Ancient fossil specimens of extinct species are genetically more distant to an outgroup than extant sister species are

There exists a remarkable correlation between genetic distance as measured by protein or DNA dissimilarity and time of species divergence as inferred from fossil records. This observation has provoked the molecular clock hypothesis. However, data inconsistent with the hypothesis have steadily accumulated in recent years from studies of extant organisms. Here the published DNA and protein sequences from ancient fossil specimens were examined to see if they would support the molecular clock hypothesis. The hypothesis predicts that ancient specimens cannot be genetically more distant to an outgroup than extant sister species are. Also, two distinct ancient specimens cannot be genetically more distant than their extant sister species are. The findings here do not conform to these predictions. Neanderthals are more distant to chimpanzees and gorillas than modern humans are. Dinosaurs are more distant to frogs than extant birds are. Mastodons are more distant to opossums than other placental mammals are. The genetic distance between dinosaurs and mastodons is greater than that between extant birds and mammals. Therefore, while the molecular clock hypothesis is consistent with some data from extant organisms, it has yet to find support from ancient fossils. Far more damaging to the hypothesis than data from extant organisms, which merely question the constancy of mutation rate, the study of ancient fossil organisms here challenges for the first time the fundamental premise of modern evolution theory that genetic distances had always increased with time in the past history of life on Earth.     

Evaluation of the fossil fish‐specific diversity in a chadian continental assemblage: Exploration of morphological continuous variation in Synodontis (Ostariophysi,Siluriformes)

In the fossil record, the quantification of continuous morphological variation has become a central issue when dealing with species identification and speciation. In this context, fossil taxa with living representatives hold great promise, because of the potential to characterise patterns of intraspecific morphological variation in extant species prior to any interpretation in the fossil record. The vast majority of catfish families fulfil this prerequisite, as most of them are represented by extant genera. However, although they constitute a major fish group in terms of distribution, and ecological and taxonomic diversity, the quantitative study of their past morphological variation has been neglected, as fossil specimens are generally identified based on the scarcest remains, that is, complete neurocrania that bear discrete characters. Consequently, a part of freshwater catfish history is unprospected and unknown. In this study, we explored the morphological continuous variation of the humeral plate shape in Synodontis catfishes using Elliptic Fourier Analysis (EFA), and compared extant members and fossil counterparts. We analysed 153 extant specimens of 11 Synodontis species present in the Chad basin, in addition to 23 fossil specimens from the Chadian fossiliferous area of Toros Menalla which is dated around 7 Ma. This highly speciose genus, which is one of the most diversified in Africa, exhibits a rich fossil record with several hundred remains mostly identified as Synodontis sp. The analysis of the outline of the humeral plate reveals that some living morphological types were already represented in the Chad Basin 7 My ago, and allows for the discovery of extinct species. Beside illuminating the complex Neogene evolutionary history of Synodontis, these results underline the interest in the ability of isolated remains to reconstruct a past dynamic history and to validate the relevance of EFA as a tool to explore specific diversity through time. J. Morphol. 277:1486–1496, 2016. © 2016 Wiley Periodicals, Inc.     

The fossil record and limb disparity of enantiornithines, the dominant flying birds of the Cretaceous

Morphometric and stratigraphic analyses that encompass the known fossil record of enantiornithine birds (Enantiornithes) are presented. These predominantly flighted taxa were the dominant birds of the second half of the Mesozoic; the enantiornithine lineage is known to have lasted for at least 60 million years (Ma), up until the end of the Cretaceous. Analyses of fossil record dynamics show that enantiornithine 'collectorship' since the 1980s approaches an exponential distribution, indicating that an asymptote in proportion of specimens has yet to be achieved. Data demonstrate that the fossil record of enantiornithines is complete enough for the extraction of biological patterns. Comparison of the available fossil specimens with a large data set of modern bird (Neornithes) limb proportions also illustrates that the known forelimb proportions of enantiornithines fall within the range of extant taxa; thus these birds likely encompassed the range of flight styles of extant birds. In contrast, most enantiornithines had hindlimb proportions that differ from any extant taxa. To explore this, ternary diagrams are used to graph enantiornithine limb variation and to identify some morphological oddities ( Otogornis , Gobipteryx ); taxa not directly comparable to modern birds. These exceptions are interesting – although anatomically uniform, and similar to extant avians in their wing proportions, some fossil enantiornithines likely had flight styles not seen among their living counterparts.     

Metric variation and sexual dimorphism in the dentition of Ouranopithecus macedoniensis

The fossil sample attributed to the late Miocene hominoid taxon Ouranopithecus macedoniensis is characterized by a high degree of dental metric variation. As a result, some researchers support a multiple-species taxonomy for this sample. Other researchers do not think that the sample variation is too great to be accommodated within one species. This study examines variation and sexual dimorphism in mandibular canine and postcanine dental metrics of an Ouranopithecus sample. Bootstrapping (resampling with replacement) of extant hominoid dental metric data is performed to test the hypothesis that the coefficients of variation (CV) and the indices of sexual dimorphism (ISD) of the fossil sample are not significantly different from those of modern great apes. Variation and sexual dimorphism in Ouranopithecus M(1) dimensions were statistically different from those of all extant ape samples; however, most of the dental metrics of Ouranopithecus were neither more variable nor more sexually dimorphic than those of Gorilla and Pongo. Similarly high levels of mandibular molar variation are known to characterize other fossil hominoid species. The Ouranopithecus specimens are morphologically homogeneous and it is probable that all but one specimen included in this study are from a single population. It is unlikely that the sample includes specimens of two sympatric large-bodied hominoid species. For these reasons, a single-species hypothesis is not rejected for the Ouranopithecus macedoniensis material. Correlations between mandibular first molar tooth size dimorphism and body size dimorphism indicate that O. macedoniensis and other extinct hominoids were more sexually size dimorphic than any living great apes, which suggests that social behaviors and life history profiles of these species may have been different from those of living species.     

Variations and asymmetries in regional brain surface in the genus Homo

Paleoneurology is an important field of research within human evolution studies. Variations in size and shape of an endocast help to differentiate among fossil hominin species whereas endocranial asymmetries are related to behavior and cognitive function. Here we analyse variations of the surface of the frontal, parieto-temporal and occipital lobes among different species of Homo, including 39 fossil hominins, ten fossil anatomically modern Homo sapiens and 100 endocasts of extant modern humans. We also test for the possible asymmetries of these features in a large sample of modern humans and observe individual particularities in the fossil specimens.This study contributes important new information about the brain evolution in the genus Homo. Our results show that the general pattern of surface asymmetry for the different regional brain surfaces in fossil species of Homo does not seem to be different from the pattern described in a large sample of anatomically modern H. sapiens, i.e., the right hemisphere has a larger surface than the left, as do the right frontal, the right parieto-temporal and the left occipital lobes compared with the contra-lateral side. It also appears that Asian Homo erectus specimens are discriminated from all other samples of Homo, including African and Georgian specimens that are also sometimes included in that taxon. The Asian fossils show a significantly smaller relative size of the parietal and temporal lobes. Neandertals and anatomically modern H. sapiens, who share the largest endocranial volume of all hominins, show differences when considering the relative contribution of the frontal, parieto-temporal and occipital lobes. These results illustrate an original variation in the pattern of brain organization in hominins independent of variations in total size. The globularization of the brain and the enlargement of the parietal lobes could be considered derived features observed uniquely in anatomically modern H. sapiens.     

Terpenoids as chemosystematic markers in selected fossil and extant species of pine (Pinus, Pinaceae)

The compositions of terpenoids from the seed cones and resins of selected fossil and modern species of pine ( Pinus , Pinaceae) were analysed using gas chromatography-mass spectrometry (GC-MS) to study the preservation of chemosystematic markers (biomarkers) in fossil conifers. Diterpenoids of the abietane, pimarane, isopimarane, and labdane types are the major components in the solvent extracts of all fossil and extant pines. Mono- and sesquiterpenoids of the bornane and cadalane classes and two phytosterols were also identified. The terpenoid patterns of the fossil species largely match the terpenoids observed in modern pines. The characteristic features of the fossil and extant Pinus species are abietane-, pimarane-, and isopimarane-type diterpenoids with a high abundance of abietic acids. Several biomolecules were preserved unaltered in the Eocene and Oligocene pine cones. In addition to the biomolecules, the fossil pines contain a series of degradation products (geomolecules) that were generated from their biological precursors before and after burial. The generation of the geomolecules is governed by various oxidative and reductive processes, such as microbial degradation and clay catalysis. In conclusion, the comparative analysis of solvent-extractable biomarkers in fossil and related extant species is a valuable tool for (palaeo)chemosystematic studies of conifers.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 154 , 129–140.     

Digit ratios predict polygyny in early apes, Ardipithecus, Neanderthals and early modern humans but not in Australopithecus

Social behaviour of fossil hominoid species is notoriously difficult to predict owing to difficulties in estimating body size dimorphism from fragmentary remains and, in hominins, low canine size dimorphism. Recent studies have shown that the second-to-fourth digit ratio (2D : 4D), a putative biomarker for prenatal androgen effects (PAEs), covaries with intra-sexual competition and social systems across haplorrhines; non-pair-bonded polygynous taxa have significantly lower 2D : 4D ratios (high PAE) than pair-bonded monogamous species. Here, we use proximal phalanx ratios of extant and fossil specimens to reconstruct the social systems of extinct hominoids. Pierolapithecus catalaunicus, Hispanopithecus laietanus and Ardipithecus ramidus have ratios consistent with polygynous extant species, whereas the ratio of Australopithecus afarensis is consistent with monogamous extant species. The early anatomically modern human Qafzeh 9 and Neanderthals have lower digit ratios than most contemporary human populations, indicating increased androgenization and possibly higher incidence of polygyny. Although speculative owing to small sample sizes, these results suggest that digit ratios represent a supplementary approach for elucidating the social systems of fossil hominins.     

Ecotypic variation in the context of global climate change: revisiting the rules

Patterns of ecotypic variation constitute some of the few 'rules' known to modern biology. Here, we examine several well-known ecogeographical rules, especially those pertaining to body size in contemporary, historical and fossil taxa. We review the evidence showing that rules of geographical variation in response to variation in the local environment can also apply to morphological changes through time in response to climate change. These rules hold at various time scales, ranging from contemporary to geological time scales. Patterns of body size variation in response to climate change at the individual species level may also be detected at the community level. The patterns underlying ecotypic variation are complex and highly context-dependent, reducing the 'predictive-power' of ecogeographical rules. This is especially true when considering the increasing impact of human activities on the environment. Nonetheless, ecogeographical rules may help interpret the likely influences of anthropogenic climate change on ecosystems. Global climate change has already influenced the body size of several contemporary species, and will likely have an even greater impact on animal communities in the future. For this reason, we highlight and emphasise the importance of museum specimens and the continued need for documenting the earth's biological diversity.     

New fossil records of Tapirus (Mammalia, Perissodactyla) from Brazil, with a critical analysis of intra-generic diversity assessments based on lower molar size variability

A large set of South American fossils belonging to the genus Tapirus has been described on the basis of differences in size and proportions of lower molariform teeth. Nevertheless, the reliability of dental proportions for the diagnosis of fossil tapir species is controversial. In this paper, we describe new fossil material of Tapirus from the Quaternary of Serra da Bodoquena, Southwestern Brazil, comparing it to other fossil and extant specimens of the genus by means of multivariate morphometric analyses of lower molariform teeth linear dimensions. The results of Principal Component Analyses indicate that some of the extant and extinct material attributed to Tapirus fall within the range of variation in size and proportions of lower molariform teeth exhibited by recent species of the genus. Therefore, part of the fossil material attributed to new species or to Tapirus sp. may be referable to the extant species Tapirus terrestris. We conclude that the sole use of lower molariform teeth size and proportions to erect new species of Tapirus may not be reliable, and therefore we advocate caution when describing fossil tapirs exclusively based on these criteria.     

Ancient DNA from pollen: a genetic record of population history in Scots pine

Assessments of plant population dynamics in space and time have depended on dated records of fossil pollen synthesized on a subcontinental scale. Genetic analyses of extant populations have revealed spatial relationships that are indicative of past spatial dynamics, but lack an explicit timescale. Synthesis of these data requires genetic analyses from abundant dated fossil material, and this has hitherto been lacking. Fossil pollen is the most abundant material with which to fill this data gap. Here we report genetic analyses of fossil pollen retrieved from Holtjärnen postglacial lake sediment in Sweden and show that plastid DNA is recoverable from Scots Pine and Norway spruce pollen grains that are 100 and 10 000 years old. By sequencing clones from two short plastid PCR products and by using multiple controls we show that the ancient sequences were endogenous to the fossil grains. Comparison of ancient sequences and those obtained from an extant population of Scots pine establishes the first genetic link between extant and fossil samples in this species, providing genetic continuity through time. The finding of one common haplotype present in modern, 100-year old and 10 000-year old samples suggests that it may have persisted near Holtjärnen throughout the postglacial period. This retrieval of ancient DNA from pollen has major implications for plant palaeoecology in conifer species by allowing direct estimates of population dynamics in space and time.     

Microscopic,chemical and molecular methods for examining fossil preservation

Advances in technology over the past two decades have resulted in unprecedented access to data from biological specimens. These data have expanded our understanding of physical characteristics, physiological, cellular and subcellular processes, and evolutionary relationships at the molecular level and beyond. Paleontological and archaeological sciences have recently begun to apply these technologies to fossil and subfossil representatives of extinct organisms. Data derived from multidisciplinary, non-traditional techniques can be difficult to decipher, and without a basic understanding of the type of information provided by these methods, their usefulness for fossil studies may be overlooked. This review describes some of these powerful new analytical tools, the data that may be accessible through their use, advantages and limitations, and how they can be applied to fossil material to elucidate characteristics of extinct organisms and their paleoecological environments.     

THE MACROEVOLUTIONARY CONSEQUENCES OF ECOLOGICAL DIFFERENCES AMONG SPECIES

Abstract:  In this paper, I examine the dynamics of species richness in a model system in which multiple species compete in a metacommunity (multiple patches linked by dispersal). Patches lie along an environmental gradient, and new species are introduced into the system by speciation of existing species. This model is used to explore how the ecological similarity of species influences the patterns in community structure that result and to determine whether patterns in fossil and systematics data may be signatures for different types of community structure. Making species more similar overall along the entire gradient or making new species that have more similar optimal positions along the gradient to their progenitor both increase the time required to drive species extinction. As a result, making species more similar ecologically to one another increases overall species richness because of an increased frequency of transient species in the system. Having more transient species in systems shifted the longevity distributions of species in the fossil record towards having a greater frequency of shorter duration species, and the age distribution of extant species that would be estimated from molecular phylogenies also had a higher frequency of younger aged species. Comparisons of these results with species longevity distributions extracted from two data sets and with species ages derived from species-level molecular phylogenies strongly suggest that transient species are an important component of real biological communities.     

Geometric morphometric analyses of worn cheek teeth help identify extant and extinct gophers (Rodentia,Geomyidae)

Studies of the biostratigraphy and palaeoecology of fossil vertebrate assemblages require large samples of accurately identified specimens. Such analyses can be hampered by the inability to assign isolated and worn remains to specific taxa. Entoptychine gophers are a diverse group of burrowing rodents found in Oligo‐Miocene deposits of the western United States. In both entoptychines and their extant relatives the geomyines, diagnostic characters of the occlusal surface of the teeth are modified with wear, making difficult the identification of many isolated fossil teeth. We use geometric morphometrics to test the hypothesis that tooth shape informs taxonomic affinities and expected levels of morphological variation across gopher taxa. We also incorporate data from microcomputer tomography to investigate changes in occlusal surface shape through wear within individuals. Our analyses demonstrate the usefulness of our approach in identifying extant geomyines to the genus, subgenus and species levels, and fossil entoptychines to the genus and, in some cases, the species level. Our results cast doubt on the validity of some species within Entoptychus and suggest future revisions to entoptychine taxonomy. The amounts of morphological divergence observed among fossil and extant genera are similar. Fossil species do not differ greatly from extant ones in that regard either. Further work evaluating the morphological variation within and across entoptychine species, including unworn teeth and osteological material, will allow revised analyses of the biostratigraphy and palaeoecology of important Oligo‐Miocene mammalian assemblages of the western United States and help to infer the phylogenetic relationships and evolution of gophers.     

Using extant patterns of dental variation to identify species in the primate fossil record: a case study of middle Eocene <Emphasis Type="Italic">Omomys</Emphasis> from the Bridger Basin,southwestern Wyoming

Patterns of extant primate dental variation provide important data for interpreting taxonomic boundaries in fossil forms. Here I use dental data from several well-known living primates (as well as data from selected Eocene forms) to evaluate dental variation in Middle Eocene Omomys, the first North American fossil primate identified by paleontologists. Measurements were collected from a sample of 148 omomyid dental specimens recovered from Bridger B localities in the Bridger Basin, Wyoming. Most of these specimens have not previously been described. Nonmetric traits were also scored for this sample. Lower molar coefficients of variation range from 4.01 for M₂ length (n = 80) to 6.73 for M₃ talonid width (n = 57). All of the nonmetric traits scored exhibit less than 100% presence in the overall sample, including traits previously described as representative of Omomys (e.g., P₄ metaconids present in 91%, n = 55; M² pericones present in 80%, n = 15). Dental traits also vary in a set of spatially restricted localities from the same fossil horizon and in a separate, single fossil locality (DMNH 868, P₄ metaconids present in 67%, n = 6). An increasing frequency in several premolar traits across time in these more restricted samples suggests an anagenetic change in Bridger B Omomys. However, this degree of morphological variability is consistent with that seen in extant primate species from single locations. Metric variation in this sample is comparable to that seen in other Eocene primates, such as new data presented here for the omomyid Arapahovius gazini from the Washakie Basin, southern Wyoming. Omomys metric variation is also comparable to that found in several samples of well-known extant primates from single localities (e.g., ring-tailed lemurs and gray–brown mouse lemurs). These metric data also correspond to the patterns of variability described in previously published studies of Omomys carteri. In sum, a single species interpretation (O. carteri) for this new Bridger B Omomys sample from southern Wyoming is affirmed, and this study illustrates the usefulness of dental data from extant primates for evaluating primate fossil samples.     

Geophilomorph centipedes from the Cretaceous amber of Burma

The only previously known Mesozoic fossils of the chilopod order Geophilomorpha are two species from the Late Jurassic and Late Cretaceous, both known from single specimens that cannot be assigned with precision to a family. Four specimens from the Late Cretaceous (earliest Cenomanian) amber of Burma include three that can be identified as conspecific, described here as Kachinophilus pereirai gen. nov. sp. nov. These specimens preserve greater morphological detail in comparison with other fossil geophilomorphs: the form and fine features of the head, the maxillary complex, the trunk sternites with associated glandular pores and the ultimate pair of legs defend the assignment of the species to the extant family Geophilidae, and most probably to a derived subgroup including well‐known extant genera such as Ribautia Brölemann, 1909. Confocal laser scanning microscopy supplements examination under incident and transmitted light to document details of high taxonomic relevance in the head and the forcipular segment. The modern appearance of this species and its membership among deeply nested extant clades are consistent with molecular estimates that most of the diversity of crown‐group Geophilomorpha originated before the Late Cretaceous.     

The evolutionary consequences of oxygenic photosynthesis: a body size perspective

The high concentration of molecular oxygen in Earth??s atmosphere is arguably the most conspicuous and geologically important signature of life. Earth??s early atmosphere lacked oxygen; accumulation began after the evolution of oxygenic photosynthesis in cyanobacteria around 3.0?C2.5 billion years ago (Gya). Concentrations of oxygen have since varied, first reaching near-modern values ~600 million years ago (Mya). These fluctuations have been hypothesized to constrain many biological patterns, among them the evolution of body size. Here, we review the state of knowledge relating oxygen availability to body size. Laboratory studies increasingly illuminate the mechanisms by which organisms can adapt physiologically to the variation in oxygen availability, but the extent to which these findings can be extrapolated to evolutionary timescales remains poorly understood. Experiments confirm that animal size is limited by experimental hypoxia, but show that plant vegetative growth is enhanced due to reduced photorespiration at lower O₂:CO₂. Field studies of size distributions across extant higher taxa and individual species in the modern provide qualitative support for a correlation between animal and protist size and oxygen availability, but few allow prediction of maximum or mean size from oxygen concentrations in unstudied regions. There is qualitative support for a link between oxygen availability and body size from the fossil record of protists and animals, but there have been few quantitative analyses confirming or refuting this impression. As oxygen transport limits the thickness or volume-to-surface area ratio??rather than mass or volume??predictions of maximum possible size cannot be constructed simply from metabolic rate and oxygen availability. Thus, it remains difficult to confirm that the largest representatives of fossil or living taxa are limited by oxygen transport rather than other factors. Despite the challenges of integrating findings from experiments on model organisms, comparative observations across living species, and fossil specimens spanning millions to billions of years, numerous tractable avenues of research could greatly improve quantitative constraints on the role of oxygen in the macroevolutionary history of organismal size.     

Craniodental variation in Paranthropus boisei: a developmental and functional perspective

What levels and patterns of craniodental variation among a fossil hypodigm are necessary to reject the null hypothesis that only a single species is sampled? We suggest how developmental and functional criteria can be used to predict where in the skeleton of fossil hominins we should expect more, or less, within-species variation. We present and test three hypotheses about the factors contributing to craniodental variation in extant primate taxa, and then apply these results to the interpretation of the P. boisei hypodigm. Within the comparative samples of extant Homo, Pan, Gorilla, Pongo, and Colobus, variables from the cranial base, neurocranium, and face that are not subject to high magnitudes of strain have consistently lower levels of intraspecific variation than variables from regions of the face subject to high levels of strain. Dental size variables are intermediate in terms of their reliability. P. boisei is found to have a low degree of variability relative to extant primates for variables shown to be generally useful for testing taxonomic hypotheses. Contrary to the claims of Suwa et al. ([1997] Nature 389:489-492), the recently discovered material from Konso falls within the range of variation of the "pre-Konso" hypodigm of P. boisei for available conventional metrical variables. Those aspects of the Konso material that appear to extend the range of the P. boisei hypodigm involve regions of the skull predicted to be prone to high levels of within-species variation. The approach used in this study focuses on craniodental data, but it is applicable to other regions of the skeleton.     

Mandibular size and shape variation in the hominins at Dmanisi, Republic of Georgia

The hominin fossils of Dmanisi, Republic of Georgia, present an ideal means of assessing levels of skeletal size and shape variation in a fossil hypodigm belonging to the genus Homo because they have been recovered from a spatially and temporally restricted context. We compare variation in mandible size and shape at Dmanisi to that of extant hominoids and extinct hominins. We use height and breadth measurements of the mandibular corpus at the first molar and the symphysis to assess size, and analyze shape based on size-adjusted (using a geometric mean) versions of these four variables. We compare size and shape variation at Dmanisi relative to all possible pairs of individuals within each comparative taxon using an exact resampling procedure of the ratio of D2600 to D211 and the average Euclidean distance (AED) between D2600 and D211, respectively. Comparisons to extant hominoids were conducted at both the specific and subspecific taxonomic levels and to extinct hominins by adopting both a more, and less speciose, hominin taxonomy. Results indicate that the pattern of variation for the Dmanisi hominins does not resemble that of any living species: they exhibit significantly more size variation when compared to modern humans, and they have significantly more corpus shape variation and size variation in corpus heights and overall mandible size than any extant ape species. When compared to fossil hominins they are also more dimorphic in size (although this result is influenced by the taxonomic hypothesis applied to the hominin fossil record). These results highlight the need to re-examine expectations of levels of sexual dimorphism in members of the genus Homo and to account for marked size and shape variation between D2600 and D211 under the prevailing view of a single hominin species at Dmanisi.