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Fossil calibrations and molecular divergence time estimates in centrarchid fishes (Teleostei: Centrarchidae).

Molecular clock methods allow biologists to estimate divergence times, which in turn play an important role in comparative studies of many evolutionary processes. It is well known that molecular age estimates can be biased by heterogeneity in rates of molecular evolution, but less attention has been paid to the issue of potentially erroneous fossil calibrations. In this study we estimate the timing of diversification in Centrarchidae, an endemic major lineage of the diverse North American freshwater fish fauna, through a new approach to fossil calibration and molecular evolutionary model selection. Given a completely resolved multi-gene molecular phylogeny and a set of multiple fossil-inferred age estimates, we tested for potentially erroneous fossil calibrations using a recently developed fossil cross-validation. We also used fossil information to guide the selection of the optimal molecular evolutionary model with a new fossil jackknife method in a fossil-based model cross-validation. The centrarchid phylogeny resulted from a mixed-model Bayesian strategy that included 14 separate data partitions sampled from three mtDNA and four nuclear genes. Ten of the 31 interspecific nodes in the centrarchid phylogeny were assigned a minimal age estimate from the centrarchid fossil record. Our analyses identified four fossil dates that were inconsistent with the other fossils, and we removed them from the molecular dating analysis. Using fossil-based model cross-validation to determine the optimal smoothing value in penalized likelihood analysis, and six mutually consistent fossil calibrations, the age of the most recent common ancestor of Centrarchidae was 33.59 million years ago (mya). Penalized likelihood analyses of individual data partitions all converged on a very similar age estimate for this node, indicating that rate heterogeneity among data partitions is not confounding our analyses. These results place the origin of the centrarchid radiation at a time of major faunal turnover as the fossil record indicates that the most diverse lineages of the North American freshwater fish fauna originated at the Eocene-Oligocene boundary, approximately 34 mya. This time coincided with major global climate change from warm to cool temperatures and a signature of elevated lineage extinction and origination in the fossil record across the tree of life. Our analyses demonstrate the utility of fossil cross-validation to critically assess individual fossil calibration points, providing the ability to discriminate between consistent and inconsistent fossil age estimates that are used for calibrating molecular phylogenies.

Animals↗

Testing the molecular clock: molecular and paleontological estimates of divergence times in the Echinoidea (Echinodermata).

The phylogenetic relationships of 46 echinoids, with representatives from 13 of the 14 ordinal-level clades and about 70% of extant families commonly recognized, have been established from 3 genes (3,226 alignable bases) and 119 morphological characters. Morphological and molecular estimates are similar enough to be considered suboptimal estimates of one another, and the combined data provide a tree that, when calibrated against the fossil record, provides paleontological estimates of divergence times and completeness of their fossil record. The order of branching on the cladogram largely agrees with the stratigraphic order of first occurrences and implies that their fossil record is more than 85% complete at family level and at a resolution of 5-Myr time intervals. Molecular estimates of divergence times derived from applying both molecular clock and relaxed molecular clock models are concordant with estimates based on the fossil record in up to 70% of cases, with most concordant results obtained using Sanderson's semiparametric penalized likelihood method and a logarithmic-penalty function. There are 3 regions of the tree where molecular and fossil estimates of divergence time consistently disagree. Comparison with results obtained when molecular divergence dates are estimated from the combined (morphology + gene) tree suggests that errors in phylogenetic reconstruction explain only one of these. In another region the error most likely lies with the paleontological estimates because taxa in this region are demonstrated to have a very poor fossil record. In the third case, morphological and paleontological evidence is much stronger, and the topology for this part of the molecular tree differs from that derived from the combined data. Here the cause of the mismatch is unclear but could be methodological, arising from marked inequality of molecular rates. Overall, the level of agreement reached between these different data and methodological approaches leads us to believe that careful application of likelihood and Bayesian methods to molecular data provides realistic divergence time estimates in the majority of cases (almost 80% in this specific example), thus providing a remarkably well-calibrated phylogeny of a character-rich clade of ubiquitous marine benthic invertebrates.

Animals↗

Origins of spondyloarthropathy in Perissodactyla.

OBJECTIVE: Spondyloarthropathy has clearly been documented as not limited in occurrence to humans. Transmammalian in nature, it is of interest to understand the antiquity, and perhaps the origins, of this disorder in animal groups sufficiently represented in the skeletal record. METHODS: Fossil and recent skeletons of perissodactylae from North America were systematically examined to determine the occurrence and population frequency of spondyloarthropathy. RESULTS: Spondyloarthropathy was the most common form of arthritis recognized in the extant and fossil records. Common in extinct families such as Brontotheriidae and Chalicotheriidae, a progressive increase in the frequency of spondyloarthropathy was observed through geologic time in Equidae and Rhinocerotidae. CONCLUSION: Erosive arthritis of the spondyloarthropathy variety is now documented as not only persisting in Perissodactyla, but as actually increasing significantly in frequency (3-6 fold). Given the unusual evolutionary penetrance of this "disease," the possibility must be considered that its persistence provides evidence for some unknown benefit to the affected host.

Animals↗

A model for diurnal patterns of carbon fixation in a Precambrian microbial mat based on a modern analog.

Microbial mat communities are one of the first and most prevalent biological communities known from the Precambrian fossil record. These fossil mat communities are found as laminated sedimentary rock structures called stromatolites. Using a modern microbial mat as an analog for Precambrian stromatolites, a study of carbon fixation during a diurnal cycle under ambient conditions was undertaken. The rate of carbon fixation depends primarily on the availability of light (consistent with photosynthetic carbon fixation) and inorganic carbon, and not nitrogen or phosphorus. Atmospheric PCO2 is thought to have decreased from 10 bars at 4 Ga (10(9) years before present) to approximately 10(-4) bars today, implying a change in the availability of inorganic carbon for carbon fixation. Experimental manipulation of levels of inorganic carbon to levels that may have been available to Precambrian mat communities resulted in increased levels of carbon fixation during daylight hours. Combining these data with models of daylength during the Precambrian, models are derived for diurnal patterns of photosynthetic carbon fixation in a Precambrian microbial mat community. The models suggest that, even in the face of shorter daylengths during the Precambrian, total daily carbon fixation has been declining over geological time, with most of the decrease having occurred during the Precambrian.

Bacteria↗

Antiquity of arthritis: spondyloarthropathy identified in the Paleocene of North America. off.

OBJECTIVE: Determination of the antiquity of spondyloarthropathy by examining the fossil record. METHODS: Fossil collections from the Paleocene and Eocene of North America were systematically examined to determine the occurrence and population frequency of spondyloarthropathy. RESULTS: Spondyloarthropathy was present in three mammalian orders dated at 30 to 50 million years. The frequency was 13% to 50% in the affected genera. CONCLUSIONS: Erosive arthritis of the spondyloarthropathy variety is now documented to have originated at least 40 million years ago in the Paleocene.

Animals↗

Glutoxylon Chowdhury (Anacardiaceae): the first record of fossil wood from Bangladesh.

This paper documents the first record of silicified fossil wood from a previously undescribed wood-rich horizon in the Sitakund Anticline, Eastern Bangladesh. The outcrop is composed of cross-stratified, fine-medium grained sandstones with bidirectional cross stratification indicative of a tidal environment, deposited ca. 5-5.2 million years before present (Miocene/Pliocene). The wood is characterised by large solitary vessels with alternate intervascular pits, banded parenchyma, uniseriate rays, and multiseriate rays with one radial canal per ray. This character combination closely resembles the wood of extant Gluta L. of the Anacardiaceae. This specimen has been assigned to the organ genus Glutoxylon Chowdhury erected for fossil woods with anatomical similarity to Gluta (including Melanorrhoea Wall.). The excellent preservation of this mature wood specimen illustrates the potential for using fossil wood from the Sitakund locality for palaeoecological studies in terms of biodiversity and adaptational response to climate change. Moreover such investigations of fossil woods from Bangladesh will compliment studies undertaken on fossil plants in other parts of Central and Southeastern Asia which will further the understanding of plant migration routes between India and Southeast Asia during the Tertiary.

Journal Article↗

Investigating human evolutionary history.

We rely on fossils for the interpretation of more than 95% of our evolutionary history. Fieldwork resulting in the recovery of fresh fossil evidence is an important component of reconstructing human evolutionary history, but advances can also be made by extracting additional evidence for the existing fossil record, and by improving the methods used to interpret the fossil evidence. This review shows how information from imaging and dental microstructure has contributed to improving our understanding of the hominin fossil record. It also surveys recent advances in the use of the fossil record for phylogenetic inference.

Animals↗

The problem of missing links: today and yesterday.

The problems of "missing links" over the years since Darwin's On the Origin of Species is analyzed in the perspective of our increased knowledge of the fossil record and our changing concepts of evolution, geology, and biology. The history of application of evolutionary theory to the fossil record and the consequences of the conflict of categorical, essential classification and evolutionary continuity are examined, as they relate to the "naive" idea of "missing links". Transitions between taxonomic categories are treated at two levels: the infraspecies-species level, and the supraspecies level. In the latter category case analyses of transitions within and between the major groups of metazoans and metaphytes are made, with special attention to the evidence of origins of groups in the fossil record. Throughout, the impacts on evolutionary theory of changes in knowledge and interpretations of the record of the history of life are considered. The problem of the existence of linkages and phylogenies at the species and generic levels has been much reduced during the last one hundred and twenty years. How this reduction supports or denies Darwin's concepts of phyletic gradualism is still a matter of interpretation of the evidence. At familial and higher levels, the establishment of linkages between categories has been much less successful, and decreasingly so at each successive higher level. Under the very best circumstances, however, morphological and stratigraphically graded transitions between classes and subclasses have been found. At the level of phyla and higher categories, any information on transitions as far as the fossil record is concerned is essentially non-existent. Fairly standard patterns of transitions between high categories can be established on the basis of the optimal cases, and these point up the continuing problems in evolutionary theory as being the interrelationships and integration of micro-evolutionary and macro-evolutionary processes.

Amphibians↗

Parting ways: Pan-Homo divergence revisited.

The timing of divergence between hominins and the bonobo-chimpanzee clade has been at the core of palaeoanthropological debate for over a century. The earliest molecular studies indicated divergence times ranging from 5 Ma to as recently as 1.3 Ma. This study critically reviews the trends of time estimates published between 1967 and 2023, and analyses how these are supported or rejected by the current molecular and fossil records. We compiled 202 divergence estimates and defined three distinct thresholds based on fossil evidence at 4.4 Ma (Australopithecus anamensis and Ardipithecus ramidus), 6.2 Ma (Orrorin tugenensis and Ardipithecus kadabba), and 7.2 Ma (Sahelanthropus tchadensis). We then used these thresholds to filter out molecular estimates that are too young to fit the fossil record. Overall, the data suggests a divergence event within the late Miocene, with each threshold pushing it further back, 8.63-6.38, 10.33-7.81, and 10.95-8.81 Ma, respectively. We use a quadratic regression to demonstrate that estimates have been slowly shifting from ~ 6 Ma to ~ 8.5 Ma over the past 56 years. A Bayesian meta-analysis of genomic estimates filtered by our most consensual threshold (i.e., assuming Australopithecus belongs to Hominini) indicates that the split must have occurred early in the late Miocene, most likely before 7 Ma (~ 99.5% posterior probability) with a pooled effect of 8.69-7.28 Ma. We conclude that, despite an initial bias towards younger estimates, the molecular timing for the last common ancestor (LCA) of Pan-Homo has been progressively approaching the intervals suggested by the current fossil record.

Animals↗

Geologic constraints on the macroevolutionary history of marine animals.

The causes of mass extinctions and the nature of taxonomic radiations are central questions in paleobiology. Several episodes of taxonomic turnover in the fossil record, particularly the major mass extinctions, are generally thought to transcend known biases in the geologic record and are widely interpreted as distinct macroevolutionary phenomena that require unique forcing mechanisms. Here, by using a previously undescribed compilation of the durations of sedimentary rock sequences, I compare the rates of expansion and truncation of preserved marine sedimentary basins to rates of origination and extinction among Phanerozoic marine animal genera. Many features of the highly variable record of taxonomic first and last occurrences in the marine animal fossil record, including the major mass extinctions, the frequency distribution of genus longevities, and short- and long-term patterns of genus diversity, can be predicted on the basis of the temporal continuity and quantity of preserved sedimentary rock. Although these results suggest that geologically mediated sampling biases have distorted macroevolutionary patterns in the fossil record, preservation biases alone cannot easily explain the extent to which the sedimentary record duplicates paleobiological patterns. Instead, these results suggest that the processes responsible for producing variability in the sedimentary rock record, such as plate tectonics and sea-level change, may have been dominant and consistent macroevolutionary forces throughout the Phanerozoic.

Animals↗

Evolutionary and preservational constraints on origins of biologic groups: divergence times of eutherian mammals.

Some molecular clock estimates of divergence times of taxonomic groups undergoing evolutionary radiation are much older than the groups' first observed fossil record. Mathematical models of branching evolution are used to estimate the maximal rate of fossil preservation consistent with a postulated missing history, given the sum of species durations implied by early origins under a range of species origination and extinction rates. The plausibility of postulated divergence times depends on origination, extinction, and preservation rates estimated from the fossil record. For eutherian mammals, this approach suggests that it is unlikely that many modern orders arose much earlier than their oldest fossil records.

Animals↗

The role of Quaternary environmental change in plant macroevolution: the exception or the rule?

The Quaternary has been described as an important time for genetic diversification and speciation. This is based on the premise that Quaternary climatic conditions fostered the isolation of populations and, in some instances, allopatric speciation. However, the 'Quaternary Ice-Age speciation model' rests on two key assumptions: (i) that biotic responses to climate change during the Quaternary were significantly different from those of other periods in Earth's history; and (ii) that the mechanisms of isolation during the Quaternary were sufficient in time and space for genetic diversification to foster speciation. These assumptions are addressed by examining the plant fossil record for the Quaternary (in detail) and for the past 410 Myr, which encompasses previous intervals of icehouse Earth. Our examination of the Quaternary record indicates that floristic responses to climate changes during the past 1.8 Myr were complex and that a distinction has to be made between those plants that were able to withstand the extremes of glacial conditions and those that could not. Generation times are also important as are different growth forms (e.g. herbaceous annuals and arborescent perennials), resulting in different responses in terms of genetic divergence rates during isolation. Because of these variations in the duration of isolation of populations and genomic diversification rates, no canonical statement about the predominant floristic response to climatic changes during the Quaternary (i.e. elevated rates of speciation or extinction, or stasis) is currently possible. This is especially true because of a sampling bias in terms of the fossil record of tree species over that of species with non-arborescent growth forms. Nevertheless, based on the available information, it appears that the dominant response of arborescent species during the Quaternary was extinction rather than speciation or stasis. By contrast, our examination of the fossil record of vascular plants for the past 410 Myr indicates that speciation rates often increased during long intervals of icehouse Earth (spanning up to 50 Myr). Therefore, longer periods of icehouse Earth than those occurring during the Quaternary may have isolated plant populations for sufficiently long periods of time to foster genomic diversification and allopatric speciation. Our results highlight the need for more detailed study of the fossil record in terms of finer temporal and spatial resolution than is currently available to examine the significance of intervals of icehouse Earth. It is equally clear that additional and detailed molecular studies of extant populations of Quaternary species are required in order to determine the extent to which these 'relic' species have genomically diversified across their current populations.

Biological Evolution↗

Live birth in Cretaceous marine lizards (mosasauroids).

Although live-bearing (viviparity) has evolved around 100 times within reptiles, evidence of it is almost never preserved in the fossil record. Here, we report viviparity in mosasauroids, a group of Cretaceous marine lizards. This is the only known fossil record of live-bearing in squamates (lizards and snakes), and might represent the oldest occurrence of the trait in this diverse group; it is also the only known fossil record of viviparity in reptiles other than ichthyosaurs. An exceptionally preserved gravid female of the aigialosaur Carsosaurus (a primitive mosasauroid) contains at least four advanced embryos distributed along the posterior two-thirds of the long trunk region (dorsal vertebrae 9-21). Their orientation suggests that they were born tail-first (the nostrils emerging last) to reduce the possibility of drowning, an adaptation shared with other highly aquatic amniotes such as cetaceans, sirenians and ichthyosaurs; the orientation of the embryos also suggests that they were not gut contents because swallowed prey are usually consumed head-first. One embryo is located within the pelvis, raising the possibility that the adult died during parturition. Viviparity in early medium-sized amphibious aigialosaurs may have freed them from the need to return to land to deposit eggs, and permitted the subsequent evolution of gigantic totally marine mosasaurs.

Animals↗

The relationships of vascular plants.

Recent phylogenetic research indicates that vascular plants evolved from bryophyte-like ancestors and that this involved extensive modifications to the life cycle. These conclusions are supported by a range of systematic data, including gene sequences, as well as evidence from comparative morphology and the fossil record. Within vascular plants, there is compelling evidence for two major clades, which have been termed lycophytes (clubmosses) and euphyllophytes (seed plants, ferns, horsetails). The implications of recent phylogenetic work are discussed with reference to life cycle evolution and the interpretation of stratigraphic inconsistencies in the early fossil record of land plants. Life cycles are shown to have passed through an isomorphic phase in the early stages of vascular plant evolution. Thus, the gametophyte generation of all living vascular plants is the product of massive morphological reduction. Phylogenetic research corroborates earlier suggestions of a major representational bias in the early fossil record. Mega-fossils document a sequence of appearance of groups that is at odds with that predicted by cladogram topology. It is argued here that the pattern of appearance and diversification of plant megafossils owes more to changing geological conditions than to rapid biological diversification.

Biological Evolution↗

Identifying the morphological signatures of hybridization in primate and human evolution.

Recent studies point to contact and possible admixture among contemporaneous hominin species during the Plio-Pleistocene. However, detection of hybridization in fossils-and especially fossil hominins-is contentious, and it is hindered in large part by our lack of understanding about how morphological hybridity is manifested in the primate skeleton. Here, we report on a study of known-pedigree, purebred yellow and olive baboons (n = 112) and their hybrids (n = 57), derived from the baboon colony of the Southwest Foundation for Biomedical Research. The hybrids were analyzed in two different groups: (1) F1 = olive x yellow first-generation hybrids; (2) B1 = olive x F1 backcross hybrids. Thirty-nine metric variables were tested for heterosis and dysgenesis. Nonmetric data were also collected from the crania. Results show that these primate hybrids are somewhat heterotic relative to their parental populations, are highly variable, and display novel phenotypes. These effects are most evident in the dentition and probably indicate the mixing of two separately coadapted genomes and the breakdown in the coordination of early development, despite the fact that these populations diverged fairly recently. Similar variation is also observed in museum samples drawn from natural hybrid zones. The results offer a strategy for detecting hybrid zones in the fossil record; implications for interpreting the hominin fossil record are discussed.

Analysis of Variance↗

Human evolution.

The common ancestor of modern humans and the great apes is estimated to have lived between 5 and 8 Myrs ago, but the earliest evidence in the human, or hominid, fossil record is Ardipithecus ramidus, from a 4.5 Myr Ethiopian site. This genus was succeeded by Australopithecus, within which four species are presently recognised. All combine a relatively primitive postcranial skeleton, a dentition with expanded chewing teeth and a small brain. The most primitive species in our own genus, Homo habilis and Homo rudolfensis, are little advanced over the australopithecines and with hindsight their inclusion in Homo may not be appropriate. The first species to share a substantial number of features with later Homo is Homo ergaster, or 'early African Homo erectus', which appears in the fossil record around 2.0 Myr. Outside Africa, fossil hominids appear as Homo erectus-like hominids, in mainland Asia and in Indonesia close to 2 Myr ago; the earliest good evidence of 'archaic Homo' in Europe is dated at between 600-700 Kyr before the present. Anatomically modern human, or Homo sapiens, fossils are seen first in the fossil record in Africa around 150 Kyr ago. Taken together with molecular evidence on the extent of DNA variation, this suggests that the transition from 'archaic' to 'modern' Homo may have taken place in Africa.

Animals↗

The microfossil record of early land plants.

Dispersed microfossils (spores and phytodebris) provide the earliest evidence for land plants. They are first reported from the Llanvirn (Mid-Ordovician). More or less identical assemblages occur from the Llanvirn (Mid-Ordovician) to the late Llandovery (Early Silurian), suggesting a period of relative stasis some 40 Myr in duration. Various lines of evidence suggest that these early dispersed microfossils derive from parent plants that were bryophyte-like if not in fact bryophytes. In the late Llandovery (late Early Silurian) there was a major change in the nature of dispersed spore assemblages as the separated products of dyads (hilate monads) and tetrads (trilete spores) became relatively abundant. The inception of trilete spores probably represents the appearance of vascular plants or their immediate progenitors. A little later in time, in the Wenlock (early Late Silurian), the earliest unequivocal land plant megafossils occur. They are represented by rhyniophytoids. It is only from the Late Silurian onwards that the microfossil/ megafossil record can be integrated and utilized in interpretation of the flora. Dispersed microfossils are preserved in vast numbers, in a variety of environments, and have a reasonable spatial and temporal fossil record. The fossil record of plant megafossils by comparison is poor and biased, with only a dozen or so known pre-Devonian assemblages. In this paper, the early land plant microfossil record, and its interpretation, are reviewed. New discoveries, novel techniques and fresh lines of inquiry are outlined and discussed.

Biological Evolution↗

Systematics of cetaceans using restriction site mapping of mitochondrial DNA.

Phylogenetic analysis of 14 cetacean species, including members from two baleen whale families and three toothed whale families, was undertaken using restriction site mapping of mitochondrial DNA and using cladistic and distance measures to infer phylogenies. The amount of between-taxa sequence divergence inferred from the data was lower than expected from the standard interpretation of the fossil record, but more consistent with some recent estimates of sequence divergence in cetacean mitochondrial DNA or nuclear DNA. This implies either that the rate of molecular evolution of cetacean DNA is much lower than that of other mammalian orders or that the fossil record of cetaceans requires reinterpretation. The incompleteness of the cetacean fossil record precludes resolution of the paradox at the present time. However, this discrepancy could in part be attributed to the sampling error inherent in the restriction site mapping technique, as comparative studies using the complete mtDNA genome and restriction site data of the blue and fin whales (genus Balaenoptera) indicate that the restriction site maps underestimate sequence divergence by about 40%. In contrast to a recent study suggesting that toothed whales were paraphyletic, with the sperm whales being more closely related to the rorquals than to the other toothed whales, the restriction data tend to support the monophyly of the baleen and the toothed whales, a finding which is consistent with a recent molecular-based study and with morphological and paleontological data. Topologies of the subfamily and generic levels are generally consistent with morphologically based schemes.

Animals↗