Jennifer E. Caselle

Variability in recruitment of coral reef fishes : The importance of habitat at two spatial scales

We investigated spatial and temporal variability in recruitment of coral reef fishes to St. Croix, U.S. Virgin Islands, with the goal of assessing the importance of habitat and physical oceanographic processes in determining patterns of distribution at two spatial scales. Recruitment was monitored visually each month, using SCUBA, at sites spaced equally around the island and on transects within each site. Spatial patterns of distribution were consistent within seasons and between years at both spatial scales. Although recruitment showed a seasonal pattern, there were few inter-annual differences at any spatial scale. At the largest scale, recruitment varied significantly among sites around the island, and a unique pattern of distribution was observed. On the windward shore, recruitment ranged from high on the eastern (upcurrent) end to low on western end of the island. Recruitment showed the opposite pattern on the leeward shore, ranging from low on the eastern end to high on the western end. At the transect scale, certain features of the habitat significantly predicted recruitment density. None of the microhabitat features that predicted recruitment at the small scale could explain patterns of recruitment observed at the large scale. In contrast, physical processes affecting larval availability or transport were more important than habitat in determining the spatial patterns of recruitment at this scale. Thus, physical oceanographic processes appeared to be responsible for patterns of recruitment to the two shores of St. Croix, and to sites along those shores. Despite consistent patterns of recruitment to sites, habitat selection did not appear to be important at the site scale. Once delivered to a site, however, settlers or new recruits distributed themselves based on habitat preferences.

Scale-dependent effects of habitat on movements and path structure of reef sharks at a predator-dominated atoll

The effects of habitat on the ecology, movements, and foraging strategies of marine apex predators are largely unknown. We used acoustic telemetry to quantify the movement patterns of blacktip reef sharks (Carcharhinus melanopterus) at Palmyra Atoll National Wildlife Refuge, in the Pacific Ocean. Sharks had relatively small home ranges over a timescale of days to weeks (0.55 +/- 0.24 km2) and showed strong site fidelity to sand-flat ledges within the west lagoon over a three-year period. Sharks showed evidence of diel and tidal movements, and they utilized certain regions of the west lagoon disproportionately. There were ontogenetic shifts in habitat selection, with smaller sharks showing greater selection for sand-flat habitats, and pups (total length 35-61 cm) utilizing very shallow waters on sand-flats, potentially as nursery areas. Adult sharks selected ledge habitats and had lower rates of movement when over sand-flats and ledges than they did over lagoon waters. Fractal analysis of movements showed that over periods of days, sharks used patches that were 3-17% of the scale of their home range. Repeat horizontal movements along ledge habitats consisted of relatively straight movements, which theoretical models consider the most efficient search strategy when forage patches may be spatially and temporally unpredictable. Although sharks moved using a direct walk while in patches, they appeared to move randomly between patches. Microhabitat quantity and quality had large effects on blacktip reef shark movements, which have consequences for the life-history characteristics of the species and potentially the spatial distribution of behaviorally mediated effects on lower trophic levels throughout the Palmyra ecosystem.

Incorporating biogeography into evaluations of the Channel Islands marine reserve network

Networks of marine reserves are increasingly a major component of many ecosystem-based management plans designed to conserve biodiversity, protect the structure and function of ecosystems, and rebuild and sustain fisheries. There is a growing need for scientific guidance in the design of network-wide monitoring programs to evaluate the efficacy of reserves at meeting their conservation and management goals. Here, we present an evaluation of the Channel Islands reserve network, which was established in 2003 off the coast of southern California. This reserve network spans a major environmental and biogeographic gradient, making it a challenge to assess network-wide responses of many species. Using fish community structure data from a long-term, large-scale monitoring program, we first identified persistent geographic patterns of community structure and the scale at which sites should be grouped for analysis. Fish communities differed most among islands with densities of individual species varying from 3- to 250-fold. Habitat structure differed among islands but not based on reserve status. Across the network, we found that, after 5 years, species targeted by fishing had higher densities (1.5×) and biomass (1.8×) inside reserves, whereas nontargeted species showed no significant differences. Examining trophic groups, piscivore and carnivore biomass was significantly greater inside reserves (1.8× and 1.3× more, respectively), whereas the biomass of planktivores and herbivores was similar inside and out. A framework for incorporating biogeographic variation into reserve network assessments is critical as we move from the evaluation of single reserves to networks of reserves.

SIZE-SELECTIVE HARVESTING ALTERS LIFE HISTORIES OF A TEMPERATE SEX-CHANGING FISH

Selective mortality, whether caused naturally by predation or through the influence of harvest practices, initiates changes within populations when individuals possessing certain heritable traits have increased fitness. Theory predicts that increased mortality rates will select for changes in a number of different life history characteristics. For example, fishing often targets larger individuals and has been shown repeatedly to alter population size structure and growth rates, and the timing of maturation. For sex-changing species, selective fishing practices can affect additional traits such as the mature population sex ratio and the timing of sexual transformation. Using historical comparisons, we examined the effects of exploitation on life history characteristics of California sheephead, Semicossyphus pulcher, a temperate protogynous (female-male sex changer) labrid that inhabits nearshore rocky environments from central California, USA, to southern Baja California, Mexico. Recreational fishing intensified and an unregulated commercial live-fish fishery developed rapidly in southern California between the historical and current studies. Collections of S. pulcher from three locations (Bahia Tortugas, Catalina Island, and San Nicolas Island) in 1998 were compared with data collected 20-30 years previously to ascertain fishery-induced changes in life history traits. At Bahia Tortugas, where fishing by the artisanal community remained light and annual survivorship stayed high, we observed no changes in size structure or shifts in the timing of maturation or the timing of sex change. In contrast, where recreational (Catalina) and commercial (San Nicolas) fishing intensified and annual survivorship correspondingly declined, males and females shifted significantly to smaller body sizes, females matured earlier and changed sex into males at both smaller sizes and younger ages and appeared to have a reduced maximum lifespan. Mature sex ratios (female:male) increased at San Nicolas, despite a twofold reduction in the mean time spent as a mature female. Proper fisheries management requires measures to prevent sex ratio skew, sperm limitation, and reproductive failure because populations of sequential hermaphrodites are more sensitive to size-selective harvest than separate-sex species. This is especially true for S. pulcher, where different segments of the fishery (commercial vs. recreational) selectively target distinct sizes and therefore sexes in different locations.

EARLY POST‐SETTLEMENT MORTALITY IN A CORAL REEF FISH AND ITS EFFECT ON LOCAL POPULATION SIZE

The relative importance of recruitment and post-recruitment factors in controlling population fluctuations and determining population sizes in open marine systems is still poorly understood. To address this issue, I conducted an experiment investigating whether very early post-recruitment mortality of a coral reef fish, Thalassoma bifasciatum, is dependent on recruitment density. To quantify the extent to which local populations are likely to be controlled primarily by recruitment or by post-settlement density-dependent mortality, I also measured natural mortality rates and recruitment variation at sites around the island of St. Croix, U.S. Virgin Islands. I manipulated densities of 1–3-d post-settlement T. bifasciatum on large natural patch reefs in Tague Bay, St. Croix, U.S. Virgin Islands. Manipulated recruits were followed for ∼30 d. The loss rate over the initial 24-h period following placement of recruits on the reef was positively related to the density at the beginning of that period. After this period, initial recruit density had no effects on mortality. Natural monthly mortality rates approximating the monthly mortality monitored during the experiment were measured for naturally recruited fish at sites around the island. Mortality was positively correlated with recruitment density. The effect was strong enough that across sites the relationship between recruit density and juvenile density the following month was not proportional; that is, juvenile density tended to reach an asymptote with increasing recruitment. The relationship between total recruitment per season at a site and adult abundance at the end of the season was similarly asymptotic, indicating the persistence of a density-dependent effect at the scale of the island. Mortality rates around the island were extremely variable and density independent at densities below ∼1 recruit/m2. Above that level, mortality rates were higher and tended to increase with further increases in recruitment density. Around St. Croix, three out of eight sites consistently received recruitment above this “threshold,” while five sites never experienced recruitment above that level during the study. Because spatial patterns of recruitment around St. Croix were consistent over time, some local populations may be controlled mainly by recruitment and density-independent mortality at all times, while others may be chronically affected by post-settlement density dependence. This study gives support to the idea that recruitment limitation and density dependence are not mutually exclusive. Rather, local populations must be characterized by the degree of recruitment, the resultant population density, and the subsequent degree of density-dependent regulation.

Global patterns of kelp forest change over the past half-century

Significance Kelp forests support diverse and productive ecological communities throughout temperate and arctic regions worldwide, providing numerous ecosystem services to humans. Literature suggests that kelp forests are increasingly threatened by a variety of human impacts, including climate change, overfishing, and direct harvest. We provide the first globally comprehensive analysis of kelp forest change over the past 50 y, identifying a high degree of variation in the magnitude and direction of change across the geographic range of kelps. These results suggest region-specific responses to global change, with local drivers playing an important role in driving patterns of kelp abundance. Increased monitoring aimed at understanding regional kelp forest dynamics is likely to prove most effective for the adaptive management of these important ecosystems. Kelp forests (Order Laminariales) form key biogenic habitats in coastal regions of temperate and Arctic seas worldwide, providing ecosystem services valued in the range of billions of dollars annually. Although local evidence suggests that kelp forests are increasingly threatened by a variety of stressors, no comprehensive global analysis of change in kelp abundances currently exists. Here, we build and analyze a global database of kelp time series spanning the past half-century to assess regional and global trends in kelp abundances. We detected a high degree of geographic variation in trends, with regional variability in the direction and magnitude of change far exceeding a small global average decline (instantaneous rate of change = −0.018 y−1). Our analysis identified declines in 38% of ecoregions for which there are data (−0.015 to −0.18 y−1), increases in 27% of ecoregions (0.015 to 0.11 y−1), and no detectable change in 35% of ecoregions. These spatially variable trajectories reflected regional differences in the drivers of change, uncertainty in some regions owing to poor spatial and temporal data coverage, and the dynamic nature of kelp populations. We conclude that although global drivers could be affecting kelp forests at multiple scales, local stressors and regional variation in the effects of these drivers dominate kelp dynamics, in contrast to many other marine and terrestrial foundation species.

Geographic variation in density, demography, and life history traits of a harvested, sex-changing, temperate reef fish

Geographic variation in ecological and environmental factors may lead to intraspecific differences among populations. For the California sheephead (Semicossyphus pulcher), an important predator in kelp forests and a target of commercial and recreational fisheries, we evaluated the degree to which different populations exhibited variation in density, demography, and life history traits. We assessed biogeographic patterns of abundance through underwater visual census at 39 sites spanning a major portion of the species range (southern California, USA, to Baja California, Mexico) and made collections from seven focal sites to investigate geographic differences in demography and life histories. California sheephead densities were significantly greater in the southern part of their range and at offshore islands than along the mainland coast. At the focal sites, we found significant spatial variation in density, fecundity, size structure, growth rates, annual survivorship, and the timing of maturation and sex ch...

Comparative analyses of animal-tracking data reveal ecological significance of endothermy in fishes

Significance Most fishes are cold-blooded, but tunas and some sharks (e.g., white sharks) maintain their exercising muscles warmer than ambient waters. This ability is a remarkable example of convergent evolution because bony and cartilaginous fishes diverged as long as 450 million years ago. What are the ecological benefits driving the evolution of warm muscles in fishes? Despite extensive discussion, no previous studies have tested a simple possibility that fishes with warm muscles may swim faster in nature. We demonstrate that fishes with warm muscles swim faster and perform larger-scale annual migrations than similar-sized cold-blooded fishes. Our results suggest that warm muscles enhance power output and, thus, cruising speeds, which may enable longer-distance migrations and potentially greater access to seasonally available resources. Despite long evolutionary separations, several sharks and tunas share the ability to maintain slow-twitch, aerobic red muscle (RM) warmer than ambient water. Proximate causes of RM endothermy are well understood, but ultimate causes are unclear. Two advantages often proposed are thermal niche expansion and elevated cruising speeds. The thermal niche hypothesis is generally supported, because fishes with RM endothermy often exhibit greater tolerance to broad temperature ranges. In contrast, whether fishes with RM endothermy cruise faster, and achieve any ecological benefits from doing so, remains unclear. Here, we compiled data recorded by modern animal-tracking tools for a variety of free-swimming marine vertebrates. Using phylogenetically informed allometry, we show that both cruising speeds and maximum annual migration ranges of fishes with RM endothermy are 2–3 times greater than fishes without it, and comparable to nonfish endotherms (i.e., penguins and marine mammals). The estimated cost of transport of fishes with RM endothermy is twice that of fishes without it. We suggest that the high energetic cost of RM endothermy in fishes is offset by the benefit of elevated cruising speeds, which not only increase prey encounter rates, but also enable larger-scale annual migrations and potentially greater access to seasonally available resources.

Distribution, size frequency, and sex ratios of blacktip reef sharks Carcharhinus melanopterus at Palmyra Atoll: a predator-dominated ecosystem

Blacktip reef sharks Carcharhinus melanopterus were the most abundant predator in the lagoons at Palmyra Atoll. They were evenly distributed throughout the lagoons, although there was some evidence of sexual segregation. Males reach sexual maturity between 940-1,020 mm L(T). Bird remains were found in some C. melanopterus stomachs. C. melanopterus at Palmyra appear to be smaller than those at other locations.

Fishers' behaviour in response to the implementation of a Marine Protected Area.

Marine Protected Areas (MPAs) have been widely proposed as a fisheries management tool in addition to their conservation purposes. Despite this, few studies have satisfactorily assessed the dynamics of fishers’ adaptations to the loss of fishing grounds. Here we used data from before, during and after the implementation of the management plan of a temperate Atlantic multiple-use MPA to examine the factors affecting the spatial and temporal distribution of different gears used by the artisanal fishing fleet. The position of vessels and gear types were obtained by visual surveys and related to spatial features of the marine park. A hotspot analysis was conducted to identify heavily utilized patches for each fishing gear and time period. The contribution of individual vessels to each significant cluster was assessed to better understand fishers’ choices. Different fisheries responded differently to the implementation of protection measures, with preferred habitats of target species driving much of the fishers’ choices. Within each fishery, individual fishers showed distinct strategies with some operating in a broader area whereas others kept preferred territories. Our findings are based on reliable methods that can easily be applied in coastal multipurpose MPAs to monitor and assess fisheries and fishers responses to different management rules and protection levels. This paper is the first in-depth empirical study where fishers’ choices from artisanal fisheries were analysed before, during and after the implementation of a MPA, thereby allowing a clearer understanding of the dynamics of local fisheries and providing significant lessons for marine conservation and management of coastal systems.