North American pollen records provide evidence for macroscale ecological changes in the Anthropocene.

Recent global changes associated with anthropogenic activities are impacting ecological systems globally, giving rise to the Anthropocene. Critical reorganization of biological communities and biodiversity loss are expected to accelerate as anthropogenic global change continues. Long-term records offer context for understanding baseline conditions and those trajectories that are beyond the range of normal fluctuation seen over recent millennia: Are we causing changes that are fundamentally different from changes in the past? Using a rich dataset of late Quaternary pollen records, stored in the open-access and community-curated Neotoma database, we analyzed changes in biodiversity and community composition since the end Pleistocene in North America. We measured taxonomic richness, short-term taxonomic loss and gain, first/last appearances (FAD/LAD), and abrupt community change. For all analyses, we incorporated age-model uncertainty and accounted for differences in sample size to generate conservative estimates. The most prominent signals of elevated vegetation change were seen during the Pleistocene-Holocene transition and since 200 calendar years before present (cal YBP). During the Pleistocene-Holocene transition, abrupt changes and FADs were elevated, and from 200 to -50 cal YBP, we found increases in short-term taxonomic loss, FADs, LADs, and abrupt changes. Taxonomic richness declined from ~13,000 cal YBP until about 6,000 cal YBP and then increased until the present, reaching levels seen during the end Pleistocene. Regionally, patterns were highly variable. These results show that recent changes associated with anthropogenic impacts are comparable to the landscape changes that took place as we moved from a glacial to interglacial world.

The Dmanisi Equus: Systematics, biogeography, and paleoecology.

The Equus datum has been established as a geochronologic 'instantaneous' migratory event of a North American Equus species into Eurasia at the beginning of the Pleistocene (2.58 Ma). A remarkable radiation of Equus followed across Eurasia and Africa. Dmanisi includes excellent remains of Equus, well calibrated between 1.85 and 1.76 Ma. Our morphologic and morphometric analyses of the augmented Dmanisi Equus sample support the co-occurrence of Equus stenonis and Equus altidens in the sequence. Dmanisi E. stenonis is found to be morphologically similar to the European E. stenonis populations and represents the best well-dated easternmost occurrence of this species in Eurasia. The Dmanisi E. altidens represents the oldest well-calibrated occurrence of this species in Western Eurasia. Our analyses demonstrate that E. altidens extended its range westward from west Asia to Greece, Germany, Italy, Spain, and possibly France. Our results do not support distinguishing multiple subspecies of E. altidens, including E. altidens altidens, E. altidens granatensis and E. stenonis mygdoniensis. The Dmanisi cranial and postcranial samples exhibit morphologies close both to extant hemiones and zebras. Equus altidens is believed to have been well adapted to newly emergent arid environments in western Eurasia during the late Early and early Middle Pleistocene. The first occurrence of E. altidens at Dmanisi marks an important turnover in the horse communities of the late Early Pleistocene, with a dispersion of this species from West Asia to West Europe ca. 1.8 Ma.

Ancient plant DNA reveals High Arctic greening during the Last Interglacial.

Summer warming is driving a greening trend across the Arctic, with the potential for large-scale amplification of climate change due to vegetation-related feedbacks [Pearson et al., Nat. Clim. Chang. (3), 673-677 (2013)]. Because observational records are sparse and temporally limited, past episodes of Arctic warming can help elucidate the magnitude of vegetation response to temperature change. The Last Interglacial ([LIG], 129,000 to 116,000 y ago) was the most recent episode of Arctic warming on par with predicted 21st century temperature change [Otto-Bliesner et al., Philos. Trans. A Math. Phys. Eng. Sci. (371), 20130097 (2013) and Post et al., SciAdv (5), eaaw9883 (2019)]. However, high-latitude terrestrial records from this period are rare, so LIG vegetation distributions are incompletely known. Pollen-based vegetation reconstructions can be biased by long-distance pollen transport, further obscuring the paleoenvironmental record. Here, we present a LIG vegetation record based on ancient DNA in lake sediment and compare it with fossil pollen. Comprehensive plant community reconstructions through the last and current interglacial (the Holocene) on Baffin Island, Arctic Canada, reveal coherent climate-driven community shifts across both interglacials. Peak LIG warmth featured a ∼400-km northward range shift of dwarf birch, a key woody shrub that is again expanding northward. Greening of the High Arctic-documented here by multiple proxies-likely represented a strong positive feedback on high-latitude LIG warming. Authenticated ancient DNA from this lake sediment also extends the useful preservation window for the technique and highlights the utility of combining traditional and molecular approaches for gleaning paleoenvironmental insights to better anticipate a warmer future.

The last meal of an Eocene pollen-feeding fly.

One of the most important trophic interactions today is that between insects and their floral hosts. This biotic association is believed to have been critical to the radiation of flowering plants and many pollinating insect lineages over the last 120 million years (Ma). Trophic interactions among fossil organisms are challenging to study, and most inferences are based on indirect evidence. Fossil records providing direct evidence for pollen feeding, i.e., fossil stomach and gut contents, are exceptionally rare.1,2 Such records have the potential to provide information on aspects of animal behavior and ecology as well as plant-animal interactions that are sometimes not yet recognized for their extant relatives. The dietary preferences of short-proboscid nemestrinids are unknown, and pollinivory has not been recorded for extant Nemestrinidae.3 We analyzed the contents of the conspicuously swollen abdomen of an ca. 47.5 Ma old nemestrinid fly of the genus Hirmoneura from Messel, Germany, with photogrammetry and state-of-the-art palynological methods. The fly fed on pollen from at least four plant families-Lythraceae, Vitaceae, Sapotaceae, and Oleaceae-and presumably pollinated flowers of two extant genera, Decodon and Parthenocissus. We interpret the feeding and foraging behavior of the fly, reconstruct its preferred habitat, and conclude about its pollination role and importance in paratropical environments. This represents the first evidence that short-proboscid nemestrinid flies fed, and possibly feed to this day, on pollen, demonstrating how fossils can provide vital information on the behavior of insects and their ecological relationships with plants.

Coping with arid environments: A critical threshold for human expansion in Europe at the Marine Isotope Stage 12/11 transition? The case of the Iberian Peninsula.

Archaeological remains have highlighted the fact that the interglacial Marine Isotope Stage (MIS) 11 was a threshold from the perspective of hominin evolution in Europe. After the MIS 12 glaciation, considered one of the major climate-driven crises experienced by hominins, the archaeological records show an increasing number of occupations, evidence of new subsistence behaviors, and significant technical innovations. Here, we used statistical and geographic techniques to analyze the amphibian- and reptile-based paleoclimate and habitat reconstructions generated from a large data set of the Iberian Peninsula to (1) investigate if temperature, precipitation, and/or forest cover may have impacted the hominin occupation of the territory during the Early and Middle Pleistocene, (2) propose an 'Iberian' ecological model before and after the MIS 12/11 transition, and (3) evaluate, based on this model, the potential hominin occupation at a European scale. The results indicate the existence of climatic constraints on human settlement related to rainfall and environmental humidity. The Early Pleistocene and the first half of the Middle Pleistocene are dominated by the occupation of relatively humid wooded areas, whereas during the second part of the Middle Pleistocene, a broadening of the earlier ecological niche is clearly observed toward the occupation of more open arid areas. Based on the estimated occupational niche for hominins, a maximum potential distribution for early hominins is proposed in Europe before and after 426 ka. Results also indicate that parts of the Iberian Peninsula may not have been suitable for early hominin occupation. Our ecological model is consistent with the pattern of hominin occupation observed in northern and central Europe, where the earliest evidence reflects only pioneering populations merely extending their ranges in response to the expansion of their preferred habitats, as compared with a more sustained occupation by 400 ka.

Community Assembly and Climate Mismatch in Late Quaternary Eastern North American Pollen Assemblages.

Plant community response to climate change ranges from synchronous tracking to strong mismatch. Explaining this variation in climate change response is critical for accurate global change modeling. Here we quantify how closely assemblages track changes in climate (match/mismatch) and how broadly climate niches are spread within assemblages (narrow/broad ecological tolerance, or "filtering") using data for the past 21,000 years for 531 eastern North American fossil pollen assemblages. Although climate matching has been strong over the last 21 millennia, mismatch increased in 30% of assemblages during the rapid climate shifts between 14.5 and 10 ka. Assemblage matching rebounded toward the present day in 10%-20% of assemblages. Climate-assemblage mismatch was greater in tree-dominated and high-latitude assemblages, consistent with persisting populations, slower dispersal rates, and glacial retreat. In contrast, climate matching was greater for assemblages comprising taxa with higher median seed mass. More than half of the assemblages were climatically filtered at any given time, with peak filtering occurring at 8.5 ka for nearly 80% of assemblages. Thus, vegetation assemblages have highly variable rates of climate mismatch and filtering over millennial scales. These climate responses can be partially predicted by species' traits and life histories. These findings help constrain predictions for plant community response to contemporary climate change.

Pollination of Cretaceous flowers.

Insect pollination of flowering plants (angiosperms) is responsible for the majority of the world's flowering plant diversity and is key to the Cretaceous radiation of angiosperms. Although both insects and angiosperms were common by the mid-Cretaceous, direct fossil evidence of insect pollination is lacking. Direct evidence of Cretaceous insect pollination is associated with insect-gymnosperm pollination. Here, we report a specialized beetle-angiosperm pollination mode from mid-Cretaceous Burmese amber (99 mega-annum [Ma]) in which a tumbling flower beetle (Mordellidae), Angimordella burmitina gen. et sp. nov., has many tricolpate pollen grains attached. A. burmitina exhibits several specialized body structures for flower-visiting behavior including its body shape and pollen-feeding mouthparts revealed by X-ray microcomputed tomography (micro-CT). The tricolpate pollen in the amber belongs to the eudicots that comprise the majority of extant angiosperm species. These pollen grains exhibit zoophilous pollination attributes including their ornamentation, size, and clumping characteristics. Tricolpate pollen grains attached to the beetle's hairs are revealed by confocal laser scanning microscopy, which is a powerful tool for investigating pollen in amber. Our findings provide direct evidence of insect pollination of Cretaceous angiosperms, extending the range insect-angiosperm pollination association by at least 50 million years. Our results support the hypothesis that specialized insect pollination modes were present in eudicots 99 million years ago.

Beetle Pollination of Cycads in the Mesozoic.

Cycads, unlike modern wind-pollinated conifers and Ginkgo, are unusual in that they are an ancient group of gymnosperms pollinated by insects [1-3]. Although it is well documented that cycads were diverse and abundant during the mid-Mesozoic, little is known about their biogeography and pollination before the rise of angiosperms. Direct fossil evidence illuminating the evolutionary history of cycads is extremely rare [4, 5]. Here we report a specialized beetle-mediated pollination mode from the mid-Cretaceous of Myanmar, wherein a new boganiid beetle, Cretoparacucujus cycadophilus, with specialized pollen-feeding adaptations in its mouthparts and legs, was associated with many pollen grains of Cycadopites. Phylogenetic analyses indicate Cretoparacucujus as a sister group to the extant Australian Paracucujus, which pollinate the cycad Macrozamia riedlei. Our discovery, along with the current disjunct distribution of related beetle-herbivore (tribe Paracucujini) and cycad-host (tribe Encephalarteae) pairs in South Africa and Australia, indicate a probable ancient origin of beetle pollination of cycads at least in the Early Jurassic, long before angiosperm dominance and the radiation of flowering-plant pollinators later in the Cretaceous.

Using Paleoecology to Inform Land Management as Climates Change: An Example from an Oak Savanna Ecosystem.

Oak savanna, a transitional ecosystem between open prairie and dense oak forest, was once widespread in Minnesota. Upon European settlement much of the oak savanna was destroyed. Recently, efforts to restore this ecosystem have increased and often include the reintroduction of fire. Though fire is known to serve an important role within oak savannas, there are currently few studies which address fire regimes on timescales longer than the last century. This research presents a paleoecological history of Sherburne National Wildlife Refuge (SNWR) in MN, USA, spanning the last ~8000 years. The objectives of this study were to use charcoal, pollen, and magnetic susceptibility of lake sediments collected from Johnson Slough (JS) within the refuge to evaluate the natural range of variability and disturbance history of the oak savanna within the refuge, assess the success of current restoration strategies, and add to the regional paleoecological history. The mid/late Holocene period of the JS record shows a period of high fire activity from ca. 6500 to 2600 cal year BP, with a shift from prairie to oak savanna occurring over this same period. A (possibly agricultural) disturbance to JS sediments affected the period from ca. 2600 cal year BP to 1963 AD, which includes the time of Euro-American settlement. However, the destruction and subsequent restoration of the oak savanna is evident in a pollen ratio of Quercus:Poaceae, indicating that current restoration efforts have been successful at restoring the oak savanna to within the natural range of variability seen just prior to destruction.

False Blister Beetles and the Expansion of Gymnosperm-Insect Pollination Modes before Angiosperm Dominance.

During the mid-Cretaceous, angiosperms diversified from several nondiverse lineages to their current global domination [1], replacing earlier gymnosperm lineages [2]. Several hypotheses explain this extensive radiation [3], one of which involves proliferation of insect pollinator associations in the transition from gymnosperm to angiosperm dominance. However, most evidence supports gymnosperm-insect pollinator associations, buttressed by direct evidence of pollen on insect bodies, currently established for four groups: Thysanoptera (thrips), Neuroptera (lacewings), Diptera (flies), and now Coleoptera (beetles). Each group represents a distinctive pollination mode linked to a unique mouthpart type and feeding guild [4-9]. Extensive indirect evidence, based on specialized head and mouthpart morphology, is present for one of these pollinator types, the long-proboscid pollination mode [10], representing minimally ten family-level lineages of Neuroptera, Mecoptera (scorpionflies), and Diptera [8, 10, 11]. A recurring feature uniting these pollinator modes is host associations with ginkgoalean, cycad, conifer, and bennettitalean gymnosperms. Pollinator lineages bearing these pollination modes were categorized into four evolutionary cohorts during the 35-million-year-long angiosperm radiation, each defined by its host-plant associations (gymnosperm or angiosperm) and evolutionary pattern (extinction, continuation, or origination) during this interval [12]. Here, we provide the first direct evidence for one cohort, exemplified by the beetle Darwinylus marcosi, family Oedemeridae (false blister beetles), that had an earlier gymnosperm (most likely cycad) host association, later transitioning onto angiosperms [13]. This association constitutes one of four patterns explaining the plateau of family-level plant lineages generally and pollinating insects specifically during the mid-Cretaceous angiosperm radiation [12].

Community ecology in a changing environment: Perspectives from the Quaternary.

Community ecology and paleoecology are both concerned with the composition and structure of biotic assemblages but are largely disconnected. Community ecology focuses on existing species assemblages and recently has begun to integrate history (phylogeny and continental or intercontinental dispersal) to constrain community processes. This division has left a "missing middle": Ecological and environmental processes occurring on timescales from decades to millennia are not yet fully incorporated into community ecology. Quaternary paleoecology has a wealth of data documenting ecological dynamics at these timescales, and both fields can benefit from greater interaction and articulation. We discuss ecological insights revealed by Quaternary terrestrial records, suggest foundations for bridging between the disciplines, and identify topics where the disciplines can engage to mutual benefit.

Early origin of parental care in Mesozoic carrion beetles.

The reconstruction and timing of the early stages of social evolution, such as parental care, in the fossil record is a challenge, as these behaviors often do not leave concrete traces. One of the intensely investigated examples of modern parental care are the modern burying beetles (Silphidae: Nicrophorus), a lineage that includes notable endangered species. Here we report diverse transitional silphids from the Mesozoic of China and Myanmar that provide insights into the origins of parental care. Jurassic silphids from Daohugou, sharing many defining characters of Nicrophorinae, primitively lack stridulatory files significant for parental care communications; although morphologically similar, Early Cretaceous nicrophorines from the Jehol biota possess such files, indicating that a system of parental care had evolved by this early date. More importantly, burying beetles of the genus Nicrophorus have their earliest first record in mid-Cretaceous Burmese amber, and document early evolution of elaborate biparental care and defense of small vertebrate carcasses for their larvae. Parental care in the Early Cretaceous may have originated from competition between silphids and their predators. The rise of the Cretaceous Nicrophorinae implies a biology similar to modern counterparts that typically feed on carcasses of small birds and mammals.

Environmental impact of geometric earthwork construction in pre-Columbian Amazonia.

There is considerable controversy over whether pre-Columbian (pre-A.D. 1492) Amazonia was largely "pristine" and sparsely populated by slash-and-burn agriculturists, or instead a densely populated, domesticated landscape, heavily altered by extensive deforestation and anthropogenic burning. The discovery of hundreds of large geometric earthworks beneath intact rainforest across southern Amazonia challenges its status as a pristine landscape, and has been assumed to indicate extensive pre-Columbian deforestation by large populations. We tested these assumptions using coupled local- and regional-scale paleoecological records to reconstruct land use on an earthwork site in northeast Bolivia within the context of regional, climate-driven biome changes. This approach revealed evidence for an alternative scenario of Amazonian land use, which did not necessitate labor-intensive rainforest clearance for earthwork construction. Instead, we show that the inhabitants exploited a naturally open savanna landscape that they maintained around their settlement despite the climatically driven rainforest expansion that began ∼2,000 y ago across the region. Earthwork construction and agriculture on terra firme landscapes currently occupied by the seasonal rainforests of southern Amazonia may therefore not have necessitated large-scale deforestation using stone tools. This finding implies far less labor--and potentially lower population density--than previously supposed. Our findings demonstrate that current debates over the magnitude and nature of pre-Columbian Amazonian land use, and its impact on global biogeochemical cycling, are potentially flawed because they do not consider this land use in the context of climate-driven forest-savanna biome shifts through the mid-to-late Holocene.

Use of pollen and ancient DNA as conservation baselines for offshore islands in New Zealand.

Islands play a key role globally in the conservation of endemic species. Many island reserves have been highly modified since human colonization, and their restoration and management usually occur without knowledge of their prehuman state. However, conservation paleoecology is increasingly being recognized as a tool that can help to inform both restoration and conservation of island reserves by providing prehuman vegetation baselines. Many of New Zealand's mammal-free offshore islands are foci for biological diversity conservation and, like many islands in the Polynesian region, were deforested following initial human settlement. Therefore, their current restoration, replanting, and management are guided either by historic vegetation descriptions or the occurrence of species on forested islands. We analyzed pollen and ancient DNA in soil cores from an offshore island in northern New Zealand. The result was a 2000-year record of vegetation change that began >1200 years before human settlement and spanned 550 years of human occupation and 180 years of forest succession since human occupation ceased. Between prehuman and contemporary forests there was nearly a complete species turnover including the extirpation of a dominant conifer and a palm tree. The podocarp-dominated forests were replaced by a native but novel angiosperm-dominated forest. There is no modern analog of the prehuman forests on any northern New Zealand island, and those islands that are forested are dominated by angiosperms which are assumed to be climax forests. The pollen and DNA evidence for conifer- and palm-rich forests in the prehuman era challenge this climax forest assumption. Prehuman vegetation records can thus help to inform future restoration of degraded offshore islands by informing the likely rate and direction of successional change; helping to determine whether natural rates of succession are preferable to more costly replanting programs; and providing past species lists if restoration replanting is desired.

Space can substitute for time in predicting climate-change effects on biodiversity.

"Space-for-time" substitution is widely used in biodiversity modeling to infer past or future trajectories of ecological systems from contemporary spatial patterns. However, the foundational assumption--that drivers of spatial gradients of species composition also drive temporal changes in diversity--rarely is tested. Here, we empirically test the space-for-time assumption by constructing orthogonal datasets of compositional turnover of plant taxa and climatic dissimilarity through time and across space from Late Quaternary pollen records in eastern North America, then modeling climate-driven compositional turnover. Predictions relying on space-for-time substitution were ∼72% as accurate as "time-for-time" predictions. However, space-for-time substitution performed poorly during the Holocene when temporal variation in climate was small relative to spatial variation and required subsampling to match the extent of spatial and temporal climatic gradients. Despite this caution, our results generally support the judicious use of space-for-time substitution in modeling community responses to climate change.

Palynological analysis of a late Holocene core from Santo Antônio da Patrulha, Rio Grande do Sul, Southern Brazil

A sedimentar core collected at Santo Antônio da Patrulha, Rio Grande do Sul State, southmost Brazil, was submitted to pollen analysis to provide the vegetational history of this region, and the paleoecological and paleoclimatic changes. A total of 98 taxa of palynomorphs was identified from 35 subsamples. Three radiocarbonic datings were obtained along a section of 115 cm depth, including the basal age of 4730 ± 50 yr BP. Pollen diagrams and cluster analysis were performed based on palynomorphs frequencies, demonstrating five distinct phases (SAP-I to SAP-V), which reflected different paleoecological conditions. The predominance of plants associated with grasslands in the phase SAP-I suggests warm and dry climate conditions. A gradual increasing of humidity conditions was observed mainly from the beginning of the phase SAP-III, when the vegetation set a mosaic of grasslands and Atlantic rainforest. Furthermore, the presence of some forest taxa ( Acacia-type, Daphnopsis racemosa, Erythrina-type and Parapiptadenia rigida-type), from the phase SAP-IV, is interpreted as an influence of the seasonal semideciduous forest in the study region. From the phase SAP-V (ca. 4000 yrs BP), the vegetation became similar to the modern one (extant Atlantic rainforest Biome), especially after 2000 yrs BP (calibrated age).

Um testemunho de sondagem coletado em Santo Antônio da Patrulha, Rio Grande do Sul, Brasil, foi submetido para análise polínica a fim de revelar a história vegetacional e mudanças paleoecológicas e paleoclimáticas. Um total de 98 táxons foi identificado a partir de 35 subamostras. Três datações radiocarbônicas foram obtidas ao longo de uma seção de 115 cm de profundidade, incluindo a idade basal de 4730 ± 50 anos AP. Diagramas polínicos e análises de agrupamentos foramrealizadas com base nas freqüências dos palinomorfos, demonstrando cinco fases distintas (SAP-I a SAP-V), as quais refletiram diferentes condições paleoecológicas. A predominância de plantas relacionadas à vegetação campestre na fase SAP-I sugere condições climáticas quentes e secas. Um gradual aumento nas condições de umidade foi observado principalmente no início da fase SAP-III, quando a vegetação conformou um mosaico de Campos e Floresta Atlântica. Além disso, a presença de certos táxons florestais (tipo- Acacia, Daphnopsis racemosa, tipo Erythrina e tipo Parapiptadenia rigida), a partir da fase SAP-IV, é interpretada como influência da Floresta Estacional Semidecidual na região de estudo. A partir da fase SAP-V (ca. 4000 anos AP) a vegetação tornou-se similar à moderna (atual Bioma da Floresta Atlântica), especialmente após 2000 anos AP (idade calibrada).