Daphne G. Fautin

Managing for ocean biodiversity to sustain marine ecosystem services

Managing a complex ecosystem to balance delivery of all of its services is at the heart of ecosystem-based management. But how can this balance be accomplished amidst the conflicting demands of stakeholders, managers, and policy makers? In marine ecosystems, several common ecological mechanisms link biodiversity to ecosystem functioning and to a complex of essential services. As a result, the effects of preserving diversity can be broadly beneficial to a wide spectrum of important ecosystem processes and services, including fisheries, water quality, recreation, and shoreline protection. A management system that conserves diversity will help to accrue more “ecoservice capital” for human use and will maintain a hedge against unanticipated ecosystem changes from natural or anthropogenic causes. Although maintenance of biodiversity cannot be the only goal for ecosystem-based management, it could provide a common currency for evaluating the impacts of different human activities on ecosystem functioning and can...

Structural diversity, systematics, and evolution of cnidae

Cnidae are secreted by the Golgi apparatus of all cnidarians and only cnidarians. Of the three categories of cnidae (also called cnidocysts), nematocysts occur in all cnidarians, and are the means by which cnidarians defend themselves and obtain prey; spirocysts and ptychocysts are restricted to a minority of major taxa. A cnida discharges by eversion of its tubule; venom may be associated with the tubule of a nematocyst. About 30 major morphological types of nematocysts are recognized, but no single nomenclature for them is accepted. Function seems not to correlate tightly with morphology--nematocysts of at least some types are used both offensively and defensively. Similarly, it is not clear if morphology correlates with toxicity. Some types of nematocysts are taxonomically diagnostic whereas others are widespread. Nonetheless, an inventory of types of cnidae (the cnidom), with their distribution and size, is an essential component of most taxonomic descriptions. Complicating the taxonomic value of cnidae are the facts that not all members of a species may have the same types of cnidae, even at the same life-cycle stage, and size of nematocysts of a species may vary geographically and with size of individual. The diversity of nematocysts is so great and the features within each major type are so variable that homologies have not been determined. Nematocyst complement, morphology, and size likely reflect both phylogeny and biology; the feedback between the two may confound analysis. Although cnidae are valuable in taxonomy of at least some groups, more understanding of the forces that affect them is needed for their systematic and phylogenetic value to be understood and their potential as indicators of evolution to be realized.

Finding NEMO: nestedness engendered by mutualistic organization in anemonefish and their hosts

The interaction structure of mutualistic relationships, in terms of relative specialization of the partners, is important to understanding their ecology and evolution. Analyses of the mutualistic interaction between anemonefish and their host sea anemones show that the relationship is highly nested in structure, generalist species interacting with one another and specialist species interacting mainly with generalists. This supports the hypothesis that the configuration of mutualistic interactions will tend towards nestedness. In this case, the structure of the interaction is at a much larger scale than previously hypothesized, across more than 180° of longitude and some 60° of latitude, probably owing to the pelagic dispersal capabilities of these species in a marine environment. Additionally, we found weak support for the hypothesis that geographically widespread species should be more generalized in their interactions than species with small ranges. This study extends understanding of the structure of mutualistic relationships into previously unexplored taxonomic and physical realms, and suggests how nestedness analysis can be applied to the conservation of obligate species interactions.

The Adaptive Hypothesis of Bleaching

Despite the perception that corals and coral reefs are limited to stable habitats distinguished by very narrow environmental parameters, the coral-algal symbiosis is capable of surviving under a variety of extreme conditions. Through the process of photoadaptation, corals and their algal symbionts adjust algal densities and pigment concentrations to function over a wide range of light levels ranging from direct exposure to full sunlight in intertidal corals to virtual darkness at the extreme limits of the photic zone (>200 m) on reef slopes (Zahl and McLaughlin 1959; Schlichter et al. 1986). Corals and reef communities in some areas (such as the Arabian Gulf) tolerate salinity and temperature conditions that are lethal when imposed rapidly on the same species in less extreme environments (Coles 1988; Sheppard 1988; Coles and Fadlallah 1991; Chap. 23, Jokiel, this Vol.). There are abundant reports of reef corals occurring in turbid, high nutrient, nearshore habitats (Larcombe et al. 2001). Coral reefs exist at the inherently variable interface between the sea, air and land (Smith and Buddemeier 1992), and reef communities have persisted over geological time through significant climate and sea-level fluctuations. Despite this, rates of speciation and extinction in scleractinian corals have been relatively low over the last 220 million years (Veron 1995).

An Overview of Marine Biodiversity in United States Waters

Marine biodiversity of the United States (U.S.) is extensively documented, but data assembled by the United States National Committee for the Census of Marine Life demonstrate that even the most complete taxonomic inventories are based on records scattered in space and time. The best-known taxa are those of commercial importance. Body size is directly correlated with knowledge of a species, and knowledge also diminishes with distance from shore and depth. Measures of biodiversity other than species diversity, such as ecosystem and genetic diversity, are poorly documented. Threats to marine biodiversity in the U.S. are the same as those for most of the world: overexploitation of living resources; reduced water quality; coastal development; shipping; invasive species; rising temperature and concentrations of carbon dioxide in the surface ocean, and other changes that may be consequences of global change, including shifting currents; increased number and size of hypoxic or anoxic areas; and increased number and duration of harmful algal blooms. More information must be obtained through field and laboratory research and monitoring that involve innovative sampling techniques (such as genetics and acoustics), but data that already exist must be made accessible. And all data must have a temporal component so trends can be identified. As data are compiled, techniques must be developed to make certain that scales are compatible, to combine and reconcile data collected for various purposes with disparate gear, and to automate taxonomic changes. Information on biotic and abiotic elements of the environment must be interactively linked. Impediments to assembling existing data and collecting new data on marine biodiversity include logistical problems as well as shortages in finances and taxonomic expertise.

Adaptive bleaching: a general phenomenon

Laboratory and field data bearing on the adaptive bleaching hypothesis (ABH) are largely consistent with it; no data of which we are aware refute it. We generalize the ABH in light of these data and observations. The population of zooxanthellae within an organism is dynamic, the diversity of zooxanthellae is both surprising and difficult to ascertain, and field experiments demonstrate both turn-over in zooxanthella types and habitat-holobiont correlations. Dynamic change in symbiont communities, and the idea of an equilibrium or optimal community that matches the environment at a particular place and time, are concepts that underlie or emerge from much of the recent literature. The mechanism we proposed to explain responses to acute bleaching appears to operate continuously, thereby enabling the host-symbiont holobiont to track even subtle environmental changes and respond promptly to them. These findings enhance the potential importance of the ABH in the outcomes of acute bleaching, which can (1) accelerate this process of holobiont change, and (2) change the set of possible trajectories for how symbiont communities might recover.

Patterns of coral bleaching: Modeling the adaptive bleaching hypothesis

Abstract Bleaching — the loss of symbiotic dinoflagellates (zooxanthellae) from animals normally possessing them — can be induced by a variety of stresses, of which temperature has received the most attention. Bleaching is generally considered detrimental, but Buddemeier and Fautin have proposed that bleaching is also adaptive, providing an opportunity for recombining hosts with alternative algal types to form symbioses that might be better adapted to altered circumstances. Our mathematical model of this “adaptive bleaching hypothesis” provides insight into how animal-algae symbioses might react under various circumstances. It emulates many aspects of the coral bleaching phenomenon including: corals bleaching in response to a temperature only slightly greater than their average local maximum temperature; background bleaching; bleaching events being followed by bleaching of lesser magnitude in the subsequent one to several years; higher thermal tolerance of corals subject to environmental variability compared with those living under more constant conditions; patchiness in bleaching; and bleaching at temperatures that had not previously resulted in bleaching.

Envisioning a Marine Biodiversity Observation Network

Humans depend on diverse ocean ecosystems for food, jobs, and sustained well-being, yet many stressors threaten marine life. Extensive research has demonstrated that maintaining biodiversity promotes ocean health and service provision; therefore, monitoring the status and trends of marine biodiversity is important for effective ecosystem management. However, there is no systematic sustained program for evaluating ocean biodiversity. Coordinating existing monitoring and building a proactive marine biodiversity observation network will support efficient, economical resource management and conservation and should be a high priority. A synthesis of expert opinions suggests that, to be most effective, a marine biodiversity observation network should integrate biological levels, from genes to habitats; link biodiversity observations to abiotic environmental variables; site projects to incorporate environmental forcing and biogeography; and monitor adaptively to address emerging issues. We summarize examples illustrating how to leverage existing data and infrastructure to meet these goals.

Why do anemonefishes inhabit only some host actinians

SynopsisThe 25 species ofAmphiprion and one ofPremnas (family Pomacentridae) are obligate symbionts of 10 species of facultatively symbiotic sea anemones. Throughout the tropical Indo-West Pacific range of the relationship, a fish species inhabits only certain of the hosts potentially available to it. This specificity is due to the fishes. Five fishes occupy six sea anemone species at Lizard Island, Great Barrier Reef, Australia.Entacmaea quadricolor harborsP. biaculeatus, A. melanopus andA. akindynos. Adults ofPremnas occur deeper than about 3 m in large, primarily solitary actinians; juveniles may occupy peripheral members ofEntacmaea clones in shallow water. Specimens ofA. melanopus live exclusively in clonal anemones, which are found no deeper than 3 m. Most individuals ofA. akindynos inEntacmaea are juveniles, occurring shallow and deep, in solitary anemones or at the margins of clones. Interspecific as well as intraspecific social control of growth may be responsible for keeping fish small at clone fringes. Conspicuous specimens ofE. quadricolor depend upon their anemonefish to survive. Actinians cleared of symbionts disappeared within 24 h, probably having been eaten by reef fishes.Entacmaea, the most abundant and widespread host actinian at Lizard Island and throughout the range of the association, is also arguably the most attractive to anemonefishes. I believe its vulnerability to predation was a factor in its evolving whatever makes it desirable to fishes. Experimental transfers pitted fish of one species against those of another, controlling for ecophenotype of host, and sex, size and number of fish. Competitive superiority was in the same order as abundance and over-all host specificity:P. biaculeatus, A. melanopus, A. akindynos. At least three factors are necessary to explain patterns of species specificity - innate or learned host preference, competition, and stochastic processes.

Host selection by shrimps symbiotic with sea anemones: A field survey and experimental laboratory analysis

In three coastal areas of the Republic of China, we found six species of anemoneshrimps and five of host sea anemones in 13 symbiotic combinations; four of the 13 combinations were previously documented. In the laboratory, we tested host preference of the three common shrimps, Periclimenes ornatus Bruce, P. brevicarpalis (Schenkel), and Thor amboinensis (De Man), for anemones of three species. Periclimenes ornatus unequivocally preferred Entacmaea quadricolor (Ruppell and Leuckart), the only anemone with which it occurs in northern Taiwan and from which the experimental specimens had been collected. We therefore consider P. ornatus a specialist on E. quadricolor. Results of experiments with P. brevicarpalis and T. amboinensis were ambiguous: for both, outcome of some experiments was affected by identity of the anemone from which the shrimp had been collected, and in other experiments, T. amboinensis exhibited no preference. We therefore consider Thor a generalist symbiont, and P. brevicarpalis intermediate in specificity. In experiments to determine sensory cue(s) used by the shrimps to locate hosts, none could locate an anemone by vision alone. A significant proportion of shrimp of all three species was chemically attracted to anemones of at least one species; as in the preference experiments, source of the shrimp affected the results for P. brevicarpalis and T. amboinensis. The proportion of shrimp chemically attracted to an anemone in the experiments on sensory cues was lower than the proportion attracted in the host preference experiments. We, therefore, infer that shrimp use more than one sensory modality to locate a host anemone.