J. Moll. Stud.

(1999), 65, 411–423 © The Malacological Society of London 1999

FIELD OBSERVATIONS ON FEEDING OF THE LAND SNAIL

HELIX ASPERSA MÜLLER

J. IGLESIAS and J. CASTILLEJO

Department of Animal Biology, Faculty of Biology, University of Santiago de Compostela,
15706 Santiago de Compostela, Spain
(Received 16 March 1998; accepted 30 November 1998)

ABSTRACT The fact that many land gastropods are capable

of eating a wide variety of living and dead
Feeding of the land snail Helix aspersa (Müller) was plants and of adjusting their diet to the avail-
observed at monthly intervals. Three natural popula- ability of the food items might determine the
tions in Galicia (NW-Spain) were studied. At two
wide distribution of many species (Pallant,
sites only a few plants constituted the bulk of the diet
and in spring the snails’ diet had the highest diversity 1972).
(H9). In the third population feeding and distribution Nevertheless, land gastropods show different
of Helix aspersa (Müller) were observed in a small levels of preference for the different com-
plot with permanent patches of Urtica dioica. Nearly ponents of their diet, as demonstrated in a
one half of feeding snails fed upon Urtica dioica. Most number of field (Grime & Blythe, 1969; Pallant,
of the other observations were on Mentha suaveolens, 1969, 1972; Wolda et al., 1971; Williamson &
Ranunculus repens and Gramineae. The diversity of Cameron, 1976; Dan, 1978; Richter, 1979;
the snails’ diet showed seasonal variation with the Speiser & Rowell-Rahier, 1991; Hatziioannou
maximum in the autumn months. Comparison between
et al., 1994) and laboratory studies (Grime,
the availability of the different plant species and their
contribution to the snails’ diet showed that the snails MacPherson-Stewart & Dearman, 1968; Grime,
did not eat at random; Urtica dioica was eaten much Blythe & Thornthon, 1970; Cates & Orians,
more than expected from its occurrence and grasses 1975; Dirzo, 1980; Richardson & Whittaker,
were strongly under-represented in the snails’ diet. 1982; Rathcke, 1985; Mølgaard, 1986; Gallois &
Temporal changes of availability were significantly Daguzan, 1989; Baur, Baur & Fröberg, 1994).
correlated with the amounts eaten in the case of Food preferences determine the food inges-
Urtica, but not for the other food plants. The distribu- ted by an animal and hence affect its physio-
tion of the snails in the plot was significantly corre- logical condition and fitness (Carefoot, 1967;
lated with that of Urtica. Chemical analyses of the
Vadas, 1977; Batzli & Lesieutre, 1991). The
food plants revealed Urtica as the species with the
higher protein, ash and calcium contents. The strong concept of ‘food preference’ usually comprises
preference of Helix aspersa for Urtica dioica could be two features of the food, attractiveness and
explained by the value of Urtica as food or by its suit- palatability. Attractiveness is experimentally
ability as habitat for the snails. The largest propor- measured with choice tests (Dan, 1978; Nicotri,
tions of green material in the snails’ diet occurred in 1980; Rathcke, 1985; Szlavecz, 1986; Gallois &
the spring and juveniles ate more green material than Daguzan, 1989). Palatability is often measured
adults in the three populations. by comparing the quantity of one food item
ingested by the consumer in relation to another
food (Grime et al., 1968; Cates & Orians, 1975;
INTRODUCTION Mølgaard, 1986; Richardson & Whittaker, 1982;
Chang, 1991; Egonmwan, 1991) or by means of
Land gastropods have traditionally been assimilation rates (Mason, 1970; Jennings &
regarded as generalist herbivores taking a wide Barkham, 1976; Williamson & Cameron, 1976;
variety of plants from the environment (Boy- Dan, 1978; Staikou & Lazaridou-Dimitriadou,
cott, 1934; Grime & Blythe, 1969; Mason, 1970; 1989). Vadas (1977) investigated several features
Wolda, Zweep & Schuitema, 1971; Richardson, related to food preference in sea urchins like
1975; Jennings & Barkham, 1975; Williamson & chemoreception, feeding rates, assimilation
Cameron, 1976; Chatfield, 1976; Chang, 1991; efficiency, calorific value of the foods growth
Speiser & Rowell-Rahier, 1991; Hatziioannou, and reproductive capacities of the consumers
Eleutheriadis & Lazaridou-Dimitriadou, 1994). on different foods, and availability of prey
412 J. IGLESIAS & J. CASTILLEJO
items. He concludes that feeding in nature is a The main observation site for population Lapido-I
compromise between preferences of the con- was an unsurfaced road 90 m long and 4 m wide, the
sumer and availability of prey items. vegetation of which consisted mainly of perennial
The habitat or microhabitat conditions which herbs including Ranunculus repens, Juncus effusus,
Mentha suaveolens, Plantago lanceolata and Sonchus
food plants provide to the consumer may influ- asper, and grasses including Holcus sp., Arrhen-
ence its feeding habits (Nicotri, 1980), a feature atherum sp., Anthoxanthum sp., Avena sp., Agrostis
which has not received much attention in gas- sp., Briza sp. and Dactylis sp.
tropods. If any plant species offers to the grazer For population Lapido-II the main observation site
protection from predators, good resting plates, was a triangular plot of 91 m2 delimited by the wall of
egg-laying sites, overwintering sites, a particular a house, by a road three meters wide and by an unsur-
microclimate or any other favourable condition faced road two meters wide; it was about 400 metres
for its life cycle, this might influence the food away from the site of population Lapido-I. The
preference of the consumer. This is specially vegetation of this plot was very dense but with low
diversity, since it consisted almost exclusively of
true in small animals like land gastropods with Mentha suaveolens, Ranunculus repens, Urtica dioica
low mobility and little capacity to defend them- and grasses including Dactylis sp., Arrhenaterum sp.,
selves against predators. Holcus sp. and Anthoxanthum sp. Some specimens of
This paper describes the feeding of the land Cyperus longus, Lotus corniculatus, Juncus effuses
snail Helix aspersa in natural populations on the and Plantago lanceolata were also present, but in very
basis of field observations. The importance of low abundance. The plot was frequently visited
direct visual observations in the field for under- before the start of sampling in July 1991, and the
standing feeding of animals in natural con- existence of a well established population of Helix
ditions has been emphasized (Richardson, 1975; aspersa and of permanent patches of Urtica dioica
were confirmed. In June 1991, five zones named A, B,
Vadas, 1977; Chang, 1991), but most previous C, D (18 m2 each) and zone E (19 m2) were delimited
studies on snails’ diet have been based on labo- with red cords tied to wooden stakes. There were no
ratory feeding experiments or analysis of faeces barriers to prevent snails moving inside the plot or to
or crop contents. Since the stinging nettle Urtica leave it.
dioica is often reported as a preferred food
plant of land gastropods (Grime et al., 1968;
Grime & Blythe, 1969; Pallant, 1969; Mason, Sampling
1970; Grime et al., 1970; Wolda et al., 1971; Cobas and Lapido-I populations were visited monthly
Cates & Orians, 1975; Jennings & Barkham, from September 1990 to September 1992 (except for
1975, 1976; Chatfield, 1976; Dan, 1978; Carter, May 1991, when only the site of Cobas was visited) on
Jeffery & Williamson, 1979; Staikou & Lazari- three consecutive nights. Lapido-II population was
dou-Dimitriadou, 1989; Lazaridou-Dimitriadou visited monthly on two consecutive nights from July
1991 to September 1992. Each observation site was
& Kattoulas, 1991; Hatziioannou et al., 1994), carefully explored for 3 hours at least, searching for
we also decided to study in situ whether Helix snails by torchlight. When a snail was seen, we
aspersa really prefers Urtica over other plant recorded the plant species and whether the snail ate a
species of the environment, and, if so, to search green or a senescent plant. In Lapido-II we recorded
for the causes of that preference. also the position (zone) of the snail. Only observa-
tions made on snails feeding upon living or senescent
plants were noted. Snails eating dead plant material
on the ground, mosses, lichens, fungi, faeces of verte-
MATERIALS AND METHODS
brates and dead arthropods were also seen, but they
were not recorded quantitatively. Observations were
Study area
noted only when the animals were not disturbed by
The study was carried out in two localities in Galicia the observer and when consumption of food was
(NW-Spain), named Cobas and Lapido. The study certain from the feeding movements of the animal
areas, the climate and the annual activity cycles of and/or the feeding marks on the plant. Plant nomen-
Helix aspersa at these localities were described in clature was based on Tutin, Heywood, Burges,
detail by Iglesias, Santos & Castillejo (1996). The Moore, Valentine, Walters & Webb (1964), and grass
three populations will be referred as Cobas, Lapido-I species were pooled. The presence of a solid reflected
and Lapido-II. The main observation site in Cobas, a lip on the peristome of the shell was taken to denote
plot of 400 m2, was abandoned land, the vegetation of adulthood of snails.
which consisted of woody species about 2 m tall, On the day previous to the first night of sampling in
including Crataegus monogyna, Prunus spinosa and Lapido-II, the relative abundance (percentage cover)
Rosa spp., herbs which included mainly Sanguisorba of the plant species was measured by means of a
minor, Potentilla splendens, Geum sylvaticum, Eryn- stratified sampling on 45 points of the plot (9 points/
gium campestre, Fragaria vesca, Origanum virens and zone). Samples of leaves of the most abundant plant
grasses including Bromus sp., Poa sp. and Lolium sp. species in this plot (Mentha, Ranunculus, Urtica and
 FEEDING OF HELIX ASPERSA IN THE FIELD 413
Gramineae) were collected on four occasions and ranges between 21 and 11, positive values indicating
analized for crude protein (Kjeldahl method), raw preference, negative values indicating avoidance and
fibre (Wender method), soluble sugars (colorimetry), C 5 0 for random feeding.
ash (550°C for 4 hours) and calcium (atomic absorp- In order to examine any relationship between the
tion). distribution of the snails and that of the plant species,
the percentage distribution of adult and juvenile
Data analysis snails and the percentage distribution of each plant
species over the five zones were compared by means
Diversity of the snails’ diet and of the vegetation of scatter diagrams. The existence of association
was determined for each sampling by means of the between the two variables was tested with Kendall’s
Shannon-Wiener index, H95 2S(pi)(log2pi), (Krebs, rank correlation coefficient, t. The same coefficient
1986) where pi was the frequency of each plant was used to investigate any relationship between
species in the snails’ diet or in the plot. To compare abundance of each plant species and the amount
the diversity of adult and juvenile snails’ diet, the eaten by the snails.
Mann-Whitney test was performed. A x2 test with
contingency tables was executed to investigate differ-
ences between adults and juveniles in relation to
green or senescent material. ANOVA and Fisher RESULTS
LSD tests, with arc-sine transformation of the data
(Sokal & Rohlf, 1981) were performed to compare
protein, fibre, soluble sugars, ashes and calcium con- Snails’ diet in the field
tents of the different plant species. In Cobas, a total of 833 observations of snails
With the data of the snails’ diet composition and feeding upon 25 plant species was made (Table
the data of relative abundance of the plants (% cover)
in the plot Lapido-II, an electivity index C, according
1). The most frequent foods in the diet of Helix
to Pearre (1982), was calculated for adults and juve- aspersa in this site were Fragaria vesca (17.28%
niles in each sampling. The index C is based on x2 and of the observations), Gramineae (15.96%),
provides a simple way to test the significance of any Trifolium angustifolium (12.24%), Hieracium
degree of selection at any sample size: the value of C pilosella (10.2%), Eryngium campestre (7.32%)

Table 1. Field observations on feeding of Helix aspersa in Cobas.

Adults Juveniles

Plants Green Senescent Green Senescent %

Fragaria vesca 23 46 36 39 17.28

Gramineae 10 58 17 48 15.96
Trifolium angustifolium 14 37 26 25 12.24
Hieracium pilosella 14 9 51 11 10.20
Eryngium campestre 6 28 8 19 7.32
Carlina corymbosa 5 32 7 16 7.20
Sanguisorba minor 5 3 25 13 5.52
Potentilla splendens 3 5 22 3 3.96
Taraxacum officinale 6 2 14 2 2.88
Plantago lanceolata 7 14 2.52
Daucus carota 16 5 2.52
Rubus spp. 3 13 2 2 2.40
Origanum virens 3 6 9 1 2.28
Crataegus monogyna 5 5 2 4 1.92
Sedum forsteramum 10 1.20
Foeniculum vulgare 5 5 1.20
Geum sylvaticum 3 4 0.84
Trifolium pratense 1 6 0.84
Lathyrus sp. 2 1 0.36
Vicia sativa 3 0.36
Prunus spinosa 3 0.36
Campanula lusitanica 1 1 0.24
Silene vulgaris 1 0.12
Lavandula stoechas 1 0.12
Rumex acetosa 1 0.12
% 13.32 32.17 31.33 23.17
414 J. IGLESIAS & J. CASTILLEJO
and Carlina corymbosa (7.2%); these six plant tion of the most frequent plant in each sampling
species represented more than 70% of the feed- was in general about 20%.
ing observations made in Cobas over the study In Lapido-II, a total of 958 observations of
period. Senescent grasses were the most abun- snails feeding were noted during the study
dant material in the diet of the snails at Cobas period. 95.6% of the observations were made
during the autumn, accounting for more than upon the four major foods, Gramineae, Mentha
30% of the observations in September 1990 and suaveolens, Ranunculus repens and Urtica
November 1991, but in spring, hardly any senes- dioica, and only a 4.4% were made upon very
cent grasses were eaten. During the spring scarce plant species in the plot like Cyperus
periods the maximum proportion of one plant longus, Lotus corniculatus, Juncus effusus and
species in the diet of the snails was Fragaria Plantago lanceolata (Table 3). The most fre-
vesca, with percentages around 20%. quent food in the diet of Helix aspersa in this
In Lapido-I, a total of 708 observations of site was Urtica dioica with 47.9% of the obser-
snails feeding upon 24 plant species was made vations; its contribution to the snails’ diet
(Table 2). Mentha suaveolens (20.48%), Ram- ranged from 61.9% in August 1991 to 29.8% in
unculus repens (15.39%), Gramineae (15.39%), November 1991.
Heracleum sphondylium (8.05%), Plantago For the whole study period, 44.65% of the
lanceolata (7.76%) and Apium nodiflorum feeding observations in Cobas were made on
(7.48%) were the most abundant foods, repre- green material and 55.35% on senescent plants;
senting more than 70% of the feeding observa- in Lapido-I 67.5% of the observations were on
tions made in this plot over the study period. green material and 32.5% on senescent plants;
There was not a clear seasonal variation in the in Lapido-II 71.5% of the observations were on
contribution of any plant species to the diet of green plants and 28.5% on senescent plants.
the snails in Lapido-I; the highest contribution The proportions of adults and juveniles feeding
of a single plant to the diet was that of Mentha upon green and senescent material during each
suaveolens in November 1991, but the contribu- sampling are shown in Fig. 1. There was a clear

Table 2. Field observations on feeding of Helix aspersa in Lapido-I.

Adults Juveniles

Plants Green Senescent Green Senescent %

Mentha suaveolens 29 29 78 9 20.48

Gramineae 13 40 34 22 15.39
Ranunculus repens 17 24 47 21 15.39
Heracleum sphondylium 33 24 8.05
Plantago lanceolata 19 36 7.76
Apium nodiflorum 6 47 7.48
Juncus effusus 12 6 6 3 3.81
Lotus corniculatus 3 15 1 2.68
Sonchus asper 7 8 1 2.25
Hypochoerys radicata 8 7 2.12
Coleostephus myconis 5 9 1.97
Trifolium repens 4 10 1.97
Cyperus longus 5 8 1.83
Rumex obtusifolius 3 9 1 1.83
Plantago major 2 8 1 1.55
Pentaglottis sempervirens 2 7 1.27
Conyza canadensis 2 7 1.27
Polygonum persicaria 2 1 2 0.70
Eupatorium cannabinum 2 1 0.42
Cardamine pratensis 3 0.42
Hypericum perforatum 3 0.42
Rubus spp. 2 1 0.42
Sesamoides pygmaea 1 1 0.28
Lithodora diffusa 1 0.14
% 19.35 19.63 48.16 12.85
 FEEDING OF HELIX ASPERSA IN THE FIELD 415

Table 3. Field observations on feeding of Helix aspersa in Lapido-II. Others includes Juncus
effusus, Cyperus longus, Lotus corniculatus and Plantago lanceolata.

Adults Juveniles

Plants Green Senescent Green Senescent %

Urtica dioica 67 39 266 87 47.91

Gramineae 32 25 88 40 19.31
Mentha suaveolens 28 14 92 28 16.91
Ranunculus repens 18 15 59 18 11.48
Others 10 2 25 5 4.38
% 16.18 9.91 55.32 18.58

Figure 1. Percentages of field observations of adult (left bars) and juvenile (right bars) snails feeding upon
green and senescent material during each sampling in the sites Cobas, Lapido-I and Lapido-II.
416 J. IGLESIAS & J. CASTILLEJO
seasonal variation in the diet with respect to and in the abundance (% cover) of Urtica
green or senescent material; snails ate mainly dioica along the study period are shown in
green plants during spring, and senescent mate- Fig. 3, from which it seems that the snails’ diet
rial during the autumn. Juveniles ate more green diversity follows an opposite trend to that of
material than adults in spring (Cobas x2 5 35.08, the vegetation. Also, the diversity of the snails’
P , 0.001; Lapido-I x2 5 25.95, P , 0.001), in diet increased when the abundance of Urtica
summer (Lapido-I x2 5 20.03, P , 0.001) and in decreased and vice versa. Kendall’s rank corre-
autumn (Cobas x2 5 14.29, P , 0.001; Lapido-I lation test showed that there was no significant
x2 5 24.73, P , 0.001). In Lapido-II juveniles association between any of these variables.
ate more green material than adults over the Mann-Whitney test showed no statistical dif-
whole study period (x2 5 14.99, P , 0.001), but ference between adults and juveniles in relation
for the different seasons differences were only to diet diversity in each site.
significant in summer (x2 5 10.89. P , 0.001).
Snails’ diet in relation to the relative abundance
Diversity of vegetation and of the snails’ diet of the plants in Lapido-II
The diversity of the snails’ diet for the whole Urtica dioica was preferred by Helix aspersa
study period was H9 5 3.7 in Cobas and H9 5 over the other plant species in the plot (Fig. 4).
3.63 in Lapido-I, but there was a seasonal varia- Ranunculus repens and Mentha suaveolens were
tion (Fig. 2) with the maximum diversity in in a similar rank of abundance as Urtica, but
spring and the minimum in autumn. their contribution to the snails’ diet was smaller.
The diversity of the diet in Lapido-II was Gramineae were the most available plant
H9 5 1.95; there was also a seasonal variation species in the plot, but they were also eaten in a
with the maximum diversity in the autumnal smaller proportion than Urtica.
months. The changes in the diversity of the For the whole population a significant posi-
snails’ diet, in the diversity of the vegetation tive association existed between the abundance

Figure 2. Changes in the diversity of the snails’ diet in the sites of Cobas and Lapido-I during the study period.

Figure 3. Changes in the diversity of the snails’ diet, in the diversity of the vegetation, and in the abundance
(% cover) of Urtica dioica along the study period in Lapido-II.
 FEEDING OF HELIX ASPERSA IN THE FIELD 417

Figure 4. Scatter diagrams for the relative abundance (% cover) and the contribution to the snails’ diet of each
plant species, for adult and juvenile snails in Lapido-II. Each point represents one sampling occasion. GRAM
5 Gramineae, MSUA 5 Mentha suaveolens, UDIO 5 Urtica dioica, RREP 5 Ranunculus repens.

Table 4. Monthly values of the electivity index C for adult and juvenile snails and for
the commonest plant species of the site Lapido-II. * 5 P < 0.05, ** 5 P < 0.01. N.S. 5
not significant.

U.dioica Gramineae M.suaveolens R.repens

Jul-91 Adults 0.15 ** 20.10 * 20.01 N.S. 0.01 N.S.

Juveniles 0.45 ** 20.21 ** 20.08 N.S. 20.03 N.S.
Aug-91 Adults 0.34 ** 20.23 ** 0.00 N.S. 20.04 N.S.
Juveniles 0.36 ** 20.26 ** 0.02 N.S. 20.01 N.S.
Sep-91 Adults 0.33 ** 20.19 ** 0.01 N.S. 0.00 N.S.
Juveniles 0.50 ** 20.29 ** 0.01 N.S. 20.04 N.S.
Oct-91 Adults 0.23 ** 20.14 ** 0.01 N.S. 0.00 N.S.
Juveniles 0.28 ** 20.31 ** 0.18 ** 0.07 N.S.
Nov-91 Adults 0.11 * 20.13 * 0.10 N.S. 0.02 N.S.
Juveniles 0.30 ** 20.21 ** 0.12 * 20.02 N.S.
Apr-92 Adults 0.24 ** 20.19 ** 0.01 N.S. 0.01 N.S.
Juveniles 0.35 ** 20.32 ** 0.12 * 0.03 N.S.
May-92 Adults 0.25 ** 20.13 ** 20.04 N.S. 0.02 N.S.
Juveniles 0.37 ** 20.29 ** 0.03 N.S. 0.00 N.S.
Jun-92 Adults 0.25 ** 20.13 ** 20.03 N.S. 0.00 N.S.
Juveniles 0.34 ** 20.17 ** 0.00 N.S. 20.07 N.S.
Jul-92 Adults 0.24 ** 20.15 ** 0.01 N.S. 20.01 N.S.
Juveniles 0.36 ** 20.16 ** 20.09 N.S. 20.03 N.S.
Aug-92 Adults 0.10 * 20.13 ** 0.06 N.S. 20.01 N.S.
Juveniles 0.43 ** 20.29 ** 20.01 N.S. 20.06 N.S.
Sep-92 Adults 0.10 * 20.10 * 0.04 N.S. 0.01 N.S.
Juveniles 0.39 ** 20.30 ** 20.01 N.S. 20.04 N.S.

of Urtica and the amount eaten by the snails The monthly values of the electivity index C
(t 5 0.636, P , 0.01, n 5 11), but for the rest for adult and juvenile snails are shown in Table
of the plant species there was no significant 4. The values of the index C, indicate that both
relationship between abundance and the adult and juvenile H. aspersa show a strong
amounts eaten. The association between abun- preference for U. dioica and a strong avoidance
dance of plant species and amount eaten was for Gramineae. For Ranunculus repens, the
significant only in the case of juveniles and values of the electivity index were always
Urtica (t 5 0.527, P , 0.05, n 5 11). around zero, indicating that this plant species
418 J. IGLESIAS & J. CASTILLEJO
was eaten proportionally to its abundance. nied by an increase in the abundance of snails
Mentha suaveolens was neither preferred nor (adults and juveniles in zone C, only juveniles in
avoided by adult snails, but juveniles showed zone E).
preference for this plant species in October There existed a significant negative relation
and November 1991 and also in April 1992, among the distribution of Urtica and that of
coinciding with low abundance of Urtica. Gramineae in the plot (t 5 20.670, P , 0.001, n
5 55), and therefore the percentage distribu-
tion of snails also showed a significant negative
association with the percentage distribution of
Spatial distribution of the snails in relation to
Gramineae among the five zones of the plot
the relative abundance of the plants in Lapido-II
(for adults, t 5 20.613, P , 0.001; for juveniles,
The scatter diagrams for the percentage distri- t 5 20.651, P , 0.001; n 5 55 in both cases).
bution of Urtica dioica and the percentage dis- Furthermore, the distribution of the snails was
tribution of snails among the five zones of the never significantly related to the distribution of
plot on each sampling occasion are shown in Mentha suaveolens and Ranunculus repens.
Fig. 5. There was a significant positive associa-
tion between the percentage distribution of
Composition of the plant species
snails among the five zones of the plot and that
of Urtica dioica, both for adult (t 5 0.742, P Table 5 shows the contents of protein, fibre,
, 0.001, n 5 55) and juvenile snails (t 5 0.734, soluble sugars, ashes and calcium for the four
P , 0.001, n 5 55). commonest plant species of the plot Lapido-II.
The distribution of snails over time and that Fisher LSD tests showed that Urtica dioica
of Urtica were strongly related; the abundance differed statistically (P , 0.05) from the other
of snails in zones A and B, where Urtica was plant species because of its high protein, ashes
relatively abundant over the study period, was and calcium contents; Gramineae were differ-
not related to the temporal changes in the entiated because of their high fibre and low
abundance of nettles; in zones C and E, where calcium contents, and Ranunculus repens was
Urtica was absent (zone C) or scarce (zone E) differentiated because of its high soluble sugars
before the winter, the increase in the abun- content. No other statistical differences were
dance of nettles after the winter was accompa- found.

Figure 5. Scatter diagrams for the percentage distribution of Urtica dioica and the percentage distribution of
adult and juvenile snails among the five zones of the plot Lapido-II. Each point represents one sampling occa-
sion; in the abscissa the percentage of Urtica in one zone with respect to the cover of Urtica in the whole plot,
and in the ordinate the percentage of snails in the same zone and the same sampling with respect to the total
number of snails in the whole plot.
 FEEDING OF HELIX ASPERSA IN THE FIELD 419
Table 5. Protein, fibre, soluble sugars, ashes and calcium contents of the four commonest plant
species of the site Lapido-II. values are % dry weight (mean 6 S.D.), sample size 5 4 in all cases except
soluble sugars in M. suaveolens and R. repens, where n 5 3. Figures within the same column followed
by the same letter are not significantly different (P , 0.05) by Fisher LSD test.

Crude Soluble
Plant species Calcium Ash protein Raw fibre sugars

U. dioica 3.83 6 0.84 a 20.09 6 4.16a 30.50 6 1.77a 12.30 6 0.97a 05.11 6 2.79a
M.suaveolens 1.62 6 0.60b 13.05 6 2.14b 22.01 6 4.63b 10.89 6 3.77a 02.57 6 0.75a
R. repens 1.21 6 0.13b 11.65 6 1.47bc 22.12 6 4.18b 12.76 6 1.40a 11.44 6 2.30b
Gramineae 0.51 6 0.13c 09.23 6 2.04c 20.49 6 4.44b 26.96 6 2.68b 03.49 6 0.87a

DISCUSSION The avoidance of grasses by slugs and snails

has frequently been shown (Grime et al., 1970;
The results of this work agree well with previ- Wolda et al., 1971; Williamson & Cameron,
ous studies on food and feeding of land gas- 1976; Carter, Jeffery & Williamson, 1979;
tropods. For a given species of land gastropod, Cottam, 1985; Szlavecz, 1986; Chang, 1991;
the range of foods eaten under natural condi- Speiser & Rowell-Rahier, 1991), although a
tions may vary widely depending on the food number of studies pointed out the grasses as the
available (South, 1992). For example, the slug main diet components of some species like
Deroceras reticulatum feeds mainly on Ranun- Cepaea nemoralis (Grime & Blythe, 1969),
culus repens and Urtica dioica in a oak grove Deroceras reticulatum (Pallant, 1972) and Helix
(Pallant, 1969), and on grasses in rough grass- aspersa (Dan, 1978). In these three studies, the
land (Pallant, 1972). Grime & Blythe (1969) Gramineae were the most abundant plants in
found that senescent grasses form the bulk of the habitat and their predominance in the diet
the diet of Cepaea nemoralis in grassland, but probably arises from high availability. Also, the
animals living in a nettle (Urtica dioica) patch three studies were based on the frequency of
eat large proportions of green Urtica. The same presence of each food item in faeces or stomach
was found for Monacha cantiana by Chatfield contents, and grasses are the most readily iden-
(1976). tifiable and detectable plants in the analyses
The fact that plants which are frequent in the due to the shape and arrangement of the stom-
field are also eaten frequently and the occur- ata and due to low degradation in the digestive
rence of seasonal changes in the diet related to process (Grime & Blythe, 1969; Chatfield, 1976;
availability have been reported for other Jennings & Carkham, 1976). The fact that
species of slugs and snails (Chatfield, 1976; grasses comprised the bulk of the diet does not
Williamson & Cameron, 1976; Richter, 1979; mean that they were preferred food. In the
Szlavecz, 1986; Speiser & Rowell-Rahier, 1991; present study grasses were also abundant in the
Hatziioannou et al. 1994). The seasonal changes three sites and they always occupied the second
in the percentages of green material in the diet place in the rank of feeding observations,
reported here probably reflect also seasonal although the values of the electivity index C
changes of the plant community and in the clearly show a strong avoidance for Gramineae
abundance of different food items: during in site Lapido-II. The rejection of grasses by
spring, when a variety of green plants were slugs and snails is usually attributed to the high
present, the snails’ diet showed the highest per- silica content of the leaves (Wadham & Wynn
centage of green material. The same applies for Parry, 1981) or to their toughness (Grime et al.,
diet diversity in the sites Cobas and Lapido-I 1968; Dirzo, 1980; Reingold & Gelperin, 1980;
where maximum diet diversity occurred in Speiser & Rowell-Rahier, 1991).
spring. On the contrary, diet diversity in On the other hand a number of studies have
Lapido-II followed an opposite trend to that of shown that Urtica dioica is one of the most pre-
diversity of vegetation, and it seems that diet ferred food plant for snails and slugs. Stinging
diversity in this plot was related to the abun- nettles are highly acceptable for Cepaea
dance of the highly preferred food Urtica nemoralis (Grime et al., 1968, 1970; Carter et al.
dioica: low abundance of Urtica coincided with 1979), Cepaea hortensis (Carter et al. 1979),
high diet diversity and vice versa, although Deroceras reticulatum (Pallant, 1969), Arion
there was no significant relationship between ater (Cates & Orians, 1975; Jennings &
these variables. Barkham, 1975), Ariolimax colombianus (Cates
420 J. IGLESIAS & J. CASTILLEJO
& Orians, 1975) and Helix aspersa (Dan, 1978; nemoralis, the existence of a strong attraction
Gallois & Daguzan, 1989). Other studies make towards the odour of Urtica. In the field, when
reference to high ingestion and/or assimilation people cut the weeds of road margins, we
rates of terrestrial gastropods with Urtica observed how land snails come in hordes to the
dioica: Mason (1970) with Discus rotundatus, remainders of Urtica, which give off a strong
Jennings & Barkham (1976) with Arion ater, odour. In the herbivorous prosobranch Litto-
Staikou & Lazaridou-Dimitriadou (1989) with rina littorea, the olfactory and/or gustatory cues
Helix lucorum and Lazaridou-Dimitriadou & which determine attractiveness of food items
Kattoulas (1991) with Eobania vermiculata. A are the primary determinant of food preference
number of studies point out Urtica doica as an (Imrie, Hawkins & McCrohan, 1989).
important component in the diet of land gas- Urtica dioica is a nitrophilous plant (Olsen,
tropods: Grime & Blythe (1969) with Arianta 1921) which possesses high nutrient quality; it is
arbustorum, Pallant (1969) with Deroceras retic- well known for its high content of proteins
ulatum, Wolda et al. (1971) with Cepaea (Pollard & Briggs, 1984), minerals (Vulink &
nemoralis, Chatfield (1976) with Monacha can- Drost, 1991), calcium (Pramila, Kumar &
tiana and Trichia striolata, and Hatziioannou Raghuvanshi, 1991) and vitamins A and C
et al. (1994) with Helix lucorum, Bradybaena (Pollard & Briggs, 1984). The analyses of the
fruticum and Monacha cartusiana. different plant species of the plot showed that
Also, the association between the distribu- Urtica differed statistically from the other
tion of nettles and that of snails has been found species because of its high protein, ash and
in several studies: Grime & Blythe (1969) make calcium contents. The optimum protein content
reference to high densities of Arianta arbusto- of the diet of Helix aspersa ranges between
rum in patches of Urtica dioica, and the same 15.5% (Bonnet, Aupinel & Vrillon, 1990) and
was seen by Mason (1970) with Trichia striolata 17.5% (Sanz Sampelayp, Fonolla & Gil
and Chatfield (1976) with Monacha cantiana; Extremera, 1991) and none of the four com-
Baur (1986) found densities of 11.3–17.3 monest plant species of the plot reach these
snails/m2 in patches of Urtica dioica for a popu- levels on a fresh-weight basis, but Urtica was
lation of Arianta arbustorum; Cain, Cook & the plant with the highest protein content. Also
Currey (1990) with Cepaea nemoralis found the higher ash and calcium contents of Urtica
higher densities of snails in nettle patches than in comparison with the other plant species of
in the surrounding grassland and they report the plot could be of prime importance in deter-
that changes in the cover of Urtica were accom- mining the preference of the snails, since
panied by changes in the snails’ density; Williamson & Cameron (1976) consider that
Oliveira Silva (1992) found 4–5 adult Helix the need for minerals can explain the food pref-
aspersa on each plant of Urtica. erence of Cepaea nemoralis in the field in better
The question that arises from this study is than other factors like the energetic value of the
whether Helix aspersa is associated with Urtica food. Calcium has been traditionally considered
dioica for the food value of the plant or whether the most important element for land snails, and
this association is caused by other reasons like the availability of calcium is directly related to
habitat suitability. The high ingestion and the vitality, growth, reproduction and survival
assimilation rates and the high acceptability of of land snails (Godan, 1983). Snails usually eat
Urtica dioica for land gastropods, seem to indi- soil and are able to extract calcium from acid or
cate that the food value of the plant may be the neutral soils (Crowell, 1973), but distribution
reason for the preference of these animals for and abundance of land snails may be limited by
the nettles. Thus, the association between the calcium (Wäreborn, 1970; Mason, 1974). Mason
distribution of nettles and Helix aspersa in the (1974) pointed out that land snails often appear
present case may be caused by Urtica dioica associated with calcium sources in non-calcare-
being the best available food plant for the ous areas. In Ceylon, where the soils are very
snails. acid, Achatina shows a great preference for the
The value of Urtica as food for land snails is tea plants, which have a high content of calcium
supported by reasons related to both attractive- (Mead, 1961). The soil of the plot studied here
ness and nutritional properties. In relation to is also acid (pH 5 5.9) and Urtica was the plant
attractiveness, the ability of land snails for species with the highest calcium content, so this
olfactory orientation towards foods has been could be a primary determinant in the snails’
demonstrated (Farkas & Shorey, 1976; Gallois preference for Urtica dioica.
& Daguzan, 1989), and Grime et al. (1970) Another aspect not studied here is the
found, in laboratory experiments with Cepaea presence of secondary compounds in the
 FEEDING OF HELIX ASPERSA IN THE FIELD 421
plants, which are important factors determining Wolda et al., 1971; Richardson, 1975; William-
food preference in Arianta arbustorum (Speiser son & Cameron, 1976; Chatfield, 1976; Richter,
& Rowell-Rahier, 1991). This could also be 1979; Carter et al., 1979; Szlavecz, 1986; Speiser
related to the preference of Helix aspersa for & Rowell-Rahier, 1991; Hatziioannou et al.,
nettles, since there is no evidence that Urtica 1994) have reported a predominance of senes-
dioica contains any internal defensive com- cent material in the natural diet of land gastro-
pounds (Pullin & Gilbert, 1989). pods, but references in the literature about
However, it is also possible that snails select different diets among adults and juveniles are
the nettle patches for reasons not related to the scarce and ambiguous: Wolda et al. (1971)
food value of the plant, and then eat Urtica for found that some plant species are eaten by juve-
its closeness and availability. Bailey (1989) nile Cepaea nemoralis rather than by adults and
found that terrestrial gastropods often remain vice versa, and Williamson & Cameron (1976)
in the same food patch and eat the first food found that juveniles of the same species show
item encountered; Paine & Vadas (1969) and significantly less food selection than adults.
Arrontes (1990) consider that food preference Hatziioannou et al. (1994), in a study of the diet
in marine herbivores is a response to availabil- and food preferences of five coexisting species
ity more than a response to the nutritive value of land snails, found that adult Bradybaena
of the food. fruticum eat more senescent material than
Animals with high mobility can select an area juveniles in autumn, but with respect to the
to live for its physical characteristics and then other species and the rest of the year, they
move to preferred feeding areas, but in less found no differences between adults and juve-
mobile animals, habitat selection may be a com- niles. To our knowledge this is the first time that
promise between the physical characteristics of a consistent difference between adult and juve-
the area and the value of the existing food. nile diets have been pinpointed. It seems obvi-
Nettle patches may be a suitable habitat for ous that the soft and tender green vegetables
land snails because Urtica dioica is a perennial may be easier to eat for the young snails, and
plant which survives well under heavy grazing green plants often contain more nitrogen, but
pressure (Pullin & Gilbert, 1989) and which can more research is needed to confirm this result
provide to the snails protection from predators and to look into the possible causes.
(Cain & Currey, 1967) and from eventual
human harvest. The stinging trichomes of ACKNOWLEDGEMENTS
Urtica are an effective deterrent for most
vertebrate herbivores (Pollard & Briggs, 1984; I am grateful to Dr. Bernhard Speiser for his com-
Pullin & Gilbert, 1989) but not for slugs and ments on a draft of the manuscript, and to the anony-
snails (Grime et al., 1968; Mølgaard, 1986; mous referees who made useful suggestions. Also
Pullin & Gilbert, 1989) and thus the potential thanks are due to the Agricultural and Phytopatho-
competition for food with other herbivores logical Laboratory of Galicia for the analyses of
must be reduced. The other plant species in the plants.
plot, Gramineae, Mentha suaveolens and
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