TABLE 3.— Collections of breeding Etheostoma squamiceps. 
Locality Collection Date Remarks 
erguson Creek, Livingston Co., Ky. 24 March 1972 Spawning when collected. 
erguson Creek, Livingston Co., Ky. 23 March-20 May 1973 Spawning when collected. 
ig Creek, Hardin Co., Il. 15 April-3 May 1971 Spawning when collected. 
ig Creek, Hardin Co., Il. 31 March-20 May 1973 Spawning when collected. 
ickory Creek, Livingston Co., Ky. 27 March 1964 Female in extreme breeding condition. 
juck River, Coffee Co., Tenn. 28 March 1965 Male in extreme breeding coloration. 
ush Creek, Posey Co., Ind. 29 March 1963 Males and females in extreme breeding condition. 
Preceding spawning, males periodically left the 
est stone to court females. When attempting to lure 
female back to the stone, a male would swim near 
female, display his bold pattern and spread fins, 
nd wag his tail at or against the female. 
After joining a male under his nest stone, a female 
sually remained passive while the male courted her. 
he male was extremely active at this time, con- 
antly darting about, nudging the female with his 
1out, and occasionally bobbing his spinous dorsal 
n up and down, the contrasting dark and clear 
reas of which created a blinking effect. 
Eventually the female began moving about, arch- 
ig her head upward, and holding her snout near 
ie underside of the stone, apparently selecting a 
te for egg deposition. 
Both male and female assumed an inverted posi- 
on when laying and fertilizing eggs. After nosing 
bout the rock for a time, the female rolled to one 
de and simultaneously rose to press her venter 
yainst the underside of the nest stone. Usually almost 
nmediately after the female had inverted, the male 
led to one side and positioned himself tightly along- 
de and slightly overlapping the female, with a 
elvic fin across her nape, in an inverted head-to-head 
sition (Fig. 7). Pressed against the stone, the 
male began quivering slightly, barely moving for- 
ard as she did so, and laying a series of two to 
ve eggs on the stone. As she laid the eggs, the 
ale trembled and released sperm. Then both fish 
nmediately returned to a right-side-up position be- 
sath the stone for a short time before courting 
y the male began the sequence again. 
Fifteen seconds to several minutes elapsed be- 
veen inversions. Each inversion lasted less than 
seconds. The eggs were concentrated in a con- 
ricted area but never were laid on top of other 
3gs. Eggs were deposited as high as 35 mm above 
ie substrate. 
The presence of a second female with a spawning 
uir was tolerated, and in fact, both females were 
yurted by the male. No instances of more than one 
the females spawning at a time were observed, 
owever. 
At least three females spawned with one male 
| an aquarium, two with another. In one instance the 
awnings with different females occurred within 2 
hours of one another; in another instance the spawn- 
ings were 5 days apart. Eggs laid by a second or 
third female were added to those already in the 
nest, enlarging the concentrated egg mass. 
Counts and estimates of the numbers of eggs in 
20 nests of E. squamiceps found in Big and Ferguson 
creeks were 8, 90, 125, 125, 160, 200, 200, 200, 200, 
250, 275, 300, 450, 475, 475, 500, 850, 1100, 1500, and 
1500, the mean being 449. Several nests contained 
many more eggs than the largest number of mature 
ova (357) found in preserved females and, as in 
aquarium nests, contained eggs from more than one 
female. Eggs in nests averaged 1.8 mm in diameter. 
Males vigorously guarded the eggs. Whenever a 
nest stone in a stream was lifted for observation, a 
male was found underneath and usually swam away 
only when provoked. When a male did dart away, 
he usually returned soon after the rock was replaced. 
In all nests which were re-examined on a later date, 
the male had returned and was again guarding the 
nest. 
Aquarium-held males were constantly active while 
guarding eggs, darting back and forth and continu- 
ously brushing the eggs with their dorsal fins. The 
knobs on the tips of the dorsal spines of E. squamiceps 
males may be an adaptation to reduce the sharpness 
of the spines, thereby reducing the probability of 
rupturing eggs, during the nest-guarding time. Peri- 
odic nipping at the eggs presumably removed invad- 
ing organisms. The breeding pattern of the male 
was retained during the egg-guarding period. 
Nests were guarded against potential egg preda- 
tors, including crayfishes. Large crayfishes introduced 
into aquaria were driven from the nest stone by the 
male E. squamiceps as he repeatedly charged the 
crayfishes and bit them or hit them on the abdomen. 
A 63-mm male E. squamiceps succeeded in driving 
from his nest a 75-mm total length Orconectes ken- 
tuckiensis but not a 90-mm Cambarus laevis, both 
common crayfishes in Big Creek. 
Nests were never tended by females. Lake (1936: 
823) believed that only E. flabellare males provide 
parental care to their eggs. 
Some areas used as breeding grounds contained 
large numbers of nests. Separate nest stones of E. 
squamiceps were found as close as 200 mm from one 
another. Two large stones with two Catonotus nests 
