Agriculture, Ecosystems and Environment 91 (2002) 25–36

Shady field boundaries and the colonisation of dung by

coprophagous beetles in Central American pastures
Finbarr G. Horgan∗
Escuela de Biologı́a, Universidad de El Salvador, Final 25 Avenida Norte, San Salvador, El Salvador
Received 1 September 2000; received in revised form 7 June 2001; accepted 29 June 2001

Abstract
This study examines how the shade from field boundaries may enhance dung decomposition in tropical pastures through its
effects on coprophagous beetles. Between August 1995 and January 1998, beetles were collected using dung-baited pitfall traps
at a cattle ranch in coastal El Salvador. Fifteen species were recorded throughout the study and, although some species were
significantly associated with field boundaries, there were no species exclusive to the boundaries. Diurnal specialist open-field
beetles arrived predominantly on the day of dung deposition whereas nocturnal generalist species colonised older dung (one
species) or colonised dung for longer (three species) particularly, in the shade. The endocoprid Aphodiinae (two species)
generally colonised dung that was 3–7 days old, at a time when activity of the larger paracoprid Coprini and Onthophagini
had diminished. Deposition of dung in the shade of field boundaries extended the period during which it was suitable for
colonisation, raising the average dung colonisation time by generalist species. The importance of shady field boundaries in
dung beetle conservation and efficient dung decomposition in pasture ecosystems is discussed.
© 2002 Elsevier Science B.V. All rights reserved.
Keywords: Central America; Dung beetles; Field boundaries; Scarabaeidae; Succession; Tropical pastures

1. Introduction et al., 1998). Understanding how the members of

The destruction of natural habitat to accommodate
farming in tropical regions has lead to losses in the
diversity of a number of insect groups (Klein, 1989;
Brown and Hutchings, 1997; Davies and Margules,
1998). Modern day intensification of farming can lead
to further declines in diversity by excluding some of
the species that have adapted to the landscapes of
traditional farms (Perfecto et al., 1997; Vandermeer
∗ Present address: Department of Biology, University of New
Brunswick, Fredericton, New Brunswick, Canada E3B 6C2.
Tel.: +1-506-452-6314.
E-mail address: boniga@hotmail.com (F.G. Horgan).
insect assemblages respond to different landscape fea-
tures in agricultural areas can be useful in the design
of successful land management strategies for insect
conservation.
Most of the native vegetation of the Pacific Coast of
Meso-America has already been cleared for agricul-
ture. The destruction of native tropical dry forests to
create permanent pastures has been particularly promi-
nent (Koop and Tole, 1997; Trejo and Dirzo, 2000).
Studies conducted throughout Tropical America have
noted local losses in coprophagous beetle diversity
in response to habitat destruction with induced pas-
tures and clear-cuts maintaining only a small portion
of the species found in native forests (Howden and

0167-8809/02/$ – see front matter © 2002 Elsevier Science B.V. All rights reserved.
PII: S 0 1 6 7 - 8 8 0 9 ( 0 1 ) 0 0 2 6 1 - 4
26 F.G. Horgan / Agriculture, Ecosystems and Environment 91 (2002) 25–36
Nealis, 1975; Klein, 1989; Estrada et al., 1998). It is
important to maintain dung beetle diversity and pro-
mote dung beetle activity in pastures given the role
that these beetles play in the efficient functioning of
pasture ecosystems (see Bornemissza, 1976; Edwards
and Aschenborn, 1987; Horgan, 2001).
Tropical forests offer beetle assemblages a diver-
sity of microhabitats related to the biophysical make
up of the forest (i.e. stands of tree species, hills and
troughs, soil types) and the availability of substrates
for oviposition and feeding (dung from forest mam-
mals, cadavers and decaying fruit and fungi). But,
tropical pastures lack the microhabitat diversity of the
original forests. However, in some parts of Central
America, many cattle farms still consist of small herds
on roughly cleared land, often with extensive-over-
grown field boundaries, small forest stands, or patches
of native grasses and mixed scrubby-growth known
as “matorral” (Marroquı́n, 1992). Such pastureland is
generally regarded as inefficient and there is usually
pressure to intensify production through the plant-
ing of exotic C4 grasses, the reduction of shade (in-
cluding the use of electrical fencing), construction of
stables and the adoption of rotated grazing systems
(Marroquı́n, 1992).
A number of dung beetle species may be depen-
dent on the presence of shady field boundaries and
isolated stands of trees (including cultivated species)
for their survival in agricultural areas (Estrada et al.,
1998). With homogenisation of the pasture land-
scape through intensification of animal production,
species that are shade specialists might become
excluded from large areas of tropical rangeland.
Understanding how dung beetles utilise dung in
semi-intensive cattle farms may point to strategies
to improve dung degradation, maintain beetle diver-
sity and facilitate the dispersal of species between
forest-fragments where pasture is a dominant habitat
matrix.
This study examines the colonisation of dung by
beetles present in pastures at a lowland semi-intensive
cattle ranch in El Salvador. It examines the effect of
shade from field boundaries in determining the nature
of the dung colonisation curves for the most abundant
beetle species and suggests mechanisms by which
such boundaries might maintain species diversity and
improve dung decomposition in induced pastures in
Central America.
2. Materials and methods
2.1. Study species
A small number of dung beetle species commonly
occur on herbivore dung in Central American pas-
tures. Beetles of the tribes Coprini and Onthophagini
(subfamily Scarabaeinae) are among the most visu-
ally apparent (Janzen, 1983; Horgan, 2001). Coprine
species are generally medium to large beetles with
heavy armament and low fecundity. Onthophagine
species are smaller beetles and the females of some
species may lay hundreds of eggs during their life-
time. Aphodius beetles are also generally common in
herbivore dung in open-fields, these are small, light
beetles that rarely demonstrate marked sexual dimor-
phism and have medium to high fecundity (Halffter
and Edmonds, 1982).
Four major dung utilisation strategies have been
identified among dung beetles. Telecoprids cut dung
from the pad and roll the dung some distance from
the source where it is then buried. Paracoprids ex-
cavate blind ending tunnels under the source pad,
which they then pack with dung. Kleptocoprids
lay their eggs in dung previously buried by other
species. Endocoprids utilise dung without displace-
ment from the pad, feeding and laying eggs di-
rectly within the pad (Halffter and Edmonds, 1982;
Doube, 1990).
2.2. Study site
Studies were carried out at the experimental
field-station “La Providencia” of the University of El
Salvador in the Department of La Paz, El Salvador
(13◦ 20 N, 87◦ 05 W). The climate of La Paz is typical
of the hot tropical plateau of the Central American
Pacific Coast with mean daily temperatures oscil-
lating between 23 and 28 ◦ C and an average annual
rainfall of 1720 mm.
The field-station is at an altitude of 50 masl and has
an area of 32.7 ha of which 5.9 ha are dedicated to cat-
tle grazing. Over the study period, between 106 and
115 Creole cattle (mixed Jersey and Zebu) and two to
seven horses grazed pastures at the field-station. Cat-
tle were kept overnight in a wooded corral. Pastures
were mainly of exotic pangolagrass Digitaria decum-
bens Stent, with usually <10% of stargrass Cynodon
 F.G. Horgan / Agriculture, Ecosystems and Environment 91 (2002) 25–36
plectostachyus (K. Schumann) Pilger. Each field had
a good supply of water for the cattle.
The fields were generally small (0.5 ha) and sep-
arated by field boundaries, which were often over-
grown. The field boundaries were generally straight,
with barbed wire fencing held to wooden posts or
attached to live trees. Trees included Ficus glabrata
H.B.K., Enterolobium cyclocarpum (Jacquin) Grise-
bach Schultes and Bursera simoruba (L.) Sargent.
Field boundaries were not cut back during 1995 and
1996, and were about 1–2 m wide at the base with
most tree species reaching a height of 10–15 m and
a canopy diameter of about 5–7 m. In 1997, the field
boundaries were cut back to increase the area suitable
for exotic grasses, this reduced the width and shade
value of the boundaries.
2.3. Field conditions
From September 1995 to October 1996 (excluding
February and March), the water content of fresh cow
dung at the station was estimated from weekly 50 g
dung samples (n > 30) taken before 8.00 h at a cor-
ral. The moisture content of horse dung was estimated
only in June 1996. Estimates were from 20 samples
(50 g) collected immediately after dung dropping in
an open-field. Soil humidity was monitored at a sin-
gle pasture in 1996 from collected soil samples to a
depth of 10 cm (n = 10). Grass height was recorded
weekly with a graduated post at 10 points in each
of three parallel 100 m transects at the same pasture.
The distance between adjacent points was about 20 m
(a total of 30 points).
2.4. Seasonal activity
Between mid August 1995 and January 1998, a sin-
gle permanent pitfall trap was established at the centre
of each of five adjacent pastures to monitor species’
seasonal activities. Each pitfall consisted of a circular
metal container, 30 cm deep, dug into the ground and
flush with the ground surface. A tightly fitting fun-
nel, 15 cm in diameter, was placed at the mouth of the
container with the dung bait suspended on a wooden
tripod some 5–10 cm over the funnel entrance. An
empty collecting jar was placed beneath the funnel in
the metal container. Baits consisted of about 1.5 kg of
homogenised cattle dung (measured in a standardised
27
bucket) collected in the morning from a corral and
wrapped in a muslin bag. Homogenising the dung for
use in baits reduced the percent coefficient of variation
in dung moisture by about half. Baits were changed
each week before 11.00 h and beetles were collected
from the traps 2 and 7 days after baiting. In 1995,
samples of the beetles were dried and weighed. In the
2 years, from 1996 to 1998, beetles were released af-
ter capture at a central location within the station to
avoid any possibility of a trapping-out effect over the
study period.
2.5. Use of habitat and succession
Throughout 1995 and 1996, pairs of pitfall traps
were placed in 10 pastures spread throughout the sta-
tion, with one trap placed in a permanently shaded
area under a field boundary, and a second trap located
in the centre of the field. Traps were baited before
11.00 h with fresh homogenised cow dung in October
and November 1995, and each month from April to
October 1996. Beetles were collected 1, 2, 4, 7, 14 and
21 days after baiting. In May 1996, the field-station
acquired a number of horses and between June and
September of 1996 a second group of 10 pitfall traps
were set-up with two traps installed in each of five
pastures. One trap was placed in the shade and the
second at the centre of each field (parallel with cow
dung-baited traps already in the field). These traps
were baited with about 1.5 kg of fresh homogenised
horse dung collected in the morning from a horse pad-
dock. Beetles were collected at the same intervals as
indicated above for traps baited with cow dung. All
traps located within the same pasture were spaced at
least 15 m apart and distances between traps in adja-
cent pastures varied from about 100 to 800 m.
2.6. Data analysis
Comparisons of the number of beetles colonising
horse or cow dung in open and shade conditions from
June until September were made using a two-way
ANOVA on square-root transformed data. Where data
could not be adequately transformed to ensure homo-
geneity and normality of residuals, the non-parametric
Scheirer, Ray and Hare ANOVA was used (Zar, 1984).
Analyses of colonisation curves for different dung
beetle species under open and shade conditions were
28 F.G. Horgan / Agriculture, Ecosystems and Environment 91 (2002) 25–36

carried out using calculated successional mean oc- 3. Results

currences (SMO) with pastures as sampling units.
The SMO is defined as the mean of the colonisation 3.1. Field conditions
curve and is expressed in days (Hanski, 1980). It is
the age of dung to which beetles arrive on average. Fresh cow dung had a yearly average water content
The SMO is calculated by the formula of 82.8% of fresh weight and varied little throughout
the year (Fig. 1). In contrast, soil was extremely dry

n and hard (<10% water per wet weight) throughout the
pi (ti − ti−1 )ti
SMO =
i=1
n (1) dry season (November until April) and became wetter
i=1 pi (ti − ti−1 ) through the rainy season (mid-April until October),
reaching levels above 35% water per wet weight in
where pi is the number of beetles trapped with bait August and September of 1996. Grass height ranged
of age ti and n the number of collections made along from <10 cm at the end of the dry season to over 1 m
the succession. Most authors have applied the SMO at peak height in September. Fields usually contained
to succession data from temperate climates, where the 5 to 25 cattle so that pastures were never intensively
endocoprid Aphodiinae dominate the dung commu- grazed. The water content of horse dung in July was
nity and sampling is generally conducted using artifi- 80.05 ± 0.43% of fresh weight.
cial sampling pads. In such conditions, the SMO is a The first rains at the station began in mid-April in
combination of colonisation time and dung residence 1995 and 1996. However, 1997 coincided with a re-
times by individual species. In the case of tropical gional drought when rains were about 1 month later
coprophagous beetles sampled using baited pitfalls, and there was less rainfall. Also, early May tempera-
the SMO does not incorporate the residence times for tures were notably higher (Maximum 38 ◦ C) and rela-
each species. It is particularly important to note this, tive humidity lower than in previous years (Fig. 2a–c).
when interpreting SMO estimates for endocoprids in
this study. Neither paracoprids nor telecoprids reside 3.2. Seasonal activity
within the pad, but may continually visit the pad over
a given time. Over the entire study 20,282 individuals from
Using monthly SMOs for each pasture, the effect 15 species were captured in pitfall traps. Most species
of temporal sampling point (i.e. month) and degree
of shading on the colonisation curves of Copris
lugubris Boheman, a nocturnal generalist species, and
Phanaeus demon Laporte–Castlenau, a diurnal spe-
cialist, was examined. These were the only species
sufficiently abundant in all shaded and open-field traps
to permit analyses. Comparisons were made using a
GLM factorial analysis for a completely randomised
block design with pastures as blocks and data ranked
within the blocks (Collin, 1998). For the remaining
species, data from individual pastures were pooled
and Wilcoxon’s signed-ranks test was used to compare
colonisation under different degrees of shading with
calculated species’ SMOs for all shaded and open-field
traps paired each month. Similarly, Friedman’s test
was used to compare SMOs for the most common
species that arrived at traps located in the open (four
Fig. 1. Changes in the moisture content of cow dung (<2 h old)
species) and the common species in traps placed in and pasture soil (top 10 cm) at La Providencia from September
shaded areas (four species) and post-hoc comparisons 1995 to October 1996 (dung n > 30; soil n = 10). Bars indicate
were made using non-parametric Tukey tests. standard errors.
 F.G. Horgan / Agriculture, Ecosystems and Environment 91 (2002) 25–36 29

Fig. 2. Temperature (a), rainfall (b) and relative humidity (c) with weekly biomass of dung beetles trapped in pitfalls baited with fresh cow
dung (d). Climatic data was recorded from La Providencia Meteorological Station and was averaged for 10-day intervals. Bars indicate
standard errors.

were represented by few individuals (Table 1). Adult
dung beetles were active throughout most of the year
(Fig. 2d), but with a lull in activity between January
and March. Beetle activity peaked between June and
August. In general, females were more frequently
trapped than males (Table 1). Fewer beetles were
captured in 1997. C. lugubris was the most abundant
species and the only species trapped between late
January and mid-April. An increase in beetle activity
during December 1995 was the result of the emer-
gence of large numbers of adult C. lugubris at that
time (Fig. 2d).
Adult activity commenced for most species after the
first rains with peak activity in mid-July or mid-August
(Fig. 3). In 1997, adult emergence and peak activities
were generally later than in 1996, and many of the
smaller species remained active throughout December.
In both years, activity of D. carolinus was greatest in
30 F.G. Horgan / Agriculture, Ecosystems and Environment 91 (2002) 25–36
 F.G. Horgan / Agriculture, Ecosystems and Environment 91 (2002) 25–36
Fig. 3. Adult activity of the common dung beetle species at La Providencia in 1996 and 1997. Triangles indicate the time of peak capture
in pitfall traps at 5 or 10 pastures.
the early rainy season whereas D. centralis was most
active at the end of the rainy season.
3.3. Use of habitat
Only nine species were abundant at the study site;
three of these specialise in utilising herbivore dung
under open-field conditions. These included Aphodius
sp. 1, O. marginicollis and P. demon (Table 1). Other
species were generally more abundant in dung located
in shaded areas. These included C. lugubris, D. caroli-
nus, D. centralis and O. batesi. The smaller Aphodius
sp. 2 occurred almost exclusively in shaded areas.
Two of the three species that showed preferences for
open-field conditions were diurnal, whereas nocturnal
species preferentially colonised dung in shaded areas
(five species). Furthermore, open-field species tended
to be dung specialists (all three species) whereas
species with higher captures in the shade were pre-
dominantly substrate generalists (four of five species)
(Table 1).
31
C. lugubris and O. marginicollis showed prefer-
ences for cow dung over horse dung in shaded and
open habitats (Table 2). Shading and dung type inter-
acted to determine colonisation by P. demon, which
preferred cow dung in the open-field and horse dung
in the shade. Only one species, A. rodriguezi, was not
recorded from cow dung.
3.4. Succession
Most of the colonisation of dung in open-field
conditions occurred before the dung was 2 days
old (Fig. 4). In the open-field, dung dries rapidly.
The diurnal beetles that prefer these conditions (i.e.
O. marginicollis, P. demon and possibly C. i. chevro-
lati) generally colonised dung that was only a few
hours old (Fig. 4).
The nocturnal species, D. carolinus, D. centralis
and O. batesi generally colonised dung only on the
first day in open habitat (Fig. 4). The two common
Aphodius species were most attracted to dung that
32 F.G. Horgan / Agriculture, Ecosystems and Environment 91 (2002) 25–36

Table 2
Occurrence of beetles in pitfall traps baited with one of two types of herbivore dung set out in open-field and shaded conditions in five
cattle pastures between June and September 1996
Species Total number of beetles captured Substratea test Degree of shadinga Interactiona test
statistic test statistic statistic
Cow dung Horse dung
Open Shaded Open Shaded
Aphodius sp. 1 18 1 7 0 H = 0.63 (ns) H = 2.41 (ns) H = 3.09 (ns)
Aphodius sp. 3 2 0 2 0 – – –
A. rodriguezi 0 0 0 1 – – –
C. i. chevrolati 0 0 1 0 – – –
C. lugubris 123 420 73 196 F1,19 = 22.13∗∗∗ F1,19 = 7.50∗ F1,19 = 1.08 (ns)
D. carolinus 5 15 4 3 H = 3.10 (ns) H = 1.30 (ns) H = 3.68 (ns)
D. centralis 0 5 3 6 H = 0.63 (ns) H = 1.00 (ns) H = 1.23 (ns)
O. batesi 3 70 14 32 F1,19 = 0.05 (ns) F1,19 = 13.14∗∗ F1,19 = 2.72 (ns)
O. höpfneri 1 0 0 0 – – –
O. marginicollis 60 15 4 2 F1,19 = 8.79∗ F1,19 = 4.07 (ns) F1,19 = 3.06 (ns)
P. demon 652 37 21 59 F1,19 = 34.99∗∗∗ F1,19 = 24.55∗∗∗ F1,19 = 49.38∗∗∗
a Data are pooled from three sampling points (ns = not significant: ∗ P < 0.05; ∗∗ P < 0.01; ∗∗∗ P < 0.001). All data square-root
transformed where parametric ANOVAs were used.

Fig. 4. Dung beetle colonisation curves for cow dung placed in open (open points) and shaded positions (closed points) in 10 pastures.
Within each pasture data are pooled from six sampling points between June and October 1996. Bars indicate standard errors. Note
differences in scales.
 F.G. Horgan / Agriculture, Ecosystems and Environment 91 (2002) 25–36 33

was 3–7 days old. Aphodius sp. 1 and C. lugubris

were the only species attracted to dung after 7 and
14 days in open and shaded areas, respectively. How-
ever, this was a rare occurrence. Colonisation times
differed significantly among species that regularly
colonised dung in the open (Friedman: χ 2 = 12.30,
d.f. = 3, P = 0.006) with Aphodius sp. 1 colonising
dung significantly later than C. lugubris, P. demon
and O. marginicollis (P < 0.05 for all comparisons),
and C. lugubris significantly later than P. demon and
O. marginicollis (P < 0.05 for all comparisons).
The mean colonisation times differed among the
four nocturnal species that were common in shaded
conditions (Friedman: χ 2 =13.56, d.f.=3, P =0.004).
C. lugubris arrived at dung significantly later than the
other three species (D. carolinus, D. centralis and O.
batesi) which all colonised dung of about the same
age (P < 0.05 for all comparisons of C. lugubris with
other species). There was no colonisation of horse
dung in either open or shade conditions after 4 days
by any species.
Interestingly, two very similar sequences of dung
beetle arrival were apparent on cow dung depending
on whether it is placed in the open-field or in the shade Fig. 5. Mean colonisation times of cow dung by coprophagous bee-
tles from October 1995 to October 1996. (a) Aphodius sp. 1 (open
of vegetative field boundaries. Generally, dung was
circles) and O. marginicollis (open triangles) in the open-field
colonised on the first day by large coprine (P. demon (solid line) with D. carolinus (closed circles) and O. batesi (closed
[open]; D. carolinus and D. centralis [shaded]) and triangles) in shaded areas (broken line). Data for all pastures were
smaller onthophagine beetles (O. marginicollis [open]; pooled to calculate SMOs (n = 1). (b) C. lugubris (solid circles)
O. batesi [shaded]). The telecoprid C. i. chevrolati and P. demon (open triangles) in shaded (broken line) and open
areas (solid line). Points indicate mean SMOs (n = 10 pastures).
also colonised dung in the open-field on the first day.
Bars indicate standard errors. Most species were absent during the
On the second night, after deposition large numbers of dry season except C. lugubris, which was the only species present
C. lugubris colonised dung in both open and shaded in large numbers in November 1995 and April 1996.
areas. After activity of the larger beetles had dimin-
ished, small endocoprids (Aphodius sp. 1 [open]; Phanaeus demon colonised dung for longer in the
Aphodius sp. 2 [shaded]) reached a peak in colonisa- later months and colonised only fresh dung in July
tion. The late colonisation of dung by the Aphodius (Fig. 5b) (month: F4,63 = 10.46, P < 0.001). Older
species suggests that they avoid the burial activities dung was colonised more frequently in open than in
of the larger coprine and onthophagine species. shaded areas (degree of shading: F1,63 = 11.93, P <
The sampling period affected the form of the 0.001; interaction: F8,63 = 0.765, P = 0.635). For
colonisation curve for each species (Fig. 5). Ear- most species, colonisation of fresher dung was more
lier colonisation of dung was more frequent among apparent towards the beginning and/or end of the dry
C. lugubris in the dry season than in the rainy sea- season and in July (Fig. 5a).
son and during the July sampling period (Fig. 5b)
(month: F8,155 = 9.551, P < 0.001). Deposition of
dung in the shade extended the colonisation curve of 4. Discussion
C. lugubris in all months except October 1996 (degree
of shading: F1,155 = 16.883, P < 0.001; interaction: Baited pitfall traps are commonly employed in eco-
F8,155 = 1.386, P = 0.209). logical studies of tropical dung beetles. However, these
34 F.G. Horgan / Agriculture, Ecosystems and Environment 91 (2002) 25–36
traps have been criticised because of the difficulty in
interpreting their results (see Southwood, 1978) and
the limited information they yield on dung community
processes (Giller and Doube, 1994). This study as-
sumes that dung baits adequately simulate some types
of large naturally occurring dung pads. It assesses the
suitability of these baits for colonisation by beetles
under different habitat conditions where the baits un-
dergo different rates of change. The study makes no
inference on subsequent acceptance of, or residence
times in dung by any of the species. Therefore, the
application of pitfall traps in this study was entirely
adequate. Because traps were baited in the mornings,
the results make inference only to dung that is dropped
in the morning. At the study site, as in many parts
of Central America, cattle were maintained in corrals
at night, therefore, all of the naturally occurring dung
pads were dropped in the pastures during the day.
In Tropical America, a relatively small number of
dung beetles occur in open pastures or forest clear-cuts
when compared with adjacent forest habitats (Howden
and Nealis, 1975; Klein, 1989; Estrada et al., 1998). In
this study, only nine species were abundant throughout
the extensive study period. Adults of all the species,
except C. lugubris, occurred almost exclusively dur-
ing the rainy season. C. lugubris occurred throughout
the year, but does not construct tunnels during the dry
season when the soil is hard (personal observation).
It is likely that soil moisture was a major factor in
determining adult beetle activity because dung mois-
ture remained relatively constant throughout the year.
The cutting-back or removal of shady field boundaries
to improve the growth of exotic grasses may lead to
further declines in species richness of coprophagous
beetle assemblages (see Estrada et al., 1998) or re-
duce beetle tunnelling activity. Dung dropped in the
open-field quickly develops a dry crust and may re-
main on the soil surface for several weeks whereas
dung dropped under shade is often completely buried
within 2–3 days (personal observation). This suggests
that dung decomposition is slower in open-fields.
There are a number of mechanisms by which re-
duced shading through its effects on dung beetle as-
semblages might reduce rates of dung decomposition
in tropical pastures. This study addresses two such
mechanisms. Firstly, a loss of beetle diversity could
affect dung decomposition where there is a relation-
ship between decomposition rates and richness of the
beetle assemblage. Such a relationship has been de-
tected in aquatic decomposer communities, however,
it has not yet been investigated in dung communities
(see Finn, 2001 and references therein).
In this study, only three of the recorded species
were exclusive to open habitat, the remaining species
were typical of forested areas, but many are now also
common in open-fields at night. There is evidence
that field boundaries do maintain some species in pas-
ture ecosystems. Estrada et al. (1998) found 10 forest
species that were associated with field boundaries, but
absent from the open-field in the pasture landscapes
of Veracruz, Mexico. In this study, three species (each
represented by a single individual) were recorded only
in association with field boundaries. Each of these
species has been captured in open-fields in other stud-
ies from the region (Janzen, 1983; Montes de Oca and
Halffter, 1995; Horgan unpublished data) so they are
unlikely to occur exclusively along field boundaries.
However, Aphodius sp. 2, a common species that is
significantly associated with field boundaries, could
be affected by the loss of field boundaries and become
excluded from large open areas. This species preferred
older dung in both the open-field and shaded areas so
that its observed preference for shaded habitat is not
an artefact from differential rates of dung-bait desic-
cation. Therefore, whereas open-field specialists are
unlikely to be affected by maintaining shade in mod-
ern Central American cattle ranches, the diversity of
generalist species may be enhanced by the presence
of shady field boundaries.
A second mechanism by which reduced shad-
ing might reduce rates of dung decomposition is
through its effects on the colonisation of dung by
pasture-tolerant dung beetles. Breshears et al. (1998)
found that shade from woody plants reduced tem-
perature fluctuations and decreased the evaporation
rates of soil in shaded areas when compared with
non-shaded patches in semi-arid woodland in New
Mexico. Intuitively dung that is placed under the
shade of trees or field boundaries will dry at a slower
rate than dung placed in open-fields. Differences in
drying rates, temperatures and light intensities will
likely affect the biochemical processes involved in
dung decomposition and the rate at which dung qual-
ity changes under the two regimes.
Changes in dung quality have been shown to in-
fluence colonisation by dung beetles. Montes de Oca
 F.G. Horgan / Agriculture, Ecosystems and Environment 91 (2002) 25–36
and Halffter (1995) demonstrated that colonisation
by Onthophagus species of dung that is dropped in
the morning is significantly higher than that of dung
placed at night whereas C. lugubris and Digiton-
thophagus gazella (Fabricius) are significantly more
abundant in nighttime dung. Furthermore, dung qual-
ity changes associated with dung ageing have been
noted as important in influencing the colonisation
and succession of Aphodius dung beetles in Europe
as this relates to the oviposition behaviour of individ-
ual species (Gittings and Giller, 1998). In this study,
dung placed under shade was attractive to nocturnal
generalists for longer. Only small numbers of D. car-
olinus, D. centralis and O. batesi colonised dung in
the open-field on the first day after dropping and there
was little subsequent colonisation of open-field dung
by these species. However, dung under the shade was
colonised by significantly higher numbers of noctur-
nal generalists and, during some months, the dung was
continually colonised for up to a week after dropping.
At least with C. lugubris, these shade effects were ap-
parent throughout the year. Some of these nocturnal
generalist species may not be truly associated with
shade, but were probably attracted to the shaded dung
because it remained moister than dung in the open.
Dung deposited in the open-field at night is affected
by direct sunlight on the second day and is, therefore,
colonised by beetles during a shorter period.
5. Conclusions
There is evidence that field boundaries are impor-
tant in maintaining populations of some dung bee-
tles in tropical pastures. The shade from field bound-
aries may also enhance dung decomposition by pro-
moting dung beetle colonisation. Diurnal open-field
specialists occur predominantly on the first day after
dung deposition and avoid dung placed under shade.
Nocturnal generalists colonise shaded dung in signif-
icantly higher numbers than dung in the open and,
whereas most generalist species avoid dung after a
single day in the open, they will continue to colonise
dung in the shade even up to a week after deposition.
Dung is seldom colonised by any paracoprids (noc-
turnal or diurnal) after it has been in direct sunlight
for 2 days. By maximising light availability for exotic
grasses, farmers may inadvertently deplete dung beetle
35
diversity and reduce the efficiency of dung decompo-
sition in tropical pastures.
Acknowledgements
The author thanks staff at the Agronomy Faculty
of the University of El Salvador for their co-operation
and interest throughout this study; L. Serrano
Cervantes for help in many aspects of the research;
A. Solı́s (Instituto de Biodiversidad [InBio], Costa
Rica) and E. Cano (Universidad del Valle, Guatemala)
for help in identifying beetles, and V. Valencia Durán,
Y. López Ventura and S. Diaz Sierra for help with the
field work. J. Finn, T. Gittings, G. Moreau, D. Quiring
and anonymous referees gave many helpful comments
to improve the manuscript. The Ministry of Livestock
and Agriculture of El Salvador gave permission to
collect dung beetles throughout this study.
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