Eur. J. Soil Biol.

37 (2001) 103−111
© 2001 Éditions scientifiques et médicales Elsevier SAS. All rights reserved
S1164556301010731/FLA

Burial of bovine dung by coprophagous beetles (Coleoptera: Scarabaeidae)

from horse and cow grazing sites in El Salvador

Finbarr Gabriel Horgan*§

Escuela de Biología, Universidad de El Salvador, Final 25 Avenida Norte, San Salvador, El Salvador
Received 22 November 2000; accepted 30 March 2001

Abstract − Dung beetles that colonise horse and cow dung were collected using baited pitfall traps at three contrasting Salvadoran
sites subject to varying degrees of livestock grazing. The sites included a lowland coastal farm, a mid-altitude farm and a high
altitude pine-grass site. The quantities of cow dung buried by each of ten tunnelling species from the sites (five Coprini and five
Onthophagini) were evaluated through laboratory experiments. The quantities of dung buried in the absence of competition and the
amount of dung provided for each egg were related to female beetle body size across species. The quantities of dung buried by pairs
of each species and the low number of beetles colonising dung at the mid-altitude site suggest that much of the dung is not buried
at the site. However, sufficient colonisation occurred at the coastal farm and at the pine-grass site for dung pads at these sites to be
completely buried, indicating that competition is normally intense. In the face of such competition, the opportunistic Onthophagini
quickly provision and lay eggs as long as favourable conditions persist while the Coprini, which are rapid dung pre-emptors, can
rarely bury sufficient dung to attain full brood production. The importance and conservation of the small number of native dung
beetles adapted to Central American pastures is discussed. © 2001 Éditions scientifiques et médicales Elsevier SAS
competition / dung beetles / dung burial / El Salvador / paracoprids / tropical pastures

1. INTRODUCTION large areas of rangeland. The large-scale introduction

The importance of dung beetles in the efficient
functioning of pasture ecosystems is widely acknowl-
edged. Dung beetles, through their feeding and repro-
ductive activities, improve soil quality by working
dung into the soil [1, 6]. The tunnels that some species
construct improve water and air circulation, reducing
the effects of soil compaction [6]. Dung beetles have
also been noted as competitors of nuisance dung
breeding flies [24, 27] and in tropical forests may
reduce the impact of seed predators by burying the
seeds present in animal dung [7]. The importance of a
specific dung beetle fauna that is adapted to pasture
landscapes and the dung of domestic herbivores has
been exemplified through the success of the Australian
dung beetle project. Many native Australian dung
beetles were unable to utilise the dung of introduced
cattle, which remained on the soil surface and fouled
*Correspondence and reprints.
E-mail address: boniga@hotmail.com (F.G. Horgan).
§
Present address: Department of Biology, University of New
Brunswick, Fredericton, New Brunswick, E3B 6C2, Canada
of mainly African dung beetles notably reduced pas-
ture fouling [1, 4].
Central America has no large native herbivorous
mammals. Cattle and horses were first introduced over
500 years ago at the time of early European colonisa-
tion. However, a recent and growing demand for beef
has led to increased cattle production and the conver-
sion of large areas of native forests to pastures [21].
The increase in size and number of cattle ranches is
currently one of the principal causes of tropical defor-
estation in Latin America [25]. This conversion of
native tropical habitat to pastureland can lead to severe
losses in the diversity of coprophagous beetles at local
scales [8, 16, 19]. A reduced species richness and
biomass of coprophagous beetles could cause the same
problems in cattle ranches in some regions of tropical
America as occurred in Australia. Pasture productivity
could decrease and soil compaction increase where the
burial of dung and the construction of tunnels by dung
beetles are reduced. Furthermore, populations of dung
breeding nuisance flies might increase in the absence
of certain dung beetles that are successful fly competi-
tors.
104 F.G. Horgan / Eur. J. Soil Biol. 37 (2001) 103–111
Apart from some work in Costa Rica [18], very little
has been published on the dung beetles of Central
American cattle pastures from Chiapas (Mexico) to
Panama. The ecological and economic importance of
the small group of beetles adapted to open pastures in
the region has been largely overlooked and, although a
number of studies describe the nesting behaviour of
some common species from tropical pastures [13, 14,
22], few studies have quantified dung burial by these
species. It is useful to quantify dung burial by indi-
vidual species since this is one of the first steps in dung
decomposition, soil improvement and competition
with nuisance flies. Furthermore, such data are useful
in helping to understand resource partitioning among
dung beetles and could aid in determining conserva-
tion strategies for certain species.
The present study compares dung beetle assem-
blages in horse and cow dung in three habitats that
represent contrasting conditions of cattle production in
El Salvador (see [21]). The sites include two predomi-
nantly agro-pastoral landscapes with small to medium
herds of cattle but at different altitudes, and a mixed
pine-grass site with a small number of cattle and
horses. Through laboratory observations, I quantified
dung burial by the most common species encountered
at the field sites. These data are used to determine dung
requirements for the laying of eggs for each species
and to examine utilisation of cow dung pads by
coprine and onthophagine beetles. As such, this study
helps determine the potential for competition among
the small group of native dung beetles that can tolerate
cattle pastures in Central America.
2. MATERIALS AND METHODS
2.1. Study species
The most common dung beetles in Central Ameri-
can pastures are included in the tribes Coprini and
Onthophagini [15, 18]. Central American coprine spe-
cies are generally medium to large-sized beetles with
heavy armament. Many species exhibit obvious sexual
dimorphism [5, 22]. Fecundity is typically low and
females of most species lay up to eight eggs during
their lifetime [13, 20]. In contrast, onthophagine spe-
cies are generally small-sized beetles and, though
many species exhibit marked sexual dimorphism, male
dimorphism is common with major males developing
enlarged horns and prominences while minor males
may lack obvious secondary sexual characteristics
[17]. Fecundity is high and female onthophagine
beetles may lay hundreds of eggs during their lifetime
[28]. Both tribes consist predominantly of paracoprid
beetles, that is, beetles that construct tunnels under the
dung pad where they bury the dung and produce their
brood.
2.2. Study sites
Dung beetles were collected at three sites including
a lowland coastal farm, a mid-altitude farm and a
higher altitude mixed pine-grass site. The latter two
sites were located in the Department of Morazán and
were separated by a distance of approximately 15 km.
The lowland coastal farm was located in the Depart-
ment of La Paz approximately 120 km from the
mid-altitude site and 110 km from the pine-grass site.
The lowland coastal farm was located at the Uni-
versity of El Salvador Experimental Field Station near
Comalapa, La Paz, (13°20’ N, 87°05’ W) at about
15 km from the Pacific coast and at an altitude of
50 m. Cattle ranches are common throughout the zone
[21]. The station had 5.9 ha of pasture where about one
hundred Creole cattle and seven horses grazed. Five
moderately grazed pastures were chosen for study.
Pastures at the station were mainly of exotic pangola-
grass Digitaria decumbens Stent, with about 10 %
stargrass Cynodon plectostachyus (K. Schum) Pilger.
Fields were separated by field boundaries of Ficus
glabrata H.B.K., Enterolobium cyclocarpum (Jacquin)
Grisebach Schultes, Bursera simoruba (L.) Sargent
and Cordia alba (Jacquin) Roem and Schultes. The
climate of La Paz is typical of the hot tropical plateau
of the Central American Pacific coast with an average
annual temperature of 26 °C and an average annual
precipitation of 1 720 mm.
Three higher altitude pastures were located at a
small farm in El Rodeo at an altitude of 700 m, about
1.5 km from Jocoaitique, Morazán (13°54’ N,
88°08’ W). Cattle farms in much of the Morazán
region are mainly small family enterprises [21] so that
open pastures at mid-altitudes are rare. The pastures
covered an area of about 1.5 ha and consisted mainly
of stargrass. Barbed wire fencing separated fields.
Occasional shade trees, including Ficus spp., Lysiloma
divaricatum (Jacquin) MacBride and chaparro, Curi-
tella americana L., occurred throughout the site.
Twenty cows and a single horse grazed at the farm.
The lands surrounding this site are dedicated mainly to
vegetable and grain production, including cassava
Manihot esculenta Crantz and maize Zea mays L.,
with abundant guava Psidium guajava L., red mombin,
Spondias purpurea L. and common bamboo Bambusa
vulgaris Schrad. The average annual temperature is
23 °C and the average annual precipitation is
2 044 mm.
The mixed pine-grass site was located on Cerro
Perquín immediately outside the town of Perquín,
Morazán (13°57’ N, 88°09’ W) at an altitude of
1 300 m. It has an average annual temperature of
20.6 °C and an average annual precipitation of
2 540 mm. The site consists of small open patches of
grasses and ferns but is predominantly covered by pine
Pinus oocarpa Schiede with small numbers of cypress
Cupressus lusitanica Mill. and with little undergrowth
vegetation. This site was used as a small family plot
for the grazing of four horses and two cows and is
 F.G. Horgan / Eur. J. Soil Biol. 37 (2001) 103–111
typical of smallholdings at higher altitudes throughout
El Salvador.
2.3. Dung beetle assemblages
To compare beetle assemblages in horse and cow
dung at the three sites, ten permanent pitfall traps were
set out at each site between May and August 1997 with
about 0.7 kg fresh horse dung (five traps) and fresh
cow dung (five traps) as baits. Traps were set out in
two transects at each of the two Morazán sites,
alternating cow and horse dung baits. Transects were
separated by about 50 m with individual traps sepa-
rated by about 20 m. At La Paz, pairs of traps (one
baited with horse dung and one with cow dung) were
located in five separate pastures. Traps were separated
by a distance of about 20 m in each pasture with
varying distances between traps in the different pas-
tures (minimum 100 m). Baits were set out at each site
before 1 100 h on the same day usually every 2 weeks.
The dung used as bait at all three sites was collected
fresh from the same source and was homogenised and
frozen the day before sampling.
Beetles were collected and identified 24 h after
setting out the baits. Some females (usually n > 30) of
each species at each site were collected and dried at
60 °C in a forced draught oven. The colonisation of
horse and cow dung by individual species was com-
pared using independent t-tests (Morazán sites) or
paired t-tests (La Paz site) on log (x + 0.01)-
transformed data. The residuals of the data were
plotted following analyses to check for normality and
homogeneity. Further occasional trapping was carried
out at each Morazán site in September and October
1997. Weekly monitoring of dung colonisation had
been carried out from August 1995 until January 1998
at the Comalapa site [15].
2.4. Dung burial
Observations of species’ burial behaviour were
made using an adaptation of the method that Doube et
al. [3] used for South African coprine and onitine
species. Experiments were carried out on the five most
abundant coprine species (Copris lugubris Boheman,
Dichotomius carolinus (L.) and Dichotomius centralis
(Harold), which are nocturnal, and Phanaeus demon
Laport-Castelnau and Phanaeus eximius Bates, which
are diurnal) and four onthophagine species (Onthopha-
gus batesi Howden & Carthwright and Onthophagus
incensus (Say), which are nocturnal, and Onthophagus
höpfneri Harold and Onthophagus marginicollis
Harold, which are diurnal). However, few individuals
of O. höpfneri were captured at the field sites and
survival and burial activity of this species were low in
the laboratory. The species was therefore excluded
from much of the analyses. One additional species,
Onthophagus championi Bates (nocturnal) was in-
cluded in the experiments. This species occurred at a
forest stand in Comalapa and frequently co-occurs
with some of the aforementioned species in herbivore
105
dung in forested coastal areas (pers. obs.). Beetles
were taken from the pitfall traps and held until
required in large holding bins with compacted soil and
supplied daily with ample fresh dung.
Experiments were carried out in an open shed
between July and October 1997. The shed was well
aired, having one complete wire-mesh wall. Condi-
tions were monitored and air-circulation improved
using a large electric fan. Normal ambient temperature
(22–25 °C) and light regimes were maintained
throughout the study. During the experiments, pairs of
beetles were randomly taken from the holding bins and
transferred to prepared arenas. Arenas consisted of
rectangular plastic containers (35 cm deep × 22 cm
long × 18 cm wide) with 30 cm depth of a compacted
moist sand-soil mix (1:3) and covered with plastic
insect screening secured by strong elastic. Each con-
tainer was supplied with fresh homogenised cow dung
(about 1.5 kg for coprine species and 0.5 kg for the
onthophagine species). The moisture content of the
dung was estimated before each trial by drying 50-g
samples in a forced draught oven at 60 °C during
1 week and then reweighing the samples. Dung mois-
ture levels varied between 80 and 84 % throughout the
experiments. Analysis of variance indicated that this
variation in dung moisture did not affect beetle activity
or dung burial by any of the species at any interval.
Experiments were run for 1, 2, 4, 7 and 14 d, and
were destructively sampled at the end of each time
period. Experiments were replicated depending on the
availability of beetles and fresh dung. There were
fewer replicates for experiments with the onthoph-
agine species because of the difficulty in finding full
males among field populations (sub-males were not
used in any trial). Experiments with diurnal species
were initiated and terminated during the mornings
whereas experiments with nocturnal species were
initiated and terminated in the late evenings.
After the experimental period, the soil in the arenas
was manually sieved to find the beetles and the buried
dung. Soil and sand was removed from the buried
dung with a light, soft paintbrush and the collected
dung was dried at 60 °C until a constant weight was
attained. Following drying, the original amount of
dung buried was estimated from the dry dung weight
and corresponding moisture estimates for the particu-
lar batch of dung that was used in the experiment. For
each species, the total quantity of dung buried and the
amount of dung provided for the brood by beetle pairs
at successive time intervals was analysed using
ANOVA on log-transformed data. Following each
analysis, the data residuals were plotted to check for
normality and homogeneity. Linear regression was
performed to determine the influence of average fe-
male dry weight for each species on the average
quantity of dung buried by the species after 1 week
and the quantity of dung provided for each egg in the
brood cells. Following experiments, beetles were re-
leased to their original sites of capture.
106 F.G. Horgan / Eur. J. Soil Biol. 37 (2001) 103–111

Table I. The number of dung beetles trapped in pitfalls at three Salvadoran sites where active cattle and horse grazing occurs. Five traps were baited
with cow dung and five with horse dung for six 24-h sampling periods each 2 weeks between May and August 1997. t-Tests failed to detect differences
in the attractiveness of horse and cow dung for each species at each site. Species recorded only during the extended trapping periods are indicated
by ‘pp’.

Habitat type Pasture Pine

Site (altitude) Comalapa (50 m) El Rodeo (700 m) Perquín (1 300 m)
Dung bait Cow Horse Cow Horse Cow Horse
Aphodiinae
Aphodius sp. 1 2 2 0 0 0 0
Aphodius sp. 2 pp 0 1 0 6 0
Aphodius sp. 3 0 0 6 4 0 0
Scarabaeinae
Canthonini
Canthon cyanellus sallei Harold 0 0 1 0 0 0
C. indigaceus Harold pp 0 0 0 0 0
Coprini
Copris laeviceps Harold 0 0 2 0 0 0
C. lugubris Boheman 99 124 195 182 6 7
Coprophanaeus telemon (Harold) 0 0 1 0 2 0
Dichotomius carolinus (Linnaeus) 3 8 39 23 186 208
D. centralis (Harold) pp 1 3 4 9 4
Phanaeus demon Laport-Castelnau 234 131 0 0 0 0
P. endymion Harold 0 0 1 0 1 0
P. eximius Bates 0 0 104 146 pp 0
P. amethystinus guatemalensis Harold 0 0 0 0 pp 0
Phanaeus sp. 0 0 1 0 0 0
Eurysternina
Eurysternus mexicanus Harold 0 0 0 0 pp 0
Onthophagini
Onthophagus batesi Howden & Cartwright 6 6 139 127 0 0
O. höpfneri Harold 5 8 0 0 0 0
O. incensus (Say) 0 0 0 0 690 339
O. marginicollis Harold 74 84 1 2 0 0
Total species (extended sampling) 7 (10) 8 (8) 13 (13) 7 (7) 7 (10) 4 (4)
Total species recorded for each site 10 13 10

3. RESULTS buried dung in the experimental containers at the

3.1. Dung beetle assemblages
Only twenty species, including ten coprine and four
onthophagine species were recorded from the field
study (table I). Each pasture site had one abundant
diurnal (P. demon or P. eximius) and one abundant
nocturnal (C. lugubris) coprine species and an abun-
dant onthophagine species (O. marginicollis or O.
batesi). At the pine site, only the two nocturnal species
D. carolinus (Coprini) and O. incensus (Onthophagini)
were common. Small numbers of Aphodius spp. were
present at each site. t-Tests failed to indicate prefer-
ences among the species for either cow or horse dung
during the study period.
3.2. Dung burial
A high percentage of coprine beetle pairs buried
dung in the experiments. In general, fewer beetles
earlier time intervals (i.e. 0–1 and 0–2 d) (figure 1). At
each time interval, only a few pairs of D. centralis
were active (< 30 %). All five species buried large
quantities of dung. Dichotomius carolinus, the largest
species, buried from 250 to 350 g when active (figure
1a). This was substantially more dung than that buried
by any of the other species and three times the amount
buried by the next most successful pre-empter, C.
lugubris (ANOVA (dung buried after 14 d):
F4,127 = 32.386, P < 0.001; Tukey P-values < 0.001
for all comparisons with D. carolinus). Most species
buried dung only on the first day of colonisation with
no significant changes in the amount of dung buried
over the 2-week period (figure 1a, b, d, e) (ANOVAs:
D. carolinus, F4,139 = 0.39, P > 0.05; D. centralis,
F4,56 = 0.60, P > 0.05; P. demon, F4,143 = 0.83,
P > 0.05; P. eximius, F4,44 = 2.01, P > 0.05). Copris
lugubris was slower than the other species at burying
dung, reaching a constant weight of buried dung on the
second day after colonisation (ANOVA: F4,186 = 3.00,
P < 0.05) (figure 1c).
 F.G. Horgan / Eur. J. Soil Biol. 37 (2001) 103–111 107

Figure 1. Dung burial and egg provisioning by five species of coprine beetles from Salvadoran pastures. Burial of fresh cow dung during 2 weeks
was monitored in the laboratory by periodic destructive sampling for pairs of (a) D. carolinus, (b) D. centralis, (c) C. lugubris, (d) P. demon and (e)
P. eximius. Solid points indicate the average total amount of dung buried after each time interval. Open points indicate the average amount of brood
containing dung after each time interval. The number of pairs that actively buried dung is indicated above each solid point and the number of pairs
that produced eggs from among all active pairs is indicated adjacent to each open point. The numbers in parentheses indicate the total number of
containers that were destructively sampled at each time period. Bars indicate standard errors. Note differences in scales.

Only one pair of D. carolinus produced an egg.
This single egg, found after 14 d, was provided
with 236 g buried dung. Pairs of D. centralis
did not produce eggs on any occasion in the experi-
ments. Some pairs of P. demon (15 %) began to form
brood balls after the second day and the amount of
dung converted to brood increased over the
following days (figure 1d) (ANOVA: F2,37 = 5.83,
P < 0.01). Brood balls of P. demon contained
68.31 ± 3.33 g (x ± SE) dung. Pairs of P. eximius
formed brood balls only after the first week (figure 1e).
Brood balls of P. eximius contained 10.42 ± 0.7 g
(x ± SE) dung. Pairs of both Phanaeus species laid a
maximum of three eggs throughout the experiments
but pairs of P. eximius tended to have larger
broods (figure 2a). Preparation of brood balls by
C. lugubris began after the second day (10 %) and
increased over the following days (figure 1c)
(ANOVA: F2,50 = 8.19, P < 0.001). Pairs of C. lugu-
bris laid a maximum of eight eggs throughout the
experiments, these were located individually in brood
balls containing 19.83 ± 0.53 g dung (x ± SE) that
were continually prepared throughout the 2-week pe-
riod (figure 2a).
108 F.G. Horgan / Eur. J. Soil Biol. 37 (2001) 103–111
All five onthophagine species buried dung in the
experiments. However, only two pairs of O. höpfneri
buried dung and survival of individuals in the experi-
ments was low. Among the remaining four species,
pairs generally had buried only a small amount of
dung by the earliest time intervals, increasing the
quantity of dung buried over time (figure 3) (ANO-
VAs: O. batesi, F3,39 = 1.82, P < 0.05; O. championi,
F3,17 = 10.27, P < 0.001; O. incensus, F4,37 = 3.08,
P < 0.05) (t-test: O. marginicollis, t = –3.26, df = 10,
P < 0.01). However, the rate of dung burial declined
after the first week (figure 3). All active pairs of
O. championi, O. incensus, O. marginicollis and
O. höpfneri laid eggs. However, pairs of O. batesi did
not lay eggs during the first day and the proportion of
actively burying pairs that had laid eggs gradually
increased from 83 % after 2 d to 100 % after 2 weeks.
Egg-laying and provisioning was continuous through-
out the trials (ANOVAs: O. batesi, F3,39 = 4.02,
P < 0.01; O. championi, F3,17 = 10.55, P < 0.001;
O. incensus, F3.37 = 3.08, P < 0.05;) (t-test: O. mar-
ginicollis, t10 = –3.50, P < 0.01;). However, O. batesi
and O. incensus did not lay eggs after the first week
(figure 2b). Eggs of O. championi and O. incensus
were provided with about twice the amount of dung
(2.65 ± 0.16 g and 2.40 ± 0.19 g, respectively) as those
of the remaining species (O. batesi, 1.50 ± 0.06 g;
O. höpfneri, 0.98 ± 0.52 g; O. marginicollis, 1.01 ±
0.17 g (x ± SE)).
The average amount of dung buried by each species
during the 0–7-d interval, when burial was most
efficient, was related to the mean female body size
across species (log dung = 0.58 + 0.60(log dry female
body weight), R2 = 0.749, P = 0.003) (figure 4a). Fur-
Figure 2. The average number of
eggs laid by pairs of (a) three
species of coprine beetle and (b)
four species of onthophagine
beetle during 2 weeks in laboratory
trials. Bars indicate standard er-
rors. Note differences in scales.
thermore, the amount of dung buried with each egg
was related to the mean body size of adult females
across species (log dung = –0.85 + 1.11(log dry female
body weight), R2 = 0.96, P < 0.001) (figure 4b).
4. DISCUSSION
Pastures throughout tropical America maintain only
a small portion of the species found in native habitats
[8, 16, 19]. In this study, a few native species were
extremely abundant in the dung of domestic herbi-
vores at three contrasting habitats. Cow dung tended to
attract a greater diversity of species. However, there
was no obvious preference for either cow or horse
dung among the species in this study (but see [15]). A
number of typical forest species [5, 22] were recorded
at both the Morazán sites. These included C. telemon,
P. endymion, D. centralis, E. mexicanus and P. am-
ethystinus guatemalensis among others.
At El Rodeo, pitfall captures were generally low
throughout the sampling period. On 7 June alone,
pitfalls captured over 60 % of the total number of
coprine beetles recorded at the site. Captures on this
date were probably influenced by noticeably cloudy
conditions, which probably slowed the rate of dung
desiccation. The only open-field specialist at the site,
O. marginicollis, occurred in small numbers. The
isolated nature of the farm and the low incidence of
pastures in the zone may have influenced the occur-
rence of open-field specialists at the site.
Many of the pastureland Coprini buried large quan-
tities of dung in 1 or 2 d. Pairs of D. carolinus buried
over 300 g dung in a single day. Pairs of P. demon and
 F.G. Horgan / Eur. J. Soil Biol. 37 (2001) 103–111
Figure 3. Dung burial by four species of onthophagine beetle from
Salvadoran pastures. Burial of fresh cow dung during 2 weeks was
monitored in the laboratory by periodic destructive sampling for pairs
of (a) O. championi (open circles) and O. incensus (solid circles) and
for (b) O. batesi (open triangles) and O. marginicollis (solid triangles).
The number of pairs that actively buried dung after each time interval
is indicated at each point with the total number of containers that were
destructively sampled indicated in parentheses. Arrows indicate the
number of active pairs and total number of containers sampled for O.
marginicollis. Bars indicate standard errors.
C. lugubris, two very common species in coastal El
Salvador, buried about 100 g dung in one (P. demon)
or two (C. lugubris) days, but some pairs of both
species had buried over 500 g on the first day after
colonisation. The low incidence of egg-laying among
the Dichotomius species in this study is notable. It is
possible that the reduced space of the experimental
containers inhibited egg-laying by D. carolinus or that
the reproductive activity of this species was reduced
during the period of study. Dichotomius carolinus is
most active during April and May [15, 23]. Burial
activity by D. centralis was very low at each time
interval (< 30 %) and no eggs were laid on any
occasion. Dichotomius centralis is most active in
September [15], at the time of study the beetles were
reproductively active as indicated by ovarian condition
(pers. obs.). Dichotomius centralis is a common spe-
cies in forested areas where it colonises horse dung in
higher numbers than cow dung (unpubl. data). It is
109
Figure 4. Relationship between (a) the log dry weight of adult female
dung beetles and the quantities of dung buried 1 week after colonisa-
tion, and (b) log dry weight of females and the amount of dung
provided for each egg. Cl, C. lugubris; Dc, D. carolinus; Dct, D.
centralis; Ob, O. batesi; Oc, O. championi; Oh, O. höpfneri; Oi, O.
incensus; Om, O. marginicollis; Pd, P. demon and Pe, P. eximius.
possible that adults will feed from cow dung but that
the dung is not a suitable substrate for egg production.
The smaller onthophagine beetles generally buried
dung at a slower rate than the coprine beetles. How-
ever, dung burial by O. incensus after 1 week and O.
championi after 2 weeks was comparable to that bur-
ied by the much larger P. eximius and D. centralis.
Therefore, even though the rate of dung burial is
slower, at a single colonisation event, pairs of these
onthophagine beetles may bury as much as some of the
coprine species. It is important to note in this study
that emigration from the experimental dung pads was
restricted and this may have resulted in some ontho-
phagine pairs burying more dung in the arenas than
they would have if emigration were permitted. Still,
the rate of dung burial after the first week declined by
at least 30 % in all three species where full data sets
were available. Furthermore, there was no increase in
the number of eggs laid by O. batesi and O. incensus
after the first week. Under natural conditions the burial
110 F.G. Horgan / Eur. J. Soil Biol. 37 (2001) 103–111
of dung by onthophagine beetles in pads of older than
1 week is probably limited [26]. Even in the case
where dung was kept in the shade of a shed, older dung
was utilised less. Therefore, in open fields where dung
desiccation is greater the period during which pairs
bury dung and lay eggs is probably even shorter.
This study evaluates dung burial by the dominant
tunnelling dung beetles from Central American pas-
tures. All seven tunnelling species from Comalapa as
well as the most common species from El Rodeo and
Perquín have been evaluated. The results for both
coprine and onthophagine beetles indicate that under
favourable conditions many of the species will bury
substantial quantities of cow dung. However, quanti-
ties of dung buried by beetles in the field will depend
on habitat conditions and in particular on the hardness
of the soil. According to this study, during the wet
season and under optimal conditions for dung burial, a
fresh dung pad weighing about 0.7 kg might support
seven to eight pairs of C. lugubris and/or P. demon, or
two to four pairs of D. carolinus. At the coastal and
pine-grass sites, pitfall trapping indicated that during
much of the year the number of beetles attracted to
0.7 kg cow dung on the first days of dropping exceeds
this amount, so that most of the dung could be buried.
Dung pads at the pine-grass site were often com-
pletely buried while much of the dung in the pastures
was left on the surface, even despite the high levels of
colonisation at Comalapa. Rapid drying of dung in the
pastures and in particular the formation of a hard crust
(pers. obs.) may deter beetle colonisation [2]. It also
reduces the effective pad volume, which may further
increase the likelihood of competition [10]. Still, the
rapid pre-emption of dung by coprine beetles suggests
that even if dung dries rapidly in the open, early
colonisers may still bury large amounts.
Dung beetles are not active during the dry season.
The heavy tropical rains at the start of the wet season
wash away dung that accumulates during these dry
months [18]. Therefore, the long-term accumulation of
dung in pastures even along the coast is unlikely.
However, other beneficial dung beetle functions may
be affected where dung is colonised during only a
short period. A lower activity of dung beetles in
heavily grazed open fields may lead to less tunnelling
activity so that the effects of soil compaction are not
countered. Currently soil compaction is a problem in
many cattle producing areas in El Salvador [9].
At each site there was an obvious diurnal and/or
nocturnal coprine-onthophagine species pair colonis-
ing domestic herbivore dung. This suggests that post-
colonisation behavioural mechanisms permit an over-
lap in dung colonisation by coprine and onthophagine
species under specific habitat conditions. It is likely
that species-specific dung burial strategies are in-
cluded among these mechanisms.
In the laboratory, some female C. lugubris laid up to
eight eggs in a single nest. Large broods only occurred
in experiments conducted in October, when young
recently emerged females were abundant. Therefore,
assuming that the beetles trapped in the field are a
random sample of the active population, then most
females do not have the opportunity to lay all their
eggs at once. On many occasions, the colonisation of
dung may be unsuccessful and no egg is laid. This will
occur in heavily colonised pads, particularly among
large species such as P. demon, which requires over
50 g fresh dung for each brood ball, or D. carolinus,
which buried over 200 g with a single egg. Successful
resource pre-emption is determined by the species’
ability to quickly locate fresh dung, its ability and
efficiency in dung burial and its body size.
The Onthophagini, because of their smaller body
size, provide considerably less dung for each egg when
compared to the coprine species. Egg-laying probably
continues in each pad until fresh dung is no longer
available for the beetles or the energetic costs of
procuring sufficient dung for a single egg become too
great. In the presence of large numbers of competitors,
female onthophagine beetles may lay only a small
number of eggs before the resource is depleted. How-
ever, at pads with little colonisation by potential
competitors, females can produce a large number of
eggs in a short time. This is similar to the mechanism
proposed to explain the presence of onitine beetles in
dung with competitively superior coprine beetles in
South Africa (see [3, 11, 12]).
There is a growing interest in the conservation of
tropical dung beetles many of which are now restricted
to small fragments of the original tropical forest
vegetation [8, 19]. However, conservation in tropical
farming landscapes has received little attention. The
role of coprophagous beetles in dung decomposition
needs to be emphasised to encourage conservation
efforts in modern Latin American farms and especially
in pastureland, which is often the dominant habitat
matrix between isolated forest patches. The efficiency
in dung burial by the few synanthropogenic species
studied here highlights their importance to agro-
pastoral ecosystems. An understanding of the require-
ments and activities of these dung beetles can help
improve management strategies for livestock grazing
in the pasture landscapes that dominate much of the
Pacific coast of Central America.
Acknowledgements. I wish to thank the landowners
and the staff at the Agronomy Faculty of the Univer-
sity of El Salvador for access to field sites. Thanks to
L. Serrano Cervantes who helped in a number of
logistic aspects of this work, to A. Solís (Instituto de
Biodiversidad [InBio], Costa Rica) and E. Cano (Uni-
versidad del Valle, Guatemala) for help in identifying
beetles, and V. Valencia Durán and R. Fuentes Morón
for help with field-work. Dan Quiring and two anony-
mous reviewers made many helpful comments on
earlier drafts of this manuscript. The General Author-
ity on Renewable Natural Resources of the Ministry of
Livestock and Agriculture (El Salvador) gave permis-
sion to collect dung beetles during this study.
 F.G. Horgan / Eur. J. Soil Biol. 37 (2001) 103–111
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