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U.S. copyright law (title 17 of U.S. code) governs the reproduction and redistribution of copyrighted material Branch Main Library Lending 2 Rapid #: -867943 (MM CALL #: Page 2 of Ariel IP: 165.230.139.63 QH359 .E94 LOCATION: IQU :: Main Library 1 ‘TYPE: Article CC:CCL q JOURNAL TITLE: Evolutionary theory USER JOURNAL TITLE: Evolutionary Theory IQU CATALOG TITLE: Evolutionary theory ARTICLE TITLE: A new evolutionary law" | ARTICLE AUTHOR: Lee VanvValen q VOLUME: 1 | IssU | YEAR: 1973 PAGES: 1-30 ISSN: 0093-4755 oct #: CROSS REFERENCE ID: 147110 VERIFIED: | BORROWER: Main Library PATRON: Peter PATRON ID: PATRON ADDRESS: PATRON PHONE: PATRON FAX: PATRON E-MAIL: PATRON DEPT: PATRON STATUS: PATRON NOTES: pimorin@rci.rutgers.edu Graduate NOTICE: This material may be protected by copyright law {Title 17 U.S. Code System Date/Time: 9/6/2006 10:54:30 AM MS hitps:/rapid2. library colostate.edu/ll/ViewQueue.aspx?ViewType=PendingByBranch&ld=48 9/6/20 Ss . ALW4O 193949 ‘A NEW EVOLUTIONARY LAW Leigh Van Valen Department of Biology ‘The University of Chicago Chicago, Illinois 60637 ABSTRACT: All groups for which data exist go extinct at a rate that is constant for @ given group. When this is recast in ecological form (the effective environment of any homogeneous eToup of organisms deteriorates at a stochasti- cally constant rete), no definite exceptions exist although a few are possible. Extinction rates are similer within some very broad categories and vary regularly with size of erea inhabited. A new unit of rates for discrete phenomena, the macarthur, is introduced, Lews are appropriate in evolutionary diology. Truth needs more than correct predictions. The Law of Extinction is evidence for ecological significance and comparability of taxa. A ncn- Markovian hypothesis to explain the lew invokes mutually incompatible optima within an adaptive zone. A self-perpetuating fluctuation results which can de stated in terms of an unstudied aspect of zero-sum game theory. The hypothesis can be derived fron a view that monentary fitness is the amount of control of resources, which remain constent in total amount. The hypothesis implies that long-term fitness has only two components and that events of mutualism ere rare. The hypothesis largely explains the observed pattern of molecular evolution. * * * Introduction During e study (Van Valen, submitted) on the effects of extinction I wanted to show that a model I was using was oversimplified. It assumed no correlation of probability of extinction with age of the group, and I thought that generally more vulnerable groups should die out first. A’test using date from Simpson (1953) showed to my astonishment that the assumption was reasonably correct in these cases. I did not believe it could be generally true and so tested these and other cases in more deteil. ‘The assumption proved to be consistent with all available data. Others (unpublished results) have now confirmed this finding for individual taxa. I will present @ more extended treatment elsewhere; the present paper is condensed. ‘The Evidence The method is an application of the survivorship curve of population ecology (including demography). It is a simple plot of the proportion of the original sample that survive for various intervals. In this case the sample is the set of all known subgroups of some larger group, no matter when in absolute time each subgroup originated. A logarithmic ordinate, standard in ecology, gives the property that the slope of the curve at any age is Proportional to the probability of extinction at that age. Simpson (19hh, 1953) compiled two well-known taxonomic survivorship curves but used an erithmetic ordinate (1-4). ‘The results (Figs. 1-5) for over 25,000 subtaxe show almost uniform Linearity for extinct taxe except for effects attributeble to sampling error (5,6). Sampling error is most noticeable at the bottom of the graphs, where Evol. Theory 1:1-30 (July 1973) AR 4B 7AK LINM CENT! IRS a UN OSPR IOI 67 1000-y@—$ DINOFLAGELLATE DINOFLAGELLATE CYSTS CYSTS 100. (EXTINCT GENERA) . (EXTINCT SPECIES) wo *s . 104 oF ‘ al 7 ° 80 wo My. 180 4 | Sas Py 8 S0__ My 100 woe, terme cenrnases ‘ Paawaces “oe rare, Boo 1000-4 - s EXTINCT 1.0" *LIVING woz 7 . ° so” My 100 . 100g, COCCOLITHOPHYCEAE . x (EXTINCT GENERA) ey . x . * 104 1 é 2 40 mv. 60 a CENTRALES ro 1000-4 . (SPECIES) ° 50 100 MY. 150 ' of FORAMINIFERA ao x (FAMILIES) . . Tora exrincr « 100-4 5 x a EXCEPT LATE PERMIAN mk, aS, ae 0-4 . a an ca " sextet 200°" 400° My. 600 | 1000-9 FORAMINIFERA o "_ __40_x._ 60 (GENERA) J CENOZOIC PLANKTONIC is EXTINCT PLANKTONIC x FORAMINIFERA So TINT MTN (EXTINCT SPECIES) 10-4 ] 1 : x $ % 2 My 30 Fig. 1. Taxonomic survivorship curves for protists. fa BRACHIOPODA (EXTINCT FAMILIES) ALL (EXCLUDING, cate "PERMIAN JEXTINCTIONS oes = oo) ee ° Mix. "300 PELECYPODA (GENERA) 1000 100 10 0 ‘© OTHER ARTICULATA BRACHIOPODA (EXTINCT GENERA) offATE PERMAN IEXTINCTIONS x OTHER INARTICULATA 100 MY ‘SEXTINCT LIVING ~~ (GENERA) RUDISTS. 100: (HIPPURITACEA) PELECYPODA (FAMILIES) BRYOZOA (FAMILIES) ‘EXTINCT = LIVING 20 My. 400 ia NAUTILOIDEA EXTINCT GENERA EXTINCT FAMILIES Wa ad bax, “mm, oo ALL ome * . aaa a ° 20 40 MY, 60 100 200 MY. 300 Goo. AMMONOIDEA ARCHAEOGASTROPODA + MONOPLACOPHORA eau ever “s

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