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Cockroache; Ecology, behavior & history - W.J. Bell

Cockroache; Ecology, behavior & history - W.J. Bell

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tactile stimulation in this species. Patrolling dominant<br />

males of N. cinerea tap members of their social group on<br />

the pronotum with their antennae, evoking a submissive<br />

posture in lower-ranking members (Ewing, 1972). Similarly,<br />

reflex immobilization in Blab. craniifer can result<br />

from antennal tapping of the pronotal shield by another<br />

individual (Gautier, 1967).<br />

COLOR<br />

As in many other insect groups, the suborder Blattaria encompasses<br />

species with both cryptic and conspicuous<br />

coloration. The former decreases the risk of detection,<br />

and the latter is often used in combination with chemical<br />

defenses and specific <strong>behavior</strong>s that discourage predators.<br />

Color patterns can vary considerably within a species,<br />

contributing to taxonomic difficulties (Mackerras,<br />

1967a), and in a few cockroaches color variation is correlated<br />

with geographic features, seasonal factors, or both.<br />

Two subspecies of Ischnoptera rufa collected at high elevations<br />

in Costa Rica and Mexico are darker than their<br />

counterparts collected near sea level (Hebard, 1916b).<br />

Adults of Ectobius panzeri in Great Britain are darker at<br />

higher latitudes, and females have a tendency to darken<br />

toward the end of the breeding season (Brown, 1952).<br />

Parcoblatta divisa individuals are typically dark in color,<br />

but a strikingly pale morph is found in Alachua County,<br />

Florida. No dark individuals were found in a series of<br />

several hundred specimens taken from this location, and<br />

the pale form has not been collected elsewhere (Hebard,<br />

1943). Color variation among developmental stages within<br />

a species may be associated with changing requirements<br />

for crypsis, mimicry, or aposematicism. Adults of Panchlora<br />

nivea, for example, are pale green, while the juvenile<br />

stages are brown (Roth and Willis, 1958b).<br />

Many cockroaches are dark, dull-colored insects, a<br />

guise well suited to both their cryptic, nocturnal habits<br />

and their association with decaying plant debris. Several<br />

species associated with bark have cuticular colors and<br />

patterns that harmonize with the backgrounds on which<br />

they rest. Trichoblatta sericea lives on Acacia trees, blending<br />

nicely with the bark of their host plant (Reuben,<br />

1988). Capucina rufa lives on and under the mottled bark<br />

of fallen trees and seems to seek compatibly patterned<br />

substrates on which to rest (WJB, pers. obs.). A cloak of<br />

background substrate enhances crypsis in some species.<br />

Female Laxta spp. may be encrusted with soil or a parchment-like<br />

membrane (Roth, 1992), and Monastria biguttata<br />

nymphs are often covered with dust (Pellens and<br />

Grandcolas, 2003).<br />

Not unexpectedly (Cott, 1940), there are dramatic differences<br />

in coloration between the cockroaches on the<br />

dayshift versus the nightshift. Day-active cockroaches<br />

tend to fall into three broad categories: first, the small, active,<br />

colorful, canopy cockroaches; second, the chemically<br />

defended, aposematically colored species; and third,<br />

those that are Batesian mimics of other taxa. Patterned,<br />

brightly colored insects active in the canopy in brilliant<br />

sunshine have a double advantage against predators. They<br />

are not only cryptic against colorful backgrounds, but<br />

they are obscured by rapidly changing contrast when<br />

moving in and out of sun flecks (Endler, 1978). A number<br />

of aerial cockroach species have translucent wing covers,<br />

tinted green or tan, that provide camouflage when<br />

they are sitting exposed on leaves (Perry, 1986).<br />

Among the best examples of aposematic coloration are<br />

in the Australian Polyzosteriinae (Blattidae). Nocturnal<br />

species in the group are usually striped yellow and brown,<br />

but the majority are large, wingless, slow-moving, diurnal<br />

cockroaches fond of sunning themselves on stumps<br />

and shrubs. They are very attractive insects, often metallically<br />

colored, or spotted and barred with bright orange,<br />

red, or yellow markings (Rentz, 1996; Roach and Rentz,<br />

1998). When disturbed, they may first display a warning<br />

signal before resorting to defensive measures. Platyzosteria<br />

castanea and Pl. ruficeps adults assume a characteristic<br />

stance with the head near the ground and the abdomen<br />

flexed upward at a sharp angle, revealing orange-yellow<br />

markings on the coxae and venter. Continued harassment<br />

results in the discharge of an evil-smelling liquid “so execrable<br />

and pungent that it drove us from the spot”<br />

(Shelford, 1912a). Elegant day-flying cockroaches in the<br />

genera Ellipsidion and Balta (Blattellidae) can be observed<br />

basking in the sun and exhibit bright orange colors<br />

suggestive of Müellerian mimicry rings (Rentz, 1996).<br />

<strong>Cockroache</strong>s in the genus Eucorydia (Polyphaginae) are<br />

usually metallic blue insects, often with orange or yellow<br />

markings on the wings (Asahina, 1971); little is known of<br />

their habits. The beautiful wing patterns of some fossil<br />

cockroaches are suggestive of warning coloration. Some<br />

Spiloblattinidae, for example, had opaque, black, glossy<br />

wings with red hyaline windows (Durden, 1972; Schneider<br />

and Werneburg, 1994).<br />

Several tropical cockroaches mimic Coleoptera in size,<br />

color, and <strong>behavior</strong>. This is evident in their specific<br />

names, which include lycoides, buprestoides, coccinelloides,<br />

dytiscoides, and silphoides. Shelford (1912a) attributes<br />

beetle-mimicry in the Blattaria to the similar body types<br />

of the two taxa. Both have large pronota and membranous<br />

wings covered by thickened elytra or tegmina.“Only<br />

a slight modification of the cockroach form is required to<br />

produce a distinctly coleopterous appearance.” Vršanský<br />

(2003) described beautifully preserved fossils of small,<br />

beetle-like cockroaches that were day active in Mesozoic<br />

forests (140 mya). Extant species of Prosoplecta (Pseudophyllodromiinae)<br />

(Fig. 1.3) have markedly convex oval or<br />

4 COCKROACHES

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