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That awkward age for butterflies: insights from the age of the butterfly subfamily Nymphalinae (Lepidoptera: Nymphalidae).

The study of the historical biogeography of butterflies has been hampered by a lack of well-resolved phylogenies and a good estimate of the temporal span over which butterflies have evolved. Recently there has been surge of phylogenetic hypotheses for various butterfly groups, but estimating ages of divergence is still in its infancy for this group of insects. The main problem has been the sparse fossil record for butterflies. In this study I have used a surprisingly good fossil record for the subfamily Nymphalinae (Lepidoptera: Nymphalidae) to estimate the ages of diversification of major lineages using Bayesian relaxed clock methods. I have investigated the effects of varying priors on posterior estimates in the analyses. For this data set, it is clear that the prior of the rate of molecular evolution at the ingroup node had the largest effect on the results. Taking this into account, I have been able to arrive at a plausible history of lineage splits, which appears to be correlated with known paleogeological events. The subfamily appears to have diversified soon after the K/T event about 65 million years ago. Several splits are coincident with major paleogeological events, such as the connection of the African and Asian continents about 21 million years ago and the presence of a peninsula of land connecting the current Greater Antilles to the South American continent 35 to 33 million years ago. My results suggest that the age of Nymphalidae is older than the 70 million years speculated to be the age of butterflies as a whole.

Animals↗

A lamprey from the Cretaceous Jehol biota of China.

Widespread nowadays in freshwater and coastal seas of the cold and temporal zones, lampreys are a jawless vertebrate group that has been in existence for more than 300 million years but left a meagre fossil record. Only two fossil lamprey species, namely Mayomyzon pieckoensis and Hardistiella montanensis, have been recognized with certainty from North American Carboniferous marine deposits. Here we report a freshwater lamprey from the Early Cretaceous epoch (about 125 million years ago) of Inner Mongolia, China. The new taxon, Mesomyzon mengae, has a long snout, a well-developed sucking oral disk, a relatively long branchial apparatus showing branchial basket, seven gill pouches, gill arches and impressions of gill filaments, about 80 myomeres and several other characters that are previously unknown or ambiguous. Our finding not only indicates Mesomyzon's closer relationship to extant lampreys but also reveals the group's invasion into a freshwater environment no later than the Early Cretaceous. The new material furthers our understanding of ancient lampreys, bridges the gap between the Carboniferous ones and their recent relatives, and adds to our knowledge of the evolutionary history of lampreys.

Animals↗

Fossilized soft tissues in a Silurian platyceratid gastropod.

Gastropod shells are common in the fossil record, but their fossil soft tissues are almost unknown, and have not been reported previously from the Palaeozoic. Here, we describe a Silurian (approx. 425 Myr) platyceratid gastropod from the Herefordshire Lagerstätte that preserves the oldest soft tissues yet reported from an undoubted crown-group mollusc. The digestive system is preserved in detail, and morphological data on the gonads, digestive gland, pedal muscle, radula, mouth and foot are also available. The specimen is preserved three-dimensionally, and has been reconstructed digitally following serial grinding. Platyceratids are often found attached to echinoderms, and have been interpreted as either commensal coprophages or kleptoparasites. The new data provide support for an attached mode of life, and are suggestive of a coprophagous feeding strategy. The affinities of the platyceratids are uncertain; they have been compared to both the patellogastropods and the neritopsines. Analysis of the new material suggests that a patellogastropod affinity is the more plausible of these hypotheses.

Animals↗

On the Pleistocene extinctions of Alaskan mammoths and horses.

The fossil record has been used to shed light on the late Pleistocene megafaunal extinctions in North America and elsewhere. It is therefore important to account for variability due to the incompleteness of the fossil record and error in dating fossil remains. Here, a joint confidence region for the extinction times of horses and mammoths in Alaska is constructed. The results suggest that a prior claim that the extinction of horses preceded the arrival of humans cannot be made with confidence.

Alaska↗

Fossil-calibrated molecular phylogenies reveal that leaf-mining moths radiated millions of years after their host plants.

Coevolution has been hypothesized as the main driving force for the remarkable diversity of insect-plant associations. Dating of insect and plant phylogenies allows us to test coevolutionary hypotheses and distinguish between the contemporaneous radiation of interacting lineages vs. insect 'host tracking' of previously diversified plants. Here, we used nuclear DNA to reconstruct a molecular phylogeny for 100 species of Phyllonorycter leaf-mining moths and 36 outgroup taxa. Ages for nodes in the moth phylogeny were estimated using a combination of a penalized likelihood method and a Bayesian approach, which takes into account phylogenetic uncertainty. To convert the relative ages of the moths into dates, we used an absolute calibration point from the fossil record. The age estimates of (a selection of) moth clades were then compared with fossil-based age estimates of their host plants. Our results show that the principal radiation of Phyllonorycter leaf-mining moths occurred well after the main radiation of their host plants and may represent the dominant associational mode in the fossil record.

Animals↗

A chicken-sized crane precursor from the early Oligocene of France.

A skeleton of a chicken-sized crane precursor is described from the Lower Oligocene of the Lubéron in Southern France. Parvigrus pohli gen. et sp. nov. is the most substantial Paleogene fossil record of the Grues (Aramidae [limpkin] + Gruidae [cranes]), and among its oldest representatives. The fossil species is classified in the new taxon Parvigruidae, which is shown to be the sister group of extant Grues. It is the first fossil record of a stem lineage representative of the Grues and, among others, differs from modern Grues in its smaller size, shorter beak, and rail-like limb proportions. Size increase in the stem lineage of the Gruidae may be related to the spread of grasslands during the Oligocene and Miocene. Occurrence of stem lineage Grues in the Lower Oligocene of Europe is in concordance with the fact that there is no evidence for the presence of crown group members of modern avian "families" in pre-Oligocene fossil deposits.

Animals↗

Metazoan phylogenies: falling into place or falling to pieces? A palaeontological perspective.

Metazoan phylogeny is in a state of ferment, stirred by the addition of new molecular trees as well as controversial interpretations of molecular 'clocks'. Concerning the latter topic, the clocks recurrently point to divergence times substantially older than the known fossil record. Some attempt reconciliation by appealing to a conveniently cryptic interval prior to the first fossils. This effectively reduces the fossil record to an erratic search-light giving only glimpses into the true evolutionary history. Other options, however, remain open. Molecular clocks may themselves run erratically and what happens in molecular history may not coincide with the emergence of body plans.

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Counting dinosaurs: how many kinds were there?

Dinosaurs figure prominently in discussions of mass extinctions and evolutionary metrics, but their usefulness is hampered by archaic taxonomy, imprecise biostratigraphy, and imperfect preservation that bias our understanding of dinosaur diversity. A critical evaluation shows that of 540 genera and 800 species of dinosaurs proposed since 1824, 285 genera and 336 species are probably valid. Nearly half of all genera are based on a single specimen, and complete skulls and skeletons are known for only 20% of all dinosaurs. Dinosaurs are known from every continent. Countries with the greatest known diversity of dinosaurs are (in descending order) the United States, Mongolia, China, Canada, England, and Argentina; the greatest future increases may be expected from Argentina and China. Nearly half of all dinosaur genera are of latest Cretaceous age (Campanian or Maastrichtian). Estimates of the average duration of a dinosaur genus range from 5 million to 10.5 million years, with the most likely value about 7.7 million years. Dinosaurs evolved as rapidly as Cenozoic mammals. Global dinosaur diversity during the Campanian and Maastrichtian is estimated at 100 genera per stage, using a logistic model to estimate future discoveries. A model of increasing diversity and a bottleneck model compensate for the biasis in the preserved fossil record. The number of dinosaurs that have ever lived is estimated at 900-1200 genera. The fossil record of dinosaurs is presently about 25% complete. Dinosaurs disappeared in the Maastrichtian near the peak of their historic diversity.

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The relative rate of DNA evolution in primates.

In 73 relative-rate tests involving the sequences of 17 genes between humans and six nonhuman primate taxa, there is only one significant (P less than 0.01) difference in evolutionary rate--i.e., that between human and Old World-monkey psi eta-globin genes. No evolutionary rate difference between humans and Old World monkeys is evident from analysis of 18 other genes with a total length of 6 kb. This and the comparison, between humans and other primate taxa, of new extended psi eta-globin sequences suggest that earlier observations of evolutionary-rate differences between humans and other primates were based on differences that are peculiar to psi eta-globin and that are not representative of the whole genome, which appears to be evolving at a stochastically uniform rate. This is supported by whole-genome single-copy DNA and mitochondrial DNA comparisons, neither of which shows any evidence of evolutionary-rate variation among primate taxa. Uniformity in the evolutionary rate of the DNA of primate and other mammalian taxa is inconsistent with current mammalian fossil-record interpretation. Either there has been a general slowing down in rate across lineages or the fossil record has been misinterpreted.

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Spine anatomy reveals the diversity of catfish through time: a case study of Synodontis (Siluriformes).

Synodontis (Mochokidae, Siluriformes) is a freshwater catfish endemic to Africa. The 118 extant species are present in almost all hydrographic basins. Some species are restricted to a single stream, whereas others have a vast distribution. Synodontis is known in the fossil record since the Miocene, and its history depends on the connections among African basins through time. The identification of species in the fossil record is essential to reconstruct this historical pattern. Catfish pectoral and dorsal spines are robust, they preserve well and they form most of the fossil remains for the genus Synodontis. Unfortunately, the criteria for the identification of extant Synodontis species are not applicable to fossil specimens. Here, we define 11 original morphological characters that permit to discriminate four extant species from the Chad-Chari hydrographic system. Six of these characters are defined on pectoral spines and five on dorsal spines. We then show that these characters can be used successfully for identifying fossil specimens. In particular, we present a case study in which we identify Synodontis cf. schall and Brachysynodontis cf. batensoda in the hominid-bearing sector Toros-Menalla (Late Miocene, northern Chad). We show that spine anatomy can be a powerful tool to recognise catfish species through time and thus to identify historical diversity pattern.

Africa↗

The diversification of Paleozoic fire systems and fluctuations in atmospheric oxygen concentration.

By comparing Silurian through end Permian [approximately 250 million years (Myr)] charcoal abundance with contemporaneous macroecological changes in vegetation and climate we aim to demonstrate that long-term variations in fire occurrence and fire system diversification are related to fluctuations in Late Paleozoic atmospheric oxygen concentration. Charcoal, a proxy for fire, occurs in the fossil record from the Late Silurian (approximately 420 Myr) to the present. Its presence at any interval in the fossil record is already taken to constrain atmospheric oxygen within the range of 13% to 35% (the "fire window"). Herein, we observe that, as predicted, atmospheric oxygen levels rise from approximately 13% in the Late Devonian to approximately 30% in the Late Permian so, too, fires progressively occur in an increasing diversity of ecosystems. Sequentially, data of note include: the occurrence of charcoal in the Late Silurian/Early Devonian, indicating the burning of a diminutive, dominantly rhyniophytoid vegetation; an apparent paucity of charcoal in the Middle to Late Devonian that coincides with a predicted atmospheric oxygen low; and the subsequent diversification of fire systems throughout the remainder of the Late Paleozoic. First, fires become widespread during the Early Mississippian, they then become commonplace in mire systems in the Middle Mississippian; in the Pennsylvanian they are first recorded in upland settings and finally, based on coal petrology, become extremely important in many Permian mire settings. These trends conform well to changes in atmospheric oxygen concentration, as predicted by modeling, and indicate oxygen levels are a significant control on long-term fire occurrence.

Air↗

Cranial remains of an Eocene tarsier.

The phylogenetic position of tarsiers relative to anthropoids and Paleogene omomyids remains a subject of lively debate that lies at the center of research into anthropoid origins. Omomyids have long been regarded as the nearest relatives of tarsiers, but a sister group relationship between anthropoids and tarsiers has also been proposed. These conflicting phylogenetic reconstructions rely heavily on comparisons of cranial anatomy, but until now, the fossil record of tarsiers has been limited to a single jaw and several isolated teeth. In this article, we describe cranial material of a fossil tarsiid from the middle-Eocene Shanghuang fissure-fillings in southern Jiangsu Province, China. This facial fragment, which is allocated to Tarsius eocaenus, is virtually identical to the corresponding anatomy in living tarsiers and differs substantially from that of early anthropoids such as Bahinia, Phenacopithecus, and Parapithecus. This new specimen indicates that tarsiers already possessed greatly enlarged orbits and a haplorhine oronasal configuration by the time they are first documented in the fossil record during the middle Eocene.

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Morphological and developmental macroevolution: a paleontological perspective.

Evidence of the morphological evolution of metazoans has been preserved, in varying degrees of completeness, in the fossil record of the last 600 million years. Although extinction has been incessant at lower taxonomic levels, genomic comparisons among surviving members of higher taxa suggest that much of the developmental systems that pattern their bodyplans has been conserved from early in their history. Comparisons between the origin of morphological disparity in the record and patterns of genomic disparity among living taxa promise to be interesting. For example, Hox cluster composition varies among major taxa, and the fossil record suggests that many of the changes in Hox clusters may have been associated with late Neoproterozoic evolution among minute benthic vermiform clades, from which crown bilaterian phyla arose just before or during the Cambrian explosion. Study of genomic differences among crown classes and orders whosetiming and mode of origin can be inferred from morphological data inthefossil record should throw further light on the timing and mode of origin of genomic disparities.

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Placental mammal diversification and the Cretaceous-Tertiary boundary.

Competing hypotheses for the timing of the placental mammal radiation focus on whether extant placental orders originated and diversified before or after the Cretaceous-Tertiary (KT) boundary. Molecular studies that have addressed this issue suffer from single calibration points, unwarranted assumptions about the molecular clock, andor taxon sampling that lacks representatives of all placental orders. We investigated this problem using the largest available molecular data set for placental mammals, which includes segments of 19 nuclear and three mitochondrial genes for representatives of all extant placental orders. We used the ThorneKishino method, which permits simultaneous constraints from the fossil record and allows rates of molecular evolution to vary on different branches of a phylogenetic tree. Analyses that used different sets of fossil constraints, different priors for the base of Placentalia, and different data partitions all support interordinal divergences in the Cretaceous followed by intraordinal diversification mostly after the KT boundary. Four placental orders show intraordinal diversification that predates the KT boundary, but only by an average of 10 million years. In contrast to some molecular studies that date the rat-mouse split as old as 46 million years, our results show improved agreement with the fossil record and place this split at 16-23 million years. To test the hypothesis that molecular estimates of Cretaceous divergence times are an artifact of increased body size subsequent to the KT boundary, we also performed analyses with a "KT body size" taxon set. In these analyses, interordinal splits remained in the Cretaceous.

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Solution to Darwin's dilemma: discovery of the missing Precambrian record of life.

In 1859, in On the Origin of Species, Darwin broached what he regarded to be the most vexing problem facing his theory of evolution-the lack of a rich fossil record predating the rise of shelly invertebrates that marks the beginning of the Cambrian Period of geologic time ( approximately 550 million years ago), an "inexplicable" absence that could be "truly urged as a valid argument" against his all embracing synthesis. For more than 100 years, the "missing Precambrian history of life" stood out as one of the greatest unsolved mysteries in natural science. But in recent decades, understanding of life's history has changed markedly as the documented fossil record has been extended seven-fold to some 3,500 million years ago, an age more than three-quarters that of the planet itself. This long-sought solution to Darwin's dilemma was set in motion by a small vanguard of workers who blazed the trail in the 1950s and 1960s, just as their course was charted by a few pioneering pathfinders of the previous century, a history of bold pronouncements, dashed dreams, search, and final discovery.

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The age of the angiosperms: a molecular timescale without a clock.

The age of the angiosperms has long been of interest to botanists and evolutionary biologists. Many early efforts to date the age of the angiosperms and evolutionary divergences within the angiosperm clade using a molecular clock have yielded age estimates that are grossly inconsistent with the fossil record. We investigated the age of angiosperms using Bayesian relaxed clock (BRC) and penalized likelihood (PL) approaches. Both of these methods allow the incorporation of multiple fossil constraints into the optimization procedure. The BRC method allows a range of values for among-lineage rate of substitution, from a nearly clocklike behavior to a condition in which each branch is allowed an optimal substitution rate, and also accounts for variation in molecular evolution across multiple genes. A topology derived from an analysis of genes from all three plant genomes for 71 taxa was used as a backbone. The effects on age estimates of different genes, single-gene versus concatenated datasets, and the inclusion and assumptions of fossils as age constraints were examined. In addition, the influence of prior distributions on estimates of divergence times was also explored. These results indicate that widely divergent age estimates can result from the different methods (198-139 million years ago), different sources of data (275-122 million years ago), and the inclusion of temporal constraints to topologies. Most dates, however, are between 180-140 million years ago, suggesting a Middle Jurassic-Early Cretaceous origin of flowering plants, predating the oldest unequivocal fossil angiosperms by about 45-5 million years. Nonetheless, these dates are consistent with other recent studies that have used methods that relax the assumption of a strict molecular clock and also agree with the hypothesis that the angiosperms may be somewhat older than the fossil record indicates.

Bayes Theorem↗

First fossil lamprey: a record from the Pennsylvanian of Illinois.

A fossil record of lampreys has previously been unknown. A new genus demonstrates the presence of this group in the Pennsylvanian. The body outline, parts of the head skeleton, rasping tongue mechanism, gill basket, and other internal organs are preserved. The fossils are very similar in structure to modern forms. The absence of hagfish characters in the fossil supports the view that the common ancestor of lampreys and hagfishes lived prior to the Pennsylvanian.

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