Mediterranean landscape evolution and degradation as multivariate
Transcription
Mediterranean landscape evolution and degradation as multivariate
Landscape Planning, 9 (1982) 125-146 Elsevier Scientific Publishing Company, Amsterdam - Printed in The Netherlands MEDITERRANEAN LANDSCAPE EVOLUTION AND DEGRADATION MULTIVARIATE BIOFUNCTIONS: THEORETICAL AND PRACTICAL IMPLICATIONS 125 AS 2. NAVEI-I Faculty of Agricultural (Israef) (Accepted Engineering, Technion-Israel Znstitute of Technology, Haifa 3200 30 March 1982) ABSTRACT Naveh, Z., 1982. Mediterranean landscape evolution and degradation as multivariate biofunctions: theoretical and practical implications. Landscape Plann., 9: 125-146. The evolution of the natural Mediterranean landscape is described as a multivariate function of independent initial and driving ecosystem state factors on their dependent soil and biotic variables. The increasingly dominant role of man as controlling state factor, by hunting and gathering and burning in the Upper Pleistocene and by agro-pastoral land uses during historical times, has turned these functions into anthropogenic biofunctions. In the latter, the introduction of man-made cultural artifacts as dependent variables has created the cultural Mediterranean landscape. The detrimental effects of disrupting these traditional agro-pastoral functions, by either increasing or completely releasing the defoliation pressures of burning, grazing and cutting and by monospecies pine afforestation in present neotechnological degradation functions, are demonstrated by quantifying the biotic variables of species composition, diversity and vegetation structure. Mediterranean ecosystems in California, in which these agro-pastoral biofunctions were introduced abruptly only in recent decades, have not only lower diversity but are lacking the adaptive agro-pastoral resilience acquired by the vegetation in the Mediterranean area. They are, therefore, more vulnerable and have succumbed to chiefly autogameous-weedy invaders which were unintentionally introduced from the Mediterranean at the time of Spanish settlement. It is concluded that the conservation of Medite~anean landscapes and their organic variety can be ensured by continuation and/or simulation of the agro-pastoral functions under which these landscapes evolved, thereby maintaining their dynamic evolutionary flow equilibrium within closely interwoven networks of multiple land-use patterns. KNTRODUCTION In the description of the evolution of cultural landscapes, long- and shortterm ecosystem dynamics should be coupled with historic events. This can be achieved by using the functional-factoral approach of Jenny’s (1958,1961, 1980) state factor equations. This approach has been introduced into plant 0304-3924/82/000~000/$02,75 o 1982 Elsevier Scientific Publishing Company 126 ecology by Major (1951), showing that vegetation (V), like soil (S) in Jenny’s soil formation (or pedogenic) equation, is dependent on the same five mathematic~ly-independent groups of factors, namely: parent material @) from which the soil originated; relief (P) or topography, referring to the slope exposure; regional climate (cl); organisms (0) or biota-flora, fauna and the human factor; and time (t) or the duration of these processes. Therefore, vegetation, like soil, can be presented as a dependent variable of the function v = fQ-v,cf,o,t) Jenny (1961) extended this equation to all dependent ecosystem properties which are related to or are a function of three major state factor groups : (1) initial state of the system or site conditions of p and r at time t = 0; (2) external flux potentials: the driving ecosystem forces of climate and biotic factors and other unspecified flux potentials, denoted by dots . . ,; and (3) the age of the system t. By differentiating these state factor equations, Jenny and Major showed that soil and vegetation and any other dependent ecosystem variables can be approximated and therefore ‘solved’ as a function or a sequence of only one major influencing independent factor. This is the case when this factor varies greatly in comp~ison with the others or the latter may be of relatively little importance in determining differences in these properties so that, save for the one function, the slopes of the others are nearly zero. In his recent book, Jenny (1980) treated the above-mentioned five groups of state factors as soil and ecosystem functions, according to the dominant and subordinate factors, as climofunction (or climosequence), biofunction, topofunction, lithofunction, chronofunction and dotfunction. Thus, a biofunction (or biosequence) of the land ecosystem (I) and its soil (s), vegetation (u) and animal (a) properties is presented as l,v,a,s = f(O,cl,r,p,t * . .)* Major (1951) called evolutionary differences in the flora of a certain landscape, caused by immigration or isolation, ‘florosequences’ and called changes introduced by various degrees of grazing pressures by domestic livestock (and therefore by man) ‘homosequences’ or homofunction Oh. In this paper, the term ‘homofunction’ has been replaced by ‘anthropogenic’ function. Its meaning has been broadened to describe biofunctions in which man (in the anthropological and not sexual sense), in the course of his cultural evolution (his ‘noogenesis’) has become a more and more independent and even dominating state factor in landscape formation. He thereby not only modifies biotic and edaphic variables, but also changes the other independent initial and driving state factors, derived from the biosphere and geosphere, and also introduces his artifacts as cultural variables derived from the noosphere. This coupling of natural and man-made variables in functions-factors landscape ecotope equations is justified by a holistic view, integrating man- 127 and-his-total environment in the ‘Total Human Ecosystem’, fit defined by Egler (1970). This concept was further deveIoped by defining landscapes as the concrete entities of the Total Human Ecosystem, or ecosphere, composed of bio-ecosystems (driven by biological conversion of energy) with the biosphere as the largest, global one and of techno-ecosystems (driven by technological conversion,of, chiefly, fossil energy) with the technosphere as the largest global one. Their smallest, mappable ecosphere landscape units or ecotopes are integrated spatially and visually but, alas, not yet functionally and structurally. To supply scientific and educational feedback for this integration is one of the major tasks of landscape ecology as a human ecosystem science (Naveh, 1978,1980,1982; Naveh and Lieberman, 1982 and Fig. I). The object of this paper is to describe the evolution and degradation of Mediterranean landscape with the help of such semi-formal multivariate ecotope functions and, on the basis of their study, to formulate a new dynamic approach to Mediterranean landscape conservation and reconstruction. Eco-sphere urban 1 &o-Sphere ‘1 i: Em Techno and -Sphere - sphere Fig. 1. Hierarchical black-box model of the ecosphere as the concrete system and global landscape of the total human ecosystem. The biosphere and technosphere and their landscape ecotopes of bio- and technoecosystems are integrated spatially and visually through the geosphere in the ecosphere. Their functional and structural integration is the goal of landscape ecology as a human ecosystem science (after Naveh, 1990). FOUR MAJOR PHASES OF MEDITERRANEAN DEGRADATION WITH THEIR MULTIVARIATE LANDSCAPE EVOLUTION AND BIOFUNCTION EQUATIONS Evolution of natural Mediterranean landscape over geological time As described in detail elsewhere (Naveh, 1973, 1977), we have every reason to suppose that natural fires, caused by lightning as well as by volcanic eruption, raged from the late Pliocene and early Pleistocene, especially since the last Wurm interglacial and interpluvial dessications when the present climatic fluctuations pattern, between wet and dry seasons, along with Mediterranean flora and fauna became established (Butzer, 1972). Therefore, fire and drough t 128 may have acted as dominant environmental agents in the evolution of these landscapes in a similar way as that recognized by Axelrod (1958) in the Madrotertiary geoflora of California. We can also suppose that grazing and browsing animals, especially ungulates, evolved together with the woody and herbaceous vegetation. These assumptions are supported by recent findings in local limestone caves, in which old bones, charcoal, ashes, fire-traced stones and reddened hearth areas show the use of fire by Paleolithic hunter-gatherers for some several hundred thousand years. By modifying above-described Jenny’s state factor equation, the evolution of the Mediterranean landscape can, therefore, be described as follows Es,” = f(P.R.Cld,fiO, . . . T< 1 000 000) (1) where T = time. In this equation, initial site conditions of parent material of the soil (P), the relief (R), the driving fluxes of climate (CI) and organisms (0), along with their most important evolutionary forces, drought (dr), fire (fi) and grazing (gr), operate in landscape genesis on dependent soil (s) and vegetation (u) variables of these landscape ecotopes (E) through geological time (T). Evolution of semi-natural Mediterranean landscape in Upper Pleistocene In the Upper Pleistocene, with the emergence of advanced and diversified mesolithic cultures, hunter-gatherers (Hhu@) became a more and more independent controlling state factor, modifying the dense forest by the intentional use of fire (bu) into more open multi-layered mixed arboraceous and herbaceous ecotopes, as claimed by Vita-Fincy and Higgs (1970) for the Palestinian Neanderthal Man of Mount Carmel and Higgs et al. (1967) for Greece. At the same time, the above-mentioned natural evolutionary forces were still operative in landscape genesis. The rich faunal collections in the final Acheulan, Levalloisian and Mousterian levels in the Carmel caves, together with ash, hearths (Garrod and Bate, 1937) and the steady increase of garigue, steppe and rock-dwelling rodents (Tchernov, 1968), are evidence of the co-existence of closed forests with open woodlands and grasslands, along with the gradual enlargement of drier, more exposed, rockier habitats induced by burning. Thus, these are the first indications of noospheric inputs of energy and cultural information by the land use of Stone Age hunter-gatherers. Therefore, eq. 2 can be considered the first anthropogenic biofunction, lasting several hundreds of thousands of years E s,v,a=f(Hbu, Evolution huga P,R,Cldr,ti,Ogr-. - TG100 of agro-pastoral cultural landscape 000) (2) in the Holocene In the Carmel caves the first agro-technological artifacts have been found: flint sickles and grinding stones apparently used by late Mesolithic, semisedentary Natufian cultures for harvesting and threshing parched kernels of 129 grasses, such as Hordeum s~~~~aneu~ and Triticum diccoeoides (the largegrained wild barley and Emmer wheat which are the progenitors of our cereals; see Zohary, 1969). These annual grasses are amongst the most prolific fire followers and post-fire collection of seeds could, logically, have been one of the first phases of their domestication. The first known centers of successful Neolithic cereal and stock breeding economies, marking the beginning of the evolution of agro-pastoral landscapes, were established in adjacent drier and very fire-prone subhumid woodlands and semi-arid grasslands, where the above-mentioned grasses are abundant. Thus fire, having operated for many thousands of years as a major force in biological evolution, became the environmental and cultural trigger of agro-pastoral evolution in the fire-induced maquis edges and the fire-swept open grasslands. As will be shown below, it also played an important role in the agro-pastoral biofunction, thereby shaping open Mediterranean landscapes for hundreds of thousands of years, until the present day. In this anthropogenic agro-pastoral (ag-pa) biofunction man, as the dominating state factor, began to control all other independent state factors. By burning, cutting, coppicing, land clearing, terracing, cultivating, grazing and browsing he converted natural and semi-natural bio-ecosystems into semi-agricultural (‘natural’ pastures) and agricultural bio-ecosystems (fields and plantations). By introducing cultural artifacts such as terraces, fences, wells, roads, houses, etc., he constructed rural, and later on urban-techno-ecosystems. In this way, agro-technological energy (including fire) and cultural information, coupled with positive feedback loops of accelerating cultural evolution due to the advent of written language and subsequent increased communication, created the cultural Mediterranean landscapes of the Jewish, Greek and Roman civilizations. These agro-pastoral landscapes and their management are well documented in the Bible, the Talmud and classical literature. They served as the cradle of our modern civilization and striking remnants can still be found in select locations in southern Europe and the other Mediterranean countries. Their evolution can, therefore, be characterized by the following eq. 3, with historical land-use cycles of at least several centuries E .$,i,?,a = f&g.pa(P,R,CltO . . . T<lOO) (3) This major cultural phase of agro-pastoral biofunctions lasted from the Neolithic revolution throughout the whole historical period, until a few decades ago. As described in more detail by Naveh and Dan (1973), it can be subdivided into several aggradation and degradation cycles, corresponding to historical land-use cycles of thousands and hundreds of years. The terracing of arable upland slopes, chiefly for olive pl~~tions and vineyards, after the uprooting of woody vegetation (trees and shrubs were apparently left along the terrace walls and on steeper slopes between the terraces), was started in the early Iron Age by the Phoenicians and reached great agro- and hydrotechnological sophistication. This was one of the few instances where man’s agricultural land uses improved the initial state factors controlling soil forma- 130 tion. By changing the original base levels into smaller secondary slopes with local base levels and by adding special gravel and soil layers near the terrace walls, man created deeper, more fertile and stable soils and increased their moisture-holding capacities. However, their abandonment and neglect in periods of political upheaval and insecurity was followed by the disintegration of the local base levels, especially if grazing was continued, and eventually led to increased catastrophic erosive degradation processes leading to the siltation of riverbeds and flood plains. This caused, in general, severe landscape dessication and, in the semi-arid zone, even desertification. On the other hand, on those steep and rocky slopes which were not suitable for cultivation even by terracing, the woody and herbaceous vegetation canopy, as long as it was not uprooted, continued to provide very efficient protection. Therefore, the shallow but fertile and fine-structured brown rendzina and terra rosa soils have suffered much less from erosion than has been generally assumed (Naveh and Dan, 1973). Throughout the long agro-pastoral phase, shrublands, woodlands and grasslands became functionally closely interwoven with terraces or patch-cultivated fields which were grazed before seeding and after harvesting, facilitating the transfer of fertility and seeds to and from adjacent untillable ecotopes. Ideal conditions were thereby created for introgression and spontaneous hybridization of wild and cultivated plants and biotypes, as shown in the case of Hordeum vulgure (the cultivated barley) which mixed with Hordeum spontaneum (the very abundant wild annual barley) in Israel (Zohary, 1969). This period was long enough to allow the evolution of genotypes better adapted to these man-modified conditions and it can be assumed that selection pressure favoured the survival of species and ecotypes, as well as plant communities, with the highest resilience to the combined impact of environmental rigour and defoliation pressures by maximizing their adaptive feedback responses. As described in more detail elsewhere (Naveh, 1975), negative-feedback responses enabled the avoidance of extreme conditions and disturbances which would have endangered plant survival, and positive-feedback responses increased physiological activities and regeneration vigour which overcame disturbance and stress. Simultaneously, the great variety in space and time, namely biological and microsite diversity, and the short-term, mostly cyclic, climatically-induced seasonal and annual fluctuations, contributed greatly to the global stability and persistence of these non-arable upland ecosystems, Their seasonal and annual fluctuations in productivity also acted as effective negative feedback in preventing overgrazing, because the numbers of livestock which could be supported during the critical period of low food availability in early winter were not sufficient to overgraze pastures during the spring flush of growth and seed-setting. At the same time, the over-use of the woody vegetation was also prevented by enforced burning and coppicing rotations to ensure sustained productivity and sufficient recovery. During the long phase of agricultural decay and population decline (until the downfall of the Ottoman Empire) these agro-pastoral biofunctions, i.e. 131 regular grazing, burning and coppicing regimes, led to the establishment of a dynamic equilibrium in the non-cultivated upland ecosystems, which were neither over-aged nor heavily coppiced. According to Naveh and Dan (1973), this man-maintained equilibrium between trees, shrubs, herbs, grasses and geophytes and between dependent and controlling state factors contributed much to the striking biological diversity and attractiveness of the hlediterranean landscape. This is, without doubt, its most important asset for recreation and tourism. Present accelerated ~e~~ec~no~o~~c#~degradation cycles The anthropogenic biofunctions are characterized by short~term and accelerating degradation cycles. These are caused by the combined, even synergistic, impacts of neotechnoIogic~ land uses (ne), the intensification of agro-pastoral land uses (ag-pa int.) in densely populated uplands along with mechanized land clearing and cultivation, or complete abandonment (ab). In addition, through urban-industry sprawl, pollution and mass recreation more and more bio-ecosystems are being biologic~ly and scenically impoverished or replaced by techno-ecosystems, with their artifacts (highways, quarries, etc.) reaching even the remotest sites. Unfortunately, not only these but also the two major land-use alternatives aimed at environmental conservation, namely complete prolonged nature protection (pr) and chiefly mono-species conifer ~fores~tion (af), are also disruptive of the dynamic equilibrium, given in eq. 3. These accelerating degradation cycles can be characterized by the following equation E su,a = ~~~~,~~-~~~t.,~b,~~~~~,~,c~,~ . . . mlo) (4) As will be shown below, these anthropogenic degradation functions are causing the depletion of organic variety and scenic beauty in the most productive and attractive ecotopes. In addition, the induced modem policy of fire prevention at all costs and at all times has disrupted the traditional cycles of periodic burning by p~tor~ists and has caused, in their stead, much hotter and more destructive wildfires, due to the greater amounts of dry fuel, along with the expansion of highly-flammable conifer forests and the increasing fire hazard of recreations and urban uses. In contrast to the above-mentioned environmental and cultural negativefeedback couplings which govern eq. 3, these degradation cycles, like all neotechnological processes, are driven by a positive feedback between fossil energymaterial production (or conversion) and consumption and cultural information. They are therefore growing at exponential rates, severely endangering the future of the open ~edi~~~ean landscape. VEGETATION COMPOSITION, STRUCTURE AND DIVERSITY AS DEPEXDENT BIOTIC VARIABLES OF RECENT BIOFUNCTIONS In order to compare these relations in some of the recent anthropogenic 132 biofunctions in Israel, either adjacent sites or those closely related in their geobotanical features, but differing in the recent dominating human management state factors, were chosen. We can, therefore, assume that in each comparison the study sites belong to the same class of ecotopes (E,). In this, with the exception of recent management (Hm, and Hmb), all other state factors of E, are held constant (Jenny, 1961), including the long time-span under which these sites have evolved under the same prior agro-pastoral biofunctions of eq. 3. Therefore, the dependent diversity parameters of sites Ex, (div a) and Exb (div b) can be considered as an anthropogenic biofunction of Hm, and Hmb Exvdiv a = f tHh)CIO,R,P,T . . . Exvdiv b = f (Hmb)Cl,O,R,P,T . . . It was further assumed that those study plots where previous traditional land uses of more-or-less moderate grazing, coppicing and burning pressures were continued without radical changes could be equalled with the agropastoral eq. 3 and therefore Hm, 2 Hag_pa However, on the other hand, all other plots of the same site class, where these traditional pressures were either very much increased (Hag-pa tit.) or reduced (Hpr), or where these sites were converted into dense pine forests (H&f), could then be equal to the present neotechnological degradation function of eq. 4, as described above. Table I represents such a comparison of the effect of these different land-use practices on the occurrence and abundance of orchid and other geophyte species; the most attractive flowering plants in the Mt. Carmel National Park. Site 1 is an open Pinus halepensis and Pinus pinea forest with a mixed, open understorey of sclerophyll shrubs, dwarf shrubs and grass located close to Bedouin homes where eq. 3, i.e. moderate grazing, browsing and collection of plants for food and brushwood for fuel, continues. Site 2, across the main road, is now almost completely protected from grazing and the process of brush encroachment is apparent. Site 3 has been converted into a dense, shaded pine forest by additional plantings of Pinus halepensis trees. Here, not only these geophytes but also most other herbaceous plants have been eliminated and the woody understorey has been reduced in species numbers and coverage. These three sites are typical for the present H,, (or Hab) and Hd biofunctions, not only in the eastern Mediterranean but also in the western Mediterranean countries, especially southern France (Schreiber and Naveh, 1980, unpublished data). Light measurement with a solarimeter on cloudless spring mornings confirmed the close relationship of these mostly heliophytic orchids to light intensities above 0.11 g cal cm-’ min“, as measured in open niches of browsed and burned brush canopy, on the edges of goat paths and in grassy spots. 133 TABLE I Species richness and relative abundance of geophytes in adjacent disturbed (l), protected (2) and afforested (3) sites on Mt. Carmel (spring 1969) Sites and relative abundance* (1) (2) 3 1 3 3 1 1 + (3) Orchid species Ophyrs sintenissii Fleisch et Bornm. Ophrys dinsmorei Schltr. Ophrys fuciflora Hal. Ophrys bornmuelleri M. Scbeuze Ophrys lutea Cav. Ophrys fusca LK. Ophrys iricoior Desf. Serapias uomeracea (Burm) Brig. Anacamptis pyramidalis (L) Rich. Orchis pa~ilionaeens L. Orchis anatolicus Boiss Orchis galilaeus Schltr. Orchis tridentatus Stop. Cephalanthera longifolia (Huds) Fritsch Limodorum abortiuum (L.) SW. Total no. species at site Other geophyte species Asphodelus microcarpws Viv. Asphodelus tenuifolius Cav. All~um ampeloprasum L. Allium neapolitanurn Cyr. Allium hirsutum Zucc. Ornithogalum narbonense L. Iris sisynchium L. Iris palaestina (Bad) Boiss. Arisarum vulgare Targ. Arum dioscoridis S. et S. Anemone coronaria L. Ranunculus asiaticus L. Cyclamen persicum Mill. Thrincia tuberosa (L) Lam. et DC Bellis silvestris Cyr. - - ; - - 2 3 2 3 12 - 2 2 + - 1 + + + + + 2 2 1 1 10 2 1 2 - - + + 2 1 1 1 - - 1 1 1 - + 1 2 1 2 2 1 1 1 1 1 1 1 Total no. species at site 13 11 5 Total no. geophytes 25 21 7 *3 = very abundant; 2 = abundant; 1 = occasional; + = rare. 1 2 2 - 134 TABLE II Effect of different anthropogenic biofunctions on stratified and overall plant diversity shrubland and woodland sites of 1000 mz in northern Israel 1975-1976 Growth-forms and diversity measuresa Perennial herbs (species no.) Woody species (species no.) H’F in Sl S’I’R” Total Ge PGr Pfo Total TS MS SS 7 5 4 5 21 13 3 2 3 8 1 6 14 16 5 42 24 13 6 20 39 - 5 5 I.0 8 28 19 14 5 3 22 2 2 6 5 4 19 15 8 3 12 23 Mt. Carmel sites Closed undisturbed (H,ro) Muhraqa Semi-open disturbed u&z*,) Forty Oaks Grove Lower Galilee sites Heavily grazed, browsed and cut wag%mirlt 1 Bosmat T&on Moderately, rotationally grazed (%z*a) Allonim aGrowth form codes and diversity measures: (HI’) = high tree; (TS, MS and SS) = medium and low shrub; (CL) = climber; (Ge) = geophytes; (PGr) = perennial grass; perennial forb; (AGr) = annual grass; (ALe) = annual legume; (AFo) * annual forb ing Iegumes). C = Simpson index of dominance concentration, using refative cover plant species (Cveg), relative cover of species within the woody strata only (Tweed) relative cover of five strata as wholes; H’ is the Shannon-Wiener index for relative ail pfant species, tog,,. bIncludes species present in more than one stratum (Naveh and Whittaker, 1979). tall, (PFo) = (excludfor ah and cover of Additional striking proof of the adverse effects on structural and floristic diversity of disruption of traditional agro-pastoral functions can also be found in the Samarian mountains. Here, in typical shallow and rocky terra rosa soil on hard limestone and dolomite, complete and prolonged protection of the Urn Rechan Forest Reserve (throughout the Mandatory, Jordanian and present Israeli rules) for more than 50 years has created a green island of several square kilometers. This is surrounded by heavily grazed, coppiced (and therefore severely impoverished and stunted) maquis shrubland, typical of Hwpa ht. preva~ing under heavy human and iivestock pressures, over large areas. However, closer inspection reveals that the protected maquis has turned into a dense and almost inaccessible brush thicket, especially on the more mesic northern slope. This is populated in the upper layer by 3-5 m-tall Quercus c~ff~~ri~os (dominant), P~flfyrea media (subdomin~t), eight other sclerophylls 135 Annual herbs (species no.) AGr ALe Total herb Cves C wood CStI K 21 0.408 0.600 0.574 0.581 Total species 1000 m3 AFo Total I_ - - 24 25 56 95 119 0.177 0.270 0.190 1.352 7 14 22 43 65 84 0.336 0.569 0.282 0.904 16 28 53 97 120 135 0.029 0.449 0.253 1.718 species - 7 8 (rare) and, in the lower shrub layers, by Pistaciu len tiscus (dominant), Cistus sa~u~~o~~~s(subdomin~t), three sub-shrubs, four climber species and a very few shade-tolerant herbaceous plants near rock edges. In spite of the favourable slope exposure and an annual rainfall of 600-700 mm, there are relatively few young shoots and most taller shrubs show signs of senescence. The large amount of highly-combustible dead material, dry branches and a thick undecomposed litter layer creates a great fire hazard. Similar results are shown in Table II. These have been derived from recent, more detailed comparisons of such contrasting biofunctions from two main site classes in northern Israel (Naveh and Whittaker, 1979). The first is from typical Quercus calliprinos maquis shrubland on Mt. Carmel in a subhumid climate of, 700-800 mm annual rainfall between October and May in shallow and rocky terra rosa on hard limestone of Cenomanian origin. The second is 136 from open (deciduous) Quercus ithaburense (Tabor oak) woodland in the foothills of the lower Galilee in a slightly drier climate, with 600-700 mm annual rainfall on shallow but fertile dark rendzina soils of Eocenic origin on soft limestone with a Nari (caliche) crust. As might be expected, in the drier and more open oak woodlands, serving as natural pastures, the number of herbaceous species is much higher. However, of greater significance are the differences within each site class induced by the different land-use practices. All the Mt. Carmel sites now belong to the Mt. Carmel National Park. The Muhraqa site, completely protected for 40 years, has turned into an almost inpenetrable one-layered tall shrub canopy dominated by Quercus calliprinos with very low numbers of total and stratified woody species and only a few shade-tolerant perennial herbs. As a result, very low values for equitability (H’) and, conversely, very high values of woody and stratified dominant concentrations (C) were recorded. In the ‘disturbed’ site of the Forty Oaks Grove, pruning and protection of ‘holy’ oak trees in the distant past has led to an overstorey of very large trees. The site is kept, by moderate to light grazing and browsing and recent burning, in a semi-open state with a well-proportioned woody cover between the different strata and species. This leads to low woody C values and favours a rich herbaceous understorey containing a wealth of flowering geophytes, as well as hemicryptophytes and therophytes in openings. This is reflected in the high H’ values and there is a total of 119 species, with 95 herb species, in the Oaks Grove as compared to only 21 species and eight herbs in the protected Muhraqa ecotope, A simultaneous zoological study (Warburg, 1977) showed that the semi-open ‘disturbed’ sites also had much greater species richness and abundance of birds, rodents, reptiles and insects. Similar striking differences can be noted in the second group of woodland ecotopes. The Bosmat-Tivon site, in the vicinity of a Bedouin settlement, is grazed and browsed very heavily throughout the year by goats, sheep and cattle and the oak trees have been cut down indiscriminately for fuel. This has resulted in an impoverished mosaic-like vegetation pattern of dense shrub patches, dominated by the unpalatable, very resilient sclerophyll Pistacia lentiscus, alternating with grassy patches of mostly unpalatable small grasses and forbs, described in detail by Whittaker and Naveh (1980). Such Pistacia lentiscusdominated shrub-grass mosaics are now typical for many maquis in Israel and other eastern Mediterranean countries, especially near villages which are exposed to the above-described recently intensified agro-pastoral biofunctions of eq. 4. On the other hand, the moderately grazed Tabor oak woodlands near the collective settlement of Allonim are still maintaining a dense, rich, chiefly annual herbaceous cover and consequently have very high alpha diversity. However, under light grazing, and even more under protection, these pastures rapidly lose their high alpha diversity and turn into species-poor, tall grass stands, dominated by Auena sterilis (Naveh and Whittaker, 1979). Under prolonged protection hemicryptophytic, perennial grasses and thistles increase but flowering geophytes, such as Cyclamen per- 137 sicum and Anemone coronaria, are smothered together with most flowering annuals, so that these protected Tabor oak woodlands lose most of their landscape amenity values (Naveh, 1971). In this way this biocline is characterized by a distinct two-slope relation of diversity to grazing pressure with minima at lowest and highest grazing pressures, typical for eq. 4, and maxima under moderate, rotational grazing, resembling the traditional agro-pastoral biofunction. We can, therefore, summarize these results with the general conclusion that E divd < E div,, < E divag-Pa > E div,,, COMPARISONS ht. WITH OTHER MEDITERRANEAN BIOMES Although derived chiefly from studies in the Mediterranean, mountainous parts of Israel, these multivariate biofunctions are applicable to all uplands around the Mediterranean basin with actual or potential evergreen sclerophyll vegetation and also, after modification, to other regions with similar Mediterranean-type climates, vegetation and landscapes. Zohary (1974) stated that the so-called maquis ‘climax’ communities in which human interference ceased were turning into very monotonous shrub thickets dominated by Quercus calliprinos in all Near Eastern countries. Horvat et al. (1974) also found that the cessation of intensive agro-pastoral utilization of the southern European Adriatic Quercus ilex forests during the last 40 years has caused the loss of their open appearance and, consequently, their floristic and fauna1 richness. In our recent studies in southern France (Schreiber and Naveh, 1980, unpublished data) we found low diversity values in the abandoned, fire-prone, nongrazed Quercus ilex maquis, Quercus coccifera garigues and Pinus halepensis forests, but slightly higher values in their recently-burned counterparts. Susmel et al. (1976) showed that the oldest Quercus ilex ‘climax’ forests in Supramonte, Sardinia were maintaining their dynamic, energetic and metabolic balance under traditional pastoral uses, in spite of the fact that 50% of the acorns were consumed by swine. On the other hand, they claimed that the ‘modern’ land-use practices of tree-cutting for timber and the cessation of grazing for ‘protection’ were leading to increased fire hazards and endangering these magnificent forests. As shown recently by Naveh and Whittaker (1979), the three truly convergent Mediterranean biomes - based on their relatively recent Pleistocene evolutionary history - are the Mediterranean proper, Chile and California, as opposed to the divergent, much older Gondowan biomes in South Africa and Australia. In these Mediterranean biomes a sequence in diversity values can apparently be found, according to the duration of the agro-pastoral biofunctions, with by far the highest values in the Mediterranean itself, followed by Chile, with lowest values in California (where the latter biofunctions were commenced only 200 years ago). In addition to influencing diversity, the long, gradually-evolving agro-pastoral biofunction, as opposed to its sudden 138 and recent rise in other Mediterranean biomes, has had far-reaching consequences on the biomes’ composition, productivity and.stability. Drought, fire and grazing-tolerant grasses and legumes, evolving under the early Mediterranean agro-pastoral functions, served as genetic stock for the most important grains and pulses and, more recently, for pasture and fodder plants also. Their gen-ecological advantages as chiefly autogamous colonizers and their opportunistic, flexible behaviour have preconditioned many of these Mediterranean annuals to become widespread weeds and to invade other areas with similar Mediterranean-type climates successfully, where these agropastoral functions were commenced only in recent decades. Thus, a comparison of Medi~rr~e~ landscapes in California and Israel (Naveh, 1967) revealed that more than 100 annual species, auto~hthonous in Israel and the Mediterranean, have naturalized in California also. Most of these are abundant in annual grasslands or as weeds in fields and on roadsides. According to Robbins (1940), more than 400 alien species can be found in Californian grasslands. An even more unreliable rainfall and temperature regime in the critical early winter period, coupled with the shorter agro-pastoral phase and consequently lower adaptive resilience, plus lower diversity, has made Californian uplands much more vulnerable. This is especially true for overgrazing, misuse of the poorer, coarse-structured non-calcic brown upland soils and for the destructive effects of complete fine protection, which has disrupted the fire cycles of the Indian hunter-gatherer biofun~tion given in eq. 2. Here, in contrast to the Mediterranean, where such burning cycles were continued also in eq. 3, fire has been used by the European settlers (with the exception of controlled burning of foothill ranges) only as a tool for land clearing prior to cultivation. However, in contrast to the Mediterranean, recent important work on fire ecology (Biswell, 1967) has already led to a reappraisal of ecologically-unsound fire protection and has initiated controlled burning and fuel management in national parks and forests in an attempt to simulate the Indian practices (Van Wagtendonk, 1975). In a recent comparative study of the closest landscape-ecological equivalents in California and Israel, namely the Quercus douglasii woodlands in the central foothills in California and the Quercus it~ubu~ense woodlands in the lower Galilee foothills in Israel (Griffin and Naveh, unpublished data), we have found a similar two-slope relation of diversity and grazing pressure, but under a considerably lower plateau of diversity. On the rested and lightly grazed pastures, annual grasses, chiefly the aggressive Mediterranean invaders Auena fatua, Avena barbata and Bromis mollis, dominate. DISCUSSION The application of multivariate functions, with man as one of the independent ecosystem state factors, is an attempt to present landscape evolution as a combined process of biogenesis and noogenesis. In this, the formation of 139 the noosphere, as perceived by Teilhard de Chardin (1966), is reflected in the equation of these functions by the more dominating role of homo sapiens as driving state factor, through which the original pedogenic and biogenic functions were gradually replaced by psychogenic ones. In this way bio-ecosystems and their concrete landscape ecotopes were converted into semi-natural ones. Man is regarded as a ‘holon’* of the biosphere which is affected by and affects it via the technosphere (his concrete noospheric systems outside the biosphere). In order to render these state factor equations useful for better ecological comprehension, and especially for landscape conservation m~agement, they should be ‘solved’ by quantitatively relating their ecosystem properties to the dominating human state factor. However, as mentioned in the introduction, these equations are only semi-formal and those relations derived from human living systems cannot be quantified in the formal mathematical sense. The Total Human Ecosystems, of which these landscape ecotopes are concrete systems, as explained in the introduction, should be recognized as “self-transcendent Gestalt systems” (Pankow, 1976). Their openness goes beyond the formal openness to flow of energy, material and information, thus they are capable of representing themselves and being recognized in their entirety only by other natural Gestalt systems, such as everyday lay language, rather than the formal scientific language of mathematics or other conceptual systems. At the same time, these equations are formal in the sense that their elements are expressed as variables and their relations as functions. These can be treated, as suggested by Zadeh (19731, as ‘fuzzy sets’ in which the different anthropogenie land-use practices are l~~istic~ly characterized and, wherever possible, relative values can be designated for their gradients. This has been attempted in this study using the example of bioclines of increasing grazing pressures. These functions are biofunctions in the widest sense and their evolutionary trends should be judged as such. Thus, in the agro-pastoral biofunctions, the evolution of ‘Homo tuber’ is reflected by the conversion of these semi-natural ecotopes into semi-agricultural ‘natural’ pastures and plantations, or they are replaced entirely by the chiefly rural techno-ecosystems of farms, villages, roads and their artifacts. In this way, the classical cultural ~edi~~~e~ landscape evolved. However, in recent years ‘Homo ~~d~s~iul~s’ has accelerated and expanded this process of replacement, intensified the agro-pastoral land uses and added new ones. The combined impact of these factors has distorted the dynamic flow equilibrium of degradation and regeneration cycles, typical for the agro-pastoral functions, and has turned these cycles into exponential degradation functions. Some of the alarming effects of these recent trends on biotic variables have been illustrated by the results of diversity studies in *‘Hoion’ (a combination of holos = whole, and proton = part), coined by Koesffer (1969) to emphasize the dichotomic nature of biosystems as intermediary structures in the ascending order of complexity, or ‘holarchy’, acting both as autonomous and self-asserting wholes towards their subordinate subsystems and as dependent integrative parts of their supersystems. 140 Israel. These are also typical for all other Mediterranean countries in which similar anthropogenic degradation functions are operating on the open landscapes. Equations 2 and 3 emphasize not only the increasingly dominant role of man, coupled with the shortening time-span of each phase, but also the lasting role of fire and grazing - first together with drought (as natural state factor) and then as anthropogenic land-use factors - in shaping Mediterranean landscapes until the present. This is in contrast to the recently introduced neotechnological land uses, including protection, afforestation and recreation. In Fig. 2, these major phases in the evolution of the cultural Mediterranean landscapes are presented graphically in an isomorphic model depicting the structural and spatial dimensions of these changes through time. From these, it seems now even more obvious that the increasing noospheric impacts are leading to increasing dominance of human artifacts and, thereby, to the replacement of biosphere by technosphere and to the disappearance of the open, unspoiled Mediterranean landscape. Unfortunately, few reliable figures are available on the speed and extent of this process. Therefore, we can only indicate the general trend: that techno-ecotopes have not only more than doubled their area but have influenced much of the modern industrial-chemical (and recently also plastic) agricultural ecotopes, in addition to the semiagricultural and few remaining semi-natural ones. At the same time, large-scale monospecies afforestation projects with conifers (which, in addition to being highly inflammable, are also very susceptible to air pollution, especially photochemical smog (Naveh et al., 1981)) have depleted organic variety and caused irreversible changes in many upland sites without ensuring socioeconomic returns which would justify these projects. In addition, some of the most attractive and valuable biotopes are also gravely endangered by commercial over-development, serving as a positive feedback for mass recreation and depletion. Unfortunately, Clementsian relay floristic climax-succession dogmas based on a deterministic axiom are still uncritically accepted by most Mediterranean ecologists and phytosociologists from the Braun Blanquet school. In this “facilitation model” (Connell and Slatyer, 1977), lower successional stages are supposedly always being replaced by more developed and more demanding ones until the final stable and mature ‘climax’ is reached. These highly hypothetical views have induced the adoption of complete and prolonged protection from fire, grazing and other human interventions to facilitate the reconstruction of the ‘maquis climax’ as the major aim of Mediterranean conservation strategies (Tomaselli, 1977). However, present Mediterranean vegetation dynamics are highly stochastic multivariate functions. Under certain conditions they behave according to the Connell and Slatyer (1977) “inhibition control model”. In this, progressive successions are arrested at the lower herbaceous and dwarf shrub stages but, after cessation of perturbations by fire, grazing and cutting there is, in general, a process of vegetative regeneration of all extant sclerophyll shrubs and trees. 141 E”C%UTlON MAl’ERlAl. LOWER FwiioQ PLElSTOCENE 4E8) GEGRAl,ATION AU9 HFCfWATiOt4 ff MDITERRANEAN WU15 FROM UPPER PLEISTOCENE THE LANDSCAPE GEOSWERE, ECOTWES EIOSPHERE HOLOCENE AGRD-PASTORAL BY ENERGY. m NOOS~MRE PRESENT NEO-TEU(NOCGGICAL INCREASING MOOtFiCATiON, OF BIO-ECOSYSTEMS AN0 OCMINANCE OF ~~~A~-~AD~ ARTIFACTS Fig. 2. EvoIution and degradation of Mediterranean Iandscape ecotopes by energy, material and information inputs from the geosphere, biosphere and noosphere. ‘I’ H E = Total Human Ecosystem. Geosphere and biosphere inputs from natural and total human ecosystems: (v) biological conversion of solar energy; (of natural material and organisms; (v) bio-physico-chemical information and control. Noosphere inputs from total human ecosystems: (LA) technological conversion of fire energy; (* technological conversion of muscle energy; (4 technological conversion of fossil energy; (v) cultural and technological information and control; (0) agricultural material and organisms; (a) rural artifacts (chiefly natural); ( l ) urban-industrial artifacts (chiefly converted and synthesised). Landscape ecotopes L%l closed forests, woodlands and shrublands of natural bio-ecosystems; m semi-open forests, woodlands and shrublands of semi-natural bio-ecosystems; m semi-open and open forests, woodlands, shrublands and grasslands of semi-a~icultur~ ecosystems; 0 terraced and cultivated fields and plantations of agro-bioecosystems; m farms, villages, roads etc. of rural techno-ecosystems; q cities, factories, roads, etc. of urban-industrial techno-ecosystems. Biofunctions: P,R,Cla,fiOo,. f Hne.agga (I) E, v = f(P,R,Cld,RO . . . . ‘2’ G 100 000); (IIfiE,l;a’- 7-4 1000 000); (II) .!$ u = f(Hbu h,, f HaBqa (P,R,Cl . , . : . 2% 1OOj; (E) Es,u,a = int.,ab.af,pr(P,ReCl . . . . . m 10). This process can be c&led “autosuccession”, after Wanes (1971). 1x1many abandoned vineyards and orchard plantations there is also direct invasion of sclerophylls from adjacent stands without any intermediate successional stages (Naveh, 1982). In addition, as explained above, in all truly Mediterranean biomes eqs. 2 and 3 have converted the natural Pleistocene landscapes into semi-natural and cultural ones. The possibility of reconstruction 142 of a hypothetical pristine Pleistocene climax is highly improbable and is neither of theoretical nor practical relevance for conservation m~agement and research. The prolonged complete exclusion of fire, man, his axe and livestock from these semi-natural landscapes cannot be regarded as the re-creation of a ‘natural’ situation, which would presumably lead to the re-establishment of a hypothetical climax of mature and stable communities. On the contrary it may iead to a less natural and, from the point of view of biological diversity and scenic attractiveness, less desirable situation. We can safely assume that the loss of structural floristic and faunistic diversity in such protected, closed, monotonous and senescent maquis and chaparral thickets will be reflected in a lower efficiency of energy interception and transfer and in a reduction of channels for nutrient and water circulation and storage capacity. In thermodynamic terms, this means an increase in entropy at the cost of productivity and over-all long-term global stability. This is manifested by the great vulnerability of dense maquis stands, as well as fire-protected chaparral in California, due to the accumulation of large masses of highly combustible dead material (Schultz, 1967). Pignatti (1978) was the first to introduce the thermodynamic concept of dissipative structures as a combination of biological and cultural processes, in a si~ific~t paper on the evolution of cultural ~edite~~ean landscapes. The role of dissipative structures as a new ordering principle in non-equilibrium systems has been defined by Prigogine (1976) and Nicolis and Prigogine (1977). This state of non-equilibrium is brought about by the effect of increasingly powerful env~onm~nt~ constraints, imposing a continuing change in entropy on the system through fluctuations and removing it further from equilibrium. These are ordered structures which, as opposed to those in equilibrium, are maintained and stabilized only by permanent energy exchange with the environment, namely “structures which dissipate energy”. Because of their tendency to move through a sequence of mutating transitions to new regimes which, in each case, generate the conditions of renewal of higher entropy production within a new higher regime of org~ization and order, they create “order through fluctuation”. According to Jan&h (‘f975), this new theory of self-organization can be regarded as a major breakthrough in our transdisciplinary conceptions of physical, biological, ecological and human systems. It can also be applied to the the~odynamic interpretation of the st~ctur~-functions inte~ation of bio- and techno-ecosystems in concrete landscape ecotopes, leading through mutation towards a dynamic regime at a higher state of compIexity. This is also supported, in general, by the biocybemetic behaviour of viable biological and ecological systems, as defined by Vester (1976). Attempts should be made to apply this theory to a dynamic interpretation of the behaviour of the ~edite~ane~ upland bio-ecosystems. These are manmodified, semi-natural and ~rni-a~cult~~, mutating from one stage to another as complex metastable systems. Both natural and man-induced flue: 143 tuations provide the major means for energy exchange with the environment, restoring the system’s capability for renewal of entropy production through rest periods (without burning, cutting, grazing, etc.) until a new peak of entropy is reached in each dynamic regime. The above-described multivariate biofunctions also fit the topological models of epigenetic landscapes used by Waddington (1975). In these, stable growth paths are canalized pathways across generalized surfaces of system development. Thus, bio- and eco-systems move through time, in multidimensional space, as locally unstable but globally long-term stable systems. Waddin~on, who was not only an eminent geneticist but also a great science and biology philosopher, called these canalized pathways of systems’ flow processes “chreods”. He urged turning away from linear and deterministic mathematical, biological and ecological models which eannot do justice to the adaptive evolutionary dynamics of biosystems, create homeostasis and, in ecological terms, a stationary climax state. He coined instead an important concept of evolutionary stability, namely “homeorhesis”, the preservation of these systems’ flow processes as a pathway of change in time or, in other words, to keep the systems altering in the same way as they have altered in the past, Accordingly, our Mediterranean bio-ecosystems and their concrete landscape ecotopes meandered in very broad chreods, comparable to “river flood plains”, as long as they were governed by the traditional agro-pastoral biofunctions. However, in present neotechnological degradation functions they are pushed out of these chreods over the ‘watershed’. Therefore the reestablishment and maintenance of their homeorhetic flow equilibrium should become the major object of dynamic conservation management. CONCLUSIONS From the results of this study and its theoretical and practical implications it is obvious that any attempt to conserve Mediterranean landscapes, especially their striking organic variety in space and time, should be directed towards continuation of and/or simulation of the traditional pyric and biotic defoliation pressures of the agro-pastoral biofunctions, which have created and maintained this variety throughout hundreds, if not thousands, of years. This can be achieved only as part of comprehensive conservation and development landscape masterplans for all non-tillable Mediterranean uplands, as outlined in more detail in a forthcoming book (Naveh and Lieberman, 1982). In these masterplans rational utilization strategies should replace not only the complete protection policy but also the even more destructive, narrow and short-sighted livestock and silvicultural crop-production-oriented land uses, regarding these uplands merely as a source of immediate revenues for private or national interests. Their major value, however, lies in their ‘noneconomic’ landscape values, i.e. in their combined bio-socio-ecologic~ ecosystem functions for which no alternative land is available. Their loss would, therefore, be final and irrevocable. 144 In nature reserves the highest priority should be given to sites with unique biological, geological or cultural value, aiming at maximum attainable ecological diversity. In nature parks and recreation areas, optimization of landscape, wildlife and recreation amenities should enable maximum enjoyment with minimum damage to natural resources. In the remaining open lands, used primarily for economic benefits, landscape management should aim at optimization of all bio- and socio-ecological and economic benefits according to site potential and socio-economic and other requirements, weighing all relevant land-use and env~onment~ variables and their mutual influences. In this way, protective and regulatory functions can be combined with plant and animal production functions in dynamic conservation management of closely interwoven networks of multiple land-use patterns (Naveh, 1919,1982), What is needed, above all, is a radical change in the attitude of decisionmakers, land planners, owners and users, from the present narrowly revenueoriented exploitation and negligence to a more far-sighted landscape-ecological determinism in land-use decision-making. 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