Holocene Man in North America: The Ecological Setting and Climatic Background

Abstract

Patterns of human occupation and vegetation are delineated on maps of North America for one thousand year intervals through the Holocene. The raw data for this review include radiocarbon-dated pollen cores and archaeological information and treering records. Dynamic changes in the Laurentide Ice limits and major ecotones are observed through the middle Holocene, when both features reach essentially post-glacial stability. Significant changes in the vegetation boundaries continue to the present, but the scale of change is much diminished.

Early Holocene occupation apparently expanded from Alaska south to California, then east, parallel to the southern boundary of the grasslands, to the Mississippi River and northeastward to the east coast. The absence of occpuation in late-Atlantic time is noted through much of the Great Plains and continued until about 4,000 BP. Within the next millennium, evidence of human occupation virtually covered the United States (except for the northwest) and much of coastal Canada.

Environmental conditions and occupation over North America are reviewed within the framework of Holocene climatic episodes.     

Beringia as a glacial refugium for boreal trees and shrubs: new perspectives from mapped pollen data

Aim Beringia, far north‐eastern Siberia and north‐western North America, was largely unglaciated during the Pleistocene. Although this region has long been considered an ice‐age refugium for arctic herbs and shrubs, little is known about its role as a refugium for boreal trees and shrubs during the last glacial maximum (LGM, c. 28,000–15,000 calibrated years before present). We examine mapped patterns of pollen percentages to infer whether six boreal tree and shrub taxa (Populus, Larix, Picea, Pinus, Betula, Alnus/Duschekia) survived the harsh glacial conditions within Beringia. Methods Extensive networks of pollen records have the potential to reveal distinctive temporal–spatial patterns that discriminate between local‐ and long‐distance sources of pollen. We assembled pollen records for 149 lake, peat and alluvial sites from the Palaeoenvironmental Arctic Sciences database, plotting pollen percentages at 1000‐year time intervals from 21,000 to 6000 calibrated years before present. Pollen percentages are interpreted with an understanding of modern pollen representation and potential sources of long‐distance pollen during the glacial maximum. Inferences from pollen data are supplemented by published radiocarbon dates of identified macrofossils, where available. Results Pollen maps for individual taxa show unique temporal‐spatial patterns, but the data for each taxon argue more strongly for survival within Beringia than for immigration from outside regions. The first increase of Populus pollen percentages in the western Brooks Ranges is evidence that Populus trees survived the LGM in central Beringia. Both pollen and macrofossil evidence support Larix survival in western Beringia (WB), but data for Larix in eastern Beringia (EB) are unclear. Given the similar distances of WB and EB to glacial‐age boreal forests in temperate latitudes of Asia and North America, the widespread presence of Picea pollen in EB and Pinus pollen in WB indicates that Picea and Pinus survived within these respective regions. Betula pollen is broadly distributed but highly variable in glacial‐maximum samples, suggesting that Betula trees or shrubs survived in restricted populations throughout Beringia. Alnus/Duschekia percentages show complex patterns, but generally support a glacial refugium in WB. Main conclusions Our interpretations have several implications, including: (1) the rapid post‐glacial migration rate reported for Picea in western Canada may be over estimated, (2) the expansion of trees and shrubs within Beringia should have been nearly contemporaneous with climatic change, (3) boreal trees and shrubs are capable of surviving long periods in relatively small populations (at the lower limit of detection in pollen data) and (4) long‐distance migration may not have been the predominant mode of vegetation response to climatic change in Beringia.     

Identification of glacial refugia in south-eastern North America by phylogeographical analyses of a forest understorey plant, Trillium cuneatum

Aim We examine several hypotheses emerging from biogeographical and fossil records regarding glacial refugia of a southern thermophilic plant species. Specifically, we investigated the glacial history and post‐glacial colonization of a forest understorey species, Trillium cuneatum. We focused on the following questions: (1) Did T. cuneatum survive the Last Glacial Maximum (LGM) in multiple refugia, and (if so) where were they located, and is the modern genetic structure congruent with the fossil record‐based reconstruction of refugia for mesic deciduous forests? (2) What are the post‐glacial colonization patterns in the present geographical range? Location South‐eastern North America. Methods We sampled 45 populations of T. cuneatum throughout its current range. We conducted phylogeographical analyses based on maternally inherited chloroplast DNA (cpDNA haplotypes) and used TCS software to reconstruct intraspecific phylogeny. Results We detected six cpDNA haplotypes, geographically highly structured into non‐overlapping areas. With one exception, none of the populations had mixed haplotype composition. TCS analysis resulted in two intraspecific cpDNA lineages, with one clade subdivided further by shallower diversification. Main conclusions Our investigation revealed that T. cuneatum survived the LGM in multiple refugia, belonging to two (western, eastern) genealogical lineages geographically structured across south‐eastern North America. The western clade is confined to the south‐western corner of T. cuneatum’s modern range along the Lower Mississippi Valley, where fossil records document a major refugium of mesic deciduous forest. For the eastern clade, modern patterns of cpDNA haplotype distribution suggest cryptic vicariance, in the form of forest contractions and subsequent expansions associated with Pleistocene glacial cycles, rather than simple southern survival and subsequent northward colonization. The north–south partitioning of cpDNA haplotypes was unexpected, suggesting that populations of this rather southern thermophilic species may have survived in more northern locations than initially expected based on LGM climate reconstruction, and that the Appalachian Mountains functioned as a barrier to the dispersal of propagules originating in more southern refugia. Furthermore, our results reveal south‐west to north‐east directionality in historical migration through the Valley and Ridge region of north‐west Georgia.     

Refugial persistence and postglacial recolonization of North America by the cold-tolerant herbaceous plant Orthilia secunda

Previous phylogeographical and palaeontological studies on the biota of northern North America have revealed a complex scenario of glacial survival in multiple refugia and differing patterns of postglacial recolonization. Many putative refugial regions have been proposed both north and south of the ice sheets for species during the Last Glacial Maximum, but the locations of many of these refugia remain a topic of great debate. In this study, we used a phylogeographical approach to elucidate the refugial and recolonization history of the herbaceous plant species Orthilia secunda in North America, which is found in disjunct areas in the west and east of the continent, most of which were either glaciated or lay close to the limits of the ice sheets. Analysis of 596 bp of the chloroplast trnS-trnG intergenic spacer and five microsatellite loci in 84 populations spanning the species' range in North America suggests that O. secunda persisted through the Last Glacial Maximum (LGM) in western refugia, even though palaeodistribution modelling indicated a suitable climate envelope across the entire south of the continent. The present distribution of the species has resulted from recolonization from refugia north and south of the ice sheets, most likely in Beringia or coastal regions of Alaska and British Columbia, the Washington/Oregon region in the northwest USA, and possibly from the region associated with the putative 'ice-free corridor' between the Laurentide and Cordilleran ice sheets. Our findings also highlight the importance of the Pacific Northwest as an important centre of intraspecific genetic diversity, owing to a combination of refugial persistence in the area and recolonization from other refugia.     

The influence of abrupt climate change on the ice‐age vegetation of the Central American lowlands

Aim To investigate the effects of abrupt climate change in the North Atlantic on the vegetation history of lowland Central America. We use palynological evidence from a Central American lake on the Yucatan Peninsula to evaluate the effects of rapid climate changes during the last ice age, between 65 and 8 ka. Location Lake Petén‐Itzá, lowlands of northern Guatemala. Methods Sediment core PI‐6 was sampled at c. 170‐year resolution for pollen and charcoal analysis in order to construct a temporal sequence of environmental change. Uni‐ and multivariate statistical analyses were performed on the pollen dataset to test whether there was an association between Heinrich events in the North Atlantic and vegetation changes in the Central American lowlands. Results Pollen analysis revealed that the composition of plant assemblages on the Yucatan Peninsula varied in phase with rapid changes in North Atlantic climate. Pine savannas were the main vegetation type between c. 60 and 47 ka. These savannas gave way to pine–oak (Pinus–Quercus) forests in the latter half of the last ice age. Marked episodes of replacement of the pine–oak assemblage by xeric‐tolerant taxa occurred during Heinrich events. The Last Glacial Maximum (LGM) was characterized by mesic conditions. Main conclusions The pollen record from Lake Petén‐Itzá showed that vegetation changes associated with Heinrich events were more significant than those associated with the LGM. Each Heinrich event produced a characteristic shift towards xeric taxa. Although colder than Heinrich events, the LGM on the Yucatan Peninsula was relatively moist, and the presumed savannization of the landscape during the maximum cooling of the last glacial was not supported by our data. Our findings suggest alternative scenarios for plant diversification and genetic interchange during glacial times, and also indicate that vegetation in tropical continental settings was not as stable as previously thought.     

Response of tundra ecosystem in southwestern Alaska to Younger-Dryas climatic oscillation

Climatic warming during the last glacial–interglacial transition (LGIT) was punctuated by reversals to glacial‐like conditions. Palaeorecords of ecosystem change can help document the geographical extent of these events and improve our understanding of biotic sensitivity to climatic forcing. To reconstruct ecosystem and climatic variations during the LGIT, we analyzed lake sediments from southwestern Alaska for fossil pollen assemblages, biogenic‐silica content (BSiO₂%), and organic‐carbon content (OC%). Betula shrub tundra replaced herb tundra as the dominant vegetation of the region around 13 600 cal BP (cal BP: ¹⁴C calibrated calendar years before present), as inferred from an increase of Betula pollen percentages from << 5% to >> 20% with associated decreases in Cyperaceae, Poaceae, and Artemisia. At c. 13 000 cal BP, a decrease of Betula pollen from 28 to << 5% suggests that shrub tundra reverted to herb tundra. Shrub tundra replaced herb tundra to resume as the dominant vegetation at 11 600 cal BP. Higher OC% and BSiO₂% values suggest more stable soils and higher aquatic productivity during shrub‐tundra periods than during herb‐tundra periods, although pollen changes lagged behind changes in the biogeochemical indicators before c. 13 000 cal BP. Comparison of our palaeoecological data with the ice‐core dδ¹⁸O record from Greenland reveals strikingly similar patterns from the onset through the termination of the Younger Dryas (YD). This similarity supports the hypothesis that, as in the North Atlantic region, pronounced YD climatic oscillations occurred in the North Pacific region. The rapidity and magnitude of ecological changes at the termination of the YD are consistent with greenhouse experiments and historic photographs demonstrating tundra sensitivity to climatic forcing.     

Multilocus phylogeography and population structure of common eiders breeding in North America and Scandinavia

Aim Glacial refugia during the Pleistocene had major impacts on the levels and spatial apportionment of genetic diversity of species in northern latitude ecosystems. We characterized patterns of population subdivision, and tested hypotheses associated with locations of potential Pleistocene refugia and the relative contribution of these refugia to the post‐glacial colonization of North America and Scandinavia by common eiders (Somateria mollissima). Specifically, we evaluated localities hypothesized as ice‐free areas or glacial refugia for other Arctic vertebrates, including Beringia, the High Arctic Canadian Archipelago, Newfoundland Bank, Spitsbergen Bank and north‐west Norway. Location Alaska, Canada, Norway and Sweden. Methods Molecular data from 12 microsatellite loci, the mitochondrial DNA (mtDNA) control region, and two nuclear introns were collected and analysed for 15 populations of common eiders (n = 716) breeding throughout North America and Scandinavia. Population genetic structure, historical population fluctuations and gene flow were inferred using F‐statistics, analyses of molecular variance, and multilocus coalescent analyses. Results Significant inter‐population variation in allelic and haplotypic frequencies were observed (nuclear DNA F_ST = 0.004–0.290; mtDNA Φ_ST = 0.051–0.927). Whereas spatial differentiation in nuclear genes was concordant with subspecific designations, geographic proximity was more predictive of inter‐population variance in mitochondrial DNA haplotype frequency. Inferences of historical population demography were consistent with restriction of common eiders to four geographic areas during the Last Glacial Maximum: Belcher Islands, Newfoundland Bank, northern Alaska and Svalbard. Three of these areas coincide with previously identified glacial refugia: Newfoundland Bank, Beringia and Spitsbergen Bank. Gene‐flow and clustering analyses indicated that the Beringian refugium contributed little to common eider post‐glacial colonization of North America, whereas Canadian, Scandinavian and southern Alaskan post‐glacial colonization is likely to have occurred in a stepwise fashion from the same glacial refugium. Main conclusions Concordance of proposed glacial refugia used by common eiders and other Arctic species indicates that Arctic and subarctic refugia were important reservoirs of genetic diversity during the Pleistocene. Furthermore, suture zones identified at MacKenzie River, western Alaska/Aleutians and Scandinavia coincide with those identified for other Arctic vertebrates, suggesting that these regions were strong geographic barriers limiting dispersal from Pleistocene refugia.     

Summer vegetation, deglaciation and the anomalous bird diversity gradient in eastern North America

Aim Geographic variation in the species richness of birds has been shown to be strongly associated with annual water and energy levels (actual evapotranspiration, AET) at the global scale. However, the gradient in eastern North America appears to be anomalous, because richness is greatest around the Great Lakes, whereas AET is highest in the south‐eastern US. Here I examine if birds may be responding to vegetation produced during the breeding season rather than to annual production. Location North America east of longitude 98° W. Methods The bird richness pattern was examined using climatic variables, remotely sensed estimates of annual and seasonal plant biomass, and time since areas were exposed by the retreating Laurentide ice sheet from 20,000 to 6000 yr bp . Results Average summer GVI (Global Vegetation Index, derived from NDVI) was found to be positively linearly associated with richness, explaining 82% of the variance, whereas the relationships between richness and annual measures of both AET and GVI were curvilinear. The pattern of retreat of the Laurentide ice sheet explained an additional 6% of the variance in richness, consistent with a previous analysis of Canadian birds. Main conclusions In eastern North America, a seasonal variable associated with plant production explains the diversity gradient rather than the annual measures, but it does not undermine a general conclusion that bird diversity is closely linked with plant biomass. Further, both contemporary and historical factors appear to influence the gradient, and an association between bird richness and the geographic pattern of glacial retreat is detectable in both climatic and plant‐biomass models of bird diversity.     

A mitogenic view on the evolutionary history of the Holarctic freshwater gadoid, burbot (Lota lota)

Climatic oscillations during the Pleistocene epoch had a dramatic impact on the distribution of biota in the northern hemisphere. In order to trace glacial refugia and postglacial colonization routes on a global scale, we studied mitochondrial DNA sequence variation in a freshwater fish (burbot, Lota lota; Teleostei, Gadidae) with a circumpolar distribution. The subdivision of burbot in the subspecies Lota lota lota (Eurasia and Alaska) and Lota lota maculosa (North America, south of the Great Slave Lake) was reflected in two distinct mitochondrial lineages (average genetic distance is 2.08%). The lota form was characterized by 30 closely related haplotypes and a large part of its range (from Central Europe to Beringia) was characterized by two widespread ancestral haplotypes, implying that transcontinental exchange/migration was possible for cold-adapted freshwater taxa in recent evolutionary time. However, the derived mitochondrial variants observed in peripheral populations point to a recent separation from the core group and postglacial recolonization from distinct refugia. Beringia served as refuge from where L. l. lota dispersed southward into North America after the last glacial maximum. Genetic variation in the maculosa form consisted of three mitochondrial clades, which were linked to at least three southern refugia in North America. Two mitochondrial clades east of the Continental Divide (Mississippian and Missourian clades) had a distinct geographical distribution in the southern refuge zones but intergraded in the previously glaciated area. The third clade (Pacific) was exclusively found west of the Continental Divide.     

Late-glacial and Holocene palaeovegetation zonal reconstruction for central and north-central North America

Aim The purpose of this study is to develop palaeovegetation zonation models for central and north‐central North America, based on late‐Quaternary and Holocene pollen stratigraphic data (n = 246 sites). A secondary purpose was to evaluate an hypothesis ( Strong & Hills, 2003 ) to explain the disjunct distribution of species in western Alberta. Location Hudson Bay‐Lake Michigan to the Rocky Mountains region, north of 36° N to the Arctic Ocean (c. 70° N). Methods Pollen profiles spanning 40 years of palaeoecological research in North America were extracted from published and unpublished archival sources. Individual profiles were subdivided into 1000‐year increments based on the assumption of a constant sedimentation rate between stratigraphic dates (e.g. surface sediments, radiocarbon ¹⁴C dates, tephra layers). The pollen composition among profiles was standardized to 54 commonly recognized taxa, with percentage composition within each stratigraphic sample prorated to 100% prior to analysis. Near‐surface sediments from these profiles were included as analogues of modern vegetation. Cluster analysis was used as a guide to the classification of 2356 temporal stratigraphic samples, which resulted in the recognition of 16 pollen groups. These groups were summarized in terms of their pollen composition, mapped, and used in combination with terrain information and an ecological knowledge of the study area to construct six physiognomically‐based palaeovegetation zonation models at 2000‐year intervals from 14,000 to 4000 yr bp (radiocarbon years before present). Results The 14,000 yr bp model placed Boreal and Cordilleran Forests proximal to the southern glacial front, whereas Arctic tundra dominated the Yukon Territory–Alaska ice‐free zone. Pollen and macrofossil evidence suggests that this Boreal Forest zone contained a mixture of coniferous and deciduous tree species. Grassland was postulated immediately south of the forest zone, with its northern extreme near 49° N latitude in the Alberta–Montana border area. Separation of the Laurentide and Cordilleran glacial fronts about 12,000 yr bp initiated the northward advance of Boreal Forests into western Canada. By the end of the Hypsithermal at about 6000 yr bp , Boreal Forests occurred near the Arctic Ocean, and Grassland and Aspen Parkland zones may have extended to 54° N and 59° N latitude in Alberta, respectively. Between 6000 and 4000 yr bp , a 5° and 1° latitudinal southward shift of the northern Boreal Forest and Grassland/Aspen Parkland boundaries occurred, respectively, near their contemporary positions with corresponding expansions of the Subarctic and Arctic zones. Modern Canadian Cordilleran Forests along the eastern slopes of the Rocky Mountains were interpreted as originating from the north‐central Montana–south‐western Alberta area. Jack pine (Pinus banksiana Lamb.), a common Boreal Forest species, appears to have entered central Canada via the north side of Lake Superior after 11,000 yr bp . Main conclusions Modern vegetation in central Canada evolved from biomes located in the northern USA during the late‐Quaternary. The Boreal Forest biome contained the same arboreal taxa as the modern vegetation, except it lacked jack pine. The proposed regional palaeovegetation models support the hypothesis of Strong & Hills (2003) , but new independent palaeoecological data will be needed for a proper evaluation.     

No‐analog climates and shifting realized niches during the late quaternary: implications for 21st‐century predictions by species distribution models

Empirically derived species distributions models (SDMs) are increasingly relied upon to forecast species vulnerabilities to future climate change. However, many of the assumptions of SDMs may be violated when they are used to project species distributions across significant climate change events. In particular, SDM's in theory assume stable fundamental niches, but in practice, they assume stable realized niches. The assumption of a fixed realized niche relative to climate variables remains unlikely for various reasons, particularly if novel future climates open up currently unavailable portions of species’ fundamental niches. To demonstrate this effect, we compare the climate distributions for fossil‐pollen data from 21 to 15 ka bp (relying on paleoclimate simulations) when communities and climates with no modern analog were common across North America to observed modern pollen assemblages. We test how well SDMs are able to project 20th century pollen‐based taxon distributions with models calibrated using data from 21 to 15 ka. We find that taxa which were abundant in areas with no‐analog late glacial climates, such as Fraxinus, Ostrya/Carpinus and Ulmus, substantially shifted their realized niches from the late glacial period to present. SDMs for these taxa had low predictive accuracy when projected to modern climates despite demonstrating high predictive accuracy for late glacial pollen distributions. For other taxa, e.g. Quercus, Picea, Pinus strobus, had relatively stable realized niches and models for these taxa tended to have higher predictive accuracy when projected to present. Our findings reinforce the point that a realized niche at any one time often represents only a subset of the climate conditions in which a taxon can persist. Projections from SDMs into future climate conditions that are based solely on contemporary realized distributions are potentially misleading for assessing the vulnerability of species to future climate change.     

Scattered late‐glacial and early Holocene tree populations as dispersal nuclei for forest development in north‐eastern European Russia

Aim Concepts about patterns and rates of post‐glacial tree population migration are changing as a result of the increasing amount of palaeobotanical information being provided by macroscopic plant remains. Here we combine macrofossil, pollen and stomata records from five sites in north‐eastern European Russia and summarize the results for the late‐glacial–early Holocene transition. The late‐glacial–early Holocene transition encompasses the first indications of trees (tree‐type Betula, Picea abies, Abies sibirica and Larix sibirica) and subsequent forest development. Considerable time‐lags between the first macrobotanical and/or stomata finds of spruce (Picea abies) and the establishment of a closed forest are reconsidered. Location Pechora basin, north‐eastern European Russia. Methods We used plant macrofossil, stomata, pollen and radiocarbon analyses to reconstruct late‐glacial and early Holocene tree establishment and forest development. The data were derived from lake sediment and peat archives. Results Palaeobotanical data reveal an early Holocene presence (11,500–10,000 cal. yr bp ) of arboreal taxa at all five sites. One site presently located in the northernmost taiga zone, shows the presence of spruce and reproducing tree birch during the late‐glacial. Given the current view of post‐glacial population dynamics and migration rates, it seems likely that the source area of these early tree populations in north‐eastern European Russia was not located in southern Europe but that these populations had local origins. Results thus support the emerging view that the first post‐glacial population expansions in non‐glaciated regions at high latitudes do not reflect migration from the south but were a result of an increase in the size and density of small persisting outlying tree populations. Main conclusions Results suggest that the area east of the margin of the Scandinavian ice sheet to the Ural Mountains had isolated patches of trees during the late‐glacial and early Holocene and that these small populations acted as initial nuclei for population expansion and forest development in the early Holocene.     

Assembly of Alaska‐Yukon boreal steppe communities: Testing biogeographic hypotheses via modern ecological distributions

Beringia (eastern Asia, Alaska, northwest Canada) has been a land‐bridge dispersal route between Asia and North America intermittently since the Mesozoic Era. The Quaternary, the most recent period of exchange, is characterized by large, geologically rapid climate fluctuations and sea‐level changes that alternately expose and inundate the land‐bridge region. Insights into how Quaternary land‐bridge geography has controlled species exchange and assembly of the North American flora comes from focusing on a restricted community with narrow ecological tolerances: species that are today restricted to isolated steppe habitats (dry grasslands) in the Subarctic. We evaluated (i) potential controls over current spatial distributions of steppe plants and their pollinators in Alaska and Yukon and (ii) their ecological distributions in relation to potential biogeographic histories. Taxa present in North America that are disjunct from Asia tended to have larger altitudinal ranges (tolerating colder temperatures) than taxa disjunct from farther south in North America, which were largely restricted to the warmest, lowest‐elevation sites. Ecological findings support the following biogeographic scenarios. Migration from Asia via the land‐bridge occurred during Quaternary glacial periods when conditions were colder and drier than today. While a corridor for migration of cold‐tolerant species of cold steppe and tundra, the land bridge acted as a filter that excluded warmth‐demanding species. Migration from North America occurred under warm, dry interglacial conditions; thermophilous North American disjuncts taking this route may have long histories in Beringia, or they may have migrated recently during the relatively warm and dry early Holocene, when forest cover was incomplete.     

Ice‐age persistence and genetic isolation of the disjunct distribution of larch in Alaska

Larix laricina (eastern larch, tamarack) is a transcontinental North American conifer with a prominent disjunction in the Yukon isolating the Alaskan distribution from the rest of its range. We investigate whether in situ persistence during the last glacial maximum (LGM) or long‐distance postglacial migration from south of the ice sheets resulted in the modern‐day Alaskan distribution. We analyzed variation in three chloroplast DNA regions of 840 trees from a total of 69 populations (24 new sampling sites situated on both sides of the Yukon range disjunction pooled with 45 populations from a published source) and conducted ensemble species distribution modeling (SDM) throughout Canada and United States to hindcast the potential range of L. laricina during the LGM. We uncovered the genetic signature of a long‐term isolation of larch populations in Alaska, identifying three endemic chlorotypes and low levels of genetic diversity. Range‐wide analysis across North America revealed the presence of a distinct Alaskan lineage. Postglacial gene flow across the Yukon divide was unidirectional, from Alaska toward previously glaciated Canadian regions, and with no evidence of immigration into Alaska. Hindcast SDM indicates one of the broadest areas of past climate suitability for L. laricina existed in central Alaska, suggesting possible in situ persistence of larch in Alaska during the LGM. Our results provide the first unambiguous evidence for the long‐term isolation of L. laricina in Alaska that extends beyond the last glacial period and into the present interglacial period. The lack of gene flow into Alaska along with the overall probability of larch occurrence in Alaska being currently lower than during the LGM suggests that modern‐day Alaskan larch populations are isolated climate relicts of broader glacial distributions, and so are particularly vulnerable to current warming trends.     

Effects of Holocene climate change on the historical demography of migrating sharp-shinned hawks (Accipiter striatus velox) in North America

DNA sequences of the mitochondrial control region were analysed from 298 individual sharp-shinned hawks (Accipiter striatus velox) sampled at 12 different migration study sites across North America. The control region proved to be an appropriate genetic marker for identification of continental-scale population genetic structure and for determining the historical demography of population units. These data suggest that sharp-shinned hawks sampled at migration sites in North America are divided into distinct eastern and western groups. The eastern group appears to have recently expanded in response to the retreat of glacial ice at the end of the last glacial maximum. The western group appears to have been strongly effected by the Holocene Hypsithermal dry period, with molecular evidence indicating the most recent expansion following this mid-Holocene climatic event 7000-5000 years before present.     

The climate,the fuel and the land use: Long‐term regional variability of biomass burning in boreal forests

The influence of different drivers on changes in North American and European boreal forests biomass burning (BB) during the Holocene was investigated based on the following hypotheses: land use was important only in the southernmost regions, while elsewhere climate was the main driver modulated by changes in fuel type. BB was reconstructed by means of 88 sedimentary charcoal records divided into six different site clusters. A statistical approach was used to explore the relative contribution of (a) pollen‐based mean July/summer temperature and mean annual precipitation reconstructions, (b) an independent model‐based scenario of past land use (LU), and (c) pollen‐based reconstructions of plant functional types (PFTs) on BB. Our hypotheses were tested with: (a) a west‐east northern boreal sector with changing climatic conditions and a homogeneous vegetation, and (b) a north‐south European boreal sector characterized by gradual variation in both climate and vegetation composition. The processes driving BB in boreal forests varied from one region to another during the Holocene. However, general trends in boreal biomass burning were primarily controlled by changes in climate (mean annual precipitation in Alaska, northern Quebec, and northern Fennoscandia, and mean July/summer temperature in central Canada and central Fennoscandia) and, secondarily, by fuel composition (BB positively correlated with the presence of boreal needleleaf evergreen trees in Alaska and in central and southern Fennoscandia). Land use played only a marginal role. A modification towards less flammable tree species (by promoting deciduous stands over fire‐prone conifers) could contribute to reduce circumboreal wildfire risk in future warmer periods.     

Phylogeographic assessment of mtDNA paraphyly and the evolution of unisexuality in Calligrapha (Coleoptera: Chrysomelidae)

The leaf beetle genus Calligrapha is one of the few examples of animals with several obligate unisexual, female‐only species. Previous work showed that each one arose independently from interspecific hybridization events involving different species. However, all of them clustered in a single mtDNA clade together with some individuals of the parental bisexual species, which appeared as deeply polyphyletic in the mtDNA genealogy of the genus. The dating of these splits using a molecular clock placed them in the Quaternary and it was hypothesized that climatic change during this period may have favored range expansions and secondary contacts required for hybridization. In this work, we test this hypothesis and the origins of unisexuality in Calligrapha examining the diversity of mitochondrial (cox1) and nuclear (wingless, Wg) genes and the Bayesian continuous mtDNA phylogeography of a sample of more than 500 specimens of two bisexual species of Calligrapha at a continental scale and two unisexual species derived from them. Besides a major topological difference, whereby each bisexual species is monophyletic for Wg but paraphyletic for cox1, both gene datasets are consistent with a minimum of seven evolutionary lineages, coherent with geography and consistent with an ordered expansion to occupy their current ranges. The results also imply their survival in well‐established glacial refuges during the Last Glacial Maximum (LGM). Thus, for bisexual C. multipunctata there are two main, southern and northern lineages. The southern lineage expanded its range in two evolutionary branches, to the Rocky Mountains and to the northern Mississippi and Ohio River basins, respectively. The northern lineage has one branch in the Upper Mississippi and one that expanded west to the Pacific Northwest and east to the northeastern North Atlantic, finding refuge in both areas. These major lineages are parapatric in the Northern Great Plains, an area consistent with them having found refuge in the so‐called Driftless region during the LGM. For bisexual C. philadelphica, one northern lineage expanded west from the northern Appalachians and one east and southwest along the axis of the Appalachians, and the timing of events is consistent with their persistence in glacial refugia at both sides of the main Great Lakes lobe of the Laurentide Ice Sheet. There is evidence that the northeastern North Atlantic lineages of both species hybridized at the edge of their ranges after the LGM. The additional, divergent mtDNA lineage of these species shows evidence of range expansions of two lineages, one for each species, in an area south of the Laurentide Ice Sheet, and giving origin to the unisexual species by way of hybridization with other species in the Alleghanian region after the LGM. Interestingly, the individuals of supposedly bisexual species in this clade are all females. This suggests that unisexuality actually predates the origin of unisexual taxa in this system and that some bisexual species in Calligrapha may be species complexes instead, with cryptic species differing in their reproductive mode.     

Biomes of western North America at 18,000, 6000 and 0 14C yr bp reconstructed from pollen and packrat midden data

A new compilation of pollen and packrat midden data from western North America provides a refined reconstruction of the composition and distribution of biomes in western North America for today and for 6000 and 18,000 radiocarbon years before present (¹⁴C yr bp ). Modern biomes in western North America are adequately portrayed by pollen assemblages from lakes and bogs. Forest biomes in western North America share many taxa in their pollen spectra and it can be difficult to discriminate among these biomes. Plant macrofossils from packrat middens provide reliable identification of modern biomes from arid and semiarid regions, and this may also be true in similar environments in other parts of the world. However, a weighting factor for trees and shrubs must be used to reliably reconstruct modern biomes from plant macrofossils. A new biome, open conifer woodland, which includes eurythermic conifers and steppe plants, was defined to categorize much of the current and past vegetation of the semiarid interior of western North America. At 6000 ¹⁴C yr bp , the forest biomes of the coastal Pacific North‐west and the desert biomes of the South‐west were in near‐modern positions. Biomes in the interior Pacific North‐west differed from those of today in that taiga prevailed in modern cool/cold mixed forests. Steppe was present in areas occupied today by open conifer woodland in the northern Great Basin, while in the central and southern Rocky Mountains forests grew where steppe grows today. During the mid‐Holocene, cool conifer forests were expanded in the Rocky Mountains (relative to today) but contracted in the Sierra Nevada. These differences from the forests of today imply different climatic histories in these two regions between 6000 ¹⁴C yr bp and today. At 18,000 ¹⁴C yr bp , deserts were absent from the South‐west and the coverage of open conifer woodland was greatly expanded relative to today. Steppe and tundra were present in much of the region now covered by forests in the Pacific North‐west.     

Climate change and evolution of the New World pitviper genus Agkistrodon (Viperidae)

Aim We derived phylogenies, phylogeographies, and population demographies for two North American pitvipers, Agkistrodon contortrix (Linnaeus, 1766) and A. piscivorus (Lacépède, 1789) (Viperidae: Crotalinae), as a mechanism to evaluate the impact of rapid climatic change on these taxa. Location Midwestern and eastern North America. Methods We reconstructed maximum parsimony (MP) and maximum likelihood (ML) relationships based on 846 base pairs of mitochondrial DNA (mtDNA) ATPase 8 and ATPase 6 genes sequenced over 178 individuals. We quantified range expansions, demographic histories, divergence dates and potential size differences among clades since their last period of rapid expansion. We used the Shimodaira–Hasegawa (SH) test to compare our ML tree against three biogeographical hypotheses. Results A significant SH test supported diversification of A. contortrix from northeastern Mexico into midwestern–eastern North America, where its trajectory was sundered by two vicariant events. The first (c. 5.1 Ma) segregated clades at 3.1% sequence divergence (SD) along a continental east–west moisture gradient. The second (c. 1.4 Ma) segregated clades at 2.4% SD along the Mississippi River, coincident with the formation of the modern Ohio River as a major meltwater tributary. A single glacial refugium was detected within the Apalachicola region of southeastern North America. Significant support was also found for a hypothesis of trans‐Gulf rafting by the common ancestor of A. piscivorus from eastern Mexico (possibly the Yucatan Peninsula) to northern Florida. There, a Mid–Late Pliocene marine transgression separated it at 4.8% SD from mainland North America. Significant range expansions followed compressive glacial effects in three (of four) A. contortrix clades and in two (of three) A. piscivorus clades, with the Florida A. piscivorus clade exhibiting significant distributional stasis. Main conclusions Pliocene glaciations, rapidly developing western aridity, and Pleistocene glacial meltwaters seemingly led to the diversification of A. contortrix and A. piscivorus in North America. Both species were pushed southwards by Pleistocene climate change, with subsequent northward expansions uninhibited topographically. The subspecific taxonomy used for A. contortrix and A. piscivorus today, however, appear non‐representative. The monophyletic Florida subspecies of A. piscivorus may be a distinct species (at 4.8% SD), whereas two western subspecies of A. contortrix also appear to constitute a single distinct species, pending additional analyses. We conclude that both species of Agkistrodon can be used as suitable ectothermic models to gauge impacts of future climate change.     

Population genomic evidence for plant glacial survival in Scandinavia

Quaternary glaciations have played a major role in shaping the genetic diversity and distribution of plant species. Strong palaeoecological and genetic evidence supports a postglacial recolonization of most plant species to northern Europe from southern, eastern and even western glacial refugia. Although highly controversial, the existence of small in situ glacial refugia in northern Europe has recently gained molecular support. We used genomic analyses to examine the phylogeography of a species that is critical in this debate. Carex scirpoidea Michx subsp. scirpoidea is a dioecious, amphi‐Atlantic arctic–alpine sedge that is widely distributed in North America, but absent from most of Eurasia, apart from three extremely disjunct populations in Norway, all well within the limits of the Weichselian ice sheet. Range‐wide population sampling and variation at 5,307 single nucleotide polymorphisms show that the three Norwegian populations comprise unique evolutionary lineages divergent from Greenland with high between‐population divergence. The Norwegian populations have low within‐population genetic diversity consistent with having experienced genetic bottlenecks in glacial refugia, and host private alleles that probably accumulated in long‐term isolated populations. Demographic analyses support a single, pre‐Weichselian colonization into Norway from East Greenland, and subsequent divergence of the three populations in separate refugia. Other refugial areas are identified in North‐east Greenland, Minnesota/Michigan, Colorado and Alaska. Admixed populations in British Columbia and West Greenland indicate postglacial contact. Taken together, evidence from this study strongly indicates in situ glacial survival in Scandinavia.