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
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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.
One of the first steps could be the
initiation of “Redbooks of Threatened
Mediterranean
Landscapes” by the
International
Union for Conservation
of Nature and Natural Resources, in
a similar line to the Redbooks of threatened plants and animals, in order to
provide more concrete figures and facts and as a vehicle for public awareness
and action. Within the Redbook, in addition to inventories of threatened
ecotopes, a special category should be devoted to those landscape in which
traditions
Medite~ne~
upland uses are still intact and in which these
semi-natural ecotopes deserve special protection
as part of natural biotic or
anthropological
reserves, within controlled interferences
and more intensively
managed agro-ecosystems.
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