Social inhibition of maturation in females of the temperate wrasse Pseudolabrus celidotus and a comparison with the blennioid Tripterygion varium

In two temperate reef fish, Pseudolabrus celidotus (Bloch and Schneider) and Tripterygion varium (Bloch and Schneider) studied near Leigh, New Zealand, most spawning activity was concentrated during the first 2 months of spawning seasons which lasted about 5 months. In P. celidotus, maturation and spawning of first-year females (0+) was delayed with respect to older (1+yr) females. Within the 0+ age class, the minimum size of maturation declined as the spawning season proceeded. However, in all females of T. varium, ripening and spawning took place over the same time ranges. Observations on the seasonal patterns of female aggression suggested that larger female P. celidotus were socially inhibiting the maturation of small females. The level of interaction was high compared to that in T. varium, and reached a peak over the onset of the spawning season. This hypothesis was tested by the removal of larger (1+yr) female P. celidotus from a field population. The remaining 0+ females increased in ovary weight and matured earlier than those in an undisturbed area. We argue from this experiment that it is of advantage for P. celidotus females to spawn early in the season and of advantage to inhibit other females from doing so. Two possible reasons are suggested, the first applying to sexchanging species such as P. celidotus, the second to temperate-water fishes in general.     

On the possible contribution of mixed species flocks to species richness in neotropical avifaunas

Summary Mixed species foraging flocks are a dominant component of the infra-structure of avian communities in neotropical forests. In Amazonia, these flocks consist of pairs of 10–20 species, many of which are permanently associated with mixed flocks. At least half of these flocking species maintain territories that correspond exactly to the flock home range. Small individuals that participate as permanent members of the flocks must adopt the large home range of the larger nucleus species. Therefore, the densities of smaller species are dependent on the availability and density of flocks rather than the availability of food resources. Single pairs of 4 small flocking species with individual body masses of 8 g occupied exclusive territories of 8–12 ha. These were the same exact territories that were defended by at least 6 other flocking species with individual body masses of up to 37 g. Because of their attachment to flocks with large territories, small species are expected to under-utilize available food resources. The under-utilization of food resources is expected to allow smaller species to coexist with greater niche overlap resulting in increased species richness. This hypothesis was tested by quantifying foraging niche in terms of foraging height, foraging maneuver, and prey substrate; and using these values in addition to body mass and bill size (length, depth and width) to determine relative niche overlap between large versus small species pairs.Smaller species had greater foraging overlap than large flocking species and particularly the three smallest species of the genus Myrmotherula; longipennis, axillaris and menetriesii had very high overlap (average foraging niche overlap for the 3 species=0.83±0.12 compared with 0.12±0.19 for all flocking species), similar body sizes (body masses differing by no more then 8%) and similar bill morphologies (maximum ratio in length=1.08, width=1.07, and depth=1.06). These results are consistent with the hypothesis that small species participating in Amazonian mixed flocks can coexist with greater niche overlap because their density is flock dependent rather than resource dependent.     

Differential use of shelter holes by sympatric species of blennies (Blennidae)

Comparative use of shelter use by three sympatric species of combtooth blenny (Ecsenius stictus, Glyptoparus delicatulus, and Salarias patzneri) was studied among micro-atolls in the lagoon at Lizard Island (14°42′S, 145°30′E), northern Great Barrier Reef, Australia. Blenny species used different sized holes; however, the average diameter and depth of holes used by the smallest and largest species differed by only 4 and 25 mm, respectively, indicating interspecific differences in suitable refuge can be very subtle. Both hole diameter and depth were positively related to total length of fish, suggesting use of holes relates to interspecific differences in body size. Total abundance of blennies was best explained by a general linear model that included either the number of holes or total habitat area on individual micro-atolls, predictor variables that were positively correlated with each other. However, the relative importance of variables differed among the three species, feeding area being most important for S. patzneri, feeding area and number of holes for E. stictus, and variance in hole diameter being the best explanatory variable for G. delicatulus abundance. The number of blenny species on a micro-atoll was best explained by variance in hole diameter, emphasizing the influence of refuge size variety in fish diversity. It is likely that subtle habitat partitioning, which relates to interspecific differences in body size, contributes to the co-existence of blenny species within the same microhabitat, but presence of holes is unlikely to regulate abundance of these fish.     

Dominance in feeding territories relates to foraging success and offspring growth in brown skuas <Emphasis Type="Italic">Catharacta antarctica lonnbergi</Emphasis>

Territoriality should lead to strict dominance, as territory holders typically control access to resources and exclude others from their use. In feeding territories, dominance should be reflected in foraging success and ultimately in reproduction differences; however, these successive links have rarely been made explicit. Therefore, we investigated a population of brown skuas Catharacta antarctica lonnbergi, in which only part of the breeding population occupied feeding territories within penguin colonies. We identified the dominance hierarchy and determined the foraging success of the participants in fights for access to penguin carcasses within the territories. Furthermore, we monitored offspring growth from parents with and without feeding territories. Our results indicated a clear dominance hierarchy with territorial birds in their own territory dominating over territorial breeders from other territories, non-territorial breeders and non-breeding birds. However, territory owners could not completely exclude others from access to food. Foraging success was positively related to dominance scores: The dominant territory owners received 63% of a carcass, whereas non-territorial pairs could get less than 10%. The link between foraging success and offspring development was less clear: Although male chicks of non-territorial parents suffered from lower growth rates and, thus, delayed fledging, there were no such differences in female chicks. Territoriality in skuas did not imply a complete occupation of food, but guaranteed optimal growth conditions for offspring. Non-territorial individuals were forced to search for alternative resources, and the restricted access to the preferred food resulted in inferior conditions for offspring development, making this foraging strategy less rewarding.     

Diel foraging patterns of the sea urchin Centrostephanus coronatus as a predator avoidance strategy

During the day, the diadematid sea urchin Centrostephanus coronatus occupies holes and crevices in shallow subtidal rocky substrata. Individuals emerge from these after sunset and forage on organisms attached to the surrounding rock surface. Each urchin travels <1 m from its shelter and returns to the same one before sunrise. The sheephead wrasse Pimelometopon pulchrum does not remove urchins from their shelters, but will attack and consume urchins placed in normal feeding locations during the daytime. The active periods of the sheephead and the urchin do not overlap; urchins begin foraging about 20 min after the diurnal sheephead retire in the evening and return to their shelters 1 to 2 h before sheephead resume feeding in the morning. We infer that the urchin's daytime crevice-dwelling and nocturnal foraging habits have evolved as a response to sheephead predation. Moreover, because shelters are limited in supply, shelter fidelity may have evolved to insure refuge from sheephead.     

Interactions between fangblennies (Plagiotremus rhinorhynchus) and their potential victims: fooling the model rather than the client?

Lepidophagous (scale-eating) blue-striped fangblennies (Plagiotremus rhinorhynchus Bleeker 1852) are often found sympatrically with the bluestreak cleaner wrasse (Labroides dimidiatus Valenciennes 1839). They have some resemblance to the juvenile L. dimidiatus and have previously been reported as aggressive cleaner wrasse mimics. We observed 14 P. rhinorhynchus on a small area in the barrier reef near Hoga Island, Indonesia to assess the effects of client size on the behaviour and attack success of fangblennies: our results suggest that fangblennies are selective with regard to victim size; fish avoided by the fangblennies are significantly larger than those not avoided and attack success is significantly higher at intermediate victim size classes. The behaviour of the victims also has a significant direct effect on the foraging success of the fangblennies; where the potential victim posed, 63.6% were ignored by the fangblenny and only 7.4% of attacks were successful on posing fish as opposed to a surprise attack success rate of 71.6%. Overall, victims which exhibited the pose behaviour were significantly smaller in size. It appears likely that the predatory strategy of these fangblennies varies with victim size and that mimicry plays a minor role in attracting potential victims. We suggest that in common with other mimetic fish the resemblance of fangblennies to juvenile bluestreak cleaner wrasse allows them to actively hunt in areas where adult cleaners are common thus, indirectly improving their feeding opportunities.     

Coordinated mechanics of feeding,swimming, and eye movements in <Emphasis Type="Italic">Tautoga onitis</Emphasis>, and implications for the evolution of trophic strategies in fishes

Fish feeding behavior results from successful coordination of the fins, jaws, and sensory systems, and the organization of this behavior may affect the fish’s foraging abilities and trophic ecology. Using quantitative kinematic methods, movements of the jaws, fins and eyes of Tautoga onitis (Teleostei: Labridae) were analyzed during feeding events. Tautog feeding events consisted of three phases: approach, strike, and recovery, each defined by a combination of kinematic events. The approach was characterized by changes in fin movements and in body position, with the eyes directed forward at the food item. The strike began with the onset of jaw opening and protrusion, then cranial elevation, with the eyes no longer looking at the food item. The end of the strike and the beginning of the recovery involved a braking maneuver with the pectoral fins; the fish turns and swims away from the original food location item after prey capture. The coordination performance variables of tautog were quantitatively compared to published data from closely related cheiline wrasses and parrotfishes, which represent different feeding ecologies within a monophyletic assemblage. Fishes feeding on molluscs and benthic invertebrates (Cheilinus fasciatus and Tautoga onitis) represented an intermediate coordination condition, with herbivores (the parrotfishes, Scarus quoyi and Sparisoma radians) at one extreme, and fishes feeding on elusive prey (Epibulus insidiator and Oxycheilinus digrammus) at the other extreme. The analysis suggests that the biomechanical demands of coordination for feeding on benthic invertebrates may represent a generalized, and perhaps ancestral behavior in the wrasses, whereas more specialized trophic niches have evolved divergent, more specialized demands. Examining the movement and coordination of the jaws, fins, and eyes during fish feeding provides a detailed mechanistic basis for behavior, and comparison of coordination patterns during feeding among different taxa can measure how these trophic strategies differ. Understanding the evolution of feeding ecologies in these demersal fishes may have implications for understanding their role within their shallow water reef community.     

Influence of territoriality on adult density in two rockfishes of the genus Sebastes

Nonterritorial Sebastes carnatus and S. chrysomelas existed, along with territorial individuals, at 3 tagging sites off southern California, USA, which were monitored for nearly 1 yr. To test the hypothesis that territoriality affected adult density in these species, territorial fish were removed and the subsequent utilization of vacated territories by other fish was monitored. Intrusion of other fish into vacated territories increased significantly in 90% of the removals. Other fish colonized both the feeding and sheltering parts of the vacated territories, indicating that the previous owners had successfully defended both parts of their territories. Many of the colonizers had already possessed territories; they expanded their territories or moved into presumably better havitat. Several previously-nonterritorial fish also moved into vacated areas, and at least some of them appeared to establish territories. These fish, then, had previously been capable of establishing territories, but were prevented from doing so by resident territory holders. Thus territoriality, rather than such other factors as predation or low recruitment, limited the number of territorial fish at each site. However, territorial fish did not inhibit the settlement of larval recruits, and the relative mortality rates of older territorial vs nonterritorial fish were not determined. Thus the question of whether territoriality was a major factor controlling total density remains unresolved.     

Interference asymmetries among age-sex classes of rufous hummingbirds during migratory stopovers

Summary Three age-sex classes of rufous hummingbirds (Selasphorus rufus) overlap temporally and defend feeding territories during migratory stopovers in the Sierra Nevada of California. We demonstrate that these classes differ in their ability to secure and maintain high-quality feeding territories for refueling, and that these differences result in differences in resource use. Data on acquisition of territories, territory characteristics, and responses of territory owners to intruders suggest that several mechanisms are involved in determining dominance, involving sex- and age-related differences in wing disc loading, coloration, and experience. We discuss the implications of these results for understanding intraspecific variation in migration strategies. Correspondence to: F.L. Carpenter     

Rules of engagement for adult salamanders in territorial conflicts with heterospecific juveniles

Coexisting species within a guild have the potential for resource overlap and consequently for interspecific competition (e.g., interspecific territoriality). When the adults are of different sizes, which frequently is the case in terrestrial salamander communities in eastern North America, competition may occur between juveniles of the larger species and adults of the smaller species. Adults of the relatively small redbacked salamander (Plethodon einereus: up to 13 cm total length) defend intra- and intersexual territories on the forest floor, and they are broadly sympatric with the larger P. glutinosus (up to 21 cm total length). Although individuals of the two species share the same forest floor habitat, we found significantly fewer juveniles of P. glutinosus sharing territories with 336 same-size adults of P. cinereus than would be expected by random chance alone. In laboratory experiments, residential adults of P. cinereus were as aggressive toward juvenile intruders of P. glutinosus as they were toward adult conspecific intruders. Therefore, adults of P. cinereus appear to defend territories against juveniles of P. glutinosus, illustrating how interference competition may depend on the symmetry of sizes between the species.     

Males are faster foragers than females: intersexual differences of foraging behaviour in the Apennine chamois

Availability of food resources and individual characteristics can influence foraging behaviour, which can differ between males and females, leading to different patterns of food/habitat selection. In dimorphic species, females are usually more selective in food choice, show greater bite rates and spend more time foraging than males. We evaluated sexual differences in foraging behaviour in Apennine chamois Rupicapra pyrenaica ornata, during the warm season, before the rut. Both sexes selected nutritious vegetation patches and spent a comparable amount of time feeding. However, males had a significantly greater feeding intensity (bite rate) and a lower search effort for feeding (step rate), as well as they spent more time lying down than females. Females selected foraging sites closer to refuge areas than males. In chamois, sexual size dimorphism is seasonal, being negligible in winter–spring, but increasing to 30–40 % in autumn. Our results suggest that males enhance their energy and mass gain by increasing their food intake rate during the warm season, to face the costs of the mating season (November). Conversely, females seem to prioritize a fine-scale selection of vegetation and the protection of offspring. A great food intake rate of males in the warm season could have developed as a behavioural adaptation leading herbivores to the evolutionary transition from year-round monomorphism to permanent dimorphism, through seasonal dimorphism.     

Lek structure and territoriality in the chryxus arctic butterfly,Oeneis chryxus (Satyridae)

Summary The territorial and mating systems of the satyrid butterfly Oeneis chryxus were studied for 3 years on individually marked populations in algonquin Provincial Park, Ontario. Males defended territories in a dry open forest clearing by means of chases and spiral flights with conspecifies. Defended areas did not contain food or oviposition sites required by females. Territories were not uniformly distributed over the study area; the highest density of territories was clustered over bare ground covered with sand and small stones rather than areas of vegetation. This pattern of distribution of territories occurred during each year of the study. The patterns of occupancy of territories showed a high degree of consistency from one day to the next, some males defending a given territory for as many as 11 consecutive days. Removal experiments showed that a surplus of non-territorial males did not exist. Vacant territories in the high-density area were absorbed by neighboring males; those in the low-density area remained empty. Male size was not correlated with territory position in the study are. Wing patterning (i.e. size of spots) differed between males occupying territories on sandy/stony versus vegetated areas. The territorial and mating systems of O. chryxus resemble lek polygyny, and are more similar to some vertebrate lek systems than to hilltopping or landmark territoriality.     

Matching and mismatching stable isotope (δ13C and δ15N) ratios in fin and muscle tissue among fish species: a critical review

Using non-lethal tissue sampling for stable isotope analysis has become standard in many fields, but not for fishes, despite being desirable when species are rare or protected, when repeated sampling of individuals is required or where removal may bias other analyses. Here, we examine the utility of fish dorsal fin membrane as an alternative to muscle for analyzing δ¹³C and δ¹⁵N ratios in two reef fish species (blue cod Parapercis colias and spotty Notolabrus celidotus) that have differing feeding modes. Both species exhibited evidence of size-based feeding from fin δ¹⁵N values, but not from muscle. Blue cod fin δ¹⁵N increased steadily throughout the sampled size range (213–412 mm fork length), whereas spotty exhibited a distinct ontogenetic diet shift at approximately 120–140 mm fork length after which size-based feeding did not occur. Fin membrane was higher than muscle in δ¹³C in both species and in δ¹⁵N for blue cod, but fin δ¹⁵N was lower than muscle in spotty. The δ¹³C and δ¹⁵N fin–muscle offsets were constant in spotty regardless of size, while in blue cod, δ¹³C was constant with fish size, but δ¹⁵N offsets increased with increasing fish size. Non-lethal sampling utilizing fin tissue can be employed to estimate stable isotope values of muscle in fishes, but it is necessary to assess relationships among tissues and the effects of fish size on isotope values a priori for each species studied. Our data indicated that fin membrane may be a more sensitive tissue than muscle for detecting size-based feeding in some fish species using stable isotopes. A critical literature review revealed inconsistencies in tissue types tested, little understanding of tissue-specific trophic shift or turnover rates, and pseudo-replicated analyses leading to erroneous postulating of 1:1 relationships between tissues.     

Seabird predation on the sea-skater Halobates sericeus (Heteroptera: Gerridae)

Regurgitated food samples were collected from 18 species of seabirds on 8 of the Northwestern Hawaiian Islands between February 1978 and February 1981. Sea-skaters (Halobates sericeus) was found in the diets of 9 species, but can be considered to be an important food item for only 4 species: the blue-gray noddy Procelsterna cerulea; the Bonin petrel Pterodroma hypoleuca; the gray-backed tern Sterna lunata; and Bulwer's petrel, Bulweria bulwerii. The blue-gray noddy, by far the most important avian predator of Halobates spp., may at times feed exclusively on this food item and may appreciably reduce the populations of sea-skaters within their foraging territories.     

Density and distribution patterns of the temperate marine fish Cheilodactylus spectabilis (Cheilodactylidae) in a reef environment

Cheilodactylus spectabilis (Hutton) is common over shallow reefs in north-eastern New Zealand. Replicated transect between-area differences in density and in size frequency. C. spectabilis is not nocturnally active. The between-area differences in density and size frequency remained constant over daylight hours, demonstrating that C. spectabilis does not undergo any systematic feeding migrations. Topographic complexity was shown to have a significant positive influence on fish density. Mean size was related to water depth at 5 localities investigated, with small (<200 mm standard length, SL) individuals being restricted to shallow water. All sizes of C. spectabilis examined had fed on small invertebrates, with gammarid amphipods predominating. No evidence of size-related differentiation in feeding patterns was observed, although small individuals spent significantly more time feeding than large ones. Small (<200 mm SL) individuals occupied feeding areas and shelter sites from which they excluded other small C. spectabilis. Large individuals showed no site-associated aggression, covered a greater area during daily movements and overlapped with other large and small C. spectabilis while feeding. No evidence of dawn or dusk peaks of feeding activity or movement was collected.     

The development of sex differences in ring-tailed lemur feeding ecology

Sex differences in feeding ecology may develop in response to fluctuations in physiological costs to females over their reproductive cycles, or to sexual size dimorphism, or function to minimize feeding competition within a group via resource partitioning. For most mammal species, it is unknown how these factors contribute to sex differences in feeding, or how the development of males and females reflects these intraspecific feeding differences. We show changes in dietary composition, diversity, overlap, and foraging behavior throughout development in ring-tailed lemurs (Lemur catta) and test how the development of sex differences in feeding is related to female costs of reproduction and year-round resource partitioning. Sex differences in dietary composition were only present when females were lactating, but sex differences in other aspects of feeding, including dietary diversity, and relative time spent feeding and foraging, developed at or near the time of weaning. Sex difference in juveniles and subadults, when present, were similar to the differences found in adults. The low year-round dietary overlap and early differences in dietary diversity indicate that some resource partitioning may begin with young individuals and fluctuate throughout development. The major differences between males and females in dietary composition suggest that these larger changes in diet are closely tied to female reproductive state when females must shift their diet to meet energetic and nutritional requirements.     

Foraging activity and resource use by three parrotfish species at Fernando de Noronha Archipelago, tropical West Atlantic

Parrotfishes are important components of the herbivore and detritivore guilds of tropical and subtropical reefs. Most of parrotfish species are protogynous hermaphrodites that change colour and sex, from initial phase females or males (IP) to terminal phase males (TP). We studied the foraging behaviour of Sparisoma amplum, S. axillare and S. frondosum, three syntopic scarids on the rocky reefs of Fernando de Noronha Archipelago, Tropical West Atlantic. The three parrotfish species differed in food selection and preference, but IP and TP individuals of the same species preferred the same food types, except for S. amplum. Feeding rates of IP individuals were higher than those of TP individuals, but the distribution of feeding frequencies throughout the day of IP and TP individuals of the same species was similar. IP individuals had higher feeding rates than TP ones, which seems related to the fact that TP individuals spend a large amount of time patrolling their territories and chasing away conspecific individuals at the study site. The general foraging pattern we found for S. amplum, S. axillare and S. frondosum is similar to patterns found for other parrotfish species in the Western Atlantic.     

Social spacing in the temperate marine goby Coryphopterus nicholsi

Coryphopterus nicholsi is a temperate marine goby that occupies rock rubble in protected subtidal areas along the Pacific coast of NOrth America. In field populations, fish of all sizes and both sexes are found in the same habitat. All observed fish over 25 mm standard length defended space during both the reproductive and the non-reproductive seasons, with territory size being directly correlated with fish size. There was peripheral overlap of territories, particularly between adjacent adult conspecifics of different size classes, but also with smaller fish that had territories centered in the interstices of larger territories. However, zones of overlap were used at different times by space-sharing fish so that defended areas remained temporally discrete. Access to areas of overlap appeared to be determined by dominance rank. More dominant (usually larger) fish used areas of overlap at will; the defense and use of shared areas by less dominant fish was contingent upon the absence of larger fish. Social organization in C. nicholsi appears to be defined by a combination of territorial behavior and dominance relationships among adjacent individuals, permitting the coexistence of all members of the population under circumstances of limited habitat availability.