Checklist of helminth parasites of the cane toad Bufo marinus (Anura: Bufonidae) from Mexico

Thirty-four adult cane toads Bufo marinus L. (12 males and 22 females) collected from 2 localities in Mexico (Cerro de Oro and Temascal Dams, Oaxaca) in September 2003 were examined for helminth parasites. In total, 14,749 helminths belonging to 14 taxa were collected. Included were 2 adult digeneans (Choledocystus hepaticus, Mesocoelium monas); 1 larval cestode (an unidentified pseudophyllidean); and 11 nematodes, including 3 species of larvae (Contracaecum sp., Physaloptera sp., Physocephalus sexalatus) and 8 species of adults (Aplectana itzocanensis, Cosmocerca sp., Cruzia morleyi, Ochoterenella digiticauda, Oswaldocruzia sp., Raillietnema sp., Rhabdias americanus, and Rhabdiasfuelleborni). Higher species richness was recorded in B. marinus from Cerro de Oro (12 taxa versus 9 in those from Temascal); hosts from both localities shared 7 taxa. There were 25 new locality records, and 2 taxa were registered in Mexico for the first time. To date, 112 helminth species have been recorded parasitizing B. marinus along its native and introduced range of distribution, with 40.5% of them reported from Mexico.     

Strongyluris panamaensis n. sp. (Nematoda: Heterakidae) and other helminths from the lizard,Anolis biporcatus (Sauria: Polychrotidae), from Panama

Strongyluris panamaensis n. sp. from the intestine of Anolis biporcatus collected in Panama is described and illustrated. Strongyluris panamaensis represents the 30th species assigned to this genus and the fourth from the neotropical region. Strongyluris panamaensis can be easily distinguished from all other neotropical species because it has the shortest spicule. In addition to S. panamaensis, 3 species of Trematoda, Mesocoelium monas, Parallellopharynx arctus, and Urotrema scabridum, and 3 species of Nematoda, Crytosomum penneri, Oswaldocruzia bainae, and Piratuba digiticauda, were identified.     

Helminths of two anuran species, Atelopus spurrelli (Bufonidae) and Dendrobates histrionicus (Dendrobatidae), from Colombia, South America

Two species of anurans from Colombia, South America, Atelopus spurrelli and Dendrobates histrionicus, were examined for helminths. A. spurrelli was found to harbor three species of Nematoda, adults of Cosmocerca podicipinus and larvae of Physocephalus sp. and Porrocaecum sp. D. histrionicus was found to harbor, in addition to C. podicipinus, Physocephalus sp. and Porrocaecum sp. and cystacanths of three species of Acanthocephala: Centrorhynchus sp., Onicola sp. and Polymorphus sp. A. spurrelli and D. histrionicus represent new host records for these helminths.     

A new species of Spinicauda (Nematoda: Heterakidae) and other endoparasites in Platymantis pelewensis (Anura: Ranidae) from the Palau Islands, Republic of Belau, Oceanica

Spinicauda fisheri n. sp. from the intestine of Platymantis pelewensis collected in the Palau Islands, Republic of Belau, is described and illustrated. Spinicauda fisheri represents the 12th species assigned to the genus and the first from Oceanica. It is most similar to Spinicauda spinicauda in that the tail ends in a filament, but it has 19 pairs of caudal papillae as compared with 5 pairs in S. spinicauda. In addition to S. fisheri, 2 species of Cestoda, cysticercoids of Joyeuxiella sp. and tetrathyridia of Mesocestoides sp., 2 species of Nematoda, Oswaldocruzia bakeri and Physocephalus sp. (larvae in cysts), 1 species of Acanthocephala, Acanthocephalus bufonis, and 1 species of Pentastomida, nymphs of Kiricephalus pattoni were identified.     

Ultraviolet exposure and vitamin D synthesis in a sun-dwelling and a shade-dwelling species of Anolis: are there adaptations for lower ultraviolet B and dietary vitamin D3 availability in the shade?

We compared the natural ultraviolet B (UV-B) exposure, dietary vitamin D, and skin-generated vitamin D synthesis for adult males of two species of Jamaican anoles. The more shade-tolerant and thermal-conforming Anolis lineotopus merope, rarely exposed to full sun, experienced less UV-B irradiation in its shady environment than the more heliophilic and thermophilic Anolis sagrei, which frequently basked in full sun during the morning hours (0800-1100 hours). Both species obtained detectable levels of vitamin D(3) in their diet, but the heliophilic A. sagrei obtained more. To compensate for less availability of UV-B and dietary vitamin D, the skin of A. lineotopus merope seems to have acquired a greater sensitivity than that of A. sagrei regarding UV-B-induced vitamin D(3) photobiosynthesis. We assessed this by observing a greater conversion of provitamin D to photoproducts in skin exposed to UV-B from a sunlamp. The reduced skin sensitivity of A. sagrei regarding vitamin D photobiosynthesis may reflect a correlated response associated with less need for vitamin D photobiosynthesis and greater need for UV-B screening capacity as an adaptation to a more damaging UV-B environment. However, the possibility that adaptations for photobiosynthesis of vitamin D and for protection from skin damage could involve independent mechanisms needs investigation. Also, the ability to behaviorally regulate UV-B exposure, as shown for the panther chameleon, would benefit both species of Anolis and should be investigated.     

Cosmocerca vrcibradici n. sp. (Ascaridida: Cosmocercidae), Oswaldocruzia vitti n. sp. (Strongylida: Molineoidae), and other helminths from Prionodactylus eigenmanni and Prionodactylus oshaughnessyi (Sauria: Gymnophthalmidae) from Brazil and Ecuador

Cosmocerca vrcibradici n. sp. and Oswaldocruzia vitti n. sp., intestinal parasites of Prionodactylus eigenmanni (type host) and P. oshaughnessyi, are described and illustrated. Of the 19 valid species of Cosmocerca, C. vrcibradici n. sp. represents the 12th Neotropical species and the first species to be reported from lizard hosts. Of the 76 valid species of Oswaldocruzia, O. vitti n. sp. represents the 14th Neotropical species and the eighth species to be reported from lizard hosts. In addition, 1 species of Digenea, Mesocoelium monas, and 1 species of Acanthocephala, Acanthocephalus saurius, were found.     

Microhabitat use, diet, and performance data on the Hispaniolan twig anole, Anolis sheplani: pushing the boundaries of morphospace

Caribbean Anolis lizards are often cited as a textbook example of adaptive radiation. Similar morphologies (ecomorphs) have originated in similar ecological settings on different large islands in the West Indies. However, relatively little is known about one of the morphologically most specialized and divergent ecomorphs: the twig anoles. Here, we investigate aspects of morphology, dewlap size, locomotor and bite performance, structural habitat and diet of the poorly known twig anole, Anolis sheplani from Hispaniola. Few observations have previously been made of this species in its natural habitat, and few quantitative data on its natural history are available. A. sheplani is an extreme twig anole with respect to its morphology, performance capacities, and ecological niche. Males and females of this species do not differ from each other in body dimensions, performance or habitat use, but males do have a bigger dewlap than females. We present data for 25 individuals and compare them with data for other Greater Antillean anoles. It becomes apparent that twig anoles constitute a large component of the morphological, functional, and ecological diversity of Anolis lizards. Small twig anoles such as A. sheplani appear to be pushing the boundaries of morphospace and are thus crucial in our understanding of the evolution of phenotypic diversity.     

Acanthocephalus saurius n. sp. (Acanthocephala: Echinorhynchidae) and other helminths from the lizard Norops limifrons (Sauria: Polychrotidae) from Costa Rica

Acanthocephalus saurius n. sp. from the small intestine of Norops limifrons collected in Costa Rica is described and illustrated. Acanthocephalus saurius n. sp. represents the third acanthocephalan species in which gravid individuals have been found in lizard hosts. In addition to A. saurius; 1 species of Trematoda, Mesocoelium monas; 3 species of Nematoda, Piratuba digiticauda, Strongyluris panamaensis, and Acuariidae gen. sp. (larvae in cysts) were found.     

Intraspecific variation in Anolis sagrei mirrors the adaptive radiation of Greater Antillean anoles

Anolis lizards of the Greater Antilles represent one of the classic examples of vertebrate adaptive radiation. The same morphological types ('ecomorphs') have evolved repeatedly in response to similar ecological pressures on different islands. We tested whether patterns of within species diversification were congruent with between species patterns and the processes leading to the adaptive radiation of Greater Antillean anoles by measuring variation in performance-related morphological characters in the brown anole, Anolis sagrei . We measured morphological and genetic variation in two different habitat types on each of five islands in the Bahamas. We estimated population structure and rates of gene flow within and among islands using eight microsatellite markers. Intraspecific variation in performance-related morphological characters was similar to the pattern of interspecific variation that characterizes the adaptive radiation of this group in the Greater Antilles. For example, limb length was correlated with perch diameter within A. sagrei as has also been shown among species of anole. Morphological divergence in traits has occurred despite relatively high levels of gene flow both within and among islands. These results are discussed in the context of the divergence-with-gene-flow model of speciation. The results provide important intraspecific evidence that the diversification of anoles has been shaped by natural selection and show how ecologically-based selection pressures explain diversification at both the population and species levels.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 189–199.     

Testing the island effect in adaptive radiation: rates and patterns of morphological diversification in Caribbean and mainland Anolis lizards

Many of the classic examples of adaptive radiation, including Caribbean Anolis lizards, are found on islands. However, Anolis also exhibits substantial species richness and ecomorphological disparity on mainland Central and South America. We compared patterns and rates of morphological evolution to investigate whether, in fact, island Anolis are exceptionally diverse relative to their mainland counterparts. Quite the contrary, we found that rates and extent of diversification were comparable--Anolis adaptive radiation is not an island phenomenon. However, mainland and Caribbean anoles occupy different parts of morphological space; in independent colonizations of both island and mainland habitats, island anoles have evolved shorter limbs and better-developed toe pads. These patterns suggest that the two areas are on different evolutionary trajectories. The ecological causes of these differences are unknown, but may relate to differences in predation or competition among mainland and island communities.     

The guest playing host: colonization of the introduced Mediterranean gecko, Hemidactylus turcicus, by helminth parasites in southeastern Louisiana.

Parasite surveys of exotic hosts offer the opportunity to examine parasite colonization on different scales (i.e., host individual, host population, host species, and new geographic locality). Ten helminths (Macracanthorhynchus ingens, Mesocestoides lineatus, Oochoristica javaensis, Haematoloechus varioplexus, Mesocoelium monas, Telorchis corti, Cosmocercoides variabilis, Oswaldocruzia leidyi, Skrjabinoptera sp., and a larval acuariid nematode) were recovered from the exotic Mediterranean gecko Hemidactylus turcicus, in southeastern Louisiana. Only 1 exotic parasite, O. javaensis, colonized a new geographic locality, but 7 local helminths colonized a new host species. Helminth communities of H. turcicus were similar in structure to what has been hypothesized or observed for lizards. Thus, communities were composed of generalists and were depauperate (i.e., colonization of individual geckos or host populations was rare for most of the helminths); however, there was significant variation in community structure among local habitats. Although the gecko's behavioral and physiological attributes predict colonization by monoxenous helminths, only 2, C. variabilis and O. leidyi, were recovered. Eight heteroxenous helminths, 2 of which (the acuariid and O. javaensis) were the most widely distributed and abundant, were the better colonizers. The gecko's generalist diet may have exposed it to a diverse parasite fauna and thus been important in determining the helminths that could colonize.     

Out of Cuba: overwater dispersal and speciation among lizards in the Anolis carolinensis subgroup

Overwater dispersal and subsequent allopatric speciation contribute importantly to the species diversity of West Indian Anolis lizards and many other island radiations. Here we use molecular phylogenetic analyses to assess the contribution of overwater dispersal to diversification of the Anolis carolinensis subgroup, a clade comprising nine canopy-dwelling species distributed across the northern Caribbean. Although this clade includes some of the most successful dispersers and colonists in the anole radiation, the taxonomic status and origin of many endemic populations have been ambiguous. New mitochondrial and nuclear DNA sequences from four species occurring on small islands or island banks (Anolis brunneus, Anolis longiceps, Anolis maynardi, Anolis smaragdinus) and one species from the continental United States (A. carolinensis) are presented and analysed with homologous sequences sampled from related species on Cuba (Anolis allisoni and Anolis porcatus). Our analyses confirm that all five non-Cuban species included in our study represent distinct, independently evolving lineages that warrant continued species recognition. Moreover, our results support Ernest Williams's hypothesis that all of these species originated by overseas colonization from Cuban source populations. However, contrary to Williams's hypothesis of Pleistocene dispersal, most colonization events leading to speciation apparently occurred earlier, in the late Miocene-Pliocene. These patterns suggest that overwater dispersal among geologically distinct islands and island banks is relatively infrequent in anoles and has contributed to allopatric speciation. Finally, our results suggest that large Greater Antillean islands serve as centres of origin for regional species diversity.     

Helminth parasites in Chaunus marinus and Cranopis valliceps (Anura: Bufonidae) from Lagunas Yalahau, Yucatan, Mexico

Eight helminth taxa were found parasitizing Chaunus marinus (n = 40) and Cranopsis valliceps (n = 40) from the Parque Estatal Lagunas Yalahau, Yucatan, Mexico. Seven taxa (2 digeneans: Langeronia macrocirra, Mesocoelium monas; 1 acanthocephalan: Oncicola sp.; 3 nematodes: Rhabdias füleborni, Aplectana itzocanensis, Cruzia morleyi; and a nematode larva) were found in C. marinus, while 4 taxa (all nematodes: Rhabdias fuelleborni, Aplectana itzocanensis, Ozwaldocruzia sp., and a nematode larva) were present in C. valliceps. Nematodes, particularly A. iztocanensis, showed high prevalence, mean abundance, and mean intensity values for both species of amphibians. The occurrence of R. fuelleborni, M. monas, L. macrocirra, and C. morleyi in these amphibians from the Yucatan Peninsula confirms their neotropical distribution, while the presence of A. itzocanensis increases its geographical distribution, suggesting a preference by neotropical, rather than neartic areas.     

Lack of convergence in aquatic Anolis lizards

Why convergent evolution occurs among some species occupying similar habitats but not among others is a question that has received surprisingly little attention. Caribbean Anolis lizards, known for their extensive convergent evolution among islands in the Greater Antilles, are an appropriate group with which to address this question. Despite the well-documented pattern of between-island convergence, some Greater Antillean anoles are not obviously part of the convergence syndrome. One example involves aquatic anoles--species that are found near to and readily enter streams-which have evolved independently twice in the Caribbean and also twice on mainland Central America. Despite being found in similar habitats, no previous study has investigated whether aquatic anoles represent yet another case of morphological convergence. We tested this hypothesis by collecting morphological data for seven aquatic anole species and 29 species from the six convergent types of Greater Antillean habitat specialists. We failed to find evidence for morphological convergence: the two Caribbean aquatic species are greatly dissimilar to each other and to the Central American species, which, however, may be convergent upon each other. We suggest two possible reasons for this lack of convergence in an otherwise highly convergent system: either there is more than one habitat type occupied by anoles in the proximity of water, or there is more than one way to adapt to a single aquatic habitat. We estimate that almost all of the 113 species of Greater Antillean anoles occupy habitats that are also used by distantly related species, but only 15% of these species are not morphologically similar to their distantly related ecological counterparts. Comparative data from other taxa would help enlighten the question of why the extent of convergence is so great in some lineages and not in others.     

New species of Allopharynx (Digenea: Plagiorchiidae) and other helminths in Uranoscodon superciliosus (Squamata: Tropiduridae) from Amazonian Brazil

Allopharynx daileyi n. sp. (Digenea: Plagiorchiidae) from the small intestine of the tropidurid lizard Uranoscodon superciliosus from Amazonian Brazil is described and illustrated. Of the 11 currently recognized species of Allopharynx, we consider Allopharynx megorchis Simha, 1961 a synonym of Allopharynx mehrai (Gogate, 1935) Price 1938 and Allopharynx puertoricensis Acholonu, 1976 a synonym of Allopharynx riopedrensis Garcia-Diaz; thus, A. daileyi becomes the 10th species assigned to the genus. The trematode species Mesocoelium monas and 2 species of Nematoda, Africana chabaudi and Cosmocerca vrcibradici, were also present.     

Molecular data reveals California as the potential source of an invasive leafhopper species, Macrosteles sp. nr. severini, transmitting the aster yellows phytoplasma in Hawaii

A species of aster leafhopper ( Macrosteles sp.) became established in 2001 on Oahu, Hawaii, and through the transmission of the aster yellows phytoplasma, caused devastating losses to the island's watercress industry. DNA sequence data were analysed from two mitochondrial genes [cytochrome oxidase subunit 1( CO1 ) and nicotinamide adenine dinucleotide 1 ( NADH1 )] and one nuclear gene (wingless, Wg ) (combined total of 1874 bp) to reconstruct phylogenetic relationships between putative US mainland source populations of aster leafhoppers and those introduced to Hawaii. These data were applied to elucidate the origin(s) and identity of Hawaiian infestations and the amount of genetic diversity within introduced invasive populations. Both phylogenetic search criteria (Bayesian and maximum likelihood models) converged onto similar tree topologies for all three gene regions and suggested that Hawaii infestations represent a single undescribed leafhopper species unrelated to the common aster leafhopper, Macrosteles quadrilineatus . An exact haplotype match was found from a specimen intercepted from watercress shipped to Hawaii from Los Angeles, California, suggesting this region as the potential source for Hawaiian infestations. Two mitochondrial haplotypes were identified in Hawaii suggesting two or perhaps just a single introduction of more than one female.     

A contribution to the Dicyrtomidae (Collembola) of Hawaii

Ten new species in three subgenera of Dicyrtoma are described from the Hawaiian Islands. Specimens were received from collections made on Hawaii, Maui, Kauai and Oahu. Species definitions are based on chaetotaxy of head, legs and the circumanal region. In addition, presence or absence of lateral spines (neosminthurid setae) on the parafurcular lobes may assist in grouping species within subgenera. Previous records of dicyrtomids from Hawaii include only one species, Ptenothrix ( Papaairioides ) daubia (Folsom). The following new species are described: Ptenothrix ( Ptenothrix ) hawaiicnaeis sp.n., Ptenothrix ( Papirioides ) kauaiensis sp.n., P. ( Papirioides ) serrata sp.n., Dicyrtoma ( Calvatomina ) sylvestratilis sp.n., D. ( Calvatomina ) brevifibra sp.n., D. ( Calvatomina ) tesselata sp.n., D. ( Calvatomina ) longidigita sp.n., D. ( Calvatomina ) bellingeri sp.n., D. ( Calvatomina ) madestris sp.n., and D. ( Calvatomina ) microdentata sp.n.     

Exploiting genomic resources in studies of speciation and adaptive radiation of lizards in the genus Anolis

Lizards in the genus Anolis have radiated extensively within and among islands in the Caribbean. Here, I provide a prospectus for identifying genes underlying adaptive phenotypic traits in anoles. First I review patterns of diversification in Anolis and the important morphological axes along which divergence occurs. Then I discuss two features of anole diversification, the repeated, convergent evolution of ecomorphs, and phenotypic divergence among populations within species, that provide opportunities to identify genes underlying adaptive phenotypic variation. While small clutch size and difficulty with captive rearing currently limit the utility of quantitative trait locus analyses, comparative analyses of gene expression, and population genomic approaches are promising.     

Evolutionary consequences of ecological release in Caribbean Anolis lizards

On the large islands of the Greater Antilles, multi-species communities of Anolis lizards are composed of species specialized to use particular habitats; similar sets of specialized species have evolved independently on each island. We studied species of anoles found on small Caribbean islands. Because these islands contain at most only one other species of anole, we predicted that species on these islands should not be as specialized as Greater Antillean species; rather, they might be expected to exhibit a generalized morphology and a greater breadth of habitat use. Our findings, however, do not confirm these predictions. Lesser Antillean species do not exhibit greater breadth of habitat use than Greater Antillean species, nor do they exhibit a generalized morphology. Most species are ecologically and morphologically similar to specialized trunk-crown anoles of the Greater Antilles, although some species exhibit morphologies unlike those seen in Greater Antillean species. Among descendants of specialized Greater Antillean species occurring on one-or two-species islands, most descendants of trunk-crown species have diverged relatively little, whereas several descendants of trunk-ground anoles have diverged considerably. Consequently, we propose that ancestral species in the Greater Antilles may have been trunk-crown anoles.