Effects of Climate History and Environmental Grain on Species’ Distributions in Africa and South America

Climate changes during the Pleistocene produced shifts, reductions, and expansions of biomes that, in turn, have been hypothesized to have driven speciation and extinction and shaped patterns of biodiversity. Here, we explore effects of Late Pleistocene climatic changes on environmentally and geographically cohesive areas mimicking species’ distributions. We analyzed persistence of these ‘species’ over the transition from the warm Last Interglacial period to the cool Last Glacial Maximum period to warm present‐day conditions, for four levels of environmental restriction (5, 10, 15 and 20% of overall variation; akin to niche breadths). African environments were overall much less conserved over these periods than those of South America, matching diversity contrasts between the two continents. Results thus indicate that biodiversity patterns relate closely to historical patterns of environmental grain and their stability through time; this view is a step toward an integral understanding of the role of environmental and geographic factors in the process of biological diversification.     

Predictable variation of range-sizes across an extreme environmental gradient in a lizard adaptive radiation: evolutionary and ecological inferences

Large-scale patterns of current species geographic range-size variation reflect historical dynamics of dispersal and provide insights into future consequences under changing environments. Evidence suggests that climate warming exerts major damage on high latitude and elevation organisms, where changes are more severe and available space to disperse tracking historical niches is more limited. Species with longer generations (slower adaptive responses), such as vertebrates, and with restricted distributions (lower genetic diversity, higher inbreeding) in these environments are expected to be particularly threatened by warming crises. However, a well-known macroecological generalization (Rapoport's rule) predicts that species range-sizes increase with increasing latitude-elevation, thus counterbalancing the impact of climate change. Here, I investigate geographic range-size variation across an extreme environmental gradient and as a function of body size, in the prominent Liolaemus lizard adaptive radiation. Conventional and phylogenetic analyses revealed that latitudinal (but not elevational) ranges significantly decrease with increasing latitude-elevation, while body size was unrelated to range-size. Evolutionarily, these results are insightful as they suggest a link between spatial environmental gradients and range-size evolution. However, ecologically, these results suggest that Liolaemus might be increasingly threatened if, as predicted by theory, ranges retract and contract continuously under persisting climate warming, potentially increasing extinction risks at high latitudes and elevations.     

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

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

Pleistocene Niche Stability and Lineage Diversification in the Subtropical Spider Araneus omnicolor (Araneidae)

The influence of Quaternary climate oscillations on the diversification of the South American fauna is being increasingly explored. However, most of these studies have focused on taxa that are endemic to tropical environments, and relatively few have treated organisms restricted to subtropical biomes. Here we used an integrative phylogeographical framework to investigate the effects of these climate events on the ecological niche and genetic patterns of the subtropical orb-weaver spider Araneus omnicolor (Araneidae). We analyzed the mitochondrial (Cytochrome Oxidase I, COI) and nuclear (Internal Transcribed Subunit II, ITS2) DNA of 130 individuals throughout the species’ range, and generated distribution models in three different climate scenarios [present, Last Glacial Maximum (LGM), and Last Interglacial Maximum (LIG)]. Additionally, we used an Approximate Bayesian Computation (ABC) approach to compare possible demographic scenarios and select the hypothesis that better explains the genetic patterns of A. omnicolor. We obtained high haplotype diversity but low nucleotide variation among sequences. The population structure and demographic analyses showed discrepancies between markers, suggesting male-biased dispersal in the species. The time-calibrated COI phylogenetic inference showed a recent diversification of lineages (Middle/Late Pleistocene), while the paleoclimate modeling indicated niche stability since ~120 Kya. The ABC results agreed with the niche models, supporting a panmictic population as the most likely historical scenario for the species. These results indicate that A. omnicolor experienced no niche or population reductions during the Late Pleistocene, despite the intense landscape modifications that occurred in the subtropical region, and that other factors beside LGM and LIG climate oscillations might have contributed to the demographic history of this species. This pattern may be related to the high dispersal ability and wide environmental tolerance of A. omnicolor, highlighting the need for more phylogeographical studies with invertebrates and other generalist taxa, in order to understand the effects of Quaternary climate changes on Neotropical biodiversity.     

A 21 000 cal years paleoclimatic record from Caçó Lake,northern Brazil: evidence from sedimentary and pollen analyses

Sedimentological studies including seismic profiles, mineralogy and organic geochemistry on two cores from the center and margin of Caçó Lake, Maranhão State, northern Brazil, revealed variable climatic and environmental conditions during the last 21 cal kyr BP. Between 21 and 17 cal kyr BP, during the Late Glacial Maximum, regional climate was predominantly dry, interrupted by short humid phases, as reflected by a succession of very thin layers of sand and organic matter. The Late Pleistocene climate was relatively wet as is suggested by rapid lake-level rise and forest expansion. The Late Pleistocene humid climate differed significantly from present conditions. We suggest that Late Glacial humid conditions were the consequence of intensification of the Inter-Tropical Convergence Zone or shifts of its position, resulting in Antarctic cold-front occurrences. The abrupt climatic changes during this period were marked by siderite deposition into Caçó Lake, which appears to be related to regional hydrologic changes linked to global/Northern Hemisphere events. The Holocene was characterized by lower moisture availability and a distinct dry period until 7 cal kyr BP, in response to South American insolation conditions.     

西伯利亚南部旧石器时代晚期哺乳动物群动态学

旧石器时代晚期至全新世发生了广泛而迅速的动物群更替,为研究动物群对快速环境变化的反应提供了有益的信息。自中新世以来中亚地区一直是欧亚大陆向更开阔和干旱环境转变的焦点。利用已发表的俄罗斯贝加尔湖和阿尔泰山区域旧石器时代晚期的文献资料,通过对大型食草动物臼齿齿冠高度、食性和体型大小的分析,绘制了环境变化图。结果表明,在旧石器时代最晚期(晚冰期),平均齿冠高度、平均体型大小均有所减小,食物取向从食草向食树叶转变,这意味着气候变得温暖,降雨量增加,森林也有可能扩展。常见种(出现于超过25%的地点)更强烈地表现出这一趋势。齿冠高度随海拔高度而增大,提示降雨量减少。从齿冠高度得出的降雨量数值与根据其他指标所估算的数值之间的偏差可能随海拔高度而增大。影响西伯利亚南部猛犸象动物群消亡的主要变化可能开始于末次冰期冰盛期(LGM)之后。猛犸象和奇蹄动物出现频率的降低与大型哺乳动物的绝灭相一致,它们和偶蹄类的差异则反映了气候的变化。人类的过度开发可能给区域环境带来了更大的压力,使得哺乳动物的种群数量下降,导致它们更迅速地绝灭。     

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.     

Ancient islands acted as refugia and pumps for conifer diversity

Island species are thought to be extinction‐prone because of small population sizes, restricted geographical distribution and limited dispersal ability. However, the topographical and environmental heterogeneity, geographical isolation and stability of islands over long timescales could create refugia for taxa whose source area is threatened by environmental changes. We address this possibility by inferring the evolution of the New Caledonia (NC) and New Zealand (NZ) conifer diversity, which represents over 10% of the world's diversity for this group. We estimate speciation and extinction rates in relation to the presence/absence on these islands, and dispersal rates between the islands and surrounding areas. We also test the Eocene submersion of NC and the Oligocene drowning of NZ by comparing the fit of biogeographical scenarios using ancestral area estimations. We find that extinction rates were significantly lower for island species, and dispersal “out of islands” was higher. A model including a diversification shift when NC emerged better explains the diversification dynamics. Biogeographical analyses corroborate that conifers experienced high continental extinctions, but survived on islands. NC and NZ have thus contributed to the world's conifer diversity as “island refugia”, by maintaining early‐diverging lineages from continents during environmental changes on continents. These ancient islands also acted as “species pumps”, providing species into adjacent areas. Our study highlights the important but neglected role of islands in promoting the evolution and conservation of biodiversity.     

Patterns of resource use in early Homo and Paranthropus

Conventional wisdom concerning the extinction of Paranthropus suggests that these species developed highly derived morphologies as a consequence of specializing on a diet consisting of hard and/or low-quality food items. It goes on to suggest that these species were so specialized or stenotopic that they were unable to adapt to changing environments in the period following 1.5 Ma. The same conventional wisdom proposes that early Homo species responded very differently to the same environmental challenges. Instead of narrowing their niche it was the dietary and behavioral flexibility (eurytopy) exhibited by early Homo that enabled that lineage to persist. We investigate whether evidence taken across eleven criteria supports a null hypothesis in which Paranthropus is more stenotopic than early Homo. In six instances (most categories of direct evidence of dietary breadth, species diversity, species duration, susceptibility to dispersal, dispersal direction, and non-dietary adaptations) the evidence is inconsistent with the hypothesis. Only one line of indirect evidence for dietary breadth-occlusal morphology-is unambiguously consistent with the null hypothesis that Paranthropus' ability to process tough, fibrous food items (e.g., leaves) was reduced relative to early Homo. Other criteria (habitat preference, population density, direct and indirect evidence of dietary breadth related to incisor use) are only consistent with the hypothesis under certain conditions. If those conditions are not met, then the evidence is either inconsistent with the hypothesis, or ambiguous. On balance, Paranthropus and early Homo were both likely to have been ecological generalists. These data are inconsistent with the conventional wisdom that stenotopy was a major contributing factor in the extinction of the Paranthropus clade. Researchers will need to explore other avenues of research in order to generate testable hypotheses about the demise of Paranthropus. Ecological models that may explain the evolution of eurytopy in early hominins are discussed.     

The effect of selection history on extinction risk during severe environmental change

Environments rarely remain the same over time, and populations are therefore frequently at risk of going extinct when changes are significant enough to reduce fitness. Although many studies have investigated what attributes of the new environments and of the populations experiencing these changes will affect their probability of going extinct, limited work has been directed towards determining the role of population history on the probability of going extinct during severe environmental change. Here, we compare the extinction risk of populations with a history of selection in a benign environment, to populations with a history of selection in one or two stressful environments. We exposed spores and lines of the green alga Chlamydomonas reinhardtii from these three different histories to a range of severe environmental changes. We found that the extinction risk was higher for populations with a history of selection in stressful environments compared to populations with a history of selection in a benign environment. This effect was not due to differences in initial population sizes. Finally, the rates of extinction were highly repeatable within histories, indicating strong historical contingency of extinction risk. Hence, information on the selection history of a population can be used to predict their probability of going extinct during environmental change.     

Dispersal provided resilience to range collapse in a marine mammal: insights from the past to inform conservation biology

Population loss is often a harbinger of species extinction, but few opportunities exist to follow a species’ demography and genetics through both time and space while this occurs. Previous research has shown that the northern fur seal (Callorhinus ursinus) was extirpated from most of its range over the past 200–800 years and that some of the extirpated populations had unique life history strategies. In this study, widespread availability of subfossils in the eastern Pacific allowed us to examine temporal changes in spatial genetic structure during massive population range contraction and partial recovery. We sequenced the mitochondrial control region from 40 ancient and 365 modern samples and analyzed them through extensive simulations within a serial Approximate Bayesian Computation framework. These analyses suggest that the species maintained a high abundance, probably in subarctic refugia, that dispersal rates are likely 85% per generation into new breeding colonies, and that population structure was not higher in the past. Despite substantial loss of breeding range, this species’ high dispersal rates and refugia appear to have prevented a loss of genetic diversity. High dispersal rates also suggest that previous evidence for divergent life history strategies in ancient populations likely resulted from behavioral plasticity. Our results support the proposal that panmictic, or nearly panmictic, species with large ranges will be more resilient to future disturbance and environmental change. When appropriately verified, evidence of low population structure can be powerful information for conservation decision‐making.     

Spatial insurance in multi‐trophic metacommunities

Metacommunity theory suggests that dispersal is a key driver of diversity and ecosystem functioning in changing environments. The capacity of dispersal to mitigate effects of environmental change might vary among trophic groups, potentially resulting in changes in trophic interactions and food web structure. In a mesocosm experiment, we compared the compositional response of bacteria, phyto‐ and zooplankton to a factorial manipulation of acidification and dispersal. We found that the buffering capacity of dispersal varied among trophic groups: dispersal alleviated the negative effect of acidification on phytoplankton diversity mid‐experiment, but had no effect on the diversity of zooplankton and bacteria. Likewise, trophic groups differed in whether dispersal facilitated compositional change. Dispersal accelerated changes in phytoplankton composition under acidification, possibly mediated by changes in trophic interactions, but had no effect on the composition of zooplankton and bacteria. Overall, our results suggest that the potential for spatial insurance can vary among trophic groups.     

Exploring the major depletions of conodont diversity during the Triassic

In this paper, we show that the Triassic fossil record reflects just two great depletions of conodont diversity before the Rhaetian, which occurred in the Smithian (Olenekian, Early Triassic) and in the Julian (Carnian, Late Triassic). By exploring this context, our results highlighted that they respond to different origination–extinction dynamics. Thus, while the Smithian diversity depletion can be interpreted as a consequence of elevated extinction, the Julian diversity depletion was triggered by fluctuations in origination regime. This evidence suggests that, despite the role of extinction on diversity losses, conodonts suffered crucial changes on the origination regimes during the Late Triassic which triggered these events. Notwithstanding, our results indicate that the end-Triassic diversity depletion of conodonts was produced by background extinction levels in a context of lower origination. This suggests that several biological factors, rather than a unique, environmental and/or cyclic cause, could have influenced the evolutionary history of conodonts during the Triassic.     

Late Permian marine ecosystem collapse began in deeper waters: evidence from brachiopod diversity and body size changes

Analysis of Permian–Triassic brachiopod diversity and body size changes from different water depths spanning the continental shelf to basinal facies in South China provides insights into the process of environmental deterioration. Comparison of the temporal changes of brachiopod diversity between deepwater and shallow‐water facies demonstrates that deepwater brachiopods disappeared earlier than shallow‐water brachiopods. This indicates that high environmental stress commenced first in deepwater settings and later extended to shallow waters. This environmental stress is attributed to major volcanic eruptions, which first led to formation of a stratified ocean and a chemocline in the outer shelf and deeper water environments, causing the disappearance of deep marine benthos including brachiopods. The chemocline then rapidly migrated upward and extended to shallow waters, causing widespread mass extinction of shallow marine benthos. We predict that the spatial and temporal patterns of earlier onset of disappearance/extinction and ecological crisis in deeper water ecosystems will be recorded during other episodes of rapid global warming.     

Fragmentation and the context-dependence of dispersal syndromes: matrix harshness modifies resident-disperser phenotypic differences in microcosms

Habitat fragmentation, the conversion of landscapes into patchy habitats separated by unsuitable environments, is expected to reduce dispersal among patches. However, its effects on dispersal should depend on dispersal syndromes, i.e. how dispersal covaries with phenotypic traits, because these syndromes can drastically alter dispersal and subsequent ecological and evolutionary dynamics. Our comprehension of whether environmental factors such as habitat fragmentation generate and/or modify dispersal syndromes (i.e. conditional dispersal syndromes) is therefore key for biodiversity forecasting. Here we tested whether habitat fragmentation modulates dispersal syndromes by experimentally manipulating matrix harshness, a critical feature of habitat fragmentation, in ciliate microcosms. We found evidence for dispersal syndromes involving multiple traits linked to morphology (elongation and size), movement (velocity and linearity) and demography (growth rate and maximal population density). More importantly, these syndromes were modified by matrix harshness, with increased differences between residents and dispersers in morphology and movement traits, and decreased differences in growth rate as the matrix became increasingly harsh. Our findings thus reveal that habitat fragmentation can mediate the intensity and form of dispersal syndromes, a context-dependence that could have important consequences for ecological and evolutionary dynamics under environmental changes.     

Dispersal and diversity in the earliest North American sauropodomorph dinosaurs, with a description of a new taxon

Sauropodomorph dinosaurs originated in the Southern Hemisphere in the Middle or Late Triassic and are commonly portrayed as spreading rapidly to all corners of Pangaea as part of a uniform Late Triassic to Early Jurassic cosmopolitan dinosaur fauna. Under this model, dispersal allegedly inhibited dinosaurian diversification, while vicariance and local extinction enhanced it. However, apomorphy-based analyses of the known fossil record indicate that sauropodomorphs were absent in North America until the Early Jurassic, reframing the temporal context of their arrival. We describe a new taxon from the Kayenta Formation of Arizona that comprises the third diagnosable sauropodomorph from the Early Jurassic of North America. We analysed its relationships to test whether sauropodomorphs reached North America in a single sweepstakes event or in separate dispersals. Our finding of separate arrivals by all three taxa suggests dispersal as a chief factor in dinosaurian diversification during at least the early Mesozoic. It questions whether a 'cosmopolitan' dinosaur fauna ever existed, and corroborates that vicariance, extinction and dispersal did not operate uniformly in time or under uniform conditions during the Mesozoic. Their relative importance is best measured in narrow time slices and circumscribed geographical regions.     

Optimal life‐history schedule in a metapopulation with juvenile dispersal

Previous models have predicted that when mortality increases with age, older individuals should invest more of their resources in reproduction and produce less dispersive offspring, as both their future reproductive value and their prospect of competing with their own sib decline. Those models assumed stable population sizes. We here study for the first time the evolution of age‐specific reproductive effort and of age‐specific offspring dispersal rate in a metapopulation with extinction‐recolonization dynamics and juvenile dispersal. Our model explores the evolutionary consequences of disequilibrium in the age structure of individuals in local populations, generated by disturbances. Life‐history decisions are then shaped both by changes with age in individual performances, and by changes in ecological conditions, as young and old individuals do not live on average in the same environments. Lower juvenile dispersal favours the evolution of higher reproductive effort in young adults in a metapopulation with extinction‐recolonization compared with a well‐mixed population. Contrary to previous predictions for stable structured populations, we find that offspring dispersal should generally increase with maternal age. This is because young individuals, who are overrepresented in recently colonized populations, should allocate more to reproduction and less to dispersal as a strategy to exploit abundant recruitment opportunities in such populations.     

Demographic estimates from the Palaeolithic–Mesolithic boundary in Scandinavia: comparative benchmarks and novel insights

Prehistoric demography has recently risen to prominence as a potentially explanatory variable for episodes of cultural change as documented in the archaeological and ethnographic record. While this has resulted in a veritable boom in methodological developments seeking to address temporal changes in the relative size of prehistoric populations, little work has focused on the manner in which population dynamics manifests across a spatial dimension. Most recently, the so-called Cologne Protocol has led the way in this endeavour. However, strict requirements of raw-material exchange data as analytical inputs have prevented further applications of the protocol to regions outside of continental Europe. We apply an adjusted approach of the protocol that makes it transferable to cases in other parts of the world, while demonstrating its use by providing comparative benchmarks of previous research on the Late Glacial Final Palaeolithic of southern Scandinavia, and novel insights from the early Holocene pioneer colonization of coastal Norway. We demonstrate again that population size and densities remained fairly low throughout the Late Glacial, and well into the early Holocene. We suggest that such low population densities have played a significant role in shaping what may have been episodes of cultural loss, as well as potentially longer periods of only relatively minor degrees of cultural change.This article is part of the theme issue ‘Cross-disciplinary approaches to prehistoric demography’.     

Rainfall seasonality predicts the germination behavior of a tropical dry‐forest vine

Seed dormancy is considered to be an adaptive strategy in seasonal and/or unpredictable environments because it prevents germination during climatically favorable periods that are too short for seedling establishment. Tropical dry forests are seasonal environments where seed dormancy may play an important role in plant resilience and resistance to changing precipitation patterns. We studied the germination behavior of seeds from six populations of the Neotropical vine Dalechampia scandens (Euphorbiaceae) originating from environments of contrasting rainfall seasonality. Seeds produced by second greenhouse‐generation plants were measured and exposed to a favorable wet environment at different time intervals after capsule dehiscence and seed dispersal. We recorded the success and the timing of germination. All populations produced at least some dormant seeds, but seeds of populations originating from more seasonal environments required longer periods of after‐ripening before germinating. Within populations, larger seeds tended to require longer after‐ripening periods than did smaller seeds. These results indicate among‐population genetic differences in germination behavior and suggest that these populations are adapted to local environmental conditions. They also suggest that seed size may influence germination timing within populations. Ongoing changes in seasonality patterns in tropical dry forests may impose strong selection on these traits.     

Late glacial and Holocene environmental changes inferred from sediments in Lake Myklevatnet,Nordfjord, western Norway

Late Glacial and Holocene environmental changes were reconstructed using physical, chemical and biological proxies in Lake Myklevatnet, Allmenningen, (5º13′17″E, 61º55′13″N) located at the northern side of Nordfjorden at the coast of western Norway. Myklevatnet (123 m a.s.l.) lies above the Late Glacial marine limit and contains sediments back to approximately 14,300 years before a.d. 2000 (b2k). Because the lake is located ~48 km beyond the margin of the Younger Dryas (YD) fjord and valley glaciers further inland, and did not receive glacier meltwater from local glaciers during the YD, the lake record provides supplementary information to Lake Kråkenes that received glacial meltwater from a local YD glacier. Lake Myklevatnet has a small catchment and is sensitive to Late Glacial and Holocene climate and environmental changes in the coastal region of western Norway. The age-depth relationship was inferred from a radiocarbon- and tephra-based smoothing-spline model with correlated ages from oxygen isotope maxima and minima in the Late Glacial sequence of the NGRIP ice core (in years b2k) to refine the basal chronology in the Myklevatnet record. The results indicate a two-step YD warming, colder early YD temperatures than in the later part of the YD, and considerably more climate and environmental variability during the late Holocene in western Norway than recorded previously in the oxygen isotopes from Greenland ice cores. The Myklevatnet record is also compared with other Late Glacial and Holocene terrestrial and marine proxy reconstructions in the North Atlantic realm.