Temporal, spatial, and body size effects on growth rates of loggerhead sea turtles (Caretta caretta) in the Northwest Atlantic

In response to a call from the US National Research Council for research programs to combine their data to improve sea turtle population assessments, we analyzed somatic growth data for Northwest Atlantic (NWA) loggerhead sea turtles (Caretta caretta) from 10 research programs. We assessed growth dynamics over wide ranges of geography (9–33°N latitude), time (1978–2012), and body size (35.4–103.3 cm carapace length). Generalized additive models revealed significant spatial and temporal variation in growth rates and a significant decline in growth rates with increasing body size. Growth was more rapid in waters south of the USA (<24°N) than in USA waters. Growth dynamics in southern waters in the NWA need more study because sample size was small. Within USA waters, the significant spatial effect in growth rates of immature loggerheads did not exhibit a consistent latitudinal trend. Growth rates declined significantly from 1997 through 2007 and then leveled off or increased. During this same interval, annual nest counts in Florida declined by 43 % (Witherington et al. in Ecol Appl 19:30–54, 2009) before rebounding. Whether these simultaneous declines reflect responses in productivity to a common environmental change should be explored to determine whether somatic growth rates can help interpret population trends based on annual counts of nests or nesting females. Because of the significant spatial and temporal variation in growth rates, population models of NWA loggerheads should avoid employing growth data from restricted spatial or temporal coverage to calculate demographic metrics such as age at sexual maturity.     

Re-creating missing population baselines for Pacific reef sharks

Sharks and other large predators are scarce on most coral reefs, but studies of their historical ecology provide qualitative evidence that predators were once numerous in these ecosystems. Quantifying density of sharks in the absence of humans (baseline) is, however, hindered by a paucity of pertinent time-series data. Recently researchers have used underwater visual surveys, primarily of limited spatial extent or nonstandard design, to infer negative associations between reef shark abundance and human populations. We analyzed data from 1607 towed-diver surveys (>1 ha transects surveyed by observers towed behind a boat) conducted at 46 reefs in the central-western Pacific Ocean, reefs that included some of the world's most pristine coral reefs. Estimates of shark density from towed-diver surveys were substantially lower (<10%) than published estimates from surveys along small transects (<0.02 ha), which is not consistent with inverted biomass pyramids (predator biomass greater than prey biomass) reported by other researchers for pristine reefs. We examined the relation between the density of reef sharks observed in towed-diver surveys and human population in models that accounted for the influence of oceanic primary productivity, sea surface temperature, reef area, and reef physical complexity. We used these models to estimate the density of sharks in the absence of humans. Densities of gray reef sharks (Carcharhinus amblyrhynchos), whitetip reef sharks (Triaenodon obesus), and the group "all reef sharks" increased substantially as human population decreased and as primary productivity and minimum sea surface temperature (or reef area, which was highly correlated with temperature) increased. Simulated baseline densities of reef sharks under the absence of humans were 1.1-2.4/ha for the main Hawaiian Islands, 1.2-2.4/ha for inhabited islands of American Samoa, and 0.9-2.1/ha for inhabited islands in the Mariana Archipelago, which suggests that density of reef sharks has declined to 3-10% of baseline levels in these areas.     

Scale dependence in species turnover reflects variance in species occupancy

Patterns of species turnover may reflect the processes driving community dynamics across scales. While the majority of studies on species turnover have examined pairwise comparison metrics (e.g., the average Jaccard dissimilarity), it has been proposed that the species-area relationship (SAR) also offers insight into patterns of species turnover because these two patterns may be analytically linked. However, these previous links only apply in a special case where turnover is scale invariant, and we demonstrate across three different plant communities that over 90% of the pairwise turnover values are larger than expected based on scale-invariant predictions from the SAR. Furthermore, the degree of scale dependence in turnover was negatively related to the degree of variance in the occupancy frequency distribution (OFD). These findings suggest that species turnover diverges from scale invariance, and as such pairwise turnover and the slope of the SAR are not redundant. Furthermore, models developed to explain the OFD should be linked with those developed to explain species turnover to achieve a more unified understanding of community structure.     

Density-dependent facilitation cascades determine epifaunal community structure in temperate Australian mangroves

Co-occurring foundation species can determine biological community structure via facilitation cascades. We examined the density dependencies of facilitation cascades, including how the density of a basal foundation species influences the density of secondary foundation species, and how the density of secondary foundation species influences community structure. The system in which we assessed density dependencies was a temperate mangrove forest in which pneumatophores trap the fucoid alga Hormosira banksii and provide substrate for the oyster, Saccostrea glomerata. The alga and oyster in turn determine benthic community structure. In the field, algal biomass was positively correlated with pneumatophore density. Oysters, by contrast, were highly over-dispersed and correlated with the presence/absence of pneumatophores. Epifaunal abundance and species richness were positively correlated with algal and oyster abundance, but their effects were independent. The positive effect of pneumatophore density on epifauna was primarily an indirect effect of trapping more algae. Pneumatophores did not directly influence invertebrate communities. Experiments revealed that, at very low pneumatophore densities, algal retention was insufficient to facilitate epifauna above that found on pneumatophores alone. At higher densities, however, increasing the density of pneumatophores increased algal retention, and the density and diversity of associated invertebrates. Shading by the mangrove canopy reduced algal biomass but did not modify the density-dependent nature of the cascade. Our results extend facilitation theory by showing that the density of both basal and secondary foundation species can be critical in triggering facilitation cascades. Our study also reveals that, where foundation species co-occur, multiple, independent cascades may arise from a single basal facilitator. These findings enhance our understanding of the role of density-dependent facilitation cascades in community assembly.     

Dissolved histamine: a potential habitat marker promoting settlement and metamorphosis in sea urchin larvae

Many species of marine invertebrate larvae settle and metamorphose in response to chemicals produced by organisms associated with the adult habitat, and histamine is a cue for larvae of the sea urchin Holopneustes purpurascens. This study investigated the effect of histamine on larval metamorphosis of six sea urchin species. Histamine induced metamorphosis in larvae of three lecithotrophic species (H. purpurascens, Holopneustes inflatus and Heliocidaris erythrogramma) and in one planktotrophic species (Centrostephanus rodgersii). Direct comparisons of metamorphic rates of lecithotrophic and planktotrophic larvae in assays cannot be made due to different proportions of larvae being competent. Histamine (10 μM) induced metamorphosis in 95% of larvae of H. purpurascens and H. inflatus after 1 h, while the coralline alga Amphiroa anceps induced metamorphosis in 40–50% of these larvae. Histamine (10 μM) and A. anceps induced 40 and 80% metamorphosis, respectively, in the larvae of H. erythrogramma after 24 h. Histamine (10 μM) and the coralline alga Corallina sp. induced 30 and 70% metamorphosis, respectively, in the larvae of C. rodgersii after 24 h. No metamorphosis of any larval species occurred in seawater controls. Larvae of two planktotrophic species (Tripneustes gratilla and Heliocidaris tuberculata) did not metamorphose in response to histamine. Seagrasses, the host plants of H. inflatus, induced rapid metamorphosis in larvae of the two Holopneustes species, and several algae induced metamorphosis in C. rodgersii larvae. Histamine leaching from algae and seagrasses may act as a habitat marker and metamorphic cue for larvae of several ecologically important sea urchin species.     

Urban marine ecology in southern California: the ability of riprap structures to serve as rocky intertidal habitat

Increasing human populations along marine coastlines has lead to increasing urbanization of the marine environment. Despite decades of investigations on terrestrial ecosystems, the effect of urbanization on marine life is not well understood. Riprap is the rocky rubble used to build jetties, breakwaters, and armored shorelines. Roughly 30% of the southern California shoreline supports some form of riprap, while 29% of the shoreline is natural rocky substrate. Astonishingly few studies have investigated this anthropogenic rocky habitat even though it rivals a natural habitat in area on a regional scale along a coastline that has been extensively studied. In this study, I compared the diversity and community structure of exposed rocky intertidal communities on four riprap and four natural sites in southern California. I ask the following questions: (1) does diversity or community composition differ between intertidal communities on riprap and natural rocky habitats in southern California, (2) if so, which organisms contribute to those differences, (3) which physical factors are contributing to these differences, and (4) do riprap habitats support higher abundances of invasive species than natural habitats? On average, riprap and natural rocky habitats in wave exposed environments in southern California did not differ from each other in diversity or community composition when considering the entire assemblage. However, when only mobile species were considered, they occurred in greater diversity on natural shores. These differences appear to be driven by wave exposure. The presence of invasive species was negligible in both natural and riprap habitats.     

Worker reproductive parasitism and drift in the western honeybee Apis mellifera

When a honeybee (Apis spp.) colony loses its queen and is unable to rear a new one, some of the workers activate their ovaries and produce eggs. When a colony has a queen (i.e., it is queenright) almost all worker-laid eggs are eaten, but when hopelessly queenless, the workers become more tolerant of worker-laid eggs and rear some of them to adult drones. This increased tolerance renders a queenless colony vulnerable to worker reproductive parasitism, wherein unrelated workers enter the colony and lay eggs. Here, we show that the proportion of unrelated (non-natal) workers significantly decreases after an Apis mellifera colony becomes queenless. The remaining non-natal workers are as likely to have activated ovaries as natal workers, yet they produce more eggs than natal workers, resulting in significantly higher reproductive success for non-natal workers. In a second experiment, we provided queenless and queenright workers with a choice to remain in their own colony or to join a queenless or queenright colony nearby. The experiment was set up such that worker movement was unlikely to be due to simple orientation errors. Very few workers joined another colony, and there was no preference for workers to drift into or out of queenless or queenright colonies, in accordance with the proportion of non-natal workers declining significantly after becoming queenless in the first experiment.     

Genetic connectivity and historical demography of the blue barred parrotfish (Scarus ghobban) in the western Indian Ocean

Studies on genetic connectivity are essential for the design of management strategies for coral reef fisheries. In this study we used a mitochondrial DNA marker to investigate population structure of the reef-associated parrotfish, Scarus ghobban, from four countries, Kenya, Mauritius, Seychelles and Tanzania, in the western Indian Ocean. We obtained nucleotide sequences of the mitochondrial control region for 117 individuals. Measures of haplotype diversity were relatively high. Pairwise population differentiation (F _ST) was low, but not always non-significant. Analysis of molecular variance (AMOVA) showed genetic differentiation between groups, when the data was partitioned into two groups consisting of samples from Mauritius and Tanzania in one group, and samples from Kenya and Seychelles in another group. Direction of gene flow was estimated using a Bayesian approach. Migration was sometimes asymmetric or directional, coinciding with the flow of major oceanic and coastal currents in the region. Mismatch distributions, based on the observed number of differences among haplotype pairs, produced a unimodal distribution, indicative of recent demographic expansion. Phylogenetic analyses revealed three clades without any geographic structure, suggesting recent migration between historically isolated lineages. We reconstructed the historical demography of S. ghobban and examined it in the context of Pleistocene climate stages and changes in relative sea level. Overall, these results showed that populations of S. ghobban are genetically diverse and have relatively high gene flow, with some genetic structuring in the western Indian Ocean.     

The ecological significance of lipid/fatty acid synthesis in developing eggs and newly hatched larvae of Pacific cod (Gadus macrocephalus)

The lipid/fatty acid composition of marine fish eggs and larvae is linked with buoyancy regulation, but our understanding of such processes is largely restricted to species with pelagic eggs. In this study, we examined developmental changes in the lipid/fatty acids of eggs and embryos of Pacific cod (Gadus macrocephalus), a species that spawns demersal eggs along coastal shelf edges, but as larvae must make a rapid transition to the upper reaches of the water column. Adult Pacific cod were collected in the Gulf of Alaska during the spawning season and eggs of two females were artificially fertilized with sperm from three males for each female. The eggs were subsequently reared in the laboratory to determine (1) how lipids/fatty acids were catabolized during egg and larval development, and (2) whether lipid/fatty acid catabolism had measurable effects on egg/embryo density. Eggs incubated at 4°C began hatching after 3-weeks and continued to hatch over a 10-day period, during which there was a distinct shift in lipid classes (phospholipids (PL), triacyglycerols (TAG), and sterols (ST)) and essential fatty acids (EFAs: 22:6n-3 (DHA), 20:5n-3 (EPA), and 20:4n-6 (AA)). In the egg stage, total lipid content steadily decreased during the first 60% of development, but just prior to hatch we observed an unexpected 2–3-fold lipid increase (~6–9 μg individual⁻¹) and a significant drop in egg density. The increase in lipids was largely driven by PL, with evidence of long-chained fatty acid synthesis. Late-hatching larvae had progressively decreasing lipid and fatty acid reserves, suggesting a shift from lipogenesis to lipid catabolism with continued larval development. Egg density measures suggest that lipid/fatty acid composition is linked to buoyancy regulation as larvae shift from a demersal to a pelagic existence following hatch. The biochemical pathway by which Pacific cod are apparently able to synthesize EFAs is unknown, therefore representing a remarkable finding meriting further investigation.     

Causes of habitat divergence in two species of agamid lizards in arid central Australia

The deserts of central Australia contain richer communities of lizards than any other arid regions, with the highest diversity occurring in sand dune habitats dominated by hummock-forming spinifex grasses. To investigate the mechanisms that permit coexistence, we studied two species of coexisting agamid lizards that exhibit striking divergence in their use of habitat in the Simpson Desert of central Australia. Here, the military dragon Ctenophorus isolepis is restricted primarily to sites providing > 30% cover of hard spinifex Triodia basedowii, whereas the central netted dragon C. nuchalis occurs in areas with much sparser (< 10%) cover. We constructed four mechanistic models to explain this pattern and then derived hypotheses to test them. One hypothesis, that competition restricts each species to its preferred habitat, was rejected after dyad encounters in field enclosures failed to elicit any habitat shift or any overt interactions between the species. Our next hypotheses were that each species exhibits preferences for different thermal environments or different prey types and that each selects the habitats that maximize access to them. Both were supported. C. isolepis preferred lower temperatures when active and specialized in eating ants < 5 mm long and selected spinifex-dominated areas where these requirements were met. In contrast, C. nuchalis preferred higher temperatures and a diversity of prey, both of which were available mostly in open areas. Finally, we used plasticine models to test the hypothesis that each species faced lower risk of predation in its selected habitat. This was partly supported, as models of both species were attacked more often in the open than under spinifex cover. The results show that habitat divergence occurs along several, probably covarying, niche axes. We suggest that different levels of spinifex cover provide the template for a broad range of ecological interactions, allowing lizard species to partition biotic and abiotic resources and achieve the extraordinarily high levels of local diversity that are observed.