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Table 1
Definitions and summary statistics of measurements taken from nests constructed by male three-spined sticklebacks
Nest character
Definition
N*
Substratum weight
Number of threads (N threads)
Number of thread ends (N ends)
42
42
42
Total area (A
tot
Bulk area (A
bulk
Neatness index, In
Abulk
Atot
Compactness index, Ic
Nends
2(Nthreads)
Mean ( SD)
1.87 g (1.77g)
148.0
( 46.6)
64.9
( 38.0)
Range
0.005.78 g
37200
0170
39
18028577 mm2
39
4293523 mm2
42
0.77 ( 0.14)
0.481.00
39
0.45 ( 0.25)
0.121.00
* 42 males built nests. Data are reduced in some cases because nest area measurements could not be made from three nests.
Figure 1
Photographs of representative nests built by male sticklebacks during the study, with diagrammatic representations showing the various
quantitative measurements made for each nest.
393
Figure 2
The relationship between the two indices of nest structure for nests
built by male sticklebacks in our study.
modal distribution with peaks around 23 and 56 days; Figure 3, main). Early nesters (those taking 3 days to initiate
construction) built neater nests than late nesters (t test, t
2.60, df 38, p .013; Figure 3, inset), but the compactness
of nests built by early and late nesters did not differ (t test, t
0.84, df 34, P .40). Males exhibiting longer construction times built more compact (r .358, n 38, p .027)
nests and incorporated more deposited substratum (r .318,
n 42, p .040), but there was no significant correlation
between construction time and nest neatness (r .206, n
38, p .197).
Nest structure and male physiological traits
Perhaps surprisingly, neither male body length nor BCF correlated with nest neatness, compactness, or with the weight of
deposited substratum (length, 0.080 r .229, all p .150;
BCF, 0.281 r .072, all p .087). The RKW of nesting
males correlated positively both with the neatness (r .360,
n 38, p .026) and compactness (r .350, n 37, p
.036) of the nests they built, but not with the weight of substrate deposited (r .164, n 36, p .325). RSW correlated
negatively and significantly with the total weight of deposited
substratum (r .366, n 37, p .026) and with nest compactness (r .397, n 35, p .018), but only marginally
with nest neatness (r .267, n 36, p .11). HSI did not
Table 2
Characteristics of nest-building male sticklebacks used in the study
Male character
Mean ( SD)
Range
42
42
42
42
42
38
37
38
42
42
44.60
0.94
2.05
0.46
84.4
0.045
0.003
0.024
4.0
6.0
3955
0.601.79
1.352.94
0.410.53
15.170.5
0.0270.078
0.0010.005
0.0110.035
117
112
(
(
(
(
(
(
(
(
(
(
3.32)
0.25g)
0.24)
0.03)
35.1)
0.011)
0.001)
0.005)
2.9)
3.9)
394
Figure 3
The relationship between nest
neatness and the length of the
pre-building lag. The main figure shows the frequency distribution of observed pre-building lag periods (the time between arrival in the laboratory
and initiation of nest building)
for the 42 male sticklebacks
that completed nests in our
study. Hatched bars show
males that initiated nest building within 3 days following
transfer to nesting tanks (early
nesters) and open bars show
those taking longer than 3 days
(late nesters). The inset figure
shows the mean ( SE) nest
neatness indices of early nesters (hatched bars) and late
nesters (open bars). Early nesters built significantly neater
nests than late nesting fish (see
text for statistical details).
rA
sq-rt asin-transformed In
sq-rt asin-transformed Ic
Weight of deposited
substratum
r .312, p .047
r .233, p .159
r .199, p .206
r .211, p .186
r .006, p .972
r .020, p .901
explain both the pre-building lag, as such fish prepare physiologically for nesting, and alsoif nest quality is an honest
correlate of male qualitythe poor quality nests. Interestingly,
Jamieson and Colgan (1992) found that late-nesting males
were more likely to become sneakers in a laboratory study.
Although we can provide no evidence to distinguish between
these two hypotheses for the correlation between building lag
and final nest quality, they could be tested experimentally.
In one of the only other studies linking nesting behavior to
male traits, male common gobies Pomatoschistus microps that
built nests with little sand cover were in poorer nutritional
condition and more likely to cannibalize their offspring than
those that covered their nests more extensively (Kvarnemo et
al., 1998). We did not detect a relationship between the
weight of deposited substratum and male body condition factor or HSI, two medium-term indices of nutritional condition,
but we did find a negative correlation with relative spleen
weight. Enlarged spleens are an indicator of immune stress
associated with disease and parasite infection in fish (Arnott
et al., 2000; Bruno et al., 1998; Byrne et al., 1998). The small
number of males infected with S. solidus parasitesimportant
pathogens in our study population (Tierney, 1991)also
tended to build less compact nests.
Sticklebacks base nest site choices on many factors including the presence or absence of cover (e.g., Cleveland, 1994;
Jenni, 1972; Kraak et al., 2000), water depth (Kraak et al.,
2000; Mori, 1994) and distance to shore (see reviews by Rowland, 1994; Whoriskey and Fitzgerald, 1994). Females are
known to prefer males that build nests close to vegetation,
and such nests are more likely to last until fry-hatching (Kraak
et al., 1999b). Yet although the selection of nest site influences
mate attractiveness and the survival of embryos and fry (e.g.,
Kynard, 1978; Moodie, 1972; Sargent and Gebler, 1980), research on avian taxa has suggested that variation in the structure of the nest itself may also provide differential resistance
to adverse environmental conditions or predation attempts
(e.g., Mller, 1990). Stickleback nests are susceptible to the
action of local water currents and disturbances from predators, and both cause losses in natural habitats (Mori, 1995).
It would be interesting to know whether certain nest characteristics (e.g., compactness, neatness, or weight of deposited
substratum) influence resistance to perturbation events or
predation attempts, or female choice. Variation in nest con-
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