Cockroaches are among the oldest and most primitive of insects. They evolved about 350 million years ago during the Silurian Period, diverging together with the manrids from an ancestral stock that also gave rise to termites (Boudreaux 1979). Cockroaches are recognized as the order Blattaria. Although the majority of species are feral and not directly associated with people, a few species have evolved in proximity to human habitations, where they have adapted to indoor environments. Their omnivorous feeding behavior, facilitated by their unspecialized chewing mouthparts, has contributed to a close physical relationship between cockroach populations and humans, with resultant chronic exposure of humans to these pests. The presence of some species in the home (e.g., German and brownbanded cockroaches) often is an indicator of poor sanitation or substandard housekeeping.
Although they are primarily nuisance pests, their presence can have important health implications. Cockroaches are generalists that feed on virtually any organic substance grown, manufactured, stored, excreted, or discarded by humans. Consequently, food supplies are at risk of contamination by pathogens associated with cockroaches. Because species that infest structures typically have high reproductive rates, humans commonly are exposed to high levels of potentially allergenic proteins associated with cockroaches, which can lead to significant respiratory ailments. Cockroaches also can serve as intermediate hosts of parasites that debilitate domestic animals.
TAXONOMY
There are about 4000 species of cockroaches worldwide. About 70 species occur in the United States, 24 of which have been introduced from other parts of the world. According to Atldnson et al. (1991), 17 of these species are pests of varying degrees. There are five cockroach families, three of which include most of the pest species: Blattidae, Blattellidae, and Blaberidae. Species in the Cryptocercidae are unusual in that they have gut symbionts similar to those found in termites, and they live in family groups in decaying logs. Members of the Polyphagidae include those dwelling in arid regions, where they are capable of moving rapidly through sand. Species in these two families are rarely pests. The family Blatfidae includes relatively large cockroaches that are the most common peridomestic pests throughout much of the world. Blattellid cockroaches range in length from less than 25 mm (e.g., Supella and Blattella) to 35-40 mm (e.g., Periplaneta and Parcoblatta spp.). Parcoblatta species are feral, occasionally invading homes but seldom reproducing indoors. Blaberid cockroaches range greatly in size and include some of the more unusual species, such as the Cuban cockroach, which is green as an adult, and the Surinam cockroach, which is parthenogenetic in North America. Nearly all of the blaberids that occur in the United States are restricted to subtropical regions and have minor medical or veterinary significance. Taxonomic keys for adults are provided by McKittrick (1964), Cornwell (1968), Roth (1985), and Heifer (1987). A pictorial key for identifying the egg cases of common cockroaches is provided by Scott and Borom (1964).
MORPHOLOGY
Cockroaches have retained their basic ancestral form. The Blattaria are distinguished from other insect orders by morphological characters associated with wing size and venation, biting/chewing mouthparts, and prominent cerci. They differ from other orthopteroid insects by having hind femora which are not enlarged, cerci typically with eight or more segments, a body that is dorsoventrally flattened and generally ovoid, and a head that is largely concealed from above by a relatively large pronotum.
A common indicator of cockroach infestations is their egg cases, or oothecae (singular ootheca), purse-shaped capsules that typically contain 5-40 embryos (Fig. 3.1). Coloration ranges from light brown to chestnut brown, depending on the degree of sclerotization. A keel that runs the anterior length of the ootheca permits transport of water and air to the developing embryos. Each embryo is contained in a separate compartment that may or may not be obvious externally. In some species (e.g., German and brownbanded cockroaches) lateral, anterior-to-posterior indentations denote the individual developing embryos. Others have only weak lateral indentations (e.g., brown and smokybrown cockroaches), and still others have no lateral indentations but differ in their symmetry (e.g., Oriental, American, and Australian cockroaches).
The mouthparts of cockroach nymphs and adults are characterized by strongly toothed mandibles for biting and chewing. Maxillary and labial palps are well developed, with five and three segments, respectively. Antennae are long and whiplike, originate directly below the middle of the compound eyes, and consist of numerous small segments. The arrangement of three ocelli near the
antennal sockets is variable: they are well developed in winged species (macropterous) but rudimentary or lacking in species with reduced wings (brachypter0us) or those lacking wings altogether (apterous).
Adults generally have two pairs of wings that are folded fanwise at rest. The front wings, called tegmina (singular tegmen), are typically hardened and translucent, with well-defined veins. The hind wings are membranous and larger. In some species, such as the wood cockroaches (e.g., Parcoblatta species), females are brachypterous and incapable of flight, whereas males are macropterous. Other species, such as the Florida woods cockroach (Eurycot# floridana), have only vestigial wing buds as adult males and females. In cockroaches, all three pairs of legs are well developed, with large coxae and slender, long segments that aid in the rapid running that is characteristic of these insects. Each femur has two longitudinal keels that typically are armed with spines. The tibiae are often heavily spined and are used for defense against predators. Each tarsus consists of five segments with a pair of claws and may bear a padlike arolium that aids in walking on smooth surfaces. Ventral pads, or pulvilli, are present on tarsomeres 1-4. A pair of caudal cerci have small ventral hairs that are sensitive to vibrations caused by lowfrequency sound and air movement; their stimulation initiates an escape response.
The posterior end of the abdomen of some nymphs and all males bears a pair of styli (singular stylus) between the cerci, arising from the sternum of the ninth abdominal segment. In winged species, the styli may be apparent only when viewed ventrally. The structure of the styli serves to distinguish males from females. Generally the males also can be recognized by their more slender bodies, with laterally tapered and dorsally flattened external genitalia (terminalia). The terminalia of the more robust females are notably broader than in males and bear a conspicuous subgenital plate that is rounded or keel-like when viewed ventrally. Associated with this plate is a relatively large genital chamber (genital pouch) in which the ootheca develops. For a more detailed description of cockroach genitalia, see McI~ttrick (1964) or Cornwell (1968). Nymphal stages are similar in appearance to adults, but they lack wings, have incompletely developed genitalia, and may vary markedly in color from the adult.
LIFE HISTORY
Cockroaches are paurometabolous insects. The immature cockroaches generally are similar in appearance to the adults except for their undeveloped sexual organs and lack of fully developed wings (Fig. 3.2). Reproduction in cockroaches is typically sexual, although parthenogenesis is reported in a few species. Comparative life history data for some of the more common cockroach pests are provided in Table I.
In cockroaches, embryogenesis and oviposition occur in one of three ways. Most species are oviparous, including
all Periplaneta species and the Oriental and brownbanded cockroaches. Eggs of oviparous species are protected inside a thick-walled, impermeable ootheca which is deposited soon after it is formed. Embryonic development occurs external to the female. The German cockroach is oviparous, but the female carries the ootheca protruding from the genital chamber until just hours before hatching occurs. The ootheca is softer than in Periplaneta species, allowing uptake of water and nutrients from the genital pouch. A few cockroaches, such as Blaberus species and the Surinam cockroach, are pseudo-ovoviviparous, in that females produce an ootheca which is extruded, rotated, and then retracted into the genital pouch. The eggs are incubated internally until hatching. The only known pseudo-viviparous species is Diploptera punctata, a pest species in Hawaii; the embryos hatch while still in the genital pouch. Embryogenesis takes 1-8 weeks, depending on the species.
The number of nymphal instars varies from 5 to 13, depending on the species, nutritional sources, and microclimate. Development of pestiferous species through the nymphal stadia requires from 6-7 weeks for German cockroaches to well over a year for Periplaneta species and other larger cockroaches. Typically, the nymphs exhibit strong aggregation tendencies, governed largely by aggregation pheromones. These pheromones act as locomotory inhibitors; when cockroaches perceive the pheromone they become relatively stationary. Studies of various species have shown that development to the adult stage is quicker when nymphs are reared in groups rather
than in isolation. However, aggregation does have a biological cost; those reared in groups typically are smaller in size, and cannibalism may occur. Longevity of cockroaches varies from several weeks to over a year.
BEHAVIOR AND ECOLOGY
Mating in cockroaches generally is preceded by courtship behavior facilitated by sex pheromones. In some species a blend of volatile compounds is produced by virgin females to attract and orient males (e.g., Periplaneta species and the brownbanded cockroach). In the German cockroach, the sex pheromone is a blend of nonvolatile and volatile cuticular components that elicits courtship by males following palpation of the female’s integument by the male’s antennae. Once courtship is initiated in the male, he turns away from the female and raises his wings to expose dorsal tergalglands; the female feeds on pheromones from these glands as the male grasps her genitalia with his pair of caudal claspers. Most species copulate in an end-to-end position. During the hour or so that follows, a spermatophore is formed and passed from the male into the genital chamber of the female. Only about 20% of females mate again after the first gonotrophic cycle.
Cockroaches can be categorized ecologically as domestic, peridomestic, or feral. Domestic species live almost exclusively indoors and are largely dependent on humans for resources (food, water, and harborage) for survival. They rarely are able to maintain themselves outdoors. Although this group contains the smallest number of species, it presents the greatest concern to human health. Domestic species include the German and brownbanded cockroaches. Peridomestic species are those which survive in or around human habitation. Although they do not require humans for their survival, they are adept at exploiting the amenities of civilization. This group is represented by American, Australian, brown, and smokybrown cockroaches (all Periplaneta species), the oriental cockroach, and the Florida woods cockroach. Feral species are those in which survival is independent of humans. This group includes more than 95% of all species in the world. Only a few occur indoors as occasional and inadvertent invaders that typically do not survive in a domestic environment. They are of little or no medical importance.
Cockroach behavior and survival are strongly influenced by their need for food, water, and safe harborage from potential predators and detrimental microclimates. They are omnivorous and will consume virtually any organic matter, including fresh and processed foods, stored products, and even book bindings and pastes on stamps and wallpaper when more typical foodstuffs are not available. Cockroaches have the same general problems with water balance as do other terrestrial arthropods. Their relatively small size results in a high surface area to volume ratio and a high risk of losing water through respiration, oral and anal routes, or the cuticle. Temperature, air flow, relative humidity, and availability of liquid water greatly affect water regulation.
As a result of these physiological considerations, physical constraints of the environment usually determine habitat preferences of cockroaches in and around structures. Oriental and American cockroaches, for example, require high moisture and occur in damp terrestrial environments such as septic tanks and municipal sewer systems. Brown, smokybrown, and Florida woods cockroaches occur in a wider range of habitats associated with trees, wood and leaf piles, wall voids, and foundation blocks of buildings. Brownbanded cockroaches are more tolerant of drier conditions and commonly occur in kitchens, pantries, and bedrooms. German cockroaches occupy harborages near food and water. Consequently, they are found primarily in kitchens and pantries, and secondarily in bathrooms, when their populations are high. In mixed populations of German and brownbanded cockroaches, the German cockroach tends to outcompete the brownbanded cockroach within 9 months.
Cockroaches are adept crawlers and are capable of rapid movement even across windows and ceilings. Flight ability varies with species. Some are incapable of flight except for crude, downward gliding used as an escape behavior. Others are weak fliers, occasionally seen flying indoors when disturbed. Still others are relatively strong fliers that are particularly active at sunset, when they may be attracted indoors by lights and brightly lit surfaces. Attraction to light is especially common in the Asian, Surinam, and Cuban cockroaches and in many of the wood cockroaches (Parc0blatta species).
Pestiferous cockroaches that occur indoors are typically nocturnal and tend to avoid lighted areas. This enables them to increase their numbers and become established in structures before human occupants even become aware of their presence.
COMMON COCKROACH SPECIES
The following 11 species of cockroaches are commonly encountered by homeowners in the United States and are the ones most frequently brought to the attention of medical entomologists.
Oriental cockroach (Blatta orientalis)
This peridomestic cockroach (Fig. 3.3) is believed to have originated in northern Africa and from there spread to Europe and western Asia, South America, and North America. It is a relatively lethargic species that prefers cooler temperatures than does the German cockroach and is primarily a concern in temperate regions of the world. Adults are black and 25-33 mm long. Males are winged but do not fly, and females are
brachypterous. Their tarsi lack aroliar pads, precluding this cockroach from climbing on smooth vertical surfaces. Oothecae are 8-10mm long, each typically containing 16 eggs. Also commonly lmown as waterbug, this species is usually associated with damp or wet conditions, such as those found in decaying wood, heavy ground cover (e.g., ivy) , water meter boxes, and the lower levels of structures. It infests garbage chutes of apartment complexes, sometimes reaching upper floors. Development is slow compared to that of most other species, requiring about a year depending on temperature conditions. Adults may live for many months. Mobility is fairly restricted, malting control easier than for most other species. This species is rarely seen during the daytime.
American cockroach (Periplaneta americana)
The American cockroach (Fig. 3.4) is a large species with adults 34-53 mm in length. It is reddish brown, with substantial variation in light and dark patterns on the pronotum. Adults are winged and capable of flight. Nymphs typically complete development in 13- 14months while undergoing 13molts. Adults live an average of 15 months, but longevity may exceed 2 years. Females drop or glue their oothecae (8 mm long) to substrates within a few hours or days of formation. Each ootheca has 12-16 embryos. A female generally produces 6-14 egg cases during her life (mean of 9).
The American cockroach is perhaps the most cosmopolitan peridomestic pest species. Together with other closely related Periplaneta species, P. americana is believed to have spread from tropical Africa to North America and the Caribbean on ships engaged in slave trading. Today this species infests most of the lower latitudes of both hemispheres and extends significantly into the more temperate regions of the world.
The habitats of this species are quite variable. American cockroaches infest landfills, municipal sewage systems, storm drainage systems, septic tanks, crawl spaces beneath buildings, attics, tree holes, canopies of palm trees, voids in walls, ships, electronic equipment, caves, and mines. Studies conducted in Arizona indicated movement by a number of individuals several hundred meters through sewer systems and into neighboring homes. This species often can be seen at night on roofs and in air stacks or vents of sewage systems, through which they enter homes and commercial buildings. Entrance also is gained to homes through laundry vent pipes and unscreened or unfiltered attic ventilation systems. This cockroach is known to move from crawl spaces of hospitals via pipe chases into operating theaters, patients’ rooms, storage facilities, and food preparation areas. Consequently, the potential of this cockroach for disseminating pathogenic microorganisms can be a significant concern for health care personnel.
Australian cockroach (Periplaneta australasiae)
Adult body coloration is similar to that of the American cockroach, but with paler lateral markings on the upper edges of the tegmina (Fig. 3.5). The pronotum is tinged with similar coloration. Adults are slightly smaller than American cockroaches, measuring 32- 35 mm in length. Females mature in about 1 year and typically live for another 4-6 months. A female can produce 20-30 oothecae during her lifetime; the ootheca is about 11 mm long and contains about 24embryos. Embryonic development requires about 40 days. Nymphs are strikingly mottled, distinguishing them from nymphs of other Periplaneta species.
This peridomestic species requires somewhat warmer temperatures than the American cockroach and does not occur in temperate areas other than in greenhouses and
other pseudotropical environs. In the United States, outdoor populations are well established in Florida and along the coastal areas of Louisiana, Mississippi, Alabama, and Georgia. It commonly is found in environments similar to those inhabited by the smokybrown cockroach. In situations where both species occur (e.g., tree holes, attics), the Australian cockroach tends to displace the smokybrown. It can be a serious pest in greenhouses and other tropical environments in more temperate latitudes, where it can cause feeding damage to plants, notably seedlings.
Brown cockroach (Periplaneta brunnea)
The brown cockroach (Fig. 3.6) is smaller than the American cockroach (33-38 mm), and its pronotal markings are more muted. The most apparent diagnostic characteristic for separating these two species is the shape of the last segment of the cercus; in the brown cockroach, the length is about equal to the width, whereas in the American cockroach the length is about 3 times the width. The ootheca of the brown cockroach usually is larger (7-13 vs 8 ram) and contains more embryos (24 vs 16). The brown cockroach affixes its oothecae to substrates using salivary secretions. They give the ootheca a grayish hue not typical of other Periplaneta species that attach their oothecae with salivary secretions. This species is more subtropical than the American cockroach, occurring throughout the southeastern United States, where it infests homes and outbuildings. It is less frequently associated with sewage than is the American cockroach. Because of its similar appearance to the American cockroach, it is often misidentified and may be more widely distributed than is commonly recognized. In Florida, P. brunnea is commonly found in canopies of palm trees and attics. It also readily infests various natural cavities and those in human-associated structures. The oothecae can be useful in differentiating species infesting buildings. Most
cockroach oothecae persist in the environment after the nymphs have emerged and provide a history of infestation.
Smokybrown cockroach (Periplanetafuliginosa)
The smokybrown cockroach (Fig. 3.7) has become a major peridomestic pest throughout the southern United States, including southern California, and extends as far north as the Midwestern states. It can be differentiated from the American cockroach by its slightly smaller size (25-33 ram) and uniform dark coloration. Mthough developmental times are quite variable, individuals mature in about 10 months. Adults may live for more than a year. Females produce several oothecae, which are 10-11 mm in length with 20 embryos, at 11-day intervals.
Primary loci for this peridomestic species in the southeastern United States are tree holes, canopies of palm trees, loose mulches such as pine straw or pine bark, and firewood piles. Within structures, P. futiginosa seeks the ecological equivalent of tree holes–areas characterized as dark, warm, protective, and moist, with little air flow and near food resources. These include the soffits (eves) of underventilated attics, behind wall panels, the interstices of block walls, false ceilings, pantries, and storage areas. From these harborages, individuals forage for food and water, generally returning to the same refugia. Mark-release-recapture studies using baited live traps have shown that the median distance traveled between successive recaptures is less than I m but that some adults may forage at distances of more than 30 m.
Florida woods cockroach (Eurycotisfloridana)
This cockroach is restricted to a relatively small area of the United States along the Gulf of Mexico from
eastern Louisiana to southeastern Georgia. It is mentioned here only because of its defensive capabilities. It is a large, dark-reddish brown to black cockroach (Fig. 3.8), 30-40 mm long. Although small wing pads are evident, adults are apterous and are relatively slow moving. Oothecae are 13-16 mm long and contain about 22 embryos. E. floridana occurs in firewood piles, mulches, tree holes, attics, wall voids, and outbuildings. Last-instar nymphs and adults, if alarmed, can spray a noxious mix of aliphatic compounds that are both odoriferous and caustic. If this is sprayed into the eyes or onto soft tissues, a temporary burning sensation is experienced. Domestic dogs and cats quicldy learn to avoid this species. Among its common names are the Florida cockroach, the Florida woods roach, the Florida stinkroach, and palmettobug. The last term also is commonly used for other Pcriplaneta species.
Brownbanded cockroach (Supella longipalpa)
Like the German cockroach, this domestic species probably originated in tropical Africa, where it occurs both indoors and outdoors. In North America and Europe it is confined almost exclusively to indoor environments of heated structures. In warm climates, infestations occur particularly in apartments without air conditioning and in business establishments with relatively high ambient temperatures, such as pet stores and animal-care facilities. Adults are similar in size to the those of the German cockroach (13-14.5 mm long) but lack pronotal stripes. Adults have two dark bands of horizontal stripes on the wings (Fig. 3.9), whereas nymphs have two prominent bands running across the mesonotum and first abdominal segment. The brownbanded cockroach derives its name from these bands.
Populations tend to occur in the nonfood areas of homes, such as bedrooms, living rooms, and closets. Male brownbanded cockroaches occasionally fly and are attracted to lights. Members of this species seek harborage higher within rooms than do German cockroaches. The ootheca is small, only 5 mm long, with an average of 18 embryos and an incubation time of 35-80 days. Females deposit their oothecae by affixing them to furniture, in closets, on or behind picture frames, and in bedding. Transporting S. longipalpa with furniture to new locales is common. Although this species occurs with other cockroaches in homes, the German cockroach often outcompetes it within a few months.
German cockroach (Blattellagermanica)
This cockroach also is known as the steamily in Great Britain. It is believed to have originated in northern or eastern Africa, or Asia, and has spread from there via commerce. The German cockroach is considered to be the most important domestic pest species throughout the developed world. Adults are about 16 mm long, with two dark, longitudinal bands on the pronotum (Fig. 3.10). It requires warm (optimally 30-33°C), moist conditions near adequate food resources. It primarily inhabits kitchens and pantries, with secondary foci in bathrooms, bedrooms, and other living spaces in heavily infested structures. Although this species is nocturnal, like most other cockroaches, some individuals may be seen moving about on walls and in cupboards during the daylight hours where infestations are heavy. Their wing musculature is vestigial, making them unable to fly except for short, gliding, downward movements. B. germanica does not readily move between buildings; however, it does occur in garbage collection containers and outbuildings near heavily infested structures.
The German cockroach has a high reproductive potential. Females produce an ootheca (6-9 ram) containing about 30 embryos within 7-10 days after molting to the adult, or about 2-3 days after mating. The female carries the egg case until a few hours before hatching of the nymphs, preventing access of any oothecal parasitoids or predators. Oothecae are produced at intervals of 20- 25 days, with a female producing 4-8 oothecae during her lifetime. Nymphs complete their development in 7- 12 weeks.
This species is the main cockroach pest in most households and apartment complexes. Control is difficult, in part because of their movement between apartments through plumbing chases in shared or adjacent walls. Researchers studying over 1,000 apartments in Florida concluded that the median number of cockroaches per apartment was >13,000. This high biotic potential makes this species a major nuisance, as well as a pest with implications for human health.
Asian cockroach (Blattella asahinai)
The Asian cockroach is closely related to the German cockroach, from which it is difficult to distinguish morphologically. In fact, Asian and German cockroaches are capable of hybridizing and producing fertile off’- spring, which further complicates their identifications. Techniques have been developed to differentiate these two species and their hybrids based on cuticular hydrocarbons in the waxy layer of the integument.
Despite their morphological similarity, B. asahinai differs from B. germanica in several aspects of its behavior and ecology. It is both a feral and a peridomestic species. Nymphs of the Asian cockroach commonly occur, sometimes in large numbers, in leaf litter and in areas of rich ground cover or well-maintained lawns. Unlike the German cockroach, the adults fly readily and are most active beginning at sunset, when they fly to light-colored walls or brightly lit areas. This behavior can make invasion a nightly occurrence in homes near heavily infested areas. Flight does not occur when temperatures at sunset are below 21°C.
Like those of the German cockroach, Asian cockroach females carry their oothecae until shortly before they are ready to hatch. The ootheca is similar in size and contains the same number of embryos as does that of the German cockroach (38-44). Nymphs are smaller than their B. germanica counterparts and are somewhat paler in appearance. Development from egg to adult requires about 65 days, with females producing up to six oothecae during their life span. Adults also are slightly smaller than those of B. germanica (average of 13 vs 16mm).
The Asian cockroach was first described in 1981 from specimens collected in sugar-cane fields on the Japanese island of Okinawa. When it was first discovered in the United States in 1986, the Asian cockroach was found only locally in three counties in Florida, from Tampa to Lakeland; populations already had become established, with densities as high as 250,000 per hectare. By 1993, this species had spread to at least 30 Florida counties and had infested citrus groves throughout the central part of the state. It feeds on succulent early growth of citrus nursery stock, tassels of sweet corn, strawberries, cabbage, tomatoes, and other agricultural products, although there has been no evidence of significant economic damage.
Infestations of apartments by B. asahinai have become common in central Florida. This cockroach also has become an increasing problem in warehouses, department stores, hotels, fast-food establishments, automobile dealerships, and other businesses with hours of operation that extend beyond dusk.
Surinam cockroach (Pycnoscelus surinamensis)
This species is believed to have originated in the Indo- Malayan region. It commonly occurs in the southeastern United States from North Carolina to Texas. The adults are fairly stout, 18-25 mm in length, with shiny brown wings and a black body (Fig. 3.11). Nymphs characteristically have shiny black anterior abdominal segments, whereas the posterior segments are dull black and roughened. In North America this species is unusual in that it is parthenogenetic, producing only female offspring; elsewhere both males and females are found. The ootheca is 12-15 mm long, is poorly sclerotized, and contains about 26 embryos. Oothecae are retained inside the genital chamber, from which the nymphs emerge in about 35 days. Females produce an average of three oothecae and live about 10 months in the laboratory. This cockroach commonly burrows into compost piles and the thatch of lawns. Transfer
of fresh mulch into the home for potting plants can result in household infestations. Adult females fly and are attracted to light. They are most likely to be noticed by homeowners at night when they fly into brightly lit television screens. This species commonly is transported in commercial mulch to more temperate areas of the United States, where it has been known to infest greenhouses, indoor plantings in shopping malls, and ZOOS.
Cuban cockroach (Panchlora nivea)
This medium-sized cockroach (22-24 mm in length) is unusual in that the adults are pale green. The nymphs are dark brown and are found in leaf litter and decaying wood piles. Adults are strong fliers and are attracted to lights. Panchlora nivea is believed to be native to the Caribbean basin, Mexico, Central America, and northern South America. In the United States it occurs commonly in Florida and coastal Louisiana and Texas. This cockroach often is seen in the evening, resting on windows and glass patio doors, apparently drawn to the brightness of indoor lighting.
PUBLIC HEALTH IMPORTANCE
Cockroaches infesting human dwellings and workplaces represent a more intimate and chronic association than do most other pests of medical/veterinary importance. High populations of any cockroach species may adversely affect human health in several ways. These include contamination of food with their excrement, dissemination of pathogens, induced allergies, psychological stress, and bites. Mthough documentation of bites is limited, there are reports of cockroaches feeding on fingernails, eyelashes, skin calluses of hands and feet, and food residues about the faces of sleeping humans, causing blisters and small wounds (Roth and Willis 1957, 1960). There are other accounts of bites around the mouths of infants in heavily infested homes and even in hospitals. American and Australian cockroaches are the most often implicated species. Bites by the Oriental cockroach have resulted in inflammation of the skin, degeneration of epithelial cells, and subsequent necrosis of the involved tissues.
While many individuals develop a tolerance for cockroach infestations, others may experience psychological stress. The level of stress tends to be proportional to the size of the cockroaches and the magnitude of the infestation. An aversion to cockroaches may be so strong that some people become irrational in their behavior, imagining a severe infestation even when there is none. This illusion of abundant cockroaches has caused some families to move out of their homes. High cockroach populations also produce a characteristic odor that can be unpleasant or even nauseating to some people. Foodstuffs may become contaminated with the excrement of cockroaches, which, on subsequent ingestion, may cause vomiting and diarrhea.
The presence of cockroaches in homes does not necessarily imply poor housekeeping. Peridomestic species such as the American and the Oriental cockroach commonly infest municipal sewage systems or septic tanks and may move into homes through sewage lines. Any of the Periplaneta species may develop high outdoor populations, inducing individuals to seek less crowded environments. At such times, they often enter homes through attic vents, breaches in construction joints, or through crawl spaces. This tends to occur in early fall. While they are active at night the smokybrown cockroach, Asian cockroach, and feral wood roaches (Parcoblatta species) often find their way into even the best-kept homes. Adults frequently alight on doors illuminated by entrance lights, or on window screens of lighted rooms. Entrance is gained once the door is opened or by squeezing past window-screen frames.
Poor housekeeping and unsanitary conditions contribute significantly to cockroach infestations. The German cockroach and, to a lesser degree, the brownbanded cockroach are the principal bane of apartment dwellers. Their survival is enhanced by crowded living quarters, associated clutter, and the accumulated organic debris associated with food preparation. Construction practices used to build apartment complexes (e.g., common wiring ducts, sewage lines, and refuse areas) can contribute to the spread of cockroaches in multiunit dwellings.
PATHOGENIC AGENTS
The significance of cockroaches in public health remains controversial despite the logical assumption that they play a role in transmitting pathogenic agents. Given that cockroaches are so closely associated with humans and poor sanitation, the potential for acquiring and mechanically transmitting disease agents is very real. They are capable of transmitting microorganisms (Fig. 3.12) and other disease agents indirectly by contaminating foods or food preparation surfaces.
Table II lists pathogenic organisms that have been isolated from cockroaches in domestic or peridomestic environments. At least 32 species of bacteria in 16 genera are represented. These include such pathogens as Bacillus subtilis, a causative agent of conjunctivitis; Escherichia coli and 9 strains of Salmonella, causative agents of diarrhea, gastroenteritis, and food poisoning; Salmonella typhi, the causative agent of typhoid; and 4 Proteus species, which commonly infect wounds. These isolations primarily have involved American, German, and Oriental cockroaches. Cockroaches also have been found harboring the eggs of 7 helminth species, at least 17 fungal species, 3 protozoan species, and 2 strains ofpoliomyelitic virus (Brenner et al,. 1987; Koehler et al., 1990, Brenner 1995). Researchers in Costa Rica have shown that Australian, American, and Madeira cockroaches become infected with the protozoan Toxoplasmagondii after eating feces of infected cats. This suggests the possibility of cockroach involvement in the maintenance and dissemination of this parasite, which causes toxoplasmosis in humans, cats, and other animals.
Mthough many pathogens have been recovered from natural populations of cockroaches, this does not necessarily mean that cockroaches serve as their vectors. Isolation of pathogens from cockroaches simply may be
indicative of the natural microbial fauna and flora in our domestic environment. Under certain circumstances, however, cockroaches have the potential for serving as secondary vectors of agents that normally are transmitted by other means. Anecdotal accounts associating diseases in humans with the occurrence of cockroaches and microbes lend some credence to the hypothesis that these pests can serve as vectors. Burgess (1982) reported the isolation from German cockroaches of a serotype of S. dysenteriae that was responsible for an outbreak of dysentery in Northern Ireland. Mackerras and Mackerras (1948) isolated S. bovis-morbificans and S. typhimurium from cockroaches captured in a hospital ward where gastroenteritis, attributed to the former organism, was common. In subsequent experimental studies, Salmonella organisms remained viable in the feces of cockroaches for as long as 40 days postinfection (Mackerras and Mackerras 1949). Some of the most compelling circumstantial evidence suggesting that cockroaches may be vectors was noted in a correlation between cases of infectious hepatitis and cockroach control at a housing project during 1956- 1962 in southern California (Tarshis 1962). The study area involved more than 580 apartments and 2800 persons; 95% of the apartments had German cockroaches and a lesser infestation of brownbanded and Oriental cockroaches. After pest control measures were initiated, the incidence of endemic infectious hepatitis decreased for i year. When treatments were discontinued during the following year because the insecticide was offensive to apartment dwellers, the cockroach population increased, accompanied by a corresponding increase in the incidence of hepatitis. Effective control measures were applied for the following 2 years, and cockroach populations and cases of infectious hepatitis dropped dramatically while hepatitis rates remained high in nearby housing projects where no pest control measures were conducted.
INTERMEDIATE HOSTS
Cockroaches can serve as intermediate hosts for animal parasites (Table III). Roth and Willis (1960) published an extensive list of biotic associations between cockroaches and parasitic organisms that potentially infest humans. The eggs of seven species of helminths have been found naturally associated with cockroaches. These include hookworms (Ancylostoma duodenale and Necator americanus), giant human roundworm (Ascaris lumbricoides), other Ascaris species, pinworm (Enterobius vermicularis), tapeworms ( Hymenolepis species), and the whipworm Trichuris trichuria. Development of these helminths in cockroaches has not been observed. These relationships probably represent incidental associations with the omnivorous feeding behavior of cockroaches.
However, cockroaches may serve as potential reservoirs and possible vectors through mechanical transfer in areas where a high incidence of these pathogens in humans is accompanied by substantial cockroach infestations. Human infestations by spirurid nematodes associated with cockroaches are known only for the cattle gullet worm (Gongylonema pulchrum) in the United States, Europe, Asia, and Africa and for the stomach worm Abbreviata caucasia in Africa, Israel, Colombia, and Chile. Human cases involving these parasites are rare and cause no pathology.
COCKROACH ALLERGIES
It is only in recent years that the importance of cockroach allergies has been recognized. Allergic reactions result after initial sensitization to antigens following inhalation, ingestion, dermal abrasion, or injection. Allergens produced by cockroaches are rapidly being recognized as one of the more significant indoor allergens of modernized societies. Among asthmatics, about half are allergic to cockroaches. This rate is exceeded only by allergies to house-dust mites. Sensitivity to cockroaches also affects about 10% of nonallergic individuals, suggesting a subclinical level of allergy.
Symptoms exhibited by persons allergic to cockroaches are similar to those described by Wirtz (1980), who reported on occupational allergies in entomologists. They include sneezing and a runny nose, skin reactions, and eye irritation in about two-thirds of the cases. In the more severe cases, individuals may experience difficulty breathing or, even more alarming, anaphylactic shock following exposure to cockroaches. Such allergic reactions can be life-threatening (Brenner et al., 1991).
In recent years, research has focused on determining the specific components of cockroaches that cause allergy. Laboratory technicians exhibit strong allergies to cast skins and excrement of German cockroaches, whereas most patients seen at allergy clinics react primarily to cast skins and whole-body extracts of German cockroaches. Once an individual has become hypersensitized, he or she may experience severe respiratory distress simply by entering a room where cockroaches are held.
Several proteins that can cause human allergies have been identified in the German cockroach. Different exposure histories are likely to result in allergies to different proteins. Cast sldns, excrement, and partially consumed food of cockroaches, in addition to living cockroaches, all produce allergenic proteins. Some are extremely persistent and can survive boiling water, ultraviolet light, and harsh pH changes, remaining allergenically potent for decades. Traditionally, whole-body extracts have been used to screen for allergens in sldn tests and in bronchial challenges for diagnosing cockroach allergies
(Fig. 3.13). However, use of more specific antigens that become aerosolized in cockroach-infested homes may be more appropriate, as this is likely to be the sensitizing material. Studies with laboratory colonies have shown that a population of several thousand German cockroaches produced several micrograms of aerosolized proteins in 48 hr. Consequently, the presence of cockroaches may have profound respiratory implications for asthmatic occupants of infested structures. For a general discussion on aerosolized arthropod allergens, see Solomon and Mathews (1988).
Development of an allergy to one insect species can result in broad cross-reactivity to other arthropods, including shrimp, lobster, crab and crawfish, sowbugs (isopods), and house-dust mites. Chronic indoor exposure to cockroach allergens, therefore, may have significant and widespread effects on human health.
VETERIN/ dkY IMPORTANCE
Cockroaches serve as intermediate hosts for a number of parasitic worms of animals (Table III). Most of these relationships are of no economic importance. The majority of the parasites are nematodes in the order Spirurida, all members of which use arthropods as intermediate hosts. Species infesting dogs and cats, among other hosts, attach to the mucosa of the gastrointestinal tract, where erosion of tissue may occur at the points of attachment.
Although serious damage seldom occurs, anemia and slow growth may result. Several cockroach-associated nematodes occur in Europe and North America. The esophageal worms Physaloptera rara and P. praeputialis are the most widespread species in the United States. They develop in the German cockroach, field crickets, and several species of beetles.
Poultry also are parasitized by nematodes which undergo development in cockroaches. The Surinam cockroach is the intermediate host for the poultry eye worms Oxyspirura mansoni and O. parvorum. Both occur in many parts of the world. In the United States, their distribution is limited to Florida and Louisiana. The German cockroach has been incriminated as the intermediate host for chicken and turkey parasites, including the stomach worms Tetrameres americana, T. fissispina, and Cyrnea colini; C. colini also develops in the American cockroach. C. colini apparently causes no significant damage to poultry, but Oxyspirura species can cause pathology ranging from mild conjunctivitis to severe ophthalmia with seriously impaired vision. T. fissispina can cause severe damage to the proventriculus of infested birds.
Several nematode parasites of rats and cattle utilize cockroaches as intermediate hosts (Table III). These include G. neoplasticum and Mastophorus muris in rodents. Both genera occur widely in the United States, where they cause no known pathological problems. The gullet worm of cattle, G. pulchrum, has been shown experimentally to undergo development in the German cockroach, although the usual arthropod hosts are coprophagous beetles.
Exotic zoo animals also can become infested with parasitic nematodes for which cockroaches serve as possible intermediate hosts. Protospirura bonnei and P. muricola, for example, have been found in cockroaches collected in cages of monkeys. In a case of “wasting disease” in a colony of common marmosets, more than 50% of German and brownbanded cockroaches captured in the animal room in which they were housed contained the coiled larvae of Trichospirura leptostoma in muscle cells (Beglinger et al., 1988).
Acanthocephalans (thorny-headed worms) commonly infest primates in zoos and research facilities. Prosthenorchis elegans and P. spirula occur naturally in South and Central America. Their natural intermediate hosts are unknown. In captivity, primates become infected after eating any of several cockroach species in which the intermediate stages of the parasite have completed development. Heavily infested primates frequently die within a few days. The proboscis of acanthocephalan adults commonly penetrates the intestines of the primate host, causing secondary infections, perforation of the gut wall, and peritonitis.
One pentastomid (tongue worm), Raillietiella hemidactyli, develops in cockroaches and reptilian hosts. In Singapore, infested geckos are a common occurrence in houses where heavy infestations of R. hemidactyli larvae have been found in American cockroaches. Remnants of cockroaches are found commonly in the guts of these lizards.
For additional information on the veterinary importance of cockroaches, see Chitwood and Chitwood (1950), Roth and Willis (1957), Levine (1968), and Noble and Noble (1976).
PREVENTION AND CONTROL
Traditionally, cockroaches have been controlled using a variety of toxic chemicals applied as residual pesticides to harborage sites or areas frequented by foraging individuals (see Ebling, 1975 and Rust et. al., 1995). Most materials are neurotoxins that disrupt the nervous system, causing locomotory and respiratory failure. These include organophosphates, carbamates, botanicals such as pyrethrins, and pyrethroids. Formulations include wettable powders, emulsifiable concentrates, crack-andcrevice aerosols, dusts, and baits. Several other materials with different modes of action also are currently in use. When ingested, boric acid (delivered as a fine powder or a dilute solution) damages the gut epithelium of cockroaches and kills them by interfering with nutrient absorption. Inorganic silica dust is absorptive, reducing cuticular lipids and causing desiccation. Active ingredients with other modes of action, such as hydramethylnon and sulfluramid, are metabolic inhibitors which disrupt the conversion of food to energy.
The use of baits containing many of the active ingredients mentioned above have been used extensively to control cockroaches. These baits are used indoors in the form of child-resistant bait stations to reduce human exposure. Other bait formulations of gels or pastes are used in crackand- crevice treatments, making them inaccessible to children and pets. Scatter baits are commonly used outdoors to treat mulches and other landscaping materials that harbor cockroaches.
Insect growth regulators (IGRs) can be used to prevent cockroaches from reaching maturity. Two commonly used IGRs are juvenile hormone analogs and chitin synthesis inhibitors. Juvenile hormone analogs regulate morphological maturation and reproductive processes. They are highly specific to arthropods, have very low mammalian toxicity, and are effective at exceptionally low rates of application. Such compounds include hydroprene and fenoxycarb. Chitin synthesis inhibitors prevent normal formation of chitin during molting. These compounds cause many of the affected nymphs to die during the molting process. Males that survive to the adult stage often have reduced life expectancies, whereas females tend to abort their oothecae.
Integrated pest management, which incorporates various control techniques, has contributed significantly to successful control of cockroaches. This approach uses nontoxic agents, such as sticky traps, vacuum devices, diatomaceous earth, or silica-gel repellents and desiccants, and manipulation of harborage sites to reduce or prevent infestations. Desiccants and dusts should be used only in geographic areas or situations with relatively low humidity; high humidity causes these materials to clump and lose their effectiveness. Building designs and construction techniques can significantly influence cockroach survival. By manipulating microclimates in discrete areas of structures frequented by cockroaches, homes and other buildings can be rendered less hospitable to pest species while at the same time greatly reducing aerosolized allergens. Nontoxic repellents can be used to deny access of cockroaches to specific areas.
Biological control of cockroaches has drawn increased attention in recent years. Among the natural agents that have been investigated are parasitic wasps, nematodes, and sporulating fungi. Females of the eulophid wasp Aprostocetus hagenowii and the evaniid wasp Comperia merceti deposit their eggs in the oothecae of certain peridomestic cockroaches. Major shortcomings in utilizing these wasps are difficulties involved in their mass production and the fact that they do not completely eliminate cockroach infestations. However, A. hagenowii has been shown to reduce populations of the peridomestic Periplaneta species following inundative or augmentative releases of this wasp. C. merceti parasitizes oothecae of the brownbanded cockroach and is the only known parasitoid of a domestic species. The use of parasitic nematodes (e.g., Steinernema carpocapsae) and several fungal pathogens that have been isolated from cockroaches has not yet proved to be effective as a practical management tool. Another drawback to their use is the allergenic nature of several components of nematodes and many sporulating fungi that can become airborne and, upon inhalation, cause asthmatic responses in humans.
Models have been developed for predicting population foci of peridomestic cockroaches based on physical characteristics of residential properties (Smith et al., 1995). However, the use of such models is limited by the scope of the data base used in its development and the complexity of the model itself. The use of traps to detect foci of cockroaches and the analysis of trap counts to determine cockroach abundance and distributional patterns can be helpful in assessing the extent of infestations and monitoring the effectiveness of control programs (Brenner and Pierce 1991).