Single Refugium Research Articles

Low Within Population Genetic Variation and High Among Population Differentiation inCyrtomium falcatum(L.f.) C. Presl (Dryopteridaceae) in Southern Korea: Inference of Population-Establishment History

Abstract In the Korean Peninsula, the current distribution of the warm-temperate and subtropical vegetation (including many homosporous ferns) is limited to southern coastal areas. Paleoecological data suggest that during the Last Glacial Maximum this vegetation retreated to glacial refugia putatively located in southern Japan and/or southern China, followed by a post-glacial recolonization. Two broad scenarios of post-glacial recolonization could be hypothesized: extant Korean populations are derived from multiple source populations (i.e., from multiple refugia); alternatively, they originate from a single refugium. To test which of these scenarios is more likely, we surveyed patterns of genetic diversity in eight (n = 307) populations of Cyrtomium falcatum from southern Korea. We found extremely low levels of allozyme variation within populations coupled with high among-population differentiation. These data best support the second hypothesis, and indicate that the current genetic diversity may be a c...

Continent-scale genetic structure in a boreal forest migrant, the Blackpoll Warbler (Setophaga striata)

ABSTRACT. We describe the range-wide phylogeography of Blackpoll Warblers (Setophaga striata), a migratory passerine with a broad breeding range in North America's boreal forest that encompasses several possible biogeographic barriers but shows no phenotypic geographic variation. We used mitochondrial control-region sequences from 304 individuals in combination with ecological niche models and coalescent simulations to test alternative historical hypotheses about the number of Pleistocene refugial populations and divergences among modern populations. Population pairwise FSTand spatial analyses of molecular variance suggested significant genetic structure among western, eastern, and Newfoundland populations, but no structure among sky-island populations at the southeastern periphery of the breeding range. Inferred gene flow fits a model of isolation-by-distance. Coalescent simulations rejected all multiple-refugia hypotheses in favor of a single refugium. Paleodistribution models and modern migratory pathw...

Phylogeography of the Korean pine (Pinus koraiensis) in northeast Asia: inferences from organelle gene sequences

Range-wide genetic variation of Korean pine (Pinus koraiensis) was assessed using maternally inherited mtDNA and paternally inherited cpDNA for 16 natural populations throughout northeast Asia in order to study its phylogeographical history during the Quaternary. The cpDNA variation indicated that there was no difference between populations on the Asian continent and those in the Japanese archipelago. In contrast, the mtDNA variation indicated that there was significant difference between the populations from the two regions, with each region having a different lineage. The continental populations exhibited no diversity in the mtDNA examined despite the species' current extensive range and large populations. Conversely, while the Korean pine is rare in Japan, the Japanese populations exhibited greater levels of mtDNA diversity (H (T)=0.502). The higher mtDNA diversity and evidence from numerous Korean pine macrofossil remains dated to the Pleistocene and recovered various sites in Japan suggest that the Japanese archipelago once served as a refugium to a much larger Korean pine population with a more extensive range than is the case today. The presence of the single mtDNA haplotype across the Asian continent suggests that the present widespread populations could have expanded from a single refugium population after the last glacial periods.

Genetic diversity of flat-headed vole (Alticola strelzowi (Kastschenko, 1899)) inferred from cytochrome b variation

Variation of the cytochrome b gene fragment was examined in 27 flat-headed voles Alticola strelzowi from different parts of the species range. A total of 15 haplotypes were described, while the species as a whole was characterized by low levels of genetic differentiation and polymorphism. The haplotypes fell into three haplogroups, one of which corresponded to the subspecies A. s. strelzowi, and the other two, to A. s. desertorum. Based on different index values, the level of genetic polymorphism in the later subspecies was considered to be higher than in the first one. Phylogeographic analysis suggested post-glacial dispersal of flat-headed voles from a single refugium located in Western Altai. Using different techniques, relatively recent colonization of the Central Altai territory was demonstrated (subspecies A. s. strelzowi), which determined low level of genetic variation in this territory.

Multiple refugia and barriers explain the phylogeography of the Valais shrew, Sorex antinorii (Mammalia: Soricomorpha)

The aim of the present study was to investigate the genetic structure of the Valais shrew (Sorex antinorii) by a combined phylogeographical and landscape genetic approach, and thereby to infer the locations of glacial refugia and establish the influence of geographical barriers. We sequenced part of the mitochondrial cytochrome b (cyt b) gene of 179 individuals of S. antinorii sampled across the entire species' range. Six specimens attributed to S. arunchi were included in the analysis. The phylogeographical pattern was assessed by Bayesian molecular phylogenetic reconstruction, population genetic analyses, and a species distribution modelling (SDM)-based hindcasting approach. We also used landscape genetics (including isolation-by-resistance) to infer the determinants of current intra-specific genetic structure. The phylogeographical analysis revealed shallow divergence among haplotypes and no clear substructure within S. antinorii. The starlike structure of the median-joining network is consistent with population expansion from a single refugium, probably located in the Apennines. Long branches observed on the same network also suggest that another refugium may have existed in the north-eastern part of Italy. This result is consistent with SDM, which also suggests several habitable areas for S. antinorii in the Italian peninsula during the LGM. Therefore S. antinorii appears to have occupied disconnected glacial refugia in the Italian peninsula, supporting previous data for other species showing multiple refugia within southern refugial areas. By coupling genetic analyses and SDM, we were able to infer how past climatic suitability contributed to genetic divergence of populations. The genetic differentiation shown in the present study does not support the specific status of S. arunchi.

Phylogeography of an Italian endemic salamander (genus Salamandrina): glacial refugia, postglacial expansions, and secondary contact

The Italian endemic genus Salamandrina has been historically regarded as monotypic but, recently, studies based on both mitochondrial and nuclear markers have indicated the existence of two distinct species of spectacled salamanders: Salamandrina perspicillata, in central and northern Italy, and Salamandrina terdigitata, in southern Italy. We analyzed nucleotide variation at mitochondrial and nuclear genes [cytochrome b, 12S and 16S rRNA, recombination activating gene (RAG 1)] in 223 individuals from 56 locations, aiming to investigate their genetic structure and recent evolutionary histories. Phylogenetic and phylogeographical analyses revealed the existence of three and two genetically distinct groups of populations in northern and southern salamander, respectively. Historical demographic analyses led to the inference of range expansion for both species in the late Pleistocene. During the last glacial stage, each salamander survived in a single refugium, namely the southern in Calabria and the northern in central Italy. At the end of this period, both lineages expanded northward and established secondary contact. Spatial distribution of RAG 1 haplotype variation revealed two differentiated population groups corresponding to the major mitochondrial (mt)DNA clades. Nuclear pattern of introgressive hybridization was more extensive than the highly limited introgression of mtDNA markers. From a conservation standpoint, southern Latium and Calabria proved to be the major genetic diversity reservoirs, thus deserving particular conservation efforts.

Postglacial recolonization of eastern Blacknose Dace, Rhinichthys atratulus (Teleostei: Cyprinidae), through the gateway of New England

During the last ice age, much of North America far south as 40°N was covered by glaciers (Hewitt 2000). About 20,000 years ago, as the glaciers retreated, the hydrologic landscape changed dramatically creating waterways for fish dispersal. The number of populations responsible for recolonization and the regions from which they recolonized are unknown for many freshwater fishes living in New England and southeastern Canada. The Blacknose Dace,Rhinichthys atratulus, is one of the freshwater fish species that recolonized this region. We hypothesize that the earliest deglaciated region, modern-day Connecticut, was recolonized byR. atratulusvia a single founding event by a single population. In this paper, we test this hypothesis phylogenetically with regard to the major drainage basins within Connecticut. The mitochondrial DNA exhibits low nucleotide diversity, high haplotype diversity, and a dominant haplotype found across the state. A small percentage of individuals in the Housatonic drainage basin, however, share a haplotype with populations in New York drainage basins, a haplotype not found elsewhere in Connecticut's drainage basins. We calculated a range for the rate of divergence for NADH dehydrogenase subunit 2 (nd2) and control region (ctr) of 4.43–6.76% and 3.84–8.48% per million years (my), respectively. While this range is higher than the commonly accepted rate of 2% for mitochondrial DNA, these results join a growing list of publications finding high rates of divergence for various taxa (Peterson and Masel 2009). The data support the conclusion that Connecticut as a whole was recolonized initially by a single founding event that came from a single refugium. Subsequently, the Housatonic basin alone experienced a secondary recolonization event.

Southern European glacial refugia: A tale of tales

AbstractInterpretations of current diversity patterns based on the contraction/expansion model forced by climatic oscillations during the last two million years are commonplace in phylogeographic literature. Of the wealth of scientific studies accumulated during the past two decades in Europe, the ones we understand best are those mostly from higher latitudes, probably because patterns were simplified to a great extent by major losses of diversity during glacial periods. In Southern European regions (or in general, in those places where ice effects were less severe) the situation is quite different and to some extent opposite. These regions are referred to as refugia because they are known to contain more genetic diversity than elsewhere. This is not only due to preservation of genotypes that went extinct in other places, however, but also to the intensity and accumulation of a number of processes in a patchy landscape across a varied topography. A lack of general phylogeographic patterns in these regions is one consequence. Speaking of a single refugium to refer to each of the peninsulas, however, is an oversimplification. Even speaking of multiple unconnected refugia does not adequately reflect the complexity of the processes that shaped the current genetic and specific diversity.

Long-term declines of wader populations at the Tagus estuary, Portugal: a response to global or local factors?

SummaryMigratory wader populations face global threats, mainly related to increasing rates of habitat loss and disturbance driven by human activities. To a large extent, the long-term survival of these populations requires the conservation of networks of sites along their migratory flyways. The Tagus estuary, Portugal, is among the most important wetlands for waders in the East Atlantic Flyway. Annual winter wader counts have been carried in this wetland since 1975 and a monthly roost-monitoring programme was implemented in 2007. Wintering populations of three out of the five most abundant species, DunlinCalidris alpina, Grey PloverPluvialis squatarolaand RedshankTringa totanus, showed significant population declines over the past three decades, which are most likely due to the loss and degradation of roost sites as a result of increasing human activity. The situation is unlikely to improve, as a high proportion of the wintering waders use roost sites that are situated in highly urbanised areas with no legal protection. The use of different roost sites by waders is highly variable both temporally and spatially, thus emphasizing the need for a network of good quality roost sites. Additionally, during migration, 60–80% of all the waders of the Tagus estuary concentrate at a single refuge, thus increasing the risk for wader populations during these periods.

Adaptive management in the U.S. National Wildlife Refuge System: Science-management partnerships for conservation delivery

Adaptive management is an approach to recurrent decision making in which uncertainty about the decision is reduced over time through comparison of outcomes predicted by competing models against observed values of those outcomes. The National Wildlife Refuge System (NWRS) of the U.S. Fish and Wildlife Service is a large land management program charged with making natural resource management decisions, which often are made under considerable uncertainty, severe operational constraints, and conditions that limit ability to precisely carry out actions as intended. The NWRS presents outstanding opportunities for the application of adaptive management, but also difficult challenges. We describe two cooperative programs between the Fish and Wildlife Service and the U.S. Geological Survey to implement adaptive management at scales ranging from small, single refuge applications to large, multi-refuge, multi-region projects. Our experience to date suggests three important attributes common to successful implementation: a vigorous multi-partner collaboration, practical and informative decision framework components, and a sustained commitment to the process. Administrators in both agencies should consider these attributes when developing programs to promote the use and acceptance of adaptive management in the NWRS.

Evolutionary history of an alpine shrub Hippophae tibetana (Elaeagnaceae): allopatric divergence and regional expansion

Increasing evidence suggests that geological or climatic events in the past promoted allopatric speciation of alpine plants in the Qinghai-Tibetan Plateau and adjacent region. However, few studies have been undertaken to examine whether such allopatric divergences also occurred within a morphologically uniform species. In the present study, we report the evolutionary history of an alpine shrub species, Hippophae tibetana, based on examining chloroplast DNA (cpDNA) and nuclear ribosomal internal transcribed spacer (ITS) DNA variations. We sequenced two cpDNA fragments (trnL-F and trnS-G) and the nuclear ITS region in 183 individuals collected from 21 natural populations. Ten chlorotypes and 17 ITS types were identified. Phylogenetic analyses of both chlorotypes and ITS sequence variations suggested two distinct lineages distributed in the eastern and western region, respectively. On the basis of the fast and low plant substitution rates, these two lineages were estimated to have diverged from each other between 1 and 4 million years ago, during the period of the major glaciations and orogenic processes. In addition, ITS has undergone the accelerated evolution in two populations in the southern Himalaya isolated by the high mountains with a surprising accumulation of the private variations. The east-west split was also supported by an analysis of molecular variance, which partitioned around 91% of the total cpDNA variance between these two groups of populations. A single chlorotype was found for most populations in eastern or western region, suggesting a recent postglacial expansion within each region. Star-phylogeny and mismatch analyses of all chlorotypes within the eastern group of populations suggested an earlier regional expansion before the Last Glacial Maximum (LGM). The local fixture of the different chlorotypes in multiple populations suggested more than one refugia remained for eastern or western region. Coalescent tests rejected the hypothesis that all current populations originated from a single refugium during the LGM. Instead, they supported hypothesis that two lineages diverged before the late Pleistocene. These findings, when taken together, suggested that this species had experienced long allopatric divergence and recent regional range expansions in response to orogenic processes and the climate changes. The evolutionary history of this shrub species highlights importance of geographical isolations to the intraspecific divergence of alpine plants occurring in the world's ruff. (c) 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102, 37-50.

Allopatric divergence and phylogeographic structure of the plateau zokor (Eospalax baileyi), a fossorial rodent endemic to the Qinghai–Tibetan Plateau

AbstractAim Most species of temperate regions are believed to have shifted to lower latitudes or elevations during the glacial periods of the Quaternary. In this study we test whether this phylogeographic assumption is also true for the plateau zokor (Eospalax baileyi), a fossorial rodent endemic to the climate‐sensitive Qinghai–Tibetan Plateau (QTP), which ranges in elevation from 2600 to 4600 m.Location The QTP of western China.Methods Phylogeographic analyses were conducted based on the mitochondrial cytochrome b gene sequences of 193 individuals from 20 populations over the entire range of the species.Results A total of 54 haplotypes identified in the present study clustered into four geographically correlated clades located in the interior of the QTP (clade A) and at the plateau edge (B, C and D). Molecular calibrations suggest that the interior plateau (A) and plateau‐edge (B–D) clades diverged at 1.2 Ma and that the three plateau‐edge clades diverged between 0.85 and 0.80 Ma. These estimates are concordant with diastrophism and glaciation events in the QTP. Coalescent tests rejected both the hypothesis that all current populations originated from a single refugium at a low elevation during the Last Glacial Maximum (LGM) and the hypothesis that the two lineages diverged during the LGM. The tests instead supported the hypothesis that there were four refugia during the LGM, and that the four clades diverged prior to the late Pleistocene.Main conclusions Our results suggest that Quaternary diastrophisms and glaciations repeatedly promoted allopatric divergence of the plateau zokor into geographical clades, and that these regional clades subsequently persisted at high elevations, rather than migrating to the low‐elevation plateau edge during subsequent glacial ages.

Mitochondrial phylogeography of the edible dormouse (Glis glis) in the western Palearctic region

This study describes in detail the phylogeographic pattern of the edible dormouse (Glis glis) a European rodent with pronounced hibernating behavior. We used sequences of 831 base pairs of the mitochondrial DNA cytochrome-b gene from 130 edible dormice collected at 43 localities throughout its distribution. Our results reveal presence of 3 main haplogroups: Sicilian, South Italian (restricted to the Calabrian region), and European (a widespread lineage corresponding to all remaining western, central, and eastern European populations). Examination of paleontological data confirms refugial regions for G. glis in the 3 Mediterranean peninsulas, although overall low genetic diversity is found. The low diversity of the European lineage is probably the result of a recent expansion (dated around 2,000 years ago) from a single refugium. Other factors, such as the ecological constraints on the species, may have caused genetic bottlenecks that reinforced the low genetic variability of G. glis. This work could have important implications for strategies to conserve the edible dormouse by defining important areas for their conservation. DOI: 10.1644/08-MAMM-A-392R1.1.

Population genetic structure of long-tailed pygmy rice rats (Oligoryzomys longicaudatus) from Argentina and Chile based on the mitochondrial control region

The rodent Oligoryzomys longicaudatus (Bennett, 1832) (Rodentia, Cricetidae) inhabits southern forests of Argentina and Chile, a region severely affected by glaciations during the Pleistocene–Holocene periods. We evaluate here the diversity of the mitochondrial control region to characterize the genetic structure of this species from forests and bushy areas of seven populations from Argentina and four populations from Chile. Statistical analyses showed shallow haplotype trees and mismatch distributions compatible with recent range expansions. The presence of “private” haplotypes indicates that current levels of gene flow among populations of each country would be low to moderate. Significant differences in haplotype frequencies were detected between eastern and western populations, indicating that the Andes mountains would be an effective geographic barrier for gene flow despite the existing valleys that could act as corridors for dispersion. A single clade containing all the haplotypes was recovered in the phylogenetic trees, suggesting postglacial dispersion from a single refugium during the Last Glacial Maximum. The higher effective size and levels of polymorphism in populations from Chile suggest that the refugium was located in this country. The asymmetric gene flow from Chile to Argentina may reflect a recent colonization of the eastern populations.

Phylogeography of 3 North Atlantic Wolffish species (Anarhichas spp.) with Phylogenetic Relationships within the Family Anarhichadidae

Phylogenetic analyses of all 4 wolffish species (Atlantic, Spotted, Northern, and Bering wolffishes) and the Wolfeel were assessed with both mitochondrial (D-loop and ND1) and nuclear (amplified fragment length polymorphism) DNA to resolve relationships within the family Anarhichadidae. Species-specific mitochondrial DNA (mtDNA) mutation rates were estimated based on 2 possible dates of divergence between the Pacific and Atlantic lineages. Phylogeographic patterns within each of the 3 North Atlantic wolffishes were investigated with Markov chain Monte Carlo simulations based on mtDNA to determine whether population size changes occurred following the last glaciation and where wolffishes likely survived glaciation. All 3 species of North Atlantic wolffishes showed evidence of postglacial expansion but did not show evidence of persistence in multiple refugia in both the eastern and western Atlantic Ocean. Rather, the data supported persistence in a single refuge, with postglacial expansion into the rest of the range. Nucleotide diversity, in particular, was low in wolffishes compared with other marine fishes, possibly related to reductions in population sizes during the last glaciation.

Comparative phylogeography of five avian species: implications for Pleistocene evolutionary history in the Qinghai-Tibetan plateau

Pleistocene climate fluctuations have shaped the patterns of genetic diversity observed in extant species. In contrast to Europe and North America where the effects of recent glacial cycles on genetic diversity have been well studied, the genetic legacy of the Pleistocene for the Qinghai-Tibetan (Tibetan) plateau, a region where glaciation was not synchronous with the North Hemisphere ice sheet maxima, remains poorly understood. Here, we compared the phylogeographical patterns of five avian species on the Qinghai-Tibetan plateau by three mitochondrial DNA fragments: the Tibetan snow finch (Montifringilla adamsi), the Blanford's snow finch (Pyrgilauda blanfordi), the horned lark (Eremophila alpestris), the twite (Carduelis flavirostris) and the black redstart (Phoenicurus ochruros). Our results revealed the three species mostly distributed on the platform region of the plateau that experienced population expansion following the retreat of the extensive glaciation period (0.5-0.175 Ma). These results are at odds with the results from avian species of Europe and North America, where population expansions occurred after Last Glacial Maximum (LGM, 0.023-0.018 Ma). A single refugium was identified in a restricted semi-continuous area around the eastern margin of the plateau, instead of multiple independent refugia for European and North American species. For the other two species distributed on the edges of the plateau (the twite and black redstart), populations were maintained at stable levels. Edge areas are located on the eastern margin, which might have had little or no ice cover during the glaciation period. Thus, milder climate may have mitigated demographic stresses for edge species relative to the extremes experienced by platform counterparts, the present-day ranges of which were heavily ice covered during the glaciation period. Finally, various behavioural and ecological characteristics, including dispersal capacities, habitat preference and altitude specificity along with evolutionary history might have helped to shape different phylogeographical structures appearing in these five species.

Sudden expansion of a single brown bear maternal lineage across northern continental Eurasia after the last ice age: a general demographic model for mammals?

The brown bear has proved a useful model for studying Late Quaternary mammalian phylogeography. However, information is lacking from northern continental Eurasia, which constitutes a large part of the species' current distribution. We analysed mitochondrial DNA sequences (totalling 1943 bp) from 205 bears from northeast Europe and Russia in order to characterize the maternal phylogeography of bears in this region. We also estimated the formation times of the sampled brown bear lineages and those of its extinct relative, the cave bear. Four closely related haplogroups belonging to a single mitochondrial subclade were identified in northern continental Eurasia. Several haplotypes were found throughout the whole study area, while one haplogroup was restricted to Kamchatka. The haplotype network, estimated divergence times and various statistical tests indicated that bears in northern continental Eurasia recently underwent a sudden expansion, preceded by a severe bottleneck. This brown bear population was therefore most likely founded by a small number of bears that were restricted to a single refuge area during the last glacial maximum. This pattern has been described previously for other mammal species and as such may represent one general model for the phylogeography of Eurasian mammals. Bayesian divergence time estimates are presented for different brown and cave bear clades. Moreover, our results demonstrate the extent of substitution rate variation occurring throughout the phylogenetic tree, highlighting the need for appropriate calibration when estimating divergence times.

Species‐wide phylogeography of North American mule deer (Odocoileus hemionus): cryptic glacial refugia and postglacial recolonization

Quaternary climatic oscillations greatly influenced the present-day population genetic structure of animals and plants. For species with high dispersal and reproductive potential, phylogeographic patterns resulting from historical processes can be cryptic, overshadowed by contemporary processes. Here we report a study of the phylogeography of Odocoileus hemionus, a large, vagile ungulate common throughout western North America. We examined sequence variation of mitochondrial DNA (control region and cytochrome b) within and among 70 natural populations across the entire range of the species. Among the 1766 individual animals surveyed, we recovered 496 haplotypes. Although fine-scale phylogenetic structure was weakly resolved using phylogenetic methods, network analysis clearly revealed the presence of 12 distinct haplogroups. The spatial distribution of haplogroups showed a strong genetic discontinuity between the two morphological types of O. hemionus, mule deer and black-tailed deer, east and west of the Cascade Mountains in the Pacific Northwest. Within the mule deer lineage, we identified several haplogroups that expanded before or during the Last Glacial Maximum, suggesting that mule deer persisted in multiple refugia south of the ice sheets. Patterns of genetic diversity within the black-tailed deer lineage suggest a single refugium along the Pacific Northwest coast, and refute the hypothesis that black-tailed deer persisted in one or more northern refugia. Our data suggest that black-tailed deer recolonized areas in accordance with the pattern of glacial retreat, with initial recolonization northward along a coastal route and secondary recolonization inland.

Phylogeography of the mitten crab Eriocheir sensu stricto in East Asia: Pleistocene isolation, population expansion and secondary contact

We examined the impact of Pleistocene glacial cycles on geographical distribution and genetic structure of the mitten crab Eriocheir sensu stricto in East Asia using sequence variation of mitochondrial cytochrome c oxidase I and cytochrome b gene segments. Phylogenies revealed four distinct but shallow structured lineages in Eriocheir s. s. Three lineages dominated the East China Sea-Yellow Sea, the Sea of Japan and the South China Sea on the margins of the region, and one lineage occurred on Okinawa Island. This geographical distribution represents a general phylogeographic pattern in East Asia, which is closely associated with the fluctuations of marginal seas and islands during the Pleistocene. The four lineages are estimated to have diverged during the mid-Pleistocene. Demographic expansions were observed in each lineage, starting within the second-to-latest interglacial period in the marginal sea lineages ( approximately 70-130ka) and within the last glacial period in the Okinawa lineage ( approximately 25-80ka). Expansions have probably taken place northward along the coast of the East China Sea-Yellow Sea, following the rise of sea levels. Centered on the southern Korean Peninsula, expansions have likely occurred northward along the west coast and eastward along the south coast of the Sea of Japan. Each marginal sea has served as a single refugium during glacial periods. Two secondary contact regions were identified, one of the East China Sea-Yellow Sea and South China Sea lineages, and another of the East China Sea-Yellow Sea and Sea of Japan lineages. Phylogeography of Eriocheir s. s. provides insights into the evolutionary history and mechanism for generating biodiversity in East Asia.

Spatial cognition in the gerbil: computing optimal escape routes from visual threats

Previous studies in our laboratory have shown that when presented with a sudden stimulus simulating an oncoming predator, Mongolian gerbils can compute the optimal trajectory to a safe refuge, taking into account the position of the threat, the location of a clearly visible refuge, and several other contextual variables as well. In the present studies, the main goal was to explore the abilities of gerbils to use mental representations of spaces that were visually occluded by opaque barriers to compute efficient escape trajectories. In all studies, gerbils were placed into a round open field containing a single refuge. On each trial, an overhead visual stimulus was caused to 'fly' overhead, eliciting robust escape movements from the gerbils. By manipulating the shape and position of a series of opaque barriers that were interposed between the gerbils and the refuge, we were able to show that gerbils can compute the shortest route to an invisible target, even when the available routes to the target are made complex by using elaborate barrier shapes. These findings suggest that gerbils can maintain representations of their locations with respect to salient environmental landmarks and refuges, even when such locations are not continuously visible.