An adaptationist programme has dominated evolutionary thought in England and the United States during the past 40 years. It is based on faith in the power of natural selection as an optimizing agent. It proceeds by breaking an organism into unitary ‘traits’ and proposing an adaptive story for each considered separately. Trade-offs among competing selective demands exert the only brake upon perfection; non-optimality is thereby rendered as a result of adaptation as well. We criticize this approach and attempt to reassert a competing notion (long popular in continental Europe) that organisms must be analysed as integrated wholes, with Baupläne so constrained by phyletic heritage, pathways of development and general architecture that the constraints themselves become more interesting and more important in delimiting pathways of change than the selective force that may mediate change when it occurs. We fault the adaptationist programme for its failure to distinguish current utility from reasons for origin (male tyrannosaurs may have used their diminutive front legs to titillate female partners, but this will not explain why they got so small); for its unwillingness to consider alternatives to adaptive stories; for its reliance upon plausibility alone as a criterion for accepting speculative tales; and for its failure to consider adequately such competing themes as random fixation of alleles, production of non-adaptive structures by developmental correlation with selected features (allometry, pleiotropy, material compensation, mechanically forced correlation), the separability of adaptation and selection, multiple adaptive peaks, and current utility as an epiphenomenon of non-adaptive structures. We support Darwin’s own pluralistic approach to identifying the agents of evolutionary change.

The spandrels of San Marco and the Panglossian paradigm: a critique of the adaptationist programme

An adaptationist programme has dominated evolutionary thought in England and the United States during the past 40 years. It is based on faith in the power of natural selection as an optimizing agent. It proceeds by breaking an organism into unitary 'traits' and proposing an adaptive story for each considered separately. Trade-offs among competing selective demands exert the onlv brake upon perfection; non-optimality is thereby rendered as a result of adaptation as well. We criticize this approach and attempt to reassert a competing notion (long popular in continental Europe) that organisms must be analysed as integrated wholes, with Bauplane so constrained by phyletic heritage, pathways of development and general architecture that the constraints themselves become more interesting and more important in delimiting pathways of change than the selective force that may mediate change when it occurs. We fault the adaptationist programme for its failure to distinguish current utility from reasons for origin (male tyrannosaurs may have used their diminutive front legs to titillate female partners, but this will not explain why they got so small); for its unwillingness to consider alternatives to adaptive stories; for its reliance upon plausibility alone as a criterion for accepting speculative tales; and for its failure to consider adequately such competing themes as random fixation of alleles, production of nonadaptive structures by developmental correlation with selected features (allometry, pleiotropy, material compensation, mechanically forced correlation), the separability of adaptation and selection, multiple adaptive peaks, and current utility as an epiphenomenon of non-adaptive structures. We support Darwin's own pluralistic approach to identifying the agents of evolutionary change.

/pdf/the-spandrels-of-san-marco-and-the-panglossian-paradigm-a-19fdlsieru.pdf

Rising atmospheric carbon dioxide (CO2), primarily from human fossil fuel combustion, reduces ocean pH and causes wholesale shifts in seawater carbonate chemistry. The process of ocean acidification is well documented in field data, and the rate will accelerate over this century unless future CO2 emissions are curbed dramatically. Acidification alters seawater chemical speciation and biogeochemical cycles of many elements and compounds. One well-known effect is the lowering of calcium carbonate saturation states, which impacts shell-forming marine organisms from plankton to benthic molluscs, echinoderms, and corals. Many calcifying species exhibit reduced calcification and growth rates in laboratory experiments under high-CO2 conditions. Ocean acidification also causes an increase in carbon fixation rates in some photosynthetic organisms (both calcifying and noncalcifying). The potential for marine organisms to adapt to increasing CO2 and broader implications for ocean ecosystems are not well known; both are high priorities for future research. Although ocean pH has varied in the geological past, paleo-events may be only imperfect analogs to current conditions.

/pdf/ocean-acidification-the-other-co-2-problem-4h18mw2cpj.pdf

Ocean Acidification: The Other CO 2 Problem

We review the biogeography of microorganisms in light of the biogeography of macroorganisms A large body of research supports the idea that free-living microbial taxa exhibit biogeographic patterns Current evidence confirms that, as proposed by the Baas-Becking hypothesis, 'the environment selects' and is, in part, responsible for spatial variation in microbial diversity However, recent studies also dispute the idea that 'everything is everywhere' We also consider how the processes that generate and maintain biogeographic patterns in macroorganisms could operate in the microbial world

/pdf/microbial-biogeography-putting-microorganisms-on-the-map-1k7e5do5pn.pdf

Microbial biogeography : putting microorganisms on the map

Darwin (1859, pp. 11-14, 143-150) stressed the evolutionary importance of covariation between characters in populations and its "most imperfectly understood" connection with correlated responses to artificial and natural selection. After the turn of the century the discoveries of pervasive pleiotropy and linkage of Mendelian factors revealed the underlying genetic mechanisms. Existing theory on the dynamics of correlated characters has been developed in the limited framework of practical plant and animal breeding. Methods of multivariate analysis, functional analysis and optimality criteria popular among evolutionists, do not account for dynamical constraints imposed by the pattern of genetic variation within populations. Consideration of phenotypic variation often does not suggest any clear mechanism connecting growth patterns or adult variation to interspecific evolution. When there is individual variation in development, no necessary correspondence exists between ontogenetic and adult variation in a population (Cock, 1966, pp. 148-15 1). It is also common for the pattern of adult variation within a species to differ from that at higher taxonomic levels (Simpson, 1953, pp. 25-29). An example which will be investigated here is the brain weight:body weight relationship. At various taxonomic levels, brain and body weights tend to follow the allometric equation

https://onlinelibrary.wiley.com/doi/pdf/10.1111/j.1558-5646.1979.tb04694.x

Quantitative genetic analysis of multivariate evolution, applied to brain:body size allometry.

We have calculated rates of evolution for 19 lineages of Neogene bivalves over time intervals ranging from 1 ma to 17 ma. Our morphometric comparisons are based on 24 variables, for which we have made more than 43,000 individual measurements normalized for shell size. We initially assessed evolutionary changes in shape for 19 early Pliocene (4 ma old) species of bivalves by comparing these forms to their closest living relatives, which in 12 cases have traditonally been assigned to the same species. To make our study unbiased and comprehensive, we included all species that met certain predetermined guidelines and that offered suitable fossil material for measurement. We compared early Pliocene and Recent populations using (1) all 24 variables treated separately, (2) multivariate distance (Mahalanobis' D), based on the full set of variables, and (3) eigenshapes for shell outlines. For these comparisons, we used as a yardstick the same measures of morphologic distance applied to pairs of geographically separated Recent populations that belong to eight of the living species to which the fossil populations were compared. As it turns out, with minor exceptions, the distribution of morphologic distances between 4 ma old and Recent populations resembled the distribution of distances between conspecific Recent populations.We calculated net rates of evolution separating pairs of populations that belong to single lineages. For all intervals of time, the distribution of differences between population means for individual variables is remarkably similar to a comparable distribution representing the comparison of pairs of conspecific Recent populations from separate geographic regions. Because morphologic differences between populations do not vary greatly with evolutionary time, measured “rates” of evolution, on the average, decrease with interval of measurement. Because these differences resemble intraspecific variability, however, the rates do not represent significant evolution. Evolution has followed a weak zigzag course, yielding only trivial net trends.The weak and reversible “trends” that we measured yield net rates averaging less than 10 millidarwins, which is much lower than most rates previously reported for marine invertebrates (average ~200 millidarwins for a 1 ma interval and ~60 millidarwins for a 10 ma interval). We attribute this disparity (1) to the fact that most previously published rates have been calculated when a significant amount of evolution was recognized in advance (often for a poorly documented lineage) and (2) to the fact that most measured variables have represented nothing more than some measure of body size. We conclude that shape, as opposed to size, has been highly stable in bivalve evolution over millions of years and 106–107 generations. We conclude that to characterize rates or evolution for any group of organisms, one must employ a large, unbiased sample of measurements for numerous well-documented lineages, and one must segregate data depicting shape from data depicting size.

Approximate evolutionary stasis for bivalve morphology over millions of years; a multivariate, multilineage study

Rates of origination and extinction for marine animal genera dropped to low levels in latest Mississippian time, immediately after massive glaciers formed in the Southern Hemisphere and a second-order mass extinction occurred. Evolutionary turnover and diversity remained low for ∼50 m.y., shifting markedly upward precisely when extensive glaciation ended in Early Permian time. All diverse taxa with good fossil records experienced low rates of origination and extinction during this major ice age. Such sluggish rates would be predicted for faunas of shallow seas on or adjacent to a heavily glaciated supercontinent such as Pangea, where cool and highly seasonal thermal regimes should dictate that species have broad ecological niches, widespread geographic distributions, and large and relatively stable populations.

Depressed rates of origination and extinction during the late Paleozoic ice age: A new state for the global marine ecosystem

A variety ofpatterns suggest that climatic cooling was the primary cause of the extinction of organic reef communities late in the Ordovician, Devonian, and Permian periods. Other potential agents exhibit serious deficiencies. Anoxic conditions, for example, cannot spread to surface waters. Eustatic sea-level lowering occurred during only two of the extinction intervals, and at other times during the Phanerozoic, lowering of sea level failed to produce mass extinction. Furthermore, sea-level drops precipitate an increase in the areas around volcanic islands available for reef growth. No iridium anomaly that might have resulted from a bolide impact has been found for any of the three Paleozoic crisis intervals, and each of these intervals spanned several million years. Patterns consistent with the idea that climatic cooling caused heavy extinction of Paleozoic reef communities are the following: 1) Non-tropical marine taxa suffered less severe extinction than tropical groups, presumably because many of their species were able to migrate equatorward. 2) During the protracted Ordovician and Permian crises, biogeographic provinces contracted toward the equator. 3) For each crisis, a disproportionately large percentage of surviving taxa were cosmopolitan in distribution. 4) In the aftermath of each event, carbonate production was reduced. 5) Reef-building was suppressed for several million years after each crisis, even though potential reef-builders survived. 6) Each mass extinction approximately coincided with the initiation of a glacial interval in a polar region.

Climatic cooling and mass extinction of Paleozoic reef communities

Trends, rates, and patterns of evolution in the Bivalvia

A revision of the best selling historical geology text, completely updated to provide a thorough but lucid introduction to the principles of historical geology. Stanley's assertion that 'the physical history and the biological history of the earth are inextricably interwined shapes his text throughout. The material is organized clearly and logically. The two-part progression resents basic principles first, then proceeds with a chronological study of the history of life. For everv interval, global events (such as general patterns of geographic change and major occurrences in the evolution and extinction of life) are reviewed within the context of plate tectonics. This review is followed bv a description of important regional events and their place within this global framework. Stanlev offers insightful observations on how environmental conditions have affected life through the ages. Numerous attractive illustrations enhance textual material. Chapter-ending reviews and exercises reinforce students' understanding and there are additional reading lists, a thorough glossary and useful appendices. Stanley has updated and improved his text to make i-t even more accessible to students. Notable changes include: Streamlined and reorganized chapter on mountain building, with new discussions of ancient plate movements Condensed and updated coverage of the Precambrian Updated and expanded discussions of mass extinctions throughout the text Additional material on isotopes and dating techniques Updated discussion of human evolution.

Steven M. Stanley

Papers

Approximate evolutionary stasis for bivalve morphology over millions of years; a multivariate, multilineage study

Depressed rates of origination and extinction during the late Paleozoic ice age: A new state for the global marine ecosystem

Climatic cooling and mass extinction of Paleozoic reef communities

Trends, rates, and patterns of evolution in the Bivalvia

Earth and life through time