Millennial-scale faunal record reveals differential resilience of European large mammals to human impacts across the Holocene

The use of short-term indicators for understanding patterns and processes of biodiversity loss can mask longer-term faunal responses to human pressures. We use an extensive database of approximately 18 700 mammalian zooarchaeological records for the last 11 700 years across Europe to reconstruct spatio-temporal dynamics of Holocene range change for 15 large-bodied mammal species. European mammals experienced protracted, non-congruent range losses, with significant declines starting in some species approximately 3000 years ago and continuing to the present, and with the timing, duration and magnitude of declines varying individually between species. Some European mammals became globally extinct during the Holocene, whereas others experienced limited or no significant range change. These findings demonstrate the relatively early onset of prehistoric human impacts on postglacial biodiversity, and mirror species-specific patterns of mammalian extinction during the Late Pleistocene. Herbivores experienced significantly greater declines than carnivores, revealing an important historical extinction filter that informs our understanding of relative resilience and vulnerability to human pressures for different taxa. We highlight the importance of large-scale, long-term datasets for understanding complex protracted extinction processes, although the dynamic pattern of progressive faunal depletion of European mammal assemblages across the Holocene challenges easy identification of ‘static’ past baselines to inform current-day environmental management and restoration.     

A long-term association between global temperature and biodiversity, origination and extinction in the fossil record

The past relationship between global temperature and levels of biological diversity is of increasing concern due to anthropogenic climate warming. However, no consistent link between these variables has yet been demonstrated. We analysed the fossil record for the last 520 Myr against estimates of low latitude sea surface temperature for the same period. We found that global biodiversity (the richness of families and genera) is related to temperature and has been relatively low during warm 'greenhouse' phases, while during the same phases extinction and origination rates of taxonomic lineages have been relatively high. These findings are consistent for terrestrial and marine environments and are robust to a number of alternative assumptions and potential biases. Our results provide the first clear evidence that global climate may explain substantial variation in the fossil record in a simple and consistent manner. Our findings may have implications for extinction and biodiversity change under future climate warming.     

Reconstructing past species assemblages reveals the changing patterns and drivers of extinction through time

Predicting future species extinctions from patterns of past extinctions or current threat status relies on the assumption that the taxonomic and biological selectivity of extinction is consistent through time. If the driving forces of extinction change through time, this assumption may be unrealistic. Testing the consistency of extinction patterns between the past and the present has been difficult, because the phylogenetically explicit methods used to model present-day extinction risk typically cannot be applied to the data from the fossil record. However, the detailed historical and fossil records of the New Zealand avifauna provide a unique opportunity to reconstruct a complete, large faunal assemblage for different periods in the past. Using the first complete phylogeny of all known native New Zealand bird species, both extant and extinct, we show how the taxonomic and phylogenetic selectivity of extinction, and biological correlates of extinction, change from the pre-human period through Polynesian and European occupation, to the present. These changes can be explained both by changes in primary threatening processes, and by the operation of extinction filter effects. The variable patterns of extinction through time may confound attempts to identify risk factors that apply across time periods, and to infer future species declines from past extinction patterns and current threat status.     

Primate extinction risk and historical patterns of speciation and extinction in relation to body mass

Body mass is thought to influence diversification rates, but previous studies have produced ambiguous results. We investigated patterns of diversification across 100 trees obtained from a new Bayesian inference of primate phylogeny that sampled trees in proportion to their posterior probabilities. First, we used simulations to assess the validity of previous studies that used linear models to investigate the links between IUCN Red List status and body mass. These analyses support the use of linear models for ordinal ranked data on threat status, and phylogenetic generalized linear models revealed a significant positive correlation between current extinction risk and body mass across our tree block. We then investigated historical patterns of speciation and extinction rates using a recently developed maximum-likelihood method. Specifically, we predicted that body mass correlates positively with extinction rate because larger bodied organisms reproduce more slowly, and body mass correlates negatively with speciation rate because smaller bodied organisms are better able to partition niche space. We failed to find evidence that extinction rates covary with body mass across primate phylogeny. Similarly, the speciation rate was generally unrelated to body mass, except in some tests that indicated an increase in the speciation rate with increasing body mass. Importantly, we discovered that our data violated a key assumption of sample randomness with respect to body mass. After correcting for this bias, we found no association between diversification rates and mass.     

Patterns of extinction and persistence of Arctotertiary flora in Iberia during the Quaternary

The native or allochthonous nature of certain extant species of the Iberian Peninsula is a matter of some controversy given our lack of knowledge regarding the survival and extinction processes to which they have been subject. The aim of the present work is to provide a review of the current knowledge regarding these processes during the Quaternary. The Middle Pleistocene Transition was a period of noticeable disappearance of Arctotertiary taxa in the Iberian Peninsula, related to the alteration in glacial cyclicity and climate change, including an increase in the severity of glaciations, a reduction in the length of the interglacial periods, and increased dryness during the coldest times. It is difficult to maintain that human activities played any important part in the complete or near-complete disappearance of some taxa during the Quaternary. In the Late Pleistocene and Holocene, however, the extinction of Cedrus, Picea and Carpinus might be related to climate change, competition with other taxa, and anthropogenic disturbances. In contrast, the extreme reduction or even complete disappearance of Platanus or Syringa seems to be due to natural causes alone. The available geological, molecular and palaeobotanical data suggest that some Arctotertiary taxa persisted over long periods of time during the Quaternary in the Iberian Peninsula. Indeed, the fossil record of one important group of taxa, including Castanea, Ceratonia, Carpinus and Juglans, shows its continued presence throughout the Quaternary.     

The influence of denitrifying seawater on graptolite extinction and diversification during the Hirnantian (latest Ordovician) mass extinction event

A continuous trench exposure within the uppermost type Vinini Formation at Vinini Creek, Roberts Mountains, Nevada, provides an unparalleled opportunity to examine the fate of graptolites, prominent Paleozoic zooplankton, during most of the Hirnantian mass extinction event. On the basis of a detailed biostratigraphic and sedimentological dataset, the relatively complete extinction record is examined in the context of ecological constraints, and it is found to reflect an ecological collapse driven by glacio-eustatic sea-level fall and associated changes in oceanic circulation. Diverse graptolite populations of the Dicranograptidae-Diplograptidae-Orthograptidae (DDO) fauna, which flourished in denitrifying waters within the oceanic oxygen-minimum zone (OMZ) during sea-level highstand, largely vanished with the loss of these conditions during glacio-eustatic sea-level fall. However, populations of one clade, the normalograptids, which inhabited the oxygenated waters of the photic zone, not only survived but diversified. These survivors gave rise to rapid recolonization and diversification with re-establishment of the oxygen-minimum and denitrifying conditions during post-Hirnantian sea-level rise. This ecological model also applies globally to other well-documented coeval stratigraphic intervals, representing both oceanic and platform sea settings.     

The Frasnian-Famennian mass extinction: insights from high-resolution sequence stratigraphy and cyclostratigraphy in South China

Two sequences (SFr, SFa), each 1-1.2 Myr in duration, are recognised in the strata across the Frasnian-Famennian (F-F) transition both in carbonate platform and interplatform basinal successions in South China. The sequence boundary between the two sequences is placed a little below the top of the Frasnian. The sequences are basically composed of coarsening-upward/bed-thickness increasing-upward cycles and shallowing-upward cycles (parasequences) in basinal and platform deposits respectively, which stack into cycle-sets (typically six to eight cycles). 10 and 12 cycle-sets are identified in sequences SFr and SFa respectively. These cycle-sets can be further grouped into larger-scale composite cycle-sets (herein termed mesocycle- and megacycle-sets with two and four cycle-sets respectively). This vertical cycle-stacking pattern and the hierarchy of cyclicity suggest a Milankovitch style of forcing such that the cycles and cycle-sets were formed in response to the orbital perturbations of precession (16-18 kyr) and eccentricity (∼100 kyr in duration), respectively. In the basinal cycles, smaller-scale rhythmic stratification beds (typically six to eight beds in a cycle) are extensive, and were likely caused by millennial-scale climatic forcing. In the lower sequence, SFr, the latest highstand deposits consist of calciturbidites and debrites in deep-water strata and fenestral limestones in shallow-water strata, representing a major (third-order) sea-level fall. Within these deposits, four cycle-sets are further identified in both coeval deep-water and platform successions. Succeeding deeper-water organic-rich facies, within which three cycles occur, are the transgressive deposits of the overlying Famennian sequence (SFa). These cycles represent three higher-frequency (16-18 kyr) sea-level fluctuations and accompanying anoxia, superimposed on a major third-order sea-level rise. The F-F boundary is placed at the top of the first cycle, based on conodont data. Thus, a major sea-level fall and then a rise occurred in the F-F transitional period. Faunal and sedimentological data reveal a massive biotic decline in concert with the major sea-level fall, and a further biotic demise coinciding with the major sea-level rise and its three superimposed higher-frequency sea-level fluctuations and accompanying anoxia. The F-F biotic crisis was therefore characterised by two episodes of step-down extinction. On the basis of Milankovitch orbital rhythms, the first major biotic extinction took place over ∼400 kyr, and the subsequent event was ∼50 kyr in duration, i.e. ∼450 kyr for the entire event. At the same time as the massive decline of normal-marine fossils during the latest Frasnian sea-level fall, there was widespread cyanobacterial growth and a thriving of planktonic calcispheres, suggesting eutrophic conditions. This situation could have caused a severe biotic loss, as a result of the deterioration of surface water clarity and formation of anoxic bottom waters due to over-consumption of oxygen through respiratory demands and decomposition by the cyanobacteria and phytoplankton. The subsequent rapid sea-level rise with superimposed higher-frequency sea-level fluctuations and accompanying anoxia could have caused rapid elevation of anoxic bottom waters and expansion of eutrophic surface waters over shallow-water platforms due to enhanced upwelling ocean currents and improved ocean circulation. This situation would have exerted further stresses upon the already-weakened biota, leading to a further biotic demise. However, a small number of organisms such as pelagic tentaculitids, small mud-adapted brachiopods and gastropods did survive into the Famennian, although with very low diversity.     

When the pond turtle followed the reindeer: effect of the last extreme global warming event on the timing of faunal change in Northern Europe

Faunal communities have been shaped in different ways by past climatic change. The impact of the termination of the last Glacial and the onset of the present (Holocene) Interglacial on large‐scale faunal shifts, extinction dynamics and gene pools of species are of special interest in natural sciences. A general pattern of climate‐triggered range expansion and local extinction of vertebrate species is known for Europe, and shows that in the modern temperate zone the main faunal change took place mainly during the Late Glacial (14 700–11 700 years ago) and Early Holocene (11 700–9 100 years ago). Based on large datasets of new radiocarbon data, we present precise temporal dynamics of climate‐driven disappearance and appearance of reindeer and pond turtle in southern Sweden. These two species are significant climate indicators in Late Quaternary biostratigraphy. Our data reveal that the reindeer disappeared from southern Sweden ca. 10 300 years ago, whereas the pond turtle colonized the area ca. 9 860 years ago, with a 450‐year gap between each species. This provides evidence for a sudden environmental turnover, causing the replacement of an arctic faunal element by a thermophilic species. The postglacial range dynamics of pond turtle and reindeer are a unique model case, allowing insights into the faunal turnover of other vertebrates during the last dramatic natural global warming event at the Pleistocene–Holocene transition.     

Conodonts Meet Cladistics: Recovering Relationships and Assessing the Completeness of the Conodont Fossil Record

A numerical cladistic analysis of the conodont family Palmatolepidae has been undertaken to determine the applicability of the technique to group-wide systematic revision. Results suggest a new hypothesis of relationships that is considerably more parsimonious than trees compatible with existing hypotheses of relationships, or trees that are even loosely constrained stratigraphically. This may occur either because the fossil record is incomplete, because taxon sampling for the cladistic analysis is low, or because the most parsimonious trees approximate the true tree less well than do stratigraphically-constrained trees (or because of a combination of these factors). Although more taxa and more characters would be preferable in choosing between these possibilities, the tree derived solely from morphological data is adopted. Thus, stratigraphic data can be used to test hypotheses of relationships and construct phylogenies; hypotheses of relationships can be used to test the completeness of the conodont fossil record. Existing schemes of classification within the Palmatolepidae are rejected because most groups within them are either polyphyletic or paraphyletic. A new scheme is presented. Character changes suggest correlated, progressive and mosaic evolution within the Palmatolepidae. Parsimony analysis of partitioned datasets indicates that more phylogenetic information can be recovered from S rather than P or M element positions, although data from all three positional groups are preferable to data from just one. Thus, multielement taxonomy is essential to the resolution of conodont interrelationships.     

Fossils and living taxa agree on patterns of body mass evolution: a case study with Afrotheria

Most of life is extinct, so incorporating some fossil evidence into analyses of macroevolution is typically seen as necessary to understand the diversification of life and patterns of morphological evolution. Here we test the effects of inclusion of fossils in a study of the body size evolution of afrotherian mammals, a clade that includes the elephants, sea cows and elephant shrews. We find that the inclusion of fossil tips has little impact on analyses of body mass evolution; from a small ancestral size (approx. 100 g), there is a shift in rate and an increase in mass leading to the larger-bodied Paenungulata and Tubulidentata, regardless of whether fossils are included or excluded from analyses. For Afrotheria, the inclusion of fossils and morphological character data affect phylogenetic topology, but these differences have little impact upon patterns of body mass evolution and these body mass evolutionary patterns are consistent with the fossil record. The largest differences between our analyses result from the evolutionary model, not the addition of fossils. For some clades, extant-only analyses may be reliable to reconstruct body mass evolution, but the addition of fossils and careful model selection is likely to increase confidence and accuracy of reconstructed macroevolutionary patterns.     

The Luoping biota: exceptional preservation, and new evidence on the Triassic recovery from end-Permian mass extinction

The timing and nature of biotic recovery from the devastating end-Permian mass extinction (252 Ma) are much debated. New studies in South China suggest that complex marine ecosystems did not become re-established until the middle–late Anisian (Middle Triassic), much later than had been proposed by some. The recently discovered exceptionally preserved Luoping biota from the Anisian Stage of the Middle Triassic, Yunnan Province and southwest China shows this final stage of community assembly on the continental shelf. The fossil assemblage is a mixture of marine animals, including abundant lightly sclerotized arthropods, associated with fishes, marine reptiles, bivalves, gastropods, belemnoids, ammonoids, echinoderms, brachiopods, conodonts and foraminifers, as well as plants and rare arthropods from nearby land. In some ways, the Luoping biota rebuilt the framework of the pre-extinction latest Permian marine ecosystem, but it differed too in profound ways. New trophic levels were introduced, most notably among top predators in the form of the diverse marine reptiles that had no evident analogues in the Late Permian. The Luoping biota is one of the most diverse Triassic marine fossil Lagerstätten in the world, providing a new and early window on recovery and radiation of Triassic marine ecosystems some 10 Myr after the end-Permian mass extinction.     

The contribution of mass drug administration to global health: past,present and future

Mass drug administration (MDA) is a means of delivering safe and inexpensive essential medicines based on the principles of preventive chemotherapy, where populations or sub-populations are offered treatment without individual diagnosis. High-coverage MDA in endemic areas aims to prevent and alleviate symptoms and morbidity on the one hand and can reduce transmission on the other, together improving global health. MDA is the recommended strategy of the World Health Organisation to control or eliminate several neglected tropical diseases (NTDs). More than 700 million people now receive these essential NTD medicines annually. The combined cost of integrated NTD MDA has been calculated to be in the order of $0.50 per person per year. Activities have recently been expanded due, in part, to the proposed attempt to eliminate certain NTDs in the coming two decades. More than 1.9 billion people need to receive MDA annually across several years if these targets are to be met. Such extensive coverage will require additional avenues of financial support, expanded monitoring and evaluation focusing on impact and drug efficacy, as well as new diagnostic tools and social science strategies to encourage adherence. MDA is a means to help reduce the burden of disease, and hence poverty, among the poorest sector of populations. It has already made significant improvements to global health and productivity and has the potential for further successes, particularly where incorporated into sanitation and education programmes. However logistical, financial and biological challenges remain.     

The high fidelity of the cetacean stranding record: insights into measuring diversity by integrating taphonomy and macroecology

Stranded cetaceans have long intrigued naturalists because their causation has escaped singular explanations. Regardless of cause, strandings also represent a sample of the living community, although their fidelity has rarely been quantified. Using commensurate stranding and sighting records compiled from archived datasets representing nearly every major ocean basin, I demonstrated that the cetacean stranding record faithfully reflects patterns of richness and relative abundance in living communities, especially for coastlines greater than 2000 km and latitudinal gradients greater than 4°. Live-dead fidelity metrics from seven different countries indicated that strandings were almost always richer than live surveys; richness also increased with coastline length. Most death assemblages recorded the same ranked relative abundance as living communities, although this correlation decreased in strength and significance at coastline lengths greater than 15,000 km, highlighting the importance of sampling diversity at regional scales. Rarefaction analyses indicated that sampling greater than 10 years generally enhanced the completeness of death assemblages, although protracted temporal sampling did not substitute for sampling over longer coastlines or broader latitudes. Overall, this global live-dead comparison demonstrated that strandings almost always provided better diversity information about extant cetacean communities than live surveys; such archives are therefore relevant for macroecological and palaeobiological studies of cetacean community change through time.     

Generic species richness and body mass in North American mammals: Support for the inverse relationship of body size and speciation rate

Generic species richness, the number of species per genus, is examined as a function of mean generic body mass for extant North American mammals. Species richness decreases as an inverse power function with increased mass, and the Spearman rank correlation coefficient of the logio transformed data is significant (r_s= ‐0.37). When the data are partitioned by trophic level, the relationship is not statistically significant for carnivores but strengthens for herbivores (r_s= ‐0.46). This interesting but incidental effect is due to the negligible number of diminutive and excessively large carnivores, which is in turn determined by foraging strategies. Alternate hypotheses for the “right‐skewed”; size distribution of modern North American mammals, such as disproportionate extinction of large species, differential species longevity, and a geographical scaling function, are rejected in favor of the proposition that elevated levels of speciation are restricted to animals of small body mass, as originally proposed by Gould and Eldredge (1977). This phenomenon is explained as a function of habitat restriction and particularly in herbivores, limited home range size. Aquatic mammals, regardless of body size, speciate rarely. Cope's Rule, the tendency of many animal groups to evolve towards large size, is understood as a probabilistic statement reflecting the phylogenetic tendencies of a disproportionately high number of small species alive at any given point in time.     

The legacy of past climate and landscape change on species' current experienced climate and elevation ranges across latitude: a multispecies study utilizing mammals in western North America

Aim Elevation and climate ranges across latitude experienced by 21 wide‐ranging mammal species in western North America were summarized to examine two questions: (1) do populations in the northern and southern portions of a species’ range experience different climates or are environments selected to remain similar to climates at the core of ranges; and (2) how do species’ elevational ranges, experienced temperature seasonality and temperature ranges change across latitude? Given the larger effects of climate oscillations in the north vs. the south, a predicted outcome is for species to conserve climate niches across latitude and to show reduced climate and elevation ranges in the north. An alternative outcome is latitudinal niche diversification and increased climate variation in the north. Location Western North America. Methods The questions above were examined using a combination of species occurrence data bases, climate data bases, simple summaries of means and standard deviations and by testing summaries against random distributions across latitude for 21 mammal species from a variety of orders. Results The results showed that: (i) most species conserve their niche strongly or weakly given overall temperature gradients from north to south; (ii) seasonality experienced by species is relatively static until the highest latitudes despite directional trends across the region; and (iii) the elevation range and temperature variation that species experience decreases from south to north. Main conclusions Populations at range edges appear to partition environments to remain closer to temperature values similar to those at the core of the range. In addition, seasonality is not a likely explanatory factor of genetic diversity in latitudinal gradients. The data are instead more consistent with predictions that a combination of higher gene‐flow, increasing environmental instability and decreasing elevation gradients in the north compared to the south may lead to negative correlations between latitude and species’ climate variation. The results corroborate risks faced by northern mammal populations to global climate changes.     

The significance of the oil palm (Elaeis guineensis Jacq.) in the late Holocene environments of west and west central Africa: A further consideration

Abundant archaeological evidence of the occurrence of the endocarp of the oil palm,Elaeis guineensis, in the rain forest and woodland savanna zones of west and central Africa from about 5000 B.P. has shown the tree to be an important element in the subsistence economy of the region; its pollen also has been recorded in most of the regional terrestrial sediments studied so far. The distinct and consistently sudden and more marked increases in this pollen during the late Holocene when compared with the late Tertiary and late Pleistocene frequencies strongly indicate that the late Holocene upsurges were due to both natural and human factors favourable for the expansion of this heliophytic tree. Reasons are given for suggesting that upsurges in oil palm pollen during the late Holocene period in this region can be used as indices of plant cultivation. While the oil palm is known from early Tertiary deposits in west Africa, its earliest palynological record from terrestrial sediments in the west central part dates back only to the early Holocene. More palynological studies of Tertiary and Quaternary terrestrial cores are required to establish with more certainty the antiquity ofE. guineensis in west central Africa.