Overlapping territory of a benthophagous fish, Goniistius zonatus (Teleostei: Cheilodactylidae)

Overlapping territories of a benthophagous morwong, Goniistius zonatus, were studied at two temperate water locations (Morode and Arakashi) in southern Japan. At both locations, total length (TL) of the fish inhabiting the study areas ranged from 12 cm to 34 cm, but fish 20–25 cm TL were not found in the Arakashi study area. At Morode, individuals maintained territories against conspecifics of similar size, and the territories broadly overlapped among three size classes (small < 20 cm medium <25 cm large). At Arakashi, large-sized fish chased individuals of both the same- and medium-sized individuals from their territories, resulting in territory overlap only between large and small size classes. The territories were apparently maintained for feeding, as the main activity of the fish within the territories was foraging and no reproductive behavior was observed at either location. All fish foraged on similar substrates in the daytime. At Morode, however, the diet composition differed among different size classes, whereas at Arakashi, fish of all sizes fed on quite similar foods. A comparison between the two populations indicates that the territory overlap among the three size classes at Morode may have resulted from food-resource partitioning.     

Group foraging by the herbivorous cichlid fish,Petrochromis fasciolatus,in Lake Tanganyika

Group foraging of the algae eating cichlid,Petrochromis fasciolatus, was studied along a rocky shore in Lake Tanganyika, in an area occupied by the feeding territories of several herbivorous cichlid species. SolitaryP. fasciolatus or schools comprising a few members only were nearly always repelled from such territories, although larger schools (40–250 individuals) ofP. fasciolatus were able to overwhelm the defenses of territory owners and enable subsequent foraging. The frequency and duration of individual foraging sessions on the algal mats were positively correlated with group size, probably owing to individuals in large schools suffering fewer attacks from territory owners. Almost all of the foraging sites ofP. fasciolatus were inside the territories of other herbivorous cichlids, especially those ofNeolamprologus moorii (more than 80% of total), in which the algal density was more than 10 times that in territories of the other species.N. moorii territories included only 4% rocky substrate, indicating that schools ofP. fasciolatus selectively chose territories containing rich resources.     

Size-specific social interactions and foraging styles in a shallow water population of mutton snapper,Lutjanus analis (Pisces: Lutjanidae), in the central Bahamas

Synopsis Field observations quantified the effects of fish size and time of day on activity patterns, intraspecific encounters, and foraging styles in mutton snapper,Lutjanus analis, during the spring and winter of 1991. Fish ranged in size from 15 to 65 cm fork length (FL), and were associated with an artificial patch reef system located on a shallow seagrass meadow in the Exuma Cays, Bahamas. The most common, non-resting diurnal activities were intraspecific chasing and displacing, and feeding. Intraspecific displacing was significantly higher during midday compared to morning or evening. The highest proportion of intraspecific encounters (combined chasing and displacing events) occurred among medium (25–35 cm FL) and large (> 35 cm FL) fish. The few large fish observed (13% of population) initiated the same proportion of encounters as the predominant (50%) medium fish. The remaining (37%) small fish (> 25 cm FL) were the least aggressive. Dark barred and dark nape color patterns were associated with displacing and chasing, respectively. Fish exhibited considerable variability in feeding behavior, Proportionally fewer fish fed during midday compared to morning or evening, although small fish fed proportionally more often than medium or large fish despite time of day. Picking was the primary feeding mode and was observed during all times of day. Winnowing was observed during midday and evening, whereas midwater strikes were confined to morning and evening. Small fish displayed proportionally higher picking and midwater strikes during morning and evening, respectively, compared to medium or large fish. However, large fish winnowed proportionally more often than small or medium fish during evening. Dark barred color patterns were associated with feeding on the substrate, whereas no color changes occurred during midwater strikes. Our results indicate thatL. analis forms dominance hierarchies and that high variability in foraging styles, according to fish size and time of day, may be a means to reduce intraspecific competition.     

Niche differentiation depends on body size in a cichlid fish: a model system of a community structured according to size regularities

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 different-sized 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.     

Interspecific territoriality in males of the tube blenny <Emphasis Type="Italic">Neoclinus bryope</Emphasis> (Actinopterygii: Chaenopsidae)

Territoriality was investigated in the tube blenny Neoclinus bryope (Actinopterygii: Chaenopsidae) at rocky intertidal areas of Banda Beach, Tateyama Bay, central Japan. Males used small holes as spawning nests, usually staying at the nest and maintaining the area while showing exclusive behaviors. Their home ranges were limited to areas within 30 cm distance from the nest for over 2 months. Four heterospecific fishes were threatened when they approached to within 6–14 cm of the nest holes, and two species of carnivorous snails were removed at points 0–30 cm from the nest entrance. There were no significant differences between the distances from the nest entrances to the points defended against fish and those used for foraging. As the four heterospecific species have similar feeding habits to those of N. bryope, the area defended against fishes may function as a foraging territory. At 24 h following the removal of nest owners, carnivorous snails had gathered to actively prey on eggs, indicating that the area defended against snails may function as a territory for protection against egg predators. Although the positions of females where males started courtship displays were significantly farther than the foraging points and the positions of threatened fishes, males displayed no territoriality against conspecific males. The fact that males did not leave the nest hole during the courtship suggests that it may be costly to maintain a courtship territory. These results show that males of tube blenny maintain territories for egg guarding and for protecting food resources around their nest holes in the spawning season.     

Nocturnal foraging habitats of French and bluestriped grunts,Haemulon flavolineatum and H. sciurus,at Tobacco Caye,Belize

Synopsis Nocturnal foraging habitats of Haemulon flavolineatum and H. sciurus were investigated in the backreef habitat around Tobacco Caye, Belize. Grunts leave the reef at dusk to forage in the grass beds and sand flats surrounding the reef. The hypothesis that French and bluestriped grunts use separate foraging habitats was examined by following tagged fishes from their diurnal territories or schooling sites to nocturnal foraging grounds. The tag consisted of a small, glowing Cyalume light stick sutured to the dorsal musculature of the fish, next to the first dorsal fin. Surveys of foraging habitats were done to support the tracking study. Large quadrats (225 m²) were set out over the sand flats and grass beds during the day. The numbers of French and bluestriped grunts feeding in each habitat were counted one hour after dark. Foraging French grunts used sand flats, whereas bluestriped grunts usually fed in grass beds. Repeated sightings of two French grunts and one bluestriped grunt in the same individual night-time locations support the hypothesis that nocturnal foraging sites may be used repeatedly by the same individuals.     

Intertidal territoriality and time-budget of the surgeonfish, Acanthurus lineatus, in American Samoa

Synopsis The herbivorous surgeonfish Acanthurus lineatus aggressively maintained feeding territories in the surf zone of the outer reef flat in American Samoa. Intertidal territories were re-established each morning, as well as after displacement by low tides or rough surf. Day-to-day site fidelity of recognizable individuals was high: 99.9% return rate per day for adults (15–20 cm), 99.6% for juveniles (8–13 cm), and 97.2% for recruits (2.5–5 cm). Fish fed on turf algae primarily in the afternoon (80% of available time), and spent 10% of their time on active territorial defense and 2–13% of their time on forays from their territory. On average, a fish defended its territory 1900 times daily and took 17 000 bites (= 7400 bites m^–2 d^–1), but rough surf reduced feeding by 60% and defense by 75%. High territorial defense requirements significantly reduced feeding rates. Although the distribution and behavior of this species in Samoa was in large part similar to that reported for it elsewhere (Australia, Indian Ocean), there were notable differences: in Samoa A. lineatus densities within colonies were greater (0.4 fish m^–2), territory size was smaller (2.3 m²), and defense rate against intruders was greater (2.5 attacks min^–1). These differences in Samoa may be related to their smaller body size, greater abundance or increased food supply caused by hurricane damage to reefs which has enhanced the algal turfs that A. lineatus feeds upon.     

Resource partitioning in two carcinophagous confamiliars: distribution and feeding habits of psychrolutid fish on the southern Patagonian shelf

Psychrolutes marmoratus and Cottunculus granulosus, two psychrolutids, inhabit the shelf and continental slope of the southern South America between 45 and 1,250 m. Over this range, the P. marmoratus is found in depths of less than 400 m, whilst C. granulosus inhabits waters of >200 m. There is a trend of increasing total length with increasing water depth for P. marmoratus and the converse for C. granulosus. The total lengths of the species are different in the depth zone that they overlap in (200–400 m), and we hypothesise that this may reduce interspecific competition. Both are specialised carcinophagous feeding on crabs, large isopods and sea spiders, but they also prey upon polychaets and gastropods. There was no difference in the feeding spectra of similar-sized fish sampled at the same depth, indicating a similar foraging strategy. In contrast to this, the feeding spectra are quite different between the species over different depths as well as at different adult sizes. The feeding niche breadth in these species is similar to other slope dwelling fish in the area.     

Male Reproductive Behavior of the Social Wasp Polistes fascatus (Hymenoptera: Vespidae)

Populations of male Polistes fuscatus simultaneously exhibit two different mate-locating tactics. Some males defend territories in female nesting and hibernation sites. These males frequently do not occupy the same territory each day, and they drag their gasters over perches, which may function to apply a secretion to the perch. Another segment of the population patrols large overlapping areas in female foraging sites. In contrast to territorial males, patrolling males do not rub their gasters on perches, and males seen on more than one day are tenacious to one area. Males in both sites are aggressive to other males and attempt to copulate with females. A laboratory study indicates that large males have an advantage in male-male competition. The mean size of patrollers is smaller than that of territorial males, indicating that patrollers are competitively inferior males. Yet there is considerable size overlap of males between the two sites, suggesting that there is also overlap in the range of probability of mating success between the two sites.     

Feeding habits and diet overlap of marine fish larvae from the peri-Antarctic Magellan region

The Magellan region is a unique peri-Antarctic ecosystem due to its geographical position. However, the knowledge about the distribution and feeding ecology of fish larvae is scarce. Since this area is characterized by low phytoplankton biomass, we hypothesize that marine fish larvae display different foraging tactics in order to reduce diet overlap. During austral spring 2009–2010, two oceanographic cruises were carried out along southern Patagonia (50–56°S). Larval fish distribution and feeding of the two most widely distributed species were studied, the smelt Bathylagichthys parini (Bathylagidae) and black southern cod Patagonotothen tessellata (Nototheniidae). Larvae of B. parini showed a lower increase in the mouth gape at size, primarily feeding during daytime (higher feeding incidence during the day) mostly on nonmotile prey (invertebrate and copepod eggs, appendicularian fecal pellets, diatoms). They showed no increase in feeding success (number, total volume of prey per gut and prey width) with increasing larval size, and the niche breadth was independent of larval size. Larvae of P. tessellata showed a large mouth gape at size, which may partially explain the predation on motile prey like large calanoid copepods (C. simillimus) and copepodites. They are nocturnal feeders (higher feeding incidence during night) and are exclusively carnivorous, feeding on larger prey as the larvae grow. Nonetheless, niche breadth was independent of larval size. Diet overlap was important only in individuals with smaller mouth gape (<890 μm) and diminished as larvae (and correspondingly their jaw) grow. In conclusion, in the peri-Antarctic Magellan region, fish larvae of two species display different foraging tactics, reducing their trophic overlap throughout their development.     

Overlapping territories ofPseudosimochromis curvifrons males and other herbivorous cichlid fishes in Lake Tanganyika

Males of the herbivorous cichlid fishPseudosimochromis curvifrons established mating territories 3–10 m in diameter, which included both spawning and feeding sites. Territorial males attacked conspecific males and also other species. Only conspecific males were chased out of the territories. Territorial males attacked other species at the spawning sites while courting or waiting for females and at other sites in their territories while patrolling and foraging. Attacks against herbivorous species were more frequent than against non-herbivores since herbivores were much more abundant. Territories of four abundant herbivorous cichlids largely overlapped the territories ofP. curvifrons males. Symbiotic relationships were not detected between the species. Instead, they were aggressive towards each other but coexisted by segregating feeding sites in the overlapping areas. The aggressive coexistence ofP. curvifrons males with other herbivores may have resulted from the energetic costs of defending their relatively large territories against all food competitors. Interspecific food-resource partitioning may also have facilitated the coexistence.     

Feeding habits of the blue shark, Prionace glauca, and salmon shark, Lamna ditropis, in the transition region of the Western North Pacific

We describe the feeding habits of 70 blue sharks (Prionace glauca) and 39 salmon sharks (Lamna ditropis) caught at 0–7 m depth at night by research drift gillnets in the transition region of the western North Pacific during April–May of 1999 and 2000. Blue sharks of 50–175 cm total length fed on a large variety of prey species, consisting of 24 species of cephalopods and 16 species of fishes. Salmon sharks of 69–157 cm total length fed on a few prey species, consisting of 10 species of cephalopods and one species of fish. Important prey for the blue sharks were large, non-active, gelatinous, meso- to bathypelagic cephalopods (e.g., Chiroteuthis calyx, Haliphron atlanticus, Histioteuthis dofleini and Belonella borealis) and small myctophid fishes. Important prey for the salmon sharks were mid-sized, active, muscular, epi- to mesopelagic squids (e.g. Gonatopsis borealis, Onychoteuthis borealijaponica and Berryteuthis anonychus). Our results suggest that blue sharks feed on cephalopods mainly during the daytime when they descend to deep water. Salmon sharks may feed opportunistically with no apparent diurnal feeding period. Blue sharks and salmon sharks have sympatric distribution in the transition region in spring; they have different feeding habits and strategies that reduce competition for food resources.     

Evidence that fish structure the zooplankton communities of turbid lakes and reservoirs

1. Visually foraging fish typically exclude large zooplankton from clear‐water lakes and reservoirs. Do fish have the same effect in turbid waters, or does turbidity provide a refuge from visual predation? 2. To test the hypothesis that fish exclude large zooplankton species from turbid sites, I searched for populations of medium or large Daphnia species in turbid, fish‐containing reservoirs of south‐central Oklahoma and north‐central Texas, U.S.A., and surveyed the literature for accounts of Daphnia species in turbid habitats worldwide. 3. Only small Daphnia species and the exuberantly spined Daphnia lumholtzi were detected in the turbid reservoirs. The Daphnia species in the reservoirs are smaller than other Daphnia species that occur in the area but were not detected. An extensive survey of the literature suggests that large Daphnia may be found in the lakes of extreme turbidity [Secchi disk depth (SD) < 0.2 m] but that only small and spiny Daphnia are likely to occur in more typical turbid locations (1.0 m > SD > 0.2 m) unless some additional factor reduces the influence of fish predation in such sites. 4. The field samples from Texas and Oklahoma together with the literature review suggest that the effect of visually foraging planktivorous fish on the size structure of turbid‐water zooplankton communities may often be as strong or even stronger than the effect of fish on clear‐water zooplankton communities.     

Nile perch,Lates niloticus,predation on the freshwater prawn,Caridina nilotica,in the Nyanza Gulf,Lake Victoria,East Africa

Synopsis The percentage of Nile perch, Lates niloticus, containing the prawn, Caridina nilotica, declined from 60–85%, for fish between 5 and 40 cm total length, to less than 10% for fish larger than 80 cm in length. The maximum number of prawns eaten by a Nile perch increased steadily, from 10 prawns for a fish in the 5–9.9 cm length group, to 913 prawns in a fish in the 60–69.9 cm length group, and then fell sharply for larger fish. The mean number of Caridina eaten by fish that were foraging on this animal followed a similar trend. Gill raker spacing increased in direct proportion to fish length and this may prevent fish larger than 70 cm in length from being effective prawn predators.     

Ontogenetic changes in foraging tactics of the piscivorous cornetfish <Emphasis Type="Italic">Fistularia commersonii</Emphasis>

Foraging behaviors of the piscivorous cornetfish Fistularia commersonii were observed at shallow reefs in Kuchierabu-jima Island, southern Japan. This fish foraged on two types of prey fishes: one was reef fish that typically dwell on or near substrata (e.g., Tripterygiidae and Labridae), and the other was pelagic fish that shoal in the water column (e.g., Clupeidae and Carangidae). The prey sizes, prey types and foraging behaviors changed as the predator size increased. Prey sizes were largely limited by gape size of the cornetfish, and small predators consumed small prey. The small cornetfish (10–30 cm in total length) fed only on reef fish captured after stalking (where the fish slowly approaches the prey and then suddenly attacks). The stalking was done either solitarily or in foraging association with conspecifics. Large fish (30–120 cm) fed on both types of fishes by stalking and/or chasing (where the fish chases the prey using its high mobility and attacks), either solitarily or in foraging association with con- or heterospecifics. Thus, chasing was only performed by the large cornetfish against pelagic prey fish in associative foraging with other con- and heterospecific predators. As their body sizes increased, F. commersonii began to show a diversification of foraging behaviors, which was strongly related not only to the habitat types and anti-predatory behaviors of the prey fishes but also to associative foraging with con- or heterospecifics, which improves their foraging success.     

Territorial Organization of the White Fronted Bee-eater in Kenya

White fronted bee-eaters (Merops bullockoides) live in extended family clans that aggregate to roost and nest in large colonies. Members of a given clan also share a common foraging territory, spatially segregated from the colony, to which they commute daily. The size of this foraging territory is positively related to clan size. Clan foraging territories are divided into a number of loosely overlapping foraging home ranges (FHRs), each occupied by an individual or mated pair of birds. Bee-eaters feed solitarily, flycatching to snap up large insects from widely dispersed perches. Each bird tolerates intrusion on its FHR by various members of its own clan, but aggressively excludes individuals belonging to other clans. Birds defend only their own FHR; however, because of the high amount of FHR overlap, the result is a loose form of group defense of the larger clan feeding area. For this reason we refer to the system as one of clan foraging territories. Birds occupying clan foraging territories located more than 1.5 to 2 km from a colony temporarily abandoned them while feeding nestlings. At such times, these birds provisioned their young by foraging near the colony. Birds that abandoned territories foraged less efficiently, provisioned nestlings at a lower rate, and had lower breeding success than did birds that continued use of their foraging territories. A model is developed relating territory abandonment to the energetics of central place foraging. Bee-eaters typically shift colony locations between successive breeding seasons. Foraging territory locations, in contrast, remain largely stable, resulting in large and unpredictable changes in the quality of any given foraging territory across years (quality being defined as distance from the currently active nesting colony). When a pair bond forms in bee-eaters, one member typically remains in its natal clan while the other moves into the clan of its partner. At this time, the new pair also establishes its own FHR, generally located within or on the periphery of the clan foraging territory of the natal member. The result of this settlement pattern is that white fronted bee-eaters live their lives spatially surrounded by members of their natal or their matrimonial clan. This, in turn, increases the likelihood of both mutualistic and nepotistic interactions among clan members. Such benefits include shared territory defense, enhanced security against predation, and maintenance of close social bonds with potential helpers. We hypothesize that the adaptive value of clan foraging territories lies in long-term familiarity with a foraging area. Such familiarity was demonstrated to lead to improved foraging efficiency and hypothesized to provide both increased security from predation and a more accurate means of monitoring temporal changes in environmental quality. The system of clan foraging territories found in white fronted bee-eaters differs from the all-purpose group territories of most other cooperative breeders studied to date in two important ways. First, foraging territories were not limiting in the sense of restricting dispersal and “forcing” offspring to remain with their natal clans. Unoccupied areas of seemingly suitable habitat were present throughout the study area at all times. Birds also showed no tendency to expand their boundaries or move into areas vacated when neighboring clans decreased in size or died off. Second, breeding status and foraging territory ownership are not linked in Merops bullockoides. All pairs defended foraging areas, yet only about 3/4 of them bred in any given year. This percentage did not differ significantly between pairs occupying high quality foraging territories (located near the active nesting colony) and pairs forced to abandon low quality foraging territories located more distantly. We conclude that foraging territories are not a critical ecological constraining factor for white fronted bee-eaters in Kenya.     

Territories of male and female terrestrial salamanders: costs,benefits, and intersexual spatial associations

Summary I used a mark-recapture study to estimate home areas for 107 red-backed salamanders (Plethodon cinereus) in a natural forest habitat. Both males and females of this species defend feeding territories, but I presume that some individuals in this relatively highdensity population (approximately 2.8 salamanders per m²) are nonterritorial floaters. Although territorial salamanders exhibited greater numbers of tail autotomies, they had significantly longer relative tail lengths. This difference suggests that territorial individuals gain benefits from territorial ownership. From the observation that home area size was inversely correlated with body size, I infer that larger animals gained higher quality foraging areas. Home areas of adults were significantly more segregated intrasexually and more aggregated intersexually than would be expected from a random distribution. Furthermore, intersexual overlap of home areas was significantly greater than intrasexual home area overlap. Territorial defense of feeding areas by male and female red-backed salamanders therefore also may play a role in mating behavior.     

Why hummingbirds have such large crops

Summary Male Anna's Hummingbirds (Calypte anna) defend territories that contain a predictable food source, floral nectar. For such a hummingbird, the meal size that maximizes long-term net energy intake is less than the maximal crop volume. Smaller meals must be consumed more frequently, but larger meals increase body mass and therefore flight cost. Individuals without territories or with inadequate territories do not have easy access to nectar and intrude on territories owned by otherC. anna, where they may be chased at nay time. It was predicted that these intruders should minimize the number of potentially risky intrusions necessary for maintenance by ingesting as much nectar as possible whenever they manage to feed without being chased (usually when owners are temporarily absent). Therefore, relative to uninterrupted feeding by owners, uninterrupted intruders should feed longer and take larger meals. Field observations supported these predictions. Intruders apparently filled their crops in all seasons, whereas owners ingested smaller amounts (0.21–0.22 ml) and fed for lengths of time consistent with the prediction of an optimization model (0.21 ml). Thus, owners may energetically optimize meal size whereas intruders fill their crops whenever they are not chased. Under most conditions, hummingbirds only fill their crops one-tenth to one-third full, leading to the question why hummingbirds have such large crops. This study demonstrates that a large crop volume may be of survival value when an individual lacks a territory or has inadequate access to resources and must poach on others' territories.     

The role of sea anemones as refuges and feeding habitats for the temperate fish Oxylebius pictus

Synopsis This study describes an association between the temperate zone fish Oxylebius pictus and sea anemones in Barkley Sound, Vancouver Island, British Columbia, Canada. Fish were observed swimming and resting unharmed among the tentacles, or next to a column of Urticina lofotensis (98% of all observations) and U. piscivora (2% of all observations). These associations were most commonly observed at moderately exposed sites where there were relatively high densities of O. pictus and anemones. Most associations occurred at night when the fish were inactive. Small young-of-year fish (< 6 cm total length) associated closely with sea anemones, larger individuals spent less time with their hosts, and most adults (> 12 cm total length) sheltered in rock holes and crevices. Experiments demonstrated that small O. pictus were more vulnerable to predators than large individuals. Urticina lofotensis and U. piscivora were large, persistent structures in the study habitats that provided effective refuge for O. pictus from predators. Anemone size, shape, tentacular adhesive force, and presence of copepod associates influenced patterns of host specificity. Oxylebius pictus fed on copepods and other crustaceans that associated with U. lofotensis and host sea anemones served as both refuges and feeding habitats for the fish. The sea anemones did not appear to receive any significant benefit from the relationship and the fish was considered to be a facultative commensal.     

Determinants of territory size in the pomacentrid reef fish,Parma victoriae

Summary Factors governing the size of territories defended by the pomacentrid reef fish, Parma victoriae, were investigated, prompted by contradictory predictions in the literature concerning the effects of food supply and competitors. Observations were carried out over the non-breeding period (March–October) on a medium density population in which territories were partially contiguous. The territory size of adult fish varied between 3 and 26 m², and was inversely correlated with local densities of conspecifics. The same range in territory size was found for both males and females, which did not differ in the time they spent on territory defence and foraging activities. No correlation existed between territory size and the abundance of algal food, body size, age or time spent on territory defence. Also, there was little variation in territory size over time, despite seasonal changes in the abundance of food algae.Experimental reduction of food supplies on isolated territories of males and females had no effect on territory size. In a higher density habitat an experiment was carried out in which population density and food abundance were simultaneously manipulated. This showed that territory size was primarily determined by intraspecific interactions, as territories exhibited considerable increases in size upon removal of neighbours. No changes in the size of defended areas resulted from either artificial increases or decreases of food levels. There were also no changes in the time spent on defence of territories, foraging time or feeding rates associated with food manipulations or territory expansion, which suggested that food was not a limited resource. This conflicted with current theories proposed to explain territory defence and expansion. It is hypothesized that intraspecific interactions constrain territory size well below the optimum in terms of the abundance of preferred food algal species.