Genetic diversity, structure, and demographic change in tanoak, Lithocarpus densiflorus (Fagaceae), the most susceptible species to sudden oak death in California

Knowledge of population genetic structure of tanoak (Lithocarpus densiflorus) is of interest to pathologists seeking natural variation in resistance to sudden oak death disease, to resource managers who need indications of conservation priorities in this species now threatened by the introduced pathogen (Phytophthora ramorum), and to biologists with interests in demographic processes that have shaped plant populations. We investigated population genetic structure using nuclear and chloroplast DNA (cpDNA) and inferred the effects of past population demographic processes and contemporary gene flow. Our cpDNA results revealed a strong pattern of differentiation of four regional groups (coastal California, southern Oregon, Klamath mountains, and Sierra Nevada). The chloroplast haplotype phylogeny suggests relatively deep divergence of Sierra Nevada and Klamath populations from those of coastal California and southern Oregon. A widespread coastal California haplotype may have resulted from multiple refugial sites during the Last Glacial Maximum or from rapid recolonization from few refugia. Analysis of nuclear microsatellites suggests two major groups: (1) central coastal California and (2) Sierra Nevada/Klamath/southern Oregon and an area of admixture in north coastal California. The low level of nuclear differentiation is likely to be due to pollen gene flow among populations during postglacial range expansion.     

DISCORDANT DISTRIBUTION OF POPULATIONS AND GENETIC VARIATION IN A SEA STAR WITH HIGH DISPERSAL POTENTIAL

Patiria miniata, a broadcast‐spawning sea star species with high dispersal potential, has a geographic range in the intertidal zone of the northeast Pacific Ocean from Alaska to California that is characterized by a large range gap in Washington and Oregon. We analyzed spatial genetic variation across the P. miniata range using multilocus sequence data (mtDNA, nuclear introns) and multilocus genotype data (microsatellites). We found a strong phylogeographic break at Queen Charlotte Sound in British Columbia that was not in the location predicted by the geographical distribution of the populations. However, this population genetic discontinuity does correspond to previously described phylogeographic breaks in other species. Northern populations from Alaska and Haida Gwaii were strongly differentiated from all southern populations from Vancouver Island and California. Populations from Vancouver Island and California were undifferentiated with evidence of high gene flow or very recent separation across the range disjunction between them. The surprising and discordant spatial distribution of populations and alleles suggests that historical vicariance (possibly caused by glaciations) and contemporary dispersal barriers (possibly caused by oceanographic conditions) both shape population genetic structure in this species.     

Phylogeography and host association in a pollinating seed parasite Greya politella (Lepidoptera: Prodoxidae)

Inferring the historical context of ecological diversification is an important step in understanding the way that population-level processes result in a diversity of species and interactions in communities. We performed a phylogeographic analysis of mitochondrial DNA haplotypes from the pollinating seed parasite Greya politella (Lepidoptera: Prodoxidae) in order to determine the degree to which populations were structured according to geographical location and host-plant association. Ninety-eight individuals were sampled from 29 locations ranging from southern California to western Idaho. Restriction-site variation in 87 individuals (27 populations) was screened by digestion with 11 endonucleases, followed by Southern blotting; 38 restriction-site positions were mapped by double digests. Haplotypes were further defined by generating fragments 251 bases in length via PCR, screening them for sequence variation using denaturing gel gradient electrophoresis (DGGE), and sequencing the resulting variants. Parsimony analysis of the resulting 12 restriction-site and 15 sequence haplotypes indicated strong geographical structuring of populations: (i) most populations were monomorphic for haplotype; (ii) haplotypes from California and the Pacific Northwest (Oregon, Washington and Idaho) formed robust monophyletic groups. Population structure was significant both within and between the two regions, as reflected by N _ST. Patterns of host-plant association and haplotype phylogeny suggest that populations have recently undergone host-plant shifts in many different parts of the species range, although the direction and number of host shifts cannot be determined at the present level of sampling resolution.     

Chloroplast DNA intraspecific phylogeography of plants from the Pacific Northwest of North America

Molecular studies of plants from the Pacific Northwest of North America suggest a recurrent pattern of genetic differentiation and geographic structuring. In each of five angiosperms and one fern species representing diverse life histories, cpDNA data indicate two clades of populations that are geographically structured. A northern group comprises populations from Alaska to central or southern Oregon, whereas populations from central Oregon southward to northern California form a southern group. In several of these species, a few populations having southern genotypes may have survived in glacial refugia further north in the Olympic Peninsula, Queen Charlotte Islands, and Prince of Wales Island. Allozyme data reveal a similar pattern of differentiation in several other plants from the Pacific Northwest. North-south partitioning of genotypes has also been reported for several animal species from this region. On a broader geographic scale, northsouth partitioning of genotypes has also been observed in other plants from western North America having a variety of geographic distributions. Some species also display a reduction of genetic variability in the northern portion of their range compared to the south. The data suggest strongly that past glaciation profoundly influenced the genetic architecture of the flora and fauna of the Pacific Northwest. Two alternative hypotheses are advanced to explain the geographic structuring of genotypes. First, past glaciation may have created discontinuities in the geographic distributions of plant species, with populations surviving in several well-isolated northern and southern refugia. Following glaciation, migration of genetically differentiated, once-isolated populations resulted in the formation of a continuous geographic distribution with a major genetic discontinuity. Alternatively, plants survived and subsequently migrated northward from a southern refugium, and a genotype became fixed in one or a few populations at the leading edge of recolonization. Subsequent long-distance dispersal from this leading edge resulted in a relatively uniform northern genotype that differs from the southern genotype(s). Whatever the underlying mechanism, Pleistocence glaciation may have molded the intraspecific genetic architecture of both plants and animals from the Pacific Northwest in a geographically similar manner. Future studies should seek to obtain a comprehensive phylogeography for regions that includes a diversity of both plants and animals.Dedicated to emer. Univ.-Prof. DrFriedrich Ehrendorfer on the occasion of his 70th birthday     

Historical and Contemporary DNA Indicate Fisher Decline and Isolation Occurred Prior to the European Settlement of California

Establishing if species contractions were the result of natural phenomena or human induced landscape changes is essential for managing natural populations. Fishers (Martes pennanti) in California occur in two geographically and genetically isolated populations in the northwestern mountains and southern Sierra Nevada. Their isolation is hypothesized to have resulted from a decline in abundance and distribution associated with European settlement in the 1800s. However, there is little evidence to establish that fisher occupied the area between the two extant populations at that time. We analyzed 10 microsatellite loci from 275 contemporary and 21 historical fisher samples (1880–1920) to evaluate the demographic history of fisher in California. We did not find any evidence of a recent (post-European) bottleneck in the northwestern population. In the southern Sierra Nevada, genetic subdivision within the population strongly influenced bottleneck tests. After accounting for genetic subdivision, we found a bottleneck signal only in the northern and central portions of the southern Sierra Nevada, indicating that the southernmost tip of these mountains may have acted as a refugium for fisher during the anthropogenic changes of the late 19^th and early 20^th centuries. Using a coalescent-based Bayesian analysis, we detected a 90% decline in effective population size and dated the time of decline to over a thousand years ago. We hypothesize that fisher distribution in California contracted to the two current population areas pre-European settlement, and that portions of the southern Sierra Nevada subsequently experienced another more recent bottleneck post-European settlement.     

Genetic structure of desert ground squirrels over a 20-degree-latitude transect from Oregon through the Baja California peninsula

The genetic structure of populations over a wide geographical area should reflect the demographic and evolutionary processes that have shaped a species across its range. We examined the population genetic structure of antelope ground squirrels (Ammospermophilus leucurus) across the complex of North American deserts from the Great Basin of Oregon to the cape region of the Baja California peninsula. We sampled 73 individuals from 13 major localities over this 2500-km transect, from 43 to 22 degrees north. Our molecular phylogeographical analysis of 555 bp of the mitochondrial cytochrome b gene and 510 bp of the control region revealed great genetic uniformity in a single clade that extends from Oregon to central Baja California. A second distinct clade occupies the southern half of the peninsula. The minimal geographical structure of the northern clade, its low haplotype diversity and the distribution of pairwise differences between haplotypes suggest a rapid northward expansion of the population that must have followed a northward desert habitat shift associated with the most recent Quaternary climate warming and glacial retreat. The higher haplotype diversity within the southern clade and distribution of pairwise differences between haplotypes suggest that the southern clade has a longer, more stable history associated with a southern peninsular refugium. This system, as observed, reflects both historical and contemporary ecological and evolutionary responses to physical environmental gradients within genetically homogeneous populations.     

Glacial vicariance in the Pacific Northwest: evidence from a lodgepole pine mitochondrial DNA minisatellite for multiple genetically distinct and widely separated refugia

The Canadian side of the Pacific Northwest was almost entirely covered by ice during the last glacial maximum, which has induced vicariance and genetic population structure for several plant and animal taxa. Lodgepole pine ( Pinus contorta Dougl. ex. Loud.) has a wide latitudinal and longitudinal distribution in the Pacific Northwest. Our main objective was to identify relictual signatures of glacial vicariance in the population structure of the species and search for evidence of distinct glacial refugia in the Pacific Northwest. A maternally inherited mitochondrial DNA minisatellite-like marker was used to decipher haplotype diversity in 91 populations of lodgepole pine located across the natural range. Overall population differentiation was sizeable ( G _ST = 0.365 and R _ST = 0.568). Four relatively homogeneous groups of populations, possibly representative of as many genetically distinct glacial populations, were identified for the two main subspecies, ssp. latifolia and ssp. contorta . For ssp. contorta , one glacial lineage is suggested to have been located at high latitudes and possibly off the coast of mainland British Columbia (BC), while the other is considered to have been located south of the ice sheet along the Pacific coast. For ssp. latifolia , two genetically distinct glacial populations probably occurred south of the ice sheet: in the area bounded by the Cascades and Rocky Mountains ranges, and on the eastern side of the Rockies. A possible fifth refugium located in the Yukon may have also been present for ssp. latifolia . Zones of contact between these ancestral lineages were also apparent in interior and northern BC. These results indicate the role of the Queen Charlotte Islands and the Alexander Archipelago as a refugial zone for some Pacific Northwest species and the vicariant role played by the Cascades and the American Rocky Mountains during glaciation.     

Phylogeography of the mountain chickadee (Poecile gambeli): diversification, introgression, and expansion in response to Quaternary climate change

Since the late 1990s, molecular techniques have fuelled debate about the role of Pleistocene glacial cycles in structuring contemporary avian diversity in North America. The debate is still heated; however, there is widespread agreement that the Pleistocene glacial cycles forced the repeated contraction, fragmentation, and expansion of the North American biota. These demographic processes should leave genetic 'footprints' in modern descendants, suggesting that detailed population genetic studies of contemporary species provide the key to elucidating the impact of the late Quaternary (late Pleistocene-Holocene). We present an analysis of mitochondrial DNA (mtDNA) variation in the mountain chickadee (Poecile gambeli) in an attempt to examine the genetic evidence of the impact of the late Quaternary glacial cycles. Phylogenetic analyses reveal two strongly supported clades of P. gambeli: an Eastern Clade (Rocky Mountains and Great Basin) and a Western Clade (Sierra Nevada and Cascades). Post-glacial introgression is apparent between these two clades in the Mono Lake region of Central California. Within the Eastern Clade there is evidence of isolation-by-distance in the Rocky Mountain populations, and of limited gene flow into and around the Great Basin. Coalescent analysis of genetic variation in the Western Clade indicates that northern (Sierra Nevada/Cascades) and southern (Transverse/Peninsular Ranges) populations have been isolated and evolving independently for nearly 60,000 years.     

Concordant molecular and phenotypic data delineate new taxonomy and conservation priorities for the endangered mountain yellow-legged frog

The mountain yellow-legged frog Rana muscosa sensu lato , once abundant in the Sierra Nevada of California and Nevada, and the disjunct Transverse Ranges of southern California, has declined precipitously throughout its range, even though most of its habitat is protected. The species is now extinct in Nevada and reduced to tiny remnants in southern California, where as a distinct population segment, it is classified as Endangered. Introduced predators (trout), air pollution and an infectious disease (chytridiomycosis) threaten remaining populations. A Bayesian analysis of 1901 base pairs of mitochondrial DNA confirms the presence of two deeply divergent clades that come into near contact in the Sierra Nevada. Morphological studies of museum specimens and analysis of acoustic data show that the two major mtDNA clades are readily differentiated phenotypically. Accordingly, we recognize two species, Rana sierrae , in the northern and central Sierra Nevada, and R. muscosa , in the southern Sierra Nevada and southern California. Existing data indicate no range overlap. These results have important implications for the conservation of these two species as they illuminate a profound mismatch between the current delineation of the distinct population segments (southern California vs. Sierra Nevada) and actual species boundaries. For example, our study finds that remnant populations of R. muscosa exist in both the southern Sierra Nevada and the mountains of southern California, which may broaden options for management. In addition, despite the fact that only the southern California populations are listed as Endangered, surveys conducted since 1995 at 225 historic (1899–1994) localities from museum collections show that 93.3% ( n =146) of R. sierrae populations and 95.2% ( n =79) of R. muscosa populations are extinct. Evidence presented here underscores the need for revision of protected population status to include both species throughout their ranges.     

Limited phylogeographic structure in the flightless ground beetle, Calathus ruficollis, in southern California

The California Floristic Province is home to more than 8000 species of beetles, yet their geographical patterns of supra- and infraspecific diversity remain largely unexplored. In this paper, we investigate the phylogeography and population demographics of a flightless ground beetle, Calathus ruficollis (Coleoptera: Carabidae), in southern California. We sampled 136 specimens from 25 localities divided into 10 populations using a fragment of the mitochondrial cytochrome oxidase I gene. We tested several hypotheses, including the association of geography with particular clades and populations, the degree of differentiation among regions, and the expansion of populations. Parsimony and Bayesian phylogenetic analyses along with nested clade analysis and amova indicate a deep split between the southern Sierra Nevada population and populations south and west. This split corresponds closely to the split between subspecies C. ruficollis ignicollis (southern Sierra Nevada) and C. ruficollis ruficollis . Populations otherwise exhibit limited geographical structure, though F st values indicate some local differentiation. Mismatch distributions and Fu's F s indicate range expansion of several populations, suggesting that some structure may have been obscured by recent exchange. The population of C. ruficollis on Santa Cruz Island, which might have been expected to be isolated, shares several haplotypes with mainland populations, appearing to represent multiple colonizations.     

Contrasting phylogeographical patterns for springtails reflect different evolutionary histories between the Antarctic Peninsula and continental Antarctica

Aim  We examined the genetic structure among populations and regions for the springtails Cryptopygus antarcticus antarcticus and Gomphiocephalus hodgsoni (Collembola) to identify potential historical refugia and subsequent colonization routes, and to examine population growth/expansion and relative ages of population divergence.
Location  Antarctic Peninsula for C. a. antarcticus ; Antarctic continent (southern Victoria Land) for G. hodgsoni .
Methods  Samples were collected from 24 and 28 locations across the Antarctic Peninsula and southern Victoria Land regions for C. a. antarcticus and G. hodgsoni , respectively. We used population genetic, demographic and nested clade analyses based on mitochondrial DNA (cytochrome c oxidase subunit I and subunit II).
Results  Both species were found to have population structures compatible with the presence of historical glacial refugia on Pleistocene (2 Ma–present) time-scales, followed by post-glacial expansion generating contemporary geographically isolated populations. However, G. hodgsoni populations were characterized by a fragmented pattern with several 'phylogroups' (likely ancestral haplotypes present in high frequency) retaining strong ancestral linkages among present-day populations. Conversely, C. a. antarcticus had an excess of rare haplotypes with a much reduced volume of ancestral lineages, possibly indicating historical founder/bottleneck events and widespread expansion.
Main conclusions  We infer that these differences reflect distinct evolutionary histories in each locality despite the resident species having similar life-history characteristics. We suggest that this has predominantly been influenced by variation in the success of colonization events as a result of intrinsic historical glaciological differences between the Antarctic Peninsula and continental Antarctic environments.     

Pseudotsuga menziesii chemical races of California and Oregon

Cortical oleoresin from more than 300 Douglas-fir trees sampled in over 30 localities throughout California and southern Oregon was analyzed for its monoterpene hydrocarbon composition. The results confirmed the existence of a distinct chemical race found in its pure unhybridized form throughout the Sierra Nevada as well as near Lompoc in the Purissima Hills of coastal California. Chemical features characterizing this race included near-absence of sabinene and presence of larger amounts of - and β-pinenes in the cortical essential oil. Elsewhere in California and southernmost Oregon, the Sierra Nevada race has extensively intergraded with the coastal Douglas-fir race from western Oregon, Washington, and British Columbia (var. menziesii), a race producing large amounts of sabinene and lesser amounts of - and β-pinenes. On the basis of chemical composition, the Sierra Nevada race was judged to be closer to the inland Douglas-fir races (var. glauca) than to the coastal race. Evolution of the Sierra Nevada race is explained by the emergence of the Great Basin arid regions during the PliocenePleistocene epochs which disjoined Western Douglas-fir populations from their eastern inland counterparts.     

Phylogeography of Pseudacris regilla (Anura: Hylidae) in western North America, with a proposal for a new taxonomic rearrangement

The Baja California populations of Pseudacris regilla, a widespread species in Western North America ranging from British Columbia to southern Baja California, are characterized by extensive geographic fragmentation. We performed phylogeographic and historical demographic analyses on 609 bp of the cytochrome b mitochondrial gene of 110 individuals representing 28 populations to determine the relative influences of current and historical processes in shaping the present distribution of genetic diversity on the Baja California Peninsula. Haplotypes from this area were nested in a clade with three well-differentiated groups. Two of these groups are from Baja California Sur and another is from California and Baja California. The estimated date for the split of these groups, between 0.9-1 Ma, fits with previously proposed hypotheses of vicariance due to different transpeninsular seaways, although successive population fragmentation and expansion due to climatic oscillations during Pleistocene glaciations cannot be discarded. Historical demographic analyses detected signs of past population expansions, especially in the southernmost group. With respect to populations north of this region, two older clades were identified, one with haplotypes mainly distributed in central California, and the other corresponding to the northern half of the species range, in what apparently is a recurrent pattern in the Pacific coast of North America. Based on the concordance between mt-DNA and available allozyme data indicating that these species have a long independent evolutionary history, we propose to consider the three major clades as distinct species: P. regilla, P. pacifica, and P. hypochondriaca.     

Long-distance dispersal vs vicariance: the origin and genetic diversity of alpine plants in the Spanish Sierra Nevada

Here, we investigated the origin and genetic diversity of four alpine plant species co-occurring in the Spanish Sierra Nevada and other high mountains in south-western Europe by analysis of amplified fragment length polymorphisms (AFLPs). In Kernera saxatilis, Silene rupestris and Gentiana alpina we found intraspecific phylogroups corresponding to mountain regions as predicted by the vicariance hypothesis. Moreover, genetic distances between Sierra Nevada and Pyrenees populations were always higher than those between populations from the Pyrenees and the south-western Alps/Massif Central. This suggests successive disruption of gene exchange between mountain ranges as postglacial climatic warming proceeded from south to north. In Papaver alpinum, our data indicate that a central Pyrenean population arose via long-distance dispersal from the Sierra Nevada, and that vicariant separation events between the Sierra Nevada and the Pyrenees and between the Pyrenees and the south-western Alps occurred simultaneously. Overall, Sierra Nevada populations of all species investigated here preserve unexpectedly high (or not exceptionally reduced) genetic diversity. This testifies to the important influence of long-term isolation, i.e. vicariance, on genetic diversity through fostering the accumulation of new mutations and/or the fixation of ancestral ones.     

Streaked horned lark <Emphasis Type="Italic">Eremophila alpestris strigata</Emphasis> has distinct mitochondrial DNA

The Streaked Horned Lark (STHL; Eremophila alpestris strigata) is a federal candidate for listing under the Endangered Species Act. We evaluated the conservation status and level of genetic diversity of the STHL using the complete mitochondrial ND2 gene. We sampled 32 STHLs from the southern Puget Sound region, the Pacific coast, and Whites Island in the Columbia River of Washington, and additional 68 horned larks from Alaska, alpine and eastern Washington, Oregon, California, and Asia (outgroups). Our Maximum Likelihood analysis of 32 haplotypes identified three geographically concordant clades in Pacific coast states: Pacific Northwest (alpine and eastern Washington, Alaska), Pacific Coast (western Washington, California), and Great Basin (eastern Oregon). Each of the three clades was supported by bootstrap values ≥86%. The distance among them varied from 0.72 to 0.79% nucleotide divergence excluding intraclade variation. The relationship among the clades was not resolved. AMOVA also showed significant structuring of haplotypes among the three clades. Differences among clades accounted for 75.7% of sequence variation, differences among localities within clades accounted for 12.1%, and differences among individuals within localities accounted for the remaining 12.2%. Although STHL populations were closely related to the Californian sample, they appeared unique and isolated. All pairwise F _st values involving the STHL samples were significant (except between themselves). STHLs appear to have remarkably low genetic diversity; all 32 STHLs shared the same haplotype. Even with small sample sizes, all other localities had multiple haplotypes. Because the STHL appears to be unique and isolated, and to have little genetic diversity our data suggest it should be a conservation priority.     

Inferring Geographic Isolation of Wolverines in California Using Historical DNA

ABSTRACT Delineating a species' geographic range using the spatial distribution of museum specimens or even contemporary detection-non-detection data can be difficult. This is particularly true at the periphery of a species range where species' distributions are often disjunct. Wolverines (Gulo gulo) are wide-ranging mammals with discontinuous and potentially isolated populations at the periphery of their range. One potentially disjunct population occurred in the Sierra Nevada Mountains, California, USA, and appears to have been extirpated by the 1930s. Many early 20th century naturalists believed that this population was connected to other populations occurring in the Cascade Range of northern California, Oregon, and Washington, USA, but a recent analysis of historical records suggests that California wolverines were isolated from other populations in North America. We used DNA extracted from museum specimens to examine whether California wolverines were isolated. Both nuclear and mitochondrial DNA data indicate that California wolverines were genetically distinct from extant populations, suggesting long-term isolation. We identified 2 new control region (mitochondrial DNA) haplotypes located only within California. We used these data and referenced sequences from the Rocky Mountains, USA, to make inferences regarding potential wolverine translocations into California. In addition, we used these genetic data to make inferences about wolverine conservation throughout western North America.     

Genetic evidence for the persistence of the critically endangered Sierra Nevada red fox in California

California is home to both the native state-threatened Sierra Nevada red fox (Vulpes vulpes necator), which historically inhabited high elevations of the Sierra Nevada and Cascade mountains, and to multiple low-elevation red fox populations thought to be of exotic origin. During the past few decades the lowland populations have dramatically expanded their distribution, and possibly moved into the historic range of the native high-elevation fox. To determine whether the native red fox persists in its historic range in California, we compared mitochondrial cytochrome-b haplotypes of the only currently-known high-elevation population (n = 9 individuals) to samples from 3 modern lowland populations (n = 35) and historic (1911–1941) high-elevation (n = 22) and lowland (n = 7) populations. We found no significant population differentiation among the modern and historic high-elevation populations (average pairwise F _ST = 0.06), but these populations differed substantially from all modern and historic lowland populations (average pairwise F _ST = 0.52). Among lowland populations, the historic and modern Sacramento Valley populations were not significantly differentiated from one another (F _ST = −0.06), but differed significantly from recently founded populations in the San Francisco Bay region and in southern California (average pairwise F _ST = 0.42). Analysis of molecular variance indicated that 3 population groupings (mountain, Sacramento Valley, and other lowland regions) explained 45% of molecular variance (F _CT = 0.45) whereas only 4.5% of the variance was partitioned among populations within these groupings (F _SC = 0.08). These findings provide strong evidence that the native Sierra Nevada red fox has persisted in northern California. However, all nine samples from this population had the same haplotype, suggesting that several historic haplotypes may have become lost. Unidentified barriers have apparently prevented gene flow from the Sacramento Valley population to other eastern or southern populations in California. Future studies involving nuclear markers are needed to assess the origin of the Sierra Nevada red fox and to quantify levels of nuclear gene flow.     

Molecular evidence for Pleistocene glacial cycles driving diversification of a North American desert spider, Agelenopsis aperta

The influence of historical climatic vs. geological changes on species diversification patterns was investigated in a widely distributed North American desert spider, Agelenopsis aperta (Araneae: Agelenidae), with particular reference to Pleistocene glacial cycles and earlier patterns of mountain building. Levels of sequence divergence obtained from the mitochondrial gene, cytochrome oxidase I, dated to the Pleistocene, eliminating Rocky Mountain orogeny as a cause of diversification, as orogeny ended 4 million years ago. The results of phylogenetic and network analyses showed the presence of three geographically defined clades, which were consistent with the presence of at least three glacial refugia: (i) east of the Rocky Mountains; (ii) between the Rocky Mountains and Sierra Nevadas; and (iii) west of the Sierra Nevadas. In addition, populations within the Rocky Mountains exhibited significantly lower genetic diversity than populations east of the Rocky Mountains and the haplotypes found within the Rockies were a subset of eastern haplotypes. These patterns suggest that a post-Pleistocene range expansion occurred out of an eastern glacial refugium into the Rocky Mountains. Examination of phylogeographical studies of other North American desert taxa indicated that mountain building explained diversification patterns more effectively for some taxa but Pleistocene climate change was more important for others, including A. aperta.     

Geographic differentiation of monoterpenes from Abies procera and Abies magnifica

Cortical oleoresins from 352 Abies magnifica and A. procera trees, collected in 35 localities, were analysed for composition of their monoterpene fractions. The monoterpenes were composed primarily of -pinene, β-pinene, limonene and β-phellandrene, with camphene and 3-carene appearing only sporadically in larger amounts. Limonene, β-phellandrene, -pinene and 3-cerene were used to study the transitional populations in northern California and southern and central Oregon and the transitional-like populations in southern Sierra Nevada. The populations segregated latitudinally into three related clusters—above 44° (A.procera), between 44° and 40° (transitional), and below 40° (A.magnifica). A.magnifica from southern Sierra Nevada and A.magnifica from Mtn. Shasta differed in a number of parameters, with southern Sierra Nevada populations being chemically much closer to typical A. magnifica from central and northern Sierra Nevada. While monoterpene and morphological data did not allow a clear Interpretation of the status of the transitional and southern populations, the paleobotanical evidence favored the recent evolution over introgression. The desirability of additional studies was indicated.     

Geological barriers and restricted gene flow in the holarctic skipper Hesperia comma (Hesperiidae)

Patterns of genetic variation within a species may be a consequence of historical factors, such as past fragmentation, as well as current barriers to gene flow. Using sequence data from the mitochondrial cytochrome oxidase subunit II region (COII) and the nuclear gene wingless, we conducted a phylogeographical study of the holarctic skipper Hesperia comma to elucidate patterns of genetic diversity and to infer historical and contemporary processes maintaining genetic variation. One hundred and fifty-one individuals were sampled from throughout North America and Eurasia, focusing on California and adjacent regions in the western United States where morphological diversity is highest compared to the rest of the range. Analyses of sequence data obtained from both genes revealed a well-supported division between the Old and New World. Within western North America, wingless shows little geographical structure, while a hierarchical analysis of genetic diversity of COII sequences indicates three major clades: a western clade in Oregon and Northern California, an eastern clade including the Great Basin, Rocky Mountains and British Columbia, and a third clade in southern California. The Sierra Nevada and the Transverse Ranges appear to be the major barriers to gene flow for H. comma in the western United States. Relatively reduced haplotype diversity in Eurasia compared to North America suggests that populations on the two continents have been affected by different historical processes.