21.03.2015 Views

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

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

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

SHOW MORE
SHOW LESS

Create successful ePaper yourself

Turn your PDF publications into a flip-book with our unique Google optimized e-Paper software.

Fig. 8.7 Nymphs of Cryptocercus punctulatus cooperatively feeding on a sliver of wood. Photo by<br />

C.A. Nalepa.<br />

wagging, or to scrape it off by dragging it along the side<br />

of the gallery (CAN, unpubl. obs.).<br />

It should be noted that in Cryptocercus there are cooperative<br />

as well as competitive <strong>behavior</strong>s among nymphs<br />

when procuring food. Wood is not only nutritionally<br />

poor and difficult to digest, but physically unyielding.<br />

Like young nymphs in aggregations, early developmental<br />

stages of Cryptocercus may need the presence of conspecifics<br />

to help acquire meals when they begin including<br />

wood in their diet. Nymphs have been observed feeding<br />

cooperatively on wood slivers pulled free by both siblings<br />

(Fig. 8.7) and adults (Nalepa, 1994; Park and Choe,<br />

2003a).<br />

Cost of Parental Care<br />

Most cockroaches that exhibit parental care are subject to<br />

risks associated with brood defense and invest time in<br />

taking care of offspring. Other costs vary with the form<br />

and intensity of parental care. Brooding, for example, is a<br />

small investment on the part of the female in relation to<br />

potential returns (Eickwort, 1981). In females that carry<br />

offspring on their bodies, the burden may hinder locomotion<br />

and thus the ability to escape from predators. Energy<br />

expended on nest construction can detract from a<br />

parent’s capacity for subsequent reproduction in those<br />

species where parental care occurs in excavated burrows.<br />

Insects that utilize nests may also invest time and energy<br />

in provisioning and hygienic activities (Tallamy and<br />

Wood, 1986). Feeding offspring on bodily secretions may<br />

drain stored reserves otherwise devoted to subsequent<br />

bouts of oogenesis. The metabolic expenditure may be<br />

particularly high in wood-feeding species, whose diet is<br />

typically low in nitrogenous materials. The high cost of<br />

parental care in Cryptocercus may account for their functional<br />

semelparity (Nalepa, 1988b), and has been proposed<br />

as a key precondition allowing for the evolution of<br />

eusociality in an ancestor they share with termites (Chapter<br />

9). It is of interest then, that, another wood-feeding<br />

cockroach (Salganea matsumotoi) that lives in biparental<br />

groups and is thought to exhibit extensive parental care<br />

appears to have more than one reproductive episode<br />

(field data) (Maekawa et al., 2005).<br />

In insects that do not nest in their food source, providing<br />

care to young may conflict with feeding opportunities,<br />

particularly in species whose diet consists of dispersed<br />

or ephemeral items that require foraging over<br />

substantial distances. One solution to is to carry one<br />

brood while gathering nutrients for subsequent brood<br />

development (Tallamy, 1994). To test this hypothesis, it is<br />

necessary to determine (1) if females feed while externally<br />

carrying nymphs, and (2) if females carrying nymphs are<br />

concurrently developing their next set of eggs, incubating<br />

eggs in the brood sac, or building reserves for the next<br />

brood. We found relevant information on two species. A<br />

Pseudophoraspis nebulosa female caught in the field with<br />

numerous neonates clinging to the undersurface of her<br />

abdomen was dissected, and her brood sac was empty<br />

(Shelford, 1906a). In Tho. porcellana, newly hatched<br />

nymphs remain in association with their mother for 45<br />

days. After partition another ootheca is formed in 15 to<br />

20 days, and gestation takes 45–52 days. There is therefore<br />

a period of time when the female is both internally<br />

incubating an ootheca in her brood sac and externally<br />

carrying nymphs on her back. However, these are sluggish<br />

insects that remain stationary in the leaves on which they<br />

148 COCKROACHES

Hooray! Your file is uploaded and ready to be published.

Saved successfully!

Ooh no, something went wrong!