Population genetic structure of the round stingray Urobatis halleri (Elasmobranchii: Rajiformes) in southern California and the Gulf of California

The round stingray, Urobatis halleri, is a viviparous elasmobranch that inhabits inshore, benthic habitats ranging from the western U.S.A. to Panama. The population genetic structure of this species was inferred with seven polymorphic microsatellite loci in samples collected at three sites in coastal southern California, one near Santa Catalina Island, California and one in the eastern Gulf of California. Urobatis halleri is relatively common, but little is known of its movement patterns or population structure. Small F_ST values (?0·0017 to 0·0005) suggested little structure among coastal populations of southern and Baja California. The population sampled at Santa Catalina Island, which is separated by a deep‐water channel from the coastal sites, however, was significantly divergent (large F_ST, 0·0251) from the other populations, suggesting low connectivity with coastal populations. The Santa Catalina Island population also had the lowest allele richness and lowest average heterozygosity, suggesting recent population bottlenecks in size.     

Characterization of seven polymorphic microsatellite loci in the Baja California endemic black‐tailed brush lizard Urosaurus nigricaudus

Seven microsatellite loci were developed for the Baja California endemic black‐tailed brush lizard Urosaurus nigricaudus, using an enriched genomic DNA library. All loci were polymorphic and overall presented high levels of variation. Number of alleles ranged from five to 16 (average 12.14), and observed heterozygosities from 0.535 to 0.923 (average 0.752). Cross‐species amplification was successful and polymorphism was detected for all the loci using the congeners Urosaurus lahtelai and Urosaurus ornatus. These markers will be useful to study fragmented populations of U. nigricaudus on agricultural landscape of the Baja California Peninsula.     

Characterization of microsatellite loci in polyploid Lavatera assurgentiflora ssp. assurgentiflora and ssp. glabra (Malvaceae)

Lavatera assurgentiflora (Malvaceae) is one of four species of the genus Lavatera native to California and Baja California. Two geographically defined subspecies are recognized: L. a. assurgentiflora on the northern islands and L. a. glabra on the southern islands. We isolated nine polymorphic microsatellite loci that amplify in both subspecies of L. assurgentiflora. Substantial levels of polymorphism were observed at many of the loci. Four loci exhibited more than 10 alleles, polymorphism information content ranged from 0.4 to 0.8, and up to six alleles were found in some individuals, supporting reports that these taxa are hexaploid. All loci also amplified in Lavatera lindsayi from Guadalupe Island, and we anticipate that they will cross‐amplify in other California Lavatera species as well.     

Seri Indian traditional knowledge and molecular biology agree: no express train for island‐hopping spiny‐tailed iguanas in the Sea of Cortés

Aim The role of human activities in species biogeography can be difficult to identify, but in some cases molecular techniques can be used to test hypotheses of human‐mediated dispersal. A currently accepted hypothesis states that humans mediated the divergence of two species of spiny‐tailed iguanas in the Ctenosaura hemilopha species complex, namely C. conspicuosa and C. nolascensis, which occupy islands in the Sea of Cortés between the peninsula of Baja California and mainland Mexico. We test an alternative hypothesis that follows the traditional knowledge of the Seri Indians and states that the divergence of these species was not mediated by humans. Location Mexico, including Baja California, Sonoran and Sinaloan coastal regions, and Isla San Esteban and Isla San Pedro Nolasco in the Sea of Cortés. Methods We analysed mitochondrial (cytochrome b and cytochrome c oxidase subunit III) DNA sequences from four species in the C. hemilopha species complex. Maximum parsimony and Bayesian inference were used to infer matriarchal genealogical relationships between the species and several outgroup taxa. Bayesian methods were used to estimate divergence times for the major nodes on the trees based on previously published, fossil‐calibrated priors. Results Our analysis indicated that lineages within the C. hemilopha species complex diverged long before human colonization of the Americas. The divergence of C. nolascensis and C. conspicuosa could not be attributed to Seri translocations. The matriarchal genealogy of the species complex currently defies a simple biogeographical interpretation. Main conclusions We conclude that humans did not mediate the divergence of C. nolascensis and C. conspicuosa. This conclusion is consistent with the traditional knowledge of the Seri people. These results demonstrate the utility of molecular techniques in investigating potential cases of human‐mediated dispersal of plants and animals, and reinforce the importance of considering traditional knowledge in the formation of scientific hypotheses and the interpretation of results.     

Revision of Chaetodipus arenarius (Rodentia: Heteromyidae)

Chaetodipus arenarius is an endemic species of the Baja California Peninsula. It has undergone taxonomic changes in the last few years. Today, two species, C. arenarius and Chaetodipus dalquesti, are formally recognized with an unknown range for each. However, genetic analysis showed the possible presence of a third species. The objectives of this study were to use molecular data (cytochrome b and cytochrome c oxidase subunit III genes) to determine how many genetic groups are present in C. arenarius in the Baja California Peninsula; if these groups have morphological characteristics that could be used to distinguish them; if the genetic groups have limited distributions; identify taxonomic implications; and establish if there are any associations amongst genetic groups and different habitats. Bayesian inference, maximum parsimony, maximum likelihood, distance optimality criteria, and minimum spanning network indicated three monophyletic groups with high genetic differences (5.54–13.59%). These groups are morphologically diagnosable based on pelage and morphometric characters and geographical range. The three genetic groups correspond to three taxa previously described as subspecies or species. Based on our results, these should be considered to be different species: C. arenarius occupies the most extreme habitats with hot summers; C. dalquesti occupies the moistest coastal areas in the southern part of the peninsula, and Chaetodipus siccus is restricted by mountain ranges to the isolated Los Planes Basin and Cerralvo Island. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010.     

GEOGRAPHICAL CORRELATION OF MORPHOLOGICAL VARIATION IN SILVETIA COMPRESSA (FUCACEAE,FUCALES, PHAEOPHYCEAE)

Silvetia compressa ( J. Agardh) Serrão et al. is a common member of the upper intertidal fucoid community on the Pacific coast of America from Humboldt County, California, to Punta Baja, Baja California, Mexico. A relatively narrow range of morphological variability is exhibited by most mainland populations, regardless of latitude, but some mainland populations and all insular populations participate in a complex pattern that we have attempted to analyze. A few populations on the Monterey Peninsula in which the fronds are atypically delicate were described by Setchell & Gardner as f. gracilis, to which was assigned a population from Santa Catalina Island. After comparing populations from various parts of the range of the species, including all of the Channel Islands, we conclude that two subspecies may be recognized. In subsp. compressa, which includes f. gracilis as a growth form and occurs chiefly on the mainland, the frond is robust with long tapered receptacles. In the variant subspecies, which is chiefly insular but also occurs on the coast of northern Baja California, the typical frond has slender axes as in f. gracilis, but is more densely branched and has short ellipsoidal receptacles. Comparison of nucleotide sequences from the ITS regions of rDNA revealed an identical pattern for subsp. compressa from Baja California and central California, including populations assignable to f. gracilis. By contrast, the pattern for the variant subspecies differed by 2 bp (0.3%) from that of subsp. compressa.     

Characterization of microsatellite markers for the mangrove tree Avicennia germinans L. (Avicenniaceae)

We describe 10 primers for amplification of microsatellite loci for the mangrove, Avicennia germinans. Eight loci were isolated from a DNA sample from the Pacific coast of Baja California, Mexico and two loci were isolated from a DNA sample from the Atlantic coast of Bermuda. Polymorphism was investigated in a population from the Mexican Pacific coast (n = 15) and in four samples scattered throughout the range of the species. Total number of alleles for the species ranged from two to 10. Observed heterozygosity in the Mexican Pacific coast population ranged from 0.27 to 0.60, with two loci having fixed alleles.     

New polymorphic microsatellite markers for California sea lions (Zalophus californianus)

Nine microsatellite loci were isolated and characterized from California sea lions (Zalophus californianus). In addition, two of five loci tested from harbour seal (Phoca vitulina) produced a single, clear band in Z. californianus, as did one out of five loci from grey seal (Halichoerus grypus) and one out of two loci from elephant seal (Mirounga sp.). No locus tested from South American fur seal (Arctocephalus australis) amplified in Z. californianus. Locus variability was assessed in California sea lions from Los Islotes rookery, Baja California Sur, Mexico. All loci were variable, with allele numbers ranging from three to 12.     

GENETICS OF INCIPIENT SPECIATION IN DROSOPHILA MOJAVENSIS. III. LIFE‐HISTORY DIVERGENCE IN ALLOPATRY AND REPRODUCTIVE ISOLATION

We carried out a three‐tiered genetic analysis of egg‐to‐adult development time and viability in ancestral and derived populations of cactophilic Drosophila mojavensis to test the hypothesis that evolution of these life‐history characters has shaped premating reproductive isolation in this species. First, a common garden experiment with 11 populations from Baja California and mainland Mexico and Arizona reared on two host species revealed significant host plant X region and population interactions for viability and development time, evidence for host plant adaptation. Second, replicated line crosses with flies reared on both hosts revealed autosomal, X chromosome, cytoplasmic, and autosome X cactus influences on development time. Viability differences were influenced by host plants, autosomal dominance, and X chromosomal effects. Many of the F₁, F₂, and backcross generations showed evidence of heterosis for viability. Third, a QTL analysis of male courtship song and epicuticular hydrocarbon variation based on 1688 Baja × mainland F₂ males also revealed eight QTL influencing development time differences. Mainland alleles at six of these loci were associated with longer development times, consistent with population‐level differences. Eight G × E interactions were also detected caused by longer development times of mainland alleles expressed on a mainland host with smaller differences among Baja genotypes reared on the Baja host plant. Four QTL influenced both development time and epicuticular hydrocarbon differences associated with courtship success, and there was a significant QTL‐based correlation between development time and cuticular hydrocarbon variation. Thus, the regional shifts in life histories that evolved once D. mojavensis invaded mainland Mexico from Baja California by shifting host plants were genetically correlated with variation in cuticular hydrocarbon‐based mate preferences.     

Characterization of nine microsatellite loci on endemic kangaroo rats Dipodomys simulans peninsularis from southern Baja California Peninsula

Nine polymorphic microsatellite loci were isolated to investigate the effects of habitat fragmentation on gene flow and genetic variation of the peninsular kangaroo rat Dipodomys simulans peninsularis on agricultural landscape from the Baja California Peninsula. The markers had an average of 9.11 alleles per locus (range 2–14), with mean observed and expected heterozygosities 0.646 (range 0.333–0.900) and 0.788 (range 0.284–0.932), respectively. No evidence of linkage disequilibrium was found across pairs of loci, and only one locus exhibited evidence of null alleles.     

Population structure of nuclear and mitochondrial DNA variation among humpback whales in the North Pacific

The population structure of variation in a nuclear actin intron and the control region of mitochondrial DNA is described for humpback whales from eight regions in the North Pacific Ocean: central California, Baja Peninsula, nearshore Mexico (Bahia Banderas), offshore Mexico (Socorro Island), southeastern Alaska, central Alaska (Prince Williams Sound), Hawaii and Japan (Ogasawara Islands). Primary mtDNA haplotypes and intron alleles were identified using selected restriction fragment length polymorphisms of target sequences amplified by the polymerase chain reaction (PCR–RFLP). There was little evidence of heterogeneity in the frequencies of mtDNA haplotypes or actin intron alleles due to the year or sex composition of the sample. However, frequencies of four mtDNA haplotypes showed marked regional differences in their distributions (Φ_ST = 0.277; P < 0.001; n = 205 individuals) while the two alleles showed significant, but less marked, regional differences (Φ_ST = 0.033; P < 0.013; n = 400 chromosomes). An hierarchical analysis of variance in frequencies of haplotypes and alleles supported the grouping of six regions into a central and eastern stock with further partitioning of variance among regions within stocks for haplotypes but not for alleles. Based on available genetic and demographic evidence, the southeastern Alaska and central California feeding grounds were selected for additional analyses of nuclear differentiation using allelic variation at four microsatellite loci. All four loci showed significant differences in allele frequencies (overall F_ST = 0.043; P < 0.001; average n = 139 chromosomes per locus), indicating at least partial reproductive isolation between the two regions as well as the segregation of mtDNA lineages. Although the two feeding grounds were not panmictic for nuclear or mitochondrial loci, estimates of long-term migration rates suggested that male-mediated gene flow was several-fold greater than female gene flow. These results include and extend the range and sample size of previously published work, providing additional evidence for the significance of genetic management units within oceanic populations of humpback whales.     

PATTERNS OF REPRODUCTION,GENETIC DIVERSITY,AND GENETIC DIFFERENTIATION IN CALIFORNIA POPULATIONS OF THE GENICULATE CORALLINE ALGA LITHOTHRIX ASPERGILLUM (RHODOPHYTA)1

The reproductive composition and genetic diversity of populations of the red seaweed Lithothrix aspergillum Gray (O. Corallinales) were studied at three southern California sites (Shaw's Cove and Treasure Island, Laguna Beach; Indian Rock, Santa Catalina Island) and at a fourth site (Bodega Bay) located in northern California. Sexually reproducing populations were confined to southern California. Diploid individuals were numerically dominant over haploid (gametophytic) individuals at all sites. Intertidal and subtidal subpopulations from Shaw's Cove differed in their reproductive profiles. Most intertidal specimens found on emersed surfaces were densely branched, turf-forming, and bore tetrasporangial (68.6%), carposporangial (11.4%), or spermatangial (5.7%) conceptacles, reflecting a sexual life history; none produced asexual bispores. In contrast, 74.3% of the larger, loosely branched subtidal specimens bore bisporangial conceptacles indicative of asexual reproduction. Nearly 70% of the Indian Rock thalli showed no evidence of conceptacle formation. Only asexual, diploid bispore-producing thalli were obtained from the Bodega Bay site. Genetic diversity (mean number of alleles per locus, percent of polymorphic loci, and average expected heterozygosity) of diploid L. aspergillum populations varied with life-history characteristics and geographic location. A total of 30 alleles was inferred from zymograms of 16 loci examined by starch-gel electrophoresis; of these loci, 11 were polymorphic. The genetic diversity of sexual, diploid populations of L. aspergillum (alleles per locus [A/L] = 1.4-1.5; percent polymorphic loci [%P] = 37.5-50.0) was relatively high compared with other red seaweeds. Lowest diversity (A/L = 1.0; %P = 0.0) occurred in the exclusively asexual Bodega Bay population which consisted of genetic clones. All sexual L. aspergillum populations deviated significantly from Hardy-Wein-berg expectations due to lower than expected heterozygosity. Genetic differentiation (Wright's F_statistic [F_ST]; Nei's Genetic Distance [D]) among sexually reproducing southern California populations was low (F_ST= 0.030) on a local scale (ca. 5 km), suggesting high levels of gene flow, but high genetic differention (F_ST= 0.390 and 0.406) occurred among southern California populations separated by ca. 70 km. Very high genetic differentiation (F_ST= 0.583–0.683) was obtained between northern and southern California populations separated by 700–760 km. Our genetic and reproductive data suggest that the L. aspergillum population from Bodega Bay is sustained by perennation, vegetative propagation, or asexual reproduction by bispores and may represent an isolated remnant or a population established by a founder event.     

Molecular genetic analysis of two native desert palm genera,Washingtonia and Brahea,from the Baja California Peninsula and Guadalupe Island

The complex geological and ecological processes that have generated high levels of biodiversity and endemism in the Baja California Peninsula have been the subject of intensive study. However, relatively little is known about phylogeography of the iconic endemic palm species of this region. We therefore analyzed a total of 2,294 bp of chloroplast and 738 bp of nuclear sequence data in 169 samples of five native palm species from Baja California, Sonora and Guadalupe Island. We found that Washingtonia and Brahea palms had low levels of genetic diversity and were highly structured, with the majority of species and major geographic regions being characterized by distinct haplotypes. We also found strong support for currently recognized species in Washingtonia, but our results were less clear cut for Brahea due to haplotype sharing. Furthermore, patterns of population structure were broadly consistent with historical vicariant events such as the inundation of the Isthmus of La Paz, the formation of the Sea of Cortez, and the more recent colonization and isolation of Guadalupe Island's palms. Our findings contribute toward a growing appreciation of the complexity of plant responses to past geological changes and also provide valuable baseline genetic data on relict American palm species.     

Factors Influencing the Selection of Basking Perches on Cardon Cacti by Spiny-Tailed Iguanas (Ctenosaura hemilopha)1

The selection of optimal basking perches to achieve preferred body temperatures and avoid predation risks is one of the most important behavioral decisions that a lizard makes. We studied perch selection by the spiny-tailed Iguana (Ctenosaura hemilopha) in Baja California Sur where they bask on elevated cacti. The more important factor determining cardon selection by iguanas was the presence of old woodpecker holes which could be used as refuges within the top 50 cm of a branch. Other factors are the height of the cardon, the presence of some other holes and the number of branches of the plant. These data suggest that predation risk is the strongest factor affecting the selection of certain cardons as basking structures by spiny-tailed iguanas. Because the holes are mainly old woodpecker nests, the choice of a cardon as a refuge-perching site has to be related to a prior selection by woodpeckers.     

Ten polymorphic microsatellite loci for the endangered Buena Vista Lake shrew (Sorex ornatus relictus)

The ornate shrew (Sorex ornatus) is restricted to the vanishing wetlands of California, USA and Baja California, Mexico. Several subspecies of ornate shrews are considered ‘mammal species of special concern’ in California by the Department of Fish and Game, and one (Sorex ornatus relictus) has recently been listed as endangered. Populations of shrews around Buena Vista Lake have been diminished or extirpated due to habitat deterioration and human development. In order to study the patterns of genetic variation in isolated populations of Buena Vista Lake shrews, we developed 10 polymorphic microsatellite loci. There were 6–27 alleles per locus, and the loci had heterozygosity values that ranged from 20 to 80%. In addition, we screened 20 different populations of S. ornatus, eight species within two subfamilies of shrews (Soricinae and Crocidurinae), as well as in a mole (Talpidae, Neurotrichus gibbsii), to determine if these loci could be informative in other species as well.     

Genotyping by sequencing reveals contrasting patterns of population structure,ecologically mediated divergence,and long‐distance dispersal in North American palms

Comparative studies can provide powerful insights into processes that affect population divergence and thereby help to elucidate the mechanisms by which contemporary populations may respond to environmental change. Furthermore, approaches such as genotyping by sequencing (GBS) provide unprecedented power for resolving genetic differences among species and populations. We therefore used GBS to provide a genomewide perspective on the comparative population structure of two palm genera, Washingtonia and Brahea, on the Baja California peninsula, a region of high landscape and ecological complexity. First, we used phylogenetic analysis to address taxonomic uncertainties among five currently recognized species. We resolved three main clades, the first corresponding to W. robusta and W. filifera, the second to B. brandegeei and B. armata, and the third to B. edulis from Guadalupe Island. Focusing on the first two clades, we then delved deeper by investigating the underlying population structure. Striking differences were found, with GBS uncovering four distinct Washingtonia populations and identifying a suite of loci associated with temperature, consistent with ecologically mediated divergence. By contrast, individual mountain ranges could be resolved in Brahea and few loci were associated with environmental variables, implying a more prominent role of neutral divergence. Finally, evidence was found for long‐distance dispersal events in Washingtonia but not Brahea, in line with knowledge of the dispersal mechanisms of these palms including the possibility of human‐mediated dispersal. Overall, our study demonstrates the power of GBS together with a comparative approach to elucidate markedly different patterns of genomewide divergence mediated by multiple effectors.     

Eight polymorphic microsatellite markers for kelp bass,Paralabax clathratus,amplified in three multiplex polymerase chain reaction sets

Kelp bass, Paralabrax clathratus, is an important sport fishery of California, USA and Baja, Mexico. Here we describe eight microsatellite loci developed for this species. Two loci were derived from known primers for other species in the family, Serranidae and six were developed anew using a clone enrichment protocol. Loci were arranged into three multiplex PCR (polymerase chain reaction) sets for fast throughput; 564 individuals from nine populations across the species’ range were genotyped. Polymorphism ranged from four to 47 alleles and all populations at all loci displayed Hardy–Weinberg and linkage equilibrium.     

Population structure of sablefish Anoplopoma fimbria using genetic variability and geometric morphometric analysis

Population structure of the sablefish (Anoplopoma fimbria) in the northeastern Pacific Ocean was determined using three approaches: geometric morphometrics (14 landmarks), mitochondrial DNA (fragment of COI gene), and nuclear DNA (four microsatellite loci). Samples came from the Bering Sea, Gulf of Alaska, offshore Oregon, and offshore the mid‐Baja California Peninsula (at San Quintin). Differences in body shape were grouped in the samples from the north (Bering Sea and Gulf of Alaska). A slight but significant population structure was also observed in allele frequencies of microsatellites, F_ST values, amova , and Bayesian individual assignment tests; however, analyses of population structure using mtDNA did not reveal any population differentiation. Differences in population structure detected by distinct approaches, in addition to the moderately high haplotype diversity and low nucleotide diversity of the COI fragment, suggest recent and developing population differentiation in the sablefish.     

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.     

Polymorphic microsatellite loci in Lithophragma maximum (Saxifragaceae), an endemic plant of San Clemente Island

In this paper, we report the isolation of 11 polymorphic microsatellite loci from Lithophragma maximum (Saxifragaceae), a rare species restricted to San Clemente Island, California. We found moderate levels of allelic variation (mean N_A= 5.5) and high levels of expected heterozygosity (mean H_E= 0.531) across sampled individuals. However, eight loci showed a significant absence of heterozygous individuals, which we attribute to small population size and demographic history of the species.