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.     

A review of hybridization in marine angelfishes (Perciformes: Pomacanthidae)

Synopsis Although hybridization of terrestrial and freshwater organisms has been well-studied, very little work has focused on hybridization among coral reef fish species. In the present paper, eleven examples of probable hybrids between marine angelfishes (Pomacanthidae) are reviewed. Evidence is presented which strongly suggests that the nominal speciesApolemichthys armitagei is invalid and that specimens previously identified as this species represent hybrids betweenA. trimaculatus andA. xanthurus. Of the remaining ten probable pomacanthid hybrids, five are inCentropyge (C. eibli x C. flavissimus, C. eibli x C. vrolikii, C. flavissimus x C. vrolikii, C. loricu0lus x C. potteri, andC. multifasciatus x C. venustus); one inHolacanthus (H. bermudensis x H. ciliaris), and four inPomacanthus (P. arcuatus x P. paru, P. chrysurus x P. maculosus, P. maculosus x P. semicirculatus, andP. sexstriatus x P. xanthometapon). An additional five examples of possible pomacanthid hybrids are described, two inCentropyge, two inChaetodontoplus and one inPomacanthus. Examination of hybrids may provide clues on reproductive behavior, dispersal capabilities, and phylogenetic relationships of species. More studies on hybridization in coral reef fish species, particularly those involving molecular techniques, are needed.     

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.     

Two alternative female tactics in the polygynous mating system of the threadsail filefish,Stephanolepis cirrhifer (Monacanthidae)

Reproductive behavior of the threadsail filefishStephanolepis cirrhifer was studied at Kashiwajima, southern Shikoku, Japan. This species spawned in pairs on the sandy bottom, the eggs being scattered over an area of about 15 cm in diameter and attached to sand particles. After spawning, males departed immediately, while the females remained at the site to guard the eggs for a few minutes. Thereafter the eggs were left unguarded for three days until hatching. Females spawned only once daily, whereas males mated with multiple females in succession. The reproductive males established territories, in which 1–4 resident females defended smaller territories from each other. The harem size changed according to some ecological conditions, such as population density. Moreover, the males also mated with visiting non-resident females. Thus, the two alternative tactics of females resulted in two mating patterns, haremic polygyny and female visiting of male territories, in a single population ofS. cirrhifer.     

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.     

Female defense polygyny in the territorial triggerfish <Emphasis Type="Italic">Sufflamen chrysopterum</Emphasis>

The occurrence of polygyny requires specific environmental conditions such as female aggregation or patchy resource distribution. However, it is difficult to determine the factors responsible for polygyny in species in which the territories of both sexes overlap. To overcome this, we performed female removal experiments in the polygynous triggerfish Sufflamen chrysopterum (Balistidae) in the Okinawa coral reef. Both sexes defended their territories exclusively against consexuals of the same species, and female aggregation was absent. Each male territory included 1–3 female territories, and nonterritorial males were significantly smaller than territorial males. Further, the body size of territorial males was positively correlated with that of the largest female in their territories, and larger males tended to mate with more females. The results of the female removal experiments (n = 10 females) indicated that females competed for better territories rather than larger mates. In contrast, males abandoned the territories once the females emigrated. These results strongly suggest that males defend females rather than sites and compete for larger and a greater numbers of females. Thus, in S. chrysopterum, female defense polygyny occurs in the absence of female aggregation.     

Overlapping territory of the benthophagous cichlid fish, Lobochilotes labiatus, in Lake Tanganyika

Synopsis Feeding territories do not usually overlap in the same fish species. We studied overlapping territories and feeding behaviours of a benthophagous cichlid fish Lobochilotes labiatus Boulenger in a coastal area of Lake Tanganyika. Forty-four individuals of both sexes, 6–31 cm in total length, maintained territories in a 21 m × 25 m area. These could be regarded as feeding territories. The territories of similar-sized owners (usually different by less than 5 cm) did not overlap each other, but those of different-sized owners overlapped widely. Aggressive interactions were rarely observed between fish distinctly different in body size. Both large and small fish took benthic animals of the same kinds and of similar size. In contrast, the size of foraging sites (crevices) was different between them: the large fish foraged at large crevices and small fish at small crevices. This partitioning of foraging sites may enable the overlap of territories in this benthivore.     

Territorial defense against various food competitors in the Tanganyikan benthophagous cichlid <Emphasis Type="Italic">Neolamprologus tetracanthus</Emphasis>

Territorial defense of nonbreeding female Neolamprologus tetracanthus, a shrimp-eating Tanganyikan cichlid, was investigated. Females defended territories (=home ranges, ca. 1m across) against a variety of intruding fishes. Conspecific females were usually attacked outside the territories, heterospecific benthivores (shrimp eaters) and omnivores near the border of the territories, and piscivores, algae and detritus feeders, and herbivores inside the territories. Females used some parts of the sandy substrate in the territories for foraging (foraging areas). Territorial defense prevented most of the conspecific females and benthivores from intruding into the foraging areas. In omnivores, piscivores, and algae and detritus feeders, about half the intruders were repelled from the foraging areas, although herbivores were infrequently repelled in the areas. Soon after removal of the resident females, many food competitors invaded the foraging areas and eagerly devoured prey, suggesting that the territories are maintained for food resource protection from these competitors. Females are likely to discriminate intruding fishes and change their territorial defense primarily on the basis of the degree of dietary overlap, resulting in monofunctional serial territories.     

Male mating success,sexual size dimorphism,and site fidelity in two species of Malacoctenus (Labrisomidae)

Synopsis In both Malacoctenus hubbsi and Malacoctenus macropus, males defended preferred oviposition sites from both other males and potential egg predators. In M. hubbsi, adult females were larger than adult males. Larger M. hubbsi males were not associated with territory parameters that were correlated with higher mating success, and male size was not correlated with mating success. Male size did affect mating success when territory parameters were statistically controlled for, but the failure of large males to associate with better territories eliminated any mating advantage for larger males. In M. macropus, males are larger than females. Larger males defended preferred oviposition sites, and had higher mating success than did smaller males. Male M. macropus also had much higher site fidelity than male M. hubbsi. These results suggest that the evolution of the differences in site fidelity and sexual size dimorphism between these two species may be due to sexual selection acting differentially in these two species.     

Feeding Territoriality in Female Northern Bats,Eptesicus nilssoni

A nursing colony of 26 Northern bats was studied in South-Central Sweden. The foraging behaviour is described with emphasis on social interactions on feeding grounds. The bats foraged in small feeding territories used night after night. The reproducing females defended feeding territories against other colony members as well as non-members by means of aggressive chases and vocalizations. A linear dominance order occurred among the females that regularly visited a feeding site. Intrusions into already occupied feeding sites resulted in territorial conflicts (47 %), passive departure by one or both opponents (43%) or mutual acceptance (11%). Conflicts occurred throughout the summer, but became less frequent in July when insect abundance increased.     

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.     

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.     

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.     

Feeding territoriality and the reproductive consequences in brown skuas<Emphasis Type="Italic"> Catharacta antarctica lonnbergi</Emphasis>

In the maritime Antarctic, brown skuas (Catharacta antarctica lonnbergi) show two foraging strategies: some pairs occupy feeding territories in penguin colonies, while others can only feed in unoccupied areas of a penguin colony without defending a feeding territory. One-third of the studied breeding skua population in the South Shetlands occupied territories of varying size (48 to >3,000 penguin nests) and monopolised 93% of all penguin nests in sub-colonies. Skuas without feeding territories foraged in only 7% of penguin sub-colonies and in part of the main colony. Females owning feeding territories were larger in body size than females without feeding territories; no differences in size were found in males. Territory holders permanently controlled their resources but defence power diminished towards the end of the reproductive season. Territory ownership guaranteed sufficient food supply and led to a 5.5 days earlier egg-laying and chick-hatching. Short distances between nest and foraging site allowed territorial pairs a higher nest-attendance rate such that their chicks survived better (71%) than chicks from skua pairs without feeding territories (45%). Due to lower hatching success in territorial pairs, no difference in breeding success of pairs with and without feeding territories was found in 3 years. We conclude that skuas owning feeding territories in penguin colonies benefit from the predictable and stable food resource by an earlier termination of the annual breeding cycle and higher offspring survivorship.Research licence: Umweltbundesamt Bonn 13.4-94003-1/5-7.     

Mating system of the lefteye flounder,Engyprosopon grandisquama

Underwater observations were conducted on the reproductive behavior and mating system of the lefteye flounder,Engyprosopon grandisquama, off Nagashima, southwest of Kyushu Island, Japan. Two types of males were found: large males, which defended territories against other large males, and small males, which did likewise but only against smaller males. Large males established territories which encompassed or ovelapped the home ranges of 1 or 2 cohabitant females. Territories of the small males, in which a smaller female often maintained a home range, overlapped those of large males. Pair spawning occurred around sunset. Mating of large males with cohabitant females was observed 36 times and that of a small male with a smaller female once. Pair formation was assorted by body size, paired males being larger than females in most cases. Thus, inE. grandisquama, gigamous large males were common, small males occurring within the former's erritorial boundaries mating with smaller females.     

Male foraging avoidance in female feeding territories in a harem polygynous cichlid in Lake Tanganyika

We studied foraging site partitioning between the sexes in Neolamprologus tetracanthus, a shrimp-eating Tanganyikan cichlid with harem-polygyny. Females maintained small territories against heterospecific food competitors within large territories of males, foraging exclusively at the inner side of their own territories (foraging areas). Males fed as frequently as females in their own territories, but mostly outside female foraging areas, although they frequently entered female territories and repelled food competitors from the territories. Soon after removal of the resident females, however, harem males, as well as many food competitors, invaded the vacant territories and intensively devoured prey of female foraging areas. This indicates that although female foraging areas appear to contain more food than outside the areas, harem males refrained from foraging there when the resident females were present. We suggest that harem males will attempt to keep female foraging areas in good condition, whereby they may get females to reside in male territories and/or promote female gonadal maturation.     

Aggressive behaviour in Montagu’s harrier Circus pygargus during the courtship period

Aggressive behaviour of Montagu’s harrier was observed during the pre-laying period in the 1992–1995 seasons on the calcareous marshes of Chelm in Eastern Poland. In total, 435 flights performed by 24 pairs of individually marked harriers were analysed. All flights were performed in relation to the territory of defence. Males performed 61% of aggressive interactions while females performed 39%. Intraspecific and interspecific aggression comprising direct attacks (58%), escorts (13%), pursuits (28%) and “mirror behaviour” (1%) were described for the first time. Interspecific aggressive behaviour occurred only near territory boundaries. The main intruders were marsh harriers Circus aeruginosus breeding on the same marshes. Occasionally, harriers attacked curlews Numenius arquata, short-eared owls Asio flammeus, magpies Pica pica, hooded crows Corvus corone cornix, common buzzards Buteo buteo or hobbies Falco subbuteo. Four cases of communal mobbing were observed. Three to five males from the neighbourhood attacked the intruders together. Intraspecific aggressive behaviour was observed in the pair territory, near the boundary or at a distance of up to 100 m from the defended area. Aggressive interactions performed by both sexes were more frequent towards birds of the same sex. Females defended their territories more aggressively against females. Similarly, males were more aggressive against males. All cases of aggressive behaviour were observed near harrier territories. Aggressive relations between birds outside breeding territories or when foraging around marshes were not observed.     

Bigamy or monogamy with maternal egg care in the triggerfish,Sufflamen chrysopterus

Reproductive behavior and mating systems of the triggerfish,Sufflamen chrysopterus (Balistidae), were studied on the fringing reef of Sesoko Island, Okinawa. Both males and females maintained territories against consexual adults, feeding on benthic animals within their own territories. Each male territory overlapped one or two female territories, with mating occurring between the cohabitants. The monogamous males were smaller and foraged more frequently than the bigamous ones, suggesting that the former allocated more energy to growth rather than to improving reproductive success. Pair spawning occurred around sunrise, females only taking care of the demersal eggs until hatching, which occurred around sunset of the same day. On spawning days females foraged less frequently than usual, but as frequently as males. Females spawned at intervals of 5–7 days, usually shifting sites within their territories. Thus both feeding and spawning sites were available for females within their territories, providing males with the opportunity to monopolize females by defending their territories.     

Intra- and interspecific dominance hierarchies and variation in foraging tactics of two species of stream-dwelling chars

Aggressive interactions, foraging behavior, habitat use and diet were studied in sympatric populations of white-sported char,Salvelinus leucomaenis, and Dolly Varden,Salvelinus malma, in a Japanese mountain stream. Underwater observations on individuals of both species revealed two distinct behavioral regimes: aggressive drift foragers and non-aggressive benthos foragers. Aggressive drift foragers defended partial territories around focal points from which they made forays to capture invertebrates drifting in the water column. Non-aggressive benthos foragers cruised around and beneath cobble in large foraging ranges that overlapped each other. Intra- and interspecific, size-dependent dominance hierarchies were recognized among aggressive drift foragers, whereas non-aggressive benthos foragers showed no such relationships. Terrestrial invertebrates were the most abundant prey in the diets of drift foragers, whereas a very small proportion of the diet of benthos foragers was made up of these taxa. Benthos foragers showed more complex diet composition than drift foragers. These results suggest that non-aggressive benthos foragers may avoid not only interference but also exploitative competition by using alternative foraging tactics. The proportion of drift foragers to benthos foragers among white-spotted char was more than 35 times that among Dolly Varden. The significant difference in the proportion of each species using the two types of foraging strategy results in interspecific food segregation in sympatric populations.     

MORPHOLOGY,TERRITORIALITY AND MATING SYSTEM OF THE PINTAILED WHYDAH VIDUA MACROURA

Savalli, U. M. 1995. Morphology, territoriality and mating system of the Pintailed Whydah Vidua macroura. Ostrich 66: 129–134.

The biology of the Pintailed Whydah Vidua macroura was studied at the Kakamega Forest, western Kenya. This species is sexually dimorphic in plumage and size (males are brighter, long tailed and larger). Males defended large (1.4 ha) territories which contained areas of bare ground (9% of total area) suitable for feeding on grass seeds such as Paspalum scrobiculatum. There were two breeding peaks: during the long rains (April-August) and the short rains (November-December). Territorial interactions were frequent; a previously unreported tail-uphill-wiping display is described. Females frequently visited male territories and were pursued and courted by the males. Male tarsus length was weakly, but positively, related to the size of feeding area (a possible indicator of territory quality), but there were no other significant correlates with territory size, or frequency of intrusions. There were no significant correlates of female visitation rates (which do correlate with copulation frequency), so the basis of female choice (if any) remains unknown. Although this species has been classified as an exploded lekker, the possibility that females are attracted to resources (such as grass seeds) cannot be ruled out. Tail streamer length was not more variable than other morphological traits when fully grown, but was much more variable at the start of the breeding season while still growing.