Miocene snakes from northeastern Kazakhstan: new data on the evolution of snake assemblages in Siberia

Abstract

The Neogene snake fauna from the central and eastern regions of Eurasia is still largely unknown. This paper reports on a unique snake fauna from the late middle Miocene of the Baikadam and Malyi Kalkaman 1 and 2 localities, northeastern Kazakhstan, which represents the best-documented Miocene snake assemblage in Central Asia. Previous studies admitted that snake fauna could be homogeneous over a large part of Eurasia during the Miocene, with the late middle to early late Miocene assemblages similar to snake assemblages that inhabited Europe in the late early and early middle Miocene. This assumption is partially supported by the presence of Texasophis bohemiacus and Coluber cf. hungaricus, as well as vipers of the ‘V. aspis’ complex. However, the presence of taxa which are (1) probably not related to European representatives (‘Colubrinae’ A and B), (2) probably never occurred in Central and Western Europe and (3) are closely related to species recently inhabiting southern Siberia (Elaphe aff. dione, Gloydius sp.) indicates that faunal dissimilarity was relatively high within Eurasia during the late middle Miocene. This assumption is in accordance with studies of small mammal assemblages which show a decreasing homogenity in the Eurasia in the course of the middle Miocene.     

MAMMALIAN EXTINCTIONS IN THE LATE PLEISTOCENE OF NORTHERN EURASIA AND NORTH AMERICA

The 'mass extinctions' at the end of the Pleistocene were unique, both in the Pleistocene and earlier in the geological record, in that the species lost were nearly all large terrestrial mammals. Although a global phenomenon, late Pleistocene extinctions were most severe in North America, South America and Australia, and moderate in northern Eurasia (Europe plus Soviet Asia). In Africa, where nearly all of the late Pleistocene 'megafauna' survives to the present day, losses were slight. Ruling out epidemic disease or cosmic catastrophe, the contending hypotheses to explain late Pleistocene extinctions are: (a) failure to adapt to climatic/environmental change; and (b) extermination by human hunters ('prehistoric overkill'). This review focuses on extinctions in northern Eurasia (mainly Europe) in comparison with North America. In addition to reviewing the faunal evidence, the highly relevant environmental and archaeological backgrounds are summarized. The latest survival dates of extinct species are estimated from stratigraphic occurrences of fossil remains, radiocarbon dates, or association with archaeological industries. The Middle and Upper Pleistocene (ca. 700,000-10,000 BP) in northern Eurasia and North America was a time of constantly changing climate, ranging from phases of extensive glaciation in cold stages, to temperate periods (interglacials). In the Lateglacial (ca. 15,000-10,000 BP), during which most extinctions occurred, there was a major reorganization of vegetation, mainly involving the replacement of open vegetation by forests. These changes were more profound than earlier in the Last Cold Stage, but similar in nature to vegetational changes that took place at previous cold stage/interglacial transitions. The archaeological record shows that humans have been present in Europe since the early Middle Pleistocene. The arrival in Europe ca. 35,000 BP of 'anatomically modern humans', with their technologically more advanced upper palaeolithic industries, was a 'quantum leap' in human history. Extinctions occurred throughout the European Pleistocene, but until the late Pleistocene most losses were replaced by the evolution or immigration of new species, and most of those lost without replacement were small mammals. In marked contrast, extinctions without replacement in the late Pleistocene were almost entirely confined to the largest mammals (greater than 1000 kg) and some medium-large species (100-1000 kg).(ABSTRACT TRUNCATED AT 400 WORDS)     

East Asian allopatry and north Eurasian sympatry in Long‐tailed Tit lineages despite similar population dynamics during the late Pleistocene

Eurasia is a large continent characterized by heterogeneous environments. Glacial cycles during the late Pleistocene have had variable impacts on the avifauna across Eurasia. Bird populations from South‐East Asia show stability through the Last Glacial Maximum (LGM), while populations from Europe exhibit evidence of post‐LGM expansion. We investigated the phylogeography of the Long‐tailed Tit (Aegithalos caudatus), which spans the longitudinal breadth of Eurasia to test how climatic history and regional topographical complexity affected populations and diversification within the species complex. Our results show that two lineages from central and southern China (lineages C and D) segregate geographically, while lineages across northern Eurasia (lineage A and B) show substantial sympatry. Bayesian estimates for the timing of diversification suggest that the four lineages diverged during the middle Pleistocene, splitting in parallel and undergoing concurrent demographic histories since divergence. A. caudatus lineages experienced similar and synchronous population size dynamics during glacial cycles before the LGM. We conclude that the difference in geo‐topologic complexity may be an important factor that led to the variation in secondary admixture between northern Eurasian and eastern Asian lineages.     

A multivariate approach to the determination of faunal structures among European butterfly species (Lepidoptera: Rhopalocera)

Multivariate analyses of 393 butterfly species over 85 geographical areas (R- and Q-data matrices) in Europe and North Africa have produced a consistent pattern of faunal structures (units and regions). Prominent features to emerge are the latitudinal zonation of geographical units and the division of the Mediterranean into western and eastern components; southwards in Europe, endemicity increases whereas faunal structures decrease in spatial dimensions. Central Europe–from the Urals to the British Isles–forms a single large faunal structure (extent unit and region). A model has been constructed to account for Pleistocene evolutionary changes and endemism in European butterflies and for the east-west taxonomic divisions in the extent faunal structure which dominates central Europe. Periodic Pleistocene climatic changes have resulted in cycles of population extinction, isolation, evolution and migration, but the nature and timing of events has depended on the environmental tolerances of species belonging to different faunal units. During Pleistocene glaciations, southern species have been relatively static and more isolated and have evolved independently. By comparison, northern species have been more mobile and have migrated over large distances. Contact and hybrid zones among cosmopolitan species in northern Europe are probably of some antiquity. They result from persistent survival and isolation of refuge populations in the west and east Mediterranean during glacial phases; dispersal from these refuges leads to their regeneration during each interglacial.     

The integration of multiple independent data reveals an unusual response to Pleistocene climatic changes in the hard tick Ixodes ricinus

In the last few years, improved analytical tools and the integration of genetic data with multiple sources of information have shown that temperate species exhibited more complex responses to ice ages than previously thought. In this study, we investigated how Pleistocene climatic changes affected the current distribution and genetic diversity of European populations of the tick Ixodes ricinus, an ectoparasite with high ecological plasticity. We first used mitochondrial and nuclear genetic markers to investigate the phylogeographic structure of the species and its Pleistocene history using coalescent‐based methods; then we used species distribution modelling to infer the climatic niche of the species at last glacial maximum; finally, we reviewed the literature on the I. ricinus hosts to identify the locations of their glacial refugia. Our results support the scenario that during the last glacial phase, I. ricinus never experienced a prolonged allopatric divergence in separate glacial refugia, but persisted with interconnected populations across Southern and Central Europe. The generalist behaviour in host choice of I. ricinus would have played a major role in maintaining connections between its populations. Although most of the hosts persisted in separate refugia, from the point of view of I. ricinus, they represented a continuity of ‘bridges’ among populations. Our study highlights the importance of species‐specific ecology in affecting responses to Pleistocene glacial–interglacial cycles. Together with other cases in Europe and elsewhere, it contributes to setting new hypotheses on how species with wide ecological plasticity coped with Pleistocene climatic changes.     

Effects of late quaternary climate change on Palearctic shrews

The Late Quaternary was a time of rapid climatic oscillations and drastic environmental changes. In general, species can respond to such changes by behavioral accommodation, distributional shifts, ecophenotypic modifications (nongenetic), evolution (genetic) or ultimately face local extinction. How those responses manifested in the past is essential for properly predicting future ones especially as the current warm phase is further intensified by rising levels of atmospheric carbon dioxide. Here, we use ancient DNA (aDNA) and morphological features in combination with ecological niche modeling (ENM) to investigate genetic and nongenetic responses of Central European Palearctic shrews to past climatic change. We show that a giant form of shrew, previously described as an extinct Pleistocene Sorex species, represents a large ecomorph of the common shrew (Sorex araneus), which was replaced by populations from a different gene‐pool and with different morphology after the Pleistocene Holocene transition. We also report the presence of the cold‐adapted tundra shrew (S. tundrensis) in Central Europe. This species is currently restricted to Siberia and was hitherto unknown as an element of the Pleistocene fauna of Europe. Finally, we show that there is no clear correlation between climatic oscillations within the last 50 000 years and body size in shrews and conclude that a special nonanalogous situation with regard to biodiversity and food supply in the Late Glacial may have caused the observed large body size.     

The first human dispersal to Europe: Remarks on the archaeological and palaeoanthropological record from Orce (Guadix-Baza basin, southeastern Spain)

The chronology of the first arrival ofHomo in Europe is a rather controversial issue, with most scholars claiming until very recently that there were no permanent human settlements before the middle Pleistocene. However, new findings at Atapuerca, Dmanisi and Orce, as well as the re-evaluation of the evidence from Java, Israel and China indicate a protracted chronology for the arrival of hominids in Eurasia, during late Pliocene/lower Pleistocene times. The systematic study of the macrovertebrate assemblages from circummediterranean sites such as Orce and Dmanisi has shown a faunal replacement at the Plio-Pleistocene boundary, marked by the arrival in Europe of African immigrants such as the hippopotamus, an equid similar to modern grevy’s zebra, a large cercopithecoid and several carnivores, including a giant hyaena, a sabre-tooth and a wild dog. An analysis of the relative frequency of bones and teeth for those species of large mammals preserved in Venta Micena indicates the improbability thatHomo sp. is represented in the fossil assemblage by several bones and no tooth remain. Finally, new data on the stratigraphy of Barranco León are offered, with remarks on the discovery in this locality of lithic artefacts and molar tooth fragment BL5-0.     

Late‐glacial recolonization and phylogeography of European red deer (Cervus elaphus L.)

The Pleistocene was an epoch of extreme climatic and environmental changes. How individual species responded to the repeated cycles of warm and cold stages is a major topic of debate. For the European fauna and flora, an expansion–contraction model has been suggested, whereby temperate species were restricted to southern refugia during glacial times and expanded northwards during interglacials, including the present interglacial (Holocene). Here, we test this model on the red deer (Cervus elaphus) a large and highly mobile herbivore, using both modern and ancient mitochondrial DNA from the entire European range of the species over the last c. 40 000 years. Our results indicate that this species was sensitive to the effects of climate change. Prior to the Last Glacial Maximum (LGM) haplogroups restricted today to South‐East Europe and Western Asia reached as far west as the UK. During the LGM, red deer was mainly restricted to southern refugia, in Iberia, the Balkans and possibly in Italy and South‐Western Asia. At the end of the LGM, red deer expanded from the Iberian refugium, to Central and Northern Europe, including the UK, Belgium, Scandinavia, Germany, Poland and Belarus. Ancient DNA data cannot rule out refugial survival of red deer in North‐West Europe through the LGM. Had such deer survived, though, they were replaced by deer migrating from Iberia at the end of the glacial. The Balkans served as a separate LGM refugium and were probably connected to Western Asia with genetic exchange between the two areas.     

Occurrence of Sapindopsis (Platanaceae) in the Cretaceous of Eurasia

Three new species of the genus Sapindopsis Fontaine from Central Asia and western Siberia are described: S. neuburgae (Vachr.) Golovn., comb. nov., S. janschinii (Vachr.) Golovn., comb. nov., and S. kryshtofovichii (I. Lebed.) Golovn., comb. nov. Geographical and stratigraphic analyses have shown that the genus evolved in Eurasia from the Middle Albian to the Cenomanian, and was mostly restricted to the subtropics of the Euro-Sinian phytogeographic area from the Middle East to Russian Primorye and northern China. The are no reliable records of Sapindopsis from Europe. The migration of Sapindopsis from North America to Eurasia through the Bering Land Bridge was most probably related to the Early-Middle Albian climatic optimum.     

Phylogeography and demographic history of Shaw's Jird (Meriones shawii complex) in North Africa

Palaeoenvironmental data and climatic reconstructions show that the Mediterranean ecoregion of North Africa underwent drastic ecological changes during the Pleistocene. Given its rich palaeontological record, North Africa is a pertinent region for documenting the role of climate change and human mediated‐habitat changes on the demography and genetic structure of faunal species. In the present study, we collected data from this species in Morocco, Algeria, and Tunisia, and we combined molecular (mitochondrial and nuclear DNA sequences, microsatellites), fossil, palaeoenvironmental, and human context data to propose an explanation for the fluctuations of populations belonging to the Meriones shawii complex in the past. Genetic and fossil data both indicate a strong bottleneck in Moroccan populations at the Middle Holocene (last interglacial optimum) compared to the Late Pleistocene. Our mitochondrial DNA data suggest a diversification event within Morocco corresponding to the 130–125 kya interglacial optimum. Given that (1) major demographic changes in the M. shawii complex coincide with the interglacial optimums, and (2) the impact of human activities on the landscape and faunal communities was moderate during the Middle Holocene (beginnings of the Neolithic culture), our results demonstrate that climate, rather than anthropogenic influences, likely explains the M. shawii complex population decline in the Holocene.     

Niche shifting in response to warming climate after the last glacial maximum: inference from genetic data and niche assessments in the chisel‐toothed kangaroo rat (Dipodomys microps)

During Pleistocene glacial‐interglacial cycles, the geographic range is often assumed to have shifted as a species tracks its climatic niche. Alternatively, the geographic range would not necessarily shift if a species can adapt in situ to a changing environment. The potential for a species to persist in place might increase with the diversity of habitat types that a species exploits. We evaluate evidence for either range shift or range stability between the last glacial maximum (LGM) and present time in the chisel‐toothed kangaroo rat (Dipodomys microps), an endemic of the Great Basin and Mojave deserts. We modeled how the species’ range would have changed if the climatic niche of the species remained conserved between the LGM and present time. The climatic models imply that if D. microps inhabited the same climatic niche during the LGM as it does today, the species would have persisted primarily within the warm Mojave Desert and expanded northwards into the cold Great Basin only after the LGM. Contrary to the climatic models, the mitochondrial DNA assessment revealed signals of population persistence within the current distribution of the species throughout at least the latest glacial‐interglacial cycle. We concluded that D. microps did not track its climatic niche during late Pleistocene oscillations, but rather met the challenge of a changing environment by shifting its niche and retaining large portions of its distribution. We speculate that this kind of response to fluctuating climate was possible because of ‘niche drifting’, an alteration of the species’ realized niche due to plasticity in various biological characters. Our study provides an example of an approach to reconstruct species’ responses to past climatic changes that can be used to evaluate whether and to what extent taxa have capacity to shift their niches in response to the changing environment – information becoming increasingly important to predicting biotic responses to future environmental changes.     

Glacial refugia of mammals in Europe: evidence from fossil records

• 1 Glacial refugia were core areas for the survival of temperate species during unfavourable environmental conditions and were the sources of postglacial recolonizations. Unfortunately, the locations of glacial refugia of animals and plants are usually described by models, without reference to facts about real geographical ranges at that time.
• 2 Careful consideration of the faunal assemblages of archaeological sites from the Younger Palaeolithic, which are precisely dated to the Last Glacial Maximum (LGM), gives indications about the distribution of species during the LGM (23 000–16 000 bp ) and provides evidence for the locations of glacial refugia for mammalian species in Europe.
• 3 In Europe, 47 LGM sites, dating from 23 000 to 16 000 bp and containing typical temperate mammal species, have been described. The geographical range of these archaeological sites clearly shows a distribution which differs from the hypothesized traditional refuge areas of the temperate fauna. A considerable number of sites situated in the Dordogne in south‐western France and the Carpathian region contain records of red deer Cervus elaphus, roe deer Capreolus capreolus, wild boar Sus scrofa and red fox Vulpes vulpes.
• 4 The faunal composition of the majority of the evaluated Palaeolithic sites in the southern European peninsulas (with the exception of Greece), as well as France and the Carpathian region, indicates the co‐occurrence of these temperate species with cold‐adapted faunal elements such as mammoth Mammuthus primigenius and/or reindeer Rangifer tarandus.
• 5 The survival of species in Central European refugia would have significant consequences for phylogeography and would be revealed by the dominant distribution of haplotypes, originating from this region. A Carpathian refuge could also be the reason for the very early records of small mammals or mustelids from the Late‐Glacial or Interstadials before the LGM in regions like southern Germany.

     

Quaternary refugia are associated with higher speciation rates in mammalian faunas of the Western Palaearctic

Quaternary climate changes have contributed to shape the biogeographic distribution of extant species. The combination of climatic niche conservatism and glacial‐interglacial cycles forced many species to retract their range limits for surviving under the advance of Pleistocene ice‐sheets. Refugia offered geographical opportunities for species to retreat, persist and, later on, begin recolonization processes under favourable environmental conditions. Here we explore the hypothesis that refugia have been not only crucial for the survival of multiple species but also acted as speciation centres for Western Palaearctic mammals. We define ‘recurrent massive refugia’ as those geographical regions that have historically accumulated the highest levels of co‐occurring species for several Quaternary cycles. Our assemblage‐level analyses identify the existence of refugia within the Iberian Peninsula and the Atlantic French margin that were recurrently selected by most mammals. The topographic heterogeneity, climatic stability and microhabitat availability of these refugial areas may have offered suitable habitat conditions for multiple species during different climatic events over time. Using a Bayesian analysis of macroevolutionary mixtures we detected that the higher level of divergence and accumulative evolutionary changes in mammals of the Western Palaearctic are found in refugia. The continuous retractions and expansions of species’ ranges during the Pleistocene promoted temporal changes in the composition and richness of communities in this biogeographic region. The reorganization of ecological composition driven by cyclical climatic events may have favoured the emergence of biotic interactions and ecological responses conducive to novel selective pressures. Our findings suggest, first, that multiple climatic changes in the form of glacial‐interglacial transitions during the Quaternary have left a detectable imprint on the observed geographical patterns of species richness in mammalian faunas of the Western Palaearctic, and second, highlight the importance of refugia for the preservation of species (‘museums’) and as centers of speciation and endemism (‘cradles’) as well.     

Deep intra‐island divergence of a montane forest endemic: phylogeography of the Puerto Rican frog Eleutherodactylus portoricensis (Anura: Eleutherodactylidae)

Aim Hypotheses proposed for lineage diversification of tropical montane species have rarely been tested within oceanic islands. Our goal was to understand how basin barriers and Pleistocene climatic fluctuations shaped the distribution of diversity in Eleutherodactylus portoricensis (Eleutherodactylidae), a frog endemic to the montane rain forests of Puerto Rico. Location The north‐eastern (Luquillo) and south‐eastern (Cayey) mountains of Puerto Rico. Methods We generated mitochondrial DNA (mtDNA) control region sequences (c. 565 bp) from 144 individuals of E. portoricensis representing 16 localities, and sequenced 646 bp of cytochrome b and 596 bp of nuclear DNA (nDNA) rhodopsin exon and intron 1 from a subset of individuals. We conducted a phylogenetic analysis on the mtDNA sequence data and explored population substructure with maximum parsimony networks, a spatial analysis of molecular variance, and pairwise F_ST analysis. Coalescent simulations were performed to test alternative models of population divergence in response to late Pleistocene interglacial periods. Historical demography was assessed through coalescent analyses and Bayesian skyline plots. Results We found: (1) two highly divergent groups associated with the disjunct Luquillo and Cayey Mountains, respectively; (2) a shallow mtDNA genetic discontinuity across the La Plata Basin within the Cayey Mountains; (3) phylogeographic congruence between nDNA and mtDNA markers; (4) divergence dates for both mtDNA and nDNA pre‐dating the Holocene interglacial (c. 10 ka), and nDNA suggesting divergence in the penultimate interglacial (c. 245 ka); and (5) historical demographic stability in both lineages. Main conclusions The low‐elevation Caguas Basin is a long‐term barrier to gene flow between the two montane frog populations. Measures of genetic diversity for mtDNA were similar in both lineages, but lower nDNA diversity in the Luquillo Mountains lineage suggests infrequent dispersal between the two mountain ranges and colonization by a low‐diversity founder population. Population divergence began prior to the Holocene interglacial. Stable population sizes over time indicate a lack of demonstrable demographic response to climatic changes during the last glacial period. This study highlights the importance of topographic complexity in promoting within‐island vicariant speciation in the Greater Antilles, and indicates long‐term persistence and lineage diversification despite late Pleistocene climatic oscillations.     

Phylogenomics unravels Quaternary vicariance and allopatric speciation patterns in temperate‐montane plant species: A case study on the Ranunculus auricomus species complex

The time frame and geographical patterns of diversification processes in European temperate‐montane herbs are still not well understood. We used the sexual species of the Ranunculus auricomus complex as a model system to understand how vicariance versus dispersal processes in the context of Pleistocene climatic fluctuations have triggered speciation in temperate‐montane plant species. We used target enrichment sequence data from about 600 nuclear genes and coalescent‐based species tree inference methods to resolve phylogenetic relationships among the sexual taxa of the complex. We estimated absolute divergence times and, using ancestral range reconstruction, we tested if speciation was enhanced by vicariance or by dispersal processes. Phylogenetic relationships among taxa were fully resolved with some incongruence in the position of the tetraploid R. marsicus. Speciation events took place in a very short time at the end of the Mid‐Pleistocene Transition (830–580 thousand years ago [ka]). A second wave of intraspecific geographical differentiation occurred at the end of the Riss glaciation or during the Eemian interglacial between 200 and 100 ka. Ancestral range reconstruction suggests a widespread European ancestor of the R. auricomus complex. Vicariance has triggered allopatric speciation in temperate‐montane plant species during the climatic deterioration that occurred during the last phase of the Mid‐Pleistocene Transition. Vegetation restructuring from forest into tundra could have confined these forest species into isolated glacial macro‐ and microrefugia. During subsequent warming periods, range expansions of these species could have been hampered by apomictic derivatives and by other congeneric competitors in the same habitat.     

Complete taxon sampling of the avian genus Pica (magpies) reveals ancient relictual populations and synchronous Late‐Pleistocene demographic expansion across the Northern Hemisphere

Previous studies have suggested that bird populations in east Asia were less affected by Pleistocene climatic fluctuations than those in Europe and North America. However, this is mainly based on comparisons among species. It would be more relevant to analyse geographical populations of widespread species or species complexes. We analyzed two mitochondrial genes and two nuclear introns for all taxa of Pica to investigate 1) which Earth history factors have shaped the lineage divergence, and 2) whether different geographical populations were differently affected by the Pleistocene climatic changes. Our mitochondrial tree recovered three widespread lineages, 1) in east Asia, 2) across north Eurasia, and 3) in North America, respectively, with three isolated lineages in northwest Africa, Arabia and the Qinghai‐Tibet Plateau, respectively. Divergences among lineages took place 1.4–3.1 million yr ago. The northwest African population was sister to the others, which formed two main clades. In one of these, Arabia was sister to Qinghai‐Tibet, and these formed the sister clade to the east Asia clade. The other main clade comprised the North American and north Eurasian clades. There was no or very slight structure within these six geographical clades, including a lack of differentiation between the two North American species black‐billed magpie P. hudsonia and yellow‐billed magpie P. nutalli. Demographic expansion was recorded in the three most widespread lineages after 0.06 Ma. Asymmetric gene flow was recorded in the north Eurasian clade from southwestern Europe eastward, whereas the east Asian clade was rooted in south central China. Our results indicate that the fragmentation of the six clades of Pica was related to climatic cooling and aridification during periods of the Pliocene–Pleistocene. Populations on both sides of the Eurasian continent were similarly influenced by the Pleistocene climate changes and expanded concomitantly with the expansion of steppes. Based on results we also propose a revised taxonomy recognising seven species of Pica.     

Status of Savi's pipistrelle Hypsugo savii (Chiroptera) and range expansion in Central and south‐eastern Europe: a review

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Interglacial microrefugia and diversification of a cactus species complex: phylogeography and palaeodistributional reconstructions for Pilosocereus aurisetus and allies

The role of Pleistocene climate changes in promoting evolutionary diversification in global biota is well documented, but the great majority of data regarding this subject come from North America and Europe, which were greatly affected by glaciation. The effects of Pleistocene changes on cold‐ and/or dry‐adapted species in tropical areas where glaciers were not present remain sparsely investigated. Many such species are restricted to small areas surrounded by unfavourable habitats, which may represent potential interglacial microrefugia. Here, we analysed the phylogeographic structure and diversification history of seven cactus species in the Pilosocereus aurisetus complex that are restricted to rocky areas with high diversity and endemism within the Neotropical savannas of eastern South America. We combined palaeodistributional estimates with standard phylogeographic approaches based on two chloroplast DNA regions (trnT‐trnL and trnS‐trnG), exon 1 of the nuclear gene PhyC and 10 nuclear microsatellite loci. Our analyses revealed a phylogeographic history marked by multiple levels of distributional fragmentation, isolation leading to allopatric differentiation and secondary contact among divergent lineages within the complex. Diversification and demographic events appear to have been affected by the Quaternary climatic cycles as a result of isolation in multiple patches of xerophytic vegetation. These small patches presently harbouring P. aurisetus populations seem to operate as microrefugia, both at present and during Pleistocene interglacial periods; the role of such microrefugia should be explored and analysed in greater detail.     

Geographic patterns of genetic diversity in Poulsenia armata (Moraceae): implications for the theory of Pleistocene refugia and the importance of riparian forest

Abstract. Tropical forests are species-diverse communities, but we know very little about the geographical distribution of genetic diversity within a species. During the late Pleistocene, lower temperatures and rainfall reduced the distribution of tropical wet forests, and in Central America lowland species may have been limited to riparian habitats. Approximately 12,000 years bp , temperature and rainfall increased in Central America, the distribution of wet forest species expanded, and today the distribution of some species extends into southern Mexico. The distribution of genetic diversity, based on RAPD markers, among ten populations of Poulsenia armata (Miq.) Standl. (Moraceae) suggests that these populations did not originate from a single refugium or region in the late Pleistocene. The Central American populations had high genetic diversity and unique bands giving support to the hypothesis that populations of P. armata occurred in Central American during the late Pleistocene. The majority of genetic diversity was partitioned among populations and there was no geographical relationship among populations, suggesting that these populations were isolated for a long period and there has been little gene flow. Populations of P. armata may have persisted in riparian zones along the Caribbean coast during the late Pleistocene. Given that riparian forest can support high levels of biodiversity in ecological time, and they have played an important role during periods of climate change over geological time, their conservation is of utmost importance particularly with the threat of a rapid shift in climatic patterns.     

Late Pliocene and Quaternary Eurasian locust infestations in the Canary Archipelago

Meco, J., Muhs, D.R., Fontugne, M., Ramos, A.J.G., Lomoschitz, A. & Patterson, D. 2010: Late Pliocene and Quaternary Eurasian locust infestations in the Canary Archipelago. Lethaia, Vol. 44, pp. 440–454. The Canary Archipelago has long been a sensitive location to record climate changes of the past. Interbedded with its basalt lavas are marine deposits from the principal Pleistocene interglacials, as well as aeolian sands with intercalated palaeosols. The palaeosols contain African dust and innumerable relict egg pods of a temperate‐region locust (cf. Dociostaurus maroccanus Thunberg 1815 ). New ecological and stratigraphical information reveals the geological history of locust plagues (or infestations) and their palaeoclimatic significance. Here, we show that the first arrival of the plagues to the Canary Islands from Africa took place near the end of the Pliocene, ca. 3 Ma, and reappeared with immense strength during the middle Late Pleistocene preceding MIS (marine isotope stage) 11 (ca. 420 ka), MIS 5.5 (ca. 125 ka) and probably during other warm interglacials of the late Middle Pleistocene and the Late Pleistocene. During the Early Holocene, locust plagues may have coincided with a brief cool period in the current interglacial. Climatically, locust plagues on the Canaries are a link in the chain of full‐glacial arid–cold climate (calcareous dunes), early interglacial arid–sub‐humid climate (African dust inputs and locust plagues), peak interglacial warm–humid climate (marine deposits with Senegalese fauna), transitional arid–temperate climate (pedogenic calcretes), and again full‐glacial arid–cold climate (calcareous dunes) oscillations. During the principal interglacials of the Pleistocene, the Canary Islands recorded the migrations of warm Senegalese marine faunas to the north, crossing latitudes in the Euro‐African Atlantic. However, this northward marine faunal migration was preceded in the terrestrial realm by interglacial infestations of locusts. □Locust plagues, Canary Islands, Late Pliocene, Pleistocene, Holocene, palaeoclimatology.