* Your assessment is very important for improving the workof artificial intelligence, which forms the content of this project
Download Possible Biological Consequences of Plate Tectonics
Post-glacial rebound wikipedia , lookup
Earth's magnetic field wikipedia , lookup
Spherical Earth wikipedia , lookup
Anoxic event wikipedia , lookup
Geomagnetic reversal wikipedia , lookup
Age of the Earth wikipedia , lookup
History of Earth wikipedia , lookup
Abyssal plain wikipedia , lookup
History of geomagnetism wikipedia , lookup
Geological history of Earth wikipedia , lookup
History of geology wikipedia , lookup
Possible Biological Consequences of Plate Tectonics Malcolm C. McKenna PLATE TECTONICS Hardly anyone nowadays is unaware of "continental drift," so thorough has been the coverage of the subject in all available media. Biologists, however, may not all be fully familiar with the basic principles of plate tectonics, of which "continental drift" is but one ~pect, so it may be useful to attempt to list the basic principles here. Geophysicists now conclude that these principles have been in operation for at least the last three billion years and probably for the whole history of the earth. A few authors still dispute these principles (e.g. Meyerhoff and Meyerhoff 1972), and no doubt there have been excesses of enthusiasm by certain proponents of "the new global tectonics," but even at this stage in its conceptual development the generality of plate tectonic theory has convinced most geologists of its essential validity. The author is Frick Curator in the Department of Vertebrate Paleontology, The American Museum of Natural History, and Associate Professor of Geology, Columbia University, New York, N.Y. September 1972 The fundamental idea of plate tectonics is that the ocean floor moves. The lithosphere of the earth (approximately the top 70 km) consists essentially of a system of large, nearly rigid plates added to by upwelling of new, relatively hot lithosphere along oceanic ridges and destroyed by subduction and remelting of old, relatively cold lithosphere beneath oceanic trenches. Continents float like scum on oceanic lithosphere and may be carried passively about, sometimes bumping into one another, sometimes breaking apart and reforming, but always part of a dynamic system. Collisions as well as separations may occur. The continents do not "drift" through the lower part of the lithosphere but are carried with it. Evidence for such a view stems from many sources; one of the joys of plate tectonics is how well the concept is consistent with data from many seemingly unrelated disciplines. Direct observation of plate creation and relative motion on the order of a dozen centimeters a year is presently out of the question along most of the globegirdling length of the mid-ocean ridge system, but direct measurements in Iceland, where the Atlantic ridge is above water, have led to the conclusion that the ridge is indeed spreading there (Decker, Einarsson, and Mohr 1971). Another method of direct measurement, involving laser reflections from corner reflectors on the moon, may ultimately enable direct measurement of relative motion of distant places on the earth's crust, but in view of the present uncertainties of 30 cm or so in laser ranging to the moon (Hammond 1970) it may be some time before the test would yield usable results. plausible coastal matches which form an important body of evidence that separation of some major land areas indeed occurred, though such fits say nothing concerning the timing of separation and contribute no information about continental collisions. Matching geology along such coasts has added muscle to the geometric arguments, and has added information about timing as well, though at a low power of resolution. Until the 1960's most non-biological arguments about "continental drift" centered about geometrical fits and geological similarities of opposing coasts. Least-squares fits of either the present coasts or of various depth contours are 6O"W ",'W o· ... AU'H B Fig. 1. A. The location of the Reykjanes Ridge and the area of B. The 1,()()()'fathom submarine contour is shown together with the Coastal Matching S()()'fathom contour for the Rockall Bank. B. Contoured symmetrical positive magnetic The classic fit of the southeast coast anomalies recorded over the Reykjanes Ridge. of South America to the Gulf of Guinea Areas of positive anomaly are shown in black. and southwest coast of Africa is the (from Vine in Proc. Amer. Phi/os. Soc. 112,5 most obvious of a growing number of after Heirtzler, Le Pinchon and Baron, 1966.) 519 Downloaded from http://bioscience.oxfordjournals.org/ at Penn State University (Paterno Lib) on September 18, 2016 Geology is currently undergoing an upheaval as profound as Darwin's reorganization of biology in the Nineteenth Century. The purpose of this essay is to examine briefly the interface between these two sciences with an eye toward new possibilities of synthesis and new ways of viewing the parent subject, natural history. A current research trend, now accelerating in activity but still inadequately pursued, deals with the biological consequences of the geological evolution of the earth as deduced from recent theoretical advances in geophysics. Inquiry into the no-man's-land between geophysics and biology is particularly encouraging at present: not since Darwin and Wallace have there been such great opportunities to mesh geological concepts of the evolution of the earth with biological ideas about the evolution of organisms. now common arguments in the literature (Bullard et a1. 1965, Merservey 1971). Magnetics The two most powerful arguments for large scale plate motions in the lithosphere are derived from two different aspects of the history of the earth's magnetic field. These are the "polar wander paths" and "zebra stripes" (Fig. 1) of the geophysical literature. Measured from each lithosphere plate, a sequence of rotational pole positions, each calculated by averaging a number of relatively contemporaneous magnetic pole positions, does not fall at random but rather in a definite "wander path" progressing in a definite direction when the calculated rotational positions are arranged according to geological age (Irving 1964, Creer 1970). The "wander paths" are somewhat different for different plates, but parts of the paths may be indistinguishable. Because the earth has only one rotational axis, the conclusion is that the plates have on occasion moved with respect to each other, as well as with respect to the rotational pole. Comparison of the calculated rotational "polar wander paths" from various plates then generates a history of plate latitudes and plate rotations (Fig. 2). The lithosphere as a whole may also "wander," and a generally westward motion relative to the deeper interior has been postulated in theoretical work (Palmer 1968, Bostrom 1971). The Eulerian pole of rotation of the lithosphere would be expected to fall close to the rotational pole of the earth as a whole. However, to the extent /~ r o. \3 ®2 ~®' T. d I 03 \02 ~o' j o' /3 /05 I ; 03I O. ~o. \, <:J Y2 D o~ 0' t o~., Ts Fig. 2. Plate motions deduced from superposed "polar wander paths;" Left: Both plates move together from time T1 to\ T4.~producing the same path. Center: Relative motion has separated the ancient wander paths, but a single position is obtained at time TS' Right: Both paths have the same shape from Time Tl to T4. but not thereafter. Circles indicate poles calculated from the square plate; dots indicate poles calculated from the triangular plate. 520 earth's present field, cause the observed anomalies in the present magnetic field from place to place. Sequences of Pleistocene and Pliocene oceanic anomalies can be correlated convincingly with data sequences of magnetic polarity changes on continents; older oceanic anomalies deep under a cover of sediment or distant from originating ridges must be identified by extrapolation, or directly dated with various degrees of resolution by borehole sampling of the volcanic basement or by dating the immediately overlying sedimentary cover. Anomaly sequences can be and have been extensively correlated from ocean to ocean and their symmetry on either side o( originating ridges is truly remarkable (Fig. 3). At present the ocean floor is rapidly being mapped at a power of resolution sufficient to delimit all the major lithosphere plates and many minor ones as well. Transform Faults, Plate Motion, and Continental Winnowing Individual plate motions also follow Eulerian geometry, with instantaneous poles of rotation for each plate. This is because plate motions occur on the surface of a sphere. If angular rotation tends to be constant, then either the rate of creation increases as a cosine function of latitude with respect to the instantaneous pole of rotation or breaks occur. Such breaks, the oceanic fracture zones, fall on arcs centered on the instantaneous pole of plate rotation and are at right angles to the "zebra stripes" of magnetic anomaly maps, i.e. parallel to the spreading direction. Secondary creation of new oceanic crust can occur along fracture zones between plates with substantially different poles of rotation. On a global scale, the geometry of plate motion is quite complex (McKenzie and Morgan 1969, Menard and Atwater 1969). Fault offsets along fracture zones between segments of oceanic ridges where new crust is being formed have been called ridge-ridge transform faults (Wilson 1965, McKenzie and Parker 1967, Atwater 1972). Relative motion of the plates on opposite sides of these faults is the reverse of that which might have been expected (rom ridge offset under the old concept in which the ocean floor did not spread (Fig. 4). Only with ocean-floor spreading from each of the offset oceanic ridge segments do these reversed offsets make sense. That BioScience VoL 22 No.9 Downloaded from http://bioscience.oxfordjournals.org/ at Penn State University (Paterno Lib) on September 18, 2016 that in the past the two poles have no~ coincided, a latitudinal component of "general lithosphere wander" should be detectable and should have a decipherable history. Relative longitude changes are not directly determinable by analysis of "polar wander paths" alone, but are accessible through studies of "hot spots" (e.g., Morgan 1972). "Hot spots" are migrating volcanic centers that appear to overlie rising plumes of material beneath the lithosphere plates. From time to time vulcanism breaks out at the surface, but in the meantime the plate has moved, generating a "wander path." "Hot spot wander paths" place important constraints on conjectures of plate motions. "Zebra stripes" are patterns appearing on maps which depict contoured magnetic anomaly data derived from magnetometers towed over oceanic crust. These were at first a great puzzle, but did not long remain so. They have turned out to be the result of alterations of the earth's past magnetic field polarity, evidence of which is frozen in new lithosphere as it appears and solidifies along oceanic ridges (Vine and Matthews 1963). The remanent magnetic field of new lithosphere appearing at such ridges reflects the geomagnetic polarity in effect when the new lithosphere cools below its Curie point. Because the polarity reverses often, still younger lithosphere will exhibit the opposite polarity, and so forth, resulting in bilaterally symmetrical, outward-moving bands of alternatingly polarized lithosphere which, as many authors have remarked, resemble two nearly identical tape recordings moving away from a recording head in opposite directions. These, superposed on the WEST KM 200 200 400 • 400 • EAST KM i r GAMMAS GAMMAS '~t yrvvM "] ~r :~ •• 'I'll. I . . . M.' , . - ...I r III M'II ): ,lib.......~"t-........~-"'---jlt-+:"""'-i-:t"2-+:~o4,-~2-+!""'+!~4~--~"""'~-!:-"""~{0 MY M.Y .• , I I~"" I I I GilBERT :GAU S~MATUYAMA:SRU NHES:MA TUYAMA:GAUSS: GllBE RT REV. !NORM; REV : NORM.: REV. ~ORM.: REV. Fig. 3. Center and upper magnetic profdes are obselVed, with east on the right, and reversed, with east on the left, respectively. The lower profde is the theoretical profde over the bodies shown. Normally magnetized bodies are black and reversely magnetized are white, and all are 2 km thick. With a spreading rate of 4.5 cm/yr the magnetized blocks correlate with the known history of the reversals to the Gilbert epoch. (from Heirtzler in Phinney: "The History of the Earth's Crust," 1968. By permission, Princeton University Press.) present-day motion along such offsets is indeed opposite in direction to the offset of the ridge segments has been a major confirmation by seismology of the plate tectonics synthesis. First-motion studies of earthquakes occurring along ridge-ridge transform faults confirm the symmetrical spreading of oceanic lithosphere from mid-ocean ridges. Further evidence of the mobility of the earth's crust comes from many additional sources such as studies of crustal heat flow and petrology. Plate boundaries and the nature of interaction at plate boundaries are especially well shown by seismological studies (Bara~angi and Dorman 1969). Continental crust, because of its relatively low density, is not so subject to subduction as is oceanic crust and does not return permanently below the lithosphere as oceanic plates do at the end of their conveyor-belt trips along the ocean floor. If it does descend deeply it tends to rise again later after remelting and differentiation, or lifts up continental crust above it to form exceptionally high mountain ranges, such as the Himalayas. Continental crust is therefore winnowed and tends to collect well away from long established spreading ridges. On a geological time scale the conveyor-belt action of the oceanic crust keeps its surface effectively swept clean of truly ancient sediments, which September 1972 explains the lack of rocks older than Mesozoic on the present deep ocean floor. The lithosphere of the present ocean floor, which covers about twothirds of the area of the earth, has been created in no more than the most recent 5 percent of the earth's history. . Until the 1960's many geologists, particularly American geologists, supported a static theory of stable continents, in which the continents had always been about where they are now. This was mainly because no mechanism for "contintneal drift" through oceanic crust could be envisaged. The current geological revolution has changed all this, putting in place of the static theory a far more general dynamic model which accounts for a much wider system of facts and interpretations that either were unknown or made no sense at all under the old scheme. It is now time to investigate how the plate tectonics concept affects biology. Epicontinental Flooding and Other Results of Spreading Maxima An increasing body of evidence suggests that at times of maximum spreading activity at any particular mid-ocean ridge more seawater is displaced by the rising ridge than at other times (Hallam 1963 and 1971, Johnson 1971). Such eustatic seawater flooding caused by activity in anyone area of the globe would tend to flood low sections of all continents simultaneously, leading to simultaneous stress in processes of adaptation, extinction, and occupation by organisms in far-flung areas. Similarly, cessation of spreading in one area would also have world-wide eustatic effects. How all this affects cyclothems and world climate is just now coming under review. Expansion and contractions of ecospace are linked to such episodes; thus, if spreading is to some extent episodic, then in some degree so are some of the major factors in evolution and extinction (Valentine and Moores 1970). If world-wide changes in plate tectonic activity are an index of changes in deep activity beneath the lithosphere (Nelson and Temple 1972), there may be an important linkage between oceanfloor spreading, vulcanism, and rates of polarity switching in the earth's mag- --=.~------------~j=-----=[- -F DEXTRAL TRANSFORM ~~ ~~ SINISTRAL BIOLOGICAL EFFECTS If we accept the principles and pro- cess of plate tectonics, then a very great deal follows, particularly with regard to the history of environments but also affecting other aspects of biology. Plate tectonics theory is not yet complete, but it is general enough to supplant SINISTRAL FAULTS TRANSCURRENT DEXTRAL FAULTS Fig. 4. Sense of displacements associated with transform faults and with transcurrent faults. Double line represents crest of mid-oceanic ridge; singte line, fracture zone. Terms "dextral" and "sinistral" denote sense of motion on active portions of faults. (from Sykes in Phinney: ''1he History of the Earth's Crust", 1968. By permission, Princeton University Press.) 521 Downloaded from http://bioscience.oxfordjournals.org/ at Penn State University (Paterno Lib) on September 18, 2016 '~J v~J~~~ previous ideas and to provide a firstorder model of the dynamics of the crust of the earth-the stage for all biological activity. netic field. Vulcanism in tum is linked to atmospheric dust levels, to albedo changes, and to availability of precipitation nuclei. Polarity reversals, passing through zero magnetic field as they must, cause Van Allen Belt collapse and changes in cosmic ray shielding, especially in the tropics. Quantitative assessments of these factors are still in a primitive state (Hays 1971). Separation Effects 522 relics of past continental separations (Griffiths 1971, Griffiths and Varne 1972). Collision Effects Most of us tend to think of "continental drift" as separation of continental masses, but in plate tectonic theory collision is as likely as separation. Past continental collisions at various times are now postulated by many geologists as the principal original processes operating in the creation of the Ural Mountains, the Appalachians, the Alpine/Himalayan chain, and several other important past and present mountain chains throughout the world. India smashed, albeit in very slow motion, into southern Asia during the Cenozoic; Australia seems to be colliding with southeast Asia at present. Many ancient collisions seems to be indicated by characteristic ophiolitic and blueschist suites of rocks that mark the sutures between former disparate continental masses. Two major environmental results of continental collision are obvious. First, marine conditions and maritime climate in the affected area would be profoundly altered and finally destroyed. Second, a range of mountains, probably a great one, would be created by marginal crumpling as enormous amounts of inertial energy were dissipated over some millions of years. A crude estimate of such energy under reasonable conditions might be in the neighborhood of 1024 ergs/year over a 2000 km front (Hales 1969). As much as 10 31 ergs might be dissipated in this case. The removal of the ocean and creation of such a range would have far-reaching climatic effects. Ecospace would be generally reduced, even though this would be partly offset by new ecospace created by mountain building. Environmental stress would be high and extinctions would be especially common because of the tremendous alteration of the environment. Mixture of terrestrial biotas of colliding land masses would occur, first by a few "sweepstakes dispersals" (Simpson 1940, 1953) as the coasts neared one another, then through a filter as they actually touched somewhere, and finally by a corridor modulated by the rising mountains which would act as a partial barrier. Ecological replacement of the less successful of similarly adapted organisms would take place on a large scale, and the biota, now a single BioScience VoL 22 No.9 Downloaded from http://bioscience.oxfordjournals.org/ at Penn State University (Paterno Lib) on September 18, 2016 Let us suppose that an originating ridge appeared or was carried beneath a continent, rafting away from each other the two continental halves. I have in mind the separation of Mrica from the Americas. The most obvious effect would be the immediate increase in length of the total continental coastline. As the continental fragments were pulled apart, the new coastlines would undergo profound environmental evolution as new continental shelves and slopes formed and as the climate changed. Expansion of total ecospace (Valentine 1969) would occur and new endemism would begin to develop, increasing total world species diversity (Valentine and Moores 1970). When the land masses first separated, a long, narrow, effectively intracontinental arm of the ocean would fill the developing gap. Evaporation would likely exceed precipitation and, where large rivers did not enter, salinity would be high until the gap widened farther. The Red Sea is a modem example. Salt deposition in the nascent split between eastern North America and the northwest coast of Africa during the Mesozoic seems to have been a similar case, the salt deposits now being widely sundered (Rona 1969, Pitman and Talwani 1972, Fig. 2). The moderating effects of large areas of water, as well as the changes in oceanic circulation, would have enormous influence on the climate as the land mass broke and its parts separated. As one of my colleagues has aptly remarked, there couldn't have been a Gulf Stream before there was an Atlantic Ocean! The eastward moving circumpolar Antarctic Current and its biological effects must also have evolved to the present situation since the breakup of the ancient supercontinent of Gondwanaland. Inasmuch as Australia has moved substantially northward, separating from Antarctica entirely within the Cenozoic, the evolution of the Antarctic Current is a Cenozoic event. Shifts in Arctic oceanic circulation as connections to other oceans were made or broken must have once had farreaching climatic effects now only dimly perceived. Other climatic changes would occur if a separating plate moved across climatic zonation, carrying a biota into a new environment. Ecospace might either increase or decrease if climatic belts were crossed. The northward movement of Australia is a good example because the motion has been toward the tropics. If Australia had not moved northward, its terrestrial biota would have shared to a significant extent the fate of the pre-glacial Antarctic terrestrial biota. Although they are probably of very minor import in paleobiogeography, guyots and atolls, carried along on the lava conveyor-belts of spreading plates, may shift geographic position without significant change relative to sea level, or they may be carried to great depths or, as has occurred in one known area, the Afar Triangle at the south end of the Red Sea, even be moved up onto land. Many of the older guyots and atolls of the Pacific are calculated to have moved with a northward component 25-30° or more since their creation (Francheteau et at, 1970). This, in conjunction with a relative, generally westward, Eulerian rotation of the lithosphere, is in harmony with various northwest-southeast lines of Pacific volcanic islands not associated with ridges' and trenches (Morgan 1972). Palebiological studies of sediment cores from guyots and atolls should be related to possible latitudinal components of plate motion. Some of the island biogeography of MacArthur and Wilson (I967) takes place on a moving, not a static, base. Although the time scale of such motion is long, its potential effects need to be considered. Separation of formerly contiguous areas would lead to disjunct distribution of organisms, though it is not the only way to achieve that phenomenon. At first taxonomic similarities would still extend to low levels, but with time and isolation, unless mitigated by dispersal across or around barriers, similarities would be confined to higher and higher taxonomic categories as endemism developed. The evolution in isolation of • the monotremes, Sphenodon, various ratite birds, and the southern distribution of various other archaic organisms affected by the fragmentation of Gondwanaland come to mind as Noah's Arks Paleogeography under the stable continent rationale was greatly influenced and shaped by two Americans, W. D. Matthew (1915, 1939) and G. G. Simpson (1940, 1943, 1946, 1947a, 1947b, 1952, 1953). Simpson, in particular, did much to develop the principles of the subjeCt and the quantify aspects of dispersal probability. Simpson developed the criteria for recognition of various types of dispersal routes, ranging in increasing order of probability from waif dispersal (or sweepstakes routes) through the filter bridge to the corridor (Simpson, 1940, 1953, Hallam, 1967). Under the stable continent rationale that was about are far as one could. If the ocean floor and continents or fragments -of continents move, however, then large' and small plates carrying continental crust and even oceanic islands rising above non-continental crust will make long fourneys if sufficient time is available, carrying samples of the biota of one area to disembark eventually at another. En route, evolution in isolation would occur, to be sure, and occasional sweepstakes dispersals might also be expected, depending in part on the distances to be traversed, but eventually the ark, to use an obvious term, would dock and the passengers get off to try to make a new life in a new land. If its land mass were large enough, say of nearly continental dimensions, such an ark might represent a delayed-action September 1972 one-way corridor. A balanced but consequences for the ark's biota before evolved sample of the biota would be docking occurred. Australia's rodents transferred from land mass A on a illustrate this point (Simpson, 1961 ; fragment of A, A I , to invade land mass Plane, 1967). After docking, one might B, but the inhabitants of B. could not expect a small ark's biota to be largely spread back all the way to A in the same swamped; the larger the ark, the more manner. Such a delayed one-way corri- likely survival would be. dor, which could also be viewed as a Motions Along Continental type of filter, would not be possible Strike-slip Faults under the basic premise of stable continents. Because of the distances to be traversed by large arks, known rates of Extensive lateral differential motions plate motion suggest that the time scale occurs in continental crust overlying of ark journeys would be of Epoch to areas of differentail motion in the lower Ear magnitude. lithosphere. The overlying continents The one-way corridor or filter prin- are fractured and formerly contiguous ciple is partly illustrated by the col- areas are carried away from each other. lision, now under way, of Australia The famous right-laterial San Andreas (once part of Gondwanaland) with Fault system of the North American southeast Asia (Cox 1970). Also illu- West Coast is an excellent example strated by the Australian collision is the (Dietz and Holden 1970, Dickinson et fact that a new corridor between two al. 1972). In this way fossil assemblages colliding land masses does not develop may be moved significant distances instantly but first passes through stages away from the locale where they were of waif dispersal and filter action. To once alive (Addicott 1967). Addicott's some Ifxtent, also, the more sedentary interpretation suggests a motion of travelers on an ark, A I , would be met between 190 and 320 km. iri California on arrival not only by the indigenous since the early Miocene. According to inhabitants of B but also by somewhat the interpretation presented by Dickinmodified descendants of A l passengers son et a1. (1972) about 300 km of who managed to jump ship while the movement has taken place there since ark was still nearing port. Likewise, the early Miocene and perhaps twice invasion of the ark by inhabitants of the that amount since the beginning of the port would begin even before docking Cenozoic. A similar right-lateral fault and conceivably would have far-reaching system in Alaska (Page and Lahr 1971) Fig. S. Calculated rotational pole positions for the earth in the early Tertiary, uncorrected for relative plate motions (Based on Creer, 1970, and other sources). All of the calculated rotational pole positions indicate that, relative to the places yielding the magnetic data utilized, the rotational pole was nearer Beringia than now. If the earth's rotational axis has not shifted appreciably, then the plates themselves have shifted positions, both individually and as part of a more general shift of the lithosphere relative to the deeper earth. 523 Downloaded from http://bioscience.oxfordjournals.org/ at Penn State University (Paterno Lib) on September 18, 2016 biota, would undergo major reorganization. Thus world species diversity woul,d be reduced by the union of a major pieces of continental crust. An important effect of continental collision, especially if separation occurs later along the same general trend of the suture, is that fossil-bearing rocks (and also rocks from which pole positions may have been calculated!) of wholly different geographic origins might be juxtaposed, as seems to have been the case along the east coast of North America (Wilson 1968). Fragments of continents get left behind, so to speak, attached like fender paint to the other party in the collision. The older the rocks, the greater the chance would be that the continent to which they now belong is composite. Unscrambling such "unnatural" juxtaposition with the aid of plate tectonic theory may be expected to' have profound effects on paleogeographic work. may have moved certain warm temperate early Cenozoic floral localities farther north than their original positions. Total lateral motion along the Alpine Fault in New Zealand is interpreted to be as much as 1200 km (Griffiths and Varne 1972). Although the distances may not be very great, still they must be taken into account in biogeographic reconstructions. If such motions continue on a time scale measured in hundreds rather than tens of millions of years, then distances on the order of thousands of kilometers are involved. "Polar wander paths" for the different plates have done much to confirm differential plate motion, but they also confirm differential motion of the lithosphere as a whole (or at least of groups of plates) with respect to the earth's rotational axis (Irving 1964, Creer 1970). Irving (I 964), Irving and Robertson (I968), Sloan (I 969), Stanley (I 970), Szalay and McKenna (1971), and others have attempted on this basis to reconstruct paleolatitudes for late Cretaceous and early Cenozoic time, and a number of authors have done the same for earlier periods in earth history. The data for early Cenozoic rotational pole positions are still rather scanty (Fig. 5) but there is little question that even during that time the eastern end of Siberia and the Bering land bridge were nearer the "north" rotational pole than they are now. Paleolatitude belts concentric around such a shifted rotational pole position relative to the lithosphere make sweeping curves on maps of present latitudes and vice versa, yet many biogeographic reconstructions are still made on the basis of the present polar coordinate system (e.g., various maps in Middlemiss et al. 1971). Both aspects of the plate tectonics systhesis should be reflected in paleogeographic base maps for biogeographic reconstructions: relative plate motion should be depicted on a coordinate system compatible with what is known of the rotational pole position for the time depicted in each reconstruction. Even as late as the beginning of the Cenozoic, latitudes were significantly tilted with respect to their present orientation and, to the extent that latitudes control climatic belts, the climatic belts would have tended to be similarly tilted with respect to the present coordinate system. 524 This short survey suggests that the next few years will be exciting ones for those who wish to synthesize a new model of biological interaction with the physical conditions of the earth throughout its long history. Paleogeography will be profoundly altered and given a primarily geophysical, rather than biological, base. Biogeography will, in general, have to adjust to and test the validity of that base, and indeed there will be significant biological constraints on some geophysical proposals. Biology will contribute information about the timing of certain events and the degree of filtering in dispersal, but will be of secondary importance in determining past configurations of the earth's lands and oceans. The physical sciences will prQvide a steadily increasing power of resolution concerning the history of environmental complexity and physical environmental stimuli affecting biological evolution. REFERENCES Addicott, W. O. 1967. Zoogeographic evidence for late Tertiary laterial slip on the San Adnreas Fault, California. U.S. Geol. Surv. Prof, Paper 593-D: D1-D12. Atwater, T. 1972. Test of New Global Tectonics: discussion. Bull. A mer. Assoc. PetroL Geol., 56: 385-388. Barazangi, M., and J. Donnan. 1969. World seismicity maps compiled from ESSA, Coast and Geodetic Survey, epicenter data, 1961-1967. Bull. Seismol. Soc. A mer. , 59: 369-380. Bostrom, R. C. 1971. Westward displacement of the lithosphere. Nature, 234: 536-538. Bullard, E., J. E; Everett, and A. G. Smith. 1965. The fit of the continents around the Atlantic. Phil. Trans. Roy. Soc. London, 258: 41-51. Cox, C. B. 1970. Migrating marsupials and drifting continents. Nature, 226: 767-770. Creer, K. M. 1970. A review of palaeomagnetism. Earth-Sci. Rev., 6: 369466. Decker, R. W., P. Einarsson, and P. A. Mohr. 1971. Rifting in Iceland: new geodetic data. Science, 173: 530-533. Dickinson, W. R., D. S. Cowan, and R. A. Schweichert. 1972. Test of New Global Tectonics: discussion. Bull. A mer. Assoc. Petrol. GeoL, 56: 375-384. Dietz, R. S., and J. C. Holden. 1970. The breakup of Pangaea. Sci. Amer., 223(4): 30-41. Francheteau, J., C. G. A.Harrison, J. G. Sclater, and M. L. Richards. 1970. Magnetization of Pacific seamounts: a preliminary polar curve for the northeastern Pacific. J. Geophys. Res., 75: 2035-2061. Griffiths, J. R. 1971. Reconstruction of the south-west Pacific margin of Gondwanaland. Nature, 234: 203-207. Griffiths, J. R., and R. Vame. 1972. Evolution of the Tasman Sea, Macquarie Ridge, and Alpine Fault. Nature (Phys. Sci.), 235: 83-86. Hales, A. L. 1969. Gravitational sliding and continental drift. Earth Planet. Sci. Lett., 6: 31-34. Hallam, A. 1963. Major epeirogenic and eustatic changes since the Cretaceous, and their possible relationship to crustal structure.Amer. J. Sci., 261:397423. _ 1967. The bearing of certain palaeozoogeographic data on continental drift. Palaeogeogr. PalaeoclimatoL Palaeoecol., 3: 210-241. _ 1971. Mesozoic geology and the opening of the North Atlantic. J. Geol., 79(2): 129-157. Hammond, A. L. 1970. Laser ranging: measuring the moon's distance. Science, 170: 1289-1290. Hays, J. D. 1971. Faunal extinctions and reversals of the earth's magnetic field. GeoL Soc. Amer. BulL, 82: 2433-2447. Heirtzler, J. R. 1968. Evidence for ocean floor spreading across the ocean basisn. In: The history of the earth's crust, R. A. Phinney (ed.). Princeton University Press, Princeton, p. 90-100. Irving, E. 1964. Paleomagnetism and its application to geological and geophysical problems. John Wiley and Sons, New York. Irving, E., and W. A. Robertson. 1968. The distribution of continental crust and its relation to ice ages. In: The history of the earth's crust, R. A. Phinney (ed,). Princeton University Press, Princeton, p. 168-177. Johnson, J. G. 1971. Timing and coordination of orogenic, epeirogenic, and eustatic events. Geol. Soc. A mer. BulL, 82: 3263-3298. MacArthur, R. H., and E. O. Wilson. 1967. The theory of island biogeography. Princeont Univ. Press, Princeton. Matthew, W. D. 1915. Climate and evolution. Ann. New YorkAcad. Sci.; 24: 171-318. _ 1939.Climate and evolution. Second edition, revised and enlarged, Spec. Publ. New York Acad. Sci., 1: 1-223. McKenzie, D. P;, and W. J. Morgan. 1969. Evolution of triple junctions. Nature, 224:' 125-133. McKenzie, D. P., and R. L. Parker. 1967. The North Pacific: an example of tectonics on a sphere. Nature, 216: 1276-1280. Menard, H. W., and T. Atwater. 1969. Origin of fracture zone topography. Nature, 222: 1037-1040. Merservey, R. 1971. The coastline fit of Africa and South America. Palaeogeogr. PalaeoclimatoL PalaeoecoL, 9: 233-243. Meryerhoff, A. A., and H. A. Meyerhoff. 1972. "The New Global Tectonics": major inconsistencies. BulL A mer. Assoc. Petrol. Geol., 56: 269-336. Middlemiss, F. A., P. F. Rawson, and G. Newell (eds.). 1971. Faunal provinces in space and time. Seel House Press, LiverpooL Morgan, W. J. 1972. Deep mantle convection plumes and plate motions. BulL A mer. Assoc. Petrol. Geol., 56: 203-213. Nelson, T. H., and P. G. Temple. 1972. Mainstream mantle convection: a geologic analysis of plate motion. Bull. Amer. Assoc. Petrol. Geol., 56: 226-246. Page, R., and J. Lahr. 1971. Measurements of fault slip on the Denali, Fairweather, and Castle Mountain faults, Alaska. J. Geophys. Res., 76: 8534-8543. BioScience Vol. 22 No.9 Downloaded from http://bioscience.oxfordjournals.org/ at Penn State University (Paterno Lib) on September 18, 2016 Tilted Latitudes CONCLUDING REMARKS Palmer, H. 1968. East Pacific Rise and westward drift of North America. Nature, 220: 341-345. Pitman, W., and M. Talwani. 1972. Sea-floor spreading in the North Atlantic. Ceol. Soc. Amer. BUll., 83: 619-646. Plane, M. 1967. Stratigraphy and vertebrate fauna of the Otibanda Fonnation, New Guinea. Bur. Min. Res. Ceol. Ceophys. (Aust.) , Bull. 86: VI, 1-64. Rona, P. A. 1969. Possible salt domes in the deep A tlantic off North-west Africa. Nature, 224 : 141-143. Simpson, G. G. 1940. Mammals and land bridges. J. Wash. Acad. Sci., 30(4): 137-163. 1943. Mannals and the nature of continents. A mer. J. Sci., 241: 1-31. __ . 1946. Tertiary land bridges. Trans. New York A cad. Sci. , 8(8): 255-258. 1947a. Holarctic mammalian faunas and continental relationships during the Cenozoic. Ceol. Soc. A mer. Bull., 58: 613-688. 1947b. Evolution, interchange, and resemblance of hte North American and Eurasian Cenozoic mammalian faunas. Evolution, 1(3): 218-220. 1952. Probabilities of dispersal in geologic time. In: The problem of land connections across the South Atlantic, with special reference to the Mesozoic, E. Mayr (ed.). Bull. A mer. Mus. Nat. Hist., 99 : 163-176. 1953. Evolution and geography. Condon Lectures, Oregon State Syst. Higher Educ., Eugene. Sloan, R. E. 1969. Cretaceous and Paleocene terrestrial communities of western North America. Proc. North A mer. Paleont. Conv., September 1969, pt. E, p. 427-453. Stanley, E. A. 1970. The stratigraphical, b i 0 ge ographical, paleoau tecological and evolutionary significance of the fossil pollen group Triprojectacites. Bull. Ceorgia Acad.Sci., 28 : 1-44. Sykes, L. R. 1968. Seismological evidence for transfonn faults, sea floor spreading, and continental drift. In: The History of the earth 's crust, R. A. Phinney (ed.). Princeton University Press, Princeton, p. 120-150. Szalay, F. S., and M. C. McKenna. 1971. Beginning of the Age of Mammals in Asia: the late Paleocene Gashato Fauna, Mongolia. Bull. A mer. Mus. Nat. Hist. , 144 : r I h ElIf¥ POLAROIC • _ _I _ _I _ _I 269~318 ; Valentine, J. W. 1969. Patterns of taxonomic and ecological structure of the shelf benthos during phanerozoic time. Paleontology, 12: 684-709. Valentine, J. W., and E. M. Moores. 1970. Plate-tectonic regulation of faunal diversity and sea level: a model. Nature, 228: 657-659. Vine, F. J. 1968. Magnetic anomalies associate with mid-ocean ridges. In: The history of the earth's crust, R. A. Phinney (ed.). Princeton, p. 73-89. Vine, F. J, and D. H. Matthews. 1963. Magnetic anomalies over oceanic ridges. Nature, 199: 949-949. Wilson, J. T. 1965. A new class of faults and their bearing on continental drift. Nature, 207: 343-347. _ _ 1968, Static or mobile earth: the current scientific revolution. Proc. A mer. Phil. Soc., 112(5): 309-320. September 1972 Shoot round circles and square squares. Some extraordinary optics in the new Bausch & Lomb Integrated Camera Series II produce images freer of distortion than eVer before. Shooting a properly aligned specimen through a StereoZoom 7 Microscope, Series II Camera and Camera Adapter, you get a picture that retains the identical shape. Circles are never out of round. Squares are square-sides don't cave in or bulge out. Compare results with any other system and convince yourself. There's more, too, like a large viewing screen that permits comfortable group viewing in ambient light. A highly sensitive exposure meter simplifies picture taking by eliminating guesswork. Your choice of three interchangeable camera bodies-the 35mm, takes the full range of 35mm films-BW, color prints and transparencies; 3~" x4~" Polaroid* and 4" x 5" which accepts plates, film packs, sheets, Polaroid sheets. Ask for a see-for-yourself, no-obligation demonstration, or write for our fact-filled Catalog 42-2396. BAUSCH & LOMB ~ StereoZoom, Reg. T.M. Bausch & Lomb 'Trademark, Polaroid Corp . SCIENTIFIC INSTRUMENT DIVISION 21609 Bausch Street, Rochester, N.Y. 14602 PLEASE CIRCLE NO.8 ON THE READER'S SERVICE CARD 525 Downloaded from http://bioscience.oxfordjournals.org/ at Penn State University (Paterno Lib) on September 18, 2016 _ _I