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

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

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Fig. 4.2 Feeding and drinking cycles in relation to the reproductive<br />

cycle of the wood cockroach Parcoblatta fulvescens.<br />

Filled circles, water consumption; open squares, food consumption;<br />

EC, egg case formation; ECD, egg case deposition.<br />

From Cochran (1986b), courtesy of Donald G. Cochran, with<br />

permission from Elsevier Press.<br />

but no water, they lived for about the same period of time<br />

as those deprived of both. If they are provided with water,<br />

most lived longer. Some species can live for 2 to 3 mon<br />

on water alone, and others significantly longer. Virgin females<br />

of Eublaberus posticus live an average of 360 days on<br />

water alone, whereas starved but mated females can live<br />

an average of 8 mon and are even able to produce 1 or 2<br />

litters, yielding about 26 young. One female mated at<br />

emergence was starved for 252 days, during which time<br />

she produced 2 litters totaling 50 nymphs. She was then<br />

given food on day 252 (and thereafter), mated again 4<br />

days later, and lived an additional 525 days, producing 5<br />

more oothecae from which 24, 18, 5, 1, and 0 nymphs<br />

hatched. Although this female had been starved for the<br />

first 8 mon of adult life, after food was made available she<br />

managed to give birth to a total of 98 offspring, which is<br />

about normal for this species (Roth, 1968c).<br />

There is a significant difference in starvation resistance<br />

between males and females in cockroach species exhibiting<br />

sexual dimorphism in body size. In Table 4.3, males<br />

and females are of similar size only in Neostylopyga rhombifolia,<br />

Eurycotis floridana, and Nauphoeta cinerea; in<br />

these cases, survival of males and females is similar. In the<br />

remaining species males are significantly smaller than females<br />

and are more vulnerable to starvation. A larger<br />

body size is correlated with bigger fat bodies and their accumulations<br />

of carbohydrates, lipids, and uric acid; these<br />

reserves can be rapidly mobilized on demand (Mullins<br />

and Cochran, 1975b; Downer, 1982). The nutrients and<br />

water housed in developing oocytes are additional resources<br />

available to starving females. The strategy for a<br />

food-deprived female of P. americana seems to involve resorption<br />

of yolk-filled eggs, storage of their yolk proteins,<br />

and then rapid incorporation of protein into eggs when<br />

feeding re-ensues (Roth and Stay, 1962b; <strong>Bell</strong>, 1971; <strong>Bell</strong><br />

and Bohm, 1975).<br />

A variety of digestive attributes help cockroaches<br />

buffer food shortages. The large crop allows an individual<br />

to consume a substantial quantity of food at one time.<br />

This bolus then acts as a reservoir during periods of fasting.<br />

When fully distended with food, the crop is a pearshaped<br />

organ about 1.5 cm in length and 0.5 cm at its<br />

widest part (in Periplaneta australasiae). It extends back<br />

to the fourth or fifth abdominal segment, crowding the<br />

other organs and distending the intersegmental membranes.<br />

A meal may be retained in the crop for several<br />

days (Abbott, 1926; Cornwell, 1968). Solid food is also retained<br />

in the hindgut of starving P. americana for as long<br />

as 100 hr, although the normal transit time is about 20 hr<br />

(Bignell, 1981); this delay likely allows microbial biota to<br />

more thoroughly degrade some of the substrates present,<br />

particularly fiber. The functional significance of intestinal<br />

symbionts increases in times of food deficiency and helps<br />

to maintain a broad nutritional versatility (Zurek, 1997).<br />

A starving cockroach is thus indebted to its microbial<br />

partners on two counts: first, for eking out all possible nutrients<br />

in the hindgut, and second, for mobilizing uric<br />

acid stored in the fat body (Chapter 5). When food is<br />

again made available, starved P. americana binge. After<br />

starving for 13 days the amount of food consumed rose<br />

to five times the normal level, then leveled off after approximately<br />

20 days. Greater consumption was accomplished<br />

by larger and longer meals, not by increasing the<br />

number of foraging trips (Rollo, 1984a).<br />

PLANT-BASED FOOD<br />

There is little evidence that any cockroach species is able<br />

to subsist solely on the mature green leaves of vascular<br />

plants. There are reports of occasional herbivory, such as<br />

that of Crowell (1946), who noted that the small, round<br />

leaves of the aquatic plant Jussiaca are included in the diet<br />

of Epilampra maya. Often, cockroaches that appear to be<br />

feeding on green leaves are actually eating either a small,<br />

dead portion at the leaf edge or around a hole, or other<br />

material on the leaf (WJB, unpubl. obs.). To test the extent<br />

to which tropical cockroaches include fresh vegetation<br />

in their diets, WJB set up a series of two-choice tests<br />

in laboratory cages at La Selva Biological Station in Costa<br />

Rica. Ten species of cockroaches were tested: Capucina<br />

sp., Cariblatta imitans, Epilampra involucris, Ep. rothi, Ep.<br />

unistilata, Latiblattella sp., Imblattella impar, Nahublattella<br />

sp., Nesomylacris sp., and X. hamata. The insects were<br />

offered a choice of green leaves versus dead leaves of the<br />

same plant species; only leaves eaten readily by local Or-<br />

66 COCKROACHES

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