The bioclimatic model: a method of palaeoclimatic qualitative inference based on mammal associations

Aim The bioclimatic model is a new method for palaeoclimatic reconstruction built on the assumption of a significant correlation between climate and mammal community composition. The goal of this approach is to infer past climatic conditions using mammal fossil associations as source data. Location The study used mammal faunas from all over the world to develop the bioclimatic model. As an example of the potential of the model, we have applied it to Quaternary faunas from Eurasia. Methods The proposed model was constructed by applying multivariate discriminant analysis to modern mammal faunas and climates from throughout the world. The model was validated with a different set of modern faunas than those used in the discriminant analysis, including some from transitional zones between different climates (ecotones). To test the reliability of the method in the Pleistocene, the results have been compared to those obtained with data from other disciplines, such as palaeobotany. Results The results obtained in the validation of the model show that more than 90% of the localities have been classified correctly. Comparisons of results in the late Pleistocene‐Holocene of Barová between a palaeobotanical study and the bioclimatic analysis show the latter to be highly accurate. The results for early Pleistocene faunas show somewhat drier and more open climatic conditions for Europe than the present day, with larger areas of steppe environments. Main Conclusions The bioclimatic model could be used to infer climatic conditions from mammal faunas. The results presented in this work provide a preliminary example of the potential that bioclimatic analysis has as a tool for palaeoclimatic inference. Finally, this method offers the opportunity to standardize data coming from vertebrate palaeontology for use in the construction and evaluation of climatic models.     

Life in proto‐Amazonia: Middle Miocene mammals from the Fitzcarrald Arch (Peruvian Amazonia)

The Middle Miocene has been identified as a time of great diversification in modern lineages now distributed in tropical South America, and when basic archetypal traits defining Amazonia appear, including climatic humid conditions, basic floral physiognomy and phylogenetic composition of modern rainforests. Nonetheless, Middle Miocene localities in South America are poorly known, especially at low latitudes where only one species‐rich locality, La Venta in Colombia, has been extensively studied. The present contribution describes the mammal fauna of Fitzcarrald, a new Middle Miocene local fauna from western Amazonia in Peru. Fitzcarrald is correlated with the Laventan South American Land Mammal Age based on the presence of taxa defining the ‘Miocochilius assemblage zone’ in La Venta. The mammalian fauna of Fitzcarrald comprises 24 taxa among cingulates, folivores, astrapotheres, notoungulates, litopterns, rodents, odontocetes and a possible marsupial. At this time, tropical South America was characterized by the presence of the Pebas megawetland, a huge lacustrine complex that provided unique ecological and environmental conditions most likely isolating northern South America from southern South America. These isolating conditions might have come to an end with its disappearance in the Late Miocene and the establishment of the subsequent Acre system, the predecessor fluvial system of modern Amazonia. Results of faunistic similarity between Fitzcarrald and other Miocene faunas throughout South America support these scenarios. The Fitzcarrald mammal fauna exhibits first appearance datums and last appearance datums of various taxa, showing that tropical South America has played a crucial role in the evolutionary history and biogeography of major clades, and revealing a more complex biological history than previously proposed, based on the record from the southern cone of the continent.     

Contrasts in the large herbivore faunas of the southern continents in the late Pleistocene and the ecological implications for human origins

Aim

Africa is renowned for the current abundance and diversity of its large mammals. The aim of this study was to assess distinctions evident in the functional composition of continental large herbivore faunas during the late Pleistocene before extinctions depleted the megafauna outside Africa.

Location

The African large herbivore fauna was compared with that formerly inhabiting South America, Australia, North America, Eurasia and tropical Asia during the late Pleistocene.

Methods

Pleistocene faunas were reconstructed from the literature in terms of their relative body size composition, grazer/browser contributions and taxonomic representations, omitting forest and island species.

Results

Although the three southern continents were closely similar in the overall species richness of large herbivores that they supported during the late Pleistocene, South America had a predominance of very large herbivores, while most of Australia's mammalian herbivores were relatively small and those of Africa were intermediate. Africa had many more grazers, especially in the size range 100–1000 kg, than other continents. The South American pattern resembled that in North America and Eurasia, while Africa and Australia diverged in different ways.

Main conclusions

Neither the total extent of savannas in each continent nor the morphological features enabling bovid radiation seemed adequate on their own to explain the greater richness of macrograzers in Africa. Africa is characterized by the widespread occurrence of arid/eutrophic savannas, which are unrepresented in other continents. The prevalence of savanna is partly attributable to the high elevation of interior eastern and southern Africa, associated with relatively low rainfall, and to the comparatively high soil fertility, related to volcanic influences. This promoted an abundance and diversity of medium‐sized grazing ruminants unrivalled elsewhere. Indigenous grasses in South America and Australia are less well adapted to withstand severe grazing than the African grasses introduced to support livestock. The locally high abundance of African ungulates presented conditions that facilitated the adaptive transition by early hominins from plant‐gatherers to meat‐scavengers.     

Human influence on distribution and extinctions of the late Pleistocene Eurasian megafauna

Late Pleistocene extinctions are of interest to paleontological and anthropological research. In North America and Australia, human occupation occurred during a short period of time and overexploitation may have led to the extinction of mammalian megafauna. In northern Eurasia megafaunal extinctions are believed to have occurred over a relatively longer period of time, perhaps as a result of changing environmental conditions, but the picture is much less clear. To consider megafaunal extinction in Eurasia, we compare differences in the geographical distribution and commonness of extinct and extant species between paleontological and archaeological localities from the late middle Pleistocene to Holocene. Purely paleontological localities, as well as most extinct species, were distributed north of archaeological sites and of the extant species, suggesting that apart from possible differences in adaptations between humans and other species, humans could also have a detrimental effect on large mammal distribution. However, evidence for human overexploitation applies only to the extinct steppe bison Bison priscus. Other human-preferred species survive into the Holocene, including Rangifer tarandus, Equus ferus, Capreolus capreolus, Cervus elaphus, Equus hemionus, Saiga tatarica, and Sus scrofa. Mammuthus primigenius and Megaloceros giganteus were rare in archaeological sites. Carnivores appear little influenced by human presence, although they become rarer in Holocene archaeological sites. Overall, the data are consistent with the conclusion that humans acted as efficient hunters selecting for the most abundant species. Our study supports the idea that the late Pleistocene extinctions were environmentally driven by climatic changes that triggered habitat fragmentation, species range reduction, and population decrease, after which human interference either by direct hunting or via indirect activities probably became critical.     

Quantitative palaeoclimatic inference based on terrestrial mammal faunas

Aim The main goal of this paper is to propose a new tool, in the form of Quantitative Bioclimatic Models (QnBMs), for reconstructing past climates based on fossil mammal associations. As a case study, European climatic conditions during the basal early Pleistocene will be inferred using QnBMs. Location The study uses faunas throughout the world to develop a quantitative bioclimatic model, which is then applied to Quaternary faunas from Eurasia. Methods The models were constructed by applying multivariate linear regression to modern mammal faunas and climates from all over the world. The models were validated with a second group of modern faunas, which includes several from transitional zones between different climates (ecotones). To test the reliability of the method when applied to fossil associations, the results obtained for the Pleistocene have been compared with those obtained from palynology. Results Validation of the models shows that as many as 11 climatic factors can be inferred with high reliability using the regression models developed in this work. Comparisons of results in the late Pleistocene–Holocene of Barová (Czech Republic) between a palynological study and the quantitative bioclimatic analysis show a high degree of similarity. The results for the early Pleistocene show colder and drier climatic conditions for Europe than today. Main conclusions The application of the quantitative bioclimatic models to present day as well as Quaternary mammal faunas proves to be a useful tool for palaeoclimatic reconstruction during the Quaternary and probably most of the Neogene. Transfer functions are presented for a complete set of climatic factors, allowing a precise estimation of the climate in a locality from its mammal fauna.     

Plate tectonics, seaways and climate in the historical biogeography of mammals

The marsupial and placental mammals originated at a time when the pattern of geographical barriers (oceans, shallow seas and mountains) was very different from that of today, and climates were warmer. The sequence of changes in these barriers, and their effects on the dispersal of the mammal families and on the faunas of mammals in the different continents, are reviewed. The mammal fauna of South America changed greatly in the Pliocene/Pleistocene, when the newly-complete Panama Isthmus allowed the North American fauna to enter the continent and replace most of the former South American mammal families. Marsupial, but not placental, mammals reached Australia via Antarctica before Australia became isolated, while rats and bats are the only placentals that dispersed naturally from Asia to Australia in the late Cenozoic. Little is known of the early history of the mammal fauna of India. A few mammal families reached Madagascar from Africa in the early Cenozoic over a chain of islands. Africa was isolated for much of the early Cenozoic, though some groups did succeed in entering from Europe. Before the climate cooled in the mid-Cenozoic, the mammal faunas of the Northern Hemisphere were much richer than those of today.     

Cenozoic environmental change in South America as indicated by mammalian body size distributions (cenograms)

A cenogram is a rank‐ordered body size distribution of non‐predatory terrestrial mammal species within a community. Studies of cenograms for modern faunas have shown that certain quantifiable attributes of cenograms are correlated with environmental variables such as rainfall and vegetation structure. Based on these correlations, cenograms of fossil communities have been used to infer palaeoenvironments and palaeoenvironmental variables. The present study uses cenogram statistics to interpret palaeoenvironmental conditions for eight Cenozoic South American mammal faunas, ranging from Eocene to Pleistocene in age. Body sizes for fossil taxa were taken either from the literature or were estimated using regressions of body size on molar length (or femoral bicondylar width) for modern mammals. Cenogram statistics are calculated for the eight fossil faunas and compared to similar statistics calculated for 16 modern South American mammal faunas, allowing palaeoenvironmental interpretations to be made. The palaeoenvironmental interpretations based on cenogram analyses sometimes support and sometimes contradict interpretations based on herbivore craniodental morphology (e.g. levels of hypsodonty). Simulations of expected errors in body size estimates for fossil taxa suggest that the discrepancies do not result primarily from erroneous body size estimates. It is possible that some of the incongruity in interpretations results from certain non‐analogue attributes of South American faunas during much of the Cenozoic (e.g. the relatively depauperate mammalian predator diversity prior to the Great American Biotic Interchange).     

Sequence of mammalian fossils, including hominoid teeth, from the Bubing Basin caves, South China

A Plio-Pleistocene to Holocene faunal sequence has been recovered from four carefully excavated caves in the Bubing Basin, adjacent to the larger Bose Basin of South China. The caves vary in elevation; we suggest that the higher caves were formed and filled with sediments prior to the lower caves. The highest deposits, which are from Mohui Cave, contain hominoid teeth and other fossilized remains of mammalian taxa most similar to late Pliocene and early Pleistocene faunas. Wuyun Cave ( approximately 50m lower in elevation than Mohui) contains a late middle Pleistocene fauna, which is supported by U-series age constraints from 350 to 200ka. Lower Pubu Cave ( approximately 23m below Wuyun) is assigned to the late Pleistocene, while the Cunkong Cave (the lowest, approximately 2m lower elevation than Lower Pubu) preserves a Holocene fauna. The four faunal assemblages indicate species-level changes in Ailuropoda, Stegodon, and Sus, the appearance of Elephas, the local disappearance of Stegodon, and the migration of Equus hemionus to South China. These initial results of our work call into question the continued value of the Stegodon/Ailuropoda Fauna, a category long used to characterize the Pleistocene faunas of South China. Excavation of karstic caves of varying elevation within the basins of South China holds promise for defining local sequences of mammalian fossils that can be used to investigate faunal variations related to climate change, biogeographic events, and evolutionary change over the past two million years. Stable isotopic analysis of a small sample of mammalian teeth from Bubing Basin caves is consistent with 100% C(3) vegetation in the Bubing/Bose region, with certain delta(13)C values consistent with a canopied woodland or forest. A preliminary assessment of the hominoid teeth indicates the presence of diverse molar and premolar morphologies including dental remains of Gigantopithecus blacki and a sample with similarities to the teeth reported from Longgupo.     

Body size structure in north-western Mediterranean Plio-Pleistocene mammalian faunas

Aim We investigated the patterns of body‐size changes of the north‐western Mediterranean Plio‐Pleistocene large mammal faunas (excluding rodents, bats, lagomorphs and insectivores) in order to identify the tempo and mode of the major shifts in body size distribution, and to put them in the context of Plio‐Pleistocene environmental changes and the development of the Mediterranean climate. Location We analysed fossil faunas of Spain, France and Italy. A set of recent regional faunas from several macroclimatic regions was selected to serve as elements for comparison of the size distribution of past faunas, consisting of: Spain, France and Italy together, Florida, California, Central Chile, Indochina, India, Korea‐Manchuria, Malawi, The Cape, North Africa, Turkey and Australia. Methods Mammal species were grouped into five body size categories for carnivores and four categories for noncarnivore species. The number of species in each size category was computed and the resulting matrix of body weight classes × regions and time intervals was used as an input matrix in a Correspondence Analysis. Results Recent and fossil faunas strongly differ in body size structure. The distribution of recent faunas within the CA seems to reflect both ecological and historic factors, intertwined in a complex fashion. No clear relationship has been observed between body size structure and environmental factors. During the late Pliocene to early Pleistocene there were only minor changes in the pattern of size distribution, although plant communities were in a transition process from subtropical forests to Mediterranean woodlands and steppes. The major change in body size structure of the north‐western Mediterranean fauna occurred at the Galerian, around 1 Ma ago. This marked the beginning of the modern fauna, and a general trend towards a larger body size, reduction in the number of medium sized herbivores, and an increase of large herbivores and megaherbivores. Main conclusions The Plio‐Pleistocene faunas lack modern analogues. The body size structure of mammalian regional faunas appears to be strongly dependent on historical factors. The only major shift in body size distribution occurred during the Plio‐Pleistocene, in the late Villafranchian‐Galerian transition, coincident with the onset of the Pleistocene high intensity glacial cycles.     

Are Chilean coastal forests pre‐Pleistocene relicts? Evidence from foliar physiognomy,palaeoclimate, and phytogeography

Aim We ask whether contemporary forests of the Chilean Coastal Range can be considered to be direct and conservative descendants of pre‐Pleistocene palaeofloras that occurred in southern South America from the Palaeogene to early Neogene periods (65–10 Ma), maintaining foliar physiognomies that do not match their present‐day climate. We also identify the most likely ancestors of present‐day coastal forests. Location Coastal Range of south‐central Chile (33–40° S). Methods We compared leaf morphology between five representative modern floras from mid‐latitude forests of the Chilean Coastal Range, and 14 Palaeogene–early Neogene palaeofloras from southern South America. We also compared the composition of biogeographical elements (defined by the modern distribution of plant genera) between fossil and present‐day assemblages. Palaeoclimatic reconstructions were based on a canonical correspondence analysis between leaf morphology of modern assemblages and eight climatic variables, and tested by a Monte Carlo permutation test. We compared the relative positions of fossil and modern floras on the environmental vector space defined by Canoco, and on axes defined by instrumental and estimated temperature and precipitation data. Results According to foliar characters, Palaeogene palaeofloras were strikingly divergent from present‐day coastal forests of central Chile. In contrast, two extant forest floras of the Chilean Coastal Range have a foliar morphology that resembles some late Eocene to early Miocene mixed palaeofloras, at least 23 Myr older. These two modern sites are representative of an area of the Coastal Range (36–37° S) that has been highlighted for its relictual character. None of the 14 fossil floras corresponded exactly to the modern composition of phytogeographic elements, although correspondence analyses showed that mixed and Neogene subtropical fossil floras were compositionally close to the extant woody floras of coastal forests in central Chile. Main Conclusions Contemporary forests of the Chilean Coastal Range exhibit strong physiognomic resemblance to the mixed palaeofloras from 33°57′ to 41°15′ S, which may be the closest ancestor of the deciduous and endemic‐rich Maulino forest, presently restricted to coastal areas between 36° and 38° S. In turn, the Neogene subtropical palaeoflora that occurred in the Proto‐Andean foothills of central Chile is the likely predecessor of Mediterranean‐type sclerophyllous forests of central Chile (32–33° S). Despite foliar resemblance between the late pre‐Pleistocene and extant forest floras, our palaeoclimatic reconstructions suggest that modern assemblages exist under climatic conditions that do not match their foliar physiognomy. We attribute this convergence in foliar morphology to the ‘evolutionary inertia’ of surviving lineages, favoured by the buffering effect of the coastal environment on climatic variability.     

Combined statistical and dynamical assessment of simulated vegetation–rainfall interactions in North Africa during the mid‐Holocene1

A negative feedback of vegetation cover on subsequent annual precipitation is simulated for the mid‐Holocene over North Africa using a fully coupled general circulation model with dynamic vegetation, FOAM‐LPJ (Fast Ocean Atmosphere Model‐Lund Potsdam Jena Model). By computing a vegetation feedback parameter based on lagged auto‐covariances, the simulated impact of North African vegetation on precipitation is statistically quantified. The feedback is also dynamically assessed through initial value ensemble experiments, in which North African grass cover is initially reduced and the climatic response analyzed. The statistical and dynamical assessments of the negative vegetation feedback agree in sign and relative magnitude for FOAM‐LPJ. The negative feedback on annual precipitation largely results from a competition between bare soil evaporation and plant transpiration, with increases in the former outweighing reductions in the latter given reduced grass cover. This negative feedback weakens and eventually reverses sign over time during a transient simulation from the mid‐Holocene to present. A similar, but weaker, negative feedback is identified in Community Climate System Model Version 2 (CCSM2) over North Africa for the mid‐Holocene.     

Mammals in Last 30 to 7 ka Interval (Late Pleistocene-Early Holocene) in Southern Uruguay (Santa Lucía River Basin): Last Occurrences,Climate, and Biogeography

Fossiliferous Quaternary sedimentary beds in the Santa Lucía Basin (southern Uruguay) are potentially useful for the study of the last occurrences of extinct taxa, as well as the environmental and climatic patterns in the late Pleistocene to the early Holocene. They have provided a chronological framework (AMS ¹⁴C and Optically Stimulated Luminescence dates), a mammalian diversity and interpretations of last occurrence, the climatic-environmental setting, and some associated biogeographic processes. The ages produced encompass the last 30 to 7 ka interval (latest Pleistocene-early Holocene). The mammalian assemblage (36 genera, 24 species) includes typical South American late Pleistocene mammals, extinct species of extant genera, and some extant species that still exist elsewhere on the continent. The preservation pattern includes articulated and semi-articulated skeletons of large and small mammals. The presence in the southern Uruguayan Pampean area of some mammals currently inhabiting Patagonia, northwest and central Argentina (Dolichotis, Galea, Microcavia, Chaetophractus, Lagostomus, and Vicugna) is explained by the predominance of open areas and cold climates associated with the Last Glacial Maximum. The mammalian record depicts local extinctions or shifting ranges occurring in latest Pleistocene or early Holocene. The sedimentary chronological framework and taphonomic features suggest the persistence into the early Holocene of Eutatus seguini, Morenelaphus brachyceros, Equus neogeus, Hemiauchenia sp., Lama sp., and Vicugna sp. Glyptodon and Glossotherium seem to persist at least to the latest Pleistocene. This pattern can substantiate the hypothesis that some megafauna and large mammals persisted for some millennia alongside people with extinction occurring before, during, and after human colonization.     

Biodiversity from a historical geology perspective: a case study from Marajó Island, lower Amazon

Assessing patterns of abundance and distribution of Amazonian species is still an overwhelming task that requires integration of multiple disciplines. This work is based on background information gathered from previous reconstructions of the geological history of the lower Amazon drainage basin, in order to analyse biodiversity patterns within the context of landscape transformation. A highly dynamic geological scenario is depicted for this area during the Plio‐Pleistocene and Holocene, which consisted of a large palaeovalley formed as a response of tectonic reactivation. This palaeovalley was filled with sediments transported by a north/northwest orientated palaeo‐Tocantins River. The palaeodrainage became abandoned as the main river course was deviated to the northeast, initiating the separation of Marajó Island from mainland. Geology had a direct impact on the modern physiognomy, with open vegetation dominating in areas with Holocene sedimentation, while closed forests prevailing in older Quaternary and, probably also, Pliocene terrains. Data from fossil and modern mammalian groups indicate the connection of Marajó Island to the mainland during the Last Glacial Maximum, when open vegetation seems to have dominated. Tectonic subsidence was responsible for the maintenance of this vegetation pattern on the eastern side of the Marajó Island, keeping it as a habitat favourable for savanna adapted faunal elements. Based on this kind of information, this work attempts to highlight the importance of integrating studies combining geological and biological events as the key to understand biodiversity patterns in Amazonia. It is expected to open new lines of research dealing with the comprehension of ecology, species and genetic diversity, biogeography, evolutionary scenarios, and speciation mechanisms.     

Body mass‐related changes in mammal community assembly patterns during the late Quaternary of North America

The late Quaternary of North America was marked by prominent ecological changes, including the end‐Pleistocene megafaunal extinction, the spread of human settlements and the rise of agriculture. Here we examine the mechanistic reasons for temporal changes in mammal species association and body size during this time period. Building upon the co‐occurrence results from Lyons et al. (2016) – wherein each species pair was classified as spatially aggregated, segregated or random – we examined body mass differences (BMD) between each species pair for each association type and time period (Late Pleistocene: 40 000 ¹⁴C–11 700 ¹⁴C ybp, Holocene: 11 700 ¹⁴C–50 ybp and Modern: 50–0 yr). In the Late Pleistocene and Holocene, the BMD of both aggregated and segregated species pairs was significantly smaller than the BMD of random pairs. These results are consistent with environmental filtering and competition as important drivers of community structure in both time periods. Modern assemblages showed a breakdown between BMD and co‐occurrence patterns: the average BMD of aggregated, segregated and random species pairs did not differ from each other. Collectively, these results indicate that the late Quaternary mammalian extinctions not only eliminated many large‐bodied species but were followed by a re‐organization of communities that altered patterns of species coexistence and associated differences in body size.     

Mammal diversity and environment evolution during the Plio-Pleistocene in East Africa

Human evolution began in East Africa four million years ago, with a transition from an arboreal state to a more terrestrial one. This evolution seems to be correlated with a large environmental change in East Africa around 2.5 m.y. due to a major climatic change leading to drier and cooler conditions. Cenogram analysis (a graphical representation of community structure) can be used to reconstruct the vegetation cover at a regional scale, and to infer the changing climatic conditions. Using cenogram sequences of different sites along the Rift Valley, we were able to determine the regional ecological context in which mammals and hominids have evolved in East Africa during the last 3 million years. Between 3.5 and 2 m.y., during a general climatic change, successive faunas of South Tanzania reflect the progressive opening of their environment. In contrast around Lake Turkana a mosaic of isolated dry and wet habitats were present throughout this period. At this time, the Rift seems to have been spatially structured in several basins isolated from one other, and isolated faunas experienced separate speciation events (particularly with the appearance ofHomo genus). After 2 m.y., the disappearance of the isolating barriers on one hand, and a regional increase in aridity, on the other hand, led to more homogenous faunas arising throughout the region. Replacements of mammal species occurred (especiallyHomo erectus replacingHomo habilis) and several others mammal species, including australopithecines, disappeared during this same period.     

Mountain uplift explains differences in Palaeogene patterns of mammalian evolution and extinction between North America and Europe

Patterns of late Palaeogene mammalian evolution appear to be very different between Eurasia and North America. Around the Eocene–Oligocene (EO) transition global temperatures in the Northern Hemisphere plummet: following this, European mammal faunas undergo a profound extinction event (the Grande Coupure), while in North America they appear to pass through this temperature event unscathed. Here, we investigate the role of surface uplift to environmental change and mammalian evolution through the Palaeogene (66–23 Ma). Palaeogene regional surface uplift in North America caused large-scale reorganization of precipitation patterns, particularly in the continental interior, in accord with our combined stable isotope and ecometric data. Changes in mammalian faunas reflect that these were dry and high-elevation palaeoenvironments. The scenario of Middle to Late Eocene (50–37 Ma) surface uplift, together with decreasing precipitation in higher-altitude regions of western North America, explains the enigma of the apparent lack of the large-scale mammal faunal change around the EO transition that characterized western Europe. We suggest that North American mammalian faunas were already pre-adapted to cooler and drier conditions preceding the EO boundary, resulting from the effects of a protracted history of surface uplift.     

Co‐declining mammal–dung beetle faunas throughout the Atlantic Forest biome of South America

The millennial–scale evolutionary relationships between mammals and dung beetles have been eroded due to several drivers of contemporary biodiversity loss. Although some evidence of co‐decline has been shown for mammals and dung beetles at some Neotropical sites, a biome‐scale analysis for the entire Atlantic Forest of South America would strengthen our understanding of how relictual sets of mammal species can affect dung beetle co‐occurrences and co‐declines. We therefore collated hundreds of assemblages of both dung beetles and medium‐ to large‐bodied mammals throughout the world's longest tropical forest latitudinal gradient to examine to what extent mammal assemblages may exert a positive influence on dung beetle species composition and functional assembly, and whether this relationship is scale dependent. We also collated several climatic and other environmental variables to examine the degree to which they shape mammal–dung beetle relationships. The relationships between local mammal and dung beetle faunas were examined using regression models, variation partitioning, dissimilarity indices and ecological networks. We found a clear positive relationship between mammal and dung beetle species richness across this forest biome, indicating an ongoing process of mammal–dung beetle niche‐mediated co‐decline. We found a strong relationship between the species composition of both taxa, in which dung beetle species dissimilarity apparently track changes in mammalian dissimilarity, typically in 80% of all cases. Co‐variables such as phytomass and climatic variables also influenced mammal–dung beetle patterns of co‐decline along the Atlantic Forest. We conclude that dung beetle diversity and community assembly are shaped by the remaining co‐occurring mammal assemblages and their functional traits, and both groups were governed by environmental features. We emphasize that ecosystem‐wide effects of mammal population declines remain poorly understood both quantitatively and qualitatively, and curbing large vertebrate defaunation will ensure the persistence of co‐dependent species.     

Historical biogeography and phenotype‐phylogeny of Chroodiscus (lichenized Ascomycota: Ostropales: Graphidaceae)

Aim To analyse the historical biogeography of the lichen genus Chroodiscus using a phenotype‐based phylogeny in the context of continental drift and evolution of tropical rain forest vegetation. Location All tropical regions (Central and South America, Africa, India, Southeast Asia, north‐east Australia). Methods We performed a phenotype‐based phylogenetic analysis and ancestral character state reconstruction of 14 species of the lichen genus Chroodiscus, using paup * and mesquite ; dispersal–vicariance analysis (DIVA) and dispersal–extinction–cladogenesis (DEC) modelling to trace the geographical origin of individual clades; and ordination and clustering by means of pc‐ord , based on a novel similarity index, to visualize the biogeographical relationships of floristic regions in which Chroodiscus occurs. Results The 14 species of Chroodiscus show distinctive distribution patterns, with one pantropical and one amphi‐Pacific taxon and 12 species each restricted to a single continent. The genus comprises four clades. DIVA and DEC modelling suggest a South American origin of Chroodiscus in the mid to late Cretaceous (120–100 Ma), with subsequent expansion through a South American–African–Indian–Southeast Asian–Australian dispersal route and late diversification of the argillaceus clade in Southeast Asia. Based on the abundance of extant taxa, the probability of speciation events in Chroodiscus is shown to be extremely low. Slow dispersal of foliicolous rain forest understorey lichens is consistent with estimated phylogenetic ages of individual species and with average lengths of biological species intervals in fungi (10–20 Myr). Main conclusions The present‐day distribution of Chroodiscus can be explained by vicariance and mid‐distance dispersal through the interconnection or proximity of continental shelves, without the need for recent, trans‐oceanic long‐distance dispersal. Phylogenetic reconstruction and age estimation for Chroodiscus are consistent with the ‘biotic ferry’ hypothesis: a South American origin and subsequent eastward expansion through Africa towards Southeast Asia and north‐eastern Australia via the Indian subcontinent. The present‐day pantropical distributions of many clades and species of foliicolous lichens might thus be explained by eastward expansion through continental drift, along with the evolution of modern rain forests starting 120 Ma, rather than by the existence of a hypothetical continuous area of pre‐modern rain forest spanning South America, Africa and Southeast Asia during the mid and late Cretaceous.     

Holocene mammal extinctions in the Carpathian Basin: a review

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