mandibles pogonomyrmecoid (similar to
ectatommoid); body hairs bifid or denticulate, not
anchor-tipped (Wheeler and Wheeler 1953, 1976).
Synonymic List of Species
Castes that have been collected are recorded
in brackets in the right column. W = worker, Q =
queen, M = male. Upper case Q and M are used if
at least some reproductives are associated through
a nest series. Lower case q and m are used if no
reproductives are linked to workers by a nest
series. An asterisk (*) is added if reproductives
were not even found in the same localities as
workers.
alzatei n. sp. [W, q]
belti Mann, 1922 [W, Q, Mj
besucheti n. sp. [W]
blanda Fr. Smith, 1858 [W, Q, M]
=foveata Kempf, 1964 (synonymy by Kempf 1965)
bruchi Santschi, 1922 [W]
brunnea Santschi, 1930 [W, q]
=caraiba Santschi, 1936 n. syn.
=cubensis Santschi, 1936 n. syn.
=habamca Santschi, 1936 n. syn.
=scabra Weber, 1934 n. syn.
cannata n. sp. [W]
cilwsa n. sp. [W, Q]
cornuta n. sp. [W]
creightoni Snelling, 1973 [W, q*[
cuneola n. sp. [W, q]
curvipubens Emery, 1894 [W, Q]
exsulans Wilson and Taylor, 1967 [W]
foreli Emery, 1894 [W, Q]
=gaigei Forel, 1914 n. syn.
=huachucana Snelling, 1973 n. syn.
germaini Emery, 1894 [W, q*]
=minensis Santschi, 1923 n. syn.
gibba n. sp. [W, q]
inermis Mann, 1922 (W, Q]
innotabilis n. sp. [W, q*]
lacertosa Kempf, 1963 [W|
leptonana n. sp. [W, q", M)
lirata n. sp. [W, q*|
megastigmatica n. sp. [W]
merenbergiaim n. sp. [W, Q]
micromma Kempf, 1961 [W]
minima Kusnezov, 1958 (Q]
neilyensis n. sp. \\\ |
nevadensis n. sp. [W]
pellecta Kempf, 1963 [wj
procera Emery, 1896 [W]
=brasiliensis Borgmeier, 1953 (Synonymy by Kempf
1962a)
pronnnula n. sp. [W]
scandens Mann, 1922 [VV, Q]
scobinata n. sp. [VV, q]
sicaria Kempf, 1962b [VV]
stigmatica Emery, 1897 [W, Q, M]
=sublevinodis Emerv, 1914 n. syn.
=manni Santschi, 1922 n. syn.
26
Journal of Hymenoptera Research
subarmata Kempf, 1962a
terescandens n. sp.
tonduzi Forel, 1899
tribrocca n. sp.
unguispma n. sp.
[W]
[W]
[W]
[W]
[W]
Names Transferred to Other Genera.
azumai Santschi 1941 (to Lordomyrma, Brown 1952)
epinotalis Mann 1919 to Lordomyrma, n. comb.
rugosa Mann 1921 to Lordomyrma, n. comb.
tortuosa Mann 1921 to Lordomyrma, n. comb.
tortuosa levifrons Mann 1921 to Lordomyrma, n. comb.
tortuosa pohta Mann 1921 to Lordomyrma, n. comb.
tortuosa stonen Mann 1925 to Lordomyrma, n. comb.
striatella Mann 1921 to Lordomyrma, n. comb.
SPECIES GROUPS
Some species of Rogeria can be assembled into
more or less distinct species groups. Others can
not be placed easily in any group, or seem to link
several groups. These incertae sedis species are
described with the group to which thev may be
most related.
The stigmatica-group may be distinct enough
to be treated as a separate genus, but for now
seems tied to the rest of Rogeria through blanda.
In the creightom species group, extensive varia-
tion and often limited and scattered collections
made it difficult to ascertain which specimens
were simple variants and which were distinct
species. At one time or another, I considered rec-
ognizing between four and 20 species in this group.
Ultimately, I tried to steer a middle course be-
tween these extremes; neither masking the varia-
tion by describing large conglomerate species, nor
encumbering the genus with a lot of dubious new
species names. As a result, I have been somewhat
more venturesome about naming new species in
this group, yet there remain a number of rather
heterogeneous species (alzatei, brunnea, creightom,
leptonana). I have described and illustrated some
of the variants in these heterogeneous species in
order to alert other taxonomists to remaining spe-
cies problems and the need for extensive collect-
ing, especially in the Caribbean Islands and
circumcaribbean countries.
In the diagnoses below, if a species is listed as
undissected, the palpal formula, pygidium, and
sting apparatus characters are unknown.
stigmatica-group: besucheti, ciliosa, gibba,
megastigmatica, prominula, stigmatica.
Diagnosis. — (prominula andmegastigmaticanot
dissected). WL 0.67-1. 30mm. Palpal formula 3,3 or
3,2. Eye with 4-41 facets. Metanotal groove distinct
and extends down sides of mesosoma. Petiolar
peduncle without a keel and inferior process a
small step or absent. Nodes and gaster wide. Gaster
with terminal segments not rotated ventrad, or
only slightly rotated (besucheti). No pygidial gland
sculpture or any evidence of reservoirs. Sting ap-
paratus: weakly sclerotized, spiracle occupying
more than 1/4 of spiracular plate; no anal plate;
valve chamber not arising abruptly from base of
sting shaft; sting shaft and lancets weak, sparulate
(except perhaps for prominula); sting shaft apex
without any flange; furcula a simple arch (no
dorsal arm). Predominantly dense areolate
macrosculpture on head (except middorsum),
mesosoma (except some stigmatica), and nodes.
Mandibles with piligerous punctures and stron-
ger, more extensive carinulate sculpture than in
other Rogeria species. No microsculpture on most
of body. Body of clypeus with a median erect hair
low near anterior margin.
Similar incertae sedis species: blanda. This species
has sculpture and pilosity like ciliosa and gibba, but
has a 2,2 palpal formula, weak metanotal groove,
dentate inferior petiolar process, and gaster, py-
gidium and sting apparatus typical of other groups.
scandens-group: scandens, subarmata, terescandens
Diagnosis. — (terescandens not dissected). WL
0.78-1. 17mm. Palpal formula 3,2 Eyes large (30-80
facets). Propodeal spiracle faces strongly caudad.
Metapleural lobes prominent, triangular. Petiolar
node low, anterior face indistinct from peduncle
(except some subarmata). Inferior petiolar process
dentate. Postpetiolar sternum large. Pygidial gland
sculpture and reservoirs present, at least in
subarmata. Sting apparatus like that of inermis (Fig.
42). Head, mesosoma, and nodes predominantly
rugose; sides of head below eye smooth. Dorsa of
head, mesosoma nodes, and gaster Tl with sparse,
stiff, untapered, erect hairs and sparse, incon-
spicuous appressed pilosity (occasional decum-
bent hairs in some subarmata). Scapes and legs
with appressed hair only.
Volume 3, 1 994
27
Similar incertae sedis species: procera, tonduzi.
These both lack the stiff hairs and inflated
postpetiole that make the group so distinctive, but
otherwise are so similar that perhaps they should
be included.
germaini-group: germaini, lirata, lacertosa.
Diagnosis. — WL 0.65-1. 05mm, usually >
0.80mm. Mandibles triangular. Palpal formula 3,2.
Body of clypeus not projecting over apron. Eye
small (6-20 facets; OI 0.09-0.14). Propodeal spines
rather long (PSI 0.18-0.23); directed caudad. Peti-
ole with prominent keel and dentate inferior pro-
cess. Petiolar node rises abruptly from anterior
peduncle and with more or less distinct anterior,
dorsal and posterior surfaces. Posterior two- thirds
of postpetiolar node with parallel or slightly con-
vergent sides. Postpetiolar sternum in side view
rather long, its anterior corner perpendicular. Py-
gidial gland sculpture and reservoirs present. Sting
apparatus similar to that of inermis, but sting shaft
and lancets weak; lancet lacks barbule. Laterodorsa,
sides, and posterior head rugose-areolate (some-
times effaced on sides). Mesosoma predominantly
rugose or vermiculate-rugose. Microsculpture
weak; intervals on head and mesosoma shiny and
nearly smooth. Mesosoma dorsum and gaster Tl
with abundant fine, tapered, erect-suberect hairs,
but very little, if any, decumbent or appressed
pilosity.
Similar incertae sedis species: pellecta, sicaria
(sicaria not dissected) These have different petiole
shapes and more decumbent hair on the mesosoma.
Rogeria sicaria also differs in clypeus, propodeal
spines, and erect pilosity.
creightoni-group:alzatei,brunnea,carinata,creightoni,
innotabilis, leptonana, merenbergiana, nevadensis,
scobinata, tribrocca, unguispina
Diagnosis. — (brunnea, nevadensis, and tribrocca
not dissected). WL 0.51-0.93mm. Mandibles trian-
gular, except as noted. Palpal formula 3,2 or 2,2.
Eyes small (6-19 facets). Petiole with a distinct
node and dentate inferior process. Pygidial gland
sculpture and reservoirs present. Sting apparatus
like that of inermis unless otherwise noted.
Mesosoma predominantly rugose or carinate.
Head dorsum, mesosoma, nodes, and gaster cov-
ered with two distinct kinds of hairs: 1) abundant
short, decumbent pilosity and 2) equally or less
abundant longer, erect-suberect hair (except for
lack of erect hair on head of some scobinata.).
Usually more than 10 pairs on mesosoma dorsum
(9 in some leptonana, 8 in carinata) and 2 or more
posterodorsally projecting hairs on each node. All
hairs tapered.
Similar incertae sedis species: inermis, belti,
cornuta, neilyensis, exsulans (cornnta and neilyensis
not dissected). These do not have two distinct
types of hair on the mesosoma dorsum.
curvipiibens-group: cuneola, curvipubens
Diagnosis. — WL 0.50-0.63mm. Mandibles tri-
angular. Palpal formula 2,2. Clypeal apron usu-
ally convex (medially flattened in Haitian
curvipubens). Eye small (4-11 facets). PSI 0.13-0.18.
Petiole with distinct node, weak keel, and dentate
inferior process. Postpetiole from above usually as
in Fig. 74. Pygidial gland sculpture and reservoirs
present. Sting apparatus as in inermis (Fig. 42),
except for sting shaft and lancets of curvipubens.
Median head and cheeks with weak longitudinal
rugose-areolate macrosculpture. Mesosoma with
predominantly rugose macrosculpture and ar-
eolate microsculpture, both often weak. Body cov-
ered with appressed-decumbent pilosity. Erect
hairs sparse: none on scapes or extensor surfaces
of legs, 0-16 on head (if present, short and limited
to posterior head), usually 2-7 pairs on mesosoma
dorsum (rarely 8), 0-1 pair posterodorsally pro-
jecting and 0-1 laterally projecting hairs on each
node. Anterior portions of gaster Tl often lack
erect hair. All hairs tapered.
Similar incertae sedis species: micromma, minima
(neither dissected). Rogeria micromma has a
subrectangular postpetiole in dorsal view and
abundant erect hair on head. Rogeria minima has
stiff, cuneate hairs and a 2,1 palpal formula.
/ore//-group: bruchi,foreli
Diagnosis. — WL 0.50-0. 80mm, usually <
0.75mm. Mandibles triangular. Palpal formula 2,2.
Eye 5-20 facets. Propodeal spines wide at base.
Petiole with distinct node and dentate inferior
process. Pygidial gland sculpture and reservoirs
present. Sting apparatus almost identical to that of
inermis (Fig. 42). Sculpture predominantly
microareolate on head, mesosoma, and waist;
28 Journal of Hymenoptera Research
macrosculpture feeble and limited to head. Most clypeus, ventral head, coxae, trochanters, sterna of
of body with short, appressed-decumbent pilosity gaster, and terminal terga of gaster.
only. Erect-suberect hairs present on mandibles,
KEY TO WORKERS OF ROGERIA
1 Pilosity appressed on mesosoma dorsum and gaster Tl. A very small number of short decumbent hairs
may also be present , 2
1 ' Erect, suberect, or subdecumbent hairs present on mesosoma or gaster Tl, often in addition to appressed
or decumbent pilosity 4
2(1) Body of clypeus strongly projecting over apron. Basal angle of mandibles very reduced, (amazonian
Brazil) (Fig. 14) prominula
2' Body of clypeus at most projecting only slightly over apron. Mandibles triangular 3
3(2') Clypeal apron emarginate. Mandible with 5 teeth. (Paraguay, Argentina) (Fig. 82) bntclii
3' Clypeal apron convex, often with faint median angle. Mandible with 6-7 teeth. (Caribbean, southwestern
United States through northern South America) (Figs. 79-81, 104-105) foreli
4(1') Very dense, flexuous, erect hairs on dorsa of head, mesosoma and gaster; no decumbent or appressed
pilosity. Promesonotum uniformly and densely areolate 5
4' Erect hair not so dense or flexuous. Decumbent hair present. Promesonotum may have some rugae or
carinae 7
5(4). Metanotum forms an abrupt declivity between promesonotum and propodeum. (lowland Colombia,
Ecuador) (Figs. 5-6) gibba
5' Shallow metanotal groove hardly interrupts mesosoma profile 6
6(5') Petiolar node relatively short and tall . Gaster larger (GW / WL 0.94-0.97). (lowland Ecuador, Venezuela)
(Figs. 2-4) ciliosa
6' Petiolar node relatively long and low. Gaster smaller (GW/ WL 0.63-0.70). (Central and South America)
(Figs. 83-84) blanda
7(4'). Ventral process of petiolar peduncle reduced to a small step in petiolar profile, or absent 8
Ventral process of petiolar peduncle dentate 10
8(7) Propodeal spines longer (PSI 0.17-0.20). Ventral process of petiolar peduncle present as a small step.
(Paraguay, Peru, Colombia) (Figs. 15-16) besucheti
8' Propodeal spines shorter (PSI 0.07-0.13). Petiolar peduncle lacking a ventral process 9
9(8') WL 0.72-0.92mm. (Pacific Islands) (Figs. 7-12) stigmatica
9' WL 1.18-1. 30mm. (Solomon Islands) (Fig. 13) megastigmatica
10(7') WL 1.28-1. 53mm. Eye large (80-100 facets). Postpetiole with an inconspicuous sternum. (Guyana,
amazonian Brazil) (Fig. 18) procera
10' WL and eyes usually much smaller. If WL and eyes nearly as large, then postpetiolar sternum
conspicuous 11
11(10) Sparse, stiff, erect-suberect hairs and short appressed hairs on dorsa of head, mesosoma, waist, and
gaster Tl. WL > 0.72mm. Eye with > 25 facets 12
11' Erect hairs more curved and tapered; shorter pilosity, if present, usually decumbent. If some erect hairs
are stiff, then WL and eye are smaller 14
12(11) Propodeal spines very short (PSI 0.09-0.12). Pygidium with a pair of median tubercles just above the
sting, (eastern Brazil) (Figs. 22-23) subarmata
12' Propodeal spines longer (PSI > 0.15). Pygidial tubercles lacking 13
13(12') Macrosculpture vestigial or absent on lateral pronotum and petiolar node. Waist slender (PpetW/PpetL
1.08-1.16). (Costa Rica) (Fig. 21) terescandens
13' Lateral pronotum with distinct rugae, petiolar node strongly rugose to areolate-rugose. Waist inflated
(PpetW/PpetL 1.38-1.61). (Central America) (Figs. 19-20) scandens
14(11) Petiole clavate. Eye with 39-48 facets. (Central America) (Figs. 85-86) tonduzi
14' Petiole with a distinct node and/or smaller eyes 15
15(14) Undamaged hair on mesosoma dorsum mostly erect to suberect. Decumbent hairs, if present, much less
abundant than longer, more erect pilosity 16
15' Mesosoma dorsum with abundant appressed or decumbent pilosity, usually in addition to longer erect
to suberect hair 21
Volume 3. 1994 29
16(15) EL/SpL > 1.00. Gaster Tl usually lacks decumbent and appressed pilosity (occasionally 1 or 2
decumbent hairs present). (Central America) (Figs. 40-42) inermis
16' EL/SpL usually < 0.90. If greater (some belti), then gaster Tl with abundant decumbent pilosity.... 17
17(16) Clypeal apron with median tooth 18
17' Clypeus emarginate to evenly convex 19
18(17) Promesonotum with thick rounded vermiculate rugae. Metapleural lobes well developed, triangular.
(Trinidad, Guyana, amazonian Peru and Colombia, Mato Grosso) (Figs. 28-30, 89-90) lirata
18' Promesonotum areolate-rugose to vermiculate rugose; rugae narrower, sharper. Metapleural lobes low,
broadly rounded. (Paraguay, Mato Grosso and Minas Gerais) (Figs. 24-27, 87-88) germaini
19(17') Eyes small and propodeal spines long and horizontal (EL/SpL < 0.46). (Belize, southern Mexico) (Fig.
45) cornuta
19' EL/SpL > 0.46. Propodeal spines more inclined 20
20(19') Eye smaller, elliptical (OI 0.13-0.14). Petiolar peduncle with lamellate keel. Sides of postpetiolar node
distinctly areolate. (southern Brazil) (Fig. 31) lacertosa
20' Eye larger, oval (OI 0.18-0.22). Petiolar keel not lamellate. Sides of postpetiolar node smooth, or nearly
so. (southern Mexico to Colombia) (Figs. 35-39, 91-92) belti
21(15') Scapes with longer erect to suberect hair in addition to shorter decumbent to appressed pilosity (in
exsidans and some brunnea, erect hairs can be sparse and little longer than decumbent hair) 33
21' No erect hair on scapes 22
22(21) WL 0.90-0.99mm. EL/SpL 0.48-0.56. (southern Brazil) (Figs. 32-33) pellecta
22' Smaller. If WL approaches 0.90mm, then EL/SpL either < 0.30 or > 0.70 23
23(22') Propodeal spines very long (PSI 0.29), strongly inclined. Eyesmall (EL/SpL0.13). (southern Brazil) (Fig.
34) sicaria
23' Propodeal spines usually much shorter. If similar in size, then not strongly inclined. EL/SpL > 0.20 ..
24
24(23') WL > 0.71mm. Pilosity on gaster not differentiated into two distinct kinds. (Costa Rica) (Fig. 43)
neilyensis
24' WL usually < 0.71mm. If similar in size, hair on gaster differentiated into short-decumbent and longer-
erect 25
25(24') Clypeal apron with median tooth. Gaster T3 with short median spine. (Colombia) (Fig. 63)
tribrocca
25' Clypeal apron without a median tooth. Gaster T3 unarmed 26
26(25) Erect-suberect hairs absent from head dorsum (or short and limited to posterior head) and sparse on
mesosoma. WL 0.51 -0.63mm 27
26' Erect hair present on middorsum and posterior head. If missing (scobinata), then > 10 pairs of erect hairs
on mesosoma dorsum 28
27(26) Postpetiolar sternum wedge-shaped in side view; node less vaulted. Sides of head and pronotum
strongly microareolate, making macrosculpture (if present) difficult to see. (central Mexico to Costa
Rica) (Figs. 77-78, 103) cuneola
27' Postpetiolar sternum not wedge-shaped; node more vaulted. Sides of head and pronotum with more or
less smooth, shiny intervals. (Caribbean Islands, northern South America; possibly Central America)
(Figs. 74-76, 101-102) curvipubens
28(26') Part of gaster Tl microareolate. Most erect hairs stiff, cuneate-fimbriate. (Argentina) (Figs. 72-73)
minima
28' Gaster Tl smooth. Stiff hairs absent (except on some micromma) 29
29(28') Rugae on pronotal dorsum and sides unbranching and nearly straight, with smooth and shiny intervals.
Sides of head below eye nearly smooth, strongly shining, (northeastern Colombia over 1000m) (Figs.
65, 94) nevadensis
29' Rugae on promesonotum and sides of pronotum undulating and with lateral spurs or branches,
sometimes forming areolate patches; intervals appear granular, not strongly shining. Sides of head
rugose-areolate 30
30(29') Eye with 2-5 facets. Mesosoma sides opaque with dense areolate microsculpture, but macrosculpture
absent. Mesosoma dorsum with 8-10 pairs of erect hairs. (Surinam, Para State of Brazil) (Fig. 71) ....
micromma
30' Eye with 4-21 facets (rarely < 7). Mesosoma sides with rugose macrosculpture in addition to weak or
distinct microsculpture. Mesosoma dorsum with > 12 pairs of erect hairs 31
30 Journal of Hymenoptera Research
31(30) Macrosculpture tuberculate on posterodorsum of head (Fig. 100). Erect hair usually absent from head
dorsum, but sometimes short, sparse and mostly limited to posterior rim. (Trinidad, South America
east of Andes to Paraguay) (Figs. 61-62, 100) scobinata
31' Posterodorsum of head rugose to areolate; no tubercles. Middorsum of head with long erect hair .32
32(31 ) Clypeal apron convex. Nuchal groove clearly visible in lateral view. Propodeal spines generally wider.
Petiolar keel distinct. (Mexico, Nicaragua; possibly into Colombia) (Figs. 55-57, 97-98) .. innotabilis
32'. Clypeal apron truncate in Central America. Nuchal groove not clearly visible in lateral view. Propodeal
spines usually slender. Petiolar peduncle with little or no keel. (Panama, South America, Dominican
Republic) (Figs. 58-60, 99) alzatei
33(21) WL 0.93-1. 02mm. Propodeal spines long (SpL > 0.20mm; PSI > 0.25), horizontal. Gaster with few or no
decumbent hairs. (Belize, southern Mexico) (Fig. 45) comuta
33' WL usually < 0.90mm. If larger (some belt i, brunnea, creightoni), then decumbent hair abundant on gaster
34
34(33) Gaster Tl with two distinct kinds of pilosity: shorter, decumbent hairs and longer, erect hairs 36
34' Hair on gaster Tl not clearly sorting into two distinct kinds 35
35(34) Pilosity on gaster Tl more dense and decumbent to subdecumbent. Petiolar peduncle with very large
keel. (Samoa) (Fig. 44) exsulans
35' Pilosity non gaster Tl more sparse and erect to subdecumbent. Petiolar peduncle with little or no keel.
(Central America) (Figs. 40-42) biennis
36(34) Eyes larger (> 19 facets), oval. Macrosculpture on pronotal disc usually predominantly areolate and
extending uninterrupted onto anterior face of pronotum (if predominantly rugose, then eye with > 25
facets), (southern Mexico to Colombia) (Figs. 35-39, 91-92) belti
36' Eyes smaller, often elliptical. Pronotal disc predominantly longitudinally rugose, vermiculate, or
carinate. Anterior edge of disc transversely rugose, rugose-areolate, or carinate 37
37(36') Promesonotum with nearly straight, longitud inal cannae; no lateral branches; intervals very smooth and
shiny. Eye small, elliptical (EL < 0.10. (Caribbean Islands) (Fig. 64) carinata
37' Promesonotum longitudinally rugose, vermiculate, or rugose-areolate. If intervals smooth and shiny,
then eyes oval and EL > 0.10 38
38(37) Clypeus evenly convex. Postpetiolar node from above subtrapezoidal; sternum long, flat, without
anterior lip. (Caribbean Islands) (Figs. 50, 93) brunnea
38' Clypeus usually emarginate or truncate. If convex, then postpetiolar node not subtrapezoidal; sternum
shorter, with distinct anterior lip 39
39(38') Eye elongate-oval. A strong transverse carina runs across pronotal shoulders. Pronotal disc and sides
with weakly undulating rugae and nearly smooth interrugal spaces. Propodeal spines often with
downcurved tips. (Venezuela) (Fig. 49) unguispina
39' Eyes not elongate. Anterior pronotum may have 1 or more transverse rugae, but not carinae. Pronotal
disc rugose or rugose-areolate. Propodeal spines straight 40
40(39') WL 0.54-0. 66mm. Mesosoma low, slender (PW < 0.37mm). Propodeal spines short, sometimes absent
(SpL < 0.11mm). Postpetiole generally widest in anterior half (Fig. 66) (southern Mexico, Central
America, Dominican Republic) (Figs. 66-70) leptonana
40' Generally larger, with MHI > 0.90, PW > 0.37mm, and SpL > 0.10mm. Postpetiole subtrapezoidal or
subrectangular (Figs. 51, 53) 41
41(40') Metanotum strongly interrupts contour of mesosoma profile. Sides of head rugose or effaced rugose-
areolate. EL/SpL > 0.65. (Colombian and Ecuadorian Andes above 1000m) (Figs. 46-48)
meretibergiana
41' Metanotum with little or no effect on overall shape of mesosoma profile. EL/SpL < 0.65. Sides of head
rugose-areolate; not effaced, (southern Texas to Panama) (Figs. 51-54, 95-96) creightoni
Volume 3, 1 994
31
SPECIES DESCRIPTIONS
Stigmatica-Group and Related Species
Rogeria ciliosa new species
Figs. 2-4
Additions to stigmatica-group diagnosis. WL
1 .06-1 .20mm. Mandibles triangular; coarsely punc-
tured, weakly carinulate. Anterior clypeus evenly
convex. Propodeal spines long (> 0.20mm),
strongly inclined and diverging. Metapleural lobes
well developed; angular. Inferior petiolar process
reduced to a small step. Dorsal head, mesosoma
and gaster densely covered with long flexuous
hairs; terminal segments of gaster with dense, stiff
erect hair.
Holotype and Paratype Workers. — TL 4. 1-4. 8 (4.1),
HL 0.90-1.02 (0.90), HW 0.78-0.89 (0.785), SL 0.69-
0.77 (0.69), EL 0.14-0.16 (0.15) (32-42 facets), PW
0.60-0.66 (0.60), WL 1.06-1.20 (1.06), SpL 0.24-0.28
(0.245), PetL 0.44-0.53 (0.44), PpetL 0.23-0.29
(0.235)mm, CI 0.84-0.87 (0.87), OI 0.17-0.19 (0.19),
SI 0.85-0.90 (0.88), PSI 0.21-0.23 (0.23). N=9
Mandibles with 7-9 teeth (3 apical teeth de-
creasing in size basad, followed by 4-6 small,
subequal teeth). Palpal formula 3,2. No clypeal
apron. Body of clypeus not projecting over ante-
rior edge. Nuchal grooves weak. Pronotal shoul-
ders rounded. Ventral petiole with a low median
carina rather than a distinct keel. Node distinct,
wider than long. Postpetiole dorsal view shape as
in Fig. 66. Postpetiolar sternum short, with a dis-
tinct anterior lip followed by a narrow sulcus.
Gaster large (GW/WL 0.94-0.97). Quadrate plate
of sting apparatus with somewhat reduced
apodeme that lacks lobes on anterodorsal corner;
oblong plate ventral arm very reduced (see also
Fig. 3 and stigmatica-group diagnosis).
Body of clypeus with fragmented longitudi-
nal rugulae surrounded by effaced areolate sculp-
ture. Longitudinal rugae on head dorsum mostly
confined to the frontal lobes, frontal area, and
midline. Frontal lobes rugose-areolate in some
paratypes. Rest of dorsum, cheeks, venter, and
posterior head densely areolate; intervals bearing
shallow piligerous punctures. Promesonorum with
the same areolate sculpture. Mesopleura,
metapleura, and sides of propodeum with more
irregular and confused areolate sculpture. Met-
anotal groove scrobiculate. Dorsal face of
propodeum transversely rugose with undulating,
smooth intervals or largely areolate with a few
carinulae between the spines. Most of petiole and
postpetiole strongly areolate as well. Sculpture on
anterior petiolar node effaced; dorsum of peduncle
smooth. First tergum and sternum of gaster rather
coarsely punctured in front and more finely punc-
tured caudad; smooth and shiny between punc-
tures.
Fine, long, flexuous, erect to suberect hair
covers middorsum of head, dorsum of mesosoma,
dorsum and sides of waist, and first segment of
gaster. On terminal segments of gaster these be-
come denser, stif fer and more erect to form brush-
like rings. Shorter, subdecumbent hairs occur on
lateral and ventral surfaces of head, dorsal sur-
faces of scapes, sides of mesosoma, and sometimes
dorsal face of propodeum. Very short, decumbent
to appressed pilosity on extensor surfaces of legs
and ventral petiole. Median hair on clypeus fine
and often obscured by surrounding paired hairs.
Body uniformly rusty-brown; appendages
lighter, more yellowish-brown.
Paratype Queen.— TL5.3,HL 1.04, HW 0.91, SL
0.80, EL0~27, PW 0.97, WL 1.50, SpL 0.30, PetL 0.58,
PpetL 0.30mm, CI 0.88, SI 0.88, PSI 0.20. N=l
Differing from paratypes only in the normal
queen characters (Fig. 4). Mandible8-toothed. Short
parapsidal furrows present on mesoscutum. Both
mesoscutum and mesoscutellum with same dense
areolate sculpture as in worker. Metanotum
vaguely microareolate. Dorsal face of propodeum
with transverse carinulae mesad; areolate laterad.
Discussion. — Rogeria gibba (Figs. 5-6) from
western Ecuador and Colombia resembles ciliosa
in size, sculpture and pilosity, but has different
mandibles, clypeal margin, propodeal spines, and
metapleural lobes. Rogeria stigmatica and
megastigmatica (Figs. 7-13) from the Pacific have
much shorter propodeal spines, less punctured
gaster, and different pilosity.
The name ciliosa refers to its dense covering of
flexuous hairs.
Distribution. — This species is known only from
lowland rain forests in the amazonian basin of
Ecuador and the Orinocan basin of Venezuela.
Material Examined. — Holotype locality. EC-
UADOR: Napo Province, Limoncocha, 250m, 18-
VI-1976, #B-348 (S. and J. Peck) [MCZ].
Paratype locality. 9 workers, 1 queen, VEN-
EZUELA: Bolivar State, Campamento Rio Grande,
8.07N61.42W, 250m, 14- VIII-1986, sifted leaf mold
32
Journal of Hymenoptera Research
and rotten wood, #8572-12 (P. S. Ward) [2 workers
dissected: mouthparts, 2 stings] [BMNH, LACM,
MCZ, MIZA, MZSP, USNM].
Rogeria gibba new species
Figs. 5-6
Additions to stigmatica-group diagnosis. WL
0.85-0.93mm. Mandibles subtriangular. Clypeal
apron slightly convex medially, with sharp cor-
ners on either side. Eyes with 16-20 facets.
Mesosoma profile humpbacked. Propodeal spines
short (< 0.15mm), not strongly inclined. Propodeal
spiracle rather large, within one diameter of pos-
terior edge of mesosoma. Metapleural lobes re-
duced to short carinae, sometimes nearly absent.
No inferior petiolar process. Dorsal head,
mesosoma and gaster densely covered with long
flexuous hairs.
Holotypeand Paratype Workers. — TL 3.5-3.8 (3.7),
HL 0.78-0.85 (0.83), HW 0.73-0.81 (0.78), SL 0.54-
0.60 (0.59), EL 0.09-0.10 (0.10) (15-22 facets), PW
0.51-0.57 (0.55), WL 0.85-0.96 (0.93), SpL 0.11-0.14
(0.11), PetL 0.40-0.49 (0.46), PpetL 0.21-0.23
(0.22)mm, CI 0.91-0.96 (0.94), OI 0.12-0.13 (0.13), SI
0.72-0.77 (0.76), PSI 0.12-0.15 (0.12). N=10
Mandibles with 5 teeth diminishing in size
basad. Palpal formula 3,3. Body of clypeus does
not project over apron. Head capsule about as
wide as long. Nuchal grooves visible from below
or behind. Promesonotal dorsum almost flat and
dropping abruptly to metanotum (Fig. 5). Petiolar
peduncle lacks inferior petiolar process. Petiolar
node distinct, bulbous, wider than long. Postpetiole
node subrectangular from above; sternum short.
Gaster large (GW/ WL 0.90-0.97). Sting apparatus
like that of ciliosa, except for: 1 ) a longer tulcral arm
on the oblong plate (like inermis Fig. 42), 2) no
companion seta on gonostylus, 3) more enlarged
lancet apex, and 4) no anterolateral processes on
sting base (Fig. 6).
Middorsum of head longitudinally rugose
becoming rugose-areolate behind level of eyes.
Laterodorsa, posterior, sides, and ventral surfaces
of head areolate with minutely granulate ridges;
intervals smooth except for some piligerous punc-
tures. Promesonotum with the same sculpture as
sides and back of head. Meso- and metapleura
with more confused areolate sculpture, but simi-
lar ridges and intervals. Metanotal groove
scrobiculate. Dorsal face of propodeum and part
of posterior face transversely rugulose. Rest of
posterior face shagreened. Petiolar peduncle
colliculate; anterior face of node smooth, except
for piligerous punctures. Rest of petiolar node and
all of postpetiolar node transversely areolate-rug-
ose, with granulate ridges and smooth to weakly
punctured intervals, as on head and prome-
sonotum. Gaster smooth and shiny, dotted by
numerous small, shallow piligerous punctures.
Dorsum of head, mesosoma, top and sides of
waist, and all sides of gaster densely covered with
long, flexuous, erect to suberect hairs and without
interspersed shorter pilosity.
Body rusty-brown, gaster slightly darker. An-
tennae, legs and mandibles yellowish-brown to
yellow.
Queens.— TL 4.2-4.3, HL 0.83-0.85, HW 0.80,
SL 0.58-0.59, EL 0.19-0.20, PW 0.72-0.73, WL 1.14-
1.15, SpL 0.15-0.17, PetL 0.49-0.53, PpetL 0.24-
0.25mm, CI 0.94-0.96, SI 0.73-0.74, PSI 0.13-0.15.
N=4
Queens differ from the workers in the usual
and the following ways. Short parapsidal furrows
present. Longitudinal areolate-rugose sculpture
on median head extends to posterior of head.
Median pronotum transversely rugose.
Mesoscutum with longitudinally rugose sculp-
ture that becomes more areolate on mesoscutellum.
Discussion. — See the ciliosa discussion for com-
parison. Like stigmatica also in many features, but
differs in pilosity and shape of promesonotum.
The name gibba is from Latin meaning humpback.
Distribution. — One worker is from the eastern
side of the central cordillera in northern Colombia,
the rest are from about 1000km away on the west-
ern slope of the Andes and the coastal range of
northern Ecuador. In all three areas they were
collected at 300-800m elevation in natural rain
forest, probably by Berlese sampling.
Material Examined. — Holotype locality. EC-
UADOR: Pichincha Province, 4 km E. Santo
Domingo de los Colorados, 22-VI-1975, #B-304 (S.
and J. Peck) [MCZ].
Paratype localities. COLOMBIA: 1 worker,
Antioquia Department, near El Bagre, Providencia,
Estacion Biologica, Zona Buenos Aires, 30-31-XII-
1977 (C. Kugler) [MCZ]; 1 worker, Choco Depart-
ment, Rio Napipi, 1968 (P. A. Silverstone) [LACM].
ECUADOR: 5 workers, holotype locality, 22- VI-
1975 and 8-VII-1976 (S. and J. Peck) [2 mouthparts,
stings] [BMNH, CKC, MCZ, MZSP]; 2 workers,
Volume 3, 1994
33
Pichincha Province, 47km S. Santo Domingo, Rio
Palenque Station, 23-V-1976 (S. and J. Peck [MCZ];
1 worker, Pichincha Province, Tinalandia, 16km
SE. Santo Domingo de los Colorados, 4-VI-1976 (S.
and J. Peck) [MCZ].
Nontype localities. ECUADOR: 3 queens,
Pichincha Province, 47km S Santo Domingo, Rio
Palenque Station, 1975 (S. and J. Peck); 1 queen,
Manabi Province, 73km NE Chone, 12-VI-1976 (S.
and J. Peck) [MCZ].
Rogeria stigmatica Emery
Figs. 7-12
Rogeria stigmatica Emery 1897:589. Syntype workers, NEW
GUINEA: Friednch-Wtlhelmshafen [=Madang] (Biro)
[MHN] (Both syntypes examined].
Rogeria stigmatica subsp. sublevinodis Emerv 1914:415. Svntvpe
workers, LOYALTY ISLANDS: Mare, Raoua [MHN]
[Both syntypes examined). N. syn.
Rogeria sublevinodis; Wilson and Taylor 1967:76, Fig. 61.
Rogeria m<mm'Santschi 1922:353. Syntype workers, SOLOMON
ISLANDS: Fulakora, Ysabel Is. (Mann) [MCZ, NMB] [1 2
syntypes examined]. N. syn.
Lordomyrma manm, Brown 1953:4.
Additions to stigmatica-group diagnosis. WL
0.72-0.92mm. Mandibles usually subtnangular.
EL usually > 0.10mm. Propodeal spiracle < half its
diameter from edge of infradental lamella.
Propodeal spines short (PSI 0.07-0.13), strongly
inclined dorsad. Metapleural lobes much reduced.
Inferior petiolar process absent. Abundant de-
cumbent hair on head, mesosoma, waist and gaster;
erect hairs rather sparse, not flexuous.
Workers.— TL 3.0-3.7, HL 0.66-0.83, HW 0.58-
0.71, SL 0.46-0.61, EL 0.05-0.15 (7-34 facets), PW
0.44-0.52, WL 0.72-0.92, SpL 0.05-0.10, PetL 0.32-
0.42, PpetL 0.19-0.23mm, CI 0.84-0.92, OI 0.08-
0.22, SI 0.79-0.87, PSI 0.07-0.13. N=28
Mandibles subtnangular (usually) to triangu-
lar; with 5 subequal teeth or 3 apical teeth followed
by 3-4 (rarely 5) smaller teeth or denticles. Palpal
formula 3,3. Little or no clypeal apron; median
clypeal margin truncate, weakly convex, or weakly
angular. Body of clypeus not projecting over clypeal
margin. Eyes oval, large (EL 0.10-0. 15mm and 17-
34 facets), except in some of the Papua New Guinea
workers (EL 0.05-0.07mm; 7-9 indistinct facets).
Nuchal groove distinct from behind, but not clear
in side view. Promesonotum with evenly convex
profile. Metanotal suture narrow, emphasized by
a sharp ridge at anterior edge of propodeum.
Node large, wider than long, more or less sym-
metrical in side view (Fig. 8). Postpetiolar node in
side view rounded front to back; usually widest in
anterior half, much as in Fig. 66, but sometimes
widest midlength. Postpetiolar sternum short.
Sting apparatus like that of ciliosa (Fig. 3), except
for: 1) spiniform medial and and lateral projec-
tions from anterodorsal corner of quadrate plates,
2) smaller valve chamber, and 3) lack of
anterolateral processes on sting base (Fig. 9).
Gonostylus sometimes with no clear sensillar gap;
sometimes lacking a companion seta. The "Rogeria
(stigmatica group) spp. 1 and 2" in Kugler (1978b)
are both stigmatica. The sting shown here is more
accurate than the previous one, which was not in
full lateral orientation when drawn.
Middorsum of head longitudinally rugose;
rest of head, including venter, coarsely areolate.
Dorsum of promesonotum coarsely areolate to
rugose (intermediate specimens predominantly
areolate, but with elongate cells or short rugae
medially; rarely, rugae also occur on shoulders).
Anterior and sides of mesosoma areolate, finely so
on neck and ventrad on meso- and metapleura.
Spaces in sculpture smooth except for piligerous
punctures. Dorsal face of propodeum areolate
along very anterior margin, followed by either
transversely rugulose or densely punctate sculp-
ture, or both in varying degrees of density and
definition. Petiolar peduncle finely colliculate or
smooth. Anterior and apex of node smooth or
areolate; posterior face and sides areolate, some-
times with a few rugae. Postpetiole transversely
rugose-areolate behind, becoming more effaced
anteriorly, often leaving anterior face smooth and
shining.
Decumbent to subdecumbent pilosity covers
most of body. Sparser erect to suberect hair also on
dorsa of scapes, head, mesosoma, nodes, and
gaster. Erect hair moderately abundant on gaster
Tl of most specimens (Fig. 8), but sparse on speci-
mens from Papua New Guinea. Body of clypeus
with strong median seta.
Color of mandibles, frontoclypeal region, an-
tennae, and legs light brownish-yellow to brown.
Rest of body light brown to blackish-brown.
Queens.— TL 3.6-4.5, HL 0.73-0.85, HW 0.66-
0.75, SL 0.53-0.62, EL 0.19-0.24, PW 0.60-0.71, WL
1.00-1.17, SpL 0.10, PetL 0.41-0.48, PpetL 0.21-
0.26mm, CI 0.87-0.90, SI 0.80-0.84, PSI 0.09-0.10.
N=6
34
Journal of Hymenoptera Research
As in worker except for the usual caste differ-
ences. Mesosoma habitus as in Fig. 10. Queen from
McAdam Park, Papua New Guinea with median
bulge on pronotum. Pronotum areolate on sides;
finer and transversely rugose-areolate mesad.
Mesoscutum longitudinally rugose; mesoscu-
tellum areolate-rugose. Metanotum smooth.
Mesosoma sides confused areolate, except for
smooth area on mesokatepisterna and costulate
metapleural gland bullae. Wing venation as in
Hrata (Fig. 30), except for Rs vein as in be/f; (Fig. 37).
Males.— TL 2.6-3.1, HL 0.45-0.54, HW 0.56-
0.66 SL 0.27-0.35, EL 0.20-0.26, PW 0.50-0.62, WL
0.84-1.04, PetL 0.22-0.30, PpetL 0.14-0. 19mm, CI
1.22-1.27, SI 0.48-0.53. N=3
Mandibles with a large apical tooth and 4
others decreasing in size basad. Posterior lobe of
clypeus projects more broadly between antennae
than in worker; anterior clypeal margin weakly
convex. Frontal lobes absent. No distinctly im-
pressed frontal area. Funicular segment 6 curved
and longer than 4 and 7; more extremely curved
and elongate on one side of the head than the
other. Posterior outline of head medially concave;
sharp crests run from ocelli to posteroventral cor-
ners of head, which project slightly and fit around
prothoracic sternum when head is retracted.
Mesosoma and waist as shown in Fig. 1 1 . Genitalia
as shown in Fig. 12. Head integument vaguely
roughened. Mesosoma and waist smooth, except
along furrows and on sides of propodeum,
metepimera, and petiolar peduncle. Gaster smooth
and shining. Pilosity all erect to suberect, except
around eyes. Propodeum nude. Color variation as
in worker.
Discussion. — According to Emery (1914) and
Wilson and Taylor (1967), sublevinodis differs from
stigmatica in having larger size, coarser sculpture
on head and mesosoma, and smooth nodes with
coarser punctures on other parts of the waist.
When two of Emery's stigmatica syntypes and two
of his sublevinodis syntypes in the MHN were
compared side by side, the TL and WL of the
stigmatica syntypes fell within the range of the
sublevinodis syntypes. The sculptural characteris-
tics were not distinct either, except on the dorsal
face of the propodeum, which is transversely ru-
gulose and very weakly punctate and shiny in
stigmatica syntypes, but densely punctate and lack-
ing rugulae in the sublevinodis syntypes. However,
intergrades with rugulae and various degrees of
punctation occur in the Solomon Islands, Irian
Jaya, and Pohnpei.
Santschi (1922) claimed that manni differed
from stigmatica in a variety of ways. After examin-
ing manni types, I could confirm only one clear
way they differ from the types of stigmatica and
sublevinodis: the presence of rugose sculpture in-
stead of areolate sculpture on the promesonotum.
However, in non-type material, I found all inter-
mediate states, sometimes within the same local-
ity. Other supposedly different characters also
intergrade or are due to the manni types being at
the small end of the size distribution.
I have too few collections from Papua New
Guinea to know if those specimens with unusually
small eyes and few erect hairs on gaster Tl might
be a distinct species.
See sister species megastigmatica description.
See also ciliosa and gibba for discussions of related
species in South America and exsulans for discus-
sion of a not so related species from the Pacific.
Distribution and Behavior. — Rogeria stigmatica
is known only from the Central and West Pacific,
from as far east as Tahiti to the western tip of the
Island of New Guinea and from about 22 S to 7°N.
Most collections come from berlesate or sift-
ing of leaf mould, rotten wood, soil, moss, or bases
of fern epiphytes in rain forest. The one nest series
with ecological data (Sorong, Irian Jaya) is from
rotten wood. Mann (1921:451) found colonies be-
neath stones and logs. Twelve specimens, were
collected on imported coconuts in Honolulu. If
nesting occurs in coconuts, colonization of
Polynesia and Melanesia from South America may
have occurred by rafting on the South Equatorial
Current.
Mann (1921:451) observed workers produc-
ing long, worm-like stands from the anal area
when the nest was disturbed.
Material Examined.— SOCIETY ISLANDS: Ta-
hiti, Punaauia District (J. Dixon). AMERICAN
SAMOA [=E. Samoa]: Tutuila [sting; whole speci-
men]; Tafuna; Alega (T. E. Woodward). WEST-
ERN SAMOA: Falepuna [sting]; Matautu; Vaipoto;
Poutasi (T. E. Woodward); Le Mafa; Gagaifomauga
(G Ettershank); Apia (H. Swale). WALLIS IS-
LANDS: Nukuione (G. Hunt) [mouthparts, sting,
whole specimen]. ILES DE HORNE [=Hoorn Is-
lands]: Futuna (G. Hunt). FIJI: Vanua Ava;
Waiyanitu; Ovalau; Lasema; Somosomo; Nagasau;
Saiaro; Munia; Nadarivatu; Labasa (W. M. Mann);
Volume 3, 1 994
35
Sigatoka (W. L. and D. E. Brown) [2 stings; male
genitalia]. VANUATU [=New Hebrides]: Espiritu
Santo (E. O. Wilson). LOYALTY ISLANDS: Mare.
SOLOMON ISLANDS: Ysabel, Fulakora [sting];
Ugi [=Uki] (W. M. Mann) [sting]. PAPUA NEW
GUINEA: New Guinea, Huon Peninsula (E. O.
Wilson); Wau, McAdam Park (S. Peck); Friedrich-
Wilhelmshafen [=Madang] (Biro). INDONESIA:
Irian Jaya, Sorong (W. L. Brown). CAROLINE
ISLANDS: Ponape [=Pohnpei] (Y. Kondo). 170
workers, 9 queens, 3 males [CUIC, MCZ, MHN,
USNM],
Rogeria megastigmatica new species
Fig. 13
Holotype and Paratype Workers. — TL 4.5-4.9
(4.5), HL ~1.01-1.09 (1.01), HW 0.90-1.00 (0.91), SL
0.77-0.83 (0.77), EL 0.16-0.19 (0.16) (41-52 facets),
PW 0.62-0.70 (0.62), WL 1.18-1.30 (1.18), SpL 0.10-
0.11 (0.11), PetL 0.51-0.52 (0.52), PpetL 0.28-0.31
(0.28)mm, CI 0.88-0.92 (0.90), OI 0.18-0.19 (0.18), SI
0.83-0.87 (0.85), PSI 0.08-0.09 (0.09). N=5
Like stigmatica in most respects, but markedly
larger (at least 35% larger than stigmatica on other
Solomon Islands). Metanotal groove not as nar-
row or sharply defined. Petiolar node strongly
asymmetrical in side view (Fig. 13).
Mandibles triangular, with at least 6 teeth,
some basal denticles may have been abraded.
Median clypeal apron convex. Mandibular
carinulae effaced. Pronotum areolate with a rug-
ose-areolate patch in center of disc. Dorsal face of
propodeum transversely rugulose. Petiolar pe-
duncle smooth; sides of node areolate; posterior
face transversely rugulose-areolate or areolate-
rugose.
Material Examined. — Holotype locality.
SOLOMON ISLANDS: Guadalcanal, Ilu Bush, 16-
III-1962, #1181 (P.J. M. Greenslade) (MCZ).
Paratype localities. SOLOMON ISLANDS: 1
worker, holotype locality [MCZ]; 3 workers, San
Cristobal, Kira Kira, 24-IV-1962, #1579 (P. J. M.
Greenslade) [MCZ].
Rogeria prominula new species
Fig. 14
Additions to stigmatica-group diagnosis. WL
0.78mm. Basal angle of mandibles greatly reduced.
Body of clypeus projects strongly over the trun-
cate clypeal apron. Frontal region elevated and
laterodorsa slightly concave. Eyes very small.
Metapleural lobes and metapleural gland bulla
reduced. Body almost devoid of erect hair.
Holotype Worker.— TL 2.88, HL 0.66, HW 0.56,
SL 0.51, EL 0.05 (4 facets), PW 0.435, WL 0.78, SpL
0.15, PetL 0.30, PpetL 0.19mm, CI 0.85, OI 0.09, SI
0.91, PSI 0.19.
Mandibles with 5 teeth, basal angle greatly
reduced (Fig. 14). Frontal lobes wide, closely ap-
proximated. Eyes small, round, sunken; facets
indistinct. Nuchal groove visible laterally as a
distinct notch. Pronotal shoulders well rounded.
Dorsal face of propodeum wide, weakly concave.
Propodeal spiracle about 2 diameters from edge of
infradental lamella. Metapleural lobes reduced to
low carinae. Metapleural gland bulla small. Peti-
olar node large, wider than long. Petiolar pe-
duncle short, with weak keel and small, dentate
inferior process. Posfpetiole subrectangular in
dorsal view. Postpetiolar sternum projects shelf-
like under articulation with petiole. GW/WL 0.84.
Terminal segments of gaster not rotated ventrad.
Shaft of undissected sting slender, with slightly
enlarged apex; lancets acute and appear strong.
Mandibles strongly carinulate for most of
length. Frontal area impressed, smooth. Median
head longitudinally rugose-areolate. Cheeks near
antennal insertions microareolate. Laterodorsa,
sides and posterior head densely macroareolate;
cells small, usually obscuring the effaced
microareola te background. Promesonotal dorsum
with same areolate sculpture as on back and sides
of head. Mesosoma sides strongly microareolate
and macroareolate; in different places one or the
other predominates. Scrobiculate sculpture in
mesosoma sutures. Dorsal face of propodeum dis-
tinctly microareolate, with branching rugulae
crossing between propodeal spines. Peduncle and
anterior face of petiole as well as sterna of petiole
and postpetiole microareolate; rest of petiole and
postpetiole macroareolate, somewhat effaced
mesad. Gaster Tl densely covered with piligerous
punctures; punctures weaker caudad. Other terga
smooth except for vaguely roughened posterior
margins.
Most of body covered with very fine, ap-
pressed pilosity. Sparse erect hairs on clypeus,
frontal lobes and nearby head middorsum. A few
short, decumbent hairs on mesosoma dorsum;
dense erect hairs on terminal segments of gaster.
36
Journal of Hymenoptera Research
No pilosity on ventral petiole.
Discussion. — This is perhaps the most aber-
rant Rogeria species, with its unusual head shape
and pilosity. But it has the characteristic Rogeria
antennae, nuchal grooves and square anteroventral
corners of the pronotum. It has affinities with
ciliosa, gibba, and stigmatica, as described in the
stigmatica-group discussion.
The name prominula, meaning little promi-
nence, describes the body of the clypeus.
Material Examined. — Holotype locality. BRA-
ZIL: Amazonas, Ig. Mananil, Rio Branco Road,
24km NE Manaus, 22-VIII-1962, #M-2 (W. L.
Brown) [MCZ].
Rogeria besucheti new species
Figs. 15-16
Additions to stigmatica-group diagnosis. WL
0.67-0.75mm. Eye small (8-10 facets). Propodeal
spiracles small, more than 3/4 diameter from
infradental lamella. PSI 0.17-0.20. Metapleural
lobes well developed. Inferior petiolar process a
small step. Head, mesosoma, nodes and gaster
with abundant decumbent pilosity and more sparse
erect hairs.
Holotyi>e and Paratype Workers. — TL 2.7-3.1
(2.85), HL 0.65-0.71 (0.66), HW 0.59-0.63 (0.60), SL
0.43-0.49 (0.44), EL 0.05-0.08 (0.06) (8-10 facets),
PW 0.42-0.46 (0.43), WL 0.67-0.75 (0.69), SpL 0.12-
0.15 (0.13), PetL 0.30-0.34 (0.32), PpetL 0.16-0.18
(0.18)mm, CI 0.87-0.90 (0.90), OI 0.08-0. 13 (0.10), SI
0.72-0.78 (0.72), PSI 0.17-0.20 (0.18). N=7
Holotype mandible with 5 visible teeth de-
creasing in size basad. In paratypes, mandibles
always with 3 apical teeth, but basal teeth may
have additional denticles or be replaced by pairs
of denticles. Palpal formula 3,2. Clypeal apron
truncate; body of clypeus projecting enough to
block view of apron in full dorsal view. Pronotal
shoulders rounded. Shallow metanotal groove
shallow present on dorsum and sides of mesosoma.
Propodeum lacking a distinct transverse carina at
anterior border. Peduncle of petiole with weak
ventral keel; inferior process reduced to a small
step. Petiolar node bulbous, wider than long.
Postpetiolar node widest in anterior half (as in Fig.
74). Postpetiolar sternum short, anterior lip not
greatly prominent. GW/WL 0.91-0.98. Terminal
segments of gaster slightly rotated ventrad, but
not enough to make T3 the distalmost point of the
gaster. Sting apparatus much like that of ciliosa
(Fig. 3), but: 1) anterior apodeme of spiracular
plate widest midlength, 2) anterodorsal corner of
quadrate plate longer, narrower, 3) anterior
apodeme of oblong plate longer, 4) gonostylus a
little longer, with two companion setae and less
distinct gap in setation and 5) sting base lower and
without anterolateral processes (Fig. 16).
Longitudinally rugose macrosculpture on
frontal lobes becomes rugose-areolate on
middorsum. Laterodorsa, sides, and posterior head
areolate with rather small areolae; intervals smooth
and shining, except for piligerous punctures.
Pronotal disc varies from all rugose-areolate to all
areolate. Rest of promesonotum slightly less
coarsely areolate. Intervals smooth, except for
piligerous punctures. Dorsal face of propodeum
densely microareolate, with or without overlying
transverse rugulae.
Dorsum and anterior face of petiolar node
smooth; rest of petiolar and postpetiolar nodes
effaced areolate. Gaster predominantly smooth
and shiny; Tl and SI with piligerous punctures
that in some specimens become weaker caudad.
Remaining terga and sterna very weakly rough-
ened and shiny.
Paraguayan specimens have suberect hair on
scapes; others do not. Head dorsum with suberect
hairs. Pilosity on mesosoma dorsum and nodes
ranges from short and decumbent to long and
erect; all curving quite strongly toward midline.
Gaster Tl similar, but with no erect hairs. Termi-
nal segments of gaster with rather dense erect hair
and decumbent pilosity. No hair on ventral peti-
ole.
Extremities and mandibles light brownish
yellow. Rest of body brown with more yellowish
than reddish accents; frontoclypeal area and ter-
minal segments of gaster lighter.
Discussion. — Rogeria besucheti differs from
ciliosa, gibba, prominula, and blanda in pilosity. It
differs from stigmatica and megastigmatica in
mesosoma shape, propodeal spine length, and
generally smaller eye size.
This species is named for Claude Besuchet,
who as director of the MHN in Geneva was most
helpful and patient in loaning material valuable
for this work.
Distribution. — Paraguayan specimens come
from gallery forest with some bamboo. Peruvian
specimens are from mixed broadleaf primary for-
Volume 3, 1 994
37
est on a steep hillside at 1000m. In both localities
collections resulted from Berlese and Winkler sam-
pling of leaf litter and rotten wood.
Material Examined. — Holotype locality. PARA-
GUAY: Alto Parana Province, Puerto Santa Teresa,
3-XI-1979 (F. Baud, et al.) [MHN].
Paratype localities. COLOMBIA: 3 workers,
Putumayo Department, Villa Garzon, 23-VII-1977
(D. Jackson) [BMNH], PERU: 2 workers, Pasco
Department, near Pozuzo, 1000m (C. Kugler and
R. R. Lambert) [mouthparts, sting] [MCZ, MZSP].
PARAGUAY: 1 worker, San Benito Province,
Itapua, 29-X-1982 (F. Baud, et al.) [MHN].
Rogeria blanda Fr. Smith
Figs. 17, 83-84
Myrmica blanda Fr. Smith 1858:131. Syntvpe workers, BRA-
ZIL: Amazonas, Ega [=Tefe] [BMNH] [4 syntypes exam-
ined].
Irogera foveaia Kempf 1964:64, Figs. 19-20. Holotype and
paratype workers, BRAZIL: Amazonas, Manaus (K.
Lenko) [MZSP] [Paratype examined]
Rogeria blanda; Kempf 1965:185.
Diagnosis. — WL0.87-l.15mm. Palpal formula
2,2. Eye rather large, oval. Propodeal spines long
(PSI 0.20-0.29). Petiolar node long and low. Gaster
with terminal segments rotated ventrad. Inferior
petiolar process dentate. Head, mesosoma, and
nodes densely areolate. Body rather densely cov-
ered with long, fine, soft, erect hair; no decumbent
hair.
Workers.— TL 3.0-4.0, HL 0.73-0.92, HW 0.65-
0.82, SL 0.49-0.62, EL 0.12-0.16 (27-59 facets), PW
0.48-0.63, WL 0.87-1.15, SpL 0.20-0.30, PetL 0.35-
0.49, PpetL 0.18-0.25mm, CI 0.85-0.91, OI 0.18-
0.21, SI 0.73-0.80, PSI 0.20-0.29. N=20
Additions to description and figures of Kempf
(1964). Mandibles triangular, with 6 large teeth
decreasing in size basad, then basal tooth larger
than neighbor. One or two denticles sometimes
added between basal teeth. Clypeal apron with a
shallow median notch. Nuchal grooves not visible
in lateral view. Metanotal groove weak (Fig. 83) to
absent. Propodeal spiracle small, strongly directed
caudad. Propodeal spines long, straight or weakly
upturned at apex; distinctly longer in the two
specimens from Ecuador (PSI 0.29) than in the
others (PSI 0.20-0.25). Metapleural lobes triangu-
lar; apex blunt to subacute. Petiolar peduncle
curved, with weak keel and dentate inferior peti-
olar process. Petiolar node widest in posterior
half. Postpetiolar node highest in posterior half;
shape from above as in Fig. 32 or Fig. 49. Gaster not
enlarged (G W/ / WL 0.63-0.70); terminal segments
rotated ventrad. Pygidial gland sculpture present.
Sting apparatus nearly identical to that of inermis
(Fig. 42).
Longitudinal rugae on frontal lobes rapidly
give way to areolate sculpture usually by mid-eye
level and continuing onto posterior head. Sides of
head strongly sculptured: rugose-areolate in front
of eye to areolate behind. Mesosoma and nodes
(Figs. 83-84) also densely areolate, except for trans-
verse carinulae between spines and smooth poste-
rior face of propodeum. Some elongate cells on
pronotal disc. Very apex of postpetiolar node some-
times smooth. No microsculpture.
Scapes with very long erect hairs and shorter
hairs ranging from suberect to decumbent. Erect
hair on terminal segments of gaster not brush-like.
Legs with little decumbent or appressed pilosity.
Head, mesosoma, and waist black to yellow-
ish-brown; appendages lighter. Gaster darker than
rest of body. Head sometimes with black patches
around and/or between eyes.
Queens.— TL 3.6-4.1, HL 0.79-0.87, HW 0.70-
0.78, SL 0.52-0.58, EL 0.19-0.20, PW 0.61-0.70, WL
1 .02-1 .17, SpL 0.24-0.30, PetL 0.40-0.52, PpetL 0.22-
0.25mm, CI 0.87-0.90, SI 0.73-0.81, PSI 0.22-0.26.
N=8
Median pronotum sometimes transversely
rugose-areolate; mesonotum longitudinally rug-
ose-areolate. Otherwise, differing from the worker
only in the normal queen characters. Wing vena-
tion like that of belti (Fig. 37).
Males.— TL 3.2-3.4, HL 0.60-0.62, HW 0.70-
0.75, SL 0.23-0.26, EL 0.32-0.35, PW 0.79-0.86, WL
1.10-1.24, PetL 0.34-0.35, PpetL 0.18-0.20mm, CI
1.16. SI 0.32-0.35. N=4
All males came from Rio Akaban, Venezuela.
Mandibles with 4 subequal teeth. Anterior edge of
clypeus with weak median notch. Frontal area a
distinctly impressed triangle. No frontal lobes.
Flagellomeres 2-11 straight, subequal in length
and width. Habitus much like male of belti (Fig.
38), but junction of dorsal and posterior faces of
propodeum has blunt lateral corners, and the
propodeal spiracle faces more caudad.
Back of head microareolate with piligerous
38
Journal of Hymenoptera Research
punctures in the pits; median vertex with addi-
tional longitudinal rugulae. Pronotum and
mesopleura largely smooth. Mesonotum densely
and finely rugulose with scattered punctures in
intervals. Posterior face of propodeum, nodes,
and gaster smooth. Genitalia as shown in Fig. 17.
Hairs shorter, less flexuous than in worker; erect
to suberect and moderately dense over much of
body. Color dark brown with lighter brown ap-
pendages.
Discussion. — Rogeria ciliosa and gibba from low-
land South America resemble blanda in having
long, soft, dense pilosity and areolate sculpture,
but see the stigmatica group diagnosis.
Distribution. — Rogeria blanda is found in south-
ern Central America and in South America east of
the Andes to southern Brazil. Elevations range
from 50m (Costa Rica) to 1000m (Venezuela). Nests
have been found in trunks of cacao trees in Costa
Rica and in a small rotten log suspended about
50cm above the ground in Peru.
Material Examined.— COSTA RICA: Heredia
Province, Puerto Viejo de Sarapiqui, La Selva Sta-
tion (L. Garling); Santa Clara Province, Hamburg
Farm (F. Nevermann); Puntarenas Province, Osa
Peninsula, Corcovado (J. Longino). PANAMA:
Barro Colorado Island (Brown and McCluskey; D.
E. Wheeler; J. Zetek). TRINIDAD: Basin Hill Re-
serve (N. A. Weber) [mouthparts, sting, whole
specimen]. VENEZUELA: Falcon State, near
Curimagua, Haitoncito; Monagas State, Caripe;
Bolivar State, Talud, south of Amarawai Tepuii
and Rio Akaban (J. Lattke) [2 male genitalia].
GUYANA [=British Guiana]: Oko River (N. A.
Weber). BRAZIL: Amazonas State, Tefe; Manaus
vicinity ( W. L. Brown; K. Lenko); Para State, Icoarci
(W. L. Brown), Belem (N. Rosa), Jacareacanga (M.
Alvarenga); Mato Grosso State, Municipal
Diamantino (W. L. Brown); Espiritu Santo State,
Linhares (M. Alvarenga). ECUADOR: Napo Prov-
ince, Limoncocha (R. Chadab). PERU: Madre de
Dios Department, Puerto Maldonado, Lake
Sandoval (C. Kugler) [mouthparts, sting]. 93 work-
ers, 8 queens, 4 males [BMNH, CKC, CUIC, MCZ,
MIZA, MZSP, USNM].
Scandens-Group and Related Species
Rogeria scandens Mann
Figs. 19-20
Macrormscha scandens Mann 1922:30, Fig. 14. Syntype work-
ers, HONDURAS: Lombardia (Mann) (USNM) [5 of b
syntypes examined].
Irogera scandens; Kempf 1962a:436, 438.
Rogeria scandens; Kempf 1965:185.
Additions to scandens-group diagnosis. WL
0.93-1. 17mm. Eye very large (about 60-80 facets).
Propodeal spines long (PSI 0.25-0.28). Petiole clav-
ate, with rather large node (PetW/PetL 0.49-0.58).
Postpetiole wide (PpetW/PpetL 1.38-1.61).
Mesosoma and petiolar node strongly macro-
sculptured. Erect hairs with dentate ends.
Workers.— TL 3.5-4.2, HL 0.83-1.02, HW 0.70-
0.85, SL 0.59-0.73, EL 0.17-0.20 (60-80 facets), PW
0.53-0.62, WL 0.93-1.17, SpL 0.23-0.30, PetL 0.42-
0.55, PpetL 0.23-0.27mm, CI 0.82-0.86, OI 0.21-
0.24, SI 0.84-0.86, PSI 0.25-0.28. N=6
Mandible in most specimens with 5 teeth de-
creasing in size until large basal tooth. A denticle
may appear between basal and penultimate tooth.
Nuchal grooves shallow, not visible in lateral view.
Metanotal groove in lateral view broad and shal-
low to absent; groove sometimes accentuated by a
low transverse ridge at anterior of propodeum.
Propodeal spines more strongly inclined in Hon-
duran (Fig. 19) than Panamanian specimens.
Metapleural lobes somewhat longer and narrower
in Honduran specimens. Petiolar node slightly
more distinct in Honduran specimens (Fig. 19).
Sting apparatus like that of biennis (Fig. 42), except
for wider anterior apodemes and a more rounded
posterodorsal corners on spiracular plates and
larger anterolateral processes on the sting base
(pygidium and anal plate lost in preparation).
Median clypeus with additional 1-2 pairs of
fairly distinct carinulae lateral to the usual pair for
the genus. Posterior head longitudinally rugose
(continuing from middorsum), transversely arch-
ing rugose, or transversely rugose-areolate.
Interrugal spaces on head distinctly to weakly
granular; microsculpture weaker and surface
shinier on posterior. Oval area on ventral half of
sides of head largely smooth and very shiny. An-
terior surface and neck of pronotum smooth;
rest of promesonotum with widely spaced longi-
tudinal rugae, which become vermiculate on
Volume 3. 1994
39
mesonotum. Rugae on Panamanian specimens
not so vermiculate on mesonotum, and with lat-
eral spurs. Sides of pronotum with parallel,
upcurved carinae, which are more numerous and
more distinct on Honduran specimens. Dorsal
face of propodeum confused areolate or areolate-
rugose. Intervals in mesosoma macrosculpture
smooth and shiny. Petiolar node heavily areolate-
rugose on sides and posterior face; sculpture
weaker dorsad; no clear microsculpture.
Postpetiolar node weakly rugose or rugose-ar-
eolate on sides and posterior surface, becoming
weaker, sometimes absent toward midline;
microsculpture vaguely microareolate to nearly
smooth.
Scapes and extensor surfaces of legs without
erect hair. Dorsa of head, mesosoma, nodes, and
gaster Tl sparsely covered with fine appressed
hair and longer, stiff, thick, erect-suberect hair
with toothed apex (Fig. 20). Mandibles, clypeus
and terminal segments of gaster with long erect
hair that is tapered and less stiff.
Color of Honduran specimens reddish-brown.
Panamanian specimens blackish-brown. Append-
ages often lighter.
Queen.— TL 4.2, HL 0.95, HW 0.82, SL 0.63, EL
0.22, PW 0.69, WL 1.20, SpL 0.28, PetL 0.50, PpetL
0.30mm, CI 0.86, SI 0.76, PSI 0.23. N=l
Differing from the Panamanian workers in the
usual queen characteristics. Pronotum transversely
rugose. Mesonotum longitudinally rugose with-
out cross-connections or vermiculate appearance.
Discussion. — The Honduran and Panamanian
specimens may be separate species. They differ
slightly in shape of mesosoma and petiole, sculp-
ture, and color, and the Honduran specimens are
a little larger (WL 1.13-1. 17mm) than the Panama-
nian specimens (WL 0.93-1. 08mm). I prefer to call
these geographic variants, however, until we have
more specimens from more localities.
See the terescandens description for compari-
sons with its sister species.
Ecology. — Some specimens from the Canal
Zone were collected from Heliconia.
Material Examined.— HONDURAS: Lombardia
(W. M. Mann). PANAMA: Canal Zone, Barro Colo-
rado Island (W. L. Brown and E. S. McCluskey; D.
Wheeler; J. Zetek) [2 workers: mouthparts, whole
specimen; sting]. 31 workers, 1 queen [CKC,
LACM, MCZ, USNM].
Rogeria terescandens new species
Fig. 21
Diagnosis. — Like that of scandens, except: 1)
Propodeal spines shorter, 2) petiolar node lower,
more slender (PetW/PetL0.40-0.41)/3)postpetiole
narrower (PpetW/PpetL 1.08-1.16), and 4)
macrosculpture very weak, especially on
mesosoma and waist, and 5) eye slightly smaller.
Holotype and Paratype Workers.— TL 3.6, HL
(0.88)-0.89, HW (0.68)-0.70, SL (0.70)-0.71, EL 0.16
(49-54 facets), PW (0.50)-0.52, WL 1.00, SpL (0.18)-
0.21, PetL (0.45)-0.47, PpetL 0.25mm, CI (0.77)-
0.79, OI 0.23-(0.24), SI 1.01-(1.03), PSI (0.18)-0.21.
N=2.
Also differing from scandens in the following
ways. Mandible with 6 teeth. Profile of mesosoma
dorsum almost evenly convex; no metanotal
groove or ridge at front of propodeum. Median
carinulae on clypeus weak; no lateral carinulae.
Head dorsum densely microareolate; overlain on
middorsum by wisps of of longitudinal rugulae,
on laterodorsa by faint reticulations, and on poste-
rior of head by fragmented, transversely arching
rugulae. Microsculpture on sides and posterior
head effaced. Mesosoma dorsum, including dor-
sal face of propodeum, densely microareolate,
with superimposed patches of fine rugulose-ar-
eolate macrosculpture. Pronotal sides shiny and
coriarious with some effaced longitudinal rugulae.
Mesopleura and metapleura also shiny near coxae,
but more opaque dorsad with microareolate and
confused rugulose sculpture. Sides of petiolar node
shiny, with effaced microareolate background and
vestigial longitudinal rugulae. Postpetiole smooth
and shiny.
Discussion. — The name of this species refers to
sculpture like that of R. scandens, but smoother, as
if rubbed (teres L., rubbed off).
Distribution. — Both specimens of terescandens
were taken from trees in lowland forest on the
Pacific side of Costa Rica. The holotype was col-
lected in a two week old treefall by general collect-
ing on trunks. The paratype was on or beneath a
thick epiphyte mat on the base of a fallen branch (J.
Longino unpublished field notes).
Material Examined. — Holotype locality.
COSTA RICA: Osa Peninsula, Sirena, 8.28N
83.35W, 50m, 31-111-1982, #0950 (J. Longino) [MCZ].
Paratype locality. 1 worker, holotype locality, 28-
V-1981, #1100 (J. Longino) [LACM].
40
Journal of Hymenoptera Research
Rogeria subarmata Kempf
Figs. 22-23
Irogera subarmata Kempf 1962a:438, Figs. 1-4. Holotype and
paratype workers, BRAZIL: Guanabara, Rio de Janeiro,
Deodoro (A. Ronna) [MZSP] [12 of 38 paratypes exam-
ined; holotype not examined].
Rogeria subarmata; Kempf 1965:185.
Rogeria subarmata; Kempf 1975:367 [new records].
Additions to scandens-group diagnosis. WL
0.78-1. 00mm. Eye with 30-53 facets. Propodeal
spines short (PSI 0.09-0.12). Pygidium with a pair
of median piligerous tubercles near caudal mar-
gin. Strong macrosculpture on mesosoma and
petiolar node. Erect hairs not as rigid as in scandens;
tips acute.
Workers.— TL 2.9-3.7, HL 0.69-0.87, HW 0.60-
0.75, SL 0.46-0.57, EL 0.12-0.16 (30-53 facets), PW
0.45-0.57, WL 0.78-1.00, SpL 0.08-0.10, PetL 0.37-
0.47, PpetL 0.20-0.26mm, CI 0.86-0.89, OI 0.20-
0.23, SI 0.73-0.77, PSI 0.09-0.12. N=6
Additions to Kempf's (1962a) description.
Mandibles usually with 5 teeth that decrease in
size basad. Sometimes basal tooth replaced by two
very small teeth, or 1-2 denticles are found be-
tween the basal and penultimate tooth. Clypeal
apron weakly notched medially to evenly convex.
Frontal lobes narrow as in scandens (Fig. 19). Nuchal
grooves shallow, forming only a weak notch in
lateral view. Figs. 22-23 show the range of
propodeal spine size and angle, but tips some-
times more rounded. Petiole clavate to rather dis-
tinctly set off from peduncles (Figs. 22-23).
Postpetiole from above much as in Fig. 21. Poste-
rior surface of pygidium with a caudal pair of
long, columnar, piligerous tubercles that are vis-
ible at 50X with a dissection microscope. Sting
apparatus nearly identical to that of inerniis (Fig.
42); sting as in pellecta (Fig. 33).
Median clypeus with 1-2 pair of fairly distinct
extra carinulae lateral to the usual pair. Posterior
head with transversely arching rugose-areolate
macrosculpture. Head covered with dense, indis-
tinctly microareolate roughening that appears
punctate or granular at lower magnifications.
Mesosoma dorsum longitudinally rugose; rugae
with numerous lateral spurs that occasionally con-
nect rugae on shoulders. Macrosculpture on sides
of mesosoma and dorsal face of propodeum con-
fusedly rugose to rugose-areolate. Mesosoma
microsculpture as on head. Petiolar node ver-
miculate-rugose to rugose-areolate. Postpetiolar
node similar, but rugae straighter, more effaced.
Microsculpture on nodes slightly weaker than on
head and mesosoma.
Scapes and extensor surfaces of legs lack erect
hair. Rest of body with both short, appressed-
decumbent and longer, erect-suberect hairs. Erect
hairs are nearly as stiff as those of scandens and
terescandens (Fig. 20), but seem to have acute tips.
Color dark brown to yellowish-brown with a
reddish tint on mesosoma, waist and middle of
gaster; appendages and ends of gaster lighter.
Distribution. — All available specimens are from
localities along the coast of Brazil. The type series
was collected from the stomach of an anteater
(Tamandua tetradactyla).
Material Examined. — BRAZIL: Para State,
Belem (N. Rosa) [mouthparts, sting]; Bahia State,
Itabuna (J. A. Winder); Espirito Santo State, Pedro
Canario near Conceicao da Barra (M. Alvarenga);
Guanabara State [=Rio de Janeiro State], Rio de
Janeiro, Deodoro (A. Ronna). 20 workers [MCZ,
MZSP].
Rogeria procera Emery
Fig. 18
Rogeria procera Emery 1896:92, Fig. 19. Holotype worker,
BRAZIL: Para, Belem [MCSN] [Holotype examined].
Rogeria {Irogera) procera; Emery 1915:191.
Macromischa brasiliensis Borgmeier 1953(1951 ):107, Figs. 1-4.
Holotype worker, BRAZIL: Para, RioCumina, Cachoeira
do Breu [MZSP] [Holotype examined). [Synonymy by
Kempf 1962a:437],
Irogera procera; Kempf 1962a:436 [partial description].
Irogera procera; Kempf 1964:66 [partial key].
Rogeria procera; Kempf 1965:185.
Diagnosis. — WL 1.28-1. 53mm. Eye very large.
Propodeal spines long, not inclined. Postpetiolar
node small, subconical; sternum inconspicuous
and without a differentiated peduncle. Middorsum
and posterior head with fine, nearly straight longi-
tudinal rugae. Mesosoma and petiolar node with
thick, vermiculate rugae. Erect hairs tapered; rarely
any appressed or decumbent hairs on mesosoma
dorsum, gaster, or legs. Scapes and extensor sur-
faces of legs with erect hairs and appressed to
decumbent pilosity. Palpal formula, propodeal
spiracle, metapleural lobes, petiole, pygidium, and
sting apparatus as in scandens-group diagnosis.
Workers.— TL 4.5-5.4, HL 1.07-1.19, HW 0.91-
1.08, SL 0.66-0.78, EL 0.19-0.23 (about 80-100 fac-
Volume 3, 1 994
41
ets), PW 0.67-0.80, WL 1.28-1.53, SpL 0.26-0.32,
PetL 0.60-0.73, PpetL 0.28-0.33mm, CI 0.85-0.91,
OI 0.20-0.22, SI 0.70-0.75, PSI 0.18-0.23. N=7
Mandibles triangular with 6 teeth; basal larger
than neighbor. Clypeal apron with shallow me-
dian notch. Nuchal grooves shallow, not visible in
lateral view. Sting apparatus like that of inermis
(Fig. 42), but spiracular plate with more rounded
anteroventral corner and gonostylus with sepa-
rate proximal and distal patches of sensilla.
Laterodorsa of head longitudinally rugose-
areolate; sides below eye smooth and shiny.
Microsculpture vaguely microareolate on anterior
laterodorsa; more effaced on rest of head, impart-
ing a vaguely granular, shiny appearance between
rugae. Promesonotum longitudinally vermiculate-
rugose. Meso- and metapleura rugose. Interrugal
spaces on mesosoma and petiole almost smooth.
Postpetiolar node shiny with vague rugae and
weak microsculpture.
Scapes and head dorsum with erect hair and
much shorter appressed to decumbent pilosity.
Mesosoma, legs, waist, and gaster with erect hairs,
but rarely any decumbent or appressed hairs.
Mesosoma and waist black to dark brown;
appendages and sometimes gaster lighter brown.
Material Examined.— GUYANA (British
Guiana): Oronoque River (N. A. Weber). BRAZIL:
Para State, Rio Cumina (A. Sampaio), Ourem (A.
Schulz); Amazonas State, Manaus (K. Lenko);
Manaus to Itacoatiara (W. L. Brown) [mouthparts,
sting]. 23 workers [CUIC, MCZ, MZSP].
Rogeria tonduzi Forel
Figs. 85-86
Rogerm tonduzi Forel 1899:53. Holotype worker, COSTA RICA
(Tonduz) (MHN) [Holotype examined).
Irogera tonduzi, Emery 1915:191.
Irogera tonduzi; Kempf 1962a:436.
Rogeria tonduzi; Kempf 1965:185.
Diagnosis.— WL 0.81-0.90mm. Eye large. Pal-
pal formula 2,2. Propodeal spiracle faces laterally.
Propodeal spines long. Postpetiolar sternum not
enlarged. Posterior head with transversely arch-
ing rugae. Rugae on mesosoma and petiolar node
thick and rounded. Decumbent hair abundant on
head dorsum and legs; little on gaster; no decum-
bent or appressed hair on mesosoma or nodes.
Scapes, head dorsum, mesosoma, nodes and gaster
with abundant flexible, tapered, erect hair; none
on extensor surfaces of legs. The following as in
scandens- group diagnosis: metapleural lobes, peti-
ole, pygidial gland sculpture, sting apparatus, and
sculpture.
Workers.— TL 3.0-3.2, HL 0.72-0.78, HW 0.61-
0.68, SL 0.48-0.53, EL 0.12-0.15 (39-48 facets), PW
0.46-0.50, WL 0.81-0.90, SpL 0.18-0.21, PetL 0.37-
0.40, PpetL 0.17-0.19mm, CI 0.85-0.88, OI 0.19-
0.24, SI 0.78-0.81, PSI 0.20-0.25. N=7
Mandibles triangular; most specimens with 6
teeth, the first 5 decreasing in size basad then a
large basal tooth. In others, the penultimate basal
is replaced by 2-3 denticles. Clypeal apron convex
with median angle or small tooth. Body of clypeus
rises perpendicularly. Eyes oval with narrow an-
terior point. Nuchal groove inconspicuous.
Pronotum in lateral view lacks a distinct angle
between anterior and dorsal faces. Metanotal
groove very weak to absent. Propodeal spines
long, weakly inclined. Postpetiole subtrapezoidal
from above; sternum flat, with a distinct peduncle.
Sting apparatus like that of inermis (Fig. 42) except
for more elongate anterolateral processes on sting
bulb.
Middorsum of head longitudinally rugose;
laterodorsa and dorsal part of sides rugose-ar-
eolate. Mesosoma (Figs. 85-86) with thicker rugae
and narrower interrugal spaces than on head.
Rugae transverse on anterior pronotum, trans-
verse to confused on metanotum and dorsal face
of propodeum, predominantly longitudinal on
sides and pronotal disc. Petiolar node longitudi-
nally rugose on sides; smooth along midline.
Postpetiole smooth. Microsculpture weak or ab-
sent throughout, integument very shiny.
Scapes and head with erect to suberect hair
along with the typical short decumbent pilosity.
Mesosoma dorsum and waist generally with erect
to suberect hair of a variety of lengths; no decum-
bent-appressed pilosity. Gaster with long, erect
hair and a few short, decumbent hairs.
Color shiny black with dark brown mandibles,
scapes and legs to reddish-brown with vellowish-
brown appendages.
Discussion. — Rogeria belti occurs in the same
localities and could be confused with tonduzi, but
belti has a distinct petiolar node and predomi-
nantly areolate pronotal sculpture. Rogeria lirata
(Figs. 28-30, 89-90) from northern South America
is similar to tonduzi in size, clypeus, and sculpture,
but lirata has smaller eyes (6-12 facets), a distinct
42
Journal of Hymenoptera Research
petiolar node, and scapes with suberect hair only.
Distribution. — With the exception of one speci-
men from Guatemala, all tonduzi specimens come
from Costa Rica at elevations of 0-100m on both
sides of the cordillera. Most specimens were col-
lected by Jack Longino as strays on ground and
vegetation. He found one worker in a Cyphomyrmex
nest and another "...on the base of a small tree,
amongst some Pheidole workers" (unpublished
field notes). Lyn Garling found a nest with a
"tubular entrance with white 'fuzz'" in a cacao
tree (field note on label).
Material examined.
GUATEMALA (no locality or collector).
COSTA RICA: Heredia Province, Puerto Viejo de
Sarapiqui, La Selva Station (L. Garling); Puntarenas
Province Carara Biological Reserve (P. S. Ward),
Osa Peninsula, Corcovado National Park (J.
Longino) [mouthparts, sting]. 24 workers [BMNH,
CKC, JTLC, LACM, MCZ, MZSP, USNM].
Gerwaim-Group and Related Species
Rogeria germaini Emery
Figs. 24-27, 87-88
Rogeria germaini Emery 1894:189. Syntype workers, BRAZIL:
MatoGrosso (Germain) [MCSN] [1 of 2 syntypes exam-
ined]
Rogeria germaini minensis Santschi 1923:1262. Lectotype and
paratype workers, BRAZIL: MinasGerais, PassaQuatro
(Reichensperger) [NMB] [Both lectotype and paratype
examined]. N. syn.
Rogeria germaini; Kempf 1962b:20, Figs. 18, 19 [Redescnbed]
Rogeria minensis; Kempf 1963:189, Figs. 1, 2 [Redescnbed,
raised to species].
Additions to gemiaini-group diagnosis. WL
0.74-0. 90mm. Clypeal apron with median tooth.
Metanotal groove very weak or absent. Metapleural
lobes reduced, not angular. Petiolar keel not lamel-
late. Sting shaft apex weak, lacks dorsal flange;
lancets spatulate. Macrosculpture effaced on side
of head below eye; sometimes nearly smooth.
Promesonotal rugae sharp and narrow like those
on head; rugae low on sides of pronotum do not
continue onto me tanotum. Macrosculpture on both
nodes and petiolar peduncle vestigial. Scapes with-
out erect hair. Mesosoma with erect-suberect only.
Head and gaster with both erect and decumbent
pilosity.
Workers.— TL 2.7-3.5, HL 0.66-0.85, HW 0.59-
0.74, SL 0.45-0.59, EL 0.06-0.10 (7-13 facets), PW
0.44-0.51, WL 0.74-0.90, SpL 0.15-0.21, PetL 0.30-
0.44, PpetL 0.19-0.23mm, CI 0.86-0.89, OI 0.09-
0.14, SI 0.75-0.80, PSI 0.20-0.23. N=9
Additions to Kempf (1962b). Dentition vari-
able; simplest pattern is 6 teeth of decreasing size,
however, it seems that any or all of the last 3 teeth
may be replaced by a pair of denticles. FLW/HW
0.32-0.36. Posterior outline of head concave to
convex. Anterior propodeum marked by a trans-
verse carina. Petiolar node distinct; Figs. 24 and 26
show extremes of shape. Postpetiolar node with a
distinct posterior face; shape from above as in
Figs. 24 or 32. Sting apparatus differing dramati-
cally from that of inermis (Fig. 42) in some features:
1) sting shaft and lancets very weak, 2) lancets
spatulate as in Fig. 29, 3) sting shaft apex with
eroded sides, and no dorsal flange (Fig. 27), and 4)
furcula with a shorter dorsal arm that broadly
merges with the lateral arms, thus appearing
broadly V-shaped in anterior view.
Posterior head transversely arching rugose to
rugose-areolate. Promesonotal dorsum varies from
predominantly areolate (with occasional elongate
cells) (Fig. 88) to predominantly vermiculate-rug-
ose. Pronotal sides rugose; rugae subparallel with
ventral edge of pronotum to diagonal.
Microsculpture vestigial, leaving irregular, but
shiny spaces in macrosculpture; sides of mesosoma
especially smooth.
Color brownish-yellow to brown with darker
gaster; appendages at times slightly lighter.
Queen.— TL 4.1, HL 0.90, HW 0.79, SL 0.73 EL
0.17, PW 0.67, WL 1.12, SpL 0.15, PetL 0.48, PpetL
0.28mm, CI 0.88, SI 0.92, PSI 0.13. N=l
Differing from the worker in the normal queen
characteristics and the following. Notch formed
by nuchal groove not so distinct in lateral view.
Anteroventral corner of pronotum not as clearly
dentate. Parapsidal furrows not distinguishable
from furrows in sculpture. Diverging rugae on
middorsum of head continue onto posterior head,
with few or no lateral spurs. Laterodorsa and sides
of head similarly rugose. Anterior face of prono-
tum transversely areolate, mesonotum with longi-
tudinal rugae diverging from an anterior point,
then parallel in posterior half; some branching,
but no cross-ridges; less vermiculate than in
worker.
Discussion. — Kempf (1963) examined the types
of germaini and minensis and noted that they were
very similar, but chose to retain both names on the
basis of a list of differences he saw in those speci-
Volume 3. 1994
43
mens. I have also examined the types as well as 1 1
specimens collected in Paraguay in 1979 and 1982.
These new collections bridge the gap between the
types oigermaini and minensis. They are all inter-
mediate in size between the smaller germaini type
and the larger minensis types, have convex poste-
rior heads like ihegermain i type, have promesonotal
sculpture varying from nearly as extensively rug-
ose as thege rmaini type, to areolate like the minensis
types; and some have petiolar nodes intermediate
between the more abruptly arising tninensis-hke
and the more obliquely arisingger??!fl;>n-like nodes.
Rogeria lirata (Figs. 28-30, 89-90) from more
northern parts of South America is germaini' s clos-
est relative (See lirata description for compari-
sons). Rogeria lacertosa (Fig. 31), also from southern
Brazil, differs in size, clypeal shape, sculpture on
side of head, and pilosity . Rogeria pellecta (Figs. 32-
33), collected further south in Brazil, differs in
clypeal shape, metapleural lobes, sting, and pilos-
ity.
Distribution. — So far germaini is known only
from southern Brazil and Paraguay. Most speci-
mens have come from Winkler apparatus collect-
ing by expeditions from the MHN in Geneva.
Specimens were extracted from rotting leaf litter
and wood in forests.
Material Examined. — BRAZIL: Minas Gerais
State, Passa Quatro (Reichensperger); Mato Grosso
State; Sao Paulo State, Anhembi [=Piramboia, 29km
E Botucatu] (Kempf et al.). PARAGUAY: Alto
Parana Province, Puerto Santa Teresa; Misiones
Province, Panchito Lopez; Itapua Province, San
Benito Island [2 workers: whole specimen; mouth-
parts, sting]; Itapua Province, Santa Maria; Cen-
tral Province, Asuncion (F. Baud et al.). 15 work-
ers, 1 queen [BMNH, CKC, MCSN, MCZ, MHN,
NMB],
Rogeria lirata new species
Figs. 28-30, 89-90
Diagnosis. — As in germainibut: 1) metapleural
lobes larger, triangular, and 2) rugae on
promesonorum more rounded and thicker than
on head; one ruga begins near anteroventral cor-
ner of pronotum and continues unbroken to the
metanotum.
Holotype and Paratype Workers. — TL 2.6-3.4
(3.3), HL 0.63-0.80 (0.78), HW 0.53-0.70 (0.67), SL
0.45-0.56 (0.55), EL 0.05-0.08 (0.08) (6-12 facets),
PW 0.39-0.51 (0.51), WL 0.65-0.87 (0.86), SpL 0.12-
0.20 (0.175), PetL 0.31-0.47 (0.44), PpetL 0.17-0.23
(0.21)mm, CI 0.83-0.88 (0.86), OI 0.09-0.12 (0.12), SI
0.79-0.85 (0.82), PSI 0.19-0.23 (0.20). N=10
Dentition as in germaini. Nuchal groove forms
a weak notch in lateral view. No clear angle be-
tween anterior and dorsal faces of pronotum. Spi-
racle faces slightly caudad, posterior edge within
one diameter of nearest edge of propodeum. Peti-
olar peduncle with a sharp, but not lamellate keel.
Petiolar node profile angular. Postpetiolar node in
dorsal view shaped as in Figs. 24 or 49; sternum
flat. Sting apparatus as described for germaini.
Divergent longitudinal rugae on frontal lobes
grade into areolate sculpture at level of eyes. Sculp-
ture of posterior head transversely arching rugose
to rugose-areolate. Laterodorsa and sides of head
longitudinally rugose-areolate; sides may be ef-
faced to nearly smooth. Head microsculpture ves-
tigial; intervals nearly smooth, quite shiny. Ante-
rior face of pronotum areolate, becoming ver-
miculate-rugose on disc (Fig. 90). Thick, rounded
rugae of promesonotum may merge, but are rarely
joined by cross-ridges. Rugae on meso- and
metapleura sharper and more separated. Anterior
and dorsal faces of petiolar node smooth. Sides
and posterior faces of node in holotype longitudi-
nally rugose-areolate, but weaker and more rug-
ose in Trinidad specimen and nearly smooth in
specimens from Peru. Microsculpture weak or
absent; integument shiny.
Scapes with decumbent to suberect hair (holo-
type) or with short, uniformly subdecumbent hair.
Head dorsum with subdecumbent to erect hair.
In most, mandibles, frontoclypeal region, an-
tennae and legs brownish-yellow; other parts rusty-
brown, becoming darker, almost black, on dorsa
of mesosoma and petiole. Trinidad and Guyana
specimens lighter on all parts.
Queens.— TL 3.6-3.7, HL 0.76-0.80, HW 0.69-
0.70, SL 0.52-0.57, EL 0.14-0.15, PW 0.58-0.60, WL
0.95-1 .00, SpL 0.18-0.20, PetL 0.46-0.52, PpetL 0.23-
0.24mm, CI 0.88-0.90, SI 0.76-0.81, PSI 0.18-0.21.
N=4.
Besides the usual features of an alate female,
differing from the worker in the following ways.
Mandible with 7 teeth or 3 teeth and 5 denticles.
Parapsidal furrows not distinct from furrows in
sculpture. Propodeal spines project caudad or
posteroventrad. Rugae on sides of pronotum al-
most vertical to shoulder, then bend across the
44
Journal of Hymenoptera Research
anterior face of the pronotum. Mesonotal rugae
longitudinal with occasional fusions and cross-
ridges. Wing venation (Fig. 30) most similar to that
of stigmatica. Radial sector and median veins long,
nearly reaching wing margin; r-m vein present.
Discussion. — See the tonduzi discussion for
comparisons with that species. The name lirata
refers to the characteristic rugae of the mesosoma,
which resemble the ridges thrown up by a plow.
Material Examined. — Holotype locality. CO-
LOMBIA: 7km N Leticia, forest litter, 10-25-11-
1972, #B-230 (S. and J. Peck) [MCZ].
Paratype localities. TRINIDAD: 3 workers, #191
(N. A. Weber) [MCZ, MZSP]; 1 worker, Mayaro,
Trinity Hills Reserve, 5-VIII-1976 (J. Noyes)
[BMHN]. GUYANA (British Guiana): 1 worker, R.
Mazaruni Forest Settlement, 20-VIII-1935, #304
(N. A. Weber) [MCZ]. COLOMBIA: 2 workers,
holotype locality [mouthparts, sting; one coated
for electron microscopy] [CKC, MCZ]. PERU: 2
workers, Loreto Department, 15km WSW
Yurimaguas, 5.59S 76.13W, 200m, 22-VII-1986,
#8701-24 and #8701-25 (P. S. Ward) [mouthparts,
sting] [LACM, MCZ].
Nontype localities. TRINIDAD: 1 queen,
Nariva Swamp, 23-IV-1935, #140 (N. A. Weber)
[MCZ]. BRAZIL: 3 queens, Mato Grosso State,
Sinop, 55.37W 12.31S, X-1974 #12314 (Alvarenga),
#12576 (Alvarenga and Roppa) [MZSP]; 2 queens,
Goias State, Jatai, XI-XII-1972, #8857, #8939 (F. M.
Oliveira) [MZSP].
Rogeria lacertosa Kempf
Fig. 31
Rogeria lacertosa Kempf 1963:194, Figs 5-6. Holotype and
paratype workers, BRAZIL: Rio Grande do Sul State,
Sinimbu (F. Plaumann) [MZSP] [All 4 paratypes exam-
ined; holotype not examined].
Additions to germaini-group diagnosis. WL
0.93-1. 05mm. Clypeal apron with median notch.
Metanotal groove weak to strong. Petiolar keel
with single lamellate carina. Sting shaft apex has
dorsal flange; lancets acute; both weak.
Macrosculprure not effaced on side of head below
eye. Promesonotal rugae sharp and narrow like
those on head. Sides of both nodes distinctly
macroareolate. Erect hairs on scapes. No decum-
bent hair on gaster; little if any on mesosoma
dorsum.
Workers.— TL 3.6-4.0, HL 0.83-0.91, HW 0.72-
0.81, SL 0.59-0.61, EL 0.10-0.11 (19-20 facets), PW
0.52-0.60, WL 0.93-1.05, SpL 0.17-0.20, PetL 0.43-
0.45, PpetL 0.23-0.26mm, CI 0.87-0.88, OI 0.13-
0.14, SI 0.75-0.79, PSI 0.18-0.20. N=4
The following supplements Kempf (1963). All
specimens at hand with 6 mandibular teeth de-
creasing in size basad, except for a large basal
tooth. Eyes elliptical. Nuchal groove forms a notch
in in lateral view of head. Anterior edge of
propodeum not marked by a transverse carina.
Metapleural lobes low and broadly rounded or
triangular with more narrowly rounded apex.
Postpetiolar node from above like that of pellecta
(Fig. 32). Sting apparatus as biennis (Fig. 42), but
sting shaft and lancets are less sclerotized (easily
twisted) and the lancets lack the barbule.
Laterodorsa of head predominantly rugose to
rugose-areolate. Back of head areolate in a trans-
versely arched pattern. Microsculpture vestigial;
intervals with a shiny, effaced granular appear-
ance. Anterior face of pronotum transversely ar-
eolate; disc with diverging, longitudinal, vermicu-
late rugae with variable number of cross-ridges
imparting a rugose-areolate appearance in places.
Sides of mesosoma predominantly longitudinally
rugose, but with occasional cross-ridges making
elongate cells. Intervals in mesosoma
macrosculprure shiny, nearly smooth, especially
on sides. Anterior and sometimes dorsal faces of
nodes weakly sculptured. Nodes slightly dulled
by vestigial microsculprure.
Head dorsum with erect-suberect hairs in
addition to the typical decumbent pilosity.
Color yellowish-brown; gaster slightly darker.
Legs and, sometimes, antennae lighter, more yel-
lowish. Mandibles often slightly darker than head
capsule.
Discussion. — Because minensis has been syn-
onymized with gertnaini, some of Kempf's (1963)
list of characters that distinguish lacertosa are no
longer valid, however, gertnaini and lacertosa are
still distinguishable on the basis of a number of
characters (see gertnaini discussion). Rogeria pellecta
(Figs. 32-33), also from southern Brazil, differs in
petiole keel, sting and lancets, promesonotal sculp-
ture, and pilosity.
Distribution. — Rogeria lacertosa is known only
from the type material collected from 100-200m
elevation in southern Brazil. No ecological data
are available.
Material Examined. — BRAZIL: Rio Grande do
Volume 3. 1 994
45
Sul State, Pardinho (F. Plaumann), Sinimbu (F.
Plaumann) [mouthparts, sting]. 4 workers [MZSP].
Rogeria pellecta Kempf
Figs. 32-33
Rogeria pellecta Kempf 1963:191, Figs. 3-4. Holotype worker,
BRAZIL: Santa Catanna State, Nova Teutonia (F.
Plaumann) [MCZ, MZSP] [6 of 28 paratype workers
examined, including 4 from holotype locality; holotype
not examined]
Diagnosis.— WL 0.90-0.99mm. Clypeus with
median notch. Metanotal groove distinct.
Metapleural lobes prominent, triangular. Petiolar
node arises gradually from peduncle. Sting and
lancets strong, acute; sting shaft with dorsal flange;
lancet with barbule. Laterodorsa of head longitu-
dinally rugose. Promesonotum coarsely areolate
to rugose-areolate. Back of petiolar node strongly
areolate; postpetiole vestigially areolate. Scapes
lack erect hair. Dorsa of head, mesosoma, nodes
and gaster with erect and decumbent pilosity.
Mandibles, palpal formula, eye, propodeal spines,
postpetiole, other aspects of petiole, sculpture,
and pilosity as in germaini-group diagnosis.
Workers.— TL 3.4-3.7, HL 0.81-0.89, HW 0.71-
0.78, SL 0.55-0.60, EL 0.09-0.11 (16-21 facets), PW
0.50-0.55, WL 0.90-0.99, SpL 0.17-0.21, PetL 0.39-
0.46, PpetL 0.20-0.22mm, CI 0.87-0.88, OI 0.12-
0.14, SI 0.77-0.79, PSI 0.19-0.22. N=6
The following supplements Kempf (1963).
Palpal formula 3,2. Anterior edge of clypeus weakly
emarginate. Eyes oval. Metanotal groove may be
bordered behind by a transverse costa. Posterior
face of petiolar node vertical or slightly concave in
lateral view. Postpetiole widest anteriorly, as in
Fig. 32, or evenly convex, as in Fig. 53. Sting
apparatus like that of inermis (Fig. 42), except for
the sting (Fig. 33), which has larger anterolateral
processes on sting base, a stronger, relatively
thicker sting shaft, and a lower dorsal flange.
Posterior head sculpture is rugose-areolate,
with rugae longitudinally diverging or transversely
arching across back of head. Rugae of laterodorsa
sometimes broken and with lateral spurs. Sides of
head areolate around eye, but effaced and nearly
smooth ventrad. Dorsal face of propodeum trans-
versely carinulate to coarsely areolate. Petiolar
node with areolate macrosculpture that becomes
weaker and less defined anteriorly. Ventral peti-
ole with a pair of longitudinal carinae arising from
an anterior keel. Head and mesosoma
microsculpture vestigial, leaving nearly smooth,
shiny intervals in macrosculpture. Sides of peti-
olar peduncle microareolate; nodes roughened by
obscure microsculpture and not as shiny as head
and mesosoma.
Color yellowish-brown; gaster slightly darker.
Legs, and sometimes antennae, lighter, more yel-
lowish. Mandibles often slightly darker than head
capsule.
Gynecoid Workers. — As described by Kempf
(1963).
Discussion. — In the southern Brazil /Paraguay
area occur related species germaini, lacertosa, and
sicaria. Comparisons with these species are found
in the "Species Groups" section and in the germaini
and lacertosa discussions.
Distribution. — Rogeria pellecta is known only
from the 33 type specimens taken in southern
Brazil from berlesate of leaf litter.
Material Examined.— BRAZIL: Santa Catarina
State, Nova Teutonia (F. Plaumann) [mouthparts,
sting]. 6 workers, 1 gynecoid [MCZ, MZSP].
Rogeria sicaria Kempf
Fig. 34
Rogeria sicaria Kempf 1962b:22, Figs. 20, 21 . Holotype worker,
BRAZIL: Sao Paulo, Agudos [19km SSE Bauru] (W W.
Kempf) [MZSP] [Holotype examined].
Diagnosis. — WL 0.75mm. Eye very small.
Clypeal apron convex. Metanotal groove strong.
Propodeal spines very long, strongly inclined dor-
sad and divergent. Petiole long (PetL/PW 0.95),
with prominent keel. Sting apex strong, with dor-
sal flange. Sides of head below eye, mesosoma,
and petiolar node areolate; coarsely so on
mesosoma. No erect hair on scapes. Head,
mesosoma dorsum, nodes, and gaster Tl with
stiff, erect-suberect hairs and shorter, more abun-
dant, appressed-decumbent hair.
Holotype Worker.— TL 2.8, HL 0.67, HW 0.56,
SL 0.48, EL 0.03 (8 facets), PW 0.37, WL 0.75, SpL
0.22, PetL 0.35, PpetL 0.17mm, CI 0.83, OI 0.05, SI
0.86, PSI 0.29.
The following supplements Kempf (1962b).
Mandible triangular, with 5 teeth decreasing in
size to base. Clypeal apron evenly convex; body of
clypeus projecting over mandibles. Frontal lobes
narrow (FLW/HW 0.29). Eyes small, circular,
somewhat sunken in head. Petiolar peduncle long
(PetL/PW 0.95), with strong keel and dentate
4h
Journal of Hymenoptera Research
inferior process. Postpetiole highest in caudal half;
dorsal view subtrapezoidal as in Fig. 49.
Posterior head areolate and more coarse than
elsewhere on head. Intervals shiny, but dulled
somewhat by vestigial microsculpture. Areolate
sculpture on petiolar node not as well defined as
on head and mesosoma and even more effaced on
postpetiolar node. Nodes roughened by vague
microsculpture.
Color uniformly golden-brown.
Discussion. — This species is still known only
from the holotype. It seems related to germaini,
pellecta, and lacertosa, also from southern Brazil.
See the "Species Groups" section for comparisons.
Creightoni-Group and Related Species
Rogeria merenbergiana new species
Figs. 46-48
Additions to creightoni-group diagnosis. WL
0.69-0. 83mm. Palpal formula 3,2. Clypeal apron
with median concavity. Eye small, elliptical; EL/
SpL 0.67-0.86. Anterior edge of pronotal disc with-
out a strong transverse carina. MHI 0.91-1.01.
Promesonotum and metanotum with distinct pro-
files— promesonotum convex, metanotum usu-
ally flat. Propodeal spines short, straight.
Postpetiole from above subtrapezoidal. Mesosoma
and sides of head rugose (sides of head sometimes
effaced rugose-areolate). Erect hair on scapes and
middle and hind tibiae.
Holotype and Paratype Workers. — TL 2.7-3.1
(3.0), HL 0.69-0.71 (0.71), HW 0.57-0.62 (0.61), SL
0.46-0.52 (0.50), EL 0.07-0.09 (0.08) (8-14 facets),
PW 0.42-0.47 (0.45), WL 0.74-0.83 (0.80), SpL 0.10-
0.14 (0.12), PetL 0.28-0.36 (0.335), PpetL 0.15-0.18
(0.16)mm, CI 0.83-0.88 (0.85), OI 0. 13-0. 15 (0.13), SI
0.80-0.84 (0.83), PSI 0.13-0.17 (0.15), MHI 0.91-1.00
(0.93). N=6
Mandibles with 6-7 teeth decreasing in size
basad and basal tooth as large or larger than
penultimate basal. If 6 teeth, may have 1-2 addi-
tional denticles among basal teeth. Body of clypeus
not strongly produced. Nuchal groove weak. Pos-
terior outline of head evenly convex. Propodeal
spiracle one diameter, or slightly less, from edge of
infradental lamella. Metapleural lobes low; edges
perpendicular. Petiolar node with a more or less
distinct angle between dorsal and posterior faces.
Petiolar peduncle with weak keel. Postpetiole with
longer anterior face and short posterior face meet-
ing at a narrow apex. Postpetiolar node from
above as in utiguispina (Fig. 49). Postpetiolar ster-
num short, weakly concave; anterior edge square.
Laterodorsa of head longitudinally rugose-
areolate. Posterior head with rugae diverging from
midline or transversely arching; few if any cross
ridges. Sides of head longitudinally rugose. Ante-
rior pronotal disc transversely rugose to rugose-
areolate; rest of promesonotal dorsum longitudi-
nally rugose. Dorsal face of propodeum areolate-
rugose. Lateral mesosoma rugose; less regularly
so on meso- and metapleura. Microsculpture on
head and mesosoma effaced; intervals quite smooth
and shiny, especially on sides and back of head.
Petiole distinctly microareolate; sides of node
vaguely rugose to vaguely areolate; posterior face
with strong ridges. Postpetiole effaced
microareolate and very sparsely and vaguely rug-
ose.
Color very dark reddish brown to yellowish
brown; appendages and often sides of mesosoma
and gaster lighter.
Nontype Workers.— TL 2.6-3.0, HL 0.63-0.71,
HW 0.53-0.60, SL 0.43-0.49, EL 0.08-0.09 (12-15
facets), PW 0.40-0.43, WL 0.69-0.77, SpL 0.10-0.13,
PetL 0.29-0.33, PpetL 0.16-0.18mm, CI 0.84-0.85,
OI 0.14-0.15, SI 0.79-0.82, PSI 0.14-0.17, MHI 0.95-
1.01. N=3
Little difference from types. Some slightly
smaller, but with slightly larger eye for size. Gen-
erally higher mesosoma height index. Posterior
head sometimes transversely arching rugose-ar-
eolate. Sides of head rugose-areolate, but effaced
posterior to eye. Promesonotal dorsum longitudi-
nally rugose to rugose-areolate. Lateral mesosoma
rugose-areolate.
Paratype And Nontype Queens. — TL 3.3-3.7, HL
0.70-0.75," HW 0.61-0.64, SL 0.48-0.52, EL 0.16-0.17,
PW 0.58-0.60, WL 0.98-1.15, SpL 0.15-0.17, PetL
0.34-0.39, PpetL 0. 19-O.20mm, CI 0.86-0.88, SI 0.78-
0.80, PSI 0.14-0.16, MHI 1.13-1.19. N=3
Mandible with 6-7 teeth; if 6, basal larger than
penultimate tooth. Clypeus evenly convex to
weakly emarginate. Posterior outline of head
strongly convex. Mesosoma and waist habitus
shown in Fig. 48. Sculpture mostly like worker.
Anterior pronotum areolate, becoming more rug-
ose on sides. Half to nearly all of ventral half of
mesopleura smooth; the rest of meso- and
metapleura and meso- and metanota longitudi-
Volume 3, 1994
47
nally rugose-carinate. Dorsal face of propodeum
transversely rugose. Intervals on mesosoma nearly
smooth; shiny. Petiole and postpetiole more
strongly microareolate and weakly areolate.
Discussion. — Rogeria unguispina (Fig. 49) from
the mountains of Venezuela (1100-2000m) is also
very similar in size, sculpture, pilosity and color to
merenbergiana, and the two species intergrade with
respect to mesosoma shape and petiole shape. It is
quite possible that these are geographic variants of
the same species, but all unguispina specimens
presently available have more elongate eyes and
have a strong carina running across the shoulders,
and specimens dissected differ in palpal formula.
Most unguispina specimens also have downcurved
propodeal spines, concave posterior face of peti-
olar node, and a broad, shallow metanotal groove.
See also the nevadensis discussion for comparisons
with that species from high elevations in Colom-
bia.
Rogeria alzatei occurs in Colombia, but so far
has not been found above 1000m. These ants are
usually smaller than merenbergiana (WL 0.51-
0.68mm), lack erect hair on the scapes, and lack a
distinct metanotal profile.
Rogeria creightoni, so far only known from
North and Central America, is related to
merenbergiana , but generally has: 1) longer, less
inclined propodeal spines, 2) less distinct metano-
tum, and 3) stronger, rugose-areolate sculpture on
sides of head (Figs. 51-54, 95-96).
Rogeria belti from Central America is the same
size and also generally has a more or less definite
step in mesosoma profile at metanotal groove, but
has: 1) larger eye, 2) longer propodeal spines, 3)
more rounded postpetiolar node, 4) more areolate
pronotum, and 5) mesosoma dorsum without two
distinct types of hairs.
This species is named in honor of Gunther
Buch and his family, who have heroically main-
tained part of their farm, Finca Merenberg, as a
nature preserve and permitted the collection of
these and other ants.
Distribution. — Rogeria merenbergiana is found
at high elevations (1300-2300m) in the Andes of
Ecuador and southern Colombia. It has been taken
in litter samples in moist broadleaf forest and
bamboo-moss forest. A nest was found in a rotten
log in a pasture.
Material Examined. — Holotype locality. CO-
LOMBIA: 14 workers, 1 queen, Huila, 12km W
Belen, Santa Leticia, Finca Merenberg, 2300m, 9-
13-1-1978, rotten jog in pasture (C. Kugler and J.
Hahn) [1 mouthparts; 4 stings] [MCZ].
Paratype localities. COLOMBIA: 13 workers, 1
queen, holotype locality [BMNH, LACM, MCZ,
MZSP, USNM]; 2 workers, Huila, Las Cuevas de
los Guacharos National Park, 1900-2300m, near
Palestina, about 20km S Pitalito, 15-17-1-1978 (C.
Kugler and J. Hahn) [whole specimen slide
mounted] [MCZ].
Nontype localities. ECUADOR: 2 workers,
Pichincha Province, 16km E Tandapi, 2000m, in
litter, 20-VI-1975, B-302 (S. and J. Peck); 3 workers,
2 queens, Pichincha Province, 3km E Tandapi,
1 300m, litter in wet ravine, VI-1975 (S. and J. Peck);
4 workers, Pichincha Province, 20-30km ENE
Alluriquin on Chiriboga Road, 1400-1800m, 1975,
B-301 (S. and J. Peck) [CKC, MCZ].
Rogeria unguispina new species
Fig. 49
Additions to creightoni-group diagnosis. WL
0.64-0.83mm. Palpal formula 2,2. Clypeal apron
with median concavity. Eye moderately large,
elongate-oval. Strong carina across shoulders.
Metanotal groove broad and shallow. Propodeal
spines usually with downcurved tips. Postpetiole
from above subtrapezoidal. Sculpture on side of
head vestigial, leaving a small or extensive smooth
area. Pronotal disc and sides with weakly undu-
lating rugae and nearly smooth, shiny interrugal
spaces. Erect-suberect hair on scapes, and middle
and hind tibiae.
Holotype and Paratope Workers. — TL 2.4-3.1
(3.0), HL 0.60-0.72 (0.71), HW 0.50-0.61 (0.60), SL
0.40-0.49 (0.47), EL 0.10-0.11 (0.11) (15-19 facets),
PW 0.36-0.46 (0.46), WL 0.64-0.83 (0.81), SpL 0.09-
0.13 (0.13), PetL 0.26-0.34 (0.34), PpetL 0.14-0.19
(0.19)mm, CI 0.82-0.86 (0.84), OI 0.12-0.16 (0.13), SI
0.79-0.82 (0.78), PSI 0.14-0.16 (0.15). N=7
Mandibles with 5-6 teeth, basal larger than
penultimate tooth; some with a denticle between
basal and penultimate teeth. Body of clypeus not
projecting over apron. Nuchal groove makes a
notch in side of head. Pronotum rounded on front
and sides. Propodeal spiracle large (Fig. 49) to
moderate; less than one-half diameter from near-
est edge of propodeum. Free edges of metapleural
lobes form an oblique angle. Posterior face of
petiole concave in holotype and some paratypes.
48
Journal of Hymenoptera Research
Petiolar peduncle lacks a distinct keel. Postpetiolar
node highest in posterior half. Postpetiolar ster-
num with a short, flat profile and square anterior
corner.
Laterodorsa of head rugose, sometimes ar-
eolate caudad (holotype). Posterior head rugose-
areolate in a more or less transversely arching
pattern. Interstices on head vaguely roughened by
effaced microsculpture; especially shiny on sides
and back of head. Promesonotal dorsum longitu-
dinally rugose in front, becoming more vermicu-
late with partial or complete cross-ridges behind.
Pronotal sides longitudinally rugose with few
cross-ridges. Meso- and metapleura with broken,
irregular macrosculpture. Dorsal face of
propodeum transversely areolate. Interstices on
disc and sides of pronotum shiny, nearly smooth;
rest of mesosoma vaguely microareolate and quite
shiny. Petiolar node weakly to vestigially
microareolate; posterior face of node with longitu-
dinal carinulae. Postpetiolar node nearly covered
with vague microsculpture to nearly smooth; al-
ways weaker than on petiole.
Color blackish-brown with yellowish-brown man-
dibles, flagellum of antennae, legs, and terminal
segments of gaster.
Discussion. — Rogena nevadensis is a similar ant
from the mountains of northeastern Colombia,
but nevadensis: 1) is usually smaller, 2) has smaller
eyes, 3) has straight propodeal spines, 4) has more
effaced sculpture, and 5) lacks erect hair on the
scapes and tibiae. See also the merenbergiana dis-
cussion.
The name unguispina refers to the claw-like
shape of the propodeal spines.
Distribution. — So far, unguispina is known only
from Venezuelan rain forest between 200m and
2000m elevation.
Material Examined. — Holotype locality. VEN-
EZUELA: Miranda state, Rio Capaya at Quebrada
El Bagre, near Salmeron, E of Caracas, 200m, nest
beneath log on forest floor in limestone gorge, 8-II-
1982, #238 (J- Lattke) [MCZ].
Paratype localities. VENEZUELA: 9 workers,
holotype locality [mouthparts, sting] [BMNH,
CKC, LACM, MCZ, MZSP, USNM]; 1 worker,
Falcon State, Sierra de San Luis [mountain range
just south of Paraguana Peninsula], Haitoncitos,
near Curimagua, 1180m, primary forest, 3- VIII-
1982, #271 (J- Lattke) [MCZ]; 1 worker, Aragua
State, Rancho Grande, 15-VI-1960, 4563a (J. qasti)
[MCZ]; 1 worker, Aragua State, Rancho Grande,
14-VIII-1967, 1100m (R. W. Poole) [MCZ]; 2 work-
ers, Lara State, vicinity of Caspo, 18km S Sanare,
1620m, forest rotten wood, 6-XII-1985 (J. Lattke
and W. L. Brown) [MIZA].
Rogeria brunnea Santschi
Figs. 50, 93
Rogena curvipubens brunnea Santschi 1930:79. Syntype work-
ers, CUBA: Pinar del Rio, Sierra de los Organos, Rangel
(A. Bierig) [NMB] [Both syntypes examined]. N. comb.
Rogena scabra Weber 1934:27, Fig. 2. Syntype workers, queen,
CUBA: Cienfuegos, Soledad, Limones Seboruco
(Darlington and Weber) (MCZ) [Queen and 1 of 2 worker
syntypes examined], N. syn.
Rogeria caraiba Santschi 1936:198, Fig. 5. Syntype workers,
CUBA: Habana, Playa de Mananao [NMB] [All 8
syntypes examined]. N. syn.
Rogena cubensis Santschi 1936: 199, Figs. 3-4. Holotype worker,
CUBA: Pinar del Rio, Sierra del Rosano (A. Bierig)
[NMB] [Holotype examined]. N. syn.
Rogena cubensis liabanica Santschi 1936:200. Holotype worker,
CUBA: Playa de Marianao (Bierig) [NMB] [Holotype
examined]. N. syn.
Additions to creightoni-group diagnosis. WL
0.61-0. 91mm. Clypeal apron convex. Eye small,
elliptical. Nuchal groove weakly visible in lateral
view. MHI 0.84-0.99. Pronotal shoulders well
rounded. Propodeal spines straight, inclined.
Propodeal spiracle less than 1 diameter from edge
of infradental lamella. Postpetiole long (PpetL/
PW 0.42-0.49), subtrapezoidal in dorsal view; ster-
num with long, flat profile and receding anterior
edge (except Dominican Republic). Both macro-
and microsculpture weak overall (stronger in
Dominican Republic). Rugae fine, low on head
and mesosoma (Fig. 93); vestigial on petiole. Erect
hair sometimes sparse and only slightly longer
than decumbent hair. Scapes have erect hairs;
extensor surfaces of legs do not.
Workers.— TL 2.3-3.3, HL 0.57-0.80, HW 0.45-
0.70, SL 0.36-0.52, EL 0.06-0.09 (9-18 facets), PW
0.35-0.49, WL 0.61-0.91, SpL 0.09-0.15, PetL 0.25-
0.34, PpetL 0.15-0.22mm, CI 0.79-0.87, OI 0.12-
0.16, SI 0.73-0.89, PSI 0.15-0.17, MHI 0.84-0.94.
N=16
Mandibles with 5-7 teeth; basal tooth equal to
or slightly larger than penultimate basal; a den-
ticle may occur between basal teeth. Body of
clypeus not projecting over apron. Posterior out-
line of head convex to weakly concave. Mesosoma
with broadly rounded shoulders; metanotal groove
Volume 3. 1 994
49
a weak impression or absent; no ridge at anterior
of propodeum. Propodeal spines narrow, moder-
ately long; a bisecting line passes dorsal to
propodeal spiracle and through or below
anteroventral corner of pronorum. Metapleural
lobes low, angular. Petiolar peduncle with strong
keel; node large and somewhat bulbous.
Postpetiolar node with long, low profile and
weakly defined posterior and anterior faces (more
distinct in Dominican Republic).
Laterodorsa and sides of head rugose-areolate;
in most Cuban specimens ridges become very
weak on sides, even absent in spots. Posterior
head with transversely arching rugose-areolate
macrosculp-rure. Anterior pronotal disc trans-
versely rugose to rugose-areolate; rest of prono-
rum longitudinally rugose, usually with incom-
plete cross-ridges between rugae. Mesonotum
more rugose-areolate. Pronotal sides basically rug-
ose (areolate-rugose in Dominican Republic);
meso- and metapleura confused rugose-areolate
with some smooth patches. Dorsal face of
propodeum transversely rugose. Vague rugosi-
ties on sides and /or posterior petiolar node;
postpetiolar node smooth.
Colorbrown to yellowish-brown, with lighter,
more yellowish appendages and frontoclypeal
region.
Queens.— TL 2.8-3.4, HL 0.59-0.74, HW 0.52-
0.65, SL 0.39-0.50, EL 0.11-0.15, PW 0.45-0.52, WL
0.81-0.95, SpL 0.10-0.16, PetL0.26 (estimated)-0.40,
PpetL 0.17-0.22mm, CI 0.86-0.89, SI 0.74-0.78, PSI
0.13-0.16, MHI (Cuban only; could not measure in
Bahamian) 0.97-1.06. N=3
Apparently collected as strays, so not defi-
nitely associated with workers described above.
The Bahamian queen distinctly smaller (all of the
smallest measurements above); Cuban queens
nearly identical in size. Mandible with 6 teeth;
basal and penultimate tooth subequal in size.
Clypeus evenly convex. Posterior outline of head
medially concave. Mesosoma of Cuban queens
robust, shaped like that of scobinata (Fig. 62), but
larger and with more prominent, angular
metapleural lobes. Mesosoma of Bahamian queen
seems like that of Cuban queens, but is obscured
by legs. Waist as in workers, except for lack of keel
on the petiolar peduncle in Cuban specimen.
PpetW/PpetL 1.05-1.14. Sculpture much as in
workers. Macrosculpture on sides of head weak.
Most of ventral half of mesopleura smooth. Pilos-
ity and color as in workers.
Discussion. — Workers of this species have two
disjunct sizes. In the smaller size range (WL 0.62-
0.70mm) are three specimens from Pinar del Rio
Province (including brunnea syntypes), one from
Soledad (scabra syntype), one from I. Pinas, three
from Dominican Republic, and one from Baha-
mas. The larger workers (WL 0.80-0. 91mm) are
from Sierra del Rosario (cubensis holotype),
Soledad, Playa Marianao (habanica holotype), Si-
erra Maestra, and Sierra Anale. Aside from size,
the smaller workers show little consistent differ-
ence from the larger workers, and in ways that
often vary within Rogeria species: 1) slightly more
macrosculpture on petiolar node, and 2) slightly
more extensive microareolate sculpture on sides
of mesosoma. The specimens from Dominican
Republic differ from the rest in having a more
compact mesosoma, more distinct postpetiolar
node and sternum in side view, and slightly stron-
ger sculpture.
Workers of creightoni from San Jose, Costa
Rica have the same size as larger brunnea and
much the same pilosity, nuchal groove, and shape
of mesosoma and petiole. Rogeria creightoni speci-
mens from Yucatan and Chiapas have size, eye
size, postpetiole, and pilosity like the larger brunnea,
and the Yucatan specimen has similar mesosoma
shape. But brunnea workers can be distinguished
by the combination of characters in the diagnosis.
Material Examined. — CUB A: Pinar del Rio Prov-
ince, Sierra de los Organos, Sierra del Rosario
(Bierig), Las Acostas (E. O. Wilson), San Vincente
(E. O. Wilson); Habana Province, Playa Marianao
(Bierig); Cienfuegos Province, Soledad (P. J.
Darlington; N. A. Weber; E. O. Wilson; D. Bates
and G. Fairchild); Las Villas Province, Caibanen
(E. O. Wilson); Sierra Anale (Bierig); Sierra Maestra,
Uvero (L. Armas). I. Pinas, S. Casas (L. B. Zayas).
DOMINICAN REPUBLIC: 16km ENE Pedemales
(P. S. Ward). BAHAMAS: Andros Island, Nassau
Island (W. M. Wheeler). 20 workers, 3 queens
[LACM, MCZ, NMB].
50
Journal of Hymenoptera Research
Rogeria creightoni Snelling
Figs. 51-54, 95-96
Rogeria creightoni Snelling 1973:2, Fig. 1. Holotype and
paratype worker, UNITED STATES: Texas, Cameron
County, La Fena (W. S. Creighton) [LACM] (Holotype
and paratype examined].
Additions to creightoni-group diagnosis. WL
0.63-0. 93mm. Clypeal apron usually with weak to
distinct median notch. Eyes small, oval to circular
(EL/SpL 0.29-0.64). Metanotal groove weak or
absent. MHI usually 0.90-1.06. Propodeal spines
long (PSI usually > 0.17), often nearly horizontal.
Postpetiolar node from above usually
subtrapezoidal, sometimes subrectangular; ante-
rior edge of sternum not prominent. Sides of head
distinctly rugose-areolate. Pronotal disc trans-
versely rugose-areolate on anterior edge and lon-
gitudinally rugose-areolate behind (Fig. 96). Scapes
and extensor surfaces of tibiae with erect-suberect
hairs.
Workers.— TL 2.4-3.5, HL 0.58-0.81, HW 0.51-
0.72, SL 0.38-0.55, EL 0.05-0.10 (8-16 facets), PW
0.37-0.52, WL 0.63-0.93, SpL 0.11-0.20, PetL 0.27-
0.40, PpetL 0.14-0.22mm, CI 0.85-0.91, OI 0.08-
0.17, SI 0.68-0.80, PSI 0.16-0.28, MHI 0.87-1.08.
N=21
Mandibles with 5-7 teeth, if 5, may have 1-2
additional basal denticles. Basal tooth subequal or
larger than penultimate basal. Palpal formula 3,2.
Body of clypeus projecting to anterior edge of
apron or beyond. Nuchal groove weak to strong.
Posterior margin of head weakly concave to con-
vex. Mesosoma habitus variable (Figs. 51-53).
Propodeal spines moderately long and angled to
very long and nearly horizontal, a bisecting line
usually passes well above anteroventral corner of
pronotum. Propodeal spiracle not especially large
or prominent; located < 1/2 diameter to almost
one diameter from edge of infradental lamella.
Metapleural lobes moderately to very prominent;
corner broadly to narrowly rounded. Petiolar node
more or less distinct (Figs. 51, 53). Petiolar keel
absent to moderately well developed. Sting appa-
ratus of specimens from four localities like that of
inermis (Fig. 42), except for less angular
anteroventral corner of spiracular plate, some-
what lower valve chamber height, and in Belize
and Oaxaca specimens the sting shaft is higher
and slightly upturned.
Laterodorsa of head longitudinally rugose to
areolate; posterior head transversely arched ar-
eolate-rugose. Pronotal sides longitudinally rug-
ose to areolate-rugose; meso- and metapleura con-
fused rugose (Fig. 95). Dorsal face of propodeum
marked anteriorly by a distinct transverse carina,
then areolate and /or transversely rugose. Head
and mesosoma microsculpture obscure; intervals
in macrosculpture moderately shiny. Top and sides
of petiolar node effaced microareolate with very
weak, indistinct macrosculpture; posterior face
with weak longitudinal rugulae. Postpetiolar node
with vague microsculpture on sides; smooth dor-
sally.
Color brownish-yellow to brownish-yellow;
dorsa of head and gaster slightly darker, append-
ages lighter.
Queen. — Uncertain; see discussion.
Discussion. — This species becomes quite het-
erogeneous with this revision. Ants from almost
every locality are different from the others in some
conspicuous way. Specimens from Belize have the
longest propodeal spines (PSI 0.24-0.28), more
prominent metapleural lobes, weak or absent peti-
olar keel and thicker ridges in macrosculpture.
The Texas specimens have similar habitus, but
have somewhat shorter propodeal spines (PSI 0.21-
0.23), less prominent metapleural lobes, more dis-
tinct petiolar keel and less thickened
macrosculpture. A Tamaulipas specimen is simi-
lar to the Texas specimens, but has longer, dis-
tinctly inclined propodeal spines. Specimens from
Yucatan, and Chiapas (Fig. 52) are the largest ( WL
0.78-0. 93mm), have a prominent clypeal body,
somewhat longer scapes, small circular eye, strong
nuchal groove, relatively shorter propodeal spines
(PSI 0.19-0.22), and head macrosculpture with
sharp ridges and unusually large areolae. Speci-
mens from La Selva, Costa Rica are similar, but
have normal clypeus and sculpture. Others from
Costa Rica (Fig. 53) have shorter, more inclined
propodeal spines (PSI 0.16-0.20), a more distinct
petiolar node and keel, and subrectangular
postpetiolar node. One specimen from Oaxaca is
more like those from Belize; another is more like
the San Jose specimens. With little material from
most localities, I did not feel confident naming a
new species for each variant, but as collections
improve, I will not be surprised if this species
undergoes fission.
At one time during the revision I also consid-
ered describing cornuta, innotabilis, leptonana, and
Volume 3, 1 994
51
alzatei as variants of creightoni. But now I believe
they can delineated. Comparison of creightoni with
leptonana is found in key couplet 40; comparisons
with conuita and alzatei, in their respective discus-
sions. All known specimens of innotabilis fall within
the geographic range of creightoni and the two
species are sympatric in at least one locality. All
specimens of creightoni differ from innotabilis (Figs.
55-57, 97-98) in several ways: 1) erect hair on
scapes and tibiae, 2) palpal formula 3,2, 3) clypeal
apron with weak to distinct median notch, and 4)
postpetiolar sternum not prominent. The regional
variants differ in additional, but inconsistent ways.
A single queen from Trinidad (N. A. Weber
#129) is very much like the three Costa Rican and
Colombian queens provisionally assigned to
innotabilis (Fig. 57) in size, general habitus, trian-
gular mandibles, convex clypeus, robust mesosoma
(MHI 1.28), propodeal spine size and shape, and
subrectangular postpetiolar node, but differs in
having erect hair on scapes and tibiae, little or no
keel on petiolar peduncle, undulate ventral profile
of postpetiole, and longitudinally oriented sculp-
ture on the posterior head. On the basis of the
pilosity, I provisionally assign the Trinidad queen
to creightoni, but because of its great similarity in
other respects to the innotabilis-like queens, and
because neither set of queens is from an area
where workers of creightoni or innotabilis have
been collected, I do not feel confident of these
assignments.
See also discussions of alzatei, belti, brunnea,
and cornuta.
Distribution. — Rogeria creightoni ranges from
southern Texas to Costa Rica; from about 1,500m
in parts of Chiapas and Costa Rica to sea level.
Types were collected in a residential area formerly
mesquite-acacia savannah. Other specimens come
from riparian woodland, palm-thorn forest, rain
forest, pine-oak forest, cecropia forest, and cacao
plantation. Most were taken in leaf litter on the
ground. Some Belize specimens were under a
termite nest; another in orchids.
Material Examined.— UNITED STATES: Texas,
Cameron County (W. S. Creighton) [LACM]; Live
Oak County (P. S. Ward) [mouthparts, sting]
[MCZ]. MEXICO: Tamaulipas, Antiguo Morelos
(S. & J. Peck); Vera Cruz, Pueblo Nuevo (E. O.
Wilson); Oaxaca, 1 mi. E Reforma, nr. Tuxtepec (A.
Newton) [mouthparts, sting]; 9 mi. E El Cameron
(A. Newton); Chiapas, 12 mi. NW Ocozocoautla
(A. Newton); Yucatan, Chichen Itza [MCZ].
BELIZE: Intercepted in Brownsville, Texas on or-
chids (Heinrich); nr. Belmopan (S. & J. Peck); Caves
Branch (S. & J. Peck) [mouthparts, sting] [BMNH,
CKC, MCZ, MZSP, USNM]. COSTA RICA:
Heredia Province, La Selva (Talbot &
VanDevender) [sting] [LACM], (D. M. Olson)
[DMOC]; San Jose (H. Schmidt) [MZSP];
Jurrucarres (A. Bierig) [MZSP]; no locality (F.
Nevermann) [MZSP]. 32 workers.
Rogeria innotabilis new species
Figs. 55-57, 97-98
Additions to creightoni-group diagnosis. WL
0.66-0. 73mm. Mandibles triangular to slightly
subtriangular. Palpal formula 2,2. Clypeal apron
convex or slightly flattened medially. Eye small,
elliptical. Nuchal groove clearly visible in lateral
view, but not notching ventral outline of head.
Mesosoma with nearly flat dorsal profile; a strong
carina marks anterior edge of propodeum; MHI
0.90-1.00. Propodeal spines straight, wide. Peti-
olar keel moderate to large. Postpetiole
subrectangular in dorsal view; sternum weakly to
strongly prominent. Spiracular plate of sting ap-
paratus with long spine on posteroventral corner.
Sides of head areolate-rugose; mesosoma predomi-
nantly rugose; microareolate sculpture distinct on
mesosoma sides. No erect hair on scapes or exten-
sor surfaces of tibiae.
Holotype and Paratope Workers. — TL 2.4-2.7
(2.7), HL b.60-0.65 (0.65), HW 0.50-0.57 (0.54), SL
0.40-0.45 (0.43), EL 0.06-0.08 (0.08) (7-13 facets),
PW 0.37-0.41 (0.41), WL 0.66-0.73 (0.72), SpL 0.10-
0.15 (0.13), PetL 0.26-0.30 (0.29), PpetL 0.14-0.16
(0.16)mm, CI 0.83-0.87 (0.84), 010.11-0.15(0.15), SI
0.76-0.80 (0.78), PSI 0.15-0.20 (0.18), MHI 0.90-1.00
(0.98) N=10
Mandible with 5-7 teeth. If 5 or 6 teeth, it may
have 1-3 denticles among proximal teeth. If 7 teeth,
4 proximals small. Basal tooth equal to or larger
than penultimate tooth. Body of clypeus not pro-
jecting to anterior edge of apron. Eye small, ellip-
tical. Posterior outline of head strongly to weakly
convex. Metanotal groove absent or weak.
Propodeal spines wide, straight; a bisecting line
passes through or below anteroventral corner of
pronotum. Petiolar keel weak or absent. Petiolar
node evenly rounded from front to back, or apex
somewhat flattened (Fig. 55). Postpetiolar node
52
Journal of Hymenoptera Research
highest in posterior half; node in dorsal view
much as in Fig. 53. Sting apparatus like that of
inermis (Fig. 42), except for spiracular plate shape
(Fig. 56).
Head macrosculpture areolate on laterodorsa;
areolate to transversely arching rugose-areolate
on posterior head. Areolae on posterior head are
larger and ridges sharper than elsewhere on head.
Head dorsum slightly dulled by vestigial
microsculpture; intervals smooth and shiny on
sides and back. Anterior pronotal disc with 1-2
transverse rugae; rest of promesonotal dorsum
longitudinally rugose with lateral spurs (Fig. 98)
to rugose-areolate. Lateral pronotum areolate;
meso- and metapleura mixed rugose and rugose-
areolate. Anterior edge of propodeum marked by
weak to strong transverse carina; dorsal face
crossed by one or more transverse, sometimes
branching, rugulae. Dorsum with vestigial
microsculpture, but ventral sides quite distinctly
microareolate (Fig. 97). Macrosculpture on sides
and back of petiolar node very weakly areolate;
microsculpture vague and effaced. Postpetiolar
node shiny, almost completely smooth.
Color golden brown with yellowish append-
ages to brown with slightly lighter frontoclypeal
area, mandibles, mesosoma sides, gaster, and legs.
Possible Queens.— TL 2.6-2.9, HL 0.59-0.61 , H W
0.52-0.54, SL 0.40-0.41, EL 0.11-0.13, PW 0.44-0.50,
WL 0.72-0.83, SpL 0.13-0.14, PetL 0.28-0.31, PpetL
0.14-0.17mm, CI 0.88-0.89, OI 0.21-0.24, SI 0.75-
0.77, PSI 0.16-19, MHI 1.15-1.22. N=3
Collected outside known range of innotabilis
workers, but have innotabilis traits: shapes of
clypeus, nuchal groove, and postpetiole, and lack
of erect /suberect hair on scapes and tibiae.
Differing from the worker by normal caste
differences and the following. Parapsidal furrows
indistinguishable from furrows in sculpture.
Pronotum with 2-3 transverse rugae mesad, be-
coming longitudinally rugose to rugose-areolate
on mesosoma sides; anterior katepisternum
smooth. Mesonotum longitudinally rugose. Ve-
nation nearly identical to that of belti (Fig. 37).
Discussion. — For comparison with related spe-
cies, see creightoni, leptonana, and alzatei discus-
sions. Remarks about the queens are found in the
creightoni and curvipubens discussions.
The name innotabilis (L., not remarkable) re-
flects my inability to identify any one salient fea-
ture on which to base the name for this species.
Distribution. — If the queens truly belong to
innotabilis, this species ranges from Chiapas to
northern Colombia, but workers have not been
collected south of Nicaragua. Workers come from
moist forest litter at 1000-1200m elevation. Collec-
tion sites for queens range from 10-1200m.
Material Examined. — Holotype locality.
MEXICO: Chiapas State, 12 mi. NW Ocozocoautla,
3200 ft., 4-5-IX-1973, forest leaf litter (A. Newton)
[MCZ],
Paratype Localities. MEXICO: 7 workers, ho-
lotype locality [mouthparts, sting, 1 coated for
SEM] [BMNH, MCZ, MZSP]; 10 workers, Chiapas,
6-XII-1951, #51 11574 (Cary) [2 mouthparts, stings]
[USNM], NICARAGUA: 2 workers, Km 139 near
Matagalpa, Hotel Selva Negra, ca. 1200m, 18- VI-
1978, forest leaf litter (C. Kugler & J. Hahn) [mouth-
parts, sting] [LACM, MCZ].
Nontype localities. COSTA RICA: Puntarenas
Province, Monteverde, 1200m, 23-V-1979, #3496
(P. Ward) [MCZ], COLOMBIA: Magdalena De-
partment, Parque Tayrona, 210m, l-X-76 (C.
Kugler) [MCZ]; Guajira, Don Diego, 25-50m, 22-
VI-76 (W. L. Brown & C. Kugler) [queen mouth-
parts, sting] [MCZ]. 3 queens.
Rogeria alzatei new species
Figs. 58-60, 99
Diagnosis. — WL 0.51-0. 70mm, most < 0.65mm.
Mandibles subtriangular (usually) to triangular.
Clypeus usually truncate. Eye usually moderately
large, oval (10-14 facets, OI 0.17-0.20 in types), but
occasionally half as large and elliptical. Nuchal
groove weak to strong. Mesosoma compact (MHI
0.90-1.07). Propodeal spines usually slender, in-
clined. Petiolar peduncle with little or no keel.
Postpetiolar node from above usually
subrectangular; anterior edge of sternum not
prominent. Macrosculpture areolate on head sides.
Posterior head transversely rugose-areolate (Fig.
99). Pronotal disc rugose to rugose-areolate. No
erect hair on scapes or legs.
Holotype and Paratype Workers. — TL 1.9-2.2
(2.2), HL 0.51-0.56 (0.54), HW 0.45-0.48 (0.47), SL
0.32-0.36 (0.35), EL 0.07-0.09 (0.09) (11-14 facets),
PW 0.32-0.37 (0.35), WL 0.51-0.61 (0.59), SpL 0.10-
0.12 (0.11), PetL 0.21-0.26 (0.23), PpetL 0.12-0.14
(0.13)mm, CI 0.87-0.88 (0.87), OI 0.17-0.20 (0.19), SI
0.72-0.74 (0.74), PSI 0.19-0.21 (0.19), MHI 1.01-1.07
(1.05). N=ll
Volume 3. 1 994
53
Mandible subtriangular; 5-6 teeth, two basal teeth
small, subequal. Palpal formula 2,2. Clypeal apron
truncate, with distinct corners; body of clypeus
does not project to edge of apron. Posterior outline
of head continuously convex. Nuchal groove in-
distinct in lateral view. Mesosoma's evenly con-
vex dorsal profile ends abruptly fore and aft by
rather sharp angles and by transverse sculpture.
Metanotal groove weak or absent. Propodeal spines
straight, narrow; a bisecting line passes through or
just above anteroventral corner of pronotum. Cor-
ner of metapleural lobes a narrowly rounded right
angle. Petiolar node evenly rounded front to back.
Pygidium in the five specimens dissected with no
piligerous tubercles.
Head laterodorsa rugose-areolate to areolate;
sides more coarsely areolate. Microsculpture ves-
tigial, producing irregular intervals on head dor-
sum and nearly smooth intervals on sides and
posterior (Fig. 99). Anterior edge of pronotal disc
with two transverse rugae. Promesonotal dorsum
longitudinally rugose with numerous lateral spurs
that sometimes connect, forming areolae. No
macrosculpture on dorsal face of propodeum.
Lateral pronotum longitudinally rugose-areolate.
Meso- and metapleura with scattered, irregular
rugae. Vestigial rugae on sides and back of peti-
olar node. Vague microsculpture makes rough-
ened intervals on mesosoma, petiole and much of
postpetiole; apex of postpetiolar node smooth.
Body yellowish-brown to reddish-brown with
slightly darker head and gaster dorsa (black to
naked eye); legs and antennae lighter, more yel-
lowish or yellowish-brown.
Nontype Workers.— TL 1.9-2.5, HL 0.50-0.66,
HW 0.42-0.56, SL 0.31-0.46, EL 0.05-0.10 (4-21 fac-
ets), PW 0.29-0.39, WL 0.51-0.70, SpL 0.09-0.13,
PetL 0.21-0.28, PpetL 0.11-0.15mm, CI 0.81-0.88,
OI 0.08-0.20, SI 0.71-0.85, PSI O.15-0.21, MHI 0.90-
1.07. N=32
Specimens from Guyana, French Guiana, and
the Caribbean coast of Colombia extend the de-
scription as follows: Mandibles triangular with 3-
4 teeth plus 4 basal denticles. Clypeal apron evenly
convex, sometimes with medial emargination.
Some with only 7-8 facets in the eyes. Some with
wider propodeal spines (Fig. 60) or slightly
downturned tips. Petiolar keel sometimes distinct
(Fig. 60). Macrosculpture on head or pronotum
more areolate.
Dominican Republic specimens have a con-
vex clypeus, a distinct metanotal groove as in Fig.
46, and subtrapezoidal postpetiolar node as in Fig.
51.
Specimens from Peru, Brazil, and Paraguay
extend the description somewhat differently: Pal-
pal formula 3,2. Clypeal apron usually truncate
with rounded corners; occasionally convex. Sev-
eral specimens with a somewhat prominent clypeal
body (Fig. 59). Generally with smaller, elliptical
eyes (OI of most 0.10-0.16), but the four workers
from the Puerto Maldonado vicinity of Peru pos-
sess both the smallest eyes (Fig. 59) and largest
eyes of the species (4 and 21 facets, OI 0.08 and
0.20). Nuchal groove forms a distinct notch in the
head of one Peruvian worker (Fig. 59). Mesosoma
generally less compact (MHI 0.93-1.03). Petiolar
keel weak to distinct (Fig. 59). Some with
postpetiolar node slightly wider in anterior half
and shape similar to Fig. 66. Posterior pygidium
with small tubercles in a Paraguayan specimen
dissected. Sting apparatus with reduced
anterodorsal corner of quadrate plate in a Brazil-
ian worker.
Paratype and Nontype Queens. — TL 2.4-2.8, HL
0.56-0.60,"hW 0.51-0.53, SL 0.35-0.38, EL 0.13-0.14,
PW 0.44-0.49, WL 0.70-0.79, SpL 0.14-0.15, PetL
0.26-0.30, PpetL 0.15-0. 1 6mm, CI 0.88-0.92, OI 0.25-
0.26, SI 0.70-0.72, PSIO.19-0.20, MHI 1.17-1. 18. N=2
Both collected in the same localities as alzatei
workers (BCI, Panama; Quebrada Susumuco, Co-
lombia), but not in nest series. They most resemble
alzatei workers in characteristic shapes of clypeus,
mandible, mesosoma, propodeal spines, petiole
and postpetiole, as well as sculpture and pilosity.
Mesoscutellum projects slightly beyond posterior
edge of metanotum. Posterior head rugose-ar-
eolate; no tubercles. Middorsum of head covered
with erect hairs.
Discussion. — The extent of variation within
this species makes me question whether this is a
single species, but external characters that can be
used in a key are either not marked enough to
discount individual variation or not concordant.
Further complicating the picture is the surprising
amount of variation within four specimens from
the same region of Peru, and unique sets of fea-
tures found in single specimens from northern
Colombia, Rio de Janeiro, and Peru. Specimens
from Dominican Republic strongly resemble
creightoni workers from the same localities, but
differ in lacking erect hair on the scapes and tibiae
54
Journal of Hymenoptera Research
and in having narrower propodeal spines.
Given the geographic variation within alzatei,
the species as a whole is difficult to distinguish
clearly from creightoni, innotabilis, and leptonana.
But if one compares only those specimens oialzatei
that are in sympatry or parapatry with those three
species, the differences are clear. The Panamanian
alzatei workers differ from Central American
creightoni, innotabilis, and leptonana in having: 1)
subtriangular mandibles, 2) truncate clypeal apron,
3) larger, elongate-oval eye (OI 0.17-0.20), 4) more
compact mesosoma (MHI 1.01-1.07), and 5) peti-
olar peduncle without a keel. They are also smaller
than creightoni and innotabilis workers (WL 0.51-
0.61mm vs. 0.66-0. 93mm in creightoni and 0.66-
0.73mm in innotabilis). Additional differences from
leptonana are palpal formula (2,2 vs. 3,2 in leptonana),
longer propodeal spines (PSI 0.19-0. 20 vs. < 0.16 in
leptonana), and lack of erect hair on scapes.
See also the description of sibling species
scobinata and the discussions of belli, curvipubens,
micromma, tribrocca and merenbergiana.
The species is named in honor of a courageous
Colombian intellectual and friend, Isaac Alzate.
Distribution. — The range of alzatei is from
Panama to Paraguay, from western slopes of the
Andes in Colombia to the Eastern coast of Brazil.
All specimens were apparently taken as strays or
in Berlese or Winkler sampling in moist forests
from sea level to 1000m.
Material Examined. — Holotype locality.
PANAMA: Canal Zone, Barro Colorado Island
(BCI), 2-VII-79, W. L. Brown [MCZ].
Paratype localities. PANAMA: 3 workers, 1
queen from holotype locality [mouthparts, sting, 1
worker coated for SEM] [CKC, MCZ]; 2 workers,
BCI, IV-V-1942, #4953 (J- Zetek) [USNM]; 2 work-
ers, BCI, VI-X-1943, #5105 (J. Zetek) [whole speci-
men slide mounted] [USNM]; 1 worker, BCI, I-
1960 (W. L. Brown & E. S. McCluskey) [MCZ]; 1
worker, BCI, 7-III-1975, FP#10 (C. Toft & S. Levings)
[LACM]; 1 worker, Punta de los Chivos, W side
Gatun Lake, 3km SW Gahin, 100m, 3-9-VII-1979
(W. L. Brown) [MCZ]; 1 worker, Bocas del Toro,
Pipeline Road, 300m, 8.53N 82.10W, 18-VII-1987,
#633 (D. M. Olson) [MZSP].
Nontype localities. DOMINICAN REPUBLIC:
16km ENE Pedernales, 800m (P. S. Ward). CO-
LOMBIA: Guajira, Rio Don Diego (W. L. Brown &
C. Kugler); Magdalena Department, San Pedro,
550m (P. Ward) [MCZ]; Choco Department, 10km
SW San Jose del Palmar, Rio Torito, Finca los
Guaduales, 800m (C. Kugler) [mouthparts, sting]
[CKC, MCZ, MZSP]; Antioquia Department,
Providencia, Estacion Biologico, 600-800m, (C.
Kugler) [MCZ]; Cundinamarca Department,
Bogota to Villavicencio, Km 79 (W. L. Brown & I.
del Polania) [MCZ]; Meta Department, Bogota to
Villavicencio, 23km NW Villavicencio, Quebrada
Susumuco, 1000m (S. & J. Peck; C. Kugler) [1
mouthparts, 2 stings, 1 whole specimen] [MCZ].
GUYANA (British Guiana): Oronque River (N. A.
Weber) [MCZ]. BRAZIL: Sao Paulo State, Agudos
(W. Kempf; C. Gilbert) [mouthparts, sting] [MZSP,
USNM]; Rio de Janeiro (T. Borgmeier) [MZSP];
Federal District (C. A. C. Seabra) [MZSP]. PERU:
Madre de Dios Department, Puerto Maldonado
vicinity, 260m (C. Kugler) [CUIC, MCZ]. PARA-
GUAY: Central Province (F. Baud, et al.) [mouth-
parts, sting] [BMNH, MCZ (voucher), MHN];
Concepcion, and Paraguari Provinces (F. Baud, et
al.) [BMNH, MHN]. 62 workers, 1 queen.
Rogeria scobinata new species
Figs. 61-62, 100
Diagnosis. — As inalzatei, except the following:
Clypeal apron truncate (none emarginate). MHI
0.96-1.14. Posterior head with tuberculate
macrosculprure (Fig. 100). Erect hair usually ab-
sent from head dorsum; if present, it is short and
usually confined to posterior margin.
Holotype and Parati/pe Workers. — TL 1.9-2.5
(2.2), HL 0.50-0.60 (0.55), HW 0.42-0.52 (0.475), SL
0.31-0.40 (0.35), EL 0.06-0.09 (0.07) (10-14 facets),
PW 0.30-0.40 (0.35), WL 0.50-0.67 (0.585), SpL 0.08-
0.12 (0.095), PetL 0.19-0.26 (0.23), PpetL 0.12-0.16
(0.14)mm, CI 0.82-0.86 (0.86), OI 0.14-0.18 (0.15), SI
0.74-0.78 (0.74), PSI 0.16-0.19 (0.16), MHI 0.96-1.09
(1.00). N=5
Nontype Workers.— TL 1.9-2.5, HL 0.50-0.61,
HW 0.44-0.52, SL 0.31-0.40, EL 0.06-0.09 (7-15 fac-
ets), PW 0.30-0.40, WL 0.51-0.68, SpL 0.08-0.13,
PetL 0.19-0.28, PpetL 0.12-0.17mm, CI 0.84-0.88,
OI 0.14-0.18, SI 0.70-0.78, PSI 0.16-0.20, MHI 1.00-
1.14. N=32
Mandibles subtriangular, 5-toothed (some-
times with 1-2 additional basal denticles), decreas-
ing in size basad; basal tooth small. Palpal formula
2,2. Median clypeus of some nontype workers
from Colombia like that of alzatei (Fig. 58), but type
specimens with less prominent corners (Fig. 61)
Volume 3, 1994
55
and other nontypes (Bolivia, some Brazil) have an
almost evenly convex clypeal apron. Body of
clypeus not projecting over clypeal apron. Poste-
rior outline of head weakly concave medially to
weakly convex. Nuchal groove clearly visible in
side view. Eye oval to elliptical. Anterior and
dorsal faces of pronotum may join smoothly, or in
a weak angle (Fig. 61). Metanotal groove broad,
slightly less to slightly more impressed than shown
in Fig. 61, bordered behind by a transverse carina.
Propodeal spines inclined. Metapleural lobes
moderately prominent; corner varies from sharply
angular (Ecuador, some Peru) to rounded as in
Fig. 44 (some Paraguay). Ventral petiolar peduncle
usually with a weak, nonlamellate keel, but some
Ecuadorian specimens with distinct keel.
Postpetiolar node in dorsal view subrectangular
as in Fig. 58. Pygidium with a pair of median,
columnar, piligerous tubercles near caudal edge
(barely visible in dissection microscope at 50X).
Laterodorsa and sides of head densely ar-
eolate. Posterior head with short triangular, blunt
tubercles in more or less distinct rows (Fig. 100).
Tuberculate sculpture usually extends across pos-
terior quarter of head, but in a few specimens from
Leticia, Benjamin Constant, and Paraguay, the
ridges between the tubercles are not always com-
pletely lost, so the posterior head appears mostly
fragmented-rugose, with only a few of the trian-
gular tubercles. Interstices on most of head some-
what dulled by indistinct areolate microsculpture,
but smoother and quite shiny between tubercles
on back of head; sides sometimes rather strongly
microareolate. Anterior edge of pronotal disc with
1-4 more or less transverse rugae. Rest of
promesonotum longitudinally rugose with fre-
quent incomplete lateral spurs. Mesosoma sides
weakly and sparsely rugose to rugose-areolate,
but more strongly microareolate than on pronotal
disc. Dorsal face of propodeum usually lacking
macrosculpture, but rather strongly microareolate.
Rest of mesosoma with indistinct microareolate
sculpture. Petiolar node with broken vestigial
macrosculpture. Postpetiole without macrosculp-
ture; nearly smooth on top. Sides of nodes with
weak microareolate sculpture that imparts a granu-
lar appearance; microsculpture usually weaker on
postpetiole.
Workers from Leticia have 8-10 erect hairs
along posterior rim of head and those from Ben-
jamin Constant have sparse, short erect hairs on
the posterior rim and along the midline.
Color brown to golden brown. Legs and an-
tennae generally lighter than rest of body; gaster
sometimes darker.
Queens.— TL 2.4-2.6, HL 0.55-0.58, HW 0.46-
0.52, SL 0.34-0.38, EL 0.11-0.13, PW 0.41-0.45, WL
0.68-0.74,SpL0.11-0.14,PetL0.25-0.27,PpetL0.14-
0.16mm, CI 0.84-0.90, SI 0.70-0.75, PSI 0.16-0.19,
MHI 0.64-0.69. N=7
Habitus shown in Fig. 62. Parapsidal furrows
indistinguishable from grooves in sculpture. An-
terior pronotum transversely rugose to rugose-
areolate, becoming longitudinal on sides.
Mesoscutum with longitudinal, often diverging
rugae; mesoscutellum rugose or rugose-areolate.
Discussion. — Rogeria alzatei is a sibling species
of scobinata, with which it is sympatric in Peru,
Brazil, and Paraguay, but can be distinguished by
characters in the diagnosis and key. The pair of
columnar tubercles on the pygidium may also be
distinctive.
The name scobinata, meaning having the na-
ture of a rasp, refers to the sculpture on the poste-
rior head, which has rows of teeth like a rasp.
Distribution. — Rogeria scobinata ranges from
the north coast of South America to Paraguay at
elevations below 1000m. All specimens were taken
as strays in tropical forest, mostly by Berlese or
Winkler sampling of leaf litter, rotten wood, or
moss.
Material Examined. — Holotype locality. PERU:
Madre de Dios Department, 3km N Puerto
Maldonado, 260m, primary forest remnant by side
of road, berlesate of leaf litter and rotten wood, 13-
16-VI-1981 (C. Kugler and R. R. Lambert) [MCZ].
Paratype localities. PERU: 2 workers, 1 queen,
holotype locality [MCZ]; 22 workers, 2 queens,
5km E Puerto Maldonado on Rio Tambopata, Finca
Medina, 260m, primary forest berlesate, 13-16- VI-
1981 (C. Kugler and R. R. Lambert) [mouthparts,
sting, 1 worker coated for SEM] [BMNH, CKC,
LACM, MCZ, MZSP, MHN, USNM].
Nontype localities. TRINIDAD: Nariva
Swamp (N. A. Weber) [MCZ]. SURINAM: La
Poulle, Vank, and Tambahredjo (J. van der Drift)
[MZSP]. COLOMBIA: 7km N Leticia (S. and J.
Peck) [MCZ]. ECUADOR: Napo Province,
Limoncocha and 20km S Tena (S. and J. Peck);
Pastaza Province, 25km N Puyo, (S. and J. Peck)
[MCZ]. PERU: Loreto Department, Ramon Castillo
(S. and J. Peck), 15km WSW Yurimaguas (P. S.
Sb
Journal of Hymenoptera Research
Ward) [PSWC]; Pasco Department, near Pozuzo,
(C. Kugler and R. R. Lambert); Madre de Dios
Department, Puerto Maldonado vicinity near Lake
Sandoval (C. Kugler and R. R. Lambert) [CKC,
CUIC, MCZ]. BOLIVIA: La Paz Department,
Caranavi (C. Kugler and R. R. Lambert) [MCZ].
BRAZIL: Para State, (collector name unreadable);
Amazonas State, Benjamin Constant and vicinity
(W. L. Brown); Mato Grosso State, Utiariti, Rio
Saueruina [=R. Papagaio] (Lenko and Pereira);
Sao Paulo State, Piracununga Municipality,
Cachoeira das Emas (EEBP) (W. L. Brown); Rio de
Janeiro (T. Borgmeier) [MCZ, MZSP]. PARA-
GUAY: Alto Parana, Amambay, Caaguazii,
Canendiyu, and Concepcion Provinces (F. Baud et
al.). 122 workers, 13 queens.
Rogeria tribrocca new species
Fig. 63
Additions to creightoni-group diagnosis. WL
0.62mm. Eye small, elliptical. Clypeal apron with
three acute processes. Metapleural lobes very low,
broadly rounded. Postpetiolar node widest in an-
terior half. Posterior edge of gaster T3 with me-
dian spine that seems to arise from a weak concav-
ity. Sides of head smooth. Promesonotal dorsum
with fine rugae diverging and branching from
anterior pronotum, becoming weak on meso- and
metanota; interrugal spaces rather wide. No erect
hair on scapes or legs.
Holotype Worker.— TL 2.3, HL 0.61, HW 0.51,
SL 0.40, EL 0.05 (7 facets), PVV 0.36, WL 0.62, SpL
0.10, PetL 0.26, PpetL 0.13mm, CI 0.83, OI 0.10, SI
0.78, MHI 0.93, PSI 0.16.
One mandible with six teeth; the other with
the penultimate basal replaced by two denticles
(Fig. 63). Nuchal groove forms a weak notch in
lateral view of head. Pronotal shoulders seen from
above subangular. Propodeal spiracles 1 diameter
from nearest edge of infradental lamella. Ventral
petiole somewhat obscured by glue, but probably
with distinct keel and dentate process. Petiolar
node narrow; nearly symmetrically rounded in
lateral view. Postpetiolar node with broadly
confluent anterior and dorsal faces and short pos-
terior face. Postpetiolar sternum with perpendicu-
lar anterior corner and flat ventral profile (Fig. 63);
node in dorsal view widest in anterior half, as in
Fig. 74. Sting shaft projecting from gaster acute,
with low dorsal flange.
Laterodorsa of head longitudinally rugose-
areolate. Posterior head with transversely arching
rugae. Intervals with weak microsculpture. Dor-
sal face of propodeum with 3 weak transverse
rugulae. Mesosoma sides with sparse, indistinct,
longitudinal rugulae. Microsculpture on
mesosoma even weaker than on head; interrugal
spaces shiny, nearly smooth. Petiolar and
postpetiolar nodes nearly smooth, shiny.
Mesosoma dorsum with eight pairs of erect
hairs (some matted down); nodes each with two
pairs of posterodorsally projecting hairs.
Color of body and appendages brownish-yel-
low; dorsum of gaster Tl slightly darker.
Discussion. — Boihciiri'ijntbens (Figs. 74-76, 101-
102) and alzatei (Figs. 58-60, 99), have also been
collected in Quebrada Susumuco or nearby
Villavicencio environs. But they do not have a
three-toothed clypeal apron or a median spine on
the third tergum of the gaster.
The name tribrocca (brocca L., projection of
teeth) refers to the three tooth-like projections of
the clypeal apron.
Material Examined. — Holotype locality. CO-
LOMBIA: Meta Department, Quebrada Susumuko
[or Susumuco], 23km NW Villavicencio, 1000m,
forest litter, 5-III-1972, #B-234 (S. and J. Peck)
[MCZ].
Rogeria carinata new species
Fig. 64
Rogeria curvipubens: M. Smith 1937:851. Misidentification.
Additions to creightoni-group diagnosis. WL
053-0.67mm. Clypeal apron convex. Eye small.
Propodeal spines small. Mesosoma low (MHI 0.86-
0.96). Postpetiole subrectangular in dorsal view;
sternum with distinct, but not projecting anterior
lip. Sides of head rugose. Promesonotal dorsum
carinate, with broad, shiny intervals. Short erect-
suberect hairs on scapes, but not on extensor sur-
faces of legs.
Holotype and Parati/pe Workers. — TL 2.0-2.5
(2.2), HL 0.51-0.61 (0.55), HW 0.44-0.54 (0.46), SL
0.35-0.42 (0.38), EL 0.05-0.06 (0.06) (7-10 facets),
PW 0.31-0.41 (0.34), WL 0.53-0.67 (0.57), SpL 0.05-
0.10 (0.08), PetL 0.19-0.24 (0.22), PetW 0.10-0.13
(0.11), PpetL 0.11-0.14 (0.12), PpetW 0.14-0.18
(0.15)mm, CI 0.86-0.89 (0.86), OI 0.11-0.12 (0.13), SI
0.78-0.80 (0.83), PSI 0.09-0.15 (0.14), MHI 0.86-0.96
(0.92). N=15
Volume 3. 1 994
57
Mandibles with 5 teeth and 0-2 denticles be-
tween the two basal teeth; basal tooth larger than
penultimate basal. Eyes oval to elliptical. Nuchal
groove weak. Posterior outline of head weakly
convex to weakly concave. Mesosoma from above
broad, angular at shoulders, but strongly con-
stricted behind; metanotum half as wide as shoul-
ders. Propodeal spiracle small, one diameter from
posterior edge of propodeum. Metapleural lobes
small, rounded. Ventral petiole with weak keel.
Postpetiole highest in posterior half. Sting appara-
tus with slightly lower valve chamber than that of
inermis (Fig. 42).
Laterodorsa of head rugose-areolate in Puerto
Rican specimens. Posterior head transversely
arched areolate-rugose. Interrugal spaces on head
weakly and irregularly roughened on dorsum;
smoother on sides and posterior. Anterior face of
pronotum with transverse carinules that continue
onto shoulders and curve dorsad. Meso- and
metapleura with irregular longitudinal rugae; in-
tervals smooth dorsally, but vestigially
microareolate below level of propodeal spiracle.
Dorsal face of propodeum mostly smooth with 0-
3 transverse rugulae. Petiolar node vaguely
microareolate; macrosculpture absent or indis-
tinct on sides. Postpetiole vaguely microareolate
on venter; smooth and shiny on dorsum. Gaster
smooth and shiny.
Mesosoma dorsum with 10 pairs of erect hairs.
Color brownish-yellow; head and mesosoma
slightly darker than rest of body. Some Puerto
Rican specimens slightly darker, with reddish hue
to head and mesosoma.
Discussion. — The carinate macrosculpture of
the promesonotum with very shiny intervals serves
to differentiate carinata from others of the creightoni-
group. See also the discussion of nevadensis, which
has similar sculpture.
Material Examined. — Holotype locality. BRIT-
ISH VIRGIN ISLANDS: Tortola, Long Bay,
berlesate from soil and leaves, 25-VII-1965 (I. Proj.
Staff) [MCZ].
Paratype localities. 1 worker, holotype local-
ity [MCZ]. PUERTO RICO: 3 workers, Guanica, 7-
IV-1982, #75 (J. A. Torres) [LACM]; 1 worker,
Cayo Ratones, 17-VII-1982, #77 (J. A. Torres)
[LACM]; 8 workers, Cayo L. Pena, Humacao, 9-X-
1982, #91 (J. A. Torres) [2 stings] [BMNH, CKC,
LACM, MCZ, MZSP]; 1 worker, Ensenada, #1409,
(no collector) [USNM].
Rogeria nevadensis new species
Fig. 65, 94
Additions to creightoni-group diagnosis. WL
0.58-0.65mm. Clypeal apron emarginate. Eye small.
Shoulders rounded; promesonotum and propo-
deum meet without a strong angle. Postpetiolar
node widest in anterior half. Sides of head largely
smooth and shiny. Promesonotal dorsum longitu-
dinally rugose, with low, rounded ridges and no
lateral spurs; interrugal spaces wide and nearly
smooth. No erect hair on scapes or extensor sur-
faces of legs.
Holotype and Paratype Workers.— TL 2.2-2.5 (2.2),
HL 0.59-6.62 (0.59), HW 0.49-0.51 (0.49), SL 0.40-
0.42 (0.40), EL 0.06-0.08 (0.06) (7-10 facets), PW
0.34-0.38 (0.34), WL 0.58-0.65 (0.58), SpL 0.09-0.10
(0.09), PetL 0.22-0.25 (0.22), PpetL 0.12-0.14
(0.12)mm, CI 0.81-0.83 (0.82), 010.13-0.15(0.13), SI
0.82, PSI 0.15, MHI 0.93-0.95 (0.93). N=3
Mandibles with 5 teeth plus two denticles, or
six teeth plus 1-2 denticles among basal teeth;
basal tooth not larger than penultimate basal. Eye
elliptical. Nuchal groove weak. Promesonotum
slopes to join dorsal face of propodeum without an
angle or transverse carina; metanotal groove weak
or absent. Propodeal spines narrow; a bisecting
line passes through anteroventral corner of prono-
tum. Propodeal spiracle within one diameter of
edge of infradental lamella. Petiolar peduncle with
indistinct keel. Postpetiolar node with a short
vertical anterior face and long, sloping dorsal face;
posterior face slightly concave in two paratypes.
Postpetiole widest in anterior half as in Fig. 66.
Postpetiolar sternum with flat ventral profile and
somewhat prominent anterior edge. Sting shaft
projecting from apex is like that otinermis (Fig. 42).
Laterodorsa of head rugose-areolate. Poste-
rior head with transversely arching rugae. Head
shiny with effaced microsculpture, especially on
sides and back. Anterior pronotum crossed by
several incomplete rugae. Lateral mesosoma (Fig.
94) with little distinct macrosculpture and smooth,
shiny intervals. Dorsal face of propodeum with
transverse rugae and effaced microareolate sculp-
ture. Both nodes with weak, effaced microareolate
sculpture and vague, vestigial rugose
macrosculpture.
Mesosoma dorsum with 10-11 pairs of erect
hairs; 2-3 pairs project posterodorsally from each
node.
58
Journal of Hymenoptera Research
Color brown to dark reddish-brown
(mesosoma and head dorsum may be darkest),
with brownish-yellow appendages and
frontoclypeal area.
Gynecoid Worker. — A nontype specimen is
worker-like in most respects, but is larger (WL
0.70mm; PSI 0.19), has distinct mesonotal and
metanotal sutures and partially formed wing at-
tachments. Pronotum transversely rugose;
mesoscutum longitudinally rugose. Sides of
postpetiolar node from above more evenly con-
vex.
Discussion. — Rogeria merenbergiana (Figs. 46-
48), which is also found at high elevations in
Colombia and Ecuador, has similar shapes of
clypeus, mesosoma and propodeal spines, and
somewhat reduced sculpture, but: 1) are a little
larger than nevadensis (WL 0.69-0. 83mm), 2) have
a metanotal impression, 3) have erect hair on
second and third tibiae and scapes, and 4) are
more distinctly sculptured, with sides of head
rugose.
Rogeria carinata (Fig. 64) from Puerto Rica and
Tortola have the same size, pilosity, shape of waist,
and extensive shiny spaces on mesosoma, but: 1)
the metanotal-propodeal junction is more angu-
lar, 2) the sides of the head are distinctly rugose, 3)
the promesonotal dorsum has straighter parallel
ridges and smoother intervals, and 4) the propodeal
spines are generally shorter (PSI 0.11-0.15).
Rogeria nevadensis takes its name from the
Sierra Nevada de Santa Marta, site of the type
localities.
Material Examined. — Holotype locality. CO-
LOMBIA: Magdalena Department, SE Santa Marta,
Cuchilla San Lorenzo, vicinity of El Campano,
1340m, rain forest, VI-1976 (W. L. Brown) [MCZ].
Paratype localities. 1 worker, same site as
holotype [coated for SEM] [MCZ]. 1 worker, CO-
LOMBIA: Magdalena, E Orihueca, 74.03W 10.55N,
San Pedro de la Sierra, 1300m, rotten log in coffee
plantation, 10-11-1977 (C. Kugler) [LACM].
Nontype locality. One gynecoid worker,
berlesate at the San Pedro de Sierra site (C. Kugler)
[MCZ].
Rogeria leptonana new species
Figs. 66-70
Additions to creightoni-group diagnosis. WL
0.54-0. 66mm. Median clypeal apron concave to
convex. Eye small, nearly circular. Propodeal spines
small to absent. Mesosoma low, slender (MHI
0.78-0.90; PW/WL 0.53-0.58). Petiole with moder-
ate to large keel. Postpetiolar node from above as
in Fig. 66. Anterior lip of postpetiolar sternum not
prominent. Sides of head distinctly macro- or
microsculptured. Mesosoma rugose, sometimes
with strongly microareolate sides. Erect hair on
scapes and usually on extensor surfaces of tibiae.
Holotype and Paratype Workers. — TL 2.1-2.3
(2.1), HL 0.50-0.55 (0.51), HW 0.43-0.46 (0.44), SL
0.32-0.34 (0.32), EL 0.05-0.07 (0.06) (7-10 facets),
PW 0.32-0.36 (0.33), WL 0.57-0.61 (0.57), SpL 0.07-
0.10 (0.07), PetL 0.20-0.23 (0.21), PpetL 0.12-0.13
(0.13)mm, CI 0.83-0.86 (0.86), OI 0.11-0.15 (0.14), SI
0.73-0.75 (0.73), PSI 0.12-0.16 (0.12), MHI 0.83-0.90
(0.83). N=4
Mandibles triangular, with 5 teeth and occa-
sional denticle; basal tooth larger than penultimate
tooth. Palpal formula 3,2. Median clypeal apron
concave. Body of clypeus projecting slightly over
edge of apron. Posterior outline of head very
weakly concave or flat. Nuchal groove indistinct
in lateral view. Mesosoma dorsal profile nearly
flat. Propodeal spines short with wide base; a
bisecting line passes just above anteroventral cor-
ner of pronotum. Spiracle about 1 diameter from
nearest edge of infradental lamella. Petiolar node
small. Postpetiole widest in anterior half (Fig. 66).
Ventral profile of sternum concave; anterior lip
not prominent.
Head rugose-areolate to areolate on
laterodorsa and sides, transversely areolate-rug-
ose on posterior head. Macrosculpture well de-
fined on posterior head, weaker on dorsum and
sides. Areolate microsculpture gives a granular
appearance to dorsum and sides of head, where it
largely obscures the weak macrosculpture; inter-
vals on posterior head nearly smooth. Anterior
edge of pronotal disc transversely rugose-areolate.
Rest of promesonotal dorsum longitudinally rug-
ose with occasional laterals; microsculpture ob-
scure. Mesosoma sides with confused areolate
macro- and microsculpture; the latter quite strong,
giving a granular appearance. Petiole appears
granular, with vestigial overlying macrosculpture.
Volume 3, 1 994
59
Postpetiolar node smooth; sides and venter ap-
pear granular.
Mesosoma dorsum with more than 12 pairs of
emergent erect hairs; nodes each with at least 2
pairs of long posterodorsally projecting hairs.
Tibiae with erect hairs.
Body yellowish-brown to brownish-yellow;
frontoclypeal area and appendages lighter, more
yellowish.
Nontype Workers.— TL 2.2-2.5, HL 0.54-0.60,
HW 0.43-0.50, SL 0.34-0.40, EL 0.04-0.08 (5-9 fac-
ets), PW 0.33-0.35, WL 0.54-0.66, SpL 0.03 (no
spines)-0.11, PetL 0.21-0.26, PpetL 0.12-0.16mm,
CI 0.80-0.83, OI 0.09-0.12, SI 0.77-0.83, PSI 0.05 (no
spines)-0.18, MHI 0.78-0.90. N=8
Type description extended as follows. Man-
dibles subtriangular in Dominican Republic speci-
mens. Clypeal apron convex in the Chiriqui,
Panama specimens and truncate in Chiapas and
Dominican Republic. Body of clypeus not project-
ing beyond clypeal apron in Chiapas specimen.
Basal tooth may equal penultimate basal in size.
Scapes slightly longer (SI 0.77-0.83). The Ocosingo,
Mexico specimen has slightly longer spines than
types (PSI 0.18); Cerro Pico Blanco, Costa Rica
specimens have very short spines (PSI 0.07-0.10)
those from Panama are devoid of armature (Fig.
67). Short propodeal spines of Costa Rican speci-
mens are more inclined than in types; a bisecting
line passes just below the axilla. Postpetiole of
Chiapas and Pedernales, Dominican Republic
specimens narrower than others (PpetW/ PpetL
1.23-1.28 vs. 1.41-1.56); Pedernales postpetioles
are widest in posterior half. Microsculpture on
head may be less distinct than in types. Panama-
nian specimens lack erect hair on extensor sur-
faces of tibiae and have only 9 pairs of erect hairs
on the mesosoma dorsum.
Possible Queens.— TL 2.6-2.8, HL 0.60-0.61, HW
0.52-0.53, SL 0.40, EL 0.12-0.13, PW 0.45, WL 0.73-
0.78, SpL 0.14-0.17, PetL 0.28-0.30, PpetL 0.14-
0.16mm, CI 0.86-0.88, OI 0.20-0.23, SI 0.75-0.77, PSI
0.19-0.22, MHI 0.92-0.97. N=2
Both queens collected on the north coast of
Colombia. Though workers of leptonana are un-
known from South America, these queens (Fig. 68)
have strongest affinities to leptonana workers:
Mandible with 5 teeth; basal as large as or larger
than penultimate basal. Clypeal apron emargin-
ate. Posterior outline of head flat. Propodeal spines
wide. Propodeal spiracle 2 diameters from edge of
infradental lamella. Mesosoma low. Petiolar pe-
duncle with prominent lamellate keel; node short.
Pygidium and sting apparatus as in workers.
Sculpture also as in workers. Microsculpture
present but indistinct on head and mesosoma.
Pronotum with 1-2 transverse rugae medially;
laterally rugose. Meso- and metanota longitudi-
nally rugose, with nearly smooth, shiny intervals.
Meso- and metapleura longitudinally rugose, ex-
cept for smooth, shiny mesokatepisterna. Dorsal
face of propodeum smooth. Petiole, including
peduncle, and postpetiole nearly smooth.
There are differences, however. Palpal for-
mula 2,2 in the dissected Guajiran queen (vs. 3,2 in
workers). One queen has a nearly subrectangular
postpetiole. Neither has erect hair on the tibiae.
Males.— TL2.9,HL0.51-0.53,HW0.55-0.56,SL
0.32-0.34, EL 0.25, PW 0.50-0.53, WL 0.91-0.95,
PetL 0.32-0.33, PpetL0.15-0.16mm, CI 1.04-1. 10, SI
0.58-0.61. N=4.
All four specimens from a nontype nest series,
Cerro Pico Blanco, near San Jose, Costa Rica. Man-
dible with 4 teeth. Clypeal margin convex. Anten-
nal flagellomeres 2-11 subequal in length and
width; not twisted. Lateral habitus shown in Fig.
69. May rian and parapsidal sutures present. Forew-
ing venation as in Fig. 30 hind wing as in Fig. 37.
Genitalia shown in Fig. 70. Mandibles smooth;
clypeus nearly so. Head dorsum areolate; sides
behind eyes rugose; posterior head rugose-areolate.
Mesosoma longitudinally rugose on lateral prono-
tum and dorsal meso- and metapleura. Anterior
and ventral mesopleura smooth. Ventral
metapleura diagonally rugose. Anterior portion
of mesonotum with vague effaced microsculpture;
longitudinally rugose macrosculpture begins at
level of wing and continues onto mesoscutellum.
Propodeum areolate. Petiole microareolate, with a
few weak, fine longitudinal rugae. Postpetiole and
gaster smooth. Abundant erect and decumbent
pilosity on scapes, head, mesosoma, waist, and
gaster. Head brown, except for yellowish-brown
frontoclypeal area and yellow mandibles. Rest of
body and appendages brownish-yellow; gaster Tl
and SI darker.
Discussion. — The nontype specimens from the
high elevations of Costa Rica may belong to a
distinct species defined by the complete absence
of propodeal spines, but presently they seem linked
to the types by the two specimens from Panama,
which have intermediate spine length.
60
Journal of Hymenoptera Research
The different palpal formula of the workers
and queens and the fact that workers and queens
specimens come from different countries cause
doubt about whether the queens are correctly
assigned. But most morphological characters fit
leptonana better than any other species. See also the
curvipubens discussion.
The range imwtabilis (Figs. 55-57, 97-98) over-
laps that of leptonana at least in southern Mexico
and Nicaragua, perhaps more, if queens are cor-
rectly assigned. Workers of innotabilis differ as
follows: 1) no erect hair on scapes, 2) generally
larger (WL 0.66-0.73mm), 3) clypeus evenly con-
vex, 4) palpal formula 2,2, 6) postpetiolar node
widest midlength and sternum moderately to
strongly prominent.
See also creightoni, curvipubens, alzatei, inermis,
and neilyensis discussions.
The name from leptos (G., slender) and nanus
(L., dwarf) describes the habitus of this species.
Distribution. — Specimens range from Chiapas,
Mexico to the northern coast of Colombia; from
sea level to 1750m. Type specimens come from
lowland rain forest, Colombian specimens from
forest receiving about 2m strongly seasonal an-
nual precipitation. Montane specimens from Costa
Rica were taken under large rocks in a pasture;
those from Panama were collected in debris under
a stump in unknown habitat.
Material Examined. — Holotype and paratype
locality. PANAMA: Canal Zone, Barro Colorado
Island; 2 workers including holotype, H-III-1943,
#5059 (J. Zetek) [USNM, lot 43 3035]; 5 workers,
VI-X-1943, #5105 (J. Zetek) [1 whole specimen
slide mounted] [USNM, lot 43-16534]; 4 workers,
14-11-1976 (A. Newton) [MCZ]; 1 worker, 27-11-
1976 (A. Newton) [MCZ]; 1 worker, 1-1960, #B-9
(W. L Brown & E. S. McCluskey) [mouthparts,
sting] [MCZ]; 2 workers, 7-III-1975, FP#6 (C. Toft
& S. Levings) [LACM].
Nontype localities. MEXICO: 1 worker,
Chiapas, Ocosingo Valley, Finca El Real, 1-7- VII-
1950 (Goodnight & Stannard). COSTA RICA: 6
workers, 4 males, Cerro Pico Blanco, near San Jose,
1750m, 15-III-1987(W. Eberhard & W. L. Brown) [2
male genitalia] [MCZ], PANAMA: 2 workers,
Chiriqui Province, near Nueva California, W of
Finca Palo Santo, 1550m, 5-III-1959, #B-377 (H. S.
Dybas) [MCZ]. COLOMBIA: 1 queen, Guajira,
near Don Diego, Quebrada Guacoche, 10m, 22-
VII-1976 ( W. L. Brown & C. Kugler) [queen mouth-
parts, sting] [MCZ]; 1 queen, Magdalena Depart-
ment, Parque Nacional Tayrona, near Pueblito,
210m, ll-XI-1976 (C. Kugler) [MCZ]. DOMINI-
CAN REPUBLIC: 3 workers 16km ENE Pedernales,
800m, 9-IX-1992 (P. S. Ward); 1 worker 4km NNW
Villa Altagracia, 200m, 12-IX-1992 (P. S. Ward)
[MCZ].
Rogeria belti Mann
Figs. 35-39, 91-92
Rogeria belti Mann 1922:31. Syntype workers, HONDURAS:
Progreso (Mann) [USNM] [2 of 12 syntypes examined].
Diagnosis. — WL 0.61-0. 94mm. Clypeal apron
usually with median notch, sometimes convex.
Eyes large, oval. Propodeal spines moderately
long, inclined (bisecting line usually passes well
below anteroventral corner of pronotum).
Promesonotal dorsum usually strongly areolate;
postpetiolar node smooth. Scapes with erect hair.
Hair on mesosoma dorsum decumbent to erect;
not clearly sorting into two kinds. Head dorsum
and gaster Tl with short decumbent and long
erect/suberect hair. No erect hair on extensor
surfaces of tibiae.
Workers.— TL 2.4-3.4, HL 0.56-0.80, HW 0.48-
0.69, SL 0.39-0.55, EL 0.10-0.14 (20-36 facets), PW
0.34-0.51, WL 0.61-0.94, SpL 0.11-0.20, PetL 0.25-
0.39, PpetL 0.13-0.21mm, CI 0.82-0.89, OI 0.18-
0.25, SI 0.76-0.89, PSI 0.14-0.24. N=31
Mandibles with 5-6 teeth; typically 5 teeth
evenly decreasing in size basad followed by a
large basal tooth, or 4 teeth are followed by 1 or 2
denticles and a large basal tooth (Fig. 35). A few
specimens in scattered localities have an addi-
tional denticle between the third and fourth teeth.
Palpal formula 3,2. Clypeal apron medially emar-
ginate in most, but evenly convex in specimens
from Miami, Costa Rica. Body of clypeus not pro-
jecting over clypeal apron. Posterior outline of
head weakly convex, sometimes with a slight
median concavity. Eyes sometimes narrower than
shown in Fig. 35; widest in Miami, Costa Rica
specimens (Fig. 36). Pronotum from the side usu-
ally subangular at junction of anterior and dorsal
faces; from above, shoulders well rounded. Met-
anotal groove either hardly visible, or a distinct
narrow groove, or a small step in mesosoma pro-
file; usually marked behind by a transverse ridge
(Fig. 35). Propodeal spiracles small, slightly more
than 1 diameter from posterior edge of propodeum,
facing slightly caudad. Metapleural lobes nar-
Volume 3, 1 994
61
rowly to broadly rounded. Petiolar keel not lamel-
late. Petiolar node usually dome-shaped (Fig. 35),
but sometimes slightly flattened on top.
Postpetiolar node highest in posterior half; dorsal
view subtrapezoidal as in Fig. 49; sternum long
and flat. Sting apparatus as in creightoni-group
diagnosis.
Longitudinal rugae on frontal lobes become
rugose-areolate by midlength of head; middle pair
of rugae usually remain free of cross-ridges until
posterior head. Cheeks and laterodorsa areolate to
rugose-areolate. Sides of head strongly areolate in
most, but nearly smooth in Miami, Costa Rica
specimens. Posterior head areolate with no clear
transverse pattern in most, but transversely arch-
ing rugose to rugose-areolate in ants from Miami,
Costa Rica and Colombia. Promesonotum gener-
ally coarsely areolate to areolate-rugose (Figs. 91-
92), but lower in relief and more rugose in speci-
mens from Miami, Costa Rica and some from
Colombia. Meso- and metapleura confused ar-
eolate to rugose. Dorsal face of propodeum pre-
dominantly transversely rugose, except for
Boquete, Panama specimens, in which dorsal face
is indistinctly areolate. Petiolar node not as heavily
sculptured as mesosoma, but distinctly areolate to
weakly rugose-areolate; dorsal and anterior faces
often free of macrosculpture. Intervals irregular
and usually shiny. Microsculprure generally weak
and indistinct, producing quite shiny intervals in
macrosculpture, especially on sides of head,
mesosoma, and on petiolar node. Head dorsum
and petiolar node of Miami, Costa Rica specimens
with more pronounced microsculprure, lending a
granular appearance.
Most dark reddish-brown with yellowish-
brown mandibles, antennae and legs. Specimens
from Boquete, Panama and Miami, Costa Rica
lighter, with more yellowish appendages and end
of gaster. Mexican specimens reddish-black with
brown appendages.
Queens.— TL 3.0-3.4, HL 0.68-0.74, HW 0.58-
0.64, SL 0.48-0.52, EL 0.18-0.20, PW 0.48-0.59, WL
0.88-1.00, SpL0.13-0.21,PetL0.29-0.37,PpetL0.18-
0.20mm, CI 0.84-0.85, SI 0.77-0.86, PSI 0.14-0.21.
N=5
Like the worker except for normal queen struc-
tures and the presence of predominantly rugose
sculpture on the mesonotum. Parapsidal furrows
indistinguishable from furrows in sculpture.
Habitus similar to that of merenbergiana queen
(Fig. 48), but mesonotum slightly more convex
and propodeal spines longer and narrower. Wings
as in male (Fig. 37).
Males.— TL 3.4-3.5, HL 0.59-0.61, HW 0.65-
0.70, SL 0.32-0.35, EL 0.32, PW 0.71-0.72, WL 1.12-
1.19, PetL 0.40-0.41, PpetL 0.19-0.20mm, CI 1.11-
1.15, SI 0.50. N=2
All five specimens from La Ceiba, Honduras.
Mandibles with 5 teeth. Clypeus weakly emargin-
ate. Flagellomeres 2-11 straight, subequal in length
and width. Wings, habitus and genitalia as shown
in Figs. 37-39. Rs vein curves away from leading
edge of wing; r-m vein lost; M vein short. Man-
dibles smooth except for piligerous punctures.
Face very finely areolate, opaque. Posterior head
longitudinally rugose. Pronotum weakly punc-
tate. Mesonotum and posterior face of propodeum
finely rugulose and densely punctured.
Mesopleura longitudinally striate right under
wing, otherwise smooth and shiny with piligerous
punctures. Metapleura and sides of propodeum
longitudinally rugose. Posterior face of propodeum
rugose-areolate. Petiolar node finely punctate to
smooth. Postpetiole and gaster smooth and shiny.
Hair as in worker, but more dense. Color brown-
ish-black, with brown legs and antennae and
brownish-yellow mandibles.
Discussion. — The 18 specimens from Miami,
Costa Rica (Fig. 36) differ from typical belti in
having: 1) an evenly convex clypeus, 2) a larger
eye (31-36 facets vs. 20-30 in typical belti), 3) smaller
propodeal spines (PSI 0.15-0.18 vs. 0.19-0.24 in
typical belti), 4) generally narrower head, 5) head
and mesosoma more rugose than areolate, and 6)
weaker macrosculpture in general, but
microsculprure stronger on head and petiolar node.
Queens from Miami, Costa Rica like their workers
have narrower heads, shorter propodeal spines
and weaker sculpture than the four from Guate-
mala, Panama and the Osa Peninsula of Costa
Rica. I considered these Miami ants a new species
until the arrival of new specimens from Costa Rica
and Colombia. One worker from Heredia, Costa
Rica has a convex clypeus like the Miami ants, but
eye and propodeal spine size just within the typi-
cal belti range. Other workers from Colombia have
eyes as large as any of the Miami specimens, but an
emarginate clypeus like typical belti, and they
have intermediate sculpture. While it is still pos-
sible that the Miami population represents a dif-
ferent species, I am not confident of that at present.
62
Journal of Hymenoptera Research
Some alzatei (Figs. 58-60, 99) specimens
strongly resemble specimens of belti from Costa
Rica in size, habitus, spine length, and sculpture,
but alzatei: 1) is smaller (WL 0.52-0.65mm), 2) has
a truncate clypeal apron in Central America, 3) has
smaller eyes (11-14 facets), and 4) lacks erect hair
on the scapes.
The geographic ranges of creightoni (Figs. 51-
54, 95-96) and belti overlap in much of Central
America and some members look quite similar in
size, sculpture, and pilosity. Workers of creightoni
can be distinguished by: 1) generally smaller, el-
liptical eyes with longer propodeal spines (< 19
facets; EL/SpL 0.29-0.64 vs. 0.62-1.23 in belti), 2)
more horizontal propodeal spines, 3) promesonotal
dorsum rugose to rugose-areolate, and 4) hair on
mesosoma dorsum clearly sorting into two dis-
tinct kinds.
See discussions of inermis, tonduzi,
merenbergiana, and exsulans for comparisons with
other similar species.
Distribution. — Rogeriabelti is so far known from
southern Mexico to the Canal Zone and southwest
Colombia. Elevations range from 200m to 1500m
in Mexico and 50m to 950m in Panama. Labels
with ecological data show them collected in rain
forest "in Philodendron," "frond sheaths on
ground," leaf litter, around an epiphyte mat on a
fallen branch, rotten wood, and in dead trees or
branches on the ground. Jack Longino found a
colony "nesting under loose bark of a still solid log
fallen across trail." On another rotten log he lured
workers from under the bark to a freshly killed
tabanid (Longino, unpublished field notes).
Material Examined. — MEXICO: Vera Cruz
State, Los Tuxtlas (P. S. Ward); Chiapas State, El
Bosque (A. Newton). GUATEMALA: Bobas (W.
M. Mann). BELIZE [=British Honduras]: Caves
Branch (S. and J. Peck). HONDURAS: Progreso [or
El Progreso] (W. M. Mann), Lancetilla and La
Ceiba (W. L. Brown) [mouthparts, sting; 2 male
genitalia]. EL SALVADOR: between Apopa and
Nejapa (W. L. Brown). COSTA RICA: Miami (un-
known collector) [mouthparts, sting] [USNM];
Heredia Province, (J. Longino); Santa Clara
[ = Limon?] Province, Hamburg Farm and
Parismina Br. (F. Nevermann); Puntarenas Prov-
ince, Manuel Antonio National Park and Reserva
Biologica Carara (P. S. Ward), Osa Peninsula,
Corcovado National Park (J. Longino; P. S. Ward).
PANAMA: Chiriqui Mountains, Boquete (F. M.
Gaige); Cerro Campana (A. Newton); El Valle (H.
S. Dybas); Canal Zone, Barro Colorado Island (J.
Zetek; A. Newton). COLOMBIA: Nariho Depart-
ment, La Guayacana (P. Silverstone) [LACM]. 117
workers, 15 queens, 5 males [CUIC, JTLC, LACM,
MCZ, MZSP, USNM].
Rogeria inermis Mann
Figs. 40-42
Rogcrm inermis Mann 1922:32, Fig. 15. Syntype workers,
HONDURAS: Progreso, Lombardia (Mann) [MCZ] [3of
5 syntypes examined].
Diagtiosis. — WL 0.60-0.85mm. Clypeal apron
medially emarginate. Eye oval, moderately large.
Nuchal grooves weakly visible in lateral view.
Propodeal spines short to absent. Propodeal spi-
racles rather large, prominent, face somewhat
caudad, usually < 1/2 diameter from edge of
infradental lamella. Promesonotal dorsum rugose
to areolate-rugose; pronotal sides rugose, shiny.
Scapes with erect hair. Head dorsum with erect
hairs and shorter, decumbent pilosity. Hair on
mesosoma and gaster Tl subdecumbent to erect;
not two distinct kinds. Pilosity on gaster rather
sparse. Extensor surfaces of tibiae with decum-
bent to suberect hairs.
Workers.— TL 2.2-3.0, HL 0.55-0.71, HW 0.48-
0.63, SL 0.38-0.50, EL 0.09-0.13 (15-28 facets), PW
0.34-0.47, WL 0.60-0.85, SpL 0.03(no spines)-0.10,
PetL 0.25-0.37, PpetL 0.14-0.20mm, CI 0.82-0.91,
OI 0.18-0.22, SI 0.73-0.88, PSI 0.04(no spines)-0.14.
N=15
Mandibles with 5 or 6 teeth, usually five de-
creasing in size basad then a large basal tooth.
Palpal formula 3,2. Median clypeal apron weakly
to strongly emarginate. Posterior outline of head
convex. Metanotal groove weak to absent. Ante-
rior edge of propodeum sometimes marked by a
transverse ridge. Propodeal spines usually short
and acute (Fig. 40), but may also be quite long (Fig.
41), blunt, or absent. Petiole (Fig. 40) with a dis-
tinct, rounded node; peduncle slender, venter with
strong tooth, but little or no keel. Postpetiolar
node highest in posterior half; shape in dorsal
view subtrapezoidal as in Fig. 49. Anterior lip of
postpetiolar sternum square, not protruding. Py-
gidium as in creightoni-gioup diagnosis.
Sting apparatus as in Fig. 42. Spiracular plates
with incomplete medial connection; spiracle not
enlarged. Anterodorsal corner of quadrate plate
Volume 3, 1994
63
prominent, with medial and lateral lobes. Oblong
plate anterior apodeme long, slender; ventral arm
tall, with with long, slender fulcral arm. Gonostylus
with dorsoterminal and companion setae; no gap
between these and other sensilla. Lancets strong,
acute, with a single minute apical barb. Sting shaft
slender, straight; apex strong, acute, with low
dorsal flange. Valve chamber and sting bulb large,
subequal in height and length. Sting base much
lower than bulb height, with weak anterolateral
processes. Furcula an inverted Y-shape. See Kugler
(1978b) for a more complete description.
Microsculpture on head and mesosoma re-
duced to vague irregularities in otherwise smooth
and shiny interrugal spaces. Middorsum of head
longitudinally rugose with lateral spurs, grading
to areolate-rugose on laterodorsa and sides. Sculp-
ture below and behind the eye sometimes vesti-
gial, leaving a shiny area. Posterior head in most
specimens transversely arching rugose (with lat-
eral spurs) to areolate-rugose, but longitudinally
rugose in two workers from the Osa Peninsula of
Costa Rica. Rugae on mesosoma are sharp, with
broad interrugal spaces that are even smoother
and shinier than on the head. Anterior pronotum
with 1-3 transverse rugae; disc with longitudinal
rugae; meso- and metanorum vermiculate-rugose
or areolate-rugose. Pronotal sides, mesopleura,
and metapleura predominantly rugose; intervals
very shiny. Dorsal face of propodeum transversely
rugose to areolate. Sides of petiolar node weakly
rugose to rugose-areolate, producing an uneven,
shiny surface; apex smooth. Postpetiolar node
largely smooth and shining with only slight un-
evenness of surface on sides.
Color varies from a reddish-brown (almost
black) body with yellowish brown appendages to
a yellowish-brown body with darker gaster and
brownish-yellow appendages.
Queens.— TL 2.9-3.0, HL 0.62-0.64, HW 0.55-
0.57, SL 0.41-0.45, EL 0.15-0.16, PW 0.47-0.48, WL
0.81-0.84, SpL 0.09, PetL 0.32-0.36, PpetL 0.18-
0.19mm, CI 0.89, SI 0.74-0.79, PSI 0.11. N=3
Mandibles with 6 teeth, or 5 teeth and 2 den-
bdes. Parapsidal furrows not discernible. Sculp-
ture on posterior head of Panamanian specimens
transversely rugose-areolate; more longitudinally
oriented in Costa Rican specimens. Pronotum
transversely rugose-areolate. Mesonotal sculpture
predominantly rugose, with some closed cells.
Dorsal face of propodeum transversely rugose.
Petiolar node with weak macrosculpture,
postpetiole smooth.
Discussion. — The two specimens from the Osa
Peninsula of Costa Rica differ from the other speci-
mens in having noticeably longer propodeal spines
and longitudinal, rather than transverse sculpture
on the back of the head. These may be members of
a sibling species, but spine length is quite variable
in inermis (e.g. PSI 0.04-0.09 in one Berlese sample).
Thus, I hesitate to name a new species until more
specimens show consistent differences.
Rogena leptonana (Figs. 66-70) from Cerro Pico
Blanco, Costa Rica and Nueva California, Panama
have reduced or absent propodeal spines, but
differ from inermis in: 1) generally smaller size
(WL 0.57-0.66mm; only 1 of 49 inermis workers
with WL < 0.67mm), 2) smaller eye (6-8 facets; OI
0.11-0.12), and 3) mesosoma and gaster Tl dorsa
with 2 distinct types of pilosity. Rogeria belti (Figs.
35-39, 91-92), often collected with inermis, has
longer propodeal spines and more areolate head
and pronotal sculpture. See also the neilyensis dis-
cussion.
Distribution. — All specimens are from Guate-
mala to Panama, at elevations from 10m to 1250m.
Some were collected in fruit bat guano; others, in
Berlese samples from rain forest leaf litter.
Material Examined. — GUATEMALA: Alta
Verapaz Department, Lanquin (S. and J. Peck).
HONDURAS: Progreso [or El Progreso];
Lombardia (W. M. Mann). COSTA RICA: (A. Bierig;
F. Nevermann); Heredia Province, Puerto Viejo,
La Selva Field Station (C. Kugler; Talbot &
VanDevender) [2 stings] ; Limon Province, Cahuita
(P. S. Ward); Puntarenas Province, Osa Peninsula
(S. Peck), Manuel Antonio National Park (P. S.
Ward), Carara Biological Reserve (P. S. Ward);
Cartago Province, Turrialba vicinity (A. Bierig)
(W. L. Brown) [mouthparts, sting]. PANAMA:
Bocas del Toro, 3 localities (D. M. Olson); Barro
Colorado Island (W. L. Brown and E. S. McCluskey
[mouthparts, whole specimen]; N. A. Weber; J.
Zetek); Gamboa (P. S. Ward); Panama City (W. L.
Brown). 49 workers, 3 queens [CKC, CUIC, L ACM,
MCZ, MZSP,USNM].
Rogeria neilyensis new species
Fig. 43
Diagnosis. — WL 0.75-0.76mm. Clypeal apron
convex, with weak median angle. Eye small.
64
Journal of Hymenoptera Research
Nuchal groove makes a deep notch in lateral view
of head. Propodeal spines short, wide. Propodeal
spiracles > 1 diameter from edge of infradental
lamella. Postpetiolar sternum not enlarged. Ter-
minal segments of gaster rotated ventrad. Sides of
head with distinct areolate-rugose to rugose
macrosculpture; mesosoma predominantly rug-
ose. Scapes and legs lack erect hairs. Head dorsum
with short, dense, decumbent pilosity and longer,
more sparse, erect-suberect hairs. Mesosoma hairs
range continuously from short, decumbent to long,
erect (i.e. not sorting into two distinct kinds as on
head.). Hair on gaster Tl dense, subdecumbent-
suberect (decumbent hairs few or absent).
Holotype and Paratope Workers. — TL 2.8-2.9
(2.8), HL 0.66-0.68 (0.67), HW 0.58-0.60 (0.59), SL
0.45, EL 0.06-0.07 (0.06) (9-13 facets), PW 0.42-0.43
(0.42), WL 0.75-0.76 (0.75), SpL 0.08-0.10 (0.08),
PetL 0.30-0.34 (0.30), PpetL 0.18-0.19 (0.18)mm, CI
0.88-0.89 (0.88), OI 0.10-0.12 (0.10), SI 0.76-0.77
(0.76), PSI 0.11-0.13 (0.11), MHI 0.96-1.00 (0.96).
N=2
Mandible 6-toothed; basal tooth larger than
penultimate tooth. On left mandible, fifth tooth
replaced by 2 denticles. Body of clypeus slightly
prominent. Frontal lobes rather wide (FLW/HW
0.43-0.44). Anteroventral corner of pronotum pro-
jecting tooth-like. No distinct metanotal groove.
Propodeal spiracles moderately large, weakly
bulging from sides. Metapleural lobes low, broadly
rounded. Petiolar peduncle with large ventral keel
and tooth. Postpetiolar node lacking a distinct
posterior face; subtrapezoidal from above, as in
Fig. 50.
Longitudinal rugae on frontal lobes become
areolate-rugose halfway tobackof head. Posterior
head transversely areolate-rugose. Laterodorsa
areolate-rugose. Intervals in macrosculpture of
head shiny, but wrinkled by vague, irregular
microsculpture. Anterior edge of pronotal disc
areolate; rest of promesonotal dorsum longitudi-
nally rugose-areolate; more rugose mesad, more
areolate laterad. Dorsal face of propodeum trans-
versely rugose. Intervals on mesosoma generally
wider and smoother than on head. Petiolar node
vaguely areolate; posterior face and peduncle with
longitudinal carinae. Surface of postpetiolar node
uneven, but predominantly smooth and shiny.
Gaster Tl and SI smooth and shiny, but other
segments progressively dulled caudad by dense
microareolate sculpture.
Color yellowish-brown, with slightly more
yellowish appendages and anterior of head; first
tergum of gaster dark brown.
Discussion. — This species is similar in size and
reduced petiolar spines to another Central Ameri-
can species, biennis (Figs. 40-42), which differs as
follows: 1) clypeal apron medially emarginate, 2)
nuchal groove weak, 3) eyes larger (OI 0.18-0.22;
15-28 facets), 4) propodeal spiracle < 1 diameter
from edge of propodeum, 5) scapes with erect hair,
and 6) mesosoma and gaster Tl with little, if any,
decumbent hair.
The leptonana (Figs. 66-70) specimens from
Cerro Pico Blanco, Costa Rica, have low, wide
propodeal spines like neilyensis, but leptonana has:
1 ) an emarginate clypeus, 2) weak nuchal grooves,
3) lower lower mesosoma profile (MHI 0.84-0.90),
and 4) scapes with erect hair.
The name neilyensis refers to the type locality.
Material examined. — Holotype locality. COSTA
RICA: Puntarenas Province, 3km N Ciudad Neily
[=Villa Neily], 8.41N 82.57W, 210m, 31-VII-1985,
#7771-5 (P. S. Ward). From sifted litter (leaf mold
and rotten wood) in second-growth rain forest
[MCZ].
Paratype locality. 2 workers, holotype locality
[LACM, MCZ].
Rogeria exsulans Wilson and Taylor
Fig. 44
Rogena exsulans Wilson and Taylor 1967:74, Fig. 60. Holotype
worker, SAMOA: Upolu (T. E. Woodward) [Holotype
MCZ; paratypes ANIC, MCZ] [Holotype and 30
paratypes examined].
Diagnosis. — WL 0.68-0. 84mm. Eye fairly large,
oval. Propodeal spiracle faces posterolaterally.
Petiole with lamellate keel and dentate inferior
process. Terminal segments of gaster rotated
ventrad. Sides of head areolate-rugose; posterior
head and promesonotum strongly areolate. Scapes
with decumbent and a few short, suberect hairs.
No erect hair on extensor surfaces of legs. Mostly
abundant decumbent to subdecumbent hair on
dorsa of head, mesosoma, nodes and gaster Tl
(sparse erect hair on head and mesosoma); not
sorting into two distinct kinds of pilosity.
Workers.— TL 2.5-3.0, HL 0.62-0.71, HW 0.52-
0.64, EL 0.08-0.11 (20-26 facets), PW 0.37-0.45, WL
0.68-0.84,SpL0.10-0.15,PetL0.26-0.33,PpetL0.15-
0.17mm, CI 0.84-0.90, OI 0.19-0.21, SI 0.81-0.85, PSI
Volume 3. 1 994
65
0.14-0.19. N=4
Some mandibles with 6 teeth (5 decreasing in
size then a large basal), but usually the small
penultimate basal tooth is replaced by 2 denticles,
or by a gap and 1 denticle. Palpal formula 3,2.
Clypeal apron medially emarginate. Back of head
in dorsal view convex or flat medially; temples
broadly rounded (Wilson and Taylor's Fig. 60).
Promesonotal profile may be evenly rounded, but
more often angular (Fig. 44). Metanotal groove
usually distinct, producing a step between
promesonotum and propodeum, but weak in two
specimens. Propodeal spiracles more than one
diameter from nearest edge of inf radental lamella.
Node short, evenly convex. Postpetiolar sternum
relatively long, flat; postpetiole from above
subtrapezoidal, as in Fig. 51. Sting apparatus as in
creightoni-group diagnosis.
Longitudinal rugae of frontal lobes and me-
dian head gradually change to strongly areolate
on posterior head. Cheeks and laterodorsa longi-
tudinally areolate-rugose. Intervals on head dor-
sum vaguely undulate, but shiny; intervals on
sides and posterior head smooth and shiny.
Promesonotum for the most part strongly areolate,
but several elongate cells may occupy the midline
of the pronotal disc, and often the pronotal sides
have weaker areolate sculpture. Meso- and
metapleura confused areolate-rugose. Mesosoma
microsculpture reduced; intervals in macrosculp-
ture uneven, but very shiny. Petiolar node with
weak to vestigial areolate macrosculpture and
vague microareolate background. Postpetiolar
node shiny, nearly smooth throughout.
Head, mesosoma dark reddish-brown, waist
and anterior and posterior ends of gaster some-
what lighter. Mandibles, clypeus, antennae, and
legs yellowish-brown.
Discussion. — The only other Pacific species,
stigmatica and megastigmatica are easily distin-
guished from exsulans by their lack of a petiolar
keel and inferior process, as well as other features
listed in the diagnoses.
The species most similar to exsulans is Central
American. Rogeria belti has some members with
similar size, habitus, eye size, clypeus shape, a
distinct petiolar keel, strong areolate sculpture,
and intergrading types of hairs on mesosoma.
However, belti workers still differ in having gener-
ally longer propodeal spines (PSI 0.19-0.24), less
prominent petiolar keel, and distinct long/erect
and short /decumbent pilosities on the gaster.
Distribution. — Rogeria exsulans has been col-
lected only at 600-700m on a single island in the
middle of the South Pacific. Most collections specify
a rain forest habitat, where it has been found under
dead bark, in rotten logs, in moss and ferns on
trees and in berlesate of moss on logs and tree
trunks.
Material Examined.— WESTERN SAMOA:
Upolu, Afiamalu (T. E. Woodward; R. W. Taylor;
E. C. Zimmerman) [2 mouthparts, stings]. 31 work-
ers [MCZ],
Rogeria cornuta new species
Fig. 45
Diagnosis. — WL 0.93-1. 02mm. Eye relatively
small. Nuchal groove makes strong notch in lat-
eral view of head. Propodeal spines very long
(EL/SpL < 0.50); not inclined dorsad; distal por-
tions subparallel with midline. Propodeal spiracles
prominent, less than 1/2 diameter from edge of
infradental lamella. Metapleural lobes prominent.
Petiole with little or no keel. Sides and posterior
head strongly areolate; mesosoma predominantly
rugose. Erect hair on scapes. Gaster Tl lacks de-
cumbent hair.
Holotifpe and Paratype Workers. — TL (3.5)-3.8,
HL (0.80)-0.89, HW (0>3)-0.78, SL (0.55)-0.58, EL
0.10 (16-18 facets), PW (0.55)-0.60, WL (0.93)-1.02,
SpL (0.25)-0.27, PetL (0.42)-0.45, PpetL (0.21)-
0.24mm, CI 0.88-(0.92), OI 0.13-(0. 14), SI 0.74-(0.75),
PSI 0.26-(0.27), MHI (0.98)-1.02.
Mandible with 6 teeth; basal larger than
penultimate basal. Clypeal apron medially emar-
ginate; body of clypeus projecting slightly over
apron. Posterior outline of head broadly and
weakly concave. Sides of pronotum with anterior
grooves for insertion of corners of head; shoulders
from above angular. No meso- or metanotal
grooves. Paratype promesonotal dorsum less con-
vex and more angular in front and back than
shown for holotype (Fig. 45). Propodeal directed
caudad so that a bisecting line would extend just
below shoulder; distal half of spines curve inward,
almost paralleling the midline. Postpetiolar node
somewhat flattened on top; subtrapezoidal in dor-
sal view. Postpetiolar node of paratype lower in
front than behind. Postpetiolar sternum long, not
projecting anteriorly.
Head macrosculpture coarse (especially be-
66
Journal of Hymenoptera Research
hind), with sharp ridges and shiny interstices.
Longitudinal rugae on front break up at midlength
of head and give way to a transversely arching
areolate-rugose pattern on the posterior head.
Laterodorsa confused rugose-areolate. Mesosoma
macrosculpture also coarse with smooth inter-
stices, but ridges are rounded. Anterior face of
pronotum transversely rugose-areolate, pronotal
disc longitudinally rugose to vermiculate-rugose
(holotype). Mesonotum vermiculate-rugose with
some cross-ridges. Pronotal sides areolate-rugose
(holotype) or broken and confused; rest of sides
confusedly longitudinally rugose with few con-
necting ridges. Anterior edge of propodeum
marked by a sharp transverse ridge. Petiolar node
weakly areolate on sides and posterior; smooth
along anterior and dorsal midline. Postpetiolar
node weakly areolate on sides; smooth along mid-
line.
Scapes, head dorsum and tibiae with short
decumbent and long erect-suberect hair. Hair on
mesosoma and waist ranges from decumbent to
erect and varies in length, but not clearly segre-
gated into two distinct types. Hair on gaster erect-
suberect.
Color dark reddish-brown, with lighter
frontoclypeal area and ends of gaster; legs and
antennae yellowish-brown.
Discussion. — Some creightoni, also from Belize,
have the same habitus as cornuta, including long,
horizontal propodeal spines, but these creightoni
members are much smaller (WL 0.63-0. 71mm)
and have abundant decumbent pilosity on the
gaster Tl. Other creightoni from La Selva, Costa
Rica are the same size as the cornuta holotype, but
have shorter propodeal spines, slightly smaller
eyes, and abundant decumbent pilosity on the
gaster Tl.
The name cornuta means horned, referring to
the long, horn-like propodeal spines.
Material Examined. — Holotype locality.
BELIZE (British Honduras): 2.5 mi. S Belmopan, 4-
VIII-1972, S. and J. Peck, #242 [MZSP].
Paratype locality. 1 worker, MEXICO: Chiapas
State, Ocosingo, 2-VI-1967, J. M. Campbell [MCZ] .
Curvipubens-Group and Related Species
Rogeria curvipubens Emery
Figs. 74-76, 101-102
Rogeria curvipubens Emery 1894:190. Worker and queen
syntypes, U. S. VIRGIN ISLANDS: St. Thomas (Eggers)
[MCSN] [Worker syntype examined].
Additions to curvipubens-group diagnosis.
Postpetiolar node strongly vaulted and with small
posterior peduncle. Anterior edge of postpetiolar
sternum not strongly produced; junction of poste-
rior and ventral edges angular (Fig. 75). Sting shaft
and lancets weak, spatulate. Sides of head smooth
and shiny. Promesonotum with vestigial
microsculprure, making interrugal spaces nearly
smooth and weakly to strongly shiny. Dorsal face
of propodeum usually with 1-5 transverse rugulae.
Workers.— TL 1.9-2.3, HL 0.48-0.58, HW 0.38-
0.49, SL 0.32-0.39, EL 0.04-0.07 (4-11 facets), PW
0.28-0.38, WL 0.50-0.63, SpL 0.05-0.11, PetL 0.20-
0.24, PpetL 0.10-0.13mm, CI 0.80-0.86, OI 0.10-
0.15, SI 0.77-0.81, PSI 0.13-0.18, MHI 0.84-1.05.
N=22
Mandibles typically with 6 teeth, but occa-
sionally with an additional denticle or tooth; basal
tooth little if any larger than penultimate basal.
Haitian specimens with 5 teeth. Anterior edge of
clypeus evenly convex or with median concavity
(Haiti). Body of clypeus, though not especially
prominent, is generally the anteriormost point of
the head in full dorsal view; little or no shelf-like
apron. Posterior outline of head generally broadly
convex, but sometimes with a weak median con-
cavity (Fig. 74). Nuchal groove weak. Eye small,
elliptical. Pronotum weakly angular or rounded
in front and on sides. Metanotal groove absent or
suggested by a broad, very shallow impression.
Anterior propodeum marked by transverse
carinula that often does not interrupt mesosoma
profile. Propodeal spines short, wide, with curved
(Fig. 74) or straight edges; a bisecting line passes
below the anteroventral corner of pronotum.
Metapleural lobes low, subangular to rounded.
Sting apparatus like that of inermis (Fig. 42) in most
respects, but lancets and sting shaft are weak and
the lancets are blunt and spatulate. Though twisted
in all preparations, the sting shaft does seem to
have a dorsoterminal flange.
Posterior head transversely arching rugose to
rugose-areolate, intervals shiny (Fig. 101). Most of
Volume 3. 1 994
67
head with vague microsculpture. Anterior edge of
pronotal disc areolate to rugose-areolate; disc lon-
gitudinally rugose (Fig. 102), sometimes effaced
on meso- and metanota. Pronotal sides with one or
more weak longitudinal rugae; meso- and
metapleura confused rugose to rugose-areolate.
Petiole and postpetiole microareolate and devoid
of macrosculpture, except for small carinulae on
petiolar peduncle of some specimens.
Microsculpture weaker on apices of nodes, espe-
cially postpetiolar node, which is shiny and nearly
smooth.
Head with 0-6 short suberect hairs; mesosoma
dorsum with 2-7 pairs. Extent of erect pilosity on
gaster Tl variable; usually covering whole sur-
face, but may cover as little as the posterior third.
Color uniformly yellow, to golden or light
brown body with lighter appendages,
frontoclypeal region and ventral gaster.
Queens.— TL 2.3-2.6, HL 0.51-0.59, HW 0.44-
0.51, SL 0.34-0.39, EL 0.09-0.12, PW 0.38-0.45, WL
0.63-0.71, SpL 0. 1 1-0.14, PetL 0.22-0.27, PpetL 0.12-
0.15mm, CI 0.83-0.90, SI 0.73-0.78, PSI 0.16-0.19,
MHI 0.96-1.17. N=6
Queens (Fig. 76) vary like workers in shapes of
head, propodeal spines, petiole and postpetiole.
Mandibles have 6 teeth in most; plus 2 denticles in
the Guatemalan specimen. Parapsidal furrows
weak or indistinguishable. Wing venation as in
belti (Fig. 37). Sting apparatus of a queen from the
north coast of Colombia like that of workers from
the region, including spatulate lancets. Sculpture,
pilosity, and color also vary similarly. Sides of
head may be partly rugose, but some portion
smooth. Pro-, meso-, and metanota rugose. Meso-
and metapleura longitudinally rugose dorsad;
anterior half of mesokatepisterna very smooth
and shiny. Microsculpture as in workers. Num-
bers of erect hairs on various parts range as in
workers, except for one queen with 10 pairs on
mesosoma dorsum.
Queens from Mexico and Guatemala are not
associated with workers. Moreover, the Guatema-
lan specimen lacks a postpetiole and gaster. Both
have the side of the head rather strongly rugose-
areolate, but I am guessing that they are cunnpubens,
rather than cuneola on the basis of the strongly
shiny mesokatepisterna and shape of the
postpetiolar sternum of the Mexican specimen.
Discussion. — Workers come from the Carib-
bean Islands and northern South America. How-
ever, if the two queens from Mexico and Guate-
mala are in fact cunnpubens, the range of cunnpubens
broadly overlaps that of sister species cuneola. The
two species are distinguished by characters in the
cuneola diagnosis and description.
Rogeria alzatei (Figs. 58-60, 99) and leptonana
(Figs. 66-70) are very similar to curvipubens and
cuneola and have been considered curvipubens in
the past, but I believe they can be distinguished,
usually by general habitus, but especially by more
abundant erect pilosity than described in the
curvipubens-group diagnosis. Moreover, side-by-
side comparisons of sympatric specimens from
the Villavicencio vicinity of Colombia and from
Barro Colorado Island reveal more differences:
the alzatei specimens have a truncate clypeal apron,
narrower propodeal spines, and generally stron-
ger sculpture. The leptonana specimens have an
emarginate clypeal apron, lower mesosoma, and
larger petiolar keel. The only question of identity
arises on the north coast of Colombia, where a
single specimen has wider propodeal spines and
convex clypeus like curvipubens, which are com-
mon in the area, but has the abundant pilosity and
stronger macrosculpture of alzatei. I call that speci-
men alzatei, since spine and clypeal shapes vary in
that species.
Assigning stray queens has also been prob-
lematic. In two localities on the north coast of
Colombia (Pueblito, Tayrona Park; Don Diego,
Guajira) I collected three distinct kinds of queens.
One set has a distinctly lower mesosoma (MHI
0.92-0.97) and larger petiolar keel than the other
two and has abundant erect pilosity on scapes,
whole head, mesosoma, waist, and gaster. These
and other characters suggest those are leptonana
queens. The other two kinds of queens have a
more compact mesosoma (MHI 1 .05-1 .22), but one
set has no erect hair on scapes, head, waist, or first
tergum of gaster, little or no erect hair on the
mesosoma dorsum, more effaced sculpture with
nearly smooth sides of head and transversely arch-
ing rugae on posterior head, and spatulate lancets,
just like the curvipubens workers that are common
in both areas. The third set of queens have distinct
macrosculpture, distinctly areolate sides and pos-
terior head, short erect hair on the whole head
dorsum and more abundant erect /suberect hair
on mesosoma, waist and gaster and acute lancets.
I have tentatively assigned them to innotabilis be-
cause of their evenly convex clypeus and
68
Journal of Hymenoptera Research
posteroventral spine on the spiracular plate of the
sting apparatus.
See also micromma, and tribrocca discussions.
Distribution. — Rogeria curvipubens has been
collected in by Berlese sampling in tropical rain
forest, secondary growth forest, and dry tropical
forest. Collection sites have all been below 500m
elevation.
Material Examined.— VIRGIN ISLANDS: St.
Thomas (Eggers; Balzan); St. Croix (I. Proj. staff).
HAITI: Aquin (no collector). JAMAICA: St. Eliza-
beth Parish, Black River (no collector). MEXICO:
Vera Cruz, Cuatatolapan (M. Abarca). GUATE-
MALA: Alta Verapaz, Trece Aguas (Schwarz and
Barber). PANAMA: Barro Colorado Island (N. A.
Weber; W. L. Brown). COLOMBIA: Guajira,
Serrania de Macuira and vicinity Rio Don Diego
(W. L. Brown and C. Kugler) [1 mouthparts, 2
stings, 1 whole specimen slide mounted];
Magdalena Department, Tigrera near Santa Marta
(W. L. Brown and C. Kugler), Tayrona National
Park (C. Kugler) [worker and queen mouthparts,
sting]; Meta Department, Caho El Buque near
Villavicencio (Kugler). VENEZUELA: Bolivar
State, Campamento Rio Grande (P. S. Ward);
Monagas State, Parque de Laguna Grande (P. F.
Kukuk). SURINAM: Tambahredjo [queen sting],
Sidoredjo [sting], Dirkshoop, and La Poulle (I. van
der Drift). 57 workers, 13 queens [BMNH, CKC,
CUIC, LACM, MCZ, MZSP, USNM].
Rogeria cuneola new species
Figs. 77-78, 103
Additions to curi'ipubens-group diagnosis.
Postpetiolar node weakly vaulted and with no
posterior peduncle. Anterior edge of sternum
strongly produced; posterior and ventral edges
merge insensibly (Fig. 78). Sting shaft and lancets
strong, acute; sting shaft with dorsal flange; lancet
with barbule. Sides of head and mesosoma with
strong microareolate sculpture that obscures weak
macrosculpture and makes intervals opaque. Dor-
sal face of propodeum without transverse rugulae.
Holotype and Paratype Workers. — TL 2.0-2.1
(2.0), HL 0.50-0.54 (0.51), HW 0.43-0.46 (0.43), SL
0.32-0.35 (0.33), EL 0.05-0.06 (0.05) (6-7 facets), PW
0.31-0.33 (0.31), WL 0.51-0.56 (0.52), SpL 0.07-0.09
(0.08), PetL 0.20-0.22 (0.20), PpetL 0.10-0.12
(O.ll)mm, CI 0.84-0.87 (0.84), OI 0.10-0.13 (0.13), SI
0.73-0.78 (0.77), PSI 0.15-0.17, MHI 0.94-1.02 (1.02).
N=7
Nontype Workers.— TL 2.0-2.3, HL 0.50-0.56,
HW 0.43-0.48, SL 0.31-0.36, EL 0.04-0.06 (5-10 fac-
ets), PW 0.30-0.37, WL 0.52-0.60, SpL 0.07-0.10,
PetL 0.21-0.24, PpetL 0.11-0.13mm, CI 0.83-0.86,
OI 0.10-0.13, SI 0.72-0.78, PSI 0.14-0.17, MHI 0.93-
1.03. N=9
Like curvipubens, but differing in the follow-
ing ways in addition to diagnosis. Relative widths
of nodes with slightly different ranges (PetW/
PetL 0.56-0.70); PpetW/PpetL 1 .38-1 .52). Sting ap-
paratus of specimens from Oaxaca (paratypes)
and Vera Cruz, Mexico with strong, acute sting
shaft and lancets as in inennis (Fig. 42).
Posterior head with transversely arching rug-
ose-areolate macrosculpture. Compared to
curvipubens, rugae on mesosoma dorsum with
more lateral spurs that may connect rugae and
create areolae on anterior pronotum and on met-
anotum. Macrosculpture on mesosoma sides ab-
sent or weakly rugose-areolate. Dorsal face of
propodeum lacks macrosculpture.
Head dorsum with 0-16 hairs suberect hairs;
mesosoma dorsum with 1-8 pairs (usually 2-7).
Erect hair on gaster Tl usually limited to posterior
margin, but entirely absent from Jalisco specimen
and entirely covering the tergum of the Yucatan
specimen. The Yucatan specimen is also unique in
having some stiff, spatulate hairs on head,
mesosoma and gaster.
Paratype and Nontype Queens. — TL 2.3-2.5, HL
0.53-0.56,HW 0.45-0.50, SL 0.35-0.38, EL 0.10-0.11,
PW 0.39-0.45, WL 0.65-0.72, SpL 0.11-0.14, PetL
0.22-0.25, PpetL0.13-0.15mm, CI 0.85-0.89, SI 0.76-
0.78, PSI 0.16-0.19. N=2
Queen as in curvipubens, except for shape of
postpetiole, sting, and sculpture as in workers of
cuneola. Mandibles with 6 or 7 teeth. Sides of head
and mesokatepisterna strongly microareolate and
opaque. Paratype queen with erect-suberect hair
over whole gaster Tl.
Discussion. — See micromma and minima dis-
cussions for comparisons with other tiny Rogeria.
The name cuneola (L., small wedge) refers to the
shape of the postpetiolar sternum in lateral view.
Distribution. — These tiny ants are most often
taken in berlesate of leaf litter and rotten wood.
Some come from sittings under termite mounds
and one was collected in a Cattelya orchid. Habitat
of most specimens is rain forest or mesic forest,
either primary or secondary growth, but one speci-
men was found in Yucatan thorn forest.
Volume 3, 1994
69
Material Examined. — Holotype locality.
MEXICO: Oaxaca State, 1 mi. E Reforma, 15-VIII-
1973, litter, tropical evergreen forest (A. Newton)
[MCZ].
Paratype localities. MEXICO: 14 workers, ho-
lotype locality [ BMNH, CKC, LACM, MCZ, MZSP,
USMN]; 1 worker, 1 queen, Oaxaca State, 1 mi. E
Reforma, near Tuxtepec,12-15-VIII-1973, litter for-
est floor (A. Newton) [3 stings, 1 worker coated for
SEM] [MCZ].
Nontype localities. MEXICO: 1 worker, San
Luis Potosi State, El Salto, 6-VII-1969, #B-166 (S.
and J. Peck) [MCZ]; 2 workers, Vera Cruz State,
Pueblo NuevonearTetzonapa,17-VIII-1953,#B.F.
(E. O. Wilson) [MCZ]; 2 workers, Vera Cruz State,
El Palmar near Tetzonapa, 8-VIII-1953, #173 (E. O.
Wilson) [sting, whole specimen] [MCZ]; 1 worker,
Jalisco State, 6km N El Tuito, 31-XII-1987, #9327-6
(P. S. Ward) [MCZ]; 7 workers, 1 queen, Chiapas
State, Palenque, 4-IX-1974 (E. M. and J. L. Fisher)
[LACM]; 1 worker, Yucatan State, Uxmal, 27- VII-
1963, #118 (E. O. Wilson) [MCZ]. BELIZE: 4 work-
ers, Belmopan, 1-15-VIII-1972, #B-243 (S. and J.
Peck). GUATEMALA: 1 worker, 14-15- V-1946[?],
SF-21214, 46-6134 (no locality orcollector) [USNM] .
HONDURAS: 2 workers, La Lima, 4-IX-1961, UCF
217-128 (C. Evers Q.) [MZSP]. COSTA RICA: 1
worker, Guanacaste Province, Santa Rosa National
Park, 14-VII-1983 (P. S. Ward) [CKC]; 3 workers,
Puntarenas Province, Manuel Antonio National
Park, 27-VII-1985, #7692-9 (P. S. Ward) [MCZ].
Rogeria micromma Kempf
Fig. 71
Rogeria micromma Kempf 1961:509 (Figs. 12-13) . Holotype
worker, SURINAM: Dirkshoop (J. vander Drift) [MZSP]
[Holotype and La Poulle paratype examined]
Diagnosis. — WL 0.45-0. 52mm. Clypeal apron
medially flattened. Eye tiny. Postpetiolar node
subrectangular in dorsal view and not strongly
vaulted in side view; anterior lip of sternum not
prominent, posterior edge angular in side view.
Sides of head and mesosoma and dorsal face of
propodeum opaque with dense microareolate
sculpture. Laterodorsa and sides of head also finely
macroareolate. Propodeum free of macrosculpture.
No erect hair on scapes or extensor surfaces of
legs. Dorsa of head, mesosoma, nodes and gaster
Tl with short, appressed hairs and longer, erect to
suberect hairs. Mesosoma dorsum with 8-10 pairs
of erect hair; each node with 2 pairs of
posterodorsally projecting hairs.
Workers.— TL 1.7-1.8, HL 0.44-0.51, HW 0.37-
0.45, SL 0.28-0.35, EL 0.02-0.04 (2-5 facets), PW
0.28-0.30, WL 0.45-0.52, SpL 0.07-0.08, PetL 0.17-
0.18, PetW 0.10-0.13, PpetL 0.10-0.11, PpetW 0.13-
0.15mm, CI 0.84-0.88, OI 0.05-0.08, SI 0.76-0.79, PSI
0.15-0.16, MHI 0.92-0.96. N=3
The following supplements diagnosis and
Kempf (1961). Mandibles 5-toothed to 6-toothed;
basal tooth not larger than penultimate basal.
Mesosoma profile of holotype interrupted by a
weak metanotal groove followed by two trans-
verse carinulae, but profiles of La Poulle and Bra-
zil specimens uninterrupted. Brazil specimen with
narrower propodeal spines than in types, and
propodeal spiracle closer to posterior edge of
propodeum. Petiole short (PetL//WL 0.35-0.38),
with ventral keel and tooth.
In Surinam specimens, vague microareolate
microsculpture densely covers head, dorsum of
mesosoma, dorsal face of propodeum, petiole and
postpetiole of types, producing a weakly shining,
granular appearance. Brazil specimen similar, ex-
cept for smoother ventral petiolar peduncle and
postpetiolar dorsum. Sides of mesosoma and pos-
terior surface of head more distinctly microareolate.
Anterior of pronotal disc with 1 to 2 transverse
rugae followed by longitudinally rugose to ar-
eolate-rugose macrosculpture that disappears into
microsculpture on meso- and metanota. Sides of
mesosoma with sparse, faint longitudinal rugulae
on meso- and metapleura. Nodes microareolate;
more effaced on postpetiole.
Erect hairs of Dirkshoop specimen all trichoid;
but thicker and stiff on the La Poulle specimen, at
least some cuneate; erect hairs on the Belem speci-
men seem intermediate.
Color brownish-yellow; legs, mandibles and
antennae sometimes slightly lighter. Frontocly peal
region not lighter than rest of head.
Discussion. — It may be that the three micromma
specimens are just unusual curvipubens or cuneola,
but at present there are enough differences to
provisionally retain this species. Eighteen
curvipubens (Figs. 74-76, 101-102) workers from
other van der Drift collections in Dirkshoop and
La Poulle, are very similar to micromma in size and
shape, but differ as follows: 1 ) clypeal apron evenly
convex, 2) sides of head and mesosoma shinier
with effaced microsculpture, 3) sides of head with
70
Journal of Hymenoptera Research
rugose macrosculpture, and 4) reduced pilosity.
Some cuneola (Figs. 77-78, 103) specimens are
also very similar to micromma in shape and size,
and one from Yucatan has stiff erect hairs on head,
mesosoma, waist and gaster Tl , but cuneola work-
ers differ in shape of the postpetiolar sternum, as
well as in clypeal shape and pilosity.
Some alzatei from Panama, Colombia, Guyana,
and French Guiana are only slightly larger (WL
0.51-0.68mm) than micromma and have the same
pilosity and similar structure and sculpture, but
they generally have distinctly larger eyes with
more than 10 facets, have a higher, more compact
mesosoma (MHI 1.00-1.04), and generally nar-
rower propodeal spines. Severalrt/za to from north-
ern Colombia have reduced eyes (7-8 facets) and
one has wider propodeal spines, but those Colom-
bian ants are larger, have a higher mesosoma, and
more abundant erect pilosity. See also the minima
discussion.
Material Examined. — SURINAM: Dirkshoop (J.
van der Drift), La Poulle 0- van der Drift). BRAZIL:
Para, Pirelli Plantation (Iriboca) nr. Belem, (W. L.
Brown). 3 workers [MCZ, MZSP].
Rogeria minima Kusnezov
Figs. 72-73
Rogeria minima Kusnezov 1958:44, Figs. 1-3. Holotypedealate
queen, ARGENTINA: Tucuman [FML] [Holotvpe ex-
amined].
Known only from a single queen mounted on a microscope
slide showing dorsal head (nearly split in half length-
wise), ventral maxillae and labium, lateral mesosoma
and petiole, and a mostly ventral view of postpetiole
and gaster.
Diagnosis. — WL of worker probably < 0.60mm.
Mandibles triangular. Palpal formula 2,1. Sting
shaft and lancets spatulate. Postpetiole widest in
anterior half; anterior lip of sternum not promi-
nent. Mesosoma predominantly rugose. No
microsculpture on head dorsum, little on
mesosoma sides; but microareolate sculpture
present on gaster Tl and SI. No erect hairs on
scapes or extensor surfaces of legs. Head dorsum
with abundant erect hair; mesosoma dorsum with
more than 12 pairs; petiolar node with 2 pair
projecting posterodorsally. Most erect hairs cu-
neate-fimbriate.
Queen.— TL 2.3, HL 0.55, HW 0.53, SL 0.34, EL
0.10, WL 0.65, SpL 0.12, PetL 0.22, PpetL 0.11mm,
CI 0.96, SI 0.64, PSI 0.18.
Mandibles with 5 teeth; basal only slightly
larger than penultimate basal. Palpal formula 2,1.
Clypeus torn; shape of apron unclear. Eyes mod-
erately large, with about 20 facets. Mesosoma as
shown in Fig. 72. Parapsidal furrows cannot be
discerned. Petiole with ventral tooth and
nonlamellate keel. Postpetiole wider (0. 18mm) than
long; sides of postpetiole seem to be convex in
front, then tapered, as in many curvipnbens (Fig.
74); sternum seems low and not prominent or
wedge-shaped. Pygidial gland sculpture present,
no tubercles on posterior surface. What is visible
of the sting apparatus looks like that of inermis
(Fig. 42), except that lancet apices lack barbules
and sting shaft seems to have little, if any, terminal
flange. Sinceboth sting shaft and lancets are folded,
they are probably weakly sclerotized.
Median head with diverging rugae continu-
ing on posterior head as diverging rugose-areolate
sculpture; laterodorsa areolate-rugose. No
macrosculpture on median pronotum; sides
weakly and incompletely areolate, especially
ventrad. Mesonotum longitudinally rugose. Dor-
sal half of mesopleura longitudinally rugose; most
of mesokatepisterna smooth, but with some weak
areolate sculpture along posterior margins.
Metapleural and propodeal sides confused ar-
eolate-rugose. Mesosoma lacks microsculpture,
except on median pronotum, sides of propodeal
spines, and metapleural lobes (metanotum and
median propodeum could not be examined). All
surfaces of petiole, at least venter and sides of
postpetiole, and at least sterna of gaster minutely
and shallowly areolate with sharp, thin partitions.
Much of dorsal aspect of gaster difficult to see, but
at least anterior and lateral portions of Tl, and the
terminal terga are also shallowly microareolate.
Body covered with appressed to decumbent
setiform pilosity. In addition, dorsa of head,
mesosoma nodes, and gaster Tl and SI with erect
cuneate-fimbriate hairs (Fig. 73). Mesosoma dor-
sum with more than 12 pairs of erect hairs; petiolar
node with 2 pairs of posterodorsally projecting
erect hairs (postpetiolar hairs hidden). All hair on
terminal segments of gaster are setiform.
Discussion. — A worker of micromma from La
Poulle, Surinam and a worker of cuneola from
Uxmal, Yucatan come very close in size, sculpture,
and in having cuneate pilosity, but the gasters of
both are smooth except for piligerous punctures. I
cannot tell if any hairs are fimbriate.
Volume 3. 1 994
71
Fore/i-Group
Rogeria foreli Emery
Figs. 79-82, 104-105
Rogeria foreli Emery 1894:191. Holotype worker, VIRGIN
ISLANDS: St. Thomas (Eggers) [MHN] [Holotype ex-
amined).
Rogeria foreli gaigei Forel 1914:617. Holotype worker, CO-
LOMBIA (Gaige) [MHN]) [Holotypeexamined], N. syn.
Rogeria huaciiucana Snelling 1973:4, Fig. 1. Holotype and
paratype worker, USA: Arizona, Cochise County
(Snelling) [LACM] [Holotype and 1 paratype exam-
ined]. N. syn.
Additions to foreli-gioup diagnosis. Basal
mandibular teeth abruptly smaller than apical
teeth. Clypeal apron convex, often with a faint
median angle. Eyes usually 10 or more facets.
Workers.— TL 1.9-2.9, HL 0.50-0.71, HW 0.43-
0.62, SL 0.32-0.51, EL 0.06-0.10 (7-20 facets), PW
0.30-0.45, WL 0.50-0.80, SpL 0.07-0.15, PetL 0.20-
0.32, PpetL 0.12-0.19mm, CI 0.83-0.89, SI 0.74-0.85,
OI 0.12-0.17, PSI 0.14-0.20. N=25
Mandibles with 4-7 teeth and 0-3 denticles.
Generally teeth 1-4 decrease in size gradually,
then teeth 5-7 (if present) abruptly smaller and
possibly interspersed with one or more denticles.
Sometimes basal tooth is distinctly larger than
penultimate tooth. Body of clypeus often project-
ing slightly over the anterior clypeal margin. Pos-
terior outline of head flat to weakly convex. Eyes
small, oval. Nuchal grooves inconspicuous in lat-
eral view. Pronotal shoulders well rounded. Met-
anotal groove generally absent (Fig. 79), but may
be weakly to distinctly (Fig. 80) visible. Anterior
border of propodeum not marked by a ridge.
Metapleural lobes small, broadly rounded. Peti-
olar node shape varies between extremes shown
in Figs. 79 and 81; smaller nodes are as long as
wide, larger nodes are longer than wide.
Postpetiolar node peaks in posterior half;
subrectangular in dorsal view. Anterior lip of
postpetiolar sternum small (Fig. 79), or prominent
(Figs. 81, 104).
Mandibles, median clypeus, legs, posterior
face of propodeum, gaster, and sometimes sides of
petiolar peduncle smooth, except for minute
piligerous punctures. Rest of body densely
microareolate or microcolliculate, often appear-
ing granular at low magnification. Microareolate
sculpture on head is more distinct near antennal
insertions and more effaced caudad, sometimes
nearly smooth on sides of head. Microareolate
pattern distinct on meso- and metapleura and
generally on dorsal face of propodeum; indistinct
on promesonotum and petiole and vestigial on
postpetiole (Figs. 104-105). Microsculprure over-
lain by very fine longitudinal rugulae on lateral
clypeus, cheeks, frontal lobes, middorsum and
sometimes laterodorsa; posterior head with very
fine transverse or diverging rugulae.
Color yellow with a slightly brownish gaster
to chestnut-brown with brownish-yellow or light
brown appendages.
Queens.— TL 2.4-3.1, HL 0.54-0.69, HW 0.48-
0.59, SL 0.36-0.49, EL 0.10-0.16, PW 0.38-0.51, WL
0.64-0.87, SpL0.12-0.17,PetL0.24-0.32,PpetL0.13-
0.20mm, CI 0.86-0.93, SI 0.72-0.83, PSI 0.17-0.20.
N=8
Differing from the workers in the normal queen
attributes and in the following. One specimen
with 3 teeth and 5 denticles; the others with the
same variation as in the workers. Posterior outline
of head with weak median concavity in some.
Parapsidal furrows are barely discernable.
Mesoscutum in all specimens longitudinally ru-
gulose; mesoscutellum vaguely rugose to areolate-
rugose.
Discussion. — Side by side comparison of the
types of foreli and gaigei revealed that the only
difference between them is size; but the gaigei type
is well within the size variation of foreli specimens.
Although the types of huachucana come from Ari-
zona, far from other known foreli specimens, and
were collected in an unusual habitat, they differ
from the foreli holotype only in having: 1 ) 7-8 facets
in the eyes (vs. 12), and 2) a weak metanotal groove
(vs. none). Since eye size and metanotal groove
development vary continuously and not concor-
dantly in foreli specimens from Central and South
America, and since equally small eyes and even
more distinct metanotal grooves are present in
those specimens, I am unable to distinguish the
huachucana specimens as a distinct species at this
time.
While working in Northern Colombia for two
years I regularly collected two kinds of foreli, which
I could distinguish at a glance by the shapes of
their postpetiolar sterna (Figs. 79, 81). Specimens
from Barro Colorado Island, Panama, and Trinidad
also have the same two types of sterna. In Colom-
bia and Panama, both morphs have been taken
from the same locality, but it is not known whether
72
Journal of Hymenoptera Research
the two morphs come from different colonies or
not. I considered calling the specimens with a
more prominent, shelf-like postpetiolar sternum a
new species, but the difference is sometimes subtle
and individuals difficult to assign. Without cor-
roboration from another character, I decided
against erecting a new species at this time.
Rogeria foreli is closely related to bruchi, which
at present is known only from Argentina and
Paraguay, much farther south than known foreli
specimens (see the bruchi description).
Distribution. — Rogeria foreli has a disjunct dis-
tribution: Panama and northern South America,
the Caribbean, and Southwestern United States.
Specimens come from Om to 610m in Panama, 0m
to 240m in Colombia and Venezuela generally
from berlesate of leaf mold and rotten wood. The
Arizona specimens were under stones in an oak-
juniper woodland at 1783-1814m.
Material Examined. — USA: Arizona, Cochise
County (R. R. Snelling). PUERTO RICO: Rio Piedras
(J. A. Torres). ST. CROIX: Buck Island. ST. THO-
MAS (Eggers). TRINIDAD: (N. A. Weber).
PANAMA: Barro Colorado Island (W. L. Brown,
W. L. Brown and E. S. McCluskey, A. Newton, N.
A. Weber, J. Zetek); Gahin (W. L. Brown); Ancon
(no collector) [sting]; Gamboa (P. S. Ward); Colon
Province, Frijoles (J. Ventocilla); Cerro Azul (A.
Newton). COLOMBIA: no locality (Gaige);
Magdalena Department, Tayrona Park (C. Kugler,
P. S. Ward) [mouthparts, sting]; Tigrera (W. L.
Brown and C. Kugler) [queen mouthparts, whole
specimen; worker mouthparts, sting] ; Guaj ira, Don
Diego (W. L. Brown and C. Kugler). VENEZU-
ELA: Barinas State, near Ciudad Bolivia (P. Ward);
Sucre State, 32km W. Campano (S. & J. Peck). 106
workers, 17 queens, 4 males [CKC, CUIC, LACM,
MCZ, MCZ, MZSP, USNM].
Rogeria bruchi Santschi
Fig. 82
Rogeria bruchi Santschi 1922:352. Holotype worker, ARGEN-
TINA, Buenos Aires, Monte Veloz (C. Bruch) [NHM]
[Holotype examined).
Workers.— TL 2.2-2.6, HL 0.56-0.65, HW 0.48-
0.57, SL 0.38-0.46, EL 0.04-0.06 (5-7 facets), PW
0.35-0.40, WL 0.58-0.70, SpL 0.10-0.17, PetL 0.21-
0.29, PpetL 0.13-0.17mm, CI 0.61-0.70, OI 0.10-
0.12, SI 0.76-0.81, PSI 0.17-0.24. N=5
The five known specimens of bruchi are just
like the foreli with nonprojecting clypeus, weak or
absent metanotal groove, and nonprojecting
postpetiolar sternum, but differ in the following
ways: 1) Mandible with five large teeth, second to
fifth subequal (Fig. 82), 2) clypeus with a median
notch, 3) eyes smaller than in most foreli.
Material Examined. — PARAGUAY: Misiones
Province, 30km S. of San Juan Bautista (F. Baud et
al.); Concepcion Province, 50km E. of San Lazaro
(F. Baud, et al.) [mouthparts, sting]. ARGENTINA:
Buenos Aires Province, Monte Veloz (C. Bruch);
Misiones Province, Posadas (A. Breyer). 5 workers
[MCZ, MHN, MZSP, NHM],
ACKNOWLEDGEMENTS
I am very grateful to the following people for loans or
gifts of specimens: Cesare Baroni Urbani (NMB), Claude
Besuchet (MHN), Barry Bolton (BMNH), Carlos Roberto
Brandao (MZSP), William L.Brown, Jr. (MCZ), John E.Lattke,
Jack Longino, Alfred Newton (MCZ), David M. Olson, Roberto
Poggi (MCSN), David Smith (USNM), Roy R. Snelling
(LACM), Philip S Ward, and E. O. Wilson (MCZ). Jack
Longino also generously sent me his unpublished field col-
lections notes.
Lily Fainter of the Virginia-Maryland Regional College
of Veterinary Medicine and Bob Honeycutt of Virginia Poly-
technic Institute and State University, Department of Forest
Products assisted with the scanning electron microscopy.
Reed R. Lambert helped collect ants in Peru and Bolivia.
Richard L. Hoffman, W. L. Brown, Jr., Andre Francoeur, and
R. W. Taylor provided valuable advice. Special thanks to Roy
Snelling, who made many helpful criticisms of a draft. None
of these bear any responsibility for the product.
The work was supported financially by NSF grant #DEB-
8022177 and Radford University.
LITERATURE CITED
Bolton, B. 1973. The ant genera of West Africa: A synonymic
synopsis with keys (Hymenoptera: Formicidae). Bulle-
tin of the British Museum (Natural History), Entomology
27(6):319-368.
Bolton, B. 1982. Afrotropical species of the myrmicine ant
genera Cardiocondyla, Leptothorax, Melissotarsus, Messor
and Cataulacus (Formicidae). Bulletin of the British Mu-
seum (Natural History), Entomology 45(4):307-370.
Bolton, B. 1987. A review of the Solenopsis genus-group and
revision of afrotropical Monomorium Mayr (Hy-
menoptera: Formicidae). Bulletin of the British Museum
(Natural History), Entomology 54(3):263-452.
Borgmeier, T. 1953 (1951). Algumas formigas do genero
Macromischa Roger. Arquwos do Museu Nacwnal Rio de
/amero 42:107-112.
Brown, W. L. 1952. Synonymous ant names. Psyche 58(3): 124.
Brown, W. L. 1953. Characters and synonymies among the
Volume 3. 1 994
73
genera of ants. Part II. Brevwra no. 18:1-13.
Brown, W. L. 1988. Data on malpighian tubule numbers in
ants (Hymenoptera: Formicidae). Pp. 17-27 in: J. C.
Trager (ed.) Advances in myrmecology . Leiden: E. J. Brill,
xxvii+551 pp.
Brown, W. L., and W. L. 1949. Wing venation and the phylog-
eny of the Formicidae. Transactions of the American Ento-
mological Society 75(3-4):113-134.
Emery, C. 1894. Studi sulle formiche della fauna neotropica.
V1-XV1. Bullettnio della Societa Entomologica Italiana26:\37-
242 + 4 pi.
Emery, C. 1896. Studi sulle formiche della fauna neotropica.
XVII-XXV. Bullettmo della Societa Entomologica ltaliana
28:33-107.
Emery, C. 1897. Formicidarum species novae vel minus
cognitae in collectione musaei nationalis Hungarici quas
in Nova-Guinea, colonia germanica, collegit L. Biro.
Termeszetrajzi Fuzetek 20:571-599 + 2 pi.
Emery, C. 1914. Les fourmis de la Nouvelle-Caledonie et des
lies Loyalty. P 415 In: F. Sarasin and J. Roux (eds.) Nova
Caledonia recherches scientifiques en Nouvelle Caledonie et
aux iles Loyalty (Zoologie). Wiesbaden: C. W. Kreidels
Verlag, l(4):389-436+13 pi.
Emery, C. 1915. Noms de sous-genres et de genres proposes
pour la sous-famille des Myrmicinae. Modifications d la
classification de ce groupe (Hymenoptera Formicidae).
Bulletin de la Societe Entomologique de France 1915:189-
192.
Forel, A. 1899. Formicidae. Biologia Centrali-Americana. Insecta.
Hymenoptera 3:1-160+4 pi.
Forel, A. 1914. Einige amerikanische ameisen. Deutsche
Entomologische Zeitschrift 1914:615-620.
Harris, Rick A. 1979. A glossary of surface sculpturing. Occa-
sional Papers in Entomology. Department of Food and Agri-
culture, State of California no. 28:1-31.
Holldobler, B. and H. Engel. 1978. Tergal and sternal glands
in ants. Psyche 85(4):285-330.
Holldobler, B., R. Stanton, R., and Engel. 1976. A new exo-
crine gland in Novomessor (Hymenoptera: Formicidae)
and its possible significance as a taxonomic character.
Psyche 83(1 ): 32-41.
Holldobler, B. and E. O. Wilson. 1990. The Ants. Cambridge,
Massachusetts: Belknap Press, xii+732 pp.
Kempf, W. W. 1961. A survey of the ants of the soil fauna in
Surinam. Studia Entomologica 4(l-4):481-524.
Kempf, W. W. 1962a (1961). Remarks on the ant genus Irogera
Emery, with the description of a new species (Hy-
menoptera, Formicidae). Rei'istaBrasileiradeEntomologia
21(4):435-441.
Kempf, W. W. 1962b. Miscellaneous Studies on Neotropical
Ants. II. (Hymenoptera, Formicidae). Studia Entomologica
5(l-4):l-38.
Kempf, W. W. 1963. Additions to the neotropical ant genus
Rogeria Emery, with a key to the hitherto recorded South
American species (Hym., Formicidae). Remsta Brasileira
de Entomologia 23(2):189-196.
Kempf, W. W. 1964. Miscellaneous studies on neotropical
ants. III. Studia Entomologica 7(l-4):45-71.
Kempf, W. W. 1965. Nota preliminar sobre algumas formigas
neotropicas, descritas por Frederick Smith (Hy-
menoptera, Formicidae). Revista Brasileira de Entomologia
25(2):181-186.
Kempf, W. W. 1972. Catalogo abreviado das formigas de
regiao neotropical (Hym. Formicidae). Studia
Entomologica 15(l-4):3-344.
Kempf, W. W. 1975. Miscellaneous studies on neotropical
ants. VI. (Hym. Formicidae). Studia Entomologica 18(1-
4):341-380.
Kugler, C. 1978a. Pygidial glands in the myrmicine ants
(Hymenoptera, Formicidae). Insectes Sociaux 25(3):267-
274.
Kugler, C. 1978b. A comparative study of the myrmicine
sting apparatus (Hymenoptera, Formicidae). Studia
Entomologica 20:413-548.
Kugler, C. 1980. The sting apparatus in the primitive ants
Nothomyrmecia and Myrmecia. Journal of the Australian
Entomological Society 19:263-267.
Kugler, C. 1986. Stings of ants of the tribe Pheidologetini
(Myrmicinae). Insecta Mundi l(4):221-230.
Kugler, C. 1992 (1991). Stings of ants of the tribe Ectatommini
(Formicidae: Ponerinae). Insecta Mundi 5(3-4):153-166.
Kusnezov, N. 1958. La posicion sistematica del genero Rogeria,
con descripcion de una nueva especie. Acta Zoologica
Lilloana 15:41-45.
Linneaus, C. 1758. Systema Naturae Regnum Animate ed. 10,
Tom 1. Stockholm, VI+824pp.
Mann, W. M. 1919. The ants of the British Solomon Islands.
Bulletin of the Museum of Comparative Zoology, Harvard
63(7):273-391 + 2 pi.
Mann, W. M. 1921. The ants of the Fiji Islands. Bulletin of the
Museum of Comparative Zoology, Harvard 64(5):401-499.
Mann, W. M. 1922. Ants from Honduras and Guatemala.
Proceedings of the United States National Museum 61(13):1-
54.
Mann, W. M. 1925. Ants collected by the University of Iowa
Fiji-New Zealand expedition. University of Iowa Studies
m Natural History ll(4):5-6.
Mayr, G. L. 1886. Notizen ueber die Formiciden-Sammlung
des British Museum in London. Verhandlungen der
Zoologisch-Botanischen Gesellschaft in Wein 36:353-368.
Roger, J. 1863. Die neu aufgefuehrten Gattungen und Arten
meines Formicidenverzeichnisses, nebst Ergaenzungen
einiger frueher gegebenen Beschreibungen. Berliner
Entomologische Zeitschrift 7:131-214.
Santschi, F. 1922. Myrmicines, dolichoderines et autre
formicides neotropiques. Bulletin de la Societe Vaudoise
Sciences Naturelles 54(205):345-378.
Santschi, F. 1923. Description de quelques nouvelles fourmis
du Bresil. Reinsta (Museu Paulista, Sao Paulo, Brazil)
13:1253-1264,
Santschi, F. 1930. Quelques fourmis de Cuba et du Bresil.
Bulletin de la Societe Royale de Botamque d'Egypte Mai
1930(2-3):75-83.
Santschi, F. 1936. Contribution & l'etude des fourmis de
l'Amerique du Sud. Revista de Entomologia, Rio de Janeiro
6(2):196-218.
Santschi, F. 1941. Quelques fourmis japonaises inedites.
Mitteilungen der Schweizerischen Entomologischen
Gesellschaft 18:273-279.
Smith, F. 1858. Catalogue of the hymenopterous insects in the
collection of the British Museum. Part VI. Formicidae. Lon-
don, 216 pp + 14 pi.
Smith, M. 1937. The ants of Puerto Rico. University of Puerto
Rico, Journal of Agriculture 20(4):819-875.
Snelling, R. R. 1973. Two ant genera new to the United States.
Contributions in Science, Natural History Museum of Los
Angeles County no. 236:1-8.
Snelling, R. R. 1989. Untitled comment. Notes From Under-
74
Journal of Hymenoptera Research
ground: A Myrmecological Newsletter. MCZ Laboratories,
Harvard, MA no. 3, pp. 7-8.
Trager, J. C. 1989. A pitch for standardized orientation of the
ant head, and related matters. Notes from Underground: A
Myrmecological Newsletter. MCZ Laboratories, Harvard,
MA. no. 1, pp. 5-7.
Weber, N. A. 1934. Notes on neotropical ants, including the
descriptions of new forms. Revista de Entomologia, Rw de
Janeiro 4(l):22-59.
Wheeler, G. C. 1989. Vive la myrmecologie. Notes from Under-
ground: a Myrmecological Newsletter. MCZ Laboratories,
Harvard, MA. no. 3, pp. 6-7.
Wheeler, G. C. and J. Wheeler. 1953. The ant larvae of the
myrmicine tribe Pheidolini. Proceedings of the Entomo-
logical Society of Washington 55(2):49-84.
Wheeler, G. C. and J Wheeler. 1973. Ant larvae of four tribes:
Second supplement (Hymenoptera: Formicidae:
Myrmicinae). Psyche 80(l-2):70-82.
Wheeler, G. C. and J. Wheeler. 1976. Ant larvae: Review and
synthesis. (Memoires of the Entomological Society of
Washington, no. 7). Washington: Entomological Society
of Washington D.C. vi + 108 pp.
Wheeler, G. C. and J. Wheeler. 1977. Supplementary studies
on ant larvae: Myrmicinae. Transactions of the American
Entomological Society 103:581-602.
Wheeler, G. C and J. Wheeler. 1986. Supplementary studies
on ant larvae: Myrmicinae (Hymenoptera: Formicidae).
journal of the New York Entomological Society 94(4):489-
499.
Wheeler, G. C. and J. Wheeler. 1988. Notes on ant larvae:
Myrmicinae. Transactions of the American Entomological
Society 114:319-327.
Wheeler, W M. 1911. A list of the type species of the genera
and subgenera of Formicidae. Annals of the Neiv York
Academy of Sciences 21:157-175.
Wilson, E. O 1955. A monographic revision of the ant genus
Lasius. Bulletin of the Museum of Comparative Zoology,
Harvard 113(1):1-201.
Wilson, E. O. and R. W. Taylor. 1967. The ants of Polynesia
(Hymenoptera: Formicidae). Pacific Insects Monograph
14:1-109.
Frontal area
Lateral clypeus
Denticles
Posterior head
Middorsum
Laterodorsum
Body of clypeus
Clypeal apron
>,/ Nuchal groove
Side
Venter
Erect
1
Suberect
Appressed
Fig. 1. Illustration of some terms and measurements used in descriptions. Dotted lines on the heads show approximate
boundaries of the regions. The dashed line is an extension of a line bisecting the propodeal spines. Where it crosses the
mesosoma outline gives a rough measure of spine inclination. SpL = spine length. WL = Weber's length. The mesosoma height
index (MHI) is distance a (mesosoma height) divided by distance b.
Volume 3, 1 994
75
gibba
Figs. 2-6. Figs. 2-4. Rogerm ciliosa. 2. Holotype profile; dorsal head. 3. Paratype sting apparatus. Above: lateral views spiracular
plate (SP), quadrate plate (QP), oblong plate (OP), gonostylus (Go), triangular plate (TP), and lancet (La). Below left: ventral
sting, furcula. Below right: lateral sting, furcula (Fu). 4. Paratype queen lateral mesosoma. Figs. 5-6. Rogeria gibba. 5. Holotype
profile; dorsal head. 6. Paratype sting with furcula in lateral and ventral views; lateral view of lancet apex.
76
Journal of Hymenoptera Research
14 prominula
Figs. 7-14. Figs. 7-12. Rogeria stigmatica. 7. Syntype dorsal mandibles, clypeus. 8. Syntype profile (pilosity reconstructed by
comparison with other specimens); nontype lateral gaster showing typical pilosity (Fulakora, Solomon Islands). 9. Lateral and
ventral views of sting with furcula (Falepuna, Western Samoa). 10. Queen mesosoma profile (Viti Levu, Fiji). 11. Male
(Sigatoka, Viti Levu). 12. Genitalia (same male). Fig. 13. Rogeria megastigmatka holotype profile. Fig. 14. Rogeria prominula
holotype profile (appressed pilosity not shown); dorsolateral view mandibles and clypeal margin; dorsal head. Same scale for
all external views.
Volume 3, 1994
77
subarmata
Figs. 15-23. Figs. 15-16. Rogeria besucheti. 15. Holotype profile; dorsal head. 16. Paratype ventral and lateral views of sting with
furcula. Fig. 17. Rogerui blanda male genitalia (Rio Akabin, Venez.). See Figs. 83-84 for worker. Fig. 18. Rogeria procera profile
(Rio Cuming, Braz. — " brasiliensis" holotype). Figs. 19-20. Rogeria scartdens syntype. 19. Profile; dorsal head; dorsal waist. 20.
Erect hair. Fig. 21. Rogeria terescandens holotype profile; dorsal waist. Figs. 22-23. Rogeria subarmata. 22. Paratype profile. 23.
Nontype lateral propodeum, petiole (Belem, Braz).
78
Journal of Hymenoptera Research
germami
lirata
lacertosa
Figs. 24-31. Figs. 24-27. Rogeria germaini. 24. Profile (mesosoma hairs had been matted down somewhat), dorsal head; dorsal
postpetiole (Passa Quatro, Brazil — "minensis" lectotype). 25. Mesosoma dorsum profile showing typical pilosity (Central
Prov., Para.). 26. Lateral petiole (hair omitted) (Central Prov., Para.). 27. Lateral sting, furcula; ventral sting shaft (San Benito
Is., Para.). Figs. 28-30. Rogeria lirata. 28. Holotype profile; dorsal head. 29. Paratype lateral lancet (nr. Yunmaguas, Peru). 30.
Nontype queen wings (Jatai, Braz.) Cubital vein (Cu), median vein (M), radio-medial crossvein (r-m), radial sector vein (Rs),
submarginal cell (SM). Fig. 31. Rogeria lacertosa paratype profile. Same scale for all external views except wings.
Volume 3, 1 994
79
Figs. 32-39. Figs. 32-33. Rogerm pellecta. 32. Paratype profile; dorsal postpetiole. 33. Paratype lateral sting, furcula. Fig. 34.
Rogeria sicaria holotype profile. Figs. 35-39. Rogeria belti. 35. Syntype profile; dorsal head. 36. Nontype lateral head, mesosoma
(hair omitted) (Miami, C. R.). 37. Male wings (La Ceiba, Hond.). 38. Male (La Ceiba, Hond.). 39. Genitalia (same male). Same
scale for all external views.
80
Journal of Hymenoptera Research
neilyensis
Figs. 40-43. Figs. 40-42. Rogeria inermis. 40. Syntype profile showing smallest propodeal spines. 41. Nontype mesosoma dorsum
profile (hairs omitted) showing longest propodeal spines (Osa Penin., C. R.). 42. Sting apparatus (La Selva Station, C. R). Right:
lateral views of spiracular plate, quadrate plate (QP), oblong plate (OP), gonostylus, triangular plate and lancet; dorsal view
anal plate (AP). Below: Lateral and ventral views sting, furcula. Fig. 43. Rogeria neilyensis holotype profile; dorsal head.
Volume 3, 1 994
81
exsulans
cornuta
merenbergiana
Figs. 44-48. Fig. 44. Rogeria exsulans holotype profile. Fig. 45. Rogeria cornuta holotype profile; dorsal head. Figs. 46-48. Rogeria
merenbergiana. 46. Holotype profile. 47. Nontype mesosoma dorsum profile (hair omitted) showing the least distinct
metanotum (Cuevas de los Guacharos, Col.). 48. Paratype queen lateral mesosoma and waist. Same scale for all figures.
82
Journal of Hymenoptera Research
unguispma
creightoni
Figs. 49-54. Fig. 49. Rogena unguispma holotype profile; dorsal head; dorsal waist. Fig. 50. Rogena brunnea profile; dorsal
postpetiole (Sierra del Rosario, Cuba — "cubensis" holotype). Figs. 51-54. Rogena creightoni. 51. Paratype profile; dorsal head,
dorsal waist. 52. Nontype profile (Ocozocoautla, Mex). 53. Nontype profile; dorsal waist (Costa Rica, Nevermann leg.). 54.
Lateral sting, furcula (Caves Branch, Belize). Same scale for all external views.
Volume 3. 1 994
83
Figs. 55-62. Figs. 55-57. Rogeria innotabilis. 55. Holotype profile; dorsal head. 56. Paratype spiracular plate from sting apparatus
(Chiapas, Mex). 57. Nontype queen profile (Parque Tayrona, Col.). Figs. 58-60. Rogeria alzatei. 58. Holotype profile, dorsal head;
dorsal waist. 59. Nontype profile (Pto. Maldonado, Lake Sandoval, Peru). 60. Nontype profile (R. Don Diego, Col.). Figs. 61-
62. Rogeria scobinata. 61. Holotype profile; dorsal head. 62. Paratype queen profile (holotype loc). Same scale for all external
views.
K4
Journal of Hymenoptera Research
tribrocca
carinata
nevadensis
leptonana
Figs. 63-70. Fig. 63. Rogeria tribrocca holotype profile; dorsal head. Fig. 64. Rogena carinata holotype profile; dorsal head. Fig.
65. Rogeria nevadensis holotype profile, dorsal head. Figs. 66-70. Rogeria leptonana. 66. Holotype profile; dorsal head; dorsal
postpetiole. 67. Nontype profile (nr. Nueva California, Pan.). 68. Nontype queen profile (Parque Tayrona, Col.). 69. Male
(Cerro Pico Blanco, C. R). 70. Male genitalia (same loc). Same scale for all external views.
Volume 3. 1 994
85
micromma
1 ^Q
curvipubens
cuneola
Figs. 71-78. Fig. 71. Rogeria micromma holotype profile. Figs. 72-73. Rogeria minima holotype. 72. Lateral mesosoma, petiole;
ventral postpetiole, gaster. 73. Erect hair. Figs. 74-76. Rogeria curvipubens. 74. Nontype profile; dorsal head; dorsal waist (BCI,
Pan). 75. Slide-mounted worker lateral petiole (Parque Tayrona, Col.). 76. Queen profile (R. Don Diego, Col). Figs. 77-78.
Rogeria cuneola. 77. Holotype profile. 78. Slide-mounted worker lateral petiole (El Palmar, Mex.). Same scale for all figures
except 73, 75, and 78.
86
Journal of Hymenoptera Research
79/
foreli
1.0 mm
bruchi
Figs. 79-82. Figs. 79-81. Rogerm foreli. 79. Holotype profile. 80. Nontype mesosoma dorsum profile showing strongest metanotal
groove (Cerro Azul, Pan.) 81. Lateral waist (Don Diego vie, Col). Fig. 82. Rogeria bruchi holotype profile; dorsal mandibles
and clypeus. All to same scale
Volume 3. 1 994
87
;W3fe|
86
tonduzi
N«
I
l» :""'
lirata
Figs. 83-90. Figs. 83-84. Rogeria blanda (L. Sandoval, Peru). 83. Lateral mesosoma, waist. 84. Dorsal head, mesosoma, petiole.
Figs. 85-86. Rogeria tonduzi (La Selva Station, C. R.). 85. Lateral mesosoma and waist. 86. Dorsal mesosoma. Figs. 87-88. Rogerw
germaini (Alto Parana Prov., Para.). 87. Lateral mesosoma. 88. Dorsal mesosoma, posterior head. Figs. 89-90. Rogeria lirata
paratype. 89. Lateral mesosoma. 90. Dorsal mesosoma. Scale lines = 0.10mm.
88
mm'*/®
'''■*IW
mm
Journal of Hymenoptera Research
creightoni
|98 |fc inotabilis
Figs. 91-98. Figs. 91-92. Rogeria belti (nr. El Bosque, Mex.). 91. Lateral head (part), mesosoma. 92. Dorsal mesosoma and waist
of a. smaller worker. Fig. 93. Rogeria brunnea lateral mesosoma (Blanco's Woods, Soledad, Cuba). Fig. 94. Rogeria nevadensis
paratype lateral mesosoma. Figs. 95-96. Rogeria creightoni (Belmopan, Belize). 95. Lateral mesosoma, petiole. 96. Dorsal
mesosoma. Figs. 97-98. Rogeria innotabilis paratype. 97. Lateral head, mesosoma. 98. Posterior head, dorsal mesosoma. Scale
lines = 0.10mm.
Volume 3, 1994
89
Figs. 99-105. Fig. 99. Rogeria alzatei paratype posterodorsal head, dorsal pronotum. Fig. 100. Rogeria scobinata paratype
posterodorsal head. Figs. 101-102. Rogeria curvipubens (R. Don Diego vie, Col). 101. Posterodorsal head. 102. Dorsal
mesosoma. Fig. 103. Rogeria cuneola paratype dorsal mesosoma. Figs. 104-105. Rogeria foreli (Parque Tayrona, Col.). 104. Lateral
waist showing prominent anterior lip of postpetiolar sternum of some specimens. 105. Posterior head, dorsal mesosoma. Scale
lines = 0.10mm.
J. HYM. RES.
Vol. 3, 1994, pp. 91-105
The Genera of Bethylinae (Hymenoptera: Bethylidae)
Andrew Polaszek and Karl V. Krombein
(AP) Department of Entomology, Wageningen Agricultural University, The Netherlands
International Institute of Entomology, c/o The Natural History Museum, London, UK
(KVK) Department of Entomology, Smithsonian Institution, Washington, D. C. USA.
Abstract- — The taxonomic status of the genera comprising the bethylid subfamily Bethylinae is reassessed using
computerised phylogenetic analysis. From this analysis seven genera are considered valid, and the following are
synonymised: Trissomalus Kieffer 1905 with Odontqjyns Kieffer 1904; and Anoxus Thomson 1862 with Bethylus Latreille
1802. Several species are transferred generically, and several new combinations are presented. The distribution and
biology of the Bethylinae are summarised.
INTRODUCTION
The aculeate family Bethylidae (Chrysidoidea)
is a moderately large family of about 1,900 de-
scribed species with probably at least as many
undescribed. Bethylids are mainly gregarious
ectoparasitoids, the Bethylinae mostly develop-
ing on larval Lepidoptera, the remainder mostly
on Coleoptera, although the hosts are known for
only a small proportion of the family. Bethylids
are frequently encountered as parasitoids of crop
pests, especially in tropical areas, and several spe-
cies have been used in attempted biological con-
trol (Gordh & Evans 1976; Hempel 1934). Success-
ful identification of Bethylidae is therefore impor-
tant for many economic entomologists, but has
been extremely difficult because the most recent
revision was published eighty years ago (Kieffer
1914) and contains many errors. Even at the ge-
neric level identification can be very difficult, par-
ticularly for the Palaeotropical species. For the
Nearctic and Neotropical species this problem has
largely been alleviated by the work of Evans (1964,
1978).
During our independent studies on African
and Asian Bethylinae we have each discovered
related undescribed species that donotfitKieffer's
(1914) concepts of the bethyline genera, but which
we do not regard as meriting description in new
genera. Kieffer's and subsequent authors' con-
cepts of the bethyline genera needed modification
to accommodate the degree of morphological varia-
tion which we had discovered. Computerised
phylogenetic analysis was selected as the most
objective method of assessing character states
within the bethyline genera, while providing a
more stable classification of the subfamily. The
following study was carried out primarily to set
new limits to some of these genera, and to facilitate
their identification.
Here we address the internal phylogeny of the
Bethylinae. In comparison to the other bethylid
subfamilies, the Bethylinae have been little af-
fected at the generic level since Kieffer's (1914)
revision. Only one genus, Neoclystopsenella Kurian
(1955), was assigned to the Bethylinae since
Kieffer's work. Neoclystopsenella was synonymised
with Tapinoma (Formicidae) by Brown (1987), but
mistakenly retained by Gordh & Moczar (1990) in
the Bethylidae. Gordh & Moczar (1990) also mis-
takenly transferred Odontepyris Kieffer to Epyrinae,
thus assigning a total of eight genera to the
Bethylinae (Table 1). When attempting to identify
Bethylinae genera using Kieffer's (1914) key, the
primary source of confusion is his treatment of the
genera Goniozus Fbrster, Parasierola Cameron and
Perisierola Kieffer. The latter two genera have since
correctly been synonymised with Goniozus (Evans
92
Journal of Hymenoptera Research
1978), but from our studies it is clear that Kieffer
intended something different from what he inad-
vertently achieved when he keyed and diagnosed
these genera in his revision. Goniozus (sensu Evans
1978) contains species that either possess, or do
not possess, a closed discoidal cell (areola, areolet)
(Fig. 1). Kieffer assigned those species with a dis-
coidal cell to Perisierola, and those without one to
Goniozus. Kieffer (1907, 1911, 1914), Brues (1907a
1907b) and Muesebeck (1940) all mistakenly se-
lected Parasierola Cameron (1883) to accommodate
species with both a closed discoidal cell and with
one or more longitudinal carinae on the
propodeum, this lattercharacter being absent from
Kieffer's concepts of the other two genera. Fur-
thermore, Kieffer (1914) transposed his concepts
of Parasierola and Perisierola when going from his
key to genera (1914:238) to his generic diagnoses
(1914:533, 542). Thus began 80 years of confusion
surrounding these bethyline genera.
METHODS
Selection of taxa
To clarify the status of the genera of Bethy linae
we analysed 11 taxa of Bethylinae for 22 morpho-
logical characters using the parsimony programme
'Hennig86' (Farris 1988). To polarise characters
the genus Lytopsenella Kieffer was selected as the
outgroup. Lytopsenella possesses all the characters
common to all the remaining bethyline genera in
their hypothetical plesiomorphic conditions (see
character selection). Characters that are prone to
reduction (e.g., number of antennal segments,
maxillary and labial palp segments, and wing
venation) are found at their maxima within
Lytopsenella. Lytopsenella has previously been cho-
sen as a basal group, not just for Bethylinae but for
Bethylidae as a whole (Evans 1964; Sorg 1988).
Representatives of each of the currently valid
bethylid genera were included in the analysis. In
cases of existing doubt or controversy surround-
ing the limits of some genera, type species of both
current genera and formerly recognised genera
were examined. Particular attention was paid to
previous authors' concepts of Goniozus,
Odontepyris, Parasierola, Perisierola and Trissomalus.
Three species of Goniozus (in the broad sense, i. e.
that of Evans 1978) were selected to cover the
range of known variation in propodeal and wing
venation characters which are important for de-
ducing the phylogeny of the subfamily. Current
interpretations of the genera Anoxus and Bethylus
differ from each other only in whether the eyes are
setose or not. This is a character that we have
observed to vary intragenerically, so only one
representative species of Bethylus was included in
the analysis. The taxa selected are given in the data
matrix below (Table 2) in the generic combinations
which have resulted from this study. Former ge-
neric combinations can be found in the treatments
of Bethylus and Odontepyris (see below).
Selection of Characters
We consider the following to represent the
ground plan characters of the subfamily Bethylinae.
This character list is based partly on the work of
Evans (1964) and of Sorg (1988) but largely on our
own independent assessments.
Ground plan characters within Bethylinae
Antennae 13-segmented; clypeus with a well-
developed keel, and frontal streak present; maxil-
lary palps 6-segmented; labial palps 3-segmented;
notauli and parapsidal lines present; scutellum
flat, with two small grooves at its proximal corners
(Fig. 2); propodeum without carinae (but see be-
low); fore femora expanded; prostigma absent;
discoidal cell (areolet) present, submarginal and
marginal cells present; subdiscoidal cell absent
(=discoidal cell of Evans 1964); sternum of petiole
with a complete keel, sexual dimorphism limited
to genital characters and head shape.
Characters Analysed
The following characters include all of those
which have been used previously for the discrimi-
nation of genera within the Bethylinae (except eye
setation, see above) as well as some which have
not been used previously. We generally agree
with Sorg (1988) concerning both the selection and
polarization of characters. However, we disagree
with Sorg's polarization of the scutellar foveae
(character 6, below; Figs. 3, 4). Sorg considers that
the occurence of scutellar foveae in the
Embolemidae (Sorg 1988: p 30) suggests
plesiomorphy. The probability is that, at least in
the Bethylinae, they are represented in their
Volume 3, 1 994
93
Table 1: Genera of Bethylinae
Kieffer (1914)
Gordh & Moczar
(1990)
This paper
(Bethylini)
(Bethylinae)
(Bethylinae)
Anoxus
Anoxus
Bethylus
Bethylus
Bethylus
Bethylus
Clyslopsenella
not mentioned
Scolebythidae (Evans, 1963)
Digoniozus
Goniozus
Goniozus
Eupsenella
Eupsenella
Eupsenella
Gomozus
Gomozus
Gomozus
Kathepyris
Pristocerinae
Pristocerinae (Evans 1964)
Lytopsenella
Lytopsenella
Lytopsenella
Neoclystopsenella (incertae sedis)
Formicidae (Brown 1987)
Odontepyris (Bethylini)
Odontepyris (Epy
rinae)
Odontepyris (Bethylinae)
Parasierola
Goniozus
Goniozus
Perisierola
Goniozus
Gomozus
Progoniozus
Goniozus
Goniozus
Prosierola
Prosierola
Prosierola
Swrola
Sierola
Sierola
Trissomalus
Trissomalus
Odontepyris
plesiomorphic condition in Lytopsenella.
Assumed primitive (plesiomorphic) charac-
ters are coded '0' with '1', '2' representing as-
sumed derived (apomorphic) states.
1. Number of antennal segments: The
plesiomorphic condition in the Bethylinae is
possession of 13-segmented antennae. In
Bethylus (including Anoxus) a reduction to
12-segmented antennae has occurred. 0= an-
tennae 13-segmented; 1= antennae 12-seg-
mented.
2. Number of labial palp segments: The
plesiomorphic condition is possession of 3-
segmented labial palps. In Bethylus and Sierola
the labial palps are 2-segmented. 0= labial
palps 3-segmented; 1= labial palps 2-seg-
mented.
3. Number of maxillary palp segments: The
plesiomorphic condition is possession of 6-
segmented maxillary palps, found in both
Eupsenella and Lytopsenella. In the remaining
taxa the number is reduced to five, except
Sierola which has 4-segmented maxillary
palps. 0= maxillary palps 6-segmented; 1 =
maxillary palps 5-segmented; 2= maxillary
palps 4-segmented.
Presence of an unsculptured streak fron-
tally: In many bethyline taxa a narrow
impunctate, sometimes shiny, streak extends
from the proximal end of the clypeal carina to
the frontal ocellus (Fig. 12, see also Sorg 1988:
121, Fig. 33d, for its presence in Epyrinae:
Chilepyris). In Bethylus (including Anoxus),
Goniozus and Sierola the sculpturing of the
head is reduced (i. e. smooth rather than
strongly rugose) and this streak is absent. 0=
frontal streak present; 1= absent.
Presence of notauli: The notauli, a pair of
longitudinal furrows on the mesoscutum,
occur throughout the Hymenoptera, and are
often reduced. In the Bethylinae they are
present only in the basal taxa, Eupsenella (Figs.
2, 3) and Lytopsenella. In all remaining
Bethylinae the notauli are absent. The notauli
should not be confused with the parapsidal
lines, which are an additional pair of shallow
grooves lateral to the position of the notauli
(Menke 1993) (Fig. 2). 0= notauli present; 1=
absent.
Presence of scutellar grooves or pits: In the
basal taxa Eupsenella and Lytopsenella the
scutellum has a pair of grooves at its anterior
corners (Figs. 2, 3, 5). In several bethyline taxa
94
Journal of Hymenoptera Research
pterostigma marginal cell
10.
11.
12.
position of prostigma
median cell
submedian cell
subdiscoidal cell
(absent in Bethylinae)
Fig. 1. Bethylidae: terminology of wing venation
these have apparently become enlarged to
form pits, often connected by a broad trans-
verse groove (Figs. 4, 6, 7). 0= scutellar pits or
foveae present as grooves; 1= scutellar foveae
enlarged.
Expansion of mesopleuron: Kieffer (1905)
characterised his genus Odontepyris by the
possession of dentate processes on the
mesopleura (Figs. 6, 14). Although an
apomorphic character in one lineage, it shows
varying degrees of development in related
taxa. 0= mesopleuron not expanded to form
a dentate process; 1= mesopleuron moder- 13.
ately expanded; 2=mesopleura with dentate
processes.
Presence of posterior transverse propodeal
carina (Figs. 2, 6, 7, 9, 10): 0= posterior trans-
verse propodeal carina present; 1= posterior
transverse propodeal carina absent.
Presence of median longitudinal propodeal
carina (Figs. 2, 5, 6, 8, 9): 0= median longitu-
dinal propodeal carina present; 1= median
longitudinal propodeal carina absent.
Presence of discal longitudinal propodeal 14.
carinae (Figs. 2, 5, 6, 7, 8, 9, 10): 0= discal
longitudinal propodeal carinae present; 1=
discal longitudinal propodeal carinae absent.
Presence of median propodeal pits or foveae
(Figs. 4, 7, 10): The presence of these struc-
tures is characteristic of the genus Prosierola.
0= median propodeal pits absent; 1= median
propodeal pits present. 15.
Development of a smooth, triangular area
on the dorsal propodeum: In several
bethyline taxa, particularly Goniozus and
Prosierola, the proximal dorsum of the
propodeum is characterised by a smooth,
unsculptured triangular area which shows
modifications in several taxa (Figs. 7, 10, 11).
It is absent in the basal groups. The develop-
ment of this character in some bethyline lin-
eages should not be confused with the
"propodeal triangle" in Apidae and
Sphecidae (Brothers 1976). 0= smooth, trian-
gular area absent from dorsal propodeum
(Figs. 2, 5, 6, 8, 9); 1= smooth, triangular area
present on dorsal propodeum (Figs. 7, 10,
11).
Petiole ventrally keeled: To our knowledge,
this character has not been used previously
in defining the genera of Bethylinae, but is
important for separating Goniozus from su-
perficially similar taxa in Odontepyris (see
key to genera, below). In most Bethylinae, a
complete longitudinal keel is present on the
ventral petiole (Fig. 13). This keel is reduced
in several taxa (e. g. Fig. 15). 0= petiole with
a complete ventral keel; 1= petiolar keel re-
duced; 2= petiolar keel absent.
Expansion of the fore femora: Expansion of
the fore femora for fossorial use is often en-
countered among the Bethylidae, and also
occurs in related chrysidoid taxa and other
aculeates. Other modifications of the fore
femora are frequent in the Aculeata. 0= fore
femora strongly expanded; 1= fore femora
less strongly expanded, half as wide as long.
Development of the prostigma: The expan-
sion of the junction of the subcostal and basal
veins into a secondary pterostigma is charac-
teristic of the genus Goniozus (Figs. 20, 25),
but also occurs elsewhere within the subfam-
Volume 3. 1994
Table 2. Character matrix:
95
Characters
1
2
3
4
5
6
7
8
9
10 11
12 13
14
15 16 17 18 19 20 21
22
Lytopsenella sp.
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Eupsenella sp.
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
1
0
0
0
0
Sierola sp.
0
1
2
1
1
0
0
0
1
0
0
0
0
0
1
0
0
0
0
1
0
1
Bethylus cq>halotes
1
1
1
1
1
0
0
0
0
0
0
0
2
0
0
0
0
0
1
1
1
1
Odontepyris flavinervis
0
0
1
0
1
1
2
1
1
1
0
0
0
1
0
1
1
0
0
1
1
0
Odontepyns transvaalensis
0
0
1
0
1
1
1
1
1
1
0
0
0
0
0
1
1
0
1
1
1
0
Odontepyns xanthoneurus
0
0
1
0
1
1
1
1
1
1
0
0
0
1
1
1
1
0
0
1
1
0
Prosierola sp.
0
0
1
0
1
1
1
1
0
1
1
1
0
0
0
1
1
0
0
1
1
0
Conwzus sp.
0
0
1
1
1
0
0
1
0
0
0
1
1
0
1
0
0
0
0
1
1
0
Goniozus gallicus
0
0
1
1
1
0
0
0
0
0
0
1
1
0
1
0
0
0
0
1
1
0
Gomozus mdicus
0
0
1
1
1
0
0
0
0
0
0
1
1
0
1
0
0
0
1
1
1
0
ily. Its presence outside Goniozus has previ-
ously led to the generic misplacement of
certain species. 0= prostigma absent; 1= 19.
prostigma present.
16. Length of rs+m: In the basal taxa, fore wing
vein rs+m forms the upper distal part of the
discoidal cell (Fig. 1). In all Goniozus species
it is present and well-developed, despite the
fact that in Goniozus (sensu stricto) the discoi-
dal cell is open. The genus Odontepyris is here
characterised partly by the reduction of this
vein (Figs. 22-24). The absence of this vein in
Bethylus is clearly convergent with its reduc- 20.
tion in Odontepyris (see below). 0= rs+m long;
as long as, or longer than, rs (Figs. 16-18, 20,
25); 1= rs+m shorter than rs, or absent (Figs.
19, 22-24).
17. Length of rs: The length of rs relative to m
(Fig. 1) characterises the Prosierola / 21.
Odontepyris clade, in which rs has become
elongated. 0= rs equal to or shorter than m
(Figs. 16-20, 25); 1= rs clearly longer than m
(Figs. 21-24).
18. Length of marginal (radial) cell: In Eupsenella, 22.
one of the basal taxa, the marginal cell is
characteristically shortened (Fig. 17). In the
remaining taxa which have a closed marginal
cell {Lytopsenella, Sierola), the marginal cell is
of normal dimensions. For those taxa in which
the marginal cell is open, it is necessary to
calculate the size of a hypothetical closed
marginal cell by the length of the radial vein
(2r - rs of Sorg 1988; r + Rs of Evans 1964). 0=
marginal cell long (Fig. 16); 1= marginal cell
shortened (Fig. 17).
Presence of discoidal cell (areolet): Within
all Bethylinae the discoidal cell appears to be
homologous, being plesiomorphically
present. In Goniozus it is either clearly present
or absent, whereas in Odontepyris various
intermediate degrees of reduction of the dis-
coidal cell are apparent (c. f. Figs. 22, 23, 24).
0= discoidal cell present (Figs. 16, 17, 18, 21,
22, 23, 25); 1= discoidal cell absent (Figs. 19,
20, 24).
Presence of submarginal cell In the
Bethylinae, the submarginal cell is present
and complete only in Eupsenella and
Lytopsenella. It is absent in all remaining taxa.
0= submarginal cell present (Figs. 1, 16, 17);
1= submarginal cell absent (Figs. 18-25).
Presence of closed marginal cell: In the basal
taxa, and in Sierola, the marginal cell is closed,
the plesiomorphic condition for Bethylinae.
0= marginal cell closed (Figs. 1, 16-18); 1 =
marginal cell open (Figs. 19-25).
Angle of radial vein: The radial vein (Fig. 1;
= vein 2r - rs of Sorg 1988; vein r + Rs of Evans
1964) is distinctly angled in Bethylus and
Sierola (Figs. 18, 19), genera respectively with
an open and a closed marginal cell. The angle
of the radial vein is thus independent of the
closure of the marginal cell. 0= radial vein
without a sharp angle; 1 = radial vein sharply
angled.
96
Journal of Hymenoptera Research
Fig. 2. Bethylidae: terminology of mesosoma.
The character matrix was analysed using the
'implicit enumeration' option of Hennig86, the
procedure which computes the maximum pos-
sible number of cladograms, or 'trees'. Multistate
characters (3: number of maxillary palps; 5: pres-
ence of notauli; 7: expansion of the mesopleuron;
13: reduction of the petiolar keel) were treated
both as unordered (non-additive) and ordered
(additive).
RESULTS
A single cladogram (tree) resulted from the
analysis, having the following characteristics:
length 35 steps, consistency index 71 and retention
index 80 (Fig. 26). Ordering the data had no effect
on the topology of the tree, only increasing its
length by one step, and reducing the consistency
index by two.
Characters Supporting Monophyly
of Bethyline Genera / Clades
1. (node A, Fig. 26): {(Prosierola + Odontepyris) +
((Sierola + Bethylus) + Goniozus))).
Monophyly of this clade is supported by the
following synapomorphies: 3: reduction of
number of maxillary palp segments from six
to five; 5: loss of notauli; 20: loss of submar-
ginal cell; 21: marginal cell open (reversed in
Sierola).
2. (node B, Fig. 26): (Prosierola + Odontepyris)
Monophyly of this clade is supported by the
following synapomorphies: 6: expansion of
the scutellar pits; 7: expansion of the
mesopleura; 8: presence of a posterior trans-
verse propodeal carina; 10: presence of discal
carinae; 16: rs+m shorter than rs; 17: rs longer
than m.
3. (node C, Fig. 26): Odontepyris
Volume 3. 1994
97
Figs. 3-4. 3, Eupsenella sp.: detail of mesosoma. 4, Prosierola sp.: scutellum and propodeum.
Monophyly of Odon tepyris is supported by the
following synapomorphy: 9: presence of a
median propodeal carina.
4. (node D, Fig. 26): Prosierola
Monophyly of Prosierola is supported mostly
by convergent characters, except for the
autapomorphic character (11) presence of
propodeal pits.
5. (node E, Fig. 26): ((Sierola + Bethylus) + Goniozns))
Monophyly of this clade is supported by a
single synapomorphy: 4: loss of the frontal
streak.
6. (node F, Fig. 26): Goniozus
Monophyly of Goniozus is supported by a
single synapomorphy: 13: reduction of the
petiolar keel. Character 12, presence of a
smooth triangular area on the propodeum is a
synapomorphy for Goniozus which occurs as a
homoplasy in Prosierola.
7. (node G, Fig. 26): (Sierola + Bethylus)
Monophyly of this clade is supported by the
following synapomorphies: 2: reduction of
the number of labial palp segments from three
to two; 22: radial vein sharply angled.
8. (node H, Fig. 26): Sierola
Monophyly of Sierola is supported by 3: max-
illary palps 4-segmented, as well as by the
closed marginal cell (21 ) which is here a rever-
sal.
9. (node I, Fig. 26): Bethylus
Monophyly of Bethylus is supported by the
following synapomorphies: 1: reduction of
the number of antennal segments from 13 to
12; 13: petiolar keel absent.
DISCUSSION
The analysis supports monophyly of our modi-
fied concept of Odontepyris, including Trissomalus
Kieffer, and Parasierola Cameron sensu Kieffer
(1914), not Cameron (1883). The currently accepted
definitions of most of the remaining genera are
also supported by the analysis.
Trissomalus (Kieffer 1905) was characterised
in a key, but not formally diagnosed until Kieffer's
(1914) generic revision. Odontepyris (Kieffer 1904)
was characterised mainly by the possession of
dentate processes on the enlarged mesopleura
98
Journal of Hymenoptera Research
Figs. 5-7. 5. Eupsenella sp.: mesosoma. 6, Odontepyris sp.: mesosoma. 7, Prcsierola sp.: mesosoma.
(Fig. 14). We have examined many described and
undescribed species belonging to the Odontepyris
I Trissomalus group, and conclude that the degree
of variation in the development of the mesopleural
processes, coupled with a successive reduction in
the size of the discoidal cell (Figs. 22-24) suggests
the group should be treated as a single genus, for
which Odontepyris Kieffer is the oldest available
name. The alternative would be to treat as new
genera all intermediate species or species-groups.
Examination olAnoxus specimens shows that
the genus differs from Bethylus only in the extent of
setation of the eyes, other characters are identical
in the two genera. Furthermore in some Bethylus
species some setation can be found on the eyes (e.
g. B. amoenus Fouts). Since we know that the de-
gree of eye setation is a character that clearly varies
interspecifically within other bethyline genera,
we do not see how retaining Anoxns as a distinct
genus can be justified, and it is therefore
synonymised below.
GENERIC SYNONYMY
ODONTEPYRIS Kieffer
Odontepyris Kieffer 1904: 378. Type species Odontepyris
flavinervis Kieffer 1904 by original designation. Holo-
type female: SUMATRA: Pangherang-Pisang x. 90 e iii.
91 (E. Modigliani) (MCSN) [examined].
Trissomalus Kieffer 1905: 105. Type species Goniozus
transvaalensis Du Buysson, 1888: 354 by subsequent des-
ignation. Holotype female: [SOUTH AFRICA:]
Transvaal, Hamman's Kraal 1893 (E. Simon) (MNHN)
[examined] syn. n.
Diagnosis. — Antennae 13-segmented. Palpal
formula 5:3. Frontal streak present. Notauli ab-
sent. Scutellar foveae well developed. Mesopleuron
expanded, sometimes developed into a dentate
prominence (Fig. 14). Median, discal and posterior
transverse carinae present on propodeum, the
longitudinal carinae occasionally reduced. Petiole
with a complete ventral keel. Prostigma present or
absent, rs + m reduced, rs elongate. Discoidal cell
present, reduced or absent. Marginal and submar-
ginal cells absent. Radial vein smoothly curved.
Included species. — Odontepyris argyriae Kurian;
O. batrae Kurian; O. cameroni (Kieffer) (comb. n.
from Trissomalus, holotype examined); O. cirphi
Kurian; O. erucarum (Szelenyi) (comb. n. from
Parasierola); O. flavinervis (Kieffer) (holotype ex-
amined); O. fuscicrus (Kieffer) (comb. n. from
Trissomalus, holotype examined); O. hypsipylae
(Kurian) (comb. n. from Goniozus); O. indicus
Kurian (comb. n. from Trissomalus); O. moldavicus
(Nagy) (comb. n. from Prosierola); O. peringueyi
(Kieffer) (comb. n. from Trissomalus); O.
quadrifoveatus (Muesebeck) (comb. n. from
Parasierola, holotype examined); O. ruficeps Kieffer;
Volume 3, 1 994
99
KEY TO GENERA OF BETHYLEMAE
1 Closed submarginal cell present (Figs. 1, 16, 17) 2
— Closed submarginal cell absent (Figs. 18-25) 3
2 Marginal cell elongate, larger than submarginal (Figs. 1, 16) Lytopsenella
— Marginal cell short, smaller than submarginal (Fig. 17) Eupsenella
3 Marginal cell closed (Figs. 1, 16-18) Sierola
— Marginal cell open (Figs. 19-25) 4
4 Antennae 12-segmented. Wing venation as in Fig. 19; radial vein at apex turned abruptly upward, but not
reaching wing margin. Fore wing with rs+m always absent Bethylus
— Antennae 13-segmented. Wing venation different; radial vein shorter and evenly curved towards wing
apex (Figs. 20-25). Fore wing with rs+m present or absent 5
5 Propodeum without well-developed lateral carinae (fig 11). Scutellum without large foveae, with small
grooves (as in Fig. 3). Petiole ventrally with a reduced, forked keel (Fig. 15) Goniozus
— Propodeum with well-developed lateral carinae (Figs. 9, 10). Scutellum with large foveae (Figs. 6, 7). Petiole
ventrally with a complete keel (Fig. 13) 6
6 Median longitudinal propodeal carina present (Fig. 9). Propodeum without median foveae
Odontepyris
— Median longitudinal propodeal carina absent (Fig. 10). Propodeum with median foveae (Fig. 10)
Prosierola
O. transvaalensis (De Buysson) (comb. n. from
Goniozus, holotype examined); O. zvaterhousei
(Kieffer) (comb. n. from Parasierola, holotype ex-
amined); O. xanthoneurus (Kieffer) (comb. n. from
Parasierola, holotype examined).
Comments. — Tryapitsin (1978) and Terayama
(1990) mistakenly included the Neotropical
bethyline genus Prosierola in their respective keys
to Russian and Japanese bethylid genera. In each
case the error was due to misidentification of an
Odontepyris species.
BETHYLUS Latreille
Bethylus Latreille, 1802: 315. Type species Omalus fusacorms
Junne 1807: 301 by subsequent designation (Interna-
tional Commision on Zoological Nomenclature, opin-
ion 153). Holotype 'female [not examined, ?lost).
Anoxus Thomson 1862: 451. Type species Anoxus boops
Thomson 1862: 452. Monotypic. Lectotype female
(Naturhistonska Rijksmuseet, Stockholm) [not exam-
ined] syn. n.
For full synonymy see Gordh & Moczar 1990.
Diagnosis. — Antennae 12-segmented. Palpal
formula 5:2. Frontal streak absent. Eyes with or
without long hairs. Notauli absent. Scutellar foveae
narrow, groove-like. Mesopleuron not expanded.
Propodeum without carinae. Petiole with the ven-
tral keel absent. Prostigma and discoidal cells
absent. Marginal and submarginal cells absent.
Radial vein usually sharply angled.
Species transferred from Anoxus
B boops (Thomson) comb. n. ; B. coniceps (Kieffer) comb. n. ,
B pilosus (Kieffer) comb. n. .
Comments. — It might appear questionable to
synonymise/iHOXKS with Bethylus without having
examined the type species of either genus. How-
ever, there has not been any controversy sur-
rounding these genera since Kieffer's (1914) ge-
neric revision, and we have examined sufficient
material conforming to the original descriptions
and Kieffer's interpretations of Anoxus and
Bethylus.
DISTRIBUTION OF BETHYLINAE
The most primitive bethyline, Lytopsenella, is
known only from two extant species, both from
Chile. Three fossil species are known from Baltic
amber (Evans 1964). Eupsenella is known so far
only from Australia.
Odontepyris is predominantly Palaeotropical,
but its distribution spans southeastern Europe to
South Africa and northern Queensland. The small
genus Prosierola is primarily Neotropical although
one species ranges into the extreme southern
United States; a fossil species from Baltic Amber is
certainly incorrectly assigned to Prosierola (Brues
1933). Goniozus, with about 150 described species,
100
Journal of Hymenoptera Research
■m
Figs. 8-11. 8, Eupsenella sp.: propodeum. 9, Odontepyris sp.: propodeum. 10, Prosierola sp.: prapodeum. 11, Goniozus sp.:
propodeum.
Volume 3. 1 994
101
Figs. 12-15. 12, CWoHf£7n/nssp.:head. 13, Odontepyns sp.: ventral
showing dentate process. 15, Goniozus sp.: ventral petiole and
is cosmopolitan although only one species is known
from the Pacific Islands. Approximately three
dozen species each were described from the Ori-
ental, Nearctic and Neotropical regions, and about
a dozen each from the Palaearctic, Ethiopian and
Australian regions. A number of species have
been introduced into other countries as biological
control agents. A few fossil species have been
petiole and sternite 2. 14, Odontepyris sp.: side of mesopleuron
sternite 2.
described from Baltic and Dominican amber.
Sierola contains almost 200 described species,
mostly from Hawaii, with three endemic species
in Australia. The genus underwent tremendous
speciation in Hawaii, and a secondary small spe-
ciation in the Marquesas Islands. Evans (1978)
suggested that the single Calif ornian species might
be introduced, and this is possibly also true of the
102
Journal of Hymenoptera Research
17
19
25
Figs. 16-25. Fore wings of various Bethylidae: 16, Lytopsenella sp. 17, Eupsenella sp. ; 18, Sierola sp. 19, Bethylus sp. 20. Coniozus
(sensu stricto) 21, Prosierola sp. 22-24, Odonttpyris spp. 25, Goniozus ("Parasierola") sp.
Volume 3, 1 994
103
26
F
■
H
G
■
■
I
Lytopsenella sp.
Eupsenella sp.
Odontepyris flavinervis
Odontepyris xanthoneurus
Odontepyris transvaalensis
Prosierola sp.
Goniozus sp.
Goniozus gallicus
Goniozus indicus
Sierola sp.
Bethylus cephalotes
Fig. 26. Cladogram of the Bethylinae.
single Chinese species. A fossil species has been
described from Baltic amber (Sorg 1988).
Bethylus, containing some 30 described spe-
cies, was previously considered as Holarctic, and
predominantly Palaearctic, only two species be-
ing definitely known from the Nearctic. Recently,
the first author examined several specimens of
Bethylus from South Africa. Homalus lamplipennis
Motschulsky was erroneously placed by Gordh &
Moczar (1990) in Bethylus; Krombein (1987) trans-
ferred it to Holepyris.
BIOLOGY OF BETHYLINAE
Bethylinae are characterized in part by para-
sitizing lepidopterous larvae, whereas members
of the other subfamilies attack primarily coleopter-
ous larvae. A few exceptions to these host prefer-
ences are documented in Bethylinae, Epyrinae
and Pristocerinae, but the few known hosts of
Mesitiinae are case-bearing coleopterous larvae.
The preferred hosts within the Bethylinae are
mostly microlepidopterous as listed by Evans
(1978) and Gordh & Moczar (1990). However,
there are a few anomalous or questionable host
records. It should be noted that, under experimen-
tal conditions, females of the epyrine Sclerodermus
immigrans Bridwell (1920) will successfully para-
sitize a variety of insect larvae other than their
normal coleopterous host. These include other
Hawaiian beetles, bees, wasps, ants, and even
braconid and chalcidoid parasites of the normal
host larva. Bridwell also reared S. immigrans from
normal workers, dealate adults, and nymphs of
the termite Neotermes castaneus (Burmeister), al-
though not from the soldiers. Perhaps some of the
questionable records below may reflect this capa-
bility in other bethylids.
Evans (1964) reported that a specimen of
Lytopsenella herbsti (Kieffer) was labelled as attack-
ing an adult cantharid beetle. This record is dubi-
ous, inasmuch as hosts of all other bethylids are
larvae. The only other records of bethylines attack-
ing a host stage other than the larva are those of
Nagy (1976). He reported Odonte^n/ris moldavicus
(Nagy) (as Prosierola moldavica) as reared from
"pupae of a nocruid moth" and Goniozus plugarui
Nagy attacking a lepidopterous pupa. These
104
Journal of Hymenoptera Research
records of bethylids reared from pupae are dubi-
ous, and the most probable explanation is that
"pupal parasites" were reared from larvae which
had spun cocoons but had not yet pupated.
Evans (1962) reported that Bethylus amoenus
Fouts was reared from an olethreutid moth and a
nitidulid beetle. The latter record was later omit-
ted from Evans' (1978) list of host records, and to
us seems doubtful. A species tentatively identified
as Goniozus gestroi (Kieffer) was reported by
Richards (1955) as being reared from larvae of the
anobiid beetle, Lasioderma. The rearing was not
questioned, only the specific identity of the
Goniozus, and we therefore accept this record as
authentic. Goniozus morindae Kurian (1952), de-
scribed from a single male, was reared from the
gall of acecidomyiid fly , Asphondyla morindae Mani
in flowers ofMorinda tinctoria. The record is anoma-
lous, and is the first for a dipteran as a host of a
bethylid. The most recent authentic record of a
hymenopterous host for a bethy line is that of Melo
and Evans (1992) who reported Goniozus
microstigmi Evans as being reared from a brood
cell of the sphecid wasp Micros tigmus xylicola Melo,
a predator of nymphal Thysanoptera. The nests
were in abandoned beetle galleries in beams of an
exposed roof, and a dozen G. microstigmi females
were collected while walking near and entering
nests of M. xylicola and M. similis Melo.
So far as is known, all Bethylinae are gregari-
ous parasitoids, laying a clutch of eggs on each
host larva. The host is stung, sometimes repeat-
edly, behind the gula. Paralysis of the host larva
may be temporary or permanent, depending upon
the species of wasp. The number of eggs per clutch
varies according to the size of the host, as well as
interspecifically (Gordh & Evans 1976), ranging
from one or two to as many as 40. Placement of the
eggs depends upon the species, eggs being depos-
ited either intersegmentally or longitudinally, and
either dorsally, ventrally or laterally. Eggs hatch
about two days after oviposition, and the larvae
complete feeding in 2 to 5 days. Each larva spins a
cocoon on the substrate near the host remains. The
pupal period varies from 8-14 days, depending
upon the species and ambient temperature. Males
emerge a day or two before females are ready to
eclose; they have been observed chewing into a
cocoon containing a female and mating with her
while she is still teneral. The progeny from a clutch
usually consists of a single male and a number of
females, and sibling mating is common (Hardy
1992). Maternal care of the larvae has been ob-
served in Bethylus and Prosierola, and the mother
may subsequently mate again with one of her
male offspring. Maternal care has also been ob-
served in Goniozus (Hardy & Blackburn 1991).
Adults of both sexes feed on honey in culture, and
females have been observed feeding on
haemolymph exuding from the paralysed host.
ACKNOWLEDGMENTS
This work was carried out while the first author was in
receipt of a grant from the Directorate General for Interna-
tional Cooperation of the Netherlands Government (DGIS).
We thank the following for the loan of material: J. Casevitz-
Weulersse and M. LaChaise (Museum National d'Histoire
Naturelle, Pans MNHN), E. De Coninck (Musee Royal de
l'Afrique Centrale, Tervuren, Belgium), V Rainen (Museo
Civico di Stona Naturale, Genoa, Italy MCSN) and M
Terayama (Biological Laboratory, Toho Institute of Educa-
tion, Tokyo). J. M. Carpenter (American Museum of Natural
History, New York), H. E. Evans (Dept of Entomology, Colo-
rado State University), I. C. W Hardy (Dept of Population
Biology, University of Leiden), A. Menke (Systematic Ento-
mology Lab. , USDA) and M. Mickevich (Maryland Center
for Systematic Entomology, University of Maryland) are
thanked for their comments on the manuscript, and R. W.
Hodges (Systematic Entomology Laboratory, USDA) for as-
sistance with Lepidoptera systematics.
LITERATURE CITED
Brothers, D. J. 1976. Modifications of the metapostnotum and
origin of the 'propodeal triangle' in Hymenoptera
Aculeata. Systematic Entomology 1: 177-182.
Brown, W. L. 1987. Neoclystopsenella (Bethylidae) a synonym
of Tamnoma (Formicidae). Psyche 94: 337.
Brues, C T. 1907. Notes and descriptions of North American
parasitic Hymenoptera IV. Bulletin of the Wisconsin Natu-
ral History Society 5: 96-1 1 1
Brues, C. T. 1933. The parasitic Hymenoptera of the Baltic
Amber. Part 1. Bernstein-Forschungen 3 (1932): 4-172.
Cameron, P. 1883. Descriptions of new genera and species of
Hymenoptera. Transactions of the Entomological Society of
London 1883: 187-197.
Du Buysson, R 1898. Voyage de M. E. Simon dans l'Afrique
australe (janvier-avril 1893). Annates de la Societe
Entomologiaue de France 66: 351-363.
Evans, H. E. 1962. The genus Bethylus in North America
Breviora 16: 1-12.
Evans, H. E. 1964 A synopsis of the American Bethylidae
(Hymenoptera: Aculeata). Bulletmof the Museum of Com-
parative Zoology, Harvard Unwersity 133:67-151.
Evans, HE. 1978. The Bethylidae of America north of Mexico.
Memoirs of the American Entomological Institute 27: 1-332.
Volume 3. 1994
105
Farns, J. S. 1988. Henmg86 Reference. Version 1. 5. User's
Manual 17 pp.
Gordh, G. and H. E. Evans. 1976. A new species of Gomozus
imported into California from Ethiopia for the biologi-
cal control of pink bollworm and some notes on the
taxonomic status of Parasierola and Gomozus (Hy-
menoptera: Bethylidae). Proceedings of the Entomological
Society of Washington 78: 479-489.
Gordh, G. and L. Moczar. 1990. A catalog of the World
Bethylidae (Hymenoptera: Aculeata). Memoirs of the
American Entomological Institute 46: 1-364.
Hardy, I. C. W. 1992. Non-binomial sex allocation and brood
sex ratio variances in the parasitoid Hymenoptera. Oikos
65: 143-158.
Hardy, I. C. W. and T. M. Blackburn. 1991. Brood guarding
in a bethylid wasp. Ecological Entomology 16: 55-62.
Hempel, A. 1934. A Prorops nasuta Waterston no Brasil.
Archizvs do lnstituto Bwlogico Sao Paolo 5: 197-212.
Junne, L. 1807. Nouvelle methode de classer les Hymenopteres et
les Dipteres. J. J. Paschaud, Geneva. 1-319.
Kieffer, J. J. 1904. Description de nouveaux Dryininae et
Bethylinae du Musee Civique de Genes. Annali Museo
Civico di Storm Naturale di Genova 41: 351-412.
Kieffer, J. J. 1905. Description de nouveaux proctotrypides
exotiques. Annales de la Societe Saentiftque de Bruxelles29:
95-142.
Kieffer, J. J. 1907 (1906). Beschreibung neuer in
Naturhistorischen Museum zu Hamburg aufbewahrter
Proctotrypiden and Evaniiden. Berliner Entomologische
Zeitschrift 51: 258-278.
Kieffer,]. J. 1911. Nouveaux Bethylides et Dry mides exotiques
du British Museum du Londres. Annales de la Societe
Scientifique de Bruxelles 35: 200-233.
Kieffer, J. J. 1914. Hymenoptera, Proctotrupoidea, Bethylidae
Das Tierreich 41: 1-595.
Krombein, K. V. 1987. Synonymic notes on the Bethylidae
described by V. de Motschulsky (Hymenoptera:
Aculeata). Proceedings of the Entomological Society of Wash-
ington 89: 356-358.
Kurian, C. 1952. Descriptions of four new and record of one
known Bethyloidea (Parasitic Hymenoptera) from In-
dia. Agra University journal of Research 1: 63-72.
Kurian, C. 1955. Bethyloidea (Hymenoptera) from India
Agra University Journal of Research 4: 67-155.
Latreille, P. A. 1802. Histoire Naturelle des Crustaces et Insectes.
Tome 3, 1-467.
Melo, G. A. R. and H. E. Evans. 1993. Two new Mtcrostigmus
species (Hymenoptera, Sphecidae), with the descrip-
tion of their parasite, Gomozus microstigmi sp. n. Proceed-
ings of the Entomological Society of Washington 95: 258-263.
Menke, A. S. 1993. Notauli and parapsidal lines: just what are
they? Sphecos 24: 9-12.
Muesebeck, C. F. W. 1940 Two new hymenopterous para-
sites of sugarcane borers in India. Proceedings of the
Entomological Society of Washington 42: 120-122.
Nagy, C. G. 1976. Bethylidae (Hymenoptera) parasitizing
orchard caterpillars. Revue Roumame de Biologie, Serie de
Bwlogie Animate 21: 103-108.
Richards, O. W. 1955. On the Bethyloidea (Hymen. ) of Israel.
Bulletin of the Research Council of Israel 4: 357-359.
Sorg, M. 1988. Zur phylogenie und systematik der Bethylidae
(Insecta: Hymenoptera: Chrysidoidea). Geologisches
lnstitut der Umversitaet zu Koeln Sonderveroeffentlichungen
63: 1-146.
Terayama, M. 1990. Keys to the Japanese Bethylidae (Hy-
menoptera, Aculeata) I. Subfamilies and Genera. Bulle-
tin of the Toho Gakuen 5: 19-43.
Thomson, C. G. 1862. Svenges Proctotruper (Fortsattning).
Svenska Vetenskapsakadenuen. Stockholm. Oefrersigt af K.
Academinens. Forhandlmgar. 18: 451-453.
Tryapitsyn, V. A. 1978. [Identification of the insects of the
European part of the USSR. Volume 3. Hymenoptera.
Bethyloidea, Bethylidae. ] Opredeliteh Faune SSSR 120: 3-
16 (in Russian).
J. HYM. RES.
Vol. 3, 1994, pp. 107-117
Variation in the Haemolymph Protein Composition of Confined
Apis Mellifera and Partial Restoration of Vitellogenin Titre by
Juvenile Hormone Analogue Treatment
M. M. G. Bitondi, Z. L. P. Simoes, A. M. do Nascimento and S. L. Garcia
Dept. Biology, Faculdade de Filosofia, Ciencias e Letras de Ribeirao Preto Umversidade de Sao Paulo, CEP. 14040-901
Ribeirao Preto, S.P., Brasil (AMN and SLG are undergraduate fellows of the Convenio CNPq/USP)
Abstract . — Haemolymph proteins, especially vitellogenin ( Vg), were investigated in confined Apis mellifera workers, that
were fed different diets and treated with juvenile hormone (JH) I, III or with pyriproxifen (PPN). Vg and total protein
concentrations were drastically decreased in the haemolymph of workers removed from the colony and confined for different
periods of time. SDS-PAGE analysis demonstrated that confinement also caused induction and repression of the synthesis of
certain haemolymph proteins. All of these changes occurred even when the confined workers were fed a protein-rich diet. In
workers with Vg deficiency induced by confinement PPN, but not JHI or JHIII, induced a partial increase in Vg concentration.
INTRODUCTION
The vitellogenin (Vg) of Apis mellifera, the
precursor of vitellin — the major protein of yolk —
is a glycolipoprotein produced in the fat body
(Harnish and White 1982; Trenczek and Engels
1986; Shapiro et al. 1988; Wheeler and Kawooya
1990). In most insects, Vg synthesis may be con-
trolled by the diet, which obviously provides the
materials and energy needed for this process
(Bianchi and Pereira 1987; Bownes 1989; Bownes
and Reid 1990). However, other factors in addition
to diet are also involved. Many studies have been
conducted on the role of neurosecretory cells (Elliott
and Gillott 1978) and neurohormones (Keeley and
Mckercher 1985; Keeley et al. 1988; Girardie et al.
1992), and of juvenile hormones and ecdysteroids
in the regulation of Vg synthesis (Adams et al.
1985; Borowsky et al. 1985; Hagedorn 1985;
Schwartz et al. 1985; Wojchowsky and Kunkel
1987; Adams and Filipi 1988; Keeley et al. 1988; Ma
et al. 1988; Roseler and Roseler 1988; Wyatt 1988;
Bownes 1989; Bownes and Reid 1990; Davis et al.
1990; Hatakeyama and Oishi 1990; Yin et al. 1990;
Agui et al. 1985, 1991; Don- Wheeler and Engelman
1991;Sochaeffl/. 1991).
In Apis mellifera, a highly eusocial insect, the
control exerted by the queen over the workers
represents an additional factor influencing Vg
synthesis. A queen pheromone inhibits oocyte
development in the workers. As a result Vg is not
incorporated into the oocytes, although Vg is de-
tected in the workers haemolymph. However, in
queenless colonies the Vg titre of workers in-
creases, reaching a level similar to that observed in
the queens, followed by oocyte growth and ovipo-
sition (Engels 1974; Engels and Fahrenhorst 1974).
How Vg synthesis is regulated in A. mellifera
still remains an interesting question, and aspects
of this regulation have been studied in queens,
workers and drones. In queens Vg synthesis does
not depend on functional corpora allata and is not
mediated by JH (Engels and Ramamurty 1976;
Kaatz 1985). Similarly, JH topical application does
not increase Vg synthesis in drones (Trenczek et al.
1989). But in workers, Rutz etal. (1976) and Flurief
al. (1977) observed that low JH doses applied on 6
day old workers stimulate Vg synthesis whereas
high doses have an inhibitory effect. Furthermore,
Rutz et al. (1976) observed a correlation in vivo
between low JH titre and Vg synthesis. Within the
first few days after worker emergence, character-
ized by low JH titres in haemolymph there is an
increase in Vg synthesis. After this period Vg
108
Journal of Hymenoptera Research
synthesis decreases while JH titre increases. This
increase in JH titre was confirmed by Huang et al.
(1991) and the temporal changes in Vg titre ob-
served by Rutz et al. (1976) were similar to those
related by Engels et al. (1990).
The investigation of factors that affect A.
mellifera Vg synthesis can help to understand how
this protein is regulated. In the present work, the
effect of some factors such as changes in social
environment (confinement of workers with or
without a queen) and diet (protein-rich or not) and
JH or PPN (pyriproxyfen, 2-[l-methyl-2(4-
phenoxyphenoxy) ethoxyl] pyridine, a JH ana-
logue, were investigated not only on worker Vg
synthesis but also on other haemolymph proteins.
MATERIALS AND METHODS
Apis mellifera
We used "wild type" Africanized Apis mellifera
bees (hybrids of European A. m. ligustica, A. m.
carriica, A. m. mellifera and the African bee A. m.
scutellata) from the Experimental Apiary of the
Department of Genetics, Faculty of Medicine of
Ribeirao Preto, University of Sao Paulo.
Combs containing workers ready to emerge
were removed from colonies and placed in an
incubator whose temperature (34°C) and R.H.
(80%) were similar to those in the colony. The
workers that emerged within 15-20 hours were
collected. About 100 newly emerged workers were
marked on the thorax and put back into a small
colony, formed by a queen, approximately 3000
workers (hive and forager bees), brood (eggs, lar-
vae and pupae) and, sometimes, a few drones.
Presence of nectar and pollen into the combs were
also checked.
Confinement
The remaining workers were separated into
groups of 150-200 and immediately submitted to
confinement in 8 x 11 x 13 cm wooden cages with
a sliding glass door and a screened bottom. The
workers in these cages were placed in an incubator
at 34°C and 80% R.H. and confined for 6 days
(short confinement) or 15-16 days (long confine-
ment). The confined bees received water and food
ad libitum. The diet consisted of 50% sugar in water
(syrup), a mixture of pollen from the comb (bee-
bread) and candy (powdered sugar and honey), or
only candy.
In three experiments, three naturally mated
queens aged 60-90 days were removed from the
respective colonies and confined with groups of
150-200 newly emerged workers. These groups
formed by workers and a queen were confined for
15-16 days, and fed on the mixture of beebread and
candy. Water was also supplied.
Treatment of confined workers
with JHIII, JHI and PPN
Some groups of 150-200 workers confined for
6 days were treated with JHIII, JHI or PPN applied
topically to the abdominal cuticle. Each worker
received 1 ul of a hormone solution in acetone at a
concentration of 1 ug/ul, administered in two
equal doses, the first immediately after emergence,
before the introduction into the cages, and the
other on the third day after rapid anesthesia with
gaseous nitrogen. Two worker groups were treated
with JHIII, two groups with JHI and nine groups
with PPN.
Three control groups were prepared in paral-
lel: the first consisted of marked workers reintro-
duced into the colony (control a), the second formed
by confined workers treated with two 1 ul doses of
acetone on the first and third days, respectively
(control b). This group was also submitted to a
rapid anesthesia with gaseous nitrogen at third
day, immediately before acetone treatment. The
third group consisted of untreated confined work-
ers (control c). All worker groups, except control a,
were allowed to feed ad libitum on the mixture of
beebread and candy. Water was also supplied.
Haemolymph
For collecting haemolymph the workers were
anesthetized with gaseous nitrogen and immobi-
lized on dissection plates. Haemolymph was ex-
tracted through a small superficial incision in the
dorsal cuticle between the 2nd and 3rd tergites.
Haemolymph was withdrawn from 6 day old
and 15-16 day old confined workers that were fed
on different diets, treated or not with hormones or
acetone and mantained with or without a queen
(Table I). Haemolymph was also extracted from
colony reared workers (6 or 15-16 day old) and
from newly emerged workers before confining or
Volume 3, 1 994
109
returning them to the colony.
Haemolymph pools were prepared from
groups of at least 80 workers obtained from the
same confinement cage. Similar pools were pre-
pared with the haemolymph of at least 20 workers
of the same age maintained in the colony and of 1 2
newly emerged workers. Phenylthiourea was
added to the pools. Haemolymph pools were cen-
trifuged at 3080 g for 10 min at 0°C and the super-
natant was stored at -20°C.
Rocket Immunoelectrophoresis
Immunoelectrophoresis was used for the quan-
titative determination of the vitellogenin fraction
in haemolymph. Monospecific vitellogenin anti-
serum produced in rabbits (Simoes 1980) was
added at a 1% concentration to 1% agarose gels
prepared with 0.06 M Tris-HCl buffer, pH 8.6.
Immunoelectrophoresis was carried out at 10°C
for 16 hours, at 0.08 V/cm gel. The same buffer
used in the gel was used in the electrode compart-
ments at a concentration of 0.3 M. The gels were
stained with Coomassie Blue R-250. The height of
the peaks detected (reported as mm) was propor-
tional to the amount of antigen. The values ob-
tained for confined workers were compared to
those for workers of the same age maintained in
colonies. The Vg peaks detected in colony reared
workers (control c) was considered to be 100%.
Total protein
Protein concentration in the haemolymph
pools was determined using bovine serum albu-
min as a standard (Bradford 1976).
SDS-PAGE
Soluble haemolymph proteins were separated
by SDS-PAGE according to the method of Laemmli
(1970) except that SDS was not used in the separat-
ing and stacking buffers. A 5-12% acrylamide
gradient was used on a 0.7-mm thick gel. Electro-
phoresis was run at 12mA constant current at 10°C
until bromophenol blue tracking dye reached the
bottom of the slab.
Haemolymph samples (5 \i\) from confined
and newly emerged workers (diluted 1: 2, v/v, in
sample buffer) and from workers maintained in
the colonies (diluted 1: 20 or 1: 40, v/v) were
applied to the acrylamide gel.
RESULTS
Confinement blocks the increase
of haemolymph Vg
The confinement of workers for a period of 6
days after emergence prevented the increase of Vg
titre in haemolymph that normally occurs in work-
ers in colonies. This occurred even when bee-
bread, the natural source of bee protein was sup-
plied to the confined bees. In some of the
haemolymph pools an extremely low Vg titre was
detected (Fig. 1, wells 5 and 6), but, on average, Vg
titres in these pools corresponded to 6.0 ± 2.1%
(Table II) of that present in workers maintained in
the colonies under natural conditions (Fig. 1 , wells
1 and 2).
A small increase in Vg titre was observed in
15-16 day confined workers (Fig. 1, wells 3 and 4).
This only occurred when the workers were fed
beebread and candy. Even with this diet, the Vg
concentration was much lower than in workers of
the same age maintained in the colonies (Fig. 1,
wells 1 and 2).
Confinement changes the protein pattern of
haemolymph obtained by SDS-PAGE
The pattern of soluble haemolymph proteins
from six day confined workers (Fig. 2, lane 4)
differed from that observed in workers main-
tained in colonies during the same period of time
(Fig. 2, lane 3). In the confined bees, in addition to
the fact that the Vg band (as determined by
Trenczek et al. 1989) was very weak, the a and c
polypeptides were not observed, whereas the b
polypeptide formed a strong band. This polypep-
tide corresponded to a weak band in the workers
living in colonies. These were the most obvious
differences, however differences between low
molecular weight polypeptides, were also ob-
served.
In Fig. 2, lanes 1, 2 correspond to the
haemolymph protein pattern of newly emerged
workers, collected immediately before the bees
were confined or returned to the colony. This
pattern changed as the bees maintained in the
colony aged (Fig. 2, lane 3). However, the changes
depended on the social environment as shown by
the protein pattern of confined workers (Fig. 2,
lane 4).
110
Journal of Hymenoptera Research
vg
^*t|l'"iWWwwi«AM*'
J |
2 3 4 5 6
Fig. 1. Rocket Immunoelectrophoresis of worker
haemolymph. 150 |il queen egg antiserum in 15 ml agarose
gel. Antigen: 2 ul of a haemolymph pool per well. Staining:
Coomassie Brilliant Blue R-250. Adult workers (6 days) main-
tained naturally in the colony (1, 2), confined for 15-16 days
(3, 4) and fed beebread and candy and confined for 6-days (5,
6) and fed the same diet.
The haemolymph pattern of workers confined
for 15-16 days and fed candy and pollen (Fig. 3,
lanes 5 and 6) did not differ from that obtained
after a short confinement (Fig. 2, lane 4). However,
a discrete increase was observed in the Vg band of
workers confined for 15-16 days. The value of diet
protein content for protein synthesis is clearly
shown in Fig. 3, lanes 3, 4, corresponding to the
protein pattern of a haemolymph pool from work-
ers maintained on a carbohydrate (syrup) diet for
15-16 days. In the same figure, the protein pattern
of haemolymph from confined workers can also
be compared with that of newly emerged ones
(lanes 7, 8) and with that of workers maintained in
the colony for 15 days (lanes 1, 2).
There is no Vg in workers confined for
15-16 days with a queen
Workers confined for 15-16 days in the ab-
sence of a queen had Vg in their haemolymph
b
c
4 5 6 7 8 9 0
M(kDa)
Fig. 2. SDS-PAGE (5.0-12%). Coomassie Blue staining. Pat-
terns of worker haemolymph proteins. (1, 2) Newly emerged
workers. Haemolymph diluted 1: 2 (v/v) in sample buffer; (3,
7) 6-day old workers maintained naturally in the colony
Haemolymph diluted (3) 1: 20 (v/v) and (7) 1! 40 (v/v) in
sample buffer. Workers confined for 6 days after emergence
and treated with PPN (10), JHI (5,9), JHIII (6), or acetone (8,
11), or untreated (4). Haemolymph diluted 1: 2 (v/v) in
sample buffer. (M) - molecular weights in kDa according to
marker proteins. Note: Columns should be numbered 1-11.
when fed on the beebread and candy mixture, as
can be seen by Immunoelectrophoresis Fig. 1, wells
3 and 4. However if a queen is confined together
with a group of 150-200 workers during this same
time interval, from emergence until 15-16 days,
the workers will not have Vg in the haemolymph.
Thus, the queen effect on Vg synthesis could be
observed even in a different environment condi-
tion, i.e. that established by confinement. The
haemolymph protein SDS-PAGE pattern of these
workers was similar to that of bees confined in the
absence of a queen, except for a weak Vg band
present in the latter (results not shown).
PPN induces Vg titre increase in workers
confined for 6 days
Workers confined for six days after emer-
Volume 3. 1994
111
vg Wt*
b
^-Htm*
180
116
.97
•66
-45
I
— 29
12 3 4 5 6 7 8 M(kDa)
Fig. 3. SDS-PAGE (5.0-12%), Coomassie Blue staining. Pat-
terns of worker haemolymph proteins. (1, 2) 15-day old
workers maintained naturally in the colony. Haemolymph
diluted 1: 20 (v/v) in sample buffer; (3, 4) workers confined
for 15-16 days after emergence and fed syrup or (5, 6) bee-
bread and candy; haemolymph diluted 1: 2 (v/v) in sample
buffer; (7, 8) Newly emerged workers. Haemolymph diluted
1: 2 (v/v) in sample buffer. (M) - molecular weights in kDa
according to marker proteins.
gence, fed beebread and candy and treated with
PPN consistently showed a significant increase
(p<0.001) in haemolymph Vg concentration (Fig.
4, wells 3, 4, and Fig. 5, wells 3, 4) when compared
to confined workers treated with acetone (Fig. 4
wells 1, 2, 7, 8; Fig. 5 well 1) or untreated (Fig. 4,
wells 9, 10). This increase corresponded to ap-
proximately 17% of the Vg levels in the
haemolymph of workers naturally maintained in
the colonies (Fig. 4 wells 11, 12; Fig. 5, well 5). To
calculate Vg percentage, we used a total of 9
haemolymph pools derived from 9 experiments,
each consisting of 150-200 workers treated with
PPN. In this case, Vg concentration in control a
(workers maintained in the colonies) was consid-
ered to be 100% (Table II). Table II also shows that
the percentage of Vg detected in acetone-treated
workers (control b, 7.4 + 3.2%) did not differ
(0.5>p>0.4) from that detected in the untreated
control c (6.0 ±2.1%).
The Vg titre detected in workers treated with
PPN was similar to that detected in workers con-
fined for 15-16 days and feeding on pollen and
candy (Fig. 1, wells 3 and 4). This Vg concentra-
tion, however, was never detected in controls c
(workers confined for six days and feeding on
pollen and candy) or b (workers confined for six
days, feeding on pollen and candy and treated
3 4
7 8 9 10 11 12 13 14 15 16
Fig. 4. Rocket Immunoelectrophoresis of worker haemolymph. 150 ul queen egg antiserum in 15 ml agarose gel. Antigen: 2
ul of a haemolymph pool per well. Staining: Coomassie Brilliant Blue R-250. Adult workers confined for 6 days after emergence
and treated with JHIII (13, 14), JHI (5, 6, 15. 16), PPN (3, 4), or acetone (control b, 1, 2, 7, 8), or untreated (control c, 9, 10). These
confined workers were fed beebread and candy. Wells 11 and 12 correspond to the Vg of adult 6-day old workers maintained
naturally in colonies.
112
Journal of Hymenoptera Research
o
1 2
I
Fig. 5. Rocket Immunoelectrophoresis of worker
haemolymph. 150 ul queen egg antiserum in 15 ml agarose
gel. Antigen: 2 (il of a haemolymph pool per well. Staining:
Coomassie Brilliant Blue R-250. Adult workers confined for
6 days after emergence, fed beebread and candy and treated
with acetone (control b, well 1), JHI (well 2), or PPN (wells 3
and 4). Well 5 corresponds to the Vg of adult 6-day old
workers maintained naturally in colonies.
with acetone). Thus, under conditions of prolonged
confinement, there is an increase in Vg concentra-
tion in haemolymph during the second week of
confinement, but this increase never occurs dur-
ing the first week of confinement unless the work-
ers are treated with PPN.
PPN does not change the SDS-PAGE
protein pattern
The protein pattern (Fig. 2) from workers con-
fined for six days and treated with PPN (lane 10)
was similar to that observed in the acetone-treated
(lanes 8, 1 1) and untreated controls (lane 4), except
by an increase in Vg band.
PPN does not significantly change the total protein
concentration in the haemolymph
There was no significant difference in
haemolymph protein content between PPN-rreated
workers and their controls: acetone treated (con-
trol b, p>0.35) and untreated (control c, p>0.15).
(Table III).
Comparison between untreated groups (con-
trol c) and the groups maintained in the colonies
(control a) demonstrated a drastic decrease in
haemolymph protein concentration in the first
group, maintained under confinement conditions,
even when beebread and candy was supplied.
JHI and jHIll did not induce Vg titre increase in
workers confined for 6 days
Treatment with JHI (Fig. 4, wells 5, 6, 15, 16
and Fig. 5, well 2) or JHIII (Fig. 4, wells 13, 14)
under the same experimental conditions as those
with PPN did not increase Vg concentration in
haemolymph. The Vg peaks detected in the work-
ers treated with JHI or JHIII did not differ statisti-
cally (p>0.15 and p>0.5, respectively) from the
acetone-treated control b (Fig. 4, wells 1, 2, 7, 8 and
Fig. 5, well 1 as showed in Table IV). However, the
Vg peaks obtained from workers treated with JHI,
a hormone not synthesized by Apis mellifera
(Robinson et al. 1987), were found to be slightly
higher than the peaks obtained for workers treated
with acetone or with JHIII, the natural hormone of
these bees. This result should be considered with
caution since it is based only on the data obtained
for two haemolymph pools from workers treated
with JHI or JHIII. But we do not exclude the
possibility of this homologue (JHI) being more
effective on Apis mellifera than JHIII.
JHI and JHIII did not change the SDS-PAGE
protein pattern
The haemolymph protein pattern of workers
confined for six days and treated with JHI (Fig. 2,
lanes 5, 9) or JHIII (Fig. 2, lane 6) was similar to that
observed in the controls treated with acetone (Fig.
2, lanes 8, 11) or untreated (Fig. 2, lane 4).
Volume 3, 1 994
113
DISCUSSION
Effect of confinement, queen presence and diet on
haemolymph protein composition
Our results show that the normal Vg titre in
workers mainly depends on social environment
established in the colony. When workers were
removed from the colony, and maintained during
6 days in an appropriate environment (where
mortality was practically zero) and on a protein-
rich diet, profound physiological changes occurred,
that inhibited Vg titre and probably Vg synthesis.
The initiation of Vg synthesis in these workers can
be observed if confinement is lenghthened for 15-
16 days. But the haemolymph Vg peak detected by
immunoelectrophoresis in these workers (after
15-16 days of confinement) is smaller than that
observed in colony reared workers. The onset of
Vg in the haemolymph of 15-16 days confined
workers is dependent on administration of a pro-
tein-rich diet: Vg is not detected in these confined
workers fed a diet without protein such as syrup,
or a low protein content diet, such as candy made
with honey. Besides if a queen is confined with the
workers for 15-16 days no Vg is produced al-
though a protein rich diet (beebread and pollen)
had been available.
The influence of factors related to the social
environment, on Vg synthesis has been studied in
social Hymenoptera, especially the interaction
among individuals of different castes in the colo-
nies. Reproductively active queens characteristi-
cally inhibit Vg synthesis and egg-laying in other
females capable of reproduction. This fact has
been well documented in A. mellifera (Engels et al.
1990), Melipona (Engels and Imperatriz-Fonseca
1990), and Camponotus festinatus (Martinez and
Wheeler 1991) among other social insects. We
verify that A. mellifera queens can inhibit Vg syn-
thesis in workers even under conditions of con-
finement, i.e., far from the normal colony environ-
ment. Engels et al. (1990) maintained groups of 25
workers for three weeks confined from emergence
on a piece of comb, in the presence and absence of
a queen. They detected that both worker groups
had Vg in the haemolymph, but the Vg titre of
workers confined in the absence of a queen was
higher than that of workers confined in the pres-
ence of a queen. In our experiments, we found that
workers confined without a queen for 15-16 days
produced Vg when fed beebread and candy. How-
ever, when the workers were confined with a
queen during this same period of time, no Vg was
detected in the haemolymph. Perhaps, the experi-
mental conditions employed by Engels et al. 1990,
with a piece of comb (with brood?) within the
confinement cage, provided a more favorable en-
vironment (maybe more similar to that of a normal
colony) that permitted Vg synthesis even in the
presence of a queen. For comparison, other condi-
tions employed in both experiments should also
be taken into account, such as the age of the queens
used, the size of the worker population and the
time of confinement.
Confinement affects not only Vg synthesis,
which is also controlled by the queen and by the
food available, but also the synthesis of other
haemolymph proteins, as determined by SDS-
PAGE. The polypeptide a showed marked reduc-
tion, the polypeptide c was not detected, whereas
another, b, showed markedly increased titer (Figs.
2 and 3). This indicates that confinement can si-
multaneously provoke contrasting gene expres-
sions. This was observed in confined workers fed
proteic or non-proteic diets. Workers maintained
on a syrup diet had a lower protein content, but
similar protein pattern in the haemolymph (Fig. 3)
when compared to workers maintained on a proteic
diet. Thus the social interactions and not the pro-
tein supply are responsible for the contrasting
gene expressions mentioned above.
Action ofPPN, JHIII and ]H1 on haemolymph protein
composition changed by confinement
A significant (p<0.001) increase in Vg titre
ocurred in workers with confinement-induced Vg
deficiency, after treatment with PPN. However
the observed induction was partial. Apparently
other factors are also involved in the regulation of
normal Vg synthesis. The action of these factors
may be prevented or impaired by the confinement
conditions employed.
PPN seems to be specific for Vg synthesis,
since the haemolymph SDS-PAGE pattern of the
other proteins in confined workers apparently did
not change when treated with this JH analogue.
However as the small increase in Vg titre induced
by PPN was not reflected in total protein measure-
ment we can not exclude the possibility that PPN
also partially inhibits other haemolymph
114
Journal of Hymenoptera Research
protein(s).
Contrary to what is observed with PPN, the
natural JHIII (Robinson et al. 1987) and its homo-
logue JHI did not significantly increase Vg con-
centration in haemolymph. JHIII and JHI also did
not seem to modify the haemolymph protein pat-
tern induced by confinement. We should consider
that PPN may have caused an increase in Vg titre
in haemolymph by being a more potent analogue.
PPN is considered to be one of the most effective
juvenile hormone analogues known for locusts
(De Kort and Koopmanschap 1991), with a strong
juvenilizing effect on these insects. PPN has a
strong morphogenetic effect when topically ap-
plied to A. mellifera larvae. Larvae treated with 1 fig
undergo metamorphosis, but the pupae show dras-
tic changes in pigmentation, especially in the eyes
and thorax, and can also die before emergence
(Bitondi et al., unpublished data). Similarly to
what occurs in Locusta, the effect of PPN on A.
mellifera larvae and pupae is more drastic when
compared to the effects induced by JHIII or JHI.
Kaatz (1985) proposed a model of regulation
of Vg synthesis in A. mellifera queens. According to
this model, Vg synthesis may be influenced by JH
but also by ecdysteroids and by a haemolymph
factor. These factors mentioned by Kaatz (1985)
may be involved in the regulation of normal Vg
synthesis in workers. However, the simple adap-
tation to workers of the model proposed for queens
should be considered with caution since some
data obtained in studies on Vg synthesis regula-
tion in queens indicate that this regulation may
differ between the two castes (Engels et al. 1990).
In summary the absence of one or more types
of social interaction induced by confinement
caused a reduction of total protein content in
haemolymph, induction and repression of protein
synthesis and impairment of Vg synthesis in A.
mellifera workers. Only haemolymph Vg titer can
be partially recovered by PPN treatment of con-
fined workers.
ACKNOWLEDGMENTS
This work was supported by grants from the Fundacao
de Amparo a Pesquisa do Estado de Sao Paulo (FAPESP)
Proc. 91/2587-8.
We thank Dr. H Oouchi, Sumitomo Chemical Co.,
Ltd, Osaka, Japan, for providing the pynproxyfen, and PR.
Epifanio and L. R. Aguiar for the technical assistance We
thank also Dr. David De Jong for correcting the English.
LITERATURE CITED
Adams, T. S. and P A. Filipi. 1988. Interaction between
]uvenile hormone, 20-hydroxyecdysone, the corpus -
cardiacum - allatum complex, and the ovaries in regulat-
ing vitellogenin levels in the housefly, Musca domestica.
journal of Insect Physiology 34: 11-19.
Adams, T. S., H. H. Hagedorn and G. D. Wheelock. 1985.
Haemolymph ecdysteroid in the housefly, Musca
domestica, during oogenesis and its relationship with
vitellogenin levels, journal of Insect Physiology 31: 91-97.
Agui, N., S. Izumi and S. Tomino. 1985. The role of ecdysteroids
and juvenoids in vitellogenin levels and follicle devel-
opment in the housefly, Musca domestica. Applied Ento-
mology and Zoology 20: 179-188.
Agui, N., T. Shimada, S. Izumi and S. Tomino. 1991. Hor-
monal control of vitellogenin mRNA levels in the male
and female housefly, Musca domestica. Journal of Insect
Physiology 37: 383-390.
Bianchi, A. G. de and S. D. Pereira. 1987. Time of synthesis of
Musca domestica vitellogenin during the first gonotrophic
cycle. Comparative Biochemistry, and Physiology 86B(4):
697-700.
Borovsky, D., B. R. Thomas, D. A. Carlson, L. R. Whisenton
and M. S. Fuchs. 1985. Juvenile hormone and 20-
hydroxyecdysone as primary and secondary stimuli of
v\te\\oger\esism Aedes aegypti. Archwes of Insect Biochem-
istry and Physiology 2'. 75-90.
Bownes, M. 1989. The roles of juvenile hormone, ecdysone
and the ovary in the control of Drosophila vitellogenesis.
Journal of Insect Physiology 35(5): 409-413.
Bownes, M and G. Reid. 1990. The role of the ovary and
nutritional signals in the regulation of fat body yolk
protein gene expression in Drosophila melanogaster. jour-
nal of Insect Physiology 36(7): 471-479.
Bradford, M.. 1976. A rapid and sensitive method for the
quantification of microgram quantities of protein utiliz-
ing the principle of protein dye binding. Analytical Bio-
chemistry 72: 248-254.
Davis, R. E., T. J. Kelly, E. P. Masler, H. W. Fescemyer, B. S.
Thyagaraja and A. B. Borkovec. 1990 Hormonal control
of vitellogenesis in the gypsy moth, Lymantria dispar
(L.): suppression of haemolymph vitellogenin by the
juvenile hormone analogue, methoprene. Journal of In-
sect Physiology 36(4): 231-238.
De Kort, C. A. D. and A. B. Koopmanschap. 1991. A juvenile
hormone analogue affects the protein pattern of the
haemolymph in last-instar larvae of Locusta nugraloria.
Journal of Insect Physiology 37(2): 87-93.
Volume 3, 1 994
115
Don-Wheeler, G. and F. Engelmann. 1991. The female - and
male - produced vitellogenins of Leucophaea maderae.
Journal of Insect Physiology 37(12): 869-882.
Elliot, R. H. and C. Gillot. 1978. The neuro-endocrine control
of protein metabolism in the migratory grasshopper,
Melanoplus sangumipes. Journal of Insect Physiology 24:
119-126.
Engels, W. 1974. Occurrence and significance of vitellogenins
in female castes of social Hymenoptera. American Zoolo-
gists 14: 1229-1237.
Engels, W. and H. Fahrenhorst. 1974. Alters - und kasten -
spezifische Veranderungen der Haemolymph - Protein
- Spektren bei Apis mellifica. Wilhelm Roux' Archiv 174:
285-296.
Engels, W. and V. L. Imperatriz-Fonseca. 1990. Caste devel-
opment, reproductive strategies, and control of fertility
in honey bees and stingless bees. In: Engels, W. (ed.)
Social Insects. An evolutionary approach to castes and repro-
duction. Springer-Verlag, Berlin, ppl67-230.
Engels, W., H. Kaatz, A. Zillikens, Z. L. Paulino-Simoes, A.
Trube, R. Braun and F. Dittrich. 1990. Honey bee repro-
duction: vitellogenin and caste-specific regulation of
fertility. In: M. Hoshi and O. Yamashita (eds.) Advances
in invertebrate reproduction 5, 495-502. Elsevier,
Amsterdam.
Engels, W and P. S. Ramamurty. 1976. Effects of carbon
dioxide on vitellogenin metabolism in unmated queen
honeybees. Journal of Apiculture Research 15: 3-10.
Fluri, P., H. Wille; L. Gerig and M. Luscher. 1977. Juvenile
hormone, vitellogenin and haemocyte composition in
winter worker honey bees (Apis mellif era). Experientia 33:
1240-1241.
Girardie, J., O. Richard and A. Girardie. 1992. Time-depen-
dent variations in the activity of a novel ovary maturat-
ing neurohormone from the nervous corpora cardiaca
during oogenesis in the locust, Locusta migratoria
migratorwides. Journal of Insect Physiology 38: 215-221.
Hagedorn, H. H.. 1985. The role of ecdysteroids in reproduc-
tion. In Comprehensive Insect Physiology, Biochemistry and
Pliarmacology. (Edited by G. A. Kerkut and L. I. Gilbert),
vol. 8, pp 205-262. Pergamon Press, Oxford.
Harnish, D. G. and B. N. White. 1982. Insect vitellins: identi-
fication, purification, and characterization from eight
orders. Journal of Experimental Zoology 320: 1-10.
Hatakeyama, M. e K. Oishi. 1990. Induction of vitellogenin
synthesis and maturation of transplanted previtellogenic
eggs by juvenile hormone III in males of the sawfly,
Athalia rosae. Journal of Insect Physiology 36: 791-797.
Huang, Z. Y., G. E. Robinson, S. S. Tobe, K. J. Yagi, C. Strambi,
A. Strambi and B. Stay. 1991. Hormonal regulation of
behavioral development in the honey bee is based on
changes in the rate of juvenile hormone biosynthesis.
Journal of Insect Physiology 37(10): 733-741.
Kaatz, H. H.. 1985. Initiation und Regulation der Vitellogenin-
Synthese bei der Bienenkbnigin (Apis mellifera L.). Inau-
gural-Dissertation, Universitat Tubingen, pp. 146-159.
Keeley, L. L. and S. C. McKercher. 1985. Endocrine regula-
tions of ovarian maturation in the cockroach Blaberus
discoidales. Comparatwe Biochemistry and Physiology 80A:
115-121.
Keeley, L. L., S. M Sowa, T. K. Hayes and]. Y. Bradfield. 1988.
Neuroendocrine and juvenile hormone effects on fat
body protein synthesis during the reproductive cycle in
female Blaberus discoidalis cockroaches. General Com-
paratwe Endocrinology 72: 364-373.
Laemmli, U. K. 1970. Cleavage of structural proteins during
the assembly of the head of bacteriophage T4. Nature
227: 680-685.
Ma, M, J. -Z. Zhang and H. Gong. 1988. Permissive action of
juvenile hormone on vitellogenin production by the
mosquito Aedes aegypti. Journal of Insect Physiology 34:
593-596.
Martinez, T. and D. Wheeler. 1991. Effect of the queen, brood
and worker caste on haemolymph vitellogenin titre in
Camponotus festinatus workers. Journal of Insect Physiol-
ogy 37(5): 347-352.
Robinson, G. E., Strambi, A., Strambi, C, Paulino-Simoes,
Z.L., Tozeto, SO., Barbosa, J. M. N. 1987. Juvenile hor-
mone titers in Africanized and European honey bees in
Brazil. Gen. Comp. Endocrinol. 66: 457-459.
Roseler, P. F. and I. Roseler. 1988. Influence of juvenile
hormone on fat body metabolism in ovanolectomized
queens of the bumblebee, Bombus terrestris. Insect Bio-
chemistry 18(6): 557-563.
Rutz, W., L. Gerig;, H. Wille and M. Luscher. 1976. The
function of juvenile hormone in adult worker honey-
bees, Apis mellifera. Journal of Insect Physiology 22: 1485-
1491.
Schwartz, M. B., T. J. Kelly, R. B. Imberski and E. C. Rubenstein.
1985. The effects of nutrition and methoprene treatment
on ovarian ecdysteroid synthesis in Drosophila
melanogaster. Journal of Insect Physiology 31: 947-957.
Shapiro, J. P., J. H. Law and M. A. Wells. 1988. Lipid transport
in insects. Annual Rei'iew of Entomology 33: 297-318.
Simoes, Z. L. P. 1980. Estudo da vitelogenina e da vitelina em
Apis mellifera L. (Hymenoptera : Apoidea). Tese de
doutoramento, Universidade de Sao Paulo, pp.: 1-108.
Socha, R., J. Sula, D. Kodrik and I. Gelbic. 1991. Hormonal
control of vitellogenin synthesis in Pyrrhocoris apterus
(L.) (Heteroptera). Journal of Insect Physiology 37( ): 805-
816.
Trenczek, T. and W. Engels. 1986. Occurrence of vitellogenin
in drone honeybees. International Journal of Invertebrate
Reprodution and Development 10: 307-311.
Trenczek, T., A. Zillikens and W. Engels. 1989. Developmen-
tal patterns of vitellogenin haemolymph titre and rate of
synthesis in adult drone honey bees (Apis mellifera).
Journal of Insect Physiology 35: 475-481.
Wheeler, D. E. and J. K. Kawooya. 1990. Purification and
characterization of honey bee vitellogenin. Archives of
Insect Biochemistry and Physiology 14: 253-267.
Wojchowski, D. M. and J. G. Kunkel. 1987. Purification of two
distinct oocyte vitellins and identification of their corre-
sponding vitellogenins in fat body and hemolymph of
Blaberus discoidalis. Insect Biochemistry 17: 189-198.
Wyatt, G. R.. 1988. Vitellogenin synthesis and the analysis of
juvenile hormone action in locust fat body. Canadian
Journal of Zoology 66: 2600-2610.
Yin, C. -M„ B. -X. Zou, S. -X. Yi and J. G. Jr. Stoffolano. 1990.
Ecdysteroid activity during oogenesis in the black blow-
fly, Phormia regina (Meigen). Journal of Insect Physiology
36: 375-382.
116
Journal of Hymenoptera Research
Table I. Haemolymph pools prepared from workers confined for 6 or 15-16 days in the presence or absence of a queen,
fed on different diets and submitted or not to hormonal treatment
No. of
Confinement
Diet
Hormonal or
pools
(days from
acetone
analysed
emergence)
treatment
1
6
Syrup
_
1
6
Candy
-
5
6
breebread and candy
-
2
6
breebread and candy
JHIII
2
6
breebread and candy
JHI
9
6
breebread and candy
PPN
6
6
breebread and candv
Acetone
2
15-16
Syrup
-
1
15-16
Candy
-
3
15-16
breebread and candy
-
3
15-16
breebread and candy
-
Queen
absent
absent
absent
absent
absent
absent
absent
absent
absent
absent
present
Table II - Vitellogenin (Vg) in haemolymph pools from confined workers treated with PPN or acetone (control b), from
untreated confined workers (control c) and from workers maintained in colonies (control a).
PPN
Acetone
Untreated
Maintainec
treatec
treated
in the
colonies
nent
(control b)
(control
c)
(control a)
Experir
Peak
Vg#
Peak
Vg#
Peak
Vg#
Peak
Vg#
No.
Heigh
(mm)
(%)
Height
(mm)
(%)
Height
(mm)
(%)
Height
(mm)
(%)
1
5.0
16.13
3.5
11.29
1.5
4.39
31.0
100
:
5.5
17.74
*
*
*
»
•
*
3
411
12.90
»
*
•
*
•
•
4
6.5
20.97
*
*
*
•
•
*
5
6.5
15.12
2.0
4.65
3.0
6.98
43.0
100
6
7.0
17.95
1 i)
2.56
3.0
7.69
39.0
100
7
8.5
21.80
3.5
8.97
3.0
7.69
+
+
8
4.5
14.52
2 5
8.06
1.0
3.22
31.0
100
9
6.5
14.61
4 (I
8.99
z
z
44.5
100
MEAN
±SD
16.9±3.0
7.4±3.2
6.0±2.1
100
# Percent Vg in relation to workers maintained in the colony (control a = 100% Vg);
* Since experiments 1-4 were performed simultaneously, the same controls (a, b and c) were used in each;
z There was no control c for experiment 9;
+ Experiments 6 and 7 were performed during subsequent weeks. The same control a was used.
Volume 3, 1 994
117
Table III. Protein titre* (ug/|il haemolymph in BSA equivalents) of 6-day old workers confined from emergence
and treated with PPN or acetone (control b), untreated (control c) and maintained in the colony (control a).
Experiment
PPN treated
Acetone
Untreated
Maintained in
No.
treated
the colonies
(control b
(control c)
(control a)
|lg/(ll
M-g/^l
Hg/|d
\ig/\d
1
6.76
7.22
5.04
53.15
2
7.03
—
—
—
3
6.81
—
—
—
4
8.38
—
—
—
5
—
—
—
33.88
6
7.0
6.62
5.64
31.05
7
7.98
6.89
7.93
—
MEANiSD
7.3+0.7
6.9±0.3
6.2+.1.5
39.4+12.0
(')Protein concentrations were only measured in 6 of the 9 experimental groups treated with PPN, in 3 of the 6
groups treated with acetone, in 3 of the 5 untreated groups, and in 3 of the 5 groups maintained in the colonies.
Table IV. Vitellogenin (Vg) percent in haemolymph pools from confined workers treated with JHIII, JHI or
acetone (control b) and from workers maintained in the colonies (control a).
Experiment
JHI
JHIII
Acetone
Maintained in
No.
treated
treated
treated
the colonies
Peak
Vg
Peak
Vg
Peak
Vg
Peak Vg
Height
Height
Height
Height
(mm)
(%)
(mm)
(%)
(mm)
(%)
(mm) (%)
1
4.5
10.00
4.0
8.89
* *
2
4.0
8.89
3.5
7.78
1.5
3.33
45 100
3
2.0
4.60
2.0
4.60
43.5 100
MEAN±
SD
9.4+0.8
6.2±2.2
5.6±2.9
* Since experiments 1 and 2 were performed on subsequent days, the same control a was used.
J. HYM. RES.
Vol. 3, 1994, pp. 119-132
A Review of the Agile Species Group of Pison
(Hymenoptera: Sphecidae: Trypoxylini)
Alexander V. Antropov
Zoological Museum of the Moscow State University, Herzen Street 6, Moscow K-9, 103009 Russia
Abstract. — The agile species group of Pison is redescribed, the 12 included species are reviewed, and a key is provided for
identification. The agile group is restricted to the Oriental and eastern Palearctic Regions, although one species, koreense, is
adventive in North America. Five species are new: agiloides from Sri Lanka; chrysoptilum from Borneo; mngyuenfuense from
southwestern China; vechti from Malaya and Indonesia; and pulawskii from India. Other species of the group are: agile (Smith)
from southern India and Sri Lanka; erythropus Kohl from western India; koreense (Radoszkowski) from eastern Asia and North
America; rothneyi Cameron from southeast Asia; browni (Ashmead) from the Philippines; different Turner from Assam, India;
and hissancum Gussakovskij from Uzbekistan and Tajikistan. Lectotypes are designated for agile, differens and rufipes (Smith)
and a neotype is designated for koreense. Pison koreense is removed from synonymy with agile.
INTRODUCTION
The genus Pison, which contains nearly 200
described species (Bohart and Menke 1976; Menke
1988), is well represented in all faunal regions
except North America where only an adventive
species, koreense, is known.
In this paper I review the agile species group
which contains 12 species, five of which are new.
Members of this assemblage have been placed in
the subgenera Pisonoides and Krombeiniellum, but
Menke (1988) used species groups rather than
subgenera for infrageneric groups. The agile group
is restricted to the eastern Palearctic and the Orien-
tal Regions, except for the east Asian species
koreerxse which was introduced into North America
presumably after World War II (Krombein 1958a).
Morphological terminology used here follows
Bohart and Menke (1976) and Menke (1988). The
following abbreviations are used in the text: OOD
= ocello-ocular distance; OD = ocellus diameter;
POD = distance between posterior ocelli.
The following museum and institutions lent
specimens for this study (abbreviations are used
in the text):
AUZM
BMNH
Universiteit van Amsterdam, Zoologisch Museum,
Amsterdam, The Netherlands (W. Hogenes).
The Natural History Museum, London, England
(C. R. Vardy, L. Ficken).
CAS California Academy of Sciences, San Francisco,
California (W. J. Pulawski).
NMNH Nationaal Natuurhistorisch Museum, Leiden, The
Netherlands (C. van Achterberg).
OUM Hope Entomological Collections, University Mu-
seum, Oxford, England (C. OToole).
USNM U.S. National Museum, Washington, DC, USA (K.
V. Krombein, A. S. Menke).
ZIN Zoological Institute, Russian Academy of Sciences,
St. Petersburg, Russia (V. I. Tobias).
ZMK Zoological Museum, Copenhagen, Denmark (O.
Lomholdt).
The specimens from the collection of the Zoo-
logical Museum of the Moscow State University
(ZMUM) were also used in the review.
THE AGILE GROUP
This group is characterized as follows: com-
pound eyes densely setose, antenna clavate, oc-
cipital carina complete or nearly so, anterior
pronotal pit small, subomaulus present, episternal
sulcus straight, forewing with only two submar-
ginal cells (true second submarginal cell lost
through diminution), hindcoxa without dorsolat-
eral carina, metapleural flange usually lamelli-
form, propodeum without lateral carina or crenu-
late ridge, male sternum VIII narrow, gonostyle
simple, volsella small, penis valve compressed
120
Journal of Hymenoptera Research
laterally and without teeth or notched ventrally.
Menke (1988) regarded the setose eyes, the
clavate antenna, the two submarginal cells, the
presence of a subomaulus, the straight episternal
sulcus, the lamelliform metapleural flange and
claw shape as apomorphies of the agile group. Its
species apparently represent a monophyletic as-
semblage. Morphologically they are very similar
and differ mainly in leg color, tergal bands, punc-
tation, vestiture and proportions of tergum I. Male
genitalia are also similar. One of the new species,
pulaivskii, stands apart from the others in the group
because its metapleural flange is narrow. I regard
it as a plesiomorphic state. Two other species have
unique autapomorphies: in browni the propodeal
dorsum is delimited laterally by a shallow, broad
sulcus and tergum I has a distinct preapical de-
pression; in agiloides submarginal cell II is open
distally.
Description. — Inner orbits of eyes moderately
emarginate, parallel (eyes equidistant at vertex
and clypeus) or slightly converging below (rarely
above); eyes covered densely with short, erect
setae (Fig. 3); clypeus convex, in female rounded
(Figs. 4a, 5a), in male angulate or prominent apically
(Figs. 4b, 5b); frons convex; antennae clavate, com-
paratively short, with distal flagellomeres wider
than long; labrum subquadrate, truncate or slightly
emarginate apically; occipital carina a complete
(or almost complete) circle, narrowly separated
from hypostomal carina; male mandible simple,
that of female with inner tooth slightly distad of
midpoint; pronotum with small round pit anteri-
orly, without lamellae; scutum and mesopleuron
moderately, uniformly punctate; episternal sul-
cus almost straight, not curved forward ventrad;
subomaulus recurved ventrad; omaulus and ac-
etabular carina absent; mesopleural sulcus paral-
leled anteriorly by a row of foveolae; metapleuron
smooth; metapleural flange usually broadly lamel-
liform posteriorly (Figs. 6-8); tegula entirely punc-
tate; forewing media diverging after cu-a; forew-
ing with two submarginal cells (Figs. 10-13), 2nd
(really 3rd) usually not petiolate; recurrent veins
received by 1st and 2nd submarginal cells or 2nd
recurrent vein interstitial between 1st and 2nd
submarginal cells; hind coxa dorsum with low
inner carina, without outer carina; legs finely sculp-
tured, without stout spines on tibiae and
tarsomeres; all tarsomeres IV with small plantulae;
tarsal claw thick to just before apex; propodeum
rounded, without lateral carinae or lines of foveae
and crenulate ridges, punctate with smooth
interspaces; propodeal dorsum not delimited
(browni with shallow lateral sulcus), with medium
furrow containing short to complete ridge; abdo-
men compact; tergum I simple or with preapical
transverse depression (Figs. 15-19); apical bands
of terga often translucent and with silvery or golden
pubescence; male sternum VIII long, narrow,
rounded or weakly notched apically; genitalia
compact, compressed laterally; volsellae small,
rounded and weakly setose; gonostyle triangular,
simple, with long, coarse lateral setae curved be-
neath; penis valve compressed laterally, consider-
ably widened apically, without teeth or notches
ventrally.
Included Species. — agile (Smith), agiloides sp. n.,
browni (Ashmead), chrysoptilum sp. n., differens
Turner, erythropus Kohl, hissaricum Gussakovskij,
koreense (Radoszkowski), ningxjuenfuense sp. n.,
pulaivskii sp. n., rothneyi (Cameron), vechti sp. n.
Biology. — Information is available for two spe-
cies of the group, koreense and erythropus. The
former was studied by Iwata (1964) in Japan and
by Sheldon (1968) in North America. The Indian
species, en/thropus, was studied by Home (1870).
These wasps construct small, clay cells, placing
them separately or in groups on variable surfaces,
but not within linear cavities (borings in wood or
empty stems of plants). The groups may include
up to 21 cells which do not merge in a common
mass but keep their independence. Prey consists
of 6-31 paralyzed, small, usually immature spi-
ders.
KEY TO SPECIES OF THE AGILE GROUP
1 All femora completely reddish 2
— Mid- and hindfemora brown, forefemora mainly brown 6
2(1) Metapleural flange narrow (Fig. 9); India pulaivskii sp. n.
— Metapleural flange broadly lamelliform posteriorly (Figs. 6-8) 3
Volume 3. 1994 121
3(2) Pronotum, scutum, scutellum, metanotum and propodeal dorsum and hindface with suberect golden
setae; Borneo chrysoptilum sp. n.
Thorax and propodeum with only silvery pubescence 4
4(3) All legs including tarsi and base of trochanters yellowish-red; western India erythropus Kohl
— Trochanters, hintibiae apically and tarsi dark brown 5
5(4) Abdominal tergum I densely punctate (punctures separated by a diameter or less), dull due to dense
microsculpture; translucent apical bands of terga I-III whitish, at middle hardly broader than hindtarsal
diameter; Uzbekistan, Tajikistan hissaricum Gussakovskij
— Abdominal tergum I finely, sparsely punctate (punctures more than a diameter apart), surface smooth,
weakly shiny in spite of microstriae; translucent apical bands of terga golden, broad, those of 1 1— 1 1 1 at
middle almost equal to diameter of hindtibia; Malaysia, Indonesia vechti sp. n.
6(1) Abdominal tergum I polished, sculpture nearly same as following terga 7
— Abdominal tergum I dull or weakly shiny in contrast to following terga 8
7(6) Tibiae reddish; scutal punctures fine, dense, less than a diameter apart; translucent apical bands of
abdominal terga golden, apical bands of terga I-III at middle at least twice as broad as diameter of
hindtarsomere I; submarginal cell II open distally (Fig. 14); Sri Lanka agiloides sp. n.
— Mid- and hindtibiae brown, foretibia partly reddish; scutal punctures coarse, more than a diameter apart;
translucent apical bands of abdominal terga mainly whitish, those on I-III equal to or hardly broader than
diameter of hindtarsomere I (Fig. 15); submarginal cell II closed; southern India and Sri Lanka
agile (Smith)
8(6) Lamelliform part of metapleural flange mainly dark; translucent apical bands of abdominal terga not
broader than diameter of hindtarsomere I 9
Lamelliform part of metapleural flange reddish at least posteriorly; translucent apical bands of abdominal
terga (at least of terga II-III) obviously broader than diameter of hindtarsomere I 10
9(8) Abdominal tergum I dull, very densely, finely sculptured, transversely depressed preapically (Fig. 17);
apical bands of abdominal terga II-III dark brown, that of tergum I narrower than hindtarsal diameter;
propodeal dorsum delimited by shallow depression at least laterally; Philippines
browni (Ashmead)
— Abdominal tergum I shiny dorsally, coarsely, sparsely punctate, faintly transversely depressed preapically;
apical bands of abdominal terga II-III yellowish, band on tergum I almost as broad as hindtarsal diameter
and half as wide as band on tergum II; propodeal dorsum not delimited by depressions; southeastern
China ningyuenfuense sp. n.
10(8) Translucent apical bands of abdominal terga I - 1 1 bright golden, of equal width; propodeum as long as wide
(seen from above), with hind surface mostly punctate apically; ocelli (especially in male) small
(OOD>OD); India: Assam differens Turner
Translucent apical bands of abdominal tergum I very narrow or absent; propodeum wider than long (seen
from above), hind surface transversely carinate apically; ocelli larger (OOD<OD=POD) 11
11(10) Abdomen comparatively short, terga II-III twice as wide as long (dorsal view); translucent apical bands of
female tergum I often absent, narrow on terga II-III (hardly broader than diameter of hindtarsomere I),
bright golden; tergum I dull due to dense microsculpture; metapleural flange with narrower, densely
pubescent lamella (Fig. 11); Russian Far East, Korea, eastern China, Japan, United States
koreense (Radoszkowski)
— Abdomen comparatively longer, terga II-III 1 .5 times as wide as long; translucent apical band of tergum
I present, those on terga II-III whitish-yellow; tergum I weakly shiny, with sparse microsculpture;
metapleural flange with broader, spoon-shaped, sparsely pubescent lamella (Fig. 8); southeast Asia,
Malaysia, Indonesia rothneyi Cameron
Pison agile (Smith) P. (Pison) agile: Bohart and Menke 1976:333 (listed).
Figs .3 4 6 10 15 ^ agile. Menke 1988:38 (member of agile group, redescnbed).
Parapison agilis Smith 1869:300. Lectotype: female, "Ceylon" Lectotype Selection.— The natural History Mu-
(now Sri Lanka) (BMNH), present designation. , f_ , . ... ~ ... , . , ...
P. (Paraxon) agile: Kohl 1885:186 (listed). SeUm haS tW0 femaleS Wlth Smith S handwritten
P (Ptsonoides) agilis: Turner 1916:616 (new combination, re- tyPe labels. The first female has the museum white
described). round label (1) with "Type" printed inside a red
122
Journal of Hymenoptera Research
koreense
MAP 2
^^
c?
Fig. 1. Geographic distribution of the Pison agile species group in the Old World.
Fig. 2. Geographic distribution of P. koreense in the New World.
Volume 3, 1 994
Figs. 3-19. Morphological features of Pison species. 3, head in frontal view of P. agile (a, female; b, male). 4-5, clypeus in frontal
view (a, female; b, male): 4, P. agile; 5, P. pulawskii. 6-9, right metapleuron in lateral view (arrow shows metapleural flange):
6, P. agile; 7, P. koreense; 8, P. rothneyi; 9, P. pulawskii. 10-13, submarginal cell area of right forewing: 10, P. agile; 11, P. browni;
12, P. vechti; 13, P. pulawskii. 14, submarginal cell area of forewing of P. agiloides (a, right; b, left). 15-19, abdominal tergite I (a,
lateral view; b, dorsal view): 15, P. agile; 16, P. koreense; 17, P. browni; 18, P. t'fdifi; 19, P. pulawskii.
124
Journal of Hymenoptera Research
oval, a pale-blue round label (2) with "India"
handwritten on the upper side and "56/150" hand-
written in two lines on the under side, a white
rectangular label (3) with "agilis Sm. Type" hand-
written in two lines, and the museum white rect-
angular label (4) with "B.M. TYPE. HYM." printed
in two lines and "21.538" handwritten below.
Another female has a pale-blue round label (1)
with "Ceylon" handwritten on the upper side and
"61 /36" on the under side, a pale-blue rectangular
label (2) with "P. agilis Smith. Type" handwritten
in two lines. Only "Ceylon" was mentioned as the
type locality in the original description so the
"India" type may not be a true type. The Natural
History Museum also has more than a dozen agile
females from "Ceylon" and three of them have the
same pale-blue round labels found on the second
"type." I believe them to be the members of the
type series and I have selected the specimen with
Smith's type label as lectotype and the three other
females as paralectotypes. I am discounting the
Indian specimen as a type.
Discussion. — Pison agile is a member of a sub-
group of species with mainly dark legs. Together
with agiloides sp. n., it differs from other such
species in having comparatively fine
microsculpture of abdominal tergum I in which
the interspaces are polished. The remaining terga
are similarly polished. Pison agile differs from
agiloides in having completely brown mid and
hind legs, sparser and coarser scutal punctures
and narrower whitish translucent apical tergal
bands.
Range. — Known only from southern India and
Sri Lanka.
Material Examined. — 2 males, 18 females. IN-
DIA: Anamalai Hills, Cinchona (BMNH, NMNH).
SRI LANKA: North Western Prov., Kurunegala
Dist, Kurunegala; Central Prov., Kandy Dist.,
Kandy, Udawattakele (BMNH, USNM, ZMUM).
Pison browni (Ashmead)
Figs. 11, 17
Pisonoides browni Ashmead 1905:961. Holotype: male, Philip-
pines, "Manila" (USNM no. 8332), examined.
P. (Pisonoides) browni: Turner 1916:617 (new combination,
listed).
P. (KrombeinieUum) browni: Bohart and Menke 1976:337 (new
combination, listed).
P. (KrombeinieUum) browni: Tsuneki 1983:81 (redescribed)
P. browni: Menke 1988:38 (member of agile group).
Discussion. — This is the darkest species in the
agile group, differing from other members by the
narrow, brown apical bands on terga II-III, by the
strong, transverse preapical depression of tergum
I and the shallow depressions that delimit the
propodeal dorsum. The last character is unique in
the group.
Range. — Philippines.
Material Examined. — 1 male, 2 females.
LUZON: Manila (USNM), Laguna (CAS). Tsuneki
(1983) recorded the species from Bontoc, Luzon;
Mambucal, Negros; Cagayan de Oro, Mindanao.
Pison differens Turner
P. (Pisonoides) differens Turner 1916:617. Lectotype: female,
India, "Shillong, Assam" (BMNH No. 21.540), present
designation.
P. (KrombeinieUum) differens: Bohart and Menke 1976:337 (new
combination, listed).
P. differens: Menke 1988:38 (member of agile group).
Lectohfpe Selection. — Turner described this spe-
cies from three females without selecting a holo-
type. They have two identical rectangular labels:
(1) "Shillong. 5.03" handwritten in two lines and
(2) "Assam. R.Turner. 1905-125" printed in three
lines ("Shillong, Assam, 5000 ft. (Turner), May"
was mentioned in the original description). One
female has in addition Turner's label "Pison
(Pisonoides) differens. Turn. Type." handwritten
in four lines and two museum labels, a round one
with "Type. H.T." printed in two lines inside a red
ring and a white rectangular one with "B.M.
TYPE.HYM." printed in two lines and "21.540"
handwritten below. I have selected the last female
as lectotype and the two others as paralectotypes.
Discussion. — Pison differens is similar to rothneyi
in color of the body and of the translucent apical
tergal bands. The female differs from rothneyi by
its comparatively small lateral ocelli and elongate
propodeum with its hindface punctate (trans-
versely carinate in rothneyi). The male also has
very small ocelli but the propodeum is not elon-
gate. In both sexes the propodeal dorsum is pol-
ished, finely sparsely punctate. The scutum is
finely densely punctate in females. I think that
differens may be a local form of rothneyi but more
material will be required to resolve this.
Range. — Known only from Assam State, In-
dia.
Material Examined. — 1 male, 3 females. IN-
DIA: Assam State, Shillong, IV. 1903, V.1903
(BMNH).
Volume 3, 1 994
125
Pison erythropus Kohl
Parapison rufipes Smith 1869:299. Lectotype: female, "India"
(BMNH No. 21.539), present designation. Nee Pisomtus
rufipes Shuckard 1838:79 (now in Pison).
Parapison rufipes Home, in Home and Smith 1870:165 (biol-
ogy)-
P. rufipes Smith, in Home and Smith 1870:188 (redesenbed).
Pison (Parapison) erythropus Kohl 1885:183 (new name for
Parapison rufipes Smith 1869, nee Ptsonitus rufipes
Shuckard 1838).
P. (Parapison) erythropus Kohl 1885:186 (listed).
P. erythropus: Bingham 1897:221 (listed).
P. Pisonoides) erythropus: Turner 1916:616 (listed).
P. (Pison) erythropus: Bohart and Menke 1976:335 (new com-
bination, listed).
P. erythropus: Menke 1988:38 (member of agile group).
Lectotype Selection. — The Natural History
Museum has three females of this species bearing
round labels with "India" handwritten on the
upper side and "98.69" on the under side. One of
them also has a rectangular label with "rufipes Sm.
Type" handwritten in two lines. I believe them to
be the members of the type series and I have
selected the last female as lectotype and the other
two females as paralectotypes.
Discussion. — Pison erythropus belongs to the
subgroup whose members have brightly colored
legs. This species is easily recognized by its en-
tirely reddish-orange legs (including tarsi and
basal parts of the trochanters).
Biology. — The nests observed by Home (1870)
in northwestern India as Parapison rufipes (Sm.)
included "a mass of loosely arranged cells of earth
attached to some hanging objects, such as a creeper,
tendril, or pendent straw, or even a curled dry
leaf." Home mentioned that the cells were "very
globular" and that their walls consisted of com-
paratively large "pellets. ..loosely attached to one
another." He also "counted eighteen ("smallest
spiders" in Home) in two chambers." Nothing
was reported about the number of cells per nest,
the species or even family of the prey nor was the
cocoon described.
Range. — Known with certainty only from west-
ern India.
Material Examined. — 2 males, 9 females. IN-
DIA (BMNH, ZMUM); Maharashtra District, West-
em Ghats, Lonavale (NMNH).
Pison hissaricum Gussakovskij
P. (Parapison) hissaricum Gussakovskij 1937:622. Holotype:
female, Tajikistan, "Stalinbad" (now Dushanbe) (ZIN),
examined
P. (Pison) hissaricum: Bohart and Menke 1976:336 (new com-
bination, listed).
P. hissaricum: Menke 1988:38 (member of agile group).
Discussion. — Pison hissaricum also belongs to
the subgroup whose members have brightly col-
ored legs. This species can be recognized by the
following combination of features: legs except
trochanters, tarsi and tibiae reddish apically; ter-
gum I dull due to dense microsculpture of
interspaces, punctures of tergum I dense; and
whitish translucent apical bands of terga I-III nar-
row.
Range. — Known from Uzbekistan and
Tajikistan.
Material Examined. — 8 males, 10 females.
UZBEKISTAN: Aman-Kutan (ZMUM). TAJI-
KISTAN: Dushanbe; Varzob Valley, Kondara (ZIN,
ZMUM).
Pison koreense (Radoszkowski)
Figs. 7, 16
Paraceramius koreensis Radoszkowski 1887:433. Holotype: fe-
male, "Koree" (Krakow, Poland. Lost). Neotype: fe-
male, Russia, Primorskiy Kray (ZMUM), present desig-
nation.
Ceramius koreensis: Morawitz in Dalla Torre 1894:3 (new
combination, listed in Vespidae).
Pison (Parapison) koreense: Kohl in Dalla Torre 1897:712 (new
combination, listed in Crabroninae).
Pison (Pisonoides) koreensis: Turner 1916:617 (new combina-
tion, listed).
P. (Pisonoides) koreensis: Yasumatsu 1935:229 (listed).
P. (Parapison) koreense: Gussakovskij 1937:622 (listed).
P. (Pisonoides) koreensis: Yasumatsu 1939:83 (listed).
P. (Paraceramius) koreense Krombein 1958a:166 (first record
from United States).
P. (Paraceramius) koreense: Krombein 1958b:189 (listed).
KrombemieUum koreense: Richards 1962:118 (new name for
Paraceramius Radoszkowski 1887, nee Saussure 1854).
P. (Pisonoides) koreensis: Iwata 1964:1 (biology).
P. (KrombemieUum) koreense: Krombein 1967:394 (new combi-
nation, listed).
P. (Krombemiellum) koreense: Menke 1968a:7 (listed).
P. (KrombemieUum) koreense: Menke 1968b: 1102 (redesenbed).
P. koreense: Sheldon 1968:107 (biology, larval morphology).
P. (KrombemieUum) koreense: Hawkins 1974:279 (biology).
P. (KrombemieUum) koreense: Bohart and Menke 1976:337
(listed).
P. (KrombemieUum) agile: Krombein, et al 1979:1641 (synony-
mized).
126
Journal of Hymenoptera Research
P. koreense: Kazenas 1980:92 (listed).
P. agile: Menke 1988:38 (listed).
Neototype Selection. — At my request, A. P.
Rasnitsyn recently searched for the type oikoreense
in Radoszkowski's collection in Krakow, Poland
but could not find it. I presume it was lost and I
have selected a specimen from the southern part of
Primorskiy Kray (32 km SE Ussurijsk, 25. VIII. 1986;
eastern Russia not far from Korea)as the neotype.
Discussion. — Pison koreense is the best known
member of the agile group because its biology has
been studied both in its native habitat in Japan
(Iwata 1964) and in the United States (Sheldon
1968) where it was introduced apparently after
World War II (Krombein 1958a).
Krombein (1979) synonymized koreense with
agile and Menke (1988) accepted this synonymy.
After comparing the syntypes of agile with mate-
rial from Sri Lanka, Japan, Russian Far East and
the United States, I have concluded that koreense is
a distinct species. It differs from agile in having a
dull tergum I that contrasts with a shiny tergum II
(tergum I shiny in agile). Pison koreense also differs
from agile in having bright golden translucent
apical bands on terga II-III (whitish in agile), largely
yellowish tibiae (brown in agile) and a transversely
carinate propodeal hind surface (mostly punctate
in agile). The metapleural flange of koreense is broad
and reddish (Fig. 7); it is narrow and dark in agile
(Fig. 6). Pison koreense differs from differens and
rothneyi in having a more compact abdomen and a
weakly developed translucent apical band on ter-
gum I.
Biology. — The wasps were studied in detail in
Japan by Iwata (1964) and in North America by
Sheldon (1968) who also provided a detailed de-
scription of the immature stages of koreense. Both
Iwata and Sheldon observed that koreense con-
structed small (6.0-10.2 X 4.0-6.0 mm), fragile,
cylindrical mud cells with finely cemented walls.
The cells are placed separately or in groups of up
to 21 independent cells. The cells are attached in
various protected places: inside a photographic
tank (Krombein 1958a); in a culvert, in small de-
pressions and cracks and under a bridge (Sheldon
1968); in the nooks of a mud wall under eaves
(Iwata 1964); and even within the old empty cells
of Trypoxylon (Trypargilmn) politum (Say) (Sheldon
1968; Hawkins 1974). Construction of cells on
plants, as in erythropns, is unknown. The cells were
provisioned with six immature spiders of the ge-
nus Aranens (Araneidae) (Iwata 1964) or by 20-31
small spiders of the genus Dictyna (Dictynidae),
including males and females of D. bellans
Chamberlin and D. sublata Hentz (Sheldon 1968).
P. formise cocoons are cylindrical (6.0-9.1 X 2.0-3.6
mm) and, according to Sheldon, were surrounded
by "delicate silken threads attaching cocoon to
inside of cell." Sheldon also mentioned a large
number of adult Melittobia chalybii Ashmead
(Eulophidae) which were reared from koreense
cocoon "from a nest constructed in an unsealed
cell of Trypoxylon politum."
I also observed nesting and hunting activities
of koreense during 1983-1986 in the Primorskiy
Kray in the Far East of Russia. The females made
their small clay cells between logs in the wall of an
old rural shed, placing them in groups of up to 20
independent cells. Five to seven specimens of
immature spider of the genus Araneus were put
into each cell. When hunting the female of koreense
looked for the prey on their webs and attacked
them from the dorsum, embracing the carapace
and applying a single sting to the venter of the
spider. After paralyzing the spider, the female
turned the prey venter up, clutched the base of its
chelicerae with her mandibles and flew with it to
her cell. I observed egg-laying only once and it was
the last operation before sealing the cell. Sheldon
(1968) also mentioned that the egg was attached to
the abdomen of the last spider in the cell.
Thus, populations of koreense in Russian Far
East, Japan and the United States have similar
biological features except prey. Spiders of the
family Dictynidae were preferred in North America
and Araneidae in eastern Asia.
Range. — Russian Far East, Korea, Japan, east-
ern China, United States: Illinois, Kansas, Mary-
land, Michigan, New York, Virginia, Wisconsin.
Material Examined. — 28 males, 18 females.
RUSSIA, Primorskiy Kray: Sichote-Alin, Vangou;
32 km SE Ussurijsk; Partizansk; 70 km ENE
Partizansk (CAS, USNM, ZMUM). CHINA,
Zhejiang: Hangchow (USNM). JAPAN, Fukui, Mt.
Haku: Ichinose; Chugu; Mie (USNM, ZMUM).
USA: Kansas, Lawrence; Virginia, McLean
(USNM).
Volume 3. 1994
127
Pison rothneyi Cameron
Fig. 8
P. (Parapison) rothneyi Cameron 1897a:81. Holotype: female,
India, West Bengal State, "Barrackpore" (now Barakpur)
(OUM), examined.
P. (Parapison) crassicorne Cameron 1897a:25. Holotype: male
(female in original description), India, West Bengal State,
"Barrackpore" (now Barakpur) (OUM), examined. Syn-
onymy by Turner (1916:617).
P. (Pisonoides) rothneyi: Turner 1916:617 (new combination,
listed).
P. (Krombeiniellum) erythropus: Tsuneki 1974:637 (new combi-
nation, misidentified).
P. (Pison) rothneyi: Bohart and Menke 1976:336 (listed).
P. rothneyi: Menke 1988:38 (member of agile group).
Discussion. — This species is similar to differens
and also koreense, differing from the first by the
narrower translucent whitish-yellow apical bands
of the terga, by the comparatively larger ocelli and
by the shorter propodeum whose dorsum is dis-
tinctly punctate and the hind surface transversely
carinate apically . Also, rothneyi has a more coarsely
sculptured scutum and deeper impressed
parapsidal lines. Pison rothneyi differs from koreense
by the comparatively longer abdominal terga with
well developed whitish-yellow translucent bands
on all terga, including tergum I. Pison rothneyi also
has a broader, spoon-shaped reddish-orange
metapleural flange (Fig. 8) and its tergum I is shiny
(semidull in differens and koreense). Tsuneki (1974)
misidentified specimens of rothneyi as erythropus.
The dark femora and tarsi of his specimens con-
firm this.
Range. — This is the most widely distributed
Oriental member of the agile group. It is recorded
from eastern India, Thailand, Viet Nam, Malaysia
and Indonesia.
Material Examined. — 1 male, 13 females. IN-
DIA, West Bengal: Barrackpore (OUM). THAI-
LAND, Chieng Mai Province: Fang Horticultural
Exp. Station; Doi Inthanon N. P.: Huai Sai Luang
(ZMK). Malaysia: Teluk Merban (ZMK); Kuala
Lumpur (BMNH). VIET NAM: Tonkin, Poste de
Dong-Dang (ZMUM). INDONESIA, Sumatra:
Pakanbaru, Solok; Java: Bogor, Semarang
(NMNH).
Pison agiloides Antropov, new species
Fig. 14
Description of Holotype Female. — Black except
the following reddish: palpi, mandible largely,
labrum, tibiae except apically, femora apically,
pronotal lobe posteriorly, tegula and metapleural
flange; translucent apicalbands of abdominal terga
I-V and sterna II-V, and terga I-V laterally yellow-
ish-white.
Clypeus, frons beneath, pronotum anteriorly,
scutellum, metanotum and propodeum with dense
erect silvery pubescence, longest setae on
propodeum laterally and posteriorly; pronotal
collar, scutum, mesopleuron and abdomen with
suberect, dense, short pubescence, longest setae
on apical band of tergum I.
Labrum slightly emarginate apically, inner
orbits almost parallel, OOD<OD.
Frons densely punctate, interspace smooth,
shiny; scutum densely, finely punctate, punctures
separated by less than a diameter; scutellum,
mesopleuron, metanotum and propodeum more
sparsely punctate, punctures 1-3 diameters apart,
interspaces polished; propodeal dorsum not en-
closed by sulcus or carina; metapleural flange
broad, spoon-shaped, translucent, with long sil-
very setae.
Right forewing (Fig. 14a) with two, left forew-
ing (Fig. 14b) with three submarginal cells, outer
submarginal cell of forewings open distally; recur-
rent vein II ending on tiny submarginal cell II (left
forewing) and interstitial on right forewing.
Abdomen simple; terga shiny, densely punc-
tate, diameter of punctures decreasing from basal
to apical terga; tergum I with microsculpture,
other terga more polished between punctures;
sternum I similar to frons, other sterna densely,
finely punctate, shiny; translucent apical bands of
abdominal segments 1 .5 to twice as broad as diam-
eter of first hind tarsomere.
Male. — Unknown.
Discussion. — The new species differs from ag-
ile in having the dense, fine punctation of the
scutum and abdomen, the longer propodeum, the
comparatively short setae of the thorax and abdo-
men, the broad translucent apical bands of the
abdominal segments and the reddish tibiae and
metapleural flange. The last features are shared
with rothneyi, but the latter has completely reddish
tibiae, tergum I is shiny and the propodeum is
elongate. Pison differens also has fine, dense scutal
punctures, but it differs from agiloides by its larger
OOD, by t its dark-brown tibiae and by the weakly
shiny surface of tergum I.
The open second submarginal cell of the unique
128
Journal of Hymenoptera Research
specimen of agiloides may prove to be individual
when more material is available. The presence of
the true second submarginal cell in the left forew-
ing is likely to be atypical also, but it illustrates that
the loss of this cell is achieved via diminution of its
size in the agile group (see also Menke 1988).
Etymology. — The ending of the species name
emphasizes the likeness of this species and agile.
Range. — Sri Lanka.
Type.— Holotype female: SRI LANKA,
Sabaragamuwa Province, Ratnapura District,
Belihuloya Resthouse, 9.IV.1978, M. D. Hubbard,
T. Wijesinhe (USNM).
Pison ningyuenfuense Antropov, new species
Description of Holotype Female. — Black except
the following reddish: palpi, mandible largely,
labrum, foretibia anteriorly, midtibia basally,
hindtibia basally and posteriorly and all spurs;
apical rim of clypeus, tarsi ventrally and tegula
reddish-brown; metapleural flange brown; trans-
lucent apical bands of abdominal terga I-V whit-
ish.
Pubescence silvery; clypeus, frons, vertex,
pronotum, scutum posteriorly, scutellum, met-
anotum and porpodeum with erect setae, longest
on pronotum laterally and on propodeum; gena,
mesopleuron and abdomen with suberect, dense,
short setae, those on gena and lateral angles of
abdominal tergum I longest; scutum with ex-
tremely short, erect setae as on femora.
Pronotum and scutum finely and densely
punctate, shiny, punctures less than a diameter
apart; mesopleuron, scutellum and metanotum
more sparsely punctate (punctures 1-2 diameters
apart); propodeum coarsely punctate (punctures
at least twice as large as those on scutum); abdomi-
nal tergum I moderately coarsely, sparsely punc-
tate (punctures similar in size to those on
propodeum), interspaces weakly shiny due to
dense microsculpture; sternum I similarly punc-
tate but shiny; following abdominal segments
densely, finely punctate (like scutellum), shiny.
Translucent apical bands of abdominal terga
II-V as broad as diameter of first hind tarsomere,
bands of tergum I and sterna II-V less than diam-
eter of first hind tarsomere; apex of tergum I
shallowly but distinctly depressed (depressed part
as broad at middle as maximal diameter of hind
tibia); terga II-IV laterally with rounded tubercles.
Recurrent vein I ending on submarginal cell I,
recurrent vein II ending on submarginal cell II
near its base (nearly interstitial ).
Length 7.6 mm.
Male. — Unknown.
Discussion. — Pison ningyuenfuense differs from
the most similar browni by having an undefined
propodeal dorsum (no limiting sulcus or carina),
by the comparatively shallowly depressed apical
part of abdominal tergum I, which is more coarsely
punctate, with interspaces only weakly shiny, and
by the whitish translucent apical bands of terga I-
V.
Etymology. — The species name is derived from
the native name of the holotype locality.
Range. — Known only from the type locality in
southeastern China.
Type. — Holotype female: CHINA, Hunan
Province, Ningyuenfu, July 24-26-28, alt. 600-
10,800, D. C. Graham (USNM).
Pison vechti Antropov, new species
Figs. 12, 18
Description of Holotype Female. — Black except
the following bright reddish-orange: palpi, man-
dible largely, fore and mid tibiae and fore femur
completely, mid and hind femora except dorsum
and hind tibia except posteriorly, tegula and
metapleural flange posteriorly, apical band on
abdominal tergum I and medial spots on terga II-
III before translucent apical bands; pronotal lobe
posteriorly and spical tarsomeres beneath yellow-
ish; translucent apicalbands of terga I-V and sterna
II- VI bright golden.
Pubescence of head and thorax silvery, mainly
golden on abdominal terga (especially on translu-
cent bands laterally); frons, vertex, gena, prono-
tum, scutum, scutellum, mesopleuron and met-
anotum with dense, short, erect setae (not longer
than mid ocellus diameter); propodeum
posteriolaterally with longer dense, erect setae
(almost twice as long as mid ocellus diameter);
abdomen with suberect or almost appressed dense,
short setae.
Labrum truncate apically; median lobe of
clypeus obtusely angulate, with narrow, shiny
apical margin; inner orbits almost parallel;
OOD<OD=POD.
Clypeus, frons, pronotum and scutum densely,
minutely punctate, weakly shiny (punctures
Volume 3. 1 994
129
mostly less than a diameter apart); scutellum,
mesopleuron and metanotum with sparses punc-
tation (punctures 1.5-2.0 diameters apart),
interspaces polished; metaplaural flange broad,
spoon-shaped, translucent posteriorly; propo-
deum posteriorly and laterall punctured like
mesopleuron but obviously coarser; propodeal
dorsum shiny, not enclosed by sulcus or carina,
with sparse punctures (2-4 diameters apart) and
with deep median furrow containing simple ca-
rina on basal half; abdominal tergum I punctate
(punctures 1-3 diameters apart), densely
microsculptured, weakly shiny, sternum I, ter-
gum and sternum II densely, finely punctate (punc-
tures of sternum I coarsest), shiny; other abdomi-
nal segments densely micropunctate, surface
smooth, weakly shiny; translucent apical bands of
abdominal terga II-IV as braod laterally and medi-
ally as hindtibial diameter (those of terga I and V
and sterna II-IV 1.5 times diameter of first
hindtarsomere (Fig. 18).
Recurrent vein I ending on submarginal cell I,
recurrent vein II ending near base of submarginal
cell II, the latter very narrow anteriorly, almost
triangular (Fig. 12).
Length 7.6 mm.
Variation in Females (18 specimens). — Femora
and tibiae all bright reddish-orange in two females
from Buitenzorg, Java (28. VI. 1932) and
Tembajangan, Kangean Island all femora dark
posteriorly in a female from Mulie, Java, and sub-
marginal cell II completely triangular in another
female from Buitenzorg (16.X.1941). Length 7.0-
7.8 mm.
Male. — As in female except: median lobe of
clypeus narrower, acutely prominent apically;
punctation sparser on scutum (punctures 1-2 di-
ameters apart) and mesopleuron (punctures 2-4
diameters apart). Length 5.4 mm.
Discussion. — Pison vechti is very similar to
hissaricum, erythropus and differens in having wid-
ened metapleural flange and largely bright red-
dish-orange legs. It is easily separated from
hissaricum and erythropus in having the broad,
golded, translucent apical bands of the terga. Fur-
thermore, it differs from erythropus in having dark-
brown trochanters and tarsi and a densely punc-
tate, weakly shiny scutum. Pison vechti differs
from differens in having the larger lateral ocelli and
a comparatively short propodeum. The color of
the abdomen of vechti is similar to rothneyi, but the
latter has mainly brown femora and tibiae. Pison
pulaivskii also has bright reddish-orange legs but
differs from all mentioned species in having a
narrow metapleural flange.
Etymology. — This species is dedicated to
Jacobus van der Vecht.
Range. — Malaysian Peninsula and southeast-
ern Asian islands (Malaysian and Indonesian).
Types.— Holotype female: INDONESIA, E.
Java, Idjen, Plateau Blawan, 900 m, 28.V.1939, J. v.
d. Vecht (NMNH). Paratypes (1 male, 19 females):
MALAYSIA, Kuala Lumpur: 27. X. 1929, H.
Pendlebury (NMNH); Tanglui Road, 20. V.1928, T.
T. Pagden (BMNH). INDONESIA, Sumatra: Fort
de Kock, 920 m, 1926, E. Jacobson ( AUZM); Sibolga,
1 42'N-98 48'E, VJVIII. 1954, W. Vergeest (NMNH);
Java: Buitenzorg, J. v. d. Vecht; Buitenzorg,
16.X.1941, 11.1953, J. v. d. Vecht; Buitenzorg,
Pebaton, 28.VI.1932, J. v. d. Vecht; Buitenzorg,
Jnsl. v. Pin., 15.VI.1929, J. v. d. Vecht; Malang,
IV. 1933, Betrem; Bogor, 1955, Hamann; Mulie; W.
Preanger, Z. Soekaboemi, IV. 1933, J. v. d. Vecht;
Ambarawa, Lundeking (NMNH); Malang, IV.1933,
Betrem (ZMUM); Mt. Tijoeng, Djampang Tengah,
1.1939, K. M. Walsh (BMNH); Kangean Isl.:
Tembajangan, 11.1936, M. E. Walsh (NMNH).
Pison chrysoptilum Antropov, new species
Description of Holotype Female. — Black except
the following reddish-orange: palpi, mandible
except apically, labrum, pronotal lobe posteriorly,
tegula mainly, widened part of metapleural flange,
all femora except posteriorly and tibiae except
apically; translucent apical bands of abdominal
terga I-V golden.
Pubescence silvery on clypeus, mesopleuron,
propodeum laterally and abdominal sterna, golden
on frons, vertex, gena, pronotum, scutum, scutel-
lum, metanotum, propodeal corsum and hindface
and abdominal terga. Clypeus, pronotal collar,
propodeal dorsum and terga with suberect or
decumbent dense setae (shortest on abdomen,
longest on pronotum); other areas with erect setae,
shortest on scutum and abdominal sterna, longest
on metanotum and propodeum posterolaterally.
Median clypeal lobe rounded, with narrow
semitranslucent brown apical margin; inner orbits
of eyes converging above; OD>POD>OOD.
Clypeus, frons, vertex, pronotum, scutum,
scutellum and mesopleuron finely, densely punc-
130
Journal of Hymenoptera Research
tate (punctures 0.5-1.5 diameters apart), shiny;
metapleural flange broad, spoon-shaped, widened
part translucent; propodeum more coarsely,
sparsely punctate laterally and dorsally (at least 2
diameters apart), hindface coarsely and densely
punctate (0.5-1 .0 diameters apart), with transverse
ridges ventrallv; propodeal dorsum not enclosed
by sulcus or carina, with narrow median furrow
containing simple carina; tergum I punctate (l-#
diameters apart), shiny in spite of microstriahon;
sternum I with coarser punctures (as on propodeal
hindface), shiny; other abdominal sclerites micro-
scopically but distinctly, densely and uniformly
punctate (approximately a diameter apart), shiny;
translucent apical margins of terga I-IV twice as
broad as diameter of first hind tarsomere (on
tergum V and sterna III-V slightly less than diam-
eter).
Recurrent vein I ending on submarginal cell I,
recurrent vein II ending near base of submarginal
cell II.
Length 8.2 mm.
Variation in Females (2 specimens). — Paratype
female differs by its smaller size (6.2 mm) and
almost parallel inner orbits of eye.
Male. — Unknown.
Discussion. — Unlike all other members of the
agile group, chrysopdlwn has golden pubescence
on the frons, vertex, thorax and propodeum. It
resembles vechti because of its brightly colored
legs and golden translucent bands on the abdomi-
nal terga. Pison chrysoptilum and vechti may be two
forms of one species, but more material will be
necessary to resolve this.
Etymology. — This species name is derived from
the Greek words din/sos (=gold) and ptilon (=down,
fluff) emphasizing the color of the pubescence.
Range. — Northern Borneo (Sarawak, Brunei).
Types. — Holotype female: MALAYSIA,
Sarawak: 4th div., Gn. Mulu, RGS Exp., 17.IX-
23.X.1977, D. Hollis (BMNH). Paratype (1 female):
BRUNEI: Ulu Temburong, Base camp hut, 300 m,
115 16'E,4 26'N, 16.II-9.III.1982, M. C. Day (BMNH).
Pison pulawskii Antropov, new species
Figs. 5, 9, 13, 19
Description of Holotype Female. — Black except
the following reddish-orange: palpi, mandible
largely, labrum, fore legs distad of coxal apex, mid
and hind legs except basal parts of coxae and
tarsomeres III-V; antennal articles beneath, apical
margin of clypeus, tegula, mid and hind tarsomeres
III-V reddish-brown.
Pubescence silvery only; clypeus, frons, ver-
tex, pronotal collar, scutum, scutellum, metano-
tum, metapleuron and propodeum with erect se-
tae, longest on propodeum posterolaterally and
shortest on scutum (scarsely longer than on
femora); gena, mesopleuron and abdomen with
suberect or appressed short setae (longest on api-
cal bands of terga and shortest on sterna).
Labrum truncate apically; clypeus compara-
tively broadly rounded (Fig. 5A); inner orbits of
eyes almost parallel; OOD=OD=POD.
Frons densely punctate, shiny; scutum very
densely, finely punctate, weakly shiny;
mesopleuron uniformly, sparsely punctate (1.0-
1.5 diameters apart), polished; metapleural flange
not lamellate (Fig. 9); propodeal dorsum obliquely
carinate basally and along median furrow, dor-
sum not enclosed by sulcus or carina, sparselv
punctate (as on propodeal side) (1-3 diameters
apart); tergum I uniformly, densely ounctate (as
on mesopleuron), shiny; tergum II more densely
and finely punctate, shiny; remaining terga weakly
shiny because of very dense microscopic punc-
tures; sternum I dull, punctured as frons; sternum
II shiny, punctate as mesopleuron; other sterna
with dense micropunctures, weakly shiny; trans-
lucent apical bands of terga I-V and sterna II-IV
whitish, as broad as diameter of hindtarsomere I
(Fig. 19).
Recurrent vein I ending on submarginal cell I,
recurrent vein II ending on submarginal cell II
near its base (Fig. 13).
Length 7.0 mm.
Variation in Females (11 specimens). — Mid tarsi
completely reddish-orange in three specimens
from holotype locality and one from
Kurumbagaram; hind tibiae and tarsi brownish
posteriorly and antennal flagellum completely
dark-brown in one specimen from Coimbatore.
Length 6.0-7.0 mm.
Males (3 specimens). — As in females except:
clypeus with acute median prominence spicallv
(Fig. 5b); labrum slightly emarginate; mandibles
without inner teeth. Length 4.8-5.2 mm.
Discussion. — The nonlamellate metapleural
flange of pulawskii sets the species apart from other
in the agile group. Other features of the species are
the form of the clypeus, the mostly reddish-orange
Volume 3. 1994
131
legs and the narrow whitish translucent apical
bands of the abdominal segments. Reddish legs
occur in erythropus and whitish abdominal bands
occur in erythropus, agile and ningyuenfitense, but
these species have a lamellate metapleural flange.
Etymology. — This species is dedicated to
Wojciech J. Pulawski.
Range. — Western and southern India.
Types. — Holotype female: INDIA, Rajasthan:
Uddaipur (24 35'N), 27.V.1989, W. J. Pulawski
(CAS). Paratypes (3 males, 9 females): same place,
date and collector as holotype (CAS); same place
and collector as holotype, 21-25. V. 1989 (CAS,
ZMUM); Gujarat: Disa (=Deesa), 4-6. VI. 1989, W. J.
Pulawski (CAS); Karnataka: Bangalore, 915 m,
26.V.1980, 30.V.1980, K. D. Ghorpade (ZMK);
Karikal Territory: Kurumbagaram, III.1947, P. S.
Nathan (USNM).
ACKNOWLEDGMENTS
I express my gratitude to all colleagues who lent mate-
rial for study. I am grateful to Dr. A. P. Rasnitsyn for his help
in searches for the type specimen of koreense in Radoszkowski's
collection. I am extremely grateful to Dr. A. S. Menke who
read earlier versions of the manuscript and suggested many
important improvements of the text and who helped me in
the English. Finally, I thank Dr. K. V. Krombein and Dr. W. J.
Pulawski who critically and attentively reviewed the entire
manuscript and who provided manv useful comments.
LITERATURE CITED
Ashmead, W. H. 1905. Additions to the recorded Hymenopter-
ous fauna of the Philippine Islands, with descriptions of
new species. Proceedings of the United States National
Museum 28(1413): 957-971.
Bingham, C. T. 1987. The fauna of British India. Hymenoptera,
vol. 1. Taylor and Francis, London. 179 pp.
Bohart, R. M. and A. S. Menke. 1976. Sphecid wasps of the
World, a generic revision. University of California Press,
Los Angeles, London, iv + 695 pp.
Cameron, P. 1897a. IV. Hymenoptera Orientalia, or contribu-
tions to a knowledge of the Hymenoptera of the Oriental
zoological region. Part V. Memoirs and Proceedings of the
Manchester Literary & Philosophical Society 41(2): 1-144.
Dalla Torre, C. G. 1894. Catalogus Hymenopterorum hucusque
descriptorum systematica et synonymicus. Vol. IX. G. En-
gelmann, Lipsiae. 181 pp.
Dalla Torre, C. G. 1897. Catalogus Hymenopterorum hucusque
descriptorum systematica et synonymicus. Vol. VIII. G.
Engelmann, Lipsiae. 750 pp.
Gussakovskij, V. V. 1937. Especes palearctictiqques des genres
Didmeis Wesm., Pison Latr. et Psen Latr. (Hymenoptera,
Sphecodeal). Travaux de I'Institut Zoohgie de I ' Academie
des Sciences de I'URSS 4(3-4): 599-695.
Hawkins, W. A. 1974. A western record for an introduced
Pison (Hymenoptera, Sphecidae) journal of the Kansas
Entomological Society 47(2): 279.
Home, C and F. Smith. 1870. Notes on the habits of some
hymenopterous insect from the north-west provinces of
India. With an appendix, containing descriptions of
some new species of Apidae and Vespidae collected by
Mr. Home. Transactions of the Zoological Society of London
7(3): 161-196.
Iwata. K 1964. Ethological notes on four Japanese species of
Pison (Hymenoptera, Sphecidae). Mushi 38: 1-6.
Kazenas, V. L. 1980. Materials to the fauna of digger wasps
(Hymenoptera, Sphecidae) of the Far East of the USSR.
Taxonomy of insects of the Far East, Vladivostok, pp. 80-94.
Kohl, F. F. 1885. Die Gattungen und Arten der Larnden
Auctorum. Verliandlungen derk. k. zoologisch-botamschen
Gesellschaft in Wien 34: 171-268.
Krombein, K. V. 1958a. Pison tParaceramius) koreense (Rad.), a
new adventive wasp in the eastern United States. Enfo-
mologtcal News 69: 166-167.
Krombein, K. V. 1958b. Superfamily Sphecoidea. In:
Krombein, K. V., et al., Hymenoptera of America North of
Mexico - Synoptic Catalog. Agriculture Monograph 2,
First Supplement, U. S. Department of Agriculture,
Washington, DC. pp. 186-204.
Krombein, K. V. 1967. Superfamily Sphecoidea. In: Krombein,
K. V., B. D. Burks et al., Hymenoptera of America North of
Mexico — Synoptic Catalog. Agriculture Monograph 2,
Second Supplement, U S. Department of Agriculture,
Washington, DC. pp. 386-421.
Krombein, K. V., P. D. Hurd, D. R. Smith and B. D. Burks.
1979. Catalog of Hymenoptera in American North of Mexico.
Vol. 2. Smithsonian Institution Press, Washington, DC.
pp. 1199-2209.
Menke, A. S. 1968a. New genera and species of wasps of the
tribe Trypoxylonini from the neotropical region (Hy-
menoptera: Sphecidae: Larnnae). Los Angeles County
Museum Contributions in Science 135: 1-9.
Menke, A. S. 1968b. A review of the New World species of
Pison. I. The subgenus Krombeiniellum (Hymenoptera:
Sphecidae). The Canadian Entomologist 100(10): 1100-
1107.
Menke, A. S, 1988. Pison in the New World: a revision
(Hymenoptera: Sphecidae. Trypoxylonini). Contribu-
tions of the American Entomological Institute 24(3): 1-171.
Radoszkowski, O. 1887. Hymenopteres de Koree. Horae
Scoitatis Entomologicae Rossicae 21(3-4): 428-436.
Richards, O. W. 1962. A revisional study of the masarid wasps.
Wm. Clowes and Sons, London. 294 pp.
Sheldon, J. K. 1968. The nesting behavior and larval morphol-
ogy of Pison koreense (Radoszkowski). Psyche 75: 107-
117.
Shuckard, W. E. 1838. Descriptions of new exotic aculeate
Hymenoptera. Transactions of the Entomological Society of
London 2(1): 68-82.
Smith, F. 1869. Descriptions of new species of the genus Pison,
and a synonymic list of those previously described.
Transactions of the Entomological Society of London. 9: 289-
300.
Tsuneki, K. 1974. A contribution to the Knowledge of
Sphecidae occuring in Southeast Asia (Hym.). Polskie
132
Journal of Hymenoptera Research
Pismo Entomologiczne 44(3): 585-660.
Tsuneki, K. 1983. Further studies on the Larrinae of the
Philippine Islands, with remarks on the Indian species
of the genus Lyroda (Hymenoptera, Sphecidae). Special
Publications of the japan Hymenopterists Association 24: 1-
116.
Turner, R. E. 1916. Notes on the wasps of the genus Pison and
some allied genera. Proceedings of the Zoological Society of
London 1916: 591-629.
Yasumatsu, K. 1935. The genus Pison Spinola of the Japanese
Empire (Hymenoptera, Trypoxylonidae). Annotationes
zoologicae Japonenses 15(2): 227-238.
Yasumatsu, K. 1939. Notes supplementaires sur le genre
Pison Spinola du Japon (Hymenoptera, Trypoxylonidae).
Festschrift zum 60 Geburtstage von Professor Dr. Embnk
Strand, Riga 5: 81-84.
J. HYM. RES.
Vol. 3, 1994, pp. 133-143
Colony Densities and Preferences for Nest Habitats of Some Social Wasps in
Mato Grosso State, Brazil (Hymenoptera, Vespidae)
IVONE R. DiNIZ AND KlNITI KlTAYAMA
Departamento de Zoologia, Institute de Biologia
Universidade de Brasilia, 70910-900 Brasilia DF, BRAZIL
Abstract — Studies of colony densities and preferences for nest habitats of some social wasps were conducted in cerrado
vegetation in Southern Mato Grosso state in Central Brazil. Wasp colony densities were estimated in three habitats (campo
umido, cerrado sensu-stricto and gallery forest). Regarding the wasp nest habitats preferences, we found 100 colonies of 30
species in 15 genera nesting in six different habitats in Southern Mato Grosso. In Northern Mato Grosso, based on Richards
(1978), we found 199 colonies of 51 species in 14 genera nesting in six different habitats. Considering both regions together,
we came out with 299 colonies of 61 species in 16 genera nesting in nine different habitats.
INTRODUCTION
METHODS AND STUDY AREA
The effectiveness and efficacy of predation by-
social wasps on many insects (Gobbi et al. 1984,
Gobbi and Machado 1985, Machado et al. 1987,
Raw 1988) confers on them a fundamental impor-
tance as biological control agents. Nonetheless,
the study of the nests of these insects has focused
mainly on their architecture (Jeanne 1975, Kojima
1982) and on the numbers of individuals per colony
(Richards 1978), with few reports on pairs of spe-
cies nesting close together (Windsor 1972, Starr
1988).
Information on preferences for nest habitats
(Richards 1978, Reed and Vinson 1979, Forsyth
1980) and on colony densities (Rau 1942, Kitayama
et al. 1989) of neotropical social wasps is still very
scarce. In central Brazil, 130 species of wasps have
been collected (Richards 1978, Raw pers. comm.
and our own collections), but studies on their
ecology are very fragmentary.
The aim of this report is to add new informa-
tion on colony densities and preference for nest
habitats of social wasps in the cerrado sensu-lato of
Central Brazil.
The study was conducted in cerrado vegeta-
tion during a five week survey (August, October
through December 1988 and June 1989) at the Rio
Manso Hydroelectric Power Station (HPS-Rio
Manso) (14°52' S and 55°50' W), in the Chapada
dos Guimaraes county, southern Mato Grosso
State in central Brazil.
Cerrado sensu-lato is a semideciduous
xeromorphic vegetation dominant in Central Bra-
zil, occupying about 20% of the whole country. It
occurs in various structures from closed forest-
like forms to pure grasslands (Eiten 1972). "Habi-
tat" is used here to refer to a vegetation subtype
within the cerrado sensu-lato.
The censuses of social wasp colonies (nests
with resident adults) were conducted in campo
umido, vereda, campo sujo, cerrado sensu-stricto,
gallery forest and surrounding dirt roads. Charac-
terizing these habitats very briefly it could be said
that campo umido is a wet grassland without
visible woody plants (Fig. 1); vereda is a broad
marshy valley bottom grassland with buriti palm
(Mauritia vinifera) galleries (Fig. 2); campo sujo is a
cerrado grassland with a few, very scattered, low
conspicuous shrubs or acaulescent palms (Fig. 3);
134
Journal of Hymenoptera Research
AiiP*:'
=*#■;*>
a-- S^^Ss
M 1 K?
Fig. 1. Area of 100 nv of campo umido where wasp nests were surveyed.
Fig. 2. Area of Vereda (Photo by Dr. Roberto Cavalcante)
135
1 v^g»HK
Fig. 3. Area of campo sujo.
Fig. 4. Area of cerrado sensu-stricto (Photo by Dr. Roberto Cavalcante).
136
Journal of Hymenoptera Research
'4MB£^4<
Fig. 5. Area of gallery forest (Photo by Dr. Roberto Cavalcante).
cerrado sensu-stricto includes forms with the total
woody plant cover of about 30-40% and with a
canopy generally less than seven meters (Fig. 4)
and gallery forest is usually a narrow evergreen
mesophytic forest following the streams (Fig. 5)
(Eiten 1972).
Searches for nests were conducted in four
different months, covering two seasons: dry sea-
son (June and August) and wet season (October
and December) in six habitats. Each habitat re-
ceived the same intensity of searching during each
of these seasons.
From each nest, adult wasps were collected to
guarantee identification of the species. All the
nests and insects sampled are deposited in the
Laboratorio de Zoologia, Departamento de
Zoologia in the Universidade de Brasilia.
The data on Xavantina and Serra do Cachimbo
(10°50'S and 51°47'W), in northern Mato Grosso
state were compiled from Professor Richards'
(1978) species descriptions of nest habitats using
only the data where there was no doubt on the
location of the nests). Richards' censuses were
conducted in campo sujo, dirt roads and clearings,
cerrado sensu-stricto, Cerradao (the medium tall
arboreal form of cerrado with a closed or semi-
open canopy with 30-40% tree crown cover, Eiten
1972), gallery forest and dry forest (deciduous and
semi deciduous mesophytic forest, Eiten 1972).
For the study on colony density, known areas
were sampled, in gallery forest (7500 m2) and in
cerrado sensu-stricto (10000 m2). Nests were lo-
cated among leaves, on branches, on the trunks of
trees and in holes in tree-trunks and in the ground.
Binoculars were used to locate nests in the canopy.
In addition, five squares of 100 m2 each were
surveyed in campo timido, where nests of wasps
were sought among tufts of grasses and sedges
(Fig.l).
The survey for estimates of the density of
colonies, within the study area, was done during
the wet season. Three people spent approximately
360 hours searching for nests. In each habitat the
time spent per square meter was about one per-
son-minute.
For the estimates of density of adult wasps/
ha, the number of adults within some of the nests
were counted and also, for some species, data
from Richards (1978) were compiled.
In each table the habitats, when it was pos-
sible, were arranged in order of increasing com-
plexity of structure because this makes it easier to
see general patterns in species preferences for
habitats.
Volume 3. 1 994
137
RESULTS AND DISCUSSION
1. Nesting Habitat Preference of Wasps
A total of 100 colonies, comprising 30 species
in 15 genera of wasps were encountered in six
different habitats in Chapada dos Guimaraes (Table
1). Among them, 18 species nested in only one
habitat, nine in two, and three (Polybia (Myrapetra)
ruficeps Schrottky — a very common species of the
region, Chartergus chartarius (Olivier) and
Clwrtegelhis communis Richards) in three habitats
(Table 1). Of the species that nested in only one
habitat, most were found in cerrado sensu-stricto
(seven species), in gallery forest (five species) and
in campo umido (four species) (Table 1). Of the 30
species collected in this study, only Polistes
(Epicnemius) pacificus liliaciosus de Saussure was
not collected by Richards (Table 1 and 2). This
subspecies has been collected only rarely in cen-
tral Brazil and, until Richards' 1978 record, its
distribution was thought to be restricted in Brazil
to the North (Amazonas, Amapa and Para states).
Polybia ruficeps was the commonest species of wasp
at Rio-Manso with 23 colonies, of which 70% were
found in cerrado sensu-stricto. Three other species,
Epipona tatua (Cuvier), Synoeca surinama (L) and
Oiartergus chartarius, with eight colonies each,
were also common (Table 1).
Richards (1978) recorded 199 colonies com-
prising 51 species in 14 genera nesting in six habi-
tats (Table 2). Most of the species he collected (25
of the 51), were found nesting only in one habitat
(eight of them in dry forest), 12 species nested in
two habitats, eight nested in three habitats, three
nested in four, and one species Polybia (Myrapetra)
platycephala Richards nested in five habitats. In
Xavantina and Serra do Cachimbo, Polybia
(Myrapetra) occidentalis (Olivier) with 24 colonies
(12%), was the most abundant, followed by Polybia
ruficeps (10%), Polybia (Apopolybia) jurinei de
Saussure (8%), Polybia (Trichothorax) ignobilis
(Haliday) (7%) and Polybia (Formicicola) rejecta (F)
(5%) (Table 2).
The data collected by us and by Richards were
combined and for the same type of habitat the data
were lumped (Table 3 representing the sum of
Tables 1 and 2). At the nine habitats 299 colonies,
comprising 61 species and 16 genera, were col-
lected. Twenty eight species (46%) nested in only
one habitat, 13 species nested in two, 10 in three,
seven in four and three in five habitats (Table 3).
The area surveyed was different in each habitat so
the number of colonies found do not represent one
absolute abundance but the relative habitat pref-
erence for nesting.
Of the total number of colonies recorded by
Richards and us from Mato Grosso, the species
with the largest number of colonies was Polybia
ruficeps (14%), followed by Polybia occidentalis (9%),
Polybia jurinei (6%), Polybia (Myrapetra) erythrothorax
Richards (5%), Polybia ignobilis (5%), Paraclmrtergus
fratemus (Gribodo) (4%) and Epipona tatua (4%).
The genus Polybia represents 60% of the colonies
reported from the two regions of Mato Grosso
State (Table 3), ranging from 67% in Xavantina and
Serra do Cachimbo (Table 2) to 45% in Rio Manso
(Table 1). Among all the colonies of the 19 species
of Polybia collected, 58% were found in cerrado
sensu-stricto (Table 3) ranging from 31% in Rio
Manso and 55% in Xavantina and Serra do
Cachimbo (Tables 1 and 2).
Richards (1978) suggested that the choice of
habitat for nesting is very characteristic and less
diverse than that used for foraging. His opinion is
confirmed in our survey at Rio Manso. For ex-
ample, both Synoeca suririama (L), which nests in
gallery forest, and Polybia (Trichothorax) sericea
(Olivier) in cerrado sensu-stricto, were collected
when they hunted in gallery forest, cerrado sensu-
stricto, campo sujo and campo umido. Of the 30
species of wasps at Rio Manso, 21 (70%) were
encountered foraging in more than one habitat (in
preparation).
A water source is important for wasps to nest
successfully, (Rau 1942, Forsyth 1980). The latter
author concluded that the greater colony density
of wasps in gallery forest was due mainly to the
presence of water. Wet habitats at Rio Manso
(gallery forest, eight/ha and campo umido 60/ha)
also had higher colony densities than did dry
habitats (cerrado sensu-stricto five/ha). Polistes
(Epicnemius) subsericeus de Saussure and
Mischocyttarus (Mischocyttarus) drewseni de
Saussure which nest in campo umido, had higher
colony density compared to other species in drier
habitats, such as cerrado sensu-stricto. Availability
of water all year round and nest protection by tufts
of grasses and sedges, could be the cause of that
high density.
Most animals have preferences for particular
habitats (Partridge, 1978). In Mato Grosso ap-
138
Journal of Hymenoptera Research
Table 1. Wasps nesting habitats in Rio Manso Chapada dos Guimaraes: Mato
Grosso, Brazil.
Species
CU
VE
cs
OF
CE
GF
TT
Apoica c.f. pollens
0
0
0
0
1
1
2
Brachygastra bilineolata
0
0
0
0
1
0
1
Cliartegellus communis
0
1
1
0
1
0
3
Clwrtergus chartarius
0
0
4
0
3
1
8
Epipona tatua
0
0
0
0
5
3
8
Metapolybia cingulala
0
0
0
0
1
0
1
Mischocyttarus cerberus
0
0
0
0
2
0
2
Mischocyttarus drewseni
2
0
0
0
0
0
2
Mischocyttarus labiatus
0
0
0
0
1
0
1
Mischocyttarus matogrossoensis
1
0
0
0
0
0
1
Mischocyttarus mcthathoracicus
0
0
0
0
1
0
1
Parachartergus fratemus
0
1
0
0
4
0
5
Polistes canadensis
1
0
0
0
0
0
1
Polistes pacificus
0
0
0
0
0
1
1
Polistes subsericeus
4
0
0
0
0
0
4
Polybia emaciata
0
0
0
0
0
1
1
Polybia erythrothorax
0
0
2
0
4
0
6
Polybia jurinei
0
0
0
0
2
0
2
Polybia occiden talis
0
0
0
1
2
0
3
Polybia paulista
0
0
0
0
0
2
2
Polybia quadricincta
0
0
0
2
0
0
2
Polybia ruficeps
(1
0
0
5
16
2
2<
Polybia sericea
0
0
0
1
2
0
3
Polybia singularis
0
11
11
0
1
2
3
Protopolybia exigua
0
0
0
1
1
0
2
Pseudochartergus chartergoides
0
0
0
0
0
1
1
Pseudopolybia compressa
0
0
0
0
0
1
1
Pseudopolybia vespiceps
0
(1
0
0
1
0
1
Stelopolybia lobipleura
0
0
1
0
0
0
1
Synoeca surinama
0
0
0
2
0
6
8
Total of nests
08
02
08
12
49
21
100
Total of species
04
02
04
06
18
11
30
CU - Campo umido
VE - Vereda
CS - Campo sujo
OF - Old field
CE - Cerrado sensu-stricto
GF - Gallery forest
TT- Total
Volume 3, 1994
139
Table 2. Wasp nesting habitats of Xavantina and Serra do Cachimbo: Mato
Grosso, Brazil (Data from Richards, 1978)
Species
CS
DC
CE
CD
GF
DF
TT
Apoica gellida
0
0
0
2
0
1
3
Apoica pollens
0
0
0
0
1
2
3
Brachygastra augustii
0
3
1
3
0
0
7
Brachygastra moebiana
0
0
1
1
0
0
2
Brachygastra scutellaris
0
0
0
0
0
2
2
Chartegellus communis
0
0
1
1
0
0
2
Cluirtergus chartarius
0
0
1
1
0
0
2
Chartergus metanotalis
0
0
1
0
0
0
1
Clypearia humeralis
0
1
0
0
0
0
1
Epipona tatna
0
1
2
0
0
0
3
Mischocyttanis flavicornis
0
2
0
0
0
0
2
Mischocyttarus latior
0
0
1
0
0
0
1
Mischocyttanis matogrossoensis
1
0
0
0
0
0
1
Mischocyttanis melanoxanthus
0
1
0
0
0
0
1
Mischocyttanis metluithoracicus
0
0
0
0
0
1
1
Mischocyttanis omicron
0
1
0
0
0
0
1
Mischocyttarus surinamensis
0
0
0
0
1
0
1
Mischocyttarus undulatus
0
0
0
0
2
0
2
Parachartergus fraternus
0
4
4
0
0
0
8
Polistes billardieri
1
0
0
0
0
0
1
Polistes cinerascens
0
0
1
0
0
0
1
Polistes goeldii
1
0
0
0
0
0
1
Polybia chrysothorax
0
0
2
0
0
0
2
Polybia dimidiata
0
0
0
2
2
2
6
Polybia emaciata
0
0
2
0
0
2
4
Polybia en/throthorax
0
1
2
6
0
0
9
Polybia gorytoides
0
1
0
0
3
0
4
Polybia ignobilis
2
0
7
0
0
5
14
Polybia jurinei
0
0
2
5
6
2
15
Polybia liliacea
0
0
0
1
0
1
2
Polybia micans
0
0
0
0
0
1
1
Polybia occidentalis
11
6
1
6
0
0
24
Polybia platycepluila
0
1
1
1
1
2
6
Polybia quadricincta
0
0
0
0
0
2
2
Polybia rejecta
0
0
1
1
2
6
10
Polybia ruficeps
0
6
7
0
7
0
20
Polybia scrobalis
0
0
0
0
2
0
2
Polybia sericea
0
3
1
2
0
0
6
Polybia singularis
0
1
0
3
0
1
5
Polybia striata
0
0
0
0
0
2
2
Protopolybia acutiscutis
0
0
0
0
0
1
1
Protopolybia exigua
1
0
0
0
0
0
1
Protopolybia sedula
0
0
1
0
0
1
2
Pseudopolybia compressa
0
0
0
1
0
0
1
Pseudopolybia vespiceps
0
0
0
1
0
0
1
140 Journal of Hymenoptera Research
Species CS DC CE CD GF DF TT
Stelopolybia angulata
0
0
0
1
2
1
4
Stelopolybia fulvofasciata
0
0
0
0
0
1
1
Stelopolybia lobipleura
1
0
0
0
0
0
1
Stelopolybia testacea
0
0
0
0
2
1
3
Synoeca chalybea
0
0
0
2
0
0
2
Synoeca surinama
0
0
0
0
0
1
1
Total of nests
18
32
40
40
31
38
199
Total of species
07
14
20
L8
12
21
51
CS - Campo sujo
DC - Dirt roads and clearings
CE - Cerrado sensu-stricto
CD - Cerradao
GF - Gallery forest
DF - Dry forest
TT- Total
Table 3. Wasp nesting habitats in Chapada dos Guimaraes and Xavantina and Serra do Cachimbo:
Mato Grosso, Brazil.
Species
CU
VE
CS
OF
DC
CE
CD
GF
DF
TT
Apoica gellida
0
0
0
0
0
0
2
0
1
3
Apoica pallens
0
0
0
0
0
1
0
2
2
5
Brachygastra augustii
0
0
0
0
3
1
3
0
0
7
Brachygastra bilineolata
0
0
0
0
I)
1
0
0
0
1
Brachygastra moebiana
0
0
0
0
0
1
1
0
0
2
Brachygastra scutellaris
0
0
0
0
0
0
0
0
2
2
Cluirtegellus communis
0
1
1
0
I)
2
1
0
0
5
Clwrtergus cliartarius
0
0
4
0
0
4
1
1
0
10
Owrtergus metanotalis
0
0
0
0
0
1
0
0
0
1
Clypearia humeralis
0
0
0
0
1
0
0
0
0
1
Epipona tatua
0
0
0
0
1
7
0
3
0
11
Metapolybia cingulata
0
0
0
0
0
1
0
0
0
1
Mischocyttarus cerberus
0
0
0
0
0
2
0
0
0
2
Mischocyttarus dreioseni
2
0
0
0
o
0
0
0
0
2
Mischocyttarus flavicornis
0
0
0
0
2
0
0
0
0
2
Mischocyttarus labiatus
0
0
0
0
0
1
0
0
0
1
Mischocyttarus latior
0
0
0
0
0
1
0
0
0
1
Mischocyttarus matogrossoensis
1
0
1
0
0
0
0
0
0
2
Mischocyttarus melanoxanthus
0
0
0
0
1
0
0
0
0
1
Mischocyttarus metliathoracicus
0
0
0
0
0
1
0
0
1
2
Mischocyttarus omicron
0
0
0
0
1
0
0
0
0
1
Mischocyttarus surinamensis
0
0
0
0
0
0
0
1
0
1
Mischocyttarus undulatus
0
0
0
0
0
0
0
2
0
2
Parachartergus frateruus
0
1
0
0
4
8
0
0
0
13
Polistes billardieri
0
0
1
0
0
0
0
0
0
1
Polistes canadensis
1
0
0
0
0
0
0
0
0
1
Volume 3. 1 994
141
Species
CU VE CS OF DC CE CD GF DF TT
Polistes cinerascens
0
0
0
0
0
1
0
0
0
1
Polistes goeldii
0
0
1
0
0
0
0
0
0
1
Polistes pacificus
0
0
0
0
0
0
0
1
0
1
Polistes subsericeus
4
0
0
0
0
0
0
0
0
4
Poly bia chrysothorax
0
0
0
0
0
2
0
0
0
2
Polybia dimidiata
0
0
0
0
0
0
2
2
2
6
Polybia emaciata
0
0
0
0
0
2
0
1
2
5
Polybia erythrothorax
0
0
2
0
1
6
6
0
0
15
Polybia gorytoides
0
0
0
0
1
0
0
3
0
4
Polybia ignobilis
0
0
2
0
0
7
0
0
5
14
Polybia jurinei
0
0
5
0
0
4
5
6
2
17
Polybia liliacea
0
0
0
0
0
0
1
0
1
2
Polybia micans
0
0
0
0
0
0
0
0
1
1
Polybia occidentalis
0
0
11
1
6
3
6
0
0
27
Polybia paulista
0
0
0
0
0
0
0
0
2
2
Polybia platycephala
0
0
0
0
1
1
1
1
2
6
Polybia qimdricincta
0
0
0
2
0
0
0
0
2
4
Polybia rejecta
0
0
0
0
0
1
1
2
6
10
Polybia ruficeps
0
0
0
5
6
23
0
9
0
43
Polybia scrobalis
0
0
0
0
0
0
0
2
0
2
Polybia sericea
0
0
0
1
3
3
2
0
0
9
Polybia singnlaris
0
0
0
0
1
1
3
2
1
8
Polybia striata
0
0
0
0
0
0
0
0
2
2
Protopolybia acutiscutis
0
0
0
0
0
0
0
0
1
1
Protopolybia exigiia
0
0
1
1
0
1
0
0
0
3
Protopolybia sedula
0
0
0
0
0
1
0
0
1
2
Pseudoclmrtergus clwrtergoides
0
0
0
0
0
0
0
1
0
1
Pseudopolybia compressa
0
0
0
0
0
0
1
0
1
2
Pseudopolybia z>espiceps
0
0
0
0
0
1
1
0
0
2
Stelopolybia angulata
0
0
0
0
0
0
1
2
1
4
Stelopolybia fitlvofasciata
0
0
0
0
0
0
0
0
1
1
Stelopolybia lobipleura
0
0
1
0
0
1
0
0
0
2
Stelopolybia testacea
0
0
0
0
0
0
0
2
1
3
Synoeca chalybea
0
0
0
0
0
0
2
0
0
2
Synoeca surinama
0
0
0
2
0
0
0
6
1
9
Total of nests
08
02
24
12
32
88
39
48
41
299
Total of species
04
02
10
06
14
29
17
19
23
61
CU — Campo umido
VE— Vereda
CS — Campo sujo
OF— Old field
DC — Dirt roads and clearings
CE — Cerrado sensu-stricto
CD — Cerradao
GF — Gallery forest
DF — Dry forest
TT— Total
142
Journal of Hymenoptera Research
Table 4. Wasp colonies density in three different habitats in Rio Manso - Chapada dos
Guimaraes: Mato Grosso, Brazil.
Campo
umido
Cerrado
sensu stricto
Gallery
forest
Area (m2)
500
Number of nests
3
Nest density/ha
60
Density of adult
wasps/ha
480
Number of species
2
10000
5
5
1300
5
7500
6
8
50000
5
proximately 50% of wasps were restricted to a
single habitat for nesting (Table 3).
Only a few ubiquitous species like Polybia
occidentalis (5), Polybia platycephala (5), Polybia
(Pedothoeca) si)igularis Ducke (5), Chartergellus
communis (4), Chartergus chartarius (4), Polybia
erythrothorax (4), Polybia rejecta (4), Polybia ruficeps
(4) and Polybia sericea (4) use more than three
habitats for nesting (Table 3).
Substrates for nesting in the cerrado is almost
unlimited (Henriques el al. 1992), but availability
of food, water and protection could be limiting
factors.
2. Density
In an area of 2500 m2 of the gallery forest (Fig.
5) three colonies of two species of wasps were
found, two of Polybia (Myrapetra) paulista H.von
Ihering and one of Polybia ruficeps. In a contiguous
area of approximately 5000 m2 three colonies of
three species (Polybia (Pedothoeca) emaciata Lucas,
Polybia (Myrapetra) sp. and Polybia (Pedothoeca)
singularis (Ducke) were found, which correspond
to an average of eight colonies and five species per
ha (Table 4). In an area of 10000 m2 of cerrado sensu
stricto (Fig. 4), near a stream, three colonies of the
following species were found: Mischocyttarus
( Kappa) metathoracicus(deSaussure),Mischoa/ttarus
(Mischocyttarus) labiatus (F) and Polybia ruficeps.
Colonies of Brachygastra bilineolata Spinola and
Stelopolybia lobipleura Richards were also found on
trees close to each other, but away from the stream
with an average of colonies of five species per ha
(Table 4). A similar result was found in the cerrado
sensu-stricto of Brasilia DF ( Ecological Reserve of
IBGE), where six colonies were found in an area of
1.5 ha corresponding to an average of five colo-
nies per ha (Henriques et al, 1992).
In an area of 500 m2 of campo umido (Fig. 1),
two colonies of Polistes subsericeus and one of
Mischocyttarus drewseni were found. Based on these
numbers estimates were 60 colonies per ha for
both species, or 40 per ha for Polistes subsericeus
and 20 per ha for Mischocyttarus drewseni (Table 4).
Although the colony density in campo umido was
larger than that found in gallery forest and cerrado
sensu-stricto, the number of individuals per nest
was smaller, due to low numbers of adults per nest
(eight per nest). In gallery forest and cerrado sensu
stricto, the colonies of Synoeca surinama, Epipona
tatua and Polybia spp, were larger, with hundreds
to thousands of individuals per nest (Table 4).
In a relatively simply-structured habitat, such
as campo umido (Fig. 1) with a small area (500 m2),
the density estimate was more accurate than in
more complex habitats. A ranking of habitats in
order of reliability of estimate is: campo umido,
cerrado sensu-stricto and gallery forest.
Volume 3. 1 994
143
ACKNOWLEDGMENTS
We acknowledge the contribution made by the late Professor
O.W. Richards to studies of social wasps in general. It is not
unusual for subsequent workers to discover interesting in-
formation in the data he published. Financial support was
provided by the North Brazilian Electricity Company
(Eletronorte) and the University of Brasilia. We are espe-
cially grateful to Dr. Anthony Raw and Dr. Linda Caldas of
the Universidade de Brasilia and the anonymous reviewer
for useful comments on the manuscript. Dr. Raw also iden-
tified several of the species of wasps. Dr. Paul M. Marsh
made some editorial changes on the manuscript. Sra. Beatnz
Mossri kindly assisted in the collection of field data. We also
thank Dr. Antonio Jose E. Brussi, for his help and encourage-
ment
LITERATURE CITED
Eiten, G. 1972. The Cerrado vegetation of Brazil. The Botanical
Review 38:201-341.
Forsyth, A. 1980. Nest site and Habitat selection by the social
wasp, Metapolybia azteca Araujo (Hymenoptera:
Vespidae) Brenesia 17: 157-162.
Gobbi, N. & Machado, V. L. L. 1985. Material capturado e
utihzado na alimentacao de Polybia (Myrapetra) paulista
Ihering 1896 (Hymenoptera-VespidaeJ./lnais da Sociedade
Entomologica do Brasil 14(2): 189-195.
Gobbi, N. Machado, V. L. L. & Tavares Filho, J. A. 1984.
Sazonalidade das presas utilizadas na alimentacao de
Polybia occidentalis (Olivier 1791) (Hymenoptera:
Vespidae). Anais da Sociedade Entomologica do Brasil 13(1):
63-69.
Henriques, R. P. B. Rocha, I. R. D. & Kitayama, K. 1992. Nest
density of some social wasps species in Cerrado vegeta-
tion of Central Brazil (Hymenoptera: Vespidae)
Entomologia Generalis 17(4): 265-268.
Jeanne, R. L. 1975. The adaptiveness of social wasp architec-
ture. Quarterly Review of Biology 50: 267-287.
Kitayama, K. Rocha, I. R. D. & Bulhoes B. M. 1989. Densidade
de ninhos de vespideos em diversos habitats de Mato
Grosso. I . Shnpdsio Latino-americano sobre Inseto Socials
Neotropicais. Rio Claro SP. R-67 P 44.
Kojima, J. 1. 1982. Nest architecture of three Ropalidia species
(Hymenoptera: Vespidade) on Leyte Island, The Philip-
pines. Biotropica 14(4): 272-280.
Machado, V. L. L. Gobbi, N. & Simoes, D. 1987. Material
capturado naalimentai;aodeSrp/opo/i/l)i(!fi«//if)fs (Olivier
1791) (Hymenoptera: Vespidae) Anais da Soctedade
Entomologica do Brasil 16 (1): 73-79.
Partridge, L. 1978. Habitat selection In Behavioural Ecology
(Eds. JR. Krebs & N.B. Davies) Blackwell Scientific
Publications, Oxford, 351-376.
Rau, P. 1942. Habitat preferences of Polistes wasps (Hy-
menoptera: Vespidae) Entomological Nezus 53: 293-295.
Raw, A. 1988. Social wasps (Hymenoptera: Vespidae) and
insect pests of crops of the Surui and Cinta Larga
Indians in Rondonia, Brazil. The Entomologist 107: 104-
109.
Reed, H. C. & Vinson, S. B. 1979. Nesting ecology of paper
wasps (Polistes) in a Texas urban area (Hymenoptera:
Vespidae). Journal of the Kansas Entomological Society 52
(4): 673-689.
Richards, O. W. 1978. The social wasps of the Americas.
British Museum of Natural History. London 580 pp.
Starr, C. K. 1988. The nesting association of the social wasps
Mischocyttarus immarginatus and Polybia spp. in Costa
Rica. Biotropica 20(2): 171-173.
Windsor, D. M. 1972. Nesting association between
twoNeotropical polybiine wasps (Hymenoptera:
Vespidade)Biofropic«. 4: 1-3.
J. HYM. RES.
Vol. 3, 1994, pp. 145-150
Descriptions of two New Species and Notes on the Genus Bakeriella Kieffer
from Brazil and Ecuador (Hymenoptera: Bethylidae)
Celso Oliveira Azevedo
Departamento de Biologia, Universidade Federal do Espinto Santo, Av. Marechal Campos 1468, Maruipe, 29040-090
Vitoria, ES Brasil
Abstract. — Bakeriella mcompleta sp.n., from rubber-tree crops of Sao Paulo, Brazil and Bgrossensissp. n. fromMatoGrosso,
Brazil are described and illustrated. Taxonomic data on B. reclusa (Evans) and B. montivaga (Kieffer) are included, and the latter
recorded from Brazil for the first time. A key to females with a transverse carina on the pronotal disc is presented.
INTRODUCTION
Bakeriella Kieffer is a neotropical genus, rang-
ing from Southern United States to Northern Ar-
gentina (Azevedo 1991). Kieffer (1910a) described
this genus based on one species from Para, Brazil,
B. flavicornis, which has the pronotum carinate
laterally and medially as well as anteriorly. This
genus is related to Epyris Westwood by having the
scutellar groove divided (Evans 1964). Kieffer
(1910b) described B. depressa from Peru which has
a carinate pronotum also. Evans (1964) expanded
the definition of Bakeriella when he described ten
species, in some of which the median or lateral
carinae of the pronotum are weak or absent. Evans
(1969), in his revision of Epyris, pointed out the
relationship of the monthwgiis-group of Epyris to
Bakeriella by the fact that both groups have similar
scutellar pits. Evans (1979) reviewed Bakeriella,
providing identification keys for both males and
females and arranging the genus into four species-
groups. In this revision, Evans described one spe-
cies, B. erythrogaster, which has the scutellar groove
similar to the genus Rliabdepyris Kieffer. Evans
also transfered members of the montivagus-group
from Epyris to Bakeriella simply by the fact that
they closely resemble each other. Curiously, he
did not include B. subcarinata Evans 1965 in his
revision. Gordh & Moczar (1990) cited in their
catalog 19 species for the world and Azevedo
(1991) described B. dentata from Sao Paulo State,
Brazil.
The present study focuses on four species of
this genus, B. montivaga (Kieffer), B. reclusa (Evans)
and two new species. Ideally what is needed is a
complete revsion of the genus. Unfortunately,
specimens are rare in collections and until more
material is available for study, I have chosen to
present these notes and new species descriptions.
Specimens for this study belong to the follow-
ing collections (abbreviations according to Arnett
& Samuelson (1986)): IBSP, Instituto Biologico de
Sao Paulo, SP, Brasil; MZSP, Museu de Zoologia
da Universidade de Sao Paulo, SP, Brasil; UFES,
Colecao de Entomologia da Universidade Federal
do Espirito Santo, Vitoria, ES, Brasil. The last col-
lection is not included in Arnett & Samuelson
(1986) because it is new.
Abbreviations used follow Evans (1964) and
are as follows: LFW, length of forewing; WH,
maxim un width of head, including eyes; LH, length
of head, measured from median apical margin of
clypeus to median point of vertex; WF, minimun
width of frons; HE, maximun height (or length) of
eye; WOT, maximun width of ocellar triangle,
including posterior ocelli; OOL, shortest ocello-
ocullar line. The nomenclature of the integument
follows Eady (1968).
146
Journal of Hymenoptera Research
Bakeriella incompleta, sp. n.
Figs. 1-2
Holotype female.— Brazil: SP,Ibitinga;05.X.1988;
yellow pan trap, rubber-tree crop, no collector
given. (UFES).
Description of holotype female. — Length of body
6.0 mm; LFW 2.85 mm; head and thorax black,
abdomen black except tip dark brownish; palpi
light brownish; mandible brownish, black at base
and with a light brownish spot outside, teeth
reddish; antennal scape dark brownish, apex
brownish, pedicel and flagellum somewhat
infuscated; tegula brownish; fore and hindcoxae
black, midcoxa darkbrownish; trochanters, femora,
tibiae and tarsi dark brownish, the anterior being
lighter; wings subhyaline somewhat brownish,
veins, prostigma and stigma dark brownish.
Head (Fig. 1). Mandible with 5 teeth, apical
tooth largest (Fig. 1). Clypeus with subangulate
median lobe, with median carina straight in lat-
eral profile, clypeus forming a small excavated
reentrance in the region of contact with frons. WH
1.03 X LH; WF 0.5 XWH;WF 1.34 X HE; WOT 1.33
X OOL. Antennal scrobe not carinate. Posterior
ocelli about 3 X their diameter from the vertex.
Vertex nearly straight and with lateral angles
rounded off. Distance from the top of eyes to crest
of vertex about 0.7 X HE. Length of first four
antennal segments in a ratio of about 11:5:4:4.
Segments 5 to 12 about as wide as long. Temple not
carinate. Frons somewhat alutaceous and bril-
liant.
Mesosoma(Fig. 1). Somewhat more alutaceous
than frons, mesoscutum polished anteriorly.
Pronotal disc slightly convex anteriorly, tranverse
anterior carina without teeth, median carina weak,
situated in a slight depression and absent anteri-
orly, not intersecting transverse carina. Posterior
margin of pronotal disc paralelled by a series of
small foveae; lateral margin of disc somewhat
sharpened, not carinate, and slightly diverging
posteriorly; median width of the disc slightly
greater than its length; disc slightly shorter than
mesoscutum and scutellum together. Notauli not
reaching the anterior margin of mesoscutum, con-
verging posteriorly, widened posteriorly, and
slightly curved anteriorly. Parapsidal furrows
straight, somewhat more enlarged posteriorly,
occupying only the posterior half of mesocurum
(fig. 1). Scutellar pits tranverse, much wider than
long, separated by a thin septum which is lower
than margin of foveae. Propodeal disc as wide as
long, with three discal carinae, median one com-
plete and the others incomplete, occupying about
two thirds of disc and converging posteriorly but
not reaching median carina; space among discal
carinae with tranverse striae; disc without
sublateral carinae; inner side of lateral carinae
paralelled by a series of small irregular depres-
sions; anterior part of space between lateral and
discal carinae occupied by a rather large depres-
sion; posterior angle of the disc foveolate; decliv-
ity of propodeum with median carina and perpen-
dicular striae. Metapleuron with longitudinally
sinuous striae. Mesopleuron granulate, with large
fovea anteriorly and small, deep pit at anterior
margin, with small pentagonal fovea and elon-
gate upper fovea (fig. 2). Mesosternum with small
median pit, anterior margin of acetabular carina
paralelled by striae. Midtibia spinose. Forefemur
2.0 X as long as thick, midfemur 1.65 X as long as
thick. Claw trifid, first tooth rounded off, the oth-
ers sharp.
Metasoma. Sterna setose on posterior of each
sternum. Terga with few setae on the posterior
half of each tergum. Terga I and II with lateral
setae. Tranverse section of metasoma circular.
Length of mesosoma 1.35 X length of metasoma.
Valvae III dark brownish with only its end setose,
setae range between one third and a half length of
the valvae.
Paratype female. — BRAZIL: SP, Balsamo;
21. IX. 1988, white pan trap, no collector given.
(IBSP).
The paratype differs from holotype as fol-
lows: WH 1.00 XLH;WF0.57XWH; WF 0.76 X HE;
OOL 1.62 X WOT; propodeal disc 1.08 as large as
long.
Remarks. — This species is known only from
the female. Bakeriella incompleta differs from other
species in the genus in having the median longitu-
dinal pronotal carina incomplete anteriorly, not
reaching the anterior transerve carina, but reach-
ing the posterior margin. In B. cristata Evans this
carina is incomplete posteriorly.
Etymology. — The specific name refers to the
fact that the median longitudinal carina of the
pronotal disc is incomplete.
Volume 3. 1 994
147
Figs 1-2. Bakeriella mcompleta, sp n. 1, head and mesosoma, dorsal view; 2, mesopleurum, lateral view. Scale = 0.5 mm.
Bakeriella grossensis, sp. n.
Holotype female. — Brazil: Barra do Tapirape,
MT; XII.1960; unknown trap; B. Malkin, col.
(MZSP).
Description of holotype female. — Length of body
3.75 mm; LFW 2.4 mm; head and mesosoma black,
metasoma black with apex somewhat brownish;
palpi brownish; mandible yellowish with teeth
darker; antenna yellowish with scape darker;
tegula yellowish; coxae and femora brown, tro-
chanters, tibiae and tarsi yellowish; wings
subhyaline, veins brown.
Head. Mandibles with 5 teeth, the basal teeth
small and rounded off, the 2 apical teeth sharp-
ened, anterior tooth largest, the second of median
size. Clypeus with median lobe angulate, without
median carina, clypeus forming a small excavated
reentrance in the region of contact with frons. Eyes
with sparse long hairs. WH 1.08 X LH; WF 0.58 X
WH; WF 1.16 X HE; WOT 1.62 X OOL. Antennal
scrobe not carinate. Distance posterior ocelli from
vertex about 2.8 X diameter of ocelli, angle of
ocellar triangule less than a right angle. Vertex
rounded, very slightly angled laterally. Distance
from the top of eyes to crest of vertex about 0.57 X
HE. Length of first four antennal segments in a
ratio about 20:8:7:7. Temple not carinate. Frons
coriaceous with punctures which are separated by
1-3 X their own diameters.
148
Journal of Hymenoptera Research
Mesosoma. Pronotal disc coriaceous, punc-
tures similar to frons, with straight anterior trans-
verse carina, without median longitudinal carina,
posterior margin paralelled by series of small
foveae; sides of disc subparalell; median width of
disc 1.17 X its length. Pronotal disc 0.77 longer
than mesoscutum and scutellum together. Ante-
rior half of mesoscutum alutaceous, posterior half
coriaceous. Notauli straight and very thin, vesti-
gial anteriorly, widened posteriorly, reaching
both anterior and posterior margin of mesoscutum.
Parapsidal furrows straight, occupying only pos-
terior half of mesoscutum and reaching its poste-
rior margin. Scutellar pits transverse, 2.0 X as wide
as long, separated by thin septum. Propodeal disc
1.05 X as wide as long, with three discal carinae,
median one complete and lateral ones converging
behind, not reaching median carina; space be-
tween discal carinae with small transverse striae,
space between discal carinae and lateral carina
polished; disc without sublateral carinae; lateral
carinae boardered by a depression with small
transverse striae; posterior angle of disc foveolate;
declivity of propodeum with median carina and
perpendicular striae. Metapleuron alutaceous,
without striae. Mesopleuron coriaceous, with large
ventral fovea occupying the lower half of
mesopleuron, with deep pit in upper margin of
lower fovea, with a fovea in the anterior part of
mesopleuron and with long fovea in upper part of
mesopleuron opened anteriorly. Mesosternum
coriaceous, with pit in the middle of posterior
region. Midtibia spinose. Forefemur 2.5 X as long
as thick, midfemur 1.7 X. Claws trifid, first tooth
rounded off, the others sharpened.
Metasoma. Wholly polished, terga and sterna
with few setae. Setae of terga on posterior and
lateral margins. Transverse section of metasoma
subcircular. Length of mesosoma slightly longer
than the metasoma. Valvae III brown with setae on
apex.
Remarks. — This species is known only from
the holotype. It runs to B. inconspicua and B. polita
in the key written by Evans (1979). B. grossensis
differs from B. inconspicua in having the scutellar
pits tran verse, the claws trifid, the eyes not strongly
setose, the metasoma black with reddish apex and
the mandible 5-toothed. B. grossensis differs from
B. polita in lacking a median carina in the clypeus,
in lacking a lateral carina on the pronotal disc, the
head being not well developed above the eyes and
the occipital carina being obscured in full frontal
view of head. In addition, the mandible of B.
grossensis has two big apical teeth, while B. polita
has one and B. inconspicua has three.
Etymology. — The specific name is in reference
to the Mato Grosso State, region of origin of the
holotype.
Bakeriella montivaga (Kieffer) 1910b
Epyris montivagus Kieffer 1910b:31; Evans 1969:324
Bakeriella montivaga: Evans 1979:261
This species is recorded from Peru, Colombia
(Kieffer 1910b), Bolivia, Ecuador, Venezuela,
Panama, Costa Rica, Honduras, Mexico (Evans
1969), and now from Brazil.
Four females have been studied: Balsamo, SP,
Brazil, one female, 28. IV. 1988, yellow pan trap,
rubber-tree crop, no collector given (UFES). The
others were from Pichincha, Centro Cientifico R.
Palangue, Ecuador, 21.XII.1980, 1.1.1981, 4.1.1981,
forest, S. Sandoval, col. (MZSP).
These specimens are similar to the type;
pronotal disc with the sides diverging posteriorly;
WH 0.99 to 1.17 X LH; WF 0.53 to 0.59 X WH; WF
1.18 to 1.46 X HE; OOL 1.63 to 1.94 X WOT;
propodeal disc slightly wider than long. The line
of small foveae near the posterior margin of the
pronotal disc is weakly angled anteriorly in the
middle. This character is common only in males.
Evans (1969) considered the possibility that B.
quinquepartita (Kieffer) and B. montivaga represent
variations of a single species. Indeed, both species
are very similar and the only differences between
them were pointed out in Evans'(1979) key to
females.
Bakeriella reclusa (Evans) 1969
Epyris reclusus Evans 1969:329
Bakeriella reclusa: Evans 1979:261
This species was described from Costa Rica
and El Salvador. Evans (1979) studied a female
from Ecuador. I have examined a male from Ecua-
dor: Morona Santiago, Los Tayos, 3. VIII. 1976,
DeVries, col. (MZSP). Measurements for this spe-
cies are: LFW = 2.13 mm, WH 1.2 X LH; WF 0.55 X
WH; WF 1 .06 X HE; WOT 1 .44 X WOT. The pronotal
disc is slightly wider than long, and the propodeal
disc is as long as wide.
Volume 3. 1994 149
The following key to females with an anterior transverse carina on the pronotal disc is modified
from Evans (1979:258) and includes B. dentata Azevedo 1991, B. subcarinata Evans 1965, B. incompleta, sp.
n., and B. grossensis, sp. n.
1 Pronotum without median longitudinal carina; head black 2
1' Pronotum usually with complete median carina; head black or dark green 7
2(1) Legs wholly yellowish; propodeal disc covered with weak transverse striations ... floridana Evans 1964
2' Coxae and femora largely black; metapleuron shining, at most with very weak sculpture 3
3(2') Transerve carina of pronotum weakly angled forward medially; median area of propodeum rather weakly
sculptured 4
3' Transerve carina of pronotum evenly arched; median area of propodeum with strong transverse rigdes
5
4(3) Mandible with 5 teeth, basal 3 small; frons with small punctures which are separated by 1.5-3.0 X their own
diameters olmeca Evans 1964
4' Mandible with only two large apical teeth; frons with somewhat stronger and more widely spaced
punctures brasiliana Evans 1964
5(3) Sides of pronotal disc carinate anteriorly; scutellar pits ovoid; mandible with 3 strong apical teeth
inconspicua Evans 1964
5' Sides of pronotal disc not carinate; scutellar pits transverse; mandible with 1 or 2 strong apical teeth
6
6(5') Sides of pronotal disc sharpened; mandible with one large apical tooth; scutellar pits slightly wider than
long polita Evans 1964
6' Sides of pronotal disc angled, but not truly sharpened; mandible with two large apical teeth; scutellar pits
2.0 X wider than long grossensis, sp. n.
7(1') Median carina of pronotal disc incomplete 8
7' Median carina of pronotal disc complete 9
8(7) Median longitudinal carina of pronotum lacking posteriorly; body black with green metallic reflexions
cristata Evans 1964
8' Median longitudinal carina of pronotum lacking anteriorly; body black without metallic reflexions
incompleta, sp. n.
9(7') Internal margin of tranverse carina of pronotum with tooth-like emargination .... dentata Azevedo 1991
9' Internal margin not like above 10
10(9) Median longitudinal carina of pronotum somewhat weak and not paralleled by a depression
subcarinata Evans 1965
10' Median longitudinal carina of pronotum well evident and situated in a depression inca Evans 1964
ACKNOWLEDGMENTS
I express thanks to Ehana Bergmann for the loan of
material from IBSP and Dr. Carlos Brandao for the loan of
material from MZSP. I also thank Oscar Shibatta for prepar-
ing the illustrations.
150
Journal of Hymenoptera Research
LITERATURE CITED
Arnett, R. H., Jr & G. A. Samuelson (eds). 1986. The insect and
spider collections of the world. Gainesville, E.J. Brill /Fauna
& Flora Publications, 220p.
Azevedo, C. O. 1991. Especie nova e notas sobre Bakeriella
Kieffer (Hymenoptera, Bethylidae) da regiao de Sao
Carlos, SP. Revista Brasileira de Entomologia 35(3): 535-8
Eady, R. D. 1968. Some illustrations of microculpture in the
Hymenoptera. Procedings of the Royal Entomological Soci-
ety of London, series A General Entomology 43(3-6): 66-72.
Evans, H. E. 1964. A Synopsis of the American Bethvlidae
(Hymenoptera : Aculeata). Bulletin of the Museum of Com-
parative Zoology, Harvard University 132(1): 1-222.
Evans, H. E. 1965. Further studies on Neotropical Epynni
(Hymenoptera: Bethylidae). Psychef Cambngde) 72(4):
265-78.
Evans, H. E. 1969. A revision of the genus Epyris in the
Americas (Hymenoptera: Bethylidae). Transactions of
the American Entomological Society 95(2): 181-352.
Evans, H. E. 1979. A reconsideration of the genus Bakeriella
(Hymenoptera: Bethylidae). Journal of the Nexv York En-
tomological Society 87(3): 256-66.
Gordh, G. and L. Moczar. 1990 A catalog of the world
Bethylidae (Hymenoptera). Memoirs of the American En-
tomological Institute 46: 1-364.
Kieffer, J. J 1910a. Description de Nouveaux Micro-
hymenopteres du Bresil. Annates de la Societe de
Entomologique de France 78: 287-348.
Kieffer, J. J. 1910b. Description de Nouveaux Microhy-
menopteres du Bresil. Annates de la Societe de
Entomologique de France 79: 31-56.
J. HYM. RES.
Vol. 3, 1994, pp. 151-156
Localization of the Female Sex Pheromone Gland in
Cotesia rubecula Marshall (Hymenoptera: Braconidae)
Scott A. Field And Michael A. Keller
Department of Crop Protection, University Of Adelaide, Waite Campus, Glen Osmond, South Australia 5064, Australia
Abstract - A combination of behavioural, histological and electron microscopic techniques was used to localize the female
sex pheromone gland in the parasitic wasp Cotesia rubecula (Marshall) (Hymenoptera: Braconidae). The genital capsule was
identified as the source of the sex pheromone by presenting males with a choice of body parts in a flight tunnel. Histological
examination and electron microscopy further reduced the possibilities for pheromone production to two structures. One of
these was a gland in a position similar to that reported for other braconid wasps, and which was associated with surface pores
but lacked any obvious secretory products. The other was an active secretory gland which is associated with the moving parts
of the ovipositor. Further studies aimed at isolating and identifying the chemicals involved are required to resolve this issue.
The presence of sex pheromone glands has
been demonstrated in a number of species of para-
sitic wasps (Weseloh 1976, 1980; Tagawa 1977,
1983). Although unable to localize the sex phero-
mone source more precisely than the abdominal
region, Vinson (1978) suggested that Dufour's
gland produced a sex identification pheromone in
Cardiochiles nigriceps. Obara and Kitano (1974) first
demonstrated that the source of the female sex
pheromone in Cotesia glomerata is the tip of the
abdomen, near the ovipositor. Tagawa (1977) later
described a pair of secretory glands located at the
base of the second valvifer on the ninth abdominal
tergite of this species, which he suggested was
responsible for sex pheromone production. A sub-
sequent study (Tagawa 1983) revealed the exist-
ence of similar glands in a corresponding position
in all seven braconid species examined. Upon
revising his initial findings (Weseloh 1976),
Weseloh (1980) likewise concluded that paired
epidermal glands on the last abdominal tergite
were the sex pheromone source in Cotesia
melanoscelus.
Previous field observations (Keller unpub-
lished) established that courtship in male Cotesia
rubecula (Marshall) (Hymenoptera: Braconidae) is
elicited by a female sex pheromone. The present
study combined behavioural tests with micro-
scopic techniques in order to identify the position
of the sex pheromone gland in C. rubecula.
MATERIALS AND METHODS
Behavioural Experiments. — C. rubecula was
reared in the laboratory on larvae of Pieris rapae L.
using the methods of Keller (1990). Females be-
tween one and four days of age were frozen at -15°
C for approximately 30 minutes before being dis-
sected.
Using the same experimental protocol, we
performed two experiments to test the respon-
siveness of males to different female body parts. In
the first, the head and mesosoma (thorax plus first
abdominal segment) were tested against the
metasoma (remaining abdominal segments), and
in the second the genital capsule (terminal ab-
dominal segment plus ovipositor) was tested
against the remaining anterior portion of the
metasoma.
A glass microscope slide was divided into two
halves and the two body parts being tested were
placed randomly in the middle of either half,
presenting a choice to the males. The slide was
clamped at its midpoint and held horizontally at a
152
Journal of Hymenoptera Research
height of 30 cm by a small metal stand. Tests were
conducted in a flight tunnel (Keller 1990) at a wind
speed of 30 cm/s. Males were kept in a separate
cage and removed for testing individually with a
minimum of disturbance. They were released 30
cm downwind of the slide and observed to deter-
mine (1) which half of the slide they landed on, (2)
whether subsequently they oriented to either body
segment and performed typical elements of court-
ship (Field and Keller 1993), and (3) whether they
attempted to copulate with either body segment.
The same males were then tested again, but
this time the female body segments were washed
in diethyl ether prior to testing. This solvent re-
moved all contaminating pheromone from the
surface of the body segment (Golub and
Weatherston 1984). Therefore any subsequent
emission of pheromone could be due only to con-
tinued leaking of the pheromone onto the surface
from an internal glandular reservoir. The prefer-
ence of courting males for a particular segment
after washing would therefore localize the phero-
mone gland to somewhere within that segment.
Most males were successful in navigating to
the slide; those that did not were discarded after a
few attempts. To lessen the effects of pheromone
contamination and depletion of pheromone re-
serves, the stand was washed with diethyl ether
and body segments and slides were replaced at
regular intervals throughout the course of the
experiments. Variation in attractiveness of females
was assumed to be negligible. Fifty males were
tested in each experiment and data were analysed
using two-tailed binomial tests (Zar 1984).
Scanning Electron Microscopy (SEM). — Two
methods were used to prepare specimens for SEM
observations. In the first, newly-dissected speci-
mens were immersed in Peterson's KAA (kero-
sene (10%), 95% alcohol (75%), glacial acetic acid
(15%)) prior to washing in 0.1 M phosphate buffer
and dehydration through an alcohol series. This
technique is commonly used to preserve larval
insects because it distends structures that may
otherwise lose shape (Smithers 1981).
The second method was to "clear" the speci-
mens by gently warming in 10% KOH for 30-60
minutes. This removed most tissue and allowed
dissection of the specimens so that both internal
and external cuticular structures could be exam-
ined under SEM. Following dehydration, the KAA-
treated and cleared specimens were critical-point
dried in an EMSCOPE CPD 750 and mounted on
TAAB aluminium stubs using Acheson Electrodag
915. They were coated by vacuum evaporation
with carbon and gold palladium and examined in
an ETEC Autoscan scanning electron microscope
at an accelerating voltage of 20 kV.
Light Microscopy (LM). — Genital capsules were
removed and immersed for 4 hours in fixative (3%
glutaraldehyde/3% formaldehyde made up in 0.1
M phosphate buffer, pH 7.4, to which had been
added 2.5% polyvinyl pyrrolidone). They were
washed in 0.1 M phosphate buffer overnight and
then dehydrated by passing through a series of
alcohols. After washing in propylene oxide, they
were infiltrated with increasing concentrations of
TAAB epoxy embedding resin over 48 hours and
then embedded in resin by curing at 60° C for a
further 48 hours.
Using glass knives in a Sorvall MT2-B "Porter-
Blum" ultramicrotome, serial transverse sections
of 0.5 urn thickness were cut, starting from near the
base of the ovipositor and proceeding anteriorly.
These were stained using 0.025% Toluidine Blue
in 0.5% Borate buffer and photographed with a
Wild MPS 45 photoautomat on Ilford Pan-f 50
ASA film. Agfa x 3 light green and Wratten 58
filters were used to achieve suitable contrast. Males
were also subject to the same processing proce-
dures and serial sections were cut from the poste-
rior tip of the abdomen.
Transmission Electron Microscopy (TEM). -Tis-
sue samples were prepared in the same mariner as
for LM. Gold to silver sections (0.08 mm) were cut
with a Diatome diamond knife using a Reichert-
Jung Ultracut and stained with saturated uranyl
acetate in 70% alcohol for 20 minutes, followed by
lead citrate for 12 minutes (Reynolds, 1963). They
were examined in a JEOL 100S transmission elec-
tron microscope at an accelerating voltage of 80
kV.
RESULTS
Behavioural Experiments. — In the first experi-
ment, neither the head/mesosoma nor the
metasoma was found to be more attractive when
the segments had not been washed in ether. How-
ever, when surface contamination had been re-
moved with ether, males responded preferentially
toward the metasoma, indicating that it was the
source of the sex pheromone (Table 1).
Volume 3. 1994
153
Fig. 1. SEM of lateral view of genital capsule in Cotesia rubecula. Arrow indicates position of hair plate and pores (see Fig. 2);
n - ninth tergite; o - ovipositor. Scale bar = 100 urn
Fig. 2. High magnification SEM of hair plate and pores. Scale bar = 10 urn
Fig. 3. Transverse section of genital capsule, showing under LM two possible sex pheromone-producing glands: gp - glands
associated with surface pores; gb - gland at base of genital capsule; hp - hair plate. Scale bar = 30 urn
Fig. 4. TEM of basal section of genital capsule, p - pore canals; s - secretory droplets. Scale bar = 5 urn
154
Journal of Hymenoptera Research
Table 1 . Response of male Cotesia rubecula to a choice between the metasoma and the
head+mesosoma of females, washed in ether or unwashed. * indicates significant
differences (P<0.001).
Treatment
Number Responding
Response
Metasoma
Head+Mesosoma
Unwashed
Washed
Land
Court
Copulate
Land
Court
Copulate
23
23
10
36
36
19
27
27
22
l"
r
r
Table 2. Response of male Cotesia rubecula to a choice between the genital capsule
and the anterior portion of the metasoma of females, washed in ether or unwashed.
* indicates significant differences (P<0.001).
Treatment
Number Responding
Response Anterior Metasoma Genital Capsule
Unwashed
Washed
Land
17
Court
18
Copulate
Land
Court
Copulate
19
3
4
3
31
27
1*
28*
21*
I
In the second experiment, there was no prefer-
ence shown by males for either the unwashed
genital capsule or the unwashed anterior portion
of the metasoma in terms of frequencies of landing
and courting (Table 2). Several males moved be-
tween the body segments after landing in close
proximity to one of them. More males attempted
to copulate after courting an unwashed metasoma
(18/18, plus one that courted to the genital capsule
initially) than an unwashed genital capsule (1/
27). After these body parts were washed with
ether, males showed a clear preference for the
genital capsule in terms of landing and courting,
but not copulating.
Microscopy. — Having identified the genital
capsule as the source of the sex pheromone, we
examined KAA-treated genital capsules under the
SEM to determine whether there were any surface
pores that could serve as outlets for pheromone
secretion. The only pores found were positioned
slightly below and posterior to a mechanosensory
hair plate (Figs. 1,2). Dissection of a cleared speci-
men revealed the internal attachment site of these
mechanosensory hairs to be at the base of the
second valvifer, which is an internal cuticular
plate comprising part of the ovipositor system (see
Snodgrass, 1935).
The LM sections revealed only one region of
tissue that was a possible candidate for the sex
pheromone source. A pair of glands was located in
a position corresponding closely to that of the
pores, i.e., just below the hairs and appearing
immediately before the hairs in serial transverse
sections (Fig 3). No pores were apparent, but this
Volume 3, 1994
155
may have been a result of the small proportion of
sections cut (approx. 1/10) that were actually re-
trieved, stained and mounted for examination.
The results of ultrastructural examination of this
gland were inconclusive and failed to offer any
indication of the organelles present or whether it
was secretory in nature.
TEM examination revealed the presence of a
second pair of secretory glands (Fig. 4), located at
thebase of the genital capsule and adjacent to folds
of membrane which are presumably stretched and
compressed during movements of the ovipositor.
The ultrastructure of these glands was also indis-
tinct, but pore canals leading to the surface and an
abundance of secretory bodies were visible on or
just under the surface of the cuticle.
DISCUSSION
The behavioural experiments clearly demon-
strate that the genital capsule is the source of the
sex pheromone in C. rubeculn. With all surface
pheromone removed, males were attracted to the
metasoma in preference to the head/mesosoma
and, when given a further choice, made their
initial approach more often to the genital capsule
rather than to the anterior portion of the metasoma.
This is indicative of the continuous emission of
pheromone from an internal reservoir located in
the genital capsule, which provides an olfactory
cue to the searching wasp.
There are, however, two anomalous results
which require explanation. Firstly, a single male
chose the washed head/mesosoma over the
metasoma in the first experiment. This could have
been due to incomplete washing of the body parts.
Secondly, although significantly more males
landed near and courted the genital capsule as
opposed to the anterior portion of the metasoma,
they attempted copulation with the genital cap-
sule less frequently. This is not considered to be
evidence that the sex pheromone gland is in the
anterior portion of the metasoma. Rather, it sug-
gests that a visual cue is involved in triggering
attempts at copulation by the male. Upwind flight,
landing, orienting and courting all appear to be
stimulated entirely by chemical cues. The pres-
ence of a particular threshold amount of chemical
may "prime" males so that they are ready to
mount and copulate after a certain amount of
courtship, but an object of appropriate size, shape
and colour in the vicinity of a pheromone source
may be required to release copulation behavior.
One male did attempt copulation with the tiny
genital capsule (0.5 mm in length), so the phero-
mone alone may be sufficient to elicit attempts at
copulation in some cases.
The histology and electron microscopy failed
to resolve the exact location of the sex pheromone
gland within the genital capsule, but narrowed the
possibilities to two structures. One of these is a
group of cells located directly underneath surface
pores which are adjacent to the mechanosensory
hair plate on the genital capsule (Figs. 1-3). The
hair plate was in turn juxtaposed to the second
valvifer, which places this gland in approximately
the same position as the putative pheromone
glands indicated by Tagawa (1977, 1983) for a
number of other closely-related braconids. Al-
though this constitutes circumstantial evidence
that this is the sex pheromone gland in C. rubecula,
the absence of any clearly defined secretory prod-
ucts in the ultrastructural examinations leaves
some uncertainty. The pores associated with these
glands required thorough washing before they
became visible by SEM.
Another possible source of sex pheromone is
a gland located further toward the base of the
genital capsule. The most striking feature of this
gland is an abundance of associated secretory
bodies amassed on and under the surface of the
cuticle which are visible under TEM (Fig. 4). One
possibility is that these secretions act as a lubricant
for the ovipositor as it is extended and retracted
from the metasoma . However, if it is the sex phero-
mone gland, then such movements would un-
doubtedly smear the secretions over the cuticle,
which would be ideal for pheromone dispersal.
An additional possibility is that the one secretion
has the dual function of lubrication and sex attrac-
tant. In the absence of isolation and identification
of the pheromone, the question of which of the two
glands is the source of sex pheromone in C. rubecula
will remain unresolved.
156
Journal of Hymenoptera Research
ACKNOWLEDGEMENTS
We thank Chris Leigh for the generous use of his facili-
ties for histological processing, preparation of TEM sections
and help with the SEM, Dr Adam Lockett for the use of his
photographic equipment, Frances FitzGibbon for printing
the micrographs and Paul Dangerfield for proofreading the
manuscript.
LITERATURE CITED
Field, S.A. and MA. Keller. 1993. Courtship and intersexual
signalling in the parasitic wasp Cotesia rubecula. journal
of Insect Behaviour 6: 737-750
Golub, MA. and Weatherston, I. 1984. Techniques for ex-
tracting and collecting sex pheromones from live insects
and from artificial sources. In Hummel, H. and Miller,
T., eds. .Techniques in pheromone research. Springer- Verlag,
New York. 464 pp.
Keller, MA. 1991 . Responses of the parasitoid Cotesia rubecula
to its host Pieris rapae in a flight tunnel. Entomologia
Experiementalis et Applicata, 57: 243-249.
Obara, M. and Kitano, H. 1974. Studies on the courtship
behaviour of Apanteles glomeratus L. -1. Experimental
studies on releaser of wing-vibrating behaviour in the
male. Kontyu.il: 208-214.
Reynolds, E.S. 1963. The use of lead citrate at high pH as an
electron-opaque stain in electron microscopy. Journal of
Cell Biology, 17: 208-212.
Smithers, C. 1981 Handbook of Insect Collecting. A.H. and A.W.
Reed, Sydney. 120 pp.
Snodgrass, RE. 1935. Principles of Insect Morphology. McGraw-
Hill Book Co., New York. 667 pp.
Tagawa, J. 1977. Localization and histology of the sex phero-
mone-producing gland in the parasitic wasp, Apanteles
glomeratus. journal of Insect Physiology, 23: 49-56
Tagawa, J. 1983 Female sex pheromone glands in the para-
sitic wasps, genus Apanteles. Applied Entomology and
Zoology, 18: 416-427.
Vinson, SB. 1978. Courtship behavior and source of a sexual
pheromone from Cardiochiles nigriceps. Annals of the
Entomological Society of America, 71: 832-837.
Weseloh, R.M. 1976. Dufour's gland: Source of sex phero-
mone in a hymenopterous parasitoid. Science, 193: 695-
697.
Weseloh, R.M. 1980. Sex pheromone gland of the gypsy moth
parasitoid, Apanteles melanoscelus : revaluation and ul-
trastructural survey. Annals of the Entomological Society
Of America, 73: 576-580.
Zar, J.H. 1984 Biostatistical Analysis. 2nd edition. Prentice-
Hall, Englewood Cliffs. 718 pp.
J. HYM. RES.
Vol. 3, 1994, pp. 157-173
A Review of the World Species of Orthomiscus Mason
(Hymenoptera: Ichneumonidae: Tryphoninae)*
Virendra K. Gupta
Entomology and Nematology Department, University of Florida, Gainesville, Florida 32611-0620, U.S.A.
Abstract. — The world species of the exentenne genus Orthomiscus Mason are reviewed. O. amurensis Kasparyan is
transferred to Kristotomus , leaving six species under Orthomiscus in the Holarctic Region: O. eridolius Kasparyan, medusae
Kasparyan, pectoralis (Hellen), -platyura Mason, simplex (Mason) and unicinctus (Holmgren). Diagnostic characters, species
relationships and a key to the species are given. Diagnostic characters are illustrated by diagrams and photographs.
INTRODUCTION
The genus Orthomiscus was described by Ma-
son (1955) to accommodate two new Nearctic
species, O. platyura and O. leptura. It belongs to the
Kristotomus-Complex of genera in the tribe
Exenterini (Gupta 1990), including Kristotomus
Mason, Kerrichia Mason and Orthomiscus Mason.
This group is characterized by having the apical
rim of the hind tibia with a fringe of long, close
bristles on the inner side and with a flat polished
area on the lower and inner sides between the
apical fringe of bristles and the tarsal socket; by
having the hind tibia widest at apex and truncate;
and by its short ovipositor which hardly surpasses
the tip of abdomen. These genera also exhibit the
usual exenterine characters such as the presence of
only one spur on the middle tibia and the absence
of spurs on the hind tibia.
Mason (1962) commented on the relationships
of the genus and included two European species:
Exenterus unicinctus Holmgren, 1858 (with which
he synonymized O. leptura) and Cteniscus pectoralis
Hellen, 1951. Kasparyan (1976, 1986, 1990) de-
scribed three additional new species from Russia:
O. medusae, O. amurensis, and O. eridolius. Gupta
(1990) transferred the Japanese Kristotomus sim-
plex Mason, 1962 to this genus. In the present
study, O. amurensis Kasparyan is transferred to the
genus Kristotomus as it exhibits most of the charac-
ters of that genus, like the tapered and slender
ovipositor and sheaths, lower valve of ovipositor
without teeth and egg with a simple stalk and
anchor. Orthomiscus thus is recognized as a Hol-
arctic genus with six species.
Biology. — Not much is known about the biol-
ogy of Orthomiscus except that O. pectoralis has
been reared from late larval cimbicids (Trichiosoma
lucorum L. and T. tibiale Stephens) that feed on
birch and ash (Betula and Crataegus) in Europe
(Jussila 1975).
Material. — This paper is based on the speci-
mens present in the collection of the American
Entomological Institute, Gainesville (GAINES-
VILLE). The types of the Russian species were
borrowed from the collections of the Zoological
Institute, Russian Academy of Science, St. Peters-
burg (ST. PETERSBURG) through the kindness of
Dr. D. R. Kasparyan. Under synonymy of various
species, the location of the type is indicated by the
'Florida Agricultural Experiment Station Journal Series R-03128.
158
Journal of Hymenoptera Research
city where the type is housed (see Townes, Momoi
& Townes 1965 for details). The key to the species
is based on the females which are more diagnostic.
Genus ORTHOMISCUS Mason
Orthomiscus Mason, 1955. Canadian Journal of Zoology, 33:
63.
Type: Orthomiscus platyura Mason; original designation.
Taxonomy: Mason 1962: 1273, Gupta 1990: 7; Kasparyan
1990: 36.
decurved, not tapered apically and not extending
beyond apex of abdomen. Upper valve of oviposi-
tor flattened dorsally in basal half. Lower valve of
ovipositor with an obliquely serrated edge (Figs.
10, 12, 18, 29). Ovipositor sheaths short, broadly
triangular or sometimes a little slender. Female
subgenital plate thin, creased medially and pinched
apically. Egg stalk short, anchor simple, button-
like or flattened and enclosing part of the egg
(Figs. 4-14).
Diagnostic characters. — Head subcuboidal,
swollen behind eyes. Mandibular teeth equal or
subequal. Occipital and hypostomal carinae com-
plete. Occipital carina bent inwards, meeting
hypostomal carina at a right angle above base of
mandible by a distance equal to the basal width of
mandible. Epomia distinct. Epicnemial carina ex-
tending at least to the middle of hind margin of
pronotum. Basal area of propodeum confluent
with areola or only partly separated from it. Legs
slender, elongate. Tarsal claws with 2-6 basal teeth.
Areolet present, oblique, sometimes second
intercubitus partly weak. Nervellus inclivous, in-
tercepted in its lower 0.1 5 to 0.3. Tergitel elongate,
usually about 2.0x as long as its apical width, its
dorsomedian and dorsolateral carinae sharp and
distinct; dorsolateral carina usually extending to
the entire length of the tergite and passing just
dorsad of spiracle and not broken there. Tergite 2
usually with a pair of strong to weak oblique
basolateral grooves. Ovipositor short, stout and
SPECIES RELATIONSHIPS
On the basis of egg structure, the six species of
Orthomiscus form two distinct species groups.
Group 1 comprises platyura Mason (type-species),
medusae Kasparyan and eridolius Kasparyan (Figs.
4, 7, 9), where the egg stalk is short and wide and
the anchor is four-pronged and encloses part of
the egg. Group 2 comprises pectoralis (Hellen),
unicinctus (Holmgren) and simplex (Mason) (Figs.
8, 12, 30) where the egg has a simple short stalk and
a short button-like anchor. In the latter group, the
stalk arises apicoventrally in pectoralis and
unicinctus while in simplex it is central and the
anchor is flat and oval.
Gupta (1990) stated that the plesiomorphic
state of the egg in Cteniscini was probably an egg
with a short stalk and anchor. This type of egg is
seen in some generalized members of the genus
Kristotomus (like claviventris, santoshae and
ctenonyx). Kristotomus also exhibits more general-
eridolius
platyura
T2 with deep
grooves
modified egg
anchor/stalk
pectoralis
Kristotomus
stalk
medusae
wide mandible
(reversal)
body
punctate
unicinctus
second segment
-of maxillary palp
swollen
simplex
simple egg
anchor/stalk
mandible
widened
apically
Diagram depicting the relationships of Orthomiscus species
Volume 3, 1994 159
ized characters from which Orthomiscus and Under group 1, Orthomiscus platura has deep
Kerrichia may be derived. The Orthomiscus species oblique grooves on tergite 2 separating it from O.
closest to Kristotomus appears to be O. simplex, medusae and eridolius. In medusae the mandibles are
which was originally described in Kristotomus and widened apically and the lower tooth is longer
considered a primitive member of the genus by than the upper (reversal). It also has the basal vein
Mason (1962), primarily because the mandible is more strongly curved medially (Fig. 38). Under
slightly widened apically in this species and in group 2, O. simplex has the mandible widened
several species of Kristotomus. O. medusae also apically with the lower tooth longer than the up-
exhibits a widened mandible but the egg structure per. This is not the condition in O. pectoralis and O.
is very different. I consider that the widened man- unicinctus. O. pectoralis has the body, including
dible of these two species arose by convergence. face, punctate and the propodeum rugose and
The relationships of all Orthomiscus species may punctate in places. O. unicinctus has the second
be depicted as follows: segment of maxillary palp swollen (Fig. 39).
KEY TO THE SPECIES
la Basal vein of fore wing slightly curved close to its junction with the medius vein. Petiolar area of propodeum
almost pentagonal, about 0.75x the combined length of areola and basal area (Fig. 3). Apical transverse
carina of propodeum angulate at its junction with lateral and median longitudinal carinae (Fig. 23).
Mandibular teeth equal in length (Figs. 2, 15). Hypostomal carina not raised at its junction with occipital
carina. Egg stalk situated apicoventrally or in the middle 2
lb Basal vein of fore wing more strongly curved, curvature in the middle of the vein. Petiolar area of
propodeum small, circular, about half the combined length of areola and basal area. Apical transverse
carina of propodeum almost evenly curved between the lateral longitudinal carinae. Mandible slightly
widened apically and the lower tooth longer than the upper (Figs. 27, 31). Hypostomal carina slightly
raised at its junction with occipital carina. Egg stalk situated in the middle of the egg (Figs. 7, 30) ... 5
2a Second segment of maxillary palp swollen, about 2. Ox as wide as the rest. Abdomen black dorsally, tergites
2-3 often with triangular yellow patches or their apical margins yellow. Pronotum yellow. Mesopleurum
partly yellow. Mesosternum yellow. Ovipositor slender (Figs. 20, 21). Egg stalk short and apicoventral
(Figs. 6, 11). Egg anchor button shaped. Holarctic 2. unicinctus (Holmgren)
2b Second segment of maxillary palp similar to the others, not swollen. Abdomen beyond first tergite black or
brown with middle segments of ten with large yellowish brown patches. Egg stalk and anchor various
(Figs. 4, 8, 9) 3
3a Face with definite punctures. Propodeum rugose and punctate in places. Tergites 4-5 as long as wide.
Ovipositor sheaths slender. Propleurum yellow. Mesopleurum largely black. Abdominal tergites black
with narrow apical yellow stripes. Egg stalk short and apicoventral (Figs. 8, 12). Egg anchor button-
shaped. Eurasia, Japan 1. pectoralis (Hellen)
3b Face polished, only with sparse punctures. Propodeum polished to subpolished, without distinct punc-
tures or rugosities. Tergites 4-5 transverse. Ovipositor thick, its sheaths triangular (Figs. 18,25). Egg oval,
its stalk very short, central. Egg anchor partly enclosing the egg (Figs. 4, 5, 9, 14) 4
4a Tergite 2 with deep oblique basolateral grooves. Abdominal tergites beyond first largely reddish brown.
Pronotum, mesopleurum ventrally and mesosternum yellow. Nervellus intercepted at its lower 0.25-0.3.
Egg stalk short (Fig. 4), anchor 4-pronged (Fig. 5). U.S.A., Canada 5. platyura Mason
4b Tergite 2 without oblique grooves. Abdominal tergites black, with margins of tergites 2-3 (sometimes 4-5
also) yellow. Mesosternum of female black. Nervellus intercepted at its lower 0.15-0.2. Egg anchor (Fig.
9) broad and partly enclosing the egg. Russia: Far East 6. eridolius Kasparyan
5a Mesopleurum of female yellow ventrally, in male with yellow spots. Tarsal claws with 4-5 teeth. Female
subgenital plate sharply creased medioventrally. Ovipositor slender (Fig. 29). Egg with a simple, oval
anchor (Fig. 30). japan 3. simplex (Mason)
5b Mesopleurum black. Tarsal claws with a pair of weak teeth. Female subgenital plate convex ventrally, not
strongly creased. Ovipositor thick (Fig. 10). Egg with a 4-lobed anchor (Fig. 7). Russia: Far East, Japan
4. medusae Kasparyan
160
Journal of Hymenoptera Research
1. Orthomiscus pectoralis (Hellen)
Figs. 8, 12
Ctemscus pectoralis Hellen 1951. Notulae Entomologicae
Helsingfors, 31: 31. F. des. Type: Female, Finland: Esbo
(HELSINKI). — Kernch 1952. Bulletin of the British
Museum (Natural History) Entomology Series, 2 (6):
432. F. des., distr.
Orthomiscus pectoralis: Mason, 1962. Canadian Entomologist,
94: 1 274. n. comb. Japan: Nagano Prefecture: Kamikochi.
— Townes, Momoi & Townes, 1965. Memoirs of the
American Entomological Institute, 5: 109. cat. — Jussila,
1975. Acta Entomologica Fennica,41 (2): 53. des. of male,
host records. — Kasparyan, 1976. Entomologicheskoe
Obozrenie, 55: 137. — Kasparyan, 1977. Nasekomye
Mongoliaca, 5: 463. Mongolia. — Kasparyan, 1990. Fauna
of USSR Insecta, Hymenoptera, 3 (2): 43. key, des., fig.
Russia.
No specimens of this species were available
for study. Kerrich (1952) and Kasparyan (1990)
have provided detailed descriptions of this spe-
cies. The description that follows is adapted from
Kerrich (1952).
Diagnosis. — This species appears different
from the others by having the face and thorax
extensively punctate and the propodeum rugoso-
punctate (cf. Kerrich, 1952 description). All other
species have only weak to indistinct body puncta-
tion. Tergites 4-5 as long as wide. The egg has a
short apicoventral stalk and a small button-like
anchor (Fig. 8).
Male and female. — Face moderately punctate.
Clypeus with fine punctures. Epomia short and
weak. Mandibular teeth equal in length. Notauli
sharply impressed. Scutellum strongly convex,
finely punctate. Epicnemial carina incomplete
medially. Sternaulus superficially impressed.
Propodeum rugose to punctate in places. Areola
about as long as wide, confluent with basal area.
Abdomen elongate and slender. Abdomen elon-
gate and slender. Tergite 1 about 1.6x as long as its
apical width, its dorsomedian and dorsolateral
carinae strong. Postpetiole with a pitlike depres-
sion bounded by carinae. Tergite 2 with small but
distinct thyridia. Tergites 4-5 as long as their basal
width. Apical abdominal segments compressed in
female. Female subgenital plate less sharply folded
than in O. unicinctus (with which Kerrich com-
pared this species). Ovipositor sheaths slender.
Ovipositor comparatively stout as is usual in the
genus (Fig. 12).
Color. — Ground color black. Mouthparts,
clypeus, face, and temples broadly, yellow. Scape
and pedicel pale brownish. Thorax and
propodeum black, with pronotum, scutellum,
metascutellum, a mark above epicnemial carina,
mark above middle coxa, tegula and subtegular
ridge, yellowish brown. Propleurum yellow. Fore
and middle coxae and trochanters yellow. Fore
and middle femora yellowish brown, their tibiae
and tarsi paler. Hind coxa reddish brown. Hind
femur, tibia and tarsus more brownish to reddish
brown. Abdomen black above, yellow below.
Tergites 1-6 with narrow, pale apical margins and
tergite 7 with an obscure, triangular yellow mark.
Ovipositor sheaths yellow, infuscate above and
rufous towards apex.
Egg. — (Fig. 8) Egg with a short apicoventral
stalk and a button-like anchor.
Length. — 5.0-10.0 mm. Fore wing 3.8-8.5 mm.;
Ovipositor about 0.5 mm.
Distribution. — Europe (Finland), Russia,
Mongolia, Japan.
Mason (1962) reported this species from Japan
and included two females from Nagano Prefec-
ture: Kamikochi, collected by Townes in 1954.
These specimens could not be located in the AEI
collections.
2. Orthomiscus unicinctus (Holmgren)
Figs. 6, 11, 19-23,36,39
Exen terusumanct us Holmgren, 1858. SvenskaVetensk.-Akad.
Handlirsch (n. f), 1: 234. M, F. key, des. Lectotype
(selected by Roman, 1914), Female, Sweden: Lapland:
Tarna (STOCKHOLM).
Exenterus macrocephalus Holmgren, 1858. Svenska Vetensk.-
Akad. Handlirsch (n. f), 1: 243. M. des. Type: Male,
Sweden (STOCKHOLM). Syn. by Roman, 1914.
Ctemscus macrocephalus:]acob &Tosquinet, 1896. Annalesde
la Societe Entomologique Beige, 34: 108. new comb.
Cteniscus unicinctus: Roman, 1914. Arkiv for Zoologie, 9 (2):
17. syn., des. — Kerrich, 1942. Transactions of the Soci-
ety for British Entomology, 8 (2): 63-64. Britain. —
Kerrich, 1952. Bulletin of the British Museum (Natural
History) Entomology Series, 2 (6): 434. key, des., fig.
Localities in Britain, France, Belgium, Germany, Swe-
den.
Orthomiscus leptura Mason, 1955. Canadian Journal of Zool-
ogy, 33: 64. M, F. key, des., fig. Type: Female, U.S.A.:
New Hampshire: Pinkham Notch (AEI, Gainesville)
Examined. Syn. by Mason, 1962.
Orthomiscus unicinctus: Mason, 1962. Canadian Entomolo-
gist, 94: 1274. syn., distr. Japan: Sapporo. — Townes,
Momoi & Townes, 1965. Memoirs of the American En-
tomological Institute, 5: 109. cat. — Mason, 1966. Cana-
dian Entomologist, 98: 48. key. Japan. — Carlson, 1979.
Catalog of Hymenoptera in America North of Mexico, 1 :
377. cat. U.S.A., Canada. — Kasparyan, 1990. Fauna of
Volume 3. 1994
161
USSR Insecta, Hymenoptera, 3 (2): 45. key, des., fig.
Russia.
Diagnostic features.— Second segment of max-
illary palp widened and flat, about 2.0x as wide as
the rest (Fig. 39). Oblique impressions on tergite 2
faint. Ovipositor comparatively more slender than
in the previous species (Figs. 20, 21). Egg with a
short apicoventral stalk and a button-like anchor
(Figs. 6, 11). Temple yellow only on its basal half.
Scutellum black except at apex. Hind coxa with
blackish brown patches. Abdomen largely black
and with yellow patches on tergites 2 and 3. Tho-
rax of male largely black, that of female generally
yellow in lower half, but color variable.
Male and female— Flagellum with 23-24 seg-
ments in male and 23-26 segments in female. First
segment about 1.4x as long as the second. Second
segment of maxillary palp widened and flat, about
2.0x as wide as the rest of the segments (Fig. 39).
Mandibular teeth equal in length but lower tooth
more prominent than the upper. Hypostomal ca-
rina of normal shape, not conspicuously raised at
its junction with occipital carina. Vertex widened
posteriorly. Interocellar distance 0.25x the
ocellocular distance.
Thorax subpolished. Mesoscutum rather
strongly convex. Notauli distinctly impressed up
to middle of mesoscutum. Lateral carinae of scutel-
lum distinct only at base of scutellum. Epomia
sharp across pronotal collar. Sternaulus and
epicnemial carina moderately impressed.
Propodeum convex, subpolished. Areola (Fig. 23)
widened at its junction with costulae, confluent
with basal area. Basal vein of fore wing uniformly
but weakly arched (Fig. 36). Nervellus intercepted
in its lower 0.25-0.33.
Abdomen subpolished. Tergite 1 slender (Fig.
19), its dorsomedian carinae strong and extending
to 0.7-0.8 the length of the tergite, its dorsolateral
carina sharp and extending to the entire length of
the tergite. Tergite 2 with distinct thyridia and
faint oblique grooves. Female subgenital plate
folded medially and pointed apically. Ovipositor
slender, of uniform diameter (Fig. 21). Ovipositor
sheaths (Figs. 20, 21) comparatively narrow. Egg
anchor small and situated apicoventrally (Fig. 6).
Color. — Ground color black. Thorax almost
wholly black in male and some females. Abdomi-
nal tergites largely black. Scape, pedicel below,
mouthparts, clypeus, face, inner orbits up to top of
eyes, malar space, and temples on lower half,
yellow. Propleurum, lower half of pronotum, its
upper margin, tegula, subtegular ridge, tip of
scutellum, epicnemium, mesosternum, lower half
of mesopleurum, abdominal venter, and sides of
abdomen, yellow . Legs yellow to yellowish brown.
Fore and middle coxae and trochanters yellow.
Hind coxa largely brownish or yellow with large
black or brown patches. Hind femur brown. Hind
coxa largely yellow in specimens from Ontario
and blackish in specimens from Alaska. Tergites 2
and 3 often with triangular yellow patches or with
their apical margins yellow. In males the yellow
along the inner orbits extending only to the level of
antennal sockets, temple yellow on its lower 0.3,
and thorax almost wholly black. The color is quite
variable.
The European specimens were not examined
to ascertain the range of variation. Kerrich (1952:
434) and Kasparyan (1990: 45) have described this
species in detail.
Egg — (Fig. 6, 11, 21). Egg small, oval, with a
short apicoventral stalk and a button-like anchor.
Length. — 6.0-8.5 mm.; fore wing 5.0-7.0 mm.;
ovipositor 0.4-0.6 mm. Kerrich (1952) stated that a
female measuring 4.0 mm. was seen from Lapland.
Specimens examined. — CANADA: Quebec:
Stoneham, 1 male, 1 female, 21.VI.1938, H. & M.
Townes. U.S.A.: New Hampshire: Pinkham Notch,
1 female, 25.VI.1938, H. & M. Townes; Mt. Madi-
son, 3 males, 24.VI.1938, H. & M. Townes. Ver-
mont: L. Willoughby, 2 male, 17, 25.VI.1945, C.P.
Alexander. (Types and paratypes in AEI,
Gainesville). Non-type material: U.S.A.: Oregon:
Corvallis, 2 males, 1 female, V-VII. 1978, H. & M.
Townes. Alaska: Anchorage, 1 female, 6-
16.VII.1976, Petre Rush. CANADA: Ontario:
Cumberland, 1 male, 3 females, VI.1975, L. Ling.
Distribution.— -U.S.A., Canada, Europe (Brit-
ain, France, Belgium, Germany, Sweden), Russia,
Japan.
3. Orthomiscus simplex (Mason)
Figs. 26-30, 42
Kristotomus simplex Mason, 1966. Canadian Entomologist, 98:
46. M, F. key, des., fig. Type: Female, Japan: Nagano
Prefecture: Kamikochi (GAINESVILLE. Examined. —
Kasparyan, 1976. Entomologicheskoe Obozrenie, 55:
150. (Entomological Review, 55: 108). key.
Orthomiscus simplex: Gupta, 1990. Contrib. Amer. Ent. Inst.,
25 (6): 8. n. comb. — Kasparyan, 1990. Fauna of USSR
Insecta, Hymenoptera, 3 (2): 44. key, des., fig. Japan.
162
Journal of Hymenoptera Research
Diagnostic features. — Mandible widened
apically, its lower tooth wider and longer than the
upper (Fig. 27). Hypostomal carina slightly raised
at its junction with occipital carina (Fig. 42). Basal
vein of fore wing curved. Egg reniform-ovate,
with a short central stalk and a simple anchor. Egg
surface coarsely granular. Head and thorax in
female yellow on lower half and black on upper
half, in male largely black. Abdominal tergites
black, their apical margins yellow.
Male and female. — Scape about 1.5x as long as
wide. Flagellum with 24 segments. First flagellar
segment 1.4x as long as the second. Face (and rest
of head also) smooth and shiny. Mandible wid-
ened apically, the lower tooth wider and slightly
longer than the upper (Fig. 27). Malar space 0.4x
the basal width of mandible. Maxillary palp seg-
ments normal, not swollen (Fig. 42). Hypostomal
carina slightly raised at its junction with occipital
carina (Fig. 42). Vertex widened behind eyes (Fig.
26). Interocellar distance 0.4x the ocellocular dis-
tance.
Thorax largely smooth and shiny. Notauli
distinct anteriorly. Lateral carinae of scutellum
distinct. Epomia normal, not very sharp.
Epicnemial carina extending to half the height of
mesopleurum. Sternaulus indistinct. Propodeum
normally areola ted. Petiolar area small and round.
Apical transverse carina uniformly curved be-
tween its junction with lateral longitudinal cari-
nae. Basal vein in fore wing medially curved, more
strongly so than in the preceding species. Second
intercubitus of areolet sometimes absent.
Abdomen smooth and shiny. Tergite 1 about
2.25x as long as its apical width, its dorsomedian
carinae extending up to about 0.75 its length, its
dorsolateral carina complete, sinuate and passing
just above the spiracle (Fig. 28). Tergite 2 with
weak basolateral oblique grooves. Female
subgenital plate strongly creased and pointed
apically. Ovipositor (Fig. 29) somewhat slender
and cylindrical, its teeth weak but present at the
tip. Ovipositor sheaths small and slender. Male
claspers broader than in other species.
Color. — Ground color black. Male compara-
tively darker than the female. Face, clypeus, man-
dibles, lower half of temples, pronotal collar, hind
corner of pronotum, whole of fore and middle
legs, hind coxae and trochanters, and abdominal
venter, yellow. Ventral side of scape and pedicel
and apex of scutellum yellowish-brown. Hind
femur, tibia and tarsus blackish brown (male) to
yellowish brown (female). Pronotum of female
more extensively yellow and mesosternum and
lower part of mesopleurum may also be yellow.
Abdominal tergites black with yellow apical
stripes, particularly on tergites 2-3.
Egg. — Reniform-ovate (Fig. 30). Stalk shorter
than its own diameter, arising near the middle of
the egg. Anchor about 0.33 as long as the egg, flat
and oval. Egg surface coarsely granular.
Length. — 4.5-6.5 mm.; fore wing 4.5-6.0 mm.;
ovipositor about 0.5 mm. long.
Specimens examined. — JAPAN: Nagano Prefec-
ture: Kamikochi, 6 males and 2 females (holotype,
allotype and paratypes), 23-31. VII. 1954, Townes
family (AEI, Gainesville).
Distribution. — Japan.
Discussion. — Mason (1966) placed this species
under Kristotomus, stating that it was the most
primitive member of the genus and closely related
to Ortlnmuscus, particularly in the structure of the
egg and development of the dorsolateral carina of
the petiole. Apart from the widened mandible,
this species shares most characters of Orthomiscus,
particularly the structure of the ovipositor, in which
the lower valve has teeth-like indentations. The
dorsolateral carina of the petiole and the egg struc-
ture also relate it to the present genus.
4. Orthomiscus medusae Kasparyan
Figs. 7, 10, 31,38, 41
Orthomiscus medusae Kasparyan, 1976. Entomologicheskoe
Obozrenie, 55: 140 (Entomological Review, 55 (1): 99).
M, F. des., fig. Type: Female, Russia: Sakhalin: Kunashir
Island (ST. PETERSBURG). Examined — Kasparyan,
1990. Fauna of USSR Insecta, Hymenoptera, 3 (2): 42.
key, des., fig. Russia.
Diagnostic features. — Mandible widened
apically, the lower tooth longer than the upper
(Fig. 31). Hypostomal carina raised at its junction
with occipital carina. Scutellum subconvex. Basal
vein of fore wing strongly curved medially (Fig.
38). Tergite 2 with very faint to indistinct oblique
grooves. Female subgenital plate convex ventrally,
not sharply creased. Egg (Figs. 7, 10) with a short
medioventral stalk and a four-pronged anchor
enclosing the egg. Thorax and abdomen black;
sometimes pronotal collar and presternum yellow
and tergites 2-3 with yellow marks, particularly in
females.
Volume 3. 1 994
163
Male and female. — Flagellum with 23-24 seg-
ments in male and 25-27 segments in female. First
flagellar segment about 1 .3x as long as the second.
Face and clypeus subpolished, with sparse punc-
tures. Malar space about 0.33-0.4.X (male) to 0.4-
0.5x (female) the basal width of mandible. Man-
dible slightly widened apically (Fig. 31), its lower
tooth a little longer than the upper. Maxillary palp
slender, normal (Fig. 41). Vertex widened posteri-
orly, smooth and shiny. In terocellar distance 0.25x
the ocellocular distance.
Epomia present but not very strong. Notauli
distinct to middle of mesoscutum. Scutellum
subconvex, its lateral carinae confined to base.
Epicnemial carina strong, ending in middle of
hind margin of pronotum. Sternaulus weakly in-
dicated anteriorly. Propodeum areolated but its
longitudinal dorsal and sublateral carinae quite
often erased beyond the costulae or absent; com-
bined length of basal area and areola approxi-
mately 1 .5x the length of apical area (according to
the original description). In one paratype and in
the Japanese specimens examined in the AEI col-
lection, the areola is narrow, 2. Ox as long as wide,
confluent with the basal area, and a little widened
at the costulae. Basal vein curved medially (Fig.
38). Tarsal claws with a pair of weak teeth at base.
Tergite 1 2.0x as long as its apical width; its
dorsomedian carinae extending to 0.75 its length
and its dorsolateral carinae strong and complete.
Postpetiole with a median oval depression. Terg-
ite 2 with rather weak oblique basolateral impres-
sions. Female subgenital plate convex, not strongly
creased ventrally, apically narrow and pinched.
Ovipositor stout (Fig. 10). Ovipositor sheaths
broad.
Color. — Ground colorblack. Scape and pedicel
reddish brown dorsally and lighter ventrally. Fla-
gellum yellow. Face, clypeus, mouthparts, and
lower half of temple, yellow. Thorax black with
pronotal collar and prosternum yellow in female.
Apex of scutellum brown. Coxae and trochanters
yellow. Fore and middle legs otherwise yellowish
brown. Hind femur, tibia and tarsus blackish
brown. Abdomen black with tergites 2 and 3 with
triangular yellow marks. Tergite 4 with a faint
yellow mark in the paratype examined. In one
male studied only yellow apical lines are present
on the abdominal tergites.
Egg-— Oval in shape (Fig. 7, 10), with a
medioventral short stalk, its anchor four-pronged,
enclosing part of the egg, somewhat similar to
that seen in O. platyura.
Length. — 7.0-9.0 mm.; fore wing 5.5-7.0 mm.;
Ovipositor about 0.5 mm.
Specimens examined. — RUSSIA: SAKHALIN:
Kunashir Island: Sernovodsk, 1 female (paratype),
26.VIII.1973, D. R. Kasparyan. JAPAN: Nagano
Prefecture: Kamikochi, 1 male, 22. VII.1954, 1 male,
24. VII. 1954 (labeled O. unicinctus Tow.), Townes
family (AEI, Gainesville).
Distribution. — Russia, Japan.
Relationships. — Structurally this species is close
to O. simplex (Mason) from which it differs mainly
in the complex structure of the egg anchor, black
mesopleurum, male tarsal claws with a pair of
weak teeth at base, female subgenital plate convex
but not strongly creased ventrally and by possess-
ing a comparatively thicker ovipositor. The egg
structure relates it to O. plati/ura.
5. Orthomiscus platyura Mason
Figs. 1-5, 15-18,37,40
Orthomiscus platyura Mason, 1955. Canadian Journal of Zool-
ogy, 33: 66. M, F. key, des., fig. Type: Female, U.S.A.:
Rhode Island: Westerly (GAINESVILLE). Examined. —
Carlson, 1979. Catalog of Hymenoptera in America
North of Mexico, 1:377. cat. U.S.A. New Hampshire,
Vermont, California. Canada: Quebec, Ontario.
Diagnostic features. — Areola narrow, parallel-
sided and confluent or partly separated from the
basal area. Tergite 2 with rather deep oblique
basolateral grooves. Ovipositor stout, its sheaths
broadly triangular. Egg with a four-lobed anchor,
as large as the egg. Scutellum mostly yellow. Hind
coxa yellow. Abdominal tergites beyond first red-
dish brown to reddish yellow.
Male and female. — Scape about 1.5x as long as
wide. Flagellum with 24 segments in male and 26
segments in female. First segment 1.32 to 1.35x as
long as the second segment. Face (Fig. 15) pol-
ished, depressed laterally. Clypeus convex and
polished. Mandibular teeth subequal, upper tooth
slightly longer than the lower. Malar space 0.45x
the basal width of mandible. Second segment of
maxillary palp slender and similar to other seg-
ments (Fig 40). Vertex polished, quadrate, wid-
ened behind eyes, 1.75x as wide as eye in dorsal
view. Interocellar distance 0.22x the ocellocular
distance. Temple in lateral view 1.7x to 1.75x as
wide as eye.
164
Journal of Hymenoptera Research
Thorax mostly smooth and shiny.
Mesoscutum convex, pubescent. Notauli deep
and distinct to middle of mesoscutum. Scutellum
flat, with lateral carina confined to base. Prono-
tum somewhat dull, epomia strong and extend-
ing close to upper edge of pronotum. Epicnemium
dull, epicnemial carina strong and extending to
0.75 the height of mesopleurum. Sternaulus indi-
cated on anterior 0.4 of mesopleurum. Propodeum
convex, shiny. Areola narrow, parallel-sided,
about 2.0x as long as wide (Fig. 3), confluent with
basal area or only partly separated from it.
Abdomen shiny. Tergite 1 2. Ox as wide at
apex as at base, about 2. Ox as long as its apical
width, its dorsomedian carinae extending up to
about 0.8 its length (carinae weaker apicaUy, wid-
ened, and enclosing a depressed area between
them), its dorsolateral carina complete and strong
(Fig. 17). Tergite 2 with deep, oblique, basolateral
grooves. Ovipositor thick, decurved (Fig. 18).
Ovipositor sheaths short, broadly triangular.
Color. — Black and yellow. LowerO. 66 of head,
scape, flagellum beyond 4th segment, pronotum,
scutellum, metascutellum, presternum, mesoster-
num, lower part of mesopleurum, fore and middle
coxae and trochanters, sternites, ovipositor sheaths,
and subgenital plate, yellow. Legs otherwise and
abdomen (except tergite 1 ), yellowish brown. Terg-
ite 1 black. Brown to blackish patches present on
abdominal tergites and pronotum. Wings hyaline.
Egg— (Figs. 4, 5, 18). Short, oval, ventrally flat
and bearing on nearly half its length a short but
wide stalk; egg anchor large and enclosing most of
the egg. Anchor bearing two long posterior and
two long anterolateral arms that extend toward
the dorsal surface of the egg.
Length. — 6.0-8.0 mm.; fore wing 5.0-6.5 mm.;
ovipositor about 0.4 mm.
Specimens examined. — CANADA: Ontario:
Cumberland, 1 female, ll.VI.1975,L. Ling. U.S.A.:
Rhode Island: Westerly, 1 male, 2 females, 9-
1 1 .VI. 1936, M. Chapman. New York: McLean Bogs,
Tompkins Co., 1 female, 30. VI. 1939, J. G.
Franclemont. New Jersey: Tabernacle, 1 male,
11.VI.1939, H. K. Townes; Millwood, 1 female,
21.VI.1936, H. K. Townes. Maryland: Takoma Park,
1 male, 2.VII.1944, H. & M. Townes. Pennsylvania:
Trout Run, 1 female, 29.V.1938, H. K. Townes;
Spring Brook, 1 female, 27.VI.1945, H. K. Townes;
Michigan: Huron Mrs., 2 males, 2 females, VI. 1961,
H. & M. Townes; Iron River, 1 female, 23. VI. 1969,
H. & M. Townes. South Carolina: Cleveland, 3
females, V-VI.1971, G. Townes. Alabama: Gulf
Shores, 1 male, 22.IV.1968, H. & M. Townes. All
these specimens are in the AEI collection at
Gainesville, and those collected prior to 1955 are
paratypes (see Mason, 1955: 67). Additional
paratypes were described by Mason from Maine,
Vermont, New Hampshire, Massachusetts, Con-
necticut, and Quebec.
Distribution. — U.S. A., Canada.
6 Orthomiscus eridolius Kasparyan
Figs. 9, 14, 24-25
Orthomiscus eridolius Kasparyan, 1990. Fauna of USSR Insecta,
Hymenoptera, 3(2):44. M, F. key, des., fig. Type: Female,
Russia: Far East: Khabozavodsk Region (ST.
PETERSBURG). Paratypes examined.
This species has been described in detail by
Kasparyan (1990). Structurally it resembles O. sim-
plex (Mason) but shows differences in the curva-
ture of the basal vein, shape of the areola, shape of
tergite 2, and in the egg structure. The diagnostic
features are given below.
Diagnosis. — Temple more buccate than in O.
simplex. Mandibular teeth equal. Epomia present,
moderately strong. Areola slightly widened at its
junction with costulae and confluent with basal
area. Apical transverse carina of propodeum
angulate at its junction with lateral and median
longitudinal carinae. Petiolar area appearing pen-
tagonal. Basal vein of fore wing only slightly curved
near its junction with the medius vein. Nervellus
intercepted at its lower 0.15-0.2. Tergite 2 without
oblique grooves. Ovipositor short and thick (Fig.
25). Ovipositor sheaths short, tapered. Egg (Figs.
9, 14, 25) oval, with a ventral flattened anchor that
encloses almost half of the egg.
Coloration essentially similar to that of O.
simplex. Head and thorax in female yellow in lower
half and black in upper half. Mesosternum of
female black or brownish-black. Face of male partly
yellow. Abdomen beyond tergite 1 brownish black,
with yellow spots. Mesosternum black in female.
Face of male largely black.
Length. — 4.5-6.0 mm.; fore wing 3.8-4.8 mm.;
ovipositor about 0.4 mm.
Distribution. — Russia: Far Eastern Region.
Orthomiscus amurensis Kasparyan, 1986 is hereby
transferred to Kristotomus.
Volume 3, 1994
165
Kristotomus amurensis (Kasparyan),
new combination
Figs. 13, 32-35
Orthomiscus amurensis Kasparyan, 1986: 55. Kasparyan, 1990.
Fauna of USSR Insecta, Hymenoptera, 3 (2): 40. key.
des., fig. Type: Female, Russia: Far East: Khabozavodsk
Region (ST. PETERSBURG). Examined.
The holotype of this species was examined in
1992. The ovipositor and its sheaths are tapered
and slender and both extend beyond the apex of
abdomen. The lower valve of the ovipositor is
without teeth. The egg has a simple stalk and
anchor. The interocellar distance is about 0.8x the
ocellocular distance. The malar space is 0.5x the
basal width of mandible. The temple is moder-
ately swollen, and is about as wide as the eye
width. The dorsolateral carina of tergite 1 is inter-
rupted just above the spiracle.
The above mentioned characters and the gen-
eral body form would place this species in
Kristotomus, to which genus this species is hereby
transferred. It appears related to Kristotomus
bitccatus Kasparyan.
ACKNOWLEDGMENTS
My thanks are due to Dr. D. R. Kasparyan (St. Peters-
burg) for the loan of the type specimens and for his com-
ments. The collections of the American Entomological Insti-
tute (AEI) were readily available for my studies, for which I
am thankful to the President of the Institute. The draft of the
paper was kindly read by and commented upon by Drs. John
Barron and Charles Porter and their comments have been
incorporated. I am also thankful to Mr. Andrei Sourakov for
translating Kasparyan's key to the Russian species of the
genus for my use. Subsequently I received a similar transla-
tion from Dr. Kasparyan. The use of published figures from
the works of Henry Townes, D. R. Kasparyan and W. R. M.
Mason is duly acknowledged and the source is cited under
figure legends.
LITERATURE CITED
Carlson, R. W. 1979. Family Ichneumonidae. In: Krombein,
Smith & others, Catalog of Hymenoptera in America North
of Mexico 1: 315-741 (Ichneumonidae pages). Washing-
ton, DC.
Gupta, Virendra K. 1990. The taxonomy of the Kristotomus-
Complex of genera and a revision of Kristotomus (Hy-
menoptera: Ichneumonidae: Tryphoninae). Contributions
of the American Entomological Institute 25 (6): 1-88.
Hellen, W. 1951. Neue Ichneumoniden aus
Ostfennoskandien. I. Notulae Entomologicae 31: 25-31.
Holmgren, A. E. 1858. Forsok till upstallning och beskrifning
av de i Sverige fauna Tryphonider [Monographia
Tryphonidum Sueciae]. Svenska Vetensk.-Akad.
Handlirsch (N.F.) 1: 93-246, 305-394.
Jacob, J. C. and J. Tosquinet. 1890. Catalogue des
Ichneumonides de la Belgique appartenant au groupe
des Tryphonides. Annales de la Societe Entomologique
Beige 34: 44-135.
Jussila, R. 1975. Ichneumonological (Hym.) reports from
Finland III. Annales Entomologici Fennici 41: 49-55.
Kasparyan, D. R. 1976. [New species of ichneumonids of the
tribe Cteniscini (Hymenoptera, Ichneumonidae) from
eastern Asia. The genera Cycasis Townes, Orthomiscus
Mason and Kristotomus Mason.] Entomologicheskoe
Obozrenie 55(1): 137-150. [English translation in Entomo-
logical Review 55 (1): 99-108.]
Kasparyan, D. R. 1977. [Ichneumonids of the subfamilies
Pimphnae and Tryphoninae (Hymenoptera, Ichneu-
monidae) new for Mongolia and Transbaikalia.]
Nasekomye Mongolica [Insects of Mongolia] 5: 456-470.
Kasparyan, D. R. 1986. [Two new species of the tribe Exentenni
(Hymenoptera, Ichneumonidae) of the Far East] In:
Ler, P. A. (Editor) [Systematics and ecology of insects from
the Far East.] Academiya Nauk SSSR, Vladivostock, 155
pages. (Pp. 54-57).
Kasparyan, D. R. 1990. Fauna of USSR Insecta, Hymenoptera
Vol. Ill, No. 2. Ichneumonidae, Subfamily Tryphoninae:
Tribe Exentenni, Subfamily Adelognathinae. Leningrad:
Nauka Publishing House. 340 pages.
Kerrich, G. J. 1942. Second review of literature concerning
British Ichneumonidae (Hym.), with notes on palaearctic
species. Transactions of the Society for British Entomology
8 (2): 43-77.
Kerrich, G. J. 1952. A review, and a revision in greater part, of
the Cteniscini of the Old World. Bulletin of the British
Museum (Natural History) Entomology Series 2 (6): 305-
460.
Mason, W. R. M. 1955. A revision of the Nearctic Cteniscini
(Hymenoptera: Ichneumonidae) I. Eudiaborus Kerrich
and a new genus. Canadian Journal of Zoology 18-73.
Mason, W. R. M. 1962. Some new Asiatic species of Exenterini
(Hymenoptera: Ichneumonidae) with remarks on ge-
neric limits. Canadian Entomologist 94: 1273-1296.
Mason, W.R.M. 1966. A primitive new species of Kristotomus
Mason (Hymenoptera: Ichneumonidae). Canadian Ento-
mologist 98: 46-49.
Roman, A. 1914. Beitrage zur schwedischen Ichneu-
monidenfauna. Arkivfor Zoologie 9 (2): 1-40.
Townes, H. 1969. The genera of the Ichneumonidae Part I.
Memoirs of the American Entomological Institute 11: 1-300.
Townes, H., S. Momoi and M. Townes. 1965. A Catalog and
reclassification of the Eastern Palearctic Ichneumonidae.
Memoirs of the American Entomological Institute 5:1-671.
166
Journal of Hymenoptera Research
Figs. 1-14. 1-5. Orthomiscus platyura Mason: 1, habitus, 2, head; 3, propodeum and tergite 1; 4, lateral view of egg; 5, ventral
view of egg. Figs. 6-9. Lateral view of egg of: 6, O unicinctus (Holmgren); 7, O medusae Kasparyan; 8, O pectoralis Hellen; 9,
O. eridolius Kasparyan. Figs. 10-14. Apex of abdomen showing subgemtal plate, ovipositor and ovipositor sheath of: 10, O.
medusae; 11, O. unicinctus; 12, 0. pectoralis; 13, 0. [=Kristotomus] amurensis (Kasparyan); 14, O. eridolius. [Figs. 1-3 after Townes,
1969; 4-6 after Mason, 1955; 7-14 after Kasparyan, 1990.]
Volume 3, 1 994
167
17
Figs. 15-18. O. platyura: 15, face and mandible; 16, tip of abdomen showing ovipositor, subgenital plate and ovipositor sheath;
17, tergite 1; 18, enlarged view of ovipositor + egg.
168
Journal of Hymenoptera Research
20
t>
(■P
21
Figs. 19-21. Orthomiscus unianctus: 19, tergite 1; 20, apical half of abdomen; 21, ovipositor with egg, ovipositor sheath and
subgenital plate. [Photos taken from specimens of O leptura Mason.]
Volume 3. 1 994
169
25
Figs. 22-25. 22, vertex of Orthomiscus unicinctus; 23, propodeum of O. unicinctus; 24, vertex of O. eridolius; 25, ovipositor + egg
of O. eridolius. [22-23 of O. leptura Mason; 24-25 from holotype of eridolius.]
170
Journal of Hymenoptera Research
28
Figs. 26:31. 26-30. Orthomiscus simplex (Mason): 26, vertex; 27, face and mandible; 28, tergite 1; 29, tip of abdomen showing
ovipositor and subgemtal plate; 30, egg. 31, Face and mandible of O medusae. [Figs. 27, 29, 30 after Mason, 1966; 26 and 28
of paratype and 30 after Kasparyan, 1990.]
Volume 3. 1994
171
32
35
Figs. 32-35. Kristotomus amurensis (Kasparyan): 32, habitus; 33, face; 34, ovipositor and sheaths; 35, abdomen. [Photos of
holotype, Orthonuscus amurensis Kasparyan]
172
Journal of Hymenoptera Research
Figs. 36-38. Fore wing showing the curvature of the basal vein in: 36, Orthomiscus unicinctus; 37, O. platyura; 38, O. medusae.
Volume 3. 1 994
173
Figs. 39-42. Maxillary palp: 39, Orthomiscus unicinctus; 40, O. platyura; 41, O. medusae; 41, O. simplex.
J. HYM. RES.
Vol. 3, 1994, pp. 175-206
Revision of West-European Genera of the Tribe Aylacini Ashmead
(Hymenoptera, Cynipidae)
J. L. Nieves-Aldrey
Museo Nacional de Ciencias Naturales (Biodiversidad), c/Jose Gutierrez Abascal 2, 28006 Madrid, Spain
Abstract. — The west European genera of Aylacini Ashmead (Hym., Cynipidae) and the genus Antistrophus Walsh from North
America are reviewed. Diagnoses of all the genera are presented and an illustrated key for the identification of the genera is
provided. Four genera are described as new: Barbotuua gen. n., Hedickwna gen. n., Iraelk gen. n. and Neaylax gen. n. The genus
Timaspis Mayr is resurrected and the following synonymies are proposed: Aylacopsis Hedicke 1923 = Timaspis Mayr 1881;
V</eldiella Ionescu and Roman 1962 = Cecconta Kieffer 1902 and Endocauloma Ionescu and Roman 1960 = Panteliella Kieffer 1902.
Five species are redescnbed and figured and lectotypes for six species are designated. Nine new combinations are proposed
as well as new synonymies for three species. A check list of the west European species of Aylacini is presented.
INTRODUCTION
The Cynipidae or gall wasps is one of the
better known families of Cynipoidea. It is esti-
mated that there are around 1,800 species distrib-
uted mainly in temperate areas of the northern
hemisphere (Dalla Torre and Kieffer 1910,
Nordlander 1984). More than 250 species have
been recorded in Europe (Nieves-Aldrey 1987).
Galls of this group of parasitic wasps are counted
among the most specialized and morphologically
complex of all zoocecidia. About 75% of all known
cynipids are gall inducers associated with
Fagaceae, almost exclusively with Querais; an-
other large group is associated with Rosaceae and
Asteraceae, while a few species are linked to
Lamiaceae, Papaveraceae, Valerianaceae,
Aceraceae, Fabaceae, and Apiaceae.
With regard to the higher classification of the
Cynipidae, the family is divided here into six
tribes, on the basis of the five tribes proposed by
Ashmead (1903); see also Kinsey (1920). All the
tribes but one, viz Eschatocerini, are present in
west-Europe. Eschatocerini is only represented by
the Neotropical Eschatocerus Mayr, gall maker on
species of Acacia and Prosopis. This genus presents
an axilla large, triangular and situated in the same
plane as scutellum; scutellar foveae and notauli
absent and reduced wing venation; all of them
very distinctive apomorphic morphological fea-
tures, supporting its monophyly. Pediaspidini in-
cludes only one genus of palaearctic distribution:
Pediaspis, Tischbein gall maker on Acer (Aceraceae).
Some authors (Weld 1952, Weidner 1968, Askew
1984) include this genus within the Aylacini but
since it exhibits a specialized scutellar morphol-
ogy and has a heterogenic life-cycle, it is better
placed in a separate tribe. Rhoditini is composed
of two genera: Diplolepis Fourcroy and Liebelia
Kieffer (Ronquist, "in press") which induce galls
on Rosa (Rosaceae). Cynipini comprises of oak gall
inducers and they are associated exclusively with
Fagaceae, mainly Qitercus. The inquiline group is
composed of genera which do not produce galls,
living as inquilines in the galls of other Cynipidae.
Some authors, especially in North America, have
given subfamily status to this group (Ashmead
1903, Burks 1979); more frequently it has been
included together with the gall inducers in a large
group without subdivisions (Weld 1952, Eady and
Quinlan 1963). Here it is regarded as a tribe.
Recent studies on phylogenetic relationships
within Cynipidae (Ritchie 1984, Ronquist in press)
support the monophyly of Synergini as well as
that of Cynipini and Rhoditini. On the other hand,
Aylacini appears to be based on symplesiomorphy
and would be an unnatural paraphyletic group.
Nevertheless, for historical reasons and consider-
ing biological characters, I prefer to treat them
here as a tribe .
176 Journal of Hymenoptera Research
KEY TO THE TRIBES OF CYNIPIDAE
1 Pronotum dorsally longer, in median dorsal line at least one-sixth, usually one-third as long as greatest
length on outer lateral margin, and frequently with truncation and pits forming pronotal plate (Figs. 1,
2); projection of ventral spine of gaster of female always short. Species usually fully alate, except male
of Plwnacis centaureae and some forms of Synergus ttmumacerus, which are brachypterous 2
— Pronotum dorsally very short, in medial dorsal line one-seventh or less as long as outer lateral margin, and
without truncation or pits (Fig. 3); projection of ventral spine of gaster of female usually longer than
broad, viewed laterally. Agamic forms of some species apterous 4
2 Scutellum dorsally flat and trapezoidal with a large, heart-shaped impression on the disk; scutellar foveae
absent (Fig. 4). One species with heterogonic life cycle, associated with Aceraceae (Acer spp)
Pediaspidini
— Scutellum oval or rounded, always dorsally convex; scutellar foveae usually present, sometimes reduced
to an arched depression. Inquilines or gall makers associated with plants other than Acer 3
3 Gula (Fig. 5) reduced to a long, narrow median strip; gular sulci united well before reaching hypostomata;
metasomal tergites 2+3 (third and fourth abdominal terga )(Fig. 8) fused, at least in the female, into one
large segment occupying nearly the whole of the gaster in the female; if showing only a false suture in
either sex (Fig. 6), then the face usually has two vertical and parallel carinae (Fig. 7); face and thorax
sculptured, sometimes very lightly, but never smooth and shining; Maxillary palps with 5 palpomeres
with 1 and 2 fused. Inquilines in galls of Quercus and Rosa Synergini
— Gular sulci (Fig. 9) usually free or meeting near the hypostomata; metasomal tergites 2-7 (Fig. 10) free in
most cases; if metasomal tergites 2+3 fused in the female into one large segment then head and thorax
almost smooth and shining; pits of pronotum usually present and sometimes associated with a light
pronotal plate; maxillary palps 5-segmented. Gall makers on plants other than Quercus, Rosa, and Acer
Aylacini
4 Axilla large, triangular and situated in the same plane as scutellum; scutellar foveae (Fig. 1 1 ) absent; notauli
absent; wing venation (Fig. 12) reduced; Rs+M and Rl not visible; radial cell of forewing with radius
quite separate from anterior margin of wing; frons (Fig. 13) with a strong medial keel. One genus from
South America, gall maker on Acacia Eschatocerini
— Axilla much smaller; not situated at the same level as the scutellum; scutellar foveae present or forming a
shallow arcuate transverse depression; (Rs+M) of fore wing present and Rs reaching or almost reaching
to anterior margin of wing; frons without medial keel or with a much weaker keel 5
5 Mesopleuron (Fig. 14) with longitudinal furrow; hypopygium of female (Fig. 15) ploughshare-shaped;
radial cell of wing usually closed anteriorly. Gall makers on Rosa Rhoditini
— Mesopleuron without longitudinal furrow; hypopygium of female never shaped as above; radial cell open
anteriorly. Gall makers on Quercus, Castanea and Lithocarpus Cynipini
The tribe Aylacini comprises a group of primi- (1963), Quinlan (1968) and Kovalev (1982), but as
five genera of cynipid gall-inducers associated far as I know no entire revision of the group has
mainly with herbaceous plants. Most of their rep- been accomplished.
resentatives are linked with species of Asteraceae The group appears to be particularly abun-
producing milky juice, but some genera and spe- dant and species-rich in the far east of Europe
cies are associated with Papaveraceae, Lamiaceae, where many taxa have been described recently
Valerianaceae and Apiaceae. Galls of the species (Zerova et al. 1988) (see also Belizin 1959). Unfor-
of this tribe are not complex ; they are produced tunately I have failed to obtain type material of
mainly in twigs or fruits and are not detachable these taxa to be included in this revision,
from the host plant tissues. The life cycles are uni-
or bivoltine but without heterogony or alternation MATERIAL AND METHODS
of generations (Folliot 1964, Askew 1984). There is
no previous full revision of the genera of this tribe. This study is supported by the examination of
SomepartialrevisionofgeneraincludedinAylaciru the types of almost all the genera represented in
were made by Weld (1952), Eady and Quinlan western Europe. The initial purpose was to in-
Volume 3, 1994
177
elude all the genera described from the western
Palaearctic region but since I failed to borrow the
type material of some genera from eastern Europe,
mainly Russia, Ukraine and Rumania, the work
was finally restricted to west-Europe. It was also
necessary, in order to clarify the taxonomic status
and relationships of some genera, to examine some
Nearctic material. These were: Antistrophus,
Gillettea and the type species oiAulacidea Ashmead.
Type species of genera of Aylacini are cited ac-
cording to Rohwer and Fagan (1917). The list of
collections from which material have been exam-
ined for this study is as follows:
MCNM Museo Nacional de Ciencias Naturales, Madrid,
Spain
MNHP Museum National d'Histoire Naturelle, Pans,
France (Mme Casevitz-Weulersse)
MZLU Museum of Zoology and Entomology Lund Uni-
versity, Lund, Sweden (R. Danielsson)
NHML The Natural History Museum (British Museum),
London, England (N. Fergusson)
NHMW Naturhistorisches Museum, Vienna, Austria (M.
Fischer)
USNM National Museum of Natural History, Smithsonian
Institution, Washington,DC, USA (A. Menke)
ZMHB Zoologisches Museum, Humboldt-Universitat,
Berlin, Germany (F. Koch)
ZMH Zoologisches Museum Hamburg, Hamburg, Ger-
many (R Abraham)
ZSBS Zoologische Sammlungen des Bayerischen Staates,
Munich, Germany (E. Diller)
MORPHOLOGICAL TERMS
The terminology mostly follows that used by
Eady and Quinlan (1963), Ritchie (1984) and
Ronquist and Nordlander (1989). The term "tho-
rax" as used here includes the propodeum and
thus is an equivalent to the "mesosoma" of the
mainly American literature. Measurements and
abbreviations used here are as follows: A3 and A4,
third and fourth antennomeres; POL (post-ocellar
distance) is the distance between the inner mar-
gins of the posterior ocelli; OOL (ocell-ocular dis-
tance) is the distance from the outer edge of a
posterior ocellus to the inner margin of the com-
pound eye; transfacial line is the shortest distance
across the face between the antennal sockets. Ab-
breviations of forewing venation are used as Eady
and Quinlan (1963). Supraclypeal area is the me-
dial area between the clypeus and the antennal
sockets. Length and width of the radial cell of the
forewing are measured following Wiebes-Rijks
(1979).
AYLACINI (Ashmead 1903)
Aulacini Ashmead 1903:147
Aylaxini (Ashmead): Quinlan 1968:275
Aylacinae (Ashmead): Kovalev 1982:85
Aulacideini (=Aylaxini) Fergusson 1988:143
Type genus: Aylax Hartig 1840 (=/WflxHartig, 1843 (unjusti-
fied emendation)), orig. desig.
Insects of relatively small size; 1-5 mm. Col-
oration usually black, brown or red-brown. Head
viewed dorsally transverse; usually two times
broader than long; in frontal view usually slightly
broader than high or as broad as high; temples not
expanded behind the eyes; ocelli small; face with
radiating striae from clypeus to antennal sockets
weak to moderately developed, medially usually
absent; subocular sulcus absent. Frons and vertex
usually with dull sculpture, coriaceous, alutaceous
or reticulate; rarely smooth and shining; frontal
carina not present; supraclypeal area raised or
slightly protuberant; antenna filiform with 12 to
14 antennomeres in females, 13 to 15 in males.
Pronotum dorsally long; in median dorsal line
one-sixth at least, usually one-third as long as
greatest length on outer lateral margin, frequently
with truncation and submedial pits forming a
light pronotal plate; sides of pronotum usually
pubescent; mesoscutum usually with coriaceous,
alutaceous or reticulate sculpture, less frequently
transversally rugulose or almost smooth; notauli
complete to almost absent; median mesoscutal
impression, if present, usually impressed only in
posterior third or less of mesoscutum; scutellar
foveae usually present, sometimes confluent and
not well defined posteriorly; mesopleuron usually
longitudinally striated, reticulate or rugose-reticu-
late. Wings well developed except the brachypter-
ous males of Phanacis centaureae Forster; radial cell
of forewing open or closed in the margin. Abdomi-
nal tergites 3 to 7 free; third abdominal tergum
covering 1/3 to 1/8 of the whole length of gaster,
laterally with or without a pubescent patch; terg-
ites with or without punctures. Ventral spine of
hypopygium very short. Tarsal claws simple, rarely
with acute basal lobe or tooth.
178 Journal of Hymenoptera Research
KEY TO WEST-PALAEARCTIC GENERA OF AYLACINI
(INCLUDING NEARCTIC GENUS ANTISTROPHUS WALSH)
1 All claws with distinct basal lobe or tooth (Figs. 16, 18). Mesopleuron, mesoscutum and vertex smooth and
shining. Associated exclusively with Rubus and Potentilla (Rosaceae) 2
— All claws simple (Figs. 19, 20). Mesoscutum and vertex rather dull, sculptured, striate or reticulate;
mesopleuron striate or reticulate. Genera not associated with Rosaceae 3
2 Claws weakly lobed (Figs. 17, 18). Subcosta and radius reaching to anterior margin of wing; radial cell
sometimes partly closed anteriorly (Fig. 21). Third and fourth abdominal terga completely fused in
females, separated in males. Gall-maker on Potentilla Xestophanes Foerster
— Claws strongly lobed (Fig. 16). Subcosta and radius not reaching to anterior margin of wing (Fig. 22); radial
cell open anteriorly. Third and fourth abdominal terga completely free in both sexes. One genus
associated with Rubus and Potentilla Diastrophus Hartig
3 Mesopleuron longitudinally striated. Pronotum always with two distinct submedial pits 7
— Mesopleuron reticulate (Figs. 24, 28, 41), rugulose-reticulate (Fig. 116) or rugulose-striate. Pronotum with
or without submedial pits 4
4 Rl and Rs not quite reaching to anterior margin of wing (Fig. 42); wing fringe absent (Fig. 42). Third antennal
segment shorter than fourth (Fig. 37) Antistrophus Walsh
— Rl and Rs reaching to anterior margin of wing and hair fringe distinct, usually long. Third antennal segment
as long as or longer than fourth 5
5 Pronotum (Fig. 23) with two distinct submedial pits. Pronotum, mesoscutum and mesopleuron strongly
reticulate (Fig. 24). Radial cell (Fig. 25) clearly open anteriorly. Radiating striae on face absent (Fig. 26).
Third antennal segment as long as fourth (Fig. 27). Gall-maker on Papaver somniferum L
Iraella gen. n.
— Pronotum usually without submedial pits, rarely with two rather small and indistinct pits. Mesopleuron
reticulate or ruguloso-reticulate. Radial cell usually closed, at least partially. Radiating striae on face
present, sometimes weak or faint. Third antennal segment longer than fourth 6
6 Mesopleuron reticulate (Fig. 28). Notauli usually faint or absent anteriorly. Gall inducer on Asteraceae..
Phanacis Foerster
— Mesopleuron rugulose-reticulate ( Fig. 116) or rugulose-striated. Notauli complete. Gall inducer on Asteraceae
Timaspis Mayr
Head, in frontal view, as high as broad, slightly higher than broad in male (Fig. 29); malar space at least as
long as the height of an eye in the female, longer in male. Scutellar foveae shallow and indistinct. Hair
fringe on apical margin of forewing long (Fig. 30). Gall-maker on Valenanella (Valerianaceae)
Cecconia Kieffer
— Head in frontal view broader than high (Figs. 44, 57); malar space usually shorter than height of an eye.
Scutellar foveae distinct. Wing fringe on apical margin, if present, usually not as long 8
8 Rl and Rs of forewing not quite reaching to anterior margin of wing; radial cell clearly open at base and
along margin (Figs. 62, 79, 110) 9
— Rs almost reaching to anterior margin of wing and Rl continuing more or less along margin of wing
(sometimes Rl only touching the margin); radial cell closed, partially closed or obsoletely closed {Figs.
34,50,55, 103) 12
9 Pronotum shorter medially (Fig. 60). Mesopleuron (Fig. 62) striate with some reticulation between striae.
Third abdominal tergum without lateral pubescent patch. Gall maker on Papaver (Papaveraceae)
Barbotinia gen. n.
— Pronotum longer medially (Figs. 75, 107). Mesopleuron (Fig. 77) finely or strongly striate. Third abdominal
tergum usually with lateral pubescent patch 10
10 Pronotum longitudinally striate and strongly pubescent (Fig. 31). Mesoscutum finely reticulate (Fig. 32).
Third female antennal segment longer than fourth (Fig. 33). Third abdominal tergum with a distinct
lateral pubescent patch (Fig. 31). Hair fringe on margin of forewing moderately long. Associated with
Glechoma (=Nepeta) (Papaveraceae) Liposthenus Forster
— Pronotum without longitudinal striae (Fig. 77), coriaceous or reticulate, only moderately pubescent.
Mesoscutum coriaceous or rugulose-coriaceous. Third female antennal segment shorter than fourth (Fig.
73). Third abdominal tergum with or without lateral pubescent patch. Hair fringe on margin of forewing
absent or very short 11
Volume 3, 1994 179
11 Hair fringe on wing margin completely absent (Fig. 110). Mesoscutum coriaceous, with scattered piliferous
punctures (Fig. 108); notauli distinct only in posterior half of mesoscutum. Gaster laterally smooth,
without punctures (Fig. 109). Gall inducer on Phlomis (Lamiaceae) Rhodus Quinlan
— Hair fringe on wing margin very short or absent. Mesoscutum without punctures (Fig. 76). Notauli almost
complete. Fourth to seventh abdominal tergites finely punctate (Fig. 78). Associated with Centaurea
(Asteraceae) Isocolus Foerster
12 Rl of forewing continuing along margin of wing almost completely closing radial cell; sometimes vein is
very lightly pigmented on the margin and radial cell appears obsoletely closed (Figs. 50, 55, 69).
Pronotum always dorsally quite long 13
— Rl usually not continuing along margin of wing, occasionally slightly so; radial cell appearing open, at least
anteriorly (Figs. 34, 103) 14
13 Mesopleuron clearly striate (Figs. 49, 54). Third abdominal tergum with lateral pubescent patch. Antenna
of female 13-segmented. Notauli usually complete. Hair fringe on margin of forewing moderately long.
Associated with Asteraceae and Lamiaceae Aulacidea Ashmead
— Mesopleuron not clearly striate (Fig. 68), with some striae on alutaceous sculpture. Third abdominal tergum
without lateral pubescent patch (Fig. 70). Antenna of female 12-segmented (Fig. 65). Hair fringe on
margin of forewing absent (Fig. 69). Gall inducer on Salvia (Lamiaceae) Hedickiana gen. n.
14 Notauli weak or obsolete even in posterior half of mesoscutum. Third abdominal tergum without lateral
pubescent patch. Gall maker on Phlomis (Lamiaceae) Panteliella Kieffer
— Notauli clearly impressed, at least in posterior half of mesoscutum. Third abdominal tergum with or
without lateral pubescent patch 15
1 5 Pronotum dorsally relatively short and pits narrowly separated, more strongly transverse (Fig. 1 ). Scutellar
foveae smaller, sculptured. Female antenna 13-segmented. Rl partially closing the radial cell (Fig. 34).
Gall maker on Papaver (Papaveraceae) Aylax Hartig
— Pronotum dorsally longer and pits rounder and more broadly separated (Fig. 100). Scutellar foveae larger,
rounded and smooth (Fig. 101). Female antenna 12-13 segmented. Rl not partially closing the radial cell
(Fig. 103). Associated with Salvia (Lamiaceae) Neaylax gen. n.
ANTISTROPHUS Walsh
Antistrophus Walsh 1869:74. Type species: Antistroplius
ligodesmiae-pisum Walsh. Monotypic (examined)
Asclepiadiphila Ashmead 1897b:263. Type species:
Asclepiadiplula steplianotidis Ashmead. Monotypic.
Diagnosis. — Head, viewed dorsally two times
broader than long; viewed frontally, slightly
broader than high; temples slightly expanded be-
hind the eyes. Frons and vertex strongly reticulate;
radiating striae present; supraclypeal area reticu-
late, slightly protuberant; antenna 13-14 segmented
in female, 14-segmented in male; A3 clearly shorter
than A4. Pronotum strongly reticulate, in dorsal
view, long; submedial pits conspicuous.
Mesoscutum reticulate; notauli slightly faint ante-
riorly; median mesoscutal impression weakly im-
pressed posteriorly; scutellar foveae large,
rounded, confluent; scutellum strongly reticulate;
mesopleuron reticulate. Wings hyaline; Rl not
quite reaching the margin of wing; radial cell
open, nearly three times as long as broad, without
hair fringe on distal margin. Third abdominal
tergum without lateral pubescent patch; follow-
ing tergites not punctate.
Comments. — The structure of the antenna,
pronotum and radial cell relate it to the European
genus Isocolus, but it differs from Isocolus in the
strongly reticulate sculpture of the mesopleuron
and mesoscutum. This feature relates it to Phanacis
but it differs from that genus in important traits
such as the presence of submedial pits in the
pronotum, scutellar foveae distinct, third anten-
nal segment shorter than fourth, radial cell open,
and hair fringe on distal margin of forewing ab-
sent.
Distribution and Biology. — Antistrophus, in spite
of being cited in Weld (1952) as a genus confined
to North America, is mentioned in the generic
keys of the same work as also present in Greece,
galling plants of Salvia (Lamiaceae). Later on,
Quinlan (1968) reiterates, without questioning it,
the record. But the truth is that there is no correct
distribution record of this species for that country,
therefore Quinlan's citation is quite clearly errone-
ous and the occurrence of the genus in Europe
quite unlikely. All data suggest that it is a genus of
exclusively Nearctic distribution.
180
Journal of Hymenoptera Research
The genus comprises nearly 10 species in North
America, associated with species of Silphium and
Chrysotlwmnus (Asteraceae), in the stems of which
they produce galls.
REVISED SPECIES
Antistrophus pisum Ashmead
Antistrophus pisum Ashmead 1885:294
Antistrophus ligodesmm-pisum Walsh 1869:74 (invalid name).
Asclepiadiphila stephanotidis Ashmead 1897b:263
Examined material. — Walsh's type series of A.
pisum in the USNM consists of 34 syntypes: 11 6\
23 9 , part of them mounted on micro-pins and part
on cards. 4 9 , with red labels printed "Type No
1523 U.S.N.M." and white handwritten
"Antistrophus pisum Walsh"; the female in better
condition, only missing the last segments of the
right antenna, is designated lectotype; 7 $ , glued
on a single card, with labels number 208, red
"Type No 1523 USNM"; white "Lygodesmia gall",
are designated paralectotypes. The type series of
A. stephanotidis Ashmead comprises 46" and 79;
1 9 , set on card point, with labels: white "4780 4-0-
91" and red "Type No 3737 USNM", is designated
lectotype.
Redescription. — FEMALE. (Figs. 35-42). Length
3 mm. Yellow-red or amber with darkened parts
mainly in vertex, mesoscutum, scutellum and
upper part of gaster; antenna and legs concolorous
with the body. Head scarcely pubescent and
strongly reticulate, viewed dorsally (Fig. 35) two
times broader than long, viewed frontally (Fig. 36)
1.1 times broader than high. POL slightly shorter
than OOL and about three times greatest diameter
of lateral ocellus; transfacial line around 2.4 times
height of eye; temples moderately expanded be-
hind eyes; malar space 1.2 times height of eye; face
laterally with radiating striae; supraclypeal area
reticulate, only slightly prominent; antenna (Fig.
37) relatively long, slightly shorter than body,
filiform, with 13-14 antennomeres; if only 13, then
top segment is very long; third segment clearly
shorter than fourth, slightly excavated in middle.
Pronotum (Fig. 39) reticulate and pubescent; dor-
sally long, with distinct submedial pits associated
to clearly visible pronotal plate; mesoscutum (Fig.
40) strongly reticulate; notauli faint in anterior
third; moderately convergent posteriorly; median
mesoscutal impression indistinct, weakly im-
pressed in posterior fourth of mesoscutum; scutel-
lar foveae large, confluent, rounded; scutellum
rounded, strongly reticulated; mesopleuron (Fig.
41) reticulate; propodeum reticulate and laterally
pubescent; propodeal carinae slightly divergent.
Wings hyaline with pale venation; Rl and Rs of
forewing not quite extending to wing margin,
radial cell open (Fig. 42); Rs rather curved ; length
of the radial cell about 2.8 times the width; areolet
indistinct; cubitalis (Rs + M) joining the basalis
(Rs+M) at a point close to junction of basalis with
Cu^ fore-wing without distinct hair fringe on wing
margins. Gaster, viewed laterally, slightly longer
than head + thorax; third abdominal tergum (Fig.
41) basally smooth without sculpture or pubes-
cence, following segments not punctate nor pu-
bescent; ventral spine of hypopygium very short.
MALE, differs from female in darker coloration,
mainly rufo-brown; antenna longer (Fig. 38), with
14 antennomeres.
AULACIDEA Ashmead
A ulacidea Ashmead 1 897a :68. Type species: Aulax mulgedncola
Ashmead (= Aulax hamngtoni Ashmead 1877) Original
designation
Diagnosis. — Head, viewed dorsally, about two
times broader than long; viewed frontally slightly
broader than high; temples not expanded or slightly
expanded behind eyes; Frons and vertex coria-
ceous; radiating striae present laterally on face;
supraclypeal area raised, coriaceous; antenna 13-
segmented in female, 14-segmented in male, A3
slightly shorter than A4. Pronotum long dorsally
and submedial pits present, broadly separated;
mesoscutum coriaceous, granulate or finely rugu-
lose; notauli usually complete; median mesoscutal
impression usually present posteriorly; scutellar
foveae distinct, usually large and rounded;
mesopleuron longitudinally striated. Radial cell
of forewing entirely closed on margin; hair fringe
present, moderately long. Third abdominal ter-
gum, in females, with lateral pubescent patch;
following tergites usually punctate.
Comments. — Morphologically, the genus is
well distinguished by its closed radial cell and
striate mesopleuron. The type species was de-
scribed from North America and is redescribed
below.
Distribution and Biology. — Genus of holarctic
distribution. The genus is represented in West-
Volume 3, 1 994
181
Europe by 12 species. Most of European species
induce galls on stems and leafs of species of
Asteraceae specially on Hieracium; one species
produce galls in achenes of Nepeta (Lamiaceae).
Aulacidea harringtoni (Ashmead)
Aulax harringtoni Ashmead 1887:146
Aulax mulgediicola Ashmead 1896:133 (examined)
Type material. — Series composed of 75 speci-
mens, individually set on card points. 1 9 , with
labels: red printed "Type No 3092 USNM"; white
handwritten "Aulax mulgediicola" , is designated
lectotype.
Redescription. — FEMALE. Figs. 43-50. Length:
2.4 mm. Reddish yellow or orange, darker on
dorsum; antenna and legs entirely yellow. Head,
in dorsal view (Fig. 43), a little less than two times
broader than long; in frontal view (Fig. 44) 1.2
times broader than high; POL: OOL as 10:9; OOL
slightly more than three times greatest diameter of
lateral ocellus; trans facial line about 1 .4 times height
of eye; temples slightly expanded behind eyes;
frons and vertex finely coriaceous; face laterally
with incomplete radiating striae; supraclypeal area
not prominent; antenna (Fig. 45) filiform with 13
antennomeres, third segment shorter than fourth.
Pronotum (Fig. 47) coriaceous and scarcely pubes-
cent; dorsally long, with distinct submedial pits
broadly separated; mesoscutum (Fig. 48) finely
rugulose or granulate; notauli complete; median
meoscutal impression visible in posterior 1/3 of
mesoscutum; scutellar foveae ovate, well defined,
separated by a septum relatively broad posteri-
orly; scutellum weakly rugose; mesopleuron (Fig.
49) finely striated longitudinally. Wings hyaline ;
radial cell of forewing (Fig. 50) closed on the
margin; length of the radial cell about 2.4 times
width; areolet indistinct; hair fringe on wing api-
cal margin distinct. Gaster (Fig. 49), viewed later-
ally, as long as head + thorax; third abdominal
tergum basally with lateral pubescent patch, fol-
lowing segments finely punctate ; ventral spine of
hypopygium very short. MALE. Differs from fe-
male in antenna (Fig. 46), with 14 antennomeres.
Comments. — The morphological features of the
type species are congruent with those exhibited by
the species represented in Europe.
Distribution and Biology. — Nearctic. The spe-
cies makes galls in stems of Lactuca spp, mainly L.
villosa (Burks, 1979).
SPECIES FROM WESTERN EUROPE
The species of this genus recorded in Western
Europe are listed in the check list (see Appendix).
The novelties are represented by some species
described by Thomson which are considered ei-
ther good species or represent new synonymies
(Nieves-Aldrey, in press). The following are other
new combinations or redescriptions of poorly
known species:
Aulacidea kerneri (Wachtl) comb. n.
Aylax kerneri Wachtl 1891:277
Examined material. — I have been not able to
examine the type material of this species. Observa-
tions come from specimens reared from galls on
Nepeta spp collected in Spain.
The following characters advise the transfer of
this species to the genus Aulacidea Ashmead: prono-
tum relatively long dorsally (Fig. 52); 13-segmented
antenna, with A3 slightly shorter than A4 (Fig. 51);
mesoscutum with complete notauli (Fig. 53); lon-
gitudinally striated mesopleuron (Fig. 54) ; closed
radial cell, although Rl is slightly pigmented on
margin (Fig. 55).
Aulacidea scorzonerae (Giraud)
Aulax scorzonerae Giraud 1859:370
Aulacidea scorzonerae (Giraud): Kieffer 1902:96
Examined material. — The type material, sentby
the MNHP, comprises 66 specimens set on micro-
pins isolated or in pairs. Most specimens are com-
plete and in good condition. All the specimens
carry a white label "Museum Paris, coll. Giraud"
and some carry labels handwritten by Giraud. 1_,
complete, set next to a male, is hereby designated
lectotype.
Redescription. — FEMALE. Figs. 88-95. Length:
2 mm. Coloration variable, from almost entirely
red-brown with darker thorax, antenna orange or
dark-yellow and legs entirely red-yellow except
coxae and base of femora darkened to dark-brown
or black with legs darker too. Head scarcely pu-
bescent and finely coriaceous; viewed dorsally
(Fig. 88) two times broader than long; viewed f ron-
tally (Fig. 89) 1 .2 timesbroader than high. POL:OOL
as 10:7; OOL about 3.5 times the greatest diameter
of lateral ocellus: transfacial line around 1.8 times
182
Journal of Hymenoptera Research
height of eye; temples not expanded behind eyes;
face laterally with incomplete radiating striae;
supraclypeal area only slightly prominent; An-
tenna (Fig. 90) filiform, relatively short, about 0.6
times length of the body; with 13 antennomeres,
A3 clearly longer than A4. Thorax scarcely pubes-
cent; pronorum (Fig. 91) dorsally long; submedial
pits clearly visible, slightly transverse; pronotum
with coriaceous sculpture; mesoscutum (Fig. 92)
finely coriaceous-alutaceous sculptured; notauli
faint anteriorly, posteriorly not broad; median
mesoscutal impression weakly impressed in pos-
terior 1/6 of mesoscutum; scutellar foveae shal-
low, shining and almost smooth, ovate, separated
by septum broader posteriorly; scutellum rounded
with coriaceous-alutaceous sculpture; mesopleu-
ron (Fig. 93) with fine longitudinal striae;
propodeum with subparallel lateral carinae; me-
dian area scarcely pubescent, almost smooth.
Forewings slightly longer than body, hyaline; ra-
dial cell (Fig. 94) closed on front margin, about 2.3
times longer than broad; hair fringe relatively
short; areolet inconspicuous. Gaster as long as
head+thorax; third abdominal tergum with a slight
lateral pubescent patch (Fig. 93); following terg-
ites without visible punctures; ventral spine of
hypopygium quite short. Tarsal claws simple.
MALE. Differs from female in 14- segmented
antenna (Fig. 95), A3 slightly broadened apically.
Comments. — The species had been very sel-
dom recorded and, consequently, was poorly
known having been incompletely described. After
examination, I confirm that it belongs to Aulacidea.
It is a species which is closely related to A.
tragopogonis Thomson, from which it differs in
host plant as well as in its incomplete notauli, less
pronounced scutellar foveae, relatively shorter
radial cell and invisible gastral punctuation.
Distribution and Biology. — Only recorded from
Austria and Hungary. Reared from galls in the
stems of Scorzonera austriaca Willd and S. humilis L.
AYLAX Hartig
Ay/fl.v Hartig 1840(1839):186, 195. Typespecies: Cynips rhoeadis
Bouche. Desig. by Ashmead 1903 (not seen)
Aulax Hartig 1843:412. Emendation.
Diagnosis. — Head, viewed dorsally, about two
times as broad as long; viewed frontally slightly
broader than high; temples not or slightly ex-
panded behind eyes; radiating striae present later-
ally on face; supraclypeal area slightly prominent;
antenna of female with 13 antennomeres; A3 as
long as A4; antenna of male with 14 segments.
Pronorum relatively short dorsally and submedial
pits strongly transverse and slightly separated;
mesoscutum coriaceous-granulate; notauli com-
plete or faint anteriorly; median mesoscutal im-
pression present in posterior 1/2 or more of
mesoscutum; scutellar foveae small, slightly trans-
verse and confluent; mesopleuron longitudinally
striated. Radial cell of forewing only partially or
obsoletely closed on the margin; Rl reaching to
anterior margin of wing; hair fringe present, mod-
erately long. Third abdominal tergum of females
with lateral pubescent patch; following tergites
without punctures.
Comments. — The genus was originally very
heterogeneous. Later on, some species were set
apart and transferred to such new genera as
Aulacidea Ashmead and Isocolus Forster. My present
conception of the genus is very restricted and I
limit it to the species which are gall makers on seed
capsules of Papaver and maybe also of Hypecoum.
Distribution and Biology. — The genus is repre-
sented in Europe by two species: A. papaveris (Perris
1839) and A. minor Hartig 1840. Aylax hipecoi Trot-
ter 1912, described from Northern Africa (Tripoli),
with galls in Hypecoum fruits, also found in Greece
and Algeria, could also be assigned to this genus.
However, the species has not been recorded again
and I am ignorant as to the location of the types, so
I have not been able to study the species.
BARBOTINIA gen. n.
Figs. 56-62
Type species. — Aylax oramensisBarbotm 1964:152. Herein des-
ignated. Derivation: Feminine gender. Dedicated to the
French cynipidologist Mr. Barbotin, discoverer of the
type species.
Description. — Head transverse; viewed dor-
sally (Fig. 56) more than two times as broad as
long; temples not expanded behind eyes; viewed
frontally (Fig. 57) trapezoid-shaped; transfacial
line almost two times height of eye; malar space
longer than height of eye; radiating striae on face
strong, complete, laterally reaching the antennal
sockets, medially absent; face without subocular
sulcus; mandibles large; frons and vertex minutely
alutaceous; antenna filiform, females with 14
antennomeres (Fig. 58), male with 15 antennomeres
Volume 3, 1994
183
(Fig. 59); pedicel as long as broad; A3 as long as A4.
Pronotum (Fig. 60) dorsally relatively short; in
median dorsal line, not more than 1/4 as long as
greatest length on outer lateral margin; submedial
pits clearly visible slightly transverse and moder-
ately separated; mesoscutum (Fig. 61) granulate-
puncruate but punctures very obsolete or indis-
tinct; notauli complete; median mesoscutal im-
pression impressed in about the posterior half of
mesoscutum; scutellar foveae large, rounded, con-
tiguous, shining and almost smooth; scutellum
granulate-rugulose; mesopleuron (Fig. 62) irregu-
larly longitudinally striated but some reticulation
present between the striae. Wings hyaline; Rt not
quite reaching the margin and radial cell of forew-
ing open (Fig. 62); distal wing margins with short
hair fringe. Gaster as long as head+thorax; third
abdominal tergum covering about 1/3 of the gaster,
basally smooth without punctures or pubescence;
ventral spine of hypopygium short. Legs with
simple tarsal claws.
Comments. — The proposed genus is closely
related to Aylax Htg. these two genera having in
common a pronotum which is relatively short
medially and submedial pits which are transverse
and scarcely separated. However, Barbotinia dif-
fers from Aylax in that the head has a trapezoidal
shape with a relatively long malar space in rela-
tion to the height of an eye; the conformation of the
scutellar foveae, very large and rounded; the stri-
ate-reticulated sculpture of the mesopleuron; the
radial cell of the forewing completely open on the
margin, with the Rl vein hardly reaching the wing
margin and the absence of a lateral pubescent
patch, in the third abdominal tergum.
Distribution and Biology. — The new genus com-
prises a single known species: Barbotinia oraniensis
(Barbotin 1964) comb, n., gall maker on capsules of
Papaver species (Papaveraceae). Its known distri-
bution area includes the mediterranean region
and Northern Africa (Barbotin 1964). In Spain it is
relatively frequent (Nieves-Aldrey 1984). In the
NHML there are six specimens and galls of this
species collected in Italy, Moladi Baxi, 1974, iden-
tified by Quinlan as Aylax sp.
CECCONIA Kieffer
Cecconia Kieffer 1902:7, 93. Type species: Aulax valerianellae
Thomson 1877. Monobasic and original designation
(examined)
We/iiid/n Ionescu and Roman 1962:551. Type species: Weldiella
aequalis Ionescu and Roman 1962. Monobasic and origi-
nal designation (not seen)
Diagnosis. — Head slightly less than two times
broader than long viewed dorsally; in frontal view
as broad as high or slightly higher than broad;
malar space as long as height of eye; clypeus
subquadrate; radiating striae well impressed;
supraclypeal area slightly prominent; female an-
tenna with 13 segments; A3 shorter than A4 or as
long as A4; male antenna with 14 antennomeres.
Pronotum dorsally not very long with distinct
submedial pits; mesoscutum with alutaceous
sculpture; notauli very lightly impressed, absent
in anterior half of mesoscutum; median mesoscutal
impression absent; scutellar foveae confluent,
rounded, not well defined posteriorly; scutellum
dorsally convex, with reticulate sculpture;
mesopleuron finely striated longitudinally. Wings
hyaline; radial cell of forewing open on the mar-
gin; about three times longer than broad; wing
margins with long hair fringe. Third abdominal
tergum with a lateral pubescent patch; following
segments not punctate.
Comments. — The genus is easily distinguished
morphologically: the head shape higher than broad
viewed frontally; presence of long hairs in the
distal wing margin; the shape of the scutellar
foveae and notauli conformation. Weldiella was
erected by Ionescu and Roman (1962) for the spe-
cies W. aequalis found in Rumania, also gall-maker
on fruits of species of Vrt/t'n'one//rt.(Valerianaceae).
On the basis of the original description (since I
have not had the chance to examine the type
species) I consider the differences used to differen-
tiate that genus from Ceconia insufficient to sup-
port the description of a separate genus. At most,
the mentioned features could indicate the exist-
ence of a second species, different from C.
valerianellae, within the genus Cecconia.
Distribution and Biology. — The genus com-
prises two known species which produce galls in
the fruits of species of Valerianella ( Valerianaceae).
C. valerianellae (Thomson) is widely distributed in
Europe from Sweden to Rumania, through the
Iberian Peninsula; C. aequalis (Ionescu & Roman),
is only recorded from Rumania.
184
Journal of Hymenoptera Research
DIASTROPHUS Hartig
Diastrophus Hartig 1840 (1839): 186. Type species: Diastrophus
rubi Hartig. Monotypic
Type material of D rubi Hartig, is housed in Munich (ZSBS)
It is composed of 1 i and 5 9 . Lectotype 9 , mounted on
micro-pin, designated by Weld (1931) (examined).
Diagnosis. — Head slightly more than two times
broader than long viewed dorsally; in frontal view
slightly broader than height; malar space slightly
shorter than height of eye; transfacial line 1.5 to 2
times height of eye; supraclypeal area rugose-
punctate with some striae; radiating striae incom-
plete; female antenna with 13 segments, male an-
tenna with 14 antennomeres; A3 slightly longer
than A4. Pronotum dorsally long with distinct
submedial pits asociated with a distinct pronotal
plate; mesoscutum mostly smooth and shining;
notauli well impressed and complete; median
mesoscutal impression variable as to its length;
scutellar foveae deep, rounded, smooth and shin-
ing; scutellum with a longitudinal furrow;
mesopleuron almost entirely smooth and shining,
medially with some fine longitudinal striae. Wings
weakly infuscate; radial cell of forewing open on
the margin; wing apical margin with hair fringe
moderately long. Third abdominal tergum with-
out lateral pubescent patch; following segments
punctate. Tarsal claws of the legs with conspicu-
ous acute basal lobe or tooth.
Comments. — The genus is quite distinct mor-
phologically and together with Xestophanes Forster
1869 and Gonaspis Ashmead 1897 it forms a sepa-
rate group within Aylacini. In fact, morphologi-
cally, they are distinguished from the rest of the
genera of this tribe because they exhibit claws
having a basal lobe or tooth; in the biological
aspect, both genera are also well characterized by
their association with species of Ritbus and Poten-
tilla (Rosaceae). Their special position within
Aylacini has been pointed out by recent phylog-
eny studies of some genera of cynipids, mainly
inquilines, by Ritchie (1984) and Ronquist (in press).
The latter study indicates that Diastrophus is the
genus of gall-wasps most closely related to the
inquilines, which form a monophyletic group.
Distribution and Biology. — In Europe
Diastrophus is represented by only two species: D.
rubi Hartig, gall-inducer in species of Rubus and D.
mayri Reinhart, 1876, which forms galls in the
stems of Potentilla argentea L. In contrast with
Europe, the genus comprises many species in North
America, most of them also associated with spe-
cies of Rubus, the others associated with Potentilla
and Fragaria.
HEDICKIANA gen. n.
Figs. 63-70
Type species: Aulaadea levantma Hedicke 1928:81. By present
designation Derivation: Feminine gender Dedicated to
the German entomologist Hedicke, discoverer of the
type species.
Examined material. — 4 9 (paratypes) deposited
in the NHML. The holotype stated by the author to
be deposited in his collection, has not been found
and is probably lost. Consequently, we designate
a lectotype amongst the studied paratypes: 1_,
mounted on micro-pin on card point, red label
"paratypus"; handwritten labels " Aulacidea
levantina 1928 Hedicke det" and "Tel Aviv Pales-
tine Bodenheimer".
Description. — FEMALE. Head transverse;
viewed dorsally (Fig. 63) more than two times
broader than long; POL almost equal to OOL;
temples slightly expanded behind eyes; viewed
frontally (Fig. 64) clearly broader than high;
transfacial line slightly more than 1.5 times height
of eye ; malar space shorter than height of eye;
radiating striae on face well impressed laterally;
supraclypeal area slightly protuberant, without
striae; frons and vertex with coriaceous-alutaceous
sculpture; antenna (Fig. 65) 12 segmented, A3
slightly more slender at the base; as long as A4.
Pronotum (Fig. 66) dorsally relatively long
submedial pits clearly visible, broadly separated
mesoscutum (Fig. 67) coriaceous-punctate; notauli
posteriorly broad but impressed only in posterior
3/4 of mesoscutum; median submedial impres-
sion visible in posterior 1 /4 of mesoscutum; scutel-
lar foveae large, rounded, contiguous, shining and
finely coriaceous; scutellum rugulose;
mesopleuron (Fig. 68) longitudinaly striated but
some reticulation present between the striae. Wings
slightly infuscated; radial cell of forewing (Fig. 69)
closed on front margin; distal wing margins with
hair fringe quite short. Gaster as long as
head+thorax; third abdominal tergum covering
about 1 /3 of gaster; basally smooth without punc-
tures or pubescence; following segments punc-
tate; ventral spine of hypopygium short. Tarsal
claws simple.
Volume 3, 1 994
185
Comments. — The genus is erected for the single
species: H. lez>antina (Hedicke), described within
Aulacidea Ashmead. In common with Aulacidea,
the new genus exhibits a closed radial cell but it
can be distinguished from Aulacidea as follows:
antenna with 1 2 segments; mesopleuron not clearly
longitudinally striated, in part irregularly reticu-
late; incomplete notauli and mesoscutum sculp-
tured with piliferous punctures; base of third ab-
dominal tergum lacking lateral pubescent patch.
Distribution and Biology. — Eastern Mediterra-
nean; recorded from Crete, Syria and Israel
(Hedicke 1928). The only known species, Hedickiana
levantina (Hedicke), induces conspicuous juicy galls
in stems of Salvia triloba L. (Lamiaceae).
IRAELLA gen. n.
Figs. 23-27
Type species: Aulax luteipes Thomson 1877:807. (= Timaspis
papavens Kieffer in Goury et Gignon). By present desig-
nation (examined)
Derivation: Feminine gender. Dedicated to my wife Ira
Description. — Head viewed dorsally scarcely
less than two times broader than long; temples not
expanded behind eyes; viewed frontally (Fig. 26)
slightly broader than high; transfacial line slightly
more than height of eye ; radiating striae on face
absent; clypeus trapezoid-shaped, moderately
projecting anteriorly, supraclypeal area slightly
prominent, reticulate; frons and vertex with re-
ticulate-coriaceous sculpture; female antenna, 13-
14-segmented, male antenna 14-15-segmented; A3
almost of equal length to A4 (Fig. 27). Pronotum
(Fig. 23) dorsally relatively short; submedial pits
present; mesoscutum reticulate-coriaceous; notauli
faint in anterior one third of mesoscutum; median
mesoscutal impression visible in posterior 1/3 of
mesoscutum; scutellar foveae transverse,
confluent; scutellum reticulate; mesopleuron (Fig.
24) reticulate. Wings hyaline; radial cell of forew-
ing (Fig. 25) open; three times longer than wide;
distal wing margins with hair fringe long. Third
abdominal tergum basally with lateral pubescent
patch; following segments without punctures.
Comments. — The following morphological fea-
tures define this new genus: face lacking visible
radiating striae; clypeus projecting anteriorly;
pronotum relatively short dorsally but with vis-
ible submedian pits; mesopleuron entirely reticu-
late; radial cell open on front margin. Some char-
acters, like the general body shape, face and
mesopleuron sculpture and shape of the scutellar
foveae, approach the new genus to the Timaspis-
Phanacis complex, but the presence of submedian
pits in the pronotum is a differentiating trait; other
features that distinguish them are: the entirely
reticulate face with total absence of radiating striae;
clypeus projecting anteriorly; third antennal seg-
ment being of almost the same length as fourth
and radial cell of the forewing open on front
margin.
The genus is erected for a single species: Iraella
luteipes (Thomson) comb. n. (= Timaspis papaveris
Kieffer). Redescription of Aylax luteipes Thomson
including statement of the synonymy indicated
above was recently published (Nieves-Aldrey, in
press).
Distribution and Biology. — Genus distributed
from Sweden to Spain and Iran. The only known
species of the genus is associated with Papaver
somniferum L. (Papaveraceae), on the stems of
which it produces inconspicuous galls. I have had
the chance to examine in the Natural History
Museum of London some specimens from Iran
obtained from stems of Papaver pseudorientale and
P. bracteatum Lindley, which might correspond to
a different species within the same genus.
ISOCOLUS Forster
Isocolus Forster 1869:330, 334 ( = Eubothrus Forster). Type spe-
cies: Diastrophus scabiosae Giraud 1859. Monobasic and
original designation (examined)
Diagnosis. — Head, viewed dorsally, about two
times broader than long; temples at most only
slightly expanded behind eyes; viewed frontally
1.3 times broader than high; transfacial line around
1.5 times the height of eye ; malar space shorter
than height of eye; radiating striae visible later-
ally, usually incomplete, sometimes present even
in the supraclypeal area; antenna of female with 1 3
antennomeres, A3 shorter than A4; antenna of
male with 14 segments. Pronotum dorsally long
and submedial pits conspicuous, broadly sepa-
rated; mesoscutum with coriaceous-rugulose
sculpture, sometimes with transverse rugae;
notauli usually complete; median mesoscutal im-
pression usually visible at least in posterior one
third of mesoscutum; scutellar foveae distinct,
well defined; scutellum rugulose; mesopleuron
with conspicuous longitudinal striae; wings hya-
186
Journal of Hymenoptera Research
line; radial cell of forewing open; Rl and Rs not
quite reaching anterior margin of wing; distal
margin of wing with hair fringe very short. Third
abdominal tergum usually without lateral pubes-
cent patch, rarely with pubescence; abdominal
tergites usually punctate, sometimes weakly, from
fourth tergite; ventral spine of hypopygium short.
Tarsal claws simple.
Comments. — The genus is defined mainly on
the basis of wing characters as follows: Rl and Rs
not quite reaching to anterior margin of wing,
radial cell clearly open and wing fringe absent or
very short.
Distribution and Biologi/. — The distribution is
Palaearctic: five species are known in Western
Europe. From the biological point of view the
genus is associated with Asteraceae; the species
inducing galls in stems, achenes or Lnvolucral
bracteae of species of Centaurea and Serrahda
(Asteraceae).
Redescription Of Some Species
Isocolus scabiosae (Giraud)
Diastrophus scabiosae Giraud 1859:368
Diastrophus areolatus Giraud 1859:369. Syn. n.
Isocolus scabiosae (Giraud): Forster 1869:334
Isocolus rogenhoferi Wachtl 1880:542. Syn. n.
Aulax scabiosae (Giraud): Mayr 1882: 10
Aylax scabiosae (Giraud): Dalla Torre and Kieffer 1910:671
Isocolus scabiosae (Giraud): Rohwer and Fagan 1917:369
Isocolus scabiosae (Giraud): Weld 1952:271
Material examined. — The type series of D.
scabiosae Giraud, in the MNHP, comprises 63 speci-
mens: 30<5, 339, mounted on micro-pins, as well
as some galls on achenes of Centaurea scabiosa. 1 9
, set next to a male, with handwritten label "3 mai"
and "Museum Paris, coll. GIRAUD", is hereby
designated lectotype. Paralectotypes, 18 6 and 12 9
Redescription.— FEMALE. (Figs. 71-79). Length
2.8-3.3 mm. Black; gaster red-brown to black; an-
tenna and legs red brown; coxa, base of femora
and first two segments of antenna darker. Head
viewed dorsally (Fig. 71) 2.2 times broader than
long; viewed frontally (Fig. 72) 1.3 times broader
than high; temples only slightly expanded behind
eyes; frons and vertex with sculpture coriaceous-
alutaceous; POL slightly shorter than OOL , OOL
about four times greatest diameter of lateral ocel-
lus; transfacial line around 1.6 times height of eye;
malar space slightly shorter than height of eye;
face laterally with radiating striae; supraclypeal
area slightly raised, with some striae; antenna
(Fig. 73) about 2/3 the length of the body, with 13
antennomeres, A3 slightly shorter than A4. Prono-
tum (Fig. 75) coriaceous, laterally pubescent; dor-
sally long, with distinct submedial pits ;
mesoscutum (Fig. 76) with coriaceous-rugulose
sculpture posteriorly with some conspicuous trans-
verse rugae; notauli complete, well impressed
posteriorly; median mesoscutal impression dis-
tinct in posterior half of mesoscutum; scutellar
foveae large, rounded; shining and almost smooth;
separated by broad posteriorly septum; scutellum
ovate, rugulose with the posterior half more or less
flat and sloping backwards; mesopleuron (Fig. 77)
longitudinally striated. Wings hyaline or very
lightly infuscated; Rl and Rs of forewing not quite
extending to wing margin and radial cell open
(Fig. 79); Rs slightly curved apically; length of the
radial cell about 3 times the width; areolet indis-
tinct; fore wing without distinct hair fringe on
wing margins. Third abdominal tergum (Fig. 78)
basally smooth without sculpture or pubescence;
posterior third of third tergite and following seg-
ments conspicuously punctate; ventral spine of
hypopygium very short; tarsal claws simple.
MALE, differs from female in antenna (Fig. 74)
with 14 antennomeres; A3 slightly curved in the
middle.
Comments. — The complete series from the
Giraud Collection, sent by the MNHP, comprises
63 individuals. In contrast with the specimens
predominantely black described above, other in-
dividuals of a lighter, chestnut-reddish shade and
some completely reddish individuals are included
in the series. The latter also exhibit other morpho-
logical differences such as a stronger mesoscutal
sculpture and less impressed notauli and medial
mesoscutal impression. Also included in the series
are some galls consisting of transformed achenes
from flower heads of Centaurea scabiosa L. and in
compliance with the handwritten labels from
Giraud: "caput centaureae scabiosae" and the date,
at least part of the specimens of the series should
have been obtained from these galls. Giraud origi-
nally described the species Diastrophus scabiosae as
obtained from conspicuous galls in the stems of
Centaurea scabiosa L. (Giraud 1859), but these galls
are not included in the series. The specimens ob-
tained from galls in achenes could belong to Isocolus
rogenhoferi Wachtl. However, the status of the
Volume 3, 1994
187
latter has not been clear, since only differences in
the coloration of the antennal flagellum together
with the different position of the galls have been
pointed out in the literature (Kieffer 1901, Eady
and Quinlan 1963) to separate it from /. scabiosae.
For these reasons, I consider that there is only one
species with a wide morphological variability be-
ing capable of inducing galls on both stems and
flower heads of species of Centaurea, mainly C.
scabiosa L. It is possible that the two types of galls
correspond to two different generations of the
same species. In the same work, Giraud described
another species, Diastrophus areolatus, from 2 fe-
males caught by netting near Vienna. The differ-
ences between the latter and I. scabiosae were stated
to be reddish coloration of the antennal flagellum
and presence of areolet in the fore-wings. In the
examined series, I found 2 specimens which could
correspond to D. areolatus labeled "captur 13 juill,
Turksch" . The presence or absence of the areola, as
well the general coloration, is quite variable in the
whole of the series, so I consider D. areolatus as
conspecific with J. scabiosae and, therefore, I estab-
lish the corresponding new synonymy.
Isocolus serratulae Mayr comb. n.
Aulax serratulae Mayr 1882:9
Aylax serratulae (Mayr): Kieffer 1901:313
Type material. — Series composed of 31 speci-
mens: 126", 19 9, most of them set on micro-pins
(NHMW). 1 9 , with handwritten labels "serratulae
ydetRogenhofer" and "Luxenburg,Rogenh 1881",
hereby designated lectotype. Other specimens
carry handwritten labels "Aid. serratulae det. G.
Mayr" and "Serratula heterophylla"; designated
paralectotypes.
Redescription.— (Figs. 80-87). FEMALE. Length
2.5 mm. Black; gaster red-brown to dark-brown;
antenna and legs light brown; coxa, base of femora
and two first segments of antenna darker. Head
viewed dorsally (Fig. 80) about 2 times broader
than long; viewed frontally (Fig. 81) 1.2 times
broader than high; temples not expanded behind
the eyes; frons and vertex with sculpture minutely
coriaceous-reticulate; POL:OOLas 11:9; OOL about
three times the greatest diameter of lateral ocellus
transfacial line around 1 .5 times the height of eye
malar space around 0.6 times the height of eye,
face laterally with radiating striae; supraclypeal
area slightly raised, without striae; antenna (Fig.
82) with 13 antennomeres, A3 clearly shorter than
A4. Pronotum (Fig. 83) reticulate-coriaceous, lat-
erally pubescent; dorsally long, with distinct
submedial pits ; mesoscutum (Fig. 84) with reticu-
late-coriaceous sculpture; notauli complete, well
impressed posteriorly; median mesoscutal impres-
sion distinct in posterior 1/5 of mesoscutum; scutel-
lar foveae large, subtriangle-shaped; not smooth;
separated by a posteriorly broader septum; scutel-
lum reticulate-coriaceous, with longitudinal fur-
row variably impressed; mesopleuron (Fig. 85)
longitudinally striated. Wings hyaline, slightly
longer than the body; Rl and Rs of forewing not
quite reaching to wing margin and radial cell open
(Fig. 86); length of the radial cell about 2.6 times
the width; areolet indistinct; fore wing without
distinct hair fringe on wing margins. Third ab-
dominal tergum (Fig. 85) basally with a lateral
pubescent patch; posterior third of third tergite
and following segments conspicuously punctate;
ventral spine of hypopygium very short; tarsal
claws simple. MALE. Differs from female in an-
tenna with 14 antennomeres (Fig. 87), A3 not
modified only slightly curved.
Comments. — This species was described by
Mayr from material collected by Rogenhofer on
Serratula heterophylla auct ross non (L.) Desf. The
gall is probably produced on this plant, but it has
not been described. Since the description, the spe-
cies has been recorded once from Ukrania, includ-
ing galls Zerova et al. (1988), so it is only known
from Austria and that country. The features of this
species fit well in the genus Isocolus, except for the
third abdominal tergum with basal pubescent
patch, so I transfer it to this genus.
Isocolus lichtensteini (Mayr) comb. n.
Aulax lichtensteini Mayr 1882:7
Aylax lichtensteini (Mayr): Kieffer 1901:297
Isocolus lavaresi Nieves-Aldrey 1984:243. Syn. n.
Type material. — Type series composed of 2 9 ;
19, mounted on micro-pin, with handwritten la-
bels " Aulax lichtensteini, det. G. Mayr" and "Cen-
taur, gall lich " designated lectotype. Paralectotype,
one female with label "Centaur, salmant. gall Lich".
Comments. — This species was described from
material collected in Montpellier (France), pro-
ducing galls in the stems of Centaurea salmantica L.
188
Journal of Hymenoptera Research
(=Microlonchus salmanticus) (Mayr, 1882). Isocolus
tavaresi Nieves-Aldrey, described from material
reared from galls on Centaurea aspera L. in Spain
(Nieves-Aldrey, 1984), later also found on Centau-
rea melitensis L., (Nieves-Aldrey, unpublished), is
conspecific with the two type specimens studied
and, therefore, I establish the corresponding syn-
onymy.
The type specimens differ from individuals
collected in Spain (Nieves-Aldrey 1984) only in
their lighter chestnut-reddish coloration, also ex-
tended for antenna and legs, including the coxae.
Therefore, the types exhibit only a slight colour
contrast between the two first antennal segments
and the flagellum, and the longitudinal furrow of
scutellum is more pronounced compared to the
Spanish specimens collected.
LIPOSTHENES Forster
Liposthenus (!) Forster 1869:332.
Liposthenes Forster 1869:336. Type species: Aulax glechomae
Hartig. Monotvpic and orig. desig. (examined). The
types of Aulax glechomae Hartig, in the ZSM, consist of 4
9 , mounted on pins. One of them carry label of lectotype
? designated by Weld in 1931 (Weld 1952).
Diagnosis. — Head, viewed dorsally, slightly
less than two times broader than long; in frontal
view slightly broader than high; transfacial line
1.5 times height of eye; malar space shorter than
height of eye; frons and vertex finely reticulate;
radiating striae incomplete, weakly impressed;
antenna of female with 13 segments, A3 slightly
longer than A4; male antenna 14-segmented.
Pronotum dorsally long, laterally strongly pubes-
cent and with some longitudinal striae or rugae;
submedial pits conspicuous, asociated with a dis-
tinct pronotal plate; mesoscurum regularly coria-
ceous and shining; notauli well impressed and
complete; median mesoscutal impression very
short ; scutellar foveae deep, rounded, smooth and
shining; mesopleuron finely and clearly striated
longitudinally . Wings hyaline; radial cell of forew-
ing open on front margin; length of radial cell
slightly less than three times width. Third ab-
dominal tergum with conspicuous lateral pubes-
cent patch; following segments not punctate. Tar-
sal claws simple.
Comments. — The genus is morphologically
similar to Isocolus Forster, but can be separated by
the pronotum being strongly pubescent laterally;
presence of wing fringe on distal margin; third
antennal segment being longer than fourth; third
abdominal tergum having a lateral pubescent patch
and in general sculpture and biology.
Distribution and Biology. — The genus com-
prises a single known species from Western Eu-
rope which causes the formation of galls in the
runners, stems and leaves of Nepeta (=Glechoma)
hederacea L. (Lamiaceae).
NEAYLAX gen. n.
Type species: Aylax salviae (Giraud 1859):369. By present
designation, (examined)
Derivation: from Neos and Aylax (new Aylax)
Description. — Head, viewed dorsally, two
times broader than long; temples not expanded
behind the eyes; radiating striae on face well im-
pressed laterally; supraclypeal area raised, with-
out striae; frons and vertex with coriaceous or
alutaceous sculpture; antenna 12-13- segmented
in females; 14-15-segmented in males; A3 as long
as A4 or shorter than A4. Pronotum dorsally long;
submedial pits clearly visible, not broadly sepa-
rated; mesoscutum coriaceous or alutaceous, some-
times punctate; notauli complete or faint anteri-
orly; scutellar foveae large, rounded; mesopleuron
longitudinaly striated. Wings hyaline; radial cell
of forewing open on front margin but Rj reaching
margin; distal wing margins with hair fringe mod-
erately long. Third abdominal tergum basally
smooth or with a lateral pubescent patch; follow-
ing segments finely punctate but punctures not
conspicuous; ventral spine of hypopygium short.
Legs with tarsal claws simple.
Comments. — Neai/lax is erected to comprise
some species formerly included in Aylax Htg, but
which do not fit well in that genus. The new
proposed genus is closely related to Isocolus Forster,
from which it differs as follows: radial cell of
forewing not completely open along the margin
and at the base; first abscissa of radius reaching the
wing margin and presence of hair fringe on the
distal margin of the wing (absent in Isocolus).
Biologically Neai/lax differs from Isocolus in its
association with Lamiaceae rather than with
Asteraceae. The dorsally long pronotum, the
rounded scutellar foveae, and the association with
Lamiaceae and not with Papaveraceae separate it
from Aylax Htg.
Volume 3, 1 994
189
Distribution and Biology. — The genus com-
prises two species distributed in centre and south
of Europe which produce galls on species of Salvia
(Lamiaceae) and a third species, only recorded
from the Iberian Peninsula whose host plant and
gall are unknown.
REVISED SPECIES
Neaylax salviae (Giraud), comb. n.
Aulax salviae Giraud 1859:369
hocolus salviae (Giraud): Nieves-Aldrey 1988:224. Syn. n.
Type material. — The type series of A. salviae, in
the NHMW, sent to me by M. Fischer, consists of
12 specimens. 1 6 , mounted on a rectangular card
with labels: red "Type" and white handwritten
"Aulax salviae Giraud", is designated lectotype.
1 8 , 1 9 , mounted on micro-pin, with labels "Col-
lect. Mayr" and "Aulax salviae Gir, det. G. Mayr"
clearly do not belong to the type series, although
they are conspecific with A. salviae Giraud. In the
same series 16, 29, collected by Mayr, deter-
mined as Aulax salviae Giraud, are cynipids but do
not correspond to that species. Finally 2 speci-
mens, identified as A. salviaeby Fahringer, belong
to the family Eucoilidae.
Redescription.— MALE. (Figs. 96-103). Length
1 .8 mm. Red-brown; antenna red-brown; legs with
coxae red-brown, tibiae and tarsi paler. Head
viewed dorsally (Fig. 96) about 2 times broader
than long; temples not expanded behind the eyes;
frons and vertex with sculpture coriaceous-
alutaceous; POL as long as OOL; OOL about 4
times the greatest diameter of lateral ocellus; face
laterally with radiating striae; supraclypeal area
almost raised, without striae; antenna (Fig. 98)
with 14 antennomeres, A3 clearly shorter than A4.
Pronotum (Fig. 100) coriaceous; dorsally long,
with distinct submedial pits; mesoscutum (Fig.
101) finely alutaceous and with some obsolete
punctures medially; notauli relatively expanded
posteriorly, complete but slightly faint anteriorly;
median mesoscutal impression indistinct in poste-
rior 1/4 of mesoscutum; scutellar foveae large,
rounded, smooth and shining, separated by a
narrow septum; scutellum weakly rugulose, with
a weak longitudinal furrow; mesopleuron (Fig.
102) longitudinally striated. Wings hyaline; radial
cell of forewing (Fig. 103) open on front margin but
Rj reaching margin; length of radial cell about
three times the width; hair fringe on distal margin
of wing long. FEMALE, differs from male as fol-
lows: antenna with 13 antennomeres (Fig. 98);
third abdominal tergum without lateral pubes-
cent patch; following segments with fine and in-
conspicuous punctures; ventral spine of
hypopygium short.
Distribution and Biology. — The species was
described from specimens collected in Dalmatia,
reared from galls in the fruits of Salvia officinalis L.
Species of apparently Eastern Mediterranean dis-
tribution.
SPECIES TRANSFERRED TO
NEAYLAX GEN. N.
Neaylax verbenacus (Nieves-Aldrey), comb. n.
Isocolus verbe>\acus Nieves-Aldrey 1988:221. Syn. n.
This species is closely related to N. salviae
(Giraud). It was described within Isocolus from
Spain. Its galls are produced in fruits of Salvia
verbenaca L.
Neaylax versicolor (Nieves-Aldrey), comb. n.
Aylax versicolor Nieves-Aldrey 1985:122. Syn. n.
This species was described under Aylax
(Nieves-Aldrey 1985). It is known only from Spain.
Gall unknown.
PANTELIELLA Kieffer
Pantelia Kieffer 1901:248 (non Bolivar 1887, Orthopf.)
Panteliella Kieffer 1901:324. Type species: Aulax fedtschenkoi
(Rubsaamen 1896). Monobasic
Endocaulonia Ionescu and Roman 1960:222. Type species:
Endocaulonia bicolor Ionescu and Roman 1960. By origi-
nal designation.
Weld (1952), stated that the types, consisting
of slide-mounts of two adults in balsam, were in
the Berlin Museum. Later on, Quinlan (1968) stated
that the type material consisted of two specimens
deposited in the Vienna Museum but Dr. Fischer
informed me that the only material of A. fedtschenkoi
housed in that museum consisted in galls. I have
had the chance to examine one of the preparations
of the Berlin Museum, consisting of a head in bad
condition, with one of the two antennae being
190
Journal of Hymenoptera Research
complete, a leg and the gaster. The preparation has
a red type label and is printed "Aulaxfedtschenkoi;
Bijuk Onlar, juni 1893; Blattgallen on Phlomis
tuberosa". The type species was redescribed by
Weld (1930) and by Quinlan (1968).
Endocanlonia was described by Ionescu and
Roman in 1960 based on E. bicolor, a species discov-
ered by these authors in Rumania producing galls
in the stems of Phlomis tuberosa L. Although I have
not been able to examine the types, it seems clear
from the detailed description that this genus is
very similar to Panteliella Kieffer only differing
slightly in the relative lengths of pedicel and third
antennal segment. The descriptions of the two
genera, except for the mentioned feature, are fully
coincident, as is the gall and the host plant, which
makes me consider that Endocaulonia Ionescu and
Roman is synonymous with Panteliella Kieffer.
The type species of Endocaulonia appears to be
synonymous with Panteliella fedtschenkoi, although
in this case I do not venture to formally establish
the synonymy without a previous examination of
the type.
I present the following diagnosis of the genus
based on Quinlan's redescription (1968).
Diagnosis. — Female antenna 14-segmented;
male 15-segmented; A3 and A4 of same length in
the two sexes. Mesoscurum reticulate; notauli and
median mesoscutal impression very faintly indi-
cated; scutellar foveae distinct separated by a broad
and striate septum; mesopleuron longitudinally
striated. Radial cell of forewing open on the mar-
gin; wing fringe on distal margin distinct. Third
abdominal tergum without lateral pubescent patch;
following tergites weakly punctate on lateral mar-
gins. Tarsal claws simple.
The genus is close to Neaylax Nieves-Aldrey
and to Rhodus Quinlan. It can be separated from
the former by the obsolete notauli, scutellar foveae
broadly separated and female antenna 14-seg-
mented. From Rhodus it is distinguished by the
distinct fringe on distal margin of forewing, the
obsolete notauli and A3 not being shorter than A4.
Distribution and Biology. — Only one known
species: Panteliella fedtschenkoi (Rubsaamen, 1896)
distributed in central and eastern Europe and
eastern Mediterranean. It produces galls in the
leaves and stems of Phlomis tuberosa L. (Lamiaceae).
Panteliella bicolor (Ionescu and Roman) is posibly a
second species of the genus, only known from
Rumania.
PHANACIS Forster
Plmnacis Forster 1860:145. Type species: Plianacis centaureae
Forster 1860. Monobasic.
Gillettea Ashmead, 1897:69. Type species: Gillettea taraxaci
Ashmead 1897. Original designation.
Weld (1952) pointed out that Forster's types in
the Berlin Museum were destroyed by dermestids
but he could observe good specimens of P. centau-
rea in Vienna. However, I have abundant Iberian
material of the type species at my disposal from
which I have produced the diagnosis of the genus.
Gillettea Ashmead was synonymized with
Phanacis Forster by Eady and Quinlan (1963). I
have examined the type species of Gillettea and I
confirm the synonymy.
Diagnosis. — Head, viewed dorsally two times
broader than long; viewed frontally slightly
broader than high; frons and vertex reticulate-
coriaceous; radiating striae of face weakly im-
pressed, incomplete; female antenna 13-seg-
mented; male antenna with 14 antennomeres; A3
clearly longer than A4. Pronotum dorsally moder-
ately long, without distinct submedial pits;
mesoscurum reticulate-coriaceous; notauli usu-
ally faint anteriorly; scutellar foveae transverse,
confluent, not well defined or indistinctly closed
posteriorly; mesopleuron reticulate. Wings fully
developed in females, sometimes reduced in males;
radial cell of forewing usually closed, sometimes
incompletely on the margin; hair fringe on distal
wing margins moderately long. Third abdominal
tergum without lateral pubescent patch. Tarsal
claws simple.
Comments. — Phanacis is easily distinguished
from other genera of Aylacini excepting Timaspis
by the absence of pronotal submedian pits on the
pronotum and the reticulate sculpture of the
mesopleuron. The genus was considered by Eady
and Quinlan (1963) to comprise also the species
included in Timaspis Mayr. However, when some
species of the latter genus not examined by Eady
and Quinlan, are included in Phanacis, the hetero-
geneity of the grouping becomes evident. This is
the case with, e.g., Timaspis cichorii Kieffer and T.
phoenixopodos Mayr. The boundary between the
two genera does not, however, seem clearly de-
fined and some species, particularly P. caulicola
(Hedicke), P. hypochoeridis (Kieffer) and Timaspis
lampsanae Perris, are not easily assigned to one or
Volume 3, 1 994
191
the other genus. All the known species of Phanacis
(s. str.) produce galls on plants of the family
Asteraceae.
Distribution and Biology. — The genus has a
palaearctic distribution with two species intro-
duced in North America. All the known species
are associated with Asteraceae. Galls are produced
mostly on stems.
Phanacis taraxaci (Ashmead)
Gillettea taraxaa Ashmead 1897:69.
Phanacis taraxaci (Ashmead): Eady and Quinlan, 1963:18.
Examined material. — The type material con-
sists of two female specimens; one in the USNM
and the other in the NHMW (both examined); the
specimen from Wien, in better condition, is desig-
nated lectotype.
The species is being studied in detail by
Shorthouse (personal communication). Appar-
ently the species has been introduced in North
America, together with the host plant Taraxacum,
from Europe. But the existence of the species in
Europe is documented only by a few gall records,
so the european origin of this species does not
seem clear. The species is widely distributed and
relatively abundant in North America (Shorthouse,
pers. comm.).
RHODUS Quinlan
Rhodus Quinlan 1968:282. Type species: Rhodus oriundus
Quinlan 1968. Monotypic and original designation (ex-
amined)
Diagnosis. — Figs. 104-110. Head viewed dor-
sally (Fig. 104) two times broader than long; viewed
frontally (Fig. 105) clearly broader than high; f rons
and vertex with reticulate sculpture; radiating
striae of face present laterally; supraclypeal area
raised; female antenna (Fig. 106) 13-segmented;
male with 14 antennomeres; A3 clearly shorter
than A4. Pronotum (Fig. 107) dorsally long, with
distinct submedial pits; mesoscutum (Fig. 108)
with dull reticulate sculpture and some scattered
piliferous punctures; notauli distinct in posterior
3/4 of mesoscutum; faint or obsolete anteriorly;
scutellar foveae large, rounded, sculptured; sep-
tum relatively broad; mesopleuron (Fig. 109) lon-
gitudinally striated. Rl and Rs not quite reaching
anterior margin of wing; radial cell of forewing
(Fig. 110) open; hair fringe on distal wing margins
absent. Third abdominal tergum without lateral
pubescent patch. Tarsal claws simple.
Comments. — The genus was based on one spe-
cies, Rhodus oriundus Quinlan, found in Rhodes
Island, galling apical buds of Phlomis cretica Presl
(Quinlan 1968). Its morphological features relate it
to the genera Isocohis Forster and Neaylax Nieves-
Aldrey; it differs from the former in the conforma-
tion of the notauli, the sculpture of the mesoscutum
and the distal margin of the wing completely
lacking hairs. The wing venation with Rl and Rs
not quite reaching the anterior margin of wing and
the absent hair fringe on distal wing margin sepa-
rate it from Neaylax.
Distribution and Biologi/. — The genus comprises
one described species, only recorded from Rhodes
(Greece) associated with Phlomis (Lamiaceae).
TIMASPIS Mayr
Timaspis Mayr 1881:18. Type species: Diastrophus lampsanae
Karsh. Monotypic. (Type. — Timaspis phoenixoptodos Mayr
1882. Designated by Ashmead 1903 p.214).
Phanacis Forster (part): Eady and Quinlan 1963:18.
Aylacopsis Hedicke 1923:81. Syn. n. Type species: Aylacopsis
heraclei Hedicke 1923. Monotypic and original designa-
tion.
The type material of Timaspis lampsanae, in the
Museum of Vienna, consists of a 2 mounted on
micro-pin on polyporous strip labeled "Tim.
lampsanae det Karsch Type"; "lampsanae Karsch
Type"; "collect G. Mayr" and red label "Holo-
type".
Diagnosis. — Radiating striae present laterally
on the face; incomplete. Female antenna of 13-14
segments; 14-15 in males; A3 relatively long; at
least 1.5 times longer than A4. Pronotum dorsally
moderately long, without distinct submedial pits;
notauli usually complete but often not well im-
pressed anteriorly; median mesoscutal impres-
sion usually present; scutellar foveae small,
confluent, usually indistinctly closed posteriorly;
mesopleuron reticulate-rugulose, rugulose-
alutaceous or striate-alutaceous. Radial cell of
forewing closed on the margin, sometimes incom-
pletely so anteriorly; hair fringe on distal margins
of wing long. Third abdominal tergum without
lateral pubescent patch. Tarsal claws simple.
Comments. — Timaspis can be distinguished
from Phanacis on the basis of its rugulose-reticu-
192
Journal of Hymenoptera Research
late sculpture of the mesopleuron and complete
notauli. Timaspis was synonymyzed with Phanacis
Forster by Eady and Quintan (1963) but these
authors' concept of Timaspis was incomplete be-
cause it was limited to the type species of the
genus, T. lampsanae. But if other species described
within Timaspis are considered, especially T.
phoenixopodos Mayr, I consider it advisable to keep
the genera separate. It must be emphasized, how-
ever, that the boundary between the two genera is
not clearly defined and some species are difficult
to assign to one genus or the other.
Aylacopsis Hedicke, was described from a spe-
cies that produces galls in the stems of Heracleum
sphondylum L. (Apiaceae) (Hedicke 1923). After
having examined the type species, I consider that
it fits well within Timaspis Mayr. Consequently, I
synonymize Aylacopsis with Timaspis Mayr.
Distribution and Biology. — As understood
herein, the genus comprises 9 species in West-
Europe. All the species, excepting one, are associ-
ated with Asteraceae
Timaspis heraclei (Hedicke), comb. n.
Aylacopsis heraclei Hedicke 1923:81. Syn. n.
The type of Aylacopsis heraclei Hedicke in the Berlin Museum
consists of a 9 specimen glued on a card triangle. The
specimen carries the following labels: white handwrit-
ten "Doicul, Leipzig 26-2-15"; "Aylacopsis heraclei 9
Hedicke" and red printed "Type".
We can contribute the following complemen-
tary data to the original description: Figs. 111-118.
Head, viewed dorsally (Fig. Ill), about two times
broader than long; viewed frontally (Fig. 112)
slightly broader than high; frons and vertex finely
coriaceous; POL slightly less than two times OOL;
transfacial line about 1.6 times height of eye; face
laterally with radiating striae; supraclypeal area
without striae; with weak median protuberance-
malar space about 0.7 times height of eye; antenna
(Fig. 113) slender, filiform, with 14 antennomeres;
A3 slightly curved in the middle, about 1.7 times
longer than A2; 1.3 times longer than A4. Prono-
tum (Fig. 114) dorsally only moderately long;
scarcely pubescent, without submedial pits;
mesoscutum (Fig. 1 15) finely reticulate-coriaceous;
notauli complete; median mesoscutal impression
impressed in posterior 1 / 3 of mesoscutum; scutel-
lar foveae confluent, indistinctly closed posteri-
orly; scutellum ovate, weakly rugulose;
mesopleuron (Fig. 116) rericulate-rugulose. Forew-
ing with radial cell (Fig. 117) open anteriorly on
margin but R, clearly reaching wing margin; length
of radial cell 2.6 times width; hair fringe on wing
margins long. Third abdominal tergum (Fig. 118)
without lateral pubescent patch; following seg-
ments not punctate; ventral spine of hypopygium
quite short, slightly visible. Tarsal claws simple.
Comments. — Within Timaspis Mayr this spe-
cies is morphologically close to Timaspis cichorii
from which it is differentiated by the coloration,
sculpture of the mesoscutum and conformation of
the scutellar foveae. According to the original
description, this species produces galls in the stems
of Heracleum sphondylum (Apiaceae) at that time
being the first cynipid species demostrated to be
associated with this family of plants. Afterwards
this species has not been recorded again and,
consequently, the host has not been confirmed. It
may, however, be pointed out that Phanacis eryngii
Diakonchuk, another Aylacini species, was re-
cently found in Eastern Europe associated with
Apiaceae (En/ngium) (Diakonchuk 1984).
Timaspis lusitanica Tavares
Timaspis lusitanicus Tavares 1904:301
Timaspis lusitanica Tavares 1904: D.T. and Kieffer 1910 Emend.
Phanacis crepidos Weidner 1965:1. Syn. n.
Timaspis lusitanica Tavares was described from
Portugal (Tavares 1904) reared from galls on
Barkausia taraxacifolia DC. (Asteraceae). More re-
cently, it has been recorded from Spain, on the
same host plant (Nieves-Aldrey 1992). Phanacis
crepidos was described from Germany (Weidner
1965) from material reared from galls on Crepis
biennis L. (Asteraceae). Having examined the type
material from the Museum of Hamburg, I have
concluded that it is the same species described by
Tavares, so I establish the corresponding new
synonymy.
Volume 3, 1 994
193
XESTOPHANES Forster
LITERATURE CITED
Xestophanes Forster 1869:332, 337. Type species: Cynips
potentillae De Villers. Monotypic and original designa-
tion.
Diagnosis. — Face laterally with radiating striae;
supraclypeal area without striae; with weak me-
dian protuberance; frons and vertex almost smooth
and shining; antenna of female 13-segmented, A3
as long as or slightly longer than A4; antenna of
male 14 segmented. Pronorum dorsally long, lat-
erally pubescent; submedial pits conspicuous;
mesoscutum smooth and shining; notauli com-
plete or faint anteriorly; median mesoscutal im-
pression slightly indicated; scutellar foveae
smooth; mesopleuron smooth and shining. Forew-
ing with radial cell open on the margin but R,
reaching to wing margin; hair fringe moderately
long. Third abdominal tergum laterally smooth
and glabrous. Tarsal claws with weakly devel-
oped basal lobe or tooth .
Comments. — The genus, together with the al-
lied genus Diastrophus, form a distinct unit within
the Aylacini: morphologically, by the tarsal claws
having an acute basal lobe or tooth and frons,
vertex and mesopleuron without sculpture; bio-
logically, by their association exclusively with
Rosaceae (the genera Potentilla and Rubus). The
genus is very likely monophyletic, based on the
fusion of third and fourth abdominal terga in the
females
Distribution and Biology. — Two species of
Xestophanes are known in Western Europe, both
gall makers on stems and runners of species of
Potentilla. .
ACKNOWLEDGMENTS
I am very grateful to the curators of the collections listed
under materials for the loan of type material and other
specimens. Special thanks to A. Menke (USNM) and R.
Danielsson (MZLU) also for sending some interesting bibli-
ography. I am also most indebted to one anonymous re-
viewer and specially to F. Ronquist for a critical revision of
the manuscript including linguistic suggestions. M. A. Alonso-
Zarazaga helped with nomenclatonal questions. I would like
to thank Inaqui Diez for the rotulation of the drawings made
by myself. Financial support was provided by the Research
project "Fauna Ibenca II", DGICY PB89 0081.
Ashmead, W. H. 1885. A Bibliographical and Synonymical
Catalogue of the North American Cynipidae, with de-
scription of new species. Transactions of the American
Entomological Society 12: 291-304.
Ashmead, W. H. 1887. On the Cynipidous galls of Florida,
with descriptions of new species and synopses of the
described species of North America. Transactions of the
American Entomological Society 14: 125-158.
Ashmead, W. H. 1896. Description of new cynipidous galls
and gall wasps in the United States National Museum.
Proceedings of the United States National Museum 19: 113-
136.
Ashmead, W. H. 1897a. Description of some new genera in
the family Cynipidae. Psyche 8: 67-69.
Ashmead, W. H. 1897b. Description of five new genera in the
family Cynipidae. Canadian Entomologist 29: 260-263.
Ashmead, W. H. 1903. Classification of the gall-wasps and
the parasitic cynipoids, or the super-family Cynipoidea.
I-IV. Psyche 10: 140-155
Askew, R. R. 1984. The biology of gall wasps, pp. 223-271 . In T.
N. Ananthakrishnan (ed.), Biology of Gall Insects. Ed-
ward Arnold, London.
Barbotin, F. 1964. Sur une nouvelle galle et deux cynipides en
provenance d'Algerie. Marcellia 31: 151-157.
Belizin, V. I. 1959. Gall wasps of the tribe Aylaxim (Hy-
menoptera, Cynipoidea) new for the fauna of the USSR.
Entomologkheskoe Obozreme 38: 662-674.
Burks, B. D. 1979. Cynipoidea, pp. 1045-1107. In K. V. Krombein,
P. D. Hurd Jr., D. R. Smith, and B. D. Burks (eds), Catalog
of Hymenoptera in North America North of Mexico. Volume
1, Smithsonian Institution Press, Washington, DC.
Dalla Torre, C. W. and J. Kieffer. 1910. Cynipidae. Das Tierreich
24: 1-891.
Diakonchuk, L. A. 1984. New species of Cynipidae (Hy-
menoptera) from Georgian SSR. Vestnik Zoology 3: 74-77
(In Russian).
Eady, R. D. and J. Quinlan. 1963 Hymenoptera: Cynipoidea.
Key to families and subfamilies and Cynipinae (includ-
ing galls). Handbooks for the Identification ofBritish,lnsects
8(la): 1-81.
Folhot, R. 1964. Contribution a l'etude de la biologie des
cynipides gallicoles (Hymenopteres, Cynipoidea).
Annates des Sciences Naturelles, Zoologie, Paris. 12 ser. 6:
407-564.
Forster, A. 1860. Die zweite Centune neuer Hymenoptera
(Eucoela et Phanacis). V erhandlungen derNaturhistorischen
Vereins der Preussischen Rhemlande und Westphalens 17:
143-146.
Forster, A. 1869. Ueber die Gallwespen. V erhandlungen der
kaiserlich-koemglichen zoologisch-botamschen Gesellschafi
in Wien 19: 327-370.
Giraud, J. 1859. Signalements de quelques especes nouvelles
de cynipides et de leurs galles. Verhandlungen der
kaiserlich-koemglichen zoologisch-botanischen Gesellschaft
in Wien 9: 337-374.
Hartig, T. 1840. Uber die familie der Gallwespen. Germar
Enlomologische Zeitung 3: 322-358.
Hartig, T. 1843. Zweiter nachtrag zur naturgeschichte der
194
Journal of Hymenoptera Research
Gallwespen. Zeitschrift fur Entomologie 4: 395-422.
Hedicke, H. 1923. Beitrage zur kenntnis dercvnipiden (Hym.)
XII. Ein neues cecidozoon an Heracleum spondylium.
Zeitschrift fur Pflanzenkrankheiten 33(1-2): 81-83.
Hedicke, H. 1928. Beitrage zur kenntnis der cynipiden (Hym.)
XIV. Der Erzeuger der levantimschen salviagalle
Deutsche Entomologische Zeitschrift 14: 81-85.
Ionescu, M. A. and N. Roman. 1960. Un genre nouveau de
Cynipides gallicoles Endocauloma bicolor n g., n. sp.
(Hymenoptera, Cynipoidea). Revue de Bwlogie.Acadenuci
Republicu Populare Ronune Bucaresti 5(3): 221-225.
Ionescu, M. A. and N. Roman. 1962. New genera of gall maker
Cynipidae in the fauna of the rumanian people's repub-
lic: Weldiella aequalis n. gen. and Cecconia valerianeliae
Thomson. Revue de Biologie Academia Republicii Populare
Romine Bucaresti 7: 551-559.
KiefferJ.J. 1901 . Monographic desCynipides d' Europe et d' Algene
Hermann edit. Paris, t.I, 687 pp.
Kieffer, J. J. 1902. Revision du genre Aulax et des genres
limitrophes d'Aulax, avec quelques notes sur divers
autres Cynipides. Bulletin de la Societe d Histoire N aturelle
deMetz. 10:91-97.
Kinsey, A.C. 1920. Phylogeny of cynipid genera and biologi-
cal characteristics. Bulletin of the American Museum of
Natural History 62: 357-402.
Kovalev, O. B. 1982. Cynipid gall-makers of the subfamily
Aylacinae (stat. n.) (Hym., Cynipidae) and their species
described within the family Figitidae. Trudy
Zoologicheskogo Instituta Akademiya Nauk SSR 110:85-93.
(in russian)
Mayr, G. 1881. Die Genera der gallenbewohnenden
Cynipiden. 20, fahresbenchte der Communal-Oberrealschule
im l. Bezirke pp. 1-38.
Mayr, G. 1882. Die europaischen arten der gallen-
bewohnenden cynipiden. 21, fahresberichte der Commu-
nal-Oberrealschule im I. Bezirke pp. 1-44.
Nieves-Aldrey, J. L. 1984. Notas sobre los Aylaxini (Hvm,
Cynipidae, Cynipinae) de la Peninsula Iberica con
descripcion de una nueva especie de Isocolus. Eos 60:
235-250.
Nieves-Aldrey, J. L. 1985. Nuevos Aylaxini (Hym., Cynipidae)
para la Peninsula Iberica con descripcion de una nueva
especie de Aylax Htg. Boletim da Sociedade Porttuguesa de
entomologia, suppl. 1, Adas do II Congresso Iberico de
Entomologia 117-128.
Nieves-Aldrey, J. L. 1987. Estado actual de conocimiento de
la subfamiha Cynipinae (Hym.. Parasitica, Cynipidae)
en la Peninsula Iberica. Eos 63: 179-195.
Nieves-Aldrey, J. L. 1988. Descripcion de una nueva especie
de Isocolus Forster con notas de otras especies de Aylaxini
nuevas para la Peninsula Iberica (Hym., Cynipidae). Eos
64: 221-227.
Nieves-Aldrey, J. L. 1992. Adiciones a la lista de cinipidos
ibericos, especialmente Aylaxini, incluyendo la
descripcion de una nueva especie de/lH/aniifa Ashmead
(Hymenoptera, Cynipidae). Boletim da Sociedade
portuguesa de entomologia, suppl n°3, Adas do II Congresso
Iberico de Entomologia v. 1: 493-500.
Nieves-Aldrey, J. L. "In press". The types of Aylax Hartig
described by C.G.Thomson (Hymenoptera, Cynipidae).
Entotnologica Scandmavica.
Nordlander, G. 1984. Vad vet viom parasitiska Cynipoidea?.
Entomologisk Tidsknft 105: 36-40
Quinlan, J. 1968. Cynipinae (Hymenoptera) occurring on
Phlomis L. Transactions of the Royal Entomological Society
of London 120:275-286.
Ritchie, A. J. 1984. A revision of the higher classification of the
mquiline gall-wasps (Hymenoptera: Cynipidae) and a revi-
sion of the Neartic species of Periclistus Forster. Ph. D.
Thesis, Carleton University, Ottawa, ON. 368 pp.
Rohwer, S. A. and M M. Fagan. 1917. The type-species of the
genera of the Cynipoidea, or the gall wasps and para-
sitic cynipoids. Proceedings of the United States National
Museum, 53: 357-378.
Ronquist, F. In press. Evolution of parasitism among closely
related species: phylogenetic relationships and the ori-
gin of inquilinism in gall wasps (Hymenoptera,
Cynipidae). Evolution.
Ronquist, F. and G Nordlander 1989. Skeletal morphologv of
an archaic cympoid, Ibalia rufipes (Hymenoptera:
Ibaliidae). Entomological scandmavica Suppl. 33: 1-60.
Tavares, J. da S. 1904. Descnpcao de un Cynipide novo.
Broteria 3: 301-302.
Thomson, C. G. 1877. Oefversigt af sveriges Cynips Arter.
Opuscula Entomologica_8: 732-841.
Wachtl, F. 1880. Beitrage zur Kenntnis der gallenerz. Insecten
Europas. Verhandlungen der kaiserlich-koeniglichen
zoologisch-botamschen Cesellscliaft in Wien 30: 531-545.
Wachtl, F. 1891. Eine neue Gallwespe. Wiener Entomologische
Zeitung pp. 277-280.
Walsh, B. D. 1869. Galls and their architects. The American
Entomologist 2: 70-74.
Weidner, H. 1965. Phanacis crepuios sp. nov., eine neue
gallwespe an Crepis biennis L. Entomologische Mttteilungen
52: 1-7.
Weidner, H. 1968. Zur Kenntnis der Gallwespentnbus
Aulacini (Hymenoptera, Cynipidae). Entomologische
Zeitschrift 10: 105-120.
Weld.L. H. 1930. Notes on Types (Hymenoptera: Cynipidae).
Proceedings of the Entomological Society of Washington 32:
8.
Weld, L. H. 1952. Cynipoidea (Hym.) 1905-50. Privately printed,
Ann Arbor, MI, 351 pp.
Wiebes-Rijks, A. A. 1979. A character analysis of the species
of Synergus Hartig, Section II (Mayr, 1872) (Hy-
menoptera, Cynipidae). Zoologische mededelingen 53(28):
297-321.
Zerova, M. D., Diakonchuck, L. A. and V. M. Ermolenko.
1988. Gall-formers of the european part of the USSR Part 2 .
Hymenoptera. Naukova Dumka, Kiev. 180 pp.
Volume 3, 1 994
195
APPENDIX
Check list of the west-European genera and species of Aylacini including their host plant genera.
AYLACINI Ashmead 1903 Host plant genus
AULACIDEA Ashmead, 1897
abdominalis (Thomson, 1877) ?
arnicae Hoffmeyer, 1930 Arnica
follioti Barbotin, 1972 Sonchus
hieracii (Bouche 1834) Hieracium
artenusiae (Thomson, 1877)
crassinervis (Thomson, 1877)
foveiger (Thomson, 1877)
kerneri (Wachtl, 1891) comb, n Nepeta
kieffen ' Cotte, 1915
laurae Nieves-Aldrey, 1992 ?
nibletti Quinlan, 1969 Hieracium
pilosellae (Kieffer, 1901) Hieracium
scorzonerae (Giraud, 1859) Scorzonera
subterminalis Niblett, 1946 Hieracium
tragopogonis (Thomson, 1877) Tragopogon
tavaresi Nieves-Aldrey, 1984 syn. n.
scabiosae (Giraud, 1859) Centaurea
rogenhoferi Wachtl, 1880 syn. n.
serratulae (Mayr, 1882) comb, n Serratula
. Nepeta
LIPOSTHENES Forster
glechomae (Linnaeus, 1758)...
latreilki (Kieffer, 1898)
NEAYMXgen.n.
salviae (Giraud, 1859) comb, n Salvia
verbenacus (Nieves-Aldrey, 1988) comb, n Salvia
versicolor (Nieves-Aldrey, 1985) comb, n ?
PANTELIELLA Kieffer, 1902
ENDOCAULONIA Ionescu and Roman, 1960 syn. n.
fedtschenkoi (Rubsaamen, 1896) Phlomis
bicolor (Ionescu and Roman, 1960) Phlomis
AYLAX Hartig, 1840
hypecoi Trotter, 1912 Hypecoum
minor Hartig, 1840 Papavcr
papaveris (Perris, 1839) Papaver
artenusiae Weidner, 1968 misident.
BARBOT1N1A gen. n.
oramensis (Barbotin, 1964) comb, n Papaver
CECCON1A Kieffer, 1902
WELD/ELM Ionescu and Roman, 1962 syn. n.
valerianellae (Thomson, 1877) Valenanella
aequalis (Ionescu and Roman, 1962) Valenanella
DIASTROPHUS Hartig, 1840
rubi (Bouche, 1834) Rubus
mayri Remhart, 1876 Potentilla
HED1CK1ANA gen. n.
levantina (Hedicke, 1928) comb, n Salvia
IRAELLA gen. n.
luteipes (Thomson, 1877) Papaver
papaveris Kieffer in Goury and Gignon, 1905
1SOCOLUS Forster, 1869
fitchi (Kieffer, 1898) Centaurea
jaceae (Schenck, 1863) Centaurea
lichtensteini (Mayr, 1882) comb, n Centaurea
PHANACIS Forster, 1860
GILLETTEA Ashmead, 1897
caulicola (Hedicke, 1939) Picris
centaureae Forster, 1860 Centaurea
punctipleuris (Thomson, 1877)
hypochoendts (Kieffer, 1887) Hypochoens
taraxaci (Ashmead, 1897) Taraxacum
RHODUS Quinlan, 1968
onundus Quinlan, 1968 ...
. Phlomis
T1MASP1S Mayr, 1881
AYLACOPSIS Hedicke, 1923 syn. n.
cichorn Kieffer, 1909 Cichonum
heraclei (Hedicke, 1923) comb, n Heracleum
lampsanae Perris, 1873 Lampsana
lusitanica Tavares, 1904 Crepis
crepidos (Weidner,1968) syn. n.
phoenixopodos Mayr, 1882 Lactuca
pilicornis (Thomson, 1877) ?
rufipes Ionescu and Roman, 1959 Crepis
sonchi (Steffarti, 1900) Sonchus
urospermi Kieffer, 1901 Urospermum
XESTOPHANES Forster, 1869
brevitarsis (Thomson, 1877) Potentilla
potentillae (Retzius in Degeer, 1873) Potentilla
foveicollis (Thomson, 1877)
abreviatus (Thomson, 1877)
196
Journal of Hymenoptera Research
Figs. 1-15. Cynipidae. 1-3, Pronotum, dorsal view. 1, Aylax papaveris, 2, Aulacidea hieracii. 3, Cynips divisa 9. 4, Scutellum,
Pediaspis acens. 5, Gula, Penchstus brandtii. 6-7, Ceroptres 6, Gaster, lateral (9)7, Head, frontal view. 8, Gula, Isocolus. 9, Gaster,
lateral, Synergus ilicinus (2). 10, Gaster, lateral, Cecconia valerianellae. (9). 11-13, Eschatocerus. 11, Mesoscutum and scutellum,
dorsal view. 12, Forewing (hairs omitted). 13, Head, dorsal view. 14-15, Diplolepis rosae. 14, Mesopleuron. 15, Gaster, lateral
(2).
Volume 3, 1994
197
Figs. 16-34. Aylacini. 16-20, Tarsal claws. 16, Diastrophus rubi (hind). 17, Xestophanes potentillae (front). 18, X. potentillae (hind).
19. Barbotmia oraniensis (hind). 20, Isocolus Hchtensteini (hind). 21, Forewing, X. potentillae. 22, Forewing, D. rubi. 23-27, lraclla
luteipes (9). 23, Pronotum. 24, Head and thorax, lateral. 25, Forewing. 26, Head, frontal view. 27, Antenna, first segments. 28,
Mesopleuron, Phanacis caulicola. 29-30, Cecconia valerianellae 8. 29, Head, frontal view. 30, Forewing. 31-33, Liposthenus
glechomae 9. 31, Head and thorax, lateral. 32, Mesoscutum and scutellum, dorsal view. 33, Antenna, first segments. 34,
Forewing At/lax papaveris (9).
198
Journal of Hymenoptera Research
Figs. 35-42. Antistrophuspisum. 35, Head, dorsal view. 36, Head, frontal view. 37, Antenna ( 9 ). 38, Antenna ( 6 ). 39, Pronotum.
40, Mesoscutum and scutellum. 41, Body, lateral. 42, Forewing (hairs omitted).
Volume 3, 1994
199
Figs. 43-55. Aulacidea. 43-50, Aulacidea liarringtom. 43, Head, dorsal view. 44, Head, frontal view. 45, Antenna ( ? ). 46, Antenna
(§).47, Pronotum.48, Mesoscutum and scutellum. 49, Body, lateral. 50, Forewing (hairs omitted). 51-55, Aulacidea kerneri. 51,
Antenna (?). 52, Pronotum, 53, Mesoscutum and scutellum. 54, Body, lateral. 55, Radial cell of forewing (hairs omitted).
200
Journal of Hymenoptera Research
Figs. 56-62. Barbotinia oraniensis. 56, Head, dorsal view. 57, Head, frontal view. 58, Antenna ( 9 ). 59, Antenna ( 6 ). 60, Pronotum.
61, Mesoscutum and scuteilum. 62, Body and wings.
Volume 3, 1 994
201
%vj&
Figs. 63-70. Hedickiana levantina. 63, Head, dorsal view. 64, Head, frontal view. 65, Antenna ( 9 )
and Scutellum. 68, Mesopleuron. 69, Forewing (hairs omitted). 70, Gaster, lateral.
66, Pronotum. 67, Mesoscutum
202
Journal of Hymenoptera Research
Figs 71-79. hocolus scabwsae. 71, Head, dorsal view. 72, Head, frontal view. 73, Antenna (9). 74, Antenna (6). 75, Pronotum.
76, Mesoscutum and scutellum. 77, Mesopleuron. 78, Gaster, lateral. 79, Forewing (hairs omitted).
Volume 3, 1 994
203
Figs. 80-95. Isocolus and Aulacidea. 80-87, Isocolusserratulae. 80, Head, dorsal view. 81, Head, frontal view. 82, Antenna (9). 83,
Pronorum. 84, Mesoscutum and scutellum. 85, Body, lateral. 86, Forewing (hairs omitted). 87, Antenna {6). 88-95, Aulacidea
scorzonerae. 88, Head, dorsal view. 89, Head, frontal view. 90, Antenna ( 9 ). 91, Pronorum. 92, Mesoscutum and scutellum. 93,
Body, lateral. 94, Forewing (hairs omitted). 95, Antenna (6").
204
Journal of Hymenoptera Research
Figs. 96-103. Neaylax salviae. 96, Head, dorsal view. 97, Head, frontal view. 98, Antenna ( 9 ). 99, Antenna ( 9 ). 100, Pronotum.
101, Mesoscutum and scutellum. 102, Body, lateral. 103, Forewing (hairs omitted)
Volume 3, 1994
205
105
104
j^Vi^?
Figs. 104-110. Rhodus oriundus. 104, Head, dorsal view. 105, Head, frontal view. 106, Antenna (9). 107, Pronotum. 108,
Mesoscurum and scutellum. 109, Body, lateral. 110, Forewing (hairs omitted).
20b
Journal of Hymenoptera Research
Figs. 111-118. Timaspis heraclei. Ill, Head, dorsal view. 112, Head, frontal view. 113, Antenna (9). 114, Pronotum, 115,
Mesoscutum and scutellum. 116, Mesopleuron. 117, Forewing (hairs omitted). 118, Gaster, lateral.
J. HYM. RES.
Vol. 3, 1994, pp. 207-226
A Review of North American Belomicrus
(Hymenoptera, Sphecidae, Crabroninae)
Richard M. Bohart
Department of Entomology, University of California, Davis, CA 95616
Abstract. — The North American Belomicrus are reorganized into five groups containing a total of 40 species. Of these, 14
are described as new: californicus (central Sierra of California), costalis (central California), darwini (sw. U.S.); desertus (s.
California); myo (Inyo Co., California), longiceps (sw. U.S.), melanus, (central Sierra of California), montanus (central Sierra of
California), oraibi (Arizona and s. Utah), palhdus (s. California), powelli (coast range mts. of California), siccatus (s. California),
sierrae (central Sierra of California), texensis (Texas and Baja California Sur). Keys, illustrations, and distributions are given.
New synonymy: quemaya Pate is raised to full species; and jurumpa Pate is synonymized under quemaya.
Belomicrus Costa (1871) is one of several
crabronine genera in the tribe Oxybelini. Bohart
and Menke (1976:359-370) gave an overview of
present day knowledge of the tribe. They pointed
out that the presence of metanotal projections
(squamae) and a propodeal projection (mucro) in
addition to a lateral carina on terga I-III, and
sometimes IV-V, characterize Belomicrus. As far as
known, distribution of the genus is holarctic and
Ethiopian. Bohart and Menke listed 63 species of
which 25 were North American. I now recognize
40 species in the latter region, of which 14 are new.
Among earlier writers the work of Pate
(1940a,b) was outstanding. His overall assessment
of Belomicrus and its relatives was brilliant, and his
descriptions were meticulous. The only criticisms
that can be made are that some of his phylogenetic
discussions suffered from lack of material, and his
descriptions contain much detail irrelevant at the
species level.
Belomicrus are ground-nesting wasps, mostly
in sandy areas. Prey of the forbesii group are
nymphal Miridae (Bohart 1956, Evans 1969). The
franciscus group provisions with adult dasytid
beetles (Williams 1936, Bohart and Menke 1976).
Prey of the other groups are unknown.
Members of the genus are few in most collec-
tions. Through museum visits and borrowings I
have been able to examine about 600 specimens.
The majority of these are in the University of
California at Riverside museum and were col-
lected by P. H. Timberlake. Diligent work by Bohart
Museum collectors at Davis has made some 3,000
more specimens available for study.
Principal cooperating museums and individu-
als are listed below. Museum designations are
identified by their city locations in capital letters.
The manuscript was read by Dr. Arnold Menke
and Dr. Eric Grissell. Their comments and correc-
tions were quite helpful, and their assistance is
gratefully acknowledged.
BERKELEY— J. Powell, P. D. Hurd (deceased), Essig Mu-
seum, University of California at Berkeley.
DAVIS — L. S. Kimsey, S. Heydon, Bohart Museum, Univer-
sity of California at Davis.
LOGAN — G. E. Bohart, Entomology Museum, Utah State
University.
NEW YORK — ]. Rozen, Entomology Museum, American
Museum of Natural History.
OTTAWA — L. Masner, Biological Resources Division, Agri-
culture Canada.
PHILADELPHIA— S. Roback, Daniel Otte, D. Azuma, Acad-
emy of Natural Sciences.
RIVERSIDE— P. H. Timberlake (deceased), S. Frommer, En-
tomology, Museum, University of California at River-
side.
SAN FRANCISCO— W. J. Pulawski, P. Arnaud, Entomology
Museum, California Academy of Sciences.
VIENNA — M. Fischer, Naturhistorische Museum, Vienna,
Austria.
WASHINGTON— A. S. Menke, K. V. Krombein, U.S. Na-
tional Museum.
208
Journal of Hymenoptera Research
Terms used in the species key and in descrip-
tions which may need explanation are: epipleural
"button", small raised area just behind pronotal
lobe; T-I, T-II, etc. terga after propodeum; S-I, S-II,
etc., sterna after propodeum; LID, least interocular
distance; PD, puncture diameter; post-tegula, basal
wing sclerite.
The generotype, Belomicrus italicus A. Costa
(1871), has a palearctic distribution and is similar
to species in the forbesii group. However, it should
be placed in a different group because the clypeus
has a considerable flattened area on the median
lobe, and the more posteriorly blunt metanotal
projection is divided by a median carina.
The ten groups of North American Belomicrus
which Pate (1940b) proposed are now reduced to
five. Bohart and Menke (1976) considered only 2
groups, forbesii and cladothricis. With the discovery
of additional species it now seems desirable to
divide the forbesii group into four according to the
following key.
The purpose of this and other keys in the
paper is to give a quick means of making identifi-
cations. It is understood that the key user should
have access to at least a representative named
collection of Belomicrus for comparative purposes.
KEY TO THE SPECIES GROUPS OF NORTH AMERICAN BELOMICRUS
Metanotal projections (squamae) divided into two distinct lobes (Figs. 28-43), basally joined or separated
but without an outer membranous area; body rarely more than 4mm long; postocular tubercles well
defined; (Figs. 18-24) cladothricis group
Metanotal projections joined to form a roughly triangular plate with a posterior notch and an outer partly
membranous edge (Figs. 11-14), postocular tubercles various 2
Mandible of females and some males with an obtuse angle subapically on lower edge (Fig. 15),mesopleuron
with an omaulus vanyume group
Mandible with at most a slight inferior angle subapically, mesopleuron without omaulus 3
Clypeus with an anchorlike apical projection in female (Fig. 16), body about 2 mm long
apache group
Clypeus simply pointed or with a deflected apical bevel, body 3.5 mm long or more 4
Clypeus medially hoodlike, often somewhat wrinkled, not deflected apically but ending in a small point
(Fig. 17) forbesii group
Clypeus with an apicomedial polished triangle or deflected bevel (Figs. 3-7) franciscus group
Belomicrus forbesii group
The only previous key to the forbesii group is
that given as part of a more extensive one by Pate
(1940b:210). The forbesii part was based on about
65 specimens, of which some 43 were forbesii and
columbianus, 10 were cookii, seven were penuti (as a
subspecies of forbesii), 3 were coloratus, and two
were querecho. I have studied about 1,500 speci-
mens of the group.
Instead of the six now valid species recog-
nized by Pate, this figure has been raised to 10,
which includes four new species.
Members of the forbesii group have relatively
few "structural" differences. However, the colora-
tion and color patterns seem to be quite constant in
long series. Therefore, I am treating columbianus as
distinct from forbesii, sierrae as distinct frompenuti,
and desertus as distinct from cookii. Details on
differences are given in the key, which follows:
Volume 3. 1994 209
KEY TO THE BELOMICRUS FORBESII SPECIES GROUP
1 T-I with basal slope all or mostly red 2
— T-I with basal slope dark reddish brown, dark brown, or black 5
2 Postocular tubercles weak and obtuse, tergal pale yellow bands usually well developed on T-I and often
on T-II, weak or faint on T-III and following montanus R. Bohart
— Postocular tubercles well developed, tergal markings various 3
3 S-II and following with distinct, close, moderately fine punctures; S-II with a broad median depression
querecho Pate
— S-II and following with indistinct, close, fine punctures or other microsculpture; S-II with at most a median
flattened area 4
4 Terga without distinct pale yellow bands, these sometimes faintly visible; band across pronotum nearly
always broken a little inside pronotal lobe, wing membrane stained powelli R. Bohart
Terga with distinct pale yellow bands on most terga, band across pronotum complete to pronotal lobes,
wing membrane nearly clear desertus R. Bohart
5 Epipleural "button" cup shaped, cup usually with a membranous outer edge, or pronotal collar all black
6
— Epipleural "button" not cup shaped, only slightly indented, pronotal collar nearly always maculate
7
6 Pronotal collar all black, T-I to IV all dark or with indistinct bands, epipleural "button" pocket without a
lighter colored edge, postocular tubercles weak sierrae R. Bohart
— Pronotal collar maculate, terga with pale bands or lateral spots usually on T-I to V, epipleural "button"
pocket deeper and with a lighter colored edge, postocular tubercles often well developed
penuti Pate
7 Tergal markings weak, pale bands indistinct at least on T-III to V; pronotal collar black laterally and
sometimes medially columbianus (Kohl)
Tergal markings prominent; pronotal collar nearly always completely banded 8
8 T-I, II and sometimes III in female with dark ground color, following terga with red ground color; both sexes
with small but sharp postocular tubercles cookii Baker
T-I and most following terga in both sexes with reddish brown to black ground color; both sexes with at
most weakly to moderately developed, but not sharp, postocular tubercles 9
9 Tergal coloration in front of yellow bands brownish red or reddish brown, postocular area without definite
tubercles coloratus Baker
Tergal coloration in front of yellow bands dark brown to nearly black, postocular area nearly always with
low but definite tubercles forbesii Robertson
Belomicrus coloratus Baker Belomicrus columbianus (Kohl)
Belomicrus coloraia Baker 1909:29. Holotype female, Ormsby Oxybelus columbianus Kohl 1892:208. Lectotype female (ex-
Co., Nevada (DAVIS), examined. - amined and here designated), Revelstoke, British Co-
lumbia (VIENNA). Treated as a subspecies of forbesii by
I have studied 422 males and 140 females from Bohart and Menke (1976). New status.
Nevada and California. They were all taken in the Oxybelus larimerensis Rohwer 1908:417. Holotype female,
months of June and July at elevations of 4,000 to Lanmer Co " Colorado (WASHINGTON), examined,
oonnr lm j j • i i ^ , Treated as a synonym of forbesii bv Bohart and Menke
8,000feet. Nevada county records include Ormsby, (1976) Revised syn;nymy.
Douglas, and Eureka. California counties are Al-
pine, El Dorado, Inyo, Lassen, Mono, Nevada, In all/ 54 males ^j 69 females have been
Placer, Sierra, and Tuolumne. A single Oregon srudied. These were taken during the months of
record is 5 mi e. Bly, Klamath Co. May (rarely), June, July, and August, mostly at
The principle diagnostic features, especially elevations above 4,000 feet. Western state records
the tergal markings, are given in the key. include British Columbia, Alberta, Colorado,
Idaho, Montana, Nevada, Oregon, Utah, and Wash-
ington. California records are: Bell Echo Camp,
210
Journal of Hymenoptera Research
Dorris, and Rattlesnake Meadow, Siskiyou Co.;
Morrison Meadow and Snowslide Park, Trinity
Co.
The new status of columbianus and its syn-
onym, larimerensis is indicated by the difference in
markings from forbesii as outlined in the key.
Belomicrus cookii Baker
Belomicrus cookii Baker 1909:29. Lectotype male, Claremont,
Los Angeles Co., California (WASHINGTON). Lecto-
type designated by Pate (1940a).
This species is known from a few southern
California localities. I have seen 8 males and 11
females from Los Angeles Co. (Claremont,
Newhall) and Riverside Co. (Riverside, Gavilan),
all taken during the months of April and May at
low elevations in foothill locations.
Belomicrus desertus R. Bohart, new species
Female holotype. — Length 5 mm. Black, red are:
clypeal apex, foretarsus partly, tegula dully, terga
mostly, sterna except for dark median blotches on
T-I to T-III; whitish yellow are: mandible basally,
scape partly, flagellum beneath, apical spots on
femora, tibiae outwardly, squamal triangle, apical
bands on T-I to III, weak on IV-V; wings nearly
clear. Pubescence moderate, silvery and appressed
on clypeus, lateral frontal band enlarged above,
short supraclypeal spot, postocular, mesopleural,
and lateral tergal areas moderately silvered. Punc-
tation fine and close, a little shiny across frons
below ocelli; pygidial plate with coarse, separated
punctures. Postocular tubercles well developed,
epipleural "button" weakly indented.
Male. — Length 4-5 mm. Whitish yellow are:
mandible mostly, clypeus across apex, basal tarsal
segment, apical bands on T-I to VII (usually).
Holotype female (DAVIS), 3 mi s. Kramer
Junction, San Bernardino Co., California, IV-6-66
(R.M. Bohart). Paratypes, 123 males, 31 females
topotypical, collected in April of various years by
R.M. Bohart, E.I. Schlinger, M.E. Irwin, D.S.
Horning, J.C. Hall, F.D. Parker. Other paratypes
from San Bernardino Co., California: 3 males, 5
females, Joshua Tree National Monument, (H.K.
Court); 13 males, 4 females, Adelanto (E.I.
Schlinger, et al.): 6 males, 2 females, Apple Valley
(W. R. Mason, P. D. Hurd). Other specimens (not
paratypes) were from California counties: San Ber-
nardino (Victorville, Morongo Valley, Red Moun-
tain), Imperial (Palo Verde), Los Angeles (Llano,
near Lovejoy Buttes), Ventura (Chuchupate Ranger
Sta., Frazier Mt.), Kern (Dove Well), inyo (Panamint
Mts., 8 mi w. Brown). Paratypes in cooperating
museums.
This species is similar to powelli which also has
well developed postocular tubercles, the basal
slope of T-I red, and the epipleural "button"
scarcely indented. However, desertus usually has
extensive whitish yellow tergal markings. Some-
times, these may be on T-I only. See also poioelli.
Etymology. — specific name derived from Latin
adjective, desertus = abandoned.
Belomicrus forbesii (Robertson)
Figs. 12, 17
Oxybelus forbesii Robertson 1889:85. Holotype male, Colo-
rado (PHILADELPHIA), examined.
I have studied 118 males and 122 females,
collected from May to August at elevations of
4,000 to 10,000 feet in the following states and
counties: California (Alpine, Lassen, Modoc, Mono,
Nevada, Sierra, Siskiyou, Trinity), Colorado
(Larimer, Denver), Montana (Missoula), Nevada
(Elko, Pine, Washoe), Utah (Box Elder, Kane, Sum-
mit), and Wyoming (Sublette, Teton, Uinta). Fe-
male face (Fig. 17).
Diagnostic characters, particularly the abun-
dant tergal markings, are given in the key.
Belomicrus montanus R. Bohart, new species
Female holotype. — Length 5 mm. Black, red are:
clypeal apex, terga and sterna mostly except for
dark blotches on S-I-II; whitish are: mandible ba-
sally, scape apically, flagellum beneath, pronotal
collar and lobe, apical spot on forefemur, tibiae
outwardly, squamal triangle, T-I apex faintly;
wings lightly stained. Pubescence light, silvery
and appressed on clypeus, lateral frontal area,
short supraclypeal spot, postocular and
mesopleural areas moderately. Punctation fine
and close, not shiny across frons below ocelli;
pygidial plate with coarse, separated punctures.
Postocular tubercles quite weak, epipleural "but-
ton" slightly indented.
Male. — Length 4-4.5 mm. Foretarsus partly
Volume 3, 1 994
211
off-white, T-I whitish apically.
Holotype female (DAVIS), White Mts., Mono
Co., California, 10,000 ft., VII-10-68 (R.M. Bohart).
Paratypes, 14 males, 13 females, topotypical but
collected from June 22 to July 23 by J. W.MacSwain,
J. Powell, and G.I. Stage. Other California
paratypes: 1 pair, Benton, Mono Co., V-23-86 (R.M.
Bohart); 3 males, Westgard Pass, Inyo Co., V-VI-37
(G.A. Hamsher, CD. Michener); 2 females, Deep
Springs, Inyo Co., V-13, V-14 (L.D. French, N.J.
Smith). Paratypes in cooperating museums as far
as possible.
There is some similarity to querecho, powelli,
and desertus, since the basal slope of T-I is mainly
red, and the epipleural "button" is scarcely in-
dented. From all three of these the weakly indi-
cated postocular tubercles of montanus are differ-
entiating. Also, it may be separated from querecho
by the distinct sternal punctation of the latter, as
well as its broad depression on S-II.
Etymology. — specific name derived from Latin
adjective, montanus = dwelling on mountains.
Belomicrus penuti Pate
Belomicrns forbesii penuti Pate 1940:27. Holotype male,
Yosemite Valley, Mariposa Co., California (PHILADEL-
PHIA). Raised to species status by Bohart and Menke
(1976).
The 1,115 males and 51 females I have studied
were collected during April to September in Cali-
fornia and neighboring Nevada at elevations
mostly above 5,000 feet. California counties are:
Alpine, El Dorado, Fresno, Lassen, Mariposa,
Mono, Nevada, Placer, Calaveras, Sierra, Tulare,
Amador, and Tuolumne. Nevada records are In-
cline Village and Little Valley, Washoe Co. Also,
Pate (1940a:28) recorded penuti from southern
Oregon: Crater Lake and Lake of the Woods, Kla-
math Co. I have not studied these specimens.
The relationships to columbianus and forbesii
are outlined in the key.
Belomicrus powelli R. Bohart, new species
Female holotype. — Length 5 mm. Black, red are:
clypeal apex, terga and sterna mostly except for
dark blotches on S-II-III; whitish are: mandible
basally, flagellum beneath, pronotal collar medi-
ally, pronotal lobe, apical spot on femora, tibiae
outwardly, foretarsus dully, squamal triangle;
wings moderately stained. Pubescence silvery
appressed on clypeus, lateral frontal area, stout
supraclypeal triangle, postocular and mesopleural
areas (weakly). Punctation fine and close, not shiny
across f rons below ocelli, somewhat shiny on terga;
pygidial plate with coarse, separated punctures.
Postocular tubercles well developed, epipleural
"button" slightly indented.
Male. — Length 4-4.5 mm. Characters about as
in female. Scape whitish at apex.
Holotype female (SAN FRANCISCO), La
Panza, 12 mi ne. Pozo, San Luis Obispo Co., Cali-
fornia, IV-29-62 (J. Powell). Paratypes, 5 males, 16
females (all from California collected in April and
May): Monterey Co.: 1 male, 5 females, Hastings
Reserve (D.L. Linsdale); 1 pair, Arroyo Seco (D.
Burdick, P. Torchio); female, 12 mi n. Cholame
(D.H. Janzen); San Luis Obispo Co.: 2 pair,
topotypical (J. Powell, R.W. Thorp); female, 5 mi
ne. Santa Margarita (R.W. Thorp); Fresno Co.: 12
mi w. Coalinga (J.W. MacSwain); Ventura Co.: 3
males, 8 females, Chuchupate Ranger Sta., Frazier
Mt. (J. Powell, P.D. Hurd). Paratypes in SAN
FRANCISCO, DAVIS.
The well developed postocular tubercles are
also found in desertus, which see. The all red terga
and broken yellow pronotal band of powelli are
distinguishing. Although primarily a coastal form,
poioelli occurs with desertus on Frazier Mt., Ventura
Co., at Chuchupate Ranger Station. In fairly long
series the differences given in the key seem to
hold. The species is named for my friend, Jerry
Powell, a lepidopterist who has also worked with
wasps.
Belomicrus querecho Pate
Belomicrus querecho Pate 1940:36. Holotype male,
Alamogordo, Otero Co., New Mexico (PHILADEL-
PHIA), examined.
This southwestern species is largely restricted
to the Chihuahuan Life Zone. I have studied 3
males and 20 females, collected in April, May, and
June in the following states: Arizona (near Apache
and Willcox, Cochise Co.; Tubac, Santa Cruz Co.;
Oak Creek Valley Road, Yavapai Co.; 10 mi w.
Jacob Lake, Coconino Co.). Utah (25 mi s. Moab,
Grand Co.). New Mexico (Skeleton Canyon,
Peloncillo Mts., and Rodeo, Hidalgo Co.;
212
Journal of Hymenoptera Research
Alamogordo, Otero Co.; 5 mi e. Las Cruces and
Leasburg Dam, Dona Ana Co.). Nevada (4 mi s.
Warm Springs, Nye Co. ) and Texas (Rankin, Upton
Co.).
Belomicrus sierrae R. Bohart, new species
Female holotype. — Length 5 mm. Black or dark
brown, red are: flagellum beneath dully, foretarsus
dully, apical half of pygidial plate; whitish yellow
are: mandible basally, femorotibial joints, tibiae
outwardly, apicomedial spot on squamal triangle
(all yellow in some paratypes); wings moderately
stained. Pubescence silvery appressed on clypeus
laterally, narrowly on lateral frontal area, short
supraclypeal spot, weakly on postocular and
mesopleural areas. Punctation fine, close, dull;
clypeus wrinkled, with a pair of coarse punctures
above polished apex, pygidial plate with coarse,
separated punctures. Postocular tubercles present
but weak, epipleural "button" deeply indented
but not covered with a thin membrane; lateral
propodeal carina flattened, somewhat bent in-
ward.
Male. — Length 4.5-5 mm. About as in female.
Pronotal lobe sometimes dully pale.
Holotype female (DAVIS), Sagehen Creek,
Nevada Co., California, 6,500 feet, VI-25-68 (R.M.
Bohart). Paratypes, 118 males, 97 females,
topotypical, V-29 to VII-25 (R.M. Bohart, J. A. Skin-
ner, L.S. Kimsey, M.E. Irwin, D.S. Horning, J.E.
Slansky, J. Powell, B. Villegas). Also paratypes, 10
males, 19 females, Grass Lake, El Dorado Co.,
California, 8,000 ft., VII-5-62 and VII-16-62 (R.M.
Bohart et ai). Other specimens (not paratypes)
have been studied from other California counties
at 6,000 ft. or above: Alpine, Calaveras, Plumas,
Lassen, Placer, and Trinity. Paratypes in all coop-
erating museums.
The dark basal slope of T-I and the deeply
indented epipleural "button" ally sierrae with
penuti. However, the "button" is not as thinly
lidded above as in penuti. Futhermore, the latter
seems always to have at least several of the terga
with whitish bands. The dark abdomen ot sierrae is
similar to that of colwnbianus, and conceivably the
two species might occur together in California
near the Oregon border. In addition to the
epipleural "button" difference, columbianns has
the lateral propodeal carina simple, whereas in
sierrae it is not sharp but flattened and bent in-
ward.
Etymology. — the specific name refers to the
California Sierra.
Belomicrus apache group
The apache group contains only the single
known species, apache Pate. It was described in
detail from a single female by Pate (1940a) and
placed in a key by Pate (1940b). This is one of the
smallest known Belomicrus, approximately 2 mm
in length. The female has a peculiar anchoriform
terminal process figured by Pate (1940a) and herein
(Fig. 16). The male clypeus is more ordinary. Fea-
tures of the group are characterized in the key. A
brief description of the species, including the pre-
viously unknown male, and additional locality
records follow:
Belomicrus apache Pate
Figs. 13, 16
Belomicrus apache Pate 1940a: 15. Holotype female, Las Cruces,
Dona Ana Co., New Mexico (PHILADELPHIA), exam-
ined.
Female. — Length 2 mm. Black with red
unhanded abdomen; legs partly whitish yellow,
frons at narrowest point about equal to eye breadth,
clypeus with an apical anchoriform projection
(Fig. 16), no postocular tubercles or omaulus,
squamal triangle translucent laterally, deeply in-
cised posteriorly (Fig. 13).
Male. — Length 2 mm. Much as in female but
LID slightly greater and clypeal bevel triangular
rather than anchoriform.
I have seen three males and seven females
from the following western states: New Mexico
(Las Cruces, Doha Ana Co.; Rodeo, Hidalgo Co.),
Arizona (Aztec, and 7 mi s. Quartzite, Yuma Co.;
Quijotoa and Sells, Pima Co.; Willcox, Cochise Co.
on Euphorbia mat), California (Cathedral City, Riv-
erside Co. on Euphorbia poly carpa). Collections were
made from August to October.
Belomicrus vanyume group
The vanyume group consists of three species of
which one is described below as new. Pate (1940a,
b) placed vanyume in a monotypic group of its
own. The vanyume group, as I consider it, is unified
by the distinct omaulus running vertically along
the widest point of the mesopleuron, and ending
Volume 3, 1994
213
in a tooth or angle. This is followed halfway to the
midcoxa by another tooth. Females have the
forefemur right-angled subbasally. Except for
vanyume, males also have this feature. Further,
females have the mandible broadened before the
apex (Fig. 15).
KEY TO SPECIES OF THE BELOM1CRUS VANYUME GROUP
Postocular area a little swollen but not tuberculate, clypeus broadly rounded, scape light yellow, squamal
unit with windowlike clear areas vanyume Pate
Postocular area with prominent tubercles, clypeus with a longitudinal raised area medially, scape dark,
squamal unit various 2
Pronotum all dark, mesopleuron completely punctate and dull, clypeus (as seen in side view) not dentate
above bevel, T-II to V or III to V black texensis R. Bohart
Pronotum usually partly whitish, pronotal lobe white, clypeus ending above bevel in a forward-directed
tooth (Fig. 15), T-I to V usually red, especially in females maricopa Pate
Belomicrus maricopa Pate
Figs. 14, 15
Belomicrus maricopa Pate 1947:54. Holotype female, Higley,
Maricopa Co., Arizona (WASHINGTON), examined.
Belomicrus mariposa Pate 1947:55. In error.
I have seen 44 males and 58 females from
Arizona (Higley, Willcox, 28 mi s. Quartzite,
Picacho Canyon), New Mexico (5 mi e. Las Cruces,
9 mi n. Cotton City), California (18 mi w. Blythe,
Palo Verde), and Sonora, Mexico (Guaymas).
This species is similar to texensis, described
below and differences enumerated. The unde-
scribed male is much like the female but is a little
smaller and the mandible is somewhat less ex-
panded. Also, the last few terga are darker. The
principal differences between the two species are
the shinier mesopleuron and apically toothed
clypeus of maricopa (Fig. 15).
Belomicrus texensis R. Bohart, new species
Female holotype. — Length 5 mm. Black, light
yellow are basal two-thirds of mandible, foretibia
outwardly; brownish are: legs partly; reddish are:
forefemur at base, T-I to III; wings clear, veins
mostly orange. Silvery appressed pubescence on
clypeus mostly, broad patch along eye margin
reaching up as far as an imaginary line drawn
across beneath midocellus, short but large
supraclypeal patch. Punctation of mesonotum and
interocellar area coarse and separated by 1-2 PD of
microsculpture, mesopleuron similar but rugose
below, tergal punctation moderate and a little
shiny, coarse on pygidial plate. LID about 1.7x
scape length, clypeus with median raised area
bearing longitudinal depression, clypeal bevel
transverse and nearly linear, frontal groove well
impressed, postocular tubercles well developed,
gena in side view bent outward at middle below,
mandible strongly expanded at distal one-third,
front dorsal margin of pronotum obtusely cor-
nered, squamal unit as in Fig. 14, forefemur
rightangled near base, pygidial plate narrowly
triangular.
Male. — Length 4 mm. Legs mostly brownish
red, forewing veins dark, clypeal bevel broadly
triangular, T-I-II red, III a little red laterally and
posteriorly, VII dully red.
Female holotype (DAVIS), Kingsville, South
Pasture, Kleberg Co., Texas, VI-16-68 (J. E.
Gillaspy). Paratypes, 4 females (DAVIS, NEW
YORK), same data as holotype but V-6-67 and V-
6-68; on Ratibida columnaris; paratype male (SAN
FRANCISCO), Mexico: Baja California Sur, e. edge
of Sierra Placeres, 111-24-84 (W. J. Pulawski).
The large size (for Belomicms\), large postocular
tubercles, expanded mandible subapically and all
dark antenna are also found in maricopa. However,
texensis has many differences: pronotum all dark,
214
Journal of Hymenoptera Research
punctation of mesopleuron and mesonotum much
closer and not shiny, female clypeus with a longi-
tudinal flattened and shallowly grooved median
ridge, clypeus also without a forward-pointing
apical tooth, wing veins of female mostly orange
instead of black, marginal cell of forewing some-
what shorter, and T-III to V mostly or all black.
Etymology- — the specific name indicates "of or
from" Texas.
Belomicrus vanyume Pate
Belomicrus vanyume Pate 1940a:17. Holotype female,
Victorville, San Bernardino Co., California
(PHILADEPHIA), examined.
In addition to the type I have seen five males
and three females, all from California counties:
Riverside (Box Canyon, Thousand Palms), San
Bernardino (Apple Valley, Kramer Hills, Adelanto,
1 1 mi w. Ludlow), Imperial (Fish Creek Mts.), Inyo
(Darwin Falls). These were all taken in April, May,
and June. The previously undescribed male is
much like the female but the last two terga may be
dark.
Belomicrus franciscus group
The absence of an omaulus, triangular median
squamal complex (Fig. 11), mandible not angled
beneath toward apex, and clypeus with an apical
deflected bevel or polished triangle, when taken
together distinguish the group. I have identified
eight species, of which three are herein described
as new.
Species characters are the extent of tergal yel-
low markings, flagellarcoloration, pronotal macu-
lation, punctation of the postmandibular area of
males, development of the mandibular midtooth,
and form of the clypeal apex. Since females may be
difficult to distinguish, the following key is based
on males.
KEY TO MALES OF THE BELOMICRUS FRANCISCUS GROUP
Ventral area of head just posterior to mandibular insertion nearly all polished, flagellum relatively short,
flagellomeres hardly longer than broad (Fig. 2) 2
Ventral area of head just posterior to mandibular insertion distinctly but not closely punctate, flagellum
various 4
Clypeal apex arched (Fig. 5), T-I usually with a subapical yellow band or spot (central California below
3,000 feet) costalis R. Bohart
Clypeal apex not much, if any, arched; terga usually all red or with only a narrow streak of yellow on T-
I 3
Flagellum pale yellow beneath on I-IX or X; mandible tooth on inner margin well developed (Fig. 9);
punctures of interocellar area close but not contiguous, area a little shiny (central Sierra of California at
4,000 to 5,000 feet) californicus R. Bohart
Flagellum orange beneath before apex; mandible tooth on inner margin small (Fig. 8), punctation of
interocellar area contiguous and dull (low elevations in California from San Luis Obispo Co. to San Diego
Co.) serrano Pate
Ground color on T-I to T-VI black, tergal markings whitish (central Sierra of California above 5,000 feet)
melanus R. Bohart
Ground color on T-I to T-III, at least, red; tergal markings pale yellow : 5
Clypeal bevel produced downward laterally (Fig. 4), terga maculate 6
Clypeal bevel narrowed laterally, maculation various 7
Flagellum unusually long, mostly orange, many flagellomeres longer than broad as viewed laterally (Fig.
1); clypeal bevel often with a small median point (Fig. 4); terga often with yellow spots but rarely with
complete yellow bands (desert areas from Inyo Co., California and Clark Co., Nevada south to Sonora,
Mexico) quemaya Pate
Flagellum rather stout, dark above (Fig. 2), flagellomeres about as broad as long in lateral view; clypeal
bevel without a median denticle; terga with complete yellow banding (San Francisco)
franciscus Pate
Clypeal apex convex overall, bevel simple below, femora often extensively red (southern California)
cahuilla Pate
Clypeal apex more nearly flat, bevel margined below by a slight inflection, femora black with a small
amount of pale yellow (central Sierra of California at 5,000-7,000 feet) mono Pate
Volume 3, 1 994
215
Belomicrus cahuilla Pate
Belonucrus cahuilla Pate 1940:39. Holotype male, Andreas
Canyon, Riverside Co., California (PHILADELPHIA),
examined.
The male differs from that of mono by the
simple and convex clypeal bevel. In the female the
bevel is more indented than that in mono or quemaya.
Both sexes have dark median areas on the terga,
which may extend all across on T-III and follow-
ing. Only a single female has been identified, so
variation cannot be assessed. The type series of
four males came from Andreas Canyon (Palm
Springs). I have also seen five males from close by
(Taquitz Canyon, F. D. Parker and L. A. Stange).
Other southern California records are: male, fe-
male, near Cajon Junction, San Bernardino Co. (J.
C. Hall, E. I. Schlinger); males, Walker Pass and
West Wofford Heights, Kern Co. (J. Powell); male,
2 mi e. Banner, Riverside Co. (H. C. Dickson).
Dates of capture ranged from April 16 to June 7
(males), and July 4 (female).
Belomicrus californicus R. Bohart, new species
Figs. 7, 9
Belomicrus franciscus Pate of Bohart, in Bohart and Menke
1976:363. Misidentification.
Male holotype. — length 4.5 mm. Black, whitish
are: mandible mostly, scape in front, flagellum
beneath except at apex, pronotal collar medially,
lobe, squamal unit, tip of mucro, femora distally,
tibiae outwardly, foremetatarsus dully; red are:
clypeal bevel mostly, abdomen except for median
dark blotches on T-III to VI; wings weakly stained.
Silvery appressed pubescence along inner eye
margin, stout supraclypeal spot, weak pubescence
on postocular area, mesopleuron, and terga. Punc-
tation sparse on mostly polished postmandibular
area of head venter, fine and close on dorsum of
body, a little shiny on interocellar area. Clypeal
bevel quite thin (Fig. 7), distinctly darkened later-
ally; mandibular inner tooth well developed.
Female. — Length 5 mm. Clypeal bevel slightly
margined below, mandible (Fig. 9).
Holotype male (DAVIS), Baxter, Placer Co.,
California, elev. 5,000 ft., VI-16-56 (R. M. Bohart).
Paratypes: 18 males, 21 associated females, all
from the California Sierra at 4,000 to 5,000 ft.
elevation during May to July, in the following
counties: Placer (Dutch Flat, Baxter, Colfax),
Tuolumne (Strawberry), El Dorado (Pyramid
Ranger Station and near Icehouse Road). Collec-
tors were J. G. Rozen, J. W. MacSwain, R. M.
Bohart, W. J. Pulawski, and H. M. Kimball Court.
Paratypes are deposited in museums listed in
acknowledgements.
This species is close to Serrano. Both have the
male postmandibular area practically impunctate,
but in californicus the mandible tooth is well devel-
oped, and punctation of the interocellar area is a
little less dense.
J. MacSwain and I observed a large nesting
area in 1956 near Pyramid Ranger Station north of
Placerville, California. Females were provisioning
ground nests with dasytine beetles, Amecocerus
cervicalis Blaisdell (Melyridae).
Etymology. — the specific name indicates "of or
from" California.
Belomicrus costalis R. Bohart, new species
Fig. 5
Male holotype. — Length 4.5 mm. Black marked
with whitish yellow: scape and mandible mostly,
flagellum beneath, pronotum all across, post-
tegula, squamal unit, large forefemoral spot, small
distal midfemoral spot, tibiae and metatarsi out-
wardly, subapical band on T-I, attenuate laterad;
red are: clypeal bevel, T-I to III mostly, T-IV to VI
except for median dark blotches, T-VII, wings
weakly stained. Silvery appressed pubescence in
strip along inner eye margin, a little expanded
above, supraclypeal spot, postocular area and
mesopleuron moderately, terga lightly. Puncta-
tion sparse on mostly polished postmandibular
area of head venter, fine and close on dorsum of
body, somewhat reflective on interocellar area.
Clypeal bevel somewhat arched (Fig. 5).
Female. — Length 5 mm. Terga all red, scape
pale in front only, flagellum reddish yellow be-
neath. Clypeal bevel flat, not margined below;
mandibular inner tooth well developed.
Holotype male (DAVIS), Cache Creek Can-
yon, Yolo Co., California, IV-30-54 (R. M. Bohart).
Paratypes (all from California at low to moderate
elevations): male (DAVIS), 4 mi nw. Lake Berryessa,
Napa Co., V-12-61 (F. D. Parker); 2 males (DAVIS,
WASHINGTON), Sacramento, Sacramento Co.,
V-19-61 (R. M. Bohart); male (BERKELEY), Alpine
Lake, Marin Co., VI-6-57 (J. Powell); 2 males
216
Journal of Hymenoptera Research
(DAVIS), Midlake, Lake Co., V-10-56 (E. I.
Schlinger); male (BERKELEY), near Quincy,
Plumas Co., Vl-22-49 (J. W. MacSwain); 2 males
(NEW YORK), Tuolumne City, Tuolumne Co., V-
30-53 (J. G. Rozen); male (DAVIS), Boca, Nevada
Co., VI-19-62 (R. M. Bohart); male (DAVIS), Kyburz
Flat, Sierra Co., VII-15-76 (R. M. Bohart); male
(DAVIS), Daffodill Hill, Amador Co., VI-5-63 (R.
M. Bohart). Also, 6 presumed and associated fe-
males from localities listed above: Cache Creek
Canyon, Boca, Daffodill Hill, Sacramento.
The band on the pronotal collar is sometimes
broken in both sexes. The arched clypeal apex (Fig.
5) differentiates males from serrano and californicus
which agree with costalis in having the
postmandibular area polished. Females are best
distinguished by association with males.
Etymology . — specific name derived from Latin
noun, costa = side. The species occurs especially on
the western side of California.
Belomicrus franciscus Pate
Figs. 2, 3, 11
Belomicrus franciscus Pate 1 331 :77.Holotype male, Lone Moun-
tain, San Francisco, San Francisco Co., California (PHILA-
DELPHIA), examined.
I have studied several of the type series. The
rather stout male flagellum (Fig. 2), extensively
yellow-banded terga, and partly punctate male
postmandibular area distinguish this species. Fe-
male face (Fig. 3). The type series was collectecd by
F. X. Williams (1936) in sandhills of San Francisco.
Williams found a colony of the species in late April
and early May of 1930. Females were provisioning
with Trichochrous antennatus Mots. (Melyridae). In
all probability the type locality has been taken
over by a housing development.
Belomicrus melanus R. Bohart, new species
Fig. 6
Male holotype. — Length 4.5 mm. Black or
brown, whitish are: scape in front, mandible mostly,
flagellum beneath, pronotum medially, pronotal
lobe, post-tegula, squamal unit, tip of mucro, outer
distal spots on fore and midfemora, tibiae and
foremetatarsus outwardly, subapical band on T-I,
weak subapical bands on T-II to VI, apex of VII;
dark red: clypeal bevel; wings weakly stained.
Silvery appressed pubescence in strip along inner
eye margin nearly to level of midocellus, stout
supraclypeal spot, postocular area, mesopleuron
moderately, terga lightly. Punctation moderate on
postmandibular area of head venter, fine and close
on dorsum of body, slightly reflective on
interocellar area. Clypeal bevel thin all across (Fig.
6).
Female. — Length 5 mm. Whitish bands on T-I-
V sometimes better developed in para types, prono-
tum sometimes white all across, pygidial plate
partly red. Postmandibular area and rest of head
venter polished. Mandibular inner tooth well de-
veloped, clypeal bevel thin but impressed over its
entire breadth.
Holotype male (DAVIS), Sonora Pass,
Tuolumne Co., California, elev. 9,624 ft., VII-6-61
(R. M. Bohart). Paratypes (all from California Si-
erra at elevations above 5,000 ft.): female, same
data as holotype; 3 females, Hope Valley, Alpine
Co., VII, 1948, 1978 (R. M. Bohart); male, Fred's
Place, El Dorado Co., VII-10-67 (R. M. Bohart);
male, Gold Lake, Sierra Co., VII-8-54 (R. M. Bohart).
Paratypes all in DAVIS museum.
This species belongs to the subgroup in which
males have the postmandibular area somewhat
punctate. It differs from other members of the
subgroup by the extensively black (rather than
red) ground color of the terga. In cahmlla the terga
may be partly black but the male clypeal bevel is
not rimmed above, and the terga are not spotted
with whitish yellow.
Etymology. — specific name derived from the
Greek melanos = black.
Belomicrus mono Pate
Belomicrus serrano mono Pate 1940:46. Holotype female, 5 mi
n. Round Valley, Mono Co. California (PHILADEL-
PHIA), examined.
Belomicrus mono Pate, of Bohart and Menke 1976:363.
The male belongs to the subgroup with punc-
tate postmandibular area on the head venter. The
characters given in the key separate it. Females are
best identified by association with males.
I have seen 83 males and 65 associated females
from 5,000 to 7,000 feet, in the California Sierra of
the following counties: Mono (Round Valley holo-
type), Placer (Lake Tahoe), Sierra (Yuba Pass, Si-
erra Valley, Independence Lake, Sattley), Nevada
Volume 3, 1 994
217
(Sagehen Creek), Glenn (Plaskett Meadows). The
Sagehen Creek locality is represented by a long
series. Collection dates are mostly in June and
July.
Belomicrus quemaya Pate
Figs. 1, 4, 10
Belomicrus quemaya Pate 1940:47. Holotype male, PalmSprings,
Riverside Co., California (PHILADELPHIA), examined.
Belomicrus jurumpa Pate 1940:53. Holotype male, Gavilan,
Riverside Co., California (PHILADELPHIA), examined.
New synonymy.
Belomicrus franciscus quemaya Pate, of Bohart and Menke
1976:363.
The punctate postmandibular area of the head
venter in the male places quemaya in its subgroup.
The unusually long and lightly colored flagellum
of both sexes and the distinctive clypeal bevel of
the male (Figs. 1, 4) are distinguishing. Wings are
nearly clear in both sexes. Females are best identi-
fied by association with males. B. jurumpa appears
to be a simple synonym based on an examination
of the holotypes.
I have studied 60 males and 30 associated
females collected from March to May. California
county records are: Riverside (Whitewater Can-
yon, Gavilan, Thousand Palms, 18 mi w. Blythe,
Palm Springs), San Bernardino (Kramer Jet.,
Adelanto), Kern (Short Canyon, Iron Canyon),
Imperial (Kane Springs, Fish Creek Mts., Palo
Verde), Inyo (Mazourka Canyon, Lone Pine), San
Diego (Borrego). Other records are Clark Co.,
Nevada (Searchlight, Jean), and Sonora, Mexico (9
mi e. Kino).
Belomicrus serrano Pate
Fig. 8
Belomicrus serrano serrano Pate 1940:42. Holotype male,
Tujunga Wash, San Gabriel Mts., Los Angeles Co., Cali-
fornia (PHILADELPHIA), examined.
The polished postmandibular area of the male
places the species in the same subgroup as costalis
and califomicus. The dull interocellar area and
weak mandible tooth (Fig. 8) separate serrano.
Also, the relatively simple male clypeal bevel, and
the indented one of the female are additional
characters.
I have seen 46 males and 27 females in addi-
tion to the type series. The species is widespread in
southern California. The range includes low el-
evation localities in the following counties: Los
Angeles, Riverside, San Bernardino, San Diego,
Kern, Ventura, and San Luis Obispo. Collection
dates are from March 27 to May 7.
Belomicrus cladothricis group
The deeply divided squamal lobes of the met-
anotum (Figs. 28-43), and short body length, char-
acterize the group. With the exception of cladothricis
Cockerell (1895), all previously known species
were described by Pate (1940a, 1947). The group
appears to be largely confined to western United
States, only a few species occurring east of the
100th meridian or in Sonora and the Baja Califor-
nian peninsula of Mexico. Considering a small
amount of synonymy and the 6 new species de-
scribed herein, the total number of known species
in the group is now 18. In most of these the squamal
lobes are separated at the base by dark integu-
ment. In some species, however, the pale color and
setal texture may be continuous basally, at least in
females. These are cladothricis, eriogoni, and sechi.
Species characters of the tiny forms in this
group are squamal structure, scutal and interocellar
punctation, frons breadth, clypeal bevel details,
and the somewhat variable coloration. In one spe-
cies, sechi, the unusual amount of silvery pubes-
cence, and shape of the male scape are distinguish-
ing.
Although little is known about the life history
of these species, they are all presumably ground
nesting (personal observation of eriogoni at Antioch,
California), and they are most frequently collected
on the flowers of Eriogonum. Other flower hosts
reported are Chilopsidis, Lepidospartum, Euphorbia,
Cladothrix, Cleomella, Rhamnus, Chrysothamnus,
Solidago, Cercidium, Prosopis, Sphaeralcea, Asclepias,
Baeria, and Salsola. Because of the small size of
species in the cladothricis group (2-4 mm long), it
can be surmised that they, like many other di-
minutive sphecids, provision with thrips.
A great deal of descriptive matter on the group
was given by Pate (1940a, b, 1947). However, he
had a limited amount of material to work with and
many of his new species were based on one or two
specimens, often of a single sex. His remarks on
geographical range must not be taken too literally.
For instance, on timberlakei he said (1940a:93),
"Known at present only from the two specimens
218 Journal of Hymenoptera Research
recorded above from Riverside, timberlakei will in Transverse Ranges district of southern Califor-
all probability eventually be found rather wide- nia". As I have shown later in this paper, Pate's
spread throughout the southern piedmont of the conclusion was far off the mark.
KEY TO SPECIES OF THE CLAD0THR1CIS GROUP
1 Scutum partly polished, punctures widely spaced 2
— Scutum with rather close punctation 3
2 Forewing veins normally dark, squamae separated by less than 3x mucro width (Fig. 34)
mescalero Pate
Forewing veins practically all white, squamae separated by about 4x mucro width (Fig. 33)
pallidus R. Bohart
3 Pronotum all dark including lobe 4
Pronotum partly whitish, at least on pronotal lobe 6
4 Distance between squamal apices about half length of scutellum (Fig. 35), area between squamae not
pointing posteriorly timberlakei Pate
Distance between squamal apices at least two-thirds length of scutellum (Figs. 41, 42), area between
squamae various 5
5 Area between squamae pointed posteriorly (Fig. 43), scutal punctures moderate and close, terga brown to
black bridwelli Pate
— Area between squame truncate (Fig. 41), scutal punctation quite coarse, T-I or T-I-I1I with some red
potawatomi Pate
6 Squamae relatively long and angled inward so that apices are not or hardly farther apart than width of
mucro (Figs. 37, 39,40), LID 1.3x to 1.6x length of scape 7
— Squamal apices not unusually long or angled inward, apices farther apart than width of mucro, LID various
9
7 Scutal punctation fine and close, not reflective, interocellar area likewise, T-I not yellow spotted or banded,
LID 1.4-1.5x length of scape pachappa Pate
— Scutal punctation a little uneven, some punctures medium fine; surface reflective, interocellar area
likewise, tergal markings and LID various 8
8 Scutal and interocellar punctation a little more coarse and distinct, terga without yellow markings
tuktum Pate
— Scutal and interocellar punctation a little finer and less distinct, T-I or T-I-II often with yellow markings
istam Pate
9 Abdomen black; squamae slender, crescent shaped (Fig. 22), scutal punctation medium coarse with
polished interspaces of 1 PD or more; scape whitish, flagellum partly pale beneath; pronotal collar
whitish all across; tibiae and tarsi whitish vierecki Pate
Abdomen with some pale markings or at least with T-I usually somewhat reddish, squamae not so crescent
shaped, scutal punctation with microsculptured interspaces, antenna and pronotal collar various, tibiae
and tarsi various 10
10 Head unusually long and narrow (Figs. 21, 24), inner eye margins nearly parallel below ocelli, postocular
tubercles prominent and sharp, female genal area depressed and densely silvery below (genal carina
undeveloped), frontal groove extending below broadly rounded brow, hindtibia and hindmeta tarsus all
whitish outwardly 11
Head not unusually long and narrow (Figs. 18, 20), inner eye margins more strongly curved, female genal
area not depressed, frontal groove and leg markings various 12
11 Flagellum extensively pale beneath, forewing veins mostly pale, T-I to III (male) or T-I to VI (female) with
prominent whitish bands, squamae somewhat curved but shorter than their interapical distance (Fig. 36)
oraibi R. Bohart
Volume 3, 1 994
219
— Flagellum dark beneath, forewing veins mostly brown, T-I-VI unbanded, squamae strongly curved within
and longer than their interapical distance (Fig. 38) longiceps R. Bohart
12. Body with exceptional amounts of silvery pubescence (Fig. 30), interocellar area and vertex silvery setose,
propodeum usually extensively red, male scape strongly swollen distally (Fig. 27) sechi Pate
— Body with moderate amounts of silvery pubescence, interocellar area and vertex not silvery, propodeum
black, male scape not swollen 13
13 Flagellum at least partly lighter colored toward base 14
— Flagellum practically all dark 16
14 Flagellum mostly light orange with some dark spots laterally, squamae moderately stout (Fig. 32)
darivini R. Bohart
— Flagellum somewhat lighter toward base beneath; squamae short and stout 15
15 Interocellar area, vertex, and scutum dull; terga red, at least toward base; female frons length below
midocellus 1.6x LID (Fig. 22) eriogoni Pate
— Interocellar area, vertex, and scutum somewhat shiny; T-I or II often pale marked, female frons length
below midocellus 1.9x LID (Fig. 18) cladothricis Cockerell
16. Frons length below midocellus 1.2-1.3x LID (Fig. 19), wing veins often mostly orange (female) or light
brown (male) rather than dark brown to black siccatus R. Bohart
— Frons length below midocellus 1.7-1.9x LID (Fig. 20), wing veins mostly dark brown to black 17
17. Mid and hindmetatarsi brown or brownish red (Fig. 26) postocular tubercles small as seen laterally or
dorsally, LID in female usually nearly equal to scape length, female scape whitish in front
inyo R. Bohart
— Mid and hindmetatarsi white (Fig. 25), female only known, postocular tubercles large, LID a little broader,
scape all whitish cucamonga Pate
Belomicrus bridwelli Pate
Fig. 43
Belomicrus bridwelli Pate 1940a:93. Male holotype, Clifton,
Fairfax Co., Virginia (PHILADELPHIA), examined.
In addition to the holotype and four topotype
paratypes, I have also studied the following: male,
Branford, Suwannee Co., Florida, VII-31-30; 2
males, female, Fort Bragg, Cumberland Co., North
Carolina; female, Dayton, Rhea Co., Tennessee,
VI-20-54; male, 15 mi nw. Big Spring, Howard Co.,
Texas, VI-1 3-63. According to these meager records,
bridwelli occurs east of 102° latitude in May, June,
and July. Female squamae and mucro (Fig. 43).
Belomicrus cladothricis (Cockerell)
Fig. 18
Oxybelus cladothricis Cockerell 1895:309. Female lectotype,
Las Cruces, New Mexico (PHILADELPHIA), examined.
Lectotype designated by Cresson (1928).
Belomicrus cladothricis prosopidis Pate 1940a:72. Male holo-
type, Palm Springs, Riverside Co., California (PHILA-
DELPHIA), examined.
Belomicrus minidoka Pate 1940a:79. Male holotype, Hagerman,
Gooding Co. Idaho (WASHINGTON), examined.
This rather wide-ranging and relatively abun-
dant species is represented in the DAVIS collec-
tion by 450 males and 198 females taken at 51
localities during March to October. All of the
captures were made south of 41.5° longitude and
west of 100° latitude. Outlying states in the range
are Nebraska, Texas, Utah, Idaho, and California.
Elevations were moderate to low and many were
in strictly desert areas. As might be expected of
such a common and widespread species, there is a
considerable amount of variation. The short and
rather broad squamae are a consistent feature.
Also, the narrow LID of females (Fig. 18) helps
identification. Many western examples have yel-
low spots on basal terga. Punctation of the scutum
and interocellar area is close but allows some
shininess not found in several related species such
as eriogoni. The legs are usually extensively red-
dish but this occurs in some other species. T-I-II
are often reddish in males but II may be all dark
brown as in the type of prosopidis, or I-II brown as
in the type of minidoka.
220
Journal of Hymenoptera Research
Belomicrus cucamonga Pate
Fig. 25, 31
Belomicrus cucamonga Pate 1940a:76. Female holotype, Camp
Baldy, San Gabriel Mts., San Bernardino Co., California
(PHILADELPHIA), examined.
In addition to the holotype I have seen three
females from Big Pine, Inyo Co., California, VII-
10-61 (R. M. Bohart); and a female from Batchelder
Springs, Inyo Co., VI-10 (H. K. Court). The more
abundant whitish of the scape and tarsi (Fig. 25), as
well as the larger postocular tubercles and broader
frons, distinguish cucamonga from inyo. Female
squamae and mucro (Fig. 31).
Belomicrus darwini R. Bohart, new species
Fig. 32
Female holotype. — Length 3 mm. Black or dark
brown, white are: mandible mostly, scape in front,
pronotal lobe, squama, mucro distally; reddish
are: clypeal bevel, flagellum (pale orange, spotted
laterally), tibia and tarsi mostly (brownish red),
hindtibia lighter in basal ring, dull apical bands on
T-I to III; wings clear, veins black. Silvery ap-
pressed pubescence extending upward along two-
thirds of inner eye margin, short supraclypeal
patch, weak on genal and mesopleural areas. Punc-
tures of scutum and interocellar area medium fine,
slightly separated by microsculpture. LID about
1.4x scape length, frontal suture relatively deep
halfway from midocellus to clypeus, clypeal bevel
margined and triangular, postocular tubercle small,
squamae short and slightly curved (Fig. 32).
Male. — Length 2.5 mm. Scape mostly brown-
ish, flagellum a little spotted laterally as viewed in
front.
Holotype female (DAVIS), Darwin Falls, Inyo
Co., California, May 17, 1970 (R. M. Bohart).
Paratypes, 2 males, 2 females, same data as holo-
type but collected by E. E. Grissell and R. M.
Bohart. Other paratypes, female (DAVIS), Wikiup,
Mohave Co., Arizona, IV-11-57 (T. R. Haig); fe-
male (RIVERSIDE), Kyle Canyon, Clark Co., Ne-
vada, V-4-41 (P. H. Timberlake); female (DAVIS),
near Boron, Clark Co., Nevada, VI-7-41 (P. H.
Timberlake).
As indicated in the key, the rather close punc-
tation, moderate LID, brownish legs and terga,
short squamae, light orange flagellum, and long
frontal suture characterize the species. The flagel-
lum is a little longer and more slender than in other
species.
Etymology. — named for Dr. Darwin French,
who discovered Darwin Falls in 1860.
Belomicrus eriogoni Pate
Fig. 22
Belomicrus cladothrias eriogoni Pate 1940a:70. Male holotype,
San Lucas, Monterey Co , California (PHILADELPHIA),
examined.
Belomicrus eriogoni Pate of Bohart and Menke (1976).
I have studied 240 males and 165 females in
the DAVIS collection. These were taken during
May to September in California, Arizona, New
Mexico, Nevada, and Utah. California records are
from Mono Co., Lassen Co., Sacramento Co., and
Contra Costa Co., south to the Mexican border.
Mexican records are from Baja California. At times
the species is abundant, especially on the flowers
of Eriogonum fasciculatum. Female facial propor-
tions (Fig. 22).
Belomicrus inyo R. Bohart, new species
Fig. 20, 26, 28
Female holotype. — Length 2.5 mm. Black, white
are: mandible mostly, scape in front, pronotal
lobe, squama, foretibia outwardly, mid and
hindtibiae on basal one- third; reddish are: clypeal
bevel, abdomen (more brownish on T-II to VI);
wings clear, veins black. Silvery appressed pubes-
cence is narrow along lower half of inner eye
margin, supraclypeal patch conical, moderate on
gena and mesopleuron. Punctures of scutum and
interocellar area medium fine, slightly separated
by microsculpture. LID about equal to scape length
(Fig. 20); frontal suture not deep; faintly visible
halfway from midocellus to clypeus; clypeal bevel
triangular, postocular tubercles small; squamae
curved, well separated distally (Fig. 28).
Male. — Length 2.5 mm. Hindtibia mostly or-
ange with dirty white basal area, mid and
hindmetatarsi orange or reddish brown. LID about
1.7x scape length.
Holotype female (DAVIS), Antelope Springs,
Inyo Co. California, V-10-61 (R. M. Bohart).
Paratypes, 2 males, 1 female (DAVIS, BERKE-
LEY), topotypes, VI-29-61, VII-1-61, VIII-24-60, on
Volume 3, 1 994
221
Solidago (P. D. Hurd, G. I. Stage).
The orange to brown metatarsi (Fig. 26), dark
flagellum, narrow female LID (Fig. 20), basally
banded mid and hindtibiae (Fig. 26), close and
dull punctation, and well separated squamae (Fig.
28) characterize the species.
Etymology. — specific name is an acronymic
noun inspired by Inyo Co., where all of the type
series was collected.
Belomicrus istam Pate
Fig- 37
Belomicrus istam Pate 1940a:81. Male holotype, 4.5 mi nw.
Edom, Riverside Co., California (PHILADELPHIA),
examined.
I have seen a total of 37 males and 62 females
collected from March to June in sandy areas of 11
localities. These range in California from Sacra-
mento to Borrego Valley, and in Nevada from
Nixon to Stillwater. I collected a long series of both
sexes lighting on a sandy slope in Sand Canyon,
Kern Co., California, on April 14, 1962. Female
squamae and mucro (Fig. 37).
Belomicrus longiceps R. Bohart, new species
Figs. 24, 38
Female holotype. — Length 2 mm. Black, white
are: mandible mostly, pedicel partly, pronotal lobe,
post-tegula, squama, mucro, tibiae outwardly,
tarsi; reddish are: clypeal bevel, abdomen; wings
clear, veins brown. Silvery appressed pubescence
present along lower half of eye margin, short
supraclypeal patch, dense on genal area and
mesopleuron. Punctation on scutum and
interocellar area fine and close but a little irregu-
lar, somewhat shiny. Head unusually long and
narrow (Fig. 24), inner eye margins nearly straight
(Fig. 24), frontal carina distinct one-third of dis-
tance from midocellus to clypeus, LID 1.7x length
of scape, clypeal bevel broader than high,
postocular tubercle prominent and sharp, genal
area flattened to a little concave inside row of erect
hairs, squamae short but curved and well sepa-
rated (Fig. 38).
Male. — length 1.8 mm. T-I-II partly black, T-
III-VII black.
Holotype female (DAVIS), 6 mi nw. Adelanto,
San Bernardino Co. California, V-ll-79, on
Spliaeralcea (R. M. Bohart). Para types collected April
to August, 4 males, female (DAVIS, WASHING-
TON), 18 mi w. Blythe, Riverside Co., California
(M. E. Irwin, F. D. Parker, R. M. Bohart, R. W.
Brooks); male (DAVIS), 15 mi w. Baker, San Ber-
nardino Co., California (N. J. Smith); female (BER-
KELEY), Surprise Canyon, Inyo Co., California (P.
D. Hurd); male (DAVIS), Patrick, Washoe Co.,
Nevada (E. E. Grissell); female (NEW YORK), 2 mi
n. Rodeo, Hidalgo Co., Nevada (J. G. Ehrenfeld).
Characteristic are the long head (Fig. 24), dark
antenna, sharp postocular tubercles, extensively
silvery female gena and mesopleuron, brown wing
veins, and unhanded abdomen. The only other
species with a rather long head (compare Figs. 21,
24) is the much more abundantly marked oraibi,
which see.
Belomicrus mescalero Pate
Fig. 34
Belomicrus mescalero Pate 1940a:87. Female holotype,
Alamogordo, Otero Co., New Mexico (PHILADEL-
PHIA), examined
This species is known to occur in April and
May. In addition to the three female types, one of
which came from Needles, California, I have stud-
ied the following: male, female, 3 mi sw. Rodeo,
Hidalgo Co., New Mexico, IV-30-65 (J- G. Rozen).
The partly polished scutum, together with the
dark wing veins, mostly red terga, and whitish-
yellow spotted T-I are distinguishing. The male is
in general agreement with the female description
given by Pate, but T- VI- VII are reddish. Female
squamae and mucro are shown in Fig. 34.
Belomicrus oraibi R. Bohart, new species
Figs. 21, 36
Female holotype. — Length 3 mm. Black, white
are: mandible mostly, scape in front, pronotal
lobe, tegula, post-tegula, metanotum, mucro dis-
tally, tibiae and tarsi outwardly (a little reddish),
T-I to VI mostly; reddish are: scape partly, flagel-
lum beneath, clypeal bevel, femora distally, tibiae
partly, terga basally; wings clear, most veins light
orange. Silvery appressed pubescence along lower
three-fourths of eye margin, short supraclypeal
patch, dense on genal area and mesopleuron. Punc-
tation fine, close and dull on scutum, a little reflec-
222
Journal of Hymenoptera Research
tive on interocellar area. Head longer than usual
(Fig. 21 ), inner eye margins nearly straight, frontal
carina distinct, LID 1.7x length of scape, clypeal
bevel broader than high, postocular tubercles
prominent and sharp, genal area slightly concave
inside row of erect hairs; squamae short, curved,
well separated (Fig. 36).
Male . — Length 2.5 mm. About as in female but
T-IV-VI black. Genal area not concave.
Holotype female (DAVIS), Oraibi, Navajo Co.,
Arizona, VIII-30-76, on Chrysothamnus (R. M.
Bohart). Paratype female (WASHINGTON),
Dinnebito Wash, 5 mi nw. Hotevilla, Navajo
Co., Arizona, VIII-31-86 (A. S. Menke); paratype
male (DAVIS), 23 mi nw. Page, Kane Co., Utah, VI-
22-70 (N. J. Smith); paratype male (SAN FRAN-
CISCO), Green River, 5 mi ne. Jensen, Uinta Co.,
Utah, VI-26-82 (W. J. Pulawski).
Although generally similar to lotigiceps, which
also has an unusually long head, oraibi differs by
its pale wing veins and extensively white terga.
The concavity of the lower gena of the female is
even more pronounced. The presumed males from
Utah agree with the female in punctation and
wing vein color. However, they have T-IV to VI
dark. The presently known range includes north-
ern Arizona and southern Utah.
Etymology. — the name is an acronymic noun
inspired by the Oraibi indians of Navajo Co.,
Arizona.
Belomicrus pachappa Pate
Fig. 40
Belomicrus pacliappa Pate 1940a:73. Female holotype, River-
side, Riverside Co., California (PHILADELPHIA), ex-
amined
I have studied 11 males and 30 females col-
lected from April to August at 4 localities in Cali-
fornia: Weed, Siskiyou Co. (J. Powell); Sand Can-
yon, Kern Co. (R. M. Bohart); Chuchupate Ranger
Station, Ventura Co. (J. Powell, P. D. Hurd); River-
side, Riverside Co. (P. H. Timberlake). Nevada
records axe from Washoe Co.: Wadsworth (R. M.
Bohart, G. I. Stage); Patrick (R. M. Bohart). The 3
species, pachappa, tuktum, and istam, have the squa-
mae elongate and nearly touching distally. The
close and dull punctation of pachappa separates it.
The female has T-I-VI red, but the male has the
abdomen dark following T-I or T-II. Female squa-
mae and mucro (Fig. 40).
Belomicrus pallidus R. Bohart, new species
Fig. 23, 33
Male holotype. — Length 2 mm. Black, white
are: mandible mostly, antenna mostly but dark
distally in front, pronotum all across, tegula and
post-tegula, wing veins, squamae and mucro, legs
beyond basal two-thirds of femora; reddish are:
legs basad of white markings, abdomen; wings
clear. Silvery appressed pubescence on face below
midocellus except for a median bare spot (Fig. 23),
genal and mesopleural areas densely silver, scutum
with scattered silvery setae. Punctation fine and
sparse on extensively polished mesonorum, a little
less so on interocellar area. LID (just above anten-
nal insertion) 1.4x scape length, inner eye margin
weakly curved below midocellus (Fig. 23); frontal
groove and clypeal bevel (if any) obscured by
silvery pubescence; postocular tubercle promi-
nent, sharp; squamae short, curved, pubescent,
well separated (Fig. 33).
Female.— Unknown.
Male holotype (SAN FRANCISCO), Borrego,
San Diego Co., California IV-25-54 (P. D. Hurd).
Paratype male (DAVIS), Blythe, Riverside Co.,
California, on Asclepias, VII-8-56 (M. Wasbauer).
The rather extensive silvery pubescence is
reminiscent of sechi, but there are many points of
difference, one of which is the simple scape in
pallidus male. General features characterizing
pallidus are the considerably polished mesonorum,
whitish wing veins, and red and white legs.
Etymology, — specific name based on the Latin
adjective pallidus = pale.
Belomicrus potawatomi Pate
Fig. 41
Belomicrus potawatomi Pate 1947:57. Female holotype, Sioux
City, Woodbury Co., Iowa (WASHINGTON), exam-
ined.
The holotype was collected July 12, 1935 by C.
N. Ainslee. The only other record of which I am
aware is: female, Regnier, Cimarron Co., Okla-
homa, June 9 (NEW YORK). This species is similar
to bridwelli which also has the squamae far apart.
However, the metanorum is posteriorly pointed
(compare Figs. 41, 43) in bridwelli and its scutal
punctation is less coarse.
Volume 3, 1994
223
Belomicrus sechi Pate
Figs. 27, 30
Belomicrus sechi Pate 1940a:60. Male holotype, 6 mi n. Palm
Springs, Riverside Co. California (PHILADELPHIA),
examined.
I have seen 17 males and five females of this
extensively silvery pubescent species. They were
taken from May to August in the following desert
localities of southern California: Palm Springs,
Magnesia Canyon, and Rancho Mirage, all River-
side Co.; 11 mi w. Ludlow, San Bernardino Co.;
and Borrego Valley, San Diego Co.; a male (SAN
FRANCISCO) comes from Arizona: Roper Lake
State Park, Graham Co., V-26-83 (W. J. Pulawski).
The male has a unique, distally swollen scape (Fig.
27). Otherwise, the female agrees with the descrip-
tion given by Pate. The female squamae and mu-
cro are shown in Fig. 30.
Belomicrus siccatus R. Bohart, new species
Figs. 19, 29
Female holotype. — Length 2.5 mm. Black, white
are: mandible basally, pronotal tubercle, squa-
mae, mucro distally; light dull orange are: wing
veins, tibiae and tarsi; dark red are: T-I-II, py-
gidium; wing membrane clear. Silvery appressed
pubescence in a broad patch reaching up along
two- thirds of inner eye margin, supraclypeal patch
undeveloped, postocular and mesopleural patches
moderate. Punctation of scutum and interocellar
area fine and a little separated, somewhat shiny.
LID 2x scape length, inner eye margin gently
curved (Fig. 19), postocular tubercles present but
not sharp, squamae nearly straight (Fig. 29).
Holotype female (DAVIS), Thousand Palms
Canyon, Riverside Co., California, IV-9-64 (R. M.
Bohart). Paratypes, from Riverside Co.: 2 males
(DAVIS), Whitewater, IV-17-87 (N. J. Smith); 7
males (DAVIS), SAN FRANCISCO, WASHING-
TON), same data as holotype; 2 male topotypes
(DAVIS), 111-29-77 (R. M. Bohart, N. J. Smith); 2
male topotypes (DAVIS), 111-31-63 (F. D. Parker);
female topotype (DAVIS), IV-11-70 (R. M. Bohart).
Also a paratype female (RIVERSIDE), Apple Val-
ley, San Bernardino Co., California, V-20-41 (P. H.
Timberlake).
The combination of unusually broad frons
(Fig. 19) black flagellum, nearly straight squamae
(Fig. 29) orange (female) to brownish (male) wing
veins, and orange-tinted leg markings character-
ize the species.
Etymology. — specific name derived from the
Latin adjective siccus = dry.
Belomicrus timberlakei Pate
Fig. 35
Belomicrus timberlakei Pate 1940a:91. Male holotype, River-
side, Riverside Co., California (PHILADELPHIA), ex-
amined.
This species has been collected from April to
September. I have studied 41 males and 1 1 females
from California, Nevada, Utah, Arizona, and New
Mexico. In California 13 localities range from
Siskiyou Co. to Riverside Co., mostly in the Tran-
sition Life Zone. One record is at 8,000 ft. on
Carson Pass, Alpine Co. Non-California records
are: Cochise Co., Arizona (Willcox, Apache, Santa
Rita Mts.); Eureka Co., Nevada (Eureka); Churchill
Co. Nevada (Frenchman); Grant Co., Utah (Moab);
Emery Co. Utah (Greenriver); Hidalgo Co., New
Mexico (Rodeo); Baja California, Mexico (10 mi e.
San Quintin).
Of the 3 species with all dark pronotum
(timberlakei, potazoatomi and bridzoelli) only
tnnberlakei has the squamae only moderately sepa-
rated (Fig. 35). The female resembles the male in
nearly all respects but its LID equals 2x the scape
length instead of 1.7.
Belomicrus tuktum Pate
Fig. 39
Belomicrus tuktum Pate 1940a:84. Male holotype 5 mi e. Edom,
RiversideCo., California (PHILADELPHIA), examined.
In addition to the holotype I have studied 24
males and 25 females collected from March to
May. The 13 localities represented include the
following California counties: Inyo (Darwin Falls,
Little Lake, Surprise Canyon), Kern (Sand Can-
yon), Ventura (Quatal Canyon), Riverside (Thou-
sand Palms, Deep Canyon, 18 mi w. Blythe,
Whitewater Canyon), San Bernardino (5 mi n.
Barstow), Imperial (Glamis), and San Diego
(Borrego Valley). This species is quite similar to
istam. Females have the abdomen essentially all
dark red, but males usually have T-I dark red, the
rest brown. Female squamae and mucro are shown
in Fig. 39.
224
Journal of Hymenoptera Research
Belomicrus vierecki Pate
Fig. 42
Belomicrus vierecki Pate 1940a:56. Male holotype, Alamogordo,
Otero Co., New Mexico (PHILADELPHIA), examined.
This is a relatively abundant and widespread
species. I have seen 272 males and 47 females
collected from March to July. I swept a long series
from flowers of Chilopsis linearis at 15 mi w. Baker,
San Bernardino Co., California in May. Other pre-
ferred hosts are Cercidium and Prosopis. The 15
localities for California are all sandy situations at
low elevations south of 38.6 longitude. Arizona
records are from Wickenburg, Tucson, and Santa
Rita Mts. New Mexico records are Alamogordo,
Las Cruces, and near Laguna, Valencia Co. Mexi-
can material has been seen from Sonora (Alamos),
Puebla (Petlalcingo), Durango (Nombre de Dios),
and Baja California Sur (La Ribera). The most
eastern record in the United States is Presidio,
Texas at about 104.5° latitude.
The dark brown abdomen, white tibiae and
tarsi, extensively polished interocellar area, nar-
row LID, and well separated crescentic squamae
(Fig. 42), make identification relatively simple.
REFERENCES
Baker, C. F. 1909. Studies in Oxybelidae I. Pomona journal of
Entomology 1:27-30.
Bohart, R. M. 1956. Prey captures of Belomicrus penuti and B
coloratus. In Bohart and Menke 1976: 363. Ibid.
Bohart, R. M. and A. S. Menke. 1976. Sphecid Wasps of the
World. A Generic Revision, ix + 695 pp. Univ. California
Press, Berkeley
Cockerell, T. D. A. 1895. The second Anacrabro and the small-
est American Oxybelus. Canadian Entomologist 27: 308-
309.
Costa, A. 1871 . Prospetto sistematico degh Immenotterologia
Itahana. Ann Mus. Zool. Univ. Napoli 6: 28-83.
Cresson, E. T. 1928. The types of Hvmenoptera in the Acad-
emy of Natural Sciences of Philadelphia other than
those of Ezra T. Cresson. Memoirs of American Entomo-
logical Society 5: 1-90.
Evans, H. E. 1969. Notes on the nesting behaviour of Pisonopsis
clypeata and Belomicrus forbesu. Journal of the Kansas
Entomological Society 42: 117-125.
Kohl, F. F. 1892. Neue Hymenopterenformen. Ann. Nat Hist
Hofmus Vienna 7: 197-234.
Pate, V. S. L. 1931 . A new Belomicrus from the West. Ent. News
42: 77-78.
Pate, V. S. L. 1940a. The taxonomy of the Oxybeline wasps I.
A review of the genera Belomicroides, Bnmocelus and
Belomicrus with particular reference to the nearctic spe-
cies. Transactions of the American Entomological Society 66:
1-99.
Pate, V. S. L. 1940b. The taxonomy of the Oxybeline wasps II.
The classification of the genera Belomicrus and
Enchenncrum . Transactions of the American Entomological
Society 66: 209-264.
Pate, V. S. L. 1947. New North American Belomicrus. Proceed-
ings of the Entomological Society of Washington 49: 54-57.
Robertson, C. 1889. Synopsis of North American species of
Oxybelus. Transactions of the American Entomological Soci-
ety 5&. 77-85.
Rohwer, S. A. 1908. Four new Hvmenoptera. Entomological
News 19:417-420.
Williams, F. X. 1936. Notes on two oxybehd wasps in San
Francisco. Pan-Pacific Entomologist 12: 1-8.
Volume 3. 1 994
225
1. quemaya
2. franciscus
3. franciscus
4. quemaya
5. costalis
6. melanus 7. californicus
8. serrano
15. maricopa
16. apache
17. forbesii
Figs. 1-17. 1, 2, male antenna xlOO; 3, female face; 4-7, male clypeus, anterior view xlOO; 8-10, female mandible x75; 11-14,
female scutellum, squamae, and mucro, not to scale; 15-17, female face, not to scale.
226
Journal of Hymenoptera Research
18. cladothricis
19. siccatus
20. inyo
21. oraibi
33. pallidus 34. mescalero 35. timberlakei 36. oraibi 37. istam 38. longiceps
39. tuktum
41. potawatomi 42. vierecki
43. bridwelli
Figs. 18-24, female face x50; 25-26, female hindtibia and metatarsus x50; 27, male antenna xlOO; 28-43, female squamae and
mucro, dorsal xlOO. Silvery scales on face and squamae shown as dark setal areas.
J. HYM. RES.
Vol. 3, 1994, pp. 227-231
Myosomatoides Gen. Nov., (Hymenoptera: Braconidae), A Neotropical Larval
Parasitoid of Stem-borer Pests, Diatraea (Lepidoptera: Pyralidae)
Donald L. J. Quicke
Department of Biology, Imperial College at Silwood Park, Ascot, Berks SL5 7PY, England
Abstract.-A new genus, Myosomatoides Quicke (type species: Myosoma pennipes Westwood 1882), from South America is
described and illustrated. Ipobracon pennipes Myers 1931 (non Westwood), originally recorded as a parasitoid of an unspecified
Diatraea species from British Guiana and subsequently reared from D. angustella Dyar, is congeneric with Myosomatoides
pennipes (Westwood) comb. nov. and is therefore a junior subjective homonym of the former. A new name, Myosomatoides
myersi is proposed for Myers's species and a key is provided to enable its separation from M. pennipes (Westwood) comb. nov.
INTRODUCTION
Diatraea, a large genus of New World pyralid
moths, whose larvae bore in the stems of
graminaceous plants, includes several species of
considerable economic importance in both the
temperate and particularly the tropical regions
where they attack sugarcane, corn and sorghum
(Elias 1970, Chippendale 1979, Agnew et al. 1988,
Pashley et al. 1990). Several genera of parasitic
wasps of the braconid subfamily Braconinae have
been recorded as attacking species of Diatraea, viz.
Bracon Fabricius (Muesebeck 1925, Shenefelt 1978),
Digonogastra Viereck (Wharton et al. 1989),
Myosoma Brulle (Quicke 1989) and Palabracon
Quicke (Quicke 1988a). All of these are believed to
be ectoparasitoids attacking relatively late instar
host larvae. In addition to these, Myers (1931)
described and illustrated three species of
Braconinae reared from Diatraea spp. in British
Guiana, all of which he placed in Ipobracon
Thomson, though he recognized that one of his
new species, I. pennipes Myers (non Westwood),
was an "isolated species". Ipobracon was subse-
quently shown to be a junior synonym of
Cyanopterus Haliday (Quicke 1985, 1987) and the
limits of that genus were restricted such that the
great majority of New World species described
under both Ipobracon and Iphiaulax Foerster, and
including the other two species described by Myers,
actually belong to a distinct, though related genus,
Digonogastra Viereck (Quicke 1988b, Wharton et
al. 1989). As Myers implied; however, I. pennipes
does not belong to Digonogastra, and in fact, it is a
member of a new genus near Myosoma Brulle
which is described below.
By coincidence, Westwood (1882) had earlier
described another species belonging to the same
new genus under the name of Myosoma pennipes,
which therefore becomes a senior, subjective hom-
onym of pennipes Myers. In addition to the type
specimens of Myosomatoides pennipes (Westwood)
comb. nov. (Hope Entomological Collections, Ox-
ford) and M. myersi comb, et nom. nov. (Natural
History Museum, London: type no. 3C 418) the
author has seen seven other specimens belonging
to the new genus from Argentina, Brazil, Colom-
bia, British Guiana, Ecuador, Paraguay, and Peru,
though none of these has any associated host data.
Both M. myersi and M. pennipes appear to be wide-
spread in South America.
Available material suggests that there are only
two species of Myosomatoides in South America,
viz. M. pennipes and M. myersi nom. nov. A key is
provided to enable their separation. Some varia-
tion is apparent for both species. In particular,
specimens of M. pennipes vary in the extent of the
black coloration on the body, the wing coloration,
the extent and length of setosity, the degree of
compression of the hind leg, the shape of the head,
the relative lengths of hindwing veins lr-m and
SC+RI, and the relative length of the ovipositor.
228
Journal of Hymenoptera Research
Because no two specimens examined were found
to be entirely similar, I consider it most likely that
they represent a single rather variable species
rather than a diverse polyspecific complex though
the latter possibility cannot be completely ex-
cluded.
Myers (1931) provided a fairly complete de-
scription of M. myersi (= pennipes Myers non
Westwood), in which he illustrated the rather
distinctive metasoma. Following Myers' original
record, the species was recorded as a parasitoid of
Diatraea, on several further occasions (Myers 1932,
Thomson 1953) and, in particular, of D. angustella
Dyar in Argentina (Parker et al. 1953, De Santis &
Esquival 1966). Members of the related genus
Myosoma are similarly idiobiontectoparasitoids of
stem-boring pyralid moth larvae, including
Diatraea and Chilo species (Quicke 1989, Quicke &
Wharton 1989). At least one species of Myosoma is
gregarious.
Myosomatoides is being described here to make
the name available to biocontrol workers in South
and Central America and for a forthcoming manual
to the New World genera of Braconidae (Marsh &
Wharton, in preparation).
Terminology follows that of Achterberg (1979,
1988).
MYOSOMATOIDES Quicke gen. nov.
Type species: Myosoma pennipes Westwood, 1882
Diagnosis. — Superficially similar to Myosoma
except that the 2nd metasomal tergite is strongly
pinched-up in the middle to form a mid-longitudi-
nal ridge (Fig. 4). In addition, the hind femur and
tibia are extremely broad and laterally compressed
and have very long, dense setosity (Fig. 3). In
Quicke & Sharkey's (1989) key to the North Ameri-
can genera of Braconinae, Myosomatoides will key
to Bracon Fabricius, from which it can be distin-
guished by its extremely compressed hind femur
and tibia.
Description. — Head. Antenna with approxi-
mately 50 flagellomeres. Terminal flagellomere
acuminate. Medial flagellomeres wider than long.
Scapus small, shorter ventrally than dorsally in
lateral aspect, emarginate apico-laterally, not emar-
ginate apicomedially . Clypeus very shallow; with-
out a transverse carina separating reflexed ventral
part from dorsal part; dorsal part punctate. Clypeus
separated from face by weak groove. Malar area
weakly impressed, punctate. Face densely setose
except for smooth, shiny, glabrous supraclypeal
triangular area. Antennal sockets approximately
level with middle of eye. Frons very weakly im-
pressed with indistinct midlongitudinal sulcus;
with moderately dense short setosity.
Mesosoma. Mesosoma smooth, shiny. Notauli
indicated only by weak depressions at anterior of
mesoscutum. Scutellar sulcus narrow, smooth.
Propodeum simple, without carinae.
Forewing. Veins 1-M and 1-SR+M straight,
vein cu-a interstitial (or virtually so), the 2nd sub-
marginal cell rather long (i.e. vein 3-SR 0.67-0.95
times SRI), vein m-cu less than or equal to 0.5 times
length of 2-SR.
Hindwing. Vein lr-m more or less straight,
slightly shorter than, to slightly longer than, vein
SC+R1. Apex of vein C+SC+R with one especially
thickened bristle. Base of wing evenly and densely
setose. Vein 2-1A absent.
Legs. Claws with large, acutely pointed, basal
lobes. Telotarsi swollen in dorsal aspect compared
to basal tarsal articles. Hind leg and often mid-leg
(though less so) extremely laterally compressed,
tibia markedly expanded in lateral aspect (Figs.
1,2,4).
Metasoma. Smooth, shiny, sparsely setose.
First metasomal tergite rather long, approximately
1.5-2.0 times longer than wide, and without cari-
nae. Second tergite transverse, with
midlongitudinal carina. Third to seventh tergites
without transverse grooves subbasally or sub-
posteriorly, without anterolateral areas, and mem-
branous posteriorly. Hypopygium not extending
beyond apex of metasoma, pointed in lateral as-
pect. Ovipositor (part exserted beyond apex of
metasoma) approximately two-thirds length of
metasoma, with a pre-apical dorsal nodus and
apicoventral serrations.
Male genitalia. Digitus with two rather small,
well separated apical teeth. Parameres not extend-
ing beyond base of digitus; setation restricted to a
fairly narrow band at apex. Volsellae virtually
glabrous. Basal ring moderately produced
medioanteriorly, approximately as long as wide.
Internal anatomy. Rectum small with four
circular rectal pads. Male with a pair of weak,
pouch-shaped, intertergal gland reservoirs be-
tween abdominal tergites 7 and 8, and with similar
but weaker intertergal glands between tergites 6
Volume 3. 1 994
229
Figs 1-4. Myosomaloides penmpes (Westwood) gen. et comb, nov., Light photomicrographs of male from Brazil. 1. Habitus,
lateral aspect. 2. Detail of hind leg. 3. Head and mesosoma lateral aspect. 4. Metasoma and hind legs, dorsal aspect. Scale lines:
1=2 mm; 2=0.6 mm; 3=0.7 mm, 4=0.8 mm.
230
Journal of Hymenoptera Research
flattened hind tibiae and general habitus simply
represent convergence cannot be totally excluded.
As with Myosomatoides, members of the genus
Myosoma are parasitoids of lepidopterous grass-
stem borers including Pyralidae and Sesiidae
(Maeto 1992).
ACKNOWLEDGMENTS
and 7; abdominal tergum 8 without obvious glands.
Etymology. — Name based on the existing ge-
neric name Myoscma and the Greek suffix -oides
indicating close affinity; gender masculine.
Distribution. — M. myersi: Argentina, Brazil,
British Guiana, Colombia, Ecuador, and Paraguay;
M. pennipes: Brazil (Amazonia) and Peru.
Biology. — M. myersi nom. nov. is a larval para-
sitoid of members of the pyralid borer genus
Diatraea. Nothing is known of the biology of M. I would like to thank Tom Huddleston (The Natural
pennipes (Westwood). History Museum, London), Paul Marsh (USNM), Chris
' O'Toole (Hope Entomological Collections, Oxford), L.Stange
(Florida State Collection of Arthropods) and David Wahl
(American Entomological Institute, Gainesville) for allowing
KEY TO THE SPECIES OF MYOSOMATOIDES me access to specimens in their care. This work was partly
supported by the American Entomological Institute, a scien-
1 Wings uniformly brown tific exchange visit to Budapest funded by the Royal Society,
M. muer si nom. nov. dr,d by a research grant jointly to DLJQ and Mike Fitton
(BMNH) from the NERC.
— Wings largely hyaline with the apical third of the
forewing brown (Fig. 1) LITERATURE CITED
M. pennipes (Westwood)
Achterberg, C. van, 1979. A revision of the subfamily Zelinae
auct. (Hymenoptera, Braconidae). Tijdschrift voor
Entomologie 122: 241-479.
Achterberg, C. van, 1988. Revision of the subfamily Blacinae
Foerster (Hymenoptera, Braconidae). Zoologische
Verliandelmgen, Leiden 249: 1-324.
Agnew, C.W., Rodriguez-del-Bosque, L. A., Smith, J. W. Jr.
1988. Misidentification of Mexican stalkborers in the
subfamily Crambinae (Lepidoptera: Pyralidae). Folia
Entomologica Mexicana 75: 63-75.
Chippendale, G.M. 1979. The southwestern corn borer,
Diatraea grandiosella: case history of an invading insect.
Bulletin of the Missouri Agricultural Experiment Station,
no. 1031, 1-52.
De Santis, L., Esquivel, L. 1966. Tercera lista de himenopteros
parasiticos y predatores de los insectos de la Republica
Argentina. RevistadelMuseo de La P/<i(<i(Secci6nZoologia
N.S.) 9: 47-215.
Elias, L.A. 1970. Maize resistance to stalk borers in Zeadiatraea
Box and Diatraea Guilding at five localities in Mexico. 172
pp. Ph.D. dissertation, Kansas State University (Univ.
Microfilms no. 70-16 627).
Maeto, K. 3992. Japanese species of the Myosoma group (Hy-
menoptera? Braconidae). Japanese Journal of Entomology
60: 714.
Mason, W. R. M. 1978. A synopsis of the Nearctic Braconini,
with revisions of Nearctic species of Coeloides and
Myosoma (Hymenoptera: Braconidae). Canadian Ento-
mologist 110: 721 -768.
Muesebeck, C. F. W. 1925. A revision of the parasitic wasps of
the genus Microbracon occurring in America north of
Mexico. Proceedings of the United States National Museum
67: no. 2580: 1-85."
Myers, J. G. 1931. Descriptions and records of parasitic Hy-
menoptera from British Guiana and the West Indies.
Bulletin of Entomological Research 22: 267-277.
DISCUSSION
Myosomatoides gen. nov. most closely re-
sembles and is probably closely related to the
small and widespread, but principally New World
genus Myosoma Brulle. Individuals of both genera
have the hind femur rather strongly laterally com-
pressed and densely clothed with setae (Mason
1978): Myosomatoides differs from Myosoma in hav-
ing an even more strongly laterally compressed
hind femur and tibia (apomorphous) (Figs. 1, 2, 4),
and in having the 2nd metasomal tergite 'pinched-
up' in the middle so as to form a more or less
complete midlongitudinal ridge (apomorphous)
(Fig. 4). In Myosoma, the 2nd tergite totally lacks
carinae, the 1st metasomal tergite is longer and
narrower than in Myosomatoides (apomorphous)
and has the flattened lateral areas absent or nearly
so (apomorphous) (Mason 1978, Quicke & Wharton
1989), and often the anterolateral part of the 2nd
metasomal tergite is only weakly sclerotized
(apomorphous). Myosomatoides also resembles
Myosoma in its complement of intertergal glands
(Quicke 1990) and in the form of the male genitalia
(Quicke 1988c), though in Myosoma there is only
one tooth on the digitus (apomorphous). Myosoma
and Myosomatoides appear therefore to be sister
groups though at present the possibility that the
Volume 3. 1 994
231
Myers, J. G. 1932. Biological observations on some Neotropi-
cal parasitic Hymenoptera. Transactions of the Royal En-
tomological Society of London 80: 121-136.
Parker, H. L., Berry P. A. Silveira Guido A. 1953. Host-
parasite and parasite-host lists of insects reared in the
South American Parasite Laboratory during the period
1940-1946. Revistade la AsociaciondelngenieraAgronomica,
Montevideo, no. 92: 1-101.
Pashley, D. P., Hardy T„ Hammond A. M, Mihm J A. 1990.
Genetic evidence for sibling species within the Sugar-
cane Borer (Lepidoptera: Pyralidae). Annals of the Ento-
mological Society of America 83: 1048-1053.
Quicke, D. L. J. 1985. Two new genera of Braconinae (Insecta,
Hymenoptera) from the Afrotropical Region. Zoologica
ScriptaU: 117-122.
Quicke, D.L.J. 1987. The Old World genera of braconine
wasps (Hymenoptera: Braconidae). Journal of Natural
History 21: 43-157.
Quicke, D. L. J. 1988a. A new genus and species of Braconinae
(Hymenoptera: Braconidae) parasitic on Diatraca (Lepi-
doptera: Pyralidae) in Ecuador. Bulletin of Entomological
Research 78: 15-18.
Quicke, D. L. J. 1988b. Digonogastra: the correct name for
Nearctic Iphiaulax of Authors. Proceedings of the Entomo-
logical Society of Washington 90: 196-200.
Quicke, D. L. J. 1988c. Inter-generic variation in the male
genitalia of the Braconinae (Insecta, Hymenoptera,
Braconidae). Zoologica Scrinta 17: 399-409.
Quicke, D. L. J. 1989. Further new host records for genera and
species of Braconinae (Hym., Braconidae). En tomologist's
Monthly Magazine 125: 199-205.
Quicke, D. L. ]. 1990 Tergal and inter-tergal metasomal
glands of ma le braconine wasps (Insecta, Hymenoptera,
Braconidae). Zoologica Scripta 19: 413-423.
Quicke, D. L. J. Sharkey M. J. 1989. A key to and notes on the
genera of Braconinae Hymenoptera: Braconidae) from
America north of Mexico with descriptions of two new
genera and three new species. Canadian Entomologist
121:337-361.
Quicke, D. L. J., Wharton R. A. 1989. Myosoma nyanzaensis
sp.n. (Hymenoptera: Braconidae) parasitic on the stem-
borer pest Chiio partellus (Swinhoe) (Lepidoptera:
Pyralidae) in Kenya. Bulletin of Entomological Research
79: 1-5.
Shenefelt, R. D. 1978. Hytnenopterorum Catalogus (nov. ed.).
Part 15. Braconidae, vol. 10. The Hague, Junk, pp. 1425-
1865.
Thomson, W. R. 1953. A catalogue of the parasites and predators
of insect pests. Section 2, host parasite catalogue. Part 2,
hosts of the Hymenoptera (Agaonidae to Braconidae).
Commonwealth Institute of Biological Control, Ottawa,
190 pp.
Westwood, J. O. 1882. Descriptions of new or imperfectly
known - Ichneumones adsciti. Tijdschrift voor Entomologie
25: 17-48.
Wharton, R. A., Smith J. W. Jr, Quicke D. L. J., Browning H. W.
1989. Two new species of Digonogastra Viereck (Hy-
menoptera: Braconidae) parasitic on Neotropical pyralid
borers (Lepidoptera) in maize, sorghum and sugarcane.
Bulletin of Entomological Research 79: 401 -410.
J. HYM. RES.
Vol. 3, 1994, pp. 233-239
Immature Stages of Aganaspis pelleranoi (Brethes)
(Hymenoptera: Cynipoidea: Eucoilidae),
a Parasitoid of Ceratitis capitata (Wied.)
and Anastrepha Spp. (Diptera: Tephritidae).
Sergio M. Ovruski
Centre de Investigaciones para la Regulation de Poblaciones de Organismos Nocivos (CIRPON), Pasa|e Caseros 1050,
C.C.C. 90, S. M. de Tucuman (4.000), Argentina.
Abstract- — The embryological-larval development of Aganaspis pelleranoi (Brethes), a larval parasitoid of Ceratitis capitata
(Wied.) and Anastrepha spp. is described and illustrated. A. pelleranoi possesses a stalked egg, eucoiliform first- and second-
instar larvae, and hymenoptenform third- and fourth-instar larvae, followed by a prepupal and pupal stage. The duration of
each stage at 25-26°C was as follows: egg, 3-4 days; first-, second-, and third-instar larvae, 2-3 days each; fourth-instar larva,
3-4 days, prepupa, 1-2 days; pupa, 9-14 days.
Aganaspis pelleranoi (Brethes) is a parasitoid of
Ceratitis capitata (Wied.) and several Anastrepha
species (Diptera: Tephritidae) in Argentina (De
Santis 1965), that attacks the final larval instar of
these fruit flies (Turica and Mallo 1961). Aganaspis
pelleranoi has also been reported from Lonchaea
spp. (Diptera: Lonchaeidae) in Brazil (De Santis
1980).
From 1941 to 1945, A. pelleranoi was released
as a control agent of fruit flies in Tucuman prov-
ince, Argentina (Nasca 1973). In 1942 this species
was introduced and reared in Peru for the control
of A. fratercuhts (Wied.) (Clausen 1978).
Ovruski (in press) carried out studies on host
detection behaviour in this fruit fly parasitoid.
Otherwise little is known of the biology of A.
pelleranoi, and descriptions of the immature stages
of this species have not been published. In fact
very little at all is published on the development of
eucoilid parasitoids of fruit flies, except that
Clausen et al. (1965) briefly described the imma-
ture stages of A. daci (Weld) (=Trybliograplia daci
Weld), a parasitoid of Dacits dorsalis Hendel, the
oriental fruit fly.
In this paper, studies on the life cycle and pre-
imaginal development of A. pelleranoi are pre-
sented.
MATERIALS AND METHODS
Aganaspis pelleranoi was reared on last instar
larvae of C. capitata in the laboratory as described
by Ovruski (in press). The host larvae were reared
on carrot/corn meal/yeast medium as described
by Nasca (1977).
The life cycle and immature stages of A.
pelleranoi were studied by exposing 20 to 30 C.
capitata larvae to individual parasitoid females.
The rearings and experimentation were conducted
at 25-26°C, 70-80% RH, and 12:12 LD photoperiod.
Host larvae and host puparia were removed at
intervals of 24 h after parasitism and dissected in
Insect Ringer's solution on depression slides. The
parasitoid larvae were removed from each host
puparium and preserved in 70% ethanol for later
examination by light microscopy. Eight fourth-
instar larvae were refixed in 4% glutaraldehyde
and 0.2 M phosphate buffer, and dehydrated in 35-
50-70-90-100% ethanol. Thereafter they were placed
in 100% acetone. These specimens were critical
point dried before examination by scanning elec-
tron microscopy (SEM). Descriptions are based on
several specimens. Thirty-four eggs in different
stages of development were mounted on slides
and examined by light microscopy. Six first-, five
234
Journal of Hymenoptera Research
second-, and five third-instar larvae were mounted
and examined by light microscopy. Eight fourth-
instar larvae were observed with SEM and fifteen
fourth-instar larvae were examined by light mi-
croscopy. Six females of A. pelleranoi of different
ages were dissected in Insect Ringer's solution to
observe the ovarian mature eggs. The terms used
for morphological structure follow Kopelman and
Chabora (1984) and Evans (1987). The drawings
were made with a camera lucida. Data are pre-
sented as range and /or mean. Specimens, in etha-
nol and mounted, are deposited in the Miguel
Lillo Institute Fundation, S. M. de Tucuman Ar-
gentina.
LIFE CYCLE
Females of A. pelleranoi deposited a single egg
in the body cavity of the host larva and one para-
sitoid adult emerged from each host puparium.
Under laboratory conditions, the period from
oviposition in the host larva to the emergence of a
parasitoid adult varied from 25 to 27 days in males
and from 26 to 30 days in females. Thus, males
tended to emerge somewhat earlier than females.
The egg stage inside the host larva lasted about 3-
4 days. Observations of the postembryological
development showed the presence of four larval
instars. The first, second, and third instar each
lasted about 2-3 days and the fourth instar 3-4 days
(9-13 days in total). The prepupal stage lasted 1-2
days, and the pupal stage from 9 to 14 days.
EMBRYOLOGICAL DEVELOPMENT
The newly laid egg is 0.32 mm-0.56 mm long
and 0.06 mm - 0.10 mm wide; the stalk is 0.42 mm
- 0.73 mm long and the embryo is not yet distin-
guishable (Fig. 1).
24 h after oviposition inside the host larva, the
parasitoid egg is 0.26 mm-0.48 mm long and 0.08
mm - 0.15 mm wide. The stalk is reduced in length,
and the embryo, vitelline membrane and the
chorion are well discernible (Fig. 2 ).
By 48 h after oviposition, the egg has length-
ened and swelled (0.52 mm-0.66 mm long and 0.24
mm - 0.26 mm wide), and the stalk has decreased
still more (Fig. 3 ). The embryo shows signs of
body segmentation, a caudal portion and thoracic
appendages.
After 72 h the parasitoid larva is completely
formed. The stalk disappears (Fig. 4 ) and the size
of the egg is similar to the 48 h old egg. Eclosion of
the first-instar larva occurs after 78-80 h. The
chorion is broken through anteriorly by the man-
dibles and posteriorly by movements of the tail
(Fig. 5 ).
LARVAL DEVELOPMENT
After hatching, the first-instar larva actively
feeds on the internal tissues of the host. For this
purpose, the mandibles are protruded from the
oral cavity, although it was not possible to verify
their use in feeding by direct observation. During
this stage, the tail becomes less prominent and the
head wider.
The second-instar larva appears on the fifth to
sixth day after oviposition, when the host pu-
parium is formed. The larva continues to feeds
inside the host pupa.
The third-instar larva appears on the eigth to
ninth day after oviposition. In this stage the para-
sitoid larva partially emerges from the host pupa
near the middle region, and it then feeds exter-
nally within the host puparium. The parasitoid
larva occupies both internal and external posi-
tions in the host pupa and develops two respira-
tory mechanisms: cuticular respiration through
submerged posterior parts of the body and tra-
cheal respiration through prothoracic spiracles.
In the fourth-instar larva, which appears on
approximately the tenth day after oviposition,
feeding continues externally on what remains of
the host pupa. The parasitoid larva is very slug-
gish and occupies 3/4 of the host puparium. After
feeding is completed, the meconium is released. It
appears as a dark crust in the posterior apex of the
puparium (Fig. 14 ).
DESCRIPTIONS OF IMMATURE STAGES
Mature ovarian egg. — (Fig. 1.) Stalked; total
length on average 0.60 mm; stalk 1 .2 X longer than
egg body, the latter portion 3.5 X longer than wide;
chorion translucent and smooth.
First-instar larva. — (Fig. 6, 7). Length 0.93 mm
- 1.39 mm; 2.2 X longer than wide; eucoiliform;
translucent; subcylindrical in shape with elongate
distinct head; mouth surrounded by several oral
papillae, inside with a pair of unidentate
Volume 3. 1 994
1
235
I
Figs. 1-7. Aganaspis pelleranoi. 1, ovarian egg. St., stalk. Scale bar = 0,12 mm. 2, egg at 24 h. after oviposition. ch., chorion; V.M.,
vitelline membrane; Em., embryo. Scale bar = 0,12 mm. 3, egg at 48 h. after oviposition. Scale bar = 0,12 mm. 4, egg at 72 h. after
oviposition. Scale bar = 0,12 mm. 5, egg at 80 h. after oviposition. Scale bar = 0,12 mm. 6, first-instar larva. OP., oral papillae;
Th. App., thoracic appendages; V.P., ventral process; 77., tail; An., anus. Scale bar = 0,12 mm. 7, mandibles of first- instar larva.
Scale bar = 0,02 mm.
236
Journal of Hymenoptera Research
n.
Ml.
G. F.
Figs. 8-9. Aganaspis pelleranoi. 8, second-instar larva. OP., oral papillae; 77., tail. Scale bar = 0,25 mm. 9, third-instar larva. Sp.
spiracle; Mt., mouth; C.F., globular fat; An., anus. Scale bar = 0,32 mm.
237
Figs. 10-12. Aganaspis pelleranoi. 10, fourth-instar larva. Lat. Sw., lateral swelling; An. , anus. Scale bar = 1,0 mm. 1 1 , anterior view
of head, fourth-instar larva. Ant. Or., antennal orbit; Ma., mandible; Mi., maxilla; La., labium; Lb , labrum. Scale bar = 0,10 mm
12, lateral view of body segment 6-8, fourth-instar larva. Sp , spiracle; Lat Sw , lateral swelling. Scale bar = 0,10 mm.
subtriangular mandibles (Fig. 7 ); ventral surface
of head with a prominent tubular projection; gut
easily discernible; each thoracic segment with a
pair of long slender ventral appendages; seven
well-defined abdominal segments; caudal segment
with a long tail and short ventral process; tail
bearing several small setae apically; ventral mar-
gin of 7th abdominal segment and basal end of
caudal segment with scalelike ornamentation; tail
6 X longer than ventral process; anus opening
dorsally.
Second-instar lari'a. — (Fig. 8). Length 1.53 mm
- 2.31 mm; 2 X longer than wide; modified
eucoiliform; whitish yellow with some white
globular fat particles; cylindrical in shape with
short and fleshy head; mouth with external oral
papillae; unidentate mandibles; gut well discern-
ible; without thoracic appendages; caudal seg-
ment with very short tail and without ventral
process; anus opening dorsally.
Third-instar larva. — Fig. 9. Length 2.78 mm-
3.45 mm; 3 X longer than wide; more typically
hymenopteriform; yellowish with several white
globular fat particles in thorax and abdomen; cy-
lindrical in shape with slender posterior portion;
broad head; small unidentate mandibles, difficult
to discern; prothoracic segment with a single pair
of spiracles; ten distinguishable body segments;
brain and gut easily discernible; without tail and
ventral process; anus opening ventrally.
Fourth-instar larva. — Figs. 10, 11,12, 13). Length
3.48 mm-4.13 mm; 2 X longer than wide;
hymenopteriform; whitish yellow with many small
white globular fat particles dispersed throughout
dorsolateral parts of thorax and abdomen; head
23S
Journal of Hymenoptera Research
13
Me.
Figs. 13-15. Aganaspis pelleranoi. 13, mandibles of fourth-instar larva. Scale bar = 0,04 mm. 14, prepupa inside the host
pupanum. Ph., pupanum; Pr., prepupa; Me , meconium. Scale bar = 1,0 mm. 15, pupa. Scale bar = 1,6 mm.
relatively large, subcircular in front view, without
setae; antennal orbits large; prominent bidentate
mandibles, 1.2 X broader than long basally, very
dark apically (figs. 1 1 and 13); labrum 3.2 X broader
than long, with rounded lateral sides and median
depression on apical margin; maxillae circular in
shape and joined to labium; body very swollen;
only eleven body segments discernible; except
prothoracic and terminal abdominal segment, all
segments with a pair of spiracles; each body seg-
ment provided with lateral voluminous swellings
behind spiracles (Fig. 12); integument smooth and
without setae; anus opening ventrally.
Prepupa.— (Fig. 14). Length 2.55 mm - 2.98
mm; width 1.18 mm - 1.56 mm; similar to fourth-
instar larva; body curved and reduced in size;
occupying 2/3 of the host puparium.
Pupa.— (Fig. 15). Length 2.68 mm - 2.99 mm,
initially white, later becoming dark; antennae 1,7
mm - 2.1 mm in length, extending to first abdomi-
nal segment.
DISCUSSION
Aganaspis pelleranoi produces a stalked egg,
typical of the Eucoilidae. The transition from en-
doparasitic to ectoparasitic life has also been re-
ported for other species of Eucoilidae (Jenni 1951 ;
Wishart and Monteith 1954; Sychevskaya 1974;
Kopelman and Chabora 1984). The embryological
Volume 3, 1994
239
and larval development is similar to other eucoilid
parasitoids: Tn/bliograplia rapae (Westwood) (James
1928; Wishart and Monteith 1954); Hexacola sp.
(Simmonds 1952); Hexacola sp. near Websteri
(Crawford) (Eskafi and Legner 1974); Leptopilina
boulardi (Barbotin et al.) (Kopelman and Chabora
1984); Eucoila frichopsila (Hartig) (Sychevskaya
1974) and Kleidotoma japonica Huz. (Huzimatu
1940).
Clausen et al. (1965) reported three larval in-
stars in A. daci . This study proves the presence of
four larval instars in A. pelleranoi. The first-, sec-
ond-, and fourth-instar larvae described here are
similar to the first-, second-, and supposed third-
instar larvae of A. daci respectively. The third-
instar larvae of A. pelleranoi is a transition between
the eucoiliform type and the hymenopteriform
type. This stage is distinguished by a single pair of
open spiracles, in the prothoracic segment, as in
the supposed third stage of T. rapae (Wishart and
Monteith 1954), the supposed fourth stage of £.
frichopsila (Sychevskaya 1974) and the supposed
fourth stage of L. boulardi (Kopelman and Chabora
1984). Further comparative work would be needed
to confirm these apparent differences between
species, as intermediate instars of apocritan larvae
are notoriously easy to overlook.
Furthermore Clausen et al. (1965) stated that
the first-instar larva of A. daci does not possesses
discernible mandibles, but the first-instar larvae of
A . pelleranoi has a pair of well developed unidentate
mandibles with their ends projecting from the oral
cavity, similar to the first-instar larvae of £.
trichopsila (Sychevskaya 1974).
Prominent bidentate mandibles in last instar
larvae have also been reported for A. daci (Clausen
et al. 1965), Trybliograplia Forster and Hexacola
Fbrster (Evans 1987).
ACKNOWLEDGMENTS
I am especially grateful to Dr. Patricio Fidalgo (CIRPON,
Argentina), Dr. Abraham Willink (CONICET-UNT, Argen-
tina), and Dr. Goran Nordlander (Swedish University of
Agricultural Sciences, Uppsala, Sweden) for suggestions and
critical reviews of the manuscript
LITERATURE CITED
Clausen, C. P. 1978. Tephritidae, pp. 320-334. In Clausen, C.
P., ed. Introduced Parasites and Predators of arthropod
pests and weeds: a World Review. U.S. Agriculture Hand
Book, N° 480. 545 pp.
Clausen, C. P., D. W. Clancy and Q. C. Chock. 1965. Biological
control of the oriental fruit fly {Dacus dorsalis Hendel)
and the other fruit flies in Hawaii. Technical Bulletin
United States Department of Agriculture, N" 1322. 102 pp.
De Santis, L. 1965. Nota sobre un parasito de la mosca
sudamencana de la fruta (Hymenoptera: Cynipidae).
Rei'ista de la Soaedad Entomologica Argentina 27 (1-4): 73-
74.
De Santis, L. 1980. Catalogo de los himenopteros brasilenos
de la Sene Parasitica, incluyendo Bethyloidea.
Universidad Federal do Parana, Curitiba. 395 pp.
Eskafi, F. M. and E. F. Legner. 1974. Descriptions of immature
stages of the cynipid Hexacola sp. near Websteri
(Eucoilinae: Hymenoptera), a larval-pupal parasite of
Hippelates eye gnats (Diptera: Chloropidae). The Cana-
dian Entomologist 106: 1043-1048.
Evans, H. E. 1987. Cynipoidea. Cynipids, gall wasp and
others, pp. 665-66. In Stehr, F. W., ed. Immature insects.
Kendall Hunt, Dubuque, Iowa. 754 pp.
Huzimatu, K. 1940. The life history of a new cynipid fly,
Kleidotoma japonica, n. sp. Science Report, Toboku Imperial
University, Sendai 15: 457-480.
James, H. C. 1928. On the life histories and economic status of
certain parasites of dipterous larvae, with descriptions
of some new larval forms. Annals of Applied Biology 15:
287-316.
Jenni, W. 1951. Beitrag zur Morphologie und Biologie der
Cynipide Pseudeucoila bocliei Weld, eines larvenparasiten
von Drosophila melanogaster Meig. Acta Zoologica 32: 177-
254.
Kopelman, A. H. and P. C. Chabora. 1984. Immature stages of
Leptopilina boulardi (Hymenoptera: Eucoilidae), a
protelean parasite of Drosophila spp. (Diptera:
Drosophilidae). Annals of the Entomological Society of
America 77(3): 264-269.
Nasca, J. A. 1973. Par^sitos de "moscas de los frutos"
establecidos en algunas zonas de Tucum^n. Revista
Agricola del Noroeste Argentino 10 (1-2): 31-43.
Nasca, J. A. 1977. Estudios relacionados con las posibilidades
del Control Integrado de las moscas de las frutas.
Universidad Nacwnal de Tucumdn, Informe CAFPTA, N"
529. 61 pp.
Ovruski, S. M. In press. Comportamiento en la deteccion del
huesped en Aganaspis pelleranoi (Brethes) (Hymenoptera:
Cynipoidea, Eucoilidae) parasitoidedelarvasde Cera fitis
capitata (Wied) (Diptera: Tephritidae). Revista de la
Sociedad Entomologica Argentina 53.
Simmonds, F. J. 1952. Parasites of the frit-fly, Oscinellafrit (L.),
in eastern North America. Bulletin Entomological Re-
search 43: 503-542.
Sychevskaya, V. L. 1974. The biology of Eucoila trichopsila
Hartig (Hymenoptera, Cynipoidea), a parasite of the
larvae of synanthropic flies of the family Sarcophagidae
(Diptera). Entomological Review 53: 36-44.
Turica, A. and R. G. Mallo. 1961. Observaciones sobre la
poblacion de las "Tephritidae" y sus endoparasitoides
en algunas regiones citricolas argentinas. IDIA 6: 145-
161.
Wishart, G. and E. Monteih. 1954. Trybliograplia rapae (Westw.)
(Hymenoptera: Cynipidae), a parasite of Hylemya spp.
(Diptera: Anthomiidae). The Canadian Entomologist 86:
145-154.
J. HYM. RES.
Vol. 3, 1994, pp. 241-277
Phylogenetic Implications of the Mesofurca
and Mesopostnotum in Hymenoptera
John M. Heraty, James B. Woolley and D. Christopher Darling
(JMH) Biological Resources division, CLBRR, Agriculture Canada, C.E.F., Ottawa, Ontario, Canada Kl A 0C6
(present address: Department of Entomology, University of California, Riverside, California, USA 92521);
(JBW) Department of Entomology, Texas A&M University, College Station, Texas, USA 77843;
(DCD) Department of Entomology, Royal Ontario Museum, Toronto, Ontario, Canada M5S 2C6
Abstract. — The skeleto-musculature of the mesofurca and the mesopostnotum is examined in Hymenoptera. Character
systems based on internal structure of the mesothorax support recent hypotheses that suggest sawflies are paraphyletic with
respect to Apocrita. Unique character states for Hymenoptera include the presence of two mesofurcal-laterophragmal
muscles, a mesofurcal-third basalare muscle, and a scutellar-metanotal muscle. Other possible apomorphies include the
medial emargination of the mesopostnotum and the formation of anterior furcal arms. The arrangement of mesofurcal muscles
that attach to the profurca and the laterophragma are described and interpreted in light of recent phylogenetic hypotheses.
Changes in attachment sites, fusion or loss of the anterior arms of the mesofurca and features of the laterophragma provide
characters that are consistent with the monophyly of Tenthredinoidea + (Cephoidea + (Siricoidea (including Anaxyelidae) +
(Xiphydriidae + Orussoidea + Apocrita))). Groundplan states for the Apocrita are proposed that include retention of a
mesofurcal bridge, retention of an anterior process on the bridge that supports the interfurcal muscles, reduction of the
mesofurcal-laterophragmal muscles from two to one, retention of the mesotergal-laterophragmal muscle, loss of the
mesofurcal-third basalare muscle, and loss of the metafurcal-spina muscle. Within Apocrita the distribution of character state
changes is less informative than in Symphyta, but provide evidence for relationships of some taxa. The mesofurcal bridge is
lost convergently in Ceraphronoidea, Pelecinidae, Platygastroidea, Mymarommatoidea, Mymandae and some Chalcidoidea.
The tergal-laterophragmal muscle and associated posterior lobe of the laterophragma are postulated to have been lost
independently in nine lineages of Apocrita. The development of the laterophragma into an axillary lever is a synapomorphy
for Vespoidea and Apoidea, and in Apiformes the lever is an independent sclerite. The distribution of states for 12 characters
is discussed for 62 families of Hymenoptera. Parsimony analysis of these data result in trees that generally agree with the
current hypotheses for Symphyta but not for Apocrita.
INTRODUCTION 1969, 1980,Matsuda 1970,Shcherbakov 1980, 1981,
Gibson 1985, Johnson 1988, Whitfield et al. 1989).
"Students of these [hymenopteran] parasites discover that The determination of homologous structures and
the thorax presents valuable characters for the determination polarity of characters in Hymenoptera are crucial
and classification of species, but they are handicapped by the r > _■• ,, .• i .• i ■
lack of reliable studies on the structure of the thorax" for understanding phylogenetic relationships, a
Snodgrass 1910 p 37 toPlc recently addressed by various authors
(Rasnitsyn 1969, 1980, 1988, Brothers 1975,
Since Snodgrass (1910) first attempted to ex- Konigsmann 1977, 1978a 1978b, Carpenter 1986,
pand our knowledge of the structure of the hy- Brothers ^d Carpenter 1993).
menopteran thorax, additional studies have de- The mesofurca is an invagination of the ster-
scribed the skeleto-musculature of single species num mto the thorax that forms a central Point of
or single families (Weber 1925, 1927, Tulloch 1935, attachment f°r the ventral longitudinal muscles,
Maki 1938, Duncan 1939, Michener 1944, Bucher the sterno-pleural muscles, the coxal and trochant-
1948, Alam 1951, Saini et al. 1982, Daly 1964, eral muscles' and the mesopostnotal muscles
Gibson 1986, 1993). Fewer studies have compared (KelseY 1957< Matsuda 1970). The mesofurca is
thoracic structures among families of Hy- comPnsed of a basal plate (discrimenal lamella)
menoptera (Snodgrass 1942, Daly 1963, Rasnitsyn that rises vertically f™ the discrimen, slopes
posteriorly to the furcal base, and divides dorsally
242
Journal of Hymenoptera Research
into two lateral arms, termed the sternal apophy-
ses (Snodgrass 1927, Chapman 1992, Lawrence et
al. 1992). The mesopostnotum is one of the pri-
mary dorsal sclerites involved in flight through
the posterior inflection of the antecosta (second
phragma) which forms the posterior attachment
of the longitudinal flight muscles. Recent studies
involving skeleto-musculature of the hy-
menopteran thorax have focused on the pleural
attachments (Shcherbakov 1980, 1981, Gibson 1985,
1993), the extrinsic musculature of the mesocoxa
(Johnson 1988), and the development of the
metapostnotum (Whitfield et al. 1992). Rasnitsyn
(1969, figs. 187-194) was the first person to com-
pare the different skeletal structures for the
mesofurca of 8 families of Symphyta. His illustra-
tions show the transformation series for Symphyta
that are discussed in this paper. Rasnitsyn (1988)
refers to the furca for features supporting
Tenthredinoidea and for Cephoidea + Siricoidea +
Apocrita. Similarly, Snodgrass (1942) presented a
pictorial evolutionary history for development of
the axillary lever of Apoidea. This work expands
upon these initial studies and extends the com-
parative aspects of these works to include most
families of Apocrita.
This study of the mesofurca and
mesopostnotum began as an attempt to under-
stand the polarity and homology of mesofurcal
structures and muscles of Aphelinidae
(Chalcidoidea) and the phylogenetic implications
of these attributes within the Chalcidoidea. Even-
tually the entire Hymenoptera needed to be sur-
veyed to resolve what we initially thought were
relatively simple questions. In this study, all
muscles attaching to the mesofurca and
mesopostnotum are identified and compared to
homologous muscle groups in Neuropterida and
Mecopterida (sensu Kristensen 1992), as they are
considered to be phylogenetically close to Hy-
menoptera (Kristensen 1992), and have a mesotho-
rax which is structured similar to the Symphyta.
Within Hymenoptera, we have concentrated our
analysis on the skeletal structure of the mesofurca
and mesopostnotum, and on the muscles attach-
ing between the thoracic furcae and the
laterophragma of the mesopostnotum. The evi-
dence provided by the mesofurca and
mesopostnotum for relationships within the
Chalcidoidea will be discussed in a subsequent
paper.
MATERIA! S AND METHODS
Terms for structures and muscles generally
follow Snodgrass (1910, 1942), Daly (1963), Gibson
(1985, 1986, 1993) and Ronquist and Nordlander
(1989). Muscles were identified using the systems
proposed by Kelsey (1957) and Daly (1963) (Table
1). Figures 1 and 2 are used to place the skeleto-
musculature within the context of the mesosoma.
Muscles and stuctures are extensively labeled in
Figs. 3 and 4. The Kelsey system uses a fixed set of
numbers and is useful for comparisons across the
Endopterygota. Daly's system is preferred for clar-
ity because the insertion-origin of attachment sites
are readily identified and new muscles can be
added to the system; for example, the new muscle
fu,-ba3 was given the abbreviation fbl for the
Kelsey system as it could not be assigned a nu-
meric value that would signify its relative position
to other muscles in the mesothorax. Terms pro-
posed by Matsuda (1970) are comprehensive and
may be referenced across orders of insects; how-
ever his abbreviated system is difficult to use and
is not followed here.
Several new classifications of families within
Hymenoptera have been proposed recently that
differ largely in placement of certain families as
separate superfamilies, families, or subfamilies.
We follow the classification of Huber and Goulet
(1993), as it represents the most current synthesis
of information across the order.
Dissections were based on specimens pre-
served in 70%ethanol or initially fixed in Dietrich's
or Kahle's solution and then transferred to etha-
nol. All specimens were critical point-dried prior
to dissection. The mesosoma of Monomachns
(Monomachidae) was rehydrated using Barber's
solution, transferred through increasing concen-
trations of ethanol to 98% and then critical point-
dried. For each dissection, the mesosoma was
anchored onto a standard SEM stub using chloro-
form-based silver paint. Dissections were made
using hooked minuten pins or fragments of razor
blades. Dried haemolymph and extraneous tis-
sues were removed from dissections using small
amounts of glue obtained by dragging a hooked
minuten pin across clear sticky tape (Gibson 1985).
Exemplar taxa were chosen to represent the
maximum variation within taxa. In some groups
(e.g. Apoidea), there was virtually no variation;
whereas within some taxa (e.g. Diapriidae) both
Volume 3. 1994
243
structure and presence of muscles varied and more
genera were dissected to characterize this varia-
tion. Our primary concern was for establishing
groundplan states for higher taxa, although
autapomorphies are discussed. The taxa exam-
ined for internal characters are listed in Table 2.
Numerous Chalcidoidea were also dissected as
part of a comprehensive study of the mesofurca in
that superfamily. Representative dissections are
housed at the Canadian National Collection
(CNCI), Royal Ontario Museum (ROM) and Texas
A&M University (TAMU). When possible, con-
specific adults of the dissected specimens are de-
posited as voucher specimens in the above collec-
tions. The majority of specimens were obtained
from the CNCI alcohol collection.
The mesofurcal-mesopostnotal complex was
broken up into 12 characters with a total of 36
character states. Seventy hymenopteran taxa were
scored (Appendix 2) based on the examination of
internal characters for 119 species (Table 2). A
single outgroup taxon was scored based on dissec-
tions of 5 families of Neuropterida and 3 families
of Mecopterida. Characters 1, 3 and 9 are postu-
lated as unique characters for Hymenoptera; in
each of these cases, the outgroup is coded as a
unique character state (state 0). The state value "?"
was used to denote uncertain homology, not miss-
ing data.
Illustrations were made with a camera lucida.
Outlines of muscles present but not illustrated are
represented by dashed lines. Some muscles not
central to this study were not consistently figured
(e.g. muscle 180 for Symphyta) and caution should
be exercised in deriving additional interpretation
from the illustrations. Abbreviations referring to
muscles are circled in all figures, skeletal charac-
ters are not. The mesofurcal-mesopostnotal com-
plex is abbreviated as MF-MPN complex.
RESULTS AND DISCUSSION
Character Analysis
Character 1. Mesofurcal bridge
The most significant modification of the
mesofurca in Hymenoptera is the fusion of the
elongate anterior arms (af) into a mesofurcal bridge
(fb) that characterizes virtually all Apocrita. In
basal groups of Symphyta (excluding Cephoidea
+ remaining Hymenoptera), the anterior furcal
arms are either absent, short, or elongate and well
separated along their entire length. The anterior
arms, or the equivalent region on the lateral arms
of the mesofurca, form the posterior attachment
sites for the ventral intersegmental muscles (muscle
124, fiij-fUj, and muscle 127, fu-rsps,). With fusion
of the anterior arms, the furca separates the gastric
and nervous systems and the ventral nerve cord
passes through the foramen bounded by the
mesofurcal bridge and the lateral arms of the
mesofurca.
In most Neuropterida and Mecopterida, the
interfurcal muscles attach directly to the anterior
face of the lateral arms of the mesofurca (state 0, no
arms) (Snodgrass 1927, Kelsey 1957, Matsuda 1970).
In Xyelidae (Fig. 3) and Pamphiliidae (Fig. 6) (and
probably Megalodontidae, cf. fig. 187, Rasnitsyn
1969), the interfurcal muscles attach to anterior
projections (af, anterior furcal arms) that are long,
robust, and separated along their entire length
(state 1). Of the outgroup taxa examined, only
Brachynemurus (Myrmeleontidae) have anterior
arms (state 1) similar to Xyelidae, and thus their
presence could be plesiomorphic or apomorphic
for Hymenoptera. Because anterior arms are
present only in a derived member of the outgroup,
we consider the presence of long anterior arms in
Xyeloidea and Megalodontoidea to be apomorphic
for Hymenoptera.
In Tenthredinoidea, the anterior arms (af) can
be reduced in size (Cimbicidae Fig. 8,
Blasticotomidae Fig. 20, and Nematus
(Tenthredinidae) Fig. 21), modified into support-
ing cup-like structures (Diprionidae, Fig. 7), or lost
entirely so that the interfurcal muscles attach to
the anterior face of the lateral furcal arms ( Argidae,
Pergidae Fig. 5, and most Tenthredinidae).
Rasnitsyn (1988) treated "fore arm short" as a
character state (his 2-3d) supporting
Tenthredinoidea and "fore arm reduced" (his 6-
8a) for Argidae + Pergidae. Short arms, as in
Blasticotomidae (Fig. 20), are probably
plesiomorphic for Tenthredinoidea, and further
modifications of the arms or complete loss are
derived within Tenthredinidae. It is possible to
code for several different character states within
Tenthredinoidea (small, absent, cup-shaped, etc.),
but this would only introduce unnecessary ho-
moplasy into the analysis (i.e. "absence" derived 2
or more times) or a series of autapomorphic char-
244
Journal of Hymenoptera Research
Table 1. Homology and terms for muscles of the mesopostnotum and mesofurca examined in Neuroptera and Hymenoptera
Letter m added to distinguish metathoracic muscles. Muscle fbl described in text.
Kelsey 1957 — Con/dalus
No Name
Hymenoptera
Matsuda Daly Johnson Present
1970 1964 1988 designation
Notes
Mesothoracic dorsal indirect muscles
112 internal (ventral] longitudinal tl4 lph-2ph
112m metathoracic longitudinal tl4 2ph-3ph
114 scutellar-metanotal tl3 t,-t,
lph-2ph longitudinal flight muscle Kelsey (1957) recognized both
ventral (internal, 112) and dorsal (external, 113) muscles
Recognition of the two muscles in Hymenoptera is unnecessary
2ph-3ph reduced and attached laterally in most Hymenoptera
t,-t, in Hymenoptera, paired medial muscles passing from meta-
notum (t,) to scutellum over mesopostnotum (PN,), external in
some sawflies From PN; in Neuroptera
116 second dorsal diagonal
Mesothoracic ventral muscles
124 mesothoracic interfurcal
127 mesofurcal-spina
t,-2ph
sl3 fu,-fu,
sl4 fu,-lsps
t,-pn, dorsomedial attachment on mesoscutum to anterior face of
laterophragma (pn,), attachment to dorsal axillar surface where
transscutal arhculahon present
fu,-fu, from lateral face of fu, to anterior face or arms of fu, Kelsey
(1957) treats as three muscles (124-126) and Matsuda (1970) as
bundles of the same muscle Hymenoptera have maximum of
two bundles and homology of each is uncertain
fu,-sps, spina of profurca to apex of lateral arm of fu,.
181 metathoracic interfurcal
sl3 fuj-fu, • fu,-fu, posterior face of lateral arms of fu; to fu, Kelsey (1957) treats as
three muscles (181-183) Hymenoptera have maximum of two
180 metafurcal-spina
sl4
fUj-spSj spina of mesofurca to apex of lateral arm of fu,
Tergopleural muscles
137 laterophragmal-basalare t-p87
Furcal muscles
150a posterior furcal-laterophragmal t-sl fu,-2ph
150b anterior furcal-laterophragmal t-sl
151 furcal-pleural arm
170 coxal arhculation-furcal
fbl furcal-basalare
p-sl pl,-fu,
s-cx2
p-s3? fu2-pl37
t -ba, posterior face of laterophragma ( = t,) to apodeme of basalare
(ba,) Synonomy with t-p8 is questionable
fu,-pn, posterior attachment on lateral arm of fu, to anterior process of
laterophragma (ap)
fu,-pn, attachment anterior to 150a on anterior or lateral arm of fu, to
posterior lobe of laterophragma (pn,)
pl,-fu,_ lateral surface of fu, to pleural ridge
pl,-fu,t lateral surface of fu2 to coxal process on pleuron
fu,-ba, apex of lateral arm of fu2 to basalare (ba,), may be homologous
with pupal muscle of Apis (Daly 1964), well developed with
dorsal cap and apodeme in Dipnon (one preparation where ba,
dissected with fu;-ba, and pl,-ba, both attached). Possibly
homologous with Matsuda's p-s3 from furca to anterior margin
of succeeding episternum
Walking and indirect flight muscles
169 anterior furcal-coxal s-cx6 fu,-cx fu-cx fu,-cx
173 posterior furcal-coxal
s-cx3 fu2-cx fu-cx fu2-cx.
anterior base of fu; to median nm of coxa.
attachment on fu, posterior to muscle 169 to posteromesal rim of
174 furcal depressor of trochanter s-trl hy h",
fu,-tr, anterior base of fu; (or arms) to trochanteral apodeme
Volume 3, 1 994
Table 2. Taxa dissected for study of the mesofurcal-mesopostnotal complex.
245
Section: NEUROPTERIDA
Mantispidae
Chrysopidae
Corydalidae
Myrmeleontidae
Rhaphidiidae
Mantispa sp.
Chrysopa sp.
Corydalus sp.
Brachynemurus sp.
Rhaphidia sp.
Section: MECOPTERIDA
Bittacidae Bittacus sp.
Meropeidae Merope tuber Newman
Panorpidae Panorpa sp.
Order: HYMENOPTERA
SYMPHYTA
Xyeloidea
Xyelidae
Macroxyelinae: Macroxyela ferruginea
(Say); Xyelinae: Pleuroneura sp., Xyela
minor Norton
Megalodontoidea
Pamphilndae Cephalciinae: Acantholyda sp.,
Pamphiliinae: Pamphilius sp.
Tenthredinoidea
Blasticotomidae
Tenthredinidae
Dipnonidae
Cimbicidae
Pergidae
Argidae
Cephoidea
Cephidae
Siricoidea
Anaxyehdae
Siricidae
Xiphydrioidea
Xiphydriidae
Orussoidea
Orussidae
APOCRITA
Stephanoidea
Stephanidae
Blasticotoma sp.
Heteranthinae: Profenusa canadensis
(Marlatt); Nematinae: Nematus sp.;
Selandriinae: Aneugmenus flavipes
(Norton), Strongylogaster fflcirn(Norton);
Tenthredininae: Filacus sp., Macrophya
sp.
Dipnoninae: Diprion similis (Hartig)
Cimbicinae: Cimbex americana Leach;
Ambnnae: Zaraea americana Cresson
Acordulecerinae: Acordulecera sp.,
Syzygoninae: Lagideus Imexicana Smith
Arginae: Arge sp., Durgoa matogrossensis
Mai.
Cqjhus cinctus Norton,
Hartigia trimaculata (Say)
Syntexis libocedrii Rohwer
Siricinae: Urocerus albicornis (Fabricius);
Tremicinae: Tremex columba (Linnaeus)
Xipiudria abdommalis Say
Orussus terminalis Newman
Megischus bicolor (Westwood)
Trigonalyoidea
Trigonalyidae
Ceraphronoidea
Megaspilidae
Ceraphronidae
Evanoidea
Aulacidae
Evaniidae
Gasteruptiidae
Ichneumonoidea
Braconidae
Ichneumonidae
Chrysidoidea
Plumarndae
Sclerogibbidae
Embolemidae
Dryinidae
Bethylidae
Chrysididae
Orthogonalys pulchella (Cresson)
Megaspilus fuscipennis
Trichosteresis sp.
Ceraphron sp.
(Ashmead),
Prislaulacus strangahae (Rohwer)
Hyptw thoracica (Blanchard), Evania sp.
Gasteruptwn sp.
Alysiinae: Coelwius sp.; Hybrizontinae:
Hybrizon sp.; Macrocentnnae:
Macrocentrus sp.; Miscogastennae:
Apanteles sp.; Rogadinae: genus ?.
Ephialtinae: genus ?, Scambus sp.,
Megarhyssa sp.; Ophioninae: Enicospilus
sp.
Plumana sp.
Probethylus sp.
Etnbolemus nearcticus (Brues)
Anteoninae (female), Gonatopodinae
(male)
Anisepyris sp., Epyris sp.
Amiseginae: Adelphe anisomorphae
Krombein; Chrysidinae: Chrysis sp.,
Parnope sp.; Cleptinae: Cleptes sp.
Vespoidea
Tiphiidae
Sapygidae
Mutillidae
Sierolomorphidae
Pompilidae
Rhopalosomatidae
Bradynobaenidae
Formicidae
Scoliidae
Vespidae
Apoidea
Crabronidae
Heterogynaeidae
Myzinum sp.
Sapyga sp.
Sphaeropthalminae (males, 3 genera)
Sierolomorpha canadensis Provancher
Pepsinae: Calicurgus hyalinatus Fabr.;
Pompilinae: Aporinella galapagensis
Rohwer, Aporus sp.
Rhopalosoma sp.
Bradynobaenus sp.
Myrmicinae: Solenopsis invicta Buren
(queen & worker); Formicinae:
Camponotus planus F. Smith (queen),
Paratrechina sp. (queen)
Scolia sp.
Eumeninae: Odynerus sp.,
Parancistrocerus sp.; Vespinae:
Dolichovespula sp., Vespula sp.
Ectetnnius sp., Larra sp.
Heterogyna sp.
continued on next page
246
Journal of Hymenoptera Research
Table 2 continued
Andrenidae
Anthophondae
Apidae
Halictidae
Megachilidae
Proctotrupoidca
Diaprndae
Monomachidae
Vanhorniidae
Helondae
Ropronhdae
Proctotrupidae
Pelecimdae
Cynipoidea
Ihalndae
Eucoihdae
AUoxvstidae
Platygastroidea
Scelionidae
Platygastndae
sP.
Andrena sp.
Nomada sp., Ceratma sp.
Apis mellifera L., Bombus sp., Trigona sp.
Agnpostemon sp., Laswglossum sp.
Megachile sp.
Ambositrinae: Dissoxylabis sp.;
Belytinae: Aclista sp., Acropiesta sp.,
Aneurrynchus sp., Oxylabis sp.,
Diaprunae: Coptera sp., Paramenia sp.
Spilomicrus sp., Trichopna sp.
Monomachus sp.
Vanhornia eucnemidarum Crawford
Helorus sp.
Roprorua sp.
Exallonyx sp., Mwta vera (Fouts)
Pelecinus polyturator (Drury)
Jbfl/ifl sp.
genus 7
Alloxysta sp.
Scehonmae: y4n/cris sp., Calotelea sp.,
Oemasfobaeus sp., Gryon sp. (wingless),
Macrcteleia absona Muesebeck, Sparasion
sp., Teleasinae: Tnmorus sp.
Inostemmatinae: hocybus sp., Inoslemma
Mymarommatoidea
Mymarommatidae Palaeomymar sp
Chalcidoidea (additional dissections for work in progress
not listed)
Mymandae Gonafoccrus sp.
Pteromalidae Cleonvminae: Cleonymus sp., Oodera sp.
acter states. Coding of character states must in part
reflect the level of analysis. At a different level, for
example in an analysis of the relationships of
Tenthredinoidea that is associated with a more
extensive survey of taxa, itmightbe appropriate to
further partition the various shapes. These modi-
fications are difficult to characterize and here we
have combined them into one apomorphic state,
the reduction of well-separated anterior arms (state
2).
A clear transformation series leading to the
development of the mesofurcal bridge is found
within the Symphy ta beginning with the Cephidae.
In Cephidae (Fig. 9), the anterior arms are narrow
and elongate and only the extreme apices of each
arm are joined (state 3). The anterior arms are
considered to be homologous with those of
Xyeloidea and Megalodontoidea; therefore, state
3 is probably derived from state 1. Rasnitsyn (1988;
his character 19b) includes Cephoidea with
Siricoidea + Vespina based on having the fore-
arms of the mesofurca long and fused for some
distance. Although true for Siricoidea + Vespina,
the arms are only apically fused in Cephidae.
In Anaxyelidae (Fig. 10) and Siricidae (Fig.
11), the anterior arms are elongate and fused along
most of their length (state 4). The anterior arms of
Siricidae are laterally flattened (Fig. lib) and in
dorsal view each arm can be distinguished along
its entire length (Fig. 11a). Anaxyelidae have a
similar structure, including anterior placement of
muscle 150b, but fusion of the arms is more com-
plete (Fig. 10a). Posteriorly, the anterior furcal
arms of Anaxyelidae are connected by a thin hori-
zontal plate of cuticle.
InXiphydrndae(Fig. 12), theanteriorarmsare
completely fused and form a transverse mesofurcal
bridge (state 5), and the entire dorsal and lateral
surfaces form the posterior attachment for muscle
124. Displacement of muscles 150a and 150b to the
lateral arms suggests that fusion of the arms in
Xiphydriidae may be independent of the fusion in
Siricidae and Anaxyelidae.
Orussidae (Fig. 13) exhibit complete fusion of
the anterior arms into a smooth and bowed
mesofurcal bridge (state 6) with a strong median
anterior process that is the attachment site for
muscles 150 and 124, as occurs in Anaxyelidae and
Siricidae.
The groundplan states for the mesofurca of
Apocrita consists of 1) a complete mesofurcal
bridge (as in Xiphydriidae and Orussidae), 2) an
anterior medial projection supporting muscle 124
(fu^-fUj) (as in Xiphydriidae and Orussidae), and
3) lateral displacement of muscle 150a (fu,-pn, )
(as in Xiphydriidae). No Apocrita have muscle 150
originating on the anterior projection of the
mesofurca as in Orussidae. Because of the lateral
displacement of muscle 150a, which is similar to
Xiphydriidae (see character 2), Apocrita with a
mesofurcal bridge are coded as character state 4.
The mesofurcal bridge is absent (state 7) in
Ceraphronoidea (Fig. 25), Pelecinidae (Fig. 31),
some Chalcidoidea (including all Mymaridae),
Mymarommatidae, and Platygastridae (Fig. 28).
We consider that absence of the bridge is an
Volume 3, 1 994
247
apomorphic loss of the type of bridge found in
Xiphydriidae (state5), which is similar to the type
found in most Apocrita. Assuming that presence
of a bridge is a groundplan state for Apocrita
(whether states 4, 5 or 6), losses within each of
these taxa are considered irreversible (unlikely
that a bridge can be regained) and probably inde-
pendent. In Ceraphronidae, Pelecinidae,
Mymarommatidae, Platygastridae, and some
Mymaridae, the mesofurca is lyre-shaped with the
lateral arm terminating in a cup-shaped process
that supports muscle 150a. When the furcal bridge
is lost in Chalcidoidea (Aphelinidae, Encyrtidae,
Rotoitidae, Signiphoridae and Tricho-
grammatidae), the shape of the mesofurca and
attachment of muscle 150a are considerably differ-
ent.
Synonymy for mesofurcal bridge:
Siricidae & Vespidae: mesofurcal ring
(Tenthredinidae, Vespa, Weber 1925). Ichneu-
monoidea: mesofurcal bridge (Stenobracon,
Alaml951). Aculeata:mesofurcalbridge(ArF2)
(Vespula, Duncan 1939); supraneural bridge
(Apis, Snodgrass 1942). Chalcidoidea: arch of
the furca (Monodontomerus, Bucher 1948); ten-
dinous arch of the mesofurca (Tetramesa, James
1926).
Character 2. Laterophragma of mesopostnotum
(pn2)
In Neuropterida, Mecopterida, Xyelidae and
Pamphiliidae, the laterophragma of the
mesopostnotum forms a lobe (pn,, Figs. 4, 6) (state
0) that extends obliquely into the mesothorax from
the lateral corners of the mesopostnotum, mediad
of the lateral attachment to the upper mesepimeron
and lateral to the second phragma. The
laterophragma in all of these taxa forms the attach-
ment site for muscle 150 (fu2-pn,, see Character 3),
muscle 1 16 (t2-pn„ see Character 4) and muscle 137
(pnj-ba,). The posterior face of the laterophragma
forms the attachment site for muscle 137 (pn,-bav
Fig. 3). Muscle 137 is usually small and difficult to
trace, but it is apparently lost in Pergidae and all
Apocrita.
In Xyelidae (Figs. 3, 4) and most Symphyta
(Figs. 6, 8, 9), an apodeme (ap) is present on the
anterolateral margin of the lobe that serves as the
attachment site for muscle 150a (fu,-pn2 ). The
laterophragma is functionally coupled with
muscles 150 and 116, and the loss of either of these
muscles is associated with a corresponding change
in structure of the laterophragma. To avoid dupli-
cation of character coding, the presence or absence
of certain structures of the laterophragma are
treated under other characters. For example, the
anterior apodeme is lacking in Neuropterida and
Mecopterida but this was not coded as a different
state for this character because it is reflected in the
differentiation of muscle 150a in Hymenoptera,
which is dealt with as Character 3. A broad
laterophragma with a small anterior apodeme
(state 0) occurs in Xyelidae, Pamphiliidae (and
probably Megalodontidae), Blasticotomidae,
Tenthredinidae, Argidae, Diprionidae, and
Cephidae.
The laterophragma of Cimbicidae (Fig. 8) pos-
sesses an exaggerated apodeme (ap) and an en-
larged posterior lobe (pn,), which is fused with the
second phragma (2ph). The laterophragma is
unique in form and apparently autapomorphic.
In Xiphydriidae (Fig. 12), Monomachidae (Fig.
14), Vanhorniidae (Fig. 15), Cynipoidea (Fig. 22),
Ceraphronoidea (Fig. 25) and most Diapriidae
(Fig. 29), the laterophragma is excised between the
elongate apodeme (ap) and the posterior lobe
(pn,) (statel). In most of these taxa the apodeme of
the laterophragma extends medially and horizon-
tally into the thoracic cavity, but in Cynipoidea (cf .
Fig. 22), the apodeme is vertical. The posterior lobe
is lost in most Apocrita (incuding some Diapriidae),
but this was not coded as an additional state
change for this characterbecause it reflects the loss
of the muscle 116 (Character 4). The anterior
apodeme (=axillary lever) or associated attach-
ment for muscle 150a is present in all Apocrita.
Additional character state changes for the
laterophragma in Apocrita are based only on
changes in the shape of the anterior apodeme.
In Pergidae (Fig. 5) and Orussidae (Fig. 13),
the laterophragma is reduced to a narrow triangu-
lar process that forms an attachment for the ten-
don of muscle 150 (state 2). This reduction is also
associated with the loss of muscle 116. State 2
could be derived from either state 0 or state 1.
In Apocrita, the axillary lever (ap) occurs in a
variety of shapes that probably have different
effects on leverage of the laterophragma with re-
spect to the fourth axillary sclerite and the second
phragma. In all Apocrita the apex of the axillary
lever maintains a connection with muscle 150a
24S
Journal of Hymenoptera Research
(fu^-piv )• The plesiomorphic lever for Apocrita is
narrow and inflected medially into the thorax as
found in Xiphidriidae (statel) and occurs either in
association with a posterior lobe (most Diapriidae,
Monomachidae (Fig. 14), Vanhorniidae (Fig. 15),
Ceraphronoidea (Fig. 25) and Cynipoidea (Fig. 22)
or without the posterior lobe (Tngonalyidae (Fig.
23), Stephanidae (Fig. 24), Diapriidae (Fig. 29),
Proctotrupidae (Fig. 32), some Ichneumonidae,
Mymarommatidae, and Chrysidoidea (Figs. 34-
39) (except some Chrysididae). The axillary lever
exists in a variety of forms in Apocrita but in
shapes that are difficult to separate into discrete
states, especially without a more thorough survey
of the apocritan taxa. In all cases where the lever
was narrow and inflected medially the character
was coded as state 1.
In Platygastridae (Fig. 28), some Scelionidae
and most Chalcidoidea, the axillary lever is de-
flected ventrally (state 3). In Mymarommatidae,
the lever is robust and inflected medially (statel).
In Sparasion (Scelionidae; Fig. 16), the lever is
reduced to a small cup-shaped lobe on the
laterophragma (autapomorphic and not coded).
In other Scelionidae and Mymaridae the lever is
short, narrow and medially inflected (statel).
The axillary lever in Stephanidae (Fig. 24) and
most Chrysidoidea (Figs. 35-38, also in Fig. 34 but
this view is slightly different) is elongate and
strongly inflected medially (state 1). This confor-
mation of the lever is likely the precursor to the
lever found in Apoidea and Vespoidea, discussed
below. However, in Cleptinae and Chrysidinae
(Chrysididae) the lever is reduced (state 4) form-
ing a short, broad process attaching to a broad
tendon of the shortened muscle 150a (Fig. 40).
In Apoidea (Figs. 45, 46, 54) and Vespoidea
(Figs. 41, 42, 44) the axillary lever is robust and
strongly appressed to the inner surface of the
second phragma (state 5). This type of lever was
found in all Apoidea examined and is associated
with a robust muscle 150a. Except in
Bradynobaenidae and Formicidae, muscle 150a is
conical and attached to the lever by a narrow
tendon or robust and nearly tubular (Mutillidae
and Sapygidae). In Sierolomorphidae, the lever is
not closely appressed to the second phragma (in-
flected medially about 30°), but this was not treated
as a different state (intermediate between states 1
and 5). In Bradynobaenidae, the lever is reduced to
a rounded knob (attaching to a narrow tendon of
muscle 150a) (state 4?). In Formicidae (Fig. 43), the
lever forms a narrow, twisted apodeme (state 1?).
Neither Bradynobaenidae and Formicidae were
broadly surveyed and other forms may exist.
In Ichneumonidae the axillary lever is either
closely appressed to the second phragma and
similar to the lever found in Vespoidea and
Apoidea (state 5), or rod-like and extending medi-
ally into the mesolhorax (state 1), or inflected
medially and dorsally (Fig. 17a). A robust lever is
not found in Braconidae, including Hybrizontinae
(Fig. 33), in which it was knob-like or short and
slightly deflected ventrally (state 1), similar to
Trigonalyidae and some Proctotrupoidea. The le-
ver of Gasteruptiidae (Fig. 26) is similar to some
Ichneumonidae but is oriented vertically with re-
spect to the second phragma. In Oodera
(Pteromalidae), the lever is robust, appressed to
the second phragma and horizontal (state 5). This
form of the lever appears to be convergent with
Ichneumonidae and Aculeata, since in Oodera
muscle 150a attaches to the entire ventral aspect of
the apodeme as in other Chalcidoidea.
All Apiformes have the axillary lever sepa-
rated (state 6) as an independent sclerite (Snodgrass
1942). This feature was verified in all of the
Apiformes examined here. The apical connection
of the mesopostnotum is weak and encased by the
cup-shaped basal process of the axillary lever
(Figs. 45,54).
Synonomy for laterophragma:
Symphyta: lateral lobe of second phragma
(2ph) (Daly 1963). Icnneumonoidea: muscle
bearing process of mesopostphragma
(Stenobracon, Alam 1951); axillary lever
(Snodgrass 1942). Cynipoidea: mesopostnotal
apodeme (Ibalia, Ronquist and Nordlander
1989); 2ph (Daly 1963). Aculeata: axillary le-
ver (Apis, Snodgrass 1957; Bombus, Pringle,
1957, 1960, 1961 ); accessory sclerite of the fourth
axillary sclerite (Apis, Snodgrass 1910); inner
process of mesopostphragma (Vespa, Weber
1925 [states that lever turns apex of
mesophragma and the connected axillary 4]);
anterior process of mesopostphragma (MPPh J
(Vespula, Duncan 1939). Hymenoptera: axil-
lary lever (Matsuda 1970).
Character 3. Furcal-laterophragmal muscle
In Neuropterida and Mecopterida, only a
Volume 3, 1994
249
single muscle (150, fu2-pn2) acts between the
mesofurca and laterophragma of the
mesopostnotum (state 0) (Kelsey 1957, Matsuda
1970). In the majority of Symphyta, including
Xyelidae and Pamphiliidae, two distinct muscles
(150a and 150b) operate antagonistically on the
enlarged laterophragma (Figs. 2, 3, 4, 6). The pre-
sumed division of muscle 150 into two muscles is
considered as an apomorphy for Hymenoptera
(state 1). Muscle 150a (fu,-pn^) originates on the
lateral arms of the mesofurca posterior or lateral to
the origin of 150b and inserts onto an anterior
apodeme or process of the laterophragmal lobe
(Figs. 4, 6, 7, 9). Muscle 150b (fu,-pn2 ) originates
anteriorly or medially on the lateral or anterior
arms of the mesofurca and broadly attaches to the
margin of the laterophragmal lobe (Figs. 4, 6, 7, 9).
The fibres of muscle 150b are often arranged so
that the anteriormost fibres attach to the
posteriormost or innermost margin of the
laterophragmal lobe, although this is dependent
on the size of the muscle and lobe.
If both muscles are present, muscle 116 (t,-pn,)
is also present. Loss of muscle 150b (state 2) occurs
in Cimbicidae, Monomachidae, Vanhorniidae,
Diapriidae, Cynipoidea, and Ceraphronoidea, all
of which have retained both the laterophragmal
lobe (presumably the former point of insertion of
muscle 150b) and muscle 116. All Apocrita have
lost muscle 150b. In Anaxyelidae (Fig.10), muscle
150b is attached to the posterior face of the poste-
rior lobe (versus the margin) and muscle 150a is
retained. Because of a similar placement and at-
tachment of muscles in Siricidae (Fig. 1 1), it can be
postulated that muscle 150a is lost and 150b re-
mains (state 3). When both the laterophragmal
lobe and muscle 116 are missing, as in Pergidae
(Fig. 5) and Orussidae (Fig. 13), it is difficult to
assess which muscle, 150a or 150b, has been lost.
We could assume that muscle 150b is lost in both
families. However, the forward attachment of
muscle 150 to the mesofurca in Orussidae suggests
that it is homologous with muscle 150b of
Anaxyelidae and Siricidae. Therefore, the con-
figuration in Pergidae and Orussidae could also
be treated as an parallel loss of muscle 150a. Be-
cause of this uncertainty, we code the reduction to
a single muscle in Pergidae and Orussidae as of
questionable homology (state ?). Based on the
presence of muscle 150a in the apocritan families
mentioned above, we assume that muscle 150b is
lost in all Apocrita without a posterior
laterophragmal lobe. Autapomorphic losses of
muscle 150a occurs in workers of Formica and
Camponotus (Markl 1966, Saini et al. 1982).
Synonymy for muscle 150:
Tenthredinidae: Ilisml (Dolerus, Schizocerus,
Tenthredo, Weber 1927), 25 & 26
(Euthomostethus, Maki 1938). Ichneumonidae:
25 (Psilopsyche, Maki 1938); 63 (Stenobracon,
Alam 1951). Aculeata: 25 (Vespa, Maki 1938);
78 (Formica, Markl 1 966; Apis, Snodgrass 1942);
IIdv2 (Vespula, Duncan 1939); Ilism (Vespa,
Weber 1925); fu2-2ph (Apis, Daly 1964). Hy-
menoptera: t-sl (Matsuda 1970).
Character 4. Second dorsal diagonal (=phragmal
flexor) muscle
Muscle 116 (t2-pn2) is found attaching between
the mesonotum and the anterior face of the
laterophragmal lobe in Neuropterida, Mecopterida
and most Symphyta (Figs. 2, 3, 12, 14, 15, 20), and
is therefore plesiomorphic for Hymenoptera (state
0). Muscle 116, and the associated posterior lobe of
the laterophragma, is lost (statel) in Pergidae,
Orussidae and most Apocrita. It seems unlikely
that a functional complex composed of both muscle
116 and the corresponding posterior lobe of the
laterophragma could be regained, and we con-
sider the apomorphic state (loss of muscle and
lobe) to be irreversible. An assumption of irrevers-
ibility has obvious consequences for models of
character state change which are discussed in the
later section on parsimony analyses. Muscle 116
and the corresponding posterior lobe are present
in Monomachidae, Vanhorniidae, Cynipoidea,
Ceraphronoidea and most Diapriidae, and it is
likely that this complex is the groundplan for
Apocrita (Daly 1963; Gibson 1985). The loss of this
complex in Pergidae and Orussidae is therefore
convergent with the loss in most Apocrita.
Synonomy for muscle 116:
Symphyta: 21 (Euthomostethus, Maki 1938).
Symphyta, Monomachidae, Diapriidae,
Vanhorniidae, Ceraphronoidea, Cynipoidea:
t,-2ph (Daly 1963; Gibson 1985). Hymenoptera:
tl2 (Matsuda 1970).
250
Journal of Hymenoptera Research
Character 5. Mesopostnotum and scutellar-
metanotal muscle
In Neuropterida and Mecopterida, the
mesopostnotum is broadly exposed medially; al-
though in Corydalus (Megaloptera) the
mesopostnotum is weakly sclerotized medially
and appears to be split. The scutellar-postnotal
muscle (muscle 114, t2-t3; absent in Bittacus) passes
internally from its origin on the scutellar area of
the mesonorum to the anterior medial margin of
the mesopostnotum (Kelsey 1957; Matsuda 1970).
Within Hymenoptera, changes in the structure of
the mesopostnotum and attachment of muscle 114
are correlated; however, the attachment of muscle
114 to the mesopostnotum is unique with respect
to the outgroup, and the outgroup is scored as a
separate state (state 0).
In Hymenoptera, muscle 114 (t,-t3) arises
dorsomedially from a fan-shaped attachment on
the mesonorum and passes medially over the
mesopostnotum (rarely under), to a narrow me-
dial attachment on the anterior margin of the
metanotum. In all Symphyta, our dissections indi-
cate a posterior attachment of muscle 114 to the
metanotum. Our observations are supported by
illustrations of attachments to the metanotum in
Weber (1925) for Vespa (his IldlmJ and Markl
(1966) for Apis (his 70). In Eutomostethus
(Tenthredinidae), muscle 114 was described as
inserting on the median membrane that divides
the mesopostnotum (Maki 1938, his 19 & 20), but
in Tenthredinidae muscle 114 passes through the
membrane to the metanotum. The attachment of
muscle 114 to the metanotum is apomorphic for
Hymenoptera.
Determination of the groundplan condition
for Hymenoptera is complicated by the presence
of different character states in the two extant sub-
families of Xyelidae. In Xyelinae (Xyelidae) (based
on Pleuroneura, Fig. 47, and Xyela), the
mesopostnotum (PNJ is broadly exposed dor-
sally and the scutellar-metanotal muscle (1 14, t;-t,)
passes under the mesopostnotum through a small
emargination of its posterior margin (state 1,
broadly exposed and 114 internal). In
Macroxyelinae (Xyelidae) (based on Macroxycla
ferrunginea (Say), Megaxyela tricolor Norton,
Megaxyela aviingrata (Dyar) and Xyelicia neurotica
Ross), Megalodontoidea (Fig. 48),Tenthredinoidea
and Cephoidea (Fig. 49), the mesopostnotum is
exposed dorsally (depending on the contortion of
the mesosoma) and muscle 114 passes externally
over the mesopostnotum as a tendon enclosed by
a sheath of connective tissue (state 2, broadly
exposed and 114 external). In groups with an
external muscle and and an exposed
mesopostnotum (state 2), the anterior medial mar-
gin may be shallowly or deeply emarginate (ap-
pearing split) underneath muscle 114. In some
genera of Tenthredinidae and Dipnonidae, the
mesopostnotum is obscured by posterior devel-
opment of the scutellum, but otherwise conforms
to state2 (see Character 6).
In Anaxyelidae, the mesopostnotum and
muscle 114 are the same as in Xyelinae (Fig. 47).
Initially, we coded this as the same character state
(state 1). However, this caused problems in devel-
oping an additive coding scheme that would force
the anaxyelid state to be aucapomorphic and not
transitional between states 2 and 4. To resolve this
problem, Anaxyelidae were assigned a new char-
acter state (state 3), which is treated as an
autapomorphy. In all of the analyses, coding for
Anaxyelidae and Xyelinae as state 1 or state 3 had
no effect on tree topology.
In Siricidae, a median vertical process on the
anterior margin of the metanotum forms the pos-
terior attachment of muscle 114, which is internal
and lacks any connective sheath. Although the
process is unique for Hymenoptera, this confor-
mation is considered to be a modification of state
2 even though the mesopostnotum is generally not
exposed. The scutellum often extends over the
metanotal process and, as in some Tenthre-
dinoidea, the internalization of muscle 114 and
mesopostnotum may be secondary. Snodgrass
(1910) reported that the mesopostnotum of Tremex
cohtmba was exposed medially; however, we ob-
served an exposed mesopostnotum only in some
specimens of Urocerus. Some Cephidae (Fig. 49)
also have a similar attachment to a peg-like pro-
cess (mp) on the metanotum.
In Xiphydriidae (Fig. 50), Orussidae and
Apocrita, muscle 114 (t2-t3) and the mesopostnotum
(medially) are completely internal and muscle 1 14
passes medially over the second phragma (state
4). An autapomorphic modification of state 4 is
found in Xiphydriidae, in which the cuticle of the
second phragma (between the lobes of the
pseudophragma) encircles the muscle tendon as it
passes through to the metanotum. Further devel-
opment of the mesopostnotum and its associated
Volume 3. 1 994
251
phragma in Apocrita is toward an even greater
degree of internalization and a lateral shift of the
points of articulation with the scutellum and met-
anotum (Snodgrass 1910) (see character 10). In
Aculeata (Fig. 53), the second phragma is almost
entirely disassociated mesally and the notal at-
tachments are reduced to lateral lamellae along
the extreme lateral margins (Snodgrass 1942).
Synonomy for muscle 114:
Tenthredinidae: 19 & 20 (Euthomostethus, Maki
1938). Ichneumonidae: 20 & 21 (Psilopsyche,
Maki 1938). Aculeata: IIdlm2 {Vespa, Weber
1925); 23 (Vespa, Maki 1938)'; Ilisl (Vespula,
Duncan 1939); 70 (Apis, Markl 1966); t,-t3 {Apis,
Daly 1963). Hymenoptera: tl3 (Matsuda 1970,
Brodskiy 1992).
Character 6. Formation of pseudophragma
The postphragma (2ph) originates at the junc-
tion of the mesopostnotum and metanotum and
forms the posterior attachment site for the longitu-
dinal flight muscles (112, lph-2ph) (Snodgrass
1910). In Mecopterida, Neuropterida, Xyelidae (Fig.
47), Megalodontoidea (Fig. 48) and Tenthre-
dinoidea, the anterior margin of the
mesopostnotum is broadly attached to the poste-
rior margin of the mesonotum (scutellum) by a
narrow conjunctiva (60, term from Ronquist and
Nordlander 1989) (state 0). In Cephidae (Fig. 9b),
Anaxyelidae, Siricidae (Fig. lib), Xiphydriidae
and Apocrita, the anterior margin of the
mesopostnotum is developed into a broad bilobed
internal plate (pseudophragma) that extends an-
teriorly and dorsally beyond the dorsal attach-
ment of the mesopostnotum to the mesonotum
(state 1). The pseudophragma is an extention of
the second phragma and both form the posterior
attachment for the longitudinal flight muscles (lph-
2ph). A pseudophragma is not apparent in
Orussidae (Fig. 13) and some Ichneumonidae and
has probably been lost.
Characters 7-8. Furcal-spina muscles
In Xyeloidea (Fig. 3) and Pamphiliidae (Fig. 2,
indicated by dashed line), muscles 127 (fu,-sps,)
and 180 (fu?-sps,) connect the spina with the furca
of the following segment (stateO). Muscle 127 is
present in both Neuropterida and Mecopterida
and muscle 180 is found only in Neuropterida
(Maki 1938, Kelsey 1957, Matsuda 1970); we veri-
fied their presence only in Corydalus. Muscle 127 is
lost (state 1, character 7) in all Hymenoptera ex-
cept the Xyeloidea and Megalodontoidea.
Rasnitsyn (1969, fig. 204) illustrates a muscle con-
nection (127?) between sps, (his sst) and fu2 in
Tremex (Siricidae), but we did not observe this in
either Tremex or Urocerus. The loss of muscle 180
(state 1, character8) is a synapomorphy of Apocrita.
Synonomy for muscle 127:
Apidae: fu,-lsps [pupal muscle only] (Apis,
Daly 1964). Hymenoptera: sl2 (Matsuda 1970;
not listed by Matsuda for Hymenoptera).
Synonomy for muscle 180:
Symphyta: living (Dolerus, Schizocerus,
Tenthredo, Weber 1927).
Character 9. Furcal-basalare muscle
The furcal-basalare muscle (fbl, fu,-ba3) ap-
pears to be found only in Hymenoptera (state 1).
The muscle is apparently absent in the outgroup
(state 0). Muscle fbl could be homologous with
Matsuda's p-s3 which connects the furca to the
anterior margin of the succeeding episternum.
Muscle p-s3 is found in Neuropterida (IIpml7,
Korn 1916 for Myrmeleon), but apparently not in
other Neuropterida or Mecopterida (Matsuda
1970). In Symphyta (Figs. 2, 3, 5-13) muscle fbl
extends from the extreme lateral apex of the
mesofurcal arm to the apodeme of the basalare of
the hindwing (ba?). This muscle was first illus-
trated, but not discussed, by Rasnitsyn (1969, fig.
204) for Tremex. In Diprionidae (Fig. 7), muscle fbl
ends in a sclerotized cap attached to the apodeme
of the basalare, along with muscle 137 (t,-ba )
which originates from the posterior face of the
laterophragma (t3).
Muscle fbl is absent (state 0) in most Apocrita.
However, what appears to be the same muscle
occurs in Mutillidae (Fig. 18), Rhopalosomatidae
(Fig. 19) and Bradynobaenidae (state 1). The muscle
in these taxa originates laterally on the mesofurcal
arms (dorsal to the furcal-trochanteral muscle,
musclel74) and narrows to a fine tendon that
enters the pleural area near the base of the hind
wing. The dorsal point of insertion was not ob-
served but the muscle enters the pleural area pos-
terior to the subalar muscle of the mesothorax and
may insert onto the basalare or axillary sclerites of
the hind wing. No such muscle was observed in
252
Journal of Hymenoptera Research
other Apocrita. The loss of muscle fbl in Apocrita
could be the result of a shift of the muscle origin
from the furca to the pleural wall. Gibson (1986)
reported a unique muscle in Eupelmidae and
Pteromalidae (pl2-3ax3a) that attached between
the lower mesepimeron and possibly the metatho-
racic third axillary sclerite (of uncertain insertion;
Gibson pers. comm. 1993). The homology of fbl
with the pleural muscles is beyond the scope of
this study.
Character 10. Lateral articulation of
mesopostnotum
In Xyeloidea, Megalodontoidea (Fig. 51) and
Tenthredinoidea and the outgroup taxa, the
mesopostnotum is broadly and evenly joined to
the upper mesepimeron (state 0). In Cephidae
(Fig. 52), Anaxyelidae and Siricidae, the juncture
is invaginated and connected by weak cuticle
within the invagination, but maintains a strong
cuticular connection at the anteriormost point of
attachment (state 1). In Xiphydnidae, Orussidae
and Apocrita, the mesopostnotum is completely
internal and the lateral connection of the
mesopostnotum to the upper mesepimeron is re-
duced to the anteriormost point of attachment
(state 2).
Character 11. Fusion of lateral arms of mesofurca
and metafurca
In most Aculeata the lateral arms of the
mesofurca and metafurca are closely appressed,
but as in most other Hymenoptera and the
outgroup taxa, they are broadly separated and
joined by the interfurcal muscle 181 (state 0) (Fig.
3). In Apoidea (Figs. 46, 53), the lateral arms of
both furcae are fused at the junction with the
mesofurcal bridge and the interfurcal muscle is
lost (state 1). The lower furcal arms form a four-
cornered brace for the furcal complex, and the
metafurcal arms are divided lateral to the furcal
bridge. A similar fusion of the mesofurca and
metafurca occurs in Amiseginae (Chrysididae; Fig.
39) and Gasteruptiidae (Fig. 26), but in both cases
muscle 181 is retained and these are considered to
be non-homologous and autapomorphic changes.
Additional coding as autapomorphic states is un-
necessary for this analysis but may be warranted
in more extensive studies of related taxa.
Character 12. Furcal process for trochanteral
muscle 174
In most Hymenoptera and the outgroup taxa,
the furcal depressor of the trochanter (muscle 174,
fu^-tr^) attaches to the lateral arms of the mesofurca
(Fig. 3) (state 0). In Chrysididae (Cleptinae and
Chrysidinae), muscle 174 attaches to anterior ex-
tensions of the lateral furcal arms (Fig. 40, vp)
(state 1). In Pompilidae (Fig. 42), Sapygidae,
Scoliidae, Sierolomorphidae and Vespidae (Fig.
41), muscle 174 attaches to similar extensions (vp)
that arise instead from the furcal bridge (state 2).
In Vespoidea, muscle 174 has two origins - the
lateral furcal arms and the ventral process (= scoop
shaped processes, schT, Weber 1925; anterior pro-
cess of mesofurcal bridge, PF„ Duncan 1939).
PHYLOGENETIC HYPOTHESIS FOR
HYMENOPTERA
Figs. 55, 56
The higher classification of Hymenoptera has
come under increasing scrutiny following reviews
published by Konigsmann (1977, 1978a) and
Rasnitsyn (1969, 1980). Konigsmann (1978a) pre-
sented evidence that Hymenoptera was comprised
of two monophyletic sister taxa, the Symphyta
(excluding Cephoidea) and Cephoidea + Apocrita,
with most apocritan families presented as a series
of unresolved lineages. Although this more tradi-
tional concept of the Symphyta is still used (Zessin
1985), it has not been accepted in more recent
papers except to present an alternate hypothesis of
relationships (Gauld and Bolton 1988). Rasnitsyn
(1980) presented evidence that Symphyta is
paraphyletic relative to Apocrita with Cephoidea
as the sister group of Apocrita. Studies subsequent
to Rasnitsyn (1980) (Gibson 1985, 1986; Naumann
and Masner 1985; Carpenter 1986) led Rasnitsyn to
modify his original hypotheses in a later paper
(Rasnitsyn 1988), in which Cephoidea were placed
as a more basal lineage of Hymenoptera and
Orussidae as the sister group of Apocrita. A far
more resolved set of relationships among the para-
sitic Apocrita (Fig. 56) were presented by Rasnitsyn
(1988). Some of Rasnitsyn's hypotheses have been
corroborated in recent morphological or phyloge-
netic treatments (Johnson 1988; Whitfield et al.
1989), although Gibson (1993) presented evidence
against the biphyletic origin of Hymenoptera from
Xyelidae and Brothers and Carpenter (1993) sup-
Volume 3, 1 994
253
ported a different hypothesis for Chrysidoidea
and Aculeata. For our data, these modifications
affect only the distribution of character 12 in the
Aculeata, and have no effect on the parsimony
arguments presented in a later section. Cladograms
comparing the hypotheses of Rasnitsyn and
Konigsmann are presented in Whitfield (1992).
Rasnitsyn (1980, 1988), Shcherbakov (1980,
1981) and Gibson (1993) have proposed Xyelidae
as the most basal extant lineage of Hymenoptera.
As well as possessing a branched Rs vein, Xyelidae
retain the anepisternum as an integral part of the
pleuron and retain the plesiomorphic condition of
a relatively equally proportioned meso- and meta-
thorax (Gibson 1993). Other than the structure of
the antenna, which may be plesiomorphic, there
are no features that support the monophyly of the
Xyelidae, and Rasnitsyn (1988) proposed that the
two subfamilies of Xyelidae belong to two lin-
eages of Hymenoptera (Hymenoptera biphyletic
with respect to the two subfamilies). Gibson (1993
and personal communication, 1993) argues that
Hymenoptera excluding Xyelidae are monophyl-
etic but that no characters support the monophyly
of the Xyelidae. The mesofurcal-mesopostnotal
complex of Pamphiliidae and both subfamilies of
Xyelidae are basically the same and we consider
them to represent the plesiomorphic condition for
Hymenoptera.
Rasnitsyn (1988) classified the suborder
Siricina into three infraorders that include the
following extant families: Xyelomorpha (Xyelidae),
Tenthredinomorpha (Argidae, Blasticotomidae,
Cimbicidae, Pergidae and Tenthredinidae [includ-
ing Diprionidae]), and Siricomorpha (including
Megalodontoidea [=Pamphilioidea]: Pamphiliidae
and Megalodontidae; Cephoidea: Cephidae;
Siricoidea: Anaxyelidae, Siricidae and
Xiphydriidae). Rasnitsyn (1988) treated Xyelidae
as paraphyletic with respect to Tenthredinoidea
and remaining Hymenoptera, and the three fami-
lies of Siricoidea are paraphyletic with respect to
the Orussidae + Apocrita (Suborder Vespina).
Rasnitsyn (1988) placed Cephoidea as basal to the
Siricoidea + Apocrita, rather than sister group to
the Apocrita as in his earlier treatments (Rasnitsyn
1969, 1980).
As a framework for evaluating our results, we
have compiled the results of Rasnitsyn (1988) and
Gibson (1985, 1993) to derive the cladogram pre-
sented in Figure 55. Contrary to Rasnitsyn (1988),
Hymenoptera excluding Xyeloidea are treated as
monophyletic based on the presence of a
postspiracular sclerite (=detached anepisternum)
(Gibson 1985, 1993) and an apically simple Rs vein
that is not bifurcate (Rasnitsyn 1988). Xyelidae are
treated as paraphyletic to allow for discussion of
Character 5. Tenthredinoidea are monophyletic
and placed here as sister group to the remaining
Hymenoptera, excluding Xyeloidea and
Megalodontoidea, based on a reduction in the
number of preapical tibial spurs (from 2-4 spurs
on the hind legs in Xyeloidea and Megalodontoidea
tolessthanl in Tenthredinoidea, l-2inCephoidea
and 0 in remaining Symphyta and Apocrita; H.
Goulet and G. Gibson, pers. comm., 1993), and the
loss of the ventral mesofurcal-spina muscle, fu2-
spSj, which is present only in Xyeloidea and
Megalodontoidea (see Character Analysis).
Remaining infraorders of extant Vespina pro-
posed by Rasnitsyn (1988) include the
Orussomorpha, Evaniomorpha (Stephanoidea,
Evanioidea), Proctotrupomorpha (Proctotru-
poidea, Cynipoidea, Chalcidoidea), Ichneu-
monomorpha (Ichneumonoidea), and
Vespomorpha (Chrysidoidea, Sphecoidea,
Pompiloidea, Scolioidea, Formicoidea and
Vespoidea). Recent classifications (Naumann 1992;
Huber and Goulet 1993) treat the five families of
Rasnitsyn's Stephanoidea as higher taxa
(Ceraphronoidea [Ceraphronidae and
Megaspilidae], Megalyroidea, Stephanoidea and
Trigonalyoidea). Following Brothers (1975) and
Huber and Goulet (1993), the Aculeata are re-
ferred to only three superfamilies (Chrysidoidea,
Apoidea and Vespoidea) and Mymarommatidae
are placed as a superfamily.
GROUP ANALYSIS
Character states are referred to in brackets as
[charactenstate]. Relationships are discussed ac-
cording to the phy logenetic hypotheses illustrated
in Figures 55 and 56. The distribution of character
states for each taxon are listed in Appendix 2.
254
Journal of Hymenoptera Research
Xyeloidea + Megalodontoidea + remaining
Hymenoptera
Figs. 1, 2, 3, 4, 6, 47, 48, 51
Four characters of the mesofurcal-
mesopostnotal (MF-MPN) complex are not found
in the outgroups and therefore support the mono-
phyly of Hymenoptera: long anterior arms of the
mesofurca [1:1]; two furcal laterophragmal muscles
[3:1]; muscle 114 attaching to the metanotum (part
of Character 5, see below); and furcal-basalare
muscle (fbl, fu,-ba.,) present [9:1].
Xyeloidea and Megalodontoidea are
plesiomorphic for all characters postulated for
Hymenoptera in the mesofurcal-mesopostnotal
complex. Megalondontidae were verified for ex-
ternal characters only and the presence of anterior
furcal arms [1:1] was based on fig. 189 in Rasnitsyn
(1969). Rasnitsyn (1969) also refers to the
mesofurco-metapre-episternal muscle as found
only in Xyela. This muscle was not identified in our
dissections, although it may be synonymous with
muscle 127 (fu^sps,).
The mesopostnotum is external and broad
medially in both Xyeloidea and Megalodontoidea.
In extant Xyelinae (Xyelidae), the mesopostnotum
is entire medially and muscle 114 is internal [5:1,
Fig. 47]. A similar conformation is found in the
outgroup, but the attachment of muscle 114 is
different [5:0]. In Macroxyelinae (Xyelidae),
Megalodontoidea and Tenthredinoidea, the
mesopostnotum is almost completely separated
medially and muscle 114 passes externally [5:2,
Fig. 48]. The deep emargination of the
mesopostnotum associated with state 5:2 is visible
externally and can be interpreted from fossil Hy-
menoptera. Based on illustrations in Rasnitsyn
(1969), external passage occurs in the fossil xyelids:
Archexyelinae: Asioxyela (fig. 34), Leioxyela (fig.
41), Triassoxyela (fig. 32), Xiphoxyela (figs. 45, 46);
Macroxyelinae: Agaridyela Rasn. (fig. 41),
Chaetoxyela Rasn. (fig. 76), Ceroxyela Rasn. (fig. 77);
and Xyelinae: Eoxyela Rasn. (fig. 59), Lydoxyela
Rasn. (fig. 68), Spathoxyela Rasn. (fig. 42), Xyelisca
Rasn. (fig. 61), and Xyela mesozoica Rasn. (fig. 62).
If correctly interpreted in fossil Xyelinae, the inter-
nal passage of muscle 114 and an entire
mesopostnotum are derived within extant
Xyelinae, and the external passage of muscle 114
and an emarginate mesopostnotum are probably
apomorphic for Hymenoptera
Tenthredinoidea + remaining Hymenoptera
Monophyly of Tenthredinoidea and the re-
maining Hymenoptera is supported by the loss of
muscle 127 (fu2-spSj) [7:1].
Tenthredinoidea
Figs. 5, 7-8, 20, 21
The anterior mesofurcal arms are reduced or
lost in all Tenthredinoidea [1:2].
Blashcotomidae (Fig. 20)
The plesiomorphic combination of muscles is
present. Anterior furcal arms are present, but they
are short and thin. As discussed earlier (analysis of
characterl ), because of uncertainties in the homol-
ogy of reduced arms, we have scored both re-
duced and absent anterior furcal arms as the same
character state (1:2). The anterior apodeme of the
laterophragma is absent and both 150a and 150b
attach to the apex of the posterior lobe (Fig. 20a).
This is considered to be an autapomorphic devel-
opment, but there is also a general similarity with
the laterophragma of Nematus (Fig. 19), in which
short anterior furcal arms are also present.
Tenthredinidae (Fig. 21)
The plesiomorphic combination of muscles is
present in all taxa examined, and the structure of
the laterophragma is the same as in Xyelidae and
Pamphiliidae. The anterior arms of the mesofurca
vary from narrow, elongate apodemes extending
from the anterior face of the lateral arms (Nematus),
to short cup-like structures similar to those found
in Diprionidae, to completely absent (most
Tenthredinidae). Muscle 180 (fu^sps,) was not
found in Eutomostethus (Blennocampinae) (Maki
1938).
Diprionidae (Fig. 7)
The plesiomorphic combination of muscles
and structures for Tenthredinoidea are present
but with the following apomorphic features: ante-
rior arms reduced to cup-like receptacles for muscle
124; anterior process of laterophragma expanded
and forming a broad attachment for 150a; and
muscle 150b originates medially to 150a on the
lateral furcal arms.
Volume 3. 1 994
255
Cimbicidae (Fig. 8)
The anterior arms are reduced to small spoon-
shaped processes, the posterior lobe of
mesopostnotum is fused to second phragma form-
ing attachment for a large muscle 116, and muscle
150b is lost [3:2]. The anterior process of the
laterophragma is enlarged and extends ventrally
into the cup-shaped lateral arm of the mesofurca
(autapomorphy).
Argidae + Pergidae
The lateral arms of mesofurca are tubular and
hollow (apomorphic), the anterior arms of the
mesofurca are absent, and the attachment site for
muscle 124 (f Uj-fiij) is to the rounded surface of the
lateral arms. Although they would suggest a sister
group relationship for these two families, these
characters were not surveyed throughout Hy-
menoptera and were not coded for the parsimony
analysis.
Argidae
The laterophragma and its associated muscle
attachments are as in Tenthredinidae.
Pergidae (Fig. 5)
This family displays a number of
autapomorphies: loss of the tergal depressor
muscle (116) [4:1] and the associated posterior
lobe of the laterophragma [2:2], and loss of one of
the furcal laterophragmal muscles (possibly 150b)
[3:?]. The laterophragma is reduced to a small
triangular lobe that ends in a narrow attachment
to muscle 150. The lateral arms form a cup-shaped
attachment for muscles 150 and fbl. A sclerotized
apodeme, probably derived from the lateral arm
and muscle 151 (pl,-fu,rt), extends laterally to the
pleural ridge. Ventrolateral extensions of the lat-
eral arms are developed as insertions for the en-
larged muscle 170 (pl2-fu2b).
Cephidae + remaining Hymenoptera
Figs. 9, 49, 52
Monophyly of the Cephidae and the remain-
ing Hymenoptera is supported by a suite of char-
acters: apical connection of the anterior mesofurcal
arms [1:3], formation of a pseudophragma [6:1],
and invagination of the mesopostnotum laterally
[10:1]. The pseudophragma is not present in all
Hymenoptera, but its complete absence in the
lower Hymenoptera and presence in Cephidae,
Anaxyelidae, Siricidae, Xiphydriidae and most
Apocrita suggest that it is a synapomorphy at this
level. There are no published accounts which make
reference to a pseudophragma occuring elsewhere
within Endopterygota.
Cephidae are plesiomorphic for other charac-
ters of the mesofurcal-mesopostnotal complex:
muscles 116, 150a, 150b, and fbl are present, and
the laterophragma is divided into a large posterior
lobe and a small anterior process. The
mesopostnotum is usually exposed medially but
is sometimes concealed by the scutellum. Muscle
114 passes externally over the mesopostnotum in
all genera (plesiomorphic) and may arise
dorsomedially from a vertical process on the
metatergum {Calameuta Konow, Ceplms Latrielle
[mp, Fig. 49], Monoloplopus Konow, Syrista Konow
and Tracheitis Jurine) as in Siricidae, or from an
emargination of the metatergum (Caenocephalus
Konow and Janus Stephens) as in Anaxyelidae.
Anaxyelidae + (Siricidae + (Xiphydriidae +
(Orussidae + Apocrita)))
Fig. 10
Monophyly of the Anaxyelidae and remain-
ing Hymenoptera is based on a more complete
fusion of the anterior arms of the mesofurca [1:4].
Fusion of the anterior mesofurcal arms in
Anaxyelidae (Fig. 10a) appears to be more com-
plete than in Siricidae (Fig. 11a) in which the arms
are fused in about the anterior half. The anterior
arms of Anaxyelidae (Fig. 10a) are still traceable in
dorsal view and they are connected posteriorly by
a thin plate of cuticle. The greater degree of fusion
of the arms is probably autapomophic in
Anaxyelidae; however, in Anaxyelidae and
Siricidae the fusion is intermediate between the
apical connection found in Cephidae and the com-
plete fusion including a strong furcal bridge found
in Xiphydriidae + Orussidae + Apocrita. In
Anaxyelidae, Siricidae and Xiphydriidae, muscle
150b inserts onto the posterior face of the
laterophragmal lobe rather than to the ventral
margin as in Xyeloidea, Megalodontoidea,
Tenthredinoidea and Cephoidea. As the muscle is
absent in Orussidae and Apocrita, the importance
of this unique attachment as a character for sup-
porting monophyly of Siricoidea (including
Anaxyelidae and Xiphydriidae) or Siricoidea +
256
Journal of Hymenoptera Research
Orussoidea + Apocrita cannot be assessed.
In Anaxyelidae, the presence of both furcal-
laterophragmal muscles (150a & 150b) and ante-
rior origin of muscle 150b on the f ureal arms is
plesiomorphic. The mesopostnotum is broadly
exposed medially and muscle 114 (t,-t3) passes
under the mesopostnotum medially (autapo-
morphy; 5:3). The dorsal enclosure of muscle 114
in Anaxyelidae may result from the extension and
medial fusion of the anterior margin of the
mesopostnotum. Rasnitsyn (1969, fig. 99) illus-
trates a split mesopostnotum [5:2] for the fossil
Anaxyela gracilis Martynov. This supports the con-
tention that an internal muscle 114 is derived
within Anaxyelidae.
Siricidae + (Xiphydriidae + (Orussidae +
Apocrita))
Fig. 11
No characters of the mesofurcal-
mesopostnotal complex demonstrate the mono-
phyly of Siricidae and the remaining Hymenoptera.
The anterior mesofurcal arms of Siricidae (Fig. 1 1 )
are vertically flattened and in dorsal view are
distinct along their entire length. The degree of
fusion of the anterior arms is intermediate be-
tween Cephidae and Xiphydriidae + Orussidae +
Apocrita, but, as discussed above, is comparable
to that found in Anaxyelidae. Siricidae are
autapomorphic for the loss of muscle 150a (along
with the anterior process of the laterophragma)
[3:3]. The posterior attachment of muscle 114 (t,-t,)
to a vertical process of the metanotum is shared
with some Cephidae but not with Xiphydriidae,
Orussidae or Apocrita, which lack a process. Inter-
nal passage of muscle 114 in Siricidae may result
from the posterior extension of the scuteOum rather
than internalization of the mesopostnotum as in
Cephidae and Tenthredinoidea.
Rasnitsyn (1988) proposed that Siricidae +
remaining Hymenoptera were monophyletic based
on four synapomorphies: 1) compound third an-
tennal segment reduced and subequal to the fol-
lowing segments, 2) head capsule with postgenae
subcontiguous, divided with narrow hypostomal
bridge, 3) transscutal articulation present, and 4)
prepectus concealed. In Anaxyelidae, the primi-
tive state of a compound third antennal segment is
found only in the fossil anaxyelid genus
Sphenosyntexis Rasnitsyn; the derived state is found
in the remaining genera of Anaxyelidae including
the only extant species, Syntexis libocedrii Rohwer.
However, the antennal flagellum is apomorphic
(reduced and divided) in Xyeloidea, Megalo-
dontoidea, Tenthredinoidea, Cephoidea and
within Anaxyelidae, and thus its reduction in
Siricidae and remaining Apocrita is not a reliable
character. The postgenae meet broadly along the
medial line to form a postgenal bridge (=genaponta
sensu Ross 1937) in Anaxyelidae, Siricidae and
Xiphydriidae; we agree with Ross (1937) that the
postgenae of Anaxyelidae and Xiphydriidae are
similar and thus Anaxyelidae cannot be excluded
from Siricoidea based on this character. Gibson
(1985) demonstrated that a transscutal articula-
tion is absent in Cephidae, Anaxyelidae and
Siricidae. The presence of a concealed prepectus is
useful for supporting the monophyly of Siricidae
+ Xiphydriidae + Orussidae + Apocrita (Rasnitsyn
1988; Gibson 1985).
Xiphydriidae + (Orussidae + Apocrita)
Fig. 12
There is strong support in the MF-MPN com-
plex for the monophyly of Xiphydriidae, Orussidae
and Apocrita. The anterior mesofurcal arms are
completely fused into an anterior furcal process
(fp) and posteriorly strengthened to form a trans-
verse mesofurcal bridge (fb) [1:4], with the excep-
tion of some Apocritan taxa in which the mesofurcal
bridge is lost. In addition, the mesopostnotum is
completely internal medially [5:4] and laterally
[10:2], and the laterophragma is strongly incised
with the anterior process (ap) developed into a
strong apodeme [2:1]. The plesiomorphic combi-
nation of muscles in the MF-MPN complex is
retained. Among the Cephoidea, Siricoidea and
Orussoidea, Xiphydriidae are unique in having
muscle 150b attached to the lateral arms of the
mesofurca (versus the anterior arms or furcal pro-
cess). Muscle 124 (fu^-fUj) is broadly attached to
the furcal process and bridge. In Xiphydriidae, the
tendon of muscle 114 (t2-t3) is enclosed medially,
and internally, by the cuticle of the second phragma
(autapomorphic).
Orussidae + Apocrita
Fig. 13
No synapomorphies were found in the
Volume 3, 1994
257
mesofurcal-mesopostnotal complex that support
the monophyly of Orussidae + Apocrita without
Xiphydriidae. It is possible that the loss of muscle
150a supports the monophyly of Orussidae +
Apocrita [3:? = 3:2] . However, the homology of the
remaining muscle 150 in Orussidae is uncertain
(see discussion of Character 3). Therefore, it is also
possible that muscle 150b is lost in Orussidae and
150a is lost in Apocrita.
Orussidae have at least three apomorphic fea-
tures in the MF-MPN complex, two of which are
also found in Pergidae and Cimbicidae. The furcal
bridge (fb) is rounded and bowed anteriorly with
a distinct median process (fp) [1:6]. The attach-
ment of muscle 150 to the furcal process is found
only in Orussidae. In Apocrita, muscle 150a is
usually attached to the furcal bridge or lateral arm.
Otherwise the furcal bridge and process in
Orussidae are similar to Xiphydriidae and other
Apocrita. Muscle 116 is lost [4:1] in Orussidae, and
there is an associated reduction of the
laterophragma to a small triangular lobe that forms
the attachment for muscle 150 [2:2]. Additional
autapomorphies in Orussidae include loss of the
pseudophragma [6:0] and the configuration of
muscle 180 (fu3-sps2) which arises from the spina
as a single narrow tendon that bifurcates about
half-way along its length to the metafurcal arms
(the plesiomorphic configuration is completely
separated along its entire length).
Apocrita
Figs. 14-19, 22-46, 53, 54
The loss of two muscles in the MF-MPN com-
plex supports the monophyly of the Apocrita: the
metafurcal-spina muscle (180, fu3-sps2) [8:1] and
the furcal-basalare muscle (fbl, fu2-ba3) [9:0].
Muscle 150b (fu2-pn2a) [3:2] is lost in all Apocrita
including those that retain the posterior lobe of the
laterophragma. However, as discussed above, the
homology of the remaining muscle 150 in
Orussidae is not clear, and therefore loss of 150a is
either a synapomorphy of Orussidae + Apocrita or
an autapomorphy of Apocrita. Loss of musclel37
may also support the monophyly of Apocrita (see
discussion for Character 2).
Shcherbakov (1981) stated that muscle 170
(pl2-fu2b, s-cx2) is connected to the coxal process of
the mesopleuron in all Hymenoptera. Based on
our observations, it is well developed in all
Symphy ta and attaches laterally to the mesofurcal
arms (Figs. 3, 5, 9b, 12b, 13). In Apocrita, the
position of muscle 170 shifts from the apex of the
furcal arms to the base of the furca and is more
nearly horizontal. This is most apparent in
Vespoidea (cf. Duncan 1939, muscle IIfpl2 (71)).
The laterophragma of Cynipoidea, some
Diapriidae, Monomachidae (Fig. 14) and
Vanhorniidae (Fig. 15a) are similar in structure
with an anterior apodeme (ap) narrowly sepa-
rated by a deep incision from the posterior lobe
(pn,) [1:5], which supports muscle 116 [4:0]. In
Megaspilidae (Fig. 25), the apodeme and lobe may
be broadly or narrowly separated. Muscle 116
[4: 1 ] is present only in the families listed above and
probably has been lost numerous times within
Apocrita (Gibson 1985). It is unlikely that this
muscle and associated posterior lobe could be
regained once lost. Under the present hypothesis
of relationships in Apocrita, muscle 116 and the
lobe are lost at least 9 times (10 if also lost in
Austroniidae) (Fig. 56). Of these, only in Diapriidae
is the muscle lost within an entire family.
Groundplan states for the Apocrita are exem-
plified by Monomachidae (Fig. 14), Vanhorniidae
(Fig. 15), most Diapriidae (Fig. 29) and Cynipoidea
(Fig. 22), in which muscle 116 and the posterior
lobe of the laterophragma are retained (Daly 1963;
Gibson 1985). Plesiomorphic attributes of Apocrita
include l)presence of a median furcal process (fp)
forming the attachment for muscle 124, 2) furcal
bridge complete (fb) [1:4], 3) muscle 150a originat-
ing laterally on the bridge or the lateral arm of the
mesofurca, 4)laterophragma divided into an ante-
rior apodeme and posterior lobe [2:1], 5)presence
of the tergal depressor muscle (116) [4:0], 6)
mesopostnotum internal [10:2] and 7) prephragma
present [6:1]. Ceraphronoidea (Fig. 25) have lost
the mesofurcal bridge [1:7] but retain a divided
laterophragma, and in some Diapriidae (Fig. 22)
the posterior lobe of the laterophragma is lost [4:1]
Apart from changes in the laterophragma,
apomorphic changes of the MF-MPN complex in
Apocrita include loss of the mesofurcal bridge
[1:7] and changes in the structure of the anterior
apodeme (=axillary lever) of the laterophragma
[2:3-6]. Changes in the shape of muscle 150a, lat-
eral arms, furcal bridge and second phragma are
homoplastic and probably related to changes in
thoracic shape. Notable exceptions in Aculeata
include development of an axillary lever [2:5] that
258
Journal of Hymenoptera Research
is separated in Apiformes [2:6], fusion of the lat-
eral arms of the mesofurca and metafurca [11:1],
development of processes on the mesofurca for
the furcal-trochanteral muscle [12:1,2], and a me-
dial reduction of the attachment points (60 & 61)
between the second phragma and terga.
The cladogram presented in Figure 56 is con-
sistent with the relationships proposed by
Rasnitsyn (1988) and Brothers and Carpenter ( 1 993)
for extant Apocrita, and is used as the model for
examination of character state change within
Apocrita. Apart from the modif ications in Aculeata,
the MF-MPN complex offers little support for
phylogenetic relationships proposed for Apocrita.
Within Apocrita a total of 22 steps are required to
explain the distribution of character states as mod-
elled on Figure 56 (excluding changes in the
Chalcidoidea + Platygastroidea + Mymarom-
matoidea trichotomy, Austroniidae and
Peradeniidae). Given that the tergal depressor
muscle and posterior lobe of the laterophragma
are present [4:0] in what are assumed to be rela-
tively derived groups (e.g., Cynipoidea, Diapriidae
and Monomachidae), this complex must be lost a
minimum of 9 times (or 10 times if it is absent in
Austroniidae). Only 8 losses are required if
Pelecinidae and Vanhorniidae are monophyletic
(Fig. 56). An additional 3 losses of muscle 116 [4:1]
are required using Konigsmann's (1978a) hypoth-
esis for Apocrita. Obviously, loss of this muscle
does not offer strong evidence for the monophyly
of Megalyroidea, Stephanoidea and Trigo-
nalyoidea (Fig. 56).
Among the parasitic families of Apocrita, the
mesofurcal complex of Gasteruptiidae (Fig. 26)
and Evaniidae (Fig. 27) were very different from
each other. However, the mesofurcal bridge of
Aulacidae and Evaniidae are similar in shape
(broad and flattened with muscle 124 attaching
under the anterior margin of the bridge). In all
three families the axillary lever is strongly in-
flected and perpendicular to the lateral wall of the
mesosoma (Figs. 26, 27), and the lever is connected
by a long thin tendon to muscle 150a. Scelionidae
and Chalcidoidea both have a similar mesofurcal
bridge (bowed with the median process virtually
absent) which may indicate a sister-group rela-
tionship between Chalcidoidea and Platy-
gastroidea. However, given that the bridge is lost
in Platygastridae, Mymarommatidae, Mymaridae
and some Chalcidoidea [1:7], it is not clear what
the groundplan state of this character is in either
superfamily. Also, the axillary lever is deflected
[2:3] in Platygastridae, some Scelionidae and most
Chalcidoidea. Almost all Chalcidoidea including
Mymaridae are unique in that muscle 150a at-
taches to the entire length of the deflected axillary
lever (versus only apical attachment).
Mymarommatidae have a medially inflected lever
[2:1] and muscle 150a is conical and attaches only
to the apex of the lever (plesiomorphic).
Proctotrupidae (Fig. 32) have a similiar axillary
lever to that of Platygastridae (Fig. 28), although it
is shorter and not as strongly deflected (Figs. 29-
32). In Heloridae (Fig. 30) and Pelecinidae (Fig. 31),
the lever is followed by a short spine (pn,?) that
has no muscle connection and may be the remnant
of the posterior lobe of the laterophragma. Some
Proctotrupidae have a blunt triangular lobe in a
homologous position. Heloridae, Pelecinidae and
Proctotrupidae are the only Hymenoptera with a
vestigial posterior lobe of the mesopostnotum.
Ropronndae were identical to Heloridae inter-
nally except for the lack of a vestigial posterior
lobe.
Most Ichneumonoidea (Fig. 17) have a
mesofurca consisting of a broad mesofurcal bridge
and a strong medial projection. The furcal bridge
of Hybrizontinae (Braconidae) (Fig. 33) lacks a
median process but this could be autapomorphic.
The axillary lever in Ichneumonidae is similar to
Vespoidea and Apoidea [2:5]. This may be evi-
dence for the close relationships to Aculeata that
have been proposed (Rasnitsyn 1988, Sharkey and
Wahl 1992), but a similar lever has not been found
in Braconidae or Chrysidoidea.
Chrysidoidea do not have any defining
apomorphies of the mesofurca or axillary lever.
The arched bridge of Plumariidae (Fig. 34),
Sclerogibbidae (Fig. 35) and Embolemidae (Fig.
36) are similar but this feature may be
plesiomorphic, as this shape is similar to
Stephanidae. The furcal bridge of the single fe-
male dryinid examined (Anteoninae) was similar
to Bethylidae (Fig. 38) (relatively straight with a
strong apical projection), and the bridge of the
male dryinid (Fig. 35) was reduced to a straight
bar. Obviously more taxa need to be surveyed to
understand the importance of furcal shape in
Chrysidoidea. Chrysididae have apomorphies that
distinguish taxa at the subfamily level: the fusion
of the lateral arms of the mesofurca and metafurca
Volume 3, 1 994
259
in Amiseginae (Fig. 39), and extensions of the
furca that support the furcal depressor of the
trochanter [12:1] in Chrysidinae (Fig. 40) and
Cleptinae. These latter two subfamilies also share
a reduction of the axillary lever [2:4]. Reduction of
the axillary lever and presence of ventral pro-
cesses in Cleptinae and Chrysidinae do not sup-
port the hypothesis of relationships among
Chrysididae proposed by Kimsey and Bohart
(1990) of Cleptinae + (Amiseginae + Chrysidinae).
Based on relationships for Chrysidoidea proposed
by Brothers and Carpenter (1993), Carpenter (1986)
and Rasnitsyn (1988), the plumariid type of bridge
is plesiomorphic, Dry inidae have an independently
derived bridge, and Chrysididae + Bethylidae have
a relatively broad, straight furcal bridge with a
strong medial projection.
Monophyly of the Vespoidea and Apoidea is
supported by the development of an axillary lever
[2:6], which is subsequently reduced in
Bradynobaenidae [2:4] and modified in Formicidae
[2:1]. Most Aculeata also have a broad, flattened
furcal bridge with muscle 124 attaching to the
dorsal and lateral surfaces (Fig. 18), although the
bridge is not as well developed in Rhopalo-
somatidae (Fig. 19), Sierolomorphidae and some
Sphecidae (Fig. 46). In Tiphiidae (Fig. 44), the furca
is convoluted and broadly excavated dorsally.
The furcal bridge has several shapes in Vespoidea
but their importance for assessing relationships
will require a much broader survey of taxa. A
strong ventral process (vp) for the furcal depres-
sor of the trochanter (174, fu,-tr,) confluent with
the furcal bridge [12:2] is present in Sapygidae,
Sierolomorphidae, Pompilidae (Fig. 42), Scoliidae
and Vespidae (Fig. 41). Under the most recent
hypothesis of relationships for Vespoidea (Broth-
ers and Carpenter 1993), the ventral process is
derived independently in Sapygidae, Pompilidae
and Scoliidae + Vespidae, although several mod-
els of state change are possible. Mutillidae (Fig.
18), Rhopalosomatidae (Fig. 19) and Bradyno-
baenidae have apparently retained muscle fbl
(fu2-ba3) [9:1] and have a characteristically shaped
mesofurca, however these families are not regarded
as sister taxa (Brothers 1975, Carpenter and Broth-
ers 1993).
Apoidea have been separated into two groups,
the Apiformes and the Spheciformes (Goulet and
Huber 1993). Heterogynaidae have been included
either within the Spheciformes (see Goulet and
Huber 1993) or as the sister group of Spheciformes
+ Apiformes (Carpenter and Brothers 1993).
Apoidea (Figs. 45, 53), including Heterogynaidae,
have the arms of the mesofurca and metafurca
fused at the junction with the mesofurcal bridge
and muscle 181 (fu3-fu2) is lost [11:1]. This feature
is unique in Hymenoptera. In Vespoidea (Figs. 41,
42, 44), Spheciformes (Fig. 46) and Heterogynaidae,
the base of the axillary lever is broadly fused to the
lateral arm of the mesopostnotum. All Apiformes
have the axillary lever separated from the
mesopostnotum as an independent sclerite [2:6]
(Fig. 45) (Snodgrass 1942).
PARSIMONY ANALYSIS
The mesofurcal-mesopostnotal complex rep-
resents a single system of inter-related characters,
and it should not be used alone to form new
hypotheses of relationships. However, it is of in-
terest to determine if our interpretations of charac-
ter state evolution are in fact the most parsimoni-
ous for these data. We tried two approaches to
explore this question. First, we superimposed the
data in Appendix 2 on existing hypotheses of
relationships. PAUP Version 3.0s (Swofford 1985)
was used to optimize the character state data in
Appendix 2 on Figure 55 for Symphyta and Fig-
ures 55 and 56 for the entire Hymenoptera (using
one fully resolved tree for Apocrita based on
Rasnitsyn (1988) and Brothers and Carpenter
(1993)). Ibaliidae was used to represent the
groundplan of Apocrita in the former case, and
each matrix included the outgroup (Appendix 2).
All most parsimonious models of character state
evolution were examined on these tree topologies.
Characters 1 and 4 are sensitive to assump-
tions about transformation series. As discussed
above (Character Analysis), it seems most reason-
able to assume that character 1 can evolve from
state 0 to state 1 (unique for Hymenoptera), from
state 1 to state 2 or state 3, from state 3 to state 4 and
from state 4 to state 5; states 6 and 7 are indepen-
dent derivations from state 5. Character 1 was
ordered using a character state tree (or "stepmatrix"
in PAUP) that specified the above transformation
series. As discussed earlier, character 4 is treated
as irreversible. The assumptions of additive cod-
ing for characters 1 and 4 are referred to as the
Ordered Characters. Other characters were al-
ways treated as nonadditive (unordered).
260
Journal of Hymenoptera Research
For Symphyta, the models of character state
change shown in Figure 55 are in fact the most
parsimonious for that tree topology. 27 steps are
required to explain the data (indices of homoplasy
are not possible for characters ordered using step
matrices). Different models of state change are
possible for character 3 (because it is scored as
missing in Pergidae and Orussidae) and character
5 (unknown polarity for ingroup). For character 5,
the presence of state 2 in fossil Xyelinae argues that
this condition is groundplan for Hymenoptera,
but it is equally parsimonious (at least for extant
taxa) to assume that state 1 is groundplan and state
2 arises in the interval below Macroxyelinae and
remaining Hymenoptera (dashed line, Fig. 55).
The latter solution would make Xyelidae
paraphyletic. Treating all characters as nonaddi-
tive (unordered) does not change the tree length
(27 steps, consistency index 0.81, retention index
0.77). In fact, the only effect is to allow for a number
of alternate models of state change for character 1 .
In this case, it is equally parsimonious to assume
any of states 0, 1, 2 or 3 in the nodes below
Tenthredinoidea + remaining Hymenoptera and
Cephidae + remaining Hymenoptera, a result that
essentially discards the information in the trans-
formation series. Optimizing the ordered data
onto alternate hypotheses of Symphyta resulted in
longer trees. Rasnitsyn's (1988) tree, with
Macroxyelinae as sister group to Tenthredinoidea,
is one step longer, because of the parallel loss of
muscle 127 [7:1]. Konigsmann's (1977) tree, with
Symphyta monophyletic and Cephidae as sister
group to Apocrita, requires 37 steps (character 3
optimized).
For Hymenoptera, similar results were ob-
tained for optimizing the Ordered Character data
onto one tree, which is summarized for Apocrita
in Figure 56. 58 steps were required to explain the
data. Different optimizations were possible for
Character 1 forChalcidoidea + Mymarommatoidea
+ Platygastroidea (if parallel loss of mesofurcal
bridge [1:7] is preferred), and Character 9 in
Vespoidea. In the former case, although it is equally
parsimonious to assume loss of the mesofurcal
bridge (1:7) followed by a gain [1:6], parallel loss of
the bridge seems more likely. Likewise, it seems
more likely to postulate a parallel loss of the f urcal-
basalare muscle [9:0] in Vespoidea. Treating all
characters as nonadditive decreases the length of
the hymenopteran tree by 5steps because of a
different optimization for Character 4 (treated as
irreversible in the ordered analysis) and reversals
(4:1 to 4:0) are favoured. For this result, the loss of
the posterior lobe of the laterophragma and asso-
ciated muscle 1 16 (4:1) would be a synapomorphy
of Orussidae and Apocrita and then the presence
of both structures (4:0) would have to be regained
at least 4 times within Apocrita. Although this
may be a more parsimonious solution in terms of
the number of steps, we feel it is an unacceptable
model of character evolution and the assumption
of irreversibility should be invoked.
Parsimony analysis using the branch and
bound algorithm (Symphyta) or branch-swapping
(Hymenoptera) was then performed on these data
to determine if another tree topology would result
in a more parsimonious solution. When characters
are ordered as above, for Symphyta, 24 trees of 27
steps result (after derooting, condensing and
rerooting the resulting trees in PAUP, which is
necessary when using directed characters). Cod-
ing Anaxyelidae as state 1 (homoplastic) or state 3
(autapomorphic) had no effect on the number of
steps or resulting trees. As should be apparent
from Figure 55, the basal taxa (Xyelinae,
Macroxyelinae and Pamphiliidae) are unresolved
in each of the 24 trees. Tenthredinoidea is always
resolved as monophyletic but with little or no
further resolution and Cephidae is consistently
placed as sister group to remaining Hymenoptera,
as in Figure 55. Anaxyelidae and Siricidae form a
trifurcation with Xiphydriidae + Orussidae +
Apocrita while the latter three taxa are essentially
unresolved. In other words, there is no more par-
simonious interpretation of these data and the
most parsimonious solutions are congruent with
the data shown on Figure 55. When characters are
unordered for Symphyta, 138 trees of 26 steps
result. The strict consensus solution for these trees
is almost completely unresolved. For Hy-
menoptera, the lack of character support in
Apocrita resulted in a considerably shorter tree
(40 steps for the Ordered Characters) but also
provided no resolution of Apocritan relationships;
the structure of the Symphyta portion remained
unchanged.
CONCLUSIONS
The mesofurcal-mesopostnotal complex is a
conservative character system that generally ex-
Volume 3. 1 994
261
hibits few changes within families of Hymenoptera
but undergoes considerable change at higher taxo-
nomic levels. The most significant changes occur
in the Symphyta, but additional characteristics of
the mesofurcal-mesopostnotal complex provide
characters that are useful in inferring relation-
ships within Apocrita.
Of the competing hypotheses for relationships
of Symphyta, those in Figure 55 are the most
parsimonious using the hypotheses of character
state transformation discussed in this paper. Exact
search methods using additive (ordered) charac-
ters reproduce these relationships with the same
number of steps and identical indices of character
homoplasy and fit. Siricoidea are recognized as a
paraphyletic group with respect to Apocrita. If
Xyelidae are paraphyletic with the rest of Hy-
menoptera (Rasnitsyn 1980, 1988) and
Macroxyelinae + Tenthredinoidea are monophyl-
etic, then a minimum of one extra step is required
for character 7 (loss of fUj-spSj). In sharp contrast,
a monophyletic grouping for Symphyta with
Cephidae as sister group to the Apocrita
(Konigsmann 1977) requires 11 additional steps
for the same data. The phy logenetic hypothesis for
Apocrita proposed by Rasnitsyn (1988; Fig. 56)
was shorter than Konigsmann's by only three
steps (for character 4). Hypotheses based on parsi-
mony are much shorter for Apocrita, even when
characters are ordered, but these hypotheses
should not be accepted until they can be included
in a larger data set with other character systems.
Monophyly of Orussidae + Apocrita (Vespina
se nsu Rasnitsyn) is supported by 10 external char-
acters of adults (Rasnitsyn 1988) and a shift of the
furcal-coxal muscles from a posterior to an ante-
rior attachment on the discriminal lamella of the
mesosoma (Johnson 1988). No features of the
mesofurcal-mesopostnotal complex discussed here
provide support for the monophyly of Orussidae
+ Apocrita, without Xiphydriidae. The mesofurca
and laterophragma of Orussidae is highly derived
and the only potential synapomorphy for Vespina
(Character 3: loss of muscle 150b, fu2-pn, ) is prob-
ably not homologous. The lack of 2ph-3ph in
Siricidae and Xiphydriidae has been proposed as
a potential synapomorphy for Xiphydriidae +
Siricidae (Whitfield et al. 1992), but this muscle has
also been lost independently in Goniozus
(Bethy lidae), Cleptes (Chrysididae), Formicidae and
Ampulex (Sphecidae) (Daly 1963). Monophyly of
Xiphydriidae, Orussidae and Apocrita is supported
by four synapomorphies. No evidence in the
mesofurcal-mesopostnotal complex supports the
inclusion of Xiphydriidae or Orussidae within
Siricoidea or the monophyly of Anaxyelidae +
Siricidae.
The monophyly of Apocrita is supported based
on losses of the muscles fu2-pn, , fu3-sps2, fu2-ba3,
pl2-fu2b, and probably t -ba . Using the evolution-
ary scheme proposed by Rasnitsyn (1988), charac-
ters of the mesofurca and mesopostnorum were
less useful for demonstrating relationships within
Apocrita. Taxa which have retained the posterior
lobe of the laterophragma and the associated
muscle 116 are treated by Rasnitsyn (1988) as
relatively derived taxa (especially Diapriidae and
Cynipoidea). Rasnitsyn's hypothesis requires at
least nine independent losses of character 4, if loss
of both lobe and muscle is irreversible. This ap-
pears to be a poor character for postulating rela-
tionships. Loss of the mesofurcal bridge [1:7] could
indicate support for the monophyly of
Ceraphronoidea, but it is homoplastic within both
Chalcidoidea and Platygastroidea. Some changes
in shape of the axillary lever or mesofurcal bridge
within Apocrita are difficult to categorize or to
place into distinct transition series. Distinctive
features such as the vestigial posterior
laterophragmal lobe of Pelecinidae, Heloridae and
Proctotrupidae are reductions and possibly con-
vergent. The development and separation of the
axillary lever, fusion of the mesofurca and
metafurca, and the retention (or redevelopment)
of muscle fbl provide strong support for relation-
ships within the Aculeata.
Changes in the mesofurcal-mesopostnotal
complex probably result from an increased em-
phasis on the fore wings for flight, reduction of the
metathorax and fusion of the first abdominal seg-
ment. Although the indirect flight muscles are the
main wing depressors, the direct muscles of the
mesothorax control flight through modifications
of the pronation and rotation of the wings during
the downstroke and by controlling the tension of
the longitudinal flight muscles, thereby affecting
the amplitude of the wing beat (Pringle 1957, 1960,
1961 ). In Apocrita, the axillary lever acts to turn the
apex of the mesophragma and the associated fourth
axillary sclerite (Weber 1925). This is a shift in
function from the lower Hymenoptera where
muscles 150a and 150b twist the posterior lobe of
262
Journal of Hymenoptera Research
the laterophragma, initiating a greater force upon
the indirect phragmal flexor (muscle 116, t;-pn2).
The excision of the laterophragma in Xiphydriidae
would reduce the effect of the anterior apodeme (=
axillary lever) on the posterior lobe. Pringle (1960,
1961 ) proposed that the pleurosternal and axillary
lever muscles control the power generated by the
indirect flight muscles and thus the amplitude of
the wing beat. The flexibility of the axillary lever
reaches a maximum in the Apiformes in which
complete separation from the laterophragma re-
sults in more powerful leverage upon the associ-
ated axillary sclerites of the forewing (Snodgrass
1942).
Homologies may not be clear in taxa from
opposite ends of a phylogenetic tree. For example,
Weber (1925) discusses the mesofurcal ring
(=mesofurcal bridge) of Siricidae and Vespidae as
non-homologous. Superficially they do look dif-
ferent, but by examining the intermediate stages
the homologies can be verified. Changes in the
mesofurca and mesopostnotum provide a num-
ber of characters that show informative transitions
from the plesiomorphic states in Xyelidae, but
only by comparing a large number of taxa do these
transformation series become apparent. Nonaddi-
tive multistate characters are not as useful in build-
ing a classification because unlinked character
states support only their inclusive members. How-
ever, a priori ordering of transformation series are
dependent on previous classifications or on the
presumed homology of sometimes very divergent
character states. For example, placement of
Cephidae as sister group to the Apocrita led to a
misinterpretation of the hypostomal bridge by
Ross (1937) and the mesofurcal bridge was not
discussed as an important character by Rasnitsyn
(1969, 1980, 1988). Present interpretations of
changes in the mesofurca and mesopostnotum
would have made little sense without the direc-
tion provided from new classifications for
Symphyta proposed by Gibson (1985) and
Rasnitsyn (1988). The current distribution of char-
acter states of the mesofurca and mesopostnotum
within Apocrita seems incongruent with other
character information. Perhaps as more characters
are used to construct improved classifications of
Apocrita, this will also change.
ACKNOWLEDGEMENTS
Gary Gibson and two anonymous reviewers provided
valuable comments and criticisms. We thank Henri Goulet,
John Huber, Lubomir Masner and especially Gary Gibson
(all CNC) for providing specimens and guidance. Jeff
Cumming (CNC) and Dave Smith (USNM) provided speci-
mens of miscellaneous families that were important addi-
tions to the material examined. This study was supported by
Hatch Project TEXO-6705 of J.B.W. for initial work carried
out at Texas A&M University and an NSERC grant to D.C.D.
for subsequent work done in Ottawa.
LITERATURE CITED
Alam, S.M. 1951. The skeleto-muscular mechanism of
Stenobracon deesae Cameron (Bracomdae, Hymenoptera)
- An ectoparasite of sugarcane and juar borers of India.
Part I. Head and thorax. Aligarh Muslim University Pub-
lications (Zoological Series) 3: 76 pp. + 9 pis.
Brodskiy, A.K. 1992. Structure, function, and evolution of the
terga of wing-bearing segments of insects. II. Organiza-
tional features of the terga in different orders of insects.
Entomological Review 71: 8-28.
Brothers, D. J. 1975. Phvlogeny and classification of the
aculeate Hymenoptera, with special reference to
Mutillidae. Kansas University Science Bulletin 50: 483-
648.
Brothers, D. J. and J. M. Carpenter. 1993. Phylogeny of
Aculeata: Chrysidoidea and Vespoidea. Journal of Hy-
menoptera Research 2: 227-304.
Bucher, G. E. 1948. The anatomy of Monodontomerus dentipes
Boh., an entomophagous chalcid. Canadian journal of
Research (D) 26: 230-281.
Carpenter, J. M 1986. Cladistics of the Chrysidoidea (Hy-
menoptera). journal of the New York Entomological Society
94: 303-330.
Chapman, R. F. 1992. General anatomy and function, pp. 33-
67. In The Insects of Australia, 2nd edition. Volume 1.
Melbourne University Press, Carlton. 1137 pp.
Daly, H. V. 1963. Close-packed and fibrillar muscles of the
Hvmenoptera. Annals of the Entomological Society of
America 56: 295-306.
Daly, H. V. 1964. Skeleto-muscular morphogenesis of the
thorax and wings of the honey bee Apis mellifera (Hy-
menoptera: Apidae). University of California Publications
in Entomology 39: 77 pp.
Duncan, C. D 1939. A contribution to the biology of North
American vespine wasps. Stanford University Publica-
tions (Biological Sciences) 8 (1): 272 pp.
Gauld, I. and B. Bolton. 1988. The Hymenoptera Oxford Uni-
versity Press, New York. 332 pp.
Gibson, G. A. P. 1985. Some pro- and mesothoracic structures
important for phylogenetic analysis of Hvmenoptera,
with a review of terms used for the structures. Canadian
Entomologist 117: 1395-1443.
Gibson, G. A. P. 1986. Mesothoracic skeletomusculature and
mechanics of flight and jumping in Eupelminae (Hy-
menoptera, Chalcidoidea: Eupelmidae) Canadian Ento-
mologist 118:691-728
Volume 3, 1994
263
Gibson, G. A. P. 1993. Groundplan structure and homology
of the pleuron in Hymenoptera based on a comparison
of the skeletomusculature of Xyelidae (Hymenoptera)
and Rhaphidndae (Neuroptera). Canadian Entomologist
165: 165-187.
Huber, J. T. and H. Goulet. 1993. Families of Hymenoptera of the
World. Research Branch, Agriculture Canada. Publication
1894/E. 668 pp.
James, H. C. 1926. The anatomy of a British phytophagous
chalcidoid of the genus Harmolita {Isosoma). Proceedings
of the Zoological Society of London [1926] 75-182.
Johnson, N.F. 1988. Midcoxal articulations and the phylog-
eny of the order Hymenoptera. Annals of the Entomologi-
cal Society of America 81: 870-881.
Kelsey, L.P. 1957. The skeleto-motor mechanism of the Dob-
son Fly, Corydalus cornutus. Part II. Pterothorax. Mem-
oirs of the Cornell University Agricultural Experimental
Station 346: 31 pp. + 55 figs.
Kimsey, L. and R.M. Bohart 1990. The Chrysidid Wasps of the
World. Oxford University Press: Oxford. 652 pp.
Korn, W. 1916. Die muskulatur des Kopfes und des thorax
von Myrmeleon europaes und ihr metamorphose.
Zoologische Jahrbucher 68: 273-330.
Konigsmann, E. 1977. Das phylogenetische System der Hy-
menoptera. Teil 2: "Symphyta." Deutche Entomologische
Zeitschrift (N.S ,) 24: 1-40.
Konigsmann, E. 1978a. Das phylogenetische System der Hy-
menoptera. Teil 3 "Terebrantes" (Unterordung
Apocrita). Deutche Entomologische Zeitschrift (N.S. )25: 1-
55.
Konigsmann, E. 1978b. Das phylogenetische System der Hy-
menoptera. Teil 4 Aculeata (Unterordung Apocrita).
Deutche Entomologische Zeitschrift (N.S.) 25: 365-435.
Knstensen, N.P. 1992. Phylogeny of extant Hexapods. pp.
125-140. In The Insects of Australia, 2nd edition. Volume
1. Melbourne University Press, Carlton. 1137 pp.
Lawrence, J.F., E.S. Nielson and I.M. Mackerras. 1992. Skel-
etal anatomy and key to orders, pp. 3-32. In The Insects of
Australia, 2nd edition. Volume 1. Melbourne University
Press, Carlton. 1137 pp.
Maki, T. 1938. Studies on the thoracic musculature of insects.
Memoirs of the Faculty of Science and Agriculture, Taihoku
Imperial University 24: 343 pp. + 17 pis.
Markl, H. 1966. Peripheres Nervensystem und Muskulatur
lm Thorax der Arbeiterin von Apis mellifera L., Formica
polyctena Foerster und Vespa vulgaris L., und der
Grundplan der Innervierung des Insektenthorax
Zoologische Jarbucher 83: 107-184.
Matsuda, R. 1970. Morphology and evolution of the insect
thorax. Memoirs of the Entomological Society of Canada 76:
431 pp.
Michener, CD., 1944. Comparative external morphology,
phylogeny, and a classification of the bees (Hy-
menoptera). Bulletin of the American Museum of Natural
History 82: 157-326.
Naumann, I.D. 1992. Hymenoptera. pp. 916-1000. In The
Insects of Australia, 2nd edition. Volume 1. Melbourne
University Press, Carlton. 1137 pp.
Naumann, I.D. and L. Masner. 1985. Parasitic wasps of the
proctotrupoid complex: a new family from Australia
and a key to world families (Hymenoptera:
Proctotrupoideasensu lata). Australian journal of Zoology
33: 761-783.
Pnngle, J. W. S. 1957. Insect Flight. University Press, Cam-
bridge. 132 pp.
Pnngle, J. W. S. 1960. The function of the direct flight muscles
in the bee. Proceedings of the 1 2th International Congress of
Entomology p. 660.
Pnngle, J. W. S. 1961. The flight of the Bumblebee Natural
History 70 (7): 20-29.
Rasnitsyn, A. P. 1969. [Origin and evolution of the lower
Hymenoptera.) Trudy Paleontologicheskogo Instituta,
Akademiya Nauk SSSR 123: 196 pp.
Rasnitsyn, A. P. 1980. [Origin and evolution of Hymenoptera. ]
Trudy Paleontologicheskogo Instituta, Akademiya Nauk SSSR
174: 190 pp.
Rasnitsyn, A. P. 1988 An outline of evolution of the hy-
menopterous Insects (Order Vespida). Oriental Insects
22: 115-145.
Ronquist, F. and G. Nordlander. 1989. Skeletal morphology
of an archaic cynipoid, Ibalia rufipes (Hymenoptera:
Ibaliidae). Entomologica Scandinavica, Supplement 33: 60
pp.
Ross, H. H. 1937. A generic classification of the Nearctic
sawflies (Hymenoptera, Symphyta). Illinois Biological
Monographs 34: 173 pp.
Saini, M.S., S.S. Dhillon, and R. Agarwal. 1982.
Skeletomuscular differences in the thorax of winged
and non-winged forms of Camponotus camelinus (Smith)
(Hym., Formicidae). Deutche Entomologische Zeitschrifte
(N.S.) 29: 447-458.
Sharkey, M. J. and D. B. Wahl. 1992. Cladistics of the
Ichneumonoidea (Hymenoptera). Journal of Hymenoptera
Research 1: 15-24.
Shcherbakov, D. E. 1980. [Morphology of the pterothoracic
pleura in Hymenoptera. 1. Groundplan]. Zoologicheskn
Zhurnal 59: 1644-1653.
Shcherbakov, D. E. 1981. [Modifications of the pterothoracic
pleura in Hymenoptera. 2. Modtficat ions of the groundplan].
Zoologicheskii Zhumal 60: 205-213.
Snodgrass, R. E. 1910. The thorax of Hymenoptera. Proceed-
ings of the United States National Museum 39: 37-91 + 16
pis.
Snodgrass, R. E. 1927. Morphology and mechanism of the
insect thorax. Smithsonian Miscellaneous Collections 80
(1): 108 pp.
Snodgrass, R. E. 1942. The skeleto-muscular mechanisms of
the honey bee. Smithsonian Miscellaneous Collections 103
(2) 120 pp.
Tulloch, G. S. 1935. Morphological studies of the thorax of the
ant. Entomologica Americana 15: 93-130.
Weber, H. 1925. Derthorax derHornisse. Zoologische jarbucher
67: 1-100 + 4 pis.
Weber, H. 1927. Die Gliedrung der Sternalregion des
Tenthredinidenthorax. Ein Beitrag zur vergleichenden
Morphologie des Insecktenthorax. Zeitschrift fur
ivissenschaftliche Insektenbiologie 22: 161-198 + 6 pis.
Whitfield, J. B. 1992. Phylogeny of the non-aculeate Apocrita
and the evolution of parasitism in the Hymenoptera.
journal of Hymenoptera Research 1: 3-14.
Whitfield, J. B„ N. F. Johnson and M. R. Hamerski. 1989.
Identity and phylogenetic significance of the
264
Journal of Hymenoptera Research
metaposmotum in nonaculeate Hymenoptera. /birni/so/ Zessin, W. 1985. Neue oberliassiche Apocnta und die
the Entomological Society of America 82: 663-673. Phylogenie der Hymenoptera (Insecta, Hymenoptera).
Deutsche Entomologische Zeitschrift, N.S. 32:129-142.
Appendix 1 List of abbreviations used for figures. Abbreviations used by Ronquist and Nordlander (1989) in brackets.
af
ap
ex
dl
fb
fp
fu,
FWB
hP
HWB
If
pn2
anterior furcal arm [91] PN,
anterior process/axillary pph
lever of laterophragma (pn;) sps
cenchrus t,
coxa t3
discrimenal lamella [dl] ume
mesofurcal bridge vp
mesofurcal process [91] lph
metafurca 2ph
fore wing base 60
horizontal plate of mesofurca [90]
hind wing base hi
lateral furcal arm [fu;]
laterophragma of mesopostnotum
(posterior face of lobe =t,) [pnap]
mesopostnotum
pseudophragma |pph]
spina
mesonorum
metanotum
upper mesepimeron
ventral process of lateral furcal arm
first phragma
second phragma
attachment line of mesopostnotum
(PN2) to scutellum (t;) [60)
attachment line of mesopostnotum
(PN,) to metanotum (t,) [61]
Appendix 2. Character states of the mesofurcal-mesopostnotal complex attributed to Hymenoptera. Characters and states are
discussed in text. Superscript values refer to other observed states within a taxon.
Characters
Character?
1
2
3
4
5
6 7
8
9
111
11
12
1
2
3 A
5 6 /
8 9
10
11
12
Outgroup
i)
0
ii
1)
0
0 0
0
0
II
"
0
Proctotrupidae
5
2
3 1 1
1 0
2
0
0
Xyehnae
1
i)
1
•>
1
0 0
0
1
0
0
n
Mvmandae
7
~i
3 1 1
1 0
2
0
0
Macroxyelinae
1
ii
1
u
1
0 0
0
1
I)
11
il
Pteromalidae
5
2
3 1 1
1 0
2
0
0
Pamphilndae
1
0
1
0
■'
0 0
0
1
(1
11
ii
Mymarommatidae7
2
3 1 1
1 0
2
0
0
Blasticotomidae
2
0
1
Ii
2
0 1
0
1
0
1.1
ii
Platygastndae
"
T
3 1
1 0
2
0
0
Tenthredinidae
2
0
1
Ii
2
0 1
0
1
0
11
ii
Scehonidae
5
3'
2
3 1 1
1 0
i
0
11
Dipnomdae
2
0
1
0
p
0 1
0
1
0
II
(i
Bracomdae
5
■>
3 1
1 0
i
1)
II
Cimbicidae
2
0
2
0
2
0 1
0
1
0
0
ii
Ichneumonidae
5
2
3 r
1 0
i
0
II
Pergidae
i
1
?
1
)
0 1
0
1
0
1)
0
Plumariidae
5
-)
3 1
1 0
2
0
0
Argidae
2
0
1
1)
2
0 1
0
1
11
II
1.1
Sclerogibbidae
5
->
3 1
1 11
2
0
0
Cephidae
3
1)
1
1)
:
0
1
1
11
II
Embolerrudae
5
T
3 1
1 0
2
0
II
Anaxvehdae
-1
(1
1
1)
3
0
1
1
11
11
Dryinidae
5
2
3 1
1 0
^
II
(1
Sincidae
4
0
3
II
3
0
1
1
II
11
Bethylidae
5
T
3 1
1 0
2
0
0
Xiphvdrndae
5
1
1
1)
1
1
2
11
II
Chrysidid.it'
Orussidae
6
2
?
1
3
1
-1
0
0
Amiseginae
5
1
2
3 1
1 0
-i
0
0
Stephanidae
5
2
1
3
0
2
II
0
Cleptinae
5
4
~>
3 1
1 0
t
11
1
Trigonalvidae
5
2
1
3
1.1
-i
II
0
Chrysidinae
5
4
T
3 1
1 0
->
1
Megaspilidae
7
2
11
i
0
2
11
II
Bembicidae
5
5
T
3 1
1 0
2
0
Ceraphronidae
7
2
11
i
II
2
II
II
Larndae
5
5
2
3 1
1 0
2
II
Aulacidae
5
)
1
3
0
2
0
0
Heteri ^vnaeidae
5
5
2
I 3 1
1 0
T
0
Evanndae
5
2
1
3
1)
-i
II
0
Andremdae
5
6
2
I 3 1
1 0
2
II
Gasteruptudae
5
-i
1
1
11
2
II
0
Anthophoridae
5
6
2
1 3 1
1 0
2
II
[baliidae
5
2
0
1
II
i
0
0
Apidae
5
6
-i
3 1
1 0
2
II
Eucoilidae
5
2
11
3
II
2
II
II
Colletidae
5
b
2
1 3 1
1 1 0
T
0
Alloxystidae
5
j
II
3
0
2
11
II
Halictidae
5
b
2
1 1 1
1 1 0
2
0
Diaprndae
Megachihdae
5
b
2
1 3 1
1 1 0
i
0
Ambositnnae
5
2
0
1
(1
-t
1.1
II
Tiphndae
5
5
T
1 3 1
1 1 0
2
11
Diaprnnae
5
2
11
3
II
2
II
11
Sapvgidae
5
5
2
1 3 1
I 1 1)
2
2
Belytinae
Mutilhdae
5
5
2
1 3 1
1 1 1
2
11
Acropiesta
5
i
II
3
II
2
11
II
Sierolomorphida*
5
5
2
1 3 1
1 1 11
2
II
2
Aneurrynchus
r.
2
II
3
0
2
II
0
IVmpihdae
5
5
T
1 3 1
1 1 0
i
11
2
Oxylabis
5
2
1
1
1.1
2
II
11
Rhopalosomatidae5
5
2
1 3 1
1 1 1
1
0
0
Monomachidae
5
i
II
3
11
2
II
II
Bradvnobaenid.u
5
4
2
1 3 1
1 1 1
->
11
0
Ropronndae
5
2
1
3
11
2
0
0
Formicidae
5
1
2
1 3 1
1 1 0
2
II
II
Helondae
5
2
1
1
(1
2
II
0
Scoliidae
5
5
2
1 3 1
1 1 11
">
1
2
Pelecinidae
"
2
1
3
II
2
II
0
Vespidae
s
5
2
1 3 1
1 1 11
2
II
2
Vanhornudae
5
->
(1
}
II
2
II
0
Volume 3, 1 994
265
dl
Figs. 1-2. Pamphiliidae. l,Pamphilius sp., lateral habitus of mesosoma; 2, Acantholyda sp., sagittal section of mesosoma. Muscle
127 removed and location indicated by dashed line. Abbreviations as listed in Table 1 and Appendix 1.
266
Journal of Hymenoptera Research
3 XYELIDAE
sps2
4 XYELIDAE
5 PERGIDAE
Figs. 3-5. 3, Xyela minor, oblique subsagittal section of mesothorax. 4, Xyela minor, skeletal components of Fig. 3, musculature
removed. 5, Acordulecera sp., frontolateral view of MF-MPN complex. Abbreviations as listed in Table 1 and Appendix 1.
Volume 3, 1 994
267
6 PAMPHILIIDAE
7 DIPRIONIDAE
8 CIMBICIDAE
Figs. 6-8. 6, Pamphilius sp.: a, dorsal view of MF-MPN complex; b, lateral view of MF-MPN complex. 7, Diprion sp., supralateral
view of MF-MPN complex. 8, Zaraea sp., dorsal view of MF-MPN complex. Abbreviations as listed in Table 1 and Appendix
268
Journal of Hymenoptera Research
9CEPHIDAE
10 ANAXYELIDAE
1 1 SIRICIDAE
Figs. 9-11. MF-MPN complex. 9, Cephus cinctus: a, dorsal view; b, supralateral view. 10, Syntexis libocedrii: a, dorsal view; b,
supralateral view. 11, Tremex sp.: a, dorsal view; b, lateral view. Abbreviations as listed in Table 1 and Appendix 1
Volume 3, 1 994
269
12 XIPHYDRIIDAE
1 3 ORUSSIDAE
14M0N0MACHIDAE
15 VANHORNIIDAE
Figs. 12-15. MF-MPN complex. 12, Xiphydria abdommalis: a, dorsal view; b, supralateral view. 13, Orussus sayii, supralateral
view. 14, Monomachus sp., supralateral view. 15, Vanhornia eucnemidarum: a, laterophragma; b, supralateral view. Abbrevia-
tions as listed in Table 1 and Appendix 1.
270
Journal of Hymenoptera Research
16 SCELIONIDAE
17 ICHNEUMONIDAE
18 MUTILLIDAE
19 RHOPALOSOMATIDAE
Figs. 16-19. MF-MPN complex. 16, Sparasion sp., supralateral view. 17, Megarhyssa sp. (Ichneumorudae): a, laterophragma
(inner view); b, supralateral view. 18, Mutilhdae (male), supralateral view. 19, Rhcpalosoma sp., supralateral view. Abbrevia-
tions as listed in Table 1 and Appendix 1.
Volume 3, 1 994
271
20 BLASTICOTOMIDAE
23 TRIGONALYIDAE
21 TENTHREDINIDAE
24STEPHANIDAE
22 IBALIIDAE
25 MEGASPILIDAE
26GASTERUPTIIDAE
27 EVANIIDAE
28 PLATYGASTRIDAE
Figs. 20-28. MF-MPN complex of Apocrita: right laterophragma (upper figure); mesofurca in dorsal view (lower figure). 20,
Blasticotoma sp.; 21, Nematus sp.; 22, Ibalm sp.; 23, Orthogonalys pukhella; 24, Megischus sp.; 25, Megasptlus sp.; 26, Gasteruption
sp.; 27, Hyptia sp.; 28, Isocybus sp.
272
J JNAI C)( HvMENOPTERA RfSLARi H
JLji
29DIAPRIIDAE
30 HELORIDAE
31 PELECINIDAE
32 PROCTOTRUPIDAE 33BRACONIDAE, 34PLUMARIIDAE
HYBRIZONTINAE
35 SCLEROGIBBIDAE
36EMBOLEMIDAE
37 DRYINIDAE
38BETHYLIDAE
Figs. 29-38. MF-MPN complex of Apocnta: right laterophragma (upper figure); mesofurca in dorsal view (lower figure). 29,
Oxylabis sp.; 30, Helorus sp.; 31, Pelecinus polyturator; 32, Proctotrupidae; 33, Hybnzon sp., 34, Plumana sp.; 35, Sclerogibbidae,
36, Embolemus sp.; 37, Gonatopodinae (male); 38, Goniozus sp.
Volume 3, 1994
273
ap —
39 CHRYSIDIDAE, AMISEGINAE
40CHRYSIDIDAE, CHRYSIDINAE
41 VESPIDAE
42 POMPILIDAE
43F0RMICIDAE
45 APIDAE
44 TIPHIIDAE
46 LARRIDAE
Figs. 39-46. MF-MPN complex of Apocrita: right laterophragma (upper figure); mesofurca in dorsal view (lower figure). 39,
Amiseginae; 40, Chn/sis sp.; 41, Vespula sp.; 42, Aporinella galapagoensis; 43, Camponotus planus, 44, Myzmuni sp.; 45, Tngona sp.;
46, Larra sp.
274
Journal of Hymenoptera Research
PN2
47 XYELIDAE
48 PAMPHILIIDAEl
■*-,.
^
.
**
50 XIPHYDRIIDAE
Figs. 47-50. Mesopostnotum, dorsal view. 47, Pleuroneura sp.; 48, Pamphilius sp, 49, Cephus cinctus; 50, X,PMn« abfommoto
Abbreviat.ons as listed in Appendix 1. Split f.gures are magnifications of highlighted area in upper figure.
Volume 3, 1 994
275
Figs. 51-54. 51-52, Mesopostnotum in lateral view with magnification of connection to upper mesepimeron. 51 , Pamphilius sp.;
52, Cqyhus ductus. Arrow indicates articulation of laterophragma with upper mesepimeron. 53-54, MF-MPN complex of
Lasioglossum, dorsolateral view. 53, fused meso- and metafurca; 54, magnification of axillary lever (ap). Other abbreviations
as listed in Table 1 andAppendix 1. Split figures are magnifications of highlighted area in upper figure
276
Journal of Hymenoptera Research
Fig. 55. Phylogenetic hypothesis for extant Hymenoptera based on Rasnitsyn (1988) and Gibson (1985, 1993), with characters
of the MF-MPN complex superimposed. Characters and states discussed in text. Solid circles are unique apomorphies; shaded
circles indicate convergence; open circle indicates reversal. Xyelidae are not demonstrably paraphyletic, but under some
optimizations character 5:2 is treated as a synapomorphv of Macroxyehnae and the remaining Hymenoptera (dotted line to
alternate placement, see text for discussion). Character 3:? was not optimized, and this tree is two steps longer than found in
the parsimony analyses (see text).
Volume 3. 1994
277
Fig. 56. Phylogenetic hypothesis for Apocrita based on Rasnitsyn (1988), with characters of the MF-MPN complex superim-
posed. Characters and states discussed in text. Placement of Peradenndae and Vanhorniidae by Masner (pers. comm., 1993).
Superfamily names follow Huber and Goulet (1993), those names in parentheses were not examined for internal characters
and question marks indicate unknown character states. Solid circles are unique apomorphies; shaded circles indicate
convergence; open circle indicates reversal. Multiple states indicate groundplan first followed by proposed derived state
changes; changes listed in Appendix 2.
J. HYM. RES.
Vol. 3, 1994, pp. 279-302
A Taxonomic Study of the genus Ascogaster in China"
(Hymenoptera: Braconidae: Cheloninae)
Yuquing Tang and Paul M. Marsh
(YT) Biological Control Research Institute, Fujian Agricultural College, Fuzhou, Fujian, People's Republic of China; (PMM)
Cooperating Scientist, Systematic Entomology Laboratory, U. S. Department of Agriculture
(present address: P. O. Box 384, North Newton, Kansas 67117)
Abstract. — The chelonine genus Ascogaster is studied for the first time from the largely unknown area of mainland China and
Taiwan. Twenty-three species are described and keyed, of which 1 3 are new to science and six are recorded for the first time from
this area.
INTRODUCTION
MATERIALS AND METHODS
Ascogaster Wesmael is a cosmopolitan bra-
conid genus whose species are parasitic upon
microlepidoptera, predominantly Tortricidae.
Some species have been recorded frequently as
parasitoids of several economically important in-
sect pests and have been considered for use in
biological control programs. About 110 species of
Ascogaster have been recorded in the world.
Huddleston (1984) revised the 30 Palaearctic spe-
cies; Tobias (1986a, 1986b, 1988) and Papp (1989)
added 12 species to this region. Shaw (1983) re-
viewed 11 species for the Nearctic Region and
described a related new genus, Leptodrepana (4
species). Walker and Huddleston (1987) recorded
12 species of Ascogaster in New Zealand. The fauna
of the Indo-Australian Region is largely unknown,
although Baker (1926) and Szepligeti (1905, 1908)
described a few species. The Ascogaster fauna of
China is especially poorly studied with only five
species recorded, three from mainland China
(Fahringer 1934; He, et al 1989) and two from
Taiwan (Sonan 1932). In this work 23 species of
Ascogaster in China are treated, of which 13 species
are new to science and six are recorded for the first
time from China.
"Throughout this paper, China includes mainland China and
Taiwan.
Approximately 1,200 specimens of Chinese
Ascogaster were examined in this study, mostly
from the collections of the Fujian Agricultural
College, the Zhejiang Agricultural University, the
Zoology Research Institute of Academia Sinica,
the Taiwan Agricultural Research Institute and
the American Entomological Institute. The fol-
lowing acronyms are used to identify collections
that provided specimens for this study and several
as the depositories for type material:
AEIG American Entomological Institute, Gainesville,
Florida, USA
BM Bishop Museum, Honolulu, Haiwaii, USA
CNC Candian National Collection, Agriculture Canada,
Ottawa, Canada
ELKU Entomological Laboratory, Kyushu University,
Fukuoka, Japan
FAC Biological Control Research Institute, Fujian Agricul-
tural College, Fuzhou, China
HNHM Hungarian Natural History Museum, Budapest,
Hungary
IRSNB Institut Royal des Sciences Naturelles de Belgique,
Brussels, Belgium
TARI Taiwan Agricultural Research Institute, Wufeng, Tai-
wan
UEI Entomological Institute, Hokkaido University,
Sapporo, Japan
USNM National Museum of Natural History, Washington,
D. C, USA
ZAU Department of Plant Protection, Zhejiang Agricul-
tural University, Hungzhou, China
2S0
Journal of Hymenoptera Research
ZRJ Zoology Research Institute, Academia Simca, Beijing,
China
We also examined available Palaearctic and
Indo-Australian material in the U. S. National
Museum of Natural History and some type mate-
rial from other institutions as follows: USNM -
abdominatorDahlbom,albitarsusRemhard,annulans
Nees von Esenbeck, atamiensis Ashmead (holo-
type, synonymized with bidentida Wesmael by
Huddleston 1984), bicarinata Herrich-Shaffer,
bidentida Wesmael, brevicornis Wesmael, canifrons
Wesmael, cinctus Baker (holotype), coelioxoides
(Baker) (holotype), consobrnw Curtis, detectus Baker
(holotype), distinctus Baker (holotype), fullawayi
(Baker) (type series), inconspicuus Baker (holo-
type), intensus Baker ( holotype), laeviventris Baker
(holotype), longulus Baker (holotype), luzonensis
Baker (holotype), maculaticeps Baker (holotype),
malayanus (Baker) (holotype), modestus Baker (ho-
lotype), philippinensis Baker (type series),
quadradentata Wesmael, reticulata Watanabe (type
series), rufidens Wesmael, rufipes (Latreille), vividus
Baker (holotype); BM - argentea Fullaway
(syntypes); rugosa Fullaway (holotype); IRSNB -
armata Wesmael (lectotype), bidentida Wesmael
(lectotype), brevicorinis Wesmael (lectotype),
canifrons Wesmael (lectotype); limitatus Wesmael
(syntypes, synonymized with scabricula (Dahlbom)
by Huddleston 1984), quadridentata Wesmael (type-
series), rufidens Wesmael (type-series), varipes
Wesmael (type series); UEI - epinotiae Watanabe
(type series, synonymized with quadridentata
Wesmael by Huddleston 1984); reticulata Watanabe
(type series); ELKU -longicornis Huddleston (holo-
type, synonymized with formosensis Sonan, new
synonymy); HNHM - nilena Papp (holotype).
The general braconid morphological termi-
nology and measurements used in this study are
mostly after van Achterberg (1988). The terminol-
ogy used for wing veins is illustrated and explaned
in Sharkey ( 1988). Microsculprure terms are based
on Harris (1979). Two character states of the ocel-
lar triangle are defined after Huddleston (1984) as
follows: where a straight line drawn between the
anterior borders of posterior ocelli also touches
the anterior ocellus, the ocelli are referred to as
being 'on line'; where a line between the posterior
ocelli passes behind the anterior ocellus without
touching it, the ocelli are referred to as being 'not
on line'. The measurements CO and OD refer,
respectively, to the ocellar-ocular distance and the
diameter of the posterior ocellus. The ratio CL/
CW is carapace length (CL) to carapace width
(CW). Shaw (1983) and Huddleston (1984) dis-
cussed the important taxonomic characters of
Ascogaster in detail and we have used most of the
specific characters used by Huddleston (1984) for
the description of new species in this study (see
discussion under generic diagnosis). All dates from
labels on specimens examined have been stan-
dardized thus, 1-1-93, to avoid confusion.
BIOLOGY
Members of Ascogaster, like all the chelonines,
are internal egg-larval parasitoids of microlepi-
doptera, principally Tortricidae. They lay their
eggs into the host egg, but their larval develop-
ment is delayed at the first instar until the host
larva is mature. Little detailed work has been done
on the biology of Ascogaster. Huddleston (1984)
and Shaw (1983) reviewed the hosts of Ascogaster
for the Palaearctic and Nearctic Regions respec-
tively. Cox (1932), Rosenberg (1934) and Boyce
(1936) gave fairly detailed accounts of the biology
of A. quadridentata Wesmael. A research group in
Japan has recently studied in some detail host-
searching, host-preference, mating and oviposi-
tion behavior of A. reticulata W atanabe, an impor-
tant parasitoid of the smaller tea tortix, Adoxophyes
sp., in Japan (Kainoh 1986, 1988; Karnoh, el al. 1982;
Kainoh, et a/.1991; Kainoh and Tamaki 1982, 1988;
Kainoh, et fl/,1990; Kainoh, et al. 1989; Kamano, et
al. 1989; Kawakami 1985; Kawakami and Kainoh
1985, 1986). Little is known at present about the
biology of the Chinese Ascogaster species, how-
ever He et al. (1987) gave a full host list of A.
reticulata in mainland China.
SYSTEMATICS
The Cheloninae are distinguished from all
other subfamlies of Braconidae by the combina-
tion of having the first three metasomal terga
fused to form a carapace (Figs. 21-26), having a
complete postpectal carina and three submarginal
cells in the forewing (Fig. 1). Two tribes of the
Cheloninae, Chelonini and Phanerotomini, are
represented in the Palaearctic and Indo- Austrialian
regions. In the Phanerotomini the carapace is di-
vided into three tergites by two transverse su-
Volume 3. 1 994
281
tures; in the Chelonini the carapace has no trace of
sutures. The Chelonini are represented in the
Palaearctic and Indo-Australian regions by three
genera, Ascogaster Wesmael, Megascogaster Baker
and Chelonus Panzer. The former two genera can
be separated from Chelonus by having the first
submarginal cell always separated from the discal
cell by vein 1-Rs+M (Rs+M of some authors) (Fig.
1). Many authors (e.g. Baker 1926; Shaw 1983;
Huddleston 1984; Walker and Huddleston 1987;
van Achterberg 1990b; Zettel 1990) state that
Ascogaster is also characterized by having hairless
eyes. However, we have described one new spe-
cies, A. setula from Taiwan, with eyes densely and
distinctly setose (Fig .8), but with most of the other
characters of typical,4scogasfer. We include it within
Ascogaster until more species with this unusal
character have been found. Megascogaster Baker is
distinguished from Ascogaster only by having a
very elongate carapace (distinctly longer than head
and thorax) and a slender stigma in the forewing.
The taxonomic position of Megascogaster Baker
must wait until more material from the Indo-
Australian region has been studied. Shaw (1983)
cited thirteen characters for distinguishing
Ascogaster and Leptodrepana, however intermedi-
ate forms of most characters occur in many unde-
scribed Indo-Australian Ascogaster which we have
examined and we have chosen to follow van
Achterberg's synonymy. This concept makes
Ascogaster an apparent polyphyletic group and we
feel that a thorough phylogenetic study of the
entire genus is needed before the correct position
of Leptodrepana and the many other unusual forms
from the Indo-Australian Region can be settled.
That is beyond the scope of this limited study
which aims only to identify species from the pre-
viously unknown area of China.
Ascogaster Wesmael
Ascogaster Wesmael 1835:226. Type-species: Ascogaster
mstabilis Wesmael \-abdonunator (Dahlbom)], subse-
quently desigated by Foerster (1862).
Cascogaster Baker 1926:482. Type-species: Cascogaster fullawayi
Baker, original desigation, syn. by Watanabe (1937).
Leptodrepana Shaw 1983:37. Type-species: Leptodrepana
opuntiae Shaw, original designation; syn. by van
Achterberg (1990b).
Diagnosis. — Distinguished from all other bra-
conid genera by having: (a) the first three
metasomal terga fused into a rigid carapace with-
out transverse sutures; and (b) the forewing with
three submarginal cells and vein 1-Rs+M present,
separating the first submarginal and discal cells.
KEY TO THE ASCOGASTER SPECIES OF CHINA
1 Face punctate or rugose-punctate, occasionally nearly smooth; mesonotum usually predominantly
punctate, notauli usually distinct 2
Face strongly areolate or areolate-rugose, at least finely areolate-rugose; mesonotum usually coarsely
areolate-rugose so that notauli often are indistinct 13
2(1) Setae on upper part of face pointing upwards; ocellar triangle acute, ocelli usually not on line or almost
on line; propodeum without mediodorsal and apicolateral tubercles 3
Setae on upper part of face pointing downwards; ocellar triangle always obtuse, ocelli usually on line;
propodeum with distinct mediodorsal and apicolateral tubercles 6
3(2) Clypeus with conspicuous dentate flanges laterally; legs completely black or sometimes fore tibia
reddish-brown; ovipositor sheaths broad, knife-like semenovi Telenga
Clypeus without dentate flanges laterally; legs mostly yellow or yellow-brown; ovipositor sheaths not
conspicuously broad and knife-like 4
4(3) Clypeus with a large median incision apically; female carapace with a conspicuous dorsal prominence
at base ditnorpha Tang and Marsh, new species
Clypeus without any medial apical tubercle or incision; carapace without a dorsal prominence at base
5
5(4) Vein r of forewing about 1.0-1.2 times as long as 3-RS; hind coxa of male completely yellow
townesi Tang and Marsh, new species
Vein r of forewing about twice as long as 3-RS; hind coxa of male mostly black
yunnanica Tang and Marsh, new species
282
Journal of Hymenoptera Research
6(2) Eyes distinctly setose; antennae short, with 26-29 flagellomeres; body 2.9-3.1 mm
setula Tang and Marsh, new species
Eyes without setae or at most with only a few minute setae; antennae longer, generally with more than
30 flagellomeres; body generally more than 3.2 mm 7
7(6) Mesopleuron completely coarsely areolate-rugose so that precoxal suture not distinct; antennae very
long, 45-49 flagellomeres; apical border of clypeus weakly marginate medially ..fortnosensis Sonan
Mesopleuron at least in part smooth or punctate, precoxal suture usually distinct (except vanpes);
antennae shorter, at most with 40 flagellomeres; apical border of clypeus either produced or with one
or two small tubercles 8
8(7) Apical border of clypeus transversely impressed and produced medially but without medial excision or
tubercles 9
Apical border of clypeus with one or two distinct teeth medially 11
9(8) Hind coxa strongly strigate; carapace short, oval in dorsal view; mandible at base with a deep,
semicircular depression varipes Wesmael
Hind coxa largely smooth or finely punctate; carapace elongate, clavate in dorsal view; mandible at base
with vertical, parallel-sided depression 10
10(9) Legs yellow except hind coxa at base black, apex of hind femur and tarsi infuscate consobrina Curtis
Legs black except tibiae at base and basal tarsus of hind leg ivory or pale yellow albitarsus Reinhard
11(8) Apical border of clypeus with two medial tubercles, generally with a small excision between them; if
excision absent, then medial clypeal border with a narrow, spatulate projection, but never a medial
tooth arisanica Sonan
Apical border of clypeus with a single medial pointed toot 12
12(11 ) Carapace short, oval, broadest at about middle point in dorsal view; ventral opening of carapace shorter
in front of apex; hind coxa completely yellow perkinsi Huddleston
Carapace longer, clavate, broadest in posterior third in dorsal view; ventral opening of carapace long at
apex of carapace; hind coxa predominantly black lint Tang and Marsh, new species
13(1) Carapace elongate, CL/CW more than 2.0 14
Carapace shorter, usually oval, CL/CW generally 1.7 or less, never more than 2.0 15
14(13) Hind coxa strigate and black; ventral opening of carapace short, reaching only slightly beyond middle
of carapace grandis Tang and Marsh, new species
Hind coxa smooth or finely punctate, yellow; ventral opening of carapace longer, reaching apical 1 /4 of
carapace macrogaster Tang and Marsh, new species
15(13) Gena long and straight in face view; hind coxa strigate and yellowc/iooi Tang and Marsh, new species
Gena more or less rounded in face view; hind coxa often punctate or if strigate, then always mostly black
16
16(15) Interantennal carina strongly raised into an erect triangular flange between scapes 17
Interantennal carina present but never strongly raised into a triangular flange between scapes 19
17(16) Carapace rounded, without a tubercle apically; clypeus matte, rugose-punctate, its apical border straight
with no medial tooth annatoides Tang and Marsh, new species
— Carapace apically pointed, with a tubercle at apex; clypeus smooth, sparsely punctate, its apical border
pointed with a medial tooth 18
18(17) Carapace oval; face and temple areolate-rugose acutus Tang and Marsh new species
— Carapace acutely pointed; face and temple strongly and coarsely rugose fullawayi (Baker)
19(16) Hind coxa smooth, or finely punctate, yellow or infuscate at base; hind tibia with a medial pale colored
band; clypeus transversly impressed apically, without a medial tooth or if with a medial tooth, then
lower part of face smoother than upper part 20
Hind coxa strigate, at least at part, always black; hind tibia generally completely black or pale at base,
but never with a medial pale-colored band medially; clypeus always with a distinct medial tooth; face
evenly areolate or strongly areolate-rugose 21
20(19) Clypeus matte, rugose-punctate, straight apically, with no medial tooth; face strongly areolate
reticulata Watanabe
Clypeus smooth, finely and sparsely punctate, with a distinct medial apical tooth; face finely areolate-
rugose rugulosa Tang and Marsh, new species
21(19) Anterior corners of middle mesonotal lobe swollen, notauli distinct; palpi black
gibbosa Tang and Marsh, new species
Volume 3. 1994
283
— Anterior corners of middle mesonotal lobe not swollen, notauli indsitinct; palpi yellow or reddish-brown
22
22(21). Temple short, about equal in width to eye in dorsal view; carapace short, oval in dorsal view, deep in
lateral view quadridentata Wesmael
— Temple rounded, distinctly longer than eye in dorsal view; carapace longer, clavate in dorsal view, not
so deep in lateral view hei Tang and Marsh, new species
Ascogaster acutus Tang and Marsh,
new species
Fig. 23
Female. — Length of forewing 2.5-2.7 mm, of
body 3.0-3.2 mm.
Head. — Antenna with 28-32 flagellomeres,
slightly dilated medially, medial flagellomeres
slightly broader than long, apical ones longer than
broad. Temple slightly contracted behind eyes,
distinctly longer than length of eye in dorsal view.
Occiput strongly concave. Ocelli small, on line,
00= 4.0 OD. Frons strongly depressed, finely
rugose or smooth, medial carina distinct, extend-
ing from uper part of face betwen antennae to
anterior ocellus and strongly expanded between
antennae into an erect triangular lamella. Eyes
protuberant, glabrous. Face slightly protuberant,
about twice as broad as high, strongly areolate-
rugose. Clypeus punctate, strongly divided from
face, apical border produced medially into a small
pointed tooth.
Mesosoma. — Pronotum slightly projecting in
front of mesonotum, dorsolaterally rugose-punc-
tate. Notauli not distinct; mesonotum completly
coarsely rugose. Precoxal suture indistinguish-
able; mesopleuron strongly rugose. Propodeum
coarsely areolate-rugose, divided by a transverse
carina which is rasied into a medial pair and a
lateral pair of dentate flanges. Hind coxa strigate.
Vein r of forewing about 1.0-1.5 times as long as 3-
RS.
Metasoma. — Carapace (Fig. 23) short, with
pointed tubercle at apex, areolate-rugose. Ventral
opening of carapace longer, not distinctly in front
of apex. Ovipositor concealed.
Color. — Black; carapace yellow at base; all legs
yellow except hind coxa at base, tibia at apex and
tarsi black.
Male. — Same as females except antenna not
dilated medially, smaller yellow spot at basal
carapace.
Holotype Female. — BEIJING CITY: Mentougou,
VIII-25-81, M. S. Shi. Deposited in ZAU.
Paratypes.— BEIJING CITY: 1 female,
Lugouqiao, IX-57, T. L. Chen; 1 male, IX-78, J. H.
He. GUIZHOU PROVINCE: 1 female, Guiyang,
V-13-85, Q. H. Luo. FUJIAN PROVINCE: 1 female,
Shaowu, VII-20-45, H. F. Chao; 1 female, Shanhan,
VII-19-21-88, Y. Ma. TAIWAN: 1 female Duruanpi,
VIII-5-82, K. C. Chou and C. N. Lin; 1 female,
Wushe, IV-13-83, H. Townes; 1 female, Wushe, rV-
19-83, H. Townes; 1 female, Wushe, V-3-83, H.
Townes; 2 females, Wushe, V-10-83, H. Townes; 1
male, Wushe, V-15-83, H. Townes. Deposited in
AEIG, FAC, TARI, ZAU and ZRI.
Host. — Unknown.
Distribution. — China (Beijing City, Guizhou
Province, Fujian Province),Taiwan.
Remarks. — This species is very close tofidlawayi
(Baker) from which it can be distinguished by the
characters mentioned in the key.
Etymology. — The specific name is from the
Latin acutus meaning pointed in reference to the
pointed tubercle at the apex of the carapace.
Ascogaster albitarsus Reinhard
Ascogaster albitarsus Reinhard 1867: 364. Lectotype male,
POLAND: Gdansk (designated by Huddleston 1984).
Shenefelt 1973: 815; Huddleston 1984: 364; Papp 1989:
297.
Ascogaster leptopus Thomson 1874: 584. Lectotype female,
SWEDEN (designated by Huddleston 1984 and syn. by
Hellen 1953)
Diagnosis. — Length of foewing 3.3-3.5 mm, of
body 3.9-4.0 mm. Antenna with 36-38
flagellomeres, slightly dilated medially (female);
ocelli almost on line; face about 1 .5 times as broad
as high, rugose-punctate; clypeus with its apical
border produced medially and slightly reflexed
forwards; notauli distinct, fovealate, coalescing
posteriorly in a large areolate-rugose area, rest of
mesonotum densely areolate-punctate; precoxal
suture strongly areolate-rugose anteriorly, rather
weak posteriorly, mesopleuron anterodorally
strongly rugose, rest of mesopleuron finely punc-
tate; hind leg predominately dark with base of
284
Journal of Hymenoptera Research
tibia and tarsus pale-yellow; hind coxa finely punc-
tate; carapace long, clavate and widest in posterior
third.
Specimens Examined.— HEBEI PROVINCE: 2
females. SICHUAN PROVINCE: 2 females.
Additional Specimens Examined. — WESTERN
EUROPE: 1 male, det. by Huddleston, 1983.
Host. — A tortricid moth on pine tree.
Distribution. — China (Heibei Province, Sichuan
Province), Korea and some European countries.
Remarks. — Our specimens from China have
the mesopleuron with a polished impuncate area
posteroventrally, but otherwise agreeing with the
redescription of Huddleston (1984)
Ascogaster arisanica Sonan
Fig. 3
Ascogaster arisanicus Sonan 1932: 79. Holotype male, TAI-
WAN: Alishan (=Arisan). Watanabe 1937: 77 (as syn. of
rufipes (Latreille), Chou 1981 : 72 (notes on the locality of
types).
Ascogaster arisanica Sonan: emended by Huddleston 1984:
365.
Diagnosis. — Length of forewing 2.9-3.2 mm, of
body 3.3-3.7 mm. Antenna with 34-38
flagellomeres, moderately dilated medially (fe-
male); ocellar triangle obtuse, ocelli almost on line;
face (Fig. 3) about 1.5 times as broad as high,
densely punctate or rugose-punctate; apical bor-
der of clypeus produced, with a distinct medial
excision flanked by two small tubercles; notauli
shallow, coarsely rugose, coalescing posteriorly in
an areolate-rugose area, rest of mesontum punc-
tate or rugose-punctate; precoxal suture broad,
shallow, foveolate; rest of mesopleuron sparsely
punctate except dorsally rugose; hind coxa finely
punctate; carapace elongate, broadest in distal
half; ventral opening of carapace conspicuously in
front of apex.
Specimens Examined.— ZHEJIANG PROV-
INCE: 16 females, 9 males. SICHUAN PROV-
INCE: 2 females. YUNNAN PROVINCE: 1 male.
GUANGXI PROVINCE: 1 female. HAINAN
PROVINCE: 1 female. TAIWAN: 115 females, 35
males.
Host. — Unknown.
Distribution. — China (Zhejiang Province,
Sichuan Province, Yunnan Province, Guangxi
Province, Hainan Province), Taiwan, Japan.
Remarks.— Huddleston (1984) stated that "This
species is structurally very close to bidentula, but it
can be distinguished by the more massive head
and the more elongate carapace." However, our
specimens generally have the heads not so mas-
sive as described by Huddleston. In Chinese speci-
mens examined here, the temple is only 1.0-1.3
times as long as eye in dorsal view. Sonan (1932)
named this species as arisanicus, however, the
holotype is labelled 'Ascogaster artsensis Sonan'.
Ascogaster armatoides Tang and Marsh,
new species
Fig. 19
Female. — Length of forewing 3.5-3.6 mm, of
body 4.2-4.4 mm.
Head. — Antenna with 42 flagellomeres, mod-
erately dilated medially, medial segments about
as broad as long. Temple constricted behind eyes,
slightly shorter than length of eye in dorsal view.
Occiput strongly concave. Ocelli very small, on
line, OO = 4.0 OD. Frons strongly excavate behind
antennae, smooth, with a distinct medial carina
which is expanded between antennae into an erect
triangular flange (Fig. 19). Eyes glabrous, strongly
protuberant. Face slightly protuberant, about 1.5
times as broad as high, strongly areolate-rugose.
Clypeus dull, rugose-punctate, not very distinctly
divided from face; apical border convex, not pro-
duced medially and without distinct tooth or tu-
bercle.
Mesosma. — Pronotum slightly projecting in
front of mesonotum, areolate-rugose laterally
Notauli indistinct; mesonotum strongly rugose.
Precoxal suture indistinct; mesopleuron com-
pletely areolate-rugose. Propodeum strongly ar-
eolate-rugose, divided into dorsal and posterior
surfaces by a transverse carina which is raised into
a pair of medial blunt teeth and laterally a pair of
less stout teeth. Hind coxa strongly strigate. Vein
r of forewing about as long as 3-RS.
Metasotna. — Carapace short, CL/CW = 1.6,
oval in dorsal view, somewhat pointed in
lateralview, areolate-rugose. Ventral opening of
carapace short, in front of apex. Ovipositor
convealed.
Male. — Unknown.
Holotype Female.— GUANGXI PROVINCE:
Longzhou, V-18-82, J. H. He. Deposited in ZAU.
Paratypes. — INDIA: 4 females, Ammatti, S.
Coorg, 11-52, P. S. Nathan. Deposited in CAN,
USNM.
Volume 3, 1994
285
Host. — Unkown.
Distribution. — China (Guangxi Province), In-
dia.
Remarks. — This species is very close to arniata
Wesmael from which it can be distinguished by its
longer antenna and less massive head.
Etymology. — The specific name is in reference
to the similarity of this species to arniata.
Ascogaster chaoi Tang and Marsh, new species
Female. — Length of forewing 3.0-3.4 mmm, of
body 3.5-4.0 mm.
Head. — Antenna with 40-42 flagellomeres,
strongly dilated medially, tapered at apex, medial
segments slightly broader than long, apical seg-
ments about as long as broad. Temple constricted
behind eyes, slightly longer than length of eyes in
dorsal view. Occiput deeply concave. Ocelli small,
OO =3.5 OD, on line. Frons moderately depressed,
finely punctate, with distinct medial carina. Eyes
slightly protuberant, glabrous without distinct
setae. Gena long and straight in face view. Face
slightly protuberant, about 1.5 times as broad as
high, areolate-rugose. Clypeus protuberant, punc-
tate, distinctly divided from face; apical border
slightly impressed laterally, produced medially
into stout dentate flange.
Mesosoma. — Pronotum projecting in front of
mesonotum, dorsolaterally areolate-rugose.
Notauli indistinguishable; mesonotum strongly
areolate-rugose. Mesopleuron completely rugose
so that precoxal suture not distinct. Propodeum
strongly areolate-rugose, with a medial transverse
carina raised into a pair of medial stout dentate
and a pair of lateral less stout dentate. Hind coxa
strigate. Vein r of forewing 1.0-1.5 times as long as
3-RS.
Metasoma. — Carapace shorter, CL/CW = 1.5-
1.7, somewhat pointed in dorsal view, rounded
and deep in lateral view, areolate-rugose. Ventral
opening of carapace short, distinct in front of apex.
Hypopygium short. Ovipositor sheath clavate.
Color. — Black; carapace yellow at base; all legs
yellow except hind femur and tibia black at apex.
Male. — Unknown.
Holotype Female.— FUJIAN PROVINCE:
Huanggangshan, VII-14-85, Y. Q. Tang. Deposited
inFAC.
Paratypes.— FUJIAN PROVINCE: 1 female,
same as holotype; 6 females, Tongmuguan
(Wuyishan), VII-11-82, J. C. Huang; 1 female,
Wuyishan, VI-80, H. F. Chao; 2 females, Shanguan
(Wuyishan), VII-5-85, D. H. Huang and G. Zheng;
1 female, Wuyishan, VII-15-86, J. S. Wang.
GUANGXI PROVINCE: 1 female, Tainlin, V-30-
82, J. H. He ; 1 female, Longsheng, VI-24-84, J. H.
He. ZHEJIANG PROVINCE: 5 females, Songyang,
VII-15-89, J. H. He and H. L. Chen; 1 female,
Tianmushan, VI-17-83, Y. Ma. ANHUI PROV-
INCE: 1 female, Huangshan, VII-30-86, S. C. Zhang.
JILING PROVINCE: 1 female, Dongliao, VII-20-
31-88, X. M. Luo. Deposited in FAC, USNM, ZAU.
Host. — Unknown.
Distribution. — China (Jiling Province, Anhui
Province, Zhejiang Province, Guangxi Province,
Fujian Province).
Remarks. — This species is close toquadridentata
from which it can be easily distinguished by its
longer antenna, straight and long gena, and yel-
low hind coxa.
Etymology. — This species is named for Prof. H.
F. Chao, Fujian Agricultural College, in apprecia-
tion for his outstanding contribution to the tax-
onomy of parasitic Hymenoptera in China.
Ascogaster consobrina Curtis
Figs. 20, 27
Ascogaster consobrmus Curtis 1837: 672. Holotype male, GREAT
BRITAIN: England. Shenefelt 1973: 819; Huddleston
1984: 367; Papp 1989: 297.
Diagnosis. — Antenna with 32-34 flagellomeres,
slightly dilated medially (female); ocellar triangle
obtuse, ocelli almost on line; face (Fig. 20) about 1 .5
times as broad as high, areolate-punctate, hairs
downwards; apical border of clypeus not retracted,
medially produced into a broad, blunt point, with-
out impression or tubercle; notauli distinct,
foveolate, rest of mesonotum punctate except with
a broad areolate-rugose area posteromedially;
precoxal suture distinct foveolate, mesopleuron
(Fig. 27) dorsally finely rugose-punctate except for
a polished and impunctate area just dorsal precoxal
suture, ventrally always sparsely punctate, shin-
ing; carapace rather long, clavate, widest in poste-
rior third, with a downwardly directed anterior
flange; hind coxa punctate, sometimes weakly
rugose dorsaly; hind leg yellow except coxa at
base black, apex of femur and sometimes of tarsus
infuscate. Length of forewing 3.3-3.6 mm, of body
3.8-4.3 mm.
286
Journal of Hymenoptera Research
Specimens Examined.— ZHEJIANG PROV-
INCE: 1 female. TAIWAN: 89 females, 12 males.
Additional Specimens Examined. — Western Eu-
ropean countries, 6 females, 10 males, det. by
Huddleston, 1983.
Hosts. — No reared material was examined.
Shenefelt (1973) recorded Chelonia caja and Gelechia
vulgella as its hosts, but these records need to be
confirmed.
Distribution. — China (Zhejiang Province), Tai-
wan., Japan, Korea, and several European coun-
tries (see Huddleston 1984: 367)
Remarks. — All specimens examined agree well
with the redescription given by Huddleston (1984)
except for the slightly shorter medial flagellar
segments of the female, which are generally about
as long as broad.
Ascogaster dimorpha Tang and Marsh,
new species
Fig. 4
Female. Length of forewing 3.8 mm, of body
4.4 mm.
Head. Antenna, with 26 flagellomeres, flagel-
lum slightly dilated medially, all segments longer
than broad. Temple more or less rounded behind
eyes,, slightly longer than eye in dorsal view. Oc-
ciput strongly concave. Ocelli large, OO = 4.5 OD;
ocellar triangle acute, ocelli not on line. Frons
behind antenna moderately excavated, smooth
anteriorly, rugose-punctate posteriorly. Eyes mod-
erately protuberant, nearly glabrous with only
few scattered minute setae. Gena contracted be-
low in face view. Face (Fig. 4) slightly protuberant,
about twice as broad as high, densely and regu-
larly rugose-punctate, moderately hairy, the hairs
on the upper part of the face pointing upwards.
Clypeus moderately protuberant, rugose-punc-
tate, but less densely than that of face, its apical
border strongly incised medially.
Mesosoma. — Pronotum projecting in front of
mesonotum, rugose-foveolate dorsolaterally.
Notauli not very distinct, mesonotum finely punc-
tate except with an areolate-rugose area
posteromedially. Precoxal suture shallow
foveolate, rest of mesopleuron punctate.
Propodeum slightly impresed and punctate medi-
ally, strongly areolate-rugose laterally, but with-
out distinct tubercles. Hind coxa smooth and
shining. Vein r of forewing about 2.5 times as long
as 3-RS.
Metasoma. — Carapace long, CL/CW = 2.2, ar-
eolate-rugose, but finely and sparsely punctate
apically, with a distinct dorsal prominence at base.
Hypopygium large and broad. Ovipositor long
slender, upcurved.
Color. — Black; legs yellow-brown except hind
coxa at base and tarsis black, and hind tibia
infuscate with a pale-yelow band medially; cara-
pace with yellow-pale spots at anterolateral sides;
papli yellow-pale,
Males. — Same as female except antenna longer,
not dilated medially, with 30-31 flagellomeres and
carapace more slender, CL/CW = 2.3-2.5, more
rounded and deeper apically, without a dorsal
prominence at base. Length of forewing 3.6-3.9
mm, of body 4.3-4.5 mm.
Holotype female.— ZHEJIANG PROVINCE:
Songyang, VII-18-31-89, J. H. He. Deposited in
ZALL
Paratypes.— ZHEJIANG PROVINCE: 3 males,
Songyang, VII-15-17-89, J. H. He. TAIWAN: 5
males, Meifeng 2150 m, -24-26-81, K. S. Lin and W.
S. Tang (TARI); 1 male, Meifeng, VI-22-26-83, K. S.
Lin and S. C. Lin. Deposited in TARI, ZAU.
Hosts. — Unknown.
Distribution. — China (Zhejiang Province), Tai-
wan.
Remarks. — This species belongs to the caucasica-
group (sensu Huddleston 1984) by virtue of its
punctate face, hair on the upper part of the face
pointing upwards and the bidentate clypeus. It
appears to be related to caucasica Kokujev and
bicarinata Herrich-Schaffer, but is easily distin-
guished from the later two species by the charac-
teristics of the carapace.
Etymology. — The specific name refers to the
sexual dimorphism of the metasomal carapace.
Ascogaster formosensis Sonan
Fig. 5
Ascogaster formosensis Sonan 1932:78. Holotype female, TAI-
WAN: Arisan, Kunkiko. Watanabe 1937:76; Shenefelt
1973: 822; Chou 1981: 72 (notes on the locality of types);
Papp 1989: 297.
Ascogaster formosanus (!): Watanabe 1934: 198.
Ascogaster longicomis Huddleston 1984: 368. Holotype fe-
male, JAPAN: Mt.Tachibana (ELKU). New synonymy.
Diagnosis. — Antenna with 45-49 flagellomeres,
medial segment weakly dilated; ocelli on line or
Volume 3, 1 994
287
almost on line; face (Fig. 5) rugose-punctate, 1.2-
1.5 times as broad as high; clypeus with its apical
border rounded except its medial 1/4 weakly
emarginate; notauli foveolate, rest of mesonotum
densely punctate except posteromedially areolate-
rugose; mesopleuron coarsely rugose so that
precoxal suture not distinct; propodeum areolate-
rugose with 4 prominent tubercles; hind coxa finely
punctate; carapace always yellow at base, oval in
dorsal view, clavate in lateral view, ventral open-
ing of carapace distinctly in front of apex. Larger
species, length of forewing 5.0-5.4 mm, of body
5.8-6.4 mm.
Specimens Examined.— YUNNAN PROVINCE:
1 female. TAIWAN: 15 females, 7 males.
Additional Specimens Examined. — NEPAL: 1
female, 1 male. INDIA: 6 females, 6 males. JAPAN:
Holotype of longicorinis Huddleston.
Hosts. — Unknown.
Distribution. — China (Yunnan Province), Tai-
wan, India, Japan, Nepal.
Remarks. — This species is easily distinguished
from other species of Chinese Ascogaster by its
conspicuously long antennae and by the charac-
teristics of its face and clypeus. It is very similar to
philippinensis Baker from which it differs only in
the sclupture of the mesonotum. We treat it here as
a valid species given no intermediate forms are
known.
Ascogaster fullawayi (Baker)
Figs. 7, 24
Cascogaster fullawayi Baker 1926: 483. Holotype female (cited
as male), PHILIPPINES: Baguio, Benguet.
Ascogaster fullawayi: Shenefelt 1973: 822.
Diagnosis. — Antenna with 34-35 flagellomeres,
slightly dilated medially; scrobes outwardly mar-
gined by high, thin and complete carina middle of
which is raised into sharply angulate teeth; ocelli
almost on line; face (Fig. 7) narrow, less 1.5 times
as broad as long, strongly and coasely rugose;
clypeus smooth and shining, sparsely punctate,
apical border acute; mesonotum between notauli
swollen, strongly areolate-rugose; mesopleuron
coarsely areolate-rugose, precoxal suture indis-
tinguishable; hind coxa rugose-punctate; carapace
(Fig. 24) acute with a tubercle at apex; ventral
opening of carapace short, distinctly in front of
apex.
Speimens Examined. — TAIWAN: 2 females.
Additional Specimens Examined. — Holotype of
fullawayi (Baker), female, PHILIPPINES: Baguio,
Benguet (USNM). Holotype of fullawayi var.
maquilingensis (Baker), female, PHILIPPINES: Mt.
Makiling, Luzon (USNM). Paratype of fullawayi
(Baker), 1 female, PHILIPPINES: Imugin, N,
Viscaya (USNM).
Hosts. — Unknown.
Distribution. — Taiwan, Philippines.
Remarks. — Two Taiwanese speicmens exam-
ined agree well with the type of fullawayi (Baker)
except that the antennae are shorter. This species
is the only Philippine Ascogaster described by Baker
(1926) found in China. It is distinguished from the
other Chinese Ascogaster by the characteristics of
the carapace and sculpture of the head.
Ascogaster gibbosa Tang and Marsh,
new species
Fig. 28
Females.- — Length of forewing 3.0-3.2 mm, of
body 3.5-3.8 mm.
Head. — Antenna with 37-40 flagellomeres,
slightly dilated medially, medial segments about
as long as broad. Temple rounded behind eye,
slightly shorter than eye in dorsal view. Occiput
concave. Ocelli small, OO = 3.5-4.0 OD, on line.
Frons moderately depressed, smooth, with a weak
medial carina. Eyes protuberant, without distinct
setae. Gena in face view contracted. Face about 1.5
times as broad as high, coarsely areolate-rugose.
Clypeus distinctly divided from face, punctate,
apical border produced medially into a strongly
pointed large tooth.
Mesosoma. — Pronotum slightly projecting in
front of mesonotum, dorsolaterally areolate-rug-
ose. Notauli present, areolate-rugose; mesonotum
(Fig. 28) between notauli strongly protuberant,
rugose-punctate, rest of mesonotum areolate-rug-
ose. Precoxal suture indistinguishable;
mesopleuron completely areolate-rugose.
Propodeum strongly areolate-rugose, divided by
transverse carina which is raised medially and
laterally into prominent dentate flanges. Hind
coxa dorsally strigate. Vein r of forewing about as
long as 3-RS.
Metasoma. — Carapace moderately long, CL/
CW = 1.6-1.7, oval in dorsal view, somewhat
pointed in lateral view. Hypopygium short.
288
Journal of Hymenoptera Research
Ovipostor short, straight.
Color. — Almost completely black except ex-
treme apex of hind coxa and of femur yellow.
Male. — Unknown.
Holotype Female. — TAIWAN: Tsuifeng, VI-3-
80, L. Y. Chou and C. C. Chen. Deposited in TARI.
Paratypes. — TAIWAN: 2 females, same as ho-
lotype; 2 females, Tsuifeng, VII-16-82. S. C. Lin and
C. N. Lin; 1 female, Tsuifeng, VI-21-79, K. S. Lin
and B. H. Chen; 1 female, Meifeng, VII-31-IX-2-82,
L. Y. Chou and K. C. Chou; 1 female, Tayuling, IX-
12-15-80, K. S. Lin and C. H. Wang. Deposited in
TARI, USNM.
Host. — Unknown.
Distribution. — Taiwan.
Remarks. — This species can be easily distin-
guished from other species of the quadridentata-
group (see Huddleston 1984:371 ) by the character-
istics of the mesonotum and by the strongly pointed
clypeus tooth and completely black body.
Etymology.— The specific name is from the
Latin gibbosus meaning hunched or humped in
reference to the strongly humped mesonotum.
Ascogaster grandis Tang and Marsh,
new species
Fig. 6, 21
Females. — Length of forewing 4.0-4.1 mm, of
body 4.8-5.0 mm.
Head. — Antenna with 37 flagellomeres; flagel-
lum dilated medially, strongly tapered at apex,
medial segments about as long as broad. Temple
slightly constricted behind eyes, at least 1.5 times
as long as eye in dorsal view. Occiput deep, con-
cave. Ocelli very small, on line, OO = 4.0 OD. Frons
strongly depressed, rugose-punctate. Eyes protu-
berant, glabrous, without distinct setae. Gena swol-
len in lateral view. Face (Fig. 6) slightly protuber-
ant, about 1.5 times as broad as high, areolate-
rugose, with medial carina dorsally. Clypeus
slightly protuberant, punctate, distinctly divided
from face, apical border impressed laterally, pro-
duced medially into a very small pointed tooth.
Mesosoma. — Pronotum strongly projecting in
front of mesonotum, dorsolaterally rugose-punc-
tate, smooth ventrally. Notauli not distinct,
mesonotum strongly areolate-rugose. Precoxal
suture indistinguishable, mesopleuron strongly
areolate-rugose. Propodeum completely coarsely
areolate-rugose, divided by a transverse carina
produced into medial and postero-lateral pairs of
stout teeth. Hind coxa strongly strigate or ar-
eolate-rugose. Vein r of forewing about as long as
3-RS.
Metasoma. — Carapace (Fig. 21) very long, CL/
CW = 2.3-2.5, areolate-rugose. Ventral opening of
carapace very short, ending almost in middle of
carapace. Ovipositor concealed.
Color. — Black; fore and middle legs yellow
except coxae and femora black basally, hind leg
black except trochanters reddish-brown; papli
pale yellow.
Male. — Same as female except antenna not
dilated medially.
Holotype Female.— ZHEJIANG PROVINCE:
Xitianmushan, VII-29-84, X. J. Wu. Deposited in
ZAU.
Paratypes.— ZHEJIANG PROVINCE: 1 male,
same as holotype; 1 female, Xitianmushan, VII-27-
84, X. J. Wu; 1 female, Xitianmushan, VII-22-87, X.
M. Lou. Deposited in USNM, ZAU.
Host. — Unknown.
Distribution. — China (Zhejiang Province).
Remarks. — Both this species and macrogaster,
new species, run into the quadridentata-group in
the key of Huddleston (1984), but both of them are
distinguished from species of this group by the
characteristics of head shape and their elongate
carapaces.
Etymologi/. — The specific name is from the
Latin grandis meaning large in reference to the
large expanded temple.
Ascogaster hei Tang and Marsh, new species
Figs. 15, 16
Females. — Length of forewing 4.0-4.2 mm, of
body 4.6-5.0 mm.
Head. — Antenna with 33-37 flagellomeres,
moderately dilated medially, medial flagellomeres
slightly broader than long. Temple rounded be-
hind eyes, distinctly longer than length of eye in
dorsal view. Occiput deeply concave. Ocelli small,
OO = 3.5-4.0 OD, almost on line. Frons (Fig. 15)
moderately depressed behind antennae, rugose-
punctate, with a weak medial carina. Eyes protu-
berant, nearly glabrous but with a few scattered
setae. Gena in face view contracted. Face (Fig. 16)
about twice as broad as high, strongly areolate-
rugose. Clypeus distinctly divided from face, punc-
tate; apical border produced medially into a small
Volume 3. 1 994
289
pointed tooth.
Mesosoma. — Pronotum projecting in front of
mesonotum, dorsolaterally areolate-rugose.
Notauli not distinct; mesonotum strongly areolate-
rugose; precoxal suture indistinguishable;
mesopleuron coarsely areolate-rugose.
Propodeum strongly areolate-rugose, divided by
a transverse carina which is raised medially and
laterally into stout dentate flanges. Hind coxa
strongly strigate. Vein r of forewing about as long
as 3-RS.
Metasoma. — Carapace longer, CL/CW =1.6-
1.7, clavate in dorsal view. Ventral opening of
carapace longer, not very distinctly in front of
apex. Hypopygium short. Ovipositor sheaths clav-
ate.
Color. — Almost completely black except fore
tibia, and sometimes middle and hind coxae testa-
ceous apically.
Male. — Same as females except antennae not
dilated medially, with 34-35 flagellomeres.
Holotype Female.— ZHEJIANG PROVINCE:
Fengyangshan, VIII-12-84, L. R. Sheng. Deposited
inZAU.
Paratypes.— JILING PROVINCE : 2 females, 3
males, Dongjiang, VII-20-31-88, X. M. Luo.
HEILONHJIANG PROVINCE: 1 female,
Qingdinzhi, V-77, Y. Y. Hun. ZHEJIANG PROV-
INCE: 1 male, same as holotype; 1 female,
Xitainmushan, VIII-3-84, L. K. Sheng; 1 female,
Songyang, VII-15-17-89, J. H. He; 1 male, Zhuji, VI-
10-85, X. X. Chen. FUJAIN PROVINCE: 1 female,
1 male, Guadun (Wuyishan), VI-11-82, J H. Xiu; 1
female, Wuyishan, VI-20-80, N. Q. Lin; 1 female,
Wuyishan, VI-30-80, J. H. Xiu; 1 female, Wuyishan,
VIII-3-83, Y. Ma; 1 male, Wuyishan, VII-30-81, J. S.
Weng. Deposited in FAC, USNM, ZAU.
Host. — Unknown.
Distribution. — China (Heilongjiang Province,
Tiling Province, Zhejiang Province, Fujian Prov-
ince).
Remarks. — This species can be distinguished
from other species of the quadridentata-group by
the more massive head, strongly strigate-rugose
hind coxa, and larger and stout body.
Etymology. — This species is named for Prof. J.
H. He, Zheijiang Agricultural University, in ap-
preciation for his helpfulness in this work.
Ascogaster lini Tang and Marsh, new species
Fig. 9
Females. — Length of forewing 3.9-4.3 mm, of
body 4.4-4.8 mm.
Head. — Antenna with 33-36 flagellomeres,
weakly dilated medially, slightly tapered at apex,
medial segments about as broad as long, basal
segments distinctly longer than broad, apical seg-
ments slightly longer than broad. Temple strongly
constricted behind eyes, slightly shorter than eye
in dorsal view. Occiput deeply concave. Ocelli
small, on line, OO = 4.0 OD. Frons with two
depressed, polished impunctate areas behind an-
tenna, median carina distinct. Eyes moderately
protuberant, glabrous. Face (Fig. 9) slightly protu-
berant, about 1.5 times as broad as high, finely
punctate, with a median carina in upper part;
Clypeus weakly divided from face, protuberant,
sparsely punctate, smoother than face; apical bor-
der of clypeus strongly impressed laterally, me-
dial area raised and produced forwards into a
distinct pointed tooth.
Mesosoma. — Pronotum projecting in front of
mesonotum, laterally shining and smooth, dor-
sally rugose-foveolate. Notauli strong, foveolate,
coalescing posteriorly in an areolate-rugose area;
rest of mesonotum rugose-punctate. Precoxal su-
ture shallow, foveolate; mesopleuron above
precoxal suture rugose-foveolate anteriorly,
smooth and punctate posteriorly, rest of
mesopleuron ventrally densely punctate.
Propodeum completely areolate-rugose divided
medially by a transverse carina with medial and
lateral tubercles, the medial pair broad and stout,
lateral tubercles more prominent. Hind coxa finely
punctate. Vein r of forewing about 1.5-2.0 times as
long as 3-RS.
Metasoma. — Carapace long, CL/CW =1.8-2.0,
clavate in dorsal view, finely areolate-rugose, with
two weak medial carinae at base. Ventral opening
of carapace longer, almost at apex of carapace.
Hypopygium short. Ovipositor sheaths clavate,
short.
Color. — Black except carapace yellow at base;
antenna infuscate except scape yellow; legs yel-
low except hind coxa at base, mid and hind femur
at apex, and all tarsi infuscate.
Males. — Same as females except medial
flagellomeres not dilated, carapace completely
black and scape infuscate as in flagellum.
290
Journal of Hymenoptera Research
Holotype Female.— TAIWAN: Tsuifeng 2300 m,
malaise trap, VI-84, K. S. Lin and K. C. Chou.
Deposited in TARI.
Paratypes. TAIWAN: 1 female, 3 males ,
Tsuifeng," VI-21-79, K. S. Lin and B. H. Chen; 3
males, Tsuifeng, VI-3-80, L. Y. Chou and C. C.
Chen; 1 male, Tsuifeng, V-8-81, K. S. Lin and S. C.
Lin; 1 female, Tsuifeng, VII-1-3-81, T. Lin and W. S.
Tang; 12 females, 4 males, VI-25-27-81, K. S. Lin
and W. S. Tang; 2 females, 1 male, Tsuifeng, V-23-
82, L. Y. Chou; 9 females, 1 male, VII-16-82, S. C.
Lin and C. N. Lin; 4 females, 5 males, Tsuifeng, VI-
23-25-83, K. S. Lin and S.C. Lin; 13 females, 14
male, same as holotype; 20 females, 8 males,
Tsuifeng, VII-XI-84, Malaise trap, K. S. Lin and K.
C. Chou; 1 male, Tsuifeng, IX-85, Malaise trap, K.
S. Lin; 1 female, 1 male, Meifeng, V-10-79, K. C.
Chou; 2 females, Meifeng, VI-20-22-79, K. S. Lin
and B. H. Chen; 2 females, 1 male, Meifeng, VI-2-
4-80, L. Y. Chou and C. C. Chen; 1 female, Meifeng,
VI-5-8-80, C. C. Chen; 1 female, Meifeng, VI-8-80,
K.S. Lin and B. H. Chen; 1 female, Meifeng, X-5-9-
80, C. C. Chen and C. C. Chien; 1 female, 10 males,
Meifeng, V-7-9-81, K. S. Lin and S. C. Lin; 2 fe-
males, Meifeng, VI-24-26-81, K. S. Lin and W. S.
Tang ; 2 males, Meifeng, V-22-82, L. Y. Chou; 1
male, Meifeng, V-3-83, H. Townes; 5 males,
Meifeng, V-10-83, H. Townes; 2 males, Meifeng, V-
15-83, H. Townes; 1 female, 4 males, V-22-83, H.
Townes; 4 females, Meifeng, VI-22-26-83, K. S. Lin
and S. C. Lin; 1 male, Meifeng, V-8-11-84, K. C.
Chou and C. C. Pan; 1 female, 3 males, Meifeng,
VI-84, Malaise trap, K. S. Lin and K. C. Chou; 1
female, Meifeng, VIII-84, Malaise trap, K. S. Lin
and K. C. Chou; 1 female, Meifeng, X-84, Malaise
trap, K. S. Lin and K. C. Chou; 1 female, Meifeng,
X-85, Malaise trap, K. S. Lin; 1 female, Sungkang
2100 m, X-84, Malaise trap, K. S. Lin and K. C.
Chou; 1 female, 2 males, Sungkang, X-XI-85, Mal-
aise trap, K. S. Lin; 8 females, 3 males, Tayuling
2560 m, VI-9-16-80, Malaise trap, K. S. Lin and B.
H. Chen; 1 female, 2 males, Alishan 2400 m, VI-12-
16-65, T. Maa and K. S. Lin; 4 females, Anmashan
2275m, VII-6-9-79, L. Y. Chou. Deposited in AEIG,
FAC, TARI, USNM.
Host. — Unknown.
Distribution. — Taiwan.
Remarks. — This species is very similar to
perkinsi in the characteristics of the face and clypeus,
but it is rather different in the shape of the carapace
and color patterns of both legs and antennae.
Etymology. — It is a pleasure to name this spe-
cies in honor of its collector, Mr. K. S. Lin, who also
collected a large amount of Braconidae in Taiwan.
Ascogaster macrogaster Tang and Marsh,
new species
Females. — Length of forewing 3.5-4.2 mm, of
body 4.3-4.9 mm.
Head. — Antenna with 33 flagellomeres, slightly
dilated medially, medial segments about as long
as broad. Temple slightly contracted behind eyes,
slightly longer than eyes in dorsal view. Occiput
concave. Ocelli larger, OO =3.0 OD, on line. Frons
slightly excavated, rugose-punctate; medial ca-
rina distinct. Eyes not protuberant, glabrous with-
out distinct setae. Gena expanded in lateral view.
Face slightly protuberant, about 1 .5 times as broad
as high, areolate-rugose. Clypeus punctate; apical
border impressed laterally, produced medially
into a small pointed tooth.
Mesosma. — Pronotum strongly projecting in
front of mesonotum, dorsolaterally areolate-rug-
ose. Notauli not distinct; mesonotum rugose-punc-
tate anteriorly, rest of mesonotum foveate.
Mesopleuron deep punctate, but precoxal suture
not distinguishable. Propodeum strongly areolate-
rugose, not distinctly divided into dorsal and pos-
terolateral surfaces. Hind coxa smooth, shining.
Vein r of forewing about 1.5 times as long as 3-RS.
Metasoma. — Carapace very long, CL/CW =
2.5, areolate-rugose, rounded apically in lateral
view. Ventral opening of carapace at 1/4 apex of
carapace. Hypopygium broad. Ovipositor short,
pointed apically; ovipostor sheath clavate.
Color. — Black; carapace sometimes yellow at
base; all legs yellow except hind femur and tibia
brownish apically; palpi pale yellow.
Male. — Same as female except antenna not
dilated medially, all flagellomeres longer than
broad, and smaller yellow spot at basal carapace.
Holotype Female.— FUJIAN PROVINCE:
Tongmugun (Wuyishan), VI-23-82, J. C. Huang.
Deposited in FAC.
Paratypes.— FUJIAN PROVINCE: 1 female,
same as holotype; 1 female, Wuyishan, VI-80, H. F.
Chao; 1 female, Wuyishan, VI-80. J. C. Huang; 1
male, Huanxi, V-30-86, N. Q. Lin. Deposited in
FAC.
Host. — Unknown.
Distribution. — China (Fujian Province).
Volume 3, 1 994
291
Remarks. — This species is very close to grandis
from which it can be distinguished not only by the
characters mentioned in the key but also by the
lack of distinct tubercles on the propodeum.
Etymology. — The specific name is from the
Greek makros meaning long and gaster meaning
stomach in reference to the long metasomal cara-
pace.
Ascogaster perkinsi Huddleston
Figs. 10, 22
Ascogaster perkmsi Huddleston 1984: 368. Holotype female,
JAPAN: Mt.Tachibana, Fukuoka.
Diagnosis. — Length of forewing 2.6-2.9 mmm,
of body 3.0-3.5 mm. Antenna with 34-36
flagellomeres, slightly dilated medially (female);
ocellar triangle obtuse, ocelli almost on line; face
(Fig. 10) about 1.5 times as broad as high, rugose-
punctate; apical border of clypeus strongly im-
pressed laterally, medial area projecting and pro-
duced forwards into a distinct pointed tooth;
notauli weak, foveolate, rest mesonotum rugose-
punctate; precoxal suture foveolate, mesopleuron
above precoxal suture areolate-foveolate anteri-
orly, sparsely punctate posteriorly, rest of
mesopleuron densely punctate; carapace short,
oval in dorsal view, clavate in lateral view, gener-
ally yellow at base; ventral opening of carapace
(Fig. 22) short, distinctly in front of apex; hind coxa
finely punctate; all legs yellow except middle tibia
dark at base, hind femur and tibia dark at base,
mid and hind tarsi infuscate.
Specimens Examined. — ZHEJIANG PROV-
INCE: 41 females, 24 males. HUNAN PROVINCE:
1 male. FUJIAN PROVINCE: 4 females, 3 males.
TAIWAN: 11 females, 6 males.
Host. — Unknown.
Distribution. — China (Zhejiang Province,
Hunan Province, Fujian Province), Taiwan, Japan.
Remarks. — All specimens examined agree well
with the description of Huddleston (1984). This is
a new species record for the fauna of China.
Ascogaster quadridentata Wesmael
Figs. 17, 18
Ascogaster quadridentatus Wesmael 1835: 237. Lectotype fe-
male, BELGIUM: Brussels (designated by Shaw 1983).
Chelonus impressus Herrich-Schaffer 1838: 153. Syntypes,
GERMANY ( syn. by Reinhard 1867)
Ascogaster nigricornis Thomson 1892: 1719. Lectotype female,
SWEDEN (designated and syn. by Huddleston 1984).
Shenefelt 1973: 826.
Ascogaster cynipum Thomson 1892: 1720. Holotype male,
SWEDEN (syn. by Huddleston 1984). Shenefelt,1973:
820.
Ascogaster egregius Kokujev 1895: 83. Holotype male, USSR
(syn. by Huddleston 1984). Fahnnger 1934: 525; Shenefelt
1973: 821.
Chelonus nigrator Szepligeti 1896: 303. Holotype female, YU-
GOSLAVIA: Buccari (syn. by Huddleston 1984).
Shenefelt 1973: 859.
Chelonus carpocapsae Viereck 1909: 43. Holotype female, USA:
Michigan (syn. by Rosenberg 1934).
Ascogaster epinotiae Watanabe, 1937: 76. Holotype female,
JAPAN: Hokkaido, Sapporo (syn. by Huddleston 1984).
Shenefelt 1973: 821.
Ascogaster quadridentata; Shenefelt 1973: 828; Shaw 1983: 32;
Huddleston 1984: 376 Walker and Huddleston 1987:
343; Papp 1989: 258.
Diag)iosis. — Antenna with 27-32 flagellomeres,
slightly dialted medially; ocelli on line or almost
on line (Fig. 17); face (Fig. 18) about twice as broad
as high, genearlly finely rugose; clypeus finely
punctate, apical border produced medially into a
pointed tooth; notauli and precoxal suture indis-
tinguishable, mesonotum and mesopleuron com-
pletely areolate-rugose; hind coxa always mostly
black and strongly strigate-rugose; carapace oval,
short, genearlly deep in lateral view, sometimes
yellow at base; ventral opening of carapace short,
usually distinctly in front of apex.
Specimens Examined. — BEIJING CITY: 1, fe-
male, 1 male. JILFNG PROVINCE: 1 male. HUNAN
PROVINCE: 2 females. JIANGXI PROVINCE: 1
male. GUANGXI PROVINCE: 1 female, 1 male.
YUNNAN PROVINCE: 2 females, 2 males.
GUIZHOU PROVINCE: 3 females, 2 males.
ZHEJIANG PROVINCE: 29 females, 30 males.
FUJIAN PROVINCE: 18 females, 11, males. TAI-
WAN: 26 females, 7 males.
Additional Specimens Examined. — Lectotype of
quadridentatus Wesmael, female, BELGIUM: Brus-
sels (IRSNB). Lectoparatypes of quadridentatus
Wesmael, 1 female, 5 males, same as lectotype
(IRSNB). USA: 1 female, 1 male, WA, Pullman,
August 30, 1989, ex. Cydia pomonella, det. by Shaw,
1990 (FAC).
Hosts.— Huddleston (1984) and Shaw (1983)
recorded the following hosts in the Palaearctic and
Nearctic Regions: Cydiafunebrana (Treitsche), Cydia
nigricana Steph., Cydia pallifrontana (Lienig &
Zeller), Cydia pomonella (L.), Epiblema uddmanniana
292
Journal of Hymenoptera Research
(L.), Grapholitlui molesta (Busck), Grapholitapnmivora
(Walsh), Spilonota ocellana (Denis & Schif fermuller)
(Lepidoptera: Tortricidae); Endopiza viteana
(Clem.); Yponomeuta padella (L.) (Lepidoptera:
Yponomeutidae).
Distribution. — China (Beijing City, Jiling Prov-
ince, Jiangsu Province, Zhejiang Province, Yunnan
Province, Guizhou Province, Guangxi Province,
Fujian Province), Taiwan, Japan, Korea, New
Zealand, West Palaearctic Region, Nearctic Re-
gion.
Remarks. — This species has been studied
widely as a parasitoid of several economically
important insect pests in Europe and North
America. Its hosts and biology were reviewed by
Huddleston (1984) and Shaw (1983). There is very
little published work on this species in China.
Fahringer (1934) recorded it in Jiangsu Province
(=Kiangfu). This species is characterized by its
coarse sculpture, short oval and deep carapace,
short temple and antenna, highly strigate and
black hind coxa, and the single tooth on its cly peus.
Ascogaster reticulata Watanabe
Asc ogaster reticulatus Watanabe 1967: 41 . Holotype male (cited
as female), JAPAN: Hokkaido, Ashigawa. He, Chen and
Ma 1989: 438.
Ascogaster reticulata: Huddleston 1984: 377; Papp 1989: 298.
Diagnosis. — Length of forewing 2.8-3.2 mm, of
body 3.4-3.8 m. Antenna with 34-36 flagellomeres,
weakly dilated medially; ocelli large, on line; face
about 1.5 times as broad as high, coarsely irregu-
larly rugose; clypeus strongly punctate, matt and
its apical border almost flat without any trace of
tooth or tubercle; mesonotum areolate-rugose,
notauli indistinct; precoxal suture indistinguish-
able, mesopleuron strongly areolate-rugose; cara-
pace short, deep, sometimes yellow at base; ven-
tral opening of carapace short, distinctly in front of
apex; hind coxa yellow, smooth or sometimes
finely punctate; hind tibia black with a pale yellow
medial band.
Specimens Examined.— BEIJING CITY: 2 fe-
males. HENAN PROVINCE: 13 females.
SHANDONG PROVINCE: 8 females, 5 males.
SHANXI PROVINCE: 8 females. SHAANXI
PROVINCE: 3 females, 1 male. ANHUI PROV-
INCE: 1 male. JIANGSHU PROVINCE: 9 females,
2 males. JIANGXI PROVINCE: 2 females.
ZHEJIANG PROVINCE: 7 females, 4 males.
YUNNAN PROVINCE: 3 females. GUANHXI
PROVINCE: 1 female. FUJIAN PROVTNCE: 1 fe-
male, 1 male. TAIWAN: 31 females, 36 males.
Additional Specimens Examined. — Holotype of
reticulatus Watanabe, male, JAPAN: Hokkaido,
Asahigawa, June 15, 1966 (UEI). Paratypes of
reticulatus Watanabe: 3 females, JAPAN, May 16-
July 20, 1966 (UEI); 1 female, 1 male, JAPAN, July
20-23, 1966 (USNM).
Hosts. — Acleris fimbriana (Thunberg),
Adoxophyes orana (Fischer von Roslerstamm),
Archips issikii Kodama, Archips oporana (L.), Archips
pulchra (Butler), Carposiatw nipponensis Walsingham
(Lepidoptera: Tortricidae).
Distribution. — China (Beijing City, Henan
Province, Shandong Province, Shanxi Province,
Shannxi Province, Anhui Province, Jiangshu Prov-
ince, Jiangxi Province, Zhejiang Province, Yunnan
Province, Guangxi Province, Fujian Province),
Taiwan, Japan, Korea, Czechoslovakia.
Remarks. — This species is easily distinguished
from other species of the quadridentata-gToup by
the matt and no tooth clypeus and by the yellow
and smooth hind coxa and the hind tibia with a
yellow-pale band medialy.
Ascogaster rugulosa Tang and Marsh, new
species
Fig. 13
Females. — Length of forewing 3.0-3.2 mm, of
body 3.4-3.7 mm.
Head. — Antenna with 31-35 flagellomeres,
slightly dilated medially, flagellomeres 1-11 longer
than broad, rest of flagellum about as broad as
long. Temple rounded behind eyes, about equal to
length of eye in dorsal view. Occiput concave.
Ocelli small, OO =3.5-4.0 OD, on line. Eyes protu-
berant, glabrous. Frons moderately concave,
smooth, with a weak medial carina. Face (Fig. 13)
about 1.5 times as broad as high, finely areolate-
rugose, generally more finely scluptured ventrally
than dorsally. Clypeus punctate, apical border
produced medially into a pointed tooth.
Mesosoma. — Pronotum little projecting in front
of mesonotum, rugose-punctate dorsolaterally.
Notauli present but not very distinct; mesonotum
rugose-punctate except an areolate-rugose area
posteromedially. Precoxal suture very weak, shal-
low foveolate; rest of mesopleuron sparsely but
coarsely punctate. Propodeum coarsely areolate-
Volume 3. 1 994
293
rugose, divided by a tranverse carina which is
raised into a pair of medial dentate flanges and a
pair of lateral dentate flanges. Hind coxa finely
punctate. Vein r of forewing about as long as or
slightly longer than 3-RS.
Metasoma. — Carapace short, oval in dorsal
view, clavate in lateral view, areolate-rugose. Ven-
tral opening of carapace less distinctly in front of
apex. Ovipositor short, tapered apically.
Color. — Black; fore and middle legs yellow
except middle tibia at apex and all tarsi infuscate,
hind leg black except apex of coxa and base of
femur yellow and with a pale yellow medial band
on tibia; carapace always yellow laterally at base,
but black medially.
Males. — Same as females except antenna not
dilated medially.
Holotype Female.— TAIWAN: Meifeng, V-7-9-
81, K. S. Lin and S. C. Lin. Deposited in TARI.
Paratypes.— ZHEJIANG PROVINCE: 1 male,
Xitainmushan, VI-2-4-90, L. G. Weng. HAINAN
PROVINCE: 1 male, Shuiman, V-26-60, X. F. Li.
TAIWAN: 7 females, 1 male, same as holotype; 1
female, Meifeng, VII-26-78, K. C. Chou; 1 female,
Meifeng, IV-8-9-78, K. S. Lin; 1 male, Meifeng, VII-
18-79, K. C. Chou; 1 female, Meifeng, V-15-22-79;
1 female, Meifeng, VI-20-21-79, K. S. Lin and B. H.
Chen; 1 female, Meifeng, X-5-9-80, C. C. Chen and
C. C. Chien; 1 male, Meifeng, VI-24-26-81, K. S. Lin
and W. S. Tang; 3 males, Meifeng, VIII-28-29-81, L.
Y. Chou and S. C. Lin; 1 female, 1 male, VII-31-IX-
2-82, L. Y. Chou and S. C. Lin; 2 females, 2 males,
Meifeng, V-8-11-82, K. C. Chou and C. C. Pan; 1
male, Meifeng, VII-30-83, L. Y. Chou; 1 female,
Meifeng, VI-22-26-83, K. S. Lin and S. C. Lin; 1
female, Meifeng, X-4-7-82, K. C Chou; 1 male, V-
8-11-84, K. C. Chou and C. C Pan; 1 female,
Meifeng, VII-84, Malaise trap, K. S. Lin and K. C.
Chou; 1 female, Tungpu, X-18-21-82, K. C. Chou
and S. C. Lin; 1 female, Tungpu, VI-20-24-83, K. C.
Chou and C. Y. Wong; 1 female, Tungpu, VII-85,
Malaise trap, K. S. Lin; 1 female, Tsuifeng, VI-21-
79, K. S. Lin and B. H. Chen; 1 female, Tsuifeng, VI-
3-80, L. Y. Chou and C. C. Chen; 1 female, Tsuifeng,
V-8-81, K. S. Lin and S. C. Lin; 4 females, 1 male,
Tsuifeng, VI-25-27-81, K. S. Lin and W. S. Tang; 1
male, Tsuifeng, VIII-27-81, L. Y. Chou and S. C.
Lin; 1 male, Tsuifeng, IX-1-3-82, L. Y. Chou and K.
C. Chou; 3 females, Tsuifeng, VI-23-25-83, K. S. Lin
and S. C. Lin; 2 females, Tsuifeng, VI- VII-84, Mal-
aise trap, K. S. Lin and K. C. Chou; 3 males,
Tsuifeng, IX-85, Malaise trap, K. S. Lin; 4 males,
Tsuiofeng, IX-12-14-84, K. S. Lin and S. C. Lin; 3
males, Wushe, IV-26-83, H. Townes;l male, Wusha,
VII-25-78, K. C. Chou; 1 female, Sungkang, VIII-6-
84, K. S. Lin; 1 female, 1 male, Sungkang, X-84,
Malaise trap, K. S. Lin and K. C. Chou; 2 males,
Sungkang, Malaise trap, IX-X-85, K. S. Lin; 1 fe-
male, 2 males, Shengkuang, IX-20-68, K. S. Lin; 1
female, Wuling, VI-27-29-79, K. S. Lin and L. Y.
Chou; 1 female, Tayuling, VI-9-16-80, K. S. Lin and
B. H. Chen; 1 male, Lishan, IX-12-68, K. C. Chou.
Deposited in AEIG, FAC, TARI, USNM, ZAU,
ZRI.
Host. — Unknown.
Distribution. — China (Zhejiang Province,
Hainan Province), Taiwan.
Remarks. — This species is close to reticulata
from which it can be distinguished not only by the
characters mentioned in the key but also by its
smooth mesopleuron, less deep carapace and
mostly black hind coxa.
Etymologic — The specific name is in reference
to the rugulose sculpturing on the clypeus.
Ascogaster semenovi Telenga
Figs. 11, 25
Ascogaster semenovi telenga 1941: 310, 453. Holotype female,
MONGOLIA: Alashan, Dyn-juan-in. Shenefelt 1973: 825;
Huddleston 1984: 352.
Ascogaster kyushuensis Yoneda 1978: 291. Holotype female,
JAPAN: Kyushu, Fukuoka Pref., Fukuoka City, Hakozaki
( syn. by Huddleston 1984 ).
Diagnosis. — Length of forewing 4.3-4.7 mm, of
body 5.2-5.6 mm. Antenna of female with 20
flagellomeres and medial segments dilated,
genearlly broader than long, male with 25-26
flagellomeres, medially not dilated, all segments
longer than broad; ocellar triangle acute, ocelli not
on line; face (Fig. 11) about twice as broad as high,
rugose-punctate, the hairs on the upper part of
face pointing upwards; clypeus with its apical
border rounded medially, produced laterally into
broad dentate flanges, without tooth or incision;
notauli distinct, f oveolate, rest of mesonotum punc-
tate with an areolate-rugose area posteromedially;
precoxal suture deep foveolate, rest of mesopleuron
punctate with a deep foveolate groove
anterodorsally; hind coxa finely and sparsely punc-
tate; carapace (Fig. 25) long, CL/CW = 2.2-2.4,
pointed in dorsal and lateral views; ovipositor
294
Journal of Hymenoptera Research
sheaths broad, knife-like. Almost completely black
except sometimes fore tibia reddish-brown.
Specimens Examined.— JIANGSU PROVINCE:
2 females, 6 males. SHANGHAI CITY: 8 females,
3 males. ZHEJIANG PROVINCE: 2 females, 8
males.
Hosts. — Unkown.
Distribution. — China (Jiangsu Province, Shang-
hai City, Zhejiang Province), Mongolia, Japan.
Remarks. This remarkable speices is easily dis-
tinguished from other species of Ascogaster by the
lateral dentate flanges on its clypeus and by the
broad and knife-like ovipositor sheaths. It is new
to the fauna of China.
Ascogaster setula Tang and Marsh, new species
Fig. 8
Females. — Length of forewing 2.4-2.7 mm, of
body 2.9-3.1 mm.
Head. — Antenna with 26-29 flagellomeres; fla-
gellum moderately dilated medially, medial seg-
ments slightly broader than long, apical segments
about as broad as long. Temple rounded behind
eyes, about equal to or slightly shorter than eye in
dorsal view. Occiput concave. Ocelli small, OO =
4.0 OD, almost on line. Frons moderately depressed
behind antenna. Eyes slight protuberant, with
dense and distinct hairs. Malar space shorter, about
0.5 time as eye high. Gena strongly constricted in
face view. Face (Fig. 8) protuberant, coarsely rug-
ose-punctate, about 1.5 times as broad as high.
Clypeus not distinctly divided from face, slightly
convex, its apical border more or less straight,
with a very small tooth medially.
Mesosoma. — Pronotum protecting in front of
mesonotum, dorsolaterally foveate. Notauli shal-
low foveolate, rest of mesonotum punctate except
an areolate-rugose area posteromedially . Precoxal
suture not very distinct, mesopleuron generally
coarsely rugose-punctate, sometimes areolate-
rugose. Propodeum coarsely areolate-rugose, di-
vided by a transverse carina which is raised into a
medial pair and a lateral pair of stout dentate
flanges. Hind coxa smooth. Vein r of forewing 1.0-
1.3 times as long as 3-RS.
Metasoma. — Carapace shorter, CL/CW = 1.5-
1.8, areolate-rugose, oval, deeper in lateral view.
Ventral opening of carapace distinctly in front of
apex. Hypopygium short. Ovipositor short, its
sheath clavate.
Color. — Black; carapace always yellow at base;
all legs yellow except hind femur and tibia apically
in fu scare.
Males. — Same as female except antennae not
dilated medially and carapace black at base.
Holotype Female.— TAIWAN: Tungpu, XI-18-
21-82, K. C. Chou and S. C. Lin. Deposited in TARI.
Paratopes. — TAIWAN: 4 females, 1 male, same
as holotype; 3 females, 2 males, Tungpu, IX-25-29-
80, L. Y. Chou and T. Lin; 1 female, Tungpu, IV-28-
V-2-81, T Lin and C. J. Lee; 2 females, 1 male,
Tungpu, X-5-8-81, T. Lin and W. S. Tang; 1 female,
Tungpu, XI-18-23-81, T. Lin and W. S. Tang; 6
females, Tungpu, X-XI-85, Malaise trap, K. S. Lin;
4 males, Tungpu, IV-16-VII-27-84, K. C. Chou and
C. H. Yung; 2 females, 2 males, Lienhuachi, III-VII-
84, Malaise trap, K. S. Lin and K. C. Chou. Depos-
ited in FAC, TARI, USNM.
Hosts. — Unknown.
Distribution. — Taiwan.
Remarks. — Previous authors (Baker 1926, Shaw
1983, Zettel 1990) treated glabrous eyes as a char-
acter to distinguish Ascogaster. Strikingly this spe-
cies has distinct and dense setae on its eyes, but
except for this character it agrees well with typical
Ascogaster. Its very small body and the character-
istics of the clypeus are also useful to distinguish
this species from other Chinese Ascogaster.
Etymology. — The specific name is the diminu-
tive form of the Latin seta meaning bristle in
reference to the unusually hairy eyes.
Ascogaster townesi Tang and Marsh,
new species
Figs. 12, 26
Females. — Length of forewing 4.3-4.5 mm, of
body 4.6-4.9 mm.
Head. — Antenna with 23 flagellomeres, me-
dial segments not dilated, all segments longer
than broad. Temple rounded behind eyes, slightly
longer than eye in dorsal view. Occiput deeply
concave. Ocelli small, 00= 4.0-4.5 OD, ocellar
triangle acute, ocelli not on line. Frons behind
antenna slightly depressed, sparsely punctate. Eyes
moderately protuberant, glabrous with few scat-
tered minute setae. Malar space short. Gena in face
view strongly constricted. Face (Fig. 12) slightly
convex, about twice as broad as high, less hairy,
the hairs on the upper part of face pointing up-
wards, finely punctate, with a medial carina on the
Volume 3, 1994
295
upper part. Clypeus slightly protuberant, more
scatteredly punctate than face; apical border
straight, without medial teeth or flanges.
Mesosoma. — Pronotum projecting little in front
of mesonotum, dorsolaterally smooth, shallow
fovea te. Notauli deep, foveolate, rest of mesonotum
punctate except with a depressed and areolate-
rugose area posteromedially . Precoxal suture shal-
low foveate; rest of mesopleuron sparsely punc-
tate, smooth except anterodorsally foveolate.
Propodeum not distinctly divided into dorsal and
posterlateral surfaces; strongly rugose with no
tubercles. Hind coxa smooth. Vein r of forewing
1.0-1.2 times as long as 3-RS.
Metasoma. — Carapace (Fig. 26) very long, CL/
CW = 2.3-2.5, clavate in dorsal view, flat in lateral
view, areolate-rugose, apically polished. Ventral
opening of carapace at apex of carapace.
Hypopyium large and broad. Ovipositor long,
slender and upcurved.
Color. — Black; legs yellow-brown except hind
coxa basally, hind and middle femora and tibia
apically and tarsi brownish; antenna infuscate
except scape reddish brown; palpi yellowish
brown.
Males. — Same as females except antennae
longer with 32 flagellomeres, flater carapace and
yellow hind coxa.
Holotype Female. — TAIWAN: Meifeng 2150 m,
V-10-83, H. Townes. Deposited in AEIG.
Paratypes.— TAIWAN: 1 male, Meifeng 2150
m, IV-19-21-83, K. C. Chou and P. Huang; 4 males,
Meifeng.2150 m, IV-26-83, H. Townes; 1 female, 30
males, Meifeng 2150 m, V-3-83, H. Townes; 2
females, 17 males, same as holotype. Deposited in
AEIG, FAC, TARI, USNM.
Host. — Unknown.
Distribution. — Taiwan.
Remarks. — This species is easily distinguished
from other species of Ascogaster by the hairs on the
upper part of the face, very flat carapace, no tu-
bercle on its clypeus, and long, upcurved oviposi-
tor.
Etymology. — It is a pleasure to name this spe-
cies in honor the late Henry Townes who collected
many of the type series and who contributed greatly
to our knowledge of the World Ichneumonoidea
during his lifetime.
Ascogaster varipes Wesmael
Fig. 14
Ascogaster imnpes Wesmael 1835: 234. Lectotype female, BEL-
GIUM: Brussels (designated by Huddleston 1984).
Shenefelt 1973: 837; Huddleston 1984: 370; Tobias 1986:
305; Papp 1989:298.
Ascogaster cavifrons Thomson 1874: 585. Lectotype female ,
SWEDEN: Skane, Torekov (desiganted and syn. by
Huddleston 1984). Shenefelt 1973: 818.
Ascogaster sternalis Thomson 1874: 587. Lectotype female,
SWEDEN: Smaland (designated by Huddleston 1984
and syn. by Telenga 1941).
Ascogaster jaroslawensis Kokujev 1895: 86. Holotype female,
USSR: 'Jaroslaw' (syn. by Huddleston 1984 ). Shenefelt
1973: 824; Tobias 1986: 304 (as a valid species).
Ascogaster variipes; Telenga 1941: 322.
Diagnosis. — Antenna with 32-36 flagellomeres,
slightly dilated medially (females); ocellar triangle
obtuse, ocelli almost on line; face (Fig. 14) gener-
ally rugose-punctate, all hairs downwards; clypeus
with its apical border transversely impressed,
without medial tooth or excision; mandibles with
a deep semicircular depression at base; notauli
distinct, foveolate, rest of mesonotum punctate
except posteriorly areolate-rugose; mesopleuron
coarsely rugose so that precoxal suture not easily
distinguished; propodeum completely areolate-
rugose; hind coxa strongly transversely strigate;
carapace short, oval, deep in lateral view, ventral
opening of carapace short, distinctly in front of
apex.
Specimens Examined.— SHANDONG PROV-
INCE: 2 females, Lao-shan, 800 m.
Additional Specimens Examined. — Lectotype of
varipes Wesmael, female, BELGIUM: Brussels
(IRSNB). Lectotype and paralectotypes of varipes
Wesmael, BELGIUM: 2 females, 3 males (IRSNB).
Hosts.— Shenefelt (1973) listed many host
records. However, much of this information is of
little value because of doubt about the accuracy of
identification of the parasite species involved (see
Huddleston 1984).
Distribution. — China (Shandong Province),
Korea, "USSR" and many other European coun-
tries (see Huddleston 1984).
Remarks. — This species is characterized on
account of the semicircular depression at the base
of the mandibles, the strongly strigate hind coxa,
the characteristics of the clypeus and the shape of
the carapace. The two female Chinese specimens
examined here differ from the type series of imripes
296
Journal of Hymenoptera Research
Wesmael in having a rugose-punctate face instead
of a finely areolate-rugose face as in the type series.
Ascogaster yunnanica Tang and Marsh, new
species
Male. — Length of forewing 2.7 mm, of body
3.3 mm.
Head. — Antenna incomplete, 1st flagellomere
about 3.0 times as long as broad, 2-6 th flagellomeres
2.0-2.5 times as long as broad. Temple constricted
behind eyes, slightly shorter than eye in dorsal
view. Occiput deeply concave. Ocelli small OO =
4.0 OD; ocellar triangle acute, ocelli not on line.
Frons behind antenna moderately depressed,
smooth. Eyes slightly protuberant, glabrous with-
out distinct setae. Malar space short. Gena in face
view strongly constricted. Face slightly convex,
finely and sparsely punctate, about twice as broad
as high, the hairs on the upper part of face pointing
upwards. Clypeus slightly protuberant, more
sparsely punctate than face, apical border straight
without flange or tooth.
Mesosoma. — Pronotum projecting little in front
of mesonotum, deep foveate dorsolateraly . Notauli
foveolate, coalescing posteriorly in a fine areolate-
rugose area; rest of mesonotum punctate. Precoxal
suture shallow foveate anterodorsally, indistinct
posteroventrally; rest of mesopleuron smooth
posteroventraly, punctate anterodorsaly.
Propodeum not distinctly divided into dorsal and
posterolateral surfaces, strongly rugose but with
no dentates. Hind coxa smooth. Vein r of forewing
about twice as long as 3-RS.
Metasoma. — Carapace very long, CL/CW =
2.5, oval in dorsal view, deeper and not so strongly
flat in lateral view, areolate-rugose, but sparsely
punctate apically. Ventral opening of carapace
long, at apex of carapace.
Color. — Black; all legs yellow except hind coxa
mostly black, hind femur and tibia apically and
tarsi infuscate; palpi yellow-brown.
Female. — Unknown.
Holotype Male.— YUNNAN PROVINCE:
Kunming, 111-30-81, J. H. He. Deposited in ZAU.
Host. — Unknown.
Distribution. — China (Yunnan Province).
Remarks. — Morphologically this species is simi-
lar to townesi from which it differs not only in the
characters mentioned in the key but aslo in the
smaller body and the less flat carapace.
Etymology. — The species name is in reference
to the type locality.
ACKNOWLEDGEMENTS
We thank the following people for loan of
types and much of unidentified specimens or for
providing valuable information: C. van Achterberg
(The Netherlands), A. Austin (Australia), H. F.
Chao (FAC), X. X. Chen (ZAU), L. Y. Chou (TARI),
P. Dessart (IRSNB), J. H. He (ZAU), T. Huddleston
(BMNH), K. Maeto (UEI), I. Nauman (Australia),
J. Papp (HNHM), D. J. Preston (BM), M. J. Sharkey
(CNC), S. R. Shaw (Wyoming), G. N. Shida (BM),
M. Suwa (UEI), D. Wahl (AEIG), J. Y. Wang (ZRI).
We also thank C. M. Liu (FAC) for help with
translations of some Russian papers. The senior
author offers special thanks to Prof. H. F. Chao for
his continued helpfulness and encouragement and
to Chunying for her patience and understanding.
This work was partly supported by a grant to Y. Q.
Tang by the National Educational Commission of
the People's Republic of China.
LITERATURE CITED
Achterberg, C. van. 1976. A preliminary key to the subfami-
lies of the Braconidae (Hymenoptera). Tijdschrift voor
Entomologie 119: 33-78.
Achterberg, C van. 1988. Revision of the subfamilv Blacmae
Foerster (Hymenoptera: Braconidae). Zoologische
Verhandelingen 249: 1-324.
Achterberg, C. van. 1990a. Illustrated kev to the subfamilies
of the Holarctic Braconidae (Hvmenoptera:
Ichneumonoidea). Zoologische Mededelmgen 64: 1-20
Achterberg, C. van. 1990b. Revision of the western Palaearctic
Phanerotomini (Hymenoptera: Braconidae). Zoologiche
Verliandelingen 255: 1-106.
Baker, C. F. 1926. Braconidae-Cheloninae of the Philippines,
Malaya, and Australia. The Philippine Journal of Science
31: 451-489.
Bovce, H R. 1936. Laboratory breeding of Ascogaster
carpocapsae Viereck with notes on biology and larval
morphology. Canadian Entomologist 68: 241-246.
Cox, J. A. 1932. Ascogaster carpocapsae Viereck, an important
parasite of the codling moth and the oriental fruit moth.
Technical Bulletin of the New York State Agricultural Ex-
periment Station 188: 1-26.
Fahringer, J. 1934. Opuscula Braconologica, Palaearktische Re-
gion 2: 321-594.
Fullaway, D. T. 1919. New genera and species of Braconidae
mostly Malayan, journal of Straits Branch, Royal Asiatic
Society 80: 39-61.
Harris, R. A. 1979. A glossary of surface sculpturing. Occa-
sional Papers in Entomology, California Department of Food
and Agriculture 28: 1-31.
Volume 3. 1 994
297
He, J. H., X. X Chen and Y. Ma. 1989. The Braconid (Hy-
menoptera) parasites of Adoxophyes orana Fischer von
Rosterstamm (Lepidoptera: Tortncidae) from China
Acta Agricultae Universitatis Zhejiangensis 15 (4): 437-439.
Huddleston, T. 1984. The Palaearctic species of Ascogaster
(Hymenoptera: Braconidae). Bulletin of the British Mu-
seum (Natural Htston/) (Entomology) 49: 341-392.
Kainoh, Y. 1986. Mating behavior of Ascogaster reticulatus
Watanbe (Hymenoptera: Braconidae, an egg-larval para-
sitoid of the smaller tea tortrix moth, Adoxophyes sp.
(Lepidoptera: Tortncidae) I. Diel patterns of emergence
and mating, and some condintions for mating. Applied
Entomology and Zoology 21: 1-7.
Kainoh, Y. 1988. Some factors influencing sex ratio in
Ascogaster reticulatus Watanabe (Hymenoptera:
Braconidae). Applied Entomology and Zoology 23:35-40.
Kainoh, Y., T. Hiyori and Y. Tamaki. 1982. Kairomone of the
egg-larval parasitoid, Ascogaster reticulatus Watanabe
(Hymenoptera: Braconidae). Applied Entomology and Zo-
ology \7: 102-110.
Kainoh, Y., T. Nemoto, K. Shimizu, S. Tatsuki, T. Kusano and
Y. Kuwahara. 1991. Mating behavior of Ascogaster
reticulatus Watanabe (Hymenoptera: Braconidae), an
egg-larval parasitoid of the smaller tea tortrix, Adoxophyes
sp. (Lepidoptera: Tortricidae) II. Identification of a sex
pheromone. Applied Entomology and Zoology 26: 543-549.
Kainoh, Y. and Y. Tamaki. 1982. Searching behavior and
oviposition of the egg- larval parasitoid, Ascogaster
reticulatus Watanabe (Hymenoptera: Braconidae). Ap-
plied Entomology and Zoology 17: 194-206.
Kainoh, Y. and S. Tatsuki. 1988. Host egg kairomones essen-
tial for egg-larval parasitoid, Ascogaster reticulatus
Watanabe (Hymenoptera: Braconidea I, Internal and
external kairomones. journal of Chemical Ecology 14: 1475-
1484.
Kainoh, Y., S. Tatsuki. and T. Kusano. 1990. Host moth scales;
a cue for host location for Ascogaster reticulatus Watanabe
(Hymenoptera: Braconidae). Applied Entomology and
Zoology 25: 17-25.
Kainoh, Y., S. Tatsuki, H. Sugie and Y. Tamaki. 1989. Host egg
kairomones essential for egg-larval parasitoid, Ascogaster
reticulatus Watanabe (Hymenoptera: Braconidae) II.
Identification of internal kairomone. Journal of Chemical
Ecology 15: 1219-1229.
Kamano, Y., K. Shimizu, Y. Kainoh and S. Tatsuki. 1989.
Mating behavior of Ascogaster reticulatus Watanabe (Hy-
menoptera: Braconidae), an egg-larval parasitoid of the
smaller tea tortix, Adoxophyes sp. (Lepidoptera:
Tortricidae). Applied Entomology and Zoology 24: 372-378.
Kawakami, T. 1985. Development of the immature stages of
Ascogaster reticulatus Watanabe (Hymenoptera :
Braconidae), an egg-larval parasitois of the smaller tea
tortrix moth, Adoxophyes sp. (Lepidopatera: Tortricidae).
Applied Entomology and Zoology 20: 380-386.
Kawakami, T. and Y. Kainoh. 1985. Host discrimination and
competition in the egg larval parasitoids, Ascogaster
reticulatus Watanabe (Hymenoptera: Braconidae). Ap-
plied Entomology and Zoology 20: 362-364.
Kawakami, T. and Y. Kainoh. 1986. Host specificity of the
egg-larval parasitoid, Ascogaster reticulatus Watanabe
(Hymenoptera: Braconidae) among five tortricid and
one noctuid species. Applied Entomology and Zoology 21 :
8-14.
Papp, ]. 1989. Braconidae (Hymenoptera) from Korea XI. Acta
Zoologica Hunganca 35(3-4): 295-326.
Quicke, D. L. J. and C. van Achterberg. 1990. Phy logeny of the
subfamilies of the family Braconidae (Hymenoptera:
Ichneumonoidea). Zoologische Verhandehngen 258: 1-95.
Rosenberg, H. F. 1934. The biology and distribution in France
of the larval parasites of Cydia pomonella L. Bulletin of
Entomological Research, 25: 201-256.
Shaw, M. R. and T. Huddleston. 1991. Classification and
biology of braconid wasps (Hymenoptera: Braconidae).
Handbooks for the Identification of British Insects 7 (11): 1-
126.
Shaw, S.R. 1983. A taxonomic study of Nearctic Ascogaster
and a description of a new genus Leptodrepana (Hy-
menoptera: Braconidae). Entomography 2: 1-54.
Shenefelt, R. D. 1973. Catalogus Hymenopterorum (Noi<. ed. part
10 Braconidae, 6:813-936. s'Gravenhage.
Sonan, J. 1932. Notes on some Braconidae and Ichneumomdae
from Formosa with descriptions of 18 new species.
Transactions of the Natural History Society of Formosa 22:
66-86.
Szepligeti, G. V. 1905. Exotische Braconiden aus den
Aethiopischen, Dnentalischen und Austtralischen
Regionen. Annates Musei Nationalis Hungarici 3: 25-55.
Szepligeti, G. V. 1908. Braconidae und Ichneumomdae. Die
Fauna Sudtoest- Australiens 1(9): 317-324.
Telenga, N. A. 1941. Insects Hymenoptera, Family Braconidae,
Subfamily Braconinae (continued) and Sigalphinae
Fauna SSSR 5 (3): 1-466.
Tobias, V. I. 1986a. New species of Subfamily Cheloninae
(Hymenoptera: Braconidae) from the Far East of the
USSR. Trudy Zoologicheskogo Instituta Akademiya Nauk
SSSR 159: 3-17 (in Russian).
Tobias, V. I. 1986b. Guide to the insect of European part of the
USSR Hymenoptera. Opredeliteli Faune SSSR, 3 (4): 1-500
(in Russsian).
Tobias, V. I. 1988. Two new species of Braconidae of the
subfamlily Cheloninae (Hymenoptera) from the pro-
tected territoties of the Liruanian SSR. Acta Entomology
Lituaman 9: 89-94.
Walker, A. and T. Huddleston. 1987. New Zealand chelonine
braconid wasps (Hymenoptera: Braconidae). journal of
Natural History 21: 339-361.
Watanabe, C. 1937. A contribution to the knowledge of the
Braconid fauna of the Empire of Japan, journal of the
Faculty of Agriculture Hokkaido Imperial University 42(1):
1-188."
Watanabe, C. 1967. Description of a new species of the genus
Ascogaster Wesmael and notes on synonym of Apanleles
species (Hymenoptera: Braconidae). Insecta
Matsumurana 29(2): 41-44.
Wesmael, C. 1835. Monographic des Braconides de Belgique.
Noui'eaux Memoires de V Academic Royale des sciences et
belles-lettres de Bruxelles 9: 1-252.
Yoneda, Y. 1978. A new species of the genus Ascogaster
Wesmael (Hymenoptera, Braconidae) from Japan.
Kontyu 46: 291-296.
Zettel, V.H.I 990. Eine Revision der Gattungen der Cheloninae
(Hymenoptera, Braconidae) mit Beschreibungen neuer
298
Journal of Hymenoptera Research
Gattungen und Arten. Annates Naturlustorisches Mu-
seum Wien 91(B): 147-196.
NOTE ADDED IN PROOF
The following paper came to our attention after
our manuscript was sent to the printer: Chen
Jiahua, Huang Juchang and Wu Zhishan. 1994.
Notes on two new species and six new records of
the genus Ascogaster Wesmael from China (Hy-
menoptera: Braconidae: Cheloninae). Journal of the
Fujian Agricultural College (Natural Sciences Edi-
tion) 23(1)51-57. Although we were not able to
study the specimens used by Chen et al., we have
made comments below on each species mentioned
by them based on a translated version of their
paper.
Ascogaster albitarsis Sonan. The characters mentioned in
their key are not significant to separate this species from
reticulatus based on our study (see discussion under
reticulatus).
Ascogaster arisanica Sonan and Ascogaster bidentula
Wesmael. During our study we examined the type ma-
terial of arisanica, bidentula and atamiensis (synonym of
bidentula). All of the material we examined under the
name arasanica agreed with the type but none agreed
with types of bidentula or atamiensis. Furthermore, we
found that the characters used by Chen et al. and
Huddelston (1984) are not realiable in distinguishing
bidentula and arisanica (see our discussion of arasanica).
Most of the material examined by Chen et al. of bidentula
and arasanica was collected at the same locality and
same date; thus we suspect that their material is prob-
ably arasanica. This problem is further complicated by
the fact that atamiensis was recorded from China by
Fahringer (1938, Ark. Zool. 30:3) and Shenefelt (1973).
Obviously this species complex needs to be studied
further
Ascogaster consobrina Curtis and Ascogaster inf acetus Chen
and Huang. The new species described in Chen et al.,
mfacetus, will run to consobrina in our key. Fig. IB in
Chen et al. which is an illustration of the mesopleuron is
not clear. Thus, we must reserve comment on this spe-
cies until the type can be observed
Ascogaster longicornis Huddleston. We treated this species
as a synonym olformosensis based on type examination.
Ascogaster perkinsi Huddleston. See notes under wuyiensis
below.
Ascogaster reticulatus Watanabe. Although this species was
mentioned in their key, it was not discussed in the text.
Thus, we must see their specimens before deciding
upon the relationship of this species and albitarsis.
Ascogaster ruf idens Wesmael. We did not find this species in
any of the material that we studied. Again, their mate-
rial should be studied before we confirm this species is
in China.
Ascogaster wuyiensis Chen and Huang. This species will
run to perkinsi in our key based on the description in
Chen etal, and the color of the clypeus and theshapeof
the yellow mark at the base of the carapace may be only
variation in perkinsi. We described a new species, lini,
which is related to wuyiensis and perkinsi. The validity of
these three species must wait until specimens of wuyiensis
are examined.
Figs. 1-2. Fore wings, somewhat diagrammatic. 1, Ascogaster. 2, Chelonus. SM=submarginal cells; D=discal cell.
Volume 3. 1994
299
Figs. 3-8. Faces of Ascogaster species. 3, arisanica Sonan. 4, dimorpha n. sp. 5 , formosensis Sonan. 6, grandis n. sp. 7,fullaivayi
(Baker). 8, setula n. sp.
300
Journal of Hymenoptera Research
Figs. 9-14. Faces of Ascogaster species. 9, lini n. sp. 10, perkinst Huddleston. 11, stmenoi'i Telenga. 12, townesi n. sp. 13, rugulosa
n. sp. 14, varipes Wesmael.
Volume 3. 1 994
301
Figs. 15-20. Heads of Ascogaster species. 15-16, hein.sp.: 15, dorsal view; 16, face. 17-18, quadridentataVJesmael. 17, dorsal view;
18, face. 19, armatoides n. sp. 20, consobrina Curtis.
302
Journal of Hymenoptera Research
■. ■ .■;.■■■ .:
' "■" -^^^Sk
,.'-' '
,--■-, . ■ -" . - .
■"- i^v^B
*>.! 'T/ >. ■* ''
■ '• ' "•'-' • ^~
• ' - r
-""^IS^sJHf
^■'^-:>L*
v.r:;;.;:- .■•.,•;..•.
^titm
K ©
Mm
mm
Figs. 21-28. Morphology of Ascogaster species. 21-26, metasomal gaster: 21, grandis n. sp.; 22, perkmsi Huddleston; 23, acutus
J. HYM. RES.
Vol. 3, 1994, pp. 303-308
Systematics of Pseudomethoca areta (Cameron): Sex association,
description of the male and a gynandromorph, and
a new synonymy (Hymenoptera: Mutillidae).
DlOMEDES QUINTERO A. AND ROBERTO A. CaMBRA T.
Museo de Invertebrados G.B. Fairchild, Universidad de Panama, Estafeta Universitaria, Panama, Repiiblica de Panama;
(DQA) Smithsonian Tropical Research Institute, Unit 0948, APO AA 34002-0948.
Abstract. — A gynandromorph of Pseudomethoca areta (Cameron, 1895) is described, and previously published cases of
gynandromorphism in Mutillidae are reviewed. Sex association permits recognition of the undescribed male of P. areta. We
place Pseudomethoca panamensis (Cameron, 1895) in New Synonymy with P. areta.
Resumen. — Se describe un individuo ginandromorfo de Pseudomethoca areta (Cameron, 1 895) y se presenta un resumen de casos
previamente descritos de ginandromorfia en Mutillidae. Se lleva a cabo la asociacion sexual y se describe al macho de P. areta,
hasta ahora desconocido. Pseudomethoca panamensis (Cameron, 1895) se coloca como Nueva Sinonimia de P. areta.
INTRODUCTION
Mutillids are solitary parasitoid wasps that
exhibit great sexual dimorphism, making sex as-
sociations difficult. The New World genus
Pseudomethoca Ashmead illustrates this problem;
only one-fifth of its 103 described species are known
from both sexes. Distinctly fewer sex associations
have been obtained for Neotropical than for Ne-
arctic species of Pseudomethoca. Seventeen out of
45 Nearctic species known (37.8%) have both sexes
recognized (Krombein, 1992). Of the remaining 28
species, 20 are known only from females, and
eight only from males. In contrast, only four of
some 58 Neotropical species of Pseudomethoca are
known from both sexes (6.9%) (Nonveiller, 1990;
Cambra & Quintero, 1992). Of the remaining 54, 46
are known from females only, and eight from
males only. Success in associating the sexes will
facilitate future biological work on the group and
will solve some of the annoying taxonomic prob-
lems. We suspect that only about one-third of the
species of Pseudomethoca in the Neotropics have
been described. Previous taxonomic work on
Pseudomethoca was done by Mickel (1924, 1935,
revision of North American species; 1952, key
to females of Guyanan species), Schuster (1945,
key to Caribbean species), and Krombein (1992).
Gynandromorphy is a developmental phe-
nomenon useful for associating the sexes in some
extremely dimorphic animals, including mutillids
(Mickel 1928, 1936, 1952; Bischoff 1931). Unfortu-
nately, gynandromorphs are rare in Mutillidae.
After examining more than 15,000 mutillid speci-
mens, we have discovered only two gynandro-
morphs. A review of the literature revealed only
six previously published cases (Table 1). We re-
port here the second known Neotropical mutillid
gynandromorph. We recently discovered a gy-
nandromorph of Timulla labdace Mickel, from
Panama, that will be described in a separate pub-
lication.
304
Journal of Hymenoptera Research
Pseudomethoca areta (Cameron)
Figs. 1 - 4
Sphaerophthalma areta Cameron, 1895: 332, pi. 14, fig. 12,
female. Bugaba, Chinqui Province, Panama, Champion
col., BM(NH), London, Type 15.822, examined;
Pseudomethoca areta: Cambra & Quintero, 1992: 474.
Sphaerophthalma panamensis Cameron, 1895: 334-35, female.
Bugaba, Chinqui Province, Panama, Champion col.,
BM(NH), London, Type 1 5.833, examined; Pseudomethoca
panamensis: Cambra & Quintero, 1992: 475. NEW SYN-
ONYMY.
Diagnosis. — In Mickel's revised key (1935) it
runs to P. vanduzei Bradley in couplet five. The
male of P. areta differs from P. vanduzei as follows:
posterior half of tegula bent downward so as to
form a posterior face at a sharp angle with the
dorsal surface (in P. vanduzei the tegula is uni-
formly convex, without a posterior face); anterior
margin of clypeus in P. areta is bidentate (it lacks
teeth in P. vanduzei); mandibles tridentate at the tip
in P. areta (bidentate in P. vanduzei); integument of
abdomen mostly ferruginous in P. areta (totally
black in P. vanduzei). Pseudomethoca areta is en-
demic to Panama, and P. vanduzei is present in the
southeastern United States (Krombein, 1979).
Description. — Integument black, except apex
of tergum one and abdominal segments two to six,
orange. Head large, subrectangular in dorsal view,
as wide as thorax, clothed with long, erect and
recumbent white pubescence; row of six to eight
long, erect, dark hairs near inner margin of eyes;
posterolateral angles of head not dentate. Man-
dibles tridentate distally; clypeus strongly
bidentate medially on the cephalic margin; disk of
clypeus densely punctate. Scape with a strong
longitudinal carina beneath; first flagellomere
equal in length to second. Front, vertex and genae
coarsely and confluently punctate. Antennal
scrobes and genae not carinate. Ocelli small, dis-
tance between eye margin and lateral ocelli equal
to approximately five times the greatest diameter
of the latter.
Pronotum, mesonotum and scutellum with
close, more or less confluent punctures, punctures
about the size of those on head. Propodeum
strongly and coarsely reticulate dorsally and pos-
teriorly. Tegula punctate throughout; posterior
part of tegula bent downward so as to form a
posterior face at a sharp angle with the dorsal
surface. Humeral angles of pronotum without any
evidence of a carina. Propleura and mesopleura
with close punctures; metapleura smooth, with-
out punctures; sides of propodeum with only a
few scattered punctures. Pronotum, scutellum,
metanotum and dorsal face of propodeum, clothed
with sparse, long erect white pubescence;
mesonotum clothed with sparse black pubescence;
tegula clothed with intermixed, black and white
pubescence; metapleura and sides of propodeum
almost bare, with only sparse white
micropubescence.
Anterior and intermediate coxae without teeth
or keels; posterior coxa with a keel on inner mar-
gin. Legs clothed with sparse white pubescence.
Calcaria pale.
Abdomen with segment one completely sessile
with second. Terga one and two with small, sepa-
rated punctures, except the apical margins with
close punctures; terga three to six with small, close
punctures. Pygidium rugose. Felt line 0.6 x as long
as lateral margin of tergum two. Sternum one
almost smooth, with only a few, sparse punctures,
and with a low, median longitudinal carina on
anterior two-fifths. Sternum two with sparse, mod-
erate punctures. Sterna three to six with close,
moderate punctures. Posterior margin of
hypopygium evenly convex. Tergum one clothed
with sparse, long erect, white pubescence. Terga
two to six with sparse long erect, orange pubes-
cence, the apical margins with a band of dense,
recumbent, orange pubescence. Last tergum
clothed with black pubescence. Sterna one to six
clothed with white pubescence, except lateral
margins of sterna two to four, with orange pubes-
cence. Last sternum clothed with white pubes-
cence and a few intermixed fuscous hairs. Wings
infuscated, especially apically; forewing with two
well defined submarginal cells and traces of a
third. Body length; 10.6 mm.
ALLOTYPE Male Information.— PANAMA:
Darien Province, Cruce de Mono, Estacion
INRENARE, 8 Feb 1993 (yellow trap). R. Cambra
T., J. Coronado, Museo de Invertebrados "G. B.
Fairchild", Universidad de Panama (MIUP).
Additional Material Examined. — PANAMA:
Darien Province, Cruce de Mono, Estacion
INRENARE, R. Cambra, J. Coronado, 144 females
and 27 males, 8 Feb - 4 Mar 1993, deposited in
MIUP, NMNH-Smithsonian Institution, Univer-
sity of Minnesota Insect Collection, and BM(NH).
Body length varies in males from 8.3 to 1 1 .0 mm, in
Volume 3, 1 994
305
Figs. 1-4. Pseudomethoca areta. 1. Male allotype, dorsal habitus. 2. Gynandromorph, dorsal habitus. 3.
Male genitalia, split drawing, dorsal = right; ventral = left. 4. Penis valve, side view.
306
Journal of Hymenoptera Research
females from 7.8 to 11.1 mm.
Comments on Sex Association and Nezo Syn-
onymy. — Sex association is based on observations
of males courting and mating with females in the
field, and has been corroborated in the laboratory
with mating experiments. Courtship is very brief,
lasting 15-40 seconds, and consists of bursts of
rapid vibrations of the wings and antennae, inter-
spersed by short hopping flights. The male climbs
onto the female and grasps her neck with his
mandibles and attempts to mate with her. We
have not seen heterospecific courtship or mating
in Psendomethoca. Erroneous heterospecific sex as-
sociations may be made if containers or outdoor
sites become contaminated by pheromones re-
leased from a female of a different species that
recently occupied that site (personal observations).
Males of Psendomethoca areta, like those of the
genotype and males of numerous other species of
Pseudomethoca we have examined, have a strong
longitudinal carina beneath the scape. Therefore,
we consider erroneous Casal's (1965) observation
that males of the ill-defined genus Darditilla Casal
(genotype is only male known, and 35 other spe-
cies, known only from females) differ from those
of Psendomethoca in having a carina beneath the
scape, said to be absent in the latter. The male
genitalia of P. areta are symmetrical (Fig. 3), as are
those of all other Neotropical sphaeropthalmine
males we have examined, and phoretic mating is
absent in this group. In contrast, the Neotropical
mutilline males of Timulla (Timnlla) present
strongly asymmetrical genitalia and phoretic mat-
ings (Cambra & Quintero, 1992). The asymmetry
in the male genitalia possibly functions to provide
a better hold, or grasp, of the female while air-
borne.
Cameron's types of areta and panamensis are
both from Bugaba, and we found them to be
identical; the name areta has page precedence over
that of panamensis. Psendomethoca areta is closely
related to P. hecate (Gerstaecker, 1874), from Costa
Rica, differing only in the integumental coloration
of the vertex and dorsolateral areas of the thorax.
We suspect that they are the same species. To
confirm the synonymy we need to examine
Gerstaecker's type specimen and to compare the
genitalia of males sexually associated with hecate
females with those of the males described here.
Gynandromorph Individual of
Psendotnethoca areta
Fig. 2
Description. — Head identical to that of a nor-
mal female, without a trace of male characters.
Thorax and legs identical to those of a normal
male, without a recognizable trace of female char-
acters. The anterior wings have abnormally thin,
translucent venation; they are torn along their
posterior half, and we suspect were never func-
tional. Abdomen with six segments, as in the
female. First tergum completely male. Second ter-
gum a mosaic: right half with coloration and pu-
bescence of male and female; left half is com-
pletely male-like. Third tergum, right half with
female characteristics only; left half is a mosaic
with integument coloration and pubescence both
of male and female. Abdominal segments four to
six are female only. Second sternum with right
half female and left half male, same as other sterna,
except for sternum three, identical to that of a
female. Body length: 11.1 mm.
Data and comments on the gynandromorph. —
The gynandromorph individual was collected on
26 February 1993, at 10:00 AM, in the general
locality of the allotype. When we first noticed it,
the individual was walking over dry leaves on the
ground. Shortly afterward, we watched a male
arrive, flying upwind, attracted by what we
thought was a normal female. The male quickly
attempted to mate but encountered indifference
on the part of the female-like individual. Female
mutillids are known to attract winged males by
means of wind-dispersed pheromones (see Cambra
& Quintero, 1993). The upwind arrival of the male
suggests that the gynandromorph individual was
secreting female pheromones. The abnormally thin
and quite battered forewings, suggest that the
animal was unable to fly, although it had perhaps
attempted to. The specimen exhibits anterior/
posterior division of male and female compo-
nents, as well as mosaic segments, a type of gynan-
dromorphy not previously described for mutillids
(see Table 1).
Volume 3. 1994
307
ACKNOWLEDGMENTS
We thank the Smithsonian Tropical Research Institute
(STRI), particularly Ira Rubinoff, for providing research fa-
cilities. We are grateful to the British Embassy in Panama, in
particular Ambassador Thomas H. Malcomson, for securing
travel funds to England for R.A.C.; to the Entomology De-
partment personnel of the Natural History Museum, Lon-
don, BM(NH) for providing R.A.C. with working facilities
and assistance during his visit. We also thank Indra
Candanedo and Roberto Arango, Instituto Nacional de
Recursos Naturales Renovables, who aided us with permits
and logistics in the Darien National Park. This project was
financed in part by Vicerectoria de Investigacion y Postgrado,
Universidad de Panama, Fund No. 1-4500-91-12. Our thanks
to Karl V. Krombein, Smithsonian Institution, Annette Aiello
and David W. Roubik, STRI, and an anonymous reviewer, for
reading and making comments on the manuscript, James
Coronado, now at STRI, for providing very valuable help in
the field, and to Angel Aguirre, STRI, for bibliographic assis-
tance and location of references on gynandromorphs.
LITERATURE CITED
Bischoff, H. 1931. Der typus der Mutilla dubia ¥.— ein
gynander. Mitteilungen deutschen entomologischen
gesellschaft 2(4): 54-56."
Cambra, R. A. and D. Quintero A. 1992. Velvet ants of
Panama: distribution and systematics (Hymenoptera:
Mutillidae), pp. 459-478. In: Quintero A., D. & A. Aiello
(eds.) Insects of Panama and Mesoamerica: Selected studies.
Oxford University Press, Oxford.
Cambra, R. A. and D. Quintero A. 1993. Studies on Timulla
Ashmead (Hymenoptera: Mutillidae): New distribu-
tion records and synonymies, and descriptions of previ-
ously unknown allotypes. Pan-Pacific Entomologist 69(4):
296-310.
Cameron, P. 1895-1896. Hymenoptera, vol. 2. Biologia Centrali
Americana, pp. 262-360, plates 13-14.
Casal, O. H. 1965. Darditilla nuevo genero Neotropical de
Sphaeropthalminae (Hym. Mutillidae). Eos, Madrid, 41:
9-18.
Dalla Torre, C. G. and C. Friese. 1899 (1898). Die
hermaphroditen und gynandromorphen Hymenop-
teren. Berichte des naturxuissenschaftlich - Medizmischen
Vereins in Innsbruck 24: 1-96.
Gerstaecker, C. E. 1874. Mutillarum Americae mendionalis
indigenarun synopsis systematica et synonymica. Archil'
fur Naturgeschiechte 40: 41-77.
Krombein, K. V. 1979. Family Mutillidae, pp. 1276-1314. In:
Krombein, K. V. et al., eds. Catalog of Hymenoptera m
America North of Mexico, Vol. 2. Smithsonian Institution
Press, Washington, D.C
Krombein, K. V. 1992. Host relationships, ethology and sys-
tematics of Pseudomethoca Ashmead (Hymenoptera:
Mutillidae, Andrenidae, Hahctidae and Anthophondae).
Proceedings of the Entomological Society of Washington
94(1): 91-106.
Maeklin, F. W. 1856. Om hermafroditism bland insekterna,
samt beskrifning ofver en i Helsingfors funnen
hermafrodit af Mutilla obscura Nyl. ofver. af Finska
Vetenskaps-Societens Forliandlingar 3: 106-112.
Mann, W. M. 1915. A gynandromorphous mutillid from
Montana. Psyche 22: 178-180.
Mickel, C. E. 1924. A revision of the mutillid wasps of the
genera Myrmilloides and Pseudomethoca ocurring in
America North of Mexico. Proceedings of the United States
National Museum 64(15): 1-51.
Mickel, C. E. 1928. Biological and taxonomic investigations
on the mutillid wasps. United States National Museum
Bulletin 143: 1-351.
Mickel, C. E. 1935. Descriptions and records of mutillid
wasps of the genera Myrmilloides and Pseudomethoca
(Hymenoptera: Mutillidae). Transactions of the American
Entomological Society 61: 383-398.
Mickel, C. E. 1936. New species and records of Nearctic
mutillid wasps of the genus Dasymutilla. Annals of the
Entomological Society of America 29: 29-60.
Mickel, C. E. 1952. The Mutillidae (wasps) of British Guiana.
Zoologica 37(3): 105-150.
Nonveiller, G. 1990. Catalog of the Mutillidae, Myrmosidae and
Bradynobaenidae of the Neotropical region including Mexico
(Insecta: Hymenoptera). SPB Academic Publishing bv,
The Netherlands, 150 pp.
Schuster, R. M. 1945. A new species of Pseudomethoca
(Mutillidae) from the West Indies. Bulletin of the
Brooklyn Entomological Society 40: 7-8.
Wheeler, W. M. 1910. A gynandromorphous mutillid. Psyclie
17: 186-190.
308
Journal of Hymenoptera Research
Table 1. Previously published cases of gynandromorphy in Mutillidae
Species
Type
Wing Country Ref.
& tegula
Mutilla europea obscura bilateral, right f, left m
Finland Maeklin 1856
DallaTorre & Friese 1899
Dasi/nmtilla cypris
[=hora]
head, thorax, abdom. segm.
5-7 m; 1-4 f
USA Mickel 1928
Dasymutilla gloriosa
[=reperticia]
Dasymutilla vestita
[=euchroa
=fulvohirta]
Pseudomethoca frigida
[=canadensis]
TraumatomutiUa dubia
not described
decussated,
head half:
right m, left f;
thorax & abdomen:
right f; left m
bilateral,
right m, left f
head, thorax, 1st
abdom. segment m,
rest abdomen mosaic
USA Mickel 1936
USA Mann 1915
USA Wheeler 1910
Am. Mer. Bischoff 1931
[Guyana] Mickel 1952
Abbreviations: f, female; m, male; +, well developed; 0, absent.
Volume 3, 1994 309
Additions and Corrections to Volume 2, Number 1, 1993
Publication date: the correct date of publication for Volume 2, Number 1, is November 17, 1993.
All ">", "<" and "=" signs were inadvertently left out of the text in the following two articles resulting
in a significant loss of information. The affected text is as follows:
Davidson, D. W. and D. McKey, "The evolutionary ecology of symbiotic ant-plant relationships," pp.
13-83.
P. 14, column 2, line 42 "(N = 8)"
P. 14, column 2, line 43 "(N = 6)"
P. 14, column 2, line 44 "(N = 4)"
P. 14, column 2, line 46 "(N = 5) and "(N = 6)"
P. 15, column 1, lines 11, 12 "(N = 8), (N = 7) and (N > 5)"
P. 15, column 1, line 35 "(N = 50-60 ant-plant species)"
P. 15, column 1, line 36 "(N = 20)"
P. 15, column 1, line 37 "(N = 15)"
P. 15, column 1, lines 38, 39 "(N = 12 species) and (N > 6)"
P. 15, column 1, line 40 "(N = 15)"
P. 15, column 1, line 42 "(N = 3)"
P. 15, column 2, line 6 "(N = 23)"
Additional corretions are as follows:
P. 69, Appendix 1 : reference to Manriquez & Dirzo 1990 should read "Iborra-Manriquez & Dirzo 1990."
P. 70, Appendix 1: entry for Barteria should read ". . . HFOWEb Y Z . . . "
P. 74, Appendix 1: entry for Maieta should read "... B;D; Vasconcelos 1990, 1991, IP; Herre et al. 1986"
P. 78, Appendix 1: entry for Cordia should read ". . . ufRIy Y,i' GH . . ."
Ward, P. S., "Systematic studies on Pseudomyrmex acacia-ants," pp. 117-168.
P. 121, column 2, line 30 "PLI2 > 0.77"
P. 121, column 2, line 45 "worker PLI > 0.71, queen PLI > 0.64"
P. 122, column 1, line 36 "HW < 0.85"
P. 122, column 1, line 37 "HW > 0.85"
P. 122, column 1, line 41 "PL/HW > 0.71"
P. 122, column 2, lines 8-9 "CI > 0.94 and/or HW > 0.96"
P. 122, column 2, line 15 "SL/HL > 0.22"
P. 122, column 2, line 16 "SL/HL < 0.21"
P. 130, line 33 "CI = 1.12"
P. 130, line 35 "CI = 0.80"
P. 132, line 40 "SL/HL < 0.21"
P. 132, line 42 "SL/HL > 0.22"
P. 133, lines 4-5 "CI > 0.94 and/or HW > 0.96"
P. 133, column 1, line 8 "MD8/MD9 = 0.70"
P. 133, column 1, line 11 "MD4/MD5 = 0.74"
P. 135, column 2, line 4 "worker REL < 0.50, queen REL < 0.48"
P. 135, column 2, lines 5-6 "worker PLI < 0.71, queen PLI < 0.63"
P. 135, column 2, lines 29-30 "queen PLI = 0.65, queen PL/HL = 0.49"
P. 145, column 1, lines 26-27 "REL < 0.45, REL2 < 0.56, EL/LHT < 0.61'
310 Journal of Hymenoptera Research
P. 145, column 1, line 30 "FCI = 0.055"
P. 145, column 2, lines 7-8 "worker PWI3 > 0.50, worker PPW1 < 1.30"
P. 146, column 2, line 10 "CI = 0.61"
P. 146, column 2, line 12 "LHT/HW = 1.12"
P. 152, column 2, lines 14-15 "(< 0.10 mm)"
P. 155, column 1, line 10 "REL = 0.66"
P. 157, column 1, lines 20-21 "PLI = 0.55"
P. 157, column 2, line 12 "MD8/MD9 = 0.70"
P. 157, column 2, line 17 "MD4/MD5 = 0.65"
P. 158, column 1, lines 7-8 "PWI3 = 0.60"
P. 158, column 1, line 41 "REL = 0.65"
P. 158, column 1, lines 42-43 "MFC = 0.02"
P. 158, column 1, line 44 "CI = 0.85"
P. 159, column 1, line 3 "CI = 1.12"
Additional corrections are as follows:
P. 163, Table 2, column 31: Pseudomyrmex nigrocinctus and P. particqis should be "1", not "0" for
character 31.
P. 121, column 1, line 41: for "public domain software" read "Shareware software".
P. 131, line 34: for "Figs. 10, 34" read "Figs. 11, 34".
Kazenas, V. L. and B. A. Alexander, "The nest, prey, and larva of Etttomosericus kaufmani", pp. 221-
226. Nearly all references to illustrations should be corrected as follows:
P. 221, column 2, line 19: for "Fig. 2" read "Fig. 4".
P. 221, column 2, line 23: for "Fig. 3" read "Fig. 6".
P. 223, column 1, lines 7-8: for "Fig. 4" read "Fig. 7".
P. 223, column 1, line 13: for "Figs. 5, 8" read "Figs. 8, 11".
P. 223, column 1, line 22: for "Fig. 8" read "Fig. 9".
P. 223, column 1, line 28: for "Figs. 5, 8": read "Figs. 8, 11".
P. 223, column 1, lines 29-30: for "Figs. 10, 11" read "Figs. 2, 3, 8".
P. 223, column 1, line 31: for "Figs. 5, 6" read "Figs. 5, 8".
P. 223, column 1, line 37: for "Fig. 6" read "Fig. 5".
P. 223, column 2, line 3: for "Fig. 7" read "Fig 10".
INSTRUCTIONS FOR AUTHORS
General Policy
The journal of Hymenoptera Research invites papers of high scientific quality reporting comprehensive research on all
aspects of Hymenoptera, including biology, behavior, ecology, systematics, taxonomy, genetics, and morphology. Taxonomic
papers describing single species are unlikely to be accepted unless a strong case is evident, such as importance in economic
entomology or with concurrent biology or ecology. Manuscript length generally should not exceed 50 typed pages; however,
no upper limit on length has been set for papers of exceptional quality and importance, including taxonomic monographs at
generic or higher level.
All papers will be reviewed by at least two referees. The referees will be chosen by the appropriate subject editor. However,
it would be helpful if authors would submit the names of two persons who are competent to review the manuscript.
The language of publication is English. Summaries in other languages are acceptable.
Format and Preparation
Three copies of each manuscript, including copies of illustrations, should be submitted on letter size or A4 paper, double
spaced, with at least 25 mm margins on all sides. On the upper left of the title page give name, address and telephone and fax
numbers of the author to whom all correspondence is to be sent.
The paper should have a concise and informative title, followed by the names and addresses of all authors. The sequence
of material should be: title, author(s), abstract, text, acknowledgments, literature cited, appendix, figure legends, figure copies
(each numbered and identified), tables (each numbered and with heading). Each of the following should start a new page: (1)
title page, (2) abstract, (3) text, (4) literature cited, (5) figure legends, (6) footnotes.
Following acceptance of the manuscript, the author should provide the editor with two copies of the manuscript
accompanied by a copy on diskette using DD, double sided computer diskettes - IBM compatible MS DOS 5.25 inch or IBM and
Macintosh 3.5 inch diskettes. (Authors who do not have access to a computer should submit three copies of the manuscript.)
The paper may be submitted in most PC and Mac word processor programs such as Microsoft Word, FullWrite Professional,
WordPerfect, WriteNow, Nisus, MacWrite, or MacWrite II; it is important that the manuscript also be formatted as an ASCII
or TEXT file. If possible, all words that must be italicized should be done so, not underscored. Use of the words male and female
is preferred to sex symbols. Tables may be formatted in a spread sheet program such as MS Works or MS Excel. Text should
be double spaced typing, with 25 mm left and right margins. Tables should be put in a separate file. Diskettes should be
accompanied by the name of the software program used (e.g., WordPerfect, Microsoft Word). Authors should keep backup
copies of all material sent to the Editor. The Society cannot be responsible for diskettes or text mislaid or destroyed in transit
or during editing.
Illustrations should be planned for reduction to the dimension of the printed page (15.3 X 22 cm, column width 7.5 cm)
and allow room for legends at the top and bottom. Do not make plates larger than 14" X 18" (35.5 X 46 cm). Individual figures
should be mounted on a suitable drawing board or similar heavy stock. Photographs should be trimmed, grouped together and
abutted when mounted. Figure numbers should be on the plate, but it is strongly recommended that names be included after
the numbers (e.g., Fig. 2, texanus). Include title, author(s) and address(es), and illustration numbers on back of each plate.
Original figures need not be sent until requested by the editor, usually after the manuscript has been accepted. Reference to
figures /tables in the text should be in the style "(Fig. ')" "(Table 1)". Measurements should be in the metric system.
All papers must conform to the International Code of Zoological Nomenclature. The first mention of a plant or animal should
include the full scientific name including the authority. Genus names should not be abbreviated at the beginning of a sentence.
In taxonomic papers type specimens must be clearly designated, type depositories must be clearly indicated, and new taxa must
be clearly differentiated from existing taxa by means of keys or differential diagnoses. Authors are required to deposit all type
material in internationally recognized institutions (not private collections). Voucher specimens should be designated for
specimens used in behavioral or autecological studies, and they should be deposited similarly.
Acceptance of taxonomic papers will not require use of cladistic methods; however, authors using them will be expected
to specify the phylogenetic program used (if any), including discussion of program options used. A data matrix should be
provided. The number of parsimonious cladograms generated should be stated and the reasons for the one adopted. Lengths
and consistency indices should be provided.
References in the text should be (Smith 1999), without a comma, or Smith (1999). Two articles by a single author should
be (Smith 1999a, 1999b) or Smith (1999a, 1999b). For papers in press, use "in press," not the expected publication date. The
Literature Cited section should include all papers referred to in the paper. Journal names should be spelled out completely.
Charges
Publication is free to members of the International Society of Hymenopterists. At least one author of the paper must be a
member. Reprints are charged to the author and must be ordered when returning the proofs; there are no free reprints. Author's
corrections and changes in proof are also charged to the author. Color plates will be billed at full cost to the author.
All manuscripts and correspondence should be sent to:
Paul M. Marsh, Editor
P. O. Box 384
North Newton, Kansas 67117
(316) 284-0990
Bound
.Tb-Wcase
FEB 00
uccTFR INDIAN^"6962!
Live Music Archive
Librivox Free Audio
Metropolitan Museum
Cleveland Museum of Art
Internet Arcade
Console Living Room
Books to Borrow
Open Library
TV News
Understanding 9/11