Comprehensive phylogenetic analysis of all species of swordtails and platies (Pisces: Genus Xiphophorus) uncovers a hybrid origin of a swordtail fish, Xiphophorus monticolus, and demonstrates that the sexually selected sword originated in the ancestral lineage of the genus, but was lost again secondarily

BMC Evolutionary Biology

Volumn 13, Issue 1, 2013, Pages

  Kang, Ji Hyoun ^a   Schartl, Manfred ^b   Walter, Ronald B ^c   Meyer, Axel ^a  

^a UNIVERSITY OF KONSTANZ   (Germany)

^b UNIVERSITY OF WÜRZBURG   (Germany)

^c Texas State University   (United States)

Author keywords

[No Author keywords available]

Indexed keywords

CYPRINID; EVOLUTIONARY BIOLOGY; GENETIC MARKER; HYBRID; HYBRIDIZATION; MOLECULAR ANALYSIS; PHYLOGENETICS; PHYLOGENY; SEXUAL SELECTION; SPECIATION (BIOLOGY); TAXONOMY;

MESOAMERICA;

PISCES; XIPHOPHORUS; XIPHOPHORUS CLEMENCIAE; XIPHOPHORUS MACULATUS; XIPHOPHORUS MAYAE; XIPHOPHORUS MIXEI; XIPHOPHORUS MONTICOLUS;

MITOCHONDRIAL DNA;

ANIMAL; ARTICLE; CELL NUCLEUS; CHIMERA; CLASSIFICATION; CYPRINODONTIFORMES; DNA SEQUENCE; EVOLUTION; FEMALE; GENETIC MARKER; GENETICS; HYBRIDIZATION; MALE; PHYLOGENY; STATISTICAL MODEL;

ANIMALS; BIOLOGICAL EVOLUTION; CELL NUCLEUS; CHIMERA; CYPRINODONTIFORMES; DNA, MITOCHONDRIAL; FEMALE; GENETIC MARKERS; HYBRIDIZATION, GENETIC; LIKELIHOOD FUNCTIONS; MALE; PHYLOGENY; SEQUENCE ANALYSIS, DNA;

MLCS; MLOWN;

EID: 84872908276     PISSN: None     EISSN: 14712148     Source Type: Journal    
DOI: 10.1186/1471-2148-13-25     Document Type: Article

References (99)

1. Middle-American poeciliid fish of the genus Xiphophorus. Rosen DE, Bull Fla State Mus Biol Sci 1960 5 57 242
2. The genus Xiphophorus in Mexico and Central America. Kallman KD, Kazianis S, Zebrafish 2006 3 271 285 10.1089/zeb.2006.3.271 18377209 (Pubitemid 44569738)
3. Fishes from the uplands and intermontane basins of Guatemala: revisionary studies and comparative geography. Rosen DE, Bull Am Mus Nat Hist 1979 162 267 376
4. Testing the viviparity-driven-conflict hypothesis: parent-offspring conflict and the evolution of reproductive isolation in a poeciliid fish. Schrader M, Travis J, Am Nat 2008 172 806 817 10.1086/592999 18950276
5. Do embryos influence maternal investment? Evaluating maternal-fetal coadaptation and the potential for parent-offspring conflict in a placental fish. Schrader M, Travis J, Evolution 2009 63 2805 2815 10.1111/j.1558-5646. 2009.00763.x 19573085
6. Molecular phylogenetic relationships and the evolution of the placenta in Poecilia (Micropoecilia) (Poeciliidae: Cyprinodontiformes). Meredith RW, Pires MN, Reznick DN, Springer MS, Mol Phylogenet Evol 2010 55 631 639 10.1016/j.ympev.2009.11.006 19922806
7. Female preference predates the evolution of the sword in swordtail fish. Basolo AL, Science 1990 250 808 810 10.1126/science.250.4982.808 17759973 (Pubitemid 120034304)
8. Phylogenetic evidence for the role of a pre-existing bias in sexual selection. Basolo AL, Proc R Soc Lond B 1995 259 307 311 10.1098/rspb.1995.0045
9. Factors governing sexual selection as an isolating mechanism in the poeciliid fish Lebistes Reticulatus. Haskins CP, Haskins EF, Proc Natl Acad Sci USA 1950 36 464 476 10.1073/pnas.36.9.464 16578341
10. Sex chromosomes and sexual selection in poeciliid fishes. Lindholm A, Breden F, Am Nat 2002 160 214 224 10.1086/341016 18707488 (Pubitemid 34805520)
11. Hybrid origin of a swordtail species (Teleostei: Xiphophorus clemenciae) driven by sexual selection. Meyer A, Salzburger W, Schartl M, Mol Ecol 2006 15 721 730 10.1111/j.1365-294X.2006.02810.x 16499697 (Pubitemid 43344788)
12. Sexual behavior, genes, and evolution in Xiphophorus. Rosenthal GG, Garcia de Leon FJ, Zebrafish 2006 3 85 90 10.1089/zeb.2006.3.85 18248248
13. The molecular basis of color vision in colorful fish: four long wave-sensitive (LWS) opsins in guppies (Poecilia reticulata) are defined by amino acid substitutions at key functional sites. Ward MN, Churcher AM, Dick KJ, Laver CR, Owens GL, Polack MD, Ward PR, Breden F, Taylor JS, BMC Evol Biol 2008 8 210 10.1186/1471-2148-8-210 18638376
14. Complementary effect of natural and sexual selection against immigrants maintains differentiation between locally adapted fish. Plath M, Riesch R, Oranth A, Dzienko J, Karau N, Schiessl A, Stadler S, Wigh A, Zimmer C, Arias-Rodriguez L, Schlupp I, Tobler M, Naturwissenschaften 2010 97 769 774 10.1007/s00114-010-0691-x 20574847
15. Tactical release of a sexually-selected pheromone in a swordtail fish. Rosenthal GG, Fitzsimmons JN, Woods KU, Gerlach G, Fisher HS, PLoS One 2011 6 16994 10.1371/journal.pone.0016994 21339807
16. Male swords and female preferences: response. Basolo AL, Science 1991 253 1426 1427 17793485
17. Darwin C, The descent of man, and selection in relation to sex London: Murray 1871
18. Sexual selection, sensory systems and sensory exploitation. Ryan MJ, Oxf Surv Evol Biol 1990 7 157 195
19. Sexual selection, receiver biases, and the evolution of sex differences. Ryan MJ, Science 1998 281 1999 2003 9748154 (Pubitemid 28475482)
20. A further examination of a preexisting bias favoring a sword in the genus Xiphophorus. Basolo AL, Anim Behav 1995 50 365 375 10.1006/anbe.1995.0252 (Pubitemid 26447842)
21. Monophyly and geography of the Rio Panuco basin swordtails (genus Xiphophorus) with descriptions of four new species. Rauchenberger M, Kallman KD, Morizot DC, Am Mus Novit 1990 2975 1 41
22. Recurrent origin of a sexually selected trait in Xiphophorus fishes inferred from a molecular phylogeny. Meyer A, Morrissey JM, Schartl M, Nature 1994 368 539 542 10.1038/368539a0 8139686 (Pubitemid 24118768)
23. Preexisting mating biases and the evolution of the sword in the genus Xiphophorus. Basolo AL, Am Zool 1990 30 80
24. A phylogenetic and biogeographic perspective on the evolution of poeciliid fishes. Hrbek T, Seckinger J, Meyer A, Mol Phylogenet Evol 2007 43 986 998 10.1016/j.ympev.2006.06.009 17185005 (Pubitemid 46777372)
25. Likelihood of ancestor states in adaptive radiation. Schluter D, Price T, Mooers AO, Ludwig D, Evolution 1997 51 1699 1711 10.2307/2410994 (Pubitemid 28104532)
26. The evolution of sexually selected traits in male swordtail fishes (Xiphophorus: Poeciliidae). Meyer A, Heredity 1997 79 329 337 10.1038/hdy.1997.161 (Pubitemid 27393028)
27. Character definitions, sexual selection, and the evolution of swordtails. Wiens JJ, Morris MR, Am Nat 1996 147 866 869 10.1086/285882
28. Xiphophorus kallmani sp. n.-a new species of swordtail from Mexico (Teleostei, Cyprinodontiformes, Poeciliidae). Meyer MK, Schartl M, Zool Abh, Staat Mus Tierk Dresden 2003 53 57 64
29. Xiphophorus mayae, a new species of swordtail from Guatemala (Teleostei: Poeciliidae). Meyer MK, Schartl M, Ichthyol Explor Freshwaters 2002 13 59 66
30. Two new species of Xiphophorus (Poeciliidae) from the Isthmus of Tehuantepec, Oaxaca, Mexico with a discussion of the distribution of the X. clemenciae clade. Kallman KD, Walter RB, Morizot DC, Kazianis S, Am Mus Novit 2004 3441 1 34
31. Arnold ML, Natural hybridization and evolution New York: Oxford University Press 1997
32. Natural hybridization in primates: one evolutionary mechanism. Arnold ML, Meyer A, Zoology 2006 109 261 276 10.1016/j.zool.2006.03.006 16945512 (Pubitemid 44572717)
33. Hybrid speciation. Mallet J, Nature 2007 446 279 283 10.1038/nature05706 17361174 (Pubitemid 46426165)
34. Introgressive hybridization between the allotetraploid Coffea arabica and one of its diploid ancestors, Coffea canephora, in an exceptional sympatric zone in New Caledonia. Mahe L, Le Pierres D, Combes MC, Lashermes P, Genome 2007 50 316 324 10.1139/G07-011 17502905 (Pubitemid 46923643)
35. Unpredictable evolution in a 30-year study of Darwin's finches. Grant PR, Grant BR, Science 2002 296 707 711 10.1126/science.1070315 11976447 (Pubitemid 34441992)
36. Hybridization and adaptive radiation. Seehausen O, Trends Ecol Evol 2004 19 198 207 10.1016/j.tree.2004.01.003 16701254 (Pubitemid 38355799)
37. Introgressive hybridization between two Iberian endemic cyprinid fish: a comparison between two independent hybrid zones. Aboim MA, Mavarez J, Bernatchez L, Coelho MM, J Evol Biol 2010 23 817 828 10.1111/j.1420-9101.2010.01953.x 20210830
38. Evidence for multiple sources of invasion and intraspecific hybridization in Brachypodium sylvaticum (Hudson) Beauv. in North America. Rosenthal DM, Ramakrishnan AP, Cruzan MB, Mol Ecol 2008 17 4657 4669 10.1111/j.1365-294X.2008. 03844.x 18627455
39. Replicated hybrid zones of Xiphophorus swordtails along an elevational gradient. Culumber ZW, Fisher HS, Tobler M, Mateos M, Barber PH, Sorenson MD, Rosenthal GG, Mol Ecol 2011 20 342 356 10.1111/j.1365-294X.2010.04949.x 21210879
40. Contrasting patterns of mitochondrial DNA and microsatellite introgressive hybridization between lineages of lake whitefish (Coregonus clupeaformis); relevance for speciation. Lu G, Basley DJ, Bernatchez L, Mol Ecol 2001 10 965 985 10.1046/j.1365-294X.2001.01252.x 11348504 (Pubitemid 32406397)
41. Speciation via introgressive hybridization in East African cichlids? Salzburger W, Baric S, Sturmbauer C, Mol Ecol 2002 11 619 625 10.1046/j.0962-1083.2001.01438.x 11918795 (Pubitemid 34235071)
42. Reticulate phylogeny of gastropod-shell-breeding cichlids from Lake Tanganyika-the result of repeated introgressive hybridization. Koblmuller S, Duftner N, Sefc KM, Aibara M, Stipacek M, Blanc M, Egger B, Sturmbauer C, BMC Evol Biol 2007 7 7 10.1186/1471-2148-7-7 17254340 (Pubitemid 46198544)
43. The role of hybridization in evolution. Barton NH, Mol Ecol 2001 10 551 568 11298968 (Pubitemid 32406342)
44. Discordance of nuclear and mitochondrial DNA phylogenies in Hawaiian Drosophila. DeSalle R, Giddings LV, Proc Natl Acad Sci USA 1986 83 6902 6906 10.1073/pnas.83.18.6902 3462736
45. Hybrid origin of the Pliocene ancestor of wild goats. Ropiquet A, Hassanin A, Mol Phylogenet Evol 2006 41 395 404 10.1016/j.ympev.2006.05.033 16837213 (Pubitemid 44428202)
46. Phylogenetic incongruence between nuclear and mitochondrial markers in the Asian colobines and the evolution of the langurs and leaf monkeys. Ting N, Tosi AJ, Li Y, Zhang YP, Disotell TR, Mol Phylogenet Evol 2008 46 466 474 10.1016/j.ympev.2007.11.008 18180172
47. Molecular phylogeography of the asp viper Vipera aspis (Linnaeus, 1758) in Italy: evidence for introgressive hybridization and mitochondrial DNA capture. Barbanera F, Zuffi MA, Guerrini M, Gentilli A, Tofanelli S, Fasola M, Dini F, Mol Phylogenet Evol 2009 52 103 114 10.1016/j.ympev.2009.02.006 19236928
48. Dissolution of sexual signal complexes in a hybrid zone between the swordtails Xiphophorus birchmanni and Xiphophorus malinche (Poeciliidae). Rosenthal GG, De La Rosa Reyna XF, Kazianis S, Stephens MJ, Morizot DC, Ryan MJ, De Leon FJG, Copeia 2003 2003 299 307 10.1643/0045-8511(2003)003[0299:DOSSCI]2. 0.CO;2 (Pubitemid 36784926)
49. Melanomas in the hybrid offspring of two species of swordtails, Xiphophorus montezumae and Xiphophorus hellerii. Gordon M, Anat Rec 1947 99 613 18935417
50. Effect of sex on the development of melanoma in hybrid fish of the genus Xiphophorus. Siciliano MJ, Perlmutter A, Clark E, Cancer Res 1971 31 725 729 5104321
51. Age-specific incidence of hereditary melanomas in the Xiphophorus fish hybrids. Ozato K, Wakamatsu Y, Carcinogenesis 1981 2 129 133 10.1093/carcin/2.2.129 7273296 (Pubitemid 11139134)
52. Genetic mapping in Xiphophorus hybrid fish: assignment of 43 AP-PCR/RAPD and isozyme markers to multipoint linkage groups. Kazianis S, Morizot DC, McEntire BB, Nairn RS, Borowsky RL, Genome Res 1996 6 280 289 10.1101/gr.6.4.280 8723721 (Pubitemid 26161690)
53. Differential regulation of msx genes in the development of the gonopodium, an intromittent organ, and of the "sword," a sexually selected trait of swordtail fishes (Xiphophorus). Zauner H, Begemann G, Mari-Beffa M, Meyer A, Evol Dev 2003 5 466 477 10.1046/j.1525-142X.2003.03053.x 12950626 (Pubitemid 37143923)
54. Deciphering the swordtail's tale: a molecular and evolutionary quest. Wilkins AS, Bioessays 2004 26 116 119 10.1002/bies.10414 14745829 (Pubitemid 38197009)
55. The role of sexual selection in the loss of sexually selected traits in the swordtail fish Xiphophorus continens. Morris MR, Moretz JA, Farley K, Nicoletto P, Anim Behav 2005 69 1415 1424 10.1016/j.anbehav.2004.08.013 (Pubitemid 40793447)
56. An organizer controls the development of the "sword," a sexually selected trait in swordtail fish. Eibner C, Pittlik S, Meyer A, Begemann G, Evol Dev 2008 10 403 412 10.1111/j.1525-142X.2008.00251.x 18638317 (Pubitemid 351991475)
57. Eine neue Xiphophorus-Art aus Vera Cruz Mexiko (Pisces: Poeciliidae). Meyer MK, Schartl M, Senkenberg Biol 1980 60 147 151
58. A phylogeographic investigation of the hybrid origin of a species of swordtail fish from Mexico. Jones JC, Perez-Sato JA, Meyer A, Mol Ecol 2012 21 2692 2712 10.1111/j.1365-294X.2012.05571.x 22494472
59. A private ultraviolet channel in visual communication. Cummings ME, Rosenthal GG, Ryan MJ, Proc R Soc Biol Sci Ser B 2003 270 897 904 10.1098/rspb.2003.2334 (Pubitemid 36561024)
60. Vertical bars on male Xiphophorus multilineatus: a signal that deters rival males and attracts females. Morris MR, Mussel M, Ryan MJ, Behav Ecol 1995 6 274 279 10.1093/beheco/6.3.274 (Pubitemid 26447841)
61. Responses to conspecific and heterospecific olfactory cues in the swordtail Xiphophorus cortezi. McLennan DA, Ryan MJ, Anim Behav 1997 54 1077 1088 10.1006/anbe.1997.0504 9398364 (Pubitemid 27520240)
62. Genetic variation and phylogeography of the swordtail fish Xiphophorus cortezi (Cyprinodontiformes, Poeciliidae). Gutierrez-Rodriguez C, Morris MR, Dubois NS, Queiroz K, Mol Phylogenet Evol 2007 43 111 123 10.1016/j.ympev.2006. 10.022 17194605 (Pubitemid 46484356)
63. Phylogeography and monophyly of the swordtail fish species Xiphophorus birchmanni (Cyprinodontiformes, Poeciliidae). Gutierrez-Rodriguez C, Shearer AE, Morris MR, de Queiroz K, Zool Scr 2008 37 129 139 10.1111/j.1463-6409.2007. 00311.x (Pubitemid 351208044)
64. Ontogeny and phylogeny in the northern swordtail clade of Xiphophorus. Marcus JM, McCune AR, Syst Biol 1999 48 491 522 10.1080/106351599260111 (Pubitemid 129479560)
65. Phylogenetic relationships among populations of northern swordtails (Xiphophorus) as inferred from allozyme data. Morris MR, de Queiroz K, Morizot DC, Copeia 2001 2001 65 81 10.1643/0045-8511(2001)001[0065:PRAPON]2.0.CO;2 (Pubitemid 32175253)
66. Arbitrarily primed DNA fingerprinting for phylogenetic reconstruction in vertebrates: the Xiphophorus model. Borowsky RL, McClelland M, Cheng R, Welsh J, Mol Biol Evol 1995 12 1022 1032 8524037
67. Microsatellites in the genus Xiphophorus, developed in Xiphophorus montezumae. Seckinger J, Brinkmann H, Meyer A, Mol Ecol Notes 2002 2 4 6 10.1046/j.1471-8286.2002.00124.x (Pubitemid 34287094)
68. Molecular phylogeny and biogeography of the Cuban genus Girardinus Poey, 1854 and relationships within the tribe Girardinini (Actinopterygii, Poeciliidae). Doadrio I, Perea S, Alcaraz L, Hernandez N, Mol Phylogenet Evol 2009 50 16 30 10.1016/j.ympev.2008.09.014 18854217
69. MEGA4: Molecular Evolutionary Genetics Analysis (MEGA) software version 4.0. Tamura K, Dudley J, Nei M, Kumar S, Mol Biol Evol 2007 24 1596 1599 10.1093/molbev/msm092 17488738 (Pubitemid 47236692)
70. jModelTest: phylogenetic model averaging. Posada D, Mol Biol Evol 2008 25 1253 1256 10.1093/molbev/msn083 18397919 (Pubitemid 351882013)
71. An approximately unbiased test of phylogenetic tree selection. Shimodaira H, Syst Biol 2002 51 492 508 10.1080/10635150290069913 12079646 (Pubitemid 35180700)
72. CONSEL: for assessing the confidence of phylogenetic tree selection. Shimodaira H, Hasegawa M, Bioinformatics 2001 17 1246 1247 10.1093/ bioinformatics/17.12.1246 11751242 (Pubitemid 33735360)
73. Testing hybridization hypotheses based on incongruent gene trees. Sang T, Zhong Y, Syst Biol 2000 49 422 434 10.1080/10635159950127321 12116420
74. Gene tree discordance, phylogenetic inference and the multispecies coalescent. Degnan JH, Rosenberg NA, Trends Ecol Evol 2009 24 332 340 10.1016/j.tree.2009.01.009 19307040
75. 170 Years of "Lock-and-Key": Genital Morphology and Reproductive Isolation. Masly JP, Int J Evol Biol 2012 2012 247352 22263116
76. Female heterogamety in the swordtail, Xiphophorus alvarezi Rosen (Pisces, Poeciliidae), with comments on a natural polymorphism affecting sword coloration. Kallman KD, Bao IY, J Exp Zool 1987 243 93 102 10.1002/jez. 1402430112 3612059
77. Ancient versus reticulate origin of a hemiclonal lineage. Mateos M, Vrijenhoek RC, Evolution 2002 56 985 992 12093033 (Pubitemid 34617021)
78. Shared preferences by predators and females for male ornaments in swordtails. Rosenthal GG, Flores Martinez TY, Garcia de Leon FJ, Ryan MJ, Am Nat 2001 158 146 154 10.1086/321309 18707343 (Pubitemid 32682541)
79. The turn of the sword: length increases male swimming costs in swordtails. Basolo AL, Alcaraz G, Proc Biol Sci 2003 270 1631 1636 10.1098/rspb.2003.2388 12908985 (Pubitemid 36960295)
80. Female disdain for swords in a swordtail fish. Wong BB, Rosenthal GG, Am Nat 2006 167 136 140 10.1086/498278 16475105 (Pubitemid 43039432)
81. Secondary reduction of preference for the sword ornament in the pygmy swordtail Xiphophorus nigrensis (Pisces: Poeciliidae). Rosenthal GG, Wagner WE, Ryan MJ, Anim Behav 2002 63 37 45 10.1006/anbe.2001.1887
82. Is UV ornamentation an amplifier in swordtails? Cummings ME, de Garcia Leon FJ, Mollaghan DM, Ryan MJ, Zebrafish 2006 3 91 100 10.1089/zeb.2006.3.91 18248249
83. Museum fish specimens and molecular taxonomy: A comparative study on DNA extraction protocols and preservation techniques. Chakraborty A, Sakai M, Iwatsuki Y, J Appl Ichthyol 2006 22 160 166 (Pubitemid 44056520)
84. Convenient single-step, one tube purification of PCR products for direct sequencing. Werle E, Schneider C, Renner M, Volker M, Fiehn W, Nucleic Acids Res 1994 22 4354 4355 10.1093/nar/22.20.4354 7937169 (Pubitemid 24322750)
85. CLUSTAL W: improving the sensitivity of progressive multiple sequence alignment through sequence weighting, position-specific gap penalties and weight matrix choice. Thompson JD, Higgins DG, Gibson TJ, Nucleic Acids Res 1994 22 4673 4680 10.1093/nar/22.22.4673 7984417 (Pubitemid 24354800)
86. BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Hall TA, Nucleic Acids Symp Ser 1999 41 95 98
87. Confidence-limits on phylogenies-an approach using the bootstrap. Felsenstein J, Evolution 1985 39 783 791 10.2307/2408678
88. Further analysis of data by Akaikes information criterion and finite corrections. Sugiura N, Commun Stat A-Theor 1978 7 13 26 10.1080/ 03610927808827599
89. New algorithms and methods to estimate maximum-likelihood phylogenies: assessing the performance of PhyML 3.0. Guindon S, Dufayard JF, Lefort V, Anisimova M, Hordijk W, Gascuel O, Syst Biol 2010 59 307 321 10.1093/sysbio/syq010 20525638
90. Bayesian phylogenetic inference using DNA sequences: a Markov Chain Monte Carlo Method. Yang Z, Rannala B, Mol Biol Evol 1997 14 717 724 10.1093/oxfordjournals.molbev.a025811 9214744 (Pubitemid 27287292)
91. Bayesian phylogenetic inference via Markov chain Monte Carlo methods. Mau B, Newton MA, Larget B, Biometrics 1999 55 1 12 10.1111/j.0006-341X.1999.00001. x 11318142 (Pubitemid 29161999)
92. MrBayes 3: Bayesian phylogenetic inference under mixed models. Ronquist F, Huelsenbeck JP, Bioinformatics 2003 19 1572 1574 10.1093/bioinformatics/ btg180 12912839 (Pubitemid 37038874)
93. MRBAYES: Bayesian inference of phylogenetic trees. Huelsenbeck JP, Ronquist F, Bioinformatics 2001 17 754 755 10.1093/bioinformatics/17.8.754 11524383 (Pubitemid 32851390)
94. PartitionFinder: Combined Selection of Partitioning Schemes and Substitution Models for Phylogenetic Analyses. Lanfear R, Calcott B, Ho SY, Guindon S, Mol Biol Evol 2012 29 1695 1701 10.1093/molbev/mss020 22319168
95. TREE-PUZZLE: maximum likelihood phylogenetic analysis using quartets and parallel computing. Schmidt HA, Strimmer K, Vingron M, von Haeseler A, Bioinformatics 2002 18 502 504 10.1093/bioinformatics/18.3.502 11934758 (Pubitemid 34284960)
96. A rapid bootstrap algorithm for the RAxML Web servers. Stamatakis A, Hoover P, Rougemont J, Syst Biol 2008 57 758 771 10.1080/10635150802429642 18853362
97. Mesquite: A modular system for evolutionary analysis. Maddison WP, Maddison DR, Version 2.74 2010 http://mesquiteproject.org
98. The maximum likelihood approach to reconstructing ancestral character states of discrete characters on phylogenies. Pagel M, Syst Biol 1999 48 612 622 10.1080/106351599260184 (Pubitemid 129479567)
99. A likelihood approach to estimating phylogeny from discrete morphological character data. Lewis PO, Syst Biol 2001 50 913 925 10.1080/106351501753462876 12116640