Evidence of density-independent mortality in a settling coral reef damselfish, Chromis viridis

To know if the variation in the number of settling fish larvae can be dampened by density-dependent postsettlement mortality, we investigated the relationship between settler density and predator-induced mortality of a coral reef damselfish, Chromis viridis. Totals of 2, 3, 5, 8, 10, 12, 14, 16, 18, and 20 fish of 10 or 20 mm total length were released in experimental cages enclosing a coral head of Porites rus (to provide settlement habitat) and five predators. The results showed that the mortality rate of both 10- and 20-mm fish was density independent.     

Population dynamics of coral-reef fishes: Controversial concepts and hypotheses

Abstract Knowledge of processes that drive the local population dynamics of coral-reef fishes is important for managing reef fisheries, and for using these species as models for understanding the ecology of demersal marine fishes in general. However, the reef-fish literature is replete with poorly defined concepts and vague hypotheses regarding the issue of population dynamics. Dichotomous arguments, such as whether or not recruitment drives population dynamics, are misdirected because they fail to incorporate several important concepts. First, changes in local population size are driven by four demographic rates (birth, death, immigration and emigration), all of which must be studied to understand population dynamics. Second, all populations that persist do so because at least one of these demographic rates operates in a density-dependent way that is both sufficiently strong and appropriately time-lagged. Therefore, identifying the source(s) of direct density dependence is critical for understanding the limits to variation in population size (i.e. population regulation). Third, regulation does not imply a simple point equilibrium in population size; density dependence in populations of reef fishes is bound to lie within a field of stochastic variation, and thus be difficult to detect. Since its formal origin in 1981, the ‘recruitment limitation’ hypothesis for explaining local population dynamics in reef fishes has undergone ambiguous changes in definition that threaten its usefulness. ‘Recruitment, ‘originally defined as the appearance of newly settled fish on a reef, more recently is often measured months after settlement, thus confounding pre- and post-settlement processes. ‘Limitation, ‘ which originally referred to recruitment being so low as to preclude local populations from reaching densities where resources were limiting, is more recently defined as an absence of any form of density dependence after settlement. The most effective means of testing whether post-settlement mortality is in fact density-independent is to examine patterns of mortality directly, rather than indirectly by interpreting the shape of the relationship between initial recruit density and subsequent adult density within a cohort (the recruit-adult function). Understanding the population dynamics of coral-reef fishes will require a more equitable focus on all four demographic rates, be they density dependent or not, as well as greater attention to identifying sources of density dependence. Such a pluralistic focus necessitates integrated studies of both pre- and post-settlement processes conducted at multiple spatial and temporal scales. For example, recent evidence suggests that density-dependent pre-dation on new recruits that have settled among reefs at different densities may prove to be an important source of local population regulation, especially via the aggregative response of transient piscivores.     

The predation gauntlet: early post-settlement mortality in reef fishes

Most marine fishes have pelagic larvae that settle to benthic juvenile/adult habitats. Ecologists have argued that mortality rates are particularly high during the settlement transition, but relevant data have been sparse. Recently, researchers have used several novel techniques to estimate the magnitude of predation mortality during the settlement transition. We used meta-analysis to determine that for 24 taxonomically diverse species in geographically widespread locations, an estimated 55.7% (CI: 43.0–65.5%) of juveniles were consumed within 1–2 days of settlement. Such high mortality highlights this brief period as a key phase in the life history of fishes and supports the view that these communities are strongly influenced by predation. Additionally, we argue that because predators have such strong effects on juvenile survival, the population and community dynamics of reef fishes may be linked to human exploitation of reef predators.     

The adaptive significance of larval dispersal in coral reef fishes

Synopsis Coral reef fishes almost universally disperse over relatively great distances during a pelagic larval phase. Barlow (1981) suggested that this dispersal is adaptive because adult fishes inhabit a patchy, uncertain environment. This reiterated an older idea that the random extinction of local populations necessarily favours dispersal, since ultimately all populations of non-dispersers will disappear. Whereas this view is based on adult survival, we emphasize a less frequent view that substantial larval dispersal may be adaptive when offspring experience patchy and unpredictable survival in the pelagic habitat. We do not address the question of why these animals ‘broadcast’ rather than ‘brood’, but suggest that species committed to pelagic offspring will be under selection to disperse siblings to spread the risk of failure among members of a cohort. Our arguments are supported by a heuristic computer simulation.     

Rethinking ecological inference: density dependence in reef fishes

We use a meta‐analysis of density dependence in reef fishes to evaluate how ecologists approach detection, inference, and estimation. We compared two groups of studies: those that detected effects of density on survival and those that did not. Distinctions between these groups have spawned heated debate about the processes that affect fish dynamics. Per capita effects of density were similar between the two groups, although total effects (and hence ambient density) were greater in studies that detected density effects. The majority of the variation in effects of density was not resolved by the classification of studies based on the authors' conclusions. These results suggest (1) that standard inferences based on null hypothesis tests may miss important sources of variation in effects and give rise to unnecessary debate; and (2) that estimation of effect sizes and model parameters (including their uncertainty) is a powerful alternative to detection of ecological processes.     

Ocean current variability and the spawning season of Hawaiian reef fishes

Synopsis The spawning patterns of four Hawaiian reef fishes with similar reproductive habits, but different biogeographic distributions were studied from September 1980 to October 1981. Two species are Hawaiian endemics [Centropyge potteri (Pomacanthidae) and Chaetodon multicinctus (Chaetodontidae)] and the other two have pan-tropical Pacific distributions [Ctenochaetus strigosus and Zebrasoma flavescens (Acanthuridae)]. All showed increased spawning activity from January to July, the prevalent pattern among a majority of coastal marine fishes in Hawaii. Environmental correlatives to the period of peak reproduction include cycles of (a) daylight length and temperature which probably function as proximate cues and (b) seasonal variations in ocean current patterns which may ultimately affect survival of larvae and dispersal. Peak reproduction takes place during months when (1) mesoscale eddies most likely occur, and (2) the probability is greatest of drifting objects remaining near the islands.     

Age-specific mortality rates in reef fishes: Evidence and implications

Abstract Mortality is a fundamental demographic rate, the nature of which has profound consequences for both the dynamics of populations and the life-history evolution of species. For example, if per capita mortality rates are age- or stage-specific, life-history traits should evolve in response to age- and stage-specific differences in selection arising from these temporally variable rates. Similarly, variation in the average mortality rate across ages and/or stages can also select for shifts in life history. Mortality rates of recently settled reef fishes can be very high and per capita mortality is commonly assumed to decrease with increasing age. A review of evidence for age-specific per capita mortality rates in reef fishes from early postsettlement up to 13 months postsettlement suggests that during this period these rates are often age invariant. The data on which these interpretations are based, however, are extremely limited both in terms of the proportion of the life cycle over which mortality rates have been sampled and the quality of these data. Nonetheless, these data do suggest that selective pressures associated with patterns of mortality may vary among species of reef fishes and that these species therefore could be more effectively used in the study of life-history evolution. At present, reef fishes are under-represented in the study of life-history evolution compared with other vertebrate taxa.     

Population regulation by post-settlement mortality in two temperate reef fishes

 Input of individuals dispersing into open populations can be highly variable, yet the consequences of such variation for subsequent population densities are not well understood. I explored the influence of variable input (”supply”) on subsequent densities of juveniles and adults in open local populations of two temperate reef fishes, the bluebanded goby (Lythrypnus dalli) and the blackeye goby (Coryphopterus nicholsii). Variable recruitment was simulated by stocking a natural range of densities of young fishes on replicate patch reefs. Density and mortality of the stocked cohorts were followed over time, until the fishes reached maturity. Over the first day of the experiments, mortality of both species was significantly density-dependent; however, there was still a very strong relationship between density on day 1 and density on day 0 (i.e., simulated recruitment was still an excellent predictor of population density). At this point in the study, the main effects of density-dependent mortality were to reduce mean densities and variation about the mean. Over the period from the start of the experiments until the time when maturity was reached by each species (about 1 and 3 months for Lythrypnus and Coryphopterus, respectively), mortality was strongly density-dependent. Such strong density-dependent mortality virtually eliminated any linear relationship between adult density and ”recruit” density. However, for both species, the relationship between these two variables was well fit by an asymptotic curve, with the asymptotic density of adults equal to c. 3/m² for Coryphopterus, and c. 10/m² for Lythrypnus. Natural recruitment (via settlement of larvae) to the reefs over the period of the study (9 months) was above the asymptotic densities of adults for the two species, even though the study did not encompass the periods of peak annual recruitment of either species. This suggests that adult populations of these two gobies may often be limited, and regulated, by post-settlement processes, rather than by input of settlers. Other studies have shown that mortality of the two species is density-independent, or only weakly density-dependent, on reefs from which predators have been excluded. Hence, it appears that predators cause density-dependent mortality in these fishes. Received: 26 November 1996 / Accepted: 5 April 1997     

Post-settlement mortality and growth of newly settled reef corals in a subtropical environment

There are few studies of post-settlement mortality of newly settled corals, particularly on subtropical reefs. This study examined the mortality and growth of spat of three broadcast spawning coral species at the Solitary Islands, eastern Australia (30°S). Mortality of spat was high, with only 0.2–2.8% surviving their first year of life. Growth was slow, with coral spat achieving a maximum diameter of 2 mm after 8 months. High post-settlement mortality and slow growth rates are likely to contribute to low rates of recruitment of broadcast spawning species at the Solitary Islands.     

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.     

Regulation of local populations of a coral reef fish via joint effects of density- and number-dependent mortality

Density-dependent mortality can regulate local populations - effectively minimizing the likelihood of local extinctions and unchecked population growth. It is considered particularly important for many marine reef organisms with demographically open populations that lack potential regulatory mechanisms tied to local reproduction. While density-dependent mortality has been documented frequently for reef fishes, few studies have explored how the strength of density-dependence varies with density, or how density-dependence may be modified by numerical effects (i.e., number-dependent mortality). Both issues can have profound effects on spatial patterns of abundance and the regulation of local populations. I address these issues through empirical studies in Moorea, French Polynesia, of the six bar wrasse (Thalassoma hardwicke), a reef fish that settles to isolated patch reefs. Per capita mortality rates of newly settled wrasse increased as a function of density and were well approximated by the Beverton-Holt function for both naturally formed and experimentally generated juvenile cohorts. Average instantaneous mortality rates were a decelerating function of initial densities, indicating the per capita strength of density-dependence decreased with density. Results of a factorial manipulation of density and group size indicate that per capita mortality rates were simultaneously density- and number-dependent; fish at higher densities and/or in groups had higher probabilities of disappearing from patch reefs compared with fish that were solitary and/or at lower densities. Mortality rates were ~30% higher for fish at densities of 0.5 fish/m² than at 0.25 fish/m². Similarly, mortality rates increased by ~45% when group size was increased from 1 to 2 individuals per patch, even when density was kept constant. These observations suggest that the number of interacting individuals, independent of patch size (i.e., density-independent effects) can contribute to regulation of local populations. Overall, this work highlights a greater need to consider numerical effects in addition to density effects when exploring sources of population regulation.     

Density dependent recruitment in the reef fish Chaetodon miliaris

Synopsis Recruitment of reef fish has generally been found to be unrelated to or positively related to adult densities. This paper reports an inverse relationship between the number of recruits of Chaetodon miliaris and the density of conspecific adults. C. miliaris are non-territorial, planktivorous butterflyfish. The study populations occurred on the flanks of patch reefs of about 30 m in diameter. At their peak densities populations consisted of 300–400 C. miliaris. Recruitment (the appearance in the stock of fish less than 3.5cm in total length) occurred primarily from April to June and corresponded to the new moon period in these months. Populations generally showed a steady decline in numbers during months of the year when recruitment was not occurring: the finite rate of decline was approximately 30% per month. Populations varied in the density of adults at the start of the spring-summer period of recruitment, and the magnitude of recruitment to a reef was inversely related to the density of these conspecific adults. Reduction of the densities of adult C. miliaris through trap fishing, resulted in increased recruitment to the fished stock. These results provide evidence that in some. species of reef fish, benthic processes may play an important role in determining the magnitude of recruitment to an adult stock, in contrast to the widely held view that recruitment is in large part a result of chance events in the plankton or that recruitment is directly related to adult densities.     

Spatial scales in the study of reef fishes: A theoretical perspective

Abstract Theoretical models imply that spatial scale derives its greatest importance through interactions between density-dependent processes and spatial variation in population densities and environmental variables. Such interactions cause population dynamics on large spatial scales to differ in important ways from predictions based on measurements of population dynamics at smaller scales, a phenomenon called the scale transition. These differences can account for large-scale population stability and species coexistence. The interactions between density dependence and spatial variation that lead to the scale transition can be understood by the process of non-linear averaging, which shows how variance originating on various spatial scales contributes to large-scale population dynamics. Variance originating below the scale of density dependence contributes less to the scale transition as the spatial scale of the variation declines, while variation originating on or above the scale of density dependence contributes independently of the spatial scale of the variation.     

Coral recruitment and juvenile mortality as structuring factors for reef benthic communities in Biscayne National Park, USA

Coral communities of Biscayne National Park (BNP) on offshore linear bank-barrier reefs are depauperate of reef corals and have little topographic relief, while those on lagoonal patch reefs have greater coral cover and species richness despite presumably more stressful environmental regimes closer to shore. We hypothesized that differences in rates of coral recruitment and/or of coral survivorship were responsible for these differences in community structure. These processes were investigated by measuring: (1) juvenile and adult coral densities, and (2) size-frequency distributions of smaller coral size classes, at three pairs of bank- and patch-reefs distributed along the north-south range of coral reefs within the Park. In addition, small quadrats (0.25 m²) were censused for colonies <2 cm in size on three reefs (one offshore and one patch reef in the central park, and one intermediate reef at the southern end), and re-surveyed after 1 year. Density and size frequency data confirmed that large coral colonies were virtually absent from the offshore reefs, but showed that juvenile corals were common and had similar densities to those of adjacent bank and patch reefs. Large coral colonies were more common on inshore patch reefs, suggesting lower survivorship (higher mortality) of small and intermediate sized colonies on the offshore reefs. The more limited small-quadrat data showed similar survivorship rates and initial and final juvenile densities at all three sites, but a higher influx of new recruits to the patch reef site during the single annual study period. We consider the size-frequency data to be a better indicator of juvenile coral dynamics, since it is a more time-integrated measurement and was replicated at more sites. We conclude that lack of recruitment does not appear to explain the impoverished coral communities on offshore bank reefs in BNP. Instead, higher juvenile coral mortality appears to be a dominant factor structuring these communities. Accepted: 9 September 1999     

Age structure and reproduction in feral New Zealand populations of the house mouse (Mus musculus), in relation to seedfall of southern beech

Abstract

After good seedfalls by southern beech (Nothofagus spp.), density indices for mice increased, the breeding season for adult females was brought forward, and winter breeding was observed. The end of the breeding season of the following summer was brought forward only slightly if the maximum density achieved was not great; incomplete data on 2 very high-density populations suggested a much earlier end to breeding. In early winter following a good seedfall the recruitment of juveniles was temporarily increased. After spring, few young mice entered the population, though breeding continued. The distribution of age classes shifted steadily upwards as population density rose. The peak population of 1976–77 comprised mainly old mice, still breeding, but without significant recruitment. No effects of density on sex ratio, litter size, or body weight could be detected from these data. In all populations studied, density decreases were ultimately due to failure of recruitment and the disappearance of ageing, non-breeding mice. The proximate causes of this mortality were probably the onset of cold weather in 1976–77 and predation by stoats in 1979–80.     

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.).     

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     

Assessing early recruitment dynamics and its demographic consequences among tropical reef fishes: Accommodating variation in recruitment seasonally and longevity

Abstract When settlement of pelagic juveniles of reef fishes is highly and predictably seasonal, annual, end-of-season surveys of surviving recruits (which are commonly used on the Great Barrier Reef) are useful for assessing recruitment dynamics and their demographic effects. However, when settlement is continuous or weakly seasonal, with patterns that vary both between species and within species among years, regular, sometimes year-round, recruitment surveys at intervals linked to short-term settlement dynamics are needed to quantify fluctuations in recruitment strength. Monthly recruitment surveys may be appropriate in the tropical northwest Atlantic, where settlement is often both lunar periodic, and broadly and variably seasonal. Use of a variety of recruit-census methods impedes comparisons of recruitment patterns and their demographic effects, because recruit densities and recruit:adult ratios cannot be directly compared when recruits (because they have widely varying post-settlement ages) have experienced very different levels of early post-settlement mortality. Examining the relationship between changes in adult populations and annual, end-of-season recruitment may be satisfactory for long-lived species with strong settlement seasonality and maturation times of approximately 1 year. However, it is inappropriate for short-lived, rapidly maturing species, particularly those that have broad and variable settlement seasons and whose populations fluctuate substantially throughout the year in response to short-term fluctuations in recruitment. Comparisons of demographic effects of recruitment among species with different longevity require the use of non-arbitrary time scales, such as the time to maturity and the adult half-life.