Applied Animal Behaviour Science 191 (2017) 17–23

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Applied Animal Behaviour Science

journal homepage:www.elsevier.com/locate/applanim

Behavioral patterns of (co-)grazing cattle and sheep on swards differing

in plant diversity
a,b,∗ c b
Mario Cuchillo Hilario , Nicole Wrage-Mönnig , Johannes Isselstein
a
National Institute of Medical Sciences and Nutrition Salvador Zubirán (INCMNSZ), Animal Nutrition Department “Fernando Pérez-Gil Romo”, Vasco de Quiroga 15,
14080, Mexico City, Mexico
b
University of Goettingen, Department of Crop Sciences, Grassland Science, Von-Siebold-Strasse 8, 37075, Goettingen, Germany
c
University of Rostock, Agricultural and Environmental Faculty, Grassland and Fodder Sciences, Justus-von-Liebig-Weg 6, 18059, Rostock, Germany

article info abstract

Article history: Both botanical composition and the presence of additional grazer species may modify the grazing effi-ciency and ingestive
Received 3 November 2016 behavior of ruminants. However, at present, potential effects of interactions between sward diversity and presence of multiple
Received in revised form 3 February 2017 animal species on the grazing behavior of cattle and sheep are poorly understood. The objective of this study was to evaluate
Accepted 15 February 2017 Available online 22
the effect of mono- or co-grazing of sheep and cattle, as well as the influence of sward botanical composition (either diverse
February 2017
or grass-dominated swards) on animal behavior patterns in a fully factorial design ( = 0.05). Sixteen core animals (two mono-
grazing animals per species on two sward types, plus two co-grazing animals per species on two sward types) were observed
Keywords:
simultaneously. The main behavior (grazing, walking, ruminating/resting) of core animals per paddock was recorded
Biodiversity
repeatedly in scan samplings every ten minutes from 6 to 22 h on twelve observation periods distributed over the grazing
Mixed grazing
Botanical composition seasons of 2009 and 2010. Four daylight quarters of four hours each were differentiated within one observation period.
Foraging behavior Additionally, bites per minute, bites per step and steps per minute were observed 15 times per core animal on each observation
Grassland date. Behavioral observations showed that cattle decreased the time allocated to grazing in the presence of sheep (co-grazing
management; P ≤ 0.036) and ruminated/rested longer (P ≤ 0.023). However, sward botanical composition had no effect on
cattle behavior (P ≥ 0.05). In contrast, sward botanical composi-tion impacted sheep behavior to a greater degree than mono-
or co-grazing management (P ≤ 0.05). Both cattle and sheep had a tendency to spend more time grazing and less time
ruminating/resting towards the end of the day. In cattle, longer grazing periods were followed by longer ruminating/resting
times, whereas sheep varied less in their time spent grazing or ruminating throughout the day. Sheep and cat-tle responded
differently in their grazing behavior to vegetation composition and grazing management showing a potential of the livestock
to adapt to a broad range of conditions. The behavior data suggests that livestock performance might gain from co-grazing
management. Cattle and sheep behavior should be considered in planning future directions for grassland management and
productivity.

© 2017 Elsevier B.V. All rights reserved.

1. Introduction
Animal grazing behavior is influenced by a range of fac-tors, including
environment, weather, temperature, geographical conditions, sward surface-
height, time at pasture, age, herbage allowance, stocking density, and
botanical composition of pastures. Botanical composition both affects and is
affected by grazers’ selec-tivity, which differs among animal species
(Benavides et al., 2009;
∗
Corresponding author.
E-mail address: mario.cuchilloh@incmnsz.mx (M. Cuchillo Hilario).
Villalba and Provenza, 2009; Dennis et al., 2012). The spatial dis-tribution of
forages and the botanical composition of grasslands modify the behavior of
ruminants (Hejcmanová et al., 2009; Lin et al., 2011; Liu et al., 2015; Ferreira
et al., 2016). For instance, in grasslands with a highly clustered distribution of
less preferred for-ages the intake rate of more palatable forages by cattle
increased, while it decreased in swards with a more even distribution of less
preferred species (Wang et al., 2010). Moreover, the time allo-cated to grazing
might also vary on diurnal and seasonal timescales, with the length of daylight
(photoperiod) and temperature playing important roles (Dumont et al., 2004;
Röver, 2006; Rutter, 2010; Pokorná et al., 2013).

http://dx.doi.org/10.1016/j.applanim.2017.02.009 0168-
1591/© 2017 Elsevier B.V. All rights reserved.
18 M. Cuchillo Hilario et al. / Applied Animal Behaviour Science 191 (2017) 17–23
Employing grazing to manage the botanical composition of grasslands is
normally accepted as a low-cost practice, because it requires little capital
investment. In addition, grazing provides several ecosystem services (Metera
et al., 2010; Wrage et al., 2011; Fraser et al., 2014), e.g. effective control of
weeds, fire con-trol, increase of plant diversity, avoidance of fodder
senescence, increase of seed dispersal, creation of micro-niches within
pastures by forage selection, enhanced generation of micro-catchments by
trampling and an efficient recycling of nutrients by manure depo-sition
(Celaya et al., 2007; Smith et al., 2010; Fraser et al., 2014; Rao et al., 2015).
Moreover, co-grazing of multiple grazer species is expected to have additional
benefits over mono-grazing man-agement. Complementary and adjusted
intake preferences from different grazer species pasturing together may lead
to a more complex vegetation structure than mono-grazing (Animut and
Goetsch, 2008; Díaz Falú et al., 2014). Besides, complementary use of
resources by co-grazing animals might result in improvements in the
performance of single or multiple animal species (Benavides et al., 2009;
Anderson et al., 2012; d’Alexis et al., 2014).
Studies of grazing behavior are essential sources of informa-tion for
designing improved grazing schemes that enhance animal productivity and
welfare while increasing grassland diversity and providing ecosystem services
(Utsumi et al., 2009; Lin et al., 2011; Meier et al., 2012). However, little is
known about animal behavior in relation to botanical composition and co-
grazing in permanent grasslands (Nolan et al., 2001; Fraser et al., 2007;
Fraser et al., 2013). The majority of studies in this context have been carried
out as cafeteria trials (Wang et al., 2010) or with few choices of forages at
pasture (Edouard et al., 2010; Ginane and Dumont, 2010; Glienke et al.,
2010), mainly on grass/clover grasslands (Nolan et al., 2001) hindering
natural selection by herbivores.
In this study, we evaluated the animal behavior patterns of cattle and
sheep pasturing alone or together at two different plant diver-sity levels
(diverse or grass-dominated swards) on semi-natural grassland. Diverse
swards display a more complex vegetation struc-ture and spatial distribution
of food items than grass-dominated swards and the interaction between
ruminant species enhances the competition for food items. Also, both animal
species utilize the maximum rumen capacity to avoid grazing events after
dusk. Therefore, we hypothesized that 1) on diverse swards, cattle and sheep
increase the time spend grazing at the expense of ruminat-ing and walking; 2)
on diverse swards, the bite and step rate of cattle and sheep is increased; 3)
co-grazing of cattle and sheep increases the grazing time of both grazer
species independent of sward com-position; and 4) grazing activity is lower in
the morning than in the evening.
2. Material and methods
2.1. Experimental set up
This trial was carried out on mesotrophic permanent grass-land in the
Solling Uplands of Lower Saxony, Germany from May to September 2009
and 2010. Species diversity of pad-docks was manipulated by the use of
herbicides [(Starane XL (Fluroxypyr-1-methylheptyl-ester) and Duplosan
((2R)-2-(4-chloro-2-methylphenoxy) propanoic acid)] in 2006 and 2009,
2
resulting in grass-dominated swards (9 species per 9 m ) as described by
Seither et al. (2012). In contrast, untreated swards had a higher level of
2
diversity (14 species per 9 m ) and were composed of grasses, forbs and
legumes. Each sward type was combined with the following three grazing
treatments in a fully factorial design: C = cattle mono-grazing; S = sheep
mono-grazing and CS = cattle and sheep co-grazing. The six treatments were
set up on 0.5 ha paddocks, replicated three times in rotationally grazed
blocks. As the experiment was established before behavior mea-surements
started, cattle and sheep had grazing experience on the same site and
management. Non-lactating cattle (German Sim-mental; 640.9 ± 48.4 kg live
weight) and sheep (Black headed and Leine breeds; 75.6 ± 12.8 kg live
weight) were stocked during three rotations per year as follows: the stocking
−1
density was 12.1 ± 0.9 and 12.7 ± 0.7 livestock units (LU) ha for rotation
one and two (May and June), respectively (LU = 500 kg of animal live
weight). Due to reduced herbage production of the swards, stocking den-sity
per paddock was reduced from July onwards (third rotation) to 8.2 ± 0.5 LU
−1
ha . In co-grazing plots, LU contributions of cattle and sheep were each half
of the LU of mono-grazed plots. Rumi-nants were moved to the next block
when the average compressed sward height measured at 50 points per
paddock (zig-zag walking) had decreased to ca. 6 cm (Castle, 1976). Each
stocking period lasted on average 8.7 ± 3.6 days. A rest period of six weeks
between the second and third rotation was allowed for animal mating. Animals
did not receive any supplementary feed during the experiment.
2.2. Animal behavior
The primary (grazing, walking, and ruminating/resting) and sec-ondary
(bites per minute, steps per minute and bites per step) behavior patterns of
ruminants were recorded in two blocks (A and B) per rotation in both years as
follows: Two core animals per species and treatment were observed. The core
animals were cho-sen randomly and the same animals were observed
throughout the study. Sixteen animals (two mono-grazing animals per species
on two sward types, plus two co-grazing animals per species on two sward
types) were observed simultaneously (per block) pasturing grass-dominated
and diverse swards. The geographical conditions of block C made it
impossible to perform the scans without disturb-ing the normal behavior of
animals; therefore, it was discarded for animal observations.
Primary and secondary behavior patterns were measured according to
Dumont et al. (2004). Briefly, the animals were con-sidered to be grazing
when they kept their heads close to the ground and were biting or searching
for forage, including move-ments as biting, chewing, ingestion and head
shaking activities. A walking bout (described as a continuous activity) was
registered when animals showed no sign of grazing and walked with the head
up. Animals were described as ruminating/resting when in stand-ing or laying
positions including activities as chewing, swallowing and regurgitation of
boluses of ingesta. Previous observation and experiences aimed to depict the
main animal behavior patterns, considered the “other behavior activities” as
drinking, grooming, idling, peer interactions etc. as negligible in terms of time
consump-tion and therefore were not considered in the ethogram (Röver,
2006). Primary behavior patterns or activity bouts were obtained by scan
samplings every ten minutes from 0600 to 2200 h on each observation period.
One observation period was completed in two days: the scan samplings were
performed from 1400 h to 2200 h during the first day and from 0600 to 1400 h
on the second day, leading to 96 recordings per animal during one period.
Primary and secondary behavior patterns of the core animals were recorded on
five periods in 2009 and seven periods in 2010 throughout the whole grazing
season. Each observation period started on the sec-ond or third day after the
animals entered a new paddock. Four day-light quarters (DLQ) of four hours
each (DLQ1 = 0600 to 1000 h; DLQ2 = 1000 to 1400 h; DLQ3 = 1400 to
1800 h and DLQ4 = 1800 to 2200 h) were used to characterize the most
common behavior pat-terns during the course of the day (Dumont et al., 2004;
Röver, 2006). For the third rotation of 2010, it was not possible to conduct
animal observations after 2100 h as darkness impeded clear
scans of animals. Thus, only three observational hours were
used for the last DLQ (from 1800 to 2100 h). The time spent
by core animals to
 M. Cuchillo Hilario et al. / Applied Animal Behaviour Science 191 (2017) 17–23 19
take either fifty bites or five steps while grazing was recorded fif- 3. Results

teen times per core animal and period to estimate the secondary
behavior patterns. 3.1. Behavioral patterns across observational days

Cattle used on average 47% of time grazing, 6% walking and 47%

2.3. Statistical analysis
Animal behavior observation was analyzed by ANOVA using the Proc
Mixed model of SAS v. 9.2 (2009). Plots were considered the experimental
unit. Blocks were treated as random, while all other factors were considered
fixed. Multiple observations of the same core animal were treated as repeated
measurements. The model
employed was Yij = + PDi + GTj + PDi × GTj + eij ; where Y is the tar-get
variable, is the overall mean, PDi = plant diversity treatment
i (diverse or grass-dominated swards), GT j = grazing type j (cattle or
sheep/mono- or co-grazing) and e = random experimental error. Comparison
of the means with a significant difference ( = 0.05) was established by
Tukey’s test using the macro PDMIX612 of SAS (Saxton, 1998).
ruminating/resting, whereas sheep spent 49% grazing, 4% walking and 47%
ruminating/resting. No differences (P ≥ 0.05) in primary activity patterns
(grazing, walking, ruminating/resting) were detected among seasons (data not
shown), thus only overall means are presented. Both ruminant species had two
major graz-ing bouts during the observational periods: peaks of grazing
activity were registered at 1100 h to 1200 h and at 1800–1900 h. ( Fig. 1). An
additional smaller peak of grazing activity was observed from 0600 to 0700 h.
Cattle spent longer times ruminating/resting after peak grazing events,
whereas sheep alternated shorter times for grazing and ruminating/resting.

Grazing Walking Ruminating Grazing Walking Ruminating

Cattle mono-grazing Sheep mono-grazing

100% 100%
80% 80%
60% 60%
40% 40%
acvies

20% 20%
Biodiverse
0% 0%

Cattle co-grazing Sheep co-grazing

100% 100%

80% 80%

60% 60%

40% 40%

20% 20%

0% 0%

Cattle mono-grazing
100% 100% Sheep mono-grazing
80% 80%
60% 60%
40%
40%
20%
20%
0%
0%
Grass-
Percentage of
spendondifferent
Biodiverse

Cattle co-grazing
100%
80%
60%
Grass-domianted

40%
me per hour
dominated

20%
0%
0011:

0013:
0014:

0017:
0018:

0020:
10:00

12:00

15:00
16:00

19:00
6:00

8:00
9:00
007:

21:00
 1 60%
0
0 40%
%
She 20%
ep 8
co- 0%
0
graz

8:00
9:00
10:00
11:00
12:00
13:00

19:00
20:00
006:
007:

0014:
0015:
0016:
0017:
0018:

21:00
ing %

Time of day

Fig. 1. Distribution of time spent grazing, walking or ruminating/resting by cattle and sheep grazing alone or together on grass swards differing in plant species diversity (diverse or grass-dominated).
Values shown are means of animal observations within the same treatments for the complete grazing seasons of 2009 and 2010 (96 observations per observation period, twelve observation periods).
20 M. Cuchillo Hilario et al. / Applied Animal Behaviour Science 191 (2017) 17–23

Walk ing 314 43.6 .39 02 3.0 3. 3.1 92. 0. .79 5 60 3461 082 648 5820 4179 32 5642 3.5 92. 0. 3.45 4.3 .39 02 0. .93 87 403 0692 672 0897 365902 2 360 538 703 9308 2810 805 1037 7319 7284 534.6 .39 02 43.0 5.3 3.4 92. 0. .43 05 105 5247 1706 6734 9304 72450 70 594 37.968 92. 0. 6.4 53.6 .39 02 v.0 erag.A 65e 35. 0–. 0. 45. 4.3 .3–
3.2. Effects of plant diversity and type of grazing

Ru minat g/restin g1 547 9.46 5.0 0.39 02 . 50.6 53. 51. 35. 92. 0. 0. 90. 07 08143 516 87 917 063 310 0 7124 42.53 42.53 7.39 0.2 40.7 42. 46.2 34. 92. 80. 06. 07 079 9843 60 8105 0678 6235 96 0217 079 9104 25 63 5.4 53.62 4.39 02 .0 50 .
Pval ue Pval ue DLQDi ve r se Gra ss- dom in at ed DFNDFDSEMPDGTPD*GTDiv ers eGrass - do mi na te dDFNDFDSEM PDGTPD*GT

90039180758190550629902042317054339.48.41.31.39.3920.0. 0.0. 46.51. 54.51. 50.3920.0. 0. 0.0469.

a 1bT le Percn otag ftime ecatl an hd pes oens rgtf razing k,w ingal orum ni gati /resting nwh nge ograzi onal rtoge oeth wner dars iferns ng plani tsp vecisd ersity v(d ersoi rg ras- do hminated rou)t ngh obau oservt npati eriod .Valu esho wn arem oean mfans alobi serv ation hsw hint esamt etram en tsfo orth mpec letg r azing seao 2nso 0f 9and 20 A10 uANO( yk,TV tesa=0 .05 ).

CattleShee

CCSCCSAv erag eSCSSCSAv erage

DL Q= qday-ligh ut arte.DF N= deg reso fredo mn oum r.DFeat D= deg reso fredo nmd ome oin r.C=at catlem on o-g razing h;S= epms on no-g Cgrazi S=; Catlen hd oeps -c grazin .SE nM= dsta eroa hro emft Pean D=. dplan ivt ersity tream Gen T=t. grazin yg pe.Et fectsin bo hld arctes esig n ifcan taP uv e<0al .05 n.With aroi w,m wean ithds ifern tsup erscip td iferatP 0< 5. .
0150. 7560.6423.
Grazin g148.37.4 7.47.4 5.3920.0.0 .1.30.65802122091141515

Average51.43.4 9.44.4 7.3–0.0.0.0 .45.3144111884 0364712

Average43.50.4 3.49.4 7.3–0.0.0.0 .50.48880098810237056

47779075500128722414004424235860252. 42.50. 41.46. 3920.0.0.0. 48.51. 50.54. 51.3920.0.0. 0.

0 0. .19 76 802 307 8617 0621 5072 63 09

p

0210. 6710.
0.062 0.090

0.202 0.132

0.115 0.033
0680. 0340.
On mono-grazed paddocks, cattle spent more time grazing (51 and 49%)
than co-grazing cattle (43 and 44%, for diverse and grass-dominated swards,
respectively). In general, the grazing (P = 0.036) and ruminating/resting (P =

6760.
0.018

0.013

0.010
0160.
7300.
0.023) behavior of cattle was affected by the presence of sheep (Table 1). In
contrast, plant diversity had no effect on cattle behavior. No significant
differences in walking bouts were observed for cattle due to differences in

160.

010.
150.
0.05

0.04

0.03
plant diversity (P = 0.307) or presence of co-grazers (P = 0.786).

92

92

–
92
92
Interestingly, sheep spent more time grazing on grass-dominated swards
(52 and 52%) than on diverse swards (45 and 49% for mono and co-grazing,

33
3

3
respectively); however, only mono-grazing on grass-dominated swards was
significantly different from diverse swards in this respect (P = 0.013). Sheep

857.

345.
337.
41.0

49.4

46.5
grazing on diverse swards tended to spend more time ruminating/resting (51
and 46%) than those grazing on grass-dominated swards (44 and 45% for
mono and co-grazing, respectively), although only mono-grazing on diverse

b
swards was significantly different from grass-dominated swards in this
respect (P = 0.010). In contrast to cattle, the time sheep spent for walking was

754.

643.
641.
46.1

51.5

45.1
affected by sward diversity, i.e. diverse swards tended to increase the time

b
sheep allocated to walking (P = 0.050).

761.

942.
032.
45.3

51.9

44.2
ab

ab
ab

ab
861. 253.

833. 741.
42.2

49.0

46.1
a 544.
3.3. Day-light quarters effect

a
Animals spent more time grazing later in the day and more time
ruminating/resting in the morning in all treatment combinations (Table 1).

0330.

0510.
The time cattle allocated to walk increased as the day pro-gressed with an
average of 4.6, 5.4, 5.7 and 8.1% for DLQ1, 2, 3 and 4, respectively. For
DLQ1 and 2, cattle spent 45 and 47% of the time grazing while in DLQ3 the

0130.

0090.
time dedicated to this activity dropped to 40% and increased at DLQ4 (60%).
The opposite happened to the time dedicated to ruminating/resting having the

4940.
lowest value (33%) at DLQ4. Also, sheep spent more time ruminating/resting 4580.
during the early morning (DLQ1) with a plateau between DLQ2 and 3 than
dur-ing the evening twilight hours (DLQ4), while their time spent for grazing

180.
190.

increased during that period. Sheep spent most time walk-ing in DLQ 2 and 4,
except when co-grazing in grass-dominated swards, when they walked most
92

92
in DLQ3 (Table 1). However, no statistical differences were observed in this
respect (P ≥ 0.05).
3

3
3.4. Secondary behavior patterns
533.
458.

Table 2 shows the secondary behavior patterns of cattle and sheep while
grazing (bites per minute, steps per minute, and bites per step). In general,
there were few effects of either sward composition or animal grazer species
ab

ab

on secondary behavior pat-terns. However, an effect of plant diversity (P =

135.
ab 158.

0.005), grazing type (P = 0.004) and their interaction (P = 0.001) was found
ab

for bites per minute of cattle. Cattle mono-grazing on diverse swards showed
629.

the smallest bite rate (57 bites per minute) while cattle co-grazing the same
b 361.

swards increased the bite rate (63 bites per minute). Sheep co-grazing on
a

diverse swards had an increased step rate (P = 0.007) when compared to the
845.
a 045.

other treatments.
b

4. Discussion

We hypothesized that diverse sward composition would increase the

grazing time of cattle and sheep at the expense of walking and
ruminating/resting because food items are more het-erogeneous. However, in
our experiment, cattle did not react to plant diversity with observable
behavior changes, i.e. the time they allocated to grazing, walking and
ruminating/resting was not influ-enced by plant diversity. A possible
explanation for this result is that cattle are limited by their anatomical
characteristics (muzzle) in the
 M. Cuchillo Hilario et al. / Applied Animal Behaviour Science 191 (2017) 17–23 21

Table 2
Bites per minute (BPM), steps per minute (SPM) and bites per step (BPS) of cattle and sheep when grazing alone or together on diverse or grass-dominated swards. Values shown are means of animal
observations within the same treatments for the complete grazing seasons of 2009 and 2010 (ANOVA, Tukey test at = 0.05).

Means P value
Diverse Grass-dominated DFN DFD SEM PD GT PD*GT

Mono grazing Co-grazing Mono grazing Co-grazing

Cattle BPM c a a b 3 76 0.00 0.005 0.004 0.001
57.4 63.4 63.1 61.2
SPM 8.5 9.4 9.1 9.2 3 76 0.01 0.545 0.174 0.278
BPS 7.2 7.0 7.1 6.8 3 76 0.01 0.710 0.458 0.818
Sheep BPM 53.1 62.6 53.1 54.6 3 76 0.26 0.441 0.309 0.449

SPM b a ab ab 3 76 0.00 0.805 0.007 0.317

7.8 9.0 8.1 8.9
BPS 7.6 7.3 7.0 6.6 3 76 0.03 0.288 0.535 0.996

DFN = degrees of freedom numerator. DFD = degrees of freedom denominator. SEM = standard error of the mean. PD = plant diversity treatment. GT = grazing type. Effects in bold characters are
significant at a P value <0.05. Within a row, means with different superscript differ at P < 0.05.

performance of acute vertical selection (selection of different plant parts),
thus being less sensitive to changes in vegetation structure than sheep or goats
(Fraser et al., 2007; Benavides et al., 2009). Spatial distribution of animals
during the grazing activity within the swards may also be related to this
outcome, since cattle favor the grazing on sites were biomass is more
abundant, consequently being less selective (Díaz Falú et al., 2014; Tóth et
al., 2016). In the same line, Díaz Falú et al. (2014) found that site selection of
both cattle and sheep co-grazing in the same area is notably affected by sward
vegetation variables even over weather, topography, water-ing points and
shade conditions. Likely, because of distinct body size, cattle need larger
ingestion rates of forage compared to small ruminants, which may help to
restrict selective grazing of bovine animals at pasture (Rutter, 2010). Also,
Tóth et al. (2016), found higher taxonomic and plant functional diversity in
paddocks grazed by cattle in comparison to paddocks grazed by sheep, which
may be related to lower selectivity of cattle. Therefore, the distinct selective
grazing patterns of domestic grazers would have profound impli-cations in the
planning of co-grazing strategies directed to preserve plant diversity inventory
of grasslands.
In contrast to our first hypothesis, sheep on diverse swards tended to
increase the time allocated to walking, but also to rumi-nating/resting. This
was at the expense of grazing time. The larger effects of plant diversity on
sheep than on cattle are probably due to the greater selectivity of sheep. Also,
though botanical compo-sition of diverse swards is more complex than on
grass-dominated swards, likely botanical composition indorsed sheep to
perform a faster intake of more nutritive food choices than sheep graz-ing on
grass-dominated swards. This is supported by Metera et al. (2010) who
concluded that the time sheep allocate to grazing can be greatly modified by
plant diversity. On diverse swards, sheep could find more options for
selectivity, which may allow them to achieve the required food intake and
nutrients faster than sheep grazing on grass-dominated swards. This was
confirmed by the findings of Seither et al. (2012) in a study completed in the
same site of the present investigation. They found no differences in for-age
biomass intake of cattle and sheep grazing the two types of swards studied
here, although swards differed in terms of for-age nutritive value. Due to a
larger abundance of legumes and forbs, diverse swards showed better forage
quality than grass-dominated swards. They proposed that foraging efficiency
in those conditions may not have been altered and may be fairly constant.
This means that the biomass consumed by sheep and cattle was similar
between diverse and grass-dominated swards, despite the different time
allocated to grazing on both botanical compositions. However, marked
differences in the post-grazing forage quality were observed. Co-grazing and
sheep mono-grazing swards were of lower quality relative to swards grazed
by cattle. The acid deter-gent fiber value increased while protein contents
decreased on swards where sheep were stocked (mono and co-grazing) while
the
opposite trend was observed on swards where cattle grazed. This explains why
sheep on grass dominated swards spend more time selecting nutritive food
choices than on diverse swards, since small ruminants need to ingest their diet
in smaller portions with higher energy contents than cattle (Benavides et al.,
2009; Rutter, 2010). Thus, we can assume that sheep tried to avoid
unnecessary energy losses looking for preferred forages, but at the same time
performed acute feed selection among available plant species, plant propor-
tions and plant heights (Ginane and Dumont, 2010; Lin et al., 2011). The
longer time sheep spent grazing and shorter time spent rumi-nating/resting in
grass-dominated compared to diverse swards can also be attributed to their
flexibility to choose more digestible for-ages even when available forages are
of low quality (Fraser et al., 2007; Edouard et al., 2010; Villalba et al., 2010).
We hypothesized that diverse swards increase bite and step rate of cattle
and sheep. However, we found no significant changes in secondary behavior
patterns due to plant diversity. Only bites per minute (BPM) performed by
cattle were influenced by plant diversity as well as type of grazing. High bite
rates (63 BPM) in co-grazed, diverse swards were related with a shorter time
for grazing (43%) and a longer time spent ruminating/resting (51%); i.e. cat-
tle tended to spend less time grazing at high ingestive rates. In co-grazed,
diverse swards, cattle satisfied their nutritional needs by quickly consuming
forage and dedicating more time for rumi-nating/resting. In contrast to sheep,
cattle use the tongue to wrap the plants, limiting the selection among single
plants and plant parts (Ginane and Dumont, 2010; Lin et al., 2011). However,
cattle might benefit from diverse swards, since forage quality from those
swards had a slightly higher nutritive value than grass-dominated swards. A
different relationship was found for cattle mono-grazing on grass-dominated
swards. A higher bite rate (63 BPM) was here related with a longer time for
grazing (49%) and a shorter time for ruminating/resting (44%). The findings
of Seither et al. (2012) on the same site help to better understand this apparent
contradic-tion. Since forage intake among treatments was similar in the same
swards, cattle mono-grazing on grass-dominated swards tended to spend more
time grazing at high bite rates but they might also have taken smaller bites. In
line with our findings, Pokorná et al. (2013) suggested that animals
consuming grass biomass satisfy their food intake requirements in a longer
time via smaller bites than when pasturing phytodiverse biomass. However,
this should be further evaluated and confirmed as in the present study we did
not evaluate bite size.
The only significant change in the secondary behavior patterns of sheep
co-grazing diverse swards was an increase in step per minute rate. Social
hierarchies in the structure of the herd might be the reason for this outcome.
Dominance of cattle over sheep may have pushed sheep to graze alternative
areas where they would not normally graze in mono-grazing management.
Thus, territori-ality of grazing areas dominated by cattle may increase the
distance
22 M. Cuchillo Hilario et al. / Applied Animal Behaviour Science 191 (2017) 17–23
traveled by sheep within the sward and further increased the step per minute
rate independent of sward composition. Glienke et al. (2010) observed that
sheep increased bite per minute rate when the availability of protein increased
and NDF percentages decreased in the herbage on offer. However, the better
nutritional value of diverse swards in our study tended to increase bite rates of
sheep. Nevertheless, the sharper selection of sheep grazing alone decreased
herbage nutritive value to a greater extent than sheep grazing with cattle,
leading to a rapid depletion of available nutri-ents. These observations are
according to the findings of Glienke et al. (2010) who demonstrated a
reduction of bite and step rate as the nutritive value of herbage was also
reduced. To equilibrate these behavior traits between forage quality and
forage intake, sheep co-grazing on diverse swards tended to spend more time
grazing at slightly higher bite rates but likely performing smaller bites.
Our results indicated that co-grazing management influenced the pattern
of grazing and rumination time of cattle over the day, in line with the reports
of Hejcmanová et al. (2009). Cattle co-grazing diverse swards increased their
intake rate (up to 63 bites per minute) and percentage of time spent
ruminating/resting (51%) while decreasing grazing time (43%) probably due
to competi-tion for food with sheep. Although the nutritive value of diverse
swards was presumed to be better (higher protein, larger dry mat-ter
digestibility and lower ADF value) than that of grass-dominated swards based
on results of Seither et al. (2012), selective grazing of sheep led cattle to
perform a fast intake rate to meet nutritional requirements and dietary
consumption needs. To better under-stand the factors that determine the
behavior of ruminants at pasture, a further analysis of stress indicators, animal
performance, intake preferences and social interactions is needed.
During different daylight quarters (DLQ), ruminants showed modified
behavior patterns. Cattle showed a tendency to spend more time grazing and
less time ruminating in DLQ4 in all treat-ments. High temperature during the
day might result in a larger time spent for resting at midday and longer time
for grazing dur-ing sunset and the beginning night. However, cattle and sheep
had grazing activity peaks from 1100 to 1200 (DLQ2) and from 1800 to 2000
(DLQ4) while the grazing activity decreased at DL3. This indicates how cattle
reacted to high temperatures by accumulat-ing grazing bouts before noon just
before the hottest part of the day and displaying a longer grazing activity in
the afternoon as suggested by Röver (2006). Also, both species had peaks in
grazing activity with six to eight hours in between, likely to maintain rumen
functioning and sufficient degradation rates of fiber. According to Röver
(2006), cattle select the less palatable forages that require longer rumination
time in DLQ 4 to maximize grazing periods dur-ing daylight and shifting
ruminating time to the night. It should also be considered that as the day
advances, forages have larger concentrations of non-structural carbohydrates
than in the morn-ing, i.e. the same forages have higher digestibility in the
afternoon and twilight (Edouard et al., 2010; Villalba et al., 2010), which may
influence main behavior patterns; however, this should be further evaluated.
This characteristic may be useful to design part-time grazing schemes to
improve ingestion rates and production of both cattle and sheep.
5. Conclusions
The research has shown that sheep and cattle respond dif-ferently in their
grazing behavior to vegetation composition and grazing management (co- vs
mono-grazing) showing a potential of the livestock to adapt to a broad range
of conditions. Cattle behavior modified by co-grazing management and sheep
behavior modi-fied by plant diversity may be useful to plan grazing strategies
to preserve botanical inventory of grasslands.
Acknowledgement
Mario Cuchillo Hilario was supported by a German Academic Exchange
Service (DAAD) grant A/07/72730.
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