Social Wasps of Florida

Social Wasps of Florida

Henry R. Hermann, Ph.D.*

Department of Mathematics and Science

Edison State College
LaBelle Center

LaBelle, Florida

Email: hrhermann@edison.edu

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I am currently working on the morphology of the venom apparatus of Polistes major (Hymenoptera: Vespidae: Polistinae) and Pseudomyrmex gracilis (Hymenoptera: Formicndae: Pseudomyrmecinae) in the Ft. Myers, Florida, area. This may lead to a study of defensive behavior in Polistes major if I can find enough nests. 
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Students of this group will find additional information and references for earlier papers at: www.henryhermannpublications.blogspot.com.
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What Will You Find At This Blogsite?

This blogsite is currently being established and maintained as a means of identifying and learning about social wasps found in the state of Florida. It is periodically updated but currently does not have a complete list of individuals working on this group. 
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All social wasp species in Florida belong to the family Vespidae. Members of this family are commonly referred to by wasp specialists simply as vespids. They are one group of eusocial (truly social) insects. Such wasps live complex social lives and have a defensive nature. Further breakdown of taxonomic categories within this family will be presented below.
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Colonies of vespid wasps, like those of other eusocial insects, are female-dominated, i.e., the queen is an egg-layer, and workers (all females) represent the workforce. The sole function of males (which are produced only at certain times during the colony cycle) is to produce spermatozoa for succeeding generations of females. They do not function in defending the nest or doing work.
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Dichotomous keys and important biological information will be offered, along with pictures of each species. Because of certain anatomical and behavioral features of members of this group, keys are often not needed for their identification. However, they may be used as an additional tool to determine which species one is looking at.
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What Are Social Wasps?

Social wasps are insects that live in colonies and demonstrate defensive behaviors that sometimes are harmful to humans. Members of a colony cooperate with carrying out the various chores during the colony cycle. Yet, a certain amount of antagonistic behavior is noticed between colony members, often in association with dominance establishment.
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Such colonies may be initiated by a single queen (i.e., the species is haplometrotic) or by a number of fertile females (i.e., the species is pleometrotic or polygynic). While these definitions are easy to understand, the actual designation of a species as haplometrotic, pleometrotic, or polygynic is occasionally confusing because significant variation exists in the way some species (e.g., Polistes annularis) found their colonies.
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In general, founding (fertile) females are poorly defensive when compared to the defensiveness of workers (infertile females). The life cycle is usually divided into: 1) a founding stage when fertile females are the only adults on the nest; and 2) an ergonomic stage when workers take charge of colony functions (other than egg-laying). The nest founding period is generally much shorter than the ergonomic period.
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Throughout the ergonomic period, adult workers emerge from cells that they occupied as an egg, larva, and pupa (they demonstrate complete metamorphosis). By the end of a season, there are usually numerous workers, unless the nest is disturbed in some way (e.g., when they are attacked by predators or parasites).
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Near the end of the ergonomic period in temperate zones, females that are potential queens and males (both of which are generally referred to as reproductives) emerge on the nest. Mating between these reproductives generally occurs in the fall of the year, and newly fertilized females do not construct a new nest at that time. They generally linger on or around the nest until winter approaches.
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As ectotherms, they enter hibernation when the weather becomes cold, and they remain in hibernation until the weather becomes warmer. Hibernation sites may be under bark, under roof tiles, in other secluded places, and sometimes in the nest itself (as with certain yellowjackets). The queens of that year die by the end of the season (in temperate zones), along with the males. During the first warm spells of the year, the hibernating wasps may leave their hibernaculum and be seen warming in the sun's rays, only to return to their hibernaculum when the weather turns cold again.
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Old nests are not used in consecutive years in temperate zones, and there are new queens each and every year. Nest founding in tropical and semitropical areas (such as in Mexico, Central America, South America, and South Florida) are more difficult to understand because queens may not hibernate in the fall of the year, and they remain as a queen over a period that may last into another year.
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Founding Females are Generally Related
In most cases, fertile pleometrotic or polygynic females that initiate a nest together are related, generally all originating from the same nest during the previous season (all are siblings and dauaghters of a single queen). Whether haplometrotic or pleometrotic, spring-time activities in temperate areas often show a considerable amount of nest shifting by fertile females who are competing for nest site selection and dominance status. Even in a species such as Polistes exclamans which is typically haplometrotic, there is a considerable amount of nest shifting by fertile females (which is a result of dominance interactions on each nest) (Willer, 1988; Willer and Hermann, 1989).
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Most members of the family Vespidae share similar anatomy, although size, the shapes of certain structures, and degree of robustness may vary. One outstanding characteristic of all members of the group that is easy to recognize is seen in the presence of longitudinally-folded wings. They remain folded when adults are not in flight and during certain defensive behaviors, but they unfold during flight and other wing movements (such as while fanning the nest).
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There are a total of four wings. The two forewings extend from the second (mesothoracic) thoracic segment, and two hindwings extend from the third (metathoracic) thoracic segment. The prothorax, as in other types of winged insects, does not have wings. The hindwings are hooked to the forewings by tiny hooks called hamuli. Anatomical features that vary most among the species are the degree of robustness, coloration, and pattern. The following picture shows their general anatomy with the wings unfolded.

The drawing above shows the generalized anatomy of a social wasp (family Vespidae). The wings are unfolded. What is not evident in this drawing is the way the abdomen is attached to the thorax. In all members of the Hymenoptera, the first abdominal segment (referred to as the propodeum) is completely fused with the last thoracic segment and thus it appears as the posterior part of the thorax. The next segment or two may be expressed in aculeate (stinging) species as a more slender petiole, giving the insect the appearance of a "waist." The remainder of the abdomen, the posterior-most, separate part, is generally called the gaster.

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Paper-Makers

Social wasps are also referred to as paper wasps since they construct their nests of masticated wood pulp which they collect from trees and buildings that have exposed wood. They are world-wide in distribution.
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The coarseness of the paper usually depends on the size of the wasp making it. At one extreme among indigenous species are nests made by the Bald-faced Hornet (Dolichovespula maculata) (the most coarse) and at the other extreme the nests of Mischocyttarus species (the least coarse). Paper made by the largest vespid in North America, the exotic European Hornet (Vespa crabro), is extremely course in texture.
To collect wood, adult wasps position themselves with their head down (body oriented vertically to the ground) on a source of wood. They use their mandibles to scrape the wood. As it is scraped, saliva is added, and eventually the collected material takes the form of a round, moist ball of wood pulp which the wasp takes back to its nest. It applies this material gradually to form the walls of paper cells, using their antennae as guides in placing the pulp. The first cell constructed is round, but as cells are added around it, the initial cell and subsequent ones take on a hexagonal shape. The outer cells may have rounded outer edges.
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Paper nests may be bag-like (calyptodomous) and contain a number of tiers of cells (as found in hornet and yellow jacket colonies) or open (gymnodomous), with a single tier (as found in polistine species).

The photo above shows a paper nest of Dolichovespula maculata, the bald-faced hornet. The bag-like nest (referred to as calyptodomous) generally has several tiers of cells inside, one positioned on top of the other. This type of nest is also found in yellow jacket species. The bald-faced hornet generally builds a suspended nest in trees or on buildings, while yellow jackets most often build their nest in the ground. Yellow jackets in Florida, as in other areas, sometimes build their nest above ground, and one species in Florida has been found to construct nests that are quite large.
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The picture above shows a paper nest of Polistes exclamans with several adults. This type of nest is attached by one or more pedicels (the number of pedicels often depending on the species) to a substrate (i.e., it is stellocyttarous). It is generally referred to as an open nest since there is a single tier of cells (this condition being referred to as gymnodomous), and the cells are exposed. The nest is protected to a degree from foraging insects by the addition of repellent chemicals applied by adult wasps to the pedicel (the process being referred to as pedicellar smearing). Being volatile, these chemicals (which are produced by abdominal glands on the ventral side of the abdomen) are periodically smeared on the nest pedicel. Since the nest cells are exposed, nest attack by birds and other flying organisms generally comes about from the direction of the exposed cells, and a vast array of pre-stinging defensive behaviors are demonstrated toward an intruder.
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How Many Different Types of Vespids Are There?
According to Akre and Davis (1978), "there are about 15,000 species of aculeate (stinging) wasps in the world. However, 95% of these are solitary and nonaggressive species that use their stings primarily for subduing prey." Thus they do not defend their nest, but they may use their sting in defense during a personal attack.
Most wasp stings experienced by humans are inflicted by social members of the family Vespidae. This family includes the subfamilies Stenogastrinae, Polistinae, and Vespinae, with about 800 species. Of these subfamilies, only the Polistinae (commonly referred to as polistine wasps) and Vespinae (commonly referred to as vespine wasps) are found in Florida.
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Vespines
Three species of indigenous vespine wasps occur in Florida: Dolichovespula maculata (the bald-faced hornet), Vespula squamosa (the southern yellow jacket), and Vespula maculifrons (the eastern yellow jacket). These three species typically construct a bag-like (calyptodomous) nest. They may be distinguished from one another by their size, coloration, and patterns.
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Polistines
The subfamily Polistinae is a diverse group of eusocial wasps, being represented in the New World by 21 genera (Carpenter 2004). Most species are tropical in distribution. Only four genera occur in North America north of Mexico. Species of wasps in the genus Polistes are the most common types of paper wasps, having over 300 recognized species.
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Polistes
According to Carpenter (1996), the cosmopolitan genus Polistes (Polistinae) comprises most of the species diversity within the family Vespidae, with 28 species occurring in the United States alone. Ten species occur in Florida. Within Florida, as in the rest of the United States, they construct petiolated (stellocyttarous), single-combed, open (gymnodomous) nests in which the cells are clearly visible.
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Nesting Period and Size
Most species of social vespid wasps in North America, including the north half of the Florida peninsula, form small, seasonal colonies (Pickett et. al., 2001). Temperate species (those in the northern half of the state) found their nests during the spring period, go through their ergonomic period of nest enlargement and production of adults throughout the warmer months, and they produce potential queens and males near the end of the season. Mating occurs during the fall and females that have mated (fertile females) enter hibernation through the winter months. They exit their hibernacula in the spring and return to the vacinity of the parental nest, but they construct a new nest for the new season. As pointed out above, seasonality is not as evident in the southern parts of Florida, and social wasps there sometimes function more like tropical species, remaining on the nest even when temperatures drop close to the freezing point. 
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Pickett et. al. point out that colonies of most polistine wasps rarely produce more than 100 contemporaneous adults, although nests of Polistes annularis colonies have been known to achieve more than 1,400 cells. The nests of vespines (hornet and yellow jackets) are characteristically more productive than those of polistines in the United States, and certain vespines are known to build collosal nests.
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Those authors report on an extremely large nest of Vespula squamosa in Southern Florida that had 476,982 cells and 38 putative egg-laying queens. It was claimed to be the largest nest made by vespid wasps found within the United States. These researchers pointed out that large polygyne (many-queened) colonies of Vespula species are not rare, attaining their large size because they persist for many years in warm environments, often with queens overwintering in the nest itself. Vespula squamosa, like other vespine species, is generally a haplometrotic species. Based on the large colony they dissected in Lee County, Florida, it appeared the proximate mechanism permitting multiple queens to inhabit such large colonies was the manner in which brood regions were separated from each other by regions of dead comb.
While the finding of large nests with multiple queens indicates that they may sometimes be polygynic or pleometrotic, if areas of separation exist in a single large colony, the nest may actually be made up of numerous separate colonies within the nest's framework, and thus each colony exists as a haplometrotic group. Thus it may be best to restrict the term polygyny to a colony in which there are multiple fertile females that all function together as queens and a pleometrotic colony as one in which there are multiple fertile females but only a single egg layer.
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Taxonomy of Paper Wasps
The taxonomy of eusocial wasps, like the taxonomy of many groups of animal species, has gone through an evolution of sorts over the years. The following classification hierarchy may help in an understanding of how social wasps are currently categorized:
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Kingdom - Animalia
Phylum - Arthropoda (arthropods have exoskeletons and jointed appendages)Class - Insecta (insects have a head, thorax and abdomen and six legs)Order - Hymenoptera (this order includes all ants, bees, and wasps)
Family - Vespidae (this family includes all of the social wasps in Florida).
Subfamily Vespinae (generally referred to as vespines) - In Florida, this subfamily includes the bald-faced hornet (genus Dolichovespula) and two species of yellow jackets (genus Vespula).

Subfamily Polistinae (generally referred to as polistines) - In Florida, this subfamily includes members of the genera Polistes and Mischocyttarus.


In summary, the genera of social wasps represented in Florida are: 1) Vespula, 2) Dolichovespula, 3) Polistes, and 4) Mischocyttarus. There are 14 indigenous species within the family Vespidae that are found in the state of Florida. These are:
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Subfamily Vespinae

Dolichovespula maculata - Bald-faced Hornet

The above photo is of a bald-faced hornet, Dolichovespula maculata. Notice the robust appearance of its body and black and white coloration which is different from all other vespids in the state of Florida. This is the largest of the indigenous social wasps in Florida. It, like other vespines, is typically a haplometrotic nest founder.

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Vespula squamosa - Southern Yellow Jacket

The above photo of a Vespula squamosa adult female shows the distribution of yellow and black coloration. It is this species that constructs huge nests in southwest Florida. 
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Vespula maculifrons - Eastern Yellow Jacket

The above drawings compare the facial regions of Vespula squamosa (left) with Vespula maculifrons, pointing out the distribution of yellow and black.
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The above photo of a Vespula maculifrons worker shows the distribution of yellow and black on the body.
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Subfamily Polistinae

Polistes annularis (Linnaeus)

The above photo is of a Polistes annularis queen on a new nest. Notice the dark coloration on the abdomen and the absence of a number of light bands that are often found in other species of Polistes. This queen has founded her nest alone, but other colonies of this same species may found their nest with multiple fertile females. When multiple fertile females of this species found a nest together, they generally undergo a week or two of dominance establishment. Following dominance establishment, the alpha (most dominant) female undergoes ovarian development and generally becomes the sole egg layer. The ovaries of more subordinate females undergo atrophy. However, they remain fertile with the spermatozoa they received during their mating swarm, and another female can become queen if the alpha female dies.
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Polistes apachus de Saussure

The above picture shows a nest of Polistes apachus on an ergonomic nest. This wasp species shows a considerable amount of yellow with a medium brown (as compared to Polistes exclamans) in the body coloration but not as much as Polistes major major.
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Polistes bahamensis Bequaert and Salt

While this picture of Polistes fuscatus shows clearly marked lighter rings on the abdomen and lighter markings on the thorax, this species is quite variable, and such markings are not always evident. This is a good shot to show the folded wings. They become unfolded when the wasp flexes its muscles during wing extension.

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Polistes bellicosus Cresson

This picture of Polistes bellicosus also clearly shows the longitudinally-folded wings characteristic of all vespid wasps. When folded, it is difficult to tell that there are two wings on each side of the meso- and meta-thorax. The rear (meta-thoracic) wings, like those of other polistines, are hooked to the meso-thoracic wings by small hooks called hamulae.
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Polistes carolina (Linnaeus)

It is clear to see that this species of Polistes lacks the banding and other patterns found in other species. Its overall light coloration is a good feature to use for its identification.
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Polistes dorsalis dorsalis (Fabricius)

The banding and alternating colors of Polistes dorsalis dorsalis make it a beautifully-patterned species. It is also a little smaller than many other polistines.
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Polistes exclamans Vierak

Polistes exclamans is probably the most common polistine species around homes in Florida. Its smaller size, characteristic banding, and readiness to nest on many types of plants help to distinguish it from other species. It appears somewhat like Polistes apachus but the yellow bands on its abdomen are not as wide. It is well known as a nest defender and may represent the greatest threat to intruders. You will notice that some of the cells in this nest are capped (closed). The larvae spin silk over the opening of their cell prior to entering the pupal stage. When first spun, the caps are an unblemished white color, but they become discolored over time when adult workers add water or head secretions to the caps.
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Polistes fuscatus (Fabricius)

This Polistes fuscatus shows well defined rings of light yellow around the abdomen and lines demarcating certain thoracic sclerites, but considerable variation occurs in this species, sometimes resulting in confusion of this species with other forms of Polistes. When confusion exists as to their identify, you may have to resort to using the dichotomous key.
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Polistes major major Palisot de Beauvois

The large size of Polistes major major adults and the extensive yellow coloration on their body helps to distinguish this species from other polistines. However, the degree of yellow coloration varies, even among the members of a single nest. Very little has been published on this species.
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Polistes metricus Say

 

 Polistes metricus lacks the extensive yellow banding on the abdomen and marks on the thorax that are demonstrated by other polistine species. Its body also doesn't appear as slim as most other species. *
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Mischocyttarus mexicana cubicola

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Paper Wasps Are Eusocial

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Like all eusocial insects, social wasps exhibit certain characteristics that define a eusocial state: 1) the young (larvae) are cared for by adult wasps; 2) there is some form of reproductive division of labor, i.e., there is usually one or more fertile females on the nest, one or more of which may deposit eggs, and a number of infertile females that are generally referred to as workers; 3) there is an overlap of generations (the queen lives long enough to coexist with her offspring).
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From Mating to Egg-Laying
Females that mate store spermatozoa they received from males in the mating in a chamber called a spermatheca. When they begin to nest, an egg passes out of the ovary of the egg-laying female and passes down her fallopian tube to the uterus. Spermatozoa are released from the spermatheca and pass down a spermathecal tube to the uterus where the egg is fertilized. This event of fertilization is carried out throughout the nest founding period and most of the ergonomic period. In late summer, eggs that pass down the fallopian tube may be fertilized (resulting in females) or spermatozoa may not be released, and the egg will develop into a male (non-fertilized eggs become males). The process of females arising from diploid eggs and males arising from haploid eggs is generally referred to as haplodiploidy.
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How Are Social Wasps Raised?

Eggs are deposited in paper cells by a single queen in haplometrotic colonies and sometimes by multiple fertile females in pleometrotic colonies. They are adhered to the bottom or lower sections of the sides of the cells. Larvae that hatch from these eggs are vertically positioned in the cells with their head down. They are raised within that paper cell and fed the flesh of some type of insect. Most members of the social wasps (those members of the subfamilies Polistinae and Polybiinae) feed on caterpillars, but a few are known to feed on other insects.

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Once larvae have fed to completion and are ready for pupation, they spin a silken cap over the opening of the cell. At this point, the silken cap is flat. The larvae then turn around and spin silk in the bottom of the cell. In the process, their caudal end pushes against the silken cap, making it balloon out from the nest. Emergence from the cell by a new adult occurs when the adult cuts an opening through the silken cap with its mandibles and escapes. Following emergence, the remainder of the silk cap is removed by adults, and cells may be used again once or twice to produce other individuals.

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Are They Beneficial or Pestiferous?

In this light, social wasps are sometimes said to be beneficial insects, ridding gardens of insect pests. A considerable amount of research has been carried out to determine if polistine species can be used in field settings to control pestiferous caterpillars. People that have established a butterfly garden may think of them as pests because wasps looking for insect prey do not distinguish between caterpillars that are desirable or undesirable in the garden.
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Life Cycle, Metamorphic Stages, and Developmental Times

In the biology of the wasp Polistes (Epicnemius) cinerascens Saussure, E. Giannotti reported on the life stages. Larvae go through stages of feeding. At the end of a particular stage, larvae molt their exoskeletons, grow larger in size, and a new exoskeleton is deposited over the body surface. Each feeding stage is referred to as an instar.
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The number of instars in this species is 5.
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The duration of the immature stages is as follows: eggs = 13.0 days, larvae = 23.7, pupae = 22.2 and total duration = 58.6 days.
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Longevity of adult wasps = 38.3 days. This is a neotropical species. The life cycle of more temperate species may vary, depending on the temperature present during their developmental period. More research is needed to determine information of this type for other polistine species.

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Details of Nest Founding and Maintenance?
Most polistine species within the United States occur in a temperate climate. They react to cold weather by entering hibernation. Late in the summer period, potential queens and males develop on the nest, and mating most commonly occurs in the fall.
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Most wasps that hibernate are fertile females that have mated during the fall mating period. When they exit from their hibernacula in the spring, they generally go back to the vicinity of their parental nest and enter some form of behavior referred to as dominance establishment.
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Species in Florida, especially in the southern parts, may remain on the nest throughout the winter period. and new nests are established as peripheral (satalite) colonies of the parental nest. This type behavior is similar to what would be found in more tropical areas. With additional information on species in southern Florida, we may determine that group founding is also present in that area.
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During the early nest-founding stage, these females may fight with one another and chase one another from the nest. Some females that lose battles on one nest may fly to another nest to enter a dominance interaction with other females. Eventually, a dominance hierarchy (pecking order) is established, and it is the dominant females that become the egg layers.
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With haplometrotic species, i.e., they found and maintain their nests with a single queen. In such cases, the queen is intolerant of other egg-laying females. With pleometrotic species, i.e., the nests are occupied by a number of fertile females, only one is generally the egg-layer. The establishment of a dominant queen (alpha female) on a nest before and during the nest founding stage generally results in the enlargement of the ovaries in the dominant queen and an atrophy of the ovaries in more subordinate fertile females, in spite of the presence of spermatozoa in their spermathecae.
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The term polygyny is often reserved for a species that typically has a number of queens (all of which are egg layers) in a single colony. The difference between pleometrotic nest founding and polygyny becomes confusing when the species may demonstrate variation in the degree of dominance establishment. For instance, Polistes annularis may demonstrate haplometrotic nest founding or pleometrotic nest founding. At times on a pleometrotic nest, the degree of dominance determines if one or more fertile females are depositing eggs.
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If a female is extremely dominant (i.e., she is despotic), she may chase all other females from the nest and thus she must function alone until workers emerge. Most researchers report that such lone queens out-produce queens on a pleometrotic nest. However, when considering the productivity of the nest as a whole, the nests of pleometrotic colonies are generally larger than those of haplometrotic colonies and better at survival. If a queen is dominant but tolerant of other females, she may become the principle or sole egg layer. In such cases, a linear dominance hierarchy (pecking order) is established, and if the queen dies, the beta female redevelops her ovaries and becomes the new queen.
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In yet other cases, there may be a poorly defined dominance hierarchy established in which more than one female may deposit eggs. In such cases, there may be a degree of oophagy (egg eating) in which females feed on each other's eggs, subsequently depositing their own eggs in a cell. Colonies that develop a poorly defined dominance hierarchy usually suffer from a lack of colony cooperation and thus the productivity is lower than it would be if cooperation ensued.
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While group founding is more characteristic of certain tropical wasp species, it is poorly understood in social wasps within the United States. As pointed out above, Polistes annularis often commences nest building with a number of fertile females (thus they may fit the description of group-founding wasps), and they maintain their nest with multiple females (i.e., they are pleometrotic). Most social wasps in Florida are solitary nest founders and thus maintain a haplometrotic status.
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In addition, wasps in South Florida usually stay on the nest during the winter and thus nest founding does not follow the typical hibernation and founding of a new nest each spring. More work on species in South Florida should prove to be interesting in this respect.

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Identifying Social Wasps

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Social wasps represent only a small number of the order Hymenoptera (ants, bees, and wasps). Of all wasps, social species represent the minority. Thus, it is important to first determine if an insect is a member of the Hymenoptera, if it is a wasp, and finally if it is a social species. The following material will help in this determination.
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General Description Of Some Families Within The Order Hymenoptera

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Wings among the various species of the order Hymenoptera, when present, number four. There are some wingless females within the order. The males of such species possess wings. Wings are transparent and membranous. The forewings are larger than the hindwings, the two pairs being hooked together by tiny hooks known as hamuli that extend from the anterior margin of the hindwings. Wings are folded longitudinally.

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Antennae are filamentous, with an elongate basal scape and a segmented distal region which is bent (geniculate). The antennae are longer than the head. The mandibles (the largest pair of mouthparts) are well developed and move transversely to enable biting. It is the mandibles that are used to collect wood to build their nest in paper wasps.

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Parasitoid wasps possess an ovipositor, through which eggs pass from the oviduct. Female aculeate (stinging) wasps possess a venom apparatus with a well-defined sting at the posterior end of the body. The sting has been derived from sclerites that are homologous to those of the ovipositor of earlier forms. Thus it is only females that have a sting. Males, of course, do not have one since males have never possessed an ovipositor. 

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Most aculeate species do not represent a threat to humans because they are poorly defensive. However, social species characteristically defend their nest and can sting in groups. Most vespid wasps are capable of stinging repeatedly because the sting lancets possess poorly-defined barbs at their distal tips. Some yellow jackets occasionally have been known to lose their sting in the skin of their victims because they possess larger lancet barbs than most stinging insects. Loss of the sting at the sting locus, as is commonly found in honey bees, is referred to as sting autotomy.
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Key To Some Families Of Insects In The Order Hymenoptera

1a. The base of the abdomen is broadly joined to the thorax - These insects belong to the suborder Symphyta which includes the sawflies and horntails.

The photo above shows a lateral view of a sawfly, a member of the order Hymenoptera that has its abdomen broadly joined to its thorax. Female sawflies and horntails also have an ovipositor through which eggs pass. The sting of aculeates, which is derived from the ovipositor of its predecessors, does not use its sting in depositing eggs.
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1b. The base of the abdomen is constricted into a slender "waistline" referred to as a petiole. The petiole is attached to the propodeum (first abdominal segment which is fused to and often indistinguishable from the last thoracic segment). These are in the suborder Apocrita and known as aculeates - Go to # 2

The photo above shows an adult Polistes which has the base of the abdomen joined to the thorax by a thin "waist." Members of the suborder Apocrita are known as aculeates, possessing a sting (which is a modified ovipositor). Eggs do not pass through the sting but exit from the female reproductive system immediately in front of the sting.
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2a. Adult females are wingless. This group includes wasps in the families Tiphiidae, Mutillidae, Bethylidae, Formicidae, and a few others. It does not include the social wasps. This seems a little confusing when considering the ants. While ant workers are wingless (apterous) females, the reproductive females in the colony are generally winged (alates). They become dealates after a mating swarm when they remove their wings before initiating a new colony.

The photo above is of a dorsal view of a female mutillid wasp. While she is wingless, males are winged. Female members of the Mutillidae generally have an extremely long and coiled sting with which they can deliver a painful sting. Most of them are also aposematically colored. However, they are not social insects.
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2b. Two pairs of wings are present - Go to #3. Be sure to look closely at this feature. Since tiny hooks (hamuli) extend from the anterior edge of the hindwings and hook on the posterior edge of the forewings, the pair on each side may appear to be a single wing.
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3a. Ovipositor or sting is present. These are female aculeates. - Go to #4

The photo above shows a lateral view of a vespid wasp (family Vespidae). While the sting is evident in this picture, it is often retracted into the abdomen and may not be visible without probing.
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3a. Ovipositor or sting is not present. These are males. - Go to #5
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4b. An ovipositor extends from the ventral part of the abdomen just anterior to the last abdominal segment. The last abdominal sternite (ventral plate) is split ventrally around the ovipositor. Ovipositor cannot be completely withdrawn into the abdomen. These characteristics belong to females of the parasitic Hymenoptera, including the Ichneumonoidea, Chalcidoidea, Evanoidea, and Cynipoidea.

The photo above shows an ichneumon wasp with an extremely long ovipositor (egg-laying tube). Eggs pass from the internal reproductive system and move down the ovipositor to be deposited in the substrate of its hosts. While it is said that ichneumons do not possess a sting, some species have shorter ovipositors and can produce pain when used on humans.
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4b. Sting may protrude from the end of the abdomen, but it is retractable. These characteristics belong to the females of aculeate (stinging) Hymenoptera. - Go to #6

The photo above shows the distal tip of a paper wasp abdomen with the sting extruded. A drop of venom has formed at the tip of the sting. When examining such female wasps, the sting is sometimes not seen because it is within the sting sheath and thus out of view.
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5a. Trochanters (first leg segment beyond the coxa) have two segments. This is characteristic of the males of parasitic Hymenoptera, including the Ichneumonoidea, Chalcidoidea, Evanoidea, and Cynipoidea.

The picture above shows the leg segments. Notice the trochanter between coxa and femur.
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5b. Trochangers have a single segment. This is characteristic of males of aculeate (stinging) Hymenoptera - Go to #6
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6a. Petiole in the "waist bears one or two dorsal nodes that are distinctly separate from the propodeum (first abdominal segment that is joined to the thorax) and gaster (the enlarged part of the abdomen that follows the petiole). These are the ants, family Formicidae.
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6b. Petiole is without any dorsal node - Go to #7
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7a. Hairs on the thorax are branched or feather-like (plumose). The basitarsus of the hind leg is often broad and flattened. Body has a bulbous appearance. These are the bees, superfamily Apoidea.
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This honey bee shows abundant hairs on the thorax, but they are found on other parts of the body as well. When examined under a dissecting microscope, the hairs are found to be branched. If wasps have hairs on their body, they are not branched and generally not as extensive as those in bees.
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7b. Hairs on the thorax are unbranched, and the body is rather slender - Go to #8
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8a. The first discoidal cell of the forewing is shorter than the submedian cell. The wings are not folded when at rest. - Go to #9 *

The discoidal cell in the wings above are #?. The submedian cell is #?.
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8b. The first discoidal cell of the forewing is much longer than the submedian cell (see drawings above). Forewings are folded (diplopterous) when they are at rest - Go to #13
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9a. The mesopleuron has one transverse suture and legs are very long. These are solitary wasps in the family Pompilidae.
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9b. The mesopleuron does not have a transverse suture, and the hind femora do not reach back to a position near the caudal tip of the abdomen - Go to #10
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10a. The lateral lobe of the pronotum (dorsal plate on the first thoracic segment) does not extend back as far as the tegula. These are solitary wasps in the family Sphecidae.
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10b. The pronotum extends back to the tegula or close to it - Go to #11
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11a. The distal margin of the wings are wrinkled to appear as very fine pleats. These are solitary wasps in the family Scoliidae.
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11b. The distal margin of the wings is not wrinkled - Go to #12
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12a. The compound eye is oval in shape, with no notch (ocular sinus) on it inner margin. These are solitary wasps in the family Tiphiidae.
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12b. The compound eye is kidney-shaped (reniform) - Go to #13
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13a. The middle tibia has one apical spur. The claws are forked (bifid). The mandibles are elongate and crossing. These are solitary wasps in the subfamily Eumeninae.
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13b. The middle tibia have two apical spurs. The claws are tapered. The mandibles are broad and not crossed. These wasps are members of the social family Vespidae.
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Key To The Subfamilies and Genera Of Social Vespidae in Florida

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1a. The clypeus is broadly truncate and usually notched at its distal tip. The hindwings lack a jugal lobe. The first gastral segment is broadly truncate. These wasps belong to the Subfamily Vespinae - Go to #2
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1b. The clypeus is rounded, bluntly or pointed at its tip. The hindwings have a jugal lobe. The wasps have a rather slender build. The first gastral segment is elongate or tapering basally. These wasps belong to the Subfamily Polistinae - Go to #4
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2a. The vertex of the genae is enlarged above and behind the compound eyes. The postocellar area of the vertex is longer than the ocellar triangle when seen in dorsal view. These are very large wasps with mahogany brown and yellow markings on the body. Vespa crabro germana Christ. If this species is in the state, it is an exotic in the northern portions. It is larger than any indigenous vespine species.
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2b. The vertex of the genae are not conspicuously enlarged behind the compound eyes. The postocellar area of the vertex is shorter than the ocellar triangle. This type of wasp has bands of black and yellow or black and white - Go to #3
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3a. The oculomalar space is at most half of the length of the penultimate antennal segment. These wasps belong to the genus Vespula.
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3b. The oculomalar space is about as long as or longer than the penultimate antennal segment. These wasps belong to the genus Dolichovespula Rower.
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4a. The first gastral segment is elongate, stalk-like, very much longer than it is broad, and only slightly widened distally. At its caudal margin, it is much narrower than the second gastral segment. These are small wasps, Mischocyttarus, in the subfamily Polybiinae. There is only one subspecies in this subfamily in Florida - Mischocyttarus mexicana cubicola.

4b. The first gastral segment is compact, cone-shaped, and not stalk-like. It gradually widens distally until it is almost as wide as the second gastral segment. These wasps are in the genus Polistes, subfamily Polistinae.
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Key to the Two Genera of Polistine Wasps in Florida
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*1a. The first gastral segment is very much longer than it is broad, only slightly widened distally and much narrower than the second gastral segment. These are relatively small wasps, that are sometimes recognized by taxonomists in their own subfamily Polybiinae. Mischocyttarus mexicana cubicola.
1b. The first gastral segment is compact, gradually widening distally until it is almost as wide as the second gastral segment. These are species of the genus Polistes.
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Key to the Species of Polistes in Florida
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Florida Species of Dolichovespula (Linnaeus) Rohwer
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Dolichovespula maculata (Linnaeus) - Bald-faced Hornet
Description - Dolichovespula maculata, the bald-faced hornet, is the only indigenous species of the hornet group within the state of Florida. Its large size and pattern of black and creamish coloration helps us to easily distinguish this species from other social wasps in Florida. It is the largest indigenous species of vespid wasp in North America and is commonly found from coastal lowlands to mountainous areas. Although it most typically constructs its calyptodomous nest on tree branches, it sometimes adheres its paper to the sides of houses, especially under the eaves.
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Biology - The nest is generally very large (40 cm diameter) and constructed of approximately five tiers of cells within a multilayered outer envelope. The nest generally has a single opening in the bottom, but weathered nests may have other openings on occasion.
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House flies are the preferred prey of this species of wasp. While it is generally not considered a scavenger, they are often attracted to refuse where they can become abundant.
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Florida Species Of Vespula - Yellow -
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Only two species of the subgenus Vespula occur in Florida: Vespula squamosa (Drury) and Vespula maculifrons (Buysson). They can be identified without the use of a dichotomous key. 
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Vespula squamosa (Drury)
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Description - The scutum has two broad, longitudinal pale stripes. Males and females have an abdomen banded with black and yellow. The queen has extensive deep orange markings.
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Biology - This species exhibits marked polymorphism in which the queen has a deep orange coloration, unlike any other species of the genus Vespula. Workers and males are banded with black and yellow. This species is considered a facultative social parasite in which queens habitually usurp newly initiated nests of Vespula maculifrons. Unlike other social parasites, Vespula squamosa subsequently completes a normal annual cycle, especially in the northern part of the state. As with other vespid species in temperate areas, the major part of the growing season is spent producing brood that develop into workers. Immatures that will develop into potential queens and males are produced in the latter part of the season.
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A typical nest, especially in the northern parts of Florida, is a rather large bag-like structure with a fairly durable grayish envelope. At maturity, approximately eight tiers of cells are constructed within the envelope.
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Colonies continue into the early winter in northern parts of Florida, followed by mating and subsequent hibernation of females destined to be queens of the following year. Gigantic multiseasonal colonies have been reported from the more southern areas of Florida where the cycle is continued throughout the winter months and a second generation of queens apparently usurp their mother or develop a separate colony somewhere in the large nest. Such nests initially appear to be polygynic (having multiple queens). However, separations between queens and their colonies by "dead" sections of comb apparently keep the various queens isolated from one another.
In many areas, these wasps are considered to be a nuisance because of several reasons: they are attracted to soft drink machines and refuse where sugar-rich liquids and other edible materials accumulate; nest sites may be a hazard for maintenance crews and mowers.
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Vespula maculifrons (Buysson)

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Description - The scutum is black, without light-colored stripes (or at most with two short, pale marks near the scutellum). The occipital carina on the back of the head extends all the way to the base of the mandible. The dorsal surface of the first gastral tergite has a sagittate black marking.
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Biology - Nests of this species are rather large, often consisting of six to eight tiers of cells within a creamish beige or tan-orange fragile outer envelope. The envelope consists of overlapping layers of paper.
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Identification of Species Of Polistes Latreille and Mischocyttarus Saussure In Florida

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General Discription - These are referred to as petiolate, aculeate, diplopterous wasps. The first discoidal cell of the forewing is longer than the submedian cell. They have two apical spurs on their middle tibia, and the pronotum extends back to the tegula. Their compound eyes are reniform, and their antennal segments number 12 in females and 13 in males. There are six visible gastral segments in females and seven in males. They are paper wasps with a slender build when compared to vespine species. A jugal lobe is present on the hind wing. The clypeal apex is narrow and pointed or rounded when examined from a frontal view.
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Mischocyttarus (Monocyttarus) mexicanus cubicola
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Description - Mischocyttarus (Monocyttarus) mexicanus (de Saussure) is one of two species in this genus in the United States (Krombein et al., 1979). It is found in Georgia, Florida, South Carolina, Cuba, and the Bahamas. The genus is primily neotropical, being treated taxonomically by Bequaert (1933), Richards (1945, 1978) and Zik (1935, 1949).
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It is smaller than any species within the genus Polistes (body length 6-9 mm). The stellocytarous nest is constructed with small cells (approximately 3 mm in diameter).
Biology - The genus Mischocyttarus is the only member of its own tribe, the Mischocyttarini. They build a relatively simple, stellocyttarous and gymnodomous (single-comb) nest. Their biology is similar to that of species in the genus Polistes, although they seem to be less defensive than most species of Polistes.
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The distribution of M. m. cubicola in the United States was reported by Hermann, Gonzalez, and Hermann (1985). Those authors visited every county in Florida to determine its presence.
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Litte (1977, 1979) initially described the biology of M. mexicanus in Florida and M. flavitarsis (de Saussure) in Arizona. The subspecies Mischocyttarus mexicanus mexicanus occurs in Texas, Mexico, and Central America. Mischocyttarus mexicanus cubicola is reported from Florida, Georgia, Alabama, South Carolina, Cuba, and the Bahamas (Hermann and Chao, 1984).
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In a study of the nesting biology and defensive behavior of this subspecies on Sapelo Island, Georgia, Hermann and Chao (1984) found nests on buildings, other man-made structures and on vegetation. Nest on man-made structures were most often wooden frames of windows, wooden support beams of metal-roofed out-buildings, and the underside of concrete arbors. In Polk County, they were found in metal and ceramic bell-like wind chimes. Nests on vegetation on Sapelo Island were found only on the underside of the leaves of the cabbage palm, Sabal palmetto (Walt).
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Based on information from 10 nests, a single egg layer was on each nest during June. Determination of who was an egg layer was based on ovary size. The pleometrotic status of this subspecies was reported by Litte (1977).
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Many shredded nests were found, evidently due to predation. Birds and ants are the most important predators of social wasps (Jeanne, 1975), but Jeanne (1970) also reported predation by bats. In Florida, Litte (1977) listed blue jays, scrub jays, carolina wren, common yellowthroat, cardinal, thrashers, mockingbirds, and robins as potential vertebrate predators. Invertebrate predators of wasp immatures were found to be the following ant species: Dorymyrmex flavopectus, Crematogaster ashmeadi, Pheidola floridana, and Camponotus floridanus.
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Defensive behavior was poorly expressed during the nest-founding stage, as is the case with most polistine species, although a form of behavior called pseudoattack (poorly-defined attack without stinging) was elicited from wasps upon strong provocation. Both pseudoattack and attack behavior was demonstrated by this species in more mature colonies, but mature colonies were relatively docile when compared to the colonies of polistine species. When provoked, other forms of defense involved the type of nest architecture, erratic flight, general nest excitability, defensive posture, wing raising, wing buzzing, wing fluttering, forward jerking, mandibular pecking, backward jerking, abdominal pumping, abdominal twisting, retreat, and escape.
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According to R. Clouse ( ), the mortality rate of new nests is astonishingly high (80% fail within 20 days of nest initiation). Surviving haplometrotic females have much higher daily rates of production than pleometrotic females. The daily per capita rate of cell addition declines as group size decreases in pleometrotic nests. Lone female production was much larger than group production in his study.
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Polistes annularis (Linnaeus)

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Description - Polistes annularis is one of the largest polistine species in the state of Florida, with a body length of 18-25 mm. Polistes major may be as large or larger than this species. Nests are also large, sometimes reaching a diameter of 25 mm or more. Sexual dimorphism and coloration among colony inhabitants vary only slightly. The clypeus of males is flatter than it is on females and more closely contiguous to the eye margin.
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Biology - Hermann and Dirks (1971) reported on their nesting behavior and dominance interactions in Georgia. There, as in the northern parts of Florida, a definite seasonality is found in the annual cycle. In those temperate locations, a number of fertile females may found a nest together during the early spring (April). In the southern part of the state, colonies probably continue through winter months, and new nests result from reproductive females initiating satallite nests from their initial colony.
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There is usually some indication of a linear dominance hierarchy amongst founding females, resulting in an alpha female becoming queen and the more subordinate females functioning as workers. If the queen is lost, the next most dominant female generally takes her place as the queen. During the maintenance of this hierarchy, the ovaries of the alpha female become very large, whereas the ovaries of subordinate females atrophy. During queen loss, the ovaries of the beta female redevelop so that she becomes the egg layer.
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Comments about variation in the establishment and maintenance of a dominance hierarchy are found in the introductory comments on polistine behavior in this blogsite. Dominance ranges from absolute despotism to poorly defined.
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While mating occurs primarily during the fall, males of this species have been found hibernating along with females in Georgia. Since they may temporarily emerge from hibernation during warm winter days in that area, it may be possible for them to mate during these brief periods of activity. Very little is known about the winter nesting and mating behavior in the southern part of the state where the winter is not as severe.
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Polistes apachus de Saussure
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Polistes bahamensis Bequaert and Salt
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Polistes bellicosus Cresson
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Polistes carolina (Linnaeus)
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Polistes dorsalis dorsalis (Fabricius)
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Polistes exclamans (Vierck)
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Polistes exclamans (Viereck) (Hymenoptera: Vespidae: Polistinae), ranges in the U. S. from New Jersey to Florida and west to Nebraska, Colorado and Texas (Cannamela, 1993; Krombein et al., 1979). It is common throughout the southeastern U. S.
Known as the guinea wasp, this species can be distinguished from most other polistines in Florida by the abundance of yellow on the legs and abdomen. The only species that has more yellow is Polistes major (see its description). Its nests are found on buildings and plants.
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Polistes exclamans is similar in it's biology and life history to other temperate zone paper wasps (Hermann et al., 1975; Strassmann, 1985a, 1985b; Strassmann et al., 1983). For an excellent in-depth look at spring behavior in temperate areas, the reader is directed to a study of Polistes exclamans by Donna E. Willer on Sapelo Island, Georgia (Willer, 1988).
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Overwintering queens initiate their nest in the spring (nest founding period), and the first workers appear several weeks later in early summer. This species is generally thought of as a haplometrotic species.
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Willer (1988) studied multiple foundress associations and nest switching in a localized subpopulation of P. exclamans on Sapelo Island, Georgia. In 1985, she found that 31 of 178 marked females switched to nests other than those on which they had been captured initially. Fifteen of 18 nests were involved in switching by these females. In 1986, she found that 97 of 239 females switched nests, involving 17 of the original 19 nests. 
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During nest initiation in both 1986 and 1987, over 50% of all nests in this subpopulation were initiated by multiple foundresses. Only two nests were founded by females bearing the same color coded marks in each year.
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The number of females returning to the exact site of their natal nest during nest initiation was negligable in both years. Viscosity in this subpopulation was high: no marked females from this location were found nesting in other locations on Sapelo Island. Multiple foundress associations and nest switching were assessed in terms of their evolutionary implications, especially in relation to knowledge of nestmate recognition among Polistes females.
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In a separate study, Willer compared thirteen colonies nesting on saw palmettos and 49 colonies nesting on a wooden outbuilding. Nest initiation on saw palmettos involved only solitary foundresses, while 30 (61.2%) of the nests on the outbuilding were initiated by multiple foundresses. Nests on saw palmettos averaged 37 cells, and those on the outbuildings averaged 85 cells at the peak of the season.
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Parasitism of wasp brood by a pyralid moth, Chalcoela pegasalis, and a eulophid wasp, Elasmus polistis, led to early decline of colonies on saw palmettos. By July of that year, only 3 were functioning; at the end of the season, those three were defunct.
Nineteen colonies on the outbuilding failed during nest initiation or early in colony formation. Birds attacked brood in 12 other colonies in this subpopulation. However, 18 of the original colonies on the outbuilding survived to the end of the season, producing large numbers of reproductives.
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One of the only studies on defensive flights in polistine wasps was carried out on this species by Barden Canamella and at the University of Georgia in 1993 (thesis), but the information was not published. It involved the use of video cameras designed to view wasps as they left the nest and stung a victim.
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Polistes fuscatus (Fabricius)
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Polistes major major Palisot and Beauvois
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Polistes metricus Say
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Glossary of Terms

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Arthropoda - A phylum of animals that have an exoskeleton and jointed appendages. The phylum includes such arthropods as crustaceans, ticks, mites, spiders, phalangids, whip scorpions, scorpions, centipedes, millipedes, and insects.
Clypeus - Best defined as the upper lip in a facial view of an insect.
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Despotic Dominance - A form of dominance in which the queen is so dominant that she chases everyone off the nest. She is intolerant of other females.
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Family - The family is a taxonomic category that ends in idae. It is position in the classification hierarchy beneath Order and above Subfamily and Genus. All paper wasps belong to the family Vespidae. Within this family are the following subfamilies: Vespinae, Polistinae, and Polybiinae.
*Gaster - The gaster is the part of the abdomen without the first segment (the propodeum), as well as the petiole and post-petiole (as in ants). See propodeum below).
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Group Founding - Group founding is a condition in which a group of fertile females leave their parental nest and establish a new nest together.
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Haplometrosis - Haplometrosis is a condition in which a fertile female establishes her nest alone, without the help of other females.
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Insects - Arthropods with three body regions (head, thorax, and abdomen) and six legs in the adult stage. The class with insects is called Insecta.
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Linear Dominance Hierarchy - A linear dominance hierarchy is a behavioral hierarchy in which the individuals can be placed in a particular order from most dominant to most subordinate. This is in contrast to a despotic condition in which the queen is so dominant that she chases all other females off the nest. In a linear hierarchy, the fertile female that is dominant to all wasps except the alpha female will become queen of the nest if the alpha is lost.
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Nest Founding - Nest founding refers to the establishment of a new nest by onr or more queens. Vespines (hornets and yellow jackets) are typically haplometrotic, establishing their nest with a single queen. Some polistine species are pleometrotic, establishing their nest with two or more fertile females, while others are haplometrotic. Tropical species of wasps often establish their nest as group founders, i.e., a number of queens fly together to a new location and establish a new nest together.
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Petiole - This is a structure found best developed in ants. It is the second swollen abdominal segment of a hymenopteran with a waist. Some ants have a petiole while others have a petiole and post-petiole. These characteristics can be used to determine what subfamily ants belong to.
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Pleometrosis - This is a term used in speaking about colonies of social wasps that have multiple fertile females, but they not represent multiple queens. Often, such a species establishes a linear dominance hierarchy in which their is a dominant queen and subordinate fertile females.
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Polygyny - This is a term that is used in speaking about colonies of social wasps that have multiple queens, all functioning on the nest, including egg laying. A polygynic species generally has a number of new queens leaving their parental nest and establishing a new nest together (a behavior referred to as group founding).
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Propodeum - The propodeum is the first abdominal segment that is firmly joined to the thorax. In paper wasps and some other members of the Order Hymenoptera, the propodeum actually looks like the posterior part of the thorax, and the rest of the abdomen, which is posterior to the thin waist, is properly called the gaster.
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Scientific Names - Scientific names are generally binomens (i.e., composed of two names), e.g., Polistes annularis. In some cases, the scientific name may be a trinomen (i.e., composed of three names) (e.g., Polistes major major and Mischocyttarous mexicanus cubicola). The first name in the scientific name is the genus, the second name is the species, and the third name is the subspecies. Scientific names often have the names of the original describer of the name following it, e.g., Polistes apachus de Saussure. If the describer's name is in parantheses, it indicates that the original name has been changed for some taxonomic reason from the original name, e.g., Polistes annularis (Linnaeus). The scientific name, whether a binomen or trinomen, is set in italics to stand out from the other type, while all other terms (e.g., family and subfamily names, as well as the describer's names) are not.
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Subfamily - Subfamily is a taxonomic category that is a subdivision of the family taxon. For instance, the family Vespidae contains three subfamilies, Vespinae, Polistinae, and Polybiinae.
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Tiers - Entire layers of cells within the nests of paper wasps. Gymnodomous nests have a single tier while calyptodomous nests have multiple tiers. Thus species that build calyptodomous nests produce nests that have more workers.
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Types of Nests - The single tier of a stelocyttarous nest is attached to the substrate by one or more pedicels. The nest is gymnodomous, i.e., its single tier is visible because it is not covered by an outer bag-like covering. Calyptodomous nests are bag-like and consist of several innter tieirs.
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Waistline - This term refers to the constriction between the apparent posterior region of the thorax and the gaster of the wasp. In more primitive wasps, there is no waist. The abdomen is broadly attached to the thorax.
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References
Akre, R. D., and H. G. Davis. 1978. Biology and pest status of venomous Wasps. Ann. Rev. Entomol., 23:215-38.
Cannamela, B. R. 1993. Defensive Behavior of Polistes exclamans (Viereck). Masters Thesis, Univ. of Georgia, Athens, GA.
Hermann, H. R., and J. T. Chao. 1984. Nesting biology defensive behavior of Mischocyttarus mexicanus cubicola (Vespidae: Polistinae). Psyche, 91:51-65.
Hermann, H. R., R. Barron, and L. Dalton. 1975. Spring behavior of Polistes exclamsns (Hymenoptera: Vespidae: Polistinae). Entomol. News 86: 173-78.
Hermann, H. R., and T. F. Dirks. 1971.
Hermann, H. R., J. M. Gonzalas, and B. S. Hermann. 1985. Mischocyttarus mexicanus cubicola (Hymenoptera), distribution and nesting plants. Florida Entomol., 68:609-14.
Krombein, K. V., P. D. Hurd, and D. R. Smith. 1979. Catalog of Hymenopptera in America North of Mexico. Vol. 3. Smithsonian Institution Press: Washington, D. C.
Litte, M. 1977. Behavioral ecology of the social wasp Mischocyttarus mexicanus. Behav. Ecol. Sociobiol., 2:229-46.
Strassmann, J. E. 1985a. Relatedness of workers to brood in the social wasp, Polistes exclamans (Hymenoptera; Vespidae). Z. Tierpsychol. 69, 141-48.
Strassmann, J. E. 1985b. Worker mortality and the evolution of castes in the social wasp, Polistes exclamans. Insectes Sociaux 32, 275-85.
*Willer, D. E. 1988. Behavioral Ecology and Population Biology of Polistes exclamans on Sapelo Island, Georgia. Doctoral Thesis, Univ. of Georgia, Athens, GA.
Willer, D. E., and H. R. Hermann. 1989. Multiple foundress associations and nest switching among females of Polistes exclamans (Hymenoptera: Vespidae). Sociobiology. 16: 197-216.
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About the Organizer of this Blogsite:
Henry Hermann received his doctorate degree at Louisiana State University. He subsequently held a research and teaching position in the Division of Biological Sciences at the University of Georgia for 30 years. While there, he was active in workshops available to pest control operators that involved the identification of pestiferous insects. 
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His main interests in research were the morphology and behavior of social wasps and ants. Most of his work on social wasps was on species within the subfamily Polistinae. Through the years, he published numerous papers on the morphology of the venom apparatus and behavior of defensive behavior. His list of publications may be accessed through the following blog site: http://henryhermannpublications.blogspot.com/. 
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Information on other blog sites, directing the reader to the most recent fiction and non-fiction titles, are given there.
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Dr. Hermann is currently teaching biology in the Department of Mathematics and Science at Edison State College. He has been a Regional Leader (SW Florida) for the Florida Writers Association and published author of a number of books in the categories of fiction and non-fiction. The following blogsites highlight his recent books:
Non-Fiction: http://henryhermannnonfiction.blogspot.com/
Fiction: http://henryhermannnovelist.blogspot.com/
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