Threat Sensitivity in Bicolor Damselfish: Effects of Sociality and Body Size

Threat sensitivity involves graded antipredator responses that reflect the degree of predatory threat encountered. Models of predatory Atlantic trumpetfish (Aulostomus maculatus) were presented to differently sized juvenile bicolor damselfish (Pomacentrus partitus) that occurred singly or in small groups to test for threat sensitivity. Bicolor damselfish displayed threat sensitivity by responding more strongly: 1. To large models than to small models; and 2. As models were brought closer. Solitary damselfish showed stronger responses than did fish in small groups. However, no habituation to threat occurred, no correlation was found between response strength and fish size, and many individuals gave strong, ungraded responses to predator models. Comparisons between juvenile bicolor damselfish and threespot damselfish suggest that bicolor are weakly threat sensitive in that they respond strongly regardless of threat (hypersensitivity). Solitary fish are more hypersensitive than grouped fish. Threespot damselfish show a stronger influence of damselfish body size: small threespot exhibit graded responses in proportion to degree of threat (pure threat sensitivity), whereas large threespot show relatively weak responses (nonchalance). These interspecific comparisons indicate the potential range that characterizes threat sensitivity and the possible influence of social and ontogenetic factors on that range.     

Do cleaning stations affect the distribution of territorial reef fishes?

We investigated the role of cleaning stations in determining the distribution of territorial reef species. Cleaner fish reduce their clients' ectoparasite loads and, therefore, proximity to cleaning stations should be advantageous for territorial fish. We focused on five damselfish species which hold permanent territories and cleaning stations occupied by cleaning gobies (Elacatinus spp.) on a Caribbean reef. Contrary to our predictions of higher densities near cleaning stations, we found that bicolor damselfish were less abundant near cleaning stations than at ecologically similar points without cleaning gobies whereas no effects were seen for longfin, dusky, yellowtail, and threespot damselfish. In addition, although damselfish densities were higher in the immediate vicinity of cleaning stations than 1.5-3 m away for most species, this was also the case at points without cleaners. Because cleaning stations are usually located on prominent coral heads or sponges, the overall significant attraction of damselfish to such structures, whether occupied by cleaning gobies or not, could reflect attraction to past or potential cleaning stations. However, it is more likely that interspecific competition and/or the low benefits of being cleaned at our study site prevent aggregation around cleaners. Cleaning stations may play only a minor role in determining the distribution of territorial reef fishes.     

Effects of the Caribbean threespot damselfish,Stegastes Planifrons (Cuvier), on algal lawn composition

Algal lawn communities within territories of herbivorous threespot damselfish, Stegastes planifrons (Cuvier) were studied in a shallow back-reef environment at La Parguera, Puerto Rico. Caging exclusion experiments, monitoring of non-manipulated lawns and gut content analyses were used to determine the effect of S. planifrons on algal species composition of the lawn. An average of 40 algal species per sampling period were found in naturally occuring lawns, with a total of 53 species being recorded throughout the course of study. Examination of algal species found in damselfish guts and comparison with natural lawns indicate that damselfish feed primarily on their algal lawns and graze the algae present in proportion to abundance. Caged algal lawns showed significant decline in algal species number and abundance over time. Non-selective feeding by Stegastes planifrons within their territorial lawns appears to result in an “intermediate disturbance” which maintains a greater algal species richness than is found in their absence.     

Juvenile settlement,survivorship and in situ growth rates of four species of Caribbean damselfishes in the genusStegastes

Synopsis FourStegastes species were visually censused to determine seasonality and size of juvelines at settlement as well as their survival and growth rates. The census was performed weekly from September 1987 to March 1991 in southwest Puerto Rico. All species had a settlement pulse during September–October. The threespot damselfish,S. planifrons, appeared to have the strongest site fidelity, longest survivorship and slowest growth of the four species. The cocoa damselfish,S. variabilis, and the bicolor damselfish,S. partitus, had similar patterns of site fidelity, life expectancy and growth rates, but these two species differed in being the most and least abundant, respectively. The beaugregory,S. leucostictus, occurred second in abundance but included the least number of new settlers. This species displayed poor site fidelity, a characteristic which probably influenced calculations indicating poor survivorship, life expectancy and growth relative to its congeners. Similarities and differences in these characteristics, as well as in the habitat preferences displayed by these four fishes, may contribute to their abilities to coexist in the same area.     

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.     

Territorial vocalization in sympatric damselfish: acoustic characteristics and intruder discrimination

Damselfishes are well known for their aggressive, territorial behaviour during which the use of vocalization behaviour has been well documented. However, agonistic acoustic signalling has been understudied in particular when the vocalizations are interspecific. In this study, we characterize and compare the previously undescribed vocalization behaviour of longfin damselfish (Stegastes diencaeus), in an agonistic context, with the closely related and sympatric dusky damselfish (Stegastes adustus). Next, we examined if these congeneric species modulate their vocalizations in a similar pattern to previously described aggressive behaviour patterns. Audio field recordings of territorial males were obtained in response to three separate stimuli: (1) conspecific male damselfish, (2) heterospecific male damselfish and (3) a common intruder, the slippery dick wrasse (Halichoeres bivittatus). The vocal repertoires of both longfin and dusky damselfish comprised the same three distinct call types: chirps, pops and pulse trains. However, temporal measures of the calls showed significant differences between species. Additionally, dusky damselfish were more vocal overall, producing more calls and spending more time calling than longfin damselfish. These responses were stimulus and species dependent, as the two species modulated acoustic response by modulating pulse number based on intruder species. These results suggest that these closely related species of damselfish use vocalization behaviours that are both unique and context dependent.     

Response to underwater laser pointer in the Orange-finned Anemonefish Amphiprion chrysopterus and three-spot damselfish Dascyllus trimaculatus

Response of orange-finned anemonefish Amphiprion chrysopterus and three-spot damselfish Dascyllus trimaculatus to red laser-pointer light was studied in Mo'orea, French Polynesia. Four magnificent anemones Heteractis magnifica and their resident fish were observed for typical behaviours (biting, chasing, hiding, posing, lunging and retreating) with and without exposure to laser-pointer light. Lunging behaviour increased significantly for both fish species upon exposure to laser-pointer light; none of the other behaviours changed significantly. We advance the hypothesis that orange-finned anemonefish and three-spot damselfish interpret laser pointer stimulation as a territorial threat.     

Parental defence on the reef: antipredator tactics of coral‐reef fishes against egg‐eating seasnakes

On coral reefs in New Caledonia, the eggs of demersal‐spawning fishes are consumed by turtle‐headed seasnakes (Emydocephalus annulatus). Fish repel nest‐raiding snakes by a series of tactics. We recorded 232 cases (involving 22 fish species) of antipredator behaviour towards snakes on a reef near Noumea. Blennies and gobies focused their attacks on snakes entering their nests, whereas damselfish (Pomacentridae) attacked passing snakes, as well as nest‐raiders (reflecting territorial defence). Biting the snake was the most common form of attack, although damselfish and blennies also slapped snakes with the tail, or (blennies only) plugged the nest entrance with the parent fish's body. Gobies rarely defended the nest, although they sometimes bit or threw sand at the snake. A snake was more likely to flee if it was attacked before it began feeding rather than after it found the eggs (82% versus 3% repelled) and if bitten on the head rather than the body (68% versus 53%). Tail‐slaps were not effective, although plugging the burrow and throwing sand often caused snakes to flee. These strong patterns reflect phylogenetic variation in fish behaviour (e.g. damselfish detect a snake approach sooner than do substrate‐dwelling blennies and gobies) coupled with intraspecific variation in snake diets. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 415–425.     

Interference behavior and ecology of threespot damselfish (Eupomacentrus planifrons)

Summary Interference behavior by threespot damselfish directed at the sea urchins, Diadema antillarum and Echinometra viridis, differed in levels of aggression and discrimination. Higher aggression towards and recognition of Diadema by threespots was correlated with the distribution of Diadema along edges of coral patches. Lower aggression levels combined with lower recognition levels of Echinometra were correlated with a distribution of Echinometra closer to damselfish algal lawns. Differences in behavior of the urchins stimulated differing levels of aggression by the damselfish directly affecting the distribution of the urchins in the back-reef environment.This is contribution number 167 from the Discovery Bay Marine Laboratory of the University of the West Indies, Jamaica     

Factors Affecting Interspecific Aggression in a Year‐Round Territorial Species,the Jewel Damselfish

Vertebrates live in complex species networks in which interspecific interactions are common. In some contexts, the aggressive behaviours shown in these interspecific interactions are very similar to those shown in intraspecific interactions. It is still an open question whether intra‐ and interspecific aggression share common causality. We studied a year‐round territorial species the jewel damselfish, (Plectroglyphidodon lacrymatus), which cultivate algae they feed on. Territory holders aggressively defend these algae that are an attractive resource for many other species. In this study, we recorded territorial aggression in free‐living individuals and recorded aggressive responses to a standardized territorial intrusion test in captive individuals. Field observations indicated that territorial aggression was selectively targeted towards food competitors. Independent of the size of the species, aggression was more frequent towards common species around their territories. This relationship was confirmed experimentally by confronting the jewel damselfish with novel objects to which the subjects were exposed either frequently or rarely. We suggest that jewel damselfish have to learn which species are competitors and therefore should be chased. In a standardized intrusion test with captive individuals, no significant differences were found in territorial responses towards intra‐ or interspecific intruders. Neither territorial aggression nor the intrusion showed any relationship with plasma androgen levels. Together, these data suggest that experience might be more important in non‐seasonal territorial aggression than circulating hormonal factors.     

Ichthyotoxicity of Chattonella marina (Raphidophyceae) to damselfish (Acanthochromis polycanthus): the synergistic role of reactive oxygen species and free fatty acids

This investigation aimed to elucidate the relative roles of putative brevetoxins, reactive oxygen species and free fatty acids as the toxic principle of the raphidophyte Chattonella marina, using damselfish as the bioassay. Our investigations on Australian C. marina demonstrated an absence or only very low concentrations of brevetoxin-like compounds by radio-receptor binding assay and liquid chromatography–mass spectroscopy techniques. Chattonella is unique in its ability to produce levels of reactive oxygen species 100 times higher than most other algal species. However, high levels of superoxide on their own were found not to cause fish mortalities. Lipid analysis revealed this raphidophyte to contain high concentrations of the polyunsaturated fatty acid eicosapentaenoic acid (EPA; 18–23% of fatty acids), which has demonstrated toxic properties to marine organisms. Using damselfish as a model organism, we demonstrated that the free fatty acid (FFA) form of EPA produced a mortality and fish behavioural response similar to fish exposed to C. marina cells. This effect was not apparent when fish were exposed to other lipid fractions including a triglyceride containing fish oil, docosahexaenoate-enriched ethyl ester, or pure brevetoxin standards. The presence of superoxide together with low concentrations of EPA accelerated fish mortality rate threefold. We conclude that the enhancement of ichthyotoxicity of EPA in the presence of superoxide can account for the high C. marina fish killing potential.     

The effect of territorial damselfish (family Pomacentridae) on the space use and behaviour of the coral reef fish, Halichoeres bivittatus (Bloch, 1791) (family Labridae)

Most demersal species of damselfish (family Pomacentridae) are territorial herbivores that aggressively chase other fishes away from their nests. This study investigates whether the aggressive territorial damselfish, Stegastes leucostictus, modifies behaviours and home range area use in a less aggressive, non-territorial species, the slippery dick wrasse, Halichoeres bivittatus (family Labridae). Damselfishes and wrasses are ubiquitous and abundant members of coral reef fish assemblages around the world; hence, this study has broader implications beyond the two Caribbean species chosen for this study. A manipulative field experiment consisted of transplanting one or three S. leucostictus into artificial shelters positioned within adult or adjacent to juvenile H. bivittatus home ranges. The introduction of damselfish had little effect on the size of home range areas of juvenile or adult wrasses, but had a significant effect on the location of their home ranges. The damselfish also affected adult microhabitat use but did not affect use by juveniles. In addition, there was an increase in damselfish-wrasse interactions that resulted in a significant reduction in the amount of time adult wrasses could spend foraging or in a foraging assemblage. It was concluded that aggressive interactions with territorial damselfish affect individual H. bivittatus space use and may consequently affect the spatial distribution of H. bivittatus populations.     

The social organization of adult blue tangs, <Emphasis Type="Italic">Acanthurus coeruleus</Emphasis>, on a fringing reef,Barbados, West Indies

Blue tangs in Barbados exhibit three distinct social modes: territorial, schooling and wandering. We compared the mobility, foraging, aggression performed and received and the use of cleaning stations of adult blue tangs among modes and among habitats within a single fringing reef in Barbados. Evidence from observed switches during focal observations and multiple observations of tagged individuals indicate that fish are either territorial or non-territorial. Non-territorial fish formed schools and wandered. However, wandering can be used during solitary movements by fish in either type. Fish in the territorial mode, not previously described in adults of this species, restricted activity to a small area overlapping the territories of other tangs. They actively chased conspecifics and were chased mostly by damselfishes. They swam more slowly and fed at higher rates than other modes. Fish in the schooling mode ranged widely in compact, polarized groups of conspecifics, congeners and other species. They were not aggressive and were attacked mostly by damselfishes. They swam rapidly and fed at intermediate rates. Fish in the wanderer mode showed neither aggression nor association with other individuals. They swam rapidly, well above the substrate, fed little, were chased by conspecifics, ocean surgeonfish, A. bahianus, and damselfish and visited cleaning stations more often then other modes. All three modes were observed in all four main zones of the reef, and their behaviour changed quantitatively with habitat type. We suggest that territoriality reduces competition for algal food, schooling allows fish to overcome the food defence by damselfish, tangs and ocean surgeonfish, and wandering permits solitary movement over the reef to cleaning stations, feeding sites and other resources.     

Short-term predator avoidance behavior by invasive and native amphipods in the Great Lakes

Understanding predator avoidance behavior by prey remains an important topic in community and invasion ecology. Recently, the Ponto-Caspian amphipod Echinogammarus ischnus (Stebbing 1898) was accidentally introduced into the Great Lakes. Since its introduction, it has displaced the native amphipod, Gammarus fasciatus (Say 1818), from several locations in the lower lakes. To assess whether behavioral differences in predator avoidance might be a causal mechanism increasing the success of the invasive amphipods, two experiments were conducted examining (1) native and invasive amphipod behavioral responses to five fish species with different foraging behaviors, and (2) amphipod responses to different densities of round gobies, a hyper-abundant benthic invertivore. Echinogammarus reduced its distance moved in the presence of all fish species tested, whereas Gammarus reduced its distance moved only after exposure to round gobies, black crappies, and rainbow darters. Both amphipod species increased the time spent motionless following exposure to round gobies, but not after encountering the scent of most of the remaining fish predators. The exception was that Echinogammarus also responded to black crappie scent whereas Gammarus did not. Although both amphipod species exhibited behavioral responses to many of the fish predators, the magnitude of their responses differed only after exposure to the brown bullhead. In the bullhead trials, Echinogammarus reduced its distance traveled significantly more than Gammarus. Both amphipod species increased their avoidance response to increasing goby density, however, the pattern of avoidance behavior was different. Invasive E. ischnus exhibited a consistently strong avoidance response to round gobies over the test duration. Native G. fasciatus initially avoided goby scent, but then either ceased their avoidance response or showed a hyper-avoidance response, depending on goby density. These results suggested (1) both species of amphipods were able to differentiate and react to a variety of fish predators, (2) invasive Echinogammarus amphipods avoided a larger range of fish predators than the native Gammarus, (3) increased avoidance behavior was associated with an increased density of fish, and (4) the avoidance response patterns of invasive Echinogammarus when faced with round goby predators might lead to increased predation on native Gammarus in habitats where they co-occur.     

An experimental removal of a territorial pomacentrid: effects on the occurrence and behavior of competitors

Synopsis Stegastes fasciolatus is the most common territorial damselfish in the shallow waters of Hawaii. Territorial defense was observed against other herbivorous fishes, especially acanthorids, scarids and one omnivorous chaetodontid. One acanthurid,Acanthurus nigrofuscus was found to differ in abundance and social behavior in areas whereS. fasciolatus was present, compared to areas where it was absent. The chaetodontid,Chaetodon quadrimaculatus was sheltered during the day in areas where the pomacentrid was abundant, apparently feeding at night. In other areas it fed during the day and at night, depending on the phase of the moon.S. fasciolatus were then experimentally removed from one study site, to test whether the differences in abundance and behavior of the other species were due to the presence of the damselfish. There was a significant increase in numbers of the surgeonfishAcanthurus nigrofuscus in the removal area, as well as changes in social behavior from schooling to defense of small territories. The butterflyfish,C. quadrimaculatus, was observed to forage during the day in the removal area. There were no significant changes in the control sites. The presence of the interspecifically territorial damselfish,S. fasciolatus, thus appears to be an important determinant of the behavior of these potential food competitors.     

Diurnal and seasonal changes in the mate-locating behavior of the satyrine butterfly <Emphasis Type="Italic">Lethe diana</Emphasis>

The time schedule of mate-locating behavior in the male satyrine butterfly Lethe diana was studied. Males displayed territorial behavior only in the afternoon, but it was not known whether they attempted to locate mates in the morning. In order for males of most butterfly species to mate with a virgin female, it would be most effective to initiate mate-locating behavior in the morning, when synchronous eclosions occur. Males of L.diana flew patrols in the morning and displayed territorial behavior in the afternoon. The cost of territorial behavior is probably less than that of patrolling, as territorial flights are shorter than those when patrolling. The temporal switching of mate-locating behavior is theoretically favored, since the reproductive value of females decreases later in the day, when males perform the less costly mate-locating behavior. Thus, the diurnal switching of male mate-locating behavior may be an adaptive strategy in response to the rapid decline in the reproductive value of females over the course of a day.     

El Niño mass coral mortality: a test of resource limitation in a coral reef damselfish population

Summary The mass mortality of reef corals in the eastern Pacific as a result of the 1982–1983 El Niño oceanographic anomaly permitted the first large scale test of resource limitation for a coral reef fish. Population densities of territorial herbivorous damselfish did not respond to the massive regional increase in space available for the cultivation of algal food following the El Niño event. The proportion of juveniles in the population was low and new recruits were uncommon, indicating that recruitment rates rather than resource supply probably control the abundance of this coral reef damselfish.     

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.     

Effects of group behavior in the predatory raid on damselfish nests by the false cleanerfish Aspidontus taeniatus

The false cleanerfish Aspidontus taeniatus, which resembles the bluestreak cleaner wrasse Labroides dimidiatus, is one of the best-known examples of mimicry in vertebrates. This mimicry system has been viewed as an aggressive mimicry to bite fish fins. However, recent field studies have reported that large individuals of the false cleanerfish often form groups and jointly raid damselfish nests to eat eggs that are guarded by their parents. The benefits of group behavior have been reported in a variety of animals. In the case of false cleanerfish, parental defense of territorial damselfishes is the main factor that constrains the availability of nutritionally valuable food resources. Here, we conducted field observations on the reefs of Okinawa, and found that the false cleanerfish formed groups of 2–12 individuals when they raided breeding nests of 13 species of damselfishes (Pomacentridae) and one species of triggerfish (Balistidae). Since the cleaner wrasse does not form such groups, the feeding groups of the false cleanerfish are assumed to reduce the effectiveness of mimicry. However, our results showed that the group behavior has two effects: a dilution effect, which reduces the risk of being attacked by egg-guarding fish, and an increase in foraging efficiency. We conclude that the false cleanerfish need to form foraging groups during egg-eating because the mimicry has no effect on parental damselfishes.     

Positive interactions between corals and damselfish increase coral resistance to temperature stress

By the century's end, many tropical seas will reach temperatures exceeding most coral species' thermal tolerance on an annual basis. The persistence of corals in these regions will, therefore, depend on their abilities to tolerate recurrent thermal stress. Although ecologists have long recognized that positive interspecific interactions can ameliorate environmental stress to expand the realized niche of plants and animals, coral bleaching studies have largely overlooked how interactions with community members outside of the coral holobiont shape the bleaching response. Here, we subjected a common coral, Pocillopora grandis, to 10 days of thermal stress in aquaria with and without the damselfish Dascyllus flavicaudus (yellowtail dascyllus), which commonly shelter within these corals, to examine how interactions with damselfish impacted coral thermal tolerance. Corals often benefit from nutrients excreted by animals they interact with and prior to thermal stress, corals grown with damselfish showed improved photophysiology (F_v/F_m) and developed larger endosymbiont populations. When exposed to thermal stress, corals with fish performed as well as control corals maintained at ambient temperatures without fish. In contrast, corals exposed to thermal stress without fish experienced photophysiological impairment, a more than 50% decline in endosymbiont density, and a 36% decrease in tissue protein content. At the end of the experiment, thermal stress caused average calcification rates to decrease by over 80% when damselfish were absent but increase nearly 25% when damselfish were present. Our study indicates that damselfish-derived nutrients can increase coral thermal tolerance and are consistent with the Stress Gradient Hypothesis, which predicts that positive interactions become increasingly important for structuring communities as environmental stress increases. Because warming of just a few degrees can exceed corals' temperature tolerance to trigger bleaching and mortality, positive interactions could play a critical role in maintaining some coral species in warming regions until climate change is aggressively addressed.