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A revision of Martin's seed classification system, with particular reference to his dwarf-seed type

Published online by Cambridge University Press:  01 March 2007

Carol C. Baskin*
Department of Biology, University of Kentucky, Lexington, KY 40506-0225, USA Department of Plant and Soil Sciences, University of Kentucky, Lexington, KY 40546-0312, USA
Jerry M. Baskin
Department of Biology, University of Kentucky, Lexington, KY 40506-0225, USA
*Correspondence: Fax: +1 859 257 1717 Email:


Martin's (1946) seed classification system has 10 types based on embryo and endosperm characteristics and two additional types based on seed size: dwarf (0.3–2.0 mm) and micro ( ≤ 0.2 mm). He listed 17 families and 12 genera (in five other families) as having dwarf seeds. Our recent discovery of morphophysiological dormancy in dwarf seeds of several taxa of Campanulaceae and one of Gentianaceae prompted an evaluation of dwarf seeds. Martin's paper contains 37 families with one to several small (0.3–2.0 mm) seeded species that he did not list as being dwarf. Comparison of Martin's dwarf families and the 37 small-seeded non-dwarf families revealed no consistent differences between the two groups in endosperm texture, seed-coat anatomy, embryo morphology, class of seed dormancy or phylogenetic position. Also, Martin's dwarf seeds include a variety of embryo morphologies. Consequently, we have revised Martin's key to seed types. The dwarf category has been removed and the micro category replaced by ‘undifferentiated’ to reflect the state of the embryo in fresh seeds. Further, the key now includes linear fully developed, linear underdeveloped, spatulate fully developed and spatulate underdeveloped seed types, which Martin illustrated but did not include in his key. In the revised key, all seeds are distinguished on the basis of embryo and endosperm characteristics.

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Copyright © Cambridge University Press 2007

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Adams, C.A., Baskin, J.M. and Baskin, C.C. (2005) Trait stasis versus adaptation in disjunct relict species: Evolutionary changes in seed dormancy-breaking and germination requirements in a subclade of Aristolochia subgenus Siphisia (Piperales). Seed Science Research 15, 161173.CrossRefGoogle Scholar
APG II (Angiosperm Phylogeny Group) (2003) An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II. Botanical Journal of the Linnean Society 141, 399436.CrossRefGoogle Scholar
Baskin, C.C. and Baskin, J.M. (1998) Seeds: Ecology, biogeography, and evolution of dormancy and germination. San Diego, Academic Press.Google Scholar
Baskin, C.C. and Baskin, J.M. (2005) Underdeveloped embryos in dwarf seeds and implications for assignment to dormancy class. Seed Science Research 15, 357360.CrossRefGoogle Scholar
Baskin, C.C., Meyer, S.E. and Baskin, J.M. (1995) Two types of morphophysiological dormancy in seeds of two genera (Osmorhiza and Erythronium) with an Arcto-Tertiary distribution pattern. American Journal of Botany 82, 293298.CrossRefGoogle Scholar
Baskin, C.C., Milberg, P., Andersson, L. and Baskin, J.M. (2002) Non-deep simple morphophysiological dormancy in seeds of the weedy facultative winter annual Papaver rhoeas. Weed Research 42, 194202.CrossRefGoogle Scholar
Baskin, C.C., Baskin, J.M. and Yoshinaga, A. (2005) Morphophysiological dormancy in seeds of six endemic lobelioid shrubs (Campanulaceae) from the montane zone in Hawaii. Canadian Journal of Botany 83, 16301637.CrossRefGoogle Scholar
Baskin, J.M. and Baskin, C.C. (1984) Germination ecophysiology of an eastern deciduous forest herb, Stylophorum diphyllum. The American Midland Naturalist 111, 390399.CrossRefGoogle Scholar
Baskin, J.M. and Baskin, C.C. (1994) Nondeep simple morphophysiological dormancy in seeds of the mesic woodland winter annual Corydalis flavula (Fumariaceae). Bulletin of the Torrey Botanical Club 121, 4046.CrossRefGoogle Scholar
Baskin, J.M. and Baskin, C.C. (2004) A classification system for seed dormancy. Seed Science Research 14, 116.CrossRefGoogle Scholar
Baskin, J.M., Baskin, C.C. and Li, X. (2000) Taxonomy, anatomy and evolution of physical dormancy in seeds. Plant Species Biology 15, 139152.CrossRefGoogle Scholar
Baskin, J.M., Baskin, C.C. and Dixon, K.W. (2006) Physical dormancy in the endemic Australian genus Stylobasium, a first report for the family Surianaceae (Fabales). Seed Science Research 16, 229232.CrossRefGoogle Scholar
Boesewinkel, F.D. (1984) Ovule and seed structure in Datiscaceae. Acta Botanica Neerlandica 33, 419429.CrossRefGoogle Scholar
Bregman, R. and Bouman, F. (1983) Seed germination in Cactaceae. Botanical Journal of the Linnean Society 86, 357374.CrossRefGoogle Scholar
Compilation Committee (2000) Seeds of woody plants in China (in Chinese). Beijing, China Forestry Press.Google Scholar
Coops, H. and van der Velde, G. (1995) Seed dispersal, germination and seedling growth of six helophyte species in relation to water-level zonation. Freshwater Biology 34, 1320.CrossRefGoogle Scholar
Corner, E.J.H. (1976) The seeds of dicotyledons. Vols I and II. Cambridge, Cambridge University Press.Google Scholar
da Silva, E.A.A., Toorop, P.E., van Aelst, A.C. and Hilhorst, H.W.M. (2004) Abscisic acid controls embryo growth potential and endosperm cap weakening during coffee (Coffea arabica cv. Rubi) seed germination. Planta 220, 251261.CrossRefGoogle ScholarPubMed
Ellison, A.M. (2001) Interspecific and intraspecific variation in seed size and germination requirements of Sarracenia (Sarraceniaceae). American Journal of Botany 88, 429437.CrossRefGoogle ScholarPubMed
Finch-Savage, W.E. and Leubner-Metzger, G. (2006) Seed dormancy and the control of germination. New Phytologist 171, 501523.CrossRefGoogle ScholarPubMed
Forbis, T.A. and Diggle, P.K. (2001) Subnivean embryo development in the alpine herb Caltha leptosepala (Ranunculaceae). Canadian Journal of Botany 79, 635642.CrossRefGoogle Scholar
Forbis, T.A., Floyd, S.K. and de Queiroz, A. (2002) The evolution of embryo size in angiosperms and other seed plants: Implications for the evolution of seed dormancy. Evolution 56, 21122125.Google ScholarPubMed
Grushvitzky, I.V. (1967) After-ripening of seeds of primitive tribes of angiosperms, conditions and peculiarities. pp. 329336+ figures 1–8 in Borriss, H. (Ed.) Physiologie, ökologie und biochemie der keimung. Greifswald, Germany, Ernst-Moritz-Arndt-Universitat.Google Scholar
Hewett, D.G. (1964) Menyanthes trifoliata L. Journal of Ecology 52, 723735.CrossRefGoogle Scholar
Hidayati, S.N., Baskin, J.M. and Baskin, C.C. (2000a) Dormancy-breaking and germination requirements for seeds of Diervilla lonicera (Caprifoliaceae), a species with underdeveloped linear embryos. Canadian Journal of Botany 78, 11991205.CrossRefGoogle Scholar
Hidayati, S.N., Baskin, J.M. and Baskin, C.C. (2000b) Morphophysiological dormancy in seeds of two North American and one Eurasian species of Sambucus (Caprifoliaceae) with underdeveloped spatulate embryos. American Journal of Botany 87, 16691678.CrossRefGoogle ScholarPubMed
Hidayati, S.N., Baskin, J.M. and Baskin, C.C. (2000c) Dormancy-breaking and germination requirements of seeds of four Lonicera species (Caprifoliaceae) with underdeveloped spatulate embryos. Seed Science Research 10, 459469.CrossRefGoogle Scholar
Hidayati, S.N., Baskin, J.M. and Baskin, C.C. (2005) Epicotyl dormancy in Viburnum acerifolium (Caprifoliaceae). The American Midland Naturalist 153, 232244.CrossRefGoogle Scholar
Horovitz, A., Bullowa, S. and Negbi, M. (1975) Germination characters in wild and cultivated Anemone coronaria L. Euphytica 24, 213220.CrossRefGoogle Scholar
Jacques-Félix, H. (1977) La graine et l'embryon chez Memecylon (Mélastomatacées) Africains. Adansonia (Series 2). 17, 193200.Google Scholar
Karlsson, L.M., Tamado, T. and Milberg, P. (2003) Seed dormancy pattern of the annuals Argemone ochroleuca and A. mexicana (Papaveraceae). Flora 198, 329339.CrossRefGoogle Scholar
Kondo, T., Okubo, N., Miura, T., Baskin, C.C. and Baskin, J.M. (2005) Ecophysiology of seed dormancy and germination in the mesic woodland herbaceous perennial Corydalis ambigua (Fumariaceae) in Japan. Canadian Journal of Botany 83, 571578.CrossRefGoogle Scholar
Kondo, T., Sato, C., Baskin, J.M. and Baskin, C.C. (2006) Post-dispersal embryo development, germination phenology, and seed dormancy in Cardiocrinum cordatum var. glehnii (Liliaceae s. str.), a perennial herb of the broadleaved deciduous forest in Japan. American Journal of Botany 93, 849859.CrossRefGoogle ScholarPubMed
Martin, A.C. (1946) The comparative internal morphology of seeds. The American Midland Naturalist 36, 513660.CrossRefGoogle Scholar
Moles, A.T., Ackerly, D.D., Webb, C.O., Tweddle, J.C., Dickie, J.B. and Westoby, M. (2005) A brief history of seed size. Science 307, 576580.CrossRefGoogle ScholarPubMed
Morgan, M.D. (1990) Seed germination characteristics of Iris virginica. The American Midland Naturalist 124, 209213.CrossRefGoogle Scholar
Netolitzky, F. (1926) Anatomie der Angiospermen-samen. Handbuch der Pflanzenanatomie. Vol. 10. Berlin, Verlag von Gebrüder Borntraeger.Google Scholar
Nikolaeva, M.G. (2004) On criteria to use in studies of seed evolution. Seed Science Research 14, 315320.CrossRefGoogle Scholar
Nikolaeva, M.G., Rasumova, M.V. and Gladkova, V.N. (1985) Reference book on dormant seed germination (in Russian). Leningrad, Nauka Publishers, Leningrad Branch.Google Scholar
Scatena, V.L. and Bouman, F. (2001) Embryology and seed development of Paepalanthus sect. Actinocephalus (Koern.) Ruhland (Eriocaulaceae). Plant Biology 3, 341350.CrossRefGoogle Scholar
Shipley, B. and Parent, M. (1991) Germination responses of 64 wetland species in relation to seed size, minimum time to reproduction and seedling relative growth rate. Functional Ecology 5, 111118.CrossRefGoogle Scholar
Spongberg, S.A. (1978) The genera of Crassulaceae in the southeastern United States. Journal of the Arnold Arboretum 59, 197244.CrossRefGoogle Scholar
Takhtajan, A. (1991) Anatomia seminum comparativa. Vol. 3. Dicotyledones. Caryophyllidae – Dilleniidae. Leningrad, Nauka Sectio Leninopoli.Google Scholar
Takhtajan, A. (1992) Anatomia seminum comparativa. Vol. 4. Dicotyledones. Dilleniidae. St Petersburg, Nauka Sectio Petropoli.Google Scholar
Takhtajan, A. (1996) Anatomia seminum comparativa. Vol. 5. Dicotyledones. Rosidae I. St. Petersburg, Mir et Semja.Google Scholar
Teryokhin, E.S. (2001) The origin of ‘dust’ seeds in parasitic and mycoparasitic angiosperms: A hypothesis for symbioses. Beitrage zur Biologie der Pflanzen 72, 381397.Google Scholar
Walck, J.L., Baskin, C.C. and Baskin, J.M. (1999) Seeds of Thalictrum mirabile (Ranunculaceae) require cold stratification for loss of nondeep simple morphophysiological dormancy. Canadian Journal of Botany 77, 17691776.CrossRefGoogle Scholar
Watson, L. and Dallwitz, M.J. (1992 onwards) The families of flowering plants: Descriptions, illustrations, identification, and information retrieval. Version: 14 December 2000. . Currently available athttp://www. Scholar
Young, J.A. and Young, C.G. (1992) Seeds of woody plants in North America (revised and enlarged edition). Portland, Dioscorides Press.Google Scholar