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Biodiversity Implications Is Biogeography: For FEATURE BIODIVERSITY IS BIOGEOGRAPHY: IMPLICATIONS FOR CONSERVATION By G. Carleton Ray THREE THEMES DOMINATEthis review. The first is similarities, as is apparent within the coastal zone, that biodiversity is biogeography. Or, as Nelson the biodiversity of which depends on land-sea in- and Ladiges (1990) put it: "Indeed, what beyond teractions. biogeography is "biodiversity' about?" Second, "Characteristic Biodiversity"--a Static View watershed and seashed patterns and their scale-re- A complete species inventory for any biogeo- lated dynamics are major modifiers of biogeo- graphic province on Earth is virtually impossible. graphic pattern. And, third, concepts of biodiver- In fact, species lists, absent a biogeographic frame sity and biogeography are essential guides for of reference, can be ecologically meaningless, be- conservation and management of coastal-marine cause such lists say little about the dynamics of systems, especially for MACPAs (MArine and environmental change. To address such dynamics, Coastal Protected Areas). biodiversity is best expressed at a hierarchy of Conservation and conservation bioecology have scales, which this discussion follows. entered into an era of self-awareness of their suc- cesses. Proponents of "biodiversity" have achieved Global Patterns worldwide recognition. Nevertheless: "Like so Hayden el al. (1984) attempted a summary of the many buzz words, biodiversity has many shades of state-of-the-art of global, coastal-marine biogeogra- • . the coastal zone, meaning and is often used to express vague and phy at the behest of UNESCO's Man and the Bios- which is the most bio- ill-thought-out concepts" (Angel, 1991), as phere Programme (MAB) and the International reflected by various biodiversity "strategies" Union for the Conservation of Nature (IUCN). The diverse portion of (WRI, IUCN, UNEP, 1992; Norse, 1993), wherein overall objective was to review coastal-marine bio- the overall goals are clear, but in which the "game Planet Earth, and geographic provinces, following Udvardy (1975) for plan'" is not. Furthermore, too little attention is the land. Purportedly, the result would enable these within which many given to biodiversity at all levels from genes to organizations to incorporate "representative," marine ecosystems (Solbrig, 1991), as well as to how this and coastal protected areas into a "global network" conservation problems broad spectrum of diversity may be encapsulated of protected areas. A fundamental requirement for most urgently lie. as "the complex mosaic of the variety of life forms any classification is that it should be comprehensive on Earth" (Angel, 1991). and comparative. Thus our physical classification This paper focuses on biogeographic pattern at described reasonably symmetrical oceanic and global to subregional scales and on the functional coastal realms (Fig. 1); that is, the oceanic realms dynamics of keystone species and ecosystems. are circumferential and the trailing and collision Emphasis is on the coastal zone, which is the most coasts are generally on the east and west sides of biodiverse portion of Planet Earth, and within continents, respectively. However, the coastal, biotic which many conservation problems most urgently provinces, derived largely from Briggs (1974) (Fig. lie. I will first describe coastal-zone biodiversity 2) cannot be symmetrical, as endemic flora and from a static point of view, then take up some eco- fauna differ from place to place and, thus, describe logical dynamics, followed by a discussion of con- unique provinces, taxonomically speaking. servation and management implications. Addition- This classification shows that the coastal zone ally, I will submit that the special features of is geomorphologically and ecologically distinct marine and terrestrial systems are subsumed by the from both upland areas and the open sea, follow- ing Ketchum's (1972) definition that the terrestrial portion of the coastal zone includes the continental G. Carleton Ray, Department of Environmental Sci- plains and the seaward portion includes the conti- ences, University of Virginia, Charlottesville, VA 22903, nental shelves. The coastal zone has been con- USA. formed by the rise and fall of sea level from the 50 OCEANOGRAPlfY*VoI.9. No. 1"1996 Fig. 1: A global classification of coastal and marine environments. Oceanic areas are indicated by let- ters. Coastal areas are distinct biogeographically and are shown symbolically as colored bands around the continents. Cross-hatched areas are enclosed seas. Note the general symmetry of both oceanic and coastal areas. See original text for full explanation. From Hayden et al. (1984); terrestrial biogeographic provinces, after Udvardy (1975). Pleistocene to present (Fig. 3). With respect to the "characteristic" diversity of ecoregions and their biodiversity, this zone has disproportionate global species assemblages. This would facilitate descrip- significance. For example, of the approximate tions of what is special about each region, as well as 22,000 fish species, about half are restricted to the enable better comparisons among regions inter alia. • . it would seem 8% of Planet Earth that is coastal zone, whereas more appropriate to -40% are freshwater and only -10% oceanic (Nel- Watersheds, Seasheds, Habitats, and Niche son, 1984), i.e., occurring beyond the continental Subglobally, a biogeographic hierarchy is facil- concentrate on what slope. Other noteworthy coastal-zone species are itated by conceiving the coastal zone as an ecotone may be termed the right, bowhead, and gray whales, many polar sea between land and sea, but one that is distinguished birds, seals, walruses, and a host of invertebrates. by strong spatial and temporal gradients, both "characteristic" diversi- Another way to address global biogeographic along and across this zone (Ray and Hayden, structure, besides biogeographic provinces, is 1992) and within the water column (Holligan and ty of ecoregions and through gradients in species richness. Whether de Boois, 1993). This zone is uniquely where land, their species assem- there is a latitudinal gradient in species richness is sea, and atmosphere intensely meet. Complex in- a matter of debate (Angel, 1991; Clarke, 1992; teractions produce a hierarchy of mosaics of habi- blages. This would Rohde, 1992; Rex et al., 1993). At the very least, tats, as represented by watersheds, estuaries and facilitate descriptions such gradients are heavily taxon dependent; that is, lagoons, wetlands, and coastal islands, within marine mammals, such as seals, are most diverse which the smallest biogeographic scale is that of of what is special in polar regions, whereas fishes are most diverse the species "niche." Thus both physical and bio- about each region, as in the tropics (Ray, 1988). Some taxa have specific logical approaches are necessary to distinguish hi- requirements favoring low temperatures--e.g., erarchical, biogeographic relationships. well as enable better many pinnipeds, cetaceans, and fishes--so that A physical approach is represented by Ray and comparisons among clearly defined gradients in species richness are Hayden (1992), who formulated a simple clas- difficult to evaluate. Comparative biogeographic sification for within-province, coastal-zone struc- regions inter alia. studies at different latitudes are needed to clarify ture for defining "coastal units in which the con- whether biotic patterns result from historical, evo- nections among mass, energy, and biota across the lutionary, behavioral, or physiological factors, or, coastal zone are stronger than along the coast, es- in all probability, from combinations among them. pecially those that are associated with fluvial inter- From both points of view, it would seem more actions" (Fig. 4A). These units allow the identi- appropriate to concentrate on what may be termed fication of several fundamental types of watershed OCEANOGRAPHY'VoI.9, No. 1"1996 5 1 The smallest-scale levels of the biogeographic hierarchy are species-specific habitat and niche. With respect to the former, such habitat types as rocky shore, estuary, lagoon, various types of ben- thos, etc., appear about equally represented world- wide; that is, there appears to be no gradient in coastal-marine habitat diversity from the tropics to the high latitudes. In fact, habitat diversity may in- crease in the polar regions due to the complicating effects of seasons and sea ice. The inference is that ecological complexity and habitat variety are not readily related to species diversity and/or ecosystem resiliency. Pattern and Process--a Dynamic View Little can be said about conservation and man- agement until and unless ecosystem dynamics are incorporated into ecological descriptions. For ex- ample, Steele (1991) has defined functional diver- sity as "the variety of different responses to envi- ronmental change, especially the diverse space and time scales with which organisms react to each other and to the environment." This approach Fig. 2: A coastal-zone classification for eastern is concordant with the major theme of the North and Central America. Letters indicate phys- IUBS/SCOPE/UNESCO Diversitas program, ical regions: (A) Subpolar; (B) Eastern temper- ate; (C) Eastern Subtropical; and (D) Tropical. Numbers are for biogeographic regions after Briggs (1974); (9) Acadian; (8) Virginian; (7) Carolinian; (6) Louisianian; (5) Caribbean, and (4) West Indian. Note that the physical and bio- geographic regions reasonably approximate one another. See original text for full
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