Phylogeography of Supralittoral Rocky Intertidal Ligia Isopods in the Pacific Region from Central California to Central Mexico

Background

Ligia isopods are widely distributed in the Pacific rocky intertidal shores from central California to central Mexico, including the Gulf of California. Yet, their biological characteristics restrict them to complete their life cycles in a very narrow range of the rocky intertidal supralittoral. Herein, we examine phylogeographic patterns of Ligia isopods from 122 localities between central California and central Mexico. We expect to find high levels of allopatric diversity. In addition, we expect the phylogeographic patterns to show signatures of past vicariant events that occurred in this geologically dynamic region.

Methodology/Principal Findings

We sequenced two mitochondrial genes (Cytochrome Oxidase I and 16S ribosomal DNA). We conducted Maximum Likelihood and Bayesian phylogenetic analyses. We found many divergent clades that, in general, group according to geography. Some of the most striking features of the Ligia phylogeographic pattern include: (1) deep mid-peninsular phylogeographic breaks on the Pacific and Gulf sides of Baja peninsula; (2) within the Gulf lineages, the northern peninsula is most closely related to the northern mainland, while the southern peninsula is most closely related to the central-southern mainland; and, (3) the southernmost portion of the peninsula (Cape Region) is most closely related to the southernmost portion of mainland.

Conclusions/Significance

Our results shed light on the phylogenetic relationships of Ligia populations in the study area. This study probably represents the finest-scale phylogeographic examination for any organism to date in this region. Presence of highly divergent lineages suggests multiple Ligia species exist in this region. The phylogeographic patterns of Ligia in the Gulf of California and Baja peninsula are incongruent with a widely accepted vicariant scenario among phylogeographers, but consistent with aspects of alternative geological hypotheses and phylo- and biogeographic patterns of several other taxa. Our findings contribute to the ongoing debate regarding the geological origin of this important biogeographic region.     

Biogeography and endemism of ants (Hymenoptera: Formicidae) in Baja California, Mexico: afirst overview

Abstract Aims Ants (Hymenoptera: Formicidae) of the Baja California peninsula are poorly known, with information based largely on scattered museum and literature records. We provide the first comprehensive account of ant species occurring on the peninsula, we examine distribution patterns, and we assess the ‘peninsular effect’ which predicts that species richness declines from the base to the tip of a peninsula. Location Peninsula of Baja California, Mexico. Methods Data collection involved examining, identifying and recording label data from c. 2350 series of ants. These records provide a provisional, if incomplete, species list. We applied the incidence‐based estimator, Chao‐2, to our data base of specimen records to estimate the total number of ant species on the peninsula. We assessed endemism by comparing our peninsular species list to those from adjacent states. The peninsular effect was tested by comparing genus and species level richness between the two states of Baja California, and across five latitudinal blocks. Results We document 170 native ant species in thirty‐three genera, plus six non‐native species, in Baja California. It seems likely that additional species remain to be discovered: the Chao‐2 estimator of species richness, at 206.0 species, is about 20% higher than our observed species richness. About 30% of the species and 20% of the genera are restricted within Baja California to the relatively mesic California Floristic Province of north‐western Baja California. Nearly all of these species also occur in California. Forty‐seven species (27.6%) are peninsula endemics. Using our entire data set, the peninsular effect appears to be strong, with about twice as many species in the northern state of Baja California than are recorded from the southern state of Baja California Sur; the ratio of genera is 33 to 24. However, this effect becomes weak at the species level and absent at the genus level when minimizing habitat effects by omitting species restricted to the California Floristic Province. At a finer scale, across latitudinal blocks of about 1.9°, the number of species declines towards central portions of the peninsula and then increases in the Cape Region. Nine ant species display strongly disjunct distributions, and these occur in two general patterns: peninsula disjuncts and peninsula–mainland disjuncts. Main conclusions The Baja California peninsula supports a diverse and distinctive ant fauna, with the proportion of endemic species similar to that displayed by plants. Patterns of species and genus richness across the five latitudinal blocks provide poor support for the peninsular effect. Moreover, habitat diversity, especially that related to topographic relief, appears to be the most important factor affecting the gradient of ant species richness in Baja California. Additional collections are needed to develop a more complete species list and to determine the boundaries and status of many species. Nevertheless, the present data base provides a useful starting point for understanding the evolution of ant assemblages in Baja California and for comparison with peninsular patterns in other taxa.     

Two new Marina (Leguminosae) from the southern Baja California peninsula, Mexico

Marina victoriae andM. brevis are newly described species from the southern Baja California peninsula, Mexico. The former is a perennial herb locally abundant in the eroded hillsides at the foothills of Sierra de la Victoria, where an oak woodland and a dry tropical forest meet. The latter is an elusive ephemeral inhabiting a narrow strip of land between the mangroves and the desert scrub on an island in the Gulf of California. These two new species belong to the seriesChrysorrhizae, a group native to the Gulf of California basin.     

Colour variation is incongruent with mitochondrial lineages: cryptic speciation and subsequent diversification in a Gulf of California reef fish (Teleostei: Blennioidei)

The Gulf of California endemic reef fish, Acanthemblemaria crockeri (Blennioidei, Chaenopsidae), reportedly has two colour morphs, one with melanic lateral spots ('Gulf' morph) and one with orange spots ('Cape' morph). In this study, we recorded colour morph in both males and females and collected mitochondrial DNA sequence data for cytochrome c oxidase I (COI) and tRNA-Pro/D-loop of specimens from throughout the Gulf to explore the genetic basis of the colour morphs. Two highly divergent (HKY + I distance = 11.9% for COI), reciprocally monophyletic lineages were identified, consistent with the presence of two parapatric species. A 30-km gap between the distributions of mitochondrial lineages roughly corresponds to a hypothesized former seaway across the Baja California peninsula north of La Paz, although the estimated divergence time (1.84 million years ago) is more recent than the hypothetical seaway (3–4 million years ago). Surprisingly, the distribution of mitochondrial species is not congruent with the distribution of either male or female colour morphs. Our analysis also revealed significant population differentiation within both species and no shared haplotypes among populations. The northern Gulf species includes four populations (NB, CB, NM and CM) corresponding to northern and central Baja and northern and central mainland sites, while the Cape species includes two populations (SB and SM) corresponding to the Baja and mainland sides of the southern Gulf. The NB/CB division corresponds to a hypothesized Plio–Pleistocene mid-peninsular seaway. The level of genetic divergence documented in this lineage is extraordinary for a marine fish with a pelagic larval stage within a semi-enclosed basin.     

Morphology of Pyrodinium bahamense Plate (Dinoflagellata) near Isla San José, Gulf of California, Mexico

The morphology of Pyrodinium bahamense south of Isla San José in the Gulf of California, Mexico was described using light and scanning electron microscopy. Morphological and dimensional examination of whole cells and thecal patterns were conducted to discriminate between P. bahamense var. compressum and P. bahamense var. bahamense. Microscopic examinations confirmed that specimens of P. bahamense from Isla San José are closely related to P. bahamense var. bahamense. The main average dimensions were: cells 41.9 μm long, 43.8 μm wide, 55.6 μm amplitude, apical horn 7.2 μm long, and the left antapical spine 19.5 μm long. The highest cell concentration was 240 cells l⁻¹ and occurred with a bloom of Pseudo-nitzschia spp. The presence of this dinoflagellate off the east coast of the Baja California Peninsula had not been previously reported. Recently, P. bahamense from Florida tested positive for saxitoxins. Paralytic shellfish poisoning from this species has occurred in Mexican waters. Here, significant aspects of taxonomy, toxicology, life cycle, and ecology are discussed.     

Tempo and mode of speciation in the Baja California disjunct fish species Anisotremus davidsonii

The Baja California region provides a natural setting for studying the early mechanisms of allopatric speciation in marine systems. Disjunct fish populations from several species that occur in the northern Gulf of California and northern Pacific coast of Baja California, but are absent from its southern shores, were previously shown to be genetically isolated, making them excellent candidates for studying allopatry. In addition, one of these species, the sargo Anisotremus davidsonii, has two pairs of congeneric Panamic trans-isthmian geminate species that allow for internal molecular clock calibration. Phylogeographic and demographic approaches based on mitochondrial (cytochrome b) and nuclear (S7 ribosomal protein) sequences showed that A. davidsonii entered the gulf from the south, and later colonized the Pacific coast, approximately 0.6-0.16 million years ago. Pacific coast colonization may have used a route either around the southern cape of Baja California or across the peninsula through a natural seaway. However, while several seaways have been described from different geological times, none matches the dates of population disjunction, yet much geological work remains to be done in that area. At the present time, there is no evidence for dispersal around the southern tip of the Baja California Peninsula. Signatures of incipient allopatric speciation were observed, such as the reciprocal monophyly of disjunct populations for the mitochondrial marker. However, other characteristics were lacking, such as a strong difference in divergence and coalescence times. Taken together, these results suggest that disjunct populations of A. davidsonii may be consistent with the earliest stages of allopatric speciation.     

Cryptic vicariance in Gulf of California fishes parallels vicariant patterns found in Baja California mammals and reptiles

Comparisons across multiple taxa can often clarify the histories of biogeographic regions. In particular, historic barriers to movement should have affected multiple species and, thus, result in a pattern of concordant intraspecific genetic divisions among species. A striking example of such comparative phylogeography is the recent observation that populations of many small mammals and reptiles living on the Baja California peninsula have a large genetic break between northern and southern peninsular populations. In the present study, I demonstrate that five species of near-shore fishes living on the Baja coastline of the Gulf of California share this genetic pattern. The simplest explanation for this concordant genetic division within both terrestrial and marine vertebrates is that the Baja Peninsula was fragmented by a Plio-Pleistocene marine seaway and that this seaway posed a substantial barrier to movement for near-shore fishes. For some fish species, the signal of this vicariance in mtDNA has been eroded by gene flow and is not evident with classic, equilibrium measures of population structure. Yet, significant divisions are apparent in coalescent analyses that jointly estimate divergence with gene flow. The genetic divisions within Gulf of California fishes also coincide with recognized biogeographic regions based on fish community composition and several environmental factors. It is likely that adaptation to regional environments and present-day oceanographic circulation limit gene exchange between biogeographic regions and help maintain evidence of past vicariance.     

Frond elongation rates of shallow waterMacrocystis pyrifera (L.) Ag. In northern Baja California,Mexico

Macrocystis pyrifera (L.) Ag.frond elongation rates were measured during autumn-winter, spring and summer in a shallow water (7.5 m depth) kelp bed in Bahía Papalote, northern Baja California. Frond elongation was maximum during spring and minimum during winter. Frond growth was significantly correlated with solar radiation, and was highest in the smallest size fronds (0–2 m). Average frond elongation rate varied between 0.3–11% d^–1 during the study period. The relationship between average frond length and elongation rate followed an exponential curve with a negative slope during autumn-winter and summer, but was best described by a straight line during spring. Standard growth rates were obtained by a graphic method. Standard growth rates had intermediate values between those reported for southern California and southern Baja California.M. pyrifera growth cycle shows a different trend from what has been previously reported.     

Phylogeography of the frog Leptodactylus validus (Amphibia: Anura): Patterns and timing of colonization events in the Lesser Antilles

The frog Leptodactylus validus occurs in northern South America, Trinidad and Tobago, and the southern Lesser Antilles (Grenada and St. Vincent). Mitochondrial DNA sequences were used to perform a nested clade phylogeographic analysis (NCPA), to date colonization events, and to analyze colonization patterns using on a relaxed molecular clock and coalescent simulations. L. validus originated on the mainland and first colonized Trinidad with subsequent independent colonizations of Tobago and the Lesser Antilles from Trinidad. The NCPA suggests a historical vicariant event between populations in Trinidad and Tobago from those in the Lesser Antilles. The colonization of Trinidad occurred 1 million years ago (mya) and the colonization of the Lesser Antillean islands occurred 0.4 mya. The coalescent approach supported the scenario where L. validus dispersed from Trinidad to St. Vincent and from there to Grenada, a dispersal event that could have been mediated by human introduction as recent as 1600 years ago.     

Island biogeography of bats in Baja California, Mexico: patterns of bat species richness in a near-shore archipelago

Aim We studied the relationship between the size and isolation of islands and bat species richness in a near‐shore archipelago to determine whether communities of vagile mammals conform to predictions of island biogeography theory. We compared patterns of species richness in two subarchipelagos to determine whether area per se or differences in habitat diversity explain variations in bat species richness. Location Islands in the Gulf of California and adjacent coastal habitats on the Baja California peninsula in northwest Mexico. Methods Presence–absence surveys for bats were conducted on 32 islands in the Gulf of California using acoustic and mist‐net surveys. We sampled for bats in coastal habitats of four regions of the Baja peninsula to characterize the source pool of potential colonizing species. We fitted a semi‐log model of species richness and multiple linear regression and used Akaike information criterion model selection to assess the possible influence of log₁₀ area, isolation, and island group (two subarchipelagos) on the species richness of bats. We compared the species richness of bats on islands with greater vegetation densities in the southern gulf (n = 20) with that on drier islands with less vegetation in the northern gulf (n = 12) to investigate the relationship between habitat diversity and the species richness of bats. Results Twelve species of bats were detected on islands in the Gulf of California, and 15 species were detected in coastal habitats on the Baja peninsula. Bat species richness was related to both area and isolation of islands, and was higher in the southern subarchipelago, which has denser vegetation. Log₁₀ area was positively related to bat species richness, which increased by one species for every 5.4‐fold increase in island area. On average, richness declined by one species per 6.25 km increase in isolation from the Baja peninsula. Main conclusions Our results demonstrate that patterns of bat species richness in a near‐shore archipelago are consistent with patterns predicted by the equilibrium theory of island biogeography. Despite their vagility, bats may be more sensitive to moderate levels of isolation than previously expected in near‐shore archipelagos. Differences in vegetation and habitat xericity appear to be associated with richness of bat communities in this desert ecosystem. Although observed patterns of species richness were consistent with those predicted by the equilibrium theory, similar relationships between species richness and size and isolation of islands may arise from patch‐use decision making by individuals (optimal foraging strategies).     

Geographical variation in population demography and life history traits of Tecate cypress (<Emphasis Type="Italic">Hesperocyparis forbesii</Emphasis>) suggests a fire regime gradient across the USA–Mexico border

Plant adaptations to fire often display spatial heterogeneity associated with geographical variation in fire regime. We examined whether populations of the Tecate cypress (Hesperocyparis forbesii Adams) in southern California and northern Baja, Mexico, exhibited spatial heterogeneity in cone serotiny, in other life history traits associated with fire-adaptation, and in population demographic structure, to assess a putative difference in fire regime across the USA–Mexico border. Demographic data, tree life history data, and tree ring series were used to compare the demographic structure and life history traits of three populations in southern California with three populations in northern Baja California. In Baja populations, a greater number of tree size classes were present (χ ² = 12,589; P < 0.05), cone serotiny was more facultative (Mann–Whitney U = 58, P < 0.05), and young adult trees had a higher reproductive output (Mann–Whitney U = 2.65, P < 0.05), suggesting that a difference in fire regime between southern California and northern Baja has existed long enough (ca 8000 years) to drive microevolutionary divergence between the two sets of populations, and is not solely the result of 20th century differences in fire management policies across the international border. The transitional area between the two different fire regimes does not appear to coincide with the border itself but may lie in a zone of ecological transition south of Ensenada. The range of phenotypic variation observed within the Tecate cypress metapopulation suggests this species has the capacity to adapt to future environmental changes.     

Uranium-series ages of marine terraces,La Paz Peninsula,Baja California Sur,Mexico

Uranium-series dating of coral samples from raised marine terrace deposits between 1.5 and 10 m above sea level in the La Paz Peninsula area, Baja California Sur, yielded ages between 123 ka and 138 ka that are in agreement with previously reported results. The stratigraphy and ages of marine units near the El Coyote Arroyo indicate the presence of two high stands of the sea during the last interglacial or oxygen isotope substage 5e at about 140 ka and 123 ka. Accepting 5 m for the sea level during the last interglacial transgression, we calculate average uplift rates for the marine terraces of about 70 mm/ka and 40 mm/ka. These slow rates of uplift indicate a relative stability of the La Paz peninsula area for the past 140 000 years. In contrast, areas of Baja California affected by major faultf experienced higher rates of uplift. Rockwell et al. (1987) reported vertical uplift rates of 180 to 300 mm/ka at Punta Banda within the Aqua Blanea fault zone in northern Baja California.     

Historical biogeography, phylogenetic relationships and intraspecific diversity of agamid lizards in the Central Asian deserts of Kazakhstan and Uzbekistan

The Central Asian agamid lizards are ecologically and morphologically diverse, occurring across a broad range of desert environments in this biogeographically important region. It is probable that past climatic shifts have significantly influenced the diversification patterns and distributions of the agamid lizards of this region. To assess this within a phylogenetic framework we sequenced a 1200 bp region of mitochondrial DNA and a 1200 bp nuclear gene (RAG-1), incorporating both inter- and intraspecific sampling across Central Asian agamids. Our topology and divergence time estimates support an Eocene origin of the Agaminae subfamily on the Indian subcontinent, coinciding with the collision of India into Eurasia. The onset of aridification in Central Asia during the Late Oligocene, resulting from the retreat of the Paratethys Sea and the intensified uplift of the Tibetan–Himalayan complex, probably played an important role in the diversification of Phrynocephalus, one of the three genera studied. Intensification of aridity and geologic events in the Plio-Pleistocene and Quaternary glacial cycling probably had a significant influence on intraspecific diversification patterns within Phrynocephalus.     

Geographic structuring of chloroplast DNA genotypes inTiarella trifoliata (Saxifragaceae)

Tiarella trifoliata comprises varietieslaciniata, trifoliata, andunifoliata, and is distributed from southeastern Alaska to northern California. We analyzed restriction site variation of chloroplast DNA (cpDNA) using 23 endonucleases in 76 populations representing the entire geographic range of the species and the three recognized varieties. We also employed comparative restriction site mapping of PCR-amplified chloroplast DNA fragments using 16 restriction endonucleases. This species exhibits low cpDNA restriction site variation. No differentiation is evident among varieties of this species based on cpDNA data; some plants of each variety were characterized by each of the two major cpDNA types detected. The two major cpDNA clades, which differ by only a single restriction site mutation, are geographically structured. A northern clade comprises populations from Alaska to central Oregon; most populations analyzed from southern Oregon and California form a southern clade. Populations that possess the typical northern cpDNA type also occur disjunctly to the south at high elevations in the Siskiyou—Klamath Mountain area of southern Oregon and northern California. Conversely, the southern cpDNA type is found disjunctly to the north in the Olympic Peninsula of Washington. Both geographic areas characterized by disjunct cytoplasms are considered glacial refugia.Tiarella trifoliata joins two other species,Tolmiea menziesii andTellima grandiflora, in having well-demarcated northern and southern cpDNA lineages. All three species have similar life-history traits and geographic distributions. We suggest that glaciation may have played a major role in the formation of the cpDNA discontinuities present in these three taxa. The pronounced relationship between cpDNA variation and geographic distribution suggests the potential applicability of intraspecific phylogeography to plants via the analysis of intraspecific cpDNA variation. These three examples also join a rapidly growing data base which indicates that cytoplasms are often geographically structured within species and species complexes.     

Use of Museum Specimens to Refine Historical Pronghorn Subspecies Boundaries

Endangered Sonoran (Antilocapra americana sonoriensis) and Peninsular (A. a. peninsularis) pronghorn persist largely because of captive breeding and reintroduction efforts. Recovery team managers want to re-establish pronghorn in their native range, but there is currently uncertainty regarding the subspecies status of extinct pronghorn populations that historically inhabited southern California, USA, and northern Baja California, Mexico. To address this uncertainty, we genotyped museum specimens and conducted phylogenetic and population genetic analyses of historical data in the context of 3 contemporary pronghorn populations. The historical northern Baja California pronghorn share the most ancestry with contemporary Peninsular pronghorn, whereas pronghorn in southern California share more ancestry with contemporary American (A. a. americana) pronghorn. For reintroductions into northern Baja California, the Peninsular subspecies is more appropriate based on museum genetic data. For reintroductions into Southern California, ecological and genetic factors are both important, as the subspecies most genetically related to historical populations (American) may not be well-adapted to the hot, low-elevation deserts of the reintroduction area. © 2019 The Wildlife Society.     

Genetic isolation and evolutionary history of oases populations of the Baja California killifish,Fundulus lima

The Baja California killifish, Fundulus lima, is found in six desert oases of the southern Baja California Peninsula, Mexico. The recent introduction of exotic fishes, particularly redbelly tilapia, have impacted the ecology of Fundulus lima such that it is now endangered. Plans of relocating F. lima to bodies of freshwater that are free of exotics have been proposed, however little is know about the genetic identity of the current populations. In this study, we examined the mitochondrial control region of F. lima samples from 4 oases, and in addition, compared these samples to their sister species, the California killifish F.␣parvipinnis. Using a combination of phylogenetic and coalescent approaches, we were able to determine that the two subspecies of the California killifish, F. p.␣brevis, and F. p. parvipinnis, and F. lima form an unresolved trichotomy that diverged between 200,000 years and 400,000 years ago. The one F. lima individual that we were able to collect in the southernmost oasis grouped with the southern subspecies of the California killifish, F. parvipinnis brevis. In contrast, we found that the 3 northern oases grouped together in a “Fundulus lima” clade. Each oasis is genetically distinct, yet there is no evidence of a␣marked genetic bottleneck in any populations (Haplotype diversity between 0.5 and 0.8). Future relocation plans will therefore need to be done cautiously to preserve the genetic identity of the original populations.     

Diversification patterns of pebble-mimic dragons are consistent with historical disruption of important habitat corridors in arid Australia

The pebble-mimic dragon lineage of Tympanocryptis is widely distributed in the stony, or ‘gibber’, deserts of Australia but is noticeably absent from intersecting areas of sand deserts. Past fluctuations in the extent and configuration of sandy desert habitat barriers are likely to have been an import factor promoting genetic differentiation in this group. We sequenced a 1400 bp region of mitochondrial DNA and a 1400 bp nuclear gene (RAG-1) to investigate phylogeographic structuring of species of pebble-mimic dragons. Our topology indicates an early split in this lineage between eastern and western parts of the arid zone that probably dates to the mid-Miocene. This split corresponds directly with large expanses of contemporary sandy habitat in the form of Great Sandy and Great Victoria Deserts. Our data indicate that this biogeographic barrier established very early on in the development of the arid zone and has persisted to present. Additional genetic structuring in the absence of recognized barriers suggests that an expanded view of potential habitat barriers in the arid zone is required.     

Studies of morphological and molecular phylogenetic divergence in spiders (Araneae: Homalonychus) from the American southwest, including divergence along the Baja California Peninsula

Comparative phylogenetic and phylogeographic analyses have revealed a pervasive midpeninsular divergence in the mitochondrial genealogies of numerous vertebrate taxa distributed on the Baja California Peninsula. In this study, we extend the investigation of regional vicariance in Baja California to an arthropod taxon by examining patterns of phylogenetic and morphological divergence in the spider genus Homalonychus (Araneae, Homalonychidae). We analyzed data from two mtDNA genes (16S rRNA and NADH dehydrogenase subunit (1) and a nuclear gene (28S rRNA) using maximum parsimony and Bayesian phylogenetic analyses, and also conducted geometric morphometric analyses employing landmark data on male and female genitalia. Genes and morphology both reveal a deep split across the Colorado River and Gulf of California, separating Homalonychus selenopoides on the east side of river from its congener Homalonychus theologus on the west side of the river, including the Baja California Peninsula. Along the north-south axis of the Baja Peninsula, an apparently more recent midpeninsular phylogenetic break is evident within H. theologus in the mitochondrial genome and in female genitalia. However, there is no measurable divergence between northern and southern populations in either nuclear DNA or male genitalia. We suggest that this discordance between datasets reflects either a difference in rates of evolution between male versus female systems, or that male-based nuclear gene flow is obscuring a phylogenetic split that is fixed in the female-based systems. Our findings provide additional support for a midpeninsular Baja divergence event, although the timing and geological evidence for such an event remain elusive.     

Structural elucidation of an exopolysaccharide from mycelial fermentation of a Tolypocladium sp. fungus isolated from wild Cordyceps sinensis

A novel polysaccharide designated EPS-1A with an average molecular weight around 40 kDa was fractionated and purified by anion-exchange and gel-filtration chromatography from the crude exopolysaccharide (EPS) isolated from fermentation broth of Cs-HK1, a Tolypocladium sp. fungus isolated from wild Cordyceps sinensis. The structural characteristics of EPS-1A were determined with various methods (e.g. GC, GC–MS, FT-IR, ¹H NMR and ¹³C NMR) and through acid hydrolysis, methylation, periodate-oxidation and Smith degradation. The results suggested that EPS-1A was composed of glucose, mannose and galactose at 15.2:3.6:1.0 M ratio. EPS-1A was a slightly branched polysaccharide and its backbone was composed of (1 → 6)-α-d-glucose residues (77%) and (1 → 6)-α-d-mannose residues (23%). Branching occurred at O-3 position of (1 → 6)-α-d-mannose residues of the backbone with (1 → 6)-α-d-mannose residues and (1 → 6)-α-d-glucose residues, and terminated with β-d-galactose residues.     

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.