Sara M. Lewis

The role of herbivorous fishes in the organization of a caribbean reef community

Experimental manipulations of grazing intensity were used to examine the role of herbivorous fishes in the families Acanthuridae (surgeonfishes) and Scaridae (parrotfishes) in determining distributions and abundances of benthic species within and among shallow tropical reef habitats. A back reef habitat along the Belizean barrier reef was characterized by a diverse benthic assemblage of algal turfs, coralline algae, and the coral Porites astreoides, but by extremely low macroalgal abundance. In contrast, several nearby shallow habitats were dominated by dense stands of several macroalgal species. Experimental reduction of herbivorous fish grazing in the back reef (achieved by constructing exclosures) rapidly and dramatically altered existing patterns of benthic species composition and species abundances. After 10 wk of reduced herbivory, total macroalgal abundance increased significantly in herbivore exclusion areas relative to unmanipulated controls, and was correlated with decreased percent cover of available space, several algal turf species, crustose coralline algae, and Porites. Some macroalgal species were able to directly overgrow and kill portions of Porites colonies within herbivore exclusion treatments. Successful recruitment and growth of several algal species under experimentally reduced herbivory indicated that macroalgal species distributions may be limited by herbivory rather than by lack of spore availability or unsuitable physical conditions. Algal turfs characteristic of many reef habitats appear to represent herbivore—tolerant assemblages, persisting under high grazing intensity but responding rapidly to reduced herbivory with increased abundances, morphological changes, and altered reproductive status. These results suggest that herbivorous fish grazing profoundly influences benthic species distributions and abundances within some tropical reef habitats. Spatial variation in herbivory appears to be of fundamental importance in determining regional patterns of benthic community structure on tropical reefs. The spatial mosaic of benthic community composition among shallow reef habitats was associated with patterns of grazing intensity by herbivorous fishes. Several reef habitats supporting dense macroalgal stands represented spatial refuges from herbivory, with low herbivorous fish densities and reduced grazing intensities. Transplant experiments revealed that algal species characteristic of these low—herbivory habitats were highly susceptible to grazing by herbivorous fishes. Spatial heterogeneity in grazing intensity may contribute to high regional diversity among tropical reef habitats by maintaining different benthic species assemblages under fundamentally distinct selective regimes.

HERBIVORE ABUNDANCE AND GRAZING INTENSITY ON A CARIBBEAN CORAL REEF

Herbivory is a primary factor in determining the structure of coral reef communities. Spatial variation among reef habitats in the intensity of herbivory has been documented, but underlying variation in species composition and abundance within the herbivore guild has received little attention. The distri- bution and relative abundances of herbivorous fishes and sea urchins across several habitats were studied on the Belizean barrier reef off the Caribbean coast of Central America. Marked variation in total herbivore density as well as major changes in the composition of the herbivore guild were found across reef habitats. Acanthurids (surgeonfishes) predominated in shallow areas (< 5 m) while scarids (parrotfishes) were dominant in deeper habitats. Significant differences among habitats in an experimental assay of grazing intensity were strongly correlated with herbivore abundance. The spatial distribution of herbivorous fishes across reef habitats does not appear to be simply explained by differences in reef topography, but may depend on complex interactions among proximity to nearby shelter, predator abundance, density of territorial competitors, and local availability of food resources.

Sources of Intraspecific Variation in Sperm Precedence in Red Flour Beetles

Sperm precedence, or differential fertilization success among mating males, is likely to be important in mediating male reproductive success in many species. Numerous studies have provided measures of sperm precedence for diverse taxonomic groups, but previous discussions have focused on species-specific mean P2 values. In this study, which systematically examined sources of variation in sperm precedence in Tribolium castaneum, consistent differences among male pairs represent a significant percentage (17.8%) of the total variation in sperm precedence. These results in T. castaneum and a reexamination of published studies reveal a high degree of intraspecific variation in sperm precedence, suggesting that mean values are insufficient to adequately characterize sperm-precedence patterns. Such individual variation in sperm precedence may profoundly influence differential reproductive success among males.

Can tropical seaweeds reduce herbivory by growing at night? Diel patterns of growth, nitrogen content, herbivory, and chemical versus morphological defenses

SummaryTropical seaweeds in the genus Halimeda reduce losses to grazing by capitalizing on diel patterns of herbivore activity. These seaweeds produce new, more herbivoresusceptible growth at night when herbivorous reef fishes are inactive. Plant portions more than 48 h old are low in food value, well defended morphologically (calcified and high in ash content), and relatively resistant to herbivory. Younger plant portions represent 3–4.5 times the food value (nitrogen or organic content) of older portions but are only moderately more susceptible to herbivores due to their high concentrations of the terpenoid feeding deterrents halimedatrial and halimedatetraacetate. Halimedatrial significantly deters grazing by both parrotfishes (Scaridae) and surgeonfishes (Acanthuridae) and occurs in high concentrations (2–4.5% of plant ash-free dry mass) in plant portions that are 4–12 h old, intermediate concentrations (0.3–2.3%) in portions that are 16–26 h old, and low concentrations (0.3%) in older plant portions. The related compound halimedatetraacetate is absent from the youngest plant portions, shows a rapid increase in concentration (from 0 to 1%) in plant material that is approximately 16 h old, and then rapidly declines to low levels (0.1 to 0.2%) in older plant portions. Thus, newly produced tissues are nutritionally valuable but contain high concentrations of defensive chemicals. As these tissues age, morphological defenses increase, the tissue becomes less valuable as a food for herbivores, and chemical defenses decrease. Additionally, new growth of Halimeda remains unpigmented until just before sunrise. Thus, the valuable, nitrogen-containing molecules associated with photosynthesis are not placed in the new, and more herbivore susceptible, growth until lights is available and they can start producing income for the plant.Experiments in a coral-reef microcosm, where diel patterns of light and water chemistry could be altered, indicated that Halimedas growth pattern is cued by the timing of light-dark cycles rather than by co-occurring diel changes in water chemistry. Although the growth patterns of Halimeda seem unusual, similar patterns appear to occur in numerous other seaweeds and in microalgae such as diatoms and dinoflagellates.

Herbivory on coral reefs: algal susceptibility to herbivorous fishes

SummaryThe susceptibility of several tropical algal species to fish grazing was studied on the Belizean barrier reef off the Caribbean coast of Central America. Short-term transplant experiments indicate that plant species vary markedly in their rates of biomass loss to grazing by a shallow-water guild of herbivorous fishes. Algal species transplanted from habitats with low grazing pressure are highly susceptible to grazing, while species occurring in habitats with high herbivore densities are highly resistant to grazing. Algal species show differential susceptibility to grazing by two major components of the tropical herbivore guild, Acanthurus (surgeonfishes) and Sparisoma (parrotfishes).Variability in plant susceptibility to grazing by herbivorous fishes was not clearly correlated with morphological or chemical characteristics that have been previously suggested as plant defenses against herbivory. Plants found to be highly resistant to fish grazing, such as Halimeda, exhibit both morphological characteristics and secondary chemical compounds which do appear to reduce herbivory. In contrast, species of Caulerpa, Sargassum, Turbinaria, and Padina, which also possess alleged morphological and/or chemical defenses, are nevertheless highly susceptible to fish grazing.

The Regulation of Morphological Plasticity in Tropical Reef Algae by Herbivory

Many tropical algal species exhibit considerable morphological variation associated with different coral reef habitats, but the factors contributing to such variation have not been identified. Two strikingly distinct morphologies are described here for the common Caribbean alga Padina jamaicensis. These distinct morphologies are character- istically found in different reef habitats, and are shown to represent phenotypic responses to different levels of herbivorous fish grazing. Experimental reduction of grazing intensity in a high-herbivory reef habitat resulted in a rapid (96 h) morphological shift from a prostrate, highly branched turf morphology to an erect foliose morphology. Transplant experiments indicated that foliose Padina plants were preferentially consumed by herbiv? orous parrotfishes. These results suggest that morphological plasticity may represent an important adaptive strategy in some tropical algal species, enabling plants to persist in intensely grazed reef habitats while maintaining the ability to respond rapidly to spatial or temporal fluctuations in herbivory.

An integrative view of sexual selection in Tribolium flour beetles

Sexual selection is a major force driving the evolution of diverse reproductive traits. This evolutionary process is based on individual reproductive advantages that arise either through intrasexual competition or through intersexual choice and conflict. While classical studies of sexual selection focused mainly on differences in male mating success, more recent work has focused on the differences in paternity share that may arise through sperm competition or cryptic female choice whenever females mate with multiple males. Thus, an integrative view of sexual selection needs to encompass processes that occur not only before copulation (pre‐mating), but also during copulation (peri‐mating), as well as after copulation (post‐mating), all of which can generate differences in reproductive success. By encompassing mechanisms of sexual selection across all of these sequential reproductive stages this review takes an integrative approach to sexual selection in Tribolium flour beetles (Coleoptera: Tenebrionidae), a particularly well‐studied and economically important model organism.

The influence of male ejaculate quantity on female fitness: a meta-analysis.

Although the primary function of mating is gamete transfer, male ejaculates contain numerous other substances that are produced by accessory glands and transferred to females during mating. Studies with several model organisms have shown that these substances can exert diverse behavioural and physiological effects on females, including altered longevity and reproductive output, yet a comprehensive synthesis across taxa is lacking. Here we use a meta‐analytic approach to synthesize quantitatively extensive experimental work examining how male ejaculate quantity affects different components of female fitness. We summarize effect sizes for female fecundity (partial and lifetime) and longevity from 84 studies conducted on 70 arthropod species that yielded a total of 130 comparisons of female fecundity and 61 comparisons of female longevity. In response to greater amounts of ejaculate, arthropod females demonstrate enhanced fecundity (both partial and lifetime) but reduced longevity, particularly for Diptera and Lepidoptera. Across taxa, multiply mated females show particularly large fecundity increases compared to singly mated females, indicating that single matings do not maximize female fitness. This fecundity increase is balanced by a slight negative effect on lifespan, with females that received more ejaculate through polyandrous matings showing greater reductions in lifespan compared with females that have mated repeatedly with the same male. We found no significant effect size differences for either female fecundity or longevity between taxa that transfer sperm packaged into spermatophores compared to taxa that transfer ejaculates containing free sperm. Furthermore, females that received relatively larger or more spermatophores demonstrated greater lifetime fecundity, indicating that these seminal nuptial gifts provide females with a net fitness benefit. These results contribute to our understanding of the evolutionary origin and maintenance of non‐sperm ejaculate components, and provide insight into female mate choice and optimal mating patterns.

Courtship Behavior in the Dwarf Seahorse, Hippocampus zosterae

The seahorse genus Hippocampus (Syngnathidae) exhibits extreme morpho- logical specialization for paternal care, with males incubating eggs within a highly vascularized brood pouch. Dwarf seahorses, H. zosterae, form monoga- mous pairs that court early each morning until copulation takes place. Daily behavioral observations of seahorse pairs (n = 15) were made from the day of introduction through the day of copulation. Four distinct phases of seahorse courtship are marked by prominent behavioral changes, as well as by differences in the intensity of courtship. The first courtship phase occurs for one or two mornings preceding the day of copulation and is characterized by reciprocal quivering, consisting of rapid side-to-side body vibrations displayed alternately by males and females. The remaining courtship phases are restricted to the day of copulation, with the second courtship phase distinguished by females pointing, during which the head is raised upward. In the third courtship phase, males begin to point in response to female pointing. During the final phase of courtship, seahorse pairs repeatedly rise together in the water column, eventually leading to females transferring their eggs directly into the male brood pouch during a brief midwater copulation. Courtship activity level (representing the percentage of time spent in courtship) increased from relatively low levels during the first courtship phase to highly active courtship on the day of copulation. Males more actively initiated courtship on the days preceding copulation, indicating that these seahorses are not courtship-role reversed, as has previously been assumed.

Female role in sperm storage in the red flour beetle, Tribolium castaneum

This study clarifies the role of female-controlled processes contributing to sperm storage in the red flour beetle, Tribolium castaneum. Evidence presented indicates that sperm motility is not affected by extreme hypoxia produced by anesthetization of the female with either carbon dioxide or nitrogen. Sperm location and motility in low-oxygen environments did not differ from that of sperm in reproductive tracts immersed in fully aerated saline. Sperm motility was unaffected by exposure to potassium cyanide, an aerobic respiratory system poison, but was inhibited by exposure to iodoacetic acid, a glycolysis poison. Based on the retention of sperm motility under extreme hypoxia, female control over sperm storage was then examined. Both anesthetized females and dead females had fewer stored sperm after mating than unanesthetized control females. These results suggest that female T. castaneum play an active role in moving sperm from the site of deposition into storage.