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Paleontology in the 21St Century David Jablonskia,1 and Neil H

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Paleontology in the 21St Century David Jablonskia,1 and Neil H SPECIAL FEATURE: INTRODUCTION The future of the fossil record: Paleontology in the 21st century David Jablonskia,1 and Neil H. Shubinb,1 ing the split between sharks and bony fishes, aDepartment of Geophysical Sciences and bDepartment of Organismal Biology and but the fossil record shows that bone was Anatomy, University of Chicago, Chicago, IL 60637 present well below that evolutionary node, and that modern sharks represent a derived statefortheclade,havinglostbonystructures In the past two decades, great progress in the fungi, and protists (5–14). The contributions widely distributed in their ancestors (30, 31). biological sciences has come from the in- of paleontology to the study of the rates and Paleontological data are invaluable for in- corporation of history in understanding basic pattern of phenotypic evolution are legion. ferring ancestral character states and the as- mechanisms in fields ranging from ecology At the species level, the fossil record has sembly of character complexes, and can now and conservation biology to physiology and famously shown that the evolutionary re- be used to test hypotheses drawn from de- developmental biology. This conceptual ex- sponsiveness of local populations on decadal velopmental or phylogenetic analyses. pansion has been promoted in large part by or centennial timescales usually translates at The discovery and analysis of fossils from theoretical, methodological, and empirical the 1- to 10-million-y timescale into stasis or keyintervalsinthehistoryoflifecaninform advances in two seemingly disparate fields. nondirectional random walks rather than the sequence, pattern, and phylogenetic The first field is molecular biology, which sustained, directional evolutionary trans- dynamics underlying the origin of major opens powerful new windows on phyloge- formation (15, 16). For higher taxa, the functional and anatomical novelties. Such netic relationships, genome structure and quantification of form within a multidi- “intermediate forms” in the fossil record can function, and developmental mechanisms. mensional morphospace was developed in serve as tests of genetic, developmental, and The second field is paleontology, which paleontology (17) and this rich literature affords a unique, direct, and expanding biomechanical hypotheses based on extant continues to find new ways to explore taxa. Indeed, some of the most fundamental source of information into the anatomies, evolutionary diversification, from formal ecologies, physiologies, and spatial and discoveries of fossil stem taxa have only visualizations of evolutionary convergence happened in recent decades. Among verte- temporal dynamics of past life. The fossil and parallelism (e.g., ref. 18) to direct record is certainly rich in incident and rife brates alone, fossils have illuminated evolu- analyses of the relation between ontogeny tionary pathways leading to the origin of with bizarre players, but an extensive body and phylogeny (19–21). of research now treats the fossil record as a vertebrates (32), tetrapods (33), turtles (34), The biological world we see around us snakes (35), mammals (36, 37), birds (38), biological laboratory for rigorously framing today is a highly pruned version of a rich and and testing hypotheses at the intersection of horses (23), whales (39, 40), hominids (24), ancient tree of life. Consequently, reliance and many other groups. These fossils are paleontology with diverse disciplines across solely on recent taxa in analyses of origins i the full range of timescales encompassed valuable on several counts: ( ) they provide can definitively mislead. The diversity and data on the rate and pattern of character by the earth and life sciences. This Special disparity of many extant clades, from ele- Feature collects some of the exciting and acquisition, which in turn illuminates the phants (22) to horses (23), and for that factors underlying dramatic transformations important new directions and insights, matter hominins (24), has demonstrably from the beginnings of life on Earth to (e.g., feathers as insulation and display be- declined in the latest Cenozoic. Those clades ii the immediate precursor to the present-day fore flight in birds); ( ) they inform mech- were hardly unique, so that fossil data are biota (Fig. 1). anistic hypotheses on underlying changes essential for a fuller understanding of the Paleontology informs the natural sciences in development, both constraining and in- rates and patterns of phenotypic change by providing unique sources of data on spiring experimental tests of rival scenarios within and among many clades (e.g., refs. important phenotypes and on the spatial and (e.g., developmental hypotheses on the ori- 25 and 26). Of course, some clades are in- temporal dynamics of biological events and gin of the wrist and ankle joints of tetra- accessible or sparsely represented as fossils, iii processes. Accordingly, we organize these pods); and ( ) they place transitions in their but even in such groups, from onychoph- contributions in terms of the important environmental context (e.g., whales on the orans to ants to penguins, fossils alter our phenotypes contained in the fossil record, shoreline of a tropical sea (39) and hominids and the analysis of dynamics of species, picture of the timing and extent of mor- arising with bipedal gaits in woodland set- clades, and communities in both space phological diversification. Formal incorpo- tings (41). Moreover, many major transi- and time. ration of sparse fossil morphologies in tions occurred in ecosystems lacking close phylogeny-based analyses of extant species modern analogs, and so can be understood Phenotypes is a promising interdisciplinary growth area ecologically only by paleontological analysis At its most fundamental level, the fossil re- (e.g., ref. 27). of the peculiar ecosystems of their times. cord is a narrative of changes to phenotypes Extinction can generate a false signal re- and their functions: the origin, persistence, garding the origin of evolutionary novelties Author contributions: D.J. and N.H.S. wrote the paper. – and demise of biological form (1 4), along when only extant taxa are analyzed (28, 29). The authors declare no conflict of interest. with changes in behavior, physiology, and life For example, phylogenetic analysis of extant 1To whom correspondence may be addressed. Email: djablons@ history of vertebrates, invertebrates, plants, chordates suggests that bone evolved follow- uchicago.edu or [email protected]. 4852–4858 | PNAS | April 21, 2015 | vol. 112 | no. 16 www.pnas.org/cgi/doi/10.1073/pnas.1505146112 Downloaded by guest on September 28, 2021 Kidwell much a document of clade failure as suc- INTRODUCTION Goswami et al. cess, and provides a direct observational SPECIAL FEATURE: Jackson & Blois window on a wide range of natural exper- White et al. iments, in deep time and directly preceding S. Huang et al. the present biological moment. D. Huang et al. Atthesametime,anotherkeyfinding Slater from the fossil record is a lack of extinction Pieretti et al. where it might be expected. For example, one Pieretti et al. of the great contributions of fossil data is the Pieretti et al. demonstration that species tend to respond Hunt et al. to climate changes by individualistic range Droser & Gehling shifts, despite the web of positive and nega- Brasier et al. tive interactions seen in every present-day Brasier et al. site. Thus, biotic associations have been dis- banded and assembled during repeated gla- cial cycles without significant extinction or Protero- Archean zoic Paleozoic Mesozoic Cenozoic evident disruption of energy flow through 3 1 500 50 5 2.5 105 novel (“non-analog”)systems(72).Thead- Ga Ma Ka dition of anthropogenic pressures—not the Geologic time least migration barriers in the form of high- Fig. 1. Distribution through geologic time of the contributions to this Special Feature. Ga, billion (109) y ago; Ka = ways, cultivated land, and cities and sub- thousand (103) y ago; Ma, million (106)yago. urbs—has the potential to overturn such resiliency, and paleontological data on what biotas “want to do” in response to climatic – Examples include the origin of vertebrates, traits, can change over time (e.g., refs. 56 59). and other drivers provide a valuable baseline arthropods, mollusks, and echinoderms This paleontological access to long-term perspective from which to view the likely (among others) in late Proterozoic and evolutionary dynamics allows empirical acceleration of future species movements. Cambrian seas with the initial establishment evaluation of multilevel evolutionary pro- Larger-scale biotic interchanges, which were of macrofaunal foodwebs (42), the Paleozoic- cesses, and the relative contribution of traits often asymmetric between donor and re- early Mesozoic diversification of major insect at organismic, species, and even clade level in cipient regions in the geologic past as they are clades in a world before flowering plants (43), the waxing and waning of clades through today (73), further allow comparative evalu- and the protracted evolution of oceanic eco- time (2, 7, 60, 61). A major area of ongoing ation of the major determinants and long- systems lacking the mineralized phytoplank- work on this general topic involves the in- term consequences of successful range shifts ton that only became major factors in the tegration of paleontological and molecular of different magnitudes. At least one marine late Mesozoic and Cenozoic (44). phylogenetic data, particularly when one or interchange
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