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AR I Fi T Ifi H Bidi Ti a Review of Interspecific Hybridization in the Order Testudines

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ARA Rev iew o fIf In terspec ific Hy bidibridiza tion in the Order Testudines Timothy R. Brophy, Wayne Frair, and Darlene Clark Classification of Extant Turtles Order Testudines (Ernst et al., 2000) Suborder Pleurodira (side-necked turtles) FamilyPelomedusidae - S.A. & Africa Family Chelidae - S.A. & Australia Suborder Crytodira (hidden -necked turtles) Superfamily Trionychoidea FilKitFamily Kinostern idae - Md&MMud & Musk Family Dermatemydidae - C.A. River FilCtthlidFamily Carettochelyidae - Pig Nose Family Trionychidae - Softshell Classification of Extant Turtles Suborder Crytodira (hidden-necked turtles) Superfamily Chelonioidea Family Cheloniidae - Marine Family Dermochelyidae - Leatherback SSpuperfamily yT Testudinoidea Family Chelydridae - Snapping Family Platysternidae - Big Headed Family Emydidae - N.W. Pond Family Geoemydidae - OWO.W. Pond Family Testudinidae - Tortoises TTBurtle Baraminology • Turtles have been the subject of much baraminological research (see Wood, 2005) – Frair (1984) – All turtle species constitute a ppyypolytypic baramin with four diversification lines (Pleuro dira, Ch el onioi dea, T rionych id ae & rest of Cryptodira). Diversification line ≈ holobaramin – Frair (1991) – All turtles descended from a created ancestor (possibly PPgroganocheys y))D. Did not discard hypothesis of four diversification lines TTBurtle Baraminology • Turtles have been the subject of much baraminological research (see Wood, 2005) – Wise (1992) – Turtles are apobaraminic. Some evidence supports Frair’s (1984) four holobaramins – Robinson (1997) – Further supports contention tha t tur tles are apo baram in ic. Che lon iidae & Gopherus are monobaraminic Turtle Baraminology • Turtles have been the subject of much baraminological research (see Wood , 2005) – WW(5)ood (2005) – TTpurtles are apobaraminic – Composed of five holobaramins: Chelidae, Pelomedusidae, Trionychoidea, other Cryptodira & Proganochelys TTHyurtle Hybridization • AAg thorough review of inters pecific hybridization, with baraminological interppypretations, has yet to be reported • Robinson (1997) incorporated the use of hbiditihybridization in hibhis baram ino log ica l ana lyses o f the marine turtles & gopher tortoises • We set out to find evidence of interspecific hyygbridization throughout the entire turtle order & use this evidence to “build” monobaramins SSyRummary of Results • EEyvidence of hybridization in 8 of 1 3 families Pelomedusidae Chelidae Kinosternidae DDyermatem1ydidae CCyarrettochel1 yidae TTyrionychidae Cheloniidae Dermochelyidae1 Chelydridae Platysternidae1 Emydddidae GddGeoemydidae TddTestudinidae Summary of Results • Crosses between 89 unique species pairs • 29 of these are intergeneric (≈33%) • No interfamilial or inter-holobaraminic ((W,5)sensu Wood, 2005) crosses • 16 small monobaramins (2-4 species) • 2 medium monobaramins (5-8 species) • 2 large monobaramins (>8 species) FFyPamily Pelomedusidae – 1sm. 1 cross 2 spp. Pelusios seychellensis Pelusios castanoides 1 Small Monobaramin FFyCamily Chelidae – 2sm. 3 crosses Chelodina longicollis 2IG2 IG Chelodina canni 4 spp. 1st Small Monobaramin Emydura albertisii Elseya novaeguineae FFyCamily Chelidae Phrynops hilari Phrynops geoffroanus 3 crosses 4spp4 spp. Phrynops williamsi Phrynops gibbus 2nd Small Monobaramin Family Kinosternidae – 1med. Kinos ternon su bru brum Kinos ternon flavescens Kinosternon baurii Sternotherus depressus Sternotherus carinatus 6 crosses Sternotherus minor Sternotherus odoratus 1 IG 7 spp. 1 Medium Monobaramin FFyTyamily Trionychidae – 1sm. 1 “cross” 2 spp. Apalone spinifera Apalone mutica 1 Small Monobaramin Family Cheloniidae – 1med. 7 crosses 6 IG, 3 IT 5 spp. Lepidochelys kempii Lepidochelys olivacea Caretta caretta 1M1 Medi um M onob arami n Eretmochelys imbricata Chelonia mydas FFyCamily Cheloniidae • What about NNp(atator depressus (flatback turtle ))? • No records of hybridization with any other species • May be due in part to its limited range in Australia and New Guinea (Karl et al., 1995) Family Cheloniidae • Tribes Carettini (Caretta, Lepidochelys & Ere tmoche lys)&Ch) & Chelonini (Che lonia) thought to have been separated for 50-75 million years • “May be among the oldest vertebrate lineages capppable of producing viable hy brids in nature” (Kar l et a l., 1995 ) Baraminology of the Cheloniidae • Robinson (1997) considered the Cheloniidae in his baram ino log ica l ana lyses: – Concluded entire family was a single monobaramin – NttNatator depressus fllithifell within ran ge o f cy toc hrome b sequence variation of hybridizing turtles, so it was included as part of the monobaramin • We reviewed several recent reports of hyyyCbridization in the family Cheloniidae – Published since or not included in Robinson (1997) – Several with molecular verification – Confirm Robinson’s (1997) conclusions Recent Reports of Sea Turtle Hybridization Species Crossed N/A? Evidence Reference Morphology, Caretta caretta x Lepidochelys kempii Natural mtDNA Barber et al., 2003 & nuclear DNA Morphology, Garman, 1888; Caretta caretta x Eretmochelys imbricata Natural mtDNA Kamezaki, 1983; Frazier, 1988; & nuclear DNA Witzell & Schmid, 2003 Kamezake et al., 1996; Morphology & Hemens, 1997; Kawata, 2003; Caretta caretta x Chelonia mydas Natural mtDNA Phillips, 2003; Jones et al., 2004 Morphology, Buden & Edward, 2001; Chelonia mydas x Eretmochelys imbricata Natural mtDNA Kawata, 2003; & scnDNA Seminoff et al., 2003 Family Testudinidae – 6sm. Testudo graeca Testudo hermanni 3 crosses 4 spp. Testudo horsfieldi Testudo marginata 1st Small Monobaramin FFyTamily Testudinidae 1 cross 2 spp. Indotestudo forstenii Indotestudo elongata 2nd SllMbSmall Monobaramin FFyTamily Testudinidae 1 cross 2 spp. Gopherus berlandieri Gopherus agassizii 3rd SllMbSmall Monobaramin Baraminology of Gopher Tortoises • Robinson (1997) considered the genus Gopherus in his baraminological analyses: – Concluded entire genus was a single monobaramin (4 spp)ecies): G. agg,assizii, G. berlandieri , G. flavomarginatus, G. polyphemus – G. flavomarginatus & G. polyphemus fell within range of cytochrome b sequence variation of hybridizing turtles, so they were included as part of the monobaramin Family Testudinidae Kinixys erosa 3 crosses 3 spp. Kinixys homeana Kinixys belliana 4th Small Monobaramin Family Testudinidae Madagascar SAmericaS. America Geochelone denticulata GhldGeochelone radiata 2 crosses, 3 spp. GhlGeochelone car bonaria ??? 1 cross, 2 spp. Geochelone sulcata Geochelone pardalis 5th & 6th Small Monobaramins EdidEdiEmydidae: Emydinae – 3sm. 1 cross 2 spp. TlTerrapene carolina Terrapene ornata 1st Small Monobaramin Fam ily Emy didae: Emy dinae 1cross1 cross 1 IG Clemmys guttata2 spp. Glyptemys muhlenbergii 2nd SllMbSmall Monobaramin Familyyy Emydidae: Em ydinae Glyptemys insculpta 4 crosses Emys blandingii 4 IG 4 spp. Emys orbicularis Actinemys marmorata 3rd Small Monobaramin Wood x Blandinggy’s Hybrid X Hybridization in the Emydinae • Emydinae = Clemmys, Glyptemys, Actinemys, Emys & Terrapene Clemmys guttata Glyptemys spp. Terrapene spp. 1 Large Monobaramin? Actinemys marmorata Emys spp. Emydidae: Deirochelyinae - 1sm.&1l. Trachemys stejnegeri 2 crosses 3 spp. ThTrachemys decorata ThTrachemys terrapen 1 Small Monobaramin Family Emydidae: Deirochelyinae P. floridana P. concinna P. rubriventris P. alabamensis P. texana C. picta P. nelsoni GGp. psuedog eog rap hica TTp. scripta G. versa GhitiG. ouachitensis G. kohnii GhiG. geographica G. barbouri G. oculifera G. nigrinoda Hybridization in the Deirochelyinae • Deirochelyinae = Graptemys, Trachemys, Chrysemys, Psuedemys, Malaclemys & Deirochelys 21 crosses Trachemys scripta Graptemys spp. 5 IG 16 spp. 1 Large Pseudemys spp. Monobaramin Chrysemys picta Family Geoemydidae – 2sm.&1l. Rhinoclemmys punctularia 3 crosses 3 spp. Rhinoc lemmys me lanosterna Rhinoc lemmys dia dema tta 1st Small Monobaramin FilGFamily Geoemy didae 1 cross 2 spp. Mauremys caspica Mauremys rivulata 2nd SllMbSmall Monobaramin Family Geoemydidae M. japonica M. reevesii M. annamensis M. mutica M. sinensis M. nigricans Cyc. shanensis H. grandis S. quadriocellata R. Pulcherimma S. pseudocellata (C. America) Mal. subtrijuga G. japonica CbiiC. amboinensis CflC. flavomarg inata C. galbinifrons C. mouhotii C. bourreti C. trifasciata Hybridization in the Geoemydidae 26 crosses, 11 IG, 19 species Heosemys grandis * Geoemyda japonica Cyclemys shanensis Mauremys spp. Cuora spp. Sacalia spp. 1 Large Monobaramin Rhinoclemmys pulcherimma *Large phylogenetic distance Malayemys subtrijuga Proposed Phylogeny ofhf the Geoemy dddidae * * Spinks et al., 1994 Discussion • 288 turtle species – 41,238 possible hybrids (0.22%) – 5,971 possible intra-fllhbd()familial hybrids (1.5%) • GGgrowing turtle monobaramins – Continue to verify records we already have – MMy,y,any, as yet, undescribed & undiscovered h ybrids (many in process of being published) – Inclusion of non-hybridizing turtles if they fall within range offhbdlf variation of hybridizing turtles • WWjW(5)e are unable to reject Wood’s (2005) hypothesis of five turtle holobaramins.
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    i The comparative ecology of Krefft’s River Turtle Emydura krefftii in Tropical North Queensland. By Dane F. Trembath B.Sc. (Zoology) Applied Ecology Research Group University of Canberra ACT, 2601 Australia A thesis submitted in fulfilment of the requirements of the degree of Masters of Applied Science (Resource Management). August 2005. ii Abstract An ecological study was undertaken on four populations of Krefft’s River Turtle Emydura krefftii inhabiting the Townsville Area of Tropical North Queensland. Two sites were located in the Ross River, which runs through the urban areas of Townsville, and two sites were in rural areas at Alligator Creek and Stuart Creek (known as the Townsville Creeks). Earlier studies of the populations in Ross River had determined that the turtles existed at an exceptionally high density, that is, they were superabundant, and so the Townsville Creek sites were chosen as low abundance sites for comparison. The first aim of this study was to determine if there had been any demographic consequences caused by the abundance of turtle populations of the Ross River. Secondly, the project aimed to determine if the impoundments in the Ross River had affected the freshwater turtle fauna. Specifically this study aimed to determine if there were any difference between the growth, size at maturity, sexual dimorphism, size distribution, and diet of Emydura krefftii inhabiting two very different populations. A mark-recapture program estimated the turtle population sizes at between 490 and 5350 turtles per hectare. Most populations exhibited a predominant female sex-bias over the sampling period. Growth rates were rapid in juveniles but slowed once sexual maturity was attained; in males, growth basically stopped at maturity, but in females, growth continued post-maturity, although at a slower rate.
  • A Case of Inversion and Duplication Involving Constitutive Heterochromatin

    A Case of Inversion and Duplication Involving Constitutive Heterochromatin

    Genetics and Molecular Biology, 36, 3, 353-356 (2013) Copyright © 2013, Sociedade Brasileira de Genética. Printed in Brazil www.sbg.org.br Short Communication Cytogenetic comparison of Podocnemis expansa and Podocnemis unifilis: A case of inversion and duplication involving constitutive heterochromatin Ricardo José Gunski1, Isabel Souza Cunha2, Tiago Marafiga Degrandi1, Mario Ledesma3 and Analía Del Valle Garnero1 1Universidade Federal do Pampa, Campus São Gabriel, São Gabriel, RS, Brazil. 2Secretaria Municipal de Meio Ambiente e Turismo, São Desidério, BA, Brazil. 3Parque Ecológico “El Puma”, Candelaria, Argentina. Abstract Podocnemis expansa and P. unifilis present 2n = 28 chromosomes, a diploid number similar to those observed in other species of the genus. The aim of this study was to characterize these two species using conventional staining and differential CBG-, GTG and Ag-NOR banding. We analyzed specimens of P. expansa and P. unifilis from the state of Tocantins (Brazil), in which we found a 2n = 28 and karyotypes differing in the morphology of the 13th pair, which was submetacentric in P. expansa and telocentric in P. unifilis. The CBG-banding patterns revealed a heterochromatic block in the short arm of pair 13 of P. expansa and an interstitial one in pair 13 of P. unifilis, suggest- ing a pericentric inversion. Pair 14 of P. unifilis showed an insterstitial band in the long arm that was absent in P. expansa, suggesting a duplication in this region. Ag-NORs were observed in the first chromosome pair of both spe- cies and was associated to a secondary constriction and heterochromatic blocks. Keywords: chromosome banding, Ag-NORs, karyotype, chelonids.