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Original article
Abstract.—The diet of an Algerian population of spur-thighed tortoise (Testudo graeca) was studied with
the aims of exploring: (i) the variation in diet among males, females and juveniles, (ii) the relationships
between consumption and relative availability of the plant species, and (iii) which plant tissues, vegeta-
tive or reproductive, are eaten by tortoises. We recorded more than forty plant species at the study area,
with monocot species (n = 7) having a greater percent cover than that of dicot species (n = 31 species)
or Gymnospermae (n = 2). Tortoise diet was studied by categorising 4422 plant and animal fragments in
faecal pellets of 20 males, 16 females and eight juveniles. Tortoises ate a wide variety of plant species,
including 13 dicots and three monocots, and occasionally invertebrates. The number of fragments for a
plant species was correlated with plant species cover, and plant vegetative tissues exceeded plant repro-
ductive tissues in the faeces. Dicots (Fabaceae, Composeae, Primulaceae, and Caryophyllaceae)
accounted for over 70% of the diet (faecal fragments). The high dietary (niche) overlap, and null model
analysis (RA3 algorithm with 30,000 Monte Carlo simulations), indicate that males, females and juve-
niles did not partition food resources; all three groups ate the same plant species.
ost extant reptiles are carnivorous (croc- instructive for understanding other aspects of
M odiles, snakes, lizards, freshwater tur-
tles), with herbivory being unusual (in fresh-
life-history and evolution of these reptiles (e.g.,
Luiselli 2006), and scientists are beginning to
water turtles), rare (in lizards), exceptionally carefully analyse the dietary habits of testu-
rare (in crocodiles), or nonexistent (in snakes; dinids (e.g., MacDonald & Mushinsky 1988;
Halliday & Adler 2002). Terrestrial chelonians, Jennings 1993; Luiselli 2003; El Mouden et al.
especially members of the family Testudinidae, 2006). Such detailed dietary studies are neces-
are unusual among reptiles in that typically sary to help us understand the main patterns of
they are herbivorous, with few species being tortoise foraging ecology.
omnivorous or mainly carnivorous (Ernst &
Barbour 1989; Hailey et al. 2001; Luiselli The spur-thighed or Moorish tortoise (Testudo
2006). Studying foraging ecology may be very graeca graeca Linnaeus, 1758) has a wide dis-
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AFRICAN JOURNAL OF HERPETOLOGY 57(2) 2008
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ROUAG ET AL. — Food choice of Algerian tortoises
identified using keys in Quezel & Santa (1963). Carl Zeiss). According to Rodde’s technique,
three microscope slides were prepared for each
Tortoise diet.—We quantified T. graeca diet by sample, and fragments were counted on three
analysing faecal pellets collected in spring horizontal lines, separated by 2 mm, on each
2007, in exactly the same period as we did slide (El Mouden et al. 2006).
plant surveys. Faecal pellet analysis has been
widely employed on tortoises (e.g., MacDonald Most plant fragments in faeces were identified
& Mushinsky 1988; Luiselli 2003) because it is to species level using a photographic atlas of all
noninvasive and unlikely to harm low-density plant species identified at the study area; some
species (Luiselli 2006). However, this method were identifiable only to higher taxonomic lev-
may pose biases in that softer materials may be els. Identification followed Baumgartner &
digested so that they are unidentifiable in the Martin’s microhistological method (see Rodde
faecal fragments (Demuth & Buhlmann 1997; 1977, Chapuis 1980), which has been widely
Gunzburger 1999; Pincheira-Donoso 2008); used for analysing diets of mammalian herbi-
this bias has not been quantified. Faeces were vores. The micro-morphological criteria for
collected from wild tortoises during random plant identification included the shape, size and
walks throughout the study area. Once cap- general feature of cells, cellular inclusions,
tured, tortoises were marked individually by density and distribution of stomata, and the
carapace scale notching, sexed, and identified morphology of external epidermal structures.
as male, female or juvenile for intraspecific We were unable to prepare a photographic
analyses (e.g., Bury & Luckenbach 1977; identification atlas for invertebrates, so we lim-
Rouag et al. 2007). Adults were distinguished ited these identifications to the presence or
from juveniles based on the appearance of nar- absence of invertebrate tissue.
rower growth annuli that occur at maturity
(Castanet & Cheylan 1979; Diaz-Paniagua et Statistical analyses.—Using diet composition
al. 2001; Rouag et al. 2007). Tortoises typical- (% of total fragments), dietary diversity (H’,
ly defecated during capture and manipulation. sensu Spellerberg & Fedor 2003) and equitabil-
Their faecal pellets were collected, placed in ity indices (E) were calculated according to
ethanol (50% v/v), and stored in plastic vials Shannon & Weaver’s formulas (1949). H’ and
for subsequent analysis. Then the tortoises E differences among groups (males, females
were released at their capture points. To avoid and juveniles) were calculated by a matrix per-
pseudoreplication, only one pellet was collect- mutation test, contrasting the actual data matrix
ed per tortoise. with data generated by 5000 random boot-
straps, and then applying the Mantel’s test
Faecal pellets were air dried, homogenised in a (Manly 1997). Food niche overlap (Ojk) among
mortar, decoloured using Javel’s water groups was calculated by Pianka’s (1986) sym-
(Massemin 1992), and sieved under a series of metric equation, with values ranging from 0
0.4-0.8 mm sieves (Chapuis 1980). Fragments (no overlap) to 1 (total overlap). To test the
longer than 0.4 mm were dried again, washed hypothesis that there was intraspecific resource
in a 50% (v/v) glycerine-water solution and partitioning for food, we employed null model
repulverised to make a fine, homogeneous analysis (e.g., Gotelli and Graves 1996) on
sample, and added back to the sample to help niche overlaps. The occurrence of non-random
produce a large number of fragments (Rodde intraspecific food resource partitioning in T.
1977; Butet 1985). Rodde’s technique (Rodde graeca was tested by contrasting the original
1977) was used to identify and count all frag- data matrices with simulated data matrices gen-
ments via light microscopy (x100; Microscope erated by Monte Carlo simulations (Gotelli &
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AFRICAN JOURNAL OF HERPETOLOGY 57(2) 2008
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ROUAG ET AL. — Food choice of Algerian tortoises
Table 1. Relative cover indices* of plant species in the 30 ha study area of the Park National d'El Kala, in
northeastern Algeria, April 2007.
Dicotyledons
Family Species
Amaranthaceae Amaranthus albus +
Apiaceae Chimaphila umbellata +
Daucus carota +
Other unidentified Apiaceae +
Apocynaceae Nerium oleander +
Asteraceae Centaurea napifolia +
Chrysanthemum coronarium 1
Evax pygmaea 1
Galactites tomentosa +
Ormenix mixta 1
Unidentified Asteraceae 2
Boraginaceae Cynoglossum sp. 1
Echium plantagineum 1
Campanulaceae Campanula dichotoma +
Caryophyllaceae Paronychia argentea 1
Cistaceae Tuberaria guttata 1
Euphorbiaceae Euphorbia sp. +
Fabaceae Coronilla scorpioides 1
Lotus ornithopodioides 1
Trifolium arvense 1
Trifolium campestre 1
Trifolium sp. 2
Lamiaceae Lamium sp. +
Mentha pulegium +
Mentha rotundifolia +
Plantaginaceae Linaria pinifolia 2
Polygonaceae Rumex bucephalophorus 1
Primulaceae Anagalis arvensis 1
Rosaceae Rubus ulmifolius 2
Rubiaceae Rubia peregrina +
Thymelaeaceae Daphne gnidium +
Monocotyledons
Family Species
Alliaceae Allium triquetrum 1
Arecaceae Chamerops humilis +
Cyperaceae Carex remota 1
Iridaceae Iris pseudacorus 1
Poaceae Cynodon dactylon 4
Hordeum murinum 4
Smilacaceae Smilax aspera 1
Gymnospermae
Pteridophyta
Family Species
Polypodiaceae Polypodium vulgare 2
Pinophyta
Family Species
Cupressaceae Juniperus oxycedrus +
* Braun-Blanquet & Pavillard (1929), the relative abundance scores represent: +, species present but very
rare (<1% of study area), and 1 to 5 respectively, for species covering less than 5% (score = 1), 5-25% (2),
25-50% (3), 50-75% (4), or 75 to 100% (5).
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AFRICAN JOURNAL OF HERPETOLOGY 57(2) 2008
Table 2. Relative abundance (%) of fragments identified in the faeces of 20 male, 16 female and eight juve-
nile Testudo greaca. The number of fragments counted was 1700, 1516 and 1206 for males, females and
juveniles, respectively. The vegetative parts included leaves and stems. The reproductive parts included
flowers and seeds.
Monocotyledons
Cynodon dactylon
leaves 11.88 8.97 9.95
stems 2.53 2.11 2.74
Hordeum murinum
leaves 8.06 10.29 7.79
stems 2.00 2.31 1.66
Carex remota
leaves 3.53 9.04 -
stems 1.71 2.84 -
Monocot vegetative 29.71 35.55 22.14
Monocot total 29.71 35.55 22.14
Dicotyledons
Anagalis arvensis
leaves 2.71 3.76 1.82
stems 0.88 1.52 0.91
seeds 0.35 1.98 1.00
Apiaceae indet.
leaves 2.47 0.73 1.82
stems - - 0.50
Asteraceae
leaves 2.65 1.58 2.82
stems 9.06 3.96 0.50
Chrysanthemum coronarium
leaves 3.06 1.65 2.82
stems - 0.20 0.83
Coronilla scorpioides
leaves 1.06 1.85 1.91
stems - 0.20 2.82
seeds 0.65 1.72 0.75
Cynoglossum sp.
leaves 3.12 0.99 2.32
stems - 0.33 1.24
Linaria pinifolia
leaves 0.71 1.32 1.24
stems - 0.13 -
seeds 0.29 1.12 0.17
Lotus ornithopodioides
leaves 4.82 1.52 1.66
stems 0.76 0.66 1.49
flowers 2.12 1.72 1.82
Ormenix mixta
leaves 0.35 - -
stems 1.53 - -
Paronychia argentea
leaves 1.18 1.19 1.41
stems - 0.13 0.66
flowers 1.53 1.32 2.07
Trifolium arvense
leaves 1.65 1.58 2.40
stems 0.47 0.66 0.75
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ROUAG ET AL. — Food choice of Algerian tortoises
Table 2 continued.
Trifolium sp.
leaves 13.24 10.03 11.44
stems 3.53 2.31 3.57
Tuberaria guttata
leaves 1.88 3.23 4.56
stems 2.70 3.82 2.82
seeds 0.59 2.11 2.57
Dicot vegetative 57.82 43.34 52.32
Dicot reproductive 5.53 9.96 8.37
Dicot total 63.35 53.30 60.70
(i.e., Cynodon dactylon and Hordeum mur- using the Bonferroni-adjusted criterion
inum) were important food items (Table 2) and (P=0.016), there were no sexual (Χ21= 14.93, P
covered considerable area (Cover index = 4). = 0.037) or adult-juvenile (Χ21= 16.05, P =
Several species, such as Paronychia argentea, 0.025) differences in seed fragments or inverte-
Cynoglossum sp., and Chrysanthemum coro- brate fragments (sexes: Χ21= 5.101, P = 0.024);
narium (Table 2), were rare in the environment adult-juvenile: Χ21= 3.887, P = 0.049).
(scored + or 1) and faeces. However, tortoises
ate some rare species disproportionate to their Diet diversity was nearly identical in males (H’
availability. For instance, among the species = 3.693), females (H’ = 3.712) and juveniles
with abundance score = 1, Carex remota and (H’ = 3.702; differences not significant, P >
Lotus ornithopodiodes were consumed regular- 0.25 with Mantel test). Equitability was also
ly (Table 2). Overall, species consumption (% relatively high, and not statistically different
of faecal fragments) was correlated to plant among males (E = 0.886), females (E = 0.908),
species availability (rs = 0.57, n = 16, P < 0.01). and juveniles (E = 0.925; P > 0.15 with Mantel
test). Food niche overlap values were very high
Dietary comparisons among groups.—The fre- between males and females (Ojk = 0.972),
quency of faecal fragments seemed similar males and juveniles (Ojk = 0.986), as well as
among groups (Table 2). For monocots, neither females and juveniles (Ojk = 0.957). The mean
the male-female nor adult-juvenile comparison observed index (Ojk = 0.972) was greater than
was significant (both Χ21< 4.47, P > 0.480). the simulated values (Ojk = 0.459±0.019) in
Similarly, the frequency of vegetative and 29,997 simulations out of 30,000. The Pobs<exp
reproductive plant tissues, whether or not = 0.999, thus rejecting the hypothesis of an
including monocots and dicots, did not differ intraspecific resource partitioning for food.
between sexes (Χ21= 2.118, P = 0.548) or Furthermore, the observed index was signifi-
between juveniles, males and females (Χ22= cantly higher than the simulated value (P <
6.097, P = 0.107). The data showed a slight 0.0001), suggesting all groups ate the same
suggestion that seeds and invertebrates were foods.
consumed differently among groups. However,
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AFRICAN JOURNAL OF HERPETOLOGY 57(2) 2008
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ROUAG ET AL. — Food choice of Algerian tortoises
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