1978] Wood — Parental Care in Guayaquila compressa 
143 
the pronotum, their cryptic coloration and lack of movement make 
them difficult to find on woody branches of the host plant (Wood 
1976b). The pronotum of female G. compressa is similar in shape 
and hardness to that of mature P. vittata, but the black females 
provide contrast to the green leaf background. Thus, rapid female 
dispersal producing a startle response in a predator such as an 
arboreal anole would be a viable alternative for an otherwise 
unprotected female. 
Herding of offspring by parent females within a host plant is 
unique to G. compressa. Female U. crassicornis and P. vittata 
remain with their offspring on the woody branches where eggs were 
deposited until nymphs reach maturity. Eggs of E. bactriana are 
deposited in leaves on herbaceous plants but nymphs move from 
these leaves and reaggregate on new ones after being deserted by 
females (Wood 1977a). Herding in G. compressa may permit 
enhanced exploitation of the host plant by reducing localized 
feeding damage. 
Alarm or escape behavior actively involves both female G. 
compressa and offspring, differing significantly from that of U. 
crassicornis or P. vittata. Alarm displays by females of the latter 
species are produced in response to injured nymphs or predators, 
but nymphs do not disperse from the feeding site, nor are they 
deserted by parent females (Wood 1976a, b). E. bactriana females 
place themselves between predator and offspring. Nymphs remain 
with females initially, but upon prolonged exposure to alarm 
pheromones or injured females, nymphs disperse from the leaf. 
Escape behavior of nymphs is also modified by the behavior of 
various ant species which provide predator protection (Wood 
1977a). Thus, the startle response produced by female G. compressa 
provides time to permit nymphal dispersal. Dispersed nymphs 
reaggregate with or without parent females and maintain an effec- 
tive nymphal escape response if the parent female is captured or dies 
from other causes. 
Females in a number of membracid species deposit eggs close to 
each other on a branch even when there appears to be an abundance 
of oviposition space, suggesting cooperative brood care (Wood, 
unpublished). G. compressa provides the first evidence to suggest an 
hypothesis for the adaptive nature of this cooperation. Normal 
herding behavior on a small host plant means 2 aggregations have a 
