Molecular phylogeny and morphological revision of Myotis bats (Chiroptera: Vespertilionidae) from Taiwan and adjacent China

Zootaxa

Volumn 3920, Issue 2, 2015, Pages 301-342

  Ruedi, Manuel ^a   Csorba, Gábor ^b   Lin, Liang Kong ^c   Chou, Cheng Han ^c,d  

^a MUSÉUM D HISTOIRE NATURELLE   (Switzerland)

^b DEPARTMENT OF ZOOLOGY   (Hungary)

^c TUNGHAI UNIVERSITY   (Taiwan)

^d ENDEMIC SPECIES RESEARCH INSTITUTE   (Taiwan)

Author keywords

Cryptic species; Cytochrome b; Multivariate analysis; Myotinae; Submyotodon; Taxonomy

Indexed keywords

CHIROPTERA; MYOTIS; MYOTIS ADVERSUS; MYOTIS FORMOSUS; MYOTIS MURICOLA; VESPERTILIONIDAE;

ANATOMY AND HISTOLOGY; ANIMAL; ANIMAL DISPERSAL; ANIMAL STRUCTURES; BAT; BODY SIZE; CHINA; CLASSIFICATION; ECOSYSTEM; FEMALE; FOSSIL; GENETICS; GROWTH, DEVELOPMENT AND AGING; MALE; ORGAN SIZE; PHYLOGENY; TAIWAN;

ANIMAL DISTRIBUTION; ANIMAL STRUCTURES; ANIMALS; BODY SIZE; CHINA; CHIROPTERA; ECOSYSTEM; FEMALE; FOSSILS; MALE; ORGAN SIZE; PHYLOGENY; TAIWAN;

EID: 84930741591     PISSN: 11755326     EISSN: 11755334     Source Type: Journal    
DOI: 10.11646/zootaxa.3920.2.6     Document Type: Article

References (94)

1. Allen, G.M. (1923) New Chinese bats. American Museum Novitates, 85, 1-8.
2. Allen, G.M. (1938) The mammals of China and Mongolia. Vol. 11. Part. 1. The American Museum of Natural History, New York, 620 pp.
3. Anderson, J. (1881) Catalogue of Mammalia in the Indian Museum, Calcutta. Indian Museum, Calcutta, 223 pp.
4. Ärnbäck-Christie-Linde, A. (1908) A collection of bats from Formosa. Annals and Magazine of Natural History, 2, 235-238. http://dx.doi.org/10.1080/00222930808692477
5. Bates, P.J.J. & Harrison, D.L. (1997) Bats of the Indian Subcontinent (Vol. 43). Harrison Zoological Museum, Sevenoaks, 258 pp.
6. Bates, P.J.J., Hendrichsen, D.K., Walston, J.L. & Hayes, B. (1999) A review of the mouse-eared bats (Chiroptera: Vespertilionidae: Myotis) from Vietnam with significant new records. Acta Chiropterologica, 1, 47-74.
7. Benda, P., Faizolâhi, K., Andreas, M., Obuch, J., Reiter, A., Ševcik, M., Uhrin, M., Vallo, P. & Ashrafi, S. (2012) Bats (Mammalia: Chiroptera) of the Eastern Mediterranean and Middle East. Part 10. Bat fauna of Iran. Acta Societatis Zoologicae Bohemicae, 76, 163-582.
8. Bickham, J.W., Patton, J.C., Schlitter, D.A., Rautenbach, I.L. & Honeycutt, R.L. (2004) Molecular phylogenetics, karyotypic diversity, and partition of the genus Myotis (Chiroptera: Vespertilionidae). Molecular Phylogenetics and Evolution, 33, 333-338. http://dx.doi.org/10.1016/j.ympev.2004.06.012
9. Blyth, E. (1863) Catalogue of the Mammalia in the Museum Asiatic Society. Savielle and Cronenburgh, London, 187 pp.
10. Borisenko, A.V., Kruskop, S.V. & Ivanova, N.V. (2008) A new mouse-eared bat (Mammalia: Chiroptera: Vespertilionidae) from Vietnam. Russian Journal of Theriology, 7, 57-69. http://dx.doi.org/10.1515/mamm.2001.65.1.63
11. Bradley, R. & Baker, R. (2001) A test of the genetic species concept: cytochrome b sequences and mammals. Journal of Mammalogy, 82, 960-973. http://dx.doi.org/10.1644/1545-1542(2001)082〈0960:atotgs〉2.0.co;2
12. Cheng, H.C., Fang, Y.P. & Chou, C.H. (2010) A photographic guide to the bats of Taiwan. The Agriculture Committee Taiwan Endemic Species Research Institute, Taipei, 144 pp. [in Chinese]
13. Chou, C.-H. (2004) Taxonomic status of the genus Myotis (Chiroptera: Vespertilionidae) in Taiwan. Department of Wildlife Ecology, Tunghai University, Tunghai, 115 pp. [in Chinese]
14. Corbet, G.B. & Hill, J.E. (1992) The mammals of the Indomalayan region: a systematic review. Oxford University Press, Oxford, 488 pp. http://dx.doi.org/10.1017/s0030605300020718
15. Csorba, G., Chou, C.-H., Ruedi, M., Görföl, T., Motokawa, M., Wiantoro, S., Thong, V.D., Son, N.T., Lin, L.-K. & Furey, N. (2014) The reds and the yellows: a review of Asian Chrysopteron Jentink, 1910 (Chiroptera: Vespertilionidae: Myotis). Journal of Mammalogy, 95, 663-678. http://dx.doi.org/10.1644/13-mamm-a-200
16. Csorba, G. & Lee, L.L. (1999) A new species of vespertilionid bat from Taiwan and a revision of the taxonomic status of Arielulus and Thainycteris (Chiroptera: Vespertilionidae). Journal of Zoology, 248, 361-367. http://dx.doi.org/10.1017/s0952836999007086
17. Dawson, M.R., Marivaux, L., Li, C.K., Beard, K.C. & Metais, G. (2006) Laonastes and the "Lazarus effect'' in recent mammals. Science, 311, 1456-1458.
18. Dobson, G.E. (1871) Notes on nine new species of Indian and Indo-Chinese Vespertilionidae, with remarks on the synonymy and classification of some other species of the same family. Proceedings of the Asiatic Society of Bengal, 1871, 210-215. http://dx.doi.org/10.1126/science.1124187
19. Dobson, G.E. (1878) Catalogue of the Chiroptera in the collection of the British Museum. Taylor & Francis, London, 567 pp. http://dx.doi.org/10.5962/bhl.title.55341
20. Dung, V.V., Gioa, P.M., Chinh, N.N., Tuoc, D., Arctander, P. & MacKinnon, J. (1993) A new species of living bovid from Vietnam. Nature, 363, 443-445. http://dx.doi.org/10.1038/363443a0
21. nd Edition. Alden Press, Oxford, 810 pp. http://dx.doi.org/10.1126/science.115.2990.431
22. Fenton, M.B. & Bogdanowicz, W. (2002) Relationships between external morphology and foraging behaviour: bats in the genus Myotis. Canadian Journal of Zoology, 80, 1004-1013. http://dx.doi.org/10.1139/z02-083
23. Findley, J.S. (1972) Phenetic relationships among bats of the genus Myotis. Systematic Zoology, 21, 31-52. http://dx.doi.org/10.2307/2412256
24. Francis, C., Borisenko, A., Ivanova, N., Eger, J., Lim, B., Guillen-Servent, A., Kruskop, S., Mackie, I. & Hebert, P. (2010) The role of DNA barcodes in understanding and conservation of mammal diversity in Southeast Asia. Plos One, 5, e12575. http://dx.doi.org/10.1371/journal.pone.0012575
25. Goodman, S.M., Maminirina, C.P., Weyeneth, N., Bradman, H.M., Christidis, L., Ruedi, M. & Appleton, B. (2009) The use of molecular and morphological characters to resolve the taxonomic identity of cryptic species: the case of Miniopterus manavi (Chiroptera: Miniopteridae). Zoologica Scripta, 38, 339-363. http://dx.doi.org/10.1111/j.1463-6409.2008.00377.x
26. Gray, J.E. (1846) Catalogue of the specimens and drawings of Mammalia and birds of the Nepal and Thibet presented by B.H. Hodgson, Esq. to the British Museum. Trustees of the British Museum, London, 156 pp. http://dx.doi.org/10.5962/bhl.title.32709
27. Han, N.J., Zhang, J.S., Reardon, T., Lin, L.K., Zhang, J.P. & Zhang, S.Y. (2010) Revalidation of Myotis taiwanensis Ärnbäck-Christie-Linde 1908 and its molecular relationship with M. adversus (Horsfield 1824) (Vespertilionidae, Chiroptera). Acta Chiropterologica, 12, 449-456. http://dx.doi.org/10.3161/150811010x538016
28. Hodgson, B.H. (1835) Synopsis of the Vespertilionidae of Nepal. Journal of the Asiatic Society of Bengal, 4, 699-701.
29. Horsfield, T. (1824) Zoological researches in Java, and the neighbouring islands. Kingsbury, Parbury & Allen, London, 328 pp. http://dx.doi.org/10.5962/bhl.title.44848
30. Howell, A.B. (1926) Three new mammals from China. Proceedings of the Biological Society of Washington, 39, 137-140.
31. Ibáñez, C., García-Mudarra, J.L., Ruedi, M., Stadelmann, B. & Juste, J. (2006) The Iberian contribution to cryptic diversity in European bats. Acta Chiropterologica, 8, 277-297. http://dx.doi.org/10.3161/1733-5329(2006)8[277:tictcd]2.0.co;2
32. Imaizumi, Y. (1962) List of type specimens of mammals which are present or had been present in the past in Japan. Proceedings of the Japanese Society of Systematic Zoology, 27, 6-10.
33. Irwin, D.M., Kocher, T.D. & Wilson, A.C. (1991) Evolution of the cytochrome b gene of mammals. Journal of Molecular Evolution, 32, 128-144. http://dx.doi.org/10.1007/bf02515385
34. Jenkins, P.D., Kilpatrick, C.W., Robinson, M.F. & Timmins, R.J. (2005) Morphological and molecular investigations of a new family, genus and species of rodent (Mammalia: Rodentia: Hystricognatha) from Lao PDR. Systematics and Biodiversity, 2, 419-454. http://dx.doi.org/10.1017/s1477200004001549
35. Jiang, T.L., Sun, K.P., Chou, C.H., Zhang, Z.Z. & Feng, J. (2010) First record of Myotis flavus (Chiroptera: Vespertilionidae) from mainland China and a reassessment of its taxonomic status. Zootaxa, 2414, 41-51.
36. Jones, G., Parsons, S., Zhang, S.Y., Stadelmann, B., Benda, P. & Ruedi, M. (2006) Echolocation calls, wing shape, diet and phylogenetic diagnosis of the endemic Chinese bat Myotis pequinius. Acta Chiropterologica, 8, 451-463. http://dx.doi.org/10.3161/1733-5329(2006)8[451:ecwsda]2.0.co;2
37. Kaneko, Y. & Maeda, K. (2002) A list of scientific names and the types of mammals published by Japanese researchers. Honyurui Kagaku (Mammalian Science), 42, 1-21.
38. Kawai, K., Nikaido, M., Harada, M., Matsumura, S., Lin, L.K., Wu, Y., Hasegawa, M. & Okada, N. (2003) The status of the Japanese and East Asian bats of the genus Myotis (Vespertilionidae) based on mitochondrial sequences. Molecular Phylogenetics and Evolution, 28, 297-307. http://dx.doi.org/10.1016/s1055-7903(03)00121-0
39. Kim, Y.M., Choi, E.H., Kim, S.K., Jang, K.H., Ryu, S.H. & Hwang, U.W. (2011) Complete mitochondrial genome of the Hodgson's bat Myotis formosus (Mammalia, Chiroptera, Vespertilionidae). Mitochondrial DNA, 22, 71-73. http://dx.doi.org/10.3109/19401736.2011.624598
40. Kishida, K. (1924) On the Formosan Chiroptera. Zoological Magazine, 36, 30-49. [Tokyo, in Japanese]
41. Kishida, K. (1932) Notes on a Formosan whiskered bat. Lansania, 4, 153-160.
42. Kitchener, A.C., Beaumont, M.A. & Richardson, D. (2006) Geographical variation in the clouded leopard, Neofelis nebulosa, reveals two species. Current Biology, 16, 2377-2383. http://dx.doi.org/10.1016/j.cub.2006.10.066
43. Koopman, K.F. (1994) Chiroptera: Systematics. In: Niethammer, J., Schliemann, H. & Starck, D. (Eds.), Handbuch der Zoologie. de Gruyter, Berlin, pp. 100-109.
44. Kruskop, S.V. & Borisenko, A.V. (2013) A new species of South-East Asian Myotis (Chiroptera: Vespertilionidae), with comments on Vietnamese 'Whiskered bats'. Acta Chiropterologica, 15, 293-305. http://dx.doi.org/10.3161/150811013x678937
45. Kruskop, S.V., Borisenko, A.V., Ivanova, N.V., Lim, B.K. & Eger, J.L. (2012) Genetic diversity of northeastern Palaearctic bats as revealed by DNA barcodes. Acta Chiropterologica, 14, 1-14. http://dx.doi.org/10.3161/150811012x654222
46. Kruskop, S.V. & Tsytsulina, K.A. (2001) A new big-footed mouse-eared bat Myotis annamiticus sp. nov. (Vespertilionidae, Chiroptera) from Vietnam. Mammalia, 65, 63-72. http://dx.doi.org/10.1515/mamm.2001.65.1.63
47. Kuo, H.C., Chen, S.F., Fang, Y.P., Flanders, J. & Rossiter, S.J. (2014) Comparative rangewide phylogeography of four endemic Taiwanese bat species. Molecular Ecology, 23, 3566-3586. http://dx.doi.org/10.1111/mec.12838
48. Kuo, H.C., Fang, Y.P., Csorba, G. & Lee, L.L. (2009) Three new species of Murina (Chiroptera: Vespertilionidae) from Taiwan. Journal of Mammalogy, 90, 980-991. http://dx.doi.org/10.1644/08-mamm-a-036.1
49. Kuroda, N. (1922) Notes on the mammal fauna of Tsushima and Iki Islands, Japan. Journal of Mammalogy, 3, 42-45.
50. Kuroda, N. (1935) Formosan Mammals preserved in the collection of Marquis Yamishina. Journal of Mammalogy, 16, 277-291. http://dx.doi.org/10.2307/1374636
51. Lack, J.B., Roehrs, Z.P., Stanley, C.E. Jr., Ruedi, M. & Van den Bussche, R.A. (2010) Molecular phylogenetics of Myotis indicate familial-level divergence for the genus Cistugo (Chiroptera). Journal of Mammalogy, 91, 976-992. http://dx.doi.org/10.1644/09-mamm-a-192.1
52. st ed.). National Museum of Natural Sciences, Taipei, 165 pp. [in Chinese]
53. nd ed.). National Museum of Natural Sciences, Taipei, 177 pp. [in Chinese]
54. Mao, X., Zhang, J., Zhang, S. & Rossiter, S.J. (2010a) Historical male-mediated introgression in horseshoe bats revealed by multilocus DNA sequence data. Molecular Ecology, 19, 1352-1366. http://dx.doi.org/10.1111/j.1365-294x.2010.04560.x
55. Mao, X.G., Zhu, G.J., Zhang, S. & Rossiter, S.J. (2010b) Pleistocene climatic cycling drives intra-specific diversification in the intermediate horseshoe bat (Rhinolophus affinis) in Southern China. Molecular Ecology, 19, 2754-2769. http://dx.doi.org/10.1111/j.1365-294x.2010.04704.x
56. Menu, H. & Sigé, B. (1971) Nyctalodontie et myotodontie, importants caractères de grades évolutifs chez les chiroptères entomophages. Comptes Rendus de l'Académie des Sciences de Paris, 272, 1735-1738.
57. Milne-Edwards, A. (1872) Mémoire de la faune mammalogique du Tibet Oriental et principalement de la principauté de Moupin. In: Milne-Edwards, H. (Ed), Recherches pour servir à l'histoire naturelle des mammifères comprenant des considérations sur la classification de ces animaux. G. Masson, Paris, pp. 231-304. http://dx.doi.org/10.5962/bhl.title.59889
58. Mori, T. (1928) Four new species of bats (Vespertilionidae) from Korea. Annotationes Zoologicae Japonensis, 11, 389-395.
59. Nylander, J.A.A. (2004) MrModeltest v. 2.3. Program distributed by the author. Evolutionary Biology Centre, Uppsala University. Available from: https://github.com/nylander/MrModeltest2 (accessed 21 October 2011)
60. Peters, W. (1870) Catalogue of Mammals of China. In: Swinhoe, R. (Ed.), Proceedings of the Zoological Society of London, 3, 615-653.
61. R-Core-Team (2014) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna. Available from: http://www.R-project.org (accessed 11 November 2014)
62. Rambaut, A. & Drummond, A.J. (2009) Tracer v1.5. Available from: http://beast.bio.ed.ac.uk/Tracer (accessed 17 September 2012)
63. Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D.L., Darling, A., Hohna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck, J.P. (2012) MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology, 61, 539-542. http://dx.doi.org/10.1093/sysbio/sys029
64. Ruedi, M., Biswas, J. & Csorba, G. (2012) Bats from the wet: two new species of Tube-nosed bats (Chiroptera: Vespertilionidae) from Meghalaya, India. Revue suisse de Zoologie, 119, 111-135.
65. Ruedi, M. & Mayer, F. (2001) Molecular systematics of bats of the genus Myotis (Vespertilionidae) suggests deterministic ecomorphological convergences. Molecular Phylogenetics and Evolution, 21, 436-448. http://dx.doi.org/10.1006/mpev.2001.1017
66. Ruedi, M., Stadelmann, B., Gager, Y., Douzery, E.J.P., Francis, C.M., Lin, L.-K., Guillén-Servent, A. & Cibois, A. (2013) Molecular phylogenetic reconstructions identify East Asia as the cradle for the evolution of the cosmopolitan genus Myotis (Mammalia, Chiroptera). Molecular Phylogenetics and Evolution, 69, 437-449. http://dx.doi.org/10.1016/j.ympev.2013.08.011
67. nd ed.). Cold Spring Harbor Laboratory, Cold Spring Harbor, 888 pp.
68. Shamel, H.H. (1944) A new Myotis from Formosa. Journal of Mammalogy, 25, 191-192.
69. Shamel, H.H. (1942) A collection of bats from Thailand (Siam). Journal of Mammalogy, 23, 317-328. http://dx.doi.org/10.2307/1375002
70. Sikes, R.S. & Gannon, W.L. (2011) Guidelines of the American Society of Mammalogists for the use of wild mammals in research. Journal of Mammalogy, 92, 235-253. http://dx.doi.org/10.1644/10-mamm-f-355.1
71. Silvestro, D. & Michalak, I. (2012) raxmlGUI: a graphical front-end for RAxML. Organisms Diversity & Evolution, 12, 335-337. http://dx.doi.org/10.1007/s13127-011-0056-0
72. Simmons, N.B. (2005) Order Chiroptera. In: Wilson, D.E. & Reeder, D.M. (Eds.), Mammal species of the world. A taxonomic and geographic reference. Johns Hopkins University Press, Washington, pp. 312-529.
73. Stadelmann, B., Herrera, G., Arroyo-Cabrales, J. & Ruedi, M. (2004a) Molecular systematics of the piscivorous bat Myotis (Pizonyx) vivesi. Journal of Mammalogy, 85, 133-139. http://dx.doi.org/10.1644/1545-1542(2004)085〈0133:msotfb〉2.0.co;2
74. Stadelmann, B., Jacobs, D., Schoeman, C. & Ruedi, M. (2004b) Phylogeny of African Myotis bats (Chiroptera, Vespertilionidae) inferred from cytochrome b sequences. Acta Chiropterologica, 6, 177-192. http://dx.doi.org/10.3161/001.006.0201
75. Stadelmann, B., Kunz, T.H., Lin, L.K. & Ruedi, M. (2007) Molecular phylogeny of New World Myotis (Chiroptera, Vespertilionidae) inferred from mitochondrial and nuclear DNA genes. Molecular Phylogenetics and Evolution, 43, 32-48. http://dx.doi.org/10.1016/j.ympev.2006.06.019
76. Stamatakis, A. (2006) RAxML-VI-HPC: Maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics, 22, 2688-2690. http://dx.doi.org/10.1093/bioinformatics/btl446
77. Swinhoe, R. (1870) Catalogue of the mammals of China (south of the River Yangtsze) and of the island of Formosa. Proceedings of the Zoological Society of London, 42, 615-652.
78. Tamura, K., Peterson, D., Peterson, N., Stecher, G., Nei, M. & Kumar, S. (2013) MEGA6: Molecular Evolutionary Genetics Analysis version 6.0. Molecular Biology and Evolution, 30, 2725-2729. http://dx.doi.org/10.1093/molbev/mst197
79. Tate, G.H. (1941) A review of the genus Myotis (Chiroptera) of Eurasia, with special reference to species occurring in the East Indies. Bulletin of the American Museum of Natural History, 78, 537-565.
80. Tate, G.H. (1947) Mammals of Eastern Asia. MacMillan Co, New York, pp. 45-66.
81. Thong, V.D., Puechmaille, S.J., Denzinger, A., Dietz, C., Csorba, G., Bates, P.J.J., Teeling, E.C. & Schnitzler, H.U. (2012) A new species of Hipposideros (Chiroptera: Hipposideridae) from Vietnam. Journal of Mammalogy, 93, 1-11. http://dx.doi.org/10.1644/11-mamm-a-073.1
82. Tiunov, M.P., Kruskop, S.V. & Feng, J. (2011) A new mouse-eared bat (Mammalia: Chiroptera, Vespertilionidae) from South China. Acta Chiropterologica, 13, 271-278. http://dx.doi.org/10.3161/150811011x624758
83. Tomes, R.F. (1858) On the characters of four species of bat inhabiting Europe and Asia, and the description of a new species of Vespertilio inhabiting Madagascar. Proceedings of the Zoological Society London, 1858, 78-90.
84. Tomes, R.F. (1859) Descriptions of six hitherto undescribed species of bats. Proceedings of the Zoological Society of London, 1859, 68-79.
85. Topál, G. (1997) A new mouse-eared bat species, from Nepal, with statistical analyses of some other species of subgenus Leuconoe (Chiroptera, Vespertilionidae). Acta Zoologica Academiae Scientiarum Hungaricae, 43, 375-402.
86. Trouessart, E.-L. (1878) Catalogue des mammifères vivants et fossiles. Chiroptera. Revue et magazine de zoologie pure et appliquée, 3, 200-254.
87. Trouessart, E.-L. (1897) Catalogus mammalium tam viventium quam fossilium. Fasciculus I.-II. Friedländer and Sohn, Berlin, 1469 pp.
88. Tsytsulina, K., Dick, M.H., Maeda, K. & Masuda, R. (2012) Systematics and phylogeography of the steppe whiskered bat Myotis aurascens Kuzyakin, 1935 (Chiroptera, Vespertilionidae). Russian Journal of Theriology, 11, 1-20.
89. Tsytsulina, K. & Strelkov, P.P. (2001) Taxonomy of the Myotis frater species group (Vespertilionidae, Chiroptera). Bonner zoologischer Beiträge, 50, 15-26.
90. Weyeneth, N., Goodman, S.M. & Ruedi, M. (2011) Do diversification models of Madagascar's biota explain the population structure of the endemic bat Myotis goudoti (Chiroptera: Vespertilionidae)? Journal of Biogeography, 38, 44-54. http://dx.doi.org/10.1111/j.1365-2699.2010.02393.x
91. Worthington-Wilmer, J. & Barratt, E. (1996) A non-lethal method of tissue sampling for genetic studies of Chiropterans. Bat Research News, 37, 1-3.
92. Yoshiyuki, M. (1989) A systematic study of the Japanese chiroptera. National Science Museum monographs, 7, 1-242.
93. Zhang, Z.Z., Tan, X.Y., Sun, K.P., Liu, S., Xu, L.J. & Feng, J. (2009) Molecular systematics of the Chinese Myotis (Chiroptera, Vespertilionidae) inferred from cytochrome-b sequences. Mammalia, 73, 323-330. http://dx.doi.org/10.1515/mamm.2009.058
94. Ziegler, R. (2003) Bats (Chiroptera, Mammalia) from Middle Miocene karstic fissure fillings of Petersbuch near Eichstätt, Southern Franconian Alb (Bavaria). Geobios, 36, 447-490. http://dx.doi.org/10.1016/s0016-6995(03)00043-3