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Volume19, Number 1 ^^^^ s^^ARIES ^^^,^^ April 2010 

ISSN #1070-9428 

SCPiMIDT, J. O. and C. K. STARR. Editorial. Persona of Roy 1 

OLIVEIRA, O. A. L. de, R B. NOLL, and J. W. WENZEL. Foraging behavior and colony cycle 

of Agelaia vicina (Hymenoptera: Vespidae; Epiponini) 4 

SOSA-CALVO, J., T R. SCHULTZ, and J. S. LAPOLLA. A review of the dacetine ants of Guy- 
ana (Formicidae: Myrmicinae) 12 

MACKAY, W. P. and F. SERNA. Two new species of the strigatus species complex of the ant 

genus Cyphomyrmex (Hymenoptera: Formicidae) from Costa Rica and Panama 44 

GUERRERO, R. J., J. H. C. DELABIE, and A. DEJEAN. Taxonomic contribution to the aurita 

group of the ant genus Azteca (Formicidae: Dolichoderinae) 51 

RICHARDS, M. H. and L. PACBCER. Social behaviours in solitary bees: interactions among in- 
dividuals in Xeralictus bicuspidariae Snelling (Hymenoptera: Hahctidae: Rophitinae). . . 66 

HOOK, A. W., J. D. OSWALD, and J. L. NEFF. Plega hagenella (Neuroptera: Mantispidae) parasi- 
tism of Hylaeus (Hylaeopsis) sp. (Hymenoptera: Colletidae) reusing nests of Trypoxylon 
manni (Hymenoptera: Crabrorudae) in Trinidad 77 

WEST-EBERHARD, M. J., J. M. CARPENTER, L. R R GELIN, and F B. NOLL. Chartergellus 
golfitensis West-Eberhard: a new species of Neotropical swarm-founding wasp (Hy- 
menoptera: Vespidae, PoUstinae) with notes on the taxonomy of Chartergellus zonatiis 
Spinola 84 


BUFFINGTON, M. L. The description of Banacuniculus Buffington, new genus (Hymenoptera: 

Figitidae: Eucoilinae) 94 


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Guest Editors: Justin O. Schmidt and Christopher K. Starr 

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Title of Publication: Journal of Hymenoptera Research. 

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This issue was mailed 16 March 2010 

Vol. 19(1), 2010, pp. 1-3 


Persona of Roy 

Volume 18, Issue 2 (2009) of this Journal was dedicated Roy R. Snelling, with 
an introduction by Longino and Snelling (2009) highlighting his life, some of his 
scientific adventures, and a bibliography of his publications. This issue of the 
Journal of Hymenoptera Research continues the dedication with seven more 
papers honoring Roy. Many of the contributors in both issues penned comments 
of inspirations Roy had given throughout their careers. On the personal side, 
everybody who knew Roy has one or many stories. Here we share a few of the 
stories exemplifying the essence of Roy. 

Who Would Think The Name ''Roy Snelling" Could Save Your Hide! - Almost four 
decades ago I was a graduate student working on my dissertation on the 
chemical ecology of Nearctic Camponotus species. In spring of 1973, 1 embarked 
on a collecting trip across the southern United States. After southern Texas, I 
headed westward collecting in West Texas, New Mexico, around Portal, 
Arizona, and westward to the Huachuca Mountains. Roy Snelling, a friend of 
my advisor, Murray Blum, had given directions to a location where I might be 
able to collect Camponotus ulcerosus Whr. I drove the rather desolate road to the 
Huachuca National Monument and remember thinking that I hoped my old car 
would not break down. When I found what I determined to be the collecting 
locality that Roy had given, I pulled off the road and began looking for foraging 
workers. I was probably a quarter to a half mile south of the dirt road when I 
located the entrance to a colony of C. ulcerosus under a rock and began to 
excavate. I probably dug a two foot deep hole 2-3 feet in diameter. 

Preoccupied with the excavating, I didn't notice the approach of a pick-up 
truck that had driven close to me through the scrub. Two men had gotten out of 
the truck and were heading toward me. One was an older rugged looking 
cowboy and the other looked about twenty. The older one said ''Stand up!" 
They surprised the daylights out of me. As I got up I could see that the younger 
one was wearing a holster with the largest revolver I had ever seen. It looked 
like the barrel was 18 inches long and the guy had his hand on its handle. The 
older guy asked me what I was doing. When I said I was digging up an ant 
colony, he asked "Why here?". I said a friend had told me about this location 
and that there was a particular species of ant that I was hoping to collect. He 
then demanded, "Who told you to come here?". I told him that he would not 
know the person, but I could see he was getting agitated. When he growled "I 
asked you who told you to collect at this location?", I blurted out 'Roy Snelling'. 
Immediately the older cowboy's demeanor changed. He turned to the younger 
guy who still had his hand on the handle of the revolver and said "He's OK, he 
is a friend of Roy's!" They had known Roy for some time and were friends. The 
rancher explained that he had been told that there was someone exhibiting 
suspicious behavior on his property. He further explained that drug dealers 
would cut his fences that parallel the US /Mexican border and drive trucks 
across the border loaded with marijuana. His cattle would inadvertently wander 
over the border into Mexico and were rustled or slaughtered as soon as they 

Journal of Hymenoptera Research: Festschrift Honoring Roy Snelling 

crossed the border. It was costing him hundreds of dollars. The rancher thought 
I might have been a spotter for these people, or that I was digging up drugs 
buried at that location. The name Roy Snelling had a very long reach. 
- Richard M. Duf field, Howard University 

Roy and Big and Halves - In the 1990s I was out many nights alone in Willcox, 
AZ working on vinegaroon behavior. I was slowly walking around with a 
headlight scanning for vinegaroons - that is, acting exactly like a sick or disabled 
prey. To make matters worse, I would crawl under trees or in brush to examine 
critters or holes. Several nights I saw mountain lions in the beam of the 
headlight that was a fixture on my forehead. Mountain lions have a beautiful 
green eyeshine; all other North America cats have yellow eyeshine. One night I 
looked to the right and saw two large green eyes, and a little later looked to the 
left and saw two small green eyes about half the height above the ground. 
Needless to say these night adventures became less pleasant and more anxious. 
It is amazing what one's mind can do when alone for hours at night in a quiet 

Roy Snelling called a few days later and I relayed my story and asked how to 
deal with this situation. He matter of factly commented that the Indians in the 
US West routinely had that problem that they solved by making full neck-length 
chokers of closely fitting elk rib bones. They worked because the cats have rather 
short teeth, cannot puncture through the rib bones, kill by piercing the cervical 
spinal cord, and will flee if the prey is not quickly killed and puts up much of a 
fight. By blocking the success of an initial surprise attack, the warrior could turn 
and punch the cat in the belly or elsewhere and it would flee. I never had to test 
Roy's theory, but did run around thereafter with a thick roll of towel around my 
neck and a bicycle helmet. Roy's wisdom gave me piece of mind and I never 
became cat food! 

- Justin O. Schmidt, Southwestern Biological Institute 

Roy on the Phone - When I was a post doc at the Smithsonian, I had no phone, 
and my rare calls came to Arnold Menke's phone. One day Arnold stuck his 
head out in the hall and yelled gruffly ''Chris, that Indian's on the phone." Well, 
it was kind of like being in England and hearing someone call "God save the 
Queen". I wouldn't ask which queen, and in this case it didn't occur to me to ask 
which Indian. So, I walked down the hall, picked up the phone and asked "You 
send-um smoke signal?", to which Roy responded "Ugh." 

- Christopher K. Starr, University of West Indies 

Horse - This final story was known to many. In 1973 Roy was attending the 
funeral for William S. Creighton, the great North American ant taxonomist of 
the middle of the 20th Century. Creighton had died of a heart attack. While at 
the service Roy found himself in the same hospital as Creighton had been in, in 
the same bed, with the same doctors, and with the same diagnosis. Having seen 
how Creighton ended up and sizing up the competence of the doctors, he 
decided it best to check himself out and return from Missouri to California, 
against the protests of the doctors who said he would never make it alive. Once 

Volume 19, Number 1, 2010 3 

back, he sent the doctors a post card featuring the south end of a horse to assure 
them he had arrived safely. That was Roy! 
- The editors 

This Festschrift would not have been possible without the generous help of 
many people and the support of the International Society of Hymenopterists. 
Many reviewers selflessly interrupted their busy schedules to facilitate rapid 
reviews, often approaching record turn around times. A hearty thanks to the 
followings reviewers: John Alcock, Jeffery R. Aldrich, Craig M. Brabant, Stephen 
Buchmann, James H. Cane, James M. Carpenter, Martin Cooper, Robin Crewe, 
Cameron Currie, Richard M. Duffield, Pierre Escoubas, Fernando Fernandez, 
Brian L. Fisher, Terry Griswold, Darryl T. Gwynne, Robert L. Jeanne, Robert A. 
Johnson, John T. Longino, William P. MacKay, Donald G. Manley, Robert L. 
Minckley, Andreas Mueller, John L. Neff, Michael Ohl, William L. Overal, John 
D. Oswald, Robert J. Paxton, Christian Rabeling, William L. Rubink, Ted R. 
Schultz, Stephen W. Taber, Richard S. Vetter, S. Bradley Vinson, Philip S. Ward, 
John W. Wenzel, Diana E. Wheeler, Alexander L. Wild, Douglas Yanega, and 
James R. Zimmerman. Special thanks to Gavin Broad, editor of the Journal of 
Hymenoptera Research, for generously and expertly shepherding the volume 
through to production. 

Justin O. Schmidt & Christopher K. Starr 

(JOS) Southwestern Biological Institute, Tucson, AZ 85745, USA; email: ponerine® 

(CKS) Department of Life Sciences, University of the West Indies, St Augustine, Trinidad 
& Tabago; email: 


Longino, J. T. and G. C. Snelling. 2009. An inordinate fondness for things that sting. Journal of Hymenoptera 
Research 18: 125-135. 

Vol. 19(1), 2010, pp. 4-11 

Foraging Behavior and Colony Cycle of Agelaia vicina 
(Hymenoptera: Vespidae; Epiponini) 

Otavio a. L. de Oliveira, Fernando B. Noll, and John W. Wenzel 

(OALdeO, FBN) Departamento de Zoologia e Botanica; Institute de Biociencias, Letras e Ciencias 

Exatas, UNESP, Sao Jose do Rio Preto, SP, Brazil 

(JWW) Department of Evolution Ecology and Organismal Biology, Ohio State University, 

Columbus, OH, USA 

Abstract. — The neotropical Agelaia vicina has the largest nest built among social w^asps, yet little is 
known about nest construction, growth and structure. In this work, the development of two nests of 
A. vicina was followed. Studies were done through analysis of images to estimate the growth of nests. 
The material collected below the nests was examined to estimate colony productivity. Nests were 
collected to analyze their architecture and structure. Colony cycle was similar in the two colonies. 
Colonies increase in size throughout the dry season and into the rainy season, with a sudden drop in 
production at the end of the rainy season. The colonies doubled in size in about six months. 

Social w^asps are important to the study 
of social evolution. They fall into tw^o 
groups according to how^ they initiate 
colonies (Jeanne 1991). In independent- 
founding species, one or a small group of 
queens start the construction of a rvew nest, 
w^ithout the aid of w^orkers. In sw^arm- 
founding species, a new^ colony is initiated 
by a large group of w^orkers and one or 
more queens. Patterns in nest construction 
vary. The independent-founders are char- 
acterized by continuous nest construction; 
that is, the size of the nest increases 
gradually throughout the founding and 
ergonomic stages of the colony cycle, and is 
closely correlated w^ith oviposition rate 
(Richards and Richards 1951; Wenzel 

In contrast, some several sv^arm-found- 
ing species engage in episodic nest con- 
struction, building the nest rapidly at the 
beginning of the founding stage. The new^ 
cells are constructed faster than the queens 
fill them with eggs. The nest is completed 
rapidly in this stage (Jeanne 1991; Wenzel 
1991), and then the nest stays static for 
weeks or months, while the colony ex- 
pands its population within the limits of its 

initial structure. Subsequent expansion of 
the nest in its following stage is equally 
quick, building hundreds or thousands of 
cells in a few days. 

In tropical environments, the groups 
most studied for foraging behavior are 
Polistes, Mischocyttarus, and Polybia (Dap- 
porto and Palagi 2006; Hermann and Chao 
1984; Hmcir et al. 2007; Jha et al. 2006; 
O'Donnell and Jeanne 2002; Richter 2000). 
The importance of these wasps is related to 
how they act in a trophic network as 
herbivores (sugar and nectar collectors) 
and predators (Raposo Filho and Rodri- 
gues 1983). Necrophorus feeding habits are 
known in Agelaia and Angiopolybia (O'Don- 
nell 1995). Agelaia is very commonly found 
at flowers (Mechi 2005) and is among the 
most abundant genera in neotropical for- 
ests (Hunt et al. 2001; Silveira et al. 2005; 
Zucchi et al. 1995), indicating its ecological 

Agelaia vicina (Saussure) has the largest 
colony size among the social wasps. Von 
Ihering (1903, 1904) first offered informa- 
tion on this, reporting an A. vicina colony of 
with more than 108,000 individuals, but 
colonies may exceed one million adults 

VOLL-^IE 19, Xl-mber 1, 2010 

(Zucchi et al. 1995). The nests are built 
commonly in cavities, such as caverns or 
tree hollows. Built in protected places, they 
lack a nest envelope, as in many other 
Agelaia (Hunt et al. 2001; Wenzel 1991). The 
nest of A. vicina is composed of vegetable 
fibers without wax or resin. Some parts, 
such as the pedicels receive additional 
glandular secretion as a presumed rein- 
forcement (Wenzel 1998). Workers build 
cells that form combs. During nest initia- 
tion, several combs are built separately, 
fixed by pedicels to the substrate (generally 
ceilings of cavities), so that the combs are 
parallel to the substrate. The combs are 
then expanded and merge to form a great 
expanse of ceUs. Besides the considerable 
amount of information regarding nest 
architecture in A. vicina, \drtually nothing 
is kno^\TL about its biolog}^ 


We obser\^ed t^vo nests in Sao Paulo 
state, Brazil, one in the municipalit}' of 
Paulo de Faria BrazH (19'S 49=W) and the 
other in Pindorama (21 ^S 48 ^W). The Paulo 
de Faria nest was located in an abandoned 
wooden guard station, 3.5 m above the 
ground. The Pindorama nest was 2.5 m 
above the ground, inside a brick structure 
in the form of a shut to^ver in the back yard 
of an abandoned house. 

Images \vere captured using a digital 
camera in order to measure nest gro^vth. 
The images of the Paulo de Faria nest were 
captured from Xovember 2005 to April 
2006, those of the Pindorama nest from 
June 2006 to Februar}^ 2007. Xest growth 
was estimated bv Axiovision, software that 
calculates area increase from the images. 

At the end of the obser\^ations, each nest 
%vas collected, weighed and dismantled 
comb by comb for a better understanding 
of its structure and composition (Fig. lA- 
B). The Paulo de Faria nest was collected 
after natural decline, and the Pindorama 
nest was killed for collection. Subse- 
quently, the combs of each nest were cut 
out in squares with areas of 100cm-. We 

used these squares for counting and 
weighing nest ceUs. 

In February 2007 we gathered 42 hours 
of video segments at the entrance of the 
Pindorama nest in order to record the 
departure and return of foragers. These 
images were studied then in slow motion. 
We recorded for one hour each at starting 
times of 06h, 14h, 18h and 24h. 

To study foraging behavior, we offered 
baits of meat 15m from the nest. Arriving 
foragers \vere marked on the thorax with 
non-toxic ink. This allowed us to estimate 
round- trip times. To determine whether 
the presence of baits increases the number 
of workers leaving the nest, we noted the 
number of marked foragers present 
throughout the day. We designated the 
start of foraging when the first forager 
arrived in the bait. 

It is known that A. vicina discards 
leftover food and the opercula of pupal 
cocoons below the nest (Zucchi et al. 1995). 
Because each operculum corresponds to an 
emergent adult, the number of opercula 
corresponds to the number of adults 
produced in a period of collection. Plastic 
trays were put below the nests in order to 
coUect the discarded material (Fig. IC). 


Xest growth was continuous during the 
period of obserA^ation. From the image 
analyses, the Paulo de Faria nest initially 
had 459,143 cells, which increased to 
approximately 956,340 cells over the course 
of six months of obser\'ations, an increase 
of 108° 0. After nest collection, it ^veighed 
13.8 kg with cells (or combs) organized in 
28 layers (Fig. IB), yielding an estimate of 
69,300 cells per kg. Only the central area 
(about 30° o) of the nest was used for the 
brood production, as seen in the presence 
of meconia in these ceUs. For the nest 
studied in Pindorama it ^vas not possible to 
estimate nest gro^vth. When the nest was 
collected, it had approximately 745,564 
cells distributed in 41 layers and weighed 
11.5kg (Fig. lA). As in the Paulo de Faria 

Journal of Hymenoptera Research: Festschrift Hoxoring Roy Snelling 


Fig. 1. Nests of Agehia vicina from Pindorama (A) and Paulo de Faria (B), SP, Brazil. Position of trays beneath 
a nest to collect material discarded from the Paulo de Faria nest (C). 

nest, a central area comprising about 30% 
of the nest was used for brood production. 
This nest had about 64,832 cells per kg. 

Foraging began early from, the Pindo- 
rama nest, so that during the 06-07h period 
when the colony was in an active phase too 
many workers were active to allow their 
number to be estimated, even with slow- 
motion playback (Table 1). Later, in the 
declining phase of the colony cycle, with 
fewer individuals and less activity, it was 
possible to identify more foragers exiting 
than returning. From 14h-15h to 18h-19h, 
there were more foragers returning. We 
observed that about 30% of the "foraging 
activity" was related to very brief absences 
that presumably brought nothing into the 
nest. This behavior gave rise to a cloud of 
workers outside the nest throughout the 

day while the colony was in an active 

Foraging times of exiting the nest, arriv- 
ing at the bait, then returning to the nest in 
Paulo de Faria averages 2 min (1.19- 
3.47 min). The increase in the number of 
foragers in the baits was linear (Fig. 2), 
suggesting the absence of recruitment 
(Hmcir et al. 2007). 

In October 2005, the colony of Paulo de 
Faria was active, producing males and 
workers, both of which are easily identified 
in this species. Using operculum numbers, 
we found a rise in the production of new 
individuals in the period from November 

2005 to March 2006, reaching a peak in 
March, following by a decrease in April 

2006 (Fig. 3A), with signs of nest desertion 
due to the absence of workers in the 

\"oll"ME 19, Xl"Mber 1, 2010 7 

Table 1. Mean ratio of Agdma vicina foragers exiting by those returning per minute (values reported as exit/ 
return) in different periods of the day at nest in Pindorama, SP, Brazil. These are recorded both when the colony 

was active and ir. apparent good health and later ^vhen it was ir. a state oi decline. 


03 06 





external area of the nest. In front or above 
the nests of A. licina we sometimes saw a 
cloud of foragers fl\TQg near the nest. This 
cloud disappeared after April 2006. 

The colony cycle at Pindorama ^vas 
similar to that from Paulo de Faria. From 
May to June 2006, there was little activit}' 
and almost no production of new individ- 
uals. After July 2006 males ^vere found in 
the population. Adult production in- 
creased to a climax in Januan.' 2007, 
remained high up to the end of Februar}' 
(2007) ^vhen the population lessened dras- 
tically (Fig. 3B). 

Agelaia licuia collects at least 10 different 
orders of insects: Lepidoptera, Coleoptera, 
Dermaptera, H}Tnenoptera, Heteroptera, 
Mantodea, Diptera, Xeuroptera, Blattodae 
and Homoptera. In addition, we found 

manv spiders (.Arachnida: .Araneaej. In the 
two studied colonies, the taxonomic com- 
position of the discarded prey parts '^vas 
similar. Lepidoptera and Coleoptera, rep- 
resented by remains of lar\-al mandibles 
and adult body parts, ^vere the most 
common. Spiders ^vere also an important 
item in their diet. The other orders \\'eie 
found at much lower levels. In addition, 
we found two different seeds, one from a 
grass [Fanicum sp.) and a Cyperus sp. 
Several balls of plant leaf hairs ^vere found, 
^vasp lar\"ae, and some small pebbles and 


Information on architecture of the ob- 

ser\-ed nests corroborates that of Wenzel 
(1991j and Zucchi et al. (1995), in that the 

Mean PIot(Spreadsheet1 6^*6c) 
Mean: Whiter. MeardD.95 Conf. Interval 

18 \ 
jj 16 

S 14 




1 10 

^ 8 




-Q- Mean 

32 "e3rttD.95 Conf. Intewal 

Fig. 2. Xumbers of Agelaia vicina foragers per minute ^mea: 
Pindorama, SP, Brazil. 

SE I that foiind and collected meat baits at 

Journal of Hymenoptera Research: Festschrift Honoring Roy Swelling 



dec/05 jan/06 feb/06 mar/06 apr/06 


Fig. 3. Number of opercula/day collected during the months of study from nests of Agelaia vicina at Paulo de 
Faria (A) and Pindorama (B), SP, Brazil. 

peripheral regions of combs do not fulfill a 
reproductive function but the act as an 
envelope. Detailed studies of nest devel- 
opment in swarm-founding wasps are 
largely limited to the genus Polybia (Jeanne 
and Bouwma 2004; Kudo et al. 2003, 2005; 
Loope and Jeanne 2008), which provides a 
baseline for comparison. In swarms of P. 
occidentalis (Oliver), for example, rapid 
construction in the founding stage com- 
pletes the initial nest in two to three weeks, 
followed by almost no change during 
much of the ergonomic stage. Renewed 
expansion may occur several weeks or 
months later Qeanne and Bouwma 2004). 

In A. vicina, initial construction of the nest 
can also be considered rapid, with the 
construction of several combs. Approxi- 
mately fifteen pedicels and initial cells may 
be built in the first five hours (Oliveira pers. 
obs). However, A. vicina then shows a 
continuous increase in the size of the nest, 
rather than alternating periods of stasis and 
sudden expansion. The growth rate found 
here shows that nests double in size in six 
months. Factors that allow nests of the great 
size of A. vicina include high growth rate, 
high population and large number of queens. 

The large number of foragers producing 
a cloud of workers during the whole 

VoLL-ME 19, Nl-mber I, 2010 

activity period of the colony is visually 
impressive. As we obser\^ed, most exits 
from the nest lead to very short flights that 
appear unconnected with foraging. Hy- 
potheses to explain this persistent cloud 
include that it a) ser\^es a defensive func- 
tion in obstructing the approach of pred- 
ators or parasites, b) ser\^es as a landmark 
in the orientation of returning foragers, or 
c) is simply a way of reducing crowding 
inside the nest during the active part of the 
day. The absence of feces on the substrate 
near the nest indicates that these are not 
defecation flights, as Richards (1978) sug- 
gested. If it is found that the cloud is 
composed of young workers who are not 
yet fully mature, this would be inconsistent 
with h}^othesis (a) and consistent with (b) 
and (c). 

No recruitment to meat baits was de- 
tected. This is in agreement with previous 
obser\^ations on Agelaia Qeanne et al. 1995) 
and contrar}^ to what was obser\^ed for 
Polybia paiilista (Ihering) (Hmcir et al. 2007) 
in which authors found recruitment for 
sugar sources. In terms of diet, our results 
are similar to that from many other social 
wasps (Edwards 1980). A. vicina has a very 
broad diet, foraging for water (O. OHveira 
pers. obs), plant tissues, proteins and 
carbohydrates. They use plant fibers (cel- 
lulose) for nest construction, proteins (from 
captured arthropods and carrion) and 
sugar from fruits (probably from the pulp 
of the seeds found) as energy source for the 
brood and adults (Akre 1982; Rossi and 
Hunt 1988; Spradbery 1973). A. vicina is a 
generaUst predator of land arthropods, 
taking spiders and a broad range of insects. 
We can infer that it plays important roles in 
their ecosystem as a predator of large 
numbers of invertebrates (see daily pro- 
ductivity, below). 

The colony cycle of the two studied 
colonies of A. vicina was similar. In the 
dry season, population increases possibly 
indicated preparations for reproductive 
swarming, which causes an abrupt fall in 
colony size, as is known in other Epiponini 

Qeanne 1991). Based on Figure 3A-B, nests 
differ greatly in the number of individuals 
produced. The Paulo de Faria nest reached 
a production of 18,000 new individuals per 
day, but the Pindorama nest reached no 
more than 300 individuals per day at peak 
production. It is hard to account for this 
very large difference. 

For less complex societies, like Polistes, 
feeding efficiency may not to be a limiting 
factor for nest size (Strassmann and Orgren 
1983). In A. vicina, however, the fall in 
worker production may be a consequence 
of the difficulty of obtaining food in the 
rainy season, associated with the implica- 
tions of a high relative moisture for the 
maintenance of the colony, an increase in 
predation, or appearance of fungus or 
parasitism (Hunt et al. 2001; Richards 
1978). Because the forest remnant of Paulo 
de Faria (435 ha) is larger than that of 
Pindorama (128 ha), it may also provide 
more resources. Nevertheless, the differ- 
ences presented regarding the production 
of new individuals may not be tied only to 
resource availability but also may relate to 
the differences in colony age and preda- 

The term ''keystone species" was coined 
by Paine (1969). A keystone species influ- 
ences several organisms in an ecosystem 
with a effects on the other species out of 
proportion to its abundance. A keystone 
species may determine the types and 
numbers of various other species in a 
communit}^ Several aspects of our results 
suggest that A. vicina may act as a keystone 
species. A. vicina, besides having the 
largest nests and colonies among social 
wasps, has a very high rate of brood 
production (up to several thousand indi- 
viduals per day). The quantitv^ of prey 
brought to the nest is evidently very 
substantial, especially if we take into 
account the rough rule of a 10% efficiency 
energy transfer between trophic levels. A. 
vicina preys upon an impressive diversity 
of arthropods and must impact their 
populations locally. We propose A. vicina 


Journal of Hymenoptera Research: Festschrift Honoring Roy Snelling 

as a candidate keystone species, so that it 
would be fruitful if future studies were to 
evaluate its influence in neotropical envi- 


We dedicate this paper to Roy Snelling, who liked 
social wasps and helped to inspire young hymenop- 
terists of every stripe. He was one of those rare people 
who excelled in doing difficult, dangerous, and 
sometimes unpleasant work if it would open doors 
to a pro\ocative new view of the fascinating insects 
around us. We think he would ha\'e enjoyed returning 
a critique of this paper. This work was funded by 
FAPESP (2005/03569-8; 2008/07633-1). 


Akre, R. D. 1982. Social wasps. Pp. 1-105 in: Hermann, 
H. R. ed. Social Insects, vol. 4. Academic, New 

Dapporto, L. and E. Palagi. 2006. Wasps in the 
shadow: Looking at the pre-hibemating clusters 
of Polistes dominuhis. Annates Zoologici Fennici 43: 

Edwards, R. 1980. Social Wasps: Their Biology and 
Control. Rentokil, East Grinstead. 398 pp. 

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and Defensive Behavior of Mischocyttarus (Mono- 
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Psyche 91: 51-65. 

Hmcir, M., S. Mateus, and F. S. Nascimento. 2007. 
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decisions in swarm-founding wasps. Behavioral 
Ecology and Sociobiology 61: 975-983. 

Hunt, J. H., S. O'Donnell, N. Chemoff, and C. 
Brownie. 2001. Obser\^ations on tw^o neotropical 
swarm-fonding wasps Agelaia yepocapa and A. 
panamaensis (Hymenoptera:Vespidae). Annals of 
the Entomological Society of America 94: 555-562. 

Ihering, R. Von. 1903. Contributions a I'etude des 
Vespides de I'Amerique su Sud. Annates de la 
Societe Entomologique de France 72: 144-155. 

. 1904. As vespas sociaes do Brasil. Revista do 

Museu Paulista 6: 97-309. 

Jeanne, R. L. 1991. The swarm-founding Polistinae. 
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eds. The Social Biology of Wasps. Cornell University 
Press, Ithaca. 

and A. M. Bouwma. 2004. Divergent patterns 

of nest construction in eusocial wasps. Journal of 
the Kansas Entomological Society 77: 429-^i47. 

, J. H. Hunt, and M. G. Keeping. 1995. Foraging 

in social wasps: Agelaia lacks recruitment to food. 
Journal of the Kansas Entomological Society 68: 

Jha, S., R. G. Casey-Ford, J. S. Pedersen, T. G. Piatt, R. 
Cervo, D. C. Queller, and J. E. Strassmann. 2006. 
The queen is not a pacemaker in the small-colony 
wasps Polistes instahilis and P. dominuhis. Animal 
Behaviour 71: 1197-1203. 

Kudo, K., R. Zucchi, and K. Tsuchida. 2003. Initial nest 
development in the swarm-founding paper wasp, 
Polybia paulista (Hymenoptera: Vespidae, Epipo- 
nini): cases of building multiple initial combs. 
Journal of the Nrcv York Entomological Society 111: 

, S. Tsujita, K. Tsuchida, W. Goi, So. Yamane, S. 

Mateus, Y. Ito, S. Miyano, and R. Zucchi. 2005. 
Stable relatedness structure of the large-colony 
swarm-founding wasp Polybia paulista. Beha- 
vioral Ecology and Sociobiology 58: 27-35. 

Loope, K. J. and R. L. Jeanne. 2008. A test of adaptive 
hypotheses for rapid nest construction in a 
swarm-founding wasp. Insectes Sociaux 55: 

Mechi, M. R. 2005. Comunidade de vespas Aculeata 
(Hymenoptera) e suas fontes florals. Pp. 256-265 
in: Pivello, V. R., and E. M. Varanda eds. O Cerrado 
Pe-de-Gigante: Ecologia e Conservagao - Parque 
Estadual Vassununga. Secretaria do Meio Am- 
biente, Sao Paulo. 

O'Donnell, S. 1995. Necrophagy by neotropical 
swarm-founding wasps (Hymenoptera: Vespi- 
dae; Epiponini). Biotropica 17: 133-136. 

and R. L. Jeanne. 2002. The nest as fortress: 

defensive behavior of Polybia emaciata, a mud- 
nesting eusocial wasp. Journal of Insect Science 2: 

Paine, R. T. 1969. A note on trophic complexity and 
community stability. American Naturalist 103: 

Raposo Filho, J. R. and V. M. Rodrigues. 1983. 
Comportamentos trdficos de Mischocyttarus 
(Monocyttarus) extinctus Zikan, 1935 (Polistinae, 
Vespidae). I. Alimentaqao proteica. Naturalia (Sao 
Paulo) 8: 101-104. 

Richards, O. W. and M. J. Richards. 1951. Observa- 
tions on the social wasps of South America 
(Hymenoptera Vespidae). Transactions of the Royal 
Eritomological Society of London 102: 1-170. 

. 1978. The Social Wasps of the Americas, Excluding 

the Vespinae. British Museum (Natural History), 
London. 580 pp. 

Richter, M. R. 2000. Social wasp (Hymenoptera: 
Vespidae) foraging behavior. Annual Review of 
Entomology 45: 121-150. 

Rossi, A. M. and J. H. Hunt. 1988. Honey supplemen- 
tation and its developmental consequences: evi- 
dence for food limitation in a paper wasp, Polistes 
metricus. Ecological Entomology 13: 437-42. 

Silveira, O. T., M. C. Esposito, J. N. Santos Jr, and F. E. 
Gemaquejr 2005. Social wasps and bees captured 
in carrion traps in a rainforest in Brazil. Entomo- 
logical Science 8: 33-39. 

Volume 19, Number 1, 2010 


Spradbery, J. P. 1973. Wasps: an Account of the Biology 

and Natural History of Social and Solitary Wasps. 

University of Washington Press, Seattle. 408 pp. 
Strassmann, J. E. and C. F. Orgren. 1983. Nest 

architecture and brood development times in 

the paper wasp Polistes exclamans (Hymenop- 

terarVespidae). Psyche 90: 237-248. 
Wenzel, J. W. 1991. Evolution of nest architecture. Pp. 

480-519 in: Ross, K. G., and R. W. Matthews eds. 

The Social Biology of Wasps. Cornell University 

Press, Ithaca. 

. 1998. A generic key to the nest of hornets, 

yellowjackets, and paper wasps worldwide (Ves- 
pidae: Vespinae, Polistinae). American Museum 
Novitates 3224: 1-39. 

Zucchi, R., S. F. Sakagami, F. B. Noll, M. R. Mechi, 
S. Mateus, M. V. Baio, and S. N. Shima. 1995. 
Agelaia vicina, a swarm-founding polistine 
with the largest colony size among wasps 
and bees (Hymenoptera: Vespidae). journal 
of the New York Entomological Society 103: 129- 

Vol. 19(1), 2010, pp. 12-43 

A Review of the Dacetine Ants of Guyana (Formicidae: Myrmicinae) 

Jeffrey Sosa-Calvo, Ted R. Schultz, and John S. LaPolla 

QS-C) Maryland Center for Systematic Entomology, Department of Entomology, University of 

Maryland, 4112 Plant Sciences Building, College Park, Maryland 20742, U.S.A. 

(JS-C, TRS) Department of Entomology, National Museum of Natural History, Smithsonian 

Institution, POB 37012, NHB, CE516, MRC 188, Washington, D. C. 20013-7012, U.S.A. 

(JSL) Department of Biological Sciences, Towson University, 8000 York Road, Towson, 

Maryland 21252, U.S.A. 

Abstract. — The dacetine ants of Guyana are reviewed. One genus, Acanthognathus, is added to the 
three genera reported previously from Guyana. A total of 42 species are reported, 32 of v^hich are 
new records for Guyana. Among these 32 species, the following five are new species: Pyramica 
dahlanae sp. n., Pyramica mariae sp. n., Struntigenys acarai sp. n., Strumigenys royi sp. n., and 
Strumigenys waiwai sp. n. Pyramica dahlanae is unusual for the genus because it lacks propodeal 
spines and possesses distinctive mandibular morphology. Pyramica mariae belongs to the gundlachi 
group and is apparently closely related to P. denticulata based on the length of the mandibles, the 
absence of spongiform tissue on the ventral margin of waist segments, general body pilosity, and 
general habitus. Pyramica denticulata is illustrated in order to show morphological differences from 
P. mariae. Strumigenys royi is a remarkable ant because its waist segments lack ventral spongiform 
tissue, it possesses short propodeal spines, its mandibles are long and with a minute denticle 
proximal to the apicodorsal tooth of the mandibular fork, and its coloration is distinctive. 
Strumigenys acarai is unusual because it possesses a minute denticle on the inner margin of the 
mandibles, distinctive rugulose sculpture on the dorsum of mesonotum that differs from sculpture 
found on other parts of the body, and a longitudinal median carina on the promesonotum. 
Strumige7iys waiwai is easily recognized by the unusual multifurcate pilosity of the cephalic dorsum 
and small body size. Modifications of Bolton's (2000) keys to Pyramica and Strumigenys are provided 
to accommodate the newly described species. 

Resumen. — En este articulo se revisaron las hormigas dacetinas presentes en Guyana. El genero 
Acanthognathus se adiciona a los tres generos de hormigas dacetinas conocidos anteriormente para 
Guyana. Se reportan un total de 42 especies, de las cuales 32 constituyen nuevos registros en este 
pais. Dentro de estos 32 nuevos registros, cinco especies son nuevas: Pyramica dahlanae sp. n., 
Pyramica mariae sp. n., Strumigenys acarai sp. n., Strumigenys royi sp. n., y Strumigenys waiwai 
sp. n. Pyramica dahlanae es facilmente reconocida por que carece de espinas en el propodeo y su 
morfologia mandibular es distintiva. Pyramica mariae pertenece al grupo gundlachi y esta 
probablemente relacionada con P. denticulata por la longitud de las mandibulas, la ausencia de 
tejido en forma de esponja en la margen ventral del peciolo y postpeciolo, la pilosidad en el cuerpo 
y por la forma del cuerpo en general. Tambien se ilustra Pyramica denticulata con el fin de mostrar 
las diferencias morfologicas que la separan de P. mariae. Strumigenys royi es una hormiga notable 
caracterizada por la carencia de tejido en forma de esponja en la parte ventral del peciolo y 
postpeciolo, por la presencia de espinas propodeales cortas, por la presencia de mandibulas largas 
y con un diminuto denticulo proximo al diente apicodorsal en la bifurcacion apical, y por la 
coloracion caracteristica. Strumigenys acarai es una hormiga poco usual porque posee un diminuto 
denticulo en el borde intemo proximo a la parte media de las mandibulas, presenta una 
caracteristica escultura rugulosa en el dorso del mesonoto la cual difiere de cualquier escultura 
presente en el resto del cuerpo, y presenta una carina media longitudinal en el promesonotum. 
Strumigenys waiwai es facilmente reconocida por la pilosidad multi-furcada poco usual en el dorso 

Volume 19, Number 1, 2010 


cefalico y por el reducido tamano de las obreras. Modificaciones a las claves de Bolton (2000) para 
identificar las species de Pyramica y de Strumigenys son presentadas para incluir las nuevas 

Key words. — Dacetini, Hymenoptera, leaf-litter sampling, Neotropics, new species, Pyramica, 
Strumigenys, taxonomy 

Species of ants in the tribe Dacetini 
(Formicidae: Myrmicinae) vary greatly in 
size, morphology, and behavior (Holldob- 
ler and Wilson 1990). They inhabit rotten 
wood, leaf litter, soil, and trees (Holldobler 
and Wilson 1990; Bolton 1998) and feed on 
a diverse variety of small arthropods 
(Wilson 1953; Dejean 1985a; Bolton 1998). 
It has been hypothesized that the bizarre 
mandibular morphology of dacetines, in- 
cluding the different mandibular modes of 
action, and the conspicuous spongiform 
tissue located mostly on the waist seg- 
ments are adaptations for attracting and 
capturing springtails (Collembola) on 
which most members of the tribe presum- 
ably feed (Brown and Wilson 1959; Dejean 
1985a, b, 1987; Dietz and Brandao 1993; 
Gronenberg 1996; Kantarovich et al. 2006; 
Masuko 1984, 2009). 

Guyana occupies a central position 
within the Guiana Shield, a large 
(-1,000,000 km^), ancient (Proterozoic, 
—2.5 billion years ago) geological area that 
was once attached to West Africa (Gibbs 
and Baron 1993) and that currently extends 
between the Amazon and the Orinoco River 
Basins. Unlike most tropical countries, 
—70% of Guyana's land, including large 
tracts of primary rainforest, remains intact 
or is only marginally affected by human 
disturbance (Funk and Richardson 2002). 
Due to the creation of new roads, the influx 
of new inhabitants (especially from Brazil), 
and increased mining and timber-harvest- 
ing activity, this situation is rapidly chang- 
ing. It is therefore imperative to gather the 
biological information necessary for identi- 
fying areas of conservation concern. 

The ant fauna of Guyana remains largely 
unknown. Wheeler (1916, 1918) and La- 

Polla et al. (2007) have produced the only 
publications specifically addressing this 
fauna. Weber (1946) studied the fungus- 
growing ants (Attini) from Guyana; Ken\pf 
(1972) and Fernandez and Sendoya (2004), 
based primarily on literature reports, re- 
corded —350 described ant species from 
Guyana. LaPolla et al.'s (2007) study 
recorded 230 ant species (44 genera) 
collected from eight localities using leaf- 
litter mini- Winkler sampling. These figures 
clearly underestimate the actual number of 
species present in the country; for example. 
La Selva, a —1500 ha Biological Reserve in 
Costa Rica, possesses at least 437 ant 
species (Longino et al. 2002). Bolton 
(2000) and Fernandez and Sendoya (2004) 
reported three dacetine genera and 10 
species for Guyana. As a result of recent 
leaf -litter surveys in Guyana (Appendix 1), 
we increase the number of Guyana's 
dacetine ant species to 42, describe two 
new species in the genus Pyramica Roger 
and three new species in the genus 
Strumigenys F. Smith, and report for the 
first time species of Acanthognathus Mayr in 
Guyana (Appendix 2). Although Bolton's 
generic level classification of dacetines has 
recently been questioned (Baroni-Urbani 
and de Andrade 2007), we choose to follow 
it here for the sake of taxonomic stability in 
the face of indecisive phylogenetic data. 

Despite Bolton's (2000) recent mono- 
graph of the dacetines, it is clear that many 
species remain to be discovered and 
described in this species-rich tribe. Fortu- 
nately, Bolton's study provides the context 
for rapidly identifying and describing new 
species as they are discovered (Deyrup 
2006; Sosa-Calvo et al. 2006; Longino 2006; 
Azorsa & Sosa-Calvo 2008, Bolton et al. 


Journal of Hymenoptera Research: Festschrift Honoring Roy Snelling 

2008). This study summarizes the current 
state of dacetine taxonomy in Guyana and 
describes several new species. While 
Guyana certainly contains many more 
dacetine species, both described and un- 
described, we believe it is important to 
begin the process of documenting Guya- 
nese dacetine diversity because (i) this 
information will facilitate the sorting and 
identification of material generated by 
ongoing ant surveys in Guyana, as well 
as in French Guiana, Suriname, and eastern 
Venezuela (Appendix 3); (ii) this informa- 
tion, combined with the information gen- 
erated by those ongoing studies, will 
provide data urgently required by conser- 
vation efforts underway in Guyana and 
Suriname (LaPolla et al. 2007; Sosa-Calvo 
2007; Alonso and Mol 2007; Alonso et al. 
2008); and (iii) this information can be 
incorporated into ongoing studies aimed at 
understanding biodiversity patterns of the 
Guiana Shield, especially those generated 
by the Smithsonian's Biodiversity of the 
Guianas Program (Funk et al. 2002; Funk 
and Richardson 2002). This study increases 
our knowledge of the species that occur in 
Guyana and complements publications on 
other genera including Acropyga Roger 
(LaPolla 2004), Lachnomyrmex Wheeler (Fei- 
tosa and Brandao 2008), Pheidole Westwood 
(LaPolla and Cover 2005), and Rogeria 
Emery (LaPolla and Sosa-Calvo 2006). 


Specimens were examined and measured 
to the nearest 0.001 mm at various magni- 
fications using a Leica MZ125 light stereo- 
microscope. All measurements are in milli- 
meters unless noted otherwise. Specimens 
were photographed using a JVC KY-F70B 
video camera mounted on a Leica M420 
stereomicroscope attached to an IBM In- 
tellistation M Pro computer on which 
composite images were assembled using 
Auto- Montage Pro Version 5.03.0018 BETA 
software® (Synoptics Ltd.). Images were 
cropped and enhanced using Photoshop 
CS2 Version 9® (Adobe Inc.). Scanning 

electron micrographs (SEM) of uncoated 
specimens (P. dahlanae, P. mariae, and S. royi) 
were taken using a Philips XL-30 ESEM 
with Lanthanum Hexaboride (LaB6) source 
and a backscatter detector. Strumigenys 
acarai and S. waiwai were sputter-coated 
with 60:40 wt% Gold:Palladium alloy on a 
Cressington Scientific 108 auto/SE sputter 
coater to a thickness of 25-20 nm. Scanning 
electron micrographs for these specimens 
were taken using an Amray 1810 SEM with 
LaB6 source. Terminology for morphologi- 
cal features and surface sculpture, as well as 
abbreviations, foUow Bolton (1994, 2000) 
and Harris (1979) with modifications where 
noted. Anatomical abbreviations are as 








Eye Length: Maximum dia- 
meter of compound eye in 
lateral view. 

Gaster Length: Length of gaster 
in lateral view from anterior- 
most point of first gastral seg- 
ment (third abdominal seg- 
ment) to posterior-most point, 
excluding sting apparatus if 

Head Length: Length of head 
in full-face (dorsal) view, in- 
cluding occipital lobes and 
anterior clypeal margin but 
excluding mandibles. 
Head Width: Maximum mea- 
surable width of head in full- 
face view, excluding eyes. 
Mandible Length: Exposed 
length of closed mandibles, in 
full-face view, from anterior 
clypeal margin to apex of 

Petiole Length: Straight line 
from posterior-most margin of 
petiole to posterior-most mar- 
gin of metapleural lobe in 
lateral view. 

Postpetiole Length: Maximum 
length of postpetiole in lateral 

Volume 19, Number 1, 2010 






Pronotal Width: Maximum 
measurable width of pronotum 
in dorsal view. 

Scape Length: Maximum 
length of antennal scape, ex- 
cluding condylar bulb. 
Total Length: HL + ML + WL + 
PL + PPL + GL. 
Weber's Length: Maximum 
length of diagonal connecting, 
in lateral view, antero-dorsal 
angle of pronotum to posterior- 
most basal angle of meta- 
pleuron. (=Alitrunk Length in 
Bolton [2000].) 

Cephalic Index: (HW/HL) X 

Mandibular Index: (ML/HL) X 

Petiolar Length Index: (PL/ 
WL) X 100. 
Scape Index: (SL/HW) X 100. 

Specimens examined were borrowed 
from and /or have been deposited in the 
following institutions: The Natural History 
Museum, London, U.K. (BMNH); Centre 
for the Study of Biological Diversity, Uni- 
versity of Guyana, Georgetown, Guyana 
(UGBC); Museum of Comparative Zool- 
ogy, Harvard University, Cambridge, MA., 
U.S.A. (MCZC); Museu de Zoologia da 
Universidade de Sao Paulo, Sao Paulo, 
Brazil (MZSP); National Museum of Nat- 
ural History, Smithsonian Institution, Wash- 
ington, D.C., U.S.A. (USNM). 


Pyr arnica dahlanae Sosa-Calvo, Schultz, 

and LaPolla, n. sp. 

(Figs 1-5) 

Material examined. — Holotype: worker, labeled: 
''GUYANA: Mabura Hill camp at end of Rd. 
from Georgetown to Lethem Rd.; 64 m; 58° 
41.982' W 5° 9.313' N; 29 x 2002; J.S. LaPolla et 
al.; primary forest; litter sample. (JSL021029- 
LS04)" USNM ENT No. 00442119 (UGBC). 
Paratype: worker, labeled: "GUYANA: Calm 

Water Creek along Essequibo River nr. Bartica; 
58° 37.16' W 6° 28.06' N; 23 ix 2002; J.S. LaPolla; 
primary forest; litter sample. GSL020923-LS02)" 
USNM ENT No. 00441577 (USNM). 

Diagnosis (worker). — Very small (TL = 
1.38-1.42); eyes absent; mandibles linear, 
elongate, and in closed position with gap 
between basal mandibular teeth and ante- 
rior portion of clypeus; propodeum un- 
armed; ventral portion of petiole lacking 
spongiform tissue. 

Description (worker). — Head: in full-face 
view, clypeus slightly concave anteriorly, 
with long apical spoon-shaped hairs ex- 
tending over mandibular gap; mandibles 
sublinear and elongate; at full closure 
mandibles contacting only in apical halves 
of their lengths, leaving gap between them 
basally; mandibles with 10 teeth, basal 
tooth acute, all other teeth rounded and 
flattened; teeth 1, 3, 5, 7, 9, and 10 (from 
base to apex) larger than other teeth; lateral 
dorsum of mandible with appressed sim- 
ple hairs; eyes absent; sculpture on clypeal 
plate imbricate; sculpture on cephalic 
dorsum areolate and covered with squa- 
mate hairs; hairs on anterior margin (lead- 
ing edge) of scape spoon-shaped and 
directed basad; antennal scape narrowed 
basally, anterior margin abruptly ex- 
panded, distinctly widest at point of 
expansion; apicoscrobal hair absent. Meso- 
soma: dorsum of anterior portion of prono- 
tum glabrous; pronotal humeral hair ab- 
sent; dorsum of promesonotum and dor- 
sum and declivity of propodeum entirely 
areolate; propodeum lacking spines or 
denticles at its posterior margin; meso- 
pleuron and metapleuron smooth and 
shining; dorsal portion of mesosoma cov- 
ered with appressed spoon-shaped hairs 
(as on head) without erect hairs of any 
kind, lateral portions glabrous. Metasoma: 
petiole lacking ventral spongiform lobe, 
petiolar disc areolate and covered with 
slightly appressed spatulate hairs; lateral 
surface of petiolar peduncle smooth and 
shining; ventral surface of side of petiole 
weakly sculptured; disc of postpetiole 


Journal of Hymexoptera Research: Festschrift Honoring Roy Snelling 

weakly sculptured and shining, covered 
with hairs similar to those on petiole but 
narrower; ventral surface of postpetiole 
with well-developed spongiform lobe that 
extends throughout its entire length; lateral 
spongiform tissue overhanging ventral 
spongiform lobe; dorsal surface of first 
gastral segment smooth with some lon- 
gitudinal basigastral costulae. Color: indi- 
viduals light yellow to dark yellow. Hairs 
throughout body lighter than integument. 

Measurements: holotype (and paratype): 
GL = 0.3 (0.32), HL = 0.34, HW = 0.27 (0.28), 
ML = 0.09, PL = 0.17, PPL = 0.12 (0.11), PW 
- 0.19 (0.18), SL = 0.16, TL = 1.42 (1.38), WL 
= 0.39 (0.36). Indexes: CI = 82 (79), MI = 26, 
PI = 47 (44), SI = 59 (57). (n = 2) 

Gyne and male. — Unknown. 

Etymology. — Named after Ms. Nor Far- 
idah Dahlan in recognition of her expertise 
and hard work in support of Smithsonian 
ant research and in gratitude for her 
consistent good will and friendship. JS-C 
is deeply grateful to Faridah for all her help 
and care when he first arrived in the 
United States. 

Comments. — Pyramica dahlanae n. sp. is 
most similar to members of the Nearctic 
perga7idei-gT0up, which includes P. angulata 
(M.R. Smith), known from the southeastern 
United States and Illinois, and P. pergandei 
(Emer\'), widely distributed in Canada and 
the United States. Pyramica dahlanae shares 
with those species the following characters: 
(i) mandibles short (MI 25-35) and, in 
frontal view, narrow and elongate, dentate 
only at the apical portion where they are in 
contact leaving an edentate gap betv\^een 
them; (ii) specialized mandibular dentition 
(alternating pattern of longer and shorter 
mandibular teeth); (iii) lateral chqpeal mar- 
gins, in dorsal view, extending beyond the 
line of the outer margin of the mandibles 
when closed; and (iv) preocular carina 
broad and conspicuous. Pyramica dahlanae 
differs from the species in the pergandei- 
group in four character states: (i) 10 
mandibular teeth (15-16 in the pergandei- 
group), (ii) absence of triangular teeth on 

the propodeum (present in the pergandei- 
group), (iii) absence of a well-developed 
spongiform tissue on the ventral portion of 
the petiole (present in the pergandei-group), 
and (iv) shorter antennal scape, SI 57-59 (SI 
65-84 in the pergandei-group). 

The mandibles of P. dahlanae are similar 
to those within the pergandei-group in that 
they contact in the apical third, producing 
a basal gap betw^een the mandibles. This 
condition is different from the one found in 
species in the ohioensis-group, in which the 
masticatory margins contact through al- 
most their entire lengths and in which the 
mandibles are triangular rather than elon- 
gate. Elongate mandibles can be found in 
the gundlachi- and argiola-giowps, the latter 
an Old World group introduced into the 
United States (P. hexamera (Brown)). Man- 
dibles in P. hexamera are highly distinctive 
with an elongate and spiniform apicodor- 
sal tooth and two long preapical teeth (see 
Bolton 2000 for further information). Spe- 
cies of the gundlachi-group share with P. 
dahlanae the absence of a spongiform lobe 
on the ventral surface of the petiole but 
differ from P. dahlanae in: (i) mandibular 
length and morphology, (ii) the presence of 
a pair of triangular teeth or short spines on 
the propodeum, and (iii) the presence of 
pronotal humeral hairs and, in almost all 
species, a pair of laterally projecting apico- 
scrobal hairs. 

Pyramica dahlanae may also be related to 
P. paradoxa Bolton, known from a single 
worker collected in Costa Rica. Both 
species share the absence of propodeal 
spines; however, P. dahlanae differs from P. 
paradoxa by the shape of the mandibles, 
and the head and mesosoma strongly 
areolate with the meso- and metapleuron 
smooth and shining. The head and meso- 
soma are mostly smooth and shining in P. 
paradoxa. Although P. dahlanae shares a 
number of character states with some 
members of the aforementioned groups, 
this species is not easily placed in any of 
the species groups defined by Bolton 

VOLL-ME 19, Nl-mber 1, 2010 


Figs 1-5. Scanning electron micrographs of the holotv^e worker of Pyramica dahlanae, new species. 1, Full-face 
(dorsal) view. 2, Closed mandibles. 3, Dorsal view of petiole and postpetiole. 4^ Dorsal view of mesosoma. 5, 
Lateral view. 


Journal of Hymenoptera Research: Festschrift Honoring Roy Snelling 


Pyramica dahlanae will not key out to any known species of Pyramica in either the Nearctic or the 
Neotropical keys of Bolton (2000). The key to Neotropical species can be modified as below to include 
P. dahlanae. The numbering of couplets follows Bolton (2000). 

7. Dorsum of postpetiole (= disc) smooth and with weak costulae 7b 

- Dorsum of postpetiole entirely reticulate-punctate couplet 12 in Bolton (2000) 

7b. Cephalic dorsum with 1 or 2 pairs of standing hairs. Apicoscrobal and pronotal 

humeral hairs present couplet 8 in Bolton (2000) 

Cephalic dorsum lacking standing hairs. Apicoscrobal and pronotal humeral hairs 

absent P. dahlanae new species 

Pyramica mariae Sosa-Calvo, Schultz, and 

LaPolla, n. sp. 

(Figs 6, 8, and 10) 

Material examined. — Holotype: worker, labeled 
"GUYANA: Mt. Ayanganna montane forest; 
1300 m; 59° 57.969' W 5° 22.483' N; 13.X.2002; 
T.R. Schultz, }. LaPolla, C. Marshall, R. Williams; 
litter sample." USNM ENT No. 00413858. 
(UGBC). Paratypes: 3 workers, same locality as 
in holotype. USNM ENT No. 00413859, 
00442882, 00442883. (USNM). 

Diagnosis (worker). — Mandibles linear, 
elongate, and narrow; inner margin of 
mandibles with two clearly defined teeth, 
which are larger than the rest; labral lobes 
short with long trigger hairs at their apices; 
metapleuron smooth and shining; ventral 
portions of petiole and postpetiole lacking 
spongiform tissue. 

Description (worker). — Possessing char- 
acters of the gundlachi-group and gundlachi- 
complex (Bolton 2000). Head: in full-face 
view nearly as broad as long; inner margin 
of elongate mandibles slightly concave to 
more or less straight, with 4 teeth on left 
mandible and 3 on right mandible, of 
which a pair of teeth are larger on each 
mandible (same in paratypes); with 2 mi- 
nute intercalary denticles between apico- 
dorsal and apicoventral fork teeth; labral 
lobes short, almost invisible in full-face 
view; trigger hairs long; eyes with 3 

ommatidia in longest row, with 6-7 om- 
matidia in total. Cephalic dorsum with two 
pairs of erect hairs: one pair located close 
to occipital margin and another pair 
located close to highest point of vertex; 
each upper scrobal margin with a short 
apicoscrobal hair that projects laterally. 
Mesosoma: pronotum with a pair of short 
humeral hairs that project laterally; meso- 
notum with a pair of short, erect, stiff hairs; 
mesopleuron and metapleuron mostly 
smooth and shining; dorsum of promeso- 
notum, propodeum, and propodeal decliv- 
ity strongly reticulate. Metasoma: peduncle 
of petiole long, length of petiole 3-3.5 times 
longer than its disc; petiolar disc reticulate- 
punctate, with a pair of erect hairs on 
posterior portion of disc; ventral portion of 
petiole lacking spongiform tissue; disc of 
postpetiole reticulate, ventral portion of 
postpetiole lacking spongiform tissue; pos- 
terior portion of postpetiole disc with a 
row of 4 erect hairs; first gastral tergite 
almost entirely reticulate except for a small 
portion at posterior portion of tergite. 
Individuals light brown to brown. 

Measurements: holotype (and paratype): 
GL = 0.59 (0.48), HL = 0.52 (0.48-0.50), 
HW = 0.42 (0.38-0.46), ML = 0.36 (0.36- 
0.38), PL = 0.28 (0.24-0.27), PPL - 0.12, 
PW = 0.27 (0.23-0.24), SL = 0.30 (0.30- 
0.31), TL = 2.47 (2.24-2.28), WL = 0.58 
(0.55-0.56). Indexes: CI = 81 (78-92), MI = 

Volume 19, Number 1, 2010 


Figs 6-11. Full-face (dorsal) and lateral views of the holotype worker of Pyramica mariae, new species (6, 8, 10) 
and P. denticulata (7, 9, 11). 

73 (72-75), PI = 48 (43^9), SI = 71 (65-82). 
(n = 4) 

Gyne and male. — Unknown 

Etymology. — Named in honor of the first 
author's mother, Maria del Carmen Calvo, 
in gratitude for her encouragement and 

Comments. — Pyramica mariae n. sp. is 
clearly a member of the gundlachi-gioup 

(refer to Bolton [2000: 176-179 p.] for 
further information). Within the gundlachi- 
group, Bolton (2000) identified two com- 
plexes, crassicornis and gundlachi. Pyramica 
mariae belongs to the gundlachi complex 
and resembles P. denticulata (Mayr), P. 
enopla Bolton, and P. vartana Bolton. Pyr- 
amica mariae shares with P. vartana the 
smooth and shining mesopleuron and 


Journal of Hymenoptera JIesearch: Festschrift Honoring Roy Snelling 

metapleuron, but P. mariae can be distin- 
guished from P. vartana by the form of the 
apicoscrobal and pronotal humeral hairs, 
both short and stiff {mariae) rather than 
long and filiform (vartana), and the disc of 
the postpetiole is reticulate {mariae) rather 
than smooth and shining {vartana). 

Pyr arnica mariae is of similar size and 
color as P. enopla. However, P. mariae 
differs from P. enopla in that the apicoscro- 
bal, humeral, and mesonotal hairs are 
short, erect, and stiff {mariae) rather than 
long and filiform {enopla); the metapleuron 
is smooth and shining {mariae) rather than 
reticulate {enopla); the dorsum of the 
petiole bears a single pair of hairs {mariae) 
rather than tw^o pairs of hairs {enopla); and 
the dorsum of the postpetiole lacks an 
anterior pair of hairs {mariae), present in 

Pyramica mariae can easily be confused 
with P. denticulata (Figs 7, 9, and 11) with 
which it shares the most character states. 
However, the species can be separated by: 
(i) mandibular dentition: P. denticulata has 
5-10 preapical denticles of similar size, 
whereas P. mariae has 3-4 preapical denti- 

cles, two of which are larger than the rest. 
In Pyramica mariae, at least in the four 
specimens examined, there are 4 teeth on 
the left mandible and 3 teeth on the right 
mandible; (ii) mesosomal sculpture: the 
metapleuron in P. denticulata is reticulate, 
whereas in P. mariae it is smooth and 
shining; (iii) petiole proportions: the petio- 
lar peduncle in P. denticulata is relatively 
shorter (PI 38-42) than in P. mariae (PI 43- 
49) (Figs 12-13). 

The four specimens known of P. mariae 
were collected in a leaf-litter sample 
extracted with a mini-Winkler. The sample 
was collected in a primary lower montane 
forest (1300 m). Other species in the 
gundlachi-group have been recorded from 
wet forest habitats and from lowland rain- 
forest to cloud forest and some in agro- 
ecosystems. Pyramica denticulata, the spe- 
cies perhaps most closely related to P. 
mariae, has been collected in lowland (< 
1000 m) forests in Panama (Sosa-Calvo et 
al. 2006) to subtropical forests in the wet 
Chaco region of Argentina (Theunis et al. 
2005). Nothing is known about the biology 
of P. mariae other than the collection data. 


In Bolton's (2000) key, Pyramica mariae keys out to P. denticulata. The key can be modified as 
below to include P. mariae. Numbering of couplets follows Bolton (2000). 



In lateral view, postpetiole lacking ventral spongiform lobe; sometimes a minute 

vestige visible; mesonotum with a pair of erect hairs 23b 

In lateral view, postpetiole with reduced ventral spongiform lobe but distinct; if lobe 
very shallow then mesonotum with pair of straggly (i.e., laid out in an irregular, 

untidy way) flagellate hairs couplet 25 in Bolton (2000) 

Mandibles long, MI 72-85. Dorsum of pronotum lacking pair of stiff erect hairs . . . 23c 
Mandibles short, MI 58-65. Dorsum of pronotum with pair of stiff erect hairs . . . eggersi 
Inner margin of mandibles with 5-10 preapical denticles of similar size. Metapleuron 

densely reticulate. Peduncle of petiole short, PI 38^2 denticulata 

Inner margin of mandibles with 3-4 preapical denticles, two distinctly larger than rest. 

Metapleuron smooth and shining. Peduncle of petiole elongate, PI 48-49 

mariae new species 

Volume 19, Number 1, 2010 


0.48 1 




1 ^K 

A A 


^ 0.4. 






• A 






■ Pyramica ikniiculata 







• Pyramica marlae 
A Pyramica enopla 


0.48 0.53 

Head Length 

0.77 - 





C 0.62 ■ 







0.52 ■ 











■ Pyramica denficulata 
• Pyramica mariae 
A Pyramica enopla 


0.15 0.17 0.19 0.21 0.23 25 0.27 0.29 0.3! 0.33 

Petiole Length 

Fig. 12. Relationship between head width and head length among Pyramica denticulata, P. mariae, and P. enopla. 
Measurements in millimeters. 

Fig. 13. Relationship between petiole length and Weber's length among Pyramica denticulata, P. mariae, and P. 
enopla. Measurements in millimeters. 

USNM ENT No. 00537288. (UGBC). Paratype: 1 
worker, same locality as in holotype. USNM 
ENT No. 00537289. (USNM). 

Diagnosis (worker). — Leading edge of 
Material examined.— Holotype: worker, labeled antennal scape with all hairs curving to 

Strumigenys royi Sosa-Calvo, Schultz, and 

LaPolla, n. sp. 

(Figs 14-25) 

''GUYANA: Kanuku Mts.: Nappi Creek, camp; 
128 m; 59°33.963' W, 3°21.018' N; 24.X.2002; ].S. 
LaPolla; forest; on tree trimk. (JSL02 1024-08)" 

apex, lacking hairs that curve to the base of 
segment; mandibles long and linear with a 
small, but conspicuous preapical tooth 


Journal of Hymenoftera Research: Festschrift Honoring Roy Snelling 

Figs 14-22. Scanning electron micrographs of the holotype worker of Strumigenys royi, new species. 14, Lateral 
view. 15, Lateral view of head. 16, Full-face view. 17, Closed mandibles. 18, Lateral view of mesosoma. 19, 
Lateral view of petiole, postpetiole, and gaster. 20, Dorsal view of mesosoma. 21, Dorsal view of mesosoma, 
waist segments, and first gastral tergite. 22, Dorsal view of waist segments (petiole and postpetiole). 

close to the apicodorsal teeth; propodeum 
with small denticles; segments of the waist 
with ventral margin lacking spongiform 
tissue of any kind; first gastral stemite 
lacking spongiform pad; body strongly 
reticulate and with area within each re- 
ticulation verrucose; coloration distinctive: 
mandibles mostly whitish, antennae and 
legs yellowish, mesosoma mostly ferrugi- 

nous, waist segments light brown, and 
head and gaster mostly dark brown or 

Description (worker). — Head: in full-face 
view, mandibles thick throughout most of 
their length and abruptly narrowing just 
before apex by sudden oblique divergence 
of inner margin, a minute but conspicuous 
preapical denticle arising on this oblique 

Volume 19, Number 1, 2010 


0.5 mm 

Figs 23-25. Automontage images of a paratype of Strumigenys royi, new species. 23, Full-face view. 24, Dorsal 
view. 25, Lateral view. 

section; mandibles with intercalary denti- 
cle that arises from dorsal base of apico- 
ventral tooth; in full-face view, anterior 
margin of clypeus transverse to very 
slightly concave and with at least 6 
narrowly spatulate elongate hairs; dorsum 
of clypeus finely reticulate-punctate and 
with short, appressed, simple hairs; ocular 
carina short, ending at eye level; leading 
edge of scapes with all hairs curved or 
inclined toward apex of scape; funicular 

segments 11 and III long [holotype: 2'''^ = 
0.073, 3'" = 0.092; paratype: 2"^^^ = 0.079, 3"^^ 
= 0.092], their lengths, when combined, 
almost as long as funicular segment IV; 
occipital margin deeply emarginate, form- 
ing prominent rounded cephalic lobes; 
dorsum of head with two pairs of erect 
fine hairs: one pair close to margin of 
occipital concavity and another close to 
highest point of vertex, clearly differing 
from appressed simple ground-pilosity 


Journal of Hymenoptera Research: Festschrift Honoring Roy Snelling 

(sensii Bolton 2000:998: referring to "the 
short pilosity often present on cephalic 
dorsum, dorsolateral margins of head, 
promesonotum and its margins. These 
hairs could be simple or spatulate to 
orbicular, usually decumbent to appressed 
and may rarely be elevated"); eye with 13- 
14 ommatidia on longest row; apicoscrobal 
hair elongate and simple (this hair lacking 
on holotype due to damage); dorsum of 
head strongly reticulate and with areas 
within each reticulation verrucose (i.e., 
containing wart-like protuberances). Meso- 
soma: humeral hair elongate and simple, 
similar in shape to apicoscrobal hair but 
longer; pilosity on dorsum of pronotum 
consisting of decumbent hairs; dorsum of 
pronotum with irregular rugae and mark- 
edly reticulate with areas within reticula- 
tion verrucose; mesonotum, in lateral view, 
raised and separated from pronotum by 
transverse or rounded carina; mesonotum, 
in lateral view, with pair of erect elongate 
simple hairs and pair of short erect simple 
hairs; dorsum of mesonotum with conspic- 
uous longitudinal rugae and strongly 
reticulate with area within reticulation 
verrucose; mesopleuron and metapleuron 
separated by deep scrobiculate constriction 
that extends to dorsum of alitrunk to form 
metanotal groove; in lateral view, constric- 
tion extends backward from propodeum 
throughout base of propodeal denticles, 
before beginning of propodeal declivity; 
propodeum with small denticles; declivity 
of propodeum with thin reticulate carina; 
mesopleuron mostly reticulate and with 
area within reticulation verrucose; proximal 
to border with metapleuron (and especially 
upper portion of katepistemum), reticula- 
tions fading leaving only verrucose sculp- 
ture visible; anepisternum mostly verru- 
cose; metapleuron reticulate and with areas 
within reticulation verrucose. Metasoma: 
waist segments lacking ventral spongiform 
tissue; petiole with anterior acute process; 
node of petiole, in lateral view, rounded; 
pilosity on petiole, in frontodorsal view, 
consisting of anterior pair of elongate. 

simple suberect hairs and posterior trans- 
verse row of four elongate, simple suberect 
hairs (hair on each side of petiole and pair 
on posterior dorsum of petiole); posterior 
margin of petiolar node with low spongi- 
form crest; disc of petiole, in dorsal view, 
rugose-reticulate and with area within 
reticulation verrucose. Postpetiole, in lateral 
view, globose and in fronto-dorsal view, 
with two transverse rows of four elongate, 
simple suberect hairs, located on anterior 
and posterior portions of postpetiole. (Dis- 
tribution of these hairs similar to that of 
posterior row on petiole.) Anterior margin 
of postpetiole, in dorsal view, concave; 
postpetiole wider than long [length = 
0.205, width = 0.238]; posterior margin of 
postpetiolar disc with small spongiform 
crest; disc of postpetiole reticulate with 
areas within reticulations verrucose. First 
gastral tergite finely reticulate-substrigulate 
with some verrucose sculpture confined to 
basigastral area; dorsum of first gastral 
tergite with widely spaced elongate erect 
simple hairs. Similar hairs on gastral ster- 
nite but more abundant; first gastral stemite 
reticulate, differing from sculpture on ter- 
gite; basigastral costulae longitudinal, 
spaced, and very short but conspicuous. 
Color: anterior portion of head yellowish 
and gradually increasing in color to ferru- 
ginous by level of eyes and dark brown on 
rest of head. Mandibles mostly whitish with 
tips ferruginous to dark brown. Mesosoma 
ferrugineous; petiole and postpetiole light 
brown; legs and antennae yellowish, 
slightly lighter in color than waist segments; 
first gastral tergite black or dark brown, 
second and third gastral tergites ferrugi- 
neous, fourth gastral tergite yellowish; first 
to third gastral sternites ferrugineous, 
fourth gastral sternite yellowish. 

Measurements: holotype (and paratype): 
EL = 0.16 (0.14), GL = 0.79 (0.78), HL = 
0.86, HW = 0.65, ML = 0.49 (0.52), PL = 
0.40 (0.37), PPL = 0.19 (0.20), PW = 0.36 
(0.35), SL = 0.57 (0.59), TL = 3.53, WL = 
0.79 (0.80). Indexes: CI = 75, MI = 57 (60), 
PI = 51 (46), SI = 88 (90). (n = 2) 

Volume 19, Number 1, 2010 


Gyne and male. — Unknown. 

Etymology. — This species is named in 
honor of Roy Snelling to acknowledge his 
numerous contributions to the taxonomy 
of ants, bees, and wasps. He will live on 
through the solid foundation he provided 
for ant taxonomy and through the thou- 
sands of specimens that he left behind for 
myrmecologists to ponder over for many 
years to come. 

Biology. — Strumigenys royi was col- 
lected from an upright, living tree trunk 
in a small dirt tunnel (likely made by 
termites) that ran up the side of the 

Comments. — This large species is easily 
distinguished from any other species in the 
genus Strumigenys (sensu Bolton 2000) by 
lacking the spongiform tissue on the 
ventral margin of the waist segments 
(petiole and postpetiole) and lacking a 
spongiform pad on the first gastral stemite. 

by having the apical fork of the mandibles 
with an intercalary denticle that arises 
from the dorsal base of apicoventral tooth, 
by having antennal funicular segments II 
and m, when combined, almost as long as 
funicular segment IV (shared with S. fair- 
childi Brown), by having a minute denticle 
close to the apicodorsal tooth (similar in S. 
lanuginosa Wheeler), and by having 
marked body sculpture. Due to this com- 
bination of characters it is difficult to place 
this species in any of the species groups 
given by Bolton (2000). 

Strumigenys royi differs from S. idiogenes 
Bolton, to which it keys out in Bolton's 
(2000) key, as the latter possesses: a 
larger and conspicuous lobe on ventral 
margin of postpetiole, a narrow spongi- 
form pad on the base of first gastral 
sternite, asymmetrical dentition on the 
mandibles, and a pair of narrow spines 
on the propodeum. 


Strumigenys royi will key out to S. idiogmes ia Bolton's (2000) "key to Neotropical-Nearctic 
Strumigenys species." The key for the species of Strumigenys can be modified as below to include S. 
royi. Numbering of couplets follows Bolton (2000). 

4. Mandibles without intercalary teeth or denticles that arise between apicodorsal and 
apicoventral teeth of apical fork, nor arise from dorsal base of apicoventral 
tooth couplet 5 of Bolton (2000) 

- Mandible with 1 or 2 intercalary teeth or denticles that arise between apicodorsal and 

apicoventral teeth of apical fork, or arise from dorsal base of apicoventral 

tooth couplet 10 of Bolton (2000) 

Mandibles without, or with only one, preapical tooth or denticle couplet 11 of Bolton 


Mandible with 2 preapical teeth or denticles couplet 15 of Bolton (2000) 

Preapical dentition consisting of single tooth on one or both mandibles; preapical tooth 

conspicuously dentiform and located close to apicodorsal tooth 12 

Preapical dentition absent from both mandibles or single minute denticle present; if 

the latter denticle located close to midlength, not near apicodorsal tooth 14 

First gastral tergite very densely clothed with long fine flagellate hairs. Dorsolateral 

margin of head with 2 freely laterally projecting long flagellate hairs, one at level of 

eye, other apicoscrobal lanuginosa 

- First gastral tergite with stout curved hairs that are rermform or apically spatulate or 

simple erect standing hairs. Dorsolateral margin of head without projecting 

flagellate hairs or with single hair, in apicoscrobal position 13 

13. Scape strongly dorsoventrally flattened and very broad; in full-face view maximum 

width of scape greater than maximum width of mandible. First gastral tergite 





Journal of Hymenoptera Research: Festschrift Honoring Roy Snelling 

unsculptured. Pronotal humeral hairs stiff and stout. With head in profile highest 
point of vertex without erect hairs platyscapa 

Scape subcylindrical; in full-face view maximum width of scape less than maximum 
width of mandible. First gastral tergite with sculpture present other than 
basigastral costulae. Pronotal humeral hairs elongate or flagellate. With head in 

profile highest point of vertex with pair of erect hairs 13b 

13b. Ventral surface of postpetiole with large and distinct spongiform lobe. Propodeal 
spines long. Mandibles with asymmetrical preapical dentition (left mandible 
without trace of preapical dentition, right mandible with small slender preapical 
tooth located close to apicodorsal tooth and slightly smaller than intercalary 
tooth) idiogenes 

Ventral surface of postpetiole without spongiform lobe or crest. Propodeal spines 
short. Mandibles with symmetrical preapical dentition (both mandibles with small 
preapical tooth located close to apocidorsal tooth) royi new species 

Strumigenys acarai Sosa-Calvo, Schultz, 

and LaPolla, n. sp. 

(Figs 26-39) 

Material examined. — Holotype: worker, la- 
beled "GUYANA: Upper Takutu-Upper Esse- 
quibo, Acarai Mountains, New Romeo Camp, 
1069 m., 58°57.828'W, ri9.938'N; 14.X.2006; 
T.R. Schultz, J. Sosa-Calvo, C.J. Marshall, R. 
Williams; 1° upland forest; leaf-litter sample. 
(JSC061014-01)" USNM ENT No. 00537294. 
(UGBC). Paratypes: 9 workers, labeled 
"GUYANA: Upper Takutu-Upper Essequibo, 
Acarai Mountains, New Romeo Camp, 1069 m., 
58^57.828'W, ri9.938'N; 14.X.2006; T.R. Schultz, 
J. Sosa-Calvo, C.J. Marshall, R. Williams; 1° 
upland forest; leaf-litter sample. (JSC06 101 4-02, 
JSC061014-03). USNM ENT No. 00537295- 
00537303. (BMNH (1), MCZC (1), and USNM 

Diagnosis (worker). — Small (TL 1.62- 
1.79); eyes vestigial, consisting of one or 
two ommatidia; masticatory margin of 
mandibles with an inconspicuous tooth, 
visible under high magnifications; leading 
edge of antennal scapes with some hairs 
that curve toward the base of scape; 
dorsum of promesonotum rugulose and 
with a conspicuous median longitudinal 
ruga that extends for entire length of 
promesonotum; petiole lacking a ventral 
process or spongiform tissue of any kind. 

Description (worker). — Head: mandibles 
elongate with outer margin convex; inner 

margin of mandibles with minute incon- 
spicuous preapical denticle in mandible's 
midlength, visible under high magnifica- 
tion; apical fork of mandibles lacking 
intercalary teeth; anterior margin of cly- 
peus slightly concave or transverse; dor- 
sum of antennal scape imbricate; anterior 
edge of antennal scape with at least 3 
narrowly spatulate hairs curving toward 
base, some hairs on scape multi-furcate 
(Fig. 30); hairs on upper margin of scrobe 
narrowly spatulate and curving anteriorly; 
apicoscrobal hair flagellate; dorsum of 
head strongly areolate; ocular carina failing 
to reach level of eyes; eyes minute, with 
only 1 (one paratype, most of them with 
two) or 2 ommatidia; dorsum of head with 
fine subdecumbent hairs, some of which 
curve medially and with pair of erect hairs 
present on cephalic margin (very difficult 
to see). Mesosoma: humeral hair flagellate; 
anterior portion of pronotum, in dorsal 
view, strongly reticulate; dorsum of pro- 
mesonotum rugulose and with conspicu- 
ous median longitudinal ruga or carina 
that extends for entire length of promeso- 
notum; areas between rugae smooth and 
shining; dorsum of promesonotum with 
subdecumbent hairs that curve medially, 
most hairs directed backwards; posterior 
half of promesonotum areolate; mesono- 
tum with pair of flagellate simple hairs; 
dorsum of propodeum and declivity of 

Volume 19, Number 1, 2010 


propodeum areolate; mesopleuron and 
metapleuron mostly smooth and shining; 
mesopleuron and metapleuron divided by 
strip of aerolate sculpture that originates at 
ventral margin of mesopleuron and meta- 
pleuron and extends dorsally in direction 
of metanotal groove, this strip incomplete, 
fading before it connects with metanotal 
groove; propodeal spines long; declivity of 
propodeum with a thin carina. Metasoma: 
dorsum and sides of petiole strongly 
areolate; ventral margin lacking spongi- 
form tissue or process of any kind; node of 
petiole, in lateral view, with two transverse 
rows each consisting of four long subde- 
cumbent and simple hairs and composed 
of two hairs medially and two hairs 
distally (Fig. 35); posterior margin of pet- 
iolar node with small spongiform crest, 
best seen in fronto-dorsal view; in dorsal 
view, lateral projections of crest conspic- 
uous and triangular; postpetiole with ven- 
tral and lateral spongiform lobes well 
developed; dorsum of postpetiole with 
longitudinal rugae, areas between rugae 
smooth and shining; base of first gastral 
stemite bearing conspicuous pad of spon- 
giform tissue; basigastral costulae lon- 
gitudinal and sharply defined, longer than 
maximum length of disc of postpetiole; 
dorsum of first gastral tergite with numer- 
ous long flagellate hairs; entire tergite 
posterior to basigastral costulae smooth 
and shining. 

Measurements: holotype (and paratypes): 
GL = 0.40 (0.35-0.41), HL = 0.42 (0.39- 
0.41), HW = 0.31 (0.29-0.33), ML = 0.25 
(0.24-0.25), PL = 0.20 (0.15-0.19), PPL = 
0.09 (0.08-0.11), PW = 0.18 (0.17-0.19), SL 
- 0.29 (0.27-0.30), TL = 1.77 (1.62-1.79), 
WL = 0.41 (0.38-0.42). Indexes: CI = 73 
(74-81), MI = 59 (58-64), PI = 48 (38-51), SI 
= 94 (88-96). (n = 10) 

Gyne and male. — ^Unknown. 

Etymology. — The name of this species 
refers to the Acarai Mountains, in the 
Upper Takutu-Upper Essequibo region of 
southern Guyana, where specimens of this 
species were collected. 

Comments. — Strumigenys acarai seems to 
belong to the S. silvestrii species group 
(sensu Bolton 2000), sharing with some 
members of that group: (i) the ventral 
margin of petiole lacking spongiform tis- 
sue; (ii) the small worker size (HL 0.39- 
0.43, HW 0.29-0.33, TL 1.62-1.79, WL 0.38- 
0.42 in S. acarai, HL 0.36-0.52, HW 0.28- 
0.44, TL 1.5-2.2, WL 0.36-0.56 in the S. 
silvestrii group); (iii) the apical fork of 
mandibles lacking intercalary denticles; 
(iv) the leading edge of the antennal scapes 
having two or more hairs that are curved 
or inclined toward the base of the scape; (v) 
the eyes minute, usually with only 1-3 
ommatidia in total; (vi) the preocular 
carina short and ending before the level 
of the eye; (vii) the propodeal spines 
usually present; and (viii) the head and 
alitrunk usually sculptured but the meso- 
pleuron and metapleuron entirely smooth 
and shining. 

Strumigenys acarai shares with S. cari- 
nithorax Borgmeier, in addition to the 
character states mentioned above, the pres- 
ence of a median fine longitudinal carina 
on the mesonotum. Strumigenys acarai 
differs from S. carinithorax, however, by 
having the ground-pilosity of the head, 
from above level of eye to close to occipital 
margin, very narrowly spatulate (almost 
simple) rather than spatulate as in S. 
carinithorax; the mandibles with a pair of 
minute inconspicuous preapical denticles 
proximal to the midlength of the mand- 
ibles rather than with a pair of spiniform 
preapical teeth as found in S. carinithorax, 
which are located in the distal third, and 
with a minute pair of denticles that may be 
difficult to see that are just proximal to the 
midlength of the mandibles (Bolton 2000); 
the leading edge of the antennal scapes 
with some multifurcated narrowly spatu- 
late hairs rather than spoon-shaped hairs 
of S. carinithorax. Strumigenys acarai shares 
with S. waiwai (described here) the pres- 
ence of multifurcated hairs. In the former 
species, however, these hairs seem to be 
restricted to the leading edge of the 


Journal of Hymenoptera Research: Festschrift Honoring Roy Snelling 

Figs 26-36. Scanning electron micrographs of a paratype worker of Strumigenys acarai, new species. 26, Head 
and mandibles in full-face view. 17, Cephalic capsule in full-face view. 28, Mandibles in dorsal view. 29, Tooth 
on inner margin of mandibles in dorsal view. 30, antennal scapes and hairs on leading edge of scape in full-face 
view. 31, Lateral view. 32, Dorsal view. 33, Lateral view of mesosoma. 34, Dorsal view of mesosoma. 35, Dorsal 
view of waist segments (petiole and postpetiole). 36, Dorsal view of postpetiole and first gastral tergite. 

antennal scapes, whereas in the latter these 
hairs are present on the dorsum of the 
head. The two species described here (S. 
acarai and S. waiwai) also differ from each 
other in mandibular dentition (inconspic- 
uous pair of teeth at midlength of mand- 
ibles in S. acarai, and having a pair of 
spiniform teeth and a minute, but conspic- 
uous pair of teeth at midlength of mand- 

ibles in S. waiwai), in the sculpture of the 
dorsum of the promesonotum (rugulose 
and with a conspicuous median longitudi- 
nal ruga in S. acarai, and strongly aerolate 
in S. waiwai), and in the length of the 
costulae on first gastral tergite (longer than 
the maximum length of the disc of post- 
petiole in S. acarai, and barely as long as the 
disc of postpetiole in S. waiwai). 

VOLL-ME 19, Xl-mber 1, 2010 


Hgs 26-36. Continued. 


In Bolton's (2000) key, Stnimigenys acarai will not ke}" out to any of the kno^sn species. The key for 
the species of Stnimigmys of the Xeotropics can be modified as below to include 5. acarai. 
Xumbering of couplets follows Bolton (2000). 

48. Cephalic dorsum ^sith two pairs of short erect hairs that differ from other cephalic 

ground-pilosit}-, one pair close to occipital margin, other close to highest point of 

\'ertex. \'entral surface of petiole -^sith curtain or fringe of spongiform tissue, or at 
least ^\ spongiform lobes linked by carina couplet 49 of Bolton (2CXX)) 


Journal of Hymenoptera Research: Festschrift Honoring Roy Snelling 

Figs 37-39. Automontage images of the holotype worker of Strumigenys acarai, new species. 37, Full-face view. 
38, Lateral view. 39, Dorsal view. 

- Cephalic dorsum w^ithout or w^ith one pair of short erect hairs that differs from other 

cephalic ground-pilosity, w^hen one present it is close to occipital margin. Ventral 
surface of petiole without spongiform tissue, sometimes w^ith rounded or angular 

anteroventral cuticular process 52a (different to couplet 52 in Bolton 2000) 

52a. Mandible w^ith minute inconspicuous denticle close to midlength .... acarai new species 

- Mandible with 1 or 2 very conspicuous spiniform preapical teeth, distal one in apical 

third, proximal one close to midlength couplet 52 in Bolton (2000) 

Strumigenys waiwai Sosa-Calvo, Schultz, 

and LaPolla, n. sp. 

(Figs 40^7) 

Material examined. — Holotype: worker, labeled 
"GUYANA: Upper Takutu-Upper Essequibo, 
Acarai Mountains, camp edge Kamoa River, 
394 m., 58 49.929' W, r32.786' N; 22.X.2006; /. 
Sosa-Calvo, T.R. Schultz; V forest; leaf-litter 
sample. (TRS 061022-LS04)" USNM ENT 
No. 00537291. (UGBC). Paratypes: 2 workers, 
same locality as in holotype. USNM ENT 

No. 00537290, 00537292; 1 worker, labeled 
"GUYANA: Upper Takutu-Upper Essequibo, 
Acarai Mountains, camp edge Kamoa River, 
530 m., 58=50.299' W, 1=33.046' N; 24.X.2006; /. 
Sosa-Calvo, T.R. Schultz, C.J. Marshall, R. Wil- 
liams; V forest; leaf-litter sample. (JSC 061024- 
LSIO)" USNM ENT No. 00537293. (USNM). 

Diagnosis (worker). — Small (TL 1.35- 
1.45); cephalic margin with multi-furcate 
hairs; leading edge of antennal scapes at 
least with one hair that curves towards 
base of scape; eyes small, consisting of two 

VOLL-ME 19, Nl-mber 1, 2010 


or three ommatidia; inner margin of 
mandibles with a pair of spiniform teeth 
and a minute but conspicuous pair of teeth 
at midlength of mandibles; ventral margin 
of petiole with angular ventral process. In 
some workers (paratypes) ventral process 
of petiole very reduced. 

Descriiption (worker). — Head: leading 
edge of antennal scapes with at least one 
hair curving toward base of antennal 
scape; hairs on leading edge of antennal 
scapes narrowly spatulate or simple; inner 
margin of mandibles with pair of preapical 
spiniform teeth close to apicodorsal tooth 
and minute but conspicuous pair of teeth 
at midlength of mandibles; anterior margin 
of clypeus slightly concave (Figs 40, 42); 
dorsum of clypeus finely reticulate; dor- 
sum of head markedly areolate and with 
multi-furcate hairs on posterior occipital 
portion (Figs 40-41, 43); upper margin of 
scrobes with row of simple hairs that cur\^e 
anteriorly and with two flagellate hairs, 
one of which is in apicoscrobal position; 
eyes very reduced, with 2 or 3 ommatidia; 
ocular carina weakly developed, short and 
not reaching level of eyes; cephalic dor- 
sum, in profile, with 2 pairs of inconspic- 
uous erect simple hairs, very difficult to see 
and perhaps fragile and easily lost but 
differing from bifurcating pilosity that 
surrounds them. Mesosoma: dorsum of 
promesonotum, propodeum, and declivit}^ 
of propodeum strongly areolate; dorsum of 
pronotum with elongate pair of hairs in 
addition to those at humeri; humeral hair 
flagellate; promesonotal spiracle in dorsal 
view projecting laterally, giving pronotum 
wider appearance and mesonotum and 
propodeum narrower appearance; mesono- 
tum with pair of elongate hairs; sides of 
pronotum and anepistemum strongly areo- 
late; mesopleuron and metapleuron mostly 
smooth and shining; dorsum of promeso- 
notum and propodeum with simple sub- 
decumbent hairs; propodeal spines minute 
and acute, subtended by broad lamella on 
declivity. Metasoma: dorsum of peduncle, 
disc, and sides of petiole strongly areolate; 

in lateral view, petiole with long ventral 
process. In one worker (paratype) ventral 
process reduced to small tooth (Fig. 43); 
petiole, in lateral view, subquadrate; sides 
of petiole, in dorsal view, with conspicuous 
triangular crest; postpetiole, in lateral view, 
with large ventral spongiform lobes; disc of 
postpetiole with some longitudinal rugae; 
areas betw^een rugae smooth and shining; 
dorsum of postpetiole with decumbent 
hairs; first gastral sternite with pad of 
spongiform tissue; first gastral tergite 
smooth and shining, with some conspicu- 
ous longitudinal basigastral costulae, barely 
as long as disc of postpetiole; first gastral 
tergite mostly with subdecumbent and 
decumbent hairs and some erect hairs 
(lacking in some paratype specimens. It is 
probable that these hairs are very fragile 
and lost easily). 

Measurements: holotype (and paratypes): 
GL = 0.31 (0.28-0.31), HL = 0.33 (0.31- 
0.34), HW = 0.26 (0.25-0.26), ML = 0.18 
(0.18-0.19), PL = 0.17, PPL = 0.08 (0.08- 
0.09), PW = 0.16 (0.14-0.16), SL = 0.21 
(0.19-0.21), TL = 1.44 (1.35-1.45), WL = 
0.36 (0.32-0.36). Indexes: CI = 79 (76-80), 
MI = 55 (53-58), PI = 48 (47-51), SI = 81 
(77-82). (n = 4) 

Gyne and male. — UnknowTi 

Etymology. — This species is named after 
the Wai-Wai indigenous people, who de- 
pend on the area where we collected this 
species for their sustenance. Without their 
guidance, support, and permission to con- 
duct research on their land, this work 
would not have been possible. 

Comments. — Strumigenys waiwai is most 
similar to members of the Neotropical 
silvestrii-gr oup. Strumigenys waiwai shares 
with most of the 18 known species in this 
group the following characters: (i) the 
smaH size (HL 0.31-0.34, HW 0.25-0.26, 
TL 1.35-1.45, WL 0.32-0.36. In members of 
the silvestrii-group HL 0.36-0.52, HW 0.28- 
0.44, TL 1.5-2.2, WL 0.36-0.56); (ii) the 
absence of intercalary^ tooth between the 
apicodorsal and apicoventral teeth in the 
apical fork of the m.andibles (this character 


Journal of Hymenoptera Research: Festschrift Honoring Roy Snelling 

Figs 40-47. Scanning electron micrograph of a paratype worker of Strumigenys waiwai, new species. 40, Head 
and mandibles in full-face view. 41, Cephalic capsule in full-face view. 42, Mandibles in dorsal view. 43, Lateral 
view. 44, Dorsal view. 45, Dorsal view of mesosoma. 46, Waist segments in dorsal view. 47, Postpetiole and first 
gastral tergite in dorsal view. 

State is shared with all the species in the 
group, except for S. xochipili Bolton from 
Mexico, which possesses a single interca- 
lary tooth); (iii) the presence, on the inner 
margin of mandibles, of a spiniform pair of 
preapical teeth close to the apicodorsal 
tooth of the apical fork and, in addition, a 
minute pair of denticles on the midlength 
of the mandibles that may be difficult to 

see; (iv) the antennal scapes short to 
moderate (SI 76-80. In members of the S. 
silvestrii species group the SI 62-91), and 
with some hairs on the leading edge of the 
antennal scape that are curved toward the 
base of the scape; (v) the eyes very small, 
commonly formed by 1-3 ommatidia in 
total (some members of the S. silvestrii 
species group with 6 or more ommatidia); 

Volume 19, Number 1, 2010 


Figs 40-47. Continued. 

(vi) the preocular carina short or weakly 
developed, ending before level of the eye; 
(vii) the propodeum, in profile, usually with 
a triangular pair of teeth that are subtended 
by a lamella or carina that extends down the 
declivity; and (viii) ventral surface of petiole 
lacking spongiform tissue instead with a 
small, but conspicuous crest or ventral 
process (members of the silvestrii-group 
usually lack spongiform tissue on ventral 
margin of petiole except for the species S. 
nastata Bolton, S. perdita Bolton, and S. 
calamita Bolton). 

Variations in the presence of standing 
erect hairs on the cephalic dorsum were 
observed in the specimens studied (holo- 
type and paratypes). In the material exam- 
ined, some workers of S. waiwai have the 
two pairs of erect simple hairs on the 
dorsum of head that differ from the 
cephalic ground-pilosity, of which one pair 
is located close to occipital margin and the 
other is located close to the highest point of 
vertex. Among the variations observed in 
other specimens are: the two pairs of 
standing hairs are present, but curving at 
the tips, or only one pair of hairs is visible, 
or the hairs are difficult to see, or the hairs 
are missing. Apparently these hairs are 
very fragile and can be easily lost; therefore 
specimens may appear to have no hairs. 
Within the silvestrii-^owp , only individ- 
uals in the species S. calamita, S. nastata, S. 
perdita, and S. timicala Bolton, all endemic 
to Central America, share with individuals 

of S. waiwai the presence of two pairs of 
erect hairs on the cephalic dorsum, but 
these species differ from S. waiwai by 
having a fringe or curtain of spongiform 
tissue on the ventral margin of the petiole, 
whereas S. waiwai has an angular crest or 
small ventral process. In addition, S. waiwai 
differs from: S. nastata by having the 
antennal scape without a projecting nar- 
row cuticular lamella that arises proximal 
to the subbasal bend; S. timicala by having 
the preapical tooth of the mandible sepa- 
rated from the apicodorsal tooth by a 
distance twice its length, rather than hav- 
ing the preapical tooth very close to the 
apicodorsal tooth; S. calamita and S. perdita 
by having the first gastral tergite generally 
with subdecumbent or decumbent simple 
hairs (in some specimens it is possible to 
see, in addition, a few erect simple hairs) 
rather than entirely short stout hairs that 
are remiform to claviform (in S. calamita) or 
simple erect and stiff (in perdita); and from 
all four species and any other species in the 
S. silvestrii-group by: (i) having the dorso- 
lateral margin of the head with two freely 
laterally projecting elongate or short fla- 
gellate hairs, one of which is located at the 
level of the vestigial eye and the other the 
apicoscrobal hair. This character state is 
also shared with S. lanuginosa but these 
hairs are shorter in S. waiwai than in S. 
lanuginosa; (ii) having hairs on the upper 
margins of the antennal scrobe simple and 
curving anteriorly rather than spoon- 

34 Journal of Hymenoptera Research: Festschrift Honorevg Roy Snelling 

shaped, spatulate, or narrowly spatulate pilosity composed of short erect or sub- 
and curving anteriorly (except for perparva decumbent multifurcated hairs rather than 
in which case these hairs are posteriorly spoon-shaped, spatulate, or narrowly spat- 
curved); and (iii) the cephalic ground- ulate hairs. 


In Bolton's (2000) key, Strumigenys zvaiivai keys out to S. perdita. The key for the species of 
Stnimigenys of the Neotropics can be modified as below to properly include S. waiwai. Numbering 
of couplets follows Bolton (2000). 

47. In full-face view upper scrobe margin with a row of 4-5 broadly spatulate to spoon- 

shaped hairs that are curved posteriorly perparva 

In full-face view upper scrobe margin with row of simple or spatulate to spoon-shaped 
hairs that are all curved anteriorly couplet 48 in Bolton (2000) 

48. Cephalic dorsum with two pairs of short erect hairs, one pair close to occipital margin, 

the other close to highest point of vertex. Ventral surface of petiole with curtain or 
fringe of spongiform tissue, or at least with spongiform lobes linked by carina. If 
spongiform tissue reduced to angular anteroventral process present, then 
dorsolateral margin of head with two freely laterally projecting short flagellate 
hairs, one at level of the vestigial eye, the other the apicoscrobal 49 

- Cephalic dorsum without or with one pair of short erect hairs, when one is present it is 

close to occipital margin. Ventral surface of petiole without spongiform tissue, 
sometimes with rounded or angular anteroventral cuticular process. Dorsolateral 
margin of head without projecting flagellate hairs or with single hair, in 
apicoscrobal position couplet 52 in Bolton (2000) 

49. Distal preapical tooth conspicuous and obviously spiniform, located markedly 

proximal of the apicodorsal tooth and at about right angle to long axis of the 
mandible. Distal preapical teeth of opposing mandibles so long that their apices 
meet or even slightly overlap when mandibles fully closed. Mandibles always with 

small denticle just proximal of the inner midlength 50 

Distal preapical tooth small, thorn-like and not obviously spiniform, located very close 
to the apicodorsal tooth and inclined toward it. Distal preapical teeth of opposing 
mandibles so short that their apices are widely separated when mandibles fully 
closed. Mandible usually without trace of denticle proximal to inner midlength but 
rarely vestigial denticle visible timicala 

50. Leading edge of scape at subbasal bend with projecting convex cuticular lamella; 

lamella originates close to scape base and terminates just distal of the bend. Distal 
preapical tooth about same distance from proximal preapical denticle as it is from 
apicodorsal tooth nastata 

- Leading edge of the scape at subbasal bend without projecting convex cuticular 

lamella. Distal preapical tooth closer to apicodorsal tooth than it is to proximal 

preapical denticle 51a 

51a. Leading edge of antennal scape with spoon-shaped or narrowly spatulate hairs. In 
full-face view upper scrobe margin with row of spatulate to spoon-shaped hairs. 
Cephalic ground-pilosity spoon-shaped or spatulate .... couplet 51 of Bolton (2000) 
Leading edge of antennal scape with simple or filiform hairs. In full-face view, upper 
scrobe margin with a row of simple or filiform hairs. Cephalic ground-pilosity 
multifurcated waiwai new species 

VOLL-ME 19, Number 1, 2010 



We would like to thank Bam- Bolton for confirming 
the new species. We thank George Else, Suzanne 
Lewis, Christine Taylor (BMNH), and Carlos Roberto 
Brandao (\1ZSP) for specimen loans. Ted Suman 
(USNM) sorted and prepared the specimens, Scott 
Whittaker (USNDvI) assisted with the scanning elec- 
tronic micrographs, and Eugenia Okonski (USXM) 
databased and labeled the specimens. For corrections 
and comments on earlier \-ersions of the manuscript 
we thank Matt Buffington, Da\'id Furth, Victor H. 
Gonzalez, Akito Kawahara, Robert Kula, and John T. 
Longino. Claudia M. Ortiz and Monica Pava-Ripoll 
read and commented on the Spanish-language ab- 
stract. We thank Brian Fisher and an anon\Tnous 
reviewer for comments that greatly improved the 
manuscript. This project was supported in part by 
XSF DEB 0110073 and DEB 0431330 to TRS, by a 
National Geographic Societ\' Committee for Research 
and Exploration grant to JSL and TRS, by the 
Smithsonian Institution USS Restricted Endowment 
Fund grant to TRS, and by the Smithsonian Institu- 
tion's Biological Diversity of the Guiana Shield 
Program (BOG). This manuscript is number 158 in 
the BDG Program's publication series. 


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Volume 19, Number 1, 2010 


Appendix 1 

Map of Guyana with locations of places sampled, modified from LaPolla et al. (2007). 
Legend (Abbreviations follow those of Appendix 1): (1) CWC; (2) MAB, MAD, MAP, 
MAU; (3) MHC; (4) IFR; (5) KMM; (6) ARC, ANR^ ANR^; (7) KRC. 


Journal of Hymenoptera Research: Festschrift Honoring Roy Snelling 

Appendix 2. Dacetine ant species currently known from Guyana. 



Notes /Reference 

Acanthognnthus hrevicornis 

M.R. Smith 
A. lentus Mann 
A. ocellatiis Mayr 
A. stipulosus? 

Brown & Kempf 
Daceton armigerum 

Pyramica alberti 

P. auctideiis Bolton 
P. beebei (Wheeler) 
P. cincinnata (Kempf) 
P. crassicornis (Mayr) 

P. dahJanae n. sp. 
P. denticulata (Mayr) 

P. dqjressiceps (Weber) 

P. glenognatha Bolton 
P. inusitata (Lattke) 
P. mariae n. sp. 
P. metopia (Brown) 
P. metrix Bolton 
P. mirabilis (Mann) 
P. stenotes Bolton 
P. subedentata (Mayr) 

P. tluixteri (Wheeler) 
P. urrhobia Bolton 
P. villiersi (Perrault) 
Strumigenys acarai 
S. cordovensis Mayr 
S. cosmostela Kempf 
S. dolichognatha 

S. dyseides Bolton 
S. elongata Roger 

S. godmani Forel 

S. pariensis Lattke & 

S. perparva Brown 

S. precava Brown 

S. royi n. sp. 
S. ruffl Bolton 



IFR, Essequibo River, Rupununi 

Savannah, Tukeit 
ANR^ Kartabo, KRC, KRT 




ANR\ ANR^ Guyanas, KRC, KRT 


ARC, ANR\ ANR^ Berbice Dubulay 
Ranch, CWC, IFR, KMM, KRC, KRT, 
MAB, MHC, Morabukea, R. Mazaruni 












IFR, Kartabo, MAB, MHC 



KRT, MAB, MHC, Morabukea, 


ARC, ANR\ ANR^ Berbice Dubulay 

Ranch, CWC, IFR, KMM, KRC, KRT, 

ANR2, Between R. Cuyuni & R. 

Mazaruni, Kamakusa, KRC, MHC, 


New record 

New record 
New record 
New record 

Bolton 2000; Fernandez & Sendoya 

2004; Wheeler 1916; this study 
Bolton 2000; Fernandez & Sendoya 

2004; this study 
New record 
New record 
New record 
Kempf 1972; Bolton 2000; 

Fernandez & Sendoya 2004; 

this study 
New species 
Bolton 2000; Fernandez & Sendoya 

2004; this study 

Bolton 2000; Fernandez & Sendoya 

New record 
New record 
New species 
New record 
New record 
New record 
New record 
New record 

New record 
New record 
New record 
New species 
New record 
New record 
Bolton 2000; Fernandez & Sendoya 

2004; this study 
New record 
Bolton 2000; Fernandez & Sendoya 

2004; this study 
Bolton 2000; Fernandez & Sendoya 

New record 

Bolton 2000; Fernandez & Sendoya 
2004; this study 

Bolton 2000; Fernandez & Sendoya 
2004; this study 

New species 
New record 

5. silvestrii Emery 
5. smilax Bolton 


S. smithii Forel 
5. trinidadensis 




S. trudifera Kempf 

& BrowTi 
S. waiwai n. sp. 


Volume 19, Nl-mber 1, 2010 39 

Appendix 2 Continued. 

Tax on Localit}' Xotes/ Reference 

New record 
New record 
New record 
New record 

New record 

New species 

Abbreviations for localities: ARC: Upper Takutu-Upper Essequibo, Acarai Mountains, Romeo Camp, 58''56.789'W, 
1'23.147'N, elev. 290 m; ANR^: Upper Takutu-Upper Essequibo, Acarai Mountains, New Romeo Camp, 
58'57.828'W, ri9.938'N, elev. 1069 m; ANR": Upper Takutu-Upper Essequibo, Acarai Mountains, New Romeo 
Camp, 58^57.49'W, 1^20.854'N, elev. 750 m; CWC: Calm Water Creek, 58^37.16'W, 6^28.06'N, elev. 20 m; IFR: 
Iwokrama Forest Reser\'e Whitewater Camp, 58'50.992'W, 4'43.890'N, elev. 60 m; KMM: Kanuku Mountains 
near Moco-Moco Falls, 59'38.376'W, 3"-17.297'N, elev. 224 m; KMN: Kanuku Mountains near Nappi Creek 
Camp, 59'33.963'W, 3-21.018'N, elev. 128 m; KRC: Upper Takutu-Upper Essequibo, Kamoa River Camp, 
58'-49.929'W, r32.786'N, elev. 530 m; KRT: Upper Takutu-Upper Essequibo, Kamoa River top mountain, 
58^50.299' W, 1"33.046'N, elev. 717 m; MAB: Base Camp Mount. Ayanganna, 59'-55.486'W, 5^20.063'N, elev. 
732 m; MAD: Dicymbe Camp Mount Avanganna, 59'54.632'W, 5'17.76b'N, elev. 717 m; MAF: Falls Camp Mount 
Ayanganna, 59'-57.563'W, 5'22.332'N, elev. 1134; MAU: Upper Forest Mount Avanganna, 59'-57.969'W, 
5^22.483'N, elev. 1300 m; MHC: Mabura FHU, 58=41.982'W, 5=09.313'N, elev. 64 m. "New record" refers to a new 
record for Guyana. 

Appendix 3 

(NOTE: This key is an adaptation of that of Bolton [2000] to include only the species that occur in 
Guyana. This key also includes modifications from Brown and Kempf [1969]; Bolton [1994]) 

1. Antenna 11-segmented 2 

- Antenna 4 to 6-segmented 6 

2(1). In lateral view, antennal scape passes below eye when in resting position. Propodeal 

node bidentate. Palpal formula 5,3 Daceton annigerum 

- In lateral view, antennal scape passes above eye when in resting position. Propodeal 

node unarmed. Palpal formula 0,1 3 

3(2). Preapical area of masticatory margin of mandibles with irregular denticles 

Acanthognathus hrevicomis 

- Preapical area of masticatory margin of mandibles lacking denticles 4 

4(3). Node of petiole with convex anterior face and flat, sloping posterior face, not evenly 

rounded Acanthognathus stipulosus 

- Node of petiole low and evenly rounded when seen in profile 5 

5(4). Fossae on posterior half of dorsum of head smaller, mostly separated by flat, smooth 

spaces Acanthognathus ocellatus 

- Fossae on posterior half of dorsum or head large, mostly contiguous or separated by 

single, simple longitudinal rugulae Acanthognathus lentus 

6(1). Mandibles inserted on sides of anterior cephalic margin and converging towards apex 
when closed. Inner margin of mandibles generally with numerous teeth or 
denticles 7 (Pyramica) 

- Mandibles inserted on median portion of anterior cephalic margin and diverging 

towards apex when closed. Inner margin of mandibles with 0-2 preapical 
teeth 25 Istrumigenys) 

40 Journal of Hymenoptera Research: Festschrift Honoring Roy Snelling 

7(6). In full-face view, mandibles sublinear to linear, elongated and narrow; when closed, 
mandibles contacting each other only in apical halves or less of their lengths; either 
with an elongate space between mandibles or their inner margins convex so that 

margins touch, or nearly touch, near midlength 8 

In full-face view, mandibles either short and trap-like, or triangular to elongate- 
triangular; when closed contacting through most or all of their exposed length, 
lacking an elongate space between mandibles or at most with diastema basally 
between basal lamella and basal tooth 15 

8(7). Disc of postpetiole smooth or with weak longitudinal costulae in parts, never densely 

reticulate-punctate Pyramica dahlanae 

Disc of postpetiole densely reticulate-punctate over most or all of its surface 9 

9(8). Inner margin of mandible with clearly defined submedian tooth or distinctly enlarged 
denticle at or just distal of midlength of mandible, this tooth or denticle obviously 
larger than any other preapical dentition that may be present distal to it, if two 
distinctive enlarged teeth present, distal located at about the apical third and 
proximal in basal third of mandible length. Labral lobes very long and slender, 

trigger hairs at apices of lobes short 10 

Inner margin of mandible without a tooth or distinctly enlarged denticle at or near the 
midlength that is obviously larger than any other preapical dentition that may be 
present distal to it, if two distinctive enlarged teeth present, both of them closer to 
preapical dentition than to midlength of mandible. Labral lobes short, trigger hairs 
at apices of lobes long 13 

10(9). Pronotal humeral hair long and flagellate. Mesonotum with single pair of long 

flagellate hairs Pyramica metopia 

Pronotal humeral hair sometimes absent usually present, short-spatulate to filliform, 
never flagellate. Mesonotum without flagellate hairs 11 

11(10). Scape narrow basally; anterior margin of scape beyond base abruptly expanded and 
almost lobate at subbasal angle, scape distinctly widest at this point. Dorsolateral 
margin of head lacking apicoscrobal hair. Postpetiole, in profile, swollen or 

subglobular Pyramica crassicomis 

Scape gradually broadening from base to apex; anterior margin convex but not 
abruptly expanded at subbasal angle, scape widest at or near its midlength. 
Dorsolateral margin of head with an apicoscrobal hair of some form. Postpetiole, in 
profile, not swollen nor subglobular 12 

12(11). Inner margin of mandible with single enlarged preapical tooth, located near midlength; 
other minutely denticles present, but without a second equally sized tooth. Larger 

species (HL 0.61-0.63, HW 0.41-0.43, AL 0.58-0.60) Pyramica stenotes 

Inner margin of mandible with two enlarged preapical teeth of approximately equal 
size; in addition to other minutely denticles. Smaller species (HL 0.50-0.52, HW 
0.34-0.36, AL 0.46-0.50) Pyramica auctidens 

13(9). In lateral view, dorsum of mesosoma with 4-6 pair of stout remiform standing hairs 
(not including those at humeri). Mandibles short (MI 49-54). In full-face view, inner 
margins of mandibles convex and, when entirely closed, touching at about 

midlength Pyramica subedentata 

In lateral view, dorsum of mesosoma with single pair of standing hairs (not including 
those at humeri). Mandibles larger (MI 72-85). In full-face view inner margins of 
mandibles more or less straight to shallowly concave 14 

14(13). Inner margin of mandibles with 5-10 preapical denticles of similar size. Metapleuron 

entirely densely reticulate. Peduncle of petiole short, PI 38-^ . . . Pyramica denticulata 

- Inner margin of mandibles with 3-4 preapical denticles, two of which are distinctly 
much larger than rest. Metapleuron in most of its surface smooth and shining. 
Peduncle of petiole elongate, PI 48-49 Pyramica mariae 

Volume 19, Number 1, 2010 41 

15(7). With head in full-face view, anterior margin of scape with projecting curved hairs, of 
which one or more, distal to subbasal bend, distinctly curve toward base of scape. 
These hairs may be simple, spatulate, spoon-shaped, or wire-like 16 

- With head in full-face view, anterior margin of scape without projecting hairs that 

distinctly curve toward base of scape. Scape edge may have elongate simple 
straight or flagellate projecting hairs present; or may have entirely anteriorly or 

apically directed short hairs; or lacking hairs 21 

16(15). Pronotal humeral hair present, may be filiform, flagellate, remiform, or clavate; 
humeral hair always distinctly differentiated from any other pilosity that may be 
present on dorsal pronotum 17 

- Pronotal humeral hair absent; humerus without a hair that is distinctly differentiated 

from any other pilosity that may be present on dorsal pronotum 20 

17(16). Ventral surface of petiole in profile with spongiform tissue reduced to absent; 
discounting anterior subpetiolar process (if present) usually with narrow non- 
spongiform cuticular carina, but if weakly spongiform strip occurs then its 
maximum depth only fraction of depth of peduncle. Disc of postpetiole usually 
sculptured at least in part, only rarely mostly smooth 18 

- Ventral surface of petiole in profile with deep, conspicuous and very obviously spongiform 

curtain, its maximum depth at least half that of peduncle and usually more. Disc of 

postpetiole completely unsculptured and glassy smooth Pyr arnica alberti 

18(17). Metapleuron and side of propodeum entirely reticulate-punctate 19 

- Metapleuron and side of propleuron mostly or entirely smooth and shinning .... 

Pyramica cincinnata 

19(18). Pronotal humeral hair elongate and freely projecting, slightly flattened apically and 
more or less straight. Scape in dorsal view slender, broadest point distinctly distal 
of midlenght. Anterior margin of clypeus very shallowly convex in full-face view. 
Disc of postpetiole not entirely densely reticulate-pimctate Pyramica urrhobia 

- Pronotal humeral hair very short, clavate. Scape in dorsal view broad and flattened, 

broadest point proximal of midlenght, at or just distal of subbasal bend. Anterior 
margin of clypeus transverse to very shallowly concave in full-face view. Disc of 

postpetiole entirely densely reticulate-punctate Pyramica metrix 

20(16). Promesonotum, side of mesosoma, and disc of postpetiole finely reticulate-punctate. 
Head in profile incredible dorsoventrally flattened; at eye level depth of head 
capsule scarcely more than twice vertical diameter of the eye. Ventral margin of 
petiole lacking curtain of lamellate or spongiform tissue .... Pyramica depressiceps 

- Promesonotum and side of mesosoma smooth and shining, disco of pospetiole not 

reticulate-punctate. Head in profile not strongly dorsoventrally flattened; at eye 
level depth of head capsule distinctly more than twice vertical diameter of eye. 
Ventral margin of petiole with lamellate curtain that extends entire length of 

segment Pyramica thaxteri 

21(15). With head in full-face view dorsolateral margin behind level of eye with laterally projecting 

hairs present; at least an apicoscrobal hair but often more along margin 22 

- With head in full-face view dorsolateral margin behind level of eye without laterally 

projecting hairs of any form; any hairs that do occur are minute and closely 

appressed, not at all projecting 24 

22(21). Midline of clypeal dorsum raised into a high-arched thick longitudinal crest that 
extends entire length of sclerite. With postpetiole in profile ventral spongiform lobe 
either completely absent or reduced to minute triangular vestige anteriorly on 
stemite Pyramica inusitata 

- Midline of clypeal dorsim:i not raised into a high longitudinal crest that extends length 

of sclerite. With postpetiole in profile ventral spongiform lobe fully developed, 
basally extending length of stemite and conspicuously convex apically 23 

42 Journal of Hymenoptera Research: Festschrift Honoring Roy Snelling 

23(22). Dorsal surface of petiole without erect cuticular lamella on peduncle and anterior face 
of node. Dorsolateral margin of propodeum without an erect lamella on each side; 
lamella on propodeal declivity narrow to cariniform, in profile its maximum width 
much less than length of postpetiole disc Pyramica villiersi 

- Dorsal surface of petiole with erect cuticular lamella that extends along peduncle and 

ascends anterior face of node, terminating just behind anterodorsal angle. Dorsolateral 
margin of propodeum with a tall erect cuticular lamella on each side that is continuous 
with extremely broad lamella on declivity; in profile maximum width of lamella on 

declivity equal to length of postpetiole disc Pyramica mirabilis 

24(21). Dorsal outline of clypeus angled down at about 45 degrees to line of vertex. Ventral 
surface of petiole with well-developed curtain, of spongiform or translucent 
lamellar tissue, that runs most or all of the length of the segment. Head elongate (CI 
68-70) Pyramica beehei 

- Dorsal outline of clypeus not angled down from line of vertex. Ventral surface of 

petiole lacking curtain of spongiform or lamellar tissue. Head short and broad (CI 

91-96) Pyramica glenognatha 

25(6). Anterior margin of scape with all hairs curved or inclined toward apex of scape, 
without hairs that curve toward base of scape, and without a series of hairs at right- 
angles to long axis of scape shaft 26 

- Anterior margin of scape either with one to many hairs that distinctly curve toward 

base of scape, or rarely with hairs that are at right angles to long axis of scape shaft; 

never with all hairs obviously curved or inclined toward apex of scape 29 

26(25). Mandibles relatively short, MI < 75. Bulla of femoral gland located in apical quarter of 
dorsum of each leg; each bulla usually appears as pale oval patch, less commonly 
as short streak 27 

- Mandibles relatively long MI > 85. Bulla of femoral gland located close to midlength on 

dorsum of each leg; each bulla appears as pale elongate streak or as oval patch ... 28 
27(26). Mandibles short and stout (MI 41-48), broad and powerful, outer margins strongly 
bowed outwards. Declivity of propodeum in profile with tooth or spine above and 
triangular lobe or tooth below, two linked by lamella. First gastral tergite glassy 
smooth behind minute to vestigial basigastral costulae Strumigenys godmani 

- Mandibles long and linear (MI 57-60), thick throughout most of their length and 

abruptly narrowing just before apex by sudden oblique divergence of inner margin, 
outer margins straight to slightly convex. Declivity of propodeum in profile with 
single tooth or spine, lacking second tooth or lobe below. First gastral tergite finely 
reticulate-substrigulate with some verrucose sculpture confined to basigastral 

area Strumigenys royi 

28(26). In full-face view distal preapical tooth of mandible is closer to proximal preapical 

tooth than it is to apicodorsal tooth Strumigenys dolichognatha 

- In full-face view distal preapical tooth of mandible is closer to apicodorsal tooth than it 

is to proximal preapical tooth Strumigenys cordovensis 

29(25). Mandible without intercalary teeth or denticles that arise between apicodorsal and 

apicoventral teeth, nor that arise from dorsal base of apicoventral tooth 30 

- Mandible with 1 or 2 intercalary teeth or denticles that arise between apicodorsal and 

apicoventral teeth, or that arise from dorsal base of apicoventral tooth 40 

30(29). Mandible without preapical teeth or denticles 31 

Mandible with 1 or 2 preapical teeth or denticles 32 

31(30). Hairs of first gastral tergite flagellate, not flattened and ribbon-like through most of 

their length Strumigenys elongata 

- Hairs of first gastral tergite flattened and ribbon-like through most of their length, 

narrowly flagellate only in apical section Strumigenys pariensis 

32(30). Mandibles very long, MI > 100 Strumigenys trudifera 

Mandibles much shorter, MI < 75 33 

Volume 19, Number 1, 2010 43 

33(32). With head in profile preocular carina extends back aknost to apex of scrobe, well 
beyond level of minute eye. Bulla of femoral gland proximal of midlength on 
dorsum of middle leg, very conspicuous. Scape relatively long, SI 111-112 .... 

Strutnigenys smilax 

With head in profile preocular carina terminates at level of eye. Bulla of femoral gland 
distal of midlength on dorsum of middle leg or inconspicuous. Scape shorter, SI < 
100 34 

34(33). Large species (HL 0.86-1.02, ML 0.50-0.56, AL 0.80-1.00). Ventrolateral margin of head 
in front of eye, and side of head above it, deeply concave; margin and side appear 

excavated or constricted in oblique dorsal view Strutnigenys precava 

Smaller species (HL 0.39-0.45, ML 0.20-0.30, AL 0.36-0.47). Ventrolateral margin of 
head in front of eye, and side of head above it, not deeply concave; margin and side 
do not appear excavated or constricted in oblique dorsal view 35 

35(34). In full-face view upper scrobe margin with row of 4-5 broadly spatulate to spoon- 
shaped hairs that curve posteriorly Strumigenys perparva 

- In full-face view upper scrobe margin with row of simple, or narrowly spatulate to 

spoon-shaped hairs that all curve anteriorly 36 

36(35). Mandible with single spiniform preapicaltooth, in distal third, or with tooth in this 

position and denticle, that may be minute and difficult to see, close to midlength ... 37 
Mandible with minute inconspicuous denticle close to midlength lacking second tooth 

of any form Strumigenys acarai 

37(36). Leading edge of antennal scape with spoon-shaped or spatulate hairs. In full-face view 

upper scrobe margin with row of spatulate to spoon-shaped hairs, and with 

apicoscrobal hair only. Cephalic ground-pilosity spoon-shaped or spatulate .... 38 

Leading edge of antennal scape with narrowly spatulate or simple hairs. In full-face 

view upper scrobe margin with row of simple hairs and with two flageUate hairs, 

one of which is apicoscrobal hair. Cephalic ground-pilosity multi-furcate 

Strumigenys waiwai 

38(37). Hairs on first gastral tergite short and stout, broadly spatulate or remiform; elongate 
slender fine hairs absent or restricted to transverse row at extreme apex of 
sclerite Strumigenys silvestrii 

- Hairs on first gastral tergite elongate and slender, finely filiform to flageUate, or 

flexuous; short stout spatulate or remiform hairs entirely absent 39 

39(38). Mandibles relatively long, MI > 60. Disc of postpetiole smooth and shinning. 

Mesonotum without pair of erect flageUate hairs Strumigenys dyseides 

- Mandibles relatively short, MI < 60. Disc of postpetiole densely punctate to reticulate- 

punctate. Mesonotum with pair of erect flagellate hairs Strumigenys rut a 

40(29). Apical fork of mandible with single intercalary tooth or denticle that arises between apico- 

dorsal and apicoventral teeth, or arises from dorsal surface of apicoventral tooth 41 

- Apical fork of mandible with two intercalary teeth or denticles that arise between 

apicodorsal and apicoventral teeth; frequently represented by distinct intercalary tooth 

accompanied by less conspicuous or minute denticle Strumigenys cosmostela 

41(40). First gastral tergite very finely and densely longitudinaUy striolate-costulate and 
opaque. Apicoscrobal hair, pronotal humeral hair and standing hairs on 
mesonotum all flageUate. Entire body dull yellow to brownish-yeUow, gaster not 
contrasting with head and aUtrunk Strumigenys trinidadensis 

- First gastral tergite glassy smooth. Apicoscrobal hair, pronotal humeral hair and 

standing hairs on mesonotum aU stiff, simple to weakly remiform. Head and 
alitrunk reddish brown to brown, gaster blackish brown to black, both 
contrasting Strumigenys smiihii 

Vol. 19(1), 2010, pp. 44-50 

Two New Species of the strigatus Species Complex of the Ant Genus 
Cyphontyrmex (Hymenoptera: Formicidae) from Costa Rica and Panama 

William P. Mackay and Francisco Serna 

Department of Biological Sciences, Centennial Museum, The University of Texas, El Paso, TX 79968, 

USA and Grupo Sistematica de Insectos Agronomia SIA, Museo Entomologico UNAB, 

Carrera 30 #45-03, Universidad Nacional de Colombia, Bogota, COLOMBIA; 

Abstract. — The strigatus species complex is defined as those workers and females of Cyphomyrmex 
in which the preocular carina extends back to the vertex, delimiting the lateral margin of a 
depressed concave scrobe. The mandibles have 6 or more teeth and there is a single medial pronotal 
tubercle. The complex was previously reported only in South America, especially southeastern 
Brasil and northern Argentina. Two new species were found in Central America: C. andersoni from 
Costa Rica, and C. snellingi from Panama. Cyphomyrmex andersoni resembles C. quehradae, but can be 
separated as the hind femur is longer than the head capsule (shorter in C. quehradae). It can be 
differentiated from the similar C. hruchi as the mesosomal tubercles are distinct (indistinct in C. 
bruchi). Cyphomyrmex snellingi has the frontovertexal corners lobate and somewhat projecting 
posteriad. It is most similar to C.faunulus, but can be easily distinguished as the anterior mesonotal 
tubercle is not more developed than the other tubercles (much larger than the others in C.faunulus). 

Resumen. — El complejo strigatus del genero Cyphomyrmex se caracteriza porque las hembras y 
obreras presentan una carina preocular que se extiende posteriormente hasta el vertex y delimita la 
margen lateral de un escrobo antenal deprimido y concavo. Estas hormigas poseen mandibulas con 
6 o mas dientes, y un solo tuberculo pronotal mesial. El complejo strigatus se conocia solo de 
Suramerica, especialmente el sureste de Brasil y norte de Argentina. Dos nuevas especies fueron 
halladas en America Central: C. andersoni de Costa Rica, y C. snellingi de Panama. Cyphomyrmex 
andersoni es similar a C. quehradae, pero se diferencia porque el femur posterior es mas largo que la 
capsula cefalica (mas corto en C. quehradae). A su vez, C. andersoni puede ser diferenciada de C. 
hruchi porque los tuberculos mesosomales son distinguibles (no distinguibles en C. hruchi). 
Cyphomyrmex snellingi tiene las esquinas frontovertexales lobosas y algo proyectadas poste- 
riormente. Esta especie es m^as similar a C. faunulus, de la cual puede ser distinguida facilmente por 
el tuberculo mesonotal anterior no tan desarrollado. 

The ant genus Cyphomyrmex belongs to head, which covers most of the head. The 

the New World fungus growing ants of the mesosoma has a series of pairs of blunt 

tribe Attini, and presently contains 40 tubercles in nearly all species. The first 

species (Bolton et al. 2007). The genus is opisthogastral tergum* (see glossary in 

divided into two species complexes, the Serna and Mackay 2010) lacks tubercles. 

strigatus complex (Kempf 1964) and the Most surfaces are dull and without sculp- 

rimosus complex (Kempf 1965; Snelling and ture; the hairs are mostly limited to 

Longino 1992). Cyphomyrmex workers and appressed, often scale-like setae that are 

females are easily recognized, as the frontal nearly always restricted to the gaster and 

carinae form a shield on the dorsum of the the head. 

Volume 19, Number 1, 2010 


Most species nest in the soil, in rotten 
logs and stumps, or in hollow dead twigs. 
This genus also nests under bark, under 
moss, and within epiphytic pseudobulbs 
(Snelling and Longino 1992). Colonies are 
small, probably not exceeding 500 workers 
(Snelling and Longino 1992). All Cypho- 
myrmex species cultivate badisiomycete 
fungi in the tribe Leucocoprineae. In the 
C. rimosus group, these fungi grow in a 
yeast form (small masses of unicellular 
fungal cells) rather than in the multicellular 
mycelial form typical for all other attine ant 
gardens (Schultz et al. 2002; Schultz and 
Brady 2008). 

Workers of the strigatus complex can be 
recognized by the closed antennal scrobe 
(sometimes with poorly defined margins), 
mandibles with six or more teeth, and with 
a single medial pronotal tubercle (appar- 
ently a fusion of two tubercles). The species 
of the strigatus complex were previously 
considered to be primarily southern South 
American in distribution, although C. 
faunulus occurs as far north as Vene- 
zuela (Mayhe Nunes and Jaffe 1998). 
An unidentified species is found in Co- 
lombia (Fernandez and Palacio 1995) 
and an apparently new species was 
found in Ecuador (Tiputini) by Kari Ryder 
Wilkie ( 
cyphom3mnex.html) . 

In comparison, the workers of the rimo- 
sus complex have an open antennal scrobe 
(anteriorly), with the preocular carina 
curved mesially in front of the eye, and 
not directed to the posterior corner of 
the head, the mandibles have five teeth, 
and the pronotum lacks medial tubercles, 
or has a pair of tubercles. The species of 
the rimosus complex are widely distribu- 
ted from the United States to South 

Two new species of the strigatus species 
complex were found in Costa Rica and 
Panama. These new species wiU be in- 
cluded in a key to the species of Cypho- 
myrmex that can be found at http://www. / leb / antgenera.htm. 


The specimens were examined with a 
Zeiss stereoscope, at 64X, and were mea- 
sured with an ocular micrometer. The 
abbreviations are as follows: 

HL Head length, measured in full 

frontal view, from anterior 
margin of medial lobe of cly- 
peus to medial posterior mar- 
gin of frons 

HW Head width, measured in full 

frontal view, maximum width 
excluding eyes (Measured near 
posterior point of head) 

SL Scape length, excluding con- 


EL Eye length, maximum diameter 

of eye 

EW Eye width, maximum width of 

eye, perpendicular to EL 

WL Weber's length, a diagonal line 

from the top of the anterior 
edge of the pronotum to the 
posterior edge of the pos- 
teropropodeal lobes. 

CI CephaUc index, HW/HL X 100 

SI Scape index, SL/HL X 100 

OI Ocular index, EW/EL X 100 

MCZC Museum of Comparative Zool- 
ogy, Harvard University 

CWEM Collection of William and 
Emma Mackay, University of 
Texas at El Paso 

Terms followed by an asterisk are 
defined at the end of this paper and 
explained in the glossary of Serna and 
Mackay (2010). 


Cyphomyrmex andersoni new species 

(Figs 1-6) 

Diagnosis. — The worker is a small (total 
length about 2.5 mm, n=2) reddish-brown 
ant. The mandibles have six teeth, the 
frontal carinae do not reach the dorsad 
occular suture, the frontovertexal* corners 


Journal of Hymenoptera Research: Festschrift Honoring Roy Snelling 

are barely extended into auricle-like struc- 
tures; the pronotum has three angulate 
processes or teeth, including the medial 
process and two lateral processes, together 
with a pair of posterior swellings, and the 
mesonotum has a pair of conical processes; 
the propodeum has a pair of anterior, blunt 
processes and two well-developed angu- 
late posterior processes; and the posterior 
1/3 of the petiole is raised into a blunt 
process that appears bidentate when seen 
obliquely from above; the postpetiole has 
two parallel raised regions on the dorsal 
surface; and the gaster lacks longitudinal 
raised areas. The posterior femur has a 
distinctive ventral angulate process, fol- 
lowed distally by a poorly defined carina. 

The female and male are unknown. 

Distribution. — Known only from the 
states of Alajuela and Guanacaste, Costa 

Worker measurements (mm). — HL 0.76- 
0.78, HW 0.64-0.66, SL 0.58-0.60, EL 0.13- 
0.14, EW 0.08-0.10 WL 0.90-0.93. Indices: 
CI 84-85, SI 74-79, OI 65-73. Mandible 
with 6 teeth; anteclypeus broadly rounded; 
paraclypeal teeth* spiniform, moderately 
developed; frontal lobes and frontal ca- 
rinae relatively narrow, extending to front- 
overtexal corner*, forming carina that fuses 
with posterolateral margin of antennal 
scrobe, preocular carina continues poste- 
riorly to form mesial margin of antennal 
scrobe; eyes relatively small, extending 
past sides of head; scape relatively short, 
barely reaches frontovertexal comer; pro- 
notum with medial protuberance, 2 lateral, 
conical tubercles and 2 posterolateral lo- 
bate processes, anteroinfra angle of latero- 
pronotum developed; 2 angulate conical 
tubercles on mesonotum (height approxi- 
mately 0.05 mm); anterior margin of 
dorsopropodeum with 2 broad processes 
(height 0.03 mm), dorsopropodeum* rela- 
tively short (0.08 mm from notopropodeal 
groove to highest point of anterior tuber- 
cles) posteropropodeum* longer (0.25 mm, 
measured from anterior tubercles to meta- 
pleural lobe), propodeal spines small 

(length 0.04 mm) and rounded; petiole 
enlarged posteriorly, forming dorsal tuber- 
cles as seen in lateral view (length 0.1 mm, 
height 0.07 mm) that appears to have two 
lateral tiny bumps; postpetiole with lon- 
gitudinal medial depression flanked by 
two longitudinal ridges; all femora swol- 
len, fore femur with poorly developed lon- 
gitudinal carina along posteroventral mar- 
gin, middle femur similar, but carina 
poorly developed, posterior femur with 
well-developed longitudinal carina form- 
ing distinct angle distad about one third 
length from body. 

Erect hairs absent, except on mandible; 
hairs on scape and head appressed, hairs 
on ventral surface of head and anterior 
margin of procoxa subdecumbent, hairs on 
mesosoma, petiole, postpetiole, legs and 
gaster appressed. 

Type series. — Holotype worker, Costa Rica, 
Alajuela, 27 k N and 8 k W west of San Ramon, 
29-vi-6-vii-1999, R. Anderson # 19901, 99-109B, 
10°13'30"N; 34°35'30"W (MCZC). 

Additional material examined. — Costa Rica, 
Guanacaste, Cacao Field Station, 15-ii-1996, R. 
Anderson # 17682 (1 worker CWEM). 

Etymology. — This new species is named 
in honor of Robert Anderson, who col- 
lected these specimens as well as thou- 
sands of other interesting specimens. 

Discussion. — This species would key to C. 
olitor Forel in Kempf (1964), found in Brazil 
and Argentina. Cyphomyrmex under soni is 
somewhat larger (HL of C. olitor 0.64, HW 
0.56 from Kempf, 1964), has fewer mandib- 
ular teeth (7-8 in C. olitor), the midpronotal 
tubercle is approximately the same size as 
the lateral tubercles (midpronotal tubercle 
much smaller than lateral tubercles in C. 
olitor), and the propodeal spines are well 
developed (poorly developed in C. olitor). 
The mandibular teeth are worn and par- 
tially hidden by the clypeus in the holotype 
and badly worn in the mandible of the other 
specimen, but this species appears to be the 
only one of the strigatus species complex 
with six mandibular teeth (and as the 
members of the rimosus species complex 

Volume 19, Number 1, 2010 



Anterior mesonotal tubercle 
Propodeal spine 


1 mm 


Figs 1-6. Cyphomyrmex andersoni holotype worker: 1, side view. 2, mesosoma as seen from above. 3, head and 
mandible, frontal views. 4, top of pronotum as seen from dorsoposterior view. 5, right posterior femur and tibia 
posterior aspect. 6, antennal scape. 

all have five mandibular, it is apparently the 
only species in Cyphomyrmex with six teeth). 
Biology. — The two specimens were col- 
lected in montane hardwood leaf litter at 
1100-1200 m elevation and in wet montane 
forest litter. 

Cyphomyrmex snellingi new species 

(Figs 7-12) 

Diagnosis. — The worker is a small (total 
length 2 mm) ferrugineous red specimen. 
The mandibles have seven teeth, the spini- 


Journal of Hymenoptera Research: Festschrift Honoring Roy Snelling 

form paraclypeal teeth on the clypeus are 
markedly well developed; the frontal lobes 
do not reach the inner borders of the eyes 
(frontal view of head). The frontovertexal 
comers form auricle-like structures; the 
scape is short, and does not reach the 
posterior margin of the scrobe. The mid 
pronotal process is angulate, the lateral 
pronotal tubercles are poorly developed. 
The anterior mesonotal tubercles are con- 
ical and posterior mesonotal tubercles 
approximately the same size. The propo- 
deum is rounded posteriorly and without 
angles or spines. The subpetiolar tooth is 
well-developed and sharp, dorsally the 
petiole extends over the base of the anterior 
part of the postpetiole, which has two 
longitudinal elevated regions, the poster- 
ior margin of the postpetiole is nearly 
straight; the first opisthogastral* tergum is 
without ridges or processes; all femora are 
swollen ventrally, with carinae, the poster- 
ior femur has a well-developed ventral 

Erect hairs are sparse, present on the 
mandibles, apex of the scape, ventral 
surfaces of the legs, ventral and posterior 
surfaces of the gaster; appressed hairs are 
abundant on the dorsum of the first 
opisthogastral tergum. 

All surfaces dull, except the region along 
base of mandibular teeth which is smooth 
and shiny. 

Distribution. — Known only from the type 
locality in Panama. 

Description. — Worker measurements 
(mm): HL 0.71-0.74, HW 0.58, SL 0.48- 
0.50, EL 0.09-0.10, EW 0.08, WL 0.85-0.86. 
Indices: CI 78-81, SI 64-70, OI 82-86. 
Mandible with 7 teeth; spiniform paracly- 
peal teeth very well developed (length 
0.07 mm), frontal carinae relatively nar- 
rowly spaced, not reaching preocular ca- 
rina which forms mesiad margin of scrobe; 
eyes extending past sides of head, with 
about 20 ommatidia; scrobe greatly extend- 
ing posteriorly, forming auricle-like struc- 
tures; scapes not reaching posterior margin 
of scrobe; tubercles on pronotum poorly 

developed; anterior and posterior mesono- 
tal tubercles moderately well developed 
and approximately same size, anterior 
tubercle with slighter broader base; dorso- 
propodeum shorter that posteropropo- 
deum, propodeum without spines or an- 
gles; subpetiolar tooth sharp and well 
developed, petiole with two distinct, lon- 
gitudinal lateral lobes, dorsum of posterior 
face extending over anterior part of post- 
petiole; postpetiole with longitudinal de- 
pressed region in dorsum of node, outlined 
by two elongated elevated areas; dorsal 
surface of gaster flat, bordered laterally by 
slightly elevated longitudinal areas; all 
femora with carinae along ventral posterior 
border, that on posterior femur more 
developed and forming lamina. 

Few erect hairs on mandibles, antecly- 
peus and frontal lobes, remainder of hairs 
simple and appressed, located mostly on 
head and especially gaster. 

All surfaces except mandiblular teeth 
and anterior edge of clypeus dull. 

Type series. — Holotype worker (MCZC), 1 
paratype worker (CWEM), Panama, Cerro 
Campana, 950 m, 5-vi-1995, R. Anderson 

Etymology. — Named in honor of the 
memory of Roy Snelling, recalling a pleas- 
ant visit to the Los Angeles County 
Museum of Natural History in May of 
2007 where we spent time with Roy, 
Gordon Snelling, Brian Brown, and Wei- 
ping Xie. 

Discussion. — Cyphomyrmex snellingi would 
key to C. faunulus in Kempf s key (1964). It 
can be easily distinguished as the anterior 
mesonotal tubercle is relatively small, as 
compared to the greatly enlarged anterior 
mesonotal tubercle of C. faunulus (Fig. 13). 
Additionally, the posterodorsal edge of the 
petiole of C. faunulus does not extend over 
the anterior face of the petiole as it does in C. 
snellingi. Cyphomyrmex faunulus also lacks 
the erect hairs on the frontal lobes. 
Although it would key to C. faunulus, the 
two species do not appear to be morpholog- 
ically similar. 

Volume 19, Number 1, 2010 



Auriculate frontovertexal 


^ carina 


Cyphomyrmex faunulus 

Figs 7-12. Cyphomyrmex snellingi holotype worker: 7, side view. 8, top view of mesosoma (based in part on the 
paratype). 9, petiole as seen from above, postpetiole as seen in anterior view. 10, postpetiole as seen from above. 
11, Head. 12, left femur as seen from posterior view. Fig. 13, Cyphomyrmex faunulus mesosoma (Reserva Ducke, 
near Manaus, Amazonas, Brasil, LACM). 

Biology. — The type series was collected 
in a leaf litter extraction from a wet 
montane habitat. 


The genus Cyphomyrmex is divided into 
two species complexes, the rimosus com- 
plex and the strigatus species complex. The 

strigatus complex uses only mycelium 
cultivars and is probably plesiomorphic 
and paraphyletic to the rimosus complex 
(Schultz et al. 2002). 

Cyphomyrmex has two centers of species 
richness: the rimosus group at about 10° 
north (Mayhe-Nunes and Jaffe 1998), 
whereas the majority of the species of the 


Journal of Hymenoptera Research: Festschrift Honoring Roy Snelling 

strigatus group is restricted to 20°and 30° 
south (Sanhudo et al. 2007). The strigatus 
group also lacks species with wide dis- 
tributions (Mayhe-Nunes and Jaffe 1998) as 
is found in the rimosus complex. 

Apparently no new species of the striga- 
tus group have been described since 
Kempf's revision (1964), although the 
recently described C. muelleri shows simi- 
larities to the strigatus species complex, and 
along with C. longiscapus, C. costatus and C. 
wheeleri may be related (Schultz et al. 2002). 
A similar new genus Mycetagroicus, with 
three new species has been recently de- 
scribed (Brandao and Mayhe-Nunes 2001). 


Anteclypeus (= ''apron''): the anterior 
portion of the clypeus attached to the 
labrum. Dorsopropodeum: the dorsal sur- 
face of the propodeum. 

Frontovertexal comer: the posterolateral 
angle between frons and vertex. 

Notopropodeal groove: (= "metanotal 
groove") a transverse groove on the noto- 
propodeal fusion. 

Notopropodeal fusion: In workers, the 
tergal fusion of the thoracic notum and the 

Opisthogaster (adj. opisthogastral): 
(= "gaster") Abd IV to pygidium. 

Paraclypeal teeth: (="parafrontal teeth" 
- Kempf 1964, 1965), referring to the anterior 
teeth-like processes on the clypeus. 


We would like to thank Roy Snelling and Weiping 
Xie for the loan of specimens of Cyphomyrmex faunulus. 
Two anonymous reviewers provided important com- 
ments. The research was supported by a grant from 
the National Geographic Society to Robert Anderson 
and the Ernst Mayr Fund of the Museum of 
Comparative Zoology. 


Bolton, B., G. Alpert, P. Ward, and P. Naskrecki. 2007. 
Bolton's Catalog of Ants of the World: 1758-2005. 
Harvard University Press, available on CD. 

Brandao, C. and A. Mayhe-Nunes. 2001. A new 
fungus-growing ant genus, Mycetagroicus gen. 
With the description of three new species and 
comments on the monophyly of the Attini 
(Hymenoptera: Formicidae). Sociobiology 38: 

Fernandez, F. and E. Palacio. 1995. Hormigas de 
Colombia IV: Nuevos registros de generos y 
especies. Caldasia 17: 587-596. 

Kempf, W. W. 1964. A revision of the Neotropical ants 
of the genus Cyphomyrmex Mayr. Part I. Group of 
strigatus Mayr (Hym. Formicidae). Studia Entomo- 
logica 7: 1^4. 

. 1965. A revision of the Neotropical fungus- 
growing ants of the genus Cyphomyrmex Mayr. 
Part II: group of rimosus (Spinola) (Hym. For- 
micidae). Studia Entomologica 8: 161-200. 

Mayhe-Nunes, A. and K. Jaffe. 1998. On the biogeog- 
raphy of Attini (Hymenoptera: Formicidae). 
Ecotropicos 11: 45-54. 

Sanhudo, C, A. Mayhe-Nunes, and C. Brandao. 2007. 
Quern sao as Cyphomyrmex (Myrmicinae: Attini). 
Biologico 69: 433-434. 

Schultz, T. and S. Brady. 2008. Major evolution- 
ary transitions in ant agriculture. Proceedings 
of the National Academy of Sciences 105: 5435- 

, S. Solomon, U. Mueller, P. Villesen, J. 

Boomsma, R. Adams, and B. Norden. 2002. 
Cryptic speciation in the fungus-growing ants 
Cyphomyrmex longiscapus Weber and Cyphomyr- 
mex muelleri Schultz and Solomon, new species 
(Formicidae, Attini). Insectes Sociaux 49: 331- 

Serna, F. and W. Mackay. 2010. A descriptive 
morphology of the ant genus Procryptocerus 
(Hymenoptera: Formicidae). Journal of Insect 
Science (in press). 

Snelling, R. R. and J. T. Longino. 1992. Revisionary 
notes on the fungus-growing ants of the genus 
Cyphomyrmex, rimosus group (Hymenoptera: For- 
micidae: Attini). Pp. 479-494 in: Quintero, D., and 
A. Aiello eds. Insects of Panama and Mesoamerica: 
selected studies. Oxford University Press, Oxford. 
692 pp. 

Vol. 19(1), 2010, pp. 51-65 

Taxonomic Contribution to the aurita Group of the Ant Genus Azteca 

(Formicidae: Dolichoderinae) 

Roberto J. Guerrero, Jacques H. C. Delabie, and Alain Dejean 

(RJG) Grupo de Investigacion en Insectos Neotropicales, ESfTROPIC, Universidad del Magdalena, 

Carrera 32 # 22-08, San Pedro Alejandrino, Santa Marta, Magdalena, Colombia; Current address: 

Programa de Doctorado en Zoologia. Instituto de Zoologia Tropical. Facultad de Ciencias. 

Universidad Central de Venezuela. Avenida Los Chaguaramos, Caracas, Venezuela; 

(JHCD) Laboratorio de Mirmecologia, Convenio UESC-CEPEC, Centro de Pesquisas do Cacau, 

CEPLAC, Caixa Postal 7, 456000-000 Itabuna-BA, Brazil; 

(AD) Ecologie des Forets de Guyane (UMR-CNRS 8172), Campus Agronomique, 97379 KOUROU 

cedex, France; 

Abstract. — We describe five new species in the aurita group of the genus Azteca: Azteca andreae 
sp. n. (French Guiana), Azteca diabolica sp. n. (Panama), Azteca laurae sp. n. (Brazil), Azteca 
linamariae sp. n. (Brazil and Colombia) and Azteca snellingi sp. n. (Panama). Four of these new 
species are based on gynes, while the last is based only on the worker caste. All of them bear the 
aurita group characteristics. The second taxon is remarkable, as it differs from all of the other 
members of the group in the exaggerated, horn-like extensions of the posterolateral vertex margins. 
Azteca snellingi sp. n. is named in honor of our colleague, Roy Snelling, in tribute to his life-long 
contribution to knowledge of the world of Hymenoptera. A key to all known species of the aurita 
group, based on gynes, is provided. We report also for the first time an Intercast case for the genus 
Azteca, based on an Azteca schimperi specimen. 

Resumen. — Se describen cinco nuevas especies de hormigas del grupo aurita del genero Azteca: 
Azteca andreae (Guyana Francesa), Azteca diabolica (Panama), Azteca laurae (Brasil), Azteca 
linamariae y Azteca snellingi (Colombia y Brasil). Cuatro de las especies son descritas basadas en 
hembras, enguanto la ultima basada en la casta obrera. Todas poseen las caracteristicas del grupo 
aurita. El segiindo taxon es el mas particiilar; es facil diferenciar esta especie de las otras ya que 
presenta los margenes posterolaterales del vertex exageradamente extendidos similares a unos 
cuemos. Azteca snellingi sp. n. es nombrada en honor a nuestro colega Roy Snelling quien consagro 
su vida al conocimiento de los himenopteros. Se presenta una clave actualizada basada en las 
hembras de las especies del grupo aurita. Reportamos tambien por primera vez un caso de 
intercasta para el genero Azteca, con un especimen en la especie Azteca schimperi. 

Key words. — Ants, Azteca, identification keys, taxonomy 

Palabras claves. — Hormigas, Azteca, claves para identificacion, taxonomia 

Azteca Forel is a genus of doHcoderine carried out and only isolated studies have 
ant whose species-level taxonomy is still allovs^ed some species to be identified 
imclear. The only revision of the entire (Longino 1989, 1991a, 1991b, 1996). Re- 
genus was conducted in the late nineteenth cently, Longino (2007) reviewed the Costa 
century (Emery 1893). Since then, no Rican fauna and included a definition and 
systematic review of the genus has been global revision of the aurita group. 


Journal of Hymenoptera Research: Festschrift Honoring Roy Snelling 

The aiirita group of Azteca is monophy- 
letic and characterized by: palpal formula 
4,3; middle and hind tibia lacking an apical 
spur; an extremely convex anteromedial 
border of the clypeus that extends well 
beyond anterolateral clypeal lobes; HLB/ 
HLA > 1.04 (Longino 2007, except latter 
trait discussed here; for further description 
of the measurements, see Materials and 
Methods). Gynes are generally small, simi- 
lar in size to major workers. The integu- 
ment is extremely smooth and shiny, glass- 
like, with an extremely dilute appressed 
pubescence (the pilosity, when present, is 
characterized by a stubble of short, stiff, 
fully erect setae). The petiole is bluntly 
subpyramidal to bilobed, never flat and 
scale-like (Longino 2007). Worker charac- 
ters are provided in the group diagnosis by 
Longino (2007). The characteristics of 
gynes in this group suggest a syndrome 
of social parasitism (Holldobler and Wil- 
son 1990). 

In this paper, we describe five new 
species of the aiirita group: Azteca andreae 
(French Guiana), Azteca diabolica (Panama), 
Azteca laurae (Brazil), Azteca linamariae 
(Colombia and Brazil) and Azteca snellingi 
(Panama). The A. diabolica specimens 
(gynes only) were collected during the 
IBISCA international project through an 
intensive tridimensional sampling of ar- 
thropods in a Panamanian tropical rain 
forest (Basset et al. 2007), while the A. 
snellingi specimens were caught later in the 
same area. The other species were collected 
using Malaise traps (winged individuals) 
or manually from nests. An updated key 
derived from the one created by Longino 
(2007), based on the gynes of known 
species of the group, is provided. 


All measurements, indices and morpho- 
logical characters are the same used by 
Longino (2007) and were made using a 
Nikon SMZ645 stereomicroscope with an 
ocular micrometer at 80 X magnification. 

Measurements (all in millimeters): 

HLA: head length, full frontal view; 
perpendicular distance from the line tan- 
gent to the anterolateral clypeal lobes to the 
line tangent to the most extreme posterior 
of the vertex lobes. This measure was 
chosen because the anterolateral clypeal 
lobes are always visible, while the anterior- 
most extent of the medial lobe may be 
obscured by the closed mandibles. 

HLB: medial head length; this is the same 
as HLA except that it is measured from the 
anteromedian rather than anterolateral lobe 
of clypeus. This measurement is important 
for the A. aurita group, where the lateral 
lobes are not well defined and the median 
lobe protrudes to a great extend. For most 
Azteca, HLA and HLB are very similar. 

HW: head width; full frontal view, max- 
imum width of head capsule above the eyes. 

SL: scape length; length of the scape 
shaft from the apex to basal flange, not 
including the basal condyle and neck. 

EL: eye length; maximum length of the eye. 

OCW: width of the median ocellus. 


CI: cephalic index; 100*HW/HLA. 

SI: scape index; 100*SL/HLA. 

MTSC: number of metatibial setae; with 
the tibia seen from the anterior, such that 
the outer (dorsal) margin is in profile, and 
the number of erect to suberect setae 
(distinct from any underlying pubescence) 
are seen projecting from the outer margin. 

Photographs were taken using a Nikon 
SMZ 1500 stereomicroscope at 40 X and 
digital camera SIGHT DS - Fi 1. The images 
were fully-focused montage images cre- 
ated with the Combine version Z5 software 
package and edited using COREL PHOTO- 
PAINT X3 version 13. 

Types will be deposited in the fol- 
low collections: California Academy of 
Sciences, San Francisco, California, USA 
(CASC); Laboratorio de Mirmecologia do 

Volume 19, Number 1, 2010 


Centro de Pesquisas do Cacau (CPDC), 
Comissao do Piano da Lavoura Cacaueira, 
Ilheus, Bahia, Brazil; Insect Collection, 
Instituto Humboldt, Claustro San Agustin, 
Villa de Leyva, Boyaca, Colombia (lAvH); 
Instituto de Ciencias Naturales, Universi- 
dad Nacional, Bogota D.C., Colombia 
(ICN-MHN); collection of John T. Longino, 
Evergreen State College, Olympia, Wash- 
ington, USA QTLC); Los Angeles County 
Museum of Natural History, Los Angeles, 
California, USA (LACM); Museu de Zool- 
ogia, Universidade de Sao Paulo (MZUSP), 
Brazil; and Royal Belgian Institute of 
Natural Sciences (RBINS). 


Azteca andreae Guerrero, Delabie & Dejean. New 

species. French Guiana. 
A. aurita Emery 1893. Panama to Amazonian 

= silvae Forel 1899. Synonymy, Longino 


A. diabolica Guerrero, Delabie & Dejean. New 

species. Panama. 
A. lallemandi Forel 1899. Panama, Colombia, 

eastern Brazil. 
= pruinosa Mann 1916. Synonymy Longino 

A. lanuginosa Emery 1893. Southern Brazil. 
A. laurae Guerrero, Delabie & Dejean. New 

species. Western Amazonian Brazil. 
Azteca linamariae Guerrero, Delabie & Dejean. 

New species. Amazonian Colombia and 

Western Amazonian Brazil. 
A. nanogyna Longino 2007. Costa Rica. 
A. pilosula Forel 1899. Costa Rica, Panama. 
= lacrymosa Forel 1899. Synonymy, Longino 

A. schimperi Emery 1893. Mexico to Argentina. 
= A. fiebrigi Forel 1909. Synonymy, Longino 

= A. clariceps Santschi 1933. Synonymy, Longino 

= A. pallida Stitz 1937. Synonymy, Longino 

A. snellingi Guerrero, Delabie & Dejean. New 

species. Panama 


1. Orange head and orange or light brown body; long scape, SI > 70 2 

Uniform brown color; short scape, SI < 70 7 

2. Erect pilosity absent on dorsum of the head, lateral margins of the mesosoma, petiole, 

and gaster 3 

Short, erect pilosity present on the dorsum of the head and mesosoma, petiole, and 
gaster 6 

3. Very pronounced vertex lobes appearing as elongate, horn-like projections (Fig. 6); a 

seemingly wide, U-shaped occipital margin diabolica n. sp. 

Angulate vertex lobes never forming horn-like projections 4 

4. Head relatively broad CI > 99 linamariae n. sp. 

Head relatively narrow CI < 99 5 

5. Sides of the head flat and sub-paraUel, only weakly diverging posteriorly; eyes more 

or less at mid-length of the head, HW < 1.30 aurita 

Sides of the head flat and not sub-parallel, strongly diverging posteriorly; eyes anterior 
to the mid-length of the head, HW > 1.30 laurae n. sp. 

6. Dense, short, erect pilosity on scape and tibiae; head relatively narrow (CI < 97) ... pilosula 
Scape and tibiae lacking erect pilosity; head relatively broad (CI > 97) .... lallemandi 

7. Gastral dorsum lacking erect setae; HLA > 1.35 8 

Gastral dorsum with erect setae; HLA < 1.35 9 

8. Head, scapes, mesosoma, legs and petiole with erect hairs; scapes relatively long, SL > 

0.90 schimperi 


Journal of Hymenoptera Research: Festschrift Honoring Roy Snelling 

Head, scapes, mesosoma, legs and petiole devoid of any erect hairs; scapes relatively 

short, SL < 0.90 andreae n. sp. 

9. HLA about 1.3mm lanuginosa 

HLA about 0.86mm nanogyna 


Azteca andreae n. sp. Guerrero, Delabie & 
Dejean (Figs 1 & 2) 

Holotype (gyne): FRENCH GUIANA, Sin- 
namary, 5°22'39"N 52°57'35"W, Carton nest in 
Cecropia sp tree, 24 Jul 2008, (A. Dejean, P-J 
Male, S. Groc and J.H.C. Delabie) [CPDCj; 
paratypes: 7 gynes, same locality, [CASC,CPDC, 

Measurements of holotype: HLA 1.42, 
HLB 1.48, HW 1.08, AHW 0.72, SL 0.82, EL 
0.34, OCW 0.08, CI 76, SI 58, MTSC 0. 

Measurements of paratypes (N= 7): HLA 
1.38-1.42, HLB 1.46-1.52, HW 1.06-1.10, 
AHW 0.70-0.76, SL 0.80-0.86, EL 0.32-0.36, 
OCW 0.06-0.08, CI 75-80, SI 57-61, MTSC 0. 

Diagnosis. — Azteca andreae is a member of 
the A. aurita group w^ith the body com- 
pletely covered w^ith small, widely scat- 
tered pits (foveate surface) bearing a very 
short white hair; scapes short, SI 57-61. 

Gyne characters. — Head: Palpal formula 
4,3. Dorsal surface of mandibles smooth 
and shiny; from an oblique angle from 
above, weak longitudinal waves can be 
observed that are not visible in full dorsal 
view, and with scattered small and widely- 
spaced holes, each with a short hair on the 
basal surface, with long hairs behind 
masticatory margin and anterior half of 
the outer margin of the mandibles; masti- 
catory margin armed with strong apical 
and blunt tooth, sub-apical tooth, followed 
by very rounded teeth extending to the 
basal margin. Clypeal plate without con- 
spicuous pilosity; medial clypeal lobe 
strongly convex and protruding, extending 
well beyond the lateral clypeal lobes. Head 
sub-rectangular, longer than wide; cephalic 
capsule in lateral view strongly convex in 

the front; posterior margin with blunt 
angulations, deeply excavated in the mid- 
dle of the V-shape. Scape not reaching 
posterior margin of cephalic comer; funi- 
culus covered with dense, long and ap- 
pressed pilosity. 

Mesosoma: Smooth and shiny, without 
conspicuous pilosity. Dorsal surface of 
propodeum much longer than posterior 
surface, the latter with a short projection at 
the base, tube-shaped and facing poste- 
riorly. Edge of metapleural gland orifice 
bears long, golden hairs. Middle and hind 
tibiae lacking apical spur. 

Metasoma: Petiolar node strongly sub- 
triangular with rounded apex; anterior face 
of petiole excavated, posterior face nearly 
twice the length of anterior face; petiolar 
posteroventral lobe weakly convex behind, 
straight in the anterior half. Tergites and 
stemites of the gaster shiny. 

Head, mesosoma, petiole and legs dark, 
reddish brown, gaster yellowish brown 
with some darker areas. Body shiny. 

Worker characters. — Measurements (N= 
5): HLA 1.18-1.36, HLB 1.20-1.46, HW 
1.20-1.36, SL 0.84-0.96, EL 0.20-0.26, CI 
100-102, SI 67-75. 

Palpal formula 4,3. Middle and hind 
tibiae lacking apical spur. Same combina- 
tion of characters as Azteca schimperi in 
Longino (2007). Minor workers with tube- 
like propodeal spiracles projecting out- 
wards, unlike those of A. schimperi. 

Male characters. — Measurements (N= 4): 
HLA 0.70-0.72, HW 0.67-0.68, SL 0.11-0.12, 
EL 0.30, CI 94-97, SI 16-17. 

Head: Mandibles sub-triangular, outer 
edge twice as long as inner edge, mastica- 
tory margin unarmed, with only a small 
projection in the middle of margins and a 
sharp apical tooth; a basal tooth differenti- 

Volume 19, Number 1, 2010 


Fig. 1. Paratype female of Azteca andreae (French 
Guiana; CASENT 0179545). A). Full frontal view of the 
head of A. andreae. B). Lateral view of the body of 
A. and\'eae. 

ates masticatory and basal margins. Cly- 
peal plate strongly convex, as medial 
clypeal lobe extends well beyond small, 
lateral clypeal lobes; surface smooth with 
small foveae near anterior ridge of clypeal 
plate. Scapes small, trapezoidal, thin at 
base and wider distally; pedicel small, 
nearly equal to maxiraum scape width; 
second funicular segment roughly twice as 
long as scape; remaining funiculus little 
longer than scape; scape with smooth 
surface and a few scattered hairs; funicular 
segments 3-11 with long, dense and 
appressed pilosity, surface densely punc- 
tate and dull. Eyes large, located near 
anterior half of the cephalic capsule, 
separated from mandible insertion by less 
than 0.1 mm; eyes break the plane on side 
of head. Lateral oceUi protruding slightly 
above vertex in dorsal view. Margins of the 

Fig. 2. Azteca andreae male from French Guiana 
(CASENT 0179546). A). Full frontal view of the head 
of A. andrea. B). Lateral view of the body of A. andreae. 

cephalic capsule, above the eyes, slightly 
convex; posterolateral comers rounded. 

Mesosoma: Parapsidal furrows weakly 
developed. Anepistemum and katepister- 
num divided by a deep mesopleural 
groove. Anterior and posterior surfaces of 
propodeum undifferentiated. Propodeal 
spiracles visibly protruding. 

Metasoma: Petiolar node rectangular 
with rounded corners and straight dorsal 
face; anterior surface much longer than 
posterior surface; petiolar node partly 
fused to gaster. Tergites and sternites 
smooth and shiny. Pygostyle thin and long, 
with distal half curved downwards and 
squared at the tip, smooth and shiny. 

Body dark brown, shiny, slightly punc- 
tate in some areas, covered with very 
sparse decumbent, long white hairs. 

Etymology. — Azteca andreae is named in 
honor of Andrea Dejean, the third author's 
wife, in acknowledgment of her consider- 
able editorial help with myrmecology 
papers in English. 


Journal of Hymenoptera Research: Festschrift Honoring Roy Snelling 

Range. — French Guiana. 

Natural History. — This species constructs 
large, conspicuous globular carton nests at 
the base of the stem or near the crown of 
Cecropia trees (Fig. 4). Alain Dejean has 
observed and collected several A. andreae 
nests from trees in French Guiana, the vast 
majority of which were Cecropia obtusa 
Trecul. A. andreae can also nest on C. 
pahnata Willd, which also often shelters 
Azteca alfari or A. ovaticeps. The nests 
observed by A. Dejean always were in 
pioneer vegetation highly-altered through 
human activity. 

All nests collected contained colonies with 
many workers, dozens of winged females 
and few males; for example, type series of 
this species came from a colony with 
hundreds of workers, hundreds of winged 
females (only 55 were collected), five male, 
several brood and pupae, although no 
queen was observed. Other collections made 
by A. Dejean produced colonies with some 
physogastric females (Fig 5). 

Comments. — A. andreae is close to A. 
schimperi but can be distinguished by 
morphological characters of gynes and 
males; the major workers of the two species 
are indistinguishable. A. andreae gynes 
conspicuously lack pilosity over the entire 
body, while A. schimperi gynes are densely 
covered with erect hairs on the head and 
mesosoma. The anteromedial portion of 
the clypeus protrudes more in A. andreae 
than in A. schimperi; the masticatory mar- 
gin of the mandibles of A. andreae is armed 
with blunt teeth, while A. schimperi has 
sharp teeth. The scapes of A. andreae are 
significantly shorter than those of A. 
schimperi (0.80-0.86 vs. 0.94-1.01 mm, re- 
spectively); the posterior margin of the 
head is much more excavated in A. andreae 
than in A. schimperi, and the posterolateral 
corners are also more angular in the first 
species; another important character 
amongst the females of both species is the 
color: A. andreae females are a dark reddish 
brown, while A. schimperi females are 
uniformly brown. 

The males of both species are very 
similar in color, type and distribution of 
hairs, and wing venation; however, when 
we examined the male genitalia of both 
species, we found that there are conspic- 
uous differences. The pygostyle on A. 
andreae is smooth and shiny, thin, long, 
and with the distal half curved downwards 
and squared at the tip, while those on A. 
schimperi are short, thick and with a 
rounded tip (Fig. 3). 

The major workers of both species are 
indistinguishable, although the workers of 
A. andreae have a relatively smaller head; 
however, the ranges of both species overlap. 
The minor workers are also undifferen- 
tiated, but A. andreae have thin, protruding 
tube-shaped propodeal spiracles, whereas 
A. schimperi are open at the propodeum. 

The females of A. lanuginosa, A. schimperi 
and A. andreae are related to each other and 
show a clear gradation in the quantity, 
distribution and type of body hair. The first 
species is the only one of them with an 
abundant, erect pilosity all over the body, 
including the gaster (Longino 2007); it 
becomes sparse in A. schimperi and disap- 
pears altogether in A. andreae (in the latter 
two species, the gaster is devoid of any hair). 

Azteca diaholica n. sp. Guerrero, 
Delabie & Dejean (Fig. 6) 

Holotype (gyne): PANAMA, San Lo- 
renzo Forest, IBISCA project, 9°16'47.58"N, 
79°58'29.94'W, Flight-interception trap in the 
canopy, 3-13 Ago 2004 (M. Rapp) [CPDCl; 
paratypes: 1 gyne, same location, Fogging #FO- 
C3-6C, 13 Oct. 2004 (J. Bail) [RBINSl; 1 gyne, same 
location. Fogging FOG-Rl-5, 20 Oct. 2003 Q. 
Schmidl) [ICNJ. 2 gynes, same location. Fogging 
#J-2, 17 Oct. 2003 G- Schmidl) [CPDC, MZUSP]. 

Measurements of Holotype: HLA 1.48, 
HLB 1.56, HW 1.12, AHW 0.72, SL 1.24, EL 
0.30, OCW 0.06, CI 76, SI 84, MTSC 0. 

Measurements of Paratypes (N= 2): HLA 
1.60-1.62, HLB 1.64-1.74, HW 1.20-1.24, 
AHW 0.78-0.80, SL 1.34-1.36, EL 0.30-0.32, 
OCW 0.06, CI 75-77, SI 84-85, MTSC 0. 

Volume 19, Number 1, 2010 


Fig. 3. Genitalia of male A. andreae and A. schimperi. 
The arrow indicates the pygostyle. 

Diagnosis. — Azteca diabolica is a member 
of the A. aurita group with a deep and 
smoothly rounded excavation at the pos- 
terior vertex margin extending to the 
corners that form posteriorly-projecting 
rounded horns. Mesosoma smooth, shiny 
and hairless. Propodeal spiracles protrud- 
ing. Gastral tergum and sternum with 
hairless, polished surface. 

Gyne characters. — Head: Palpal formula 
is 4,3. Dorsal and ventral surfaces of head 
hairless. Dorsal surface of mandibles 
mostly smooth and shiny, with fine lon- 
gitudinal striae near masticatory margin; 
masticatory margin armed with five teeth 
and two denticles, with no angle or tooth 
separating it from basal margin; basal 
margin slightly serrated; surface of man- 
dibles with scattered, sub-decumbent long 
hairs. Clypeal plate with sub-decumbent. 

Fig. 4. A. andreae nests built in Cecropia spp. trees. 

Sparse pilosity; medial clypeal lobe 
strongly convex and protruding, extending 
well beyond lateral clypeal lobes. Head 
almost rectangular, somewhat swollen be- 
tween ocellar region and compound eye; 
posterior margin highly angular, horn-like 
laterally, deeply excavate medially. When 
laid back, scape reaches prolongations of 
vertex at apex of posterolateral projection; 
scape and funiculus provided with abun- 
dant, nearly erect pilosity. 

Mesosoma: Smooth and shiny, without 
appressed hairs. Middle and hind tibiae 
lacking apical spur. Dorsal surface of 
propodeum shorter than posterior surface; 
propodeal spiracles protruding. 

Metasoma: Petiolar node bluntly trian- 
gular, posterior surface straight, twice as 
long as anterior surface; posteroventral 
petiolar lobe very low, very shallowly 
convex, ending posteriorly in a somewhat 


Journal of Hymenoptera Research: Festschrift Honoring Roy Snelling 


Fig. 5. Physogastric A. andreae queens from French 
Guiana. A. Physogastric A. andreae queen attended by 
workers. B. Preserved physogastric A. andreae queen. 

Fig. 6. Head of Azteca diabolica (Paratype fen\ale, 
Panama; ICN-022611), full frontal view. 

mandibles and legs. Some species in the 
aurita group have a strongly pronounced 
lateral vertex margin, but none is as 
pronounced as in Azteca diabolica. 

Azteca laurae n. sp. Guerrero, 
Delabie & Dejean (Fig. 7) 

abrupt shelf. Tergites and stemites hairless, 
smooth and shiny. 

Body reddish brown, surface smooth 
and reflective. 

Worker and male. — Unknown 

Etymology. — The name refers to the form 
of the head of the gyne which suggests 
popular representations of the Devil. 

Range. — Panama. 

Natural History. — The five specimens 
were collected from the rain forest canopy, 
first with a flight-intercept trap, later by 
applying a chemical treatment to vegeta- 
tion. Three gynes were collected by fog- 
ging; one from a tree where both Azteca 
sp.2 chartifex group and Azteca instabilis (Fr. 
Smith) occurred, the two others from a tree 
with Azteca sp.2 chartifex group. This 
suggests that A. diabolica may be a social 
parasite, in particular of carton-nesting 
species of the chartifex group. 

Comments. — The gyne differs from those 
of other species in the A. aurita group in 
being almost hairless, having only sparse, 
short and decumbent hairs on the clypeus. 

Holotype (gyne): BRAZIL, Rondonia, Par- 
que Estadual Guajara Mirim, 10°19'17"S, 
64°33'47"W, #5256, Malaise trap, 02 Mar. 1998 
a.R.M. Santos) [CPDCl. 

Measurements of Holotype: HLA 1.56, 
HLB 1.62, HW 1.46, AHW 0.90, SL 1.24, EL 
0.36, OCW 0.06, CI 94, SI 79, MTSC 0. 

Diagnosis. — Azteca laurae is a member of 
the A. aurita group with an inverted, cone- 
like (cuneiform) head whose sides strongly 
diverge from the lateral region; surface of 
head smooth and shiny, with very thin 
and weak punctations visible laterally by 
tilting the specimen (dorsal-oblique view), 
although these one more noticeably visible 
in the ocellar region (full frontal view). 

Gyne characters. — Head: Palpal formula 
4,3. Dorsal and ventral surface devoid of 
any type of hair, although very short 
and sparse hair covers a small portion of 
the genae. Dorsal surface of mandibles 
completely smooth and shiny, clearly 
seen in spaces between the sparse, long 
hairs. Masticatory margin of mandibles 
armed with four teeth and two denticles. 

\ OLL-ME 19, Xl-mber 1, 2010 


Fig. 7. Holot\-p€ female of Azteca laiirae (Brazil; 
CPDC-5256). A). Full frontal \-iew of the head of .4. 
laurae. B). Lateral \iew of the body of A. laurae. 

Clypeal plate covered ^vith abundant short, 
nearly erect pilosit}'; medial ch^peal lobe 
strongly convex, projecting outwards, ^vith 
hairless anterior ridge extending well 
beyond lateral ch^eal lobes. Vertex with 
prominent, rounded comers deeply exca- 
vated in middle in U-shape with gently- 
rounded tips. Ocelli in a loose clump, 
forming a dark stain in dorsoposterior 
region. Scapes barely reaching extensions 
of vertex; scapes and funiculus covered 
with abundant, short, sub-decumbent pi- 
losit}', shorter than maximum width of 

Mesosoma: Smooth and shiny, ^vith no 
conspicuous hairs, only a few ver}- short, 
appressed hairs becoming sparser to- 
wards katepistemal and propodeal region. 
Dorsal side of propodeum almost equal in 
length to posterior surface, nearly undiffer- 
entiated due to absence of a defined 
boundar}-; propodeal spiracles weakly pro- 
truding. Middle and hind tibiae lacking 

Metasoma: Petiolar node triangular, 
sloping gently posteriorly; posterior mar- 
gin almost twice as long as anterior; 
petiolar lobe ^veakly convex behind; ven- 
tral surface roughly parallel to dorsal 
surface. Gaster hairless, surface polished 
and ver}' shinv. 

Body reddish brown, surface smooth 
and reflective. 

y^orker and male. — Unkno^vn 

Eh/mology. — The name is in honor of 
Laura Mariano Delabie, the second 
author's daughter. 

Range. — Western Amazonian Brazil. 

Xatural History. — The holot}'pe ^vas col- 
lected in the mature forest of the Parque 
Estadual Guajara Mirim, Rondonia, Brazil, 
using a Malaise trap. 

Comments. — The g}Tie of this species is 
closely similar to A. aiirita, differing in the 
amount and distribution of the hairs on the 
dorsum of the head and the scapes. A. 
laurae has ver}' fe^v short hairs on the 
genae, while the anterior part of the head 
of A. aurita is covered -with a uniform 
vestiture of short, dense, %vhite pilosit\\ 
The scapes of A. laurae are sparsely covered 
^vith short hairs, ^vhile those of A. aurita 
are densely covered by the same t}~pe 
of pilosit}' present in the dorsal region 
of the head. Another notable trait is the 
shape of the head, the sides of which 
are almost parallel in A. aurita (Fig. 8), 
^vhile in A. laurae they diverge posteriorly, 
resulting in a cuneiform-shaped head 
(Pig. 7); the vertexal margin in A. laurae 
is ^vider and slightly more concave than in 
A. aurita. 

Azteca linamariae n. sp. Guerrero, 
Delabie & Dejean (Pig. 9) 

Holotype (gyne): COLOMBIA, \'aupes, Esta- 
cion Biologica Mosiio-Itajura (Caparu), .Antigua 
Cabafia, 1^ 4'S, 69' 3'W, 60 m. Malaise trap, 18- 
27 mar 2003 (J. Pinzon), M.3610 - Insects of 
Colombia project [lAvFi]; paratype (gyne): 
BR.\ZIL, Rondonia, Paxque Estadual Guajara 
Mirim, 10^19'17"S, 64^33'471V, =5248, Malaise 
trap, 28 Jan. 1998 (J.R.M. Santos) [CPDC]. 


Journal of Hymexoptera Research: Festschrift Honoring Roy Snelling 

Fig. 8. Head of an Azteca aiirita female from 
Panama, full frontal view. 

Measurements of Holotype: HLA 1.86, 
HLB 2.0, HW 1.84, AHW 1.18, SL 1.48, EL 
0.40, OCW 0.14, CI 99, SI 80, MTSC 8. 

Measurements of Parat}^pe: HLA 1.82, 
HLB 1.86, HW 1.82, AHW 1.16, SL 1.46, EL 
0.44, OCW 0.14, CI 100, SI 80, MTSC 8. 

Diagiiosis. — Azteca linamariae is a mem- 
ber of the A. aurita group with the dorsal 
and ventral surfaces of the head, mesonotal 
dorsal region and gastral tergum and 
sternum covered with abundant, very thin, 
short, white, scale-like setae. It is the 
largest queen of any known species in the 
aurita group. 

Gyne characters. — Head: Palpal formula 
4,3. Ventral surface with abundant pilosity, 
as well as long, very closely spaced, erect 
hairs covering all head margins and back 
of foramen magnum. Mandibles smooth 
and shiny, with abundant decumbent 
pilosity, longer hairs toward masticatory 
margin. Clypeal plate covered with short, 
abundant pilosity; medial clypeal lobe 
strongly convex, but projected slightly 
toward the front, extending well beyond 
lateral clypeal lobes. Head nearly rectan- 
gular, slightly diverging laterally, flat in 
the ventral region; posterior margin deeply 
excavated, sharply angled with rounded 
comers. Scapes a significant distance from 
corners of vertex with nearly erect, 
short hairs approximately equal to half 
maximum width of scapes; funiculus cov- 

Fig. 9. Azteca linamariae (Holot\-pe female, Colom- 
bia; lAvH- M 3610). A). Full frontal \'iew of the head 
of A. linamariae. B). Lateral view of the body of 
A. linamariae. 

ered with abundant, appressed, sub-de- 
cumbent pilosity much shorter than that of 

Mesosoma: Smooth and opaque, with 
some sparse and scattered long hairs, 
mainly on promesonotal dorsum, mostly 
appressed below propodeal spiracle. Sides 
of propodeum shorter than posterior re- 
gion; propodeal spiracles projecting 
slightly outward. Middle and hind tibiae 
lack spurs. 

Metasoma: Petiolar node triangular, 
with front straight and nearly as long as 
posterior surface; posteroventral lobe often 
convex, inverted, and hump-like. Caster 
smooth and polished, without long hairs. 

Head reddish brown, mesosoma brown, 
gaster yellowish-brown with a highly 
polished and reflective surface. 

Y^orker and male. — Unknown 

Etymology. — This name is in honor of 
Lina Maria Ramos, the first author's wife, 
busv and active like an ant. 

\^OLL-ME 19, Number 1, 2010 


Range. — Amazonian Colombia and Bra- 

Natural Histoiy. — The holoU^pe was col- 
lected in the upland Am.azonian forest at 
the Mosiro-Itajura (Caparu) biological sta- 
tion in Vaupes, Colombia. This area is 
mainly covered by primary forest. The 
parat\^e was collected in a Malaise trap 
in the Par que Estadual Guajara Nlirim, 
Rondonia state, Brazil. 

Comments. — The g}TLes of this species are 
similar to those of A. pilosula, from which it 
is distinguished by hair characters. A. 
pilosiila has long, dense, white hairs on all 
sides of the head and on other regions of 
the body, while A. linamariae has very thin, 
short, white, scale-like setae, as well as 
some emerging long hairs (like those of A. 
pilosula) covering the head margins and the 
back of the foramen magnum and prome- 
sonotal dorsum, but not the dorsum of the 
head, lateral margins of the mesosoma, 
petiole or gaster. A. linammiae is the largest 
knowTi species in the group, and the g}Tie 
is much darker in color than that of A. 

The holotype and paratype differ only 
in the venation of the forewings. The 
forewings of both specimens have an r-rs 
cross-vein starting in the anteroinferior 
portion of the stigma; however, this 
cross-vein in the paratype is attached to 
Rsl and Rs2 base veins. The holotype r- 
rs cross-vein, on the other hand, is 
attached to the Rs short vein. The latter 
diverges in Rsl and Rs, posteriorly; 
moreover the paratype has \:wo cross- 
veins, 1 cu-a and 2cu-a, forming a small 
cell, while in the holot}^pe there is only a 
crossvein, 1 cu-a, and no small cell. 
Despite this difference in the pattern of 
venation of the forewing and the great 
distance (around 1.350 km) between the 
two capture sites, these t^vo specimens 
undoubtedly belong to the same species 
described above. Turther material collec- 
tion should confirm this identification 
and elucidate the small differences pre- 
sented here. 

Azteca snellingi n. sp. Guerrero, Delabie 
& Dejean (Tig. 10) 

Holotype (major worker): PANAMA, Colon, 
San Lorenzo Forest (SLPA), area metropoHtana, 
IBISCA project, 9=16.793'N, 79'- 58.499'W, man- 
ual collection in the canopy, 26 Feb 2008 (X. B. 
Espirito Santo & S. P. Ribeiro) [CPDC]; para- 
types: 6 major ^vo^ke^s, same data as for 
holot>T5e [1, CASC; 1, ICN; 1, JTLC; I, LACM; 
1, MZSP; 1, RBINSj. 

Measurements of Holotype: FTLA 1.56, 
HLB 1.66, ¥m 1.60, SL 1.30, EL 0.30, CI 103, 
SI 83. 

Measurements of Parat}3)e (N=6): HLA 
1.44-1.58, HLB 1.52-1.66, HW 1.44-1.60, SL 
1.18-1.32, EL 0.24-0.30, CI 99-101, SI 81-88. 

Diagnosis. — Azteca snellingi is a member 
of the A. aurita group with a large head, 
sHghtly wider than long, with margins 
strongly convex; metanotal groove wide 
and deep; workers have reddish brown 
head and dark brown body. 

'Worker characters. — Head: Palpal formula 
4,3. Mandibles completely flat, apical tooth 
much larger than anterior; dorsal surface 
with dense longitudinal sculpture; surface 
rough and opaque. Median clypeal lobe 
strongly convex, extending well beyond 
lateral clypedil lobes. Sides of head strongly 
cur\^ed, comers of posterolateral margins 
angled; posterior margin (vertex) strongly 
concave. Scapes cun^ed, not reaching pos- 
terolateral comers of head. 

Mesosoma: In lateral view, pronotum 
weakly convex or straight toward anterior, 
without a posterior face. Mesonotum 
strongly convex, rising well above prono- 
tum like a hump; globular in front, gently 
flattened posteriorly. Metanotal sulcus 
large and deep. In lateral view, propodeum 
roughly flat, dorsal face much larger that 
the posterior face. Middle and hind tibiae 
lacking spurs. 

Metasoma: Petiolar node large, with 
rounded end, sloped at front; posterior 
face nearly t^vice as large as the anterior 
face. Ventral lobe conspicuously uniformly 


Journal of Hymenoptera Research: Festschrift Honoring Roy Snelling 

1 mm 

Fig. 10. Azteca sneUingi (Paratype, worker major, 
Panama; CASENT 0179543). A). Full frontal view of 
the head of A. sneUingi. B). Lateral view of the body of 
A. sneUingi. 

Body entirely covered with fine, dense 
punctations. Pubescence thin and whitish, 
appressed, covering each tagma; some setae 
conspicuous near base of ventral lobe of 
petiole and stemites 2-4. Body with shiny, 
weakly reflective surface. Part of head, scape 
and mandibles dark reddish-brown. Poster- 
ior third of the head dark brown. Mesosoma, 
legs, petiole and gaster dark brown. 

Gyne and male. — Unknown 

Etymology. — This ant is named in honor 
of Roy Snelling, our colleague and friend, 
who contributed notably to the knowledge 
of Hymenoptera. 

Range. — Panama. 

Natural History. — Specimens of the type 
series were collected at different heights on 
the canopy in the San Lorenzo forest, 
Panama. Two were collected at 21m from 
a Pouteria caimito (Ruiz & Pav.) Radlk. 
(Sapotaceae) tree. Others were collected at 
20m and 23.7m height from Luehea seeman- 
nii Planch. & Triana. (Tiliaceae). 

Comments. — This species is close to A. 
aurita and A. pilosula. While workers of A. 
aurita and A. pilosula are undistinguishable 
(Longino 2007), those of A. sneUingi have a 
distinctly larger head, as long as it is wide. 
Posterior margin of the head with a deep 
concavity and continuous while the other 
two species has a V-shaped concavity. In 
general A. sneUingi is a species with work- 
ers (reproductive castes unknown) larger 
than A. aurita and A. pilosula workers (CI 
99-101 vs. A. aurita: CI 85-88, A. pilosula: CI 
89-90). Sides of head strongly convex in A. 
sneUingi, however, some A. pilosula workers 
(A. lacrimosa syntype and lectotype, syno- 
nymized by Longino 2007) have curved 
margins but not become as convex as in A. 
sneUingi. Azteca sneUingi mandibles, with 
the dorsal surface as those of Azteca aurita, 
but those of A. sneUingi are more opaque. 
Furthermore, the anterior region of the 
mesonotum is higher and more globular in 
A. sneUingi. Metanotal groove is wide and 
deep in A. sneUingi whereas other two 
species is absent or inconspicuous. 

Azteca sneUingi also differs from A. aurita 
and A. pilosula in the distribution and kind 
of the hairs on the body. A. sneUingi 
workers have no erect hairs in the petiolar 
node in contrast A. pilosula workers have 
erect hairs on the anterior face and apex of 
the petiole. Lateral margins of petiole 
lacking of any pilosity whereas A. pilosula 
has erect, conspicuous and scattered setae. 
A. sneUingi workers have no erect hairs on 
the posterior margin of the head whereas 
A. pilosula workers posterior margin of 
head with sparse, very short erect setae 
grading into white pubescence (Longino 
2007). A. sneUingi is reddish-brown and the 
body is entirely dark brown, which is 
notable in this species as no other known 
worker in the aurita group presents this 
coloration pattern. 

This species will key to couplet 2 in 
Longino's (2007) key to A. aurita group 
workers. The following modifications 
to the key will accommodate the new 

VoLL-ME 19, Xl-mber 1, 2010 

2. Head relatively broad (Q > 105); posterolateral margins of the \-ertex rounded and 

cordate, not bluntly angulate A. lallemandi 

Head relatively narro^v (CI < 106); posterolateral margins of the vertex bluntly 
angulate 3 

3. Head light orange bro\\n; mesosoma, legs, and gaster darker reddish-brown .... 

A. aurita, A. pilosiila 

Scape, mandibles and a part of the head dark reddish-browTi; posterior third of the 

head is dark bro^vn. Mesosoma, legs, petiole and gaster dark bro^vn 

A. snellingi n. sp. 

4. Pubescence dilute and tightl\" appressed; color usually bro^sn ^sith an orange head . . . 

A. schimperi 

Pubescence more abundant, giving a some^vhat wooly appearance; color all 
bro\sn A, laniigmosa 

Other relevant material relative to this 
group ^vas also studied: 

Azteca aurita Emer}- (Fig. 8): COLOMBIA, 
Putumayo, PXX La Paya, riparian forest, OTS, 
74=36'\V 320m, Malaise trap (M2440), 19 Sep-1 
Oct 2001 (R Cobete) - 1 g}ne [IAyU]. BK\ZIL, 
-Amazonas, Manaus, 12 sep 1962, = 3414, (K. 
Lenko) - 1 male, 3 minor ^vorkers [MZSP]; Mato 
Grosso, Sinop, 12^31 'S, 55'321V, Oct 1974, 
=12458, (M. Alvarenga) - 3 major workers 
[MZSP]; Mato Grosso, Vila Vera, Oct 1973, 
=10348 (M. .Vlvarenga) - 1 g^ne [MZSP]; Para, 
Belem, 12-19 Aug 1962 (K Lenko) - 1 dealate 
g}ne [MZSP]. VASASIA, Colon, San Lorenzo 
Forest, IBISCA project, 9^16.793'X, 79=58.499'\V, 
flight-interception trap in the canopy, 3-13 Ago 
2004 (M. Rapp) - 2 g}nes [CPDC]; same location, 
fogging (FO-R3-05C),' 13 Oct 2004 (J. Bail) - g\ne 
[CPDC]; same location, fogging (FOG-Rl-4),' 20 
Oct 2003 (J. Schmid) - 1 g^ne [CPDC]. PERU, 
Cusco, Pillcopata, 7 Dec 1974, =15, (J.A. Esca- 
lante) - 1 minor ^vorker, 2 major workers [MZSP]. 

Azteca lalleiJiandi Forel: BRAZIL, Bahia, nheus, 
Praia do Xorte, 27 Jun 2004 (J.H.C. Delabie) - 1 
g}ne [CPDC]; same location, 04 Dec 2004 (J.H.C. 
Delabie) - 3 g}nes [CPDC]; same location, 27 
Dec 1994 (l.C. Xasdmento; J.H.C. Delabie) - 5 
g^nes [CPDC]. PANAMA, San Lorenzo Forest, 
IBISCA project, 9-16'47"X, 79^58'\V, mosaic, Oct 
2003 (A. Dejean, J. Orivel, B. Corbara, H.-P. 
Aberlenc k. M. Leponce) - 1 g>ne [CPDC]; same 
location, fogging (FO-R3-05re, FO-R3-01), 18 
May 2004 (J. Schmidl & J. Bail) - 2 g^nes 
[CPDC]; same location, Hght trap, (LC3-C3-3), 
20 May 2004 (A. Comejo et al) - queen [CPDC]. 

Azteca lanuginosa Emer}": BR.AZIL, Rio de 
Janeiro, Cascadura,! Jan 1906, = 2317 - 2 minor 
'^N-orkers [MZSP]. 

Azteca pilosula Forel: BRAZIL, Bahia, Eheus, 
nheus - Para, (J.H.C. Delabie) - 1 g>ne [CPDC]. 

^Azteca schimperi Emer}*: BRAZIL, Amazonas, 
Manaus, 14 Apr 1981, Carton nest in Cecropia 
concolor, (IXPA =428F) - 1 alated ^-ne, 1 male 
(measured), 2 minor ^vorkers [MZSP]; measure- 
ments of Azteca schimperi male: HLA 0.74, H\V 
0.66, SL 0.10, EL 0.28, CI 89, SI 14. COSTA RICA, 
San Jose de Costa Rica, (H. Schmidt) - 1 major 
^vorker [MZSP]. 


Longino (2007) proposed four features 
that distinguish the species in the aurita 
group from other species of ants of the 
genus Azteca (see introductor}' section), but 
one of those, the proportion HLB/HLA > 
1.04, is not a consistent and stable feature 
^vithin some of the females studied here 
{e.g., A. liiiamariae parat}-pe). This trait, 
therefore, should not continue to be used 
as diagnostic tool for the aurita group while 
all the other traits are strongly consistent: 
the palpal formula is 4,3; the mdddle and 
hind tibia lack an apical spur; the ante- 
romedial border of the ch-peus is strongly 
convex and extends well beyond the 
anterolateral ch*peal lobes. These traits 
can, ho-^vever, still be of great taxonomic 
value for separing the aurita group from 
other groups of species in the genus A^zteca. 


Journal of Hymenoptera Research: Festschrift Honoring Roy Snellevjg 

Fig. 11. y42f ^crt sdz/mper/ major worker with vestigial 
wings, Costa Rica. 

Until now, Tapinoma nigerrimum (Ny lan- 
der) is the only species in the Dolichoder- 
inae subfamily for which winged or 
stumped workers have been reported 
(Scupola 2008), something that is always 
considered to be a characteristic of gynes. 
Unexpectedly, the examination of the 
Azteca aurita group material coming from 
MZSP allowed us to observe a major 
worker of A. schimperi coming from Costa 
Rica with all of the morphological char- 
acters of a worker except for its vestigial 
wings (Fig. 11). This worker presents some 
characteristics that differentiate it from 
normal workers: the head is a little larger 
than the normal major workers from the 
same species (HLA: 1.56 vs. 1.16-1.51 mm), 
the median ocellus a little more developed 
and similar to that of the gynes of A. 
schimperi, and there is the presence of wing 
rudiments on the mesosoma. The elements 
that determine these kinds of morphologi- 
cal anomalies are generally considered to 
be environmental factors, as these are 
essential for the determination and devel- 
opment of the casts (Scupola 2008); these 
environmental factors, which have a deci- 
sive function in the formation of a hybrid 
phenotype, are poor nutrition and different 
chemicals (Heinze 1998). Nevertheless, 
simple genetic accidents, viruses or para- 
sites can also provoke the development of 
abnormalities. This is the first report of the 
Intercast syndrome in the genus Azteca and 
the second for the Dolichoderinae subfam- 

ily, after the results reported by Scupola 
(2008) for Tapinoma nigerrimum. 


We thank Chris Starr and Fernando Fernandez for 
inviting us to participate in this tribute to our friend 
and colleague Roy Snelling, Jack Longino and the 
journal's reviewers for useful comments on the paper, 
Beto Brandao and Rodrigo Feitosa for loans of 
specimens, from the MZSP, Andrea Dejean for help 
with the English, Jenny Galvis for access to optical 
equipment, and Brian Fisher and the AntWeb team for 
images of Azteca snellingi. Funding for this study was 
provided by the IBISCA program, with special thanks 
to Maurice Leponce, Jerome Orivel, Bruno Corbara, 
Yves Basset, Nadia Barbosa do Espirito Santo e Servio 
Ponte Ribeiro who made the interesting IBISCA 
material available for study. This study was made 
possible through the Jovenes Investigadores program 
[COLCIENCIAS - University of Magdalena agreement 
# 122 to RJGF] and the Insects of Colombia Project 
[NSF Grant No DEB No 9972024 to Mike Sharkey 
(University of Kentucky) and Brian Brown (LACM)]. 
The first author would like to thank the Movilidad 
Intemacional program of COLCIENCIAS and the 
University of Magdalena for funding his visit to the 
CEPLAC. J.H.C.D. acknowledges his research grant 
from CNPq; and AD was partially funded by the 
Programme Amazonie II of the French Centre National 
de la Recherche Scientifique (project 2ID) and the 
Programme Convergence 2007-2013, Region Guyane 
from the European Community (project DEGA). 


Basset, Y., B. Corbara, H. Barrios, P. Cuenoud, M. 
Leponce, H.-P. Aberlenc, J. Bail, D. Bito, J. R. 
Bridle, G. Castahomeneses, L. Cizek, A. Comejo, 
G. Curletti, J. H. C. Delabie, A. Dejean, R. K. 
Didham, M. Dufrene, L. L. Fagan, A. Floren, D. 
M. Frame, F. Halle, O. J. Hardy, A. Hernandez, R. 
L. Kitching, T. M. Lewinsohn, O. T. Lewis, M. 
Manumbor, E. Medianero, O. Missa, A. W. 
Mitchell, M. Mogia, V. Novotny, F. 0degaard, 
E. Gama de Oliveira, J. Orivel, C. M. P. Ozanne, 
O. Pascal, S. Pinzon, M. Rapp, S. P. Ribeiro, Y. 
Roisin, T. Roslin, D. W. Roubik, M. Samaniego, J. 
Schmidl, L. L. Sorensen, A. Tishechkin, C. van 
Osselaer, and N. N. Winchester. 2007. IBISCA- 
Panama, a large-scale study of arthropod beta- 
diversity and vertical stratification in a lowland 
rainforest: rationale, description of study sites 
and field methodology. Bulletin de I'lnstitut Royal 
des Sciences Naturelles de Belgique, Entomologie 77: 

Emery, C. 1893. Studio monografico sul genere Azteca 
Forel. Memorie della Reale Accademia delle Scienze 
deU'Istituto di Bologna 5: 119-152. 

Volume 19, Number 1, 2010 


Forel, A. 1899. Formicidae. Biologia Centrali-Americana 

3: 1-160. 
HoUdobler, B. and E. O. Wilson. 1990. Vie Ants. Hansard 

University Press, Cambridge, MA. 732 pp. 
Heinze, J. 1998. Intercastes, intermorph, and ergatoid 

queens: who is who in ant reproduction? Insect 

Sociaux 45: 113-124. 
Longino, J. T. 1989. Taxonomy of the Cecropia- 

inhabiting ants in the Azteca alfari species group: 

evidence for two broadly sympatric species. 

Contributions in Science (Natural History Museum 

of Los Angeles County) 412: 1-16. 
. 1991a. Azteca ants in Cecropia trees: taxonomy, 

colony structure, and behavior. Pp. 271-288 in: 

Ant-Plant Interactions, Huxley, C, and D. Cutler 

eds., Oxford University Press, Oxford. 

. 1991b. Taxonomy of the Cecropw-inhabiting 

Azteca ants. Journal of Natural History 25: 

. 1996. Taxonomic characterization of some live- 
stem inhabiting Azteca (Hymenoptera: Formicidae) 
in Costa Rica, with special reference to the ants of 
Cordia (Boraginaceae) and Triplaris (Polygonaceae). 
Journal of Hymenoptera Research 5: 131-156. 

. 2007. A taxonomic re\dew of the genus Azteca 

(Hymenoptera: Formiddae) in Costa Rica and a 
global re\asion of the aurita group. Zootaxa 1491: 1-63. 

Scupola, A. 2008. Presence of brachypterous inter- 
castes in Tapinoma nigerrimum (Nylander, 1856) 
(Hymenoptera, Formicidae). Bollettino del Museo 
Civico di Storia Naturale di Verona, Botanica e 
Zoologia 32: 157-160. 

Vol. 19(1), 2010, pp. 66-76 

Social Behaviours in Solitary Bees: Interactions Among Individuals in 
Xeralictus bicuspidariae Snelling (Hymenoptera: Halictidae: Rophitinae) 

Miriam H. Richards* and Laurence Packer 

(MHR) Department of Biological Sciences, Brock University, 500 Glenridge Ave., St. Catharines, 

ON, Canada L2S 3A1 
(LP) Department of Biology, York University, 4700 Keele St., Toronto, ON, Canada M3J 1P3 

Abstract. — Understanding behavioural interactions among ancestrally solitary species is key to 
understanding the evolutionary origins of group living and cooperation. Previously, Packer (2006) 
showed that circle tube arenas can be used to evaluate the social status of species for which nesting 
data are unavailable. We used circle tube arenas to study the behaviour among 30 female dyads of 
the solitary halictid bee, Xeralictus bicuspidariae Snelling, a member of the subfamily Rophitinae, all 
members of which are ancestrally solitary. Overall, 75.2% of frontal encounters resulted in 
avoidance, 20.7% in aggression, and 4.1% in a successful pass, values which are similar to those 
previously observed in solitary halictids. Although passing events, which are interpreted as 
cooperative behaviour, were rare, they were significantly correlated with bees' rates of approach 
and avoidance, and also with differences between dyad members in rates of ovarian development. 
Rates of aggression were not correlated with physical traits of females or with other behaviours. We 
compare the circle tube behaviour of X. bicuspidariae to previously studied solitary and social 
halictids, and provide statistical support for this method of assessing social status. 

The origin of eusociality is one of the 
major events in the evolutionary history of 
life (Szathmary and Maynard Smith 1995), 
yet our understanding of w^hat transpires 
during transitions to sociality remains 
poor. One reason is the great age at which 
most solitary to eusocial transitions took 
place - over 100 million years ago for 
termites, ants, and vespid w^asps (Wenzel 
1990; Martinez-Delclos and Martinell 
1995), perhaps somewhat less in the bees 
(Michener and Grimaldi 1988), and around 
20 million years ago in the three main 
lineages of eusocial Halictinae (Brady et al. 
2006). The great age of these social lineages 
means that to investigate the evolutionary 
origins of sociality, we must often use 
comparative methods based on detailed 
knowledge of the behaviour of extant 
species. However, a second reason for our 
incomplete understanding of the origins of 

•Author for correspondence: 

eusociality stems from our poor knowl- 
edge of solitary species, whose behaviour 
is most likely to represent the ancestral 
forms from which sociality evolved. 

Sweat bees (Halictidae: Halictini and 
Augochlorini) are the most socially vari- 
able group of animals on earth, including 
species that run the gamut from obligately 
solitary to obligately social, with sociality 
varying from communal to semisocial and 
eusocial forms (Schwarz et al. 2007). There 
are even examples of intraspecific social 
polymorphism, in which solitary or social 
behaviour is expressed within or among 
populations, often in response to variabil- 
ity in environmental conditions (Schwarz 
et al. 2007). The ecological processes that 
shape the social behaviour of modern 
halictines are often considered to be analo- 
gous to those that shaped the evolution of 
major social transitions in the subfamily as 
a whole, including at least three origins of 
eusociality and multiple reversions to 

Volume 19, Number 1, 2010 


solitary behaviour (Danforth et al. 2003; 
Danforth et al. 2008). 

Recent behavioural studies provide in- 
triguing evidence that behavioural transi- 
tions from solitary to social behaviour may 
occur easily and rapidly. Jeanson et al. 
(2005) observed that in forced associations 
of solitary L. (Chilalictus) NDA-1 or of 
communal L. hemichalceum, both domi- 
nance interactions and division of labour 
arose as natural outcomes of normal 
solitary behavioural patterns expressed in 
the context of novel, social environments, 
the effect being stronger in the solitary than 
in the communal species. A similar phe- 
nomenon occurred in experiments on 
solitary Ceratina carpenter bees (Apidae, 
Xylocopinae) when females were forced to 
nest in social associations (Sakagami and 
Maeta 1977). This suggests that at the very 
beginrung of evolutionary transitions to 
caste-based sociality, 'emergent' social be- 
haviour by solitary bees could provide the 
behavioural substrate upon which natural 
selection acts, before the evolution of caste- 
based sociality. This fascinating possibility 
makes understanding the behaviour of 
obligately solitary bees all the more critical. 

Although behavioural interactions among 
individuals of social and socially poly- 
morphic sweat bees have been studied in 
detail on numerous occasions (Breed et al. 
1978; Buckle 1984; McConnell-Garner and 
Kukuk 1997; Wcislo 1997; Soucy 2002), 
solitary species have received less attention. 
Consequently, we know little about the 
potential for social interactions among 
individuals of solitarily nesting species, 
and how naturally occurring variation in 
individual behaviour might impinge upon 
the development of sociality remains ob- 
scure. Of course, one problem with study- 
ing the behaviour of soUtary bees is a dearth 
of opportunities for observing known in- 
dividuals under natural conditions at suffi- 
cient frequency to permit detailed analysis. 
Fortunately, a recent comparative study 
suggests that the circle tube arena, a circle 
of clear plastic tubing in which bees are 

forced to interact (Breed et al. 1978), is one 
route to obtain sufficient behavioural data 
on interactions among individuals of soli- 
tary species (Packer 2006). 

In this paper we analyse the results of a 
detailed study of interactions among in- 
dividuals in a solitary species of the bee 
family Halictidae. Xeralictus bicuspidariae 
Snelling is a member of the halictid 
subfamily Rophitinae with several advan- 
tages as a study organism. First, the 
phylogenetic position of rophitine bees 
suggests that their solitary behaviour is 
ancestral, i.e. there is no evidence that there 
has been any sociality in the evolutionary 
history of the entire subfamily (Danforth et 
al. 2008; Patiny et al. 2008), so any potential 
for social interactions that might be in- 
duced experimentally, is part of its solitary 
ground plan. Second, this bee exhibits 
considerable variation in colour of the 
metasoma of females (Snelling and Stage 
1995) such that pairs can easily be chosen 
to permit individual recognition without 
the intervention of artificially marking the 
bees (marking has been shown to influence 
interactions among individuals; Packer 
2005). Third, it is a large bee, facilitating 
observations of behaviours. 


Xeralictus bicuspidariae was studied at 
Dome Rock Road, La Paz County, Arizona, 
USA, in April 2005. Female bees were 
collected from flowers of Mentzelia (Loasa- 
ceae) and retained in microcentrifuge tubes 
for no more than 30 minutes before 
behavioural observations commenced. This 
duration between capture and observation 
was maintained to reduce the effect of 
captivity-induced physiological changes 
upon behaviour (Pabalan et al. 2000). Two 
bees were then placed simultaneously in a 
clean, plastic circle tube of internal diam- 
eter 7 mm and length 20 cm. Simultaneous 
entry precludes ownership effects (Wcislo 
1997), and this tube diameter was sufficient 
for the two individuals to pass one another 
and to turn around (Packer 2005), but 


Journal of Hymenoptera Research: Festschrift Honoring Roy Snelling 

narrow enough that one bee could block an 
attempted pass by the other. Observations 
lasted for fifteen minutes, a time period 
sufficient for differences in behaviour 
between individuals within a pair to be 
detected, and took place outdoors in the 
shade. The metasomal colour of females 
varied from entirely brick red to entirely 
dark brown (Snelling and Stage 1995); 
pairs were set up with one red and one 
dark female that could be easily differen- 
tiated by the observer without being 
artificially marked. Several experiments 
were terminated when discrimination be- 
tween the individuals was found to be 
more difficult than expected. 

An approach was taken to have occurred 
when individuals came within a distance 
of one body length of each other (Kukuk 
1992; Packer 2005). Both frontal (head to 
head) and front-to-back (head to tail) 
encounters were assessed and their out- 
comes classified into categories: ap- 
proaches, aggression, avoidance and 
passes. Aggressive interactions included 
nudges, lunges and C-postures. Interac- 
tions that resulted in avoidance arose when 
one individual moved away from a sta- 
tionary individual or they both moved 
away from each other. A pass was scored 
when the two bees manoeuvred to permit 
one to move past the other, or they both 
moved past one another simultaneously. 
All behavioural observations were carried 
out by LP, and are therefore directly 
comparable to the data presented in Packer 
(2006). For more detailed descriptions of 
individual behaviours, see Batra (1966). 

All bees were measured and assessed for 
relative age and reproductive condition as 
follows. Head width was measured as the 
greatest distance across the compound 
eyes; this was the greatest diameter of the 
head in dorsal view. Relative wear was 
assessed from mandibles, scored from 
(unworn) to 6 (worn to the base of the 
subapical mandibular tooth), and from 
wings (the total number of nicks along 
the margin of the left fore wing). 

Reproductive status was estimated based 
upon dissection of the metasoma. The 
spermatheca was inspected for the presence 
of sperm and ovarian development was 
assessed by estimating the size of oocytes in 
each ovariole relative to the size of a fully 
developed oocyte, and summing the result- 
ing proportions across all six ovarioles. As 
expected for a solitary bee during nest 
provisioning, all females had mated and 
so matedness was not considered further. 

Statistical analysis. — In circle tubes, the 
behaviour of each member of a pair is 
affected by the behaviour of the second 
member of the pair. This creates a problem 
of statistical non-independence between 
members of each dyad. A second problem 
is variation in behavioural rates among 
pairs - some pairs are very active and some 
do almost nothing. A common approach 
has been to standardize focal behaviors by 
the encounter rate, which in effect means 
all the behaviours are analyzed as ratios, so 
information related to absolute frequency 
is lost and the statistical problems of 
analysing ratios are gained. To address 
these issues, we present an approach 
somewhat different than in previous circle 
tube studies. First, when behavioural pat- 
terns of individuals are considered, we 
analyse only one individual per dyad (red 
bees or dark bees), which avoids inflating 
the number of degrees of freedom in each 
measurement. Second, when properties of 
dyads are considered, we analyse both 
behavioural frequencies and physical traits 
in terms of differences between each bee in 
a pair. Correlations between trait differ- 
ences can be interpreted in the same way 
as correlations between the traits them- 
selves. For instance, a negative correlation 
between head width difference and wear 
difference would indicate that larger bees 
tended to be less worn. All differences 
between pair members were calculated as 
(value for red bee) - (value for dark bee), 
except for head width (HW) difference, 
which was calculated as (red HW - dark 
HW) / average HW. 

Volume 19, Number 1, 2010 


Table 1. Physical characteristics of adult female Xeralictus bicuspidariae used in circle tube experiments. 
Signed rank tests were used to compare the physical characteristics of red and dark females in each dyad; non- 
significant (n.s.) results indicate that overall, red and dark females were equivalent. 





Signed rank test 

Head width (mm) (n=56) 




S = -22.5, n.s. 

Mandibular wear (n=56) 




S = 31.5, n.s. 

Wing wear (n=48) 




S - 4.0, n.s. 

Total ovarian score (n=55) 




S = 8.0, n.s. 

All variables, including differences, were 
checked for normality using the array of 
tests in SAS 9.1 (PROC UNIVARIATE); 
since several variables were non-normally 
distributed, we mainly used non-para- 
metric statistical methods. Additionally 
we used principal components analysis 
(PCA) to further explore and confirm 
relationships among physical and behav- 
ioural variables in X. bicuspidariae. Initially, 
the PCA was based on eight variables 
(entered as untransformed differences be- 
tween females in mandibular wear, wing 
wear, head width, total ovarian score, 
approach frequency, aggressive frequency, 
avoidance frequency, and pass frequency). 
However, since Kaiser's Measure of Sam- 
pling Adequacy (MSA) with all eight 
variables had a value of only 0.467, the 
variable with the lowest communality 
measure (head width) was dropped from 
the PCA. With the remaining seven vari- 
ables, MSA =0.63, which exceeds the 0.6 
criterion. We present both factor loading 
scores (the degree to which each variable 
influences the inferred factors) and com- 
munality estimates (a reliability score 
which estimates the proportion of variance 
in each variable that is jointly explained by 
all three factors). 

Packer (2006) argued that the social 
status of halictine bees can be accurately 
assessed using circle tube assays of fe- 
males, even in the absence of nesting data. 
Solitary bees should be characterized by 
high levels of avoidance behaviour, com- 
munal bees by high levels of cooperative 
behaviour (passing) and low levels of 
aggression, and semisocial and eusocial 

bees by low levels of cooperation and 
high levels of both aggression and avoid- 
ance. We used discriminant functions 
analysis (DFA) to assess how accurately 
X. bicuspidariae and 21 other species (refer- 
ences in Packer 2006) can be categorized as 
solitary, communal, or semi and eusocial, 
based on the percentages of avoidance, 
aggression, and passing behaviours in 
circle tubes. 


Circle tube assays. — Physical traits of the 
60 females used in 30 circle tube trials are 
presented in Table 1. There were no sig- 
nificant correlations among body size, 
degree of wear, and degree of ovarian 
development within individuals used in 
the behavioural tests, although degree of 
mandibular wear was positively correlated 
with degree of wing wear (Pearson correla- 
tion coefficient, r=0.55, n=48, p<0.0001). 
All females had at least one V4-size, 
developing oocyte, and 25 of 52 (48%) 
dissected females contained a full-size 
oocyte, ready to lay. The mean difference 
between red and dark females in each 
dyad for each of these characteristics, 
was zero (Table 1), so bee colouration had 
no significance other than providing a 
convenient identification tool for the ob- 

The frequencies of each of the four 
classes of behaviour per dyad and per 
female are given in Table 2. The most 
frequent behaviours were approaches 
(32.5 per dyad) and avoidance (25.1 per 
dyad), followed by aggressive behaviours 
(6.7 per dyad) and passing (1.2 per dyad). 


Journal of Hymenoptera Research: Festschrift Honoring Roy Snelling 

Table 2. Behavioural frequencies for approach, avoid, pass, and aggressive behaviours. Note that 
'aggression' includes C-postures, biting, and pushing. Since the behavioural rates of each member of a dyad 
are non-independent, only one bee per dyad is used to provide an estimate of behavioural frequencies per 
indi\idual. Measurements of mutual behaviour refer to simultaneous performance of that behaviour by both 
members of a dyad. Sample size N=30 dyads, except where otherw^ise noted. 






Frontal encounters 

Dyad total 





Dyad total ' 




Red female ' 




Dark female 









Dyad total 




Red female 




Dark female 









Dyad total 




Red female 




Dark female 









Dyad total 




Red female 




Dark female 





Overall, 75.2% of frontal encounters re- 
sulted in avoidance, 20.7% in aggression, 
and 4.1% in a successful pass. 

Aggressive acts were observed in all 30 
pairs, and by 52 of the 60 (87%) individuals 
assayed. Withdrawals were also observed 
in all 30 pairs; only 1 bee of 60 (2%) did not 
display a unilateral withdrawal, but she 
did take part in a mutual (bilateral) with- 
drawal. Passing or cooperative acts were 
rare, being observed in only 22 of 30 (73%) 
pairs. Of a total of 35 passes, 20 {57%) were 
bilateral (both bees moved past each other) 
and 15 (43%) were unilateral (1 bee moved 
past the other bee). 

Based on behavioural frequency differ- 
ences (red bee - dark bee), three behav- 
iours, approach, avoid, and pass were 
found to be mutually positively correlated 
(i.e. the bee that did one behaviour more 
frequently also did the other behaviour 
more frequently; Spearman rank correla- 
tions: approach vs. avoid, r= 0.798, n=29, 
p<0.0001, approach vs. pass: r = 0.577, 
n=29, p=0.001; avoid vs. pass: r=0.460, 
n=30, p =0.010), but none was correlated 
with the frequency of aggression (aggress 

vs. approach: r= —0.224, n=29, n.s.; aggress 
vs. pass: r=— 0.106, n=30, n.s.; aggress vs. 
avoid: r= -0.193, n=30, n.s.). 

Differences between bees with respect to 
head width, wing wear, and mandibular 
wear were not significantly correlated with 
differences in behavioural frequency for 
any of the behaviours. Differences in total 
ovarian score did correlate positively with 
the rates of approach and pass, although 
not with either avoidance or aggressive 
frequencies (Table 3, Fig. 1). In other 
words, the female with greater ovarian 
development was almost significantly 
likely to approach and was significantly 
more likely to pass than the female with 
lesser ovarian development. 

A principal components analysis (PCA) 
further describes behavioural and physical 
variation among female interactants in 
circle tubes. As outlined in the Methods, 
the PCA (Table 4) included all variables 
except head width, which contributed little 
to understanding variation among the 
dyads. Three factors had eigenvalues > 
1.0 and were retained, explaining 77.9% of 
the variation among dyads. Factor 1 was 

Volume 19, Number 1, 2010 


Table 3. Influence of female physical status on behaviour. Spearman rank correlations were based on 
differences in both female traits and differences in behaviour frequencies (red bee - dark bee). Positive 
correlations indicate that the bee with the greater trait value exhibited the behaviour more frequently. 

Behaviour (N= number of dvads) 

Physical trait of females 





Head width 

0.197 (27) 

0.018 (28) 

-0.054 (28) 

0.292 (28) 

Wing wear 

0.020 (20) 

-0.123 (21) 

-0.092 (21) 

0.077 (21) 

Mandibular wear 

-0.007 (27) 

-0.100 (28) 

-0.068 (28) 

0.215 (28) 

Total ovarian score 

0.451 (26), p = 


0.250 (27) 

0.435 (27), p= 


-0.132 (27) 

most influenced by non-aggressive behav- 
iour and ovarian development, reflecting 
the previously noted positive association 
between ovarian development and ap- 
proach and passing frequencies. Factor 2 
was influenced mainly by mandibular and 
wing wear; thus Factor 2 describes varia- 

bility in wear differences among dyads, 
and so does not reflect behavioural varia- 
tion. Factor 3 was influenced mainly by 
aggression. PCA based only on the four 
behavioural frequency differences, re- 
sulted in two factors that together ex- 
plained 79.7% of the variation among 

I o aggress | 

jo avoid I 


O O 

o o o 





°o a 

' o 

o o 








D -5- 






: approach 

I A cooperate I 

)5 ^ ><l 





A4 A A 






-1 ■ 


▲ -3 

1 0.5 1 




Fig. 1. Influence of ovarian development (OD, horizontal axes) on different behaviours (vertical axes), scored 
in terms of the differences between individuals (red-dark). Positive values on the horizontal and vertical axes 
indicate a greater value for the red bee, whereas negative values indicate a greater value for the dark bee. Top 
left: OD vs. aggression. Top right: OD vs. avoidance. Bottom left: OD vs. approach frequency. Bottom right: OD 
vs. cooperation (pass). 


Journal of Hymenoptera Research: Festschrift Honoring Roy Snelling 

Table 4. Principal components analysis describing variation annong dyads based on differences between 
interactants in both physical and behavioural traits. Three factors were retained with eigenvalues > 1, 
explaining a cumulative total of 77.9% of the variation among dyads. Relatively strong factor loading scores 
(>0.6) are indicated in boldface. Communality estimates describe the proportion of variance in each trait that is 
jointly explained by Factors 1, 2 and 3. Kaiser's overall Measure of Sampling Adequacy (MSA) was 0.6312. 

Trait (difference between females) 

Factor 1 

Factor 2 

Factor 3 

Communality estimate 

Mandibular wear 





Wing wear 





CKarian score 





























Variance explained 




dyads. Factor 1, which explained 55.0% of 
the behavioural variation among dyads, 
was strongly influenced by approaches 
(loading score 0.910), avoidance (0.866), 
and passes (0.752). Factor 2, which ex- 
plained 24.6% of the variation, was 
strongly influenced by aggression (loading 
score 0.959). 

Comparison of X. bicuspidariae with other 
halictines. — Fig. 2 compares X. bicuspidariae 
to 21 other species, in terms of the propor- 
tion of avoidance, aggressive, and coopera- 
tive (passing) behaviours observed in circle 
tube assays. It most closely resembles 
Penapis toroi, another solitary rophitine. 
Discriminant functions analysis (DFA) 
based on four putative categories (solitary, 
communal, semisocial, and eusocial) per- 
fectly assigned solitary and communal 
species, but failed to distinguish between 
the latter two, assigning 1 eusocial species 
to the semisocial category and 1 semisocial 
species to the eusocial category. DFA based 
on three putative categories (solitary, com- 
munal, and caste-based social) reassigned 
each species into the category presented in 
Fig. 2. Moreover, when Caenohalictus pygo- 
sinuatum was categorized as communal 
(Michener et al. 1979), then DFA assigned 
it to the solitary group (as suggested by 
Packer 2006). The success of the DFA 
approach is based on significant differ- 
ences among solitary, communal, and 

caste-based social bees in the proportions 
of aggressive behaviour (ANOVA, 
F=50.32, df=2,19, p<0.0001) and avoid- 
ance behaviour (ANOVA, F= 15.15, 
df =2,19, p<0.0001), as well as significantly 
more frequent passing behaviour in com- 
munal species, as compared to both soli- 
tary and social species (ANOVA, F= 62.55, 
df=2,19, p<0.0001). 


Solitary behaviour ofX. bicuspidariae. — In 
X. bicuspidariae, differences between circle 
tube interactants in head width, wing 
wear, and mandibular wear were not 
associated with differences in behaviour, 
suggesting that neither body size nor wear 
(and possibly age) structured interactions 
among adult females. Differences in ovar- 
ian development (OD) also did not predict 
differences in either aggression or avoid- 
ance, but were associated with rates of 
approach and pass behaviours, these being 
exhibited more frequently by the bee with 
greater ovarian development. Why would 
high OD females be more likely to ap- 
proach and especially, to pass? One possi- 
bility is that the closer a female is to laying 
an egg, the more active she is likely to be. 
Under natural circumstances, a female 
halictine getting ready to lay an egg should 
be spending considerable time readying a 
brood cell and provisioning it. In a circle 

Volume 19, Number 1, 2010 








X. blcuspidariae 

Pe. toroi 

T. prognathus 

Ca. pygosinuatum 

Ca. cuprellus 

L platycephalum 

L (Ct.) NDA-1 

L. ancense 

L. figueresi 

Co corinogaster 

Co. melanocladius 





Co. chloris 

Co. herbsti 

Co. patagonica 

■ avoidance 
n aggression 

■ pass 


mv , 



H. lanei 

L zephymm 

L hemichalceum 
R. mutabilis 
R. proxima 
Ca. dolator 
Ca. sp. A 
Ps. pississi 

L (Ct.) NDA-1 ' 
L. hemichalceum 







Fig. 2. Comparison of circle tube behaviour of X. blcuspidariae with literature values (partially redrawn from 
Packer 2006, which also contains a complete list of references). Solitary bee species are characterized by high 
rates of avoidance (withdrawals), communal bees by high rates of cooperation or tolerance (passing), and 
semisocial and eusocial species by high rates of aggression, coupled with very low rates of passing. Differences 
in behavioural profiles of bees tested in circle tubes versus vertical tubes (Jeanson et al. 2005) are evident. Genera 
represented (top to bottom) are Xeralictus (X.), Penapis (Pe.), ThrincoJmlictus (T.), Caenohalictus (Ca.), Lasioglossum 
(L.), Cojynura (Co.), Halictus (H.), Ruizantheda (R.), and Pseudagapostemon (Ps.). 

tube, heightened activity rates make it 
more likely that a bee will approach the 
second bee, and then perhaps continue 
right past her. In other words, bees with 
higher ovarian development may be more 
motivated to remain active, which in a 
circle tube would result in higher rates of 
approaching and passing. 

The lack of correlation between ovarian 
status and aggression might seem surpris- 
ing, but is consistent with observations in 
other species, including the solitary halic- 
tine, L. (Ctenonomia) NDA-1, and the 
communal species, L. (Chilalictus) 'platyceph- 
alum, in which ovarian status was not 
associated with aggression (McConnell- 
Gamer and Kukuk 1997). In contrast, in 
Lasioglossum figueresi the female with larger 
ovaries was often first to be aggressive, and 

the bee with smaller ovaries was often first 
to withdraw (Wcislo 1997). Even in obli- 
gately eusocial species like Halictus ligatus, 
dominance, aggression, and defensive be- 
haviours are most likely and most severe 
when the two members of a dyad both 
have relatively high ovarian development, 
for example, when a queen is paired with a 
worker with highly developed ovaries 
(Pabalan et al., 2000). This suggests that 
correlations between OD and aggression 
might have more to do with reactions to 
the threat of egg replacement than with 
dominance behaviour per se. In solitary 
bees, we should not then expect to see a 
correlation between ovarian development 
and aggressive behaviour (or withdrawal 
behaviour) except perhaps in species with 
high rates of intraspecific egg parasitism. 


Journal of Hymenoptera Research: Festschrift Honoring Roy Snelling 

Our experiments uncovered consider- 
able variability among individuals. Ran- 
dom pairings of individuals with different 
behavioural tendencies (personalities) 
must then have contributed to variation 
among dyads, creating behavioural scenar- 
ios that bear a marked resemblance to 
those expressed by social bees. Indeed, this 
v^ould seem to be the basis of the phenom- 
enon of 'emergent' sociality as described 
by Jeanson et al. (2005). However, the use 
of the term 'emergent' to describe forced 
social interactions among solitary bees is 
somewhat problematic, even in those 
which like X. bicuspidariae are ancestrally 
and monomorphically solitary. This is 
because solitary bees may also experience 
social interactions that insect sociobiolo- 
gists do not usually categorize as 'social', 
such as interactions between foragers on 
flowers, between nest residents in dense 
nesting aggregations, between nest resi- 
dents and would-be nest usurpers, or 
between residents and egg kleptoparasites. 
Moreover, group living may occur at very 
low frequencies in some solitary species 
without extensive nest observations, as has 
recently been found for several species of 
the apid genus, Cemtina (Rehan et al. 2009). 
In other words, many solitary bees, both 
ancestrally solitary and ancestrally social, 
may have considerable scope for intraspe- 
cific social behaviour, even if they rarely or 
never nest in multifemale groups. The 
variability in behavioural syndromes of 
solitary halictines (Fig. 2) suggests that 
eventually it may be possible to detect 
differences among obligately solitary, so- 
cially polymorphic, and reversed solitary 
species, especially based on the frequency 
of avoidance and aggession. 

Behavioural changes in social transitions. — 
One caveat to the use of artificial arenas for 
observing bee behaviour is that the fre- 
quencies of circle tube behaviours may or 
may not represent the frequencies of same 
or similar behaviours in natural settings. 
Indeed, there are obvious differences in 
behavioural frequencies assessed using 

horizontal circle tubes versus vertical lin- 
ear tubes (Fig. 2), implying that major 
differences in behavioural frequencies are 
produced by different experimental meth- 
odologies. Nevertheless, the interspecific 
consistency of behavioural syndromes ob- 
served in circle tube assays of solitary, 
communal, and caste-based social species 
is striking and statistically supportable, 
suggesting that when circle tube assays 
are used consistently, they uncover funda- 
mental differences in behaviour among 
solitary, communal, and semisocial and 
eusocial species. These differences, if not 
the behavioural frequencies themselves, 
can be used to infer general behavioural 
tendencies in bees of different social levels. 

In halictids, the ancestral trait of intoler- 
ance is suggested by high rates of avoid- 
ance in solitary bees such as X. bicuspidariae 
and another solitary rophitine species, 
Penapis toroi, in which avoidance behav- 
iours comprise about 75% of encounters 
(Fig. 2). Transitions to communal versus 
caste-based social behaviour may be quite 
different. Circle tube assays imply that 
solitary-communal transitions involve sig- 
nificant decreases in both aggression and 
avoidance, whereas transitions to caste- 
based eusociality involve a significant 
increase in aggression, coupled with a 
decrease in avoidance. To the extent that 
passes represent cooperative interactions, 
solitary-communal transitions would ap- 
pear to involve huge increases in coopera- 
tion whereas transitions to caste-based 
eusociality involve little change or perhaps 
even a decrease in cooperative behaviour. 
It will be important in future studies of 
both solitary and social halictines, to assess 
the degree of behavioural concordance 
between natural versus artificial contexts 
whenever possible, so that we can actually 
understand how representative circle tube 
behaviour is for those species for which 
nesting data are unobtainable. 

Given that one of the most outstanding 
features of the eusocial insects is their 
frequent and sophisticated cooperative 

VOLL-X^Z If X"_^.-^ZR 


b€ha\'iour, the h}"pothesis that transitions 
to caste-based socialit}' should involve 
decreases in cooperation coupled with 
increases in aggression seems contradic- 
tor'. However, semisocial and eusocial 
halictines not only interact \s'ith. n\any 
more indi\iduals than solitary- bees do, 
but they must also cope %\-ith dominance- 
subordinance relationships, m.any of which 
are regulated bv aggressive beha\"ioui 
(Kukuk and May f991; Pabalan et al. 
2000). Semisocial and eusocial bees must 
be able to exercise both tolerance and 
aggression with the same individuals. 
-\lthough aggressive beha\iours by soli- 
tar}* and caste-based social bees in circle 
tubes may appear to be similar, a m.ajor 
difference in natural settings is that ag- 
gressive beha^-iouT by the latter is likely 
modulated by nest-mate recognition, such 
that encounters \\*ith non-nestmates ^sill 
likely provoke aggression, whereas en- 
counters ^N-ith nestmates m.av engender 
aggression, tolerance, or cooperation (Peso 
and Richards 2010), depending on the 
immediate behavioural context. 

Transitions to social behaviour, especially 
to caste-based sociality' , are rarer in halic- 
tines than reversals to solitary- behav- 
iour (DaTLtorth et al. 2003). Recent evidence 
suggests that reversals to soHtar}' behav- 
iour do not necessarily retrace the original 
evolutionary* steps that led to socialitv'. For 
instance, reversed-solitarv* Lasioglossum 
have retained the social nesting character- 
istic of constructing brood ceUs close to the 
m.ain burrow, facilitating both maternal 
inspection and care of the cells (Plateaux- 
Quenu 2008), and the potential for social 
interactions among newly emerged, adult 
brood. Thus reversed soHtarv* bees mav 
have lost caste-based sodalitv*, but may 
have retained the context-dependent ability' 
to discriminate nestmates from non-nest- 
mates. Circle tube comparisons of ances- 
trally sohtarv' species Like X hiaispidariae, 
and reversed-solitar}- species like L. figiier- 
esi, may help to iQuminate and distinguish 
the evolutionarv- sequences involved in 

fonvard and reverse social transitions, 
especially where these involve the expres- 
sion of context-dependent behaviour. 


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r :r -j-f manuscript and 
: z : : : : : . 5 suggestions for 
refeir:!' 15 funded by 
—rir.g Refearch Council of 
to both authors. 


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J. H\'\I. RES. 
Vol. 19(1), 2010, pp. 77-83 

Plega hagenella (Neuroptera: Mantispidae) Parasitism of Hylaeiis 

(Hylaeopsis) sp. (Hymenoptera: Colletidae) Reusing Nests of Tn/poxylon 

manni (Hymenoptera: Crabronidae) in Trinidad 

All.\x "W. Hook, John D. Oswald, axd Johx L. Xeff 

(AWTi) Department of Biological Sciences, St Edward's Universin-, Austin, TX 78704-6489, USA; 


(JDO) Department of Entomolog}-, Texas A&M Universit}-, College Station, TX 77845-2475, USA 

(JLX) Central Texas Melittological Institute, 7307 Running Rope, Austin, TX 78731, USA 

Abstract. — Two adult specimens of Flega Imgmella were reared from a nest of the crabronid wasp 
Tryipoxxjlon manni collected in Trinidad in 2008. The mantispids developed by feeding on the 
immature stages of a colletid bee, Hylaeiis (Hylaeopsisj sp., which had secondarily occupied the cells 
of the aerial mud nest of T. manni. T^vo dead Hylaeiis pupae were found ^sithin the nest, one of 
\\'hich had six dead mantispid lar\'ae attached. Numerous insect egg chorions, interpreted as 
belonging to P. Imgenella, were found in clusters inserted into small cavities in the mud of the outer 
surface of the nest. Five additional adult P. Imgenella ^vere collected in microhabitats where other 7. 
nuinni nests were collected. These new obser\'ations confirm the presence of P. liagenella in Trinidad, 
establish the presence of the subgenus Hylaeopsis in Trinidad, and document Flega liagenella as a 
new parasite of bees in the genus Hylaeiis. 

The insect order Xeuroptera - lace^vings, 
antlions and their relatives - currently 
comprises approximately 5,730 valid extant 
species placed in about 17 families. The 
family Mantispidae - mantispids or man- 
tis-flies - form a distinctive clade of about 
395 species that is ^vell kno^\Ti for the 
raptorial forelegs of its predator}' adults, 
and the larval life histor}^ strategy of 
species in the subfamily Mantispinae as 
spider egg parasites. Less ^vell kno^\Ti are 
the lar\'al strategies of species in the three 
other extant mantispid subfamilies - the 
Symphrasinae, Drepanicinae and Calo- 
mantispinae - none of ^vhich are known 
to be associated with spiders. 

Of particular interest are lar\'al associa- 
tions of species belonging to the subfamily 
Symphrasinae, ^vhich is generally consid- 
ered to be the sister-group to all other 
mantispid subfamiHes collectively (Penny 
and da Costa 1983; Lambkin 1986). The 
biologies of symphrasine species are 
poorly knowTL and, consequently, ne\v host 

records are of considerable interest. The 
S\Tnphrasinae contains three extant genera 
- Anchieta, Trichoscelia and Plega - all of 
which are currently restricted to the Xew 
World (Penny 1982), and for each of which 
few lar\'al host/ feeding records are avail- 
able. Anchieta includes five species kno^\Ti 
from Brazil and French Guiana, only one of 
^vhich has a reported host: A. fiimosella 
(West^vood), which has recently been 
reared from cocoons of Tn/poxxflon (Tnjpar- 
giliim) aestival Richards in southeastern 
Brazil (Buys 2008). Trichoscelia contains 13 
species distributed from southern Mexico 
south to Uruguay, Argentina and southern 
Brazil (Penny 1982; Ohl 2004). Trichoscelia 
varia (Walker) has been reared from nests 
of the vespid wasps, Polybia ruficeps 
Schrottk}' and P. scutellaris (\\lrd\e) (Penny 
1982); and Dejean and Canard (1990) 
provide an interesting account of Trichos- 
celia santareni (Xavas) invading a colony of 
Polybia digiietana Buysson on the Yucatan 

78 Journal of Hymenoptera Research: Festschrift Honoring Roy Snelling 

Table 1. Plega melitomae species-group host records. 

Pkya sp. 

Host record 




Trypoxylon albitarse (Crabronidae) 


Penny 1982 


Hylaeus (Hylaeopsis) sp. (Colletidae), in 
Trypoxylon manni nest 


(this work) 


Melitoma segnientaria (as M. euglossoides) 

Mexico (Chiapas) 

Linsley and MacSwain 1955 

sp. near 

Tn/poxylon sp. 

Mexico (Veracruz) 

Parker and Stange 1965 



Megachile exaltata (Megachilidae) 

Mexico (Yucatan) 

Parker and Stange 1965 

Plega includes 14 species distributed 
from the southwestern United States, south 
to Bolivia and Brazil (Penny 1982). Two 
species groups are generally recognized - 
the melitomae and signata groups. Members 
of the P. melitomae group (P. beardi, 
hagenella, melitomae, paraense and yucatanae; 
see Penny 1982) have been reported as 
parasites of several bees and aculeate 
wasps (Table 1). In contrast to the hyme- 
nopteran feeding records of the melitomae 
group, members of the signata group have 
been reared from subterranean insects 
(Parker and Stange 1965). Plega signata 
Hagen cocoons have been found inside 
subterranean cocoons of the noctuid moth 
Egira curialis (Grote) (Woglum 1935; as 
Xylomyges curialis). Werner and Butler 
(1965) presented circumstantial evidence 
of Plega banksi Rehn devouring a scarab 
pupa as well as an asilid pupa associated 
with a scarab pupa in Arizona. They 
suggested that the larvae live in soil as 

The global crabronid genus Trypoxylon, 
which figures prominently in several host 
records for the P. melitomae group, contains 
more than 600 species and is well repre- 
sented in the New World. Trypoxylon manni 
Richards is known from Brazil and Trini- 
dad and commonly attaches its nests to 
rootlets under dirt banks next to road cuts 
in the Northern Range, Trinidad, West 
Indies (Vesey-FitzGerald 1936). Nests of 
T. manni are often communal and may 
contain up to six females and 64 cells 
(Hook and Starr unpubl.). Trypoxylon 

manni is a member of the T. fabricator 
group, in which nest sharing by females is 
known in at least three species: T. fabricator 
Smith (Sakagami et al. 1990), T. maidli 
Richards and T. manni (Hook and Starr 
pers. obs.). 


Between 2003 and 2008 the senior author 
field collected five specimens of Plega 
hagenella in several places on the island of 
Trinidad. Three of the specimens were 
collected in 2008 under dirt banks harbor- 
ing Trypoxylon manni nests. One female 
was taken on 18 July along the Paria Trail, 
another female was collected on 23 July up 
the Maracas Valley in the Northern Range 
and a male was taken on 25 July in the 
Arena Forest Reserve. A fourth specimen 
(male) was collected on 26 July 2003 up the 
Caura Valley (Northern Range) and a fifth 
specimen (female) was collected on 25 
August 2005 at U.W.I. Flats (apartments) 
in St. Augustine (Trypoxylon spp. were 
observed nesting around the apartments). 

In July of 2008 AWH collected 10 nests of 
Trypoxylon manni - seven from along the 
upper reaches of the Paria Trail (10.746°N 
61.285°W; connecting the village of Brasso 
Seco to Paria Bay on the north coast) on 18 
July, two in the Maracas Valley (10.705°N 
61.368°W) on 23 July and one on 25 July 
along a road entering the Arena Forest 
Reserve (10.562°N 61.256°W). All nests 
were collected into and maintained iso- 
lated in separate plastic bags in order to 
rear their contents. Two adults of Plega 

Volume 19, Number 1, 2010 


hagenella, and numerous presumed eggs 
and larvae of the same species, were later 
found to be associated with one of the nests 
collected along the Paria Trail (AWH field 
note 36-2008). Most of the cells of this nest 
were subsequently dissected and examined 
for evidence of insect occupation and 
usage. Observations made during the ex- 
amination of this nest are summarized 


Observations on Trypoxylon and Hylaeus. — 
The mantispid-parasitized nest had 14 
cells, of which nine were open when 
collected, two were closed with mud and 
three were closed with a tough, transpar- 
ent, membrane indicative of cell reutiliza- 
tion by a colletid bee, subsequently identi- 
fied as Hylaeus (Hylaeopsis) sp. Of nine cells 
opened by dissection, Hylaeus had reuti- 
lized eight, as evidenced by typical Hy- 
laeus-type transparent cell linings and 
partitions found within the original mud 
cells of the Trypoxylon manni nest. Hylaeus 
reused 11 of the 14 cells this nest contained. 
Trypoxylon species do not line their nest 
cells with secretions, but this is a charac- 
teristic feature of colletid bee cells (Al- 
meida 2008). Two dead Hylaeus pupae 
were found in these lined exterior cells. 
Of these, one pupa was unparasitized but 
the other had six dead mantispid larvae 
attached to its thorax and abdomen (the 
head was missing). 

Identification of the bee species as 
Hylaeus (Hylaeopsis) sp. is based on the 
dead pupae found in the nest and on bee 
rearings from 46 additional Trypoxylon 
manni nests collected previously in 1996 
and 1999 (primarily along the Blanchis- 
seuse Road in the Northern Range of 
Trinidad). Interestingly, a diversity of 
cleptoparasites was reared from 19 of the 
earlier 46 nests, but no mantispids. Six 
(13%) of the 46 nests collected during this 
earlier period had some level of Hylaeus 
cell reutilization. Those six nests contained 
a total of 93 cells, of which 16 (17%) were 

reused by Hylaeus, with one to two or 
possibly three Hylaeus cells per T. manni 

Observations on Plega. — Two adult Plega 
hagenella, one male and one female, were 
subsequently found in the plastic bag 
containing the parasitized Trypoxylon nest, 
which had been maintained indoors, dry, 
and at ambient room temperatures since its 
collection. The male was discovered nearly 
dead on 26 July 2008. The nest was 
subsequently checked irregularly until 18 
August. The next inspection was not until 9 
October, when a dead female was discov- 
ered. The adult Plega hagenella specimens 
were identified using the keys and descrip- 
tions of Penny (1982), who noted the 
existence of previously-collected females 
resembling, but not conclusively identifi- 
able as, P. hagenella from Trinidad. The 
present material confirms the presence P. 
hagenella in Trinidad based on definitively 
identifiable male specimens and associated 

Subsequent examination of the nest 
revealed two mantispid pupal exuviae, 
one protruding from each of two nest cells, 
both of which were marked by the char- 
acteristic cellophane-like lining of Hylaeus. 
A loose double-walled cocoon of typical 
neuropteran form was extracted from one 
of these cells. In another cell, six neurop- 
teran larvae of at least two sizes were 
found attached to a dead Hylaeus pupa. 

Further inspection of the partially dis- 
sected nest revealed the existence of 
numerous, small, whitish, insect egg cho- 
rions associated with its outer layers. While 
some of the eggs may have been deposited 
on the exposed outer surface of the nest, 
most appear to have been inserted into 
small cavities in the nest's irregular mud 
surface, often in small groups of 3-6 per 
cavity. The small cavities are natural 
features of the original mud nest, the result 
of incomplete joining or smoothing of 
adjacent, rounded, mud boluses used in 
its construction. At least 29 empty egg 
chorions were found in or on the nest, but 


Journal of Hymenoptera Research: Festschrift Honoring Roy Snelling 

it seems likely that additional eggs were 
lost during the dissection of the nest, or 
were inserted so deeply into crevices that 
they were hidden from view. The eggs 
were narrowly lacrimiform in shape (with 
the micropylar process terminating the 
narrowed end), ca. 0.7 mm long, simple 
(i.e., stalkless), and marked externally with 
a network of raised polygonal ridges. All of 
the eggs appeared to be empty, their larvae 
having emerged through longitudinal slits 
located near the micropylar ends of each 


Hylaeus (Hylaeopsis). — This Neotropical 
subgenus of 15 species has been reported 
from Mexico south to Paraguay and south- 
em Brazil, but it has not previously been 
reported from Trinidad, the neighboring 
Guyanas, or Venezuela (Urban and Moure 
2007). Ten or more undescribed Hylaeopsis 
species are suspected to be present in the 
Neotropical fauna (Michener 2007). Adults 
of the Trinidad Hylaeopsis are much smaller 
than adult Plega hagenella, suggesting that 
Plega larvae must consume multiple bee 
larvae and /or pupae to reach adulthood. If 
this is true, the pattern of multiple bee 
cells per wasp cell and gregarious nesting 
(also reported in H. (Hylaeopsis) tricolor 
(Schrottky) by Sakagami and Zucchi 1978) 
may contribute to making this bee species a 
suitable host for Plega hagenella. Not all 
Hylaeopsis species nest gregariously, how- 
ever, and the hyperdispersed, single-celled 
nests of H. (Hylaeopsis) grossus (Cresson) 
(Michener and Brooks 2003) may be adap- 
tive in helping to avoid attacks by larger 
nest parasites like Plega. 

Plega. — Although definitive, reared, lar- 
vae and eggs of P. hagenella have yet to be 
described, detailed observations derived 
from the parasitized Trypoxylon nest re- 
ported here, together with existing knowl- 
edge of mantispid immatures, present a 
strong circumstantial case that the eggs 
and larvae noted above are those of P. 
hagenella. The rearing of definitively iden- 

tified P. hagenella adults from the Trypox- 
ylon nest renders plausible the discovery of 
immature P. hagenella stages in the same 
nest. The general morphology of the 
recovered larvae is consistent with their 
determination as first-instar mantispid 
larvae. The larvae are morphologically 
similar to the first-instar larvae of both 
Plega yucatanae (described by Parker and 
Stange 1965) and other described first- 
instar mantispids (e.g., Climaciella brunnea 
[as Mantispa brunnea], Dicromantispa inter- 
rupta [as Mantispa interrupta], Dicromantispa 
sayi [as Mantispa sayi], Leptomantispa pul- 
chella [as Mantispa pulchella] and Zeugoman- 
tispa virescens [as Mantispa viridis], see 
Hoffman and Brushwein 1992; Tuberonotha 
strenua [as Climaciella magna] and Mantispa 
japonica, see Kuroko 1961). 

The eggs noted above, in addition to 
being physically associated with the man- 
tispid-parasitized nest, are also consistent 
both morphologically and behaviorally 
with a mantispid identification. Their size 
is appropriate to that of the smaller 
mantispid larvae observed in the nest. 
The deep insertion of many of the eggs 
into fine crevices in the mud surface of the 
nest is consistent with oviposition via a 
slender, elongate, ovipositor, the presence 
of which is, within the order Neuroptera, a 
synapomorphy of the mantispid subfamily 
Symphrasinae, to which Plega hagenella 
belongs. Although at least two Plega 
species have been reared from egg to adult 
(P. dactylota and P. signata; see MacLeod 
and Redborg 1982), no published descrip- 
tions or illustrations of Symphrasine man- 
tispid eggs exist. Interestingly, however, 
the eggs noted here bear a strong resem- 
blance to, though are somewhat more 
slender and elongate than, those illustrated 
by Minter (1990, fig. 1) for Mucroberotha 
vesicaria - a species belonging to a group of 
several genera of uncertain phylogenetic 
placement that have in recent years been 
placed in either the family Mantispidae or 
Berothidae (as Rhachiberothinae), or as a 
separate family (as Rhachiberothidae). The 

Volume 19, Number 1, 2010 


presence of sessile eggs in both rhachiber- 
othines/ids and symphrasine mantispids 
has potentially interesting implications for 
the interpretation of the stalked eggs found 
in both mantispine mantispids and ber- 
othine berothids (both of which are rela- 
tively derived subfamilies within their 
families). If sessile eggs are found to be 
symplesiomorphic in rhachiberothines/ids 
and symphrasine mantispids, the stalked 
eggs found in berothines and mantispines 
are likely to be independent, derived 

Parasite Biology. — Because the Plega ha- 
genella adults emerged from the mud nest 
of Trypoxylon manni, it was initially as- 
sumed that P. hagenella was a parasite of T. 
manni. However, several observations and 
lines of evidence suggest that, at least in 
this case, P. hagenella was parasitizing the 
Hylaeus (Hylaeopsis) sp., not T. manni. First, 
no remnants of Trypoxylon immatures were 
found in the nest, suggesting that the 
original nest builders had vacated the nest 
prior to its occupation by Hylaeus and 
Plega. Second, all evidence of P. hagenella 
cell occupation (i.e., pupal exuviae, cocoon, 
presumed larvae) was found in nest cells 
with membranous linings, indicating an 
association with the cells reused by Hy- 
laeus, rather than the uncoated cells of 
Trypoxylon. Third, the presumed larvae of 
P. hagenella were clearly found in associa- 
tion with a Hylaeus pupa in the nest. 
Several of the larvae appeared to have 
their jaws successfully inserted into the 
cuticle of the bee pupa, and differences in 
the sizes of some of the larvae suggest that 
at least some had fed successfully. What 
actually killed the discovered bee pupae 
and mantispid larvae is unknown. Finally, 
a bee host for P. hagenella is consistent with 
the known bee hosts of at least two other 
members of the Plega melitomae species 
group (P. melitomae and yucatanae; see 
Table 1). In fact, the three-species interac- 
tion documented here - Plega hagenella 
parasitizing a Hylaeus (Hylaeopsis) species 
reutilizing a Trypoxylon manni nest - calls 

into question the accuracy of previous host 
records of Trypoxylon species for other P. 
melitomae group species (Table 1). The 
discovery of a mantispid-parasitized bee 
in a reutilized aculeate wasp nest makes it 
apparent that accurate host records for P. 
melitomae group mantispids cannot be 
inferred from published associations un- 
less the possibility of nest reuse is explicitly 

The observation that P. hagenella adults 
can be collected in sheltered, exposed-soil, 
situations favorable for Trypoxylon nesting 
provides a new focal point for field 
collecting Plega adults, which are rather 
rarely collected in tropical regions. It 
remains to be determined, however, 
whether these Plega individuals are pref- 
erentially targeting bees reusing Trypoxylon 
nests in such microhabitats, or whether 
they are attracted to such areas for general 
shelter and /or for access to a potentially 
larger array of subterranean-, surface- and 
aerial-nesting aculeate Hymenoptera that 
would likely be attracted to the same 
sites because of their suitability for nest- 

Because of the paucity of available host 
records, the degree of host-specificity of 
individual Plega species is currently un- 
clear, though the possible division of wild 
hosts proposed by Parker & Stange (1965) - 
i.e., P. melitomae group species on aculeate 
Hymenoptera and P. signata group species 
on a larger array of subterranean insects 
(e.g., larvae and /or pupae of Coleoptera, 
Lepidoptera and Diptera) - still appears as 
a broad generalization. It should be noted, 
however, that this generalization may not 
apply to captive individuals reared under 
artificial conditions (MacLeod and Red- 
borg 1982). 

All Mantispinae with known biologies 
have spider-associated larvae, and adults 
with highly r-selected reproductive strate- 
gies, each female producing hundreds to 
thousands of minute, stalked, eggs that are 
deposited in the environment with appar- 
ently little or no attempt to oviposit in sites 


Journal of Hymenoptera Research: Festschrift Honoring Roy Snelling 

that might increase the probability of 
emerging larvae encountering suitable 
hosts. The finding of Plega hagenella eggs 
directly deposited on a nest containing the 
immature stages of its host suggests that P. 
hagenella, and possibly other symphrasine 
mantispids, employs a different, more 
targeted, oviposition strategy that actively 
places eggs in closer proximity to potential 
hosts. Strategies such as this suggest 
the existence of more complex adult 
behaviors, particularly higher levels of 
host-searching ability in females. Within 
the Mantispidae, extraordinary intraspeci- 
fic and oviposition behaviors are also 
found in the symphrasine species Trichos- 
celia santareni, whose males and females 
engage in lekking behavior, followed by 
the females flying to, entering and ovipos- 
iting within active vespid nests (Dejean 
and Canard 1990). 


This paper is dedicated to the memory of Roy 
Snelling. Chris Starr and Carl Fitz-James graciously 
made their homes available during my stay in 
Trinidad, and the Department of Life Sciences, The 
University of the West Indies provided office and 
laboratory facilities. The Wildlife Section of the 
Forestry Division, St. Joseph, Trinidad issued collect- 
ing and export permits. Voucher specimens are 
deposited in the University of Texas and Texas 
A&M University insect collections. 


Almeida, E. A. B. 2008. CoUetidae nesting biology 
(Hymenoptera: Apoidea). Apidologie 39: 16-29. 

Buys, S. C. 2008. Observations on the biology of 
Anchieta fumosella (Westv^ood 1867) (Neuroptera: 
Mantispidae) from Brazil. Tropical Zoology 21: 

Dejean, A. and M. Canard. 1990. Reproductive 
behavior of Trichoscelia santareni (Navas) (Neu- 
roptera: Mantispidae) and parasitism of the 
colonies of Polybia diguetana R. du Buysson 
(Hymenoptera: Vespidae). Neuroptera Interna- 
tional 6: 19-26. 

Hoffman, K. M. and J. R. Brushwein. 1992. Descrip- 
tions of the larvae and pupae of some North 
American Mantispinae (Neuroptera: Mantispi- 
dae) and development of a system of larval 
chaetotaxy for Neuroptera. Transactions of the 
American Entomological Society 118: 159-196. 

Kuroko, H. 1961. On the eggs and first-instar larvae of 
two species of Mantispidae. Esakia. Occasional 
Papers of the Hikosan Biological Laboratory 3: 25-32. 

Lambkin, K. J. 1986. A revision of the Australian 
Mantispidae (Insecta: Neuroptera) with a con- 
tribution to the classification of the family. I. 
General and Drepanicinae. Australian Journal of 
Zoology, Supplementary Series 116: 1-142. 

Linsley, E. G. and J. W. MacSwain. 1955. Two new 
species of Plega from Mexico. Pan-Pacific Entomol- 
ogist 31: 15-19. 

MacLeod, E. G. and K. E. Redborg. 1982. Larval 
Platymantispine mantispids (Neuroptera: Plani- 
pennia): possibly a subfamily of generalist pred- 
ators. Neuroptera International 2: 37-41. 

Michener, C. D. 2007. Bees of the World. Johns Hopkins 
University Press, Baltimore, xvi + 557 pp. 

and R. W. Brooks. 2003. Dispersal of brood cells 

in a Mesoamerican hylaeine bee: a probable risk- 
spreading behavior (Hymenoptera, Colletidae). Pp. 
151-152 in: Melo, G. A. R., and I. A. dos Santos eds. 
Apoidea Neotropica: homenagem aos 90 anos de Jesus 
Santiago Moure. Editora UNESC, Criucuma, Brazil. 

Minter, L. R. 1990. A comparison of the eggs and first- 
instar larvae of Mucroberotha vesicaria Tjeder with 
those of other species in the families Berothidae 
and Mantispidae (Insecta: Neuroptera). Pp. 
115-129 in: Mansell, M. W. and H. Aspock eds. 
Advances in Neuropterology. Proceedings of the Third 
International Symposium on Neuropterology (3-4 
February 1988, Berg en Dal, Kruger National 
Park, South Africa). South African Department of 
Agricultural Development, Pretoria. 298 pp. 

Ohl, M. 2004. Annotated catalog of the Mantispidae of 
the World (Neuroptera). Contributions on Entomol- 
ogy, International. 5: 131-262. 

Parker, F. D. and L. A. Stange. 1965. Systematics and 
biological notes on the tribe Platymantispini 
(Neuroptera: Mantispidae) and the description 
of a new species of Plega from Mexico. Canadian 
Entomologist 97: 604-612. 

Penny, N. D. 1982. Neuroptera of the Amazon Basin. 
Part 6. Mantispidae. Acta Amazonica 12: 415-463. 

and C. A. da Costa. 1983. Mantispideos do 

Brasil (Neuroptera: Mantispidae). Acta Amazonica 
13: 601-687. 

Sakagami, S. F., N. Gobbi, and R. Zucchi. 1990. 
Nesting biology of a quasisocial sphecid wasp 
Trypoxylon fabricator. Japanese Journal of Entomol- 
ogy 58: 846-862. 

and R. Zucchi. 1978. Nests ofHylaeus (Hylaeopsis) 

tricolor: the first record of non-solitary life in colletid 
bees, with notes on communal and quasisocial 
colonies (Hymenoptera: Colletidae). Journal of the 
Kansas Entomological Society 51: 597-614. 

Vesey-FitzGerald, D. F. 1936. Nesting habits of 
Trypoxylon (Hym. Sphec.) from Trinidad. Proceed- 
ings of the Royal Entomological Society of London. 
Series /I 11: 111-115. 

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Urban, D. and J. S. Moure. 2007. Hylaeini Viereck, Werner, F. G. and G. D. Butler, Jr. 1965. Some notes on 

1916. Pp. 710-722 in: Moure, J. S., D. Urban, and the life history of Plega hanksi (Neuroptera: 

G. A. R. Melo eds. Catalogue of Bees (Hymenoptera, Mantispidae). Annals of the Entomological Society 

Apoidea) in the Neotropical Region. Sociedade of America 58: 66-68. 

Brasileira de Entomologia, Curitiba. xiv + 1058 Woglum, R. S. 1935. Symphrasis signata Hagen. Pan- 

pp. Pacific Entomologist 11: 119. 

Vol. 19(1), 2010, pp. 84-93 

Chartergellus golfitensis West-Eberhard: a new species of Neotropical 

swarm-founding wasp (Hymenoptera: Vespidae, Polistinae) with notes 

on the taxonomy of Chartergellus zonatus Spinola 

Mary Jane West-Eberhard, James M. Carpenter, Luiz Fernando F. Gelin, and 

Fernando B. Noll 

(MJW-E) Smithsonian Tropical Research Institute, c/o Escuela de Biologia, Universidad de Costa 

Rica, Costa Rica; injwe@sent.coin 
(JMC) Department of Entomology, American Museum of Natural History, Central Park West at 

79th Street, New York, New York 10024, U.S.A.; 

(LFFG) Departamento de Zoologia e Botanica, Instituto de Biociencias, Letras e Ciencias Exatas, Sao 

Paulo State University, Rua Cristovao Colombo, 2265 CEP 15054-000 Sao Jose do Rio Preto, SP, 


(FBN) Departamento de Zoologia e Botanica, histituto de Biociencias, Letras e Ciencias Exatas, Sao 

Paulo State University, Rua Cristovao Colombo, 2265 CEP 15054-000 Sao Jose do Rio Preto, SP, 


Abstract. — Chartergellus golfitensis West-Eberhard new species, is described from Central 
An\erica and compared with C. zonatus Spinola, a species heretofore inadequately described. 

Chartergellus is a genus of Epiponini 
(Hymenoptera, Vespidae, PoHstinae) rang- 
ing from southeastern Brazil to Costa Rica 
(West-Eberhard et al. 2006). The taxonomic 
background of Chartergellus is somewhat 
troublesome. It was described by Bequaert 
(1938) as a subgenus of Chartergus sensu 
Bequaert. The use of the name Chartergus 
was due to overlooking the type designa- 
tion for that genus, and Bequaert (1943) 
pointed out that the name Parachartergus 
von Ihering, 1904 was correct. Bequaert 
(1938) designated as type species Vespa 
frontalis Fabricius, 1804, the only species 
included in Chartergellus. Richards (1978) 
raised Chartergellus to genus, and pointed 
out that Vespa frontalis Fabricius was 
preoccupied by Vespa frontalis Latreille 
1802. He proposed C. amazonicus as a 
replacement name, and described five 
new species: C. atectus Richards 1978, C. 
communis Richards 1978, C. nigerrimus 
Richards 1978, C. punctatior Richards 

1978, and C. sanctus Richards 1978. He 
treated Chartergus zonatus Spinola 1851, as 
an unrecognized species and did not 
include it in his key, because the descrip- 
tion ''does not agree fully with any speci- 
mens I have seen of Chartergellus'' (Rich- 
ards 1978: 217). Most recently an eighth 
species has been added, C. afoveatus 
Cooper 1993. 

With the exception of C. atectus (see 
Richards 1978, and below), the species of 
Chartergellus build nests consisting of mul- 
tiple combs attached laterally to the sub- 
strate by pedicels and covered by an 
envelope of a single sheet, with the 
entrance a short spout (Wenzel 1998). The 
envelope may be irregular, with the lines of 
construction evident and contributing to 
camouflaging of the nest. Chartergellus is 
characterized by having a prominent 
curved bristle on the third labial palpo- 
mere, the maxillary palpi five-segmented 
and labial palpi three-segmented, lacking 

Volume 19, Number 1, 2010 


Figs 1-8. Chartergellus golfitensis West-Eberhard new species. Holotype. 1. Habitus. Lateral view. 2. Dorsal 
view. 3. Head, frontal view. 4. Head, lateral view. 5. Head, dorsal view. 6. Mesosoma, dorsal view. 7. Mesosoma, 
lateral view. 8. Scutellum, metanotum and propodeum, dorsal view. Scale bars = 1 mm. 

an occipital carina, the mesepisternum 
lacking a dorsal groove, and the metano- 
tum rounded (Carpenter 2004). 

Previously, only Chartergellus atectus was 
recorded for the Costa Rican fauna (Rich- 
ards 1978). We describe here Chartergel- 
lus golfitensis, a new species from Costa 

Chartergellus golfitensis West-Eberhard, 

new species 

(Figs 1-9) 

Diagnosis. — Chartergellus golfitensis is dis- 
tinguished by the presence of a raised 
black-lipped arc at the base of the man- 
dible; clypeus in short contact with eye; 
pale markings on the scutellum; frons with 
three-pointed reddish brown mark with 
the central point nearly reaching the 
median ocellus and in the intraocular space 
to the center of the ocular sinus; and a 
broad pale stripe along the entire length of 
the gena, usually without red or black 
region below, which when present is no 

wider than the diameter of the median 

Description. — Female. Holotype fore wing 
length 6.72 mm. 

Color. Black. Reddish brown: clypeus, 
first antennal segment and flagellum be- 
neath, malar space, lower 3/4 of the head, 
mandibles except teeth, fore tibia, tips of 
tarsi. Whitish anterior stripes on metano- 
tum curved at margin of propodeum, 
pronotal carina, and broadly along entire 
length of gena. Wings hyaline, venation 
brownish. Head. Eyes with short sparse 
hairs, touching clypeus for short distance. 
Clypeus wider than long (median length x 
width at middle = 0.74). Entire surface of 
head pubescent except for malar space and 
gena posterior to it, and clypeus below 
level of eyes, where there are long bristles. 
Frons with sparse and shallow punctures. 
Malar space present, about as wide as 
length of fourth antennomere. Gena with 
width equal to width of compound eye. 
Widest part of eye seen from front only 
half length of first antennomere. Vertex 


Journal of Hymenoptera Research: Festschrift Honoring Roy Snelling 

Figs 1-8. Continued. 

Figs 1-8. Continued. 

Volume 19, Number 1, 2010 


Figs 1-8. Continued. 

hairs present, sparser and shorter than 
those of frons which are nearly as long as 
bristles of cl3^eus below level of eyes. Base 
of mandible with raised rim, forming an 

arc with a black basal lip, dorsal third 
covered by lateral clypeal lobe. Mandible 
smooth, outer surface with a few long 
bristles. Mesosoma. Pronotum, scutum and 

Figs 1-8. Continued. 

Journal of Hymenoptera Research: Festschrift Honore^g Roy Snelling 

Figs 1-8. Continued. 

Figs 1-8. Continued. 

Volume 19, Number 1, 2010 


Figs 1-8. Continued. 

scutellum with fine, dense punctation and 
covered with straight hairs. Mesepister- 
num with scrobal furrow beginning at 
margin beneath tegula and curving poste- 
riorly to form a gentle arc about half as 
long as the mesepisternum, which is 
rounded anterior to the furrow. Dorsal 
pronotal carina well developed, ending 
close to the posterior margin. Pronotal 

fovea present. Propodeum without poster- 
ior concavity, with hairs on entire surface, 
propodeal valve narrow throughout. 


Color. As in female but metasoma (gas- 
ter) and legs dark brown. Reddish-brown 
of face fills entire ocular sinus and lacks 
three medial peaks on frons. Whitish genal 
stripe touching eye. Additional white: 

Fig. 9. Male genitalia of Chartergellus golfitensis. From left to right, aedeagus in ventral view, aedeagus in 
lateral view, digitus, cuspis and paramere. Scale bar = 1 mm. 


Journal of H^-mexopter-a Research: Festschrift Honoring Roy Snelling 

Stripe anterior to scrobal furrow, and 
coxae. Head. Mandibles with silvery ap- 
pressed hairs on basal half of outer surface, 
apical half with a few long bristles, under- 
surface covered with long hairs. Ch^eus as 
long as wide (L:W = 1.06), in extensive 
contact with eyes except for a short 
distance dorsally, and covered with silver}^ 
pubescence, ventrally forming a dense 
band of long appressed bristles which 
gleam in direct light as do similar hairs 
on lateral portions of frons, white portions 
of coxae, and a thin line on cur\^ed distal 
margin of last metasomal sternum. Malar 
space about half as wide as length of fourth 
antennomere. Gena only about half width 
of eye. Widest part of eye seen from front 
as long as first antennomere; saber-like 
ventral mandibular tooth longer and sharp- 
er-edged than in female. 

Male genitalia (Fig. 9). Aedeagus serrate 
beneath. Ventral process (at base of serrate 
portions of aedeagus) forming a "U" with 
outer end sharply pointed, as in C. commu- 
nis (Richards 1978, fig.88b). Serrations: 
proximal nine very fine and triangular 
(pointed), more distal 25 broader, curved 
on the posterior edge and pointing for- 
ward. Serrate edge slightly longer than 
distance from end of serrations to tip 
of aedeagus. Parameral spine hairless, 
slightly curved laterad, with elongate 
lateral opening near tip. Inner surface of 
digitus with long fine hairs, longer than 
width of digitus seen in profile, and shorter 
toward tip, which is bare. 

Distribution. — Known only from Pacific 
lowlands of Costa Rica. 

Habitat. — The tv\^o obser\'ed living colo- 
nies of C. golfitensis (the colony of the 
holotype and another from the same 
locality), and all other collected specimens, 
were found in areas of lowland humid 
forest. Nests were built on a vertical tree 
trunk and the wall of a building. 

Types.— Holotype, 9, COSTA RICA, Punta- 
renas Prov., Refugio de Vida Silvestre Golfito, 
83 11'55"W, 8-39'25"N, 500m from the sea level. 
2.i.2006 (L. Chavarria), deposited in the Mu- 

seum of Zoolog}' of the Uni\ersit\' of Sao Paulo, 
Sao Paulo, Brazil. Paratypes. In Museo de 
Zoologia, Escuela de Biologia, University of 
Costa Rica: Puntarenas Prov. 40 females, 17 
males, aU with the same locality and collector as 
the holot\^e. Golfo Duke, 24 km W. Piedras 
Blancas, alt. 200m, iii 1992, I9 (Paul Hanson); 
3 km SW Rincon, VI 1991, 2 99 (Paul Hanson); 
III-IV 1992, 19 (Paul Hanson). In Instituto 
Nacional de Biodiversidad (INBio), Costa Rica: 
Puntarenas Prov. Est. Agujas, Sendero Zamia, 
300m. 22-30 AGO 1996. L_S_276750_526550 
^8318, 19 (E. Retes, A. Azofeifa, M. Lobo); 
Albergue Cerro de Oro, 200m., 5-9 MAY 1995, L 
N 279650 ^4745, I9 (B. Gamboa); 4-14 May 
1995, L_S_280450_517500 #5919, I9 (E. Alfaro); 
Rancho Quemado, 200m., Die 1992, L-S 
292500_511000, I9 (F. Quesada); Abr 1992, I9 
(K. Hores); 299 (D. Brenes); lun 1992, I9 (F. 
Quesada); Est. Esquinas, Peninsula de Osa, 
200m. Abr 1993, L S 301400.542200 #2076, I9 
Q.F. Quesada); Est. Sirena, 0-lOOm, P.N. Cor- 
covado, Jun 1991, L-S270500_508300, I9 (G. 
Fonseca); P.N. Manuel Antonio, Quepos, 80m. 
Abr 1993, L S 370900_448800 #2140, !£ (G. 
Vaiela); Oct 1992, L-S 370900_448800, I9 (G. 

Etymology. — The name is given after the 
type locality, the Golfito region in Costa 
Rica, Central America. 

Remarks. — INBio collecting stations Est. 
(Estacion) Agujas, Est. (Estacion) Esquinas, 
Albergue Cerro de Oro, and Rancho 
Quemado are all in the Osa Peninsula of 
southwestern Costa Rica. 

The holot}''pe and paratypes were com- 
pared with paratypes of three other species 
of Chartergellus: C. punctatior Richards, C. 
communis, and C. zonatus Spinola. The 
parat}^pe of C punctatior Richards, labeled 
as such by Richards, is from the same nest 
as a specimen (COLOMBIA: Valle, Anchi- 
caya, 27.vii.73, M.J.W. Eberhard) mista- 
kenly listed by Richards (1978: 222) as a 
paratype of C. nigerrimus. Richards (1978) 
listed no specimens of C. punctatior from 
Colombia in the description, but included 
Colombia in the distribution of this species 
in his key, further indicating his intention 
to consider the Colombian specimen 
(which fits the description of punctatior) a 

VoLLAiE 19, Nl-mber 1, 2010 


paratype of this species. Besides this speci- 
men, two other Colombian (Anchicaya) 
specimens of C. punctatior (28.ix.76, M. J. 
West-Eberhard) were examined by Rich- 
ards in London. One was retained in the 
Natural History Museum, London, and the 
other, from the same colony, is in West- 
Eberhard's collection, labeled by Richards 
as a paratype of C. atectus. But it is clearly a 
specimen of C. punctatior that conforms in 
structure (mandibles with strong basal rim, 
and first metasomal tergum with broad 
yellow apical band) and nest type (un- 
exceptional for the genus: combs on a rock 
rather than leaves, covered by a full 
envelope) to the published description of 
punctatior, not to those of the holotype of C. 
atectus collected at the same locality and 
described as having a nest unusual for the 
genus in being constructed betw^een leaves 
and lacking a full envelope (Richards 1978). 
The paratype of C. communis examined here 
(BRAZIL: MT, Ilha de Bananal, Santa 
Isabella [recte: do Bananal, Santa Isabel], 
25.viii.68, W. D. Hamilton) was deposited in 
the West-Eberhard collection by Richards. 
Along with other West-Eberhard voucher 
specimens, these paratypes will eventually 
be deposited in the US National Museum, 
Smithsonian Institution, Washington DC. 
We also compared a paratype female of C. 
golfitensis wilh three female syntypes of the 
Spinola type series of C. zonatus in the 
collection of the Museo di Zoologia Siste- 
matica delTUniversita di Torino, now 
housed in the Museo Regionale di Scienze 
Naturali in Turin. 

Richards (1978) described three Charter- 
gellus species (amazonicus, punctatior, and 
communis) as having a ''raised rim" at the 
base of the manidible. In the communis 
paratype this is a rim-like ridge that rises 
abruptly forming a dark-brown-edged arc 
near the inner margin of the mandible, 
continued as a straight black line to the 
overlapping comer of the clypeus. In the 
punctatior parat\^es and in C. golfitejtsis the 
arc arises less abruptly and is not contin- 
ued as a line. 

Color variation. — ^The colony of the t\^pe 
specimen contained three queens and two 
readily distinguished age cohorts: young 
individuals (16 females and 17 males), 
distinguished by wide clear metasomal 
apodeme margins with only a very thin 
black marginal line, as is characteristic of 
recently emerged vespid adults (West- 
Eberhard 1975), and 22 older females 
(including the holotype), with wide black 
apodeme margins. The young females 
resembled the queens in being lighter in 
color: meso- and metasoma and some parts 
of legs that are black in the older females 
are dark to light brown in the young 
females and queens, suggesting that the 
recently emerged females, although lack- 
ing sperm in the spermatheca and with 
undeveloped ovaries, are young queens. 
Queens have light brown first antenno- 
mere and nearly white mandibles. White 
lines at the posterior margin of the prono- 
tum and the scrobal furrow are present in 
some females, absent in some (including 
the holotype), and a brownish spot in 
some, variation not correlated with age or 
caste. A white stripe on the distal margin of 
the first metasomal tergum is complete or 
absent in some, or at the lateral margins 
only in most females. A detailed morpho- 
metric analysis of both sexes and castes 
will be published elsewhere. 

The Sendero Zamia, Rancho Quemado, 
and Manuel Antonio specimens have dark 
brown rather than reddish brown facial 
markings, and light markings are brown or 
yellowish brown rather than whitish. 


Spinola (1851) referred to four female 
specimens collected by M. Ghiliani in Para, 
with the male unknowm, but only three 
specimens are present in the Turin collec- 
tion and there is no record of the location 
of the missing female. Examination of the 
remaining specimens indicate that Rich- 
ards' (1978: 218) doubts regarding the 
Spinola (1851) description of C. zonatus. 


Journal of Hymenoptera Research: Festschrift Honoring Roy Snelling 

resulting in its status as incertae sedis 
(Carpenter and van der Vecht 1991; Cooper 
1993), are largely unjustified, though a few 
inaccuracies were evident (see below). 
Richards stated that ''the description of 
. . .Chartergus zonatus does not agree fully 
with any specimens I have seen of Charter- 
gellus" but the Spinola specimens accord 
with Richards' (1978) generic description 
for Chartergellus in having the propodeal 
valve linear rather than in-curving, prono- 
tal fovea present, widely separated from 
carina; hind-wing vein cu-a much longer 
than 1 Cui; labial palpi with three seg- 
ments (the maxillary palpi could not be 
seen); scrobal furrow strong; margin of 
mesoscutum opposite tegula strong; scu- 
tellum rounded in profile; metanotum not 
vertical; and occipital carina lacking. 
Species level characters are as follows: 
Color. Black. Dark brown: antennae 
(lighter brown at tip and beneath), legs 
and wing venation. Reddish- brown: cly- 
peus and three-pointed area of frons to 1/3 
distance from antennal insertions to me- 
dian ocellus. Yellow: intraocular space to 
ocular sinus; broad genal stripe (about half 
width of gena) from mandible nearly to 
dorsal margin of eye; propodeal carina; 
stripe anterior to scrobal furrow and about 
Va its length; anterior margins of scutellum, 
metanotum; thin stripes on margins of all 
sclerites bordering mesoscutum; posterior 
margins of terga 1, 2 (one female) or terga 
1, 2, 3 (two females). Other variation: one 
female, labeled with a small green tag 
marked "9," is darker in color than the 
other two: all of the markings that are 
yellow in the other two specimens and in 
the Spinola description, are brown, and all 
brown markings are darker brown in this 
specimen, which is one of those with three 
metasomal stripes. A female labeled "Pun- 
gono fortemente" (stung strongly) was 
affected by mold, so the remaining speci- 
men, with no special label, and having 
yellow coloration like that described by 
Spinola, was used for the description 
reported here. 

Other characters: eyes with sparse short 
hairs; mandible with four teeth, base with 
raised basal arc, scutal punctures separated 
by one diameter or more, scrobal furrow rises 
to gentle ridge at anterior side, coarsely and 
irregularly punctuate, with appressed hairs 
on posterior part and striate anterior ridge. 

As suspected by Richards regarding a 
strongly yellow-marked species, the man- 
dibles are not black; they are brown, with 
only the teeth dark brown or black, in- 
cluding in the darkest of the three speci- 
mens seen. The scutal margins are not 
yellow, as stated by Spinola; instead, it is 
the margin of the adjacent sclerites that are 
yellow; and only about half of the metano- 
tum is yellow, not the entire metanotum. 

Spinola (1851) cited Ghiliani's notes as 
stating that the nest of this species, in 
contrast to that of Chartergus nitidulans 
Fabricius, is found on the trunks of trees, 
and that the sting is very painful. Spinola 
was sent a sample of the nest carton, which 
he described as "teme et grisatre," (dull 
and grayish) but this has evidently been 
lost. There is no holotype labeled, hence all 
three specimens examined are syntypes. 


The authors thank the Ministerio de Ambiente, 
Energia y Telecomunicaciones de Costa Rica (MIN- 
AET) for the collecting license, all the personnel of the 
Refugio de Vida Silvestre Golfito for collaboration, the 
Universidad de Costa Rica for hospitality in Golfito, 
and Jose Chaves Jimenez for help during the collection 
of colonies made by L. Chavarria in Costa Rica. We 
also thank J. Diniz for helping with the dissection of 
male genitalia. MJWE thanks Prof. Maria Cristina 
Lorenzi, Guido Pagliano, and Dr. Stefania Fucini for 
help and hospitality during work on the Spinola 
specimens in Turin; and Dr. Paul Hanson of the 
University of Costa Rica and Manuel Solis of the Costa 
Rican National Biodiversity Institute (INBio) for help 
with use of collections at their institutions. 


Bequaert, J. C. 1938. A new Charterginus from Costa 
Rica, with notes on Charterginus, Pseudochartergus, 
Pseudopolybia, Epipona, and Tatua (Hymenoptera, 
Vespidae). Revista Entomologia Rio de Janeiro 9: 
99-117, 1 pi., Ifig. 

Volume 19, Number 1, 2010 


. 1943. The generic names Chartergus, Parachar- 

tergus, Epipona and Tatua. A correction (Vespidae, 
Hymenoptera). Bulletin Brooklyn Entomological 
Society 38: 4. 

Carpenter, J. M. 2004. Synonymy of the genus Mar- 
imbonda Richards, 1978, with Leipomeles Mobius, 
1856 (Hymenoptera: Vespidae; Polistinae), and a 
new key to the genera of paper wasps of the New 
World. American Museum Novitates 3465: 1-16. 

and J. van der Vecht. 1991. A study of the 

Vespidae described by William J. Fox (Insecta: 
Hymenoptera), with assessment of taxonomic 
implications. Annals of the Carnegie Museum 60: 

Cooper, M. 1993. A new species of Chartergellus (Hym., 
Vespidae, Polistinae, Polybiini) from Bolivia. En- 
tomologist's Monthly Magazine 129: 165-166. 

Fabricius, J. C. F. 1804. Systema Piezatorum. 439 pp. 

Ihering, H. von. 1904. As vespas sociaes do Brasil. 
Revista do Museu Paulista 6: 97-309, 5pls, 3 figs. 

Latreille, P. A. 1802. Historic naturalle, generale et 
particulere des Crustacees es des Insectes. (Sonnini's 
suites a Buffon), Paris. 3 xxi + 468 pp. 

Richards, O. W. 1978. The Social Wasps of the Americas, 
Excluding the Vespinae. British Museum (Natural 
History), London. 580 pp + 4 pp plates. 

Spinola, M. 1851. Compte rendu des Hymenopteres 
inedits provenant du voyage entomologique de 
Ghiliani dans le Para en 1846, 3-78. [Preprint of 
Memorie della Reale Accademia della Scienze di 
Torino (2)13: 19-94,1853 [Spinola (1853) is avail- 
able at 

Wenzel, J. W. 1998. A generic key to the nests of 
hornets, yellowjackets, and paper wasps world- 
wide (Vespidae: Vespinae, Polistinae). American 
Museum Novitates 3224: 1-39. 

West-Eberhard, M. J. 1975. Estudios de las avispas 
sociales (Himenoptera, Vespidae) del Valle del 
Cauca. I. Objetivos, metodos y notas para facilitar 
la identificacion de especies comunes. Cespedesia 4 
(16): 245-267. 

, J. M. Carpenter, and P. E. Hanson. 2006. 

Familia Vespidae. 617-644 in: Hanson, P. E., and 
I. D. Gauld eds., Hymenoptera de la Region 
Neotropical. Memoirs of the American Entomological 
Institute 77: 1-994. 

Vol. 19(1), 2010, pp. 94-112 

The Description of Banacuniculus Buffington, New Genus 
(Hymenoptera: Figitidae: Eucoilinae) 

M. L. Buffington 

Systematic Entomology Lab, ARS/USDA, c/o Smithsonian Institution, National Museum of 
Natural History, 10*^ «& Constitution Ave PO Box 37012, MRC-168, NW, Washington DC 20013;;; 

Abstract. — The new eucoiline genus Banacuniculus is described to accommodate several species 
previously placed in Ganaspidium: Banacuniculus hunteri (Crawford), new combination; B. 
merickeli (Miller), new combination; B. nigrimanus (Kieffer), new combination; B. utilis 
(Beardsley), new combination; these species are all redescribed. The following new species are 
described: Banacuniculus brautigani, B. beardsleyi, B. dis, and B. strykeri. As is the case with 
Ganaspidium, species of Banacuniculus are parasitoids of some of the most pestiferous species of leaf- 
mining Agromyzidae (Diptera) and have been investigated for use as biological control agents of 
Liriomyza trifolii (Burgess). Additional phylogenetic, host range, and distributional data, and a key 
to all species are provided. 

Key words. — Liriomyza, Agromyzidae, new species, new genus, species revision, parasitoid. 

While revising Ganaspidium Weld (Hy- 
menoptera: Figitidae: Eucoilinae), Buffing- 
ton (2010) determined that some species of 
eucoiline v^asps that had been historically 
included w^ithin Ganaspidium would render 
that genus polyphyletic if they remained 
within the genus. Buffington et al. (2007) 
supported this observation, where Ganas- 
pidium pussillae Weld was recovered as a 
separate clade from G. hunteri (Crawford) 
and G. utilis Beardsley. Buffington (2010) 
further examined the characters of the 
mesosoma in both G. hunteri and G. utilis, 
revealing additional data supporting the 
removal of these two species from Ganas- 
pidium. Buffington (2010) placed these two 
species, as well as G. merickeli (Miller) and 
G. nigrimanus (Kieffer), as incertae sedis 
pending the erection of a new genus to 
accommodate them. This paper provides 
the description of Banacuniculus Buffing- 
ton, new genus, to accommodate these 
species. Included in this description is a 
redescription of the previously named 

species Banacuniculus hunteri (Crawford), 
new combination; B. merickeli (Miller), new 
combination; B. nigrimanus (Kieffer), new 
combination; and B. utilis (Beardsley), new 
combination. Four new species are also 
described: Banacuniculus brautigani Buf- 
fington, B. beardsleyi Buffington, B. dis 
Buffington, and B. strykeri Buffington. 

Species of Banacuniculus are koinobiont 
endoparasitoids of agromyzid fly larvae 
(Diptera: Agromyzidae). Beardsley (1986, 
1988) provided the first taxonomic work 
on eucoiline parasitoids of agromyzids in 
Hawai'i, and associated host remains with 
several species that he considered Ganaspi- 
dium, chiefly G. hunteri and G. utilis. The latter 
was described in Beardsley (1988) and has 
been shown to be an instrumental species in 
controlling pestiferous Liriomyza Qohnson 
1987; Lynch and Johnson 1987; Mason and 
Johnson 1988; Rathman et al. 1991, 1995). 

Banacuniculus is nested within the Gro- 
notoma genus group (Buffington et al. 2007) 
and shares several synapomorphies with 

Volume 19, Number 1, 2010 


those genera, including a complete para- 
scutal impression, a large scutellar plate 
with a centrally placed midpit, protuber- 
ance on the cl3^eal and malar spaces, and 
remnants of notauli on the mesoscutum. 
Only the clypeal and malar protuberances 
are shared with Ganaspidium; all the other 
features are shared with other Gronotoma 
group genera (further discussed below). 

List of depositories 

BPBM -Bernice B. Bishop Museum, 

Honolulu, HI, USA (N. Even- 

CASC -California Academy of Sci- 

ences, San Francisco, CA, USA 
(R. Zuparko). 

CNCI -Canadian National Collection 

of Insects, Ottawa, Canada (J. 

KSCU -Kansas State University, Man- 

hattan, KS (G. Zolnerowich). 

TAMU -Texas A&M University Insect 
Collection, College Station, TX, 
USA (E. Riley). 

UCDC -Bohart Museum, University of 
California at Davis, Davis, CA, 
USA (L. Kimsey). 

UCRC -University of California at 
Riverside, Riverside, CA, USA 
(S. Triapytsin). 

USNM -National Museum of Natural 
History, Smithsonian Institu- 
tion, Washington, D.C., USA 
(M. Buffington). 

Additional sources of specimens. — Exten- 
sive collections from Texas were made 
available for my examination by Ricardo 
Hernandez (Department of Entomology, 
TAMU) and from Kansas by Robert Kula 
(Systematic Entomology Laboratory, Wash- 
ington, DC) and Gregory Zolnerowich (Kan- 
sas State University, Manhattan, KS). 

Specimen illustration and observation. — 
Methods for the imaging of specimens 
using light microscopy follow those of 

Buffington and van Noort (2007, 2009). 
For specimens too small for light micro- 
scopy, an environmental scanning electron 
microscope (ESEM) was used to image 
characters; specimens were shot uncoated 
in a Philips XL-30 ESEM machine, with 
lanthanum hexaboride electron source 
(LaB6). The instrument was operated in 
low vacuum mode with water vapor as the 
imaging gas and backscatter imaging with 
one-half of the diode active. Specimen 
mounting and lighting techniques follow 
Buffington and Gates (2009). Slide mounts 
were prepared with PVA mounting med- 
ium, cured for 71 hours at 43 °C, and 
photographed with a JVC KW-75C camera 
on a Leica DMRB compound microscope. 
Specimens were examined with a Leica 
MIO stereomicroscope, illuminated with a 
pair of florescent desk lights. 

Descriptive format. — Diagnoses focus on 
features that are easily recognizable by 
other observers, and closely related species 
that may have similar gross morphologies 
are distinguished. Terminology for all 
descriptive characters as well as phyloge- 
netic characters are defined in Buffington 
(2009) and are not repeated here; sur- 
face sculpture terminology follows that of 
Harris (1979). Following the descriptions 
are summaries of general distribution, 
biology and comments on nomenclatural 
issues (when applicable). The species de- 
scriptions are generated by a database 
application, vSysLab (Johnson 2008), de- 
signed to facilitate the generation of taxon 
by character data matrices and to export 
the data both as text and as input files for 
other applications. A link to a distribution 
map is included in each species descrip- 
tion. New species are designated are 
accompanied by a ZooBank registration 


Banacuniculus Buffington, new genus 


Journal of Hymenoptera Research 

Type species: Banacunictdtis Imnteri 
(Crawford), by present designation. 

Diagnosis. — Malar space and ventral cly- 
peal margin with distinct conical protuber- 
ances. Notauli lacking, setal tracks present 
where notauli are located in related taxa. 
Parascutal impression complete, terminat- 
ing anteriorly at the point of origin of the 
notaular setal tracks. Scutellar plate nearly 
circular with midpit located in center of 
plate; plate lacking tubercles; setae-bearing 
pits present, encircling midpit. Setal band 
at base of syntergum of metasoma com- 
plete. Similar to Ganaspidium but easily 
separated based on the morphology of the 
scutellar plate: Ganaspidium always has a 
pair of tubercles on the anterior half of the 
plate, and the plate is not circular but 
oblong. Banacuniculus is also similar to 
Microstilba, Nordlanderia, and Disorygma, 
but distinguished based on notauli being 
absent and the hairy ring of syntergum 
present (notauli present and hairy ring of 
metasoma absent in Microstilba, Nordlan- 
deria, and Disorygma); similar to Agrostocy- 
nips, but with pronotol plate less than 
one-half the width of the head (nearly as 
wide as head in Agrostocynips) and genal 
carina absent (present ventrally in Agrosto- 

Description. — Head. Nearly glabrous with 
a few scattered setae on lower face, 
clypeus, inner margins of compound eyes 
and gena; ocellar hair patches absent. 
Ventral 1/4 of lower face with admedial 
clypeal furrows converging towards the 
clypeus; point of convergence resulting in 
the formation of distinct conical protuber- 
ance protruding from anterior margin of 
clypeus. Orbital furrows absent. Malar 
sulcus ranging from simple to compound. 
Malar space smooth to distinctly strigose, 
with large conical protuberance present. 
Genal carina absent. 

Antennae: Female: 13 segments, monili- 
form, clavate; segments 3-13 sub-equal in 
length; rhinaria present only on the last 7 
segments. Male: 15 segments; segments 3- 

15 sub-equal in length; rhinaria present on 
segments 3-15. Segment 3 modified, 
curved outwardly, excavated laterally. 

Pronotum: Pronotal plate narrow, with 
setae present along dorsal margin; dorsal 
margin rounded; pronotal fovea open. 
Lateral pronotal carina absent. Pronotal 
triangle absent. Pronotal impression ab- 
sent. Lateral aspect of pronotum smooth, 
glabrous in most species. 

Mesoscutum: Smooth and glabrous; no 
sculpture present. Parascutal impression 
complete, lined with setae, extending from 
the tegula to point along anterior margin of 
mesoscutum where notaular setal line 
originates. Notauli, mesoscutal keel, para- 
psidal ridges and parapsidal hair lines 
absent; notaular setal line present. 

Mesopectus: Upper and lower part of 
mesopleuron completely smooth and glab- 
rous. Mesopleural triangle present, faintly 
indicated (often only visible in the space 
immediately anterior to the mesopleural 
spiracle). Mesopleural carina simple. Pre- 
coxal carina of lower part of mesopleuron 
present anteriorly and posteriorly, absent 
ventrally. Surcoxal depression reduced, 

Scutellum: Scutellar plate ranging from 
small to large; midpit centrally placed; rim 
of plate translucent; dorsal surface of plate 
smooth in most species; setae-bearing pits 
present around midpit. Dorsal surface of 
the scutellum reticulate to smooth; 
rounded posteriorly and laterally; poster- 
ior carina present or absent. Laterodorsal 
and posterior projections absent. Lateral 
bars as long as wide; ventral lobe absent. 
Scutellar fovea oval, smooth and deep. 

Metapectal-propodeal complex: Posterior 
1/3-1/4 of metapectus setose. Spiracular 
groove with a well-defined dorsal margin, 
reduced ventral margin. Posterior margin 
of metapectus smooth, not ridged. Meta- 
pleural ridge and submetapleural ridge 
absent. Anterior impression of metepi- 
meron absent; anterior impression of me- 
tepisternum present, reduced. Anteroven- 
tral cavity rounded, setose. Propodeum 

Volume 19, Number 1, 2010 


covered in dense, appressed setae. Lateral 
propodeal carinae semi-parallel, bowed at 
junction with auxiliary propodeal carinae; 
auxiliary propodeal carinae distinct. Nu- 
cha glabrous, reticulate. 

Wings: Hyaline; setose. Ri complete, 
pigmented along anterior margin of wing; 
marginal cell not truncate, as deep as long. 
Apical fringe present, short (Fig. 3C). 

Legs: Fore and mid coxa subequal in size, 
hind coxa twice the size of either fore or 
mid coxa. Fore coxa variously setose; mid 
and hind coxa with distinct anterior and 
posterior dorsoventral setal bands. Femora 
with sparse setal lines; tibiae and tarsome- 
res with dense, appressed setae. Length of 
hind tarsomere 1 equal to 0.5 X the com- 
bined length of remaining hind tarsomeres. 

Metasoma: Female: Subequal in size to 
mesosoma. Base of syntergum with hairy 
ring present, comprised of dense, ap- 
pressed setae and a ring of thin, erect 
setae; remainder of metasoma glabrous. 
Micropunctures present on posterior 1/3 of 
the syntergum, and on remaining terga. 
Terga posterior to syntergum gradually 
directed ventrally, resulting in a 70 degree 
angle between S3mtergum and remaining 
terga. Ovipositor with series of sub-apical 
serrations (seen only in large specimens). 
Male: as in female with the terga posterior 
to syntergum abruptly angled ventrally, 
resulting in a 90 degree angle between 
syntergum and remaining terga. 

Distribution. — Neotropical Region: Chile, 
Argentina, Panama, Costa Rica, Southern 
Mexico; Nearctic Region: Northern Mexico, 

continental United States, southern Can- 
ada; Hawaiian Islands. 

Biology. — Several species of Liriomyza 
have been recorded as hosts (Beardsley 
1986; Johnson 1987; Hara and Matayoshi 
1990; Acosta and Cave 1994; present 
study). Banacuniculus hunteri and B. utilis 
have been evaluated for their usefulness in 
biological control of pestiferous leaf- 
mining flies (Johnson 1987; Lynch and 
Johnson 1987; Mason and Johnson 1988; 
Rathman et al. 1991; Rathman et al. 1995). 
Petcharat and Johnson (1988) studied the 
larval stages of Banacuniculus utilis. 

Etymology. — The name translates in Latin 
to 'miner hunter': bana, hunter; cuniculus, 
miner. The name refers to the host pref- 
erence of species of Banacuniculus, which 
putatively all attack leaf mining agromyzid 
flies. The gender is masculine. 

Included species. — 
Banacuniculus heardsleyi Buffington, new 

B. brautigani Buffington, new species. 
B. dis Buffington, new species. 
B. hunteri (Crawford), Beardsley (1986). Eucoila 

hunteri Crawford, 1913: 310, holotype in 

USNM. Ganaspidium hunteri (Crawford): 

Beardsley (1986, 1988), Buffington (2004, 2010). 
B. merickeli (Miller), new combination. Nordlan- 

deria merickeli Miller, 1989: 158-159, 162, 

holotype lost. 
B. nigrimanus (Kieffer). Eucoela nigrimanus Kief- 

fer, 1907:138, holotype in CASC Ganaspidium 

nigrimanus (Kieffer): Buffington (2004). 
B. strykeri Buffington, new species. 
B. utilis (Beardsley). Ganaspidium utilis Beardsley, 

1988: 44-46, holotype in BPBM. 


Dorsal surface of scutellum completely smooth and glabrous (Figs 2F and 3B); 
posterior margin of scutellum lacking a carina (Fig. 2F) 2 

Dorsal surface of scutellum distinctly sculptured, ranging from rugulose to striate 
(Fig. ID); posterior margin of scutellum with (Fig. IE) or without (Fig. ID) a 
distinct carina 4 

In lateral view, scutellar plate extending to posterior margin of scutellum (Fig. 3B); in 
dorsal view, the scutellar plate completely obscures the dorsal surface of the 
scutellum 3 


Journal of Hymenoptera Research 

In lateral view, scutellar plate not extending to the posterior margin of scutellum 
(Fig. 2E); in dorsal view, much of the dorsal surface of the scutellum visible . . . 
B. nigrimanus (Kieffer) 

3. Dorsal surface of the scutellar plate horizontally strigose anterior of scutellar 

midpit B. heardsleyi, n. sp. 

Dorsal surface of scutellar plate entirely smooth anterior to the scutellar midpit . . . 

B. utilis (Beardsley) 

4. Posterior margin of the scutellum without a well-developed and distinct posterior 

carina, resulting in a broadly rounded posterior margin of the scutellum, lack- 
ing a distinct transition from the dorsal surface to the latero-postero surface 

(Fig. ID) 5 

Posterior margin of the scutellum with a well-developed and distinct posterior carina, 
separating the dorsal surface of the scutellum from the latero-postero surface (PC, 
Fig. IE) 6 

5. Lateral aspect of pronotum and ventral half of mesopleuron covered with long, thin, 

white setae (PHP, Fig. IB); metasoma at least twice as long as mesosoma .... 

B. brautigani n. sp. 

Lateral aspect of pronotum and ventral half of mesopleuron largely glabrous (in some 
specimens, there may be short setae present along the anterior margin of the 
pronotum) (Fig. IC) B. dis, n. sp. 

6. Dorsal surface of scutellum distinctly rugulose to coriaceous over its entirety (Fig. 2 A) 7 
Dorsal surface of scutellum longitudinally striate, frequently smooth and glabrous 

anteriorly just posterior to the scutellar fovea B. merickeli (Miller) 

7. Scutellar plate flat to slightly convex surrounding the midpit (Fig. 2A) 

B. hunteri (Crawford) 

Scutellar plate distinctly concave surrounding the midpit (Fig. 2C) . . B. strykeri, n. sp. 

Banacuniculus heardsleyi Buffington, new 

Fig. IF. 

Description. — Malar sulcus simple. Malar 
space smooth. Malar protuberance smooth, 
short, not extending beyond length of 
ventral margin of malar space. Clypeal 
protuberance short, not overhanging ante- 
rior margin of clypeus. Tubercles of scu- 
tellar plate absent. Dorsal surface of scu- 
tellar plate flat, horizontally striate, setal 
bearing pits present surrounding midpit. 
Carina along posterior margin of scutellum 
absent. Dorsal surface of scutellum entirely 
smooth. Midpit of scutellar plate in center 
of plate; plate large, obscuring dorsal sur- 
face of scutellum v/hen viev^ed dorsally. 

Mesopopleural setal patch absent. Meso- 
pleuron entirely smooth. Lateral aspect of 
pronotum w^ith some short setae anteriorly, 
remainder glabrous. Marginal cell of 
forew^ing as deep as long. 

Metasoma sub-equal in size to meso- 
soma (in lateral viev^). 

Diagnosis. — This species resembles both 
B. nigrimanus and B. utilis in the morphol- 
ogy of the dorsal surface of the scutellum, 
but is easily separated from these species 
by the presence of the horizontally striate 
scutellar plate 

Etymology. — Named in honor of my late 
mentor Jack Beardsley. 

Link to Distribution Map. — http//hol.osu. 

Biology. — Unknown. 

Type Material. — Holotype, female: MEXICO: 
Mexico, SIN, Mazatlan, 27.III.1979, L. D. French, 
USNM ENT 00655548 (deposited in UCDC). 

Volume 19, Number 1, 2010 


Banacuniculus brautigani Buffington, 
new species 

Figs 1 A and B. 

Description. — Malar sulcus compound. 
Malar space partially striate, striations 
extending 1/2 to 2/3 distance from ventral 
margin of malar space to base of com- 
pound eye. Malar protuberance smooth, 
elongate, extending beyond length of ven- 
tral margin of malar space. Clypeal protu- 
berance elongate, overhanging anterior 
margin of clypeus. Tubercles of scutellar 
plate absent. Dorsal surface of scutellar 
plate gently convex, smooth; setal bearing 
pits surrounding midpit. Carina along 
posterior margin of scutellum absent. 
Dorsal surface of scutellum entirely rugu- 
lose /wrinkled. Midpit of scutellar plate in 
center of plate; plate small, revealing 
dorsal surface of scutellum when viewed 
dorsally. Mesopopleural setal patch pre- 
sent. Mesopleuron entirely smooth. Lateral 
aspect of pronotum covered in long, thin, 
white setae. Marginal cell of forewing as 
deep as long. Metasoma distinctly larger 
than mesosoma (longer and deeper) when 
viewed laterally. 

Diagnosis. — This species differs from all 
other Banacuniculus by the presence of an 
entirely setose lateral aspect of the prono- 
tum and the presence of a metasoma that is 
roughly two times the size of the meso- 
soma (PHP, Fig. IB). 

Etymology. — Named in honor of the 
author Richard Brautigan who wrote many 
poems and short stories on the nature and 
culture of California. 

Link to Distribution Map. — http://hol. 

Biology. — Unknown. 

Type Material. — Holotype, female: UNITED 
STATES: CA, Riverside Co., 3.5mi S Palm 
Desert, Nance (Coyote) Canyon, alluvial soil, 
Philip L. Boyd Deep Canyon Research Center, 

12.IV.1975, J. D. Pinto, USNM ENT 00655348 
(deposited in UCRC). 

Banacuniculus dis Buffington, new spe- 

Figs 1 C-D and 3 E-F. 

Description. — Malar sulcus compound. 
Malar space partially striate, striations ex- 
tending 1/2 to 2/3 distance from ventral 
margin of malar space to base of compound 
eye. Malar protuberance smooth, elongate, 
extending beyond length of ventral margin 
of malar space. Clypeal protuberance elon- 
gate, overhanging anterior margin of cly- 
peus. Tubercles of scutellar plate absent. 
Dorsal surface of scutellar plate gently 
convex, smooth; setal bearing pits sur- 
rounding midpit. Carina along posterior 
margin of scutellum absent. Dorsal surface 
of scutellum entirely rugulose /wrinkled. 
Midpit of scutellar plate in center of plate; 
plate small, revealing dorsal surface of 
scutellum when viewed dorsally. Mesopo- 
pleural setal patch absent. Mesopleuron 
entirely smooth. Lateral aspect of pronotum 
with some short setae anteriorly, remainder 
glabrous. Marginal cell of forewing as deep 
as long. Metasoma sub-equal in size to 
mesosoma in lateral view. 

Diagnosis. — This species can be confused 
with Banacuniculus hunteri, but can be 
separated by the lack of a posterior carina 
of the dorsal surface of the scutellum. 

Etymology. — Named after Dis, Dante 
Alighieri's name for the City of Hell in 
The Inferno. As used here, the name refers 
to the hell-like conditions this species 
seems to thrive in the desert Southwest. 

Link to Distribution Map. — http://hol. 

Biology. — ^Unknown. 

Type Material. — Holotype, female: UNITED 
STATES: CA, Imperial Co., 5km from Gordons 
Well exit at 1-8, Sand Highway, Algodones 


Journal of Hymenoptera Research 

Fig. 1. A, Banacuniculus brautigani, n. sp., lateral habitus of holotype; B, B. braiitigani n. sp., closeup of head and 
mesosoma; C, B. dis n. sp., lateral habitus; D, B. dis n. sp., poster-dorsal view of scutellum and scutellar plate; E, 
B. merickeli (Miller), closeup of head and mesosoma; F, B. heardselyi n. sp., lateral habitus of holotype. 
Abbreviations: PHP, pronotal hair patch; PC, posterior carina of the scutellum. 

Dunes, 32=45.5'N 114'57'W, 24.VII.2008, S. L. 
Heydon, USNM ENT 00655716 (deposited in 
UCDC). Paratypes: (2 females, 3 males) UNIT- 
Big Pine, 24.V.1994, S. L. Heydon (1 female, 
USNM ENT 00655523 (UCDC)); Inyo Co., 
Eureka Valley, Joshua Flat, 24.V.1994, S. L. 
Heydon (1 male, USNM ENT 00655522 

(UCDC)); Riverside Co., 2mi NW Oasis, 
15.V.1974, N. J. Smith (1 female, USNM ENT 
00655486 (UCDC)); Solano Co., 6km SE Suisun 
City, Suisun Marsh, 14. VI. 1993, S. L. Heydon & 
L. Guo (1 male, USNM ENT 00655454 (UCDC)). 
MONTANA. Silver Bow Co., Butte, 23.VII.1983, 
J. D. Pinto (1 male, UCRC ENT 196933 

Volume 19, Number 1, 2010 


Banacuniculus hunteri (Crawford), new 

Eucoila hunteri Crawford, 1913: 310. Holotype 
in USNM. 

Ganaspidium hunteri (Crawford): Beardsley 
(1986), Buffington (2010). 

Figs 2 A and B. 

Description. — Malar sulcus compound. 
Malar space partially striate, striations ex- 
tending 1/2 to 2/3 distance from ventral 
margin of malar space to base of compound 
eye. Malar protuberance striate, elongate, 
extending beyond length of ventral margin 
of malar space. Clypeal protuberance elon- 
gate, overhanging anterior margin of cly- 
peus. Tubercles of scutellar plate absent. 
Dorsal surface of scutellar plate flat, 
smooth; setal bearing pits present sur- 
rounding midpit. Carina along posterior 
margin of scutellum delicate, defining 
transition from dorsal surface of scutellum 
from posterior surface. Dorsal surface of 
scutellum entirely rugulose/ wrinkled. Mid- 
pit of scutellar plate in center of plate; plate 
small, revealing dorsal surface of scutellum 
when viewed dorsally. Mesopopleural setal 
patch absent. Mesopleuron entirely smooth. 
Lateral aspect of pronotum with some short 
setae anteriorly, remainder glabrous. Mar- 
ginal cell of forewing as deep as long. 
Metasoma subequal in size to mesosoma 
in lateral view. 

Diagnosis. — This species is sepa- 
rated from all other Banacuniculus by 
simultaneously possessing a densely cre- 
nulate dorsal surface of the scutellum 
and a distinct posterior carina of the 

Link to Distribution Map. — http://hol. =253204 

Biology. — Recorded parasitizing Lirio- 
myza trifolii (Burgess) and L. sativae Blanch- 
ard (Lynch and Johnson 1987). 

Material Examined. — Holotype, female: UNI- 
TED STATES: TX, Dallas Co., cotton, Dallas, 
19.V.1912, Hunter, USNM ENT 00655719 (de- 

posited in USNM). Paratype: UNITED STATES: 
TX, Dallas Co., cotton, Dallas, 19.V.1912, Hunter 
(1 female, USNM ENT 00653537 (USNM)). Other 
material: (23 females, 12 males) CANADA: 2- 
25Y Bm-91, no date (1 female, USNM ENT 
00652564 (CNCI)). ALBERTA. Calgary, 1. 
VII.1956, O. Peck (1 female, USNM ENT 
00655545 (CNCI)); Lethbridge, Oldman River, 
22.VI.1956, O. Peck (8 females, 2 males, 
USNM ENT 00652563, 00653555, 00653558, 
00653559, 00653560-00653562, 00653566, 00653568, 
00653569 (CNCI)); Lethbridge, Oldman River, 
22.VI.1956, sweeping, O. Peck (1 female, USNM 
ENT 00653570 (CNCI)); grass, Elkwater Lake, 
21.VII.1956, O. Peck (2 males, USNM ENT 
00653567, 00653578 (CNCI)); montaine prairie, 
Waterton Lakes National Park of Canada, 5.VII- 
16.VII.1991, H. Goulet (1 male, USNM ENT 
00653585 (CNCI)); montaine prairie, Waterton 
Lakes National Park of Canada, 5.Vin.l991, H. 
Goulet (1 female, 1 male, USNM ENT 00653581, 
00653582 (CNCI)); montaine prairie, Waterton 
Lakes National Park of Canada, 8.VII.1991, H. 
Goulet (1 male, USNM ENT 00653565 (CNCI)); 
nr. mouth of Blakiston Creek, flowery prairie, 
Waterton Lakes National Park of Canada, 
8.VII.1991, H. Goulet (2 females, 3 males, USNM 
ENT 00653572-00653576 (CNCI)); nr. mouth of 
Blakiston Creek, flowery prairie, Waterton 
Lakes National Park of Canada, no date, H. 
Goulet (1 female, USNM ENT 00653580 
(CNCI)). BRITISH COLUMBIA. 57km N Prince- 
ton, hwy. 5, pine /grass, Kentucky- Alleyne 
Park, 10.VII.1986, H. Goulet (1 female, USNM 
Kouchibouguac National Park of Canada, 
7.VII.1977, M. Ivanochko (1 female, USNM 
ENT 00653579 (CNCI)). NOVA SCOTIA, sand- 
bar. South Harbour, 28.VI.1983 (1 male, USNM 
COLORADO. Boulder Co., University of Colo- 
rado Mountain Research Station (Science 
Lodge), 5.VIL6.VII.1961, W. R. M. Mason (1 
female, USNM ENT 00653556 (CNCI)); Clear 
Creek Co., Mt. Evans, Doolittle Ranch, 
8.VII.1961, S. M. Clark (1 female, USNM ENT 
00653557 (CNCI)); Grand Co., 7km E Winter 
Park, Rollins Pass Road, 3.VIII.1999, S. L. 
Heydon & S. M. L. Heydon (1 male, USNM 
ENT 00655515 (UCDC)). IDAHO. Cassia Co., 
#8, Burley, 27.VI.1932, D. E. Fox (1 female, 
Valencia Co., 20mi W Los Lunas, along 


Journal of Hymenoptera Research 

streambed, Carrizo Creek, l.VIII-23. VIII. 1977, 
Malaise trap, S. Peck & J. Peck (1 female, USNM 
ENT 00653584 (CNCI)). OREGON. Lake Co., 
15km N Lakeview, Bull Creek Campground, 
20.VII.1994, S. L. Heydon (1 female, USNM ENT 
00655445 (UCDC)). UTAH. Uintah Co., Vernal, 
1912, C. N. Ainslie (1 female, USNM ENT 
00653538 (USNM)). 

Banacuniculus merickeli (Miller), new 

Nordlanderia merickeli Miller, 1989: 158-159, 
162. Holotype lost. 

Figs IE and 3D. 

Description. — Malar sulcus compound. 
Malar space partially striate, striations 
extending 1/2 to 2/3 distance from ventral 
margin of malar space to base of com- 
pound eye. Malar protuberance smooth, 
elongate, extending beyond length of ven- 
tral margin of malar space. Clypeal protu- 
berance elongate, overhanging anterior 
margin of clypeus. Tubercles of scutellar 
plate absent. Dorsal surface of scutellar 
plate gently convex, smooth; setal bearing 
pits surrounding midpit. Carina along 
posterior margin of scutellum delicate, 
defining transition from dorsal surface of 
scutellum from posterior surface. Dorsal 
surface of scutellum smooth anteriorly, 
longitudinally striate posteriorly. Midpit 
of scutellar plate in center of plate; plate 
small, revealing dorsal surface of scutellum 
when viewed dorsally. Mesopopleural se- 
tal patch absent. Mesopleuron entirely 
smooth. Lateral aspect of pronotum with 
some short setae anteriorly, remainder 
glabrous. Marginal cell of forewing as deep 
as long. Metasoma subequal in size to 
mesosoma in lateral view. 

Diagnosis. — This species is separated 
from all other Banacuniculus by having a 
longitudinally striate dorsal surface of the 
scutellum; in all other species in the genus, 
the dorsal surface of the scutellum is either 
entirely smooth, or variously crenulate- 

Link to Distribution Map. — http://hol. 
Biology. — Unknown. 

Material Examined. — Other material: (13 fe- 
males, 9 males) CANADA: ALBERTA, grass, 
Writing-On-Stone Provincial Park, 24. VIII. 1990, 
McCorquodale (1 female, USNM ENT 00653589 
(CNCI)); montaine prairie, Waterton Lakes 
National Park of Canada, 14.VII.1991, H. Goulet 
(2 males, USNM ENT 00653587, 00653588 
(CNCI)); montaine prairie, Waterton Lakes 
National Park of Canada, 5.VII-16.VII.1991, H. 
Goulet (1 female, USNM ENT 00653583 
(CNCI)); montaine prairie, Waterton Lakes 
National Park of Canada, 5.Vn.l991, H. Goulet 
(1 female, 1 male, USNM ENT 00653586, 
00655612 (CNCI)); nr. mouth of Blakiston Creek, 
flowery prairie, Waterton Lakes National Park 
of Canada, 8.VII.1991, H. Goulet (1 female, 
USNM ENT 00655580 (CNCI)). MEXICO: DF, 
12mi W Texcoco de Mora, 2300m, 28.X.1982, 
screen sweeping, A. Gonzalez (1 female, UCRC 
ENT 196943 (UCRC)). MOR, El Zarco, 1.VI.1954 
(1 unknown, USNM ENT 00653532 (USNM)). 
NICARAGUA: Rivas Dept., San Juan del Sur, 
11°15'N 85°52'W, 21.VII.1994, L. J. Clark (1 
female, USNM ENT 00655504 (UCDC)). UNIT- 
Co., S of Barton Flats, 2090m, 19.VI-26.VI.2007, 
Malaise trap, F. Renter (1 male, USNM ENT 
00655317 (USNM)); Stanislaus Co., romaine 
lettuce, Modesto, 8.XI.1944, C. Weber (1 female, 
Larimer Co., Fort Collins, 25.V.1995, C. F. Baker 
(1 female, USNM ENT 00655344 (USNM)); 
Larimer Co., Fort Collins, VI-1895, C. F. Baker 
(1 female, USNM ENT 00653528 (USNM)). 
NORTH DAKOTA. Hettinger, Mott, no date, 
sweeping, C. N. Ainslie (1 female, USNM ENT 
00653519 (USNM)). NEW MEXICO. Quay Co., 
along rt. 66, within city limits, Tucumcari, 
4.rV.2003, sweeping, M. Buffington (1 male, 
USNM ENT 00655334 (USNM)); Quay Co., 
along rt. 66, within city limits, general vegeta- 
tion, Tucumcari, 4.VI.2003, sweeping, M. Buf- 
fington (1 male, USNM ENT 00655331 (USNM)). 
NEVADA. Lander Co., summit above Austin, 
2430m, 8.VIII.1999, S. L. Heydon & S. M. L. 
Heydon (1 female, USNM ENT 00655440 
(UCDC)). TEXAS. Val Verde Co., Seminole 
Canyon State Historical Park, 1400ft, 
15.IV.1999, J. M. Heraty (1 female, UCRC 

Volume 19, Number 1, 2010 


ENT 196938 (UCRC)). WYOMING. Big Horn 
Co., northern Big Horn Mts., alpine meadow. 
Sheep Mountain, 22.VII.1988, H. Goulet 
(1 male, USNM ENT 00655581 (CNCI)); 
Platte Co., Chugwater Creek, Chugwater, 
16.VIII.1986, J. D. Pinto (1 male, USNM ENT 
00655614 (USNM)); Platte Co., town center, 
Chugwater, 16.Vin.l986, J. D. Pinto (1 male, 
UCRC ENT 196936 (UCRC)); Platte Co., town 
center, Chugwater, 16.VIIL1996, J. D. Pinto (1 
male, UCRC ENT 196935 (UCRC)); Sheridan 
Co., Bighorn Mountains, 24mi W Dayton, 
20.VIIL1983, G. Gordh (1 male, UCRC ENT 
196934 (UCRC)); Teton Co., Granite Canyon, 
1987/060, Teton National Forest, 3.VII.1987, J. 
A. Jackson (1 female, USNM ENT 00655347 

Comments. — Miller (1989) described two 
species of eucoiline wasps that he placed in 
Nordlanderia Quinlan. Though the location 
of the type specimens for these two species 
is unknown (Miller, pers. comm.), it is clear 
from the scanning electron micrographs 
accompanying the descriptions that one 
species, B. merickeli (Miller) possesses the 
diagnostic features of Banacuniculus but not 
all of the defining features of Nordlanderia; 
therefore, this species is transferred to 
Banacuniculus. Communication with the 
original describer of the species suggests 
the holotype may yet surface; I am reticent 
to designate a neotype at the present time 
until it can be fully verified that the 
holot)^e is lost. 

Banacuniculus nigrimanus (Kieffer), new 


Eucoela nigrimanus Kieffer, 1907: 138. Holo- 
type in CASC. 

Ganaspidium nigrimanus (Kieffer): Buffington 
(2004, submitted). 

Figs 2 D-F. 

Description. — Malar sulcus simple. Malar 
space smooth; partially striate, striations 
extending to 1/4 distance from ventral 
margin of malar space to base of com- 
pound eye. Malar protuberance smooth, 
short, not extending beyond length of 

ventral margin of malar space. Clypeal 
protuberance short, not overhanging ante- 
rior margin of clypeus. Tubercles of scu- 
tellar plate absent. Dorsal surface of scu- 
tellar plate flat, smooth; setal bearing pits 
present surrounding midpit. Carina along 
posterior margin of scutellum absent. 
Dorsal surface of scutellum entirely 
smooth. Midpit of scutellar plate in center 
of plate; plate small, revealing dorsal sur- 
face of scutellum when viewed dorsally. 
Mesopopleural setal patch absent. Meso- 
pleuron entirely smooth. Lateral aspect of 
pronotum with some short setae anteriorly, 
remainder glabrous. Marginal cell of 
forewing as deep as long. Metasoma 
subequal in size to mesosoma in lateral 

Diagnosis. — This species is very similar 
to B. utilis but can be easily separated by 
the small relative size of the scutellar plate; 
in B. utilis the plate is enormous, covering 
the entire dorsal surface of the scutellum 
when viewed dorsally. In B. nigrimanus, the 
plate is much smaller, revealing the major- 
ity of the dorsal surface of the scutellum. 

Link to Distribution Map. — http://hol. 

Biology. — Recorded parasitizing Lirio- 
myza huidobrensis (Blanchard) (based on 
label data). 

Material Examined. — Holotype, male: [first 
label] Claremont Cal. Baker, [second label] 
5695, [third label] Eucoela nigrimanus Kieffer 
(in Kieffer's hand), [fourth label] California 
Academy of Science Type No. 10573. [Note: this 
'Baker number' could not be located in the 
Baker notes housed at the USNM]. Other 
material: (60 females, 14 males) CANADA: 
ALBERTA. Medicine Hat, 14.Vn.l956, O. Peck 
(1 female, USNM ENT 00655592 (CNCI)); 
Waterton Park, 18.VI.1956, O. Peck (1 male, 
USNM ENT 00655579 (CNCI)); montaine 
prairie, Waterton Lakes National Park of Can- 
ada, 5.Vni-16.Vni.l991, Malaise trap, H. Goulet 
(1 female, USNM ENT 00655620 (USNM)); 
montaine prairie, Waterton Lakes National Park 
of Canada, 8.VII.1991, H. Goulet (1 male, USNM 
ENT 00655613 (CNCI)); nr. Mt. Galway, mon- 
taine prairie, Waterton Lakes National Park of 


Journal of Hymenoptera Research 

Canada, 5.VIII-16.VIII.1991, Malaise trap, H. 
Goulet (1 female, USNM ENT 00655622 
(CNCI)); nr. Mt. Galway, subalpine prairie, 
Waterton Lakes National Park of Canada, 
9.VII.1991, Malaise trap, H. Goulet (1 female, 
USNM ENT 00655621 (CNCI)). MEXICO: BCS, 
road, vicinity of La Ventana, 8.m.l963, P. H. 
Amaud (1 male, USNM ENT 00655424 (CASC)). 
SON, tomatoes, Heroica Nogales, 23.111.1943 (1 
female, USNM ENT 00653520 (USNM)). UNIT- 
ED STATES: ARIZONA. Graham Co., desert, 
2.4km W on hwy 366 from hwy 191 (666), 
1160m, 27.VI-28.VI.1991, Malaise trap, J. E. 
O'Hara (2 females, USNM ENT 00655596, 
00655597 (CNCI)). CALIFORNIA. Amador Co., 
3km N Silver Lake, Martin Meadow, 
22.VII.1993, L. S. Kimsey & R. B. Kimsey (1 
female, USNM ENT 00655488 (UCDC)); Imper- 
ial Co., 11.3km NW hwy 78, Coachella Canal 
Road, Algodones Dunes, 25.111.2008, sweeping, 
S. L. Heydon (1 female, USNM ENT 00655525 
(UCDC)); Inyo Co., 31km ENE Big Pine, 
25.V.1994, S. L. Heydon (6 females, 2 males, 
USNM ENT 00655498, 00655499, 00655506- 
00655507, 00655508, 00655509, 00655510- 
00655511 (UCDC)); Inyo Co., Grays Meadow 
Campground, 6000ft, 17.VII.1985, A. S. Menke 
(1 female, USNM ENT 00655343 (USNM)); 
Nevada Co., 6km NW Hobart Mills, Sagehen 
Creek Field Station, 29.Vn.2002, sweeping, S. L. 
Heydon (2 females, USNM ENT 00655434- 
00655435 (UCDC)); Nevada Co., Hobart Mills, 
10.VII.1978, R. M. Bohart (1 female, USNM ENT 
00655449 (UCDC)); Riverside Co., vegetation/ 
wash, Wiley Well Campground, 176m, 1.V.2008, 
sweeping, M. Gates (1 female, USNM ENT 
00653530 (USNM)); San Bernardino Co., Imi N 
of crossing of 2N93 Service Road and hwy 38, 
H97-20, San Bernardino Mountains, 2350m, 
24.VI.1997, J. M. Heraty (1 male, UCRC ENT 
197000 (UCRC)); Solano Co., 6km SE Suisun 
City, Suisun Marsh, 14.VI.1993, S. L. Heydon 
& L. Guo (3 females, 1 male, USNM 
ENT 00655453, 00655455-00655456, 00655469 
(UCDC)). COLORADO. Grand Co., 22km 
NNW Kremmling, Chimney Rock, 4.Vin.l999, 
S. L. Heydon & S. M. L. Heydon (3 females, 
USNM ENT 00655436, 00655441, 00655501 
(UCDC)); Grand Co., 22km NNW Kremmling, 
hwy 40 & road 27, 4.Vm.l999, S. L. Heydon & S. 
M. L. Heydon (2 females, USNM ENT 00655439, 
00655539 (UCDC)); Larimer Co., Fort Collins, 
18.Vin.l893, Baker (1 female, USNM ENT 

00653512 (USNM)); Park Co., 6km S Lake 
George, along Fish Creek, 7.VII.1992, S. L. 
Heydon (1 female, 1 male, USNM ENT 
00655470, 00655475 (UCDC)). FLORIDA. Char- 
lotte Co., Placida, 11.IV.1952, O. Peck (1 female, 
USNM ENT 00655594 (CNCI)). KANSAS. Geary 
Co., watershed 2D, Konza Prairie Biological 
Station, 5.V-12.V.2005, Malaise trap, Zolnero- 
wich & Metlevski (1 female, USNM ENT 
00655711 (USNM)); Geary Co., watershed C, 
Konza Prairie Biological Station, 26. VIII- 
2.IX.2005, Malaise trap, Zolnerowich & Metle- 
vski (1 female, USNM ENT 00655712 (USNM)). 
NEW MEXICO. Dona Ana Co., Mesilla, no date 
(1 female, USNM ENT 00653536 (USNM)); 
Quay Co., along rt. 66, within city limits, general 
vegetation, Tucumcari, 4.VI.2003, sweeping, M. 
Buffington (2 females, USNM ENT 00655332- 
00655333 (USNM)); Valencia Co., 20mi W Los 
Lunas, along streambed, Carrizo Creek, 
23.Vm.1977, Malaise trap, S. Peck & J. Peck (3 
females, USNM ENT 00655573-00655574, 
00655583 (CNCI)). NEVADA. White Pine Co., 
45km SSE Eureka, 19.Vn.l995, S. L. Heydon & 
R. M. Bohart (1 female, USNM ENT 00655513 
(UCDC)). OREGON. Lake Co., 15km N Lake- 
view, Bull Creek Campground, 20.VII.1994, 
S. L. Heydon (10 females, 2 males, USNM 
ENT 00655442-00655444, 00655446-00655447, 
00655448, 00655457, 00655458, 00655459, 
00655460-00655461, 00655465 (UCDC)); Lake 
Co., Alkali Lake, 21.VII.1994, S. L. Heydon (1 
female, 1 male, USNM ENT 00655477, 00655492 
(UCDC)). TEXAS. Brewster Co., lowland desert 
springs. Big Bend National Park, 21.VII.1977, L. 
Masner (1 male, USNM ENT 00655600 (CNCI)); 
Hidalgo Co., Jalapeno M hot peppers, Weslaco, 
30.IV.2008, Hernandez (1 unknown, USNM 
ENT 00655029 (USNM)); Presidio Co., Big Bend 
Ranch State Park, 27.X-1.IV.1989, Malaise trap, 
D. Judd (1 female, USNM ENT 00655571 
(CNCI)). UTAH. Emery Co., nr. Goblin Valley 
State Park, Wild Horse Creek, 2.Vni-7.Vin.l997, 
Malaise trap, M. Wasbauer & J. Wasbauer (1 
male, USNM ENT 00655450 (UCDC)); Emery 
Co., nr. Goblin Valley State Park, Wild Horse 
Creek, 2.VIII-7.VIII.1997, M. Wasbauer & J. 
Wasbauer (1 female, USNM ENT 00655524 
(UCDC)); Washington Co., Pinto, no date, C. 
N. Ainslie (1 female, USNM ENT 00653531 
(USNM)); Wayne Co., 6mi W Caineville, along 
Fremont River, 4700ft, 29.VI.1993, J. D. Pinto (1 
female, UCRC ENT 196937 (UCRC)); Wayne 

Volume 19, Number I, 2010 


Co., vegetation, 11 km E Torrey, 7.Vin.l996, L. 
A. Baptiste (1 female, USNM ENT 00655438 
(UCDC)). WYOMING. Big Horn Co., northern 
Big Horn Mts., alpine meadow. Sheep Moun- 
tain, 22.Vn.1988, H. Goulet (4 females, USNM 
ENT 00655575-00655577, 00655578 (CNCI)). 

Banacuniculus strykeri Buffington, new 

Fig. 2C. 

Description. — Malar sulcus compound. 
Malar space partially striate, striations ex- 
tending 1/2 to 2/3 distance from ventral 
margin of malar space to base of compound 
eye. Malar protuberance striate, short, not 
extending beyond length of ventral margin 
of malar space. Clypeal protuberance short, 
not overhanging anterior margin of clypeus. 
Midpit of scutellar plate in center of plate; 
plate large, obscuring dorsal surface of 
scutellum when viewed dorsally. Tubercles 
of scutellar plate absent. Dorsal surface of 
scutellar plate concave, radially striate; setal 
bearing pits surrounding midpit. Carina 
along posterior margin of scutellum pres- 
ent, distinctly cleft, defining transition from 
dorsal surface of scutellum from posterior 
surface. Dorsal surface of scutellum entirely 
rugulose/wrinkled. Mesopopleura setal 
patch absent. Mesopleuron striate within 
confines of mesopleural triangle, remainder 
of sclerite smooth. Lateral aspect of prono- 
tum with some short setae anteriorly, 
remainder glabrous. Marginal cell of fore- 
wing as deep as long. Metasoma sub-equal 
size to mesosoma in lateral 

Diagnosis. — Differs from other species by 
the large scutellar plate, the presence of the 
radial striations on the surface of the 
scutellar plate, and by the reduced malar 
and clypeal protuberances. 

Biology. — Reared from an isolated pu- 
parium of Liriomyza lathyri Seghal in Mar- 
quette, MI (holotype specimen). 

Etymology. — Named in honor of my son, 
Stryker Buffington. 

Link to Distribution Map. — http://hol. 

Type Material. — Holotype, female: United 
States, MI, Marquette Co. 46=50. 627'N 
87=51.300'W, 16 August 2006, R. Priest. Lot # 
R1P1755.2.1, USNM ENT 00655717. Deposited in 
USNM. Paratypes: UNITED STATES: 1 female, 
2 males: COLORADO. Fremont Co., Phantom 
Canyon, 29-31/5/1987, G. Hevel, 1 female 
Washtenaw Co., Ann Arbor, 7/12-21/1982, R. 
Wharton, 2 males USNM ENT 00655520- 
00655521 (TAMU). 

Comments. — The scutellar morphology 
and the distribution patterns of this species 
are striking features. More Banacuniculus 
species are likely to occur in the Mid- 
western United States. 

Banacuniculus utilis (Beardsley), new 

Ganaspidium utilis Beardsley, 1988: 44-46, 
holotype in BPBM. 

Synonymized with Ganaspidium nigrimanus 
(Kieffer) by Buffington (2004); revised status: 
Buffington (2010). 

Figs 3 A-C. 

Redescription. — Malar sulcus simple. Ma- 
lar space smooth; partially striate, stria- 
tions extending 1/2 to 2/3 distance from 
ventral margin of malar space to base 
of compound eye. Malar protuberance 
smooth, short, not extending beyond 
length of ventral margin of malar space. 
Clypeal protuberance short, not overhang- 
ing anterior margin of clypeus. Midpit of 
scutellar plate in center of plate; plate large, 
obscuring dorsal surface of scutellum 
when viewed dorsally. Tubercles of scu- 
tellar plate absent. Dorsal surface of scu- 
tellar plate flat, smooth, setal bearing pits 
present surrounding midpit. Carina along 
posterior margin of scutellum absent. 
Dorsal surface of scutellum entirely 
smooth. Mesopopleural setal patch absent. 

Journal of H^aiexopteila Research 

Fig. 2. A, Ba7iacimicuhis hunteri (Crawford), posterodorsal view of mesosoma; B, B. hunieri, lateral habitus of 
holotype with inset of specimen labels; C, B. stnjkeri, n. sp., postero-dorsal view of scutellum and scutellar plate; 
D, 6. nigrimanus (Kieffer), lateral habitus of holotv'pe with inset of specimen labels; E, 6. nigrimanus, lateral 
habitus of non-type specimen; F, 6. nigrimanus, postero-dorsal \iew of scutellum, scutellar plate, 
and propodeum. 

Mesopleuron entirely smooth. Lateral as- 
pect of pronotum with some short 
setae anteriorly, remainder glabrous. Mar- 
ginal cell of forewing as deep as long. 
Metasoma sub-equal size to mesosoma in 
lateral view. 

Diagnosis. — Differs from Bajjacuniculus 
nigrimanus by having a large scutellar 
plate, and from B. beardsleyi by having a 
smooth dorsal surface of the scutellar plate; 
from the other species of Banacuniculus by 
characters in the beginning of the key. 

Volume 19, Number 1, 2010 


Distribution. — Canada: Alberta; Guam; 
Guatemala: Esquintia; Mexico: Baja Cali- 
fornia, Chiapas, Morelos, Oaxaca, Sonora, 
and Zacatecas; Nicaragua: Rivas; United 
States: Arizona, California, Kansas, New 
Mexico, Oregon, Texas and Utah. 

Link to Distribution Map. — http://hol. 

Biology. — Recorded parasitizing Lirio- 
myza trifolii (Hara and Matayoshi 1990; 
Johnson 1987; Beardsley 1986; this study), 
L. sativae (Johnson 1987) and L. huidobrensis 
(Blanchard) (Johnson 1987). This species 
has also been evaluated for its usefulness 
in biological control of pestiferous leaf- 
mining flies Qohnson 1987; Lynch and 
Johnson 1987; Mason and Johnson 1988; 
Rathman et al. 1991; Rathman et al. 1995). 
Petcharat and Johnson (1988) studied the 
larval stages. 

Material Examined. — Holotype, female: [first 
label] USA. HI, Oahu. Nanakuli, X.3.1977, 
[second label] P.D. Mothershead, reared ex 
Liriomyza pupae from cucumber leaves. Depos- 
ited in BPBM. Other material: (176 females, 41 
males) Other material: (176 females, 40 males, 1 
unknown) CANADA: ALBERTA. Waterton 
Park, 18.VI.1956, O. Peck (1 female, USNM 
ENT 00655599 (CNCI)). GUAM: watermelon 
field, n-1989, pan trap, L. Yudin (3 females, 
UCRC ENT 196967-196969 (UCRC)). watermel- 
on field, IX-1989, pan trap, L. Yudin (2 females, 
UCRC ENT 196970-196971 (UCRC)). GUATE- 
MALA: Escuintla Dept., Escuintla, 20.Vin.l975, 
N. L. H. Krauss (1 male, USNM ENT 00653533 
(USNM)). MEXICO: BC, 57km S Bahia de los 
Angeles, 22.Vn.1994, S. L. Heydon (1 male, 
USNM ENT 00655487 (UCDC)). BCS, 10km W 
San Ignacio, 24.111.1980, E. Fisher & J. Pinto (1 
female, UCRC ENT 196944 (UCRC)). BCS, 
Barracas, 18.V.1985, pan trap, P. DeBach (1 
female, UCRC ENT 196940 (UCRC)). BCS, 
Barracas, 20.V.1985, pan trap, P. DeBach (1 
female, UCRC ENT 196939 (UCRC)). BCS, La 
Paz, 20.X.1983, J. D. Pinto (1 female, 1 male, 
UCRC ENT 196941-196942 (UCRC)). Chiapas, 
14.Vn.l983, A. Gonzalez (1 female, UCRC ENT 
196951 (UCRC)). MEX, Naucalpan de Juarez, 
18.V.1984, G. Gordh (1 female, UCRC ENT 
196950 (UCRC)). MOR, Cuemavaca, m-1965 - 
V-1965, N. L. H. Krauss (1 female, 1 male. 

USNM ENT 00653515, 00653522 (USNM)). NL, 
Cienega de Hores, 10.Vn.l983, A. Gonzalez (1 
female, 1 male, UCRC ENT 196952-196953 
(UCRC)). OAX, Puerto Escondido, 29.V.1963, 
E. R. Oatman (1 female, UCRC ENT 196954 
(UCRC)). OAX, Yagul, 13.Vn.l984, G Gordh (1 
female, 1 male, UCRC ENT 196948-196949 
(UCRC)). SON, 1.XI.1947 (1 unknown, USNM 
ENT 00653534 (USNM)). SON, San Jose de 
Guaymas, 4.X.1900, L. O. Howard (1 female, 
USNM ENT 00653535 (USNM)). ZAC, Monte 
Escobedo, 12.Vn.l983, G. Gordh (3 females, 
UCRC ENT 196945-196947 (UCRC)). NICARA- 
GUA: Rivas Dept., San Juan del Sur, 11=15'N 
85=52'W, 15.IV.1998, L. J. Clark (1 female, 
USNM ENT 00655502 (UCDC)). Rivas Dept., 
San Juan del Sur, 11=15'N 85=52'W, 2.n.l998, 
Malaise trap, J. Clark (1 female, 1 male, USNM 
ENT 00655534, 00655537 (UCDC)). Rivas Dept., 
San Juan del Sur, 11=15'N 85=52'W, 25.VI.1998, 
L. J. Clark (1 female, USNM ENT 00655503 
chise Co., Huachuca Mts., Ash Canyon Road, 
15.m-30.IV.1994, Malaise trap, McFarland (1 
female, USNM ENT 00655595 (CNCI)); Graham 
Co., desert, 2.4km W on hwy 366 from hwy 191 
(666), 1160m, 27.VI-28.VI.1991, Malaise trap, J. 
E. O'Hara (1 female, USNM ENT 00655585 
(CNCI)); Pima Co., Tucson, 11.IV.1896, Baker (2 
females, USNM ENT 00653526, 00653539 
(USNM)); Santa Cruz Co., Patagonia, 4.VI.1995, 
E. Wilk & B. Brown (1 female, USNM ENT 
00655586 (CNCI)); Santa Cruz Co., juniper/ 
oak/grassland, Sonoita, 29.IX-13.X.2006, Ma- 
laise trap, E. E. Grissell (1 female, USNM ENT 
00655316 (USNM)). CALIFORNIA. Imperial Co., 
14.4km WNW Glamis, 6.4km NW hwy 78, 
2008A1126, Algodones Dunes, 22.IX-15.XI.2008, 
Malaise trap, E. Dreyfus (1 female, USNM ENT 
00655519 (UCDC)); Los Angeles Co., Eaton 
Canyon, oak/ scrub, Pasadena, 1000ft, 26. VI. 
2002, A. Ow^ens & J. George (1 female, UCRC 
ENT 56868 (UCRC)); Los Angeles Co., Forrestal 
Nature Preserve, coastal sage scrub, Rancho 
Palos Verdes, 20.rV-24.V.2003, Malaise trap, 
J. George (6 females, 4 males, USNM 
ENT 00655318, 00655319, 00655320-00655325, 
00655329, 00655330 (USNM)); Los Angeles Co., 
Forrestal Nature Preserve, coastal sage scrub, 
Rancho Palos Verdes, 5.IV-20.rV.2003, Malaise 
trap, J. George (1 female, USNM ENT 00655339 
(USNM)); Orange Co., T6S RlOW S17, Hunting- 
ton Beach, 3.1^.1969, lab reared, R. D. Gbeden & 


Journal of Hymenoptera Research 

D. W. Ricker (1 female, 1 male, UCRC ENT 
196959-196960 (UCRC)); Riverside Co., Moreno 
Valley, University of California Experiment 
Station, 5.IX.1978, J. LaSalle (1 female, UCRC 
ENT 196963 (UCRC)); Riverside Co., N of Oasis, 
G95/155, Thousand Palms Canyon, 17.IX.1995, 
M. Gates (1 male, UCRC ENT 196962 (UCRC)); 
Riverside Co., PEET 001-08-14-01 /MT3, Santa 
Rosa Plateau Ecological Reserve, 500m, SO.VII- 
14.Vin.2001, Malaise trap (4 females, USNM 
ENT 00655335-00655336, 00655338, 00655342 
(USNM)); Riverside Co., Santa Rosa Mts., Santa 
Rosa Spring Campground, 6400ft, 10.IX.1964, E. 
I. Schlinger (1 female, UCRC ENT 196958 
(UCRC)); San Bernardino Co., N of Silverw^ood 
Lake, Summit Valley, 27.Vn.1996, M. Gates (1 
female, UCRC ENT 196961 (UCRC)); San Diego 
Co., Rancho Santa Fe, 8.Vin.l979, C. Melton (2 
females, UCRC ENT 196965-196966 (UCRC)); 
San Diego Co., Torrey Pines Park, 60m, 
19.VI.1996, D. C. Hawks (1 female, UCRC ENT 
196999 (UCRC)); San Diego Co., melon, Escon- 
dido, 4.VI.1964, E. R. Oatman (1 female, UCRC 
ENT 196957 (UCRC)); San Diego Co., pole 
beans, Escondido, 5.IX.1963, E. R. Oatman (1 
female, UCRC ENT 196955 (UCRC)); San Diego 
Co., pole beans, Escondido, 9.VII.1964, E. R. 
Oatman (1 female, UCRC ENT 196956 (UCRC)); 
Santa Barbara Co., 45km NW Santa Barbara, 
Sedgwick Reserve, 308m, 24.VI-8.Vn.1997, Ma- 
laise trap, E. S. Schlinger (2 females, USNM ENT 
00655541, 00655549 (UCDC)); Santa Barbara Co., 
Santa Barbara, 11.VT1997, E. Schlinger (1 
female, USNM ENT 00655538 (UCDC)). HA- 
WAII. Honolulu Co., tomato field, O'ahu (Oahu) 
Isl., 28.x. 1976, R. Buckhart (8 females, 3 males, 
USNM ENT 00655601-00655607, 00655608, 
00655609, 00655610-00655611 (USNM)). KAN- 
SAS. Geary Co., watershed 20B, Konza Prairie 
Biological Station, 12.IX-21.IX.2005, Malaise 
trap, Zolnerowich & Metlevski (1 female, 
USNM ENT 00655701 (USNM)); Geary Co., 
watershed 20B, Konza Prairie Biological Station, 
16.Vin-26.Vin.2005, Malaise trap, Zolnerowich 
& Metlevski (1 female, USNM ENT 00655702 
(USNM)); Geary Co., watershed 20B, Konza 
Prairie Biological Station, 2.IX-12.IX.2005, Ma- 
laise trap, Zolnerowich & Metlevski (1 female, 
USNM ENT 00655703 (USNM)); Geary Co., 
watershed 20B, Konza Prairie Biological Station, 
20.VII-30.VII.2005, Malaise trap, Zolnerowich & 
Metlevski (1 female, USNM ENT 00655705 
(USNM)); Geary Co., watershed 20B, Konza 

Prairie Biological Station, 27.V-6.VI.2005, Ma- 
laise trap, Zolnerowich & Metlevski (1 female, 
USNM ENT 00655706 (USNM)); Geary Co., 
watershed 20B, Konza Prairie Biological Station, 
30.IX-11.X.2005, Malaise trap, Zolnerowich & 
Metlevski (1 female, USNM ENT 00655700 
(USNM)); Geary Co., watershed 20B, Konza 
Prairie Biological Station, 30.Vn-9.Vni.2005, 
Malaise trap, Zolnerowich & Metlevski (2 
females, USNM ENT 00655707, 00655708 
(USNM)); Geary Co., watershed 20C, Konza 
Prairie Biological Station, 18.V-22.V.2006, Ma- 
laise trap, Zolnerowich & Metlevski (1 female, 
Valencia Co., 20mi W Los Lunas, along 
streambed, Carrizo Creek, 23.VnL1977, Malaise 
trap, S. Peck & J. Peck (6 females, 2 males, 
USNM ENT 00655582, 00655584, 00655587, 
00655588, 00655589, 00655590, 00655593, 
00655598 (CNCI)). OREGON. Lake Co., nr. 
pond, 22km N Lakeview, 20.Vn.l994, S. L. 
Heydon (1 female, USNM ENT 00655433 
(UCDC)). TEXAS, no date, Hernandez (8 fe- 
males, 2 males, USNM ENT 00655227, 00655228, 
00655229, 00655230-00655236 (USNM)); Ca- 
meron Co., Brownsville, no date, R. A. Vickery 
(1 female, USNM ENT 00653521 (USNM)); 
Cameron Co., TAM Veracruz hot pepper, 
Brownsville, 17.X.2007, Hernandez (1 female, 
USNM ENT 00655184 (USNM)); Hidalgo Co., 
Cuban Hots hot pepper, Edinburg, 12.XI.2007, 
Hernandez (1 female, USNM ENT 00655281 
(USNM)); Hidalgo Co., Cuban Hots hot pepper, 
Edinburg, 27.XI.2007, Hernandez (1 female, 
USNM ENT 00655271 (USNM)); Hidalgo Co., 
Cuban Hots hot pepper, Edinburg, 30.X.2007, 
Hernandez (1 male, USNM ENT 00655262 
(USNM)); Hidalgo Co., Jalapeno M hot peppers, 
Weslaco, 1.IV.2008, Hernandez (Ifemale, USNM 
ENT 00655116 (USNM)); Hidalgo Co., Jalapeno 
M hot peppers, Weslaco, 10.L2008, Hernandez (3 
females, USNM ENT 00655259, 00655265, 
00655268 (USNM)); Hidalgo Co., Jalapeno M 
hot peppers, Weslaco, 12.XL2007, Hernandez (9 
females, 2 males, USNM ENT 00655243- 
00655247, 00655276, 00655280, 00655282, 
00655283, 00655284, 00655287 (USNM)); Hidalgo 
Co., Jalapeno M hot peppers, Weslaco, 
13.Xn.2007, Hernandez (3 females, 1 male, 
USNM ENT 00655239, 00655266, 00655270, 
00655272 (USNM)); Hidalgo Co., Jalapeno M 
hot peppers, Weslaco, 14.V.2008, Hernandez (2 
females, USNM ENT 00655043, 00655057 

Volume 19, Nl-mber 1, 2010 


Fig. 3. A. Banacuniculus utilis (Beardsley), lateral habitus of holot}-pe with inset of specimen labels; B, B. utilis, 
lateral habitus of non-t\^e specimen; C, B. utilis forewing, non-t\'pe specimen; D, habitat photo of the Konza 
Prairie Presence, Kansas; E, habitat photo of Algodone Dunes, California; F, habitat photo of Cold Canyon 
Preser\^e, California. 

(USNM)); Hidalgo Co., Jalapeno M hot peppers, 
Weslaco, 16.X.2007, Hernandez (2 females, 1 
male, USNM ENT 00655251, 00655286, 
00655296 (USNM)); Hidalgo Co., Jalapeno M 
hot peppers, Weslaco, 19.IX.2007, Hernandez (14 
females, 4 males, USNM ENT 00655196, 00655197, 
00655198, 00655199, 00655202-00655207, 00655208, 
00655209, 00655215, 00655255-00655257, 00655258, 
00655263 (USNM)); Hidalgo Co., Jalapeno M hot 

peppers, Weslaco, 19.X.2007, Hernandez (3 fe- 
males, 1 male, USNM ENT 00655185, 00655186, 
00655250, 00655285 (USNM)); Hidalgo Co., Jala- 
pefio M hot peppers, Weslaco, 2.XI.2007, Her- 
nandez (3 females, USNM ENT 00655248, 
00655290, 00655291 (USNM)); Hidalgo Co., Jala- 
pefio M hot peppers, Weslaco, 20.XII.2007, 
Hernandez (5 females, USNM ENT 00655238, 
00655264, 00655267, 00655269, 00655273 


Journal of Hymenoptera Research 

(USNM)); Hidalgo Co., Jalapeno M hot peppers, 
Weslaco, 25.X.2007, Hernandez (1 female, USNM 
ENT 00655249 (USNM)); Hidalgo Co., Jalapeno 
M hot peppers, Weslaco, 26.VI.2008, Hernandez 
(1 female, 1 male, USNM ENT 00655058, 
00655072 (USNM)); Hidalgo Co., Jalapeno M 
hot peppers, Weslaco, 28.XI.2007, Hernandez (1 
female, USNM ENT 00655279 (USNM)); Hidalgo 
Co., Jalapeno M hot peppers, Weslaco, 29.V.2008, 
Hernandez (1 female, USNM ENT 00655070 
(USNM)); Hidalgo Co., Jalapeno M hot peppers, 
Weslaco, 30.X.2007, Hernandez (4 females, 1 
male, USNM ENT 00655288, 00655289, 
00655292, 00655293, 00655294 (USNM)); Hidalgo 
Co., Jalapeno M hot peppers, Weslaco, 
30.XI.2007, Hernandez (2 females, 2 males, 
USNM ENT 00655240-00655242, 00655275 
(USNM)); Hidalgo Co., Jalapeno M hot peppers, 
Weslaco, 6.IX.2007, Hernandez (4 females, 
USNM ENT 00655210-00655212, 00655254 
(USNM)); Hidalgo Co., Jalapeno M hot peppers, 
Weslaco, 8.X.2007, Hernandez (9 females, 2 males, 
USNM ENT 00655187, 00655188, 00655189, 
00655190, 00655191, 00655192, 00655194, 
00655195, 00655252-00655253, 00655295 (USNM)); 
Hidalgo Co., Jalapeno M hot peppers, Weslaco, 
9.VI.2008, Hernandez (2 females, USNM ENT 
00655059, 00655071 (USNM)); Hidalgo Co., Wes- 
laco, 17.Xn.l968, F. F. Smith (1 female, USNM 
ENT 00653518 (USNM)); Hidalgo Co., serrano hot 
pepper, San Juan, 12.XI.2007, Hernandez (1 male, 
USNM ENT 00655260 (USNM)); Hidalgo Co., 
serrano hot pepper, San Juan, 18.X.2007, Her- 
nandez (1 female, 1 male, USNM ENT 00655140- 
00655141 (USNM)); Hidalgo Co., serrano hot 
pepper, San Juan, 3.X.2007, Hernandez (2 fe- 
males, USNM ENT 00655200-00655201 (USNM)); 
Jim Wells Co., 8mi W Ben Bolt, area 4, N end of 
La Copita Ranch, 29.IX.1990, J. B. Woolley (2 
females, USNM ENT 00655340, 00665341 
(USNM)); Jim WeUs Co., Ben Bolt, La Copita 
Ranch, 26.IX-30.IX. 1990, Malaise trap, Wharton 
& Woolley (1 female, USNM ENT 00655591 
(CNCI)). UTAH. Emery Co., nr. Goblin Valley 
State Park, Wild Horse Creek, 2.VIII-7.VIII.1997, 
Malaise trap, M. Wasbauer & J. Wasbauer (4 
females, USNM ENT 00655533, 00655536, 
00655540, 00655542 (UCDC)); Emery Co., nr. 
Goblin Valley State Park, Wild Horse Creek, 
4.VII-7.VIL1997, Malaise trap, M. Wasbauer & J. 
Wasbauer (1 male, USNM ENT 00655535 
(UCDC)); Emery Co., nr. Goblin Valley State 
Park, Wild Horse Creek, 4.VII-7.VIL1997, M. 

Wasbauer & J. Wasbauer (2 females, 1 male, 
USNM ENT 00655531-00655532, 00655543 
(UCDC)); Wayne Co., vegetation, 11 km E 
Torrey, 7.VIL1996, sweeping, L. A. Baptiste (1 
female, USNM ENT 00655544 (UCDC)); train, 
3.VI.1924, Timberlake (1 female, UCRC ENT 
196964 (UCRC)). 

Comments. — After reconsideration of the 
holotypes of both Banacuniculus nigrimanus 
(Kieffer) and B. utilis Beardsley, it is clear 
that these are two distinct and readily 
diagnosed species. The morphology of the 
scutellum in B. utilis is unique within the 


The species diversity of Banacuniculus is 
likely to be much higher than is presently 
recognized. Key locations in North Amer- 
ica needing further fieldwork to collect 
and /or rear additional species include the 
upper Midwest and the Great Basin. Host 
habitat for Banacuniculus species appears to 
be rather variable; this is epitomized by the 
distribution of B. dis, whose hosts range 
from extremely dry conditions of the High 
Desert of California to moister, interior 
scrub oak habitats of Central California 
(Fig. 3 E-F). The distribution B. utilis 
demonstrates the ability of this species to 
be a New World tramp, ranging from the 
Konza Prairie in Kansas (Fig. 3D) and 
Algodone Dunes of California (Fig. 3E), to 
cultivated crops in southern Texas and 
Hawai'i. One of the preferred hosts for this 
species is Liriomyza trifolii, itself a tramp 
species of considerable agricultural impor- 
tance in commodities such as tomato, 
cucumber and melon (Johnson 1987). 

The phylogenetics of Banacuniculus spe- 
cies are not quantitatively analyzed here, 
but there are some morphological features 
that allow for a cautious suggestion of 
relatedness among species. The entirely 
smooth dorsal and lateral surface of the 
scutellum (Fig. 2F) in B. heardsleyi, B. 
nigrimanus, and B. utilis is not only rare 
within Eucoilinae, but unites these species. 
Based on the relative size of the scutellar 

Volume 19, Number 1, 2010 


plate to the scutellum, the following 
relationship is suggested: (B. nigrimanus 
(B. beardsleyi + B. utilis)), and for conve- 
nience, this clade is referred to as the utilis 
species group. Banacuniculus merickeli is 
unique within the genus for having a 
striate dorsal surface of the scutellum, but 
the species does possess a distinct posterior 
carina of the scutellum. Banacuniculus dis 
and B. brautigani both lack the posterior 
scutellar carina, a character state also 
shared with the utilis group; but unlike 
the utilis group, the scutellum is not 
smooth. Hence, B. dis and B. brautigani 
form their own clade, referred to here as 
the dis species group. Banacuniculus hunteri 
and B. strykeri are morphologically very 
similar, differing only in the morphology 
of the scutellar plate and, possibly, their 
host geographic distribution (the latter 
may be a collection artifact); these species 
are referred to here as the hunteri species 
group. Given these observations, the fol- 
lowing relationships are proposed: (hunteri 
group (B. merickeli {dis group + utilis 
group))). In this scenario, a complete 
posterior carina of the scutellum and 
rugose /crenulate dorsal surface of the 
scutellum is pleisiomorphic; B. merickeli 
retains the carina, but possesses the striate 
state for the dorsal surface of the scutellum; 
the clade dis group + utilis group all lack 
the posterior carina of the scutellum, and 
the utilis group possesses the entirely 
smooth state for the dorsal surface of the 
scutellum. This scenario of relationships is 
merely meant to summarize what appears 
to be a parsimonious summary of character 
evolution within Banacuniculus; a more 
thorough survey and analysis of characters 
is required to corroborate and confirm 
these relationships. 


Thanks are given to Smithsonian Institution intern 
Jaime Choi and Systematic Entomology Laboratory 
Museum Specialist David Adamski for entering label 
data and labeling specimens; Joe Cora (Ohio State 
University) assisted with data management and 

digital map generation. Ricardo Hernandez (Texas 
A&M University) sent specimens of Banacuniculus 
utilis to me for identification and allowed me to keep 
the specimens in the USNM; Robert Kula (Systematic 
Entomology Lab) and Gregory Zolnerowich (Kansas 
State Univeristy) assisted with obtaining specimens 
from the Konza Prairie Reserve; Jennifer Read 
(Canadian National Collection of Insects), Robert 
Zuparko (California Academy of Science), Doug 
Yanega (Entomology Research Museum, UC River- 
side), Ed Riley (Texas A&M University) and Steve 
Heydon (Bohart Museum, UC Davis) assisted with the 
loans of specimens critical to the success of this 
research. Rich Pyle (Bishop Museum, Honolulu, HI) 
provided the ZooBank registration numbers. Smith- 
sonian Institution interns Cristy Falcone and Nick 
Olson provided the environmental SEM image in 
Fig. 2. I also thank Robert Wharton, Jack Beardsley 
and Richard Brautigan for advice and positive 
influences over the years. Finally, I extend thanks to 
Michael Pogue and Thomas Henry (Systematic En- 
tomology Laboratory) and Scott Solomon (Depart- 
ment of Entomology, Smithsonian Institution) for 
constructive and useful comments to earlier drafts of 
this paper. Habitat images were provided by Greg 
Zolnerowich (Konza Prairie, KS), Lynn Kimsey 
(Algodone Dunes, CA), and Steve Heydon (Cold 
Canyon, CA). Initial research for this project was 
begun under NSF PEET Grant # DEB9712543 
awarded to Robert Wharton and James Woolley 
(Texas A&M University). 


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parasitoids attacking Liriomyza spp. (Diptera: 
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diella semirufa (Kieffer), new combination, with 
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. 2009. Description, circumscription and phylo- 

genetics of the new tribe Zaeucoilini (Hymenop- 
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(Diptera: Agromyzidae) infesting commercial 
watermelon plantings in Hawaii. Journal of 
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Johnson, N. 2008. vSysLab, 
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Cynipiden aus Zentral- und Nord-Amerika. En- 
tomologische Zeitschift 21: 70-162. 

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sampling of Liriomyza spp. (Diptera: Agromyzi- 
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Rathman, R. J., M. W. Johnson, and B. E. Tabashnik. 
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Vol. 19(1), 2010, pp. 113-120 

Phylogenetic Relationships of Phiroides porteri, a New Genus and 
Species of Plumariidae from Argentina (Hymenoptera: Chrysidoidea) 

Patricla Diez, Arturo Roig-Alsixa, axd Patricio Fidalgo 

(PD, PF) CRIL.AR, Entre Rios y Mendoza s/n, 5301 .-Ynillaco, La Rioja, Argentina; 


(AR-A) Museo .Aigentino de Ciencias Xatuxales "Bernardino Rivadavia", Av. Angel Gallardo 470, 

1405 Buenos Aires, Argentina; arturo@macn. gov. ar 

Abstract — Phiroides porteri, a new genus and species of plumariid -^vasp from the provinces of 
Catamarca, La Rioja and San Juan in western .Argentina, is described. The new genus Phiroides 
together wdth Maplnroides, Maplurius and Plimiaroides, belong in a strongly supported South 
American clade, ^vhich is the sister-group to the African Mynnecopteriiiella. 

Key ZLvrds. — Plumariidae, taxonomy, phylogeny, Argentina 

Fhe Plumariidae are a group of ^vasps 
^vith apterous females of subterranean 
habits and ^vinged males strongly attracted 
to lights at night. This family is of 
particular interest, since it represents one 
of the basalmost lineages ^vithin the Hy- 
menoptera Aculeata. These wasps inhabit 
desertic and semidesertic areas in southern 
Africa and South America. Although they 
are conspicuous faunal elements in some 
areas, their biology is still unkno^vn. 

The family is represented by t^vo genera 
in Africa {Myrmecopterina Bischoff and 
Mynnecopterinella Day) and by four genera 
in South America (Plumariiis Philippi, 
Pliimaroides Brothers, Maplurius Roig- 
Alsina and Mapluroides Diez, Fidalgo and 
Roig-Alsina) (Brues 1924; Bradley 1972; 
Brothers 1974; Dav 1977; Roig-Alsina 
1994; Diez et al. 2007). 

Among South American plumariids, the 
genus Plumarius is the most speciose and 
has the broadest distribution. Species of 
Plumarius range from Ecuador to southern 
Argentina (Evans 1966; Xag}- 1973; Broth- 
ers 1974) with one species recently^ de- 
scribed from northeastern Brazil (Pen- 
teado-Dias and Scatolini 2003). The other 
three genera are restricted to western 

Argentina, from the province of Salta in 
the north to northern Patagonia in the 
south (Brothers 1974; Roig-Alsina 1994; 
Diez et al. 2007; Diez 2008). Plumaroides 
has three described, as weU as several 
undescribed species (Diez 2008), while the 
other t^vo genera are monot^-pic. Females 
of only t^vo genera have been discovered to 
date, those of Plumarius (Evans 1966), and 
Plumaroides (Diez 2008). 

The purpose of the present contribution 
is to describe a ne^v genus and species 
from the provinces of Catamarca, La Rioja, 
and San Juan in Argentina. Both the 
generic and specific descriptions are based 
on the male sex. The relationships of the 
ne^v genus are studied, taking into account 
previous contributions and recently de- 
scribed taxa. 


The specimens studied were collected at 
night ^vith a camping lantern provided 
^vith a fluorescent tube, 360 degrees bright 
Hght. The lantern ^vas placed on a ^vhite 
cloth extended on the ground. A few 
specimens were obtained ^vith a trap 
designed to collect myrmecophilous in- 


Journal of H^:nlexopter.a Research 

Specimens are deposited at: Museo 
Argentine de Ciencias Xaturales "Bernar- 
dino Rivadavia," Buenos Aires, Argentina 
fMACX); Institute Fundacion Miguel Lillo, 
Tucuman, Argentina (IFNIL); Museo de La 
Plata, La Plata, Argentina (MLP); Florida 
State Collection of Arthropods, USA 
(FSCA); Universit}^ of California, Riverside, 

Terminology follows Brothers (1975). 

Pliiroides new genus 

T}^e species: Pliiroides porteri sp. nov. 

Description. — Preoccipital carina absent. 
Antenna with 11 flagellomeres; scape as 
long as \vide, with short radicle, ventrall\' 
with tuberculiform s^velling (Fig. 3); flagel- 
lomeres with decumbent, short setae, the 
longest 0.25 X as long as diameter of 
flagellum. Mandible \vith three teeth, pre- 
apical ones small, blunt, of similar size. 
Palpal formula: 5:1. Ch'peus with episto- 
mal suture distinct; apical margin with 
weak emargination medially, cur\'ed at 
sides; apical margin bent backv\^ards. Pro- 
sternum visible in ventral vie^v, subtrian- 
gular (Fig. 7). Forewing with first nebulous 
vein arising from marginal cell one third 
below middle of apical margin (Fig. 8). 
Hind wing with vannal (anal) lobe 3.3 X as 
long as submedian cell (Fig. 9). Claws 
simple, arolium present only on foretarsus. 
First metasomal tergum with distinct ante- 
rior vertical surface, dorsal surface as long 
and wide as second tergum. First metaso- 
mal sternum with a longitudinal median 
keel on anterior two thirds, longer than 
second sternum. Seventh tergum subtrian- 
gular, apically rounded; posterior margin 
forming flat, sclerotized, polished flange, 
one third as long as tergum (Fig. 10). 
Se\'enth sternum broad, with apex weakly 
bilobed (Fig. 11). 

Pluroides porteri new species 

(Figs 1-13) 

Description. — Holotype male. Colour, pale 

brown, head darker. Total length 3.5 mm 

(paratypes, 2.7 to 4.5 mm). Habitus as in 

figure 1. Head: hypognathous; in frontal 
view 1.3X as wide as high (Fig. 2), vertex 
rounded. Eye hemispherical, protruding, 
glabrous, without pre- or postorbital car- 
inae. Ocell ocular distance 3.8 X diameter of 
lateral ocellus; postocellar distance 1.7x 
ocellocular distance. Antennal socket with 
lower rim elevated. Antennocular distance 
0.3 x diameter of antennal socket; inter- 
antennal distance 6.7 x antennocular dis- 
tance. Gena without furrows or carinae. 
Area between and below sockets weakly 
convex. Antenna tapering to apex. Pedicel 
with narro^v base, as long as \vide. Propor- 
tions of flagellomeres (length:width): 
11:7,5; 13:7; 15:7; 14:6; 14:6; 12:5; 13:5; 13:5; 
12:5; 11:5; 15:5. Sensor}^ plates sub-oval, 
scarcely visible, present on flagellomeres 
1-6; plates more numerous on basal fla- 
gellomeres. Ch~peus ^vith several setae of 
variable size. Labrum with concave apical 
margin, with two setae at each side. 
Mandible with broad base and setae of 
variable size. Labium subrectangular, 
^vider basally than apically, with rounded 
apex; palp unsegmented (Fig. 4). Maxillar}^ 
palp \vith five segments, proportions of 
segments (length:width): 67:22; 40:21; 
33:20; 30:20; 31:20 (Fig. 5). 

Mesosoma: 1.5 x longer than maximum 
width. Proportions of lengths of mesoscu- 
tum, mesoscutellum, metanotum, meta- 
postnotum and propodeum in dorsal view, 
along median axis: 25: 21: 12: 2: 10. 
Pronotum not visible dorsally, except for 
small part of pronotal lobe in front of 
tegula; in frontal view (head removed) 
medially forming narro'sv transverse band 
\vhich broadens laterally to four times 
median height. Pronotal lobe flattened, 
truncate; posteroventral angle of pronotum 
narrowly rounded (Fig. 6). Propleuron ex- 
tended anteriorly bevond pronotum. Pro- 
sternum visible ventrally as reduced trian- 
gular area; most of prostemal surface 
vertical, hidden by coxae (Fig. 7). Tegula 
semicircular. Mesoscutum \vith parapsidal 
line and notaulus distinct. Mesal area of 
axillae and scutellum forming nearly hor- 

Volume 19, Xl-?vser 1, 2010 


Figs 1-7. PluToides porteri sp. nov.l, habitus of male; 2, head, frontal \iew; 3, scape, pedicel and first three 
flageUomeres; 4, labium, ventral \ie%v; 5, right maxilla, \'entral \-ie^N-; 6, pronotum and mesoscutum, anterolateral 
\ie^v; 7, propleura and prostemum fpst), ventral \ie%v. Scale bars = 0.1 mm. 

izontal, posteriorlv directed triangular sur- 
face ^vhich steeplv slants postero-laterallv; 
axillar sutures indicated b}^ conspicuous 
dark, contrnuous line. Prepectus much 
reduced, hidden bv pronotum. H\-poepim- 
eral area of mesopleuron limited inferiorly 

by dark line running from mesopleural 
scrobe to meso-metapleural suture; mese- 
pistemal groove present, as short dark line 
running anteroventrally from mesopleural 
scrobe. Metanotum subrectangular. Meta- 
postnotum broader medially and narrow- 





:^§j — Nest of ants 

Figs 8-14. Pluroides porteri sp. nov. 8, forewing; 9, hind %Ning; 10, seventh metasomal tergum, dorsal \ie\v; 11, 
h\-popygium, ventral view; 12, genital capsule, \entral view, setae depicted on left side only; 13, aedeagiis, 
dorsal view, apophysis (A), lamina (L); 14, trap for m\Tmecophilous insects. Scale bars = 0.1 mm. 

ing at sides, narrowest at level of propo- 
deal spiracles. Propodeum convex, trans- 
verse; propodeal spiracle narrow, removed 
from anterior margin of propodeum by less 
than its length. Wings: forewing 2.5 X as 
long as maximum width. Pterostigma 
swollen apically. Marginal cell with ante- 
rior margin 2.1 X longer than posterior 
margin; anterior and posterior margins 
diverging apically; apical margin 1.6X as 

long as basal margin, latter slightly cur\'ed 
(Fig. 8). Hind wing with vannal (anal) lobe 
3.3 X as long as submedian cell; \vith four 
hamuli (Fig. 9). Legs: slender; tibiae and 
tarsi with weak, sparse setae. Foretibia 
with about 13 spiniform setae on outer 
surface, mainly on apex; spiniform setae 
fewer on mid tibia and absent on hind 
tibia. Tibial spurs 1-2-2; anterior tibial spur 
with approximately 21-22 spines. 

Volume 19, Number \, 2010 


Metasoma: in dorsal view 2.1 X longer 
than maximum width, tapering apically. 
Cercus well developed. Genital capsule as 
in figure 12; aedeagus with lamina sur- 
passing apophyses basally (Fig. 13). 

Etymology. — The species is named after 
Charles C. Porter, distinguished hymenop- 
terist, who has greatly contributed to the 
knowledge of neotropical wasps, and who 
has participated in the collection of Plu- 
mariidae in the field. 

Type material. — Holotype male: ARGENTINA: 
province of La Rioja, Ruta 7, 25 Km East of 
Anillaco, 850 m a.s.l., at light, 22-11-2006, col. P. 
Fidalgo & G. Fidalgo (MACN). The foUowing 
are paratypes. La Rioja: 1 male, same data as 
holotype (MACN); 1 male, Ruta 7, 7 Km East of 
Anillaco, 1200 m a.s.l., at light, 17-n-2006, col. J. 
Torrens & P. Fidalgo (IFML); 2 males, 5 Km 
South of Udpinango, 1000 m a.s.l., at light, 21-0- 
2006, col. P. Diez, J. Torrens & P. Fidalgo 
(MACN, sHde); 1 male, Santa Teresita, 736 m 
a.s.l., at light, 18-0-2006, coU. P. Fidalgo & G. 
Fidalgo (IFML); 1 male, Ruta 40, Km 395 
(between San Bias de Los Sauces and Pituil), 
1230 m a.s.l., at ligth, 9-Xn-06, col. P. Fidalgo 
(MLP). San Juan: 1 male, Ruta 141, Km 173 near 
Caucete, 580 m a.s.l., at Hght, 14-1-2006, col. P. 
Fidalgo (MLP). Catamarca: 33 males, Ruta 46, 
km 64, entre Belen y Andalgala, 965 m a.s.l., at 
light, 1/2-XI-06, col P. Fidalgo & P. Diez, 
(MACN); 3 males, Ruta 46, Km 204, East of 
Belen, 965 m a.s.l., at Hght, 6-Xn-06, col. C. 
Nieto, G Fidalgo & P. Fidalgo (MACN); 3 
males, Ruta 46, Km 64/66 (between Andalgala 
and Belen), 965 m a.s.l., pit-fall traps and trap 
for myrmecophiles, 2-XI-06 / 6-Xn-06 (MACN). 

Distribution. — ARGENTINA: Catamarca, 
La Rioja and San Juan provinces. 

Comments. — This new species has been 
collected in moderate quantities (15 speci- 
mens per night) between the localities of 
Belen and Andalgala in Catamarca pro- 
vince and in minor quantities (one or two 
specimens per night) in different localities 
of San Juan and La Rioja provinces. Three 
specimens of P. porteri, together with two 
of Plumarius sp. and two of Plumaroides 
andalgalensis , were obtained in a trap 
specially designed to catch emerging myr- 

mecophilous insects from the nest (Fig. 14) 
of an undetermined species of ant of the 
genus Acromyrmex Mayr (Formicidae, At- 
tini). It is not clear yet whether the life cycle 
of plumariids may be related to ant nests, 
or if they use these nests merely as 
emerging routes. 


The relationships among genera of Plu- 
mariidae were studied by Roig-Alsina 
(1994), based on 13 morphological char- 
acters. His study found that Plumarius and 
Myrmecopterina form a clade which is the 
sister-group to other plumariids {Myrme- 
copterinella (Plumaroides Maplurius)). Car- 
penter (1999) reanalyzed the data pre- 
sented by Roig-Alsina, adding four new 
characters taken from Brothers (1974) and 
Day (1977), supporting the relationships 
previously found. We present here a more 
comprehensive analysis, including the new 
genus Pluroides, as well as the recently 
described genus Mapluroides. The analysis 
is based on 32 morphological characters, 
and considers species as terminal taxa for 
the ingroup, not genera as in previous 
studies. Other families of Chrysidoidea 
have been used for outgroup comparison, 
since the Plumariidae is the sister group to 
all other chrysidoids. For this purpose the 
phylogeny of the superfamily presented by 
Carpenter (1999) was taken into account, 
and the ground-plan states established for 
the superfamily were used to polarize 
characters within the Plumariidae. Char- 
acters not studied by Carpenter (1999) 
were polarized in a similar way, through 
comparison with other families of Chrysi- 
doidea (Table 1). 

Species examined for this study are 
Plumarius hirticomis (Andre), Plumarius stria- 
ticeps (Andre), Plumarius spp. (several uni- 
dentified species from xeric western Argen- 
tina), Myrmecopterina filicornis Bischoff, Myr- 
mecopterina sp. from Northern Cape Pro- 
vince, South Africa, Maplurius spatulifer 
Roig-Alsina, Plumaroides andalgalensis Broth- 
ers, Plumaroides brother si Diez and Roig- 


Journal of Hymenoptera Research 

Table 1. Data matrix for the 32 characters used in the phylogenetic analysis. 

other Chrysidoidea 00000000 000000000 

Plumarius hirticomis 1 00111200 110000001 

Plumarius striaticeps 1 00111200 110000001 

Myrinecopterina filicomis 1 00010000 110000001 

Myrmecopterinn sp. 1000100000110000001 

Myrmecopterinella okahandja 4 20000011 1 001007110 

Maplurius spatulifer 3 10011101 1 001101200 

Pluroides porteri 3 11000101 1 001101200 

Mapluroides ogloblini 2 11000101 1 001112200 

Plumaroides andalgalensis 2 11000101 1 001111200 

Pluniaroides brothers! 2 11000101 1 001111200 




10 10 10 

110 1 




1 2 1 

1 1 1 

1 1 1 

1 1 1 



10 1 






2 1 



Alsina, Plumaroides typhlus Diez, Mapluroides 
ogloblini Diez, Fidalgo and Roig- Alsina, and 
Pluroides porteri n. sp. Character states for 
Myrmecopterinella okahandja Day were taken 
from the literature, because specimens were 
not available for study. 

List of characters (based on the male sex) 

1. Labial palpus. Four segments (0). 
Three segments (1). Two segments 
(2). One segment (3). Absent (4). 

2. Maxillary palpus. Six segments (0). 
Five segments (1). Three segments (2). 

3. Clypeus, apical margin. Straight or 
projecting, not bent backwards (0). 
Weakly emarginate, bent backwards (1). 

4. Antennal socket. Removed from epi- 
stomal suture by one socket diameter or 
less (0). Removed from epistomal su- 
ture by 1.5 socket diametes, or more (1). 

5. Antennal pedicel and flagellomeres 
1-10, vestiture. Clothed with short 
setae, at most 0.25 times thickness of 
flagellomeres (0). Clothed with long 
setae, as long as or longer than 
thickness of flagellomeres (1). 

6. Antennal pedicel and flagellomeres 
1-10, transverse rows of setae. Absent 
(0). Present (1). 

7. Antennal scape. Simple, without 
swellings or projections (0). With 
apico-ventral projection, varying from 
a distinct swelling to a digitiform 









projection (1). With a basiventral 

enlargement (2). 

Number of flagellomeres. Eleven (0). 

Ten (1). 

Occipital carina. Present (0). Absent (1). 
Pronotal collar. Present (0). Absent (1). 
Ventral angle of pronotum. Rounded 
(0). Pointed (1). 

Dorsal area between propleural scler- 
ites. Membranous (0). Anterior por- 
tions of propleura expanded dorsally 
forming tubular neck (1). 
Epimeral area of propleuron. Present, 
set off by sulcus above forecoxa (0). 
Absent (1). 

Pronotal lobe. Globose, posteriorly 
rounded (0). Flattened, posteriorly 
truncate (1). 

Pronotal lobe. Posterior margin of prono- 
tum continued laterally around pronotal 
lobe (0). Posterior margin of pronotum 
laterally forming carina superimposed 
on pronotal lobe, giving to it bilobate 
aspect (Fig. 6 in Diez et ah 2007) (1). 
Prosternum, ventral view. Well devel- 
oped, with distinct apophyseal pit (0). 
Visible as triangular sclerite, without 
apophyseal pit (1). Reduced, scarcely 
visible (2). 

Prepectus. Well developed, broadest 
medially and with carinate margins 
(0). Reduced, as slender bar (1). Re- 
duced, upper half narrow, widest at 
top, and lower half filiform (2). 





18. Scutelluir-. Xormal, flat (0). Produced 
poster ?-d?r5 =."->.' ?..s a sharp edged 

19. \:e: = r:5-:-;" l:r.^c5: mediaZy and 
nairo'^viTLg lO'^var as r r : r c a ea. spira- 
cles; posterior margir. c.isrlnc: C . 
"^Videning to^-rards prorodeal spira- 
cles; posterior iTLaigin distinct medi- 
allv, but limit betw'een metapostnotum 
and propodeum indistinct laterally (1). 

20. Fore%\~ing, second submarginal ceU. 
Present, moderate (0). Present, but 
reduced (1). Absent (2). 

21. Marginal ceU of fore^sing. Anterior 
margin as long as, or longer than 
maximum ^\idth of pterostigma (0). 
Cell ver\- short, anterior margin one 
third to half as long as maximum 
width of pterostigma (1). 

22. Prestigma (first absdsa of Rl). Linear, 
parallel sided except sometimes w id- 
ened at tip (0). Wide, considerably 
widened on apical tiiird (1). 

23. Vannal lobe of hind ^sing. Moderate, 
less than tvsice (1.3-1.6) as long as 
submedian cell (0). Large, more 
than t^sice (Zl— 2.3) as long as sub- 
median ceU (1). Exceedingly large (3.3 
times as long as submedian cell) (2). 
The last state is autapomorphic for 

24. Arolia of mid and hind tarsi. Present 
(0). Absent (1). 

25. Qa^ss. Dentate (0). Simple (1). 

26. Mid-tibia! spurs. Present (0). Absent 

27. Hind femur. Simple, %\-ithout apical 
projection at each side of tibial articu- 
lation (0). Projected at each side of 
tibial articulation (1). 

28. Ventral surface of hind coxa, specialized 
area of setae. Absent (0). Present (1). 

29. First and second metasomal terga. 
Rrst tergiun in dorsal vie\s' narro\\'er 
than second id. First tergum in dorsal 
\ie >v as .'. ide as second (1). 

30. Pirst metasomal sternum, median 
longitudinal keeL Absent (0). Present 

31. Seventh metasomal tergum. Simple, 
■^\"ithout carinae (0). Longitudinally 
carinate (1). With large, flat, sclero- 

tized apical flange as large as 13 of 
tergum (2). 

32. Seventh metasomal tergum. Simple, 
without carinae nor expanded apically 
(0). Longitudinally carinate fl). With 
large, flat, sclerotized apical flange as 
large as 1/3 of tergum (2). 

An exact anal}"sis '^^'. as conducted with the 
program TXT G : . : r : :: et al. 2(X)7) using 
implicit enumeranon. M-oltistate characters 
^vere run as unordered. A single most- 
parsimonious tree resulted ipig. 15 . de- 
picted wiih the aid of the rrc^r^jr. Clados 
(Nixon 1992). The lengthi is -J 5:ers. vritr. 
consistenc}- index r.S5. arid re:er.7:r. index 
0.92. Autap>omorphies are includea m the 
tree to show distinctiveness of the taxa 
(Pig. 15, black squares). "WTien autapomor- 
phies are excluded from the analysis the 
statistics are: length 39 steps, consistenc}" 
index 0.82, and retention index 0.92. Pive 
characters (1, 2, 16, 17, and 25) have states 
^ ambiguous optimizations; these are 
plotted in the figure using the accelerated 
transformation optimization. 

The results of our analysis reinforce the 
support for Ihe recognition of two major 
lineages within the Plumariidae, as sug- 
gested by preidous analyses (Roig-Alsina 
1994; Carpenter 1999). One of the lineages 
is formed by the American Plumarius and 
the African Myrmecopterina, both genera 
dearly monophyletic. 

The ne^N' genus Pluwides and the recently 
described genus Mapluroides, together ^sith 
Maplurius and Plumaroides, belong ir a 
strongly supported Soutii American dade, 
\^hich is the sister-group to the African 
Myrmecopterinella. The geniis Plumaroides 
has three described species, but current 
sur\-eys have revealed tiiat it is a spedose 
group, with several undescribed spedes. In 
the tree it appears as supported by three 
homoplasious states, but at least two of 
them are dearly convergences, and repre- 


Journal of Hymen»ter_-. R£5e.\rck 

Fig. 15. aadogram for genera ar.d species oi Plumariidae. Giaracter numbers are above the hashmarks, arA 
states are sho\NTi below. WTiite squares indicate nonhomoplasious states gray squares indicate homoplasious 
states, and black squares indicate autapomorphies. 

sent independent deri\*ations: the presence 
of caxinae on the seventh tergiun, and the 
\-er\' small marginal cell. The other three 
genera in this South American clade are 
monot\-pic, supported by their owti auta- 
pomorphies. The new genus Pluroides is 

distinguished by the extremely 

large van- 

nal (anal) lobe of the hind ^ving, and the 
large, flat, polished flange of the seventh 
tergum, one third as long as the tergum. 


This work was financed by PIP Project 6361 from 
the Consejo Nadonal de Investigadones Cientificas y 
Tecnicas, Argentina. We thank Ja\-ier Torrens and 
Gonzalo Hdalgo for their valuable collaboration given 
in the nocturnal surve\3. We also thank ti\e financial 
support of Charles Porter (USDA Florida, Gaines\Tlle, 
USA) who accompanied us on se\eral coUecting trips. 


Bradley, J. C. 1972. Notes or. the distribution of the 
genus Plumarius (H\-menoptera, Plumariidae). 
Entomological Neics 83: 135-139. 

Mothers, D. J. 1974. The genera of Plvunariidae, with 
description of a new genus and spedes from 
Argentina (H\Ti>enoptera: Bethyloidea). Journal of 
tiKentomohgiail Society of southern Africa 37: 351-356. 

. 1975. Phylogeny and classification of the 

aculeate H\-menoptera, with special reference to 
Mutillidae. The University cf Kansas Science Bulletin 

Brues, C. T. 1924. Some South African parasitic 
H\Tnenoptera of the families Exaiuidae, Braconi- 
dae, Alvsiidae and Plumariidae in the South 

African Museum, \%-ith a catalogue of the known 
spedes. Aii->iuls oftiie South .African Museum 19: 1-50. 

Carpenter, J. M. 1999. What do we know about 
chrvsidoid (Hymenc^Jtaa) relationships? Zoolo- 
gica Scripta 28: 215-231. 

Day, M. C. 1977. A new genus of Plimiariidae from 
Southern Africa, \Nith notes on Scoleb\-tiudae 
(H\Tnenoptera: Chr\-sidoidea). Cimbebasi^, A 4: 

Diez, P. 2008. A new spedes of the genus Plumaroides 
Brothers (H\Tnenoptera, Chrvsidoidea, Plumarii- 
dae) from Santiago del Estero, Argentina: male 
and female descriptions. Zootaxa 1891: 25-30. 

, P. Fidalgo, and A. Roig-Alsina. 2007. A new 

ga\us and spedes of Plumariidae rH\TnenoptOTa, 
Chn.-sidoidea) from western \eric .Argentina. 
Zootaxa 1467: 35-41. 

En ans, H. E. 1966. Discover)- of the female Plumarius 
(H\-menoptera, Plumariidae). Psycfie 73: 229-237. 

Goloboff, P., J. Farris, and K. NLxon. 2007. TNT, Tree 
analysis using new technology. Program and 
documentation. Version 1.1. 

Xag\-, C. G. 1973. Re\isionar\- Studies on the Family 
Plimiariidae Bischoff (H\-menoptera, Heterogy- 
noidea). Folia Entomoldgica Hungdrica 26 SuppL: 

XLxon, K. C 199Z Clados, version 121. 

Penteado-Dias, A. M. and D. Scatolini. 2003. A new 
spedes of the genus Plumarius Philippi (H\Tne- 
nc^teia: Plumariidae) from Brasil. Zoologische 
Mededdingeii . Leiden 77: 545-550. 

Perez D'Angelo, \ . 1974. Plumarius coquimbo n. ^. y 
primer registro de la hembra de Plumarius jMffa 
Chile (H\Tnenoptera: Plumariidae). Revista chUena 
de entomologia 8: 139. 

Roig-Alsina, A. 1994. A new genus of Plumariidae, 
\\ith notes on the relationships anK>ng the genera 
of the family. Mitteilungen der MundKner erttomo- 
logisches Gesdlsdu^ 84: 91-%. 

Vol. 19(1), 2010, pp. 121-127 

The Transcaucasian Species of Coccobius Ratzeburg 1852 (Chalcidoidea: 
Aphelinidae), with the Description of Three New Species from Georgia 

George Japoshvili and Ismail Karaca 

(GJ) Entomology and Biocontrol Research Centre, Ilia State University, Tbilisi, Georgia; 

giorgij 70@y ahoo .com 
(IK) Department of Plant Protection, Faculty of Agriculture, Suleyman Demirel University, Isparta, 


Abstract. — The species of Coccobius Ratzeburg from Transcaucasia are listed and Coccobius kato, C. 
nunu and C. otnari are described as new. Information on distribution, synonyms and hosts for eleven 
species is given. A key to the females of the Coccobius species from the Transcaucasus region is provided. 

Key words. — Coccobius, Transcaucasia, Georgia, new species, Parasitoids, Biocontrol 

The genus Coccobius Ratzeburg contains 
81 species worldwide (Noyes 2009). They 
are parasitoids of Diaspididae (Hemiptera: 
Coccoidea) and some of them have been 
used in the biocontrol of harmful pests. For 
example, Coccobius testaceus (Masi) has been 
used as a control agent against Lepidosaphes 
ulmi (Linnaeus) and other armoured scale 
insects in the former USSR and Western 
Europe (Yasnosh 1968). This species was 
introduced into the USA for the control of 
Lepidosaphes beckii (Newman) (Flanders 1942; 
Myartseva 1995). Coccobius varicornis (Flow- 
ard) was used against Diaspidiotus pernicio- 
sus (Comstock), Aspidiotus destructor Cock- 
ereU and other armoured scales in Califor- 
nia. Coccobius odonaspidis (Tachikawa) is a 
host-specific parasitoid of Odonaspis secreta 
(Cockerell) in Japan (Yasnosh 1968). 

The genus Coccobius is rather distinctive 
and is not likely to be confused with any 
other aphelinid genus. All knov^m species 
show a high degree of structural similarity 
(Hayat 1985). Prior to our studies on 
Coccobius from the Transcaucasus, the 
following species were known from the 
region: C. contigaspidis (Yasnosh), C. ephe- 
draspidis (Yasnosh), C. granati Yasnosh & 
Mustafaeva, C. indefinitus (Yasnosh & 
Myartseva), C. mesasiaticus (Yasnosh & 
Myartseva), C. noaeae (Yasnosh), C. pistaci- 

colus (Yasnosh), C. subterraneus (NikoF 
skaya) and C. testaceus (Masi). The genus 
is almost cosmopolitan in distribution and 
recent additions of species from the Pa- 
laearctic, Nearctic, Neotropical, Afrotropi- 
cal and Oriental regions indicate that there 
should be more undescribed species. 

The generic names Physcus Howard, 
Encyrtophyscus Blanchard and Physculus 
Yasnosh were placed in synonymy with 
Coccobius by Hayat (1983), but all the 
species known in these genera were not 
then specifically transferred to Coccobius 
until Hayat's new publication (Hayat 
1985), where he transferred all the species 
to the present genus as new combinations. 


This work is based upon specimens 
deposited in the following collections, with 
abbreviations used in text: St. Petersburg 
Museum of Zoology (ZIN); personal col- 
lection of V. Yasnosh housed in the L. 
Kanchaveli Plant Protection Institute of the 
Georgian Academy of Agriculture (VYC); 
personal collection of the first author, 
housed in the Institute of Zoology of Ilia 
Chavchavadze state University, former 
Institute of Zoology, Georgian Academy 
of Sciences, Tbilisi, Georgia (IZGAS) and 
the Natural History Museum, London, 

122 Journal of Hymenoptera Research 

U.K. (BMNH). Descriptions and terminol- truncation; maxillary palpi two-segmen- 

ogy follow Hayat (1998). ted, labial palpi unsegmented; pronotum 

narrow in dorsal view; median lobe of 

KhbULlb mesoscutum large, with numerous setae; 

^ , . „ . , axilla small with one seta; scutellum large, 

Coccobtus Ratzeburg ^ . . , , j j r 

Coccobius Ratzeburg, 1852: 195. Type: Coccobius P^^*^^^^^ ^^'^f ^^^^ y ^^^^.^^^^' ^^^^ 

anmilicornis Ratzeburg, designated by Gahan ^^^S ^^^^out Imea calva; disc usually 

and Faean 1923- 37. setose to base; submarginal vein with 4 or 

Physcus Howard, 1895: 43. Type: Coccophagus n\ore setae; tarsal formula 5-5-5; mid tibial 

varicornis Howard, by monotypy. spur large (compared to other aphelinid 

Encyrtophyscus Blanchard, in De Santis, 1948: genera, except Marietta Motschulsky); gas- 

192. Type: Physcus flavoflagellatus De Santis, ter at least as long as thorax; hypopygium 

by original designation. not reaching apex of gaster. 

Physculus Yasnosh, 1977: 1115. Type: Physculus M^le: similar to female, except antenna 

rffl«2/gfle Yasnosh, by original designation. ^-^j^ g segments (1.1.6) and genitalia. 

Diagnosis. — Female: antenna with one which is elongate, phallobase almost trun- 

anellus between pedicel and Fl and with cate, narrowed in basal third and with apex 

7 segments; mandible with two teeth and bifid (Hayat 1985). 


1. Brachypterous subterraneus (Nikorskaya) 

Macropterous 2 

2. Antenna uniformly coloured 3 

Antenna differently coloured 4 

3. Clypeus margin with brown band, mesoscutum with medial vertical line 

omari Japoshvili sp. n. 

Clypeus margin without brown band, mesoscutum entirely fulvous 

ephedraspidis (Yasnosh) 

4. Antennae with Fl and clava very slightly dusky nunu Japoshvili sp. n. 

Antennae with Fl and clava dark brown 5 

5. Thorax and usually the head, mostly yellow to brownish-yellow, with minimum of 

dark markings 6 

Thorax and usually the head, mostly dark brown to black 9 

6. Fl slightly shorter than than pedicel, clava longer than funicle 7 

Fl usually somewhat longer than pedicel or subequal, clava usually shorter than 

funicle testaceus (Masi) 

7. Fl subquadrate or very slightly longer than wide .... granati Yasnosh et Mustafaeva 
Fl almost 1.5X as long as wide 8 

8. F3 1.25X as long as wide. Clava length equal to all funicle length together 

indefinites (Yasnosh & Myartseva) 

F3 1.56X as long as wide. Clava slightly longer than funicle . . . kato Japoshvili sp. n. 

9. Submarginal vein at most with 6 and marginal vein with 9 setae 10 

Submarginal vein with at least 7 and marginal vein with 10 setae contigaspidis (Yasnosh) 

10. Marginal fringe at the apex of fore wing almost 0.25 X as long as width of forewings, 

mesoscutum and scutellum with small cellular sculpture . . . pistacicolus (Yasnosh) 
Marginal fringe at the apex of forewings shorter than 0.25 X as long as width of 

forewings, mesoscutum and scutellum with bigger cellular sculpture 

noaeae (Yasnosh) 

Volume 19, Number 1, 2010 


Coccobius contigaspidis (Yasnosh 1968) 

Distribution. — Armenia (Yasnosh 1968). 

Host. — Contigaspis kochiae Borchsenius 
(Diaspididae) (Yasnosh 1968; Ben-Dov et 
al. 2008). 

Coccobius ephedraspidis (Yasnosh 1968) 

Distribution. — Georgia, Turkmenistan 
(Yasnosh 1968). 

Host. — Dynaspidiotus ephedrarum (Lindin- 
ger) (Diaspididae) (Yasnosh 1968; Ben-Dov 
et al. 2008). 

Coccobius granati Yasnosh & Mustafaeva 

Distribution. — Azerbaijan (Yasnosh and 
Mustafaeva 1992). 

Host. — Lepidosaphes granati Koroneos 
(Diaspididae) (Yasnosh and Mustafaeva 

Coccobius indefinitus (Yasnosh & Myart- 
seva 1972). 

Distribution. — Armenia, Tajikistan, Turk- 
menistan (Yasnosh and Myartseva 1972). 

Host. — Chlidaspis asiatica (Archangels- 
kaya) (Diaspididae) (Yasnosh and Myart- 
seva 1972). 

Coccobius noaeae (Yasnosh 1968) 

Distribution. — Azerbaijan, Georgia (Yas- 
nosh 1978). 

Host. — Duplachionaspis noaeae (Hall) (Dia- 
spididae) (Yasnosh 1978; Ben-Dov et al. 

Coccobius pistacicolus (Yasnosh 1968) 

Distribution. — Azerbaijan, Georgia (Yas- 
nosh 1978). 

Host. — Lepidosaphes pistaciae Archangels- 
kaya (Diaspididae) (Yasnosh 1978). 

Comments. — Dr V. Yasnosh recorded 3 
females (Yasnosh 1968), which we have not 
been able to find, unfortunately, in St. 
Petersburg (ZIN) or in her personal collec- 
tion. There has also been no record of this 
species since then. We suppose that this 
species could be a synonym of C. testaceus. 
However, further study is needed to verify 
whether this species is a junior synonym of 
C. testaceus. 

Coccobius subterraneus (Nikol'skaya 

Distribution. — Armenia (NikoTskaya and 
Yasnosh 1966). 

Host. — Chortinaspis subterranea (Lindin- 
ger) (Diaspididae) (Nikol'skaya and Yas- 
nosh 1966; Ben-Dov et al. 2008). 

Coccobius testaceus (Masi, 1909) 

Distribution. — Azerbaijan, Croatia, Czech 
Republic, Egypt, France, Georgia, Ger- 
many, Hungary, Iran, Italy, Lebanon, Mol- 
dova, Montenegro, China, Poland, Roma- 
nia, Spain, Turkey, Ukraine, UK, USA, 
Uzbekistan (Noyes 2009). 

Hosts. — Chionaspis salicis (L.), Chlidaspis 
asiatica (Archangelskaya), Contigaspis ko- 
chiae Borchsenius, Diaspidiotus prunorum 
(Laing), D. transcaspiensis (Marlatt), D. gigas 
(Thiem & Gerneck), D. jaapi (Leonardi), D. 
perniciosus (Comstock), D. slavonicus 
(Green), Furchadaspis zamiae (Morgan) 
(Noyes 2009; Ben-Dov et al. 2008), Lepido- 
saphes beckii (Newman) (Myartseva 1995), 
L. conchiformis (Gmelin), L. malicola Borch- 
senius, L.ulmi (L.), Lineaspis striata (News- 
tead), Parlatoria oleae (Colvee), Eriococcus 
spurious (Modeer) (Noyes 2009; Ben-Dov et 
al. 2008). 

Coccobius kato Japoshvili sp. n. 

Material Examined. — Holotype, 9, GEORGIA: 
Vashlovani, ex Lineaspis striata (Newstead) on 
Thuja sp. 2.VI. - 15.VII.2003, G. Japoshvili 
(IZGAS). Paratypes, 3 99, same data as holo- 
type (IZGAS). Homoeotype: 9 with label: Phys- 
cus sp. ajf. pistacicolus Jasnosh ex Lineaspis 
striata collected on Juniperus foetidissima, 

Female.— Length, 0.44-0.70 mm (Holo- 
t3^e: 0.56 mm). 

Entire body yellow, legs and antenna 
slightly paler. Fl slightly brownish and 
clava brown. Head 1.8 X as wide as FV. 
Head in front view almost as wide as high. 
Ocelli with apical angle obtuse. Eyes long, 
2x as long as malar space. Toruli with 
upper margins level with lower eye mar- 
gins. Antenna as Fig. lA. Pedicel 1.8X, Fl - 
1.56X, F2 - 1.4X, F3 - 1.45X, CI - 1.2X, C2 


Journal of H^aiexoptefla Rese.arch 

Fig. 1. Coccobius kato Japoshvili, sp.n. antenna (A), left fore wing (B), ovipositor (C); C. omari Japoshvili, sp.n. 
ovipositor (D), antenna (F), left fore wing (G); C. nu7iu Japoshvili sp.n. ovipositor (E), antenna (H), right fore 
wing (I). 

- 2x as long as wide. Scape slightly more 
than 4x as long as wide. Mesoscutum (26) 
longer than scutellum (22). Scutellum 1.3X 
as wide as long. 

Fore wings 2.6X as long as wide 
(Fig. IB). Marginal fringe about 0.25 X as 
long as wing width. Submarginal: mar- 
ginal: stigmal veins as following 37:27:7. 
Hind wing slightly more than 5.5 X as long 

as wide. Marginal fringe almost equal to 
that of wing width. Gaster slightly shorter 
(11:14) than head plus thorax. Pygostyles 
inserted at the 0.6 distance from base. 
Ovipositor (Figure IC) second valvifer 
0.7X as long as ovipositor and third 
valvula 0.4 X as long as second valvifer. 
CKipositor 1.5X longer than midtibia. 
Male. — Unknown. 

Volume 19, Number 1, 2010 


Comments. — The new species is close 
to C. indefinitus Yasnosh & Myartseva, 
but differs by the characters given 
in Table 1. The species mentioned as 
Physcus sp. aff. pistacicolus by Yasnosh 
(1972), is probably conspecific with this 
new species. 

Coccobius omari Japoshvili, sp.n. 

Material Examined. — Holotype, 9, GEORGIA: 
Tbilisi, Gldanula, ex Prodiaspis tamaricicola (Mal- 
enotti) on Tamarix sp. 28.VI. - 20.Vn.l998, G. 
Japoshvili (IZGAS). Paratypes, IO99, same data 
as holotype (IZGAS). 

Female.— Length, 0.62-0.89 mm. (Holo- 
type: 0.67 mm). 

Head with metallic lustre infuscated 
yellow or testaceous-brownish yellow. 
Anterior part of pronotum, mesoscutum 
in the middle vertically, and tergites, with 
hardly noticeable brownish lines. All body 
yeUow. Femora and tibia brownish. Face 
above clypeus with dark brown band 
joining to eyes on margin. Antennae uni- 
formly coloured. 

Head 2.1 X as wide as FV. Head in front 
view almost 1.13X as wide as high. Ocelli 
with apical angle obtuse. Eyes 1.2X as long 
as malar space. Toruli with upper margins 
below lower lower eye margins. Antennae 
as Figure IF. Pedicel 2X, Fl - 1.27X, F2 - 
1.4X, F3 - 1.33X, CI - 1.29X, C2 - 2.33X as 
long as wide. Scape slightly more than 4X 
as long as wide. Mesoscutum (25) longer 
than scutellum (20). Scutellum 1.5X as wide 
as long. Fore wings 2.3 X as long as wide 
(Figure IG). Marginal fringe about 0.2 X the 
length of wing width. Submarginal: mar- 
ginal: stigmal veins as follows: 38:24:5. Hind 
wing slightly more than 5.16 X as long as 
wide. Marginal fringe almost 0.6 X as long 
as wing width. Caster slightly longer (15:13) 
than head plus thorax. Pygostyles inserted 
at the 0.74 distance from base. Ovipositor 
(Figure ID) second valvifer 0.7X as long as 
ovipositor and third valvula 0.41 X as long 
as second valvifer. Ovipositor 1.76X longer 
than midtibia. 

Male: Unknown. 

Comments: The species is close to C. 
sumbarensis Myartseva but differs by the 
morphological characters given in Table 1., 
also C. sumbarensis is entirely brown, Fl 
and clava also brown. The new species 
when it has some brownish coloration, 
then it is hardly noticeable. 

Etymology: This species is named in 
honour of the Georgian ichthyologist and 
zoologist, Dr Omar Japoshvili. 

Coccobius nunu Japoshvili, sp.n. 

Material Examined. Holotype, 9, GEOR- 
GIA: Tbilisi, Gldanula, ex Prodiaspis tamar- 
icicola (Malenotti) on Tamarix sp. 28.VI. - 
20.Vn.l998, G. Japoshvili (IZGAS). Para- 
types, 3O99, same data as holotype (IZGAS). 

Female: Length, 0.67-0.84 mm. (Holo- 
type: 0.76 mm). 

Coloration of body similar to that of 
Coccobius omari sp.n., the only difference 
being that this species has some brownish 
areas on the body. Head 1.85X as wide as 
FV. Head in front view almost 1.15X as wide 
as high. OceUi with apical angle obtuse. Eyes 
slightly more than 2 X as long as malar space. 
Toruli with upper margins level with lower 
eye margins or slightly lower. Antennae as 
Figure IH. Pedicel 2X, Fl - 1.67X, F2 - 
1.45X, F3 - 1.36X, CI - 1.54X, C2 - 2.9X as 
long as wide. Scape 3.4 X as long as wide. 
Mesoscutum (29) longer than scutellum (25). 
ScuteUum 1.36X as wide as long. Forewings 
2.3 X as long as wide. Marginal fringe about 
0.14 X of wing width. Setation and venation 
as Figure II. Submarginal: marginal: stigmal 
veins as foUows: 44:29:6. Hind wing 5.2 X as 
long as wide. Marginal fringe almost 0.57 X 
as long as wing width. Caster longer (18:15) 
than head plus thorax. Pygostyles inserted at 
the 0.67 distance from base. Ovipositor 
second valvifer 0.75 X as long as ovipositor 
and third valvula 0.37X as long as second 
valvifer. Ovipositor (Figure IE) 1.67X 
longer than midtibia. 

Male. — Unknown. 

Comments. — The new species is close to C. 
kurbani Myartseva, but differs by the char- 
acters given in Table 1. This species is also 


Journal of Hymenoptera Research 


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VoLL-^iE 19, Xl-mber 1, 2010 


close to C. viggianii Yasnosh (Yasnosh 1974) 
and C. fiircus Huang (Huang 1994), but 
differs from both by the sculpture on 
mesoscutum and scutellum.. The sculpture 
in C. viggianii is ^vith elongated cells in the 
middle on both mesoscutum and scutellum 
and C. fiircus with polygonal ceUs on both 
mesoscutum and scutellum, but in C. mimi 
sp.n. the sculpture on the mesoscutum is 
polygonal and on the scuteUum a little 
elongated. Also, there are less than 10 setae 
on the submarginal vein in C. viggianii and 
C. fiircus, \vhile there are at least setae on C. 
nunu is at least 10. 


We did not include Coccobius mesasiaticus 
(Yasnosh and Myartseva 1971) in the key 
and Hst of species of Coccohius of the 
Transcaucasian region. Coccohius mesasiati- 
cus ^vas described from Turkmenistan, and 
mentioned as also being distributed in 
Azerbaijan (Yasnosh 2000), ho^vever it was 
impossible to find any material from Azer- 
baijan to confirm its distribution there. 
Yasnosh (2000) cited distributional data 
based on Mustafaeva's doctoral thesis, 
but later Rzaeva (2002), in her monographic 
work on chalcidoids of Azerbaijan, did not 
mention this species. Also, G. Japosh\ili 
could not find any information about the 
material from Azerbaijan (Mustafaeva pers. 
comm.). We would like to note that dis- 
tributional data concerning Afganistan and 
Iran (Hayat 1985) is also doubtful, in as 
much as Hayat did not mention the exam- 
ined material and sources of data. 


Special thanks to Dr Takumasa Kondo for c±iecking 
tiie English to promote this paper. 


Ben-Dov, Y. D. R. Miller, and G. A. P. Gibson. 2008. 

ScaleXet http:/ /\N-%\-\ 

Flanders, S. E. 1942. The introduction of Physcus 

testaceiis Masi into California. Journal of Economical 

Entomology 35: 290-291. 

Hayat, M. 1983. The genera of Aphelinidae (H\-me- 
noptera) of the World. Svstejimtic Entomology 8: 

. 1985. Notes on some species of Coccohius and 

Prophysais (H}Tnenoptera: Aphelinidae), with 
special reference to Girault and Ho%vard t\-pes. 
Oriental Insects 18: 289-334. 

. 1998. Aphelinidae of India (H\Tnenoptera: 

Chalcidoidea): a taxonomic Re\*ision. Memoirs on 
Entomology, Intenmtional 13: 1—416. 

Huang, J. 1994. Systematic studies on Aplielinidae of 
China. Chonqing publishing house, China. 

Myartseva, S. X. 1995. Xe%v spedes of Aphelinids 
(H\Tnenoptera, Aphelinidae) - parasites of scale 
insects on Tamarix in Turkmenia. Ejitomological 
Rerieic 1\: 432-440 (In Russian). 

Nikol'skaya, M. and V. Yasnosh. 1966. Aplielijiids of 
European part of USSR and Caucasus. Xauka, 
Leningrad. 295 pp. 

Noyes, J. 2009. Universal Chalcidoidea Database (on- 
line), http: / /\N'\ 
research/projects / chalddoids / . 

Rzaeva, L. 2002. Clmlcids of eastern Azerbaijan and their 
economic importance. ELM, Baku. 354 pp. 

Yasnosh, V. A. 1968. Spedes of the genus Physcus 
Ho^vard (H\-menoptera, Chalddoidea), parasites 
of scale insects in the USSR. Ejitomologicheskoe 
Obozraiie 47: 205. 

. 1972. Chalddoid parasites (H\-menoptera, 

Chalddoidea) of scale insects (Homoptera, Coc- 
coidea) from Georgian arid light forests. Proceed- 
ings of All-union Entomological society. Xauka, 
Leningrad. 55: Pp. 217-247. 

. 1974. A new spedes of Physcus Ho^vard 

(H}Tnenoptera, Aphehnidae) from Italy. Bolletino 
del laboratorio di Entomologia Agraria "di Portici" 
31: 105-108. 

. 1978. ApheUnids. Pp. 469-501 in: Med\-edev, G. 

S. ed. Keys to tlie Insects of the European part of USSR. 
Hymenoptera Vol. III. Part 2. Xauka, Leningrad. 

. 2000. Aphelinid parasitoids (H\-menoptera, 

Aphelinidae) of armored scale insects (Cocdnea, 
Diaspididae) in the Caucasus. Proceedings of tlie 
Institute c^ Zoology, Academy of Sciences of 
Georgia 20: 159. 

and G. A. Mustafaeva. 1992. Xe^v parasitoid of 

Lepidosaphes granati, Coccobius granati sp. n. 
(H\"menoptera, Aphelinidae). Zoologichesldy Zhur- 
iml 71: 142-143. 

and S. X. Myartseva. 1971. Two New aphelinid 

spedes (Chalddoidea, Aphelinidae) - parasites of 
armoured scales (Coccoidea, Diaspididae) from 
middle Asia. Bulletin of Academy of Sciences of 
Turhnenistan SSR 6: 35-41 (In Russian). 
— and S. N. Myartseva. 1972. New parasite 
of Asiatic scale - Physcus indefinites, sp. n. 
(Chalddoidea, ApheUnidae). Bulletin of Academy 
of Sciences of Turkmenistan SSR 5: 51-55 (In 

Vol. 19(1), 2010, pp. 128-138 

Host Range and Offspring Quantities in Natural Populations of Nasonia 
vitripennis (Walker, 1836) (Hymenoptera: Chalcidoidea: Pteromalidae) 

Ralph S. Peters 

Zoologisches Museum Hamburg, Universitat Hamburg, Martin-Luther-King-Platz 3, 
20146 Hamburg, Germany; 

Abstract. — The gregarious ectoparasitic wasp Nasonia vitripennis is a well-established model 
organism in various aspects of genetics. In the field it is the dominant species in the parasitoid 
community found in birds' nests. In these nests N. vitripennis parasitizes the puparia of 
cyclorrhaphous flies. The most frequently recorded natural field host species are two 
omithoparasitic bird blow flies Protocalliphora azurea and P. falcozi, the necrophagous Calliphora 
vicina (all: Calliphoridae) and the sapro-necrophagous Potamia littoralis (Muscidae). These field host 
records along with additional laboratory rearings allowed us to define the host range of N. 
vitripennis: It is restricted to Cyclorrhapha with further restrictions in size, shape and surface 
structure of the host species. The host acceptance excludes Hippoboscidae, Fanniidae and small 
species like heleomyzids and drosophilids, but includes parasitism of tachinids and sarcophagids 
or facultative hyperparasitism of Alysiinae (Hymenoptera: Braconidae) in calliphorid priniary host 
puparia. The mean number of parasitoids emerging from field-collected specimens of the four 
regular host species ranged from 9.3 to 25.7 and the sex ratio was female-biased with proportions of 
males ranging from 0.13 to 0.34. This study shows that there are significant differences between 
host species in the number of individuals that emerged and the proportion of males. A correlation 
betw^een the number of individuals and host size was indicated, but not between host size and sex 

Nasonia vitripennis (Walker, 1836) (Hy- 
menoptera: Pteromalidae) is a gregarious 
cosmopolitan idiobiontic parasitoid species 
and one of the most prominent model 
organisms in speciation and developmen- 
tal genetics (e.g. Gadau et al. 1999; Perfectti 
and Werren 2001; Werren et al. 2004), but 
little is know^n about its natural life history. 
The most comprehensive work on this 
species is still the review by Whiting 
(1967), focused on behaviour and labora- 
tory tests. In Europe Nasonia vitripennis is 
the only species of the genus Nasonia, in 
Northern America two additional species 
are present: Nasonia giraulti Darling, 1990 
and Nasonia longicornis Darling, 1990 (Dar- 
ling and Werren 1990). 

Nasonia vitripennis has been frequently 
recorded from birds' nests (e.g. Abraham 
1985; Darling and Werren 1990; Molbo and 

Parker 1996; Peters and Abraham 2004) 
and repeatedly recorded from carrion (e.g. 
Blanchot 1995; Grassberger and Frank 
2004; Marchiori 2005). Carrion, as a habitat, 
overlaps with birds' nests as dead birds 
can be regularly found in nests. Within 
both habitats, N. vitripennis parasitizes 
numerous species of cyclorrhaphous flies 
such as Calliphoridae and Sarcophagidae. 
Records from dung show that N. vitripennis 
is not a dominant species in this habitat 
(Floate et al. 1999; Skovgard and Jespersen 
2000; Kaufman et al. 2001; Birkemoe et al. 
2008) and cannot be considered an eco- 
nomically important antagonist of synan- 
thropic flies (Legner 1967; McKay and 
Galloway 1999). 

Abraham (1985) and Abraham and Pe- 
ters (2008) hypothesized that birds' nests 
are the primary habitat of N. vitripennis. 

Volume 19, Number 1, 2010 


Peters (2007) corroborated that the habitat 
specialist N. vitripennis is the key species in 
the birds' nest parasitoid web. First field 
studies on N. vitripennis in birds' nests 
(Abraham 1985; Schlein 2002) were mostly 
made with sentinel replacement host pu- 
paria, which helped to reconstruct the 
parasitoids' phenology but gave little evi- 
dence regarding the natural hosts appear- 
ing in the nests. Subsequent studies (Peters 
and Abraham 2004; Abraham et al. 2005) 
reported field host records and defined a 
preliminary host range of N. vitripennis but 
left differences in life history patterns 
concerning number of offspring and sex 
ratio in relation to host species unclarified. 
Theoretical considerations predict a cor- 
relation between host size and the number 
of offspring produced (Waage 1986; God- 
fray 1994). The correlation between host 
size and number of parasitoids has already 
been described for N. vitripennis under 
laboratory conditions by Wylie (1967) and 
Rivers and Denlinger (1995). In parasitoids 
it is generally advantageous if the sex ratio 
is shifted in favour of the females (Godfray 
1994). However, parasitoid sex ratio is 
influenced by various variables and has 
been subject of numerous studies in the 
past and present. Host-quality (or host- 
size) model predicts that in smaller, less 
suitable hosts, more males are produced as 
the males' fitness is less dependent on host 
quality (Chamov et al. 1981; Waage 1986; 
Ueno 1999). Werren (1983), Molbo and 
Parker (1996) and Grillenberger et al. 
(2008) observed that other factors such as 
the parasitoid population size and the 
number of ovipositing females exert a 
strong influence on sex ratio which might 
suppress the predicted correlation between 
sex ratio and host size. These results are 
connected to local mate competition (LMC) 
theory (Hamilton 1967). LMC describes 
that male-male competition for mates is 
restricted to the natal patch. N. vitripennis 
exhibits typical characteristics predicted 
for a species with LMC: Sex ratio is highly 
female-biased if females oviposit alone. 

more males are produced by superparasi- 
tizing females to maximize the opportu- 
nities of their offspring to be able to mate 
(Werren 1980). To a lower extent asym- 
metric larval competition between sexes 
(Sykes et al. 2007) and characteristics of 
mating males (Shuker et al. 2006) have 
been demonstrated in laboratory studies to 
influence sex ratio in N. vitripennis. Sex 
ratio in relation to field host species of 
different size might give further insight 
into sex ratio mechanisms in N. vitripennis. 
In contrast to previous studies, this 
study presents field data on offspring 
quantities of N. vitripennis reared from 
different natural host species collected as 
puparia from birds' nests. Data are exam- 
ined for any differences between host 
species in number of individuals per host 
and proportion of males and for any 
correlations between the size of host 
puparia and the number of parasitoids 
reared and between host size and sex ratio. 
Furthermore, this study utilized field col- 
lection of host puparia and additional 
laboratory rearings to define the host range 
of N. vitripennis. These results can circum- 
scribe host acceptance and rejection cues of 
N. vitripennis such as shape and size of the 
host which are known to be used by 
parasitoids (Wylie 1967; Cooperband and 
Vinson 2000). 


Nests of Parus spp. (tits) were examined 
for the presence of puparia of cyclorrha- 
phous flies parasitized by Nasonia vitripen- 
nis. The nine collection sites were located 
in Germany and included: Hamburg (5 
sites), (1) Hamburg-Eissendorf, "Staats- 
forst Haake" forest N53.4548 E09.9207, (2) 
Hamburg-Eissendorf, garden N53.4540 
E09.9391, (3) Hamburg-Rotherbaum, "Stem- 
schanzenpark" N53.5655 E09.9709, (4) Ham- 
burg-Bramfeld, "Umweltzentrum Karls- 
hohe" park N53.6295 E10.1092, (5) Ham- 
burg-Poppenbiittel (garden in the district 
of Poppenbiittel without exact locality); 
Schleswig-Holstein (3 sites), (6) "Linauer 


Journal of Hymenoptera Research 

Forst" forest N53.6734 E10.4897, (7) Elms- 
horn, garden N53.7656 E09.6725, (8) Hasel- 
dorf, garden N53.6352 E09.5986; Baden- 
Wiirttemberg (1 site), (9) Bad Mergentheim, 
''Stadtw^ald" forest N49.5083 E09.7722 (re- 
ference system WGS84). 

Puparia were stored in Petri dishes at 
room temperature until the emergence of 
parasitoids. These were preserved in 70% 
EtOH in pools of multiple puparia or in 
single puparium tubes. The latter will be 
referred to as "separately stored puparia" 
within this paper. The corresponding host 
puparia were taped to the EtOH tubes. The 
parasitoids that emerged were counted 
and sex was determined. Maximum length 
and maximum width of host puparia were 
measured. For puparia species identifica- 
tion voucher specimens of puparia and 
corresponding identified imagines were 
used that were collected during our studies 
on birds' nest fly fauna (Peters and Abra- 
ham 2004; Peters 2007). Voucher specimens 
are deposited at Zoologisches Museum 
Hamburg (ZMH). 

A laboratory stock of N. vitripennis was 
maintained on Calliphora vomitoria (Lin- 
naeus, 1758) puparia in Petri dishes at 
room temperature. The stock was origin- 
ally reared from Proto calliphora azurea 
(Fallen, 1816) puparia collected from a nest 
of Panis sp. in Hamburg-Rotherbaum. For 
laboratory rearing tests on host range 
parasitoid females were put on puparia or 
pupae of a variety of Diptera and Lepi- 
doptera species. Parasitoids were taken 
from the laboratory stock. They were 3- 
6 days old, mated and fed on moistened 
raisins. 10 females were put on 30 to 50 
puparia or pupae until they died. Hosts 
included puparia of Triarthria setipennis 
(Fallen, 1810) (Diptera: Tachinidae) reared 
from Forficula auricularia Linnaeus, 1758 
collected in Hamburg-Rotherbaum, pu- 
paria of Protophormia terranovae (Robi- 
neau-Desvoidy, 1830), Calliphora vomitoria, 
Lucilia sericata (Meigen, 1826) (all: Diptera: 
Calliphoridae; obtained as larvae from bait 
shops) and Drosophila melanogaster Meigen, 

1830 (Diptera: Drosophilidae) and pupae of 
Galleria mellonella (Linnaeus, 1758) (Lepi- 
doptera: Pyralidae) presented with their 
cocoons. G. mellonella larvae were reared 
on artificial medium containing cereals, 
glycerin, milk powder and honey. AH hosts 
were 4-6 days old. In 2 of the 4 tests with 
D. melanogaster >100 puparia of different 
ages from a laboratory mass-rearing were 

Additionally, parasitoids were reared on 
freeze-killed C. vomitoria puparia, which 
were stored in -28°C 4-6 days after 
pupation and thawed before a new rearing. 
All rearings were made in Petri dishes at 
24-26 ^C. If the rearing resulted in a viable 
Fl, it was considered successful, regardless 
of the number of parasitoid specimens 
reared. If it did not result in any offspring 
after four attempts, it was considered 

All data analyses were performed with 
SPSS 16.0 for Windows. Tests on normal 
distribution of data were made with 
Kolmogorov-Smirnov-tests. All signifi- 
cance levels of comparisons were Bonfer- 
roni corrected. 


Host records. — Nasonia vitripennis was 
reared from six host species that were 
collected from birds' nests in the field. 
These included two species of dipteran 
bird parasites Protocalliphora spp., the 
necrophagous Calliphora vicina (all: Dip- 
tera: Calliphoridae) and Sarcophaga sp. 
(Sarcophagidae) and the polyphagous 
muscid Potamia littoralis. Furthermore, in- 
traordinal hyperparasitism of the diptero- 
phagous braconid Alysia manducator was 
recorded. In the laboratory, rearing was 
possible on three additional calliphorid 
host species and on Triarthria setipennis 
(Tachinidae). Calliphora vomitoria puparia 
that were freeze-killed before and then 
thawed for rearing tests are also suitable 
hosts for N. vitripennis. Drosophila melano- 
gaster was the only presented Diptera 
species on which no rearing of N. vitripen- 



Table 1. Host records for Nasonia Tntripennis from field collections and laboraton- rearings; * h^-perparasitism 
in Calliphora viciria. 

Field host records 

Laborator)- rearings 

ProtocaUiphora azurea (Fallen, 1816) (Calliphoridae) 
Protocalliphora falcozi Seguy, 1928 (Calliphoridae) 
Calliphora vicbia Robineau-Des\-oidy, 1830 (Calliphoridae) 
Potamia littoralis Robineau-Desvoidy, 1830 (Muscidae) 
Sarcoplmga sp. (Saicophagidae) 
Alysia manducator (Panzer, 1799) (H\Tn.: Braconidae) * 


Protophormia terranovae (Calliphoridae) 
Calliphora vomitoriu (Calliphoridae) 
Lucilia sericata (Calliphoridae) 
Triarthria seiipeimis (Tachinidae) 
Calliphora vomitoria (freeze-kiUed puparia) 

Drosophila melanogaster (Drosophilidae) 
Galleria mellonella (Lepidoptera: Pyralidae) 

nis was possible. Also, rearing ^vas un- 
successful on the pupae of the greater wax 
moth Galleria mellonella (Lepidoptera: Pyr- 

The recorded field hosts of N. vitripennis 
in birds' nests, potential hosts that were 
confirmed in successful laborator}^ rearings 
and examples of species outside the para- 
sitoids' host range are listed in Table 1. 

Offspring numhers and sex ratio. — Speci- 
mens of iV. vitripennis emerging from the 
host species Calliphora vidua, Protocalliphora 
azurea, P. falcozi and Potamia littoralis \vere 
counted. In total 5333 individuals from 490 
puparia from 17 Parus spp. nests were 
examined (Table 2). All puparia and 
ging parasitoids were used for calculation 
of overall mean of individuals per host and 
an overall sex ratio (Table 2). 140 puparia 
were separately stored and were used for 

comparisons of number of parasitoids per 
host and proportion of males. These 
puparia included Protocalliphora azurea (N 
= 16), P. falcozi (N = 15), C. vicina (N - 72) 
and P. littoralis (N = 37). From these 
puparia 1897 parasitoids emerged. 

Mean comparisons made of the para- 
sitoids of separately stored puparia 
sho^ved that there are significant interspe- 
cific differences in numbers of individuals 
per puparium. The number \vas higher in 
Protocalliphora azurea (median 23.0) and P. 
falcozi (median 22.0) than in C. vicina 
(median 10.0) and P. littoralis (median 8.0) 
(U-test, p < 0.001). The difference between 
neither the two Protocalliphora-species (Li- 
test, p > 0.7) nor between C. vicina and P. 
littoralis (U-test, p > 0.1) ^vas significant. 

For mean comparison of proportion of 
males, puparia from ^vhich only males 

Table 2. The total number, the number per puparium and the proportion of males of Nasonia ritripmnis in 
field host species; * = puparia -VNith emergence of only males excluded; ** = recorded from separately stored 
puparia wdth reduced sample size. 

host speues 

Callxplwra ricimz 

CallipJwra vicina * 

Protocalliplwra azurea 

PTOtocalUpiwra falcozi 

Potamia littoralis 

total no. of host puparia 






total no. of N. vitripennis 






total no. of 






total no. of 5 






overall mean no. per host 






overall mean c 






o\-erall mean £ 






median no. per host ** 





maximimi no. per host ** 





overall sex ratio 3:9 






overall proportion of £ 






median proportion of £ ** 







Table 3. Size of puparia of four field host species of N. vitripmnis. 



ProtoaiUiphcra azurea 

ProtocaUtphora fake:: 

Pctamia Uttaralis 

puparium length (mm) 





interquartile range 
minimum; ma.ximum 

4.6; 9.5 

6.3; 8.4 

7.7; 9.1 

6.0; 7.5 







mean ::: std 

2.37 - 


2.1 = 0.11 

2.14 = 


2.82 = 0.14 

emerged (possibly due to unmated females 
in the arrhenotokous hymenopterans) were 
excluded (only in C. vicma). The only 
significant difference after Bonferroni cor- 
rection is found bet^veen C. vidua (X = 61) 
and P. Uttoralis (U-test, p = 0.001). C. vicma 
puparia exhibit a higher proportion of 
male parasitoids (Table 2). The proportions 
of males in P. falcozi and P. Uttoralis are 
lower than in P. azurea, but these differ- 
ences are not significant (U-test, p = 0.04, p 
= 0.047). There is no difference betv^^een P. 
falcozi and P. Uttoralis (U-test, p > 0.6). The 
difference between C. vidua and P. azurea is 
also not significant (U-test, p > 0.2). The 
overall sex ratio measured as the number 
of females per male was higher in P. falcozi 
and Pctamia Uttoralis than in all other 
species (Table 2; x^ p < 0.001). The lowest 
value compared to all other species was 
found in the C. vicina puparia (p < 0.001). 
There was no difference if the puparia 
from which only males emerged, \vere 
excluded (p > 0.1). 

The host puparia were of different size 
depending on species (Table 3). The pu- 
paria of the Protocalliphora species were 
particularly big, wide and massive. P. 
falcozi puparia are significantly longer than 
P. azurea puparia (U-test, p < 0.001). 
Puparia size was remarkably variable in 
C. vidua. The smallest and the largest 
puparia in the study belong to this species. 
The mean length of C. vidua puparia was 
significantly greater than P. Uttoralis (U- 
test, p < 6.001), but smaller than Proto- 
calliphora spp., although the length differ- 
ence with P. azurea is not significant (U- 

test, p > 0.1). On average the puparia of P. 
Uttoralis are the smallest. Their shape is 
rather slim which is shown in the high 
length to \vidth ratio (Table 3). 


Host rauge. — Dominant field host species 
of xV. vitripeuuis from the specific primary 
habitat (birds' nests) were bird blowfUes 
{Protocalliphora azurea and P. falcozi), the 
necrophagous blowfly Calliphora vidua and 
the polyphagous muscid Potamia Uttoralis. 
The puparia of the parasitic and the 
necrophagous blowflies appear in the nests 
when either Live or dead nestUngs are 
available. The activit}^ of N. vitripeuuis is 
Hnked to the birds' breeding season and 
largely restricted to the summer months 
(Schlein 2002). During this time the para- 
sitoid can use the blowfly hosts, the 
predominantly necrophagous summer 
generation of P. Uttoralis and, additionally, 
other necrophagous taxa occurring less 
frequently like SarcopJiaga spp. (Table 1). 
There is no evidence of parasitism of other 
nidicolous taxa, such as lepidopterans, in 
this study nor in Xoyes (2007). 

Successful rearing on the tachinid Triar- 
thria setipeuuis, a parasitoid of ear^vigs, and 
on other calliphorid species like Calliphora 
vomitoria, Lucilia sericata and Protophormia 
terrauovae supports the general use of 
cyclorrhaphous hosts (Table 1). It demon- 
strated that N. vitripeuuis can parasitize 
taxonomically related hosts, with which 
the species will not have contact under 
natural conditions. Further examples of 
suitable cyclorrhaphous hosts of X. vitri- 

VOLL-ME 19, Nl-mber I, 2010 


pennis that have been recorded in various 
studies are Mtisca domestica Linnaeus, 1758, 
Muscina stabulans (Fallen, 1817) and Sto- 
moxys calcitrans (Linnaeus, 1758) (all: Mus- 
cidae) (e.g. Rivers and Denlinger 1995; 
Blanchot 1995; Gibson and Hoate 2004). 
Rearing on the pupae of the wax moth 
Galleria mellonella was not possible. All 
results imply that Lepidoptera are outside 
the host range of N. vitripennis. The very 
few records of lepidopteran hosts (Noyes 
2007) should be seen as misinterpretations 
or accidental events. 

The first limitation of host range within 
the Cyclorrhapha is host size. Puparia are 
not parasitized if they are too smaU, even if 
they appear in suitable habitats, Hke the 
frequent birds' nest species Tephrodamys 
tarsalis (Zetterstedt, 1847) (Heleomyzidae) 
(Noyes 2007; Peters 2007). It was not 
possible to rear N. vitripennis on Drosophila 
melanogaster in the laboratory, which cor- 
roborates the observ^ations of Rivers and 
Denlinger (1995). A threshold in host 
suitability regarding puparium size might 
be represented by the cheese-fly Piophila 
casei (Noyes 2007) (puparia length 4-5 mm). 
The sm.aUest parasitized puparium in this 
study was 4.6 mm (Table 3). 

A second limiting factor of host range is 
the shape of puparia: Aberrant forms Hke 
the puparia of fanniids, with their conspic- 
uous appendages, are not or not regularly 
parasitized (only two records: Legner et al. 
1967; Blanchot 1995); the almost circular 
louse fly puparia (Hippoboscidae) are not 
suitable hosts either (only one record from 
Pseudolynchia canariensis (De Santis 1967)). 
Like in N. vitripennis, host shape as an 
important host acceptance cue is known 
from e.g. Melittobia digitata (Eulophidae) 
(Cooperband and Vinson 2000). 

A third decisive factor is the host surface 
structure: N. vitripennis hyperparasitizes 
Alysia manducator (Hymenoptera: Braconi- 
dae: Alysiinae) inside Calliphora vicina 
puparia as the surface structure indicates 
a suitable and intact host. The same can be 
foimd to explain the parasitism of freeze- 

killed hosts (Table 1). Although the content 
of the puparium is completely different 
from a Hve pupa in shape and consistency, 
the host is accepted. An examination of the 
host pupae with the ovipositor, as de- 
scribed for N. vitripennis by Edwards 
(1954), is therefore unlikely, at least for its 
impact on host acceptance. The discrimina- 
tion of hosts using cues of puparia and not 
cues of their content was recorded for N. 
vitripennis by Smith (1969). In a more 
general statement Rivers (1996) concluded 
from his studies that host acceptance in N. 
vitripennis is related to exterior cues of the 
puparia. These cues are now specified as 
size, shape and surface structure. 

In summary the host range of Nasonia 
vitripennis is defined as: 

Polyphagous; Cyclorrhaphous Diptera, 
especially Calliphoridae, Muscidae and 
Sarcophagidae at least 4-5 mm in size, 
excluding puparia with appendages (Fan- 
niidae) and aberrant shapes (Hippobosci- 
dae), including hyperparasitism of Alysii- 
nae in suitable host puparia. 

Offspring numbers and sex ratio. — The 
Protocalliphora puparia showed the highest 
parasitoid numbers while the puparia of 
Calliphora vicina and Potamia littoralis had 
on average significantly fewer (Table 2). 
Puparia of Protocalliphora spp. are larger, 
which indicates a correlation between host 
size and number of individuals in the field, 
although it appears nonlinear: The C. vicina 
puparia are often smaller than the Proto- 
calliphora puparia (Table 3), but the size 
difference is not as big as the difference in 
parasitoid numbers. Furthermore, C. vicina 
puparia are larger than the P. littoralis 
puparia (Table 3), but the number of 
parasitoids does not differ. Taken together, 
C. vicina individuals seem to be a less 
suitable host, independent of host size. The 
reasons for this are unclear. One possible 
influential factor is that puparia of C. vicina 
show high variation of puparia size (Ta- 
ble 3), explained by limited food resources 
(carrion) during the lar\'al development, 
which results in small specimens and 


Journal of Hymenoptera Research 

maybe lower host quality while the para- 
sitic Protocalliphora spp. and the small P. 
littoralis have distinctly less variation of 
puparia size (Table 3). 

The correlation between host size and 
number of parasitoids was also recorded 
by Wylie (1967) and Rivers and Denlinger 
(1995) under laboratory conditions. Conse- 
quently there is a preference of N. vitripen- 
nis for larger hosts if choice is possible, 
recorded by Wylie (1967) and corroborated 
in field studies, whereas the parasitism rate 
of the smaller P. littoralis increases if the 
larger calliphorids are missing from a 
birds' nest (Peters and Abraham 2004). 

Field records of parasitoid numbers of N. 
vitripennis in the literature correspond 
quite well with the average parasitoid 
number from this study (16.1 parasitoids 
per host) but are rather rare and restricted 
to few larger hosts. Gold and Dahlsten 
(1989) recorded between 15 and 20 para- 
sitoids from Protocalliphora spp., Draber- 
Monko (1995) collected P. azurea puparia 
from tree sparrow nests and recorded a 
mean number of 20 N. vitripennis, but the 
sample size was small. Marchiori (2005) 
collected puparia of Peckia chrysostoma 
(Sarcophagidae) and noted a mean number 
of 15.2 N. vitripennis specimens. Molbo 
and Parker (1996) did not differ between 
Protocalliphora and Calliphora hosts and 
recorded a mean of 26 N. vitripennis 
from both species in the field. The results 
of Schlein (1998, 2001) using Calliphora 
vomitoria as host species indicate a signifi- 
cant difference between field and labora- 
tory conditions: An average of 33 N. 
vitripennis specimens were reared from 
Calliphora vomitoria in the laboratory 
(Schlein 2001). In the field numbers were 
distinctly lower with a mean of 15.7 speci- 
mens from the same host species (sentinel 
puparia placed inside the nest boxes) 
(Schlein 1998). 

The results show that there are inter- 
specific differences between host species 
and indicate that a correlation of parasitoid 
numbers and host size is present under 

natural conditions, but taxonomically re- 
stricted exceptions exist. Identifying the 
factors that underlie the differences in 
parasitoid numbers per host needs further 
experiments with different host species 
and values like parasitoid individual body 
size, developmental time and mortality 
rate to be included. 

Some differences in proportion of males 
emerging from puparia were recorded 
between host species, but no consistent 
correlation was indicated between sex ratio 
and host size (Table 2 and 3). Host quality 
model predictions (larger hosts exhibit 
more parasitoid females) cannot be found 
in our field data. The field recorded 
proportion of males especially in the 
abundant C. vicina and P. azurea are higher 
than expected assuming host-quality con- 
trol. The sex ratio in the smallest host, P. 
littoralis, is more female-biased than ex- 
pected. Other factors than host size must 
be considered for sex ratio determination. 
Accordant with the predictions for a 
species that shows local mate competition 
(LMC), Werren (1984) recorded a higher 
proportion of males in previously para- 
sitized hosts (superparasitism) for N. vitri- 
pennis. A further factor affecting sex ratio, 
which is also related to LMC theory, is the 
number of ovipositing females in the host 
patch, with increasing proportion of males 
if more females are present (Chamov et al. 
1981; King and Skinner 1991; Grillenberger 
et al. 2008). However, Burton-Chellew et al. 
(2008) showed that offspring sex ratio in 
natural populations was not directly influ- 
enced by number of females in the patch 
but only by relative clutch size. Besides 
LMC theory there are other demonstrated 
influences on sex ratio in N. vitripennis: 
Asymmetric larval competition between 
sexes is shown to favour less female-biased 
sex ratios as under larval competition 
females are smaller when a higher propor- 
tion of females was competing within a 
host (Sykes et al. 2007). Smaller females 
then produce less offspring (Chamov and 
Skinner 1984; Sykes et al. 2007). Shuker et 

Volume 19, Number 1, 2010 


al. (2006) showed that not only females but 
also males can have an effect on sex ratio, 
although underlying reasons for this effect 
are still unclear. The effect might be due to 
differences in sperm quality between male 
strains or due to active male influence in 
order to increase female offspring, i.e. to 
increase fertilisation and therefore increase 
contribution to next generation. However, 
both influencial factors are considered 
weak when compared to LMC (Shuker et 
al. 2006; Sykes et al. 2007). Constrained 
females which can produce only males 
obviously have effect on sex ratio. In this 
study hosts from which only males 
emerged were excluded from comparisons, 
but in superparasitized hosts we could not 
control the effect. An indirect effect of 
constrained females was shown to be 
absent in N. vitripennis by King and 
D'Souza (2004): The presence of a con- 
strained female did not influence offspring 
sex ratios of non-constrained females. 
Abraham and Konig (1977) studied the 
influence of temperature during oviposi- 
tion on sex ratio in N. vitripennis. At lower 
temperatures less eggs are laid per host 
with a more female-biased sex ratio. They 
explain these results with differential mor- 
tality: less females and more of the smaller 
males are able to finish development the 
more larvae are developing within one 

Recently, Grillenberger et al. (2009) 
studied the influence of multiparasitism 
on sex ratio in N. vitripennis and the closely 
related N. giraulti. They showed that multi- 
parasitim in this case has no effect on sex 
ratio. In our studies we recorded no 
multiparasitism although multiparasitism 
with Pachycrepiodeus vindemmiae (Rondani, 
1875) is known to occur in the Central 
European study area (Peters 2007). An- 
other factor that might indeed influence 
sex ratio is host age which was recorded as 
influencial for the solitary Spalangia sp. 
(King 2000) and might also have an impact 
on sex ratio in the gregarious N. vitripennis. 
In this study the only known factor among 

all these reported influences is host species 
and size. Number of females, degree of 
superparasitism, differences between mat- 
ing males, oviposition temperature etc. are 

Other known data on the sex ratio of N. 
vitripennis show a tendency towards lower 
proportions of males than in this study 
(especially when compared to the overall 
proportion of males. Table 2) but also 
display a wide range depending on various 
factors. In Schlein's (2001) laboratory rear- 
ings of N. vitripennis on C. vomitoria, the 
proportion of males was 0.17. In the studies 
of Abraham and Konig (1977) on another 
calliphorid host, Phormia regina, the ratio 
ranges from 0.125 to 0.2 depending on the 
temperature. Comparable ratios were 
found by Rivers and Denlinger (1995) for 
three larger sarcophagid host species in the 
laboratory. The exception in their studies 
was a proportion of males of 0.41 within 
the smaller Musca domestica. Two studies 
on the sex ratio of N. vitripennis in natural 
populations and natural host species re- 
corded a mean proportion of males of 0.19 
in unidentified Calliphora sp. and Protocal- 
liphora sp. (Molbo and Parker 1996) or 
report a wide range dependant on wasp 
population size (Werren 1983; host species: 
various necrophagous Cyclorrhapha). The 
recent studies of Grillenberger et al. (2008) 
and Burton-Chellew et al. (2008) on natural 
populations of N. vitripennis were done 
with a mixture of bait hosts and unidenti- 
fied natural hosts and thus provide no 
additional information on the role of host 

As discussed above, factors affecting 
parasitoid sex ratios are complex and 
cannot be solved here. However, our 
results indicate that host species should 
be considered when studying the influ- 
ences on proportion of males in host 
patches and parasitoid populations. This 
study initially shows data of N. vitripennis 
under field conditions and in identified 
field hosts which have been very rarely 
studied before. 


Journal of Hymenoptera Research 


The author would like to thank Bemhard Misof 
(Hamburg) and Joshua Gibson (Tempe, USA) for 
helpful comments on the manuscript; Gavin Broad, 
Mark Shaw and two anonymous referees for review- 
ing the manuscript; and Rudolf Abraham (Hamburg) 
and Helmut and Frieder Klopfer (Bad Mergentheim) 
for providing birds' nest material. 


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Vol. 19(1), 2010, pp. 139-143 

Rhopalum nasale, a New Species from Australia 
(Hymenoptera: Crabronidae) 


California Academy of Sciences, 55 Music Concourse Drive, Golden Gate Park, San Francisco, 
California 94118, USA; 

Abstract. — The new species Rhopalum nasale from Southern Australia is mainly characterized 
by its unique clypeal process, the expanded occipital carina, and the pronotal and propodeal 

Of the 278 currently recognized species of 
Rhopalum, 98 occur in Austraha (Leclercq, 
1997). This nuniber, however, if far from 
complete. I have recently seen in the 
Australian National Insect Collection, Can- 
berra, a dozen species that cannot be 
identified using Leclercq's key and that 
are apparently undescribed. They are re- 
presented mostly by one or a few specimens 
each. One of them, represented by a series of 
10 females, is so unusual that I cannot stand 
the temptation of describing it. 

The following are the abbreviations used 
in the text: 

ANIC: Austrahan National Insect Col- 
lection, Canberra, A.C.T., Aus- 

CAS: California Academy of Scien- 

ces, San Francisco, California, 

Rhopalum nasale Pulawski, sp. n. 

Name derivation. — The Latin adjective 
nasalis (neuter: nasale) means ''referring to 
the nose", or ''with a big nose"; with 
reference to the clypeal process of the 

Diagnosis. — The new species can be 
easily recogiuzed by the imique, narrow, 
conspicuous process emerging from the 
upper part of the clypeus (Fig. Ic, d), and 

the conspicuously expanded dorsolaterally 
occipital carina (Fig. la, b). The unusual 
pronotum and the propodeal side (see 
description below) are also diagnostic. 

Description. — Head as seen from above 
transverse (Fig. la); vertex thick (distance 
between hindocellus and occipital carina 
twice as long as ocellocular distance), its 
lateral margins nearly parallel. MidoceUus 
slightly smaller than hindoceUus; distance 
between hindocelli 2.6 X distance between 
midocellus and hindocellus; ocellocular 
distance 1.4 X hindocellar width. Orbital 
fovea well defined. Interantennal area flat, 
without specialized structures; interanten- 
nal distance about 1.1 X antennal socket 
width; distance between antennal socket 
and orbit about 1.5 antermal socket width. 
Clypeus with median process emerging 
from its dorsal area (Fig. Ic, d); process 
narrow, asetose, slightly broadening to- 
ward apex, emarginate apically; area be- 
tween process and clypeal ventral margin 
concave, asetose, delimited dorsally by 
carina that extends from each side of 
process. Mandibular apex unidentate. Oc- 
cipital carina joining h3rpostomal carina, 
conspicuously enlarged subdorsally but 
not enlarged dorsally (Fig. la, b). Upper 
frons and postocellar area finely punctate, 
punctures less than one diameter apart. 
Ventral portion of gena (on each side of 
hypostomal carina) without tubercle, un- 


Journal of H^t^lexoptefla Rese.arch 

Fig. i. K':c}-:i:U^': nasale sp. n., 9: a - head in dorsal view (arrow indicates orbital fovea); b - head in oblique 
view (arrow indicates occipital carina); c - clypeus in frontal \iew (arrow indicates projection); d - chpeus in 
lateral oblique view (arrow indicates projection); e - pronotum dorsallv (arrow indicates sublateral carina); f - 
pronotum laterally (arrow indicates sublateral carina). 

sculptured except for a few sparse punc- 
tures; posterior part of gena (behind eye) 
minutely punctate, punctures almost con- 
tiguous. Pronotum with pair of sublateral 
carinae (Fig. le) that diverge both anterad 
and posterad, nearest to each other at top 
of collar; anterior portion of each carina 

convex dorsally, concave subdorsally, and 
irregularly convex ventrally; pronotal side 
with deep, narro\v sulcus anterad of lobe, 
with glabrous, unsculptured area dorsally 
that is delimited by carina anteriorly and 
dorsally, and by sulcus posteriorly 
(Fig. If). Scutum finely punctate, punctures 

VoLL-ME 19, Nl-mber 1, 2010 


Fig. 2. RJiopalum rmsale sp. n., 9: a - thorax and propodeum in lateral view (arrow indicates longitudinal carina 
on propodeal side); b - propodeal dorsum; c - tergmn I and base of tergum 11 in dorsal vies (arrow indicates 
reddish bro\\Ti basal spot on tergum U); d - tergum I laterally; e - pygidial plate in dorsal \ie%v (arrow indicates 
bifid apical process). 

averaging about one diameter apart except 
more than one diameter apart at center; 
interspaces unsculptured. Mesopleural 
punctures larger than those on scutum, 
averaging less than one diameter apart 
except vertical subdorsal area adjacent to 

metapleuron unsculptured (Fig. 2a); epi- 
sternal sulcus crenulate; prepectus 
rounded. Propodeal enclosure not delim- 
ited, microareolate, dull, longitudinally 
ridged basally, without median sulcus 
(Fig. 2b); propodeal side with longitudinal 


Journal of Hymenoptera Research 

Fig. 3. Collecting localities of Rhopalum nasale, sp. n. 

carina that joins metapleural flange ante- 
riorly and ends just in front of hindcoxal 
base (Fig. 2a); area beneath carina unsculp- 
tured, shiny, area above carina with traces 
of longitudinal ridges. Recurrent vein join- 
ing submarginal cell at about two thirds of 
its length. Submarginal cell obliquely trun- 
cate. Forebasitarsus with two preapical 
rake spines that are about as long as 
basitarsus width. Hindtibia and basitarsi 
not swollen, hindtibia with well-defined 
spines. Tergum I relatively short, con- 
stricted apically (Fig. 2c), distance between 
basal tendon and imaginary line connect- 
ing spiracles half that of distance between 
line and apex; posterior half convex in 
profile (Fig. 2d); tergum length about 1.2 X 
maximum width, its maximum width 0.4- 
0.5 X that of tergum II. Tergum II with pair 
of dark reddish brown, asetose, elongate 
spots basally (Fig. 2c). Pygidial plate broad 
basally, gradually narrowing posterad, 
apically elongate as narrow, bifid process; 
with well-defined, sparse punctures ba- 
sally, and conspicuous setae next to pro- 
cess (Fig. 2e). 

Head, thorax, propodeum, and gaster 
black except the following: scape pale 
yellow, flagellum brown dorsally, light 
brown ventrally, mandible yellowish 
brown, dark apically, pronotal lobe pale 
yellow, tegula pale yellow (partly translu- 
cent), humeral plate dark brown mesally, 
pale yellow along margin. Femora black, 
forefemur pale yellow apically, largely 
yellowish brown in some specimens. Fore- 
tibia yellow; midtibia brown or yellowish 
brown, pale yellow basally and apically; 
hindtibia brown, pale yellow basally and 
apically. Tarsi yellow. 

Male. — Unknown. 

Geographic distribution (Fig, 3). — South- 
eastern South Australia. 

Records.— Holotyfe: 9, AUSTRALIA: 
South Australia: 32 km N Renmark at 
33"53'S 140°44'E, 2-29 Mar 1995, K.R. 
Pullen (ANIC). Paratypes (all collected 
by K.R. Pullen): AUSTRALIA: South 
Australia: 32 km N Renmark at 33°53'S 
140^44'E, 2-29 Mar 1995 (2 9, ANIC; 1 9, 
CAS); 31 km NW Renmark at 33°59'S 
140 30'E, 1-30 Mar 1995 (1 9, ANIC; 1 9, 

Volume 19, Number 1, 2010 


CAS) and 30 Mar - 2 May 1995 (1 9, ANIC; 
2 9, CAS); 14 km WNW Renmark at 34=07'S 
140°37, 28 Feb - 28 Mar 1995, ( 1 9, ANIC). 


generated the illustrations using Auto-Montage soft- 
ware package by Syncroscopy, and Lindsay Irving 
produced the distribution map. 


I sincerely thank Robert L. Zuparko for his critical 
comments on the manuscript. Helen K. Court con- 
firmed that the species is undescribed. Erin Prado 

Leclercq, J. 1997. Hymenopteres Sphecides d'Australie 
du genre Rhopalum Stephens, 1829. Notes Fauni- 
ques de Gembloux 32: 3-101. 

Vol. 19(1), 2010, pp. 144-158 

Colony Social Organisation of Halictus confusus in Southern Ontario, 
with Comments on Sociality in the Subgenus H. (Seladonia) 

M. H. Richards, J. L. Vickruck, and S. M. Rehan 
Department of Biological Sciences, Brock University, St. Catharines, Ontario 

Abstract. — Halictus (Seladonia) confusus Smith is one of the most common bees in North America. 
Classified as eusocial, its colony social organization is known only from qualitative descriptions of a 
population in Indiana. We studied the phenology and social behaviour of this bee in the Niagara 
Region of southern Ontario, using nest excavations, dissections and n\easurements of adult females, 
and pan trap samples of foraging bees to elucidate key elements of colony social organisation. The 
colony cycle in Niagara is typical of temperate-zone halictines, with overwintered foundresses 
producing a first brood of worker-sized females and a few males, followed by production of Brood 
2, consisting of gynes and more males. Many Brood 1 females become reproductive: about one- 
quarter of Brood 1 females dissected exhibited levels of ovarian development rivalling queens. In 
contrast, only about one-quarter of Brood 1 females become classically altruistic, sterile workers. 
High rates of worker reproductivity may result from early queen mortality and supersedure or 
from the inability of viable queens to control worker behaviour - the average queen-worker size 
difference was only 5.6%, and queens were not always larger than the workers in their own nests. 
Comparisons with the Indiana population suggest a geographic component to variation in colony 
social organisation. Comparisons with other members of the subgenus for which detailed 
information is available, suggest that in Seladonia, as in other eusocial halictines, queen control of 
worker behaviour depends on the ability of queens to dominate small numbers of small-bodied 

Key words. — Halictidae, eusociality, pan traps, sweat bee 

In halictine bees, evolutionary transi- understood, this in turn may help to 

tions from solitary to eusocial behaviour illuminate patterns observed at higher 

involve two components, a demographic taxonomic levels, such as differences 

change from univoltine to multivoltine among subgenera or genera. For instance, 

colony phenology, and a behavioural socially polymorphic sw^eat bees such as 

change from maternal care by a lone Lasioglossum calceatum and Halictus rubi- 

foundress, to associations between mothers cundus, exhibit solitary, univoltine colony 

and daughters that raise brood coopera- cycles in regions w\i\\ short breeding 

tively (Schw^arz et al. 2007). Likev^ise, seasons, and eusocial, bivoltine (actually, 

evolutionary transitions from eusociality double-brooded) colony cycles in regions 

to solitary behaviour, involve the reverse w^ith long breeding seasons (Sakagami and 

changes in demography and behaviour. Munakata 1972; Eickw^ort et al. 1996). There 

Therefore, to understand evolutionary are also obHgately eusocial species, such as 

transitions between solitary and social H. ligatus and L. malachurum that exhibit 

behaviour, it may be particularly fruitful considerable demographic variation, with 

to examine species that exhibit intraspecific colonies growing to larger sizes in areas 

variability in either or both of these traits. If with longer breeding seasons (Michener 

the adaptive significance of intraspecific and Bennett 1977; Knerer 1992). These 

demographic and social variability can be intraspecific patterns suggest that one 

Volume 19, Number 1, 2010 


cause of the phylogenetic lability of social 
behaviour observed in several halictine 
genera, might be geographic or temporal 
variability in the harshness of local envi- 
ronmental conditions. Indeed, this predic- 
tion is borne out by recent evidence that 
halictine sociality may have first evolved 
during a period of global climate warming 
(Brady et al. 2006), when it would have 
been possible for univoltine halictine 
lineages to adopt bivoltine or multivoltine 
nesting phenologies. 

One of the most common eusocial 
halictines in North America is Halictus 
(Seladonia) confusus Smith, but detailed 
information on its nesting and social 
biology are distinctly lacking. Dolphin 
(1966) studied the nesting biology and 
social behaviour of this bee in Indiana, 
USA, from 1963-1965. Although many 
crucial details were never published. Dol- 
phin suggested that H. confusus was demo- 
graphically and socially polymorphic. His 
study population contained nests that 
produced one, two, or three broods, com- 
prising both solitary and eusocial colonies. 
Eickwort et al. (1996) commented that H. 
confusus, presumed by Knerer and Atwood 
(1962) to be solitary in boreal Ontario, is 
social in New York. These tantalizing 
descriptions suggest that H. confusus may 
exhibit considerable demographic and so- 
cial variability within and between popula- 
tions. Understanding the ecological factors 
associated with such variation is key to 
investigating h3^otheses about the origins 
and extinctions of sociality in bees. 

In this paper, we describe the colony 
phenology and social organisation of H. 
confusus in southern Ontario. We studied a 
mixed nesting aggregation of halictine 
bees, including a small number of nests of 
H. confusus. We also used pan traps to 
collect adult females and males throughout 
the breeding season, in order to supple- 
ment the information from colony excava- 
tions. We show that while H. confusus is 
predominantly eusocial in southern On- 
tario, there is evidence that large numbers 

of Brood 1 females become reproductives, 
rather than sterile workers, suggesting that 
the population contains a mix of solitary 
and social strategies, as well as univoltine 
and bivoltine phenologies. We also com- 
pare H. confusus to other well studied 
members of the subgenus Seladonia, in 
order to assess the level of social variation 
in the subgenus as a whole. 


Study sites. — All study sites were on or 
within walking distance of the Brock 
University Campus in St. Catharines, On- 
tario (W 79 14' 57" N 43 07'11"). We 
excavated nests from a small nesting 
aggregation on the north shore of Lake 
Moodie that contained nests of Halictus 
confusus and H. ligatus, and hibemacula of 
H. rubicundus. The nests were on a gentle, 
south-facing slope. Nests were excavated 
using a standard technique in which baby 
powder was blown in at the nest entrances 
to coat the sides of the burrows, which 
were then carefully exposed using a 
kitchen knife. Nests were excavated in the 
morning before the entrances were open in 
or in the late afternoon after they were 
closed. All adult occupants were preserved 
in 95% ethanol, while brood were placed in 
wax-lined petri dishes indented with small 
chambers and brought back to the lab to be 
raised to adulthood. When these died or 
emerged as adults, they also were pre- 
served in ethanol. 

In addition to nest excavations, we used 
pan traps to capture flying bees at six sites 
on the Brock University campus and at the 
contiguous Glenridge Quarry Naturaliza- 
tion Site; pan trap sites were within 2 km of 
the nesting aggregation. At each site, 30 
pan traps were laid out in an X or other 
space-filling pattern, alternating yellow, 
white, and blue pans at 10m intervals, 
according to standard protocols (Lebuhn et 
al. 2003). Pans were set out weekly from 1 
May to 30 September 2006 at six locations. 
Bees caught in pan traps were used to 
determine the timing of important pheno- 


Journal of Hymenoptera Research 

logical events, including nest founding, the 
first and second brood-provisioning 
phases, and brood emergence from the 
nests. Since trapping effort was constant 
over the course of the summer, the num- 
bers of bees caught per week should 
provide a consistent estimate of bee den- 
sity and flight activity. Weeks were num- 
bered starting with 1 May 2006 as the 
beginning of week 1. 

Dissections. — Adult bees were measured, 
assessed for wear, and females were dis- 
sected. Body size was measured in terms of 
head width (HW, the distance across the 
widest part of the head, including the 
compound eyes) and length of the fore- 
wing costal vein (CVL, from the stigma to 
the end of the marginal cell); the head 
widths of pupae were also measured. 
Queen-worker size difference was calcu- 
lated as (queen HW - worker HW) / (queen 
HW) * 100. Mandibular wear (MW) was 
assessed on a scale of 0-5, with represent- 
ing completely unworn mandibles with 
sharp teeth and 5 representing mandibles 
so worn as to be completely blunted. Wing 
wear (WW) was also assessed on a scale of 
0-5, representing wings with no damage 
to the margin and 5 representing wings 
with the margin completely obliterated by 
nicks and tears. A total wear (TW) score was 
obtained by summing mandibular and 
wing wear scores for each female. As wings 
can be nicked during handling and because 
unworn mandibles sometimes appear 
somewhat blunt, bees were categorized as 
worn if TW>2. 

Females were dissected to determine 
mating status (whether the spermatheca 
was opaque, indicating that it was filled 
with sperm, or transparent, indicating that 
it was empty) and ovarian development. 
For the latter, all developing oocytes were 
assigned fractional scores of %, Vi, Va, or 1, 
indicating their size relative to a fully 
developed oocyte. These scores were then 
summed to make a total ovarian develop- 
ment (OD) score. Females with undeve- 
loped or only thickened ovaries but no 

visible oocytes, were assigned OD scores of 
or 0.1, respectively. 

Caste assignments for females were 
based primarily on seasonal activity pat- 
terns and secondarily on body size, based 
on the assumption that in Niagara, Halictus 
confusus would exhibit the bivoltine phe- 
nology typical of primitively eusocial ha- 
lictines in the temperate zone (Schwarz et 
al. 2007). The term 'foundress' is used for 
overwintered females that excavate bur- 
rows and forage in spring. The term 
'worker' is used for Brood 1 females. After 
workers emerge, a foundress may be 
referred to as a 'queen'. A gyne is a Brood 
2 female that will overwinter and found a 
nest the following spring. A 'replacement 
queen' is a Brood 1 worker that takes over 
the role of queen from a dead or moribund 

Caste designations were assigned to 
females caught in pan traps and nests 
based on the following criteria. When 
newly emerged from hibernation, foun- 
dresses are unworn, becoming progres- 
sively more worn as they excavate nests 
and provision brood cells. Thus in mid- 
summer, we can use wear scores to 
distinguish worn, late-foraging foun- 
dresses from unworn, early workers. In 
late summer, we used wear scores to 
distinguish worn workers from unworn 
gynes. Ovarian development was not used 
to assign caste designations, thus avoiding 
teleological complications in comparisons 
of the reproductive status of queens and 
workers. All adult females caught in nest 
excavations, as well as the majority of pan- 
trapped females (all foundresses, all gynes, 
and 100 workers) were measured and 

Interspecific comparisons. — To examine in- 
terspecific variation in the subgenus Sela- 
donia, the best approach would be to map 
these data onto a phylogeny and then 
investigate evolutionary correlations 
among the various traits (Felsenstein 
1988). However, in the absence of a 
phylogeny, several authors have used 

VOLL-ME 19, Nl-mber I, 2010 


principal components analysis (PCA) to 
quantify social variation among halictine 
bees and to construct hypotheses about 
how social traits co-evolve (Michener 1974; 
Breed 1976; Packer and Knerer 1985). 
Hypotheses constructed without a phylo- 
genetic framework, can then be tested 
when an appropriate phylogeny becomes 
available. For comparisons among Selado- 
nia populations, we used five variables 
commonly assessed in studies of halictine 
sociality: the proportion of males in Brood 
1, the number of workers per nest (or the 
number of females produced in Brood 1), 
the proportion of workers with developing 
ovaries, the proportion of mated workers, 
and the queen-worker size difference 
based on head width. Values for each 
variable were either taken directly from 
the literature, recalculated from figures in 
the literature, or recalculated as midpoints 
of ranges. The initial PCA was based on all 
five variables, retaining factors with eigen- 
values > 1.0. However, since Kaiser's 
Measure of Sampling Adequacy (MSA) 
with aU five variables had a value of only 
0.55, the variable with the lowest commim- 
ality measure (proportion of workers ma- 
ted) was dropped from the PCA. With the 
remaining four variables, MSA =0.77, 
which exceeds the 0.6 criterion. We present 
both factor loading scores (the degree to 
which each variable influences the inferred 
factors) and communality estimates (a 
reliability score which estimates the pro- 
portion of variance in each variable that 
is jointly explained by aU three factors). 
Note that the interspecific comparisons 
based on the PCA are presented in the last 
section of the Discussion, rather than in the 


Colony cycle. — In southern Ontario, H. 
confusus exhibits a foraging and nesting 
cycle t3rpical of temperate zone, eusocial 
halictines (Fig. 1). The beginning of the 
foundress foraging period was marked by 
the capture on 1 May 2006 of an over- 

wintered foundress. Since only two foun- 
dresses were captured in the first 3 weeks, 
they likely emerged from overwintering 
diapause in late April and early May, but 
mostly did not venture out of their nests 
until mid-May when the weather became 
more suitable. Foundresses continued to be 
caught in pan traps for about eight weeks, 
with the last foraging foundress caught on 
28 June (week 9). Most foundress foraging 
and provisioning of Brood 1 probably 
occurred from weeks 4-8. 

There was a sharp increase in the 
number of females caught beginning in 
week 8, many of them small and unworn. 
Large numbers of brood continued to be 
caught until week 11 after which pan trap 
catches declined. Weeks 8-11 thus repre- 
sented the peak emergence period of Brood 
1 and the peak worker foraging period. In 
the population as a whole, there was no 
quiescent period between the foundress 
and worker foraging periods, as the first 
Brood 1 females (which were small and 
unworn) were caught on 21 June (week 8) 
when clearly identifiable foundresses 
(large, worn females) were still flying. 
The first Brood 1 males were caught in 
week 9, so emergence of Brood 1 was 
slightly protog}^nous. Based on pan trap 
samples from weeks 8-11, the proportion 
of males in Brood 1 was about 1.9%. 

The emergence of Brood 2 was marked 
by a small increase in trap numbers of both 
males and females beginning around week 
15 (7-13 August), with the majority of 
Brood 2 emerging between weeks 18-20 
(Fig. 1). Week 15 was marked not only by 
the appearance of large, unworn females 
from Brood 2, but also by the last capture 
of smaU, unworn females deemed to be 
from Brood 1, suggesting that the last of 
Brood 1 had emerged as adults by week 15. 
The worker foraging period was mostly 
finished by week 17, although one small, 
worn forager was captured in week 19. 
Based on pan trap samples from weeks 12- 
20, the proportion of males produced in 
Brood 2 was about 22%. 


Journal of Hymexoptera Research 

D Females 
■ Males 



10 11 12 13 14 15 16 17 18 19 20 


■ Foundresses 
D Workers 
D Gynes 

2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 


Fig. 1. Right phenolog)' of H. confiisus based on 2006 pan trap samples. Top: Phenolog}' of all adult bees 
collected in pan traps. The sharp rise in numbers of females caught in mid-summer (beginning with week 8) 
corresponds with the first appearance of males in week 9, suggesting that this mid-summer peak marks the 
emergence of Brood 1. Bottom: Timing of female caste emergence based on size and wear patterns. Foundresses 
emerge in early May and continue to forage until mid-summer, slightly overlapping with females of Brood 1 
(workers). Gynes first begin to appear in week 15. Sample size differences between top and bottom graphs are 
because only 100 of the workers caught in pan traps was dissected. 

Nest contents. — Fourteen nests were ex- 
cavated in total, four prior to worker 
foraging and ten later in the summer. A 
single nest excavated during week 5 con- 
tained a foundress and three brood cells, 
comprising one provision mass with an 
egg, one medium larva, and one early stage 

pupa (damaged during excavation). In 
week 8, three nests were excavated. The 
first nest contained a queen and 3 worker 
pupae; the second nest contained a queen, 
one worker with worn mandibles, three 
worker pupae, and an unfinished provi- 
sion mass; and the third nest contained a 

Volume 19, Number 1, 2010 



4.4 J 

E 4.2 

^ 4 


A A 

O □ A 

A A 

■ □ 

A Q 
O A 
A A 

A A 


A Q 

D D 

□ O A 

A A 

■ □ 

n D 

A O 

O D 



■ Foundresses 
A Brood 1 females 



A A 









Head width (mm) 

Fig. 2. Body size distributions of pan trapped H. confusus foundresses. Brood 1 females, and gynes based on 
head width and wing length (measured as costal vein length, CVL). 

queen, two workers with worn mandibles, 
five female pupae, four male pupae, one 
larva that had completely consumed its 
provisions, one provision mass with an 
egg, and one unfinished provision mass. 
The average brood size of these three nests 
was 6.3, and 22% of the sexable brood were 
males. The latter figure is considerably 
higher than the estimate of 2% males based 
on pan traps and implies that males are 
under-represented in pan trap samples. 
The presence of workers in these nests, 
together with evident age gaps between 
younger (eggs and larvae) and older brood 
(pupae), indicates that the younger brood 
were from Brood 2 and that within 
individual nests there is a hiatus or 
quiescent period between Broods 1 and 2. 
Ten nests were excavated during weeks 
17 and 18. None contained a live foundress 
(queen). Four nests contained a total of six 
worn adult workers. Dissections showed 
that three of these had undeveloped 
ovaries but were mated (the other three 

were poorly preserved and could not be 
scored). The youngest brood were pigmen- 
ted pupae, so it is likely that older adult 
brood had already dispersed from their 
natal nests. The average number of brood 
per nest had fallen to 3.4, and only 3% of 
sexable brood were males (as compared 
with 22% in pan trap samples). Of the 23 
gynes in these nests, 21 had mated and 20 
had noticeable fat deposits in their abdo- 
mens. There was no evidence that gynes 
had begun digging hibemacula below their 
natal nests. 

Female body size. — Foundresses and 
gynes were very similar in size, and both 
were larger than Brood 1 females; there 
was no indication of a body shape differ- 
ence between the gyne and worker castes 
(ANOVA, F=8.56, df=2,147, p=0.0003; 
Fig. 2 and Table 1). Queen and worker size 
measurements were available for two nests 
(both excavated in week 8). In Nest 166, the 
queen was larger than all four of her 
workers (adults and pupae), and the 


Journal of Hymenoptera Research 

Table 1. Caste characteristics of females caught in pan traps. Smaller sample sizes for ovarian and mating 
success reflect technical difficulties with dissections. Females were considered as worn if MW>2 or WW ^2, 
and ready to lay if they contained at least one, y4-developed oocyte. Statistical comparisons of foundresses 
versus Brood 1 females were based on ANOVA (F statistics), Kruskal-Wallis tests (H statistics), and chi 
square tests. 


Brood 1 females 


Statistical comparison 



(n = 100) 


(foundresses vs. Brood 1 females) 

HW (mm ± 1 sd) 

2.09 ± 0.07 

2.03 ± 0.08 

2.09 ± 0.09 

F=11.22, df=l, p=0.0011 

CVL (mm ± 1 sd) 

4.33 ± 0.16 

4.15 ± 0.16 

4.25 ± 0.17 

F=21.91, df=l, p=0.0001 

Proportion with worn 

13/22 (60%) 

32/100 (32%) 

0/28 (0%) 

X'=5.68, df=l, p<0.0171 


Proportion with worn wings 

1/22 (4%) 

8/100 (8%) 

0/28 (0%) 

X'=0.32, df=l, n.s. 

OD score (mean and range) 

1.82 (0.5-2.75) 

0.58 (0-2.25) 

0.03 (0-0.1) 

F=21.64, df=l, p=0.0001 

Proportion ready to lay 

16/22 (73%) 

23/100 (23%) 

0/28 (0%) 

X'=20.50, df=l, p<0.0001 

Proportion mated 

18/18 (100%) 

39/80 (49%) 

17/28 (61%) 


queen-worker size difference was 7.2% 
based on head width and 9.9% based on 
wing length. In Nest 168 the situation was 
very different. The small, worn queen was 
the same size as one worker, but smaller 
than four others (two worker pupae were 
not measured), resulting in a queen-worker 
size difference of negative 4.0% based on 
head width and negative 1.2% based on 
wing length. Since the above calculations 
were based on females from only two 
nests, we also calculated the average size 
differences for pan trapped foundresses 
and workers: these were 2.9% based on 
head width and 4.2% based on wing 

Wear and reproductive status. — Based on 
females caught in pan traps, foundresses 
sustained higher levels of mandibular wear 
than Brood 1 females (Table 1). Few fe- 
males had worn wings, but one notable 
exception was the queen of Nest 168 
(excavated in week 8), with a total wear 
score of 10; this female was so much more 
worn than other bees examined that she 
might have been nesting for the second 
time, having overwintered twice. 

Potential for reproduction by foundresses 
and Brood 1 females is compared in Table 1. 
All foundresses dissected (18 from pan 
traps and 4 from nest excavations) had 
sperm in their spermathecae, whereas only 
about half of the Brood 1 females examined 

had sperm in their spermathecae. Foun- 
dresses also had significantly higher OD 
scores than Brood 1 females, and were more 
likely to have at least one oocyte ready or 
almost ready to lay. 

Four types of 'workers' could be distin- 
guished based on wear and ovarian devel- 
opment, each category comprising about 
25% of the total among Brood 1 females 
caught in pan traps (Table 2). The first 
group comprised unworn (TW < 1) 
females with undeveloped ovaries (OD < 
0.1); these were evidently newly eclosed 
workers. The second group were worn 
(TW>2) but exhibited no ovarian develop- 
ment, suggesting that they were engaged 
in nest maintenance or foraging activities, 
but were not laying eggs; these bees were 
categorized as sterile altruists. The third 
group were queen-like, at least in terms of 
their readiness to lay eggs: most of these 
(18.3% of all Brood 1 females) contained at 
least one fully developed oocyte ready to 
lay, while the remainder contained at least 
one y4 -developed oocyte. The remaining 
group of Brood 1 females can be categor- 
ized as potentially reproductive workers, 
exhibiting a distinct degree of wear and 
some ovarian development, but not suffi- 
cient to be ready to lay eggs. These workers 
likely provision both queen-laid and some- 
times their own eggs, and could also be 
referred to as 'partial altruists'. 

Volume 19, Number 1, 2010 


Table 2. Comparison of ovarian development and wear in Brood 1 females collected in pan traps. Unworn 
females had total wear ( I TW = MW + WW) scores of or 1, whereas worn females had TW > 2. Percentages 
represent proportions of the total (n=93). Four categories of workers can be distinguished: 'newly eclosed' 
females that have not yet accumulated either wear or ovarian development; 'altruists', worn, working females 
with no ovarian development, 'queen-like' females with very high rates of ovarian development, and the 
remainder, with intermediate levels of wear and ovarian development, that can be referred to simply 
as 'workers'. 

Size of largest oocyte 





24 (25.8%) 

21 (22.6%) 

45 (48.4%) 

Newly eclosed 



6 (6.5%) 

10 (10.8%) 

16 (17.2%) 

Potentially reproductive 

Potentially reproductive 




2 (2.2%) 

7 (7.5%) 

9 (9.7%) 

Potentially reproductive 

Potentially reproductive 




2 (2.2%) 

4 (4.3%) 

6 (6.5%) 




6 (6.5%) 

11 (11.8%) 

17 (18.3%) 




40 (43.0%) 

53 (57.0%) 

93 (100%) 

Roughly half of Brood 1 females caught 
in pan traps were mated (this value under- 
estimates the rate of worker mating as it 
includes newly eclosed individuals that 
might not yet have met males). Those with 
developed ovaries (OD scores > 0.25) were 
more likely to have mated than females 
with no ovarian development (Likelihood 
ratio chi-square, G= 14.46, df=l, p<0.0001; 
Table 3), and this was significant even 
when newly eclosed females are excluded 
from consideration (Likelihood ratio chi- 
square, G=4.978, df=l, p=<0.0257). De- 
gree of ovarian development was not 
correlated with body size (head width: 
p=— 0.01, n=95, n.s.; costal vein length: 

Table 3. Association between mating status and 
ovarian development in 80 H. confusus Brood 1 
females collected from pan traps. The minimum OD 
score for a female with at least one visibly developed 
oocyte is 0.25. Statistical analysis is given in the text. 

Mating status 

Ovarian score 

Unmated Mated 


OD < 0.1 
OD > 0.25 

29 (36%) 11 (14%) 
12 (15%) 28 (35%) 
41 (51%) 39 (49%) 

40 (50%) 
40 (50%) 
80 (100%) 

p=0.02, n=95, n.s.), even when females 
assumed to be newly eclosed were ex- 

Comparisons of queens to the workers in 
their own nests suggest that queens domi- 
nated but did not completely monopolize 
oviposition. Nest 166 was excavated on 22 
June 2006 (week 8), and contained a queen, 
one adult worker, three worker pupae, and 
an unfinished provision mass. The queen 
(TW=6) had an OD score of 2.75, including 
three y4 -developed oocytes but no fully 
developed oocytes. The adult worker was 
slightly worn (TW=3) and had probably 
collected the pollen provisions. She was 
mated and her OD score was 0.75, com- 
prising a single y4 -developed oocyte. Evi- 
dently, the queen or the worker could have 
had a mature oocyte to lay by the time the 
provision mass was completed. In nest 168, 
excavated on the same day, the queen, 
which was the most worn bee we found 
(TW=10) had an OD score of 1.75, com- 
prising one fully developed and one y4 
developed oocyte. Of the two adult work- 
ers in the nest, the one smaller than the 
queen was worn (TW=3), was unmated 
and had only slightly thickened ovaries. 


Journal of Hymenoptera Research 

while the one larger than the queen was a 
bit less worn (TW=2), was mated and had 
an OD score of 1.0, including a y4 -devel- 
oped oocyte. Since both workers were 
worn, they were probably both foragers, 
but only the former would be categorized 
as a 'sterile altruist'. 

Dissections of pan-trapped bees revealed 
that 4/22 foundresses, 14/100 workers, 
and 0/28 gynes had been parasitized by 
conopid larvae, many of them large en- 
ough to fill their host's abdominal cavity. 
Perhaps noteworthy is the fact that two 
foundresses caught in pan traps in late 
May contained conopid parasites so large 
as to prevent any ovarian development. 
Two gynes from nests excavated in late 
August were also parasitized by conopids. 


Phenology and colony social organization in 
southern Ontario. — In southern Ontario, 
Halictus confusus exhibits a colony cycle 
which in broad terms, is typical of eusocial, 
temperate zone halictines (Schwarz et al. 
2007). In spring, large females excavate 
new burrows, then provision a first brood 
that is composed mainly of workers and a 
few males. Foundresses cease provisioning 
shortly after the summer solstice, and then 
are replaced as small Brood 1 females 
emerge from their nests, and begin to 
provision Brood 2. Most individuals of 
Brood 2 are provisioned by the end of July, 
emerging as adults until mid-September. 
Since many queens evidently survive until 
mid-summer when workers emerge and 
begin foraging, this suggests that many 
surviving colonies become eusocial. As in 
other halictine bees (Packer 1992; Richards 
et al. 1995; Paxton et al. 2003; Richards et al. 
2005), foundress queens likely produce the 
majority of Brood 2 gynes and males. 
Dissections indicate that queens have 
higher reproductive potential on average, 
and that workers can have high rates of 
ovarian development even in queen-right 
nests. The relatively large numbers of 
unworn workers with highly developed 

ovaries caught in midsummer, suggest that 
when foundress queens die or become 
moribund, they are replaced by one of 
their Brood 1 daughters, and colonies 
become parasocial. 

Halictus confusus nests are probably 
founded haplometrotically (singly), as the 
few (n=4) nests that we excavated in 
spring each contained a single foundress. 
Haplometrotic nest founding is more likely 
when gynes overwinter away from the 
summer nesting sites, while pleometrotic 
co-founding is more likely when gynes 
overwinter together near the nesting site 
(Packer 1993; Richards and Packer 1998). 
Atwood (1933) and Dolphin (1966) sug- 
gested that H. confusus gynes overwinter 
away from their natal nests, and nests that 
we excavated near the end of August 
contained newly eclosed gynes but no 
evidence that these were preparing hiber- 
nacula. Nevertheless, pleometrosis cannot 
be ruled out entirely, as we did excavate a 
nest in which the queen was smaller than 
most of her workers. In eusocial halictines, 
queens control worker body size by ma- 
nipulating the size of larval provision 
masses (Richards and Packer 1994), mak- 
ing workers that are almost always smaller 
than themselves (Richards and Packer 
1996), so the finding of a very worn queen 
smaller than some of her own workers 
suggests that she may have been a small 
subordinate co-foundress that outlived a 
larger dominant (Packer 1986; Richards 
and Packer 1996). 

In H. confusus, it appears that females 
produced in Brood 1 may adopt one of 
three or four different reproductive op- 
tions. Some Brood 1 females become 
classical, sterile, altruistic workers that 
provide provisions for eggs laid by the 
queen but produce no offspring of their 
own. Some Brood 1 females become re- 
productive workers, a category that com- 
prises workers that collect provisions upon 
which a queen will lay eggs, but whose 
developing ovaries suggest that they also 
will lay eggs given the chance. For many. 

Volume 19, Number 1, 2010 


perhaps most, of these 'reproductive work- 
ers', egg-laying opportunities may never 
present themselves, so observations that 
many workers have ovarian development 
do not necessarily translate into high rates 
of worker oviposition in queen-right nests 
(Packer 1992; Packer and Owen 1994). 
Nevertheless, worker maternity in queen- 
right nests does occur even in strongly 
eusocial halictines (Richards et al. 2005) so 
in H. confusus, it is likely that at least some 
reproductive workers, successfully pro- 
duce brood, even in queen-right nests. 
The workers with queen-like ovaries 
would almost certainly be egg-layers, and 
most likely were replacement queens. We 
found no new H. confusus nests in mid- 
summer after the first emergence of work- 
ers, so it is unlikely that workers with 
queen-like ovaries were Brood 1 females 
that leave their natal nests to found new 
nests in summer, either solitarily or com- 
munally (Sakagami and Hayashida 1968; 
Richards et al. 2003). 

A curious feature of the flight phenology 
of H. confusus in Niagara was the small 
number of females captured in late sum- 
mer, following emergence of Brood 2, 
compared to the far greater numbers 
captured in midsummer following emer- 
gence of Brood 1. Several explanations 
present themselves. First, gynes might 
have been imder-represented in pan traps 
relative to workers, due to changes in 
flower and forage availability. Pan traps 
are known to capture relatively fewer 
foragers when flower availability increases 
(Roulston et al. 2007). Pan traps may 
therefore be less attractive to gynes (and 
males) because they are not active provi- 
sioners, and because flower availability 
may be higher after midsummer than 
before. It is also possible that the pattern 
of lower gyne than worker densities is real. 
If so, then one explanation would be high 
rates of colony failure prior to worker 
emergence (Richards and Packer 1995a). 
Another possibility is that some Brood 1 
females leave their natal nests to enter 

diapause preparatory to becoming foun- 
dresses the following spring, a phenom- 
enon known as differential diapause and 
well documented m Halictus rubicundus 
(Yanega 1988). It would be interesting to 
compare pan trap phenologies with de- 
tailed nesting data for several species with 
different colony cycles, in order to assess 
concordance in the patterns inferred using 
the two types of information. 

Geographic variation in colony social orga- 
nisation. — Demographic differences be- 
tween Indiana (Dolphin 1966) and Ontario 
likely stem from differences in the timing 
of important colony events. In Indiana, 
foundresses emerge from hibernation as 
early as March or April and complete 
foraging by late May or early June, with 
first brood workers emerging from mid- 
May to early June, second brood workers 
emerging in mid to late July, and gynes 
emerging from mid-July to early Septem- 
ber. In Ontario, foundresses emerge from 
hibernation in late April and forage until 
about the third week of June, with workers 
emerging from about June until the end of 
July, and gynes from mid-August to mid- 
September. This suggests that Dolphin's 
population experienced a breeding season 
about three weeks longer than we observed 
in Niagara in 2006. In Indiana, many 
colonies produced two worker broods. 
This seems unlikely for our Ontario popu- 
lation, as pan traps suggested that the 
majority of Brood 1 workers emerged 
between weeks 8 and 13, a six week period 
that is only slightly shorter than the period 
encompassing most foundress foraging 
activity between weeks 3 and 9. However, 
the intriguing, small peak in captures of 
females around weeks 14 and 15, might 
have signaled the emergence of a second- 
ary worker brood. We did capture some 
small, unworn females at this time, which 
we categorized as gynes, but which were 
possibly workers. The ability to interpolate 
a second worker brood in areas with long 
enough breeding is well known in Lasio- 
glossum malachurum, which produces a 


Journal of Hymenoptera Research 

single worker brood at the northerly edge 
of its range, but two or three worker 
broods in warmer environs and at the 
southerly extent of its range (Knerer 1992; 
Wyman and Richards 2003; Weissel et al. 
2006). Augochlorella striata apparently has 
sufficient behavioural flexibility that it can 
respond to annual weather conditions by 
producing workers when conditions will 
create a long breeding season or omitting 
workers and producing gynes directly 
when conditions will create a short breed- 
ing season (Packer 1990). 

Another phenological difference be- 
tween Ontario and Indiana H. confusus 
was the absence in Ontario of a distinct 
quiescent period between the foundress 
and worker foraging periods. Not only was 
there no quiescent period, but there was at 
least a week of overlap in the foraging 
periods of foundresses and workers. This 
was somewhat unexpected as a quiescent 
period between the two flight periods is 
typical of many primitively eusocial halic- 
tines, even when multiple worker broods 
are produced. The overlap suggests an 
extended rather than a synchronized per- 
iod of nest establishment in spring, with 
the result that some foundresses continued 
to provision brood as long as 2-3 weeks 
after the earliest foundresses had com- 
pleted their first broods. However, lack of 
synchronicity may not be typical of Nia- 
gara H. confusus, if weather conditions in 
the spring of 2006 led to early nesting 
activity by some foundresses. In H. ligatus 
nesting near Victoria, Ontario, unusually 
warm conditions in spring 1991 led to an 
extended nest founding period that ob- 
literated the usually predictable hiatus 
between the foundress and worker fora- 
ging periods (Richards and Packer 1995b). 

Sociality in the subgenus Seladonia. — Ta- 
ble 4 compares H. confusus to other species 
of the subgenus Seladonia for which suffi- 
ciently detailed sociobiological data are 
available. All members of the subgenus 
are thought to be primarily social (Packer 
et al 2007), including H. tumulorum which 

originally was thought to be solitary 
(Sakagami 1974). For three species, H. 
confusus, H. hesperus, and H. lucidipennis, 
social data are available for intraspecific 
comparisons between populations. Varia- 
tion between populations suggests consid- 
erable geographic variability in rates of 
worker mating and ovarian development. 
In H. confusus, there seems to be a link 
between breeding season length and col- 
ony size, as additional worker broods were 
interpolated before the gyne brood in 
Indiana. A similar pattern seems to occur 
in H. lucidipennis, which produces more 
workers in southern than in northern India 
(Batra 1966). 

In the absence of a phylogeny for 
behaviourally known members of the sub- 
genus Halictus (Seladonia), we used princi- 
pal components analysis (PCA) to quanti- 
tatively explore correlations among five 
sociobiologically important variables (Ta- 
ble 4). Only six populations (H. confusus 
from Ontario, H. aerarius, H. hesperus from 
Mexico, H. lucidipennis from northern and 
southern India, H. tumulorum from France, 
and H. vicinus from southern India) could 
be included as basic information was 
missing for the remainder. We did not 
attempt a hierarchical analysis to distin- 
guish between inter and intraspecific var- 
iation, although it is possible that intra- 
and interspecific patterns might differ (this 
could eventually be tested phylogeneti- 
cally). As noted in the methods, proportion 
of workers mated explained little of the 
variation among populations and was 
dropped from the analysis. Its lack of 
explanatory power could stem either from 
difficulties in data collection (Table 4 sug- 
gests considerable variability depending 
on when workers are captured for dissec- 
tion) or could reflect a genuine lack of 
relevance to explaining behavioural varia- 
tion among populations. The remaining 
four variables together explained 76% of 
variability among populations, ranked 
along a single eigenvector (principal com- 
ponent) with eigenvalue = 3.051. The 

Volume 19, Number 1, 2010 


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Journal of Hymenoptera Research 

Table 5. Results of principal components analysis 
for 7 populations of Seladonin {H. confusus from 
Ontario, H. aerarius, H. hesperus from Mexico, H. 
lucidipeimis from northern and southern India, H. 
tumulorum from France, and H. vicinus) compared in 
Table 4. Only the first principal component 
(eigenvalue = 3.051) was retained based on the 
criterion that eigenvalue > 1.000. 

Factor 1 



loading score 


Proportion of males in 



worker brood 

Number of workers per nest 



Proportion of workers with 



developed ovaries 

Queen-worker size 




results showed that about 76% of the 
variation among these populations and 
species was explained by this principal 
component (Table 5). The factor loading 
scores indicate strong negative correlations 
between queen size relative to workers 
versus number of workers produced in 
Brood 1 (which are usually provisioned by 
the queen herself), the proportion of males 
in this brood, and rates of worker ovarian 
development. Taken together, this suggests 
that in the subgenus Seladonia, that soci- 
ality, especially the degree of queen- 
worker reproductive skew, is related to 
the ability of queens to dominate workers. 
This is a well-known pattern in eusocial 
halictines, in which greater skew occurs 
when queens must contend with fewer 
and /or smaller workers (Schwarz et al. 

Two species not included in the PCA, H. 
hesperus and H. lutescens (Wille and Mich- 
ener 1971; Brooks and Roubik 1983; Saka- 
gami and Okazawa 1985), can develop 
extraordinarily large colony sizes number- 
ing in the many hundreds. It seems 
unlikely that such large colonies result 
from egg production by only one female 
(but see Plateaux-Quenu (1962) for an 
important exception), and more likely 
these colonies contain multiple egg-layers. 
This suggests that queens might dominate 

oviposition early in the colony cycle, but 
eventually are superseded or lose control 
of worker reproduction as colonies grow. 
This switch to 'worker' reproduction 
would allow colonies to grow even larger 
and would also blur the distinction be- 
tween queens and reproductive workers. 
Interestingly, queen supersedure was sug- 
gested by Dolphin (1966) for H. confusus in 
areas with colony cycles longer than 
average queen lifespan. Another Seladonia 
species not included in our comparative 
analysis due to a lack of nesting data, is H. 
lanei. In this species, the degree of queen- 
worker size dimorphism is extraordinary, 
with queens being as much as eight times 
larger than their workers (Janjic and Packer 
2001). This exceptional degree of size 
differentiation may allow queens to dom- 
inate oviposition and effectively control 
worker behaviour even with large colony 
sizes. Clearly, more behavioural data on 
more species in this interesting subgenus 
are required for us to better understand 
how queen control and reproductive skew 


Across Halictus (Seladonia), all species 
thus far studied exhibit sociality (Packer et 
al. 2007). However, the social behaviour of 
H. confusus suggests a high degree of 
intraspecific social variability. Moreover, 
specimens have been collected as far north 
as Alaska and Finland, where breeding 
seasons may be too short to allow foun- 
dresses to produce workers, so there is a 
high probability that at least some Seladonia 
populations or species are monomorphi- 
cally solitary. Social polymorphism, co- 
occurrence of solitary and social nests 
within populations, might represent an 
ecological intermediate between solitary 
populations in areas with very short 
breeding seasons and social populations 
in areas with longer seasons (Packer 1990). 
Behaviourally, variation in the strength of 
dominance hierarchies and in queen- 
worker reproductive skew may also be 

Volume 19, Number 1, 2010 


taken as evidence of intermediate stages in 
social evolution. The behavioural variability 
that we have observed in H. confusus and 
which others have observed in other mem- 
bers of the subgenus, make it likely that 
further study of H. (Seladonia) populations 
will shed considerable light on the ecologi- 
cal and evolutionary factors that promote 
social transitions in halictine bees. Species 
like H. confusus, with wide geographic 
ranges encompassing very short to rela- 
tively long breeding seasons, would be ideal 
candidates for studies of reproductive skew 
using microsatellite markers to specifically 
investigate the circumstances under which 
queens lose control of worker reproduction. 


We gratefully acknowledge the helpful critiques of 
two anonymous reviewers. This project was funded 
by an NSERC Discovery Grant to MHR, a Brock 
University Deans' Graduate Fellowship to JLV, and an 
NSERC Undergraduate Summer Research Award to 


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Vol. 19(1), 2010, pp. 159-178 

Ultrastructure of Scutellar Sensilla in Aphytis melinus (Hymenoptera: 
Aphelinidae) and Morphological Variation across Chalcidoidea 

Christina A. Romero and John M. Heraty* 
Department of Entomology, University of California, Riverside, CA 92521 

Abstract. — Paired, disc-like campaniform sensilla occur on the scutellum of many minute 
parasitic wasps in the superfamily Chalcidoidea (Hymenoptera). The ultrastructure of the sensilla 
is examined in Aphytis melinus DeBach (Aphelinidae). Each sensillum consists of a bilayered 
cuticular cap directly covering a tubular body with microtubules extending at a right angle to the 
cuticle. A large electron-dense mass attached to the tubular body extends laterally beneath the 
cuticle. Other structures occupying the space between the scutellum and longitudinal flight muscles 
include the paired mesoscutello-metanotal muscles and a previously undescribed layer of oblong 
structures lining the cuticle throughout the thorax. Among 23 additional species examined, the 
sensilla range in diameter from l.Sljim to 5.79 |Lim, with no apparent relationship between diameter 
of the sensilla and size of the scutellum. The function of the sensilla is unknown, but the consistent 
presence of the sensilla in small chalcidoids and the frequent absence in the largest species suggests 
a possible association with specialized flight peculiar to small insects obliged to utilize the clap-and- 
fling flight mechanism. 

Key words. — campaniform sensilla, morphology, sensory structures 

Chalcidoidea is a diverse superfamily of 
parasitic Hymenoptera whose species 
range in length from the smallest known 
insect (0.11 mm) to relatively large wasps 
(45 mm), with most specimens averaging 
2-4 mm in length (Heraty and Gates 2003). 
Over 22,000 species of Chalcidoidea are 
described, making it second to Ichneumo- 
noidea in diversity, but with an estimated 
100,000 to 400,000 undescribed species, it 
may well prove to be the largest super- 
family of Hymenoptera (Gibson et al. 1999; 
Gordh 1975a; Heraty and Gates 2003; 
Noyes 2000, 2003). Despite over 200 years 
of taxonomic work, phylogenetic relation- 
ships at the family and subfamily levels 
remain unclear (LaSalle et al. 1997). Diffi- 
culties in understanding chalcidoid phylo- 
genetics are due in part to the vast 
numbers of undescribed species and the 
poor preservation of many curated speci- 
mens (Heraty 2004; LaSalle 1993). Addi- 

"Corresponding author: 

tionally, with the vast majority of chalci- 
doids measuring less than 4 mm, there is 
often a paucity of reliable phylogenetically 
informative morphological structures. 

Sensillar structures have proven to be a 
rich source of morphological characters, 
and there have been numerous investiga- 
tions into the structure and function of 
sensilla found within Chalcidoidea (Baaren 
et al. 1996; Barlin and Vinson 1981; Olson 
and Andow 1993; Schmidt and Smith 1985, 
1987). The antenna has been the focal point 
of the majority of sensillar investigations in 
chalcidoids due to the high concentration 
and diversity of antennal sensilla (Barlin 
and Vinson 1981; Basibuyuk et al. 2000; 
Olson and Andow 1993; Walther 1983). 
These studies have focused largely on 
classifying types of antennal sensilla based 
on ultrastructural morphology (Amornsak 
et al. 1998; Baaren et al. 1996; Barlin and 
Vinson 1981; Basibuyuk and Quicke 1999; 
Consoh et al. 1999; Isidoro et al. 1996; 
Olson and Andow 1993). Other investiga- 


Journal of Hymenoptera Research 

Fig. 1. Location and appearance of scutellar sensilla in Aphytis melinus. a: whole specimen, b: scutellum, c: 
right scutellar sensillum. White boxes in (a) and (b) indicate area magnified in following figure. 

tions have examined sensory structures of 
the ovipositor (Consoli et al. 1999; Le Ralec 
et al. 1996; Le Ralec and Rabasse 1988; 
Veen and Wijk 1985), male genitalia 
(Chiappini and Mazzoni 2000) and wings 
(Schmidt and Smith 1985). The paired 
sensilla of the scutellum (Fig. la-c) have 
been identified as phylogenetically impor- 
tant (Hayat 1998; Heraty and Polaszek 
2000; Kim 2003; Schauff et al. 1996), but 
there has been no investigation into their 
ultrastructure, possible function or distri- 
bution across Chalcidoidea. 

The scutellar sensilla are a feature 
frequently overlooked in taxonomy and 
have received only sparse attention in the 
literature. Domenichini (1969) was one of 
the first morphologists working with Chal- 
cidoidea to point out the scutellar sensilla, 
noting their occurrence in several different 
families and recommending that their 
function and taxonomic value be studied. 
Rosen and DeBach (1979) also noted the 
sensilla in their treatise on Aphytis Howard 
(Aphelinidae), mentioning in each of their 
species descriptions the location of the 
scutellar sensilla relative to the anterior 
and posterior scutellar setae. They ob- 
served that, in slide preparations, the 
sensilla can be mistaken for empty setal 
sockets due to the thinness of the cuticle 
over the sensilla. They also noted rare 
mutations involving the sensilla in which 
one or both sensilla are replaced by an 

extra seta, by a pair of setae, or where there 
appears to be one, three, or four sensilla in 
place of the normal pair of sensilla. Both 
Schauff et al. (1996) and Hayat (1998) 
incorporated the placement of the sensilla 
in their keys of Encarsia Forster (Aphelini- 
dae). Heraty and Polaszek (2000) used the 
close placement of the sensilla on the 
scutellum as a defining characteristic of 
the Encarsia strenua group. Placement of the 
sensilla on the scutellum was also used by 
Schauff (1984) as a character in his phylo- 
geny of Mymaridae. Other allusions to the 
sensilla in the literature are limited to 
inclusion in illustrations and an occasional 
mention in species descriptions. 

Herein we demonstrate that these scu- 
tellar structures are campaniform sensilla, 
which are circular to oval in shape and 
innervated by just one sense cell, or 
neuron, that partially penetrates the thin- 
domed cuticle (Hicks 1857; Berlese 1909; 
Snodgrass 1935; Mclver 1985). Campani- 
form sensilla have a mechanoreceptive 
function targeted at sensing tension or 
torsion in the associated cuticle (Pringle 
1938a; Mclver 1985; Zill and Moran 1981). 

In chalcidoids, campaniform sensilla 
have been identified on the antenna 
(Amomsak et al. 1998; Olson and Andow 
1993), ovipositor (Consoli et al. 1999; Le 
Ralec et al. 1996; Le Ralec and Wajnberg 
1990), male genitalia (Chiappini and Maz- 
zoni 2000), wing (Schmidt and Smith 1985; 

Volume 19, Nl-mber 1, 2010 


Weis-Fogh 1973), pretarsus (Gladun and 
Gumovsky 2006) and legs (Schmidt and 
Smith 1987), but the internal ultrastructure 
of these sensiUa in the superfamily has 
been examdned only in male genitalia of 
Mymaridae (Chiappini and Mazzoni 2000). 

In the current study, Aphytis melimis 
DeBach (Hymenoptera: Aphelinidae: 
Aphehninae) was chosen to examine the 
ultrastructure of the scutellar sensilla. 
Aphytis melinus range in size from 
0.78 mm to 1.21 mm (Rosen and DeBach 
1979) and have been well studied because 
of their success as biological agents con- 
trolling the California Red Scale, Aonidiella 
aurantii (Maskell) (Lenteren 1994). Prior to 
this study, paired scutellar sensilla were 
recorded only in the smallest Chalcidoidea 
such as Aphelinidae (Hayat 1984, 1997; 
Huang 1994; Heraty and Polaszek 2000; 
Babcock et al. 2001; Kim 2003; Noyes and 
Valentine 1989; Schauff et al. 1996), En- 
cyrtidae (Hayat 2003; Noyes 1988; Noyes et 
al. 1997; Prinsloo 1997), Mymaridae 
(Schauff 1984), Signiphoridae (Noyes and 
Valentine 1989) and Trichogrammatidae 
(Doutt and Viggiani 1968; Noyes and 
Valentine 1989), with most attention being 
given to scutellar sensilla in Aphelinidae 
(Heraty and Polaszek 2000; Kim 2003; 
Rosen and DeBach 1979). 

No previous work has sought to examine 
the ultrastucture of the scutellar sensilla 
found in Chalcidoidea. This study seeks to 
survey variation in external appearance of 
the scuteUar sensilla found in Chalcidoi- 
dea, examine the ultrastucture of the 
sensilla in A. melinus, and accurately 
determine the category of sensilla to which 
they belong. 


Terminology. — Terms and abbreviations 
foUow Gibson (1997) and Kim (2003) for 
the structures of the mesonotum, Krog- 
mann and Vilhelmsen (2006) and Vilhelm- 
sen (2000) for muscles and internal mor- 
phology, and Harris (1979) for cuticular 
sculpturing. The paired campaniform sen- 

Table 1. Abbreviations used in figures. 

112 longitudinal flight n:\uscles 

114 mesoscutello-metanotal muscle 

2ph second phragma 

ass anterior scutellum setae 

cs campaniform sensilla 

cut cuticle 

edm electron dense mass 

edr electron dense ring 

ela electron lucent area 

epd epidermal cells 

fb fat body 

fl flange 

LI outer layer 

L2 inner layer 

m mitochondria 

ml midline between left and right longitudinal 

flight muscles 

ms mesoscutum 

pss posterior scutellum setae 

scl scutellum 

ssr scutoscutellar ridge 

sss scutoscutellar suture 

tb tubular body 

tsa transscutal articulation 

siUa on the scutellum have been termed 
campaniform sensilla (cs), and terminology 
specific to structures of the campaniform 
sensilla foUows Mclver (1985). Abbrevia- 
tions are Hsted in Table 1. 

Specimens. — Aphytis melinus for scanning 
electron microscopy (SEM) and transmis- 
sion electron microscopy (TEM) were 
obtained from a colony reared on Aspidio- 
tus nerii Bouche (Diaspididae) at the Uni- 
versity of California, Riverside. An addi- 
tional 30 specimens representing 23 species 
from ten families of Chalcidoidea, and one 
specimen of Mymarommatoidea were im- 
aged with SEM. A list of Chalcidoidea used 
for SEM imaging is given in Table 2; all 
material is represented by vouchers depos- 
ited at the University of California, River- 
side Entomology Research Museum 
(UCRC). The external morphology of the 
sensilla in Chalcidoidea and outgroups 
were more broadly surveyed, but this wiU 
be treated separately (Romero and Heraty, 
in prep.). ScuteUar sensillae have not been 
documented outside of Chalcidoidea and 


Journal of Hymenoptera Research 

Table 2. Sensillum diameters from SEM images, n indicates number of sensilla examined for that species. 


Shape of 

Maximum diameter 
mean ± SD (range) 

Average area of 


Aphytis melinus DeBach 7 circular 

Marietta sp. 4 circular 

Ablerus americanus Girault 2 circular 

Cales noacki Howard 3 circular 

Eretomocerus sp. 2 circular 

Eriaphytis sp. 1 circular 


Comperiella bifasciata Howard 
Microterys nietneri (Motschulsky) 


Orasema minutissima (Howard) 
GoUiimieUa antennata (Gahan) 

Pnigalio sp. 
Pnigalio agraules (Walker) 


Gonatocerus ashmeadi Girault 

Philotrypesis sp. 

Asaphes sp. 

Nasonia vitripennis (Walker) 


Signiphora sp. 

Tanaostigma sp. 

Epiclerus sp. 1 circular 


Megastigmus transvaalensis (Hussey) 1 circular 
Trichogra mmatidae 

Aphelinoidea sp. 2 circular 

Haeckeliania sp. 2 circular 

Hayatia sp. 2 circular 

Total 50 

4.94 |im 

2.25 ^im 
2.86 ^m 
2.99 ^im 
4.11 ^m 
5.10 ^im 

4 circular 4.10 )im 

2 circular 5.35 )im 

2 circular 1.87 ^im 

2 circular 3.47 |im 

3 subcircular 3.39 |im 
2 subcircular 3.93 |im 

1 circular 4.87 |im 

2 subcircular 3.70 |im 
1 subcircular 3.66 |im 
1 subcircular 4.34 ^im 

0.40 (3.68-4.94 ^m) 
0.27 (1.93-2.60 |im) 
0.37 (2.60-3.12 ^im) 
0.58 (2.32-3.39 ^m) 
0.17 (3.99^.23 ^m) 

0.38 (4.14-4.83 |im) 
0.62 (4.92-5.79 ^im) 

0.08 (1.81-1.93 um) 
0.45 (2.24-3.79 ^m) 

0.82 (2.82-4.44 |im) 
0.17 (3.80-4.05 [im) 

0.11 (3.78-3.62 [im) 

2 circular 2.99 ^im ± 0.20 (2.85-3.13 |im) 

1 circular 3.82 |im 

4.86 |im 

5.39 ^im 

5.07 nm ± 0.14 (4.97-5.17 ^m) 
5.05 nm ± 0.12 (4.96-5.13 ^m) 
3.82 ^im ± 0.30 (3.60-4.03 ^m) 

3.89 ^m ± 1.01 (1.81-5.79 [im) 

14.84 mm 

11.18 mm 

6.48 mm 

6.29 mm 

9.58 mm 

26.83 mm 

23.71 mm 
42.66 mm 

27.06 nm\ 
45.9 mm 

28.36 mm 
34.43 mm 

48.76 mm 

52.76 mm 

30.07 mm 


26.54 mm 

56.54 mm 

17.80 mm 

101.49 mm 

7.99 mm 

10.14 mm 

5.34 mm 

27.91 mm 

the majority of outgroup Hymenoptera 
examined had no trace of sensilla. How- 
ever, sensilla were found in species from 
four outgroup families, Ceraphronidae 
{Ceraphron sp.), Diapriidae {Trichopria sp.), 
Mymarommatidae (Mymaromma anomalum 
(Blood & Kryger)) and Scelionidae (Tele- 
nomus sp.). These families represent three 
different superfamilies from the subdivi- 

sion Proctotrupomorpha, which includes 
Chalcidoidea, and Ceraphronidae, repre- 
senting the more distantly related subdivi- 
sion Evaniomorpha. 

SEM. — Specimens selected for SEM were 
collected in 70% ethanol then dried in 
hexamethyldisilazane (HMDS) (Heraty 
and Hawks 1998). Some specimens were 
gradually rehydrated through a series of 

Volume 19, Number 1, 2010 


increasingly dilute ethanol baths, rinsed in 
two baths of deionized water, then di- 
gested in 10% KOH for 5-30 min according 
to the size of the specimen in order to clean 
the specimen of debris. Specimens were 
again rinsed in deionized water and 
dehydrated through a series of increas- 
ingly concentrated ethanol baths, then 
chemically dried in HMDS. Once dry, 
specimens were either dissected or placed 
whole onto SEM mounting stubs. Speci- 
mens were Au/Pd coated using a Cres- 
sington 108 Auto® sputter coater set for 60- 
90 seconds, then examined and digitally 
imaged under a XL30 PEG scanning 
electron microscope at 10 or 15 kV. 

Measurements. — Scutellar and sensillar 
measurements were taken in ImageJ 
1.38 X using the digital SEM images. Width 
measurements of the scutellum were made 
across the broadest point of the scutellum, 
excluding the axillula, and length measure- 
ments along the longest medial part of the 
scutellum including the frenum. Area 
measurements were made using the free- 
hand tool in ImageJ. Measurements of the 
differentiated area of the sensillum were 
taken along the longest axis and excluding 
the encircling ring, if present. To determine 
if there is a correlation between the size of 
the scutellum and the diameter of the 
sensilla, the length, width and area of the 
scutella of 50 specimens were measured 
(Table 2). A regression line was calculated 
for each of the three measurements of the 
scutellum that were graphed, and the 
coefficient of determination (R-squared) 
value calculated. 

TEM. — Live A. melinus were decapitated 
while immersed in Karnovsky's fixative 
(Karnovsky 1965). After approximately 
two hours they were placed in sodium 
cacodylate buffer, dehydrated in ethanol 
and embedded in Spurr resin (Spurr 1969). 
Sections approximately 60-70 ^im thick 
were cut using a diamond knife on a 
Leica Ultracut microtome. Sections were 
mounted on Electron Microscopy Sciences 
nickel slot grids coated with formvar/ 

carbon. Sections were then post stained 
using the SynapTec GridStick® system as 
follows. The uranyl acetate stain was 
diluted in methanol and the lead citrate 
stain mixed using 0.3 grams lead citrate, 0.3 
grams lead nitrate, 0.3 grams lead acetate 
and 0.6 grams sodium citrate dissolved in 
24.6 ml pre-boiled double distilled deio- 
nized water using a sonicator; after sonica- 
tion 5.4 ml of IN NaOH solution was 
added to the lead stain. Grids were initially 
stained for 5 minutes in uranyl acetate 
followed by two rinses in 100% methanol, 
one rinse each in 75%, 50% and 25% 
methanol, and four rinses in pre-boiled, 
double-distilled deionized water. The grids 
were then immediately stained for 10 min- 
utes in the lead stain followed by a 
30 second rinse in 0.02 N NaOH and 
30 minutes of rinsing with water changed 
every 5 minutes. Sections were examined 
with a Philips Tecnai 12 transmission 
electron microscope and digitally imaged 
using a model 780 Gatan DualVision 300 

Slide Mounts. — Aphytis melinus were col- 
lected in 70% ethanol and gradually hy- 
drated through a series of increasingly 
dilute ethanol baths, rinsed in two baths 
of deionized water and then digested in 
10% KOH for 10 minutes. Following 
digestion, specimens were rinsed in deio- 
nized water and dehydrated through a 
series of increasingly concentrated ethanol 
baths to 100% ethanol. They were then 
placed in a well plate with three drops of 
clove oil and the ethanol allowed to 
evaporate completely. The antennae, head, 
wings and body were separated from each 
specimen and arranged on the slide in 25% 
Canada Balsam and 75% clove oil (Noyes 
2003). As the clove oil evaporated, the 
Canada Balsam was gradually built up 
until the structures were covered and four 
5 mm coverslips applied. 


In most Apocrita, the mesonotum is 
divided by the transscutal articulation 


Journal of Hymenoptera Research 

/^^^ ^ 

Fig. 2. Structure of scutellum and scutellar sensilla in Aphytis melinus. a: scutellum, b: scutellar sensillum, c: 
underside of scutellum with tissue removed and both sensilla visible, d: underside of sensillum with tissue 
removed. Black arrows = 4.84fim, indicate equal distance in both (b) and (d). 

(Fig. 2a: tsa) into an anterior mesoscutum 
and a posterior scutellar-axillar complex 
(Gibson 1997). The medially located scu- 
tellum is separated from the anterolateral 
axilla by the scutoscutellar suture (Fig. 2a: 
sss). With the exception of Signiphoridae, 
in which the scutellum is reduced to a 
transverse band, all Chalcidoidea possess a 
scutellum that is a prominent plate of 
variable size and shape. The scutellum 
can be roughly circular, oval, shield or 
teardrop shaped and can also vary in 
topography. For example, some Encyrtidae 
have a rounded scutellum with sharply 
rising sides that form a dorsal hump, while 
some Mymaridae have a flat planar scu- 
tellum. Many chalcidoids (i.e. Euchariti- 
dae, Mymaridae, Pteromalidae, Tetracam- 

pidae and Torymidae) have a transverse 
sulcus or change in sculpture differentiat- 
ing a posterior region of the scutellum 
termed the frenum. In many taxa, lateral 
axillular grooves separate the axillula from 
the main portion of the scutellum, but this 
is often more apparent in lateral view. The 
scutellum of many smaller Chalcidoidea 
often has two pairs of prominent setae: the 
anterior scutellar setae (Fig. 2a: ass) and 
the posterior scutellar setae (Fig. 2a: pss). 
When present, these setae are used as 
reference points for the campaniform sen- 
silla on the scutellum. 

Structure of scutellum and sensilla in 
Aphytis. — A. melinus has a roughly oval 
scutellum with a pair of circular sensilla 
located medially to the four primary 

Volume 19, Number 1, 2010 


scutellar setae (Fig. 2a). Each campaniform 
sensillum appears externally as a smooth 
dome in the cuticle surrounded by a raised 
ring that interrupts the imbricate sculptur- 
ing of the scutellum (Fig. 2b). Internally, the 
cuticle forms a raised ring around an area of 
reticulate cuticle with an elliptical central 
depression oriented diagonally to the long- 
itudinal axis of the body (Fig. 2c-d). This 
ellipse-shaped thinning of the cuticle prob- 
ably creates a weakness along the long axis 
of the ellipse and enhances movement along 
the short axis conferring directional sensi- 
tivity similar to that obtained through an 
elliptically shaped cuticular cap (Moran and 
Rowley 1975). Across all of the specimens 
surveyed, the elliptical depression, which is 
also visible in slide mounts, was found only 
in Aphytis and Aphelinus (Aphelininae). 

Internally, the scutellum is bordered by 
several ridges forming a differentiated 
region directly above the longitudinal 
flight muscles. Along with the mesoscu- 
tello-metanotal muscles (Fig. 3a-e: 114) 
and randomly distributed fat body 
(Figs 3e: fb), this space also contains 
several unidentified structures. In certain 
dissections examined with SEM (Figs 3c, 
4a-b), there appears to be membranous 
divisions that run through this area defin- 
ing irregular sections as large as 20 fxm in 
diameter, however these divisions were 
not apparent in TEM preparations. Just 
below the cuticle, and between these 
divisions, there is a single, or sometimes 
double, layer of elongate epidermal cells 
(Fig. 4a-i: epd). While tightly packed, these 
cells appear independent of each other in 
SEM preparations (Fig. 4a-c) and in TEM 
preparations appear hollow due to a lack 
of penetration by the resin. Similar im- 
penetrable epidermal cells also appear in 
sections of the male antennae prepared by 
Romani et al. (1999) in their TEM investi- 
gation of the male antennae of A. melinus. It 
may be that in the adult wasp the 
epidermal cells have died leaving a thick 
waxy cell membrane that is impermeable 
to resin. These are not likely artifacts of 

dried haemolymph which is apparent in 
the layer of "tissue'" surrounding muscle 
114 and the sensillar stem (cs) in Fig. 3c. 
These cells line the entire internal surface 
of the cuticle, including ventral surfaces 
(Fig. 4i: epd) and internal apodemes 
(Fig. 4h: epd), but are absent where the 
scutellar sensilla attach to the cuticle 
(Fig- 4g). 

Mesoscutello-metanotal muscles. — In A. me- 
linus, a pair of muscles traverse the length 
of the scutellum between the longitudinal 
flight muscles and the dorsal surface of the 
scutellum (Fig. 3a-b and e), which are 
S5monymous with Kelsey's (1957) muscle 
114 and Vilhelmsen's (2000) mesoscutello- 
metanotal muscle. The muscles attach to 
the anterior portion of the scutellum just 
posterior to the scutoscutellar ridge 
(Fig. 3a-c: 114 and ssr). From this point of 
origin they narrow and are slightly angled 
medially to a posterior insertion to the 
anterior edge of the metanotum above the 
margin of the second phragma (2ph) to the 
anterior edge of the metanotum (Fig. 3a-b: 
114). In cross section, the longitudinal 
flight muscles have clearly defined axon 
bundles interspersed with mitochondria 
(Fig. 3d-e: 112), whereas the mesoscu- 
tello-metanotal muscle has mitochondria 
restricted to the periphery. Consequently 
axon bundles are not as easily distin- 
guished (Fig. 3d-e). These mucles may 
affect longitudinal tension of the scutellar 
disc and possibly deformation of shape in 
small soft-bodied chalcidoids. 

Ultrastructure of the sensillar cuticular 
cap. — In A. melinus, there are several dis- 
tinct features of the cuticular portion of the 
scutellar sensilla evident through electron 
microscopy. In cross sections there is a thin 
outer layer of solid cuticle. This layer 
(Fig. 5a-e: LI) sits external to a thicker 
layer of mesh-like cuticle (Fig. 5a-f: L2). 
These two layers of the cap are consistent 
with the cuticular structure found in 
campaniform sensilla observed in other 
studies where 2 or 3 layer-caps are re- 
ported (Mclver 1985) and it is nearly 


Journal of Hymenoptera Research 


Fig. 3. Mesoscutello-metanotal muscles in Aphytis melinus. a: underside of scutellum with most tissue 
removed leaving mesoscutello-metanotal muscles (114), b: same view in (a) with slightly different results from 
the chemical drying process, c: dorsal tissue found just beneath scutellum, d: cross section through mesoscutello- 
metanotal muscle, e: cross section through dorsal portion of scutellum. 

identical to the structure observed by 
Bromley et al. (1980) in aphid antennae. 
The bilayered cap is encircled by a flange 
that protrudes internally. This flange was 
observed by Mclver and Siemicki (1975) in 
the mosquito, and in the cockroach by 

Moran and Rowley (1975), who called the 
structure a cuticular collar. Moran and 
Rowley also suggested that it provides 
structural support and rigidity for the cap 
of the sensilla and enables the cap to move 
as a unit in response to cuticular deforma- 

Volume 19, Number 1, 




Fig. 4. Epidermal cells in Aphytis melinus. a: underside of scutellum with most tissue removed leaving 
epidermal cells, b: underside of scutellum with campaniform sensillum tissue and epidermal cells attached, c: 
underside of sensilla with epidermal cells attached, d-e: cross section through scutellum, f-g: cross section 
through scutellum with a campaniform sensillum, h: cross section through notal ridge, i: cross section through 
ventral portion of mesosoma. White boxes indicate area magnified in following figure. 

tion. Just dorsal to the flange, layer 1 is nation seems appropriate since it is at this 

attached to the cuticle by a ring of dark junction point, at the base of the flange 

staining cuticle (Fig. 5a-c) that Mclver and where the cap and cuticle meet, that the 

Siemicki (1975) called a hinge. This desig- cuticle would presumably bend. 


Journal of Hymenoptera Research 

Tubular body and electron dense mass. — 
One of the most distinctive features of a 
campaniform sensilla is the tubular body at 
the distal end of the nerve cell (Mclver 
1985), which is a bundle of microtubules 
set in an electron dense material that 
functions as the site of transduction 
(Thurm 1964). In A. melinus, the tubular 
body is a striated cap that inserts into layer 
two of the cuticle, almost extending to 
layer one (Fig. 5d: tb). The tubular body 
consists of microtubules perpendicularly 
oriented to the surface of the cuticle and set 
in an electron-dense material. An electron- 
lucent area located at the proximal end 
separates the tubular body from the elec- 
tron-dense mass beneath (Fig. 5c: ela and 
edm). In some preparations, the tubular 
body appears to have a slightly indented 
tip in the very center of its distal end 
(Fig. 5e). The proximal end of the cap-like 
tubular body is nested in an electron-dense 
mass (Fig. 5c: edm). This dense material 
surrounds the tubular body and is directly 
adjacent to the modified portions of the 
cuticular cap, completely filling the sunken 
areas below layer two and surrounding the 
flange (Fig. 5b-c). It also extends beyond 
the campaniform sensillum, particularly in 
the lateral direction, to form a large matt 
beneath the cuticle (Fig. 6a-d). The elec- 
tron-dense mass appears to consist of 
microtubules or lamella similar to the 
tormogen cell associated with campani- 
form sensilla found on mosquito palps 
(Mclver 1985), but appears to lack other 
cellular structures indicative of a tormogen 
cell such as a membrane bound nucleus 
(Thurm and Kiippers 1980). No other 
dendritic cells were identified in associa- 
tion with the campaniform sensilla. 

Distribution of sensilla across Chalcidoi- 
dea. — The paired scutellar sensilla are 
found in most families of Chalcidoidea 
and in exemplars of four outgroup families 
(Ceraphronidae, Mymarommatidae [single 
sensillum], Scelionidae and Diapriidae). In 
prepared slides, the sensilla appear as pale 
spots or thin areas in the cuticle and are 

readily identified in smaller taxa such as 
Aphytis (Rosen and DeBach 1979). In SEM 
preparations, they appear externally as 
differentiated areas of the cuticle that break 
the cuticular pattern and typically are 
ringed by raised or depressed cuticle 
(Figs 7a-h, 8a-h).The location and shape 
of the sensilla on the scutellum are highly 
variable across Chalcidoidea, but there is 
consistency within taxonomic groups at the 
family, tribe, genus and species levels 
(Romero and Heraty, in prep.). The loca- 
tion of the sensilla varies from medially 
abutting in some Aphelinidae, Encyrtidae 
and Mymaridae, to a lateral location within 
5 |j,m of the edge of the scutellum in some 
Pteromalidae and Eulophidae. Sensillar 
location also varies along the longitudinal 
axis from an anterior location contiguous 
with the scutoscuteUar suture (sss) in some 
Aphelinidae and Mymaridae to a posterior 
location within a few microns of the poster- 
ior margin in some Mymaridae. The sensilla 
are always found anterior to the frenal line 
when a frenum is present. The most 
common location is generally central and 
just medial of the anterior and posterior 
scutellar setae when present (Fig. 2a). The 
shape of the sensilla range from circular 
(Fig. 7a), to longitudinally oblong (Fig. 7d), 
to transversely oblong (Fig. 7c), with circu- 
lar being the predominant shape. For the 
subset of representative specimens mea- 
sured, the diameter of the sensiUa ranges 
from 1.81)im in Orasema sp. (Eucharitidae) 
to 5.79 |im in Microterys nietneri 
(Motschulsky) (Encyrtidae) (Table 2). 

Comparisons of sensillar diameter and 
scutellar length, width and area revealed 
that scutellar size accounts for very little 
variation in sensillar diameter (Fig. 9a-c). 
R-squared values were low with the high- 
est value at 0.081 (Fig. 9a). Comparisons of 
scutellar length and sensillar diameter had 
an R-squared value of 0.024 (Fig. 9b), and 
scutellar area and sensillar diameter had 
an R-squared value of 0.057 (Fig. 9c). The 
low R-squared values for the regression 
lines indicate that the variation in the size 

Volume 19, Number 1, 2010 


Fig. 5. Aphytis melinus, multiple specimens, a-c: cross sections through portions of campanifrom sensilla 
shown in (f), d: cross section through the tubular body, e: cross section through the tubular body showing divot 
at tip, f: the imderside of a campaniform sensilliom with tissue removed, dashed lines indicate the general 
location of cross sections in (a), (b) and (c). 


Journal of Hymexoptera Research 

Fig. 6. Electron-dense mass associated with the campaniform sensillum to the left of the midline in Aphytis 
meliniis. a: cross section anterior to campaniform sensillum, b-c: cross sections just anterior to the main structure 
of the campaniform sensillum, d: cross section through the center of the campaniform sensillum. 

of the scutellum accounts for less than 9% 
of the variation in the diameter of the 
sensilla. While there does not appear to be 
a relationship betv\^een size of scutellum 
and size of sensilla, the sensilla are absent 
or undetectable in chalcidoid families with 
the largest species, which have a scutellar 
size far beyond that indicated in Fig. 9 (i.e. 
Chalcididae and Perilampidae, >2 mm) 
(Romero and Heraty, in prep.). Thus our 
correlations are based only on taxa that are 
normally small in size, not those taxa that 
are large. A similar situation occurs in the 

outgroups, and we failed to find evidence 
of the sensilla in the majority of species 
which are usually larger or more heavily 
sclerotized. Strong correlations do appear 
within species (i.e. A. melinus, Fig. 9), but 
these were not correlated across subfamily 
or family groups. 


This is the first study to examine the 
morphology of scutellar sensilla of Chalci- 
doidea. The presence of sensilla has been 
noted sporadically in the literature, and 

Volume 19, Nl-mber 1, 2010 


Fig. 7. Variations in external appearance of scutellar sensilla. a: Aphytis melinus Q (Aphelinidae), b: A. melinus 
(S, c: Dirphys sp. (Aphelinidae), d: Pnigalio sp. (Eulophidae), e: Orasema minutissima (Eucharitidae), f: GoUumiella 
antennata (Eucharitidae), g: Coniperiella bifaciata Howard (Enc\Ttidae), h: Microterys nietneri (Encyrtidae). 

often included in illustrations without on the scutellum has been used in several 

comment in the text. The most attention keys to discriminate both individual spe- 

this feature has received is in Encarsia cies and species groups (Hayat 1998; 

(Aphelinidae), where relative placement Hemandez-Suarez et al. 2003; Schauff et 


Journal of Hymenoptera Research 

IGonatocerus ashmeadi 


^ Asaphes sp. 











2 pm 

2 pm 

Philotrypesis sp 

_ Palaeomymar sp 

9 >*-■ 

Fig. 8. Variations in external appearance of scutellar sensilla. a: Gonatocerus ashmeadi (Mymaridae), b: 
Philotrypesis sp. (Pteromalidae), c: Asaphes sp. (Pteromalidae), d: Nasonia vitripennis (Pteromalidae), e: 
Tanaostigma sp. (Tanaostigmatidae), f: Epiderus sp. (Tetracampidae), g: Megastigmus transvaalensis (Torymidae), 
h: Mymaromma anomalum (Mymarommatidae). Arrow indicates location of sensillum. 

Volume 19, Nl-mber 1, 2010 


al.l996). The sensilla have been referred to 
as scolopophorous sensilla (Gordh 1975b), 
and more recently as scutellar sensilla 
(Babcock et al. 2001; Hayat 1998; Herat}^ 
and Polaszek 2000), but most often as 
placoid sensilla (Annecke and Doutt 1961; 
Hayat 1998; Kim 2003; Rosen and DeBach 
1979; Schauff 1984). Both scolopophorous 
and placoid refer to specific types of 
sensilla described by Snodgrass (1935). 
Scolopophorus sensilla, also known as 
chordotonal organs, are composed of bun- 
dles of sensory cells that attach to a specific 
point on the cuticle in order to detect 
vibration. Placoid sensilla are composed of 
multiple sense cells and function as olfac- 
tory sensilla, often with numerous pores 
through the cuticle. The sensilla found on 
the chalcidoid scuteUum do not possess 
pores, and are innen'ated by a single ner\'e 
ceU as indicated by the lone tubular body 
(Fig. 6a). They are best defined as campani- 
form sensilla, which function as mechan- 
oreceptors (Pringle 1938a). 

ScuteUar sensilla are found in chalcidoid 
families with small-bodied species, but 
generally not in families with the largest- 
bodied species. There does not seem to be 
an absolute size at which the sensilla are 
consistently absent, but rather a trend 
where chalcidoid families with consistently 
small members such as ApheHnidae, En- 
cyrtidae and Trichogrammatidae possess 
the sensilla, those with consistently large 
members such as Chalcididae and Peri- 
lampidae do not, and those with inter- 
mediate-sized members such as Eulophi- 
dae and Torymidae have members with 
and without the sensilla (Romero and 
Heraty, in prep.). Presence does not appear 
to be necessarily correlated with degree of 
body sclerotization, as sensilla are retained 
in both larger members of soft-bodied 
Eulophidae and small well-sclerotized 
members of some Pteromahdae and Eu- 
charitidae. Scutellar sensilla were found in 
the outgroup families Ceraphronidae, Dia- 
priidae, SceUonidae and Mymarommati- 
dae. In these families the sensilla are onlv 

present in smaller species as is the case 
within Chalcidoidea. This would indicate 
that presence of scutellar sensilla is plesio- 
morphic for Chalcidoidea, and their sub- 
sequent loss derived. 

In chalcidoids that possess scutellar 
sensilla, the sensilla vary in size, shape 
and location. The shape varies from circu- 
lar to oval, with both shapes commonly 
obser^'ed in campaniform sensilla recorded 
from other studies (Mclver 1985). The size 
of the sensilla in the subset of taxa 
measured ranges from 1.81 |im to 5.79 .um. 
While this is over a 3 fold difference, 
campaniform sensilla have been recorded 
as small as 1 \xm (Hawke et al. 1973) and as 
large as 30 um in other insects (Hustert et 
al. 1981). Chalcidoid scutellar sensilla gen- 
erally fall in the 1.5-10 fim range as 
reported for most campaniform sensilla 
(Amomsak et al. 1998; Blaney and Chap- 
man 1969; Chevalier 1969; Mclver 1985; 
Moran and Rowley 1975; Schmidt and 
Smith 1985, 1987). 

Variation in size of the sensiQa does not 
seem to be closely tied to the size of the 
scuteUum (Fig. 9a-c). The low R-squared 
values (< 0.081) indicate that size of the 
scuteUum is a poor predictor of the size of 
the sensiUa. In several specimens such as 
Golliimiella antennata (Gahan) (Euchariti- 
dae) and Microtenjs nietneri, there is as 
much as a 0.87 um difference between the 
left and right sensiQa on the same speci- 
men. In species represented by multiple 
specimens, such as A. melinus, there ap- 
pears to be a relation between size of 
sensiUa and size of scuteUum with larger 
specimens possessing larger sensilla 
(Fig. 9a-c: A. melinus). This intraspecific 
trend between sensUlar size and specimen 
size was also obser\-ed by Schmidt and 
Smith (1985) in their examination of wing 
sensiUa in 18 specimens of Trichogramma 
minutum RUey (Trichogrammatidae). 

The shape of the campaniform sensiUa 
has been shown to confer directional 
sensitivit}^ (Pringle 1961). Circular sensiQa 
do not have directional sensitivity whereas 


Journal of Hymenoptera Research 

Width of scutellum in }im 


O 3 

^ 2 



X ".•=A+* 





■ — ' — 

O ^ 


? +. 


4- ■ 

▲ = 



R^ = 


50 100 150 200 250 300 

Length of scutellum in jim 



.5 2 J 

A = 

R'= 0.05733 

— r- 



— r— 

Area of scutellum in mm 



♦ Aphytis melimts 

= Marietta sp. 

▲ Ablerus americanus 

X Cales noacki 
% Eretomocerus sp. 
Eriaphytls sp. 


+ Comperiella bifasciata 

D Microtefys nietneri 


— Orasema sp. 

♦ Golliimiella antennata 


■ Pnigalio sp. 
Pnjgalio agraules 

X Gonatocerus ashmeadi 

>|^ Philotiypesis sp. 

O Asaphes sp. 

+ SignipJiora sp. 

O Tanaostigma sp. 


A Epi clems sp. 


♦ Megastigmus transvaalensis 

Aphelinoidea sp. 

— Haeckeiiania sp. 
+ Hayatia sp. 

Fig. 9. Diameter of scutellar sensilla and size of scutellum for various chalcidoids. a: Scutellum width and 
sensillum diameter, b: Scutellum length and sensillum diameter, c: Scutellum area and sensillum diameter. 

Volume 19, Number 1, 2010 


oval sensilla have more sensitivity to 
cuticular strain along the major axis 
(Moran and Rowley 1975). In chalcidoids, 
the scutellar sensilla can be circular or oval, 
with oval sensilla found oriented transver- 
sely in some Aphelinidae and longitudin- 
ally in some Eulophidae and Pteromalidae 
(Romero and Heraty, in prep.). The varia- 
bility in the shape as well as in the location 
of the sensilla on the scutellum is most 
likely related to the variable morphology of 
the scutellum and the mesonotum as a 
whole. Different combinations of morpho- 
logical features such as thickness and 
shape of the scutellum, placement of 
internal ridges, and relation of the scutel- 
lum to other sclerites could be associated 
with diverse complimentary positions of 
mechanoreceptors such as the scutellar 
sensilla. The consistent presence of the 
sensilla across entire families seems to 
indicate that there is strong selective 
pressure to maintain the sensilla. The 
symmetric variation in shape and position 
of the sensilla and consistency within 
phylogenetic lineages (Romero and Heraty, 
in prep.), further support the hypothesis 
that variation in the sensilla is tied to 
optimal functionality. 

Scutellar sensilla are notably absent in 
taxa with larger members. One explanation 
for the exclusive presence of the sensilla in 
smaller chalcidoids is that these smaller 
insects employ unique flight techniques 
not often utilized by larger insects. In his 
paper on flight and lift production, Weis- 
Fogh (1973) observed a novel wing motion 
mechanism in small (1-2 mm) Encarsia 
formosa Gahan (Aphelinidae), later termed 
clap-and-fling. This mechanism involves 
the right and left wing meeting, or clap- 
ping, at the end of the up-stroke and 
beginning of the down-stroke and has been 
observed in other small insects with low 
Reynolds numbers such as the greenhouse 
whitefly (Weis-Fogh 1975) and thrips (El- 
lington 1984). Miller and Peskin (2005) 
have shown that clap-and-fling provides 
less of a lift enhancing effect for insects 

with intermediate Re3molds numbers, such 
as the fruit fly, Drosophila melanogaster 
Meigen (Diptera: Drosophilidae) (Re =64). 
While larger insects are known to use clap- 
and-fling when tethered (Vogel 1966; Gotz 
1987; Zanker 1990), when carrying large 
loads (Marden 1987), and when perform- 
ing certain steering maneuvers (Cooter and 
Baker 1977; Ellington 1984), it seems very 
small insects are obliged to utilize clap- 
and-fling on a regular basis (Lehmann et al. 
2005; Miller and Peskin 2005). The scutel- 
lum is uniquely placed between the junc- 
tion of the wing base and the lateral 
anterior connection of the posterior attach- 
ment of the longitudinal flight muscles (2'"'^ 
phragma) and thus could play an impor- 
tant role in monitoring flight activity. As 
mechanoreceptors, the campaniform sen- 
silla are likely measuring changes in the 
torsion and tension of the scutellar cuticle. 
It specifically relates to a small insect flight 
specialization such as clap-and-fling, and 
consequently scutellar sensilla are consis- 
tently retained in the smallest of chalci- 
doids and other Hymenoptera. 


We would like to thank Eric Ragsdale and 
CFAMM for support in microscopy preparation, and 
Johan Liljebad, Roger Burks, Jeremiah George, James 
Munro and Jason Mottern for discussions over 
sensillar distribution patterns. We would like to 
acknowledge support from the National Science 
Foundation grants TOL EL-0341149 and FEET DEB- 


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Vol. 19(1), 2010, pp. 179-186 

A New Species of Lysiphlebus Forster 1862 (Hymenoptera: Braconidae, 

Aphidiinae) Attacking Soybean Aphid, Aphis glycines Matsumura 

(Hemiptera: Aphididae) from China 

Petr Stary, Ehsan Rakhshani, Zeljko Tomanovic, Kim Hoelmer, 
NiCKOLAS G. Kavallieratos, Juanjuan Yu, Mengqing Wang and George E. Heimpel 

(PS) Institute of Entomology, Biology centre. Academy of Sciences of the Czech Republic, 

Branisovska 31, 37005 Ceske Budejovice, Czech Republic; 

(ER) Department of Plant Protection, College of Agriculture, University of Zabol, Zabol, P.O. Box: 

98615-538, 1. R. Iran; 
(ZT) Institute of Zoology, Faculty of Biology, University of Belgrade, Studentski trg 16, 11000 

Belgrade, Serbia; 

(KH) USDA Agricultural Research Service, Beneficial Insects Introduction Research Unit, 501 South 

Chapel Street, Newark, DE 19713-3814, USA; 

(NGK) Laboratory of Agricultural Entomology, Department of Entomology & Agricultural 

Zoology, Benaki Phytopathological Institute, 8 Stefanou Delta str, 145 61 Kifissia, Attica, Greece; 

nick_kaval@hotmail . coni 

(JY) Chinese Academy of Science, Institute of Zoology, 1 Beichen West Road Chaoyang District, 

Beijing, 100101, China; 

(MW) State Key Laboratory for Biology of Plant Diseases and Insect Pests, Institute of Plant 

Protection, Chinese Academy of Agricultural Science, 12 South Zhongguancun Rd., Beijing, 100081, 

China; and USDA-ARS Sino- American Biological Control Laboratory, 12 South Zhongguancun Rd., 

Beijing, 100081, China; 

(GEH) Department of Entomology, University of Minnesota, 219 Hodson Hall, 1980 Folwell Ave., 

St. Paul, MN 55108, USA; 

Abstract. — Lysiphlebus orientalis sp. n. is described from China. The new species was reared 
from Aphis glycines Matsumura/ Glycine max association. On the basis of the fore wing venation 
pattern (short Rl vein) and the number of maxillary and labial palpomeres, we can preliminarily 
classify L. orientalis sp.n. as a member of the "testaceipes Cresson" species-group. Laboratory 
populations of L. orientalis are thelytokous, the first record of this phenomenon in this species 

Key words. — Lysiphlebus orientalis sp.n., aphid parasitoids. Glycine max 

The soybean aphid. Aphis glycines Mat- genie viruses (Halbert et al. 1986; van den 

sumura (Hemiptera: Aphididae), is native Berg et al. 1997; Wang et al. 1994). 
to northeast Asia (China, Korea and Japan) In Asia, the soybean aphid is attacked by 

but was discovered infesting soybean braconid and aphelinid parasitoids (Chang 

fields in North America beginning in et al. 1994; Wu et al. 2004b; Liu et al. 2004; 

summer 2000 (Venette and Ragsdale 2004) Miao et al. 2007). The main natural enemies 

and has become a serious pest throughout of soybean in North America are native 

soybean-growing areas of the Midwest and naturalized predators, including lady 

(Ragsdale et al. 2004, 2007). These aphids beetles, minute pirate bugs and predatory 

not only devastate soybean plants by direct flies (Fox et al. 2004, 2005; Rutledge et al. 

feeding, but they also spread plant-patho- 2004; Rutledge and O'Neil 2005; Costa- 


Journal of Hymenoptera Research 

magna and Landis 2006, 2007; Desneux et 
al. 2006; Donaldson et al. 2007; Gardiner 
and Landis 2007; Chacon et al. 2008; 
Costamagna et al. 2008; Gardiner et al. 
2009). Parasitoids attacking soybean aphid 
in North America have been rare, on the 
other hand (Lin and Ives 2003; Kaiser et al. 
2007; Noma and Brewer 2008; Pike et al. 
2007), with the exception of relatively high 
parasitism by aphidiine braconids reported 
in New York state (Nielsen and Hajek 
2005) and also by Aphelinus certus, an 
accidentally-introduced species, in eastern 
North America (Heraty et al. 2007; Heim- 
pel et al. in press). 

Initial biological control efforts directed 
at the soybean aphid have resulted in the 
importation of several aphid parasitoids 
and predators from China, Japan and 
South Korea into quarantine, including a 
strain of an aphelinid parasitoid from 
Japan in 2001 (Heimpel et al. 2004; Wu et 
al. 2004a) and at least two species of the 
aphidiine braconid genus Binodoxys (Wyc- 
khuys et al. 2007; Desneux et al. 2009a; 
Desneux et al. 2009b). In addition to these 
species, recent ongoing research on the 
introduction of the braconid parasitoids of 
soybean aphid from China has yielded the 
discovery of a new species of Lysiphlebus 
Forster contributing to our current revi- 
sionary work on the subtribe Lysiphlebina 
Mackauer. Here we describe the new 
species and discuss its identity and possi- 
ble distributional pattern. 


Parasitoids were obtained by collecting 
samples of Glycine max plants colonized by 
A. glycines in commercial and experimental 
fields in China. Collections at field sites in 
northeastern Chinese provinces were made 
by K. Hoelmer, J. Yu and M. Wang during 
June, July and August of 2006 in the 
vicinity of Harbin, Heilongjiang province, 
and Xiuyan, Liaoning province. Leaves 
and stems of plants with aphid colonies 
were cut, placed into plastic zip-lock bags 
and held in chilled picnic coolers while in 

the field, then transferred to the laboratory 
where they were kept in containers cov- 
ered with nylon mesh at room temperature 
during sample processing. Emergent 
adults were collected in vials and mummi- 
fied aphids containing developing para- 
sitoids were placed individually in wells of 
plastic microtiter plates and sealed with 
corks for shipment. Vials with adults were 
streaked with honey, and each microtiter 
plate well was given a small droplet of 
honey to sustain adults that emerged 
during transit. Parasitoids were shipped 
to the USDA ARS biological control quar- 
antine laboratory in Newark, Delaware, 
USA. Quarantine cultures of A. glycines 
parasitoids were initiated at Newark from 
these shipments and maintained on A. 
glycines on soybean. After establishment, 
a portion of the cultures were transferred 
to the University of Minnesota, USA to 
support host range evaluations. The new 
species of Lysiphlebus was discovered dur- 
ing these evaluations. Slides were made of 
dissected specimens using Hoyer's media. 
External morphology was studied using an 
Olympus SZX9 stereomicroscope. Illustra- 
tions were made based on slide-mounted 
specimens using an Olympus BH2 Phase- 
contrast microscope with a drawing tube. 
Morphological terminology follows Stary 
(1973) and Sharkey and Wharton (1997). 
Subdivisions of the flagellum are referred 
to as segments in order to maintain 
consistency with other taxonomic works 
on Aphidiinae. Measurements in the de- 
scription were taken using an ocular 
micrometer. See Tomic et al. (2005) for 
more details regarding measurements. 


Diagnosis. — On the basis of the fore wing 
venation pattern (short Rl vein, which is 
equally or slightly shorter than the stigma) 
(Fig. 5) and the number of maxillary and 
labial palpomeres (three maxillary and two 
labial palpomeres) (Fig. 1), we prelimina- 
rily classify Lysiphlebus orientalis sp.n. as a 
member of the "testaceipes Cresson'' spe- 

Volume 19, Number 1, 2010 


Figs 1-9. Lysiphlebus orientalis sp. n., female paratype. 1, head and mouthparts; 2, mesoscutum; 3, first and 
second flagellar segments; 4, antenna; 5, fore wing; 6, propodeum; 7, petiole; 8, hind leg; 9, genitalia. 

cies-group. Lysiphlebus orientalis sp. n. 12-seginented antennae vs. 13-14-segmen- 

differs from the nominate species L. testa- ted antennae of L. testaceipes). Also, flagel- 

ceipes by having a smaller number of lomeres 1 and 2 of L. orz£7tffl/zs sp. n. bear 1- 

antennal segments (L. orientalis sp. n. has 2 and 2-3 longitudinal placodes, respec- 


Journal of Hymenoptera Research 

tively, but L. testaceipes has 4-6 longitudinal 
placodes on flagellomere 1 and 5-7 on 
flagellomere 2. Lysiphlebus orientalis sp. n. 
has an elongately triangular stigma (stigma 
length/width ratio of 2.9-3.2), but L. 
testaceipes has a widely triangular stigma 
(stigma length/width ratio of 2.4-2.6). In 
addition, L. orientalis sp. n. differs from all 
other species in having short marginal fore 
wing setae. All other species of the ''testa- 
ceipes " species-group have long marginal 
fore wing setae. 

Lysiphlebus orientalis Stary & Rakhshani 
sp. n. 

(Figs 1-12) 

Female: Head (Fig. 1) transverse, wider 
than mesosoma at tegulae, bearing sparse 
setae. Eyes medium sized, oval, laterally 
prominent. Face laterally pubescent. Ten- 
torial index (tentoriocular line/intertentor- 
ial line) 0.49-0.50, Clypeus slightly pro- 
truding with 5-6 long setae. Labrum 
distinct, with 2 short setae on outer margin. 
Malar space equal to 0.28-0.30 of long- 
itudinal eye diameter. Mandible bidentate, 
with 7-9 setae in outer surface. Maxillary 
palpi with 3 palpomeres, labial palpi with 2 
palpomeres. Antenna 12-segmented (scape 
and pedicel as primary segments and 10 
flagellomeres), filiform (Fig. 4). Pedicel 
subsphaerical. Fi (Fig. 3) equal or slightly 
longer than F2 and 2.2-2.6 X as long as its 
maximum width. Fi and F2 bearing 1-2 
and 2-3 longitudinal placodes respectively 
(Fig. 3). Flagellomeres covered uniformly 
with semi-erect setae. 

Mesosoma - Mesoscutum (Fig. 2) smooth, 
covering pronotum from above; notaulices 
distinct in ascedent portion of anterolateral 
margin, effaced dorsally, with 6-7 long 
setae along laterodorsal part of mesoscu- 
tum. Scutellum subquadrate, bearing 2 
long setae at each lateral margin. Propo- 
deum (Fig. 6) smooth, with two divergent 
carinae at base, (some specimens manifest 
indications of ''pseudo-carination" or rug- 

osities in upper part of propodeum, estab- 
lishing incomplete central areola). Upper 
and lower parts of propodeum with 
2-3 and 1-2 long setae on each side. 
Fore wing (Fig. 5) densely pubescent, 
lower marginal setae short, equal to 
those on surface; stigma, 2.9-3.2 X as long 
as its width, 1.00-1.27 as long as Rl vein 
(= metacarpus). Ri vein 1. 35-1. 40 X as long 
as R2 vein, 2.00-2.10 X as long as rs-m vein. 
Hind femur with short adpressed setae 
(Fig. 8). 

Metasoma - Petiole (Fig. 7) elongate, 
smooth, slightly convex dorsally, with 
lateral depression after prominent spiracu- 
lar tubercles, positioned midsegment; its 
length 2.00-2.25 X its width at spiracles, 
1.50-1.70 X its width at base; 4-5 setae 
positioned on posterior laterodorsal mar- 
gin, one long seta posterior to spiracles. 
Ovipositor sheath (Fig. 9) short, wide at 
base, dorsally slightly convex, narrowed 
toward tip, apically truncated, bearing four 
long setae at tip and 4-5 shorter scattered 
setae on lateral and dorsal surface. Length 
of ovipositor sheath 1.8-2.0 X its maximum 
width at base, 4.2-4.4 X its minimum width 
at tip. Second valvula with smooth dorsal 

Body length: 1.5-1.7 mm 

Coloration: General body color light 
brown, head and antenna light brown, 
mouthparts except tips of mandible yel- 
lowish. Pronotum brown; mesoscutum and 
mesopleuron dark brown. Propodeum 
brown. Legs yellow, hind leg with brown 
dorsal outlines. Wings hyaline, venation 
yellowish brown. Propodeum brown. Pe- 
tiole yellow, other metasomal terga light 
brown, dorsally darker. Ovipositor sheath 
dark brown. 

Male: Antenna 14-segmented. Maxillary 
palpi with 3 palpomeres, labial palpi with 2 
palpomeres. Fore wing venation as in 
female (Fig. 10). Fore wing lower marginal 
setae distinctly longer than those on sur- 
face; stigma widely triangular, 2.8-3.1 X as 
long as its width. Petiole (Fig. 11) elon- 
gately quadrangular, 1.75-1.85 X its width 

Volume 19, Number 1, 2010 


■"^i^S^^^^"'" ' 

Figs 10-12. Lysiphlebus orientalis sp. n., male. 10, fore wing; 11, petiole; 12, genitalia. 

at spiracles, with lateral depression after 
spiracular tubercles. Aedeagus subtriangu- 
lar (Fig. 12) with subparallel posterolateral 
margins and short tip. Body darker than 
female, head and thorax black brown, 
antenna dark brown, mouthparts yellow- 
ish brown. Wings slightly translucent. Legs 
brown with light yellow patches at ventral 
and tip of segments. Petiole light brown, 
other metasomal segments greyish brown. 
Body length: 1.5-1.6 mm. 


Holotype: 9 reared from Aphis glycines 
Matsumura on Glycine max (L.) Merrill, CHINA, 

Harbin, VIII 2006, Leg. K. Hoelmer, laboratory 
culture, reared on Aphis glycines on Glycine max, 
University of Minnesota, USA, 2008, G. E. 
Heimpel (Collection of United States National 
Museum of Natural History). 

Paratypes (same sampling data as holotype): 
39 and S paratypes are deposited in the 
collection of United States National Museum 
of Natural History. 69 parat}j»es are deposited 
in the collection of P. Stary (Ceske Budejovice). 
39 paratypes are deposited in the collection of 
Institute of Zoology, Chinese Academy of 
Sciences, Beijing, China. 79 and 29 paratypes 
are deposited in the collection of Institute of 
Zoology, Faculty of Biology, University of 
Belgrade (Serbia) and in collection of University 
of Zabol (Iran), respectively. 


Journal of Hymenoptera Research 

Additional material: 209 and 2o with same 
sampling data as holotype deposited in the 
collection of Institute of Zoology, Faculty of 
Biology, Universit\' of Belgrade (Serbia). 


Lysiphlebus orientalis n. sp. is a new 
member of the " testaceipes" species group, 
a tentative taxon which has previously 
been classified within subgenus Phlebus 
(Stary 1975). The group includes the 
species distributed within a specific geo- 
graphic area of which only L. testaceipes 
Cresson has a wider distribution, assumed 
to be due to introduction and expansion of 
its range in combination with its opportu- 
nistic host range (Kavallieratos and Ly- 
kouressis 1999, 2004; Kavallieratos et al. 
2001; Pons et al. 2004; Stary et al. 2004). 
Originally believed to be a North American 
species, L. testaceipes has also been recov- 
ered from the east Palaearctic (Stary et al. 
2002). The known distribution of L. orienta- 
lis is northeast China, but further research 
may document a broader distribution. 
Other taxa which should be preliminarily 
included in the "testaceipes" group are: L. 
fritzmuelleri Mackauer (Europe), L. deser- 
torum Stary (Central Asia), L. ussuriensis 
Kiriac (Far East) and L. utahensis (Smith) 

Examination of the field-collected speci- 
mens revealed a highly skewed female: 
male sex ratio. However, laboratory cul- 
tured material comprised a mostly or 
completely uniparental population, a phe- 
nomenon that has not been previously 
recorded in any member of the testaceipes 
group, although it occurs in other Lysiphle- 
bus species, namely L. fabarum (Marshall), 
L. cardui (Marshall), and L. confusus Trem- 
blay and Eady (Belshaw et al. 1999; Stary 
1999; Stary et al. 2002), which belong to the 
subgenus Phlebus Stary. Further investiga- 
tions are needed to elucidate the nature 
and distribution of thelytoky in the respec- 
tive groups. 

Our ongoing research on the subtribe 
Lysiphlebina Mackauer reveals several 

Lysiphlebus species in Europe that are 
putatively related to L. orientalis sp.n. on 
the basis of morphological characters. We 
shall resolve the taxonomic status of the 
aforementioned Lysiphlebus taxa and their 
possible relations with L. orientalis sp.n. 
using molecular markers in a future con- 


We thank Dr. Ge-Xia Qiao (Institute of Zoology, 
Chinese Academy of Science, Beijing) and Hongyin 
Chen (Institute of Plant Protection, Chinese Academy 
of Agricultural Science, Beijing, and the USDA ARS 
Sino- American Biological Control Laboratory, Beijing) 
and their students for assistance with the logistics and 
processing of field collections, Kathryn Lanier and 
Keith Hopper (USDA ARS, Newark, Delaware, USA) 
for initiating and maintaining the L. orientalis culture 
at the Beneficial Insects Introduction Research Unit in 
Newark, and Zeynep Sezen for help with rearing in 
Minnesota. This study was also supported by the 
Ministry of Science and Technological Developments 
of the Republic of Serbia (143006B), the Entomology 
Institute Project Z50070508 (Academy of Sciences of 
the Czech Republic) and grant No. 86-19, University 
of Zabol, Iran, by the North-Central Soybean Research 
Program in the United States, and by the Minnesota 
Agricultural Experiment Station. 


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Pike, K. S., P. Stary, M. I. Brewer, T. Noma, S. Langley, 
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de Lysiphlehus testaceipes (Cresson) (Hym., Braco- 
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Ragsdale, D. W., B. P. McComack, R. C. Venette, B. D. 
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K. D. Johnson, R. J. O'Neil, C. D. Difonzo, T. E. 
Hunt, P. A. Glogoza, and E. M. CuUen. 2007. 
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nniptera : Aphididae). Journal of Economic Ento- 
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, D. J. Voegtlin, and R. J. O'Neil. 2004. Soybean 

aphid biology in North America. Annals of the 
Entomological Society of America 97: 204-208. 

Rutledge, C. E., R. J. O'Neil, T. B. Fox, and D. A. 
Landis. 2004. Soybean aphid predators and their 
use in integrated pest management. Annals of the 
Eyitomological Society of America 97: 240-248. 

and R. J. O'Neil. 2005. Orius insidiosus (Say) as 

a predator of the soybean aphid. Aphis glycines 
Matsumura. Biological Control 33: 56-64. 

Sharkey, M. J. and R. A. Wharton. 1997. Morphology 
and terminology. Pp. 19-37 in: Wharton, R. A., P. 
M. Marsh, and M. J. Sharkey eds. Manual of the 
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Stary, P. 1973. A review of the Aphidius species 
(Hymenoptera: Aphidiidae) of Europe. Annota- 
tiones Zoologicae et Botanicae 84: 1-85. 

. 1975. The subgeneric classification of Lysiphle- 
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. 1999. Biology and distribution of microbe- 
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, J. Havelka, and J. Y. Choi. 2002. New species 

and populations of Lysiphlehus Foerster - aphid 
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, B. Lumbierres, and X. Pons. 2004. Opportu- 
nistic changes in the host range of Lysiphlehus 
testaceipes (Cr.), an exotic aphid parasitoid ex- 
panding in the Iberian Peninsula. Journal of Pest 
Science 77: 139-144. 

Tomic, M., Z. Tomanovic, N. G. Kavallieratos, 
P. Stary, C. G. Athanassiou, V. Tomic, and L. 
Lucie. 2005. Morphological variability of several 
biotypes of Ephedrus plagiator (Nees) (Hymenop- 
tera: Braconidae: Aphidiinae) with the descrip- 
tion of a new species. Zoologischer Anzeiger 244: 

van den Berg, H., D. Ankasah, A. Muhammad, R. 
Rush, A. Widayanto, H. B. Wirasto, and I. Yully. 
1997. Evaluating the role of predation in popula- 
tion fluctuations of the soybean aphid. Aphis 
glycines in farmers' fields in Indonesia. Journal of 
Applied Entomology 34: 971-984. 

Venette, R. C. and D. W. Ragsdale. 2004. Assessing the 
invasion by soybean aphid (Hemiptera: Aphidi- 
dae): where will it end? Annals of the Entomological 
Society of America 97: 219-228. 

Wang, X. B., C. H. Fang, X. P. Zheng, Z. Z. Lin, L. R. 
Zhang, and H. D. Wang. 1994. A study on the 
damage and economic threshold of the soybean 
aphid at the seedling stage. Plant Protection 20: 

Wu, Z., K. R. Hopper, R. J. O'Neil, D. J. Voegtlin, D. R. 
Prokrym, and G. E. Heimpel. 2004a. Reproductive 
compatibility and genetic variation between two 
strains of Aphelinus albipodus (Hymenoptera: 
Aphelinidae), a parasitoid of the soybean aphid. 
Aphis glycines (Homoptera: Aphididae). Biological 
Control 31: 311-319. 

, D. Schenk-Hamlin, W. Zhan, D. W. Ragsdale, 

and G. E. Heimpel. 2004b. The soybean aphid in 
China-an historical review. Annals of the Entomo- 
logical Society of America 97: 209-218. 

Wyckhuys, K. A. G., K. R. Hopper, K.-M. Wu, C. 
Straub, C. Cratton, and G. E. Heimpel. 2007. 
Predicting potential ecological impact of soybean 
aphid biological control introductions. Biocontrol 
News and Information 28 (2): 30-34. 

Vol. 19(1), 2010, pp. 187-188 


Chromosomes of Blastophaga psenes (Hymenoptera: Agaonidae) 

Vladimir E. Gokhman, Andrey P. Mikhailenko and Victor N. Fursov 

(VEG, APM) Botanical Garden, Moscow State University, Moscow 119991 Russia 
(VNF) Schmalhausen Institute of Zoology, National Academy of Sciences of Ukraine, 

Kiev 01601 Ukraine 

Parasitic w^asps are one of the largest and 
most taxonorrdcally complicated groups of 
insects (Rasnits3m 1980). They play a very 
important role in food chains as parasitoids 
of many insect pests of agriculture and 
forestry. In addition, certain species of 
'parasitic' Hymenoptera are associated 
with plants, either as pests or pollinators 
(Quicke 1997). Chromosomes of about 420 
species of parasitic w^asps have been 
studied (Gokhman 2009). However, chro- 
mosomes of the medium-sized family 
Agaonidae that is associated with fruits of 
the plant genus Ficus, were never exam- 
ined before. We have managed to study the 
karyotype of Blastophaga psenes (Linnaeus), 
the sole pollinator of the edible fig, Ficus 
carica Linnaeus. The description of the 
karyotype is given below. 

Syconia of the cultivated form of F. carica 
that contained immature stages of B. psenes, 
were collected by V.N. Fursov at Nikitsky 
Botanical Garden, Ukrainian Academy of 
Agrarian Sciences (about 5 km E Yalta, the 
Crimea, South Ukraine) on 2-10 October 
2008, preserved at 10-12°C for three to four 
months and then incubated for a few days 
at room temperature. Cerebral ganglia of 
prepupae were used for karyotyping ac- 
cording to the technique developed by 
Imai et al. (1988). Chromosomes of a single 
male and five females were studied. 
Micrographs of chromosomes were ob- 
tained using Zeiss Axioskop 40 EL optic 
microscope fitted with Zeiss AxioCam 
MRc digital camera. Chromosomes of five 
diploid metaphase plates were measured 
on digital micrographs using Zeiss AxioVi- 

sion; all chromosomes were then arranged 
according to the classification provided by 
Levan et al. (1964). Voucher adult speci- 
mens of B. psenes are deposited in the 
Zoological Museum, Moscow State Uni- 
versity, Moscow, Russia. 


The chromosomal study of female in- 
dividuals of B. psenes has revealed a 
chromosome set of 2n = 12 (Eig. 1). An 
analogous study of the male individual has 
yielded very few metaphases with n = 6 
(not shown here). The haploid karyotype of 
this species comprises five large meta- 
centric chromosomes and a smaller sub- 
telocentric one (Eig. 2, Table 1). 

The families Agaonidae (at least those 
belonging to Agaoninae; Rasplus et al. 
1998), Torymidae and Ormyridae are 
usually believed to form a common clade 
(Noyes 1990, see also Boucek 1988 and 
Gibson et al. 1999). The karyotype structure 
similar to that found in B. psenes (five large 
metacentrics and a smaller subtelocentric/ 
acrocentric) is also characteristic of many 
Torymidae (including most species of the 
less advanced subfamily Toryminae) and 
one of the two studied species of the 
Ormyridae that belong to the genus Or- 
myrus (see Gokhman 2009 for review). 

w w K a w 




Fig. 1. Kar)^ogTam of the diploid karyotype of 
Blastophaga psenes. 


Journal of Hymenoptera Research 

Table 1. 

Parameters of chromosomes of Blastophaga 

Fig. 2. Ideogram of the haploid karyotype of 
Blastophaga psenes. 

Moreover, karyotypes of a few species of 
the Torymidae and Ormyridae that contain 
only five metacentric chroraosomes are 
obviously derived from the preceding ones 
through tandem fusions, analogous, for 
example, to certain Eulophidae with simi- 
lar chromosome sets (Gokhman 2009). The 
karyotype structure of B. psenes therefore 
represents the ground plan feature of the 
common clade of the Torymidae, Ormyr- 
idae and Agaonidae. On the other hand, 
karyotypes of many Pteromalidae (another 
group that is probably related to Agaoni- 
dae S.I.; Campbell et al. 2000) also comprise 
five biarmed chromosomes, and those 
chromosome sets could originate as well 
from a karyotype with an additional sub- 
telocentric/ acrocentric through chromoso- 
mal fusion (Gokhman 2009). 


The authors are grateful to the Director and staff of 
Nikitsky Botanical Garden (Dr. V.N. Ezhov and Drs. 
A.V. Smykov, E.L. Shishkina and A.N. Kazas respec- 

Chromosome no. 

Relative length 

Centromere index 



















tively) for their kind permission to collect syconia of F. 
carica at the Garden. The present study was partly 
supported by the research grant no. 07-04-00326 from 
the Russian Foundation for Basic Research to VEG 
and APM. 


Boucek, Z. 1988. Australasian Chalcidoidea (Hymenop- 
tera). A biosystematic revision of genera of fourteen 
families, with a reclassification of species. CAB 
International, Wallingford. 832 pp. 

Campbell B., J. Heraty, J.-Y. Rasplus, K. Chan, J. 
Steffen-Campbell, and C. Babcock. 2000. Molecu- 
lar systematics of the Chalcidoidea using 28S-D2 
rDNA. Pp. 59-73 in: Austin, A. D., and M. 
Dowton, eds. Hymenoptera: Evolution, Biodiversity 
and Biological Control. CSIRO Publishing, Colling- 
wood. XI + 468 pp. 

Gibson, G. A. P., J. M. Heraty, and J. B. Woolley. 1999. 
Phylogenetics and classification of Chalcidoidea 
and Mymarommatoidea - a review of current 
concepts (Hymenoptera, Apocrita). Zoologica 
Scripta 28: 87-124. 

Gokhman, V. E. 2009. Karyotypes of Parasitic Hymenop- 
tera. Springer Science+Business Media B.V., Dor- 
drecht. XIII + 183 pp. 

Imai, H. T., R. W. Taylor, M. W. J. Crosland, and R. H. 
Crozier. 1988. Modes of spontaneous chromoso- 
mal mutation and karyotype evolution in ants 
with reference to the minimum interaction hy- 
pothesis. Japanese Journal of Genetics 63: 159-185. 

Levan, A., K. Fredga, and A. A. Sandberg. 1964. 
Nomenclature for centromeric position on chro- 
mosomes. Hereditas 52: 201-220. 

Noyes, J. S. 1990. A word on chalcidoid classification. 
Chalcid Forum 13: 6-7. 

Quicke, D. L. J. 1997. Parasitic Wasps. Chapman and 
Hall, London. XVII + 470 pp. 

Rasnitsyn, A. P. 1980. Origin and evolution of 
hymenopterous insects. Trudy Paleontologicheskogo 
Instituta AN SSSR 174: 1-191 (in Russian). 

Rasplus, J.-Y., C. Kerdelhue, I. Le Clainche, and G. 
Mondor. 1998. Molecular phylogeny of fig wasps 
(Hymenoptera). Agaonidae are not monophy- 
letic. Comptes Rendus de V Academic des Sciences de 
Paris III 321: 517-527. 

Vol. 19(1), 2010, pp. 189-198 


Ian David Gauld 
25 May 1947-12 January 2009 

David B. Wahl, Andrew M. R. Bennett, Gavin R. Broad, Ilari E. SaaksjArvi, and 

Ronald J. Zuniga Ramirez 

(DBW) American Entomological Institute, 3005 SW 56*^ Ave., Gainesville, PL 32608-54047, USA; 

d wahl4@cox .net 

(AMRB) Canadian National Collection of Insects, Ottawa, Ontario 

(GRB) Natural History Museum, London, UK 

(lES) University of Turku, Finland 

(RJZR) Instituto Nacional de Biodiversidad, Costa Rica 

The generalities of Ian Gauld's life and 
career have been covered in Hanson et al. 
(2009a,b). He was a hymenopterist, from 
the beginning to the end, and examination 
of his 100 or so life publications (Appendix 
A) shows two major themes: overviews of 
the Hymenoptera and detailed studies of 
Ichneumonidae. The first theme is exem- 
plified by Gauld and Bolton (1988), Hanson 
and Gauld (1995), and Hanson and Gauld 
(2006). Such detailed compendia had never 
before been gathered together for the 
order. Ichneumonids, however, were the 
focus of Gauld's life work, and there are 
three main interests: 1) selected subfami- 
lies, 2) evolutionary biology of the family, 
and 3) faunal studies. 

The Anomalordnae, Labeninae, Ophioni- 
nae, and Pimphnae were of great interest to 
Gauld. A revision of the genera of Anom- 
alordnae was the topic of his Master's thesis 
(Gauld 1976), and he completed several 
faunal treatments of the group. Pimplinae, 
especially with regard to the subfamily's 
diverse biologies, fascinated him, and those 
who did not share his enthusiasm for the 
group were nevertheless subjected to long 
expositions. Labeninae and Ophioninae 

lan's place of burial is in Spratton, 52=19'39.09"N, 
0°57'15.53"W [52.327521 °N, -0.954318°W] 

were the subject of his doctoral dissertation 
i^'A taxonomic study of the Ophioninae and 
Labeninae (Hymenoptera; Ichneumonidae))" ; 
submitted to the Council for National 
Academic Awards, Nov. 1983; The City of 
London Polytechnic and the British Muse- 
um (Natural History)). The labenine section 
was published as Gauld (1983) and the 
ophionine section as Gauld (1985a). His 
interest in Labeninae was continued 
through a series of faunal studies and 
cuhninated in Gauld and Wahl (2000). 

Prior to his work on the Ophioninae, it is 
safe to say that species limits in this 
subfamily were not well understood. Pe- 
rusal of the American Entomological Insti- 
tute collection reveals that even Henry 
Townes did not understand morphological 
variation within the subfamily, nor even 
which characters were essential for species 
delimitation. Gauld's thousands of hours 
of ophionine study 'cracked the code', so to 
speak, and has allowed a much more 
accurate assessment of ophionine diversity. 
This is in addition to his work on ophio- 
nine genera (Gauld 1979; Gauld 1985b), 
which greatly changed the Townesian 

Gauld (1988) was the first discussion of 
ichneumonid biology from an evolutionary 
perspective. This topic captivated him, and 


Journal of Hymenoptera Research 

Table 1. Top ten most prolific ichneumonid 
taxonomists based on number of taxa described. 


Ichneumonid taxa described 





















*will increase to greater than 2150 names following 
posthumous publication. 

it was a focal point of his studies. As 
cladistic methodology was incorporated 
into his research, biology was increasingly 
tied to explicit hypotheses of relationship 
(examples being: Gauld 1983; Gauld and 
Janzen 1994; Wahl and Gauld 1998; Gauld 
and Wahl 2000). 

The regional comprehensive treatment 
of an ichneumonid fauna became an 
organizing principle of his research, start- 
ing with his study of the Australian genera 
(Gauld 1984). He became increasingly 
disillusioned with the monographic treat- 
ment of genera at the World level, seeing 
such studies as useful to only a small 
group of museum workers at major insti- 
tutions (pers. comm. to DBW). The fortu- 
itous combination of factors that led him to 
Costa Rica has been covered elsewhere; the 
last two decades of his life were mostly 
focused upon the Costa Rican fauna. The 
results of this work have been wonderfully 
stimulating to New World ichneumonid 
studies in and out of the tropics, both in 
terms of providing a baseline for other 
countries and encouraging local systematic 

In terms of Gauld's relative impact on 
our knowledge of Ichneumonidae, Table 1 
lists the top ten most prolific ichneumonid 
taxonomists based on the number of taxa 
described. Currently, Gauld lies in fifth 
position with 1,602 (1,516 species, 82 

genera and 4 tribes) (Yu et al. 2008). With 
the planned, posthumous publication of 
Gauld's last work {The Ichneumonidae of 
Costa Rica, 5) which treats the subfamily 
Campopleginae, this number will increase 
to over 2,150 making him the most prolific 
ichneumonid taxonomist in history. 
Whereas the number of taxa described 
does not always equate to the quality of 
the taxonomist, in the case of Ian Gauld, 
both quantity and quality were of the 
highest order. At the time of writing, only 
10 species, 3 genera and 1 tribe of Gauld's 
have been synonymized (less than 1% of aU 
Gauld names). Recent work using DNA 
barcoding on the ophionines of Guana- 
caste, Costa Rica have shown his keen 
sense of morphological species limits to be 
almost always correct Qanzen and collab- 
orators, work in progress). 

Certainly, part of the reason for Gauld's 
exceptionally high level of accuracy can be 
attributed to his encyclopaedic knowledge 
of the world fauna at a species level. Many 
were the times when we would present Ian 
with an enigmatic specimen or potential 
new character and, following the obligato- 
ry stroking of the beard and pensive stare 
into the distance, he would extract the 
relevant information from his prodigious 
memory, thereby allowing us to place the 
specimen or character in its correct context. 
But needless to say, he did not describe 
over 2,000 taxa with simply a good 
memory. Ian was an extremely efficient 
taxonomist and a hard worker. By ''effi- 
cient", we do not mean to imply that his 
descriptions were cursory. On the contrary 
- they are generally model examples of the 
correct balance between brevity and com- 
pleteness. For many taxa (not only ophio- 
nines), he introduced new character sys- 
tems that either reinforced previous hy- 
potheses of relationship or supported new 
ones. Note that it should not be suggested 
that Ian worked entirely by himself. Many 
people contributed to his research, and 
foremost among these were his late wife 
Pam Mitchell (Gauld and Mitchell 1978, 

Volume 19, Number 1, 2010 


1981) and his long-time technician Sondia 
Ward (Ward and Gauld 1987; Gauld 199" 
2000). Perhaps the greatest legac}- of Ian 
Gauld is that he empirically demonstrated 
that the estimate of To^vnes (1969) of 60,000 
species of ichneumonids ^vorld^vide ^vas 
likely much too lo^v. Gauld 's last estimate 
of total species of ichneumonids was 
120,000 (w-^N-^v. This num- 
ber was reached by consideration of 
ichneumonid species richness in tropical 
regions ^vhich To^NTies clearlv underesti- 
mated. In so doiag, Gauld dispelled the 
h}"pothesis that ichneumonid species rich- 
ness decreases ^vith decreasing latitude 
(Owen and Owen 1974; Gauld 1986) and 
reinforced the fact that ichneumonids are a 
m.ajor constituent of biodiversitA" in all 
regions of the ^vorld. 

Da\td B. Wahl 
Andrfw M. R. Bexxett 

As friends, coUeagues and past students 
of lan's (formally or ver\" informallv), ^ve 
would like to take this opportunity' to 
record a few words of appreciation for 
Ian, the old ^vasp taxonomist, as he used to 
sign his emails: 

W^en Ian Gauld passed a^vay we abrupt- 
ly lost a huge stock of ichneumonid ^visdom. 
lan's enthusiasm and kno'wledge ^vere e\'en 
larger than his physical presence and we 
have ^NTitten this obituar\^ as students and 
coUeagues "who gained from this enthusiasm 
and kno^vledge, through formal and infor- 
mal project supen-ision and vei}' pleasant 
collaboration. The Ichneumonidae is a large 
and complex farmly and when starting from 
a ver\' limited kno^vedge-base, to come 
under the ^\ing of Ian was a tremendous 
help. lan's achievements ^vere immense in 
producing taxonomic monographs and in 
pushing f onvards the theme of biodiversit}' 
in the neotropics. Suffice to sav that Ian ^vas 
the ^vorld authorit\' and reall}' opened up 
the study of Ichneumonidae to the next 

Ian was able to s}Tithesise huge amounts 
of information and make it readilv acces- 
sible, something he achieved through 
taxonomic monographs and through intro- 
ductory' texts. Even' working dav we reach 
for Gauld volumes from the shelf; there is 
no higher recommendation of his ^vork 
than that it is essentially practical and 
useful. It is remarkable that for manv 
subfamilies of Ichneumonidae it is no^v 
easier to identih' species from Costa Rica 
than it is from northern Europe, despite 
centuries more effort in describing the 
European fauna. If it ^vasn't for the efforts 
of Ian and his feUo^v Costa Rican pioneers 
we ^vould have ver}' little idea of the 
potential species richness of ichneumonids 
in the neotropics, ^vhere they used to be 
thought of as a rather species-poor group 
compared to their north temperate riches. 

We have aU been out for manv long, 
sHghtiy alcohol-lubricated, dinners ^vith 
Ian. He ^vas al^vays keen to give his time 
to^vards those he considered ^vorth en- 
couraging and ^vas incredibly generous 
when it came to dinners and trips to the 
American Entomological Institute. Infor- 
mation on ichneum.onids, geography, flora 
and fauna, cuisine, all sorts of topics, 
^vould tlo^v forth and ^\'e learnt a lot. 
Ichneumonids ^vere always foremost 

Gamx R. Broad 

We wlQ al^vavs remember Ian as a good 
friend. He was a humble and ^varm- 
hearted man, al^vays ready to encourage 
the research projects of his friends and 
coUeagues. Ian ^vas the world authority' on 
Ichneumonidae but never tried to prove to 
others ho^v good he actuaUy ^vas. In this 
respect, he was a perfect teacher and 
supenisor. He encouraged us to resolve 
taxonomic and systematic problems in our 
^vay and ^vas al^vays there \\'hen support 
^vas needed. 

Ian ^vas one of fe^v ichneumonologists 
capable of dealing with some taxonomicaUy 


Journal of Hymenoptera Research 

Fig. 1. Ian Gauld at the Natural History Museum in 1975 or 1976 and at his home in 2005 (R. Zufiiga). 

extremely difficult subfamilies - a true star 
in the world of parasitoid Hymenoptera 
research. He, for example, supervised his 
younger colleagues with a smile in his face 
by saying that "nobody can escape from the 
Campopleginae for long!", and attacked 
this subfamily, avoided by most of us, with 
all his enthusiasm. lan's positive attitude 
towards life and ichneumonid taxonomy 
and systematics is something that we will 
always remember and miss. 

In taxonomy, new species are often 
named in honour of distinguished re- 
searchers of the field. For example, at least 
13 species and one genus of ichneumonoid 
and one species of chalcidoid have been 
named after Ian. In contrast, Ian named a 
large number of species after people, 
whether they were academics or not, who 
had helped somehow, e.g. in the field 
inventories of Costa Rica. As a conse- 
quence, from lan's books one can find 
species named after people who helped Ian 
with Spanish language tuition or joined the 
first biodiversity training course for the 
national parks of Costa Rica. It was lan's 
way of thanking the people he had met 
during the years. 

He loved to spend time with his friends 
and readily treated these nights to his 

younger colleagues. The dinners were 
always finished with sweet desserts, which 
were his favourite. We all learned to know, 
for examples, creme brule and suspiro limena, 
the latter being a very sweet and delicious 
Peruvian dessert. Ian took his friends and 
colleagues often to his home, in the 
beautiful village of Spratton in the English 
countryside close to the city of North- 
ampton. It was always nice to stay with Ian 
and his wife Pam. They were a lovely and 
loving couple, always taking good care of 
their friends. Many of us have spent some 
truly nice days at their house studying 
ichneumonids in lan's fully equipped 
home office, sharing lovely meals prepared 
by Pam and excellent wines which were 
appreciated by both of them. In their way, 
Ian and Pam were like Henry and Marjorie 
Townes - the founders of the American 
Entomological Institute. Pam often joined 
Ian in his research and they provided a 
lovely environment for visiting colleagues. 
Pam died less than one year before Ian. It 
was a hard time for Ian and he wrote us 
"Thank goodness I still have my work". 
Their marriage lasted 35 happy years and 
they leave behind lan's son, Darren. We all 
remember Ian and Pam with love and 
affection, and with many shared years, we 

Volume 19, Number 1, 2010 


will always have a treasure trove of good 

Ilari E. Saaksjarvi 

Ian Gauld was a person full of charisma 
in entomology, his passion and dedication 
to his work led him to make great 
discoveries. He was simple and humble, 
willing to listen to anyone who had an idea 
to tell. Ian was a great teacher, but more a 
great friend, often giving advice to im- 
prove as individuals and as professionals. 

lan's way of working is an example to 
follow, with enthusiasm and joy, a lover of 
good food, talks and desserts. In Costa 
Rica, he spent many years teaching and 
working in difficult places, but always 
with a tremendous joy for what he did. 
Pam, his wife, at his side helping him in his 
huge task of preparing the material, that 
Ian would later describe. He is remem- 
bered as the great teacher and great friend; 
our obligation is to follow in his footsteps 
and continue working the way he would 
have wanted to. Working day to day is the 
best way to remember him. 

We can write hundreds of anecdotes 
about Ian, aU filled with joy; his house was 
a university classroom, he was a man who 
never failed to surprise us, who lived as he 
wanted, enjoying all he did. 

Ronald J. Zuniga 


Gauld, I. D. 1976. The classification of the Anomalo- 
ninae (Hymen op tera: Ichneumonidae). BuUeti?! of 
the British Museum (Natural Histon/) (Entomology) 
33: 1-135. 

. 1979 (1980). An analysis of the classification of 

the Ophion genus-group (Hymenoptera: Ichneu- 
monidae). Systematic Entomology 5: 59-82. 

. 1983. The classification, evolution and distri- 
bution of the Labeninae, an ancient southern 
group of Ichneumonidae (H\Tnenoptera). System- 
atic Entomology 8: 167-178. 

. 1984. An Introduction to the Ichneumonidae of 

Australia. British Museum (Natural Histor\0: 
London. 413 pp. 

. 1985a. The phylogeny, classification and 

evolution of parasitic wasps of the subfamily 

Ophioninae (Ichneumonidae). Bulletin of the Brit- 
ish Museum (Natural History) (Entomology) 51: 

— . 1985b. A preliminary surv-ey of the Ophioni- 
nae (Hymenoptera; Ichneumonidae) of Brunei. 
Brunei Museum Journal 6: 169-188. 

— . 1986. Latitudinal gradients in ichneumonid 
species-richness in Australia. Ecological Entomolo- 
gy 11: 155-161. 

— . 1988. Evolutionary' patterns of host utilization 
by ichneumonoid parasitoids (Hymenoptera: 
Ichneumonidae and Braconidae). Biological Jour- 
nal of the Linnean Society 35: 351-377. 

— . 1997. The Ichneumonidae of Costa Rica, 2. 
Memoirs of the American Entomological Institute 57: 

— . 2000. The Ichneumonidae of Costa Rica, 3. 

Memoirs of tlie American Entomological Institute 63: 

Gauld, I. D., and B. Bolton, eds. 1988. Vie Hymenoptera. 
British Museum (Natural Piistory)/ Oxford Uni- 
versity Press: London. 332 pp. 

Gauld, I. D. and D. H. Janzen. 1994. The classification, 
evolution and biology of the Costa Rican species 
of Cryptophion (Hymenoptera: Ichneumonidae). 
Zoological Journal of the Linnean Society 110: 

and P. A. Mitchell. 1978. The taxonomy, 

distribution and host preferences of African parasitic 
wasps of the subfamily Ophioninae (Hymenoptera: 
Ichneumonidae). Commonwealth Agricultural Bu- 
reaux: Slough, United Kingdom. 287 pp. 

and P. A. Mitchell. 1981. The taxonomy, 

distribution and host preferences of Indo-Papuan 
parasitic wasps of the subfamily (yphioninae (Hyme- 
noptera: Ichneumonidae). Commonwealth Agricul- 
tural Bureaux: Slough, United Kingdom. 611 pp. 

and D. B. Wahl. 2000. The Labeninae (Hvme- 

noptera: Ichneumonidae): a study in phylogenetic 
reconstruction and evolutionary biolog)'. Zoolog- 
ical Journal of the Linnean Society 129: 271-347. 

Hanson, P. E., and I. D. Gauld, eds. 1995. The 
Hymenoptera of Costa Rica. Oxford University 
Press: Oxford. 893 pp. 

Hanson, P. E., and I. D. Gauld, eds. 2006. Hymenop- 
tera de la region neotropical. Memoirs of the 
American Entomological Institute 77: 1-994. 

Hanson, P., Coronado, J., Ugalde, J., Godoy, C, and 
Zuniga, R. 2009a. Ian David Gauld (1947-2009). 
Su legado a la biodiversidad entomologica de 
Costa Rica y el mundo. Revista de Biologia Tropical 
57 (Suppl. 1): XXV-XXX. 

, Janzen, D., Knapp, S., Scoble, M., Wahl, D., 

and West-Eberhard, M. J. 2009b. Ian Gauld FLS 
(1947-2009). Master taxonomist and morpholo- 
gist. The Linnaean 25 (2): 46-47. 

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


Journal of Hymenoptera Research 

Townes, H. K. 1969. The genera of Ichneumonidae, 
Part 1. Memoirs of the American Entomological 
Institute 11: 1-300. 

Wahl, D. B. and I. D. Gauld. 1998. The cladistics and 
higher classification of the Pimpliformes (Hyme- 
noptera: Ichneumonidae). Systematic Entomology 
23: 265-298. 

Ward, S. and I. D. Gauld. 1987. The callajoppine 
parasitoids of sphingids in Central America 
(Hymenoptera: Ichneumonidae). Systematic Ento- 
mology 12: 503-508. 

Yu, D., van Achterberg, K., and Horstmann, K. 2008. 
World Ichneumonoidea 2004. Taxonomy, biology, 
morphology and distribution. [CD/DVD]. Taxa- 
pad®, Vancouver, Canada. [Available for sale at: 
http : / / ww^v^ .taxapad .com] . 

Appendix A: Complete Publication List 
of Ian D. Gauld 

Bartlett, R., J. Pickering, L Gauld, and D. 
Windsor. 1999. Estimating global biodiversi- 
ty: tropical beetles and w^asps send different 
signals. Ecological Entomology 24: 118-121. 

Billany, D. J., T. J. Winter, and L D. Gauld, 1985. 
Olesicampe monticola (Hedw^ig) (Hymenop- 
tera: Ichneumonidae) redescribed together 
w^ith notes on its biology as a parasite of 
Cephalcia lariciphila (Wachtl) (Hymenoptera: 
Pamphiliidae). Bulletin of Entomological Re- 
search 75: 267-274. 

Coronado-Rivera, J., A. Gonzalez-Herrera, I. D. 
Gauld, and P. Hanson, P. 2004. The enigmatic 
biology of the ichneumonid subfamily Lycor- 
ininae. Journal of Hymenoptera Research 13: 

Danthanarayana, W., D. Farrugia, and I. D. 
Gauld. 1977. Studies on the biology and 
systematic position of a new species of 
ichneumonid parasitising the light brown 
apple moth, Epiphyas postvittana (Walker) 
(Lepidoptera: Tortricidae), in Australia. Bul- 
letin of Entomological Research 67: 607-617. 

Fitton, M. G. and I. D. Gauld. 1976. The family- 
group names of Ichneumonidae (excluding 
the Ichneumoninae). Systematic Entomology 1: 

Fitton, M. G. and I. D. Gauld. 1978. Further 
notes on family-group names of Ichneumo- 
nidae (Hymenoptera). Systematic Entomology 
3: 245-247. 

Fitton, M. G. and I. D. Gauld. 1980. A review of 
the British Cremastinae (Hymenoptera: Ich- 
neumonidae) with keys to the species. Ento- 
mologist's Gazette 31: 63-71. 

Fitton, M. G. and I. D. Gauld. 1991. [Comments 
on the proposed conservation of Cryptus 
Fabricius, 1804 (Insecta, Hymenoptera)]. Bul- 
letin of Zoological Nomenclature 48: 326-327. 

Fitton, M. G., I. D. Gauld, L. I. N. Roberts, and 
A. K. Walker. 1983. An African ichneumonid 
(Hymenoptera) in Australasia. Bulletin of 
Entomological Research 73: 465-468. 

Fitton, M. G., I. D. Gauld, and M. R. Shaw. 1982. 
The taxonomy and biology of the British 
Adelognathinae (Hymenoptera: Ichneumoni- 
dae). Journal of Natural History 16: 275-283. 

Fitton, M. G., M. R. Shaw, and I. D. Gauld. 1988. 
Pimpline ichneumon-flies (Hymenoptera, 
Ichneumonidae, Pimplinae). Handbook for the 
identification of British insects 7(1): 1-110. 

Gamez, R. L. and I. D. Gauld. 1993. Costa Rica: 
an innovative approach to the study of 
tropical biodiversity. In: LaSalle, J. and I. D. 
Gauld, eds. Hymenoptera and Biodiversity. CAB 
International: Wallingford, UK. 348 pp. 

Gaston, K. J. and I. D. Gauld. 1993. How many 
species of pimplines (Hymenoptera: Ichneu- 
monidae) are there in Costa Rica? Journal of 
Tropical Ecology 9: 491-499. 

Gauld, I. D. 1970. Some records of Ichneumo- 
noidea (Hymenoptera) collected at light, 
during autumn 1969 in Perthshire and south- 
ern Inverness-shire, including a species new 
to Britain. Entomologist's Gazette 21: 282-284. 

Gauld, I. D. 1973a. Notes on the British 
Ophionini (Hymenoptera: Ichneumonidae) 
including a key to species. Entomologist's 
Gazette 24: 55-66. 

Gauld, I. D. 1973b. Notes on some Anomaloni- 
nae (Hymenoptera: Ichneumonidae) on Can- 
nock Chase during Augiast 1973. Entomolo- 
gist's Gazette 24: 304. 

Gauld, I. D. 1974a. Further notes on the British 
Ophionini (Hymenoptera: Ichneumonidae). 
Entomologist's Gazette 25: 147-148. 

Gauld, I. D. 1974b. Notes on the species of 
Heteropelma Wesmael (Hymenoptera: Ichneu- 
monidae) occurring in Australia. Bulletin of 
Entomological Research 64: 541-544. 

Gauld, I. D. 1976a. The classification of the 
Anomaloninae (Hymenoptera: Ichneumoni- 
dae). Bulletin of the British Museum (Natural 
History) (Entomology) 33: 1-135. 

Gauld, I. D. 1976b. A revision of the Anom- 
aloninae (Hymenoptera: Ichneumonidae) of 
Australia. Australian Journal of Zoology 24: 

VoLL-ME 19, Nl-mber 1, 2010 


Gauld, I. D. 1976c. Notes on the British 
Ophioninae (Hymenoptera: Ichneumonidae). 
Part 3. The identit}^ of the species described 
by Morley, 1915 and Thomson, 1888 and a 
checklist of British species. Eiitouiologist's 
Gazette 27: 113-117. 

Gauld, I. D. 1976d. The taxonomy of the genus 
Heteropebna Wesmael (Hymenoptera: Ichneu- 
monidae). Bulletin of the British Museum 
(Natural History) (Entomology) 34: 153-219. 

Gauld, I. D. 1976e. Notes on the species of 
Triclwmma Wesmael (H^inenoptera: Ichneu- 
monidae) occurring in the Indian subcontinent. 
Bulletin of Entomological Research 65: 643-649. 

Gauld, I. D. 1976f. Description of a ne^v genus of 
Ichneumonidae (Hymenoptera) of economic 
importance from Sabah. Bulletin of Entomo- 
logical Research 66: 1-4. 

Gauld, I. D. 1977. A revision of the Ophioninae 
(Hymenoptera: Ichneumonidae) of Australia. 
Australian journal of Zoology ( Supplementanj 
Series) 49: 1-112. 

Gauld, I. D. 1978a. Notes on the British 
C)phiordnae (Hymenoptera: Ichneumonidae). 
Part 4. A re\ised key to species of the genus 
Ophion Fabricius. Entomologist's Gazette 29: 

Gauld, I. D. 1978b. A revision of the Ethiopian 
species of Agrypon Foerster (H\Tnenoptera: 
Ichneumonidae). Joimial ofXatural History 12: 

Gauld, I. D. 1978c. A re\ision of the Anomalo- 
ninae (Hymenoptera: Ichneumonidae) of 
Melanesia I. The genera Anomalon Panzer to 
Aphanistes Foerster. Bulletin of Entomological 
Research 68: 501-519. 

Gauld, I. D. 1978d. A revision of the .Ajiom- 
aloninae (H}Tnenoptera: Ichneumonidae) of 
of Melanesia 11. The genera Perisphincter 
Townes and Agrypon Foerster. Bulletin of 
Entomological Research 68: 543-557. 

Gauld, I. D. 1979 (1980). An analysis of the 
classification of the Ophion genus-group 
(Hymenoptera: Ichneumonidae). Systematic 
Entomology 5: 59-82. 

Gauld, I. D. 1980a. A synopsis of the .Anom- 
aloninae (H}Tnenoptera: Ichneumonidae) of 
sub-Sahaxan Africa with special reference to 
species of potential economic importance. 
Bulletin of Entomological Research 70: 33-41. 

Gauld, I. D. 1980b. Notes on an economically 
important species of Temelucha Foerster (Hy- 
menoptera: Ichneumonidae) and a prelimi- 

nary key to Australian species. Bulletin of 
Entomological Research 70: 43-47. 

Gauld, I. D. 1980c. Notes on the Ne\v Zealand 
Anomaloninae (Hymenoptera: Ichneumoni- 
dae) with description of a new species of 
Aphanistes Foerster of possible economic 
importance in forestr\\ Nnv Zealand Entomol- 
ogist 7: 130-134. 

Gauld, I. D. 1982. A revised key to the 
Enicospilus antefurcalis (Szepligeti) (Hyme- 
noptera: Ichneumonidae) species-group of 
the Afrotropical region. Bulletin of Entomolog- 
ical Research 72: 33-38. 

Gauld, I. D. 1983a (1982). Netelia species of the 
subgenera Apatagium Enderlein and Mono- 
macrodon Cushman (Hymenoptera: Ichneu- 
monidae) of Brunei with a re-assessment of 
the supraspecific classification. Brunei Muse- 
um Journal 5(2): 123-143. 

Gauld, I. D. 1983b. The classification, evolution 
and distribution of the Labeninae, an ancient 
southern group of Ichneumonidae (H\Tne- 
noptera). Systematic Entomology 8: 167-178. 

Gauld, I. D. 1984a. The Australian Ophioninae 
(Insecta; Hymenoptera): a historical biogeo- 
graphic studv. journal of Biogeography 11: 269- 

Gauld, I. D. 1984b. An Introduction to the 
Ichneumonidae of Australia. British Museum 
(Natural Histor\'): London. 413 pp. 

Gauld, I. D. 1984c. The Pimplinae, Xoridinae, 
Acaenitinae and Lycorininae (H}Tnenoptera: 
Ichneumonidae) of Australia. Bulletin of the 
British Museum (Natural History) (Entomology) 
49: 235-339. 

Gauld, I. D. 1985a. The phylogeny, classification 
and evolution of parasitic wasps of the 
subfamily Ophioninae (Ichneumonidae). Bul- 
letin of the British Museum (Natural History) 
(Entomology) 51: 61-185. 

Gauld, I. D. 1985b. A preliminary- sur\'ey of the 
Ophioninae (Hymenoptera; Ichneumonidae) 
of Brunei. Brunei Museum Journal 6: 169-188. 

Gauld, I. D. 1986a. Latitudinal gradients in 
ichneumonid species-richness in Australia. 
Ecological Entomology 11: 155-161. 

Gauld, I. D. 1986b. Taxonomy, its Kmitations 
and its role in understanding parasitoid 
biolog}^ In: VVaage, J. and D. Greathead, 
eds. Insect Parasitoids. Academic Press: Lon- 
don. 389 pp. 

Gauld, I. D. 1987a. Some factors affecting the 
composition of tropical ichneumonid faunas. 


Journal of Hymenoptera Research 

Biological Journal of the Linnean Society 30: 299- 

Gauld, I. D. 1987b. Chapter 5: Ichneumonidae. 
The ichneumonid parasitoids associated with 
South-east Asian Limacodidae. In: Cock, M. J. 
W., H. C. J. Godfray, and J. D.Holloway, eds. 
Slug and nettle caterpillars: the biology, taxonomy 
and control of the Limacodidae of economic 
importance on palms in South-east Asia. Com- 
monwealth Agricultural Bureaux: WaUing- 
ford. United Kingdom. 270 pp. 

Gauld, I. D. 1988a. The species of the Enicospilus 
americanus (Hymenoptera: Ichneumonidae) 
complex in eastern North America. Systematic 
Entomology 13: 31-53. 

Gauld, I. D. 1988b. A sur\'ey of the Ophioninae 
(Hymenoptera: Ichneumonidae) of tropical 
Mesoamerica with special reference to the faima 
of Costa Rica. Bulletin of the British Museum 
(Natural History)(Entomology) 57: 1-309. 

Gauld, I. D. 1988c. Evolutionary patterns of host 
utilization by ichneumonoid parasitoids (Hy- 
menoptera: Ichneumonidae and Braconidae). 
Biological Journal of the Linnean Society 35: 351- 

Gauld, I. 1991. The Ichneumonidae of Costa 
Rica, 1. Memoirs of the American Entomological 
Institute 47: 1-589. 

Gauld, I. D. 1996. The cost of data repatriation 
\'ersus the potential of biosystematic capacity 
building. In: McNeely, J. A. and Y. S. 
Soucherita, eds. Biodiversity in Asia: Challenges 
and Opportunities for the Scientific Community. 
Ministry of Science, Technology and Envi- 
ronment: Bangkok. 

Gauld, I. D. 1997. The Ichneumonidae of Costa 
Rica, 2. Memoirs of the American Entomological 
Institute 57: 1^85. 

Gauld, I. D. 2000a. The re-definition of pimpline 
genus Hymenoepimecis (Hymenoptera: Ich- 
neumonidae) with a description of a plesio- 
morphic new Costa Rican species. Journal of 
Hymenoptera Research 9: 213-219. 

Gauld, I. D. 2000b. The Ichneumonidae of Costa 
Rica, 3. Memoirs of the American Entomological 
Institute 63: 1^53. 

Gauld, I. D., ed. 2002. The Ichneumonidae of 
Costa Rica, 4. Memoirs of the American 
Entomological Institute 66: 1-768. 

Gauld, I. D. and B. Bolton, eds. 1988. The 
Hymenoptera. British Museum (Natural His- 
tory) /Oxford University Press: London. 332 

Gauld, I. D. and J.M. Carter. 1983. The Ophio- 
ninae of the Galapagos Islands (Hymenop- 
tera: Ichneumonidae). Journal of Natural His- 
tory 17: 145-155. 

Gauld, I. D., N. M. CoUins, and M. G. Fitton. 1990. 
The biological significance and conser\'ation of 
Hymenoptera in Europe. Nature and Environ- 
ment Series (Coimcil of Europe) 44: 1^7. 

Gauld, I. D. and ]. Dubois. 2006. Phylogeny of 
the Polysphincta group of genera (Hymenop- 
tera: Ichneumonidae; Pimplinae): a taxonom- 
ic revision of spider ectoparasitoids. System- 
atic Entomology 31: 529-564. 

Gauld, I. D. and M. G. Fitton. 1980 (1979). The 
British species of Phrudinae (Hymenoptera: 
Ichneumonidae) Entomologist's Monthly Mag- 
azine 115: 197-199. 

Gauld, I. D. and M. G. Fitton. 1981. Keys to the 
British xoridine parasitoids of wood-boring 
beetles (Hymenoptera: Ichneumonidae). En- 
tomologist's Gazette 32: 259-267. 

Gauld, I. D. and M. G. Fitton. 1987. Sexual 
dimorphism in Ichneumonidae: a response to 
Hurlbutt. Biological Journal of the Linnean 
Society 31: 291-300. 

Gauld, I. D. and K. J. Gaston. 1994. The taste of 
enemy-free space: parasitoids and nasty 
hosts. In: Hawkins, B.A. and W. Sheehan, 
eds. Parasitoid Community Ecology. Oxford 
University Press: Oxford. 516 pp. 

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

Gauld, I. D. and G. A. HoUoway. 1983. A new 
genus of endaseine Ichneumonidae from 
Australia (Hymenoptera). Contributions of the 
American Entomological Institute 20: 191-197. 

Gauld, I. D. and G. A. Holloway. 1985. 
Australian ichneumonids of the tribes Labe- 
nini and Poecilocryptini. Bulletin of the British 
Museum (Natural History) Entomology 53: 107- 

Gauld, I. D. and T. Huddleston. 1976. The 
Nocturnal Ichneumonoidea of the British 
Isles, including a key to genera. Entomologist's 
Gazette 27: 35-49. 

Gauld, I. D. and D. H. Janzen. 1994. The 
classification, evolution and biology of the 
Costa Rican species of Cryptophion (Hyme- 
noptera: Ichneumonidae). Zoological Journal of 
the Linnean Society 110: 297-324. 

Volume 19, Number 1, 2010 


Gauld, I. D. and D. H. Janzen. 2004. The 
systematic and biology of the Costa Rican 
species of parasitic wasps in the Thyreodon 
genus-group (Hymenoptera: Ichneumoni- 
dae). Zoological Journal of the Linnean Society 
141: 297-351. 

Gauld, I. D. and D. Lanfranco. 1988. Los generos 
de Ophioninae de Centro y Sudamerica. 
Revista Biologia Tropical 35: 257-267. 

Gauld, I. D. and J. LaSalle. 1992. Parasitic 
Hymenoptera and the Biodiversity Crisis. 
Redia 74: 315-334. 

Gauld, I. D., R. Menjivar, M. O. Gonzalez, and 
A. Monro. 2002. Guia para la identificacion de 
los Pimplinae de cafetales bajo sombra de El 
Salvador (Hymenoptera: Ichneumonidae) . Natu- 
ral History Museum: London. 76 pp. 

Gauld, I. D. & P. A. Mitchell. 1976. The 
ichneumonid genus Pseudanomalon (Hyme- 
noptera). Systematic Entomology 1: 117-122. 

Gauld, I. D. and P. A. Mitchell. 1977. Nocturnal 
Ichneumonidae of the British Isles: the genus 
Alexeter Foerster. Entomologist's Gazette 28: 51- 

Gauld, I. D. and P. A. Mitchell. 1978. The 
taxonomy, distribution and host preferences of 
African parasitic wasps of the subfamily Ophio- 
ninae (Hymenoptera: Ichneumonidae). Common- 
wealth Agricultural Bureaux: Slough, United 
Kingdom. 287 pp. 

Gauld, I. D. and P. A. Mitchell. 1981. The 
taxonomy, distribution and host preferences of 
Indo-Papuan parasitic wasps of the subfamily 
Ophioninae (Hymenoptera: Ichneumonidae). Com- 
monwealth Agricultural Bureaux: Slough, 
United Kingdom. 611 pp. 

Gauld, I. D. and L. A. Mound. 1982. Homoplasy 
and the delineation of holophyletic groups. 
Systematic Entomology 7: 73-86. 

Gauld, I. D., M. A. O'Neill, and K. J. Gaston. 
2000. Driving Miss Daisy: the performance of 
an automated insect identification system. In: 
Austin, A. D. and M. Dowton. Hymenoptera: 
Evolution, Biodiversity and Biological Control. 
CSIRO: Collingwood, Australia. 468 pp. 

Gauld, I. D., J. A. Ugalde, and P. Hanson. 1998. 
Guia de los Pimplinae de Costa Rica (Hyme- 
noptera: Ichneumonidae). Revisita de Biologia 
Tropical 46 (Supplement 1): 1-189. 

Gauld, I. D. & G. Underwood. 1986. Some 
applications of the LeQuesne compatibility 
test. Biological Journal of the Linnean Society 29: 

Gauld, I. D. and D. B. Wahl. 2000. The 
Labeninae (Hymenoptera: Ichneumonidae): 
a study in phylogenetic reconstruction and 
evolutionary biology. Zoological Journal of the 
Linnean Society 129: 271-347. 

Gauld, I. D. and D. B. Wahl. 2001. The Town- 
esioninae: a distinct subfamily of Ichneumo- 
nidae (Hymenoptera) or a clade of the 
Banchinae? Transactions of the American Ento- 
mological Society 126: 279-292. 

Gauld, I. D. and D. B. Wahl. 2002. The 
Eucerotinae: a Gondwanan origin for a 
cosmopolitan group of Ichneumonidae? Jour- 
nal of Natural History 36: 2229-2248. 

Gauld, I. D. and D. B. Wahl. 2006. The 
relationship and taxonomic position of the 
genera Apolophus and Scolomus (Hymenop- 
tera: Ichneumonidae). Zootaxa 1130: 35-61. 

Gauld, I. D., D. B. Wahl, and G. R. Broad. 2002. 
The suprageneric groups of the Pimplinae 
(Hymenoptera: Ichneumonidae): a cladistic 
re-evaluation and evolutionary biological 
study. Zoological Journal of the Linnean Society 
136: 421-485. 

Gauld, I. D. and R. Zuniga. 2003. Apechoneura 
naturaeconservatio Gauld & Zufiiga. INBio: 
San Jose, Costa Rica. [Note: a small card 
proposing a species without a formal de- 
scription, hence a nomen nudum] 

Hanson, P. E. and I.D. Gauld, eds. 1995. The 
Hymenoptera of Costa Rica. Oxford University 
Press: Oxford. 893 pp. 

Hanson, P. E. and I. D. Gauld, eds. 2006. 
Hymenoptera de la region neotropical. Mem- 
oirs of the American Entomological Institute 77: 

Huddleston, T. and I. D. Gauld. 1988. Parasitic 
wasps (Ichneumonoidea) in British light- 
traps. Entomologist 107: 134-154. 

Janzen, D. H., A. K. Walker, J. B. Whitfield, G. 
Delvare, and I. D. Gauld. 2003. Host-specific- 
ity and hyperparasitoids of three new Costa 
Rican species of Microplitis Foerster (Hyme- 
noptera: Braconidae: Microgastrinae), para- 
sitoids of sphingid caterpillars. Journal of 
Hymenoptera Research 12: 42-76. 

Jerman, E. J. and I. D. Gauld. 1988. Casinaria, a 
paraphyletic ichneumonid genus (Hymenop- 
tera), and a revision of the Australian species. 
Journal of Natural History 11: 589-609. 

LaSalle, J. and I. D. Gauld, eds. 1993a. Hyme- 
noptera and Biodiversity. CAB International: 
Wallingford, UK. 348 pp. 


Journal of Hymenoptera Research 

LaSalle, J. and I. D. Gauld. 1993b. Hymenoptera: 
their diversity and their impact on the 
diversity of other organisms. In: LaSalle, J. & 
I. D. Gauld, eds. Hymenoptera and Biodiversity. 
CAB International: Wallingford, UK. pp. 1-26. 

Memmott, J., H. C. J. Godfray, and I. D. Gauld. 
1994. The structure of a tropical host-parasit- 
oid community. Journal of Animal Ecology 63: 

Saaksjarvi, I. E., S. Haataja, S. Neuvonen, I. D. 
Gauld, R. Jussila, J. Salo, and A. M. Burgos. 
2004. High local species richness of parasitic 
wasps (Hymenoptera: Ichneumonidae; Pim- 
plinae and Rhyssinae) from the lowland 
rainforests of Peruvian Amazonia. Ecological 
Entomology 29: 735-743 

Saaksjarvi, I. E., E. Palacio, I. D. Gauld, R. 
Jussila, and J. Salo. 2003. A new genus and six 
new species of the tropical Camptotypus 
genus-group (Hymenoptera: Ichneumonidae; 
Pimplinae) from northern South America. 
Zootaxa 197:1-18. 

Saaksjarvi, I. E., I. D. Gauld, and J. Salo. 2004: 
Phylogenetic evaluation of the tropical Camp- 
totypus genus-group (Hymenoptera: Ichneu- 

monidae), with a key to the world genera. 
Journal of Natural History 38: 2759-2778. 

Wahl, D. B. and I. D. Gauld. 1998. The cladistics 
and higher classification of the Pimpliformes 
(Hymenoptera: Ichneumonidae). Systematic 
Entomology 23: 265-298. 

Ward, S. and I. D. Gauld. 1987. The callajoppine 
parasitoids of sphingids in Central America 
(Hymenoptera: Ichneumonidae). Systematic 
Entomology 12: 503-508. 

Weeks, P. J. D., I. D. Gauld, K. J. Gaston, and M. 
A. O'Neill. 1997. Automating the identifica- 
tion of insects: a new solution to an old 
problem. Bulletin of Entomological Research 87: 

Weeks, P. J. D, M. A. O'Neill, K. J. Gaston, and I. 
D. Gauld. 1999a. Species-identification of 
wasps using principal component associative 
memories. Image and Vision Computing 17: 

Weeks, P. J. D., M. A. O'Neill, K. J. Gaston, and 
I. D. Gauld. 1999b. Automating insect iden- 
tification: exploring the limitations of a 
prototype system. Journal of Applied Entomol- 
ogy 123: 1-8. 

Vol. 19(1), 2010, p. 199 


This is my final issue as editor of Joimial 
of Hymenoptera Research. There has been 
much discussion within the societ}' re- 
cently about the direction that Journal of 
Hymenoptera Research should take and we 
now have a good idea of ho^v to adapt the 
journal. Ho^vever, I think it is best that a 
fresh pair of hands guides our journal 
through these changes. We are still short of 
copy but we are confident that Journal of 
Hymenoptera Research has a bright future 
and I hope that, as we change our format, 
more members will submit their manu- 
scripts here. 

I would like to take this opportunit}^ to 
record my thanks to ever^'body ^vho made 
this job much easier than it could have 
been, and often even enjoyable. We depend 
on the many reviewers who almost always 
say 'yes' when approached and make the 
journal what it is. It is ver\' pleasing that so 
many of you continue to devote some of 
your increasingly stretched time to review- 
ing. The subject editors, of course, have 
been unstintingly generous in devoting 

their time and efforts towards the journal, 
ver}^ frequently going out of their way to 
improve manuscripts that have merit but 
were not quite there yet, and I have relied 
upon them much more than I would 
normally care to admit. 

One of the more gratif}'ing aspects of 
editing JHR has been learning just how 
frequently hymenopterists are willing to 
devote considerable time and effort to- 
wards improving promising manuscripts 
and providing constructive criticism. 
Another boon of the job has been corre- 
sponding with so many hymenopterists 
around the world. I've also learned that 
almost nobody can format a manuscript 
precisely in accordance ^vith the instruc- 
tions. It has certainly made me a better 

Gavin Broad 

Department of Entomolog}', the Xatural 
Histor}' Museum, Cromwell Road, Lon- 
don SW7 5BD, UK, 


General Policy. The Journal of Hymenoptem 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 acceptable if the species has 
economic importance or provides new data on the biology or evolution of the genus or higher taxon. 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. This journal is ISI-listed. 

The deadline for receipt of manuscripts is 1 September (for the April issue) and 1 March (for the October 

Format and Preparation. Authors are strongly encouraged to submit manuscripts electronically to the editor 
at the email address below, and in the format specified below. On the upper left of the title page give name, 
address, telephone and fax rtumbers, and email address 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) 

Upon final acceptance of a manuscript, the author should provide the editor with an emailed IBM formatted 
electronic version. CD-ROMs or 3.5 inch floppy disks are acceptable. Because symbols and tables are not always 
correctly translated it is best to also send a printed copy of the manuscript. Preferred word processing programs 
are Microsoft Word and WordPerfect. If possible, all words that must be italicized should be done so, not 
underscored. 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. CDs and 
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 (14 x 20.5 cm, column 
width 6.7 mm) and allow room for legends at the top and bottom. Do not make plates larger than 14 x 18 in. (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. 
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-l)" "(Table 1)". Measurements should be in the metric system. 

Electronic plates may be submitted on disc, via email or uploaded to an ftp site (instructions will be given). 
They must be fully composited, labeled, and sized to fit the proportions of the journal page. Line art should be 
scanned at 1200 dpi (minimum input resolution is 600 dpi). Color or grayscale (halftone) images should have a 
dpi of 300-350. Color files should be in CMYK and not RGB. Graphics should be submitted as TIFF, Adobe 
Illustrator or EPS files. No PowerPoint or Word /WordPerfect files with images embedded in them are 

All papers must conform to the International Code of Zoological Nomenclature. The first mention of a plant or 
animal name 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 recognized institutions (not 
private collections). Voucher specimens should be designated for specimens used in behavioral or autecological 
studies, and they should be deposited similarly DNA sequences must be deposited in GenBank/EMBL/DNA 
Databank of Japan. 

Acceptance of taxonomic papers will not require use of cladistic methods; however, authors using them will 
be expected to specify the phylogenetic program used, including discussion of program options used. A data 
matrix should be provided for morphological characters. Cladograms must be hung with characters and these 
should include descriptors (not numbers alone) when feasible. The number of parsimonious cladograms 
generated should be stated and reasons given for the one adopted. Lengths and consistency indices should be 
provided. Adequate discussions should be given for characters, plesiomorphic conditions, and distributions of 
characters among outgroups when problematical. 

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 multiple authors, use the word "and," not the 
symbol "&" (Smith and Jones 1999). 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 and in italics. 

Charges. Publication charges are $10.00 per printed page. At least one author of the paper must be a member 
of the International Society of Hymenopterists. 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 addressed to: 

Dr Gavin Broad 

Dept. of Entomology 

The Natural History Museum 

Cromwell Road 

London SW7 5BD, UK 

Phone: +44(0)207 9425938; Fax: +44(0)207 9425229; Email: 


3 9088 01553 1874 

(Continued from front cover) 

DIEZ, P., A. ROIG-ALSINA, and P. FIDALGO. Phylogenetic relationships of Pluroides por- 
teri, a new genus and species of Plumariidae from Argentina (Hymenoptera: 
Chrysidoidea) 113 

JAPOSHVILI, G. and I. KARACA. The Transcaucasian species of Coccobius Ratzeburg 1852 (Chal- 

cidoidea: Aphelinidae), with the description of three new species from Georgia 121 

PETERS, R. S. Host range and offspring quantities in natural populations of Nasonia vitripennis 

(Walker, 1836) (Hymenoptera: Chalcidoidea: Pteromalidae) 128 

PULAWSKI, W. J. Rhopalum nasale, a new species from Australia (Hymenoptera: Crabroni- 

dae) 139 

RICHARDS, M. H., J. L .VICKRUCK, and S. M. REHAN. Colony social organisation of Halictus 
confiisus in southern Ontario, with comments on sociality in the subgenus H. {Sela- 
donia) 144 

ROMERO, C. A. and J. M. HERATY. Ultrastructure of scutellar sensilla in Aphytis melinus (Hy- 
menoptera: Aphelinidae) and morphological variation across Chalcidoidea 159 

J. YU, M. WANG and G. E. HEIMPEL. A new species of Lysiphlebus Forster 1862 (Hy- 
menoptera: Braconidae, Aphidiinae) attacking soybean aphid. Aphis glycines Mat- 
sumura (Hemiptera: Aphididae) from China 179 


GOKHMAN, V. E., A. P MIKHAILENKO and V. N. FURSOV. Chromosomes of Blastophaga 

psenes (Hymenoptera: Agaonidae) 187 


Ian David Gauld 25 May 1947-12 January 2009 189