Genetic variation in Costa Rican populations of the tropical timber species Cedrela odorata L., assessed using RAPDs

Cedrela odorata L. (Spanish cedar), an economically important timber species native to the American tropics, is the focus of increasing conservation concern due to high rates of deforestation within its native range. To assess the extent of the genetic diversity within and between populations of this species, samples were obtained from 10 widely dispersed populations within Costa Rica and analysed for random amplified polymorphic DNA (RAPD) variation. Fourteen 10-mer primers were used to generate 97 polymorphic RAPD bands. Presence/absence data for all bands were subjected to a pairwise genetic distance analysis, according to Jaccard's coefficient, then neighbour-joining cluster analysis was performed on these distances, as was an analysis of molecular variance ( AMOVA ), to assess levels of differentiation between populations and regions, and Shannon's Diversity Index was used to quantify levels of diversity within and between populations. Results indicated highly significant genetic differentiation ( P < 0.004, AMOVA ) between populations originating from the North Pacific and Atlantic/South Pacific regions of Costa Rica, with 35.3% of the total variation attributable to a difference between these areas. Little differentiation was recorded between populations from within the same region ( P = 0.757, AMOVA ), and 65.1% of the total variance was attributable to variation within populations. Estimated values for within-population diversity, calculated as H _pop/ H _sp by means of Shannon's Diversity Index, were found to vary greatly between primers, but the overall within-population component of genetic diversity was 0.45. Possible reasons for the high degree of intraspecific genetic variation within this species are discussed and the implications of these results for the conservation and use of its genetic resources are described.     

Chemical variation in defensive secretions of four species of Nasutitermes

Nineteen monoterpenes and 13 diterpenes were distinguished in soldier defensive secretions of four species of Nasutitermes. Samples of N. columbicus, N. corniger, N. ephratae and N. nigriceps were collected in Costa Rica and Panama. Interspecifically, these species can be differentiated by monoterpenes, which exhibit a complex pattern. Intraspecifically, there is detectable variation in monoterpenes and diterpenes, each of which allows discrimination of four populations of N. corniger. Similarity between two Pacific populations of N. corniger from different environments, a tropical dry forest and a rainforest, suggests there is no climatic influence on chemical compositions of defensive secretions. In addition, results from Costa Rica provide evidence that the Atlantic population of N. corniger is derived from Pacific populations.     

Reproductive plasticity in <Emphasis Type="Italic">Petrolisthes armatus</Emphasis> (Anomura,Porcellanidae): a comparison between a Pacific and an Atlantic population

The porcellanid crab Petrolisthes armatus, with a known geographic distribution covering a wide range of latitudes, was selected to conduct a comparative study on egg production and reproductive output between two populations from Pacific and Atlantic coasts. Ovigerous females were collected between September and November 2005 in Punta Morales, Pacific coast of Costa Rica (n = 137) and from March 2005 to July 2006 in Araçá region, São Paulo, Brazil (n = 46). The mean size of females was statistically different between both populations with larger females from Brazil than Costa Rica (7.9 ± 1.31 and 6.6 ± 0.95 mm CW, respectively). Likewise, egg production was significantly different in both populations and started in Pacific Costa Rica at a smaller female size than in Brazil. Specimens from Pacific Costa Rica produced roughly three times more eggs than equally sized females from Brazil, while newly extruded embryos were larger in Brazil than in Costa Rica (0.045 and 0.039 mm³, respectively). Egg volume increased during embryogenesis by 112.8 and 164.5% in Costa Rican and Brazilian populations, respectively. The egg water content increased steadily in both populations; however, eggs produced in Brazil contained always more water than those from the same developmental stage in Costa Rica. Average reproductive output (based on dry weight) was substantially lower in Brazil (0.031) than in Costa Rica (0.065). Our results confirm an impressive intraspecific plasticity of reproductive features in an intertidal decapod. The observed phenotypical variability might be related to local environmental conditions as well as to the location of the studied population concerning its geographic range of distribution.     

Genetic variation of introduced Hawaiian and native Costa Rican populations of an invasive tropical shrub, Clidemia hirta (Melastomataceae)

Clidemia hirta is one of the most common woody invasive plants in mesic to wet forests in Hawaii, where it was introduced around 1940. The species is relatively uncommon by comparison in its native range of Central and South America and some Caribbean Islands. We examined genetic variation in allozymes of 20 C. hirta populations on four Hawaiian Islands to determine the introduction history. For comparison, we measured genetic variation in 20 native populations across Costa Rica. Mean levels of genetic variation in Hawaiian and Costa Rican populations were low compared to other woody or introduced plants (11.5-12.5% polymorphic loci, 2.05-2.50 alleles per polymorphic locus, and 0.045-0.063 expected heterozygosity). Most genetic diversity was held within rather than among populations in both areas (G(ST) = 0.120 and 0.271 in Hawaii and Costa Rica, respectively). Hawaiian populations had a high degree of genetic similarity, and no genetic differentiation was found among the four Hawaiian Islands sampled. These patterns of genetic variation in Hawaii suggest that no intraspecific hybridization of genotypes from different parts of the native range has occurred and that introductions to the different islands came from the same or similar source populations. The low levels of genetic diversity in parts of both the native and introduced ranges suggest that genetic variation is unrelated to invasiveness in C. hirta.     

Where are the survivors? Tracking relictual populations of endangered frogs in Costa Rica

Aim We used abiotic environmental variables and historical locality records to infer distributions of endangered anuran species of Costa Rica to promote efficient strategies for future amphibian surveys. Location Costa Rica. Methods We used a Maximum Entropy Algorithm (Maxent) to predict potential distribution maps for 17 species of endangered anurans and create a consensus map of species richness. We compared the environmental conditions from localities where relictual amphibian populations were recently rediscovered with the conditions across their historical range to evaluate the possibility that these relictual populations might occur in specific climatic conditions that could explain their persistence. We used a multicriteria analysis considering the following factors: the intersection zones between the consensus map, conservation areas, potential Batrachochytrium dendrobatidis (Bd) distribution, collecting effort and areas within the precipitation range at which reappearances had occurred to locate sites for future surveys. Results The resulting predictions suggest that suitable areas for the highest number of species occur between 1300 and 2500 m.a.s.l and are concentrated along the Pacific slopes of the Cordillera de Talamanca and Cordillera Volcánica Central. Around 45% of the high potential richness area is under protection. Relictual populations of declined species seem to persist mainly in highly humid localities (2500–3500 mm of mean annual precipitation). Around 240 km² has an ideal environment for the rediscovery of relictual populations. The multicriteria analysis showed that around 0.5% of the Costa Rican territory should be surveyed exhaustively for frogs. Main conclusions Many of the potential refugia we identified here have not been surveyed since 2000, the areas identified by the best model predictions correspond well with the localities of the relictual populations recently reported. We suggest future surveys of missing amphibian species should focus on these areas. The discovery of populations of endangered species can be used to propose conservation areas.     

Genetic patterns and conservation of the Scarlet Macaw (<Emphasis Type="Italic">Ara macao</Emphasis>) in Costa Rica

Once widely distributed throughout the lowland forests of Costa Rica, scarlet macaws (Ara macao) have been reduced to two major, geographically separated, populations along the Pacific slope. Past demographic declines raise conservation concerns regarding the detrimental effects of population fragmentation. This investigation aimed to evaluate the current status of scarlet macaws along the Pacific slope by examining levels of genetic variation and patterns of genetic structure within and among remnant populations. Statistical analyses using multilocus genotypes revealed strong differentiation between Central and South Pacific populations, suggesting local geographic barriers have historically restricted gene flow between these localities. High genetic diversity suggests neither population suffers from genetic erosion, likely resulting from relatively large population sizes and high dispersal capacity and longevity. However, evidence of disequilibrium within the Central Pacific population infers anthropogenic threats have disrupted natural population dynamics. These results advocate on focusing available resources on habitat restoration and nest protection, as a means to assist in reestablishing demographic stability and maintain the genetic health of wild scarlet macaws in Costa Rica.     

Marine biodiversity of Costa Rica: Crustacea: Infraorder Anomura

The anomuran crabs are among the best known crustacean groups from the Pacific coast. However, this group is poorly known from the Caribbean coast of Costa Rica. In this compilation based on the literature and the collection at the Zoology Museum, Biology School, University of Costa Rica, we report the presence of 114 species of the Infraorder Anomura for Costa Rica, 20 species from the Caribbean, 96 species from the Pacific (two are present on both coasts). Twenty-nine species are new reports for Costa Rica, 15 from the Caribbean coast (74% of the total of species from that coast) and 14 from the Pacific (15% of the total from the Pacific). The range often species is extended to Costa Rica, siete from the Caribbean and three from the Pacific. Six species are reported for the first time from Cocos Island, where there are also four endemic species.     

Marine biodiversity of Costa Rica: class Polychaeta (Annelida)

Three hundred and thirty two species of polychaetes belonging to 50 families are reported for Costa Rica based on previously published reports. In a few families, available material was re-examined and appropriate changes in species identifications were made. Only three species, (Glycera oxycephala Ehlers, 1887; Hemipodia pustulata (Friedrich 1956); and Scolepis (Scolepis) squamata (Miller, 1806)) have been reported from the Caribbean coast of Costa Rica, and these three species have also been reported from the Pacific Coast. The families with more than 15 species are the Spionidae (26), Cirratulidae (24), Nereididae (21), Lumbrineridae (19) and Paraonidae (16). Despite extensive sampling on the Pacific coast over the last 25 years, the Costa Rican polychaete fauna, especially that of the Caribbean coast, remains poorly known.     

Diversity of the free-living marine and freshwater Copepoda (Crustacea) in Costa Rica: a review

The studies on marine copepods of Costa Rica started in the 1990’s and focused on the largest coastal-estuarine systems in the country, particularly along the Pacific coast. Diversity is widely variable among these systems: 40 species have been recorded in the Culebra Bay influenced by upwelling, northern Pacific coast, only 12 in the Gulf of Nicoya estuarine system, and 38 in Golfo Dulce, an anoxic basin in the southern Pacific coast of the country. Freshwater environments of Costa Rica are known to harbor a moderate diversity of continental copepods (25 species), which includes 6 calanoids, 17 cyclopoids and only two harpacticoids. Of the +100 freshwater species recorded in Central America, six are known only from Costa Rica, and one appears to be endemic to this country. The freshwater copepod fauna of Costa Rica is clearly the best known in Central America. Overall, six of the 10 orders of Copepoda are reported from Costa Rica. A previous summary by 2001 of the free-living copepod diversity in the country included 80 marine species (67 pelagic, 13 benthic). By 2009, the number of marine species increased to 209: 164 from the Pacific (49% of the copepod fauna from the Eastern Tropical Pacific) and 45 from the Caribbean coast (8% of species known from the Caribbean Basin). Both the Caribbean and Pacific species lists are growing. Additional collections of copepods at Cocos Island, an oceanic island 530 km away of the Pacific coast, have revealed many new records, including five new marine species from Costa Rica. Currently, the known diversity of marine copepods of Costa Rica is still in development and represents up to 52.6% of the total marine microcrustaceans recorded in the country. Future sampling and taxonomic efforts in the marine habitats should emphasize oceanic environments including deep waters but also littoral communities. Several Costa Rican records of freshwater copepods are likely to represent undescribed species. Also, the biogeographic relevance of the inland copepod fauna of Costa Rica requires more detailed surveys.     

Phytogeography of the trees of the Osa Peninsula,Costa Rica

A phytogeographic analysis of the distributions of 454 species of trees native to the Osa Peninsula in 22 families revealed that 4.8% of the species are endemic to the Osa Peninsula and the adjacent mainland of Costa Rica. However, nearly one-fourth of the species might be regionally endemic to Central-South Mesoamerica (Costa Rica, Nicaragua, and Panama). Our sample suggests that 53.6% of the species occur in some part of Mesoamerica and sometimes range into northwestern South America, and that 44.5% of the species have wide distributions throughout tropical America. There is a strong affinity with the flora of northwestern South America, with 46.2% of the species on the Osa also found there. In addition, 50.6% of the tree species on the Osa occur on both the Atlantic and Pacific slopes of Central America or, if they reach South America, are sometimes found on both sides of the Andes. Major contributions to the tree flora of the Osa have been made by species arriving in the Osa by 1) dispersal from South and North America to islands in proto Central America before the formation of a dry-land connection between the two continents, and 2) migration from South America and North America after the closure of the Panamanian isthmus was made. This analysis demonstrates the importance of the Osa as a regional refuge for protecting species with distributions limited to the Osa and parts of Panama, Costa Rica, or Nicaragua. The Osa is also important because it harbors the last expanse of tropical wet forest on the Pacific slope of Central America large enough to ensure the survival of the Central American populations of widely distributed plants and animals.     

Analysis of genetic diversity and structure in Ethiopian populations of Phytolacca dodecandra using RAPD

The genetic diversity and structure in 17 wild populations (249 individuals) of Phytolacca dodecandra (endod) sampled along altitudinal gradients of 1600-3000 meters above sea level (m.a.s.l.) in Ethiopia was studied using random amplified polymorphic DNA (RAPD). A total of 70 polymorphic loci (P) scored from 12 RAPD primers were used to calculate different diversity indices within and between populations, habitats, geographical regions, climatic zones and altitude groups. The number of polymorphic loci and overall Shannon information measure (H) in the populations varied from 30 to 55 and from 0.228 to 0.418, respectively. In general, differences in population variability were found significantly correlated to effective population size. Both P and H were significantly higher in an undisturbed than in a disturbed habitat, and in the lowland and central-highland than in the highland altitude group. However, for both parameters the differences were not statistically significant between regions and climatic zones. Genetic distance between populations varied from 0.301 to 0.628. Cluster analysis performed using the genetic distance matrix revealed a clear separation of the highland populations (2501-3000 m.a.s.l.) from those of the lowland/central-highlands (1600-2500 m.a.s.l.) irrespective of their geographical regions and climatic zones. Analysis of molecular variance (AMOVA) indicated that differences in habitat, geographical regions and climatic zones explained 4.6%, 2.5% and 4.6%, respectively. But none of these differences were significant. Altitude explained 17.2% of the total variance and was highly significant. The data, therefore, clearly indicated the association of genetic structure in endod with altitude. The proportion of RAPD variation found among populations (21.2-35.0%) was somewhat intermediate between values reported for selfing and outcrossing species. The fixation index (FST) values (0.350 to 0.384) indicated very high genetic differentiation among populations.     

Population Genetic Structure of Neotropical Mangrove Species on the Colombian Pacific Coast: Pelliciera rhizophorae (Pellicieraceae)1

Pelliciera rhizophorae is a Neotropical mangrove species whose distribution is mostly restricted to the Pacific Coast, between the Gulf of Nicoya (Costa Rica) and the Esmeraldas River (Ecuador). In the Caribbean, patches of Pelliciera have been found in Nicaragua, Panama, and Colombia. The genetic variation and the population structure of P. rhizophorae were evaluated in six zones of the Colombian Pacific, using AFLP molecular markers. Of the 225 amplified fragments produced, 155 (69%) were polymorphic in 57 individuals of P. rhizophorae collected in the localities of Virudó, Charambirá, La Plata Island, Tumaco, Milagros, and Chontal. Genetic diversity within populations varied significantly, with the lowest levels of within‐population variation (H_ep= 0.081 and 0.090, respectively) in La Plata Island and Tumaco; and the highest level of variation (H_ep= 0.187) in Chontal. In the Colombian Pacific, P. rhizophorae was significantly structured, with 26.5 percent of the variation found among populations, which was an evidence of substructure within populations. Genetic differentiation was not related to the geographic distance between zones. This suggests that the population dynamics of P. rhizophorae could be associated with historic processes influenced by ecological and environmental factors such as the movement of pollen by birds, the displacement of propagules by marine currents, and the spatial distribution of favorable habitats.     

Sexual signal evolution and patterns of assortative mating across an intraspecific contact zone

Contact zones provide important insights into the evolutionary processes that underlie lineage divergence and speciation. Here, we use a contact zone to ascertain speciation potential in the red-eyed treefrog (Agalychnis callidryas), a brightly coloured and polymorphic frog that exhibits unusually high levels of intraspecific variation. Populations of A. callidryas differ in a number of traits, several of which are known sexual signals that mediate premating reproductive isolation in allopatric populations. Along the Caribbean coast of Costa Rica, a ~100 km contact zone, situated between two phenotypically and genetically divergent parent populations, contains multiple colour pattern phenotypes and late-generation hybrids. This contact zone provides the opportunity to examine processes that are important in the earliest stages of lineage divergence. We performed analyses of colour pattern variation in five contact zone sites and six parental sites and found complex, continuous colour variation along the contact zone. We found discordance between the geographic distribution of colour pattern and previously described genomic population structure. We then used a parental site and contact zone site to measure assortative mating and directional selection from naturally-occurring amplectant mating pairs. We found assortative mating in a parental population, but no assortative mating in the contact zone. Furthermore, we uncovered evidence of directional preference towards the adjacent parental phenotype in the contact zone population, but no directional preference in the parent population. Combined, these data provide insights into potential dynamics at the contact zone borders and indicate that incipient speciation between parent populations will be slowed.     

Recruitment limitation in a tropical intertidal barnacle: Tetraclita panamensis (Pilsbry) on the Pacific coast of Costa Rica

On the central Pacific coast of Costa Rica, populations of Tetraclita panamensis (Pilsbry) were studied to determine the relative importance of pre- and postrecruitment processes in determining patterns of distribution and abundance. Populations at 12 sites on two shores were censused photographically from 1983 to 1985 at 1–3-months intervals, yielding data on recruitment, growth, and mortality. During the 2.5-yr study, recruitment replaced < 24% of the number dying at 10 of the 12 sites. Most populations were sparse and declining. Preemption of settlement was unlikely because bare space was always abundant and recruitment occurred mostly on the shells of adults. Individuals reached a maximum size in 1–3 yr, after which there was little growth. Estimated longevity ranged from 3.6 to 14.7 yr among sites. Mortality was unrelated to submergence, density or the abundance of an important predator, the gastropod Acanthina brevidentata (Wood). Patterns of distribution and abundance among the 12 sites and along the central coast of Costa Rica are probably a function of past variation in recruitment. Many apparently suitable habitats are unoccupied.     

Persistent leatherback turtle migrations present opportunities for conservation

Effective transboundary conservation of highly migratory marine animals requires international management cooperation as well as clear scientific information about habitat use by these species. Populations of leatherback turtles (Dermochelys coriacea) in the eastern Pacific have declined by >90% during the past two decades, primarily due to unsustainable egg harvest and fisheries bycatch mortality. While research and conservation efforts on nesting beaches are ongoing, relatively little is known about this population of leatherbacks' oceanic habitat use and migration pathways. We present the largest multi-year (2004-2005, 2005-2006, and 2007) satellite tracking dataset (12,095 cumulative satellite tracking days) collected for leatherback turtles. Forty-six females were electronically tagged during three field seasons at Playa Grande, Costa Rica, the largest extant nesting colony in the eastern Pacific. After completing nesting, the turtles headed southward, traversing the dynamic equatorial currents with rapid, directed movements. In contrast to the highly varied dispersal patterns seen in many other sea turtle populations, leatherbacks from Playa Grande traveled within a persistent migration corridor from Costa Rica, past the equator, and into the South Pacific Gyre, a vast, low-energy, low-productivity region. We describe the predictable effects of ocean currents on a leatherback migration corridor and characterize long-distance movements by the turtles in the eastern South Pacific. These data from high seas habitats will also elucidate potential areas for mitigating fisheries bycatch interactions. These findings directly inform existing multinational conservation frameworks and provide immediate regions in the migration corridor where conservation can be implemented. We identify high seas locations for focusing future conservation efforts within the leatherback dispersal zone in the South Pacific Gyre.     

Marine biodiversity of Costa Rica: the phyla Sipuncula and Echiura

Fourteen species of Sipuncula belonging to 9 genera have been reported from Costa Rican waters, mostly from the Pacific coast. Three of these species are new records for Costa Rica (Phascolion strombus (Montagu 1804), Aspidosiphon (Aspidosiphon) muelleri Diesing 1851, and Aspidosiphon (Aspidosiphon) gracilis schnehageni (W. Fisher 1946)). One species of Echiura, Thalassema steinbecki Fisher 1946, in the order Echiuroinea, has been reported from the Pacific coast of Costa Rica.     

Effectiveness of Protected Areas for Representing Species and Populations of Terrestrial Mammals in Costa Rica

Costa Rica has one of the greatest percentages (26%) of protected land in the world. The National Protected Areas System (NPAS) of Costa Rica was established in 1976 and currently includes >190 protected areas within seven different protection categories. The effectiveness of the NPAS to represent species, populations, and areas with high species richness has not been properly evaluated. Such evaluations are fundamental to understand what is necessary to strengthen the NPAS and better protect biodiversity. We present a novel assessment of NPAS effectiveness in protecting mammal species. We compiled the geographical ranges of all terrestrial Costa Rican mammals then determined species lists for all protected areas and the estimated proportion of each species’ geographic range protected. We also classified mammal species according to their conservation status using the IUCN Red List of Threatened Species. We found almost complete representation of mammal species (98.5%) in protected areas, but low relative coverage (28.3% on average) of their geographic ranges in Costa Rica and 25% of the species were classified as underprotected according to a priori representation targets. Interestingly, many species-rich areas are not protected, and at least 43% of cells covering the entire country are not included in protected areas. Though protected areas in Costa Rica represent species richness well, strategic planning for future protected areas to improve species complementarity and range protection is necessary. Our results can help to define sites where new protected areas can have a greater impact on mammal conservation, both in terms of species richness and range protection.     

Tectonic survey of the Central Pacific coast of Costa Rica

Three fossil coral samples (Pocillopora elegans and Pocillopora damicornis) were collected from a beachrock in Puerto Escondido, central Pacific coast of Costa Rica. The age of the samples ranged from 1272+/-60 to 2010+/-60 years BP. They represent some of the few Holocene fossil corals known from the Pacific coast of the Americas. The uplift rate for the central Pacific coast of Costa Rica was calculated in 0.4 to 2.1 mm year(-1).     

Distribution of plant species in Israel in relation to spatial variation in rainfall

Abstract. Relations between the distribution of herbaceous and woody species from the flora of Israel and the variation (25–900 mm) in mean annual rainfall are described using a grid-based approach integrating multivariate techniques (cluster analysis and DCA-ordination) and GIS. The analysis was based on an extensive data base of vascular plant records in 10 km × 10 km grid cells representing the climatic zones of Israel. Cluster analysis revealed three geographically distinct clusters of grid cells which could be arranged along the main rainfall gradient in the study area. The main subdivision is between cells from the Mediterranean region and cells from the more arid region. The second subdivision separated cells from the arid region into two distinct groups which differed significantly in rainfall. Clustering of herbaceous and woody species was similar, but boundaries separating adjacent clusters based on woody species were consistently linked to more rainy areas. Cell scores on the first DCA-axis were significantly correlated with mean annual rainfall, but this relationship was not linear. Log-transformation of the rainfall data resulted in a higher correlation between the DCA-scores and rainfall, indicating that variation in rainfall had a stronger effect on species composition in relatively dry than in more mesic regions. Slopes of linear regression models relating cell scores on the first DCA-axis to mean annual rainfall, as well as the corresponding R²-values, were highest in the desert, intermediate in the dry Mediterranean, and lowest in the mesic Mediterranean territory. These results indicate that both the per-unit effect of rainfall on plant species composition and its relative importance as a determinant of compositional variation decreased from relatively dry to more mesic regions. The length of the first axis, as well as the slopes of regression models relating cell scores on the first DCA-axis to mean annual rainfall were consistently higher in the case of woody species which are characterized by a higher rate of compositional variation. Due to lack of comparable data, it is difficult to evaluate whether our results can be generalized to other regions. However, an analysis of data obtained from different parts of the world points to a considerable variability in the degree to which rainfall affects patterns of floristic variation. Our findings that compositional responses to rainfall may vary between different groups of species and may depend on the part of the rainfall range which is being considered may partially explain this inconsistency.