PRODUCTION AND USE OF SECRETIONS PASSED 
BY MALES AT COPULATION IN 
PIERIS PROTODICE (LEPIDOPTERA, PIERIDAE)* 
By Ronald L. Rutowski 1 
Department of Zoology 
Arizona State University 
Tempe, AZ 85287 
Introduction 
During copulation in many species of insects the male passes to 
the female sizeable quantities of accessory gland secretions with the 
sperm (Thornhill and Alcock, 1983). Recently it has been shown in 
the Lepidoptera that these secretions may represent a nutrient 
investment by the male that is used by the female in the production 
of eggs and in somatic maintenance (Boggs and Gilbert, 1979; 
Boggs, 1981; Boggs and Watt, 1981; Goss, 1977; Greenfield, 1982). 
Typically these secretions are contained within a spermatophore 
with the sperm or they may be passed as loose secretion. The 
perspective that these secretions may constitute a nutrient invest- 
ment by males has some important implications for how male and 
female lepidopterans should behave (Marshall, 1982; Rutowski, 
1982; see also, Gwynne, 1982). 
In spite of recent interest in these secretions there are few studies 
of the patterns of their production and use within and between 
species of Lepidoptera (e.g., Boggs, 1981; Greenfield, 1982; Rutow- 
ski et al., 1983). This paper reports data on patterns of variation in 
the quantity of material passed to females during copulation by 
males of the checkered white butterfly ( Pieris protodice Boisduval 
and LeConte) and the disappearance of these secretions from the 
female’s reproductive tract. The secretions are received by the 
female in the bursa copulatrix which in this butterfly species is 
composed of two sacs, the corpus bursa and the appendix bursa. 
These are connected in series with the appendix bursa being furthest 
from the copulatory opening. During copulation the appendix 
bursa is filled with a whitish fluid and the corpus bursa is filled with 
* Manuscript received by the editor March 5, 1984. 
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