Coconut varieties in Mexico

Cocos nucifera Linn, was introduced to Mexico by Spanish navigators in the middle of the 16th century, from Cape Verde and Santo Domingo to the Atlantic coast, and from the Solomon Islands and the Philippines to the Pacific coast. These populations remained isolated from one another for almost 500 years. Morphological studies on fruit characteristics show that coconuts from the two populations are contrastingforms. The Atlantic coast populations show a wild type fruit, with angular shape, thick husk, thick shell, thick endosperm and low water content. The Pacific populations have a domesticated type fruit, with fruit and nut more spherical, with a much thinner husk and shell, and higher water content. The higher variability observed in the Pacific coast raises the possibility of later introductions, also from the Philippines or other Pacific islands.     

The presence of coconut in southern Panama in pre-Columbian times: clearing up the confusion

Background

The pre-Columbian presence of coconut on the Pacific coast of Panama is attested by a number of independent written accounts. However, recent papers question their accuracy and conclude that coconut was introduced to the region by the Spaniards after their conquests.

Scope

In order to examine the value of such claims, an extensive search was conducted of the relevant historical accounts of coconut in America and in the Orient.

Key Results

The Spanish chronicler Oviedo (1478–1557) is found to have effectively used fruit and seed size to distinguish coconut from other palms. In addition, it is shown that he has been inaccurately faulted with incorrectly representing a cluster of coconuts. The original drawing, a cluster of a native Bactris, was in the marginalia and was only assigned to coconut after Oviedo''s death. Finally, the location is identified of a coastal Panamanian site described by Pedro Mártir de Anglería and where tidal dispersal of coconuts was observed.

Conclusions

This previously overlooked evidence confirms the pre-historical presence of coconut in Panama. Genetic data indicate that it must have been brought there directly or indirectly from the Philippines. But when, where and by whom remains a subject of research. Further molecular marker studies, computer simulation of natural drift and archaeological research could contribute to this research.     

Evaluation and application of methods for biological assessment of streams: summary of papers

The false mussel Mytilopsis adamsi, originally described from the Pacific coast of Panama, is reported for the first time as an introduced species in the Urías estuary and an adjacent shrimp farm, on the Pacific coasts of Mexico. In the 19th century, this species was transported from the Pacific coast of Panama, reaching the Indo-Pacific Ocean, but it had not been previously reported in other coasts of the American Pacific. Its recent and irregular presence in this small estuary near shore the Mazatlán harbor and an adjacent shrimp farm suggests that the species was introduced into the area but it cannot still be considered as a harmful invader. The presence of the species in the shrimp farm may indicate that it was introduced along with shrimp stocks from Central America, although its passive transportation to the estuary via ballast water or attached to hulls is not discounted.     

Possible Ballast Water Transfer of Lionfish to the Eastern Pacific Ocean

The Indo-Pacific Red Lionfish was first reported off the Florida coast in 1985, following which it has spread across much of the SE USA, Gulf of Mexico, and Caribbean Sea. Lionfish negatively impact fish and invertebrate assemblages and abundances, thus further spread is cause for concern. To date, the fish has not been reported on the Pacific coast of North or Central America. Here we examine the possibility of ballast water transfer of lionfish from colonized areas in the Atlantic Ocean to USA ports on the Pacific coast. Over an eight-year period, we documented 27 commercial vessel-trips in which ballast water was loaded in colonized sites and later discharged untreated into Pacific coast ports in the USA. California had the highest number of discharges including San Francisco Bay and Los Angeles-Long Beach. A species distribution model suggests that the probability of lionfish establishment is low for the western USA, Colombia and Panama, low to medium for Costa Rica, Nicaragua, El Salvador and Guatemala, medium to high for mainland Ecuador, and very high for western Mexico, Peru and the Galapagos Islands. Given the species’ intolerance of freshwater conditions, we propose that ballast water exchange be conducted in Gatún Lake, Panama for western-bound vessels carrying ‘risky’ ballast water to prevent invasion of the eastern Pacific Ocean.     

Unexpected molecular and morphological diversity of hemichordate larvae from the Neotropics

The diversity of tropical marine invertebrates is poorly documented, especially those groups for which collecting adults is difficult. We collected the planktonic tornaria larvae of hemichordates (acorn worms) to assess their hidden diversity in the Neotropics. Larvae were retrieved in plankton tows from waters of the Pacific and Caribbean coasts of Panama, followed by DNA barcoding of mitochondrial cytochrome c oxidase subunit I (COI) and 16S ribosomal DNA to estimate their diversity in the region. With moderate sampling efforts, we discovered six operational taxonomic units (OTUs) in the Bay of Panama on the Pacific coast, in contrast to the single species previously recorded for the entire Tropical Eastern Pacific. We found eight OTUs in Bocas del Toro province on the Caribbean coast, compared to seven species documented from adults in the entire Caribbean. All OTUs differed from each other and from named acorn worm sequences in GenBank by >10% pairwise distance in COI and >2% in 16S. Two of our OTUs matched 16S hemichordate sequences in GenBank: one was an unidentified or unnamed Balanoglossus from the Caribbean of Panama, and the other was an unidentified ptychoderid larva from the Bahamas. The species accumulation curves suggest that nearly all the species have been collected and only one more species might still remain undetected in the Pacific. In contrast, the Caribbean species accumulation curve suggests that further sampling could yield more than 10 additional OTUs. Tornaria from the 14 OTUs exhibited typical planktotrophic morphologies, and, in some cases, may be distinguished by differences in pigmentation and by the number of telotrochal ciliary bands, but in general, few diagnostic differences were detected.     

Documenting neotropical diversity of phoronids with DNA barcoding of planktonic larvae

Phoronid larvae, actinotrochs, are beautiful and complicated organisms which have attracted as much, if not more, attention than their adult forms. We collected actinotrochs from the waters of the Pacific and Caribbean coasts of Panama, and used DNA barcoding of mtCOI, as well as 16S and 18S sequences, to estimate the diversity of phoronids in the region. We discovered three operational taxonomic units (OTUs) in the Bay of Panama on the Pacific coast and four OTUs in Bocas del Toro on the Caribbean coast. Not only did all OTUs differ from each other by >10% pairwise distance in COI, but they also differed from all phoronid sequences in GenBank, including the four species for which adults have been reported for the Pacific of Panama, Phoronopsis harmeri, Phoronis psammophila, Phoronis muelleri, and Phoronis hippocrepia. In each ocean region, one common OTU was more abundant and occurred more frequently than other OTUs in our samples. The other five OTUs were relatively rare, with only one to three individuals collected during the entire project. Species accumulation curves were relatively flat but suggest that at least one more species is likely to be present at each site. Actinotrochs from the seven sequenced OTUs had morphologies typical of species with non‐brooded planktotrophic development and, in some cases, may be distinguished by differences in pigmentation and the arrangement of blood masses. We found one larva with morphology typical of brooded planktotrophic larvae for which sequencing failed, bringing the total number of potential species detected to eight and representing >50% of the adult species currently recognized globally.     

DNA barcoding of echinopluteus larvae uncovers cryptic diversity in neotropical echinoids

Surveys of larval diversity consistently increase biodiversity estimates when applied to poorly documented groups of marine invertebrates such as phoronids and hemichordates. However, it remains to be seen how helpful this approach is for detecting unsampled species in well‐studied groups. Echinoids represent a large, robust, well‐studied macrofauna, with low diversity and low incidence of cryptic species, making them an ideal test case for the efficacy of larval barcoding to discover diversity in such groups. We developed a reference dataset of DNA barcodes for the shallow‐water adult echinoids from both coasts of Panama and compared them to DNA sequences obtained from larvae collected primarily on the Caribbean coast of Panama. We sequenced mitochondrial cytochrome c oxidase subunit I (COI) for 43 species of adult sea urchins to expand the number and coverage of sequences available in GenBank. Sequences were successfully obtained for COI and 16S ribosomal DNA from 272 larvae and assigned to 17 operational taxonomic units (OTUs): 4 from the Pacific coast of Panama, where larvae were not sampled as intensively, and 13 from the Caribbean coast. Of these 17 OTUs, 13 were identified from comparisons with our adult sequences and belonged to species well documented in these regions. Another larva was identified from comparisons with unpublished sequences in the Barcode of Life Database (BOLD) as belonging to Pseudoboletia, a genus scarcely known in the Caribbean and previously unreported in Panama. Three OTUs remained unidentified. Based on larval morphology, at least two of these OTUs appeared to be spatangoids, which are difficult to collect and whose presence often goes undetected in standard surveys of benthic diversity. Despite its ability to capture unanticipated diversity, larval sampling failed to collect some species that are locally common along the Caribbean coast of Panama, such as Leodia sexiesperforata, Diadema antillarum, and Clypeaster rosaceus.     

Re-Evaluation of early observations on coconut in the New World

Oviedo’s sixteenth century reports from the New World, relating the characteristics, uses, cultivation and geographical distribution Cocos nucifera, are analyzed in terms of their veracity and implications for the evolution of this highly important economic species. The presence of coconut on the Pacific coasts of Costa Rica, Panama, and Colombia is supported by the documentary evidence. The variability reported suggests a gene pool which includes the “wild” and “domesticated” morphotypes in populations similar to those distributed in the South Pacific region. The diffusion to America could have involved a large number of seeds. A close association between coconuts and humans, including a variety of uses and its cultivation is reported in the early sixteenth century literature. Evidence suggests both natural diffusion and diffusion by the intervention of humans. Studies of the genetic diversity in the region will identify valuable genotypes for improvement and conservation.     

Shell utilization,predation pressure,and thermal stress in panamanian hermit crabs: An interoceanic comparison

Shell preference patterns of two common hermit crabs from hard bottom reef flats on the Caribbean coast of Panama are examined in relation to the predation pressures and physical stresses of their habitat. Clibanarius antillensis Stimpson lives in the high intertidal habitat and minimizes exposure to predators by seeking refuge during high tides. It prefers high-spired shells which maximize protection from thermal stress. Calcinus tibicen Herbst avoids tidal emersion and prefers low-spired shells which enhance resistance to the predators common on Caribbean reef flats.The results are compared with similar results from the tropical eastern Pacific Bay of Panama. Shell-crushing predation on Caribbean hermit crabs is suggested to differ quantitatively and qualitatively from predation on hermit crabs in the Bay of Panama. Predation on hermit crabs in the Bay of Panama is more intense and effects larger individuals than predation on Caribbean reef flat hermit crabs. In addition, shell-crushing predation on hermit crabs in the Bay of Panama is primarily from teleost fish predators (Diodon spp.), while predation on Caribbean hermit crabs is primarily by bottom-dwelling crustaceans.Differences in predation pressures and tidal regimes between the Caribbean and Pacific coasts of Panama are reflected in the shell preferences and behavior of hermit crabs from the two areas.     

A new species of Trinectes (Pleuronectiformes: Achiridae), with comments on the other eastern Pacific species of the genus

A new eastern Pacific achirid, Trinectes xanthurus, has a distinctive yellow or light caudal fin, without pattern, to contrast with its relatively dark body. This species also has black, posterior margins of the dorsal and anal fins and 21-22 caudal vertebrae, versus 19-20 for its Pacific coast congeners. It usually is found in shallow, continental shelf waters on sand or mud bottoms and ranges from Colombia to El Salvador. The genus Trinectes is represented in the eastern Pacific by four species. A fifth species usually occurs in Pacific slope drainages (freshwater) in Panama. Of these species, only Trinectes fonsecensis is found throughout the tropical eastern Pacific, while the other three coastal species are confined to tropical waters south of the Golfo de Tehauntepec.     

PHYLOGEOGRAPHY OF THE PANTROPICAL SEA URCHIN EUCIDARIS IN RELATION TO LAND BARRIERS AND OCEAN CURRENTS

The pantropical sea urchin genus Eucidaris contains four currently recognized species, all of them allopatric: E. metularia in the Indo-West Pacific, E. thouarsi in the eastern Pacific, E. tribuloides in both the western and eastern Atlantic, and E. clavata at the central Atlantic islands of Ascension and St. Helena. We sequenced a 640-bp region of the cytochrome oxidase I (COI) gene of mitochondrial DNA to determine whether this division of the genus into species was confirmed by molecular markers, to ascertain their phylogenetic relations, and to reconstruct the history of possible dispersal and vicariance events that led to present-day patterns of species distribution. We found that E. metularia split first from the rest of the extant species of the genus. If COI divergence is calibrated by the emergence of the Isthmus of Panama, the estimated date of the separation of the Indo-West Pacific species is 4.7–6.4 million years ago. This date suggests that the last available route of genetic contact between the Indo-Pacific and the rest of the tropics was from west to east through the Eastern Pacific Barrier, rather than through the Tethyan Sea or around the southern tip of Africa. The second cladogenic event was the separation of eastern Pacific and Atlantic populations by the Isthmus of Panama. Eucidaris at the outer eastern Pacific islands (Galapagos, Isla del Coco, Clipperton Atoll) belong to a separate clade, so distinct from mainland E. thouarsi as to suggest that this is a different species, for which the name E. galapagensis is revived from the older taxonomic literature. Complete lack of shared alleles in three allozyme loci between island and mainland populations support their separate specific status. Eucidaris galapagensis and E. thouarsi are estimated from their COI divergence to have split at about the same time that E. thouarsi and E. tribuloides were being separated by the Isthmus of Panama. Even though currents could easily convey larvae between the eastern Pacific islands and the American mainland, the two species do not appear to have invaded each other's ranges. Conversely, the central Atlantic E. clavata at St. Helena and Ascension is genetically similar to E. tribuloides from the American and African coasts. Populations on these islands are either genetically connected to the coasts of the Atlantic or have been colonized by extant mitochondrial DNA lineages of Eucidaris within the last 200,000 years. Although it is hard to explain how larvae can cross the entire width of the Atlantic within their competent lifetimes, COI sequences of Eucidaris from the west coast of Africa are very similar to those of E. tribuloides from the Caribbean. F_ST statistics indicate that gene flow between E. metularia from the Indian Ocean and from the western and central Pacific is restricted. Low gene flow is also evident between populations of E. clavata from Ascension and St. Helena. Rates of intraspecific exchange of genes in E. thouarsi, E. galapagensis, and E. tribuloides, on the other hand, are high. The phylogeny of Eucidaris confirms Ernst Mayr's conclusions that major barriers to the dispersal of tropical echinoids have been the wide stretch of deep water between central and eastern Pacific, the cold water off the southwest coast of Africa, and the Isthmus of Panama. It also suggests that a colonization event in the eastern Pacific has led to speciation between mainland and island populations.     

Secondary contact in the Sea of Japan: the case of the <Emphasis Type="Italic">Careproctus rastrinus</Emphasis> species complex (Liparidae)

The Careproctus rastrinus species complex, widely known from the North Pacific, has been revealed recently to include nine genetically divergent groups on the basis of mitochondrial DNA (mtDNA) sequence variations. Herein we describe an AFLP analysis that focuses on three closely related groups in order to clarify the evolutionary history of the species complex in the Sea of Japan and off the Pacific coast of Japan. A principal coordinate analysis indicated the absence of nuclear divergence in two groups defined by mtDNA variations in the Sea of Japan, whereas another group from the Pacific coast of northern Japan was clearly distinct. This suggests extensive gene flow between two groups in the Sea of Japan as a result of secondary contact.     

New records of benthic marine algae and Cyanobacteria for Costa Rica,and a comparison with other Central American countries

We present the results of an intensive sampling program carried out from 2000 to 2007 along both coasts of Costa Rica, Central America. The presence of 44 species of benthic marine algae is reported for the first time for Costa Rica. Most of the new records are Rhodophyta (27 spp.), followed by Chlorophyta (15 spp.), and Heterokontophyta, Phaeophycea (2 spp.). Overall, the currently known marine flora of Costa Rica is comprised of 446 benthic marine algae and 24 Cyanobacteria. This species number is an under estimation, and will increase when species of benthic marine algae from taxonomic groups where only limited information is available (e.g., microfilamentous benthic marine algae, Cyanobacteria) are included. The Caribbean coast harbors considerably more benthic marine algae (318 spp.) than the Pacific coast (190 spp.); such a trend has been observed in all neighboring countries. Compared to other Central American countries, Costa Rica has the highest number of reported benthic marine algae; however, Panama may have a similarly high diversity after unpublished results from a Rhodophyta survey (Wysor, unpublished) are included. Sixty-two species have been found along both the Pacific and Caribbean coasts of Costa Rica; we discuss this result in relation to the emergence of the Central American Isthmus.     

The Central American squirrel monkey (Saimiri oerstedii): introduced hybrid or endemic species?

Although squirrel monkeys (Saimiri spp., Primates: Cebidae) are abundant and widespread in South America, the disjunct Central American species, Saimiri oerstedii, has been restricted to the Pacific wet lowlands of Costa Rica and Panama since the earliest historical records. This taxon is now endangered in Costa Rica and nearly extinct in Panama because of habitat loss, development, and the pet trade. Conservation efforts have been hampered because of the influential, but untested, speculation that S. oerstedii represents a hybrid species introduced by prehispanic Amerind traders from multiple localities in South America. Using nuclear and mitochondrial DNA sequence data to reconstruct the phylogenetic relationships among Saimiri from Central and South America, we reexamine the taxonomic status of squirrel monkeys from different geographic regions. The sequence data support P. Hershkovitz's (1984, Am. J. Primatol. 6: 257-281) taxonomy advocating four distinct species. Combining this information with evidence from the fossil record to date the divergence times among sister taxa, we test and reject the hypothesis that Central American squirrel monkeys are the result of human introduction.     

Systematics of myliobatoid elasmobranchs: with emphasis on the phylogeny and historical biogeography of neotropical freshwater stingrays (Potamotrygonidae: Rajiformes)

The neotropical freshwater family Potamotrygonidae appears to be the only stingray group that has radiated in a non-marine environment. To assess the affinities of potamotrygonids to other rays, a phylogenetic analysis was undertaken using 39 morphological characters from 18 stingray groups. The single tree produced (CI = 0.80, RI = 0.88) suggests that neotropical freshwater rays are a monophyletic group, and that within Potamotrygonidae, Paratrygon is basal to a clade composed of Plesiotrygon and Potamotrygon. The sister group to potamotrygonids was determined to be amphi-American Himantura —these taxa share synapomorphies of the ventral mandibular musculature and the hyomandibular/mandibular articulation. The topology suggests that potamotrygonids are derived from a freshwater-invading ancestor that was distributed along the northern coast of South America (Pacific and Caribbean) prior to the emergence of the isthmus of Panama. This hypothesis conflicts with parasite-based biogeographic scenarios of a stricdy Pacific origin for potamotrygonids. General systematic results concerning urolophids, dasyatids, and pelagic myliobatoid stingrays are also discussed.     

Genetic diversity of the mangrove‐associated alga Bostrychia radicans/Bostrychia moritziana (Ceramiales,Rhodophyta) from southern Central America

Previous studies suggested that the biodiversity of the mangrove‐associated Bostrychia radicans/Bostrychia moritziana species complex on the Pacific coast of Central America, based on genetic and reproductive data, were low compared with similar areas on the Atlantic coast. Evolutionary scenarios were proposed based on either a recent introduction to the Pacific, or a more uniform environment leading to genetically connected populations and low differentiation between populations. We sampled more extensively in southern Mexico, Guatemala and El Salvador and sequenced the samples for the RuBisCo spacer. Our results show that genetic diversity is high in these populations. Many haplotypes retrieved are also found in the Atlantic Ocean (USA, Brazil), an observation not made before. Data suggest that populations are highly differentiated with little evidence of isolation‐by‐distance. The population at La Puntilla, El Salvador is highly differentiated from other populations. Data also suggest that diversity is reduced in a northerly direction, with only one haplotype, unique to Pacific Central America, found north of Chiapas, Mexico. This could be due to northern expansion of this unique genotype as sea surface temperatures ameliorated following the last glacial maximum. Our data do not support the previous proposition of low diversity in the east central Pacific and suggest that much of the Pacific Central America diversity is from before the closure of the Isthmus of Panama.     

Revision of the eastern Pacific species of Gobulus (Perciformes: Gobiidae), with description of a new species

Gobulus birdsongi is described as a new species from the Pacific coast of Panama. The two other known species of Gobulus from the eastern Pacific, Gobulus crescentalis and G. hancocki are redescribed. Gobulus birdsongi differs from other species in the genus in having more numerous second dorsal and anal rays. Gobulus hancocki differs from G. crescentalis in having a much smaller eye. The genus is distinctive in having reversed countershading, with the ventral surface of the body darker than the dorsal surface.     

Diversity of the <Emphasis Type="Italic">CO1</Emphasis> Gene of Mitochondrial DNA in Representatives of Genus <Emphasis Type="Italic">Antimora</Emphasis> (Moridae,Gadiformes) in the World Oceans

The frequency of occurrence of the COI gene of mitochondrial DNA in the Pacific flatnose Antimora microlepis and blue antimora Antimora rostrata (Moridae, Gadiformes) was analyzed in samples collected in different areas of the World Ocean. The revealed maximum haplotype diversity of COI in the blue antimora in the North Atlantics may indicate that this species emerged in this region, from which it widely distributed in the World Ocean. The Pacific flatnose Antimora microlepis originated from Antimora rostrata. Antimora might penetrate into the North Pacific by several routes: through the Panama Strait, along the coast of Antarctica, or through the Indian Ocean along the coast of Australia.     

Hemicyclops columnaris sp.n. (Copepoda, Poecilostomatoida, Clausidiidae) Associated with a Coral in Panama (Pacific Side)

Hemicyclops columnaris sp.n. is described from the scleractinian coral Porites lobata Dana on the Pacific coast of Panama. The new species may be distinguished from its 23 congeners by a combination of the body size and the shape of the genital segment in the female. Although certain members of the genus Hemicyclops are associated with crustaceans, polychaetes and other marine invertebrates, this is the first record of a species of this genus from a hard coral.     

Preliminary survey of a nemertean crab egg predator,Carcinonemertes, on its host crab,Callinectes arcuatus (Decapoda,Portunidae) from Golfo de Nicoya,Pacific Costa Rica

The possible presence of egg predators in brood masses of portunid crabs from Pacific Central America has not been studied yet. This survey reports the finding of a nemertean crab egg predator on the portunid crab, Callinectes arcuatus, from the Golfo de Nicoya, Pacific Costa Rica. Nemerteans were found in the egg masses of 26 out of the 74 crabs for a prevalence of 35%. The intensity (mean number of worms/ infected crab) was estimated to be 18 with a variance of 1–123 worms/infected crab. No nemerteans were observed either in the 19 Callinectes arcuatus from Golfo Dulce (southern Pacific coast) and the 10 Portunus asper from Herradura-Jaco (central Pacific coast). This nemertean is a member of the genus Carcinonemertes, which has been reported from the Caribbean coast of Panama. However, the encountered Carcinonemertes sp. is the first published finding and report from Costa Rica and Pacific Central America.