Phylogenetic structure of the extinction and biotic factors explaining differential survival of terrestrial notosuchians at the Cretaceous–Palaeogene crisis

Although the clade Crocodylomorpha is represented by few extant species (Crocodylia), it has a rich fossil record. Hundreds of species, adapted to terrestrial, semi-aquatic and marine environments, have existed over more than 200 million years. Numerous studies have attempted to characterize the factors driving the diversification and extinction events of Crocodylomorpha, resulting in ambiguous and even contradictory conclusions, which points to the need for phylogenetically and temporally smaller-scaled studies. Here, we focus on differential survival at the Cretaceous–Palaeogene (K–Pg) crisis of Notosuchia, a diverse clade of mostly terrestrial Crocodylomorpha that achieved great diversity during the Cretaceous. More precisely, we tested the effect of body size and palaeotemperatures on notosuchian survival probability during the K–Pg crisis as well as the effect of diet on the evolution of their body size. We find that Notosuchia showed an evolutionary trend towards larger body sizes through time, associated with a shift from an omnivorous to a carnivorous diet. This may explain why sebecids were the only notosuchians to survive the K–Pg crisis. We also corroborate the conclusions of previous studies that detected a Lagerstätten effect occurring in the Adamantina Formation (Upper Cretaceous, Brazil, Bauru Group). This work confirms the value of more finely-scaled macroevolutionary studies for understanding the history of a rich and complex group such as Crocodylomorpha.     

Facies of two important Early Triassic gastropod lagerstätten: implications for diversity patterns in the aftermath of the end-Permian mass extinction

Two important lagerstätten of Early Triassic gastropods, the Sinbad Limestone (Utah, USA) and the Gastropod Oolite (North Italy) yield about 40% of all described Early Triassic species. This great contribution to the global diversity and the exceptional good preservation render high information content, which characterizes fossil lagerstätten. The Smithian Sinbad Limestone contains the most diverse Early Triassic gastropod fauna. At the type locality, it occurs in single, probably storm-induced shell bed within a series of high energy deposits underlain by intertidal microbial mats and subtidal oolite/peloid shoals. The main shell bed contains about 40 invertebrate taxa. Gastropods, scaphopods, and bivalves are most abundant and form an assemblage, which is dominated by small neritaemorphs, the opisthobranch Cylindrobullina convexa and the scaphopod Plagioglypta (annulated tubes). This assemblage lived on shallow, subtidal soft-bottoms based on sedimentological and ecological characteristics. The Dienerian (to Smithian?) Gastropod Oolite Member (North Italy) has extremely abundant, probably salinity-controlled gastropod faunas with low species richness. Almost monospecific assemblages of Pseudomurchisonia kokeni as well as assemblages with about four species are present in the Gastropod Oolite. Modern hydrobiid mudsnail faunas which are adapted to strongly fluctuating salinity in intertidal to shallow subtidal coastal areas form probably a suitable model for the Gastropod Oolite biota. Gastropods from the Werfen- and Moenkopi-Formation lagerstätten are well preserved compared to other Early Triassic deposits. The high contribution to the global diversity of just two sites suggests very incomplete sampling and preservational bias. However, the low richness of the major faunas reflects depauperate Early Triassic faunas and slow recovery from the Permian/Triassic crisis.     

Mammal survival at the Cretaceous–Palaeogene boundary: metabolic homeostasis in prolonged tropical hibernation in tenrecs

Free-ranging common tenrecs, Tenrec ecaudatus, from sub-tropical Madagascar, displayed long-term (nine months) hibernation which lacked any evidence of periodic interbout arousals (IBAs). IBAs are the dominant feature of the mammalian hibernation phenotype and are thought to periodically restore long-term ischaemia damage and/or metabolic imbalances (depletions and accumulations). However, the lack of IBAs in tenrecs suggests no such pathology at hibernation T_bs > 22°C. The long period of tropical hibernation that we report might explain how the ancestral placental mammal survived the global devastation that drove the dinosaurs and many other vertebrates to extinction at the Cretaceous–Palaeogene boundary following a meteorite impact. The genetics and biochemistry of IBAs are of immense interest to biomedical researchers and space exploration scientists, in the latter case, those envisioning a hibernating state in astronauts for deep space travel. Unravelling the physiological thresholds and temperature dependence of IBAs will provide new impetus to these research quests.     

Ostracod recovery in the aftermath of the Permian–Triassic crisis: Palaeozoic–Mesozoic turnover

Results of a detailed bathymetric survey of Crater Lake conducted in 2000, combined with previous results of submersible and dredge sampling, form the basis for a geologic map of the lake floor and a model for the filling of Crater Lake with water. The most prominent landforms beneath the surface of Crater Lake are andesite volcanoes that were active as the lake was filling with water, following caldera collapse during the climactic eruption of Mount Mazama 7700 cal. yr B.P. The Wizard Island volcano is the largest and probably was active longest, ceasing eruptions when the lake was 80 m lower than present. East of Wizard Island is the central platform volcano and related lava flow fields on the caldera floor. Merriam Cone is a symmetrical andesitic volcano that apparently was constructed subaqueously during the same period as the Wizard Island and central platform volcanoes. The youngest postcaldera volcanic feature is a small rhyodacite dome on the east flank of the Wizard Island edifice that dates from 4800 cal. yr B.P. The bathymetry also yields information on bedrock outcrops and talus/debris slopes of the caldera walls. Gravity flows transport sediment from wall sources to the deep basins of the lake. Several debris-avalanche deposits, containing blocks up to 280 m long, are present on the caldera floor and occur below major embayments in the caldera walls. Geothermal phenomena on the lake floor are bacterial mats, pools of solute-rich warm water, and fossil subaqueous hot spring deposits. Lake level is maintained by a balance between precipitation and inflow versus evaporation and leakage. High-resolution bathymetry reveals a series of up to nine drowned beaches in the upper 30 m of the lake that we propose reflect stillstands subsequent to filling of Crater Lake. A prominent wave-cut platform between 4 m depth and present lake level that commonly is up to 40 m wide suggests that the surface of Crater Lake has been at this elevation for a very long time. Lake level apparently is limited by leakage through a permeable layer in the northeast caldera wall. The deepest drowned beach approximately corresponds to the base of the permeable layer. Among a group of lake filling models, our preferred one is constrained by the drowned beaches, the permeable layer in the caldera wall, and paleoclimatic data. We used a precipitation rate 70% of modern as a limiting case. Satisfactory models require leakage to be proportional to elevation and the best fit model has a linear combination of 45% leakage proportional to elevation and 55% of leakage proportional to elevation above the base of the permeable layer. At modern precipitation rates, the lake would have taken 420 yr to fill, or a maximum of 740 yr if precipitation was 70% of the modern value. The filling model provides a chronology for prehistoric passage zones on postcaldera volcanoes that ceased erupting before the lake was filled.     

Type-Maastrichtian gastropod faunas show rapid ecosystem recovery following the Cretaceous–Palaeogene boundary catastrophe

The study of the global mass extinction event at the Cretaceous–Palaeogene (K/Pg) boundary can aid in understanding patterns of selective extinction, and survival and dynamics of ecosystem recovery. Outcrops in the Maastrichtian type area (south-east Netherlands, north-east Belgium) comprise a stratigraphically expanded K/Pg boundary succession that offers a unique opportunity to study marine ecosystem recovery within the first few thousand years following the mass extinction event. A quantitative analysis was performed on systematically sampled macrofossils of the topmost Maastrichtian and lowermost Danian strata at the former Ankerpoort-Curfs quarry (Geulhem), which represent ‘snapshots’ of the latest Cretaceous and earliest Palaeogene marine ecosystems, respectively. Molluscs in particular are diverse and abundant in the studied succession. Regional ecosystem changes across the K/Pg boundary are relatively minor, showing a decline in suspension feeders, accompanied by an ecological shift to endobenthic molluscs. The earliest Paleocene gastropod assemblage retains many ‘Maastrichtian’ features and documents a fauna that temporarily survived into the Danian. The shallow, oligotrophic carbonate platform in this area was inhabited by taxa that were adapted to low nutrient levels and resistant to starvation. As a result, the local taxa were less affected by the short-lived detrimental conditions related to K/Pg boundary perturbations, such as darkness, cooling, starvation and ocean acidification. This resulted in relatively high survival rates, which enabled rapid recolonization and recovery of marine faunas in the Maastrichtian type area.     

Stepwise and large-magnitude negative shift in δ13Ccarb preceded the main marine mass extinction of the Permian–Triassic crisis interval

Large perturbations to the global carbon cycle occurred during the Permian–Triassic boundary mass extinction, the largest extinction event of the Phanerozoic Eon (542 Ma to present). Controversy concerning the pattern and mechanism of variations in the marine carbonate carbon isotope record of the Permian–Triassic crisis interval (PTCI) and their relationship to the marine mass extinction has not been resolved to date. Herein, high-resolution carbonate carbon isotope profiles (δ¹³C_carb), accompanied by lithofacies, were generated for four sections with microbialite (Taiping, Zuodeng, Cili, and Chongyang) in South China to better constrain patterns and controls on δ¹³C_carb variation in the PTCI and to test hypotheses about the temporal relationship between perturbations to the global carbon cycle and the marine mass extinction event. All four study sections exhibit a stepwise negative shift in δ¹³C_carb during the Late Permian–Early Triassic, with the shift preceding the end-Permian crisis being larger (> 3‰) than that following it (1–2‰). The pre-crisis shifts in δ¹³C_carb are widely correlatable and, hence, represent perturbations to the global carbon cycle. The comparatively smaller shifts following the crisis demonstrate that the marine mass extinction event itself had at most limited influence on the global carbon cycle, and that both Late Permian δ¹³C_carb shifts and the mass extinction must be attributed to some other cause. Their origin cannot be uniquely determined from C-isotopic data alone but appears to be most compatible with a mechanism based on episodic volcanism in combination with collapse of terrestrial ecosystems and soil erosion.     

Mutation and evolutionary rates in adélie penguins from the antarctic

Precise estimations of molecular rates are fundamental to our understanding of the processes of evolution. In principle, mutation and evolutionary rates for neutral regions of the same species are expected to be equal. However, a number of recent studies have shown that mutation rates estimated from pedigree material are much faster than evolutionary rates measured over longer time periods. To resolve this apparent contradiction, we have examined the hypervariable region (HVR I) of the mitochondrial genome using families of Adélie penguins (Pygoscelis adeliae) from the Antarctic. We sequenced 344 bps of the HVR I from penguins comprising 508 families with 915 chicks, together with both their parents. All of the 62 germline heteroplasmies that we detected in mothers were also detected in their offspring, consistent with maternal inheritance. These data give an estimated mutation rate (micro) of 0.55 mutations/site/Myrs (HPD 95% confidence interval of 0.29-0.88 mutations/site/Myrs) after accounting for the persistence of these heteroplasmies and the sensitivity of current detection methods. In comparison, the rate of evolution (k) of the same HVR I region, determined using DNA sequences from 162 known age sub-fossil bones spanning a 37,000-year period, was 0.86 substitutions/site/Myrs (HPD 95% confidence interval of 0.53 and 1.17). Importantly, the latter rate is not statistically different from our estimate of the mutation rate. These results are in contrast to the view that molecular rates are time dependent.     

Gradualistic characean lineages in the Upper Cretaceous–Palaeocene of southern Europe

Two charophyte lineages from the Upper Cretaceous–Palaeocene of southern Europe are described here, constituting a useful biostratigraphic tool for correlating non-marine stratigraphic sequences. The Peckichara pectinata lineage, ranging from the middle Campanian to the middle–upper Maastrichtian, consists of four successive gyrogonite morphotypes that displayed progressively more complex gyrogonite ornamentation and increasingly smaller gyrogonites with time. The Peckichara sertulata lineage, ranging from the middle Campanian to the Danian, consists of three morphotypes that followed a more common evolutionary trend, developing progressively larger gyrogonites without any changes in ornamentation. Intermediate forms between the successive morphotypes suggest that they corresponded to anagenetic varieties of the same evolutionary species rather than to separate species. The different trends observed appear to be palaeoenvironmentally controlled. The grades of the P. pectinata lineage first occurred in permanent lakes that later shifted to fluvial influenced floodplain ponds, this shift accounting for the reduction in gyrogonite size. By contrast, the P. sertulata lineage mainly occurred in permanent lakes, where stable palaeoenvironmental conditions enabled a steady rise in gyrogonite size to increase the space available for the zygote.     

Discovery of charophyte flora across the Cretaceous–Paleocene transition in the Jiaolai Basin

Charophyte assemblages from the Cretaceous/Paleogene (K/Pg) boundary transition in the JZK-2 borehole of the Jiaolai Basin (eastern China) were studied. The charophyte flora includes Tolypella grambastii, Peckichara praecursoria, Microchara cristata, Microchara prolixa, Chara changzhouensis, Lamprothamnium ellipticum, Nodosochara (Turbochara) specialis, Lychnothamnus aff. vectensis, and Lychnothamnus lanpingensis n. comb. The discovery of the ostracod species Porpocypris sphaeroidalis Guan in the upper section of the JZK-2 borehole strongly indicates the arrival of the Paleocene. Three floras of the latest Cretaceous to the early Paleocene were compared among the Jiaolai, Pingyi, and Songliao basins. The Pingyi Basin is special because it is the only carbonate palaeolake that occurred throughout the K/Pg transition in China. It is characterized by the appearance of the Cretaceous brackish water element, Feistiella anluensis (Wang), and is dominated by the Paleocene species, Peckichara varians Grambast. The Songliao and Jiaolai floras grow in clastic lakes; the Jiaolai Basin is thought to correspond to an altitude lake, which was formed in the coastal mountains with high palaeoelevation (≥ 2.0 km), while the Songliao Basin is presumed to have been formed in a low elevation site according to previous studies. This difference in altitude explains why the Songliao Basin is more species-rich than the Jiaolai Basin.     

Biodiversity and biogeography of middle–late liassic ammonoids: implications for the early Toarcian mass extinction

In Western Tethyan areas, the Toarcian stage begins with two important evolutionary events in ammonite faunas: (1) the disruption of Tethyan–Boreal provinciality; (2) a biological crisis linked with the oceanic anoxic event OAE. The analysis of these events has been addressed by constructing curves of ammonoid diversity (species richness, origination and extinction rates) in the Late Pliensbachian (= Domerian)–Early Toarcian interval in selected localities. Two diversity drops are recognized. The first one is recorded at the end of the Dactylioceras mirabile subzone and reflects the disruption of Tethyan–Boreal provinciality, through the progressive extinction of the Boreal endemic family Amaltheidae that occupied the north-western European seas during the whole Pliensbachian on the one hand, and the extinction of Late Domerian Ammonitina endemic to the Mediterranean areas on the other hand. The Early Toarcian homogeneization of Mediterranean and north-western European ammonoid faunas was reached via elimination of both Boreal and Mediterranean endemics with differential rates of extinction in the two palaeogeographic domains and the subsequent geographical expansion of Tethyan-derived ammonoids. The second, dramatic drop in ammonite diversity in the upper part of the Dactylioceras semicelatum subzone coincided with the onset of OAE. It also affected epioceanic ammonoid clades like Phyllocerataceae and Lytocerataceae. These two drops are interpreted as two distinct extinctions and not as episodes of a single, stepwise event. Complex relations between ammonoid diversity and sea-level changes are suggested by trends in endemism, which may be reversed during either a single transgression or a single regression.     

Can the Lilliput Effect be detected in the brachiopod faunas of South China following the terminal Ordovician mass extinction?

In the immediate aftermath of global extinctions, organisms were normally much smaller than those prior to these events. This ‘Lilliput Effect’ can be subdivided into two types: 1) a specific type, following the original definition of the effect which targets species-level taxa associated with inhospitable environments, and 2) a more general type, related to the reactions of higher-rank taxa above the species-level. The body sizes of brachiopods from South China through the Ordovician and Silurian transition (Late Katian, Hirnantian, and earliest Rhuddanian) are compared at generic, superfamilial, ordinal, and class levels. The results indicate that the body sizes of the taxa of lower rank (e.g. genus-level) are highly variable within these different intervals. The type of evidence for the Lilliput Effect through the end Ordovician mass extinction is thus quite different from that of the end Permian mass extinction probably reflecting differences in the intensity of these two major bioevents. However, the relationships between the contrasting trends in body-size change of some taxa of higher rank (e.g. at the ordinal-level) and the relative dominance of these taxa in the latest Ordovician and earliest Silurian suggest that the brachiopods of the two major Ordovician groups, the strophomenoids and orthoids, adopted different survival strategies during and immediately after the crisis from those of the pentamerides and rhynchonellides, that were common in Silurian assemblages.     

Tracing the evolutionary routes of plant–microbiota interactions

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Temporal change in the morphometry–body mass relationship of polar bears

Accurate information on animal body mass is often an essential component of wildlife research and management. However, for many large-bodied species, obtaining direct scale weights from individuals may be difficult. In these cases, morphometric equations (e.g., based on girth or length) may provide accurate and precise estimates of body mass. We developed predictive equations to estimate the body mass of free-ranging polar bears (Ursus maritimus) in western Hudson Bay, Canada. Using multiple linear and non-linear regression, we identified a strong relationship between polar bear body weight and linear measures of straight line length and axillary girth. The mass–morphometry relationship appeared to change over time and we developed separate equations for polar bears measured during 2 time periods, 1980–1996 and 2007–2009. Non-linear models were more accurate and provided body mass estimates within 5.8% (R² = 0.98) and 6.1% (R² = 0.98) of scale weight in the earlier and later time periods, respectively. Earlier equations developed for polar bears in this subpopulation performed poorly when applied to recently sampled individuals. In contrast, some contemporary equations from other regions performed reasonably well, suggesting that temporal changes within a subpopulation may be more pronounced than regional differences and can render earlier predictive equations obsolete. Our results have important implications for current and future studies of polar bear body condition and the effects of ongoing climate warming. © 2011 The Wildlife Society.     

Liquid chromatography–tandem mass spectrometry for the determination of low-molecular mass aldehydes in human urine

A simple and sensitive method is proposed for the determination of seven low-molecular mass aldehydes in human urine samples using liquid chromatography with tandem mass spectrometric detection. Urine samples diluted twofold with 0.3 M hydrochloric acid are aspirated into a LiChrolut EN solid-phase extraction column impregnated with 2,4-dinitrophenylhydrazine for cleanup, derivatization and preconcentration of the aldehydes. After elution of the hydrazones with acetonitrile, an aliquot is injected directly into the chromatograph. Identification and quantification of aldehydes was performed with electrospray in negative ion mode by selected reaction monitoring. By using synthetic urine samples, linearity is established over the concentration range 0.1–30 μg/l and limits of detection from 15 to 65 ng/l. The intra- and inter-day precision (RSD, %) of the aldehydes ranged from 2.9% to 6.4% and 3.6% to 9.3%, respectively, and specific uncertainties were ca. 5.0 ± 0.3 ng for all aldehydes. Average recoveries performed on two levels by enriching synthetic urine samples ranged between 92% and 100%. The method was also validated in terms of study sample stability including long-term and short-term analyte stability, freeze–thaw and extract stability. In summary, the method proposed surpasses other recent chromatographic alternatives in terms of the limit of detection and sample requirements for analysis.     

Mating systems and protein–protein interactions determine evolutionary rates of primate sperm proteins

To assess the relative impact of functional constraint and post-mating sexual selection on sequence evolution of reproductive proteins, we examined 169 primate sperm proteins. In order to recognize potential genome-wide trends, we additionally analysed a sample of altogether 318 non-reproductive (brain and postsynaptic) proteins. Based on cDNAs of eight primate species (Anthropoidea), we observed that pre-mating sperm proteins engaged in sperm composition and assembly show significantly lower incidence of site-specific positive selection and overall lower non-synonymous to synonymous substitution rates (d_N/d_S) across sites as compared with post-mating sperm proteins involved in capacitation, hyperactivation, acrosome reaction and fertilization. Moreover, database screening revealed overall more intracellular protein interaction partners in pre-mating than in post-mating sperm proteins. Finally, post-mating sperm proteins evolved at significantly higher evolutionary rates than pre-mating sperm and non-reproductive proteins on the branches to multi-male breeding species, while no such increase was observed on the branches to unimale and monogamous species. We conclude that less protein–protein interactions of post-mating sperm proteins account for lowered functional constraint, allowing for stronger impact of post-mating sexual selection, while the opposite holds true for pre-mating sperm proteins. This pattern is particularly strong in multi-male breeding species showing high female promiscuity.     

Invasion waves and pinning in the Kirkpatrick–Barton model of evolutionary range dynamics

Journal of Mathematical Biology - The Kirkpatrick–Barton model, well known to invasion biologists, is a pair of reaction–diffusion equations for the joint evolution of population...     

Is specialization an evolutionary dead end? Testing for differences in speciation,extinction and trait transition rates across diverse phylogenies of specialists and generalists

Specialization has often been claimed to be an evolutionary dead end, with specialist lineages having a reduced capacity to persist or diversify. In a phylogenetic comparative framework, an evolutionary dead end may be detectable from the phylogenetic distribution of specialists, if specialists rarely give rise to large, diverse clades. Previous phylogenetic studies of the influence of specialization on macroevolutionary processes have demonstrated a range of patterns, including examples where specialists have both higher and lower diversification rates than generalists, as well as examples where the rates of evolutionary transitions from generalists to specialists are higher, lower or equal to transitions from specialists to generalists. Here, we wish to ask whether these varied answers are due to the differences in macroevolutionary processes in different clades, or partly due to differences in methodology. We analysed ten phylogenies containing multiple independent origins of specialization and quantified the phylogenetic distribution of specialists by applying a common set of metrics to all datasets. We compared the tip branch lengths of specialists to generalists, the size of specialist clades arising from each evolutionary origin of a specialized trait and whether specialists tend to be clustered or scattered on phylogenies. For each of these measures, we compared the observed values to expectations under null models of trait evolution and expected outcomes under alternative macroevolutionary scenarios. We found that specialization is sometimes an evolutionary dead end: in two of the ten case studies (pollinator‐specific plants and host‐specific flies), specialization is associated with a reduced rate of diversification or trait persistence. However, in the majority of studies, we could not distinguish the observed phylogenetic distribution of specialists from null models in which specialization has no effect on diversification or trait persistence.     

Discontinuities and alternative scalings in the density–mass relationship of anuran larvae

Size–density relationships (SDRs) frequently follow a power-law relationship, with exponents that compensate for the increase in population-level metabolic demand—the energetic equivalence rule. However, these exponents present a range of values, and elucidating its methodological and biological determinants has become a main issue. So far, a restricted set of potential relationships, mechanisms, and taxa have been considered. Here, we analyzed SDR in a population of tadpoles inhabiting a network of 16 temporal ponds. Alternative scaling regimes were detected using pacewise regressions and estimating exponents with maximum likelihood (ML). If discontinuities in the SDR are ignored, a scaling close to values reported elsewhere is observed. However, estimated slopes between discontinuities are steeper (threefold to fivefold) than those often reported, but congruent with the performance predicted for ML and the biases reported for other methods. Our estimations largely deviate from an energetic equivalence, suggesting that large individuals use less energy per unit area. The detection of different SDRs in the same database, with a strong decay in abundance with body size, points to a pattern poorly considered in previous studies, widening the range of patterns, mechanisms, and ecological, or evolutionary consequences of the SDRs.