Caribbean damselfish with varying territory quality: correlated behaviors but not a syndrome

The behavioral syndrome hypothesis suggests that individualanimals within a population behave differently due to specificbehavioral types, and these should be consistent across behaviorsor in different contexts. In contrast, for animals that livewithin an environment in which territory quality can changeover time, natural selection should have favored behavioralflexibility and modulation of the cost of defense in relationto territory quality. This would require assessment of the territoryfollowed by displays of appropriate types and intensities ofbehavior. We examined the territorial behavior of male beaugregorydamselfish (Stegastes leucostictus) by enhancing territory qualityusing artificial breeding sites and comparing their behaviorto males on lower quality natural sites. When male fish weredefending high-quality artificial territories, they had higherlevels of aggression toward male conspecifics and courtshiptoward females than when on low-quality natural territories.We also found that aggression and courtship behaviors were correlatedon natural sites but not on artificial sites. Behaviors werenot correlated within individuals when males switched from naturalto artificial territories or from artificial to natural territories.These results indicate that males assess their current territoriesand adjust behaviors accordingly and that courtship and aggressivebehaviors are not linked within a permanent behavioral syndrome.     

Short-term behavioral consequences of territory relocation in a Caribbean damselfish, Stegastes diencaeus

Home-range relocation occurs during natal and breeding dispersal,ontogenetic habitat shifts, and the maintenance of resource-or density-dependent patterns of distribution. Relocating animalsare expected to change their behavior to compensate for limitedfamiliarity with the new home range and with neighboring conspecifics;such changes may indicate some of the costs of relocation. Littleis known, however, about the magnitude and duration of the changesor about the types of behavior affected. We investigated theshort-term (2 day) behavioral changes associated with relocationin the highly territorial longfin damselfish, Stegastes diencaeus.We compared the behavior of newcomers settling into experimentallycreated vacancies in an established neighborhood with that ofthe original residents of the same territories. The greatestdifference was an increase in the rate of agonistic interactions.Newcomers also used smaller territories, moved more, and fedless. Neighboring damselfishes were less aggressive toward neighborsthat expanded into vacant territories than toward "strangers"that relocated from elsewhere. The behavior of newcomers approachedthat of original residents within 2 days but territory sizedid not. These observations suggest that relocating a territoryincreases energy expenditure and decreases energy intake. Suchcosts could explain the philopatry of reef fish when alternativelocations are of uncertain quality or are only slightly better.Nevertheless, they are unlikely to outweigh the long-term benefitsof obtaining a superior territory—especially for individualsfrom adjacent territories.     

Habitat choice in adult longfin damselfish: territory characteristics and relocation times

Habitat selection by coral reef fish during initial settlement has been shown to depend on various biotic and abiotic characteristics. However, relatively little is known of the factors influencing habitat choice by adults during post-settlement processes such as relocation or migration. In this study, we first characterised the habitat of longfin damselfish (Stegastes diencaeus Jordan and Rutter) territories to quantify territory variability. Characteristics such as percentage cover of rock, sand, live coral and distance from sand were highly variable, while territory area, turf and macro algae cover were relatively uniform across territories.We then assessed the importance of specific habitat characteristics by experimentally removing damselfish and measuring recolonisation times in relation to these characteristics. The presence of nest sites markedly increased the speed of territory recolonisation after experimental removals. Other variable territory characteristics such as substrate type, rugosity and the presence of cleaning stations did not affect recolonisation speed. In general, males recolonised territories faster than females, and males were more likely to recolonise territories previously owned by males with an active nest site. Thus, intraspecific competition for high-quality nest sites may generate sex differences in territory relocation and highly stable sex-specific patterns of adult distribution.     

The deterrent effect of bird song in territory defense

Using the responses of territory owners to playback to inferthe territorial function of acoustic signals is common practice.However, difficulties with interpreting the results of suchexperiments have obscured our understanding of territorial signalling.For instance, a stronger response to playback is often interpretedas more aggressive, but there is no consensus as to whetherthis should be in response to the least or most threateningsimulated intruder. Rather than following a gradual increaseor decrease, the relationship between signal intensity and responsestrength may instead describe a peaked curve. We manipulatedbanded wren (Thryophilus pleurostictus) songs to simulate low-,median-, and high-performance singers and used these songs asstimuli in playback experiments. Banded wrens were less likelyto approach the high-performance stimulus compared with thelow- and median-performance stimuli. However, the birds thatdid approach the high-performance stimulus sang more than thosethat approached the low-performance stimulus. In addition, birdswere more likely to match the songs when exposed to the median-and high-performance stimuli compared with the low-performancestimuli, and song matching predicted approach behavior. Theseresults are in accordance with theoretical models of aggressiveencounters in which low-performance opponents are challengedwithout further assessment. Median- and high-performance opponents,however, may require further assessment, and the latter maybe perceived as too intimidating for approach.     

Territorial defense, territory size, and population regulation

The carrying capacity of an environment is determined partly by how individuals compete over the available resources. To territorial animals, space is an important resource, leading to conflict over its use. We build a model where the carrying capacity for an organism in a given environment results from the evolution of territorial defense effort and the consequent space use. The same evolutionary process can yield two completely different modes of population regulation. Density dependence arises through expanding and shrinking territories if fecundity is low, breeding success increases gradually with territory size, and/or defense is cheap. By contrast, when fecundity is high, breeding success sharply saturates with territory size, and/or defense is costly, we predict fixed territory sizes and regulation by floaters. These surplus individuals form a buffer against population fluctuations. Yet floaters can also harm breeder performance, and by comparing population growth of a territorial population to a nonterritorial (and individually suboptimal) alternative, we can quantify the harmful effect of evolutionary conflict on population performance. Territoriality has often been found to increase population stability, but this may come at a cost of reduced equilibrium densities.     

Neuromolecular correlates of cooperation and conflict during territory defense in a cichlid fish

Cooperative behavior is widespread among animals, yet the neural mechanisms have not been studied in detail. We examined cooperative territory defense behavior and associated neural activity in candidate forebrain regions in the cichlid fish, Astatotilapia burtoni. We find that a territorial male neighbor will engage in territory defense dependent on the perceived threat of the intruder. The resident male, on the other hand, engages in defense based on the size and behavior of his partner, the neighbor. In the neighbor, we find that an index of engagement correlates with neural activity in the putative homolog of the mammalian basolateral amygdala and in the preoptic area, as well as in preoptic dopaminergic neurons. In the resident, neighbor behavior is correlated with neural activity in the homolog of the mammalian hippocampus. Overall, we find distinct neural activity patterns between the neighbor and the resident, suggesting that an individual perceives and processes an intruder challenge differently during cooperative territory defense depending on its own behavioral role.     

The importance of species identification and location on interspecific territorial defense by the damselfish,Stegastes fasciolatus

Synopsis The Hawaiian damselfish, Stegastes fasciolatus, defends an all-purpose territory on shallow-water reefs. By conceptually dividing each territory into 3 horizontal and 2 vertical zones, each intruder into the territory was recorded by species, zones violated, and reaction of the resident. Intruders included 3 herbivorous, 7 omnivorous, and 2 carnivorous species. A BMDP 4F analysis for multiway tables was applied to examine the relationships between the proximate causal factors eliciting the chase response by a resident. Species of intruder and their position in the territory had an additive effect on the chase response, with position in the territory as the less important factor. Distance from the bottom was not indicated as an important factor. The ecological implications of these results are discussed.Senior author     

Modification of benthic communities by territorial damselfish: a multi-species comparison

The effects of territorial damselfish on coral reef benthos have been well-studied for a few relatively large-bodied species with visually distinct territories. Despite a growing body of research demonstrating their abundance, and their effects on algae, corals and other grazers, there has been little research on the effects of the territorial damselfish community as a whole. This study investigated the space occupation, territory composition, and diet of ten damselfish species at three locations: Magnetic and Orpheus Islands (Great Barrier Reef), and Kimbe Bay (Papua New Guinea). Territories were measured, and the composition of benthic communities inside and outside territories was assessed both in situ and from algal collections. The stomach contents of territorial damselfishes were also quantified. Although the larger, previously well-studied species had the most visible effect on the benthic community in their territories, all the smaller species also significantly affected the algal composition, normally with an increase of palatable algae. However, the composition of algal assemblages inside the territories of different species varied considerably. Damselfish territories were highly individual, not just among species, but also among locations. Diets were diverse and indicated a greater degree of omnivory and detritivory than previously thought. At all locations, territories occupied a substantial proportion of the substratum: >25% on Magnetic Island, >50% at Orpheus Island, and ∼50% in Kimbe Bay. Within individual zones, this figure was as high as 70%. The contribution of territorial damselfishes to a range of benthic patterns and processes is considerable, and future benthic studies may need to distinguish more closely between territory and non-territory areas. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Size-dependent hyaluronate degradation by cultured cells

Hyaluronate degradation was examined in cultures of vascular wall cells (bovine aortic endothelial cells, rat aortic smooth muscle cells) and in nonvascular cells (chick embryo fibroblasts). The three cell types examined all produced hyaluronidase activity in culture which had a strict acidic pH requirement for activity. This suggested that the enzyme was active only within an acidic intracellular compartment and therefore that hyaluronate degradation occurred at an intracellular site. This was supported by the observation that the presence of hyaluronidase activity alone was not sufficient to ensure degradation of extracellular hyaluronate. Rather, the key limiting factor in this process appeared to be hyaluronate internalization, and this was found to be hyaluronate size-dependent and to a degree, cell-specific. The relationship of these results to morphogenesis and tissue remodeling is discussed.     

Size-dependent flight initiation by a lotic mayfly in response to a predatory fish

1. Laboratory experiments were performed to determine whether flight initiation by lotic mayfly larvae of Baetis tricaudatus exposed to the longnose dace ( Rhinichthys cataractae ) is affected by an individual's size and its physiological state.
2. We used a three factorial ANOVA design to test whether flight initiation distances (FIDs) were affected by Baetis size (small, large), starvation level (low, high), and the length of a predator-free foraging period (short, long).
3. FIDs were significantly affected by the interaction between Baetis size and starvation level, and the main level effects of starvation and Baetis size. FIDs of small larvae were unaffected by starvation level and the length of the predator-free feeding period, whereas starvation reduced FIDs of large larvae 2-fold.
4. Subsequent experiments showed that size-dependent differences in FIDs could not be attributed to size-specific differences in the economics of rates of food intake or gut evacuation. For both small and large larvae, food intake rates declined with increasing time spent foraging and both small and large larvae consumed the majority (70–75%) of food within the first 3 h of the 12-h feeding period. Similarly, gut evacuation declined with increasing starvation time and rates of evacuation did not differ between small and large larvae. These data suggest that size-dependent differences in flight initiation by Baetis larvae do not involve the energetics of food intake or processing.     

Size-dependent flight initiation by a lotic mayfly in response to a predatory fish

1. Laboratory experiments were performed to determine whether flight initiation by lotic mayfly larvae of Baetis tricaudatus exposed to the longnose dace ( Rhinichthys cataractae ) is affected by an individual's size and its physiological state.
2. We used a three factorial ANOVA design to test whether flight initiation distances (FIDs) were affected by Baetis size (small, large), starvation level (low, high), and the length of a predator-free foraging period (short, long).
3. FIDs were significantly affected by the interaction between Baetis size and starvation level, and the main level effects of starvation and Baetis size. FIDs of small larvae were unaffected by starvation level and the length of the predator-free feeding period, whereas starvation reduced FIDs of large larvae 2-fold.
4. Subsequent experiments showed that size-dependent differences in FIDs could not be attributed to size-specific differences in the economics of rates of food intake or gut evacuation. For both small and large larvae, food intake rates declined with increasing time spent foraging and both small and large larvae consumed the majority (70–75%) of food within the first 3 h of the 12-h feeding period. Similarly, gut evacuation declined with increasing starvation time and rates of evacuation did not differ between small and large larvae. These data suggest that size-dependent differences in flight initiation by Baetis larvae do not involve the energetics of food intake or processing.     

Muscle transplantation by fascicular territory

In some muscles, distinct and separate portions of the muscle are each under the control of a different fascicle of the motor nerve. Although there is some slight overlap in areas, this unit, a single fascicle muscle territory, is present in the gracilis muscle. Microneurovascular techniques have improved the reliability of muscle transplantation for the reconstruction of facial paralysis. The amount of movement obtained depends on many factors, including the amount of muscle transplanted and the adequacy of its reinnervation. The ability to transplant a small segment of a muscle based on the fascicular territory enables the surgeon to supply the amount of movement that each individual patient requires.     

Suppression of coral populations by filamentous algae within damselfish territories

Transplanted pieces of the coral Acropora palifera Lamarck simulating colonies 2–4 years old were adversely affected by dense algal-sediment mats developed in response to reduced grazing pressures within territories of the damselfish Dischistodus perspicillatus (Cuvier). Coral growth was initially prolonged in autumn, but was subsequently suppressed during winter and spring. There was heavy mortality only in mid-winter when corals died rapidly from the base. It is suggested that the algal mat caused the corals to expend energy to keep their surfaces clean, so that death followed the exhaustion of metabolic reserves during periods of minimum nutrient availability, and that the abundance of D. perspicillatus is sufficient to explain the paucity of small Acropora palifera colonies in an otherwise favourable habitat.     

Nest guarding in a damselfish: evidence of a role for parasites

A potential effect of parasitism on male egg loss while guarding was tested using damselfish Stegastes planifrons . Despite an apparent low energetic cost of the parasite on the male condition, there was a significant positive relationship between egg loss and parasite load. These data provide the first evidence that parasites could play a role in male guarding behaviour. The effect of the parasite could have resulted in low male condition compensated by an increase in filial cannibalism or in a lesser ability to protect the nest from predators.     

Size-dependent gender modification in a hermaphroditic perennial herb

The size-advantage model predicts that hermaphroditic organisms adjust sex allocation depending on their resource status. We investigated the relationship between size and sex allocation in the co-sexual perennial herbs Trillium erectum and Trillium grandiflorum at two sites in southern Ontario, Canada by measuring pollen and ovule production and biomass allocation at flowering and fruiting. In both species, there was a strong relationship between size and gender; larger plants allocated proportionately more biomass to female reproduction and produced fewer pollen grains relative to ovules than smaller plants. Variation in gender was better explained by size than age, although age and size were correlated. While the relationship between size and gender was similar between species, T. erectum allocated proportionately more to female reproduction than T. grandiflorum, independent of size. In the absence of pollen limitation, there was no evidence of secondary adjustment of gender at fruiting. The results are discussed in the context of models predicting size-dependent gender modification in animal-pollinated plants. Evidence about the pollination and seed dispersal biology of Trillium spp. suggests that the relative effects of local mate and resource competition may be important in driving size-dependent sex allocation in these species.     

Interspecific competition and territory defense mechanisms of Coptotermes formosanus and Coptotermes gestroi (Isoptera: Rhinotermitidae)

Coptotermes formosanus Shiraki and C. gestroi (Wasmann) are the most widely distributed species of the genus and occur sympatrically in the subtropics. Results of two bioassays in the current study showed that C. gestroi was more aggressive than C. formosanus. In the petri-dish bioassays, C. gestroi won most of the agonistic encounters over C. formosanus. In the two-dimensional foraging arena bioassays, over 73% tunnel interceptions observed in the 18 replications were caused by progressing tunnels of C. gestroi encountering the tunnels of C. formosanus. Tunnel interception of the two species resulted in minor agonistic interactions. Both species quickly buried the connected tunnel at multiple locations. Termite cadavers resulting from agonistic behavior appeared to have induced sand deposition that resulted in tunnel blockages and deterred reopening of these blockages. Sealing individual tunnels in response to encounters with other species acts to prevent further agonism and mortality, and on a broad scale, the aggregate of such blocked tunnels may come to define the borders between adjacent colonies.     

Versatile habitat conditioning by damselfish cultivating turf algae on giant clams

In this paper, a monitoring and modelling concept for ecological optimized harbour dredging and fine sediment disposal in large rivers is presented. According to the concept, first a preliminary assessment should be performed previous to the dredging and dumping procedure to derive knowledge about the current status in hydrodynamics, morphology and instream habitat quality. During the performance of the maintenance work, a high-resolution monitoring program has to be organized to measure flow velocities, the suspended sediment concentrations and the extent of the occurring plume. These data can then be compared with natural suspended sediment conditions and serve as input data for numerical sediment transport modelling. Furthermore, bathymetric surveys and biotic sampling enable the detection of possible effects of dredging and disposal in the post-dumping stage. Based on sediment transport modelling approaches, short- to mid-term developments of the sediment plume can be predicted with an additional and final habitat evaluation at the end of the project. This concept was applied and optimized during the maintenance work at the case study winter harbour Linz at the Danube River. The findings of the presented study highlight the necessity of integrated monitoring and modelling approaches for harbour dredging especially in large river systems.     

Short-term changes in plasma levels of hormones during establishment and defense of a breeding territory in male song sparrows, Melospiza melodia

When territorial male song sparrows are captured and removed from their territories, previously unmated and nonterritorial males will take over those vacant territories within 12-72 hr. Plasma levels of testosterone are elevated in these replacement males as well as in their neighbors. Since the latter already have territories, it is suggested that the agonistic interactions over territory boundaries, or behavioral stimuli from challenging males, rather than ownership of a territory per se, stimulates secretion of testosterone. To test this hypothesis further, male song sparrows were challenged by experimental simulation of a territorial intrusion. This procedure involved placing a caged male song sparrow in the center of the subject's territory and playing tape recorded conspecific song through an adjacent speaker. Responding males were then captured at intervals after onset of the intrusion. Plasma levels of testosterone were significantly higher in males exposed to experimental territorial intrusion than they were in controls, supporting the hypothesis that behavioral stimuli emanating from an intruding male can act as supplementary information stimulating secretion of testosterone.     

Recruiting juvenile damselfish: the process of recruiting into adult colonies in the damselfish Stegastes nigricans

Juveniles of Stegastes nigricans occur in adult colonies, solitarily, and occasionally in juvenile colonies. We concentrated on solitary juveniles and those in adult colonies. We examined the costs and benefits of different settlement strategies, quantified the territory requirements of adults, and investigated the process of how juveniles make the transition to adult territorial fish. An adequate adult territory lies next to those of other adults, is proportional in area to the size of the adult, and contains a refuge tunnel whose entrance is sufficiently large. Compared with solitary juveniles, those <4 cm total length inhabiting adult colonies experienced reduced heterospecific competition for algal food and consequently benefited from a greater density of algae. A cost of recruiting into an adult colony, however, was increased attacks by adults. Juveniles that settled in adult colonies avoided attacks by retreating into small holes inaccessible to adults. As juveniles in adult colonies grew, they were chased less often by adults, whereas they themselves chased adults and heterospecific fish more often. Because territory size correlated with fish size in adult colonies, its area had to expand as the young fish grew, and that expansion was done at the expense of neighbors. Obtaining the space needed by an adult may be possible only when the juvenile settles directly into an adult colony. Juveniles that first settle down solitarily, or in juvenile colonies, may later attempt to enter adult colonies. However, because they do so as larger juveniles, they would have difficulty insinuating themselves into small refuges, which is essential for retreat from the adults. Received in revised form: 4 January 2001 Electronic Publication