The effect of community on the territorial behavior of the threespot damselfish

Synopsis I examined the hypothesis that animals alter their behavior in response to changes at the community scale of organization, rather than simply reacting to a sequential set of independent interactions with other organisms. The focal species was the threespot damselfish, Stegastes planifrons,a territorial coral reef fish that experiences a high degree of variation in community composition. Field observations of individual threespot damselfish showed a significant increase in the amount of time spent on active territorial defense when the community contained greater proportions of food competitors. Territorial behavior also increased concurrently with higher species diversity in the community. However, no behavioral change was observed in response to the total number of fish entering the defended territory, suggesting that threespot damselfish are responding to more complex environmental cues associated with community structure rather than simple density.     

Brood defence and parental roles in a biparental cichlid fishLamprologus toae in Lake Tanganyika

Brood defence of a cichlid fish,Lamprologus toae, was investigated in its natural habitat in Lake Tanganyika. Both parents guarding a brood attacked both conspecific and heterospecific intruder fishes at different locations. The heterospecific intruder fishes could be classified into three groups on the basis of the locations at which the attacks against each species took place. The distinction of groups by the parents seemed to be primarily based on food habits and feeding behaviour of the intruder fishes. The piscivorous species which were more dangerous for the brood were attacked by both parents at more distant locations from the brood. Parental defence of breeding territory changed with the development of the young. The frequency of attacks against each group decreased after the young reached the size too large for the fishes of the group to prey on. Division of labour in the territorial defence was recognized between male and female. The male parent mainly defended the peripheral region of the territory and the female parent defended the inner region. Significance of the selective attack against intruders and the division of labour between the two sexes in brood defence is discussed.     

An Extreme Case of Interspecific Territoriality: Male Anthidium manicatum (Hymenoptera,Megachilidae) Wound and Kill Intruders

Anthidium manicatum males defend patches of flowering plants as mating territories and copulate with females that enter the territory to obtain nectar or pollen. Territorial males defend their territories not only against conspecific males but also against many other insect species that attempt to utilize the flowers inside the territory. In our study area, a territorial male defended its territory against conspecific males on average only twice per h but it attacked other species of insects on average 70 times per h. Territorial males distinguished between different species of intruders and attacked them unequally. During the hour following experimental removal of the territory holder the number of intruders of some insect species tripled but the number of intruders of those species that were rarely attacked remained approximately the same. A. manicatum males, but not females, bear long spines on the last two segments of the abdomen. Territorial males rammed intruding insects at high speed and shortly before the moment of impact they curved the abdomen forward to hit with the spines. Intruders could be seriously damaged or killed by these attacks. Abdominal spines of A. manicatum males may have evolved as weapons to increase the effectiveness of interspecific territoriality.     

Prior residence, body size and the dynamics of territorial disputes between male freshwater angelfish

Territorial interactions between pairs of size mismatched, sexually mature male angelfish Pterophyllum scalare were investigated in three different conditions: with the larger fish resident (the large resident condition), with the smaller fish resident (the small resident condition) and in a neutral territory (the neutral condition). In the two resident conditions, approximately half of the intruders had previously held territories and half had not. In all categories of fight, one fish showed submissive postures and lost the fight; eventual losers performed both attack and threat at a lower rate than eventual winners. Attack rate declined as the encounter progressed, while rate of performance of threat postures increased. In fights on neutral territories, the larger fish won all fights. In all fights with a resident-intruder asymmetry, the resident fish won the encounter, regardless of relative size. In eventual winners but not in eventual losers, levels of attack were lowest in the neutral encounters. In the small resident condition, levels of attack (corrected for activity of the resident) were lower in intruders that had previously held a breeding territory. Relative size influenced behaviour shown during fights, in that overall intensity was correlated negatively with size differential in all conditions. Thus although prior residence is the primary determinant of the outcome of territorial encounters in this species, both relative body size and prior possession of territory also influence the nature of the interaction.     

Coexistence of permanently territorial cichlids of the genus Petrochromis through male-mating attack

Coexistence mechanisms of permanently territorial fishes proposed hitherto have been mainly based on interspecific competition. To test the hypothesis of the coexistence through male-mating aggression (Kohda, 1995a), spacing patterns of feeding territories of three coexisting herbivorous cichlids, Petrochromis polyodon, P. trewavasae and P. famula, were investigated in a homogeneous habitat at a shallow rocky shore in Lake Tanganyika. All three species maintained individual feeding territories that were defended against both conspecifics and congenerics and rarely overlapped either intra- or interspecifically. Territories of three species were scattered and covered almost all rocky bottoms. Territorial attacks usually occurred near the border of feeding territories. But male P. polyodon, the largest and socially most dominant fish, attacked conspecific males far beyond their feeding territories, which were widely separated. Removal of some territory owners suggested that such attacks result in the separate distribution of male P. polyodon territories. The interstitial space between P. polyodon feeding territories resulting from their mating aggression can be occupied by subordinate congeneric fishes. Male-mating attack of dominant males of Petrochromis will facilitate the coexistence of other congeners.     

Microhabitat characteristics of <Emphasis Type="Italic">Stegastes planifrons</Emphasis> and <Emphasis Type="Italic">S. adustus</Emphasis> territories

Stegastes adustus and Stegastes planifrons are two species of damselfishes commonly found in the Caribbean. These territorial fishes have been widely studied due to their major ecological role on coral reef in controlling the growth of macroalgae that compete with corals for space and, inversely, on their deleterious role in destroying coral tissues to impulse the development of algae. However, few studies were conducted on the biotic and abiotic components of their territories. In the present study, territory size and surfaces of benthic components (macroalgae, algal turf, massive corals, branching corals, Milleporidae, sponges, sand and rubbles) were estimated for the two species at two contrasted sites. At Ilet Pigeon site (IP), the two damselfishes were found at different depth and exhibited different territory sizes. S. adustus defended a larger territory characterized by massive corals, sand and Milleporidae, while S. planifrons territories were smaller, deeper and characterized by branching corals, sponges and rubble. At Passe-à-Colas site (PC), the two fish species coexisted in the same depth range and defended territories of similar size. Their territories presented higher proportions of macroalgae, but smaller surfaces of Milleporidae than at IP. At PC, the main difference between the two species was a higher surface of massive corals inside S. planifrons territories than S. adustus territories. Differences in microhabitat characteristics between the two Stegastes seemed mostly site related. This resulted from the high plasticity of two species, allowing them to persist on Caribbean coral reefs after the decline of most branching acroporids, their former favorite habitats.     

Sex Differences in Territorial Behavior Exhibited by the Spotted Hyena (Hyaenidae, Crocuta crocuta)

Spotted hyenas ( Crocuta crocuta ) are gregarious carnivores that defend group territories against encroachment by neighboring conspecifics. Here we monitored the behavior of members of one clan of free-ranging spotted hyenas during border patrols, 'wars' with neighboring clans, and other interactions with alien intruders, to document differences between the sexes in territorial behavior in this species. We also examined the possibility that the probability or rate of attack on alien hyenas encountered within the clan's territory would vary with the sex of the intruders. Initiation and leadership of most cooperative territorial behaviors were by adult female clan members, although border patrols were occasionally conducted by groups composed exclusively of resident males. The vast majority of alien intruders into the territory of the study clan were males. Resident females were more likely to attack intruding females than intruding males, but hourly rates of aggression directed by females towards aliens did not vary with intruder sex. Resident males were more likely than resident females to attack alien males, and resident males directed significantly higher hourly rates of aggression towards intruding males than females. Although female leadership in most cooperative territorial behaviors distinguishes spotted hyenas from many mammalian carnivores, other sex differences in the territorial behavior of spotted hyenas resemble those documented in other gregarious predators. Sex differences observed in hyena territoriality are consistent with the hypothesis that male and female clan members derive different selective benefits from advertisement and defense of group territories.     

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.     

The effect of patch reef size on fish species richness in a shallow coral reef shore zone where territorial herbivores are abundant

Small patch reefs can harbor many reef fishes because most fishes have a drifting larval phase to randomly disperse over patchy habitats. We examined the species–area relationship (SAR) of damselfish (Pomacentridae) assemblages over 84 small patch reefs (0.05–45.4 m²) using an enlarged section of a high-resolution color aerial photograph as a field map (1/2500) in a shallow coral reef shore zone (<2 m deep, 3.6 ha, Shiraho Reef, Ishigaki Island, Japan). This study confirmed that the logarithmic function is better than other functions (including the power function) to explain the SAR in this scale. Actual species richness (24) over the entire study site was much higher than the species richness (15.4) extrapolated from the regression line in semi-log space. Better estimates were obtained using random placement models and computer simulations. These results suggest that several small patch reefs are likely to have higher species richness than a single large reef of equivalent area at the study site. The total number of individuals of the four most abundant territorial herbivores increased almost linearly with patch reef area, but that of other species roughly increased with the square root of the area. While no territorial species were found in the smallest reefs, the large territorial herbivore, Hemiglyphidodon plagiometopon, was abundant and had negative effects on species richness in large reefs. Although the well-known single-large-or-several-small (SLOSS) debate has largely been settled, this dichotomy can be important in places where territorial herbivores do not occupy the smallest reefs.     

Previous agonistic experience determines both foraging behavior and territoriality in the limpet Lottia gigantea (Sowerby)

Lottia gigantea, the owl limpet, is an algal gardener. Territorial individuals actively defend gardens, which consist of clearedareas in the intertidal zone upon which a thick algal filmdevelops and upon which the territory holder feeds. Smaller,nonterritory holders raid these gardens and graze the algalfilm. Territorial individuals must obtain an adequate ration without compromising the productivity of the garden. In contrast,a nonterritory holder grazing on another limpet's territorymust obtain an adequate ration before it contacts the territoryholder and is driven off. In the laboratory, replicate setsof 10 limpets were trained to behave territorially and nonterritorially.Training mimicked natural encounters between territorial andnonterritorial L. gigantea. Limpets given territorial trainingleft significantly (t = -4.92, df = 9, p =.00041) more algalcover behind when grazing (on average 71%) than did limpetstrained to be nonterritorial (on average 50%). Territorial limpets seldom grazed over the same area more than once (4% of the grazedarea). In contrast, nonterritorial limpets frequently foragedover an area more than once; of the area grazed, 20% had beenvisited more than once. Previous agonistic experience determinesboth territorial behavior and foraging strategies, two of thecritical behaviors necessary for successful gardening behavior.Nonterritorial limpets maximize consumption per unit area, whereas territorial limpets appear to forage prudently, leaving a significantly greater proportion of the plant biomass behind.     

Risk-sensitive resource defense in a territorial reef fish

As coral reefs are home to dense aggregations of a variety of species, aggressive territoriality is often a critical component of individual behavior. Identification and assessment of the risk posed by intruders is crucial to defending a territory, and fishes on coral reefs have been found to attend to body shape, body size, and coloration when responding to intruders. We examined the extent to which dusky damselfish (Stegastes adustus) discriminate among distinct categories of intruders by measuring the distance at which a fish attacks an intruder and the relative intensity and frequency of those attacks. We found that S. adustus discriminated among perceived threats, attacking conspecifics more intensely and more often than egg-predators and herbivores, and showing a trend of attacking those groups more often than invertebrate-feeders, which do not compete with damselfish for resources. Furthermore, territory holders attacked initial-phase wrasses from a farther distance than terminal-phase wrasses, suggesting that they can discriminate among classes of individuals within a species other than their own. Dusky damselfish thus exhibit the ability to make fine distinctions among intruders in a diverse ecosystem.     

Habitat selection by recruits establishes local patterns of adult distribution in two species of damselfishes: Stegastes dorsopunicans and S. planifrons

Local patterns of adult distribution in organisms that disperse young as pelagic larvae can be determined at the time of recruitment through habitat selection or, shortly thereafter, through post-recruitment processes such as differential juvenile survivorship and interspecific competition. This study addresses the importance of habitat selection by recruits in establishing the local pattern of adult distribution in two sympatric Caribbean damselfish species, Stegastes dorsopunicans and S. planifrons. Both species inhabit shallow reefs but show little overlap in their distribution; S. dorsopunicans predominates in the reef crest and S. planifrons occurs primarily on the reef slope. Furthermore, S. dorsopunicans is associated with rocky substrate, while S. planifrons occupies live coral. The substrate cover follows a similar pattern with coral being much less common on the reef crest than on the reef slope. Monitoring recruitment every other day in reciprocal removal experiments and artificial reefs indicates that the observed pattern of local adult distribution is a product of habitat selection for both species. The presence or absence of conspecifics did not influence recruitment patterns for either species. Stegastes dorsopunicans recruited primarily to shallow, rocky areas, appearing to cue on both substratum type and depth. Stegastes planifrons recruited exclusively to coral substratum independent of depth. These results indicate that local adult patterns of distribution can be explained by habitat selection at recruitment, and that substrate type and depth may be important cues. Received: 27 May 1997 / Accepted: 4 January 1998     

Conspecific agonistic behaviour in the Mediterranean parrotfish

acta ethologica - Aggressive behaviour in fishes, particularly in territorial species, is a common trait used to defend resources such as food or mates. Territorial males of the Mediterranean...     

Territoriality and nutritional condition in Cynotilapia afra (Gunther) and Pseudotropheus zebra (Boulenger), cichlidae in Lake Malawi National Park, Malawi

The standard length, diurnal activities, territory sizes, and areas over which individuals foraged and the nutritional condition of territorial and non-territorial Cynotilapia afra and Pseudotropheus zebra were compared. Results show that territorial tenureship in these fishes does not depend on the male size, implying that aggressiveness, experience and motivation are more important in the maintenance of territory. However, terri-toriality in C. afra (t = 10.93, P < 0.05) and P. zebra (t = 3.31, P < 0.05) manifests itself in the reduction of nutritional condition because territorial males engage in energetically more demanding activities, e.g. courtship displays, fertilizing ripe females and chasing intruders that trespass into their territories. Territorial C. afra (t = 4.77; P < 0.05) and P. zebra (t = 5.89; P < 0.05) also fed over significantly smaller areas and spent significantly less time feeding than did non-territorial males. The biological significance of territoriality in these fish species is therefore not food intake, but mate attraction and reproductive success for which they trade off their nutritional condition. However, there might be a nutritional threshold below which the cost out-weighs the benefit. Hence, territorial males in poorer health abandon their territories in order to regain their condition.     

Response of Pleistocene Coral Reefs to Environmental Change Over Long Temporal Scales

SYNOPSIS. TWO studies from the Pleistocene coral reef fossilrecord demonstrate the sensitivity of reef communities to bothlocal environmental parameters and habitat reduction. In thefirst study, Pleistocene reef coral assemblages from Papua NewGuinea show pronounced constancy in taxonomic composition andspecies diversity between 125 and 30 ka (thousand years). Spatialdifferences in reef coral community composition during successivehigh stands of sea level were greater among sites of the sameage than among reefs of different ages, even though global changesin sea level, atmospheric CO₂ concentration, tropical benthichabitat area, and temperature varied at each high sea levelstand. Thus, local environmental variation associated with runofffrom the land had greater influence on reef coral communitycomposition than variation in global climate and sea level.Proportional sampling from a regional species pool does notexplain the temporal persistence and local factors likely playeda major role. Examination of coral reef response to global changeshould not only involve regional diversity patterns but alsolocal ecological factors, and the interactive effects of localand global environmental change. In the second study, Pleistocene extinction of two widespread,strictly insular species of Caribbean reef corals, Pocilloporacf. palmata (Geister, 1975) and an organ-pipe growth form ofthe Montastraea "annularis" species complex, was natural anddid not involve gradual decrease in range and abundance, butwas sudden (thousands of years) throughout the entire range.One explanation is that sea level drop at the Last Glacial Maximum(LGM—18 ka) resulted in a threshold of habitat reduction,and caused disruption of coral metapopulation structure. Thresholdeffects predicted by metapopulation dynamics may also explainthe apparent paradox of the large amount of degraded modernreef habitat without any known modern-day reef coral extinctions.The rapid extinction of widespread Pleistocene species emphasizesthe vulnerability of reef corals in the face of present rapidenvironmental and climatic change.     

Matter of Size or Matter of Residency Experience? Territorial Contest in a Green Hairstreak,Chrysozephyrus smaragdinus (Lepidoptera: Lycaenidae)

Territorial residents usually win asymmetrical owner‐intruder contests and a variety of hypotheses have been proposed to explain this phenomenon. In the butterfly Chrysozephyrus smaragdinus (Lepidoptera: Lycaenidae), male territorial residents defended their territories against intruders during numerous contests and kept them for many successive days. Field observations and experiments were conducted to examine the factors related to this superiority of residents. Forewing length did not differ between residents and intruders, suggesting that body size is not correlated with resource holding potential. Removal–replacement experiments demonstrated that residency did not serve as an arbitrary means for contest settlement, and did not support the recently presented alternative hypothesis that males with higher body temperature are more likely to win. New residents fought longer in defense of the territory as their residence duration in the territory increased. I discuss these results in light of game theory and suggest that the superiority of residents in C. smaragdinus may be based on the asymmetry of resource (territory) value for residents and intruders.     

Male territoriality,fighting and assessment of fighting ability in the Australian frog Uperoleia rugosa

Males of Uperoleia rugosa form aggregations around the breeding pond. Within the aggregations, each calling male maintains an exclusive zone in which it does not tolerate another calling male. However, attacks on silent intruders, satellite males or females were never observed. Territorial males warned calling intruders by changing from advertisement calls to encounter calls (a display); if the intruder persisted in calling the resident would either attack or retreat and call elsewhere. Fights were usually won by the heavier males, and males of similar weight were more likely to fight, whereas males retreated from opponents that were substantially heavier. Calling was energetically costly and territorial males lost weight and thus their fighting abilities declined, whereas silent satellite males gained weight and were able to oust lighter territorial males. There was thus a dynamic interchange between territorial and satellite males depending on their relative weights. The playback of recorded advertisement calls showed that males can assess each others' fighting abilities on the basis of their vocalizations. Males retreated from a loudspeaker broadcasting the calls of a larger male but they attacked the loudspeaker if the calls of smaller or similar sized males were played (N=39, P<0·001, one-tailed binomial test). Dominant frequency was probably the call parameter used in assessment as it was correlated with the weight (r=−0·71, P<0·001, N=100) and hence the fighting ability of the caller, but this hypothesis needs further testing.     

Prehistoric fisheries in the Caribbean

We studied faunal remains from archaeological sites on five Caribbean islands, each with an early (1,850-1,280 years B.P.) and late (1,415-560 years B.P.) occupation. On each of these islands (Puerto Rico, St. Thomas, St. Martin, Saba, and Nevis), the mean size of reef fishes in the faunal remains declined from the early to the late occupation. The large samples from sites on St. Thomas and Nevis allowed examination of the size distribution of individual taxa. Samples of obligate reef fishes (Scaridae, Acanthuridae, Lutjanidae, and Serranidae) showed large reductions in size between the early and late occupations. Samples of facultative reef fishes (Carangidae and Clupeidae) showed little change in size frequency distribution. The percentage of estimated reef fish biomass in the total aquatic faunal record sharply declined in the samples from four of the islands, while on Nevis there was a slight increase. The mean trophic level of reef fishes declined from the early to the late occupations on each island. Together these patterns suggest that populations of reef fishes adjacent to occupation sites on these islands were heavily exploited in prehistoric times. Such exploitation resulted in shifts in size structure and species composition among the reef fish fauna. On some islands the decline in reef fish resources corresponded with a shift towards greater exploitation of pelagic species.     

Field experiments on species recognition by the threespot damselfish, Eupomacentrus planifrons, (Pisces: Pomacentridae)

The threespot damselfish, Eupomacentrus planifrons, defends differently sized territories against individuals of different species. Tests were conducted in the field to determine the mechanism(s) by which this interspecific recognition occurs. It was found to be based largely on visual cues, the most important of which was body form. Both colour and movement had different effects for intra- and inter-specific intruders. Based on these data, it is proposed that different mechanisms underlie the recognition of and territorial defence against the two groups. Further, interspecific territoriality need not be an ‘accidental’ form of intraspecific territoriality nor a transitional phase in the divergence of already established species, but, rather, may be a stable system based on the simple need to defend an area against non-related competitors.     

Behavioral interactions between salamanders and centipedes: competition in divergent taxa

Red-backed salamanders, Plethodon cinereus, use territorialadvertisement in the form of agonistic displays and pheromonalscent marking as a mechanism for intraspecific interferencecompetition. Although ecological and behavioral interactionsamong species of salamanders have been well studied, littleis known about the interactions between territorial P. cinereusand other ecologically similar species, such as large predatoryinvertebrates. Our field data indicate that P. cinereus anda large syntopic centipede, Scolopocryptops sexspinosus, exhibitnegative spatial associations in natural habitats, possiblyindicating interspecific territoriality. Only seven instancesof salamander/centipede co-occurrence were recorded from a fieldsample of 247 occupied cover objects. Cover object size waspositively correlated with salamander SVL (tip of the snoutto the anterior end of the cloaca), but there was no correlationof cover object size to centipede length. Data on the abilityof P. cinereus to differentiate among chemicals on the substratesuggest that visual cues are not necessary to elicit a territorialresponse from intruding salamanders. Although in laboratorytrials salamanders behaved similarly toward intruders of bothspecies, biting was directed only toward centipedes. Salamandersspent significantly more time approaching centipedes than theydid approaching other salamanders. Approach behavior was oftenassociated with nose tapping and may be an investigative, ratherthan aggressive, behavior. We suggest that territorial P. cinereusrespond similarly to intruding salamanders and centipedes, butthat they escalate more readily to biting centipedes becauseS. sexspinosus is sightless and thus unable to respond to visualsignals.