Journal of Arid Environments, (1978) 1,63-69.
Rodent species diversity in mesic and xeric habitats in the Mediterranean region of northern Israel M. R. Warburg,* A. Ben-Horin* & Dina Rankevich* Rodent species diversity was studied in eight experimental plots located along a climatic gradient within the Mediterranean region in northern Israel. One plot was located at Mt Meron (precipitation 1000 mm), three plots were selected on Mt Carmel (precipitation 750 mm), three in the Lower Galil near Allonim (precipitation 650 mm), and one on the Gilboa Mts (precipitation 500 mm). The largest numbers of both species and specimens were found in the xeric habitat on the Gilboa Mts, The two most abundant species there were Acomys cahirinus and Gerbilus dasyurus, both confined mostly to rocky habitats. Apodemus mystacinus and A. sylvaticus appear to be the most typical rodents of the oak-woodland. They were abundant whenever tree cover was densest (on Mt Meron). However, otherwise no good correlation could be established between the various vegetation factors (PVC, PSD) and rodent species diversity.
Introduction In recent years, several studies on species diversity have appeared, most of which analysed bird species diversity (BSD). BSD was found to be correlated with foliage height diversity (FHD) and not with plant species diversity (MacArthur, 1965; Whittaker, 1972; Goodman, 1975). In general there is a linear relationship between BSD and vegetation components, especially in woodland habitats (Willson, 1974), but less so in grassland communities (Wiens, 1973) or in desert scrub (Tomoff, 1974). In the latter it appears that plant coverage is of greater significance than plant species diversity (Tomoff, 1974). Even in woodland habitats, the percentage of vegetation cover (PVC) is apparently of greater importance than previously thought. There, BSD was found to have a curvilinear relationship with PVC (Willson, 1974). Whereas the correlations between BSD and components of the vegetation are comparatively simple, studies on reptilian species diversity have so far not established any simple relationShip, and with mammals the situation is even more difficult. Thus, lizard species diversity studied mostly by Pianka (1973) was found to be related to several factors including length of the growth season, amount of productivity, heterogeneity of habitat, etc. Several of these studies took place in true desert or in semi-arid grassland habitats. A number of studies are known from the tropical forest (Schoener, 1969) but they, too, did not establish good criteria relating reptilian species diversity with a measurable vegetation factor. Only recently, in a stUdy on lizard species diversity in the Mediterranean region of Chile, Hurtubia & DiCastri • Department of Biology, Technion-Israel Institute of Technology, Haifa, Israel.
64
M. R. WARBURG, A. BEN-HORIN & D. RANKEVICH
(1973) have shown a definite relationship between both trophic diversity and habitat diversity
and lizard species diversity. Only a few studies have dealt with rodent species diversity (Rosenzweig & Winakur, 1969). In some of the desert habitats studied, rodent species diversity was found to be correlated with precipitation (Brown, 1973), and indirectly with herbaceous perennials (Beatley, 1976), which are directly correlated with the amount of precipitation. For some rodent species shrub volume diversity appeared to be the best measure showing a correlation with rodent species diversity (McCloskey, 1976).
Materials and methods Experimental plots were chosen in eight different localities in northern Israel. These localities all within the Mediterranean region included both grassland, shrubland and oakwoodland. The localities were situated along a climatic gradient. One plot was located in the Upper Galil on Mt Meron at an elevation of 900 m, where the precipitation was about 1000 mm. This area was densely covered by oak woodland (Quercus calliprinos and Pistacia palaestinai. Three more plots were chosen on Mt Carmel, all in one area where the precipitation was 750 mm but differed in vegetation cover. Thus, one plot was densely covered by Maqui of the oak Q. calliprinos (Muhraqa), in another the oaks were less dense and included open patches of the shrub P. lentiscus (Bet Oren), and the third was covered by rather tall, old oak trees Q. calliprinos ('40 Oaks' Grove). Three additional plots were chosen in the Lower Galil where precipitation reached about 650 mm. These included an area with large oak Q. ithaburensis and patches of open grassland (Allone Abba). A second plot was chosen in an area sparsely covered by Q. ithaburensis trees and grassland in between (Allonim). These two plots were moderately grazed by cattle. The third plot within this area was covered by shrubs of P. lentiscus and was grazed by goats (Basmat Tivon). Finally, the last plot was in a semi-arid climate though still within the Mediterranean region, bordering with the arid Irano-Turanian region. This plot, located on Mt Gilboa, with a precipitation of 500 mm was mostly grassland sparsely covered by the shrub P. lentiscus. An area of 10 ha was surveyed. This was done especially as most rodents are known to be considerably mobile, ranging sometimes for a long distance. Rodents and other small mammals were trapped using a vegetable spread as bait. Traps were set 5 m apart in two transects 20 m long. Trapping took place in each plot approximately during the same period. The trapped animals were then identified, marked, released and sometimes recaptured. The study lasted over two years during 1974, 1975 and 1976, and included also a botanical study which will be reported elsewhere.
Results Altogether ten species of rodents were found in the Mediterranean region studied (Table 1). Of them, seven species were found on the Gilboa Mts which is bordering with a semi-arid region (Fig. 1). In this area the largest number of specimens was found as well (91). Mt Meron was second in abundance in both rodent species (4) and in the number of specimens (52) (see Fig. 2). A similar pattern was noticeable when the percentage of specimens that had been trapped was calculated. Here again the figures for Mt Meron and Mt Gilboa were the highest among all the study sites (12,41 and 10·96 per cent respectively). All other plots yielded less rodent species and only a few specimens in each. Thus, in the three Mt Carmel sites a similar number of species was found (4--6) with only a few specimens of each species (Fig. 3). To our great surprise, the oak-woodland area in the Lower Galil was rather poor in rodents. The four most abundant species of rodents were the Spiny mouse (Acomys cahirinus), two species of Wood mice (Apodemus mystacinus and A. sylvaticus), and the Rock gerbile (Gerbilus
Acomys cahirinus Apodemus mystaeinus Apodemus sylvaticus Rattus rattus Rattus noroegicus Mus musculus Meriones tristrami Gerbilus dasyurus Microtus guentheri Cricetulus migratorius
-
-
-
-
3
-
1 Total No. of species Total No. of specimens 3 491 Total No. of traps 0-61 % of specimens trapped (x 10-1) 0 Diversity (H') (~P,log. P, where P, = proportion of each species in the sample)
1. 2. 3. 4. 5. 6. 7. 8. 9. 10.
(F)
Allonim
-
-
1 1 625 0·16 0
-
-
-
-
377
-
-
-
-
-
-
1
-
-
-
(C)
Basmat Tivon (D)
Allone Abba
.-
4 9 732 1·23 1·27
-
5 10 664 1·51 1·23
-
-
-
-
1
-
1 6 1 1
(E)
Bet Oren
2
-
-
2 4 1
'40 Oaks' Grove (B)
6 9 832 1·08 1·73
-
-
1 2 2 2 1 1
(H)
Muhraqa
Table 1. Rodent species diversity at different localities in the Mediterranean region
4 52 419 12·41 0·78
-
1
-
-
-
27 23 1
Mt Meron (G)
7 91 830 10·96 1·23
3
-
6 3 27
-
49 1 2
(K)
Gilboa
10 175 4970 3·52 1·75
56 40 29 5 1 10 3 27 1 3
Total
M. R. WARBURG, A. BEN-HORIN & D. RANKEVICH
66
dasyurus) (Fig. 4). The two species of Wood mice were particularly abundant on Mt Meron,
but were also found in plots on Mt Carmel and on the Gilboa Mts. The other two species were most abundant on the Gilboa Mts with one of them occasionally found elsewhere as well. A rare species, Cricetulus migratorius, was found on the Gilboa Mts.
8
'" Q)
u
Rodents
7 I-
6
~ 5
_'" 4 o
.. 3
Q)
..0
E
i
2 I
o
~~
K H E G B C F Locolites
Figure 1. Number of rodent species in the various localities (letters indicate localities as in Table 1).
..
~
55 50
Gilboa
Mt. Meron
45
E 40
...
t 35
:. 30 o 25
..
~ 20 rE ~ 15 10
5
o
8
mf!1l~~_ 6
7 10 3
. .
2
--
234 9
Species
Figure 2. Number of specimens of each rodent species found on Mt Meron and Mt Gilboa. Numerals indicate the rodent species. 1. Acomys cahirinus. 2. Apodemus mystacinus. 3. Apodemus syloaticus, 4. Rattus rattus, S. Rattus noruegicus, 6. Mus musculus. 7. Meriones tristrami. 8. Gerbilus dasyurus. 9. Microtus guentheri. 10. Cricetulus migratorius.
..!. ...... c:
Q.
...
Muhraqa
Bet Oren
'40 Oaks'
20 /5
'0 10
.a
E z
"
5 0
2
6
I
3
2
I
3 4 6 SPlcles
2
3 4
I
5
Figure 3. Same as Fig. 2 but for the Mt Carmel plots.
6
67
RODENT SPECIES DIVERSITY
..c
e
..
'u ~
'0 ..
~ Z
55 50 45 40 35 30 25 20 15
Acomys cohirinus
Hal/us rol/us
Mus musculus
.
10
5~
o
Apodtmus syll'oficus
Apodtmus mysfocinus
m ...
KF B E H
mmn . .
G E B H K
I!III!II_-
G K H B E
m
H C G E
In
K B E H
localities
Figure 4. Abundance of the most common rodent species (letters as in Table 1).
Discussion Among the rodent species that have been trapped during this study we recognize two species that are mostly typical of Mediterranean oak woodland, A. sylvaticus and A. mystacinus. They are found only when tree cover is comparatively high (105 per cent in Mt Meron, 106 per cent in Muhraqa; Table 2). However on the whole, total PVC does not seem to be an important factor for rodent species diversity. Thus, to our great surprise, in Basmat Tivon, where the PVC was rather high (117 per cent), no rodent was found. It seems also that plant species richness (PSR) was not correlated with rodent species diversity. (Thus PSR in Allonim was 106 with one rodent species and in Muhraqa PSR was low at 18 with RSD at 1·75.) On the other hand, some of the other species of rodents were found in different habitats irrespective of vegetation factors provided they contained certain elements. Thus A. cahirinus and G. dasyurus are rodent species occurring typically in rocky habitats and indeed total percentage of stone coverage (PSC) was highest in Gilboa (44 per cent, Table 2). It seems that rodent species could be similar to reptiles in their dependence upon the heterogeneity of !he habitat (Pianka, 1973). Thus both vegetation, climatic and edaphic factors are all of tmportance. This was recently shown by Brown (1973) and Beatley (1976) who found that, for desert rodents, climatic factors (mostly precipitation) that regulated plant growth and primary productivity have a good correlation with rodent species diversity. No indication for a similar correlation was found in the present study for the Mediterranean region. It is possible that for some species studied here PVC is of significance but not FHD or PSC. This study wassupported by a grant from the U.S.-Israel BinationalScience Foundation (BSF), Jerusalem, Israel. The botanical studies were conducted by a team headed by Professor Z. Naveh of this Institute. Some of the field work (during 1974 and part of 1975)was carried out by Mr D. Feigin, to whom we wish to express our thanks.
References BeatIey, J. C. (1976). Rainfall and fluctuating plant populations in relation to distributions and numbers of desert rodents in southern Nevada. Oecologia (Berlin), 24: 21-42. Brown, J. H. (1973). Species diversity of seed-eating desert rodents in sand dune habitats. Ecology, 54: 775-787. Goodman, D. (1975). The theory of diversity-stability relationships in ecology. Quarterly Review of Biology, 50: 237-266. HUrtubia, J. & DiCastri, F. (1973). Segregation of lizard niches in the Mediterranean region of Chile. In DiCastri, F. & Mooney, H. A. (Eds). Mediterranean Type Ecosystems, Origin and Structure, pp. 349-360. Berlin: Springer. MacArthur, R. H. (1965). Patterns of species diversity. Biological Reviews, 40: 510-533.
-
2·0
105·0
Mt Meron
Galil Mts
\
4·1
5·2 6·9 4·0
108 103 18 37
0·7
0·1
8·0
(%)
Stones
-
I
106 100 90
129
No. of plant species
PVC: percentage of vegetable cover; PSC: percentage of stone cover.
107·0
-
nooo)
109·8 98·1 108·5
13·4 17·8
5·7 8·3 2·5
90·7 72·0 106·0
'40 Oaks' Bet Oren Muhraqa
Carmel (750)
80·5 74·0 117·5
53·1 37·2 23'5
3·8 2·9 18·5
23·6 33·9 75·5
Allonim Allone Abba Basmat Tivon
Lower Galil (650)
66·4
6·9
29·0
30·5
Total
(%)
Annual plants
Herbaceous plants <0·5m
Woody plants >0·5m
•
Gilboa
experimental plots
Locality of
Perennials (%)
.
PVC
Gilboa Mts (5OO)
nun)
Region (precipitation,
I
I
Table 2. Vegetation, edaphic and climatic factors
17·2
21·3
5·9 19·0 7·7
6·0 1·5 12·4
5·9 1'5 11·7 0·7 12·1 3·7
34·0
Total stones and rocks 26'0
(%)
Rocks
•
PSC \
RODENT SPECIES DIVERSITY
69
McCloskey, R. T. (1976). Community structure in sympatric rodents. Ecology, 57: 728-739. Pianka, E. R. (1973). The structure of lizard communities. Annual Review of Ecology and Systematics, 4: 53-74. Rosenzweig, M. L. & Winakur, J. (1976). Population ecology of desert rodent communities: habitats and environmental complexity. Ecology, 50: 558-572. Schoener, T. W. (1969). Size patterns in West Indian Anolis lizards: 1. Size and species diversity. Systematic Zoology, 18: 386-401. Tomoff, C. S. (1974). Avian species diversity in desert scrub. Ecology, 55: 396-403. Whittaker, R. H. (1972). Evolution and measurement of species diversity. Taxonomy, 21: 213-251. Wiens, J. A. (1973). Pattern and process in grassland bird communities. Ecological Monographs, 43, 237-270. Willson, M. F. (1974). Avian community organization and habitat structure. Ecology, 55: 1017-1029.