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Helminths of 12 Species of Anolis Lizards (Polychrotidae) from the Lesser Antilles, West Indies

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J. Helminthol. Soc. Wash. 64(2), 1997 pp. 248-257 Helminths of 12 Species of Anolis Lizards (Polychrotidae) from the Lesser Antilles, West Indies STEPHEN R. GOLDBERG,1-3 CHARLES R. BuRSEY,2 AND HAY CHEAM! 1 Department of Biology, Whittier College, Whittier, California 90608 (e-mail: [email protected]) and 2 Department of Biology, Pennsylvania State University, Shenango Campus, Sharon, Pennsylvania 16146 (e-mail: [email protected]) ABSTRACT: Twelve species of anoles (Anolis aeneus, A. extremus, A. gingivinus, A. griseus, A. luciae, A. mar- moratus, A. oculatus, A. richardi, A. roquet, A. sabanus, A. trinitatis, and A. wattsi) from the Lesser Antilles were examined for helminths. Twelve species of helminths were found: Mesocoelium monas, Oochoristica maccoyi, Oswaldocruzia marechali, Parapharyngodon cubensis, Spauligodon caymanensis, Trichospirura teix- eirai, Abbreviata sp., Ascarops sp., Physaloptera sp., Physocephalus sp., Porrocaecum sp., and Centrorhynchus sp. Twenty-nine new host records are reported. The highest prevalence (75%) was P. cubensis in A. sabanus; greatest mean intensity (56.8) was S. caymanensis in A. marmoratus. The highest diversity of helminths was found in Anolis gingivinus, which harbored 9 species; the lowest diversity occurred in A. trinitatis, which harbored 1 species. Islands with the greatest numbers of helminth species are located in the northern Lesser Antilles. KEY WORDS: Anolis, Polychrotidae, Acanthocephala, Cestoda, Nematoda, Trematoda, Lesser Antilles. The genus Anolis Daudin contains some 300 A. aeneus in Trinidad and Guyana; A. bimacu- species that inhabit Central America, southern latus in Bermuda; A. extremus in St. Lucia, Ber- U.S.A. northern South America, and the Carib- muda, and Caracas, Venezuela; A. roquet in Ber- bean islands (Roughgarden, 1995). About 130 muda; and A. trinitatis in Trinidad (Schwartz species are known from the West Indies alone and Henderson, 1991). Although anoles of the (Schwartz and Henderson, 1991). The isolation Lesser Antilles have been studied extensively necessary for such diversity to evolve was pro- (see Roughgarden, 1995), there are reports of vided by the splitting of ancestral stock into helminths from only A. bimaculatus, A. ferreus, some 3,000 separate populations at the end of A. gingivinus, A. lividus, A. marmoratus, A. ocu- the last Ice Age (Roughgarden, 1995). The larg- latus, A. sabanus, and A. wattsi (Dobson et al., er islands of the Greater Antilles have more spe- 1992; Ben Slimane et al., 1995; Goldberg and cies of Anolis; there are over 35 in Cuba, over Bursey, 1996; Goldberg et al., 1996a). The pur- 35 in Hispaniola, 11 in Puerto Rico, and 7 in pose of this paper is to report helminths from Jamaica; small islands, as typified by the Lesser Anolis aeneus, A. extremus, A. gingivinus. A. Antilles, have 1 or 2 species (Roughgarden, griseus, A. luciae, A. marmoratus, A. oculatus, 1995). A. richardi, A. roquet, A. sabanus, A. trinitatis, Sixteen species of anoles occur on the islands and A. wattsi and to compare helminth infections of the Lesser Antilles, namely, Anolis aeneus among anoles of the Lesser Antilles. Gray, 1840, A. bimaculatus Sparrman, 1874, A. extremus Garman, 1888, A. ferreus Cope, 1864, Materials and Methods A. gingivinus Cope, 1864, A. griseus Garman, All anoles examined in this study (N = 261) were 1888, A. lividus Garman, 1888, A. luciae Gar- museum specimens (for museum accession numbers, man, 1888, A. marmoratus Dumeril and Bibron, see Appendix 1). Anole species, collection location, 1837, A. nubilis Garman, 1888, A. oculatus sample size, and mean snout-vent length in millime- ters are given in Table 1. The islands of the Lesser Cope, 1879, A. richardi Dumeril and Bibron, Antilles labeled with the species of Anolis we exam- 1837, A. roquet Lacepede, 1788, A. sabanus ined for helminths are shown in Figure 1. The body Garman, 1887, A. trinitatis Reinhardt and Liit- cavity was opened by a longitudinal incision from vent ken, 1863, and A. wattsi Boulenger, 1894. Five to throat, and the digestive tract was removed by cut- of these have been introduced to other regions: ting across the anterior esophagus and rectum. The esophagus, stomach, and small and large intestines were slit longitudinally and examined under a dissect- 3 Corresponding author. ing microscope. The gallbladder, liver, and body cavity 248 Copyright © 2011, The Helminthological Society of Washington GOLDBERG ET AL.—ANOLIS LIZARDS 249 Table 1. Samples of Anolis lizards examined from the Caribbean. x snout-vent length ± SD Range Species Locality N (mm) (mm) Anolis aeneus Grenada 20 62.2 ± 3.7 54-68 Anolis extrernus Barbados 5 64.0 ± 8.0 53-70 St. Lucia 5 61.4 ± 3.4 57-66 Anolis gingivinus Anguilla 64 50.6 ± 6.8 39-63 St. Barthelemy 14 58.4 ± 4.9 43-64 Anolis griseus St. Vincent 10 91.7 ± 13.1 74-107 Anolis luciae St. Lucia 34 64.5 ± 12.3 31-82 At olis marmoratits Guadeloupe 25 60.8 ± 4.6 48-68 At olis oculatus Dominica 11 69.1 ± 9.7 52-80 At olis richardi Grenada 20 91.5 ± 11.7 66-110 At olis roquet Martinique 4 58.0 ± 7.2 51-68 At olis sabanus Saba 12 53.0 ± 4.8 45-60 At olis trinitatis St. Vincent 17 52.4 ± 8.8 37-68 At olis watt si Antigua 20 41.3 ± 3.4 36-47 were also searched for helminths. Each helminth was land of Redonda (ca. 19 km NW of Montserrat), initially placed in a drop of glycerol on a glass slide. is unlisted and remains to be examined. For Nematodes were identified from these temporary mounts. Cestodes were stained with hematoxylin, these anoles, known helminths consist of 2 spe- mounted in balsam, and identified. Acanthocephalans cies of trematodes, Alloglyptus crenshawi Byrd, were cleared in xylene, mounted in balsam, and as- 1950, and Mesocoelium monas; 1 species of ces- signed to a genus. tode, Oochoristica maccoyi; 14 species of nem- Results atodes, Abbreviata sp., Ascarops sp., Physalop- tera sp., Physocephalus sp., Porrocaecum sp. The helminths found in the 12 anole species represented by larvae only, Oswaldocruzia dor- examined in this study consisted of 1 species of sarmata Ben Slimane, Durette-Desset, and Cha- trematode, Mesocoelium monas (Rudolphi, baud, 1995, O. jeanbarti Ben Slimane, Durette- 1819); 1 species of cestode, Oochoristica mac- Desset and Chabaud, 1995, O. marechali, O. coyi Bursey and Goldberg, 1996; 9 species of mauleoni Ben Slimane, Durette-Desset and Cha- nematodes, Oswaldocruzia marechali Ben Sli- baud, 1995, Parapharyngodon cubensis, Rhab- mane, Durette-Desset, and Chabaud, 1995, Para- dias sp., Spauligodon caymanensis, Spinicauda pharyngodon cubensis (Barus and Coy Otero, spinicauda (Olfers, 1819), and Trichospirura 1969), Spauligodon caymanensis Bursey and teixeirai; and 1 species of acanthocephalan, Goldberg, 1995, Trichospirura teixeirai (Barus Centrorhynchus sp., represented by cystacanths and Coy Otero, 1968), Abbreviata sp., Ascarops sp., Physaloptera sp., Physocephalus sp., and only. With the exception of Oochoristica mac- Porrocaecum sp., and 1 species of acanthoceph- coyi, Oswaldocruzia dorsarmata, O. jeanbarti, alan, Centrorhynchus sp. The first 6 species O. marechali, and O. mauleoni, all of these hel- were represented by mature individuals; only minths have been reported from other amphibian immature individuals of the latter 6 species were or reptilian host species, some of which are out- found. Occurrences of helminths and 29 new side the Lesser Antilles (Baker, 1987). host records are listed in Table 2. Helminths The life cycles of Alloglyptus crenshawi and were placed in vials of 70% ethanol and depos- Mesocoelium monas have not yet been investi- ited in the United States National Parasite Col- gated, but they are the only trematodes so far lection [USDA], Beltsville, Maryland (Appen- reported from anoles of the Lesser Antilles and dix 2). could be expected to exhibit typical life cycles. Alloglyptus crenshawi was described by Byrd Discussion (1950) from 30 specimens found in the small The known helminth fauna for anoles of the intestine of 1 Anolis carolinensis collected in Lesser Antilles is presented in Table 3. Only Georgia. It has been found in 3 Anolis caroli- Anolis nubilis, which is restricted to the tiny is- nensis collected in Putnam County, Florida Copyright © 2011, The Helminthological Society of Washington 250 JOURNAL OF THE HELMINTHOLOGICAL SOCIETY OF WASHINGTON, 64(2), JULY 1997 zju j^urUN/\ v_/r inn nc.jL.wiiN i i~n_n^v_«ji^rtL ov^v^io i i v^r vvrtomrvvj i VJIN, ot(z), JULI iyy/ I I 63 60C ANGUILLA _1g A.gingivinus 18°- ST. MARTIN o ST BARTHELEMY SABA A. gingivinus A. sabanus o ^ BARBUDA ST. EUSTATIUS^ ST. CHRISTOPHER MONTSERRAT 0 GUADELOUPE A. marmoratus \) ^ .. OMARIE-GALANTE DOMINICA A. oculatus 1-15 15°- MARTINIQUE ST. LUCIA LESSER ANTILLES A. extremus 0 50 100 I i i « ST. VINCENT BARBADOS (_/ A gr/seus ?\ extremus A. trinitatis (-J GRENADA -12C /4. aeneus A. richardi 63 I Figure 1. Islands of the Lesser Antilles with species of Anolis labeled next to the island from which they were collected. All other named islands are sites with published records of helminths from Anolis spp. Copyright © 2011, The Helminthological Society of Washington GOLDBERG ET AL.—ANOLIS LIZARDS 251 Table 2. Prevalence and mean intensity of helminths found in Anolis spp. examined in this study. Mean inten- Species Helminth Prevalence sity Range Site Anolis aeneus Parapharyngodon cubensis* 25% (5/20) 1.2 1-2 Large intestine Physocephalus sp. (larvae)* 5% (1/20) 2.0 — Encysted stomach wall Centrorhynchus sp. (cystacanths)* 15% (3/20) 1.0 — Coelom Anolis extremus Ascarops sp. (larvae)* 30% (3/10) 6.7 1-12 Encysted stomach wall Parapharyngodon cubensis* 20% (2/10) 4.0 2-6 Large intestine Trichospirura teixeirai* 20% (2/10) 5.5 3-8 Gallbladder Anolis gingivinus Mesocoelium monas l%(l/78) 1.0 — Small intestine Oochoristica maccoyi 8% (6/78) 1.5 1-3 Small intestine Abbreviate* sp.
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    Climate Change and Collapsing Thermal Niches of Mexican Endemic Reptiles White Paper for the Environmental Working Group of the UC-Mexico Initiative Barry Sinervo1, Rafael A. Lara Reséndiz1, 2, Donald B. Miles3, Jeffrey E. Lovich4, Joshua R. Ennen5, Johannes Müller6, Robert D. Cooper1, 7, Philip C. Rosen8, Joseph A. E. Stewart1, Juan Carlos Santos9, Jack W. Sites Jr.9, Paul M. Gibbons10, Eric V. Goode10, L. Scott Hillard7, 11, Luke Welton9, 12, Mickey Agha4, 13, Gabriel Caetano1, 14, Mercy Vaughn15, Cristina Meléndez Torres16, Héctor Gadsden17, Gamaliel Casteñada Gaytán18, Patricia Galina Tessaro19, Fernando I. Valle Jiménez19, Jorge Valdez Villavicencio20, Norberto Martínez Méndez21, Guillermo Woolrich Piña22, Víctor Luja Molina23, Aníbal Díaz de la Vega Pérez24, Diego M. Arenas Moreno2, Saúl Domínguez Guerrero2, Natalia Fierro2, Scott Butterfield25, Michael Westpha26, Raymond B. Huey27, William Mautz28, Víctor Sánchez Cordero29, and Fausto R. Méndez de la Cruz29 1. The Institute for the Study of the Ecological and Evolutionary Climate Impacts, University of California Santa Cruz 2. Laboratorio de Herpetología, Instituto de Biología, Universidad Nacional Autónoma de México 3. Department of Biological Sciences, Ohio University 4. USGS, Southwest Biological Science Center, 5. Tennessee Aquarium Conservation Institute 6. Museum für Naturkunde, Leibniz-Institut für Evolutions 7. Department of Ecology and Evolutionary Biology, University of California, Los Angeles 8. School of Natural Resources & the Environment, University of Arizona 9. Department of Biology, Brigham Young University 10. Turtle Conservancy 11. Turner Endangered Species Fund 12. University of Kansas Biodiversity Institute 13. Department of Wildlife, Fish, and Conservation Biology, University of California, Davis 14. Departamento de Zoologia, Instituto de Ciências Biológicas, Universidade de Brasília 15.
  • Lizards As Model Organisms for Linking Phylogeographic and Speciation Studies

    Lizards As Model Organisms for Linking Phylogeographic and Speciation Studies

    Molecular Ecology (2010) 19, 3250–3270 doi: 10.1111/j.1365-294X.2010.04722.x INVITED REVIEW Lizards as model organisms for linking phylogeographic and speciation studies ARLEY CAMARGO,* BARRY SINERVO† and JACK W. SITES JR.* *Department of Biology and Bean Life Science Museum, Brigham Young University, Provo, UT 84602, USA, †Department of Ecology and Evolutionary Biology, University of California, Santa Cruz, CA 95064, USA Abstract Lizards have been model organisms for ecological and evolutionary studies from individual to community levels at multiple spatial and temporal scales. Here we highlight lizards as models for phylogeographic studies, review the published popula- tion genetics ⁄ phylogeography literature to summarize general patterns and trends and describe some studies that have contributed to conceptual advances. Our review includes 426 references and 452 case studies: this literature reflects a general trend of exponential growth associated with the theoretical and empirical expansions of the discipline. We describe recent lizard studies that have contributed to advances in understanding of several aspects of phylogeography, emphasize some linkages between phylogeography and speciation and suggest ways to expand phylogeographic studies to test alternative pattern-based modes of speciation. Allopatric speciation patterns can be tested by phylogeographic approaches if these are designed to discriminate among four alterna- tives based on the role of selection in driving divergence between populations, including: (i) passive divergence by genetic drift; (ii) adaptive divergence by natural selection (niche conservatism or ecological speciation); and (iii) socially-mediated speciation. Here we propose an expanded approach to compare patterns of variation in phylogeographic data sets that, when coupled with morphological and environmental data, can be used to discriminate among these alternative speciation patterns.