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M O N O G R a P February 2012 VOL. 82 • NO. 1 • FEBRUARY 2012 ISSN 0012-9615 VOL. Ecological CONTENTS 82, MONOGRAPHS Volume 82 No. 1 1 NO. A PUBLICATION OF THE ECOLOGICAL SOCIETY OF AMERICA Editorial 1, 1–148 Articles 3 Determinants and dynamics of banded vegetation pattern migration in arid climates • VINCENT DEBLAUWE, PIERRE COUTERON, JAN BOGAERT, AND NICOLAS BARBIER 23 Floral and environmental gradients on a Late Cretaceous landscape • SCOTT L. WING, CAROLINE A. E. STRÖMBERG, LEO J. HICKEY, FLEUR TIVER, BRIAN WILLIS, ROBYN J. BERNHAM, AND ANNA K. BEHRENSMEYER 49 Resilience and regime change in a southern Rocky Mountain ecosystem during the past 17 000 years • T. A. MINCKLEY, R. K. SHRIVER, AND B. SHUMAN 69 Wildfire provides refuge from local extinction but is an unlikely driver of outbreaks by mountain pine beetle • ERINN N. POWELL, PHILIP A. TOWNSEND, AND KENNETH F. RAFFA 85 Control of arthropod abundance, richness, and composition in a heterogeneous desert city • CHRISTOFER BANG, STANLEY H. FAETH, AND JOHN L. SABO 101 Controls on carbon dynamics by ecosystem structure and climate for southeastern U.S. slash pine plantations • ROSVEL BRACHO, GREGORY STARR, HENRY L. GHOLZ, TIMOTHY A. MARTIN, WENDELL P. CROPPER, AND HENRY W. LOESCHER 129 Framework to improve the application of theory in ecology and conservation • DON A. DRISCOLL AND DAVID B. LINDENMAYER Ecological Archives Appendices and Supplements are available online: http://esapubs.org/archive ESA Data Registry Available online: http://data.esa.org Instructions to Authors Available online: http://esapubs.org FEBRUARY Articles Determinants and dynamics of banded vegetation pattern migration in arid climates 2012 Floral and environmental gradients on a Late Cretaceous landscape Cover Photo: Fossil of Anemia fremontii (Knowlton), Big Cedar Ridge, Wyoming, USA (~73 Ma). This Resilience and regime change in a southern Rocky Mountain ecosystem in situ fossil flora documents that vegetation co-dominated by ferns and angiosperms persisted in sub- tropical, mid-latitude North America for tens of millions of years following the major angiosperm radia- during the past 17 000 years tion. Diversity of ferns was also high compared with modern vegetation in similar habitats (see Wing et al. pp. 23–47). Photo credit: S. L. Wing. Ecological Monographs, 82(1), 2012, pp. 23–47 Ó 2012 by the Ecological Society of America Floral and environmental gradients on a Late Cretaceous landscape 1,7 2 3 4 5 6 SCOTT L. WING, CAROLINE A. E. STRO¨MBERG, LEO J. HICKEY, FLEUR TIVER, BRIAN WILLIS, ROBYN J. BURNHAM, 1 AND ANNA K. BEHRENSMEYER 1Department of Paleobiology, NHB121, Smithsonian Institution, Washington, D.C. 20013-7012 USA 2Department of Biology and Burke Museum of Natural History and Culture, University of Washington, Seattle, Washington 98195-1800 USA 3Department of Geology and Geophysics, Yale University, P.O. Box 208109, 210 Whitney Ave., New Haven, Connecticut 06520 USA 4School of Natural and Built Environments, Mawson Lakes Campus, University of South Australia, South Australia 5095 Australia 5Clastic R and D, Chevron Energy Technology Company, 1500 Louisiana Street, Houston, Texas 77002 USA 6Museum of Paleontology, University of Michigan, 1109 Geddes Avenue, Ann Arbor, Michigan 48109-1079 USA Abstract. We describe an in situ fossil flora of Late Cretaceous age (;73 Ma [mega- annum or million years]) from Big Cedar Ridge in central Wyoming, USA, which we sampled using a modified line-intercept method to quantify the relative abundances of 122 taxa at 100 sites across 4 km of exposed sedimentary deposits. We also measured three physical variables at each site: paleotopographic level, grain size, and total organic content. Paleoenvironmental conditions and paleofloral composition at Big Cedar Ridge covary strongly and are highly heterogeneous on small spatial scales. The reconstructed vegetation has some similarities with extant topogenous fens, but also important differences. Non-monocot angiosperms were abundant only on wet, mineral substrates that had been disturbed shortly before preservation, consistent with the weedy life histories that are inferred for their Early Cretaceous ancestors. Many non-monocot angiosperms grew in small, dispersed populations, consistent with the hypothesis that they were biotically pollinated. Overall, non-monocot angiosperm abundance was low compared with many modern wetlands. A single species of coryphoid palm was the dominant on moist, stable, moderately organic-rich sites, a pattern seen in some subtropical to tropical wetlands in the present day. Fern thickets at Big Cedar Ridge occupied highly organic, possibly low-nutrient substrates, and were dominated by Dipteridaceae, Gleichenia- ceae, Schizaeaceae, and Matoniaceae. The overall high diversity and abundance of pteridophytes is unusual in the context of modern vegetation, regardless of climate zone, and probably represents a late occurrence of pteridophyte-dominated vegetation that was common earlier in the Mesozoic. Plant distributions at Big Cedar Ridge combine aspects of pre-angiosperm and modern vegetation in a way that suggests both niche conservatism and niche evolution on geological time scales. Key words: angiosperms; Big Cedar Ridge, Wyoming, USA; disturbance; ferns; nutrient stress; paleoecology. INTRODUCTION understory sites in wet climates (Feild et al. 2004, 2009), to the proposition that angiosperm-dominated forests As phylogenetic data, techniques, and understanding arose in the mid-Cretaceous (Davis et al. 2005, Wang et have improved rapidly in recent years, it has become al. 2009), and to the idea that the most diverse clades of increasingly common to reconstruct the ecological living ferns radiated in low-light environments under history of groups using phylogenetic trees derived from angiosperm canopies in the Late Cretaceous and early analyses of living species (e.g., Feild et al. 2004, 2009, Cenozoic (Schneider et al. 2004, Schuettpelz and Pryer Davis et al. 2005, Wang et al. 2009). The basic procedure 2009). Although the phylogenetic approach has many is to develop a phylogenetic tree from genetic sequence advantages, the ecological traits of living species exert a data, calibrate it to geological time using well-dated strong influence on conclusions about the ecology of fossils of known phylogenetic position, score the extinct taxa. In particular, if the ecological traits or terminals for an ecological trait, and then infer from preferences of the extinct species in a group were outside parsimony or likelihood analyses the ecological prefer- the range encompassed by their living descendants, ences of dated nodes. Phylogenetic insights of this kind phylogenetic reconstructions will be incorrect, and this is have led, for example, to the hypothesis that the earliest particularly likely in lineages that are old enough to have angiosperms were small woody plants of disturbed, many extinct species. Discovering if and how the past really is different from the present still requires data Manuscript received 15 May 2011; revised and accepted 9 August 2011; final version received 6 September 2011. from the fossil record. Corresponding Editor: A. H. Lloyd. The purpose of this paper is to demonstrate the large 7 E-mail: [email protected]. amount of paleoecological information that can be 23 24 SCOTT L. WING ET AL. Ecological Monographs Vol. 82, No. 1 deduced from study of an in situ fossil flora, and to give striking example of phylogenetic niche conservatism insight into landscape-scale patterns of floral distribu- (Prinzing et al. 2001, Crisp et al. 2009) during a major tion in the Late Cretaceous. We are particularly evolutionary radiation. interested in Cretaceous vegetation because it is the In this paper we describe the floral and environmental geological period during which angiosperms evolved, gradients preserved on a Late Cretaceous (;73 Ma) radiated, and became common in terrestrial vegetation. landscape at Big Cedar Ridge in central Wyoming, The oldest widely accepted angiosperm fossils are pollen USA. The distribution and abundance of fossil plants grains from the Valanginian Stage of the Early and their correlations with local environmental condi- Cretaceous (140–134 Ma [Hughes 1994]). Crown group tions yield strong inferences about their ecological angiosperms began radiating in the Aptian (125–112 Ma preferences that are independent of both phylogenetic [Hickey and Doyle 1977, Hochuli et al. 2006, Friis et al. position and functional interpretation of morphological 2010]), and the largest proportional increase in angio- traits. Although vegetational patterns observed along an sperm diversity occurred in the Albian-Cenomanian irregular two-dimensional transect in a single deposit, (112–94 Ma), concurrent with origination of the major recording an instant in time, obviously cannot represent extant clades within the most diverse subgroup, the the whole globe, the patterns of plant distribution at Big eudicots (Lidgard and Crane 1988, Lidgard and Crane Cedar Ridge do give direct evidence of the ecological 1990, Crepet et al. 2004, Friis et al. 2010). roles and preferences of major plant groups on this Late The effect of angiosperms on Cretaceous vegetation is Cretaceous landscape. We hope future studies will less well understood than the timing of their diversifi- establish the generality of these patterns. cation. Early angiosperm fossils are rare, and assuming no overwhelming bias against their preservation, it Environmental context seems unlikely they were abundant in pre-Aptian The Big Cedar Ridge
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