Distribution and species richness of woody dryland legumes in Baja California,Mexico

Abstract. We analysed the biogeographic patterns of woody legumes in the Baja California peninsula, NW Mexico. From the specimen labels of eight herbaria, we digitized 4205 records from 78 species, and projected them onto a grid of 205 cartographic cells (20’ longitude × 15’ latitude). Most species followed distribution patterns that coincide with floristic subdivisions of the peninsula. Endemism is high, reaching 60–70% in the centre of the peninsula, where the driest deserts are found and where significant floristic changes took place during Pleistocene glacial events. The number of cartographic cells (i.e. their geographic ranges) were log‐normally distributed, as has been reported for many other taxa. Floristic richness was found to be clumped around some cells where the observed richness is significantly higher than could be expected from chance variation. We tested the hypothesis that these ‘hotspots’ could be attributable to great collection efforts or to large land surfaces, but we still found 16 cells where richness is significantly high once these two factors are accounted for. Species richness and micro‐endemism increase towards the south, conforming to Rapoport's rule that predicts that species ranges become smaller towards the equator while richness increases. The floristic hotspots for woody legumes in Baja California occur in the Cape Region and along the Sierra de la Giganta in the southern Gulf Coast, where 77% of the total peninsular legume flora can be found. These hotspots are mostly unprotected, and should be considered priority areas for future conservation efforts.     

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.     

Population genetic structure of annual and perennial populations of Zostera marina L. along the Pacific coast of Baja California and the Gulf of California

The Baja California peninsula represents a biogeographical boundary contributing to regional differentiation among populations of marine animals. We investigated the genetic characteristics of perennial and annual populations of the marine angiosperm, Zostera marina, along the Pacific coast of Baja California and in the Gulf of California, respectively. Populations of Z. marina from five coastal lagoons along the Pacific coast and four sites in the Gulf of California were studied using nine microsatellite loci. Analyses of variance revealed significant interregional differentiation, but no subregional differentiation. Significant spatial differentiation, assessed using θ_ST values, was observed among all populations within the two regions. Z. marina populations along the Pacific coast are separated by more than 220 km and had the greatest θ_ST (0.13–0.28) values, suggesting restricted gene flow. In contrast, lower but still significant genetic differentiation was observed among populations within the Gulf of California (θ_ST = 0.04–0.18), even though populations are separated by more than 250 km. This suggests higher levels of gene flow among Gulf of California populations relative to Pacific coast populations. Direction of gene flow was predominantly southward among Pacific coast populations, whereas no dominant polarity in the Gulf of California populations was observed. The test for isolation by distance (IBD) showed a significant correlation between genetic and geographical distances in Gulf of California populations, but not in Pacific coast populations, perhaps because of shifts in currents during El Niño Southern Oscillation (ENSO) events along the Pacific coast.     

Genealogical concordance and the specific status of Peromyscus sejugis

Peromyscus sejugis, a peripheral isolate of Peromyscus maniculatus, is a threatened taxon endemic to 2 small islands in the Sea of Cortés. Although its insularity makes the specific recognition of P. sejugis inherently problematic, resolution of this problem has important conservation implications. To evaluate the specific validity and evolutionary history of P. sejugis, we compared sequence variation (ND3/ND4L/ND4) in mtDNA for both island populations of P. sejugis with that for 8 populations of P. maniculatus from mainland Baja California. Each island population of P. sejugis had a single haplotype (0.7% sequence divergence), whereas 11 different haplotypes (mean sequence divergence = 0.68%) were obtained for the populations of P. maniculatus. The mean sequence divergence between the populations of the 2 species was 2.0%. Nested clade analysis supports the conclusion that P. sejugis is an insular isolate of P. maniculatus from mainland Baja California. Although our analysis confirms a low level of mtDNA divergence between P. sejugis and P. maniculatus from Baja California, the genealogical concordance of morphological, chromosomal, microsatellite, and mtDNA haplotype distinctiveness supports the conclusion that the 2 island populations of P. sejugis constitute independent evolutionarily significant units and together represent a phylogenetic species distinct from the P. maniculatus from Baja California.     

Phytosociological,bioclimatic and biogeographical classification of woody climax communities of western North America

Abstract. In this study, combining the Braun-Blanquet phytosociological approach with numerical methods (cluster analysis and principal coordinate ordination), woody climax associations of western North America are described, ranging from NW Baja California (Mexico) to Alaska (USA), and including some interior zones of Idaho and Montana (USA), Alberta, British Columbia and the Yukon Territory (Canada). 95 floristic associations are identified, and each is characterized by floristic combination, biogeographical range, and bioclimatic conditions.     

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.     

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.     

Eco-Evolutionary Processes Generating Diversity Among Bottlenose Dolphin, <Emphasis Type="Italic">Tursiops truncatus</Emphasis>, Populations off Baja California,Mexico

For highly mobile species that nevertheless show fine-scale patterns of population genetic structure, the relevant evolutionary mechanisms determining structure remain poorly understood. The bottlenose dolphin (Tursiops truncatus) is one such species, exhibiting complex patterns of genetic structure associated with local habitat dependence in various geographic regions. Here we studied bottlenose dolphin populations in the Gulf of California and Pacific Ocean off Baja California where habitat is highly structured to test associations between ecology, habitat dependence and genetic differentiation. We investigated population structure at a fine geographic scale using both stable isotope analysis (to assess feeding ecology) and molecular genetic markers (to assess population structure). Our results show that there are at least two factors affecting population structure for both genetics and feeding ecology (as indicated by stable isotope profiles). On the one hand there is a signal for the differentiation of individuals by ecotype, one foraging more offshore than the other. At the same time, there is differentiation between the Gulf of California and the west coast of Baja California, meaning that for example, nearshore ecotypes were both genetically and isotopically differentiated either side of the peninsula. We discuss these data in the context of similar studies showing fine-scale population structure for delphinid species in coastal waters, and consider possible evolutionary mechanisms.     

Bioclimates and vegetation along the Pacific basin of Northwestern Mexico

We describe relationships between the bioclimates and vegetation of the northwest of Mexico, including the Baja California peninsula and the stretch of the Pacific basin between the states of Sonora and Colima, a transition zone from the most arid deserts of North America to tropical rainforests. Links were inferred from temperature and rainfall data obtained from 453 weather stations that were used to construct climatograms, calculate several climate indices and describe main latitudinal and longitudinal trends, and from sampling the vegetation around these stations. Across this extensive area, spanning some 14° of both latitude and longitude as well as three zonobiomes and two zonoecotones, two macrobioclimates were identified, Mediterranean and Tropical, within which we were able to distinguish six bioclimates. For each of these bioclimates, we find different types of zonal and azonal vegetation, whose floristic composition and physiognomy are described here.     

Where to conserve? Plant biodiversity and endemism in mediterranean Mexico

Six regions of northwestern (NW) Baja California (Sierra de Juárez, Sierra de San Pedro Mártir, Punta Banda, Colonet, San Quintín and Valle Tranquilo) were compared for their floristic diversity. Checklists for each region were analyzed by their total, native, and endemic floras to give measures of floristic similarity and overlap, and to identify the strongest gradients affecting species distributions. Each region was floristically distinct, with significant variation in the distribution of state endemic taxa. The six regions are readily differentiated by their geographical position in a Principal Components Analysis. The strongest gradients were (a) the W–E gradient from the coast to the mountains and (b) the latitudinal gradient from N to S. These six adjacent regions are found within a local and global biodiversity hotspot that is subject to intense conservation challenges. Conservation of many areas is essential to adequately preserve the diversity of locally endemic taxa with restricted ranges, yet the coast lacks any protected areas at the state and federal level. Private reserves such as Reserva Natural San Quintín may be critical to the conservation of regionally endemic taxa.     

New records of large branchiopods (Branchiopoda: Anostraca,Notostraca, and Spinicaudata) in Mexico

This paper reports new distribution records of large branchiopods for Mexico following a three year survey of the Baja California peninsula. The occurance of the anostracans Thamnocephalus mexicanus (Linder, 1941) and T. platyurus (Packard, 1877), the notostracan Lepidurus lemmoni (Holmes, 1894), and the spinicaudatans Eulimnadia cylindrova (Belk, 1989) and E. texana (Packard,1871), all represent the first records for the peninsula. An undescribed species of the anostracan genus Streptocephalus is recorded from the state of Baja California (Norte). The occurrence of the notostracan genus Triops and four other anostracan species on the peninsula is also confirmed. The conchostracan Cyclestheria hislopi (Baird, 1859) is recorded from the state of Quintana Roo. The collections of Lepidurus and Cyclestheria are the first records for México. These records increase the number of species of large branchiopods reported fromMéxico to 36: 20 Anostraca, 3 Notostraca, 11 Spinicaudata, and 2Laevicaudata. This revised version was published online in August 2006 with corrections to the Cover Date.     

A GRADIENT IN BENTHIC INTERTIDAL ALGAL ASSEMBLAGES ALONG THE SOUTHERN CALIFORNIA COAST1

Based on an analysis floristic data, a gradient appears to exist in the composition of intertidal algae along the 450 km of southern California coastline immediately south of Point Conception. Reciprocal averaging ordination of the algal flora at 51 sites in this area suggests that the gradient is not strictly latitudinal. Variation from a latitudinal gradient occurs ca. 60-80 km south of Point Conception. An ordination of frequently occurring species indicated that compositional changes are substantial when sites at the extreme north and south are compared, but that the changes are gradual and continuous. Possible explanations for the gradient are discussed, and these include: sea temperature, upwelling, sand movement, human disturbance (i.e., pollution, trampling), and wave action.     

Biological conservation of Isla de Cedros,Baja California,Mexico: assessing multiple threats

This paper gives a general overview of the biological conservation problems of Isla de Cedros, off the central part of the Pacific coast of the Baja California peninsula, Mexico. The island's conservation problems can be summarized as: lack of knowledge, habitat degradation, introduction of exotic species, overhunting, and (presumably) commercial capture of reptiles.     

Zonobiomes, zonoecotones and azonal vegetation along the Pacific coast of North America

In this study of the Pacific coast of North America, from Baja California to Alaska, we evaluated the hypothesis that the floristic composition of azonal vegetation determines areas and distribution limits similar to those of the corresponding zonobiomes (ZB), and does so in response to the same macroclimatic changes occurring on the continental scale. To this end, 686 vascular plants of the different habitats found in 279 sites along this coastal strip were recorded. Using an objective classification system (Average Linkage Clustering) and factorial analysis, floristic data acquired in fieldwork were classified into groups, which were in turn related to regional macroclimates. Our main finding was that the azonal coastal vegetation follows a distribution model that is closely linked to the corresponding macroclimate. The four ZB of the northern Pacific coast show a flora and azonal vegetation characteristic to each zonobiome; the latitudinal limits of the azonal vegetation practically coinciding with those already established for the zonal vegetation. The Boreal and Temperate ZB show high percentages of broadly distributed elements. The floristically richest zonobiome in terms of endemic taxa is the Mediterranean zonobiome, whereas the flora of Baja California is characterized by a high number of taxa related to Neotropical flora, especially to those showing links with South America. Data on the geographical distribution and habitats of the 247 most significant coastal species are also provided.     

INTRASPECIFIC GENETIC DIFFERENTIATION IN CALIFORNIA SEA LIONS (ZALOPHUS CALIFORNIANUS) FROM SOUTHERN CALIFORNIA AND THE GULF OF CALIFORNIA

Intraspecific patterns of mitochondrial DNA sequence variation were determined among California sea lions ( Zalophus califomianus californianus ) from three colonies along the Pacific coast of southern and Baja California and one colony in the Gulf of California. We found no variation in 368 base pairs (bp) of cytochrome b sequence among 40 sea lions from these localities, but analysis of 360 base pairs of control region revealed eleven genotypes. The four genotypes found in the Gulf of California population were unique and phylogenetically distinct from those found in sea lions along the Pacific coast. The average sequence divergence between Gulf and Southern California genotypes was 4.3%, suggesting a relatively long period of isolation. However, colonies along the Pacific coast, which are less than 200 km apart, shared mtDNA genotypes, indicating that recent genetic exchange has occurred between them. Therefore, we suggest that regional female philopatry exists in California sea lions. Regional boundaries may be related to oceanic currents or patchiness in the distribution of resources. Further research is needed to better understand the underlying causes of genetic differentiation in the California sea lion.     

Geographic genetic architecture of pocket gopher (Thomomys bottae) populations in Baja California, Mexico

Phylogenetic analyses of complete mitochondrial cytochrome b sequences support the monophyly of pocket gopher (Thomomys bottae) populations from the 1000 km length of the Baja California peninsula of Mexico, relative to other geographical segments of the species range in western North America. The Baja California peninsula is an area that encompasses considerable ecomorphological and infraspecific diversity within this pocket gopher species. However, detailed population analyses encompassing 35 localities distributed over the southern half of the peninsula reveal only trivial phylogeographical structure. Rather, most of the 72 unique 500-base pair haplotypes examined from 142 individuals is restricted to single populations, although a few haplotypes are shared broadly across geography. Individual populations are typically comprised of haplotype sets from different branches in a network of relationships. Analysis of molecular variance (amova) indicates that approximately half of the total pool of variation is contained among individuals within local populations, and that only about 25% can be explained by the regional subdivisions of current subspecies distributions or physiographic realms. A hypothesized historical vicariant event that has been causally linked to the phylogeographical structure of other, codistributed species has had little influence on these pocket gopher populations, explaining only 13% of the total variation. The temporal depth, estimated by coalescence parameters, of the haplotype lineage in Baja California is relatively recent, approximately 300,000 generations; both the mismatch distribution of pairwise comparisons and a significantly positive exponential growth estimate support a recent history of expanding populations; but current, or recent past, migration estimates have remained small, are largely unidirectional from north to south, and weak isolation by distance is present. All data suggest that pocket gophers have relatively recently invaded the southern half of peninsular Baja California, with the genetic signature of expansion still evident but with sufficient time having lapsed to result in a weak isolation by distance pattern. The geographical assemblage of sampled populations thus appears as a meta-population, with limited gene flow contrasting with random haplotype loss due to drift in small, localized populations.     

The peninsula effect on species diversity: a reassessment of the avifauna of Baja California

The peninsula effect, a decrease in species diversity from the base to the tip of peninsulas, has been proposed to explain the relatively poor species diversity of mammals on North American peninsulas. Subsequent work has questioned both the existence of peninsular declines in diversity, as well as the proposed cause (immigration-extinction dynamics). Previous studies of the Baja California avifauna have shown a gradual decrease in the diversity of breeding birds from the base to the tip of the peninsula. Using newly published data on the breeding land birds, I found a decrease only from the base to the middle of the peninsula, with a slight increase in diversity from the middle to the tip. This result is similar to that for other highly vagile taxa (e.g., Chiroptera. Lepidoptera) and is largely due to the coneave diversity gradient of montane species along the peninsula. Habitat associations of the Baja avifauna and the location of potential source populations suggest that: 1) local habitat heterogeneity is likely the single most important factor influencing the avian diversity gradient along the peninsula; and 2) limited immigration of Neotropical species from mainland areas, and of Nearctic species from the base of the peninsula to the montane southern tip is partly responsible for the form of the diversity gradient along the southern half of the peninsula. My results along with those from previous studies, suggest that rather than colonization/extinction dynamics, habitat heterogeneity and the vagility of the taxa considered have the greatest impact on the observed patterns of species diversity along peninsulas,     

Multinational Tagging Efforts Illustrate Regional Scale of Distribution and Threats for East Pacific Green Turtles (Chelonia mydas agassizii)

To further describe movement patterns and distribution of East Pacific green turtles (Chelonia mydas agassizii) and to determine threat levels for this species within the Eastern Pacific. In order to do this we combined published data from existing flipper tagging and early satellite tracking studies with data from an additional 12 satellite tracked green turtles (1996-2006). Three of these were tracked from their foraging grounds in the Gulf of California along the east coast of the Baja California peninsula to their breeding grounds in Michoacán (1337-2928 km). In addition, three post-nesting females were satellite tracked from Colola beach, Michoacán to their foraging grounds in southern Mexico and Central America (941.3-3020 km). A further six turtles were tracked in the Gulf of California within their foraging grounds giving insights into the scale of ranging behaviour. Turtles undertaking long-distance migrations showed a tendency to follow the coastline. Turtles tracked within foraging grounds showed that foraging individuals typically ranged up to 691.6 km (maximum) from release site location. Additionally, we carried out threat analysis (using the cumulative global human impact in the Eastern Pacific) clustering pre-existing satellite tracking studies from Galapagos, Costa Rica, and data obtained from this study; this indicated that turtles foraging and nesting in Central American waters are subject to the highest anthropogenic impact. Considering that turtles from all three rookeries were found to migrate towards Central America, it is highly important to implement conservation plans in Central American coastal areas to ensure the survival of the remaining green turtles in the Eastern Pacific. Finally, by combining satellite tracking data from this and previous studies, and data of tag returns we created the best available distributional patterns for this particular sea turtle species, which emphasized that conservation measures in key areas may have positive consequences on a regional scale.     

Size and age structure of the striped marlin Tetrapturus audax (Pisces: Xiphidae), at Cabo San Lucas, Baja California sur, México

The striped marlin (Tetrapturus audax) is found in temperate and tropical waters of the Pacific and Indian Ocean. It is particularly abundant in the southern tip of the Baja California peninsula. Specimens brought to Cabo San Lucas port (22 degrees 53' N y 109 degrees 54' W) by the sport fishery fleet that operates withing a radius of 54 km from 1988 to 1993 were sexes, measured and weighed. A total 1030 individuals were sampled. Dorsal fin spines (389) were collected for age estimation. The spines were sectioned and hyaline-opaque bands counted. The relation of length to spine radius was similar for males and females. The fourth dorsal spine reflected the growth of the fish. The rhythm of growth mark formation is yearly and related to the sea surface temperature. Ten age groups were found and the seventh group was the most abundant, followed by groups "6" and "8". The age structure of the striped marlin was stable in the study period. The observed sizes were 160-280 cm jaw length with 90% of the organisms measuring between 190 and 225 cm. The small individuals recruit to the fishery from June to December.