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Amer. Zool., 20:597-608 (1980) the Ecological Strategies of Sea

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Amer. Zool., 20:597-608 (1980) the Ecological Strategies of Sea Amer. Zool., 20:597-608 (1980) The Ecological Strategies of Sea Turtles1 John R. Hendrickson Departmentof Ecology and EvolutionaryBiology, University of Arizona, Tucson, Arizona 85721 Synopsis. By employing concepts such as "option" and "strategy" from game theory, this study derives an ecologically-oriented dendrogram of the probable evolutionary history and the present relationships of sea turtles. An "armored tank" strategy is seen as differ- entiating the first ancestral testudines from the stem reptiles and providing enduring advantages while simultaneously imposing basic restrictions on all later forms. A "flipper" strategy is postulated as basic to development of the sea turtle line, again imposing limi- tations while conferring selective advantage. Modern sea turtle species are grouped into three lineages representing strategies of habitat-type resource partitioning (a split-habitat, migratory pattern, a neritic residence pattern, and a pelagic residence pattern). Within the split-habitat, migratory group, further resource-partitioning by food-type separates the herbivorous Chelonia mydas populations from omnivorous Eretmochelysimbricata and the (apparently) carnivorous Chelonia depressa. Herbivory is seen as integral to the split- habitat, migratory strategy and C. mydas is considered the most "traditional" species, with the migratory habit secondarily lost in the other two. At the same time, the enhanced philopatry selected for by the migration strategy is viewed as responsible for the fact that C. mydasseems to have the most active race-formation ofthe three species. Further habitat- type partitioning in the neritic group, together with food-type partitioning, separates Caretta caretta from the two Lepidochelysspecies. L. kempi is represented as a consequence of Panamanian separation from L. olivacea following the last establishment of the isthmus as a land barrier. The third, pelagic residence, strategy is represented by Dermochelys coriacea, with little further differentiation of the line. The paper attempts to show that the evolution of sea turtles has been ecologically logical, that most conceivable niches for marine turtles are presently filled successfully, and that some predictions may be made with regard to gaps in our existing information. Introduction gained currency in discussion of ecological and recent Although I suspect that few biologists evolutionary phenomena; pub? lications 1957, is the earliest will be in much doubt about the subject of (Waddington, reference I to know this discussion, the title itself is trouble- happen about) speak some because of the use of the word "strat? of reproductive strategies, foraging strat? sex ratio and r and egies." The word stems from the Greek egies, gene strategies, K and stratos (army, host) and agein (to lead); its strategies, ecological evolutionary even and English meaning is either strictly military strategies, community ecosystem ... all with reference to entities or a transferred meaning from this, and it strategies and which we cannot make is misused here. It implies rational choice, processes agree rational choices. Louw and an example of correct usage is a com- (1979) correctly ment on the conduct of medical research points out that these usages of the word are and attributed by Bates (1951) to Alan Gregg "strategy" semantically wrong I of Rockefeller Foundation: "Strategy is the philosophically misleading?besides, that most of the users art of deciding when and on what one will strongly suspect mean trick for engage his strength, and tactics is the skill, "stratagem" (synonym: a rather than economy, promptitude and grace with achieving goal) "strategy" of What we are which one utilizes his strength to attain the (art deciding, etc). really about in most cases are ends chosen by strategy." Yet the word has talking complexes of adaptive evolutionary changes, each of which has relative positive and negative 1 values in any one ecological circumstance. From the Symposium on Behavioral and Reproduc? These adaptive tive Biology of Sea Turtles presented at the Annual complexes (phenotypes) are moves in an "existential Meeting of the American Society of Zoologists, 27- game" (Slo- 30 December 1979, at Tampa, Florida. bodkin and Rapoport, 1974), the rules of 597 598 John R. Hendrickson which are the principles of natural selec? tral lines, and include speculation in the tion and probability, and the object of form of "best guesses" when describing the which is to "turn resources into babies" ecological niches of extant species. The (Colinvaux, 1973) so that recruitment rate point is to present, not documented fact, at least equals death rate; past moves, pos? but a way of thinking about sea turtles sibly under different ecological circum? which is consistent with present evolution? stances, place varying limitations on pres? ary and ecological theory and which ent options; there are clearly defined squares in a logical way with sea turtle bi? losers, and the winner's reward is the op- ology as we presently understand it. If the portunity to keep on playing. There is no main framework is correct, it should ac- cognition, no rational choice, no predic? cept embellishment in detail without dam? tion?chance and necessity are all-control- age, and it may in itself predict the content ling. of various information gaps which now ex? Smith (1979) coins the term "phenotype ist. It is conceivable that a haemoglobin set" which I would have preferred to chemist or an endocrinologist might weave "strategy," but then goes on to give this the same fabric from different lines of evi? useful phrase a distinct hierarchical con? dence. text and to speak of the possible behavioral The General Chelonian "Strategy" strategies within the phenotype set. Mitch- ell and Williams (1979) note the difference Obviously, the extant species of sea tur? between the dictionary definition and the tles did not have unlimited possibilities for biological usage of the word "strategy," but developing the phenotypes (ecological pat? then go on to consolidate the usage. I re- terns and attendant morphologies) which luctantly agree with Murtagh (1979) that they show today; earlier "options" selected it is a necessary evil for lack of a simpler during their evolutionary history con- way to label evolutionary designs in con? ferred certain. genetic parameters within text with present-day thought on the sub? which the present species had to develop. ject. It has become jargon, and, while I la- Whether of positive, negative, or neutral ment the birth of this bastard in our family value in any one life circumstance, these of words, I have little choice but to ac? parameters from the past are basic to what knowledge its existence as a counterpart of we can detect today. the nomina conservanda of The selected the " systematics. prime "option" by 'When I use a word/ Humpty Dumpty sprawl-legged ancestral chelonian line said, in rather a scornful tone, 'it means more than 200 million years ago was the just what I choose it to mean?neither "armored tank" mode of body form?the more nor less'^Carroll, 1935). drastic modification of the entire anatomy In the discussion which follows, I at? to provide for encasement in a rigid box tempt an heuristic approach to the evolu? of defensive armor, presumably for de? tionary history of sea turtles, speaking as fense against large, toothed predators. of a complex game played with nature Pritchard (1979) gives a succinct account over the past 200 million years. I try to of early chelonian evolution and mentions show how, given various mixes of oppor- a number of important references. I cher- tunities and limitations implicit in each ish Romer's (1945) statement that: "The "move" of the game, and given the strat? chelonians are the most bizarre, and yet in egies "chosen" by the ancestral sea turtle many respects the most conservative, of line, a logical outcome seems to be the reptilian groups. Because they are still liv? present mix of species and their various ing, turtles are commonplace objects to us; niche spaces. Limited knowledge and lim? were they entirely extinct, their shells?the ited space permit no more than an account most remarkable defensive armor ever as? which hits only the high spots of the game sumed by a tetrapod?would be a cause for and is simplistic in many respects?I con? wonder. From the Triassic the turtles have sider only a few major aspects of body come down to present times practically un- form and attendant function in the ances- changed; they have survived all the vicis- Ecological Strategies of Sea Turtles 599 situdes which have swamped most of the mode to occupy new niche-space, it did reptilian groups and are in as flourishing produce a new set of life challenges a condition today as at any time in the which?under natural selection?had to be past." (In 1945, and with his long view of accommodated by further phenotypic paleontological history, Romer can hardly "strategy." The fossil record appears to be faulted for not recognizing the disas- give evidence of a number of separate at? trous effects of man on these otherwise- tempts by chelonian lines to invade the flourishing species.) sea; I believe that a single group was ul- The highly successful "armored tank" timately successful to the extent of having defense mode adopted by the ancestral representatives living today.2 chelonians involved a number of drastic Compared with the swamp, river, and anatomical modifications which committed land habitats in which the chelonian "ar? the line irrevocably to several important mored tank" strategy proved so successful, limitations on their evolutionary future the vast extent of the ocean environment (see Zangerl, 1969). About a dozen verte- and its relative monotony (lack of local spa? brae were eliminated from the trunk re? tial heterogeneity) must have presented a gion; the ribs were freed from the sternum major new challenge; mobility must have and spread laterally, and the limb girdles become a prime factor for selection.
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