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

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

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Fig. 7.7 Diagram of presumed sequence of stages in the evolution of ovoviviparity from oviparity in<br />

two subfamilies of Blattellidae. Note the difference in the orientation of the ootheca between the two<br />

subfamilies. Current evidence suggests that the oothecal rotation exhibited by the Blattellinae and by<br />

the ovoviviparous Blaberidae originated in a common ancestor.<br />

dwelling insect to carry or internalize must be either<br />

squashed dorsoventrally or rotated so that it is as flat as<br />

the female (Fig. 7.7). Intermediate stages in parity mode<br />

are conspicuous in the Pseudophyllodromiinae. Sliferia<br />

is considered ovoviviparous; nonetheless the egg case is<br />

partially exposed while it is carried. Initially it was<br />

thought that these females were collected while still forming<br />

the ootheca. Now this condition is considered the<br />

norm, and points up the continuum of reproductive<br />

modes in this subfamily (Roth, 2003b).<br />

All species in the ovoviviparous family Blaberidae<br />

carry a rotated egg case in their brood sac and are thought<br />

to have evolved from a Blattella-like ancestor (Roth and<br />

Willis, 1955c; Roth, 1967a; Mullins et al., 2002). Except for<br />

retraction of the egg case into the body, B. germanica exhibits<br />

all characteristics of an ovoviviparous cockroach<br />

(Roth and Willis, 1958a; Roth, 1970a). The oothecal case<br />

is thinner and less darkly colored than in other oviparous<br />

cockroaches, there is flow of water and other materials between<br />

mother and unhatched offspring, and oogenesis is<br />

suspended while females are carrying egg cases. The evolution<br />

of ovoviviparity would require only a minor transition<br />

from that starting point. Ovoviviparity evolved independently<br />

two or three times in cockroaches, but only<br />

in the blattellid/blaberid lineage (Roth, 1970a, 1989a):<br />

once in the Pseudophyllodromiinae, and once or twice in<br />

the clade that includes Blattellinae and Blaberidae. Viviparity<br />

evolved once, in D. punctata of the monogeneric<br />

subfamily Diplopterinae. Some authors also include Calolampra<br />

or Phoetalia in this subfamily (Roth, 2003c), so<br />

these genera may be logical targets for comparative study.<br />

Worldwide, Blattellidae is the largest cockroach family<br />

with about 1740 described species; there are approximately<br />

1020 species of Blaberidae. The oviposition <strong>behavior</strong><br />

is known in relatively few genera and species of<br />

these two families (Roth, 1982a).<br />

Reduction and Loss of the Egg Case<br />

In most oviparous type A cockroaches, the ootheca is a<br />

hard, dark, stiff structure completely covering the eggs.<br />

The dorsal keel is structurally complex, and the outer covering<br />

contains calcium oxalate crystals. These crystals<br />

comprise 8–15% of the dry weight of the ootheca in P.<br />

REPRODUCTION 125

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