Phylogeny and Biogeography of Ratite Birds Inferred from DNA Sequences of the Mitochondrial Ribosomal Genes

Total Page:16

File Type:pdf, Size:1020Kb

Phylogeny and Biogeography of Ratite Birds Inferred from DNA Sequences of the Mitochondrial Ribosomal Genes Phylogeny and Biogeography of Ratite Birds Inferred from DNA Sequences of the Mitochondrial Ribosomal Genes Marcel van Tuinen,* Charles G. Sibley,² and S. Blair Hedges* *Department of Biology and Institute of Molecular Evolutionary Genetics, Pennsylvania State University; and ²Santa Rosa, California The origin of the ¯ightless ratite birds of the southern continents has been debated for over a century. Whether dispersal or vicariance (continental breakup) best explains their origin depends largely on their phylogenetic rela- tionships. No consensus has been reached on this issue despite many morphological and molecular studies. To address this question further we sequenced a 2.8-kb region of mitochondrial DNA containing the ribosomal genes in representative ratites and a tinamou. Phylogenetic analyses indicate that Struthio (Africa) is basal and Rhea (South America) clusters with living Australasian ratites. This phylogeny agrees with transferrin and DNA hybrid- ization studies but not with sequence analyses of some protein-coding genes. These results also require reevaluation of the phylogenetic position of the extinct moas of New Zealand. We propose a new hypothesis for the origin of ratites that combines elements of dispersal and vicariance. Introduction The living ratites include two species of ostriches cause a UPGMA tree joined Rhea and Struthio, whereas (Struthio) in Africa and formerly in Asia, the Australian trees constructed with the Fitch-Margoliash (1967) al- emu (Dromaius), three species of cassowaries (Casuar- gorithm joined Rhea with the Australasian clade (Sibley ius) in New Guinea and northeastern Australia, three and Ahlquist 1990). In contrast, mitochondrial DNA se- species of forest-dwelling kiwis (Apteryx) in New Zea- quence data (12S rRNA; 400 bp) supported a basal po- land, and two rheas (Rhea) in South America (Sibley sition for Rhea (Cooper et al. 1992). However, reanaly- 1996). All lack a keel on the sternum, a character as- sis of those sequence data by Rzhetsky, Kumar, and Nei sociated with ¯ightlessness. Based on anatomical anal- (1995) and analysis of an additional 600 bp from a nu- yses, ratite phylogeny has been controversial for over a clear gene (Cooper and Penny 1997) indicate that such century. This stems from subjective interpretations of small data sets are unlikely to resolve the early history anatomical characters and from dif®culties in determin- of ratites. Therefore, we sequenced the complete mito- ing polarity in characters shared by ratite birds (Kuroch- chondrial ribosomal genes (2.8 kb) from representative kin 1995). Sibley and Ahlquist (1990) provide a histor- ratite species to address the question of the nearest living ical review of ratite systematics. relative of the Australasian ratites. These genes have Over the last two decades, several molecular stud- proven useful for addressing higher-level phylogenetic ies of ratites have clari®ed some aspects of ratite phy- questions in birds and other vertebrates (e.g., Hedges logeny. There is general agreement that living ratites are 1994; Hedges and Sibley 1994; Springer et al. 1997). monophyletic and that the weakly ¯ying tinamous are their closest living relatives (Prager et al. 1976; Sibley Materials and Methods and Ahlquist 1981, 1990; Caspers, Wattel, and De Jong 1994). Monophyly of the living Australasian ratites is A 2.8-kb region of mitochondrial DNA was se- also supported by molecular data (Prager et al. 1976; quenced for each of three ratite species (Rhea ameri- Sibley and Ahlquist 1981; 1990; Cooper et al. 1992), cana, Struthio camelus, and Dromaius novaehollandiae) although consensus on this issue has not been reached and a gray tinamou (Tinamus tao). We assumed mono- with anatomical data (Cracraft 1974; Bledsoe 1988; Ku- phyly of the living Australasian ratites based on molec- rochkin 1995). ular evidence (Prager et al. 1976; Sibley and Ahlquist The relationships of the three major lineages, rheas, 1990; Cooper et al. 1992), with Dromaius as the rep- ostriches, and Australasian ratites, have a direct bearing resentative of the living Australasian ratite clade. The on the biogeographic history of the ratites (Cracraft sequenced region includes the entire 12S rRNA, t- 1974), but these relationships remain unclear. Analysis RNAVal, and 16S rRNA genes. The corresponding se- of transferrin immunological data grouped the Rhea quences from a domestic fowl (Gallus gallus; accession with the Australasian clade and thus identi®ed Struthio number X52392, sites 1274±3994 in Desjardins and as the most basal living ratite (Prager et al. 1976). The Morais 1990) and an American alligator (Alligator mis- relationships of these three lineages with DNA-DNA hy- sissippiensis, accession number L28074) were obtained bridization data were considered to be unresolved be- from GenBank for comparison. The new sequences re- ported here have been deposited in EMBL with acces- Key words: molecular phylogeny, ratite evolution, vicariance, dis- sion numbers AJ002921±AJ002924. persal, Gondwana. DNA ampli®cation was performed with the follow- Address for correspondence and reprints: S. Blair Hedges, De- ing primer pairs:12L9/12H2, 12L9/12H5, 12L10/12H5, partment of Biology, 208 Mueller Laboratory, Pennsylvania State Uni- 12L1/12H4, 12L7/16H11, 16L11/16H5, 16L17/16H17, versity, University Park, Pennsylvania 16802. E-mail: [email protected]. 16L10/16H10, 16L9/16H3, 16L8/16H13, and 16L4/ Mol. Biol. Evol. 15(4):370±376. 1998 16H12 (Hedges 1994; Hedges and Sibley 1994; Hedges q 1998 by the Society for Molecular Biology and Evolution. ISSN: 0737-4038 et al. 1995). Primers not previously described are 370 Phylogeny of Ratites 371 [IUPAC]: 12L7 (GAA GGW GGA TTT AGY AGT AAA), 12L9 (AAA GCA HRR CAC TGA ARA TGY YDA GA), 12L10 (CMC AMG GGA MWC AGC AGT GAT WAA HAT T), 12H5 (TTA GAG GAG CCT GTC CTA TAA TCG), 16L17 (CCW AMC GAR CYT RGT GAT AGC TGG TT), and 16H17 (TGT TTA CCA AAA ACA TMY CCY YYS GC). DNA was PCR-ampli®ed as follows: 30±40 cycles (948C for 15 s, 508Cor558C for 15 s, 728C for 45 s). Double-stranded DNA was gel-puri®ed and served as template for another PCR run under slightly different conditions: 25±35 cycles (948C for 15 s, 558Cor608C for 15 s, 728C for 45 s). A hot start was performed at 808C for all PCR runs. Prior to sequencing, double- FIG. 1.ÐNeighbor-joining tree of three representative ratite birds stranded DNA was ®ltered as described earlier (Hedges (Struthio camelus, Rhea americana, and Dromaius novaehollandiae), and Sibley 1994). Both complementary L- and H- a tinamou (Tinamus tao), domestic fowl (Gallus gallus), and American strands were sequenced for all primer sets. Cycle se- alligator (Alligator mississipiensis) inferred from the mitochondrial 12S, 16S rRNA, and tRNAVal genes (2.8 kb). Con®dence values are quencing reactions were performed using 39 dye-labeled indicated between brackets on the nodes in the tree (left, interior- dideoxynucleotide triphosphates (¯uorescent dye termi- branch test; middle, interior-branch test for transversions only; right, nators) and run on an ABI PRISM 377 DNA Sequencer maximum-parsimony bootstrap values). (Perkin-Elmer ABI, Foster City, Calif.). Alignments were performed with ESEE (Cabot and Beckenbach 1989). For phylogenetic analysis, neighbor- accordance with the number of varied sites and those joining (Saitou and Nei 1987) analyses were performed informative for parsimony for the three genes (16S: 705 in MEGA (Kumar, Tamura, and Nei 1994) using a Ki- varied sites, 262 sites informative for parsimony; 12S: mura (1980) two-parameter distance with transitions and 440 varied sites, 135 sites informative for parsimony; transversions included unless otherwise noted. Maxi- tRNAVal: 28 varied sites, 12 sites informative for par- mum-parsimony analyses were conducted with PAUP simony). The 12S rRNA and tRNAVal data support the (Swofford 1993), and maximum-likelihood analyses Rhea-Dromaius clade, with 71% and 70% con®dence were performed with Nucml (HKY) in MOLPHY (Ada- values, respectively, for the interior-branch analysis. chi and Hasegawa 1996). In MOLPHY, we used the de- After this study was completed, a similar study by fault transition : transversion ratio of 4:1, which is sim- Lee, Feinstein, and Cracraft (1997) was published on ilar to that estimated for mitochondrial DNA (Kumar ratite relationships inferred from DNA sequences of sev- 1996), and 100:1 (ùtransversions only). Sites corre- eral noncoding and protein-coding mitochondrial genes, sponding to alignment gaps were excluded. The interior- including a reexamination of the morphological evi- branch (SE) test was performed to assess con®dence in dence. There was partial overlap between the sequenced the reliability of the neighbor-joining trees by testing if regions (rRNA genes) in their study and ours. Compar- interior branch lengths deviate signi®cantly from zero. ison of our Struthio sequence with their sequence and In PAUP, the bootstrap method was applied (Felsenstein that of HaÈrlid, Janke, and Arnason (1997) indicated 11± 1985) with 2,000 replications. 12 nucleotide differences between each of the three se- quences. All differences between our sequence and that Results of HaÈrlid, Janke, and Arnason (1997) were transitions, whereas both of those sequences shared six transversion Out of 2,900 sites in the initial alignment of six differences with the Struthio sequence of Lee, Feinstein, taxa, 2,429 were analyzed after elimination of gaps and and Cracraft (1997). These differences did not affect the sites showing ambiguous alignment. Of those sites, there phylogeny as shown in ®gure 1. However, the sequence were 1,173 varied sites and 409 sites informative for the analysis (combined genes) of Lee, Feinstein, and Cra- parsimony method. The phylogenetic tree (®g. 1), rooted craft (1997) supported a basal Rhea (rather than Stru- with Alligator, shows high con®dence for a Rhea-Dro- thio) lineage, an unexpected discordance that we inves- maius clade. Con®dence values are signi®cant (99%) for tigated here. the interior-branch test and almost signi®cant when only An overview of the present molecular evidence (ta- transversions are used (92%).
Recommended publications
  • Download Vol. 11, No. 3
    BULLETIN OF THE FLORIDA STATE MUSEUM BIOLOGICAL SCIENCES Volume 11 Number 3 CATALOGUE OF FOSSIL BIRDS: Part 3 (Ralliformes, Ichthyornithiformes, Charadriiformes) Pierce Brodkorb M,4 * . /853 0 UNIVERSITY OF FLORIDA Gainesville 1967 Numbers of the BULLETIN OF THE FLORIDA STATE MUSEUM are pub- lished at irregular intervals. Volumes contain about 800 pages and are not nec- essarily completed in any one calendar year. WALTER AuFFENBERC, Managing Editor OLIVER L. AUSTIN, JA, Editor Consultants for this issue. ~ HILDEGARDE HOWARD ALExANDER WErMORE Communications concerning purchase or exchange of the publication and all manuscripts should be addressed to the Managing Editor of the Bulletin, Florida State Museum, Seagle Building, Gainesville, Florida. 82601 Published June 12, 1967 Price for this issue $2.20 CATALOGUE OF FOSSIL BIRDS: Part 3 ( Ralliformes, Ichthyornithiformes, Charadriiformes) PIERCE BRODKORBl SYNOPSIS: The third installment of the Catalogue of Fossil Birds treats 84 families comprising the orders Ralliformes, Ichthyornithiformes, and Charadriiformes. The species included in this section number 866, of which 215 are paleospecies and 151 are neospecies. With the addenda of 14 paleospecies, the three parts now published treat 1,236 spDcies, of which 771 are paleospecies and 465 are living or recently extinct. The nominal order- Diatrymiformes is reduced in rank to a suborder of the Ralliformes, and several generally recognized families are reduced to subfamily status. These include Geranoididae and Eogruidae (to Gruidae); Bfontornithidae
    [Show full text]
  • Ratite Molecular Evolution, Phylogeny and Biogeography Inferred from Complete Mitochondrial Genomes
    RATITE MOLECULAR EVOLUTION, PHYLOGENY AND BIOGEOGRAPHY INFERRED FROM COMPLETE MITOCHONDRIAL GENOMES by Oliver Haddrath A thesis submitted in confonnity with the requirements for the Degree of Masters of Science Graduate Department of Zoology University of Toronto O Copyright by Oliver Haddrath 2000 National Library Biblioth&que nationale 191 .,,da du Canada uisitions and Acquisitions et Services services bibliographiques 395 Welington Street 395. rue WdKngton Ottawa ON KIA ON4 Otîâwâ ON K1A ûN4 Canada Canada The author has granted a non- L'auteur a accordé une iicence non exclusive licence allowing the exclusive permettant A la National Library of Canada to Bihliotheque nationale du Canada de reproduce, loan, distribute or sell reproduire, @ter, distribuer ou copies of diis thesis in microfonn, vendre des copies de cette thèse sous paper or electronic formats. la forme de microfiche/fïîm, de reproduction sur papier ou sur format 61ectronique. The author retains ownership of the L'auteur conserve la propriété du copyright in this thesis. Neither the droit d'auteur qui protège cette tbése. thesis nor substantial exûacts fiom it Ni la thèse ni des extraits substantiels may be priated or otherwise de celle-ci ne doivent être imprimés reproduced without the author's ou autrement reproduits sans son permission. autorisation. Abstract Ratite Molecular Evolution, Phylogeny and Biogeography Inferred fiom Complete Mitochoncîrial Genomes. Masters of Science. 2000. Oliver Haddrath Department of Zoology, University of Toronto. The relationships within the ratite birds and their biogeographic history has been debated for over a century. While the monophyly of the ratites has been established, consensus on the branching pattern within the ratite tree has not yet been reached.
    [Show full text]
  • High Metabolic Rates in Running Birds
    scientific correspondence humidity and climate variations related to 9. Minnis, P., Ayers, J. K. & Weaver, S. P. Surface-Based Observations 40 11,12 of Contrail Occurrence Frequency over the U. S., April 1983–April the North Atlantic oscillation , showed Rhea that none of them, taken individually, is a 1994 (NASA Reference Publication 1404, 1997). Wolf 10.Jensen, E. J. & Toon, O. B. Geophys. Res. Lett. 19, 1759–1762 (1992). Coyote 30 good explanation for the observed positive 11.Hurrell, J. W. Science 269, 676–679 (1995). Pony trend in cirrus occurrence and its regional 12.Mächel, H., Kapala, A. & Flohn, H. Int. J. Climatol. 18, 1–22 (1998). Fox distribution. 13.Hartmann, D. L., Ockert-Bell, M. E. & Michelsen, M. L. J. Clim. Budgerigar d 5, 1281–1304 (1992). t 20 Ostrich s Raven · Humming- 14.Warren, S. G., Hahn, C. J., London, J., Chervin, R. M. & Jenne, E The trends in cirrus ‘amount when pre- / o bird R. L. Global Distribution of Total Cloud Cover and Cloud Type c o Emu sent’ over the previously defined regions of l · Pigeon E Turkey Amounts over the Ocean (NCAR Technical Note TN-317 + STR, 10 Most high fuel consumption are 11.9% and Boulder, Colorado, 1988). mammals Penguin 14.2% for land and ocean, respectively 15.Hahn, C. J., Warren, S. G. & London, J. J. Clim. 8, 1429–1446 Stork Penguin (1995). (Table 1). The combination of a large posi- 0 Supplementary information is available on Nature’s World-Wide 0.001 0.01 0.1 1 10 100 1,000 tive trend in cirrus occurrence associated Web site (http://www.nature.com) or as paper copy from the with a negative trend in the cirrus amount London editorial office of Nature.
    [Show full text]
  • Biogeography of the Llanos De Moxos Roberto Langstroth Plotkin 183
    MF Geographica Helvetica Jg. 66 2011/Heft 3 Biogeography of the Llanos de Moxos Roberto Langstroth Plotkin 183 Biogeography of the Llanos de Moxos: natural and anthropogenic determinants Roberto Langstroth Plotkin, South Riding Bactris, Ceiba, Coccoloba, Ficus, Genipa, Guarea, Hura, Inga, Maclura, Margaritaria, Salacia, Spondias, Sterculia, Swartzia, Syagrus, Tabebuia, Trichilia, Tripla- 1 Introduction ris, and Vitex (Beck 1983; Langstroth 1996). Prior to the arrival of Europeans in the Americas, the Semialturas are levee backslopes and splays with human inhabitants of the Llanos de Moxos constructed brief, shallow inundations and vegetation contingent diverse earthworks such as mounds and causeways, upon the fire regimes. Semialturas may support largely raised agricultural fields in the savannas and managed deciduous forest or woodland (genera such as Acroco- the landscape using fire and other tools (Denevan mia, Astronium, Coccoloba, Copernicia, Cordia, Cupa- 1966; Langstroth 1996; Lombardo & Prümers 2010; nia, Enterolobium, Geoffroea, Guazuma, Piptadenia, Lombardo et al. 2011). Erickson (2008) considers the Pithecellobium, Randia, Samanea, Sterculia, Tabebuia, Llanos de Moxos to be an example of an Amazonian and Zanthoxylum), Cerrado («campo cerrado» or «domesticated landscape» and, based on evidence «campo sujo», genera listed below), or pampa with from Moxos, claims that «nature in Amazonia more scattered fire tolerant trees Pseudobombax,( Tabe- closely resembles a garden than a pristine, natural buia) and Copernicia palms (Beck 1983; Langstroth wilderness.» These arguments presume that Moxos 1996). Termite mounds are frequent and present small is representative of Amazonia and also discount the woody islands with Celtis, Cereus, Coccoloba, Coper- roles of longer-term physical and biological processes nicia, Cordia, Machaerium, Rhamnidium, and Sorocea in play since the Miocene when extensive non-forest (Beck 1983; Langstroth 1996).
    [Show full text]
  • Aquila 23. Évf. 1916
    A madarak palaeontologiájának története és irodalma. Irta : DR. Lambrecht Kálmán. Minden ismeret történetének eredete többé-kevésbbé homályba vész. Az els úttörk még maguk is csak tapogatóznak; leírásaik — a kezdet nehézségeivel küzdve — nem szabatosak, több bennük a sej- dít, mint a positiv elem. Fokozottan áll ez a palaeontologiára, amely- nek gyakran bizony igen hiányos anyaga gazdag recens összehasonlító anyagot és alapos morphologiai ismereteket igényel. A palaeontologia legismertebb történetíróinak, MARSH-nak^ és ZiTTEL-nek2 chronologiai beosztásait figyelmen kívül hagyva, ehelyütt Abel3 szellemes beosztását fogadjuk el és megkülönböztetünk a madár- palaeontologia történetében 1. phantasticus, 2. descriptiv és 3. morpho- logiai és phylogenetikai periódust. Nagyon természetes, hogy a fossilis madarak ismerete karöltve haladt a recens madarak osteologiájának megismerésével, 4 mert a palaeon- tologus csakis recens comparativ anyag és vizsgálatok alapján foghat munkához. De viszont igaz az is, hogy a morphologus sem mozdulhat meg az si alakok vázrendszerének ismerete nélkül, nem is szólva arról, hogy a gyakran nagyon töredékes fossilis maradványok mennyi érdekes morphologiai megfigyelésre vezették már a búvárokat. A phantasticus periódus. Ez a periódus, amely — összehasonlítás hiján — túlnyomóan speculativ alapon mvelte a tudományt, a XVIll. századdal, vagyis CuviER felléptével végzdik. Eltekintve Albertus MAGNUS-nak (1193—1280, Marsh szerint 1 Marsh, 0. C, Geschichte und Methode der paläoiitologischen Entdeckungen. — Kosmos VI. 1879.
    [Show full text]
  • Convergent Regulatory Evolution and the Origin of Flightlessness in Palaeognathous Birds
    bioRxiv preprint doi: https://doi.org/10.1101/262584; this version posted February 8, 2018. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission. Convergent regulatory evolution and the origin of flightlessness in palaeognathous birds Timothy B. Sackton* (1,2), Phil Grayson (2,3), Alison Cloutier (2,3), Zhirui Hu (4), Jun S. Liu (4), Nicole E. Wheeler (5,6), Paul P. Gardner (5,7), Julia A. Clarke (8), Allan J. Baker (9,10), Michele Clamp (1), Scott V. Edwards* (2,3) Affiliations: 1) Informatics Group, Harvard University, Cambridge, USA 2) Department of Organismic and Evolutionary Biology, Harvard University, Cambridge, USA 3) Museum of Comparative Zoology, Harvard University, Cambridge, USA 4) Department of Statistics, Harvard University, Cambridge, USA 5) School of Biological Sciences, University of Canterbury, New Zealand 6) Wellcome Sanger Institute, Wellcome Genome Campus, Cambridge, UK 7) Department of Biochemistry, University of Otago, New Zealand 8) Jackson School of Geosciences, The University of Texas at Austin, Austin, USA 9) Department of Natural History, Royal Ontario Museum, Toronto, Canada 10) Department of Ecology and Evolutionary Biology, University of Toronto, Toronto, Canada *correspondence to: TBS ([email protected]) or SVE ([email protected]) bioRxiv preprint doi: https://doi.org/10.1101/262584; this version posted February 8, 2018. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission. The relative roles of regulatory and protein evolution in the origin and loss of convergent phenotypic traits is a core question in evolutionary biology.
    [Show full text]
  • (Aves: Palaeognathae) from the Paleocene (Tiffanian) of Southern California
    PaleoBios 31(1):1–7, May 13, 2014 A lithornithid (Aves: Palaeognathae) from the Paleocene (Tiffanian) of southern California THOMAS A. STIDHAM,1* DON LOFGREN,2 ANDREW A. FARKE,2 MICHAEL PAIK,3 and RACHEL CHOI3 1Key Laboratory of Vertebrate Evolution and Human Origins, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing 100044, China; e-mail: [email protected], corresponding author. 2Raymond M. Alf Museum of Paleontology, Claremont, California 91711, USA. 3 The Webb Schools, Claremont, California 91711, USA The proximal end of a bird humerus recovered from the Paleocene Goler Formation of southern California is the oldest Cenozoic record of this clade from the west coast of North America. The fossil is characterized by a relatively large, dorsally-positioned head of the humerus and a subcircular opening to the pneumotricipital fossa, consistent with the Lithornithidae among known North American Paleocene birds, and is similar in size to Lithornis celetius. This specimen from the Tiffanian NALMA extends the known geographic range of lithornithids outside of the Rocky Mountains region in the United States. The inferred coastal depositional environment of the Goler Formation is consistent with a broad ecological niche of lithornithids. The age and geographic distribution of lithornithids in North America and Europe suggests these birds dispersed from North America to Europe in the Paleocene or by the early Eocene. During the Paleogene the intercontinental dispersal of lithornithids likely occurred alongside other known bird and mammalian movements that were facilitated by climatic and sea level changes. Keywords: bird humerus, fossil, Lithornithidae, Goler Formation, Tiffanian, California INTRODUCTION largely unknown in North America.
    [Show full text]
  • Ancient DNA Suggests Dwarf and 'Giant' Emu Are Conspecific
    Ancient DNA Suggests Dwarf and ‘Giant’ Emu Are Conspecific Tim H. Heupink, Leon Huynen, David M. Lambert* Griffith School of Environment and School of Biomolecular and Physical Sciences, Griffith University, Nathan, Australia Abstract Background: The King Island Emu (Dromaius ater) of Australia is one of several extinct emu taxa whose taxonomic relationship to the modern Emu (D. novaehollandiae) is unclear. King Island Emu were mainly distinguished by their much smaller size and a reported darker colour compared to modern Emu. Methodology and Results: We investigated the evolutionary relationships between the King Island and modern Emu by the recovery of both nuclear and mitochondrial DNA sequences from sub-fossil remains. The complete mitochondrial control (1,094 bp) and cytochrome c oxidase subunit I (COI) region (1,544 bp), as well as a region of the melanocortin 1 receptor gene (57 bp) were sequenced using a multiplex PCR approach. The results show that haplotypes for King Island Emu fall within the diversity of modern Emu. Conclusions: These data show the close relationship of these emu when compared to other congeneric bird species and indicate that the King Island and modern Emu share a recent common ancestor. King Island emu possibly underwent insular dwarfism as a result of phenotypic plasticity. The close relationship between the King Island and the modern Emu suggests it is most appropriate that the former should be considered a subspecies of the latter. Although both taxa show a close genetic relationship they differ drastically in size. This study also suggests that rates of morphological and neutral molecular evolution are decoupled.
    [Show full text]
  • Diet Preference and Density of the Greater Rhea (Rhea Americana) in Grasslands of the Flooding Pampa, Argentina
    Revista Brasileira de Ornitologia, 24(1), 13-20 ARTICLE March 2016 Diet preference and density of the Greater Rhea (Rhea americana) in grasslands of the Flooding Pampa, Argentina Viviana Comparatore1,3 and Cristina Yagueddú2 1 Laboratorio de Vertebrados, Departamento de Biología, Facultad de Ciencias Exactas y Naturales, Universidad Nacional de Mar del Plata; & Instituto de Investigaciones Marinas y Costeras (IIMyC, CONICET). Funes 3250, Mar del Plata (B7602AYJ), Provincia de Buenos Aires, Argentina. 2 Laboratorio de Botánica, Departamento de Biología, Facultad de Ciencias Exactas y Naturales, Universidad Nacional de Mar del Plata. Funes 3250, Mar del Plata (B7602AYJ), Provincia de Buenos Aires, Argentina. 3 Corresponding author: [email protected] Received on 15 April 2015. Accepted on 28 March 2016. ABSTRACT: The Greater Rhea (Rhea americana) has been greatly affected by habitat alteration and hunting. Density of rheas was estimated by monthly surveys along transects from June 1996 to November 1997 in 760 ha of coastal grasslands in the Flooding Pampa, Buenos Aires Province, Argentina. Fresh feces and vegetation samples were collected in spring 1996, and in winter and spring 1997 to study diet and resource selection. Feces were macroscopically processed, and the vegetal fraction was prepared for micro histological analysis. Vegetation was sampled in quadrants, separated by species and dried to obtain the percentage of dry weight to estimate availability. Mean density was 0.22 ± 0.04 rheas/ha. Dicots were always preferred in relation to their availability in the grassland, and monocots were not preferred in neither of the two spring seasons studied. Monocots were consumed more frequently than dicots in winter.
    [Show full text]
  • Ratites in Zoos
    1 Feeding Guidelines for Ratites in Zoos James Sales, Ph.D(Agric.); Ph.D(Fish. Sci.) Laboratory of Animal Nutrition Dept. of Animal Nutrition, Genetics, Breeding and Ethology Faculty of Veterinary Medicine Ghent University Heidestraat 19 B-9820, Merelbeke BELGIUM Tel: +32 9 2647824 Fax: +32 9 2647848 E-mail: [email protected] 2 Description of the animal Ratites (Order Struthioniformes) are flightless birds, with a raft-like breastbone devoid of a keel, and can be classified into five different families (Table 1). Table 1 Different families of living ratites (Drenowatz et al., 1995) Family Species Common name Adult weight (kg) Struthionidae Struthio camelus Ostrich 90-130 Rheidae Rhea americana Greater rhea 25 Pterocnemia pennata Lesser rhea 25 Casuariidae Casuarius casuarius Double -wattled cassowary 55 Casuarius unappendiculatus Single -wattled cassowary Casuarius bennetti Dwarf cassowary Dromiceiidae Dromaius novaehollandiae Emu 41 Apterygidae Apteryx austalis Brown kiwi 2 Apteryx hoastii Great spotted kiwi Apteryx awenii Little spotted kiwi Despite their similarities to other birds, ratites have evolved unique characteristics in order to survive in their natural habitat. Of interest from a nutritional standpoint are the modifications in the gastrointestinal tract and the functional characteristics these changes allow (Angel, 1996). Ratites do not have teeth or a crop, which is a feed storage organ in other avian species. While fibre fermentation appears to take place in the large intestine (colon) of the ostrich, the distal ileum serves as a fermentation organ in the emu. The most distinctive characteristic of the gastrointestinal tract of the rhea is the relative large cecum (Table 2).
    [Show full text]
  • Latitudinal Patterns in the Diet of Andean Condor (Vultur Gryphus) In
    Science of the Total Environment 741 (2020) 140220 Contents lists available at ScienceDirect Science of the Total Environment journal homepage: www.elsevier.com/locate/scitotenv Latitudinal patterns in the diet of Andean condor (Vultur gryphus)in Chile: Contrasting environments influencing feeding behavior Melanie Duclos a,f,⁎, Pablo Sabat a,b, Seth D. Newsome c, Eduardo F. Pavez d, Cristóbal Galbán-Malagón e, Fabian M. Jaksic a, Verónica Quirici f,⁎⁎ a Center of Applied Ecology and Sustainability (CAPES), Santiago, Chile b Departamento de Ciencias Ecológicas, Facultad de Ciencias, Universidad de Chile, Casilla 653, Santiago, Chile c Department of Biology, University of New México, Albuquerque, NM 87131, USA d Bioamérica Consultores, Avenida Nueva Providencia 1881, Of. 2208, Providencia, Santiago, Chile e Centro GEMA, Genómica Ecología y Medio Ambiente, Universidad Mayor, Huechuraba, Santiago, Chile f Centro de Investigación para la Sustentabilidad (CIS), Universidad Andres Bello, República 440, Santiago, Chile HIGHLIGHTS GRAPHICAL ABSTRACT • Anthropized environments alter avail- ability of resources for Andean condors. • Little evidence as to how condor's diet vary along their large latitudinal range • Introduced domestic and exotic species are common Andean condor prey across Chile. • Condors in anthropized areas use C4- based food from landfills/corn-fed live- stock. • Anthropic subsidies may help stabilize populations, also enhance mortality risks. article info abstract Article history: Human-dominated environments alter the availability and quality of resources for many species, especially for Received 18 April 2020 scavengers that have large home ranges and plastic foraging behaviors that enable them to exploit novel re- Received in revised form 12 June 2020 sources. Along the western slope of the Andes, the modification of natural landscapes have resulted in significant Accepted 12 June 2020 declines in native prey, the introduction of non-native species, and an increase in the availability of anthropo- Available online 21 June 2020 genic resources.
    [Show full text]
  • Husbandry Guidelines For
    Kelly Swarbrick 1068 Certificate III in Captive Animals 16/11/09 RUV30204 Husbandry Guidelines for Kelly Swarbrick 2008 Emus Dromaius novaehollandiae Aves: Casuariidae Compiler: Kelly Swarbrick th Date of Preparation: 16 November 2009 Western Sydney Institute of TAFE, Richmond Course Name: Certificate III in Captive Animals Course Number: 1068 Lecturers: Graeme Phipps, Jacki Salkeld, and Brad Walker 1 Kelly Swarbrick 1068 Certificate III in Captive Animals 16/11/09 RUV30204 DISCLAIMER This Emu Husbandry Manual is intended to present the current scientific, experiential and practical understanding of the captive care of Emus. Some contributions lend themselves to scientific rigor, where material presented is supported by peer-reviewed literature. Other contributions are based, out of necessity, on the collective experience of professional keepers, because relevant scientific literature is scant or non-existent. The author cannot be, and is not, legally, financially or in any other way, responsible for the application of techniques described within the Manual. When undertaking any procedures or techniques outlined in the Manual, it is up to individual workers to assess the unique circumstances of their situation, apply common sense, and subsequently apply any procedures or techniques at their own risk. In all cases, the reader of this Manual is cautioned not to use this manual as an exact step-by-step guide, but rather as a starting reference point for further case-specific studies. 2 Kelly Swarbrick 1068 Certificate III in Captive Animals 16/11/09 RUV30204 OCCUPATIONAL HEALTH AND SAFETY RISKS Exhibiting Emus falls under the medium risk category (hazardous). This is due to their powerful legs that could deliver a nasty kick.
    [Show full text]