Summary 1.Communities of different species are often structured according to niche differentiation associated with competitive interactions. We show that similar principles may apply on an ecological time-scale when individuals of a species having a wide size variation compete for resources, using the Lake Tanganyika cichlid Lobochilotes labiatus (5-30 cm). This species has a mouth especially adapted to suck up invertebrates from rock crevices. 2. Individuals defended feeding territories against similar-sized conspecifics, but not against differentsized ones. Thus, territories of similar-sized fish rarely overlapped, but up to a total of seven individuals (of seven size-ranks) had broadly overlapping territories with dissimilar-sized individuals. Comparison with expectation from the null model demonstrated clearly that observed size ratios between adjacent size rank were determined non-randomly regardless of sexual combinations. 3. Larger individuals took larger prey types of larger average size, but more importantly used wider rock crevices from which to suck food than smaller individuals. We calculated pairwise values of Schoener's index of diet overlap C d and the values of Levin's index of diet breadth B d (prey type and prey size) and the same for the width of the rock crevices used for foraging ( C r and B r ). C d remained high among all combinations of the seven ranks. In contrast, C r declined strongly in combinations of adjacent ranks (to 0·27), and was low or zero among further different size ranks. This shows that fish with overlapping territories divided the food resources largely through foraging site partitioning. Accordingly, B d did not depend on the size difference to the nearest two coinhabiting fish, whereas B r did. 4. We conclude that this L. labiatus community is structured non-randomly: body size-dependent effects on foraging site usage result in competition with, and territorial exclusion of, similar-sized individuals, but not of dissimilar-sized individuals that were accepted as coinhabitants. Accordingly, mean body size ratios (large/small) between two adjacent ranks were consistently approximately 1·28 [standard deviation (SD) = 0·07, n = 104], while approximately 1·34 from the null model (SD = 0·34, n = 10 400 simulations). We discuss our results as an example of Hutchinson's rule, applied originally to size ratios of different species.
Diet studies have shown Humboldt squid Dosidicus gigas to be aggressive opportunistic predators, yet this approach has provided only a limited and potentially biased view of their trophic feeding behavior. As an alternative, we measured the δ 13 C and δ 15 N of D. gigas and their prey from the northern California Current ecosystem (NCC) and applied stable isotope Bayesian mixing models (Stable Isotope Analysis in R [SIAR]) to assess if D. gigas isotopically matched NCC or southern California Current (SCC) migratory end-members and to examine the proportional trophic contributions of prey groups from the NCC to their diet. For the trophic SIAR model, cluster analysis of prey taxa by their respective δ 13 C and δ 15 N values was first applied to consolidate prey into groups, which were then incorporated into the model as source groups to the diet mixture. Model results from examination of NCC and SCC migratory end-members indicated greatest contributions from the NCC system, indicating D. gigas was more integrated with the regional NCC isotopic signature. From the trophic SIAR model, the results indicated mixed but lower trophic-level feeding by D. gigas relative to previous diet-based studies, with greatest contributions from macrozooplankton, ichthyoplankton, and nekton such as juvenile rockfish, market squid, sand lance, and juvenile Pacific hake. Sensitivity analyses of the SIAR model based on varying isotopic fractionation factors of δ 13 C and δ 15 N showed that proportional contributions of prey to squid diets were resilient to change.
Theories suggest that, in cooperatively breeding species, female control over paternity and reproductive output may affect male reproductive skew and group stability. Female paternity control may come about through cryptic female choice or female reproductive behaviour, but experimental studies are scarce. Here, we show a new form of female paternity control in a cooperatively polyandrous cichlid fish (Julidochromis transcriptus), in which females prefer wedge-shaped nesting sites. Wedge-shaped sites allowed females to manipulate the siring success of the group member males by spawning the clutch at the spot where the large males were just able to enter and fertilize the outer part of the clutch. Small males fertilized the inner part of the clutch, protected from the large aggressive males, leading to low male reproductive skew. Small males provided more brood care than large males. Multiple paternity induced both males to provide brood care and reduced female brood care accordingly. This is, to our knowledge, the first documented case in a species with external fertilization showing female mating behaviour leading to multiple male paternity and increased male brood care as a result.
The plasticity of the sex roles in the blenniid fish Petroscirtes breviceps, a nest brooder with exclusive paternal care, was studied throughout an 8 month breeding season. Males performed most courtships early and late in the breeding season, whereas females performed most in the middle of the season. These results indicated that the sex of individuals initiating courtship changed seasonally, with courtship role reversal in the middle of the season. Intrasexual aggression in both sexes occurred much more frequently in mid-season than in the early and late seasons. Males frequently fought when available nest sites were limited, regardless of the presence of females, suggesting that males competed for nests in order to qualify to mate (resource competition). In contrast, courting females fought only in mid-season, when females' relative success in entering nests decreased, indicating that females competed for limited mating opportunities (mating competition). The reversed courtship roles and female mating competition in mid-season suggested that the sex roles in P. breviceps changed seasonally from the conventional roles to reversed roles and back again during one breeding season. This study provides the first empirical evidence of multiple changes in the sex roles of animals within a breeding season.
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