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Historians have debated whether pathways and events from the past to the present are influenced largely by contingency, the dependence of outcomes on particular prior conditions, or whether there is long-term emergent directional change. Previous arguments for predictability in evolutionary history relied on the high frequency of convergence, but the repeated evolution of widely favored adaptations need not imply long-term directionality. Using evidence from the fossil record and arguments concerning the metabolic evolution of organisms, I show here that power (total energy taken up and expended per unit time) has increased stepwise over time at ecosystem-level and global scales thanks to the ratchet-like, cumulative effects of competition and cooperation and to the disproportionate influence of powerful top competitors and opportunistic species on emergent ecosystem properties and processes. The history of life therefore exhibits emergent directionality at large ecosystem-wide scales toward greater power.
Previous studies of cranial shape have established a consistent interspecific allometric pattern relating the relative lengths of the face and braincase regions of the skull within multiple families of mammals. In this interspecific allometry, the facial region of the skull is proportionally longer than the braincase in larger species. The regularity and broad taxonomic occurrence of this allometric pattern suggests that it may have an origin near the base of crown Mammalia, or even deeper in the synapsid or amniote forerunners of mammals. To investigate the possible origins of this allometric pattern, we used geometric morphometric techniques to analyze cranial shape in 194 species of nonmammalian synapsids, which constitute a set of successive outgroups to Mammalia. We recovered a much greater diversity of allometric patterns within nonmammalian synapsids than has been observed in mammals, including several instances similar to the mammalian pattern. However, we found no evidence of the mammalian pattern within Therocephalia and nonmammalian Cynodontia, the synapsids most closely related to mammals. This suggests that the mammalian allometric pattern arose somewhere within Mammaliaformes, rather than within nonmammalian synapsids. Further investigation using an ontogenetic series of the anomodont Diictodon feliceps shows that the pattern of interspecific allometry within anomodonts parallels the ontogenetic trajectory of Diictodon. This indicates that in at least some synapsids, allometric patterns associated with ontogeny may provide a “path of least resistance” for interspecific variation, a mechanism that we suggest produces the interspecific allometric pattern observed in mammals.
The estimation of origination and extinction rates and their temporal variation is central to understanding diversity patterns and the evolutionary history of clades. The fossil record provides the only direct evidence of extinction and biodiversity changes through time and has long been used to infer the dynamics of diversity changes in deep time. The software PyRate implements a Bayesian framework to analyze fossil occurrence data to estimate the rates of preservation, origination, and extinction while incorporating several sources of uncertainty. Building upon this framework, we present a suite of methodological advances including more complex and realistic models of preservation and the first likelihood-based test to compare the fit across different models. Further, we develop a new reversible jump Markov chain Monte Carlo algorithm to estimate origination and extinction rates and their temporal variation, which provides more reliable results and includes an explicit estimation of the number and temporal placement of statistically significant rate changes. Finally, we implement a new C++ library that speeds up the analyses by orders of magnitude, therefore facilitating the application of the PyRate methods to large data sets. We demonstrate the new functionalities through extensive simulations and with the analysis of a large data set of Cenozoic marine mammals. We compare our analytical framework against two widely used alternative methods to infer origination and extinction rates, revealing that PyRate decisively outperforms them across a range of simulated data sets. Our analyses indicate that explicit statistical model testing, which is often neglected in fossil-based macroevolutionary analyses, is crucial to obtain accurate and robust results.
Preferential dissolution of the biogenic carbonate polymorph aragonite promotes preservational bias in shelly marine faunas. While field studies have documented the impact of preferential aragonite dissolution on fossil molluscan diversity, its impact on regional and global biodiversity metrics is debated. Epicontinental seas are especially prone to conditions that both promote and inhibit preferential dissolution, which may result in spatially extensive zones with variable preservation. Here we present a multifaceted evaluation of aragonite dissolution within the Late Cretaceous Western Interior Seaway of North America. Occurrence data of mollusks from two time intervals (Cenomanian/Turonian boundary, early Campanian) are plotted on new high-resolution paleogeographies to assess aragonite preservation within the seaway. Fossil occurrences, diversity estimates, and sampling probabilities for calcitic and aragonitic fauna were compared in zones defined by depth and distance from the seaway margins. Apparent range sizes, which could be influenced by differential preservation potential of aragonite between separate localities, were also compared. Our results are consistent with exacerbated aragonite dissolution within specific depth zones for both time slices, with aragonitic bivalves additionally showing a statistically significant decrease in range size compared with calcitic fauna within carbonate-dominated Cenomanian–Turonian strata. However, we are unable to conclusively show that aragonite dissolution impacted diversity estimates. Therefore, while aragonite dissolution is likely to have affected the preservation of fauna in specific localities, time averaging and instantaneous preservation events preserve regional biodiversity. Our results suggest that the spatial expression of taphonomic biases should be an important consideration for paleontologists working on paleobiogeographic problems.
It was once thought that the endemic carnivorous mammals of South America, the metatherian sparassodonts, were driven extinct by North American carnivorans through competitive exclusion. However, sparassodonts went extinct before most groups of carnivorans entered South America; only the endemic Cyonasua-group procyonids (Cyonasua and Chapalmalania), which immigrated to South America nearly 4 million years earlier than other carnivorans, significantly overlapped with sparassodonts in time. In this study, we examine the functional morphology of the dentition of Cyonasua and Chapalmalania through quantitative analysis to determine the dietary habits of these taxa and the degree to which they may have ecologically overlapped sparassodonts and large predatory Neogene didelphimorphians. We find Cyonasua and Chapalmalania to be more carnivorous than extant procyonids, other than Bassariscus, in agreement with previous studies, but more omnivorous than most other carnivorans and all meat-eating South American metatherians, including sparassodonts. The extreme ecological dissimilarity between Cyonasua-group procyonids and members of the endemic South American predator guild may explain why procyonids were able to successfully establish themselves in South America several million years earlier than most other northern mammals (including all other carnivorans): they moved into a previously unoccupied ecological niche (large omnivore) and avoided direct competition with incumbent native species, a situation similar to that documented in historical cases of biological invasion. The omnivorous diets and climbing/swimming abilities of procyonids may have increased their chances for a successful over-water dispersal relative to other carnivorans, further favoring their successful establishment in South America.
Earliest Paleocene megafloras from North America are hypothesized to be low diversity and dominated by long-lived cosmopolitan species following the Cretaceous/Paleogene (K/Pg) mass extinction. However, megafloras used to develop this hypothesis are from the Northern Great Plains (NGP) of North America, and relatively little is known about floras from southern basins. Here, we present a quantitative analysis of an earliest Paleocene megaflora (<350 kyr after K/Pg boundary) from the Ojo Alamo Sandstone in the San Juan Basin (SJB), New Mexico. The megaflora, comprising 53 morphotypes, was dominated by angiosperms, with accessory taxa composed of pteridophytes, lycophytes, and conifers. Diversity analyses indicate a species-rich, highly uneven, and laterally heterogeneous flora. Paleoclimate estimates using multivariate and univariate methods indicate warm temperatures and relatively high precipitation consistent with a modern tropical seasonal forest.
When compared with contemporaneous floras from the Denver Basin (DB) of Colorado and the Williston Basin (WB) of North Dakota, the SJB flora had significantly higher species richness but lower evenness. Paleoclimate estimates from the SJB were 7–14°C warmer than the estimates for the DB and WB, indicating a shift from a temperate forest in the NGP to a tropical forest in the SJB. These results demonstrate the presence of a latitudinal floral diversity and paleoclimatic gradient during the earliest Paleocene in western North America. We hypothesize that the warm, wet conditions in the earliest Paleocene SJB drove rapid rates of speciation following the K/Pg boundary, resulting in a diverse and heterogeneous flora.
The work of Arthur J. Boucot (vale, 1924–2017) is foundational to Devonian biogeography. Cited more than 1000 times and mentioned in excess of 105,000 times, his classification of the Old World, Eastern Americas, and Malvinokaffric realms has become the framework for Devonian biogeographic and faunal studies. This study is the first to test and quantify the biotic area classification using the distributional data of trilobite, brachiopod, and fish taxa across the Lower, Middle, and Late Devonian. The biotic similarity analysis of 676 genera (ca. 30,000 occurrences) of late Silurian (Ludlow–Pridoli) and Devonian taxa represents the largest quantification of distributional data undertaken in Devonian biogeography. The temporal area approach (TAAp), applied here for the first time, failed to recover in the area phenogram many of the realms and regions used by Devonian biogeographers, excepting the Malvinokaffric realm, across the entirety of the Devonian. The Old World and Western Gondwana realms are found to be artificial and in need of revision. Without natural areas, bioregionalizations (area classifications) become unstable and unusable, resulting in authors creating further artificial regions and provinces. This study offers a better solution, namely, to identify artificial areas in existing area classifications using TAAp and biotic similarity analysis and revise them. Area classifications are modeled on taxonomic process (Wilson and Brown 1953; Starrett 1958) and the movement toward revision is necessary to break the cycle of reinvention of realms and regions within bioregionalization.
Archaeocyaths are the most abundant sponges from the Cambrian period, having formed the first animal reef communities more than 500 million years ago. The Archaeocyatha are index fossils for correlating rocks of similar ages globally because of their abundance, extensive geographic distribution, detailed anatomy, and well-established taxonomy. Their ecological significance remains incompletely explored, yet they are known to have strongly competitively interacted, unlike modern sponges. This study examines the feeding ecology of the fossil remains of Siberian archaeocyath assemblages. As suspension feeders, archaeocyaths filtered plankton from the water column through pores in their outer wall. Here we outline a new method to estimate the limit on the upper size of plankton that could be consumed by an archaeocyath during life. The archaeocyaths examined were predominantly feeding on nanoplankton and microplankton such as phytoplankton and protozooplankton. Size-frequency distributions of pore sizes from six different Siberian archaeocyath assemblages, ranging from Tommotian to Botoman in age, reveal significantly different upper limits to the prey consumed at each locality. Some of the assemblages contain specimens that could have fed on larger organisms extending into the mesoplankton, including micro-invertebrates as a possible food resource. These results show that during the establishment of the first animal reef systems, prey partitioning was established as a way of reducing competition. This method has applicability for understanding the construction and the functioning of the first reef systems, as well as helping to understand modern reef systems and their development though time and space.
Bone healing is an important survival mechanism, allowing vertebrates to recover from injury and disease. Here we describe newly recognized paleopathologies in the hindlimbs of the early tetrapods Crassigyrinus scoticus and Eoherpeton watsoni from the early Carboniferous of Cowdenbeath, Scotland. These pathologies are among the oldest known instances of bone healing in tetrapod limb bones in the fossil record (about 325 Ma). X-ray microtomographic imaging of the internal bone structure of these lesions shows that they are characterized by a mass of trabecular bone separated from the shaft's trabeculae by a layer of cortical bone. We frame these paleopathologies in an evolutionary context, including additional data on bone healing and its pathways across extinct and extant sarcopterygians. These data allowed us to synthesize information on cell-mediated repair of bone and other mineralized tissues in all vertebrates, to reconstruct the evolutionary history of skeletal tissue repair mechanisms. We conclude that bone healing is ancestral for sarcopterygians. Furthermore, other mineralized tissues (aspidin and dentine) were also capable of healing and remodeling early in vertebrate evolution, suggesting that these repair mechanisms are synapomorphies of vertebrate mineralized tissues. The evidence for remodeling and healing in all of these tissues appears concurrently, so in addition to healing, these early vertebrates had the capacity to restore structure and strength by remodeling their skeletons. Healing appears to be an inherent property of these mineralized tissues, and its linkage to their remodeling capacity has previously been underappreciated.