Summary
Between 1974 and 1978 structure and diversity of shrublands and woodlands of northern Israel were studied along climatic and human-disturbance gradients using 0.1 ha vegetation samples. Diversity increased along the moisture gradient, with highest woody and herb species richness in open Pistacia shrubland on the xeric border of the Mediterranean region, and highest equitability and lowest dominance concentration in sub-humid, moderately grazed, open oak woodlands. Semi-open disturbed shrublands were rich in herbs and had much higher structural, plant species, and animal species diversities than the closed, mature, ‘climax’ maquis. Diversity showed a two-slope response to grazing with highest species numbers in heavily (but not the most severely) grazed woodlands and shrublands. These communities have some of the highest plant alpha diversities in the world; the richness of their floras (especially in annual plants) is the product of relatively rapid evolution under stress by drought, fire, grazing, and cutting.
Comparative data on diversity and growth-form composition are compiled for mediterranean communities: Israeli' shrublands and woodlands, California chaparral and woodlands, Chilean matorral, South African fynbos, and Australian heath and mallee. Communities of three of these areas are of more recent (primarily Pleistocene) development and share some similarities; these threc form a sequence (California, Chile, and Old World Mediter-ranean) of increasing length of human disturbance and consequent species diversity. The southwest Australian heath or kwongan and the South African fynbos are, in contrast, derived from ancient Gondwanan heath like communities and are adapted to very old, nutrient-poor soils. The Gondwanan communities are quite different in growth-form structure and soil and nutrient relationships from communities of the three more recent mediterranean areas; the Gondwanan communities are almost lacking in annual species and are exceedingly rich in woody species. The richest temperate plant communities known — grazed Mediterranean pastures vs. fynbos and Australian heath — are in almost polar contrast in their growth-form structures and the bases of their species diversities.
This study, sponsored by the U.S.-Israel Binational Science Foundation (No. 450), was carried out by Z. Naveh as principal investigator with R.H. Whittaker as American collaborator. We thank Mr. A. Mann, S. Burmil, Mrs. Chaim, and Mrs. A. Kleen for botanical field work and statistical computations, Mr. D. Feigin and S. Ben Ezrah for technical assistance, Mr. S. Asherow for identification of young plants, and the Nature Reserve Authorities, the Neve Yaar Experimental Station, the Agricultural School Kfar Hanoar Hadati, and Kibbutzim Allonim and Allone Abba for allowing us to use their land for this study. The work by R.H. Whittaker was supported by the U.S. National Science Foundation, Australian National University, and Canberra Botanic Gardens; and we thank all the collaborators in this work.
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References
Aloni, R. 1972. The vegetation of the Lower Galil. M. Sc. Thesis, Hebrew University, Jerusalem. (Hebrew). 128 pp.
Aschmann, H. 1973. Distribution and peculiarity of mediterranean ecosystems. In: F. di Castri & H.A. Mooney. (eds). Mediterranean Type Ecosystems. Ecol. Stud. 7: 11–19.
Aschmann, H. & C. Bahre. 1977. Man's impact on the wild landscape. In: H.A. Mooney (ed.) Convergent Evolution in Chile and California, US/IBP Synthe. Ser. 5: 73–84.
Auclair A.N. & F.G. Goff. 1971. Diversity relations of upland forests in the western Great Lakes area. Amer. Nat. 105: 499–528.
Axelrod D.I. & P.H. Raven. 1978. Late Cretaceous and Tertiary vegetation history of Africa, in: M.J.A. Werger (ed.) Biogeography and Ecology of Southern Africa, p. 77–130. Junk, The Hague.
Bazzaz F.A. 1975. Plant species diversity in old-field successional ecosystems in southern Illinois. Ecology 56: 485–488.
Beard J.S. 1976. An indigenous term for the western Australian sandplain and its vegetation. J.R. Soc. W. Austr. 59: 55–57.
Burbidge N.T. 1960. The phytogeography of the Australian region. Austr. J. Bot. 8: 75–209.
Cain S.A. 1950. Life-forms and phytoclimate. Bot. Rev. 16: 1–33.
Castri F.di & H.A. Mooney, eds. 1973. Mediterranean Type Ecosystems: Origin and Structure. Ecol. Stud. 7: 1–405. Springer, New York.
Christensen N.L. & C.H. Muller. 1975. Effects of fire on factors controlling plant growth in Adenostoma chaparral. Ecol. Monogr. 45: 29–55.
Cody M.L. 1975. Towards a theory of continental species diversities: bird distributions over mediterranean habitat gradients, pp. 214–257 in: M.L. Cody & J.M. Diamond (eds.). Ecology and Evolution of Communities, Harvard Univ. Press, Cambridge.
Cody M.L. & H.A. Mooney. 1978. Convergence versus non-convergence in mediterranean-climate ecosystems. Ann. Rev. Ecol. Syst. 9: 265–321.
Connell J.H. & E. Orias. 1964. The ecological regulation of species diversity. Amer. Nat. 98: 399–414.
Cowling, R. & B. Campbell. 1980. Convergence in vegetation structure in the mediterranean communities of California, Chile, and South Africa. Vegetatio (in press).
Eig A. 1933. A historical-phytosociological essay on Palestinian forests of Quercus aegilops L. ssp. ithaburensis (Desc.) in past and present. Beih, Bot. Contr. 51, Abt. II: 225–272.
Glenn-Lewin D.C. 1977. Species diversity in North American temperate forests. Vegetatio 33: 153–162.
Good R. 1974. The Geography of the Flowering Plants. 4th ed. Longmans, London.
Grassle J.F. & H.L. Sanders. 1973. Life histories and the role of disturbance. Deep-Sea Res. 20: 643–659.
Grime J.P. 1973. Control of species density in herbaceous vegetation. J. Environ. Mangt. 1: 151–167.
Grime J.P. 1979. Plant strategies and vegetation processes. Wiley, New York. 222 pp.
Harner R.F. & K.T. Harper. 1976. The role of area, heterogeneity, and favorability in plant species diversity of pinyonjuniper ecosystems. Ecology 57: 1254–1263.
Harper, J.L. 1969. The role of predation in vegetational diversity. In: G.M. Woodwell (ed.). Diversity and stability in ecological systems, Brookhaven Symp. Biol. 22: 48–62.
Hill M.O. 1973. Diversity and evenness: a unifying notation and its consequences. Ecology 54: 427–432.
Hopper S.D. 1979. Biogeographical aspects of speciation in the southwest Australian flora. Ann. Rev. Ecol. Syst. 10: 399–422.
Karr J.R. & R.R. Roth. 1971. Vegetation structure and avian diversity in several New World areas. Amer. Nat. 105: 423–435.
Lamont B.B., S. Downes, & J.E.D. Fox. 1977. Importance-value curves and diversity indices applied to a species-rich heathland in western Australia. Nature 265: 438–441.
Levyns M.R. 1964. Migrations and origin of the Cape flora. Trans. R. Soc. S. Afr. 37: 85–107.
Loucks O.L. 1970. Evolution of diversity, efficiency, and community stability. Amer. Zool. 10: 17–25.
Lubchenco J. 1978. Plant species diversity in a marine intertidal community: importance of herbivore food preference and algal competitive abilities. Amer. Nat. 112: 23–39.
Maarel E.van der. 1975. Man-made natural ecosystems in environmental management and planning, in: W.H.van Dobben & R.H. Lowe-McConnell (eds.) Unifying Concepts in Ecology: Rept. Plen. Sess., 1st Int. Congr. Ecol., The Hague, 1974, Junk, The Hague, & Pudoc, Wageningen.
MacArthur R.H. 1964. Environmental factors affecting bird species diversity. Amer. Nat. 98: 387–397.
MacArthur R.H. 1965. Patterns of species diversity. Biol. Rev. 40: 510–533.
Margalef R. 1968. Perspectives in Ecology Theory. Univ. Chicago Press, Chicago. 111 pp.
May R.M. 1975. Patterns of species abundance and diversity in: M.L. Cody & J.M. Diamond (ed.) Ecology and Evolution of Communities p. 81–120, Harvard Univ. Press, Cambridge.
McNaughton S.J. 1968. Structure and function in California grasslands Ecology 49: 962–972.
McPherson J.K. & C.H. Muller. 1969. Allelopathic effects of Adenostoma fasciculatum, ‘chamise’, in the California chaparral. Ecol. Monogr. 39: 177–198.
Miller, P.C., D.E. Bradbury, E. Hajek, V. LaMarche, & N.J.W. Thrower. 1977. Past and present environment. In: H.A. Mooney. (ed.) Convergent Evolution in Chile and California, US/IBP Synth. Ser. 5: 27–72.
Mooney H.A., ed. 1977. Convergent Evolution in Chile and California: Mediterranean Climate Ecosystems. US/IBP Synthesis Series 5: 1–224. Dowden, Hutchinson & Ross, Stroudsburg, Pa.
Mooney, H.A., et al. 1977. The producers — their resources and adaptive responses. In: Convergent Evolution in Chile and California, ed. H.A. Mooney. US/IBP Synth. Ser. 5: 85–143.
Moral R.del. 1972. Diversity patterns in forest vegetation of the Wenatchee Mountains, Washington. Bull. Torrey Bot. Club 99: 57–64.
Morris M.G. 1973. The effects of seasonal grazing on the Heteroptera and Auchenorhyncha (Hemiptera) of chalk grasslands. J. Appl. Ecol. 10: 761–780.
Muller C.H., R.B. Hanawalt, & J.K. McPherson. 1968. Allelopathie control of herb growth in the fire cycle of California chaparral. Bull. Torrey Bot. Club 95: 225–231.
Murdoch W.W., F.C. Evans, & C.H. Peterson. 1972. Diversity and pattern in plants and insects. Ecology 53: 819–829.
Naveh Z. 1955. Some aspects of range improvement in a mediterranean environment. J. Range Mangt. 8: 265–270.
Naveh Z. 1967. Mediterranean ecosystems and vegetation types in California and Isreal. Ecology 48: 445–459.
Naveh, Z. 1971. The conservation of ecological diversity of mediterranean ecosystems through ecological management. In: E. Duffey & A.S. Watt (eds.) The Scientific Management of Animal and Plant Communities for Conservation, Symp. Brit. Ecol. Soc. 11: 605–622.
Naveh Z. 1975. The evolutionary signiticance of fire in the Mediterranean Region. Vegetatio 29: 199–208.
Naveh, Z. 1977. The role of fire in the mediterranean landscape of Israel, in: Proc. Symp. Environmental Consequences of Fire and Fuel Management in Mediterranean Ecosystems. Palo Alto, Calif., Aug. 1977, p. 299–306.
Naveh, Z. & J. Dan. 1973. Human degradation of mediterranean landscapes in Israel. In: F. de Castri & H.A. Mooney (eds). Mediterranean Type Ecosystems, Ecol. Stud. 7: 373–390.
Naveh, Z. & J. Kinsky. 1975. The effect of climate and management on species diversity of Tabor oak savanna pastures. Proc. 6th Sci. Conf. Isr. Ecol. Soc., p. 284–296.
Naveh Z. & R.H. Whittaker. 1980. Measurements and relationships of plant species diversity in Mediterranean shrublands and woodlands. In: F. Grassle, G.P. Patil, W. Smith, & C. Taillee (eds.) Ecological Diversity in Theory and Practice, Statistical Ecology Volume S6. Internat. Coop. Publ. Housé, Fairland, Md.: 219–239.
Odum E.P. 1969. The strategy of ecosystem development. Science 164: 262–270.
Parsons D.J. & A.R. Moldenke. 1975. Convergence in vegetation structure along analogous climatic gradients in California and Chile. Ecology 56: 950–957.
Peet R.K. 1974. The measurement of species diversity. Ann. Rev. Ecol. Syst. 5: 285–307.
Peet R.K. 1978. Forest vegetation of the Colorado Front Range: patterns of species diversity. Vegetatio 37: 65–78.
Pickett S.T.A. 1976. Succession: an evolutionary interpretation. Amer. Nat. 110: 107–119.
Raunkiaer C. 1934. The Life Forms of Plants and Statistical Plant Geography. Clarendon, Oxford. 632 pp.
Raven, P.H. 1973. The evolution of mediterranean floras. In: F. di Castri & H.A. Mooney (eds.). Mediterranean Type Ecosystems, Ecol. Stud. 7: 212–224.
Solbrig, O.T., M.L. Cody, E.R. Fuentes, W. Glanz, J. H. Hunt, & A.R. Moldenke. 1977. The origin of the biota. In: H.A. Mooney (ed.). Convergent Evolution in Chile and California, US/IBP Synth. Ser. 5: 13–26.
Taylor H.C. 1972. Fynbos. Veld & Flora 2: 68–75.
Taylor, H.C. 1977. The Cape floral kingdom: an ecological view, pp. 19–33 in: Proc. 2n Natn. Weeds Conf. S. Afr.
Taylor H.C. 1978. Capensis, in: M.J.A. Werger (ed) Biogeography and Ecology of Southern Africa, p. 171–229 Junk, The Hague.
Warburg M. 1977. Plant and animal species diversity along environmental gradients in a Mediterranean landscare of Israel. (B.S.F. No. 450). Animal species diversity. Technion, 1sr. Inst. Technol., Haifa, Israel. (Mimeographed).
Warburg M.R., A. Ben-Horin, & D. Rankevich. 1978. Rodent species diversity in mesic and xeric habitats in the Mediterranean region of northern Israel. J. Arid Environ. 1: 63–69.
Werger M.J.A. 1972. Species-area relationship and plot size; with some examples from South African vegetation. Bothalia 10: 583–594.
Werger M.J.A., F.J. Kruger, & H.C. Taylor. 1972. Pflanzensoziologische Studie der Fynbosvegetation am Kap der Guten Hoffnung, (Engl. summ.). Vegetatio 24: 71–89.
Westman W.E. 1975. Pattern and diversity in swamp and dune vegetation, North Stradbroke Island. Aust. J. Bot. 23: 339–354.
Whittaker R.H. 1972. Evolution and measurement of species diversity. Taxon 21: 213–251.
Whittaker R.H. 1975. Communities and Ecosystems. 2nd ed. Macmillan, New York. 385 pp.
Whittaker R.H. 1977a. Evolution of species diversity in land communities. Evol. Biol. 10: 1–67.
Whittaker R.H. 1977b. Animal effects on plant species diversity, pp. 409–425 in: Vegetation und Fauna, ed. R. Tüxen. Ber. Symp. Int. Ver. Vegetskunde, Rinteln, 1976. Cramer, Vaduz.
Whittaker R.H. & Z. Naveh. 1980. Analysis of two-phase patterns. In: G.P. Patil & M. Rosenzweig (ed.) Contemporary Quantitative Ecology and Related Ecometrics, Statistical Ecology Volume S12. Internat. Coop. Publ. House, Fairland, Md.: 157–165.
Whittaker R.H. & W.A. Niering. 1965. Vegetation of the Santa Catalina Mountains, Arizona. (II). A gradient analysis of the south slope. Ecology 46: 429–452.
Whittaker R.H. & W.A. Niering. 1975. Vegetation of the Santa Catalina Mountains, Arizona. V. Biomass, production, and diversity along the elevation gradient. Ecology 56: 771–790
Whittaker R.H., W.A. Niering, & M.D. Crisp. 1979. Structure, pattern, and diversity of a mallee community in New South Wales. Vegetatio 39: 65–76.
Willson M.F. 1974. Avian community organization and habitat structure. Ecology 55: 1017–1029.
Zeevalking H.J. & L.F.M. Fresco. 1977. Rabbit grazing and species diversity in a dune area. Vegetatio 35: 193–196.
Zinderen BakkerSr., E.M.van. 1976. The evolution of Late-Quaternary palaeoclimates of southern Africa. Palaeoecology of Africa 9: 160–202.
Zohary M. 1962. Plant Life of Palestine: Israel and Jordan. Ronald Press, New York. 262 pp.
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Nomenclature follows Zohary et al. (1948).
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Naveh, Z., Whittaker, R.H. Structural and floristic diversity of shrublands and woodlands in Northern Israel and other Mediterranean areas. Vegetatio 41, 171–190 (1980). https://doi.org/10.1007/BF00052445
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DOI: https://doi.org/10.1007/BF00052445