VARIATION IN LEAFLET MORPHOLOGY AMONG POPULATIONS OF CARIBBEAN CYCADS (ZAMIA)

Caribbean cycads (Zamia L.) are well known for variation in leaflet morphology. Here, I examine the variation in leaflet morphology among five populations of Zamia in Florida. Variables measured were leaflet length and width, leaflet length: width ratio, leaflet surface area, rachis and petiole lengths, number of leaflets per leaf, and total leaf area. In addition to comparisons among the Florida populations, these populations are compared with three previously studied populations in Puerto Rico (Newell, 1986). For the most part, the populations exhibited significant differences in all variables. In spite of the extensive morphological variation in this group of plants, leaflet morphology has long been important in Zamia taxonomy. The Florida populations would be considered by some to be either one, two, or three different species. The Florida and Puerto Rico populations together might be considered as few as one species or as many as five species. Based upon the data presented here, the five Florida populations appear to represent a single species; and the three Puerto Rico populations appear to represent two additional species. Further clarification of the taxonomy of Caribbean Zamia will require additional characters such as cone morphology and protein or DNA patterns.     

Two distinct, geographically overlapping lineages of the corallimorpharian Ricordea florida (Cnidaria: Hexacorallia: Ricordeidae)

We examined the genetic variation of the corallimorpharian Ricordea florida; it is distributed throughout the Caribbean region and is heavily harvested for the marine aquarium trade. Eighty-four distinct individuals of R. florida were sequenced from four geographically distant Caribbean locations (Cura?ao, Florida, Guadeloupe, and Puerto Rico). Analysis of the ribosomal nuclear region (ITS1, 5.8S, ITS2) uncovered two geographically partially overlapping genetic lineages in R. florida, probably representing two cryptic species. Lineage 1 was found in Florida and Puerto Rico, and Lineage 2 was found in Florida, Puerto Rico, Guadeloupe, and Cura?ao. Because of the multi-allelic nature of the ITS region, four individuals from Lineage 1 and six from Lineage 2 were cloned to evaluate the levels of hidden intra-individual variability. Pairwise genetic comparisons indicated that the levels of intra-individual and intra-lineage variability (<1%) were approximately an order of magnitude lower than the divergence (~9%) observed between the two lineages. The fishery regulations of the aquarium trade regard R. florida as one species. More refined regulations should take into account the presence of two genetic lineages, and they should be managed separately in order to preserve the long-term evolutionary potential of this corallimorpharian. The discovery of two distinct lineages in R. florida illustrates the importance of evaluating genetic variability in harvested species prior to the implementation of management policies.     

Colonization of islands in the Mona Passage by endemic dwarf geckoes (genus Sphaerodactylus) reconstructed with mitochondrial phylogeny

Little is known about the natural history of the Sphaerodactylus species endemic to the three islands located in the Mona Passage separating the Greater Antillean islands of Hispaniola and Puerto Rico. In this study, parts of two mitochondrial genes, 16S rRNA and 12S rRNA, were sequenced to determine the relationships between the sphaerodactylids that live in the Mona Passage and other Caribbean species from the same genus. While the main goal was to identify the biogeographical origin of these species, we also identified a genetically distinct type of dwarf gecko that warrants future evaluation as a possible new species. According to the reconstructed phylogenies, we propose a stepwise model of colonization wherein S. nicholsi from southwestern Puerto Rico or a very close ancestor gave rise through a founder event to Sphaerodactylus monensis on Mona Island. In a similar fashion, S. monensis or a very close ancestor on Mona Island gave rise to S. levinsi on Desecheo Island. This study also suggests that the most recent common ancestor between the species from the islands in the Mona Passage and Puerto Rico existed approximately 3 MYA.     

Response of sponges with autotrophic endosymbionts during the coral-bleaching episode in Puerto Rico

An updated list of sponges with algal endosymbionts including new records for Puerto Rico and the Caribbean, indicates that thirty-five species of common Caribbean sponges possess photosynthetic endosymbionts. Of these, 23 (67.6%) species in seven orders, were found with unicellular chroococcoid cyanobacteria (Aphanocapsa-like) and 5 (14.7%) hadromerid species were found with zooxanthellae. Sponges with other algae as symbionts occur less frequently (6%). Thirty-one common sponge species were inspected for bleaching during coral-bleaching months (July-September 1987; January 1988) in Puerto Rico. Anthosigmella varians, Xestospongia muta and Petrosia pellasarca bleached partially, but only few individuals within any given population became bleached and the bleaching of sponges was very localized. Adaptations between cyanobacterial symbionts and sponges, acquired during the long evolutionary history of these two taxa may explain the paucity of bleached sponges when compared to the high incidence of bleached corals reported.     

Athecate hydroids from Puerto Rico and the virgin Islands

About 31 athecate hydroids collected from Puerto Rico and the vicinity are recorded, 9 for the first time in the Caribbean Sea. Two new genera, Heterocoryne and Millardiana, and four new species, Heterocoryne caribbensis, Millardiana longitentaculata, Silhouetta puertoricensis and Stylactis sandrae, are described.     

Jacquinia pauciflora (Theophrastaceae), a new species from Puerto Rico

A new species,Jacquinia pauciflora, is described from subtropical moist forest on karst hills (mogotes) in northwestern Puerto Rico.Jacquinia pauciflora is similar toJ. stenophylla Urban, from which it differs in several floral and leaf characters, but is possibly most closely related toJ. umbellata A. DC. A key to the species ofJacquinia occurring in Puerto Rico is provided.     

Four New Ladybug Species Belonging to Decadiomus Chapin (Coleoptera: Coccinellidae) from Puerto Rico

While searching for native natural enemies attacking invasive insect pests in Puerto Rico, we found four undescribed ladybug species belonging to the Caribbean ladybug genus Decadiomus Chapin. In this article, we describe the following species from Puerto Rico: Decadiomus seini n. sp., Decadiomus ramosi n. sp., Decadiomus hayuyai n. sp., and Decadiomus martorelli n. sp. Illustrations of the dorsal habitus, shape of prosternal carinae, and drawings of male and female genitalia are presented. We also present a key for Diomini of Puerto Rico and discuss their importance as potential biocontrol agents.     

Regeneration filters in post-agricultural fields of Puerto Rico and Ecuador

To better understand how rainforest regenerates after agriculture I sampled the seed rain and seed bank, and set out seeds and seedlings, on microsites defined by distance to the forest in fields both in Puerto Rico and Ecuador. I found that (1) total seeds, species richness and life-form richness were twice as great in the Ecuador seed rain compared to Puerto Rico but Puerto Rico seeds were more evenly distributed among species and (2) total seedlings from the seed bank were similar between Puerto Rico and Ecuador, (3) the majority of seeds were lost to predation among all species and study sites, (4) seed disease was absent in P. aduncum and Miconia prasina, and no seeds germinated for Gonzalagunia spicata and P. riparia, (5) in Ecuador pathogenic disease claimed more seeds than germinated for all species, and Solanum ovalifolium was the only species that had seeds germinate but did not lose seeds to disease, (6) also in Ecuador, insect predation was significantly lower in the forest border for P. aduncum, and seed disease was significantly greater at the 10 m micro site for S. ovalifolium, (7) distance has a significant effect on seedling height and basal diameter, (8) losses of leaf area due to herbivory and pathogens were always low and (9) biomass and leaf specific mass were significantly reduced in the border and forest microsites. I conclude that Ecuador fields had more seeds, species, and life-forms than Puerto Rico fields, predation was the most severe post-dispersal seed filter in all fields, seeds that survived predation on Puerto Rico were lost either to disease or germination but to both mechanisms in Ecuador, all three seed mechanisms in Ecuador fields showed distance effects of seedling growth but not survivorship.     

A new species of Lepanthes (Orchidaceae) from Puerto Rico

A new species,Lepanthes caritensis, is described and illustrated. There are now nine species ofLepanthes known to occur in Puerto Rico. Plant habitat and floral morphology ofL. caritensis is most similar to that ofL. sanguinea. A survey of the area where the new species is located suggests that it is rare and should be protected under the Federal Endangered Species Act. Una especic nueva,Lepanthes caritensis, es descrita e ilustrada para Puerto Rico. Con ésta hay ahora nueve especies deLepanthes en Puerto Rico. El hábito de la planta y la morfología de la flor deL. caritensis es más similar aL. sanguinea. Un muestreo del área donde se encuentra la nueva especie sugiere que ésta es rara y que debería ser protegida por el Federal Endangered Species Act.     

A new species of Litarachna (Acari,Hydrachnidia, Pontarachnidae) from a Caribbean mesophotic coral ecosystem

New records of pontarachnid mites (Acari: Hydrachnidia) from the Caribbean island of Puerto Rico are presented. Litarachna lopezae sp. n., from substrata collected from Bajo de Sico, a mesophotic coral reef ecosystem in Mona Passage off Puerto Rico, is described as new to science. The new species was collected from nearly 70 m depth, the greatest depth from which pontarachnid mites have been found until now. In addition, a Litarachna sp. was also found in association with the tube of the polychaete Sabellastarte magnifica (Shaw, 1800) at the shallow waters of north Puerto Rico.     

Metapeyssonnelia tangerina (Peyssonneliaceae,Rhodophyta), a new species associated with coral reef habitats in Puerto Rico,Caribbean Sea

A new Metapeyssonnelia species that comprises up to 7% bottom cover at shallow‐water reef habitats in southwest Puerto Rico is described herein. It forms conspicuous orange encrustations on hard substrata and does not grow on living coral as does its two Caribbean congeners. The new species possesses conspicuous, to 30 cm in extent, tightly adherent crusts up to 950 μm thick, only hypobasal calcification, hypothallial cells arranged in broad flabellules and superficial (raised) tetrasporangial and carposporangial nemathecia. Tetrasporangia are pedicellate, borne laterally from cup‐like cells that are derived from basal paraphysal cells. Tetrasporangia measure up to 120 μm long and individual carposporangia to 80 μm long. The new species differs from other Metapeyssonnelia species developmentally in that perithallial cells at mid thallus height will divide laterally to form a new hypothallium. Small subunit gene sequences relate the new species to the two Metapeyssonnelia species that are previously known from Puerto Rico.     

Immigration history of amphidromous species on a Greater Antillean island

Aim To use molecular data to test for dispersal structuring in the immigration history of an amphidromous community on an island. Location The Caribbean island of Puerto Rico. Methods Mitochondrial DNA sequences were obtained from 11 amphidromous species, including shrimps, fish and a gastropod, sampled from throughout the island. The timing of population expansion (T_E) in each species was calculated using nucleotide variation and molecular clock dating methods. The order of species accumulation was then reconstructed (oldest to most recent estimate for T_E), and groups of species with non‐overlapping estimates for T_E were identified. The temporal span and average immigration rate for each group were calculated and compared with expectations of two previously published models of island immigration [the ‘dispersal‐structured model of island recolonization’ ( Whittaker & Jones, Oikos, 1994 , 69 , 524–529), which predicts short phases of rapid immigration followed by extended phases with relatively slow immigration rates; and the ‘colonization window hypothesis’ ( Carine, Taxon, 2005 , 54 , 895–903), which suggests that opportunities for island colonization are temporally constrained to discrete waves of colonization]. Results The molecular data indicated the immigration history of Puerto Rican amphidromous fauna from the late Pleistocene through the Holocene and identified two groups of species with non‐overlapping estimates for T_E and one group that overlapped with the other two groups. The temporal span, average immigration rate and lack of discreteness between all three groups indicated a continuum of immigration rather than distinct phases of species arrivals. Main conclusions This study did not support the expectations of the immigration models and suggested that amphidromous species from Puerto Rico comprise a single class of marine‐based dispersers. The immigration sequence we report probably reflects a recolonization chronology in this community, in keeping with the notion of species turnover through time. Four areas of future research into the immigration history of amphidromous species on islands are identified, and indicated the possibility that equilibrium processes govern long‐term community change in amphidromous biota on islands.     

Saurian malaria in the Caribbean: Plasmodium azurophilum sp. nov., a malarial parasite with schizogony and gametogony in both red and white blood cells

A survey of 466 Caribbean lizards found Plasmodium parasites present in Anolis species only of five islands. Parasites presently considered to be P. floridense occurred on Grand Cayman, North Bimini, Jamaica, Hispaniola (Haiti), and Puerto Rico. A second species, P. azurophilum, is described as new from Anolis cybotes of Haiti, A. krugi of Puerto Rico, and A. lineatopus and A. grahami of Jamaica. It lacks visible pigment in erythrocytic host cells but can produce it occasionally. Both asexual and sexual forms occur in a variety of white blood cells, notably in azurophil granulocytes and polymorphonuclear leucocytes. Experimental infections indicate that the leucocytic phase occurs after the acute erythrocytic infection declines, thus suggesting that the schizogonic and gametogonic cycles in white cells may represent an adaptive defense against immune mechanisms of the host. Mean numbers of nuclei in schizonts and mean gametocyte size are influenced by host species and type of host cell.     

PROGRESSIVE LOSS OF ANT-RELATED TRAITS OF CECROPIA PELTATA ON SELECTED CARIBBEAN ISLANDS

Cecropia of South and Central America, and Trinidad and Tobago, maintain a symbiotic relationship with Azteca ants. The plant provides a domicile and food supply and in return the ants defend the plant from insect attack and/or vine overgrowth. Cecropia is present on certain Caribbean islands, north of Trinidad and Tobago, but does not maintain a relationship with ants. The Cecropia on Puerto Rico have lost all structures normally associated with the production of the ant food. On the islands between Trinidad and Puerto Rico, stages in the loss of traits related to the ant symbiosis are present.     

Origin of the green iguana (Iguana iguana) invasion in the greater Caribbean Region and Fiji

Invasive populations of green iguanas (Iguanidae: Iguana iguana) are widely established beyond their native Central, South American, and Lesser Antillean range in various islands of the Pacific, Florida USA, and in the Greater Caribbean Region. Although widespread, information about these invasions is scarce. Here we determine the origin of invasive populations of green iguanas in Puerto Rico, Fiji, The Caymans, Florida USA, The Dominican Republic, the US Virgin Islands (USVI) of St. Thomas and St. Croix, and a U.S.A pet store. We sampled 120 individuals from these locations and sequenced one mitochondrial (ND4) and two nuclear (PAC and NT3) loci. We also include a preliminary characterization of population structure throughout Puerto Rico using six microsatellite loci to genotype individuals across 10 sampling sites. Comparing the genealogical relationships of all our samples to published sequencing data from the native range, we found that sampled populations were largely a product of populations from Colombia and El Salvador; two countries with multiple, industrial-size pet iguana farming operations. Notably, we found that haplotypes detected exclusively in the USVI and Puerto Rico’s outlying island of Vieques are closely linked to green iguanas native to Saba and Montserrat (Lesser Antilles); a clade not reported in the pet trade. Our population genetic analyses did not reveal isolation among sampling sites in Puerto Rico, rather the evidence supported admixture across the island. This study highlights the roles of the pet trade and lack of regulation in the spread of green iguanas beyond their native range.

     

Molecular phylogeny of an endemic radiation of Cuban toads (Bufonidae: Peltophryne) based on mitochondrial and nuclear genes

Aim In this study we present a molecular phylogenetic and phylogeographical analysis of Peltophryne (Anura: Bufonidae), an endemic genus of Antillean toads, to investigate the spatial and temporal origins of the genus, with particular focus on the eight Cuban species. Location Greater Antilles, with extensive sampling of the Cuban archipelago. Methods We obtained DNA sequence data from two mitochondrial genes, cytochrome c oxidase subunit I (COI) and ribosomal RNA (16S), for 124 toads representing all eight Cuban species, and combined this with published data from Hispaniola (one of three species) and Puerto Rico (one of one species) to establish a molecular phylogeny for Peltophryne. In addition, we explored the phylogeographical structure of widespread Cuban species. For a subset of 42 toads we also obtained DNA sequence data from two nuclear genes, recombination activator‐1 (RAG‐1) and chemokine receptor 4 (CXCR‐4). We combined our molecular data with published DNA sequences from a global sample of bufonid toads to place the spatial and temporal origins of Peltophryne in the Caribbean within a fuller geographical and phylogenetic context. Results All phylogenetic analyses supported the monophyly of West Indian toads. The ancestor of Peltophyrne diverged from its mainland source around the Eocene–Oligocene boundary, with a subsequent radiation across the Caribbean islands taking place during the Miocene. Cuban species are monophyletic with a basal split in the early–middle Miocene that separates extant small‐bodied from large‐bodied species. Extensive mitochondrial DNA (mtDNA) sampling within widespread Cuban species revealed contrasting phylogeographical patterns. Peltophryne taladai and P. empusa showed deeply divergent lineages, whereas no geographical structure was observed in the widespread P. peltocephala. Main conclusions Our timeline for Peltophryne diversification is consistent with a biogeographical model requiring no long‐distance overwater dispersal. Although confidence intervals on divergence time estimates are wide, the stem age of Peltophyrne coincides with the hypothesized GAARlandia landspan or archipelago, which may have connected South America briefly with the Antilles. The ages of Peltophryne for Puerto Rico, Hispaniola and Cuba are consistent with a recently proposed vicariance scenario for the region. Our molecular results support the recognition of all eight species in Cuba, and provide evidence of possible cryptic species.     

A capture–recapture model of amphidromous fish dispersal

Adult movement scale was quantified for two tropical Caribbean diadromous fishes, bigmouth sleeper Gobiomorus dormitor and mountain mullet Agonostomus monticola, using passive integrated transponders (PITs) and radio‐telemetry. Large numbers of fishes were tagged in Río Mameyes, Puerto Rico, U.S.A., with PITs and monitored at three fixed locations over a 2·5 year period to estimate transition probabilities between upper and lower elevations and survival probabilities with a multistate Cormack–Jolly–Seber model. A sub‐set of fishes were tagged with radio‐transmitters and tracked at weekly intervals to estimate fine‐scale dispersal. Changes in spatial and temporal distributions of tagged fishes indicated that neither G. dormitor nor A. monticola moved into the lowest, estuarine reaches of Río Mameyes during two consecutive reproductive periods, thus demonstrating that both species follow an amphidromous, rather than catadromous, migratory strategy. Further, both species were relatively sedentary, with restricted linear ranges. While substantial dispersal of these species occurs at the larval stage during recruitment to fresh water, the results indicate minimal dispersal in spawning adults. Successful conservation of diadromous fauna on tropical islands requires management at both broad basin and localized spatial scales.     

The Systematics and Reproductive Biology of the Galeus arae Subspecific Complex (Chondrichthyes: Scyliorhinidae)

Galeus arae is currently classified as a complex of three subspecies (Galeus arae arae, Galeus arae antillensis, and Galeus arae cadenati). Morphometric and meristic analyses, size at maturity, and variation in color patterns, support the recognition of these taxa as distinct species. All species have well-developed nidamental glands and are oviparous. Galeus arae comprises two geographically disjunct populations that are not distinguishable by the characters we examined. A northern population occurs along the east and Gulf coasts of North America from South Carolina to the Mississippi delta, and the northern coast of Cuba to the north-eastern tip of the Yucatan. A southern population occurs along the Caribbean coasts of Honduras, Nicaragua, and Costa Rica, and some neighboring islands. Galeus antillensis occurs on the northern coasts of Hispaniola and Puerto Rico, and off many of the Leeward Islands. Galeus cadenati occurs off the Caribbean coasts of Panama and Colombia. Distributional data suggest that the three species are distributed allopatrically.