FHTET-2006-08

November 2006

ALEIODES WASPS OF EASTERN FORESTS:

A GUIDE TO PARASITOIDS AND ASSOCIATED

MUMMIFIED CATERPILLARS

Forest Health Technology

Enterprise Team

TECHNOLOGY 

TRANSFER Biological

Control

SCOTT R. SHAW

UNIVERSITY OF WYOMING

FHTET Forest

Service

U.S. Department 

of Agriculture
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On the cover:

Top: Acronicta americana mummy (induced by Aleiodes stigmator). 

Photo by David Wagner.

Center: Aleiodes indiscretus parasitizing Lymantria dispar (gypsy 

moth). Photo by Scott Bauer.

Bottom: Orgyia leucostigma mummy (induced by Aleiodes pallidator). 

Photo by Scott Shaw.
ALEIODES WASPS OF EASTERN FORESTS:

A GUIDE TO PARASITOIDS AND ASSOCIATED MUMMIFIED CATERPILLARS

Scott R. Shaw

Professor of Entomology and Curator, University of Wyoming Insect Museum 

Department of Renewable Resources (3354) 

University of Wyoming 

1000 East University Avenue 

Laramie, Wyoming 82071, USA

For additional copies of this publication, contact:

Richard Reardon

USDA Forest Service–FHTET

180 Canfield Street

Morgantown, WV 26505

Phone: (304) 285-1563

Fax: 304-285-1564

rreardon@fs.fed.us
ii 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

ACKNOWLEDGMENTS

Thanks are given to all those who supplied mummified caterpillars, especially: Pedro Barbosa, 

Paul Gross, Scott A. Lewins, Daniel M. Pavuk, Alex Segarra, David Wagner, and David Wahl. 

This work was supported by a research award from the USDA Forest Service, West Virginia 

University, and the McIntire-Stennis Project WYO-362-02. Support for computers was provided by a grant from the Wyoming NASA Space Grant Consortium. Field studies were supported by a grant from the University of Wyoming National Park Service (UW-NPS) Research 

Station. National Science Foundation grant NSF-BSI-03-46729 funded my graduate assistant 

Andrew (Drew) Townsend, provided support for my laboratory, and is allowing us to extend 

our Aleiodes research to the South American continent. John Strazanac and Richard Reardon provided helpful editing suggestions on the manuscript and Lee Dyer and Clark Pearson 

proofread the final publication for consistency and clarity. Thanks to Dr. Zhaojie Zhang, of 

the University of Wyoming Microscopy Core Facility, for providing very helpful advice with 

camera settings and to Danny Dalton for his assistance with digital photograpy of some mummified caterpillars. Special thanks to my mentor Dr. Paul M. Marsh for his support and collaboration on revisions of the North American Aleiodes fauna over many years. Drew Townsend 

read early versions of the manuscript and provided helpful suggestions.

Funding for publication was provided by the USDA Forest Service’s Forest Health Technology Enterprise Team (FHTET). Layout was done by Mark Riffe, International Technology 

Experts, for FHTET.
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

iii

Contents

INTRODUCTION .....................................................................................................1

WHY STUDY ALEIODES WASPS? ...............................................................................2

BIOLOGY OF ALEIODES SPECIES .................................................................................3

COLLECTING CATERPILLARS AND MUMMIES ..................................................................4

REARING ALEIODES WASPS FROM MUMMIFIED CATERPILLARS ...........................................6

SPECIMEN PRESERVATION .........................................................................................7

NAMES OF WASPS: SCIENTIFIC AND COMMON ............................................................8

SPECIES TREATMENTS ...............................................................................................9

Aleiodes abdominalis (Cresson) ..................................................................................... 10

Aleiodes aciculatus Cresson .......................................................................................... 12

Aleiodes akidnus Marsh and Shaw ................................................................................ 14

Aleiodes albitibia (Herrich-Schaeffer) ............................................................................. 15

Aleiodes angustipennis Marsh and Shaw ....................................................................... 17

Aleiodes burrus Cresson ................................................................................................ 18

Aleiodes cameronii (Dalla Torre) .................................................................................... 20

Aleiodes ceratomiae Marsh and Shaw ........................................................................... 22

Aleiodes earinos Shaw ................................................................................................... 24

Aleiodes femoratus Cresson .......................................................................................... 25

Aleiodes flavitarsus Marsh and Shaw ............................................................................. 26

Aleiodes gastritor (Thunberg) ......................................................................................... 28

Aleiodes geometrae (Ashmead) ..................................................................................... 30

Aleiodes granulatus (DeGant) ........................................................................................ 32

Aleiodes graphicus Cresson ........................................................................................... 33

Aleiodes indiscretus (Reardon) ...................................................................................... 35

Aleiodes itamevorus Shaw and Marsh ........................................................................... 37

Aleiodes laphygmae (Viereck) ........................................................................................ 39

Aleiodes lymantriae (Watanabe) .................................................................................... 41

Aleiodes malacosomatos (Mason) .................................................................................. 43

Aleiodes mandibularis (Cresson) .................................................................................... 46

Aleiodes maritimus Shaw and Marsh ............................................................................. 48

Aleiodes melanopodus Marsh and Shaw ....................................................................... 50

Aleiodes molestus (Cresson) .......................................................................................... 52

Aleiodes nigristemmaticum (Enderlein) .......................................................................... 54

Aleiodes nolophanae (Ashmead) ................................................................................... 56

Aleiodes notozophus Marsh and Shaw .......................................................................... 58

Aleiodes pallidator (Thunberg) species-complex ............................................................ 60

Aleiodes palmatoides Marsh and Shaw ......................................................................... 62
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ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Aleiodes palmatus (Walley) ........................................................................................... 63

Aleiodes parasiticus Norton ........................................................................................... 65

Aleiodes pardalotus Marsh and Shaw ............................................................................ 67

Aleiodes platypterygis (Ashmead) .................................................................................. 68

Aleiodes politiceps (Gahan) ........................................................................................... 70

Aleiodes preclarus Marsh and Shaw .............................................................................. 72

Aleiodes pseudoterminalis Marsh and Shaw .................................................................. 74

Aleiodes quadratus Shaw and Marsh ............................................................................. 76

Aleiodes quebecensis (Provancher) ................................................................................ 78

Aleiodes rileyi Cresson .................................................................................................. 80

Aleiodes rugulosus (Nees) ............................................................................................. 82

Aleiodes sanctihyacinthi (Provancher) ............................................................................ 84

Aleiodes scrutator (Say) ................................................................................................. 86

Aleiodes simillimus (Ashmead) ...................................................................................... 88

Aleiodes smithi Marsh and Shaw ................................................................................... 90

Aleiodes stigmator (Say) ................................................................................................ 92

Aleiodes terminalis Cresson ........................................................................................... 95

Aleiodes texanus Cresson .............................................................................................. 97

Aleiodes vierecki Marsh and Shaw ................................................................................ 99

Aleiodes virginiensis Marsh and Shaw ......................................................................... 100

Aleiodes wahli Marsh and Shaw .................................................................................. 101

PHOTOGRAPHY CREDITS ......................................................................................102

GLOSSARY .......................................................................................................102

LITERATURE ......................................................................................................109

INDEX .............................................................................................................113
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

1

INTRODUCTION

The parasitic wasp genus Aleiodes is found all 

around the world, but the greatest diversity of 

species occur in northern Europe, Asia, and 

North America. In North America alone, 

there are at least 90 Aleiodes species, and new 

species are still being discovered, especially 

from the western United States and northern 

Canada. On the other hand, the Aleiodes of 

the eastern United States are comparatively 

better known, and a field guide for the most 

common species is now feasible. This work 

is intended to provide an introduction to the 

more common Aleiodes species of eastern U.S. 

forests for forest scientists, ecologists, students, and others interested in the ecology of 

forest caterpillars and their parasitoids.

For purposes of this guide, the “eastern” 

United States is defined simply as including 

all areas east of the Mississippi River. This 

was purely an arbitrary decision on my part 

to allow some limits on what to include, and 

because this area seems to be well-enough 

studied to make this guide possible. The actual distributions of Aleiodes species vary quite 

a bit. Some of the treated species, such as A. 

virginiensis and A. wahli, are known only 

from sites in the eastern United States. Most 

of the species treated here have broader distributions, with ranges extending into Canada, 

the western United States, Mexico (DelfínGonzález and Wharton 2002), and sometimes 

even Central and South America (Shaw et al.

1997, 1998b; Marsh and Shaw 1998, 1999). A 

few species in the northeast (A. pallidator and

A. gastritor) have cirumpolar distributions 

and occur also in Canada, Europe, and northern Asia.

The decision as to what constitutes a “forest 

insect” was also somewhat arbitrary. Aleiodes

species vary quite a bit in their behaviors. 

Many are found in association with caterpillars that feed in the canopy of trees or large 

shrubs, so would be defined as arboreal, and 

therefore, are true forest insects. In some 

cases, this characterization may not be so 

clear: in the eastern United States, A. malacosomatos is clearly arboreal, mainly attacking 

eastern tent caterpillars in wild cherry trees; 

the same species occurs in the western United 

States, where the wasp attacks western tent 

caterpillars on very low shrubs in windswept 

meadows with no trees. Many other Aleiodes

species are associated with caterpillars feeding 

on grasses, low herbaceous plants, and low 

shrubs such as Vaccinium. These are species 

that, although not arboreal, are forest-associated, being found in the forest understory, 

in meadows, and in marshy areas. Many 

Aleiodes species might also occur in woody 

urban landscapes or in agroecosystems. But 

the natural habitat of virtually every eastern 

Aleiodes species is either forest or habitats 

near forest, so a practical decision was made 

to include essentially all known eastern 

Aleiodes in this guide.

Several distinct forest cover types occur in the 

eastern United States (Eyre 1980; Smith et al.

2000). Aleiodes species also seem to vary in 

their association with forest types. A few (A. 

nigristemmaticum, A. wahli) might be found 

only in association with oak-gum-cypress 

hammocks of far South. Others, such as A. 

gastritor, might only be found in the maplebeech-birch forests of the far North. Many 

others (e.g., A. burrus, A. texanus, A. virginiensis) favor the oak-hickory forests of the 

mid-Atlantic states.
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ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

WHY STUDY ALEIODES WASPS?

Aleiodes wasps attack and kill various species 

of leaf-feeding caterpillars. Included among 

their host caterpillars are several well-known 

forest pests, including the gypsy moth, eastern tent caterpillar, forest tent caterpillar, fall 

webworm, tussock moths, dagger moths, 

prominents, cutworms, and loopers. Aleiodes

species are highly beneficial because they 

serve as biological control agents for the natural suppression of populations of these forest 

insect pests. In many other cases, the host 

caterpillars are not usually regarded as pests, 

but might have the potential to become forest 

pests were it not for the natural suppression 

caused by Aleiodes parasitism.

Aside from their great economic importance, 

there is another very good reason to study 

Aleiodes wasps. They demonstrate a particularly unusual and distinctive method of 

parasitism that renders them especially valuable as research organisms: the host caterpillar is mummified. Caterpillars in the wild are 

killed by a wide variety of organisms, including predators, pathogens, and parasitoids; in 

most cases the caterpillar is either completely 

consumed or the remains shrivel and decay 

beyond recognition. Parasitism by Aleiodes is 

unique because the caterpillar cadaver remains intact and identifiable as the Aleiodes

larva feeds inside the caterpillar and eventually pupates within the somewhat shrunken 

remains. The resulting host mummy remains 

intact after the adult wasp emerges, and such 

caterpillar mummies provide valuable research 

tools. 

There are several advantages to studying these 

organisms. First, the parasitized caterpillars 

can be readily identified before the parasitic 

wasp emerges, and in most cases, the wasp 

species can be correctly identified before it 

emerges. Second, since the host mummy 

remains intact indefinitely, the host remains 

can be associated with the parasitic wasp in 

research collections, and the host species can 

be identified from the mummified remains. 

Some of the mummified caterpillars photographed for this manual are museum specimens that are more than 100 years old! Third, 

since Aleiodes are the only common parasitoids that cause this type of mummification, 

in many cases the parasitoid species involved 

can be reliably and easily identified. As there 

are many unanswered questions about host 

associations and host ranges, it is therefore 

possible for even amateur collectors to make 

important new discoveries. As a child growing up in Michigan, I can remember several 

times seeing these mummified caterpillars, 

but for many years I did not understand what 

had caused this phenomenon; I hope that this 

work will make these fascinating insects more 

readily accessible to field studies by future 

generations of budding young entomologists. 

Finally, since the mummification process is 

unique to particular species, if something 

other than Aleiodes emerges from one of these 

mummies, it can safely be assumed to be a hyperparasitoid. Thus, the study of hyperparasitoid complexes can be more reliably done by 

examining these organisms.
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

3

always through a circular postero-dorsal hole. 

Irregular exit holes formed in other regions of 

the mummy are invariably formed by hyperparasitoids.

The physiological basis for host mummification has not been experimentally investigated 

in Aleiodes, but Shaw et al. (1997) hypothesized that it might be caused by the physical 

elimination of the host’s corpora allatum by 

the developing parasitoid larva. Removal of 

the corpora allatum would reduce the juvenile hormone levels in the host caterpillar and 

induce the formation of a pupal cuticle. This 

hypothesis is consistent with the observation that larval feeding by Aleiodes is located 

initially in the prothoracic region of the 

caterpillar, as evidenced by the opening of the 

glue-hole.

Host range is quite variable, depending on 

the Aleiodes species. Only a few species are 

strictly monophagous—attacking only one 

caterpillar species. An example of this is A. 

ceratomiae, a parasitoid of Ceratomia amyntor hornworm caterpillars on various broadleaf trees. By attacking only a single species, 

such parasitoids by definition have the “narrowest” of all possible host ranges. Other 

Aleiodes species are oligophagous, attacking 

a small set of multiple caterpillar species. An 

example of this is A. texanus, a parasitoid 

of three different Sphingidae caterpillars on 

several broadleaf trees and vines. Oligophages 

are also regarded as having “narrow” host 

ranges as they are limited to a small set of 

host species. Finally, some Aleiodes species 

are polyphagous, attacking many different 

caterpillar species. An example of this is A. 

simillimus, a parasitoid of at least seven geometrid species on several broadleaf trees. It is 

fair to say that host range is an important, yet 

Some very detailed studies of European and 

Asian Aleiodes species have been done by 

Mark Shaw in Scotland (Shaw 1983, 1994, 

2002; Shaw and Huddleston 1991; Quicke and 

Shaw 2005). The following account of biology is condensed from these sources and from 

Shaw et al. (1997). By comparison, there have 

been only a few detailed studies of the biology 

of North American Aleiodes (for example, 

Pennington 1916).

The species of Aleiodes are internal parasitoids 

of caterpillars, especially macrolepidoptera of 

the superfamilies Noctuoidea and Geometroidea, and to a lesser extent, Arctioidea, Sphingoidea, and Papilionoidea. All are koinobionts, which means that the host caterpillar is 

not permanently paralyzed but continues to 

feed and grow for a period of time after it is 

parasitized. The method of parasitism, unique 

to the tribe Rogadini (including Aleiodes), is 

very distinctive and noteworthy (Quicke and 

Shaw 2005). The Aleiodes larva completes its 

feeding and pupates within the shrunken and 

mummified remains of the host caterpillar. 

In all known cases, the form of the mummy 

caused by a particular Aleiodes species is 

characteristic for that host and parasitoid, 

so the mummified caterpillar remains are of 

considerable diagnostic value and should be 

retained with the parasitoid, when reared. 

Host caterpillar mummies are usually found 

attached to the host plant substrate on leaves, 

stems, or bark. Most caterpillar mummies are 

firmly attached to the substrate by a glue-like 

substance that exudes through a prosternal 

hole chewed by the parasitoid larva. The 

inside of the mummy is lightly lined with silk 

by the parasitoid larva, but the main support 

for the mummy seems to come from a premature host pupal cuticle below the remaining 

larval cuticle. Exit from the host mummy is 

BIOLOGY OF ALEIODES SPECIES
4

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

poorly studied, aspect of the biology of most 

North American Aleiodes. There are several 

cases where an Aleiodes species has only one 

known host, but this does not prove that the 

species is really monophagous. It may be that 

there are other hosts that we have simply not 

discovered yet. Host range is an interesting 

aspect of the biology of these organisms to 

which students and amateur collectors can 

make valuable contributions to further our 

knowledge.

The host range of any given Aleiodes species 

is influenced by two factors. One important 

factor is phylogeny of the host caterpillars: 

that is to say, one parasitoid species may attack several caterpillar species that are closely 

related to each other. There are many examples of this in Aleiodes. Aleiodes quebecensis

attacks four different species of Noctuidae 

on various broadleaf trees, but all of them are 

members of the genus Acronicta. Aleiodes 

malacosomatos attacks two species of Malacosoma (tent caterpillars) in the eastern United 

States and other related species in the West. 

Aleiodes ceratomiae, A. texanus, and other 

members of the A. praetor species-group are 

entirely restricted to parasitism of hornworm 

caterpillars (family Sphingidae). The second 

factor that affects host range may be aspects 

of host ecology. In some situations, a parasitoid species may attack two or more caterpillar 

species that are not particularly closely related, but that share some similarities of appearance, feeding situation, or habitat preference. 

Aleiodes stigmator attacks various hosts in 

the Noctuidae and Notodontidae. The hosts 

are not particularly closely related, but all are 

rather large, apparent, setose caterpillars on 

wetland (river bottom) vegetation, such as 

willows and cattails.

COLLECTING CATERPILLARS AND MUMMIES

cies involved. The easiest to find involve host 

caterpillars that are gregarious and can be 

found in predictable circumstances. The best 

example of a very common mummy wasp is 

Aleiodes malacosomatos, which attacks young 

caterpillars of the eastern tent caterpillar. The 

mummies can be easily found in early June by 

peeling apart the silk layers of the tents. It’s 

not unusual to find 10-15 mummies inside one 

silk shelter. In other cases, the caterpillar attacked is solitary, but still is very conspicuous. 

This is the case with some of the large, hairy, 

noctuid caterpillars attacked by the gregarious parasitoid Aleiodes stigmator. These can 

be easily found by walking through willow thickets in August or September. These 

parasitized caterpillars tend to die in a vertical 

Caterpillars can be collected in various ways, 

as outlined in Covell (1984), Wagner (2005), 

Wagner et al. (1997), and many other sources, 

though generally, there are two approaches 

to finding and raising caterpillars that are 

parasitized by Aleiodes wasps. The first 

approach is to seek dead caterpillar mummies 

from which the Aleiodes has not yet emerged. 

The second approach is to collect live young 

caterpillars, keep them fed, and watch for 

those that become mummified later. The first 

approach is very important, because there is 

a great need for data on how mummies are 

formed in natural conditions. 

Finding mummified caterpillars can be very 

easy or very difficult depending on the spe-
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

5

position on twigs well above the ground, and 

they are large enough to be easily seen from 

quite a distance away (up to 50 feet). This, 

however, is an unusual situation. Most caterpillar mummies are small, involve solitary 

species, and may be extremely cryptic. When 

attached to a twig or stem, they may resemble buds or galls, so close and careful visual 

inspection is needed to find them. Those 

mummies formed on leaves and thus harder to 

locate can be collected by beating foliage and 

examining the deadfall. In some cases, such 

as Aleiodes albitibia, the mummy is not glued 

to a substrate and falls to the ground; such 

mummies might be found, through close inspection, by sifting leaf litter. In many cases, 

mummified caterpillars could most easily be 

obtained by raising them from infected caterpillars that are still alive and feeding.

When collecting caterpillar mummies, few 

tools are needed. A small knife or pruning 

shears are useful, as well as a beating sheet. 

If possible, a photograph should be taken of 

the mummy in its natural condition. Field 

samples can be collected in plastic bags, but 

as soon as possible, these should be separated 

into individual containers. When found attached to foliage, the twig or leaf should be 

cut rather than detaching the mummy from 

the substrate. Field notes should also be taken 

regarding the natural situation and included 

in the data labels. Removing the mummy 

from the substrate, if done carefully, does not 

kill the developing wasp inside, but the specimen is more informative if still attached to its 

natural substrate. 

Because the mummies are usually glued to a 

substrate, mummies up to several years old 

may be found. These may appear weathered 

(head capsule and setae will fall away as it 

ages), and emergence holes of the parasitoid 

may be visible. Solitary Aleiodes (most species) emerge postero-dorsally. Gregarious A. 

stigmator form numerous small round emergence holes, evenly spaced, across the dorsum 

of the mummy. Irregular holes on the sides or 

bottom of a mummy are indicative of hyperparasitoids. Even if the Aleiodes has emerged 

already, it is still valuable to collect older, 

mummified caterpillars: the caterpillar may 

still be identifiable, and it might give clues to 

previously unknown host relationships.

When seeking live caterpillars parasitized by 

Aleiodes wasps, it is important to consider 

certain issues. Aleiodes only attack caterpillars during certain stages of development. 

Usually, the second or third larval instar is 

attacked, and the wasp later emerges from the 

penultimate (next to the last) larva instar, so 

the best way to find infected caterpillars is to 

seek young caterpillars that are still feeding. 

You won’t find Aleiodes by rearing caterpillars from eggs, and large caterpillars seeking 

pupation retreats are not likely parasitized by 

Aleiodes. The best way to find infected caterpillars is by visual inspection of foliage where 

feeding damage is found. Other productive 

methods include beating foliage and searching beneath bands of material tied around 

tree trunks. The later method is particularly 

important for finding caterpillars that feed at 

night, but hide during the day.
6

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

preferable to collect data on natural parasitoid 

emergence rather than simply to obtain adult 

wasp specimens from lab rearings.

Once the adult wasp emerges, it can either be 

preserved for museum studies or kept alive 

for biological observation. Adult life may 

be prolonged by placing a sugar-water or 

honey-water solution in the container. Small 

vials with cotton-stopper wicks work well for 

feeding. If both males and females are reared, 

they can be placed in the same container to 

observe mating behavior. If live (potential 

host) caterpillars are available, female wasps 

can be placed in containers with caterpillars to 

attempt to rear more wasps or to test host acceptance on alternate hosts. Female wasps do 

not have to mate to lay fertile eggs—though 

unfertilized eggs will produce only males. 

When the studied wasp dies, the specimen 

should be preserved immediately.

When raising live caterpillars, the most important factors are to keep the caterpillars fed 

daily with fresh food, to keep the cages clean 

of frass and fungi, and to inspect the caterpillars daily for evidence of parasitism. When it 

is evident that a caterpillar is being parasitized, 

it should be carefully removed (with its substrate) to a separate rearing container. Careful observations, notes, and photos should be 

made regarding the mummification process in 

terms of its duration and how the caterpillar 

changes in appearance. Otherwise, once the 

mummified caterpillar is identified, the methods for handling the mummy are the same as 

outlined above.

Practical methods for rearing parasitic Hymenoptera from infected host insects were 

outlined by Mark Shaw (1997), an excellent 

publication that interested readers will find 

helpful. Some key ideas relating to Aleiodes

are mentioned here.

When already-mummified caterpillars are collected from the field, the mummies should be 

isolated into individual rearing containers as 

soon as possible. By looking for emergence 

holes, it can be quickly determined if the 

parasitoid has already emerged or not. With 

wild-collected mummies, there is no way to 

know how soon the parasitoid may emerge. 

Unless the parasitoid is over-wintering in the 

caterpillar mummy, the adult wasp can emerge 

within a few minutes, hours, days, or weeks, 

so it is very important to inspect the mummies 

promptly and place each mummy in isolation. 

Any small container will serve, but I prefer to 

use plastic petri dishes, 90 mm diameter by 30 

mm deep, lined with 90 mm white filter paper 

circles. Tops should be secured with a rubber 

band. Such containers can be easily set out on 

lab countertops for ease of viewing.

It is important for emergence to approximate 

natural conditions. If possible, the containers 

should be kept in an area that matches natural 

conditions of light, temperature, and humidity. For example, if the parasitoid is over-wintering in the mummified caterpillar, a prolonged cold period may be necessary to break 

diapause, and the containers might best be 

kept in an outdoor shed or unheated garage; 

mummies kept in a heated laboratory might 

not promote emergence. In any event, it is 

REARING ALEIODES WASPS FROM MUMMIFIED CATERPILLARS
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

7

The simplest method for preserving emerging Aleiodes wasps is to place the insect immediately in 70 percent to 95 percent ethanol 

in a small vial (2 dram vials with polyethelene 

screw-caps are ideal). If you kept the wasp 

alive for biological observations, it is important to preserve the specimen as quickly as 

possible after it dies: once the insect dies, it 

begins to desiccate, and fungi may soon grow 

on it. 

To achieve the best preservation, it is best to 

place the live wasp directly into the alcoholfilled vial. Eventually, the wasp specimen will 

need to be dry-mounted on point or card. 

But unless you have considerable experience with point-mounting small insects, most 

specialists (including myself) would probably prefer to get an intact, clean specimen in 

alcohol and do the dry-mounting themselves. 

In any event, the specimen should be clean, 

and better results can be obtained by placing 

wasps in alcohol first. The wasps can later be 

critical-point dried or chemically dried using 

chloroform (or similar methods) to achieve 

life-like, clean specimens. 

DO NOT pin Aleiodes wasps directly through 

the body. Although some of the larger species 

are large enough to get a pin into, doing so 

destroys some features of the mesosoma and 

renders the specimen less useful for photography.

Mummified caterpillars require no special 

methods of preservation as they will not 

deteriorate further if kept dry. Having said 

that, it is remarkable how many museum 

specimens have been improperly prepared 

by thoughtless methods. DO NOT pin the 

mummy directly. Although convenient, pinning the mummy may break it and puts two 

more holes in the mummy, rendering it less 

desireable for photography. If removed from 

the pin later, the pin holes may be mistaken as 

emergence holes. DO NOT glue the mummy 

to a data label. The best method is to keep the 

mummy dry by placing it alone in a small vial 

or empty gelatin capsule. Gelatin capsules 

or genitalia vials containing mummies can be 

affixed directly to the pin and associated with 

a given wasp. The mummy should be clearly 

labeled (for instance, with a code number) to 

indicate its association with a particular wasp 

specimen. If possible, preserve the mummy 

attached to the substrate (leaf, twig, etc.). DO

NOT remove the mummy from the substrate 

for preservation. Mummies on leaf parts, 

stems, etc. can be pinned by the plant material 

and kept dry in a collection. If the mummy 

is loose in a vial, it may be useful to put some 

cotton at either end to keep it from rattling 

about in handling or shipment. If the head 

capsule has fallen away (as normally happens 

with Sphingidae mummies and a few others), the head capsule should be kept with the 

mummy. 

SPECIMEN PRESERVATION
8

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

By historical tradition, parasitoid wasps 

(including Aleiodes) are called by their Latin 

scientific names (genus and species). Some of 

the larger stinging wasps do have well-known 

common names—such as mud-daubers, 

paper wasps, yellow jackets, and bald-faced 

hornets—but very few parasitoid wasps have 

been given common names. This is a somewhat curious situation, as many parasitoid 

wasps are quite common insects, and many 

are very beneficial insects of significant economic importance. Certainly, they deserve 

to have common names. To some extent, this 

seems to be only a matter of tradition within 

particular groups: for instance, many small 

moths and beetles have been given names even 

though they are not particularly common, 

while some insects with common names are 

actually quite rare. And so, one curious thing 

about the family Braconidae is that, although 

they are one of the largest family-groups of 

all insects (more than 50,000 species worldwide)—and one of the most beneficial and 

highly economically important—no one has 

bothered to give them common names. 

Several years ago, I suggested the common 

name “death-wasp” for insects in the family Braconidae as they are all parasitoids, and 

their key defining feature, biologically, is that 

they kill their host. For Aleiodes in particular, 

the common name “mummy-wasp” is appropriate. Some of my students have adopted 

the practice of using common names for 

braconids, but it has not yet caught on widely. 

This is probably because most writing about 

Braconidae is still done in the scientific journals and not in the popular press. Still, there 

are some situations where common names 

for insects might be useful or even preferable, 

such as for informal communication among 

amateur collectors and articles for the general 

public. Because this field guide is directed, in 

part, at a non-scientist audience (amateur collectors and students), I’ve included suggested 

common names for the wasp species. I apologize in advance to those who may not like certain names or would prefer others, but simple 

common names for these insects are provided 

in the hope that it will encourage more people 

to study these wasps and discuss them more 

widely. Readers are reminded that, in all cases 

of scientific publications and rigorous scientific correspondence, that the full scientific 

names should be used. I hope that my readers 

will find these common names to be useful; 

time will tell.

NAMES OF WASPS: SCIENTIFIC AND COMMON 
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

9

SPECIES TREATMENTS

The following pages present descriptions of the Aleiodes wasps and photographs of both the 

wasp and its mummified host.
10 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Aleiodes abdominalis (Cresson)

Orange-bellied mummy-wasp

Diagnostic features of adult wasp. 

Body mostly two-colored 

with contrasting orange and 

black. Head and mesosoma 

black. Metasomal terga 1-3 

orange, remaining metasomal 

terga varying from orange 

to black. Antenna and legs 

orange. Hind tarsomeres 

orange to brown, similar in 

color to tibia. Wings lightly 

pigmented. Tegula yellow. 

Body length 6.0-8.0 mm. 

Antenna with 54-63 segments. Oral opening circular, diameter about equal to 

basal width of mandible. Malar space 

longer than basal width of mandible. 

Ocelli small, distance between eye and 

lateral ocellus distinctly broader than 

ocellus width. Median length of pronotum shorter than distance between 

occipital carina and lateral ocellus. 

Mesopleuron smooth but setose medially, subalar sulcus rugose, sternaulus absent. Fore wing vein 1cu-a beyond 

vein 1M by less than 3 times its length. Vein RS of hind wing slightly curved 

downwards, marginal cell of hind wing gradually widening toward apex. 

Hind wing vein m-cu present. Apex of hind tibia without a row of flattened 

setae along inner margin, setae normal and hair-like. Tarsal claws strongly 

pectinate over entire length. First metasomal tergum not parallel-sided, 

wider apically than at base. Apex of metasoma not compressed in female. 

Metasomal tergum 3 costate at base, remainder smooth. Males with metasomal terga 4-6 densely setose (subdivided medially). 
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

11

Host caterpillars. One specimen from Maryland has been reared from an undetermined mummy of Noctuidae.

Diagnostic features of host mummies. Very dark brown, smooth, and about 1 cm 

long.

Wasp distribution. Widely distributed in eastern North America, from Quebec and 

Ontario south to North Carolina, westward to South Dakota and Arizona. 

More commonly encountered in the eastern parts of its range.

Biology. Adults appear in early June in the northern parts of its range.

Similar species. This species is similar to Aleiodes flavitarsus, but differs by its larger 

body size, fore wing with vein 1cu-a beyond vein 1M by less than three times 

its length, hind tarsomeres orange to brown, and hind wing vein m-cu present.

Best reference. Shaw et al. (1998a).

Comments. The smooth dark mummy would be highly cryptic if formed on a plant 

substrate, such as a twig, where it would resemble a bud. This might account 

for the scarcity of rearing records for a wasp that is fairly common in museum collections. Noctuidae sp.
12 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Aleiodes aciculatus Cresson

Finely-scratched mummy-wasp

Diagnostic features of adult wasp. Body 

mostly or entirely honey-yellow 

to light orange, except antenna 

apically dark brown to black. 

Male with apex of metasoma 

dark brown to black. Wings 

clear to lightly pigmented. Stigma two-colored, dark brown to 

black medially, pale yellowishwhite basally and apically. Body 

length 3.0-7.0 mm. Antenna 

with 36-52 segments. Oral opening small and circular, diameter 

about equal to basal width of 

mandible. Malar space longer 

than basal width of mandible. 

Head rugulose dorsally. Ocelli small, distance between eye and lateral ocellus 

distinctly broader than ocellus width. Pronotum long, median length of pronotum longer than distance between occipital carina and lateral ocellus. Hind 

wing vein RS slightly sinuate, marginal cell of hind wing narrowest in middle. 

Apex of hind tibia without a row of flattened setae along inner margin, setae 

normal and hair-like. Tarsal claws simple, not pectinate. First metasomal 

tergum not parallel-sided, wider apically than at base. Apex of metasoma not 

compressed in female. Metasomal terga 1-4 longitudinally costate or aciculate.

Host caterpillars. This wasp has been recorded as attacking several species of Noctuidae, including Feltia ducens, Feltia subgothica, and Mythimna (=Pseudaletia) 

unipuncta, and the geometrid Euchlaena serrata.

Diagnostic features of host mummies. Very dark brown, slender, smooth, and about 

7 mm long.

Euchlaena serrata
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

13

Wasp distribution. Widely distributed over the eastern United States, from Massachusetts south to Florida, and westward to South Dakota, Utah, and Texas.

Similar species. Aleiodes aciculatus is distinctive from most species because it has 

sculpturing on the fourth metasomal tergum resembling fine longitudinal 

lines, as though scratched by a needle. In appearance, it is somewhat similar 

to A. angustipennis, which also has similar sculpture on the fourth tergum. 

Aleiodes aciculatus can be distinguished by its two-colored stigma and a 

shorter pronotum than is found in A. angustipennis. 

Biology. Not known in detail. Adult wasps have been collected from April through 

September, indicating that the species is probably multivoltine.

Best reference. Marsh and Shaw (2003).

Comments. This species is very distinctive in having the fourth metasomal tergum 

strongly sculptured and distinctly costate or aciculate, resembling fine longitudinal lines as though scratched by a needle.
14 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Aleiodes akidnus Marsh and Shaw

Small-eyed mummy-wasp

Diagnostic features of adult wasp. 

Antennal flagellum brown. 

Body color honey-yellow,

legs somewhat lighter. 

Ocellar triangle black. 

Fore wing stigma brown. 

Body length 4.0 mm. Antenna with 43-45 segments. 

Compound eye small, malar space equal to 2/3 eye 

height. Ocelli small, ocellocular distance slightly 

more than twice diameter of 

lateral ocellus. Fore wing 

vein r equal to or slightly 

longer than vein 3RSa. Second submarginal cell of fore wing small and nearly square. Hind wing vein 

RS weakly sinuate. Hind wing marginal cell narrowest in middle. Apex of 

hind tibia with a row of flat setae on inner side.

Host caterpillars. Unknown.

Diagnostic features of host mummies. Unknown.

Wasp distribution. Southern Florida.

Biology. Adult wasps fly from July to October. They are attracted to blacklights.

Similar species. Of the species with a fringe of flat setae on the hind tibia, A. akidnus is 

quite distictive by its very small ocelli, small eyes, and wide malar space. It is 

not likely to be confused with any other species in the eastern United States. 

Aleiodes akidnus is most similar to the Caribbean species, A. sanctivincentensis.

Best reference. Marsh and Shaw (1998).

Comments. Known from Big Pine Key and from the Tall Timbers Research Station.
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

15

Aleiodes albitibia (Herrich-Schaeffer)

White-legged mummy-wasp

Diagnostic features of adult wasp. 

Body color quite variable. 

Head and antenna mostly 

black except malar space 

and mouthparts yellow. 

Mesosoma with pronotum 

black, mesonotum yellow 

to black, scutellum yellow, mesopleuron mostly 

yellow, but sometimes 

with black markings, and 

propodeum black dorsally 

and yellow laterally, or 

sometimes entirely yellow. 

Legs mostly yellow but

hind tibia yellowish-white 

on basal half. Hind tarsus brown. Metasomal terga usually mostly black, 

but first tergum with narrow yellow median area and second tergum widely 

yellow medially. Sometimes, first and second tergum almost entirely yellow. 

Wings clear. Stigma black, sometimes with light spots, basally and apically. 

Body length 4.5-5.5 mm. Antenna with 41-45 segments. Oral opening small 

and circular, diameter about equal to malar space. Malar space short, slightly 

less than basal width of mandible. Ocelli large, distance between eye and lateral ocellus about half ocellus width. Mesopleural disc above middle leg and 

below wing base smooth and shining. Vein RS of hind wing arched medially, 

marginal cell of hind wing narrowest in middle. Apex of hind tibia without 

a row of flattened setae along inner margin, setae normal and hair-like. Tarsal claws with a wide gap between apical claw and basal comb of four or five 

spine-like setae. First metasomal tergum not parallel-sided, wider apically 

than at base. Apex of metasoma not compressed in female.

Host caterpillars. Attacks mainly caterpillars of Notodontidae and possibly some species of Geometridae. Recorded hosts include Nadata gibbosa, Nerice bidentata on Ulmus americana, Notodonta dromedarius, and Phaeosia species.
16 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Diagnostic features of host mummies. Caterpillars mummified by this species are 

smooth, very dark, and have an unusual expanded bubble-like area anteroventrally. These mummies are also unusual in that there is no antero-ventral 

“glue-hole,” and the mummy is not attached to a substrate. The mummification process typically causes decapitiation of the caterpillar, so the head 

capsule separates and falls away from the mummy. There has been some 

speculation that the unusual bubble-like area may allow this mummy to float 

in flooded areas. As the mummy does not have a glue-hole to attach the 

mummy to a plant surface, it would tend to fall to the ground (where floating 

might be adaptive).

Wasp distribution. This wasp is a widely occurring holarctic species found across 

Europe, Northern Asia, Canada, and the northern United States. In eastern 

North America, the species is found from Quebec southward to Georgia.

Biology. Shaw (1983) suggested that the antero-ventral swollen area may serve as an 

“air sac,” allowing the host mummies to float in water.

Similar species. This species can be easily distinguished from all other North American species by the central disc of the mesopleuron, which is smooth, highly 

polished, and without setae.

Best reference. Shaw et al. (1998b).

Comments. North American specimens of A. albitibia exhibit more range of color 

variation (often with more yellow or orange) than European specimens 

(which have more black areas). Nadata gibbosa

Nerice bidentata
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

17

Aleiodes angustipennis Marsh and Shaw

Narrow-winged mummy-wasp

Diagnostic features of adult wasp. Body 

color entirely yellow. Fore wing 

stigma yellow. Body length 7.0 

mm. Antenna with 49-50 segments. Pronotum long and flat, 

with median length about twice 

ocell-ocular distance. Fore wing 

long and narrow, width about 

1/5 its length. Fourth metasomal 

tergum sculptured, costate basally.

Host caterpillars. Unknown.

Diagnostic features of host mummies. 

Unknown.

Wasp distribution. Known only from Florida.

Biology. The caterpillar hosts of this wasp species 

are not yet known. In Florida, the adult 

wasps have been collected in March, 

April, July, August, and September. 

This suggests that the wasp is probably 

multivoltine.

Similar species. The combination of long, narrow wings and a long, flat pronotum easily distinguishes A. angustipennis from most other wasp species. It is closely 

related to A. granulatus, but can be easily distinguished as A. angustipennis

lacks dark color markings and lacks extensive granular sculpture. Aleiodes 

angustipennis is superficially similar to A. aciculatus (which is also yellow), 

but A. angustipennis has a yellow stigma, longer pronotum, and longer wings.

Best reference. Marsh and Shaw (2003).

Comments. The wings of this species are longer and more narrow than any other U.S. 

species.
18 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Aleiodes burrus Cresson

Little-horse mummy-wasp

Diagnostic features of adult wasp. Body color orangish brown in female. Males twocolored: black and orangish brown. Antenna orangish brown basally, brown 

to black apically, without a white annulus. Body length 6.0-8.0 mm. Females with 45-52 antennal segments; 55-60 antennal segments in males. Oral 

opening oval, diameter equal to or slightly greater than malar space. Malar 

space wide, greater than basal width of mandible. Ocelli small, ocell-ocular 

distance greater than width of lateral ocellus. Fore wing vein 1cu-a beyond 

1M by distance distinctly greater than length of 1cu-a. Hind wing vein RS 

straight or bent downwards slightly near middle. Hind wing marginal cell 

narrowest at base, widening to wing 

apex. Tarsal claws simple, not pectinate. 

First metasomal tergum coarsely rugose.

Host caterpillars. This species has been reared from the cutworm Feltia subgothica

(Haw.) and Agrotis species. It has been recorded in the literature (Marsh 

1979a) as being associated with Acronicta hasta Gn. and A. lobeliae Gn., but 

these records are questionable and need confirmation. These two Acronicta

species are more likely to be hosts of A. quebecensis.
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

19

Diagnostic features of host mummies. Rather large, thick-bodied mummies about 

14-15 mm long and 4-5 mm wide. Smooth and rather dirty- to dark-brown. 

In dorsal view, the mummy widens posteriorly and is widest near the posterior end. The sides of the mummy have a slightly-flattened raised ridge, visible 

in lateral view as a longitudinal line.

Wasp distribution. This species is widely distributed, from eastern Canada southward 

to Virginia and westward to Alberta, Wyoming, and Arizona.

Biology. Aleiodes burrus is a common species in the eastern United States, but its biology has not been carefully studied.

Similar species. Similar to A. smithi, but A. burrus does not have a white annulus on 

the antenna.

Best reference. Marsh and Shaw (2001).

Comments. Cresson (1869) did not explain the meaning of this species name. The 

Latin word burrus means “red” but the wasp is mostly orangish-brown, not 

red. The name is possibly a variation of the Latin word burricus, meaning “a 

small horse.” Perhaps Cresson thought that this brown, fairly thick-bodied 

wasp looks like a little horse. I suppose it does, if you have a vivid imagination. Another possibility is that the name could be a now-obscure reference to a historic person: Sextus Afranius Burrus, a prefect of the Praetorian 

Guard during the times of emperors Claudius and Nero. Burrus is remembered for his military leadership and died in 62 A.D. (possibly by poison). 

However, this similar name may be just an interesting coincidence. I think 

it is unlikely that Cresson named this species after a person as, in his original paper, he did not capitalize the species name, whereas, in the same paper, 

he did capitalize the name of the species named after C.V. Riley (the species 

known to us now as Aleiodes rileyi was named “Aleiodes Rileyi” by Cresson). 

The custom of capitalizing species names based on a proper name has since 

been discontinued (now the species name is always in lowercase), but that 

was a common custom in Cresson’s time. It seems likely that if Aleiodes burrus were actually named after a person, then Cresson would have capitalized 

the name.

Feltia subgothicaAgrotis sp.
20 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Aleiodes cameronii (Dalla Torre)

Cameron’s pitted mummy-wasp

Diagnostic features of adult wasp. Body color honey-yellow, except antenna and ocellar triangle brown. Body length 7.0-9.0 mm. Antenna long, with 60-65 segments. Flagellomeres all short and compact. Malar space short, about equal 

to half of mandible width. Oral space small and circular, about twice as wide 

as malar space. Ocelli very large, lateral ocellus nearly touching eye, ocellocular distance only one-twelfth diameter of lateral ocellus. Occipital carina 

complete. Antero-lateral margin of propodeum granulate. Tarsal claws 

strongly pectinate. Males with large setose pits medially on terga 5-7.

Host caterpillars. In Maryland, this species has been reared from several exposedfeeding arboreal caterpillars on willow (Salix nigra). Confirmed hosts include a geometrid, Eutrapela clemataria (J.E. Smith), two catocaline noctuids, 

Zale lunata (Drury) and Catocala cara Gn., and one unidentified species of 

Notodontidae.
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

21

Diagnostic features of host mummies. The mummy formed from Catocala cara is 

quite distinctive: about 1.5 cm long, dark brown, densely wrinkled on apical 

half and ending with two long caudal prolegs that form a conspicuous forked 

“tail” at the tip of the mummy.

Wasp distribution. Aleiodes cameronii is widely distributed in the eastern and southern United States and ranges southward into Mexico and Costa Rica. In the 

United States, it occurs from Massachusetts southward to Florida and across 

the southern states westward to southern California.

Biology. Barbosa et al. (2001) found A. cameronii to be one of the dominant parasitoids of exposed-feeding arboreal caterpillars on willow in riparian habitats 

in Maryland. There, the species is a moderate-generalist, utilizing at least 

four different host species from three different lepidopteran families, but all 

willow-feeders. The adult wasps have very large ocelli and are attracted to 

lights, indicating that they are nocturnally active. This might be an adaptation for searching for larvae that feed at night. 

Similar species. A very distinctive species, not likely to be confused with any other. 

Only a few other species have setose pits on the male metasoma, and the pits 

in A. cameronii are much larger than in any other known species. It is most 

similar to A. earinos and A. geometrae, but in those species, the occipital 

carina is not complete, and the tarsal claw is not so fully pectinate as in A. 

cameronii.

Best reference. Shaw et al. (1997).

Comments. The function of the very large setose pits on the male abdomen is not 

known, but Shaw et al. (1997) hypothesized that they might serve to disperse 

sex pheromones. Studies of the courtship behavior of this, and related species, might be very interesting. Catocala cara Catocala cara
22 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Aleiodes ceratomiae Marsh and Shaw

Big-eyed mummy-wasp

Diagnostic features of adult wasp. Body color black with yellow to organge markings. Metasoma partly or entirely black. Body length 9 mm. Antenna with 

65-66 segments. Malar space extremely short, about three-fourths mandible 

width. Eyes and ocelli enormous, covering most of head. Ocell-ocular distance very narrow, lateral ocellus approaching eye margin. Lateral ocellus 

four to six times wider than ocell-ocular distance. Hind wing vein RS sinuate and strongly arched, RS approaching wing margin medially. Hind wing 

marginal cell very narrow in the middle.

Host caterpillars. Reared from a mid-instar larva of the waved sphinx moth, Ceratomia undulosa (Walker), on ash. Also recorded as associated with the elm 

sphinx moth, Ceratomia amyntor (Geyer), but that record is questionable; in 

this case, the larva was feeding on ash, the usual host plant of C. undulosa.
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

23

Diagnostic features of host mummies. About 15 mm long, smooth, pale brownish-white with bands of tiny reddish-brown dots. Broadly attached to the 

substrate by an antero-ventral glue-hole. Head capsule decapitated from 

mummy or attached by a small piece of cuticle that is prone to break as the 

mummy ages. Posteriorly, has a distinct caudal horn about 4 mm long or a 

distinct pit if the caudal horn has fallen off. 

Wasp distribution. Known only from the northeastern United States (Connecticut, 

New Hampshire, New York) and eastern Canada (Ontario).

Biology. The host mummy is only about 15 mm long, indicating that the caterpillar is 

killed at an early stage of development—probably the third instar. This wasp 

is an arboreal species associated with ash and possibly other broadleaf trees 

and shrubs, including oaks, hawthorns, privets, lilacs, and elm. The wasp has 

been collected at lights, and the extremely large ocelli are indicative of nocturnal activity. Collecting dates for the wasp range from late April to midAugust, indicating that the wasp is active for much of the summer.

Similar species. Most similar to A. texanus, but A. ceratomiae can be easily distinguished by its exceptionally large ocelli and black metasoma, whereas A. 

texanus has smaller ocelli and a reddish-orange metasoma.

Best reference. Shaw et al. (1998b).

Comments. Aleiodes ceratomiae has large eyes and larger ocelli than any other North 

American Braconidae. Large ocelli are presumed to be an indication of 

nocturnal activity—presumably, the female wasp searches for host caterpillars at night. This species is a member of the praetor species-group, the 

only Aleiodes group known to exclusively attack Sphingidae. The caterpillar mummies are particularly distinctive because of the caudal horn and the 

tendency of the head capsule to be decapitated during the mummification 

process. Ceratomia undulosa

C. undulosa head capsule
24 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Aleiodes earinos Shaw

Spring mummy-wasp

Diagnostic features of adult wasp. Body 

color yellowish to reddish-brown, 

except antenna black and tegula 

orange. Body length 6.0-7.0 

mm. Antenna long, with 57-

65 segments. Flagellomeres all 

short and compact. Malar space 

short, about equal to, or slightly 

narrower than, basal width of 

mandible. Oral space small and 

circular, about twice as wide as 

malar space. Ocelli large, ocellocular distance about 1/4 of the 

diameter of lateral ocellus. Occipital carina incomplete, weak, 

or absent on vertex. Propodeum 

granulate anteriorly, grading to rugose posteriorly. Tarsal claws pectinate at 

extreme base, with a wide gap between apical claw and basal pecten. First 

metasomal tergum slightly longer than wide. Males with small setose pits 

medially on terga 4-6.

Host caterpillars. Unknown.

Diagnostic features of host mummies. Unknown.

Wasp distribution. Known only from Florida, Arkansas, and Texas.

Biology. Adult wasps are active in the spring during April and May and are attracted 

to lights.

Similar species. Aleiodes earinos is similar to A. geometrae, with which it has been 

often confused. Similarities with A. geometrae include claws with a wide gap 

between the apical claw and basal pectination, weak to incomplete occipital carina, and small setose pits on the male terga. Differences in A. earinos

include the larger body size, darker reddish-brown color, longer flagellum, 

less broad gap in the occipital carina, coarser propodeal sculpture, and much 

longer first metasomal tergum.

Best reference. Shaw et al. (1997).

Comments. Although the host caterpillars are not known for A. earinos, the activity 

of the wasp in the early spring and the close relationship of this species to A. 

geometrae both suggest the possibility of Geometridae hosts.
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

25

Aleiodes femoratus Cresson

Banded-leg mummy-wasp

Diagnostic features of adult wasp. Body 

color mostly honey-yellow. Propleuron, mesopleuron, and hind 

femur marked with dark brown. 

Fore wing stigma unicolored 

brown. Body length 6.0-7.0 mm. 

Antenna with 47-52 segments. 

Ocelli large, ocell-ocular distance 

distinctly smaller than diameter 

of lateral ocellus. Hind wing vein 

RS straight. Hind wing marginal 

cell gradually widening to apex. 

Apex of hind tibia with a row of 

flat setae on inner side.

Host caterpillars. Unknown.

Diagnostic features of host mummies. Unknown.

Wasp distribution. Found only in the eastern United States, from Illinois and West 

Virginia southward to Florida.

Biology. This species has been collected in Illinois and Virginia in mixed deciduous 

forests dominated by oaks, hickories, and maples. Adult wasps fly from midJune to mid-August.

Similar species. Similar to A. nigristemmaticum, but A. femoratus is distinguished by 

its solid dark brown stigma, gradually widening hind wing marginal cell, and 

with dark markings on the hind femur.

Best reference. Marsh and Shaw (1998).

Comments. This species has been known for more than 100 years, but no one has yet 

reared it from a caterpillar host. It has been collected several times by Malaise trap, indicating that it flies low to the ground.
26 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Aleiodes flavitarsus Marsh and Shaw

Yellow-footed mummy-wasp

Diagnostic features of adult wasp. Body 

mostly two-colored with contrasting orange and black. Head 

and mesosoma black. Metasomal 

terga 1-3 mostly orange, remaining metasomal terga (4-8) black. 

Antenna light brown. Legs 

mostly yellow except for black 

apex of tarsus, apical fourth of 

hind femur, and apical half of 

hind tibia. Wings clear. Tegula 

yellow. Body length 4.5-5.5 mm. 

Antenna with 46-49 segments. 

Oral opening oval, diameter 

about equal to basal width of 

mandible. Malar space equal 

to basal width of mandible. Ocelli small, 

distance between eye and lateral ocellus 

equal to, or distinctly broader than, ocellus 

width. Median length of pronotum shorter 

than distance between occipital carina and 

lateral ocellus. Mesopleuron smooth but 

setose medially, subalar sulcus rugose, 

sternaulus absent. Forewing vein 1cu-a beyond vein 1M by nearly three times its length. Vein RS of hind wing straight, 

marginal cell of hind wing gradually widening toward apex. Hind wing vein 

m-cu absent. Apex of hind tibia without a row of flattened setae along inner margin, setae normal and hair-like. Tarsal claws strongly pectinate over 

entire length. First metasomal tergum not parallel-sided, wider apically than 

at base. Apex of metasoma not compressed in female. Metasomal tergum 3 

costate at base, remainder smooth. Males with metasomal terga 4-6 densely 

setose (subdivided medially).

Host caterpillars. This species has been reared in Canada from an unknown species of 

the genus Autographa (Noctuidae) feeding on alder.
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

27

Diagnostic features of host mummies. Mummy smooth, with scattered small setae 

and irregular brown and whitish tan markings (as pictured).

Wasp distribution. This is a boreal species that is widely distributed across Canada, 

from Newfoundland westward to British Columbia, and extends southward 

into the northern United States, including Washington and Minnesota. In 

the eastern United States, A. flavitarsus has only been recorded in Maine and 

Michigan.

Biology. Adult wasps are active from late June to September. There is one record of 

an adult wasp emerging in March, but this may have been under laboratory 

conditions. The only confirmed host association is with an Autographa species feeding on alder.

Similar species. Aleiodes flavitarsus is similar to A. abdominalis but differs by having a 

smaller body size, vein 1cu-a of the fore wing beyond vein 1M by about three 

times its length, by the absence of vein m-cu in the hind wing, and by the 

hind tarsomeres 1-4 being yellow.

Best reference. Shaw et al. (1998a).

Comments. The mostly yellow legs, with the hind tarsi being mostly yellow, are quite 

distinctive.

Autographa sp.
28 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Aleiodes gastritor (Thunberg)

Thunberg’s mummy-wasp

Diagnostic features of adult wasp. Body color mostly honey-yellow, except ocellar 

triangle, propodeum, base of tergum 1, and sides of terga 2 and 3 dark brown 

to black. Stigma two-colored but with darkened medial area not discrete, 

rather translucent, with dark color grading into lighter basal and apical areas. 

Body length 4.0-5.0 mm. Antenna with 35-36 segments. Hind wing with 

vein RS arched in the middle, marginal cell narrowest near middle.

Host caterpillars. Recorded in Europe as attacking a wide range of arboreal Geometridae and one drepanid. Confirmed hosts in eastern North America include 

the geometrids Alsophila pometaria, the fall cankerworm, and Hypagyrtis 

unipunctata.

Diagnostic features of host mummies. Mummies about 9-10 mm long, mostly light 

whitish to yellowish-brown, with fine mottled black markings and tranverse 

black bands in prothoracic area. Smooth and slightly tapered at both ends. Alsophila pometaria
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

29

Wasp distribution. This is a very widely distributed circumpolar boreal species found 

in northern Europe, northern Asia, Canada, and the northeastern United 

States.

Biology. Aleiodes gastritor is a generalist species attacking various arboreal Geometridae and sometimes Drepanidae.

Similar species. Aleiodes gastritor is part of a sibling-species complex including A. 

laphygmae, A. malacosomatos, A. nolophanae, A. platypterygis, A. simillimus, 

and A. stigmator. These species are quite similar in appearance and are most 

reliably identified based on reared material.

Best reference. Shaw (1994).

Comments. Although fairly common in northern states, the presence of A. gastritor

in North America has not been generally recognized, and for many years this 

species has been confused with A. stigmator. Previous records of A. stigmator attacking Alsophila pometaria are erroneous (e.g., Marsh 1979a) and 

actually refer to Aleiodes gastritor. When reared, the two species are easy to 

distinguish: Aleiodes gastritor is a solitary species, attacking mostly small arboreal geometrids, while A. stigmator is a gregarious species, attacking much 

larger noctuids and notodontids.
30 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Aleiodes geometrae (Ashmead)

Geometrid mummy-wasp

Diagnostic features of adult wasp. Body color honey-yellow to orange, antenna black 

and tegula yellow. Body length 5.0-6.0 mm. Antenna long, with 43-56 segments. Flagellomeres all short and compact. Malar space short, about equal 

to, or slightly narrower than, basal width of mandible. Oral space small 

and circular, about twice as wide as malar space. Ocelli large, ocell-ocular 

distance about two-thirds of the diameter of lateral ocellus. Occipital carina incomplete, weak, or absent on vertex. Propodeum mostly granulate

anteriorly, grading to rugose posteriorly. Tarsal claws pectinate at extreme 

base, with a wide gap between apical claw and basal pectin. First metasomal 

tergum shorter than wide. Males with small setose pits medially on terga 4-6.

Host caterpillars. Reared from geometrids, including the spring cankerworm, Paleacrita vernata (Peck); Semiothisa ocellinata (Gn.); and an unidentified geometrid host on honey locust.
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

31

Diagnostic features of host mummies. Mummies are about 9-10 mm long, forming a 

dark reddish-brown, smooth capsule with narrow transverse pale yellowish 

bands. Head capsule and anal prolegs are pale yellowish-white. Mummies 

sometimes found glued to the underside of host plant leaves.

Wasp distribution. Widely distributed from Ontario and New York, southward to 

Florida, and westward to North Dakota, Colorado, and Texas.

Biology. Adult wasps fly from mid-March to mid-August. This wasp is arboreal, 

nocturnally-active, and appears to be oligophagous on exposed tree-feeding 

geometrids. Hosts are associated with a variety of trees, including maple, 

honey locust, elm, birch, and Prunus.

Similar species. Aleiodes geometrae is similar to the southern A. earinos, but A. geometrae is a more widespread and common species. Similarities with A. earinos include claws with a wide gap between the apical claw and basal pectination, weak to incomplete occipital carina, and small setose pits on the male 

terga. Differences between A. geometrae and A. earinos include the former’s 

smaller body size; lighter, mostly yellow body color; shorter flagellum; wider 

gap in the occipital carina; more granular propodeal sculpture; and a shorter, 

wider first metasomal tergum.

Best reference. Shaw et al. (1997).

Comments. One of the host caterpillars, Paleacrita vernata, is sometimes a serious defoliating pest. Aleiodes geometrae may be a natural control agent for spring 

cankerworm.

Geometridae sp.Geometridae sp.
32 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Aleiodes granulatus (DeGant)

Granular mummy-wasp

Diagnostic features of adult wasp. Body 

multi-colored, black, yellow, 

and orange. Head black, antenna and mouthparts yellow. 

Mesosoma sometimes entirely 

black, pronotum and mesonotum 

always black, mesopleuron and 

propodeum sometimes orange. 

Metasoma with first tergum 

orange or black, second and 

third terga orange, remainder of 

terga black. Legs orange, except 

apical half of hind femur and 

tibia black. Stigma two-colored: 

brown with yellow at base and 

apex. Body length 4.5-5.0 mm. 

Antenna with 45-50 segments. Body with 

extensive granular microsculpture on head and 

mesosoma. Pronotum long, with median length 

greater than ocell-ocular distance. Fore wing 

vein 1cu-a beyond 1M by distance less than 

length of 1cu-a. Metasomal terga 1-4 longitudinally costate.

Host caterpillars. Reported as associated with the noctuid Trichoplusia ni.

Diagnostic features of host mummies. None available for study.

Wasp distribution. Found in the northeastern United States and Canada, from New 

Brunswick south to North Carolina and westward to Wisconsin and Ohio.

Biology. Unknown.

Similar species. Aleiodes granulatus is closely related to A. angustipennis but differs by 

having the fore wing vein 1cu-a close to vein 1M and by its black and orange 

body color.

Best reference. Marsh and Shaw (2003).

Comments. This species is very distinctive by its extensive granular microsculpture.
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

33

Aleiodes graphicus Cresson

Black-winged mummy-wasp

Diagnostic features of adult wasp. Body mostly yellowish orange, except ocellar 

triangle, palpi, and legs apically black. Wings black. Body length 7.0-8.0 

mm. Antenna with 55-60 segments. Malar space short, slightly shorter than 

basal width of mandible. Eyes and ocelli large. Ocell-ocular distance narrow, 

lateral ocellus near eye margin. Lateral ocellus one to two times wider than 

ocell-ocular distance. Hind wing vein RS sinuate and strongly arched, RS 

approaching wing margin medially. Hind wing marginal cell very narrow in 

middle.

Host caterpillars. This wasp species has been reared from at least two species of 

Sphingidae, but the host mummies have not been identified beyond the 

family level. The host mummy from Florida (pictured) was associated with 

Camellia.

Diagnostic features of host mummies. About 12-15 mm long, smooth, pale brownish-tan, sometimes with dark lateral markings, and with scattered tiny spinules. Broadly attached to the substrate by a large antero-ventral glue-hole. 

Head capsule either decapitated from mummy or attached by a small piece of 

cuticle that is prone to breaking away as the 

mummy ages. Has a distinct posterior caudal horn about 4 mm long or a distinct pit if 

the caudal horn has fallen off. The formation of a large irregular emergence hole by 

the emerging wasp may cause the posterior 

end of the mummy to fall away (resulting in 

Sphingidae sp.

a loss of the caudal horn).
34 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Wasp distribution. Aleiodes graphicus is widely distributed across the southern 

United States from Arizona to Florida, northward to Colorado and South 

Dakota. It is more common in the southwest and central United States and 

is relatively uncommon in the East. The only collection record in the eastern 

states is from Florida.

Biology. Mummies are firmly glued down by a very large antero-ventral glue-hole, 

which usually results in decapitation of the caterpillar. Thus, wild-collected 

mummies usually lack a head capsule. The caudal horn of sphingid hosts is 

quite distinctive, but may fall off as the mummy ages or overwinters, leaving 

a visible pit. Adult wasps are attracted to lights and are probably nocturnally 

active.

Similar species. Most closely related to A. texanus, but that species has extensive black 

markings on the head and mesosoma. Aleiodes graphicus can be easily distinguished by its almost entirely orange body and black wings. The only species 

with superficially similar color is A. politiceps, but A. graphicus can be distinguished from that species by its strongly curved hind wing RS vein and very 

large ocelli.

Best reference. Shaw et al. (1998b).

Comments. Aleiodes graphicus is seldom collected in the eastern United States, where 

it may be restricted to the far south.
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

35

Aleiodes indiscretus (Reardon)

Indiscrete mummy-wasp

Diagnostic features of adult wasp. 

Body color mostly honeyyellow except ocellar triangle 

black, dorsum of mesosoma 

and metasoma marked with 

light brown. Body length 

6.0-8.0 mm. Antenna with 

50 segments. Oral opening 

small and circular. Ocelli 

very large, ocell-ocular distance about one-third of the 

diameter of lateral ocellus.

Host caterpillars. Lymantria dispar

(L.) and Dasychira species.

Diagnostic features of host mummies. Dasychira mummies are about 1.5 cm long 

and densely hairy, with white and black setae. Gypsy moth caterpillars 

mummified by this species were not available for study.

Wasp distribution. This species is native to India, but was introduced to the eastern 

United States (Connecticut and Massachusetts) in the late 1960s for suppression of the gypsy moth (Reardon 1970).

Biology. In India, this species is a parasitoid of Lymantria obfuscata Walker, a relative 

of the gypsy moth (Rao 1964). It was cultured in the laboratory on gypsy 

moth, L. dispar, and released for control of that pest. Details of its biology 

are provided by Reardon et al. (1973). Recently, it has been reared from 

Dasychira species in Maryland. A. indiscretus on gypsy moth

Dasychira prob. basiflava
36 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Similar species. This species is very similar to A. pallidator, and careful studies of both 

species are needed to determine whether they are really two distinct species. 

Aleiodes indiscretus differs from A. pallidator by having the temples more 

strongly converging behind the eyes; by having slightly more coarse sculpture on the mesopleuron, propodeum, and first tergum; and by the body 

color being somewhat darker and more reddish-brown. Aleiodes indiscretus

reproduces sexually, while A. pallidator reproduces parthenogenetically.

Best references. Reardon (1970), Reardon et al. (1973), and Marsh (1979b).

Comments. This exotic species now appears to be established in the eastern states 

from Massachusetts southward to Maryland. Mummies of Dasychira induced by this species are virtually identical to those of A. pallidator. 
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

37

Aleiodes itamevorus Shaw and Marsh

Itame mummy-wasp

Diagnostic features of adult wasp. Body mostly black with yellow markings on the 

face and orbits of the eyes. Legs yellow. Body length 3.5-4.5 mm. Antenna 

with 41-44 segments. Metasoma carapace-like, in dorsal view with no segments visible beyond tergum 4. Vein m-cu of the hind wing distinctly tubular.

Host caterpillars. Itame brunneata (Thunberg).

Diagnostic features of host mummies. Mummies are 8-9 mm long, pale whitish to 

yellowish-brown, mottled with irregular black markings, and very smooth. 

The mummy is formed along small stems of the host plant and is very cryptic.

Wasp distribution. Known only from Ontario, Canada, but given the distribution of 

the host, this wasp is probably present throughout boreal parts of the northeastern areas of the United States and Canada.

hind wing

Itame brunneata
38 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Biology. This wasp has been reared from the geometrid Itame brunneata (Thunberg) 

on Vaccinium angustifolium.

Similar species. Very similar to A. maritimus, but that species lacks a tubular vein mcu in the lower part of the hind wing.

Best reference. Shaw and Marsh (2004).

Comments. This small but distinctive wasp is a resident of northern blueberry bogs.
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

39

Aleiodes laphygmae (Viereck)

Armyworm mummy-wasp

Diagnostic features of adult wasp. 

Body color entirely 

honey-yellow. Antenna 

brown. Wings transparent. Pterostigma twocolored: brown with 

yellow spots at base and 

apex. Body length 4.0-5.0 

mm. Antenna with 33-35 

segments. Ocell-ocular 

distance equal to, or 

slightly wider than, lateral 

ocellus width. Angle between vein r of fore wing 

and pterostigma about 

160 degrees, vein r often 

appearing nearly on same line as hind margin of pterostigma.

Host caterpillars. Recorded from several small noctuid caterpillars including the armyworm moth, Mythimna (=Pseudaletia) unipuncta (Haw.); the beet armyworm moth, Spodoptera exigua (Hbn.); the fall armyworm moth, S. frugiperda (Smith); and the yellow-striped armyworm moth, S. ornithogalli (Guen.).

Diagnostic features of host mummies. Although formed from several caterpillar species, mummies resulting from A. laphygmae parasitism are similar in 

appearance. Very early stages of the caterpillars are attacked, so the resulting mummies are quite small, being only about 8-9 mm long. They are very 

slender, smooth, and usually light brown or gray in color, with a darker head 

capsule. Mummies of caterpillar species with longitudinal stripes, such as S. 

ornithogalli, retain a visible striped pattern. The mummy lays very flat on its 

substrate or only slightly inclined upward from the front end. Spodoptera frugiperda Spodoptera ornithogalli
40 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Wasp distribution. Found in the southeastern United States from South Carolina 

southward to Florida and westward to Texas. Also recorded from Cuba and 

Nicaragua. Introduced to Hawaii.

Biology. This species is an oligophagous feeder on early stages of cutworm and armyworm caterpillars feeding in low vegetation. Associated naturally with open 

grassy meadows with mixed herbaceous vegetation. In forest areas, restricted 

to the undergrowth and open meadows. This species also occurs commonly 

in agroecosystems.

Similar species. Aleiodes laphygmae is very similar to A. nolophanae, but can be distinguished from that species by its shorter antenna, larger ocelli, and by the 

orientation of the vein r in the fore wing.

Best reference. Viereck (1912).

Comments. This species can be quite abundant, but because of its small size and the 

cryptic brown color of the mummies, its presence can be easily overlooked.
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

41

Aleiodes lymantriae (Watanabe)

Gypsy moth mummy-wasp

Diagnostic features of adult wasp. 

Body color mostly dark 

brown to black, except legs 

mostly yellow, and with 

1 or 2 yellow spots medially on the dorsum of the 

metasoma. Body length 

6.0-8.0 mm. Antenna with 

55 segments. Ocelli very 

large, diameter of lateral 

ocellus about three times 

the ocell-ocular distance. 

Host caterpillars. A parasitoid of the gypsy moth, Lymantria dispar.

Diagnostic features of host mummies. Although parasitized at an early stage, mummies of the gypsy moth induced by this species are rather large (11-12 mm 

long) and quite distinctive. The head capsule is yellowish-brown. The body 

is much darker and densely covered by raised, round, blue or brown protuberances, each of which is thickly set with long hairs. Many of the hairs 

project out to the sides of the mummy by as much as 1 cm.

Lymantria disparLymantria disparOrgyia recens (Japan)
42 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Wasp distribution. Aleiodes lymantriae is native to Japan but has been introduced into 

the eastern United States for control of the gypsy moth. 

Biology. Adults occur in mid-summer. In addition to attacking the gypsy moth, in 

Japan this species overwinters inside caterpillars of Orgyia recens. In a laboratory setting, it has been raised on caterpillars of O. thyellina (Marsh 1979b).

Similar species. This species is most similar to A. indiscretus, but is easily distinguished 

by its much darker color with yellow markings. Both species have very large 

ocelli and are associated with caterpillars of the gypsy moth.

Best references. Watanabe (1937) and Marsh (1979b).

Comments. It is not known whether this species has become established in the eastern 

United States or, if so, how widely. It might now occur in the eastern states 

wherever the gypsy moth is found.
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

43

Aleiodes malacosomatos (Mason)

Tent caterpillar mummy-wasp

Diagnostic features of adult wasp. 

Body color mostly yellowish orange, except metanotum and propodeum 

dark brown to black. 

Metasoma usually entirely orange but sometimes 

with dark brown or black 

basally on first tergum. 

Stigma two-colored, dark 

brown to black medially, 

pale yellowish-white basally and apically. Body 

length 5 mm. Antenna 

with 33-35 segments in 

female, 37-39 segments 

in male. Frons, temple, and vertex with fine 

granular sculpture. Malar space longer than 

basal width of mandible. Tarsal claws weakly 

pectinate basally, appearing simple. First metasoma tergum wider at apex than long. Hind 

wing with vein RS arched in the middle, marginal cell narrowest near middle.

Host caterpillars. In the eastern United States this species attacks young larvae of the 

eastern tent caterpillar (Malacosoma americanum) and the forest tent caterpillar (M. disstria). In the western United States and Canada, M. californicum is 

also parasitized. Mummies of M. americanum are commonly formed inside 

the silk tents of the host or on nearby vegetation. Large numbers of mummies can sometimes be found by peeling apart the layers of the silk tents in 

early June. Mummies retain cuticular color patterns characteristic of the different Malacosoma species. Malacosoma americanum
44 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Diagnostic features of host mummies. Mummy about 1 cm long, head capsule and 

body mostly black, with pale bluish white median line and scattered lateral 

bluish white marks. Covered with long yellowish-white hairs.

Wasp distribution. Aleiodes malacosomatos is widely distributed across the northern 

United States and Canada, from New Brunswick southward to New York 

and westward to Idaho and British Columbia. In the eastern United States, 

this species is common in Massachusetts, Michigan, and New York.

Biology. Aleiodes malacosomatos is an arboreal species associated with broadleaf trees 

and shrubs, especially wild cherry. It parasitizes the gregarious larvae of tent 

caterpillars, but the wasp is always solitary, with only one wasp emerging 

from the mummy of the half-grown caterpillar.

Similar species. Aleiodes malacosomatos is quite similar morphologically to A. stigmator, but has coarser and less shining sculpture on the head and mesosoma, and 

its fore wing stigma is distinctly two-colored. The habits of the two species 

are quite different, as A. stigmator is a gregarious parasitoid of much larger 

noctuid and notodontid caterpillars.

Best reference. Mason (1979).

Comments. Although A. malacosomatos can be locally quite abundant, it is not often 

collected unless it is reared from the host mummies. The mummified caterpillars can be easily collected in June by peeling apart the silk layers of the 

shelters of the host caterpillars. During the spring of 1988 these parasitoids 

were very common in tents of M. americanum on wild cherry trees at Rock 

Meadow, Belmont, Massachusetts. However, during the following spring of 

1989, none could be found at the same sites. My studies of this wasp in Wyoming indicate that the mummies are subject to high rates of hyperparasitism 

by Encyrtidae, Pteromalidae, and Ichneumonidae.

 This species presents a curious puzzle relating to its seasonal occurrence. 

Because the host Malacosoma species overwinter as egg masses on twigs and 

Aleiodes always oviposit into young larvae, it can be assumed that the adult Malacosoma americanum Malacosoma disstria
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

45

wasps are present in the very early spring (late April or early May), although 

oviposition has not been observed. As adult wasps emerge from mummies of parasitized Malacosoma in June and July, either the species has an 

alternate host later in the summer or the adults are extremely long-lived (as 

Malacosoma larvae will not be available for attack until the following spring). 

Overwintering as adults cannot be ruled out, but it seems unlikely that 

emergence would be in June and July unless there were a second generation 

on an alternate host. Also, in the laboratory, the adult wasps live only for a 

week or two. A search was made of wild cherry trees at Rock Meadow in 

July of 1988 for possible alternate hosts, but without success. The only likely 

prospect found was the ugly nest caterpillar, which is also a gregarious feeder 

making a large, prominent, silken shelter. These were presented to female A. 

malacosomatos wasps in the laboratory, but the parasitoids made no attempt 

to oviposit.
46 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Aleiodes mandibularis (Cresson)

Large-jawed mummy-wasp

Diagnostic features of adult wasp. Body orange with brown antennae and clear to 

slightly dusky wings. Body length 8.5-10.0 mm. Antenna with 55-65 segments. Oral opening large and oval, width greater than height of face. Malar 

space short, its width equal to or slightly less than width of mandible at base. 

Mandibles large and robust. Width of ocell-ocular 

distance slightly less than width of lateral ocellus. 

Tarsal claws strongly pecinate.

Host caterpillars. Unknown.

Diagnostic features of host mummies. Unknown.

Wasp distribution. This species is widely distributed in the central United States, but 

it is usually rather scarce and seldom collected. It ranges from Ohio southward to Georgia and westward to Nebraska and Texas.

Biology. Unknown. It is attracted to lights, suggesting nocturnal activity. Other 

members of the melanopterus species-group (of which this species is a member) are known to attack caterpillars of Noctuidae.

Similar species. Aleiodes mandibularis is large and distinctive, and it probably would 

not be confused with any other species. The wasp superficially resembles 

A. politiceps because both have a large orange body, but can easily be distinguished from that species by its larger oral space, larger mandibles, and 

clear wings. It resembles A. melanopodus, but differs by having orange legs, 

whereas the legs of A. melanopodus are black.
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

47

Best reference. Marsh and Shaw (1999).

Comments. This is a very large and quite distinctive Aleiodes species, but it is rather 

rare and seldom collected. It has larger, thicker mandibles than any other 

eastern U.S. Aleiodes species. The reasons for its scarcity are not known.
48 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Aleiodes maritimus Shaw and Marsh

Maritime mummy-wasp

Diagnostic features of adult wasp. 

Body mostly black with 

yellow markings on the 

face and vertex of the 

head. Legs yellow. Body 

length 3.5-4.5 mm. Antenna with 42-43 segments. Metasoma carapace-like, in dorsal view 

with no segments visible 

beyond tergum 4. Vein 

m-cu of the hind wing absent or indicated only by a 

weakly pigmented line.

Host caterpillars. Semiothisa granitata (Guenée), S. sexmaculata Packard, and S. signaria dispuncta (Walker).

Diagnostic features of host mummies. Mummies are 8-9 mm long, yellowish tan to 

brown, mottled with irregular black markings, and very smooth. When the 

mummy is formed along plant stems it is very cryptic.

Wasp distribution. The majority of specimens collected so far are from the Canadian 

maritime provinces of Newfoundland, New Brunswick, and Nova Scotia. 

However, a few specimens from central and eastern Canada and Minnesota 

are known, indicating that this species is probably widespread in northern 

North America

Biology. Specimens have been reared from the following geometrids: Semiothisa 

granitata, S. sexmaculata, and S. signaria dispuncta. The host caterpillars feed 

on various coniferous trees, including pitch pine, scrub pine, firs, hemlocks, 

spruces, and larch.

Semiothisa sexmaculata
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

49

Similar species. This species is quite similar to A. itamevorus, but differs from that 

species by lacking a tubular vein m-cu in the lower part of the hind wing and 

having more coarse sculpture on the basal terga of the metasoma.

Best reference. Shaw and Marsh (2004).

Comments. Most likely, this species occurs in the northeastern states of New England, but it has not been collected there yet.
50 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Aleiodes melanopodus Marsh and Shaw

Black-footed mummy-wasp

Diagnostic features of adult wasp. Body and base of legs (coxae and trochanters) orange. Legs (beyond trochanters) black. Wings dusky blackish. Body length 

8.0-9.0 mm. Antenna with 53-58 segments. Oral space very wide and oval, 

its width about two times height of face. Malar space very short, its width 

about three-fourths width of mandible at base. Diameter of lateral ocellus 

about three-fourths ocell-ocular distance. Tarsal claws strongly pectinate. 

Ovipositor sheaths about 1.5 times longer than hind basitarsus.

Host caterpillars. Unknown.

Diagnostic features of host mummies. Unknown.

Wasp distribution. Known only from the eastern and southern United States. This 

species has been collected in Maryland, Virginia, North Carolina, Kentucky, 

and Louisiana.

Biology. Details unknown. Adult wasps fly in the late summer and early fall (September and October). Other members of the melanopterus species-group, of 

which this species is a member, are known to attack caterpillars of Noctuidae.

Similar species. This species is somewhat similar to A. mandibularis in color and shape 

of the mouthparts. It differs from A. mandibularis by having somewhat 

smaller mandibles, smaller ocelli, longer ovipositor, and black legs. 

Best reference. Marsh and Shaw (1999).
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

51

Comments. This species, although large and distinctive, was not named until 1999. 

The first specimen that I saw was collected by a student in my Introductory 

Entomology class as part of a student collection. This demonstrates that this 

is still an area in which students and amateur collectors can make significant 

new discoveries and contributions.
52 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Aleiodes molestus (Cresson)

Troublesome mummy-wasp

Diagnostic features of adult 

wasp. Body two-colored: black and orange. 

Head black. Mesosoma 

mostly orange, except 

propodeum variously 

marked with black and 

legs mostly black. Metasomal terga 1-3 always 

orange, remaining terga 

varying from orange to 

black. Body length 5.0-

7.0 mm. Antenna with 

45-47 segments. Malar 

space long, greater than 

basal width of mandible. 

Ocelli small. Tarsal claws strongly pectinate with 6-8 stout teeth. Marginal 

cell of hind wing narrowest basally, gradually widening to wing apex. Males 

with terga 4-7 densely covered with silvery setae (subdivided medially).

Host caterpillars. Reared from several species of noctuine and plusiine noctuids, 

including Autoplusia egena (Gn.); the soybean looper, Pseudoplusia includens 

(Wlkr.); the cabbage looper, Trichoplusia ni (Hbn.); and the variegated cutworm, Peridromia saucia (Hbn.). The known hosts are all generalist feeders 

on a variety of types of low vegetation, including several agricultural crops.
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

53

Diagnostic features of host mummies. Mummy about 1 cm long, with body forming 

a mostly smooth, pale yellowish-white capsule. Darker and densely wrinkled 

at anterior end. The head capsule is darker yellowish-brown and pulled flat 

to the substrate with anterior face upwards. Mummies sometimes formed on 

a dense silk mat spun by the host larva on the host plant.

Wasp distribution. This species is quite common in midwestern and southwestern 

states, ranging from South Dakota south to Arkansas, Louisiana, and Texas 

and westward to Wyoming, Utah, Arizona, and southern California. In 

the eastern United States it has been recorded from Florida, but it probably 

ranges across the Gulf Coast. It is also known from Mexico and Costa Rica.

Biology. Adult wasps fly from late April through mid-September in the southern 

states. Aleiodes molestus favors open meadows, fields with low vegetation, 

and agroecosystems.

Similar species. Quite similar to the northern species, A. parasiticus, but not likely to 

be confused as their ranges do not overlap. Aleiodes molestus differs from A. 

parasiticus by having the legs entirely black or dark brown beyond the coxa 

(in A. parasiticus, the hind femur and tibia are banded, orange and black).

Best reference. Shaw et al. (1998a).

Comments. The name “molestus” is Latin for “troublesome.” The wasp is not annoying to humans, but rather the name refers to the fact that the wasp makes 

trouble for the caterpillars it attacks. The wasp should be regarded as highly 

beneficial as it attacks several significant pest insects. prob. Pseudoplusia includens
54 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Aleiodes nigristemmaticum (Enderlein)

Dark-marked mummy-wasp

Diagnostic features of adult 

wasp. Entire body light 

yellow, except top of 

head (ocellar triangle) 

marked with black. 

Fore wing stigma yellow. Body length 6.5-

7.5 mm. Antenna with 

50-57 segments. Ocelli 

large, ocell-ocular 

distance distinctly less 

than diameter of lateral 

ocellus. Occipital carina complete. Fore wing 

vein r nearly as long as 

vein 3RSa. Hind wing 

vein RS parallel to wing margin on basal third, then suddenly curved downward. Hind wing marginal cell suddenly widened at apex. Apex of hind tibia 

with a row of flat setae on inner side.

Host caterpillars. Mocis latipes (Gn.) and other Mocis species (Noctuidae).

Diagnostic features of host mummies. The mummified caterpillar is about 1 cm 

long, dark brown to black, smooth, and tapering at both ends. Mummies 

are formed vertically on grasses, twigs, and low vegetation, attached with the 

head-end downward. The remainder of the mummy is unattached and bends 

away from the substrate at a 45° angle. The resulting mummy is extremely 

cryptic, very much resembling a plant part (seed or sepal). Mocis latipes
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

55

Wasp distribution. In the eastern United States, known only from southern Florida 

and Mississippi. The species is widely distributed around the Gulf of Mexico 

and the Caribbean, being recorded also from Mexico, Honduras, Costa Rica, 

Venezuela, Cuba, and sites in the West Indies. 

Biology. This species is found in early-successional forests, among low vegetation 

in the understory, or in grassy areas along forest margins. The process of 

mummy formation was studied in Costa Rica by Les Price, and is described 

in Marsh and Shaw (1998).

Similar species. Aleiodes nigristemmaticum is similar to A. femoratus, but can be distinguished by its yellow stigma and by the lack of dark markings on the hind 

femur. Aleiodes nigristemmaticum is entirely light yellow, except for a single 

black mark on the top of the head (hence the species name).

Best reference. Marsh and Shaw (1998).

Comments. The recorded host in Honduras, Mocis latipes, is a common moth (and 

caterpillar) in the southern United States. It is a generalist feeder on grasses 

and other low vegetation, including several crop plants such as corn, rice, 

beans, and turnips. 
56 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Aleiodes nolophanae (Ashmead)

Appleworm moth mummy-wasp

Diagnostic features of adult wasp. Body color entirely honey-yellow. Antenna 

brown. Wings transparent. Pterostigma two-colored: brown with yellow 

spots at base and apex. Body length 5.0-6.0 mm. Antenna with 39-45 segments. Ocell-ocular distance equal to, or slightly wider than, lateral ocellus 

width.

Host caterpillars. Recorded as attacking the many-dotted appleworm moth, Balsa 

malana (Fitch), and other small noctuids, including the green cloverworm, 

Plathypena scabra (F.), Bomolocha palparia, Lithophane sp., and Caenurgia 

erechtea.

Diagnostic features of host mummies. Although formed from several caterpillar species, mummies resulting from A. nolophanae parasitism are similar in appearance. Very early stages of the caterpillars are attacked, so the resulting mummies are quite small, being only about 8-9 mm long. They are very slender, 

smooth, and usually light brown or gray in color, with a darker head capsule. Bomolocha palparia Lithophane sp.
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

57

Wasp distribution. This species occurs throughout the eastern half of the United 

States.

Biology. This species is a polyphagous feeder attacking several small noctuid species 

feeding on low vegetation or small trees, such as apple and boxelder. Lentz 

and Pedigo (1974) provided a detailed study of the biology of this species.

Similar species. Quite similar to A. laphygmae, but this species is distinguished by its 

longer antenna and larger body size.

Best references. Lentz and Pedigo (1974) and Barbosa et al. (2001).

Comments. Barbosa et al. (2001) recorded this species as a dominant parasitoid of 

small noctuid caterpillars on boxelder.
58 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Aleiodes notozophus Marsh and Shaw

Southwestern mummy-wasp

Diagnostic features of adult wasp. Head including antenna dark brown to black. 

Apex of palpi light brown to nearly white. Mesosoma except propodeum 

brown to dark brown. Propodeum always light brown. Metasoma and legs 

light brown. Body length 6.5-7.0 mm. Antenna with 51-54 segments. Flagellomeres all short and compact. Malar space very short, about equal to 2/3 

basal width of mandible, or narrower. Oral space small and circular, about 

as wide as basal width of mandible. Ocelli very large, ocell-ocular distance 

at most 1/4 diameter of lateral ocellus, lateral ocellus often nearly touching compound eye. Occipital carina incomplete, weak, or absent on vertex. 

Propodeum mostly granulate anteriorly, grading to rugose posteriorly. Tarsal 

claws strongly pectinate with seven to eight large spines. First metasomal tergum longer than wide. Males with small setose pits medially on terga 4-7.

Host caterpillars. Unknown.

Diagnostic features of host mummies. Unknown.

Wasp distribution. In the eastern United States, this species is recorded only from 

Florida. It occurs in the southwestern United States (Arizona and California) 

and in Costa Rica.
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

59

Biology. The holotype specimen from California is labeled as reared from an “oak 

gall.” The plant association is probably correct, but it seems unlikely that 

this wasp would emerge from a gall. Most likely a mummified host caterpillar was confused with a gall. Some other members of this species-group (A. 

geometrae, for example) form mummies that are very smooth, dark brown, 

and might be mistaken for a gall.

Similar species. Aleiodes notozophus is a member of the pulchripes species-group, 

which includes other species with males that have setose pits on the metasomal terga (A. cameronii, A. earinos, A. geometrae). Aleiodes notozophus is 

easily distinguished from these by its dark-colored head.

Best reference. Shaw et al. (1997).

Comments. In Florida, this species has been collected in May. Adult wasps are attracted to blacklights (ultraviolet light).
60 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Aleiodes pallidator (Thunberg) species-complex

Pallid mummy-wasp

Diagnostic features of adult wasp. Body 

color entirely golden yellow. 

Antenna with 53-57 segments. 

Ocelli very large, ocell-ocular 

space distinctly narrower than 

width of lateral ocellus. Parasites 

of Lymantriidae caterpillars.

Host caterpillars. The satin moth, Leucoma salicis (L.). In Maryland, 

possibly other Lymantriidae, including Orgyia definita Packard; 

the white-marked tussock moth, 

Orgyia leucostigma (J.E. Smith); 

and Dasychira obliquata (Grote 

and Robinson).

Diagnostic features of host mummies. The body of the mummified host caterpillar is 

about 1 cm long, with hair tufts projecting both forward and backward such 

that the whole mummy is up to 2 cm long when hairs are measured as well as 

body. The lymantriid hosts are so densely hairy that dead mummified hosts 

look very much like living caterpillars except for the posterior emergence 

hole of the wasp. Mummies retain the color and hair characteristics of living 

caterpillars.

Wasp distribution. This species is native to Europe, where it is reported to be a monophagous parasitoid of the satin moth. During the 1930s, it was introduced 

and released in the northeastern United States (Maine, Massachusetts, New 

Hampshire, Rhode Island, and Connecticut) and the Pacific northwest 

(Washington) for biological control of the satin moth. More recently, it has 

been reported from Maryland (Barbosa et al. 2001).

Dasychira obliquataLeucoma salicis

Orgyia leucostigma
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

61

Biology. Prior to its release into the United States, the biology of this species was 

studied in great detail. There is one generation per year, with the parasitoid 

overwintering as a larva inside overwintering caterpillars. Dowden (1938) 

studied this species (as Rogas unicolor) and reported that it is a monophagous 

feeder on caterpillars of Leucoma salicis. He also noted that, in Europe as 

well as the populations released into the United States, reproduction by this 

species is parthenogenetic, with females producing females without males. 

More recent studies by Mark Shaw confirm that, in Europe, the species is a 

strict monophage, reproducing parthenogenetically (Shaw 2002). The material recently found in Maryland poses an interesting puzzle as it includes both 

males and females and has been reared from other lymantriid hosts, including

Orgyia definita Packard; the white-marked tussock moth, Orgyia leucostigma (J.E. Smith); and Dasychira species. It is not yet clear whether this is a 

sexually reproducing population of A. pallidator or whether this population 

is a previously unrecognized sibling species.

Similar species. Very similar to A. indescretus, a parasitoid of the gypsy moth. However, A. pallidator is lighter in color and has less dense sculpture, especially 

notable on the side of the mesopleuron where there is a smooth shiny area. 

By comparison, A. indiscretus is more reddish-brown and has coarser sculpture on the mesopleuron.

Best reference. Dowden (1938) and Shaw (2002).

Comments. It is not yet clear whether the Maryland population on Orgyia and Dasychira is a sexually reproducing population of A. pallidator or whether this 

population is a previously unrecognized sibling species.
62 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Aleiodes palmatoides Marsh and Shaw

Palm-like mummy-wasp

Diagnostic features of adult wasp. Body 

color honey-yellow with brown 

markings on the mesosoma and 

metasoma. Body length 4.0-5.5 mm. 

Antenna with 33-35 segments. First 

metasomal tergum long, narrow, and 

parallel-sided. Apex of metasoma 

laterally compressed in females.

Host caterpillars. Unknown.

Diagnostic features of host mummies. Unknown.

Wasp distribution. Known only from mixed 

forest localities in Virginia and 

North Carolina.

Biology. Unknown. Adult wasps are active from July to October.

Similar species. Similar to the more northern species, A. palmatus. Easily distinguished from that species by the dark brown markings on the body.

Best reference. Marsh and Shaw (2003).

Comments. The compressed apex of the metasoma places this species in the compressor species-group. Studies of A. compressor in Europe indicate that the compressed metasoma is an adaptation for parasitizing host caterpillars that feed 

in the concealment of leaf shelters tied by silk (Shaw 1994). Therefore this 

species might also be attacking concealed host caterpillars.
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

63

Aleiodes palmatus (Walley)

Walley’s palmate mummy-wasp

Diagnostic features of adult wasp. Body color entirely honey-yellow. Body length 

6.0-7.0 mm. Antenna with 33-35 segments. First metasomal tergum long, 

narrow, and parallel-sided. Apex of metasoma laterally compressed in females.

Host caterpillars. Hosts include the noctuids Nycteola cinereana Neumoegen and 

Dyar and Enargia decolor (Wlk.).

Diagnostic features of host mummies. Rather non-descript smooth mummies about 

9-10 mm long, yellowish-brown with fine yellowish transverse wrinkles.

Wasp distribution. This species is known to range across Canada from Ontario to 

British Columbia and south into California, Utah, Minnesota, and Wisconsin. Given the known hosts and the association with Populus tremuloides 

forests, it seems likely that the species will eventually also be discovered in 

northern Michigan and Maine.
64 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Biology. Recorded as a solitary parasitoid of the noctuid Nycteola cinereana. Based 

on museum specimens, it has also been reared from the noctuid Enargia decolor and an unknown caterpillar on Populus tremuloides (pictured).

Similar species. Very similar to A. palmatoides, but that species is smaller and has 

brown markings on the body; A. palmatus is a bit larger and entirely yellow.

Best reference. Walley (1941) (as a species of Petalodes) and Marsh and Shaw (2003).

Comments. This species and the closely related A. palmatoides are the only two U.S. 

Aleiodes species with a strongly compressed metasoma in the females. They 

are seldom collected, but are not likely to be confused with other species.
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

65

Aleiodes parasiticus Norton

Parasitic mummy-wasp

Diagnostic features of adult wasp. Body two-colored: black and orange. Head black. 

Mesosoma mostly orange, but mesopleuron and propodeum black. Legs 

orange and black. Metasomal terga 1-2, and basal half of tergum 3, orange. 

Apical half of tergum 3 and remaining terga black. Body length 5.0-7.0 mm. 

Antenna with 49-55 segments. Malar space long, greater than basal width of 

mandible. Ocelli small. Tarsal claws strongly pectinate with five or six stout 

teeth. Marginal cell of hind wing narrowest basally, gradually widening to 

wing apex. Males with terga 4-7 densely covered with silvery setae (subdivided medially).

Host caterpillars. Reared from caterpillars of plusiine noctuids, including Anagrapha 

falcifera (Kby.) and Syngrapha epigaea (Grt.). Anagrapha falcifera is a generalist-feeder on low plants including clover and blueberries, while S. epigaea is 

a generalist feeding on conifers such as pine, spruce, and fir.

Diagnostic features of host mummies. Mummies are about 12 mm long, cream-colored, and fairly smooth, except at anterior end. Cuticle is densely wrinkled 

at anterior end, behind head capsule. Anagrapha falcifera
66 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Wasp distribution. Eastern Canada and the northeastern United States, southward to 

Maryland, and westward to North Dakota, Wyoming, and Colorado.

Biology. Aleiodes parasiticus is a distinctly northern species associated with boreal 

forests. Adult wasps are active for most of the summer, from late May to 

early September. The confirmed host records are from caterpillars of plusiine 

noctuids. Norton (1869) and Shenefelt (1975) report this species as emerging 

from a diprionid sawfly, Neodiprion abietis (Harris), on Abies, but that seems 

highly unlikely.

Similar species. Aleiodes parasiticus is most similar to the southern species A. molestus, 

but the ranges of the two species do not overlap. Aleiodes parasiticus differs 

by having the hind femur and tibia banded, orange on the basal half and black 

on the apical half, while the legs of A. molestus are mostly black.

Best reference. Shaw et al. (1998a)

Comments. Aleiodes parasiticus has a very distinctive black-orange-black-orangeblack color pattern. It cannot be confused with any other northern species, 

and can be reliably spot-identified in the forest, even without a microscope.
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

67

Aleiodes pardalotus Marsh and Shaw

Leopard-spotted mummy-wasp

Diagnostic features of adult wasp. Body color mostly yellow but with dark brown 

and black spots on the head, mesosoma, and metasoma. Fore wing stigma 

two-colored: brown with yellow at base and apex. Body length 5.5 mm. Antenna with 50 segments. Ocelli large, ocell-ocular distance about one-third 

the diameter of lateral ocellus. Hind wing vein RS slightly sinuate. Hind 

wing marginal cell narrowest in middle. Apex of hind tibia with a row of flat 

setae on inner side.

Host caterpillars. Unknown.

Diagnostic features of host mummies. Unknown.

Wasp distribution. Known only from Archbold Biological Station in southern Florida.

Biology. The caterpillar hosts of this wasp species are not yet known. Adult wasps fly 

in late January.

Similar species. Similar to A. femoratus and A. nigristemmaticum in body form, but A. 

pardalotus is easily distinguished from these and all other U.S. species by the 

very distictive pattern of leopard-like dark-colored spots.

Best reference. Marsh and Shaw (1998).

Comments. The forest habitat at Archbold Biological Station is fairly open and dry, 

and is dominated by low oak scrub and some taller pine flatwoods (Mark 

Deyrup, pers. comm.).
68 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Aleiodes platypterygis (Ashmead)

Broad-winged mummy-wasp

Diagnostic features of adult wasp. Body color mostly honey-yellow, but ocellar triangle dark brown to black. Stigma two-colored but with darkened medial area 

not discrete, rather translucent, with dark color grading into lighter basal and 

apical areas. Body length 4.0-5.0 mm. Antenna with 34-36 segments. Hind 

wing with vein RS arched in the middle, marginal cell narrowest near middle.

Host caterpillar. Masked birch caterpillar, Drepana arcuata (Walker).

Diagnostic features of host mummies. The most distinctive feature of the mummy 

is the head capsule, which is yellowish-brown with two dark brown transverse bands. The mummy is about 9-10 mm long, light brown with irregular 

dark brown intersegmental color, and mostly smooth, but with some small 

segmental tubercles set with simple hairs. Mummies are formed attached to 

leaves of the host trees (aspen or birch), sometimes in association with silk 

webbing, and the body of the mummy is slightly inclined away from the 

substrate. Drepana arcuata
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

69

Wasp distribution. New Jersey southward to Virginia and westward to Iowa and 

Louisianna.

Biology. Recorded as a solitary parasitoid of the drepanid Drepana arcuata (Walker) 

on alder and birch.

Similar species. Aside from color differences (this species is paler), A. platypterygis is 

virtually identical in appearance to A. gastritor. The two species can only be 

reliably identified by association with host caterpillars.

Best reference. Ashmead (1889).

Comments. Although A. platypterygis has been recognized as a distinct species for 

more than 100 years, it is not clear if it should be held as distinct from A. gastritor. Careful field studies are needed to determine if this is truely a sibling 

species.
70 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Aleiodes politiceps (Gahan)

Smooth-headed mummy-wasp

Diagnostic features of adult wasp. 

Body entirely orange but 

antenna, apex of hind tibia, 

and tarsi black. Wings 

black. Body length 7.0-9.0 

mm. Antenna with 60-65 

segments. Oral opening 

large and oval. Tarsal claws 

strongly pectinate. First, 

second, and basal half of 

third metasomal tergum 

with distinctive porcate 

sculpture, consisting of 

many longitudinal raised 

lines, surface appearing 

as though scratched by a 

comb.

Host caterpillars. The known hosts include noctuid larvae such as the green cutworm, 

Anicla infecta (Ochs.); and the armyworm moth, Mythimna (=Pseudaletia) 

unipuncta (Haw.).

Diagnostic features of host mummies. Mummies are about 1 cm long, entirely 

smooth, with a light yellowish-brown head capsule, and body mosty dark 

reddish-brown.

Wasp distribution. Found from Virginia southward to Florida and westward to Arkansas and Texas. Also recorded from Costa Rica.

Biology. This species attacks cutworm-type noctuid larvae in low mixed vegetation, 

grassy meadows, and forest understories. The adults are nocturnally active 

and commonly attracted to lights. Mythimna unipuncta
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

71

Similar species. Superficially similar to A. mandibularis, but this species can be distinguished by its smaller mandibles and black wings.

Best reference. Gahan (1917) and Marsh and Shaw (1999).

Comments. This is perhaps the most common Aleiodes species found in the southern 

states. The orange body with black wings and coarsely sculptured metasoma 

are highly distinctive. It can be reliably identified in the field, even without a 

microscope.
72 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Aleiodes preclarus Marsh and Shaw

Beautiful mummy-wasp

Diagnostic features of adult wasp. Body brightly colored with yellow, orange, and 

black markings. Antennal flagellum black. Body length 5.5-6.0 mm. Antenna with 52-55 segments. Ocelli large, ocell-ocular distance half the diameter 

of lateral ocellus. Occipital carina incomplete, interrupted on vertex. Fore 

wing vein r shorter than vein 3RSa. Hind wing vein RS parallel to wing margin on basal third, then suddenly curved downward. Hind wing marginal 

cell suddenly widened at apex. Apex of hind tibia with a row of flat setae on 

inner side.

Host caterpillar. A solitary parasitoid of the painted lichen moth, Hypoprepia fucosa 

Hbn., an arctiid that feeds on lichens.

Diagnostic features of host mummies. Mummies are about 13 mm long, black, dorsally with segmental raised tubercles set with black spines. These spines are 

brittle, and break easily as the mummy ages, so they may be missing if the 

mummy is weathered. Along the sides are smaller whitish bristles. Hypoprepia fucosa
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

73

Wasp distribution. Known only from the states of West Virginia, Virginia, and Kentucky.

Biology. This species has been reared from the lichen-feeding arctiid Hypoprepia 

fucosa from lichens found on white oak, red oak, hickory, and red maple. 

Caterpillars can be found by wrapping burlap bands around such trees with 

lichens. The caterpillars seek shelter under the bands, and will form mummies there.

Similar species. Similar to A. wahli but distinguished by its black antennal flagellum.

Best reference. Marsh and Shaw (1998).

Comments. The common name refers to the attractive body colors of this brightly 

patterned species. It is a resident of eastern oak, hickory, and maple deciduous forests where lichens are abundant.
74 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Aleiodes pseudoterminalis Marsh and Shaw

False terminal mummy-wasp

Diagnostic features of adult wasp. 

Body mostly two-colored 

with contrasting orange 

and black. Head and mesosoma black. Metasomal 

terga 1 orange or red, remaining metasomal terga 

black. Antenna brown. 

Hind tibia orange with 

black apex. Hind tarsomeres dark brown. 

Wings clear but lightly 

pigmented. Tegula yellow. Body length 7.0-8.0 

mm. Antenna with 56-59 

segments. Oral opening 

circular, diameter about equal to basal width of mandible. Malar space equal 

to basal width of mandible. Ocelli small, distance between eye and lateral 

ocellus about equal to ocellus width. Median length of pronotum shorter 

than distance between occipital carina and lateral ocellus. Mesopleuron 

smooth except subalar sulcus and sternaulus rugose. Fore wing vein 1cu-a 

beyond vein 1M by more than twice its length. Vein RS of hind wing straight, 

marginal cell of hind wing narrowest at base and widening toward apex. 

Hind wing vein m-cu present as short stub. Apex of hind tibia without a row 

of flattened setae along inner margin, setae normal and hair-like. Tarsal claws 

not pectinate. First metasomal tergum not parallel-sided, wider apically than 

at base. Apex of metasoma not compressed in female. Metasomal tergum 3 

costate at base, remainder smooth. Males with metasomal terga normal, not 

densely setose. Ovipositor at least two-thirds the length of hind basitarsus, 

sometimes nearly equal to basitarsus length.

Host caterpillars. Reared from the noctuid Eupsilia devia (Grote).
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

75

Diagnostic features of host mummies. Mummies induced by this species are 10-

12 mm long, very dark brown, and the cuticle is compacted and somewhat 

wrinkled at the anterior end, just behind the head capsule.

Wasp distribution. Found in the northeastern United States and eastern Canada from 

the Great Lakes area, southward to Tennessee and Virginia.

Biology. This species in a solitary parasitoid of the noctuid Eupsilia devia (Grote). 

Mummies are similar to those formed by A. terminalis but less wrinkled, and 

may be formed in sandy soil, not attached to the food plants.

Similar species. Very similar to A. terminalis, but differing by the black second metasomal segment (orange in A. terminalis).

Best reference. Marsh and Shaw (2001).

Comments. For many years this species was confused with A. terminalis. Eupsilia devia
76 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Aleiodes quadratus Shaw and Marsh

Quadrate mummy-wasp

Diagnostic features of adult wasp. 

Body mostly or entirely 

honey-yellow to light 

orange, except antenna 

partly to entirely dark 

brown to black. Wings 

clear to lightly pigmented. 

Stigma two-colored, dark 

brown to black medially, 

pale yellowish-white on 

basal third. Body length 

6.0-7.0 mm. Antenna with 

53-56 segments. Oral 

opening small and circular, 

diameter about equal to 

basal width of mandible. 

Malar space longer than basal width of mandible. Head rugulose dorsally. Ocelli small, 

distance between eye and lateral ocellus distinctly broader than ocellus width. Propodeum 

with small blunt tubercles on posterior corners. 

Second submarginal cell of fore wing short and 

quadrate, sometimes nearly square. Vein RS of 

hind wing slightly sinuate, marginal cell of hind 

wing narrowest in middle. Apex of hind tibia 

without a row of flattened setae along inner margin, setae normal and hairlike. Tarsal claws simple, not pectinate. First metsomal tergum not parallel-sided, wider apically than at base. Apex of metasoma not compressed in 

female. Metasomal terga 1-4 longitudinally rugose striate.

Host caterpillars. This wasp is recorded as attacking the arctiids Grammia blakei 

(Grote) and Spilosoma virginica (F.).

propodeal tubercle
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

77

Diagnostic features of host mummies. Mummies of Grammia are about 1 cm. long, 

black, and densely hairy.

Wasp distribution. This wasp is common in the eastern United States, and it occurs 

from Vermont south to Florida and westward to the Rocky Mountains.

Similar species. Aleiodes quadratus is similar to A. aciculatus, but can be distinguished 

by its short and quadrate second submarginal cell and by the presence of 

tubercles on the posterior corners of the propodeum. 

Biology. Not known in detail. Adult wasps have been collected from April through 

October, indicating that the species is long-lived as an adult or possibly is 

multivoltine.

Best reference. Shaw et al. (2006).

Comments. This species is very distinctive in having the second submarginal cell of 

the fore wing very short and quadrate, sometimes nearly square. Grammia blakei
78 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Aleiodes quebecensis (Provancher)

Quebec mummy-wasp

Diagnostic features of adult wasp. Body color mostly honey-yellow or light brown. 

Antenna usually black on basal half, yellowish-white to orange on apical half. 

Hind tarsomeres 1-4 brightly colored, yellowish-white or white. Tegula yellow. Body length 6.0-8.0 mm. Antenna long, with 45-55 segments. Flagellomeres all short and compact. Malar space very short, narrower than basal 

width of mandible. Oral space small and circular, about twice as wide as 

malar space. Ocelli large, ocell-ocular distance about one-third diameter of 

lateral ocellus. Occipital carina incomplete, weak or absent on vertex. Propodeum mostly granulate anteriorly, grading to rugose posteriorly. Tarsal 

claws strongly pectinate. Hind wing with vein RS slightly arched at apical 

two-thirds; marginal cell narrowest at apical two-thirds, then abruptly widening toward wing apex.

Host caterpillars. Reared from Acronicta furcifera Gn., Acronicta grisea Wlk., and 

Acronicta hasta Gn. Reared eastern specimens have also been associated with 

Prunus sordinia and with chokecherry, indicating that several other Acronicta

are potential hosts. David Wagner has collected mummies of Acronicta lobeliae from Quercus ilicifolia and Acronicta morula from elm, and these are 

consistent with the style of mummification caused by A. quebecensis, but the 

adult wasps did not emerge.

Diagnostic features of host mummies. Mummies are about 12-14 mm long and very 

firmly glued to the substrate, usually a leaf or twig. The head capsule is pale 

yellowish-white, while the body of the mummy is mostly dark brown to 
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

79

blackish. The body of the mummy is mostly smooth on top, but with some 

small setose tubercles and with some long setae or hairs around the lower 

margins. As the mummy forms the body shrinks, pulling the head capsule 

backward until it is tilted nearly face upwards. Thus, when viewed from 

directly above, the entire anterior face of the head capsule is clearly visible. 

When formed on a twig, these mummies are rather cryptic, somewhat resembling a bud or gall.

Wasp distribution. Although first discovered in Quebec, A. quebecensis is widely 

distributed from Quebec southward to Florida and westward through the 

northern states and Canada to Wisconsin, South Dakota, Oregon, and British 

Columbia.

Biology. The period of flight activity for adult wasps ranges from early June to midAugust. Wasps have been collected at lights and have very large ocelli, indicating that the species is nocturnally active. Aleiodes quebecensis is an arboreal species that appears to be oligophagous on young Acronicta caterpillars 

in several broadleaf trees.

Similar species. Most similar to A. geometrae, but A. quebecensis is very distinctive 

and can be distinguished from A. geometrae and all other members of the 

pulchripes species-group by the arched vein RS in the hind wing and the 

brightly colored yellowish or white hind tarsus.

Best reference. Shaw et al. (1997).

Comments. I’ve seen mummies formed both on leaves and on twigs, but the ones on 

leaves may have been lab-reared. Mummies formed on twigs are more cryptic than those on leaves. Acronicta lobeliae Acronicta hasta Acronicta morula Acronicta morula
80 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Aleiodes rileyi Cresson

Riley’s mummy-wasp

Diagnostic features of adult wasp. Body entirely honey-yellow or orange. Antenna 

black. Body length 5.5-8.0 mm. Antenna with 53-55 segments. Malar space 

about equal to basal width of mandible. Ocelli large, ocell-ocular distance 

equal to, or slightly less than, diameter of lateral ocellus. Tarsal claws strongly pectinate with seven to eight stout teeth. Marginal cell of hind wing narrowest in middle, widening toward wing apex. Males with terga 4-6 densely 

covered with golden setae (subdivided medially).

Host caterpillars. Recorded from a variety of caterpillars, including the noctuids 

Acronicta oblinita (J.E. Smith), Melanchra picta (Harr.), and Nephelodes minians Gn.; the lymantriid Dasychira vagans (B. and McD.); and possibly the 

pyralids Ostrinia obumbratalis (Led.) and O. penitalis (Grt.). Most of these 

records need confirmation. I’ve only seen associated mummies of the smartweed caterpillar (=smeared dagger moth), A. oblinita.

Diagnostic features of host mummies. Mummies of the smartweed caterpillar, Acronicta oblinita, are 15-17 mm long, mostly black, with lighter yellowishwhite along sides, and with mottled inter-segmental yellowish-white markings. The head capsule is black and widely separated from body of mummy 

by a very large glue-hole. Segments have bands of round, raised tubercles set 
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

81

with long yellowish-white spines, but these tend to break off as the mummy 

ages. There are many long yellowish-white spines compressed into the area 

behind the head capsule. Mummies vary greatly in the number of spines remaining and in the extent of dark color. Light-colored parts appear to darken 

with age. The loss of spines on mummies and darkening of light colors 

indicate aging of the mummies and exposure to weather. Probably, the wasp 

over-winters inside a mummified host caterpillar.

Wasp distribution. Recorded from Connecticut southward to Florida and westward 

to Michigan, Kansas, and Saskatchewan. Probably present through most of 

the eastern United States.

Biology. All the recorded hosts are generalist-feeders on a variety of low vegetation, 

shrubs, and small trees, including Prunus and willow. Adult wasps are active 

in Florida as early as January, but in northern parts of its range, the wasp flies 

in late summer. It appears to be a polyphagous species, utilizing several unrelated hosts on a variety of low plants. In Illinois, the wasp has been found in 

pitcher plants.

Similar species. Aleiodes rileyi is a member of the apicalis species-group, distinguished 

by their strongly pectinate claws and males with dense setae on the posterior 

metasomal terga. Aleiodes rileyi is quite distinctive in being the only member 

of this group that is entirely orange, without black markings on the body.

Best reference. Shaw et al. (1998a)

Comments. This wasp is named after Charles Valentine Riley, one of the most famous 

American entomologists of the late 1800s. Acronicta oblinita
82 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Aleiodes rugulosus (Nees)

Wrinkled mummy-wasp

Diagnostic features of adult wasp. 

Body color almost entirely 

black except for some orange 

on legs and sometimes at base 

of metasoma. Body length 8 

mm. Antenna with 65-70 segments. First two metasomal 

segments extremely coarsely 

sculptured.

Host caterpillars. In Europe this species attacks Oxicesta geographica, Simyra albovenosa, and 

several species of Acronicta. 

Its hosts in the northeastern United States are not yet 

known, but presumably the host range would include these and perhaps 

other related genera of acronictine Noctuidae. In Canada, this species has 

been reared from Acronicta distans and A. oblinita.

Diagnostic features of host mummies. Mummies formed by this species are 1.0-1.5 

cm long, with black head capsules and numerous long, whitish hairs arising 

from raised segmental tubercles.

Acronicta oblinitaAcronicta distans

Acronicta rumicis
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

83

Wasp distribution. This species is native to Europe, but is now found in Canada and 

the northeastern United States. 

Biology. Details of the biology in Europe are given by Mark Shaw (2002). The species is oligophagous on several acronictine noctuid caterpillars, only found on 

low plants.

Similar species. This species has extremely coarse metasomal sculpture and cannot be 

confused with any other North American species. It is the only member of 

the rugulosus species-group that occurs in North America, so it can be keyed 

to species using the species-group key provided by Shaw et al. (1997).

Best reference. Shaw et al. (1997) and Shaw (2002).

Comments. Aleiodes rugulosus was studied in Europe during the early years of the 

gypsy moth problem. Apparently, material was also studied in New England, but there is no record of releases. Subsequent collections in eastern 

North America indicate that it may have been released, or perhaps the species has a natural holarctic distribution, but has only been recently noticed in 

North America.
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Aleiodes sanctihyacinthi (Provancher)

Saint Hyacinth mummy-wasp

Diagnostic features of adult wasp. Body mostly two-colored orange and black. Eye 

posteriorly margined with orange. Pterostigma entirely dark brown. Body 

length 4.5-5.5 mm. Antenna with 46-48 segments. Top of head with rough 

rugose sculpture. Fourth metasomal tergum with fine granular sculpture.

Host caterpillar. Hyphantria cunea (Drury).

Diagnostic features of host mummies. The mummified caterpillars are 10-12 mm 

long, with a black head capsule, and a body with dark blackish dorsal longitudinal band and yellowish sides. There are numerous segmental round 

tubercles, each set with long, mostly white hairs. The resulting mummy is 

densely hairy, but the body color is still easily visible. Mummies are found 

inside the silk shelters of the host. Hyphantria cunea

Hyphantria cunea
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85

Wasp distribution. Found in northeastern North America (New Brunswick, Nova 

Scotia, Quebec, and Maine) westward to Wisconsin, Wyoming, and Utah. 

The southern extent of its range is not known, but A. sanctihyacinthi probably occurs throughout the range of its host caterpillar. This species has been 

introduced to Yugoslavia for biological control under the name Rogas hyphantriae.

Biology. This species is an apparently monophagous parasitoid of the fall webworm 

caterpillar, Hyphantria cunea (Drury). The host caterpillar is a common eastern pest known to feed on more than 100 tree species, including ash, hickory, 

maple, oak, walnut, and apple. The caterpillars form large communal silk 

webs near the tips of branches, and the mummified caterpillars can be found 

inside these silk shelters. Dunstan (1922) studied the biology of this species 

and found some silk webs with 30-40 percent of all caterpillars parasitized. 

He noted that young caterpillars, about 10 days old, are attacked. The caterpillars are mummified during the fourth larval instar, shrinking to 10-12 

mm long. Adult wasps emerge after about two weeks. Dunstan noted that, 

because the season is well-advanced by the time adults emerge, they probably 

overwinter in the adult stage. He also speculated that, since a long period of 

time must elapse between the emergence of adults in the fall and egg-deposition into Hyphantria during the following summer, the species might well 

have another host in the spring. If so, it has not yet been discovered. During 

the introduction of this species to Europe for biological control purposes, the 

biology of this species was studied by Bogavac (1955) and Tadic (1959) under 

the name Rogas hyphantriae.

Similar species. Aleiodes sanctihyacinthi is similar to A. aciculatus (both species have 

rough sculpture on the top of the head). It can be distinguished from A. 

aciculatus by its entirely brown pterostigma and by the granular sculpture of 

the fourth metasomal tergum.

Best references. Gahan (1922) and Dunstan (1922) (as Rogas hyphantriae).

Comments. The scientific and common names for this species refer to the fact that it 

was first discovered in St. Hyacinth, Quebec (Provancher 1880). Most of the 

historical literature refers to this species as Rogas hyphantriae.
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Aleiodes scrutator (Say)

Yellow-marked mummy-wasp

Diagnostic features of adult wasp. 

Body two-colored: honeyyellow with dark brown 

markings. Metasoma dorsally dark brown with a large 

yellow or yellowish-white 

medial spot. Pterostigma 

two-colored: brown medially, yellow at base and apex. 

Body length 4.5-6.0 mm. 

Antenna with 37-45 segments. Ocelli rather small, 

ocell-ocular distance equal to 

or wider than ocellus width.

Host caterpillars. Recorded (based on museum specimens) as attacking the choreutid 

Caloreas augustella (Clarke), the tortricid Endopiza viteana (Clemens), the 

geometrid Lambidina fervidaria athasaria (Walker), the notodontid Clostera 

inclusa (Hübner), and the noctuid Plathypena scabra Fabricius.

Diagnostic features of host mummies. Some mummies formed by this species are 

small (about 8 mm long), smooth, and rather non-descript yellowish-brown.

Wasp distribution. This species is widely distributed across the United States and 

southern Canada. It appears to be more common in the northern parts of its 

range and at higher elevations.

Biology. Not studied in any detail, but the recorded host records seem to indicate that 

this species is widely polyphagous on a variety of small caterpillars feeding 

on many trees, including aspen, poplar, white oak, and willow. prob. Platypena scabra
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

87

Similar species. Most similar to the European species Aleiodes circumscriptus. 

Aleiodes scrutator is the only common member of the circumscriptus species-group in eastern North America, and in some forest conditions, it can be 

quite common. It can be identified using the species-group key provided by 

Shaw et al. (1997).

Best reference. Shaw et al. (1997).

Comments. All the host records for this species need confirmation.
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Aleiodes simillimus (Ashmead)

Similar mummy-wasp

Diagnostic features of adult wasp. Body color mostly honey-yellow, except ocellar 

triangle, propodeum, base of tergum 1, and sides of terga 2+3 dark brown to 

black. Stigma two-colored but with darkened medial area not discrete, rather 

translucent, with dark color grading into lighter basal and apical areas. Body 

length 4.0-5.0 mm. Antenna with 40-45 segments. Eyes and ocelli large, ocellocular distance about half the diameter of lateral ocellus. Hind wing with 

vein RS arched in the middle, marginal cell 

narrowest near middle.

Host caterpillars. Recorded from several small arboreal geometrids, including Narraga fimentaria Walker, Semiothisa cyda (Druce), S. colorata Grote, Tornos scolopacinarius (Guenée), Glena cribrataria (Gueneé), Itame pustularia (Guenée), 

Erannis tillaria (Harr.), and Lomographa vestaliata (Guenée).

Diagnostic features of host mummies. Mummies about 9-10 mm long, mostly light 

whitish to yellowish-brown, with fine mottled black markings and tranverse 

black bands in prothoracic area. Smooth and slightly tapered at both ends.

Erannis tillariaSemiothisa sp.
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

89

Wasp distribution. This species is found throughout the eastern United States from 

New Hampshire south to Florida and westward to Michigan and Texas.

Biology. Aleiodes simillimus is a highly polyphagous parasitoid of small arboreal Geometridae caterpillars.

Similar species. As the name implies, this is a rather non-descript small yellowish species that is quite similar to several others in the gastritor species-group. In 

the east, it is most similar to A. gastritor, but can be separated by its larger 

ocelli and longer antennae. It is also similar to A. buoculus Marsh, a parasitoid of geometrid caterpillars on jojoba, but that species only occurs in the 

southwestern United States (Marsh 1989).

Best references. Ashmead (1889), Marsh (1989).

Comments. This species is very similar to A. gastritor, and careful field studies of the 

host ranges are needed.
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ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Aleiodes smithi Marsh and Shaw

Smith’s banded mummy-wasp

Diagnostic features of adult 

wasp. Body color 

mostly honey-yellow, 

sometimes with brown 

markings on dorsum 

of mesosoma. Antennal flagellum black with 

white band consisting 

of 5-12 segments. Body 

length 6-8 mm. Antenna 

with 44-50 segments. 

Oral opening oval, diameter equal to or slightly 

greater than malar space. 

Malar space equal to or 

wider than mandible 

width. Ocelli small. Tarsal claws simple, not pectinate. Hind wing marginal 

cell narrowest at base, widening to apex.

Host caterpillars. Reared from the noctuid Lascoria ambigualis Walker.

Diagnostic features of host mummies. Mummies are about 1 cm long, with the body 

of the mummy mostly very smooth and very dark reddish-brown, almost 

black. The head capsule is brown, mottled with light yellowish-white irregular spots. Behind the head, there is a short prothoracic collar that is narrower 

than the head or the remainder of the mummy, giving the appearance of a 

short neck. The front end of the mummy is densely wrinked. The wrinkles 

are lighter yellow, and under close inspection, give the appearance of many 

fine yellow cross-bands. Lascoria ambigualis
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91

Wasp distribution. This species occurs widely through the eastern United States. It 

has been found from Ontario and Massachusetts southward to Florida and 

westward to Wisconsin and Missouri. It also occurs in Mexico and Costa 

Rica.

Biology. The only known host is the ambiguous moth, Lascoria ambigualis, a caterpillar known to feed on ragweed, chysanthemum, and horseradish. This species 

is found in forest meadows, understory vegetation, and along streambeds.

Similar species. This species is most closely related to A. ecuadoriensis Brues, a South 

American species that does not occur in North America.

Best reference. Marsh and Shaw (2001).

Comments. Aleiodes smithi is the only Aleiodes species in the eastern United States 

that has a white-banded antenna, so it can be easily identified even without a 

microscope. The species is named after sawfly researcher David Smith, who 

collected many specimens of this species in Virginia.
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Aleiodes stigmator (Say)

Stigmata mummy-wasp

Diagnostic features of adult wasp. Body color mostly yellowish orange, except metanotum and propodeum brown to black. Metasoma usually entirely orange, 

but sometimes with dark brown or black basally on first tergum. Stigma 

two-colored, translucent brown to black medially, pale yellowish-white basally and apically. Body length 4.0-5.0 mm. Antenna with 31-32 segments. 

Frons, temple, and vertex with fine granular sculpture. Malar space longer 

than basal width of mandible. Tarsal claws weakly pectinate basally, appearing simple. First metasoma tergum wider at apex than long. Hind wing with 

vein RS arched in the middle, marginal cell narrowest near middle.

Host caterpillars. Host range includes several acronictine noctuids, including Acronicta americana Harris, A. dactylina (Grote), and A. hastulifera (Smith), as well 

as the notodontid Simyra henrici Grote. I’ve also seen a Catocala mummy 

that appeared to be attacked by this wasp species.

 Previous records of this wasp species attacking the geometrid Alsophila pometaria are erroneous and actually refer to Aleiodes gastritor. Previous records 

of this wasp species as attacking Malacosoma species are also erroneous and 

refer to the species now known as Aleiodes malacosomatos.
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

93

Diagnostic features of host mummies. The mummies vary in appearance depending on the species, but all are exceptionally large for Aleiodes, ranging from 

2-3.5 cm. in length. Most are hairy mummies, but the hairs may fall off as 

the mummy is exposed to weather. Once the wasps emerge, there can be no 

doubt about the identity of the species as this is the only gregarious Aleiodes

species in eastern North America. It is not unusual for 40-50 wasps to 

emerge from a single mummy. The emergence holes are evenly spread across 

the top of the mummy, from front to back, leaving a quite distinct appearance, as though pelted evenly by shotgun pellets.

Wasp distribution. This species is widely distributed across Canada and the northern 

United States. It is associated with marsh areas and is usually found near 

plants such as willows and cattails.

Simyra Acronicta americana

henrici Acronicta dactylina

A. stigmator emergingfrom Acronicta dactylina

Acronicata dactylina
94 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Biology. Since the work of Mason (1979), this species has been interpreted as a polyphagous strictly gregarious parasitoid utilizing relatively large caterpillars. 

Hosts attacked include several acronictine noctuids such as Acronicta americana Harris, A. dactylina (Grote), and A. hastulifera (Smith), as well as the 

notodontid Simyra henrici Grote and some Catocala species. Adult wasps 

emerge in late summer or early autumn. Whether they overwinter as adults 

or have another host caterpillar for overwintering is still unknown.

Similar species. Aleiodes stigmator is quite similar morphologically to A. malacosomatos, but has less coarse and more shining sculpture on the head and mesosoma, and the fore wing stigma is less distinctly two-colored. The habits of the 

two species are quite different, as A. stigmator is a gregarious parasitoid of 

much larger noctuid and notodontid caterpillars. This is the only gregarious 

Aleiodes species in the eastern United States, so when reared from caterpillars, it is quite easy to identify.

Best reference. Mason (1979).

Comments. This is the oldest known North American Aleiodes species, having been 

discovered by the first American entomologist, Thomas Say, in 1824. Despite 

this long history, the biology of the species has never been carefully studied, 

and much could be learned by detailed field research. The name “stigmata 

mummy-wasp” is a reference to the exit holes that are produced in the host 

caterpillar. Thomas Say thought they looked like the stigmata in the hands 

and feet of the Christ.
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95

Aleiodes terminalis Cresson

Terminal mummy-wasp, the Terminator

Diagnostic features of adult wasp. Body mostly two-colored: contrasting orange and 

black. Head and mesosoma black. Metasomal terga 1-2 (sometimes tergum 

3 partly) orange; remaining metasomal terga black. Antenna brown medially 

to black apically. Hind tibia black with light yellowish-white band on basal 

third. Hind tarsomeres black. Wings clear but lightly pigmented. Tegula 

yellow. Body length 5.0-8.0 mm. Antenna with 55-62 segments. Oral opening circular, diameter about equal to basal width of mandible. Malar space 

equal to basal width of mandible. Ocelli small, distance between eye and 

lateral ocellus about equal to ocellus width. Median length of pronotum 

shorter than distance between occipital carina and lateral ocellus. Mesopleuron smooth except subalar sulcus and sternaulus rugose. Forewing vein 1cu-a 

beyond vein 1M by more than twice its length. Vein RS of hind wing straight, 

marginal cell of hind wing narrowest at base and widening toward apex. 

Hind wing vein m-cu present as short stub. Apex of hind tibia without a row 

of flattened setae along inner margin, setae normal and hair-like. Tarsal claws 

not pectinate. First metasomal tergum not parallel-sided, wider apically than 

at base. Apex of metasoma not compressed in female. Metasomal tergum 3 

costate at base, remainder smooth. Males with metasomal terga normal, not 

densely setose.
96 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Host caterpillars. Attacks several species of “cutworm-like” Noctuidae larvae, including Mythimna (=Pseudaletia) unipuncta (Haworth), Nephelodes minians

Guenée, Spodoptera frugiperda (Smith), S. ornithogalli (Guenée), and Xestia 

smithii (Snellen).

Diagnostic features of host mummies. Mummies induced by this species are 10-

12 mm long, very dark brown, and the cuticle is compacted and strongly 

wrinkled at the anterior end, just behind the head capsule. The head capsule 

sometimes falls away in older mummies exposed to weather. When formed 

on a twig or small branch, the mummy is highly cryptic as it resembles a bud 

or stem gall.

Wasp distribution. This species is widely distributed across Canada and the continental United States as far south as Mexico. It is very common in eastern forests 

and probably the most frequently encountered Aleiodes species in the East.

Biology. The biology of this species was studied by Pennington (1916). In Maryland, 

A. terminalis usually has four, and possibly as many as six, generations per 

year. Adult wasps emerge about 17 days after the mummy is formed. Mating takes place immediately after emergence, and females are able to begin 

laying eggs immediately after mating. Females live for an average of 32 days, 

but may live as long as 75 days. Males only live about half as long as females, 

averaging about 16 days. Females oviposit only into second or third instar 

hosts larvae. Overwintering is inside mummified host.

Similar species. Superficially similar to other large red and black Aleiodes species, such 

as A. abdominalis, A. pseudoterminalis, and A. ufei. Aleiodes terminalis can 

be easily distinguished from similar species by the light colored band on the 

anterior third of the hind tibia.

Best reference. Marsh and Shaw (2001).

Comments. Adults of this species are nocturnally active and commonly collected at 

lights. Perhaps the most commonly encountered Aleiodes species in the eastern states. Noctuidae sp.
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

97

Aleiodes texanus Cresson

Texas mummy-wasp

Diagnostic features of adult wasp. 

Body two-colored: black 

and reddish orange. Metasoma entirely reddish orange. Wings dusky. Body 

length 7.0-8.5 mm. Antenna with 58-62 segments. 

Malar space short, about 

equal to mandible width. 

Eyes and ocelli large. 

Ocell-ocular distance narrow, lateral ocellus near 

eye margin. Lateral ocellus one to two times wider 

than ocell-ocular distance. 

Hind wing vein RS sinuate and strongly arched, RS approaching wing margin medially. Hind wing 

marginal cell very narrow in middle.

Host caterpillars. In the East this species has been reared from the walnut sphinx, 

Amorpha (=Laothoe) juglandis (J.E. Smith); the hog sphinx, Darapsa myron

(Cramer); the Nessus sphinx, Amphion nessus floridensis B.P. Clarke; and 

other unidentified Sphingidae. In Arizona, this species has been reared from 

Ceratomia sonorensis.

Diagnostic features of host mummies. About 12-15 mm long, smooth, pale brownish-tan with with dark brown bands, and densely covered with tiny spinules. 

Broadly attached to the substrate by an antero-ventral glue-hole. Head capsule either decapitated from mummy or attached by a small piece of cuticle 

that is prone to breaking away as the mummy ages. Has a a distinct posterior 

caudal horn about 4 mm long or a distinct pit if the caudal horn has fallen off.

Ceratomia sonorensisDarapsa myron
98 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Wasp distribution. The name refers to the fact that the species was first discovered in 

Texas. It is actually very widely distributed in the eastern United States from 

Michigan and Massachusetts southward to Florida and the Gulf Coast and 

westward to Montana, Texas, and northern Mexico.

Biology. Mummies are firmly glued down by a very large antero-ventral glue-hole, 

which usually results in decapitation of the caterpillar. Thus, wild-collected 

mummies usually lack a head capsule. The caudal horn of sphingid hosts is 

quite distinctive, but may fall off as the mummy ages or overwinters, leaving a visible pit. Adult wasps are arboreal, attracted to lights, and probably 

nocturnally active. Aleiodes texanus is associated with sphingid hosts on 

several broadleaf trees and vines, including walnut, ash, and grape. The wasp 

is probably univoltine in the North and multivoltine in the South. 

Similar species. Most closely related to A. ceratomiae, but A. texanus is easily distinguished by its smaller ocelli and bright reddish orange metasoma.

Best reference. Shaw et al. (1998b).

Comments. Aleiodes texanus is one of the few species that is large enough and with 

sufficiently distinctive markings that it can easily be spot-identified even 

without magnification. The black and orange body and blackish wings are 

distinctive.
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99

Aleiodes vierecki Marsh and Shaw

Viereck’s mummy-wasp

Diagnostic features of adult wasp. Body color entirely honey-yellow. Wings clear. 

Tegula yellow. Body length 7.0-8.5 mm. Malar space equal to basal width 

of mandible. Temple narrow, about half the eye width. Oral opening small, 

circular. Diameter of lateral ocellus equal to, or slightly greater than, ocellocular distance. Tarsal claws not pectinate. Basal third of hind wing vein RS 

nearly parallel to wing margin, then curving downward, marginal cell suddenly widening.

Host caterpillars. Unknown.

Diagnostic features of host mummies. Unknown.

Wasp distribution. Known from Kansas, Oklahoma, and Michigan.

Biology. The adult wasp flies early in the summer, from April to mid-June and is attracted to lights.

Similar species. This species is similar to A. atricornis, a common species of midwestern grasslands and prairies. It differs from A. atricornis by having larger 

ocelli, narrow temple, and hind wing RS vein slightly curved.

Best reference. Marsh and Shaw (2001).

Comments. Aleiodes vierecki is a midwestern species that is not common in the East. 

In the East, it has only been collected from central Michigan. The wasp is 

named after H. L. Viereck, a student of the Braconidae who worked in the 

early 1900s.
100 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Aleiodes virginiensis Marsh and Shaw

Virginia mummy-wasp

Diagnostic features of adult wasp. 

Entire body including 

antenna and legs entirely 

honey-yellow to light 

orangish-brown, without 

black markings. Tegula 

yellow. Fore wing stigma 

two-colored, brown medially with yellow at base 

and apex. Body length 

7.0-8.0 mm. Antenna with 

58-63 segments. Ocelli 

large, ocell-ocular distance 

half the diameter of lateral 

ocellus. Occipital carina 

incomplete, interrupted on 

vertex. Fore wing vein r shorter than vein 3RSa. Hind wing vein RS parallel 

to wing margin on basal third, then suddenly curved downward. Hind wing 

marginal cell suddenly widened at apex. Apex of hind tibia with a row of flat 

setae on inner side.

Host caterpillars. Unknown.

Diagnostic features of host mummies. Unknown.

Wasp distribution. So far, only known from Essex County, Virginia.

Biology. The caterpillar hosts of this wasp species are not yet known. In Virginia, 

adult wasps fly from mid-July to mid-September.

Similar species. This species is similar to A. preclarus and A. wahli, but can be distinguished by its entirely honey-yellow to orangish-brown body without black 

markings.

Best reference. Marsh and Shaw (1998).

Comments. According to David Smith, the collector of the type-series, the area where 

this species was found is dominated by mixed hardwoods and pine (mostly 

tulip tree, sweet gum, oaks, hickories, and Pinus taeda) in the coastal plain, in 

mostly sandy soils.
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

101

Aleiodes wahli Marsh and Shaw

Wahl’s mummy-wasp

Diagnostic features of adult wasp. Antennal flagellum yellow. Mesosoma yellow with black 

or dark brown markings. Tegula brown. 

Fore wing stigma two-colored, brown medially with yellow at base and apex. Hind 

coxa brown on lower end. Hind femur 

brown on apical three-fourths. Metasoma

light yellowish-brown, without black 

markings. Body length 5.5-6.0 mm. Antenna with 56-59 segments. Ocelli large, 

ocell-ocular distance half the diameter of 

lateral ocellus. Occipital carina incomplete, 

interrupted on vertex. Fore wing vein r 

shorter than vein 3RSa. Hind wing vein 

RS parallel to wing margin on basal third, 

then suddenly curved downward. Hind 

wing marginal cell suddenly widened at 

apex. Apex of hind tibia with a row of flat 

setae on inner side.

Host caterpillars. Unknown.

Diagnostic features of host mummies. Unknown.

Wasp distribution. Known so far only from localities in Florida and North Carolina.

Biology. The caterpillar hosts of this wasp species are not yet known. In Florida, the 

adult wasps have been collected on dates in late August through early January.

Similar species. This species is similar to A. virginiensis and A. preclarus, but can be 

distinguished by its yellow antenna.

Best reference. Marsh and Shaw (1998).

Comments. Most of the collecting records for this species are from hammocks of 

southern Florida. The species is named after Dr. David Wahl of the American Entomological Institute in Gainesville.
102 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

PHOTOGRAPHY CREDITS

Aleiodes indiscretus attacking gypsy moth larva: photo by Scott Bauer, USDA/ARS Image 

Gallery, image number K7656-1

Aleiodes stigmator parasitizing Acronicta americana: photo by David L. Wagner, University 

of Connecticut, DLW Lot 2004H97.

Aleiodes aciculatus mummy Euchlaena serrata, Aleiodes albitibia mummy Nerica sp., 

Aleiodes pallidator mummy Dasychira sp., Aleiodes sanctahyacinthi mummy Hyphantria 

cunea (left-hand photograph on page 84), Aleiodes simillimus mummy Erannis tillaria, 

and Aleiodes smithi mummy on Lascoria ambigualis (right-hand photograph on page 90): 

photos by Daniel Dalton, University of Wyoming.

All other photographs: Scott Shaw, University of Wyoming.

GLOSSARY

abdomen The posterior section of an insect body. In Hymenoptera, including parasitoid wasps, the apparent “abdomen” is usually referred 

to as the “metasoma” because part of the true abdomen is fused 

to the thorax to form the middle section of the body, the “mesosoma.”

aciculate Microscopic surface sculpturing resembling fine longitudinal lines, 

appearing as though scratched by a needle. Seen, for example, on 

the metasomal terga of A. aciculatus under high magnification.

annulus A light-colored band, usually white or yellowish-white, on the 

antenna. Only one eastern U.S. species (A. smithi) shows this 

characteristic, but it is common in some tropical Braconidae.

arboreal Tree-dwelling. Some Aleiodes species attack hosts living in trees, 

while others attack low-dwelling hosts in the understory or among 

herbaceous plants.

banding A method for collecting caterpillars in which bands of material, 

paper, or cardboard are wrapped around a tree trunk to create protected shade and attract caterpillars. Useful for sampling caterpillars that feed at night and hide during the day on tree-trunks or in 

the leaf-litter.

beating sheet A square of material supported by a wooden-frame, used for collecting caterpillars. The sheet is placed under a branch that is then 

shaken or beaten with a stick or net-handle to dislodge caterpillars, 

which fall on the sheet. Useful for dislodging mummies. A lightcolored umbrella works just as well.
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

103

body length Size of an Aleiodes wasp measured from the front of the head to 

the end of the metasoma, excluding the antenna and ovipositor.

 broad host range Indicates when a parasitoid wasp is able to use many different caterpillar species as hosts.

caudal horn A large spine-like structure on the posterior end of caterpillars 

of the moth family Sphingidae, which is a characteristic feature 

of sphingid larvae (horn-worms). This feature is preserved in 

sphingids mummified by Aleiodes of the praetor species-group (A. 

ceratomiae and A. texanus).

clypeus Lower part of the head just below the face and just above the oral 

opening. The upper corners of the clypeus are marked by the anterior tentorial pits, which appear as small round holes.

 compressed metasoma The posterior section of the wasp’s body is pressed together from 

side to side and so appears narrow when viewed from above. This 

condition is regarded as a potential adaptation for attacking host 

caterpillars that are concealed in narrow spaces, such as in leaf 

rolls.

concave labrum The labrum is the lower part of the head just above the mouth and 

below the clypeus. In Aleiodes, the labrum is concave and forms 

the back wall of the (scooped out) oral opening.

costate Microscopic surface sculpturing with longitudinal raised ridges.

death-wasp A common name for any species in the parasitic wasp family 

Braconidae.

 endoparasitoid A parasitic larva, such as Aleiodes, that feeds inside the body of its 

host caterpillar.

 first metasomal tergum The first segment of the apparent abdomen of a wasp, as viewed 

from above. This body part varies in shape, color, and surface 

sculpturing, and so provides useful diagnostic features for several 

Aleiodes wasp species.

frass Caterpillar droppings.

 hind wing vein RS The radial sector vein of the hind wing, which is located in the outermost upper section of the wing. In Aleiodes species, the RS vein 

varies in shape and direction. Sometimes, it is gently sinuate (Sshaped), in which case the marginal cell appears narrowest toward 
104 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

the middle. Sometimes, the RS is nearly straight and oriented toward the lower outer edge of the wing, in which case the marginal 

cell is broader as it approaches the wing margin. See the diagram 

at the end of the Glossary.

 host caterpillar A lepidopterous larva that is attacked by a parasitoid and serves as 

its food source.

host plant The plant on which a host caterpillar feeds, so the parasitoid species is also associated with it.

host range The set of caterpillar species (one to several) that a parasitoid wasp 

is able to attack successfully in a natural setting.

 ichneumon-fly An old-fashioned common name, formerly used for wasp species 

of the family Ichneumonidae and Braconidae. Not used much 

since the early 1900s, but appearing in much older literature.

koinobiont A parasitoid species, as in Aleiodes wasps, whose attacking parasitoid adult injects an egg but does not permanently paralyze the host 

larva. This allows for the attack of rather small host caterpillars. 

After being parasitized, the host caterpillar continues to feed and 

grow, often acting and appearing normal for some period of time 

before the parasitoid larva completes its development and emerges.

lateral ocellus Either one of the simple eyes on the top of the head that is situated 

laterally, closest to the margin of the compound eye.

Malaise trap A type of flight-intercept trap that is commonly used for collecting adult wasps. It consists of several panels of fine-mesh material 

topped by a tent-like portion that directs flying insects into a collecting jar.

malar space The space between the lower margin of the compound eye and the 

mandible. Usually measured relative to the width of the mandible 

at its base.

 marginal cell of hind wing The outermost area of the hind wing along the leading (anterior 

margin) that is bordered by the radius (R) and radial sector (RS) 

veins. Because the RS vein varies in shape and orientation, the 

marginal cell appears to have different shapes in different Aleiodes

species. Usually it has one of two shapes: broad and gradually 

wider toward the wing apex or very narrow near the middle as the 

RS vein curves toward the leading wing margin. See the diagram at 

the end of the Glossary.
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

105

m-cu vein Small cross-vein located in the lower middle area of the hind wing 

of Aleiodes wasps. It may be present or absent, depending on the 

species. See the diagram at the end of the Glossary.

mesopleuron In wasps, the section of the middle body located below the front 

wing and above the middle leg. Surface sculpturing of this body 

part varies among different Aleiodes species.

mesosoma The middle section of a wasp’s body (apparent thorax), where the 

legs and wings are attached. In bees, ants, and wasps, this area is 

called the “mesosoma” rather than “thorax” because it consists of 

the true thorax and also part of the abdomen that is fused to form 

the propodeum at the end of the mesosoma.

metasoma The posterior section of an insect body. In Hymenoptera, including parasitoid wasps, the apparent “abdomen” is usually referred 

to as the “metasoma” because part of the true abdomen is fused 

to the thorax to form the middle section of the body, the “mesosoma.”

metasomal terga The segmental plates on the top posterior part of a wasp’s body. In 

Aleiodes species, these terga vary in size, shape, color, density of 

setae, and surface sculpture, and so provide many useful diagnositic features.

multivoltine Having more than one generation per year. Some Aleiodes species 

may have four to six generations per year.

 mummification The process of forming a caterpillar mummy.

mummy A caterpillar that has been parasitized by an Aleiodes wasp larva. 

The remains of the host shrink somewhat but remain inflated and 

intact (mummified). The host remains, if protected from weather, 

do not degrade further and can be kept indefinitely in a museum 

collection without preservatives.

 mummy-wasp A common name for any wasp in the genus Aleiodes.

 narrow host range Indicates when a parasitoid wasp is able to use only one or a few 

caterpillar species as hosts.

ocell-ocular distance The shortest distance between a lateral ocellus and the margin of 

the compound eye. The ocell-ocular distance is commonly measured relative to the width of a lateral ocellus.
106 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

ocellar triangle The triangular area on the top of the head circumscribed by the 

three ocelli (simple eyes). In some species (e.g., A. nigristemmaticum), this area may be a different color than the remainder of the 

head. 

ocelli (singular: ocellus) Three simple eyes located on the top of the 

wasp’s head. The middle ocellus is the median ocellus. The other 

two are the lateral ocelli.

 occipital carina A narrow rim or microscopic ridge that borders the back side of 

the head in Aleiodes wasps. If the side parts of the carina meet at 

the top of the head, the occipital carina is said to be “complete.” 

If the side parts are separated, not meeting at the top of the head, 

then the occipital carina is said to be “broken” or “incomplete.” 

The carina can be complete, narrowly broken, or widely broken, 

depending on the wasp species.

oral opening Not the true mouth, but the concave space formed by the lower 

margin of the clypeus, the concave labrum, and the mandibles. In 

Aleiodes wasp species, this space varies in shape (circular to oval) 

and size depending on the wasp species.

ovipositor The egg-laying device and “stinger” of a female wasp. Aleiodes

females use this to inject temporary venoms and eggs. Aleiodes do 

not sting humans.

oviposition The process of injecting an egg into a host caterpillar via the ovipositor.

pectinate Comb-like: referring to the condition of the tarsal claws when 

there is a microscopic set of extra spines or setae, giving the claw a 

comb-like appearance under high magnifications.

porcate Coarse microscopic surface sculpturing resembling several raised 

parallel ridges separated by distinct furrows. Seen, for example, 

on the metasomal terga of A. politiceps, easily visible even at low 

magnifications.

pronotum The most anterior upper part of the mesosoma, located just behind 

the head. This structure varies in size and shape in several Aleiodes

species.

propodeum The most posterior of the four segments of the mesosoma. The 

propodeum is actually the first true abdominal segment, which is 
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

107

fused to the thorax and separated from the rest of the abdomen by 

a constriction.

Rogas Name of a genus of wasps closely related to Aleiodes and the typegenus of the subfamily Rogadinae in which Aleiodes is classified. 

Many North American Aleiodes species were formerly classified as 

Rogas species, so the name occurs frequently in much of the older 

literature.

rugose Microscopic surface sculpturing that is wrinkled, rough, and irregular.

setae Hair-like or bristle-like structures of the insect cuticle. Unlike 

mammalian “hairs,” these are skeletal features, and they do not 

grow.

sinuate Gently ‘S’-shaped, curving like a snake: refers to the condition of 

the hind wing RS vein in certain species.

sternaulus Located on the side of the body above the middle leg, this is a 

distinct area of the mesopleuron that is sometimes indicated by a 

patch of rough sculpturing. The presence and texture of sculpturing or its absence varies among species.

stigma A large prominent expanded area in the fore wing venation, along 

the leading edge of the wing. See the diagram at the end of the 

Glossary.

 subalar sulcus A sculpture depression just below the base of the front wing on 

the upper side of the body: the upper mesopleuron.

tarsal claws A claw-like structure located at the tip of each leg, at the end of the 

fifth tarsomere. The true claw is at the tip of the leg. Below the 

tarsal claw, there may be several additional setae, spines, pegs, or 

teeth forming a pectin, or comb-like structure, on the lower side 

of the claw. These features, although microscopic, vary greatly 

among Aleiodes species and provide valuable diagnostic features. 

If the claw is not visibly pectinate, then it is called “simple.”

tarsomeres One of the five small segments of the tarsus, the end section of a 

wasp’s leg, between the tip of the tibia (where the two tibial spurs 

are situated) and the tarsal claws at the apex of the leg.

tegula Small flap located over the base of the front wing in wasps. In 

Aleiodes species, the tegula varies in color.
108 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

tibia The fifth, longest segment of an Aleiodes wasp leg. The hind tibia 

provides useful diagnostic features for some species, including 

color variation and variation in the form of setae at the inner apex.

univoltine Having only one generation per year.

vein r A small vein of the fore wing that emerges from the lower middle 

part of the pterostigma. It’s orientation or size is diagnostic in 

some Aleiodes species. See the diagram at the end of the Glossary.

wing The following diagram shows venation patterns of the fore and 

hind wings (top and bottom, respectively), labelled with diagnostic 

features for identification of Aleiodes species. 

Wing diagram of Aleiodes wasp. Drawing provided by Paul M. Marsh.
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

109

LITERATURE

Ashmead, W.H. 1889 (1888). Descriptions of new Braconidae in the collection of the U.S. 

National Museum. Proceedings of the United States National Museum 11:611-671.

Barbosa, P., A.E. Segarra, P. Gross, A. Caldas, K. Ahlstrom, R.W. Carlson, D.C. Ferguson, E.E. 

Grissell, R.W. Hodges, P.M. Marsh, R.W. Poole, M.E. Schauff, S.R. Shaw, J.B. Whitfield, and N.E. Woodley. 2001. Differential parasitism of macrolepidopteran herbivores on two deciduous tree species. Ecology 82(3):698-704.

Bogavac, M. 1955. Rogas hyphantriae Gahan. Zastita Bilja 32:17-30.

Covell, C.V., Jr. 1984. A field guide to the moths of eastern North America. Houghton Mifflin Company, Boston. 496 pp.

Cresson, E.T. 1869. American Hymenoptera. List of the North American species of the genus Aleiodes Wesmael. Transactions of the American Entomological Society 2:377-382.

Delfín-González, H. and R.A. Wharton. 2002. Distribution of species and species-groups 

of Aleiodes (Hymenoptera: Braconidae) in Mexico. Folia Entomologia Mexicana

41(3):215-227.

Dowden, P. B. 1938. Rogas unicolor (Wesmael), a braconid parasite of the satin moth. Journal 

of Agricultural Research 56:523-535.

Dunstan, A.G. 1922. A histological account of three parasites of the fall webworm (Hyphantria cunea Drury). Proceedings of the Acadian Entomological Society 8:73-95.

Eyre, F.H. 1980. Forest cover types in the United States and Canada. Society of American Foresters, 148 pp.

Gahan, A. B. 1917. Description of some new parasitic Hymenoptera. Proceedings of the 

United States National Museum 53:195-217.

Gahan, A. B. 1922. Descriptions of miscellaneous new reared parasitic Hymenoptera. Proceedings of the United States National Museum 61:1-25.

Lentz, G.L. and L.P. Pedigo. 1974. Life history phenomena of Rogas nolophanae and Winthemia sinuata, parasites of the green cloverworm. Annals of the Entomological Society of 

America 67:678-680.

Marsh, P.M. 1979a. Family Braconidae. In: Krombein, K.V., P.D. Hurd, D.R. Smith, and B.D. 

Burkes (eds.), Catalog of Hymenoptera in America North of Mexico, Vol. 1. Smithsonian Institution Press, Washington, D.C. Pp. 144-313.
110 

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Marsh, P.M. 1979b. The braconid (Hymenoptera) parasites of the gypsy moth, Lymantria 

dispar (Lepidoptera: Lymantriidae). Annals of the Entomological Society of America

72:794-810.

Marsh, P.M. 1989. Notes on Braconidae (Hymenoptera) associated with jojoba (Simmondsia 

chinensis) and descriptions of new species. Pan-Pacific Entomologist 65:58-67.

Marsh, P.M. and S.R. Shaw. 1998. Revision of North American Aleiodes Wesmael (Part 3): the 

seriatus (Herrich-Schaeffer) species-group (Hymenoptera: Braconidae: Rogadinae). 

Proceedings of the Entomological Society of Washington 100:395-408.

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melanopterus (Erichson) species-group (Hymenoptera: Braconidae: Rogadinae). Journal of Hymenoptera Research 8:98-108.

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307.

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the compressor Herrich-Schaeffer, ufei (Walley), gressetti (Muesebeck), and procerus

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Entomological Society of Washington 105:698-707.

Mason, W.R.M. 1979. A new Rogas (Hymenoptera: Braconidae) parasite of tent caterpillars 

(Malacosoma spp. Lepidoptera: Lasiocampidae) in Canada. Canadian Entomologist

111:783-786.

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Uroceridae of North America. Transactions of the American Entomological Society 2: 

321-368.

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with description of a new species and survey of mummy types within Rogadinae s. str. 

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Rao, V.P. 1964. Survey for natural enemies of gypsy moth report for the period January to 

December, 1963. U.S. PL-480 Project. (A7FS-8) Commonwealth Institute of Biological Control, Bangalore, India. 40 pp.

Reardon, R.C. 1970. A new species of Rogas parasitic on the gypsy moth, Porthetria dispar 

(Hymenoptera: Braconidae). Proceedings of the Entomological Society of Washington

72:473-475.

Reardon, R.C., M.W. Statler, and W.H. McLane. 1973. Rearing techniques and biology of five 

gypsy moth parasites. Environmental Entomology 2:124-127.

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Shaw, M.R. 1994. Parasitoid host ranges. In: Hawkins, B.A. and W. Sheehan (eds.), Parasitoid Community Ecology. Oxford University Press, Oxford. Pp. 111-144.

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25, Orpington, U.K. 47 pp.

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Evolution, Systematics, Biodiversity and Biological Control. Agroinforum, Budapest. Pp. 321-327.

Shaw, M.R. and T. Huddleston. 1991. Classification and biology of braconid wasps (Hymenoptera: Braconidae). Handbooks for the Identification of British Insects, Vol. 7, Part 

11. Royal Entomological Society of London. Pp. 1-126.

Shaw, S.R. and P.M. Marsh. 2004. Two new eastern North American species of the Aleiodes 

coxalis (Spinola) species-Group (Hymenoptera: Braconidae, Rogadinae) reared from 

Geometridae. Zootaxa 656:1-10.

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Braconidae: Rogadinae). Journal of Hymenoptera Research 6:10-35.

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(Part 8): the coxalis Spinola species-group (Hymenoptera: Braconidae: Rogadinae). 

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Shenfelt (eds.), Hymenopterorum Catalogus (novo editio), W. Junk B.V., The Hague. 

Pp. 1115-1262,

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the Fourth International Congress of Crop Protection 1:874.

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species of ichneumon-flies. Proceedings of the U.S. National Museum 43:575-593.

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Natural History. Princton University Press, 496 pp.

Wagner, D.L., V. Giles, R.C. Reardon, and M.L. McManus. 1997. Caterpillars of eastern forests. USDA, Forest Health Technology Enterprise Team, Morgantown, West Virginia, FHTET-96-34, 113 pp.

Walley, G.S. 1941. On the genus Petalodes, with descriptions of two new North American 

species (Hymenoptera: Braconidae). Canadian Entomologist 73:213-215.

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Japan (Hymenoptera). Journal of the Agriculture Faculty at the Hokkaido Imperial 

University, Sapporo 42:1-188.
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INDEX

femoratus 25, 55, 67

flavitarsus 11, 26, 27

gastritor 1, 28, 29, 69, 89, 92

geometrae 21, 24, 30, 31, 59, 79

granulatus 17, 32

graphicus 33, 34

indiscretus 35, 36, 42, 61, 102

itamevorus 37, 49

laphygmae 29, 39, 40, 57

malacosomatos 1, 4, 29, 43, 44, 45, 92, 94

mandibularis 46, 50, 71

maritimus 38, 48

melanopodus 46, 50

melanopterus 46, 50, 109

molestus 52, 53, 66

nigristemmaticum 1, 25, 54, 55, 67

nolophanae 29, 40, 56, 109

notozophus 58, 59

pallidator 1, 36, 60, 61, 102

palmatoides 62, 64

palmatus 62, 63, 64

parasiticus 53, 65, 66

pardalotus 67

platypterygis 29, 68, 69

politiceps 34, 46, 70

praetor 4, 23, 103, 111

preclarus 72, 100, 101

pseudoterminalis 74, 96

pulchripes 59, 79, 111

quadratus 76, 77

quebecensis 4, 18, 78, 79

rileyi 80, 81

rugulosus 82, 83

sanctihyacinthi 84, 85

scrutator 86, 87

simillimus 3, 29, 88, 89, 102

smithi 19, 90, 91, 102

stigmator 4, 5, 29, 44, 92, 93, 94, 102

terminalis 75, 95, 96, 110

texanus 1, 3, 4, 23, 34, 97, 98, 103

ufei 96, 109

vierecki 99

A

abdominalis, Aleiodes 10, 27, 96

Abies 66

abietis, Neodiprion 66

aciculatus, Aleiodes 12, 13, 17, 77, 85, 102

Acronicta 4, 18, 78, 79, 80, 81, 82, 92, 93, 

94, 102

americana 92, 93, 94, 102

dactylina 92, 93, 94

distans 82

grisea 78

hasta 18, 78, 79

hastulifera 92, 94

lobeliae 18

morula 78, 79

oblinita 80, 81, 82

Agrotis 18, 19

akidnus, Aleiodes 14

albitibia, Aleiodes 5, 15, 16, 102, 111

albovenosa, Simyra 82

alder 26, 27, 69

Aleiodes

abdominalis 10, 27, 96

aciculatus 12, 13, 17, 77, 85, 102

akidnus 14

albitibia 5, 15, 16, 102, 111

angustipennis 13, 17, 32

apicalis 81, 111

atricornis 99

buoculus 89

burrus 1, 18, 19

cameronii 20, 21, 59

ceratomiae 3, 4, 22, 23, 98, 103

circumscriptus 87

compressor 62, 109

earinos 21, 24, 31, 59

ecuadoriensis 91
114

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

virginiensis 1, 100, 101

wahli 1, 73, 100, 101

Alsophila pometaria 28, 29, 92

ambigualis, Lascoria 90, 91, 102

americana, Acronicta 92, 93, 94, 102

americana, Ulmus 15

americanum, Malacosoma 43, 44

Amorpha (=Laothoe) juglandis 97

Amphion nessus floridensis 97

amyntor, Ceratomia 3

amytor, Ceratomia 22

Anagrapha falcifera 65

angustifolium, Vaccinium 38

angustipennis, Aleiodes 13, 17, 32

Anicla infecta 70

apple 85

appleworm moth mummy-wasp 56

arctiid 72, 73

Arctioidea 3

arcuata, Drepana 68, 69

armyworm 39, 40, 70

armyworm mummy-wasp 39

ash 22, 23, 85, 98

aspen 68, 86

atricornis, Aleiodes 99

augustella, Caloreas 86

Autographa 26, 27

Autoplusia egena 52

B

Balsa malana 56

banded-leg mummy-wasp 25

basiflava, Dasychira 35

beautiful mummy-wasp 72

beech 1

beet armyworm 39

bidentata, Nerice 15, 16

big-eyed mummy-wasp 22

birch 1, 31, 68, 69

black-footed mummy-wasp 50

black-winged mummy-wasp 33

blakei, Grammia 76, 77

Bomolocha palparia 56

boxelder 57

braconid 108, 109, 110, 112

Braconidae 8, 23, 99, 102, 103, 104, 108, 

109, 110, 111

broad-winged mummy-wasp 68

brunneata, Itame 37, 38

buoculus, Aleiodes 89

burrus, Aleiodes 1, 18, 19

C

Caenurgia erechtea 56

californicum, Malacosoma 43

Caloreas augustella 86

Camellia 33

Cameron’s pitted mummy-wasp 20

cameronii, Aleiodes 20, 21, 59

Catocala 20, 92, 94

cara 20, 21

catocaline 20

Ceratomia

amyntor 3, 22
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

115

sonorensis 97

undulosa 22, 23

ceratomiae, Aleiodes 3, 4, 22, 23, 98, 103

chokecherry 78

choreutid 86

cinereana, Nycteola 63, 64

circumscriptus, Aleiodes 87

clemataria, Eutrapela 20

Clostera inclusa 86

colorata, Semiothisa 88

compressor, Aleiodes 62, 109

cribrataria, Glena 88

cunea, Hyphantria 84, 85, 102, 108

cutworm 18, 40, 52, 70, 96

cyda, Semiothisa 88

cypress 1

D

dactylina, Acronicta 92, 93, 94

dagger moth 80

Darapsa myron 97

dark-marked mummy-wasp 54

Dasychira 35, 36, 61, 80, 102

basiflava 35

obliquata 60

vagans 80

decolor, Enargia 63, 64

definita, Orgyia 60, 61

devia, Eupsilia 74, 75

diprionid 66

dispar, Lymantria 35, 41, 109, 110

disstria, Malacosoma 43, 44

distans, Acronicta 82

Drepana arcuata 68, 69

drepanid 28, 69

dromedarius, Notodonta 15

ducens, Feltia 12

E

earinos, Aleiodes 21, 24, 31, 59

eastern tent caterpillar 1, 2, 4, 43

egena, Autoplusia 52

elm 23, 31, 78

elm sphinx moth 22

Enargia decolor 63, 64

Encyrtidae 44

Endopiza viteana 86

epigaea, Syngrapha 65

Erannis tillaria 88, 102

erechtea, Caenurgia 56

Euchlaena serrata 12, 102

Eupsilia devia 74, 75

Eutrapela clemataria 20

exigua, Spodoptera 39

F

falcifera, Anagrapha 65

fall armyworm 39

fall cankerworm 28

fall webworm 2, 85, 108, 111

false terminal mummy-wasp 74

Feltia ducens 12

Feltia subgothica 12, 18, 19
116

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

femoratus, Aleiodes 25, 55, 67

fervidaria athasaria, Lambidina 86

fimentaria, Narraga 88

finely-scratched mummy-wasp 12

flavitarsus, Aleiodes 11, 26, 27

forest tent caterpillar 2, 43

frugiperda, Spodoptera 39, 96

fucosa, Hypoprepia 72, 73

furcifera, Acronicta 78

G

gastritor, Aleiodes 1, 28, 29, 69, 89, 92

geographica, Oxicesta 82

geometrae, Aleiodes 21, 24, 30, 31, 59, 79

geometrid 3, 12, 20, 30, 38, 86, 89, 92

Geometridae 15, 24, 28, 29, 31, 89, 111

geometrid mummy-wasp 30

Geometroidea 3

gibbosa, Nadata 15, 16

Glena cribrataria 88

Grammia blakei 76, 77

granitata, Semiothisa 48

granular mummy-wasp 32

granulatus, Aleiodes 17, 32

grape 98

green cloverworm 56, 109

green cutworm 70

grisea, Acronicta 78

gum 1, 100

gypsy moth 2, 35, 41, 42, 61, 83, 102, 109, 

110

gypsy moth mummy-wasp 41

H

hasta, Acronicta 18, 78, 79

hastulifera, Acronicta 92, 94

henrici, Simyra 92, 93, 94

hickory 1, 25, 73, 85, 100

hog sphinx 97

honey locust 30, 31

hornworm 3, 4

Hymenoptera 6, 102, 105, 108, 109, 110, 

111, 112

Hypagyrtis unipunctata 28

Hyphantria cunea 84, 85, 102, 108

hyphantriae, Rogas 85, 108, 111

Hypoprepia fucosa 72, 73

I

Ichneumonidae 44, 104

ilicifolia, Quercus 78

includens, Pseudoplusia 52, 53

inclusa, Clostera 86

indescretus, Aleiodes 61

indiscrete mummy-wasp 35

Itame

brunneata 37, 38

pustularia 88

itamevorus, Aleiodes 37, 49

Itame mummy-wasp 37

J

jojoba 89, 109

juglandis, Amorpha (=Laothoe) 97
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

117

L

Lambidina fervidaria athasaria 86

laphygmae, Aleiodes 29, 39, 40, 57

large-jawed mummy-wasp 46

Lascoria ambigualis 90, 91, 102

latipes, Mocis 54, 55

leopard-spotted mummy-wasp 67

Leucoma salicis 60, 61

leucostigma, Orgyia 60, 61

lichen 72, 73

Lithophane 56

little-horse mummy-wasp 18

lobeliae, Acronicta 18, 78, 79

Lomographa vestaliata 88

looper 52

cabbage 52

soybean 52

lunata, Zale 20

Lymantria

dispar 35, 41, 109, 110

obfuscata 35

lymantriae, Aleiodes 41, 42

lymantriid 60, 61, 80

Lymantriidae 60, 109

M

Malacosoma 4, 43, 44, 45

americanum 43, 44

californicum 43

disstria 43, 44

malacosomatos, Aleiodes 1, 4, 29, 43, 44, 

45, 92, 94

malana, Balsa 56

mandibularis, Aleiodes 46, 50, 71

many-dotted appleworm 56

maple 1, 31, 73, 85

red 73

maritime mummy-wasp 48

maritimus, Aleiodes 38, 48

melanopodus, Aleiodes 46, 50

minians, Nephelodes 80, 96

Mocis 54

latipes 54, 55

molestus, Aleiodes 52, 53, 66

morula, Acronicta 78, 79

myron, Darapsa 97

Mythimna (=Pseudaletia) unipuncta 12, 39, 

70, 96

N

Nadata gibbosa 15, 16

Narraga fimentaria 88

narrow-winged mummy-wasp 17

Neodiprion abietis 66

Nephelodes minians 80, 96

Nerice bidentata 15, 16

nessus floridensis, Amphion 97

Nessus sphinx 97

ni, Trichoplusia 32, 52

nigra, Salix 20

nigristemmaticum, Aleiodes 1, 25, 54, 55, 67

noctuid 4, 32, 39, 44, 57, 64, 70, 74, 75, 83, 

86, 90, 94

acronictine 82, 83, 92, 94

noctuine 52

plusiine 52, 65, 66

Noctuidae 4, 11, 12, 26, 46, 50, 54, 82, 96
118

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

Noctuoidea 3

nolophanae, Aleiodes 29, 40, 56, 109

Notodonta dromedarius 15

Notodontidae 4, 15, 20

notozophus, Aleiodes 58, 59

Nycteola cinereana 63, 64

O

oak 1, 59, 67, 73, 85, 86

red 73

white 73, 86

oak gall 59

obfuscata, Lymantria 35

oblinita, Acronicta 80, 81, 82

obliquata, Dasychira 60

obumbratalis, Ostrinia 80

ocellinata, Semiothisa 30

orange-bellied mummy-wasp 10

Orgyia 61

definita 60, 61

leucostigma 60, 61

ornithogalli 42

recens 41, 42

thyellina 42

ornithogalli, Spodoptera 39, 96

Ostrinia penitalis 80

Ostrinia obumbratalis 80

Oxicesta geographica 82

P

painted lichen moth 72

Paleacrita vernata 30, 31

pallidator, Aleiodes 1, 36, 60, 61, 102

pallid mummy-wasp 60

palm-like mummy-wasp 62

palmatoides, Aleiodes 62, 64

palmatus, Aleiodes 62

palparia, Bomolocha 56

Papilionoidea 3

parasiticus, Aleiodes 53, 65, 66

parasitic mummy-wasp 65

pardalotus, Aleiodes 67

penitalis, Ostrinia 80

Peridromia saucia 52

Petalodes 64, 111

Phaeosia 15

picta, Melanchra 80

pine 48, 65, 67, 100

Pinus taeda 100

Plathypena scabra 56, 86

platypterygis, Aleiodes 29, 68, 69

politiceps, Aleiodes 34, 46, 70

pometaria, Alsophila 28, 29, 92

Populus tremuloides 63

praetor, Aleiodes 4, 23, 103, 111

preclarus, Aleiodes 72, 100, 101

prominent 45, 107

Prunus 31, 78, 81

sordinia 78

Pseudoplusia includens 52, 53

pseudoterminalis, Aleiodes 74, 96

Pteromalidae 44

pustularia, Itame 88
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

119

Q

quadrate mummy-wasp 76

quadratus, Aleiodes 76, 77

quebecensis, Aleiodes 4, 18, 78, 79

Quebec mummy-wasp 78

Quercus ilicifolia 78

R

recens, Orgyia 41, 42

Riley’s mummy-wasp 80

rileyi, Aleiodes 80, 81

Rogadini 3

Rogas

hyphantriae 85, 108, 111

unicolor 61, 108

rugulosus, Aleiodes 82, 83

S

Saint Hyacinth mummy-wasp 84

salicis, Leucoma 60, 61

Salix nigra 20

sanctihyacinthi, Aleiodes 84, 85

satin moth 60, 108

saucia, Peridromia 52

sawfly 66, 91

scabra, Plathypena 56, 86

scolopacinarius, Tornos 88

scrutator, Aleiodes 86, 87

Semiothia

granitata 48

sexmaculata 48

signaria dispuncta 48

Semiothisa

colorata 88

cyda 88

Semiothisa ocellinata 30

serrata, Euchlaena 12, 102

sexmaculata, Semiothisa 48

signaria dispuncta, Semiothisa 48

similar mummy-wasp 88

simillimus, Aleiodes 3, 29, 88, 89, 102

Simyra

albovenosa 82

henrici 92, 93, 94

small-eyed mummy-wasp 14

smartweed caterpillar 80

smeared dagger moth 80

Smith’s banded mummy-wasp 90

smithi, Aleiodes 19, 90, 91, 102

smithii, Xestia 96

smooth-headed mummy-wasp 70

sonorensis, Ceratomia 97

sordinia, Prunus 78

south-western mummy-wasp 58

sphingid 34, 98, 103

Sphingidae 3, 4, 7, 23, 33, 97, 103

Sphingoidea 3

Spilosoma virginica 76

Spodoptera

exigua 39

frugiperda 39, 96

ornithogalli 39, 96

spring cankerworm 30, 31

spring mummy-wasp 24

stigmata mummy-wasp 92, 94
120

ALEIODES WASPS OF EASTERN FORESTS ____________________________________________________________

stigmator, Aleiodes 4, 5, 29, 44, 92, 93, 94, 

102

subgothica, Feltia 12, 18, 19

sweet gum 100

Syngrapha epigaea 65

T

taeda, Pinus 100

tent caterpillar mummy-wasp 43

terminalis, Aleiodes 75, 95, 96, 110

terminal mummy-wasp 95

Terminator 95

texanus, Aleiodes 1, 3, 4, 23, 34, 97, 98, 103

Texas mummy-wasp 97

Thunberg’s mummy-wasp 28

thyellina, Orgyia 42

tillaria, Erannis 88, 102

Tornos scolopacinarius 88

tortricid 86

tremuloides, Populus 63, 64

Trichoplusia ni 32, 52

troublesome mummy-wasp 52

tulip tree 100

tussock moth 60, 61

U

ufei, Aleiodes 96, 109

ugly nest caterpillar 45

Ulmus americana 15

undulosa, Ceratomia 22, 23

unicolor, Rogas 61, 108

unipuncta, Mythimna (=Pseudaletia) 12, 39, 

70, 96

unipunctata, Hypagyrtis 28

V

Vaccinium 1, 38

angustifolium 38

vagans, Dasychira 80

variegated cutworm 52

vernata, Paleacrita 30, 31

vestaliata, Lomographa 88

Viereck’s mummy-wasp 99

vierecki, Aleiodes 99

Virginia mummy-wasp 100

virginica, Spilosoma 76

virginiensis, Aleiodes 1, 100, 101

viteana, Endopiza 86

W

Wahl’s mummy-wasp 101

wahli, Aleiodes 1, 73, 100, 101

Walley’s palmate mummy-wasp 63

walnut 85, 97, 98

walnut sphinx 97

waved sphinx moth 22

white-legged mummy-wasp 15

white-marked tussock moth 60, 61

willow 4, 20, 21, 81, 86

wrinkled mummy-wasp 82

X

Xestia smithii 96
____________________________________________________________ ALEIODES WASPS OF EASTERN FORESTS

121

Y

yellow-footed mummy-wasp 26

yellow-marked mummy-wasp 86

yellow-striped armyworm 39

Z

Zale lunata 20