Habitat structure,conspecific presence and spatial variation in the recruitment of a temperate reef fish

Pronounced spatial variation in recruitment occurs in many marine invertebrate and fish populations and is thought to be critical to the demography of these species. In this study I examined the importance of habitat structure and the presence of conspecific residents to spatial variation in larval settlement and recruitment in a temperate fish Tautogolabrus adspersus. I define settlement as the movement of individuals from the water column to the benthic habitat, while I refer to recruitment as numbers of individuals surviving some arbitrary period of time after settlement. Experiments in which standard habitats were stocked with conspecifics showed that resident conspecifics were not an important factor contributing to small-scale variability in recruitment. Further correlative analyses demonstrated that large-scale variation in recruitment could not be explained by variability in older age classes. By contrast, manipulations of macroalgal structure within a kelp bed demonstrated that recruitment was significantly higher in habitats with a dense understory of foliose and filamentous algae than in habitats with only crustose algae. Understory algae varied in their pattern of disperison among sites, and the dispersion of fish matched that of the plants. In order to determine the effects of differences in patterns of algal dispersion on the demography of associated T. adspersus populations, I used experimental habitat units to manipulate patterns of dispersion. Settlement was significantly greater to randomly placed versus clumped habitats; however, no differences in recruitment between random and clumped habitats were detected. Because recruitment is a function of the numbers of settlers minus the subsequent loss of settlers, rates of mortality or migration must have been higher in the randomly placed habitats. These results are counter to the current paradigm for reef fishes which suggests that larval settlement is the crucial demographic process producing variability in population abundance. In this experiment patterns of settlement were modified by varying the patch structure of the habitat.Contribution number 278 from the Center for Marine Biology, University of New Hampshire     

Interspecific variation in life history relates to antipredator decisions by marine mesopredators on temperate reefs

As upper-level predatory fishes become overfished, mesopredators rise to become the new 'top' predators of over-exploited marine communities. To gain insight into ensuing mechanisms that might alter indirect species interactions, we examined how behavioural responses to an upper-level predatory fish might differ between mesopredator species with different life histories. In rocky reefs of the northeast Pacific Ocean, adult lingcod (Ophiodon elongatus) are upper-level predators that use a sit-and-wait hunting mode. Reef mesopredators that are prey to adult lingcod include kelp greenling (Hexagrammos decagrammus), younger lingcod, copper rockfish (Sebastes caurinus) and quillback rockfish (S. maliger). Across these mesopredators species, longevity and age at maturity increases and, consequently, the annual proportion of lifetime reproductive output decreases in the order just listed. Therefore, we hypothesized that the level of risk taken to acquire resources would vary interspecifically in that same order. During field experiments we manipulated predation risk with a model adult lingcod and used fixed video cameras to quantify interactions between mesopredators and tethered prey (Pandalus shrimps). We predicted that the probabilities of inspecting and attacking tethered prey would rank from highest to lowest and the timing of these behaviours would rank from earliest to latest as follows: kelp greenling, lingcod, copper rockfish, and quillback rockfish. We also predicted that responses to the model lingcod, such as avoidance of interactions with tethered prey, would rank from weakest to strongest in the same order. Results were consistent with our predictions suggesting that, despite occupying similar trophic levels, longer-lived mesopredators with late maturity have stronger antipredator responses and therefore experience lower foraging rates in the presence of predators than mesopredators with faster life histories. The corollary is that the fishery removal of top predators, which relaxes predation risk, could potentially lead to stronger increases in foraging rates for mesopredators with slower life histories.     

Integrating life history and physiology to understand latitudinal size variation in a damselfly

Our understanding of latitudinal life history patterns may benefit by jointly considering age and mass at maturity and growth rate. Additional insight may be gained by exploring potential constraints through pushing growth rates to their maximum and scoring physiological cost‐related variables. Therefore, we reared animals of a univoltine Spanish and Belgian population and of a semivoltine Swedish population of the damselfly Enallagma cyathigerum (spanning a latitude gradient of ca 2350 km) in a common environment from the eggs until adult emergence and exposed them to a transient starvation period to induce compensatory growth. Besides age and mass at maturity and growth rate we also scored investment in energy storage (i.e. triglycerides) and immune function (i.e. total activity of phenoloxidase). At emergence, body mass was greater in Spain and Sweden and lower in Belgium, suggesting a genetic component for the U‐shaped latitudinal pattern that was found also in a previous study based on field‐collected adults. The mass difference between univoltine populations can be explained by the shorter development time in the Belgian population, and this despite a higher growth rate, a pattern consistent with undercompensating countergradient variation. In line with the assumed shorter growth seasons, Belgian and Swedish animals showed higher routine growth rates and compensatory growth after transient starvation. Despite a strong link with metabolic rates (as measured by oxygen consumption) populations with higher routine growth rates had no lower fat content and had higher immune function (i.e. immune function decreased from Sweden to Spain), which was unexpected. Rapid compensatory growth did, however, result in a lowered immune function. This may contribute to the absence of perfect compensating countergradient variation in the Belgian population and the lowest routine growth rates in the Spanish population. Our results underscore the importance of integrating key life historical with physiological traits for understanding latitudinal population differentiation.     

Community structure in north temperate ants: temporal and spatial variation

Summary Ant communities in Vermont and New York woods were sampled in four time periods to determine species composition, relative abundances, and nest locations in space. The Vermont community was richer, containing more species and higher nest densities than New York. Both communities followed the geometric distribution of species abundances, suggesting that a single resource was mediating competition. The resource most clearly implicated was suitable nest sites, principally pre-formed plant cavities. Nonrandom species associations, underdispersion in every season, and the occurrence of incipient nests overwintering aboveground all implicated shortage of such cavities. Furthermore, microhabitat differences which produce suitable nest sites occur over a very small scale in these communities.     

Small-scale recruitment variation in a temperate fish: the roles of macrophytes and food supply

Synopsis The availability of reef-related resources, particularly food and shelter can play a significant role in determining the distribution and abundance of reef fishes. Much of the structure on temperate reefs is provided by macroalgae, and variability in the density of temperate reef fishes at large spatial scales (100's of meters) can often be explained by variation in macroalgal cover or density. In this study I investigated the role of macrophytes and associated food resources on the recruitment of a temperate fish, Tautogolabrus adspersus, at a small spatial scale (0.25 m²). No relationship between the density of new recruits and the percent cover of kelp, foliose or filamentous algae was observed. Multiple regressions revealed that less than 8% of variability in recruitment could be explained by variability in macroalgal cover. However, recruits were found in higher abundance in patches containing many functional forms of seaweeds than in patches dominated by a single form. A wide variety of prey were available for use by cunner recruits; however, crustaceans and mussels were the only common components of their diet, and crustaceans were clearly the most preferred prey. The prey composition in patches where fish were present was compared to randomly selected patches. Significantly greater numbers of isopods, amphipods and newly settled mussels were present in patches where fish were present than in randomly selected patches. The data presented in this study contradict previous work that has shown algal structure to be important in determining patterns of abundance and food supply to be of little significance. A conceptual model is proposed suggesting that settling fish select habitats in a hierarchical manner largely based on their dispersal tendencies. Hierarchical selection of habitats results in different attributes of the habitat being selected during different life-history intervals.     

Functional genomics of life history variation in a butterfly metapopulation

In fragmented landscapes, small populations frequently go extinct and new ones are established with poorly understood consequences for genetic diversity and evolution of life history traits. Here, we apply functional genomic tools to an ecological model system, the well-studied metapopulation of the Glanville fritillary butterfly. We investigate how dispersal and colonization select upon existing genetic variation affecting life history traits by comparing common-garden reared 2-day adult females from new populations with those from established older populations. New-population females had higher expression of abdomen genes involved in egg provisioning and thorax genes involved in the maintenance of flight muscle proteins. Physiological studies confirmed that new-population butterflies have accelerated egg maturation, apparently regulated by higher juvenile hormone titer and angiotensin converting enzyme mRNA, as well as enhanced flight metabolism. Gene expression varied between allelic forms of two metabolic genes (Pgi and Sdhd), which themselves were associated with differences in flight metabolic rate, population age and population growth rate. These results identify likely molecular mechanisms underpinning life history variation that is maintained by extinction-colonization dynamics in metapopulations.     

A social basis for the development of primary males in a sex-changing fish

An example of alternative male strategies is seen in diandric protogynous (female first) hermaphrodites, where individuals either mature directly as male (primary males) or first reproduce as female and then change sex to male (secondary males). In some sex-changing fishes, the testes of primary males appear anatomically similar to those of non-sex-changing species, whereas the testes of secondary males have anatomical evidence of their former ovarian function. Here, we provide evidence that in the bluehead wrasse, Thalassoma bifasciatum, these strikingly different male phenotypes arise from differences in the ontogenetic timing of environmental sex determination, timing that can be experimentally altered through changes in the social circumstances. Juveniles differentiated almost exclusively as females when reared in isolation, regardless of whether they were collected from a reef with a high proportion of primary males or from a reef with a low proportion of primary males. In contrast, one individual usually differentiated as a primary male when reared in groups of three. Our results indicate that primary males of the bluehead wrasse are an environmentally sensitive developmental strategy that has probably evolved in response to variation in the reproductive success of primary males in populations of different sizes.     

Immune indexes of larks from desert and temperate regions show weak associations with life history but stronger links to environmental variation in microbial abundance

Abstract Immune defense may vary as a result of trade-offs with other life-history traits or in parallel with variation in antigen levels in the environment. We studied lark species (Alaudidae) in the Arabian Desert and temperate Netherlands to test opposing predictions from these two hypotheses. Based on their slower pace of life, the trade-off hypothesis predicts relatively stronger immune defenses in desert larks compared with temperate larks. However, as predicted by the antigen exposure hypothesis, reduced microbial abundances in deserts should result in desert-living larks having relatively weaker immune defenses. We quantified host-independent and host-dependent microbial abundances of culturable microbes in ambient air and from the surfaces of birds. We measured components of immunity by quantifying concentrations of the acute-phase protein haptoglobin, natural antibody-mediated agglutination titers, complement-mediated lysis titers, and the microbicidal ability of whole blood. Desert-living larks were exposed to significantly lower concentrations of airborne microbes than temperate larks, and densities of some bird-associated microbes were also lower in desert species. Haptoglobin concentrations and lysis titers were also significantly lower in desert-living larks, but other immune indexes did not differ. Thus, contrary to the trade-off hypothesis, we found little evidence that a slow pace of life predicted increased immunological investment. In contrast, and in support of the antigen exposure hypothesis, associations between microbial exposure and some immune indexes were apparent. Measures of antigen exposure, including assessment of host-independent and host-dependent microbial assemblages, can provide novel insights into the mechanisms underlying immunological variation.     

Role of temperature in regulation of the life cycle of temperate fish

The review provides information on adaptive responses of fish metabolic processes in response to temperature reduction of habitat, including the direction of adjustment of energy metabolism and mechanisms to improve the adaptive capacity and reparative capacity of tissues in response to lower temperatures. It presents data on the effect of temperature on the reproductive function of fish and fish development processes in the early stages of ontogeny.     

Ontogenetic and spatial variation in size‐selective mortality of a marine fish

Although body size can affect individual fitness, ontogenetic and spatial variation in the ecology of an organism may determine the relative advantages of size and growth. During an 8‐year field study in the Bahamas, we examined selective mortality on size and growth throughout the entire reef‐associated life phase of a common coral‐reef fish, Stegastes partitus (the bicolour damselfish). On average, faster‐growing juveniles experienced greater mortality, though as adults, larger individuals had higher survival. Comparing patterns of selection observed at four separate populations revealed that greater population density was associated with stronger selection for larger adult size. Large adults may be favoured because they are superior competitors and less susceptible to gape‐limited predators. Laboratory experiments suggested that selective mortality of fast‐growing juveniles was likely because of risk‐prone foraging behaviour. These patterns suggest that variation in ecological interactions may lead to complex patterns of lifetime selection on body size.     

Complexity and variation in loggerhead sea turtle life history

Juvenile loggerhead sea turtles spend more than a decade in the open ocean before returning to neritic waters to mature and reproduce. It has been assumed that this transition from an oceanic to neritic existence is a discrete ontogenetic niche shift. We tested this hypothesis by tracking the movements of large juveniles collected in a neritic foraging ground in North Carolina, USA. Our work shows that the shift from the oceanic to neritic waters is both complex and reversible; some individuals move back into coastal waters and then return to the open ocean for reasons that are still unclear, sometimes for multiple years. These findings have important consequences for efforts to protect these threatened marine reptiles from mortality in both coastal and open-ocean fisheries.     

Dimensionless numbers and life history variation in Brown Trout

Summary Dimensionless numbers, made up from components of life history as defined by growth, mortality and maturation, may provide fresh insights into life history evolution. Most studies have previously shown that these numbers are more or less constants within taxa. The variation between taxa may clarify the evolution of different life histories. We examine the variation in three dimensionless numbers using data from 29 populations of Brown TroutSalmo trutta from Norway, and find that the dimensionless numbers are not constants for the Brown Trout populations. We find that the relationship betweenK of the von Bertalanffy growth equation and the mortality rate (M) increased with increasing growth rate. Also, relative length at maturity (L /L _inf) increased with increasing asymptotic length (L _inf). We suggest that more such data should be collected from a large number of species and taxonomic groups, to allow a more detailed assessment of why these dimensionless numbers appear to be constants in some taxa and not in others.     

Rocky intertidal fish communities of California: temporal and spatial variation

Synopsis We examined data from our own and published collections of intertidal and shallow littoral fishes of the North American Pacific Coast with respect to temporal and spatial trends in species composition and dominance. We compared (1) recent and past intertidal collections made five and seven years apart, respectively, for two California localities, (2) intertidal collections from twelve localities in California, Oregon, and British Columbia, and (3) intertidal versus subtidal collections at one California locality. Temporal comparisons indicated substantially lower abundance of the cottid Oligocottus snyderi at two California localities during 1984, at one locality due to depressed abundance of young. Interannual differences in abundances suggested that other tidepool fish assemblages undergo significant changes as well. Geographical comparisons indicated general similarity in species composition, with cottids predominating in tidepools although several other families also were well represented (e.g., Stichaeidae, Scorpaenidae). Eel-shaped stichaeids and pholids occurred at high densities in exposed boulder fields. Both the tidepool and boulder field assemblages showed north-south changes in species abundances. Comparison of collections from the intertidal and subtidal zones at one California locality demonstrated that fishes of these habitats form two essentially distinct assemblages, with most species restricted to or concentrated in one or the other habitat.     

The genomic and physiological basis of life history variation in a butterfly metapopulation

Unravelling the mechanisms underlying variation in life history traits is of fundamental importance for our understanding of adaptation by natural selection. While progress has been made in mapping fitness-related phenotypes to genotypes, mainly in a handful of model organisms, functional genomic studies of life history adaptations are still in their infancy. In particular, despite a few notable exceptions, the genomic basis of life history variation in natural populations remains poorly understood. This is especially true for the genetic underpinnings of life history phenotypes subject to diversifying selection driven by ecological dynamics in patchy environments--as opposed to adaptations involving strong directional selection owing to major environmental changes, such as latitudinal gradients, extreme climatic events or transitions from salt to freshwater. In this issue of Molecular Ecology,Wheat et al. (2011) now make a significant leap forward by applying the tools of functional genomics to dispersal-related life history variation in a butterfly metapopulation. Using a combination of microarrays, quantitative PCR and physiological measurements, the authors uncover several metabolic and endocrine factors that likely contribute to the observed life history phenotypes. By identifying molecular candidate mechanisms of fitness variation maintained by dispersal dynamics in a heterogeneous environment,they also begin to address fascinating interactions between the levels of physiology, ecology and evolution.     

Large scale spatial and temporal variation in recruitment to fish populations on coral reefs

Summary Visual census was used to sample young of the year of fish species recruited to each of two habitats on seven lagoonal platform reefs of the Capricorn-Bunker Group, Great Barrier Reef. The reefs sampled span an area 70 km in extent. In 1983, 62 species from 13 families were detected as recruits on reef slope sites. The total number of cruits, and the number of each of 6 of 16 species tested, differed significantly among reefs, despite the fact that differences among sites within reefs did not exist, and that sampled slopes were chosen to be hydrographically, and physiographically as similar as possible. Lagoonal patch reefs were sampled in two years. In 1982, 76 species of 11 families occurred as recruits. In 1983, 86 species of 12 families were recorded. All of 22 species common enough to test showed some significant variation in abundance among reefs, years, or both. For 9 species, significant year x reef interactions occurred, demonstrating that relative recruitment success among reefs varied between years. Reasons for the substantial levels of variability are discussed, and implications for the organisation of reef fish communities are considered.     

The effect of landscape composition and configuration on the spatial distribution of temperate demersal fish

In contrast to the terrestrial environment, where the use of landscape analyses has been clearly demonstrated, the influence of landscape composition and configuration on the abundance and spatial distribution of marine organisms remains poorly understood. Development of this area of marine research has been limited by the lack of accurate benthic habitat maps, particularly for marine environments deeper than can be penetrated by optical remote sensing (<10 m). However, the recent availability of detailed (1:25 000) and accurate habitat maps derived from hydroacoustic surveys of deeper marine waters has redressed this situation. The aim of this research was to establish the strength and significance of derived landscape composition and configuration indices on the demersal fish assemblage structure and spatial distribution. A combination of depth and 6 landscape measures were found to explain 34.8% of the variation in the fish assemblage. Depth contributed the highest percentage of the variance explained, accounting for 23.7%. Distance to reef was the most important landscape index explaining 7.1% of the variability, followed by total length of edge environment (the interface between different benthic substrates) explaining 1.5%. Species responsible for driving the structure of the fish assemblage were also examined and found to respond not only to the proximity of reef but also to the configuration of the reef. For example, Parika scaber and Pseudolabrus psittaculus characterised more heterogeneous landscapes offering clusters of small interconnected patches of reef with a lot of edge environment, whilst juvenile Trachyurus novaezelandiae and Platycephalus caeruleopunctatus preferred more homogeneous landscapes offering little reef or edge environment. This research demonstrated that a broad scale landscape analysis, employing indices of landscape composition and configuration, is important for understanding demersal fish assemblage structure and spatial distribution.     

Quantifying structural variation in contour feathers to address functional variation and life history trade‐offs

Body feathers are important to many interactions birds have with their physical and social environments, such as streamlining the body for flight, thermoregulation, and social signaling. Birds differ dramatically in the texture of their body plumage depending on species and age class, likely reflecting different functional demands and age‐related trade‐offs in feather production. Despite the important insights potentially offered by studying variation in the structure of body feathers, there is no clear system for quantifying this variation. We present methods for quantifying age and species differences in the structure of body feathers. Most variation in our measures is due to species and age‐class differences, with little variance attributable to individual birds or to differences among feathers sampled from the same bird. We use our measures to test the hypothesis that the loosely‐textured plumage characteristic of many juvenile passerines reflects a trade‐off between investment in feather quality and rapid body growth that promotes early fledging. The structure of juvenile feathers was negatively correlated with duration of the nestling period among ten species of New World warblers (Parulidae), suggesting a trade‐off between investment in feathers and investment in rapid somatic development promoting fledging. Systematic studies of variation in the structure of body feathers will likely offer numerous other insights into avian biology.     

Resolution of the early life history of a reef fish using otolith chemistry

We used the elemental signatures in otoliths of the damselfish Pomacentrus coelestis as a proxy for conditions experienced prior to settlement. Fish from the southern Great Barrier Reef (GBR) differed in their pre-settlement otolith chemistry, indicating that they had occupied different water masses during their pelagic stage, thus suggesting multiple larval sources. Fish from reefs in the northern GBR did not differ greatly in their pre-settlement otolith chemistry, suggesting a single larval source. Using near-natal signatures, we determined that ~67% of these signatures matched the signature established for Lizard Island (LZ), suggesting LZ could be a source reef. However, these results could also be the result of poor separation among reefs caused by reefs sharing water masses. Otolith chemistry also revealed that 50–70% of all fish examined settled on the reef the day they encountered it, while some fish spent up to 4 days near the reef prior to settlement.     

Demographic and life history response of the cladoceranBosmina longirostris to variation in predator abundance

Population dynamics, demography and body size of the cladoceranBosmina longirostris were examined in an experimental study in which the abundance of its predator (the cyprind fishPhoxinus eos) was varied in an unproductive lake. Four densities of fish were used, encompassing the biomass of fish in the lake.Bosmina was most abundant at low and medium fish densities (1.06 and 2.12 g fish biomass · m^-3) and less abundant when fish were either absent or present at high density (3.71 g fish biomass · m^-3). The unimodal response to predator abundance resulted from effects on both birth and death rates.Bosmina birth rates increased as fish biomass increased, in response to increasing food (phytoplankton) biomass. Death rates were highest at high fish biomass (because of fish predation) and in the absence of fish (because of predation by the dipteranChaoborus, which was most abundant in the absence of fish). Size-frequency distributions revealed that fish eliminated the larger size classes ofBosmina, and mean carapace length ofBosmina populations was inversely proportional to fish biomass.Bosmina initiated reproduction at smaller size in the presence of fish than in their absence, and size at maturity was inversely proportional to fish biomass. Size at birth also tended to decrease with increasing fish biomass, but this trend was not as strong as that of size at maturity. Decreased size at maturity apparently allowedBosmina individuals to reproduce before becoming vulnerable to fish predation. Flexibility in size at maturity, together with low abundance of invertebrate predators and large herbivores (which were preyed upon by fish), allowedBosmina to become abundant in low and medium fish treatments. In the high fish treatment, mortality due to fish predation was too severe to be offset by decreased size at maturity, andBosmina population density was low. The net response ofBosmina populations to fish predation results from interactive effects of predation on mortality, natality, and life history traits.