15 
Life History of Pinewoods Darter 
E. kennicotti, E. smithi), and males of these species were also strongly 
territorial at spawning. Another tendency common in territorial percids 
is that females usually outnumber males (Lachner et al. 1950), as also 
noted in E. mariae (Table 4). The difference between number of males 
(99) and females (204) was highly significant (chi-square P < 0.001, 
expected ratio 1:1). Although we did not observe territoriality, the 
larger size of mature males, preponderance of females, and sexual di- 
chromatism during the breeding season suggest that such behavior 
occurs. 
The oldest E. mariae were 36 months (age class 2). Two fish had just 
formed a third annulus. It is doubtful that E. mariae lives four years. 
The majority (58.7%) were in age class 0, 30.1% in age class 1, and 
11.2% in age class 2 (Table 4). Making the same assumptions as Page 
(1974), we found that the survival values as based on Table 5 indicated a 
low survival rate in the third year of life. Males had a 53% survival to 
age class 1 and a 10% survival to age class 2. Females had a 54% survi- 
val to age class 1, but a survival to age class 2 (25%) that was much 
greater than that of males (Table 5). Male E. mariae exhibited a faster 
growth rate and a higher mortality rate in the older ages than did 
females. 
Food Habits . — Based on stomach contents of 321 fish (11 to 59 mm 
SL), 13 animal taxa were identified (Table 6). Only 25 (7.8%) stomachs 
were empty. The majority of E. mariae ate larval dipterans (in 86.8% of 
stomachs). Other commonly ingested invertebrates were ephemerop- 
teran nymphs (28.0%), trichopteran larvae (26.0%), and plecopteran 
nymphs (22.0%) (Table 6). Food habits of E. mariae were similar to 
those of the few Southeastern darters studied (see references in Page 
1983). 
Seasonal variation in diet was evident (Table 6). Dipterans, primar- 
ily Chironomidae and Simuliidae, were eaten in all months. Chirono- 
mids were the most commonly occurring food in all months except May, 
June, and October, when simuliids were most frequent. Ephemerop- 
terans were important in May and June. Trichopterans, mainly Hydro- 
psychidae, were common in April and plecopterans were commonly eaten 
in January, March, and December. The number of taxa represented in 
the stomachs per fish per month was lowest in the coldest months, 
November through March (3-5 taxa) excepting December, and highest 
during summer through mid-fall (6-8 taxa) (Table 6). 
Prey taxa differed by fish size groups (Table 7). Chironomids were 
most important in the diet of small fish (< 40 mm). Larger darters also 
ate large numbers of chironomids but consumed even more of the larger 
simuliids. Plecopterans, ephemeropterans, and trichopterans were eaten 
