1996. Thomas, J.D..
Chapter 24. Using Marine Invertebrates to Establish Research
and Conservation Priorities (357-388). In Biodiversity
II: Understanding and Protecting Our Biological Resources.,
M.L. Reaka-Kudla, D.E. Wilson, and E.O. Wilson, editors, National
Academy Press, 551 pp.
Current methods and applications used to identify and select
coral reefs for conservation efforts are seldom based on scientific
methodology. Instead, protection efforts are focused in a series
of coral reefs under direct or imminent threat of impact or alteration
resulting in a reactive policy approach. What is needed in light
of current changes occurring in coral reef systems are programs
with a strong preventative component designed to establish research
and conservation priorities in coral reefs before they come under
significant levels of threat. In this chapter I outline inadequacies
of current approaches to identifying and managing biodiversity
in coral reefs, recommend new ways to establish selective criteria
through taxonomic surveys and inventories, and provide an example
from a coral reef system of exceptional biodiversity in Madang
Lagoon, Papua New Guinea that, while not visually spectacular,
houses remarkable levels of marine invertebrate biodiversity.
Coral reefs have provided scientists with a rich source of facts and theory and have helped to forge fundamental views on evolution, biodiversity, and geology in the ocean realm. Scientists such as Darwin and Wallace recognized distinct patterns in biological distribution of coral reef organisms and attributed part of this pattern to geological events. With the acceptance of plate tectonic theory and accurate radiometric dating, these patterns have assumed biogeographic importance. Tectonic plate movements over fixed mantle hotspots produce accurate plate circuitry measurements (Yan and Kroenke, 1993), producing linear volcanic arcs and island archipelagos that encode a history of past geological events. According to Pandolfi (1992), a correlation exists between biogeographic pattern and geological history, and that modern marine distribution patterns can best be interpreted by incorporating the geological history of the area under study.
Numerous hypotheses have been proposed to explain biodiversity patterns in coral reefs. These include dispersal models (Kay, 1984), the Pacific plate vicariance theory (Springer, 1982), a "Pacifica" continental fragmentation theory (Nur and Ben-Avrahm, 1977), an expanding earth theory (Carey, 1958, 1976), and a variety of ecological explanations (Vermeij, 1990). However, the systematic study of comparative endemism levels of coral reef invertebrates to see which paradigm, or combination of paradigms, best explains biogeographical patterns remains to be engaged by the scientific community. Fundamental knowledge of species-level biodiversity is a prerequisite for such investigations.
Integrated studies of marine biodiversity are just beginning. To date, many taxonomic and most biogeographic studies of reef systems have been random and opportunistic, focusing on easily accessible islands. Taxonomic literature on tropical marine invertebrates is scattered, dealing mostly with large, spatially obvious components such as fish, scleractinian corals, and molluscs, thus creating a biological bias for larger organisms that may not be the best indicators or measures of biodiversity (Thomas, 1992). Platnick (1992) refers to the condition of concentrating on the obvious spatial components as the "megafauna bias." The most informative invertebrate groups in terms of biogeography are those without a dispersive (pelagic) larval stage. Wide dispersal capabilities can mask small-scale distribution patterns. Thus, animals with restricted distributions more accurately reflect levels of endemism. Information on those groups of invertebrates with restricted distributions is needed to integrate the many competing hypotheses on distribution mechanisms.
While coral reefs contain the highest levels of biodiversity in any ecosystem, virtually nothing is known about possible extinctions or natural trends in biodiversity.
Worldwide reports of changes in coral reef systems are largely anecdotal, based primarily on ecological research. Human impact is suspected as a contributing factor in these changes, but the interaction of natural change versus human-induced impacts remains speculative (Roberts, 1988; Williams and Williams, 1990). The rate at which marine areas are now being designated for protected status as sanctuaries, refuges, preserves, parks, etc., creates a problem in setting priorities and selecting among the many candidates suggested for protection. The goal of maintaining native levels of biological diversity requires that knowledge regarding historical sources and levels of biodiversity be known to the extent that enables competent decisions about research, conservation, and management action.
Above the generic level, the marine environment is more diverse than terrestrial systems (Ray and Grassle, 1991), yet most of what we know about biodiversity comes from studies in tropical moist forest systems. Biodiversity studies in marine environments, particularly the tropics, suffer from diffusion of scientific effort and an inability to identify critical sites based on biodiversity. Present management and conservation efforts in marine systems are driven mainly by concepts other than biodiversity, e.g., species of special human interest, areas of spectacular natural beauty, and economics (Thomas, 1993). Such an approach places the highest priority on unusual areas with regard to biodiversity levels and trends. In some instances where biodiversity surveys have been conducted, emphasis is on "spatially obvious" organisms such as fish and corals that may not be the best indicators to identify the processes of environmental change (Angermeier and Karl, 1994).
Successful management of marine ecosystems must rely on relevant
information about biodiversity levels and trends, information
that is almost entirely lacking in current management approaches.
Biodiversity survey and inventory programs designed to identify
and select those marine areas of highest scientific value are
needed. Resources such as existing collections in natural history
museums and coordination of ongoing taxonomic surveys and inventories
need to be organized according to selective criteria. The limited
number of active field systematists and taxonomists currently
working in marine environments must be deployed effectively.
Training programs to allow a wider dissemination of taxonomic
expertise are also a critical element in a unified approach to
marine biodiversity. Use of data from disciplines outside taxonomy
and systematics must be incorporated into a marine biodiversity
program for the 21st century. Information from fields such as
plate tectonics, paleogeography, cladistics, ecology, and anthropology
must be brought together in concert to fully understand what is
happening in marine environments and what can be done to improve
our understanding of system processes. At the administrative
level, specific shifts in policy goals to include reliance on
preventative rather than reactive management strategies must become
a priority (Angermeier and Karl, 1994).
TAXONOMY AND BIODIVERSITY IN CORAL REEFS
Available taxonomic data tell us more about varying attention given to different groups of animals, the "taxonomy of taxonomists," than about the level of taxonomic knowledge (May, 1994). Information summarized by May (1994) illustrates the great disparity of attention received by different groups. Roughly one-third of taxonomists work on plants, while the remaining two-thirds spilt roughly equally between invertebrates and vertebrates. The estimated total number of vertebrate species is 40,000; of plants 300,000; and invertebrates about 1 million, with estimates up to 10 million (Grassle and Maciolek, 1992). Therefore, for every n vertebrate taxonomists, there are 0.1n plant taxonomists and 0.01n specializing in invertebrates. When we consider that a majority of invertebrate taxonomists study a single group, the insects, the great disparity within the current taxonomic work force that specialize in marine invertebrates becomes apparent. Estimates of millions of new species at an estimated novelty rate of 40-80% for undescribed taxa places an absolute accounting of all marine species outside the realm of possibility in any time frame to make any significant difference in the current biodiversity crisis. I suggest what is needed is sufficient efforts directed to taxonomic inventories of selected groups of bioindicators that are chosen by established taxonomic protocols. Such a selection process would target not only on areas that are threatened or in crisis, but would incorporate objective scientific criteria to predict possible centers of evolutionary diversification that act as genetic sources for existing biodiversity (Thomas, 1992). Approaches that focus existing taxonomic expertise on areas that may be historical sources of biodiversity will help us understand how species-level biodiversity patterns are maintained and replenished.
Despite extensive reports of large-scale change in coral reef systems, the scientific, conservation, and management community does not currently have the capability to investigate every region or problem that gains public attention.
Lack of comprehensive marine biodiversity protocols to identify areas of particular scientific, conservation, and management value have lead to a diffuse approach to the systematic investigation of coral reef biodiversity. Most research taking place now on reefs is ecological in nature, with little or no systematic effort directed to taxonomic surveys and inventories of reef systems worldwide. This lack of focus is further compounded by the shortage of experienced field systematists and taxonomists. Training programs to increase the level of personnel available for surveys and inventories in coral reefs while frequently discussed, have resulted in few active programs. The need is especially acute for the marine invertebrates, particular groups of which are sensitive indicators of change in coral reefs (Thomas, 1993).
The scientific community must develop a process to assess biodiversity that uses selective criteria and sets priorities, in effect a form of "environmental triage." Every reef system cannot be investigated with current personnel and funding levels. Difficult choices must be made that maximize existing personnel, equipment, and organizational structure. Numerous coral reef sites already enjoy protected status, while numerous additional sites are being suggested for protection. In many cases the site selection process is driven from a perception that a particular site is in a stressed or deteriorating condition. This approach focuses limited resources on a series of crisis situations and perpetuates a reactive management posture. Ideally, the scientific community should provide theoretical guidance and taxonomic information regarding appropriate site selection. The conservation community, capable of rapid response and adept at raising public awareness, should then use scientific data to target specific sites or regions for protection. The resource management community in effect does not become established until an administrative structure is created. Every effort should be made to incorporate high-quality science in management approaches to documenting biodiversity. Coral reef sites that enjoy preexisting protected status should be encouraged to adopt programs that adequately document biodiversity levels if such information is lacking. These major groups--scientists, conservationists, and resource managers--must work in concert along established guidelines to maximize the application of diffuse resources and funding to help identify and protect coral reefs.
Priorities vary depending on individuals, agencies, and processes
involved. For example, taxonomists place the highest priority
for research on reefs that contain the highest levels of species-level
biodiversity or are the least impaired or impacted by man. Conservation
groups may target areas that house species of special interest,
or endemics, while resource managers might focus on reef areas
that seem threatened by human impacts. It must be emphasized
that most coral reef systems, except for limited areas in the
Caribbean, and the Great Barrier Reef in Australia, are claimed
by developing countries with little or no scientific or administrative
resources. Therefore, any system of establishing priorities must
take into account a variety of factors.
Developing selection strategies:
(1) Identify centers of evolutionary diversification, or major centers of genetic biodiversity by surveys and inventories, existing museum collections, and cladistics.
(2) Target areas where geological history indicates a history of vicariant events that might lead to high levels of endemic species or areas of composite biodiversity.
(3) Implement surveys and inventories to determine biodiversity baselines. While entire protected areas must be surveyed initially, ongoing monitoring of biodiversity could be restricted to special protected areas such as replenishment zones located within a protected area (Bohnsack, 1993). It is important to stress that our understanding of biodiversity in coral reefs is rudimentary, therefore, initial surveys must incorporate a representative cross section of the entire area. Once biodiversity levels have been determined, decisions can be made as to the location of permanent monitoring sites and schedules.
(4) Publish comprehensive identification guides to biota. Most information regarding marine invertebrates from coral reefs is uneven in terms of taxonomic and geographic coverage. Computer-based identification manuals should be a priority for development and distribution.
(5) Establish parataxonomic training programs to augment survey
and inventory capabilities. Basic biodiversity surveys and inventories
of many protected areas remain to be initiated and are not currently
a primary management concern. Two of the largest marine protected
areas in the world, the Florida Keys National Marine Sanctuary
and the Great Barrier Reef Marine Park, Australia, have yet to
institute systematic biotic inventories, despite stated management
concerns to "monitor biodiversity."
The term "hotspots," a geological term originally used to describe fixed volcanic sources, has recently gained popularity in the biological sense to indicate areas of high biodiversity. This unfortunate amalgamation of terminology confuses the original use of the term. In this chapter, the term "hotspot" is used in a geological context.
Recent efforts to test the hypothesis that geologic hotspots accurately record the passage of lithospheric plates have generally been accepted (Duncan, 1981, 1991; Duncan and Richards, 1991; Fleitout and Moriceau, 1992; Morgan, 1980, 1983). If hotspots are indeed fixed with respect to one another over significant periods of geological time then they would constitute an accurate frame of reference that could be used to determine precise lithospheric plate movement and allow reconstruction of plate positions back into time. Morgan (1981) estimated that hotspots migrate less than 5 mm per year relative to each other. Hotspot trails have been clearly delineated for the time span zero-100 million years, thus allowing reconstruction of southwest Pacific tectonic elements. This is done by first determining linkages between individual rifted continental fragments and island arcs and then by assigning correct plate motion as derived from hotspot models, thus providing a "plate circuit" (Yan and Kroenke, 1993). Such approaches to tectonic plate circuitry allow use of this methodology in visualizing complex movements of tectonic plates through space and time. Yan and Kroenke (1993) provide a CD-ROM containing animated sequences of the southwest Pacific plate movements from zero-100 million years ago in 0.5-million year increments that helps visualize complex plate movements. This information provides additional insight on how and when marine organisms may have dispersed or been constrained by various geological events over the last 100 million years.
The island of New Guinea, composed of Papua New Guinea (PNG) and Irian Jaya, is located on the leading edge of the Australian plate that has been moving rapidly northward. The country of Papua New Guinea comprises a landmass slightly larger than California (457,000 km2). The north coast of PNG has been formed by collision events along the north edge of the Australian plate and subsequent "docking" of the east Papuan composite and other terranes in mid-late Miocene 15-20 million years ago (Pigram and Davies, 1987). I suggest the composite marine fauna of the Madang Lagoon is the result of the accretionary process along the north coast of PNG, involving rapid volcanic uplift as islands, archipelagos, submerged plateaus, and continental terranes are thrust rapidly upward in the collisional process. This process differs from other areas of the circum-Pacific where terranes are predominantly of oceanic affinity.
This "docking" process introduced a number of previously discrete biotic assemblages that then intermingled with established floral and faunal elements.
Reefs in the Madang Lagoon illustrate the importance of taxonomic
data in identifying areas of scientific concern. At first glance
the reefs in the Madang Lagoon appear "less spectacular"
(in a visual sense) than their southern counterparts on the Great
Barrier Reef. However, taxonomic surveys of marine invertebrates
suggest that Madang reefs are some of the most biologically diverse
reefs yet documented. Sustaining native levels of biodiversity
in these reefs as a potential genetic seed source for other South
Pacific reefs is important in the larger context of regional biodiversity.
MARINE INVERTEBRATES OF THE MADANG LAGOON
Over the past several years, increasing numbers of scientists have focused their research efforts in the Madang Lagoon in an attempt to document the unusual levels of biodiversity found there. The following information summarizes briefly some of the results and trends that have been documented. A more complete discussion can be found in the Proceedings of the Seventh International Coral Reef Symposium (Richmond, 1992).
While much attention has been given to scleractinian corals of
eastern Australia (Veron & Pichon, 1976, 1979, 1982; Veron
and Wallace, 1984; Veron et al., 1977) the nature of reef corals
from northern New Guinea was not well understood due to lack of
distributional data. Potts suggests the Madang Lagoon may prove
to be the single most diverse site for scleractinian corals (Potts,
personal communication, 1994). Hoeksema's recent treatment of
fungiid corals (1992) represents the only reef coral family on
which detailed distribution and taxonomic data are available for
the Indo-Pacific. According to Hoeksema (1992), northern New
Guinea appears to have the highest fungiid biodiversity (39 species),
with a fauna most similar to that of the Philippines and eastern
Indonesia (37 species). Diversity levels of eastern Australia
were the next highest (31 species), followed by western Australia
and Taiwan (26 species), northwest Java (25 species), southern
Papua New Guinea (24 species), and northeast Borneo (19 species).
Hoeksema (1992) stresses that the generic diversity of hermatypic
corals in the Indo-Pacific region is quite large, and that lists
of genera are not as informative as species diversity and monospecific
genera. Because all taxon rankings above the species level are
arbitrary, lists that include species and monospecific genera
are more likely to reflect evolutionary history, and thus allow
for a more precise comparison of biodiversity levels.
Winston (1988) estimates that only 50% of the octocoral fauna
from the Indo-Pacific is known at present. Like most other marine
invertebrate groups, our basic taxonomic knowledge of the Indo-Pacific
octocorals is poorly known (Williams, 1992). A recently described
species of octocoral from the Madang Lagoon was unlike any species
of the genus previously recorded (Bayer, 1994).
Thomas (1992) found the coral reef amphipod fauna of the Madang
Lagoon to exhibit exceptional levels of species diversity. The
amphipod fauna of Madang reefs is a composite, consisting of approximately
180 species, 60% of which are new to science and exhibit multiple
biogeographic affinities. The Madang Lagoon amphipod fauna is
taxonomically distinct from other South and Indo-Pacific sites,
and indications are that amphipod biodiversity on the north coast
of PNG exceeds that of any coral reef area studied thus far.
However, many coral reef systems in the Indo-Pacific have never
been systematically analyzed for smaller crustaceans. The author
herein suggests that future biodiversity inventories and surveys
be undertaken with regard to selective criteria. Due to the limited
dispersion capabilities and habitat specificity of amphipods,
amphipods may be of use in biogeography and environmental monitoring
in coral reef systems.
Messing (1992) reported 39 species of comatulid crinoids from
the Madang Lagoon, and with limited sampling, found the crinoid
fauna of the Madang Lagoon comparable to other more intensively
studied sites such as Lizard Island and Davies Reef (Australia,
Great Barrier Reef), and Palau.
Working in the Madang Lagoon, Gosliner (1992) found that the north coast of PNG supports a more diverse fauna of opistobranch gastropods (538 species) than has been reported from any single geographical area studied thus far. The next richest tropical site is Guam (395 species), followed by Hawaii (244 species), the Caribbean (232 species), and Japan (184 species). The importance of Gosliner's faunal records are significant because of intensive field efforts in numerous tropical localities by snorkeling and SCUBA that enable comparative studies. Other areas that are known or suspected to house high diversity have not been adequately studied or sampled to allow comparisons of opistobranch biodiversity (Gosliner, 1992).
Kristian Fauchald (personal communication, 1994) reported the
polychaete fauna of the Madang Lagoon exceeded that of any area
yet sampled. Clyde Roper and Mike Sweeney (personal communications,
1994) reported similar findings for cepahlopod molluscs.
In a preliminary survey of the marine algae of the Madang Lagoon, Mark and Diane Littler (personal communications, 1994) reported that not only were there more species of algae collected, but the number of undescribed species surpassed that of any region previously sampled.
The geological history that may have contributed to the extraordinary levels of marine invertebrates in the shallow waters of the north coast of PNG may extend into deeper waters of the region. Studying a collection of deep-sea crustaceans from the Bismarck Sea region (1,200 m), Austin Williams of the National Marine Fisheries Service, Systematics Lab, reports unusual levels and types of biodiversity (personal communication, 1994). More investigation on the deep-sea component of this region are warranted in light of this preliminary information.
Most current approaches to protecting biodiversity place emphasis on areas or species of special human interest and natural beauty, or areas with novel biological features (endemics). Therefore, highest value is placed on unique and unusual, with little regard to the nature and quality of biodiversity on a larger scale. It is imperative to develop priorities by geological past, ecosystem, region, and available specialists.
(1) Initiate surveys and inventories that document biodiversity in those areas (known or suspected to be) of extraordinary scientific or conservation value as determined by a rigorous selection process.
(2) Establish biologically based monitoring programs using selected groups of organisms that have an established reputation as high-quality bioindicators.
(3) Inventory existing historical collections from reef systems deposited in natural history museums. Many museums house historical collections of organisms from tropical regions. Such collections could serve as a "biodiversity baseline."
(4) Publish primary taxonomic monographs, identification guides, keys, and manuals, especially computerized, graphically based manuals for nonspecialists and resource managers.
(5) Develop parataxonomic training programs targeted for specific taxa and geographic regions.
(6) Implement scientific taxonomic training programs coordinated through a network of natural history museums, academic institutions, government agencies, and other organizations.
Oceanic islands and their associated coral reefs have provided scientists with a wealth of biogeographic information. Biodiversity levels on modern coral reefs provide a window on past evolutionary events detailing the correlation between biogeographic pattern and geological history. Reefs of the Madang Lagoon in Papua New Guinea exhibit unprecedented levels of biodiversity exceeding all other reef systems studied thus far. Because of this, the Madang Lagoon represents a scientific and conservation resource of the highest priority. However, while biodiversity research on coral reefs is in its infancy, the need for this information is acute.
Within PNG a complex pattern of land ownership and negligible developmental pressures and resource exploitation have allowed the reefs to remain relatively unaffected by anthropogenic impacts. That situation promises to change as the country seeks to modernize its largely subsistence economy, and as rich mineral deposits and timber resources are developed. The rugged topography of the interior of the country virtually assures that the majority of this developmental pressure and impact will be in the coastal region, adjacent to reefs.
The reefs of the north coast of PNG provide an unequaled opportunity
to study marine biogeography and in what is probably a major source
of biodiversity for a large area of the South Pacific. Research
and conservation efforts must be focused on this invaluable biotic
resource before significant impacts occur to alter yet unstudied
and undocumented groups of organisms.
The author wishes to thank the Christensen Research Institute in Madang for research support on coral reef systems in an around the waters of Madang. Former Director Matthew Jebb and current Director Larry Orsak have graciously provided facilities and equipment to marine researchers working in the Madang region.
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Coral reef systems are undergoing unprecedented changes throughout the world. A major problem in understanding this change is the almost total lack of information on biodiversity from reef systems. Comprehensive biotic surveys and inventories at selected sites must become a priority for the scientific, resource management, and conservation communities. Currently diffuse scientific efforts to document biodiversity must be focused. A comprehensive plan to establish conservation priorities and develop selective criteria for selection of protected areas based on identifying areas of evolutionary diversification rather than aesthetics should be implemented. Marine protected areas already designated should initiate biodiversity surveys and inventories as a biological measure of management success.
Successful documentation of coral reef biodiversity levels can be achieved only with a concerted effort by researchers. Difficult choices must be made on which areas should be protected and why. Protection of areas primarily for aesthetics or economic reasons does not assure conservation of less spectacular sites that may serve as sources of biodiversity at varying seascape levels. Current planning processes incorporate actions that emphasize preventative rather than reactive strategies. Long-term protection of coral reefs will be realized only with the understanding of where and how biodiversity levels are established and maintained.
Marine invertebrates from coral reefs in the Madang Lagoon on the north coast of Papua New Guinea exhibit the highest levels of biodiversity yet documented. This unprecedented accumulation of biota may be explained in part by the complex geological past of the region which rests on no less than five active lithospheric plates. Biogeographic affinities of the Madang Lagoon fauna reflect the composite nature of the accumulated collisional forces that shaped the region.