The relative importance of predation and competition in two reef fishes

Competition and predation may both strongly influence populations of reef fishes, but the importance of these processes relative to one another is poorly understood. I quantified the effects of predation and competition on the growth and survival of two temperate reef fishes, Lythrypnus dalli and Coryphopterus nicholsii, in field experiments in which I manipulated the densities of the two species and the abundance of predators (using exclosure cages) on small replicate patch reefs. I also evaluated the influence of predators on the behavior of the two species to help interpret the mechanisms of any predatory influences on growth or survival. Predation was much more important than competition (inter- or intraspecific) in Lythrypnus. For Coryphopterus, neither competition nor predation were particularly important. Behaviorally, both species responded to predators by reducing foraging rate and hiding. This altered behavior, however, had no repercussions for growth or survival of Coryphopterus. In contrast, Lythrypnus grew more slowly and suffered greater mortality when exposed to predators. Interspecific competition did not significantly influence either species. Intraspecific competition did not affect the growth of Coryphopterus, but survival tended to be lower at high densities. Growth of Lythrypnus was depressed by intraspecific competition, but survival was not, except that, in the presence of predators, survival was density dependent. In contrast to the historical emphasis placed on the role of competition, this study indicates that predation can be more important than competition in determining patterns of abundance of some reef fishes. For example, predators not only influenced foraging of both Lythrypnus and Coryphopterus, but they also reduced growth and survival of Lythrypnus, and therefore appear to help maintain the marked habitat segregation between the two species. Received: 16 June 1997 / Accepted: 3 December 1997     

黄果厚壳桂种内与种间竞争的数量关系

采用Hegyi单木竞争指数模型对鼎湖山季风常绿阔叶林建群种黄果厚壳桂的种内、种间竞争强度进行定量分析.结果表明,黄果厚壳桂所受到的竞争强度随着林木径级的增大而逐渐减小.黄果厚壳桂种内竞争较与伴生树种云南银柴的种间竞争弱.黄果厚壳桂种内和种间竞争强度的顺序为:云南银柴>黄果厚壳桂种内>荷木>白颜树>肖蒲桃>锥栗>红车>臀形果>柏拉木>水石梓>窄叶半枫荷>厚壳桂.整个林分及黄果厚壳桂种内竞争木对对象木的竞争强度与对象木的胸径大小之间的关系近似服从幂函数关系,而伴生树种竞争木对对象木的竞争强度与对象木的胸径大小之间的关系近似服从对数函数关系.竞争强度和对象木的胸径大小呈显著负相关关系.     

Sexual dimorphism in intra- and interspecific competitive ability of the dioecious herb Mercurialis annua

Males and females of dioecious plant species often show different responses to competition with individuals of the same or opposite gender, but almost no data are available on the outcome of competition with members of other species. Here, we show that male and female individuals of the wind-pollinated herb Mercurialis annua are sexually dimorphic in both their intraspecific and interspecific competitive abilities. In a controlled experiment, we found that both sexes of M. annua were negatively affected by interspecific competition, but the sensitivity of males and females depended on the identity of their competitor species, with females tending to suppress the aboveground growth of competitor species more than males. Further, we found that intrasexual and intersexual competition affected the aboveground growth of males but not that of females: only males showed a significant reduction in growth when growing with conspecific competitors (male or female). We discuss our results with reference to related studies that suggest that males and females of M. annua have different resource requirements for reproduction, which in turn affect their competitive abilities.     

Interactive effects of interspecific competition and microhabitat on early post-settlement survival in a coral reef fish

Microhabitat type and the competition for microhabitats can each influence patterns of abundance and mortality in coral reef fish communities; however, the effect of microhabitat on the intensity and outcome of competition is not well understood. In Kimbe Bay, Papua New Guinea, surveys were used to quantify microhabitat use and selectivity in two live-coral specialist damselfishes (Pomacentridae), Chrysiptera parasema, and Dascyllus melanurus. A patch reef experiment was then conducted to test how intra- and interspecific competition interacts with two types of microhabitat to influence survival of recently settled C. parasema. Surveys demonstrated that C. parasema and D. melanurus recruits utilized similar coral microhabitats; 72% of C. parasema and 85% of D. melanurus used corymbose and bottlebrush growth forms of Acropora. One microhabitat type, Pocillopora sp. coral, was commonly used by D. melanurus but rarely by C. parasema. The patch reef experiment revealed that both microhabitat and interspecific competition influence abundance of recently settled C. parasema. Microhabitat had the strongest influence on survival of C. parasema. In the absence of interspecific competitors, ~85% of C. parasema survived for 5 days after transplantation to high-complexity bottlebrush Acropora reefs when compared to only 25% survival of Pocillopora reefs. In both microhabitats, interspecific competition with D. melanurus, but not intraspecific competition, significantly decreased the survival of C. parasema. Taken together, these results suggest that the observed distribution of C. parasema results from specialized microhabitat requirements and competition for space in those microhabitats. This study demonstrates that interspecific competition and microhabitat type can interact to influence early post-settlement survival in coral reef fishes, though, whether and how these factors influence survival will depend on the behavioural attributes and strength of habitat associations among potential competitors. Communicated by Environment Editor Prof. Rob van Woesik     

The role of herbivorous fishes and urchins in coral reef communities

Synopsis Herbivorous fishes and invertebrates are conspicious elements of coral reef communities where they predominate both in numbers and biomass. Herbivores and the coral reef algae on which they feed represent a co-evolved system of defense and counter-defense. Algal species have developed toxic, structural, spatial and temporal defense or escape mechanisms, while the herbivores employ strategies that involve anatomical, physiological and behavioral adaptations. Current research demonstrates that many reef fishes are highly selective in the algae they consume. Food selection in these fishes may be correlated with their morphological and digestive capabilities to rupture algal cell walls. Sea urchins select more in accordance with relative abundance, although certain algal species are clearly avoided.The determinants of community structure on coral reefs have yet to be established but evidence indicates a strong influence by herbivores. Reef herbivores may reduce the abundance of certain competitively superior algae, thus allowing corals and cementing coralline algae to survive. We discuss how the foraging activities of tropical marine herbivores affect the distribution and abundance of algae and how these activities contribute to the development of coral reef structure and the fish assemblages which are intimately associated with reef structure.This paper forms a part of the proceedings of a mini-symposium convened at Cornell University, Ithaca, N.Y., 18–19 May 1976, entitled Patterns of Community Structure in Fishes (G. S. Helfman, ed.).     

Analysis of interspecific competition in perennial plants using Life Table Response Experiments

The impact of interspecific competition is usually measured by its effect upon plant growth, neglecting impacts upon other stages of the life cycle such as fecundity which have a direct influence upon individual fitness and the asymptotic population growth rate of a population (λ). We used parameterized matrix models for three perennial plant species grown with and without interspecific competition to illustrate how the methodology of Life Table Response Experiments (LTRE) can be used to link any change in population dynamics to changes in any part of the life cycle. Plants were herbaceous grassland species grown for two years in a field experiment at Rothamsted Experimental Station, England. Interspecific competition reduced λ by over 90% in all species. Survival and growth were slightly affected by competition whereas plant fecundity was greatly reduced. Nearly all of the observed difference in λ between the competition treatments was explained by the fecundity terms, and more precisely by a large difference in the number of seeds, and a high sensitivity of λ to the germination rate. Whereas most competition studies focus on the measurement of change in individual fitness, our study illustrates how informative it is to take account not only of the effect of competition upon vital rates but also of how different vital rates affect population growth rate.     

Patterns of intra- and interspecific association in leaf-mining insects on three oak host species

Abstract. 1. We determined mortality and distributional patterns of leaf miners on three oak host species (Quercus falcata, Q.nigra and Q.hemisphaerica) in northern Florida, U.S.A.
2. Patterns of intra- and interspecific occurrence within leaves, and mortality of five most abundant leaf miner species were analysed as a test of competition.
3. Miners co-occurred on leaves more often that expected by chance (P<0.05) in six of ten possible species combinations and log-linear model analysis showed no negative higher-order interactions.
4. All five miner species had highly clumped distributions between leaves (P<0.01).
5. Leaf miner survival was less than expected for four of five species when co-occurring on leaves with conspecifics than when mining with heterospecifics or alone (P<0.05).
6. We conclude that interspecific competition is unapparent within this leaf miner guild and that intraspecific competition occurs in four of the five major leaf miner species. We discuss leaf miner selection of common leaves, perhaps based on chemical/physical leaf characters, as a cause of intra- and interspecific aggregation.     

Effects of nitrogen on interspecific competition between two cell-size cyanobacteria: Microcystis aeruginosa and Synechococcus sp.

Micro-cyanobacteria and pico-cyanobacteria coexist in many lakes throughout the world. Their distinct cell sizes and nutrient utilization strategies may lead to dominance of one over the other at varying nutrient levels. In this study, Microcystis aeruginosa and Synechococcus sp. were chosen as representative organisms of micro- and pico-cyanobacteria, respectively. A series of nitrate and ammonia conditions (0.02, 0.1, 0.5, and 2.5 mg N L⁻¹) were designed in mono- or co-cultured systems, respectively. Growth rates of the two species were calculated and fitted by the Monod and Logistic equations. Furthermore, the interspecific competition was analyzed using the Lotka–Volterra model. In mono-cultures, the two cyanobacteria displayed faster growth rates in ammonia than in nitrate. Meanwhile, Synechococcus sp. showed faster growth rates compared to M. aeruginosa in lower N groups (≤ 0.5 mg N L⁻¹). However, in the highest nitrate treatment (2.5 mg N L⁻¹), M. aeruginosa achieved much higher biomass and faster growth rates than Synechococcus sp.. In co-cultures, Synechococcus sp. dominated in the lowest N treatment (0.02 mg N L⁻¹), but M. aeruginosa dominated under the highest nitrate condition (2.5 mg N L⁻¹). Based on the analysis of Raman spectra of living cells in mono-cultures, nitrate (2.5 mg N L⁻¹) upgraded the pigmentary contents of M. aeruginosa better than ammonia (2.5 mg N L⁻¹), but nitrogen in different forms showed little effects on the pigments of Synechococcus sp.. Findings from this study can provide valuable information to predict cyanobacterial community succession and aquatic ecosystem stability.     

Mixed-species brooding between two Baikal sculpins: field evidence for intra- and interspecific competition for reproductive resources

In an examination of the distribution of nesting spaces of two Baikal sculpins, Paracottus kneri and Cottus kessleri , along two transects offshore at Bolshye Koty, a total of 43 P. kneri nests, 32 C. kessleri nests and one Batrachocottus baicalensis nest was found. Males of these species established breeding nests under overhanging stones in the same manner as most freshwater sculpins, but nests of each species were clearly partitioned in separate zones. Cottus kessleri nests were found where the lake floor was mostly covered with overhanging stones. Paracottus kneri nests, however, were found where the lake floor was moderately covered with overhanging stones. Two mixed-species brooding nests with a C. kessleri guarding male were found in an area bounded by the two zones. Each of these nests comprised one P. kneri egg mass and several C. kessleri egg masses, guarded by a C. kessleri male. The nest distribution of these fishes appears to reflect intra- and interspecific competition for overhanging stones as a reproductive resource rather than interspecific difference of nest selection.     

栖息环境和种间竞争对中华蜜蜂群体分布的影响

在植被结构和生态条件不同的皖南山区和皖西大别山区、江淮地区及淮北平原,栖息环境的改变和种间竞争的生存竞争等因素是影响中蜂群体分布的主要因素．皖南山区和皖西大别山区自然植被完整,生态条件好,蜜粉源植物丰富,中蜂群体数量多,分布区域广,分布密度高,中蜂蜂王自然交配极少受到干扰,种间竞争处于优势,是中蜂栖息与繁衍的理想场所．江淮地区和淮北平原自然植被少,生态平历受到不同程度破坏,蟹粉源植物种类少,花期短而集中,中蜂蜂王自然交配受到意蜂雄蜂干扰,种间竞争处于劣势,中蜂群体数量锐减,分布区域缩小,群体分布密度大幅度下降,淮北平原比江淮地区更显著．     

Recent changes in coral assemblages of a South African coral reef, with recommendations for long-term monitoring

Two-mile reef, Sodwana, South Africa is an unusual coral reef, being situated on a submerged fossilized sand dune and being very southerly (27°54). It is a popular Scuba diving venue receiving about 100000 dives year^–1. The line-intercept transect method, as recommended by the global coral reef monitoring network (GCRMN), was used to determine soft coral, hard coral and other benthos percentage cover. Physical coral damage, disease and bleaching were also recorded. Results were compared with those of B. Riegl (1993 – unpublished PhD thesis) 5 to 7 years earlier. The reef appears to be ecologically and highly dynamic. In the interim, there has been an increase in living benthos cover of 22.3% but also an increase in coral bleaching from 0% in 1993 to 1% in 1998. Physical damage, despite the large number of dives on the reef was minimal (1.52%), although it appears as if coral diseases may be increasing. The 20-m transects recommended by GCRMN are too long for this highly rugose reef with its distinct ridges and gullies. It is recommended that benthos cover, coral damage, bleaching and disease should be monitored annually using 40 5-m transects on the reef ridges and 40 5-m transects on the reef slopes.     

The stability of P in coral reef fishes

The constancy of phenotypic variation and covariation is an assumption that underlies most recent investigations of past selective regimes and attempts to predict future responses to selection. Few studies have tested this assumption of constancy despite good reasons to expect that the pattern of phenotypic variation and covariation may vary in space and time. We compared phenotypic variance-covariance matrices (P) estimated for populations of six species of distantly related coral reef fishes sampled at two locations on Australia's Great Barrier Reef separated by more than 1000 km. The intraspecific similarity between these matrices was estimated using two methods: matrix correlation and common principal component analysis. Although there was no evidence of equality between pairs of P, both statistical approaches indicated a high degree of similarity in morphology between the two populations for each species. In general, the hierarchical decomposition of the variance-covariance structure of these populations indicated that all principal components of phenotypic variance-covariance were shared but that they differed in the degree of variation associated with each of these components. The consistency of this pattern is remarkable given the diversity of morphologies and life histories encompassed by these species. Although some phenotypic instability was indicated, these results were consistent with a generally conserved pattern of multivariate selection between populations.     

Influence of size and spatial competition on the bioactivity of coral reef sponges

Sponges biosynthesize a wealth of secondary metabolites, many with novel structures and strong biological activity. Such compounds may serve multiple ecological roles including anti-predation, anti-fouling functionalities and are implicated in border defense or attack during spatial competition. Relative size of benthic organisms may also play an important role in competitive interactions. To determine if a relationship exists between individual size and bioactive metabolite production in the context of spatial competition, we examined three sponge species with different morphologies: the massive Coscinoderma matthewsi, the club-shaped branching Hyrtios erecta, and the fan-shaped Ianthella basta. Extracts from sponges of various sizes and competitive environments were examined using a cell based bioassay as a proxy of bioactivity. For I. basta, sponge size was correlated with bioactivity; the largest individuals generally being the most bioactive. In contrast, there was no correlation between size and bioactivity for either C. matthewsi or H. erecta. Bioactivity of sponges in this study were however highly variable among individuals, regardless of levels of competition. The prevalence of encroaching organisms was not correlated with sponge size for any of the three sponge species, suggesting that potential bioactivity is not influenced by surrounding competition.     

Evidence of a geographically variable competitive mimicry relationship in coral reef fishes

In Kimbe Bay, Papua New Guinea, juvenile surgeonfish Acanthurus pyroferus have been shown to gain access to food resources defended by the damselfish Plectroglyphidodon lacrymatus by mimicking a pygmy angelfish, Centropyge vrolikii , that does not compete with the damsel for food. I tested whether A. pyroferus juveniles gain the same competitive advantage from mimicking a different pygmy angelfish, Centropyge flavissima , in Moorea, French Polynesia. Through abundance and substrate surveys, behavioral observations and stomach content analyses, I demonstrate that in Moorea, mimicry of Ce. flavissima does not provide A. pyroferus with access to damselfish Stegastes nigricans territories; Ce. flavissima models are always attacked upon territory entry and A. pyroferus mimics avoid damsel territories. Damselfish aggression toward the model angelfish cannot be attributed to overabundance of the deceptive mimic; instead, aggression can best be explained by the fact that Ce. flavissima competes with damsels in Moorea by consuming their algal turfs, making them inappropriate models for competitive mimics. Juveniles of many Indo-Pacific surgeonfishes appear to mimic pygmy angelfishes; I suggest that these mimics' success in gaining access to damselfish territories is geographically variable and may be determined by the extent to which mimics, models and receivers overlap in resource use at a given site. This mimicry complex may thus present an excellent illustration of the geographic mosaic model of coevolution.     

Two-species asymmetric competition: effects of age structure on intra- and interspecific interactions

1. The patterns of density-dependent resource competition and the mechanisms leading to competitive exclusion in an experimental two-species insect age-structured interaction were investigated. 2. The modes of competition (scramble or contest) and strength of competition (under- to overcompensatory) operating within and between the stages of the two species was found to be influenced by total competitor density, the age structure of the competitor community and whether competition is between stages of single or two species. 3. The effect of imposed resource limitation on survival was found to be asymmetric between stages and species. Environments supporting both dominant and subordinate competitors were found to increase survival of subordinate competitors at lower total competitor densities. Competitive environments during development within individual stage cohorts (i.e. small or large larvae), differed from the competitive environment in lumped age classes (i.e. development from egg-->pupae). 4. Competition within mixed-age, stage or species cohorts, when compared with uniform-aged or species cohorts, altered the position of a competitive environment on the scramble-contest spectrum. In some cases the competitive environment switched from undercompensatory contest to overcompensatory scramble competition. 5. Such switching modes of competition suggest that the relative importance of the mechanisms regulating single-species population dynamics (i.e. resource competition) may change when organisms are embedded within a wider community.     

Phylogeography and the conservation of coral reef fishes

Here we present a review of how the study of the geographic distribution of genetic lineages (phylogeography) has helped identify management units, evolutionary significant units, cryptic species, and areas of endemism, and how this information can help efforts to achieve effective conservation of coral reefs. These studies have confirmed the major biogeographic barriers that were originally identified by tropical species distributions. Ancient separations, identified primarily with mtDNA sequence comparisons, became apparent between populations on each side of the barriers. The general lack of correlation between pelagic larval duration and genetic connectivity across barriers indicates that life history and ecology can be as influential as oceanography and geography in shaping evolutionary partitions within ocean basins. Hence, conservation strategies require a recognition of ecological hotspots, those areas where habitat heterogeneity promotes speciation, in addition to more traditional approaches based on biogeography. Finally, the emerging field of genomics will add a new dimension to phylogeography, allowing the study of genes that are pertinent to recent and ongoing differentiation, and ultimately providing higher resolution to detect evolutionary significant units that have diverged in an ecological time scale.     

Behavioural acclimation to cameras and observers in coral reef fishes

Observer presence can bias behavioural studies of animals in both the wild and the laboratory. Despite existing evidence for significant observer effects across several taxa, little is known about the minimum periods of acclimation that should precede behavioural observations. To date, most studies either do not report any acclimation periods or include a non‐specific period without empirically quantifying its appropriateness. Here, we conducted in situ behavioural observations of two species of demersal coral reef fishes using cameras and/or observers to examine the biases associated with either approach. For both treatments, we generated 25 min time series of a number of vigilance‐associated behaviours (i.e., distance from shelter and mate, time out of shelter, swimming activity) and estimated the point of acclimation using changepoint analysis. In the camera trials, acclimation in both species appeared to occur between 2 and 7 min for different behaviours. When an observer was present, however, no apparent acclimation occurred until the observer left the area. Overall, our findings demonstrate that (i) behavioural studies of wild fishes conducted by an observer may be biased due to permanent observer effects, and (ii) when using video equipment, a species‐ and behaviour‐specific acclimation period should precede behavioural scoring.     

Increasing Himalayan tahr and decreasing chamois densities in the eastern Southern Alps, New Zealand: evidence for interspecific competition

There is anecdotal evidence that increasing densities of Himalayan tahr (Hemitragus jemlahicus) are associated with declining densities of chamois (Rupicapra rupicapra) in the Southern Alps, New Zealand. To examine this phenomenon, densities of tahr and chamois were measured at 53 sites within their sympatric range in the eastern Southern Alps during 1978–1979. In sites where only one species was present, tahr density was significantly higher than chamois density (P=0.032), probably reflecting species differences in sociality. Chamois density was higher in catchments without tahr than in those with tahr (P=0.012). Similarly, tahr density was significantly higher at sites without chamois than at those with chamois (P=0.033). Sites with both species present (termed sympatric) were significantly larger than those with only chamois or tahr (P<0.001). Following the prohibition of aerial hunting of tahr in 1983, 16 of the 17 sites where tahr and chamois were sympatric during 1978–1979 were recounted during 1991–1996. There was a 6-fold increase in the mean density of tahr between the two counts (P=0.001), whereas chamois density had declined significantly (P=0.006). Chamois persisted at only three sites, two of which had the highest chamois densities in 1978–1979. This is evidence that increasing densities of tahr exclude chamois from all but the `best' habitats. We conclude that intensive aerial hunting of tahr during 1967–1983 reduced tahr densities such that chamois could co-exist with tahr. Received: 20 March 1997 / Accepted: 8 September 1997