Coexistence: Symbiotic sharing of feeding territories and algal food by some coral reef fishes from the Western Indian Ocean

A symbiotic relationship between a damselfish (Stegastes fasciolatus) and 2 surgeonfishes (Acanthurus lineatus and A. leucosternon) is described. The damselfish, which is about 1/10 the size of the surgeonfishes, is concentrated in, and appears to prefer to be in, the feeding areas of the surgeonfishes. There is an average of one adult damselfish per adult surgeonfish feeding area. All 3 species defend their feeding areas against conspecifics and various other fishes that have similar diets, but there are few aggressive interactions between the cohabitants. Most of the interspecific defense of cohabited areas is provided by the surgeonfishes. The cohabitant species eat the same types of benthic microalgae, but only the damselfish eats microcrustaceans. It appears that the net cost to a surgeonfish of having a damselfish in its feeding area is very low because (1) biomass density of the damselfish is low, (2) it uses some food that the surgeonfishes do not use, and (3) it makes a small contribution to the defense of shared feeding areas. Even if there is a cost to the surgeonfish, the small size of the damselfish would allow it to take shelter from the surgeonfish, thus making it too costly for the surgeonfish to exclude it. Such cohabitation relationships, which may be quite common among reef fishes, represent a means by which the coexistence of species that use the same limiting resources is achieved.     

Foraging activity and resource use by three parrotfish species at Fernando de Noronha Archipelago, tropical West Atlantic

Parrotfishes are important components of the herbivore and detritivore guilds of tropical and subtropical reefs. Most of parrotfish species are protogynous hermaphrodites that change colour and sex, from initial phase females or males (IP) to terminal phase males (TP). We studied the foraging behaviour of Sparisoma amplum, S. axillare and S. frondosum, three syntopic scarids on the rocky reefs of Fernando de Noronha Archipelago, Tropical West Atlantic. The three parrotfish species differed in food selection and preference, but IP and TP individuals of the same species preferred the same food types, except for S. amplum. Feeding rates of IP individuals were higher than those of TP individuals, but the distribution of feeding frequencies throughout the day of IP and TP individuals of the same species was similar. IP individuals had higher feeding rates than TP ones, which seems related to the fact that TP individuals spend a large amount of time patrolling their territories and chasing away conspecific individuals at the study site. The general foraging pattern we found for S. amplum, S. axillare and S. frondosum is similar to patterns found for other parrotfish species in the Western Atlantic.     

The costs and benefits of territory sharing for the Caribbean coral reef fish,Scarus iserti

Summary Along the Caribbean coast of Panama, groups of unrelated female striped parrotfish, Scarus iserti, co-defend a common feeding territory. Field manipulations of group size and composition were performed to examine the benefits and costs accrued by dominant fish within these shared territories. Dominant fish benefit from the presence of relatively large subordinates because they share in the defense of the territory. Removals of these fish caused increases in defense time and decreases in feeding time for dominant group members. Dominants benefit from the presence of small subordinates because they increase the foraging efficiency of dominants. Removals of smaller subordinates caused reductions in the feeding time of dominant fish, although no changes in defense time occurred. Concurrently, dominant fish reduce costs of resource depletion by displacing subordinate group members from good food patches. Dominance interactions within a group reduce the amount of time subordinates spend feeding (subordinate individuals fed at higher rates following the removal of a dominant) and limit a subordinate's access to high quality resources. This combination of benefits and reduced costs ultimately contributes to the economic defensibility of a striped parrotfish territory and has led to the evolution of group territorial behavior in the absence of kin selection and cooperative parental care.     

Genetic connectivity and historical demography of the blue barred parrotfish (Scarus ghobban) in the western Indian Ocean

Studies on genetic connectivity are essential for the design of management strategies for coral reef fisheries. In this study we used a mitochondrial DNA marker to investigate population structure of the reef-associated parrotfish, Scarus ghobban, from four countries, Kenya, Mauritius, Seychelles and Tanzania, in the western Indian Ocean. We obtained nucleotide sequences of the mitochondrial control region for 117 individuals. Measures of haplotype diversity were relatively high. Pairwise population differentiation (F _ST) was low, but not always non-significant. Analysis of molecular variance (AMOVA) showed genetic differentiation between groups, when the data was partitioned into two groups consisting of samples from Mauritius and Tanzania in one group, and samples from Kenya and Seychelles in another group. Direction of gene flow was estimated using a Bayesian approach. Migration was sometimes asymmetric or directional, coinciding with the flow of major oceanic and coastal currents in the region. Mismatch distributions, based on the observed number of differences among haplotype pairs, produced a unimodal distribution, indicative of recent demographic expansion. Phylogenetic analyses revealed three clades without any geographic structure, suggesting recent migration between historically isolated lineages. We reconstructed the historical demography of S. ghobban and examined it in the context of Pleistocene climate stages and changes in relative sea level. Overall, these results showed that populations of S. ghobban are genetically diverse and have relatively high gene flow, with some genetic structuring in the western Indian Ocean.     

Vigilance of mustached tamarins in single-species and mixed-species groups—the influence of group composition

Species that participate in mixed-species groups (MSG) may have complementary roles in antipredator strategies. We studied vigilance in mustached tamarins (Saguinus mystax), small arboreal primates that form stable mixed-species groups with saddleback tamarins (Saguinus fuscicollis), in order to examine how the direction of vigilance changes with different species group compositions and whether the division of labor between the two species can be confirmed. We did so by comparing quantitative and qualitative differences in vigilance behavior between same individuals in and out of association (case A); MSG and single-species groups of the same total group size from two different populations (case B); and MSG of the same group size but with a different ratio of conspecifics to heterospecifics (case C). We predicted that individuals would increase downward scanning when heterospecifics are absent or their percentage is low, but total vigilance would increase only in case A due to the group size effect. However, mustached tamarins increased total vigilance due to horizontal scanning in cases A and C, and the predictions were confirmed only in small-sized groups in case B. Thus, we found indications that associating tamarin species in MSG might complement each other in the direction of vigilance, but the division of labor alone does not satisfactorily explain all the findings. There appear to be other mechanisms at work that define how direction of vigilance changes with group size and species composition. Complementarity of species probably occurs due to species vertical stratification rather than differences in the direction of vigilance.     

A new apparatus for force measurement in marine bioadhesion

In an experimental study on the effect of parrotfish (probably Scarus taeniurus) grazing on the structure of benthic reef communities, fishes in densities of 0.6 to 1.5 parrotfish per m² or 9 to 17 g wet weight of fish per m² of feeding surface were found to have an optimum effect, resulting in the greatest benthic species richness and biomass on 2-dimensional surfaces. The presence of refuges (3-dimensional habitats), however, has a greater impact on bemthic community structure (number of species and biomass) than does just the density of parrotfish in such an experimental system. Coral recruitment is enhanced by the presence of refuges and, like coralline algae, is more successful under increased grazing pressure. These optimum densities of parrotfishes relate well to observed field densities where, in a collection from a Hawaiian patch reef, there were 1.1 fish or 10.8 g wet weight of parrotfish per square meter of collection area. The success of coralline algae and corals under high grazing pressure may have important consequences for the stability and structure of modern coral reefs.     

Impacts of sublethal lead exposure on the hemato-immunological parameters in tilapia (Oreochromis mossambicus)

The effects of exposure of tilapia (Oreochromis mossambicus) to sublethal concentrations of lead on hematological and immunological parameters were investigated. The fish were semistatically exposed to low (L) (0.5 mg L^?1), medium (M) (2.5 mg L^?1), and high (H) (5 mg L^?1) lead concentrations for 14 days. At the end of the trial, red blood cell (RBC) count, hematocrit (Hct) ratio and hemoglobin levels showed significant decreases in M and H groups, while RBC was significantly increased in L group compared to control group. Lysozyme activity showed a decrease in all groups on day 14 compared to control. A decrease in myeloperoxidase activity was seen in M and H groups on day 7, and in all treatment groups on day 14. In conclusion, it was found that exposure of O. mossambicus to lead concentrations affected hematological and immunogical parameters adversely.     

Schooling tendency and foraging benefit in the ocean surgeonfish

Summary Field observations of individuallytagged ocean surgeonfish (Acanthurus bahianus) indicated that schooling provided a foraging benefit: an increase in relative forging time. The magnitude of this increase differed among individuals, primarily due to time budgets of the fish when they were not in schools. The proportion of nonschooling time devoted to foraging was positively correlated with fish size, while the proportion of schooling time that was spent forging was similar for all individuals. Schooling tendency (percentage of time in schools) was positively related to magnitude of foraging benefit and negatively related to fish size. When fish size was held constant by partial correlation, schooling tendency remained positively related to foraging benefit. These results provide evidence that fish which derive a greater foraging benefit from school membership spend more time in schools.     

Feeding ecology and niche segregation in oceanic top predators off eastern Australia

We examined the feeding ecology and niche segregation of the ten most abundant fish species caught by longline operations off eastern Australia between 1992 and 2006. Diets of 3,562 individuals were examined. Hook timer data were collected from a further 328 fish to examine feeding behaviour in relation to depth and time of day. Prey biomass was significantly related to predator species, predator length and year and latitude of capture. Although the fish examined fed on a mix of fish, squid and crustacea, fish dominated the diet of all species except small albacore (Thunnus alalunga) which fed mainly on crustacea and large swordfish (Xiphias gladius) and albacore which fed mainly on squid. Cannibalism was observed in lancetfish (Alepisaurus spp.). Multidimensional scaling identified three species groups based on their diet composition. One group consisted of yellowfin tuna (T. albacares), striped marlin (Tetrapturus audax) and dolphinfish (Coryphaena hippurus); a second group consisted of bigeye tuna (T. obesus), swordfish and albacore; and a third consisted of southern bluefin tuna (T. maccoyii) and blue shark (Prionace glauca). Of note was the separation of mako shark (Isurus oxyrhynchus) and lancetfish from all other predators. Prey length generally increased with increasing predator length although even large predators fed on a wide range of prey lengths including very small prey. Overall, differences in prey type and size, feeding times and depths were noted across the range of species examined to the extent that predators with overlapping prey, either in type or size, fed at different times of the diel period or at different depths. Taken together these data provide evidence for feeding niche segregation across the range of oceanic top predators examined.     

Group size and foraging efficiency in yellow baboons

Summary I studied the foraging behaviour of adults in three different-sized groups of yellow baboons (Papio cynocephalus) at Amboseli National Park in Kenya to assess the relationship between group size and foraging efficiency in this species. Study groups ranged in size from 8 to 44 members; within each group, I collected feeding data for the dominant adult male, the highest ranking pregnant female, and the highest ranking female with a young infant. There were no significant differences between groups during the study in either the mean estimated energy value of the food ingested per day for each individual (385±27 kJ kg^-1 day^-1) or in the estimated energy expended to obtain that food (114±3 kJ kg^-1 day^-1). Mean foraging efficiency ratios, which reflect net energy gain per unit of foraging time, also did not vary as a function of the size of the group in which the baboons were living. There was substantial variation between days in the efficiency ratios of all animals; this was the result of large differences in energy intake rather than in the energy expended during foraging itself. The members of the smallest group spent on the average only one-half as much time feeding each day as did individuals in the two larger groups. However, they obtained almost as much energy while foraging, primarily because their rate of food intake while actually eating tended to be higher than the rate in the other groups. The baboons in the small group were observed closer to trees that they could climb to escape ground predators, and they also were more likely to sit in locations elevated above the ground while resting. Such differences would be expected if the members of the small group were less able to detect approaching predators than individuals that lived in the larger groups. The results of this study suggest that predator detection or avoidance, rather than increased foraging efficiency, may be the primary benefit of living in larger groups in this population.     

Standing crops and vertical distribution of four groups of marine planktonic ciliates in relation to phytoplankton chlorophyll a

Standing crops and the vertical distribution of four groups of ciliates, autotrophic naked ciliates (ANC), heterotrophic naked ciliates (HNC ), mixotrophic naked ciliates (MNC) and loricated ciliates (LC ), were analysed in relation to phytoplankton chlorophyll a in the western Pacific. Data were gathered from the upper 60 to 200 m of water in the subarctic North Pacific in spring, in the subtropical North Pacific in spring, in Toyama Bay in summer, off eastern Australia in spring and off Sanriku in spring and fall. Of the four groups, the standing crop of HNC (cells l⁻¹) showed the highest correlation to phytoplankton chloro- phyll a [CHL, μg l⁻¹; r=0.66, n=365 (HNC=490 ×CHL ^0.653)]. Depths of the maximum layers of HNC and MNC were usually shallower than that of chlorophyll a, while those of ANC and LC occurred frequently beneath the chlorophyll maximum layer. This indicates that these four ciliate groups are substantially different from each other, and that each group holds a different ecological position in the marine microbial food web. Received: 27 October 1997 / Accepted: 27 June 1998     

Stable isotopic evidence for trophic groupings and bio-regionalization of predators and their prey in oceanic waters off eastern Australia

Muscle tissue was collected for stable isotope analysis (SIA) from the main fish predators and their fish and cephalopod prey from oceanic waters off eastern Australia between 2004 and 2006. SIA of δ¹⁵N and δ¹³C revealed that the species examined could be divided into three main trophic groups. A “top predator” group consisted mainly of large billfish (Xiphias gladius and Tetrapturus audax), yellowfin (Thunnus albacares), bigeye (T. obesus) and southern bluefin (T. maccoyii) tunas and sharks; with mako (Isurus oxyrinchus) the highest. Below this tier was a second group composed of mid-trophic level fishes including albacore tuna (Thunnus alalunga), lancet fish (Alepisaurus ferox), mahi mahi (Coryphaena hippuris) and ommastrephid squid. Underlying both groups was a grouping of small fishes including myctophids, small scombrids and nomeids as well as surface fishes including macrorhamphosids. These groupings were based largely on mean animal size which showed a positive linear relation to δ¹⁵N (r ² = 0.58). Some species showed significant ontogenetic variation in either δ¹⁵N (swordfish, lancet fish, yellowfin and albacore tuna) or δ¹³C (mako shark). We also noted a consistent latitudinal change in δ¹⁵N and δ¹³C at ~28°S for the top predator species, particularly albacore and yellowfin tuna. The differences were consistent with a change from oligotrophic Coral Sea to nutrient rich Tasman Sea waters. These differences suggest that predatory fishes may have extended residence time in distinct regions off eastern Australia.     

Predators selectively graze reproductive structures in a clonal marine organism

Although the fitness consequences of herbivory on terrestrial plants have been extensively studied, considerably less is known about how partial predation impacts the fitness of clonal marine organisms. The trophic role of Caribbean parrotfish on coral reefs is complex: while these fish are important herbivores, as corallivores (consumers of live coral tissue), they selectively graze specific species and colonies of reef-building corals. Though the benefits of parrotfish herbivory for reef resilience and conservation are well documented, the negative consequences of parrotfish grazing for coral reproductive fitness have not been previously determined. We examined recently grazed colonies of Montastraea annularis corals to determine whether grazing was positively associated with coral reproductive effort. We measured gonad number, egg number and size, and proportional reproductive allocation for grazed and intact coral colonies 2–5 days prior to their annual spawning time. We found that parrotfish selectively grazed coral polyps with high total reproductive effort (number of gonads), providing the first evidence that parrotfish selectively target specific tissue areas within a single coral colony. The removal of polyps with high reproductive effort has direct adverse affects on coral fitness, with additional indirect implications for colony growth and survival. We conclude that chronic grazing by parrotfishes has negative fitness consequences for reef-building corals, and by extension, reef ecosystems.     

Polymorphism in gigantobacterial symbionts in the guts of surgeonfish (Acanthuridae: Teleostei)

The morphology and partial cytology of a group of bacteria-related organisms from the guts of surgeonfish are described. These gigantobacteria symbionts are encountered in the guts of various species of this fish family throughout the Indo-Pacific region and the Red Sea. They have been tentatively grouped and placed in a “family”, Epulopiscidae, according to the first species described, Epulopiscium fishelsoni Montgomery and Pollak, 1988. In addition to a morphological comparison between the various forms from different species of surgeonfish, suggestions are presented with regard to their possible morphological diversification during evolutionary and host-species specificity. Received: 10 June 1998 / Accepted: 29 October 1998     

Nestmate recognition for eggs in the honeybee (<Emphasis Type="Italic">Apis mellifera</Emphasis> L.)

Colony integrity is fundamental to social insects and is threatened by the reproduction of non-nestmates. Therefore, discrimination between eggs derived from nestmates and non-nestmates would constitute an adaptation to prevent exploitation of the entire cooperative group by unrelated individuals. The removal of nestmate and non-nestmate queen and worker-laid eggs was evaluated in honeybees using colonies of Apis mellifera capensis to test female and of A. m. scutellata to test male eggs. The data show that honeybees can distinguish between nestmate and non-nestmate eggs of both sexes. Moreover, non-nestmate female queen-laid eggs were removed significantly faster than nestmate female worker-laid eggs in A. m. capensis, indicating that nestmate recognition cues can override caste-specific ones. While the experimental manipulation accounts for 37.2% (A. m. scutellata) or 1.6% (A. m. capensis) of variance in relation to egg removal, nestmate recognition explains 33.3% for male eggs (A. m. scutellata) and 60.6% for female eggs (A. m. capensis), which is almost twice as high as the impact of caste (16.7% A. m. scutellata; 25% A. m. capensis). Our data show a stronger effect of nestmate recognition on egg removal in the honeybee, suggesting that cues other than caste-specific ones (viability/kin) can dominate egg removal behavior. In light of intraspecific social parasitism, preventing the reproduction of unrelated individuals (group selection) rather than preferring queens’ eggs (kin selection) appears to be the driving force behind the evolution of egg removal behavior in honeybees.     

No evidence for offspring sex-ratio adjustment to social or environmental conditions in cooperatively breeding purple-crowned fairy-wrens

When fitness returns or production costs vary between male and female offspring, selection is expected to favor females that adjust offspring sex ratio accordingly. However, to what extent vertebrates can do so is the subject of ongoing debate. Here, we explore primary sex ratios in 125 broods of cooperatively breeding purple-crowned fairy-wrens Malurus coronatus. We expected that females might adjust offspring sex ratio because this passerine species experiences considerable variation in social and environmental conditions. (1) However, although helpers substantially increase parental fitness, females (particularly in pairs and small groups) did not overproduce philopatric males (helper-repayment hypothesis). (2) Sex-ratio adjustment based on competition among individuals (helper-competition hypothesis) did not conceal helper-repayment effects or drive sex allocation on its own: while high-quality territories can accommodate more birds, brood sex ratios were independent of territory quality, alone or in interaction with group size. (3) Additionally, males are larger than females and are possibly more costly to produce (costly sex hypothesis), and (4) female offspring may benefit more from long-term effects of favorable conditions early in life (Trivers–Willard hypothesis). Nonetheless, large seasonal variation in food abundance was not associated with a consistent skew in primary sex ratios. Thus, overall, our results did not support the main hypotheses of adaptive sex-ratio adjustment in M. coronatus. We discuss that long-term differential costs and benefits may be insufficient to drive evolution of primary sex-ratio manipulation by M. coronatus females. More investigation is therefore needed to determine the general required sex differences in long-term fitness returns for mechanisms of primary sex-ratio manipulation to evolve.     

Threat-sensitive predator avoidance in damselfish-trumpetfish interactions

Summary Predatory threat can vary during a predator-prey interaction as an attack escalates or among predators at different times. A Threat-sensitivity hypothesis is presented which predicts that prey individuals will trade-off predator avoidance against other activities by altering their avoidance responses in a manner that reflects the magnitude of the predatory threat. This hypothesis was tested in the field by presenting prey (threespot damselfish, Stegastes planifrons) with models of foraging predators (Atlantic trumpetfish, Aulostomus maculatus). During a presentation, damselfish displayed progressively stronger avoidance as predator models were brought nearer; response waned rapidly once predator models passed overhead. Larger predator models and those oriented in a strike pose evoked stronger avoidance reactions than smaller and non-attacking models, intermediate responses were evoked by size and orientation combinations that were intermediate in threat, and habituation was more common to weakly-threatening presentations. Smaller damselfish showed stronger avoidance of models than did larger damselfish. Nonavoidance activities, such as feeding and territorial defense, were curtailed during presentations or were more common during weakly threatening presentations. Approaches to the models, equated with mobbing, were more common among large damselfish, again reflecting degrees of vulnerability among different size prey individuals. These initial results indicate that damselfish threatened by predators respond in a graded manner that reflects the degree of threat posed by the predator, in accordance with the Threat-sensitivity hypothesis.     

Phylogeography of the Indo-Pacific parrotfish Scarus psittacus: isolation generates distinctive peripheral populations in two oceans

Phylogenetic, phylogeographic, population genetic and coalescence analyses were combined to examine the recent evolutionary history of the widespread Indo-Pacific parrotfish, Scarus psittacus, over a geographic range spanning three marine biogeographic realms. We sequenced 164 individuals from 12 locations spanning 17,000 km, from 55oE to 143oW, using 322 base pairs of mitochondrial control region (D-loop). S. psittacus displayed high haplotype (h = 0.83–0.98), but low nucleotide (<1%) diversity. Most (>83%) genetic variation was within populations. AMOVA revealed significant partitioning and identified five geographic groups. These included one central population and four populations peripheral to the centre. The central population occupied reefs from Western Australia to Tahiti and represented the central Indo-Pacific biogeographic realm. Cocos Keeling was distinct from central and western Indo-Pacific biogeographic realms occupying a position intermediate to these. Peripheral populations (Hawaii, Marquesas) represented the eastern Indo-Pacific biogeographic realm, while Seychelles represented the western Indo-Pacific biogeographic realm. All but the central population expanded (<163 kya). Whilst all populations experienced major sea level and SST changes associated with Pleistocene glaciation cycles, the genetic structure of the central population was relatively homogenous unlike the remaining genetically distinctive populations.     

Social discrimination tuning in ants: template formation and chemical similarity

To investigate the role of template plasticity in shaping nest-mate recognition processes in ants, we constructed experimental mixed-species groups of Manica rubida with either Myrmica rubra, Tetramorium bicarinatum or Formica selysi. Selecting Ma. rubida as the focal species, we observed the behaviour within mixed-species groups and the transfer rates of cuticular hydrocarbons (CHC) onto the focal ants, and we also tested the aggression of the focal species reared either alone or in association with each of the three different species. We show that Ma. rubida workers were always amicable towards their mixed group members, as towards members of the respective parental colonies, irrespective of the associated species. They did, however, express different levels of aggression towards single-species groups of the other species tested, depending on the species with which they were reared. The study suggests that similarity in CHC profiles in two species leads to a narrow template in mixed groups, while dissimilarity is followed by lower levels of aggression (a broader template), at least against species with similar CHC compound compositions (i.e. both a broader template in the focal ants and familiarity with the compound groups of the tested individuals operate together). This refutes the hypothesis that ants reared in mixed-species groups are systematically more tolerant. It also demonstrates that heterospecific information is not treated equally during development. We suggest that post-imaginal learning, template reforming and decision making are more precisely tuned when the two species' chemical complexes are similar.     

New Genetic Approach to Detecting Individuals of Rare and Endangered Species

Abstract: Many rare and endangered species are difficult to locate, observe, and study. Consequently, many individuals, breeding pairs, and even populations of such species could remain undetected. Genetic markers can potentially be used to detect the existence of undiscovered individuals and populations, and we propose a method to do so that requires 3 conditions. First, sampling of the known population(s) of the target species must be comprehensive. Second, the species must display a reasonable level of philopatry and genetic structuring. Third, individuals must be able to be caught outside of breeding locations (e.g., at courtship or feeding areas, in flight), and the level of recapture must be reasonably high. We applied our method to the Chatham Island Taiko (Pterodroma magentae), one of the world's most endangered seabirds. We sequenced the Taiko mitochondrial cytochrome b gene and both copies of a fragment of the duplicated domain I of the control region. Twenty‐one haplotypes were revealed, including 4 (19%) not found in birds at known burrows. These results suggest there are more burrow groups yet to be located. The species is a pelagic gadfly petrel that inhabits land only in the breeding season during which it is nocturnal and nests in burrows. Taiko burrows are situated in dense forest in a remote area of Chatham Island, and are consequently difficult to locate and study. It is important that all Taiko burrows be discovered to enable monitoring and protection of the birds from exotic predators.