Amphibian secondary metabolites are well known chemically, but their ecological functions are poorly understood--even for well-studied species. For example, the eastern newt (Notophthalmus viridescens) is a well known secretor of tetrodotoxin (TTX), with this compound hypothesized to facilitate this salamander's coexistence with a variety of aquatic consumers across the eastern United States. However, this assumption of chemical defense is primarily based on observational data with low replication against only a few predator types. Therefore, we tested the hypothesis that N. viridescens is chemically defended against co-occurring fishes, invertebrates, and amphibian generalist predators and that this defense confers high survivorship when newts are transplanted into both fish-containing and fishless habitats. We found that adult eastern newts were unpalatable to predatory fishes (Micropterus salmoides, Lepomis macrochirus) and a crayfish (Procambarus clarkii), but were readily consumed by bullfrogs (Lithobates catesbeianus). The eggs and neonate larvae were also unpalatable to fish (L. macrochirus). Bioassay-guided fractionation confirmed that deterrence is chemical and that ecologically relevant concentrations of TTX would deter feeding. Despite predatory fishes rejecting eastern newts in laboratory assays, field experiments demonstrated that tethered newts suffered high rates of predation in fish-containing ponds. We suggest that this may be due to predation by amphibians (frogs) and reptiles (turtles) that co-occur with fishes rather than from fishes directly. Fishes suppress invertebrate consumers that prey on bullfrog larvae, leading to higher bullfrog densities in fish containing ponds and thus considerable consumption of newts due to bullfrog tolerance of newt chemical defenses. Amphibian chemical defenses, and consumer responses to them, may be more complex and indirect than previously appreciated.
There has been much discussion and some controversy regarding the role of nutrient enrichment versus other factors, such as altered rates of herbivory, in the degradation of coral reef ecosystems. The resolution of this controversy has been hampered by the lack of manipulative field studies testing the effects of ecologically relevant levels of nutrient enrichment on coral reef communities. We present a new method for adding ecologically relevant levels of nutrients to experimental substrates on coral reefs. The method elevates nutrients for sustained periods of time (>41 d without replenishment of nutrients) and allows for testing interactions of nutrients and altered levels of herbivory. Results from an offshore reef in Key Largo, Florida, show strong effects of excluding large herbivorous fishes, negligible effects of nutrient enrichment–;or effects that are opposite of predictions, and no interaction between nutrient levels and herbivory in affecting algal abundance. Patterns observed for this reef did not confirm predictions of previously proposed models that frondose macroalgal or crustose algal abundance would be enhanced with nutrient enrichment or that dominance of filamentous turfs would be greater in unenriched conditions. In contrast to previous predictions, the abundance of larger macroalgae at this site was not increased by elevating nutrients above predicted threshold response levels of 1.0 µM for total inorganic nitrogen or 0.10 µM for soluble reactive phosphate. Also conflicting with some models, filamentous, nitrogen‐fixing cyanobacteria were enhanced, rather than suppressed, by nutrient enrichment.
Physical stress to seaweeds and hunger stress of herbivores can influence the outcome of chemically mediated seaweed-herbivore interactions. The unpalatable brown seaweed Dictyota ciliolata produces the diterpenoid secondary metabolites pachydictyol A, dictyol B acetate, and dictyodial. At natural concentrations, pachydictyol A deterred the sea urchin Arbacia punctulata but did not inhibit feeding by the pinfish Lagodon rhomboides or the amphipod Ampithoe longimana until concentrations were 2.5-5 times natural levels. Dictyol B acetate deterred the urchin, amphipod, and pinfish at, or far below, natural concentrations. Dictyodial was too unstable to assay directly, but indirect experiments suggested that natural concentrations deterred the urchin, but not the pinfish or amphipod. Mild desiccation of D. ciliolata reduced concentrations of the different secondary metabolites by 7-38% and plants became 2.6-3.4 times more susceptible to urchin and amphipod grazing. The combined concentrations of pachydictyol A and dictyol B acetate found in undesiccated Dictyota ciliolata deterred feeding by urchins, but this deterrent effect was lost at concentrations found in the desiccated plants. Desiccated and undesiccated plants did not differ in nutritive value (as measured by protein and total N content) or toughness. Thus, desiccated plants became more palatable because chemical defenses were lost, not because nutritive value was increased. The stress of near-surface ultraviolet radiation also caused significant physiological changes in Dictyota ciliolata. UV-exposed blades bleached, senesced, and grew 84% less than blades protected from UV radiation. Tissue loss and minimal growth of UV-stressed plants constrained our sample sizes, but the limited assays that could be run suggested that UV stress may lower chemical defenses and increase plant susceptibility to herbivores. Because many previous investigations of herbivore feeding patterns used animals that had been starved for days before an assay, we tested the effects of this commonly used procedure on feeding discrimination. Recently fed urchins always avoided food containing natural concentrations of pachydictyol A during separate feeding trials performed on each of four consecutive days. In contrast, urchins deprived of food for 3 d before this assay did not avoid the treated food on days 1 and 2 of feeding trials, but did avoid it on days 3 and 4 after their hunger was reduced by feeding during days 1 and 2. If we had used only starved urchins (a common procedure in previous investigations), we could have concluded, with apparent justification, that urchins were unaffected by pachydictyol A (if the assays were run for only 1-2 d) or that they needed 2 d of exposure to the compound in order to learn to avoid it. Both of these conclusions would have been incorrect.
Reef habitats in the South Atlantic Bight comprise only a small proportion of the available benthic habitat, but support enormous densities of fishes, invertebrates, and algae. Because these reefs are distant from shore and in deeper waters (usually >20 m), we know very little about the factors affecting the distribution of fishes or the structure of fish assemblages on these reefs. Our goal in this study was to determine how variance in macroalgal structure and species composition on reefs off the coast of North Carolina affects reef fishes. Manipulations of algal density and species composition in artificially created 1.5 m2 plots indicated that: (1) patches with Sargassum filipendula had greater numbers and higher species richness of reef fishes than patches without algae; (2) fishes responded to variability in Sargassum density with high fish abundance in patches with higher algal density, but this response varied among fish species; (3) greater numbers of individuals and species of fish used Sargassum patches compared to patches of 2 other brown algae common in this region (Zonaria and Dictyopteris); (4) fish densities were higher in patches with taller algae and greater percent cover of algae. Many of the fishes using experimental patches were small, young-of-the-year individuals. Our results indicate that variability in the structure of seaweed beds on reefs in the South Atlantic Bight can influence the distribution and abundance of reef fishes. Although the algal beds in the South Atlantic Bight are more diminutive and less extensive than kelp beds of cold temperate waters, they may be a critical habitat for juvenile reef fishes in this region, and the algal species dominating the beds will have a large effect on their value as juvenile habitat.
Numerous studies have assessed the individual effects of prey nutritional quality or chemical defenses on consumer feeding behavior. However, little is known about how these traits interact to affect consumer feeding and performance. We tested the separate and interactive effects of prey chemical defenses and nutritional quality on the feeding behavior and fitness of six sympatric crustacean mesograzers. Natural concentrations of diterpene alcohols (dictyols) from the brown alga Dictyota menstrualis were incorporated, or not incorporated, into lower quality and higher quality foods to create artificial diets mimicking prey of variable value and defense. Five amphipods (Ampithoe longimana, A. valida, Cymadusa compta, Gammarus mucronatus, and Elasmopus levis) and one isopod (Paracerceis caudata), representing a continuum of closely to distantly related organisms, were fed intact algae or lower and higher quality diets containing or lacking dictyols. All six mesograzers preferred the green alga Enteromorpha intestinalis to the dictyol producing alga Dictyota menstrualis. In assays allowing consumers to choose between simultaneously available foods, dictyols deterred feeding by all five amphipods, but not the isopod; this occurred for both lower and higher quality foods. In no-choice assays, where consumers were confined with only one of our four treatment diets, effects on feeding became more complex. Nutritional quality alone affected feeding by five of the six species. These grazers compensated for lower quality by increasing consumption. Dictyols suppressed feeding for four of the six species. More interestingly, there were significant dictyol × quality interactions for three species. Dictyols decreased feeding more when placed in lower quality foods than higher quality foods. Two amphipods deterred by dictyols in the choice assays readily consumed dictyol-containing foods in no-choice situations and suffered few negative effects of doing so. Although all amphipods were deterred by dictyols in choice assays, dictyols decreased fitness (survivorship, growth, or reproduction) for only four of the five species. These effects included large and immediate decreases in survivorship, dramatic effects on reproduction, and modest effects on female growth. Dictyols enhanced survivorship of the isopod. Thus, the effects of secondary metabolites on feeding in choice situations vs. fitness in long-term assays were inconsistent. For three amphipods, certain effects of food quality, dictyols, or their interaction were detected only for females. In general, negative effects of dictyols on fitness were greater in lower than in higher quality foods, suggesting that prey nutritional value may counteract the effects of defensive metabolites. For example, when G. mucronatus consumed dictyols in lower quality foods, mortality was >80% by day 5; for dictyols in higher quality foods, 80% mortality took 28 days to develop. Lower quality foods alone significantly decreased growth for the isopod, three of the amphipods, and the females of a fourth amphipod, concomitantly reducing fecundity for four of the five amphipods studied. The effects of both chemical defenses and nutritional quality were unrelated to consumer phylogeny; responses of congeners varied as much, or more, than responses of more distantly related consumers. Understanding mechanisms and consequences of food selection requires that the interactive effects of both chemical defenses and prey nutritional characteristics be considered explicitly. Corresponding Editor: J. D. Witman.
MEPS Marine Ecology Progress Series Contact the journal Facebook Twitter RSS Mailing List Subscribe to our mailing list via Mailchimp HomeLatest VolumeAbout the JournalEditorsTheme Sections MEPS 225:17-28 (2002) - doi:10.3354/meps225017 Plankton tethering to assess spatial patterns of predation risk over a coral reef and seagrass bed Stephan G. Bullard1, Mark E. Hay2,* 1Institute of Marine Sciences, University of North Carolina at Chapel Hill, 3431 Arendell St., Morehead City, North Carolina 28557, USA 2School of Biology, Georgia Institute of Technology, Atlanta, Georgia 30332-0230, USA *Corresponding author. E-mail: mark.hay@biology.gatech.edu ABSTRACT: Difficulties associated with manipulating plankton in situ have limited the ability of investigators to assess among-habitat variation in predation risk for plankton. We used plankton tethering units (PTUs) to tether zooplankton in a variety of reef and seagrass habitats, and used field and laboratory assays to test PTUs for tethering artifacts. Tethering did not affect the survivorship of 5 species of plankton (sizes <1 to 6 mm), indicating that the method works with a range of planktonic organisms. We then used the reef mysid Mysidium columbiae in additional assays and found that: (1) mysids remained on PTUs unless they were attacked by predators; (2) PTUs did not prevent planktivorous fishes from consuming tethered mysids; (3) untethered mysids commonly evaded predators, while mysids on PTUs did not; (4) the same types of predators consumed untethered and tethered mysids in the field; and (5) fishes were neither attracted to nor repelled from PTUs. We used PTUs and mysids to assess predation risk for plankton over various coral reef or seagrass habitats. Risk of attack varied among habitats and was correlated with abundance of planktivorous fishes. On the reef, attack rates were most intense over a topographically complex reef ledge, less intense over the less structurally complex center of the reef, and least intense over a structurally simple sand plain. Within the seagrass bed, attack rates were highest at the edge of the bed and less intense to the center of the seagrass bed and over an adjacent sand plain. Thus, attack rates at these sites varied tremendously over small spatial scales (meters). KEY WORDS: Invertebrate larvae · Mysids · Mysidium columbiae · Planktivory · Tethering · Zooplankton Full text in pdf format PreviousNextExport citation RSS - Facebook - Tweet - linkedIn Cited by Published in MEPS Vol. 225. Online publication date: January 11, 2002 Print ISSN: 0171-8630; Online ISSN: 1616-1599 Copyright © 2002 Inter-Research.
Coral reefs are in dramatic global decline, with seaweeds commonly replacing corals. It is unclear, however, whether seaweeds harm corals directly or colonize opportunistically following their decline and then suppress coral recruitment. In the Caribbean and tropical Pacific, we show that, when protected from herbivores, ~40 to 70% of common seaweeds cause bleaching and death of coral tissue when in direct contact. For seaweeds that harmed coral tissues, their lipid-soluble extracts also produced rapid bleaching. Coral bleaching and mortality was limited to areas of direct contact with seaweeds or their extracts. These patterns suggest that allelopathic seaweed-coral interactions can be important on reefs lacking herbivore control of seaweeds, and that these interactions involve lipid-soluble metabolites transferred via direct contact. Seaweeds were rapidly consumed when placed on a Pacific reef protected from fishing but were left intact or consumed at slower rates on an adjacent fished reef, indicating that herbivory will suppress seaweeds and lower frequency of allelopathic damage to corals if reefs retain intact food webs. With continued removal of herbivores from coral reefs, seaweeds are becoming more common. This occurrence will lead to increasing frequency of seaweed-coral contacts, increasing allelopathic suppression of remaining corals, and continuing decline of reef corals.
