Phylogenetic density dependence and environmental filtering predict seedling mortality in a tropical forest

Negative density dependence (NDD) and environmental filtering (EF) shape community assembly, but their relative importance is poorly understood. Recent studies have shown that seedling's mortality risk is positively related to the phylogenetic relatedness of neighbours. However, natural enemies, whose depredations often cause NDD, respond to functional traits of hosts rather than phylogenetic relatedness per se. To understand the roles of NDD and EF in community assembly, we assessed the effects on seedling mortality of functional similarity, phylogenetic relatedness and stem density of neighbouring seedlings and adults in a species-rich tropical forest. Mortality risks increased for common species when their functional traits departed substantially from the neighbourhood mean, and for all species when surrounded by close relatives. This indicates that NDD affects community assembly more broadly than does EF, and leads to the tentative conclusion that natural enemies respond to phylogenetically correlated traits. Our results affirm the prominence of NDD in structuring species-rich communities.     

Diversity of settlement-stage reef fishes captured by light-trap in a tropical south-west Atlantic Ocean coastal reef system

This study reports the results of 5 years of monitoring reef fish post-larvae using light traps in the Bay of Tamandaré, north-east Brazil. An annotated checklist of pre-settlement fish species, their frequency of occurrence and taxonomic characteristics are provided. In total, 4,422 post-larval fishes belonging to 36 families, 56 genera and 76 species were captured. The most species-rich families were Carangidae (7), Lutjanidae (6) and Pomacentridae (4), while the families Gerreidae (30.47%), Holocentridae (16.54%), Blenniidae (12.01%), Labrisomidae (8.36%), Lutjanidae (8.29%) and Acanthuridae (5.95%) were the most abundant. This is the first study of the taxonomic diversity and assemblage structure of settlement-stage reef fishes in the tropical south-west Atlantic Ocean. Although a few common species were not captured due to selectivity of light traps, the composition and taxonomic diversity of this first collection suggests that light traps are useful for studies of the early life history of a wide range of pre-settlement reef fishes.     

Global ecological success of Thalassoma fishes in extreme coral reef habitats

Phenotypic adaptations can allow organisms to relax abiotic selection and facilitate their ecological success in challenging habitats, yet we have relatively little data for the prevalence of this phenomenon at macroecological scales. Using data on the relative abundance of coral reef wrasses and parrotfishes (f. Labridae) spread across three ocean basins and the Red Sea, we reveal the consistent global dominance of extreme wave‐swept habitats by fishes in the genus Thalassoma, with abundances up to 15 times higher than any other labrid. A key locomotor modification—a winged pectoral fin that facilitates efficient underwater flight in high‐flow environments—is likely to have underpinned this global success, as numerical dominance by Thalassoma was contingent upon the presence of high‐intensity wave energy. The ecological success of the most abundant species also varied with species richness and the presence of congeneric competitors. While several fish taxa have independently evolved winged pectoral fins, Thalassoma appears to have combined efficient high‐speed swimming (to relax abiotic selection) with trophic versatility (to maximize exploitation of rich resources) to exploit and dominate extreme coral reef habitats around the world.     

The significance of variable and density-independent post-recruitment mortality in local populations of reef fishes

Abstract In this paper I focus on how post-settlement mortality may modify initial patterns of settlement in reef fish. Infrequent recruitment surveys may underestimate the role of early post-settlement mortality as most mortality in reef fishes occurs shortly after settlement. Consequently, results from infrequent recruitment surveys shed little light on the mechanisms producing patterns of abundance because these surveys ignore early post-settlement mortality. Variation in density-independent mortality may be a common mechanism that can prevent a positive relationship between larval settlement and subsequent population abundance. Although density-dependent mortality is the most commonly recognized mechanism that can disrupt the correlation between settlement and adult abundance, density-independent mortality’ can also destroy this correlation if the variance associated with post-settlement mortality is greater than variance in settlement. This point is illustrated with a simulation model in which I modelled two populations: a piscivorous fish population that was recruitment-limited with constant mortality, and a prey population that had variable recruitment and mortality that was a function of the size of the predator population. The results of this model indicate that even when mortality of prey is density-independent, predation can determine prey abundance when variation in piscivore recruitment is high relative to prey recruitment. Thus, initial patterns of prey settlement can be modified by a recruitment-limited predator population.     

Development of reproductive potential in protogynous coral reef fishes within Philippine no-take marine reserves

Empirical evidence for increases in the reproductive potential (egg output per unit area) of coral reef fish in no-take marine reserves (NTMRs) is sparse. Here, we inferred the development of reproductive potential in two species of protogynous reef fishes, Chlorurus bleekeri (Labridae: Scarinae) and Cephalopholis argus (Epinephelidae), inside and outside of Philippine NTMRs. We estimated key reproductive parameters and applied these to species-specific density and length data from 17 NTMRs (durations of protection 0–11 years) and paired fished sites (controls) in a space-for-time substitution approach. For C. argus, we also used density and length data collected almost annually over 29 years from a NTMR and an adjacent control at Apo Island. The results suggest that C. bleekeri can develop 6.0 times greater reproductive potential in NTMRs than controls after 11 years of protection, equivalent to approximately 582,000 more eggs produced 500 m⁻² inside NTMRs. Enhancement of reproductive potential in C. argus was not evident after 11 years in the space-for-time substitution. At Apo Island NTMR, reproductive potential of C. argus increased approximately 6-fold over 29 years but NTMR/control ratios in reproductive potential decreased through time (from 3.2 to 2.4), probably due to spillover of C. argus from the NTMR to the control. C. argus was estimated to produce approximately 113,000 more eggs 500 m⁻² inside Apo Island NTMR at the 29th year of protection. Ratios of reproductive potential between NTMR and controls in C. bleekeri and C. argus were often greater than corresponding ratios in density or biomass. The study underscores the importance of species-specific reproductive life history traits that drive variation in the development of larval fish subsidies that originate from NTMRs.     

Influences of wind-wave exposure on the distribution and density of recruit reef fishes at Kure and Pearl and Hermes Atolls,Northwestern Hawaiian Islands

This paper describes the results of a field survey designed to test the prediction that the density of benthic juveniles of shallow-reef fishes is greater on wind-wave “exposed” sectors of a pair of isolated oceanic atolls (Kure, Pearl and Hermes) at the far northwestern end of the Hawaiian Islands, an archipelago in which east-northeasterly trade winds dominate onshore water flow and transport by surface currents. The densities of recruits (juveniles ≤5 cm total length) were higher overall on windward versus leeward sectors of carbonate rock-rubble back reefs at both atolls, and the pattern was stronger for smaller (likely younger, more recently settled) recruits of four of the five most abundant species and the remainder pooled as an “Other” taxon. The windward-leeward disparity was four-fold greater at Pearl Hermes (the atoll with a three-fold longer perimeter) than at Kure. Resident predator biomass also was correlated with recruit densities, but habitat (benthic substratum) effects were generally weak. The distribution and abundance of recruits and juveniles of the primarily endemic reef fishes on shallow back reefs at these atolls appear partly influenced by relative rates of water flow over windward vs. leeward sectors of barrier reef and by the size, shape, and orientation of habitat parcels that filter out postlarval fishes with relatively weak swimming capabilities like labroids. Whole-reef geomorphology as well as fine-scale habitat heterogeneity and rugosity should be considered among the suite of many factors used to interpret observed spatial patterns of post-settlement juvenile fish distribution at atolls and perhaps some other tropical reefs. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The shape of abundance distributions across temperature gradients in reef fishes

Improving predictions of ecological responses to climate change requires understanding how local abundance relates to temperature gradients, yet many factors influence local abundance in wild populations. We evaluated the shape of thermal‐abundance distributions using 98 422 abundance estimates of 702 reef fish species worldwide. We found that curved ceilings in local abundance related to sea temperatures for most species, where local abundance declined from realised thermal ‘optima’ towards warmer and cooler environments. Although generally supporting the abundant‐centre hypothesis, many species also displayed asymmetrical thermal‐abundance distributions. For many tropical species, abundances did not decline at warm distribution edges due to an unavailability of warmer environments at the equator. Habitat transitions from coral to macroalgal dominance in subtropical zones also influenced abundance distribution shapes. By quantifying the factors constraining species’ abundance, we provide an important empirical basis for improving predictions of community re‐structuring in a warmer world.     

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.     

Elevational variation in density dependence in a subtropical forest

Density‐dependent mortality has been recognized as an important mechanism that underpins tree species diversity, especially in tropical forests. However, few studies have attempted to explore how density dependence varies with spatial scale and even fewer have attempted to identify why there is scale‐dependent differentiation. In this study, we explore the elevational variation in density dependence. Three 1‐ha permanent plots were established at low and high elevations in the Heishiding subtropical forest, southern China. Using data from 1200 1 m² seedling quadrats, comprising of 200 1 m² quadrats located in each 1‐ha plot, we examined the variation in density dependence between elevations using a generalized linear mixed model with crossed random effects. A greenhouse experiment also investigated the potential effects of the soil biota on density‐dependent differentiation. Our results demonstrated that density‐dependent seedling mortality can vary between elevations in subtropical forests. Species found at a lower elevation suffered stronger negative density dependence than those found at a higher elevation. The greenhouse experiment indicated that two species that commonly occur at both elevations suffered more from soilborne pathogens during seed germination and seedling growth when they grew at the lower elevation, which implied that soil pathogens may play a crucial role in density‐dependent spatial variation.     

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.     

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.     

Quantifying density dependence in a bird population using human disturbance

Although density dependence has long been recognised as vital to population regulation, there have been relatively few studies demonstrating it spatially in wildlife populations, often due to the confounding effects of variation in habitat quality. We report on a study of woodlarks Lullula arborea, a species of European conservation concern, breeding on lowland heath in Dorset, England. We take the novel approach of utilising the birds’ response to human disturbance, which resulted in much of the variation in density but had no direct impact on demographic rates. Within years, in sites with greater density there were smaller mean chick masses, lower post-fledging survival, and higher rates of nestling mortality attributed to starvation. The effects on clutch size and fledging success were confounded by the area of grassland within a site. There was no effect on brood size. Density dependence also operated within sites between years: as density increased there were reductions in mean chick mass and post-fledging survival, while nestling mortality attributed to starvation increased. Density-dependent effects on clutch size were only weakly regulatory, whereas density-dependent starvation and post-fledging mortality rates contributed strongly to differences in overall breeding output. Heavier chicks (when 7 days old) were significantly more likely to fledge and less likely to starve. Broods with heavier chicks were more likely to supply recruits to the breeding population. Nestling mass was not a factor in survival in the immediate post-fledging period, suggesting that density-dependent processes act independently on this stage. We conclude that the number of birds per hectare of suitable habitat is a valid means of expressing density, and that habitat acts as a surrogate for food abundance through which density dependence operates on the woodlark population.     

Intrageneric variation in antipredator responses of coral reef fishes affected by ocean acidification: implications for climate change projections on marine communities

Our planet is experiencing an increase in the concentration of atmospheric carbon dioxide (CO₂) unprecedented in the past 800 000 years. About 30% of excess atmospheric CO₂ is absorbed by the oceans, thus increasing the concentration of carbonic acid and reducing the ocean's pH. Species able to survive the physiological stress imposed by ocean acidification may still suffer strong indirect negative consequences. Comparing the tolerance of different species to dissolved CO₂ is a necessary first step towards predicting the ecological impacts of rising CO₂ levels on marine communities. While it is intuitive that not all aquatic species will be affected the same way by CO₂, one could predict that closely related species, sharing similar life histories and ecology, may show similar tolerance levels to CO₂. Our ability to create functional groups of species according to their CO₂ tolerance may be crucial in our ability to predict community change in the future. Here, we tested the effects of CO₂ exposure on the antipredator responses of four damselfish species (Pomacentrus chrysurus, Pomacentrus moluccensis, Pomacentrus amboinensis and Pomacentrus nagasakiensis). Although being sympatric and sharing the same ecology and life history, the four congeneric species showed striking and unexpected variation in CO₂ tolerance, with CO₂‐induced loss of response to predation risk ranging from 30% to 95%. Using P. chrysurus as a model species, we further tested if these behavioural differences translated into differential ability to survive predators under natural conditions. Our results indicate that P. chrysurus larvae raised under CO₂ levels predicted by 2070 and 2100 showed decreased antipredator responses to risk, leading to a five‐ to sevenfold increase in predation‐related mortality in the first few hours of settlement. Examining ocean acidification, along with other environmental variables, will be a critical step in further evaluating ecological responses to predicted climatic change.     

The replicability crisis in science and protected area research: Poor practices and potential solutions

The reliability of some published research from well-funded disciplines of medicine and psychology has been brought into question. This is because some researchers failed to achieve consistent results after replicating published studies using the same methodology. Researchers have referred to this as the ‘replicability in science crisis’ and have identified several practices contributing to unreliable science. Protected area and other conservation researchers are unlikely to be immune from these poor practices given they use the same scientific approaches as other disciplines. Fortunately, there are solutions to the poor practices contributing to unreliable science. In this paper I identify those poor practices and describe solutions as identified by researchers from a range of disciplines. These solutions are transferable to protected area science and related conservation disciplines. Most solutions are not costly or demanding to implement. Adopting these solutions can improve the reliability of both published and unpublished research.     

Comment arising from a paper by Wolda and Dennis: using and interpreting the results of tests for density dependence

We argue that tests for density dependence are useful in analyses of population dynamics and suggest guide lines for their use and interpretation of results which avoid many of the problems discussed by Wolda and Dennis (1993). Processes other than density dependence per se can cause statistical tests to indicate the presence of density dependence (Wolda and Dennis 1993 and unpublished simulations). Tests for density dependence cannot reveal the mechanism of regulation, but they do indicate the nature of long-term population dynamics. Tests for density dependence give misleading results if sampling is not at generation intervals; however, this problem is avoided if we only use tests on data collected in each generation (Holyoak 1993a). Similarly, species should be semelparous. Non-delayed density dependence should not be considered without looking for delayed density dependence, since the presence of delayed density dependence can lead to over-detection of non-delayed density dependence (Woiwod and Hanski 1992; Holyoak 1993b). The partial autocorrelation function and knowledge of life-history are more useful than tests for density dependence for indicating whether any density dependence is delayed or not (Royama 1992; Holyoak 1993b). Estimation error with a constant upper size limit causes tests for density dependence to overestimate the frequency of delayed density dependence; however we do not know whether estimation error is bounded in real populations. Work in progress suggests that 20–40 generations (depending on the nature of population dynamics) gives a moderate level of accuracy with tests for density dependence, and >40 generations are necessary for tests to be accurate in their assessment of the strength of density dependence. We conclude that tests are useful indicators of whether density dependence, or other feedback mechanisms are likely to be acting.     

Non‐reef habitats in a tropical seascape affect density and biomass of fishes on coral reefs

Nonreef habitats such as mangroves, seagrass, and macroalgal beds are important for foraging, spawning, and as nursery habitat for some coral reef fishes. The spatial configuration of nonreef habitats adjacent to coral reefs can therefore have a substantial influence on the distribution and composition of reef fish. We investigate how different habitats in a tropical seascape in the Philippines influence the presence, density, and biomass of coral reef fishes to understand the relative importance of different habitats across various spatial scales. A detailed seascape map generated from satellite imagery was combined with field surveys of fish and benthic habitat on coral reefs. We then compared the relative importance of local reef (within coral reef) and adjacent habitat (habitats in the surrounding seascape) variables for coral reef fishes. Overall, adjacent habitat variables were as important as local reef variables in explaining reef fish density and biomass, despite being fewer in number in final models. For adult and juvenile wrasses (Labridae), and juveniles of some parrotfish taxa (Chlorurus), adjacent habitat was more important in explaining fish density and biomass. Notably, wrasses were positively influenced by the amount of sand and macroalgae in the adjacent seascape. Adjacent habitat metrics with the highest relative importance were sand (positive), macroalgae (positive), and mangrove habitats (negative), and fish responses to these metrics were consistent across fish groups evaluated. The 500‐m spatial scale was selected most often in models for seascape variables. Local coral reef variables with the greatest importance were percent cover of live coral (positive), sand (negative), and macroalgae (mixed). Incorporating spatial metrics that describe the surrounding seascape will capture more holistic patterns of fish–habitat relationships on reefs. This is important in regions where protection of reef fish habitat is an integral part of fisheries management but where protection of nonreef habitats is often overlooked.