Quaternary phylogeography: the roots of hybrid zones

The older history of hybrid zones is explored through consideration of recent advances in climatology, paleontology and phylogeography in the Late Cenozoic, particularly the Quaternary Period with its major climatic cycles. The fossil record shows that these ice ages and their nested millennial oscillations caused substantial changes in species distributions and with genetic evidence allows deduction of refugia and colonization routes in arctic, temperate, desert and tropical regions. The age of divergence between hybridizing lineages varies from the Late Pleistocene to the Late Miocene, implying much range change and varying selection on sister lineages. Hybridizing lineages in the Tropical and Temperate regions range in age from young to old, but those studied in the Arctic are no more than a few ice ages old and their refugial roots are not clear. Mid to low latitude regions often show parapatric patchworks of lineages and multiple refugia stable through many climatic oscillations. Particular hybrid zones may have formed more than once; while some expansions were not the same, producing reticulation and introgression in previous glacial cycles. Hybrid-zone roots are complex and deep, and considerations of their complexity can reveal evolutionary pathways of species. They are indeed windows on evolution.     

Mid-Tertiary elapid snakes (Squamata, Colubroidea) from Riversleigh, northern Australia: early steps in a continent-wide adaptive radiation

Vertebral and cranial remains of elapid snakes have been collected from fossil assemblages at Riversleigh, north-west Queensland, Australia; most are Miocene but one may be late Oligocene and another as young as Pliocene. The oldest specimen (probably the oldest elapid yet known anywhere) is a vertebra that can be referred provisionally to the extant taxon Laticauda (Hydrophiinae, sensu Slowinski and Keogh, 2000), implying that the basal divergences among Australasian hydrophiine lineages had occurred by the early Miocene, in contrast to most previous estimates for the age of this geographically isolated adaptive radiation. Associated vertebrae and jaw elements from a Late Miocene deposit are described as Incongruelaps iteratus nov. gen. et sp., which has a unique combination of unusual derived characters otherwise found separately in several extant hydrophiine taxa that are only distantly related. Associated vertebrae from other sites, and two parietals from a possibly Pliocene deposit, suggest the presence of several other taxa distinct from extant forms, but the amount of material (and knowledge of variation in extant taxa) is currently insufficient to diagnose these forms. The Tertiary elapids of Riversleigh thus appear to be relatively diverse taxonomically, but low in abundance and, with one exception, not referable to extant taxa below the level of Hydrophiinae. This implies that the present diversity of hydrophiine elapids (31 recognized terrestrial genera, and approximately 16 marine) represents the result of substantial extinction as well as the “cone of increasing diversity” that could be inferred from phylogenetic studies on extant forms.     

Hammer-toothed 'marsupial skinks' from the Australian Cenozoic

Extinct species of Malleodectes gen. nov. from Middle to Late Miocene deposits of the Riversleigh World Heritage Area, northwestern Queensland, Australia are enigmatic, highly specialized, probably snail-eating marsupials. Dentally, they closely resemble a bizarre group of living heterodont, wet forest scincid lizards from Australia (Cyclodomorphus) that may well have outcompeted them as snail-eaters when the closed forests of central Australia began to decline. Although there are scincids known from the same Miocene deposits at Riversleigh, these are relatively plesiomorphic, generalized feeders. This appears to be the most striking example known of dental convergence and possible competition between a mammal and a lizard, which in the long run worked out better for the lizards.     

A systematic overview of fossil osmundalean ferns in China: Diversity variation,distribution pattern,and evolutionary implications

The order Osmundales is a unique fern taxon with extensive fossil records in geological past. Diverse osmundalean fossils have been reported from China, ranging in age from the Late Palaeozoic to the Cenozoic. Most of them are based on leaf impressions/compressions, but permineralized rhizomes are also well documented. In this study, we provide a systematic overview on fossil osmundalean ferns in China with special references on diversity variations, distribution patterns, and evolutionary implications. Fossil evidence indicates that this fern lineage first appeared in the Late Palaeozoic in China. The Late Triassic to Middle Jurassic interval was the radiation stage. From the Late Jurassic onward, fossil diversity declined rapidly. Cenozoic osmundalean taxa are represented by the relict species of Osmunda. Geographically, osmundalean fossils are found from both the Northern and Southern phytoprovinces of China, though variations are documented for geographical ranges. The Chinese fossil records cover almost all important stages for the macroevolution of the Osmundales, and contribute to further understanding of evolutionary processes of this peculiar fern lineage.     

Lack of stasis in late Cenozoic marine faunas and communities, central California

Pliocene strata in the Kettleman Hills of west-central California were deposited in the broad San Joaquin embayment as a cyclic succession of parasequences during approximately three million years. Depositional environments within each cycle ranged from relatively open marine to brackish and non-marine. Although the strata were deposited in similar, recurrent environments, the fauna changed gradually rather than during brief intervals separating periods of stasis. Although environmental gradients and community structure in the Pliocene San Joaquin Embayment and in the present-day San Francisco Bay are similar, species compositions of the faunas and of parallel communities at the two sites are markedly different. In addition, times of origination and extinction of species in the Pliocene strata of the Kettleman Hills and in San Francisco Bay do not document coordinated stasis within the Late Cenozoic. In contrast, Silurian and Devonian faunas and communities of the Appalachian Basin persisted with little change in ecological-evolutionary units that lasted for up to eight million years. Relatively brief intervals of great biotic change separate these intervals of stasis. One intriguing explanation of this pattern of coordinated stasis within an ecologic-evolutionary unit is ecological locking, in which interaction between species within the community is sufficiently strong that only major changes in the environment are able to change community and faunal composition. Probably the late Cenozoic fauna underwent rapid evolution as a result of rapidly changing environmental conditions within a complex and changing shallow, inshore marine paleogeography. In contrast, coordinated stasis in the lower Paleozoic probably resulted from negligible evolution during long periods of stable to gradually changing environments in an outer shelf setting, punctuated by brief episodes of abrupt environmental change and large-scale turnover. The independent assortment of species in late Cenozoic parallel communities indicates that ecological locking did not exist.     

Puycelci, nouveau site à vertébrés de la série molassique d’Aquitaine. Densité et continuité biochronologique dans la zone Quercy et bassins périphériques au Paléogène

After the previous lacking evidence allowed the interpretation of a non-deposition episod, the newly recorded mammal fauna from Puycelci (Tarn, SW France, close South to the Quercy paleokarstic area) now shows the MP 26 standard level (Late Oligocene, Early Chattian) being represented within the Eastern Aquitaine molassic basin filling. According to the evolutionary stage of known mammal lineages, the most significant Puycelci fossils are the Issiodoromys pauffiensis rodent and the Metriotherium cf. sarelense nov. sp. (described in this paper) artiodactyl ungulate. This new dating exemplifies the improvement of the mammalian biochronologic record as regards both the Quercy paleokarst and the surrounding lacustrine Tertiary basins W, S, and S-E to the Quercy, as well as, directly E and N-E, the smaller tectonic Tertiary basins within the crystalline Massif Central basement. The updated record shows the strong extension in time of the paleokarstic data, down to the Middle and Early Eocene, and up to the Early Miocene. Regarding the classically reported Late Eocene to Late Oligocene span, as well in the paleokarst and the peripheral basins, the available biochronological data now attains a valuable density level, the variation of which is significant. On one side there are periods (Late Eocene, Early Oligocene, Early Late Oligocene) with high biochronologic density, allowing improved time resolution using evolutionary stages within mammalian lineages (e.g. numerical ages). Such periods corresponds to almost continuous sedimentary processes, with only possible short breaks. On the other side, there are periods with unreported data, shared in the paleokarst and peripheral basins (Late Oligocene as best example). Such periods may well correspond to some non-deposition episods as once alleged by geologists. Then the surrounding biochronologic data would bear constraining value regarding these episods.     

Climate‐driven body‐size trends in the ostracod fauna of the deep Indian Ocean

Abstract: Body size is a common focus of macroevolutionary, macroecological and palaeontological investigations. Here, we document body‐size evolution in 19 species‐level ostracod lineages from the deep Indian Ocean (Deep Sea Drilling Program Site 253) over the past 40 myr. Body‐size trajectories vary across taxa and time intervals, but most lineages (16/19) show net gains in body size. Because many modern ostracod taxa are larger in colder parts of their geographical range, we compared the timing and magnitude of these size changes to established Cenozoic deep‐water cooling patterns confirmed through δ¹⁸O measurements of benthic foraminifera in the samples studied. These data show a significant negative correlation between size changes and temperature changes (ostracods get larger as temperatures get colder), and that systematic size increases only occur during intervals of sustained cooling. In addition, statistical support for an explicit temperature‐tracking model exceeds that of purely directional evolution. We argue that this Cope’s Rule pattern is driven by secular changes in the environment, rather than any universal or intrinsic advantages to larger body sizes, and we note some difficulties in the attempts to link Cope’s Rule to observations made within a single generation.     

Explosive radiation of Malpighiales supports a mid-cretaceous origin of modern tropical rain forests

Fossil data have been interpreted as indicating that Late Cretaceous tropical forests were open and dry adapted and that modern closed-canopy rain forest did not originate until after the Cretaceous-Tertiary (K/T) boundary. However, some mid-Cretaceous leaf floras have been interpreted as rain forest. Molecular divergence-time estimates within the clade Malpighiales, which constitute a large percentage of species in the shaded, shrub, and small tree layer in tropical rain forests worldwide, provide new tests of these hypotheses. We estimate that all 28 major lineages (i.e., traditionally recognized families) within this clade originated in tropical rain forest well before the Tertiary, mostly during the Albian and Cenomanian (112-94 Ma). Their rapid rise in the mid-Cretaceous may have resulted from the origin of adaptations to survive and reproduce under a closed forest canopy. This pattern may also be paralleled by other similarly diverse lineages and supports fossil indications that closed-canopy tropical rain forests existed well before the K/T boundary. This case illustrates that dated phylogenies can provide an important new source of evidence bearing on the timing of major environmental changes, which may be especially useful when fossil evidence is limited or controversial.     

Biogeography of Triassic tetrapods: evidence for provincialism and driven sympatric cladogenesis in the early evolution of modern tetrapod lineages

Triassic tetrapods are of key importance in understanding their evolutionary history, because several tetrapod clades, including most of their modern lineages, first appeared or experienced their initial evolutionary radiation during this Period. In order to test previous palaeobiogeographical hypotheses of Triassic tetrapod faunas, tree reconciliation analyses (TRA) were performed with the aim of recovering biogeographical patterns based on phylogenetic signals provided by a composite tree of Middle and Late Triassic tetrapods. The TRA found significant evidence for the presence of different palaeobiogeographical patterns during the analysed time spans. First, a Pangaean distribution is observed during the Middle Triassic, in which several cosmopolitan tetrapod groups are found. During the early Late Triassic a strongly palaeolatitudinally influenced pattern is recovered, with some tetrapod lineages restricted to palaeolatitudinal belts. During the latest Triassic, Gondwanan territories were more closely related to each other than to Laurasian ones, with a distinct tetrapod fauna at low palaeolatitudes. Finally, more than 75 per cent of the cladogenetic events recorded in the tetrapod phylogeny occurred as sympatric splits or within-area vicariance, indicating that evolutionary processes at the regional level were the main drivers in the radiation of Middle and Late Triassic tetrapods and the early evolution of several modern tetrapod lineages.     

Taking into account phylogenetic and divergence‐time uncertainty in a parametric biogeographical analysis of the Northern Hemisphere plant clade Caprifolieae

Aim To examine the biogeographical history of the angiosperm clade Caprifolieae (Caprifoliaceae) using parametric biogeographical reconstruction methods. The existing parametric method was extended to evaluate biogeographical reconstructions over the distribution of dated phylogenies. This method provides a framework for reconstructing large‐scale biogeography with parametric methods, while accounting for uncertainty in the phylogenetic relationships and divergence time. Location Asia, Europe and North America. Methods The biogeographical history of the major lineages of Caprifolieae was reconstructed over the posterior distribution of dated trees generated from Bayesian divergence‐time analyses. Results from a model with no geological constraints were compared with those from one where movement is disallowed across the North Atlantic after the Eocene. The most plausible scenarios were segregated at each node to test whether particular scenarios were reconstructed for particular divergence times. The parametric biogeographical method was also extended to estimate connectivity between areas so that the probability of dispersal between the major areas of the Northern Hemisphere could be explored. Results Phylogenetic results for Caprifolieae agreed with previous estimates using smaller sampling, but uncertainty remained despite efforts to resolve the relationships of the four genera within this clade using multiple markers. In addition to topological uncertainty, there were few fossils available for calibrations, resulting in large confidence intervals for divergence times. Divergence‐time analyses put the diversification of Caprifolieae at between 36 and 51 Ma and showed that Caprifolieae probably originated in Asia, with multiple movements into Europe and western and eastern North America. Main conclusions Newly developed parametric methods for biogeographical reconstruction incorporate more data and better models. Here, the parametric biogeographical reconstruction method has been extended to allow for topological and divergence‐time uncertainty. The analyses of Caprifolieae demonstrated that biogeographical hypotheses can be explored even where there are large confidence intervals on divergence times and uncertainty in topology. These results add to the growing evidence that Asia was an important source of Northern Hemisphere diversity throughout the Cenozoic.     

3D Morphometric Analysis Reveals Similar Ecomorphs for Early Kangaroos (Macropodidae) and Fanged Kangaroos (Balbaridae) from the Riversleigh World Heritage Area,Australia

Understanding feeding ecology of extinct kangaroos is fundamental to understanding the evolution of kangaroos and the Australia paleoenvironment during the Oligo-Miocene. Comparisons with extant species have suggested that the macropodiforms of the Oligo/Miocene (kangaroos and allies) from the Riversleigh World Heritage Area, northern Australia, were predominantly folivorous browsers or fungivores, unlike the majority of extant species. To further test this hypothesis, we investigate the relationship between variation in cranial and mandibular shape of extant and extinct macropodiforms and ecological factors such as diet, locomotion, and body mass using 3D geometric morphometric analysis of 42 living species and eight extinct species from two radiations (the extinct clade of Balbaridae and some early representatives of the extant Macropodidae. Dietary class (fungivore, browser, grazer, and mixed feeder) correlated strongly with variation in cranial shape (20–25% of variance explained). There was also significant association between cranial shape, and both locomotor mode and body mass. In a principal component analysis of shape variation for crania (including the shape of the molar row), Riversleigh macropodiforms cluster with extant folivorous browsers on principal components (PC) 1 and 3, providing support for previous interpretations of these species as browsing kangaroos. However, as a group and regardless of phylogenetic association, the shape centroid of extinct species differs significantly from that of extant species. Riversleigh macropodiforms cluster with regular hoppers or arboreal tree kangaroos, but this may be a result of the correlation between diet and locomotor mode in kangaroos. Their similarity to extant browsers supports previous interpretations of rainforest and woodland environments at Riversleigh during the early and middle Miocene, respectively. Procrustes ANOVA Analysis of the full shape dataset and diet also shows that diet accounts for a significant portion of variation; however, when phylogeny is taken into account these results become nonsignificant. In analyses of dentary shape, some balbarid species cluster with extant mixed feeders, although this may reflect phylogenetic differences rather than ecological signal.

     

Leaves and pollen of bamboos from the Polish Neogene

Recently discovered fossil bamboo leaves and pollen are described from the Neogene of Poland. Morphological analysis of fossil leaves from the Late Miocene deposits of the Bełchatów Lignite Mine, Central Poland have allowed them to be determined as “Bambusa” lugdunensis Saporta. It is the first record of this species in the Cenozoic of Poland. The fossils have been compared with other records of bamboos from the Cenozoic of Europe. Pollen grains from the Middle Miocene (Badenian) deposits from the Legnica brown coal deposit (Lower Silesia) provide additional evidence of bamboos. These are assigned to Graminidites bambusoides Stuchlik, and are similar to those of recent Arundinaria. We infer that plant (plants?) that produced the pollen of G. bambusoides could have grown in swamp forests and/or reed marshes. This report provides a general survey of the occurrence of bamboos in the European Neogene, from which the leaves and pollen documented here present important data on the distribution of this group of monocots in Poland and eastern Europe.     

Near Eastern Neolithic genetic input in a small oasis of the Egyptian Western Desert

The Egyptian Western Desert lies on an important geographic intersection between Africa and Asia. Genetic diversity of this region has been shaped, in part, by climatic changes in the Late Pleistocene and Holocene epochs marked by oscillating humid and arid periods. We present here a whole genome analysis of mitochondrial DNA (mtDNA) and high‐resolution molecular analysis of nonrecombining Y‐chromosomal (NRY) gene pools of a demographically small but autochthonous population from the Egyptian Western Desert oasis el‐Hayez. Notwithstanding signs of expected genetic drift, we still found clear genetic evidence of a strong Near Eastern input that can be dated into the Neolithic. This is revealed by high frequencies and high internal variability of several mtDNA lineages from haplogroup T. The whole genome sequencing strategy and molecular dating allowed us to detect the accumulation of local mtDNA diversity to 5,138 ± 3,633 YBP. Similarly, theY‐chromosome gene pool reveals high frequencies of the Near Eastern J1 and the North African E1b1b1b lineages, both generally known to have expanded within North Africa during the Neolithic. These results provide another piece of evidence of the relatively young population history of North Africa. Am J Phys Anthropol, 2009. © 2009 Wiley‐Liss, Inc.     

Molecular evidence for the age, origin, and evolutionary history of the American desert plant genus Tiquilia (Boraginaceae)

Although the deserts of North America are of very recent origin, their characteristic arid-adapted endemic plant lineages have been suggested to be much older. Earlier researchers have hypothesized that the ancestors of many of these modern desert lineages first adapted to aridity in highly localized arid or semi-arid sites as early as the late Cretaceous or early Tertiary, and that these lineages subsequently spread and diversified as global climate became increasingly arid during the Cenozoic. No study has explicitly examined these hypotheses for any North American arid-adapted plant group. The current paper tests these hypotheses using the genus Tiquilia (Boraginaceae), a diverse North American desert plant group. A strongly supported phylogeny of the genus is estimated using combined sequence data from three chloroplast markers (matK, ndhF, and rps16) and two nuclear markers (ITS and waxy). Ages of divergence events within the genus are estimated using penalized likelihood and a molecular clock approach on the ndhF tree for Tiquilia and representative outgroups, including most of the major lineages of Boraginales. The dating analysis suggests that the stem lineage of Tiquilia split from its nearest extant relative in the Paleocene or Eocene ( approximately 59-48 Ma). This was followed by a relatively long period before the first divergence in the crown group near the Eocene/Oligocene boundary ( approximately 33-29 Ma), shortly after the greatest Cenozoic episode of rapid aridification. Divergence of seven major lineages of Tiquilia is dated to the early-to-mid Miocene ( approximately 23-13 Ma). Several major lineages show a marked increase in diversification concomitant with the onset of more widespread semi-arid and then arid conditions beginning in the late Miocene ( approximately 7 Ma). This sequence of divergence events in Tiquilia agrees well with earlier researchers' ideas concerning North American desert flora assembly.     

A phylogeographic, demographic and historical analysis of the short-tailed pit viper (Gloydius brevicaudus): evidence for early divergence and late expansion during the Pleistocene

The impact of quaternary glaciation in eastern China on local fanua and flora has been a topic of considerable interest. We used mitochondrial DNA (mtDNA) sequence data and coalescent simulations to test two general biogeographic hypothesis related to the effects of Pleistocene climatic fluctuations for a widespread ophidian species (Gloydius brevicaudus) in eastern China and Korean Peninsula. The phylogenetic analysis revealed three major lineages, the southeast Coastal, Yangtze and North Lineages. The latter two are closely related and jointly form a continental lineage. Divergence dating and coalescent simulations indicate a Late Pliocene to Early Pleistocene divergence between lineages from the southeast coast and continental interior, followed by a mid-to-late Pleistocene divergence between lineages from the north and the middle-lower Yangtze Valley across East China, suggesting that all these lineages predated the last glacial maximum. An overlapping range between the two lineages within the continental lineage and a secondary contact associated with ecological transition zones on the margins of the North China Plain were also observed. These results show that vicariance patterns dominated the history of G. brevicaudus. Though the climatic events of the Pleistocene have had a marked effect on the historical distribution and intra-specific divergence of reptiles in China, coalescent and non-coalescent demographic analyses indicate that all lineages of G. brevicaudus seem not to have been adversely affected by glacial cycles during the Late Pleistocene, presumably because of an increase in the amount of climatically mild habitat in East Asia due to a decline in elevation and the development of monsoons since the Mid-End Pleistocene.     

Plenasium xiei sp. nov. from the Cretaceous of Northeast China: Additional evidence for the longevity of osmundaceous ferns

A new species of the Osmundaceae, Plenasium xiei sp. nov., is herein described from the Cretaceous of Northeast China. The specimens examined here represent the earliest unequivocal record of the extant genus Plenasium in Eurasia based on fossil rhizomes. The rhizome consists of a central stem with a mantle of petiole bases and adventitious roots. The stem contains an ectophloic‐dictyoxylic siphonostele and a two‐layered cortex. The C‐shaped leaf trace bears two protoxylem bundles at the point of separation from the stele. The pith is heterogeneous. The parenchymatous inner cortex is thinner than the sclerenchymatous outer cortex. Lobed sclerenchyma bands occur at the adaxial sides of the stem xylem strands, in the concavity of the leaf trace, and along the adaxial side of the vascular bundles of the petiole base. In distal petiole portions, the sclerenchyma band splits into several groups in the transverse view. Sclerenchyma rings are heterogeneous with an abaxial sclerenchymatous arc of thick‐walled fibers. Numerous sclerenchyma strands of thick‐walled fibers appear in the petiolar inner cortex and the stipular wing. These fossils provide unambiguous evidence for the existence of subgenus Plenasium of modern Plenasium by at least the Late Cretaceous, demonstrating the longevity of this extant subgenus. Altogether the leaf and rhizome fossil records of Plenasium indicate that this genus was widely distributed across North America and Eurasia from the Early Cretaceous to the Early Cenozoic, followed by a range restriction to Eurasia in the Late Cenozoic. Extant Plenasium species are only known from East and Southeast Asia.     

Permian–Triassic Osteichthyes (bony fishes): diversity dynamics and body size evolution

The Permian and Triassic were key time intervals in the history of life on Earth. Both periods are marked by a series of biotic crises including the most catastrophic of such events, the end‐Permian mass extinction, which eventually led to a major turnover from typical Palaeozoic faunas and floras to those that are emblematic for the Mesozoic and Cenozoic. Here we review patterns in Permian–Triassic bony fishes, a group whose evolutionary dynamics are understudied. Based on data from primary literature, we analyse changes in their taxonomic diversity and body size (as a proxy for trophic position) and explore their response to Permian–Triassic events. Diversity and body size are investigated separately for different groups of Osteichthyes (Dipnoi, Actinistia, ‘Palaeopterygii’, ‘Subholostei’, Holostei, Teleosteomorpha), within the marine and freshwater realms and on a global scale (total diversity) as well as across palaeolatitudinal belts. Diversity is also measured for different palaeogeographical provinces. Our results suggest a general trend from low osteichthyan diversity in the Permian to higher levels in the Triassic. Diversity dynamics in the Permian are marked by a decline in freshwater taxa during the Cisuralian. An extinction event during the end‐Guadalupian crisis is not evident from our data, but ‘palaeopterygians’ experienced a significant body size increase across the Guadalupian–Lopingian boundary and these fishes upheld their position as large, top predators from the Late Permian to the Late Triassic. Elevated turnover rates are documented at the Permian–Triassic boundary, and two distinct diversification events are noted in the wake of this biotic crisis, a first one during the Early Triassic (dipnoans, actinistians, ‘palaeopterygians’, ‘subholosteans’) and a second one during the Middle Triassic (‘subholosteans’, neopterygians). The origination of new, small taxa predominantly among these groups during the Middle Triassic event caused a significant reduction in osteichthyan body size. Neopterygii, the clade that encompasses the vast majority of extant fishes, underwent another diversification phase in the Late Triassic. The Triassic radiation of Osteichthyes, predominantly of Actinopterygii, which only occurred after severe extinctions among Chondrichthyes during the Middle–Late Permian, resulted in a profound change within global fish communities, from chondrichthyan‐rich faunas of the Permo‐Carboniferous to typical Mesozoic and Cenozoic associations dominated by actinopterygians. This turnover was not sudden but followed a stepwise pattern, with leaps during extinction events.     

BOOM AND BUST: ANCIENT AND RECENT DIVERSIFICATION IN BICHIRS (POLYPTERIDAE: ACTINOPTERYGII), A RELICTUAL LINEAGE OF RAY‐FINNED FISHES

Understanding the history that underlies patterns of species richness across the Tree of Life requires an investigation of the mechanisms that not only generate young species‐rich clades, but also those that maintain species‐poor lineages over long stretches of evolutionary time. However, diversification dynamics that underlie ancient species‐poor lineages are often hidden due to a lack of fossil evidence. Using information from the fossil record and time calibrated molecular phylogenies, we investigate the history of lineage diversification in Polypteridae, which is the sister lineage of all other ray‐finned fishes (Actinopterygii). Despite originating at least 390 million years (Myr) ago, molecular timetrees support a Neogene origin for the living polypterid species. Our analyses demonstrate polypterids are exceptionally species depauperate with a stem lineage duration that exceeds 380 million years (Ma) and is significantly longer than the stem lineage durations observed in other ray‐finned fish lineages. Analyses of the fossil record show an early Late Cretaceous (100.5–83.6 Ma) peak in polypterid genus richness, followed by 60 Ma of low richness. The Neogene species radiation and evidence for high‐diversity intervals in the geological past suggest a “boom and bust” pattern of diversification that contrasts with common perceptions of relative evolutionary stasis in so‐called “living fossils.”     

Ocean basin and depth variability of oxygen isotopes in cenozoic benthic foraminifera

One of the longest, most detailed quantitative records of oceanographic change in the Cenozoic is that provided by oxygen isotope measurements made on the tests of foraminifera. As indicated by measurements on benthic foraminifera, the deep waters of the world ocean have undergone an overall cooling of about 10°C in the Cenozoic. This change has been neither monotonic nor gradual. Rather, it is evidenced by a few, relatively rapid increases in the ¹³O content of the benthic shells. These “steps” in the isotopic record have been associated with major evolutionary changes in the mean state of the deep ocean. The variance around this mean state has also changed through the Cenozoic. From relatively high variance in the Middle Eocene, the oceans showed low variance in the Late Eocene and Oligocene. In the Miocene the variance of the isotopic measurement again increased, reaching a maximum of short duration in the Middle Miocene. This maximum as well as that which occurred during the Late Pliocene and Quaternary, may be attributable to fluctuations in the isotopic composition of the oceans caused by growth and decay of large ice sheets.In the Late Miocene the benthic oxygen isotopes in Atlantic sites less than 3000 m deep have a higher variance than sites at similar depths in the Pacific and Indian Oceans. It is thought that this high variance results from long-term changes in the importance of the cool and salty North Atlantic Deep Water relative to that of the warmer and less saline Antarctic Intermediate Water at Atlantic sites between 1000 m and 3000 m water depth. Such significant differences in benthic isotopic variance between the ocean basins have been demonstrated only in post-Middle Miocene intervals.